Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment

Summary By removing older males from their breeding territories, we tested the hypothesis that age-related dominance behavior influenced the pattern of habitat selection by breeding American redstarts Setophaga ruticilla (Aves: Parulinae). Fifteen older male redstarts removed in five experimental replicates during three breeding seasons were replaced by ten yearling and five older males; no redstart males of either age colonized the control sites during the same time periods, although two yearlings disappeared. Significantly more yearling males (67%, n=9) colonized the vacated areas than were present in the redstart population at large (26.8%, n=209). We reject the alternative hypothesis that yearling male redstarts occupy different habitats from older males because of age-related (innate) habitat preferences. Redstarts that colonized the territories made vacant by our removals (i.e., floaters) were a behaviorally heterogeneous group of animals. The presence of both yearling and older male floaters indicates that suitable habitat is limiting for this species and that intraspecific competitive interactions are important in habitat distribution, and potentially in population regulation.     

Partitioning of \alpha and \beta diversity using hierarchical Bayesian modeling of species distribution and abundance

Diversity partitioning is becoming widely used to decompose the total number of species recorded in an area or region \((\gamma )\) into the average number of species within samples \((\alpha )\) and the average difference in species composition \((\beta )\) among samples. Single-value metrics of \(\alpha \) and \(\beta \) diversity are popular because they may be applied at multiple scales and because of their ease in computation and interpretation. Studies thus far, however, have emphasized observed diversity components or comparisons to randomized, null distributions. In addition, prediction of \(\alpha \) and \(\beta \) components using environmental or spatial variables has been limited to more extensive data sets because multiple samples are required to estimate single \(\alpha \) and \(\beta \) components. Lastly, observed diversity components do not incorporate variation in detection probabilities among species or samples. In this study, we used hierarchical Bayesian models of species abundances to provide predictions of \(\alpha \) and \(\beta \) components in species richness and composition using environmental and spatial variables. We illustrate our approach using butterfly data collected from 26 grassland remnants to predict spatially nested patterns of \(\alpha \) and \(\beta \) based on the predicted counts of butterflies. Diversity partitioning using a Bayesian hierarchical model incorporated variation in detection probabilities by butterfly species and habitat patches, and provided prediction intervals for \(\alpha \) and \(\beta \) components using environmental and spatial variables.     

Simulation of herbicide impacts on a plant community: comparing model predictions of the plant community model IBC-grass to empirical data

Background

Semi-natural plant communities such as field boundaries play an important ecological role in agricultural landscapes, e.g., provision of refuge for plant and other species, food web support or habitat connectivity. To prevent undesired effects of herbicide applications on these communities and their structure, the registration and application are regulated by risk assessment schemes in many industrialized countries. Standardized individual-level greenhouse experiments are conducted on a selection of crop and wild plant species to characterize the effects of herbicide loads potentially reaching off-field areas on non-target plants. Uncertainties regarding the protectiveness of such approaches to risk assessment might be addressed by assessment factors that are often under discussion. As an alternative approach, plant community models can be used to predict potential effects on plant communities of interest based on extrapolation of the individual-level effects measured in the standardized greenhouse experiments. In this study, we analyzed the reliability and adequacy of the plant community model IBC-grass (individual-based plant community model for grasslands) by comparing model predictions with empirically measured effects at the plant community level.

Results

We showed that the effects predicted by the model IBC-grass were in accordance with the empirical data. Based on the species-specific dose responses (calculated from empirical effects in monocultures measured 4 weeks after application), the model was able to realistically predict short-term herbicide impacts on communities when compared to empirical data.

Conclusion

The results presented in this study demonstrate an approach how the current standard greenhouse experiments—measuring herbicide impacts on individual-level—can be coupled with the model IBC-grass to estimate effects on plant community level. In this way, it can be used as a tool in ecological risk assessment.

     

Reproductive,growth and life-history contrasts between two species of grapsid crabs,Sesarma cinereum and S. reticulatum

Two grapsid crabs, Sesarma cinereum (Bosc) and S. reticulatum (Say), were studied in the region of Beaufort, North Carolina, USA, during the summers of 1977, 1983, and 1984. The two grapsid species showed differences in social organization, habitat preferences, and number and size of first-stage larvae. We analyzed several other characters (growth rates, size at sexual maturity, reproductive effort per clutch) to determine if these differences could be related to a coherent pattern of life-history trends. While some relationships between characters were consistent with those found among other crab species, few seemed peculiarly matched to the ecological contrasts represented by our species. Our results suggest that other factors (phyletic predisposition; decoupling of selection between larval and adult stages; equally adaptive alternatives) may obscure the rules governing relationships between life-history characters and evolutionary trends.     

Experimental assessment of sensory modalities of coral-reef fish larvae in the recognition of their settlement habitat

One of the great mysteries of coral-reef fish ecology is how larvae locate the relatively rare patches of coral-reef habitat on which they settle. The present study aimed to estimate, by experiments in aquaria, the sensory modalities of coral-reef fish larvae for senses used in searching for their species settlement habitat. Larval recognition of settlement habitat can be based on the detection of conspecifics and/or of characteristics of coral habitat using visual, chemical and mechanical cues. For this study, larvae were captured with crest nets and were then introduced into experimental tanks that allowed testing of each type of cue separately (visual, chemical or mechanical cues). Among the 18 species studied, 13 chose their settlement habitat due to the presence of conspecifics and not based on the characteristics of coral habitat, and 5 species did not move toward their settlement habitat (e.g. Scorpaenodes parvipinnis, Apogon novemfasciatus). Among the different sensory cues tested, two species used the three types of cues (Parupeneus barberinus and Ctenochaetus striatus: visual, chemical and mechanical cues), six used two types (e.g. Myripristis pralinia: visual and chemical cues; Naso unicornis: visual and mechanical cues), and five used one type (e.g. Chrysiptera leucopoma: visual cues; Pomacentrus pavo: chemical cues). These results demonstrate that many coral-reef fish larvae could in practice use sensory cues for effective habitat selection at settlement, and have the ability to discriminate species-specific sensory cues.Communicated by J. Krause     

Statistical inference on patch-specific survival and movement rates from marked animals

Empirical estimates of patch-specific survival and movement rates are needed to parametrize spatially explicit population models, and for inference on the effects of habitat quality and fragmentation on populations. Data from radio-marked animals, in which both the fates and habitat locations of animals are known over time, can be used in conjunction with continuous-time proportional hazards models to obtain inferences on survival rates. Discrete-time conditional logistic models may provide inference on both survival and movement rates. We use Monte Carlo simulation to investigate accuracy of estimates of survival from both approaches, and movement rates from conditional logistic regression, for two habitats. Bias was low (relative bias < 0.04) and interval coverage accurate (close to the nominal 0.95) for estimates of habitat effect on survival based on proportional hazards. Bias was high ( relative bias 0.60) and interval coverage poor ( = 0.26 vs. nominal 0.95) for estimates of habitat effect based on conditional logistic regression; bias was especially influenced by heterogeneity in survival and the shape of the hazard function, whereas both bias and coverage were affected by ‘memory’ effects in movement patterns. Bias estimates of movement rate was low ( relative bias < 0.05), but interval coverage was poor ( = 0.48–0.80), possibly as a result of poor performance of a Taylor series estimate of variance. An example is provided from a radio-telemetry study of 47 wintering American woodcock (Scolopax minor), illustrating practical difficulties in field studies to parametrize these models. We also discuss extensions of continuous-time models to explicitly include a movement process, and further examine tradeoffs between continuous and discrete models.     

Seasonal changes in movements and habitat preferences of the scalloped hammerhead shark (Sphyrna lewini) while refuging near an oceanic island

Movements and habitat preferences of sharks relative to a central location are widely documented for many species; however, the reasons for such behaviors are currently unknown. Do movements vary spatially or temporally or between individuals? Do sharks have seasonal habitat and environmental preferences or simply perform movements at random at any time of the year? To help understand requirements for the designation of critical habitats for an endangered top predator and to develop zoning and management plans for key habitats, we examined vertical and horizontal movements, and determined habitat and environmental preferences of scalloped hammerhead sharks (Sphyrna lewini). We tracked seven hammerheads for 19–96 h at Wolf Island (1.38ºN, 91.82ºW) between 2007 and 2009 using ultrasonic transmitters with depth and temperature sensors, and we profiled temperature through the water column. Movements of individual hammerheads fell in two classes: constrained (remaining near the island) and dispersive (moving offshore to pelagic environments). The central activity space or kernel off the southeast side of Wolf Island was small and common to most, but the area varied among individuals (mean ± SE 0.25 ± 0.2 km²), not exceeding 0.6 km² for any of the sharks, and not changing significantly between seasons. In general, hammerheads showed preference for the up-current habitat on the eastern side of Wolf Island in both the warm and cold seasons. However, the depth of sharks varied with season, apparently in response to seasonal changes in the vertical structure of temperature. Hammerheads performed frequent vertical excursions above the thermocline during offshore movements and, in general, were observed to prefer temperatures of 23–26 °C found above the thermocline. At times, though individuals moved into the thermocline and made brief dives below it. Our results provided evidence that hammerheads (1) are highly selective of location (i.e., habitat on up-current side of island) and depth (i.e., top of the thermocline) while refuging, where they may carry out essential activities such as cleaning and thermoregulation, and (2) perform exploratory vertical movements by diving the width of the mixed layer and occasionally diving below the thermocline while moving offshore, most likely for foraging.     

Role of competition and predation in determining habitat occupancy of Cerithiidae (Gastropoda: Prosobranchia) on the rocky,intertidal, Red Sea coasts of Sinai

Six cerithiid snail species occur on rocky intertidal flats along the Sinai coasts of the Red Sea: Clypeomorus moniliferum, Cerithium caeruleum, C. scabridum, C. columna, Clypeomorus tuberculatum and Cerithium echinatum. The present study, conducted at 22 stations covering almost the whole length of Sinai, covered the 4 yr period from October 1972 to August 1976, and describes the habitat occupancy of 5 of these species (C. echinatum is excluded for lack of data). Several of these species (sometimes all) often occur together, and in such cases are distincly segregated by habitat. However all species considerably overlap in their distribution along the axes of four major interrelated abiotic gradients, thus excluding the possibility that habitat segregation is determined by larval settlement preferences. Other distributional patterns observed at some sites, such as lack of overlap or contact between belts of the various species and the relative abundance of food available to all species, make postlarval competitive interactions unlikely. The existence and the degree of proximity of a coral reef with its associated predatory fishes, influence the cerithiids' distributional patterns. Differences between the cerithiid species in their vulnerability to fish predation, associated with differences between sites in the abundance and the accessibility of predatory fishes, and in the availability of refuges for each cerithiid species, can satisfactorily explain the observed distributional patterns including co-occurrence with habitat segregation. It is proposed that habitat segregation is caused by predation on young stages by generalist fishes which may totally eliminate a certain species at a given site; the same site may provide refuges for recruits of another species, allowing these to survive to an advanced age. In general, the flat's structural complexity is associated with its diversity of refuges from predation, and hence with the number of co-existing species. This mechanism for co-existence and habitat segregation in tropical Cerithiidae may also be instrumental in maintaining the high species diversity of other tropical benthic communities.Paper No. 12 in the series Colonization of the Eastern Mediterrancan by Red Sea species immigrating through the Suez Canal     

Monitoring waterbird abundance in wetlands: The importance of controlling results for variation in water depth

Wetland use by waterbirds is highly dependent on water depth, and depth requirements generally vary among species. Furthermore, water depth within wetlands often varies greatly over time due to unpredictable hydrological events, making comparisons of waterbird abundance among wetlands difficult as effects of habitat variables and water depth are confounded. Species-specific relationships between bird abundance and water depth necessarily are non-linear; thus, we developed a methodology to correct waterbird abundance for variation in water depth, based on the non-parametric regression of these two variables. Accordingly, we used the difference between observed and predicted abundances from non-parametric regression (analogous to parametric residuals) as an estimate of bird abundance at equivalent water depths. We scaled this difference to levels of observed and predicted abundances using the formula: ((observed − predicted abundance)/(observed + predicted abundance)) × 100. This estimate also corresponds to the observed:predicted abundance ratio, which allows easy interpretation of results. We illustrated this methodology using two hypothetical species that differed in water depth and wetland preferences. Comparisons of wetlands, using both observed and relative corrected abundances, indicated that relative corrected abundance adequately separates the effect of water depth from the effect of wetlands.     

Modelling skewed data with many zeros: A simple approach combining ordinary and logistic regression

We discuss a method for analyzing data that are positively skewed and contain a substantial proportion of zeros. Such data commonly arise in ecological applications, when the focus is on the abundance of a species. The form of the distribution is then due to the patchy nature of the environment and/or the inherent heterogeneity of the species. The method can be used whenever we wish to model the data as a response variable in terms of one or more explanatory variables. The analysis consists of three stages. The first involves creating two sets of data from the original: one shows whether or not the species is present; the other indicates the logarithm of the abundance when it is present. These are referred to as the presence data and the log-abundance data, respectively. The second stage involves modelling the presence data using logistic regression, and separately modelling the log-abundance data using ordinary regression. Finally, the third stage involves combining the two models in order to estimate the expected abundance for a specific set of values of the explanatory variables. A common approach to analyzing this sort of data is to use a ln (y+c) transformation, where c is some constant (usually one). The method we use here avoids the need for an arbitrary choice of the value of c, and allows the modelling to be carried out in a natural and straightforward manner, using well-known regression techniques. The approach we put forward is not original, having been used in both conservation biology and fisheries. Our objectives in this paper are to (a) promote the application of this approach in a wide range of settings and (b) suggest that parametric bootstrapping be used to provide confidence limits for the estimate of expected abundance.     

Nonparametric estimation of Shannon’s index of diversity when there are unseen species in sample

A biological community usually has a large number of species with relatively small abundances. When a random sample of individuals is selected and each individual is classified according to species identity, some rare species may not be discovered. This paper is concerned with the estimation of Shannons index of diversity when the number of species and the species abundances are unknown. The traditional estimator that ignores the missing species underestimates when there is a non-negligible number of unseen species. We provide a different approach based on unequal probability sampling theory because species have different probabilities of being discovered in the sample. No parametric forms are assumed for the species abundances. The proposed estimation procedure combines the Horvitz–Thompson (1952) adjustment for missing species and the concept of sample coverage, which is used to properly estimate the relative abundances of species discovered in the sample. Simulation results show that the proposed estimator works well under various abundance models even when a relatively large fraction of the species is missing. Three real data sets, two from biology and the other one from numismatics, are given for illustration.     

Rarefaction and nonrandom spatial dispersion patterns

Rarefaction estimates how many species are expected in a random sample of individuals from a larger collection and allows meaningful comparisons among collections of different sizes. It assumes random spatial dispersion. However, two common dispersion patterns, within-species clumping and segregation among species, can cause rarefaction to overestimate the species richness of a smaller continuous area. We use field studies and computer simulations to determine (1) how robust rarefaction is to nonrandom spatial dispersion and (2) whether simple measures of spatial autocorrelation can predict the bias in rarefaction estimates. Rarefaction does not estimate species richness accurately for many communities, especially at small sample sizes. Measures of spatial autocorrelation of the more abundant species do not reliably predict amount of bias. Survey sites should be standardized to equal-sized areas before sampling. When sites are of equal area but differ in number of individuals sampled, rarefaction can standardize collections. When communities are sampled from different-sized areas, the mean and confidence intervals of species accumulation curves allow more meaningful comparisons among sites. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

Comparing model averaging with other model selection strategies for benchmark dose estimation

Model averaging (MA) has been proposed as a method of accommodating model uncertainty when estimating risk. Although the use of MA is inherently appealing, little is known about its performance using general modeling conditions. We investigate the use of MA for estimating excess risk using a Monte Carlo simulation. Dichotomous response data are simulated under various assumed underlying dose–response curves, and nine dose–response models (from the USEPA Benchmark dose model suite) are fit to obtain both model specific and MA risk estimates. The benchmark dose estimates (BMDs) from the MA method, as well as estimates from other commonly selected models, e.g., best fitting model or the model resulting in the smallest BMD, are compared to the true benchmark dose value to better understand both bias and coverage behavior in the estimation procedure. The MA method has a small bias when estimating the BMD that is similar to the bias of BMD estimates derived from the assumed model. Further, when a broader range of models are included in the family of models considered in the MA process, the lower bound estimate provided coverage close to the nominal level, which is superior to the other strategies considered. This approach provides an alternative method for risk managers to estimate risk while incorporating model uncertainty.

Fitness consequences of body-size-dependent host species selection in a generalist ectoparasitoid

In insect parasitoids, offspring fitness is strongly influenced by the adult females choice of host, particularly in ectoparasitoids that attack non-growing host stages. We quantified the fitness consequences of size-dependent host species selection in Dirhinus giffardii, a solitary ectoparasitoid of tephritid fruit fly pupae. We first showed a positive correlation between the size of emerged D. giffardii wasps and the size of their host fruit fly species (in order of decreasing size): Bactrocera latifrons, B. cucurbitae, B. dorsalis or Ceratitis capitata. We then manipulated individual wasps to show that the parasitoid preferred to attack the largest (B. latifrons) to the smallest (C. capitata) host species when provided with a choice, and laid a greater proportion of female eggs in B. latifrons than in C. capitata. There were no differences in developmental time or offspring survival between individuals reared from these two host species. Finally, we compared the foraging efficiency of large versus small wasps (reared from B. latifrons vs C. capitata) under two different laboratory conditions: high versus low host habitat quality, given that realized fecundity in parasitoids may be influenced by either egg-limited or time-limited factors. Under both conditions, large wasps parasitized more hosts than did small ones as a consequence of high searching efficiency in the host-poor habitat, and high capacity for adjusting egg maturation in response to host availability in the host-rich habitat. Considering the flexibility of body growth, the apparent lack of cost of achieving large body size in either development or survival, and the strong dependence of realized reproductive success on a females size, we argue that body size may be a key to understanding evolution of host species selection in ectoparasitoids. We also discuss constraints upon the evolution of size-dependent host species selection in parasitoids.Communicated by D. Gwynne     

Fish feeding on mobile benthic invertebrates: influence of spatial variability in habitat associations

Patterns of habitat association and foraging were examined for a group of tropical goatfishes (family Mullidae) that feed on mobile benthic invertebrates at Lizard Island (Great Barrier Reef). All goatfish possess barbels that disturb the substratum during feeding. Foraging methods were examined for the six most common species and used in conjunction with data on habitat associations to estimate the distribution and potential impact on the benthic invertebrate assemblage of foraging-related disturbance. Particular species exhibited broad habitat associations which differed little over two surveys (January 1989, January 1990). All species showed different preferences for the substrata they foraged. Preferences for substrata exhibited by the most common reef-associated species, Parupeneus multifasciatus, differed among locations separated by 1 km, between sites 150 m apart, and between depths (shallow and deep). Habitat preferences changed with ontogeny. Based on their habitat associations and foraging preferences, species were divided into habitat generalists and specialists. Specialists associated primarily with soft sediments. Habitat generalists, such as P. multifasciatus and P. cyclostomus, are likely to have an impact on their mobile invertebrate prey that is localised, diffuse and transitory, making any experimental analysis difficult and expensive. Habitat specialists form a guild of fishes with complementary feeding modes that efficiently exploit soft sediments and are more amenable to experimental manipulation. Experiments designed to detect the impact of foraging by these fishes must be repeated at different locations and times and must account for depth differences in foraging pressure.     

A robust weighted EM algorithm for use-availability data

Wildlife sampling for habitat selection often combines a random background sample with a random sample of used sites, because the background sample could contain too few used sites to be informative for rare species. This approach is referred to as use-availability sampling. Two variants are considered where there is: (1) a random background sample including used and unused sites augmented with a sample of used sites, and (2) a sample of used sites augmented with a contaminated background sample, i.e. use is not recorded. A weighted estimator first proposed by Manski and Lerman (Econometrica 45(8):1977?C1988, 1977) forms the basis for our suggested approach. The weighted estimator has been shown to perform better than the usual unweighted approach with uncontaminated data and mis-specified logit models (Xie and Manski in Sociol Methods Res 17(3):283?C302, 1989). A weighted EM algorithm is developed for use with contaminated background data. We show that the weighted estimator continues to perform well with contaminated data and maintains its robustness to model mis-specification. The weighted estimator has not been previously used for use-availability sampling due to reliance on the assumption that only the intercept is biased, which is valid for a correct logit model. We show that adjusting the intercept may not eliminate the bias with an incorrect logit model. In this case, the weighted estimator is a relatively simple and effective alternative.     

Are anthropogenic factors affecting nesting habitat of sea turtles? The case of Kanzul beach,Riviera Maya-Tulum (Mexico)

Marine coast modification and human pressure affects many species, including sea turtles. In order to study nine anthropogenic impacts that might affect nesting selection of females, incubation and hatching survival of loggerhead (Caretta caretta) and green turtle (Chelonia mydas), building structures were identified along a 5.2 km beach in Kanzul (Mexico). A high number of hotels and houses (88; 818 rooms), with an average density of 16.6 buildings per kilometer were found. These buildings form a barrier which prevents reaching the beach from inland, resulting in habitat fragmentation. Main pressures were detected during nesting selection (14.19% of turtle nesting attempts interrupted), and low impact were found during incubation (0.77%) and hatching (4.7%). There were three impacts defined as high: beach furniture that blocks out the movement of hatchlings or females, direct pressure by tourists, and artificial beachfront lighting that can potentially mislead hatchlings or females. High impacted areas showed lowest values in nesting selection and hatching success. Based on our results, we suggest management strategies to need to be implemented to reduce human pressure and to avoid nesting habitat loss of loggerhead and green turtle in Kanzul, Mexico.     

Spatial and temporal variability in the assemblage structure of fishes associated with mangroves (<Emphasis Type="Italic">Avicennia marina</Emphasis>) and intertidal mudflats in temperate Australian embayments

Fishes using mangrove (Avicennia marina) and mudflat habitat were sampled using three different types of gear (seine, fyke, and gill nets) at three sites within each of two large embayments through time (quarterly) between January 2002 and November 2002. At least 41 species of fish were sampled, of which 78% were marine, 17% were estuarine, and 5% were freshwater. Juveniles were sampled in 41% of the species, and 5 and 6 species occurred exclusively in mangrove and mudflat habitats, respectively. The assemblage structure of fishes varied significantly between habitats (for both fyke and seine catches) and between spring and summer (seine catches), but only in one bay. Most of the variability between habitats and times of the year could be explained by differences in abundances of atherinids, mugulids, gobiids, tetraodontids, pleuronectids, and clupeids. Fyke nets sampled mainly juvenile and smaller species of fish. Fish abundance was always greater in mangroves than mudflats (but significantly so at four of the six study sites) and varied significantly between times of the year at one site, while the number of species varied significantly between times of the year at three sites. Gill nets sampled mostly adult/subadult fishes and abundances were greater in mudflats than mangroves at two sites, and in mangroves over mudflats at one site, while species abundance varied between times of the year at two sites. The seine net sampled mainly early post-settlement and small (<20 mm) fishes, more species of which were sampled in mudflat than mangrove during winter and spring, whereas the opposite pattern occurred in summer and autumn. The number of fish sampled with the seine net only varied significantly between habitats in one embayment during summer and spring, when they were larger and smaller, respectively, in mangroves than mudflats. Mangrove habitat in temperate Australian waters supports a richer juvenile fish assemblage than adjacent mudflats, but there is little difference between habitats for the subadult/adult assemblage. Ultimately, the value of mangrove habitats to fishes depends strongly on when and where (bays and sites within bays) the study is done.Communicated by M.S. Johnson, Crawley     

Changes of microbial composition during wastewater reclamation and distribution systems revealed by high-throughput sequencing analyses

This study employed 454-pyrosequencing to investigate microbial and pathogenic communities in two wastewater reclamation and distribution systems. A total of 11972 effective 16S rRNA sequences were acquired from these two reclamation systems, and then designated to relevant taxonomic ranks by using RDP classifier. The Chao index and Shannon diversity index showed that the diversities of microbial communities decreased along wastewater reclamation processes. Proteobacteria was the most dominant phylum in reclaimed water after disinfection, which accounted for 83% and 88% in two systems, respectively. Human opportunistic pathogens, including Clostridium, Escherichia, Shigella, Pseudomonas and Mycobacterium, were selected and enriched by disinfection processes. The total chlorine and nutrients (TOC, NH₃-N and NO₃-N) significantly affected the microbial and pathogenic communities during reclaimed water storage and distribution processes. Our results indicated that the disinfectant-resistant pathogens should be controlled in reclaimed water, since the increases in relative abundances of pathogenic bacteria after disinfection implicate the potential public health associated with reclaimed water.     

Sex-specific energy requirements in nestlings of an extremely sexually size dimorphic bird, the European sparrowhawk (Accipiter nisus)

Allocation of parental investment is predicted to be equal at the population level between both sexes of offspring, and should lead to sex ratio biases in species that exhibit a sex-difference in parental care. Sex-differences in parental care are rarely quantified. We measured daily energy expenditure in free-living nestlings of the extremely sexually size dimorphic European sparrowhawk (Accipiter nisus), using the doubly labelled water method. These data were combined with measured growth characteristics to estimate daily and total metabolised energy intake of male and female young during the nestling stage. Females reached an asymptotic body mass 1.6 times higher than males. This resulted in a total metabolised energy an estimated 1.4 times higher for the nestling stage. Furthermore, we observed a decline in daily metabolised energy with an increase in brood size, which was significantly stronger for females than for males. These results are discussed in the context of Fishers equal allocation theory. Empirical evidence of a sex ratio bias at the end of parental care, with an overall excess of males, is lacking in this species. Consequently, our data do not support the idea of equal allocation between the sexes. The observed sex difference in daily metabolised energy in response to brood size may give scope for sex ratio bias at the level of the individual brood.