Building Hierarchical Models of Avian Distributions for the State of Georgia

Abstract To predict the distributions of breeding birds in the state of Georgia, USA, we built hierarchical models consisting of 4 levels of nested mapping units of decreasing area: 90,000 ha, 3,600 ha, 144 ha, and 5.76 ha. We used the Partners in Flight database of point counts to generate presence and absence data at locations across the state of Georgia for 9 avian species: Acadian flycatcher (Empidonax virescens), brown-headed nuthatch (Sitta pusilla), Carolina wren (Thryothorus ludovicianus), indigo bunting (Passerina cyanea), northern cardinal (Cardinalis cardinalis), prairie warbler (Dendroica discolor), yellow-billed cuckoo (Coccyzus americanus), white-eyed vireo (Vireo griseus), and wood thrush (Hylocichla mustelina). At each location, we estimated hierarchical-level-specific habitat measurements using the Georgia GAP Analysis18 class land cover and other Geographic Information System sources. We created candidate, species-specific occupancy models based on previously reported relationships, and fit these using Markov chain Monte Carlo procedures implemented in OpenBugs. We then created a confidence model set for each species based on Akaike's Information Criterion. We found hierarchical habitat relationships for all species. Three-fold cross-validation estimates of model accuracy indicated an average overall correct classification rate of 60.5%. Comparisons with existing Georgia GAP Analysis models indicated that our models were more accurate overall. Our results provide guidance to wildlife scientists and managers seeking predict avian occurrence as a function of local and landscape-level habitat attributes.     

Width of riparian buffer and structure of adjacent plantations influence occupancy of conservation priority birds

Conservation of biodiversity on forest landscapes dominated by plantations has become an increasingly important topic, and opportunities to maintain or enhance biodiversity within these forests need to be recognized and applied. Riparian buffers of mature forest retained along streams in managed forest landscapes offer an opportunity to enhance biodiversity across these landscapes. However, influence of buffer width and structure of adjacent plantations on habitat use by birds is not well understood. We modeled probability of occupancy, while accounting for variable detection probabilities, for 16 bird species of regional conservation importance in the Ouachita Mountains of Arkansas, USA. We examined occurrence of breeding birds in streamside management zone (SMZ) buffers embedded in three structural classes of pine plantation: young open-canopy, closed-canopy, and older thinned plantations. Our occupancy models included a positive association with SMZ width for nine bird species associated with mature forests. Models for three early successional species (prairie warbler [Dendroica discolor], white-eyed vireo [Vireo griseus], and northern bobwhite [Colinus virginianus]) included a negative association with SMZ width. Occupancy models for Acadian flycatcher (Empidonax virescens), summer tanager (Piranga rubra), pine warbler (Dendroica pinus), prairie warbler, and northern bobwhite also included structural condition of adjacent plantations, but most species did not appear affected by condition of surrounding plantations. We found diverse responses among species to width of retained SMZs and structure of adjacent plantations; some species apparently benefitted from SMZs >100 m wide, while others benefitted from narrow buffers. Furthermore, most species traditionally associated with mature forests were common in narrow SMZs, regardless of width. Thus, optimal width of SMZs relative to avian conservation depends on the species of greatest conservation interest.     

Bird Response to Clear Cutting in Missouri Ozark Forests

Abstract: We evaluated changes in breeding bird density and shifts in territory distribution with respect to clear cutting and timber stand improvement (TSI) of even-aged stands on >300 ha experimental management units as part of the Missouri Ozark Forest Ecosystem Project. After one harvest entry, clear cutting had positive effects on density of indigo bunting (Passerina cyanea), prairie warbler (Dendroica discolor), and yellow-breasted chat (Icteria virens) and negative effects on density of Acadian flycatcher (Empidonax virescens), ovenbird (Seiurus aurocapilla), and worm-eating warbler (Helmitheros vermivorus). In buffer regions within 100 m of clearcuts, indigo bunting, hooded warbler (Wilsonia citrina), wood thrush (Hylocichila mustelina), and Kentucky warbler (Oporornis formosus) densities increased and ovenbird density decreased. Breeding bird densities did not change in interior regions > 100 m from clearcuts except for a small increase for wood thrush. Breeding Acadian flycatcher and ovenbird showed greater use of stands not treated with TSI. We recommend combining adjoining stands to keep clearcut sizes between 8 ha and 13 ha to reduce negative effects on ovenbirds by cutting. We suggest a 7-year offset between the timing of clear cutting and TSI to reduce their combined effects on ovenbird.     

Polymorphic microsatellite loci from the American goldfinch (Carduelis tristis) and their cross‐amplification in a variety of passerine species

Cardueline finches have become important models in studies of sexual selection. Here I describe eight polymorphic microsatellite loci isolated from the American goldfinch (Carduelis tristis) that are useful in parentage and heterozygosity studies. The number of alleles per locus ranged from eight to 38 based on 252 goldfinches. The markers are polymorphic in at least four other Cardueline finch species, and several amplify in nonfinch passerines including red‐winged blackbird (Agelaius phoeniceus), black‐throated blue warbler (Dendroica caerulescens) and tufted titmouse (Baeolophus bicolor), indicating they should be useful in paternity and population studies of a variety of species.     

Short-term responses of birds to prescribed fire in fire-suppressed forests of California

Prescribed fire is one tool for restoring fire-suppressed forests, but application of fire during spring coincides with breeding and arrival of migrant birds. We examined effects of low-severity prescribed fires on counts of birds in a managed forest in the Sierra Nevada of California immediately, 1 year, and 3–6 years after fire was applied in spring. Of 26 species analyzed, counts of 3 species increased after fire (Pacific-slope flycatcher [Empidonax difficilis], brown creeper [Certhia americana], and American robin [Turdus migratorius]), and 6 species decreased after fire (Anna's hummingbird [Calypte anna], Hutton's vireo [Vireo huttoni], warbling vireo [Vireo gilvus], golden-crowned kinglet [Regulus satrapa], Nashville warbler [Vermivora ruficapilla], hermit warbler [Dendroica occidentalis]). Black-throated gray warbler (Dendroica nigrescens) increased in the first year following fire but decreased 3–6 years after fire. When grouped into guilds, habitat association and foraging guild best explained responses to fire, with the greatest changes occurring for oak-associated species (negative), riparian-associated species (positive), aerial foragers (positive), and bark foragers (positive). Lastly, when we compared our counts to those collected during the 1910s, changes were consistent with those predicted from fire suppression and species' affinity for burned forests, suggesting that results from contemporary fire studies should be interpreted within an ecological context that includes effects of fire suppression. We found that low-severity prescribed fires applied in spring served to drive the bird community towards pre-suppression conditions. © 2011 The Wildlife Society.     

Implications of Climate Change for Bird Conservation in the Southwestern U.S. under Three Alternative Futures

Future expected changes in climate and human activity threaten many riparian habitats, particularly in the southwestern U.S. Using Maximum Entropy (MaxEnt3.3.3) modeling, we characterized habitat relationships and generated spatial predictions of habitat suitability for the Lucy’s warbler (Oreothlypis luciae), the Southwestern willow flycatcher (Empidonax traillii extimus) and the Western yellow-billed cuckoo (Coccyzus americanus). Our goal was to provide site- and species-specific information that can be used by managers to identify areas for habitat conservation and/or restoration along the Rio Grande in New Mexico. We created models of suitable habitat for each species based on collection and survey samples and climate, biophysical, and vegetation data. We projected habitat suitability under future climates by applying these models to conditions generated from three climate models for 2030, 2060 and 2090. By comparing current and future distributions, we identified how habitats are likely to change as a result of changing climate and the consequences of those changes for these bird species. We also examined whether land ownership of high value sites shifts under changing climate conditions. Habitat suitability models performed well. Biophysical characteristics were more important that climate conditions for predicting habitat suitability with distance to water being the single most important predictor. Climate, though less important, was still influential and led to declines of suitable habitat of more than 60% by 2090. For all species, suitable habitat tended to shrink over time within the study area leaving a few core areas of high importance. Overall, climate changes will increase habitat fragmentation and reduce breeding habitat patch size. The best strategy for conserving bird species within the Rio Grande will include measures to maintain and restore critical habitat refugia. This study provides an example of a presence-only habitat model that can be used to inform the management of species at intermediate scales.     

Interspecific variability of prevalence in blood parasites of adult passerine birds during the breeding season in Alaska

Blood parasite prevalence based on microscopic examination of stained blood smears was determined in adults of 11 passerine bird species sampled during their breeding season (May and June 1997-98) in interior Alaska (USA). These species included primarily Nearctic migratory species such as the dark-eyed junco (Junco hyemalis) and neotropical migratory species such as the blackpoll warbler (Dendroica striata), alder flycatcher (Empidonax alnorum), Swainson's thrush (Catharus ustulatus), northern waterthrush (Seiurus noveboracensis), and bank swallow (Riparia riparia) as well as one long-distance palearctic migrant, the arctic warbler (Phylloscopus borealis). The more prevalent parasites were Leucocytozoon dubreuili (73% of the sampled turdinids), L. fringillinarum (42% of the sampled fringillids and parulids), and Trypanosoma avium (39% of the sampled hosts). Other parasites (H. fallisi: 18% of the sampled turdinids; Haemoproteus paruli: 14% of the sampled parulids; H. fringillae: 5% of the sampled fringillids; microfilariae: 4% of the sampled hosts) were observed less frequently. Plasmodium vaughani was found only in two yellow warblers (Dendroica petechia). Overall parasite prevalence varied between 0% in the alder flycatcher to >80% in Swainson's thrush, arctic warbler, and Townsend's warbler (Dendroica townsendi). Prevalence of various hematozoa also was bird species-dependent. No relationship was observed between prevalence and either foraging (aerial versus trees/shrubs) or nesting habits (ground versus arboreal) or general location of the wintering area of the different species examined. Prevalence also was unrelated to average dates of arrival on breeding grounds and, therefore, to potential duration of exposure to local insect vectors before capture. Differences in blood parasite prevalence among species breeding in a same region and in the same type of habitat may result from differences in host specificity such as immunological resistance to infection or blood meal preference by potential vectors and/or in behavioral adjustments/physiological traits that alter exposure to vectors.     

An indicator system to assess ecological integrity of managed forests

Ecological integrity of managed forests includes the ability of an ecosystem to support a community of organisms with a similar species composition and functional organization as found in nearby natural systems. We developed an indicator system for ecological integrity based on simulated natural disturbance and indicator species to test if forest condition and habitat in managed forests are similar to that found or expected in natural systems. We then applied the method in an area of the boreal forest (Ontario, Canada) where the objective of Ontario's strategic forest management planning approach is, in part, to conserve ecological integrity through the emulation of the natural disturbance process. Forest condition controls the supply of habitat to support the diversity of native organisms, and historically in boreal forests the natural disturbance process drove forest condition. We selected indicators of forest condition (landscape pattern and compositional mosaic) and habitat function (occupancy rates for a broad range of forest birds), and applied our assessment system to test whether indicators of forest condition and habitat function reflect outcomes expected if natural disturbance processes were successfully emulated. We collected occupancy data in natural and managed forest disturbance types using autonomous acoustic recorders, applied occupancy/detection modeling to estimate corrected occupancy rates (ψ), and then tested for differences in ψ between disturbance types. Some indicators of forest condition were within the range expected under natural disturbance, but we found relatively less old conifer, more young deciduous and greater edge density in managed forests relative to forests of natural disturbance origin. Most species (11 of 14) occurred with equal ψ in habitat originating from the two disturbance types. Brown creeper (Certhia americana), bay-breasted warbler (Mniotilta varia) and red-eyed vireo (Vireo olivaceus) differed between disturbance types. Brown creeper uses older conifer and occurred at lower rates in managed forest, while red-eyed vireo uses a range of deciduous forest ages, and occurred at higher rates in managed forest. Differences in quantity and/or quality of specific habitat types likely explain the responses. The results suggest what directional changes in the forest pattern and compositional mosaic would improve ecological similarity with natural systems, but also indicate what further research is required. We believe this approach to assessing ecological integrity can be adapted to study the effectiveness of conservation management strategies in other systems, and will contribute to adaptive management approaches and evidence-based policy development.     

How important are climate‐induced changes in host availability for population processes in an obligate brood parasite,the European cuckoo?

In recent years, populations of long‐distance migrant birds have declined markedly. Resource availability, both on breeding and wintering grounds, is likely to be important particularly since changing climates are affecting the timing and synchrony of such resources. We use novel analytical methods to examine whether large‐scale population declines in the brood‐parasite common cuckoo Cuculus canorus are the result of changes in the abundance or timing of breeding of its host species. We find that, due to climate‐induced changes in the timing of breeding, availability of dunnock Prunella modularis nests has decreased, but that availability of reed warbler Acrocephalus scirpaceus has increased. Although there is no evidence that the timing of breeding of cuckoo has changed, these changes are likely to have had only a minimal impact on its population trend, but may explain an increase in the rate of parasitism of reed warbler nests in recent decades.     

Reservoir water levels do not influence daily mass gain of warblers at a riparian stopover site

ABSTRACT Hydroelectric dam operations that lead to fluctuations in the water levels of reservoirs can influence the amount of riparian habitat available for migrating songbirds and may impact the use and quality of remaining habitat. Our objective was to determine if use of riparian habitats and mass gain by five warbler species at the Columbia River‐Revelstoke Migration Monitoring Station in British Columbia, Canada, were influenced by water levels in the surrounding Arrow Lakes Reservoir. We analyzed fall migration data collected from 1998 to 2006. Capture rates of American Redstarts (Setophaga ruticilla), Common Yellowthroats (Geothlypis trichas), Orange‐crowned Warblers (Vermivora celata), Wilson's Warblers (Wilsonia pusilla), and Yellow Warblers (Dendroica petechia) varied between years and weeks of the migration period, but were not affected by annual or weekly variation in water levels. Annual variation in capture rates was driven by hatch‐year birds (>80% of individuals captured were juveniles) and could reflect conditions on the breeding grounds that influence productivity. We found that mass gain by the five species of warblers varied between 0.32% and 0.98% of lean body mass/hour. Mass gain did not vary between years or across weeks of the migration period and was not influenced by annual or weekly variation in reservoir water levels. Although the amount of available riparian habitat was reduced when reservoir water levels were high, we found no evidence that this loss of habitat influenced either the number of warblers or the mass gain of warblers using the riparian habitat that remained. Body mass at the time of first capture varied between years and across weeks for all five species. For American Redstarts and Orange‐crowned Warblers, body mass declined as average weekly water levels increased, a pattern that could arise if water levels influenced either their settlement decisions or length of stay.     

Differential habitat use between demographic states of black bears in managed timber forests

The long-term effects of intensive forest harvest on sensitive demographic stages of the American black bear (Ursus americanus) have been often overlooked. Much of Maine, USA, is covered in forests that are hospitable to bears and commercial timber harvest. To investigate the potential effects of differing intensities of disturbance on black bears, and on females with cubs particularly, we designed a large-scale natural experiment with 197 motion-sensitive camera sites dispersed over representative forest stands in northern and central Maine. Using multi-state occupancy models, we distinguished the overall trends in space use by females with young versus adult bears without young. Forest disturbance at large spatial scales was positively associated with the probability of use for both demographic groups and the availability of hardwood trees was an additional important factor for habitat use by females with young. Our study illustrates the use of motion-sensitive cameras to monitor and understand habitat use by distinct life-history stages of animals living in human-modified landscapes, and results indicate that managers can maintain black bear habitat in areas of active forest harvest by ensuring the availability of hardwood species.     

Evidence of a Minimum Patch Size Threshold of Reproductive Success in an Endangered Songbird

ABSTRACT That area-sensitive songbirds breed only in relatively large patches suggests that there may be a minimum patch size threshold in which they will breed, even when controlling for the total amount of habitat in the landscape. We searched for minimum patch size thresholds of presence, territory establishment by males, pairing success, and reproductive success for 2 migratory songbirds that differed in sensitivity to patch size: golden-cheeked warblers (Dendroica chrysoparia) and white-eyed vireos (Vireo griseus). We assessed 2 potential limiting factors: brown-headed cowbird (Molothrus ater) parasitism and arthropod biomass (food resource). We determined whether either factor was related to patch size and compared measurements of each above and below the observed thresholds. We monitored 24 golden-cheeked warbler and 47 white-eyed vireo territories in 12 patches. We found evidence of a minimum patch size threshold (between 15.0 ha and 20.1 ha) of reproductive success for golden-cheeked warblers, but not for white-eyed vireos. We found no minimum patch size thresholds for presence, territory establishment by males, or pair formation for either species. Conservation practices based on thresholds of presence, territory establishment, or pair formation might not address issues of reproduction for golden-cheeked warblers. We failed to find evidence that cowbird parasitism or arthropod biomass were limiting factors. The ability to identify patch size thresholds of reproductive success for target species could be useful in conservation and management for setting goals for retention and restoration of target species’ habitat patch size.     

Eggshell characteristics and yolk composition in the common cuckoo Cuculus canorus: are they adapted to brood parasitism?

The developmental rate of cuckoo embryos and their hatching size is greater than that of host species, which may require more nutrient resources in the egg and more intensive gas exchange during development. In the present study, we compared various egg characteristics of a brood parasite, the common cuckoo Cuculus canorus, and its frequent host, the great reed warbler Acrocephalus arundinaceus. As maternally‐derived testosterone is known to enhance growth rate of embryos and hatchlings, cuckoo eggs are expected to contain higher concentration of testosterone than host eggs. In addition, we expected higher concentration of antioxidants in cuckoo eggs to protect embryos from oxidative stress associated with accelerated growth. Our results showed that cuckoo eggs had thicker shells and higher pore density than great reed warbler eggs. Yolk was significantly heavier in cuckoo eggs and contained higher concentrations of carotenoids and vitamin E, however, yolk androgen and immunoglobulin concentrations were lower in cuckoo eggs as compared to great reed warbler eggs. We also examined whether eggshell colour was associated to egg quality, and detected a positive association between blue‐green chroma and yolk antioxidant concentration in both species, suggesting that eggshell colour reflects the antioxidant investment of the female into the eggs. Our results suggest that cuckoo females increase the size, growth rate and competitive ability of their young by providing them with more nutrients and more dietary antioxidants for embryonic development, and not through elevated yolk testosterone or antibody levels. In addition, increased porosity of cuckoo eggshells may allow embryos to develop more rapidly because of a greater capacity of gas exchange.     

Novel microsatellite loci for the study of the black‐capped vireo (Vireo atricapillus)

We identified 14 novel polymorphic microsatellite loci in the black‐capped vireo (Vireo atricapillus). We also attempted to amplify and genotype these loci in other Vireo species, including the white‐eyed vireo (Vireo griseus), red‐eyed vireo (Vireo olivaceus), and blue‐headed vireo (Vireo solitarius). In 33 genotyped black‐capped vireos from two locations, total alleles ranged from six to 20, with observed heterozygosity ranging from 0.58 to 0.91 and expected heterozygosity from 0.65 to 0.93. Two loci had detectable levels of null alleles. Many of the loci were able to be amplified in the related Vireo species.     

Response of shrubland birds to regenerating clearcut area and shape

Conservation of birds that breed in early-successional forests will require an understanding of their response to patch size and shape during the early seral stages following disturbance. We modeled the effects of patch area, patch shape, and time period following harvest on territory densities of 5 shrubland bird species and bird community composition in 36 clearcuts from 1997 to 2014 in the Missouri Ozarks, USA. Our best-supported models indicated that densities of indigo bunting (Passerina cyanea), yellow-breasted chat (Icteria virens), and prairie warbler (Setophaga discolor) declined from the early time period (1–7 years post-harvest) to the late time period (12–18 years post-harvest), regardless of clearcut area or shape. Yellow-breasted chat density and bird species richness were positively related to clearcut shape complexity. Bird species richness increased, and bird species density decreased with clearcut area during both time periods. Non-metric multidimensional scaling indicated that bird community composition was strongly related to time since harvest. Within the spatial scale of our study, successional stage is more influential than clearcut size and shape, and some bird species may benefit from relatively small or irregularly shaped clearcuts. © 2019 The Wildlife Society.     

Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

Hybrid origin of Audubon's warbler

Several animal species have recently been shown to have hybrid origins, but no avian examples have been documented with molecular evidence. We investigate whether the Audubon’s warbler (Dendroica auduboni), one of four visually distinct species in the yellow‐rumped warbler complex, has originated through hybridization between two other species in this group, the myrtle warbler (D. coronata) and black‐fronted warbler (D. nigrifrons). Analysis of nuclear amplified fragment length polymorphism (AFLP) and sequence markers shows that Audubon’s warblers are genetically intermediate and carry a mixture of alleles otherwise found only in one or the other of their putative parental species. Audubon’s warblers also carry two deeply divergent mitochondrial DNA lineages, each shared with only one putative parental form. Broad clines between Audubon’s and black‐fronted warblers in AFLP markers call into question the validity of these two forms as full species; nevertheless, our results suggest that the Audubon’s warbler probably originated through hybridization between two long‐diverged species. It is likely that more cases of avian species of hybrid origin will be revealed by surveys of variation in nuclear DNA and other traits.     

Western Yellow-Billed Cuckoo Nest-Site Selection and Success in Restored and Natural Riparian Forests

The western distinct population segment of yellow-billed cuckoo (Coccyzus americanus; western cuckoo) has been extirpated from most of its former breeding range in the United States because of widespread loss and degradation of riparian cottonwood (Populus spp.)-willow (Salix spp.) forests. Restoration and management of breeding habitat is important to the recovery of this federally threatened species, and identification of high-quality breeding habitat can help improve the success of recovery. In 2005, the Lower Colorado River Multi-Species Conservation Program, a long-term, multi-agency effort, was initiated to maintain and create wildlife habitat within the historical floodplain of the lower Colorado River (LCR) for federally endangered and threatened species, including western cuckoos. We conducted an empirical, multi-scale field investigation from 2008–2012 to identify habitat characteristics selected by nesting western cuckoos along the LCR. Multiple logistic regression models revealed that western cuckoos selected nest sites characterized by increased densities of small, native, early successional trees measuring 8–23 cm diameter at breast height, and lower diurnal temperature compared to available habitat in restoration and natural forests. Nesting cuckoos selected sites with increased percent canopy closure, which was also important for nest success in restoration sites along the LCR. Our results show habitat components selected by nesting western cuckoos in restoration and natural riparian forests and can help guide the creation, enhancement, and management of riparian forests with habitat conditions necessary to promote nesting of western cuckoos. © 2021 The Wildlife Society.     

REED WARBLER HOSTS FINE‐TUNE THEIR DEFENSES TO TRACK THREE DECADES OF CUCKOO DECLINE

Interactions between avian hosts and brood parasites can provide a model for how animals adapt to a changing world. Reed warbler (Acrocephalus scirpaceus) hosts employ costly defenses to combat parasitism by common cuckoos (Cuculus canorus). During the past three decades cuckoos have declined markedly across England, reducing parasitism at our study site (Wicken Fen) from 24% of reed warbler nests in 1985 to 1% in 2012. Here we show with experiments that host mobbing and egg rejection defenses have tracked this decline in local parasitism risk: the proportion of reed warbler pairs mobbing adult cuckoos (assessed by responses to cuckoo mounts and models) has declined from 90% to 38%, and the proportion rejecting nonmimetic cuckoo eggs (assessed by responses to model eggs) has declined from 61% to 11%. This is despite no change in response to other nest enemies or mimetic model eggs. Individual variation in both defenses is predicted by parasitism risk during the host's egg‐laying period. Furthermore, the response of our study population to temporal variation in parasitism risk can also explain spatial variation in egg rejection behavior in other populations across Europe. We suggest that spatial and temporal variation in parasitism risk has led to the evolution of plasticity in reed warbler defenses.