Are Declines in North American Insectivorous Songbirds Due to Causes on the Breeding Range?

Previous studies have reported a recent decline in breeding populations of migratory songbirds in eastern and central North America. Several explanations have been suggested: deforestation on the wintering grounds in the tropics and habitat loss, increased predation pressure, and increased cowbirds parasitism on the breeding range. We used these factors to assign 47 species of insectivorous passerines to groups with contrasting vulnerability, and then used the North America Breeding Bird Survey to analyze population trends in these groups on a large continental scale. Variables indexing susceptibility to predation on the breeding ground were most strongly correlated with population trends form 1968 to 1987. During the period from 1978 to 1987, migratory status was also significantly associated with population trends long-distance migrants to the neotropics exhibited a small, nonsignificant decreasing trend, whereas residents and short-distance migrants increased strongly. During the same time period, the group of species with low nest location, open nest, and high cowbird parasitism declined significantly. Although it is difficult to separate the effects of multiple factors, our analyses suggest that predation on the breeding ground in North America has played a larger role in the decline of migratory songbirds than deforestation on the wintering grounds in the tropics.     

Changes in size and trends of North American sea duck populations associated with North Pacific oceanic regime shifts

Broad-scale multi-species declines in populations of North American sea ducks for unknown reasons is cause for management concern. Oceanic regime shifts have been associated with rapid changes in ecosystem structure of the North Pacific and Bering Sea. However, relatively little is known about potential effects of these changes in oceanic conditions on marine bird populations at broad scales. I examined changes in North American breeding populations of sea ducks from 1957 to 2011 in relation to potential oceanic regime shifts in the North Pacific in 1977, 1989, and 1998. There was strong support for population-level effects of regime shifts in 1977 and 1989, but little support for an effect of the 1998 shift. The continental-level effects of these regime shifts differed across species groups and time. Based on patterns of sea duck population dynamics associated with regime shifts, it is unclear if the mechanism of change relates to survival or reproduction. Results of this analysis support the hypothesis that population size and trends of North American sea ducks are strongly influenced by oceanic conditions. The perceived population declines appear to have halted >20 years ago, and populations have been relatively stable or increasing since that time. Given these results, we should reasonably expect dramatic changes in sea duck population status and trends with future oceanic regime shifts.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Monarch butterfly migration and parasite transmission in eastern North America

Seasonal migration occurs in many animal systems and is likely to influence interactions between animals and their parasites. Here, we focus on monarch butterflies (Danaus plexippus) and a protozoan parasite (Ophryocystis elektroscirrha) to investigate how host migration affects infectious disease processes. Previous work showed that parasite prevalence was lower among migratory than nonmigratory monarch populations; two explanations for this pattern are that (1) migration allows animals to periodically escape contaminated habitats (i.e., migratory escape), and (2) long-distance migration weeds out infected animals (i.e., migratory culling). We combined field-sampling and analysis of citizen science data to examine spatiotemporal trends of parasite prevalence and evaluate evidence for these two mechanisms. Analysis of within-breeding-season variation in eastern North America showed that parasite prevalence increased from early to late in the breeding season, consistent with the hypothesis of migratory escape. Prevalence was also positively related to monarch breeding activity, as indexed by larval density. Among adult monarchs captured at different points along the east coast fall migratory flyway, parasite prevalence declined as monarchs progressed southward, consistent with the hypothesis of migratory culling. Parasite prevalence was also lower among monarchs sampled at two overwintering sites in Mexico than among monarchs sampled during the summer breeding period. Collectively, these results indicate that seasonal migration can affect parasite transmission in wild animal populations, with implications for predicting disease risks for species with threatened migrations.     

An unusual trophic subsidy and species dominance in a tropical stream

In classical theory, species are assumed to achieve dominance through competitive exclusion, but if food resources are limiting, cross-habitat trophic subsidies could also underpin dominance. The impact of dominant species on community dynamics may depend on the energy base of population size. We report on an unusual, spatially subsidized population of a tropical, stream-dwelling crab that dominates the benthic fauna of a Kenyan stream. Diet and stable isotope analyses indicated that this crab is a true omnivore, with terrestrial subsidies dominating both plant and animal resources. Unusually, the animal prey included almost no aquatic invertebrates. Instead, a single species of ant constituted approximately 35% of the annual diet (stomach contents analysis) and up to 90% of assimilated nitrogen (estimates from stable isotope analysis). Ants may be pivotal to enabling crab dominance, and this crab may be largely disconnected from the local trophic network for its dietary needs. The paucity of other invertebrates in the stream community suggests that this super-dominant crab is a strong interactor that suppresses aquatic invertebrate populations. Common stabilizing attributes of spatially subsidized food webs (e.g., asynchronous prey availability, wide feeding niche, consumer migration) were absent from this system, and although apparently stable, it may be vulnerable to disturbance in the donor habitat.     

Building a better baseline to estimate 160 years of avian population change and create historically informed conservation targets

Globally, anthropogenic land-cover change has been dramatic over the last few centuries and is frequently invoked as a major cause of wildlife population declines. Baseline data currently used to assess population trends, however, began well after major changes to the landscape. In the United States and Canada, breeding bird population trends are assessed by the North American Breeding Bird Survey, which began in the 1960s. Estimates of distribution and abundance prior to major habitat alteration would add historical perspective to contemporary trends and allow for historically based conservation targets. We used a hindcasting framework to estimate change in distribution and abundance of 7 bird species in the Willamette Valley, Oregon (United States). After reconciling classification schemes of current and 1850s reconstructed land cover, we used multiscale species distribution models and hierarchical distance sampling models to predict spatially explicit densities in the modern and historical landscapes. We estimated that since the 1850s, White-breasted Nuthatch (Sitta carolinensis) and Western Meadowlark (Sturnella neglecta) populations, 2 species sensitive to fragmentation of oak woodlands and grasslands, declined by 93% and 97%, respectively. Five other species we estimated nearly stable or increasing populations, despite steep regional declines since the 1960s. Based on these estimates, we developed historically based conservation targets for amount of habitat, population, and density for each species. Hindcasted reconstructions provide historical perspective for assessing contemporary trends and allow for historically based conservation targets that can inform current management.     

Declines in migrant shorebird populations from a winter‐quarter perspective

Many long‐distance migrating shorebird (i.e., sandpipers, plovers, flamingos, oystercatchers) populations are declining. Although regular shorebird monitoring programs exist worldwide, most estimates of shorebird population trends and sizes are poor or nonexistent. We built a state‐space model to estimate shorebird population trends. Compared with more commonly used methods of trend estimation, state‐space models are more mechanistic, allow for the separation of observation and state process, and can easily accommodate multivariate time series and nonlinear trends. We fitted the model to count data collected from 1990 to 2013 on 18 common shorebirds at the 2 largest coastal wetlands in southern Africa, Sandwich Harbour (a relatively pristine bay) and Walvis Bay (an international harbor), Namibia. Four of the 12 long‐distance migrant species declined since 1990: Ruddy Turnstone (Arenaria interpres), Little Stint (Calidris minuta), Common Ringed Plover (Charadrius hiaticula), and Red Knot (Calidris canutus). Populations of resident species and short‐distance migrants increased or were stable. Similar patterns at a key South African wetland suggest that shorebird populations migrating to southern Africa are declining in line with the global decline, but local conditions in southern Africa's largest wetlands are not contributing to these declines. State‐space models provide estimates of population levels and trends and could be used widely to improve the current state of water bird estimates.     

Patterns of mtDNA diversity in North Atlantic populations of the mussel Mytilus edulis

Patterns of (female) mitochondrial DNA diversity were investigated in the blue mussel Mytilus edulis. Mytilus edulis is a ubiquitous member of contemporary North Atlantic hard-substrate communities and well represented in studies of this region. Mytilus edulis was surveyed in North America and Europe, as well as mid-Atlantic sites in Greenland, Iceland, and the Faroe Islands. Mitochondrial DNA sequences revealed considerable population structure but no monophyly of haplotypes between any major regions. Coalescent analyses suggest that migration across the Atlantic Ocean has prominently been from North American source populations and that Greenland was colonized recently and exclusively from North America. In North America, there was support for two regional groups along the North American coastline. Surprisingly, we also found evidence of recombination between some Mytilus edulis and Mytilus galloprovincialis female mtDNA sequences, particularly in northern Europe. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Extinction of Mammal Populations in Western North American National Parks

Patterns of local extinction of mammal populations in western North American parks were examined in relation to current biogeographic and population lifetime models. The analysis was based on species sighting records as of 1989. While western North American parks are obviously not true isolates, patterns of mammal extinction in them are nonetheless consistent with two predictions of the land-bridge island hypothesis. First, the number of extinctions has exceeded the number of colonizations since park establishment, and, second, the rate of extinction is inversely related to park area. Factors influencing the lifetime of mammal populations were evaluated using a stepwise multivariate survival analysis procedure for censored data. Survival time for mammal populations was positively related to estimated initial population size. After accounting for population size, species within the order Lagomorpha were particularly prone to extinction. Finally, after controlling for population size and taxon variation, survival time was positively related to age of maturity, indicating that species with longer generation times—age of maturity and generation time are highly correlated in mammals—persist longer in absolute time.     

Conservation Biology of Caecilian Amphibians

Abstract:  Most of the available data on declining populations of amphibians pertain to frogs and, to a lesser extent, salamanders. In keeping with their generally less understood biology, the population trends and conservation status of caecilian amphibians (Gymnophiona) are also much less known. We reviewed reports of threats to and declines of populations of caecilians. Despite a lack of field-study details (e.g., localities, dates, and sampling methods) and quantitative data, there are several recent reports of threats to and declines and extinctions of caecilians. A range of causal explanations (habitat loss, pollution, chytridiomycosis, and scientific collecting) for these perceived declines have been proposed but little or no associated evidence has been given. Although caecilians are often considered rare and thought to require pristine habitat, published, quantitative data demonstrate that at least some species can occur in high abundance in disturbed, synanthropic environments. Few estimates of caecilian population parameters have been made and very few field methods have been tested, so the assumed rarity of any taxa remains inadequately demonstrated. Distribution and taxonomic data are also inadequate. Because they are generally poorly known and often cryptic, caecilians can be overlooked in standard faunal surveys, meaning that lack of opportunistic collection over several years might not represent evidence of decline. The conservation status of most species must be considered data deficient. More precise assessments will require a substantial increase in all areas of caecilian research, especially those involving new fieldwork. Future reports of caecilian conservation biology need to be explicit and more quantitative.     

Application of the One-Migrant-per-Generation Rule to Conservation and Management

Abstract:  Endangered species are commonly found in several (partially) isolated populations dispersed on different fragments of a habitat, natural reserve, or zoo. A certain level of connectivity among such populations is essential for maintaining genetic variation within and between populations to allow local and global adaptation and for preventing inbreeding depression. A rule of thumb widely accepted by the conservation community is that one migrant per generation (OMPG) into a population is the appropriate level of gene flow. This rule is based on Wright's study of his island model under a long list of simplifying assumptions. I examined the robustness of the OMPG rule to the violation of each of the many assumptions, quantifying the effect with population genetics theory. I showed that, when interpreted as one effective migrant per generation, OMPG is generally valid for real populations departing from the ideal model in the discrepancies of actual (  N ) and effective (  N_e  ) population sizes and actual ( m ) and effective ( m_e  ) migration rates. I also addressed the issue of converting the effective number of migrants (  M_e= N_em_e  ) into the actual number of migrants ( M = Nm  ) of a certain age and sex. In particular, N_e < N , a case common for natural populations, did not necessarily require M > M_e to maintain a certain level of differentiation among populations. Rather, translating the elusive M_e into the manageable M depends on the specific causes (e.g., biased sex ratio, reproductive skew) that lead to N_e < N .     

Are there physiological races of the sea anemone Diadumene luciae?

The chemical activators of feeding behaviour for coelenterates have been generally regarded as species specific. Recently, however, and English (North Sea) population of the sea anemone Diadumene (Haliplanella) luciae (Verrill) and an American (Californian) population of the same species were found to respond to different chemical activators. The results of further examination of this apparent anomaly show that a third population (from the English Channel, selected because of differences in colour variety and type of habitat from the North Sea population) shared the same activators as the other English population. There was no correlation between the identities of the feeding-behaviour activators and either the type of habitat or coloration of the three populations. Since the only physical factor shared by anemones with similar activators (i.e., the two English populations) was geographical, it is tentatively concluded that physiological races of D. luciae may be evolving, independently of morphological or differently coloured races, in different parts of the world.     

Avian Survival Rates and the Extinction Process on Barro Colorado Island, Panama

Abstract: Selective extinction following isolation of habitat patches may be due to biogeographical (e.g., island size or isolation) and ecological (species natural histories, interspecifc interactions) factors, or their interactions. Among the demographic and life history attributes commonly associated with high extinction probability are small populations, large size of individuals, and population variability. Long-term capture-recapture data from forest habitat in central Panama permit an examination of the association between mainland survival rates and extinction on a nearby land-bridge island Species of birds that no longer occur on Barro Colorado Island (BCI), Panama, have, on average, lower survival rates on the adjacent mainland than species that have persisted on BCI. Moreover, of the species that no longer occur on BCI, those with lower mainland survival rates generally disappeared earlier from the island. My analysis provides little evidence of a relationship between extinction and population size. Recolonization of BCI from the adjacent mainland by the forest undergrowth species studied here is unlikely. Reduced reproductive success on BCI combined with naturally low adult survival rates seems to be responsible for these BCI extinctions. High nest predation and/or altered landscape dynamics are probable agents in the low reproductive success. The methods used here could be employed in other circumstances to identify fragmentation-sensitive species.     

Effects of anthropogenic landscapes on population maintenance of waterbirds

Anthropogenic impacts have reduced natural areas but increased the area of anthropogenic landscapes. There is debate about whether anthropogenic landscapes (e.g., farmlands, orchards, and fish ponds) provide alternatives to natural habitat and under what circumstances. We considered whether anthropogenic landscapes can mitigate population declines for waterbirds. We collected data on population trends and biological traits of 1203 populations of 579 species across the planet. Using Bayesian generalized linear mixed models, we tested whether the ability of a species to use an anthropogenic landscape can predict population trends of waterbird globally and of species of conservation concern. Anthropogenic landscapes benefited population maintenance of common but not less-common species. Conversely, the use of anthropogenic landscapes was associated with population declines for threatened species. Our findings delineate some limitations to the ability of anthropogenic landscapes to mitigate population declines, suggesting that the maintenance of global waterbird populations depends on protecting remaining natural areas and improving the habitat quality in anthropogenic landscapes. Article impact statement: Protecting natural areas and improving the quality of anthropogenic landscapes as habitat are both needed to achieve effective conservation.     

Genetic Diversity and Population Size in Four Rare Southern Appalachian Plant Species

Allozyme diversity was examined in four rare, high-montane plant species from the Appalachian Mountains of southeastern North America. These species may represent relictual members or descendants of an alpine community that was more widespread during the late Pleistocene. We sampled five populations of Geum radiatum (Rosaceae), Carex misera (Cyperaceae), Trichophorum cespitosum (Cyperaceae), and the four known populations of Calamagrostis cainii (Poaceae). Genetic diversity was low for all species but was typical of that found for plant species with limited ranges. Low genetic diversity may reflect historical events associated with changes in the species' biogeography. As the Pleistocene climate warmed, suitable habitat decreased in areal extent and became fragmented, probably resulting in smaller, more-isolated populations. In recent times these species, which co-occur in fragile rock outcrop habitats, have been adversely affected by human activities. Genetic analyses revealed reduced diversity in populations of decreasing size for three species. Estimates of gene flow were low ( Nm < 1.0) in all four species. Positive associations between genetic diversity and population size, evidence of recent population declines, and the low estimates of gene flow suggest that genetic drift may play a prominent role in shaping the present-day genetic composition of these species. Furthermore, these data suggest that the genetically depauperate populations are unlikely to regain genetic variation without human intervention.     

Rapid declines of large mammal populations after the collapse of the Soviet Union

Anecdotal evidence suggests that socioeconomic shocks strongly affect wildlife populations, but quantitative evidence is sparse. The collapse of socialism in Russia in 1991 caused a major socioeconomic shock, including a sharp increase in poverty. We analyzed population trends of 8 large mammals in Russia from 1981 to 2010 (i.e., before and after the collapse). We hypothesized that the collapse would first cause population declines, primarily due to overexploitation, and then population increases due to adaptation of wildlife to new environments following the collapse. The long‐term Database of the Russian Federal Agency of Game Mammal Monitoring, consisting of up to 50,000 transects that are monitored annually, provided an exceptional data set for investigating these population trends. Three species showed strong declines in population growth rates in the decade following the collapse, while grey wolf (Canis lupus) increased by more than 150%. After 2000 some trends reversed. For example, roe deer (Capreolus spp.) abundance in 2010 was the highest of any period in our study. Likely reasons for the population declines in the 1990s include poaching and the erosion of wildlife protection enforcement. The rapid increase of the grey wolf populations is likely due to the cessation of governmental population control. In general, the widespread declines in wildlife populations after the collapse of the Soviet Union highlight the magnitude of the effects that socioeconomic shocks can have on wildlife populations and the possible need for special conservation efforts during such times. Declinación Rápida de las Poblaciones de Mamíferos Mayores después del Colapso de la Unión Soviética     

Genetic population structure in a commercial marine invertebrate with long-lived lecithotrophic larvae: <Emphasis Type="Italic">Cucumaria</Emphasis><Emphasis Type="Italic">frondosa</Emphasis> (Echinodermata: Holothuroidea)

The patterns of genetic diversity and connectivity were investigated in Cucumaria frondosa, the most abundant sea cucumber in the North Atlantic, to assist in the management and conservation of this ecologically important marine invertebrate, which is the target of an emerging fishery. Mitochondrial DNA COI sequences of 334 C. frondosa were obtained and analyzed, mainly from its western North Atlantic range, where the commercial fishery is being developed, with complementary sampling in the mid- and eastern North Atlantic. Analysis of molecular variance showed no significant (P > 0.05) differences among subpopulations in the western region suggesting that it constitutes one panmictic population. The same analysis showed low, but significant differences between eastern and western Atlantic populations. Coalescent analyses using isolation with migration models and a Bayesian skyline plot indicated historical divergence and a general increase in population size prior to the last glacial maximum and highly asymmetric gene flow (nearly 100 times lower from west to east) between sea cucumbers from North America and Norway. Results suggest that subpopulations of C. frondosa within the western North Atlantic have been highly connected. We propose that aided by the high-connectivity local subpopulations can recover rapidly from natural (i.e., ice ages) or anthropogenic (i.e., overfishing) population declines through recruitment from deep refugia.     

Population Trends in Pacific Oceanic Sharks and the Utility of Regulations on Shark Finning

Accurate assessment of shark population status is essential for conservation but is often constrained by limited and unreliable data. To provide a basis for improved management of shark resources, we analyzed a long‐term record of species‐specific catches, sizes, and sexes of sharks collected by onboard observers in the western and central Pacific Ocean from 1995 to 2010. Using generalized linear models, we estimated population‐status indicators on the basis of catch rate and biological indicators of fishing pressure on the basis of median size to identify trends for blue (Prionace glauca), mako (Isurus spp.), oceanic whitetip (Carcharhinus longimanus), and silky (Carcharhinus falciformis) sharks. Standardized catch rates of longline fleets declined significantly for blue sharks in the North Pacific (by 5% per year [CI 2% to 8%]), for mako sharks in the North Pacific (by 7% per year [CI 3% to 11%]), and for oceanic whitetip sharks in tropical waters (by 17% per year [CI 14% to 20%]). Median lengths of silky and oceanic whitetip sharks decreased significantly in their core habitat, and almost all sampled silky sharks were immature. Our results are consistent with results of analyses of similar data sets. Combined, these results and evidence of targeted fishing for sharks in some regional fisheries heighten concerns for sustainable utilization, particularly for oceanic whitetip and North Pacific blue sharks. Regional regulations that prohibit shark finning (removal of fins and discarding of the carcass) were enacted in 2007 and are in many cases the only form of control on shark catches. However, there is little evidence of a reduction of finning in longline fisheries. In addition, silky and oceanic whitetip sharks are more frequently retained than finned, which suggests that even full implementation of and adherence to a finning prohibition may not substantially reduce mortality rates for these species. We argue that finning prohibitions divert attention from assessing whether catch levels are sustainable and that the need for management of sharks should not be addressed by measures that are simple to implement but complex to enforce and evaluate. Tendencias Poblacionales de Tiburones del Océano Pacífico y la Utilidad de Regulaciones sobre Cercenamiento de Aletas     

Directional selection by fisheries and the timing of sockeye salmon (Oncorhynchus nerka) migrations.

The timing of migration from feeding to breeding areas is a critical link between the growth and survival of adult animals, their reproduction, and the fitness of their progeny. Commercial fisheries often catch a large fraction of the migrants (e.g., salmon), and exploitation rates can vary systematically over the fishing season. We examined daily records of sockeye salmon (Oncorhynchus nerka) in the Egegik and Ugashik management districts in Bristol Bay, Alaska (USA), for evidence of such temporally selective fishing. In recent years, the early migrants have experienced lower fishing rates than later migrants, especially in the Egegik district, and the median migration date of the fish escaping the fisheries has been getting progressively earlier in both districts. Moreover, the overall runs (catch and escapement) in the Egegik district and, to a lesser extent the Ugashik district, have been getting earlier, as predicted in response to the selection on timing. The trends in timing were not correlated with sea surface temperature in the region of the North Pacific Ocean where the salmon tend to concentrate, but the trends in the two districts were correlated with each other, indicating that there may be some common environmental influence in addition to the effect of selection. Despite the selection, both groups of salmon have remained productive. We hypothesize that this resilience may result from representation of all component populations among the early and late migrants, so that the fisheries have not eliminated entire populations, and from density-dependent processes that may have helped maintain the productivity of these salmon populations.     

Combining multistate capture-recapture data with tag recoveries to estimate demographic parameters

Matrix population models that allow an animal to occupy more than one state over time are important tools for population and evolutionary ecologists. Definition of state can vary, including location for metapopulation models and breeding state for life history models. For populations whose members can be marked and subsequently reencountered, multistate mark-recapture models are available to estimate the survival and transition probabilities needed to construct population models. Multistate models have proved extremely useful in this context, but they often require a substantial amount of data and restrict estimation of transition probabilities to those areas or states subjected to formal sampling effort. At the same time, for many species, there are considerable tag recovery data provided by the public that could be modeled in order to increase precision and to extend inference to a greater number of areas or states. Here we present a statistical model for combining multistate capture-recapture data (e.g., from a breeding ground study) with multistate tag recovery data (e.g., from wintering grounds). We use this method to analyze data from a study of Canada Geese (Branta canadensis) in the Atlantic Flyway of North America. Our analysis produced marginal improvement in precision, due to relatively few recoveries, but we demonstrate how precision could be further improved with increases in the probability that a retrieved tag is reported.