Research Article |
Corresponding author: Jian-Jun Gao ( gao-leyun@263.net ) Academic editor: Rudolf Meier
© 2020 Run-Jie Jiao, Li-Hong Bai, Jian-Jun Gao.
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:
Jiao R-J, Bai L-H, Gao J-J (2020) Descriptions of two new species of the genus Colocasiomyia (Diptera, Drosophilidae) breeding on Rhaphidophora host plants in Yunnan, China. ZooKeys 968: 127-141. https://doi.org/10.3897/zookeys.968.56677
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The genus Colocasiomyia de Meijere (Diptera, Drosophilidae) is known to include 30 described and nearly 60 undescribed species classified into six species groups. Among these, the C. gigantea group of seven known species (two Southeast Asian and five Chinese) proved to be peculiar for its specificity on monsteroid (subfamily Monsteroideae, family Araceae) host plants. In this paper, two new species, C. todai Jiao & Gao, sp. nov. and C. liae Jiao & Gao, sp. nov., are described as members of the C. gigantea group with specimens collected from inflorescences of the monsteroid host species Rhaphidophora peepla (Roxb.) Schott and R. crassicaulis Engl. & Krause, respectively, in Yunnan, China. The two new species are delimitated, in comparison with all known species, based on not only morphological but also DNA barcode (partial sequence of the mitochondrial COI, i.e., cytochrome c oxydase subunit I, gene) data. A revised key to all the nine species of the C. gigantea species group is provided.
DNA barcoding, key to species, morphology, neighbor-joining tree, taxonomy
The genus Colocasiomyia de Meijere, 1914 is among a few well known anthophilic genera in the family Drosophilidae (
Two host plant species of Colocasiomyia flies A–D Rhaphidophora peepla (Roxb.) Schott (Ertaipo, Mt. Gaoligongshan, Yunnan, China) E–H R. crassicaulis Engl. & Karause (Qimaba, Lüchun, Yunnan, China) A, E plants climbing on tall trees B, F inflorescence buds C, G inflorescences D, H infructescences (with leaves shown together in H).
Colocasiomyia specimens were collected in western (Baoshan) and southern (Lüchun) Yunnan using an insect net (for adults) or by dissecting host inflorescences or infructescences (for eggs or dormant larvae within egg capsules on host infructescences), and immediately preserved in 70% (for morphological observation) or 100% (for DNA sequencing) ethanol (Table
Colocasiomyia specimens (noted in cases of egg or larval stage) used for DNA sequencing in the present study.
Species | Voucher specimens (voucher #/Stage/Gender) and collection data |
---|---|
C. hailini | 01518−01522/larva/-, 01727−01731/egg/-, 01295/egg/-, 01296/egg/-; Laomengshan, Baihualing, Baoshan, Yunnan, China; ex. spadices or spathes of R. decursiva |
C. liae sp. nov. | 10485/adult/♂, 10486/adult/♀; Qimaba, Lüchun, Yunnan, China; ex. inflorescences of R. crassicaulis |
09562−09267/adult/♂, 09568−09272/adult/♀; Qimaba, Lüchun, Yunnan, China; reared from infructescences of R. crassicaulis | |
C. longifilamentata | 01133/adult/♂, 01252/egg/-, 0158/egg/-, 01588/larva/-; Laomengshan, Baihulaling, Baoshan, Yunnan, China; ex. inflorescences of R. decursiva |
C. longivalva | 01722/adult/♀; Laomengshan, Baihulaling, Baoshan, Yunnan, China; ex. inflorescences of R. decursiva |
10103/adult/♂, 10111/adult/♂, 10114/adult/♂, 10120/adult/♀, 10121/adult/♂, 10124/adult/♀, 10127/adult/♂, 10132/adult/♀, 10133/adult/♂, 10134/adult/♀, 10135/adult/♀, 10139/adult/♂, 10143/adult/♀, 10145/adult/♀, 10146/adult/♀; Ertaipo, Baihualing, Baoshan, Yunnan, China; ex. inflorescences of R. peepla | |
C. todai sp. nov. | 10100/adult/♀, 10102/adult/♀, 10105−10110/adult/♀, 10112/adult/♀, 10113/adult/♀, 10115−10118/adult/♂, 10122/adult/♂, 10128/adult/♀, 10129/adult/♂, 10130/adult/♀, 10131/adult/♂, 10136−10138/adult/♀, 10140−10142/adult/♀, 10144/adult/♀; same collection data as above |
C. yini | 10123/adult/♂; same collection data as above |
We observed external morphology and detailed structures of dissected organs following the methods in
Adult Colocasiomyia specimens, either newly obtained or collected previously, were first identified and sorted into species of the C. gigantea group in light of morphology. DNA barcodes (i.e., the 658-bp barcoding region of the mitochondrial cytochrome c oxydase I, COI, gene) were determined for representing specimens (adults, eggs, or larvae) of each morpho-species (Table
A total of 60 adult specimens of the C. gigantea group were morphologically sorted into five species, including three known (C. longifilamentata, C. longivalva, and C. yini) and two new ones (C. todai sp. nov. and C. liae sp. nov.) (Table
The two NJ trees based on p- and K2P-distances showed the identical topology, but slightly differed in terms of BP: the tree based on p-distances yielded overall higher BPs (Fig.
Unrooted neighbor-joining tree of the C. gigantea species group built based on p-distances between COI sequences. Label of each operational taxonomic unit (OTU) is given in the form of “voucher number-GenBank accession number”. Numbers beside nodes are bootstrap percentages (shown when ≥ 50; BP based on p-distance/BP based on K2P-distance).
Table
Intra- and interspecific p-distances in the Colocasiomyia gigantea species group.
# | Species name | Number of sequences | Intraspecific distances ([Minimal, Maximal], Mean ± SEa) | Interspecific distancesb | ||||||||
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | ||||
1 | C. gigantea | 4 | [0.0000, 0.0058], 0.0029 ± 0.0016 | 0.0898 | 0.1020 | 0.0940 | 0.1120 | 0.0912 | 0.0890 | 0.0907 | 0.1170 | |
2 | C. hailini | 20 | [0.0000, 0.0065], 0.0017 ± 0.0007 | 0.1006 | 0.0807 | 0.0802 | 0.0904 | 0.0797 | 0.0846 | 0.0768 | 0.0560 | |
3 | C. liae sp. nov. | 13 | [0.0000, 0.0063], 0.0015 ± 0.0008 | 0.1178 | 0.0912 | 0.0857 | 0.0822 | 0.0802 | 0.0923 | 0.0715 | 0.0938 | |
4 | C. longifilamentata | 15 | [0.0000, 0.0130], 0.0047 ± 0.0015 | 0.1081 | 0.0975 | 0.0990 | 0.0706 | 0.0130 | 0.0802 | 0.0595 | 0.1057 | |
5 | C. longivalva | 18 | [0.0000, 0.0049], 0.0006 ± 0.0003 | 0.1186 | 0.1037 | 0.0952 | 0.0796 | 0.0706 | 0.0822 | 0.0711 | 0.1174 | |
6 | C. rhaphidophorae | 9 | [0.0000, 0.0053], 0.0017 ± 0.0008 | 0.0985 | 0.0912 | 0.0978 | 0.0268 | 0.0796 | 0.0857 | 0.0579 | 0.1017 | |
7 | C. scindapsae | 6 | [0.0000, 0.0100], 0.0059 ± 0.0021 | 0.1042 | 0.1053 | 0.1090 | 0.0978 | 0.0952 | 0.0990 | 0.0819 | 0.1177 | |
8 | C. todai sp. nov. | 29 | [0.0000, 0.0030], 0.0003 ± 0.0002 | 0.0983 | 0.0896 | 0.0791 | 0.0729 | 0.0785 | 0.0658 | 0.1058 | 0.1057 | |
9 | C. yini | 6 | [0.0000, 0.0000], 0.0016 ± 0.0010 | 0.1217 | 0.0657 | 0.1047 | 0.1217 | 0.1258 | 0.1141 | 0.1342 | 0.1139 |
C. gigantea (Okada, 1987); C. rhaphidophorae Gao & Toda and C. scindapsae Fartyal & Toda in
This key is updated from that of
1 | Aedeagus not pubescent; aedeagal apodeme as long as or longer than aedeagus (fig. 4CF13, fig. 40L14, fig. 47L14) | 2 |
– | Aedeagus pubescent (except for C. liae sp. nov.); aedeagal apodeme distinctly shorter than aedeagus. | 4 |
2 | Foreleg tarsomere II with seven or eight pegs (fig. 4BF13). Epandrial apodeme medially narrower than epandrium (fig. 4CF13). Aedeagal apodeme much longer than aedeagus (fig. 4CF13). Distal, narrow part of oviscapt much shorter than proximal, broad part, apically shaped like arrowhead, with a pair of stout, peg-like ovisensilla at apex (fig. 4BF13) | C. scindapsae Fartyal & Toda |
– | Foreleg tarsomere II with six pegs (fig. 14L14, fig. 15L14). Epandrial apodeme well developed into distally tapering, triangular extension strongly projected anteriad, twice as long as epandrial width (fig. 38L14, fig. 45L14). Aedeagal apodeme slightly longer than aedeagus (fig. 40L14, fig. 47L14). Distal, narrow part of oviscapt rod-shaped, slightly shorter than proximal, broad part, with only trichoid ovisensilla (fig. 43L14, fig. 50L14) | 3 |
3 | Wing C3F index < 2/3. Distance between antennal sockets same as socket width. Distal, narrow part of oviscapt constricted subbasally on dorsal margin (fig. 43L14) | C. hailini Li & Gao |
– | Wing C3F index > 2/3. Distance between antennal sockets larger than socket width. Distal, narrow part of oviscapt finger-like, not constricted subbasally on dorsal margin (fig.50L14). | C. yini Li & Gao |
4 | Labellum with 14 pseudotracheae per side. Distal, narrow part of oviscapt broadly truncate apically, much shorter than proximal, broad part (fig. 1FO87, fig 2HF13). | C. gigantea (Okada) |
– | Labellum with 17 or more pseudotracheae per side. Distal, narrow part of oviscapt not or only slightly truncate apically, ca 2/5 or longer the length of proximal, broad part. | 5 |
5 | Epandrium notched above basal corner of epandrial ventral lobe (fig. 30L14). Ventral lobe apically with a grooved, finger-like peg (fig. 32L14). Distal, narrow part of oviscapt twice as long as proximal, broad part (fig. 36L14) | C. longivalva Li & Gao |
– | Epandrium not notched along posterior margin. Ventral lobe apically with an ungrooved, inward-curved peg or thick spine. Distal, narrow part of oviscapt as long as or shorter than proximal, broad part. | 6 |
6 | Labellum with 17 pseudotracheae per side. Epandrium broad, with short ventral lobe apically inlaid with thick, long, inward-curved, spine-like seta (Fig. |
C. todai Jiao & Gao, sp. nov. |
– | Labellum with ≥ 21 pseudotracheae per side. Epandrium with long ventral lobe apically inlaid with short peg. | 7 |
7 | Epandrium with setae on posterior margin in addition to those on ventral lobe (fig. 24L14). Distal, narrow part of oviscapt nearly as long as proximal, broad part (fig. 28L14) | C. longifilamentata Li & Gao |
– | Epandrium with no setae on posterior margin above ventral lobe (fig. 3CF13, Figs |
8 |
8 | Labellum with 21−22 pseudotracheae per side. Ventral lobe of epandrium well developed, narrowing distally in lateral view, scabbard-like, with 3 long setae on dorsosubbasal margin, apically inlaid with a short peg; cercus with slightly projected ventral apex (fig. 3CF13). Aedeagal apodeme ca 3/4 the length of aedeagus (fig. 3DF13). Distal, narrow portion of oviscapt narrowing distally, gently curved dorsad (fig. 3GF13) | C. rhaphidophorae Gao & Toda |
– | Labellum with 34 pseudotracheae per side. Ventral lobe of epandrium prolonged like a rod, distally slightly broadened in lateral view, with 2 long and 1 medium-length setae on its insertion, apically inlaid with a relatively long, claw-like peg; circus ventrally lacking projected apex (Fig. |
C. liae Jiao & Gao, sp. nov. |
This species closely resembles C. longivalva in external morphology and structure of male genitalia, but can be distinguished from the latter by epandrium with short ventral lobe apically inlaid with a thick, long, inward-curved, spine-like seta (Fig.
(♂, ♀). Head: Supracervical setae about 17 per side. Dorsomedial arm of tentorial apodeme about 2/3 as long as dorsolateral arm. Eye red, somewhat roundish, lacking interfacetal setulae. Frontal vitta mat, black. First flagellomere not concave on inner margin. Facial carina trapeziform, medially twice as wide as first flagellomere, as long as pedicel and first flagellomere combined. Palpus convex on ventrodistal portion. Cibarium posterior sensilla minute, 1 or 2 per side. Labellum with 19 pseudotracheae per side.
Thorax
(Fig.
Adult males of the new species: lateral habitus, head (anterior view), head and thorax (dorsal view), wing (ventral view of left one in D dorsal view of right one in I), and fore leg (right one, inner view) A–E Colocasiomyia todai Jiao & Gao, sp. nov. (#10122) F–J C. liae Jiao & Gao, sp. nov. (#10485). Scale bars: 1.0 mm except for B, E, G and J (0.5 mm).
Wing
(Fig.
Legs
(Fig.
Abdomen
(Fig.
Male terminalia
(Fig.
Colocasiomyia todai Jiao & Gao, sp. nov. Adult male (holotype #10122) and female (paratype, #10100) from Ertaipo, Gaoligong Mountains, Baoshan, Yunnan, China A periphallic organs (lateral view) B periphallic organs (ventral view) C surstylus (right one, inner view) D phallic organs (dorsal view) E phallic organs (lateral view) F oviscapt (lateral view). Abbreviations: aed = aedeagus, aed a = aedeagal apodeme, aed b p = aedeagal basal process, cerc = cercus, epand = epandrium, epand a = epandrial apodeme, hypd = hypandrium, pm = paramere, 10S = tenth sternite. Scale bars: 0.1 mm.
Female terminalia
(Fig.
Measurements : BL = 3.53 (range in 5♂ paratypes: 3.27−3.38; range in 5♀ paratypes: 3.25−3.70) mm, ThL = 1.67 (1.56−1.72; 1.49−1.73) mm, WL = 3.38 (3.09−3.45; 3.15−3.45) mm, WW = 1.50 (1.25−1.48; 1.30−1.50) mm.
Indices : arb = 0/0 (5♂, 5♀, or less if noted, paratypes: 0/0), FW/HW = 0.56 (0.56−0.59), ch/o = 0.59 (0.45−0.57), prorb = 0.87 (0.75−0.99), rcorb = 0.38 (0.35−0.49), orbito = 0.68 (0.64−0.95), vb = 0.46 (0.34−0.49), dcl = 0.51 (4♂, 5♀: 0.49−0.56), dcp = 1.09 (4♂, 5♀: 1.02−1.19), sterno = 0.97 (0.75−0.98), sctl = 0.75 (4♂, 5♀: 0.73−0.84), sctlp = 1.47 (1.28−1.47), C = 1.82 (1.82−2.11), 4c = 1.26 (1.05−1.25), 4v = 2.02 (1.76−2.04), 5x = 0.91 (0.89−1.06), ac = 3.45 (3.09−3.82), M = 0.46 (0.38−0.48), C3F = 0.80 (0.73−0.85).
Holotype
♂ (#10122): China: ex inflorescence of Rhaphidophora peepla (Roxb.) Schott, Ertaipo, Gaoligong Moutains, Baoshan, Yunnan, China, 25°18.0'N, 98°47.0'E, ca 2200 m, 31.vii.2019, Jian-Jun Gao and Xue-Lin Ye (
China (Yunnan).
Rhaphidophora peepla (Roxb.) Schott (Fig.
Patronym, in honor of Professor Masanori J. Toda (Hokkaido University), who dedicated himself to the studies of taxonomy and flower-visiting/breeding behaviors of Colocasiomyia flies.
This species closely resembles C. rhaphidophorae in external morphology and structure of male and female genitalia, but can be distinguished from the latter by epandrial ventral lobe rod-like, distally slightly broadened in lateral view, apically inlaid with a relatively long, claw-like peg (Fig.
Colocasiomyia liae Jiao & Gao, sp. nov. Adult male (holotype #10485) and female (paratype, #10486) from Qimaba, Lüchun, Yunnan, China. A periphallic organs (lateral view) B periphallic organs except cerci (ventral view) C tenth sternite (posteroventral view) D phallic organs (dorsal view) E phallic organs (lateral view) F oviscapt (lateral view). Abbreviations: aed = aedeagus, aed a = aedeagal apodeme, aed b p = aedeagal basal process, cerc = cercus, epand = epandrium, epand a = epandrial apodeme, hypd = hypandrium, pm = paramere, 10S = tenth sternite. Scale lines: 0.1 mm.
(♂, ♀). Head: Supracervical setae about 9−10 per side. Dorsomedial arm of tentorial apodeme about 1/3 as long as dorsolateral arm. Eye red, somewhat roundish, with very sparse interfacetal setulae. Frontal vitta mat, black. First flagellomere not concave on inner margin. Facial carina broad trapeziform, medially twice as wide as first flagellomere, as long as pedicel and first flagellomere combined. Palpus convex on ventrodistal portion. Cibarium posterior sensilla minute, 2 or 3 per side. Labellum with 34 pseudotracheae per side.
Thorax
(Fig.
Wing
(Fig.
Legs
(Fig.
Abdomen
(Fig.
Male terminalia
(Fig.
Female terminalia
(Fig.
Measurements : BL = 2.45 (1♀ paratype: 2.60) mm, ThL = 1.10 (0.93) mm, WL = 2.08 (1.80) mm, WW = 0.91 (0.75) mm.
Indices : arb = 0/0 (1♀ paratype: 0/0), FW/HW = 0.57 (0.58), ch/o = 0.47 (0.51), prorb =1.14 (1.05), rcorb = 0.28 (0.45), orbito = 0.67 (0.86), vb = 0.34 (0.35), dcl = 0.53 (0.53), dcp = 0.97 (0.96), sterno = 0.75 (0.68), sctl = 0.56 (0.59), sctlp = 1.20 (1.03), C = 1.75 (1.64), 4c = 1.12 (1.18), 4v = 1.58 (1.70), 5x = 0.88 (0.92), ac = 3.94 (3.68), M = 0.36 (0.34), C3F = 0.82 (0.74).
Holotype
♂ (#10485): China: ex inflorescence of Rhaphidophora crassicaulis Engl. & Krause, Qimaba, Lüchun, Yunnan, China, 22°48.0'N, 102°15.0'E, ca 750 m, 6.vii.2020, Jian-Jun Gao and Run-Jie Jiao (
China (Yunnan).
Rhaphidophora crassicaulis Engl. & Krause (Fig.
Patronym, in honor of Professor Heng Li (Kunming Institute of Botany, Chinese Academy of Sciences), who helped us with the identifications of various aroid host plants of Colocasiomyia flies.
Numbers of adults of this species were obtained by rearing infructescences of R. crassicaulis collected on November 1, 2018 from the type locality, indicating that this species breeds on inflorescences/infructescences of this host plant. These adults were not defined as type specimens due to obviously insufficient body pigmentation and sclerotization, though some of them were used for DNA barcoding.
We thank the Administrations of the Huanglianshan National Nature Reserve and the Gaoligongshan National Nature Reserve for permission and supports of the field works within the Reserves. We also thank Mr Shi-Bin Jiao and Mr Xue-Lin Ye for their help in the field. This work was supported by the National Natural Science Foundation of China (Nos. 31572238).