Klimov, Pavel B. ¹ ; Kolesnikov, Vasiliy B. ² and OConnor, Barry M. ³

¹✉ Lilly Hall of Life Sciences, Purdue University, G-225, 915 W State St., West Lafayette, IN 47907, USA.
²Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, 152742 Borok, Yaroslavl Region, Russia & Zoological Institute, Russian Academy of Sciences, 199034, Saint Petersburg, Russia.
³Museum of Zoology, University of Michigan, 3600 Varsity Drive, Ann Arbor, MI 48108, USA.

2026 - Volume: 66 Issue: 4 pages: 946-1062

ZooBank LSID: C19A8243-BCEA-4732-B68F-62F4D7195C0F

Original research

Keywords

Abstract

This monograph was begun in the year 2000 and completed 25 years later — a span longer than both the Trojan War and Odysseus' homeward voyage. It therefore seems only fitting that the new species herein bear the names of Homeric heroes. And if this long labor says anything, it is the old truth that humans, in their vanity, often toil far beyond what the gods — or fate — ever intended.

Introduction

The family Acaridae sensu lato is among the most ecologically diverse groups of Astigmata, with species occupying a wide range of habitats and often utilizing phoretic associations with invertebrate and vertebrate hosts for dispersal (OConnor 2009). Although this taxon has been recognized on morphological grounds for nearly 90 years, molecular data did not support its monophyly. Instead, Acaridae sensu lato has recently been shown to comprise several monophyletic lineages, including Acaridae sensu novo and Rhizoglyphidae sensu novo (Klimov et al. 2026). These lineages can also be distinguished morphologically, the principal diagnostic character being the presence of seta aa on tarsus I in Acaridae sensu novo and its absence in Rhizoglyphidae sensu novo.

Members of each of these families occupy a great diversity of habitats (Türk and Türk 1957; Samšiňák 1961; Griffiths 1970; Hughes 1976; Fain et al. 1982; Olsen 1983; OConnor 1984; Nakao and Kurosa 1988; Klimov 1998b; Diaz et al. 2000; Klimov and OConnor 2003, 2008; Klimov et al. 2004, 2022a, b, 2023; Fan and Zhang 2004; Kasuga and Amano 2006; Rojas and Klimov 2007; Kolesnikov et al. 2024). Among Acaridae sensu novo, typical habitats include stored food and vertebrate nests (Acarus Linnaeus, 1758, Tyrophagus Oudemans, 1924), forest litter (Tyroborus Oudemans, 1924), compost, dung, and bodies of scarab beetles (Sancassania Oudemans, 1916), fungal mycelia (Tyrophagus), fungal sporocarps Sancassania (subgenus Rhizoglyphoides), nests of social insects including ants and bees (Forcellinia Oudemans, 1924, Froriepia Vitzthum, 1919, Cosmoglyphus Oudemans, 1932, Kuzinia Zachvatkin, 1941). Among Rhizoglyphidae sensu novo, typical habitats include subterranean plant organs such as bulbs and tubers (Rhizoglyphus Claparède, 1869), fungal sporocarps (Umakefeq Klimov,1999, Mezorhizoglyphus Kadzhaya, 1966, Boletoglyphus Volgin, 1953 and Capillaroglyphus Klimov, 1999), phytotelmata (Naiadacarus Fashing, 1974), freshwater habitats (Schwiebea Oudemans 1916), tree sap flows (Histiogaster Berlese, 1883), subcortical habitats (Schwiebea, Histiogaster, Reckiacarus Kadzhaya, 1972, Thyreophagus Rondani,1874), and terrestrial or freshwater crustaceans (Ewingia Pearse, 1929, Hoogstraalacarus Yunker, 1970, Askinasia Yunker, 1970).

Among Rhizoglyphidae sensu novo, polypore-associated mites occupy several distinct niches: (i) several species of Mezorhizoglyphus inhabit the context of perennial polypore sporocarps (the dense sterile tissue situated between the pileus surface and the tube layer); (ii) Umakefeq occurs in the hymenophoral tubes of thin, annual polypores (e.g., Bjerkandera spp.); and (iii) the best-known representatives, Boletoglyphus and Capillaroglyphus, inhabit the hymenophoral tubes of massive, long-lived perennial polypores, where they consume spores and hymenial tissue (Fig. 1) (Klimov 1998a, 1998b). Whereas the feeding stages of Boletoglyphus and Capillaroglyphus are trophically specialized on polypores, their non-feeding heteromorphic deutonymphs are phoretic on beetles associated with the same fungal hosts (Fig. 2) (Fain and Ide 1976; Khaustov et al. 2018; Klimov 1998a, 1998b). Most of these coleopteran hosts belong to the tribe Bolitophagini (Tenebrionidae) and occasionally to Ciidae (Klimov 1998a, 1998b). Only four species of these two genera are currently known, all from the Holarctic region: Boletoglyphus boletophagi (F. Turk et S. Turk, 1952), B. extremiorientalis Klimov, 1998, B. ornatus (Fain and Ide, 1976), and Capillaroglyphus polypori Klimov, 1998 (Turk & Turk 1952; Fain & Ide 1976; OConnor, 1984; Klimov 1998a, 1998b).

As a result of a large-scale survey of phoretic mites associated with beetles of the tribe Bolitophagini and other potential hosts, we discovered 15 new mite species belonging to Boletoglyphus (eight new species), Capillaroglyphus (four new species), and three new monotypic genera. To accommodate these five genera we propose the new tribe Boletoglyphini. Although most of these mites are associated with polypore bracket fungi, some presumably early derivative lineages (Sphaerotheriopus gen. nov., Alegoriopus gen. nov.) appear not to be associated with polypore fungi, possibly representing ancestral ecologies.

In this work, based on shared morphological characters and comparative analysis, we establish a new tribe, Boletoglyphini trib. nov. For this lineage, we infer a phylogenetic tree based on 19 ingroup taxa and 86 deutonymphal characters, with four outgroup genera. We describe the 15 new species and three new genera, and we redescribe the previously known species. Finally, we provide an identification key to all species of Boletoglyphini worldwide and discuss their geographic distributions and host associations.

Material and methods

Specimen sources

Deutonymphal mites were collected in the field from fungal sporocarps and from associated beetles. Bolitophagine beetle specimens were examined for mites in museum collections: CUIC, EMEC, IBSS, FMNH, NHMUK and ZIN (full names of the collections are in the following paragraph). Post-deutonymphal stages were collected by all authors and collaborators from fungal sporocarps in the field.

Morphological terminology

Idiosomal chaetotaxy followed Griffiths et al. (1990); the terminology of coxisternal setae followed Norton (1998); and the chaetotaxy and solenidiotaxy of appendages followed Grandjean for palps (Grandjean 1946) and legs (Grandjean 1939). Designations of tarsal dorsoapical setae on legs III–IV followed Klimov et al. (2020). The terminology of the spermatheca follows that of Witaliński et al. (1990) and Klimov and OConnor (2008). Continuous morphological characters were measured using an ocular micrometer mounted on a Zeiss Axioskop microscope. All measurements are given in micrometers (µm). Measurements of the holotype are given first, followed by the range for paratypes in parentheses. In taxonomic descriptions, we abbreviate the term ''heteromorphic phoretic deutonymph'' as HDN, while in the remainder of the text we use the shorter term ''phoretic deutonymph''. For the main Homer reference, we used the English translation of The Odyssey by S. H. Butcher and A. Lang (Homer, 1950).

Types

Types and voucher specimens are deposited in the following institutions: CNC - Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada; CUIC - Cornell University Insect Collection, Ithaca, New York, USA; EMEC - Essig Museum of Entomology, University of California, Berkeley, California, USA; FMNH - Field Museum of Natural History, Chicago, Illinois, USA; IBSS - Institute of Biology and Soil Science (Institute of Biology and Pedology), Vladivostok, Russia; MICH - University Herbarium, University of Michigan, Ann Arbor, Michigan, USA; NHMUK - The Natural History Museum, London, England, UK; NMSA - KwaZulu-Natal Museum, Pietermaritzburg, South Africa; QM - Queensland Museum, Brisbane, Australia; UMMZ - Museum of Zoology, University of Michigan, Ann Arbor, Michigan, USA; ZIN - Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.

Imaging

Line drawings were made by PBK and VBK using a camera lucida attached to a Zeiss Axioskop microscope. A few illustrations were prepared in Photoshop using processed microphotographs as the background. Microphotographs were captured by PBK and VBK with a Bioptic C-400 microscope equipped with DIC optics and a Euromex Color HD-Ultra digital camera. Images were taken from multiple focal planes and assembled in Helicon Focus 7.6.4 Pro (algorithm B, occasionally A), with subsequent manual editing to restore missing fine details from individual focal planes. Final images were combined and refined in Adobe Photoshop 22.2.0.

Phylogenetic analyses

Given that adults are only known for a few taxa (four species), our phylogenetic analyses were based on phoretic deutonymphs only. We used 86 characters (see Appendix 1) and 22 taxa, comprising 18 ingroup species of Boletoglyphini and four outgroups (Appendix 2). The outgroups, all from the family Rhizoglyphidae sensu novo, were chosen from species of the genera Rhizoglyphus, Histiogaster, Horstiella Turk, 1948, and Reckiacarus. Because molecular data indicate a distant phylogenetic position of Thyreophagus Rondani, 1874 (Klimov et al. 2026), this genus was not included in the analysis, as its inclusion destabilized the tree. For the outgroups (except Reckiacarus), data were obtained from the literature (Fan and Zhang, 2004; Klimov et al. 2022b; Ochoa and OConnor, 2000). The remaining taxa were examined by the authors using available specimens.

Phylogenetic analyses were done in PAUP v.4a168 under a maximum parsimony framework with all characters unordered. Maximum parsimony (MP) analyses were performed with a heuristic search strategy. The analysis included 100 random sequence addition replicates with tree-bisection-reconnection (TBR) branch swapping. Multiple trees were retained per replicate, with a maximum of 5,000 trees overall. The search was conducted without increasing the number of trees beyond this limit. Polymorphic characters were treated under the mstaxa = polymorph setting. Consensus trees were calculated using both strict and majority-rule criteria. Shortest MP trees with branch lengths were saved for further examination. Bootstrap support was assessed with 400 replicates using a heuristic search, and support values were reported for nodes present in ≥50% of bootstrap trees. Descriptive statistics including the consistency index, retention index, and diagnostic characters were recorded. For the above analysis, the following commands were used:

BEGIN PAUP;
log file = mp.log;
Pset mstaxa = polymorph;
delete Thyreophagus_potawatomorum;
Outgroup 1-4;
set autoclose = yes maxtrees = 5000 increase = no;
hsearch addseq=random nreps=100 multrees=yes hold=1 swap=tbr;
contree / strict=yes majRule=yes indices treeFile=′MR_consensus.tre′;
saveTrees file=′mp_shortest_trees.s.tree′ brLens=yes taxaBlk;
BOOTSTRAP NREPS=400 METHOD=HEURISTIC CONLEVEL=50;
Des 1/Ap=Yes La=Yes Diag=Yes;
log stop;
END;

Results

Phylogenetic analysis based on phoretic deutonymphs

Our analysis yielded 12 most-parsimonious trees (length = 290, Rohlf's CI = 0.770). The strict consensus tree supported the monophyly of Boletoglyphini (Fig. 3) and recovered the following relationships: Sphaerotheriopus, Alegoriopus, (Bolitoxenopus, (Capillaroglyphus, Boletoglyphus)). All genera were recovered as monophyletic (Fig. 3).

Within Boletoglyphus, the basal split separated B. anticleia (from Vietnam) from all remaining species. The next dichotomy separated a monophyletic clade comprising the two Palearctic species, B. boletophagi and B. extremiorientalis, from the remaining taxa. Subsequently, the Nearctic species B. ornatus was recovered as sister to a clade containing the remaining species from the Oriental and Australian regions (Fig. 3). Our phylogeny suggests that the occurrence of B. nausicaa in Australia resulted from a secondary dispersal event from Southeast Asia (Fig. 3).

Geographical distribution

Species of the tribe Boletoglyphini occur on all continents except Antarctica. Mite collections known to date largely reflect collecting effort, with major areas (e.g. China) remaining unsampled. The highest species richness is observed in Southeast Asia (Oriental region), where 12 species representing three genera have been recorded (Fig. 4). In the Palearctic region, three species belonging to two genera are known: Boletoglyphus boletophagi, B. extremiorientalis, and Capillaroglyphus polypori. Only a single species has been recorded from each of the following regions: B. ornatus in the Nearctic, B. nausicaa in Australia, Sphaerotheriopus odysseus in Africa (South Africa), and Alegoriopus demodocus in the Neotropical region.

The most widely distributed species is Boletoglyphus boletophagi, which occurs throughout the western Palearctic, extending eastward to eastern Siberia and southward to the Caucasus (Bugrov 1997; Haitlinger 2004; Haitlinger and Lupicki 2012; Khaustov et al. 2018; Klimov 1998b; Penttinen et al. 2013; Samšiňák 1957; Turk and Turk 1952; Volgin 1953). The northernmost records of this species are from Scotland and Finland. Boletoglyphus boletophagi has not been found in the Eastern Palearctic, where it is replaced by B. extremiorientalis, whose distribution is restricted to Eastern mainland Russia and the islands of Sakhalin and Kunashir in Russia and Hokkaido and Honshu in Japan (Klimov 1998a). The Nearctic species B. ornatus is known from eastern regions of the United States and Canada, while its occurrence in western North America remains unknown.

In Southeast Asia, species of Boletoglyphus are recorded mainly from the Philippine Islands (four species), Malaysia (two species, in eastern Borneo and the Malay Peninsula), and Singapore. The Malaysian species B. circe and B. eurycleia do not occur in the Philippines, and none of the Philippine species of Boletoglyphus have been found in Malaysia. A unique position within the genus is occupied by the single Australian species, B. nausicaa sp. nov., which occurs on the eastern coast of the continent.

Species of the genus Capillaroglyphus are also predominantly distributed in Southeast Asia, with the exception of C. polypori, which has a distribution similar to that of B. extremiorientalis (Klimov 1998a). In Southeast Asia, Capillaroglyphus species are known exclusively from Malaysia (two species from eastern Borneo) and the Philippines (two species). As in Boletoglyphus, Malaysian species of Capillaroglyphus do not occur in the Philippines, and vice versa.

This pattern suggests the presence of a well-defined biogeographic boundary between the Philippine Islands and Borneo affecting the distributions of both Boletoglyphus and Capillaroglyphus. In contrast, B. circe occurs both on Borneo and the Malay Peninsula without apparent barriers to dispersal. The only species of Boletoglyphini recorded from both Malaysia (Malay Peninsula) and the Philippines is Bolitoxenopus polyphemus.

Since both Sphaerotheriopus and Alegoriopus occupy basal positions on the phylogeny of Boletoglyphini, albeit with unresolved relationships, it can be inferred that the tribe likely originated in the Southern Hemisphere. From there, the remaining lineages appear to have migrated to the Northern Hemisphere, where they diversified extensively, particularly in the Oriental region, followed by secondary colonization of Australia (B. nausicaa) (Fig. 4).

Phoretic associations

With the exception of Sphaerotheriopus odysseus, an early-diverging taxon associated with Diplopoda, all mite species of the tribe Boletophagini are associated with beetles of the family Tenebrionidae (mostly from the tribe Bolitophagini), and less frequently with Ciidae (Table 1). Some beetles in both families are obligate or facultative fungivores whose larvae develop within the sporocarps of polypore fungi, where they feed on hymenial tissues and associated mycelium (Lawrence 1973; Nilsson 1997; Midtgaard et al. 1998; Jung et al. 2007; Arunraj et al. 2013). Adult beetles typically remain in close proximity to the same fungal substrates, using them for feeding, shelter, mating, and oviposition. The beetles usually feed on the same carpophore until the fungi are completely destroyed and decayed (Nilsson 1997; Rukke and Midtgaard 1998; Jung et al. 2007; Kobayashi and Sota 2021). This close ecological coupling between beetles and polypore fungi provides a stable habitat and predictable dispersal pathway for mites, whose feeding stages occupy the same fungal tissues and whose phoretic deutonymphs exploit adult beetles for transport between sporocarps.

When multiple beetle species compete for the same fungal resource, mites may use more than one phoretic host (see below). Likewise, if a phoretic host is not species-specific and feeds on multiple fungal species, this may facilitate the expansion of mites' fungal host ranges. However, this possibility requires further study, as fungal hosts are currently known for only 10 of the 19 mite species: Boletoglyphus boletophagi, B. extremiorientalis, B. ornatus, B. calypso, B. penelope, B. nausicaa, Capillaroglyphus polypori, C. eurymachus, C. hermes and Bolitoxenopus polyphemus (Klimov 1998a, 1998b; OConnor 1984).

The majority of tenebrionid coleopteran hosts belong to the tribe Bolitophagini; only a single species, the early-diverging Alegoriopus demodocus, was found on a representative of the tribe Ulomini (Alegoria sp.). Alegoria species are not associated with mushrooms, but live in a particular biotope - decaying banana trees. The diversity of stages of both larvae and adults, from immature to pregnant females, found in decaying pseudo-stems indicates that they spend their life there. They feed on the petioles surrounding the decomposing center of the tree and were never found associated with the dry petioles of healthy banana trees (Dutrillaux et al. 2010).

For six mite species, a host association has been recorded with only a single species of Bolitophagini (Table 1), suggesting that these mites are probably monoxenous species as per their phoretic host usage (Fain and Ide 1976; Mahunka 1977; Klimov 1998b). However, five mite species—Boletoglyphus boletophagi, B. extremiorientalis, B. circe, B. calypso, and Capillaroglyphus polypori—were recorded from multiple species of Bolitophagini, suggesting that they are polyxenous species. Among these, B. boletophagi, B. extremiorientalis, and C. polypori were also found on Ciidae. The highest number of beetle associations was observed in B. extremiorientalis (five host species) and C. polypori (four host species) (Table 1).

Ten species of Boletoglyphini exhibit synhospitality, occurring on the same beetle host species, although not necessarily on the same individual beetle: B. boletophagi and B. extremiorientalis on Bolitophagus reticulatus (Linnaeus, 1767); B. extremiorientalis and C. polypori on Boletoxenus incurvatus Lewis, 1894, Boletoxenus bellicosus Lewis, 1894, Atasthalomorpha dentifrons (Lewis,1894), and Cis jacquemartii Mellié, 1848; B. circe and Bolitoxenopus polyphemus on Boletoxenus spectabilis (Gebien, 1914); B. calypso and C. eurymachus on Boletoxenus timmi Gebien, 1922; B. circe and B. eurycleia on Byrsax sp.; and C. eumaeus and C. telemachus on Bolitonaeus vacca (Motschusky, 1858).

Finally, the following mite species were observed co-occurring on the same host individual: B. circe and B. eurycleia (host species Byrsax sp.); B. extremiorientalis and C. polypori (host species Atasthalomorpha dentifrons).

In addition to the above records there are some from what we consider to be accidental hosts: B. extremiorientalis, C. polypori on Cis jacquemartii (Coleoptera, Ciidae) (Klimov 1998a, 1998b), B. boletophagi on Ips typographus Linnaeus, 1758 (Coleoptera, Curculionidae, Scolytinae) (Penttinen et al. 2013; Khaustov et al. 2018) and Cis lineatocribratus Mellié, 1849 (Coleoptera, Ciidae) (Klimov 1998b), B. ornatus on Chrysolina coerulans (Scriba, 1791) (Hymenoptera, Chrysididae) and Trox foveicollis Harold, 1872 (Coleoptera, Trogidae) (personal observations by BMOC and PBK in UMMZ). In this study, we use the term accidental host for a beetle species on which a mite has been recorded incidentally but that is not considered a regular or ecologically established phoretic host. Such associations may result from incidental contact between mites and non-host beetles sharing the same habitat or substrate and do not, by themselves, demonstrate a stable phoretic relationship. We recognize that associations with Ciidae, which are generally small beetles associated with bracket fungi, may fall within a gray area of this definition, as additional study is needed to determine whether these associations represent established phoretic relationships or incidental occurrences. We do not consider the single record of Sphaerotheriopus with two deutonymphs to be accidental solely because it is based on a single collection; its status is considered in the context of the ecological association and the available evidence for phoresy.

In conclusion, the phoretic associations of Boletoglyphini species reflect a close ecological coupling with fungivorous beetles, particularly members of the tenebrionid tribe Bolitophagini, which provide both a reliable habitat and a means of dispersal between polypore sporocarps. While most mite species appear to be monoxenous, using a single beetle host species, several are clearly polyxenous, exploiting multiple beetle species, and occasional accidental associations with Ciidae indicate flexibility in phoretic host use. Synhospitality is widespread among Boletoglyphini, with multiple mite species co-occurring on the same beetle species, and in some cases even on the same individual, suggesting that these mites can coexist without strict competitive exclusion. These patterns indicate the importance of beetle–fungus interactions in shaping the ecology and dispersal of Boletoglyphini and indicate the need for further studies to document fungal host ranges and host specificity more comprehensively, particularly for lesser-known mite species and early-diverging taxa such as Alegoriopus demodocus.

Fungal Host Associations

For ten species of Boletoglyphini, associations with fungi of the order Polyporales are known, whereas for all remaining species no feeding association has yet been established (Table 1). Seven species have been recorded from fungi of the family Polyporaceae. A close association between Boletoglyphus boletophagi and Fomes fomentarius has been documented (Klimov 1998a, 1998b), and a similar specificity has been noted for B. ornatus and Ganoderma applanatum (OConnor 1984). One species, B. extremiorientalis, was found on Fomitopsis pinicola (Fomitopsidaceae) (Klimov 1998a), and three species (B. extremiorientalis, B. ornatus, Capillaroglyphus polypori) have been recorded from Ganoderma applanatum (Polyporaceae) (Klimov 1998a, 1998b). Finally, the Australian species, B. nausicaa have been found on Ganoderma australe (this study). The fungal hosts of the remaining species from this family have not been identified to species level. Thus, different species of Bolitophagini-associated mites may feed on the same fungal species, and conversely, a single mite species may exploit multiple fungal hosts.

Systematics

Suborder Oribatida van der Hammen, 1968

Hyporder Astigmata Canestrini, 1891

Family Rhizoglyphidae Oudemans, 1923

Tribe Boletoglyphini trib. nov.

Type genus — Boletoglyphus Volgin, 1953

ZOOBANK: 95C4F1D5-9965-4967-9A3B-54AE4D29A894

Diagnosis

Females (based on the three genera with known females: Bolitoxenopus, Capillaroglyphus, Boletoglyphus). Prodorsal shield without anterior incisions. Gnathosoma moderate in size to massive. Setae si present. Setae c₁, c₂, c₃, d₁, e₁ present or alveolar (c_1-3). Posterior apodemes II present. Anus ventral. Setae ps_1-3 present, filiform or alveolar. Leg podomeres (especially tarsi) short. Grandjean's organ narrow, finger-like. Setae ba I-II present (Boletoglyphus, Capillaroglyphus) or absent (Bolitoxenopus). Famulus elongated (at least 2 times shorter than ω₂, longer than ba I), spiniform or widened distally (never widest at middle). Tarsal seta aa I absent. Setae hT I-II absent. Ratio σ′/σ″I 2–2.5. Solenidion σ III present.

Males (based on the two genera with known males: Capillaroglyphus, Boletoglyphus). Posterior hysterosoma with sclerotization. Para-anal suckers ad₃ present, simple, disk-like; ps₁ and ps₂ short, filiform, ps₂ anterior to ps₃ at level of half of anal opening length (Capillaroglyphus) or close to ps₃ (Boletoglyphus). Tarsal solenidion ω₃ I subequal to ω₁. Tibial solenidion φ IV not modified in comparison with females. Tarsus IV with suckers d and e.

Heteromorphic phoretic deutonymph (HDN) (based on all included genera, see below). Anterior margin of propodosoma rounded or with a rostrum, completely covers gnathosoma. Vertical setae vi and ve (if ve present) situated on anterioventral propodosoma. Bases of supracoxal setae scx covered by prodorsal shield and not visible dorsally. Eyespots, when present, situated near setae se, without lenses; eyespots absent in Sphaerotheriopus, Alegoriopus, and Bolitoxenopus. Hysterosomal setae c₁, c₂, c₃, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃ present. Coxal fields III closed, anterior coxal apodeme III connected to ventral medial apodeme. Attachment organ with 2 pairs of conoids (ps₁ , ps₂ ) arranged in a transverse row, completely posterior to median suckers or rarely replaced by short, filiform setae. Famulus elongated.

Remarks

Females of Boletoglyphini differ from those of the remaining free-living Rhizoglyphidae (e.g., Histiogaster, Mezorhizoglyphus, Reckiacarus, Rhizoglyphus, Schwiebea, Thyreophagus) by setae ba I being shorter than the famulus (or ba I may be absent in Bolitoxenopus) (distinctly longer than the famulus in the remaining Rhizoglyphidae) and by the unmodified tibial solenidion φ IV in males (widened and shortened in comparison to females in the remaining Rhizoglyphidae). Phoretic deutonymphs differ from those of the remaining Rhizoglyphidae by having all conoidal setae (ps₁ -ps₂ ) of the attachment organ arranged in a transverse row situated posterior to the median suckers (not completely posterior in other Rhizoglyphidae), and by eyespots, when present, usually present near setae se (eyespots, when present, not associated with setae se in the remaining Rhizoglyphidae).

Females of Boletoglyphini differ from those of the tribe Thyreophagini (with a single genus, Thyreophagus) by the presence of idiosomal filiform setae si, c₁, c₂, c₃, d₁, e₁ (c_1-3 may be alveolar), posterior coxal apodemes II, and genual solenidion σ III present (these structures are absent in Thyreophagus). Grandjean's organ narrow (more than three times as long as it is wide), finger-like, without extensions (wide, less than twice as short as the width, with several finger-shaped extensions in Thyreophagus). Prodorsal shield without anterior incisions (incisions present in Thyreophagus). The famulus is elongated, never widest at middle (short, widest at the middle in Thyreophagus).

Phoretic deutonymphs of some Boletoglyphini (Capillaroglyphus and Boletoglyphus species), like those of Thyreophagus, have eyespots and a complete set of idiosomal setae (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). However, Boletoglyphini is distinguished by the absence of eye lenses (present in Thyreophagus); the anterior edge of the prodorsal shield overhanging the gnathosoma and the bases of the supracoxal setae, which conceals the gnathosoma (does not overhang the gnathosoma or the bases of the supracoxal setae in Thyreophagus); setae vi and ve (if present) ventral (setae vi not ventral in Thyreophagus, ve absent). Coxal fields III are closed, with anterior apodemes III connected to the ventral medial apodeme (coxal fields III are free, anterior apodeme III are not connected to the ventral medial apodeme in Thyreophagus). Conoidal setae ps₁ -ps₂ of the attachment organ are arranged in a transverse row completely posterior to the median suckers (never completely posterior to the median suckers in Thyreophagus).

Genera included

Boletoglyphus Volgin, 1953 (known from deutonymphs and adults of both sexes), Capillaroglyphus Klimov, 1998 (known from deutonymphs and adults of both sexes), Alegoriopus gen. nov. (known from deutonymphs only), Bolitoxenopus gen. nov. (known from deutonymphs and adult females) and Sphaerotheriopus gen. nov. (known from deutonymphs only).

Sphaerotheriopus gen. nov.

Type species — Sphaerotheriopus odysseus sp. nov.

ZOOBANK: 62A702A5-B91F-4213-8584-AC928CB85B56

Diagnosis

Phoretic deutonymph. Gnathosoma short, subcapitulum and palps fused, setae a, cm, sup and alveoli of h present. Dorsal surface of idiosoma with pattern of fine, evenly distributed pores. Anterior margin of propodosoma rounded, without radial pattern. Eyespots absent. Setae si situated posteriorly to se. All dorsal idiosomal setae filiform and smooth. Dorsomedian ''ridge'' on hysterosoma absent. Ratio hysteronotal shield/prodorsal shield length 2.46–2.52. Setae vi and ve situated on ventral rostrum. Supracoxal setae scx short. Sternum bifurcated, touching anterior coxal apodemes II, ends far from free ends of anterior coxal apodemes II. Sternal and ventrogenital shields touching. Medial apodeme III bifurcates posteriorly; ventral medial apodeme bifurcates anteriorly. Coxal setae 1a, 3a and 4a conoidal. Attachment organ distinctly separated from posterior edge of body. Cuticular suckers of attachment organ absent. Conoids ps₁ and ps₂ situated at posterior edge of attachment organ; sclerites of ps₂ and ps₁ distinctly separated. Leg setation: trochanter 1-1-1-0, femur 1-1-0-1, genu 2-2-1-0, tibia 2-2-1-1, tarsus 8-9-8-8. Tarsal setae aa, ba I absent. Tarsal setae f, ra, la, p, q I-IV and w III-IV with widened, leaf-shaped apices; setae e I-II with an apical ''saucer'', setae ba II, d I-IV, e I-II filiform, setae wa I alveolar, wa II spiniform. Setae hT I-II present. Seta nG III present. Solenidion ω₃ shifted anteriorly, situated in the middle of tarsus I (not situated in same cuticular field with ω₁). Famulus ε spiniform, with forked apex. Genual solenidia σ′, σ″ I, σ II, and σ III present; tibial solenidia φ I-IV present.

Etymology

The name Sphaerotheriopus is derived from the millipede host genus Sphaerotherium and the stem -op, commonly used in astigmatid systematics as a contraction of ''hypopus'', the historical term for the phoretic deutonymph. Gender masculine.

Remarks

The presence of a rounded anterior margin on the propodosoma overhanging the gnathosoma, ventral internal vertical setae vi, posterior position of both ps₁ and ps₂ and closed coxal fields I-IV in phoretic deutonymphs indicate that Sphaerotheriopus belongs to Boletoglyphini. However, Sphaerotheriopus differs from other genera of Boletoglyphini by the presence of seta cm on the gnathosoma, the bifurcated medial apodeme III and ventral medial apodeme, two solenidia on genu I (σ′, σ″ I), genual seta nG III present, and tarsal solenidia ω₃ and ω₁ separated (not situated in the same cuticular field).

Sphaerotheriopus odysseus sp. nov.

(Figs 5, 6)

ZOOBANK: cDE199D2-DE30-40AB-A327-B143207D8708

Type material

Holotype — HDN – South Africa KwaZulu Natal, Kranskop Forest (28°58′S, 030°51′E), ex Sphaerotherium sp. (Diplopoda, Sphaerotheriida, Sphaerotheriidae), 1994, M. Hamer (NMSA). Paratype: 1 HDN – same data (UMMZ).

Description

Phoretic deutonymph (Figs 5, 6; measurements: Table 2). Body ovoid (Fig. 5), 1.3 times longer than wide, widest in sejugal region; idiosomal length 256 (295) width 197 (n = 2).

Gnathosoma (Fig. 6A) short (length of gnathosoma approximately 0.98 (1.3) times width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h present, anterio-lateral (sup), posterior-lateral (a) and anteroventral (cm) setae present, filiform.

Dorsum (Fig. 5B). Dorsal surface of idiosoma with pattern of fine, evenly distributed pores. Anterior margin of propodosoma rounded, without radial pattern. Setae si approximately equal to se; both filiform, smooth, si situated posteriorly to se. Eyespots absent. Sejugal furrow well developed. Prodorsal shield 74 (83.8), hysteronotal shield 182 (211.5), ratio hysteronotal shield/prodorsal shield length = 2.46 (2.52). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of elongated, filiform, smooth setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). All setae (except h₃) shorter than femur I, seta c₂ 14 (13.5). Alveoli of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than length of corresponding setae. Cupules ia large, posterior to line connecting setae c₂ and c_p , ip not observed.

Venter (Fig. 5A). Setae vi and ve situated in ventral part of rostrum: seta vi 3.3 (3.8) times longer than minute ve, smooth. Supracoxal setae of legs I (scx) short, filiform, with slightly extended base, positioned ventrally. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between its end and ventrogenital shield longer than its length. Posterior edge of sternal shield and anterior edge of ventrogenital shield not observed. Ventrogenital shield not elongated. Sternal and ventrogenital shields touching, posterior apodemes II fused or in contact with anterior apodemes III. Coxal fields III closed, medial apodeme size equal to ventral medial apodeme, bifurcate posteriorly, ventral medial apodeme bifurcate anteriorly. Coxal setae 1a, 3a and 4a conoidal, small but well-developed, 4b filiform, both setae present. Seta 4a situated posteriorly to g. Genital seta g not touching edge of ventrogenital shield. Attachment organ shifted from posterior edge of body (distance between them more than attachment organ length). Cuticular suckers of attachment organ absent. Setae ps₁ and ps₂ conoidal, well-developed, small; ps₂ shifted posteriorly and medially. Sclerites and alveoli ps₁ and ps₂ well separated from each other. Anterior suckers (ad₃) without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih smaller than im; central spot well-developed on both structures.

Legs (Figs 6B–H) short, all segments free. Trochanters I–III each with long filiform seta, pR I–II, sR III. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform. Genual setation 2-2-1-0; setae mG I–II and cG I-II weakly spiniform, smooth, seta nG III present, filiform. Tibial setation 2-2-1-1; setae hT I–II, gT I-II and kT III-IV spiniform. Tarsal setation 8-9-8-8. All pretarsi consisting of thin and slightly hooked empodial claws (without tooth) attached to short paired condylophores. Tarsus I with setae ra, la, f, p, q with leaf-like terminal dilations, e with ''saucer''; seta d filiform, shorter than ra or la; setae s and wa represented by alveoli; setae aa and ba absent. Tarsus II similar to tarsus I except very short setae ba and wa present, wa represented by very small remnant. Tarsi III and IV with setae r, w, s, f, e, q, p with leaf-like terminal dilations, d filiform, smooth, longer than leg; p and q extending to end of pretarsus. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I longer but thinner than ω₁ I, ω₃ I tapering, situated on the middle of tarsus. Solenidion ω₂ longer than famulus, with rounded apex. Famulus (ε) spiniform, not inflated proximally, with forked apex. Solenidia of tibiae I elongate, tapering; φ I longer than tarsus I with empodium; φ II longer than tarsus II without empodium; φ III shorter than tarsus II without empodium; φ IV short, with rounded apex. Genu I with two solenidion; σ′ I twice as long as σ″ I, both with rounded apices, σ′ reaching base of φ I. Solenidion σ II not reaching base of φ II, with pointed apex. Genu III with short solenidion σ.

Distribution

South Africa.

Host

Sphaerotherium sp. (Diplopoda, Sphaerotheriida, Sphaerotheriidae).

Etymology

Etymology. The new species is named after Odysseus, the protagonist of Homer's Odyssey, king of Ithaca, and a participant in the Trojan War. His return home after the war was delayed for ten years by the wrath of Poseidon and numerous misadventures. The name is a noun in apposition.

Type deposition

Holotype in NMSA, paratype in UMMZ.

Alegoriopus gen. nov.

Type species — Alegoriopus demodocus sp. nov.

ZOOBANK: 23B8A643-20C8-48E8-8629-0914C314A7AE

Diagnosis

Phoretic deutonymph. Gnathosoma long, setae sup and a present. Dorsal surface of idiosoma with a pattern of coarse pores. Anterior margin of propodosoma rounded, with a fine radial pattern. Setae si situated posteriorly to se. All dorsal idiosomal setae filiform and smooth. Dorsomedian ''ridge'' on hysterosoma absent. Setae vi situated on ventral rostrum; setae ve absent. Eyespots absent. All coxal fields closed. Sternum bifurcate, touching anterior coxal apodemes II, ending near ends of anterior coxal apodemes II (both ends almost on the same level); distance between sternum end and ventrogenital shield much shorter than sternum length. Coxal setae 1a, 3a and 4a conoidal. Distance between attachment organ and posterior edge of body equal or more the half of attachment organ length. Anterior, posterior paired and posterior unpaired cuticular suckers of attachment organ present. Suckers ps₁ and ps₂ conoidal, ps₂ shifted posteriorly and medially; sclerites of ps₁ and ps₂ separate from each other. Leg setation of trochanters 1-1-1-0, femora 1-1-0-1, genua 2-2-0-0, tibiae 2-2-1-1, tarsi 8-8-8-8. Setae wa I-II present, aa, ba I-II absent. Setae ra, la, f, p, q I-II, f, e, q III-IV, e III and p IV widened, with leaf-shaped apices. Setae hT I-II present. Seta nG III absent. Setae d III-IV with fine barbs near base. Solenidia ω₁ and ω₃ situated in same cuticular base. Famulus ε not inflated proximally. Genu I with one solenidion, genu III without a solenidion.

Etymology

The genus name is derived from the name of the host beetle genus Alegoria and the stem -op, commonly used in astigmatid systematics as a contraction of hypopus, the historical term for the phoretic deutonymph. Gender masculine.

Remarks

This is the only genus in Boletoglyphini that lacks external vertical setae ve and has barbed setae d on tarsi III-IV. However, the presence of a rounded anterior propodosomal margin overhanging the gnathosoma, ventral internal vertical setae vi, posterior position of both of ps₁ and ps₂, tarsal solenidia ω₁ and ω₃ situated in the same cuticular field, and closed coxal fields I-IV indicate that this genus belongs to Boletoglyphini. Like Sphaerotheriopus, this genus has tibial setae hT I-II and gnathosomal seta a.

Alegoriopus demodocus sp. nov.

(Figs 7, 8)

ZOOBANK: 3D8B4A00-6CD5-4A34-B928-55749FC38F7C

Type material

Holotype — HDN – Panama Panama Prov., Parque Nacional Soberanía, Pipeline Rd, km 8, (09°09′04″N, 079°46′50″W), ex Alegoria sp. (Coleoptera, Tenebrionidae, Ulomini) ventral prothorax, on Heliconia mariae Hook f., 1864 (Zingiberales, Musaceae), 4 May 1991, B. M. OConnor, BMOC 91-0504-019 (UMMZ). Paratypes: 2 HDN – same data (UMMZ). Host in UMMZ.

Additional material — 16 HDN - Peru, Cadena, Rio Ampiyacu, 10 km NW Pevas (02°50′S, 072°30′W), ex Alegoria sp., 14 Nov 1961, B. Malkin, BMOC 01-0327-014 (FMNH, UMMZ). Host in FMNH. 15 HDN - Costa Rica Cartago, CATIE, 3 km SE Turrialba, 600m. (09°53′25″N, 083°39′05″W), ex Alegoria dilatata (LaPorte, 1840) (Coleoptera, Tenebrionidae), ventral thorax and abdomen, 1 Jun 1985, J.T. Doyen, BMOC 15-0207-090 (EMEC, UMMZ). Host in EMEC.

Description

Phoretic deutonymph (Figs 7, 8; measurements: Table 2). Body elongated, 1.67 (1.73–1.83) times longer than wide, widest in sejugal region; idiosomal length 293.3 (294.8–298.8) width 176 (163.5–170.3) (n = 3).

Gnathosoma (Fig. 8A) long (length of gnathosoma approximately 2.8 (2.8–3.0) times longer than wide), subcapitulum and palps fused, bearing short (shorter than length of gnathosoma) palpal solenidia ω, alveoli of seta h present or absent (holotype), if present, situated approximately at the middle of transverse levels of posterolateral and anterolateral setae, anterolateral seta (sup) filiform, posterolateral (a) represented by elongated spine, filiform, anteroventral (cm) seta absent.

Dorsum (Fig. 7B). Dorsal surface of idiosoma with pattern of coarse pores (2.0–3.0), denser on the sides; in central part pores larger than peripheral ones, smoothly punctate between them. Anterior margin of propodosoma rounded, with fine radial pattern. Setae si much shorter than se; both filiform, smooth, si situated posteriorly to se. Eyespots absent. Sejugal furrow well developed. Prodorsal shield 110 (111.8–112.8), hysteronotal shield 183.3 (182–187), ratio hysteronotal shield/prodorsal shield length = 1.67 (1.61–1.67). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of elongated, filiform, smooth setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). Setae c₂ and e₁ approximately equal to femur I length, h₃ longer than femur I, c_p slightly shorter than femur I, seta c₂ 26 (20). Transverse levels of c₁ and d₁ near each other, distance between them shorter than femur I length. Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than length of corresponding setae. Cupules ia large, situated dorsally between setae c₂, c_p, ip situated dorsolaterally anteriomediad of seta f₂, without distinct median spot.

Venter (Fig. 7A). Setae vi situated on anteroventral part of rostrum, smooth. Setae ve absent. Supracoxal setae of legs I (scx) short, filiform, with slightly extended base, positioned ventrally. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends near free ends of anterior coxal apodemes II (both ends almost on the same level); distance between its end and ventrogenital shield much shorter than its length. Sternal shield closed posteriorly, posterior edge concave anteriorly. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance (0.8–1.0). Coxal fields III closed, medial apodeme longer than ventral medial apodeme, ventral medial apodeme short. Coxal setae 1a, 3a and 4a conoidal, well-developed, 4b filiform, both setae present. Seta 4a posterior to g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body equal or more the half of attachment organ length. Anterior, posterior paired and posterior unpaired cuticular suckers of attachment organ present. Setae ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Alveoli of ps₁ and ps₂ represented by sclerotized rings around corresponding conoids, not extending beyond posterior edge of attachment organ; sclerites (apodemes) of ps₁ and ps₂ separated from each other. Anterior suckers (ad₃) without transparent marginal edge; central part has dotted sculpture, while peripheral part with fine radial pattern. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common heavily pitted sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih and im subequal; median spot well-developed on both structures.

Legs (Figs 8B–I) short, all segments free. Trochanters I–III each with long filiform seta, pR I–II, sR III. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform. Genual setation 2-2-0-0; setae mG I–II filiform, weakly barbed, cG I-II spiniform, smooth, seta nG III absent. Tibial setation 2-2-1-1; setae hT I–II, gT I-II and kT III-IV spiniform, barbed. Tarsal setation 8-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. Tarsus I with setae ra, la, f, p, q with leaf-like terminal dilations, e with ''saucer'', wa spiniform, without attenuated tips; seta d filiform, shorter than ra or la; seta s represented by alveolus; setae aa and ba absent. Tarsus II similar to tarsus I except seta d with widened bases. Tarsus III with setae s, f, e, q foliate, w, r, p filiform, d filiform, with fine barbs near base, slightly shorter than leg; p and q extending to end of pretarsus. Tarsus IV similar to tarsus III except seta r spiniform, e with leaf-like terminal dilations and p with leaf-like terminal dilatation. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I shorter and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ longer than famulus, with rounded apex. Famulus (ε) not inflated proximally. Solenidion φ of tibia I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, relatively long, reaching base of φ I. Solenidion σ II not reaching base of φ II, with rounded apex. Genu III without solenidion.

Distribution

Neotropical: Panama, Costa Rica, Peru.

Host

Alegoria dilatata, Alegoria sp. (Coleoptera: Tenebrionidae)

Etymology

The new species is named after Demodocus, the blind bard who performed at the court of Alcinous, king of the Phaeacians on the island of Phaeacia (Scheria). During Odysseus' stay on Scheria, before revealing his identity, Demodocus performed three songs recounting episodes related to the Trojan War and Odysseus' adventures, moving Odysseus to tears. According to the Odyssey, the Muse deprived Demodocus of sight but granted him the gift of song. The specific epithet refers to the absence of eyes in the new species, in contrast to many species of Boletoglyphini, which possess eyespots. Noun in apposition.

Type deposition

Holotype and paratypes are in UMMZ; additional specimens in EMEC, FMNH, UMMZ.

Bolitoxenopus gen. nov.

Type species — Bolitoxenopus polyphemus sp. nov.

ZOOBANK: 255A59BF-9A75-4655-803E-52A42EB03798

Diagnosis

Female. Prodorsal shield wider than its length, smoothly punctate, without incisions. Setae vi separated, ve vestigial. Supracoxal setae (scx) smooth, thin. Idiosoma with 13 pairs of setae (se, si, c₂, c_p, c₃, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae c₂ and d₂ short, seta c₁ absent. Four pairs of fundamental cupules (ia, im, ip and ih) present. Posterior part of opisthosoma without sclerotized shield. Ventral idiosoma with four pairs of coxal setae (1a, 3a, 4a and 4b), one pair of genital setae, three pairs ps_1-3 and three pairs short ad_1-3. Leg setation: trochanter 1-1-1-0, femur 1-1-0-1, genu 2-2-0-0, tibia 1-1-1-1, tarsus 10-10-10-10. Setae ba I-II, aa I, wa I-II absent, Setae e, s, u, v, p and q spiniform, other setae without terminal dilations. Solenidion ω₂ present. Genu I with two solenidia, III with one solenidion. Famulus (ε) with rounded apex, slightly widened distally.

Male. Unknown.

Phoretic deutonymph. Gnathosoma elongated, subcapitulum and palps separated, only setae sup present. Dorsal surface of idiosoma without pattern of coarse pores. Form of anterior margin of propodosoma rounded, with more or less developed rostrum, without radial pattern. Eyespots absent. Setae si situated posterior to se, all dorsal idiosomal setae filiform and smooth. Dorsomedian ''ridge'' on hysterosoma absent. Prodorsal shield short, ratio hysteronotal shield/prodorsal shield length 5.5–8.75. Setae vi and ve situated on ventral part of rostrum. Supracoxal setae scx short. Sternum not bifurcated, ending freely. Coxal setae 1a, 3a and 4a conoidal. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Posterior paired cuticular suckers of attachment organ present, anterior and unpaired posterior suckers absent. Suckers ps₁ and ps₂ conoidal, ps₂ shifted posteriorly and medially, separated from ps₁. Legs with setation of trochanter 1-1-1-0, femur 1-1-0-1, genu 2-2-0-0, tibia 1-1-1-1, tarsus 7-7-8-8. Setae wa, aa, ba I-II absent. All leg setae (except r IV) without terminal dilations. Seta nG III absent. Solenidion ω₃ situated in same cuticular field as ω₁. Famulus (ε) solenidion-shaped with rounded tip. Genu I with one solenidion, genu III without solenidion.

Etymology

The name Bolitoxenopus is derived from the coleopteran host genus Boletoxenus and the stem -op, commonly used in astigmatid systematics as a contraction of ''hypopus'', the historical term for the phoretic deutonymph. Gender masculine.

Remarks

Females of Bolitoxenopus differ from Boletoglyphus and Capillaroglyphus females by the absence of tarsal setae ba I-II. However, the presence of setae c₂, c₃, d₁, e₁, ps_1-3, a prodorsal shield without anterior incisions, and a clavate famulus indicates that this genus belongs to Boletoglyphini.

Phoretic deutonymphs of Bolitoxenopus differ from those of all other genera of Boletoglyphini by the very short prodorsal shield and separated palps on the gnathosoma. However, the presence of a rounded anterior margin of the prodorsal shield overhanging the gnathosoma, ventral setae vi, posterior displacement of ps₁ and ps₂, placement of solenidion ω₃ in the same cuticular field as ω₁, and closed coxal fields on phoretic deutonymph indicate that this genus belongs to Boletoglyphini.

Bolitoxenopus polyphemus sp. nov.

(Figs 9–11)

ZOOBANK: cF495B56-6355-401A-851E-6417EF89BD2B

Type material

Holotype — HDN – Singapore, Atkinson coll. 92-3, ex Boletoxenus spectabilis (Gebien, 1814) (Coleoptera, Tenebrionidae, Bolitophagini), no date and collector name, host in NHMUK, BMOC 82-1221-003 (NHMUK). Paratypes: 4 HDNs – same data (NHMUK, UMMZ); 1 female, 4 HDNs – Philippines, Romblon Prov., Sibuyan Is., 2.4 km. S, 4 km. E Magdiwang, 325 m., Mt. Guiting-Guiting, 12°26′46″N, 122°33′13″E, 6 May 1989, S. M. Goodman, ex very decayed Polyporaceae (Polyporales), BMOC 89-1550-011 (host fungus in MICH) (UMMZ); 3 HDN – Philippines, Romblon Prov., Sibuyan Is., 5.25 km. S, 3.5 km. E Magdiwang, 725 m., Mt. Guiting-Guiting, 12°25′10″N 122°33′15″E, 21 May 1989, S. M. Goodman, ex undetermined Polyporaceae (Polyporales), BMOC 89-1550-014 (host fungus in MICH) (UMMZ); 20 HDN - Malaysia Perak, Tapah Hills Forest Reserve, 04°19′N, 101°22′E, 31 May 2020, no collector, ex Boletoxenus sp., BMOC 22-0418-001 (UMMZ).

Description

Female (Figs 10, 11). Idiosoma rounded, slightly elongate (Figs 10A, B), 500 x 300 (n = 1), 1.66 times longer than wide. Idiosomal cuticle smooth.

Gnathosoma with subcapitular setae (h), lateral dorsal palp tibial setae (sup), dorsal palp tarsal seta (cm) filiform, h longer than other this seta, palp tibial setae a absent; supracoxal seta elcp absent; terminal palp tarsal solenidion short; external part of terminal eupathidium ul'' dome-shaped, small; terminal eupathidium ul′ not observed. Palps slender. Chelicera chelate, with one seta.

Prodorsal shield 82 long, 153 wide, 0.5 times longer than wide, with setae vi situated at anterior part of shield, alveoli separated (distance between them significantly greater than their diameter), shield without incisions (except small midlateral incisions - alveoli of vestigial setae ve). Prodorsal shield smoothly punctate. Supracoxal setae (scx) smooth, thin, sword-shaped, widened and flattened, tapering at tip. Grandjean's organ not observed. Idiosomal setae (vi, se, si, c_p, d₁, e₁, e₂, f₂, h₁, h₂, h₃) smooth, filiform, approximately equal size, setae c₂ and d₂ short filiform, setae c₃ longer than c₂ but shorter than c_p. Seta c₁ not observed (anterior region of hysterosoma partially obscured by gut contents). Four pairs of fundamental cupules (ia, im, ip and ih) present. Posterior part of opisthosoma without sclerotized shield.

Ventral idiosoma with four pairs of coxal setae (1a, 3a, 4a and 4b) and one pair of genital setae (g). Shape of coxal sclerites as in Fig. 10A, sejugal apodeme present. Genital region situated between coxal fields III and IV; genital valves form an inverted Y; epigynal and medial apodemes well-developed. Genital papillae rounded, blunt. Anal opening ventral, in posterior part of idiosoma. Three pairs of setae ps_1-3, filiform and three pairs adanal setae ad_1-3, ad₃ slightly longer than minute ad_1-2. Copulatory tube not developed, copulatory opening and spermathecal duct not observed.

Legs (Figs 11) short, all segments free. Trochanters I–III each with short, filiform seta, pR I–II, sR III; trochanter IV without seta. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform. Genual setation 2-2-0-0; setae mG and cG I–II short, slightly spiniform; seta nG III absent. Tibial setation 1-1-1-1; setae gT I–II filiform, kT III–IV weakly spiniform, setae hT I-II absent. Tarsal setation 10-10-10-10; pretarsi consist of hooked empodial claws attached to short paired condylophores. Tarsus I with setae ra, la, f and d filiform, s, e, u, v, p and q spiniform, seta e and u longer than other spiniform setae, setae aa, ba and wa absent. Tarsus II similar to tarsus I except setae e and v longer than other spiniform setae. Tarsi III and IV with setae f, d, r and w filiform, s, e, u, v, p and q spiniform, p represented by small triangular rudiments. Solenidion ω₁ on tarsus I cylindrical, with clavate apex, not bent; solenidion ω₁ on tarsus II simple, cylindrical, with clavate apex, longer and wider than ω₁ on tarsus I. Solenidion ω₂ on tarsus I shorter than ω₁ and approximately equal to famulus, cylindrical, with rounded apex, slightly widened distally, situated slightly distal to ω₁. Solenidion ω₃ on tarsus I cylindrical, with slightly tapering tip, slightly longer than ω₁. Famulus (ε) of tarsus I wide, with rounded apex, slightly widened distally. Solenidia φ of tibiae I–III elongate, tapering, extending beyond apices of respective tarsi with ambulacra; solenidion φ IV tapering, shorter than tarsus IV (without ambulacra). Solenidia σ″ and σ′ on genu I elongate, σ″ longer than σ′, σ″ reaching base of φ I. Solenidion σ on genua II and III more than 6–7 times longer than its width, with rounded tip.

Male unknown.

Phoretic deutonymph (Figs 9; measurements: Table 2). Body ovoid (Figs 9A, B), 1.44 (1.12–2.0) times longer than wide, widest in sejugal region; idiosomal length 109.5 (100.5–117) width 76.1 (72–99) (n = 5).

Gnathosoma (Fig. 9C) elongated (length of gnathosoma approximately 2.0 (1.6–3.0) times the width), subcapitulum and palps separated, bearing palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) filiform, posterolateral (a) and anteroventral (cm) seta absent.

Dorsum (Fig. 9B). Dorsal surface of idiosoma without pattern of coarse pores, smoothly punctate. Form of anterior margin of propodosoma rounded, with more or less developed rostrum, without radial pattern. Setae si 1.2-1.3 times shorter than se; both filiform, smooth, si situated posteriorly to se. Eyespots absent. Sejugal furrow well developed. Prodorsal shield 14 (12–16), hysteronotal shield 95.5 (88–105), ratio hysteronotal shield/prodorsal shield length = 6.82 (5.5–8.75). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of filiform, smooth setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). All setae (except h₃) shorter than femur I, seta c₂ 3.4 (2.7–4). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ greater than length of corresponding setae. Cupules ia large, situated between c₁ - c_p, ip large, situated dorsolaterally posterior to setae e₂, with distinct median spot.

Venter (Fig. 9A). Setae vi and ve situated on ventral part of rostrum: seta vi 5.29 (4.47–6.8) times longer than minute ve, smooth. Supracoxal setae of legs I (scx) short, filiform, with slightly expanded base, positioned ventrally above leg I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum not bifurcated, not touching anterior coxal apodeme II, ending near free ends of anterior coxal apodemes II; distance between its end and ventrogenital shield much shorter than its length. Sternal shield not closed posteriorly. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance. Coxal fields III closed, its medial apodeme shorter than ventral medial apodeme; ventral medial apodeme long. Coxal setae 1a, 3a and 4a conoidal, well-developed, 4b filiform, both setae present. Seta 4a placed posteriorly to g. Genital seta g not touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length (distance between them less than 3/4 the attachment organ length). Posterior paired cuticular suckers of attachment organ present. Anterior and unpaired posterior suckers absent. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ not touching each other. Anterior suckers (ad₃) without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih smaller than im; median spot well-developed on both structures.

Legs (Figs 9D–J) short, all segments free. Trochanters I–III each with long filiform seta, pR I–II, sR III. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform. Genual setation 2-2-0-0; setae mG I–II filiform, smooth, cG I-II weakly spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent, gT I-II filiform, kT III-IV filiform with widened bases. Tarsal setation 7-7-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. Tarsi I and II with setae ra, la, f, d, filiform, setae p, q, slightly widened terminally, e with rounded terminal dilation (''saucer''); seta d shorter than ra or la; seta s represented by alveolus; setae aa, wa and ba absent. Tarsus III with setae r, w, s, f, e, d, without leaf-like terminal dilations, q, p slightly expanded distally, all with widened bases, d shorter than leg; p and q protruding beyond pretarsus. Tarsus IV similar to tarsus III except seta r spiniform, q massive. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I size equal to ω₁ I but thinner than ω₁ I, with rounded apex, situated close to ω₁. Solenidion ω₂ slightly shorter than famulus, with rounded apex. Famulus (ε) solenidion-like with rounded tip, or very slightly inflated with acute tip. Solenidion φ of tibia I elongate, tapering; φ I and III longer than tarsi I and III with empodium; φ II shorter than tarsus II without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, reaching base of φ I. Solenidion σ II not reaching base of φ II, with pointed apex. Genu III without solenidion.

Distribution

Oriental: Singapore and Philippines.

Host

Boletoxenus spectabilis, Boletoxenus sp.

Etymology

The new species is named after Polyphemus, a Cyclops and son of Poseidon. During his return to Ithaca from the Trojan War, Odysseus encountered Polyphemus and blinded him to escape being eaten. The specific epithet alludes to the absence of eyes in the deutonymph of the new species, in contrast to many other species of Boletoglyphini that possess eyespots. Noun in apposition.

Type deposition

Holotype is in NHMUK, paratypes are in NHMUK, UMMZ.

Genus Capillaroglyphus Klimov, 1998

Capillaroglyphus Klimov, 1998a: 6; Klimov and Tolstikov, 2011: 257.

''Unnamed acarid mite'' Kurosa, 2000: 942, Fig. 29.

Type species — Capillaroglyphus polypori Klimov, 1998, by original designation

Diagnosis

Female. Body elongated. Setae si and se long. Setae c_1-3, d₁ and d₂ short, filiform, ps_1-3 present, short, other idiosomal setae long, filiform. Posterior part of hysterosoma covered by opisthosomal shield, shaped as ''visor'' concealing copulatory opening. Anal opening shifted posteriad, with its posterior end reaching the level of posterior margin of hysterosoma and extending somewhat dorsally. Internal opening of spermatheca as transverse plate with short efferent ducts on edges. Legs short. Setae ba I-II spiniform, short. Setae wa I-II, wF IV present. Genu III with solenidion.

Male. Posterior part of hysterosoma with sclerotization. Suckers ad₃ present, simple, disk-like; ps₁ and ps₂ short setiform, ps₂ situated anterior to ad₃ at level of half of anal opening. Solenidion ω₃ I long, φ IV long. Tarsus IV with suckers d and e.

Phoretic deutonymph. Prodorsal shield long (ratio hysteronotal shield/prodorsal shield length less than 3.0), with trapezoidal rostrum. Eyespots present. Seta ve present. Gnathosoma elongated, well-developed, subcapitulum and palps fused, gnathosomal solenidia shorter than subcapitulum; setae sup spiniform or knife-like, setae a and cm absent. Sejugal furrow well developed. Dorsal surface of idiosoma with pattern of small pores, dorsomedian ''ridge'' on hysterosoma absent. Dorsal idiosomal setae relatively long. Sternal shield closed posteriorly. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Posterior cuticular suckers of attachment organ may be present. Setae hT I-II and nG III absent. Genu I with one solenidion. Famulus clavate.

Species included

Capillaroglyphus eumaeus sp. nov., C. eurymachus sp. nov., C. hermes sp. nov., C. polypori, C. telemachus sp. nov.

Capillaroglyphus eumaeus sp. nov.

(Figs 12, 13)

ZOOBANK: 281EB059-6765-446F-A455-47FCCA12BF40

Type material

Holotype — HDN – Philippines Mindanao, [Zamboanga Sibugay],''Pt. Bange,'' [Port Banga], F. C. Fletcher, ex Bolitonaeus vacca (Motschulsky, 1858) (Coleoptera, Tenebrionidae), no date, BMOC 79-0301-004 (host beetle in CUIC). Paratypes: 6 HDNs – same data as for holotype(CUIC, UMMZ).

Description

Phoretic deutonymph (Figs 12, 13; measurements: Table 2). Body ovoid (Figs 12), 1.4 (1.34–1.46) times longer than wide, widest in sejugal region; idiosomal length 241 (234–256) width 170 (164–182) (n = 7).

Gnathosoma (Fig. 13A) short, length of gnathosoma approximately 1.57 (1.53–1.7) times width, subcapitulum and palps fused, bearing short (shorter than length of subcapitulum) palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, spiniform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Fig. 12A). Dorsal surface of idiosoma with pattern of fine (0.4–0.7), evenly distributed pores, pores absent in medial part of hysterosoma, smoothly punctate. Anterior margin of propodosoma with short rostrum, no radial pattern. Setae si approximately equal to se; both filiform, smooth or very slightly pectinate, si situated posteriorly to se. Eyespots present, without lenses, situated near bases of seta se. Sejugal furrow well developed. Prodorsal shield 80 (74–81), hysteronotal shield 160.5 (153–178), ratio hysteronotal shield/prodorsal shield length = 2.01 (1.89–2.28). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of elongated, filiform setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), at least c₁, c₂, d₁, d₂, e₁, h₁, h₂ very slightly pectinate. All setae approximately equal in length and slightly longer or approximately equal to femur I, seta c₂ 26.5 (31–38). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ shorter than or approximately equal to length of corresponding setae. Cupules ia large, situated between c₂, c_p, ip situated dorsolaterally between e₂, f₂, without distinct median spot.

Venter (Fig. 12B). Setae vi and ve situated in ventral part of rostrum: seta vi 5.15 (4.25–4.57) times longer than minute ve, smooth. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated posteriorly, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between its end and ventrogenital shield approximately 1/3 of its length. Sternal shield closed posteriorly, posterior edge straight. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance, almost touching each other. Coxal fields III closed, its medial apodeme short. Ventral medial apodeme present. Coxal setae 1a and 4a conoidal, well-developed, 3a vestigial, with attenuated tips, 4b filiform, short, both setae present. Seta 4a posterior to g. Genital seta g not touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Anterior paired and posterior unpaired cuticular suckers of attachment organ absent; posterior paired suckers fused with attachment organ shields. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ surrounding corresponding conoids, not protruding posteriorly, separated from each other. Anterior suckers (ad₃ alveoli) without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih equal in size to im; median spot well-developed on both structures.

Legs (Figs 13B–I) short, all segments free. Trochanters I–III each with short, filiform seta, pR I–II, sR III; pR I–II on protrusions. Femoral setation 1-1-0-1; setae vF I–II and wF IV filiform, vF I–II on protrusions, medium length, wF IV short. Genual setation 2-2-0-0; setae mG I–II and cG I-II spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III-IV spiniform, smooth. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsus I with setae ra, la, f, d filiform, long, without terminal dilations, e spiniform, p and q spiniform, massive, with attenuated tips; seta d longer than ra or la; seta s represented by alveolus; setae aa, ba and wa absent. Tarsus II similar to tarsus I except seta ba present, filiform, with widened base, situated close to ω₁. Tarsus III with setae f, d filiform, smooth, without terminal dilations, w and r with widened bases, e, s, q and p spiniform with attenuated tips, s size equals q, massive; p not protruding beyond short pretarsal ambulacrum, q protruding beyond pretarsus; d smooth, slightly shorter than leg. Tarsus IV similar to tarsus III except seta r spiniform with attenuated tip. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I approximately 1.1 times longer and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ longer (approximately twice as long) than famulus, with rounded apex. Famulus (ε) inflated distally. Solenidion φ of tibia I elongate, tapering, longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV short, with rounded apex. Genu I with one solenidion, σ I with rounded apex, reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion.

Distribution

Oriental: Philippines.

Host

Bolitonaeus vacca.

Etymology

The new species is named after Eumaeus, the faithful swineherd of Odysseus, who remained loyal to Odysseus, Penelope, and Telemachus during his master's absence. With his help, Odysseus defeated the suitors and returned home. Noun in apposition.

Type deposition

Holotype and paratypes in CUIC, other paratypes in UMMZ.

Remarks

This species differs from all others in its massive setae s III-IV with short, attenuated tips, setae s III-IV in size and proportions not different from q III-IV. All other species of the genus have s III-IV with elongated attenuated tips.

Capillaroglyphus eurymachus sp. nov.

(Figs 14, 15)

ZOOBANK: 922D37C5-81A5-4388-9307-40F13FFC603D

Type material

Holotype — HDN – Philippines, Rombolon Prov., Sibuyan Is., 2.4 km S, 4 km E Magdiwang, 325 m., Mt. Guiting-Guiting, 12°26′46″N, 122°33′13″E ex Polyporaceae (Polyporales), 6 May 1989, S. M. Goodman, BMOC 89-1550-011 (UMMZ). Paratypes: 4 HDNs, 1 pharate HDN, 3 protonymphs - same data as for holotype (UMMZ); 2 HDNs – same general locality as for holotype 5.25 km S, 3.5 km E Magdiwang, 725 m., Mt. Guiting-Guiting, 12°25′10″N, 122°33′15″E, ex Polyporaceae (Polyporales), 22 May 1989, S. M. Goodman, BMOC 89-1550-015 (UMMZ); 5 females, 2 males, 6 HDNs, 2 protonymphs, – Philippines Negros Oriental, Lake Balinsasayao, 3 km N 14 km W Dumaguete, 9°22′N 123°11′E, 980 m, ex Polyporaceae (Polyporales), 9 Nov 1982, P. D. Heideman #1102, BMOC 83-1800-043 (UMMZ); 12 HDNs – same locality, ex Boletoxenus timmi Gebien, 1922 (Coleoptera: Tenebrionidae), 9 Nov 1982, P. D. Heideman #1103, BMOC 83-1800-044, in previously noted polypore (beetle in UMMZ), (UMMZ).

Description

Phoretic deutonymph (Figs 14, 15; measurements: Table 2). Body elongated (Figs 14), 1.84 (1.72–2.05) times longer than wide, widest in sejugal region; idiosomal length 213 (206–238) width 116 (104–126) (n = 10).

Gnathosoma (Fig. 15A) short (length of gnathosoma approximately 1.5 (1.45–1.6) times width), subcapitulum and palps fused, bearing short (shorter than length of subcapitulum) palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, spiniform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Fig. 14A). Dorsal surface of idiosoma with pattern of fine (0.4–0.6), evenly distributed pores, pores are absent in medial part of hysterosoma, smoothly punctate between pore fields. Anterior margin of propodosoma with short rostrum, no radial pattern. Setae si approximately equal to se; both filiform, smooth, si situated posteriorly se. Eyespots present, without lenses, situated near bases of seta se. Sejugal furrow well developed. Prodorsal shield 60 (50–74), hysteronotal shield 153 (154–170), ratio hysteronotal shield/prodorsal shield length = 2.55 (2.16–3.08). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of elongated, filiform, smooth setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). All setae approximately equal in length and slightly shorter or approximately equal to femur I length, seta c₂ 19 (20–23). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than length of corresponding setae. Cupules ia large, situated between c₂ and c_p, ip situated dorsolaterally, near seta f_2, without distinct median spot.

Venter (Fig. 14B). Setae vi and ve situated on ventral part of rostrum: seta vi 7.5 (5.4–10.5) times longer than minute ve, smooth. Supracoxal setae of legs I (scx) filiform, with slightly extended bases, positioned ventrally dorsal to legs I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between end of sternum and ventrogenital shield approximately 1/3 of its length. Sternal shield closed posteriorly, posterior edge straight. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance, almost touching each other. Coxal fields III closed, its median apodeme short. Ventral medial apodeme present. Coxal setae 1a and 4a conoidal, well-developed, 3a vestigial, with attenuated tips, 4b filiform, short, both setae present. Seta 4a posterior to g. Genital seta g not touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Anterior paired and posterior unpaired cuticular suckers of attachment organ absent; posterior paired suckers fused with attachment organ shields. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ surround corresponding conoids, not protruding posteriorly, separated from each other. Anterior suckers (ad₃) without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, at level of c₃. Cupules ih size equal to im; median spot well-developed on both structures.

Legs (Figs 15B–I) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III; pR I–II on protrusions. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform, vF I–II on protrusions. Genual setation 2-2-0-0; setae mG I–II and cG I-II spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III-IV spiniform, smooth. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsus I with setae ra, la, f, d filiform, long, without terminal dilations, e, p and q spiniform, massive, with attenuated tips; seta d longer than ra or la; seta s represented by alveolus; setae aa, ba and wa absent. Tarsus II similar to tarsus I except seta ba present, filiform, with widened bases, situated close to ω₁. Tarsus III with setae r, f, d filiform, smooth, without terminal dilations, w with widened bases, e, s, q and p spiniform with attenuated tips; p not protruding beyond pretarsal ambulacrum, q protruding beyond pretarsal ambulacrum; d smooth, slightly shorter than leg. Tarsus IV similar to tarsus III except seta r spiniform with attenuated tip. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ longer (approximately twice as much) than famulus, with rounded apex. Famulus (ε) inflated distally. Solenidion of tibiae I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV very short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III with short solenidion, with rounded apex.

Adult female and male. Specimens in poor condition and not suitable for description.

Distribution

Oriental: Philippines.

Host

Boletoxenus timmi.

Morphological abnormalities

In a single deutonymph (BMOC 89-1550-011, 3), ω₃ on one tarsus is shifted distally.

Etymology

The new species is named after Eurymachus, one of the leading suitors of Penelope in Homer's Odyssey. Noun in apposition.

Type deposition

Holotype and paratypes are in UMMZ.

Remarks

Deutonymphs of this species differ from all other species of the genus in the elongated body (1.72–1.97 times longer than wide vs. 1.34–1.6 in other species). Note that the fungal host (BMOC 83-1800-043) was also the host for the predatory mite, Neochelacheles corpusrarosae Bochkov & OConnor, 2004 (Cheyletidae), which probably preys on boletoglyphine mites.

Capillaroglyphus hermes sp. nov.

(Figs 16, 17)

ZOOBANK: b1A7B15A-740F-47E2-A652-E1C8AC9D5B65

Type material

Holotype — HDN – Malaysia Sabah, Mt. Kinabalu National Park, 1500-2000 m. elev., ex polypore fungus (Polyporales, Polyporaceae), S. M. Goodman, 7-9 Apr 1998, BMOC 88-0927-005 (UMMZ). Paratypes: 8 HDNs – same data as for holotype BMOC 88-0927-004 (UMMZ).

Additional material — 9 HDN – Philippines Palawan, Roxas, Magara Barangay, 10°11′57″N, 119°09′24″E, ex Boletoxenus sp. (Coleoptera, Tenebrionidae), N. Layron, January 2024, BMOC 24-0402-001 (UMMZ).

Description

Phoretic deutonymph (Figs 16, 17; measurements: Table 2). Body slightly elongated (Figs 16), 1.55 (1.5–1.6) times longer than wide, widest in sejugal region; idiosomal length 235 (223–251) width 152 (140–167) (n = 9).

Gnathosoma (Fig. 17A) short (length of gnathosoma approximately 1.57 (1.5–1.92) times exceeds width), subcapitulum and palps fused, bearing short (shorter than length of subcapitulum) palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, spiniform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Fig. 16A). Dorsal surface of idiosoma with pattern of fine (0.4–0.6), evenly distributed pores, pores are present in medial part of hysterosoma, smoothly punctate between them. Anterior margin of propodosoma with elongated rostrum, no radial pattern. Setae si approximately equal to se; both long, band-like, flattened, smooth dorsoventrally, and pectinate laterally; si situated posteriorly to se. Eyespots present, without lenses, situated near bases of setae se. Sejugal furrow well developed. Prodorsal shield 80 (77.3–82.5), hysteronotal shield 155 (146–163), ratio hysteronotal shield/prodorsal shield length = 1.94 (1.88–1.98). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), all setae (except filiform c_p and h₃) long, flattened dorsoventrally and pectinate laterally. All setae longer than femur I, seta c₂ 50 (43–55). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ shorter than length of corresponding setae. Cupules ia large, situated between setae c₂, c_p, ip situated dorsolaterally, between e₂, f₂, without distinct median spot.

Venter (Fig. 16B). Setae ve situated on ventral part of rostrum, ve on dorsal part of rostrum: seta vi 7.5 times longer than minute ve, smooth. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above leg I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated (branches weakly sclerotized), touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between its end and ventrogenital shield approximately 1/4-1/5 of its length. Sternal shield closed posteriorly, posterior edge straight. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance, almost touching each other. Coxal fields III closed, its medial apodeme short. Ventral medial apodeme present. Coxal setae 1a and 4a conoidal, well-developed, 3a vestigial, with attenuated tips, 4b filiform, short, both setae present. Seta 4a placed posterolateral to g. Genital seta g not touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Anterior paired and posterior unpaired cuticular suckers of attachment organ absent; posterior paired suckers almost fused with attachment organ shields. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ surround corresponding conoids, not protruding posteriorly, separated from each other. Anterior suckers (ad₃) without transparent marginal edges. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih equal in size to im; median spot well-developed on both structures.

Legs (Figs 17B–I) short, all segments free. Trochanters I–III each with long filiform seta, pR I–II, sR III; pR I–II on protrusions. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform, vF I–II on protrusions, medium length. Genual setation 2-2-0-0; setae mG I–II and cG I-II spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III-IV spiniform, smooth. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsus I with setae ra, la, f, d filiform, long, e spiniform, p and q spiniform, massive, without attenuated tips; seta d longer than ra or la; seta s represented by alveolus; setae aa, ba and wa absent. Tarsus II similar to tarsus I except seta ba present, filiform, with widened base, situated close to ω₁. Tarsus III with setae r, f, d filiform, smooth, without terminal dilations, w, e, s, q and p spiniform with attenuated tips, s slightly longer than q; p not protruding beyond pretarsal ambulacrum, q protruding beyond ambulacrum; d smooth, slightly shorter than leg. Tarsus IV similar to tarsus III except seta r spiniform with attenuated tip. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter (1.1) and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ more than twice as long as famulus, with rounded apex. Famulus (ε) inflated distally. Solenidion φ of tibia I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion.

Distribution

Oriental: East Malaysia, Philippines.

Host

Boletoxenus sp.

Etymology

The new species is named after Hermes, the Greek messenger god, associated with trade, good fortune, cunning, theft, youth, and eloquence. Noun in apposition.

Type deposition

Holotype and paratypes are in UMMZ.

Remarks

This is the only species in the genus that has dorsal idiosomal setae distinctly flattened and pectinated, and internal vertical setae vi situated in a dorsal position.

Capillaroglyphus polypori Klimov, 1998

Capillaroglyphus polypori Klimov, 1998a: 8, Figs. 1-32; Klimov and Tolstikov, 2011: 257.

''Unnamed acarid mite'' Kurosa, 2000: 942, Fig. 29.

(Figs 18–20)

Type material

Holotype — female – Russia, Primorskii Krai, Vladivostok, Botanical Garden, 43°13′N, 122°33′E, mixed conifer-broadleaf forest, in hymenium of decaying bracket-fungus growing on Betula sp., 27 Aug 1995 (IBSS) (Klimov 1998a). Paratypes: 55 females, 13 males, 13 HDNs – same date as holotype (IBSS); 41 HDNs – same locality, 12 May 1995 (IBSS); 15 females, 5 males, 12 HDNs – same locality, 23 May 1995 (IBSS); 9 females, 10 males, 1 HDN – same locality, 30 Jun 1995 (IBSS); 1 female – same locality, 20 Aug 1995 (IBSS); 9 females, 2 males – Russia, Primorskii Krai, Kedrovaya Pad' reserve, 43°06′N, 131°29′E, same habitat, 8 Jun 1997 (IBSS); 1 female – same locality, 16 Jun 1997 (IBSS); 2 HDNs – same locality, ex Cis jacquemartii Mellié, 1849 (Coleoptera, Ciidae), 9 Jun 1997 (IBSS); 1 HDN – Japan Hokkaido, Nopporo Forest Park, 10 km SE Sapporo, 43°02′N, 141°31′E, ex Boletoxenus bellicosus (Lewis, 1894) (Coleoptera, Tenebrionidae), 2 Jul 1992, G. Lafer (IBSS) (Klimov 1998a).

Additional material — 1 HDN – Japan, Hokkaido, Sapporo-shi, Minami-ku, Minamisawa, 42°59′N, 141°18′E, ex Boletoxenus incurvatus (Lewis,1894) (Coleoptera, Tenebrionidae), 31 May 2017, M. Endo (ZIN); 3 HDNs – Japan, Hokkaido, Abutarun, Kimobetsu-cho, Mt. Sorioi, 45°45′N, 140°53′E, ex Boletoxenus incurvatus (Lewis, 1884), 17 Jun 2017, M. Endo (ZIN); 1 HDN – Japan, Honshu, Kanagawa Pref., Mt. Tanzawa, 35°28′27″N, 139°09′46″E, ex Boletoxenus bellicosus 25 Jul 1965, H. Nakamura (ZIN historical collection) (ZIN); 28 HDN's - Japan, no further data, G. Lewis ex Atasthalomorpha dentifrons BMOC 82-1221-004 (host in NHMUK) (NHMUK, UMMZ); 30 HDN's Japan, Honshu, Tanagawa, Hakone, 35°11′22″N, 139°01′39″E, ex Atasthalomorpha dentifrons (Lewis, 1894), 17-19 Apr 1880, G. Lewis, BMOC 82-1221-001 (host in NHMUK) (NHMUK, UMMZ); 4 HDNs – Russia, Kunashir Is., Mendeleyevo, 43°58′35″N, 145°42′08″E, ex Atasthalomorpha dentifrons, 15 Jul 1975, B. Korotaev (ZIN historical collection) (ZIN).

Redescription

Phoretic deutonymph (Figs 18–20; measurements: Table 2). Body ovoid (Figs 18, 20A, B), 1.47 times longer than wide, widest in sejugal region; idiosomal length 224–241 width 152–163 (n = 4).

Gnathosoma (Figs 19G, 20F) short (length of gnathosoma approximately 1.26–1.5 times width), subcapitulum and palps fused, bearing short (shorter than length of subcapitulum) palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, spiniform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Figs 18A, 20A, C–E). Dorsal surface of idiosoma with pattern of fine (0.6–0.8), evenly distributed pores, smoothly punctate between them. Anterior margin of propodosoma with short rostrum, no radial pattern (Figs 20D, C). Setae si approximately equal to se; both long, band-like, flattened, smooth dorsally/ventrally, and pectinate laterally; si situated posterior to se. Eyespots present, without lenses, situated near bases of setae se. Sejugal furrow well developed. Prodorsal shield 67–78, hysteronotal shield 147–177, ratio hysteronotal shield/prodorsal shield length = 1.88–2.64. Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of long, slightly dorso-ventrally flattened setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃). All setae approximately equal in length and longer than femur I, seta c₂ 34–52. Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ shorter than length of corresponding setae; distance h₂-h₂ slightly shorter. Cupules ia large, situated between c₂, c_p, ip situated dorsolaterally between e₂, f₂, without distinct median spot.

Venter (Figs 18B, 20B,G,H). Setae vi and ve situated on ventral part of rostrum: seta vi 6.6–10.0 times longer than minute ve, smooth. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated (branches very weakly-sclerotized), touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between its end and ventrogenital shield more than 1/3 of its length. Sternal shield closed posteriorly, posterior edge straight. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance, almost touching each other. Coxal fields III closed, its medial apodeme short. Ventral medial apodeme present. Coxal setae 1a and 4a conoidal, well-developed (Fig. 20H), 3a with thick cylindrical basal part and tapering apically (Fig. 20G), 4b filiform, both setae present. Seta 4a posterior to g. Genital seta g slightly on edge of ventrogenital shield. Distance between attachment organ (Fig. 20H) and posterior edge of body shorter than half of attachment organ length. Anterior paired and posterior unpaired cuticular suckers of attachment organ absent; posterior paired suckers almost fused with attachment organ shields. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ surround corresponding conoids, not protruding posteriorly, separated from each other. Anterior suckers (ad₃) with transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih and im equal in size; median spot well-developed on both structures.

Legs (Figs 19A–F, 20D) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III; pR I–II on protrusions (Fig. 20F). Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform, vF I–II on protrusions. Genual setation 2-2-0-0; setae mG I–II and cG I-II spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III-IV spiniform, smooth. Tarsal setation 7-8-8-8. All praetarsi consisting of hooked empodial claws (without tooth) attached to short, paired condylophores. Tarsus I with setae ra, la filiform, long, thinly expanded terminally, d, f filiform, without terminal dilation, e spiniform, p and q spiniform with attenuated tips; seta d slightly longer than ra or la; seta s represented by alveolus; setae aa, ba and wa absent. Tarsus II similar to tarsus I except seta ba present, filiform, with widened base, situated close to ω₁. Tarsus III with setae w, r, f, d filiform, smooth, without terminal dilations, e and p spiniform without attenuated tips, q and s spiniform with attenuated tips; p not protruding beyond pretarsal ambulacrum, q protruding beyond ambulacrum; d smooth, slightly longer than leg. Tarsus IV similar to tarsus III except seta r spiniform without attenuated tip, e smooth, with widened base and attenuated tip. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ longer (approximately twice as much) than famulus, with rounded apex. Famulus (ε) inflated distally. Solenidion φ of tibiae I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV very short, with rounded apex. Genu I with one solenidion σ; σ I with rounded apex, not reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III with short solenidion, with rounded apex.

Adults. Described in Klimov (1998a).

Distribution

Eastern Palearctic: Far East of Russia (Klimov 1998a; Klimov and Tolstikov 2011) and Japan (Klimov 1998a; Klimov and Tolstikov 2011; Kurosa 2000; this work).

Hosts. Boletoxenus bellicosus (Klimov 1998a; this study), Cis jacquemartii (Klimov 1998a), Boletoxenus incurvatus and Atasthalomorpha dentifrons (this work).

Remarks

The initial description indicated that setae ve in phoretic deutonymphs are not visible; however, the study of all new specimens showed their presence. We also observed that the sternum is bifurcated, but branches are very weakly-sclerotized (not simple, as originally described). In the original description, it was stated that tarsus I contains ba, but in fact that is ω₃. Also, the original work (see Klimov 1998a, Fig. 23) shows that solenidion σ I reaches the base of φ I, whereas it is clearly shorter in new specimens.

Capillaroglyphus telemachus sp. nov.

(Figs 21–24)

ZOOBANK: 764151EE-46E4-406F-ACD6-857CE54B346D

Type material

Holotype — HDN – Malaysia Sabah, Kuala Penyu district, Tiga Is., 05°34′19″N, 115°38′59″E., ex Bolitonaeus vacca (Motschusky, 1858) (Coleoptera: Tenebrionidae), 27-29 Apr 2012, A. Kompantsev (ZIN). Paratypes: 2 HDNs – same data as for holotype (ZIN).

Description

Phoretic deutonymph (Figs 21–24; measurements: Table 2). Body ovoid (Figs 21, 23A, B), 1.53 (1.46–1.54) times longer than wide, widest in sejugal region; idiosomal length 203 (205–210), width 133 (136–140) (n = 3).

Gnathosoma (Figs 22G, 24E) short (length of gnathosoma approximately 1.7 (1.6–1.7) times exceeds width), subcapitulum and palps fused, bearing short (shorter than length of subcapitulum) palpal solenidia ω, alveoli of seta h absent, anterio-lateral seta (sup) present, spiniform, posterior-lateral (a) and anterio-ventral (cm) setae absent.

Dorsum (Figs 21A, 23A,C, 24A,B). Dorsal surface of idiosoma with pattern of fine (0.8–1.0 on propodosoma, 0.4–0.6 on hysterosoma), evenly distributed pores, pores present in medial part of hysterosoma, smoothly punctate between them (Fig. 23C). Anterior margin of propodosoma with elongated rostrum, no radial pattern. Setae si approximately equal to se; both filiform, smooth or very slightly pectinate, si situated posteriorly to se. Eyespots present, without lenses, situated near bases of seta se. Sejugal furrow well developed. Prodorsal shield 68 (67, 68), hysteronotal shield 129 (128–130), ratio hysteronotal shield/prodorsal shield length = 1.9 (1.91). Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of elongated, filiform, setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), at least c₁, c₂, d₁, d₂, e₁, e₂, f₂, h₁, h₂ very slightly pectinate. All setae approximately equal in length and slightly longer or approximately equal to femur I, seta c₂ 19 (18, 19). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than or approximately equal to length of corresponding setae. Cupules ia large, situated between c₂, c_p, ip situated dorsolaterally between e₂, f₂, without distinct median spot.

Venter (Figs 21B, 23B, 24C–F). Setae vi and ve situated on ventral part of rostrum: seta vi 4.5 (4.0–5.0) times longer than minute ve, smooth. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above bases of legs I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes; distance between its end and ventrogenital shield approximately 2.6 of its length. Sternal shield closed posteriorly, posterior edge straight. Ventrogenital shield not elongated. Sternal and ventrogenital shields separated by very short distance, almost touching each other. Coxal fields III closed, its medial apodeme short. Ventral medial apodeme present. Coxal setae 1a and 4a conoidal, well-developed, 3a small, with attenuated tips, 4b filiform, short, both setae present. Seta 4a placed posteriorly to g. Genital seta g barely touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body shorter than half of attachment organ length. Anterior paired and posterior unpaired cuticular suckers of attachment organ absent; posterior paired suckers fused with attachment organ shields. Suckers ps₁ and ps₂ conoidal, well-developed; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ surround corresponding conoids, not protruding posteriorly, separated from each other. Anterior suckers (ad₃) without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, posterior to c₃. Cupules ih equal in size to im; median spot well-developed on both structures.

Legs (Figs 22A–F, 24A–C,E,F) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III; pR I–II on protrusions. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform, vF I–II on protrusions, medium length. Genual setation 2-2-0-0; setae mG I–II and cG I-II spiniform, smooth, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III-IV spiniform, smooth. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. Tarsus I with setae ra, la, f, d filiform, long, with weak terminal dilations, e spiniform, p and q spiniform, massive, with attenuated tips; seta d slightly longer than ra or la; seta s represented by alveolus; setae aa, ba and wa absent. Tarsus II similar to tarsus I except seta ba present, filiform, with widened base, situated close to ω₁. Tarsus III with setae f, d, r filiform, smooth, without terminal dilations, w with widened base, e, s, q and p spiniform with attenuated tips, s longer than q; p not protruding beyond pretarsal ambulacrum, q protruding beyond pretarsus; d smooth, shorter than leg. Tarsus IV similar to tarsus III except seta r spiniform with attenuated tip and p protruding beyond pretarsus. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter (1.1) and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ longer (1.0–1.2) than famulus, with rounded apex. Famulus (ε) inflated distally. Solenidion φ of tibiae I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III with short solenidion, with rounded apex.

Distribution

Oriental: East Malaysia.

Host

Bolitonaeus vacca.

Etymology

The new species is named after Telemachus, son of Odysseus and Penelope, and a principal character in Homer's Odyssey. Noun in apposition.

Type deposition

Holotype and paratypes are in ZIN.

Remarks

The deutonymph of the new species, like C. hermes, has an elongated rostrum, but differs from it in not having flattened dorsal setae and presence of a solenidion on genu III. The new species, like C. eumaeus, was found on Bolitonaeus vacca, but C. telemachus differs from C. eumaeus by the elongated (longer than wide) rostrum (shorter than wide in C. eumaeus), shorter solenidion ω₂ (1.0–1.2 times longer than famulus vs. two times longer in C. eumaeus) and pattern of pores present in medial part of hysterosoma (absent in C. eumaeus).

Genus Boletoglyphus Volgin, 1953

Boletoglyphus Volgin, 1953: 262; Černý and Samšińák, 1971: 508; Fain and Mahunka, 1990: 110; OConnor, 1991: 312; Bugrov, 1997: 151; Klimov, 1998a: 5; Klimov, 1998b: 1122; Klimov and Tolstikov, 2011: 257.

Fantovia Samšińák, 1957: 112 (Type species — Schwiebea boletophagi F. Turk and S. Turk, 1952, by original designation). Synonymized by Černý & Samšińák (1971).

Ellipsopus Fain and Ide, 1976: 233 (Type species — Ellipsopus ornatus Fain and Ide, 1976, by original designation). Synonymized by OConnor and Pfaffenberger (1987) and Fain and Mahunka (1990).

Lindquistia Mahunka, 1977: 69 (Type species — Lindquistia bolitotheri Mahunka, 1977 (=Boletoglyphus ornatus (Fain and Ide, 1976)), by original designation). Synonymized by Fain and Mahunka (1990).

Schwiebea (non Oudemans, 1916): F. Turk and S. Turk, 1952: 501.

Type species — Boletoglyphus cribrosus Volgin, 1953 (=Schwiebea boletophagi F. Turk and S. Turk, 1952), by original designation.

Diagnosis

Female. Body elongated. Grandjean's organ narrow, finger-shaped process. Supracoxal setae (scx) sword shaped. Setae si present, short or minute, setae se long. Setae c_1-3 alveolar, d₁ and f₂ short, filiform, d₂ absent or microsetae; ps_1-3 alveolar. Setae c_p, e₁, e₂, h_1-3 long, filiform. Posterior hysterosoma covered by opisthosomal shield; posterior portion of shield shaped as a ''visor'' over copulatory opening. Anal opening reaching posterior margin of hysterosoma and extending somewhat dorsally. Spermatheca with an inner canal. Base of spermatheca with a long transverse sclerite, with large, U-shaped sclerites of efferent ducts situated on its edges. Legs short. Setae ba I-II spiniform, short. Setae wa I-II, wF IV present. Setae hT I-II and nG III absent. Genu III with solenidion σ III present.

Male. Posterior hysterosoma with sclerotization. Para-anal suckers ad₃ present, simple, disk-like; setae ps_1-3 short setiform, ps₃ situated close to suckers ad₃. Solenidion ω₃ I long, φ IV short. Tarsus IV with suckers d and e.

Phoretic deutonymph. Gnathosoma short, subcapitulum and palps fused, gnathosomal solenidia distinctly longer than subcapitulum; setae sup filiform (spiniform in B. anticleia) or absent, setae a and cm absent. Prodorsal shield long (ratio hysteronotal shield/prodorsal shield length less than 3.0), rounded anteriorly, separate rostrum absent. Eyespots present, lenses absent. Seta ve present. Sejugal furrow well developed. Dorsal surface of idiosoma with pattern of coarse pores, dorsomedian ''ridge'' on hysterosoma present or absent. Dorsal idiosomal setae of various shapes. Posterior coxal apodemes II not touching each other medially. Attachment organ situated far from posterior margin of body; distance between them equal to or exceeding the half of attachment organ length. Cuticular suckers of attachment organ absent. Famulus cylindrical, not clavate. Setae hT I-II and nG III absent. Genu I with a single solenidion σ. Genu III with solenidion σ III absent.

Species included

Boletoglyphus boletophagi, B. extremiorientalis, B. ornatus, B. athena sp. nov., B. circe sp. nov., B. calypso sp. nov., B. penelope sp. nov., B. helena sp. nov., B. nausicaa sp. nov., B. eurycleia sp. nov., B. anticleia sp. nov.

Boletoglyphus anticleia sp. nov.

(Figs 25–27)

ZOOBANK: 9095D357-4960-4751-A45E-BECDDBA0762C

Type material

Holotype — HDN – Vietnam Đăk Ung, 04.2025 (15°28′N, 107°45′E), ex. Boletoxenus sp. (Coleoptera: Tenebrionidae), local collector (ZIN). Paratypes – 3 HDNs – same data (ZIN)

Description

Phoretic deutonymph (Figs 25–27; measurements: Table 3). Body elongate (Figs 25, 27A, B), 1.5 (1.5–1.59) times longer than wide, widest in sejugal region; idiosomal length 198 (187–211) width 125 (122–132) (n = 4).

Gnathosoma (Figs 26E, 27D) short (length of gnathosoma approximately 1.4 (1.46–1.5) times exceeds width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, spiniform, postero-lateral (a) and anteroventral (cm) setae absent.

Dorsum (Figs 25A, 27A,C). Dorsal surface of idiosoma with pattern of coarse pores surrounded by refracting ring (diameter of medial pores 1.0–2.0 on propodosoma and hysterosoma); smooth irregular punctations situated between coarse pores. Anterior edge of propodosoma rounded, without radial pattern (Fig. 27D). Setae si about 1.0–1.2 times longer than se; both long, filiform to slightly flattened, smooth, si and se situated on the same transverse level, longer than femur I, distance si–si shorter than length of corresponding si setae. Eyespots present, without lenses, situated near bases of seta se. Sejugal furrow well developed. Propodosomal shield 67 (66–76), hysteronotal shield 129 (119–133), ratio hysteronotal shield/prodorsal shield length = 1.92 (1.75–1.8). Median ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs long, filiform, smooth setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae c₁, c₂, d₁, d₂, e₁, h₁, h₃ approximately same length, h₂, c_p, e₂, and f₂ slightly shorter. All setae (except c_p, e₂ and f₂) longer than femur I, seta c₂ 24. Alveoli of idiosomal setae (except c₃ and h₃) enlarged. Distances d₁–d₁, e₁–e₁ distinctly shorter than length of corresponding setae, distance h₂–h₂ slightly longer than length of h₂. Cupules ia situated between c₂, c_p, ip small, without median spot.

Venter (Figs 25B, 27B, D–F). Setae vi and ve situated on ventral part of anterior edge of propodosoma; seta vi 4.4 (3.7–4.3) times longer than ve, smooth; seta ve smooth, weakly spiniform. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above bases of legs I. Coxal fields sclerotized, punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum not bifurcated, touching anterior coxal apodeme II, ending far from free ends of anterior coxal apodemes II; distance between its end and ventrogenital shield slightly shorter than its length. Sternal shield not enclosed posteriorly. Ventrogenital shield elongated. Sternal and ventrogenital shields separated by small distance, 2.8 (4.2–5.4). Coxal fields III closed. Ventral medial apodeme present. Coxal setae 1a large spiniform, 3a and 4a small spiniform, widened in middle length and with attenuated tips (Figs 27D–F), 4b filiform, longer than other coxal setae, both 4b setae present. Seta 4a anterior to g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ (Fig. 27F) and posterior edge of body equal or more the half of attachment organ length. Cuticular suckers of attachment organ absent. Setae ps₁ and ps₂ conoidal; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ large, surrounding corresponding conoids, slightly protruding posteriorly, not touching each other. Anterior suckers (ad₃) round, with transparent marginal edges. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, at the level of setae c₃. Cupules ih equal in size to im; median spot well-developed on both structures.

Legs (Figs 26A–D, 27D, F) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III, on short protrusions. Femoral setation 1-1-0-0; setae vF I–II long, filiform, on short protrusions. Genual setation 2-2-0-0; setae mG I–II and cG I–II spiniform, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I–II and kT III-IV spiniform. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsus I with setae ra, la, d, f filiform, with slightly widened basal part, e with large widened basal part; seta d longer than ra or la; seta s represented by alveolus; p and q spiniform, not protruding beyond pretarsus; setae aa, wa and ba absent. Tarsus II similar to tarsus I except seta ba present, filiform, situated close to ω₁. Tarsi III and IV with setae w, r, e, f and d filiform, smooth, s, q and p spiniform, q and p subequal, spiniform, barely, extending beyond pretarsal ambulacrum; seta d III shorter than leg, d IV distinctly longer than leg. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ present. Famulus (ε) solenidion-shaped, with rounded tip, slightly shorter than ω₂. Solenidion φ of tibia I elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV very short, with rounded apex. Genu I with one solenidion σ; σ I with rounded apex, reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion.

Distribution

Vietnam.

Host

Bolitoxenus sp.

Etymology

The new species is named after Anticleia, the mother of Odysseus and wife of Laertes in the Odyssey. She appears in the Underworld episode of the epic, where she informs Odysseus about events in Ithaca during his absence and dies of grief for her son. Noun in apposition.

Remarks

This species exhibits a combination of morphological features similar to those of Capillaroglyphus: a spiniform seta sup; short, spiniform proral setae p and q on legs I–II that do not protrude past the pretarsus; and short projections on femora and trochanters I–II. A unique feature of this species is the presence of a spiniform seta 1a, which has not been observed in any other species within the tribe.

Boletoglyphus athena sp. nov.

(Figs 28, 29)

ZOOBANK: 4A3B6568-BE0C-4E6F-A9CA-9DC6330119B3

Type material

Holotype — HDN – Philippines Luzon Is., ''Baltalason'' (possibly Balbalasang-Balbalan National Park, Kalinga Prov. 17°27′N, 121°09′E), ex Byrsax tuberculatus Gravely, 1915 (Coleoptera: Tenebrionidae), no date and collector name, (FMNH) BMOC 96-1203-006 (FMNH). Paratypes: 40 HDNs – same data (FMNH, UMMZ); 30 HDNs – same locality and host, no date, F. C. Fletcher, (CUIC) BMOC 79-0301-006 (CUIC, UMMZ).

Additional material — 3 HDNs – data same as for holotype (UMMZ).

Description

Phoretic deutonymph (Figs 28, 29; measurements: Table 3). Body elongate (Figs 28), 2.31 (1.88–2.33) times longer than wide, widest in sejugal region; idiosomal length 124 (116–129), width 54 (53–62) (n = 10).

Gnathosoma (Fig. 29A) short (length of gnathosoma approximately 0.6 (0.9–1.1) times width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h present, anterolateral seta (sup) present, longer than length of gnathosoma, filiform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Fig. 28B). Dorsal surface of idiosoma with pattern of coarse pores (diameter of medial pores 1.0–2.0), not approaching the bases of setae d₁, e₁ and h₁ laterally, uniformly punctate between coarse pores. Anterior margin of propodosoma rounded, with radial pattern. Setae si about 1.5 times longer than se, lanceolate, finely serrate, long (longer than femur I); se filiform, slightly longer than femur I; si situated posteriorly to se. Eyespots present, without lenses, situated lateral to bases of seta se. Sejugal furrow well developed. Prodorsal shield 44 (41–45), hysteronotal shield 80 (73–84), ratio hysteronotal shield/prodorsal shield length = 1.82 (1.70–1.98). Dorsomedian ''ridge'' on hysterosoma present. Hysterosoma with 11 pairs of setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae h₃ slightly longer than c_p, other setae distinct shorter than c_p. Setae e₁ and h₁ lanceolate, finely serrate, long (e₁ broken on holotype); other setae simple, filiform. Several opisthosomal setae (c_p, e₁, h₁, h₃) are longer than femur I, seta c₂ short 7 (6–8). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁ and h₂-h₂ longer than length of corresponding setae; distance e₁-e₁ shorter than length of e₁. Cupules ia large, situated openly, not in small hollow, ip small, without distinct median spot.

Venter (Fig. 28A). Setae vi and ve smooth, situated on ventral part of anterior margin of propodosoma; seta vi 3.5 (2.3–3.1) times longer than ve. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum not bifurcated, not touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes II; distance between its end and ventrogenital shield approximately equal to its length. Sternal shield not closed posteriorly. Ventrogenital shield elongated. Sternal and ventrogenital shields separated by large distance (10 (9–15)). Coxal fields III closed. Ventral medial apodeme present. Coxal setae 1a, 3a and 4a vestigial, with attenuated or rounded tips, 4b filiform, longer than other coxal setae, both 4b setae present. Seta 4a situated anteriorly to g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body equal or more than attachment organ length. Cuticular suckers of attachment organ absent. Suckers ps₁ and ps₂ conoidal; ps₂ shifted posteriorly and medially, situated close to ps₁. Sclerites of ps₁ and ps₂ large, rounded, touching each other. Anterior suckers (ad₃) round, without transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, slightly posterior to setae c₃. Cupules im, ih equal in size; median spot well-developed on both structures.

Legs (Figs 29B–M) short, all segments free. Trochanters I–III each with long, filiform setae, pR I–II, sR III. Femoral setation 1-1-0-0; setae vF I–II long, filiform, wF IV absent. Genual setation 2-2-0-0; setae mG I–II slightly dilated in basal part, filiform, mG II longer than mG I, cG I and II filiform, short, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III–IV filiform. Tarsal setation 7-8-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsi I and II with setae ra, la, p, q, d, f, e filiform, long; seta d I slightly longer than ra and la; seta s represented by alveolus; setae aa, wa and ba absent. Tarsi III and IV with setae w, r, s, p, q, e, f, s and d filiform, smooth, q III shorter than p III, q III not protruding beyond pretarsus; q and p IV approximately of equal length, protruding beyond pretarsus; setae d shorter than legs. Solenidion ω₁ on tarsus I cylindrical, with slightly clavate apex, without or sometimes with membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁ I, with rounded apex, situated close to ω₁. Solenidion ω₂ slightly shorter than famulus, cylindrical, with slightly clavate apex. Famulus (ε) solenidion-like, with rounded tip. Solenidion ω₁ on tarsus II cylindrical, with slightly clavate apex. Solenidia of tibiae I–III elongate, tapering; φ I and III longer than tarsus with empodium; φ II shorter than tarsus II without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, not reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion (spot present).

Distribution

Oriental: Philippines.

Hosts

Byrsax tuberculatus.

Etymology

Boletoglyphus athena is named after Athena, the ancient Greek goddess who aided Odysseus in his adventures. Noun in apposition.

Type deposition

Holotype and paratypes are in FMNH, additional paratypes in CUIC, UMMZ.

Remarks

The host beetle also carried Neochelacheles mendicus Bochkov & OConnor, 2004 (Cheyletidae), which is a likely a predator of B. athena.

Boletoglyphus boletophagi (F. Turk and S. Turk, 1952)

Schwiebea boletophagi F. Turk and S. Turk, 1952: 501, Figs 33, 34; Hughes, 1957: 300.

Fantovia boletophagi: Samšiňák, 1957: 112, Fig. 113.

Boletoglyphus boletophagi: Klimov, 1998a: 1123, Fig. 1; Klimov, 1998b: 1123, Fig.1; Haitlinger, 2004: 15; Klimov and Tolstikov, 2011: 257; Haitlinger and Lupicki, 2012: 39; Penttinen et al. 2013: 11, Fig.2f; Khaustov et al, 2018: 30, Fig. 19c, d.

Boletoglyphus cribrosus Volgin, 1953: 263, Figs A-C; Bugrov, 1997: 151; (synonymized by Černý and Samšińák, 1971).

(Figs 30–36)

Specimens examined

Two females, 9 HDNs – Scotland, Highlands Dist., Glen Affric, 57°14′09″N, 005°09′12″W, ex Fomes fomentarius (L.) Fr. 1849 (Polyporales, Polyporaceae), 5 Sep 1982, B. M. OConnor, BMOC 82-0905-001, (Topotypes) (UMMZ); 1 female, 8 HDNs, 1 protonymph – Scotland, Highlands Dist., 4.8km S Invergarry (57°02′N, 004°47′W) ex Fomes fomentarius, 5 Sep 1982, B.M. OConnor, BMOC 82-0905-002, (UMMZ); 28 HDNs – Russia, Irkutsk Province, Bratsk, 56°07′N, 101°36′E, ex decomposing hymenophore of a polypore from fallen birch tree, 15 Oct 1995, P.B. Klimov (IBSS); 15 HDNs – Russia, Irkutsk Province, Bratsk, ex decomposing hymenophore of a polypore from fallen birch tree and ex Cis lineatocribratus Mellié, 1848 (Coleoptera, Ciidae), 15 Oct 1995, P.B. Klimov (IBSS); 5 females, 3 males, 43 HDNs – Russia, Irkutsk Province, Bratsk District, Podvyezdnoe (56°17′0″N, 102°3′0″E), ex Bolitophagus reticulatus (Linnaeus, 1767) (Coleoptera: Tenebrionidae), 15 Aug 1996, P.B. Klimov, BMOC 98-0110-003 (IBSS) (Klimov 1998b); 5 females, 10 HDNs – Russia, Smolensk Region, Pronino village (54°48′30″N, 031°52′31″E), on old Fomes fomentarius, 10 Apr 2025, V.B. Kolesnikov (ZIN); 12 HDNs – Russia, Voronezh Region, Ramnosky District, Aidarovo village, 51°55′29″N, 39°17′55″E, ex Bolitophagus reticulatus, 15 Jul 2023, V.B. Kolesnikov (ZIN); 1 HDN – Croatia (exact location unknown), ex Bolitophagus interruptus Illiger, 1800 (ZIN); 5 HDNs – Croatia, Veledit Dist., Paklenica, 45°19′N, 017°02′E, ex Bolitophagus reticulatus, 4 Jun 1916, Novak coll. (CUIC); 4 HDNs – Azerbaijan, Talysh, Bilasuvar, 39°27′30″N, 048°32′42″E, 800-1000 m, ex Bolitophagus reticulatus, 7 Jun 1936 (ZIN); 19 HDNs – Czech Republic, Vychodocesky Region, Kostelec nad Orlici, 50°08′N, 016°14′E, ex Bolitophagus reticulatus, May 1963, Marek coll., BMOC 79-1011-002 (EMEC); 19 HDNs – Slovakia, Vychodoslovensky Dept., Vyhorlat Mts. 48°50′N, 022°15′E, 7 Jun 1964, O. Marek, BMOC 79-1011-002 (EMEC); 19 HDNs – Crimea, Yalta municip. Ai-Petri mountain, 44°27′04″N, 034°03′10″E, ex Bolitophagus reticulatus, no date, BMOC 96-1203-001 (FMNH); 32 HDNs – Romania, Caras-Severin county, Anina, 45°05′N, 021°51′E, ex Bolitophagus reticulatus, no collector, BMOC 96-1203-007 (FMNH); 5 females, 2 males, 7 HDNs, 12 protonymphs – Sweden Uppsala Dist., Angsbacka, 8km WNW Uppsala, 59°52′N, 017°38′W, ex Fomes fomentarius, 9 Sep 1997, T. Nilsson, BMOC 97-0920-001 (UMMZ); 5 HDN - same data, ex Bolitophagus reticulatus, BMOC 97-0920-002 (UMMZ).

Redescription

Female (Figs 30–33). Described in Klimov (1998a). Idiosoma elongate (Figs 30, 32), 580–600 x 190–200 (n = 6), 3.0–3.1 times longer than wide. Idiosomal cuticle smooth.

Gnathosoma. Chelicerae massive, with one seta (Fig. 33I). Subcapitular setae (h), palp tibial setae (a), lateral dorsal palp tibial setae (sup), dorsal palp tarsal setae (cm) filiform; h longer than other setae; supracoxal seta elcp present, needle-shaped; terminal palp tarsal solenidion ω short; external part of terminal eupathidium ul″ dome-shaped; terminal eupathidium ul′ minute, rounded (Fig. 31F).

Prodorsal shield 90–105 long, 80–90 wide; 1.1 times longer than wide; with setae vi situated at anterior part of shield; without anterior incisions; midlateral incisions (alveolar ve) present. Prodorsal shield rounded posteriorly. Prodorsal shield smoothly punctate (Figs 30A, 33A). Alveoli of vi well separated (distance between them distinctly greater than their diameter). Grandjean's organ (GO) a narrow, finger-like process, bifurcated apically (Figs 31G, 33B,C). Supracoxal seta scx finely barbed, sword-shaped, widened and flattened, tapering at tip (Figs 31G, 33B,C). Idiosomal setae (vi, se, si, c_p, d₁, e₁, e₂, f₂, h₁, h₂, h₃) smooth, filiform and short, si, c_p, d₁ and f₂ shorter than other, se longer than others. Seta c_1-3 represented by alveoli. Four pairs of fundamental cupules (ia, im, ip and ih) present, ip and ih close together. Posterior part of opisthosoma with sclerotized shield (Figs 30, 33F).

Ventral idiosoma with three pairs of coxal setae (1a, 3a and 4a) and one pair of genital setae (g) (Fig. 30B), seta 4b not seen. Shape of coxal sclerites as in Fig. 30B, sejugal apodeme absent – present only as isolated sclerites. Genital region situated between coxal fields III and IV; genital valves form an inverted Y; epigynal and medial apodemes well-developed (Figs 30B, 33D). Genital papillae rounded, blunt. Anal opening ventral, in posterior part of idiosoma. Copulatory tube not developed; copulatory opening situated under rounded edge of opisthosomal shield. Inseminatory canal (ic) of spermatheca short, noticeably widened at base. Base of spermatheca (hs) in the form of transverse plate (Figs 31H, 33E). Sac of spermatheca rounded, folded. Inside the sac there is a soft canal-fold from the inseminatory canal to the top of sac. Paired efferent ducts (ed) short with short stem.

Legs (Figs 31A–E, 33G–L) short, all segments free. Trochanters I–III each with short, filiform seta, pR I–II, sR III; trochanter IV without setae. Femoral setation 1-1-0-1; setae vF I–II long, filiform, wF IV short, filiform. Genual setation 2-2-0-0; setae mG and cG I–II short, slightly spiniform; seta nG III absent. Tibial setation 1-1-1-1; setae gT I–II and kT III–IV elongate, somewhat spiniform, setae hT I-II absent. Tarsal setation 12-12-10-10; pretarsi consist of hooked empodial claws attached to short paired condylophores; membranous ambulacrum very small. Tarsi I and II with setae ra, la, f and d filiform, ba, wa, s, e, u, v spiniform, p and q represented by small triangular rudiments, setae ba shorter than ω₂. Tarsi III and IV with setae f, d, r and w filiform, s, e, u and v spiniform, p and q represented by small triangular rudiments. Solenidion ω₁ on tarsus I cylindrical, with clavate apex, not bent; solenidion ω₁ on tarsus II simple, cylindrical, with clavate apex, not bent, shorter and wider than ω₁ on tarsus I. Solenidion ω₂ on tarsus I shorter than ω₁, cylindrical, with rounded apex, slightly widened distally and slightly curved, situated slightly distal to ω₁. Solenidion ω₃ on tarsus I cylindrical, with slightly tapering tip, slightly longer than ω_1, positioned dorso-apically. Famulus (ε) of tarsus I wide, with rounded apex, slightly widened distally. Solenidia φ of tibiae I–III elongate, tapering, extending well beyond apices of respective tarsi with ambulacra; solenidion φ IV shorter, shorter than tarsus IV (without ambulacra). Solenidia σ″ and σ′ on genu I elongate, σ′ longer than σ″, σ′ reaches bases of φ I. Solenidion σ on genua II and III more than 6–7 times longer than wide, with rounded tip.

Male. Described in Klimov (1998a). Idiosoma elongate, 449–453 x 350–360 (n = 3), 0.8 times longer than wide. Gnathosoma, prodorsal shield and setation of idiosoma as in female, except short setiform ps_1, ps₂ and ps₃ (close to anterior side of ad₃) para-anal suckers ad₃ present and rounded. Aedeagus thick, slightly protruding beyond anterior edge of supporting sclerite.

Legs I-IV as in female, except setae d and e as suckers on tarsus IV.

Phoretic deutonymph (Figs 34–36; measurements: Table 3). Body elongate (Figs 34, 36A, B), 1.5–1.61 times longer than wide, widest in sejugal region; idiosomal length 290–315 width 187–204 (n = 6).

Gnathosoma (Figs 35G, 36D) short (length of gnathosoma approximately 1.28–1.58 times width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h present, anterolateral seta (sup) present, filiform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Figs 34A, 36A, C). Dorsal surface of idiosoma with a pattern of coarse pores (diameter of medial pores 3.0–4.5), and with irregular smooth punctations between them. Anterior margin of propodosoma rounded, without radial pattern. Setae si and se slightly dilated, smooth; si and se equal in length, se longer than femur I; si and se situated on the same transverse level. Eyespots present, without lenses, situated near bases of seta se. Sejugal furrow well developed. Prodorsal shield 104–120, hysteronotal shield 182–198, ratio hysteronotal shield/prodorsal shield length = 1.57–1.75. Dorsomedian ''ridge'' on hysterosoma absent. Hysterosoma with 11 pairs of simple, filiform (h₁ slightly dilated) setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae h₃ distinctly longer than others. All setae (except h₃) shorter than femur I (c_p approximately equal femur I), seta c₂ 14–16. Bases of idiosomal setae situated openly on cuticle. Distances d₁–d₁, e₁–e₁, and h₂–h₂ longer than length of corresponding setae. Cupules ia large, situated between c₂, c_p, ip small, with median spot.

Venter (Figs 34B, 36B,E,I). Setae vi and ve smooth, situated on anteroventral margin of propodosoma; seta vi 4.4–5.5 times longer than ve. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above bases of legs I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, ends far from free ends of anterior coxal apodeme II; branches well-sclerotized, small, touching anterior coxal apodeme II; distance between sternum end and ventrogenital shield approximately 0.7 of sternum's length. Anterior coxal apodemes II not touching each other after connection with sternum. Sternal shield open posteriorly, posterior edge concave medially. Ventrogenital shield elongated. Sternal and ventrogenital shields separated by short distance (7–11). Coxal fields III closed. Ventral medial apodeme present. Coxal setae 1a, 3a and 4a short, filiform, 4b filiform, longer than other coxal setae, both 4b setae present (in one specimen both 4b setae were absent). Seta 4a situated anteriorly g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ (Fig. 34E) and posterior edge of body equal or more than half of attachment organ length. Cuticular suckers of attachment organ absent. Setae ps₁ and ps₂ conoidal; ps₂ shifted posteriorly and medially, separated from ps₁. Sclerites of ps₁ and ps₂ large, surrounding corresponding conoids, protruding posteriorly, not touching each other. Anterior suckers (ad₃) round, with transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, at level of setae c₃. Cupules ih equal in size to im; median spot well-developed on both structures.

Legs (Figs 35A–F, 36F–H) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III. Femoral setation 1-1-0-1; setae vF I–II and wF IV long, filiform. Genual setation 2-2-0-0; setae mG I–II filiform, mG II longer than mG I, cG I spiniform, cG II weakly spiniform, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I spiniform, straight, smooth; setae gT II spiniform, curved, serrated; setae kT III–IV weakly spiniform. Tarsal setation 8-9-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal dilations. Tarsus I with setae ra, la, p, q, d, f, e, wa filiform, long; seta d shorter than ra or la; seta s represented by alveolus; setae aa and ba absent. Tarsus II similar to tarsus I except seta ba present, filiform, situated close to ω₁. Tarsi III and IV with setae w, r, s, p, q, e, f and d filiform, smooth, q and p protruding beyond pretarsus, approximately of equal length; s slightly dilated in basal part; seta d longer than leg. Solenidia ω₁ on tarsi I–II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁, with rounded apex, situated close to ω₁. Solenidion ω₂ present, slightly shorter than famulus. Famulus (ε) solenidion-like, with rounded tip. Solenidia of tibiae I–III elongate, tapering; φ I longer than tarsus I with empodium; φ II shorter than tarsus II without empodium; φ III longer than tarsus III without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, not reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion.

Distribution

Western Palearctic: United Kingdom (Scotland) (F. Turk et S. Turk 1952, this study), Czech Republic (this study), Slovakia (Samšiňák 1957), Poland (Haitlinger 2004; Haitlinger and Lupicki 2012), Romania (this study), Sweden (this study), Finland (Penttinen et al. 2013), Azerbaijan (this study), Siberia, and the European part of Russia (Bugrov 1997; Klimov 1998b; Volgin 1953, Khaustov et al. 2018; this study).

Hosts

Bolitophagus reticulatus (F. Turk et S. Turk, 1952; Volgin 1953; Samšiňák 1957; Bugrov 1997; Klimov 1998b; Haitlinger 2004; Haitlinger and Lupicki 2012; this study), Bolitophagus interruptus (this study), Cis lineatocribratus (Klimov 1998b); Ips typographus (Linnaeus, 1758) (Penttinen et al. 2013; Khaustov et al. 2018). The fungal host is Fomes fomentarius, a widespread polypore associated with Betula species and other tree species. Examination of many sporocarps of this fungus in North America did not yield any Boletoglyphus.

Remarks

Adult stages of this species were first described by Klimov (1998a). Given that females of Boletoglyphus were only known for B. boletophagi and B. extremiorientalis, our comparisons are based on female morphology of these two taxa. Females of Boletoglyphus boletophagi differ from those of B. extremiorientalis as follows: (i) posterior part of the prodorsal shield is triangulated, narrowed posteriorly, without protruding rounded lateral sides in B. boletophagi (widened, with round medial and round lateral protrusions in B. extremiorientalis); (ii) setae si are short in B. boletophagi (vestigial in B. extremiorientalis); and (iii) the inseminatory canal of the spermatheca is narrowed near the base of spermatheca in B. boletophagi (diameter uniform along its entire length in B. extremiorientalis).

Phoretic deutonymphs are similar to those of B. extremiorientalis but differ by the following two features: (i) setae se and si, which are inserted at the same transverse level in B. boletophagi (si situated posterior to se in B. extremiorientalis); and (ii) the common sclerite of the posterior median suckers, which extends beyond the posterior margin of the attachment organ in B. boletophagi (not extending in B. extremiorientalis).

Boletoglyphus calypso sp. nov.

(Figs 37–39)

ZOOBANK: a4DA77C1-E9FB-4164-B3E8-D9072305722E

Type material

Holotype — HDN – Philippines, Catanduanes, 8.5 km W Gigmoto, 250m., ex Polyporales, Polyporaceae, 13°47′N, 124°19′E, 27 Feb – 2 Mar 1988, S. M. Goodman, BMOC 88-0801-001 (UMMZ). Paratypes: 2 HDNs – same data (UMMZ); 58 HDNs – Philippines, Luzon Is., Nueva Vizcaya prov., Imugan, 16°09′32″N, 120°54′10″E, ex Boletoxenus ditylus Gebein, 1921 (Coleoptera, Tenebrionidae), F. C. Fletcher, BMOC 79-0301-005 (CUIC, UMMZ); 8 HDNs – Philippines Negros Oriental, Lake Balinsasayao, 3 km N 14 km W Dumaguete, °9°22′N, 123°11′E, 980 m., ex Boletoxenus timmi Gebien, 1922 (Coleoptera: Tenebrionidae), 9 Nov 1982, P. D. Heideman #1103, BMOC 83-1800-044 (UMMZ).

Additional material — 4 HDN – Philippines Luzon Is., Nueva Viscaya, Santa Fe municipality, 2015, ex Boletoxenus ditylus (Coleoptera: Tenebrionidae), local collector (ZIN).

Description

Phoretic deutonymph (Figs 37–39; measurements: Table 3). Body elongate (Figs 37, 39A), 2.24 (1.88–2.19) times longer than wide, widest in sejugal region; idiosomal length 190 (206–232) width 85 (95–124) (n = 10).

Gnathosoma (Figs 38A, 39E) short (length of gnathosoma approximately 0.8–1.0 times the width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, shorter than length of gnathosoma, filiform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Figs 37A, 39A–C). Dorsal surface of idiosoma with pattern of coarse pores (diameter of medial pores 1.0–2.0), except on dorsomedian ''ridge″, smoothly punctate between coarse pores. Anterior margin of propodosoma rounded, with radial pattern (Fig. 39D). Setae si approximately equal to se, both lanceolate, barbed, situated on same transverse level, longer than femur I. Eyespots present, without lenses, lateral to bases of seta se. Sejugal furrow well developed. Prodorsal shield 70 (71–81), hysteronotal shield 120 (134–151), ratio hysteronotal shield/prodorsal shield length = 1.7 (1.82–1.91). Dorsomedian''ridge'' on hysterosoma present, not bifurcated, thin ridges present between h₁ and h₂. Hysterosoma with 11 pairs setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae h₃ distinctly longer than others. Setae h₁ lanceolate, barbed (Figs 37L, 30C); c₁, c₂, d₁, d₂, e₁, f₂, h₂ slightly flattened, c₂, h₂ and sometimes d₁, e₁ finely serrate, short, c_p, e₂ and h₃ filiform. Several opisthosomal setae (c_p, h₁, h₃) longer than femur I, seta c₂ 7 (7–11). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than length of corresponding setae. Cupules ia large, situated in small hollow between c₂, c_p, not seen in all specimens; ip not observed.

Venter (Figs 37B, 39D, F). Setae vi and ve smooth, situated on ventral part of anterior margin of propodosoma; seta vi 3.9 (3.5–4.1) times longer than ve. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above bases of legs I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes II (14-18); distance between its end and ventrogenital shield approximately equal to its length. Sternal shield open posteriorly. Ventrogenital shield elongated. Sternal and ventrogenital shields separated by small distance (8 (8–14)). Coxal fields III closed. Ventral medial apodeme present. Coxal setae 1a, 3a and 4a vestigial very small, with attenuated or rounded tips, 4b filiform, longer than other coxal setae, both 4b setae present. Seta 4a situated anteriorly to g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ and posterior edge of body equal or more than attachment organ length. Cuticular suckers of attachment organ absent. Setae ps₁ and ps₂ conoidal; ps₂ shifted posteriorly and medially, situated close to ps₁ in transverse row. Sclerites of ps₁ and ps₂ large, rounded, touching each other. Anterior suckers (ad₃) round, with wide transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, slightly posterior to setae c₃. Cupules ih size equal to im; median spot well-developed on both structures.

Legs (Figs 38A–M) short, all segments free. Trochanters I–III each with long, filiform seta, pR I–II, sR III. Femoral setation 1-1-0-0; setae vF I–II long, filiform, wF IV absent. Genual setation 2-2-0-0; setae mG I–II filiform, mG II longer than mG I, cG I and II spiniform, short, seta nG III absent. Tibial setation 1-1-1-1; setae hT I–II absent; setae gT I-II and kT III–IV filiform. Tarsal setation 8-9-8-8. All pretarsi consisting of hooked empodial claws (without tooth) attached to short paired condylophores. All tarsal setae without terminal widenings. Tarsus I with setae ra, la, p, q, d, f, e, wa filiform, long; seta d longer than ra or la; seta s represented by alveolus; setae aa and ba absent. Tarsus II similar to tarsus I except seta ba present, filiform, situated close to ω₁. Tarsi III and IV with setae w, r, s, p, q, e, f, s and d filiform, smooth, q III distinctly shorter than p III, q III not protruding beyond pretarsus; q and p IV approximately of equal length, protruding beyond pretarsus; setae d longer than legs. Solenidia ω₁ on tarsi I and II cylindrical, with slightly clavate apices, without membranous appendage; ω₃ on tarsus I slightly shorter and thinner than ω₁ I, with rounded apex, situated close to ω₁. Solenidion ω₂ slightly shorter than famulus, cylindrical, with slightly clavate apices. Famulus (ε) solenidion-like, with rounded tip. Solenidia φ of tibiae I–III elongate, tapering; φ I and III longer than tarsus with empodium; φ II shorter than tarsus II without empodium; φ IV short, with rounded apex. Genu I with one solenidion; σ I with rounded apex, not reaching base of φ I. Solenidion σ II short, with rounded apex. Genu III without solenidion.

Distribution

Oriental: Philippines.

Hosts

Boletoxenus ditylus and B. timmi.

Morphological variation and abnormality

One mG I is absent in holotype. One ps₂ and accompanying sclerite absent in one specimen. Alveoli of ad₂ and ad₃ are separated or fused on median suckers of attachment organ.

Etymology

The new species is named after Calypso, a nymph of Ogygia, where Odysseus, after being shipwrecked, spent seven years in her company. Noun in apposition.

Type deposition

Holotype and paratypes are in UMMZ.

Remarks

Boletoglyphus calypso is very similar to B. circe but differs from it in the shape of the dorsomedian ''ridge'' on the hysterosoma (not bifurcated to level of h₁ vs. distinct bifurcated to level of h₁ in B. circe) and lanceolate seta h₁ (vs. slightly flattened and short in B. circe).

Boletoglyphus circe sp. nov.

(Figs 40–43)

ZOOBANK: fAD32FC7-CFD7-472C-8827-87A06BD20741

Type material

Holotype — HDN – Malaysia, Sabah, Kinabatangan Dist., SE Dewhurst Bay 05°35′N 118°35′E, ex Byrsax sp. (Coleoptera, Tenebrionidae), 6-14 May 1950, R. F. Inger & D. D. Davis BMOC 96-1203-004 (FMNH). Paratypes: 30 HDNs – same data (FMNH, UMMZ); 32 HDNs – Singapore, 01°17′N, 103°50′E, ex Boletoxenus gibber Motschulsky, 1858 (Coleoptera: Tenebrionidae), 1889, Wallace, (F. Bates coll. 81-19), BMOC 82-1221-002 (NHMUK, UMMZ); 29 HDNs – Singapore, ex Boletoxenus spectabilis (Coleoptera, Tenebrionidae), Atkinson coll. 92-3, no date and collector name, BMOC 82-1221-003 (NHMUK, UMMZ).

Additional material — 3 HDNs – data as for the holotype (UMMZ); 1 HDN – Singapore, ex B. spectabilis, Atkinson coll. 92-3, no date and collector name, (NHMUK), BMOC 82-1221-003 (UMMZ); 12 HDNs – Malaysia, Perak, Batang Patang, ex Byrsax sp., 04°22′09″N, 101°20′00″E, 18 Mar 2019, M. Endo (ZIN); 9 HDN's – Philippines, Palawan, Roxas, Magara Barangay, 10°11′57″N, 119°09′24″E, ex Boletoxenus sp. (Coleoptera, Tenebrionidae), N. Layron, Jan 2024, BMOC 24-0402-001 (UMMZ).

Description

Phoretic deutonymph (Figs 40–43; measurements: Table 3). Body elongate (Figs 40, 42), 2.14 (1.88–2.09) times longer than wide, widest in sejugal region; idiosomal length 214 (195–223) width 100 (100–118) (n = 10).

Gnathosoma (Figs 41L, 43C) short (length of gnathosoma approximately 1.0 (0.8–1.0) times width), subcapitulum and palps fused, bearing palpal solenidia ω, alveoli of seta h absent, anterolateral seta (sup) present, longer than length of gnathosoma, filiform, posterolateral (a) and anteroventral (cm) setae absent.

Dorsum (Figs 40A, 42A, 43A,B). Dorsal surface of idiosoma with pattern of coarse pores (diameter of medial pores 1.0–2.0), except on dorsomedian ''ridge″, smoothly punctate between coarse pores. Anterior margin of propodosoma rounded, with radial pattern. Setae si length approximately equal to se, both lanceolate, barbed, situated on the same transverse level, longer than femur I. Eyespots present, without lenses, lateral to bases of seta se. Sejugal furrow well developed. Prodorsal shield 76 (72–81), hysteronotal shield 138 (123–143), ratio hysteronotal shield/prodorsal shield length = 1.82 (1.71–1.82). Dorsomedian''ridge'' on hysterosoma present, bifurcated posterior to e₁, two lateral ''ridges'' also present on hysterosoma (Figs 40A, 42A). Hysterosoma with 11 pairs of setae on hysteronotal shield (c₁, c₂, c_p, d₁, d₂, e₁, e₂, f₂, h₁, h₂, h₃), setae h₃ distinctly longer than others. Setae c₁, c₂, d₁, d₂, e₁, f₂, h₁, h₂ slightly flattened, c₂, h₂ and sometimes d₁, h₁ finely serrate, short, c_p, e₂ and h₃ filiform (setae e₁ and h₁ broken on holotype. All setae (except c_p and h₃) shorter than femur I, seta c₂ 12 (10–12). Bases of idiosomal setae situated openly on cuticle. Distances d₁-d₁, e₁-e₁ and h₂-h₂ longer than length of corresponding setae. Cupules ia large, situated in small hollow between c₂, c_p, difficult to see in some specimens, ip small, without distinct median spot.

Venter (Figs 40B–M, 42B, 43C,D). Setae vi and ve smooth, situated on anteroventral propodosoma; seta vi 3.7 (3.3–4.2) times longer than ve. Supracoxal setae of legs I (scx) filiform, with slightly extended base, positioned ventrally above bases of legs I. Coxal fields sclerotized, smoothly punctate. Anterior apodemes of coxal fields I fused forming sternum; sternum bifurcated, touching anterior coxal apodeme II, ends far from free ends of anterior coxal apodemes II (14-18); distance between its end and ventrogenital shield more than its length. Sternal shield open posteriorly. Ventrogenital shield elongated. Sternal and ventrogenital shields separated by small distance (4.5 (5.5–10)). Coxal fields III closed. Ventral medial apodeme present. Coxal setae 1a, 3a and 4a, very small, with attenuated or rounded tips, 4b filiform, longer than other coxal setae, both 4b setae present. Seta 4a situated anteriorly to g. Genital seta g touching edge of ventrogenital shield. Distance between attachment organ (Fig. 43D) and posterior edge of body equal or more than attachment organ length. Cuticular suckers of attachment organ absent. Setae ps₁ and ps₂ conoidal; ps₂ shifted posteriorly and medially, situated close to ps₁ in transverse row. Sclerites of ps₁ and ps₂ large, rounded, touching each other. Anterior suckers (ad₃) round, with wide transparent marginal edge. Median suckers (ad₁₊₂) distinctly larger, with paired vestigial alveoli (situated on common sclerite); pair of small spots anterolateral to median suckers (ps₃). Anus situated between anterior suckers (ad₃). Opisthonotal gland openings (gla) situated ventrally on hysteronotal shield, slightly posterior to setae c₃. Cupules ih equal in size to im; median spot well-developed on both structures.