Reconnaissance for future breeding sites by spotted sandpipers

We studied the sex-role-reversed, polyandrous spotted sandpiper(Actitis macularia) from 1974 to 1990 in northern Minnesota,USA. After peak arrival of breeding birds and before peak departureat the end of the breeding season, there were many short-termvisitors (transients) to the study site. Stepwise discriminantfunction analysis (DFA) was used to determine the importanceof absolute sex ratio (males/female), sex of the transient bird,number of nests, and number of breeding males and females duringthe week of visit in predicting whether a visiting bird wouldreturn the following year. In addition, multiple regressionwas used to determine how much variability in the number oftransient birds returning in subsequent yearscould be explainedby annual values during the year of transience for numbers ofbreeding males and females, numbers of eggs laid and hatched,and absolute sex ratio. Annual recruitment of foreign adultsranged from 1 to 20 birds, of which 0–56% were seen visitingin previous years. Female recruits were more likely than malesto have been observed previously as transients. Twenty-two chickshatched at our study site returned and bred for the first timemore than 1 year after hatching. Of these, 9 (41%) were seenas transients between the year of hatch and breeding. The DFAshowed that transient females returned more often than transientmales and that the number of transients returning in subsequentyears was positively associated with the absolute sex ratioduring the week visited. When the sexes were analyzed separately,none of the weekly variables significantly discriminated femalereturn, but sex ratio was positively associated with male return.Regression showed that the number of transient birds returningin subsequent years was positively associated with the numberof male breeders at our study site during the year a bird visited.Percentage return the year after transience was positively associatedwith the number of eggs laid at our study site during the yeara bird visited. When sexes were analyzed separately, the higherthe number of female breeders during the year a bird visited,the greater the number of males returning in subsequent years,and greater numbers of breeding males were positively associatedwith transient female return. Based on our results, we suggestthat transient birds were searching for better breeding areasfor future breeding and that intrasexual competition made thisinformation more important to females than to males.     

Survival of cohorts in a fluctuating population of the vole Microtus townsendii

Survival patterns of cohorts are described during a population cycle of the vole Microtus townsendii near Vancouver, British Columbia, Canada. A two–year live–trapping study on both enclosed and unfenced populations showed that cohorts during the increase phase of growth lived longest and had the best survival. Smaller voles in the peak density spring cohort had poor survival, but survival increased during the peak density summer. Survival of cohorts in the decline phase breeding season was very poor. The suggestions are made that changes in spacing behaviour may cause changes in cohort survival and that the causes of rapid changes in survival need to be determined.     

Adult survival and numbers in a coastal breeding population of Redshank Tringa totanus in northwest England

The results of a long-term capture-mark-recapture ringing programme carried out on a coastal population of breeding Redshanks Tringa totanus between 1974 and 1988 are presented.
Both sexes were equally likely to be recaptured in subsequent years, as were birds captured for the first time compared with those that had been captured previously. Older birds were more frequently recaptured than were young birds. There was no significant difference in male and female adult survival rates, with a mean of 72% of females and 75% of males surviving each year.
The breeding population fluctuated annually with estimated breeding densities of 122–285 pairs per km². Variation in breeding numbers (males), but not survival, was partially attributable to winter air temperature.     

The challenges of conservation for declining migrants: are reserve‐based initiatives during the breeding season appropriate for the Pied Flycatcher Ficedula hypoleuca?

Creating conservation policies for declining migrant species in response to global change presents a considerable challenge. Migrant species are affected by factors at breeding grounds, overwintering areas and during migration. Accordingly, reserve-based management during the breeding season is not always a suitable conservation strategy. Recent Pied Flycatcher population decline typifies the pattern for many migrants. The UK population has declined by 43% in the past decade, but explanations, and possible solutions, remain elusive. We use 15 years of data (1990–2004) from a declining British population to establish possible reasons for decline, considering: (1) breeding performance (including the influences of competition and predation); (2) weather patterns caused by the winter phase (December–March) of the North Atlantic Oscillation (NAO), which modify conditions experienced at wintering grounds and on migration; and (3) possible impacts of climate change on spring temperatures. We conclude that decreasing breeding performance is contributing to decline, but that non-breeding factors are more important. Winter NAO index is a strong predictor of breeding population, probably because it influences food abundance in Africa and at migratory stopover points. Importantly, however, year itself enhances the predictive model, indicating that influences on population remain unaccounted for by current research. Management strategies based on increasing breeding productivity cannot fully address population decline because non-breeding factors appear important. However, as breeding performance is declining, breeding-based strategies remain useful conservation tools. To this end, our research indicates that optimal placement of nestboxes as regards orientation and habitat management to increase larval food supplies could increase productivity significantly.     

Numbers and breeding success of Capercaillies Tetrao urogallus and Black Grouse T. tetrix at Abernethy Forest,Scotland

Capsule In most years, changes in numbers are associated with variations in breeding success.

Aims To describe the annual variation in numbers and breeding success of Capercaillies Tetrao urogallus and Black Grouse Tetrao tetrix at Abernethy Forest, and their inter‐relationships.

Methods Numbers and breeding success of Capercaillies and Black Grouse were recorded annually at Abernethy Forest (a Scots Pine Pinus sylvestris forest comprising ancient native, or semi‐natural, pinewood and plantations) during 1989–2009. Indices of abundance and densities of Capercaillies were obtained along transects, while counts of males at leks were obtained for both species. Breeding success (number of chicks per female) was obtained using dogs to locate females and chicks.

Results Capercaillie: the index of abundance increased to a peak in winter 1996/97 (2.7 times greater than in 1992/93) and then declined. There was no long‐term trend. In winters 2003/04 and 2004/05, the mean density was 4.2 per km² (95% CLs 3.1–5.7). The total number was 140 (95% CLs 100–220) for 2003/04 and 170 (95% CLs 110–280) for 2004/05, comprising about 8% of the Scottish population. The number of males at leks peaked in 1995 (to 46 males) and again in 2004 (41 males), and there was no long‐term trend. The annual breeding success varied from 0 to 2.93 chicks per female (mean = 0.64). The mean was within the 95% CLs of an independent estimate of the productivity required to maintain numbers. In a free‐running model, annual breeding success and survival rates (which were assumed to improve when mortality owing to fence collisions was removed) largely accounted for the annual variation in the index of abundance, as measured from winter counts along transects during 1990/91 to 2002/03. If mortality associated with collisions with fences had continued, the index would have declined, assuming no immigration.

Black Grouse: The number of male birds at leks increased to a peak in 1997 (to 165 males), before falling back to a smaller number (about 50 males) in the early 2000s. There was a smaller peak in 2007. The annual breeding success varied from 0 to 4.71 chicks per female (mean = 1.76).

Conclusion Numbers of Capercaillies and Black Grouse varied over a 19‐year period at Abernethy Forest, but did not show either upward or downward trends, while the national population of Capercaillies dipped to a low level in 1998/99, and the Black Grouse population continued to decline. In most years, changes in numbers of both species were associated with variation in breeding production. Mortality caused by collisions with fences would have led to a decline in Capercaillie numbers if fences had not been removed.     

Population dynamics of Gyrodactylus sp. (Monogenea) infecting the sand goby in the Oslo Fjord, Norway

Prevalence and abundance of Gyrodactylus sp. on the 1993 year-class of sand gobies Pomatoschistus minutus in the Oslo Fjord, Norway, decreased slightly during the cold months, increased as temperature increased in spring, reaching a peak in June, then decreased significantly, corresponding with a decline in the number of hosts from August. Abundance and, especially, prevalence increased again in September, and decreased in late autumn. Death of the most heavily infected hosts, and/or host resistance might be responsible for the decline in parasite numbers on post-spawners in summer. The young of the year (1994 year-class) were probably infected shortly after they commenced benthic life in July, and prevalence and abundance increased significantly throughout late summer and autumn. Parasite abundance on males was significantly higher than on females at the height of the breeding season in June, but significantly lower at the end of the breeding season in July. Two-thirds of the parasites infected the gills, oral cavity and pharynx. The proportion of parasites on the body, head and fins increased on both sexes, but significantly more so on males, during the breeding season of the host, when parasite intensity was highest. Possible routes of transmission of the parasite are discussed.     

Age variation in a fluctuating population of the common vole

We analysed variation in age in a fluctuating population of the common vole (Microtus arvalis) in southern Moravia, Czech Republic, to test the assumption of the senescence hypothesis that the age of voles increases with increasing population density. Between 1996 and 1998, we monitored the demographic changes by snap-trapping and live-trapping in a field population passing through the increase, peak and decline phase of the population cycle. We used the eye lens mass method to determine the age of snap-trapped animals and those that died in live-traps. The average age of winter males was clearly higher after the peak phase breeding season than before it. No such phase-dependent shift in age, however, was observed in the female component. Male age continued to increase from autumn to spring over the pre-peak winter, and the highest age was in spring of the peak phase year. However, after the peak phase breeding season the highest age was achieved in winter, with the decline phase males during the next spring tending to be younger. The average age of females in spring populations was always lower than in winter populations. The average age of voles from live-traps was always higher than voles from snap-traps, particularly in winter and spring populations, suggesting the presence of senescent animals. Although the density-dependent changes in age are consistent with those observed for other voles, they provide only weak evidence that population cycles in the common vole are accompanied by pronounced shifts in individual age, particularly in female voles.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

The status and recent breeding performance of the critically endangered Northern Bald Ibis Geronticus eremita population on the Atlantic coast of Morocco

The main wild population of the Northern Bald Ibis Geronticus eremita breeds on the Atlantic coast of Morocco in the Agadir region. This paper describes the numbers of breeding pairs over the last two decades, the recent breeding performance, the causes of egg and chick losses and their conservation implications. Since 1980 there has been no overall decline in numbers with, over the last 5 years, 59–74 pairs breeding and a peak prebreeding population of around 220 birds. In contrast with the now extinct Northern Bald Ibis populations in Turkey and elsewhere in Morocco, the birds are present in the Agadir region throughout the year and do not appear to migrate from the area outside the breeding season. Breeding performance is highly variable from one year to the next but does not appear to be related to rainfall in the vicinity of the colonies as has been reported elsewhere. It is suggested that coastal fogs in this region may buffer the adverse impacts of low rainfall and may in part account for the year-round residency of the birds. The main causes of breeding failures have been loss of eggs to predators and, most importantly, poor chick survival as a result of starvation and predation. Conservation action to date has focused on reducing the negative influences on breeding success but it is recognized that for such a long-lived bird adult survival is also likely to be an important limiting factor on the population size.     

Some population dynamics of the Bank vole, Clethrionomys glareolus and the Wood mouse, Apodemus sylvaticus in mixed woodland

The population dynamics of Clethrionomys glareolus Schr.and Apodemus sylvaticus (L.) in mixed woodland in County Durham, were studied from March 1963 to January 1965. Two areas of 0.9 ha (2.25 acres) each were trapped monthly and information was obtained on population size, reproduction, survival and growth from marked animals.
Clethrionomys populations increased from June to an autumn peak; then declined, at first rapidly, but then more slowly in winter, before reaching a spring trough. The breeding season of Clethrionomys was from May to December; juveniles were caught from mid-June to December. Survival was in general poor during the breeding season but good at other times. Survival of young born early in the summer was particularly good on one of the areas and some individuals lived long enough to breed in two successive years. On both areas young born early in the year matured rapidly and bred in the year of their birth. Young born in late summer and early autumn ceased growing at a weight of about 14.5 g, remained immature, and formed the bulk of the overwintering population. Growth was completed at the time of sexual maturation the following spring.
Trapping failed to provide adequate samples of juvenile Apodemus in summer to account for subsequent recruitment. The possibility that a substantial proportion of the adult population of this species was also either trap shy or had emigrated temporarily is discussed. Breeding occurred from April until the following January, but the numbers trapped remained very low throughout the early months of the breeding season. Large scale recruitment of young fecund animals into the trap-revealed population occurred during the autumn. Apodemus males continued to grow rapidly during the winter.     

Habitat loss and raptor predation: disentangling long- and short-term causes of red grouse declines

The number of red grouse (Lagopus lagopus scoticus) shot in the UK has declined by 50% during the 20th century This decline has coincided with reductions in the area of suitable habitat and recoveries in the populations of some avian predators. Here we use long-term records of shooting bags and a large-scale manipulation of raptor density to disentangle the effects of habitat loss and raptor predation on grouse populations. The numbers of grouse harvested on the Eskdale half of Langholm Moor in southern Scotland declined significantly during 1913-1990 and grouse bags from the whole moor from 1950 to 1990 exhibited an almost identical but non-significant trend. Hen harriers (Circus cyaneus) and peregrine falcons (Falco peregrinus) were absent or bred at low densities on this moor throughout this period but heather-dominant vegetation declined by 48% between 1948 and 1988. Harrier and peregrine breeding numbers on Langholm Moor increased to high levels following protection in 1990 whilst grouse density and grouse bags declined year after year until shooting was abandoned in 1998. The prediction of a peak in grouse bags on Langholm Moor in 1996 based on the patterns of bags during 1950-1990 was supported by the observed peaks in 1997 on two nearby moors with few raptors which formerly cycled in synchrony with Langholm Moor. This study demonstrates that, whilst long-term declines in grouse bags were most probably due to habitat loss, high levels of raptor predation subsequently limited the grouse population and suppressed a cycle. This study thus offers support to theoretical models which predict that generalist predators may suppress cycles in prey populations.     

The status of the Corncrake Crex crex in Britain in 1998

A census of Corncrakes throughout Britain was carried out in 1998. National Grid squares of 10 km in the Hebrides, Northern Isles and the mainland of Highland Scotland known to have held the species during the breeding season since 1988 were searched systematically by night for singing males. A sample of such squares elsewhere in Scotland was also searched. In addition, bird-watchers and the general public were encouraged by press publicity to contribute records of singing birds, which were checked for accuracy. A total of 589 singing males was counted in 93 10-km squares. Ninety percent of the population was in the Hebridean Islands. The total count was 23% larger than in the previous census in 1993 and 3% larger than in 1988. All surveys of distribution and censuses between the late 19th century and 1993 recorded a progressive decline in the national population and range. The average rate of population decline between 1988 and 1993 was 3.5% per year, but between 1993 and 1998 the population increased by an average of 4.2% per year. However, annual counts from 1993 to 1998 in core areas that held more than 90% of the population, showed that this overall increase resulted from four successive annual increases of 4–16% followed by a decline of 15% between 1997 and 1998. There was considerable variation among islands and areas in the rate of change of their Corncrake numbers.     

Annual cycle in female minnows Phoxinus phoxinus (L.) from an upland Welsh lake

A population of Phoxinus phoxinus in a small, upland, Welsh lake was sampled at monthly intervals between February 1976 and January 1977. Mature females belonged to the 1972, 1973 and 1974 year classes. Spawning took place in May and June over gravel at the lake's edge. Regression analysis provided estimates of the mean carcase, ovary and liver weights expected for a female of length 65 mm. The ovarian weight increased rapidly in spring but dropped sharply between May and August. Carcase and liver weight declined during the breeding season, but recovered rapidly on completion of breeding. The increase in the weight of the ovary was accompanied by an increase in the proportion of dry matter, while the carcase and liver became relatively hydrated during the breeding season. Also at this time, the energy and lipid content of the carcase was low, but the ash content was high. Thus ovarian maturation and breeding were associated with a depletion of the carcase and liver, but this was quickly reversed at the end of the breeding season. Preliminary estimates of the changes in total energy in the ovary and carcase during ovarian maturation were made which suggested that the decline in the energy content of the carcase represented a significant proportion of the gain in energy content of the ovary. The sampling period coincided with an unusually severe drought and the possible consequences of this on the population are considered.     

AN AFTERNOON WITH THE WADERS

Houston, D. C. 1974. Mortality of the Cape Vulture. Ostrich 45: 57–62.

An attempt was made to analyse the ringing returns of the Cape Vulture Gyps coprotheres from Transvaal colonies. Although it is not yet possible to determine annual mortality rates, some conclusions can be deduced from the results. Mortality during the early age classes is high and is over 500; during the first year. Mortality must decline rapidly with age, but it seems probable that recruitment to the breeding population at these colonies is insufficient to balance adult mortality and the population may be declining.     

Demography of two lemming species on Bylot Island, Nunavut, Canada

Lemmings play a key role in the tundra food web and their widely reported cyclic oscillations in abundance may have a strong effect on other components of the ecosystem. We documented seasonal and annual variations in population density, reproductive activity, survival, and body mass of two sympatric species, the brown (Lemmus trimucronatus) and collared lemmings (Dicrostonyx groenlandicus), over a 2-year period on Bylot Island, Nunavut, Canada. We live trapped and marked lemmings on two grids throughout the summer and we estimated demographic parameters using three different capture–recapture methods. All three methods are based on robust estimators and they yielded similar population density estimates. The density of brown lemmings declined markedly between the 2 years whereas that of collared lemmings was relatively constant. For brown lemmings, 2004 was a peak year in their cycle and 2005 a decline phase. Density of brown lemmings also decreased during the summer, but not that of collared lemmings. The recruitment of juvenile brown lemmings in the population increased during the summer and was higher in the peak year than in the year after, but no change was detected in collared lemmings. Survival rates of both species tended to be lower during the peak year than in the following year and body mass of brown lemmings was higher in the peak year than in the following year. We conclude that both changes in adult survival and juvenile recruitment occur during the population decline of brown lemmings.     

The summer decline of the collared lemming, Dicrostonyx groenlandicus, in high arctic Greenland

I studied the population structure and dynamics of the collared lemming, Dicrostonyx groenlandicus Traill, on Traill Island (NE Greenland) in 1998 to 2000. This is the first comprehensive study of a Greenlandic collared lemming population. During this period, population density continuously declined (>100 fold-decrease) from more than 10 ind/ha (peak year 1998) to less than 0.1 ind/ha (low year 2000), with the exception of one period of increase in the winter 1998–99. Several population parameters were correlated with these changes in density. Average body mass was greater in the peak year than in the intermediate year (1999) and greater in August than in July in both years. Growth rates were only positive in 1998 and only for young lemmings. Females had unexpectedly larger home ranges than males and were more mobile than males during the decline phase (1999). Poor recruitment and high predation rates (up to 3.4% per day for females in 1998) were the proximate causes of the summer declines in the study area. A simple calculation shows that even with a maximal rate of increase, this lemming population would not have been able to compensate for the high predation rate, and hence it necessarily faced a predation-induced summer decline during the part of population oscillation studied.     

Population dynamics of Clethrionomys and Peromyscus in southwestern Yukon 1973-1989

Clethrionomys rutilus and Peromyscus maniculatus occur together in the boreal forest of southwestern Yukon and we studied their population dynamics on unmanipulated live-trapping grids for 17 yr. Peromyscus showed a regular seasonal change in numbers each year with little interannual variation. Clethrionomys by comparison, showed low density fluctuations during one period (1976-1982) followed by changes in number in the next period that are consistent with a typical 3-4 yr microtine cycle. During one of two years of peak numbers adult males had unusually heavy body weights. The sexual maturation of juvenile Clethrionomys varied inversely with density. In most years Clethrionomys showed higher rates of population increase from spring to late summer than Peromyscus and this may be partly attributed to the vole's longer breeding season. Juvenile female Clethrionomys often reached sexual maturity during their first summer when population density was low but Peromyscus never matured during their first summer. Finally, there was no correlation between the two species in year to year changes in overwinter survival rates. This is the first report of a cyclic Norh American population of Clethrionomys and it underlines the need to search for hypotheses of population regulation that explain both annual and multi-annual cycles within the same population.     

On the demography of Rattus sordidus colletti in monsoonal Australia

Results of a mark-release study of Rattus sordidus colletti (Gould) on sub-coastal, treeless plains in the monsoonal north of the Northern Territory of Australia are given for 5 years. R. s. colletti is the dominant component of the small mammal fauna of these plains, with only small numbers of Melomys spp. and Planigale maculata also occurring. Two classes of peak densities were observed. Localized peaks resulted from non-breeding adult R. s. colletti refuging onto the marginally higher levees during flooding of the plains, and also from refuging into a lower-lying area at the end of a dry-season drought. Generalized peak densities resulted from peaks in reproductive effort. In 1972 and 1974, reproduction was confined to the period immediately following the monsoons, but in 1973 breeding continued throughout the dry season, following unseasonal rain in June. Thus, generalized peak densities were observed in one year at the beginning of the dry season, and in another at its end. The relatively shallower flooding and mildness of the 1972–3 wet season resulted in commencement of breeding earlier in 1973 than in 1974, while rapid severe flooding of the plains in December 1974 resulted in mass mortality and failure of the refuging populations of adults to recolonize the lower plains in 1975. R. s. colletti remained extremely rare until November 1976, when the study was terminated. The good conditions which allowed reproduction during the dry season of 1973 were reflected in rapid growth rates at that time, compared with rates approaching zero in the 1974 dry season. R. s. colletti is the most fecund form of Australian Rattus species. This high fecundity appears to compensate for the restriction, by dry-season aridity and wet-season flooding, of the breeding season in most years to a short period immediately following the monsoons. In unusual years with rainfall during the dry season, the high fecundity gives R. s. colletti populations the capacity to reach very high densities.     

Is the large‐scale decline of the starling related to local changes in demography?

The decline of one farmland bird, the migratory European starling, has been attributed to both agricultural intensification and farmland abandonment and to factors operating both during the winter and during the breeding season. We analysed population data from thirty‐three Swedish nestbox colonies over more than two decades to determine if the national decline was caused by a common factor affecting all colonies or by local changes in the breeding grounds affecting starling colonies. We found that numbers of breeding starling had declined significantly, but at different rates in different colonies. The local population sizes were affected by previous years’ productivity at both national and local scales, suggesting that changes in habitat quality at both scales could affect local population trends. There were no long‐term trends in reproductive output, but fledgling production was lowest at intermediate years. The local population changes were positively related to local changes in reproductive output, but only when including complete nest‐failures. A relationship between population declines and low mean local productivity was the result of the association between population sizes and reproductive success over time, since decline rates of starlings were not related to the average success during the first part of the study, but to the average success during the later part of the study. The relationship between population change and changes in reproductive output was evident, but fledgling production showed negative density‐dependence. In conclusion this study suggests that the decline of the starling population in Sweden has been affected by processes at small spatial scales during the breeding season affecting reproductive success, but does not exclude an additional role for processes at large spatial scales or outside the breeding season.     

Population regulation and dispersion of the smoky mouse,Pseudomys fumeus II. Spring decline,breeding success and habitat heterogeneity

The smoky mouse (Pseudomys fumeus) undergoes large annual fluctuations in abundance, characterized by a dramatic decline in numbers just before the summer breeding season. The population on Mount William has two components which are differentially affected by this decline. The first, consisting of animals living in preferred microhabitat, have high breeding success and are unaffected by the decline. Females normally breed in two seasons, and in the second season breed earlier than first year females. The second component lives in non-preferred microhabitat, is severely affected by the decline and shows high levels of fighting during the decline period. This difference may result from the lack of high quality food resources in non-preferred microhabitat following the seasonal cessation of production of fungal sporocarps. The role of spatial and temporal heterogeneity in the availability of food resources in the population processes of small mammals is discussed.     

Estimation and limitation of numbers of floaters in a Eurasian Sparrowhawk population

Over a 20‐year period, the numbers of Eurasian Sparrowhawk Accipiter nisus nests found in a 200 km² area in south Scotland remained relatively stable (mean 33.3 pairs, CV = 10.6%). Nest numbers fluctuated from year to year in a manner expected of a population subject to density‐dependent regulation. The numbers of non‐breeders (floaters) could not be counted directly, but the number of female floaters was estimated, using known mortality rates, from the numbers of females recruited to the breeding population each year at different ages. Female floater numbers were estimated by two methods: Method A assumed that birds bred for the first time in their first, second or third year, in the same ratio as they were found breeding for the first time in the study area; and Method B assumed that all third‐year birds found breeding for the first time in the study area had bred previously, unknown to us, outside the area. Under Method A, floaters consisted of some 1‐year and some 2‐year birds; while under Method B, floaters consisted only of some 1‐year birds. Under both methods, the estimated number of female floaters fluctuated greatly from year to year, but under Method A they averaged 0.90 per female breeder, and under Method B they averaged 0.28 per female breeder. The Method B estimate was most consistent with other data and with the finding that some birds found breeding in the study area were likely to have bred previously outside the area (because of territory changes). Moreover, the mean and variance in female floater numbers estimated by Method B were similar in magnitude to the values obtained in a simulation model. In this model, breeding density was given a fixed ceiling, while breeding success was allowed to vary from year to year within the limits observed in the study area. It was concluded that (a) recruitment of floaters to the breeding sector was density dependent with respect to breeder numbers, i.e. broadly speaking, floaters filled gaps in the territorial system left by the deaths and movements of established breeders; and (b) floater numbers themselves were probably not regulated in a density dependent manner, but depended on whatever was the balance between annual additions (from reproduction and immigration) and subtractions (from mortality, emigration and entry to the breeding sector).