Contributions to Zoology 89 (2020) 210-244 CTOZ brill.com/ctoz
Phylogenetic relationships of the bacchine flower flies (Diptera: Syrphidae) based on molecular characters, with a description of a new species of Melanostoma (Schiner, 1860)
Ximo Mengual Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany [email protected]
Abstract
The phylogenetic relationships among the genera of the tribe Bacchini sensu lato (i.e., Syrphinae with simple, unsegmented aedeagus) were inferred using molecular evidence. The mitochondrial protein- coding gene cytochrome c oxidase subunit I (COI) and the nuclear ribosomal 28S and 18S rRNA genes for 54 bacchine taxa were analyzed using Bayesian inference and Maximum Likelihood. Among the analyzed taxa there is a new species of Melanostoma (Schiner) from Cameroon, Melanostoma janeceki Mengual, sp. nov., which is described in full. This new species has a complete metasternum without excavation, a characteristic that is not present in other species of Melanostoma, usually with an excavated meta- sternum, but it is the diagnostic character of the genus Afrostoma Skevington, Thompson & Vockeroth. Based on the phylogenetic placement of Melanostoma janeceki Mengual, sp. nov. the taxonomic status of Afrostoma as a genus is discussed and a new generic classification of Bacchini stat. rev. and Melanosto- mini stat. rev. is presented in the light of the inferred phylogeny. Results did not recover Bacchini sensu lato monophyletic, but into two groups as follows: Melanostoma and related genera, and Baccha, Platy- cheirus and related genera. Pseudoplatychirus van Doesburg is considered junior synonym of Platycheirus Le Peletier & Audinet-Serville, and Afrostoma Skevington, Thompson & Vockeroth as junior synonym of Melanostoma. Pyrophaena Schiner and Eocheilosia Hull comb. nov. are ranked as valid genera, and con- sequently, the genus Platycheirus is divided into four subgenera: Platycheirus (Carposcalis) (Enderlein), Platycheirus (Pachysphyria) (Enderlein), Platycheirus (Platycheirus), and Platycheirus (Tuberculanostoma) (Fluke) comb. nov.
Keywords
Afrostoma – bacchine – Bacchini – melanostomine – Melanostomini – new synonym
© Mengual, 2019 | doi:10.1163/18759866-20191410 This is an open access article distributed under the terms of the CC-BY 4.0 License. Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Introduction Syrphinae needs a revision based on recent molecular studies, where Bacchini and Syr- Commonly known as flower flies or hoverflies, phini were never resolved as monophyletic syrphids (Diptera: Syrphidae) are distributed (Mengual et al., 2008a, 2012, 2015; Mengual & worldwide, absent only in remote oceanic is- Thompson, 2011; Mengual, 2015; Young et al., lands and Antarctica (Thompson & Rotheray, 2016a; Pauli et al., 2018). 1998). While adults feed on pollen and nectar, Classically, Bacchini is defined as Syrphi- larvae have a large array of natural histories nae taxa with the aedeagus of the male genita- (Ureña & Hanson, 2010; Rotheray & Gilbert, lia rather simple, undivided (Vockeroth, 1969, 2011; Pérez-Lachaud et al., 2014; Fleischmann 1992), and face and scutellum entirely black. et al., 2016). Syrphid species have been used as The coloration of the face and scutellum is no bioindicators (Sommaggio, 1999; Tscharn- longer a defining characteristic for Bacchini tke et al., 2005; Ricarte et al., 2011; Sommag- as there are Bacchini species with pale parts gio & Burgio, 2014), and they contribute to of the face (Vockeroth, 1990; Thompson, 1999) very important ecosystem services, such as and Syrphini species with face and scutellum pollination, biological control of pests or de- entirely black (Vockeroth, 1969, 1990; Huo, composition of organic matter (Lardé, 1989; 2014). The tribe Bacchini sensu lato, hereafter Schmidt et al., 2004; Bergh & Short, 2008; Bugg the members of Syrphinae with a simple ae- et al., 2008; Ssymank & Kearns, 2009; Morales deagus, has undergone several modifications & Wolff, 2010; Martínez-Falcón et al., 2012; in its definition, mostly due to the concept of Nelson et al., 2012; Inouye et al., 2015). the genus Baccha Fabricius, 1805, which ended The family is currently divided into four up to an extremely diverse tribe (Vockeroth, subfamilies, namely Microdontinae, Eristali- 1969). Bigot (1883) was the first to use a name nae, Pipizinae, and Syrphinae (Mengual et al., to refer a suprageneric group, Bacchidae. Since 2015), although some authors prefer to con- Williston (1885) separated them into Mela- sider the microdontines as a different family nostominae and Bacchinae, in almost every (Thompson, 1969; Speight, 1987, 2018). Phy- case where the author distinguished between logenetically, Pipizinae and Syrphinae form Melanostomini (also wrongly written as Mela- a clade (Ståhls et al., 2003; Hippa & Ståhls, nostomatini; see Sabrosky, 1999) and Bacchini, 2005; Mengual et al., 2015; Young et al., 2016a) the latter included several genera and species whose members have primarily predatory lar- of Syrphini placed under Baccha (Williston, vae (Rotheray, 1993; Rojo et al., 2003; Downes 1885; Hull, 1949; Wirth et al., 1965; Vockeroth, et al., 2017). However, some Syrphinae spe- 1969; Shatalkin, 1975). Thompson et al. (1976) cies have become phytophagous secondarily made a major rearrangement redefining the (Nishida et al., 2002; Weng & Rotheray, 2008; genus Ocyptamus Macquart, 1834 and placing Reemer & Rotheray, 2009; Zuijen & Nishida, it within Syrphini, which helped in the cur- 2011; Dumbardon-Martial, 2016). Vockeroth rent definition of Bacchini, but the current (1969) divided the subfamily Syrphinae into concept of Bacchini sensu lato is based mostly six tribes, namely Bacchini, Melanostomini, on Vockeroth (1992) and Thompson (1999). Paragini, Toxomerini, Chrysotoxini and Syr- Although the members of Bacchini were phini. Later, Vockeroth (1992) rearranged his accepted without major controversy, generic tribal system with the currently accepted four definitions have been unclear and the system- tribes: Bacchini, Paragini, Toxomerini, and atic status of some supraspecific taxa was un- Syrphini. This current tribal classification of stable (Thompson & Rotheray, 1998; Mengual
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Figure 1 50% majority rule consensus cladogram produced from Bayesian analysis of COI data. Bayesian posterior probabilities are shown at each node.
Outgroups and the Syrphini taxa included in Microdon mutabilis (Linnaeus, 1758) was con- this work were selected based on previous strained as outgroup as all previous phylog- phylogenetic works and current knowledge. enies resolved MicrodontinaeDownloaded from Brill.com09/25/2021 as the sister 11:12:33AM via free access
Locality information Det.: M. S.M. Hewitt. Leg.: IV.2004. Whiborrow, UK: England, Cumbria, Reemer. 199–521, 31.VII.2007. co., Wiechen, A’foort De THE NETHERLANDS: Zuijen. van Zuijen. Det.: M.P. van M.P. Leg.: de l‘Ill, 184 m., 23.VI.2005. Bas-Rhin, Selestat, Foret FRANCE: Dep. Steenis. van Zuijen & J. van M.P. Leg.: Vujić. Det.: A. Vujic. A. Leg.: Samos island, IV.2011. GREECE: Johansson. N. 23.VI.2010. Leg.: Skirö, SWEDEN: Småland, Skärvete, Ståhls. Det.: G. Vujić. Det.: A. Vujić. A. Leg.: 08.V.2007. Agiassos, nr. Lesvos, GREECE: nr Duncan Lake, hilltop Twp., Masham Quebec, Gatineau Co., CANADA: & A. Skevington. J. 21.VII.2001. Leg.: 3591:6777, 05.VII.2007. Leg.: Riikanmaa, KKJ-Y FINLAND: Ka, Joutseno, Ståhls. Det.: G. Steenis. van & J. W. Zuijen & van M.P. RN 652492–9 130725–9, naturreservat, SWEDEN: Ds Skållerud, Ryrs Ståhls. Det.: G. Nils Ryrholm. 29.VI.2006. Leg.: 12°51′S centre, Sachavacayoc Tambopata, rio de Dios, Madre PERU: Smit. T. J. 4.IX.2009. Leg.: 69°22′W, AMG 55 circuit, Lake Shadow St. Clair NP, Lake Tasmani, AUSTRALIA: Steenis. van W. 429–5337, 850 m., 22.I.2006. Leg.: 37°42.5′N 7 km S Escalante, Wash, Alvoy Utah, Garfield Co., USA: Det.: F.C. Parker. M.E. Irwin & F. 1990 m., 29.VI.2002. Leg.: 111°37.8′W, Thompson. Fluke, 1937 Fluke,
(Curran, 1921) (Curran, (Panzer, 1804) (Panzer, (Violovitsh, 1985) (Violovitsh, Bigot, 1884 Bigot, (Strobl, 1909) (Strobl, (Linnaeus, 1758) (Linnaeus, (Say, 1823) (Say, (Zetterstedt, 1843) (Zetterstedt, Loew, 1848 Loew, (Harris, 1780) (Harris, Violovitsh, Violovitsh, 1981 Taxa used in the molecular analyses, including GenBank accession numbers. GenBank accession numbers ofin including GenBank accession numbers. sequences (this study) are used in the molecular analyses, newly generated Taxa boldface
Table 1 Taxa OUTGROUP mutabilis Microdon ERISTALINAE tenur Neoascia ovatus Eumerus PIPIZINAE Claussenia hispanica notabila Cryptopipiza heringi Heringia intensica Neocnemodon quadrimaculata Pipiza certa Pipizella lasiotibialis Trichopsomyia fulvicornis Triglyphus SYRPHINAE obliqua Allograpta
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Locality information 2200 San Antonio, del Cauca. Cali, Cerro Valle Dpto COLOMBIA: X. X. Mengual. Det.: 24.II.2006. Leg.: m., 03°29.137′N 76°33.596′W, Mengual. 2175 m., San Antonio, del Cauca, Cali, Cerro Valle Dpto COLOMBIA: X. Mengual. X. Mengual. Det.: 15.II.2006. Leg.: Oso near 2nd landslide, Mal Trocha Wayqecha, Est. Biol. Cuzco, PERU: 2806 m., 8–11.XII.2011. WP 532, 13°11.07′S 71°35.075′W, 6, aise trap X. Mengual. Det.: & Nolazco. Sutton Steck, Norrbom, Leg.: Jujuy Prov., ARGENTINA: damp wash, in wooded, Malaise trap Las Lanzas, Arroyo 36 km S Jujuy, M.E. Irwin, F.D. Leg.: 1278 m., 27.X–14.XI.2003. 24°27.25′S 65°17.83′W, X. Mengual. Det.: Parker. San Sector Zamora, Cantón Zamora-Chinchipe Prov., ECUADOR: 2600 m., 26.VII–11.VIII.2012, Malaise, 3°59.488′S Páramo, Francisco, X. Mengual. X. Mengual. Det.: Leg.: 79°7.655′W. Thomp F.C. 14.I.2005. Leg.: de la Muerte, Cerro RICA: Páramo, COSTA X. Mengual. son. Det.: 2800 m, 13.I.2005. Estación La Esperanza, Tapantí, RICA: PN COSTA Thompson. Det.: F.C. Det.: G. Jakovlev. J. 2004. Leg.: Trap. Malaise Vesijako, Ta, FINLAND: Ståhls. Ulu Range, Crocker Distr., Sabah (Borneo), Penampang MALAYSIA: 5°51.383′N 116°18.484′E, 1350 m., tree, Dipterocarp Kalanggan, X. Mengual. M. Hauser & S. Gaimari. Det.: 20.X.2011. Leg.: X. X. Mengual. Det.: Leg.: 18.V.2007. Torretes, Ibi, E.B. Alicante, SPAIN: Mengual. ) Meigen, 1822 Meigen, (Williston, (Williston, (Walker, (Walker, Fluke, 1945 Fluke, (Curran, (Curran, ( Walker, 1852 Walker, (Fabricius, 1775) (Fabricius, sp. nov.1 sp. nov.2 sp. Taxa used in the molecular analyses, including GenBank accession numbers. GenBank accession including GenBank accession numbers. used in the molecular analyses, Taxa
Table 1 Taxa catabomba Argentinomyia 1891) longicornis Argentinomyia 1836) luculenta Argentinomyia neotropica Argentinomyia 1937) Argentinomyia Argentinomyia Argentinomyia elongata Baccha maculata Baccha intermedium Chrysotoxum Downloaded from Brill.com09/25/2021 11:12:33AM via free access
(Philippi, ] complex) (Fallén, 1817) (Fallén, [previously [previously (Williston, 1891) (Williston, (De Geer, 1776) (De Geer, (Hull, 1947) (Fallén, 1817) (Fallén, (Nishida, 2003) (Wiedemann, 1830) (Austen, 1893) (Austen, gracilis (Fabricius, 1794) (Fabricius, Loew, 1854 Loew, harrisi sp. ( sp. nov.1 sp. aff. Dasysyrphus albostriatus Dasysyrphus intermedia Didea Eocheilosia (Eocheilosia) Platycheirus conopida Eosalpingogaster 1865) balteatus Episyrphus syrphoides Eriozona corollae Eupeodes centropogonis Fazia Ischiodon aegyptius balboa Leucopodella bigoti Leucopodella gracilis Leucopodella Leucopodella Leucopodella
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-
(cont.) Locality information 530 m., 1.I.2015. Leg.: 10°07′N 69°14′W, El Cercado, VENEZUELA: Lara, X. Mengual. Det.: E. Arcaya. Reserva Las Palomas, Corrg. Manizales, Caldas, Dpto COLOMBIA: Leg.: 18.II.2006, 2200–2500 m., 5°04′N 75°26.2′W. Río Blanco. Natural X. Mengual. Det.: C. Gutierrez. 2200 m., San Antonio, del Cauca, Cali, Cerro Valle Dpto COLOMBIA: X. Mengual. Det.: C. Gutiérrez. 15.II.2006. Leg.: Oso ner 2nd landslide, Mal Trocha Wayqecha, Est. Biol. Cuzco, PERU: 2806 m., 8–11.XII.2011. WP 532, 13°11.07′S 71°35.075′W, 6, aise trap X. Mengual. Det.: & Nolazco. Sutton Steck, Norrbom, Leg.: Ståhls. Det.: G. Ståhls. G. VII.2001. Leg.: Venosta, Val Tirol, South ITALY: Det.: Ståhls. G. Leg.: V.2004. Karkalinniemi, Karislojo, FINLAND: Ab, Ståhls. G. Det.: Ståhls. G. Leg.: Mäntsälän Mustametsä, 10.V.2003. FINLAND: N, Ståhls. G. Ranomafana, to Valbio from road Prov., Fianarantsoa MADAGASCAR: X. Mengual. X. Mengual. Det.: 25.XI.2004. Leg.: region, Sakaroa Ranomafana N.P., Prov., Fianarantsoa MADAGASCAR: X. Mengual. X. Mengual. Det.: 26.XI.2004. Leg.: 5km SE Big Bamenda Highlands, Prov., North-West CAMEROON: 6°5.434′N 10°18.15′E, 2100–2200 m., Babanki, Mendong Buo area, X. Mengual. Det.: S. Janeček. XI.2012. Leg.: 5km SE Big Bamenda Highlands, Prov., North-West CAMEROON: 6°5.434′N 10°18.15′E, 2100–2200 m., Babanki, Mendong Buo area, X. Mengual. Det.: S. Janeček. XI.2012. Leg.:
(Macquart, Hull, 1941 (Linnaeus, 1758) (Linnaeus, Mengual, sp. nov. Mengual, sp. nov. Mengual, sp. Taxa used in the molecular analyses, including GenBank accession numbers. GenBank accession including GenBank accession numbers. used in the molecular analyses, Taxa sp. nov.2 sp. nov.3 sp. nov.4 sp. nov.5 sp.
Table 1 Taxa Leucopodella Leucopodella Leucopodella Leucopodella lucorum (Leucozona) Leucozona ( Linnaeus, 1758) erraticus Megasyrphus lasiophthalma Melangyna (Melangyna) 1843) (Zetterstedt, annulipes Melanostoma 1842) diffusum Melanostoma janeceki Melanostoma janeceki Melanostoma Downloaded from Brill.com09/25/2021 11:12:33AM via free access
CAMEROON: North-West Prov., Bamenda Highlands, 5km SE Big Bamenda Highlands, Prov., North-West CAMEROON: 6°5.434′N 10°18.15′E, 2100–2200 m., Babanki, Mendong Buo area, X. Mengual. Det.: S. Janeček. XI.2012. Leg.: X. Mengual. 2015. Det.: GERMANY: Det.: Ståhls. 15.VIII.2006. Leg G. Lentuankoski, FINLAND: Ok, Kuhmo, Ståhls. G. le Mt Atara, iti – sentier vers Tahiti Tahiti, FRENCH POLYNESIA: 825 m. Leg.: 17°47.375′S 149°14.899′W, 20.IX.2012 (fauchage), X. Mengual. Det.: Ramage. T. 00°14.13′N 34°51.87′E, Forest, Kakamega Prov., Western KENYA: Thompson. R. Copeland. Det.: F.C. Leg.: 30.VIII–5.IX.2000, Malaise trap. (USNM ENT 00036401) region, Talatakely Ranomafana N.P., Prov., Fianarantsoa MADAGASCAR: X. Mengual. X. Mengual. Det.: 18.XI.2004. Leg.: Center. Valbio [Lagoon]. Langowan Tumarates, Sulawesi, INDONESIA: Northern & C. Dien. Det.: M.F. M. Meray, 1°08′N 124°50′E, 13–20.VIII.2004. Leg.: X. Mengual. A. 6733:222, 21.VII.2004. Leg.: Perkko, Mietoinen, FINLAND: Ab, Ståhls. Det.: G. Haaarto. Korenov, mountains, REPUBLIC: Bohemia, PLA Jezerske CZECH L. Mazánek. Det.: 12.VI.2005. Leg.: 15–21.I.2005, Malaise INBioparque, Heredia, RICA: San José, COSTA Thompson. Det.: F.C. trap. Vieja, Carretera Zamora, Cantón Zamora-Chinchipe Prov., ECUADOR: X. Mengual. Leg.: 1, 1500 m., 7.VIII.2012, 3°57.334′S 79°1.633′W. Point Det.: X. Mengual. Vujić. Det.: A. Vujić. A. 2000. Leg.: Alicante, SPAIN: - - (Skev ] Mengual & (Wiedemann, (Linnaeus, 1758) (Linnaeus, Hull, 1941 Mengual, sp. nov. Mengual, sp. (Fabricius, 1794) (Fabricius, (Walker, 1852) (Walker, (Fallén, 1817) (Fallén, (Zetterstedt, 1843) (Zetterstedt, Macquart, 1834 Meigen, 1822 Meigen, Melanostoma janeceki Melanostoma mellinum Melanostoma scalare Melanostoma polynesiotes Melanostoma 2018 Ramage, quadripunctatum Melanostoma Thompson, 2014) [previously & ington quadripunctatum Afrostoma sylvarum Melanostoma univittatum Melanostoma 1824) guttata Meligramma cinctella Meliscaeva funebris Ocyptamus calda Orphnabaccha haemor (Pandasyopthalmus) Paragus rhous
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(cont.) Locality information GERMANY: Nordrhein-Westfalen, NP Eifel, 50°30.484′N 6°15.034′E, Nordrhein-Westfalen, GERMANY: Leg. 533 m., 17.V.2012. Kalterherberg, Aue, Perlenbachtal Gut Heistert, A. Ssymank. Det. Cape, 2500 Eastern Maclear, Nek, Naude’s Woodcliffe, SOUTH AFRICA: Det.: A. Ssymank. Vujić. A. m., 30°43.9′S 28°8.216′E, 09.II.2016. Leg.: El 8, Sector del Ruiz, Villamaría, via al Nevado Caldas, Dpto COLOMBIA: X. Mengual. Det.: C. Gutierrez. 3500 m., 19.II.2006. Leg.: km 8, 2530 m., Villamaría, via La Esperanza Caldas, Dpto COLOMBIA: X. Mengual. Det.: C. Gutierrez. 19.II.2006. Leg.: SWMA, Harrison 6 mi. NNE, Gilbert Baker NE, Sioux Co., USA: Steenis. van W. Det. Steenis. van W. & J. Leg.: 18.IV.2004. Steenis. van Det.: J. Steenis. van J. SWEDEN: 2000. Leg.: Det.: A.V. Barkalov. A.V. VII.2005. Leg.: Ukok, Altai, Plateau RUSSIA: Barkalov Steenis. van Det.: J. Steenis. van J. SWEDEN: 2000. Leg.: Det.: G. Ståhls. G. 76200:35219, 12.VII.2007. Leg.: FINLAND: Li, Ivalo. Ståhls. El 8, Sector del Ruiz, Villamaría, via al Nevado Caldas, Dpto COLOMBIA: X. Mengual. Thompson. Det.: F.C. 3500 m., 19.II.2006. Leg.: X. Mengual. Thompson. Det.: F.C. 2008. Leg.: COLOMBIA:
] -
sp. sp. Pseudo ) [previously ) [previously (Zetterstedt, (Zetterstedt, Vockeroth, 1973 Vockeroth,
] ] Tuberculanostoma ) ) [previously ) [previously Fluke, 1943 Fluke, Taxa used in the molecular analyses, including GenBank accession numbers. GenBank accession including GenBank accession numbers. used in the molecular analyses, Taxa (
Williston, 1887 Williston, 2007 Barkalov, Table 1 Taxa Parasyrphus macularis Parasyrphus 1843) nigrifacies Pelloloma chalconotus (Carposcalis) Platycheirus (Philippi, 1865) spinipes (Carposcalis) Platycheirus 1990 Vockeroth, coerulescens (Pachysphyria) Platycheirus ( albimanus (Platycheirus) Platycheirus 1781) (Fabricius, glupovi (Platycheirus) Platycheirus ( glupovi platychirus nielseni (Platycheirus) Platycheirus 1990 Vockeroth, peltatus (Platycheirus) Platycheirus 1822) (Meigen, (Tuberculanostoma) Platycheirus [previously nov.1 (Tuberculanostoma) Platycheirus antennatum Tuberculanostoma Downloaded from Brill.com09/25/2021 11:12:33AM via free access
) (Strobl, (Strobl, (Forster, 1771) (Forster, (Say, 1823) (Say, (Fabricius, 1798) (Fabricius, (Linnaeus, 1758) (Linnaeus, (Fabricius, 1787) (Fabricius, ) [previously ) [previously Schiner, 1868 Schiner, Sack, 1938 Meigen, 1822 Meigen, Mengual, 2019 ] Fluke, 1945 Fluke, ( (Linnaeus, 1758) (Linnaeus, Fluke, 1943 Fluke, ( Platycheirus (Tuberculanostoma) (Tuberculanostoma) Platycheirus pectinis Tuberculanostoma granditarsa Pyrophaena rosarum Pyrophaena alpina Rohdendorfia bella Rohdendorfia nigra Salpingogaster pyrastri Scaeva ambulans Spazigaster scripta Sphaerophoria claviventris Syrphocheilosia 1910) vitripennis Syrphus fervida Talahua marginatus Toxomerus agrolas (Xanthandrus) Xanthandrus 1849) (Walker, bucephalus (Xanthandrus) Xanthandrus (Wiedemann, 1830)
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Locality information CHINA: Sichuan Prov., Ya’an, Baoxing, Dengchigou, christian church christian church Dengchigou, Baoxing, Ya’an, Sichuan Prov., CHINA: of 30°22.534′N 102°49.733′E, 1600–1800 m., priest Armand David, Det.: Tkoč. M. Růžička, Hájek, J. J. Leg.: 13.VI.2014, sweeping. X. Mengual. Ståhls. Det.: G. & C. Pérez. S. Rojo Leg.: Lesbos island, IV.2001. GREECE: 2200 m., San Antonio, del Cauca, Cali, Cerro Valle Dpto COLOMBIA: X. Mengual. Det.: 24.II.2006. Leg.: 03°29.137′N 76°33.596′W, X. Mengual. AMG 55 Track, Belcher Lake Mt. Field np, Tasmania, AUSTRALIA: Steenis. van W. Zuijen & van M.P. 468–5273, 1100 m., 21.I.2006. Leg.: Steenis. van W. Det.: Det.: S. Rojo. & C. Perez. S. Rojo Leg.: Lesbos island, IV.2001. GREECE:
(Harris, (Harris, Taxa used in the molecular analyses, including GenBank accession numbers. GenBank accession including GenBank accession numbers. used in the molecular analyses, Taxa
Table 1 Taxa Xanthandrus (Xanthandrus) talamaui (Xanthandrus) Xanthandrus 1924) (Meijere, comtus (Xanthandrus) Xanthandrus 1780) (Harris, plaumanni (Xanthandrus) Xanthandrus 1937 Fluke, ruficorne (Xanthandrus) Xanthandrus 1884) (Bigot, pedissequum Xanthogramma 1776) Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Figure 2 Maximum-likelihood tree based on the combined dataset (COI, 28S, and 18S) using Garli v.2.1.17 and the structural alignment for 28S and 18S. Bootstrap support values (above) and Bayesian posterior probabilities (below) are depicted at the nodes (only >50 or >0.5, respectively). BS = Bootstrap support values; PP = Bayesian posterior probabilities. Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Bayesian inference using the Markov Chain model GTR+I+G was selected for 28S and the Monte Carlo algorithm as implemented in preferred model for 18S was TVM+I+G. The MrBayes 3.2.6 (Huelsenbeck & Ronquist, inclusion of the COI sequence of Afrostoma 2001; Ronquist & Huelsenbeck, 2003) was quadripunctatum did not change the model performed using a parallelized version of the selection. software. Data were not divided into parti- Data was analyzed under the recommend- tions following Thompson & Skevington ed models using Garli v.2.1.17 (Zwickl, 2006, (2014). The best model of evolution for the 2011) under Maximum Likelihood (ML) crite- data was determined using jModelTest 2.1.5 rion. The author conducted 24 independent (Darriba et al., 2012) under the Akaike Infor- runs using scorethreshforterm = 0.05 and sig- mation Criterion (AIC). The model chosen nificanttopochange = 0.0001 settings and the was GTR+I+G. Four chains (three hot, one automated stopping criterion, terminating cold) were run simultaneously for 500,000 the search when the ln score remained con- generations, which were sufficient to bring stant for 50,000 consecutive generations. The the convergence (average standard deviation) tree with the highest likelihood was retained to a value <0.009 (Ronquist et al., 2005). Trees and is presented here (fig. 2). Bootstrap sup- were sampled every 100 generations, the tem- port values (BS) were estimated from 500 rep- perature was set to 0.2 and the number of licates using the same independent models in runs to 6. The initial 1,250 trees (25%) were Garli. discarded as burn-in and Bayesian posterior Bayesian inference (BI) using the Markov probabilities (PP) were calculated using a 50% Chain Monte Carlo algorithm as implement- majority-rule consensus tree inferred from the ed in MrBayes 3.2.6 (Huelsenbeck & Ronquist, data (fig. 1). 2001; Ronquist & Huelsenbeck, 2003) was performed using a parallelized version of the Phylogenetic analyses software. Data were divided into the above The Maximum Likelihood analysis and the five partitions and a separate GTR+I+G model Bayesian inference were performed two times for each partition was specified in the analy- each, one without the COI barcode sequence sis where each partition has its own set of of Afrostoma quadripunctatum Skevington & parameters. Priors were applied with default Thompson, 2014, and a second time with the values. Six runs, with four chains each (one inclusion of this short sequence (647 bp) to “cold” chain and three heated chains; temp evaluate the phylogenetic position of this spe- = 0.2), were performed simultaneously for cies even with only a partial sequence of one 20,000,000 generations, which were sufficient of the three sequenced genes. For the analyti- to bring the convergence (average standard cal runs the data set was divided in five par- deviation) to a value <0.007 (Ronquist et al., titions: first codon position of COI, second 2005), sampling trees every 2,500 generations. codon position of COI, third codon position The program Tracer 1.7.1 (Rambaut et al., 2018) of COI, 28S gene and 18S gene. The best evo- was used to check convergence and accept- lutionary model that fit the data was chosen able mixing. The initial 2,000 trees (25%) were each partition using jModelTest 2.1.5 (Dar- discarded as burn-in and Bayesian posterior riba et al., 2012) under the Akaike Information probabilities (PP) were calculated using a Criterion (AIC), as recommended by Posada 50% majority-rule consensus tree inferred and Buckley (2004). The model chosen for from the data (fig. 2). position 1 of COI was GTR+I+G, TIM1+I+G for Maximum Likelihood analysis and Bayes- position 2 and TIM3+I+G for position 3. The ian inference wereDownloaded performed from Brill.com09/25/2021 on the CIPRES 11:12:33AM via free access
Science Gateway (Miller et al., 2010). All trees specimen D227 and A for specimens D226 and were drawn with the aid of FigTree v.1.4.4 D228). (Rambaut, 2018), Adobe Illustrator® and Ado- be Photoshop® CS5. Phylogenetic analyses No topological differences were found when Afrostoma quadripunctatum [= Melanostoma Results quadripunctatum comb. nov. in fig. 2] was excluded from the analysis, and the same re- Barcoding analysis lationships among taxa were recovered with The Bayesian phylogram (fig. 1) based only both dataset, with or without the DNA bar- on the DNA barcode region (the 5′ end of code sequence of Afrostoma quadripuncta- the COI gene) compares favorably with the tum. The ML and BI analyses resulted in high- cladogram from Thompson & Skevington ly similar tree topologies and are summarized (2014, figure 2). The same groupings and the together. The ML tree including the Afros- same inferred relationships were found in toma barcode with the best likelihood score the present analysis with a minor nomencla- (−28846.913085) is presented in fig. 2 with the tural change: Platycheirus concinnus (Snow, PP from the Bayesian analysis and the BS val- 1895) appears as Platycheirus pictipes (Bigot, ues from the ML analysis. 1884) following the synonymy proposed by Although the taxonomic sampling ef- Young et al. (2016b). The novelty in the pres- fort was done to include all the diversity of ent study is the placement of the new spe- the Bacchini sensu lato, it is worth to men- cies without metasternal excavation, Mela- tion the other syrphine genera and pipizines nostoma janeceki sp. nov., as sister-group of were resolved in agreement with previous Afrostoma quadripunctatum [= Melanostoma studies (Mengual et al., 2008a, 2012, 2015; quadripunctatum comb. nov. in fig. 1] with the Mengual, 2015; Young et al., 2016a; Mengual highest Bayesian posterior probability. Note et al., 2018), with members of Pipizinae re- that Afrostoma quadripunctatum appears as solved as sister-group of Syrphinae. The sub- Afrostoma vockerothi in the figure 2 of Thomp- family Syrphinae was resolved monophyletic son & Skevington (2014), and hereafter this with genera grouped into three major clades. species is referred as Afrostoma quadripunc- The first major clade comprises all the Syr- tatum or as Melanostoma quadripunctatum phini genera, including the representatives comb. nov. The group (Melanostoma janeceki of Toxomerini and Paragini. The second ma- sp. nov. + Afrostoma quadripunctatum) has a jor clade comprises the genera Leucopodella PP of 1, and the barcode similarity based on Hull, 1949, Pelloloma Vockeroth, 1973, Talahua uncorrected pairwise distance between Mela- Fluke, 1945, Xanthandrus Verrall, 1901, Argen- nostoma janeceki sp. nov. and Afrostoma quad- tinomyia Lynch Arribalzaga, 1891, Afrostoma ripunctatum is 96.136% with the D226 and and Melanostoma. In this clade, Leucopodella D228 specimens (25 nucleotide differences) species were placed as sister-group of two and 96.059% with the D227 specimen (26 nu- smaller clades: (Pelloloma + (Talahua + Xan- cleotide differences). thandrus)) and (Argentinomyia + Melanos- The DNA barcodes for the three speci- toma), including Afrostoma. It is important mens of Melanostoma janeceki sp. nov. were to point out that the two studied species almost identical. The only difference among without metasternal excavation, i.e., Melanos- the sequences of Melanostoma janeceki sp. toma janeceki sp. nov. and Afrostoma quadri- nov. was an ambiguous call in one base (W for punctatum, were resolvedDownloaded fromas sisterBrill.com09/25/2021 taxa and 11:12:33AM via free access
embedded among the species of the genus Type locality. Cameroon: Northwest Prov- Melanostoma, which all have an excavation ince, Bamenda Highlands, ca. 5 km south- in the metasternum. east of Big Babanki, Mendong Buo area, The third major clade as resolved in the 6.592699°N, 10.189999°E, 2100–2200 m. present study has the representatives of the Bacchini, with Baccha, Platycheirus and all Description the other genera related to Platycheirus, i.e., Male: Head: Face straight with distinct facial Tuberculanostoma, Pyrophaena, Syrphochei- tubercle, entirely black, pale pilose, sparsely losia Stackelberg, 1964, Spazigaster Rondani, grey pruinose except tubercle shiny; lunule 1843, and Rohdendorfia Smirnov, 1924. Within black, shiny; frontal triangle black, sparsely this clade, Baccha was resolved as the sister- grey pruinose, brownish pilose; eye contigu- group of the other genera, Tuberculanostoma ity long, longer than vertical triangle; ocellar was placed embedded in the Platycheirus triangle dark pilose; antenna brown, basofla- radiation, and (Pyrophaena + (Rohdendorfia + gellomere darker apically; arista pilose, with (Syrphocheilosia + Spazigaster))) were re- pile shorter than the arista width at base; oc- solved as a clade sister to Platycheirus, includ- ciput black, silver grey pruinose, white pilose ing Tuberculanostoma. on ventral 1/2, brownish pilose on dorsal 1/2. Thorax: Black except postalar callus slight- Melanostoma janeceki Mengual, sp. nov. ly paler, brownish black; scutum shiny black, sparsely brownish pruinose, with short and Melanostoma sp. in Vlašánková et al. (2017). long yellow pile with some darker pile; post- Figs. 3, 5C–E. pronotum white/grey pruinose, bare; scutel- lum black, yellow pilose with some darker pile Differential diagnosis. Black, straight face with intermixed, subscutellar fringe long, yellow. small, round facial tubercle; black thorax and Pleuron black, pale pilose, sparsely pale pru- abdomen black with terga 3–5 with two lat- inose; calypter yellow, yellow pilose; plumula eral, small yellow maculae on anterior mar- yellow; halter pedicel and capitulum yellow; gin (fig. 3). This species of Melanostoma with posterior spiracular fringes yellow; metaster- a complete metasternum (fig. 5C, D) is only num complete, bare. Scutum and pleuron known from Cameroon, and it is very similar with a golden iridescence. Wing: brownish, to Melanostoma quadripunctatum (Skeving- entirely microtrichose; alula broader than ton & Thompson, 2014) comb. nov., known costal cell. Legs: pro- and mesofemora brown; from Kenya and Uganda (fig. 5A, B). They dif- pro- and mesotrochanter yellow; pro-and me- fer by the COI gene sequence, sterna color- sofemora yellow; pro- and mesotibia yellow ation in females (entirely yellow in M. janeceki with a medial black annulus; protarsi dark, sp. nov.; entirely black with two lateral yellow probasitarsomere yellow basally; apical three maculae on anterior margin of terga 2–5 in mesotarsomeres black, basal two mesotarso- M. quadripunctatum; fig. 5E, F) and by the leg meres yellow; metacoxa black and metatro- coloration: M. janeceki sp. nov. has pro- and chanter yellow; metafemur black with basal mesofemora yellow in both sexes, and meta- 1/4–1/3 yellow and femoral-tibial joints yel- femora black except yellow on basal 1/4–1/3 low; metatibia black with basal 1/4–1/3 yellow; in males or metafemora entirely yellow in fe- metatarsi black. males (fig. 3); and M. quadripunctatum males Abdomen: Parallel-sided, without lateral and females have all femora black with the margin, entirely black excep t for the two lateral, very apex pale (fig. 5A). small yellow maculaeDownloaded on from anterior Brill.com09/25/2021 margin 11:12:33AM of via free access
Figure 3 Melanostoma janeceki Mengual, sp. nov. A, male holotype (ZFMK-DIP-00015940), lateral view; B, female paratype (ZFMK-DIP-00015958), lateral view; C, male holotype, habitus; D, female paratype (ZFMK-DIP-00015958), habitus; E, male holotype, frontal view; F, female paratype (ZFMK- DIP-00015958), frontal view; G, male holotype, labels; H, female paratype (ZFMK-DIP-00015958), labels. Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Figure 4 A–B: Melanostoma sp. from Cameroon (ZFMK-DIP-00015959) with complete metasternum. A, lateral view; B, habitus. C–H: Platycheirus solitarius (van Doesburg, 1955) comb. nov. C, female holotype, lateral view; D, female holotype, habitus; E, female holotype, frontal view; F, female holotype, labels; G,
female paratype, lateral view; H, female paratype, head, lateral view.Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Figure 5 A–B: Melanostoma quadripunctatum (Skevington & Thompson, 2014) comb. nov., male holotype. A, lateral view; B, habitus. C, Melanostoma janeceki Mengual, sp. nov., male holotype, ventral view. D, Melanostoma janeceki Mengual, sp. nov., male paratype (ZFMK-DIP-00015941), detail of metasternum (ms). E, Melanostoma janeceki Mengual, sp. nov., female paratype (ZFMK-DIP-00015957), ventral view. F, Melanostoma quadripunctatum, female (ZFMK-DIP-00015952), ventral view. terga 3–4; tergum 5 has also two lateral, small to male genitalia of M. quadripunctatum yellow maculae but less evident. Terga dull (Thompson & Skevington, 2014, figure 3). black pruinose, with long yellow pile laterally Female: Similar to male except for normal and short black pile medially; sterna black ex- sexual dimorphism and as follows: frons black cept sterna 3–4 with a yellow fascia on pos- with two lateral silver pruinose maculae on terior margin and sternum 5 with two lateral eye margin; pro- and mesolegs yellow except yellow maculae. Male genitalia: very similar apical tarsomeres; metafemur yellow; terga Downloaded from Brill.com09/25/2021 11:12:33AM via free access
2–5 black with two lateral yellow maculae ZFMK-DIP-00015955]; 1♂, CAMEROON: North- (those on tergum 2 elongated along lateral West Prov., Mezan Division, Bafut-Ngemba, margin); sterna yellow. env. Maison Forest, 2000 m., 10.x.1970, L. Matile [MNHN; ZFMK-DIP-00015954]; 1♂, CAM- Variation: No variation has been observed in EROON: North-West Prov., Mezan Division, the studied specimens, although some vari- Bafut-Ngemba, Vallée de la Haute Ngemba, ability in the extension of the abdominal 2100 m., 27.x.1975, Mission Cameroun C.N.R.S. maculae and in the coloration of the legs at RCP 318 [MNHN; ZFMK-DIP-00015956]; 1♀, some extent might be possible. CAMEROON: North-West Prov., Mezan Divi- sion, Bafut-Ngemba, Vallée de la Haute Ngem- Length (N = 3): Body, 9.17 mm (8.3–10.2 mm); ba, galerie, marigot avaut lac, 1800–1900 m., wing, 7.9 mm (7.2–8.6 mm). 30.x.1975, Mission Cameroun C.N.R.S. RCP 318 [MNHN; ZFMK-DIP-00015957]; 1♀, CAMER- Geographical distribution. Species known OON: North-West Prov., Mezan Division, Ba- from Cameroon. fut-Ngemba, Vallée de la Haute Ngemba, prés Maison forestiére, marigot, 2000 m., 25.x.1975, Etymology. The new species is named after its Mission Cameroun C.N.R.S. RCP 318 [ZFMK; collector, Dr. Štěpán Janeček. Species epithet ZFMK-DIP-00015957]. to be treated as a noun in the genitive case. Material of Melanostoma quadripunctatum Biology. See Vlašánková et al. (2017, as Melanos- (Skevington & Thompson, 2014) comb. nov. toma sp.) for its relationship with Impatiens KENYA: Kakamega Forest, 1585 m. [=5200 burtonii var. burtonii Hook. f. (Balsaminaceae). ft.], 18.xii.1970, E.E. Stubbs [4♂ 2♀, BMNH; 2♂ 1♀, ZFMK; ZFMK-DIP-00019927, …00015944, Material examined: Type material. Holotype, …00015945, …00015947, …00015950, …00015951, male, pinned with genitalia in microvial, …00015943, …00015946, …00015952]; …, 19.xii. deposited in the Zoologisches Forschungs- 1970, … [1♀, BMNH; ZFMK-DIP-00015953]; …, museum Alexander Koenig (ZFMK), Bonn, 20.xii.1970, … [2♂, BMNH; ZFMK-DIP-00015948, Germany, and labelled: “CAMEROON: North- …00015949]. West Prov., // Bamenda Highlands, 5km SE Big Remarks. Prior to this study, female speci- // Babanki, Mendong Buo area, // 6.592699°N, mens of M. quadripunctatum were unknown. 10.189999°E // 2100–2200 m., XI.2012. // Leg.: S. They are very similar to females of M. janeceki Janeček.” “DNA voucher specimen // ZFMK, sp. nov., but they can be distinguished by ster- Lab code // D226 // Bonn, Germany” “HO- nal and leg coloration as indicated in the diag- LOTYPE // Melanostoma // janeceki // Det. X. nosis (fig. 5F). Mengual 2017” [red] “ZFMK DIP 00015940” [barcode]. Paratypes: 1♂, same data as ho- lotype, DNA voucher ZFMK-D227 [ZFMK; Discussion ZFMK-DIP-00015941]; 1♀, same data as ho- lotype, DNA voucher ZFMK-D228 [ZFMK; Inferred generic relationships ZFMK-DIP-00015942]; 1♂, CAMEROON: Pelloloma, Talahua, Tuberculanostoma and Eo- North-West Prov., Mezan Division, Bafut- cheilosia have been included in a multi-gene Ngemba Forest Reserve (35 km from Bamen- phylogenetic analysis for the first time (see da), in the way to the Lake (env. Chemin fig. 2). In its original description, Vockeroth du Lac), 1500 m., 12.x.1970, L. Matile [ZFMK; (1973) placed theDownloaded genus fromPelloloma Brill.com09/25/2021 within 11:12:33AM the via free access
Syrphini, but already mentioned the possibil- produced face and a clear facial tubercle, but ity that it could be placed among the Mela- there are many differences from the Neotropi- nostomini. Kassebeer (2000a) explained using cal Tuberculanostoma as stated by Thompson morphological characters the reason to move & Skevington (2014). Results from Young et al. Pelloloma to Melanostomini. Present results (2016b) using the 5′ end of the COI gene re- (fig. 2) support the placement of this genus in solved Tuberculanostoma embedded in Platy- the Melanostomini. cheirus, somehow related with the stegnus Fluke (1945) described a new Melanostoma subgroup, referred as subgenus Platycheirus subgenus, Melanostoma (Talahua), for the (Carposcalis) by other authors (see Vockeroth, species Melanostoma fervidum Fluke, 1945, 1990; Thompson & Skevington, 2014; Young which is characterized by large male genitalia. et al., 2016b). Previously, Vockeroth (1990) Hull (1949) followed Fluke (1945) and consid- suggested the possibility to include Tubercu- ered Talahua as a Melanostoma subgenus, but lanostoma in Platycheirus as an alternative Thompson et al. (1976) elevated Talahua to placement. Present results (fig. 2) support the full generic status as it has a complete meta- suggestion of Vockeroth (1990) and corrobo- sternum, not reduced as defined for Mela- rate the results of Young et al. (2016b). Based nostoma. More recently, Thompson (1999) on the current evidences, the genus Platychei- placed a second species in the genus Talahua, rus comprises four subgenera: Platycheirus Melanostoma palliatum Fluke, 1975, which has (Carposcalis), Platycheirus (Pachysphyria), normal-sized male genitalia. Hereafter, the Platycheirus (Platycheirus), and Platycheirus size of the male genitalia was not diagnostic (Tuberculanostoma) comb. nov. Nevertheless, for Talahua, but only the character of meta- as stated by Young et al. (2016b), there is the coxa with a pile tuft postero-medially as stated possibility that other monophyletic species in the identification key by Thompson (1999). groups within the genus Platycheirus could be Our analysis resolved Talahua and Xanthan- recognized as distinct in the future. The pres- drus as sister taxa and Argentinomyia as sister ent results differ from the proposed subgen- taxon of Melanostoma (fig. 2), supporting the eric classification of Thompson & Skevington decision of Thompson et al. (1976). (2014) as Pyrophaena and Eocheilosia are sup- When Fluke (1943) described his new genus ported as valid genera and Pseudoplatychirus Tuberculanostoma from Ecuador, he stated is recognized as a junior synonym of Platychei- the similarity with Carposcalis Enderlein, 1938 rus (new synonym). Platycheirus (Pachysphyr- (at the time most Carposcalis species were ia) is equivalent to the Platycheirus ambiguus treated as Melanostoma) based on the pres- species group sensu Vockeroth (1990), Nielsen ence of curved bristles on the profemur. This (2004, 2014) and Young et al. (2016b). Based on similarity was also pointed out by Hull (1949). molecular characters (Mengual et al., 2008; Fluke (1943) also mentioned some diagnos- this study) and the morphological characters tic characters, such as face produced forward of the original descriptions, the Pseudoplaty- into a snout and a prominent facial tubercle, chirus species are Platycheirus species with and the preferential habitat in high elevations. elongated antennae, dichoptic males, wid- Years later, van Doesburg (1955) described Tu- ened protarsi, profemur with long, strong, berculanostoma solitarium van Doesburg, 1955 curly setae and projected face. Broadened from the Karakorum Mountains (a mountain protarsi, projected face and modified setae on range spanning the borders of Pakistan, India, the profemur are characteristics also found in and China) (see fig. 4C–H). This species has a other species groups within Platycheirus. All
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Table 2 New synthesis of generic and subgeneric names of Bacchini stat. rev. and Melanostomini stat. rev. (Syrphidae: Syrphinae) Tribe Genus Subgenus Author, year Type species Melanostomini Afroxanthandrus Kassebeer, 2000 Afroxanthandrus congoensis (Curran, 1938) Melanostomini Argentinomyia Lynch Arribalzaga, Argentinomyia testaceipes Lynch 1891 Arribalzaga, 1891 Melanostomini Leucopodella Hull, 1949 Leucopodella lanei (Curran, 1936) = Leucopodella bigoti (Austen, 1893) Melanostomini Melanostoma Schiner, 1860 Melanostoma mellinum (Linnaeus, 1758) Melanostomini Pelloloma Vockeroth, 1973 Pelloloma nigrifacies Vockeroth, 1973 Melanostomini Talahua Fluke, 1945 Talahua fervida (Fluke, 1945) Melanostomini Xanthandrus Androsyrphus Thompson, 1981 Xanthandrus (Androsyrphus) setifemoratus Thompson, 1981 Melanostomini Xanthandrus Xanthandrus Verrall, 1901 Xanthandrus (Xanthandrus) comtus (Harris, 1780) Bacchini Baccha Fabricius, 1805 Baccha elongata (Fabricius, 1775) Bacchini Eocheilosia Hull, 1949 Eocheilosia ronana (Miller, 1921) Bacchini Platycheirus Carposcalis Enderlein, 1937 Platycheirus (Carposcalis) steg- nus (Say, 1829) Bacchini Platycheirus Pachysphyria Enderlein, 1937 Platycheirus (Pachysphyria) dexter (Harris, 1780) Bacchini Platycheirus Platycheirus Lepeletier & Serville, Platycheirus (Platycheirus) 1828 scutatus (Meigen, 1822) Bacchini Platycheirus Tuberculanostoma Fluke, 1943 Platycheirus (Tuberculanostoma) antennatus (Fluke, 1943) Bacchini Pyrophaena Schiner, 1860 Pyrophaena rosarum (Fabricius, 1787) Bacchini Rohdendorfia Smirnov, 1924 Rohdendorfia dimorpha Smirnov, 1924 Bacchini Spazigaster Rondani, 1843 Spazigaster apennini Rondani, 1843 = Spazigaster ambulans (Fabricius, 1798) Bacchini Syrphocheilosia Stackelberg, 1964 Syrphocheilosia aterrima Stack- elberg, 1964 = Syrphocheilosia claviventris (Strobl, 1910)
* Pseudoplatychirus van Doesburg, 1955 is a junior synonym of Platycheirus Le Peletier & Audinet-Serville, 1828 and Afrostoma Skevington, Thompson & Vockeroth, 2014 is a junior synonym of Melanostoma Schiner, 1860. male genitalia, and without male external sec- modified protarsi, protibiae and/or with dis- ondary characters as all other Melanostoma tinctive bristles or pile tufts on profemora, species, but with an entire metasternum. but they are so variable that cannot be used Two morphological characters were diag- as diagnostic characters to define generic nostic to separate Melanostoma and Platy- groups (Vockeroth, 1990; Young et al., 2016b). cheirus species (Andersson, 1970; Barkalov, In a series of papers devoted to Bacchini 2009; Haarto & Ståhls, 2014): the shape of the sensu lato, Fluke (1937, 1943, 1945, 1957) stud- metasternum and the morphology of male ied in deep this group of genera and was the genitalia, especially the shape of surstyli and first author to study the male genitalia of postgonites. Most Platycheirus males have these taxa (Fluke, 1957). When male genitalia Downloaded from Brill.com09/25/2021 11:12:33AM via free access
Brink, 1973), and probably also in the other nize and name theDownloaded two groups from Brill.com09/25/2021 with simple, 11:12:33AM via free access
1967; Thompson, 1972; Vockeroth, 1973, 1990, and elongate surstyliDownloaded with from a Brill.com09/25/2021small basomedial 11:12:33AM via free access
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