# A new feather mite species of the genus Temnalges (Acariformes: Psoroptoididae) from coots, Fulica spp. (Gruiformes: Rallidae)

Sergey V Mironov, Simon Haarder, Terry D Galloway

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Sergey V Mironov, Simon Haarder, Terry D Galloway. A new feather mite species of the genus Temnalges (Acariformes: Psoroptoididae) from coots, Fulica spp. (Gruiformes: Rallidae). Acarologia, 2024, 64 (4), pp.1106-1116. 10.24349/i1su-eqk9. hal-04764579

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# A new feather mite species of the genus *Temnalges* (Acariformes: Psoroptoididae) from coots, *Fulica* spp. (Gruiformes: Rallidae)

Sergey V. Mironov IDª, Simon Haarder IDᵇ, Terry D. Galloway IDᶜ

## Original research

### ABSTRACT

A new feather mite species, *Temnalges fulicae* sp. n. (Psoroptoididae: Pandalurinae), is described from two species of coots, *Fulica americana* Gmelin, 1789 (type host), in Canada, and *F. atra* Linnaeus, 1758, in Denmark. The new species is close to other species of the genus associated with rails (Rallidae), especially to *T. megalonyx* Gaud, 1968, in having a long spur-like process on tibia III in males, and the prodorsal and hysteronotal shields separated by a wide area of striated tegument in females. Males of *Temnalges fulicae* are most clearly distinguished from *T. megalonyx* in having the opisthosoma rounded and without terminal and lateral clefts, and the adanal shields narrow and short. In females, the prodorsal shield has a convex posterior margin, the hysteronotal shield is roughly rectangular, and setae 4a extend to the bases of setae ps3. Additionally, we provide a world checklist and a key to *Temnalges* species.

**Keywords** Astigmata; feather mites; *Temnalges*; systematics; new species; Rallidae

**Zoobank** http://zoobank.org/53EF0F9C-F027-411F-A29B-CB0263A186E1

### Introduction

Feather mites of the genus *Temnalges* Gaud & Atyeo, 1967 (Analgoidea: Psoroptoididae: Pandalurinae) are small-sized psoroptoidids, about 200–300 micrometers long (Gaud and Atyeo 1967; Gaud 1968). As for most psoroptoidids, these mites live mainly in the downy parts of the contour feathers of their avian hosts, although they can be found on the distal parts of the contour feathers and on the flight feathers, and can even occur on the skin (Mironov 2011; Mironov *et al.* 2011, 2018; Hernandes *et al.* 2014). To date, this genus has included only seven species (Gaud and Atyeo 1967, 1982; Gaud 1968, 1976; Shereef and Rakha 1982; Hernandes and Mironov 2015). Although known species of *Temnalges* are not numerous, they are associated with avian orders being rather distant from each other: Anseriformes, Charadriiformes, Gruiformes, Opisthocomiformes, and Pelecaniformes (Table 1). It is interesting to note that one of the hosts, the hoatzin, *Opisthocomus hoazin* (Müller) (Opisthocomidae), bears two co-infesting species of this genus (Hernandes and Mironov 2015). The only common feature of these host orders is their ecological association with aquatic biotopes. Erratic distribution of *Temnalges* species among so many different avian orders and, at the same time, a small number of currently described species lead us to believe that this genus probably has an ancient origin and its extant fauna could actually be much more numerous and diverse.

Received 17 July 2024
Accepted 08 October 2024
Published 30 October 2024

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

Academic editor
Akashi Hernandes, Fabio

https://doi.org/10.24349/i1su-eqk9

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

<figure><img src="image_2.png" /></figure>

Mironov S. V. *et al.*

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 Ornebjergvej 43, 4760 Vordingborg, Denmark.</footnote>

<footnote>ᶜ Department of Entomology, University of Manitoba, Winnipeg, Manitoba, R3T 2N2, Canada.</footnote>
In the present work, we describe a new *Temnalges* species from two species of coots, *Fulica americana* Gmelin and *F. atra* Linnaeus, in Canada and Denmark, respectively. This species was first reported as *Temnalges* sp. by Galloway *et al.* (2014), along with five additional species of feather mites infesting *F. americana* in Canada. We also provide a checklist and a key to currently known species of the genus.

## Material and methods

Specimens for this study were collected from dead specimens of American coots by TDG in Manitoba, Canada and Eurasian coots by SH in Denmark. American coots used for the collections of feather mites were obtained from the Wildlife Haven (Manitoba Wildlife Rehabilitation Organization, MWRO, Île des Chênes, Manitoba, Canada; case numbers for bird specimen numbers are identified below as /CEN/). Specimens from American coots were collected by the washing technique for extracting feather mites from frozen birds described in detail in our previous publications (Mironov and Galloway 2019, 2021). Mites from road-killed Eurasian coots were collected using a simplified version of the above-mentioned technique (birds were only washed once and sieve diameter was 63 micrometers). Mites extracted from birds were fixed in 70-95% ethanol and then were mounted on microscope slides using Hoyer's medium according to standard techniques used for small mites (Krantz and Walter 2009). Drawings and measurements (in micrometers, µm) were made with a Leica microscope (DM2500, Leica Microsystems, Inc.) equipped with differential interference contrast optics (DIC), an ocular with calibrated ruler, and a *camera lucida*.

The species description follows the modern format for psoroptoidids (Mironov 2004; Mironov *et al.* 2011, 2018; Hernandes and Mironov 2015). Idiosomal chaetotaxy follows Gaud and Atyeo (1996), with additions by Norton (1998); leg chaetotaxy is that of Grandjean (1939). To construct the key to *Temnalges* species, we examined specimens deposited in the collection of the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (*T. atelodiscus* Hernandes & Mironov, 2015, *T. hoazin* Hernandes & Mironov 2015, and *T. mesalgoides* Gaud & Atyeo, 1967). Regarding other previously known species (Table 1), we relied on previous taxonomic descriptions (Gaud 1968, 1976; Shereef and Rakha 1982). Since descriptions of some previously described species are a bit incomplete from the modern taxonomic point of view, in the key, we used only reliable characters, which were exactly described and/or illustrated.

Systematics and scientific names of birds follow the IOC World Bird List, v 14.1 (Gill *et al.* 2024). 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,

<table><caption>**Table 1** Checklist, host associations and distribution of *Temnalges* species.</caption><thead><tr><th>Mite species</th><th>Type host</th><th>Host family</th><th>Locality</th><th>Reference</th></tr></thead><tbody><tr><td><i>T. amalicus</i> Shereef & Rakha, 1982</td><td><i>Plegadis falcinellus</i> (Linnaeus)</td><td>Threskiornithidae</td><td>Egypt</td><td>Shereef and Rakha 1982</td></tr><tr><td><i>T. atelodiscus</i> Hernandes & Mironov, 2015</td><td><i>Opisthocomus hoazin</i> (Müller)</td><td>Opisthocomidae</td><td>Brasil: Pará, Rondônia</td><td>Hernandes and Mironov 2015</td></tr><tr><td><i>T. bothrioplax</i> (Gaud, 1958)</td><td><i>Chlidonias niger</i> (Linnaeus)</td><td>Laridae</td><td>Morocco</td><td>Gaud 1958, 1976</td></tr><tr><td><i>T. fulicae</i> sp. n.</td><td><i>Fulica americana</i> Gmelin, JF</td><td>Rallidae</td><td>Canada: Manitoba</td><td>Present work</td></tr><tr><td>“</td><td><i>F. atra</i> Linnaeus</td><td>Rallidae</td><td>Denmark</td><td>Present work</td></tr><tr><td><i>T. hoazin</i> Hernandes & Mironov, 2015</td><td><i>Opisthocomus hoazin</i> (Müller)</td><td>Opisthocomidae</td><td>Brasil: Pará</td><td>Hernandes and Mironov 2015</td></tr><tr><td><i>T. mahranus</i> Shereef & Rakha, 1982</td><td><i>Spatula querquedula</i> (Linnaeus)</td><td>Anatidae</td><td>Egypt</td><td>Shereef and Rakha 1982</td></tr><tr><td><i>T. megalonyx</i> Gaud, 1968</td><td><i>Zapornia flavirostra</i> (Swainson)</td><td>Rallidae</td><td>Cameroon</td><td>Gaud 1968</td></tr><tr><td><i>T. mesalgoides</i> Gaud & Atyeo, 1967*</td><td><i>Porphyrio madagascariensis</i> (Latham)*</td><td>Rallidae</td><td>South Africa: Transvaal</td><td>Gaud and Atyeo 1967; Gaud 1968</td></tr></tbody></table>

Remark. *— Gaud (1968) mentioned this host under the name *Porphyrio porphyrio* (Linnaeus).
University of Manitoba, Winnipeg, Manitoba, Canada); ZISP—Zoological Institute of the
Russian Academy of Sciences (Saint Petersburg, Russia).

**Systematics**

**Family Psoroptoididae Gaud, 1958**

**Subfamily Pandalurinae Gaud & Atyeo, 1982**

**Genus *Temnalges* Gaud & Atyeo, 1967**

*Temnalges fulicae* sp. n.

Zoobank: OF0BCCF6-24B5-4A27-A5DE-B102296F39C1

(Figures 1–5)

**Type material**

**Holotype** — From *Fulica americana* Gmelin, 1789 (Gruiformes: Rallidae) AMCO/1297/CEN/98, CANADA, Manitoba, Richer, 15 August 1998, coll. T.D. Galloway: male holotype, 6 male and 7 female paratypes; other paratypes: 10 males and 4 females, AMCO/1710/CEN/99, CANADA, Manitoba, Winnipeg, 23 October 1999, coll. T.D. Galloway.

**Depository** — Holotype, 4 male and 4 female paratypes – CNC; remaining paratypes – WRME and ZISP.

**Additional material** — From *Fulica atra* Linnaeus, 1758 (Gruiformes: Rallidae), DEN-MARK, Zealand, Jyllinge, 55°44′18.3″ N 12°07′41.7″ E, 16 February 2023, coll. S. Haarder: 3 males; DENMARK, Zealand, Kirke Hvalsø, 55°37′20.8″ N 11°49′21.0″ E, 18 February 2019, coll. S. Haarder: 6 males, 4 females, and 1 tritonymph.

**Description**

**Male** — (holotype, range for 10 paratypes in parenthesis) (Figures 1, 2, 5A-E). Idiosoma, length × width, 220 (215–225) × 150 (140–150), length of hysterosoma 145 (130–150). Prodorsal shield: shaped as longitudinal rectangle slightly enlarged posteriorly, length 68 (65–70), width at posterior margin 35 (30–35); lateral margins without extensions, posterior margin straight; posterior corners rounded, longitudinal ridges divergent posteriorly and as long as 3/4 of the shield. Setae *vi*, *ve* absent. Setae *si*, *se* off prodorsal shield, bases of setae *se* separated by 48 (45–48). Distance between prodorsal and hysteronotal shields 5 (5–8). Hysteronotal shield: length from anterior margin to level of setae *h3* 145 (135–150), width at anterior margin 68 (63–70); anterior margin convex, anterior corners widely rounded; lateral margins straight. Setae *c1*, *d1* absent. Hysteronotal gland openings *gl* on hysteronotal shield near its lateral margins and closer to level of setae *d2* than to setae *e2*. Setae *e1* closer to level of setae *e2* than to openings *gl*. Setae *e2* and *d2* short filiform, shorter than the half-width of hysteronotal shield. Supranal concavity long and narrow, 38 (32–40) long. Opisthosoma widely rounded, opisthosomal lobes not expressed, bases of setae *f2*, *h2*, *ps2*, *h3* and *ps1* situated on posterolateral margins of opisthosoma *f2*, terminus of opisthosoma with small scarcely distinct concavity between bases of setae *ps1*. Distances between idiosomal setae: *c2:* *d2* 28 (25–28), *d2:* *e2* 73 (63–705), *e2:* *h3* 40 (38–42), *h2:* *h2* 50 (49–52), *h3:* *h3* 25 (25–28), *ps1:* *ps1* 10 (10–12), *ps2:* *ps2* 40 (37–40), *f2:* *f2* 60 (57–60), *d2:* *d2* 60 (57–62), *e2:* *e2* 47 (45–50). Lengths of dorsal setae: *c2* 102 (95–105), *c3* 80 (65–80), *d2* 22 (20–25), *e2* 20 (15–20), *f2* 75 (65–75), *ps1* 75 (75–80), *ps2* 55 (55–65).

Epimerites I free, bow-shaped, with divergent posterior ends. Tips of epimerites II distant from epimerites III. Anterior tips of combined epimeral sclerites IIIa+IV not enlarged and bear bases of setae 4b. Coxal fields I–IV open. Epimerites IV with sclerotized band flanking bases of trochanters IV and bearing setae 4a. Genital apparatus situated at level of anterior end of trochanters III, genital arch 12.5 (10–12.5) × 15 (14–16); aedeagus minute, shorter than genital arch (Figure 2). Epiandrum (pregenital apodeme) as low bow, 10 (7.5–10) long and 17 (15–17)
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Temnalges fulicae* sp. n.: dorsal view of male.</figcaption></figure>

wide. Adanal shield represented by a pair of narrow and oblique sclerites. Setae 3a near tips of epimerites III, represented by macrosetae 125 (110–130) long. Setae 3a and 4b approximately at the same transverse level. Setae g between levels of anterior and posterior genital papillae. Setae 4a posterior to level of setae g. Diameter of adanal suckers 12.5 (11–12.5). Distances between ventral setae: 4b:4b 24 (22–26), 4b:g 25 (20–25), 4b:4a 28 (25–30), g:ps3 55 (55–58), ps3:h3 23 (22–24).

Tarsi I, II with small apical claw-like projection and with ventral membranous spine. Tibiae I, II with a pair of ventral spines (Figure 5B, C). Tibia III with long paraxial spur 24 (24–26).
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Temnalges fulicae* sp. n.: ventral view of male.</figcaption></figure>

long; anti-axial spur absent. Tarsus IV about 1/3 the length of tibia IV, with a pair of spine-like dorsal projections; setae *d*, *e* of tarsus IV barrel-shaped with cap, situated apically; ambulacral disc much smaller than on tarsi I–III (Figure 5D, E). Lengths of solenidia: ωII 24–26, ω3I 28–32, ωIIΙ 22–25, σ2I 67–72, σII 5–6, σIII 50–55, φI 56–60, φII 68–70, φIII 55–60, φIV 30–34.

**Female** — (range for 10 paratypes) (Figures 3, 4, 5F, G). Idiosoma, length × width, 230–245 × 135–145, length of hysterosoma 140–150. Prodorsal shield: shaped as longitudinal trapezoid, posterior corners rounded, posterior margin convex with small median extension,
<figure><img src="image_5.png" /><figcaption>**Figure 3** *Temnalges fulicae* sp. n.: dorsal view of female.</figcaption></figure>

length 87–92, greatest width 63–68. Setae *vi*, *ve* absent. Setae *si*, *se* off prodorsal shield, bases of *se* separated by 62–65. Subhumeral setae *c3* short, approximately as long as trochanters III. Distance between prodorsal and hysteronotal shield along midline 20–25. Hysteronotal shield: rectangular in shape with slightly concave margins, greatest length 95–100, width at anterior margin 56–60, width at posterior margins 54–58 (Figure 3). Setae *d2* and *e2* on lateral margins of this shield. Hysteronotal gland openings on striated tegument near lateral margins of hysteronotal shield, slightly closer to level of setae *d2* than to setae *e2*. Setae *e1* closer to level of openings *gl* than to level of setae *e2*. Distances between dorsal setae: *c2*:d2 32–35, *d2*:e2 68–75, *e2*:h2 32–40, *d2*:d2 56–60, *e2*:e2 52–56, *h2*:h2 42–48. Lengths of dorsal setae: *c2* 9–11, *d2* 7–10, *e2* 10–12.

Epimerites I free, bow-shaped, with slightly divergent posterior ends (Figure 4). Epigynum semi-ovate, with parallel lateral branches, 45–48 long, 42–45 wide, anterior margin extending beyond tips of epimerites II, posterior ends extending to level of epimerites III tips; lateral margins with small extensions directed towards tips of epimerites III. Genital papillae near
<figure><img src="image_6.png" /><figcaption>**Figure 4** *Temnalges fulicae* sp. n.: B—ventral view of female.</figcaption></figure>

inner margins of epigynum. Setae *g* situated on posterior tips of epigynum, almost extending to level of setae *ps3*. Setae *3a* short, filiform. Setae *4a* extending to or slightly beyond setae *ps3*. Copulatory opening situated terminally on short frustum-like extension. Distance between ventral setae: *4b:3a* 27–33, *4b:g* 33–37, *g:4a* 15–18. Lengths of ventral setae: *g* 55–65, *3a* 13–22, *4a* 65–75, *4b* 10–12. Setae *ps1, ps2, f2, h3* short filiform, 8–15 long.

Legs I and II as in the male, except paraxial ventral extensions of tibiae I, II smaller than in the male. Setae *r, s and w* of tarsi III and setae *w* and *r* of tarsi IV filiform. Setae *d* of tarsi III, IV filiform, much longer than other tarsal setae (Figure 5F, G). Lengths of tarsi III and IV 29–31 and 32–34, respectively. Tarsal seta *dIII* slightly longer than combined length of tibia-tarsus III, seta *dIV* approximately equal to combined length of tibia-tarsus IV. Lengths of solenidia: ωI 28–30, ω3I 26–28, ωII 23–25, σ2I 44–46, σII 6–7, σIII 26–26, φI 58–60, φII 68–72, φIII 48–53, φIV 38–43.
<figure><img src="image_7.png" /><figcaption>**Figure 5** *Temnalges fulicae* sp. n., details: A — opisthosoma of male, ventral view; B, C — legs I, II of male, respectively; D — genu, tibia and tarsus III of male; E — leg IV of male; F, G — tibia and tarsus III, IV of female, respectively.</figcaption></figure>
## Differential diagnosis

Males of the new species, *Temnalges fulicae* sp. n., are closest in appearance to *T. megalonyx* Gaud, 1968 in having a long paraxial spur of tibia III, which is as long as 2/3 the length of tarsus III, the prodorsal and hysteronotal shields are separated by a wide area of soft tegument. *Temnalges fulicae* differs from *T. megalonyx* in the following features: in males of *T. fulicae*, the posterior end of opisthosoma is rounded, without a distinct terminal cleft and the lateral clefts between the bases of setae *f2* and *ps2*; the inner tips of epimerites III are simple, without a pair of acute extensions; the epiandrum is low, with the tips not extending to the base of the genital arch; the adanal shields are short, with posterior ends not extending to the level of the adanal suckers; in females, the hysteronotal shield is roughly rectangular, with slightly concave margins; the humeral setae *cp* are situated on the inner margins of the humeral shields, and the lateral branches of the epigynum are parallel-sided. In males of *T. megalonyx*, the opisthosoma has a small terminal cleft between setae *ps1* and a pair of lateral concavities between the bases of setae *f2* and *ps2*; the inner tips of epimerites III have a pair of acute unequal extensions; the tips of the epiandrum extend to the base of the genital arch, the adanal shields are long, bow-shaped and extend beyond the level of the adanal suckers; in females, the hysteronotal shield is trapezoidal with a noticeably narrowed anterior part; the humeral setae *cp* are distant from the inner margins of the humeral shields, and the lateral branches of the epigynum are divergent.

## Etymology

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

## Key to *Temnalges* species

(Males and females)

<table><tr><td>1. Vertical setae vi absent</td><td>2</td></tr><tr><td>— Vertical setae vi present</td><td>6</td></tr></table>

<table><tr><td>2. In both sexes, femur II with large and rounded, hook-like lateral process. In males, pretarsus of leg IV absent. In females, tarsus IV strongly reduced to small cylindrical segment without pretarsus and with seta d modified in long whip-like macroseta</td><td>T. atelodiscus Hernandes & Mironov, 2015</td></tr><tr><td>— In both sexes, femur II without lateral process. In males, pretarsus of leg IV present although smaller than on tarsi I-III. In females, tarsus IV with normally developed pretarsus, as in tarsus III</td><td>3</td></tr></table>

<table><tr><td>3. In males, epiandrum absent, paraxial spur-like process of tibia III very short, about 1/8 the length of corresponding tarsus. In females, tips of epigynum not extending to bases of setae g and level of epimerites IIIa, setae g much shorter than the distance between their bases and posterior end of opisthosoma</td><td>T. hoazin Hernandes & Mironov, 2015</td></tr><tr><td>— In males, epiandrum present, paraxial spur-like process of tibia III at least as long as 1/3 the length of corresponding tarsus. In females, tips of epigynum extending to bases of setae g and level of epimerites IIIa, setae g equal to or longer than half the distance between their bases and posterior end of opisthosoma</td><td>4</td></tr></table>

<table><tr><td>4. In males, paraxial spur-like process of tibia III as long as 1/3 the length of corresponding tarsus. In females, prodorsal and hysteronotal shields almost touching, setae se, si situated on prodorsal shield</td><td>T. mesalgoides Gaud & Atyeo, 1967</td></tr><tr><td>— In males, paraxial spur-like process of tibia III as long as 2/3 the length of corresponding tarsus. In females, prodorsal and hysteronotal shields separated by the distance 20–40, setae se,</td><td></td></tr></table>
<table><tr><td>si situated off prodorsal shield</td><td>5</td></tr></table>

5. In males, opisthosoma with small terminal cleft between bases of setae ps1 and with small lateral concavities between bases of setae f2 and ps2, adanal apodemes represented by narrow bow-shaped sclerites with posterior ends extending to level of adanal suckers. In females, prodorsal shield with straight posterior margin, hysteronotal shield trapezoidal, with anterior end distinctly attenuate. *T. megalonyx* Gaud, 1968
— In males, opisthosoma rounded, without terminal cleft and lateral concavities between bases of setae f2 and ps2, adanal shields represented by narrow oblique elongated sclerites with posterior ends not extending to level of adanal suckers. In females, prodorsal shield with convex posterior margin, hysteronotal shield roughly rectangular, with slightly concave lateral margins. *T. fulicae* sp. n.

6. In males, posterior part of hysteronotal shield represented by narrow sclerotized bands along lateral margins of opisthosoma separated by large semi-ovate incision extending to midlevel of this shield; paraxial spur-like process on tibia III absent, epiandrum and paragenital apodemes absent. In females, hysteronotal shield trapezoidal, epigynum with convex lateral margins and tips slightly curved to midline, setae g and 4a much shorter than epigynum. *T. bothrioplax* (Gaud, 1958)
— In males, posterior part of hysteronotal shield covering entire surface of opisthosoma, tibia III with paraxial spur-like process, only epiandrum or paragenital apodemes fused with epiandrum present. In females, hysteronotal shield roughly rectangular in shape, epigynum with parallel or slightly divergent branches, setae g and 4a equal to or longer than epigynum. 7

7. In both sexes, setae *se* and *si* on prodorsal shield, setae *si* situated slightly posterior to level of setae *se*. In males*, epiandrum and paragenital apodemes fused in sclerite shaped as an inverted U surrounding genital apparatus, setae *1a* over 100 long, spur-like process of tibia III as long as 1/3 of corresponding tarsus. In females, hysteronotal shield with lateral margins strongly concave, 1.2–1.3 times longer than wide at anterior margin. *T. amalicus* Shereef & Rakha, 1981
— In both sexes, setae *se* and *si* off prodorsal shield, setae *si* slightly anterior to level of setae *se*. In males*, small bow-shaped epiandrum flanks anterior end of genital apparatus, setae *1a* about 30 long, spur-like process of tibia III half as long of corresponding tarsus. In females, hysteronotal shield with lateral margins almost straight, about 2–2.3 times longer than wide at anterior margin. *T. mahranus* Shereef & Rakha, 1982

*Remark. It is clear from the text of the corresponding original descriptions that the figures illustrating the males of *T. amalicus* and *T. mahranus* (Shereef and Rakha 1982: figs. 3, 5, respectively) are actually mixed up, while the drawings of females are correct. In diagnostic characters for males in the key, we followed the characteristics given in the text for original descriptions of these species.

## Acknowledgements

The taxonomic part of the study was supported by the Ministry of Science and Higher Education of the Russian Federation (project No. 122031100263-1) to SVM. 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 for the care with which they treated birds that were a component in this study. Scientific Permits to allow possession and examination of birds were issued to TDG by the Canadian Wildlife Service, Environment and Climate Change Canada / Government of Canada. The authors wish to thank Jakob Skov and Lars Holm Hansen for providing specimens of *Fulica atra* from Denmark.
ORCID

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

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

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

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