Mironov, Sergey V. ¹ ; Stormer, Hannah ² and Proctor, Heather C. ³

¹✉ Zoological Institute, Russian Academy of Sciences, Universitetskaya embankment 1, 199034, Saint Petersburg, Russia.
²Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada.
³Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada.

2025 - Volume: 65 Issue: 2 pages: 482-496

ZooBank LSID: 0936B9ED-6025-4C6D-AFB6-555A4511F154

Original research

Keywords

Abstract

Introduction

The feather mite genus Ingrassia Oudemans, 1905, with 30 currently known species, is the most species-rich genus in the subfamily Ingrassiinae (Analgoidea: Xolalgidae) (Oudemans 1905; Černý 1967; Gaud 1972; Gaud and Atyeo 1981a; 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). As for all representatives of the family Xolalgidae, these mostly small feather mites (with body size of adults about 300 μm) mainly inhabit the downy and covert feathers in the plumage of their avian hosts. Species of this genus have been recorded so far from hosts of five orders of aquatic birds: Charadriiformes, Phaethontiformes, Podicipediformes, Procellariiformes and Sphenisciformes. The majority of known Ingrassia species (20 species) are associated with charadriiforms, seven species live on procellariiforms, and one species is known from each of the three remaining host orders (Table 1).

In their generic revision of the family Xolalgidae, Gaud and Atyeo (1981a, 1981b) provided a modern taxonomic diagnosis of the genus Ingrassia and a review of its species content. Investigations on the systematics and diversity of Ingrassia in the past fifty years have been mainly focused on species associated with hosts from the order Charadriiformes (Gaud 1972; Vasyukova and Mironov 1986, 1991; Chirov and Mironov 1990; Dabert and Ehrnsberger 1991; Dabert 2000; Mironov and Palma 2006; Han et al. 2021). To date, identification keys to charadriiform-associated Ingrassia species are available for those in Africa (Gaud 1972) and in northern Eurasia (Chirov and Mironov 1990; Vasyukova and Mironov 1991), while all descriptions of the species associated with birds from other orders are scattered in various taxonomic papers (Černý 1967; Mironov and Proctor 2008; Stefan et al. 2013).

In the present paper, we redescribe the species Ingrassia aequinoctialis (Trouessart, 1899), which has been recorded from three species of tropicbirds (Phaethontidae): the Red-billed Tropicbird Phaethon aethereus Linnaeus, 1758 (type host), White-tailed Tropicbird P. lepturus Daudin, 1802 (=P. candidus Brisson, 1760), and Red-tailed Tropicbird P. rubricauda Boddaert, 1783 (=P. phoenicurus Gmelin, 1789). Although I. aequinoctialis is readily recognized because of its extremely large size compared to other Ingrassia species, it has never been illustrated or carefully redescribed since its brief initial text-only description by Trouessart (1899: 20). Apparently for these reasons, this mite species for a long time remained in the genus Megninia Berlese, 1881 (Analgidae) (Radford 1953, 1958), and was placed in Ingrassia only in the revision of the family Xolalgidae (Gaud and Atyeo 1981a). In addition to this redescription, we provide an updated world checklist of Ingrassia species together with those of three closely related genera, and discuss relationships and host associations of the Ingrassia genus-group.

Material and methods

Feather samples from Red-tailed Tropicbirds, Phaethon rubricauda, were collected under permit from the US Fish and Wildlife Service in June 2002 at Johnston Island, a part of Johnston Atoll National Wildlife Refuge, located in the central Pacific Ocean, a part of the United States. Capturing, handling and feather sampling was done by Julian P. Donahue and Dr. E. A. Schreiber, both from the Los Angeles County Museum of Natural History. Collected material was preserved in vials with 70% ethanol. These samples were subsequently shipped to the Proctor Lab at the University of Alberta. Mite specimens were cleared for 24 hours in lactic acid, and then mounted by HP and HS on microscope slides in commercially prepared PVA medium (BioQuip Products, Rancho Dominguez, California). Slides were cured for four days on a slide warmer set at ~40°C prior to being examined using DIC light microscopy on a Leica DMLB compound microscope. Slide-mounted specimens of Ingrassia aequinoctialis were sent to SM for examination and illustration.

The redescription and measuring techniques follow the standard formats used for ingrassiine feather mites (Mironov and Palma 2006; Mironov and Proctor 2008; Stefan et al. 2013; Mironov et al. 2017; Han et al. 2021). Morphological terms follow Gaud and Atyeo (1981a, 1996); the idiosomal and leg chaetotaxies also follow these authors with corrections for coxal setae by Norton (1998). Primary drawings and measurements were made using Leica light microscopes (DM2500 and DM 5000B, Leica Microsystems, Inc.) with DIC illumination and camera lucida; and final digital drawings were prepared using a Wacom Cintiq 22 graphics tablet (Wacom Co., Ltd). All the measurements are in micrometres (μm). Systematics and scientific names of birds follow the IOC World Bird List, v 14.2 (Gill et al. 2024). The examined material is deposited at the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia.

Systematics

Family Xolagidae Dubinin, 1953

Subfamily Ingrassiinae Gaud & Atyeo, 1981

Genus Ingrassia Oudemans, 1905

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

Ingrassia aequinoctialis (Trouessart, 1899)

(Figures 1–6)

Megninia aequinoctialis Trouessart 1899: 20; Radford 1953: 206, 1958: 112.

Ingrassia aequinoctialis: Gaud and Atyeo, 1981a: 71.

Material examined

3 males and 3 females (ZISP 23748–23753) from Phaethon rubricauda Boddaert, 1783 (Phaethontiformes: Phaethontidae), Johnston Island, Johnston Atoll National Wildlife Refuge (16°44′13″N, 169°31′26″W), U.S.A.; June 2002; coll. J. P. Donahue and E. A. Schreiber (see Material & Methods for more details).

Description

Male — (range for 3 specimens) (Figs. 1, 2, 5A-E). Idiosoma: length from anterior end to bases of setae h3 475–500, greatest width 405–415, length of hysterosoma 300–330. Prodorsal shield: nearly rectangular longitudinal plate slightly attenuate anteriorly, posterior margin almost straight or slightly convex, length 115–120, width at posterior margin 60–65, anterior end with longitudinal crest about 10 long, posterior end extending slightly beyond level of scapular setae se. Scapular setae se, si situated on striated tegument at the same transverse level; base of setae se separated by 135–150. Scapular shields: wide, inner margins of shield convex with narrow suprategumental fold, posterior end with angular suprategumental extension. Distance between prodorsal and hysteronotal shield 10–20. Hysteronotal shield: anterior margin convex and slightly sinuous, length of shield from anterior end to bases of setae h3 330–350, surface uniformly punctured, traces of fusion with humeral shields indistinct. Setae c2, d2, e2 and c3 short filiform, shorter than trochanter III. Setae d1 and setae e1 situated posterior to levels of setae c2 and d2, respectively. Supranal concavity circular, open posteriorly into terminal cleft. Terminal cleft long and roughly triangular, extending to level of trochanters III, length of the cleft from anterior end of concavity to bases of setae h3 155–160, greatest width at level of seta ps1 85–90. Terminal extensions on interlobar membrane wide triangular, with rounded apices, length of extensions from base of setae h3 to apices 50–55; incision between extensions wide triangular with rounded anterior extending to level of setae h3, length of incision 53–55. Setae h3 situated posterior to level of setae h2. Setae ps1 situated approximately at level of setae h2, not extending to apices of terminal extensions of interlobar membrane. Setae f2 extending slightly beyond terminal extensions. Distances between dorsal setae: c2:d2 70–80, d2:e2 85–98, e2:h3 125–140, c2:c2 245–250, h2:h2 120–140, h3:h3 100–105, ps1:ps1 85–88. Lengths of dorsal setae: c2 45–50, d2 and e2, 55–60, c3 85–100, f2 105–135, ps1 50–60.

Epimerites I fused in a Y, with stem (sternum) approximately half as long as total length of epimerites I (Fig. 2). Epimerites III rudimentary. Inner ends of epimerites IIIa free (connected by narrow sclerotized bridge in one specimen). Epimerites IVa long, almost completely enclosing coxal fields IV. Central part of coxal fields IV bearing setae 4a not sclerotized. Setae 1a short filiform, extending to level of tips of epimerites II. Setae 3a extending to level of trochanters IV. Setae 4b situated on inner ends of epimerites IIIa, represented by macrosetae and extending approximately to midlength of opisthosomal lobes. Setae ps3 and 4a approximately at same transverse level. Epiandrum (pregenital apodeme) bow-shaped, 15–20 long, 36–38 wide. Genital apparatus small, 11–13 × 22–23; aedeagus minute, barely extending from genital arch. Genital shields roughly L-shaped, with strongly irregular margins (Fig. 5E). Genital setae g on anterior ends of genital shields. Adanal shields absent, setae ps3 on soft tegument anterior to adanal suckers. Adanal suckers circular, without denticles on anterior margin, 30–33 in diameter; surrounding membrane with extensive radial striation. Distances between ventral setae: 4b:3a 32–38, 4b:g 55–73, g:ps3 30–35, ps3:h3 165–170, 4b:4b 55–75, g:g 24–28, ps3:ps3 45–50. Lengths of ventral setae: 1a 27–33, 3a 65–80, 4a 42–48, 4b 220–230, g 25–28, ps2 38–45.

Tarsi I, II, each with a long apicodorsal claw-like projection, approximately 2.5 times longer than wide at base (Fig. 5A, B). Setae s of tarsi I, II spiculiform with short filiform apex. Tibia I, II, each with spine-like ventral processes, rounded apically; setae gT of tibiae I, II spiniform with short filiform apex. Seta mG of genu I spiniform with filiform apex, seta mG of genu II long filiform. Setae cG I, II with narrow membranous extension in basal part. Tibia III without angular extension at base of solenidion φ (Fig. 5C). Tarsus III 130–135 long, slightly curved, with crest-like inner and outer margins, and with short finger-like apical projection. Tarsus IV 75–78 long, with short finger-like apical extension bearing setae d, e; seta w spiculiform with short filiform apex, extending to midlength of ambulacral disc; modified setae d, e button-like, with minute central nipple (Fig. 5D). Legs IV excluding pretarsus 250–255 long, with tarsus and distal half of tibia extending beyond lobar apices (level of setae h3). Lengths of solenidia: ω1I 38–40, ω1II 58–63, σI 82–85, σII 10–12, σIII 62–65.

Female — (range for 3 specimens) (Figs. 3, 4, 5F, G). Idiosoma, length × width, 480–485 × 325–335; length of hysterosoma 275–280. Prodorsal shield: longitudinal plate shaped as very tall trapezoid; anterior end with short longitudinal crest about 5 long; posterior margin straight, posterior end slightly extending beyond level of scapular setae, length 115–125, width at posterior margin 78–83. Scapular setae se, si situated on striated tegument at the same transverse level, bases of setae se separated by 135–140. Scapular shields as in the male. Hysteronotal shield shaped as large longitudinal plate occupying median area of hysterosoma and stretching from midlevel of scapular shields to level slightly beyond trochanters IV, anterior margin straight, lateral margins strongly attenuate to posterior end, posterior margin concave, greatest length 240–250, greatest width of anterior part 145–150, width at posterior margin 60–68. Distance between prodorsal and hysteronotal shield 10–22. Setae c2, d2, e2 filiform, shorter than trochanters III, situated on striated tegument. Setae c3 slightly longer than trochanters III. Setae d1, e1 situated on hysteronotal shield, posterior to levels of setae c2 and d2, respectively. Setae e2 approximately at level of posterior margin of hysteronotal shield. Distances between dorsal setae: c2:d2 100–110, d2:e2 125–130, e2:h3 65–70, h2:h2 60–65, h3:h3 45–58. Lengths of dorsal setae: c2, d2 25–30, setae e2 35–45, c3 55–63. Macrosetae h3 about 3/4 the length of macrosetae h2.

Epimerites I fused into a Y, with stem about 1/3 the total length of epimerites I (Fig. 4). Epigynum shaped as almost straight narrow transverse sclerite between bases of setae 4b, 25–30 wide. Apodemes of oviporus with posterior ends extending beyond trochanters III and with large lateral wing-like projection. Setae g situated at level of setae 3a or slightly anteriorly. Setae 4b long, extending to level of trochanters IV; setae 4a half as long as distance between their bases and posterior end of opisthosoma. Copulatory opening subterminal, covered by posterior ends of flaps of anal opening; spermatheca indistinct. Distances between ventral setae: 4b:3a 38–45, 4b:g 35–38, g:4a 83–88, 4a:4a 110–130. Lengths of ventral setae: 1a 35–40, 3a 45–52, 4a 80–85, 4b 160–180, g 8–10.

Legs I, II as in the male. Legs IV with tarsus and distal half of tibia extending beyond posterior margin of opisthosoma (Fig .3). Tarsi III, IV with small and rounded apicodorsal spine, 80–85 and 88–95 long, respectively (Fig. 5F, G). Trochanteral setae sRIII long, approximately equal to summed length of corresponding femur, genu and tibia. Seta w of tarsus III and setae r, w of tarsus IV thick spiculiform, with short filiform apices. Lengths of solenidia: ω1I 28–35, ω1II 50–53, σI 70–78, σII 10–11, σIII 40–48.

Remarks

Ingrassia aequinoctialis is an unmistakable species of the genus Ingrassia. This is an uncommonly large species with body length 480–500 µm, while in all other species of the genus, the length of idiosoma ranges from 250–350 µm. Additionally, Ingrassia aequinoctialis differs from other known species of the genus in the following features in both sexes: the prodorsal shield is a longitudinal plate shaped as a rectangle (male) or tall-trapezoid (female), and tarsi I, II have a long claw-like apicodorsal extension, about ¼ the total length of corresponding tarsi. In other known species of the genus, the prodorsal shield is shaped as a narrow median band with pointed posterior end or narrow longitudinal plate with an angular or rounded posterior margin; and the apicodorsal extension of tarsi I, II is shaped as a short acute or rounded spine. It is interesting to note that the strongly widened idiosoma and the convex anterior margin of the hysteronotal shield in the males of I. aequinoctialis and the hysteronotal shield enlarged anteriorly in the females of this species resemble those in mites of the genus Vingrassia Vasyukova & Mironov, 1991. However, in males of Vingrassia species, the anterior margin of the hysteronotal shield is bluntly angular (vs. rounded in I. aquinoctialis), and in females, the hysteronotal shield is roughly T-shaped (vs. lateral margins of anterior part convex).

It is interesting to note that all three species of tropicbird harbor another feather mite of extremely large size: Laminalloptes phaetontis (Fabricius, 1775) (Analgoidea: Alloptidae). The length of the idiosoma in the males and females of L. phaetontis can reach 1100 and 900 µm, respectively, making this species one of the largest among feather mites (Atyeo and Peterson 1967; Mironov and Stefan 2016). The phenomenon of the origin of very large feather mites from different families on tropicbirds remains enigmatic.

Discussion

In the discussion below, we consider taxonomic characteristics of the genera comprising the Ingrassia genus-group, their host associations and possible relationships. Authorities for host species mentioned in the Discussion are shown in Table 1. The genus Ingrassia was originally established by Oudemans (1905) for a single species, Megninia veligera Oudemans, 1904 from Tringa flavipes (Charadriiformes). Vitzthum (1921, 1922) described two new species, I. oceanica Vitzthum, 1921 from Hydrobates pelagicus (Procellariiformes) and I. tringae Vitzthum, 1922 from Calidris minuta (Charadriiformes), and transferred to this genus one more species, Analges velatus Mégnin, 1877 from Anas platyrhynchos (Anseriformes). Soon after, Hull (1934) established the genus Leptosphyra Hull, 1934 based on a single species Analges centropodus Mégnin, 1877 from Vanellus vanellus (Charadriiformes). Both Ingrassia and Leptosphyra were listed in the world checklists of feather mites summarized by Radford (1953, 1958).

For unclear reasons, Gaud ignored the genus Ingrassia in his early works on feather mites from Africa and described all ingrassine species from charadriiforms in the content of the genus Leptosphyra (Gaud 1952, 1953, 1958; Gaud and Mouchet 1959). Later on, in a taxonomic review of feather mites associated with charadriiforms in Africa, Gaud (1972) recognized the priority of the genus Ingrassia and significantly reformed its species content. Along with descriptions of 11 new species, this author enlarged the content of Ingrassia by including Leptosphyra as a subgenus and establishing the new subgenus Tectingrassia Gaud, 1972.

In their generic review of the family Xolalgidae, Gaud and Atyeo (1981a, 1981b) provided uniform diagnoses for all genera of the family and revised their species contents. In the part dealing with the subfamily Ingrassiinae, these authors (Gaud and Atyeo, 1981a) elevated the subgenera Ingrassia, Leptosphyra and Tectingrassia to full generic rank, but stressed their very close affinity to each other. Within the subfamily Ingrassiinae, these three genera can be referred to as members of the Ingrassia genus-group. Along with restriction to particular aquatic bird orders, these genera share the following combination of morphological features. In both sexes, idiosomal setae c1 are absent, tarsus I has 4 ventral setae (la, ra, wa, and s), tarsus II has two ventral setae (ra and s), and tibiae I, II have ventral spine-like projections; in males, the opisthosoma has a pair of long lobes separated by the deep terminal cleft and surrounded by lateral and interlobar membranes, the distal ends of interlobar membrane spreading onto the lobar apices form angular or semi-ovate terminal projections, legs III are hypertrophied, and the inner tips of epimerites IIIa bear the bases of setae 4b; in females, idiosomal setae ps2 and ps3 are absent.

The genera Ingrassia, Leptosphyra and Tectingrassia differ from each other by the following characters. In representatives of the genus Tectingrassia, the prodorsal shield in both sexes is shaped as a trapezoid, the posterior angles of which incorporate the bases of scapular setae se and si, while in two other genera the prodorsal shield is a narrow longitudinal band or plate with angular or rounded posterior margin and scapular setae are situated on soft tegument or on small teardrop-like sclerites. Additionally, in females of Tectingrassia, the hysteronotal shield occupies the entire median area of the hysterosoma and its anterolateral angles are close to or fused with the humeral shields. In females of Ingrassia and Leptosphyra, the hysteronotal shield is variable in size, from a very small plate (as in Ingrassia micronota Stefan, Gomez-Diaz & Mironov, 2013) to a relatively large longitudinal rectangle (as in many Ingrassia species from charadriiforms), but its anterolateral parts are never close to the humeral shields (see Gaud 1972; Gaud and Atyeo 1981a; Chirov and Mironov 1990; Vasyukova and Mironov 1991).

Mites of the genus Leptosphyra differ from those of two other genera in having a retrograde hook-like lateral projection on femora II, while in the two other genera, this projection is absent or barely developed. Representatives of the genus Ingrassia are distinguished by the absence of the aforementioned character states characterizing Leptosphyra and Tectingrassia, which are obviously apomorphic. Compared to the two latter genera, members of Ingrassia are more diverse in morphological features, in particular, in the shape of the prodorsal shield in both sexes, which varies from a narrow median band (in species from Procellariiformes) to a longitudinal plate with lateral margins parallel or divergent posteriorly (in species from Charadriiformes), and in the shape and size of the hysteronotal shield in females (see Gaud, 1972; Chirov and Mironov 1990; Vasyukova and Mironov 1986, 1991; Mironov and Proctor 2008; Stefan et al. 2013; Han et al. 2021)

Vasyukova and Mironov (1991) established Vingrassia, one more genus of this genus-group, which initially included only Ingrassia velata (Mégnin, 1877), which they transferred from Ingrassia. The genus Vingrassia, representatives of which are associated with anseriform hosts, differs from other genera of the Ingrassia genus-group in having epimerites I fused as a V and the idiosoma noticeably widened in both sexes, the hysteronotal shield with blunt-angular extension on the anterior margin in males, and T-shaped hysteronotal shield in females. In contrast, the other three genera of the group have epimerites I fused in the form of a Y with a very long stem, the anterior margin of the male hysteronotal shield is straight, wavy or concave, and the hysteronotal shield in females is of another form and in most species is not enlarged anteriorly. The second described species of Vingrassia, V. cygni Mironov & Galloway, 2002, has epimerites I fused as a Y with a very short stem (Mironov and Galloway 2002). In their review of supraspecific feather mite taxa of the world, Gaud and Atyeo (1996) did not recognize Vingrassia as a genus and considered it as a synonym of Ingrassia. Unlike these authors, we recognize Vingrassia as a morphologically distinct genus and herein include one more species with a new name Vingrassia latior (Trouessart, 1899) comb. n. (Table 1). This species, originally described from one male and named Megninia centropodos latior Trouessart, 1899, was collected from Anas bahamensis (Anseriformes) in Central America. Although it was described extremely briefly, the original description indicates that it is a wide-bodied mite, with the opisthosomal membrane forming two widely rounded lobes (Trouessart 1899: 57). These features, which are also stressed in the subspecific name ''latior″, along with association with an anseriform host, give evidence that this mite is close to other species from anseriforms, like Vingrassia velata and V. cygni, rather than to narrow-bodied Ingrassia species living on charadriiforms. Gaud and Atyeo (1981a) conventionally referred this species to the genus Ingrassia.

Phylogenetic analysis of the family Xolalgidae based on morphological characters (Mironov 2005) supported the monophyly of the cluster corresponding to the Ingrassia genus-group and clades corresponding to the genera Leptosphyra, Tectingrassia and Vingrassia. However, the genus Ingrassia in this analysis appeared to be paraphyletic, because the clades of three latter genera branched out of the crown of the Ingrassia group cluster.

Considering host associations of Ingrassia and related genera, we follow the current taxonomic concept (Gaud and Atyeo 1981a; Mironov and Galloway 2002) and recognize four genera in the Ingrassia genus-group (Table 1). This updated world checklist provides data on type hosts of mites and corresponding references containing adequate descriptions and taxonomic notes. It is clear that three genera of the group are restricted to particular orders: Leptosphyra and Tectingrassia are restricted to Charadriiformes, and Vingrassia species to Anseriformes. At the same time, mites of the genus Ingrassia are distributed on members of the Charadriiformes, Phaethontiformes, Podicipediformes, Procellariiformes and Sphenisciformes. Within Neoaves, these five orders belong to the major clade Aequornithes uniting aquatic birds except Anseriformes (e.g. Prum et al. 2015). This host distribution suggests that the genus Ingrassia probably originated on the ancestors of Aequornithes and further evolved on some lineage of this major clade, while Leptosphyra and Tectingrassia were formed and evolved on representatives of Charadriiformes. The genus Vingrassia is restricted to Anseriformes, which belongs to the archaic avian clade Galloanserae. The origin of this genus on these hosts is enigmatic, because any representatives of the family Xolalgidae are absent on galliforms, a sister lineage of anseriforms in Galloanserae. It is highly likely that primary appearance of Vingrassia on the ancestor of Anseriformes is the result of a horizontal transfer event from some representatives of Aequornithes.

Taking into account the relatively weak morphological features differentiating members of the Ingrassia genus-group, their host distributions, and the results of morphologically based phylogenetic analysis of Xolalgidae, it is possible to hypothesize that the genera Leptosphyra, Tectingrassia and Vingrassia could represent derived lineages of Ingrassia that originated within corresponding bird orders. If this turns out to be the case, perhaps it would be expedient to decrease the rank of Leptosphyra, Tectingrassia and Vingrassia and include them as subgenera in Ingrassia. The taxonomic structure of the Ingrassia genus-group certainly needs careful revision, and for this purpose, molecular based phylogenetic analysis would be quite desirable.

Acknowledgements

The authors thank Julian P. Donahue and Dr. Betty Elizabeth Anne Schreiber from the Los Angeles County Museum of Natural History Museum of Los Angeles County, who provided us with mites from their collecting activities. We are also grateful to Dmitri Logunov (The Manchester Museum, U.K.) for assisting with delivery of slide-mounted mites to SM. The study was supported in part by a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to HP. 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.

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