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.     

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.     

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.     

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.     

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.     

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.     

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.     

Impact of extreme flooding on the annual survival of a riparian predator,the Brown Dipper Cinclus pallasii

The predicted effects of global climate change include altered patterns of precipitation and more extreme weather events, leading to an increase in the severity and frequency of episodic disturbances such as floods. These changes may affect lotic prey communities, which could indirectly affect aquatic and riparian predators through trophic linkages. We assessed whether extreme flooding affected the apparent survival of Brown Dippers Cinclus pallasii in Taiwan using mark–resighting data and climate data. The probability of survival was negatively correlated with the degree of flooding, and survival of first‐year birds was lower than that of adults. Previous analyses of this system suggest that the main, indirect mechanism driving such patterns is the impact of flood disturbance on the aquatic invertebrate prey of Brown Dippers. Our results show that changes in prey communities induced by flooding have the potential to affect predators in aquatic and adjacent riparian habitats. This highlights the importance of considering cross‐ecosystem linkages when identifying conservation and management goals in the face of future climate uncertainty.     

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.     

Calculating census efficiency for river birds: a case study with the White-throated Dipper Cinclus cinclus in the Pyrénées

Using the binomial law we modelled field data to estimate the probability ( ̂ ) of detecting pairs of breeding White-throated Dippers, and the population size ( ̂ ± confidence limits). The model was divided into two parts according to whether the actual size of the population under study was known or not; in the latter case the truncated binomial model was used. Dipper abundance data were collected from three 4-km-long river tracts in the Pyrénées (France) during the breeding seasons of different years. Goodness-of-fit tests indicated that the binomial model fitted the data well. For a given visit during the survey, the estimated probability of detecting any pair of Dippers if they were present was always high (0.63–0.94) and constant from year to year but not between sites. Estimations ( ̂ ) of the size of the population provided by the binomial model were very close to that derived from mapping techniques. This study provides the first ever quantification of the number of visits required to detect birds on linear territories: three visits were necessary to detect the whole breeding population.     

Diet of Dippers Cinclus cinclus during an early winter spate and the possible implications for Dipper populations subjected to climate change

The prey composition of the diet of Dippers Cinclus cinclus was shown to differ between baseflow and spate conditions. In southwest Ireland, faecal pellets were collected during a heavy spate in early winter 1991 and during a period of baseflow conditions three weeks later. The diet composition differed significantly between the two periods. During baseflow conditions diet was dominated by trichopteran larvae, especially Limnephilidae, whereas during the spate period diet was composed primarily of ephemeropteran nymphs, particularly Baetidae, and dipteran larvae of the family Simuliidae. Diet during the spate period also contained greater proportions of prey taxa of terrestrial origin. Enhanced hydrological cycles with greater precipitation, predicted under current climate change models, may have an effect on the diet of Dippers and this is discussed.     

Assessing the impact of extreme adverse weather on the biological traits of a European storm petrel colony

Climate change affects the climatic disturbance patterns and regimes and is altering the frequency and intensity of subtropical cyclones. These events can affect population dynamics of seabirds (e.g., survival, reproduction). In this work we tested the effect of adverse weather on a colony of European storm petrels (Hydrobates pelagicus) located in a small islet (Aketx) in northern Spain. Over a long-term monitoring period (1993–2014) we ringed 3728 petrels. From 2003 onwards we also monitored breeding success, the percentage of immature individuals and moult scores. We used Cormack-Jolly-Seber models and Underhill and Zucchini models to analyze the effects of climatic conditions on a number of biological traits (survival, breeding parameters, moulting patterns). Our analyses revealed a constant value of adult survival over the 26-year monitoring period. Recapture probability, however, tended to be positively influenced by NAO conditions in spring, and negatively influenced by NAO conditions in winter (although this would only affect to a fraction of first-captured birds). Moreover, the impact of adverse weather, especially in 2011 and 2014, resulted in an increasing proportion of yearlings in the breeding population, a lower breeding success and a delayed onset of moult. These effects were similar to those observed during the Prestige oil spill catastrophe.     

Evidence of a highly complex phylogeographic structure on a specialist river bird species, the dipper (Cinclus cinclus)

This study details the phylogeographic pattern of the white-throated dipper (Cinclus cinclus), a Palearctic, temperate, passerine bird that is exclusively associated with flowing water. Our results reveal a complex phylogeographic structure with at least five distinct lineages for the Western Palearctic region. As for many species of the Western Palearctic fauna and flora, this genetic structure is probably linked to the isolation of populations in different southern refuges during glacial periods. Furthermore, the isolation of populations in Scandinavia and/or Eastern regions, but also in Morocco and probably in Corsica, was accentuated by ecological and biogeographic barriers during Quaternary interglacial periods. During glacial periods, Italy, Sicily and the Balkano-Carpathian region acted as major refuge zones for the dipper. At the end of the last ice age, Western Europe was repopulated by dippers from an Italian refuge, while Eastern Europe was recolonised by Balkano-Carpathian birds. A large contact zone between these two lineages was evidenced and extends from Luxembourg to Hungary. Finally, our results indicate the need to clarify the taxonomic status of the dipper, especially concerning the European subspecies whose validity appears uncertain.     

On the ecology of wintering Dippers(Cinclus cinclus) in northern Sweden

Summary Wintering Dippers were studied in the coastal region of the province Västerbotten in northern Sweden with special emphasis on winter movements, winter site-tenacity and population dynamics. In three different streams the number of wintering Dippers varied markedly during the winter, with maximum numbers in mid-winter (December–January). Sex and age ratio also varied during the winter. and adult birds outnumbered and juvenile birds during most of the winter. The wintering Dippers showed a good site-tenacity and about 17 % of the birds returned to the same stream the following season. Two long-distance recoveries and one control of a Dipper banded in Finland might indicate the origin of the wintering Dippers as well as the migratory routes of the species in northern Scandinavia, all in consistence with earlier studies of the migration of the species. Body-weight and wing-length data corresponded well with earlier investigations in southern Scandinavia.It is suggested that the structure of the wintering Dipper population can be explained by a mechanism of dominance-influenced migration proposed byGauthreaux (1978).

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Zur Winterökologie der Wasseramsel(Cinclus cinclus) in Nordschweden

Zusammenfassung 1975–1979 wurden Wasseramseln in drei verschiedenen Flüssen im Küstenbereich der Provinz Västerbotten untersucht unter besonderer Berücksichtigung der Wanderungen, Ortstreue und Populationsstruktur im Winter.Die Anzahl der Wasseramseln zeigte klare jahreszeitliche Unterschiede mit einem Maximum in den Monaten Dezember und Januar. Die Geschlechter- und Altersverteilungen variierten deutlich während des Winters; und Altvögel überwogen vor allem am Ende des Winters gegenüber und Jungvögeln. Die überwinterten Wasseramseln zeigten eine ausgeprägte Ortstreue; ca. 17 % der Vögel kehrten im folgenden Jahre zu den gleichen Flüssen zurück.Zwei Wiederfunde von Wasseramseln, die im Untersuchungsgebiet beringt worden waren, aus Norwegen und eine Kontrolle aus Finnland deuten auf Herkunft und Zugwege der in Nordschweden überwinternden Wasseramseln hin. Diese Wiederfunde stimmen mit den Untersuchungen vonAndersson &Wester (1976) überein.Die Körpergewichte und Flügellängen zeigten gute Übereinstimmung mit Untersuchungen in Südschweden (Andersson &Wester 1971, 1972).Es wird vorgeschlagen, die Populationsstruktur überwinternder Wasseramseln anhand eines Modells der dominance-influenced migration (Gauthreaux 1978) zu erklären.

     

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.     

The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long-lived seabird

1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.     

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.     

Assessing the impact of climate variation on survival in vertebrate populations

Abstract The impact of the ongoing rapid climate change on natural systems is a major issue for human societies. An important challenge for ecologists is to identify the climatic factors that drive temporal variation in demographic parameters, and, ultimately, the dynamics of natural populations. The analysis of long-term monitoring data at the individual scale is often the only available approach to estimate reliably demographic parameters of vertebrate populations. We review statistical procedures used in these analyses to study links between climatic factors and survival variation in vertebrate populations. We evaluated the efficiency of various statistical procedures from an analysis of survival in a population of white stork, Ciconia ciconia, a simulation study and a critical review of 78 papers published in the ecological literature. We identified six potential methodological problems: (i) the use of statistical models that are not well-suited to the analysis of long-term monitoring data collected at the individual scale; (ii) low ratios of number of statistical units to number of candidate climatic covariates; (iii) collinearity among candidate climatic covariates; (iv) the use of statistics, to assess statistical support for climatic covariates effects, that deal poorly with unexplained variation in survival; (v) spurious detection of effects due to the co-occurrence of trends in survival and the climatic covariate time series; and (vi) assessment of the magnitude of climatic effects on survival using measures that cannot be compared across case studies. The critical review of the ecological literature revealed that five of these six methodological problems were often poorly tackled. As a consequence we concluded that many of these studies generated hypotheses but only few provided solid evidence for impacts of climatic factors on survival or reliable measures of the magnitude of such impacts. We provide practical advice to solve efficiently most of the methodological problems identified. The only frequent issue that still lacks a straightforward solution was the low ratio of the number of statistical units to the number of candidate climatic covariates. In the perspective of increasing this ratio and therefore of producing more robust analyses of the links between climate and demography, we suggest leads to improve the procedures for designing field protocols and selecting a set of candidate climatic covariates. Finally, we present recent statistical methods with potential interest for assessing the impact of climatic factors on demographic parameters.     

Nonlinear impact of climate on survival in a migratory white stork population

1. There is growing evidence that ongoing climate change affects populations and species. Physiological limitation and phenotypic plasticity suggest nonlinear response of vital rates to climatic parameters, the intensity of environmental impact might be more pronounced while the frequency of extreme events increases. However, a poor understanding of these patterns presently hampers our predictive capabilities. 2. A recent climatic shift in the Sahel, from droughty to less severe condition, offers a good opportunity to test for an influence of the climatic regime on the response of organisms to their environment. Using a long-term capture-mark-recapture data set on a white stork (Ciconia ciconia) population wintering in Sahel, we investigated potential change in the impact of environmental conditions on survival and recruitment probabilities between 1981 and 2003. 3. We observed a decrease in the strength of the link between survival and Sahel rainfall during the last decade, down to a nondetectable level. Whether Sahel climate was found to affect the survival of storks under droughty conditions, individuals did not seem to respond to climatic variation when precipitation was more abundant. 4. This result gives evidence to a nonlinear response of a migrant bird to wintering environment. Present climate seems to fluctuate within a range of condition providing enough resources to maximize stork's survival. It suggests that whereas inter-annual variability impacted individuals, pluri-annual average condition affected the intensity of this impact. Such pattern may be more widespread than thought, and its modelling will be crucial to predict the impact of future climate change on population dynamics.     

Exploitation of prey by a river bird, the dipper Cinclus cinclus (L.), along acidic and circumneutral streams in upland Wales

SUMMARY. 1. The diet of the Eurasian dipper Cinclus cinclus. a riverine bird from a globally widespread genus, was assessed through all stages of its annual cycle using published data and field studies from streams of contrasting chemistry. Time-activity budgets were also compiled throughout the year and used to estimate annual energy requirements from the stream ecosystem. 2. The annual energy requirements for a territorial pair ranged from 148,000 to 158,000 kJ yr^?1 depending on whether one or two broods were reared. After allowing for assimilation efficiency, these requirements were estimated to represent 10.5–11.0 kg dry mass of fish and invertebrates. Using representative values for territory size (4680–11,250 m²), annual exploitation of secondary production was estimated at 0.93–2.35 g dry mass m^?2. 3. Several features combined to focus the predatory load on certain organisms over different stages of the annual cycle. These included the availability and selection of alternative prey, the need to provision nestlings with large items such as trichopteran larvae, and the use by females of calcium-rich prey such as fish prior to egg formation. 4. Across their range of territory size, annual exploitation (dry mass) by dippers was estimated at 0.06–0.29 g m^?2 for Plecoptera. 0.02–0.22 g m^?2 for Ephemeroptera. 0.59–1.11 g m^?2 for Trichoptera and 0–0.78 g m^?2 for fish. Exploitation of Ephemeroptera. I richoptera and fish were all highest in circumneutral streams, but exploitation of Plecoptera was highest in acidic streams because other prey were scarce. 5. Cottids dominated the fish component of the diet. Influences on their density could be substantial according to available data on production in Welsh streams. Hydropsychids and limnephilids dominated the trichopteran component, with exploitation again representing potentially substantial amounts of production. Contemporaneous data are required on benthic production and exploitation by the birds. 6. The ecological role of birds in rivers particularly, and aquatic ecosystems generally, is currently neglected but worthy of considerable research effort.