Research Article |
Corresponding author: Elisabeth Stur ( elisabeth.stur@ntnu.no ) Academic editor: Gunnar Kvifte
© 2019 Fabio Laurindo da Silva, Elisabeth Stur.
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:
Silva FL, Stur E (2019) Pentaneurella katterjokki Fittkau & Murray (Chironomidae, Tanypodinae): redescription and phylogenetic position. ZooKeys 833: 107-119. https://doi.org/10.3897/zookeys.833.30936
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The monotypic genus Pentaneurella Fittkau & Murray was originally described based on larvae, pupal exuviae and pharate males. The latter prevented the observation of key features, such as wing dimensions, abdominal coloration pattern, and hypopygial apodemes (sternapodeme and phallapodeme), and the description of the adult male was considered incomplete by the authors. Herein, the adult female of Pentaneurella katterjokki is described for the first time, and the adult male, pupa and larva are redescribed and figured based on specimens recently collected in Germany and Norway. We also discuss the phylogenetic position of Pentaneurella.
DNA barcodes, immature stages, non-biting midges, Palearctic, Pentaneurini, taxonomy
Non-biting midges of the genus Pentaneurella are medium-sized dipterans with a Palearctic distribution. Larvae are only known from springs and spring-fed streams in Swedish Lapland and from a mountain stream in northern Norway (
Fourth instar larvae and pupae were sampled with hand nets, while adults were collected using emergence- and Malaise traps. Associations between different life stages were established using DNA barcoding. Alcohol-preserved specimens were dissected, the bodies cleared in 8% KOH, and slide-mounted in Euparal®. Measurement methods are according to
Pentaneura spec. Katterjokk Fittkau, 1962: 372 (description of male)
Pentaneurella katterjokki Fittkau & Murray, 1983: 62 (description of male and immature stages)
Type material
: Holotype pharate male (
Additional material : NORWAY, Oppland, Rondane National Park: Adult male (NTNU-VM slide 201765), Skranglehaugen (P4), 1110 m asl, 61.98270N, 9.80360E, 14–21.vii.2008, leg. T. Ekrem, [BOLD ID: ATNA328]. Adult male (NTNU-VM slide 201767), as previous except for Skranglehaugen (P3), 1115 m asl, 61.98219N, 9.80451E, [BOLD ID: ATNA333]. Adult female (NTNU-VM slide 201768), as previous except for 07–14.vii.2008, leg. T. Hoffstad, [BOLD ID: ATNA335]. Adult female (NTNU-VM slide 201766), Skranglehaugen (P2), 1119 m asl, 61.98141N, 9.80480E, 14–21.vii.2008, leg. T. Ekrem, [BOLD ID: ATNA 331]. Pupa (NTNU-VM slide 201769), Skranglehaugen (P5), 1105 m asl, 61.98346N, 9.80384E, 07–14.vii.2008, leg. T. Hoffstad, [BOLD ID: ATNA338]. Larva (NTNU-VM slide 201764) Skranglehaugen, 1117 m asl, benthos, 61.99186N, 9.80454E, 23.vi.2008, leg. E. Stur, [BOLD ID: ATNA122]. Pupa (NTNU-VM slide 201771) Dørålseter, 1032 m asl, kick sample 3, 61.99347N, 9.80343E, 10.viii.2015, leg. K. Hårsaker, T. Ekrem and M. Majaneva, [BOLD ID: EBAI-Ch122]. Larva (NTNU-VM slide 201770) as previous, [BOLD ID: EBAI-Ch66]. GERMANY, Bayern, Berchtesgaden National Park: Adult male, (NTNU-VM slide 201774), Herrenrointquelle 308, 1250 m asl, 47.57778N, 12.97222E, 26.vii-09.viii.2005, leg. F. Eder, [BOLD ID: ES147]. Adult male, (NTNU-VM slide 201772), Schapbachquelle 360a, 1140 m asl, 47.58278N, 12.95806E, 27.v.-14.vi.2005, leg. F. Eder, [BOLD ID: ES46]. Adult male, (NTNU-VM slide 201773), as previous except for 28.vi-12.vii.2005, leg. F. Eder & A. Schellmoser [BOLD ID: ES82].
Pentaneurella katterjokki differs from other Pentaneurini species by the combination of the following characters. Adult male: thorax with a scutal tubercle, tibial spur on fore leg with long outer tooth and shorter side teeth, anal point apically notched. Adult female: gonapophysis VIII triangular, tergite IX without setae, coxosternapodeme strongly curved, postgenital plate broadly rounded, labia with inconspicuous microtrichia. Pupa: plastron plate moderately large, corona absent, anal macrosetae with adhesive sheaths, genital sac symmetrically tapered. Larva: dorsally DP absent, peg sensilla large, firmly fused with the margin of antennal segment 2, forming a fork-like process.
Adult male (n = 3, except where otherwise stated). Size. Total length 5.2 (1) mm. Wing length 3.0–3.1 mm. Total length/wing length 1.75 (1). Wing length/profemur length 2.09–2.22 (2).
General coloration. Head pale brown with darker occipital margin; pedicel and antenna brown; maxillary palp pale brown. Thorax pale brown. Wing membrane transparent without marks. Legs brown to pale brown. Abdominal tergite I–VI white, T VII with continuous pale brown transverse band near proximal margin, VIII pale brown; hypopygium pale brown.
Head. Temporal setae 17–19, uniserial. Eye ratio 0.47–0.59. Tentorium 235–289 μm long. Clypeus 132–189 μm long, 97–111 μm wide at widest part, bearing 12–22 setae. Cibarial pump 284–301 μm long. Lengths of palpomeres 1–5 (in μm): 77–82; 84–97; 163–178; 171–207; 324–342. Antenna 1250–1297 μm long, diameter of pedicel 185–188 (2) μm. AR 1.22–1.31.
Thorax. Antepronotals 6–10. Acrostichals 30–52, double staggered row which diverges posteriorly to join the dorsocentral row; dorsocentrals 24–38, biserial anteriorly and uniserial posteriorly; prealars 11–12 (2); supraalar 1 (1). Anapleural suture ratio 0.48–0.55. Scutellars 10–14. Scutal tubercle present.
Wing (Fig.
Legs (Fig.
Hypopygium (Fig.
Pentaneurella katterjokki Fittkau & Murray, adult male (A–F), adult female (G–H). A Wing B fore tibial spur C mid tibial spurs D hind tibial spurs E tarsal claw F hypopygium, left: ventral aspect, right: dorsal aspect G female genitalia, dorsal aspect H female genitalia, ventral aspect. Scale bars: 500 µm (A); 100 µm (G, H).
Lengths (in μm) and proportions of leg segments in Pentaneurella katterjokki Fittkau & Murray, male (n = 2 or 3).
fe | ti | ta1 | ta2 | ta3 | |
---|---|---|---|---|---|
p1 | 1385–1487 | 1433–1634 | 616–630 | 551–568 | 385–409 |
p2 | 1246–1503 | 1414–1757 | 935–1110 | 577–693 | 420–510 |
p3 | 1279–1304 | 1434–1519 | 1025–1163 | 684–732 | 471–507 |
ta4 | ta5 | LR | BV | SV | |
p1 | 225–240 | 146–155 | 0.38–0.44 | 2.59–2.78 | 4.63–4.90 |
p2 | 241–284 | 154–182 | 0.63–0.75 | 2.34–2.77 | 2.22–3.12 |
p3 | 268–295 | 167–186 | 0.71–0.77 | 2.28–2.35 | 2.42–2.65 |
Adult female (n = 2, except where otherwise stated). Size. Total length 5.2 (1) mm. Wing length 3.3–3.4 mm. Total length/wing length 1.02–1.14. Wing length/profemur length 2.55–2.74.
General coloration. Head pale brown with darker occipital margin; pedicel and antenna brown; maxillary palp pale brown. Thorax pale brown. Wing membrane transparent without marks. Legs brown to pale brown. Abdominal tergites and genitalia pale brown.
Head. Temporal setae 22–24, irregularly uniserial. Eye ratio 1.12–1.33. Tentorium 186–287 μm long. Clypeus 157–176 μm long, 108–121 μm wide at largest part, bearing 25–28 setae. Cibarial pump 281–305 μm long. Lengths of palpomeres 1–5 (in μm): 52–57; 98–102; 180–181; 188–193; 333–334. Antenna 897–920 μm long, diameter of pedicel 98–100 μm. AR 0.36–0.39.
Thorax. Antepronotals 7. Acrostichals 44–48, double staggered row which diverges posteriorly to join the dorsocentral row of setae; dorsocentrals 36–48, biserial anteriorly and uniserial posteriorly; prealars 15–16; supraalars 2. Anapleural suture ratio 0.49 (1). Scutellars 8–10. Scutal tubercle present.
Wing. Width 1.00–1.10 mm. Costa 3.3–3.4 mm long. VR 0.96–0.98. WW 0.30–0.31. Brachiolum with 3 setae. Squama with 22–25 setae.
Legs. Fore leg: width at apex of tibia 71–74 μm, tibia with single, apical and pectinate spur 27–28 μm long, with 4 lateral teeth; ta1-4 without preapical pseudospurs. Mid leg: width at apex of tibia 54–68 μm, tibia with two apical spurs 27–28; 29–32 μm long, with 4–5 lateral teeth; ta1-4 with preapical pseudospurs. Hind leg: width at apex of tibia 65–67 μm, tibia with two apical spurs 30–35; 44–48 μm long, with 4 lateral teeth; comb not observed; ta1-4 without preapical pseudospurs. Claws slender, distally recurved and pointed and with large basal protuberance. Lengths and proportion of leg segments as in Table
Genitalia (Fig.
Lengths (in μm) and proportions of leg segments in Pentaneurella katterjokki Fittkau & Murray, female (n = 2).
fe | ti | ta1 | ta2 | ta3 | |
---|---|---|---|---|---|
p1 | 1235–1343 | 1539–1625 | 1149–1151 | 745–747 | 508–512 |
p2 | 1349–1414 | 1520–1621 | 822–1165 | 586–729 | 417–515 |
p3 | 1238–1448 | 1596–1879 | 837–1171 | 583–765 | 421–553 |
ta4 | ta5 | LR | BV | SV | |
p1 | 290–306 | 175–196 | 0.71–0.75 | 2.28–2.34 | 2.41–2.58 |
p2 | 232–317 | 165–188 | 0.54–0.72 | 2.36–2.68 | 2.55–3.57 |
p3 | 258–286 | 158–182 | 0.52–0.62 | 2.40–2.74 | 2.66–3.64 |
Pupa (n = 2, except where otherwise stated). Size. Abdomen 3.5–4.3 mm long in male.
General coloration. Exuviae mostly pale brown without any distinctive patterns; thoracic horn brown.
Cephalothorax (Fig.
Abdomen (Fig.
4th instar larva (n = 2, except where otherwise stated). General coloration. Head golden yellow, postoccipital margin brown. Ligula pale yellow, with apex brown. Abdomen pale yellow. Procercus pale brown along anterior margin.
Head (Fig.
Antenna (Fig.
Maxilla (Fig.
Mandible (Fig.
Mentum and M appendage (Fig.
Hypopharyngeal complex (Fig.
Body (Fig.
In their comprehensive analyses of the Chironomidae subfamily Tanypodinae,
The male of Pentaneurella is morphologically similar to Larsia Fittkau, Pentaneura Philippi, Trissopelopia, and Zavrelimyia Fittkau. The bases of the lyrate tibial spurs are similar to the ones of Larsia and Pentaneura, and the absence of setae on tergite IX resembles Trissopelopia (
Regarding the immature stages, the pupa of Pentaneurella shows certain similarities to Krenopelopia Fittkau and Monopelopia Fittkau (
In the Palaearctic, the subfamily Tanypodinae is represented by 29 genera, of which Anatopynia, Johannsen, Telmatopelopia Fittkau and Pentaneurella currently are unique to the region. The latter is a relatively common genus of non-biting midges initially recorded from northern Scandinavia. Currently, the genus has been recorded in Finland (
Little is known about the ecology of Pentaneurella. Immature stages seem to be cold stenothermic rheophiles and krenophiles. Larvae of Pentaneurella have been recorded inhabiting springs and spring-fed streams in Sweden and the Bavarian Alps as well as mountain streams in northern and Central Norway (
Thanks to the team at the Canadian Centre for DNA Barcoding for help with DNA barcode analysis. We would also like to thank Martin Spies, Torbjørn Ekrem, Karstein Hårsaker and Markus Majaneva for supplying us with the material examined in this study. Thanks to Torbjørn Ekrem for comments on the manuscript. F.L. Silva was supported by fellowships from the Coordination for the Improvement of Higher Education Personnel (CAPES - 2014/9239-13-8) and São Paulo Research Foundation (FAPESP - 2016/07039-8). We are indebted to the reviewers for their valuable comments on the manuscript. DNA barcode data in this publication were generated in collaboration with the Norwegian Barcode of Life Network (NorBOL) funded by the Research Council of Norway and the Norwegian Biodiversity Information Centre.