Habitat selection and reproductive success of Lewis's Woodpecker (Melanerpes lewis) at its northern limit.

Lewis's Woodpecker (Melanerpes lewis) has experienced population declines in both Canada and the United States and in 2010 was assigned a national listing of threatened in Canada. We conducted a two-year study (2004-2005) of this species at its northern range limit, the South Okanagan Valley in British Columbia, Canada. Our main objective was to determine whether the habitat features that influenced nest-site selection also predicted nest success, or whether other factors (e.g. cavity dimensions, clutch initiation date or time of season) were more important. Nest tree decay class, density of suitable cavities and total basal area of large trees were the best predictors of nest-site selection, but these factors were unrelated to nesting success. Estimates of demographic parameters (mean ± SE) included daily nest survival rate (0.988±0.003, years combined), nest success (0.52±0.08), clutch size (5.00±0.14 eggs), female fledglings per successful nest (1.31±0.11), and annual productivity (0.68±0.12 female fledglings per nest per year). Although higher nest survival was associated with both early and late initiated clutches, early-initiated clutches allowed birds to gain the highest annual productivity as early clutches were larger. Nests in deep cavities with small entrances experienced lower predation risk especially during the peak period of nest predation. We concluded that nest-site selection can be predicted by a number of easily measured habitat variables, whereas nest success depended on complicated ecological interactions among nest predators, breeding behaviors, and cavity features. Thus, habitat-based conservation strategies should also consider ecological factors that may not be well predicted by habitat.

Species loss, delayed numerical responses, and functional compensation in an antbird guild.

When a community loses species through fragmentation, its total food consumption may drop. Compensatory responses of remaining species, whereby survivors assume roles of extinct competitors, may reduce the impact of species loss through numerical or functional responses. We measured compensatory responses in two remaining antbird species on Barro Colorado Island, Panama, four decades after the loss of their dominant competitor, the Ocellated Antbird, Phaenostictus mcleannani. We compared current abundances and behavior of these two species on Barro Colorado to those reported before the island lost Ocellated Antbirds, and to those in a nearby mainland population where all three species still exist as a space-for-time substitution. The smaller, more subordinate Spotted Antbird, Hylophylax naevioides, responded far more strongly than the larger Bicolored Antbird, Gymnopithys leucaspis, which is functionally more like the Ocellated Antbird. Islandwide density of Spotted Antbirds has more than doubled since the loss of Ocellated Antbirds. Moreover, Spotted Antbirds now spend so much more of their time following ant swarms that their metabolic biomass at these swarms has more than tripled since Ocellated Antbirds disappeared. These responses in Spotted Antbirds were apparently delayed by >20 years. Bicolored Antbirds have not increased substantially in islandwide density or metabolic biomass at ant swarms. We hypothesize that behavioral flexibility, as shown by Spotted Antbirds on Barro Colorado Island, is a major factor governing the extent to which fragmented ecosystems can buffer the impacts of species loss.

Food and predators affect egg production in song sparrows.

Although the possibility that food and predators may interact in limiting avian populations has long been recognized, there have been few attempts to test this experimentally in the field. We conducted a manipulative food addition experiment on the demography of Song Sparrows (Melospiza melodia) across sites that varied in predator abundance, near Victoria, British Columbia, Canada, over three consecutive breeding seasons. We previously showed that food and predators had interactive effects on annual reproductive success (young fledged per female). Here, we report the effects on egg production. Our results show that food limits the total number of eggs laid over the breeding season ("total egg production") and that interactive food and predator effects, including food effects on nest predation, determine how those eggs are "parceled out" into different nests. Food addition alone significantly affected total egg production, and there was no significant interannual variability in this result. At the same time, both food and predators affected the two determinants of total egg production: "clutch number" (total number of clutches laid) and average clutch size. Both clutch number and size were affected by a food x predator x year interaction. Clutch number was lower at low-predator locations because there was less nest predation and thus less renesting. Food addition also significantly reduced nest predation, but there was significant interannual variation in this effect. This interannual variation was responsible for the food x predator x year interactions because the larger the effect of food on nest predation in a given year, the smaller was the effect of food on clutch number; and the smaller the effect of food on clutch number, the larger was the effect of food on clutch size. Potential predator and year effects on total egg production were thus cancelled out by an inverse relationship between clutch number and clutch size. We suggest that combined food and predator effects on demography could be the norm in both birds and mammals.

Combined food and predator effects on songbird nest survival and annual reproductive success: results from a bi-factorial experiment.

Food and predators have traditionally been viewed as mutually exclusive alternatives when considering factors affecting animal populations. This has led to long controversies such as whether annual reproductive success in songbirds is primarily a function of food-restricted production or predator-induced loss. Recent studies on both birds and mammals suggest many of these controversies may be resolved by considering the combined effects of food and predators. We conducted a 2x2 manipulative food addition plus natural predator reduction experiment on song sparrows (Melospiza melodia) over three consecutive breeding seasons. Food and predators together affected partial clutch or brood loss, nest survival (total clutch or brood loss) and annual reproductive success. When combined, our two treatments reduced partial losses by more than expected if the effects of food and predators were independent and additive. Food and predators also interacted in their effects on nest survival since food addition significantly reduced the rate of nest predation. While annual reproductive success was highly correlated with nest predation (r2=0.71) the strength of this relationship was reinforced by the indirect effects of food addition on nest predation. A stepwise multiple regression showed that the residual variation in annual reproductive success was explained by food effects on the total number of eggs laid over the season and the combined effects of food and predators on partial losses noted above. We conclude that annual reproductive success in song sparrows is a function of both food-restricted production and predator-induced loss and indirect food and predator effects on both clutch and brood loss. We highlight the parallels between our results and those from a comparable bi-factorial experiment on mammals because we suspect combined food and predator effects are likely the norm in both birds and mammals.

Fitness correlates of song repertoire size in free-living song sparrows (Melospiza melodia).

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male's own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free-living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male's song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte's population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.

Balancing food and predator pressure induces chronic stress in songbirds.

The never-ending tension between finding food and avoiding predators may be the most universal natural stressor wild animals experience. The 'chronic stress' hypothesis predicts: (i) an animal's stress profile will be a simultaneous function of food and predator pressures given the aforesaid tension; and (ii) these inseparable effects on physiology will produce inseparable effects on demography because of the resulting adverse health effects. This hypothesis was originally proposed to explain synergistic (inseparable) food and predator effects on demography in snowshoe hares (Lepus americanus). We conducted a 2 x 2, manipulative food addition plus natural predator reduction experiment on song sparrows (Melospiza melodia) that was, to our knowledge, the first to demonstrate comparable synergistic effects in a bird: added food and lower predator pressure in combination produced an increase in annual reproductive success almost double that expected from an additive model. Here we report the predicted simultaneous food and predator effects on measures of chronic stress in the context of the same experiment: birds at unfed, high predator pressure (HPP) sites had the highest stress levels; those at either unfed or HPP sites showed intermediate levels; and fed birds at low predator pressure sites had the lowest stress levels.

Synergistic effects of food and predators on annual reproductive success in song sparrows.

The behaviour literature is full of studies showing that animals in every taxon balance the probability of acquiring food with the risk of being preyed upon. While interactions between food and predators clearly operate at an individual scale, population-scale studies have tended to focus on only one factor at a time. Consequently, interactive (or 'synergistic') effects of food and predators on whole populations have only twice before been experimentally demonstrated in mammals. We conducted a 2 x 2 experiment to examine the joint effects of food supply and predator pressure on the annual reproductive success of song sparrows (Melospiza melodia). Our results show that these two factors do not operate in an additive way, but instead have a synergistic effect on reproduction. Relative to controls, sparrows reared 1.1 more young when food was added and 1.3 more when predator pressure was low. When these treatments were combined 4.0 extra young were produced, almost twice as many as expected from an additive model. These results are a cause for optimism for avian conservation because they demonstrate that remedial actions, aimed at simultaneously augmenting food and reducing predators, can produce dramatic increases in reproductive success.

NATURAL SELECTION ON BEAK AND BODY SIZE IN THE SONG SPARROW.

We documented temporal patterns of natural selection on beak and body traits in a song sparrow population. We looked for evidence of selection in association with reproduction and overwinter survival in order to identify the conditions under which size in beak and body traits is adaptive. We also attempted to identify the specific traits most closely associated with fitness under these conditions. Selection was observed in association with both survival and reproduction. Patterns of selection differed between the sexes. Selection on males was weak and stabilizing in association with overwinter survival. Selection on females was strong, was both stabilizing and directional, and was associated with both survival and reproduction. In females, traits that enhanced juvenile survival also reduced reproductive success; i.e., there was a trade-off between survival and reproduction. Patterns of selection in the song sparrow parallel those reported for the Galápagos finch, Geospiza fortis. However, in song sparrows, selection occurred mainly on tarsus length and beak length, and not on beak depth or width as in G. fortis. This difference may occur because most North American sparrows partition food resources by habitat, while most Galápagos ground finches partition food by seed size.

A winter feeding experiment on an island Song Sparrow population.

Supplemental food, in the form of millet seed, was provided to half of an island Song Sparrow population during the 1978-1979 winter to test if winter food influenced: (1) overwinter survival; (2) winter wights; (3) breeding density in 1979 and (4) 1979 breeding performance.Territorial males were most dominant at feeders and may have restricted access of young to feeders. Young females were most subordinate at feeders. Adult survival was not affected by supplementary food, but young survival was higher than in 6 previous years and young seen to visit feeders may have survived better than young not seen at feeders. Young females were more variable in weight on the unfed half of the island than on the fed end. The breeding population increased by 38% from 1978 to 1979, but it is not known how much of this increase resulted from food addition. Pairs of birds with feeders on their territories began to lay 25 days earlier in 1979 than control pairs, but delayed longer than controls before a second breeding attempt. One-year old females began to lay significantly later than adults on the control area, but not on the fed area. Other measures of breeding performance were not affected by supplemental food. Winter food may be more important and male territorial behaviour less important than previously supposed in limiting numbers in the Mandarte Island Song Sparrow population.