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.

Transitional chelal digit patterns in saprophagous astigmatan mites.

Changes in the functional shape of astigmatan mite moveable digit profiles are examined to test if Tyrophagus putrescentiae (Acaridae) is a trophic intermediate between a typical micro-saprophagous carpoglyphid (Carpoglyphus lactis) and a common macro-saprophagous glycyphagid (Glycyphagus domesticus). Digit tip elongation in these mites is decoupled from the basic physics of optimising moveable digit inertia. Investment in the basal ramus/coronoid process compared to that for the moveable digit mastication length varies with feeding style. A differentiated ascending ramus is indicated in C. lactis and in T. putrescentiae for different trophic reasons. Culturing affects relative investments in C. lactis. A markedly different style of feeding is inferred for the carpoglyphid. The micro-saprophagous acarid does not have an intermediate pattern of trophic functional form between the other two species. Mastication surface shape complexity confirms the acarid to be heterodontous. T. putrescentiae is a particularly variably formed species trophically. A plausible evolutionary path for the gradation of forms is illustrated. Digit form and strengthening to resist bending under occlusive loads is explored in detail. Extensions to the analytical approach are suggested to confirm the decoupling of moveable digit pattern from cheliceral and chelal adaptations. Caution is expressed when interpreting ordinations of multidimensional data in mites.

Phytoseiid mites benefited from organic fertilization by increasing the population of Tyrophagus mites in apple orchards.

This study explores sustainable agricultural practices by examining the role of organic materials in enhancing native predatory mites for controlling spider mites in apple orchards. Developing techniques to conserve indigenous natural enemies is vital for sustainable agricultural production. Phytoseiid mites can control spider mites, which are among the most significant pests in apple production. To conserve phytoseiid mite populations, it is important to identify alternative prey and to determine their role in phytoseiid mite proliferation. We demonstrated that the concurrent use of specific organic fertilizers and coconut husks can increase prey Tyrophagus mites, thereby enhancing phytoseiid mite density. Our research was conducted using sticky traps at the Miyagi Prefectural Agriculture and Horticulture Research Center in Japan. The occurrence of Tyrophagus mites was significantly correlated with the occurrence of phytoseiid mites in 2 years. In laboratory experiments, the use of organic fertilizers increased the density of Tyrophagus mites by 83 × within 4 weeks. Several species of phytoseiid mites were able to lay between 0.25 and 1.03 eggs per day per female by preying on Tyrophagus larvae. A 2-year field survey revealed that the use of organic fertilizers more than doubled the density of phytoseiid mites on apple leaves, likely through promoting Tyrophagus mite proliferation on the ground. These results highlight the potential of organic fertilizers not only to enhance soil nutrients, but also to boost phytoseiid mite populations, thereby contributing to more sustainable apple production.

Small-scale genetic structure of populations of the bulb mite Rhizoglyphus robini.

Bulb mites are an economically significant pest of subterranean parts of plants and a versatile laboratory animal. However, the genetic structure of their populations remains unknown. To fill this gap in our knowledge of their biology, we set up a field experiment in which we allowed mites to colonize onion bulbs, and then determined the genetic structure of colonisers based on a panel of microsatellite loci. We found moderate but significant population structure among sites separated by ca. 20 m (FST range 0.03-0.21), with 7% of genetic variance distributed among sites. Allelic richness within some bulbs was nearly as high as that in the total population, suggesting that colonisation of bulbs was not associated with strong population bottlenecks. The significant genetic structure we observed over small spatial scales seems to reflect limited dispersal of mites in soil.

Mite secretions from three traditional mite-ripened cheese types: are ripened French cheeses flavored by the mites (Acari: Astigmata)?

The opisthonotal glands of Astigmata contain monoterpenes, aromatics, aliphatics, and other volatile compounds; some of these compounds act as pheromones and have antifungal effects. This study analyzed volatile compounds secreted by mites on three traditional mite-ripened cheeses from producers (Milbenkäse from Germany, Mimolette and Artisou from France). The mites obtained from various traditional ripened French cheeses (Mimolette, Laguiole, Salers, and Cantal vieux) from stores were also investigated. The gas chromatography (GC) profiles of all their hexane extracts, except the Cantal vieux one, showed almost no differences and were identical to that of Tyrolichus casei Oudemans except for trace components. Based on the GC results, the mites of Cantal vieux were identified as Acarus siro L. For the Artisou and Cantal vieux, not studied before, the influence of the mite secretions on their characteristics was investigated by analyzing the headspace volatiles from the cheeses. According to the results, neral secreted from T. casei is the main compound responsible for the lemon-like flavor of the mite-ripened cheeses, which is, hence, due to a component of the mite secretions rather than the fermentation of the cheese itself. Moreover, the compounds secreted by the mites are not directly added to the cheese through ripening as they were not detected in the odors of the Artisou and Cantal vieux after the mites were removed. However, the consumers of the Artisou usually eat also the cheese rind, and thus, can enjoy its lemon-like flavor fully.

Do 'cheese factory-specific' mites (Acari: Astigmata) exist in the cheese-ripening cabinet?

To determine whether the mites used in the ripening process of traditional cheeses are genetically unique to cheese factories, we investigated mites from three types of traditional cheeses, that use mites in the ripening process: 'Würchwitzer Milbenkäse' from Germany and 'Mimolette' and 'Artisou' from France. In addition, traditional ripened cheeses were purchased from cheese specialty stores in France (Mimolette) and Japan ('Laguiole' from France) as well as stores in temporary markets in France ('Salers' and 'Cantal vieux') and the mites obtained from those cheeses were analyzed in this study. Partial sequences of the 28S rRNA gene (28S) were determined and used to reconstruct a phylogenetic tree. Tyrolichus casei, the dominant cheese mite species from the ripening cabinets of three traditional cheese producers and two cheese specialty stores in France and Japan, had identical partial 28S sequences. All specimens from Cantal vieux from a store in the temporary market in France had an identical sequence with Acarus siro and Acarus immobilis in the determined region of the 28S sequences. Mite individuals from Salers from a store in the temporary markets in France shared the same haplotype as Acotyledon paradoxa. For the T. casei individuals from five different localities (19 individuals in total), the nuclear loci were obtained using MIG-seq. More than several thousand genomic regions are amplified simultaneously by multiplex PCR, and targeting regions surrounded by inter-simple sequence repeats (ISSRs) in the genome were sequenced using the MiSeq system (Illumina). SNPs extracted from this genome-wide analysis showed that no genetic structure existed in the populations from any region. Among the five samples from the three regions, which were more than 500 km apart and from completely different environments, the mites had no geographic bias, but all mite individuals were genetically nearly identical. Thus, we found no evidence to support the existence of 'cheese factory-specific' T. casei mites, at least in terms of genetic analysis.

New feather mites of the Nycteridocaulus generic group (Acariformes: Proctophyllodidae) from passerines (Passeriformes) in Panama.

Three new feather mite species of the Nycteridocaulus generic group (Proctophyllodidae: Proctophyllodinae) are described from passerines in Panama: Atrichophyllodes myrmotherulae sp. n. from the Slaty Antwren, Myrmotherula schisticolor (Lawrence) (Thamnophilidae), Nycteridocaulus apanaskevichi sp. n. from the Grey-breasted Wood Wren, Henicorhina leucophrys (Tschudi) (Troglodytidae), and N. empidonicus sp. n. from the Yellowish Flycatcher Empidonax flavescens Lawrence (Tyrannidae). Nycteridocaulus apanaskevichi, presenting the second record of the genus from a host of oscine passerines, differs from N. guaratubensis Hernandes, 2014 in having the anterolateral extensions of the prodorsal shield rounded and the hysteronotal shield lacking any ornamentation. Males of N. empidonicus differ from N. myiobius Mironov, 2017 in having the supranal concavity open posteriorly and tarsus IV with rounded apical process; and females are distinguished by macrosetae h2 having long filiform apices. Males of A. myrmotherulae most clearly differ from A. mentalis Hernandes et al. 2007 in having the terminal lamellae rectangular and tarsus IV with triangular ventral process, and females are distinguished in having a noticeably longer idiosoma, 400-430 µm long. Comments on systematics and host associations of the genera Atrichophyllodes and Nycteridocaulus are provided.

Whole genomic sequencing and sex-dependent abundance estimation of Cardinium sp., a common and hyperabundant bacterial endosymbiont of the American house dust mite, Dermatophagoides farinae.

The two common species of house dust mites (HDMs), Dermatophagoides farinae and D. pteronyssinus, are major sources of allergens in human dwellings worldwide. Many allergens from HDMs have been described, but their extracts vary in immunogens. Mite strains may differ in their microbiomes, which affect mite allergen expression and contents of bacterial endotoxins. Some bacteria, such as the intracellular symbiont Cardinium, can affect both the sex ratio and biochemical pathways of mites, resulting in abundance variations of mite allergens/immunogens. Here, we investigated the bacterial microbiomes of D. farinae and D. pteronyssinus males and females using barcode 16S rDNA sequencing, qPCR, and genomic data analysis. We found a single species of Cardinium associated with D. farinae strains from the USA, China and Europe. Cardinium had high abundance relative to other bacterial taxa and represented 99% of all bacterial DNA reads from female mites from the USA. Cardinium was also abundant with respect to the number of host cells-we estimated 10.4-11.8 cells of Cardinium per single female mite cell. In a European D. farinae strain, Cardinium was more prevalent in females than in males (representing 92 and 67% of all bacterial taxa in females and males, respectively). In contrast, D. pteronyssinus lacked any Cardinium species, and the microbiomes of male and female mites were similar. We produced a Cardinium genome assembly (1.48 Mb; GenBank: PRJNA555788, GCA_007559345.1) associated with D. farinae. The ascertained ubiquity and abundance of Cardinium strongly suggest that this intracellular bacterium plays an important biological role in D. farinae.

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.

PCR identification and phylogenetic analysis of the medically important dust mite Suidasia medanensis (Acari: Suidasiidae) in Malaysia.

The occurrence of Suidasia medanensis (= S. pontifica) mites in Malaysian house dust was first reported in 1984. The taxonomy of this storage mite is, however, quite confusing. Therefore, we need an accurate identification to resolve morphological problems due to its minute size and some overlapping characters between species. The purpose of this study was to demonstrate the application of partial mitochondrial cytochrome c oxidase subunit I (COI) sequences for the identification of S. medanensis by PCR. Identity of the mite was first determined by observing morphological characters under a light microscope. Genomic DNA of S. medanensis mites was successfully extracted prior to PCR and DNA sequencing using COI universal primers. The length of the COI sequences obtained was 378 bp. BLAST analysis of amplicon sequences showed that local S. medanensis COI region had 99% maximum identity with S. medanensis nucleotide sequence (AY525568) available in the GenBank. As the phylogenetic tree generated indicated, COI sequences from this study were clustered with S. medanensis from Korea and the UK in one major clade, supported with high bootstrap value (> 85%). Results of the phylogenetic analysis of this COI gene were congruent with the morphological identification and provided strong support for a single clade of local S. medanensis.

Scent of a mite: origin and chemical characterization of the lemon-like flavor of mite-ripened cheeses.

Cheese infested with cheese mites is usually treated as unpalatable. Nevertheless, some traditional cheese manufactories in Germany and France intentionally use mites for fermentation of special varieties (i.e. Milbenkäse and Mimolette). While their production includes different mite species, both are characterized by a "lemon-like" flavor. However, the chemical nature and origin of this flavor-component is unknown. The cheese mites possess a pair of opisthosomal glands producing blends of hydrocarbons, terpenes and aromatics. Here, we describe the chemical profiles of the astigmatid mite species Tyrolichus casei (Milbenkäse) and Acarus siro (Mimolette). Although the chemical profiles differ in several aspects, both mite species produce neral (a volatile flavor component of lemon oil), which was absent from the headspace of both cheeses without mites. We conclude that the lemon-like flavor of mite cheese is not a consequence of fermentation of the cheese itself but a component from secretions of the cheese mites.

A new feather mite species of the genus Mycterialges Gaud & Atyeo, 1981 (Acari, Xolalgidae) from the Oriental Stork, Ciconiaboyciana (Ciconiiformes, Ciconiidae) in Korea.

A new feather mite species, Mycterialgesboycianaesp. nov. (Xolalgidae), was identified from the Oriental Stork, Ciconiaboyciana Swinhoe, 1873, in Korea. Males of M.boycianaesp. nov. are distinguished from Mycterialgesmesomorphus Gaud & Atyeo, 1981, in having a single triangular prodorsal shield, sinuous margins of the opisthosoma located between setae e2 and h2 on the hysteronotal shield, an oval-shaped epiandrum without posterior extensions, a shorter tibia + tarsus IV than femoragenu IV, and an absent ambulacral disc of leg IV. Females differ in having a prodorsal shield with a posterior margin that is blunt-angular, and a concave posterior margin of the hysteronotal shield with posterior extensions. This study presents the first record of the feather mite genus Mycterialges in birds of the genus Ciconia (Ciconiidae). Additionally, we determined the phylogenetic relationship among Ingrassiinae using the mitochondrial cytochrome c oxidase subunit (COI).

Population Genomics of Pooled Samples: Unveiling Symbiont Infrapopulation Diversity and Host-Symbiont Coevolution.

Microscopic symbionts represent crucial links in biological communities. However, they present technical challenges in high-throughput sequencing (HTS) studies due to their small size and minimal high-quality DNA yields, hindering our understanding of host-symbiont coevolution at microevolutionary and macroevolutionary scales. One approach to overcome those barriers is to pool multiple individuals from the same infrapopulation (i.e., individual host) and sequence them together (Pool-Seq), but individual-level information is then compromised. To simultaneously address both issues (i.e., minimal DNA yields and loss of individual-level information), we implemented a strategic Pool-Seq approach to assess variation in sequencing performance and categorize genetic diversity (single nucleotide polymorphisms (SNPs)) at both the individual-level and infrapopulation-level for microscopic feather mites. To do so, we collected feathers harboring mites (Proctophyllodidae: Amerodectes protonotaria) from four individual Prothonotary Warblers (Parulidae: Protonotaria citrea). From each of the four hosts (i.e., four mite infrapopulations), we conducted whole-genome sequencing on three extraction pools consisting of different numbers of mites (1 mite, 5 mites, and 20 mites). We found that samples containing pools of multiple mites had more sequencing reads map to the feather mite reference genome than did the samples containing only a single mite. Mite infrapopulations were primarily genetically structured by their associated individual hosts (not pool size) and the majority of SNPs were shared by all pools within an infrapopulation. Together, these results suggest that the patterns observed are driven by evolutionary processes occurring at the infrapopulation level and are not technical signals due to pool size. In total, despite the challenges presented by microscopic symbionts in HTS studies, this work highlights the value of both individual-level and infrapopulation-level sequencing toward our understanding of host-symbiont coevolution at multiple evolutionary scales.

The mite Acarus farris inducing defensive behaviors and reducing fitness of termite Coptotermes formosanus: implications for phoresy as a precursor to parasitism.

BACKGROUND: The ecology and evolution of phoretic mites and termites have not been well studied. In particular, it is unknown whether the specific relationship between mites and termites is commensal or parasitic. High phoretic mite densities have often been found to occur in weak termite colonies, suggesting that the relationship is closer to that of parasitism than commensalism. RESULTS: To examine this, Coptotermes formosanus was used as a carrier, and Acarus farris as the phoretic mite. We used video recordings to observe termite social immunity behaviors and bioassay to examine termite fitness. Our results showed that the attachment of the mite on the termite can enhance termite social immunity behaviors like alarm vibration and grooming frequency while decreasing the duration of individual grooming episodes in phoretic mites. Further, A. farris phoresy led to a 22.91% reduction in termite abdomen volume and a 3.31-fold increase in termite mortality. CONCLUSIONS: When termites groom more frequently, the consequence is short duration of grooming bouts. This may be indicative of a trade-off which provides suggestive evidence that frequent social behaviors may cost termites energy. And this caused phoretic behavior hastened termites' death, and helped propagate the population of mites feeding on dead termites. So, it provides a case for phoresy being a precursor to parasitism, and the specific relationship between A. farris and C. formosanus is closer to parasitism than to commensalism.

Life stages and phylogenetic position of the wool carder bee mite Sennertionyx manicati (Acari: Acaridae).

In mites of the family Acaridae, usually two developmental stages, adult (males, females) and deutonymphal stage (=hypopus) are used for diagnostic descriptions. Because these taxonomically important stages live in different habitats (i.e. deutonymphs are phoretic on a hymenopteran host, while adults live inside the nest of the host), one of these can be missing or they cannot be confidently corelated with each other. Sennertionyx manicati (Acari: Acaridae) was only known from hypopus. Herein, we describe all post-embryonic stages of S. manicati based on material obtained from a nest of Anthidium manicatum in Auckland, New Zealand, and provide a key to all life stages of this species. We also conducted a phylogenetic analysis of the 18S gene and the D2/D3 region of 28S genes from field-collected specimens and GenBank data. The molecular analysis indicated the New Zealand specimen shared the identical 18S and 99.5% identity of the D2D3 region of 28S sequences with S. manicati from Italy, and clustered together in the phylogenetic trees.

Feather mites of the genus Trouessartia (Acariformes: Trouessartiidae) from passerines (Aves: Passeriformes) in Georgia, USA.

Ten new species of the feather mite genus Trouessartia Canestrini, 1899 (Analgoidea: Trouessartiidae) are described from various passerines of the superfamily Passeroidea in Georgia: Trouessartia americana sp. n. from Setophaga americana (Linnaeus), T. helmitheros sp. n. from Helmitheros vermivorum (Gmelin, JF), T. mniotilta sp. n. from Mniotilta varia (Linnaeus), T. pensylvanica sp. n. from Setophaga pensylvanica (Linnaeus) (type host) and S. palmarum (Gmelin, JF), T. ruticilla sp. n. from S. ruticilla (Linnaeus), T. seiurus sp. n. from Seiurus aurocapilla (Linnaeus), T. tigrina sp. n. from Setophaga tigrina (Gmelin, JF) (Parulidae), T. passerinae sp. n. from Passerina caerulea (Linnaeus) (type host) and P. cyanea (Linnaeus), T. ciris sp. n. from P. ciris (Linnaeus) (Cardinalidae), and T. spizellae sp. n. from Spizella passerina (Bechstein) (Passerellidae). Based on a specific combination of morphological characters, all new species and six previously known species are arranged into a new species group capensis in the genus Trouessartia. The most important diagnostic characters of this species group include: in both sexes, the dorsal hysterosomal apertures are absent; in males, the postgenital plaque is well developed and genital setae g have cylindrical articulation rings; in females, the external copulatory tube is straight, stylet- or finger-like, and situated on the margin of the interlobar membrane, and the head of spermatheca has a semi-ovate extension without indentations. A key to all species referred to the capensis group is provided and host associations of this group with passerines are summarized and briefly discussed. It is hypothesized that this species group originated on the ancestors of New World nine-primaried oscines (Emberizoidea) and diverged in close relation with this group of hosts.

Ontogenetic stages of Oulenziella bakeri (Hughes) (Acari: Winterschmidtiidae).

The ontogenetic stages of mite family Winterschmidtiidae are rarely studied. Here we provide detailed descriptions and illustrations of all developmental stages of Oulenziella bakeri (Hughes). A comparison of the idiosomal and leg chaetotaxy of all stages are presented. New data on hosts and distribution of O. bakeri are provided.

Feather mites of the genus Trouessartia Canestrini (Acariformes: Trouessartiidae) from swallows (Passeriformes: Hirundinidae) in Canada.

Seven species of the feather mite genus Trouessartia Canestrini, 1899 (Astigmata: Trouessartiidae) have been recorded on swallows and martins (Passeriformes: Hirundinidae) in Manitoba (Canada). Of them, five are described as new species: Trouessartia ampulicaudata sp. n. and T. petrochelidon sp. n. from the American cliff swallow Petrochelidon pyrrhonota (Vieillot), T. bochkovi sp. n. from the tree swallow Tachycineta bicolor (Vieillot), and T. cryptocaudata sp. n. and T. progne sp. n. from the purple martin Progne subis (Linnaeus). A new species group stelgidopteryx, incorporating Trouessartia species living on hirundinids of the genera Progne, Tachycineta and Stelgidopteryx, is established. Renewed diagnoses of four Trouessartia species groups (appendiculata, crucifera, minutipes, and stelgidopteryx) restricted to hirundinids and a key to species recorded so far and potentially expected on swallows of North America are proposed. Host associations of Trouessartia species with swallows in North America are briefly discussed.

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.

Soil bulb mites as trace evidence for the location of buried money.

This study reports for the first time the use of soil micro-invertebrates, mites, as trace evidence to localise buried objects such as money. The case relates to a crime in Germany, where a large sum of banknotes had been hidden in an unknown location, likely abroad. In 2016, part of the money (approx. €500,000 in €500 value notes) was confiscated by the police. After analysis in the forensic lab, it was discovered that the notes were covered with small particles of a sort of debris, later identified as specimens of Rhizoglyphus howensis Manson, a non-European, nor Mediterranean species of root or bulb mites (Acaridae: Rhizoglyphinae). The restricted biogeographic distribution of R. howensis, originated in unspoiled forest soil in the Australasian region, limited the search for the money to the areas visited by the perpetrators during their trips to the region. Rhizoglyphus howensis biology provided further clues on the whereabouts of the banknotes, as they are specialist plant feeders, exclusively feeding on seeds of palm trees and on roots of Quercus patula in the Australasian region. This report aims to highlight the importance of the correct identification of the microscopic organisms associated with a crime scene and the immediate retrieval of micro-invertebrate trace evidence. This is the first record of R. howensis from Europe, and from banknotes.