# A new feather mite species of the genus Ingrassia
## (Acariformes: Xolalgidae) from the Jacksnipe
## Lymnocryptes minimus (Charadriiformes: Scolopacidae)
## in Faroe Islands

Sergey V Mironov, Simon Haarder

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► To cite this version:

Sergey V Mironov, Simon Haarder. A new feather mite species of the genus Ingrassia (Acariformes: Xolalgidae) from the Jacksnipe Lymnocryptes minimus (Charadriiformes: Scolopacidae) in Faroe Islands. Acarologia, 2025, 65 (4), pp.1018-1031. 10.24349/xegi-38o4. hal-05387033

HAL Id: hal-05387033

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Acarologia

Open Science
in Acarology

# A new feather mite species of the genus *Ingrassia* (Acariformes: Xolalgidae) from the Jacksnipe *Lymnocryptes minimus* (Charadriiformes: Scolopacidae) in Faroe Islands

Sergey V Mironov IDª, Simon Haarder IDᵇ

## Original research

### ABSTRACT

A new feather mite species *Ingrassia lymnocrypti* sp. n. is described from the Jacksnipe, *Lymnocryptes minimus* (Brünnich), collected in Faroe Islands. The new species is close to *I. limnodromi* Vasyukova and Mironov, 1986 and readily differs from the latter in having the much longer terminal projections of interlobar membrane in males, and the shorter length of dorsal setae *c2* and smaller size of the hysteronotal shield in females. A checklist with host associations of mite species of the genus *Ingrassia* is compiled. It is concluded that the fauna of the genus *Ingrassia* on charadriiforms is studied relatively well, while diversity of these mites on other host orders is known quite superficially. Comparison of the feather mite faunas known on different selected species of the tribe Scolopacini (dowitchers, snipes and woodcocks) shows that bird species of this tribe demonstrate a gradual loss of the primary feather mite fauna specific to the family Scolopacidae (Charadriiformes) and even its substitution with mites transferred from another host order.

**Keywords** Astigmata; feather mites; Ingrassiinae; *Ingrassia*; new species; Jacksnipe; Charadriiformes

**Zoobank** http://zoobank.org/AF644823-CDB7-4BE1-AC63-72288DBC7D4E

### Introduction

The feather mite genus *Ingrassia* Oudemans, 1905 (Analgoidea: Xolalgidae), currently including 30 species, and is the most specious genus in the subfamily Ingrassiinae (Oudemans 1905; Gaud 1972a; Gaud and Atyeo 1981; Chirov and Mironov 1990; Dabert and Ehrnsberger 1991; Vasyukova and Mironov 1991; Dabert 2000; Mironov and Palma 2006; Mironov and Proctor 2008; Stefan *et al.* 2013; Han *et al.* 2021; Mironov *et al.* 2025). Representatives of this genus are mostly small feather mites with the body size of adults 300-400 µm and with weakly sclerotized cuticular covers. In the plumage of avian hosts, mites of this genus, as for all xolalgids, inhabit the downy feathers and body covert feathers, where the barbs are not hooked together forming dense vanes.

Of 16 genera recognized in the subfamily Ingrassiinae (Gaud and Atyeo 1981, 1996; Mironov and Galloway 2002; Dabert *et al.* 2007), the genus *Ingrassia* and three more genera, *Leptosphyra* Hull, 1934, *Tectingrassia* Gaud, 1972 and *Vingrassia* Vasyukova & Mironov, 1991, constitute the *Ingrassia* generic group (Mironov *et al.* 2025). Representatives of this group are associated with aquatic bird orders. Only mites of the genus *Ingrassia* are distributed

Received 26 July 2025
Accepted 19 September 2025
Published 09 October 2025

Corresponding author
Sergey V Mironov ID:
sergei.mironov@zin.ru

Academic editor
Akashi Hernandes, Fabio

https://doi.org/10.24349/xegi-38o4

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_3.png" /></figure>

Mironov S. V. and Haarder S.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ª Zoological Institute, Russian Academy of Sciences, Universitetskaya embankment 1, 199034, Saint Petersburg, Russia.</footnote>

<footnote>ᵇ Independent researcher, Ornebjergvej 43, 4760 Vordingborg, Denmark.</footnote>
on hosts from five different orders, Charadriiformes, Phaethontiformes, Podicipediformes, Procellariiformes and Sphenisciformes (Table 1), while other genera are restricted to particular bird orders. Mites of the genera *Leptosphyra* and *Tectingrassia* occur on waders, or shorebirds (Charadriiformes: Charadrii), and representatives of the genus *Vingrassia*—on ducks and geese (Anseriformes). Key to species of these genera have been constructed only for species associated with charadriiform birds from Africa and Eurasia (Gaud 1972a; Chirov and Mironov 1990; Vasyukova and Mironov 1991); descriptions of species from other bird orders are scattered across various taxonomic publications (Vitzthum 1921; Černý 1967; Mironov and Proctor 2008; Stefan *et al.* 2013).

Mironov *et al.* (2025) recently provided the main diagnostic characteristics, world checklist and type hosts for all species of the *Ingrassia* generic group. This paper also noticed that the relatively weak differential features of genera and the pattern of host associations give evidence that the *Ingrassia* generic group certainly needs a revision of diagnostic characters and taxonomic borders of recognized genera (Gaud 1972a; Gaud and Atyeo 1981; Chirov and Mironov 1990). Phylogenetic study of xolalgids based on morphological features supported the monophyly of the group within the subfamily Ingrassiinae, but showed that the genus *Ingrassia* appears to be paraphyletic (Mironov 2005).

In the present work, we describe a new *Ingrassia* species found on the Jacksnipe, *Lymnocryptes minimus* (Brünnich, 1764), from Faroe Islands, Denmark. Additionally, we provide the list of host associations of the genus *Ingrassia* and compare the feather mite fauna of selected species of snipes (tribe Scolopacini).

## Material and methods

Mite specimens used in this study were collected by SH from three individuals of the Jacksnipes from Faroe Islands, Denmark. Dead birds were collected by Jens-Kjeld Jensen (Nólsoy, Faroe Islands) and kept in his freezer until sampling of mites and other ectoparasites. Some birds were killed by cats while the cause of death of others was unknown. Wing and body covert feathers were examined individually with a dissection microscope, and recovered mites were placed in tubes with 70% ethanol. Collected mite specimens were mounted on microscope slides in Hoyer's medium according to the standard technique used for small mites (Krantz and Walter 2009).

The species description follows the format used for feather mites of the subfamily Ingrassiinae (Mironov and Palma 2006; Mironov and Proctor 2008; Stefan *et al.* 2013; Mironov *et al.* 2017, 2025; Han *et al.* 2021). General morphological terms, idiosomal and leg chaetotaxies follow Gaud and Atyeo (1981, 1996) with minor corrections for coxal setae by Norton (1998). All the measurements are in micrometers (µm). Primary drawings and measurements were made using a Leica light microscope DM2500 (Leica Microsystems, Inc.) with DIC illumination and camera lucida; and final digital drawings were prepared with a Wacom Cintiq 22 graphics tablet (Wacom Co., Ltd). Systematics and scientific names of birds follow the IOC World Bird List, v 15.1 (Gill *et al.* 2025). Type material is deposited at the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZISP) and in the Department of Ecology and Zoology of the Universidade Federal de Santa Catarina, Florianópolis, Brazil (ECZ-UFSC); additional unmounted specimens are kept in ethanol in the private collection of the junior author and will eventually be deposited in the Acari collection at the National History Museum of Denmark, Copenhagen, Denmark (NHMD).
**Systematics**

**Family Xolagidae Dubinin, 1953**

**Subfamily Ingrassiinae Gaud & Atyeo, 1981**

**Genus Ingrassia Oudemans, 1905**

Type species: *Megninia veligera* Oudemans, 1904 by original designation.

**Ingrassia lymnocrypti** sp. n.

Zoobank: 164B46AC-D207-4D2B-967D-286389110B94

(Figures 1–4)

**Type material**

Male holotype (ZISP 23875), 5 male and 6 female paratypes (ZISP 23876–23886) from *Lymnocryptes minimus* (Brünnich, 1764) (Charadriiformes: Scolpacidae). Denmark, Faroe Islands, Torshavn, 13 September 2019, coll. S. Haarder; other paratypes: 5 male and 5 female paratypes (ZISP 23887–23896), same host, Denmark, Faroe Islands, Stremoy, 31 December 2001, coll. S. Haarder; 6 male and 3 female paratypes (ZISP 23897–23905), same host, Denmark, Faroe Islands, Torshavn, 14 February 2005, coll. S. Haarder.

**Type depository**

Holotype, 13 male and 11 female paratypes—ZISP, remaining paratypes—ECZ-UFSC.

**Description**

**Male** — (holotype, range for 10 paratypes in parentheses), (Figures 1, 3A-E, 4A, B). Idiosoma length (from anterior end to bases of setae *h3*) 410 (395–440), greatest width 200 (200–240), length of hysterosoma 259 (250–280). Prodorsal shield: longitudinal plate gradually enlarging from anterior end to level of scapular setae; narrowed anterior end rectangular, with 3-5 erratically orientated minute ridges; posterior part extending beyond scapular setae angular or semi-ovate, length 90 (88–100), greatest width 38 (35–40) (Figures 1A, 4A). Scapular setae *se* situated on teardrop-shaped sclerites base of setae *se* separated by 73 (70–80); setae *si* on striated tegument slightly posterior to level of setae *se*. Scapular shields: wide; inner margins of shield oblique, widely rounded posteriorly, without suprategumental extensions. Distance between prodorsal and hysteronotal shields along midline 20 (18–23). Hysteronotal shield: anterior margin slightly concave, length of shield from anterior margin to bases of setae *h3* 255 (255–285), surface without ornamentation, traces of fusion with humeral shields indistinct. Setae *c3* filiform, longer than trochanter III. Setae *c2, d2, e2* represented by macrosetae; all extending approximately to midlength of terminal cleft. Setae *d1* and *e1* posterior to levels of setae *c2* and *d2*, respectively. Supranal concavity narrowly ovate, connected posteriorly with terminal cleft by short and narrow grove. Opisthosomal lobes long, approximately 3–3.5 times longer than wide at base. Terminal cleft long triangular, with anterior end extending to level of trochanters IV, length of the cleft from anterior end to bases of setae *h3* 110 (110–125), greatest width anterior to setae *ps1* 50 (50–58). Terminal extensions on interlobar membrane long and narrowly triangular, with pointed apices, length of extensions from bases of setae *h3* to apices 60 (55–70); incision between extensions narrow, almost parallel-sided, with rounded anterior end extending to level of setae *h2*, and 78 (75–88) long. Setae *h3* posterior to level of setae *h2* and situated on apices of opisthosomal lobes. Setae *ps1* and *ps2* at the same transverse level anterior to setae *h2*, setae *f2* anterior to both pairs of pseudanal setae. Setae *f2* and *ps1* extending to apices of terminal extensions, setae *ps2* barely extending to midlength of these extensions. Distances between dorsal setae: *c2:d2* 50 (50–55), *d2:e2* 60 (60–68), *e2:h3* 135 (130–155), *c2:c2* 140 (135–165), *h2:h2* 78 (75–95), *h3:h3* 57 (53–63), *ps1:ps1* 40 (40–48).
<figure><img src="image_4.png" /><figcaption>**Figure 1** *Ingrassia lymnocrypti* sp. n., male. A – dorsal view, B – ventral view.</figcaption></figure>

Lengths of dorsal setae: c2 175 (170–205), c3 85 (85–110), d2 170 (160–185), e2 95 (90–100), f2 90 (90–120), ps1 85 (85–105).

Epimerites I fused in a Y, with stem (sternum) approximately half as long as total length of epimerites I. Epimerites III rudimentary. Inner ends of epimerites IIIa with sclerotized fields connected by narrow sclerotized bridge. Epimerites IVa long, encircling anterolaterally adanal area, inner ends of these epimerites with large triangular sclerotized fields flanking laterally postgenital area (Figures 1B, 4B). Coxal fields IV open anteriorly, central part of
coxal fields IV not sclerotized. Genital apparatus small, 10 (10–11) × 12 (12–13); aedeagus minute, barely extending from genital arch. Epiandrum (pregenital apodeme) shaped as low bow, 7 (7–9) long, 28 (28–32) wide. Genital shields represented by small longitudinal plates, with setae *g* on anterior ends. Adanal shields absent, setae *ps3* on soft tegument anterior to adanal suckers. Adanal suckers longitudinally ovate, with small denticle on anterior margin, 24 (24–28) in greater diameter; surrounding membrane with radial striation. Setae *1a* short filiform, not extending to level of tips of epimerites II. Setae *3a* filiform, barely extending to level of posterior margin of trochanters IV. Setae *4a* filiform, situated on striated tegument or on small sclerotized area of coxal field IV. Setae *4b* situated on inner ends of epimerites IIIa, represented by macrosetae, and extending beyond midlength of opisthosomal lobes. Setae *ps3* and *4a* approximately at same transverse level. Distances between ventral setae: *4b:3a* 25 (25–35), *4b:g* 60 (58–65), *g:ps3* 28 (25–32), *ps3:h3* 140 (135–155), *4b:4b* 25 (25–28), *g:g* 20 (20–24), *ps3:ps3* 23 (23–25). Lengths of ventral setae: *1a* 25 (25–35), *3a* 60 (58–62), *4a* 35 (35–50), *4b* 195 (190–210), *g* 28 (25–30), *ps2* 50 (48–58).

Tarsi I, II, each with a small nearly rectangular apicodorsal projection (Figure 3A, B). Setae *s* of tarsus II spiculiform with short filiform apex, remaining ventral setae of tarsi I, II filiform. Tibiae I, II, each with rounded ventral processus; setae *gT* of tibiae I, II spiculiform. Seta *mG* of genu I spiniform with filiform apex, seta *mG* of genu II long filiform. Setae *cGI*, *cGII* spiniform basally, with filiform apex. Femoragenu II with dorsal retrograde spine-shaped projection. Tibia III with large angular extension bearing solenidion *φ* (Figure 3C). Tarsus III 85 (83–98) long, curved, and with small finger-like apical projection. Tarsus IV 43 (38–48) long, with slightly convex inner margin, with slightly curved finger-like apical projection bearing setae *d* and *e*; setae *w* thickened and extending to ambulacral disc; modified setae *d*, *e* button-like, with barely distinct central nipple (Figure 3D). Legs IV excluding pretarsus 150 (145–155) long, with distal half of tarsus extending beyond lobar apices (level of setae *h3*), but not reaching terminal extensions of interlobar membrane. Lengths of solenidia: *ωI* 33 (30–35), *ωI* 45 (44–48), *σI* 42 (40–44), *σII* 8 (7–8), *σIII* 45 (43–50), *φIV* 44 (42–50). Ambulacral discs of tarsi III, IV approximately half as long as those on tarsi I, II and more ovate in shape.

**Female** — (range for 10 paratypes) (Figures 2, 3F, G; 4C, D). Idiosoma, length × width, 340–375 × 135–150; length of hysterosoma 215–250. Prodorsal shield: generally shaped as in the male, except lateral margins of anterior one third with distinct ledge and anterior end with 2-3 erratically orientated minute ridges; length 75–80, greatest width 36–40. Scapular setae *se* situated on teardrop-shaped sclerites, bases of setae *se* separated by 66–70; setae *si* on striated tegument slightly posterior to level of setae *se*. Scapular shields: wide; inner margins straight, rounded posteriorly, with suprategumental angular extension at level of anterior one third. Distance between prodorsal and hysteronotal shields along midline 50–54. Hysteronotal shield: shaped as narrow longitudinal plate in median area of hysterosoma, stretching from humeral shields to midlength of opisthosoma; anterior margin convex; lateral margins slightly sinuous; posterior margin deeply concave; posterior corners elongated and pointed, extending beyond level of setae *e2*; greatest length 155–165, width of anterior part 48–55, width at posterior margin 55–60 (Figures 2A, 4C). Setae *c3* filiform, shorter than trochanters III. Setae *c2*, *d2*, *e2* filiform, equal to or slightly longer than trochanters III; setae *c2* and *e2* on striated tegument, setae *d2* on hysteronotal shield near lateral margins. Microsetae *d1* and *e1* on hysteronotal shield, posterior to levels of setae *c2* and *d2*, respectively. Macrosetae *h3* about 3/4 the length of macrosetae *h2*. Distances between dorsal setae: *c2:d2* 75–80, *d2:e2* 82–90, *e2:h3* 70–75, *c2:d2* 62–65, *h2:h2* 58–64, *h3:h3* 48–52. Lengths of dorsal setae: *c2* 25–38, *c3* 15–20, *d2* 18–30, *e2* 17–20.

Epimerites I fused into a Y, with stem half as long as total length of epimerites I. Epigynum (pregenital apodeme) shaped as slightly curved transverse sclerite between bases of setae *4b*, 25–28 wide. Apodemes of oviporus long, with posterior ends extending to midlength of epimerites IV (Figures 2B, 4D). Setae *1a*, *3a*, *4a*, *4b* and *g* short filiform. Setae *g* situated posterior to level of setae *3a*. Distance between setae *4a* approximately equal to that between *4b*. Copulatory opening situated terminally; spermatheca indistinct. Distances between ventral
<figure><img src="image_5.png" /><figcaption>**Figure 2** *Ingrassia lymnocrypti* sp. n., female. A – dorsal view, B – ventral view.</figcaption></figure>

setae: 4b:3a 20–24, 4b:g 30–34, g:4a 52–58, 4a:4a 28–34. Lengths of ventral setae: 1a 15–18, 3a 18–24, 4a 12–17, 4b and g 10–12.
<figure><img src="image_6.png" /><figcaption>**Figure 3** *Ingrassia lymnocrypti* sp. n., details. A, B – legs I, II of male, dorsal view; C, D – tibiae and tarsi III, IV of male, dorsal view; E – genital and adanal areas of male; F – leg III of female, dorsal view, G – tibia and tarsus IV of female, dorsal view.</figcaption></figure>
Legs I, II as in the male, except femoragenu II without retrograde spine-like extension.
Legs IV with pretarsus barely extending beyond posterior margin of opisthosoma (Figure 2A,
B). Tarsi III, IV without distinct apicodorsal spine, 80–85 and 88–95 long, respectively (Figure
3F, G). Trochanteral setae sRIII 28–36 long, approximately equal to length of femoragenu III.
Setae *w* of tarsi III, IV thickened basally, with short filiform apices. Lengths of solenidia: ωI 1
28–30, ωII 40–45, σI 38–42, σII 7–8, σIII 10–12, φIII 33–35, φIV 18–25.

Remarks

The new species *Ingrassia lymnocrypti sp. n.* is most close to *I. limnodromi* Vasyukova & Mironov, 1986 from the long-billed dowitcher, *Limnodromus scolopaceus* (Say) (Vasyukova and Mironov 1986: 11, figs 5–7, 1991: 71, figs 46, 47, 56a). These species share the following combination of characters: in both sexes, the prodorsal shield is gradually enlarged to the level of scapular setae, with posterior ends extending beyond these setae angular or semi-ovate; in males, setae *c2*, *d2* and *e2* are represented by the macrosetae, the anterior ends of epimerites IIIa are connected with transverse bridge, tibia III with apicodorsal angular projection; in females, the hysteronotal shield is shaped as a longitudinal plate occupying the median area of hysterosoma, with deeply concave posterior margin, elongated long and pointed posterior corners extending beyond the level of setae *e2*. *Ingrassia lymnocrypti sp. n.* differs from the latter species in the following clear features: in males, the anterior margin of the hysteronotal shield is almost straight, the anterior ends of epimerites IVa bear triangular sclerotized fields, the terminal extensions of interlobar membrane are long and extend to the articulation of tibia and tarsus III, and legs IV reach only the midlenth of terminal extensions; in females, setae *c2* are filiform (25–35 long), the anterior end of the hysteronotal shield extends to the level of humeral shields, setae *e2* are situated off the lateral margins of the hysteronotal shield, and the oviporus lacks additional lateral sclerites. In males of *I. limnodromi*, the anterior margin of the hysteronotal shield is convex, the anterior ends of epimerites IVa lack sclerotized fields, the terminal extensions of interlobar membrane extend to the proximal one third of tibia III, and legs IV with entire tarsus extend beyond the apices of terminal extensions; in females, setae *c2* are represented by macrosetae (100–120 long), the anterior end of the hysteronotal shield extends far anterior to the humeral shields, setae *e2* are situated on the lateral margins of the hysteronotal shield, and the oviporus has a pair of longitudinal additional sclerites bearing bases of setae 4b (Vasyukova and Mironov, 1986: 11, figs. 5–7).

Etymology

The specific name is derived from the generic name of the type host and is a noun in the genitive case.

Discussion

The recently published world checklist of the *Ingrassia* generic group presented valid species
and only a type host for each mite species (Mironov *et al.* 2025). In the present work, we
have compiled the list of currently known species of the genus *Ingrassia* supplemented with
all reported hosts and localities (Table 1). Based on the number of charadriiform species
of the suborders Charadrii and Scolopaci known to date as hosts of *Ingrassia* (Gaud 1972a;
Vasyukova and Mironov 1990, 1991), it is possible to evaluate that the fauna of this genus on
shorebirds has been studied more or less well; while that on procellariiforms is obviously known
quite superficially (Stefan *et al.* 2013). On tropicbirds (Phaethontiformes: Phaethontidae),
*I. aequinoctialis* is the only species known from all three species of the genus *Phaethon*
comprising this order (Mironov *et al.* 2025). On grebes (Podicipediformes), the only *I. colymbi*
from the little grebe *Tachybaptus ruficollis* is currently known (Gaud 1974), although *Ingrassia*
species could probably be expected on other grebe species of this order. The record of *I.
eudyptula* on a penguin *Eudyptula minor* (Sphenisciformes) is still enigmatic, and this finding
<figure><img src="image_7.png" /><figcaption>**Figure 4** *Ingrassia lymnocrypti* sp. n., details. A – prodorsum of male; B – opisthosoma of male, ventral view; C – dorsal shields of female; D – oviporus of female, ventral view. Scale bar for photos A–D.</figcaption></figure>
<table><caption>Table 1 Host associations and distributions of species of the genus *Ingrassia*.</caption><thead><tr><th>Mite species</th><th>Host</th><th>Host family</th><th>Host order</th><th>Locality</th><th>Reference</th></tr></thead><tbody><tr><td>I. aquinoctialis (Trousart, 1889)</td><td>Phaethon acanthus Linnaeus*</td><td>Phaethontidae</td><td>Phaethontiformes</td><td>Tropical seas</td><td>Trousart 1899</td></tr><tr><td>"</td><td>P. lepturus Daudin</td><td>Phaethontidae</td><td>Phaethontiformes</td><td>Tropical seas</td><td>Trousart 1899; Mironov et al. 2025</td></tr><tr><td>I. americana Dabert & Ehmsberger, 1991</td><td>P. rubicunda Boddau</td><td>Phaethontidae</td><td>Charadriformes</td><td>Tropical seas, Pacific Ocean (Johnston Island)</td><td>Dabert & Ehmsberger 1991</td></tr><tr><td>I. antarctica (Gaud, 1952)</td><td>Tringa melanotouca (Gmelin)</td><td>Phaleoconides georgicus Murphy & Harper</td><td>Procellariiformes</td><td>Unknown (North America?)</td><td>Gaud, 1952</td></tr><tr><td>I. aphrius Dabert & Ehmsberger, 1991</td><td>Paleconoides georgicus Murphy & Harper</td><td>Procellariiformes</td><td>Charadriformes</td><td>France; Kerguelen Islands</td><td>Dabert and Ehmsberger 1991</td></tr><tr><td>I. arenaviae (Gaud, 1958)</td><td>Calidris virgata (Gmelin)</td><td>Procellariiformes</td><td>Charadriformes</td><td>Unknown (North America?)</td><td>Gaud 1958; Chirov 1979; Vasyukova and Mironov 1990</td></tr><tr><td>I. calonectris Stelan, Gomez-Diaz & Mironov, 2013</td><td>Arenaria interpre (Linnaeus)</td><td>Procellariiformes</td><td>Charadriformes</td><td>Russia (European pt.), Kirghizia, Morocco, Portugal (Azores Archipelago)</td><td>Stelan et al. 2013</td></tr><tr><td>I. centotibia Gaud, 1972</td><td>Calonectris borealis Cory</td><td>Procellariiformes</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Actitis hypoleuca (Linnaeus) (?)</td><td>Charadriformes</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Anachthus alexandrius (Linnaeus)</td><td>Charadriformes</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Charadrius dubius Scopoli</td><td>Charadriformes</td><td>Charadriformes</td><td>Morocco</td><td>Gaud 1972a; Vasyukova and Mironov 1990</td></tr><tr><td>"</td><td>Charadrius haiciatula Linnaeus</td><td>Charadriformes</td><td>Charadriformes</td><td>Russia (European pt.), Morocco, Cameroon</td><td>Gaud 1972a; Chirov 1979</td></tr><tr><td>"</td><td>Himantopus himantopus (Linnaeus)*</td><td>Charadriformes</td><td>Charadriformes</td><td>Morocco</td><td>Chirov 1979</td></tr><tr><td>"</td><td>Phalaropus lobatus (Linnaeus) (?)</td><td>Charadriformes</td><td>Charadriformes</td><td>Kirghizia</td><td>Han et al. 2021</td></tr><tr><td>I. chionis Han, Mironov, Kim & Min, 2021</td><td>Chionis albus (Gmelin)</td><td>Charadriformes</td><td>Charadriformes</td><td>Antarctica (King George Island)</td><td>Han et al. 2021</td></tr><tr><td>I. colynbi Gaud, 1974</td><td>Tachyparus ruficollis (Pallas)</td><td>Charadriformes</td><td>Charadriformes</td><td>Morocco</td><td>Dublin 1974</td></tr><tr><td>I. dubinini Černý, 1967</td><td>Ardenna temminckii (Temminck) *</td><td>Podicipedidae</td><td>Charadriformes</td><td>Russia (Wrangel Island)</td><td>Dublin 1949; Černý 1967</td></tr><tr><td>I. eudypala Mironov & Proctor, 2008</td><td>Hydrobates pelagicus (Linnaeus) (?)</td><td>Procellariiformes</td><td>Procellariiformes</td><td>Russia (Sea of Okhinsk)</td><td>Dublin 1949; Černý 1967</td></tr><tr><td>I. fassitarsa (Gaud, 1958)</td><td>Endypula minor (Forster)</td><td>Sphesicidae</td><td>Procellariiformes</td><td>Australia (Fraser Island)</td><td>Mironov and Proctor 2008</td></tr><tr><td>I. folicera (Gaud, 1953)</td><td>Gallinago gallinago (Linnaeus)</td><td>Charadriformes</td><td>Procellariiformes</td><td>Russia (European pt.), Morocco</td><td>Gaud 1958; Mironov 1981</td></tr><tr><td>I. forcipata (Haller, 1882)</td><td>Hydrobates castro (Haller)</td><td>Hydrobathidae</td><td>Procellariiformes</td><td>Sengal</td><td>Gaud 1953</td></tr><tr><td>"</td><td>Actitis macularia (Linnaeus)*</td><td>Scopopacidae</td><td>Charadriformes</td><td>Canada</td><td>Haller 1882</td></tr><tr><td>"</td><td>Actitis hypoleuca (Linnaeus)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Kirghizia</td><td>Chirov 1979; Vasyukova and Mironov 1990, 1991</td></tr><tr><td>"</td><td>Tringa glareola Linnaeus (?)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Cameroon, Namibia</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Tringa nebularia (Gunnens) (?)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Tringa stagnatilis (Bedstein) (?)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr><tr><td>"</td><td>Tringa totanus (Linnaeus) (?)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Cameroon</td><td>Hull 1934; Gaud 1972a</td></tr><tr><td>I. limnodromi Vasyukova & Mironov, 1986</td><td>Limnodromus scolopacae (Say)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Russia (Yakutia)</td><td>Vasyukova and Mironov 1986, 1991</td></tr><tr><td>I. limnoe Gaud, 1972</td><td>Limosa lapponica (Linnaeus)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Morocco</td><td>Gaud 1972a</td></tr><tr><td>I. lymocrypti Sp. n.</td><td>Lymocryptes minima (Brunlich)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Russia (Yakutia)</td><td>Vasyukova and Mironov 1991</td></tr><tr><td>I. micronota Stelan, Gomez-Diaz & Mironov, 2013</td><td>Bubneria butleri (Jardine & Selby)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Dennmark (Faroe Islands)</td><td>Present work</td></tr><tr><td>I. oceanica (Vitzhum, 1921)</td><td>Hydrobates pelagicus (Linnaeus)</td><td>Procellariiformes</td><td>Procellariiformes</td><td>Cape Verde (Ilheu Grande)</td><td>Stelan et al. 2013</td></tr><tr><td>I. oceanodromae Černý, 1967</td><td>Hydrobates leucorhous (Viehlot)</td><td>Hydrobathidae</td><td>Procellariiformes</td><td>Germany (Helgoland)</td><td>Vitzhum 1921</td></tr><tr><td>I. phalaropi Gaud, 1972</td><td>Phalaropus falcarius (Linnaeus)</td><td>Scopopacidae</td><td>Procellariiformes</td><td>Dennmark (Faroe Islands)</td><td>Černý 1967</td></tr><tr><td>"</td><td>Phalaropus lobatus (Linnaeus)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Morocco</td><td>Gaud 1972a</td></tr><tr><td>I. philomachi Gaud, 1972</td><td>Calidris pugnax (Linnaeus)</td><td>Charadriformes</td><td>Charadriformes</td><td>Russia (European pt., W. Siberia)</td><td>Vasyukova and Mironov 1990</td></tr><tr><td>I. platyspina Mironov & Palma, 2006</td><td>Prosopiona parvirostris (Peale)</td><td>Charadriformes</td><td>Charadriformes</td><td>Russia (European pt., Yakutia), Kirghizia, Cameroon</td><td>Gaud 1972a; Chirov 1979; Vasyukova and Mironov 1990, 1991</td></tr><tr><td>I. semidissa Gaud, 1972</td><td>Glareola praticola (Linnaeus)</td><td>Glareolidae</td><td>Charadriformes</td><td>French Polynesia (Tuamotu Island)</td><td>Mironov and Palma 2006</td></tr><tr><td>I. slonskiana Dabert, 2000</td><td>Tringa ochroptera (Linnaeus)</td><td>Glareolidae</td><td>Charadriformes</td><td>Zambia</td><td>Gaud 1972a</td></tr><tr><td>I. spinata (Gaud & Mouchet, 1959)</td><td>Glareola nuchalis Gray</td><td>Glareolidae</td><td>Charadriformes</td><td>Poland</td><td>Dabert 2000</td></tr><tr><td>I. strictor (Berlese, 1898)</td><td>Calidris pugnax (Linnaeus)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud and Mouchet 1959</td></tr><tr><td>I. tridacryla (Gaud & Mouchet, 1959)</td><td>Acrophilornis atricuspis (Gmelin)</td><td>Jacanidae</td><td>Charadriformes</td><td>Italy</td><td>Berlese 1898</td></tr><tr><td>I. tringae (Vitzhum, 1922)</td><td>Calidris minima (Leeser)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Bulgaria, Russia (European pt., Yakutia), Zambia, South Africa</td><td>Gaud and Mouchet 1959</td></tr><tr><td>I. veligera (Oudemans, 1904)</td><td>Tringa flaviipes (Gmelin)*</td><td>Scopopacidae</td><td>Charadriformes</td><td>Giyuna</td><td>Vitzhum 1922; Gaud 1972a; Vasyukova and Mironov 1990, 1991</td></tr><tr><td>"</td><td>Tringa glareola Linnaeus</td><td>Scopopacidae</td><td>Charadriformes</td><td>Russia (European pt., Yakutia), Kirghizia</td><td>Oudemans 1904</td></tr><tr><td>"</td><td>Tringa nebularia (Gunnens)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Russia (European pt., Yakutia)</td><td>Chirov 1979; Vasyukova and Mironov 1990, 1991</td></tr><tr><td>"</td><td>Tringa stagnatilis (Bedstein)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Poland, Russia (European pt.)</td><td>Vasyukova and Mironov 1990; Dabert, 2000</td></tr><tr><td>"</td><td>Tringa totanus (Linnaeus)</td><td>Scopopacidae</td><td>Charadriformes</td><td>Kirghizia</td><td>Chirov 1979</td></tr><tr><td>I. xiphidiopeti Gaud, 1972</td><td>Vanellus albiiceps Gould</td><td>Charadriformes</td><td>Charadriformes</td><td>Cameroon</td><td>Gaud 1972a</td></tr></tbody></table>

Remarks: *—type host, if a mite species is known from several avian species, (?)— questionable host association.
could represent the result of natural horizontal transfer from a procellariform host (Mironov and Proctor 2008).

Most of currently known *Ingrassia* species are monoxenous ectoparasites. Much lesser number of species are stenoxenous, i.e. associated with various species of the same genus (*Ingrassia aequinoctialis*, *I. limosae*, *I. phalaropi*, and *I. veligera*). Associations of *I. forcipata* with hosts from different genera and *I. centrotibia* with hosts from different families look highly doubtful. Perhaps, the reports of too different hosts for these mite species could be caused by contamination or misidentification of similar species.

The species *I. centrotibia* was originally found on the black-winged stilt, *Himantopus himantopus* (Recurvirostridae), and plovers of the genera *Anarhynchus* and *Charadrius* (Charadriidae) (Gaud 1972a). Therefore, the records of this mite on some scolopacids (Table 1), the common sandpiper *Actitis hypoleucos* and the red-necked phalarope *Phalaropus lobatus* having their own specific *Ingrassia* species, seem highly likely misidentifications. Additionally, although plovers and stilts are closely related families, the reports of *I. centrotibia* from *H. himantopus* and plovers of the genera *Anarhynchus* and *Charadrius* allow suggesting that even on these hosts this mite could potentially be a complex of closely related species.

*Ingrassia forcipata* was originally described from the spotted sandpiper *Actitis macularia* by Haller (1882) and further found on the common sandpiper *A. hypoleucos* (Vasyukova and Mironov 1990). Reports of this mite species from shanks and tattlers (Scolopacidae: *Tringa*) by Gaud (1972a) rise doubts, because from most of Eurasian and North American species of the genus *Tringa* was also reported *I. veligera*, which is known exclusively from tattlers and shanks (Oudemans 1904; Vasyukova and Mironov 1991; Dabert 2000). Indeed, the species *I. forcipata* and *I. veligera* are very similar in general appearance (Vasyukova and Mironov 1991: fig. 49 vs. fig. 50); therefore, it is possible to suggest that specimens from tattlers from Africa and Europe were misidentified as "*I. forcipata*" by Gaud (1972a) and Mironov (1981).

With the description of a new *Ingrassia* species, the feather mite fauna of the jacksnipe now includes three species inhabiting its plumage: *Alloptes lymnocrypti* Mironov, 1981 (Alloptidae), *Ingrassia lymnocrypti* (Xolalgidae), and *Rafalskiata rackae* (Gaud, 1972) (Avenzoariidae). It is interesting to compare its fauna with those of close species from the tribe Scolopacini, like snipes, woodcocks and dowitchers, and other scolopacids. According to fundamental studies of feather mite fauna of Charadriiformes (e.g. Dubinin 1951, 1956; Gaud and Till 1961; Gaud 1972a; Vasyukova and Mironov 1986, 1990, 1991; Dabert and Ehrnsberger 1999; Dabert 2003),

<table><caption>**Table 2** Feather mites inhabiting the plumage in some waders of the family Scolopacidae.</caption><thead><tr><th>Host species</th><th>Alloptidae</th><th>Avenzoariidae</th><th>Xolalgidae</th><th>Pterolichidae</th><th>Syringobiidae</th><th>Proctophyllodidae</th><th>References</th></tr></thead><tbody><tr><td><strong>Tringini</strong><br><em>Tringa nebularia</em></td><td><em>Alloptes atelesthetus</em><br>Gaud, 1972</td><td><em>Avenzoaria totani</em><br>(Canestrini, 1878);<br><em>Bregetovia obtusolobata</em><br>Dubinin, 1951</td><td><em>Ingrassia veligera</em><br>(Oudemans, 1904)</td><td><em>Montchadskiana ulitae</em><br>Dubinin, 1951</td><td><em>Plutarchusia chelopus</em><br>(Oudemans, 1904);<br><em>Syringobia parachuteopus</em><br>Vasyukova & Mironov, 1986</td><td>—</td><td>Dubinin 1951, 1956; Gaud 1972; Vasyukova and Mironov 1990, 1991; Mironov 1992; Dabert and Ehrnsberger 1999; Dabert 2003</td></tr><tr><td><strong>Scolopacini</strong><br><em>Limnodromus scolopaceus</em></td><td><em>Alloptes limnodromi</em><br>Vasyukova & Mironov, 1991</td><td><em>Avenzoaria macrorhamphi</em><br>Dubinin, 1956</td><td><em>Ingrassia limnodromi</em><br>Vasyukova & Mironov, 1986</td><td>—</td><td><em>Phyllochaeta gracilis</em><br>Vasyukova & Mironov, 1986;<br><em>Ph. latior</em><br>Vasyukova & Mironov, 1986</td><td>—</td><td>Dubini 1956; Vasyukova and Mironov 1986, 1990, 1991; Dabert 2003</td></tr><tr><td><em>Lymnocryptes minimus</em></td><td><em>Alloptes lymnocrypti</em><br>Mironov, 1981</td><td><em>Rafalskiata rackae</em><br>(Gaud, 1972)</td><td><em>Ingrassia lymnocrypti</em><br><strong>sp. n.</strong></td><td>—</td><td>—</td><td>—</td><td>Gaud 1972b; Mironov 1981; Mironov and Dabert 1997</td></tr><tr><td><em>Gallinago gallinago</em></td><td>—</td><td><em>Capelloptes flagellicaulus</em><br>(Trouessart & Neumann, 1888)</td><td><em>Ingrassia fissitarsa</em><br>(Gaud, 1958)</td><td>—</td><td>—</td><td>—</td><td>Dubinin 1951; Gaud 1958, 1972a</td></tr><tr><td><em>Scolopax rusticola</em></td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td><em>Proctophyllodes scolopacinus</em><br>(Koch, 1842)</td><td>Atyeo and Braasch 1966</td></tr></tbody></table>
the set of mite species on most of typical scolopacids, such as sandpipers, tattlers and shanks (tribes Tringini and Arenariini), is rich and commonly includes at least one species from each of the following families: Alloptidae, Avenzoariidae, Xolalgidae (Analgoidea), Pterolichidae and Syringobiidae (Pterolichoidea). The set of mite species associated with well-explored host species of the tribe Scolopacini, most of which are common game birds, is shown in Table 2. The data for the common greenshank, *Tringa nebularia* (Tringini), having a typical set of feather mites for tringines, are presented for comparison. It is easy to see that the feather mite fauna on birds of the tribe Scolopacini is reduced compared to those on typical waders, because all these hosts have lost representatives of the family Pterolichidae and almost all have lost mites of the family Syringobiidae. Of these four species of scolopacines, two bird species, the jacksnipe and long-billed dowitcher, *Limnodromus scolopaceus*, harbor mite species from three analgoidean families, Alloptidae, Avenzoariidae, and Xolalgidae. Additionally, the long-billed dowitcher retains quill-inhabiting feather mites of the family Syringobiidae (Pterolichoidea). The common snipe, *Gallinago gallinago*, has lost representatives of the family Alloptidae and harbors only mites of the families Avenzoariidae and Xolalgidae. Finally, the Eurasian woodcock, *Scolopax rusticola*, has entirely lost the primary feather mite fauna being specific to charadriiforms and has accepted a new feather mite fauna obviously received from some passerines—*Proctophyllodes scolopacinus* (Koch, 1842) (Proctophyllodidae). Mites of the family Proctophyllodidae are predominately distributed on passerines, and *P. scolopacinus* being specific to the woodcocks (*Scolopax*) is the closest to *P. corvorum* Vitzthum, 1922 associated with various corvids of the genus *Corvus* (Atyeo and Braasch 1966). The feather mite fauna of the woodcock is a rare and bright example of horizontal transfer of ectoparasites between hosts of different avian orders that gives a wide field of speculation, how the ancestor of woodcocks could accept a feather mite species from corvids. In general, the considered avian hosts of the tribe Scolopacini demonstrate a model of gradual loss of the primary feather mite fauna of Scolopacidae and the appearance of a new fauna by the horizontal transfer.

## Acknowledgements

The authors wish to thank Jens-Kjeld Jensen (Nólsoy, Faroe Islands) for providing access to the studied bird skins. The taxonomic part of the study was supported by the Ministry of Science and Higher Education of the Russian Federation (project No. 125013001089-0) for SM.

## ORCID

Sergey V Mironov https://orcid.org/0000-0003-0417-5687

Simon Haarder https://orcid.org/0000-0001-5981-738X

## References

* Atyeo W.T., Braasch N.L. 1966. The feather mite genus *Proctophyllodes* (Sarcoptiformes: Proctophyllo-didae). Bull. Univ. Nebraska State Mus., 5: 1-354.

* Berlese A. 1898. *Megninia centropodos* var. *strictior*. Acari, Myriopoda et Scorpiones hucusque in Italia reperta, Padova, Fascicolo 87, No 6.

* Černý V. 1967. Trois espèces nouvelles des Acariens plumicoles (Analgoidea) parasites des Procellariiformes. Folia Parasitol., 14: 87-91.

* Chirov P.A. 1979. Mites of the superfamily Analgoidea, which live on birds of Kirgizia. Entomologich-eskie issledovaniia v Kirgizii, Akademiia nauk Kirgizskoi SSR, 49-54. [In Russian]

* Chirov P.A., Mironov S.V. 1990. Feather mites of the subfamily Ingrassiinae of limicolines and ducks. Izvestiya Akademii nauk Kirgizkoi SSR, Khimiko-Teknologicheskie i Biologicheskie Nauki, 3: 74-83. [In Russian]

* Dabert J. 2000. Feather mites (Acari: Astigmata) of water birds of the Slońsk nature reserve with description of a new species. Biol. Bull. Poznań, 37: 303-316.

* Dabert J. 2003. The feather mite family Syringobiidae Trouessart, 1896 (Acari, Astigmata, Pterolichoidea). I. Systematics of the family and description of new taxa. Acta Parasitol., 48 (Supplement): S1-S184.
Dabert J., Badek A., Skoracki M. 2007. New feather mite species (Acari, Astigmata) from the Sulphur-crested Cockatoo *Cacatua galerita* and Yellow-crested Cockatoo *C. sulphurea* (Psittaciformes, Cacatuidae). Acta Parasitol., 52(3): 250-267. https://doi.org/10.2478/s11686-007-0034-z

Dabert J., Ehrnsberger R. 1991. Zwei neue Federmilben-Arten aus der Gattung *Ingrassia* Oudemans, 1905 (Analgoidea; Xolalgidae; Ingrassiinae). Osnabrücker Naturwissenshaftliche Mitteilungen, 17: 127-142.

Dabert J., Ehrnsberger R. 1999. Systematics of the feather mite genus *Montchadskiana* Dubinin, 1951 (Pterolichoidea, Pterolichidae, Magimeliinae) with description of five new species. Acta Zool. Cracov., 42 (2): 219-249.

Dubinin V.B. 1949. Feather mite fauna of birds of the order Procellariiformes and its features. *Parazitologicheskii Sbornik*, 11: 201-228. [In Russian]

Dubinin V.B. 1951. Feather mites of birds of the Baraba Steppe. Report I. Feather mites of waterfowl and wading birds of the orders of rails, grebes, palmipedes, anserines, herons, gulls, and limicoles. *Parazitologicheskii Sbornik*, 13: 120-256. [In Russian]

Dubinin V.B. 1956. Feather Mites (Analgesoidea). Part 3. Family Pterolichidae. Fauna of the USSR, Arachnida, Vol. 6, Issue 7. Publisher: Akademia Nauk SSSR, Moscow-Leningrad, 813 pp. [In Russian]

Gaud J. 1952. Acariens plumicoles (Analgesidae) de quelques oiseaux des Îles Kerguelen (Récolte P. Paulian). Mém. Inst. Sci. Madagascar, Sér. A, 7: 161-166.

Gaud J. 1953. Sarcoptides plumicoles des oiseaux d'Afrique occidentale et centrale. Ann. Parasitol. Hum. Comp., 28: 193-226. https://doi.org/10.1051/parasite/1953283193

Gaud J. 1958. Acariens plumicoles (Analgesoidea) parasites des oiseaux du Maroc. II. Analgidae. Bull. Soc. Sci. Natur. Phys. Maroc, 38: 27-49.

Gaud J. 1972a. Acariens Sarcoptiformes plumicoles (Analgoidea) parasites sur les oiseaux Charadriiformes d'Afrique. Mus. Roy. Afr. Centr., Ann., Sér. in-8°, Sci. Zool., 193: 1-116.

Gaud J. 1972b. Les Sarcoptiformes plumicoles (Analgoidea) parasites des Scolopacinae, avec la description d'une espèce nouvelle: *Avenzoaria rackae* n. sp. Entomologische Mitteilungen aus dem Zoologischen Museum, Hamburg, 4: 241-247.

Gaud J. (1973) 1974. Quelques espèces nouvelles de Sarcoptiformes plumicoles (Analgidae et Dermoglyphidae) parasites d'oiseaux d'Europe. Acarologia, 15: 727-758.

Gaud J., Atyeo W.T. 1981. La famille Xolalgidae Dubinin, nouveau statut (Sarcoptiformes plumicoles, Analgoidea). I. Sous-famille Ingrassiinae, n. sub. fam. Acarologia, 22: 63-79.

Gaud J., Atyeo W.T. 1996. Feather mites of the World (Acarina, Astigmata): the supraspecific taxa. Mus. Roy. Afr. Centr., Ann., Sci. Zool., 277 (Pt. 1): 1-193 (text) and (Pt. 2): 1-436 (illustrations).

Gaud J., Mouchet J. 1959. Acariens plumicoles (Analgesoidea) parasites des oiseaux du Cameroun. II. Analgesidae. Ann. Parasitol. Hum. Comp., 34: 149-208. https://doi.org/10.1051/parasite/1959341149

Gaud J., Till W.M. 1961. Suborder Sarcoptiformes. In: Zumpt F. (Ed.). The Arthropod Parasites of Vertebrates in Africa South of the Sahara (Ethiopian Region). Vol. I. Chelicerata. Publications of the South African Institute of Medical Research, No 1 (Vol. IX), Johannesburg, pp. 180-352.

Gill F., Donsker D., Rasmussen P. (Eds.). 2025. IOC World Bird List (v 15.1). Available from http://www.worldbirdnames.org/ (accessed 25 June 2025). https://doi.org/10.14344/IOC.ML.15.1.

Haller G. 1882. Zur Kenntniss der Dermaleichiden. Archiv für Naturgeschichte, 48: 47-79.

Han Y.-D., Mironov S.V., Kim J.-H., Min, G.-S. 2021. Feather mites (Acariformes: Astigmata) from marine birds of the Barton Peninsula (King George Island, Antarctica), with descriptions of two new species. Zookeys, 1061: 109-130. https://doi.org/10.3897/zookeys.1061.71212

Hull J.E. 1934. Concerning British Analgidae (Feather-mites). Transact. Northern Naturalists Union, 1: 200-206.

Krantz G., Walter D. (Eds.). 2009. A Manual of Acarology, 3rd Edition. Texas Tech University Press, Lubbock, Texas, 807 pp.

Mironov S.V. 1981. Feather mites (Acarina, Sarcoptiformes, Analgoidea) of the birds of the family Charadriidae from the Curonian Spit. In: Balashov Yu.S. (Ed.). Morphological Peculiarities of Mites and Arachnids. Trudy Zoologicheskogo Instituta AN SSSR, 106: 66-75. [In Russian]

Mironov S.V. 1992. The review of species of the feather mite genus *Bregetovia* in the USSR's fauna (Analgoidea: Avenzoariidae). *Parazitologicheskii Sbornik*, 37: 126-150. [In Russian]

Mironov S.V. (2004) 2005. Phylogeny of the feather mite family Xolalgidae (Astigmata: Analgoidea) and revolutionary trends with non-passerine birds. Phytophaga, 14: 433-449.

Mironov S.V., Dabert J. 1997. Taxonomic notes on the feather mite subfamily Avenzoariinae with establishing of two new genera (Acarina: Analgoidea: Avenzoariidae). Genus, 8 (1): 75-79.

Mironov S. V., Galloway T.D. 2002. Four new species of feather mites (Acari: Analgoidea). Canadian Entomol., 134: 605-618. https://doi.org/10.4039/Ent134605-5

Mironov S.V., Palma R.L. 2006. Two new feather mite species (Acari: Analgoidea) from the Tuamotu sandpiper *Aechmorhynchus parvirostris* (Charadriiformes: Scolopacidae). Tuhinga - Records of the Museum of New Zealand Te Papa Tongarewa, 17: 49-59.

Mironov S.V., Proctor H.C. 2008. The probable association of feather mites of the genus *Ingrassia* Oudemans, 1905 (Analgoidea: Xolalgidae) with the blue penguin *Eudyptula minor* (Aves: Sphenisciformes) in Australia. J. Parasitol., 94: 1243-1248. https://doi.org/10.1645/GE-1579.1

Mironov S.V., Ehrnsberger R., Dabert J. 2017. Feather mites of the genera *Dubininia* and *Cacatualges* (Acari: Xolalgidae) associated with parrots (Aves: Psittaciformes) of the Old World. *Zootaxa*, 4272 (4): 451-490. https://doi.org/10.11646/zootaxa.4272.4.1

Mironov S.V., Stormer H., Proctor H.C. 2025. Redescription of the feather mite *Ingrassia aequinoctialis* (Trouessart, 1899) (Acariformes: Xolalgidae) with notes on systematics and host associations of the *Ingrassia* genus-group. Acarologia, 65(2): 482-496. https://doi.org/10.24349/f5zx-leve
Norton R. 1998. Morphological evidence for the evolutionary origin of Astigmata (Acari: Acariformes). Exp. Appl. Acarology, 22: 559-594. https://doi.org/10.1023/A:1006135509248

Oudemans A.C. 1904. Acarologische Aanteekeningen XIV. Entomol. Bericht., 20: 190-195. https://doi.org/10.5962/bhl.part.1123

Oudemans A.C. 1905. Acarologische Aanteekeningen XVII. Entomol. Bericht., 23: 222-226. https://doi.org/10.5962/bhl.part.1126

Stefan L.M., Gomez-Diaz, E., Mironov S.V. 2013. Three new species of the feather mite subfamily In-grassiinae (Acariformes: Xolalgidae) from shearwaters and petrels (Procellariiformes: Procellariidae). Zootaxa, 3682 (1): 105-120. https://doi.org/10.11646/zootaxa.3682.1.4

Trouessart E.L. (1898) 1899. Diagnoses préliminaires d'espèces nouvelles d'Acariens plumicoles. Additions et corrections à la sous-famille des Analgésinés. Bull. Soc. Étud. Sci. Angers, 28: 1-62.

Vasyukova T.T., Mironov S.V. 1986. New species of feather mites of Siberia. Publisher: Nauka, Siberian Dept., Novosibirsk, 72 pp. [In Russian]

Vasyukova T.T., Mironov S.V. 1990. Fauna and ecology of feather mites of Anseriformes and Charadriiformes of Yakutia. Publisher: Yakutskii Nauchnyi Tsentr, Sibirskoye Otdelenie Akademii Nauk SSSR, Yakutsk, 96 pp. [In Russian]

Vasyukova T.T., Mironov S.V. 1991. Feather mites of Anseriformes and Charadriiformes of Yakutia. Systematics. Publisher: Nauka, Siberian Dept., Novosibirsk, 201 pp. [In Russian]

Vitzthum H. 1921. Acarologische Beobachtungen. (5. Reihe). Archiv für Naturgeschichte, 87A (4): 1-77.

Vitzthum H. 1922. Acarologische Beobachtungen. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 44 (6): 517-564.