Research Article |
Corresponding author: Avar Lehel Dénes ( avar.lehel@gmail.com ) Corresponding author: Levente-Péter Kolcsár ( kolcsar.peter@gmail.com ) Corresponding author: Lujza Keresztes ( keresztes2012@gmail.com ) Academic editor: Jukka Salmela
© 2016 Avar Lehel Dénes, Levente-Péter Kolcsár, Edina Török, Lujza Keresztes.
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:
Dénes A-L, Kolcsár L-P, Török E, Keresztes L (2016) Taxonomic revision of the Carpathian endemic Pedicia (Crunobia) staryi species–group (Diptera, Pediciidae) based on morphology and molecular data. ZooKeys 569: 81-104. https://doi.org/10.3897/zookeys.569.7458
|
Three new species of the genus Pedicia, subgenus Crunobia (Diptera: Pediciidae) belonging to the staryi group are described on the basis of a combination of molecular and morphology datasets, and a key to discriminate between species of the subgenus Crunobia is added. Geographic projection of the identified taxa suggests insular-like distribution and shows the importance of the Carpathians as a genetic center which is home to an exceptionally high aquatic diversity in Europe.
Crunobia , cryptic lineage, new species, re-description, endemism, identification key
The Holarctic genus Pedicia Latreille, 1809 is a small taxonomic unit with three subgenera, Amalopis Haliday, 1856 (with 12 species group taxa, including 2 subspecies of P. tenuiloba Alexander, 1957) and Crunobia Kolenati, 1859 (with 16 species). The distribution of these two subgenera is limited within the Palaearctic area, while the remaining subgenus Pedicia (with 33 species) extends to the whole Holarctic region (
Adult specimens were collected between 2003 and 2015 (Suppl. material
Molecular techniques. Tissue samples were collected from 152 individuals of the P. staryi species group (83 individuals of P. staryi, 17 of P. apusenica, 9 of P. lobifera, 6 of P. spinifera and 37 of P. straminea) and deposited in 96 well plates containing 30 μl of 96% ethanol. Four species (P. littoralis Meigen, 1804, P. riedeli Lackschewitz, 1940, P. nielseni Slipka, 1955 and P. zernyi Lackschewitz, 1940) representing the littoralis species group were used as outgroups for this study. The molecular processing—DNA extraction, PCR amplification, gel electrophoresis for PCR product checking, PCR cycle sequencing and sequencing—of 126 specimens and of the outgroup species was done at the Canadian Centre for DNA Barcoding (
The sequences were downloaded and aligned using Clustal W in MEGA6 (
The COI alignment of the five species belonging to the P. staryi species group was represented by 44 haplotypes. The ML and BI analysis resulted in congruent tree topologies with high posterior probability (PP) and bootstrap (BP) values, showing a clade that includes four well-differentiated lineages of P. staryi (Fig.
Bayesian inference (BI) tree with hypopygium profiles of species. Posterior probabilities (PP) and bootstrap values (BP, %) for the nodes are shown under the branches. Carpathian endemic species of staryi group are mapped. Color codes: Pedicia lobifera (gray square), P. staryi (brown point), P. apusenica (blue point), P. carpianica (purple point), P. costobocica (green point), P. roxolanica (orange point).
Pairwise K2P distances between species within Pedicia staryi species group.
P. straminea | P. spinifera | P. lobifera | P. apusenica | P. roxolanica | P. staryi | P. costobocica | |
---|---|---|---|---|---|---|---|
P. spinifera | 16,2 | ||||||
P. lobifera | 12,8 | 12,4 | |||||
P. apusenica | 12 | 11,8 | 9,61 | ||||
P. roxolanica | 11,5 | 12,5 | 9,82 | 6,16 | |||
P. staryi | 13,2 | 13,1 | 11,4 | 6,1 | 6,85 | ||
P. costobocica | 13,3 | 13,4 | 11,2 | 9,25 | 8,58 | 9,05 | |
P. carpianica | 14 | 13,4 | 11,6 | 8,94 | 8,77 | 9,34 | 4,91 |
Holotype male and three paratype males collected in Romania, Apuseni Mountains, Padiș Protected Area, 1 km west of Poiana Vărășoaia, near the Cetatea Rădesei Cave, 1320 m, 46°37.806'N, 22°42.486'E, 21.July 1999, leg. L. Ujvárosi (L. Keresztes). The holotype (CN: TI96) and paratypes (CN: TI97, TI98, TI99) are deposited in the Museum of Zoology of the Babeș–Bolyai University (MZBBU), Cluj Napoca, Romania.
Romania: Apuseni Mountains, Padiș Protected Area, 1 km west of Poiana Vărășoaia, near the Cetatea Rădesei Cave, 1320 m, 46°37.800'N, 22°42.480'E, 03.Aug.2003, 9 ♂♂, leg. L. Keresztes, 20.Aug.2013, 12 ♂♂, leg. A.L. Dénes; Apuseni Mountains, Stâna de Vale, 1140 m, 46°41.250'N, 22°36.546'E, 19.Aug.2013, 1♂ 1♀ leg. A.L. Dénes; Apuseni Mountains, 2 km west of Rogojel, 1290 m, 22°48.528'E, 22°48.528'E, 14.July.2014, 3 ♂♂ leg. L.P. Kolcsár; Apuseni Mountains, Boga, Boga Valley, 700 m, 46°36.576'N, 22°40.674'E, 15.Aug.2015, 1 ♂, leg. E. Török. Materials are stored in 96% ethanol or pinned dry and are deposited in the Diptera Collection of the Faculty of Biology and Geology, Cluj-Napoca, Romania
The species is distinguished from all other species of the Pedicia staryi group by the following combination of characteristics: all flagellomeres are almost uniformly colored, and there are no dark lines between antennae; the abdominal stripe starts from the second segment; the tip of the last palpus segment is darker than other segments; 9th tergite has a rounded median lobe, with a small apical emersion.
Large species of a yellowish orange color (Fig.
2–5 Pedicia apusenica male: 2 lateral habitus 3 inner lateral view of the gonocoxite 4 male hypopygium dorsal view 5 male hypopygium ventral view 6–9 Pedicia roxolanica sp. n. holotype male: 6 lateral habitus 7 inner lateral view of the gonocoxite 8 male hypopygium dorsal view 9 male hypopygium ventral view.
Holotype male and two paratype males collected in Romania, Baiu Mountains, Azuga, Limbășel Valley, 1200 m, 45°29.574'N, 25°35.910'E, 26.Aug.2014, L.P. Kolcsár. The pinned dry holotype (Catalog Number–CN: TI101) and paratypes (CN: TI102, TI103) are deposited in the Museum of Zoology of the Babeș–Bolyai University (MZBBU), Cluj Napoca, Romania.
Romania: Bucegi Mountains, Sinaia, Cota 1400, 1400 m, 45°21.258'N, 25°31.278'E, 21.July.2004, 4 ♂♂, leg. L. Keresztes; Iezer–Păpușa Mountains, Lerești, Voina Hut, 970 m, 45°26.526'N, 25°2.670'E, 03.Aug.2006, 4 ♂♂, leg. L. Keresztes; Bucegi Mountains, Sinaia, Peleș Valley, 1300 m, 45°22.092'N, 25°30.978'E, 04.Aug.2006, 1 ♂, leg. L. Keresztes; same site, 26.July. 2013, 6 ♂♂, leg. L. Keresztes and Á. Péter; Baiu Mountains, Azuga, Casariei Valley, 1025 m, 45°26.868'N, 25°34.260'E, 20.June.2013, 1 ♂, leg. E. Török & L. Keresztes; Bucegi Mountains, Moroeni, Dichan Hut, 1575 m, 45°19.506'N, 25°27.294'E, 26.July.2013, 1 ♂, leg. L. Keresztes & Á. Péter; Baiu Mountains, Azuga, Limbășel Valley, 1200 m, 45°29.574'N, 25°35.910'E, 26.Aug.2014, 3 ♂♂, leg. L.P. Kolcsár; Bucegi Mountains, Moroeni, Cheile Orzei, 1366 m, 45°17.682'N, 25°25.434'E, 18.Aug.2015, 2 ♂♂, leg. L. Keresztes. All materials are stored in 96% ethanol and deposited in the Diptera Collection of the Faculty of Biology and Geology, Cluj-Napoca, Romania.
The species is named after an ancient population from the southern border of the Carpathians, suggesting its ancient origin, which is revealed by deep genetic and morphological structuring.
The new species is distinguished from all other species of the Pedicia staryi group by the following combination of characteristics: all flagellomeres are almost uniformly colored; the abdominal stripe starts from the first segment; the tip of last palpus segment is the same color as the other segments; 9th tergite has a rounded or rarely five angled median lobe, with a notch on the tip or rarely with a small apical emergence.
Large species of a yellowishorange color (Fig.
Female is unknown.
Holotype male and paratype male collected in Romania, Gutâi Mountains, Baia Sprie, Arinieși brook, 1015 m, 47°43.068'N, 23°44.628'E, 15.May.2014, leg. L.P. Kolcsár. The pinned dry holotype (CN: TI104) and paratype (CN: TI105) are deposited in the Museum of Zoology of the Babeș–Bolyai University (MZBBU), Cluj Napoca, Romania.
Romania: Gutâi Mountains, Baia Sprie, Gutâi Pass, 1070 m, 47°41.898'N, 23°49.128'E, 26.May.2012, 2 ♂♂, leg. E. Török, L.P. Kolcsár & L. Keresztes; Gutâi Mountains, Baia Sprie, Gutâi Pass, 47°41.634'N, 23°47.226’, 15.May.2013, 1 ♂ 1 ♀, E. Török & L. Keresztes; Gutâi Mountains, Baia Sprie, Arinieși brook, 1015 m, 47°43.068'N, 23°44.628'E, 15.May.2014, 7 ♂♂, leg. L.P. Kolcsár. All material is stored in 96% ethanol and deposited in the Diptera Collection of the Faculty of Biology and Geology, Cluj-Napoca, Romania.
The species is named after an ancient population from the northern part of the Eastern Carpathians suggesting its ancient origin revealed by deep genetic and morphological structuring.
The new species is distinguished from all other species of the Pedicia staryi group by the following combination of characteristics: the last 1–2 antennal segments are darker than others; black line between antennae; the abdominal stripe starts from the second segment; mean body size reach 13 mm; pedicel and scape darker than first flagellomere, face dark brown; 9th tergite has a rounded median lobe, usually with a notch on the tip.
Medium sized species of a yellowish orange color (Fig.
10–13 Pedicia costobocica sp. n. holotype male: 10 lateral habitus 11 inner lateral view of the gonocoxite 12 male hypopygium dorsal view 13 male hypopygium ventral view 14–17 Pedicia carpianica sp. n. holotype male: 14 lateral habitus 15 inner lateral view of the gonocoxite 16 male hypopygium dorsal view 17 male hypopygium ventral view.
Female has a body length of 12 mm, wing length 11 mm, antenna 1.6 mm. General color is yellowish. Head is dark orange, frontal part is light brown mostly around the antennas, a narrow dark line is present between antennas. Pedicel and scape are dark orange. The antenna has 14 yellowish orange flagellomeres. Palpus is uniformly yellowish orange. The dorsal and lateral parts of the thorax are yellowish. Legs have coxae and trochanters yellowish orange. Wings are well-developed, having the ability to fly. Wing venation is yellowish orange, small spots are present at Sc2, and at around r–m. Abdomen is uniformly light orange. Female terminalia has wide cercus, dagger-like, with tip raised upward. Hypovalvae are each wide, darker than the tenth sternite with nine pairs of curved knitting needle-like strong setae at the dorsal margin. The distal seta is isolated from the rest. Two pairs of sensory setae are visible at the end of the hypovalvae, and they extend beyond the end of hypovalvae.
Holotype male and two paratypes males collected in Romania, Rodnei Mountains, Anieș, Tomnaticul Valley, 700 m, 47°27.768'N, 24°45.696'E, 19.Aug.2014, leg. L.P. Kolcsár. The pinned dry holotype (CN: TI106) and paratypes (CN: TI107, TI108) are deposited in the Museum of Zoology of the Babeș–Bolyai University (MZBBU), Cluj Napoca, Romania.
Romania: Rodnei Mountains, Rodna, Vinului Valley, 1000 m, 47°30.918'N, 24°50.094'E, 15.Aug.2008, 2 ♂♂, leg. L. Keresztes; Rodnei Mountains, Cormaia, Cormaia Valley, 750 m, 47°26.328'N, 24°39.702'E, 26.Aug.2010, 11 ♂♂, leg. R. Vaida; Rodnei Mountains, Anieș, Izvorul Mare Valley, 1220 m, 47°32.274'N, 24°40.362'E, 28.July.2011, 1 ♂, leg. R. Vaida; Rodnei Mountains, Anieș, Cepelor Spring, 1165 m, 47°31.404'N, 24°45.024'E, 19.Aug.2014, 3 ♂♂ leg. L.P. Kolcsár; Rodnei Mountains, Anieș, Tomnaticul Valley, 700 m, 47°27.768'N, 24°45.696'E, 19.Aug.2014, 11 ♂♂ leg. L.P. Kolcsár; Rodnei Mountains, Valea Mare, Rotunda Pass, 1165 m, 47°31.812'N, 25°0.810'E, 19.Aug.2014, 1 ♂, leg. L.P. Kolcsár. Material is stored in 96% ethanol or pinned dry and deposited in the Diptera Collection of the Faculty of Biology and Geology, Cluj-Napoca, Romania.
The species is named after an ancient population from the Eastern Carpathians suggesting its ancient origin revealed by deep genetic and moprhological structuring.
The new species is distinguished from all other species of the P. staryi group by the following combination of characteristics: the last 1–2 antennal segments are darker than the others, black line between antennae; the abdominal stripe starts from the second segment; mean body size reaches 15.4 mm; pedicel and scape have the same color as the first flagellomeres; only the lump is darker than other parts of the head; 9th tergite has a rounded or five angled median lobe, usually with a notch on the tip.
Large species with yellowish orange color (Fig.
18–24 Pedicia staryi male: 18 lateral habitus 19 inner lateral view of the gonocoxite 20 male hypopygium dorsal view 21 male hypopygium ventral view 22 male hypopygium caudal view 23 head dorsal view 24 antenna 25–26 Pedicia roxolanica sp. n.: 25 head dorsal view 26 dorsal view of the cranial part of abdomen 27 Pedicia carpianica sp. n.: dorsal view of the cranial part of abdomen.
Female is unknown.
Holotype male Ukraine: Ivano–Frankivsk Oblast, environs settlements Vorokhta, Verkhovinsk region, 18.VI 1962 leg. E.N Savtshenko (Savchenko); deposited in the National Museum of Natural History, Kiev, Ukraine. Based of photos taken by Valery A. Korneyev.
Ukraine: Gorgan Mt., Bukovel, 1120 m, 48°23.340'N, 24°26.010'E, 29.July.2014, 21 ♂♂ 1 ♀, leg. E. Török & L.P. Kolcsár. Romania: Rodnei Mountains, Borșa, Cailor Waterfall, 1700 m, 47°35.292'N, 24°39.510'E, 16.June.2010, 1 ♂, leg. R. Vaida; Maramureș Mountains, Borșa, Vișeu River, 1000 m, 47°37.374'N, 24°48.582'E, 17.May.2013, 1 ♂ leg. E. Török; Rodnei Mountains, Gura Lalei, Lalei Valley, 1200–1800 m, 47°32.070'N, 24°54.930'E, 20.July.2013, 13 ♂♂, leg. L. Keresztes & L.P. Kolcsár; Maramureș Hills, Leordina, 490 m, 47°46.752'N, 24°14.574'E, 17.May.2014, 3 ♂♂, leg. L.P. Kolcsár; Maramureș Mountains, Repedea, Repedea Valley, 790 m, 47°53.202'N, 24°23.418'E, 17.May.2014, 20 ♂♂ 1 ♀, leg. L.P. Kolcsár; Maramureș Mountains, Gura Lalei, Bistrița River, 1025 m, 47°33.858'N, 25°1.800'E, 20.Aug.2014, 5 ♂♂ leg. L.P. Kolcsár. Material stored in 96% ethanol or pinned dry and is deposited in the Diptera Collection of the Faculty of Biology and Geology, Cluj-Napoca, Romania.
The species is distinguished from all other species of the Pedicia staryi group by the following characters combination: the last 1–3 antennal segments are darker than others; black line between antennae; two brown spots on the scutum; abdominal dorsal stripe starts from first abdominal segment; 9th tergite has a rounded or five angled median lobe, sometimes with a notch on the tip.
Large species having general color yellowish orange (Fig.
1 | Gonostylus has dorsally only two or three big thorn-like dark spines (Figs |
2 |
– | Gonostylus has dorsally 5–11 claw-like dark spines (Figs |
(staryi species group) 11 |
2 | Gonostylus extends ventrally and forms a big obtuse projection, densely covered by black spinule (Figs |
3 |
– | Gonostylus without such projection (Figs |
8 |
3 | Wings strongly reduced, much shorter than the abdomen | P. (C.) semireducta Savchenko, 1978 |
– | Wings well-developed and as long or sometimes longer than the abdomen (Fig. |
4 |
4 | Gonostylus ventral projection short, not wider than gonocoxite, 9th abdominal tergite with a small triangular lobe with a big notch in the middle of the distal margin (Fig. |
P. (C.) tjederi Mendl, 1974 |
– | Gonostylus ventral projection long, wider than the gonocoxite, 9th abdominal tergite is differently shaped (Figs |
5 |
5 | Gonostylus ventral projection (in caudal view) is densely covered by black spinule only in the ventral parts (Fig. |
P. (C.) zangheriana Nielsen, 1950 |
– | Gonostylus ventral projection (in caudal view) uniformly covered by black spinule (Fig. |
6 |
6 | Lateral side of scutum, prescutum and wings’ base is lighter (Fig. |
P. (C.) pallens Savchenko, 1978 |
– | Lateral side of scutum, prescutum and wings’ base is darker | 7 |
7 | Wings are transparent, with more or less conspicuous pattern, 9th abdominal tergite is narrow, narrower at its base (Fig. |
P. (C.) persica Alexander, 1975 |
– | Wings are yellowish, only with an obscure dark pattern; 9th tergite wider at its base | P. (C.) zernyi (Lackschewitz, 1940) |
8 | Ochre–yellow species. Femora uniformly colored, the tip of femora without dark rings (Fig. |
P. (C.) littoralis (Meigen, 1804) |
– | Dark colored species. Femora brown, dark–brown, basally lighter (Fig. |
9 |
9 | Thorax mainly brownish yellow, with brown or yellowish brown prescutal stripes, 9th abdominal tergite lobe with a big notch (Fig. |
P. (C.) dispar Savchenko, 1978 |
– | Thorax mainly gray, with grayish–brown prescutal stripes; 9th abdominal tergite lobe without a notch (Figs |
10 |
10 | Medial lobe of the 9th abdominal tergite triangular, narrowed towards the tip (Figs |
P. (C.) riedeli (Lackschewitz, 1940) |
– | Medial lobe of the 9th abdominal tergite hexagonal, slightly narrow at the base. Wings usually yellowish (Figs |
P. (C.) nielseni (Slipka, 1955) |
11 | Gonostylus extended ventrally and forms a big obtuse projection (Figs |
P. (C.) straminea (Meigen, 1838) |
– | Gonostylus without this projection (Figs |
12 |
12 | Interbases with a spine-like outgrowth (Figs |
P. (C.) spinifera Starý, 1974 |
– | Interbases without spine-like outgrowth (Fig. |
13 |
13 | Gonocoxite has on the top a conspicuous, isolated rounded lobe (figs 50, 51) | P. (C.) lobifera Savchenko, 1986 |
– | Gonocoxite on the top without such rounded lobe (Figs |
14 |
14 | All flagella almost uniformly colored, none darker, no black or dark brown line between antennae (Fig. |
15 |
– | The last 1–3 antennal segments are darker than others (Fig. |
16 |
15 | Abdominal dorsal stripe starts from the second abdominal segment (Fig. |
P. (C.) apusenica Ujvárosi & Starý, 2003 |
– | The abdominal stripe starts from the first segment (Fig. |
P. (C.) roxolanica Kolcsár, Keresztes & Dénes, sp. n. |
16 | Two brown spots on the scutum, abdominal dorsal stripe starts from first abdominal segment (Fig. |
P. (C.) staryi Savchenko, 1978 |
– | No brown spots on scutum, stripe starts from second abdominal segment (Fig. |
17 |
17 | Smaller species, mean body size reach 13 mm; pedicel and scape darker than first flagellomere, face dark brown | P. (C.) costobocica Kolcsár, Keresztes & Dénes, sp. n. |
– | Larger species, mean body size is 15.4 mm; pedicel and scape is the same color as the first flagellomere, only the lump is darker than other parts of the head | P. (C.) carpianica Kolcsár, Keresztes & Dénes, sp. n. |
28–31 Pedicia pallens male: 28 lateral habitus 29 inner lateral view of the gonocoxite 30 male hypopygium dorsal view 31 male hypopygium caudal view 32–35 Pedicia littoralis male: 32 lateral habitus 33 inner lateral view of the gonocoxite 34 male hypopygium dorsal view 35 male hypopygium caudal view.
36–38 Pedicia nielseni male: 36 lateral habitus 37 inner lateral view of the gonocoxite 38 male hypopygium dorsal view 39–40 Pedicia riedeli: 39 inner lateral view of the gonocoxite 40 male hypopygium dorsal view 41–43 Pedicia zangheriana male: 41 inner lateral view of the gonocoxite 42 male hypopygium dorsal view 43 male hypopygium caudal view.
44–47 Pedicia straminea: 44 male lateral habitus 45 inner lateral view of the gonocoxite 46 male hypopygium dorsal view 47 male hypopygium caudal view 48–49 Pedicia spinifera: 48 inner lateral view of the gonocoxite 49 male hypopygium dorsal view 50–51 Pedicia lobifera: 50 inner lateral view of the gonocoxite 51 male hypopygium dorsal view.
The three newly described species, P. costobocica, P. carpianica and P. roxolanica can clearly be attributed to the “staryi” species group sensu
The rhitral biome in Central Europe is home to an exceptionally high aquatic diversity with an important number of endemics, in contrast with the arboreal elements, which are related mostly with some classical Mediterranean core areas (
We would like to express our sincere thanks to Valery A. Korneyev for the photos of Pedicia staryi and P. lobifera holotypes. We are also grateful to Nikolai Paramonov for translating the original Russian descriptions of P. staryi and P. lobifera. We also thank Jukka Salmela and the two reviewers, Chen Young and Herman de Jong for their helpful comments. This study was financially supported partly by a grant of the Romanian Ministry of National Education, CNCS–UEFISCDI–nr. PN–II–ID–PCE–2012–4–0595. Additional funding comes from the research project number P 23687-B17, Pl: J. Waringer founded by the Austrian Science Found (FWF). During preparation of the manuscript AL Dénes and LP Kolcsár received financial support with scholarships from the Collegium Talentum, Hungary. We thank the staff at the Canadian Centre for DNA Barcoding at the University of Guelph for molecular analysis. The English proofreading of the article was done by Thomas Cooper.
Sampling information
Data type: Collection data
Explanation note: Information for sampling of the five microendemic Carpathian species (Pedicia lobifera, P. staryi, P. apusenica, P. carpianica, P. costobocica, P. roxolanica), containing the name of the species, the date of collection, with localities, mountains, locations and geographic coordinates