Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Jon Peder Lindemann (jon.p.lindemann@uit.no), Geir Søli (geir.soli@nhm.uio.no), Jostein Kjærandsen (jostein.kjarandsen@uit.no)
Academic editor: Vladimir Blagoderov
Received: 08 Apr 2021 | Accepted: 13 Aug 2021 | Published: 24 Sep 2021
© 2021 Jon Peder Lindemann, Geir Søli, Jostein Kjærandsen
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:
Lindemann JP, Søli G, Kjærandsen J (2021) Revision of the Exechia parva group (Diptera: Mycetophilidae). Biodiversity Data Journal 9: e67134. https://doi.org/10.3897/BDJ.9.e67134
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Exechia is a diverse genus of small fungus gnats, widespread in the Holarctic Region, while the fauna is largely unknown elsewhere, such as in the Afrotropical and Oriental Region. Members of Exechia can be arranged into several species groups, based on homologies in the male and female terminalia. The Exechia parva group is delimited, based on male terminalia possessing a pair of gonocoxal lobes on the apicoventral gonocoxal margin. Eight previously-described species can be placed in this group, of which six are from the Holarctic Region, while one is recorded each from the Oriental and the Afrotropical Regions.
The Exechia parva group was reviewed and found to include 33 species, of which 24 were described as new to science and six were re-described. Identification keys to 32 species for males and nine species for females are provided together with illustrations and photos of male and female terminalia. Species delimitations were based on morphological examination of 94 male and female specimens, as well as DNA barcodes obtained from 124 specimens. Molecular and morphological species delimitations were mostly congruent, except in two cases where two species were delimited within a single Barcode Index Number (BIN). We found that each species is only known from a single zoogeographical region and that several species complexes are largely congruent with zoogeographical divisions, indicating that intercontinental barriers may have a strong impact on the species diversity of the group.
fungus gnats, Exechia, parva, repanda, taxonomy, DNA barcodes
The genus Exechia Winnertz, 1863 is known from all zoogeographical regions, except the Neotropical Region and Antarctica (Bechev 1999). With 158 valid species (
While most of the species have been described from the Holarctic Region, where the collecting effort traditionally has been greatest, it is likely that a large part of the fauna from other regions still remains to be described. It is estimated that two thirds of the Afrotropical Diptera diversity is undescribed and Mycetophilidae is considered to be one of the lesser known Diptera families of the Region (
Species identification is mainly based on comparison of the male or female terminalia. Some identification keys have been published (
A few species groups in Exechia have been sorted out and characterised.
In this study, we will revise the Exechia parva group, which involves descriptions of 24 new species and re-descriptions of six species. We apply an integrative approach, based on morphological and molecular delimitation methods, to revise the species and identify species boundaries.
The study is based on material from North-Western Europe, North America, East Africa, South Africa, Madagascar, Japan, Nepal and Bhutan, obtained from several different insect collections. The material was collected in the time span between 1971 and 2019, mostly by sweep net and malaise traps. Material from Nepal was collected by the Kyushu University Expedition to the Himalayas in 1971 and 1972 and by U. Emoto during an expedition to East Nepal in 1981. Material from Bhutan was collected by T. Saigusa during his collecting trip to Bhutan in 1993. Afrotropical material has partly been received from M. Mostovski and M. Jaschhof, collected during their travel in South Africa in 2005 and from one of the authors, J. Kjærandsen, as well as a loan from National Museum, Bloemfontein to the Natural History Museum in Oslo.
Specimens from the following collections have been examined:
For molecular species delimitation, the mitochondrial gene, cytochrome oxidase subunit 1 (CO1), was used as the DNA barcode. Legs from selected individuals were sent to the Canadian Center for DNA Barcoding (CCDB) in Guelph for subsequent DNA extraction, amplification and sequencing. DNA amplification was performed with the primer pair C_LepFoIF and C_LepFoIR (
Molecular species delimitation was mainly based on the Barcode Index Number (BIN) system (
Male and female terminalia were dissected from specimens and heated in lactic acid to remove soft tissue. Temporary slide mounts were made using glycerol as medium, so that the terminalia later could be stored on microvials in glycerol, together with the pinned specimens. For most male specimens, the gonostyli were dissected. Slides were photographed using a Zeiss Axio imager M2 microscope, together with an Axiocam 506 colour camera. Stacked images were rendered in the image-stacking software Helicon Focus 7, sharpened in Topaz Sharpen AI and post-edited in Adobe Photoshop CS6. Illustrations were made in Adobe Illustrator.
The morphological terminology follows
The following abbreviations will be used throughout the text:
Terminology used to describe male and female terminalia is shown in Figs
Male terminalia. Abbreviations: ae = aedaegus, ag = aedaegal guides, ce = cercus, db = dorsal gonostylus branch, gc = gonocoxite, gl = gonocoxal lobe, hl = hypandrial lobe, hy = hypandrium, ib = internal gonostylus branch, mb = medial gonostylus branch, pm = paramere, tg = tergite, vb = ventral gonostylus branch.
Description:
Colouration. Pale to dark brown individuals. Head usually darker than scutum. Palpus, scape, pedicel and basal part of first flagellomere usually paler than rest of head and antennae. Scutum uniformly coloured, sometimes with pale lateral and frontal margins; mesonotal stripes rarely present. Legs usually whitish-yellow. Halteres usually whitish-yellow, apically sometimes slightly darker. Wings usually hyaline with yellow or pale brown tint, rarely with weak dark marks. Abdomen pale brown to dark brown, sometimes tergites I-III slightly paler or with lateral pale area. Female tergites II-VI usually with lateral and/or dorsolateral pale areas, at the anterior margin of the tergites. Terminalia yellow to brown. Head. Two ocelli, touching margins of compound eyes. Vertex with row of 6 orbital bristles along each eye margin; 12 frontal bristles. Frons and vertex covered with short, usually pale setae, extending from level of frontal bristles to occiput. Clypeus covered with thin pale setae, usually most dense on ventral part. Antennae with 14 flagellomeres, covered with fine setae. Thorax. Scutum with strong pre-alar and postalar bristles and three rows of discal bristles. Scutellum with one pair of strong bristles, usually curving anteriorly. Bristles on scutum and scutellum slightly forked apically, with one branch shorter than the other (as illustrated in
Diagnosis:
Males. Distinguished from species in the E. cincta group in having the hypandrial lobe with branches diverging in a wide angle (Fig.
Females. Distinguished from species in the E. fusca group in having the hypogynal valves apicoventrally emarginate, truncate or obtuse (Fig.
Exechia adenaparva Chandler, 2000: 282 ♂ (
Afrotropical, Yemen (Fig.
Distribution map of species in the E. parva group occurring in the Afrotropical Region, based on localities from studied material, type material and DNA barcoded material. Different species are represented by individual colours. Altitudes are indicated in shades from grey (lowland) to white (mountains).
We did not have the opportunity to re-describe this species, but the male terminalia are well illustrated in
Exechia adenaparva Chandler, 2000. Abbreviations: db = dorsal gonostylus branch, gl = gonocoxal lobe, ib = internal gonostylus branch, mb = medial gonostylus branch, vb = ventral gonostylus branch.
Male: Body length 5.2 mm. Wing length 4.0 mm. Colouration (Dry specimen). Head dark brown; eye margin, face and clypeus yellow; labellum yellow; palpus yellow-brown, segments 4-5 dark brown. Antennae with scape and pedicel yellow; flagellum brown, first flagellomere with yellow base. Thorax with scutum brown, pale mesonotal stripes present and anterior and lateral margins broadly yellow; lateral sclerites pale brown to dark brown; propleura brown; halteres whitish-yellow, apically slightly darker. Legs whitish-yellow. Abdomen dark brown, tergites I-III with yellow lateral area. Terminalia pale brown. Head. Frons and vertex covered with short black setae. Clypeus covered with short black setae, evenly distributed; flagellomeres longer than broad, with sixth flagellomere 1.2 times as long as wide. Thorax. Scutum covered with short black setae. Legs. Mid-tibia with 23 anterior, 9 posterodorsal, 10 posterior and 6 posteroventral bristles. Hind tibia with 14 anterodorsal, 8 posterodorsal and 8 posterior bristles. Abdomen. Tergites covered with black setae. Terminalia (Fig.
Female unknown.
Distinguished from all species in the E. parva group in having the gonocoxal lobes entirely covered with setae and entire length tapering (Fig.
From afro, relating to the Afrotropical Region and parva, relating to the resemblance to E. parva.
Afrotropical, Burundi (Fig.
Adult collected in afromontane forest (2237 m a.s.l.).
Male: Wing length 3.2 mm. Colouration (Dry specimen). Head dark brown; face and clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, first flagellomere with yellow base. Thorax with scutum brown, lateral margin broadly yellow; lateral sclerites pale brown; propleura yellow; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-III with lateral yellow area. Terminalia. Head. Frons and vertex covered with pale setae. Clypeus covered with pale brown setae. Antenna long, 2.3 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, with sixth flagellomere 1.4 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 0.93 times as long as first tarsomere. Mid-tibia with 20 anterior, 4 posterodorsal, 8 posterior and 4 posteroventral bristles. Wings. Vein r-m 3 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Female: Unknown.
Distinguished from all species in the E. parva group in having the gonocoxal lobes distinctly curved interiorly (Fig.
From Latin arcus, bow, relating to the shape of the gonocoxal lobe.
Afrotropical, Kenya (Fig.
Adult collected in bamboo forest (3050 m a.s.l.).
Male: Body length 3.2 mm. Wing length 2.5 mm. Colouration (Dry specimen). Head face and clypeus dark brown, almost black; labellum brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, basal half of first flagellomere yellow. Thorax with scutum dark brown, lateral margin pale brown; lateral sclerites and propleura brown; halteres whitish-yellow, apically slightly darker. Legs whitish-yellow. Abdomen dark brown, tergites II-III with slightly paler lateral area. Terminalia yellow. Head. Frons and vertex covered with long pale brown setae. Clypeus covered with pale setae, evenly distributed; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with long, pale brown setae. Legs. Fore leg with tibia 0.76 times as long as first tarsomere. Mid-tibia with 17 anterior, 4 posterodorsal, 9 posterior and no posteroventral bristles. Hind tibia with 7-8 anterodorsal, 5 posterodorsal and 5 posterior bristles. Abdomen. Tergites covered with long, dark brown setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. penicillata and E. sambai in having the dorsal gonostylus branch short and squared with apico-internal corner right-angled (Fig.
Named in honour of Dr. Ashley Kirk-Spriggs, the collector of the holotype, who also collected other invaluable material of several species described in this revision.
Afrotropical, Burundi (Fig.
Adult collected in afromontane forest (2237 m a.s.l.).
Male: Body length 3.7 mm. Wing length 3.2 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum brown; palpus dark brown. Antenna with scape and pedicel yellow; flagellum dark brown, basal half of first flagellomere pale. Scutum and lateral sclerites dark brown; propleura brown; halteres whitish-yellow. Wings hyaline with two weak dark marks, one reaching from apical part of costal cell to middle of cell r4+5 and the other covering area posterior to cubital fork (Fig.
Exechia bifasciata sp. n.
Female: Unknown.
Distinguished from all species in the E. parva group in having the wings with dark marks (Fig.
From Latin fascia, band, with the Latin prefix bi-, two, relating to the specific wing pattern, forming two dark bands.
East Palaearctic, Japan (Fig.
Distribution map of species in the E. parva group occurring in the East Palaearctic Region, based on localities from studied material, type material and DNA-barcoded material. Different species are represented by individual colours. Altitudes are indicated in shades from grey (lowland) to white (mountains).
Unknown.
Male: Body length 3.2 mm. Wing length 2.5 mm. Colouration (Dry specimen). Head and face dark brown; clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown with first flagellomere pale brown. Thorax with scutum and lateral sclerites brown; propleura pale brown; halteres whitish-yellow, apically slightly darker. Legs whitish-yellow. Abdomen dark brown, tergites II-III with lateral yellow area. Terminalia pale brown. Head. Frons and vertex covered with long dark brown setae. Clypeus covered with brown setae, more dense towards ventral side; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with long dark brown setae. Legs. Fore leg with tibia 0.87 times as long as first tarsomere. Mid-tibia with 20 anterior, 5 posterodorsal, 8 posterior and no posteroventral bristles. Hind tibia with 7-8 anterodorsal, 8-9 posterodorsal and 4 posterior bristles. Abdomen. Tergites covered with long dark brown to black setae. Terminalia (Fig.
Female: Unknown.
Distinguished from all species in the E. parva group in having the dorsal branch of the gonostylus with an elongated and narrow baso-internal lobe extending in an almost straight angle and curving distally (Fig.
From Latin, brachiatus, branched, relating to the shape of the gonostylus with bilobed dorsal and medial branches and trilobed internal branch.
Afrotropical, Madagascar (Fig.
Unknown.
Male (n = 2): Body length 2.9-3.2 mm. Wing length 2.3-2.5 mm. Colouration (Dry specimen). Head dark brown; face and clypeus brown; labellum pale brown; palpus whitish-yellow. Antenna with scape and pedicel brown; flagellum brown. Scutum, lateral sclerites and propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown. Terminalia yellow. Head. Frons and vertex covered with pale setae. Clypeus covered with few (15-17) pale setae, evenly distributed. Antenna short, 1.45-1.5 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere 0.8-0.9 times as long as wide. Thorax. Scutum covered with short pale brown setae. Legs. Fore leg with tibia 0.96-1.0 times as long as first tarsomere. Mid-tibia with 20-21 anterior, 3-4 posterodorsal, 7-9 posterior and (n = 1) 2 posteroventral bristles. Hind tibia with 5 anterodorsal, 4-5 posterodorsal and 4-5 posterior bristles. Wings. Vein r-m 2.46-2.6 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Exechia breviflagellata sp. n.
Female (n = 1): Body length 3.1 mm. Wing length 2.7 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum pale brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown. Scutum, lateral sclerites and propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown, tergites II-VI with paler lateral areas, not extending notably dorsally. Terminalia yellow. Head. Frons and vertex and clypeus covered with pale setae. Antenna short, 1.4 times as long as length from vertex to ventral margin of clypeus; flagellomeres broader than long, with sixth flagellomere 0.7 as long as wide. Thorax. Scutum covered with pale brown setae. Legs. Fore leg with tibia as long as first tarsomere. Hind tibia with 6 anterodorsal and 6 posterodorsal bristles. Wings. Vein r-m 2.8 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Distinguished from E. brevilobata in having the gonostylus with the dorsal branch only 1.37-1.42 times longer than broad, with its apical lobe broader and shorter, 0.18-0.2 of the total dorsal branch length (Fig.
From Latin brevis, short and flagellum, whip, relating to the short antennae of the species.
Nearctic, Canada (Fig.
Distribution map of species in the E. parva group occurring in the Nearctic Region, based on localities from studied material, type material and DNA-barcoded material. Different species are represented by individual colours. Altitudes are indicated in shades from grey (lowland) to white (mountains).
Adults collected in wetland habitats.
Male (n = 9): Body length (n = 8) 3.5-3.8 mm. Wing length 2.7-3.2 mm. Colouration (Dry specimen; n = 2). Head brown to dark brown, face and clypeus pale brown to dark brown; labellum yellow to brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum yellow to brown with basal part of first segment pale. Scutum brown to dark brown, anterolateral areas distinctly paler; lateral sclerites pale brown to dark brown, propleura yellow to dark brown; halteres whitish-yellow. Legs yellow. Abdomen brown to dark brown. Terminalia yellow. Head. Frons and vertex covered with whitish to pale brown setae. Clypeus covered with thin pale setae, evenly distributed. Antenna short, 1.5-1.65 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with pale brown setae. Legs. Fore leg with tibia 1.0 times as long as first tarsomere. Mid-tibia with 20-22 anterior, (n = 3) 4 posterodorsal, 8 posterior and 1-2 posteroventral bristles. Hind tibia (n = 3) with 7-8 anterodorsal, 4 posterodorsal and 4-5 posterior bristles. Wings. Vein r-m 2.3-2.5 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Exechia brevilobata sp. n. male terminalia.
Female: Unknown.
Distinguished from E. breviflagellata in having the gonostylus with the dorsal branch 1.8-2 times longer than broad, with its apical lobe narrower and longer, 0.21-0.25 of the total dorsal branch length (Fig.
From Latin brevis, short and lobatus, with lobes, relating to the short apical lobe of the dorsal branch of the gonostylus.
East Palaearctic, Norway, Sweden (Fig.
Distribution map of species in the E. parva group occurring in the West Palaearctic Region, based on localities from studied material, type material and DNA-barcoded material. Different species are represented by individual colours. Altitudes are indicated in shades from grey (lowland) to white (mountains).
Unknown.
Material of this species from Jokkmokk, Sweden, have earlier been identified as E. repandoides (
Adult male: Body length 2.6 mm. Wing length 2.1 mm. Colouration (Dry specimen). Head dark brown; face and clypeus yellow; labellum yellow; palpus yellow, with segments 4-5 yellow to brown. Antenna with scape and pedicel yellow; flagellum brown. Thorax with scutum brown; lateral sclerites pale brown; propleura yellow; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites I-III with slightly paler lateroventral area. Terminalia pale brown. Head. Frons and vertex covered with short pale brown setae. Clypeus covered with only a few (about 11) black setae, mostly on ventral side; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with short pale brown setae. Legs. Fore leg with tibia 0.95 times as long as first tarsomere. Mid-tibia with 16 anterior, 4 posterodorsal, 7 posterior and 2 posteroventral bristles. Hind tibia with 11 anterodorsal, 4 posterodorsal and 3 posterior bristles. Abdomen. Tergites covered with long pale brown to yellow setae. Terminalia (Fig.
Female: Unknown.
Distinguished from all species in the E. parva group in having the gonocoxal lobe with apical setae distinctly splaying (Fig.
From Burundi, the country where the holotype was collected, with Latin ending -ensis, belonging to.
Afrotropical, Burundi (Fig.
Adult collected in afromontane forest (2237 m a.s.l.).
Exechia capillata Johannsen, 1912:73 ♂♀ (
Male (n = 4): Body length 2.7-3.5 mm. Wing length 2.4-3.0 mm. Colouration (Dry specimen). Head dark brown; face and clypeus brown to dark brown; labellum brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum brown to dark brown, basal part of first segment pale. Scutum and lateral sclerites brown to dark brown; propleura yellow to dark brown; halteres whitish-yellow; wings hyaline tinged brown. Legs whitish-yellow. Abdomen dark brown, tergites I-III slightly paler, sometimes tergite II with distinct pale lateral area covering ventral half. Terminalia yellow. Head. Frons and vertex covered with pale short setae. Clypeus covered with pale setae, evenly distributed. Antenna (n = 2) 2.0-2.1 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate or slightly longer than broad, sixth flagellomere 1.0-1.15 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 2) 0.9-0.94 times as long as first tarsomere. Mid-tibia with 15-19 anterior, 4-5 posterodorsal, 5-7 posterior and 2-4 posteroventral bristles. Hind tibia with (n = 3) 5-7 anterodorsal, (n = 3) 3-5 posterodorsal and (n = 3) 4 posterior bristles. Wings. Vein r-m 2.5-2.9 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (n =2, Fig.
Exechia capillata Johannsen, 1912
Female (n = 4): Body length 2.6-2.7 mm. Wing length 2.2-2.3 mm. Colouration (Dry specimen). Head dark brown; face and clypeus brown to dark brown; labellum brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, basal part of first flagellomere pale. Thorax with scutum brown to dark brown; lateral sclerites brown; propleura pale brown; halteres whitish-yellow; wings hyaline tinged with brown. Legs whitish-yellow. Abdomen pale brown to dark brown, tergites II-VI with lateral pale areas extending somewhat dorsally along anterior margin of tergites III and VI, but not forming any complete band. Terminalia pale brown. Head. Vertex, frons and clypeus covered with pale setae. Antenna (n = 3) 1.8-1.9 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 3) 0.95-1.0 times as long as first tarsomere. Mid-tibia with (n = 3) 17-19 anterior, 3-4 posterodorsal, (n = 3) 5-6 posterior and (n = 3) 2-3 posteroventral bristles. Hind tibia with 4-5 anterodorsal, 5-6 posterodorsal and 4 posterior bristles. Wings. Vein r-m (n = 1) 2.4-2.6 times longer than stem of M-fork. Abdomen. Tergites covered with brown setae. Terminalia (Fig.
Distinguished from all species in the E. parva group in having the dorsal gonostylus branch with the medio-external margin forming a short distally projected process (Fig.
Nearctic, Canada, USA (Fig.
Reared from fruitbody of Collybia dryophila (Bull. : Fr.) P. Kumm. = Gymnopus dryophilus (Bull. : Fr.) Murrill (
Holotype with whole specimen, except terminalia mounted on slide in Canada balsam with one wing detached. Terminalia in alcohol, transferred to glycerine and associated with empty pin with labels in type collection. A second tube with terminalia of E. cf. dorsalis misplaced in same vial - this was also transferred to glycerine in a microvial and pinned separately.
Male: Body length 3.3 mm. Wing length 2.8 mm. Colouration (Dry specimen). Head, face dark and clypeus dark brown; labellum pale brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, basal half of first flagellomere pale. Scutum, lateral sclerites and propleura brown; halteres yellow. Legs yellow. Abdomen dark brown. Terminalia yellow. Head. Frons and vertex covered with pale setae. Clypeus covered with pale setae, evenly distributed. Antenna long, 2.2 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, with sixth flagellomere 1.5 as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 0.83 times as long as first tarsomere. Mid-tibia with 21 anterior, 3 posterodorsal, 10 posterior and 3 posteroventral bristles. Hind tibia with 9 anterodorsal, 4 posterodorsal and 6 posterior bristles. Wings. Vein r-m 2.9 times longer than stem of M-fork. Abdomen. Tergites covered with short, pale setae. Terminalia (Fig.
Distinguished from all species in the E. parva group in having the medial gonostylus branch with an apical finger-like process with 2 setae at the apex (Fig.
From Latin chirotheca, mitten, relating to the shape of the dorsal and medial lobes of the gonostylus, resembling mittens.
Oriental, Nepal (Fig.
Distribution map of species in the E. parva group occurring in the eastern Himalayas (Oriental Region), based on localities from studied material, type material and DNA-barcoded material. Different species are represented by individual colours. Altitudes are indicated in shades from grey (lowland) to white (mountains).
Adult collected in the eastern Himalayas (2300 m a.s.l.).
Exechia cinctiformis Storå, 1941: 2 ♂♀ (
Exechia sp. Storå, 1941:3 ♀ (
Exechia dahli Nielsen, 1966: 8 ♂♀ (
West Palaearctic, Madeira (Fig.
We did not have the opportunity to re-describe this species, but the male terminalia are well illustrated in
Male: Fig.
Exechia cinctiformis Storå, 1941, male terminalia. Abbreviations: ce = cercus, db = dorsal gonostylus branch, gl = gonocoxal lobe, ib = internal gonostylus branch, mb = medial gonostylus branch, tg = tergite, vb = ventral gonostylus branch.
Male: Body length 3.9 mm. Wing length 3.2 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum dark brown; palpus whitish-yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal half of first flagellomere pale. Thorax with scutum pale brown, lateral margin paler; lateral sclerites brown; propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-III with a lateral pale area. Terminalia brown. Head. Frons and vertex covered with pale setae. Clypeus covered with only few (about 17) pale setae. Antenna long, 2.1 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, sixth flagellomere 1.5 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 0.91 times as long as first tarsomere. Mid-tibia with 19 anterior, 4 posterodorsal, 7 posterior and 3 posteroventral bristles. Wings. Vein r-m 2.14 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. serrae in having the gonocoxal lobe extended beyond the apex of the apical hypandrial setae (Fig.
From Latin columna, column, relating to the shape of the hypandrium forming a large distally-extended lobe.
Oriental, Nepal (Fig.
Adult collected in the eastern Himalayas (3500 m a.s.l.).
Male: Body length 3.6 mm. Wing length 3.2 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum pale brown; palpus yellow to pale brown. Antenna with scape and pedicel yellow; flagellum dark brown, basal half of first flagellomere yellow. Thorax with scutum dark brown, except narrow yellow anterolateral margin; lateral sclerites dark brown; propleura brown; halteres whitish-yellow. Legs yellow. Abdomen dark brown, tergites II-III with yellow laterodorsal area. Terminalia pale brown with MB dark brown. Head. Frons and vertex covered with brown setae. Clypeus covered with pale setae, most dense towards ventral side; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with brown setae. Legs. Fore leg with tibia 0.7 times length of first tarsomere. Mid-tibia with 23 anterior, 5 posterodorsal, 10 posterior and 5 posteroventral bristles. Hind tibia with 10 anterodorsal, 6 posterodorsal and 4 posterior bristles. Wings. Vein r-m 3.3 times longer than stem of M-fork. Abdomen. Tergites covered with long brown setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. trunciseta by the shape of the medial gonostylus branch (Fig.
From Latin crassus, stout and seta, bristle, relating to the shape of the seta on apicoventral margin of the gonocoxites and apically on the dorsal branch of the gonostylus.
Oriental, Nepal (2900-3000 m a.s.l., Fig.
Unknown
Male (n = 1): Body length 3.2 mm. Wing length 2.8 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum dark brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal half of first flagellomere pale. Thorax with scutum dark brown, anterior and lateral margin paler; lateral sclerites and propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown with pale lateral area confined to ventral margin of tergite II and basolateral part of tergite III. Terminalia yellow. Head. Vertex, frons and clypeus covered with pale setae. Antenna 1.9 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, with sixth flagellomere 1.2 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 1.05 times as long as first tarsomere. Mid-tibia with 20 anterior, 4 posterodorsal, 10 posterior and 3 posteroventral bristles. Hind tibia with 6 anterodorsal, 5 posterodorsal and 3 posterior bristles. Wings. Vein r-m 2.22 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (n = 2, Fig.
Exechia curvata sp. n.
Female (n = 4): Body length 2.6-2.8 mm. Wing length 2.4-2.5 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal half of first flagellomere pale. Thorax with scutum brown to dark brown; lateral sclerites brown; propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown to dark brown, tergites II-VI with pale lateral areas extending dorsally at anterior fourth to third of tergite III and anterior third to half of tergite IV, forming two distinct pale bands. Terminalia pale brown. Head. Vertex, frons and clypeus covered with pale setae. Antenna 1.75-1.9 times as long as length from vertex to ventral margin of clypeus; flagellomeres slightly longer than broad, with sixth flagellomere (n = 2) 1.0-1.1 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia as long as first tarsomere. Mid-tibia with 20-24 anterior, (n = 3) 3 posterodorsal, (n = 3) 6-8 posterior and 1-3 posteroventral bristles. Hind tibia with (n = 3) 6-7 anterodorsal, 4 posterodorsal and 3-4 posterior bristles. Wings. Vein r-m 2.0-2.2 times longer than stem of M-fork. Abdomen. Tergites covered with brown setae on dark areas and pale setae on pale areas. Terminalia (Fig.
Distinguished from E. repanda, E. subrepanda and E. neorepanda in having the dorsal gonostylus branch more curved and with external margin forming a distinct protruding angle, with the external row of setae covering only about one sixth of the total dorsal gonostylus branch length (Fig.
From Latin curvus, curved, relating to the shape of the dorsal branch of the gonostylus.
Nearctic, Canada, USA (Fig.
Adult collected in temperate mixed forest.
Exechia longichaeta Wu, Xu & Yu, 2004: 555 ♂ (
Palaearctic, Oriental, China
We did not have the opportunity to re-describe this species; however, the male terminalia are well illustrated in
Male (n = 4): Body length 3.0-3.1 mm. Wing length 2.5 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum yellow; palpus yellow. Antenna with scape and pedicel yellow; flagellum yellow. Thorax with scutum and lateral sclerites brown; propleura pale brown; halteres whitish-yellow, apically slightly darker. Legs yellow. Abdomen brown with tergite II slightly paler or with pale area covering its ventral half. Terminalia yellow. Head. Frons, vertex and clypeus covered with pale setae. Antenna short, 1.53-1.64 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere 0.9-0.95 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 0.96-1.1 times as long as first tarsomere. Mid-tibia (n = 1) with 19 anterior, 4 posterodorsal, 7 posterior and 2 posteroventral bristles. Hind tibia (n = 1) with 7 anterodorsal and 2 posterodorsal bristles. Wings. Vein r-m 2.5-3.1 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. spatulata in having the dorsal gonostylus branch more narrow, not spathulate, only 3-3.2 times longer than broad (Fig.
From Latin longus, long and lobatus, with lobes, relating to the long apical part of the dorsal branch of the gonostylus.
West Palaearctic, Sweden (Fig.
Adults collected in limestone quarry.
Male (n = 11, Fig.
Female: Unknown.
Distinguished from E. curvata, E. repanda and E. subrepanda in having the dorsal gonostylus branch only slightly curved, the external margin without a distinct angle and with the external row of setae covering one-third to half of the total dorsal gonostylus branch length (Fig.
From Greek neos, new and repanda, relating to the resemblance of the species to E. repanda. The name indicates that the species has commonly been identified as E. repanda.
West Palaearctic, Austria, Belgium, Britain, Corsica, Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Norway, Poland, Russia, Sweden, Switzerland, The Netherlands (Fig.
Reared from fruitbody of Lyophyllum loricatum (Fr.) Kühner, Mycena galericulata (Scop.) Gray, Kuehneromyces mutabilis (Schaeff.) Singer & A.H.Sm., Gyromitra esculenta (Pers.) Fr., Inocybe goodeyi Gillet, and Calocybe gambosa (Fr. : Fr.) Donk (
Adults have been reported hibernating in hollow, usually broken umbelliferous stems (
This Palaearctic species is morphologically very close to the two Nearctic species, E. repanda and E. subrepanda and species determination should be conducted with care or confirmed by their distinctly-separated DNA barcodes and BINs. We assume that previous records of E. repanda in the West-Palaearctic Region constitute what we define as E. neorepanda sp. n.
Exechia pararepanda Kallweit in Kallweit & Martens, 1995: 243 ♂ (
Male (n = 6): Body length (n = 3) 2.5-3.2 mm. Wing length (n = 3) 2.1-2.7 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown to dark brown, basal half of first flagellomere yellow. Scutum brown to dark brown, with anterolateral margin yellow; lateral sclerites brown to dark brown; propleura pale brown to brown; halteres whitish-yellow to yellow. Legs whitish-yellow to yellow. Abdomen brown to dark brown. Terminalia yellow to pale brown. Head. Frons and vertex covered with short, pale setae. Clypeus covered with few (7-15) thin pale setae, mainly on ventral half. Antenna long, (n = 2) 2.0-2.1 times as long as length from vertex to ventral margin of clypeus ; flagellomeres longer than broad, with sixth flagellomere (n = 5) 1.2-1.4 times as long as wide. Thorax. Scutum covered with short pale setae. Legs. Fore leg with tibia (n = 3) 0.88-0.91 times as long as first tarsomere. Mid-tibia with (n = 5) 15-19 anterior, (n = 5) 2-4 posterodorsal, (n = 5) 6-8 posterior and (n = 5) 1-4 posteroventral bristles. Hind tibia with 4-8 anterodorsal, 4 posterodorsal and 2-3 posterior bristles. Wings. Vein r-m (n = 3) 1.81-2.2 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Exechia pararepanda Kallweit, 1995, male terminalia.
Female: Unknown.
Distinguished from all species in the E. parva group in having the gonocoxal lobe apex with a row of 5 setae extending down apico-exterior margin (Fig.
Oriental, Bhutan, Nepal (Fig.
Adults collected in the eastern Himalayas (2400-3400 m a.s.l.).
Exechia parva Lundström, 1909: 50 ♂ (
Male (n = 4): Body length 3.0-3.8 mm. Wing length 2.5-3.2 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow to pale brown. Antenna with scape and pedicel yellow; flagellum dark brown to pale brown, basal half of first flagellomere pale. Scutum dark brown; lateral sclerites and propleura brown to dark brown; halteres whitish-yellow. Legs yellow to whitish-yellow. Abdomen brown to dark brown, tergites I-III slightly paler, sometimes tergite II with pale area confined to a narrow stripe along ventral margin. Terminalia yellow. Head. Vertex, frons and clypeus covered with pale setae. Antenna (n = 2) 1.8-2 times as long as length from vertex to ventral margin of clypeus ; flagellomeres longer than broad, with sixth flagellomere 1.1-1.3 times as long as wide. Thorax. Scutum covered with pale to pale brown setae. Legs. Fore leg with tibia (n = 1) 0.95 times as long as first tarsomere. Mid-tibia with 18-22 anterior, 3-4 posterodorsal, 4-8 posterior and (n = 2) 2-3 posteroventral bristles. Hind tibia with (n = 3) 6-8 anterodorsal, (n = 2) 5 posterodorsal and (n = 2) 4 posterior bristles. Wings. Vein r-m (n = 2) 2.45-2.5 times longer than stem of M-fork. Abdomen. Tergites covered with pale to pale brown setae. Terminalia (Fig.
Exechia parva Lundström, 1909
Female (n = 4): Body length 3.3-3.9 mm. Wing length 2.9-3.2 mm. Colouration (Dry specimen). Head, face and clypeus brown to dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown to brown, basal part of first flagellomere pale. Thorax with scutum brown to dark brown; lateral sclerites pale brown to brown; halteres whitish-yellow; wings hyaline tinged with brown. Legs whitish-yellow. Abdomen brown to dark brown, tergites III-VI with lateral pale areas, extending from lateral margin to about half tergal height. Terminalia pale brown. Head. Vertex, frons and clypeus covered with pale setae. Antenna (n = 2) 1.7-1.8 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with pale brown setae. Legs. Fore leg with tibia 0.93-1.0 times as long as first tarsomere. Mid-tibia with 23-26 anterior, 3-4 posterodorsal, 6-8 posterior and 2-3 posteroventral bristles. Hind tibia with 5-6 anterodorsal, 5 posterodorsal and 3-5 posterior bristles. Wings. Vein r-m (n = 2) 2.7-2.8 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (n = 3, Fig.
Distinguished from all species in the E. parva group in having the gonocoxal lobe evenly tapered and entirely covered with setae (Fig.
East Palaearctic, West Palaearctic, Austria, Belgium, Britain, Bulgaria, Croatia, Czech Republic, Estonia, Finland, France, Germany, Ireland, Latvia, Norway, Poland, Russia, Slovenia, Sweden, Switzerland, The Netherlands (
Reared from fruitbody of Nematoloma sp. [? udum (Pers. : Fr.) Karst.] = Hypholoma sp. [? udum (Pers. : Fr.) Kühner], Ptychoverpa bohemica (Krombh.) Boud. = Verpa bohemica (Krombh.) J. Schröt, and Russula ochroleuca (Pers.) Fr. (
Adults have frequently been reported hibernating in hollow, usually broken, umbelliferous stems (
Male: Body length 3.5 mm. Wing length 2.7 mm. Colouration (Dry specimen). Head and face dark brown; clypeus, labellum and palpus brown. Antenna with scape and pedicel yellow; flagellum brown. Thorax with scutum brown; lateral sclerites pale brown; propleura yellow; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-IV with a lateroventral yellow area. Terminalia pale brown. Head. Frons and vertex covered with short, brown setae. Clypeus covered with dense thin pale setae. Antennae elongate, 2.3 times as long as length from vertex to ventral margin of clypeus ; flagellomeres slightly longer than broad, with sixth flagellomere 1.1 times as long as wide. Thorax. Scutum covered with short, brown setae. Legs. Fore leg with tibia and first tarsomere equally long. Mid-tibia with 20 anterior, 4 posterodorsal, 7 posterior and 3 posteroventral bristles. Hind tibia with 8 anterodorsal, 6 posterodorsal and 4 posterior bristles. Wings. Vein r-m 2.1 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Exechia penicillata sp. n.
Female: Body length 3 mm. Wing length 2.5 mm. Colouration (Dry specimen). Head with vertex dark brown; face and clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, basal part of first segment yellow. Thorax with scutum brown; lateral sclerites brown to pale brown; propleura yellow; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-VI with lateral yellow areas. Terminalia brown. Head. Frons and vertex and clypeus covered with brown setae. Antenna 1.7 times as long as length from vertex to ventral margin of clypeus ; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with short dark brown setae. Legs. Mid-tibia with 22 anterior, 5 posterodorsal, 5 posterior and 2 posteroventral bristles. Hind tibia with 8 anterodorsal, 5 posterodorsal and 4 posterior bristles. Wings. Vein r-m 2.5 times longer than stem of M-fork. Abdomen. Tergites covered with brown setae. Terminalia (Fig.
Distinguished from E. sambai and E. ashleyi in having the dorsal gonostylus branch 2.8 times longer than broad, with apico-internal corner acuminate and apico-external corner right-angled (Fig.
From Latin penicillus, paint brush, relating to the shape and placement of setae on the internal lobe of the gonostylus, resembling a brush.
Afrotropical, South Africa (Fig.
Collected in mist-belt forest (1325 m a.s.l.).
Male: Body length 3.2 mm. Wing length 2.8 mm. Colouration (Dry specimen). Head, face and clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal half of first flagellomere pale. Thorax with scutum brown, anterior and lateral margin paler; lateral sclerites and propleura brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown with a pale area covering ventral and apicolateral margins of tergite II and ventral and basolateral margins of tergite III, in dorsal view appearing as a pale band narrowly broken medially. Terminalia yellow. Head. Frons, vertex and clypeus covered with pale setae. Antenna 1.8 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 1.05 times as long as first tarsomere. Mid-tibia with 26 anterior, 3 posterodorsal, 7 posterior and 2 posteroventral bristles. Hind tibia with 7 anterodorsal, 4 posterodorsal and 6 posterior bristles. Wings. Vein r-m 2.7 times longer than stem of M-fork. Abdomen. Tergites covered with long, pale setae. Terminalia (Fig.
Female: Unknown.
Distinguished from other species in the E. parva group in having the gonocoxal lobes extending almost without tapering, with only the basal tenth covered with setae (Fig.
From Latin rectus, straight and loba, lobe, relating to the shape of the gonocoxal lobes.
East Palaearctic, Japan (Fig.
Unknown.
Exechia repanda Johannsen, 1912: 73 ♂♀ (
Male (n = 3): Body length 3.0-3.2 mm. Wing length 2.4-2.6 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum pale brown to yellow; palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal half of first flagellomere pale. Thorax with scutum brown, sometimes anterior and lateral margin paler; lateral sclerites pale brown to brown; propleura yellow to pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown, tergites II-III slightly paler than the rest or sometimes a distinct pale area covering lateral part of tergite II and anterolateral part of tergite III. Terminalia yellow to pale brown. Head. Frons, vertex and clypeus covered with pale setae. Antenna 1.83-1.93 times as long as length from vertex to ventral margin of clypeus; flagellomeres slightly longer than broad, with sixth flagellomere 1.0-1.1 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia as long as first tarsomere. Mid-tibia with 20-22 anterior, 2-3 posterodorsal, 7 posterior and (n = 1) 1 posteroventral bristles. Hind tibia with 8-7 anterodorsal, 5 posterodorsal and 4 posterior bristles. Wings. Vein r-m 2.7-3 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Figs
Exechia repanda Johannsen, 1912
Female (n = 3): Body length 3.1-3.3 mm. Wing length 2.4-2.8 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum yellow to pale brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, basal part of first segment pale. Thorax with scutum dark brown, sometimes anterior and lateral margin paler; lateral sclerites brown; propleura yellow to brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown to dark brown, tergites II-VI with pale lateral areas extending dorsally at anterior fifth to third of tergites III and IV, forming two more or less distinct pale bands. Terminalia pale brown. Head. Frons, vertex and clypeus covered with pale setae. Antenna 1.7-1.8 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate or longer than broad, with sixth flagellomere 1.0-1.2 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 2) 1.1-1.13 as long as first tarsomere. Mid-tibia with (n = 1) 20 anterior, (n = 1) 3 posterodorsal and (n = 1) 5 posterior bristles. Hind tibia with (n = 1) 6 anterodorsal, (n = 1) 4 posterodorsal and (n = 1) 4 posterior bristles. Wings. Vein r-m 1.5-2.8 times longer than stem of M-fork. Abdomen. Tergites covered with pale to pale brown setae. Terminalia (Fig.
Distinguished from E. subrepanda in having the dorsal gonostylus branch only 2.3-2.4 times longer than broad and with the external row of setae covering one-third to half of the total dorsal gonostylus branch length (Fig.
Nearctic, Canada, USA (Fig.
Adults collected in grassland, savannah and wetland habitats.
The species is very close to E. neorepanda and E. subrepanda and species determination should be conducted with care or by use of DNA barcoding. It has also been reported from Chokotka, East Russia (
Holotype in rather poor condition - both wings and left legs mounted on slide. Antennae almost intact (one tip lost). Terminalia in fairly good condition - transferred to glycerine in microvial and associated with pinned specimen.
Exechia repandoides Caspers, 1984: 180 ♂ (
Male (n = 3): Body length (n = 2) 3.0-3.3 mm. Wing length 2.8 mm. Colouration (Dry specimen; n = 2). Head pale brown to brown; labellum and palpus yellow. Antenna yellow. Scutum pale brown to brown, anterolaterally distinctly paler; lateral sclerites yellow to pale brown; halteres yellow. Legs yellow to whitish-yellow. Abdomen pale brown to brown, tergite II slightly paler. Terminalia pale brown. Head. Frons and vertex and clypeus covered with pale setae. Antenna 1.8-1.95 times as long as length from vertex to ventral margin of clypeus; flagellomeres elongate, with sixth flagellomere 1.25 times longer than broad. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia 0.96-1.0 times as long as first tarsomere. Mid-tibia with 20-24 anterior, 3-4 posterodorsal, 7-8 posterior and 2-3 posteroventral bristles. Hind tibia (n = 2) with 7-8 anterodorsal, 4-6 posterodorsal and 3-5 posterior bristles. Wings. Vein r-m 2.0-2.3 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Exechia repandoides Caspers, 1984, male terminalia.
Female: Unknown.
Distinguished from E. sphaerata in having the dorsal gonostylus branch apico-internally forming a short angular projection (Fig.
West Palaearctic, Austria, Britain, Corsica, Czech Republic, Estonia, France, Germany, Hungary, Italy, Sweden, Switzerland (Fig.
Reared from fruitbody of Tricholoma sejunctum (Sowerby) Quél. and Cortinarius sp. (
The earlier records of this species from Jokkmokk, Sweden (
The pinned specimens have been preserved in alcohol for some years before being dry-mounted and may, therefore, be considerably paler compared to fresh material.
Holotype compared with and found to be conspecific with TSZD-JKJ-207679.
Exechia rohdendorfi Zaitzev, 1996: 68 ♂ (
Male (n = 4): Body length 3.4-3.6 mm. Wing length 3-3.1 mm. Colouration (Dry specimen). Head brown to dark brown; face yellow to brown; clypeus pale brown; labellum yellow; palpus yellow. Antenna with scape and pedicel yellow; flagellum yellow to brown, first segment with yellow base. Scutum pale brown with a narrow yellow anterolateral margin; lateral sclerites pale brown; propleura whitish-yellow; halteres whitish-yellow, apically slightly darker. Legs whitish-yellow. Abdomen pale brown to brown, tergite III sometimes with a small yellow lateral spot. Terminalia pale brown with MD and apico-internal lobe of DB dark brown (Fig.
Exechia rohdendorfi Zaitzev, 1996, male terminalia.
Female: Unknown.
Distinguished from E. toyoheii in having the hypandrium with apical pair of setae elongate (Fig.
East Palaearctic, Japan, Russia (Fig.
Unknown
We have recognised one specimen (TSZD-JKJ-108464) with slightly deviating shape of the medial and internal gonostylus branch (Fig.
We have studied the holotype, based on images of the terminalia, provided by courtesy of Dr. Andrey Ozerov, Moscow State University. The holotype is in good condition and terminalia not dissected.
Male (n = 2): Body length 3.3-3.4 mm. Wing length 2.8-2.9 mm. Colouration (Dry specimen). Head and face brown; clypeus pale brown; labellum yellow; palpus yellow to whitish-yellow. Antenna with scape and pedicel yellow; flagellum pale brown to brown, basal half of first flagellomere pale. Thorax with scutum pale brown, anterior and lateral margin slightly paler; lateral sclerites and propleura pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown. Terminalia yellow. Head. Frons and vertex covered with pale setae. Clypeus covered with 13-16 pale setae, evenly distributed. Antenna long, 2.0-2.25 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, with sixth flagellomere 1.2-1.3 times as long as wide. Thorax. Scutum covered with short, pale setae. Legs. Fore leg with tibia 0.91-0.95 times as long as first tarsomere. Mid-tibia with 22-23 anterior, 3-5 posterodorsal, 5-7 posterior and 2-4 posteroventral bristles. Hind tibia with 6-8 anterodorsal, 4 posterodorsal and 3-5 posterior bristles. Wings. Vein r-m 2.75-3 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Exechia sambai sp. n.
Female: Body length 3.5 mm. Wing length 3.2 mm. Colouration (Dry specimen). Head, face and clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal part of first flagellomere pale. Thorax with scutum, lateral sclerites and propleura pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown, tergites I-IV with pale lateral areas. Terminalia yellow. Head. Frons and vertex covered with short pale setae. Clypeus covered with pale setae, evenly distributed. Antenna 2 times as long as length from vertex to ventral margin of clypeus. Thorax. Scutum covered with short pale setae. Legs. Fore leg with tibia 0.92 times as long as first tarsomere. Mid-tibia with 24 anterior, 4 posterodorsal, 8 posterior and 3 posteroventral bristles. Hind tibia with 8 anterodorsal, 5 posterodorsal and 4 posterior bristles. Wings. Vein r-m 2.9 times longer than stem of M-fork. Abdomen. Tergites covered with dark brown setae. Terminalia (Fig.
Distinguished from E. penicillata in having the dorsal gonostylus branch 2 times longer than broad and with the apico-external corner extended into a rounded lobe (Fig.
Named in honour of the contemporary artist and painter Chéri Samba, relating to the shape of the internal lobe of the gonostylus, resembling a paintbrush.
Afrotropical, Kenya (Fig.
Adult collected in bamboo forest (3050 m a.s.l.)
Male: Body length 3.5 mm. Wing length 3.2 mm. Colouration (Dry specimen). Head, face and clypeus brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown. Scutum and lateral sclerites pale brown; propleura yellow; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-III with a lateral pale area. Terminalia yellow. Head. Frons and vertex covered with short pale setae. Clypeus covered with pale setae. Antenna 2 times as long as length from vertex to ventral margin of clypeus; flagellomeres longer than broad, with sixth flagellomere 1.4 times as long as wide. Thorax. Scutum covered with short, pale setae. Legs. Fore leg with tibia 0.78 times as long as first tarsomere. Mid-tibia with 18 anterior, 4 posterodorsal, 10 posterior and 4 posteroventral bristles. Hind tibia with 8 anterodorsal, 3 posterodorsal and 4 posterior bristles. Wings. Vein r-m 3.7 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. columna in having the gonocoxal lobes not reaching beyond the apical hypandrial setae (Fig.
From Latin serra, saw, relating to the shape of the medial lobe of the gonostylus.
Oriental, Nepal (Fig.
Adult collected in the eastern Himalayas (2500 m a.s.l.).
Male (n = 12): Body length 2.9-3.2 mm. Wing length 2.4-2-5 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum dark brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum dark brown, basal part of first flagellomere pale. Thorax with scutum, lateral sclerites and propleura dark brown; halteres whitish-yellow. Legs yellow. Abdomen dark brown. Terminalia brown. Head. Vertex, frons and clypeus covered with pale setae. Antenna 1.78 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with pale setae. Legs (n = 3). Fore leg with tibia as long as first tarsomere. Mid-tibia with 20 anterior, 3 posterodorsal, 4 posterior and 2 posteroventral bristles. Hind tibia with 6 anterodorsal, 5 posterodorsal and 3 posterior bristles. Wings. Vein r-m 2.8 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (n = 4, Fig.
Female: Unknown.
Distinguished from E. longilobata in having the dorsal gonostylus branch apically spathulate and about 3.5 times longer than broad (Fig.
From Latin spatula, flat piece, relating to the the apical shape of the dorsal branch of the gonostylus.
West Palaearctic, Norway, Sweden (Fig.
Unknown.
Male (n = 8): Body length 2.7-3.6 mm. Wing length 2.3-2.9 mm. Colouration (Dry specimen). Head brown to dark brown; face and clypeus dark brown; labellum yellow to pale brown; palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown to dark brown. Thorax with scutum and lateral sclerites brown; propleura brown to dark brown; halteres whitish-yellow to yellow. Legs whitish-yellow to yellow. Abdomen dark brown, sometimes with a pale area covering basolateral part of tergite III. Terminalia yellow. Head. Frons, vertex and clypeus covered with pale setae. Antenna (n = 6) 1.8-2.0 times as long as length from vertex to ventral margin of clypeus; flagellomeres slightly longer than broad, with sixth flagellomere (n = 6) 1.0-1.1 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 5) 1.0-1.05 times as long as first tarsomere. Mid-tibia with (n = 5) 16-19 anterior, (n = 5) 3-4 posterodorsal, (n = 5) 5-7 posterior and (n = 5) 1-3 posteroventral bristles. Hind tibia with 7-9 anterodorsal, 4 posterodorsal and 3-6 posterior bristles. Wings. Vein r-m (n = 6) 2.1-2.4 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Exechia sphaerata sp. n.
Female (n =) : Body length 2.9-3.7 mm. (n = 3) Wing length 2.5-2.9 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown. Thorax with scutum and propleura brown; lateral sclerites dark brown; halteres whitish-yellow. Legs yellow. Abdomen brown, tergites III-VI with lateral pale areas. Terminalia yellow. Head. Frons, vertex and clypeus covered with pale setae. Antenna (n = 3) 1.75-1.9 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate, with sixth flagellomere as long as broad. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 4) 1.0-1.05 times as long as first tarsomere. Mid-tibia with (n = 3) 16-19 anterior, (n = 1) 4 posterodorsal, (n = 1) 5 posterior and (n = 1) 2 posteroventral bristles. Hind tibia with (n = 3) 7-9 anterodorsal, (n = 3) 4-5 posterodorsal and (n = 3) 4-5 posterior bristles. Wings. Vein r-m (n = 3) 2.1-2.4 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Distinguished from E. repandoides in having the dorsal branch of the gonostylus with apex rounded (Fig.
From Latin sphaera, sphere, relating to the shape of the dorsal branch of the gonostylus.
Nearctic, Canada (Fig.
Adults collected in different types of forest and wetland habitats.
Male (n = 3): Body length 3.0-3.2 mm. Wing length 2.3-2.6 mm. Colouration (Dry specimen). Head brown to dark brown; face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown to brown, basal part of first flagellomere pale. Scutum and lateral sclerites brown; propleura pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen brown. Terminalia yellow to pale brown. Head. Vertex, frons and clypeus covered with pale setae. Antenna 1.55-1.7 times as long as length from vertex to ventral margin of clypeus; flagellomeres quadrate or longer than broad, with sixth flagellomere 1.0-1.2 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 1) as long as first tarsomere. Mid-tibia with (n = 2) 15-20 anterior, (n = 2) 3-4 posterodorsal, (n = 2) 6 posterior and (n = 1) 1 posteroventral bristles. Hind tibia with (n = 2) 5-7 anterodorsal, 4 posterodorsal and 3 posterior bristles. Wings. Vein r-m (n = 2) 2.2-2.7 times longer than stem of M-fork. Abdomen. Tergites covered with pale setae. Terminalia (Fig.
Exechia subrepanda sp. n.
Female (n = 2): Body length 2.6-3.2 mm. Wing length 2.2-2.6 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum pale brown, basal part of first segment pale. Scutum and lateral sclerites brown; propleura pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen pale brown to brown, tergites III-VI with yellow lateral areas extending dorsally at anterior tenth to eighth of tergites III and IV, forming two more or less distinct pale bands. Terminalia yellow. Head. Vertex, frons and clypeus covered with pale setae. Antenna 1.65-1.67 times as long as length from vertex to ventral margin of clypeus; flagellomeres slightly longer than broad, with sixth flagellomere 1.1-1.2 times as long as wide. Thorax. Scutum covered with pale setae. Legs. Fore leg with tibia (n = 1) as long as first tarsomere. Mid-tibia with (n = 1) 16 anterior, (n = 1) 3 posterodorsal and 4-5 posterior bristles. Hind tibia with (n = 1) 7 anterodorsal, (n = 1) 5 posterodorsal and (n = 1) 4 posterior bristles. Wings. Vein r-m 2-2.7 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Distinguished from E. repanda in having the dorsal gonostylus branch 2.4-2.7 times longer than broad and with external row of setae covering only one-fifth to one-fourth of the total dorsal gonostylus branch length (Fig.
Named after the species Exechia repanda, with Latin prefix sub-, below, relating to the close resemblance to E. repanda.
Nearctic, Canada (Fig.
Unknown
The species is very close to E. neorepanda and E. repanda and species determination should be conducted with care or with aid of DNA barcoding.
Male (n = 9): Body length (n = 8) 3.5-3.9 mm. Wing length 2.9-3.4 mm. Colouration (Dry specimen). Head dark brown; face and clypeus dark brown to pale brown; labellum yellow, sometimes pale brown; palpus yellow to pale brown, segments 4 and 5 usually somewhat darker. Antenna with scape and pedicel yellow; flagellum dark brown to pale brown, basal part of first flagellomere yellow. Thorax with scutum dark brown to brown, anterolateral margin paler; lateral sclerites pale brown to dark brown; propleura pale brown to brown; halteres whitish-yellow. Legs whitish-yellow to yellow, coxa sometimes slightly darker. Abdomen dark brown, tergites II-III with a yellow laterodorsal area. Terminalia brown with MB and internal lobe of DB dark brown (Fig.
Female: Unknown.
Distinguished from E. rohdendorfi in having the hypandrium with a short apical pair of setae (Fig.
Named in honour of Professor Emeritus Toyohei Saigusa, the collector of two of the paratypes, who also provided us with the loan of other invaluable material of several species described in this revision.
East Palaearctic, Japan (Fig.
Unknown
Male: Wing length 3.3 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum yellow; palpus yellow with segments 4 and 5 pale brown. Antenna with scape and pedicel yellow; flagellum dark brown, basal half of first flagellomere yellow. Scutum dark brown with yellow anterolateral margin; lateral sclerites and propleura brown; halteres whitish-yellow. Legs yellow. Abdomen dark brown, tergites II-III with a yellow laterodorsal area. Terminalia brown with MB dark brown (Fig.
Exechia trunciseta sp. n., male terminalia.
Female: Unknown
Distinguished from E. crassiseta by the shape of the medial gonostylus branch (Fig.
From Latin truncatus, truncated and seta, bristle, relating to the shape of the seta on posteroventral margin of the gonocoxites and on apical margin of the dorsal lobe of the gonostylus.
Oriental, Nepal (2700 m a.s.l., Fig.
Unknown.
Male: Body length 4.1 mm. Wing length 3.4 mm. Colouration (Dry specimen). Head, face and clypeus dark brown; labellum and palpus yellow. Antenna with scape and pedicel yellow; flagellum brown, first segment with yellow base. Thorax with scutum dark brown, lateral margin broadly yellow; lateral sclerites brown; propleura pale brown; halteres whitish-yellow. Legs whitish-yellow. Abdomen dark brown, tergites II-III with lateroventral yellow area. Terminalia pale brown. Head. Frons and vertex covered with pale setae. Clypeus covered with pale brown setae. Antenna long, 2.3 times as long as length from vertex to ventral margin of clypeus; flagellomeres slightly longer than broad, with sixth flagellomere 1.1 times as long as wide. Thorax. Scutum covered with short pale brown setae. Legs. Fore leg with tibia 0.89 times as long as first tarsomere. Mid-tibia with 26 anterior, 4 posterodorsal, 10 posterior and 3 posteroventral bristles. Hind tibia with 12 anterodorsal, 6 posterodorsal and 5 posterior bristles. Wings. Vein r-m 2.8 times longer than stem of M-fork. Abdomen. Tergites covered with pale brown setae. Terminalia (Fig.
Female: Unknown.
Distinguished from E. arcuata in having the gonocoxal lobe straight (Fig.
From KwaZulu-Natal, the Province where the holotype was collected, with Latin suffix -ensis, belonging to.
Afrotropical, South Africa (Fig.
Collected in mistbelt forest (1325 m a.s.l.).
Males Note that couplet 21 can be reached from both couplet 14 and 20. E. longichaeta has not been included in the key due our lack of material of the species, as well as our inability to distinguish this species from E. pararepanda, based on published illustrations. |
||
1 | Wings with dark spots (Fig. |
E. bifasciata |
– | Wings without spots. Each cercus kidney-shaped (Fig. |
2 |
2 | In ventral view, GLs entirely setose and not evenly tapered (Figs |
3 |
– | In ventral view, GLs with apical part more or less bare (Fig. |
5 |
3 | Hypandrium extended distally, forming a large medial lobe that reaches far beyond apicoventral gonocoxal margin, apically with two stout setae (Fig. |
4 |
– | Hypandrium not forming a lobe, with four stout setae (Fig. |
E. brachiata |
4 | GL reaching beyond apical hypandrial setae (Fig. |
E. columna |
– | GL shorter, not reaching beyond apical hypandrial setae (Fig. |
E. serrae |
5 | MB with a short thumb-like process with two small setae placed on the apex (Fig. |
E. chirotheca |
– | MB not as described above. DB at most apically emarginate, not bifurcate. | 6 |
6 | MB relatively dark (Figs |
7 |
– | MB not dark (Fig. |
10 |
7 | DB with a large and usually darkened internal lobe extending in interior direction (Fig. |
8 |
– | DB with a much shorter, not darkened lobe, located closer to the base (Fig. |
9 |
8 | DB apically with several stout setae (Fig. |
E. toyoheii |
– | DB apically with at most one or two stout setae (Fig. |
E. rohdendorfi |
9 | MB as in Fig. |
E. crassiseta |
– | MB as in Fig. |
E. trunciseta |
10 | IB with 3 or 4 apical setae gathered together on or close to the apex (Figs |
11 |
– | IB with one seta on or close to the apex and a pair or row of 2-3 setae placed some distance from the apical seta (Figs |
16 |
11 | GL entirely covered with setae, with entire length tapering (Fig. |
E. afroparva |
– | GL with at least some of the length bare and not tapering (Figs |
12 |
12 | GL with two apical setae (Figs |
13 |
– | GL with 3-4 apical setae (Fig. |
14 |
13 | DB about 2 times longer than broad; apico-external corner distinctly protruding, forming a rounded lobe (Fig. |
E. sambai |
– | DB about 2.8 times longer than broad; apico-external corner right-angled or virtually so, not distinctly protruding (Fig. |
E. penicillata |
14 | Apical hypandrial setae about as long as DB (Figs |
15 |
– | DB almost twice as long as the apical hypandrial setae or longer (Figs |
21 |
15 | GLs with apical setae splaying (Fig. |
E. burundiensis |
– | GLs with apical setae parallel (Fig. |
E. ashleyi |
16 | GL entirely covered with setae, with entire length tapering (Figs |
17 |
– | GL with at least some of the length bare and not tapering (Fig. |
19 |
17 | GL very short (Fig. |
E. cinctiformis |
– | GL longer (Figs |
18 |
18 | DB round with a short apical lobe (Fig. |
E. parva |
– | DB elongate, apical margin emarginate (Fig. |
E. adenaparva |
19 | Apical hypandrial setae reaching about as far as or beyond GLs (Figs |
20 |
– | Apical hypandrial setae not reaching beyond three-fourths of the GLs (Figs |
22 |
20 | DB acute, 2.2-2.5 times longer than broad (Fig. |
E. pararepanda |
– | DB spathulate, about 3.3 times longer than broad or longer (Fig. |
21 |
21 | GL short and slender, distinctly curved interiorly, apical setae very short (Fig. |
E. arcuata |
– | GL stouter, more or less straight (Fig. |
E. zuluensis |
22 | DB with a distinctly protruding external margin, forming a short process, bearing row of setae (Fig. |
E. capillata |
– | DB with external margin less protruding (Figs |
23 |
23 | DB short, at most 2.1 times longer than broad, if apical lobe distinct, then at most 0.2 of total DB length (Figs |
24 |
– | DB more elongate, at least 2.2 times longer than broad, always with a well defined apical lobe with length at least 0.22 of total DB length (Figs |
27 |
24 | DB strongly curved interiorly, without a distinct apical lobe (Figs |
25 |
– | DB only slightly curved interiorly, with a well defined apical lobe (Figs |
26 |
25 | DB apically acute (Fig. |
E. repandoides |
– | DB apically rounded, without apicoventral projection (Fig. |
E. sphaerata |
26 | DB short and round, 1.3-1.5 times longer than broad, apical lobe broader than long (Fig. |
E. breviflagellata |
– | DB more elongate, 1.8-2.0 times longer than broad, apical lobe longer than broad (Fig. |
E. brevilobata |
27 | DB at least 3 times longer than broad (Figs |
28 |
– | DB shorter (Figs |
29 |
28 | DB spathulate, about 3.5 times longer than broad (Fig. |
E. spatulata |
– | DB not spathulate, apex acute or somewhat rounded, about 3.1 times longer than broad (Fig. |
E. longilobata |
29 | GL almost without a tapered basal part, virtually bare (Fig. |
E. rectiloba |
– | GL with a distinctly tapered basal part, at least basal fourth covered with setae (as in Fig. |
30 |
30 | DB with external margin evenly rounded, without a distinct angle; external row of setae covering one-third to half of total DB length (Fig. |
E. neorepanda |
– | DB external margin with a distinct angle (Fig. |
31 |
31 | DB about 2.23 times longer than broad; external margin with somewhat protruding angle; external row of setae only about one sixth of total DB length (Fig. |
E. curvata |
– | DB at least 2.3 times longer than broad, external margin of DB with angle not protruding (Figs |
32 |
32 | DB 2.3-2.4 times longer than broad; external row of setae covering one-third to half of total DB length (Fig. |
E. repanda |
– | DB 2.4-2.6 times longer than broad; external row of setae covering one-fifth to one-fourth of total DB length (Fig. |
E. subrepanda |
Females |
||
1 | Sternite VIII with two large bare areas expanding into basal part of setae cover, lateral margins straight and parallel, cleft shallow and somewhat v-shaped (Fig. |
E. sambai |
– | Sternite VIII without bare area extending into seta cover, lateral margins rounded (e.g. Fig. |
2 |
2 | Gonapophysis IX with basolateral part expanding almost in a right angle (Fig. |
E. penicillata |
– | Gonapophysis IX with basolateral part expanding in a relatively obtuse angle (e.g. Fig. |
3 |
3 | Spermathecal eminence in ventral view trifurcate with middle branch large and round and lateral branches short (Fig. |
E. parva |
– | Spermathecal eminence in ventral view cross-shaped (Fig. |
4 |
4 | Tergite VIII with apicolateral margin forming a distinctly-protruding angle (Fig. |
5 |
– | Tergite VIII with apicolateral margin forming an obtuse angle or virtually straight (Fig. |
6 |
5 | Sternite VIII with cleft shallow, depth about 0.25 times the length between the apical lobes (Fig. |
E. curvata |
– | Sternite VIII with cleft deeper, depth about 0.5 times the length between the apical lobes (Fig. |
E. sphaerata |
6 | Sternite VIII with cleft shallow and v-shaped (Fig. |
E. breviflagellata |
– | Sternite VIII with cleft not v-shaped, usually deeper (Figs |
7 |
7 | Spermathecal eminence in ventral view unbranched (Fig. |
E. capillata |
– | Spermathecal eminence in ventral view cross-shaped (as in Fig. |
8 |
8 | If pale areas on abdomen distinct, then pale bands of tergites III and IV covering anterior fifth to third in dorsal view. Nearctic. | E. repanda |
– | If pale areas on abdomen distinct, then pale bands narrower, covering anterior tenth to eighth in dorsal view or broken medially. Nearctic. | E. subrepanda |
Of the 33 species included in this study, we were able to obtain DNA barcodes for 20 species. The remaining 12 species were delimited, based on morphology alone. DNA barcodes from two species, E. ashleyi and E. burundiensis, were not evaluated for inclusion into BINs as the sequence lengths were shorter than 500 bp. DNA barcodes from the remaining 18 species were divided into 16 BINs, of which 14 were congruent with our species delimitations; two BINs, however, were incrongruent as each included two species as delimited on morphology.
Genetic distances between species in the CO1 sequence data ranged from 0.012 between E. brevilobata and E. breviflagellata and 0.148 between E. arcuata and E. rohdendorfi, with an average of 0.076 ± 2SD [0.015-0.137] (Suppl. material
The Bayesian tree, based on the CO1 data, indicates a monophyletic and well-supported E. parva group (Fig.
Bayesian tree of 20 species in the E. parva group, based on mitochondrial CO1 sequences. Ingroup clades are coloured according to zoogeographical origin of their respective terminal branches. Outgroup branches and tip labels indicated in grey. Posterior probabilities (≥ 0.95) indicated at the nodes. Terminal branches labelled with BOLD process IDs as voucher codes. Clades (indicated with bars) and singletons constituting species are labelled with species names. Scale = 0.01 divergence in the CO1 sequence.
The species treated in this study can mostly be delimited, based on distinct characters in the male terminalia. In some cases, however, the differences are less apparent. Most notably is the close similarities amongst the four species, E. curvata, E. neorepanda, E. repanda and E. subrepanda, which above all can be distinguished, based on slight allometric variations in the shape of the dorsal branch of the gonostylus (Figs
In two cases, the molecular species delimitation, based on the BIN system, were not consistent with our delimitations. In both cases, two species ended up within a single BIN, where the BIN exhibits an Holarctic distribution, while the two included species each represent a Nearctic and a West Palaearctic distribution (Fig.
Our results fall in line with the general perception that female Exechia are more challenging than males to distinguish at the species level. Female terminalia have frequently been illustrated in earlier works (
Intercontinental barriers appear to have a strong impact on species segregation in the E. parva group, as each of the species is not found in more than a single zoogeographical region. Although two BINs in the E. parva group were shared between the West Palaearctic and Nearctic Regions, the intergrative solution in both cases was to treat them as different species. Similar patterns are known from other genera in the family (e.g.
We have further recognised several complexes in the group that are more or less congruent with the zoogeographic divisions. E. rohdendorfi, E. toyoheii, E. crassiseta and E. trunciseta can be clustered, based on the medial gonostylus branch darkened and with at least three elongated apical setae (from couplet 7, male key). This group can further be divided into two distinct clusters, based on the shape of the dorsal branch of the gonostylus and the setae on the ventral margin of each gonocoxite. The first cluster consisting of E. rohdendorfi and E. toyoheii (from couplet 8, male key), has an east Palaearctic distribution, with a centre in Japan where both species are collected (Fig.
The Afrotropical species, E. afroparva, E. sambai, E. penicillata, E. burundiensis and E. ashleyi, although highly variable in the male terminalia, can be clustered, based on the internal branch of the gonostylus apically with 3 or 4 apical setae gathered together on or close to the apex (from couplet 11, male key). The Bayesian tree suggests that also E. arcuata and E. zuluensis are part of this cluster, which is not monophyletic, but constitute two clades forming a paraphyletic assemblage (Fig.
All species keying out in the male key from couplets 17 and 22, including most of the previously-described members of the E. parva group, occur in the Holarctic Region (Figs
Additionally, the species E. columna and E. serrae, both collected in the eastern Himalayas (Fig.
The biological information about the group is very scarce for many species partly due to the limited material available for, for example, phenological studies and partly to the fact that few rearing experiments or larval studies have been carried out. The studied material is generally associated with forested, more or less damp and cool areas and, outside temperate and boreal regions (the Afrotropical and Oriental Regions), all material has been sampled from higher altitudes. As far as we know, most of the material has been collected in forest habitats, with the exception of E. longilobata, which is only known from a limestone quarry in Sweden. A connection to more open woodland with grasses and/or mires is noted for some of the northern species, especially late in the season when they seek umbelliferous stems for hibernation (see Introduction). What we know about host preferences, only based on records from the Holarctic species, E. parva, E. neorepanda, E. capillata and E. repandoides, indicates that the group attacks a wide range of fungi within Agaricales and occasionally also species of Russulales and Pezizales (for references, see the biology-sections below taxon treatments).
Some of the terms used in this article are used with a certain reservation, at least when considering homologous traits. According to
Additionally, the spermathecal eminence is here used to describe the sclerotised structure at the opening (gonopore) of the spermatheca (Figs
The sampling efforts in the different regions are highly divergent. Accordingly, most of the Afrotropical and Oriental species, described here, are based on singletons, i.e. only known from a single collecting event. Moreover, the geographical coverage in these Regions is very low, for example, the material from the Afrotropical Region is limited to five unique collecting events, leaving huge areas blank. Considering the low sampling effort in these Regions, one must assume that only a fraction of the total diversity has been recorded. Species representing other species groups in Exechia are well known from the Oriental and Afrotropical Regions (own materials,
We want to give special thanks the Canadian Center for DNA Barcoding (CCDB), to Prof. Emeritus Toyohei Saigusa and Kuyushu University Museum in Japan (KUEC) and to Ashley Kirk-Spriggs and the National Museum, Bloemfontein (BMSA), for their kind help and support and for granting us insect material on loan for this revision. We are grateful to Cornell University Insect Collection (CUIC), LUOMUS, Finnish Museum of Natrual History (MZHF) and Zoologische Staatssammlung München (SZMC) for help with and access to their type collections. We are also very grateful to Andrey Ozerov, Zoological Museum, Moscow State University, for imaging the type of E. rohdendorfi and translating the label text and Erica McAlister, Natural History Museum, London, for imaging the holotype of E. adenaparva. Further, we are indebted to Peter Chandler, Olavi Kurina and Jan Ševčík for reading through and commenting on the manuscript. Finally, many thanks to Peter Chandler for granting permission to reprint the figures of E. adenaparva and E. cinctiformis and to Janet Graham for permission to use her photo of the male terminalia of E. repandoides. Funding for DNA barcoding of Norwegian insect material came from the Norwegian Barcode of Life (NorBOL) and fieldwork was supported by the Norwegian Biodiversity Infomation Centre (NBIC). The publication charges for this article have been funded by a grant from the publication fund of UiT The Arctic University of Norway.
Genetic distances between species
Genetic distances within species
R scripts for analysing CO1 sequence data