Ecological correlates of feather mite prevalence in passerines

The relationship between host ecology and feather mite prevalence was analysed in birds. Feather mites are small arthropods (fam. Pterolichoidea and Analgoidea) commonly found on birds, although the nature of their interactions with the host (commensalism, mutualism or parasitism), still remains unclear. Host body mass and migratory behaviour were unrelated to feather mite prevalence. Contrary to expectation, there was no differences in mite prevalence between colonial and solitary-breeding species. However, winter sociality was associated with increased prevalence, suggesting that winter and breeding sociality affected the distribution patterns of feather mites in different ways. Plumage dichromatism was negatively correlated with feather mite prevalence, a result that is opposite to that predicted by the Hamilton and Zuk hypothesis for the evolution of host secondary sexual characteristics in relation to parasitism.     

DNA barcoding and minibarcoding as a powerful tool for feather mite studies

Feather mites (Astigmata: Analgoidea and Pterolichoidea) are among the most abundant and commonly occurring bird ectosymbionts. Basic questions on the ecology and evolution of feather mites remain unanswered because feather mite species identification is often only possible for adult males, and it is laborious even for specialized taxonomists, thus precluding large‐scale identifications. Here, we tested DNA barcoding as a useful molecular tool to identify feather mites from passerine birds. Three hundred and sixty‐one specimens of 72 species of feather mites from 68 species of European passerine birds from Russia and Spain were barcoded. The accuracy of barcoding and minibarcoding was tested. Moreover, threshold choice (a controversial issue in barcoding studies) was also explored in a new way, by calculating through simulations the effect of sampling effort (in species number and species composition) on threshold calculations. We found one 200‐bp minibarcode region that showed the same accuracy as the full‐length barcode (602 bp) and was surrounded by conserved regions potentially useful for group‐specific degenerate primers. Species identification accuracy was perfect (100%) but decreased when singletons or species of the Proctophyllodes pinnatus group were included. In fact, barcoding confirmed previous taxonomic issues within the P. pinnatus group. Following an integrative taxonomy approach, we compared our barcode study with previous taxonomic knowledge on feather mites, discovering three new putative cryptic species and validating three previous morphologically different (but still undescribed) new species.     

Phylogeny and co-speciation in feather mites of the subfamily Avenzoariinae (Analgoidea: Avenzoariidae)

A cladistic analysis of phylogenetic relationships is carried out for the feather mite subfamily Avenzoariinae. The analysis is made at two different taxonomic levels, for 19 genera of the all family Avenzoariidae and for taxa of species rank for the Avenzoaria and Bychovskiata generic groups. A subsequent comparative analysis of phylogenetic hypotheseis for the subfamily Avenzoariinae and recently accepted phylogenetic hypotheses of the shorebirds Charadriiformes indicates co-speciation of feather mites with their hosts. As a result of the comparative analysis it is suggested, that the subfamily Avenzoariinae originated from an ancestor of the order Charadriiformes and co-speciated with this host order. The expected pattern of parallel evolution is disturbed by different evolutionary events, such as host shifts, extinction of mites and differential evolutionary rates of mite lineages in different phyletic branches of feather parasites.     

Dynamics of infection of Fringilla coelebs chaffinch nestlings with feather mites (Acari: Analgoidea)

A process of infecting the chaffinch nestlings Fringilla coelebs with three analgoid feather mites, Analges passerinus L., 1758, Monojoubertia microphylla (Robin, 1877), and Pteronyssoides striatus (Robin, 1977), commonly occurred on this bird species was investigated. 15 nests contained totally 65 nestlings, from 2 to 6 individuals in a brood, have been examined from the day of hatching till 11th day. Observations were held in the neighbourhood of the bird banding station "Rybachy" (Russia, Kaliningrad Province) in June of 1982. Number of mites on alive nestlings taken temporarily from their nest was counted by means of binocular lens under the magnification x12.5 and x25. The nestlings receive the mites from the chaffinch female during the night time, when the female sits together with the young birds and heats them. In the condition of this prolonged direct contact the mites migrate from the female onto the nestlings. As it was shown in our study of seasonal dynamics of mites on the chaffinch (Mironov, 2000), the chaffinch female only gives its mites to young generation and looses about three quarter of its mite micropopulation during the nesting period (June), hile in the chaffinch males the number of mites continues to increase during all summer. The infections with three feather mite species happen in the second part of the nestling's stay in the nest. The starting time of this process, its intensity, and sex and age structure of mite micropopulations on the nestlings just before their leaving the nest are different in the mite species examined. These peculiarities of feather mite species are determined by the biology of examined species, and first of all by their morphological characteristic and specialisation to different microhabitats, i.e. certain structural zones of plumage. Pteronyssoides striatus (Pteronyssidae) is rather typical mite specialised to feathers with vanes. In adult birds with completely developed plumage this species occupies the ventral surface of the big upper coverts of primary flight feathers. This species appears on the chaffinch nestlings in a significant number on 7th day. The mites occupy the basal parts of primary flight feathers represented in that moment by the rods only. They sit on practically open and smooth surface of this microhabitat, which is uncommon for them, because the vanes of the big upper coverts are not yet open and also represented by thin rods. During the period of the last 5 days (from 7 to 11th day) the mean number of mites per one nestling increases from 2.3 +/- 0.5 to 17.1 +/- 1.8 mites. Just before the day, when the nestling leave the nest, the tritonymphs absolutely predominate (82.4%) in the micropopulation of P. striatus. Analges passerinus (Analgidae) is specialised to live in the friable layer formed by numerous not-engaged thread barbles of the down feathers and basal parts of the body covert feathers. Mites have special hooks on legs used for hard attaching to the barbles and for fast moving in the friable layer of feathers. On the chaffinch nestlings, these mites appear usually on 8th day, when the rod-like body covert feathers begin to open on apices and form short brushes; however some individuals occur on the skin of nestlings even on 6th day. The mean number of mites per nestling on the 11th day reaches 16.5 +/- 1.4 individuals. The micropopulation of A. passerinus is represented on the nestlings mainly by the females (45.5%), tritonymphs (23.6%) and males (11.5%). Monojobertia microphylla (Proctophyllodidae) is a typical dweller of feathers with large vanes. Mites of this species commonly occupy the ventral surface of primary and secondary flight feathers and also respective big upper covert feathers of wings. M. microphylla appears on the nestlings in a significant number (7.1 +/- 1.2 mites) on 9th day, only when the primary flight feathers already have short vanes about 10 mm in length. In next three days the number of mites increases very fast and reaches on 11th day 60.3 +/- 5.7 mites per nestling. In the micropopulation of this species, the tritonymphs count 38.3%, and the quota of males and females is 25.3% each. The migration of this species goes most intensively, than in two other species. An analitic selection of logistic curves shows, that the increasing of mite number during the process of infection with three mite species may be most adequately described by the sigmoid curves with clearly recognizable levels of saturation, which can be theoretically reached. Indeed, the number of mite individuals being able to migrate onto the nestlings is limited by their number on a respective chaffinch female. In a contrast, the increasing of plumage indices, for instance the length of flight feathers, has almost linear character during the period of observation. The beginning of mite migration is determined by the development of respective microhabitats in the plumage of nestlings, or at least by the development of certain structure elements of plumage, where mites are able to attach for a while, before that moment, when the nestlings will develop the plumage completely and begin to fly. In three mite species examined, the process of infection was performed by older stages, namely by the imago and/or tritonymphs. This can be explained by two reasons. On the one hand, the older stages are most active in their movement, resistible and able to survive successfully on new host individuals. On the other hand, the older stage are ready for the reproduction or will be ready after one moulting. The older stages of mites can quickly create a large and self-supporting micropopulations on the birds, therefore this strategy ensures a successful subsequent existence of the parasite species. In cases, when mites (A. passerinus, M. microphylla) migrate into the respective microhabitats structurally corresponding to their normal microhabitats on adult birds, the micropopulations of these mite species include a significant or dominant quota of females and males. When the normal microhabitat is not yet formed, feather mites migrate into neighboring structure elements of plumage, where they can survive and wait for the development of normal microhabitat, to which they are well adapted. Therefore, in the case of P. striatus, its micropopulations on the chaffinch nestlings are represented mainly by the tritonymphs.     

Feather mites on birds: costs of parasitism or conditional outcomes?

Feather mites (suborder Astigmata, superfamilies Analgoidea, Pterolichoidea and Freyaniidae) are among the commonest ectosymbionts of birds. Most researchers have assumed they are parasites, having negative effects on hosts. Here we present evidence that suggests that feather mites may not be parasites. We develop a framework for considering conditional outcomes in these interspecific associations, dealing with different kinds of relationships between symbionts. The non-parasitic status of feather mites is supported by a literature review as well as by preliminary data on mites' food. We illustrate symbiotic relationships with a graphical model showing different scenarios in which hosts' cost-benefit relations are determined by the interactions among their symbionts.     

Feather mite abundance increases with uropygial gland size and plumage yellowness in Great Tits Parus major

Plumicolous feather mites are ectosymbiotic organisms that live on bird feathers. Despite their abundance and prevalence among birds, the ecology of the interaction between these organisms and their hosts is poorly known. As feather mites feed on oil that birds spread from their uropygial gland, it has been hypothesized, but never tested, that the number of feather mites increases with the size of the uropygial gland of their hosts. In this study the number of feather mites is considered with respect to uropygial gland size in a breeding population of Great Tits Parus major in order to test this hypothesis. As predicted, the number of feather mites correlated positively with the uropygial gland size of their hosts, showing for the first time that uropygial gland size can explain the variance in feather mite load among conspecifics. Previous studies relating feather mite load to plumage colour have suggested that feather mites may be parasitic or neutral. To confirm this, the yellowness of breast feathers was also assessed. However, the results ran in the opposite direction to that expected, showing a positive correlation between mite load and plumage yellowness, which suggests that further work is needed to give clear evidence for a specific nature of feather mites. However, Great Tits with higher mite loads had lower hatching and breeding success, which may support the idea that feather mites are parasites, although this effect must be taken with caution because it was only found in males. Age or sex effects were not found on the number of feather mites, and it is proposed that hormonal levels may not be sufficient to explain the variation in feather mite loads. Interestingly, a positive correlation was detected between uropygial gland size and plumage brightness, which could be a novel factor to take into account in studies of plumage colour.     

Different scales of spatial segregation of two species of feather mites on the wings of a passerine bird

The "condition-specific competition hypothesis" proposes that coexistence of 2 species is possible when spatial or temporal variations in environmental conditions exist and each species responds differently to those conditions. The distribution of different species of feather mites on their hosts is known to be affected by intrinsic host factors such as structure of feathers and friction among feathers during flight, but there is also evidence that external factors such as humidity and temperature can affect mite distribution. Some feather mites have the capacity to move through the plumage rather rapidly, and within-host variation in intensity of sunlight could be one of the cues involved in these active displacements. We analyzed both the within- and between-feather spatial distribution of 2 mite species, Trouessartia bifurcata and Dolichodectes edwardsi , that coexist in flight feathers of the moustached warbler Acrocephalus melanopogon. A complex spatial segregation between the 2 species was observed at 3 spatial levels, i.e., "feather surfaces," "between feathers," and "within feathers." Despite certain overlapping distribution among feathers, T. bifurcata dominated proximal and medial regions on dorsal faces, while D. edwardsi preferred disto-ventral feather areas. An experiment to check the behavioral response of T. bifurcata to sunlight showed that mites responded to light exposure by approaching the feather bases and even leaving its dorsal face. Spatial heterogeneity across the 3 analyzed levels, together with response to light and other particular species adaptations, may have played a role in the coexistence and segregation of feather mites competing for space and food in passerine birds.     

The phenomenon of phylogenetic synhospitality in acariform mites (acari: acariformes)--the permanent parasites of vertebrates

The term synhospitality means the association of two or more closely related parasite species with one host species (Eichler, 1966). The cases of two or three synhospitalic species are known from the same host species, and especially ones where parasites were recorded from different parts of the host range, are quite common. The most ordinary reason causing synhospitality in permanent parasites is the host switching. Nevertheless, there are a number of synhospitality cases, where the parasite complex is monophyletic because evolved on a single host species. The special term--"phylogenetic synhospitality" (FS) is proposed for these cases of synhospitality. Most known cases of FS in acariform mites, permanent parasites of vertebrates, are analysed. It is found out that both astigmatan and prostigmatan parasite mites demonstrate a numbers of FS. The majority of these examples represent parasitism of two or three synhospitalic parasite species. Impressive examples of FS involving a number of synhospitalic species is shown by only astigmatan mites inhabiting the fur of mammals or plumage of birds. Most known examples involving four or more mite species are discussed: 51 mite species of the genus Schizocarpus (Chirodiscidae) parasitizing Castor fiber and C. canadensis (Castoridae); 6 species of Listrophorus spp. (Listrophoridae) from Ondatra zibethicus (Cricetidae); 23 species of Listrophoroides s. 1. (Atopomelidae) from Maxomys surifer (Muridae); 21 species of Cytostethum (Atomelidae) from Potorous tridactylus (Potoridae); 4 species of Listrophoroides (Afrolistrophoroides) from Malacomys longipes (Muridae); 7 species of Fainalges (Xolalgidae) from Aratinga holochlora (Psittacidae); 4 species of Zygepigynia (Pteronyssidae) from Chrysocolaptes lucidus (Picidae). The main reason of FS is that, in spite of the Fahrenholz's rule, the speciation of many parasites proceeds much more intensively than in their hosts because of the more rapid replacement of the parasitic generations. The first factor causing FS is the mite speciation it temporary segregated populations of the host (allopatric speciation). In this case, the "multispecies complexes" appeared after the subsequent reintegration of the host populations formerly isolated. The second factor is the speciation due to the specialization of mites to local microhabitats in the fur or plumage of host (sympatric or synxenic speciation). The second way of speciation is most characteristic for mites with highly specialized attaching structures. The phenomenon of FS more resides in ectoparasites of mammals rather than in feather mites in spite of much more structural complicacy of plumage rather than the fur. The high mobility of birds and wide dispersion of their new generations probably embarrass the process of sympatric speciation in their parasites. As a rule, only really significant geographical barriers play role for population isolation in birds. Thus, it could be concluded that two independent factors or their combination lead to FS. (i) The complex and/or disjunctive host range giving a possibility for allopatric speciation in parasites. (ii) The deep mite specialization to local microhabitats on the host body causing sympatric (synxenic) speciation. Fur of mammals and plumage of birds are very complicated in structure and microconditions and provide a considerable number of different microhabitats for mites inhabiting them. The prevalence of one of these two factors depends on the biological peculiarities of both parasites and their hosts. In mites with lesser specialized attaching organs, for example in atopomelids, allopatric speciation dominates. In mites with strongly specialized attaching organs, for example in listrophorids or chirodiscids, both pathways of speciation may take place. In feather mites, sympatric speciation should be more probable due to quite complicate and various structure of feathers in avian hosts. In fur mites, sympatric speciation is more likely in mites parasitizing hosts with peculiar ecology, for example in semiaquatic rodents possessing quite different fur structure in different parts of the body.     

Phylogeny and evolution of parasitism in feather mites of the families Epidermoptidae and Dermationidae (Acari: Analgoidea)

A phylogenetic reconstruction of feather mites of the epidermoptid complex (Analgoidea: Epidermoptidae, Dermationidae, and Knemidocoptidae) was carried out by methods of parsimony-based cladistics. The epidermoptid complex splits into two major branches, Epidermoptidae and Dermationidae. The family Dermationidae is monophyletic, while the Epidermoptidae, as previously defined, is paraphyletic. The family Knemidocoptidae is reduced to the subfamilial rank because it arises from the core of the Epidermoptidae. The subfamily Myialginae Trouessart, 1906 stat. resur. is restored within the Epidermoptidae. The aberrant genera Lukoschuscoptes and Apocnemidocoptes are moved from Knemidocoptidae to Epidermoptidae and Dermationidae, respectively. A hypothesis explaining main trends in morphological and ecological adaptations to parasitism on birds within the epidermoptid complex is proposed. New taxonomic diagnoses for higher taxa (families and subfamilies) are provided, and three new genera—Archemyialges gen. n., Hemimyialges gen. n. (Epidermoptidae), and Trochiloptes gen. n. (Dermationidae)—and a new subfamily—Apocnemidocoptinae—are established.     

Feather mites play a role in cleaning host feathers: New insights from DNA metabarcoding and microscopy

Parasites and other symbionts are crucial components of ecosystems, regulating host populations and supporting food webs. However, most symbiont systems, especially those involving commensals and mutualists, are relatively poorly understood. In this study, we have investigated the nature of the symbiotic relationship between birds and their most abundant and diverse ectosymbionts: the vane‐dwelling feather mites. For this purpose, we studied the diet of feather mites using two complementary methods. First, we used light microscopy to examine the gut contents of 1,300 individual feather mites representing 100 mite genera (18 families) from 190 bird species belonging to 72 families and 19 orders. Second, we used high‐throughput sequencing (HTS) and DNA metabarcoding to determine gut contents from 1,833 individual mites of 18 species inhabiting 18 bird species. Results showed fungi and potentially bacteria as the main food resources for feather mites (apart from potential bird uropygial gland oil). Diatoms and plant matter appeared as rare food resources for feather mites. Importantly, we did not find any evidence of feather mites feeding upon bird resources (e.g., blood, skin) other than potentially uropygial gland oil. In addition, we found a high prevalence of both keratinophilic and pathogenic fungal taxa in the feather mite species examined. Altogether, our results shed light on the long‐standing question of the nature of the relationship between birds and their vane‐dwelling feather mites, supporting previous evidence for a commensalistic–mutualistic role of feather mites, which are revealed as likely fungivore–microbivore–detritivore symbionts of bird feathers.     

Feather mites (Acari: Astigmata) and body condition of their avian hosts: a large correlative study

Feather mites are arthropods that live on or in the feathers of birds, and are among the commonest avian ectosymbionts. However, the nature of the ecological interaction between feather mites and birds remains unclear, some studies reporting negative effects of feather mites on their hosts and others reporting positive or no effects. Here we use a large dataset comprising 20 189 measurements taken from 83 species of birds collected during 22 yr in 151 localities from seven countries in Europe and North Africa to explore the correlation between feather mite abundance and body condition of their hosts. We predicted that, if wing‐dwelling feather mites are parasites, a negative correlation with host body condition should be found, while a mutualistic interaction should yield positive correlation. Although negative relationships between feather mite abundance and host body condition were found in a few species of birds, the sign of the correlation was positive in most bird species (69%). The overall effect size was only slightly positive (r =0.066). The effect of feather mite abundance explained <10% of variance in body condition in most species (87%). Results suggest that feather mites are not parasites of birds, but rather that they hold a commensalistic relationship where feather mites may benefit from feeding on uropygial gland secretions of their hosts and birds do not seem to obtain a great benefit from the presence of feather mites.     

Symbiotic feather mites synchronize dispersal and population growth with host sociality and migratory disposition

Some symbiotic taxa may have evolved to track changes in the level and quality of food resources provided by the host to increase reproduction and dispersal. As a consequence, some ectosymbionts synchronize their reproduction and activity with particular stages of their host's living cycle. In this article we examined temporal patterns of variation in prevalence and abundance of feather mites living on pre‐migratory barn swallows Hirundo rustica. Feather mites in the lineages Pterolichoidea and Analgoidea are the most common arthropod ectosymbionts living at the expenses of feather oil. We investigated whether the seasonal variations in levels of several measures of physiological condition associated with host migration were related to changes in prevalence and abundance of mites. The results suggest that the variation in prevalence of feather mites, and thus probably the mode of acquisition and dispersal of these symbionts, is linked to an increase in host sociality before migration. Physiological dynamics of hosts after the breeding season point at two clearly identifiable periods: a post‐breeding period when physiological condition remains stationary or decreases, and a pre‐migratory period characterized by a rapid increase in several measures of physiological condition. Mite population dynamics were synchronized with migratory disposition during the period of highest host gregariousness. These synchronized processes occurred in both study years, although dynamics of migratory disposition and mite prevalence and abundance differ somewhat between years for adult and juvenile hosts. Mite population increase before host migration may be a response to a higher quantity of food provided by the host, namely oil from the urpoygial gland which is stimulated by hormones. Therefore, mites might have evolved to adjust their reproduction to the time when they have more chance of dispersal through horizontal transmission. In addition, body mass of juvenile and adult hosts were positively related with mite abundance in both years after allowing for several influencing factors. Body mass variation may reflect adequately fitness of host or their current physiological state, for instance, differences in the secretion of lipids on feathers or a more adequate microclimate to these symbionts.     

Parasites and the blackcap's tail: implications for the evolution of feather ornaments

Although parasites may impair the expression of tail ornaments in birds, the importance of parasitism in driving the evolution of the initial stages of tail ornamentation is not well understood. Parasites could have negatively affected the expression of nonexaggerated, functional traits before these evolved ornaments, or they could have played a relevant role only after tails became ornamental and hence too costly to produce. To shed light on this issue, we studied the correlation between the abundance of feather mites (Acari, Proctophyllodidae) and the size, quality, growth rate and symmetry of tail feathers of blackcaps ( Sylvia atricapilla ), a non-ornamented passerine. Tail length was not correlated with mite load, yet blackcaps holding many mites at the moment of feather growth (fledglings) had lighter and more asymmetric feathers that grew at relatively lower rates. In blackcaps whose mite load was measured one year after feather growth (adults), only the negative correlation between mite intensity and feather symmetry remained significant. Changes in mite load since the moult season could have erased the correlation between condition-dependent feather traits and current parasite load in adults. Our results support the idea that different traits of non-ornamental feathers can signal parasite resistance. Therefore, parasitism could have played a central role in the evolution of tail ornamentation ever since its initial stages.  © 2002 The Linnean Society of London. Biological Journal of the Linnean Society , 2002, 76 , 481–492.     

The feather mite (Astigmata) fauna of some passerine birds (Passeriformes) in the south of Western Siberia

Feather mites (Astigmata) are specialized parasites living on the plumage and skin of birds. The paper presents data on infestation of some passerines (Passeriformes) by feather mites in the south of Western Siberia (Omsk and Tyumen Provinces). We found 24 species of feather mites belonging to the families Analgidae, Dermoglyphidae, Pteronyssidae, Trouessartiidae, and Proctophyllodidae on 16 bird species. Among them, 19 species are common parasites of the passerine birds examined; five species were detected on atypical hosts. Ten mite species were recorded for the first time on the passerine species examined. Analysis of the distribution of abundant and common mite species on their hosts has demonstrated that the majority of the bird parasites possess a specific distribution pattern in the host plumage with preference for certain feather types. We have also obtained new data on host associations of several mite species.     

Adaptive learning in arthropods: spider mites learn to distinguish food quality

Many herbivorous arthropods have been shown to possess learning capabilities, yet fitness effects of learning are largely unknown. In this paper, we test whether two-spotted spider mites (Tetranychus urticae) learn to distinguish food quality in choice tests, and whether this results in fitness benefits. Food consisted of cucumber plants with one of three degrees of feeding damage: undamaged (no mites), mildly damaged (infested by a mite strain adapted to tomato) and heavily damaged (infested by a mite strain adapted to cucumber). Mites were subjected to one choice test in a greenhouse and three sequential choice tests on leaf disks. Thereafter, individual mite performance was measured as oviposition rate over four days. In the course of the three small-scale choice tests, preference shifted towards less damaged food. The performance tests showed that learning was adaptive: mites learned to prefer the food type that yielded the higher oviposition rate. Interestingly, innate preferences in the greenhouse tests were close to those shown after learning in the small-scale tests. Given that both strains of mites had not experienced cucumber for several years, we hypothesize that the preference in the greenhouse was due to avoidance of mite odours rather than odours of damaged plants. Through its effect on foraging behaviour, adaptive learning may promote the evolution of host plant specialization in herbivorous arthropods.     

Comprehensive investigation of ectoparasite community and abundance across life history stages of avian host

Fitness consequences of ectoparasitism are expressed over the lifetime of their hosts in relation to variation in composition and abundance of the entire ectoparasite community and across all host life history stages. However, most empirical studies have focused on parasite species-specific effects and only during some life history stages. We conducted a systematic, year-long survey of an ectoparasite community in a wild population of house finches Carpodacus mexicanus Müller in south-western Arizona, with a specific focus on ecological and behavioral correlates of ectoparasite prevalence and abundance. We investigated five ectoparasite species: two feather mite genera – both novel for house finches – Strelkoviacarus (Analgidae) and Dermoglyphus (Dermoglyphidae), the nest mite Pellonyssus reedi (Macronyssidae), and the lice Menacanthus alaudae (Menoponidae) and Ricinus microcephalus (Ricinidae). Mite P. reedi and louse Menacanthus alaudae abundance peaked during host breeding season, especially in older birds, whereas feather mite abundance peaked during molt. Overall, breeding birds had more P. reedi than non-breeders, molting males had greater abundance of feather mites than molting females and non-molting males, and young males had more feather mites than older males. We discuss these results in relation to natural history of ectoparasites under study and suggest that ectoparasites might synchronize their life cycles to those of their hosts. Pronounced differences in relative abundance of ectoparasite species among host's life history stages have important implications for evolution of parasite-specific host defenses.     

Enabling large-scale feather mite studies: an Illumina DNA metabarcoding pipeline

Feather mites are among the most common and diverse ectosymbionts of birds, yet basic questions such as the nature of their relationship remain largely unanswered. One reason for feather mites being understudied is that their morphological identification is often virtually impossible when using female or young individuals. Even for adult male specimens this task is tedious and requires advanced taxonomic expertise, thus hampering large-scale studies. In addition, molecular-based methods are challenging because the low DNA amounts usually obtained from these tiny mites do not reach the levels required for high-throughput sequencing. This work aims to overcome these issues by using a DNA metabarcoding approach to accurately identify and quantify the feather mite species present in a sample. DNA metabarcoding is a widely used molecular technique that takes advantage of high-throughput sequencing methodologies to assign the taxonomic identity to all the organisms present in a complex sample (i.e., a sample made up of multiple specimens that are hard or impossible to individualise). We present a high-throughput method for feather mite identification using a fragment of the COI gene as marker and Illumina Miseq technology. We tested this method by performing two experiments plus a field test over a total of 11,861 individual mites (5360 of which were also morphologically identified). In the first experiment, we tested the probability of detecting a single feather mite in a heterogeneous pool of non-conspecific individuals. In the second experiment, we made 2?×?2 combinations of species and studied the relationship between the proportion of individuals of a given species in a sample and the proportion of sequences retrieved to test whether DNA metabarcoding can reliably quantify the relative abundance of mites in a sample. Here we also tested the efficacy of degenerate primers (i.e., a mixture of similar primers that differ in one or several bases that are designed to increase the chance of annealing) and investigated the relationship between the number of mismatches and PCR success. Finally, we applied our DNA metabarcoding pipeline to a total of 6501 unidentified and unsorted feather mite individuals sampled from 380 European passerine birds belonging to 10 bird species (field test). Our results show that this proposed pipeline is suitable for correct identification and quantitative estimation of the relative abundance of feather mite species in complex samples, especially when dealing with a moderate number (>?30) of individuals per sample.     

Feather mites (Acarina) of the parakeet, Melopsittacus undulatus (Shaw) (Aves: Psittacidae)

The parakeet (or budgerigar) has been transported to many regions of the world. Two species of feather mites, Protolichus lunula (Robin) (Pterolichoidea: Pterolichidae) and Dubininia melopsittaci n. sp. (Analgoidea: Xolalgidae), are specific to this host; from distribution records, these ectoparasites probably have been distributed worldwide with their hosts.     

Adaptive learning in arthropods: spider mites learn to distinguish food quality

Many herbivorous arthropods have been shown to possess learning capabilities, yet fitness effects of learning are largely unknown. In this paper, we test whether two-spotted spider mites (Tetranychus urticae) learn to distinguish food quality in choice tests, and whether this results in fitness benefits. Food consisted of cucumber plants with one of three degrees of feeding damage: undamaged (no mites), mildly damaged (infested by a mite strain adapted to tomato) and heavily damaged (infested by a mite strain adapted to cucumber). Mites were subjected to one choice test in a greenhouse and three sequential choice tests on leaf disks. Thereafter, individual mite performance was measured as oviposition rate over four days. In the course of the three small-scale choice tests, preference shifted towards less damaged food. The performance tests showed that learning was adaptive: mites learned to prefer the food type that yielded the higher oviposition rate. Interestingly, innate preferences in the greenhouse tests were close to those shown after learning in the small-scale tests. Given that both strains of mites had not experienced cucumber for several years, we hypothesize that the preference in the greenhouse was due to avoidance of mite odours rather than odours of damaged plants. Through its effect on foraging behaviour, adaptive learning may promote the evolution of host plant specialization in herbivorous arthropods. This revised version was published online in July 2006 with corrections to the Cover Date.     

Climate-Driven Variation in the Intensity of a Host-Symbiont Animal Interaction along a Broad Elevation Gradient

Gradients of environmental stress may affect biotic interactions in unpredictable ways responding to climate variation, depending on the abiotic stress tolerance of interacting partners. Here, we study the effect of local climate on the intensity of feather mites in six mountain passerines along a 1400 m elevational gradient characterized by shifting temperature and rainfall. Although obligatory symbionts of warm-blooded organisms are assumed to live in mild and homeothermic environments, those inhabiting external, non-blood-irrigated body portions of the host organism, such as feather mites, are expected to endure exposure to the direct influence of a fluctuating climate. As expected, feather mite intensity declined with elevation in all bird species, a pattern that was also found in cold-adapted passerines that have typical alpine habits. The elevation cline was mainly explained by a positive effect of the average temperature upon mite intensity in five of the six species studied. Precipitation explained less variance in mite intensity than average temperature, and showed a negative correlation in half of the studied species. We found no climate-driven migration of mites along the wings of birds, no replacement of mite species along elevation gradients and no association with available food resources for mites (estimated by the size of the uropygial gland). This study suggests that ectosymbionts of warm-blooded animals may be highly sensitive to climatic variation and become less abundant under stressful environmental conditions, providing empirical evidence of the decline of specialized biotic interactions among animal species at high elevations.