Research Article
Print
Research Article
Two new species of Macropelopia (Diptera, Chironomidae) from Oriental China, delineated with morphology and COI sequences
expand article infoXiangliang Fang‡, Zigang Xu§‡, Yuanyuan Yao|, Yue Fu‡
‡ Huanggang Normal University, Huanggang City, China
§ Yangtze University, Hubei, China
| Tianjin Natural History Museum, Tianjin City, China

Corresponding author: Yue Fu ( fuyue20190125@163.com )

Academic editor: Fabio Laurindo da Silva
© 2024 Xiangliang Fang, Zigang Xu, Yuanyuan Yao, Yue Fu.
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: Fang X, Xu Z, Yao Y, Fu Y (2024) Two new species of Macropelopia (Diptera, Chironomidae) from Oriental China, delineated with morphology and COI sequences. ZooKeys 1210: 287-298. https://doi.org/10.3897/zookeys.1210.127183
ZooBank: urn:lsid:zoobank.org:pub:1D0A3DB9-3CD4-4DD5-97DA-471BCF3FAD8A
Open Access

Abstract

Two new species, Macropelopia (Macropelopia) excavata Xu & Fu, sp. nov. and Macropelopia (Macropelopia) quadrimacula Xu & Fu, sp. nov., are described as male adults. A key to identify the males of Macropelopia from China is provided. Furthermore, in order to ascertain the genetic distance between these species and their morphological characteristics, mitochondrial cytochrome c oxidase subunit I gene sequences were uploaded to the National Center for Biotechnology Information. These COI sequences were then utilized to infer the relationships between the species, employing the neighbor-joining method.

Key words

DNA barcoding, Macropelopia, morphology, non-biting midge, taxonomy

Introduction

Thienemann and Kieffer (1916) established the genus Macropelopia, with Isoplastus bimaculatus Kieffer as the type species. Macropelopia is classified in the tribe Macropelopiini and is further divided into two subgenera: Bethbilbeckia Fittkau et Murray and Macropelopia s. str. (Cranston and Epler 2013). Previous studies described a total of 18 species within the subgenus Macropelopia s. str. and three species in the subgenus Bethbilbeckia. The main difference between the two subgenera in the adult stage is that Macropelopia s. str. has a tibial comb, while Bethbilbeckia does not. Among these, 11 species of the subgenus Macropelopia are found in the Palearctic region (Kieffer 1912, 1916; Tokunaga 1937; Fittkau 1962; Lencioni and Marziali 2005), one in the New North region (Roback 1971), two in the Oriental region (Tang and Niitsuma 2020; Hazra and Chaudhuri 2001), three in the Neotropical region (Andersen 2018; Silva and Pinho 2020; Dantas et al. 2023), and one in the African region (Freeman 1955).

According to Wang (2000), only one species of Macropelopia, M. nebulosa (Meigen), has been described based on adult, and four other species have been recorded based on larvae. Wang et al. (2011) recorded three additional species of Macropelopia, namely M. galbina Wang, Cheng & Wang, and M. grandivolsella Wang, Cheng & Wang which were both found in Hubei Province, and M. rotunda Wang, Cheng & Wang, which was discovered in Fujian Province. However, recent examinations conducted by Tang and Niitsuma (2020) have resulted in significant taxonomic revisions. The species M. grandivolsella has been synonymized with Macropelopia paranebulosa Fittkau, while Macropelopia rotunda is now referred to as M. kibunensis (Tokunaga), with these two species considered synonymous. Additionally, M. galbina was transferred to the genus Brundiniella. Furthermore, Tang and Niitsuma (2020) have described a previously unknown species, Macropelopia (Macropelopia) pergrandis, originating from Yunnan Province.

In this study, we report the discovery of two new species within the subgenus Macropelopia s. str., sourced from the remarkable Dabie Mountain National Nature Reserve situated in Hubei Province. The discovery holds significant importance as it contributes to the diversity of the genus. In addition, we have also assembled a key for distinguishing and identifying the known adult males of the subgenus in China. Moreover, we have conducted an analysis utilizing the mitochondrial cytochrome c oxidase subunit I (COI) gene to infer genetic distance and determine the differences between the species within the genus Macropelopia,further enhancing our understanding of their morphological characteristics.

Material and methods

Specimens were collected using the light trap induction method and preserved in 85% alcohol. Subsequently, they were sent to the laboratory for preliminary species identification and assigned individual numbers under a microscope. Images of slide mounts were obtained using a Nexcope NE930 compound microscope equipped with Capture 2.1 software. Genomic DNA was extracted from the thorax and legs of the specimens using the Qiagen DNA Blood & Tissue Kit. PCR amplification of the COI gene was performed following the primers and temperature regimes described by Folmer et al. (1994). After DNA extraction, the transparent exoskeleton was rinsed with 96% ethanol and mounted in Euparal on microscope slides, along with the corresponding antennae, head, wings, and legs, following the protocol outlined by Sæther (1969). Morphological nomenclature adheres to Sæther (1980), and measurements include the minimum, maximum, and average values for at least three specimens. All specimens are currently housed at the College of Biology and Agricultural Resources, Huanggang Normal University, China. Evolutionary analyses were conducted using MEGA 11.

The main abbreviations and corresponding English terms used in this study (with the value of ‘N’ representing the number of measured specimens mentioned in the text) are as follows: TL, total length; WL, total wing length; Pfe, length of the forefoot; VR, Venarum ratio = length of Cu / length of M; Cu1 cubitus 1; AR, Antennal ratio = length of ultimate flagellomere / combined lengths of flagellomeres one to penultimate; Fe, femur; ta1–ta5, tarsomeres ta1–ta5; LR, ta(basal segment) / ti; BV, Fe + ti +ta1 / ta2 + ta3 + ta4 + ta5; SV, Fe + ti / ta1; R, Radius; R1, Radius 1; R4+5, Radius four and five; BR, largest bristle/width of ta1 about 1/3 from distal end; HR, length of gonocoxite / length of gonostylus; HV, total length (TL) / length of gonostylus × 10.

Taxonomy

Macropelopia Thienemann

Macropelopia Thienemann in Thienemann & Kieffer, 1916: 497. Fittkau 1962: 102; Roback 1971: 87, 1978: 159; Fittkau and Murray 1986: 50; Murray and Fittkau 1989: 61; Epler 2001: 4.53; Niitsuma et al. 2004: 44; Cranston and Epler 2013: 53; Silva and Pinho 2020: 575; Tang and Niitsuma 2020.

Bethbilbeckia Fittkau et Murray, 1988: 253; Murray and Fittkau 1989: 46; Epler 2001: 4.29.

Type species

Isoplastus bimaculatus Kieffer [=Tanypus nebulosus Meigen] by original designation.

Macropelopia excavata Xu & Fu, sp. nov.

https://zoobank.org/2AA64D33-4F08-444C-AC77-FA080DBFBCAF
Fig. 1A–K

Type material

Holotype male (HNU: Cdbs60), China: Hubei Province, Huanggang City, Yingshan County, Dabie Mountain, Longtan Gorge, light trap, 31.0867°N, 115.8138°E, 486.71 m a. s. l., 7. IX. 2022, Zigang Xu.

Etymology

The new species is named “excavata” derived from the Latin term “excavatus”, meaning concave, which aptly describes the inward concave shape of tergite IX’s posterior edge.

Diagnostic characters

The distinguishing characteristics of this new species are the presence of two prominent longitudinal thick spots positioned in the middle of tergites II to IV, and the wing with brown markings on the distal end of Cu1 and basal part of cell an. Additionally, the tergites from V to IX display a distinctive brown hue, adding to their identification. The posterior edge of tergite IX is concave in shape, and the anal point is absent. The gonostylus is prominently curved at a right angle.

Description

Adult male (N = 1)

Total length 4.69 mm, thorax length 1.45 mm, wing length 2.66 mm, TL/WL 1.76, WL/Pfe 2.19.

Coloration (Fig. 1D). The head and thorax are uniformly dark brown. The femur of the legs is also dark brown, while the remaining parts of the legs are yellow. The wings exhibit two significant gray spots positioned near the Cu1 and An veins. There is a longitudinal color spot present in the middle of tergites II to IV, the tergites V to IX and hypopygium are all brown in coloration.

Figure 1. 

Macropelopia (Macropelopia) excavata Xu & Fu, sp. nov., male imago A head B antenna C wing D whole body of male adult E spotted shape of the tergites F thorax G fore tibial apex H hypopygium, dorsal view I photo of hypopygium, dorsal view J hypopygium, ventral view K gonostylus.

Head (Fig. 1A). Antenna (Fig. 1B) with 13 flagellomeres, with an antennal ratio (AR) measuring 2.24. The flagellomeres taper towards their ends and ultimate flagellomere with a single apical seta. Temporal setae 24, comprising 8 inner verticals, 12 outer verticals, and 16 postorbitals. Tentorial length 234 µm, width 43 µm. Stipes length 192 µm, width 82 µm. Clypeus with 16 setae. Length of palpomeres (in µm): 69; 100; 161; 210; 342. Length ratio of palpomeres 5/3: 2.13.

Thorax (Fig. 1F). Antepronotals with 9 setae, the acrostichals and dorsocentrals arranged in multiple columns, acrostichals with 41 setae, dorsocentrals with 62 setae. Postnotals with 6 setae, humerals with 8 setae, prealars with 32 setae. Scutal tubercle present and obvious, scutellars with 32 setae.

Wing (Fig. 1C). The wing membrane area hairy, distinctive gray spots present near Cu1 and An veins, the arculus hook-like shape, and the anal lobe mainly developed. VR: 0.91, brachiolum with 3 setae, veins with many setae, squama with 34 setae. Costal extension 128 µm.

Legs. The fore tibia possesses a single spur measuring 85 µm in length and features 15 side teeth. The width of the fore tibia at its apex is 84 µm, and the fore tibial comb consists of 5 setae (Fig. 1G). Two spurs of mid tibia are observed, measuring 87 µm and 51 µm long, bearing 16 and 11 lateral teeth, respectively, and has a width at the apex of 73 µm. The hind tibia exhibits two spurs, measuring 75 µm and 45 µm, with 18 and 12 lateral teeth, respectively. The hind tibial comb composed of 10 setae, with the longest seta 73 µm and the shortest 45 µm. The width of the hind tibia at its apex is recorded as 80 µm. The lengths and proportions of each leg are shown in Table 1.

Table 1.
Download as
CSV
XLSX

Lengths (in µm) and proportions of leg segments of male M. excavata Xu & Fu, sp. nov. (N = 1).

fe ti ta1 ta2 ta3 ta4
P1 1215 1502 1168 600 428 293
P2 1295 1461 886 428 317 222
P3 1193 1654 1144 592 414 267
ta5 LR BV SV BR
P1 208 0.78 2.54 2.32 3.96
P2 164 0.61 3.23 3.11 4.45
P3 184 0.69 2.74 2.49 4.56

Hypopygium (Fig. 1G, H). The anal point absent. The phallapodeme, although short, measures at a clearly discernible length of 69 µm. The gonocoxite showcases a cylindrical shape and spans 206 µm in length. The gonostylus (Fig. 1K) is 125 µm long and exhibits a curved inward shape at approximately two-thirds of its length. It displays protrusions on both the inside and outside, gradually narrowing towards the tip. The megaseta is 16 µm long. The inferior volsella unconspicuous, along the inside of gonocoxite, and contains concentrated long inner microhairs. HR: 1.64, HV: 3.74.

Remarks

This new species is similar to M. kibunensis (Tokunaga) because their anal point is absent and the inferior volella is undeveloped, but can be distinguished by AR 2.24, wing with brown markings on distal end of Cu1 and basal part of cell an, otherwise unmarked, and a concave rear edge line of tergite IX, while AR 1.7–1.9, wing with brown markings on distal end of Cu1, M3+4 and M1+2. The pairwise distance based on the COI sequence of M. kibunensis and M. excavata sp. nov. is 0.105–0.107, further distinguishing them from each other.

Distribution

Hubei Province, Oriental China.

Macropelopia quadrimacula Xu & Fu, sp. nov.

https://zoobank.org/13C57663-77BF-4852-988D-F05C38B51B33
Fig. 2A–K

Type material

Holotype , male (HNU: Cdbs7602), China: Hubei Province, Huanggang City, Yingshan County, Dabie Mountain, Longtan Gorge, 31.0867°N, 115.8138°E, 486.71 m a. s. l., 8. VI. 2022, light trap, leg. Zigang Xu. Paratypes: 3 males (HNU: Cdbs7601, Cdbs7603, Cdbs7604), same as holotype; 3 males (HNU: Cdbs8901, Cdbs8902, Cdbs8903), Hubei Province, Huanggang City, Yingshan County, Dabie Mountain, Wujiashan National Forest Park, 31.1047°N, 115.7913°E, 931.91 m a. s. l., 9. VI. 2022, light trap, leg. Zigang Xu.

Etymology

The name of this new species is derived from the Latin words “quartri” and “macula”, meaning “four” and “spot”, “stain” or “mark”, respectively. The name specifically pertains to the presence of four distinctive black spots found on the tergites of this species.

Diagnostic characters

This species has two short longitudinal striped spots on each side of tergites II to V, as well as two elliptical spots in the center. Wing with brown markings on apical of Cu1, M3+4 and basal part of cell an. Additionally, tergite IX with a triangular anal point beyond the margin of tergite IX. Lastly, the gonostylus is curved inward at two-thirds of its length, and the apex is markedly tapered.

Description

Adult males (N = 7)

Total length: 4.49–5.55, 5.02 mm, Wing length 2.72–3.27, 2.99 mm, TL/WL 1.65–1.76, 1.71, WL/Pfe 2.08–2.59, 2.34.

Coloration (Fig. 2D). The head and thorax of this species are uniformly brown. The femurs of all legs are also brown, while the other sections display a yellow coloration. The wings are adorned with various color spots. Notably, there is a longitudinal color band on both sides of tergites II to V, accompanied by two elliptical spots at the center. Tergites VI to VII exhibit a dark brown hue, while tergites VIII to IX and the hypopygium are brown in color.

Figure 2. 

Macropelopia (Macropelopia) quadrimacula Xu & Fu, sp. nov., male imago A head B antenna C wing D whole body of male adult E spotted shape of the tergites F thorax G fore tibial apex H hypopygium, dorsal view I photo of hypopygium, ventral view J hypopygium, ventral view K gonostylus.

Head (Fig. 2A). Antenna with 13 flagellomeres (Fig. 2B), AR: 2.01–2.05, 2.03, The end of flagellomere narrow, with approximately one-third of the end easily foldable, and the ultimate flagellomere possessing a single apical seta. Temporal setae 20–22, 21. The inner verticals and outer verticals are arranged in two or three columns, with 4–8, 6 inner verticals, 10–16, 13 outer verticals, and 12–18, 15 postorbitals. Tentorial length 250–258, 254 µm, width 36–43, 40 µm. Stipes length 271–303, 287 µm, width 62–95, 78 µm. Clypeus with 16–18, 17 setae. Length of palpomeres (in µm): 67–78, 72; 91–105, 98; 173–190, 182; 241–279, 260; 334–378, 356. Length ratio of palpomeres 5/3: 1.93–1.99, 1.96.

Thorax (Fig. 2F). Thorax length 1.36–1.73, 1.55 mm, antepronotals with 12–12, 12 setae. The acrostichals and dorsocentrals are arranged in multiple columns, acrostichals with 40–60, 50 setae, dorsocentrals with 60–80, 70 setae. Anepisternals with 2–3, 3 setae, postnotals with 4–6, 5 setae, humerals with 8–14, 11 setae, prealars with 22–26, 24 setae. Scutal tubercle was present and obvious, scutellars with 32–40, 36 setae.

Wing (Fig. 2C). The wing membrane area is hairy, Cu1 and An with obvious gray spots around the veins, arculus hook-shaped, and anal lobes particularly developed. VR: 0.91–0.96, 0.94, brachiolum with 3 setae, R1 with 30–40, 35 setae, R with 40–60, 50 setae, R4+5 with 50–60, 55 setae, squama with 54–60, 57 setae. Costal extension 157–160, 158 µm.

Legs. The fore tibia possesses a single spur measuring 85–112, 98 µm in length and features 19–20, 20 side teeth. The width of the fore tibia at its apex is 79–86, 82 µm, and fore tibial comb consists of 5–6 setae (Fig. 2G). Two spurs on mid tibia are observed, measuring 87–95, 91 µm and 50–56, 53 µm long, bearing 18–20, 19 and 11–16, 14 lateral teeth, respectively, and has a width at the apex of 75–81, 78 µm. The hind tibia exhibits two spurs, measuring 88–96, 92 µm and 49–53, 51 µm, with 19–21, 20 and 15–17, 16 lateral teeth, respectively. The hind tibial comb is composed of 13–15,14 setae, with the longest comb seta 73–81, 77 µm and the shortest 40 –46, 43 µm. The width of the hind tibia at its apex is recorded as 76–86, 81 µm. The lengths and proportions of each leg are shown in Table 2.

Table 2.
Download as
CSV
XLSX

Lengths (in µm) and proportions of leg segments of male M. quadrimacula Xu & Fu, sp. nov. (N = 6).

fe ti ta1 ta2 ta3 ta4
P1 1260–1321, 1291 1606–1803, 1705 1195–1346, 1271 628–712, 670 455–517, 486 300–342, 321
P2 1342–1526, 1434 1546–1732, 1639 944–1086, 1015 474–533, 504 343–394, 368 229–257, 243
P3 1285–1338, 1311 1795–1935, 1865 1167–1355, 1261 637–723, 680 455–498, 477 285–319, 302
ta5 LR BV SV BR
P1 192–215, 203 0.73–0.75, 0.74 2.47–2.61, 2.54 2.28–2.46, 2.37 2.57–3.84, 3.21
P2 165–184, 175 0.61–0.63, 0.62 3.16–3.21, 3.19 3.01–3.11, 3.06 2.56–4.37, 3.47
P3 176–204, 190 0.63–0.71, 0.67 2.65–2.83, 2.74 2.42–2.67, 2.55 2.84–3.47, 3.16

Hypopygium (Fig. 2H–K). The anal point is small, forming an obtuse triangle. Phallapodeme short and obvious, 72–81, 77 µm long. Gonocoxite cylindrical shape, 253–265, 259 µm long. Gonostylus (Fig. 2K) 126–146, 136 µm long, curved inward at two-thirds of its length and possessing protrusions both on the inner and outer sides, the inner protrusion was located at a quarter of the apex of the gonostylus, while the outer protrusion was located halfway along the apex. Tergite IX with 16–20, 18 setae, megaseta 16–17 µm long. Inferior volsella small and protuberant. HR: 1.91–2.09, 2.00. HV: 3.56–3.94, 3.75.

Remarks

This new species can be identified by the presence of two short longitudinal color bands on the sides of tergites II to V, along with two elliptical spots in the middle. These distinctive characteristics set it apart from other species within the genus. However, the abdominal spots of this species may sometimes be indistinct and appear blurry. In tergites III to V, these spots may be partially obscured by brown spots, but tergite II consistently displays four clearly visible elliptical spots. The overall shape of this new species is similar to that of M. kibunensis, and it shares the same gonostylus morphology. However, M. kibunensis lacks an inferior volsella and anal point, while this new species possesses a protrusion on the inferior volsella. The shape of the inferior volsella is comparable to that of M. excavata sp. nov., but this new species can still be differentiated by the presence of color spots on the tergites, the presence of the anal point, and a higher HR value (1.91–2.09) compared to M. excavata sp. nov. Based on COI sequences, the pairwise distances between M. quadrimacula and M. kibunensis, and between M. quadrimacula and M. excavata, are 0.119–0.125 and 0.131, respectively, further setting it apart from them.

Distribution

Hubei Province, Oriental China.

Key to adult males of genus Macropelopia from China

1 Wings with distinct color spots 2
– Wings without color spots; only dark markings on cross-vein r-m M. (M.) notata (Meigen)
2 Tergites with brown spots 3
– Tergites without brown spots 6
3 Foretibial comb with 9–15 small bristles, inferior volsella significantly larger M. (M.) paranebulosa (Fittkau)
– Foretibial comb with 5–7 small bristles, inferior volsella absent or undeveloped 4
4 Tergites IX is uncovered gonocoxite, base of the anal point is wide, presenting a triangular anal point; posterior setae on tergite IX are mainly distributed in the central part M. (M.) quadrimacula Xu & Fu, sp. nov.
– Tergites IX covered parts of gonocoxite, anal point absent; posterior setae on tergite IX are mainly distributed at the posterior margin 5
5 Coxa, trochanter, part femur and apex of tibia dark brown, and apex of femur pale; antennal ratio 1.7–1.9; whole wing with dense setae on membrane M. (M.) kibunensis (Tokunaga)
– Coxa, trochanter, femur and apex of tibia dark brown; antennal ratio 2.24; upper part of the wings with dense setae on membrane, but they are significantly reduced below the CU vein and M2+3 vein M. (M.) excavata Xu & Fu, sp. nov.
6 Tergites pale brown, foretibial comb with 11–15 small bristles, and anal point absent M. (M.) pergrandis (Tang & Niitsuma)
– Tergites all brown, foretibial comb with 6 bristles and anal point present M. (M.) decedens (Walker)

Discussion

Based on the statistical data presented in this study, it has been revealed that there are currently seven species of the genus Macropelopia known to be distributed in China. However, it is important to note that the species Macropelopia (M.) notata and M. (M.) decedens, as reported by Wang et al. (2020), lack sufficient morphological characteristics for definitive identification. While we have included these species in the key provided in this study to reflect the current research records, further investigations are necessary to verify their distribution with more certainty.

We successfully obtained eight COI sequences for two new species and downloaded an additional eight sequences for seven species from the National Center for Biotechnology Information (NCBI). Their taxonomic names and GenBank accession numbers can be found in Fig. 3. By utilizing the neighbor-joining method (Tamura et al. 2021) for constructing a phylogenetic tree, our analysis revealed that M. excavata sp. nov. and M. kibunensis are closely related, as they appeared on the same branch of the tree. This finding is consistent with their shared morphological characteristics, such as the similar coloration of the tergites, anal point absent, and thorax features. Macropelopia quadrimacula sp. nov. is shown to be the sister group to (M. excavata sp. nov. + M. kibunensis); the primary distinguishing features between M. quadrimacula sp. nov. and M. excavata sp. nov. lie in their hypopygium and the presence or absence of a dorsal stripe on their tergites. These results exemplify a strong congruence between the molecular and morphological data. Interestingly, our findings contrast with the description of M. kibunensis, which includes yellow femora and the wing with dense setae. In contrast, the two new species possess brown femora and a dorsal stripe on the tergites. Thus, within the genus Macropelopia, key criteria for morphological classification encompass the characteristics of the hypopygium and dorsal stripe patterns on the tergites, followed by the markings and macrotrichia of the wing, and the color and features of the legs.

Figure 3. 

Neighbor-joining tree based on cytochrome c oxidase subunit I (COI) of sixteen Macropelopia specimens. Numbers on branches refer to the evolutionary distances computed using the Kimura 2–parameter method (Kimura 1980) and represent number of base substitutions per site. Taxa names include scientific names and GenBank accession numbers of corresponding COI gene fragments.

Chironomids offer the advantage of having three distinct stages (larvae, pupae, and adults); the morphology of the larvae and pupae of Macropelopia also plays an important role in species delineation (Fittkau 1962, Roback 1978, Fittkau and Murray 1986, Tang and Niitsuma 2020). This study primarily focuses on morphological differences in the adult stage; collecting specimens from multiple life stages simultaneously remains a challenge. However,previous studies have demonstrated that COI is suitable for summarizing sequence diversity and detecting taxonomically challenging species within Macropelopia (Silva and Pinho 2020). Therefore, we anticipate further studies on species delineation using the COI gene segment to enhance the reliability of new species establishment. The analysis of partial DNA barcode sequences supports Macropelopia excavata sp. nov. and Macropelopia quadrimacula sp. nov. as valid species.

Acknowledgments

The project was supported by the National Natural Science Foundation of China (NSFC) (Grant No. 32070483, 31460572), and the National Science and Technology Fundamental Resources Investigation Program of China (Grant No. 2019FY101800). We thank the reviewers and editors for their valuable comments on the revision of the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

No funding was reported.

Author contributions

Conceptualization: YF. Data curation: ZX. Formal analysis: XF. Funding acquisition: YF. Investigation: YY, XF. Methodology: ZX. Supervision: YF. Visualization: ZX. Writing - original draft: XF. Writing - review and editing: YF.

Author ORCIDs

Yue Fu https://orcid.org/0000-0002-4045-4352

Data availability

All of the data that support the findings of this study are available in the main text.

References

  • Andersen T (2018) Macropelopia (Bethbilbeckia) chilensis n. sp. (Diptera, Chironomidae) from Cajón del Maipo, Chile. Norwegian Journal of Entomology 65: 85–90.
  • Cranston PS, Epler JH (2013) 5. The larvae of Tanypodinae (Diptera: Chironomidae) of the Holarctic Region—Keys and diagnoses. In: Andersen T, Cranston PS, Epler JH (Sci. Eds) The larvae of Chironomidae (Diptera) of the Holarctic Region—Keys and diagnoses. Insect Systematics & Evolution 66 (Supplement), 39–136.
  • Epler JH (2001) Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. Special Publication SJ2001–SP13. Department of Environmental and Natural Resources, Division of Water quality, Raleigh and St. Johns River Water Management District, Palatka, 526 pp.
  • Fittkau EJ (1962) Die Tanypodinae (Diptera: Chironomidae) (Die Tribus Anatopyniini, Macropelopiini and Pentaneurini). Abhandlungen zur Larven Systematik der insekten 6: 1–453.
  • Fittkau EJ, Murray DA (1986) 5. The pupae of Tanypodinae (Diptera: Chironomidae) of the Holarctic region—Keys and diagnoses. Entomologica scandinavica 28 (Supplement): 31–113.
  • Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.
  • Freeman P (1955) A study of the Chironomidae (Diptera) of Africa south of the Sahara. Part I Bulletin of the British Museum of Natural History. Entomology 4: 1–67.
  • Hazra N, Chaudhuri PK (2001) Two species of Macropelopia Thienemann (Diptera: Chironomidae), first report from Darjeeling-Sikkim Himalayas of India. Aquatic insects 23(4): 297–309. https://doi.org/10.1076/aqin.23.4.297.4880
  • Kieffer JJ (1916) Tendipedides (Chironomides) de Formose conserves au Museum National Hongrois de Budapest et determines par J. J. Kieffer. Annales Historico. Naturales Musei Nationalis Hungarici 14: 81–121.
  • Kimura M (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111–120. https://doi.org/10.1007/BF01731581
  • Lencioni V, Marziali L (2005) A new species of Macropelopia Thienemann (Diptera, Chironomidae) from the Italian Alps. Italian Journal of Zoology 72(4): 317–320. https://doi.org/10.1080/11250000509356692
  • Murray DA, Fittkau EJ (1989) The adult males of Tanypodinae (Diptera: Chironomidae) of the Holarctic Region—Keys and diagnoses. In: Chironomidae of the Holarctic Region. Keys and diagnoses. Part 3. Adult males (T. Wiederholm, ed.). Entomologica Scandinavica 34 (Supplement): 37–123.
  • Roback SS (1971) The adults of the subfamily Tanypodinae (= Pelopinae) in North America Diptera: Chironomidae). Monograph of the Academy of Natural Sciences of Philadelphia 17: 1–410.
  • Roback SS (1978) The immature chironomids of the eastern United States: III. Tanypodinae—Anatopyniini, Macropelopiini and Natarsiini. Proceedings of the Academy of Natural Sciences 129: 151–202.
  • Sæther OA (1969) Some Nearctic Podonominae, Diamesinae and Orthocladinae (Diptera: Chironomidae). Bulletin of the Fisheries Research Board of Canada 170: 1–154.
  • Sæther OA (1980) Glossary of chironomid morphology terminology (Diptera: Chironomidae). Entomologica scandinavica Supplement 14: 1–51.
  • Tang HQ, Niitsuma H (2020) Revision of the Chinese Macropelopia (Diptera: Chironomidae: Tanypodinae), with description of a new species. Zootaxa 4834(2): 207–218. https://doi.org/10.11646/zootaxa.4834.2.3
  • Thienemann A, Kieffer JJ (1916) Schwedische Chironomiden. Archiv für Hydrobiologie Planktonkunde Supplement 2: 483–554.
  • Tokunaga M (1937) Chironomidae from Japan (Diptera), IX. Tanypodinae and Diamesinae. Philippine Journal Science 62: 21–65.
  • Wang XH (2000) A revised checklist of Chironomidae from China (Diptera), In: Hffrichter O (Ed.) Late 20th century research on Chironomidae. An anthology from 13th international symposium on Chironomidae. Shaker Verlag, Aachen, 629–652.
  • Wang Q, Cheng M, Wang XH (2011) Three new species of Macropelopia Thienemann from China (Diptera: Chironomidae: Tanypodinae). African Journal of Agricultural Research 6(17): 4122–4130.
  • Wang XH, Liu WB, Lin XL, Song C, Sun BJ, Yan CC, Qi X (2020) Chironomidae. In Yang D, Li Z, Liu QF. Species Catalogue of China, Volume 2 Animals, Insecta (V), Diptera (1), Nematocera. Science Press, Beijing, 221–289.
login to comment