Research Article |
Corresponding author: Lujza Keresztes ( keresztes2012@gmail.com ) Academic editor: Gunnar Kvifte
© 2021 Lujza Keresztes, Jürgen Kappert, Mária Henning, Edina Török.
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:
Keresztes L, Kappert J, Henning M, Török E (2021) Helen’s twins in the Balkans: discovery of two new Paraptychoptera Tonnoir, 1919 species closely related to P. helena Peus, 1958, with systematic revision of the “lacustris” group (Diptera, Ptychopteridae). ZooKeys 1071: 63-81. https://doi.org/10.3897/zookeys.1071.58598
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Ptychoptera castor Keresztes & Kappert, sp. nov. and P. pollux Keresztes & Török, sp. nov. both belong to the subgenus Ptychoptera (Paraptychoptera)
Cladistics analyses, identification key, male genital structures, new species, phantom craneflies, Ptychoptera, TNT phylogeny
Ptychopteridae or phantom crane flies are medium- to large-sized flies with slender shiny black body, sometimes with yellow or reddish markings, and long legs with tipuloid appearance, however they differ by several characters including their having a small membranous lobe at the base of the halter (
Extant representatives are classified in two subfamilies, Ptychopterinae with a single genus, Ptychoptera (about 70 species) and Bittacomorphinae with two other genera, Bittacomorpha (with only 2 species) and Bittacomorphella (8 species) (
Paraptychoptera is a monophyletic western Palaearctic group of Ptychopteridae, largely European in distribution, and with only a few species being present in Western Asia and North Africa. Only ten species are considered here, following
Among Paraptychoptera, the lacustris group was first proposed by
The type material of Paraptychoptera that was used in this study was acquired through field collections by the present authors. Seventy (70) male specimens belonging to nine different species originating from different parts of Europe were investigated (Fig.
Taxa | Nr. ind | Source of material | Coordinates | Collectors |
---|---|---|---|---|
P. (Pa.) agnes Krzemiński & Zwick, 1993 | 0 | literature data: Krzemiński & Zwick, 1993, Fasbender, 2014 | – | – |
P. (Pa.) castor sp. nov. | 1 | Albania, Vlora, Tragjas, Repet y Izvorit, 17 m, 30.iv.2019 | 40.323132°N, 19.510031°E | leg. Henning, M. |
P. (Pa.) delmastroi Zwick and Starý, 2003 | 0 | literature data: Zwick and Starý, 2003 , Fasbender, 2014 | – | – |
P. (Pa.) handlirschi (Czizek, 1919) | 3 | Romania, Baia Sprie, Gutin Mts., 955 m, 15.v.2014 | 47.698860°N, 23.794682°E | leg. Keresztes L. |
P. (Pa.) helena Peus, 1958 | 1 | paratype, NMBG Germnay | – | – |
P. (Pa.) lacustris Meigen, 1830 | 1 | Bulgaria, Berkovitsa, small brook, 616 m, 28.v.2013 | 43.218602°N, 23.071343°E | leg. Keresztes L. |
4 | Bulgaria, Dabravka, Dabravka lake shore, 364 m, 30.iv.2012 | 43.651059°N, 22.628539°E | leg. Török E. | |
1 | Bulgaria, Velingrad, Rhodope Mts., 870 m, 12.vi.2008 | 41.986014°N, 23.971926°E | leg. Keresztes L. | |
7 | Hungary, Szobi, Ipolytölgyes, Börzsönyi Mts., 358 m, 2.vi.2016 | 47.910624°N, 18.821948°E | leg. Török E. | |
1 | Romania, Cluj, Gilău Mts., Pecica brook, 443 m, 16.iv.2016 | 46.733137°N, 23.552135°E | leg. Keresztes L. | |
3 | Romania, Poiana Mărului, Șureanu Mts., 1444 m, 16.vi.2008 | 45.316562°N, 22.517324°E | leg. Keresztes L. | |
1 | Romania, Rimetea, Bedeleuilui Mts., 531 m, 28.V.2007 | 46.448873°N, 23.570063°E | leg. Keresztes L. | |
8 | Romania, Sasca Română, Almașului Mts., 266 m, 8.v.2009 | 44.926365°N, 21.782738°E | leg. Keresztes L. | |
9 | Romania, Valea Iadului, Bihor Mts., Leșu lake, 691 m, 21.v.2006 | 46.745951°N, 22.556599°E | leg. Keresztes L. | |
2 | Serbia, Voivodina, Šušara, Deliblatska Peščara, 111 m, 10.vii.2013 | 44.831943°N, 21.111992°E | leg. Török E. | |
P. (Pa.) longicauda (Tonnoir, 1919) | 3 | Romania, Luncavița, Măcin Mts., 151 m, 1.vi.2006 | 45.221240°N, 28.320892°E | leg. Keresztes L. |
P. (Pa.) paludosa Meigen, 1804 | 1 | Austria, Wien, Hermannskoegel, 326 m, 21.v.2013 | 48.261830°N, 16.302293°E | leg. Graf, W. |
3 | Bulgaria, Fotinovo, Rhodope Mts., 1495 m, 16.vii.2012 | 41.870489°N, 24.344398°E | leg. Keresztes L. | |
4 | Hungary, Nagybörzsöny, Börzsönyi Mts., 350 m, 1.v.2016 | 47.939197°N, 18.859785°E | leg. Török E. | |
8 | Hungary, Szobi, Ipolytölgyes, Börzsönyi Mts., 358 m, 2.vi.2016 | 47.910624°N, 18.821948°E | leg. Török E. | |
2 | Romania, Sândominic, Hăghimaș Mts., Babos Laka, 809 m, 9.vi.2019 | 46.573520°N, 25.822180°E | leg. Keresztes L. | |
1 | Romania, Voșlobeni, După Luncă peat bog, 757 m, 6.vi.2017 | 46.670458°N, 25.659906°E | leg. Keresztes L. | |
2 | Serbia, Fruška Gora National Park, Čerević, 501 m, 5.vii.2018 | 45.156725°N, 19.738838°E | leg. Keresztes L. | |
P. (Pa.) pollux sp. nov. | 1 | North Macedonia, Novo Selo, Mavrovo lake, 990 m, 29.vi.2017 | 41.721355°N, 20.830103°E | leg. Török E. |
P. (Pa.) resseli Theischinger, 1978 | 0 | literature data: Fasbender, 2014 | – | – |
P. (Pa.) silvicola Zwyrtek & Rozkosny, 1967 | 3 | Romania, Voșlobeni, După Luncă peat bog, 757 m, 6.vi.2006 | 46.670458°N, 25.659906°E | leg. Keresztes L. |
P. (Pa.) surcoufi Séguy, 1926 | 0 | literature data: Fasbender, 2014 | – | – |
P. (Pt.) contaminata (Linnaeus, 1758) | 3 | Romania, Florești, marshy area, Someșul Mic, 370 m, 18.v.2019 | 46.749129°N, 23.476040°E | leg. Keresztes L. |
2 | Bulgaria. Primorsko, Ropotamo Nature Reserve, 9 m. 11.vii.2018 | 42.301909°N, 27.727464°E | leg. Keresztes L. | |
Total | 75 |
Low resolution photos of the whole fly and the wings were taken with a stereomicroscope (Zeiss Stemi 2000-C) and a consumer digital SLR camera (Cannon 1100D). Photos of different genital structure parts were taken with a compound microscope (Motic 310 BA) that was equipped with standard plan-achromatic objectives and additionally with objectives from the inverse microscope of the same manufacturer, for work on glycerol without coverslip. The camera was of the high-resolution USB CCD type (Imaging Source Europe GmbH DFK 51AU02). As stacking software, we used the Hugin suite (SourceForge.net). Male genitalia were left overnight in 10% potassium hydroxide (KOH) and for one hour in undiluted glacial acetic acid, to neutralise and wash out the soap that was created from the soft tissues. The male genitalia were then transferred to a larger amount of glycerol to wash out the acid. Afterwards, they were transferred to a drop of glycerol on a slide with rounded excavation. The genitals were dissected, the parts were oriented using the stereomicroscope, and then the slide was carefully transferred to the compound microscope for the taking of photos. Finally, the parts were washed again in 100% isopropanol and embedded permanently in artificial Canada Balsam (Malinol), whereby high resolution photos were taken. Stacking results in general consist of 5-10 single exposures with the stereomicroscope and of 10-50 exposures with the compound microscope.
The types of P. castor sp. nov. and P. pollux sp. nov. are deposited in the Diptera collection of the Faculty of Biology and Geology, Babeş-Bolyai University, Cluj-Napoca (UBB), Romania (DCFGB). The study of P. helena was based on a paratype male from the Zoological Research Museum Alexander Koenig (ZFMK), Bonn, Germany, Museum-Id ZFMK-DIP-00015966.
Paraptychoptera agnes Krzemiński & Zwick, 1993, P. delmastroi Zwick & Stary, 2003, and P. ressli Theischinger, 1978 were not available during the present study, but the detailed morphological description of wing and male terminalia, based on
Morphological characters of male Paraptychoptera specimens used in the phylogenetic analyses.
1 | Antennal segments: (0), segment 3 equal with segments 4+5; (1) segment 3 shorter than segments 4+5 |
2 | Wing R2+3+4: oblique straight (0); curved or angled in the middle (1) |
3 | Wing: R2+3+4 length: (0) > 2 × length of rm; (1) < 2 × length of rm |
4 | Male auxiliary sexual organ on abdominal sternite III: only cluster of setae (0); cluster of setae and distal lobes (1) |
5 | Male auxiliary sexual organ, distal lobe: absent or weakly developed (0), well developed, ventrally projected outer lip depressed in the middle (1) |
6 | Male auxiliary sexual organ, distal lobe: outer lip truncate or straight: (0) absent; (1) present |
7 | Median sclerotized strip of the auxiliary sexual organ with a transverse ornamentation: (0) absent; (1) present |
8 | Epandrial clasper: well sclerotized with squared basal lobe and complex ventral lobe (0); poorly sclerotized without basal lobe (1) |
9 | Epandrial clasper configuration: (0), simple cylindrical; (1), cylindrical with additional subterminal ventral lobes |
10 | Epandrial clasper configuration: (0), simple cylindrical; (1), cylindrical with basal lobes |
11 | Epandrial clasper apex: rounded (0), pointed and curved (1) |
12 | Epandrial subapical process: absent (0); present (1) |
13 | Epandrial subapical process bilobate: absent (0); present (1) |
14 | Epandrial subapical process bilobate, inferior arm chitinous and curved: (0) absent; (1) present |
15 | Epandrial subapical process bilobate, superior arm laterally compressed, lobate, inferior arm truncate: (0) absent; (1) present |
16 | Epandrial subapical process bilobate, both arms chitinous and curved: (0) absent; (1) present |
17 | Epandrial subapical process bilobate, superior arm beak-shaped, inferior arm triangular: (0) absent; (1) present |
18 | Epandrial subapical process with chitinous process having curved apex: (0) absent; (1) present |
19 | Epandrial subapical process with a rounded apex, and strong basal thorn, longer than subapical process: (0) absent; (1) present |
20 | Epanadrial subapical process with rounded aped and strong basal thorns, shorter than subapical process: (0) absent; (1) present |
21 | Hypoproct: reduced to paired rectangular plates in subepandrial membrane (0); hypoproct well developed, triangular or tongue like (1) |
22 | Hypoproct short triangular lobe with rounded apex: (0) absent; (1) present |
23 | Hypoproct long, tongue like process: (0) absent (1) present |
24 | Hypoproct long, tongue like process, apex covered with dense hear: (0) absent (1) present |
25 | Hypoproct long, tongue like process, apex glabrous and bilobate: (0) absent (1) present |
26 | Hypoproct long, tongue like process, apex pointed, harpoon-shaped: (0) absent (1) present |
27 | Hypoproct long, tongue like process, apex truncate or slightly depressed: (0) absent (1) present |
28 | Gonostylus, shape of anterior lobule: (0) scythe-like; (1), lobe-like with rostrum |
29 | Gonostylus, shape of medial lobule: lobe like (0); scythe-like (1) |
30 | Gonostylus, apical stylus of gonostylus apex with strong spines: absent (0); present (1) |
31 | Gonostylus, apical stylus or gonostylus: simple (0); inflated (1) |
32 | Gonostylus, secondary lobe of apical stylus: present (0); absent (1) |
33 | Hypandrium apex terminal division spade like, without trichoid sensilla: (0) present; (1) absent; |
34 | Hypandrium apex terminal division wide spade like, with trichoid sensilla (0) absent; (1) present |
35 | Hypandrium apex terminal division long lobe like process, with rounded apex: (0) absent; (1) present |
36 | Hypandrium apex terminal division long lobe like process with bilobate apex: (0) absent; (1) present |
37 | Hypandrium eversible sac extended anteriorly nearly to margin: (0), absent; (1), present |
38 | Hypandrium basal scale: (0), absent; (1) present |
39 | Hypandrium basal scale chitinous, hat shape: (0), absent; (1) present |
40 | Hypandrium basal scale chitinous, hat shape, with medial lobe (0), absent; (1) present |
41 | hypandrium: basal scale lobe like, well developed (0), absent; (1) present |
42 | Hypandrium basal scale lobe like, weakly developed (0), absent; (1) present |
43 | Aedeagus: ejaculatory apodeme size: (0), larger, than sperm sac; (1), subequal |
44 | Aedeagus subapical plate wide spatulate: (0) present; (1) absent |
45 | Aedeagus subapical plate narrow, apex rounded or pointed: (0) present; (1) absent |
46 | Aedeagus subapical plate narrow, apex depressed: (0) present; (1) absent |
47 | Paramere apical lobes filiform process: (0) present; (1) absent |
48 | Paramere apical lobes with a notch in the middle: (0) present; (1) absent |
49 | Paramere apical lobes tip pointed: (0) present; (1) absent |
50 | Paramere apical lobes tip rounded: (0) present; (1) absent |
51 | Paramere lateral arms reduced: (0) present; (1) absent |
52 | Paramere lateral arms long, beak shaped: (0) present; (1) absent |
53 | Paramere lateral arms short, truncate: (0) present; (1) absent |
Morphological characters were selected based on the world phylogenetic revision of Ptychopteridae (Fasbender, 2014), but completed with new morphological data. Ptychoptera (Ptychoptera) contaminata was considered as an outgroup taxon. Missing data were coded as ‘?’. The list of morphological characters is presented in Table
Matrix of the 53 morphological items of data used in the phylogenetic analyses. For the description of characters and character states see text. Missing data were coded as ‘?’.
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 | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
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 | |
agnes | ? | 1 | 1 | 1 | ? | ? | ? | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | |
castor sp. nov. | 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 | |
delmastroi | ? | ? | ? | ?1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
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 | |
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 | |
lacustris | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | |
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 | |
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 | |
pollux sp. nov. | 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 | |
resseli | ? | ? | ? | ? | 0 | ? | ? | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | |
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 | |
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 | |
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 |
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 |
castor sp. nov. | 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 |
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 |
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 |
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 |
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 |
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 |
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 |
pollux sp. nov. | 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 |
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 |
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 |
The morphological data matrix was managed with Mesquite 3.5 (
Holotype . male, Albania: Tragjas municipality, Rrepet e Izvorit, Vlora district, sweeping the vegetation near a limnocren karst spring with large basin and muddy shore with reeds, 30.iv, 2019, 17 m, leg. M. Henning, 40.323132°N, 19.510031°E. Institutional id for specimen is DCFBG-PT-0002.
. Ptychoptera (Paraptychoptera) castor sp. nov. is known only from a single male collected near a limnocrene karst spring with muddy shore invaded by rich vegetation at Repet y Izvorit, Tragjas, Albania (Fig.
Ptychoptera helena, paratype male (ZFMK) a habitus male b head and thorax dorsal c right wing d epandrium, dorsal e subapical lobe of epandrium, ventral f gonocoxite and gonostylus complex, dorsal g gonostylus anterior and medial lobules, caudal h hypandrium, caudal i paramere, ventral j aedeagal complex, dorsal.
. Medium-sized species, body length 7.3 mm, wing length 8 mm. Head and thorax shiny black, almost glabrous, pleuron almost uniformly brownish, some obvious pale setae only above halter. Head shiny brownish, labrum pale brownish to yellow. Antennae with 15 segments. Scape elongate cylinder, pedicel globular, yellowish, as the half of the first flagellar segment. Remainder flagellomeres blackish brown (Fig.
Thorax dorsally black with metallic blue shining, narrow pronotum, base of postnotum and large parts of episternum, epimeron, and metapleuron pale brownish. Coxae and legs yellowish, apex of femur, narrow base and apex of tibia, tarsal segments brownish. Wing with three transverse bands of well-developed confluent dark spots close to anterior margin on basal, middle and distal part of otherwise clear or pale yellowish membrane. Additionally, isolated dark spots are present on both sides of the middle dark band at the level of Sc and at the distal end of R3 (Fig.
First abdominal tergite blackish to dark brown with metallic shining, only a narrow yellow stripe near the distal, tergite 2 large part yellowish with brown spot in the middle, distal part shiny black, tergite 3 brownish, tergite 4 and all distal tergites brownish black. Genitalia pale brown. Narrow sternites pale brown at base, becoming yellowish towards the auxiliary sexual organ on segment III. Sternites 4-7 medially reduced to a narrow band with a deep notch in the middle at proximal margins.
Auxiliary sexual organ less developed than in the other members of Paraptychoptera, excepting P. lacustris and P. helena. Sternite 3 with thin long golden hair fringes on sides; its bare middle part lacking the transversally sculptured median sclerite, but distal brownish patch is present at distal end, close to the deep pouch of the auxiliary sexual organ. Distally sternite 3 with deep pouch of the auxiliary sexual organ. Two caudal lips of the pouch less developed, one smooth lateral lobe on each side, covered with dense hair fringe in their interior part. Lateral lips separated by a deep furrow leading to a small oval sclerite inside the pouch and two lateral lobes covered with fine sculptures.
Male terminalia. Epandrium with distinct collar, deeply and widely emarginated behind, hypoproct long lobe-like and densely hairy (Fig.
Female unknown.
. The specific epithet is named after Castor, a god from Greek mythology, the twin brother of Helena, because of its close morphological similarity with P. helena.
Holotype . male, North Macedonia: Mavrovo, Novo Selo, sweeping the vegetation near a small outflow from Mavrovo Lake, 29. vi, 2017, 990 m, leg. E. Török, 41.721355°N, 20.830103°E. Institutional id for specimen is DCFBG-PT-0003.
. Paraptychoptera pollux sp. nov. is known only from a single male collected near a small overflow of the Mavrovo Lake with muddy shore invaded by rich vegetation (Fig.
. Medium-sized species, highly similar to its sibling species, P. castor sp. nov. and P. helena. Body length 7.9 mm, wing length 8.5 mm. Head and thorax similar to P. castor. Antennae with 15 segments. Scape elongate, cylindrical, yellowish brown, pedicel globular, pale brown, flagellar segments uniformly dark brown (Fig.
Thorax dorsally brownish black with metallic blue shining, narrow pronotum, base of postnotum and large parts of episternum, epimeron and metapleuron pale brownish. Coxae orange, legs yellowish, apex of femur, narrow base and apex of tibia, tarsal segments brownish. Wing with three transverse bands of well-developed confluent dark spots in the anterior part of base, middle and distal part of otherwise clear or pale yellowish membrane. Basal spot of the wing more reduced. Distal band interrupted close to ventral edge. Isolated dark spots are present at distal end of Sc and R3 (Fig.
First abdominal tergite blackish to dark brown with a metallic blue shining, narrow yellow stripe close to distal end, well developed yellow band in tergite 2 with a black spot in the middle. Tergite 3 black, covered with yellow setae, the remaining tergites brownish black. Genitalia pale brown. Narrow sternites pale brown at base, becoming yellowish towards the auxiliary sexual organ on segment III. Sternites 4-7 medially reduced to a narrow band with a deep notch in the middle at proximal margins.
Auxiliary sexual organ highly similar to P. castor sp. nov. and P. helena. Male terminalia. Epandrium with distinct collar, deeply and widely emarginated behind, hypoproct long tongue-like and furry (Fig.
Female unknown.
. The specific epithet is named after Pollux, the twin brother of Castor in Greek mythology, known together as the Dioscuri, both twin brothers of Helena, because together with P. castor they share close morphological similarity with P. helena and all together they form a distinct monophyletic unit among Paraptychoptera, as was recovered by our cladistic analysis (Fig.
.The parsimony analyses of the 53 different morphological characters selected in the present study resulted in a single most parsimonious tree (Fig.
As shown in our parsimony analyses (Fig.
Among this group, a distinct lineage is represented by P. agnes, highly different from all other members of the clade by the presence of a conspicuous epandrial lobe, with a ventral lobe close to the base, and a transverse projection with a comb of long setae, unique only to this species, in addition to the particularly shaped hypoproct and epandrial subapical process, and also by the uniquely shaped secondary lobe of the apical stylus inflated and globulose at apex, besides the details of hypandrium and parameres (characters 8, 12, 13, 23, 30, 37, 47).
The present cladistic analyses recovered the “lacustris” species group as a monophyletic unit, which was also noted by
According to
A key to the Paraptychoptera species was recently provided by Fasbender in 2014 in his global revision of the world Ptychopteridae, including only four Paraptychoptera species, instead of the twelve currently known from the Western Palaearctic area. The present key is mostly based on his comprehensive morphology data, but incorporates additional morphological details from the remainder of the Paraptychoptera species, including the two newly described species, P. castor sp. nov. and P. pollux sp. nov. The North African P. surcoufi is not included the key because no material was available for the current investigation, nor was there any detailed information in the literature to the best of our knowledge, but its distinctness from the P. helena and related species is obvious according to
1 | Apical stylus of gonostylus reduced | P. ressli Theischinger, 1978 |
– | Apical stylus of gonostylus well developed | 2 |
2 | Epandrial clasper lacking lateral swelling, basal division of hypandrium with a weakly developed membranous basal scale | 3 |
– | Epandrial clasper with lateral swelling (mostly reduced in P. longicauda), basal division of hypandrium with a well-developed, chitinous projection of different shapes | 7 |
3 | Epandrial clasper lobes complex, with a transverse projection with a comb of long setae at apex and the presence of a subterminal ventral extension, secondary lobe of the apical stylus expanded into a large balloon-shaped structure | P. agnes Krzemiński & Zwick, 1993 |
– | Epandrial lobes simple, apical stylus simple, with apex tapering and rounded | 4 |
4 | Subapical sclerite present at base of epandrial clasper, hypoproct long, tongue-like | 5 |
– | Subapical sclerite absent, hypoproct short, triangular | P. lacustris Meigen, 1930 |
5 | Hypoproct furcate apically | P. helena Peus, 1958 |
– | Hypoproct not furcate apically | 6 |
6 | Hypoproct apex rounded, with a shallow depression in the middle, apical stylus and secondary lobe subequal in length | P. pollux Keresztes & Török, sp. nov. |
– | Hypoproct apex tapered with pointed apex, apical stylus twice as long as the secondary lobe | P. castor Keresztes & Kappert, sp. nov. |
7 | Epandrial clasper lobe very long, more than twice as long as epandrium length. Apical stylus of gonostylus pendulant, apex of terminal division of hypandrium elongate ribbon-like projection with bifurcate tip, basal scale of hypandrium with no medial triangular projection | P. longicauda Tonnoir, 1919 |
– | Epandrial clasper lobe short, shorten than epandrium length. Apical stylus of gonostylus not pendulant, apex of terminal division of hypandrium widened, spatulate, basal scale of hypandrium with medial triangular projection | 8 |
8 | Secondary lobe of apical stylus absent | P. paludosa Meigen 1804 |
– | Secondary lobe of apical stylus present | 9 |
9 | Subapical sclerite hook-like | P. delmastroi Zwick & Stary, 2003 |
– | Subapical sclerite with a paddle-like ventrally projected division | 10 |
10 | Subapical sclerite with the paddle-like ventrally projected division inflated at apex. With a short hook subterminally and dorsally projected division triangular | P. silvicola Zwyrtek & Rozkosny, 1967 |
– | Subapical sclerite with ventrally projected arms paddle-like, rounded at apex, without subterminal hooks, dorsally projected division rectangular | P. handlirschi Czizek, 1919 |
We give thanks to the curator, Dr. Ximo Mengual, head of the Section Diptera, ZFMK, Bonn, for the loan of the male specimen of P. helena paratype and granting permission to dissect the genitalia for closer examination. We also thank to Anna Dénes, Babes-Bolyai University, Cluj Napoca, Romania for the habitat photograph of P. castor sp. nov. Furthermore, Mr. Matthew Copley kindly reviewed the English version of the draft manuscript.
The present work received financial support from a national grant offered by the Executive Unit for Financing Higher Education, Research, Development and Innovation, UEFSCDI, of the Ministry of Education and Research, PN-III-P2-2.1-PED-2019-0214; nr. 476PED/2020.