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Fire adaptation in Neblinaria celiae (Theaceae), a high-elevation rosette shrub endemic to a wet equatorial tepui

Neblinaria celiae (Theaceae), a rosette shrub endemic to the exceedingly rainy summit of remote Cerro de la Neblina in southern Venezuela, has a previously undescribed set of adaptations to fire. Its growth form entails sparse branching, massive terminal leaf rosettes, and thick bark. It is highly fire-tolerant, with a survival rate of 93% in a stand recently ignited by lightning, vs. 0% in seven co-occurring woody species. Survival increases sharply with rosette height, favoring a sparsely branched habit that would maximize the rate of upward growth through the sparse fuel layer supported by a sterile substrate. Thick bark and massive rosettes help protect cambial and foliar meristems from brief exposure to high temperatures. Rosettes on shorter plants are exposed to greater damage from fire near the ground and, as expected, are bigger and impound more rainwater; the greater number of leaves nearly balances the greater leaf mortality caused by fire. We relate Neblinaria's growth form to its dominance atop Neblina, to a general model for the evolution of sparse branching, and to the evolution of growth form in other tepui plants.

Oecologia

Crassulacean acid metabolism in the seasonally submerged aquatic Isoetes howellii

Evidence to date is consistent with the hypothesis that the submerged aquatic Isoetes howellii Engelmann possesses crassulacean acid metabolism. Quantitative 14 C uptake studies indicate that CO 2 assimilation in both the light and dark are functions of pH and total inorganic carbon level. In both the light and dark, maximum uptake rates in 0.6 mM NaHCO 3 were double the rates in 0.3 mM NaHCO 3 . At both carbon levels there was a large drop in carbon assimilation rate between pH 6 and 8. In nature water pH and inorganic carbon level fluctuated diurnally thus complicating the determination of the contribution of light vs dark CO 2 uptake to the total carbon gain. On a sunny day between 0600 and 1200 h water chemistry changed markedly with ∼40% reduction in total carbon, ∼2 pH unit rise resulting from ∼100% depletion of free CO 2 . Under such conditions daytime deacidification in Isoetes leaves was 88% complete by noon. In contrast, on an overcast day, reduction of carbon in the water was much slower, deacidification was only 46% complete by noon and substantial malic acid levels remained in the leaves at the end of the day. Upon emergence crassulacean acid metabolism was largely lost in Isoetes leaves. Preliminary estimates suggest that under natural submerged conditions, early morning photosynthetic rates may be substantially higher than dark CO 2 uptake rates, though uptake rates throughout much of the day could be substantially lower than nightime CO 2 assimilation.

Oecologia

Crassulacean acid metabolism in Isoetes bolanderi in high elevation oligotrophic lakes

Isoetes bolanderi dominates the littoral flora of Siesta (elevation 2,440 m) and Ellery (2,905 m) lakes in the Sierra Nevada Range of California, USA. Both lakes are sparsely vegetated and I. bolanderi maintained aboveground oven dry weight of 30–50 m −22 through most of the 1981 summer growing season. Plants at the higher elevation Ellery Lake were half as large as plants at Siesta Lake and had substantially more biomass in corms. Titratable acidity levels in Isoetes leaves showed a diurnal fluctuation <50 μeq g 1 fresh weight early in the season at the highest elevation site but this increased to ∼300 μeq g 1 FW by mid-summer; starch and chlorophyll levels likewise increased in the leaves over this time. Throughout the season the magnitude of the diurnal acid change was comparable in Isoetes from both lakes but the dynamics of daytime deacidification were not. Averaged over the season, total daytime deacidification at Ellery Lake was 65% complete by noon whereas at Siesta Lake it was only 22% complete by noon. It is suggested that this may be related to the fact that Siesta Lake was more acidic and thus more carbon was in the form of free CO 2 . In both lakes water chemistry showed no consistent diurnal fluctuation in pH or free CO 2 though total inorganic carbon levels were at the extreme low end for aquatic habitats. The studies reported here suggest that under extremely low inorganic carbon levels there may be selection for nighttime CO 2 assimilation. Consistent with this hypothesis is the observation that emergent I. bolanderi plants, resulting from fluctuating water levels, initiated leaves with stomata (unlike adjacent submerged plants) and, although these leaves had substantially higher chlorophyll levels, they showed an order of magnitude less acid fluctuation than submerged leaves.

California

Differential seed predation on two species of Arctostaphylos (Ericaceae)

The fire-prone California chaparral contains two sympatric species of shrubs: Arctostaphylos glauca and A. glandulosa . A previous study showed that in a stand where both species had similar amounts of coverage, A. glauca had fewer seeds in the soil. We attempt to answer the questions: 1) Could ground-foraging seed predators produce the lower population of A. glauca seeds in the soil? 2) Do predators select fruits randomly with respect to fruit size? 3) Do the fruits of the two species differ in the proportions of fruit components (i.e. seeds, endocarp, mesocarp, and exocarp) in ways that could be important to seed predators? Predation was measured on artificial caches of fruits, for 17 weeks. Selection by predators was examined by comparing weights of fruits recovered from soil samples with newlymatured fruits on the shrubs. Fruits components were characterized by dividing fruits into 3 fractions and weighing. More fruits of A. glauca were removed from the caches. Fruits of both species recovered from the soil were lighter than those on the shrubs. The weights of seeds, stony and fleshy fruit layers were all larger in A. glauca . Within fruits of A. glandulosa , the weights of the three components, various combinations, and ratios were all significantly correlated, while in A. glauca no other component, combination of components, or ratio examined was significantly correlated with the weights of the seeds.

California

Carbon, oxygen and hydrogen isotope abundances inStylites reflect its unique physiology

Natural abundances of oxygen-18, carbon-14, carbon-13 and deuterium were measured in Stylites andicola , a terrestrial vascular plant lacking stomates, and two terrestrial plant species having stomates that grew nearby. No substantial differences in oxygen-18 and carbon-13 abundances were observed among the three species. The deuterium concentration in Stylites was much higher than in the other species, confirming a previous report of CAM in Stylites. Stylites was depleted in carbon-14 relative to the present day atmosphere, consistent with the proposal that it fixes CO 2 derived from decomposing peat.

Oecologia

Nitrogen dynamics in an Alaskan salt marsh following spring use by geese

Lesser snow geese ( Anser caerulescens caerulescens ) and Canada geese ( Branta canadensis ) use several salt marshes in Cook Inlet, Alaska, as stopover areas for brief periods during spring migration. We investigated the effects of geese on nitrogen cycling processes in Susitna Flats, one of the marshes. We compared net nitrogen mineralization, organic nitrogen pools and production in buried bags, nitrogen fixation by cyanobacteria, and soil and litter characteristics on grazed plots versus paired plots that had been exclosed from grazing for 3 years. Grazed areas had higher rates of net nitrogen mineralization in the spring and there was no effect of grazing on organic nitrogen availability. The increased mineralization rates in grazed plots could not be accounted for by alteration of litter quality, litter quantity, microclimate, or root biomass, which were not different between grazed and exclosed plots. In addition, fecal input was very slight in the year that we studied nitrogen cycling. We propose that trampling had two effects that could account for greater nitrogen availability in grazed areas: litter incorporation into soil, resulting in increased rates of decomposition and mineralization of litter material, and greater rates of nitrogen fixation by cyanobacteria on bare, trampled soils. A path analysis indicated that litter incorporation by trampling played a primary role in the nitrogen dynamics of the system, with nitrogen fixation secondary, and that fecal input was of little importance.

Alaska

Leaf δ 13 C variability with elevation, slope aspect, and precipitation in the southwest United States

Leaves from several desert and woodland species, including gymnosperms and angiosperms with both C 3 and C 4 physiology, were analyzed to detect trends in &delta; 13 C leaf with elevation and slope aspect along two transects in southeastern Utah and south-central New Mexico, USA. The main difference between the two transects is the steeper elevational gradient for mean annual and summer precipitation in the southern transect. For any given species, we found that isotopic differences between individual plants growing at the same site commonly equal differences measured for plants along the entire altitudinal gradient. In C 3 plants, &delta; 13 C leaf values become slightly enriched at the lowest elevations, the opposite of trends identified in more humid regions. Apparently, increasing water-use efficiency with drought stress offsets the influence of other biotic and abiotic factors that operate to decrease isotopic discrimination with elevation. For some species shared by the two transects (e.g., Pinus edulis and Cercocarpus montanus ), &delta; 13 C leaf values are dramatically depleted at sites that receive more than 550 mm mean annual precipitation, roughly the boundary (pedalfer-pedocal) at which soils commonly fill to field capacity in summer and carbonates are leached. We hypothesize that, in summer-wet areas, this may represent the boundary at which drought stress overtakes other factors in determining the sign of &delta; 13 C leaf with elevation. The opposition of isotopic trends with elevation in arid versus humid regions cautions against standard correction for elevation in comparative studies of &delta; 13 C leaf .

Oecologia

Avian fruit preferences across a Puerto Rican forested landscape: Pattern consistency and implications for seed removal

Avian fruit consumption may ensure plant reproductive success when frugivores show consistent preference patterns and effectively remove and disperse seeds. In this study we examined avian fruit preferences and their seed-removal services at five study sites in north-central Puerto Rico. At each site, we documented the diet of seven common fruit-eating avian species from February to September 1998. Using foraging observations and area-based estimates of fruit abundance, we examined preference patterns of birds. We found that 7 out of 68 fleshy-fruited plant species were responsible for most of the fruit diet of birds. Seventeen plant species were preferred and four of them were repeatedly preferred across several study sites and times by at least one avian species. Preferred plant species comprised a small percentage of fleshy fruits at each site (<15% in four out of five study sites), but showed extended phenology patterns. The quantity of seeds removed by frugivore species was not strictly related to preferences. Some frugivores showing no preference could effectively remove more seeds from plants at some locations than species exhibiting constancy in their patterns of preference. Only two frugivores, Euphonia musica and Vireo altiloquous, removed most of the seeds of plants for which they exhibited repeated preference across the landscape. Preference patterns, particularly those exhibiting consistency in space and time for plant species having prolonged fruiting periods, may have important mechanistic consequences for the persistence, succession, and regeneration of tropical plant communities.

Oecologia

Nitrogen limitation of growth and nutrient dynamics in a disturbed mangrove forest, Indian River Lagoon, Florida

The objectives of this study were to determine effects of nutrient enrichment on plant growth, nutrient dynamics, and photosynthesis in a disturbed mangrove forest in an abandoned mosquito impoundment in Florida. Impounding altered the hydrology and soil chemistry of the site. In 1997, we established a factorial experiment along a tree-height gradient with three zones, i.e., fringe, transition, dwarf, and three fertilizer treatment levels, i.e., nitrogen (N), phosphorus (P), control, in Mosquito Impoundment 23 on the eastern side of Indian River. Transects traversed the forest perpendicular to the shoreline, from a Rhizophora mangle -dominated fringe through an Avicennia germinans stand of intermediate height, and into a scrub or dwarf stand of A. germinans in the hinterland. Growth rates increased significantly in response to N fertilization. Our growth data indicated that this site is N-limited along the tree-height gradient. After 2 years of N addition, dwarf trees resembled vigorously growing saplings. Addition of N also affected internal dynamics of N and P and caused increases in rates of photosynthesis. These findings contrast with results for a R. mangle -dominated forest in Belize where the fringe is N-limited, but the dwarf zone is P-limited and the transition zone is co-limited by N and P. This study demonstrated that patterns of nutrient limitation in mangrove ecosystems are complex, that not all processes respond similarly to the same nutrient, and that similar habitats are not limited by the same nutrient when different mangrove forests are compared.

Florida

Feedback dynamics of grazing lawns: Coupling vegetation change with animal growth

We studied the effects of grazing by Black Brant (Branta bernicla nigricans) geese (hereafter Brant) on plant community zonation and gosling growth between 1987 and 2000 at a nesting colony in southwestern Alaska. The preferred forage of Brant, Carex subspathacea, is only found as a grazing lawn. An alternate forage species, C. ramenskii, exists primarily as meadow but also forms grazing lawns when heavily grazed. We mowed plots of ungrazed C. ramenskii meadows to create swards that Brant could select and maintain as grazing lawns. Fecal counts were higher on mowed plots than on control plots in the year after plots were mowed. Both nutritional quality and aboveground biomass of C. ramenskii in mowed plots were similar to that of C. subspathacea grazing lawns. The areal extent of grazing lawns depends in part on the population size of Brant. High Brant populations can increase the areal extent of grazing lawns, which favors the growth of goslings. Grazing lawns increased from 3% to 8% of surface area as the areal extent of C. ramenskii meadows declined between 1991 and 1999. Gosling mass was lower early in this time period due to density dependent effects. As the goose population stabilized, and area of grazing lawns increased, gosling mass increased between 1993 and 1999. Because larger goslings have increased survival, higher probability of breeding, and higher fecundity, herbivore-mediated changes in the distribution grazing lawn extent may result in a numerical increase of the population within the next two decades.

Oecologia

Phenotypic variation and vulnerability to predation in juvenile bluegill sunfish (Lepomis macrochirus)

Bluegill sunfish (Lepomis macrochirus) are known to diversify into two forms specialized for foraging on either limnetic or littoral prey. Because juvenile bluegills seek vegetative cover in the presence of largemouth bass (Micropterus salmoides) predators, natural selection should favor the littoral body design at size ranges most vulnerable to predation. Yet within bluegill populations, both limnetic and littoral forms occur where vegetation and predators are present. While adaptive for foraging in different environments, does habitat-linked phenotypic variation also influence predator evasiveness for juvenile bluegills? We evaluate this question by quantifying susceptibility to predation for two groups of morphologically distinct bluegills; a limnetic form characteristic of bluegills inhabiting open water areas (limnetic bluegill) and a littoral form characteristic of bluegills inhabiting dense vegetation (littoral bluegill). In a series of predation trials, we found that bluegill behaviors differed in open water habitat but not in simulated vegetation. In open water habitat, limnetic bluegills formed more dense shoaling aggregations, maintained a larger distance from the predator, and required longer amounts of time to capture than littoral bluegill. When provided with simulated vegetation, largemouth bass spent longer amounts of time pursuing littoral bluegill and captured significantly fewer littoral bluegills than limnetic fish. Hence, morphological and behavioral variation in bluegills was linked to differential susceptibility to predation in open water and vegetated environments. Combined with previous studies, these findings show that morphological and behavioral adaptations enhance both foraging performance and predator evasiveness in different lake habitats.

Oecologia

Response of desert biological soil crusts to alterations in precipitation frequency

Biological soil crusts, a community of cyanobacteria, lichens, and mosses that live on the soil surface, occur in deserts throughout the world. They are a critical component of desert ecosystems, as they are important contributors to soil fertility and stability. Future climate scenarios predict alteration of the timing and amount of precipitation in desert environments. Because biological soil crust organisms are only metabolically active when wet, and as soil surfaces dry quickly in deserts during late spring, summer, and early fall, the amount and timing of precipitation is likely to have significant impacts on the physiological functioning of these communities. Using the three dominant soil crust types found in the western United States, we applied three levels of precipitation frequency (50% below-average, average, and 50% above-average) while maintaining average precipitation amount (therefore changing both timing and size of applied events). We measured the impact of these treatments on photosynthetic performance (as indicated by dark-adapted quantum yield and chlorophyll a concentrations), nitrogenase activity, and the ability of these organisms to maintain concentrations of radiation-protective pigments (scytonemin, beta-carotene, echinenone, xanthophylls, and canthaxanthin). Increased precipitation frequency produced little response after 2.5 months exposure during spring (1 April-15 June) or summer (15 June-31 August). In contrast, most of the above variables had a large, negative response after exposure to increased precipitation frequency for 6 months spring-fall (1 April-31 October) treatment. The crusts dominated by the soil lichen Collema, being dark and protruding above the surface, dried the most rapidly, followed by the dark surface cyanobacterial crusts (Nostoc-Scytonema-Microcoleus), and then by the light cyanobacterial crusts (Microcoleus). This order reflected the magnitude of the observed response: crusts dominated by the lichen Collema showed the largest decline in quantum yield, chlorophyll a, and protective pigments; crusts dominated by Nostoc-Scytonema-Microcoleus showed an intermediate decline in these variables; and the crusts dominated by Microcoleus showed the least negative response. Most previous studies of crust response to radiation stress have been short-term laboratory studies, where organisms were watered and kept under moderate temperatures. Such conditions would give crust organisms access to ample carbon to respond to imposed stresses (e.g., production of UV-protective pigments, replacement of degraded chlorophyll). In contrast, our longer-term study showed that under field conditions of high air temperatures and frequent, small precipitation events, crust organisms appear unable to produce protective pigments in response to radiation stress, as they likely dried more quickly than when they received larger, less frequent events. Reduced activity time likely resulted in less carbon available to produce or repair chlorophyll a and/or protective pigments. Our findings may partially explain the global observation that soil lichen cover and richness declines as the frequency of summer rainfall increases. ?? Springer-Verlag 2003.

Oecologia

What limits the Serengeti zebra population?

The populations of the ecologically dominant ungulates in the Serengeti ecosystem (zebra, wildebeest and buffalo) have shown markedly different trends since the 1960s: the two ruminants both irrupted after the elimination of rinderpest in 1960, while the zebras have remained stable. The ruminants are resource limited (though parts of the buffalo population have been limited by poaching since the 1980s). The zebras' resource acquisition tactics should allow them to outcompete the ruminants, but their greater spatial dispersion makes them more available to predators, and it has been suggested that this population is limited by predation. To investigate the mechanisms involved in the population dynamics of Serengeti zebra, we compared population dynamics among the three species using demographic models based on age-class-specific survival and fecundity. The only major difference between zebra and the two ruminants occurred in the first-year survival. We show that wildebeest have a higher reproductive potential than zebra (younger age at first breeding and shorter generation time). Nevertheless, these differences in reproduction cannot account for the observed differences in the population trends between the zebra and the ruminants. On the other hand, among-species differences in first-year survival are great enough to account for the constancy of zebra population size. We conclude that the very low first-year survival of zebra limits this population. We provide new data on predation in the Serengeti and show that, as in other ecosystems, predation rates on zebras are high, so predation could hold the population in a "predator pit". However, lion and hyena feed principally on adult zebras, and further work is required to discover the process involved in the high mortality of foals.

Serengeti

Geographic variation in patterns of nestedness among local stream fish assemblages in Virginia

Nestedness of faunal assemblages is a multiscale phenomenon, potentially influenced by a variety of factors. Prior small-scale studies have found freshwater fish species assemblages to be nested along stream courses as a result of either selective colonization or extinction. However, within-stream gradients in temperature and other factors are correlated with the distributions of many fish species and may also contribute to nestedness. At a regional level, strongly nested patterns would require a consistent set of structuring mechanisms across streams, and correlation among species' tolerances of the environmental factors that influence distribution. Thus, nestedness should be negatively associated with the spatial extent of the region analyzed and positively associated with elevational gradients (a correlate of temperature and other environmental factors). We examined these relationships for the freshwater fishes of Virginia. Regions were defined within a spatial hierarchy and included whole river drainages, portions of drainages within physiographic provinces, and smaller subdrainages. In most cases, nestedness was significantly stronger in regions of smaller spatial extent and in regions characterized by greater topographic relief. Analysis of hydrologic variability and patterns of faunal turnover provided no evidence that interannual colonization/extinction dynamics contributed to elevational differences in nestedness. These results suggest that, at regional scales, nestedness is influenced by interactions between biotic and abiotic factors, and that the strongest nestedness is likely to occur where a small number of organizational processes predominate, i.e., over small spatial extents and regions exhibiting strong environmental gradients. ?? Springer-Verlag 2004.

Oecologia

The relationship between species detection probability and local extinction probability

In community-level ecological studies, generally not all species present in sampled areas are detected. Many authors have proposed the use of estimation methods that allow detection probabilities that are < 1 and that are heterogeneous among species. These methods can also be used to estimate community-dynamic parameters such as species local extinction probability and turnover rates (Nichols et al. Ecol Appl 8:1213-1225; Conserv Biol 12:1390-1398). Here, we present an ad hoc approach to estimating community-level vital rates in the presence of joint heterogeneity of detection probabilities and vital rates. The method consists of partitioning the number of species into two groups using the detection frequencies and then estimating vital rates (e.g., local extinction probabilities) for each group. Estimators from each group are combined in a weighted estimator of vital rates that accounts for the effect of heterogeneity. Using data from the North American Breeding Bird Survey, we computed such estimates and tested the hypothesis that detection probabilities and local extinction probabilities were negatively related. Our analyses support the hypothesis that species detection probability covaries negatively with local probability of extinction and turnover rates. A simulation study was conducted to assess the performance of vital parameter estimators as well as other estimators relevant to questions about heterogeneity, such as coefficient of variation of detection probabilities and proportion of species in each group. Both the weighted estimator suggested in this paper and the original unweighted estimator for local extinction probability performed fairly well and provided no basis for preferring one to the other.

Oecologia

Effects of behavioral and morphological plasticity on risk of predation in a Neotropical tadpole

Predator-induced phenotypic plasticity is widespread among aquatic animals, however the relative contributions of behavioral and morphological shifts to reducing risk of predation remain uncertain. We tested the phenotypic plasticity of a Neotropical tadpole (Rana palmipes) in response to chemical cues from predatory Belostoma water bugs, and how phenotype affects risk of predation. Behavior, morphology, and pigmentation all were plastic, resulting in a predator-induced phenotype with lower activity, deeper tail fin and muscle, and darker pigmentation. Tadpoles in the predator cue treatment also grew more rapidly, possibly as a result of the nutrient subsidy from feeding the caged predator. For comparison to phenotypes induced in the experiment, we quantified the phenotype of tadpoles from a natural pool. Wildcaught tadpoles did not match either experimentally induced phenotype; their morphology was more similar to that produced in the control treatment, but their low swimming activity was similar to that induced by predator cues. Exposure of tadpoles from both experimental treatments and the natural pool to a free-ranging predator confirmed that predator-induced phenotypic plasticity reduces risk of predation. Risk of predation was comparable among wild-caught and predator-induced tadpoles, indicating that behavioral shifts can substantially alleviate risk in tadpoles that lack the typical suite of predator-induced morphological traits. The morphology observed in wild-caught tadpoles is associated with rapid growth and high competition in other tadpole species, suggesting that tadpoles may profitably combine a morphology suited to competition for food with behaviors that minimize risk of predation. ?? Springer-Verlag 2004.

Oecologia

Density-dependent habitat selection by brown-headed cowbirds (Molothrus ater) in tallgrass prairie

Local distributions of avian brood parasites among their host habitats may depend upon conspecific parasite density. We used isodar analysis to test for density-dependent habitat selection in brown-headed cowbirds (Molothrus ater) among tallgrass prairie adjacent to wooded edges, and prairie interior habitat (>100 m from wooded edges) with and without experimental perches. Eight study sites containing these three habitat treatments were established along a geographical gradient in cowbird abundance within the Flint Hills region of Eastern Kansas and Oklahoma, USA. The focal host species of our study, the dickcissel (Spiza americana), is the most abundant and preferred cowbird host in the prairie of this region. Cowbird relative abundance and cowbird:host abundance ratios were used as estimates of female cowbird density, whereas cowbird egg density was measured as parasitism frequency (percent of dickcissel nests parasitized), and parasitism intensity (number of cowbird eggs per parasitized nest). Geographical variation in cowbird abundance was independent of host abundance. Within study sites, host abundance was highest in wooded edge plots, intermediate in the experimental perch plots, and lowest in prairie interior. Cowbirds exhibited a pattern of density-dependent selection of prairie edge versus experimental perch and interior habitats. On sites where measures of cowbird density were lowest, all cowbird density estimates (female cowbirds and their eggs) were highest near (???100 m) wooded edges, where host and perch availability are highest. However, as overall cowbird density increased geographically, these density estimates increased more rapidly in experimental perch plots and prairie interiors. Variation in cowbird abundance and cowbird:host ratios suggested density-dependent cowbird selection of experimental perch over prairie interior habitat, but parasitism levels on dickcissel nests were similar among these two habitats at all levels of local cowbird parasitism. The density-dependent pattern of cowbird distribution among prairie edge and interior suggested that density effects on perceived cowbird fitness are greatest at wooded edges. A positive relationship between daily nest mortality rates of parasitized nests during the nestling period with parasitism intensity levels per nest suggested a density-dependent effect on cowbird reproductive success. However, this relationship was similar among habitats, such that all habitats should have been perceived as being equally suitable to cowbirds at all densities. Other unmeasured effects on cowbird habitat suitability (e.g., reduced cowbird success in edge-dwelling host nests, cowbird despotism at edges) might have affected cowbird habitat selection. Managers attempting to minimize cowbird parasitism on sensitive cowbird hosts should consider that hosts in otherwise less-preferred cowbird habitats (e.g., habitat interiors) are at greater risk of being parasitized where cowbirds become particularly abundant. ?? Springer-Verlag 2004.

Oecologia

Individual quality, survival variation and patterns of phenotypic selection on body condition and timing of nesting in birds

Questions about individual variation in “quality” and fitness are of great interest to evolutionary and population ecologists. Such variation can be investigated using either a random effects approach or an approach that relies on identifying observable traits that are themselves correlated with fitness components. We used the latter approach with data from 1,925 individual females of three species of ducks (tufted duck, Aythya fuligula ; common pochard, Aythya ferina ; northern shoveler, Anas clypeata ) sampled on their breeding grounds at Engure Marsh, Latvia, for over 15 years. Based on associations with reproductive output, we selected two traits, one morphological (relative body condition) and one behavioral (relative time of nesting), that can be used to characterize individual females over their lifetimes. We then asked whether these traits were related to annual survival probabilities of nesting females. We hypothesized quadratic, rather than monotonic, relationships based loosely on ideas about the likely action of stabilizing selection on these two traits. Parameters of these relationships were estimated directly using ultrastructural models embedded within capture-recapture-band-recovery models. Results provided evidence that both traits were related to survival in the hypothesized manner. For all three species, females that tended to nest earlier than the norm exhibited the highest survival rates, but very early nesters experienced reduced survival and late nesters showed even lower survival. For shovelers, females in average body condition showed the highest survival, with lower survival rates exhibited by both heavy and light birds. For common pochard and tufted duck, the highest survival rates were associated with birds of slightly above-average condition, with somewhat lower survival for very heavy birds and much lower survival for birds in relatively poor condition. Based on results from this study and previous work on reproduction, we conclude that nest initiation date and body condition covary with both reproductive and survival components of fitness. These associations lead to a positive covariance of these two fitness components within individuals and to the conclusion that these two traits are indeed correlates of individual quality.

Oecologia