Research Article |
Corresponding author: Jian-Jun Gao ( gao-leyun@263.net ) Academic editor: Rudolf Meier
© 2018 Takehiro K. Katoh, Guang Zhang, Masanori J. Toda, Awit Suwito, 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:
Katoh T, Zhang G, Toda M, Suwito A, Gao J (2018) A revision of the subgenus Dudaica Strand of the genus Drosophila Fallén, with descriptions of six new species (Diptera, Drosophilidae). ZooKeys 781: 19-50. https://doi.org/10.3897/zookeys.781.27354
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The subgenus Dudaica Strand of the genus Drosophila Fallén has been known to comprise only two species: Drosophila (Dudaica) senilis Duda, 1926 (recorded from Indonesia, Philippines, Vietnam, Bhutan, and India) and D. malayana (Takada, 1976) (recorded from Malaysia). In the present study, this subgenus is revised, with D. malayana redescribed and six new species discovered and described from China, Malaysia, and Indonesia: gracilipalpis Katoh & Gao, sp. n., puberula Katoh & Gao, sp. n., albipalpis Katoh, Toda & Gao, sp. n., qiongzhouensis Katoh & Gao, sp. n., orthophallata Katoh, Toda & Gao, sp. n., and dissimilis Katoh & Gao, sp. n. Both morphological and molecular data (DNA barcodes) are used to distinguish the above species. A key to species of this subgenus is provided.
China, DNA barcoding, Southeast Asia, taxonomy
The phylogenetic position of the subgenus Dudaica remains unresolved.
In this paper, we revise the subgenus Dudaica, and add six new species discovered from China, Malaysia, and Indonesia to this subgenus, by identifying them with the aid of DNA sequences of the 658-bp barcoding region of the mitochondrial COI (cytochrome c oxidase subunit I) gene. We also redescribe the known species D. malayana, based on specimens newly collected from Malaysia and Indonesia. Finally, a key to all the eight species of Dudaica is given.
All specimens employed in the present study were collected from China, Malaysia, and Indonesia (Table
Species | Sex | Voucher # a | Collection site | Elevation (m) | Collection date | GenBank accession # |
---|---|---|---|---|---|---|
malayana (Takada, 1976) | ♂ | #03903 | Poring, Sabah, Malaysia | 600 | 20.iii.2008 | MH410612 |
♂ | #03904 | Ditto | 600 | 13.iii.2008 | MH410613 | |
♀ | n/a | Gunung Poteng, West Kalimantan, Indonesia | 220 | 4.xii.1996 | n/a | |
gracilipalpis sp. n. | ♂ | #00033 | Xishuangbanna Tropical Botanical Garden, Mengla, Xishuangbanna, Yunnan, China | 650 | 19.iii.2006 | MH410614 |
♂ | #03423 | Ditto | 650 | 27–28.ix.2011 | MH410620 | |
♂ | #06001 | Ditto | 650 | 28.ix.2011 | MH410624 | |
♂ | #00484 | Wangtianshu, Mengla, Xishuangbanna, Yunnan, China | 670 | 22–25.iv.2007 | MH410615 | |
♂ | #00491 | Ditto | 670 | 30.ix.2011 | MH410617 | |
♂ | #03364 | Ditto | 670 | 30.ix.2011 | MH410619 | |
♂ | #03424 | Ditto | 670 | 30.ix.2011 | MH410621 | |
♂ | #03425 | Ditto | 670 | 30.ix.2011 | MH410622 | |
♀ | #00485 | Ditto | 670 | 22–25.iv.2007 | MH410616 | |
♀ | #00492 | Ditto | 670 | 30.ix.2011 | MH410618 | |
♂ | #03902 | Bogor, West Java, Indonesia | 260 | 14–15.xi.2009 | MH410623 | |
puberula sp. n. | ♂ | #03365 | Xishuangbanna Tropical Botanical Garden, Mengla, Xishuangbanna, Yunnan, China | 650 | 27–28.ix.2011 | MH410626 |
♀ | #03426 | Ditto | 650 | 19.iii.2006 | MH410631 | |
♂ | #00480 | Wangtianshu, Mengla, Xishuangbanna, Yunnan, China | 670 | 22–25.iv.2007 | MH410625 | |
♂ | #03366 | Ditto | 670 | 30.ix.2011 | MH410627 | |
♂ | #03367 | Ditto | 670 | 30.ix.2011 | MH410628 | |
♂ | #03368 | Ditto | 670 | 30.ix.2011 | MH410629 | |
♂ | #03369 | Ditto | 670 | 30.ix.2011 | MH410630 | |
albipalpis sp. n. | ♂ | #03908 | Cikaniki, Gunung Halimun, West Java, Indonesia | 1050 | 4.xi.2009 | MH410632 |
qiongzhouensis sp. n. | ♂ | #03310 | Jianfengling National Nature Reserve, Ledong, Hainan, China | 750 | 17–20.iv.2008 | MH410633 |
♂ | #03311 | Ditto | 750 | 17–20.iv.2008 | MH410634 | |
♂ | #03312 | Ditto | 750 | 17–20.iv.2008 | MH410635 | |
♂ | #03418 | Ditto | 750 | 17–20.iv.2008 | MH410639 | |
♂ | #03419 | Ditto | 750 | 17–20.iv.2008 | MH410640 | |
♂ | #03420 | Ditto | 750 | 17–20.iv.2008 | MH410641 | |
♀ | #03313 | Ditto | 750 | 17–20.iv.2008 | MH410636 | |
♀ | #03314 | Ditto | 750 | 17–20.iv.2008 | MH410637 | |
♀ | #03315 | Ditto | 750 | 17–20.iv.2008 | MH410638 | |
♀ | #03422 | Ditto | 750 | 17–20.iv.2008 | MH410642 | |
orthophallata sp. n. | ♂ | #00177 | Ulu Senagang, Crocker Range, Sabah, Malaysia | 540 | 18.x.1999 | n/a |
♀ | #03905 | Park Headquarters, Mt. Kinabalu, Sabah, Malaysia | 1700 | 11.iii.2008 | MH410644 | |
♀ | #03906 | Ditto | 1700 | 11.iii.2008 | MH410645 | |
dissimilis sp. n. | ♂ | #00430 | Hesong, Xiding, Menghai, Xishuangbanna, Yunnan, China | 1900 | 7.iv.2011 | MH410646 |
Specimens were first identified as of the subgenus Dudaica, based on their overall resemblance to the two known species of this subgenus, D. senilis and D. malayana, especially in body color pattern, shape of palpus, and structures of male/female terminalia. The holotype specimen of D. malayana was examined for the reference. As for D. senilis, we referred to
The specimens were then subjected to DNA barcoding analysis (
For species illustration, the external morphology and detailed structures of male and female terminalia, and head and mouth parts were microphotographed using a Dino-Lite® Microscope Eyepiece Camera (ANMO Electronics Corporation). We followed
A total of 34 COI sequences of 658-bp were determined in this study (Table
Figure
Species | Intraspecific distance | Interspecific distanceb | |||||||
---|---|---|---|---|---|---|---|---|---|
Mean (SEa) | Range | 1 | 2 | 3 | 4 | 5 | 6 | 7 | |
1. malayana (Takada) | 0.0163 (n/a) | 0.0163 | 0.0796−0.0906 | 0.1279−0.1378 | 0.1195−0.1248 | 0.1262−0.1356 | 0.1343−0.1392 | 0.1230−0.1244 | |
2. gracilipalpis sp. n. | 0.0057 (0.0014) | 0.0000−0.0144 | 0.0832 (0.0106) | 0.1260−0.1386 | 0.1125−0.1172 | 0.1225−0.1279 | 0.1431−0.1528 | 0.1293−0.1335 | |
3. puberula sp. n. | 0.0089 (0.0023) | 0.0015−0.0152 | 0.1321 (0.0128) | 0.1329 (0.0132) | 0.0949−0.1004 | 0.1231−0.1262 | 0.1254−0.1336 | 0.1227−0.1277 | |
4. albipalpis sp. n. | n/a | n/a | 0.1221 (0.0129) | 0.1143 (0.0129) | 0.0964 (0.0114) | 0.0844−0.0861 | 0.0870−0.0896 | 0.1215 | |
5. qiongzhouensis sp. n. | 0.0005 (0.0005) | 0.0000−0.0015 | 0.1304 (0.0130) | 0.1245 (0.0115) | 0.1248 (0.0123) | 0.0858 (0.0113) | 0.1135−0.1184 | 0.1178 | |
6. orthophallata sp. n. | 0.0035 (n/a) | 0.0035 | 0.1361 (0.0138) | 0.1456 (0.0143) | 0.1297 (0.0137) | 0.0883 (0.0114) | 0.1154 (0.0134) | 0.1360−0.1385 | |
7. dissimilis sp. n. | n/a | n/a | 0.1237 (0.0133) | 0.1313 (0.0127) | 0.1260 (0.0126) | 0.1215 (n/a) | 0.1178 (0.0124) | 0.1373 (0.0144) |
Bayesian tree of seven species of the subgenus Dudaica based on COI gene sequences, with indication of the results of molecular species delimitation by the ABGD (solid bars) and GMYC (open bars) analyses. Label of each operational taxonomic unit (OTU) is given in the format of “voucher number (gender) province of collection locality” (bold voucher numbers: holotype specimens). Numbers beside nodes are posterior probabilities.
Among them, D. qiongzhouensis sp. n. and D. albipalpis sp. n. are very similar in morphology to each other. The diagnoses for these species are supplemented with “pure” molecular diagnostic characters, which are defined as sites with fixed status in the COI sequence alignments within the focal species but differing from the other species (
Selected diagnostic nucleotide sites (indicated in square brackets) for D. albipalpis sp. n. and D. qiongzhouensis sp. n. in the COI sequences. Sequences of the other five species in the subgenus Dudaica are shown for comparison.
Species | Sequence | Diagnostic nucleotide sites | |||
---|---|---|---|---|---|
92 | 226 | 391 | 589 | ||
albipalpis sp. n. | #03908 | T | T | [C] | [T] |
qiongzhouensis sp. n. | #03310 | [C] | [C] | T | C |
#03311 | [C] | [C] | T | C | |
#03312 | [C] | [C] | T | C | |
#03313 | [C] | [C] | T | C | |
#03314 | [C] | [C] | T | C | |
#03315 | [C] | [C] | T | C | |
#03418 | [C] | [C] | T | C | |
#03419 | [C] | [C] | T | C | |
#03420 | [C] | [C] | T | C | |
#03422 | [C] | [C] | T | C | |
malayana (Takada) | #03903 | T | T | T | C |
#03904 | T | T | T | C | |
gracilipalpis sp. n. | #00484 | T | T | T | C |
#00033 | T | T | T | C | |
#00485 | T | T | T | C | |
#00491 | T | T | T | C | |
#00492 | T | T | T | C | |
#03364 | T | T | T | C | |
#03423 | T | T | T | C | |
#03424 | T | T | T | C | |
#03425 | T | T | T | C | |
#03902 | T | T | T | C | |
#06001 | T | T | T | C | |
puberula sp. n. | #00480 | T | T | T | C |
#03365 | T | T | T | C | |
#03366 | T | T | T | C | |
#03367 | T | T | T | C | |
#03368 | T | T | T | C | |
#03369 | T | T | T | C | |
#03426 | T | T | T | C | |
orthophallata sp. n. | #03905 | T | T | T | C |
#03906 | T | T | T | C | |
dissimilis sp. n. | #00430 | T | T | T | C |
Dudaica Strand, 1943: 212. New name for Macropalpus Duda.
Macropalpus Duda, 1926: 63. Type species: Drosophila senilis Duda, 1926. Preoccupied by Macropalpus Ratzeburg, 1844 (Braconidae). Proposed as a subgenus.
Head, scutum, and scutellum mostly milky white, contrasting with mostly dark brown thoracic pleura (Figures
Head (anterior view). A malayana (Takada) (#03903) B gracilipalpis sp. n. (paratype #00492) C puberula sp. n. (paratype #03366) D albipalpis sp. n. (holotype #03908) E qiongzhouensis sp. n. (paratype #03313) F orthophallata sp. n. (paratype #03905) G dissimilis sp. n. (holotype #00430). Abbreviation: plp = palpus. Scale bars: 0.5 mm.
Head (Figures
Postocciput (caudal view). A malayana (Takada) (#03903) B gracilipalpis sp. n. (paratype #00492) C puberula sp. n. (paratype #03366) D albipalpis sp. n. (holotype #03908) E qiongzhouensis sp. n. (paratype #03313) F orthophallata sp. n. (paratype #03905) G dissimilis sp. n. (holotype #00430). Abbreviations: poc s = postocellar setae, spc s = supracervical setae. Scale bars: 0.1 mm.
Cibarium of Dudaica species (part 1; dorsal and lateral views). A malayana (Takada) (#03903) B gracilipalpis sp. n. (paratype #00492) C puberula sp. n. (paratype #03366) D albipalpis sp. n. (holotype #03908). Abbreviations: cly = clypeus, m s = medial sensilla, p s = posterior sensilla. Scale bars: 0.1 mm.
Prementum (ventral and lateral views). A malayana (Takada) (#03903) B gracilipalpis sp. n. (paratype #00492) C puberula sp. n. (paratype #03366) D albipalpis sp. n. (holotype #03908) E qiongzhouensis sp. n. (paratype #03313) F orthophallata sp. n. (paratype #03905) G dissimilis sp. n. (holotype #00430). Scale bars: 0.1 mm.
Palpus (lateral and dorsal views; only A is photographed at right side and flipped horizontally). A malayana (Takada) (#03903) B gracilipalpis sp. n. (paratype #00492) C puberula sp. n. (paratype #03366) D albipalpis sp. n. (holotype #03908) E qiongzhouensis sp. n. (paratype #03313) F orthophallata sp. n. (paratype #03905) G dissimilis sp. n. (holotype #00430). Scale bars: 0.1 mm.
Thorax (Figures
Wing (Figures
Legs (Figures
Drosophila (Dudaica) malayana (Takada) (A, K #03903 B–J #03904 L–N female specimen). A tibia of right foreleg (apical setae indicated with red arrowheads; ventral view) B, C periphallic organs (caudal and caudolateral view, respectively) D surstyli (caudoventral view) E surstylus and epandrial ventral lobe (caudolateral view) F tenth sternite and ventral protrusions of cerci (ventral view) G–I phallic organs (ventral, ventrolateral, and lateral view, respectively) J pubescence on caudolateral lobe of hypandrium (indicated with red arrowhead; caudoventral view) K sensilla (indicated with red arrowheads) and apical portion of parameres (ventral view) L, M oviscapt (lateral and ventral view, respectively) N spermatheca. Abbreviations: aed = aedeagus, aed a = aedeagal apodeme, cerc = cercus, epand = epandrium, hypd = hypandrium, pm = paramere, sur = surstylus. Scale bars: 0.1 mm.
Drosophila (Dudaica) gracilipalpis sp. n. (A, M–O paratype #00492 B–K paratype #03902 L paratype #00491). A tibia of right foreleg (apical seta indicated with red arrowhead; ventral view) B, C periphallic organs (caudal and caudolateral view, respectively) D surstyli (caudoventral view) E surstylus and epandrial ventral lobe (caudolateral view) F tenth sternite and ventral protrusions of cerci (ventral view) G–I phallic organs (ventral, ventrolateral, and lateral view, respectively) J pubescence on caudolateral lobe of hypandrium (indicated with red arrowhead; caudoventral view) K, L sensilla (indicated with red arrowheads) and apical portion of parameres (ventral view; Indonesian and Chinese specimen, respectively) M, N oviscapt (lateral and ventral view, respectively) O spermatheca. Scale bars: 0.1 mm.
Abdomen (Figures
Male terminalia (Figures
Drosophila (Dudaica) puberula sp. n. (A–J paratype #03366; K holotype #03365 L–M paratype #03426). A, B periphallic organs (caudal and caudolateral view, respectively) C surstyli (caudoventral view) D surstylus and epandrial ventral lobe (caudolateral view) E tenth sternite and ventral protrusions of cerci (ventral view) F–H phallic organs (ventral, ventrolateral, and lateral view, respectively) I, J pubescence (indicated with red arrowheads) and apical portion of parameres (ventrolateral and lateral view, respectively) K sensillum (indicated with red arrowhead) and apical portion of parameres (ventral view) L, M oviscapt (lateral and ventral view, respectively) N spermatheca. Scale bars: 0.1 mm.
Drosophila (Dudaica) albipalpis sp. n. (A–K holotype #03908). A, B periphallic organs (caudal and caudolateral view, respectively) C surstyli (caudoventral view) D surstylus and epandrial ventral lobe (caudolateral view) E tenth sternite and ventral protrusions of cerci (ventral view) F–H phallic organs (ventral, ventrolateral, and lateral view, respectively) I apical portion of aedeagus with fine serrations (indicated with red arrowhead; ventrolateral view) J, K sensilla (indicated with red arrowheads) and apical portion of parameres (ventral and lateral view, respectively). Scale bars: 0.1 mm.
Drosophila (Dudaica) qiongzhouensis sp. n. (A–K paratype #03418 L–N paratype #03313). A, B periphallic organs (caudal and caudolateral view, respectively) C surstyli (caudoventral view) D surstylus and epandrial ventral lobe (caudolateral view) E tenth sternite and ventral protrusions of cerci (ventral view) F–H phallic organs (ventral, ventrolateral, and lateral view, respectively) I apical portion of aedeagus with fine serrations (indicated with red arrowhead; ventrolateral view) J, K sensilla (indicated with red arrowheads) and/or apical portion of parameres (ventral and lateral view, respectively) L, M oviscapt (lateral and ventral view, respectively) N spermatheca. Scale bars: 0.1 mm.
Drosophila (Dudaica) orthophallata sp. n. (A–H holotype #00177 I, J paratype #03906). A, B periphallic organs (caudal and caudolateral view, respectively) C surstyli, tenth sternite, and ventral protrusions of cerci (ventral view) D epandrial ventral lobe (caudolateral view) E–G phallic organs (ventral, ventrolateral, and lateral view, respectively) H sensilla (indicated with red arrowheads) and apical portion of parameres (ventral view) I, J oviscapt (lateral and ventral view, respectively). Scale bars: 0.1 mm.
Drosophila (Dudaica) dissimilis sp. n. (A–J holotype #00430). A, B periphallic organs (caudal and caudolateral view, respectively) C surstyli (caudoventral view) D surstylus and epandrial ventral lobe (caudolateral view) E tenth sternite (ventral view) F–H phallic organs (ventral, ventrolateral, and lateral view, respectively) I, J sensilla (indicated with red arrowheads) and apical portion of parameres (ventral and lateral view, respectively). Scale bars: 0.1 mm.
Female terminalia (Figures
senilis Duda, malayana (Takada), gracilipalpis Katoh & Gao, sp. n., puberula Katoh & Gao, sp. n., albipalpis Katoh, Toda & Gao, sp. n., qiongzhouensis Katoh & Gao, sp. n., orthophallata Katoh, Toda & Gao, sp. n., and dissimilis Katoh & Gao, sp. n.
In this key, not only morphological characters but also the selected “pure” diagnostic nucleotide sites of COI are used to distinguish between albipalpis sp. n. and qiongzhouensis sp. n. (see also Table
1 | Palpus sinuate, with several stout setae only on basal portion (Figures |
2 |
– | Palpus nearly straight, with several stout setae scattered on entire length (Figures |
4 |
2 | Cercus without distinct caudoventral process (figure G&S90–2E) | senilis Duda, 1926 |
– | Cercus with distinct caudoventral process (Figures |
3 |
3 | Foreleg tibia with two apical setae (Figure |
malayana (Takada, 1976) |
– | Foreleg tibia with one apical seta (Figure |
gracilipalpis Katoh & Gao, sp. n. |
4 | Palpus medio- to baso-laterally dark grayish brown, with one apical, stout, prominent seta (Figure |
dissimilis Katoh & Gao, sp. n. |
– | Palpus entirely white, without apical, prominent seta (Figure |
5 |
5 | Palpus with 3–4 prominent, stout setae on outer, lateral surface (Figure |
puberula Katoh & Gao, sp. n. |
– | Palpus with many setae varied in size on outer, lateral surface (Figure |
6 |
6 | Aedeagus apically without small, acute claw, subapically swollen in lateral view (Figure |
orthophallata Katoh, Toda & Gao, sp. n. |
– | Aedeagus apically with small, acute claw, subapically not swollen in lateral view (Figures |
7 |
7 | Palpus broad, flat, not so rod-shaped in lateral view (Figure |
albipalpis Katoh, Toda & Gao, sp. n. |
– | Palpus slender, not so flat, somewhat rod-shaped in lateral view (Figure |
qiongzhouensis Katoh & Gao, sp. n. |
Zygothrica malayana Takada, 1976: 68.
Drosophila (Dudaica) malayana
:
MALAYSIA: 2♂ (holotype and paratype of Zygothrica malayana Takada, 1976), near Kuala Lumpur, Peninsular Malaysia, 3.vii.1972, H Takada (
Palpus long, sinuate, with several stout setae only on basal portion (Figure
Adult ♂ and ♀. Head (Figures
Thorax (Figure
Legs (Figures
Abdomen (Figure
Male terminalia (Figure
Female terminalia (Figure
Measurements (in mm): BL (straight distance from anterior edge of pedicel to tip of abdomen) = 2.13–2.39/2.27 (range in 2♂/1♀ specimens), ThL (distance from anterior notal margin to apex of scutellum) = 0.98–1.08/1.01, WL (distance from humeral cross vein to wing apex) = 1.78–1.92/1.73, WW (maximum wing width) = 0.89–0.96/0.86.
Indices. FW/HW (frontal width/head width) = 0.56–0.60 (range in 2♂ and 1♀ specimens), ch/o (maximum width of gena/maximum diameter of eye) = 0.08–0.12, prorb (proclinate orbital seta/posterior reclinate orbital seta in length) = 0.61–0.62, rcorb (anterior reclinate orbital seta/posterior reclinate orbital seta in length) = 0.29–0.32, vb (subvibrissa/vibrissa in length) = 0.41–0.59, orbito (distance between proclinate and posterior reclinate orbital setae/distance between inner vertical and posterior reclinate orbital setae) = 0.50–0.67, dcl (anterior dorsocentral seta/posterior dorsocentral seta in length) = 0.64–0.71, sctl (basal scutellar seta/apical scutellar seta in length) = 0.95–0.98, sterno (anterior katepisternal seta/posterior katepisternal seta in length) = 0.59–0.74, dcp (distance between ipsilateral dorsocentral setae/distance between anterior dorsocentral setae) = 0.39–0.44, sctlp (distance between ipsilateral scutellar setae/distance between apical scutellar setae) = 0.94–1.09, C (2nd costal section between subcostal break and R2+3/3rd costal section between R2+3 and R4+5) = 1.75–1.81, 4c (3rd costal section between R2+3 and R4+5/M1 between r-m and dm-cu) = 1.25–1.36, 4v (M1 between dm-cu and wing margin/M1 between r-m and dm-cu) = 1.75–1.95, 5x (CuA1 between dm-cu and wing margin/dm-cu between M1 and CuA1) = 2.12–2.25, ac (3rd costal section between R2+3 and R4+5/distance between distal ends of R4+5 and M1) = 2.56–3.00, M (CuA1 between dm-cu and wing margin/M1 between r-m and dm-cu) = 0.75–0.86, C3F (length of heavy setation in 3rd costal section/length of 3rd costal section) = 0.50–0.62.
Malaysia (Peninsular Malaysia, Sabah*), Indonesia* (West Kalimantan). *New records.
Drosophila malayana was originally described based only on male specimens collected from Peninsular Malaysia, with illustrations but only a very brief description of male terminalia (
Holotype ♂ (#00484): CHINA: Wangtianshu, Mengla, Yunnan, ca. 670 m a.s.l., 22‒25.iv.2007, JJ Gao (
Paratypes. INDONESIA: 1♂ (#03902), Bogor, West Java, 14‒15.xi.2009, MJ Toda (
Palpus long, sinuate, with several stout setae only on basal portion (Figure
(characters in common with D. malayana not repeated). Adult ♂ and ♀. Head (Figures
Thorax (Figure
Abdomen (Figure
Male terminalia (Figure
Female terminalia (Figure
Measurements (in mm): BL = 2.20 in holotype (range in 8♂/2♀ paratypes: 1.63–2.50/2.03–2.63), ThL = 1.00 (0.88–1.06/0.98–1.06), WL = 1.84 (1.60–2.15/2.03–2.10), WW = 0.92 (0.83–1.04/1.02–1.06).
Indices. FW/HW = 0.59 (range in 8♂ and 2♀, or less if noted, paratypes: 0.58–0.63), ch/o = 0.12 (0.07–0.13), prorb = 0.65 (6♂, 2♀: 0.46–0.63), rcorb = 0.30 (7♂, 2♀: 0.20–0.37), vb = 0.45 (7♂, 2♀: 0.22–0.60), orbito = 0.63 (0.50–0.64), dcl = 0.72 (0.63–0.79), sctl = 1.26 (7♂, 1♀: 0.94–1.12), sterno = 0.56 (0.44–0.71), dcp = 0.38 (0.41–0.57), sctlp = 0.95 (0.91–1.00), C = 1.70 (1.56–2.00), 4c = 1.45 (1.32–1.58), 4v = 2.05 (1.83–2.26), 5x = 2.38 (1.80–2.43), ac = 2.90 (2.87–3.29), M = 0.90 (0.75–0.95), C3F = 0.56 (0.52–0.59).
Indonesia (West Java), China (Yunnan).
Referring to the long and thin palpus.
This species closely resembles the foregoing species, D. malayana, in having the long, sinuate palpus and pubescent caudolateral lobe of hypandrium, but can be distinguished from the latter by having only one apical seta on foreleg tibia.
Holotype ♂ (#03365): CHINA: Xishuangbanna Tropical Botanical Garden, Mengla, Yunnan, ca. 650 m a.s.l., 27–28.ix.2011, JJ Gao (
Paratypes. CHINA: 1♀ (#03426), same except for 19.iii.2006, MJ Toda and K Takenaka (
Palpus slightly shorter than arista, with 3–4 prominent, stout setae on outer lateral surface (Figure
(characters in common with D. gracilipalpis sp. n. not repeated). Adult ♂ and ♀. Head (Figures
Thorax (Figure
Legs (Figure
Male terminalia (Figure
Female terminalia (Figure
Measurements (in mm): BL = 2.30 in holotype (range in 5♂/1♀ paratypes: 1.92–2.43/2.69), ThL = 0.88 (0.98–1.14/1.30), WL = 1.86 (1.82–2.10/2.35), WW = 0.98 (0.88–1.10/1.20).
Indices. FW/HW = 0.57 (range in 5♂ and 1♀, or less if noted, paratypes: 0.55–0.58), ch/o = 0.11 (0.07–0.18), prorb = 0.61 (0.54–0.58), rcorb = 0.30 (5♂: 0.27–0.38), vb = 0.34 (0.33–0.41), orbito = 0.63 (0.50–0.75), dcl = 0.65 (4♂, 1♀: 0.55–0.74), sctl = 1.05 (4♂, 1♀: 1.00–1.10), sterno = 0.67 (5♂: 0.42–0.67), dcp = 0.52 (0.42–0.55), sctlp = 1.05 (0.78–1.00), C = 1.67 (1.50–1.79), 4c = 1.43 (1.36–1.58), 4v = 2.00 (1.75–2.16), 5x = 1.70 (1.79–2.11), ac = 3.00 (2.83–3.58), M = 0.81 (0.76–0.89), C3F = 0.55 (0.53–0.59).
China (Yunnan).
Referring to the pubescence of hypandrium on the portion fused to paramere in the new species.
Holotype ♂ (#03908): INDONESIA: Cikaniki, Gunung Halimun, West Java, 4.xi.2009, MJ Toda (
Palpus broad, flat, with a few moderate setae on outer lateral surface (Figure
(characters in common with D. puberula sp. n. not repeated). Adult ♂. Head (Figures
Thorax (Figure
Legs (Figures
Male terminalia (Figure
Measurements (in mm): BL = 2.50 in holotype, ThL = 1.08, WL = 2.10, WW = n/a.
Indices. FW/HW = 0.59, ch/o = 0.09, prorb = 0.63, rcorb = 0.24, vb = n/a, orbito = 0.50, dcl = 0.67, sctl = 1.02, sterno = 0.70, dcp = 0.50, sctlp = 0.91, C = 1.71, 4c = 1.31, 4v = 1.65, 5x = 1.38, ac = 2.83, M = 0.69, C3F = 0.49.
Indonesia (West Java).
Referring to the white palpus in the new species.
Holotype ♂ (#03420): CHINA: Jianfengling National Nature Reserve, Ledong, Hainan, ca. 750 m a.s.l., 17–20.iv.2008, JJ Gao (
Paratypes. CHINA: 5♂, 4♀ (#03310–15, #03418, #03419, #03422), same data as holotype (
Palpus slender, not so flat, somewhat rod-shaped in lateral view, with a few moderate setae on outer lateral surface (Figure
(characters in common with D. puberula sp. n. not repeated). Adult ♂ and ♀. Head (Figures
Thorax (Figure
Legs (Figure
Male terminalia (Figure
Female terminalia (Figure
Measurements (in mm): BL = 2.72 in holotype (range in 5♂/4♀ paratypes: 1.99–2.36/2.08–3.00), ThL = 1.22 (0.90–0.98/0.82–1.19), WL = 2.30 (1.74–1.82/1.62–2.33), WW = 1.26 (0.88–0.96/0.82–1.24).
Indices. FW/HW = 0.58 (range in 5♂ and 4♀, or less if noted, paratypes: 0.55–0.58), ch/o = 0.09 (0.08–0.13), prorb = 0.68 (5♂, 3♀: 0.50–0.80), rcorb = 0.36 (0.27–0.45), vb = 0.40 (0.32–0.55), orbito = 0.70 (0.50–0.67), dcl = 0.67 (3♂, 4♀: 0.58–0.68), sctl = 0.96 (0.89–1.06), sterno = 0.47 (0.54–0.68), dcp = 0.49 (0.42–0.52), sctlp = 0.83 (0.69–1.00), C = 1.66 (1.48–1.87), 4c = 1.46 (1.20–1.61), 4v = 1.88 (1.62–2.00), 5x = 1.36 (1.45–1.90), ac = 2.92 (2.90–3.44), M = 0.73 (0.68–0.83), C3F = 0.61 (0.51–0.62).
China (Hainan).
Pertaining to the type locality, Hainan (formerly known as “Qiongzhou”).
Holotype ♂ (#00177): MALAYSIA: Ulu Senagang, Crocker Range, Sabah, 18.x.1999, MJ Toda (
Paratypes. MALAYSIA: 2♀ (#03905, #03906), Park Headquarters, Mt. Kinabalu, Sabah, 11.iii.2008, MJ Toda (
Palpus broad, flat, with a few moderate setae on outer lateral surface (Figure
(characters in common with D. puberula sp. n. not repeated). Adult ♂ and ♀. Head Figures
Thorax (Figure
Legs (Figure
Male terminalia (Figure
Female terminalia (Figure
Measurements (in mm): BL = 2.34 in holotype (range in 2♀ paratypes: 2.20–2.60), ThL = 1.18 (1.18–1.40), WL = 2.00 (2.35–2.63), WW = 1.06 (1.28–1.37).
Indices. FW/HW = 0.56 (range in 2♀, or less if noted, paratypes: 0.56–0.63), ch/o = 0.09 (0.12), prorb = 0.68 (1♀: 0.67), rcorb = 0.34 (1♀: 0.31), vb = n/a (1♀: 0.50), orbito = 0.65 (0.40–0.55), dcl = 0.65 (0.71–0.75), sctl = 1.00 (1♀: 1.04), sterno = 0.58 (0.63–0.71), dcp = 0.44 (0.43), sctlp = 0.72 (0.73–0.85), C = 1.72 (1.76–1.78), 4c = 1.25 (1.27–1.28), 4v = 1.61 (1.59–1.67), 5x = 1.33 (1.33–1.44), ac = 3.56 (2.93–3.04), M = 0.63 (0.67–0.72), C3F = 0.56 (0.54–0.58).
Malaysia (Sabah).
Referring to the straight aedeagus in the new species.
The paratype female specimens #03905 and #03906 were identified as conspecific with the holotype male specimen #00177 (DNA sequence data unavailable), based on close morphological affinity between them. This species can also be distinguished from the other Dudaica species by oviscapt valve with 17–19 marginal ovisensilla (Figure
Holotype ♂ (#00430): CHINA: Hesong, Xiding, Menghai, Yunnan, ca. 1,900 m a.s.l., 7.iv.2011, JJ Gao (
Palpus short, club-shaped, medio- to baso-laterally dark grayish brown, with one prominent seta apically and several long setae ventrally (Figure
(characters in common with D. orthophallata sp. n. not repeated). Adult ♂. Head (Figures
Thorax (Figure
Wing (Figure
Legs (Figure
Abdomen (Figure
Male terminalia (Figure
Measurements (in mm): BL = 2.33 in holotype, ThL = 1.04, WL = 2.35, WW = 1.16.
Indices. FW/HW = 0.60, ch/o = 0.08, prorb = n/a, rcorb = n/a, vb = n/a, orbito = 0.78, dcl = n/a, sctl = n/a, sterno = 0.62, dcp = 0.65, sctlp = 0.94, C = 2.09, 4c = 1.21, 4v = 1.95, 5x = 1.24, ac = 2.83, M = 0.64, C3F = 0.53.
China (Yunnan).
Referring to the morphological difference from the other species in the subgenus Dudaica.
This species is the most different in morphology from the other Dudaica species, such as the shape of palpus and parameres, apical prominent seta on palpus, and number of sensilla on parameres. Those characters are also seen in many other species than Dudaica, suggesting the plesiomorphic states of these characters.
We thank Dr. Shin-ichi Tanabe, Dr. Maklarin B. Lakim and Dr. Maryati Bte Mohamed for their help in field works and Economic Planning Unit of Malaysian Government for research permissions (UPE:40/200/19 SJ. 732 and UPE: 40/200/19 SJ.1194 and 1195) for the study in Sabah, Malaysia, and Prof. E. S. Ratna of Bogor Agricultural University for her help in field works in West Kalimantan, Indonesia. We also thank the Xishuangbanna Tropical Botanical Garden, CAS, the Xishuangbanna National Nature Reserve, and the Jianfengling National Nature Reserve for permitting us to collect flies in the Garden/Reserves. This work was supported by NSFC (Nos 31160429, 31572238), JSPS KAKENHI Grants Numbers 15255006, 21570085, 24370033 and the 21st Century Center of Excellence Program (E-01) of the Ministry of Education, Culture, Sports, Science and Technology, Japan.