Biomechanics of biting in loggerhead shrikes: jaw-closing force, velocity and an argument for power.

Differences in the physical and behavioral attributes of prey are likely to impose disparate demands of force and speed on the jaws of a predator. Because of biomechanical trade-offs between force and speed, this presents an interesting conundrum for predators of diverse prey types. Loggerhead shrikes (Lanius ludovicianus) are medium-sized (∼50 g) passeriform birds that dispatch and feed on a variety of arthropod and vertebrate prey, primarily using their beaks. We used high-speed video of shrikes biting a force transducer in lateral view to obtain corresponding measurements of bite force, upper and lower bill linear and angular displacements, and velocities. Our results show that upper bill depression (about the craniofacial hinge) is more highly correlated with bite force, whereas lower bill elevation is more highly correlated with jaw-closing velocity. These results suggest that the upper and lower jaws might play different roles for generating force and speed (respectively) in these and perhaps other birds as well. We hypothesize that a division of labor between the jaws may allow shrikes to capitalize on elements of force and speed without compromising performance. As expected on theoretical grounds, bite force trades-off against jaw-closing velocity during the act of biting, although peak bite force and jaw-closing velocity across individual shrikes show no clear signs of a force-velocity trade-off. As a result, shrikes appear to bite with jaw-closing velocities and forces that maximize biting power, which may be selectively advantageous for predators of diverse prey that require both jaw-closing force and speed.

Do birds select the plastics debris used for nest construction? A case study in a Mediterranean agricultural landscape.

Plastic pollution is becoming a global problem due to its ubiquitous occurrence and the impacts detected for many species. However, the research about plastics in nests of terrestrial bird species has remained relatively overlooked in comparison to those devoted to marine ecosystems. Here we study the occurrence and patterns of use of anthropogenic material in nests of two passerine birds, the Eurasian magpie (Pica pica) and the European serin (Serinus serinus), breeding in an orange tree cultivation in Mediterranean Spain. Our results show that both species use extensively plastic debris as nest material; almost 71% of the European serin nests and 96% of nests of Eurasian magpies contained plastic debris. Furthermore, by analyzing the plastic debris availability in the agricultural landscape surveyed we confirmed a selection pattern in the two species. Thus, both species preferably select plastic filaments over other plastic debris. The Eurasian magpie does not select plastic based on size or color but the European serin avoid black plastics prefer smaller fragments in comparison to the average size available. Moreover, we suggest the apparent similarity of plastic filaments with the natural materials typically used by these species, as well as how they use the plastic in their nests could influence their selection behavior. More studies focused on terrestrial birds inhabiting human modified habitats could offer a deeper approach to how plastic debris interacts with wildlife in different ways.

Avian mud nest architecture by self-secreted saliva.

Mud nests built by swallows (Hirundinidae) and phoebes (Sayornis) are stable granular piles attached to cliffs, walls, or ceilings. Although these birds have been observed to mix saliva with incohesive mud granules, how such biopolymer solutions provide the nest with sufficient strength to support the weight of the residents as well as its own remains elusive. Here, we elucidate the mechanism of strong granular cohesion by the viscoelastic paste of bird saliva through a combination of theoretical analysis and experimental measurements in both natural and artificial nests. Our mathematical model considering the mechanics of mud nest construction allows us to explain the biological observation that all mud-nesting bird species should be lightweight.

On the feeding biomechanics of nectarivorous birds.

Nectar-feeding birds employ unique mechanisms to collect minute liquid rewards hidden within floral structures. In recent years, techniques developed to study drinking mechanisms in hummingbirds have prepared the groundwork for investigating nectar feeding across birds. In most avian nectarivores, fluid intake mechanisms are understudied or simply unknown beyond hypotheses based on their morphological traits, such as their tongues, which are semi-tubular in sunbirds, frayed-tipped in honeyeaters and brush-tipped in lorikeets. Here, we use hummingbirds as a case study to identify and describe the proposed drinking mechanisms to examine the role of those peculiar traits, which will help to disentangle nectar-drinking hypotheses for other groups. We divide nectar drinking into three stages: (1) liquid collection, (2) offloading of aliquots into the mouth and (3) intraoral transport to where the fluid can be swallowed. Investigating the entire drinking process is crucial to fully understand how avian nectarivores feed; nectar-feeding not only involves the collection of nectar with the tongue, but also includes the mechanisms necessary to transfer and move the liquid through the bill and into the throat. We highlight the potential for modern technologies in comparative anatomy [such as microcomputed tomography (µCT) scanning] and biomechanics (such as tracking BaSO4-stained nectar via high-speed fluoroscopy) to elucidate how disparate clades have solved this biophysical puzzle through parallel, convergent or alternative solutions.

Effects of nest-box environment on fledgling success rate and pathogen load.

Nest boxes have been used for many decades as tools for conservation and to study avian population dynamics. Plastic is increasingly used as a material for nest boxes, but no studies have investigated effects of this different material. Two consecutive studies were conducted to investigate effects of nest-box environment on nidicolous parasites, bacteria and fungi, as well as nest success, in blue tits Cyanistes caeruleus and great tits Parus major. The first compared microclimate and parasite and pathogen load in plastic and wooden nest boxes. The second tested the nest protection hypothesis ­ that birds naturally incorporate aromatic herbs into nests to decrease nest parasites and pathogens ­ by comparing parasite and pathogen load in plastic nest boxes to which aromatic or non-aromatic plant material was added. No significant difference in nest-box temperature or relative humidity was found between plastic and wooden boxes. Wooden boxes, however, contained 30-fold higher numbers of fleas and a higher total bacterial load on chicks. Fledging success for blue tit broods was significantly higher in wooden boxes. Parasites and bacteria did not decrease by the inclusion of aromatic herbs. The results increase the evidence base for nest-box design in support of plastic, which can provide an appropriate alternative nest-box material to wood, with apparently no difference in microclimate and no increase in the load of measured parasites and pathogens.

A kestrel without hooked beak and talons is not a kestrel for the red-backed shrike (Lanius collurio).

Birds are strongly motivated to recognize various predators to secure survival and reproductive success. Thus, predator recognition provides a useful tool for the investigation of the process of discrimination and categorization in non-trained animals. The most important role in this process is usually attributed to the prominent unique features shared by all members within the wider category of predators (sharp teeth or beaks, claws, talons and conspicuous eyes). However, birds are also able to discriminate between particular predator species according to their species-specific features (based mainly on colouration). We manipulated general raptor salient features (hooked beak, talons with claws and eyes with supraorbital ridges) and/or species-specific plumage colouration on the dummy of a Eurasian kestrel (Falco tinnunculus) to investigate their importance in the recognition process of the red-backed shrikes (Lanius collurio) during nest defence. We showed that the presence of general raptor salient features is necessary for the recognition of a predator. The kestrel dummy with natural species-specific colouration with altered raptor salient features was treated by shrikes as a harmless bird. Nevertheless, pronounced changes in colouration also prevented successful recognition of the dummy as a kestrel, even when raptor salient features of a raptor were present.

Temperature has a causal and plastic effect on timing of breeding in a small songbird.

Phenotypic plasticity is an important mechanism by which an individual can adapt its seasonal timing to predictable, short-term environmental changes by using predictive cues. Identification of these cues is crucial to forecast the response of species to long-term environmental change and to study their potential to adapt. Individual great tits (Parus major) start reproduction early under warmer conditions in the wild, but whether this effect is causal is not well known. We housed 36 pairs of great tits in climate-controlled aviaries and 40 pairs in outdoor aviaries, where they bred under artificial contrasting temperature treatments or in semi-natural conditions, respectively, for two consecutive years, using birds from lines selected for early and late egg laying. We thus obtained laying dates in two different thermal environments for each female. Females bred earlier under warmer conditions in climate-controlled aviaries, but not in outdoor aviaries. The latter was inconsistent with laying dates from our wild population. Further, early selection line females initiated egg laying consistently ∼9 days earlier than late selection line females in outdoor aviaries, but we found no difference in the degree of plasticity (i.e. the sensitivity to temperature) in laying date between selection lines. Because we found that temperature causally affects laying date, climate change will lead to earlier laying. This advancement is, however, unlikely to be sufficient, thereby leading to selection for earlier laying. Our results suggest that natural selection may lead to a change in mean phenotype, but not to a change in the sensitivity of laying dates to temperature.

Plastic but repeatable: rapid adjustments of mitochondrial function and density during reproduction in a wild bird species.

Most of the energy fluxes supporting animal performance flow through mitochondria. Hence, inter-individual differences in performance might be rooted in inter-individual variations in mitochondrial function and density. Furthermore, because the energy required by an individual often changes across life stages, mitochondrial function and density are also expected to show within-individual variation (i.e. plasticity). No study so far has repeatedly measured mitochondrial function and density in the same individuals to simultaneously test for within-individual repeatability and plasticity of mitochondrial traits. Here, we repeatedly measured mitochondrial DNA copy number (a proxy of density) and respiration rates from blood cells of female pied flycatchers (Ficedula hypoleuca) at the incubation and chick-rearing stages. Mitochondrial density and respiration rates were all repeatable (R = [0.45; 0.80]), indicating high within-individual consistency in mitochondrial traits across life-history stages. Mitochondrial traits were also plastic, showing a quick (i.e. 10 days) downregulation from incubation to chick-rearing in mitochondrial density, respiratory activity, and cellular regulation by endogenous substrates and/or ATP demand. These downregulations were partially compensated by an increase in mitochondrial efficiency at the chick-rearing stage. Therefore, our study provides clear evidence for both short-term plasticity and high within-individual consistency in mitochondrial function and density during reproduction in a wild bird species.

Two new feather mites of the genus Proctophyllodes Robin, 1868 (Acari: Proctophyllodidae) from European passerines (Aves: Passeriformes).

Two new species of the feather mite genus Proctophyllodes Robin, 1868 (Analgoidea: Proctophyllodidae) are described from two passerine birds (Passeriformes) in Europe: Proctophyllodes markovetsi n. sp. from the tawny pipit Anthus campestris (L.) (Motacillidae) and P. loxiae n. sp. from the red crossbill Loxia curvirostra (L.) (Fringillidae). Males of P. markovetsi are most clearly distinguished from the closely related P. tchagrae Atyeo & Braasch, 1966 by having greater terminal lamellae (30-40 × 20-25 µm), the tips of genital arch curved medially, and the corolla of the anal sucker with 14-15 denticles; females of this species are characterised by the terminal appendages distinctly longer than the lobar region width. Males of P. loxiae differ from the closest species, P. fuchsi Mironov, 1997, by having smaller terminal lamellae (45-50 × 22-28 µm), the genital organ extending beyond the posterior margin of lamellae by half their length; females can be distinguished by having the terminal cleft noticeably wider than long (28-30 × 35-40 µm).

Cranial osteology of the genus Sclerurus (Passeriformes: Furnariidae).

The Furnariidae encompasses 293 species and has been recognized as an example of continental adaptive radiation. They inhabit biomes from deserts to humid forests at all strata and show morphological heterogeneity unparalleled among birds at any taxonomic level. Sclerurus is a uniform genus of cryptic, mainly dark brown furnariids, with short black tails which are found solitary on or near the ground inside humid forest. The aim of the present study was to describe and to compare the cranial osteology of all six Sclerurus species (S. scansor, S. mexicanus, S. guatemalensis, S. caudacutus, S. rufigularis, and S. albigularis) to identify osteological characters that are (1) unique to each species, (2) shared among species, and (3) that are exclusive to the genus when compared to other members of Furnariidae. For this, bone structures and measurements were done following standard methodologies. The results showed that Sclerurus differs from other Furnariidae in the following characteristics: a narrowed caudal portion of the nostril with a more rounded shape allowing upper's jaw greater mobility, used when foraging on soft substrates; the development of the post-orbital process may be related to digging behavior, as the presence of a short parsphenoid rostrum projection, a reduced cerebellar prominence, and the tapered caudal portion of the nostrils. Among the species, the interorbital width is larger in S. caudacutus and S. rufigularis, than in the remaining species. The development of the post-orbital process may be related to the behavior of digging nests in earthen banks; the narrowing of the caudal portion of the nostril allows for the greater mobility of the superior maxilla, which is used by Sclerurus when foraging in soft substrates on forest grounds.

Morphological and molecular sexing of the monochromatic Barbados bullfinch, Loxigilla barbadensis.

The bullfinch Loxigilla barbadensis is an endemic passerine on the Caribbean island of Barbados that has only recently been taxonomically split from the Lesser Antillean bullfinch L. noctis. The trait that most clearly distinguishes L. barbadensis from L. noctis is the absence in the male of sexually dimorphic coloration of the body and throat feathers, with L. barbadensis males and females sharing the same dull brown plumage. Here we report, in 64 individuals netted throughout the island, the results of a discriminant analysis on two (wing length and tail length) to four morphological traits showing very high (97%) concordance with sexing via PCR using blood samples. Females also show a paler lower mandible, a trait that yields an 80% concordance with PCR sexing. We found one L. barbadensis male that had a noctis-like reddish throat patch, supporting the idea that sexual dichromatism is the ancestral condition and that male Barbados bullfinches have evolved cryptic coloration that now makes the species monochromatic.

Macroscopic, histologic, and ultrastructural lesions associated with avian keratin disorder in black-capped chickadees (Poecile atricapillus).

An epizootic of beak abnormalities (avian keratin disorder) was recently detected among wild birds in Alaska. Here we describe the gross, histologic, and ultrastructural features of the disease in 30 affected adult black-capped chickadees (Poecile atricapillus). Grossly, there was elongation of the rhamphotheca, with varying degrees of lateral deviation, crossing, and gapping between the upper and lower beak. Not uncommonly, the claws were overgrown, and there was alopecia, scaling, and crusting of the skin. The most prominent histopathologic features in the beak included epidermal hyperplasia, hyperkeratosis, and core-like intrusions of necrotic debris. In affected birds, particularly those with moderate to severe beak overgrowth, there was remodeling of premaxillary and mandibular bones and various dermal lesions. Lesions analogous to those found in beaks were present in affected claws, indicating that this disorder may target both of these similar tissues. Mild to moderate hyperkeratosis occurred in other keratinized tissues, including skin, feather follicles, and, occasionally, sinus epithelium, but typically only in the presence of microbes. We did not find consistent evidence of a bacterial, fungal, or viral etiology for the beak lesions. The changes observed in affected birds did not correspond with any known avian diseases, suggesting a potentially novel hyperkeratotic disorder in wild birds.

Thrushes (Aves: Passeriformes) as indicators of microplastic pollution in terrestrial environments.

Microplastic pollution is one of the leading global conservation issues. The aim of this study was to investigate the occurrence of microplastics in the gastrointestinal tracts of Common Blackbirds Turdus merula (N = 16) and Song Thrushes Turdus philomelos (N = 18), songbirds with exceptionally terrestrial lifestyles and a wide distribution range. We searched for microplastics of over 100 µm in size and assessed whether their contamination differed regarding the age of the birds and the time of year. We used birds that had died as a result of collision with anthropogenic infrastructure, which were sampled during wildlife monitoring of anthropogenic infrastructures and citizen science projects in north-eastern Poland. We found that all the analysed individuals contained microplastic in their gastrointestinal tracts, which were classified as fibers, fragments, films and pellets. A total number of 1073 microplastics were observed, mostly consisting of fibers (84 %) and films (10 %) below 1 mm in size. The dominant colours of microplastics were transparent (75 %) and brown (14 %). The species average microplastic concentration was higher in Song Thrushes (40.1) than Common Blackbirds (21.9), however the difference was not statistically significant. Moreover, we found no seasonal or age-related differences in microplastic ingestion in either species. While our results show a ubiquity of microplastics in terrestrial environments, they also indicate that thrushes have the potential to be used as indicators of microplastic pollution in terrestrial ecosystems.

Microplastic accumulation in the gastrointestinal tracts of nestling and adult migratory birds.

This study examines the abundance and types of microplastic (MP) content in the digestive system of different bird species to help us better understand MP transfer to birds from their environments. The lower GI tracts of six bird species (Tree Swallow (Tachycineta bicolor), Ovenbird (Seiurus aurocapilla), Dark-eyed Junco (Junco hyemalis), Hermit Thrush (Catharus guttatus), Tennessee Warbler (Leiothlypis peregrine), White-throated Sparrow (Zonotrichia albicollis)) were collected for MP analysis. Tree Swallows were collected from nine locations along three drainages in the Milwaukee area and at a remote lake in Wisconsin, USA and consisted of nestlings not yet fledged. The five other bird species were adult migratory bird fatalities from window strikes at the Loyola University Chicago's Lake Shore Campus. Results of plastic analysis showed that the lower GI tracts of all bird species contained different types of MPs (i.e., fibers, fragment, beads). Fiber was the most dominant, followed by fragment and beads. Microplastics were polyethylene, polyethylene terephthalate, nylon, and polyvinyl base materials. There was no statistically significant difference in the site average MP concentration for nestling Tree Swallows or in the species average concentration for adult migratory birds. However, except for Ovenbird, species of adult migratory birds had a species average MP concentration (20.1) that was statistically higher than the site-average MP concentration for Tree Swallow nestlings (5.9). The presence of MPs in the lower GI tracts of unfledged swallows could suggest that MPs had been transferred to the birds from the environment via their diet, such as flying insects-the food source of the birds. Further study should be conducted to investigate the potential transfer of MPs from aquatic insects to birds.

Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions.

Ixodes brevisetosus n. sp. from Papua New Guinea, Ixodes contrarius n. sp. from Costa Rica, Ixodes guglielmonei n. sp. from Argentina and Chile, Ixodes insulae n. sp. from Australia, Ixodes moralesi n. sp. from Guatemala and Panama, Ixodes rio n. sp. from Brazil, Ixodes robbinsi n. sp. from Canada and the United States and Ixodes tinamou n. sp. from Peru (Acari: Ixodidae) are described based on females, nymphs and larvae from various avian hosts (Charadriiformes: Charadriidae; Falconiformes: Falconidae; Galliformes: Cracidae, Odontophoridae, Phasianidae; Passeriformes: Acanthizidae, Cardinalidae, Emberizidae, Fringillidae, Furnariidae, Melanocharitidae, Orthonychidae, Paridae, Passeridae, Petroicidae, Thamnophilidae, Troglodytidae, Turdidae; Strigiformes: Strigidae; Tinamiformes: Tinamidae) and a rodent (Rodentia: Muridae). Females, nymphs and larvae of all new species are similar to those of Ixodes auritulus Neumann, 1904, but can be distinguished by the shape of the scutum and basis capituli, relative length of idiosomal setae, auriculae and internal projection on palpal segment I, degree of development of dorsoapical and mesodorsal spurs on palpal segment I, and measurements and proportions of various structures. Females, nymphs and larvae of I. auritulus s. str. are redescribed based on specimens from passerine birds (Furnariidae, Troglodytidae, Turdidae) in Chile. The subspecies I. auritulus zealandicus Dumbleton, 1961 is elevated to species status, Ixodes zealandicus Dumbleton, 1961. Based on our extensive survey and morphological analyses, we evaluate the actual species richness in the I. auritulus species group and discuss future directions in the systematics of this group of ticks.

Sense and sensitivity: responsiveness to offspring signals varies with the parents' potential to breed again.

How sensitive should parents be to the demands of their young? Offspring are under selection to seek more investment than is optimal for parents to supply, which makes parents vulnerable to losing future fitness by responding to manipulative displays. Yet, parents cannot afford to ignore begging and risk allocating resources inefficiently. Here, we show that parents may solve this problem by adjusting their sensitivity to begging behaviour in relation to their own likelihood of breeding again, a factor largely neglected in previous analyses of parent-offspring interactions. In two carotenoid-supplementation experiments on a New Zealand passerine, the hihi Notiomystis cincta, we supplemented adults to enhance their propensity to breed again, and supplemented entire broods to increase their mouth colour, thus enhancing their solicitation display. We found that adults that attempted two breeding attempts a season were largely insensitive to the experimentally carotenoid-rich gapes of their brood, whereas those that bred just once responded by increasing their rate of provisioning at the nest. Our results show that parents can strategically vary their sensitivity to begging in relation to their future reproductive potential. By restricting opportunities for offspring to influence provisioning decisions, parents greatly limit the potential for offspring to win parent-offspring conflict.

Why the long beak? Phylogeny, convergence, feeding ecology, and evolutionary allometry shaped the skull of the Giant Cowbird Molothrus oryzivorus (Icteridae).

Cowbirds are a successful group of obligate brood parasites in the Neotropical passerine family Icteridae that offer an interesting model to explore the factors behind the evolution of the bird craniomandibular complex. The Giant Cowbird, Molothrus oryzivorus, stands out from its congeners, among other features, in diet (feeds mostly on fruit, nectar, and arthropods, instead on seeds), its larger body size, and longer, more robust beak with a much broader bony casque than in other cowbirds. In turn, Giant Cowbirds show a remarkable resemblance in these features to the distantly related caciques and oropendolas (some are its breeding hosts). However, the causes behind the latter resemblance and the distinctiveness among cowbirds have not yet been elucidated. We aim to explore the factors involved in the diverging morphology of the Giant Cowbird from its congeners and the convergence with caciques and oropendolas, surveying their skull and lower jaw under an explicit evolutionary framework. Using geometric morphometrics and comparative methods, we assessed the signal of phylogeny, convergence, feeding ecology, and size in skull shape. Our results indicated that evolution of the craniomandibular complex of icterids in general, and of the beak morphology in the Giant Cowbird in particular, are shaped by multiple factors, with phylogeny being largely overridden by changes in size (evolutionary allometry), primarily, and feeding ecology, secondarily. However, the evolution of a broad bony casque in the Giant Cowbird, otherwise a hallmark of caciques and oropendolas, does not appear to have primarily been ruled by evolutionary allometry. Instead, taking into account the unique extreme convergence between Giant Cowbirds and some of its caciques hosts, it might be consequence of selective regimes associated with parasite-host interactions acting on top of other evolutionary processes. This suggests chick mimicry as a reasonable explanation for this peculiar morphology that would require further investigation.

Toward a resolution of a taxonomic enigma: first genetic analyses of Paradoxornis webbianus and Paradoxornis alphonsianus (Aves: Paradoxornithidae) from China and Italy.

Paradoxornis webbianus and Paradoxornis alphonsianus naturally occur in South-East Asia. Due to a recent introduction, a mixed population currently occurs in northern Italy. A preliminary phylogeographic analysis using samples from Italy and China found little genetic differentiation between the two taxa and revealed the existence of two molecular lineages, sympatric in some part of China, that do not correspond to the morphological classification. Possible taxonomic changes and preliminary inferences on the relationships between Chinese and the Italian populations and on the likely provenance of the founders introduced in Italy are also discussed.

Chelex-based DNA isolation procedure for the identification of microbial communities of eggshell surfaces.

Microbial communities present on eggshell surfaces of wild birds are weakly studied, especially their influence on embryo infection and, thus, egg viability. Bacterial density of wild bird eggshells is very low, and most DNA extraction protocols are frequently unsuccessful. We have efficiently adapted a chelex-based DNA isolation method for 16S ribosomal gene amplification from the total communities of eggshell surfaces from six avian species.

Mosquito blood-meal analysis for avian malaria study in wild bird communities: laboratory verification and application to Culex sasai (Diptera: Culicidae) collected in Tokyo, Japan.

We conducted laboratory experiments to verify molecular techniques of avian malaria parasite detection distinguishing between an infected mosquito (oocysts on midgut wall) and infective mosquito (sporozoites in salivary glands) in parallel with blood-meal identification from individual blood-fed mosquitoes prior to application to field survey for avian malaria. Domestic fowl infected with Plasmodium gallinaceum was exposed to a vector and non-vector mosquito species, Aedes aegypti and Culex pipiens pallens, respectively, to compare the time course of polymerase chain reaction (PCR) detection for parasite between competent and refractory mosquitoes. DNA of the domestic fowl was detectable for at least 3 days after blood feeding. The PCR-based detection of P. gallinaceum from the abdomen and thorax of A. aegypti corresponded to the microscopic observation of oocysts and sporozoites. Therefore, this PCR-based method was considered useful as one of the criteria to assess developmental stages of Plasmodium spp. in mosquito species collected in the field. We applied the same PCR-based method to 21 blood-fed C. sasai mosquitoes collected in Rinshi-no-mori Park in urban Tokyo, Japan. Of 15 blood meals of C. sasai successfully identified, 86.7% were avian-derived, 13.3% were bovine-derived. Plasmodium DNA was amplified from the abdomen of three C. sasai specimens having an avian blood meal from the Great Tit (Parus major), Pale Thrush (Turdus pallidus), and Jungle Crow (Corvus macrorhynchos). This is the first field study on host-feeding habits of C. sasai in relation to the potential role as a vector for avian malaria parasites transmitted in the Japanese wild bird community.