Richard T. Holmes

Links between worlds: unraveling migratory connectivity

Migration is the regular seasonal movement of animals from one place to another, often from a breeding site to a nonbreeding site and back. Because the act of migration makes it difficult to follow individuals and populations year round, our understanding of the ecology and evolution of migrating organisms, particularly birds, has been severely impeded. Exciting new advances in satellite telemetry, genetic analyses and stable isotope chemistry are now making it possible to determine the population and geographical origin of individual birds. Here, we review these new approaches and consider the relevance of understanding migratory connectivity to ecological, evolutionary and conservation issues.

Foraging Behavior of Forest Birds: The Relationships Among Search Tactics, Diet, and Habitat Structure

The different searching tactics of passerine birds foraging for arthropods among the foliage of a northern hardwoods forest result in the capture of different kinds of prey. Five major searching modes are employed by the 11 foliage-foraging bird species in the Hubbard Brook Exper- imental Forest, New Hampshire. These are distinguished primarily by the rates and distances moved by the searching birds and by the types and forms of their prey-attacking maneuvers. These in turn reflect how large an area is scanned, how thoroughly it is searched, and how the bird moves from perch to perch in its search for prey. Mean searching and prey-attacking flight distances are positively correlated, indicating that birds move just far enough on average to take them into areas they have not previously searched visually. Likewise, birds that move rapidly while searching make significantly more prey attacks per unit time and hence encounter prey more often. Slow searchers scrutinize substrates more thoroughly and seem to take more cryptic and often larger prey. The results suggest that there are limitations on the ways that birds can search for and capture arthropod prey among foliage. We hypothesize that constraints imposed by the structure of the vegetation and by the types and abundances of prey determine the available foraging opportunities. Such habitat parameters may affect, in ecological or evolutionary time, the foraging traits of birds that can successfully exploit a particular habitat, and hence influence the patterns of bird habitat selection and community structure.

The use of isotope tracers for identifying populations of migratory birds

Abstract To determine whether stable isotopes can be used for identifying the geographic origins of migratory bird populations, we examined the isotopic composition of hydrogen (deuterium, δD), carbon (δ13C), and strontium (δ87Sr) in tissues of a migratory passerine, the black-throated blue warbler (Dendroica caerulescens), throughout its breeding range in eastern North America. δD and δ13C values in feathers, which are grown in the breeding area, varied systematically along a latitudinal gradient, being highest in samples from the southern end of the species’ breeding range in Georgia and lowest in southern Canada. In addition, δD decreased from east to west across the northern part of the breeding range, from New Brunswick to Michigan. δ87Sr ratios were highest in the Appalachian Mountains, and decreased towards the west. These patterns are consistent with geographical variation in the isotopic composition of the natural environment, i.e., with that of precipitation, plants, and soils for δD, δ13C, and δ87Sr, respectively. Preliminary analyses of the δD and δ13C composition of feathers collected from warblers in their Caribbean winter grounds indicate that these individuals were mostly from northern breeding populations. Furthermore, variances in isotope ratios in samples from local areas in winter tended to be larger than those in summer, suggesting that individuals from different breeding localities may mix in winter habitats. These isotope markers, therefore, have the potential for locating the breeding origins of migratory species on their winter areas, for quantifying the degree of mixing of breeding populations on migratory and wintering sites, and for documenting other aspects of the population structure migratory animals – information needed for studies of year-round ecology of these species as well as for their conservation. Combining information from several stable isotopes will help to increase the resolution for determining the geographic origins of individuals in such highly vagile populations.

SITE‐DEPENDENT REGULATION OF POPULATION SIZE:A NEW SYNTHESIS

The nature and extent of population regulation remains a principal unanswered question for many types of organisms, despite extensive research. In this paper, we provide a new synthesis of theoretical and empirical evidence that elucidates and extends a mechanism of population regulation for species whose individuals preemptively use sites that differ in suitability. The sites may be territories, refuges from predation, oviposition sites, etc. The mechanism, which we call site dependence, is not an alternative to density dependence; rather, site dependence is one of several mechanisms that potentially generate the negative feedback required for regulation. Site dependence has two major features: (1) environmentally caused heterogeneity among sites in suitability for reproduction and/or survival; and (2) preemptive site occupancy, with the tendency for individuals to move to sites of higher quality as they become available. Simulation modeling shows that these two features, acting in concert, generate nega...

Habitat-specific demography of breeding black-throated blue warblers (Dendroica caerulescens) : implications for populations dynamics

The distribution of individuals among habitats and their relative success in those habitats can have important consequences for population dynamics. To examine these processes for a long-distance migratory bird species, we studied the population structure, age-specific reproductive output, and local survival of black-throated blue warblers (Dendroica caerulescens, Gmelin) in two breeding habitats differing in shrub density within northern hardwoods forests in New Hampshire, USA. On forest plots with dense shrubs, warblers occurred at higher densities, and fledged significantly more young per capita per season than those occupying areas with lower shrub density. This differential productivity was due to higher reproductive output, mainly through double-brooding, of older (≥ 2 years of age) individuals, which were disproportionately more abundant in high shrub density sites. Clutch initiation dates, clutch sizes, and predation rates at individual nests did not differ significantly between habitats. Mean body mass of nestlings on day 6 following hatching were higher on average on plots with high shrub density, but differences were not significant. Annual return rates, as indices of local survival, did not differ between habitats for older males or for females. Yearling males, however, returned in subsequent years at a significantly lower rate to low shrub density plots, a result of either lower survival or, more likely, dispersal to more suitable habitat in their second year of breeding. Parental age and habitat suitability interact in that older individuals, through their experience and/or dominance, acquire sites of higher quality, which results in higher reproductive output and probably higher survival. These differences between habitats in density, reproductive performance and local survival are consistent with an ideal-despotic/preemptive distribution of individuals, and suggest that this population could be regulated by the availability, distribution, and extent of high and low quality breeding habitats.

Winter Habitat Quality, Population Limitation, and Conservation of Neotropical-Nearctic Migrant Birds

Recent declines in Neotropical-Nearctic migrant songbird populations are often attributed to events during the nonbreeding season, such as tropical habitat conversion and drought. Support for this hypothesis in most species, however, is largely anecdotal or conjectural. There is a dearth of demographic information about migrants on their Neotropical winter grounds. Such data are needed to identify specific ecological factors influencing survival, dispersal, and, ultimately, population abundances aggregated over multiple habitats at regional spatial scales. In this paper, we review several lines of evidence, emphasizing results of our research on paruline warblers in Jamaica, which indicate that migrant passerines often compete intraspecifically in winter for preferred quality habitats and that their populations may be limited at least in part by ecological conditions in winter. The demographic and ecological evidence supporting this hypothesis for migrant passerines includes : (1) differing densities among habitats, suggesting variation in habitat suitability ; (2) strong territoriality, site attachment, and site fidelity ; (3) experimental demonstrations of habitat saturation ; (4) nonrandom distributions of sex and age classes among habitats ; (5) overwinter decline of body mass by individuals occupying the most drought-stressed habitats ; and (6) different residence times among habitats, suggesting differences in survival or dispersal. We review ecological and behavioral explanations for these demographic patterns, and make conservation recommendations based on our understanding of how local demographic circumstances affect broader scale population processes.

Bird Predation on Forest Insects: An Exclosure Experiment

Exclusion experiments show that birds significantly reduce densities of larval Lepidoptera on forest understory vegetation. When insect densities are already low, bird predation may act both as a population regulator and as a strong agent of natural selection.

Bird Community Dynamics in a Temperate Deciduous Forest: Long‐Term Trends at Hubbard Brook

Changes in species composition and abundance of birds breeding in an unfragmented temperate deciduous forest in New Hampshire, USA, were studied intensively during 16 consecutive breeding seasons, 1969-1984. The number of species breeding in the 10-ha study area in any one year varied from 17 to 28, and averaged 24. Total numbers of individuals breeding on the 10-ha plot ranged from 214 to 89, with many species (70%) declining during the 16-yr period. Overall, there was significant positive covariation among population trends of all species, suggesting a major, perhaps single, factor affecting population levels, such as weather or food supply. No two species, however, had identical patterns of change across all 16 yr. Thus, populations in this forest fluctuated largely independently of one another, suggesting a different combination of regulatory factors for each species. The major factors pulsing or stressing bird populations in this forest included: (1) changes in food abundance due largely to irruptions of defoliating Lepidoptera (most bird species, but especially vireos and warblers), (2) harsh late spring and summer weather (Scarlet Tanager, American Redstart), (3) changes in habitat structure related to forest succession (Least Flycatcher, Philadelphia Vireo), (4) interspecific interactions, particularly interference competition (Least Flycatcher-American Redstart, Red-eyed Vireo-Philadelphia Vireo), and (5) mortality during winter for both resident and migrant populations (Hermit Thrush, Dark-eyed Junco, permanent resident species such as woodpeckers and nuthatches. Some, but not all, irruptions of defoliating Lepidoptera significantly influenced many bird popu- lations in this forest and contributed to the observed positive covariation in abundances. These irruptions produced pulses of food for breeding birds, but occurred at long and variable time intervals. Between outbreaks, food may regularly limit reproductive output of these forest birds, and low food abundance, along with the effects of predators, weather, and other mortality factors, contributes to high variability in bird reproductive success, which ultimately influences population size. Competition may also beome important during these periods of food scarcity. We therefore propose that birds in these temperate deciduous forests experience periods of prolonged food limitation, interrupted by relatively brief periods of superabundant food, a pattern contrasting with that described for other temperate systems (e.g., by Wiens 1977). From this examination of bird community dynamics on one site over many years, it is apparent that each species responds to its environment in a unique way, as determined by a variety of influences on its populations. Some of these operate on a local scale (e.g., vegetation structure, food abundance, interspecific competitors), while others function at regional (e.g., some weather effects) and global (e.g., winter events) geographic scales. These influences also differ as to their temporal persistence and predictability, and thus are difficult to detect in short-term studies. This pluralistic view of community structure for birds in temperate forests represents a compromise between nonequilibrial and com- petitionist models, and argues against the existence of a tightly organized community at any one spatial or temporal scale.

Results of Experimental and Natural Food Reductions for Breeding Black-Throated Blue Warblers

We examined effects of natural and experimentally created reductions in food abundance on the reproductive ecology of Black-throated Blue Warblers (Dendroica caerulescens). The study was carried out between 1982 and 1985 on four 30-ha plots of temperate deciduous forest within and near the Hubbard Brook Experimental Forest, New Hampshire, USA. Experimental food reduction using aerially sprayed Bacillus thuringiensis created significant differences in caterpillar biomass between one sprayed and two unsprayed plots throughout the breeding season in 1983. Where caterpillar abundance was reduced, Black-throated Blue Warblers made significantly fewer nesting attempts and diets of nest- lings included fewer caterpillars. Clutch size, hatching success, and number of young fledging per nest did not differ among the food reduction site and controls. Also, the reduced number of nesting attempts per pair on the food reduction site in 1983 did not significantly lower production of young per pair. No detectable differences in caterpillar biomass between sprayed and unsprayed sites were created in 1984 or 1985 because natural caterpillar abundances were already low. Natural declines in food abundance for Black-throated Blue Warblers occurred from 1982 through 1985, primarily because of a decrease in caterpillar abundances. Significant reductions in number of young fledging per nest, nestling growth rates and survival, and number of nests attempted per pair corresponded with the natural decline in food abun- dance. Of those measures, a reduced number of nesting attempts per pair lowered annual production the most. When effects of food limitation were calculated separately from those of nest predation, food limited annual breeding productivity to below that needed to balance annual mortality in at least one of the four years of this study. These findings indicate that neotropical migrant bird species are probably limited periodically by food when breeding in north-temperate habitats.

Tree species preferences of foraging insectivorous birds in a northern hardwoods forest

SummaryBirds searching for insects in the canopy of a northern hardwoods forest depart significantly from random in their use of tree species, even when these trees are generally similar in life form. All 10 foliage-dwelling bird species in the Hubbard Brook forest showed preferences for Yellow Birch, most had an aversion to Beech and Sugar Maple, and a few had special preferences for conifers or White Ash. Birds that glean prey from leaves had stronger tree species preferences than those that often hover for their prey, and were more influenced by tree species differences in foliage structure. The less common bird species and those for which northern hardwoods are marginal habitat had the most pronounced tree-species preferences. Food densities which are higher on Yellow Birch and specific adaptations to foraging in trees with particular foliage structures are considered major factors responsible for the observed tree species preferences. The implications of these findings for bird community structure and for forest management practices are discussed.