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.

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.

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.

Molecular phylogeny of North American Branchiobdellida (Annelida: Clitellata).

Branchiobdellidans, or crayfish worms, are ectosymbiotic clitellate annelids associated primarily with freshwater crayfishes. The main objectives of our study were to infer a molecular phylogeny for the North American Branchiobdellida, examine its congruence with morphology-based hypotheses of relationships at the subfamily and genus level, and use our dataset to assess consistency of GenBank-archived branchiobdellidan sequences. We used nucleotide sequence data from two mtDNA genes (COI and 16S rDNA) and three nuclear genes (28S rDNA, 18S rDNA, and ITS1) to estimate phylogenetic relationships among 47 described and one undescribed species of Branchiobdellida. We recovered a monophyletic branchiobdellidan clade with generally short branch lengths, suggesting that a large portion of the taxon has likely undergone a recent and rapid radiation in North America. Results from our phylogenetic analyses indicate that current taxonomic groupings are largely unsupported by the molecular data. All four subfamilies are either paraphyletic or polyphyletic, and only three of seven sampled non-monotypic genera were monophyletic. We found a high rate (49%) of inconsistency in GenBank-archived sequences, over 70% of which can be attributed to field- or laboratory-based error.

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).

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.

Can freshwater mites act as forensic tools?

Determination of post-mortem interval often employs analysis of age structure and diversity of saprophilic arthropods (including mites) that have colonized corpses. The majority of research has focused on decomposition processes in terrestrial situations, with relatively few studies on the utility of freshwater invertebrates as forensic agents. Most freshwater mites are predators, detritivores or algivores, and hence seem unlikely candidates as tools for aging or determining original placement of corpses or other bodily remains. The main exceptions to this are some aquatic Astigmata, which have occasionally been observed feeding on the tissues of moribund aquatic animals. Here I investigate Canadian law literature and published forensic research to determine how frequently freshwater mites are included in court cases or are found attending dead bodies. I found only one questionable report of aquatic mites in over 30 years of material from legal databases. Three published research papers reported mites associated with vertebrate flesh in fresh water. Only one paper provided an identification of mites finer than 'Acari' or 'water mites'. In this case, the mites were identified as Hydrozetes (Oribatida). In none of these papers were mites reported to be high in abundance or biomass, and in two of the three publications methodological problems and/or poor reporting of data raised doubts about interpretation of results. I conclude that based on their biology, there is little expectation that freshwater mites should be of great value as forensic tools, and this survey of legal and scientific literature supports my argument.

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.

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.

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.

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.

Geographical structuring of feather mite assemblages from the Australian brush-turkey (Aves: Megapodiidae).

Populations of a host species may exhibit different assemblages of parasites and other symbionts. The loss of certain species of symbionts (lineage sorting, or "missing-the-boat") is a mechanism by which geographical variation in symbiont assemblages can arise. We studied feather mites and lice from Australian brush-turkeys (Aves: Megapodiidae: Alectura lathami) and expected to observe geographical structuring in arthropod assemblages for several reasons. First, because the brush-turkey is a sedentary ground-dwelling bird, we predicted that geographically close host populations should share more similar arthropod assemblages than distant ones. Second, because brush-turkeys do not brood their young, vertical transfer of arthropods is unlikely, and brush-turkeys probably acquire their mites and lice at social maturity through contact with other birds. Young birds could disperse and found new populations without carrying complete sets of symbionts. We predicted that young birds would have fewer species of arthropods than older birds; in addition, we expected that males (which are polygynous) would have more species than females. Birds were sampled from 12 sites (=populations) along the east coast of Queensland, Australia, that were separated by a distance of 12.5-2,005 km. In total, 5 species of mites from the Pterolichidae and 1 species from the Ascouracaridae were found. Two species of lice were collected but in numbers too low to be statistically useful. Differentiation of mite assemblages was evident; in particular, Leipobius sp. showed 100% prevalence in 3 host populations and 0% in the remaining 9. A dendrogram of brush-turkey populations based on mite assemblages showed 2 geographically correlated clusters of sites, plus 1 cluster that contained 2 sites near Brisbane and 1 approximately at a distance of 1,000 km. There was no strong effect of host age or sex on number of mite species carried. Horizontal transfer of feather mites by hippoboscid flies, in addition to physical contact between hosts, may play a role in homogenizing symbiont assemblages within populations.

Effects of an ecosystem engineer on belowground movement of microarthropods.

Ecosystem engineers affect other species by changing physical environments. Such changes may influence movement of organisms, particularly belowground where soil permeability can restrict dispersal. We investigated whether earthworms, iconic ecosystem engineers, influence microarthropod movement. Our experiment tested whether movement is affected by tunnels (i.e., burrows), earthworm excreta (mucus, castings), or earthworms themselves. Earthworm burrows form tunnel networks that may facilitate movement. This effect may be enhanced by excreta, which could provide resources for microarthropods moving along the network. Earthworms may also promote movement via phoresy. Conversely, negative effects could occur if earthworms alter predator-prey relationships or change competitive interactions between microarthropods. We used microcosms consisting of a box connecting a "source" container in which microarthropods were present and a "destination" container filled with autoclaved soil. Treatments were set up within the boxes, which also contained autoclaved soil, as follows: 1) control with no burrows; 2) artificial burrows with no excreta; 3) abandoned burrows with excreta but no earthworms; and 4) earthworms (Lumbricus rubellus) present in burrows. Half of the replicates were sampled once after eight days, while the other half were sampled repeatedly to examine movement over time. Rather than performing classical pairwise comparisons to test our hypotheses, we used AIC(c) to assess support for three competing models (presence of tunnels, excreta, and earthworms). More individuals of Collembola, Mesostigmata, and all microarthropods together dispersed when tunnels were present. Models that included excreta and earthworms were less well supported. Total numbers of dispersing Oribatida and Prostigmata+Astigmata were not well explained by any models tested. Further research is needed to examine the impact of soil structure and ecosystem engineering on movement belowground, as the substantial increase in movement of some microarthropods when corridors were present suggests these factors can strongly affect colonization and community assembly.

Diversity of feather mites (Acari: Astigmata) on Darwin's finches.

Feather mites are a diverse group of ectosymbionts that occur on most species of birds. Although Darwin's finches are a well-studied group of birds, relatively little is known about their feather mites. Nearly 200 birds across 9 finch species, and from 2 locations on Santa Cruz Island, Galápagos, were dust-ruffled during the 2009 breeding season. We found 8 genera of feather mites; the most prevalent genus was Mesalgoides (53-55%), followed by Trouessartia (40-45%), Amerodectes and Proctophyllodes (26-33%), Xolalgoides (21-27%), Analges and Strelkoviacarus (0-6%), and Dermoglyphus (2-4%). There was no evidence for microclimatic effects (ambient temperature and relative humidity) on mite diversity. Host body mass was significantly correlated with mean feather mite abundance across 7 of 8 well-sampled species of finches. Certhidea olivacea, the smallest species, did not fit this pattern and had a disproportionately high number of mites for its body mass.

Mesostigmatid mites (Acari: Mesostigmata) on rainforest tree trunks: arboreal specialists, but substrate generalists?

Predatory mites (Acari: Mesostigmata) on tree trunks without significant epiphytic growth in a subtropical rainforest in Eastern Australia were assessed for habitat specificity (i.e. whether they are tree trunk specialists or occupying other habitats) and the influence of host tree and bark structure on their abundance, species richness and species composition. The trunks of nine tree species from eight plant families representing smooth, intermediate and rough bark textures were sampled using a knockdown insecticide spray. In total, 12 species or morphospecies of Mesostigmata (excluding Uropodina sensu stricto) were collected, most of which are undescribed. Comparison with collections from other habitats indicates that epicorticolous Mesostigmata are mainly represented by suspended soil dwellers (six species), secondarily by generalists (four species) and a bark specialist (one species). A typical ground-dwelling species was also found but was represented only by a single individual. In terms of abundance, 50.5% of individuals were suspended soil dwellers, 40.7% bark specialists, and 8.3% generalists. Host species and bark roughness had no significant effect on abundance or species richness. Furthermore, there was no clear effect on species composition. The distribution of the most frequently encountered species suggests that most mesostigmatid mites living on bark use many or most rainforest tree species, independent of bark roughness. These findings support the hypothesis that some epicorticolous Mesostigmata use tree trunks as 'highways' for dispersing between habitat patches, while others use it as a permanent habitat.

THE EVOLUTION OF COPULATION IN WATER MITES: A COMPARATIVE TEST FOR NONREVERSING CHARACTERS.

Although copulation is a widespread behavior with multiple origins, hypotheses about selective forces behind its evolution have not been tested by the comparative method. Because copulation has arisen many times in the water mites (Acari: Parasitengona), they are good subjects for a comparative study of copulation. I determined that copulation evolved 91 times in the 343 extant genera. There was no evidence of reversals to noncopulation; therefore, Ridley's (1983) contingency-table comparative test was not appropriate. I designed a comparative test for instances in which there is no loss of the derived trait of interest. This test determines whether independent evolutions of a trait cooccur with a predictor more often than expected by chance. Two hypotheses were tested: 1) that copulation would be selected for in running-water habitats because of disruption of pheromonal communication; 2) that copulation would be selected for in swimming mites because females are less likely to contact spermatophores deposited on a substrate. Independent evolutions of copulation among running-water mites were not more frequent than expected by chance (P > 0.4), but there were more evolutions of copulation than expected among swimming mites (P < 0.005). Endoparasitism, secondary loss of swimming hairs, production of spermatophore fields, courtship, and benthivorous habits may explain why some mites copulate but do not swim while others swim but do not copulate.

Feather mites (Acari: Astigmata): ecology, behavior, and evolution.

Birds host many lineages of symbiotic mites, but the greatest diversity is shown by the three superfamilies of astigmatan feather mites: Analgoidea, Pterolichoidea, and Freyanoidea. Members of this diphyletic grouping have colonized all parts of the avian integument from their ancestral nidicolous habitat. Whereas some clearly feed on feather pith or skin, acting as parasites, other feather mites are paraphages and consume feather oils without causing structural damage. Sexual dimorphism in feather mites is often extreme, and little is known of the function of many elaborate male structures. Abundance and location of vane-dwelling mites is affected by season, temperature, light, humidity, and host body condition. Because transmission between hosts usually depends on host body contact, it is unsurprising that feather mite phylogeny often parallels host phylogeny; however, recent cladistic analyses have also found evidence of host-jumping and "missing the boat" in several mite lineages.

Red, distasteful water mites: did fish make them that way?

Water mites (Acari: Hydrachnida) are unusual among the typically cryptic freshwater fauna in that many species are brightly colored red or orange, and also appear to be distasteful to fish. This apparent aposematism (use of color to warn predators) has been previously explained as the evolutionary end-product of pressure from fish predation. The fish-predation argument has been supported by observations that fish spit out red mites, powder made from red water mites is more distasteful to fish than powder made from non-red mites, and red mites appear to be more abundant than non-red mites in water bodies where fish are present. In this paper, we challenge the hypothesis that fish were the sole driving force behind the evolution of aposematism in water mites. We show that non-red mites actually dominate in water bodies with fish, and that red mites are more abundant in temporary, fishless water bodies. We also demonstrate that powder made from red, terrestrial velvet mites (Trombidiidae) was as distasteful to fish as powder made from red water mites. We suggest that the main role of red and orange carotenoid pigments may be to act as photoprotectants, and hypothesize that redness originated in the terrestrial ancestors of water mites and has been retained in certain lineages of water mites after the invasion of the aquatic habitat. We also suggest that distastefulness evolved subsequent to bright coloration in response to increased conspicuousness to predators. Relaxed selection for redness has occurred when adults and/or larvae are less exposed to sunlight, either through occupying more protected habitats, parasitizing more nocturnal hosts, or parasitizing hosts for a short period of time. Our ability to test this alternative hypothesis is hampered by lack of knowledge of the source and mode of action of distastefulness, and of phylogenetic relationships among the Parasitengona.

Species coextinctions and the biodiversity crisis.

To assess the coextinction of species (the loss of a species upon the loss of another), we present a probabilistic model, scaled with empirical data. The model examines the relationship between coextinction levels (proportion of species extinct) of affiliates and their hosts across a wide range of coevolved interspecific systems: pollinating Ficus wasps and Ficus, parasites and their hosts, butterflies and their larval host plants, and ant butterflies and their host ants. Applying a nomographic method based on mean host specificity (number of host species per affiliate species), we estimate that 6300 affiliate species are "coendangered" with host species currently listed as endangered. Current extinction estimates need to be recalibrated by taking species coextinctions into account.