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Estimating population change from count data: Application to the North American Breeding Bird Survey

For birds and many other animal taxa, surveys that collect count data form a primary source of information on population change. Because counts are only indices to population size, care must be taken in using them in analyses of population change. Temporal or geographic differences in the proportion of animals counted can be misinterpreted as differences in population size. Therefore, temporally or geographically varying factors that influence the proportion of animals counted must be incorporated as covariables in the analysis of population parameters from count data. We describe the North American Breeding Bird Survey (BBS) for illustration. The BBS is a major, landscape-level survey of birds in North America; it is typical of many count surveys, in that the same sample units (survey routes) are sampled each year, and change is modeled on these routes over time. We identify covariables related to observer ability, the omission of which can bias estimation of population change from BBS data. Controlling for observer effects or other potential sources of confounding requires the specification of models relating counts to population size. We begin with a partial model specification relating expected counts to population sizes; we describe estimators currently in use in relation to this partial specification. Additional assumptions lead to a class of over-dispersed multinomial models, for which we describe estimators of population change and procedures for parsimonious model selection. We illustrate the use of over-dispersed multinomial models by an application to data for Carolina Wren (Thryothorus ludovicianus).

Ecological Applications↗

Long-term demographic consequences of habitat fragmentation to a tropical understory bird community

Tropical deforestation continues to cause population declines and local extinctions in centers of avian diversity and endemism. Although local species extinctions stem from reductions in demographic rates, little is known about how habitat fragmentation influences survival of tropical bird populations or the relative importance of survival and fecundity in ultimately shaping communities. We analyzed 22 years of mark-recapture data to assess how fragmentation influenced apparent survival, recruitment, and realized population growth rate within 22 forest understory bird species in the Usambara Mountains, Tanzania. This represents the first such effort, in either tropical or temperate systems, to characterize the effect of deforestation on avian survival across such a broad suite of species. Long-term demographic analysis of this suite of species experiencing the same fragmented environment revealed considerable variability in species' responses to fragmentation, in addition to general patterns that emerged from comparison among species. Across the understory bird community as a whole, we found significantly lower apparent survival and realized population growth rate in small fragments relative to large, demonstrating fragmentation effects to demographic rates long after habitat loss. Demographic rates were depressed across five feeding guilds, suggesting that fragmentation sensitivity was not limited to insectivores. Seniority analyses, together with a positive effect of fragmentation on recruitment, indicated that depressed apparent survival was the primary driver of population declines and observed extinctions. We also found a landscape effect, with lower vital rates in one mountain range relative to another, suggesting that fragmentation effects may add to other large-scale drivers of population decline. Overall, realized population growth rate (&lambda;) estimates were < 1 for most species, suggesting that future population persistence even within large forest fragments is uncertain in this biodiversity hotspot.

Usambara Mountains↗

Potential environmental contaminant risks to avian species at important bird areas in the northeastern United States

Environmental contaminants can have profound effects on birds, acting from the molecular through population levels of biological organization. An analysis of potential contaminant threats was undertaken at 52 Important Bird Areas (IBAs) within the northeastern Atlantic coast drainage. Using geographic information system methodology, data layers describing or integrating contamination (impaired waters, fish or wildlife consumption advisories, toxic release inventory sites, and estimates of pesticide use) were overlaid on buffered IBA boundaries, and the relative threat at each site was ranked. The most threatened sites include Jefferson National Forest (NF), Stewart B. McKinney National Wildlife Refuge (NWR), Great Dismal Swamp NWR, Blue Ridge Parkway, Shenandoah National Park (NP), Adirondack Park, Edwin B. Forsythe NWR, George Washington NF, Green Mountain NF, Long Island Piping Plover Beaches, and Merrymeeting Bay. These sites exhibited moderate to high percentages of impaired waters and had fish consumption advisories related to mercury and polychlorinated biphenyls, and were located in counties with substantial pesticide use. Endangered, threatened and Watch List bird species are present at these sites. The Contaminant Exposure and Effects--Terrestrial Vertebrates database was searched within buffered IBA boundaries, and for a moderate number of sites there was concordance between the perceived risk and contaminant exposure. Several of the IBAs with apparently substantial contaminant threats had no avian ecotoxicological data (e.g., George Washington NF, Shenandoah NP). Based upon this screening level risk assessment, contaminant biomonitoring is warranted at such sites, and data generated from these efforts should foster natural resource management activities.

Integrated Environmental Assessment and Management↗

Use of buccal swabs for sampling DNA from nestling and adult birds

We evaluated the feasibility and efficiency of using swabs to collect buccal epithelial cells from small (2‐ to 13‐ g ) birds as a source of DNA for genetic studies. We used commercially available buccal swab kits to collect samples from 42 adult and 39 nestling (4‐ to 8‐day‐old) black‐capped chickadees (Poecile atricapillus) and from 6 4‐day‐old nestling boreal chickadees (P. hudsonica). We compared DNA from buccal epithelial samples to that from blood samples from the same individuals. We extracted sufficient quantities of DNA for analysis from all buccal samples, and samples remained viable even after being stored in original plastic sampling tubes at room temperature for up to 18 months. Yields were equivalent whether extracted using the proprietary quick‐extraction solution provided with buccal swab kits or using a salt‐extraction process with inexpensive reagents. Yields of DNA from buccal samples were consistently lower than those from blood samples, but quantities were sufficient for all analyses. Assignment of sex, based on DNA extracted from paired buccal and blood samples, was identical for all 87 birds. We found no difference in the genotypes obtained from buccal and blood samples for 12 individuals tested using 5 microsatellite loci and found perfect concordance in sequencing of an 823‐base‐pair segment within the control region of mitochondrial DNA for 7 individuals tested. Use of buccal swabs is highly recommended as a rapid, noninvasive technique for sampling avian genomic DNA, especially for extremely young altricial nestlings or small‐bodied adults, or for any birds for which blood sampling may be impossible or stressful.

Wildlife Society Bulletin↗

Effects of prescribed fire on vegetation and passerine birds in northern mixed-grass prairie

Prescribed fire is used widely to manage grasslands on National Wildlife Refuges and other public lands in the northern Great Plains, but its effects on habitat use or production of wildlife in the region are poorly understood. During 19982003, we used point counts to examine effects of prescribed fire on vegetation and passerines in a mixed-grass prairie complex in north-central North Dakota, USA (n 7 units, each 4070 ha). Vegetation structure and, to a lesser extent, plant community composition varied with year of study (likely related to changes in annual precipitation) and with number of growing seasons since fire. Fire altered plant structure, especially the amount of residual vegetation, which in turn influenced bird species richness and abundance. The number of indicated pairs for sedge wren (Cistothorus platensis), clay-colored sparrow (Spizella pallida), Le Conte's sparrow (Ammodramus leconteii), Savannah sparrow (Passerculus sandwichensis), and bobolink (Dolichonyx oryzivorus) were lowest during the first postfire growing season but generally increased and stabilized within 23 postfire growing seasons. Our results support the premise that grassland passerines are well-adapted to frequent, periodic fires, generally corresponding to those occurring prior to EuroAmerican settlement of the region. Prescribed fire is important for reducing tree and shrub invasion, restoring biological integrity of plant communities, and maintaining or enhancing populations of grassland-dependent bird species. Managers in the northern mixed-grass prairie region should not be overly concerned about reductions in bird abundances that are limited mostly to the first growing season after fire. 2010 The Wildlife Society.

Journal of Wildlife Management↗

A variable circular-plot method for estimated bird numbers

A bird census method is presented that is designed for tall, structurally complex vegetation types, and rugged terrain. With this method the observer counts all birds seen or heard around a station, and estimates the horizontal distance from the station to each bird. Count periods at stations vary according to the avian community and structural complexity of the vegetation. The density of each species is determined by inspecting a histogram of the number of individuals per unit area in concentric bands of predetermined widths about the stations, choosing the band (with outside radius x) where the density begins to decline, and summing the number of individuals counted within the circle of radius x and dividing by the area (Bx2). Although all observations beyond radius x are rejected with this procedure, coefficients of maximum distance.

Condor↗

Controlling for varying effort in count surveys: An analysis of Christmas Bird Count data

The Christmas Bird Count (CBC) is a valuable source of information about midwinter populations of birds in the continental U.S. and Canada. Analysis of CBC data is complicated by substantial variation among sites and years in effort expended in counting; this feature of the CBC is common to many other wildlife surveys. Specification of a method for adjusting counts for effort is a matter of some controversy. Here, we present models for longitudinal count surveys with varying effort; these describe the effect of effort as proportional to exp(B effortp), where B and p are parameters. For any fixed p, our models are loglinear in the transformed explanatory variable (effort)p and other covariables. Hence, we fit a collection of loglinear models corresponding to a range of values of p and select the best effort adjustment from among these on the basis of fit statistics. We apply this procedure to data for six bird species in five regions, for the period 1959-1988.

Journal of Agricultural, Biological, and Environme↗

Neoplasms identified in free-flying birds

Nine neoplasms were identified in carcasses of free-flying wild birds received at the National Wildlife Health Laboratory; gross and microscopic descriptions are reported herein. The prevalence of neoplasia in captive and free-flying birds is discussed, and lesions in the present cases are compared with those previously described in mammals and birds.

Avian Diseases↗

Demography of birds in a neotropical forest: Effects of allometry, taxonomy, and ecology

Comparative demographic studies of terrestrial vertebrates have included few samples of species from tropical forests. We analyzed 9 yr of mark—recapture data and estimated demographic parameters for 25 species of birds inhabiting lowland forests in central Panama. These species were all songbirds (Order Passeriformes) ranging in mass from 7 to 57 g. Using Jolly—Seber stochastic models for open populations, we estimated annual survival rate, population size, and recruitment between sampling periods for each species. We then explored relationships between these parameters and attributes such as body size, phylogenetic affiliation, foraging guild, and social behavior. Larger birds had comparatively long life—spans and low recruitment, but body size was not associated with population size. After adjusting for effects of body size, we found no association between phylogenetic affiliation and any demographic trait. Ecological attributes, especially foraging guild, were more clearly associated with interspecific variation in all demographic traits. Ant—followers had comparatively long life—spans, but species that participate in flocks did not live longer than solitary species. The allometric associations we observed were consistent with those demonstrated in other studies of vertebrates; thus, these relationships appear to be robust. Our findings that ecological factors were more influential than phylogenetic affiliation contrasts with comparative studies of temperate—zone birds and suggests that the relative importance of environmental vs. historical factors varies geographically.

Ecology↗

Interspecific competition among Hawaiian forest birds

The object of this study was to determine whether interspecific competition modified local geographic distribution, after taking into account the effect of habitat structure. The tendencies for 14 passerine birds to have positive or negative associations were examined, using 7861 sample points in seven native forests on the islands of Hawaii, Maui, and Kauai. All birds were at least partly insectivorous and were fairly common in forested areas, although some fed chiefly on nectar or fruit. Species—pairs were classified as primary or secondary potential competitors based on general dietary similarity. To evaluate the association between species and to account for the effect of individual species habitat preferences, partial correlations were computed for each species—pair in a study area from the simple correlations between the species and 26 habitat variables plus two quadratic terms to represent nonlinearity. The partial correlations represented a short—term ("instantaneous") assessment of the strength of competitive interactions, and did not reflect the accumulation of competitive displacement through time. Of 170 partial correlations in the analysis, only 10 indicated significant negative association. The general pattern was of positive association (76 significantly positive partials), which probably resulted from flocking and from attraction of birds to areas of resource superabundance. Two species showed consistent patterns of negative partial correlations over several adjacent study areas, the Japanese White—eye/Iiwi in montane Hawaii, and the Japanese White—eye/Elepaio in windward Hawaii; both patterns could be reasonably attributed to direct competition. Species—pairs were grouped by the native or exotic status of the component species. Native/exotic pairs had a significantly greater proportion of negative partial correlations (37%) than either native/native pairs (8%) or exotic/exotic pairs (0%). This pattern was consistent across the seven study areas and appeared to reflect the occurrence of interspecific competition along a broad and diffuse ecological "front" between a co—evolved native avifauna and recently introduced exotic species. The role of competition in the pattern was corroborated by the significantly higher proportion of negative partial correlations among species—pairs of primary potential competitors than among those of secondary potential competitors. Our results suggested that °47% of the primary potential competitors among native/exotic species—pairs may experience at least small depressions in local population density due to competition. Although the negative correlations were for the most part small (average negative r = 0.06), one species could eventually replace another as spatial displacement accumulated through time. The Japanese White—eye appeared to have a principal role in native/exotic interactions, with 62% of the partial correlations between it and native primary potential competitor species being negative. Noteworthy implications were that (1) it was important to account for the habitat responses of individual species when studying the role of interspecific competition in modifying small—scale geographic distribution; (2) competition was frequently sporadic in its geographic occurrence and in the species affected, thus supporting Wiens' (1977) theory of competition; and (3) as a consequence, the role of interspecific competition in modifying distribution may be difficult to detect statistically with small data sets.

Hawaii↗

Cavity-nesting birds and the cavity-tree resource in plains cottonwood bottomlands

Densities of, and potential nesting substrates for, cavity-nesting birds were examined in a mature plains cottonwood ( Populus sargentii ) community in northeastern Colorado. Although snag (dead tree) densities were low (0.66/ha), the cavity-nesting guild included 7 species with densities ≤ 463 birds/100 ha. This finding suggests that cavity nesters are not limited by snag densities. Most (94.2%) of the nest substrate for cavity-nesting birds was provided by live trees with large, dead limbs (≥ 10 cm in diam). Both total dead limb length and the number of trees with dead limbs were highly correlated ( P < 0.001) with the number of cavities excavated. Large trees (>55 cm in diam at breast ht [dbh]) and dead limbs 15-30 cm in diameter were preferred for cavity excavation. Because snags were a minor component of potential nest substrate, snag management may not be a useful concept for cavity-nesting birds in cottonwood bottomlands, and snag retention aspects of habitat models would be relatively unimportant for this forest type. Live-tree management is recommended.

Journal of Wildlife Management↗

Scrub-successional bird community dynamics in young and mature longleaf pine-wiregrass savannahs

Public agencies are required to manage for threatened and endangered species and for biodiversity. However, at times, management for threatened and endangered species precludes consideration of other species. We investigated how managing for red-cockaded woodpeckers (Picoides borealis) and biodiversity at the Savannah River Site (SRS), South Carolina, affected communities of bird species that use early-successional scrub habitat (hereafter, scrub-successional species). Management for red-cockaded woodpeckers at the SRS involved both (1) manipulating mature longleaf pine (Pinus palustris)-wiregrass (Andropogon spp.) stands via canopy thinning, removal of midstory trees, and prescribed burning; and (2) even-aged timber harvesting. The former management practice encouraged red-cockaded woodpeckers to establish new colonies in previously unoccupied stands (hereafter, 'recruitment' stands). The latter management practice is used to remove off-site planted pines and replant with preferred longleaf pines. We conducted a constant-effort mist net study in recruitment and regenerating stands (stands clearcut and planted with longleaf pine) during the breeding seasons of 1995-96. We hypothesized that the scrub-successional bird community in recruitment stands would have greater species richness and higher survival and reproductive rates per species than in regenerating stands. However, recruitment stands always had fewer scrub-successional species (1995:36 species; 1996:31 species) than regenerating stands (1995:54 species; 1996:55 species), and all species that occurred in recruitment stands also occurred in regenerating stands. Species which commonly occurred in both recruitment and regenerating stands had similar adult:juvenile ratios (P > 0.15) and relative proportion of adults in breeding condition (P > 0.05). We detected no difference in survival rates of Bachman's sparrows (Aimophila aestivalis), indigo buntings (Passerina cyanea), and of 'combined' scrub-successional birds between stand types (P > 0.05). We found that even-aged forestry is an important management practice for maintaining and increasing avian biodiversity on public lands, as well as an acceptable management practice for red-cockaded woodpeckers.

Journal of Wildlife Management↗

Habitat relationships and nest site characteristics of cavity-nesting birds in cottonwood floodplains

We examined habitat relationships and nest site characteristics for 6 species of cavity-nesting birds--American kestrel ( Falco sparverius ), northern flicker ( Colaptes auratus ), red-headed woodpecker ( Melanerpes erythrocephalus ), black-capped chickadee ( Parus atricapillus ), house wren ( Troglodytes aedon ), and European starling ( Sturnus vulgaris )--in a mature plains cottonwood ( Populus sargentii ) bottomland along the South Platte River in northeastern Colorado in 1985 and 1986. We examined characteristics of cavities, nest trees, and the habitat surrounding nest trees. Density of large trees (>69 cm dbh), total length of dead limbs ≥10 cm diameter (TDLL), and cavity density were the most important habitat variables; dead limb length (DLL), dbh, and species were the most important tree variables; and cavity height, cavity entrance diameter, and substrate condition at the cavity (live vs. dead) were the most important cavity variables in segregating cavity nesters along habitat, tree, and cavity dimensions, respectively. Random sites differed most from cavity-nesting bird sites on the basis of dbh, DLL, limb tree density (trees with ≥1 m dead limbs ≥10 cm diameter), and cavity density. Habitats of red-headed woodpeckers and American kestrels were the most unique, differing most from random sites. Based on current trends in cottonwood demography, densities of cavity-nesting birds will probably decline gradually along the South Platte River, paralleling a decline in DLL, limb tree density, snag density, and the concurrent lack of cottonwood regeneration.

Journal of Wildlife Management↗

Mourning dove population trend estimates from Call-Count and North American Breeding Bird Surveys

The mourning dove ( Zenaida macrour a) Call-count Survey and the North American Breeding Bird Survey provide information on population trends of mourning doves throughout the continental United States. Because surveys are an integral part of the development of hunting regulations, a need exists to determine which survey provides precise information. We estimated population trends from 1966 to 1988 by state and dove management unit, and assessed the relative efficiency of each survey. Estimates of population trend differ ( P < 0.05) between surveys in 11 of 48 states; 9 of 11 states with divergent results occur in the Eastern Management Unit. Differences were probably a consequence of smaller sample sizes in the Callcount Survey. The Breeding Bird Survey generally provided trend estimates with smaller variances than did the Callcount Survey. Although the Callcount Survey probably provides more withinroute accuracy because of survey methods and timing, the Breeding Bird Survey has a larger sample size of survey routes and greater consistency of coverage in the Eastern Unit.

continental United States↗

Breeding bird response to juniper woodland expansion

In recent times, pinyon ( Pinus spp. )-juniper ( Juniperus spp. ) woodlands have expanded into large portions of the Southwest historically occupied by grassland vegetation. From 1997-1998, we studied responses of breeding birds to one-seed juniper ( J. monosperma ) woodland expansion at 2 grassland study areas in northern Arizona. We sampled breeding birds in 3 successional stages along a grassland-woodland gradient: un-invaded grassland, grassland undergoing early stages of juniper establishment, and developing woodland. Species composition varied greatly among successional stages and was most different between endpoints of the gradient. Ground-nesting grassland species predominated in uninvaded grassland but declined dramatically as tree density increased. Tree- and cavity-nesting species increased with tree density and were most abundant in developing woodland. Restoration of juniper-invaded grasslands will benefit grassland-obligate birds and other wildlife.

Journal of Range Management↗

Removal and repopulation of breeding birds in a spruce-fir forest community

IN 1949, while engaged in population studies of birds in northern Maine, the authors accumulated considerable information concerning population dynamics of birds inhabiting the Spruce-Fir forest community. This information was obtained in connection with investigations of the effective control by breeding birds of an infestation of the spruce budworm, Choristoneura fumiferana Clem.

The Auk↗

Avian influenza virus and free-ranging wild birds

Recent media and news reports and other information implicate wild birds in the spread of highly pathogenic avian influenza in Asia and Eastern Europe. Although there is little information concerning highly pathogenic avian influenza viruses in wild birds, scientists have amassed a large amount of data on low-pathogenicity avian influenza viruses during decades of research with wild birds. This knowledge can provide sound guidance to veterinarians, public health professionals, the general public, government agencies, and other entities with concerns about avian influenza.

Asia, Eastern Europe↗

Effects of management practices on grassland birds: Greater Prairie-Chicken

Information on the habitat requirements and effects of habitat management on grassland birds were summarized from information in more than 5,500 published and unpublished papers. A range map is provided to indicate the breeding distribution of Greater Prairie-Chicken in the United States and southern Canada. Although birds frequently are observed outside the breeding range indicated, the maps are intended to show areas where managers might concentrate their attention. It may be ineffectual to manage habitat at a site for a species that rarely occurs in an area. The species account begins with a brief capsule statement, which provides the fundamental components or keys to management for the species. A section on breeding range outlines the current breeding distribution of the species in North America. The suitable habitat section describes the breeding habitat and occasionally microhabitat characteristics of the species, especially those habitats that occur in the Great Plains. Details on habitat and microhabitat requirements often provide clues to how a species will respond to a particular management practice. A table near the end of the account complements the section on suitable habitat, and lists the specific habitat characteristics for the species by individual studies. A special section on prey habitat is included for those predatory species that have more specific prey requirements. The area requirements section provides details on territory and home range sizes, minimum area requirements, and the effects of patch size, edges, and other landscape and habitat features on abundance and productivity. It may be futile to manage a small block of suitable habitat for a species that has minimum area requirements that are larger than the area being managed. The section on breeding-season phenology and site fidelity includes details on spring arrival and fall departure for migratory populations in the Great Plains, peak breeding periods, the tendency to renest after nest failure or success, and the propensity to return to a previous breeding site. The duration and timing of breeding varies among regions and years. Species&rsquo; response to management summarizes the current knowledge and major findings in the literature on the effects of different management practices on the species. The section on management recommendations complements the previous section and summarizes specific recommendations for habitat management provided in the literature. If management recommendations differ in different portions of the species&rsquo; breeding range, recommendations are given separately by region. The literature cited contains references to published and unpublished literature on the management effects and habitat requirements of the species. This section is not meant to be a complete bibliography; a searchable, annotated bibliography of published and unpublished papers dealing with habitat needs of grassland birds and their responses to habitat management is posted at the Web site mentioned below.

Kansas, Oklahoma, Colorado, Missouri, Illinois, Wi↗