Research Article |
Corresponding author: Ho-Yeon Han ( hyhan@yonsei.ac.kr ) Academic editor: Marc De Meyer
© 2019 Ho-Yeon Han, Kyung-Eui Ro.
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:
Han H-Y, Ro K-E (2019) DNA barcoding reveals a species group of the genus Campiglossa (Diptera, Tephritidae, Tephritinae) with recognition of a new species from East Asia and previously unknown females of Campiglossa coei (Hardy). ZooKeys 899: 1-36. https://doi.org/10.3897/zookeys.899.46779
|
While analyzing DNA barcodes of all the Korean and some East Asian tephritid species in conjunction with the barcode sequences available from BOLD Systems (www.boldsystems.org), the large and taxonomically enigmatic genus Campiglossa was recovered as a monophyletic clade, together with the genera Dioxyna and Homoeotricha, which are here synonymized for that reason. Ten major lineages are also recognized within the Campiglossa clade: producta group, loewiana group, sororcula group, irrorata group, achyrophori group, difficilis group, luxorientis group, magniceps group, arisanica group, and misella group. Here, more detailed taxonomic accounts are provided for the misella group, including four DNA analysis-recovered members: C. coei, C. misella, C. paramelaena sp. nov., and C. melaena. A single morphological synapomorphy is proposed for this species group: the presence of a large mid-anterior dark wing marking in males with associated structural modification (more apically positioned crossvein R-M than in females). Based on the morphological characteristics, two presumptive members that are only known from male specimens are further recognized: C. pishanica and C. propria from China. A full description of C. paramelaena sp. nov., and a redescription of C. coei, for which only males were previously known, are provided. For all the included species, a taxonomic key, diagnoses, and photographs to aid their accurate identification are given. Finally, C. favillacea is synonymized with C. coei and C. roscida with C. misella, and C. coei and C. pishanica resurrected from the synonymy of C. misella.
Campiglossa, Dioxyna, Homoeotricha, misella group, Tephritini
Tephritidae is a relatively recently diverged fly family that might have arisen around the Late Eocene (~36 mya;
The genus Campiglossa Rondani, 1870, is one of those species-rich genera, and is estimated to have approximately 200 described species (
Due to their high intra-specific variation, low inter-specific variation, sexual dimorphism and seasonal variation, systematic investigation of Campiglossa is considered very difficult (A. Freidberg, V. Korneyev, S. Masahiro, B. Merz, pers. comm.). Examination of their male and female postabdominal structure has been somewhat helpful for defining species and species groups (
In the process of analyzing DNA barcodes of all the Korean and some East Asian tephritid species in conjunction with the barcode sequences available from BOLD Systems (www.boldsystems.org), we recovered the genus Campiglossa as a monophyletic clade together with the genera Dioxyna and Homoeotricha. We also recognized ten major lineages within the Campiglossa clade, each of which can be regarded as a monophyletic species group. In this study, we provide more detailed taxonomic accounts for the misella group, including four DNA analysis-recovered members: C. coei (Hardy, 1964), C. misella (Loew, 1869), C. paramelaena sp. nov., and C. melaena (Hering, 1941). Based on the morphological characteristics, we further recognize two presumptive members that are only known from male specimens: C. pishanica (Wang, 1996) and C. propria (Chen, 1938) from China. We provide a full description of C. paramelaena sp. nov., and a redescription of C. coei, for which only males were previously known. For all included species, we provide a taxonomic key, diagnoses, and photographs to aid their accurate identification.
The terminology and morphological interpretations used in this study follow the glossary of
The molecular methods follow
The collection and voucher information for the Campiglossa flies sequenced for the DNA barcoding analysis. The status of the voucher specimens and the GenBank accession numbers are indicated in parentheses.
C. absinthii (Fabricius, 1805) | 1. ♂, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 4.VIII.2005, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915027; GenBank Acc. Nr. MN445522). |
2. ♀, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 24.VII.2005, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915028; GenBank Acc. Nr. MN445523). | |
3. ♀, RUSSIA: Primorsky-Krai, Khasansky-District, Barabash, 43°10'46.9"N, 131°28'20.0"E, 22.VI.2008, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201102; GenBank Acc. Nr. MN445524). | |
C. achyrophori (Loew, 1869) | 1. ♀, SWITZERLAND: Valais 1787–2041 m, Pointe de Bellevue, Morgins, 28.VII.2004, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW140201037; GenBank Acc. Nr. MN445525). |
C. albiceps (Loew, 1873) | 1. ♂, USA: North Carolina, Haywood Co, Great Smoky Mountains National Park, in meadow 250 m N of house at Purchase Knob, 1444 m (both wings glued on a rectangular card; YSUW090915005; GenBank Acc. Nr. MN445526). |
C. bidentis (Robineau-Desvoidy, 1830), comb. nov. from Dioxyna | 2. ♂, KOREA: Gangwondo, Jeongseon, Nammyeon, Mt. Mindungsan, from Yupyeongri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 16.VII.2005, H.-Y. Han et al. (specimen with the abdomen detached; YSUW130901095; GenBank Acc. Nr. MN445527). |
3. ♀, KOREA: Gyeongsangbuk-do, Bonghwa Myeongho-myeon, Mt. Cheongnyangsan, 29.IX.2007, Coll. H.-S. Lee et al., ex Bidens biternata (Lour.) flower, em. 3–12.X.2007 (specimen with the abdomen detached; YSUW130901096; GenBank Acc. Nr. MN445528). | |
C. coei (Hardy, 1964) | 1. ♂, CHINA: Yunnan, Mengsong, Manlvcunhanzudazhai, small hilltop, 22°07'44.0"N, 100°28'51.7"E, 1690 m, 12.VII.2011, H.-Y. Han & S.-W. Suk (specimen with the abdomen detached; YSUW 130901058; GenBank Acc. Nr. MN445530). |
2. ♀, CHINA: Yunnan, Mengsong, Bengangxizhai, in forest, 22°10'34.5"N, 100°35'06.8"E, 1725 m, 11.VII.2011, H.-Y. Han & S.-W. Suk (specimen with the abdomen detached; YSUW 130901059; GenBank Acc. Nr. MN445531). | |
3. ♂, CHINA: Yunnan, Mengsong, Manlvcunhanzudazhai, small hilltop, 22°07'44.0"N, 100°28'51.7"E, 1690 m, 12.VII.2011, H.-Y. Han & S.-W. Suk (specimen with the abdomen detached; YSUW YSUW140201034; GenBank Acc. Nr. MN445532). | |
4. ♀, CHINA: Yunnan, Mengsong, Manlvcunhanzudazhai, small hilltop, 22°07'44.0"N, 100°28'51.7"E, 1690 m, 12.VII.2011, H.-Y. Han & S.-W. Suk (specimen with the abdomen detached; YSUW 140201035 6; GenBank Acc. Nr. MN445533). | |
C. deserta (Hering, 1939) | 1. ♂, KOREA: Gangwon-do, Pyeongchang-gun, Doam-myeon, Hoenggye-ri, Daegwallyeong Samyang pasture, col. 7.X.2004, em. 1–21.VI.2005, ex Aster sp., flower, H.-Y. Han & H.-W. Byun (both wings glued on a rectangular card; YSUW090915029; GenBank Acc. Nr. MN445534). |
2. ♀, KOREA: Gangwon-do, Jeongseon-gun, Gohan-eup, Mt. Hambaeksan, Recreation forest to Manhang-jae, col. 10.X.2003, em. 24–31.V.2004, ex Aster ciliosus Kitamura ?, flower, H.-Y. Han & K.-E. Ro (both wings glued on a rectangular card; YSUW090915030; GenBank Acc. Nr. MN445535). | |
3. ♀, RUSSIA: Primorsky-Krai, Nadezhdinsky-District, Vol’no-Nadezhdinskoye, 43°22'31.6"N, 132°01'43.1"E, 22.VI.2008, Coll. H.-Y. Han & H.-S. Lee (specimen with the abdomen detached; YSUW140201103; GenBank Acc. Nr. MN445536). | |
1. ♀, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, col. 6.X.2001, em. 24–26.X.2001, ex Lactuca indica var. laciniata flower, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW08100129; GenBank Acc. Nr. MN445537). | |
2. ♀, KOREA: Jeju-do, Jeju-si, Aewol-eup, along rt 1117, col 19.X.2005, em. 23–31.X.2005, ex Lactuca indica var. laciniata flower, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW08100130; GenBank Acc. Nr. MN445538). | |
C. difficilis (Hendel, 1927) | 1. ♂, SWITZERLAND: Valais 1689–1950 m, Portes du Soleil, Morgins, 27.VII.2004, H.-Y. Han & K.-E. Ro. (specimen with the abdomen detached; YSUW140201038; GenBank Acc. Nr. MN445539). |
C. guttella (Rondani, 1870) | 1. ♀, SWITZERLAND: Valais 1787–2041 m, Pointe de Bellevue, Morgins, 28.VII.2004, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW140201039; GenBank Acc. Nr. MN445540). |
C. hirayamae (Matsumura, 1916) | 1. ♀, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 24.VI.2005, Han et al. (both wings glued on a rectangular card; YSUW06010914; GenBank Acc. Nr. MN445541). |
2. ♂, KOREA: Gangwon-do, Pyeongchang-gun, Yongpyeon-myeon, S. Valley of Mt. Gyebangsan, 3.X.2003, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW08100131; GenBank Acc. Nr. MN445542). | |
3. ♀, KOREA: Gyeongsangbuk-do, Yeongju-si, Sunheung-myeon, Mt. Sobaeksan, Choamsa to Gukmangbong (1421 m), 27.V.2005, H.-W. Byun (both wings glued on a rectangular card; YSUW08100132; GenBank Acc. Nr. MN445543). | |
C. loewiana (Hendel, 1927) | 1. ♀, MONGOLIA: Tuv Prov., Tusgalt Valley, Forestry Research-Training Center, Ntn. Univ. of Mongolia, 48°15'37"N 106°51'11"E, 1277 m, 4.VII.2013, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201075; GenBank Acc. Nr. MN445544). |
2. ♂, MONGOLIA: Tuv Prov., Tusgalt Valley, Forestry Research-Training Center, Ntn. Univ. of Mongolia, 48°15'37"N 106°51'11"E, 1277 m, 4.VII.2013, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201076; GenBank Acc. Nr. MN445545). | |
3. ♂, MONGOLIA: Tuv Prov., Tusgalt Valley, Forestry Research-Training Center, Ntn. Univ. of Mongolia, 48°15'23"N, 106°50'23"E, 1522 m, 5.VII.2013, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201081; GenBank Acc. Nr. MN445546). | |
C. longipennis Shiraki, 1933, comb. nov. from Homoeotricha | 1. ♂, RUSSIA: Sakhalin, Yuzhno-Sakhalinsk Vestochka, 46°51'58.3"N, 142°50'54.9"E, 18.VII.2008, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW090915062; GenBank Acc. Nr. MN445547). |
C. luxorientis (Hering, 1940) | 1. ♀, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 29.VIII.2005, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915035; GenBank Acc. Nr. MN445548). |
2. ♂, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 29.VIII.2005, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915036; GenBank Acc. Nr. MN445549). | |
C. melaena (Hering, 1941) | 1. ♂, RUSSIA: Primorsky-Krai, Khasansky-District, Barabash, 43°10'46.9"N, 131°28'20.0"E, 22.VI.2008, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201105; GenBank Acc. Nr. MN445560). |
2. ♂, RUSSIA: Primorsky-Krai, Nadezhdinsky-District, Vol’no-Nadezhdinskoye, N43°22'31.6”, E132°01'43.1”, 22.VI.2008, H.-Y. Han & H.-S. Lee (specimen with the abdomen detached; YSUW140201106; GenBank Acc. Nr. MN445561). | |
C. melanochroa (Hering, 1941) | 1. ♀, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119 m peak, 37°16'15"N, 128°46'30"E, col. 6.X.2001, em. 26–30.X.2001, ex Aster ageratoides Turcz. flower, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915039; GenBank Acc. Nr. MN445554). |
2. ♂, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeong-ri to 1,119m peak, 37°16'15"N, 128°46'30"E, col. 25.IX.2003, em. 13–20.IX.2003, ex Aster tataricus L. flower, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915040; GenBank Acc. Nr. MN445555). | |
C. messalina (Hering, 1937) | A1. ♂, KOREA: Gangwon-do, Pyeongchang-gun, Yongpyeon-myeon, S. Valley of Mt. Gyebangsan, 5.VIII.2005, H.-Y. Han & H.-S. Lee (both wings glued on a rectangular card; YSUW08100133; GenBank Acc. Nr. MN445550). |
A2. ♀, KOREA: Gangwon-do, Jeongseon-gun, Gohan-eup, Mt.Hambaeksan, Recreation Forest to Manhang-jae, col. 10.X.2003, em 3–6.V.2004 ex Artemisia sp. flower, H.-Y. Han & K.-E. Ro (both wings glued on a rectangular card; YSUW08100134; GenBank Acc. Nr. MN445551). | |
B1. ♂, KOREA: Gangwon-do, Jeongseon-gun, Gohan-eup, Mt.Hambaeksan, Recreation forest to Manhang-jae, col.10.X.2003, em. 3–6.V.2004, ex Artemisia sp. flower, H.-Y. Han & K.-E. Ro (both wings glued on a rectangular card; YSUW090915037; GenBank Acc. Nr. MN445552). | |
B2. ♀, KOREA: Gangwondo, Jeongseon, Nammyeon, Mt. Mindungsan, from Yupyeongri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 29.VIII.2005, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915038; GenBank Acc. Nr. MN445553). | |
C. misella (Loew, 1869) | 1. ♀, HUNGARY: Bdaors, Odvas hg., 18.VI.1991, Merz & Adams (both wings glued on a rectangular card; YSUW94082638; GenBank Acc. Nr. MN445556). |
C. misella (Loew, 1869) | 2. ♀, SWITZERLAND: Valais, Leuk-Rotafen, 46°18'59"N, 7°40'18"E, 640 m, 22.VII.2004, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW130901215; GenBank Acc. Nr. MN445557). |
3. ♂, SWITZERLAND: Valais, Leuk-Rotafen, 46°18'59"N, 7°40'18"E, 640 m, 22.VII.2004, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW140201041; GenBank Acc. Nr. MN445558 | |
4. ♀, SWITZERLAND: Valais, Visperterminen-Kreuz, 46°15'17"N, 7°53'52"E, 1500 m, 21.VII.2004, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW140201042; GenBank Acc. Nr. MN445559). | |
C. paramelaena sp. nov. | 1. Holotype ♂, KOREA: Gyeongsangbuk-do, Bonghwa-gun, Myeongho-myeon, Mt. Cheongnyangsan, 36°46'43.6"N, 128°55”30.8’E, 600 m, 30.VI.2007, H.Y. Han et al. (specimen with the abdomen detached; YSUW090915094; GenBank Acc. Nr. MN445564). |
2. Paratype ♀, RUSSIA: Primorsky-Krai: between Chernyatino and Pokrovk, 43°57'32.7"N, 131°32'24.1"E, 55 m, 26.VI.2008, H.Y. Han & H.S. Lee (both wings glued on a rectangular card; YSUW090915019; GenBank Acc. Nr. MN445565). | |
3. Paratype ♂, RUSSIA: Khasansky-District, Kedrovaya Pad, 43°05'09.4"N, 131°35'06.0"E, 22m, 23.VI.2008, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201108; GenBank Acc. Nr. MN445566). Paratype. | |
4. Paratype ♀, RUSSIA: Khasansky-District, Barabash, 43°10'46.9"N 131°28'20.0"E, 61m, 22.VI.2008, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW090915068; GenBank Acc. Nr. MN445567). Paratype. | |
C. producta (Loew, 1844) | 1. ♀, ISRAEL, Golan Heights, Mt. Hermon, 2000 m, 29.V.2000, H.-Y. Han & K.-E. Ro (specimen with the abdomen detached; YSUW130901194; GenBank Acc. Nr. MN445568). |
C. quadriguttata (Hendel, 1927) | 1. ♂, KOREA: Gangwon-do, Jeongseon-gun, Nam-myeon, Mt. Mindungsan, from Yupyeongri to 1,119 m peak, 37°16'15"N, 128°46'30"E, 19.VII.2005, H.-Y. Han et al. (specimen with the abdomen detached; YSUW090915089; GenBank Acc. Nr. MN445569). |
2. ♀, KOREA: Gangwon-do, Pyeongchang-gun, Yongpyeon-myeon, S. Valley of Mt. Gyebangsan, 3.X.2003, H.-Y. Han et al. (specimen with the abdomen detached; YSUW090915090; GenBank Acc. Nr. MN445570). | |
C. sabroskyi (Novak, 1974) | 1. ♂, USA: Utah: Grand Co., La Sal Mt. Warner Lake, 7.IX.1992, A.L. Norrbom, ex flower of Senecio sp. (1♂, 1♀ from same collecting lot; HAN115; GenBank Acc. Nr. MN445529). |
C. shensiana (Chen, 1938) | 1. ♂, KOREA: Gangwon-do, Wonju-si, Gwirae-myeon, Unnam-ri, col. 13.X.2001, em. 2–16.V.2002, ex Chrysanthemum boreale, flower, D.-S. Choi et al. (both wings glued on a rectangular card; YSUW090915041; GenBank Acc. Nr. MN445571). |
2. ♀, KOREA: Gangwondo, Jeongseon, Nammyeon, Mt. Mindungsan, from Yupyeongri to 1,119m peak, 37°16'15"N, 128°46'30"E, col. 6.X.2001, em 9.V.2002, ex Chrysanthemum makinoi, flower, H.-Y. Han et al. (both wings glued on a rectangular card; YSUW090915042; GenBank Acc. Nr. MN445572). | |
3. ♀, KOREA: Gangwon-do, Samcheok-si, Geunsan-dong, Mt. Geunsan, 37°24'28"N, 129°8'9"E, 4.V.2012, H.-Y. Han et al. (specimen with the abdomen detached; YSUW130901200; GenBank Acc. Nr. MN445573). | |
C. sororcula (Wiedemann, 1830), comb. nov. from Dioxyna | 4. ♀, JAPAN: Kyushu, Kagoshima-shi, Hirakawa-cho, Goino, 31°27'53"N 130°30'01"E, 66 m, 10.VII.2010 H.-Y. Han & S.-W. Suk (specimen with the abdomen detached; YSUW130901083; GenBank Acc. Nr. MN445574). |
5. ♀, MALAWI: Nyika National Park, Chelinda, 15kmW, 10°35.036’S 33°44.096’E, 2234 m, 31.XII.2009, H.-Y. Han (specimen with the abdomen detached; YSUW130901145; GenBank Acc. Nr. MN445575). | |
C. sp. near guttella | 1. ♀, MONGOLIA: Tuv Prov., Tusgalt Valley, Forestry Research-Training Center, Ntn. Univ. of Mongolia, 48°15'23"N, 106°50'23"E, 1522 m, 5.VII.2013, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201077; GenBank Acc. Nr. MN445562). |
2. ♂, MONGOLIA: Tuv Prov., Tusgalt Valley, Forestry Research-Training Center, Ntn. Univ. of Mongolia, 48°15'23"N, 106°50'23"E, 1522 m, 5.VII.2013, H.Y. Han & H.S. Lee (specimen with the abdomen detached; YSUW140201082; GenBank Acc. Nr. MN445563). | |
C. spenceri (Hardy, 1973) | 1. ♂, VIETNAM: Lam Dong Prov., Mt. Lang Biang, N of DaLat, 12°02'50.1"N 108°26'26.5"E, 12.XII.2013, H.Y. Han et al. (specimen with the abdomen detached; YSUW140201110; GenBank Acc. Nr. MN445576). |
Photographs of pinned specimens were captured with a Panasonic (Osaka, Japan) DMC G5 camera with a Panasonic Lumix 45–175 mm lens and a Raynox (Yoshida Inc., Tokyo, Japan) MSN-202 macro conversion lens. The consecutive digital images in different focal planes (usually 50–100 shots per a single figure) were Z-stacked using Helicon Focus software (Helicon Soft, Ltd., Kharkov, Ukraine). Photographs of live specimens (kept in a glass cage) were taken with a Nikon (Tokyo, Japan) D7000 camera with a macro lens and extension tubes. Photographs of postabdominal structures were taken with a Nikon (Tokyo, Japan) D90 camera mounted on an Olympus (Tokyo, Japan) CX41 compound microscope.
Most of the specimens used in this study are deposited in the Division of Biological Science and Technology, Yonsei University, Wonju, Korea (YSUW), and some in the National Institute of Biological Resources, Incheon, Korea (
The genus Campiglossa is a morphologically homogeneous taxon, and their monophyly has been suggested based on at least two possible synapomorphies: the elongated proboscis and the spinulose phallic preglans area (
As a result of our ongoing DNA barcoding study of the family Tephritidae, we assembled a large dataset of 7,223 individuals, 543 species, and 80 genera publicly available from BOLD systems (www.boldsystems.org), as well as our own dataset of 55 individuals and 26 Campiglossa species. The combined dataset contained 7,278 individuals, 543 species and 80 genera. Our simple neighbor-joining analysis recovered a monophyletic cluster of the genera Campiglossa, Dioxyna, and Homoeotricha together (only this portion of the tree is shown in Fig.
The genus Campiglossa portion of the neighbor-joining tree based on the Kimura 2-parameter distances of 7,223 tephritid DNA barcode sequences mostly extracted from BOLD Systems (www.boldsystems.org, as of Jan 2019), including 55 newly obtained Campiglossa sequences (names prefixed with Z). All 211 Campiglossa, Homoeotricha, and Dioxyna (regarded to be the genus Campiglossa, sensu lato, in this study) sequences were recovered as a monophyletic clade in this analysis. Putative species group names (in red) are marked on the respective branches.
We also analyzed a scale-down dataset of 32 species and 76 individuals of the genus Campiglossa as well as four species and ten individuals of the genus Tephritis that is known to be closely related to Campiglossa (
MEGA X analysis produced a maximum-likelihood (ML) phylogram of the 76 selected Campiglossa (ingroup) and ten Tephritis (outgroup) DNA barcode sequences using the general time reversible model. The first number on each branch is the bootstrap support from ML analysis (pb); the second number represents posterior probability (pp) from Bayesian inference (BI). Samples in green letters were extracted from BOLD systems (www.boldsystems.org).
The producta group was originally recognized by
The loewiana group includes ca. 30 Holarctic species that have white frontal setulae, and white postocular and posterior notopleural setae (
The sororcula group was previously known as the genus Dioxyna, which is synonymized here with Campiglossa. Both our analysis, as well as
The irrorata group, sensu stricto. In our dataset, this group is only represented by a single species, C. hirayamae (Matsumura, 1916), which has a peculiar wing pattern, including the pterostigma with two hyaline spots and the wing margin between apices of veins R1 and Cu1 with rather regularly arranged nine or ten round hyaline spots. These characteristics seem to be shared by at least the following four species: C. amurensis Hendel, 1927; C. grandinata (Rondani, 1870); C. irrorata (Fallén, 1814); and C. venusta Dirlbek & Dirlbeková, 1971. In BOLD Systems (boldsystems.org), our identification attempt using a C. hirayamae sequence indeed recovered two closely related species, C. irrorata (1.84–2.00 % barcode distance) and C. grandinata (1.83–2.15 %). The sequences of these two species were not included in our phylogenetic analyses because they were not open for public download. The name, irrorata group, was originally used by
The achyrophori group was originally recognized by
The difficilis group was defined by
The luxorientis group was originally named by
The magniceps group was defined as such by
The arisanica group was previously known as the genus Homoeotricha, which is here synonymized with Campiglossa. Our DNA barcoding analyses recovered C. longipennis Shiraki, 1933, within the Campiglossa clade. This species closely resembles C. arisanica (Shiraki, 1933), which is the type species of the East Asian genus Homoeotricha (the senior author examined the holotype ♀ of C. longipennis and the syntype ♂♀ of C. arisanica in
The misella group is named and reviewed in detail below.
Our DNA barcoding analyses (Figs
Diagnosis. Members of the misella group can be diagnosed as follows, including the remarkable sexually dimorphic wing pattern: Head with paravertical and genal setae whitish. Thorax with both notopleural setae dark; apical scutellar setae at most half as long as basal setae; anepisternum with upper seta strong, dark, but lower seta approx. half as long, whitish; katepisternal seta strong, dark; anepimeral seta strong, whitish. Legs with both mid and hind coxal setae whitish. Male wing (except for some European populations of C. misella that show small sexual wing dimorphism) with large dark mid-anterior marking (roughly elliptic to inverted triangular shape; e.g., Fig.
Distribution. All the recognized species of the misella group are distributed in East Asia including Nepal, China, the Russian Far East, and Korea, but the widespread C. misella extends its distribution to Central Asia and to Europe.
Biology. Campiglossa misella is the only species with known biology.
Remarks. In addition to the large mid-anterior wing marking, the position of the crossvein R-M is more apically placed in males of all three species measured both sexes (male vs. female vein M ratios of C. coei 0.4–0.45 vs. 0.62–0.76; C. misella, 0.26–0.28 vs. 0.4–0.49; C. paramelaena sp. nov., 0.29–0.43 vs. 0.41–0.53). Such a structural modification seems to be associated with the male wing pattern modification of the misella group. We posit that the large mid-anterior dark marking with associated structural modification present only in males is a good candidate for a morphological synapomorphy of this species group. Interestingly, the wing cell r1 of C. propria (Chen, 1938) male is further modified (see the Diagnosis of C. propria and Fig.
Since the females of the misella group do have more typical Campiglossa wing patterns and there are a good number of Campiglossa species currently known only by females, there might be some more species of the misella group not recognized in this study. A further survey of East Asian Campiglossa species in conjunction with DNA barcoding analyses is required.
1 | Legs entirely yellow-brown (Fig. |
2 |
– | Legs dark (Fig. |
3 |
2 | Width of cell r1 measured on axis of crossvein R-M as wide as or slightly wider than cell r2+3 (as in Fig. |
C. coei |
– | Width of cell r1 measured on axis of crossvein R-M approx. twice as wide as cell r2+3 (Fig. |
C. propria * ♂ |
3 | Scutum dark brown (Fig. |
C. melaena |
– | Scutum ash-grey (Fig. |
4 |
4 | Cell br posterior to fork of vein Rs hyaline (Fig. |
C. paramelaena sp. nov. |
– | Cell br posterior to fork of vein Rs with dark area (Fig. |
5 |
5 | Cell r1 posterior to pterostigma with two hyaline spots (Fig. |
C. pishanica * ♂ |
– | Cell r1 posterior to pterostigma with three hyaline spots (Fig. |
C. misella |
Tephritis coei
Hardy, 1964: 164 (Type-locality: NEPAL, Taplejung Dist., N of Sangu, above river bank, ca. 5000 ft, holotype ♂,
Campiglossa coei:
Campiglossa favillacea
Ito, 2011: 29 (Type-locality: NEPAL, Taplejung Dist., Kharu Pokhar, 3,000 m, holotype ♂,
Type series of C. favillacea Ito, 2011 (
This light-colored species can be diagnosed by the following characteristics. Head largely yellow-brown with grey upper occiput. Thorax with scutum entirely matte whitish grey without any outstanding dark spots or stripes; scutellum mostly matte whitish grey but ca. apical 1/3 yellow-brown. Legs entirely yellowish brown without any dark marking; fore femur with six or seven strong, brown postero-ventral setae. Wing with basal area (basal 1/3 anteriorly and basal 1/2 posteriorly) largely hyaline with only few small dark spots, especially cell br with area posterior to fork of vein Rs completely hyaline (Fig.
This species appears similar to C. pishanica (with only males known) but the latter species can be readily separated by the dark femora and more extensive mid-anterior wing marking with pterostigma completely dark (Figs
Genitalia of Campiglossa coei A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G spermatheca.
Body
(Fig.
Head yellow-brown with whitish pruinosity except for dark brown ocellar triangle and grey upper occiput; head ratio 0.78–0.90, frons-head ratio 0.46–0.50, eye ratio 0.71–0.77, gena-eye ratio 0.17–0.23, antenna-head ratio 0.40–0.44, arista-antenna ratio 1.3–1.6; vertex yellow-brown; dark brown inner vertical seta approx. as long as longest diameter of eye; outer vertical seta white, 0.4× inner vertical seta; post ocellar seta white, 0.4× post ocellar seta; paravertical seta white, 0.7–0.8× post ocellar seta; ocellar seta dark brown, 3.3–4.0× ocellar triangle length; frons almost bare with frontal angle ca. 115 degree; with two dark brown frontal setae; white posterior orbital seta 0.6× dark brown anterior orbital seta; scape and pedicel yellow-brown with short brown setulae; first flagellomere 1.4–1.8× pedicel length, apically rounded, yellow-brown; arista entirely short pubescent, brown except yellow-brown basal area; face yellow-brown without distinct antennal groove; parafacial 0.4× as wide as first flagellomere; facial ridge with fine pale yellow setulae; gena with strong white genal seta and relatively long white setulae; postgena swollen with strong white postgenal seta and relatively long white setulae; postocular setae with two thick white setulae plus over ten shorter brown setulae, extended 0.5× distance from upper eye margin to lower eye margin; supracervical setae white; mouthparts geniculated with yellow-brown setulose labella; palpus with brown setulae apically and white setulae on remaining area.
Thorax largely dark brown ground color with very heavy whitish pruinosity, generally appearing matte whitish grey; postpronotal lobe with single dark brown seta, yellow-brown in ground color, but appearing similar color as nearby areas due to heavy whitish pruinosity; scutum matte whitish grey with five faint brownish longitudinal bands traceable in clean specimens; two pairs of white scapular setae; acrostical setae widely separated each other, situated midway between levels of intra-alar setae and postsutural supra-alar setae; post-alar setae same level as intra-alar setae; dorsocentral setae same level as or slightly lower than transverse suture; presutural supra-alar setae approximately the same level as anterior notopleural setae; two notopleural setae dark brown with posterior seta0.5× anterior seta; scutellum mostly matte whitish grey but ca. apical 1/3 yellow-brown, slightly convex, almost bare except marginal tiny white setulae; basal scutellar setae more or less parallel, 2.3–3.5× as long as scutellum; apical scutellar setae crossed near apex, 0.9–1.3 as long as scutellum; pleura largely matte whitish grey; proepisternum with 3–5 white setulae; anepisternum matte grey with posterior 2/3 white setulose, with one strong dark brown seta and one half as long white seta ventral to it; katepisternum matte grey with a strong dark brown seta, upper area sparsely with short white setulae and lower area with long white setulae; mediotergite matte grey.
Legs entirely yellow-brown with slight grey pruinosity and brown to dark brown setae and setulae; fore coxa anteriorly with white setulae, posteriorly bare; mid coxa anteriorly with few long white setulae, posteriorly bare; hind coxa with strong white lateral seta, posteriorly largely membranous; front femur with six or seven strong brown posteroventral setae; tibiae and tarsi entirely yellow-brown; midtibial spur dark brown, 1.2–1.4 as long as tibial width.
Wing
(Fig.
Wing dimorphism.
Male (Fig.
Male abdomen.
Preabdomen slightly longer than wide, almost entirely matte pale grey; tergites 2–5 with white setulae, but tergite 5 also with 4–7 dark brown marginal setae; tergites 3–5 each with pair of pale brown submedian spots. Postabdomen (Fig.
Genitalia of Campiglossa misella A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G Spermatheca.
Female abdomen. Preabdomen slightly longer than wide, almost entirely matte grey; tergites 2–6 with white setulae, and tergite 6 especially with 4–7 dark brown marginal setae; tergites 3–6 each with pair of pale brown submedial spots. Postabdomen (Fig.
Nepal, China (Yunnan).
The male wing pattern of C. coei is atypical for the genus Campiglossa (Fig.
Oxyna misella
Loew, 1869: 19 (Type-locality: RUSSIA, Sarepta [Volgograd Region]. Syntype ♂♀,
Tephritis lusoria
Nowicky, 1869: 145 (Type-locality: UKRAINE, “Podolu, Sinkowie”; and Skale [Skala Podilska]. Syntype ♂,
Paroxyna kunlunica
Wang, 1996: 185 (Type-locality: CHINA, Yecheng, Xinjiang. Holotype ♂,
Campiglossa roscida
Ito, 2011: 28 (Type-locality: NEPAL, Taplejung Dist., Walungchung Gola, 3,350 m. Holotype ♀,
Campiglossa misella:
Paroxyna misella:
HUNGARY: Bdaors, Odvas hg., 18.VI.1991, B. Merz and Adams, 1♀ (YSUW). ITALY: Aosta, St. Pierre, M. Torrette, 800–850 m, 22.IV.2003, B. Merz and F. Amiet, 1♂ (YSUW). NEPAL: Taplejung: Walungchung Gola, 3,350 m, 14.VI.1962, T. Yasuda, holotype ♂ of C. roscida (
Males of C. misella usually have distinct sexually dimorphic wing patterns [e.g., Fig.
Europe, Central Asia, China (Xinjian, Shanxi, Sichuan, Xizang, Yunnan), Nepal.
This is the only species of the misella group with host feeding biology known. Interestingly,
We resurrected C. coei and C. pishanica from the synonymy of C. misella by Korneyev (2014). Our study indicates that C. coei is a valid species (Figs
Holotype
♂: KOREA: Gyeongsangbuk-do, Bonghwa-gun, Myeongho-myeon, Mt. Cheongnyangsan, 36°46'43.6"N, 128°55'0.8"E, 600 m, 30.VI.2007, H.Y. Han et al. (
The specific epithet is derived from the closely related species melaena prefixed with para.
This new species can be diagnosed by the following characteristics. Head largely yellowish brown with grey upper occiput. Thorax with scutum entirely ash-grey with five faint brownish longitudinal stripes (Fig.
A–E Campiglossa paramelaena sp. nov. A male, lateral view B male, dorsal view C male wing D female, lateral view E female, dorsal view F female, wing G–K C. melaena G male, lateral view H male, dorsal view I male, wing J holotype male, wing K female, wing J, K Reproduced from
Campiglossa paramelaena sp. nov., appears similar to C. misella but the former species can be readily separated by the almost hyaline basal area of the wing, and the area posterior to the fork of vein Rs in particular is completely hyaline while the latter species has a distinctly dark spot on that area (Fig.
Body
(Fig.
Head yellow-brown with whitish pruinosity except for dark grey ocellar triangle and upper occiput; head ratio 0.85–0.92, frons-head ratio 0.47–0.53, eye ratio 0.75–0.83, gena to eye ratio 0.17–0.22, antenna-head ratio 0.41–0.46, arista-antenna ratio 1.3–1.7; vertex yellow-brown; dark brown inner vertical seta approximately as long as longest diameter of eye; outer vertical seta white, 0.4× inner vertical seta; post ocellar seta white, 0.3–0.4× post ocellar seta; paravertical seta white, 0.7–0.9× post ocellar seta; ocellar seta dark brown, 3.0–3.5× ocellar triangle length; frons almost bare with frontal angle 110–115 degree; with two dark brown frontal setae; white posterior orbital seta 0.6–0.8× dark brown anterior orbital seta; scape and pedicel yellow-brown with short dark brown setulae; first flagellomere 1.5–2.1× pedicel length, apically rounded, yellow-brown but with greyish tinge in some individuals; arista entirely short pubescent, dark brown except yellow-brown basal area; face yellow-brown without distinct antennal groove; parafacial 0.4–0.5× as wide as first flagellomere; facial ridge with fine pale yellow setulae; gena with strong white genal seta and relatively long white setulae; postgena swollen with strong white postgenal seta and relatively long white setulae; postocular setae with two thick white setulae plus ten or more shorter dark brown setulae, extended 0.6× distance from upper eye margin to lower eye margin; supracervical setae white; mouthparts geniculated with labella yellow-brown setulose; palpus with brown setulae apically, white setulae on remaining area.
Thorax largely dark brown in ground color with heavy whitish grey pruinosity, generally appearing ash-grey; postpronotal lobe with single dark brown seta, yellow-brown in ground color, therefore, appearing paler than nearby areas; scutum ash-grey with five faint brownish longitudinal bands traceable in clean specimens; two pairs of white scapular setae; acrostical setae widely separated, situated midway between levels of intra-alar setae and postsutural supra-alar setae; post-alar setae same level as intra-alar setae; dorsocentral setae approximately same level as transverse suture; presutural supra-alar setae slightly above level of anterior notopleural setae; two notopleural setae dark brown with posterior seta 0.5× anterior seta; bases of acrostichal, dorsocentral, intra-alar, basal scutellar setae dark brown; scutellum mostly ash-grey but ca. apical 1/5 yellow-brown, slightly convex, almost bare except marginal tiny white setulae; basal scutellar setae more or less parallel, 3.1–3.6× (in males) and 2.4–3.0× (in females) as long as scutellum; apical scutellar setae crossed near apex, 1.1–1.4× (in males) and 0.9–1.1× (in females) as long as scutellum; pleura largely ash-grey; proepisternum with 3–5 white setulae; anepisternum ash-grey with posterior 2/3 white setulose, with single strong dark brown seta and one seta half as long and white ventral to it; katepisternum ash-grey with a strong seta, upper area sparsely covered with short white setulae and lower area with long white setulae; mediotergite ash-grey. Legs yellow-brown ground color with ash-grey pattern and brown to dark brown setae and setulae; fore coxa yellow-brown with posterobasal 1/3 grey, anteriorly with white setulae, posteriorly bare; midcoxa yellow-brown, anteriorly with few long white setulae, posteriorly bare; hind coxa greyish yellow-brown, with white lateral seta, posteriorly largely membranous; femora largely ash-grey except yellow-brown apices; tibiae and tarsi entirely yellow-brown; midtibial spur dark brown, 1.0–1.3× as long as wide.
Wing
(Fig.
Wing dimorphism.
Male (Fig.
Male abdomen.
Preabdomen slightly longer than wide, almost entirely ash-grey; tergites 2–5 with white setulae, but tergite 5 also with 5–7 dark brown marginal setae; tergites 3–5 each with pair of brown submedian spots. Postabdomen (Fig.
Genitalia of Campiglossa paramelaena sp. nov. A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G spermatheca.
Female abdomen.
Preabdomen slightly longer than wide, almost entirely ash-grey; tergites 2–6 with white setulae, tergite 6 especially with dark brown marginal setae; tergites 3–6 each with pair of brown submedial spots. Postabdomen (Fig.
Korea, the Russian Far East.
Individuals of C. paramelaena sp. nov., have DNA barcodes (Figs
Sinotephritis melaena
Hering, 1941: 27 (Type-locality: China: Manchuria, Sjaolin. Holotype ♂, allotype ♀,
Campiglossa melaena:
. Russia: Primorsky-Krai: Khasansky-District, Barabash, 43°10'46.9"N, 131°28'20.0"E, 61m, 22.VI.2008, H.Y. Han and H.S. Lee, 3♂ (YSUW); Nadezhdinsky-District, Vol’no-Nadezhdinskoye, grassland near restaurant, 43°22'31.6"N, 132°01'43.1"E, 61m, 22.VI.2008, H.Y. Han and H.S. Lee, 3♂ (YSUW).
This is the darkest species of the misella group, showing the least wing dimorphism (Fig.
North east China, the Russian Far East.
The following two species are tentatively placed in the misella group based only on the superficial male characters available from the original and subsequent descriptions. In the future, their memberships should be confirmed by the female characters as well as a DNA barcoding analysis.
Tephritis pishanica
Wang, 1996: 188 (Type-locality: CHINA, Xinjian Province, Pishan, holotype ♂, paratype 2♂,
This is an interesting species showing the characteristics of both C. coei and C. misella. The only known C. pishanica male wing pattern is very similar to that of C. coei (Fig.
A Holotype male of Campiglossa favillacea Ito, 2011 (new synonym of C. coei,
Only three males (the type series) known from China (Xinjian).
Sinotephritis propria
Chen, 1938: 149 (Type-locality: China, s.e. Gansu, Mi-tching-ngai, holotype ♂,
Campiglossa propria:
We are not sure if this species actually belongs to the misella group, because the only known male (holotype) does not show close similarity to any known member of the group except for its large mid-anterior dark wing marking (Fig.
Only known from the type locality (Gansu, China).
The genus Campiglossa currently includes ca. 200 similar looking species with their larvae usually feeding in the capitula of Asteraceae plants (
The male and female flies of most Campiglossa species seem to stay close to their host plants (
Capitula infesting tephritids including Campiglossa are the easiest tephritids to rear. Mature flower heads of Asteraceae plants should be collected and kept in mesh bags (similar to insect net bags). These bags should be stored in a sheltered area which maintains an approximate outdoor temperature, and examined for emerging flies. Once the plant materials dry, proper moisture should be maintained by misting with sterile water once or twice a week. In Korea, fall-collected flower heads, after harvesting fall-emerged tephritids, are kept in a 4 °C refrigerator between early December and early April. Overwintered flower heads, if infested by overwintering immature tephritids, usually yield adult flies until early June. Emerged flies should always be kept alive for a few days for hardening and coloring of their cuticles. Each puparium may be separated and kept in a gelatin capsule to match the emerged adult and its own puparium (see
Due to the postmortem deterioration of the Campiglossa specimens, it is desirable to take high resolution photographs while they are still alive or just after euthanasia. Most of the figures presented in this study have been made in this manner. A collapsible glass cage and a simple hand-made macro-photography stacking station are useful for taking such pictures during a multi-day collecting trip (Fig.
As demonstrated in this study, DNA barcode sequences of the genus Campiglossa form a strong monophyletic clade when analyzed phylogenetically. Therefore, any tephritids of uncertain identity clustering together within this clade should be regarded as Campiglossa. For this reason, we synonymize the genera Dioxyna and Homoeotricha, which were clearly placed within this clade (Figs
This work could not have been possible without the generous assistance from Drs. Bernhard Merz and Valery Korneyev, who are the world authorities of the Campiglossa taxonomy. Dr. Bernhard Merz provided many reference specimens of a number of European Campiglossa species. He also kindly identified many Korean and exotic Campiglossa species in our possession when he visited our laboratory in 2005. Dr. Valery Korneyev kindly reviewed the initial draft of our manuscript and made some important comments that helped us improve our manuscript. He also permitted us to use his published illustrations of C. melaena wings, and provided high quality scanned files of the original drawings. We are thankful to Dr. Xiaolin Chen, who invited the senior author for a collecting trip in 2011 in Yunnan Province, China, where we found a series of C. coei used in this study. We thank Dr. Xing-Jian Wang for permitting us to use his published wing pictures of C. pishanica and C. propria. We also thank professor Minoru Ishii for allowing us to examine the late Dr. Syusiro Ito’s tephritid collection housed in the Osaka Prefecture University (