# A new species of the feather mite genus Allodectes (Acariformes: Proctophyllodidae) from the ruby-throated hummingbird, Archilochus colubris (Apodiformes: Trochilidae), in Canada

Sergey V Mironov, Terry D Galloway

► To cite this version:

Sergey V Mironov, Terry D Galloway. A new species of the feather mite genus Allodectes (Acariformes: Proctophyllodidae) from the ruby-throated hummingbird, Archilochus colubris (Apodiformes: Trochilidae), in Canada. Acarologia, 2023, 10.24349/vejw-1k8v. hal-04142816

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# A new species of the feather mite genus *Allodectes* (Acariformes: *Proctophyllodidae*) from the ruby-throated hummingbird, *Archilochus colubris* (Apodiformes: *Trochilidae*), in Canada

Sergey V. Mironov IDª, Terry D. Galloway IDᵇ

## Original research

### ABSTRACT

A new feather mite species, *Allodectes archilochus* sp. n. (Pterodectinae: Rhamphocaulini) is described from the ruby-throated hummingbird, *Archilochus colubris* (Apodiformes: Trochilidae), in Manitoba, Canada. This is the first record of the genus *Allodectes* on hummingbirds in the northern part of North America. The new species is close to *A. thaluraniae* Hernandes, 2017, and distinguished from the latter in the following features: in both sexes, the anterior margin of the hysteronotal shield is deeply concave; in males, setae *d2* are situated at the midlevel of metapodosomal sclerites, and tarsus IV is without extension on the inner margin; in females, the anterolateral extensions of the prodorsal shield do not extend to the lateral margins of propodosoma, and the anteromedian area of the hysteronotal shield is covered with dense striation. A checklist of the 15 currently known *Allodectes* species is provided.

**Keywords** Astigmata; feather mites; Proctophyllodidae; *Allodectes*; systematics; Trochilidae

**Zoobank** http://zoobank.org/FF8C2172-101E-4367-BC12-B6D8F2C65EF6

### Introduction

Feather mites of the genus *Allodectes* Gaud & Berla, 1963 (Proctophyllodidae: Pterodectinae) belong to the tribe Rhamphocaulini, which is restricted to hummingbirds (Apodiformes: Trochilidae) (Park & Atyeo 1971b, 1972a, 1972b 1973a, 1973b, 1974a, 1974b, 1975; Mironov 2009; Hernandes 2013; Klimov et al. 2017). Of eight genera currently referred to the tribe Rhamphocaulini, the genus *Allodectes* and three additional genera, *Rhamphocaulus* Park & Atyeo 1971, *Schizodectes* Park & Atyeo 1972, and *Sclerodectes* Park & Atyeo 1973, inhabit quill cavities of flight feathers, mainly the primaries, in contrast to all remaining representatives of Proctophyllodidae which occupy surfaces of the flight and tail feathers (Gaud & Atyeo 1996). Sometimes, individuals of these quill-inhabiting genera can be found outside quills in the basal part of rachis (Park & Atyeo 1973a). This allows the possibility of collecting these mites from dry museum skins by a scraping technique commonly used for collecting mites inhabiting flight feathers (Gaud & Atyeo 1996). A distinct morphological feature of rhamphocauline genera which live inside quills is the extensively sclerotized coxal fields I-IV, though the functional significance of this feature is not clear.

Received 30 May 2023
Accepted 20 June 2023
Published 27 June 2023

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

Academic editor
Akashi Hernandes, Fabio

https://doi.org/10.24349/vejw-1k8v

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

<figure><img src="image_6.png" /></figure>

Mironov S. V. and Galloway T. D.

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

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

<footnote>ᵇDepartment of Entomology, University of Manitoba, Winnipeg, Manitoba, R3T 2N2, Canada.</footnote>
The genus *Allodectes* was originally established for one species, *Proctophyllodes (Alloptes)*
*norneri* Trouessart, 1885 (Gaud & Berla 1963). Park and Atyeo (1972a) described ten
new *Allodectes* species from Mexico, Central and South Americas, and established a new
subfamily, Allodectinae, based on this genus. Further, this subfamily was synonymized with
Rhamphocaulinae, based on the genus *Rhamphocaulus* Park & Atyeo, 1971, representatives of
which also live in quills of hummingbirds (Gaud & Atyeo 1996). Mironov (2009), based on
the cladistic analysis of morphological characters, reduced the subfamily Rhamphocaulinae to
the rank of tribe in the subfamily Pterodectinae. Additionally, four genera, *Syntomodectes* Park
& Atyeo, 1973, *Toxerodectes* Park & Atyeo, 1973, *Trochilodectes* Park & Atyeo, 1971, and
*Xynonodectes* Park & Atyeo, 1975, also restricted to hummingbirds and considered by Gaud
and Atyeo (1996) as the members of Pterodectinae rather than Rhamphocaulinae, were placed
by Mironov (2009) in the tribe Rhamphocaulini. This taxonomic decision was well supported
by the phylogenetic study of proctophyllodid mites based on molecular data (Klimov *et al.*
2017).

In the past fifty years, only three new *Allodectes* species have been described (Černý 1974; Hernandes 2013), increasing the number of known species to 14 (Table 1). This paper is a continuation of our studies of feather mites in Canada (e.g., Mironov & Galloway 2002, 2019, 2021; Galloway *et al.* 2014), and describes a new *Allodectes* species from the ruby-throated hummingbird collected in Manitoba, Canada. This is the first description of an *Allodectes* species from a hummingbird in the northern part of North America.

**Material and methods**

Frozen bird specimens used for the collections of feather mites were obtained from the
Wildlife Haven (Manitoba Wildlife Rehabilitation Organization, MWRO, Île des Chênes,
Manitoba, Canada; in bird specimen numbers are identified as /CEN/) and the Prairie Wildlife
Rehabilitation Centre (PWRC, St. Adolphe, Manitoba, Canada). All birds from the MWRO
and PWRC were the casualties of various accidents or infections.

<table><caption>**Table 1** Checklist of *Allodectes* species of the World.</caption><thead><tr><td>Mite species</td><td>Type host</td><td>Locality</td><td>Reference</td></tr></thead><tbody><tr><td>A. amadoni Park & Atyeo, 1972*</td><td>Colibri t. thalassinus (Swainson, 1827)</td><td>Mexico</td><td>Park & Atyeo 1972b</td></tr><tr><td>A. amaziliae Park & Atyeo, 1972*</td><td>Saucerottia tobaci erythronotus (Lesson, RP, 1829)</td><td>Trinidad and Mexico</td><td>Park & Atyeo 1972b<br>Park & Atyeo 1972b</td></tr><tr><td>A. amplissimus Park & Atyeo, 1972*</td><td>Anthracothorax p. prevostii (Lesson, RP, 1832)</td><td>Canada:</td><td>present work</td></tr><tr><td>A. archilochus sp. n. *</td><td>Archilochus colubris (Linnaeus, 1758)</td><td>Brazil</td><td>Park & Atyeo 1972b</td></tr><tr><td>A. bipeltatus Park & Atyeo, 1972</td><td>Phaethornis philippi (Bourcier, 1847)</td><td>Panama</td><td>Park & Atyeo 1972b</td></tr><tr><td>A. elatarcus Park & Atyeo, 1972</td><td>Phaethornis longirostris cephalus (Bourcier & Mulsant, 1848)<br>(=P. superciliosus cassinii)</td><td></td><td></td></tr><tr><td>A. longisetosus Park & Atyeo, 1972*</td><td>Heliomaster constantii leocadiae (Bourcier & Mulsant, 1852)</td><td>Mexico,<br>Colombia</td><td>Park & Atyeo 1972b<br>Park & Atyeo 1972b</td></tr><tr><td>A. nanus Park & Atyeo, 1972</td><td>Phaethornis s. striicularis Gould, 1854 (=P. longuemareus<br>striicularis)</td><td></td><td></td></tr><tr><td>A. norneri (Trouessart, 1885)*</td><td>Aglaiocercus kingii mocoa (Delattre & Bourcier, 1846)</td><td>Bolivia, Peru</td><td>Trouessart 1885; Gaud & Berla<br>1963; Park & Atyeo 1972b</td></tr><tr><td>A. paulicaulus Park & Atyeo, 1972*</td><td>Pampa c. curvipennis (Deppe, 1830)</td><td>Mexico</td><td>Park & Atyeo, 1972b</td></tr><tr><td>A. prolaticaulus Park & Atyeo, 1972</td><td>Phaethornis longirostris cephalus (Bourcier & Mulsant, 1848)</td><td>Panama</td><td>Park & Atyeo 1972b</td></tr><tr><td>A. sejugaspis Hernandes, 2013*</td><td>Thalurania glaucopis (Gmelin, JF, 1788)</td><td>Brazil: Paraná</td><td>Hernandes 2013</td></tr><tr><td>A. similis Černý, 1974</td><td>Chionomesa fimbriata (Gmelin, JF, 1788)</td><td>Surinam</td><td>Černý 1974</td></tr><tr><td>A. thaluraniae Hernandes, 2013*</td><td>Thalurania glaucopis (Gmelin, JF, 1788)</td><td>Brazil: Paraná</td><td>Hernandes 2013</td></tr><tr><td>A. wetmorei Park & Atyeo, 1972*</td><td>Phaethornis m. mexicanus Hartert, EJO, 1897</td><td>Mexico</td><td>Park & Atyeo 1972b</td></tr></tbody></table>

Remark: * – species of the group characterized by the absence of apical claw-like processes on tarsi I, II in both sexes.
The washing technique for collection of feather mites from frozen bird specimens was described in detail in our previous publications (Mironov and Galloway 2002, 2019, 2021). Collected mites fixed in 95% ethanol were cleaned and softened in 5-10% lactic acid at room temperature for 2-3 days, then mounted on microscope slides using Hoyer's medium (Krantz & Walter 2009), and dried in an oven at 60 °C for 7 days. Drawings and measurements were made with a Leica microscope (DM2500, Leica Microsystems, Inc.) equipped with differential interference contrast optics (DIC) and a *camera lucida*.

Description of the new species follows the format proposed by Hernandes (2013) for the genus *Allodectes*. General morphological terms and idiosomal chaetotaxy follow Gaud & Atyeo (1996), with minor corrections for coxal setae by Norton (1998). Leg chaetotaxy is that of Grandjean (1939). Regarding the homology of opisthosomal chaetotaxy in males, we follow the original concept of Park and Atyeo (1972a) and treat the median pair of setae on the posterior margin of the anal field as setae *ps3*, and the long filiform setae arising at bases of macrosetae *h2* as setae *h3*. Hernandes (2013) treated these setae as *h3* and *ps2*, respectively. All measurements are in micrometers (μm).

Scientific names of birds and supraspecific classification follow Gill *et al.* (2023). The abbreviations used in accession numbers for specimens and depositories are as follows: CNC – Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Eastern Cereal and Oilseeds Research Centre, Ottawa, Ontario, Canada; WRME – J.B. Wallis/R.E. Roughley Museum of Entomology (Department of Entomology, University of Manitoba, Winnipeg, Manitoba, Canada); ZISP – Zoological Institute of the Russian Academy of Sciences (Saint Petersburg, Russia).

## Systematics

**Family Proctophyllodidae Trouessart & Mégnin, 1884**

**Subfamily Pterodectinae Park & Atyeo, 1971**

**Tribe Rhamphocaulini Park & Atyeo, 1971**

**Genus Allodectes Gaud & Berla, 1963**

Type species: *Proctophyllodes* (*Alloptes*) *norneri* Trouessart, 1885, by original designation.

**Allodectes archilochus Mironov & Galloway sp. n.**

Zoobank: 063EF62A-61D0-42C1-9298-AC02082BA64D

(Figures 1-4)

### Type material

From *Archilochus colubris* (Linnaeus, 1758) (Apodiformes: Trochilidae), CANADA, Manitoba: male holotype, 2 male and 3 female paratypes (RTHU/408/CEN/12, SM 2253), Vassar, 3 May 2012, coll. T.D. Galloway and S.R. Repa; other paratypes: 2 males and 3 females (RTHU/555/CEN/01, SM 526), Winnipeg, 18 Jun. 2001 coll. T.D Galloway; 2 females, (RTHU/861/CEN/08, SM 1620), Winnipeg, 30 Aug. 2008, coll. T.D. Galloway and D. Holder; 2 females (RTHU/555/PWRC/13, SM 2391), Winnipeg, 26 Aug. 2013, coll. T.D. Galloway; 1 female (RTHU/866/CEN/14, SM 2503), Winnipeg, 3 Sep. 2014, coll. T.D. Galloway.

**Depository** — Holotype, 2 male and 3 female paratypes – CNC; remaining paratypes – WRME and ZISP.

### Description

**Male** — (holotype, range for 4 paratypes in parentheses) (Figures 1, 3A–D, 4A). Idiosoma, length × width, 325 (330–345) × 195 (195–210), length of hysterosoma 225 (225–235).
<figure><img src="image_7.png" /><figcaption>**Figure 1** *Allodectes archilochus* sp. n., male: A – dorsal view, B – ventral view.</figcaption></figure>

Prodorsal shield: anterolateral extensions reaching margins of propodosoma and fused with epimerites Ia, posterior margin strongly concave, greatest length 105 (100–110), length along midline 88 (85–90), width at posterior margin 148 (150–160), anterior part with sparse longitudinal striae, posterior part with sparse short transverse striae. *Setae* ve minute. Scapular setae *se* separated by 65 (65–70). Scapular shields present dorsally, with inner margin shaped as
blunt angle. Sejugal sclerites on striated tegument between prodorsal and anterior hysteronotal
shields absent. Humeral shields well developed, with setae c2 on their anterior margins.
Subhumeral setae c3 spiculiform, 50 (43–50) long. Metapodosomal shields present. Distance
between prodorsal and hysteronotal shields long midline 32 (32–35). Hysteronotal shield:
greatest length 220 (220–235), width in anterior margin 140 (140–155), anterior margin
concave, almost entire surface except for lateral and posterior margins covered with dense,
mostly longitudinal striation. Supranal concavity ovate and well outlined. Opisthosoma
gradually narrowed posteriorly; posterior end generally rounded; opisthosomal lobes small;
terminal cleft slit-like, lateral margins almost touching, 12 (12–17) long. Posterolateral margins
of lobes with narrow terminal membranes having smooth and convex free margin. Lateral
margins of opisthosoma, posterior to bases of setae ps2, with small triangular lateral membranes.
Setae d2 at midlength of metapodosomal shields. Setae h1 slightly anterior to level of setae
e2. Setae h2 represented by macrosetae approximately as long as body length. Setae f2 at
level of setae h2. Setae ps2 thin spiculiform, 20 (18–23) long, situated on lateral margins of
opisthosoma. Setae h3 setiform, 40 (40–45) long, situated posterolateral to bases of setae h2.
Setae ps1 minute filiform, situated at level of setae h3, close to slit-like terminal cleft. Distance
between dorsal setae: c2:d2 90 (90–98), d2:e2 85 (88–92), e2:h3 32 (32–38), d1:d2 60 (55–65),
e1:e2 25 (22–28), h1:ps2 25 (25–32), h2:h2 28 (26–28), h3:h3 38 (37–40), ps2:ps2 58 (57–60).

Epimerites I fused into a narrow V, with posterior tips connected by thin commissure. Coxal
fields I partly sclerotized, lateral 1/3 of these fields bordering on epimerites II covered with
striated tegument; coxal fields II–IV entirely sclerotized. Transventral sclerite, connecting
inner borders of coxal fields III, 17 (13–18) along midline. Anterior end of epiandrum fused
with transventral sclerites; free branches of epiandrum thin and acute, laterally flanking anterior
part of genital apparatus. Rudimentary sclerite rEpIIa absent. Epimerites IVa present, fused
with coxal fields IV, not extending to bases of setae 4a. Base of genital apparatus at midlength
of trochanters III; genital arch short, not extending beyond transventral sclerite, 32 (32–35) ×
25 (25–28). Aedeagus (from anterior bend to apex) approximately half as long as genital arch,
16 (17–20) long. Genital papillae at midlength of genital arch. Genital shields present, bearing
bases of setae g. Adanal suckers 17 (17–19) in diameter, corolla with 9–11 acute denticles.
Opisthoventral shields well developed, strongly widened anterior to anal opening and almost
completely encircling anal field. Setae 1a off sclerotized areas of coxal fields I; setae 4b
anterior to setae 3a; setae g situated on small genital shield anterior to level of setae 4a, setae
ps3 on posterior margin of anal field and close to each other. Distance between ventral setae:
4b:3a 12 (12–18), 4b:g 50 (43–50), g:4a 12 (16–17), g:ps3 125 (125–135), ps3:ps3 8 (8–10).

Tarsi I, II without apical claw-like process; tarsi III, IV with apical claw-like process.
Femora I, II with small ventral crest near base of setae vF. Solenidion ω1 of tarsus II slightly
longer than solenidion ω1 of tarsus I. Solenidion σ of genu I about 1.5 times longer than this
segment, situated at its midlength. Genual setae cGI, II, and mGI, II, setiform. Seta sR of
trochanter III spiculiform, 50 (50–54) long. Genu IV with small spine-like process on paraxial
margin and with longitudinal crest along lateral margin. Tarsus III 38 (38–40) long. Tarsus IV
50 (50–52) long, without extension on inner margin; setae e vestigial, represented by alveolus
near apex of segment, seta d indistinct. Length of solenidia: σI 56 (56–60), σIII 40 (40–42),
φIII 72 (65–78), φIV 48 (46–50).

**Female** — (range for 10 paratypes) (Figures 2, 3E–F, 4B). Idiosoma, length × width, 415–445 × 175–195. Prodorsal shield: anterolateral extensions not reaching lateral margins of propodosoma and not fused with epimerites Ia, posterior margin strongly concave, greatest length 120–125, along midline 110–115, width at posterior margin 145–165, anterior part with sparse longitudinal striae, posterior part without ornamentation. Setae *ve* minute. Scapular setae *se* separated by 75–82. Scapular shields narrow, with concave inner margins. Sejugal sclerites between prodorsal and anterior hysteronotal shields absent. Humeral shields narrow, bearing setae *c2* on their anterior ends. Setae *c3* narrowly lanceolate, 20–24 × 4.5–5. Distance between prodorsal and hysteronotal shields along midline 33–40. Hysteronotal shield completely split into anterior and lobar pieces, dorsally separated by narrow arched forward band of
<figure><img src="image_8.png" /><figcaption>**Figure 2** *Allodectes archilochus* sp. n., female: A – dorsal view, B – ventral view.</figcaption></figure>

soft tegument. Anterior hysteronotal shield: anterior and posterior margins deeply concave, greatest length 205–225, width at anterior margin 130–150, anterior and median parts with dense striation, posterolateral areas with several small circular lacunae or without them. Lobar region: anterior margin strongly convex, surface with small circular lacunae, greatest length 105–115, width 88–92. Supranal concavity small, circular, poorly sclerotized. Setae *h1* and *f2*
<figure><img src="image_9.png" /><figcaption>**Figure 3** *Allodectes archilochus* sp. n., legs: A–C – legs I–III of male, D – genu, tibia and tarsus IV of male, E – leg III of female, F – tibia and tarsus IV of female.</figcaption></figure>
<figure><img src="image_10.png" /><figcaption>**Figure 4** *Allodectes archilochus* sp. n., details: A – opisthosoma of male, ventral view, B – spermatheca and spermataducts. Abbreviations: co – copulatory opening, hs – head of spermatheca, os – opisthoventral shield, pd – primary spermataduct, sd – secondary spermataduct, tm – terminal membrane, tvs – transventral sclerite.</figcaption></figure>

in trapezoidal arrangement. Terminal cleft shaped as a narrow inverted U, 58–63 long, 7–10 wide. Macrosetae *h2* strongly widened in basal half, 120–150 long, 10–12 wide. Setae *h3* setiform, 67–70 long, about half the length of terminal appendages. Distance between dorsal setae: *c2:d2* 110–125, *d2:e2* 78–87, *e2:h2* 42–45, *h2:h3* 38–48, *d1:d2* 60–75, *e1:e2* 25–35, *h1:h2* 38–40, *h2:ps1* 23–28, *h2:h2* 78–82, *h1:h1* 18–20.

Epimerites I fused in a narrow V, with posterior tips connected with thin commissure. Epimerites IVa present, heavily sclerotized. Coxal fields I partly sclerotized, lateral 1/3 bordering on epimerites II covered with striated tegument; coxal fields II–IV entirely sclerotized. Epigynum without lateral extensions, greatest width 68–78. Apodemes of oviporus separated from epimerites IIIa. Translobar apodemes fused to each other anterior to terminal cleft. Copulatory opening situated ventrally, posterior to anal opening, and covered with posterior ends of anal flaps. Primary spermataduct with small ampuliform enlargement near head of spermatheca, secondary spermataducts 25–30 long (Figure 4B). Setae *1a* off sclerotized areas
of coxal fields I. Pseudanal setae ps2 and ps3 setiform, in trapezoid arrangement; setae ps2 at midlength of anal opening; distances between these setae: ps2:ps2 47–50, ps3:ps3 15–21, ps2:ps3 18–22.

Femur II with small ventral crest near base of setae vF; other leg segments I, II without processes. Solenidia of genu and tarsi I, II as in the male. Genual setae cGI, II, and mGI, II setiform. Seta sR of trochanter III narrowly lanceolate 28–30 long, 3–3.5 wide. Length of solenidia: σI 53–58, σIII 21–23, φIII 56–60, φIV 32–35.

Differential diagnosis

The new species, *Allodectes archilochus sp. n.*, belongs to the species group characterized by the absence of the claw-like apical processes on tarsi I, II in both sexes, and in having the vestigial terminal membrane in males (Park & Atyeo 1972b) (Table 1, species marked with asterisk). In this group, the new species is most similar to *A. thaluraniae* Hernandes, 2013 in having, in males, the median area of the hysteronotal shield being striated, setae c3 and sRIII subequal in length, the spine-like process on the distal margin of genu III, and the relatively elongate idiosoma (1.6–1.7 times longer than wide). *Allodectes archilochus sp. n.* is distinguished from *A. thaluraniae* in the following features: in both sexes, the anterior margin of the hysteronotal shield is deeply concave, and the lateral areas of coxal fields I (approximately 1/3) are not sclerotized, and covered with soft striated tegument; in males, setae d2 are at the midlevel of the metapodosomal sclerites, and tarsus IV lacks the blunt-angular extension on the inner margin; in females, the anterior part of the prodorsal shield bears sparsely disposed longitudinal striae, the anterolateral extensions of this shield do not extend to the lateral margins of propodosoma, and the anteromedian area of the hysteronotal shield is covered with dense striation. In both sexes of *A. thaluraniae*, the anterior margin of the hysteronotal shield is almost straight or slightly concave, the lateral areas of coxal fields I are extensively sclerotized, and only narrow bands along epimerites II are covered with soft tegument; in males, setae d2 are at the level of posterior ends of the metapodosomal sclerites, and tarsus IV has a blunt-angular extension on the inner margin near the base of seta w; in females, the prodorsal shield lacks striation, the anterolateral extensions of this shield extend to the lateral margins of the propodosoma, and the anterior half of the hysteronotal shield is without striation.

Etymology

The specific epithet is taken from the generic name of the type host and is a noun in apposition.

Acknowledgements

The author thank Fabio A. Hernandes (Universidade Federal de Santa Catarina, Florianópolis, Brazil) for his useful comments on the manuscript. The taxonomic part of the study was supported by the Ministry of Science and Higher Education of the Russian Federation (project No. 122031100263-1) for SM. This research was partly funded by National Science and Engineering Research Council of Canada Discovery Grants to TDG. Special thanks go to the dedicated staff at Wildlife Haven and the Prairie Wildlife Rehabilitation Hospital for the care with which they treated hummingbirds that ultimately were a component in this study. Scientific Permits to allow possession and examination of hummingbirds were issued by the Canadian Wildlife Service, Environment and Climate Change Canada / Government of Canada. The continued support of the Department of Entomology and the Faculty of Agricultural and Food Sciences at the University of Manitoba is greatly appreciated.

ORCID

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

Terry D. Galloway https://orcid.org/0000-0002-0124-8601
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