Research Article |
Corresponding author: Katarína Fogašová ( katarina.fogasova@smail.unipo.sk ) Academic editor: Gunnar Kvifte
© 2023 Libor Dvořák, Katarína Fogašová, Jozef Oboňa, Edina Török, Peter Manko.
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:
Dvořák L, Fogašová K, Oboňa J, Török E, Manko P (2023) Two new Ptychoptera Meigen, 1803 (Diptera, Ptychopteridae) from the Western Palaearctic. ZooKeys 1166: 91-102. https://doi.org/10.3897/zookeys.1166.96193
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Ptychoptera xanthopleura Dvořák, Oboňa & Manko, sp. nov. from Azerbaijan and Georgia, and Ptychoptera staryi Dvořák, Oboňa & Manko, sp. nov. from Bulgaria are described. P. xanthopleura sp. nov. differs from the other member of the lacustris group mainly by having almost completely yellow pleurae, and by the shape of the epandrium and gonocoxites. The diagnostics of P. staryi sp. nov. and P. incognita Török, Kolcsár & Keresztes, 2015 based on male genitalia are provided.
Balkan Peninsula, Caucasus, distributional data, new species, phantom crane flies, Ptychopteridae
Ptychopteridae comprises two extant subfamilies: Bittacomorphinae Shiner and Ptychopterinae Alexander. Bittacomorphinae is found in the Nearctic, East Palearctic, and Oriental regions (e.g.,
The genus Ptychoptera Meigen, 1803 comprises more than 90 recent species worldwide (e.g.,
During the last ca. 40 years, eight new Ptychoptera were described from the Western Palaearctic region; one from Italy (P. delmastroi Zwick & Starý, 2003), four from the Balkans and neighbouring areas (P. agnes Krzemiński & Zwick, 1993 from Hungary, P. incognita Török, Kolcsár & Keresztes, 2015 from Romania and Bulgaria, and P. castor Keresztes & Kappert, 2021 and P. pollux Keresztes & Török, 2021 both from the south Balkan area), and three from the Caucasus (P. peusi Joost, 1974 from Russian Caucasus, P. ressli Theischinger, 1978 from Iran, and P. alina Krzemiński & Zwick, 1993 from Armenia).
The adults live in marshy and moist habitats, near suitable substrates for the larvae as well as quite some distance from the larval habitat (
Our present study reports the description of two new Ptychoptera species, one from the Balkans and one from the Caucasus.
The terminology of male genitalia follows
All voucher specimens are deposited and accessible in the private collection of the 1st author. Genitalia of Ptychoptera staryi sp. nov. and P. xanthopleura sp. nov. were macerated in 10% KOH and dehydrated using a series of dehydrating alcoholic solutions (70%, 80%, 96%). After that, parts of genitalia were mounted on permanent slides using Canada balsam as mounting medium.
Photographs of specimens were taken using a Pentax K-50 camera and a reverse-mounted Vivitar 28 mm 1: 2.0 MC lens, Motic SMZ-1 68 stereomicroscope equipped with Canon EOS 1200D camera and EOS utility software, and with Leica M205C stereomicroscope equipped with Leica DFC295 digital camera. Focus stacking was performed using Adobe Photoshop.
We undertook phylogenetic analyses to understand the relationships between the species in the subgenus Paraptychoptera. Cladistic analyses of 53 morphological characters on antennae, wing, and male terminalia (Table
Matrix of the 53 morphological data (based on the
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | ||
P. contaminata | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | |
P. agnes | ? | 1 | 1 | 1 | ? | ? | ? | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | |
P. castor | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | |
P. delmastroi | ? | ? | ? | ?1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. handlirschi | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. helena | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | |
P. lacustris | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. longicauda | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. paludosa | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. pollux | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | |
P. resseli | ? | ? | ? | ? | 0 | ? | ? | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | |
P. silvicola | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
P. xanthopleura sp. nov. | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | |
27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 | 42 | 43 | 44 | 45 | 46 | 47 | 48 | 49 | 50 | 51 | 52 | 53 | |
P. contaminata | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
P. agnes | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 1 |
P. castor | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 |
P. delmastroi | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 |
P. handlirschi | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 |
P. helena | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 |
P. lacustris | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 1 |
P. longicauda | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 |
P. paludosa | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 |
P. pollux | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 |
P. resseli | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 |
P. silvicola | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 |
P. xanthopleura sp. nov | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 |
The morphological data matrix was created and managed with Mesquite 3.5 (
For visualisation of the phylogenetic tree, we used FigTree v. 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/), and the character state and statistical values visualisation were plotted onto the trees using Adobe Photoshop.
Medium sized to large (7–15 mm), slender, black lustrous Nematocera, often with lighter markings on thorax and/or abdomen. Antennae, wings, abdomen, and legs long and slender. Ocelli absent; antenna with 15 to 16 segments. Thorax with deep, posteriorly directed transverse suture. Wing with markings, in particular along the crossveins and where veins bifurcate; spurious vein present on either side of crossvein R-M and wing membrane with a distinct fold between veins A1 and CuA2 (
Holotype : 1 ♂: Azerbaijan, Qum, sidebrook/small tributary of the Ardavacaj (Ardavachay) River + wetland, 845 m a. s. l., 41°28'10.3"N, 46°55'57.2"E, 8.V.2019, leg. J. Oboňa & P. Manko. Paratypes: 2 ♂♂: Georgia, border of Imereti and Samtskhe–Javakheti regions, brook and spring, south slope of Zekari pass, 2 050 m a. s. l., 41°49'23"N, 42°51'09"E, 17.VII.2019, leg. G. Vinçon.
Male. Head: Frons, vertex, and occiput black with metallic blue shine, mouthparts including palpi pale yellow, scape and pedicel yellowish orange, antennal flagellomeres a somewhat darker, tending to pale brown.
Thorax : Scutum, paratergite, and mediotergite blackish with metallic blue shine; scutellum, pleurotergite, katepisternum, and katepimeron brownish black with lighter metallic blue shine; other parts yellow. Halteres yellow with light brown knob. Legs yellow except brown extreme apex of femora and tibiae, tarsi somewhat darkened.
Wing length 10 mm (holotype, Fig.
Abdomen : Tergum 1 dark shiny brown with yellow apex, sternum 1 yellow. Tergum 2 brown basally and apically, yellow in middle, sternum 2 yellow. Tergum 3 yellow basally, brown apically, sternum 3 yellow. Remaining terga and sterna brown, sternum 4 yellow basally. Auxiliar copulatory organ yellow.
Male genitalia
(Fig.
Ptychoptera xanthopleura sp. nov. terminalia a epandrium b paramere c aedeagal complex d gonostylus e hypandrium. Abbreviations: ECP = epandrial clasper, EL = epandrial lobe, EPI = epiproct, GBM = medial lobe of basal lobe of gonostylus, GAS = apical stylus of gonostylus, ALP = lateral ejaculatory process.
Female. The authors have an immature female which was sampled in Lesser Caucasus (Georgia, Kakheti region, Ilto river, above (N of) the Chart’ala village, 790 m a. s. l., 42°8'18"N, 45°7'32"E, 8.VII.2019, leg. P. Manko & G. Vinçon). The characters correspond to the above-described new species. However, its identity cannot be confirmed in this stage of ontogenesis/development and could be solved after collecting more specimens of the genus Ptychoptera from the Transcaucasia.
The name reflects predominantly yellow pleurae (Fig.
According to the presence of auxiliary sexual organ and shining pleurae, P. xanthopleura sp. nov. belongs to the subgenus Paraptychoptera and according to male genitalia and the maximum parsimonious tree based on 53 morphological characters (see Fig.
Holotype : 1 ♂: Bulgaria, Rhodopes, Yundola, 1 300 m a. s. l., 42°3'47"N, 23°51'17"E, 30.VI.2016, leg. M. Barták et Š. Kubík.
Male. Head: Frons, vertex, and occiput black, mouthparts including palpi pale yellow, scape and pedicel yellowish orange, antennal flagellomeres greyish.
Thorax : Predominantly black with silvery pubescent pleurae. Pronotum, epimeron 3 and metanotum 3 yellow. Fore and mid coxae and trochanters yellow, hind coxae brownish black basally, yellow apically, coxae also yellow. Almost all legs are missing. Halteres whitish yellow with a darker knob.
Wing length 12 mm (Fig.
Abdomen : Tergum 1, tergum 2, apical ¹⁄5 of tergum 3, almost whole tergum 4 except base, and whole terga 5–7 black; sternum 1 black, sternum 2 brown, sterna 5–7 black; the remainder orange yellow.
Male genitalia : similar to P. incognita. Epandrial clasper slightly curved outwards with simple obtuse apex; anterior projection of ventromesal lobe sharp, posterior projection bow-shaped backwards; space between both projections is rounded, almost semi-circular. Gonocoxite and gonostylus: apical process of paramere with a U-shaped dark structure with thick edges; paramere base rounded, convex; width to height ratio of dorsal gonocoxal lobe 0.5; dorsal gonocoxal lobe with dense tiny dark hairs. Aedeagus: sides of lateral ejaculatory process distinctly convex, basal projections markedly convergent; transition to lateral ejaculatory process smooth, undulate. Hypandrium: width to length ratio 1.2; apex of hypogynial valves start under basal division of hypandrium. See also differential diagnosis.
Female. Unknown.
The name is dedicated to our colleague Jaroslav Starý and his life jubilee. (Jaroslav discovered the holotype of new species in his own material and provided it for the description).
The new species is very similar to P. incognita Török, Kolcsár & Keresztes, 2015 (see also Table
Epandrium, gonocoxite and gonostylus, and hypandrium of P. staryi sp. nov., P. incognita, and P. albimana. Abbreviations: ECP = epandrial clasper, EL = epandrial lobe, EPI = epiproct, GBM = medial lobe of basal lobe of gonostylus, GCT = gonocoxite, GAS = apical stylus of gonostylus, GAT = tertiary lobe of apical stylus of gonostylus, PPA = apical process of paramere, HBD = basal division of hypandrium, HMW = membranous window of terminal division of hypandrium, HTD = terminal division of hypandrium.
Diagnostics of P. staryi sp. nov., P. incognita, and P. albimana based on male genitalia (see also Fig.
P. staryi sp. nov. | P. incognita | P. albimana | |
---|---|---|---|
Epandrium | Shape of the plate on ventral site and the space between projections rounded, almost semi-circular. | Projections of the plate on ventral site in about right angle, space between projections quadrangular. | Projections of the plate on ventral site in obtuse angle, space between projections rounded, more than semicircle. |
Gonocoxite and gonostylus | Width to height ratio of dorsal gonocoxal lobe 0.5; dorsal gonocoxal lobe with dense tiny dark hairs. The hairs of margins of gonocoxite and gonostylus much less dense and finer. Chitinisation of the proximo-lateral processes of the gonocoxite not developed, processes light coloured. | Width to height ratio of dorsal gonocoxal lobe 0.6; dorsal gonocoxal lobe with sparse small hairs. The hairs of margins of gonocoxite and gonostylus dense and stronger. Chitinisation of the proximo-lateral processes of the gonocoxite strongly chitinised, dark. | Width to height ratio of dorsal gonocoxal lobe 0.4; dorsal gonocoxal lobe with tiny dark hairs at the top. The hairs of margins of gonocoxite and gonostylus dense and stronger. Chitinisation of the proximo-lateral processes of the gonocoxite not developed, processes pale coloured. |
Aedeagus | Sides of lateral ejaculatory process distinctly convex, basal projections markedly convergent; transition to lateral ejaculatory process smooth, undulate. | Sides of lateral ejaculatory process slightly convex, basal projections divergent; transition to lateral ejaculatory process steep, in a right or acute angle to the axis of aedeagus. | Sides of lateral ejaculatory process slightly convex, basal projections almost parallel; transition to lateral ejaculatory process steep, in an obtuse angle to the axis of aedeagus. |
Hypandrium | Width and length ratio 1.2; apex of hypogynial valves start under basal division of hypandrium. | Width and length ratio 0.9; apex of hypogynial valves start on bases of basal division of hypandrium. | Width and length ratio 0.9; apex of hypogynial valves start over basal division of hypandrium. |
We are grateful to Gilles Vinçon (independent researcher, Grenoble, France) for his help in collecting and for the material he provided us from his own collections and Jaroslav Starý (Olomouc, Czech Republic) for providing us with the specimen for description, which was collected by Miroslav Barták et Štěpán Kubík (both - Czech University of Life Science, Prague, Czech Republic).
No conflict of interest was declared.
No ethical statement was reported.
This work of was financially supported by the Grant Agency of University Prešov in Prešov under the contract No. GaPU 1/2022 and by the Slovak Scientific Grant Agency, contract No. VEGA-1/0012/20.
LD is the leading person in preparing the description of both new species. PM took photographs, edited and prepared figures, PM, ET prepared the morphological data matrix for phylogenetic analyses, participated in the description, PM participated in the differential diagnosis, organized and provided funding for sampling trips to the Caucasus, LD, ET, PM, JO, KF contributed substantially to the writing of the manuscript and its revisions.
Libor Dvořák https://orcid.org/0000-0002-4712-3679
Katarína Fogašová https://orcid.org/0009-0003-9604-1885
Jozef Oboňa https://orcid.org/0000-0002-1185-658X
Edina Török https://orcid.org/0000-0001-5982-7078
Peter Manko https://orcid.org/0000-0003-1862-9117
All of the data that support the findings of this study are available in the main text.