When and where to count? Implications of migratory connectivity and nonbreeding distribution to population censuses in a migratory bird population

Migratory connectivity is a metric of the co-occurrence of migratory animals originating from different breeding sites, and like their spatial dispersion, can vary substantially during the annual cycle. Together, both these properties affect the optimal times and sites of population censusing. We tracked taiga bean geese (Anser fabalis fabalis) during 2014–2021 to study their migratory connectivity and nonbreeding movements and determine optimal periods to assess the size of their main flyway population. We also compared available census data with tracking data, to examine how well two existing censuses covered the population. Daily Mantel's correlation between breeding and nonbreeding sites lay between 0 and 0.5 during most of the nonbreeding season, implying birds from different breeding areas were not strongly separated at other times in the annual cycle. However, the connectivity was higher among birds from the westernmost breeding areas compared to the birds breeding elsewhere. Daily Minimum Convex Polygons showed tracked birds were highly aggregated at census times, confirming their utility. The number of tracked birds absent at count sites during the censuses however exceeded numbers double-counted at several sites, indicating that censuses might have underestimated the true population size. Our results show that connectivity can vary in different times during the nonbreeding period, and should be studied throughout the annual cycle. Our results also confirm previous studies, which have found that estimates using marked individuals usually produce higher population size estimates than total counts. This should be considered when using total counts to assess population sizes in the future.     

THE SYSTEMATIC POSITION OF THE SOUTH AFRICAN GANNET

Summary

Randall, R. M., Randall, B. M., Cooper, J. &; Frost, P. G. H. 1986: A new census method for penguins tested on Jackass Penguins Spheniscus demersus. Ostrich 57: 211–215

A census method for Jackass Penguins based on their moulting cycle is described. Population estimates were substantially higher with this method than with total counts, but were not dissimilar to maximum estimates obtained from strip counts. The moult method estimates the size of the entire population and is independent of the breeding cycle and breeding status of both individuals and population. It is not plagued by the problem of absenteeism encountered during most phases of the penguin annual cycle and is independent on their diel activity. The drawbacks to the method are that it is laborious and protracted, that it does not differentiate between the breeding population and the whole population, and is unsuited to some species and some habitats.     

A statistical procedure for estimating Grey seal pup production from a single census

The variable start and duration of the Grey seal breeding season makes the estimation of total pup production from a single census very difficult. Classifying the count into morphological age classes enables the form and timing of the birth rate curve and estimates of pup mortality rates to be elucidated. A simulation technique is described which enables the duration of each morphological stage to be determined from a series of such classified counts taken over one season. A further statistical technique uses these estimates to calculate the mean timing and duration of the breeding season from a single classified count taken from similar populations in subsequent years. This information allows total pup production to be calculated for any appropriate breeding colony. Some guidance is given as to the optimal timing of that single census which would yield the best estimate of production, although the precise date is not critical to the success of the technique. Results from single census estimates obtained in this way are compared with known production data from more detailed surveys for a number of different colonies.     

Nest and breeding population abundance of Least Terns: assessing bias and variation in survey timing and methods

Methods commonly used to estimate the number of nests and size of the breeding population at colonies of Least Terns (Sternula antillarum) and other waterbirds include walk‐through counts of nests (ground‐nest counts) and counts of incubating adults from the colony perimeter (incubating‐adult counts). The bias and variance of different methods and the comparability of repeated surveys versus once‐annual censuses are poorly understood. Our objectives were to assess (1) the potential bias and variation of the more rapid incubating‐adult counts compared to the time‐intensive, and presumably more accurate, ground‐nest counts, and (2) how accurately a once‐annual census captured peak nesting abundance. We studied nine Least Tern colonies at Cape Lookout National Seashore (CALO), North Carolina, from April to August 2010–2012. We analyzed observer and survey method agreement with concordance correlation coefficients (ρ_c). We deployed time‐lapse cameras at 156 nests and used repeated‐measures logistic regression to determine if the proportion of time spent incubating varied with colony, time of day, or time of season. We found substantial agreement in abundance estimates of Least Tern nests and incubating adults between observers and survey methods, and among different times of day and seasons (all comparisons ρ_c > 0.97). Least Terns incubated eggs 94% of the time on average during daylight hours, irrespective of colony, nesting stage, or month. Although the nesting peak at CALO occurred during the recommended census period for Least Terns, abundance estimates for surveys conducted at different times during that period varied by as much as 39%. We recommend conducting incubating‐adult counts to estimate nest and breeding population abundance of Least Terns or other waterbirds when vegetation or dunes do not obstruct views of nesting colonies. In addition, given the variation in abundance estimates for surveys conducted at different times during the recommended survey period, incubating‐adult counts should be performed at least twice during the census period, with the maximum count reported as peak nest abundance.     

Effects of re‐breeding rates on population size estimation of biennial breeders: results from a model based on albatrosses

Estimating total breeding populations (I) for species that exhibit biennial breeding is generally done from counts of individuals that breed in each year (N), but can be complicated by the fact that the proportion of individuals breeding varies from year to year. Partly, this reflects the proportion of individuals that re‐breed in successive years (re‐breeding rate, p), which is largely, although not exclusively, governed by reproductive failure. Here we show that variation in counts of breeding individuals not only reflects changes in total breeding population but can be sensitive to and powerfully driven by variation in p. A simulation of annual field counts of a bird exhibiting biennial breeding was constructed to explore the effect of re‐breeding attempts on estimations of the total breeding population. The model was used to simulate the consequences of adult mortality and different annual patterns of nesting failures on total breeding population estimates, and to explore the consequences of variation in p on N, when total breeding population remains constant. N is shown to be very sensitive to variations in p, so that even short‐term fluctuations in p can cause changes in N that oscillate for many years ahead. We compare our modelled results with real data for Grey‐headed Albatrosses Thalassarche chrysostoma and demonstrate that, when I is held constant in the model, actual counts may be simulated by variations in p only. Normally, I is unknown and is extrapolated from N on the assumption that N mirrors changes in the size of the total population. Consequently, applying average values of p can result in misleading estimates of total breeding population. We recommend that annual counts of breeding individuals are supplemented with annual estimates of p. Field protocols that aim to estimate annual breeding population size from counts of breeding individuals should be complemented by independent measures of rates of re‐breeding and nest failure.     

Effective number of breeding adults in Bufo bufo estimated from age-specific variation at minisatellite loci

Estimates of the effective number of breeding adults were derived for three semi-isolated populations of the common toad Bufo bufo based on temporal (i.e. adult-progeny) variance in allele frequency for three highly polymorphic minisatellite loci. Estimates of spatial variance in allele frequency among populations and of age-specific measures of genetic variability are also described. Each population was characterized by a low effective adult breeding number ( N _b) based on a large age-specific variance in mini-satellite allele frequency. Estimates of N _b (range 21–46 for population means across three loci) were ≊ 55–230-fold lower than estimates of total adult census size. The implications of low effective breeding numbers for long-term maintenance of genetic variability and population viability are discussed relative to the species' reproductive ecology, current land-use practices, and present and historical habitat modification and loss. The utility of indirect measures of population parameters such as N _b and N _e based on time-series data of minisatellite allele frequencies is discussed relative to similar measures estimated from commonly used genetic markers such as protein allozymes.     

The genetic effective and adult census size of an Australian population of tiger prawns (Penaeus esculentus)

This study compares estimates of the census size of the spawning population with genetic estimates of effective current and long-term population size for an abundant and commercially important marine invertebrate, the brown tiger prawn (Penaeus esculentus). Our aim was to focus on the relationship between genetic effective and census size that may provide a source of information for viability analyses of naturally occurring populations. Samples were taken in 2001, 2002 and 2003 from a population on the east coast of Australia and temporal allelic variation was measured at eight polymorphic microsatellite loci. Moments-based and maximum-likelihood estimates of current genetic effective population size ranged from 797 to 1304. The mean long-term genetic effective population size was 9968. Although small for a large population, the effective population size estimates were above the threshold where genetic diversity is lost at neutral alleles through drift or inbreeding. Simulation studies correctly predicted that under these experimental conditions the genetic estimates would have non-infinite upper confidence limits and revealed they might be overestimates of the true size. We also show that estimates of mortality and variance in family size may be derived from data on average fecundity, current genetic effective and census spawning population size, assuming effective population size is equivalent to the number of breeders. This work confirms that it is feasible to obtain accurate estimates of current genetic effective population size for abundant Type III species using existing genetic marker technology.     

Population viability analysis of Walia ibex (Capra walie) at Simien Mountains National Park (SMNP), Ethiopia

Population viability analysis (PVA) has been applied to the management of many threatened populations. The objective of this study was, therefore, to estimate the PVA of Walia ibex at the Simen Mountains National Park, in the north‐central highlands of Ethiopia, with respect to population growth parameters, the probability of the population reaching a lower extinction threshold and the mean time to extinction. Direct census of the population was carried out in 2009. Secondary census data were also collected from park authorities and the literature reviews. The result revealed that the estimates of the infinitesimal mean, μ (0.04117) was greater than the infinitesimal variance, σ² (0.0219). The probability that the population reaches the extinction threshold was very low (0.15%). The mean time required for the counts to decline from the existing population size to one individual animal was 160 years. But threatened species are adversely affected by changes in landscape. These changes can be brought by short‐ and long‐term human and climate change impacts, respectively. Therefore, with the absence of environmental and demographic stochasticity and, with the application of appropriate reproductions and habitat management, the population of Walia ibex will be viable and reaches its mean time of extinctions after 160 years.     

What affects the magnitude of change in first arrival dates of migrant birds?

We analysed which among four factors (mean first arrival date, migration distance, changes in population size, detectability of species) influenced the magnitude of change (regression coefficient) in the first arrival dates of 30 migrant bird species in western Poland during 1983–2003. An examination suggested that several of these factors could be important: the regression coefficient was positively related to mean first arrival date (early species advancing their arrival date more) and negatively with change in population size (species in decline changing less). Moreover, significant differences in regression coefficient were detected between short and long distance migrants and between low detectable and highly detectable species. Undertaking a principal components analysis on the four factors produced an axis explaining 59% of the variance and whose positive values were associated with late arriving, long distance and low detectable species which were more likely to be in decline. However, the multi-collinearity of these factors is a problem that cannot be resolved here and we recommend that further work from different areas is needed to tease apart these effects.     

Is the detection of the first arrival date of migrating birds influenced by population size? A case study of the red-backed shrike Lanius collurio

Many analyses do not consider the problems associated with the effects of population size on encounter recording. Population size could impact on the detection of bird arrival time as there is a higher probability of observing earlier arrival when the population size is greater and the song activity of birds is increased, as occurs with a larger population. As a case study, we have analysed data on the red-backed shrike Lanius collurio collected in Western Poland during 1983–2000. In this period the red-backed shrike’s return to its breeding sites became significantly earlier whilst the contemporary population size increased significantly. To eliminate linear trends through time we have worked on the standardised residuals from regression of both arrival time and population size on year. The correlation between arrival time and population size residuals was significantly negative, further supporting the link between detection and population size. This finding suggests that, in studies of avian migration and its changes over time, the relationship between arrival date and population size needs to be considered. Received: 25 October 2000 / Revised: 5 September 2001 / Accepted: 5 September 2001     

Effective Sizes for Subdivided Populations

Many derivations of effective population sizes have been suggested in the literature; however, few account for the breeding structure and none can readily be expanded to subdivided populations. Breeding structures influence gene correlations through their effects on the number of breeding individuals of each sex, the mean number of progeny per female, and the variance in the number of progeny produced by males and females. Additionally, hierarchical structuring in a population is determined by the number of breeding groups and the migration rates of males and females among such groups. This study derives analytical solutions for effective sizes that can be applied to subdivided populations. Parameters that encapsulate breeding structure and subdivision are utilized to derive the traditional inbreeding and variance effective sizes. Also, it is shown that effective sizes can be determined for any hierarchical level of population structure for which gene correlations can accrue. Derivations of effective sizes for the accumulation of gene correlations within breeding groups (coancestral effective size) and among breeding groups (intergroup effective size) are given. The results converge to traditional, single population measures when similar assumptions are applied. In particular, inbreeding and intergroup effective sizes are shown to be special cases of the coancestral effective size, and intergroup and variance effective sizes will be equal if the population census remains constant. Instantaneous solutions for effective sizes, at any time after gene correlation begins to accrue, are given in terms of traditional F statistics or transition equations. All effective sizes are shown to converge upon a common asymptotic value when breeding tactics and migration rates are constant. The asymptotic effective size can be expressed in terms of the fixation indices and the number of breeding groups; however, the rate of approach to the asymptote is dependent upon dispersal rates. For accurate assessment of effective sizes, initial, instantaneous or asymptotic, the expressions must be applied at the lowest levels at which migration among breeding groups is nonrandom. Thus, the expressions may be applicable to lineages within socially structured populations, fragmented populations (if random exchange of genes prevails within each population), or combinations of intra- and interpopulation discontinuities of gene flow. Failure to recognize internal structures of populations may lead to considerable overestimates of inbreeding effective size, while usually underestimating variance effective size.     

ICE-BASED CENSUS OF BOWHEAD WHALES MIGRATING PAST POINT BARROW, ALASKA, 1978-1983

Bowhead whale ( Balaena mysticetus ) census data obtained during the northward spring migration are summarized for 1978–1983. Population size estimates are derived from counts made by observers standing on the seaward edge of shorefast ice in the vicinity of Point Barrow, Alaska, from mid-April to early June. The research design utilized two counting stations: South Perch, the primary counting station, and North Perch, used to determine the number of whales missed by South Perch observers. The percentage missed is estimated for each visibility category and used here to correct the census counts. Each season's population estimate is calculated as the sum of the number of trials of several independent multinomial distributions representing different visibility conditions. Corrections are applied for unwatched hours and hours with inferior visibility. A mean estimate of the number of whales passing within view of the census station was computed as 3,674 ± 299. This estimate was based on data collected in 1978, 1981, 1982, and 1983, years with the least apparent biases. Aerial survey data provide estimates of the proportion of whales passing at various distances seaward of the census sites as follows: 0.58 from the ice edge to 2 km, 0.76 to 3 km, and 0.80 to 4 km. Correcting for whales too far offshore to be seen by the ice-based observers results in a population estimate of over 4,200 bowheads.     

Temporal genetic variation of mitochondrial DNA and the female effective population size of red drum (Sciaenops ocellatus) in the northern Gulf of Mexico

We studied genetic drift of mitochondrial DNA (mtDNA) haplotype frequencies in a natural population of red drum (Sciaenops ocellatus) from the northern Gulf of Mexico (Gulf). The amount of genetic drift observed across temporally adjacent year classes (1986–89) was used to estimate variance effective (female) population size (N_ef). N_ef was estimated to be 14 308 and the ratio of female effective size to adult female census size was approximately 0.004, which is among the lowest value reported for vertebrate animals. Low effective size relative to census size among red drum in the northern Gulf may result from yearly fluctuations in the number of breeding females, high variance in female reproductive success, or both. Despite low genetic effective size relative to census size, the genetic effective population size of red drum in the northern Gulf appears sufficiently large to preclude potentially deleterious effects of inbreeding.     

Genetic diversity, population structure, effective population size and demographic history of the Finnish wolf population

The Finnish wolf population (Canis lupus) was sampled during three different periods (1996-1998, 1999-2001 and 2002-2004), and 118 individuals were genotyped with 10 microsatellite markers. Large genetic variation was found in the population despite a recent demographic bottleneck. No spatial population subdivision was found even though a significant negative relationship between genetic relatedness and geographic distance suggested isolation by distance. Very few individuals did not belong to the local wolf population as determined by assignment analyses, suggesting a low level of immigration in the population. We used the temporal approach and several statistical methods to estimate the variance effective size of the population. All methods gave similar estimates of effective population size, approximately 40 wolves. These estimates were slightly larger than the estimated census size of breeding individuals. A Bayesian model based on Markov chain Monte Carlo simulations indicated strong evidence for a long-term population decline. These results suggest that the contemporary wolf population size is roughly 8% of its historical size, and that the population decline dates back to late 19th century or early 20th century. Despite an increase of over 50% in the census size of the population during the whole study period, there was only weak evidence that the effective population size during the last period was higher than during the first. This may be caused by increased inbreeding, diminished dispersal within the population, and decreased immigration to the population during the last study period.     

The consequences of the variance-mean rescaling effect on effective population size

The effective population size (N_e), and the ratio between N_e and census population size (N) are often used as measures of population viability. We show that using the harmonic mean of population sizes over time – a common proxy for N_e– has some important evolutionary consequences and implications for conservation management. This stems from the fact that there is no unambiguous relationship between the arithmetic and harmonic means for populations fluctuating in size. As long as the variance of population size increases moderately with increasing arithmetic mean population size, the harmonic mean also increases. However, if the variance of population size increases more rapidly, which existing data often suggest, then the harmonic mean may actually decrease with increasing arithmetic mean. Thus maximizing N may not maximize N_e, but could instead lower the adaptive potential and hence limit the evolutionary response to environmental change. Large census size has the clear advantage of lowering demographic stochasticity, and hence extinction risk, and under certain conditions large census size also minimizes the loss of genetic variation. Consequently, maximising census size has served as a useful dogma in ecology, genetics and conservation. Nonetheless, due to the intricate relationships among N_e, population viability and the properties of population fluctuations, we suggest that this dogma should be taken only as a rule of thumb.     

Arrival date, age and breeding success in white stork Ciconia ciconia

Early arrival to breeding grounds is a life history trait in birds that can result in fitness benefits. We studied the relationship between arrival date and breeding success of individuals in a central Iberian population of white stork Ciconia ciconia , between 1999 and 2005, and the ways in which other potential factors, such as age or sex, affect this relationship. Our results showed that age was the factor most closely related to arrival date and breeding success. Older individuals returned earlier to the breeding grounds, achieved larger clutch sizes and produced more chicks than younger birds. After controlling statistically for age effect, breeding probability (laid eggs or not) and laying date were still significantly explained by arrival date. A higher probability of failure to reproduce (no eggs laid) was found in birds arriving later than in those arriving early. However, clutch size and nestling success (number of nestlings in the nest 40 days after hatching) were not correlated with arrival date. Food availability in the study area throughout the breeding cycle, due to a nearby rubbish dump, could be the factor mitigating differences in clutch size and nestling success related to individual arrival date.     

Recruitment rates in forest plots: Gf estimates using growth rates and size distributions

1 In censuses of tree populations in permanent plots, short census intervals and small population size lead to uncertainty in the observed recruitment rate of a minimum size. Increasing the census interval, however, underestimates the rate because of unrecorded 'recruit and die' events.
2 We propose a new Gf procedure for estimation of recruitment rates. Recruitment rate per area is obtained by multiplication of the density in the smallest size class (f) and the average size growth rate in that class (G) divided by the width of the class. This procedure is valid when the size distribution of the population examined is continuous with size.
3 When tree size structure is negative‐exponentially distributed, as is often the case in natural rain forest populations, the Gf estimate of the recruitment rate for a given size class was least biased close to the midpoint size of this class.
4 Gf estimates agreed well with census estimates of recruitment rate from permanent plots in rain forests. A tendency for Gf estimates to be larger than census estimates disappeared when census estimates were corrected for mortality after recruitment.
5 The effects of plot size, census interval and variation in growth rate on estimates of recruitment rate were simulated using model populations. Small plot size caused substantially more among‐plot deviation for the census count of recruitment events than for the Gf estimate. The census recruitment rate also showed larger variation among plots for shorter intervals than the Gf estimate, which was independent of census interval. The Gf estimates were therefore more accurate than census counts in many situations. More than several tens of trees were needed in a size class to allow a reliable Gf estimates.     

Comparison of census methods for black-browed albatrosses breeding at the Ildefonso Archipelago,Chile

A breeding population of black-browed albatrosses has been known to exist at the Ildefonso Archipelago, Chile, for >90 years but the population has never been censused using scientifically defendable methods. To estimate population size, and examine the accuracy and practicality of various census methods, the population was censused in the 2002/2003 breeding season using (a) ground-truthed aerial photography, (b) yacht-based photography, (c) ground counts, (d) quadrat sampling and (e) point-distance sampling. Compared to ground-truthed aerial photography (judged the most accurate) yacht-based photography underestimated population size by 55%, ground counts by 13%, quadrat sampling by 11% and point-distance sampling by 9%. Ground-truthed air photography revealed that in the 2002/2003 breeding season 47,000 pairs of black-browed albatrosses bred at the Ildefonso Archipelago. A repeat aerial census in 2006 suggested the size of the breeding population had not changed in the 4 years between the two censuses. After the Falkland Islands/Islas Malvinas, South Georgia and Diego Ramirez, the Ildefonso Archipelago holds the fourth largest population of black-browed albatrosses in the world.     

Adjustment of the annual cycle to climatic change in a long-lived migratory bird species

Climate change has advanced the phenology of many organisms. Migratory animals face particular problems because climate change in the breeding and the wintering range may be asynchronous, preventing rapid response to changing conditions. Advancement in timing of spring migration may have carry-over effects to other parts of the annual cycle, simply because advancement of one event in the annual cycle also advances subsequent events, gradually causing a general shift in the timing of the entire annual cycle. Such a phenotypic shift could generate accumulating effects over the years for individuals, but also across generations. Here we test this novel hypothesis of phenotypic response to climate change by using long-term data on the Arctic tern Sterna paradisaea. Mean breeding date advanced by almost three weeks during the last 70 years. Annual arrival date at the breeding grounds during a period of 47 years was predicted by environmental conditions in the winter quarters in the Southern Ocean near the Antarctic and by mean breeding date the previous year. Annual mean breeding date was only marginally determined by timing of arrival the current year, but to a larger extent by arrival date and breeding date the previous year. Learning affected arrival date as shown by a positive correlation between arrival date in year (i+1) relative to breeding date in year (i) and the selective advantage of early breeding in year (i). This provides a mechanism for changes in arrival date being adjusted to changing environmental conditions. This study suggests that adaptation to changing climatic conditions can be achieved through learning from year to year[Current Zoology 55(2):92-101,2009].     

Pre-incubation feeding activities and energy budgets of Snow Geese: can food on the breeding grounds influence fecundity?

The potential contribution of early spring feeding conditions in the Arctic to clutch size variation was examined in a population of Lesser Snow Geese Anser caerulescens caerulescens. Behavioural observations were combined with energetic analyses of food material to derive an estimate of the energy budgets of pre-laying and laying females. Food intake of females between arrival on the breeding grounds and incubation was considerable; estimated energy gains in this period were in the same magnitude as the cost of one or several eggs. The pre-laying period spent on the breeding grounds can thus be energetically beneficial rather than costly. Accumulation of resources for reproduction in Snow Geese is a continual process including the breeding grounds, and nutrient limitation after arrival in the Arctic cannot sufficiently explain the environmental component of clutch size variation. The timing of migration and follicle development is such that clutch size decisions are sometimes made during the late stages of migration and some-times after arrival. In the latter case food conditions on the breeding grounds may greatly influence clutch size; in the former case they may still influence readjustments of clutch size after the initial decision. The universal negative correlation between clutch size and laying date in Snow Geese can be explained by negative fitness consequences of late hatching, which outweigh the benefits of delayed laying and further nutrient accumulation. Food shortage on the breeding grounds may sometimes be a secondary factor contributing to seasonal clutch size decline.