Research Article |
Corresponding author: Elisabeth Stur ( elisabeth.stur@ntnu.no ) Academic editor: Fabio Laurindo Da Silva
© 2015 Elisabeth Stur, Torbjørn Ekrem.
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Citation:
Stur E, Ekrem T (2015) A review of Norwegian Gymnometriocnemus (Diptera, Chironomidae) including the description of two new species and a new name for Gymnometriocnemus volitans (Goetghebuer) sensu Brundin. ZooKeys 508: 127-142. https://doi.org/10.3897/zookeys.508.9874
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Examination of the syntypes of Metriocnemus volitans Goetghebuer, 1940 revealed that these specimens belong to the genus Chaetocladius and are not con-specific with Gymnometriocnemus volitans (Goetghebuer, 1940) sensu
Chironomidae, Orthocladiinae, DNA barcodes, new species, taxonomy, non-biting midges
The orthoclad genus Gymnometriocnemus was suggested by
Metriocnemus volitans was described by
Subgenus Gymnometriocnemus is characterised by a short virga and a weakly developed crista dorsalis in the adult male hypopygium (
DNA barcoding using partial cytochrome c oxidase subunit 1 sequences (COI) (
The motivation for this study was to clarify the identity of Gymnometriocnemus volitans (Goetghebuer), describe hitherto unknown species of Gymnometriocnemus and to present the DNA barcodes of Norwegian Gymnometriocnemus as a resource for future studies of this genus.
We examined eight syntypes of Metriocnemus volitans Goetghebuer from the Royal Belgian Institute of Natural Sciences (RBINS), five male and three female adults mounted between cellophane strips on two separate pins. Both pins bear the label “Env. d. Abisko, Aout 1939, Dr. Thienemann” and “Metriocnemus volitans n sp” (Figs
We also examined the male holotype and a male paratype of Gymnometriocnemus (Raphidocladius) acigus Sæther, 1983 (University Museum of Bergen, Norway (ZMBN)) and two male syntypes and a female syntype of Gymnometriocnemus (Gymnometriocnemus) terrestris (RBINS).
Additional material of Gymnometriocnemus was collected using a variety of methods in different biosurveillance projects: Malaise traps, sweep netting and fogging of oak canopies (Supplementary file 1). This material is deposited in the NTNU University Museum insect collection (NTNU-VM). One to three legs were dissected off the specimens and submitted to the Canadian Centre for DNA Barcoding. Metadata, photos, sequences and trace-files are available in the Barcode of Life Data Systems (BOLD, www.boldsystems.org) through the dataset DS-GYMNO with doi: 10.5883/DS-GYMNO. GenBank accessions are given in Supplementary file 1.
DNA extracts and partial COI gene sequences were generated using standard primers and bi-directional Sanger sequencing with BigDye 3.1 termination at the Canadian Centre for DNA Barcoding in Guelph. Protocols and original trace-files are available through the dataset DS-GYMNO in BOLD. Alignments were done on amino acid sequences and was trivial as indels were absent; only sequences > 300bp were used in the final alignment. The taxon ID-tree was generated using neighbour joining analysis and 1000 bootstrap replicates on Kimura 2-parameter (K2P) genetic distances in MEGA 6 (
Morphological terminology and abbreviations follow Sæther (1980). Antennal and fore leg ratios of Norwegian Gymnometriocnemus are given in Table
Antennal ratios (AR) and fore leg ratios (LR1) of male Gymnometriocnemus from Norway.
Species | AR | LR1 |
---|---|---|
G. (Raphidocladius) kamimegavirgus | 0.88–1.14, 1.04 [n=5] | 0.51–0.52, 0.51 [n=3] |
G. (Raphidocladius) brumalis | 1.03–1.31, 1.19 [n=5] | 0.50–0.53, 0.51 [n=5] |
G. (Raphidocladius) autumnalis | 0.87–1.00, 0.94 [n=4] | 0.56–0.58, 0.57 [n=4] |
G. (Gymnometriocnemus) subnudus | 1.00–1.08, 1.06 [n=3] | 0.58–0.63, 0.61 [n=3] |
G. (Gymnometriocnemus) pallidus | 1.05–1.10, 1.07 [n=5] | 0.62–0.69, 0.65 [n=5] |
G. (Gymnometriocnemus) marionensis | 1.00–1.08, 1.05 [n=4] | 0.60–0.68, 0.65 [n=4] |
Metriocnemus volitans Goetghebuer, 1940: 59.
Metriocnemus volitans Goetghebuer in
Several of the type specimens are quite damaged and many characters are difficult or impossible to observe (Figs
We have only seen two females from eastern and central Norway. The two specimens fit Edwards’ description for G. brevitarsis and represent the first records of this species in Norway. The wing and antenna are photographed (Fig.
Female Gymnometriocnemus. A–C Gymnometriocnemus brevitarsis (CH-OSF33): A Wing B Antenna C genitalia D–F Gymnometriocnemus (G.) pallidus sp. n. (CH-eik47): D Wing E Antenna F genitalia. Specimen codes in parenthesis correspond to codes in BOLD and in figure 6. Scale bar: 200 µm (A, D); 100 µm (B, C, E, F).
Only one specimen of this species from Norway has been available to us. It fits Sæther’s description of G. (G.) marionensis in having very slightly larger megasetae on the gonostyli than specimens of G. (G.) subnudus (Figs
Hypopygia of Norwegian Gymnometriocnemus. A Gymnometriocnemus (G.) marionensis (Finnmark06) B Gymnometriocnemus (G.) pallidus sp. n. (CH-eik131) C Gymnometriocnemus (G.) subnudus (ATNA398) D Gymnometriocnemus (R.) autumnalis sp. n. (Finnmark201) E Gymnometriocnemus (R.) brumalis (Finnmark75) F Gymnometriocnemus (R.) kamimegavirgus (Finnmark76). Scale bar = 50 µm. Specimen codes in parenthesis correspond to codes in BOLD and in figure 6.
Distal part of wing for Norwegian Gymnometriocnemus. A Gymnometriocnemus (G.) marionensis (Finnmark06) B Gymnometriocnemus (G.) pallidus sp. n. (CH-eik131) C Gymnometriocnemus (G.) subnudus (ATNA98) D Gymnometriocnemus (R.) autumnalis sp. n. (Finnmark201) E Gymnometriocnemus (R.) brumalis (CH-OSF70) F Gymnometriocnemus (R.) kamimegavirgus (ATNA269). Scale bar = 200 µm. Specimen codes in parenthesis correspond to codes in BOLD and in figure 6.
Holotype: Male adult (NTNU-VM slide 143840), Norway, Hordaland, Kvam, Berge, oak canopy fogging, tree #3, 60.30921°N; 6.16453°E, 23.vi.2011, leg. Karl H. Thunes [BOLD ID: CH-eik131]. 5 Paratypes: 2 male adults as holotype except tree #1 60.314°N; 6.167°E, 21.vi.2011; 1 female adult as holotype except tree #18, 59.201°N; 9.920°E, 5.vii.2012; 2 male adults, Norway, Telemark, Porsgrunn, Brevik, Frierflauene, 59.0579°N; 9.66485°E, Malaise trap, 30.vi–27.vii.2010, leg. Geir Søli.
The species is named “pallidus”, Latin adjective meaning pale, referring to the conspicuous pale body colour compared to other Holarctic Gymnometriocnemus.
Gymnometriocnemus (G.) pallidus can be separated from other Gymnometriocnemus species by the following combination of characters in the adults: body pale yellow-green; male with short inconspicuous virga, gonostylus with convex outer margin and weakly developed crista dorsalis. Female with setae on most of wing surface, including numerous in cell m; antenna with apical flagellomere pointed and longer than flagellomere 4; genitalia with long rami, about the same length as notum.
Male adult (n = 5 unless otherwise stated). Wing length 1.21–1.30, 1.28 mm. Colouration pale yellow-green body, legs and antennae; slightly darker bands on scutum; postnotum, dorsal side of head, ventral part of preepisternum pale brown; eyes dark brown.
Head. Antennal ratio in Table
Thorax. Antepronotum with 2-6, 3 setae. Dorsocentrals 10-17, 13; acrostichals 7-12, 10, minute and difficult to discern; prealars 3-4; scutellars 6-7.
Wing (Fig.
Legs. Fore tibia with one spur, 35 µm long; mid tibia with two spurs ca. 20-25 µm long; hind tibia with well-developed comb and 2 spurs, ca. 20 and 35 µm long. Fore leg ratios in Table
Hypopygium (Fig.
Female adult (n = 1). Wing length 1.23 mm. Colouration as male.
Head. Antenna (Fig.
Thorax. Antepronotum with 6 setae. Dorsocentrals 18; acrostichals 11; prealars 3; scutellars 6.
Wing (Fig.
Legs. Fore tibia with one spur, 20 µm long; mid tibia lost; hind tibia with well-developed comb and 2 spines, ca. 35–40 µm long. Tarsus of fore leg lost (LR1 not measurable).
Genitalia (Fig.
Immature stages unknown
The species is morphologically similar to G. (G.) subnudus and G. (G.) johanasecundus, but paler (see whole specimen figures in BOLD dataset DS-GYMNO). Males and females are almost completely yellow-green with pale brown posterior side of head and postnotum; pale brown scutal bands and ventral side of preepisternum. Gymnometriocnemus (G.) pallidus is also similar to these species in having a short, triangular anal point and a small virga, but the hypopygium of G. (G.) pallidus has a more prominent inferior volsella than G. (G.) johanasecundus and considerably stronger anal tergite setae than G. (G.) subnudus (Figs
We have seen specimens from eastern and central Norway that fit well with the original and later descriptions of the species, except for having a lower AR (1.0-1.1) compared to what
Holotype: Male adult (NTNU-VM slide no. 136299), Norway, Finnmark, Porsanger, small pond near Gaggavann, 69.8306°N; 25.1856°E, 107 m a.s.l., 03.ix.2010, leg. Alyssa Anderson [BOLD ID: Finnmark201]. 3 Paratypes, male adults: 1 Norway, Finnmark, Vardø, Nedre Domen, lake and pond at road E75, 70.3215°N; 31.0341°E, 120 m a.s.l., 05.ix.2010, leg. Alyssa Anderson; 1 Norway, Finnmark, Nordkapp, Nordkapp-Plateau, 71.1446°N; 25.7641°E, 220 m a.s.l., 01-ix-2010, leg. Trond Andersen; 1 Norway, Oppland, Dovre, Rondane National Park, Vidjedalsbekken (upper), Malaise trap, 61.9717°N; 9.83606°E, 1280 m a.s.l., 15.ix.2008, leg. Terje Hoffstad.
The species is named “autumnalis”, Latin adjective meaning belonging to autumn, referring to the time of the year when the type material was collected.
Gymnometriocnemus (R.) autumnalis can be separated from other Gymnometriocnemus species by the following combination of characters in the adult male: body brown, dark brown; virga long and conspicuous with strong lateral sclerotization, anal tergite without dorsal anal point or ridge, gonostylus with convex outer margin and well-defined median crista dorsalis.
Male adult (n = 4 unless otherwise stated). Wing length 1.30–1.52, 1.43 mm. Colouration completely brown, dark brown except for pale transverse bands posteriorly on abdominal tergites V–VIII, narrower on tergite V.
Head. Antennal ratio in Table
Thorax. Antepronotum with 2 setae. Dorsocentrals 10-11; acrostichals 8-9, minute and difficult to discern; prealars 4-6; scutellars 2-5.
Wing (Fig.
Legs. Fore tibia with one spur, 40 µm long; mid tibia with two spurs ca. 20 µm long; hind tibia with well-developed comb and 2 spines, ca. 50 µm long. Fore leg ratios in Table
Hypopygium (Fig.
Female and immature stages unknown.
The species is morphologically similar to G. (R.) brumalis and G. (R.) kamimegavirgus, but different in lacking an anal point or ridge on the anal tergite. This character is similar to characters reported for G. terrestris and G. tairaprimus, but these two species can according to original descriptions be separated by having a higher AR (1.4 in G. terrestris) and a different shape of the superior volsella (
We have barcoded specimens from eastern, central and northern Norway that fall within the same genetic cluster although with quite large intraspecific divergence (0–6.8%, mean 3.42% K2P-distance) (Fig.
Gymnometriocnemus kamimegavirgus Sasa & Hirabayashi (
Gymnometriocnemus volitans (Goetghebuer), misidentifications (e.g.
Gymnometriocnemus (Raphidocladius ?) volitans (Goetghebuer) sensu
Gymnometriocnemus (Raphidocladius) volitans (Goetghebuer) sensu
Gymnometriocnemus (R.) kamimegavirgus can be separated from other species of the genus Gymnometriocnemus by having well-developed, long virga (about the length of the gonocoxite); AR 0.9-1.1 (n=5); LR1 about 0.53-0.56 (n=3); wing membrane with setae at the apex only, occasionally with 1-2 setae proximally in cell an; R2+3 situated in the middle between R1 and R4+5; dark brown almost blackish thorax and head, slightly paler abdomen and legs.
Our examined material is from eastern, central and northern Norway, frequently collected near streams, rivers and moors. Male adults fit well with Brundin’s description of G. volitans, and Sasa & Hirabayashi’s description of G. kamimegavirgus except for slightly fewer setae on the abdominal tergites (
The species Gymnometriocnemus brevitarsis is only known as female and therefore not included in the key.
1 | Large, needle-like virga well-developed, sometimes with strong lateral sclerotization (Fig. |
2 |
– | Virga small and inconspicuous, without lateral sclerotization (Fig. |
5 |
2 | Anal tergite without ridge or anal point (Fig. |
3 |
– | Anal tergite with at least a median triangular ridge (Fig. |
4 |
3 | Inferior volsella with obvious dorsal and ventral lobe (Japan) | G. (R.) tairaprimus |
– | Inferior volsella with single lobe (Norway) (Fig. |
G. (R.) autumnalis |
4 | Wing membrane with setae on wing tip only; often strong sclerotization laterally of virga (Fig. |
G. (R.) kamimegavirgus |
– | Wing membrane with setae on at least half of wing; no strong sclerotization laterally of virga (Fig. |
G. (R.) brumalis |
6 | Body pale yellow-green with pale brown scutal markings | G. (G.) pallidus |
– | Body completely brown, or when yellowish ground colour with dark brown scutal markings | 7 |
7 | Edge of anal tergite broadly rounded (possibly an anal point, but difficult to discern in syntypes); gonostylus strongly curved inwards | G. (G.) terrestris |
– | Anal point present, triangular; gonostylus at most with a weakly convex outer margin (Fig. |
8 |
8 | Ground colour of thorax yellowish (Japan) | G. (G.) johanasecundus |
– | Ground colour of thorax brown | 9 |
9 | Anal point moderately well developed, c. 38 µm long (Fig. |
G. (G.) marionensis |
- | Anal point weakly developed, c. 17 µm long (Fig. |
G. (G.) subnudus |
As a result of this study, there are now 17 species of Gymnometriocnemus registered worldwide and the genus is present in all major biogeographical regions except Antarctica. Our findings through moderate sampling in Norway indicate that the number of species could be considerably higher also on a global scale and show that molecular data can be a great advantage in diversity assessments of targeted groups. Moreover, our study also highlights the importance of consulting type material for correct identification of Chironomidae if we are to avoid long term misconceptions of species.
Thanks to the team at the Canadian Centre for DNA Barcoding for help with DNA barcode analysis and to the Norwegian Biodiversity Information Centre for funding biosurveillance projects through the Norwegian Taxonomy Initiative from which most of the material used in this study originates. DNA barcode data in this publication was generated in collaboration with the Norwegian Barcode of Life Network (NorBOL) funded by the Research Council of Norway and the Norwegian Biodiversity Information Centre. Thanks also to Pol Limbourg and Wouter Dekoninck at RBINS for the loan of the Goetghebuer types and to Trond Andersen and Steffen Roth at the University Museum Bergen for loan of specimens of G. (G.) marionensis and types of G. (R.) acigus. We are grateful for the help provided by Martin Spies in obtaining copies of relevant literature from the library at the Zoologische Staatssammlung München. Thanks to Rick Jacobsen and two anonymous reviewers for comments on the submitted manuscript.
Examined and DNA barcoded Gymnometriocnemus species from Norway
Data type: Excel spreadsheet with specimen data and GenBank accessions
Explanation note: The spreadsheet contains the following information for all examined species: sample id, catalogue number, voucher status, locality, date of collection, collector and GenBank accession.