Host space, not energy or symbiont size, constrains feather mite abundance across passerine bird species.

Comprehending symbiont abundance among host species is a major ecological endeavour, and the metabolic theory of ecology has been proposed to understand what constrains symbiont populations. We parameterized metabolic theory equations to investigate how bird species' body size and the body size of their feather mites relate to mite abundance according to four potential energy (uropygial gland size) and space constraints (wing area, total length of barbs and number of feather barbs). Predictions were compared with the empirical scaling of feather mite abundance across 106 passerine bird species (26,604 individual birds sampled), using phylogenetic modelling and quantile regression. Feather mite abundance was strongly constrained by host space (number of feather barbs) but not by energy. Moreover, feather mite species' body size was unrelated to the body size of their host species. We discuss the implications of our results for our understanding of the bird-feather mite system and for symbiont abundance in general.

Cryptic carnivores: Intercontinental sampling reveals extensive novel diversity in a genus of freshwater annelids.

Freshwater annelids are globally widespread in aquatic ecosystems, but their diversity is severely underestimated. Obvious morphological features to define taxa are sparse, and molecular phylogenetic analyses regularly discover cryptic diversity within taxa. Despite considerable phylogenetic work on certain clades, many groups of freshwater annelids remain poorly understood. Included among these are water nymph worms of the genus Chaetogaster (Clitellata: Tubificida: Naididae: Naidinae). These worms have diverged from the detritivorous diet of most oligochaetes to become more predatory and exist as omnivores, generalist predators, parasites, or symbionts on other invertebrates. Despite their unusual trophic ecology, the true diversity of Chaetogaster and the phylogenetic relationships within the genus are uncertain. Only three species are commonly referenced in the literature (Chaetogaster diaphanus, Chaetogaster limnaei, and Chaetogaster diastrophus), but additional species have been described and prior molecular data suggests that there is cryptic diversity within named species. To clarify the phylogenetic diversity of Chaetogaster, we generated the first molecular phylogeny of the genus using mitochondrial and nuclear sequence data from 128 worms collected primarily across North America and Europe. Our phylogenetic analyses suggest that the three commonly referenced species are a complex of 24 mostly cryptic species. In our dataset, Chaetogaster "diaphanus" is represented by two species, C. "limnaei" is represented by three species, and C. "diastrophus" is represented by 19 species. North American and European sequences are largely interspersed across the phylogeny, with four pairs of clades involving distinct North American and European sister groupings. Overall, our study demonstrates that the species diversity of Chaetogaster has been underestimated and that carnivory has evolved at least twice in the genus. Chaetogaster is being used as a model for symbiotic evolution and the loss of regenerative ability, and our study indicates that researchers must be careful to identify which species of Chaetogaster they are working with in future studies.

New feather mites of the genus Neodectes (Acariformes: Proctophyllodidae) from honeyeaters (Passeriformes: Meliphagidae) in Australia.

Five new species of the genus Neodectes Park and Atyeo, 1971 (Proctophyllodidae: Pterodectinae) are described from honeyeaters (Passeriformes: Meliphagidae) in Australia: Neodectes cissomelae sp. n. from Cissomela pectoralis (Gould) (type host) and Melithreptus gularis laetior Gould; N. hallidayi sp. n. from Anthochaera carunculata (Shaw) (type host) and A. chrysoptera (Latham); N. manorinae sp. n. from Manorina melanocephala (Latham); N. ophioglossus sp. n. from Conopophila rufogularis (Gould) (type host) and Lichmera indistincta (Vigors and Horsfield); and N. walteri sp. n. from Anthochaera phrygia (Shaw). A key to species and a world checklist to Neodectes species are provided for the first time. A new combination, Neodectes dicranochaetus (Gaud, 1968) comb. n., is proposed for Proterothrix dicranochaeta Gaud, 1968, which is transferred herein to the genus Neodectes from Proterothrix Gaud, 1968 (Proctophyllodidae).

Application of DNA barcoding and morphometric analysis in differentiating cystacanths of Polymorphus species (Acanthocephala: Polymorphidae) from central Alberta, Canada.

Acanthocephalans are multi-host endoparasites, many of which use freshwater amphipods as intermediate hosts for their larval stages (e.g., cystacanths) while adults live in the intestines of vertebrates, including waterfowl. In central Alberta, Canada, several co-occurring species of the acanthocephalan genus Polymorphus use the amphipod Gammarus lacustris Sars, 1863 as an intermediate host. We applied DNA barcoding and morphometric analysis to differentiate cystacanth larvae from G. lacustris sampled from 17 Albertan water bodies. We slide-mounted specimens and measured morphological traits relating to proboscis hooks. We sequenced the standard DNA barcoding region of the mitochondrial cytochrome c oxidase subunit I gene (COI). Morphometric analysis suggested that the acanthocephalans we collected belonged to four morphologically different groups that keyed to Polymorphus contortus (Bremser, 1821) Travassos, 1926; P. marilis Van Cleave, 1939; P. paradoxus Connel et Corner, 1957; and P. strumosoides (Lundström, 1942) Amin, 2013. Our Bayesian tree based on COI sequences generally corroborated the morphological results and supported that the specimens assigned to P. cf. contortus and P. cf. strumosoides belong to two distinct species. In contrast, the Bayesian tree showed that specimens of P. cf. marilis were nested as a cluster within the P. cf. paradoxus clade. Similarly, small pairwise genetic distance (< 2%) between specimens identified as P. cf. contortus and P. cf. strumosoides suggests that they are conspecific. Future studies should use morphology and sequence data from adult acanthocephalans to assess the taxonomic identity of the cystacanth-based Polymorphus taxa. Our study is the first to provide genetic information for the four Polymorphus taxa and emphasizes the importance of applying multiple approaches to differentiate parasite species.

Parasite prevalence in intermediate hosts increases with waterbody age and abundance of final hosts.

Prevalence of a parasite may be influenced by age of the habitat (= time available for hosts and parasites to colonize habitats), assemblage composition of host and non-host species, or biotic and abiotic habitat features. For a trophically transmitted parasite, the intermediate host may be consumed by both final hosts and 'dead-end' predators that are incompetent to host the parasite. We tested biotic and abiotic factors that might influence parasite prevalence in a freshwater host-parasite system using a dataset collected from 36 water bodies in the vicinity of Edmonton, Alberta, Canada. In this system, eggs of thorny-headed worms (Acanthocephala: Polymorphus spp.) are eaten by intermediate-host amphipods (Gammarus lacustris Sars), which are then consumed by final vertebrate hosts (certain aquatic birds and muskrats) and various non-host waterbird species. We found that acanthocephalan prevalence in amphipods was positively correlated with waterbody age and with abundance of final-host species. In contrast, abundance of the intermediate host G. lacustris was less important and was negatively correlated with parasite prevalence ('encounter-dilution effect'). Similarly, parasite prevalence showed a marginally significant and negative correlation with abundance of 'dead-end' Gammarus-eating birds. We conclude that in our study system, time available for colonization and abundance of final hosts are more important for parasite prevalence in intermediate hosts than is abundance of intermediate and dead-end hosts.

The Distribution of Quill Mites (Betasyringophiloidus seiuri) Among Flight Feathers of the Ovenbird (Seiurus aurocapilla).

Quill mites of the family Syringophilidae (Acariformes: Prostigmata) are permanent ectoparasites that live inside the hollow quills of feathers and use their long chelicerae to pierce the quill wall to feed on living tissue of the quill follicle. Ovenbirds (Parulidae: Seiurus aurocapilla (Linnaeus)) are host to the quill mite Betasyringophiloidus seiuri (Clark), which has been previously reported to infest only 5.1% of this host's tail feathers; however, this species has also been reported to inhabit the wing feathers of ovenbirds, but without any data on prevalence or intensity. We examined all 1,008 flight feathers from 21 dead ovenbirds from Canada for quill mites and calculated infestation parameters per feather location. Nine of the 21 birds were infested with mites (prevalence of 42.9%). Feathers with the highest prevalence were Primaries 1 and 2 and Secondaries 1, 2, and 5. Only 2 of the 9 infested birds had mites in their tail feathers; in each case, only a single feather was infested. Mean intensity was 28.4 mites/feather with a range of 2-135.6, and ∼88% of the adult mites were female. To determine if quill volume and quill wall thickness correlated with mite presence and abundance, we took quill measurements from the flight feathers of 3 additional ovenbirds. There was a strong positive correlation between quill volume and mean mite intensity. Feathers that had quill walls thicker in some areas than the mites' extended chelicerae had lower prevalences than did feathers with walls consistently thinner than the length of the chelicerae. We conclude that B. seiuri is much more likely to be found in wing than in tail feathers and that it has greater reproductive success in quills with both large volumes and thin walls; however, whether foundress mites preferentially choose to colonize these feathers requires more study.

Life history of Macrocheles muscaedomesticae (Parasitiformes: Macrochelidae): new insights on life history and evidence of facultative parasitism on Drosophila.

Macrocheles muscaedomesticae is a cosmopolitan macrochelid mite whose populations have likely diverged considering the many locations they inhabit, but most of the work published on this mite species has been on the basis of their association with the house fly, Musca domestica. Here, we studied several aspects of the biology of M. muscaedomesticae associated with drosophilid flies collected in Alberta, Canada. We assessed the degree of divergence of our populations from others, compared their life history to other published populations and experimentally tested whether M. muscaedomesticae feeds on Drosophila hydei hosts by comparing the body mass of mites that attached to hosts to those that did not. There was no strong phylogenetic differentiation among any of the M. muscaedomesticae specimens, suggesting multiple recent introductions of this species to Canada. Compared to other populations, our mites exhibited lower fecundity, which may have been a result of the temperature or nematode-only diet in which they were maintained. Finally, mites that attached to hosts for 4 h weighed significantly more than those that did not. Without direct evidence for host tissue transfer to the mites, it is difficult to determine whether the mites are indeed feeding on their hosts while attached. However, the existing evidence for the costs fly hosts endure at the expense of these mites makes this relationship antagonistic.

DNA barcodes expose unexpected diversity in Canadian mites.

Mites (Arachnida: Acariformes, Parasitiformes) are the most abundant and species-rich group of arthropods in soil, but are also diverse in freshwater habitats, on plants, and as symbionts of larger animals. However, assessment of their diversity has been impeded by their small size and often cryptic morphology. As a consequence, published estimates of their species richness span more than two orders of magnitude (0.4-114 million). In this study we employ DNA barcoding and the Barcode Index Number (BIN) system to investigate mite diversity at over 1,800 sites across Canada, primarily from soil and litter habitats with smaller contributions from freshwater, plants, and animal hosts. Barcodes from 73,394 specimens revealed 7,077 BINs with representatives from all four orders (Ixodida, Mesostigmata, Sarcoptiformes, Trombidiformes) and 60% (186) of the known families. The BIN total is 2.4 times the number of species previously recorded from Canada (2,999), reflecting the unexpectedly high richness of several families. Richness projections suggest that more than 28,000 BINs occur at the sampled locations, indicating that the Canadian mite fauna almost certainly includes more than 30,000 species-a total similar to that for the most diverse insect order in Canada, Diptera. This unexpected diversity was partitioned into highly dissimilar, spatially-structured assemblages that likely reflect dispersal limitation and environmental heterogeneity. Further sampling of a greater diversity of habitats will refine understanding of mite diversity in Canada, but similar analyses in other geographic regions will be essential to ascertain their diversity at a global scale.

Acari of Canada.

Summaries of taxonomic knowledge are provided for all acarine groups in Canada, accompanied by references to relevant publications, changes in classification at the family level since 1979, and notes on biology relevant to estimating their diversity. Nearly 3000 described species from 269 families are recorded in the country, representing a 56% increase from the 1917 species reported by Lindquist et al. (1979). An additional 42 families are known from Canada only from material identified to family- or genus-level. Of the total 311 families known in Canada, 69 are newly recorded since 1979, excluding apparent new records due solely to classification changes. This substantial progress is most evident in Oribatida and Hydrachnidia, for which many regional checklists and family-level revisions have been published. Except for recent taxonomic leaps in a few other groups, particularly of symbiotic mites (Astigmata: feather mites; Mesostigmata: Rhinonyssidae), knowledge remains limited for most other taxa, for which most species records are unpublished and may require verification. Taxonomic revisions are greatly needed for a large majority of families in Canada. Based in part on species recorded in adjacent areas of the USA and on hosts known to be present here, we conservatively estimate that nearly 10,000 species of mites occur in Canada, but the actual number could be 15,000 or more. This means that at least 70% of Canada's mite fauna is yet unrecorded. Much work also remains to match existing molecular data with species names, as less than 10% of the ~7500 Barcode Index Numbers for Canadian mites in the Barcode of Life Database are associated with named species. Understudied hosts and terrestrial and aquatic habitats require investigation across Canada to uncover new species and to clarify geographic and ecological distributions of known species.

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.

Nasal mites (Mesostigmata: Rhinonyssidae) in African penguins (Spheniscus demersus).

Rhinonyssids are obligate haematophagous mites that parasitize the nasal cavity of vertebrates, and occur in a wide range of birds worldwide. Two species of nasal mites are known to occur in penguins: Rhinonyssus sphenisci, which has been recorded from Humboldt and Magellanic penguins (Spheniscus humboldti and S. magellanicus, respectively), and Rhinonyssus schelli, which has been recorded in Adélie and Gentoo penguins (Pygoscelis adeliae and P. papua, respectively). We examined the nasal cavity of African penguins (Spheniscus demersus) that died while under care at a rehabilitation centre (N = 40) or that were found dead at breeding colonies (N = 67). Nasal mites were found in the nasal cavity and/or paranasal of sinuses of 21 penguins, some of which had signs of mild-to-moderate sinusitis. Prevalence was higher in juveniles (29%) and adults (27%) than in chicks (10%). Mean intensity was 5.9 ± 12.9 mites per infected host (range 1-60). The mites presented morphological characteristics that were at times consistent with either R. sphenisci or R. schelli, and therefore we conservatively classified them as 'R. sphenisci sensu lato'. Our morphometric results raise the question of whether the specific status of R. schelli is justified.

Hyperparasitism of an Avian Ectoparasitic Hippoboscid Fly, Ornithomya anchineuria, by the Mite, Myialges Cf. Borealis, in Alberta, Canada.

Hippoboscid flies (Diptera: Hippoboscidae) include species that are ectoparasites of birds in the Northern Hemisphere, but little is known regarding their taxonomy, parasites, avian host associations, or geographical distribution in North America. In late August of 2013 and 2014, we collected hippoboscid flies from live birds trapped in mist nets as part of a banding study in Cypress Hills Interprovincial Park in southeastern Alberta, Canada. A total of 113 birds comprising 9 species was examined in 2013. Of these, 18 individuals were infested with 1-3 Ornithomya anchineuria Speiser (n = 22 flies; prevalence = 15.9%). Eight of these flies carried 1-8 adult female epidermoptid mites anchored to their ventral, posterior abdomens. Each female was associated with clusters of up to 30 stalked eggs. The first pair of tarsi on adult female mites was highly modified as anchors, indicating permanent attachment through the host cuticle. Morphological traits identified these mites as Myialges cf. borealis Mironov, Skirnisson, Thorarinsdottier and Nielsen. Cytochrome c oxidase subunit 1 ( COX1) gene sequences obtained for 2 mites were distinct from those previously reported for species of Myialges, being most similar to Myialges trinotoni Cooreman. The paucity of available gene sequences for Myialges and related genera of epidermoptid mites prevents any further conclusions regarding taxonomy. These findings extend previous reports of O. anchineuria from Pacific and Atlantic coasts of Canada inland to the central migratory flyway of the Northern Great Plains and expand the limited information available for Myialges spp.

High diversity and low genetic structure of feather mites associated with a phenotypically variable bird host.

Obligate symbionts may be genetically structured among host individuals and among phenotypically distinct host populations. Such processes may in turn determine within-host genetic diversity of symbionts, which is relevant for understanding symbiont population dynamics. We analysed the population genetic structure of two species of feather mites (Proctophyllodes sylviae and Trouessartia bifurcata) in migratory and resident blackcaps Sylvia atricapilla that winter sympatrically. Resident and migratory hosts may provide mites with habitats of different qualities, what might promote specialization of mite populations. We found high genetic diversity of within-host populations for both mite species, but no sign of genetic structure of mites between migratory and resident hosts. Our results suggest that, although dispersal mechanisms between hosts during the non-breeding season are unclear, mite populations are not limited by transmission bottlenecks that would reduce genetic diversity among individuals that share a host. Additionally, there is no evidence that host phenotypic divergence (associated with the evolution of migration and residency) has promoted the evolution of host-specialist mite populations. Unrestricted dispersal among host types may allow symbiotic organisms to avoid inbreeding and to persist in the face of habitat heterogeneity in phenotypically diverse host populations.

Parasite biodiversity faces extinction and redistribution in a changing climate.

Climate change is a well-documented driver of both wildlife extinction and disease emergence, but the negative impacts of climate change on parasite diversity are undocumented. We compiled the most comprehensive spatially explicit data set available for parasites, projected range shifts in a changing climate, and estimated extinction rates for eight major parasite clades. On the basis of 53,133 occurrences capturing the geographic ranges of 457 parasite species, conservative model projections suggest that 5 to 10% of these species are committed to extinction by 2070 from climate-driven habitat loss alone. We find no evidence that parasites with zoonotic potential have a significantly higher potential to gain range in a changing climate, but we do find that ectoparasites (especially ticks) fare disproportionately worse than endoparasites. Accounting for host-driven coextinctions, models predict that up to 30% of parasitic worms are committed to extinction, driven by a combination of direct and indirect pressures. Despite high local extinction rates, parasite richness could still increase by an order of magnitude in some places, because species successfully tracking climate change invade temperate ecosystems and replace native species with unpredictable ecological consequences.

Phylogeography of a migratory songbird across its Canadian breeding range: Implications for conservation units.

The objectives of this study were to describe and evaluate potential drivers of genetic structure in Canadian breeding populations of the Ovenbird, Seiurus aurocapilla. We performed genetic analyses on feather samples of individuals from six study sites using nuclear microsatellites. We also assessed species identity and population genetic structure of quill mites (Acariformes, Syringophilidae). For male Ovenbirds breeding in three study sites, we collected light-level geolocator data to document migratory paths and identify the wintering grounds. We also generated paleohindcast projections from bioclimatic models of Ovenbird distribution to identify potential refugia during the last glacial maximum (LGM, 21,000 years before present) as a factor explaining population genetic structure. Birds breeding in the Cypress Hills (Alberta/Saskatchewan) may be considered a distinct genetic unit, but there was no evidence for genetic differentiation among any other populations. We found relatively strong migratory connectivity in both western and eastern populations, but some evidence of mixing among populations on the wintering grounds. There was also little genetic variation among syringophilid mites from the different Ovenbird populations. These results are consistent with paleohindcast distribution predictions derived from two different global climate models indicating a continuous single LGM refugium, with the possibility of two refugia. Our results suggest that Ovenbird populations breeding in boreal and hemiboreal regions are panmictic, whereas the population breeding in Cypress Hills should be considered a distinct management unit.

Behavioural responses to potential dispersal cues in two economically important species of cereal-feeding eriophyid mites.

Passively dispersing organisms should optimise the time and direction of dispersal by employing behaviours that increase their probability of being successfully transported by dispersal agents. We rigorously tested whether two agriculturally important passively-dispersing eriophyoid species, wheat curl mite (WCM) and cereal rust mite (CRM), display behaviours indicating their readiness to depart from current host plants in the presence of potential dispersal cues: wind, an insect vector and presence of a fresh plant. Contrary to our expectations, we found that both species decreased their general activity in the presence of wind. When exposed to wind, WCM (but not CRM) significantly increased behaviour that has previously been considered to facilitate dispersal (in this case, standing vertically). Our study provides the first sound test of the function of what have been interpreted as dispersal-related behaviours of eriophyid mites. The low proportion of WCM exhibiting dispersal behaviour suggests there may be predisposed dispersers and residents in the population. Moreover, we found that WCM was generally more active than CRM, which is likely a contributing factor to its high invasive potential.

Global associations between birds and vane-dwelling feather mites.

Understanding host-symbiont networks is a major question in evolutionary ecology. Birds host a great diversity of endo- and ectosymbiotic organisms, with feather mites (Arachnida: Acariformes: Analgoidea, Pterolichoidea) being among the most diverse of avian symbionts. A global approach to the ecology and evolution of bird-feather-mite associations has been hampered because of the absence of a centralized data repository. Here we present the most extensive data set of associations between feather mites and birds. Data include 12 036 records of 1887 feather mite species located on the flight feathers of 2234 bird species from 147 countries. Feather mites typically located inside quills, on the skin, or on downy body feathers are not included. Data were extracted from 493 published sources dating from 1882 to 2015. Data exploration shows that although most continents and bird families are represented, most bird species remain unexplored for feather mites. Nevertheless, this is the most comprehensive data set available for enabling global macroecological analyses of feather mites and their hosts, such as ecological network analyses. This metadata file outlines the structure of these data and provides primary references for all records used.

Higher-level molecular phylogeny of the water mites (Acariformes: Prostigmata: Parasitengonina: Hydrachnidiae).

With nearly 6000 named species, water mites (Hydrachnidiae) represent the largest group of arachnids to have invaded and extensively diversified in freshwater habitats. Water mites together with three other lineages (the terrestrial Erythraiae and Trombidiae, and aquatic Stygothrombiae), make up the hyporder Parasitengonina, which is characterized by having parasitic larvae and predatory nymphs and adults. Relationships between the Hydrachnidiae and other members of the Parasitengonina are unclear, as are relationships among the major lineages of water mites. Monophyly of water mites has been asserted, with the possible exception of the morphologically distinctive Hydrovolzioidea. Here we infer the phylogeny of water mites using multiple molecular markers and including representatives of all superfamilies of Hydrachnidiae and of almost all other Parasitengonina. Our results support a monophyletic Parasitengonina including Trombidiae, Stygothrombiae, and Hydrachnidiae. A monophyletic Hydrachnidiae, including Hydrovolzioidea, is strongly supported. Terrestrial Parasitengonina do not form a monophyletic sister group to water mites. Stygothrombiae is close to water mites but is not nested within this clade. Water mites appear to be derived from ancestors close to Stygothrombiae or the erythraoid group Calyptostomatoidea; however, this relationship is not clear because of extremely short branches in this part of the parasitengonine tree. We recovered with strong support all commonly accepted water mite superfamilies except for Hydryphantoidea, which is clearly paraphyletic. Our data support the previously proposed clades Protohydrachnidia (Hydrovolzioidea and Eylaoidea), Euhydrachnidia (all remaining superfamilies), and the euhydrachnid subclade Neohydrachnidia (Lebertioidea, Hydrachnoidea, Hygrobatoidea, and Arrenuroidea). We found that larval leg structure and locomotory behavior are strongly congruent with the molecular phylogeny. Other morphological and behavioral characters, including host choice, are not as strongly correlated with phylogeny. Molecular dating suggests that the Hydrachnidiae arose about 235MYA, and that Neohydrachnidia began to diversify about 155MYA. Our results provide a strong framework for classification and for further elaboration at finer taxonomic scales, which will allow testing of ecological and behavioral hypotheses associated with the transition from terrestrial to aquatic life.

Niche Partitioning of Feather Mites within a Seabird Host, Calonectris borealis.

According to classic niche theory, species can coexist in heterogeneous environments by reducing interspecific competition via niche partitioning, e.g. trophic or spatial partitioning. However, support for the role of competition on niche partitioning remains controversial. Here, we tested for spatial and trophic partitioning in feather mites, a diverse and abundant group of arthropods. We focused on the two dominant mite species, Microspalax brevipes and Zachvatkinia ovata, inhabiting flight feathers of the Cory's shearwater, Calonectris borealis. We performed mite counts across and within primary and tail feathers on free-living shearwaters breeding on an oceanic island (Gran Canaria, Canary Islands). We then investigated trophic relationships between the two mite species and the host using stable isotope analyses of carbon and nitrogen on mite tissues and potential host food sources. The distribution of the two mite species showed clear spatial segregation among feathers; M. brevipes showed high preference for the central wing primary feathers, whereas Z. ovata was restricted to the two outermost primaries. Morphological differences between M. brevipes and Z. ovata support an adaptive basis for the spatial segregation of the two mite species. However, the two mites overlap in some central primaries and statistical modeling showed that Z. ovata tends to outcompete M. brevipes. Isotopic analyses indicated similar isotopic values for the two mite species and a strong correlation in carbon signatures between mites inhabiting the same individual host suggesting that diet is mainly based on shared host-associated resources. Among the four candidate tissues examined (blood, feather remains, skin remains and preen gland oil), we conclude that the diet is most likely dominated by preen gland oil, while the contribution of exogenous material to mite diets is less marked. Our results indicate that ongoing competition for space and resources plays a central role in structuring feather mite communities. They also illustrate that symbiotic infracommunities are excellent model systems to study trophic ecology, and can improve our understanding of mechanisms of niche differentiation and species coexistence.

Different space preferences and within-host competition promote niche partitioning between symbiotic feather mite species.

Obligate symbionts (including parasites, commensals and mutualists) often share host species and host-based food resources. Such symbionts are frequently distributed unequally among hosts with different phenotypic features, or occupy different regions on a host. However, the processes leading to distinct within-host symbiont distributions remain obscure. We aimed to test whether distinct in-host symbiont distributions arise as the outcome of species-specific habitat preferences or interspecific competition, and how host phenotype influences such processes. To this end, we studied the distribution within and among individual bird hosts of two feather mites (Proctophyllodes sylviae and Trouessartia bifurcata) of migratory and sedentary European blackcaps, Sylvia atricapilla, wintering in sympatry. Trouessartia bifurcata was mostly restricted to resident blackcaps, while P. sylviae was abundant on both host types. Within hosts, each species tended to settle on different feather sectors (proximal or distal, respectively), which they filled by spreading on the wing following ordered but opposite patterns, thereby supporting the view that spatial segregation was primarily the outcome of dissimilar space preferences. However, we also found evidence of competition finely tuning mite distributions: when P. sylviae increased abundance and expanded onto the range of T. bifurcata, abundances of the two species were negatively correlated in the shared areas. In addition, the presence of T. bifurcata on a host was associated with a more restricted distribution of P. sylviae. Our results show that both species-specific preferences and interspecific interactions contribute to shaping mite distributions among and on individual hosts, a situation likely mirrored by other host-multi-symbiont systems.