Research Article |
Corresponding author: Pavel Sroka ( pavel.sroka@centrum.cz ) Academic editor: Ben Price
© 2021 Pavel Sroka, Zohar Yanai, Dmitry Palatov, Jean-Luc Gattolliat.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sroka P, Yanai Z, Palatov D, Gattolliat J-L (2021) Contribution to the knowledge of the genus Takobia Novikova & Kluge, 1987 (Ephemeroptera, Baetidae) in Central Asia. ZooKeys 1071: 127-154. https://doi.org/10.3897/zookeys.1071.71582
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Based on the original type material, the nymphal stage of the mayfly Takobia maxillare is redescribed; in parallel, a lectotype is designated. Takobia maxillare is the type species of the genus Takobia, and an accurate and complete knowledge of its morphology is crucial to the delimitation of this problematic genus and clarification of its phylogenetic affinities. Ambiguous characters, previously reported for this species in the literature are clarified. Furthermore, two new species in the same genus are described, namely Takobia sinusopalpata sp. nov. and Takobia shughnonica sp. nov. based on the morphology of nymphs from Central Asia, supplemented with COI sequences. Implications for the systematics of Takobia and related taxa are discussed and the need for an extensive phylogenetic study of this group is stressed.
Alainites, mayflies, new species, Nigrobaetis, redescription, systematics, taxonomy
Baetidae encompass more than 1150 species nested in 115 genera, making it the most speciose family of all mayflies. Its systematics is accordingly complicated, and subject to frequent changes mainly related to generic concepts and species delimitation. Some taxa within Baetidae have particularly complex histories, such as Nigrobaetis Novikova & Kluge, 1987, Alainites Waltz & McCafferty, 1994, and Takobia Novikova & Kluge, 1987, which have all been subject to several synonymies and changes in rank between species groups, subgenera, and genera (Müller-Liebenau 1969;
In this study, we focus on Takobia maxillare (Braasch & Soldán, 1983), the type species of the problematic taxon Takobia. For any future extensive analysis aimed at assessing the relevance and extent of Takobia, detailed knowledge of its type species is of crucial importance. The species was originally described by
Subsequently,
Since T. maxillare represents the type species of Takobia, an accurate knowledge of its morphology is a key prerequisite for the precise delimitation of the genus and is of significant importance for the generic attribution of all species of Alainites (also including species previously described in Acerbaetis Kang & Yang, 1994, in
Therefore, the present study aims to provide an updated redescription of T. maxillare and analyze discrepancies between the descriptions of
The original type material of T. maxillare was obtained from the collection of the Biology Centre of the Czech Academy of Sciences, Institute of Entomology, České Budějovice, Czech Republic (
Some specimens were mounted on slides with HydroMatrix (MicroTech Lab, Graz, Austria). Drawings were made using a stereomicroscope Leica M205 C and a microscope Olympus BX41, both equipped with a drawing attachment. Photographs were made using a Canon EOS 6D camera and the Visionary Digital Passport imaging system and processed with Adobe Photoshop Lightroom (http://www.adobe.com) and Helicon Focus version 5.3 (http://www.heliconsoft.com). Photographs were subsequently enhanced with Adobe Photoshop CS6. For scanning electron microscopy, samples were gradually transferred to acetone, critical point dried, and coated with gold by sputtering using a Baltec SCD050 Sputter Coater. Observations were made on the Jeol JSM 7401F at 4 kV scanning microscope at the Biology Centre CAS, České Budějovice, Czech Republic.
DNA was extracted from two individuals per species of Takobia maxillare (failed) and of the two newly described species. In addition, we extracted DNA for the first time from two additional species from central Asia: Alainites talasi (Novikova & Kluge, 1994) and A. kars (Thomas & Kazancı, in
Taxa used for genetic distance analysis (mitochondrial COI sequences) with GenBank accession numbers (novel sequences are highlighted in bold font).
Species | GenBank accession numbers |
---|---|
Takobia shughnonica sp. nov. (paratype) | MZ983793 , MZ983794 |
Takobia sinusopalpata sp. nov. (paratype) | MZ983795 , MZ983796 |
Alainites albinatii | HG934994, HG934995 |
Alainites kars | MZ983797 , MZ983798 |
Alainites muticus | HG934999, JN299112 |
Alainites talasi | MZ983799 , MZ983800 |
Alainites yixianii | GU479735 |
Nigrobaetis bacillus | MH823363, MH823364 |
Nigrobaetis digitatus | JN164308, JN164309, LT626141 |
Nigrobaetis gracilis | JN164320 |
Nigrobaetis minutus | HM417038 |
Nigrobaetis niger | JN164310, JN164311, KC158570, KC158571 |
Nigrobaetis paramakalyani | LC056973 |
Nigrobaetis vuatazi | HE651544 |
Related taxa were added to the analysis, based on published sequences in GenBank database (https://www.ncbi.nlm.nih.gov/; see Table
Takobia maxillare can be easily separated from other related species by the combination of the following characters: 1) maxillary palp highly developed with the segment I widened apically and segment II straight; 2) labrum dorsally covered with numerous setae, none of them arranged in a row; 3) right prostheca reduced, apically bifid; 4) labial palp segment III quadrangular, asymmetrical, with a short projection lateroapically; 5) claw edentate, subequal to 1/2 of corresponding tarsus; 6) paraproct with a short prolongation bent ventrally.
Length. Female body 6.8–9.1 mm; cerci 4.6–5.4 mm; median caudal filament 3.4–4.5 mm; male body 5.0–7.2 mm; cerci 4.2–5.8 mm; median caudal filament 2.5–3.5 mm.
Coloration and texture. General coloration brown (Fig.
Takobia nymphs, habitus photographs A T. maxillare, habitus in dorsal view B T. sinusopalpata sp. nov., habitus in dorsal view C T. shughnonica sp. nov., habitus in dorsal view D T. maxillare, habitus in lateral view E T. sinusopalpata sp. nov., habitus in lateral view F T. shughnonica sp. nov., habitus in lateral view. Scale bar:1 mm.
Head. Antennae close to each other, with a narrow interantennal carina; scape and pedicel with V-shaped scale insertions and sparse setae. Dorsal surface of labrum (Fig.
T. maxillare, nymph A labrum (left side dorsal view, right side ventral view) B left mandible (dorsal view) C right mandible (dorsal view) D right mandible, detail of incisors and prostheca (dorsal view) E left mandible (dorsal view), detail of incisors and prostheca (dorsal view) F maxilla (dorsal view) G labium (left side ventral view, right side dorsal view) H foreleg (dorsal view) I middle leg (dorsal view) J hind leg (dorsal view) K–Q gills.
Thorax. Forelegs (Fig.
Abdomen. Tergites (Fig.
Holotype . mature male nymph (in EtOH): Kyrgyzstan, Chuy Region. Spring – left tributary of the Adygene Riv., 144 m a.s.l., 42°34.19'N, 74°28.57'E, 29.4.2016, Palatov leg., locality code: 17Kyrg. Paratypes. 39 nymphs: same data as holotype (33 in EtOH, 2 on slides with HydroMatrix mounting medium, 2 on SEM stubs, 2 DNA voucher specimens). 1 nymph (in EtOH): Kyrgyzstan, Chuy Region, Korumdy Riv., 300 m upstream its mouth to Suusamyr Riv., 2214 m a.s.l., 42°12.40'N, 73°41.48'E, 1.5.2016, Palatov leg., locality code: 19Kyrg. 3 nymphs (in EtOH): Kyrgyzstan, Talas Region, Oshibulag Riv. – right tributary of Chychkan Riv., 1629 m a.s.l., 42°05.77'N, 72°48.19'E, 2.5.2016, Palatov leg., locality code: 25Kyrg. 1 nymph (in EtOH): Kyrgyzstan, Talas Region, Chon-chychkan Riv., ca. 1.5 km upstream Talas-Bishkek highway bridge, 1924 m a.s.l., 42°25.76'N, 72°44.03'E, 11.5.2016, Palatov leg., locality code: 60Kyrg. 12 nymphs (11 in EtOH, 1 on slide with HydroMatrix mounting medium): Kyrgyzstan, Talas Region. Otmek Riv. 2801 m a.s.l., 42°19.08'N, 73°05.77'E, 12.6.2016, Palatov leg., locality code: 65Kyrg. 6 nymphs (in EtOH): Kyrgyzstan, Osh Region, Kulun Riv., upstream from confluence of Kulaimende and Dungar Riv., 2229 m a.s.l., 40°30.46'N, 74°14.37'E, 1.5.2017, Palatov leg., locality code: 74 Kyrg.
Holotype and 30 paratypes are deposited in
Takobia sinusopalpata sp. nov. can be separated from other related species by the combination of the following characters: 1) maxillary palp highly developed with the segment I straight and segment II sinusoidal; 2) labrum dorsally covered with numerous setae, one central and two lateral forming the traditional disto-lateral arc of setae; 3) right prostheca reduced, basally bifid; 4) labial palp segment III quadrangular, slightly asymmetrical; 5) claw with one row of small teeth, subequal to 1/3 of corresponding tarsus; 6) paraproct with a short bent prolongation.
Length. Female body 6.8–7.4 mm; cerci and median caudal filament partially broken off, cerci assumed ca. 5 mm, medial caudal filament ca. 3.5 mm; male body 6.0–6.7 mm; cerci 4–5.2 mm; median caudal filament 2.6–3.4 mm.
Coloration and texture. General coloration brown (Fig.
Head. Antennae close to each other, with a narrow interantennal carina; scape and pedicel with V-shaped scale insertions and sparse setae. Dorsal surface of labrum (Fig.
Takobia sinusopalpata sp. nov., nymph A labrum (left side dorsal view, right side ventral view) B left mandible (dorsal view) C right mandible (dorsal view) D right mandible, detail of incisors and prostheca (dorsal view) E left mandible (dorsal view), detail of incisors and prostheca (dorsal view) F maxilla (dorsal view) G labium (left side ventral view, right side dorsal view) H foreleg (dorsal view) I middle leg (dorsal view) J hind leg (dorsal view) K–Q gill plates.
Thorax. Forelegs (Fig.
Abdomen. Tergites (Fig.
The name of the new species, sinusopalpata, refers to the sinusoidal shape of the second segment of the maxillary palps, very pronounced and characteristic for this species.
So far known from several localities in the Tien Shan Mountains (Kyrgyzstan). Nymphs were collected from stones and boulders sometimes covered with algae and moss in mountain springs, streams, and small rivers located at altitudes of 1600–2800 m a.s.l., at flow rates of 0.5–1.0 m/s, with water temperatures ca. 10–12°C (Fig.
Holotype . mature female nymph (in EtOH): Tajikistan, Roshtqal’a District. Spring near Sezhd village, 2966 m a.s.l., 37°12.65'N, 72°04.44'E, 2.7.2016, Palatov leg., locality code: 243Tj. Paratypes. 39 nymphs (33 in EtOH, 2 on slides with HydroMatrix mounting medium, 2 on SEM stubs, 2 DNA voucher specimens): same data as holotype. 13 nymphs (in EtOH): Tajikistan, Shughnon District, unnamed river, right tributary of Gunt Riv., ca. 500 m S from Dehmiyona village, 2700 m a.s.l., 37°42.88'N, 71°53.61'E, 23.5.2012, Palatov leg., locality code: 15Tj. 28 nymphs (in EtOH): Tajikistan, Shughnon District. Vuzh-dara Riv., 3 km upstream Dehmiyona village, 2500 m a.s.l., 37°42.47'N, 71°57.29'E, 24.5.2012, Palatov leg., locality code: 31Tj. 2 nymphs (in EtOH): Tajikistan, Shughnon District, unnamed river, tributary of Gunt Riv. near Shitam village, 2500 m a.s.l., 37°44.30'N, 72°2.19'E, 31.5.2012, Palatov leg., locality code: 76Tj. 54 nymphs (in EtOH): Tajikistan, Shughnon District, stream on the slope of Gunt Riv. valley, ca. 3 km downstream from Ver village, 2875 m a.s.l., 37°43.27'N, 72°1.85'E, 5.6.2012, Palatov leg., locality code: 93Tj. 1 nymph (in EtOH): Tajikistan, Shughnon District, unnamed river near Tong village, 2480 m a.s.l., 37°35.78'N, 71°43.79'E, 8.6.2012, Palatov leg., locality code: 113Tj. 1 nymph (in EtOH): Tajikistan, Shughnon District, spring on slope of the Bogev-dara gorge. 2578 m a.s.l., 37°31.13'N, 71°41.98'E, 9.6.2012, Palatov leg., locality code: 120Tj. 17 nymphs (in EtOH): Tajikistan, Shughnon District, right source of the Bogev-dara Riv., 2928 m a.s.l., 37°29.89'N, 71°44.36'E, 10.6.2012, Palatov leg., locality code: 123Tj.
Holotype and 34 paratypes are deposited in
Takobia shughnonica sp. nov. can be separated from other related species by the combination of the following characters: 1) maxillary palp highly developed with the segment I straight and segment II slightly sinusoidal; 2) labrum dorsally covered with numerous setae, one central and two lateral forming the traditional disto-lateral arc of setae; 3) right prostheca reduced, basally bifid; 4) labial palp segment III symmetrical and almost conical; 5) claw with one row of teeth increasing in size toward the apex, subequal to 1/3 of corresponding tarsus; 6) paraproct with a short bent prolongation.
Length. Female body 6.4–7.4 mm; cerci 4.2–5.1 mm; median caudal filament 3.4–4.2 mm; male body 5.6–6.6 mm; cerci 3.5–3.6 mm; median caudal filament 2.7–3.0 mm.
Coloration and texture. General coloration brown (Fig.
Head. Antennae close to each other, with a narrow interantennal carina; scape and pedicel with V-shaped scale insertions and sparse setae. Dorsal surface of labrum (Fig.
Takobia shughnonica sp. nov., nymph A labrum (left side dorsal view, right side ventral view) B left mandible (dorsal view) C right mandible (dorsal view) D right mandible, detail of incisors and prostheca (dorsal view) E left mandible, detail of incisors and prostheca (dorsal view) F maxilla (dorsal view) G labium (left side ventral view, right side dorsal view) H foreleg (dorsal view) I middle leg (dorsal view) J hind leg (dorsal view) K–Q gill plates.
Thorax. Forelegs (Fig.
Abdomen. Tergites (Fig.
The species is named shughnonica after the local ethnicity and the historical region of Shughnon, where the species was discovered.
So far known from several localities in the Pamir Mountains (Tajikistan). Nymphs were collected from stones and boulders sometimes covered with algae and moss in mountain springs and streams located at altitudes of 2480–2928 m a.s.l., at flow rates of 0.5–1.0 m/s, with water temperatures ca. 10–12°C (Fig.
Takobia spp., examples of habitats A locality of T. sinusopalpata sp. nov. (Otmek Riv., locality code: 65Kyrg) B type locality of T. sinusopalpata sp. nov. (left tributary of the Adygene Riv., code: 17Kyrg) C locality of T. shughnonica sp. nov. (right tributary of Gunt Riv., code: 15Tj) D locality of T. shughnonica sp. nov. (stream on the slope of Gunt Riv. valley, code: 93Tj) E locality of T. shughnonica sp. nov. (right source of the Bogev-dara Riv., code: 123Tj).
The monophyly of the two new Takobia species, as well as that of A. talasi and A. kars, were confirmed (Table
Kimura 2 parameter distance among sequences of the mitochondrial COI gene of selected Takobia, Alainites, and Nigrobaetis species (presenting mean and min–max distance for each group with >1 individual; for number of samples per each taxon see Table
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | ||
1 | T. shughnonica | 0.0 | |||||||||||||
2 | T. sinusopalpata | 13.5 (13.5–13.5) | 0.0 | ||||||||||||
3 | A. albinatii | 22.5 (22.2–22.8) | 22.0 (21.9–22.1) | 1.5 | |||||||||||
4 | A. kars | 22.8 (22.7–22.9) | 23.2 (23.1–23.3) | 20.1 (19.9–20.3) | 0.1 | ||||||||||
5 | A. muticus | 25.0 (24.2–25.7) | 24.8 (24.7–24.9) | 17.0 (16.6–17.5) | 22.3 (22.1–22.5) | 17.4 | |||||||||
6 | A. talasi | 25.4 (25.4–25.4) | 23.1 (23.1–23.1) | 27.3 (27.1–27.6) | 24.9 (24.8–25.0) | 26.0 (24.3–27.8) | 0.0 | ||||||||
7 | A. yixianii | 25.9 (25.9–25.9) | 24.7 (24.7–24.7) | 20.8 (20.5–21.1) | 20.3 (20.2–20.4) | 23.7 (23.1–24.3) | 24.2 (24.2–24.2) | NA | |||||||
8 | N. bacillus | 26.6 (26.3–26.8) | 26.1 (25.9–26.3) | 22.3 (21.8–22.8) | 22.1 (21.7–22.5) | 25.4 (24.1–26.8) | 24.9 (24.6–25.3) | 0.8 (0.5–1.1) | 0.6 | ||||||
9 | N. digitatus | 22.5 (21.4–24.4) | 22.8 (22.1–24.2) | 23.3 (22.6–24.0) | 24.2 (23.9–24.6) | 25.3 (23.4–27.6) | 24.6 (24.3–25.1) | 23.5 (21.8–24.5) | 24.4 (21.9–26.1) | 9.9 (0.6–14.6) | |||||
10 | N. gracilis | 28.3 (28.3–28.3) | 27.7 (27.7–27.7) | 23.6 (23.6–23.6) | 25.1 (25.0–25.3) | 25.1 (24.7–25.4) | 24.8 (24.8–24.8) | 25.4 | 26.5 (26.0–27.0) | 24.5 (24.2–25.2) | NA | ||||
11 | N. minutus | 25.1 (25.1–25.1) | 22.5 (22.5–22.5) | 25.5 (25.0–26.1) | 24.5 (24.4–24.6) | 27.1 (26.7–27.5) | 25.4 (25.4–25.4) | 24.3 | 25.7 (25.4–26.1) | 23.9 (23.6–24.6) | 21.8 | NA | |||
12 | N. niger | 26.7 (25.8–27.6) | 25.5 (24.8–26.4) | 23.1 (22.0–24.2) | 22.0 (21.6–22.4) | 24.6 (23.4–25.8) | 24.9 (24.3–25.7) | 22.8 (22.0–23.0) | 24.6 (23.2–25.8) | 20.2 (19.4–21.3) | 23.2 (22.4–24.8) | 24.1 (22.9–25.3) | 1.0 (0.5–1.9) | ||
13 | N. paramakalyani | 25.5 (25.5–25.5) | 24.1 (24.1–24.1) | 26.5 (25.9–27.1) | 25.0 (24.9–25.1) | 27.3 (27.2–27.5) | 28.0 (28.0–28.0) | 26.8 | 28.0 (27.7–28.4) | 24.5 (22.8–25.3) | 18.7 | 20.2 | 24.7 (24.3–25.5) | NA | |
14 | N. vuatazi | 24.8 (24.8–24.8) | 23.6 (23.6–23.6) | 24.4 (24.1–24.6) | 22.7 (22.6–22.9) | 28.0 (27.7–28.4) | 24.0 (24.0–24.0) | 23.7 | 25.4 (25.1–25.0) | 21.9 (21.4–22.3) | 7.5 | 18.1 | 23.8 (22.9–24.4) | 17.8 | NA |
The original type series consisted of nymphal material collected on a single locality (“Uzbekische SSR, Kuk-kul-See, S von Fergana, 20.5.1980, leg. T. SOLDÁN et M. TONNER”, as given in
The collection of Dietrich Braasch is now housed mostly in the Stuttgart State Museum of Natural History, Germany, and partially in the Natural History Museum Potsdam and Senckenberg German Entomological Institute, Müncheberg, Germany. According to our inquiry to the curators of all these collections, there is no material of T. maxillare in any of them. Thus, either the types from D. Braasch should be considered lost or the type material was never split and all the types remained in T. Soldán’s collection.
The collection of Tomáš Soldán is now housed in the Biology Centre CAS, Institute of Entomology (
The exact location of the T. maxillare type locality is unclear. The original publication specifies a lake in Uzbekistan, south of the town Fergana. However, the border with Kyrgyzstan is ca. 20 km south of Fergana (Farg’ona), and there is no substantial water body south of Fergana within the main territory of Uzbekistan. Nevertheless, there is a small exclave of Uzbekistan within Kyrgyzstan further south. Two lakes are situated nearby, although not directly within the exclave, but a few hundred meters past the border in Kyrgyzstan. The borders have however shifted compared to where they were in the 1980s when the lakes were administratively located in Uzbekistan. Soldán and Tonner, the collectors of the original material, probably would not have realized that they had crossed the administrative border anyway, being technically still within the USSR in 1980.
The above-mentioned lakes are named Qurbonko’l and Ko’kko’l in Kyrgyzian sources, Курбан-Кёль (Kurban-Kiol’) and Кок- Кёль (Kok-Kiol’) in Russian, although exact transliteration varies. We are convinced that “Kuk-Kul” in the original publication is a version of Ko’kko’l and the present location in Kyrgyzstan instead of Uzbekistan is caused by the close proximity to the Uzbekistan exclave and the recent changes in the administrative borders in the area. Thus, we define the type locality of Takobia maxillare as follows: Kyrgyzstan, Ko’kko’l Lake, near border with Uzbekistan, Shakhimardan (Shohimardon) town, S of Fergana (Farg’ona), 39°56.10'N, 71°51.00'E.
Based on the identical locality, morphology, and deposition of the material and despite the lack of proper labeling, we consider the material located in T. Soldán’s former collection as the type material of T. maxillare; all the specimens constitute syntypes according to ICZN Article 73.2. To ensure the stability of the species, we thus designate a lectotype (female mature nymph) and paralectotypes (173 nymphs, same data as lectotype) according to the ICZN Article 74. The material was collected by T. Soldán and M. Tonner on 20.5.1980 in the type locality specified above. For the deposition of the material, see the chapter “Material examined”.
When comparing the original description of T. maxillare by
A very distinctive feature of T. maxillare is the elongated maxillary palps with the first segment widened apically and the second segment distinctly narrow in diameter (Fig.
In the type material of T. maxillare, the claws are apparently longer compared to the claws depicted in
The shape of the gills is also different between
Based on the characters compared above, we are rather confident that
These three species possess a distinctive paraproct with a short bent prolongation. The paraproct presents similar but more pronounced prolongation in various species historically assigned to Alainites (
The elongated maxillary palp is a character shared by T. maxillare, T. sinusopalpata sp. nov., and T. shughnonica sp. nov., and is much less developed in the various species assigned to Alainites. The shape of the palp significantly differs between these three species: in T. maxillare, the first segment is widened apically and curved outwards, while it is almost straight in T. sinusopalpata sp. nov. and T. shughnonica sp. nov.; the second segment is sinusoidal in T. sinusopalpata sp. nov., slightly sinusoidal in T. shughnonica sp. nov., and straight in T. maxillare.
It seems that the nymphal morphology of T. sinusopalpata sp. nov. and T. shughnonica sp. nov. is somewhat intermediary between Alainites sensu
The presence of claw denticles is considered as a plesiomorphic condition in Baetidae, being subject to reduction in several non-related lineages. On the other hand, the elongated maxillary palp is almost unique within Baetidae and probably represents a synapomorphy of T. maxillare, T. sinusopalpata sp. nov., and T. shughnonica sp. nov. Therefore, we assign both new species described herein to the genus Takobia, primarily based on this character. The three species are also closely distributed geographically, therefore they may form a single lineage restricted to Central Asia.
In the recent Baetidae phylogeny by
However, the inconsistencies in the description of T. maxillare and later redescription by
In conclusion, our results prove the existence of several Central Asian mayfly species closely related to T. maxillare. Two of them are newly described herein and another one erroneously assigned to T. maxillare in the literature. This was tested by the study of the original type material. The fact that Takobia hitherto consisted of a single species was only a consequence of our poor knowledge of the Central Asian mayfly fauna rather than T. maxillare really being something unique. The reclassification of all Alainites species based on such a premise is undesirable. Therefore, we refrain for the moment to follow the nomenclatural changes proposed by
We thank Roman J. Godunko for consultations regarding the position of the T. maxillare type locality. Furthermore, we are thankful to Marion Podolak (Museum of Zoology Lausanne) for her lab work and preparation of the COI barcodes. We also thank the colleagues from several museums who checked their collections for the original types of T. maxillare: A.H. Staniczek and M. Pallmann (Stuttgart State Museum of Natural History), D. Berger and C. Kuhlisch (Natural History Museum Potsdam), and S. Blank and A. Köhler (Senckenberg German Entomological Institute, Müncheberg). We are grateful to Chris Steer for the English language correction and to the reviewers (J. Webb, A.V. Martynov, and one anonymous referee) for their constructive comments, which helped to improve the manuscript. The study was funded by the institutional support of Institute of Entomology (Biology Centre of the Czech Academy of Sciences) RVO: 60077344 for PS.