Neumann’s Warbler Hemitesia neumanni (Sylvioidea): the sole African member of a Palaeotropic Miocene avifauna

We present molecular evidence that Neumann’s Warbler Hemitesia neumanni is deeply nested within the Cettiidae. The species’ distribution in the Albertine Rift of East Africa is intriguing, as the family Cettiidae is principally an Asian radiation. This disjunct distribution could be a result of colonization of Africa by long‐distance dispersal, or the Cettiidae may at some point in the past have had a much larger geographical distribution that also covered parts of Africa.     

Conflicting patterns of mitochondrial and nuclear DNA diversity in Phylloscopus warblers

Molecular variation is often used to infer the demographic history of species, but sometimes the complexity of species history can make such inference difficult. The willow warbler, Phylloscopus trochilus, shows substantially less geographical variation than the chiffchaff, Phylloscopus collybita, both in morphology and in mitochondrial DNA (mtDNA) divergence. We therefore predicted that the willow warbler should harbour less nuclear DNA diversity than the chiffchaff. We analysed sequence data obtained from multiple samples of willow warblers and chiffchaffs for the mtDNA cytochrome b gene and four nuclear genes. We confirmed that the mtDNA diversity among willow warblers is low (pi = 0.0021). Sequence data from three nuclear genes (CHD-Z, AFLP-WW1 and MC1R) not linked to the mitochondria demonstrated unexpectedly high nucleotide diversity (pi values of 0.0172, 0.0141 and 0.0038) in the willow warbler, on average higher than the nucleotide diversity for the chiffchaff (pi values of 0.0025, 0.0017 and 0.0139). In willow warblers, Tajima's D analyses showed that the mtDNA diversity, but not the nuclear DNA diversity, has been reduced relative to the neutral expectation of molecular evolution, suggesting the action of a selective sweep affecting the maternally inherited genes. The large nuclear diversity seen within willow warblers is not compatible with processes of neutral evolution occurring in a population with a constant population size, unless the long-term effective population size has been very large (N(e) > 10(6)). We suggest that the contrasting patterns of genetic diversity in the willow warbler may reflect a more complex evolutionary history, possibly including historical demographic fluctuations or historical male-biased introgression of nuclear genes from a differentiated population of Phylloscopus warblers.     

Morphological traits influence prey selection by coexisting species of New World warblers (Parulidae)

New World wood warblers (Parulidae) represent one of the most dramatic adaptive radiations in North America. However, the ecological bases for the morphological differences among these species remain poorly understood, especially considering how many foraging and habitat studies the family has inspired. We hypothesized the existence of relationships between parulid morphology and diet. We combined a principal component analysis (PCA) of 18 external morphological traits of 11 species of warblers with stomach-content data from coexisting species in one breeding community in Louisiana and three wintering communities in Jamaica. The primary morphological differences, corresponding with the first three PCA axes, were body size, morphological adaptations for aerial foraging versus gleaning, and arboreal versus ground-foraging adaptations. Our analysis revealed little morphological overlap among the 11 species. Differences in diet among the warblers showed a significant relationship to the first two PCA axes of morphological traits. For five coexisting, foliage-gleaning species wintering in Jamaican wet limestone forest, larger warblers ate larger beetles and Orthopterans but not larger ants. In analyses including all four communities, species of warblers with aerial foraging morphologies consumed a greater proportion of winged insects than other warbler species. These findings document prey selection relevant to multiple subtle morphological differences among coexisting species. Overall, our results indicate that food and foraging have likely played an integral role in the morphological diversification and coexistence of species in the family Parulidae.     

Ongoing movement of the hermit warbler X Townsend's warbler hybrid zone

Background

Movements of hybrid zones – areas of overlap and interbreeding between species – are difficult to document empirically. This is true because moving hybrid zones are expected to be rare, and because movement may proceed too slowly to be measured directly. Townsend''s warblers (Dendroica townsendi) hybridize with hermit warblers (D. occidentalis) where their ranges overlap in Washington and Oregon. Previous morphological, behavioral, and genetic studies of this hybrid zone suggest that it has been steadily moving into the geographical range of hermit warblers, with the more aggressive Townsend''s warblers replacing hermit warblers along ∼2000 km of the Pacific coast of Canada and Alaska. Ongoing movement of the zone, however, has yet to be empirically demonstrated.

Methodology/Principal Findings

We compared recently sampled hybrid zone specimens to those collected 10–20 years earlier, to test directly the long-standing hypothesis of hybrid zone movement between these species. Newly sampled specimens were more Townsend''s-like than historical specimens, consistent with ongoing movement of the zone into the geographical range of hermit warblers.

Conclusions/Significance

While movement of a hybrid zone may be explained by several possible mechanisms, in this case a wealth of existing evidence suggests that movement is being driven by the competitive displacement of hermit warblers by Townsend''s warblers. That no ecological differences have been found between these species, and that replacement of hermit warblers by Townsend''s warblers is proceeding downward in latitude and elevation – opposite the directions of range shifts predicted by recent climate change – further support that this movement is not being driven by alternative environmental factors. If the mechanism of competitive displacement is correct, whether this process will ultimately lead to the extinction of hermit warblers will depend on the continued maintenance of the dramatic competitive asymmetry observed between the species.     

Presence of Plasmodium and Haemoproteus in breeding prothonotary warblers (Protonotaria citrea: Parulidae): temporal and spatial trends in infection prevalence

Prothonotary warbler (Protonotaria citrea) has shown a long-term decline in abundance in the United States. As a long-range migrant, these warblers are exposed to parasites in both tropical and temperate regions. The focus of this study was to use molecular techniques to examine the temporal prevalence patterns of heamosopridian parasites Plasmodium and Haemoproteus in breeding prothonotary warblers. The prevalence (presence or absence) of Plasmodium and Haemoproteus species was assayed using primer sets for the cytochrome b gene of the mitochondrial DNA. Blood samples were obtained from 187 adult prothonotary warblers collected at 3 central Virginia, U.S.A., breeding sites. The relationship between haemosporidian parasite infections and reproductive success also was examined. We found that 71% of captured prothonotary warblers were infected with haemosporidian parasites, specifically, with 36% prevalence for Haemoproteus spp. and 44% prevalence for Plasmodium spp., during the 2008 breeding season; for both parasites, prevalence increased throughout the season. We found significant variation in haemosporidian parasite prevalence across the breeding season that was strongly site specific. Conversely, we found no significant effects of haemosporidian parasite infections on the reproductive success of prothonotary warblers. This is in sharp contrast to recent reports suggesting considerable effects of these parasites on the reproductive success of wild birds.     

Potential Role of Spruce Budworm in Range-Wide Decline of Canada Warbler

ABSTRACT The Canada warbler (Wilsonia canadensis) is one of many common neotropical migrants whose populations are in decline across their range. Influences of habitat loss and degradation on breeding or wintering grounds have been postulated as possible causes, but few empirical data exist to support a specific cause. Based on previous studies linking abundances of Canada warbler and spruce budworm (Choristoneura fumiferana), we hypothesized that the Canada warbler may be influenced by a persistent decline in spruce budworm throughout the bird's breeding range, a hypothesis that has received little attention. This hypothesis makes 5 predictions: 1) budworm outbreaks and warbler detections should be spatially and temporally coincident; 2) the relationship between Canada warbler and spruce budworm outbreaks should be similar to relationships for other warblers known to be spruce budworm associates; 3) the relationship should be stronger than for warblers lacking an association with spruce budworm; 4) because temporal trends of both spruce budworm and Canada warblers have varied throughout Canadian provinces, declines in Canada warblers should be seen only in provinces where spruce budworm also declined; and 5) variation in Canada warbler abundance should reflect variation in supply of preferred habitat for the spruce budworm if habitat rather than budworm abundance is the key. Our analyses supported predictions 1–4, suggesting that Canada warbler may be even more closely associated with spruce budworm than are known associated species, a phenomenon noted in the literature but previously unexplained. Prediction 5 was not supported, because budworm habitat (area of mature and older balsam fir [Abies balsamea] and white spruce [Picea glauca]) remained constant in Ontario while warbler abundance declined. Although the correlative nature of these results precludes inference of a causal relationship between the declines of the Canada warbler and spruce budworm, we postulate that potential links may exist directly, where spruce budworm outbreaks provide elevated levels of insect prey items for breeding Canada warblers, or indirectly through changes in forest structure and composition following outbreaks. These results have implications when considering long-term trends in Canada warbler populations, because it may be impossible to alter population trends for species linked to the timing and magnitude of spruce budworm outbreaks.     

Plumage and mitochondrial DNA haplotype variation across a moving hybrid zone

We analyze variation in phenotypes and mitochondrial DNA (mtDNA) haplotypes over the breeding ranges of hermit and Townsend's warblers and across two of their three hybrid zones. Within these two hybrid zones, we demonstrate that the placement, shape, and width of transitions in seven plumage characters are remarkably similar, suggesting that a balance between dispersal and sexual selection keeps these hybrid zones narrow. A consistent asymmetry in these character transition curves suggests that Townsend's warblers have a selective advantage over hermit warblers, which is presumably due to the aggressive superiority of Townsend's over hermit males (Pearson and Rohwer 2000). An association between plumage and mtDNA haplotypes shows that pure Townsend's warblers, but not pure hermit warblers, immigrate into these hybrid zones, further supporting the competitive superiority of Townsend's warblers over hermit warblers. The mitochondrial haplotype transitions across these hybrid zones are much wider than the phenotypic transitions and provide no indication that the mtDNA haplotypes representing these two warblers are selectively maintained. More importantly, the phenotypically pure populations of Townsend's warblers throughout a 2,000-km coastal strip north of the Washington hybrid zones contain a preponderance of hermit warbler mtDNA haplotypes. This result suggests massive movement of the hybrid zone between these warblers during the 5,000 years since their most recent interglacial contact. We develop a model to explain the phenotypic and genetic divergence between these warblers and the evolution of their dramatic differences in aggressiveness; we also show how differences in male aggression, in combination with biased pairing patterns, can explain the haplotype footprint recording the historical movement of this hybrid zone.     

Intermediate habitat associations by hybrids may facilitate genetic introgression in a songbird

Hybridization or the interbreeding of genetically discrete populations or species can occur where ranges of genetically distinct units overlap. Golden‐winged warblers Vermivora chrysoptera, a species that has been in steady decline for decades, highlight the potential population‐level consequences of hybridization. A major factor implicated in their decline is hybridization with their sister species, the blue‐winged warbler Vermivora cyanoptera, which has likely been exacerbated by historic and current land‐use practices. We examined habitat associations of golden‐winged and blue‐winged warblers, phenotypic hybrids, and cryptic hybrids (i.e. mismatch between plumage phenotype and genotype as identified by mitochondrial DNA) in an area of relatively recent range overlap and hybridization in northern New York, USA. To explore the robustness of these results, we then compared the patterns from New York with habitat associations from the central Pennsylvanian Appalachian Mountains where blue‐winged warblers either do not occur or are in very low abundance, yet cryptic golden‐winged warbler hybrids are present. From 2008 to 2011, we captured 122 birds in New York and 28 in Pennsylvania and collected blood samples, which we used to determine maternal ancestry. For each bird captured, we measured territory‐level (50‐m radius circles) habitat, and later used remote‐sensing data to quantify habitat on the territories and in surrounding areas (100‐, 250‐, and 500‐m radius circles). In New York, golden‐winged warblers occupied structurally heterogeneous territories surrounded by homogeneously structured, contiguous deciduous forest, far from urban areas. Blue‐winged warblers showed opposite associations, and hybrids’ habitat associations were typically intermediate. In Pennsylvania, the habitat associations of golden‐winged warblers and their cryptic hybrids were remarkably similar to those in New York. These findings suggest that patterns of habitat occupancy by hybrids may promote contact with golden‐winged warblers and thus likely facilitate genetic introgression, even in areas where the parental species are not sympatric.     

Asymmetries in male aggression across an avian hybrid zone

Recent studies suggest that competitive asymmetries are causingthe hybrid zones between hermit and Townsend's warblers to move,such that Townsend's warblers are replacing hermit warblers.Here we examine the contribution of male aggression to thiscompetitive asymmetry by measuring aggressive responses to mounts.We presented male mounts of the two parental species to Townsend'sand hermit warblers outside the zone and to hybrids within the zone.Outside the zone, Townsend's males are more aggressive to both conspecificand heterospecific mounts than are hermit males. This asymmetry shouldmove the zone in the direction inferred from previous studies.Hybrids fall between parentals in their aggressiveness, whichshould accelerate the movement of the zone. Remarkably, we foundno relationship between phenotype and aggression in individualmales at a locality within the hybrid zone. The forces of selectionand dispersal that maintain narrow hybrid zones should generatesuch a correlation if aggressive differences between the parental speciesare genetically controlled. We resolve this conflict by proposinga behavioral model of competitive sorting within localities.If birds are sorted across the hybrid zone according to competitiveabilities, and competitive interactions within neighborhoodsare more or less complete, then the correlation between phenotypeand aggression within any given neighborhood will be eliminated.We tested this model by examining the relationship between phenotypeand aggression across the zone. Warblers in hybrid neighborhoodson the Townsend's side of the zone are more aggressive thanwarblers in hybrid neighborhoods on the hermit side, indicatingthat competitive sorting is occurring.     

Breeding latitude and timing of spring migration in songbirds crossing the Gulf of Mexico

Each spring, millions of songbirds migrate across the Gulf of Mexico on their way to breeding sites in North America. Data from radar and migration monitoring stations have revealed broad patterns in the spatial and temporal course of trans-Gulf migration. Unfortunately, we have limited information on where these birds have previously spent the winter and where they are migrating to breed. Here we measure stable-hydrogen isotopes in feathers (δD_f) to infer the breeding latitude of five species of songbirds – hooded warblers Wilsonia citrina , American redstarts Setophaga ruticilla , black-and-white warblers Mniotilta varia , ovenbirds Seiurus aurocapilla , and northern waterthrushes S. noveboracensis – that were captured at a stopover site along the coast of southwestern Louisiana in spring 2004. Values of δD_f across all species ranged from −163 to −35‰ (n=212), and within most species the range was consistent with the latitudinal extent of known breeding sites in central and eastern North America. Individuals that arrived first along the northern Gulf coast had δD_f values indicative of southerly breeding sites in hooded warblers, American redstarts, black-and-white warblers, and ovenbirds, but no relationship was found between passage timing and δD_f for northern waterthrushes. Our findings suggest that spring passage is often timed to coincide with the emergence of suitable conditions on breeding areas, with southern breeding birds migrating first.     

Importance of Montane Riparian Forest and Influence of Wildfire on Nest-Site Selection of Ground-Nesting Birds

Abstract: We studied breeding populations of 2 coexisting ground-nesting birds, the red-faced warbler (Cardellina rubrifrons) and yellow-eyed junco (Junco phaeonotus), in high-elevation (>2,000 m) forested drainages of the Santa Catalina Mountains, Arizona, USA. From 2004 to 2005, we 1) estimated density and nesting success of breeding populations of red-faced warblers and yellow-eyed juncos, 2) identified nest-site characteristics for each species (i.e., used sites vs. random plots), 3) compared nest-site characteristics between the 2 species, and 4) examined effects of a recent (2003) wildfire on distribution of nests of both species. In addition, we estimated the areal extent of montane riparian forest (the preferred breeding habitat of both species) within high-elevation forests of the Santa Catalina Mountains. We found that red-faced warblers and yellow-eyed juncos were the 2 most common ground-nesting birds within our study area with an average density of 2.4 and 1.4 singing males/ha, respectively, along drainage bottoms. Compared to random plots, most red-faced warbler and yellow-eyed junco nests were located close (≤30 m) to drainage bottoms within a strip of montane riparian forest characterized by abundant brush, small woody debris, ferns, and forbs (both species), high number and diversity of saplings and small trees (red-faced warblers), and abundant shrubs and downed logs and less canopy cover (yellow-eyed juncos). Although both species nested in close proximity within montane riparian forest, nest-site characteristics differed between the 2 species, especially at finer spatial scales. For example, most yellow-eyed juncos nested adjacent to grass (principally Muhlenbergia spp.), whereas red-faced warblers nested adjacent to a variety of plant species, including grass, bigtooth maple (Acer grandidentatum), white fir (Abies concolor), and Douglas-fir (Pseudotsuga menziesii). Both red-faced warblers and yellow-eyed juncos avoided nesting in areas burned during a recent wildfire. In addition, nesting success was low for red-faced warblers (13%) and yellow-eyed juncos (19%) following the wildfire, suggesting an indirect negative effect of fire on breeding populations in the short-term. Montane riparian forest appears to provide important breeding habitat for red-faced warblers and yellow-eyed juncos. However, little research or conservation planning has been directed toward montane riparian forest in the region, even though this forest type is limited in its areal extent (<11% of high-elevation forest in the Santa Catalina Mountains) and increasingly threatened by disturbance. Results from our study can be used to facilitate the management and conservation of breeding populations of red-faced warblers and yellow-eyed juncos in high-elevation forests of the southwestern United States.     

Experiments and observations of prothonotary warblers indicate a lack of adaptive responses to brood parasitism

It has been suggested that prothonotary warblers, Protonotaria citrea, respond adaptively to brood parasitism by brown-headed cowbirds, Molothrus ater, even though they lack historical habitat and range overlap with cowbirds. I studied behaviours functioning as potential defences against brood parasitism in the prothonotary warbler, a cavity-nesting host species. Opening sizes preferred by prothonotary warblers were not small enough to exclude cowbirds, and warblers were parasitized heavily in nests with larger openings. Male and female prothonotary warblers were always away from their nests before sunrise when cowbirds laid eggs in their nests. Prothonotary warblers infrequently (∼6% of 560 nests) deserted nests that were parasitized during the egg-laying period, but frequently (56% of 151 nests) deserted nests that were parasitized before a female warbler laid her first egg. Prothonotary warblers also deserted 60-70% of nests where a cowbird egg, warbler egg or die were experimentally added before egg laying. However, the experimental addition of one of these three objects during the egg-laying period did not elicit desertion. The desertion of parasitized nests was not affected by nest site availability as has been reported elsewhere in the literature. This lack of a response to brood parasitism by prothonotary warblers may be an example of evolutionary lag, because it is likely that they have only recently been exposed to widespread parasitism, and they accept parasitism at a high cost to their own reproductive success. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Hostile encounters among spruce-woods warblers (Dendroica: Parulidae)

Intraspecific interactions were initiated with high frequency by four species of sympatric warblers (Dendroica: Parulidae) early in the breeding season. They declined during the incubation and nestling stages and then increased during the fledging stage in two species, the black-throated green (Dendroica virens) and yellow-rumped (D. coronata) warblers. Interspecific interactions occurred infrequently early in the season, but increased during the incubation and nestling stages in the black-throated green and blackburnian (D. fusca) warblers and during the fledging stage in the magnolia warbler (D. magnolia). Fights and chases predominated over supplanting attacks in both intraspecific and interspecific encounters. Male black-throated green and yellow-rumped warblers initiated hostile encounters more frequently than did their females, but male and female blackburnian and magnolia warblers did not differ in this regard. A somewhat equivocal interspecific social dominance hierarchy occurred: magnolia > black-throated green > yellow-rumped > blackburnian. Reversals to this trend were frequent and may have resulted from the strong defences initiated by these species in certain parts of their territories, particularly about their nests.     

Blood parasites in migrating wood‐warblers (Parulidae): effects on refueling,energetic condition,and migration timing

Blood parasites can negatively affect energetic condition, arrival date, and reproductive performance of breeding birds, yet their impact on migrating birds is poorly understood. We quantified haematozoan infection and examined its relationship to migration timing, energetic condition, and refueling rate (plasma triglyceride concentration) for three species of wood‐warblers (Parulidae) migrating through northwestern Ohio, USA from 11 April–3 June of 2004 and 2005. Prevalence of infection for magnolia Dendroica magnolia (36.3%; n=383), yellow‐rumped D. coronata (58.3%; n=518), and yellow warblers D. petechia (12.7%; n=205) was approximately two times higher than reported in other migration studies and similar to infection rates found in breeding areas. Probability and intensity of infection were higher for later migrating yellow‐rumped warblers, especially in younger (second‐year) birds. Furthermore, we found a negative trend between intensity of infection and energetic condition for second‐year yellow‐rumped warblers. Haematozoan infection was not associated with refueling rate for any species. Our results indicate that for some migrants, especially younger birds, probability and intensity of haematozoan infection are related to later migration timing and reduced energetic condition during spring migration, and thus may be important determinants of subsequent reproductive performance in breeding areas.     

Nestling sex ratio of golden‐winged warblers Vermivora chrysoptera in an introgressed population

Sex ratio biases in avian species remain controversial, although several studies have documented apparent facultative adjustment of offspring sex ratios. While hybridizing pied and collared flycatchers have exhibited sex ratio skews that may be a response to sex‐based costs associated with hybridization, this appears not to be true of a hybridized population of blue‐winged Vermivora pinus and golden‐winged V. chrysoptera warblers. We examined the primary sex ratio of nestlings in a population of hybrid and introgressed golden‐winged warblers. The sex ratio of 298 nestlings from 81 nests in the population was approximately 50:50. We conducted paternity assignments and analyzed groups of nestlings with shared genetic parents (“genetic broods”) and found no difference from the expected binomial distribution, and no statistically significant relationship between parental species phenotype and nestling sex ratio. We saw no evidence of preferential production of male or female nestlings, and female hybrids were found to mate and breed in the population. This suggests that heterogametic (female) hybrids are both viable and fertile, and thus that Haldane's Rule does not apply to this system. While populations of hybridizing golden‐winged warblers should be monitored for evidence of costs of heterospecific pairings, it is unlikely that adjustment of sex ratios would be the form of compensation for sub‐optimal mating conditions. Our results provide support for the emerging hypothesis that hybrids suffer no disadvantage relative to golden‐winged and blue‐winged warblers.     

A phylogeny for the Cisticolidae (Aves: Passeriformes) based on nuclear and mitochondrial DNA sequence data, and a re-interpretation of an unique nest-building specialization

Based on some general similarities in feeding adaptations, a large number of Old World passerine birds were in the past lumped in one broad family, the Sylviidae. Recent molecular studies, starting with the DNA-DNA hybridization work by Sibley et al. [Sibley, C.G., Ahlquist, J.E., 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution, Yale University Press, New Haven, CT], have revealed that this group is in fact a paraphyletic assemblage, mainly in the superfamily Sylvioidea, and within this assemblage a distinct group (the Cisticolidae) can be identified around the genus Cisticola. In this study we try to define natural lineages within it, based on DNA sequence data from 35 ingroup taxa representing 12 putative genera. Both nuclear myoglobin intron II (630 bp in our study) and mitochondrial ND2 (1041 bp) genes were sequenced, and 1671 bp were aligned and subjected to parsimony, maximum likelihood and Bayesian analyses. The results strongly support the monophyly of a cisticolid clade, with the Malagasy warblers Neomixis constituting the deepest branch within the clade. Three major clades receive statistical support, but not all relationships between and within these are well resolved. All species of the genus Bathmocercus belong to the Cisticolidae but in two different clades. The tailorbirds appear also polyphyletic with most species of the genus Orthotomus (but O. cucullatus falling in the outgroup) and the African metopias being in two different clades. Also the genus Apalis is polyphyletic, but all other included genera seem to be confirmed as natural units. Based on these findings we resurrect the genera Scepomycter and Artisornis. Calamonastes is confirmed to be in the Cisticolidae and grouped with Camaroptera. Main basic nest types do not follow the phylogenetic branching, and notably the peculiar "tailorbird" technique of stitching leaves together around the nest is found in different parts of the phylogeny. The basic types of nests seem to be found in particular environments, and the sewing may therefore have evolved in some ancestor of the Cisticolidae and was later lost or modified in some genera or species following the spread of drier habitats from the mid-Miocene.     

Rejection of artificial cuckoo (Cuculus canorus) eggs in relation to variation in egg appearance among reed warblers (Acrocephalus scirpaceus)

Passerines that are exposed to brood parasitism can evolve reduced intraclutch variation in egg appearance to facilitate recognition and rejection of the parasitic egg. This has been shown to be true for European passerine species that are assumed to have participated in an evolutionary arms race with the cuckoo (Cuculus canorus). However, few investigations have been carried out with the aim of finding out whether there is a relationship between these two traits within a species. In this study, we compare the level of intraclutch variation in egg appearance and the rejection of an unlike parasitic egg within a population of reed warblers (Acrocephalus scirpaceus) in the south-eastern part of the Czech Republic. We parasitized reed warbler nests with an artificial non-mimetic cuckoo egg, and then monitored the reaction of the hosts. In 27 out of 48 nests (56.3%) the parasitic egg was rejected. The rejecter pairs had a statistically significantly lower intraclutch variation in egg appearance than the acceptor pairs. We discuss possible explanations for the observed relationship between rejection of unlike eggs and intraclutch variation in egg appearance within this population of reed warblers. The results are consistent with the evolutionary arms race hypothesis, but the intermediate rejection rate found in this population could also be maintained by an equilibrium between acceptors and rejecters due to rejection costs.     

Habitat and Landscape Effects on Brood Parasitism,Nest Survival,and Fledgling Production in Swainson's Warblers

ABSTRACT Numerous factors, including nest predation and brood parasitism, may limit populations of neotropical migratory birds. However, nest predation and brood parasitism are not constant, and temporal, biological, habitat, and landscape factors can affect the likelihood of these events. Understanding these patterns is important for species of conservation concern for which managers seek to provide quality habitat. One such species, the Swainson's warbler (Limnothlypis swainsonii), is a neotropical migrant that breeds primarily in bottomland hardwood forests of the southeastern United States. Little is known of factors influencing reproductive success of this rare, yet locally abundant, species. From 2004 through 2007, we examined factors influencing reproductive success of Swainson's warblers at 2 sites in eastern Arkansas, USA, St. Francis National Forest and White River National Wildlife Refuge. We used 2-stage modeling to assess the relationship between 1) temporal and biological, and 2) habitat and landscape factors and brood parasitism, nest survival, and fledgling production. Brood parasitism was greater in this population (36%) than reported elsewhere (≤ 10%), but decreased throughout the breeding season. Nest survival was comparable to or lower than in other populations of this species and increased throughout the breeding season. The probability of brood parasitism was greater near forest edges. Although nests of Swainson's warblers were often associated with giant cane (Arundinaria gigantea), nest survival had a weak negative association with cane density. For nests that were successful, the best predictor of number of Swainson's warblers fledged was brood-parasitism status: nonparasitized nests fledged 2.75 young, whereas parasitized nests fledged 0.60 Swainson's warblers. Our findings suggest that managing and restoring relatively high-elevation bottomland forests that are located far from agricultural edges should increase Swainson's warbler productivity.