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Citation: Vilkamaa, P.; Burdíková, N.;
Ševˇcík, J. The Genus Spinopygina gen.
nov. (Diptera, Sciaridae) from Western
North America: Preliminary Molecular
Phylogeny and Description of Seven
New Species. Insects 2023,14, 173.
https://doi.org/10.3390/insects
14020173
Academic Editor: Ding Yang
Received: 9 January 2023
Revised: 29 January 2023
Accepted: 4 February 2023
Published: 9 February 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
insects
Article
The Genus Spinopygina gen. nov. (Diptera, Sciaridae) from Western
North America: Preliminary Molecular Phylogeny and Description
of Seven New Species
Pekka Vilkamaa 1,* , Nikola Burdíková2and Jan Ševˇcík2,3
1
Finnish Museum of Natural History, Zoology Unit, University of Helsinki, P.O. Box 17, FI-00014 Helsinki, Finland
2Department of Biology and Ecology, Faculty of Science, University of Ostrava, Chittussiho 10,
CZ-710 00 Ostrava, Czech Republic
3Silesian Museum, NádražníOkruh 31, CZ-74601 Opava, Czech Republic
*Correspondence: pekka.vilkamaa@helsinki.fi
Simple Summary:
Black-winged fungus gnats (family Sciaridae) are one of the most species-rich
groups of flies (Diptera), with about 3000 described species worldwide. Their taxonomy is notori-
ously challenging, as many genera are difficult to delimit and species identification relies almost
exclusively on the male genitalia. Existing phylogenetic hypotheses based on DNA sequences are
often contradictory, presumably due to low taxon or gene sampling. In this paper, a new multigene
phylogenetic analysis is presented to support the concept and monophyly of a new genus, containing
eight species, seven of them being described as new to science.
Abstract:
The genus Spinopygina gen. nov. (type species Camptochaeta uniceps Hippa & Vilkamaa,
1994) from western North America is described. The genus includes the following eight species:
Spinopygina acerfalx sp. nov.; S. aurifera sp. nov.; S. camura sp. nov.; S. edura sp. nov.; S. peltata
sp. nov.; S. plena sp. nov.; S. quadracantha sp. nov.; and S. uniceps (Hippa & Vilkamaa, 1994) comb.
nov., transferred from Corynoptera Winnertz. The new species are described and Spinopygina uniceps
is re-diagnosed. The species are keyed and illustrated. In the maximum-likelihood phylogenetic
hypothesis based on four gene fragments (28S, 18S, 16S and COI), Spinopygina gen. nov. appears as
the sister group of Claustropyga Hippa, Vilkamaa & Mohrig, 2003. In the same analysis, a remarkable,
undescribed species is placed within Camptochaeta Hippa & Vilkamaa clade.
Keywords: Sciaridae; black-winged fungus gnats; new taxa; phylogeny; Nearctic Region
1. Introduction
Sciaridae (black-winged fungus gnats) is one of the little studied terrestrial Dipteran
families with about 3000 described species in about 100 genera and subgenera, occurring
in all continents [
1
]. Most species live in shady forested and moist habitats. Larvae are
predominantly saprophagous and live in soil and litter or under bark of dead wood. The
taxonomy is notoriously challenging, as many genera are difficult to delimit and are rich
in small and rather similar species. The species identification relies almost exclusively
on the male hypopygium, and in most genera the females are unidentifiable. Some trials
to reconstruct phylogenies extensively over the family or focused on a limited number
of taxa have been made, either using morphological characters [
1
–
8
] or using molecular
markers
[9–12]
. Often the hypotheses obtained have been contradictory, presumably
due to variable choice of morphological characters, insufficient number of molecular
characters used or because of low taxon sampling. The paper by Shin et al. [
9
] is the
most comprehensive of the molecular analyses, but it misses many crucial genera from the
Holarctic region and lacks almost completely the extra-Holarctic taxa in its taxon sampling.
Shin et al. [9] divided the family into three subfamilies, and later [10] named a fourth, but
Insects 2023,14, 173. https://doi.org/10.3390/insects14020173 https://www.mdpi.com/journal/insects
Insects 2023,14, 173 2 of 21
the large and morphologically very diverse group of genera (the ‘Pseudolycoriella group’)
appeared non-monophyletic and was not ranked as a subfamily.
Of the Holarctic fauna of Sciaridae, the Nearctic region is much less studied than
the Palaearctic fauna. Mohrig et al. [
13
] in their revision listed 166 valid species from the
Nearctic region (excluding Greenland), and subsequent authors [
14
–
26
] have added 70 new
species,making a total of 236 known species in the Nearctic to date. The number of known
Palaearctic species is much higher, about 2000. Based on the COI (barcode) gene, Hebert
et al. [
27
] counted 2200 different BINs for Sciaridae from Canada, but proper taxonomic
work is waiting to identify and name the taxa. Other biogeographic regions are even less
studied than the Holarctic, but studies made on different genera indicate that the taxonomic
richness of Sciaridae in the tropics is enormous [2,28–35].
Hippa and Vilkamaa [
36
] described Camptochaeta uniceps from Canada (British Columbia)
and included the species in their new genus Camptochaeta because it has the lambda-shaped
basomedial sclerotization in the gonostylus, regarded as a synapomorphy of Camptochaeta.
Subsequently, Menzel and Mohrig [
1
] in their revision of the Palaearctic Sciaridae did
not accept the concept of Camptochaeta, but transferred two of the included species into
Keilbachia
Mohrig, 1987 and all other species lacking the apical tooth of the gonostylus into
the Corynoptera parvula group and into the C. spinifera group of the large but obviously non-
monophyletic Corynoptera Winnertz, 1867. Later, the Nearctic Camptochaeta uniceps Hippa
&Vilkamaa, 1994 was transferred to the Corynoptera spinifera group by Mohrig et al. [13].
Here, in this study, we report on several species undoubtedly related to C. uniceps
from western North America. We were able to study the species complex in more detail,
describe the new species and consider the phylogenetic position and the taxonomic status
of the group. Simultaneously, the taxonomic placement of a new species of Sciaridae
(Camptochaeta sp., voucher SCI87) with an unusual tegmen was searched for.
2. Materials and Methods
2.1. Morphological Methods
The material originated from Malaise trap samples, and all specimens were detected
and picked out from unsorted sciarid or insect material stored in ethanol. The holotype of
Camptochaeta uniceps was obtained from Canadian National Collection, Ottawa, Canada
(CNC) and a paratype from Royal British Columbia Museum, Victoria, Canada (RBCM),
the other paratypes having been deposited in Zoological Museum, Finnish Museum of
Natural History. The specimens were mounted on microscope slides in Euparal, after
dehydrating them in absolute ethanol. Only males were studied: the females are unknown.
The terminology and methods of measuring structures follow Hippa and Vilkamaa [
28
]
and Hippa et al. [
3
,
37
]. The photographs of the slide-mounted specimens were taken with
a Leica MC170 HD camera mounted on Leica DM 4000 B LED research microscope. The
habitus of Spinopygina uniceps was photographed with Canon EOS 5DS digital SLR camera
with a Canon MP-E 65 mm macro lens. The Figures were processed with Photoshop version
CS5, CorelDraw2017 and CorelPhotopaint2017. The type material of the new species is
deposited in Zoological Museum, Finnish Museum of Natural History, Helsinki, Finland
(MZH). The specimen data of the MZH specimens will be added into the specimen database
of the Finnish Museum of Natural History (Luomus) and will be found with the specimen
codes https://luomus/GE.####.
This published work and the nomenclatural acts it contains have been registered in
ZooBank, the online registration system for the ICZN. The LSID for this publication is:
urn:lsid:zoobank.org:pub:9962DD66-13CC-4191-B6FF-0A7DD1934F75.
2.2. Molecular Methods
For the ingroup, representatives of taxa from the three subfamilies of Sciaridae were
chosen as well as selected terminals from the non-monophyletic ‘Pseudolycoriella group’
of genera proposed by Shin et al. [
9
,
10
]. From the latter group, terminals were chosen to
adequately cover the morphological diversity of the group.
Insects 2023,14, 173 3 of 21
The specimens used for DNA analysis (Table 1) were alcohol-preserved (70% to 99.9%
ethanol). The DNA was extracted using NucleoSpin Tissue Kit (Macherey-Nagel, Düren,
Germany) following manufacturers’ protocols. PCRs were performed using primers listed
by Shin et al. [9].
Table 1.
List of specimens used for the phylogenetic analysis, with GenBank accession numbers. The
voucher codes are stated only in the newly sequenced species.
Species Voucher 16S 28S 18S COI
Bradysia hilaris JQ613922 JQ613726 JQ613630 JQ613822
Camptochaeta camptochaeta JQ613970 JQ613773 JQ613675 JQ613870
Camptochaeta exquisita SCI84 OQ024861 OQ024849 OQ024870 OQ024762
Camptochaeta mixta SCI89 OQ024864 OQ024852 OQ024873 OQ024765
Camptochaeta pellax SCI85 OQ024862 OQ024850 OQ024871 OQ024763
Camptochaeta sp. SCI87 OQ024863 OQ024851 OQ024872 OQ024764
Catotricha subobsoleta MG554124 MG554155 KP288784 KT316873
Chaetosciara umbalis JQ613905 JQ613709 JQ613613 JQ613805
Claustropyga abblanda MG554121 MG554146 MG554134 MG554165
Claustropyga brevichaeta SCI113 OQ024865 OQ024853 OQ024874 OQ024766
Claustropyga corticis SCI122 - - - OQ024769
SCI123 OQ024867 OQ024856 OQ024877 -
Claustropyga refrigerata N/A KU949253 N/A KU923133
Corynoptera blanda JQ613965 JQ613768 JQ613670 JQ613865
Corynoptera boletiphaga JQ613974 JQ613777 JQ613679 JQ613874
Corynoptera deserta KU949092 KU949259 N/A KU923138
Corynoptera obscuripila - - - MZ625500
Corynoptera spinifera BOLD N/A N/A N/A SCINO533-15
Corynoptera subdentata SCI118 OQ024866 OQ024855 OQ024876 OQ024768
Corynoptera subtilis JQ613978 JQ613781 JQ613682 JQ613878
Corynoptera verrucifera SCI114 N/A OQ024854 OQ024875 OQ024767
Cratyna vagabunda JQ613968 JQ613771 JQ613673 JQ613968
Cratyna ambigua JQ613929 JQ613733 JQ613637 JQ613829
Diadocidia ferruginosa MG554126 MG554157 KP288786 KC435634
Ditomyia fasciata MG554125 MG554156 MG554141 MG554168
Epidapus absconditus N/A KU949293 N/A KU923211
Epidapus atomarius JQ613971 JQ613774 JQ613676 JQ613871
Exechia fusca MG554126 MG554158 MG684611 MG684785
Hemineurina flavicornis JQ613885 JQ613689 JQ613593 JQ613786
Heterotricha takkae MG554128 MG554159 MG684612 MG684786
Keilbachia subacumina JQ613951 JQ613755 N/A JQ613851
Keroplatus testaceus MG554129 MG554160 KP288746 KT316834
Leptosciarella trochanterata JQ613941 JQ613745 JQ613648 JQ613841
Lycoriella ingenua JQ613901 JQ613705 JQ613609 JQ613802
Pseudolycoriella porotaka MK906478 MK906568 N/A MK906375
Scatopsciara atomaria JQ613973 JQ613776 JQ613678 JQ613873
Scatopsciara vagula JQ613967 JQ613770 JQ613672 JQ613867
Schwenckfeldina carbonaria MG554120 MG554145 MG554133 MG554164
Sciara helvola JQ613911 JQ613715 JQ613619 JQ613811
Spinopygina acerfalx SCI80 OQ024859 OQ024847 OQ024868 OQ024760
Spinopygina peltata SCI82 OQ024860 OQ024848 OQ024869 OQ024761
Trichosia splendens JQ613969 JQ613772 JQ613674 JQ613869
Xylosciara betulae JQ613963 JQ613766 JQ613668 JQ613863
Zygoneura sciarina JQ613909 JQ613713 JQ613617 JQ613809
All amplified products were purified using Gel/PCR DNA Fragments Extraction Kit
(Geneaid, New Taipei City, Taiwan) and subsequently, the samples were sequenced by
Eurofins (Hamburg, Germany). All sequences were assembled, manually inspected, and
primers trimmed in SeqTrace [
38
]. New sequences were deposited in the GenBank database,
with their accession numbers listed in Table 1.
Insects 2023,14, 173 4 of 21
All genes were aligned using MAFFT version 7 [
39
] on the MAFFT server (http://mafft.
cbrc.jp/alignment/server/). To remove all unreliably aligned regions, the GBLOCKS 0.91b
program [
40
] was used (http://phylogeny.lirmm.fr/phylo_cgi/one_task.cgi?task_type=
gblocks) with conditions set as follows: allow smaller blocks, allow gap positions within
the final blocks, allow less strict flanking positions and do not allow many contiguous
nonconserved positions. The final molecular dataset consists of 4287 bp, and the lengths of
individual alignments were: 18S = 1843 bp, 28S = 1308 bp, 16S = 478 bp, COI = 658 bp.
The final concatenated dataset was partitioned by gene and codon position and
subsequently analysed using ML method with IQtree [
41
]. Best-fitting substitution models
were chosen automatically by the IQ-TREE software (GTR+F+I+G4: 16S, TPM3u+F+I+G4:
18S, TVM+F+I+G4:28S, TIM3+F+G4: COI_1, TIM2+F+I+G4: COI_2, TIM3+F+I+G4: COI_3),
without free-rate heterogeneity. Branch supports were evaluated using 1000 ultrafast
bootstrap [
42
]. All other settings were left as default. The resulting phylogenetic tree
(consensus tree) was visualized using Interactive Tree of Life (iTOL; [43]).
3. Results
3.1. Desription of the New Genus
Spinopygina gen. nov.
Type species: Camptochaeta uniceps Hippa & Vilkamaa, 1994.
LSID urn:lsid:zoobank.org:act: DBFE5207-337A-4817-B955-14847009D23B.
Etymology.
The name is formed from the Latin word spine, thorn, and the Greek
word pyge, rump, referring to the outstanding megasetae of the gonostylus of the species.
Diagnosis.
Small to medium sized Sciaridae, wing length 1.6–2.2 mm. Antenna
long, flagellomeres with long setae and necks. Mouth parts small, maxillary palpus with
2 segments, 2nd segment much reduced. Anal lobe of wing small, halter with long stalk.
Body setosity dark, long and strong. Legs long and slender, fore tibial organ poorly
differentiated. Intergonocoxal area long or moderate, without medial lobe. Gonocoxae
united medially, setae at medial margins short. Gonostylus impressed or deeply excavated
medially, with lambda-shaped basomedial sclerotization; with strong megasetae with basal
bodies, including (in most species) strongly procurved basal megaseta; without apical tooth.
Tegmen modified, with flat apicolateral parts, without detectable aedeagal teeth.
Description.
Head (Figure 1). Normal, roundish. Eye bridge 1 to 2 facets wide.
Anterior vertex non-setose. Face setose. Clypeus with 1–2 setae or non-setose. Antennal
scape and pedicel normal. Antennal flagellum long, with 14 flagellomeres, bodies of
flagellomeres subcylindrical, with smooth surface, body of 4th flagellomere 2.25–3.20
×
as
long as subapically wide, necks longer than wide. Flagellomeral vestiture rather sparse,
setae longer than width of flagellomeres. Mouth parts small. Maxillary palpus with
2 segments, 2nd segment strongly reduced, variably between species and even specimens;
1st segment with 1, rarely with 2 sharp setae; with long sparse sensilla dorsally.
Thorax (Figure 2). Brown; setae dark. Acrostichal setae few anteriorly, dorsocentral
and lateral setae in indistinct rows of a few long and strong and short and fine setae.
Scutellum with 2 long and strong setae and a few short and fine setae. Posterior pronotum
non-setose. Anterior pronotum and prothoracic episternum with a few setae, other pleural
sclerites non-setose. Mesothoracic katepisternum high with oblique anterior margin.
Wing (Figure 3A–C). Fumose. Length 1.6–2.2 mm. Anal lobe small. Veins distinct,
except stM. Membrane non-setose, veins C, R and R1 setose, R5 with only dorsal setae, bM,
r-m, stM, M and stCuA non-setose. c/w 0.70–0.90, R1/R 0.70–1.15, R1 joining C well before
level of base of M-fork. Halter yellow, with long stalk.
Insects 2023,14, 173 5 of 21
Insects 2023, 14, x FOR PEER REVIEW 5 of 25
Figure 1. (A). Head, frontal view (holotype MZH GE.1941 of Spinopygina plena sp. nov.). (B). Anten-
nal flagellomeres 3–4, frontal view (paratype MZH GE.1940 of S. peltata sp. nov.). (C). Antennal
flagellomeres 2–4, frontal view (paratype MZH GE.1936 of S. camura sp. nov.). (D). Mouth parts,
frontal view (holotype MZH GE.1932 of S. aurifera sp. nov.). (E). Mouth parts, frontal view, (holotype
MZH GE.1941 of S. plena sp. nov.). Scale bars for (A–C) = 0.1 mm, for (D,E) = 0.05 mm.
Thorax (Figure 2). Brown; setae dark. Acrostichal setae few anteriorly, dorsocentral
and lateral setae in indistinct rows of a few long and strong and short and fine setae. Scu-
tellum with 2 long and strong setae and a few short and fine setae. Posterior pronotum
non-setose. Anterior pronotum and prothoracic episternum with a few setae, other pleural
sclerites non-setose. Mesothoracic katepisternum high with oblique anterior margin.
Figure 1.
(
A
). Head, frontal view (holotype MZH GE.1941 of Spinopygina plena sp. nov.). (
B
). An-
tennal flagellomeres 3–4, frontal view (paratype MZH GE.1940 of S. peltata sp. nov.). (
C
). Antennal
flagellomeres 2–4, frontal view (paratype MZH GE.1936 of S. camura sp. nov.). (
D
). Mouth parts,
frontal view (holotype MZH GE.1932 of S. aurifera sp. nov.). (
E
). Mouth parts, frontal view, (holotype
MZH GE.1941 of S. plena sp. nov.). Scale bars for (A–C) = 0.1 mm, for (D,E) = 0.05 mm.
Insects 2023, 14, x FOR PEER REVIEW 6 of 25
Figure 2. (A). Habitus, ventral view, specimen MZH GE.1969 of Spinopygina uniceps (Hippa & Vilka-
maa; 1994). (B). Thorax, lateral view (paratype MZH GE.1931 of S. acerfalx sp. nov.). Scale bars for
(A) = 1.0 mm, for (B) = 0.50 mm.
Wing (Figure 3A–C). Fumose. Length 1.6–2.2 mm. Anal lobe small. Veins distinct,
except stM. Membrane non-setose, veins C, R and R1 setose, R5 with only dorsal setae,
bM, r-m, stM, M and stCuA non-setose. c/w 0.70–0.90, R1/R 0.70–1.15, R1 joining C well
before level of base of M-fork. Halter yellow, with long stalk.
Figure 2.
(
A
). Habitus, ventral view, specimen MZH GE.1969 of Spinopygina uniceps (Hippa & Vilka-
maa; 1994). (
B
). Thorax, lateral view (paratype MZH GE.1931 of S. acerfalx sp. nov.). Scale bars for
(A) = 1.0 mm, for (B) = 0.50 mm.
Insects 2023,14, 173 6 of 21
Insects 2023, 14, x FOR PEER REVIEW 7 of 25
Figure 3. (A–C). Wings, ventral view. (A). Paratype MZH GE.1929 of Spinopygina acerfalx sp. nov.
(B). Paratype MZH GE.1933 of S. aurifera sp. nov. (C). Paratype MZH GE.1981 of S. plena sp. nov.
(D). Foreleg, prolateral view (holotype MZH GE.1938 of S. peltata sp. nov.) (E). Apical part of fore
tibia, prolateral view (paratype MZH GE.1942 of S. plena sp. nov. Scale bars for (A–C) = 1.0 mm, for
(D) = 0.5 mm, for (E) = 0.05 mm.
Legs (Figure 3D,E). Yellow or pale brown, long, femora slender; setae dark; tibial
spurs 1:2:2, fore tibial spur as long as tibial width or slightly longer, middle and hind left
and right tibial spurs subequal in size, longer than tibial width. Fore tibia without spinose
setae, middle tibia rarely with 1 spinose seta, hind tibia with dorsal row of strong spinose
setae. The retrolateral apical setae of hind tibia fine. Fore tibial organ not impressed or
proximally bordered, with small patch of a few setae. Tarsi unmodified, long, length of
fore basitarsomere/length of fore tibia 0.47–0.63. Tarsal claws unmodified, without teeth.
Abdomen (Figures 4–9). Pale brown or yellowish, normal, slender; setae dark, long
and strong. Hypopygium brown or yellowish, as abdomen. Intergonocoxal area long or
moderate, without lobe(s). Gonocoxa unmodified, longer than gonostylus, gonocoxae
fused ventromedially; rather richly setose, setae at ventromedial margin short, medial
membrane non-setose; with 1 elongated seta ventrally and dorsally at apicomedial corner.
Gonostylus elongated, widest medially and distinctly narrowed towards apex or evenly
wide and slightly narrowed towards apex, strongly excavated or at least slightly
Figure 3.
(
A
–
C
). Wings, ventral view. (
A
). Paratype MZH GE.1929 of Spinopygina acerfalx sp. nov.
(
B
). Paratype MZH GE.1933 of S. aurifera sp. nov. (
C
). Paratype MZH GE.1981 of S. plena sp. nov.
(
D
). Foreleg, prolateral view (holotype MZH GE.1938 of S. peltata sp. nov.) (
E
). Apical part of fore
tibia, prolateral view (paratype MZH GE.1942 of S. plena sp. nov. Scale bars for (A–C) = 1.0 mm, for
(D) = 0.5 mm, for (E) = 0.05 mm.
Legs (Figure 3D,E). Yellow or pale brown, long, femora slender; setae dark; tibial spurs
1:2:2, fore tibial spur as long as tibial width or slightly longer, middle and hind left and
right tibial spurs subequal in size, longer than tibial width. Fore tibia without spinose
setae, middle tibia rarely with 1 spinose seta, hind tibia with dorsal row of strong spinose
setae. The retrolateral apical setae of hind tibia fine. Fore tibial organ not impressed or
proximally bordered, with small patch of a few setae. Tarsi unmodified, long, length of fore
basitarsomere/length of fore tibia 0.47–0.63. Tarsal claws unmodified, without teeth.
Abdomen (Figures 4–9). Pale brown or yellowish, normal, slender; setae dark, long
and strong. Hypopygium brown or yellowish, as abdomen. Intergonocoxal area long or
moderate, without lobe(s). Gonocoxa unmodified, longer than gonostylus, gonocoxae fused
ventromedially; rather richly setose, setae at ventromedial margin short, medial membrane
non-setose; with 1 elongated seta ventrally and dorsally at apicomedial corner. Gonostylus
elongated, widest medially and distinctly narrowed towards apex or evenly wide and
Insects 2023,14, 173 7 of 21
slightly narrowed towards apex, strongly excavated or at least slightly impressed medially,
with lambda-shaped basomedial sclerotization; without apical tooth; with apical, subapical
and medial megasetae, or only apical or subapical megasetae present; megasetae strong and
with distinct basal bodies, in some species basalmost megaseta distinctly procurved; with
some elongated setae medially and apically. Tegmen variable in shape, truncate or slightly
curved apically, slightly curved basolaterally, constricted subapically, membranous or with
sclerotized rim apically, with flat apicolateral parts; parameres sclerotized, approaching
each other and joining to sclerotized rim or ending at apex separately. Apodemes of tegmen
and aedeagal apodeme usually short, aedeagal teeth not detectable. Tergite 9 rather narrow,
with long and strong setae. Tergite 10 (cercus) normal.
Female unknown.
Distribution. USA: California, Oregon; Canada: British Columbia.
Insects 2023, 14, x FOR PEER REVIEW 8 of 25
impressed medially, with lambda-shaped basomedial sclerotization; without apical tooth;
with apical, subapical and medial megasetae, or only apical or subapical megasetae pre-
sent; megasetae strong and with distinct basal bodies, in some species basalmost megaseta
distinctly procurved; with some elongated setae medially and apically. Tegmen variable
in shape, truncate or slightly curved apically, slightly curved basolaterally, constricted
subapically, membranous or with sclerotized rim apically, with flat apicolateral parts;
parameres sclerotized, approaching each other and joining to sclerotized rim or ending at
apex separately. Apodemes of tegmen and aedeagal apodeme usually short, aedeagal
teeth not detectable. Tergite 9 rather narrow, with long and strong setae. Tergite 10 (cer-
cus) normal.
Female unknown.
Distribution. USA: California, Oregon; Canada: British Columbia.
Figure 4. Hypopygia, ventral view. (A). Holotype MZH GE.1928 of Spinopygina acerfalx sp. nov. (B).
Holotype MZH GE.1932 of S. aurifera sp. nov. Scale bars = 0.1 mm.
Figure 4.
Hypopygia, ventral view. (
A
). Holotype MZH GE.1928 of Spinopygina acerfalx sp. nov.
(B). Holotype MZH GE.1932 of S. aurifera sp. nov. Scale bars = 0.1 mm.
Insects 2023,14, 173 8 of 21
Insects 2023, 14, x FOR PEER REVIEW 9 of 25
Figure 5. Hypopygia, ventral view. (A). Holotype MZH GE.1935 of Spinopygina camura sp. nov. (B).
Holotype MZH GE.1937 of S. edura sp. nov. Scale bars = 0.1 mm.
Figure 5.
Hypopygia, ventral view. (
A
). Holotype MZH GE.1935 of Spinopygina camura sp. nov.
(B). Holotype MZH GE.1937 of S. edura sp. nov. Scale bars = 0.1 mm.
Insects 2023, 14, x FOR PEER REVIEW 10 of 25
Figure 6. Hypopygia, ventral view. (A). Holotype MZH GE.1938 of Spinopygina peltata sp. nov. (B).
Holotype MZH GE.1941 of S. plena sp. nov. Scale bars = 0.1 mm.
Figure 6.
Hypopygia, ventral view. (
A
). Holotype MZH GE.1938 of Spinopygina peltata sp. nov.
(B). Holotype MZH GE.1941 of S. plena sp. nov. Scale bars = 0.1 mm.
Insects 2023,14, 173 9 of 21
Insects 2023, 14, x FOR PEER REVIEW 11 of 25
Figure 7. Hypopygium of Spinopygina quadracantha sp. nov. (A). Ventral view (holotype MZH
GE.1944). (B). Dorsal view (paratype MZH GE.1956). Scale bars = 0.1 mm.
Figure 7.
Hypopygium of Spinopygina quadracantha sp. nov. (
A
). Ventral view (holotype MZH
GE.1944). (B). Dorsal view (paratype MZH GE.1956). Scale bars = 0.1 mm.
Insects 2023, 14, x FOR PEER REVIEW 12 of 25
Figure 8. Hypopygium of Spinopygina uniceps (Hippa & Vilkamaa, 1994). (A). Ventral view (holo-
type CNC). (B). Dorsal view (specimen MZH GE.1962). Scale bars = 0.1 mm.
Figure 8.
Hypopygium of Spinopygina uniceps (Hippa & Vilkamaa, 1994). (
A
). Ventral view (holotype
CNC). (B). Dorsal view (specimen MZH GE.1962). Scale bars = 0.1 mm.
Insects 2023,14, 173 10 of 21
Insects 2023, 14, x FOR PEER REVIEW 13 of 25
Figure 9. Tegmina, ventral view. (A). Holotype MZH GE.1928 of Spinopygina acerfalx sp. nov. (B).
Holotype MZH GE.1932 of S. aurifera sp. nov. (C). Paratype MZH GE.1936 of S. camura sp. nov. (D).
Holotype MZH GE.1937 of S. edura sp. nov. (E). Holotype MZH GE.1938 of S. peltata sp. nov. (F).
Holotype MZH GE.1941 of S. plena sp. nov. (G). Holotype MZH GE.1944 of S. quadracantha sp. nov.
(H). Holotype CNC of S. uniceps (Hippa & Vilkamaa, 1994) Scale bars = 0.05 mm.
Figure 9.
Tegmina, ventral view. (
A
). Holotype MZH GE.1928 of Spinopygina acerfalx sp. nov.
(
B
). Holotype MZH GE.1932 of S. aurifera sp. nov. (
C
). Paratype MZH GE.1936 of S. camura sp. nov.
(
D
). Holotype MZH GE.1937 of S. edura sp. nov. (
E
). Holotype MZH GE.1938 of S. peltata sp. nov.
(
F
). Holotype MZH GE.1941 of S. plena sp. nov. (
G
). Holotype MZH GE.1944 of S. quadracantha
sp. nov. (H). Holotype CNC of S. uniceps (Hippa & Vilkamaa, 1994) Scale bars = 0.05 mm.
Insects 2023,14, 173 11 of 21
Insects 2023, 14, x FOR PEER REVIEW 14 of 25
Figure 10. Maximum likelihood hypothesis for relationships among selected taxa of Sciaridae (Dip-
tera) based on DNA sequence data (18S, 28S, 16S, and COI), 4287 characters. Support numbers refer
to ultrafast bootstrap values (ufboot) over 50.
3.2. Comparative Diagnosis of Spinopygina gen. nov.
Mohrig et al. [13] transferred Camptochaeta uniceps Hippa &Vilkamaa, 1994 into the
Corynoptera spinifera group sensu [1]. Spinopygina gen. nov. resembles the C. spinifera group
in having a rather similar lambda-shaped basomedial sclerotized structure of the impressed
or excavated gonostylus, in lacking the apical tooth of gonostylus, in having the megasetae
Figure 10.
Maximum likelihood hypothesis for relationships among selected taxa of Sciaridae
(Diptera) based on DNA sequence data (18S, 28S, 16S, and COI), 4287 characters. Support numbers
refer to ultrafast bootstrap values (ufboot) over 50.
3.2. Comparative Diagnosis of Spinopygina gen. nov.
Mohrig et al. [
13
] transferred Camptochaeta uniceps Hippa &Vilkamaa, 1994 into the
Corynoptera spinifera group sensu [
1
]. Spinopygina gen. nov. resembles the C. spinifera group in
having a rather similar lambda-shaped basomedial sclerotized structure of the impressed or
excavated gonostylus, in lacking the apical tooth of gonostylus, in having the megasetae with
strong basal bodies, in having an apical gonostylar megaseta in all and a strongly curved
basal gonostylar megaseta in part of the species. Spinopygina differs from the C. spinifera
group as well as from the Corynoptera parvula group sensu [
1
] in important characters: the
long necks of antennal flagellomeres, the 2-segmented maxillary palpus without the sensory
pit, the long stalk of the halter, the indistinct, not impressed fore tibial organ, the basoven-
Insects 2023,14, 173 12 of 21
trally united gonocoxae with distinctly shortened setae at their ventromedial margins and
the tegmen with flat apicolateral parts and the apparent lack of the aedeagal teeth; see, e.g.,
Figures 4,5and 30A,B in Hippa & Vilkamaa (1994) [
36
]. Moreover, Spinopygina differs from
Camptochaeta Hippa &Vilkamaa, 1994 in the sense of [
1
] in the lack of the apical tooth of the
gonostylus in addition to the characters mentioned above.
Spinopygina resembles Claustropyga Hippa, Vilkamaa & Mohrig, 2003 in having the
maxillary palpus 2-segmented (in part of Claustropyga), the sensilla scattered on the dorsal
side of 1st segment of the maxillary palpus, in having a weakly modified fore tibial organ, in
having the medial part of the gonostylus impressed or excavated, in having distinct basal
bodies of the gonostylar megasetae and in having the megasetae strong at least in part of the
species, in having the gonocoxae united basoventrally with the intergonocoxal area long or
at least moderate. Spinopygina differs in having the basomedial part of gonostylus impressed
with the lambda-shaped sclerotized structure (the impression lacks in Claustropyga), in having
long flagellomeral necks and stalk of halter and in the lack of the apical tooth of gonostylus
(lacks also in one known species of Claustropyga). Moreover, even if long, the intergonocoxal
area of Spinopygina is not lobe-like produced as in most species of Claustropyga.
Spinopygina resembles Xylosciara Tuomikoski, 1960 in having a long intergonocoxal
area, in having strong gonostylar megasetae including apical ones and in having a poorly
differentiated patch of setae on the fore tibial organ. Spinopygina differs most strikingly in
having 2-segmented maxillary palpus, longer necks and setae of antennal flagellomeres, in
having strong and dark body vestiture and in having a modified tegmen and a strongly
hollowed gonostylus.
Some species of Spinopygina resemble Keilbachia Mohrig, 1987 in having a strong
procurved megaseta basally or subbasally on the gonostylus. Some species of Keilbachia
lack the apical tooth of gonostylus and some species have a strongly modified tegmen
reminiscent of that of Spinopygina, but Keilbachia has a 3-segmented maxillary palpus with
a sensory pit on the 1st palpal segment, the fore tibial organ in depression at least in part of
the species and it has ventromedially separated gonocoxae.
By the reduced maxillary palpus with sensilla scattered, long antennal flagellomeral
bodies and their necks, long and slender legs, the long and oblique mesothoracic katepis-
ternum, the strongly reduced anal lobe of wing and the poorly modified fore tibial organ,
Spinopygina resembles Epidapus Haliday, 1851 and in lacking the apical tooth of gonostylus,
especially Epidapus (Pseudoaptagogyna)absconditus group sensu [
1
]. Epidapus absconditus
(Vimmer, 1926) has, like Spinopygina, strong apical-subapical gonostylar megasetae with
distinct basal bodies. However, the megasetae of Epidapus absconditus are stout, straight
and symmetrically narrowed at apex, not long and curved as in Spinopygina. The four
megasetae of the related Epidapus quadrispinosus Mohrig & Mamaev, 1990 are all recurved.
Moreover, Spinopygina differs from the E. absconditus group in having subequal left and
right spurs on middle and hindtibia (unequal), in having the intergonocoxal area long
with the gonocoxae united basoventrally (short and separate), with shortened setae at the
ventromedial margin (equal), in having the tegmen strongly modified and with distinctly
sclerotized parameres (simple, laterally and apically roundish and completely membra-
nous tegmen in Pseudaptagogyna Vimmer, 1926) and in having the gonostylus impressed or
excavated (in Pseudoaptagogyna convex medially) and in having (in part of the species) a
curved basal megaseta in the gonostylus.
Spinopygina differs from all above-mentioned genera in having the tegmen with flat api-
colateral parts and in lacking the aedeagal teeth (the teeth not detectable using a compound
microscope.
3.3. Key to Species of Spinopygina gen. nov.
1. Gonostylus with 5 megasetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. camura sp. nov.
−
Gonostylus with 3–4 megasetae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2
2
Gonostylus with 4 megasetae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3
−
Gonostylus with 3 megasetae
. . . . . . . . . . . . . . . . . .
.
. . . . . . . . . . . . . . . . . .
.
. . .
. 7
Insects 2023,14, 173 13 of 21
3.
All megasetae at apical half of gonostylus, tegmen broad apically
. . .
.
. . . . . . . . . . . .
4
−
One megaseta at basal half of gonostylus, tegmen narrow apically
. . . . . .
.
. . .
. 5
4.
Gonostylus much wider dorsally than ventrally, tegmen with large roundish apicolat-
eral parts
. . .
.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.
. . . . . .
.
. . .
.
. . .
. .
S. plena sp. nov.
−
Gonostylus about as wide ventrally and dorsally, tegmen with small acuminate
apicolateral parts
... ... ... ... ...
.
. . .
.
. . .
.
. . .
.
. . .
.
. . .
.
. . .
.
S. peltata sp. nov.
5.
Tegmen distinctly constricted medially, roundish apically, parameres joining membra-
neous apex separately
. . . . . . . . . . . . . . . . . . . . . . . . . . .
.
. . .
.
. . .
. .
S. aurifera sp. nov.
−
Tegmen slightly constricted subapically, truncate apically, parameres joining short
sclerotized rim apically
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. 6
6.
Two medial megasetae at ventral margin of gonostylus, basalmost megaseta procurved
and with long basal body, arising from medial excavation
. . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . .
..
. . . . . . . . . . . . . . . . . . . . .
. . .
S. uniceps (Hippa& Vilkamaa, 1994)
−
One medial megaseta and basalmost megaseta at ventral margin of gonostylus,
basalmost megaseta recurved and with moderate basal body
. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . .
.
. . . . . . . . . . . . . . . . . . . . . . . . . . .
.
S. quadracantha sp. nov.
7.
Tegmen small, its length less than half of length of gonostylus, parameres separately
joining broad sclerotized rim apically, apex of tegmen largely sclerotized
. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. edura sp. nov.
−
Tegmen large, its length 2/3 of length of gonostylus, parameres united apically,
joining narrow sclerotized rim apically, apex of tegmen largely membraneous
. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. acerfalx sp. nov.
3.4. Descriptions of New Species
Spinopygina acerfalx sp. nov. Figures 2B, 3A, 4A and 9A.
LSID urn:lsid:zoobank.org:act: 008D0F6D-919A-4915-99A8-8AB979A3DAD2.
Comparative diagnosis.
By its tegmen, Spinopygina acerfalx sp. nov. is very similar to
Spinopygina camura sp. nov., S. quadracantha sp. nov. and S. uniceps (Hippa & Vilkamaa). All
have the tegmen narrowed subapically, with flat apicolateral parts and with the parameres
joining a shortly sclerotized rim apically. Spinopygina acerfalx is similar to S. edura sp.
nov. and differs from all above-mentioned species in having only three, not four or five,
gonostylar megasetae. Moreover, in the form of the gonostylus and in the arrangement of
the gonostylar megasetae Spinopygina acerfalx resembles S. edura, but the latter is distinct in
having the tegmen much smaller and more sclerotized basally and in having the parameres
wide apart, joining the broadly sclerotized rim separately at the apex.
Etymology.
The name is a Latin noun in apposition, formed from acer, sharp, and falx,
sickle, referring to the sharp and curved gonostylar megasetae.
Material examined.
Holotype: USA, Oregon,
♂
; Benton County; Corvallis; 1460 SW
Allen St.; 44.550860
◦
N, 121.270189
◦
W; 4 March–20 April 2015; S. Fitzgerald leg.; Malaise
trap; in Euparal; MZH GE.1928. Paratypes: USA, Oregon, 2
♂♂
; Same collection data
as for holotype; in Euparal; MZH GE.1929, GE.1945. Oregon, 1
♂
; same collection data
as for holotype but 5 April–3 June 2015; in Euparal; MZH GE.1976. Oregon, 2
♂♂
; same
collection data as for holotype but 11 December 2014–14 January 2015; used for DNA
extraction (samples No. SCI80, SCI81); in Euparal; MZH GE.1978, GE.1979. Oregon,
1
♂
; Benton County; 5 miles up Woods Creek Road from jct Highway 20; 44.544
◦
N,
123.50
◦
W; 25 April–15 May 2014; S. Fitzgerald leg.; Malaise trap; mixed forest; in Euparal;
MZH GE.1930. Oregon, 1
♂
; Benton County; Corvallis; 6.4 miles up Woods Creek Road
from jct Highway 20; 3 March–12 April 2015; Malaise trap; fir/alder/maple; in Euparal;
MZH GE.1946. Oregon, 1
♂
; Coos County; Seven Devils’ Road; 43.3109
◦
N, 124.3484
◦
W;
7 February–15 May 2016; E. Boyd leg; Malaise trap; in Euparal; MZH GE.1931.
Description.
Male. Head. Face and antenna uniformly brown, maxillary palpus pale
brown. Eye bridge 1–2 facets wide. Body of 4th antennal flagellomere 2.60–3.20
×
as long
Insects 2023,14, 173 14 of 21
as subapically wide, the neck longer than wide, the longest setae longer than the width of
flagellomere. Face with 5–9 dark long and short setae. Clypeus non-setose or with 1 dark
seta. Maxillary palpus with 2 segments; 2nd segment reduced; 1st segment with 1 sharp
seta, with large indistinct dorsal patch of sensilla, 2nd segment with 1–5 setae.
Thorax (Figure 2B). Brown; setae dark. Anterior pronotum with 2 setae. Prothoracic
episternum with 2–3 setae.
Wing (Figure 3A). Fumose. Length 1.8–2.0 mm. Anal lobe small. Width/length
0.40–0.45
. R1/R 0.75–0.85. c/w 0.75–0.80. stM slightly longer than M-fork, bM shorter than
or as long as r-m, stCuA shorter or as long as bM. bM and r-m non-setose. Halter yellow,
with long stalk.
Legs. Yellow, long. Fore femur slender. Fore tibial organ not impressed or proximally
bordered, with small patch of a few setae. Fore tibial spur longer than tibial width. Length
of fore basitarsomere/length of fore tibia 0.63.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 4A) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa longer than gonos-
tylus; setae rather short, shorter towards medial margin. Gonostylus elongated, curved,
the medial side strongly impressed; with a few elongated setae apically; without apical
tooth, with 3 megasetae, 1 apical in ventral position, 1 subapical in dorsal position and
1 medial in ventral position, the megasetae long and strong, slightly recurved, with long
basal bodies. Tegmen (Figure 9A) slightly shorter than wide, truncate apically, sharply
narrowed subapically; with parameres joining short sclerotized rim apically; with flat
apicolateral parts. Aedeagal apodeme short, aedeagal teeth not detectable.
Spinopygina aurifera sp. nov. Figures 1D, 3B, 4B and 9B.
LSID urn:lsid:zoobank.org:act: B9089867-2FA5-4A03-92E4-5C99E60BD592.
Comparative diagnosis.
In having the basal megaseta of the gonostylus strongly
procurved, Spinopygina aurifera sp. nov. resembles S. camura sp. nov. and S. uniceps (Hippa
and Vilkamaa) but by its form of the gonostylus and the tegmen, S. aurifera is not very
similar to any other known species of the genus: the apicalmost megaseta is shifted to
a subapical position and the two medial megasetae are in a lobe-like extension of the
ventromedial margin of the gonostylus. The tegmen is characteristic in being strongly
constricted at the middle, and in having the apex wide and roundish and in having the flat
apicolateral parts small and triangular. Spinopygina aurifera differs from all other species of
the genus in having a peculiar conical lobe apicodorsally in the gonocoxa.
Etymology.
The name is a Latin noun in apposition, from the words auris, ear, and the
suffix -fer, bearing, referring to the ear-like apicolateral parts of the tegmen.
Material examined.
Holotype: USA, Oregon,
♂
; Linn County; Hackleman Creek;
0.6 miles E of Tombstone Pass; 44.397501
◦
N, 122.131401
◦
E; 29 June–1 August 2016;
S. Fitzgerald leg; Malaise trap; in Euparal; MZH GE.1932. Paratypes: USA, Oregon,
2♂♂
; same collection data as for holotype; in Euparal; MZH GE.1933 and GE.1934. Oregon,
1♂; same collection data as for holotype; in Euparal; MZH GE.1970.
Description.
Male. Head. Face and antenna uniformly brown, maxillary palpus pale
brown. Eye bridge 1–2 facets wide. Body of 4th antennal flagellomere 2.25–2.55
×
as long
as subapically wide, the neck longer than wide, the longest setae longer than the width of
flagellomere. Face with 5–7 dark long and short setae. Clypeus non-setose or with 1 dark
seta. Maxillary palpus (Figure 1D) with 2 segments; 2nd segment reduced; 1st segment with
1(2) sharp setae, with an indistinct dorsal patch of sensilla, 2nd segment with 3–5 setae.
Thorax. Brown; setae dark. Anterior pronotum with 2 setae. Prothoracic episternum
with 2–4 setae.
Wing (Figure 3B). Fumose. Length 1.8–2.0 mm. Width/length 0.40–0.45. Anal lobe
small. R1/R 0.75–0.90. c/w 0.70–0.75. stM slightly longer than M-fork, bM longer than r-m,
stCuA as long as or shorter than bM. bM and r-m non-setose. Halter yellow, long.
Legs. Yellow, long. Fore femur slender. Fore tibial organ not impressed or proximally
bordered, with small patch of a few setae. Fore tibial spur as long as tibial width. Length of
fore basitarsomere/length of fore tibia 0.47.
Insects 2023,14, 173 15 of 21
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 4B) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa wide, slightly longer
than gonostylus; setosity normal, at medial margin shorter. Gonostylus elongated, slightly
curved, strongly excavated medially; with a few elongated setae apically and medially;
without apical tooth, with 4 megasetae, 1 subapical and 2 medial at ventral margin and
1 basally arising from medial excavation, the megasetae long and strong, nearly straight
except the basalmost one procurved, all with basal bodies. Tegmen (Figure 9B) longer than
wide, curved apically, constricted laterally, with parameres ending wide apart at apex; with
small triangular apicolateral parts. Aedeagal apodeme rather short, aedeagal teeth not
detectable.
Spinopygina camura sp. nov. Figures 1C, 5A and 9C.
LSID urn:lsid:zoobank.org:act: 144A1DDD-712D-4B81-862A-735709641561.
Comparative diagnosis.
Spinopygina camura sp. nov. is distinguished from all other
species of the genus in having five instead of three or four gonostylar megasetae, the
basalmost of which is strongly procurved (a paratype of S. quadracantha has five megase-
tae on one of its gonostyli) Spinopygina uniceps (Hippa and Vilkamaa) and S. aurifera sp.
nov. also have a procurved basal megaseta on their gonostylus but they have only four
megasetae. See also under S. acerfalx sp. nov. and S. quadracantha sp. nov.
Etymology.
The name is a Latin adjective, camura, curved, referring to the curved
basal megaseta of the gonostylus.
Material examined.
Holotype: USA, Oregon,
♂
; Benton County; 6.4 miles up Woods
Creek Road from jct Highway 20; 6 March–12 April 2015; S. Fitzgerald leg.; Malaise trap;
fir/alder/maple forest; in Euparal; MZH GE.1935. Paratype: USA, Oregon, 1
♂
; same
collection data as for holotype; in Euparal; MZH GE.1936.
Description.
Male. Head. Face and antenna uniformly brown, maxillary palpus
pale brown. Eye bridge 2 facets wide. Body of 4th antennal flagellomere (Figure 1C)
2.75–2.90
×
as long as subapically wide, the neck longer than wide, the longest setae longer
than the width of flagellomere. Face with 8–10 dark long and short setae. Clypeus with 1
dark seta. Maxillary palpus with 2 segments; 2nd segment reduced; 1st segment with 1
sharp seta, with an indistinct dorsal patch of sensilla, 2nd segment with 2–3 setae.
Thorax. Brown; setae dark. Anterior pronotum with 2 setae. Prothoracic episternum
with 2–4 setae.
Wing. Fumose. Length 2.0 mm. Width/length 0.40–0.45. Anal lobe small. R1/R 0.80.
c/w 0.80. stM slightly longer than M-fork, bM shorter than or as long as r-m, stCuA shorter
than bM. bM and r-m non-setose. Halter yellow, long.
Legs. Yellow, long. Fore femur slender. Fore tibial organ not impressed or proximally
bordered, with small patch of a few setae. Fore tibial spur longer than tibial width. Length
of fore basitarsomere/length of fore tibia 0.60.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 5A) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa longer than gonosty-
lus; setae rather short, at medial margin shorter. Gonostylus elongated, curved laterally,
strongly impressed medially; with a few elongated setae apically; without apical tooth,
with 5 megasetae, 1 apical in ventral position, 1 subapical in dorsal position, 2 medially
in ventral position and 1 basally arising from the medial excavation, megasetae long and
strong, slightly recurved, the basalmost megaseta strongly procurved, all with long basal
bodies. Tegmen (Figure 9C) slightly longer than wide, truncate apically, roundish later-
ally, with parameres joining to short sclerotized rim apically; with flat apicolateral parts.
Aedeagal apodeme short, aedeagal teeth not detectable.
Spinopygina edura sp. nov. Figures 5B and 9D.
LSID urn:lsid:zoobank.org:act: AE559992-0563-426C-B08C-7CEB91256997.
Etymology.
The name is a Latin adjective, edura, hard, referring to the strongly
sclerotized tegmen.
Comparative diagnosis.
By its gonostylus with a sharp apex and three megasetae
Spinopygina edura sp. nov. is very similar to S. acerfalx sp. nov. although the former has the
Insects 2023,14, 173 16 of 21
medial megaseta slightly closer to apex of the gonostylus. Spinopygina edura differs from all
the species of the genus in its small and more sclerotized tegmen with concave parameres
joining a broadly sclerotized rim apically.
Material examined.
Holotype: USA, Oregon,
♂
; Benton County; Corvallis; 1460 SW
Allen St.; 44.550860
◦
N, 121.270189
◦
W; ex larva, 19 October 2014, emerged 14 November
2014; S. Fitzgerald; in Euparal; MZH GE.1937.
Description.
Male. Head. Face and antenna uniformly brown, maxillary palpus pale
brown. Eye bridge 1–2 facets wide. Antennae in poor condition in the specimen studied,
the neck longer than wide. Face with 10 dark long and short setae. Clypeus with 1 dark
seta. Maxillary palpus with 2 segments; 2nd segment reduced; 1st segment with 1 sharp
seta, with an indistinct dorsal patch of sensilla, 2nd segment with 3–4 setae.
Thorax. Brown; setae dark. Anterior pronotum with 1 seta. Prothoracic episternum
with 3 setae.
Wing. Fumose. Length 2.2 mm. Width/length 0.40. Anal lobe small. R1/R 0.70.
c/w 0.80. stM slightly longer than M-fork, bM shorter than r-m, stCuA shorter than bM.
bM and r-m non-setose. Halter yellow, long.
Legs. Yellow, long. Fore femur slender. Fore tibial organ not impressed or proximally
bordered, with small patch of a few setae. Fore tibial spur slightly longer than tibial width.
Length of fore basitarsomere/length of fore tibia 0.60.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 5B) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa longer than gonosty-
lus; setae rather short, at medial margin shorter. Gonostylus elongated, curved, the medial
side impressed; with a few elongated setae apically; without apical tooth, with 3 megasetae,
1 at very apex in dorsal position, 1 subapically in ventral position and 1 medially in ventral
position; the megasetae long and strong, slightly recurved, with long basal bodies. Tegmen
(Figure 9D) slightly shorter than wide, truncate apically, concave laterally, with strong
basolateral sclerotizations, with concave parameres joining separately broad sclerotized
rim apically; with large flat apicolateral parts. Aedeagal apodeme short, aedeagal teeth not
detectable.
Spinopygina peltata sp. nov. Figures 1B, 3D, 6A and 9E.
LSID urn:lsid:zoobank.org:act: E7434F57-4965-4501-B26C-3A0C49D11302.
Comparative diagnosis.
In having four gonostylar megasetae all at the apical half
of half of the gonostylus and in having a wide tegmen, Spinopygina peltata sp. nov.
resembles
S. plena
sp. nov. but differs in having the tegmen constricted subapically and
with acuminate flat apicolateral parts. Moreover, Spinopygina plena has the dorsal side of its
gonostylus greatly enlarged.
Etymology.
The name is formed from the Latin word pelta, shield, referring to the
wide tegmen.
Material examined. Holotype: USA, Oregon,
♂
; Benton County; Corvallis; 1460 SW
Allen St.; 44.550860
◦
N, 121.270189
◦
W; 4 March–20 April 2015; S. Fitzgerald leg.; Malaise
trap; in Euparal; MZH GE.1938. Paratypes: USA, Oregon, 6
♂♂
; same collection data as for
holotype; MZH GE.1939, GE.1947–GE.1951. Oregon, 1
♂
; same collection data; in Euparal;
MZH GE.1971. Oregon, 1
♂
; same collection data but 11 December 2014–15 January 2015;
in Euparal; MZH GE.1980. Oregon, 2
♂♂
; same data as previous, used for DNA extraction
(samples No. SCI82, SCI83), MZH GE.1973, GE.1974. Oregon, 1
♂
; Coos County; Seven
Devils’ Road; 43.3109
◦
N, 124.3484
◦
W; 7 February–15 May 2016; E. Boyd leg; Malaise
trap; in Euparal; MZH GE.1940. Oregon, 1
♂
; Coos County; Charleston; Seven Devils’
Road; 43.3132
◦
N, 124.3485
◦
W; 7 February–15 May 2016; S. Fitzgerald leg; Malaise trap; in
Euparal; MZH GE.1972.
Description.
Male. Head. Face brown, darker than antenna, maxillary palpus pale
brown. Eye bridge 2 facets wide. Body of 4th antennal flagellomere 2.75–3.2
×
(Figure 1B)
as long as subapically wide, the neck longer than wide, the longest setae longer than the
width of flagellomere. Face with 5–9 dark long and short setae. Clypeus with 1 dark seta.
Insects 2023,14, 173 17 of 21
Maxillary palpus with 2 segments; 2nd segment reduced; 1st segment with 1–2 sharp setae,
with an indistinct dorsal patch of sensilla, 2nd segment with 2 setae.
Thorax. Brown; setae dark. Anterior pronotum with 1–2 setae. Prothoracic episternum
with 2–5 setae.
Wing. Fumose. Length 1.7–2.1 mm. Width/length 0.40–0.45. Anal lobe small. R1/R
0.80–1.15. c/w 0.75–0.90. stM slightly longer than M-fork, bM longer than or as long as r-m,
stCuA shorter than r-m. bM and r-m non-setose. Halter yellow, long.
Legs (Figure 3D). Yellow, long. Fore femur slender. Fore tibial organ not impressed or
proximally bordered, with small patch of a few setae. Fore tibial spur longer than tibial
width. Length of fore basitarsomere/length of fore tibia 0.58.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 6A) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa rather narrow, much
longer than gonostylus; setae rather short, at medial margin shorter. Gonostylus elongated,
rather straight, the medial side impressed; with a few elongated setae apically; without
apical tooth, with 4 megasetae, 1 at apex in dorsal position, 1 subapically in ventral position
and 2 slightly more mediad in dorsal and ventral positions, the megasetae long and strong,
nearly straight, with short basal bodies. Tegmen (Figure 9E) shorter than wide, hyaline
and slightly produced apically, roundish basolaterally and slightly constricted subapically,
with parameres ending separately wide apart at apex; with flat apicolateral parts. Aedeagal
apodeme long, aedeagal teeth not detectable.
Spinopygina plena sp. nov. Figure 1A,E, Figures 3E, 6B and 9F.
LSID urn:lsid:zoobank.org:act: 8F5CAF1D-108C-4008-A048-35CA415C8E19.
Comparative diagnosis.
Spinopygina plena sp. nov. resembles S. peltata sp. nov., see
under the latter.
Etymology.
The name is a Latin adjective, plena, plump, referring to the tumid gonos-
tylus.
Material examined.
Holotype: USA, California,
♂
; Los Angeles County; 9 km N of
La Canada; 34
◦
25
0
N, 118
◦
19
0
W; 900 m; 14 December 1994; B.V. Brown leg.; Malaise trap;
ravine with oak forest; in Euparal; MZH GE.1941. Paratypes: USA, California, 2
♂♂
; same
collection data as for holotype; in Euparal; MZH GE.1942, GE.1943. California, 1
♂
; same
collection data as for holotype but 20 January–2 February 1995; B.V. Brown leg.; in Euparal;
MZH GE.1981.
Description.
Male. Head (Figure 1A). Face brown, darker than antenna, maxillary
palpus pale brown. Eye bridge 2 facets wide. Body of 4th antennal flagellomere 2.3–2.9
×
as
long as subapically wide, the neck longer than wide, the longest setae longer than the
width of flagellomere. Face with 6–10 dark long and short setae. Clypeus non-setose or
with 1 dark seta. Maxillary palpus (Figure 1E) with 2 segments; 2nd segment reduced; 1st
segment with 1–2 sharp setae, with an indistinct dorsal patch of sensilla, 2nd segment with
1–2 setae.
Thorax. Brown; setae dark. Anterior pronotum with 2–4 setae. Prothoracic episternum
with 3–5 setae.
Wing. Fumose. Length 1.6–1.7 mm. Width/length 0.40. Anal lobe small. R1/R
0.80–0.95. c/w 0.75–0.80. stM slightly longer than M-fork, bM longer than r-m, stCuA
shorter than r-m. bM and r-m non-setose. Halter yellow, long.
Legs. Yellow, long. Fore femur slender. Fore tibial organ (Figure 3E) not impressed or
proximally bordered, with indistinct patch of a few setae. Fore tibial spur longer than tibial
width. Length of fore basitarsomere/length of fore tibia 0.50.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 6B) brown,
as abdomen. Intergonocoxal area long, with short setosity. Gonocoxa longer than gonos-
tylus; setosity rather sparse, at medial margin shorter. Gonostylus wide, curved laterally,
strongly impressed medially, distinctly lobe-like produced dorsomedially, with a few elon-
gated setae apically without apical tooth, with 4 megasetae, 1 at the very apex, 2 subapically,
1 medially, the megasetae rather long and strong, nearly straight, with distinct basal bodies.
Tegmen (Figure 9F) slightly shorter than wide, truncate apically, roundish laterally, with
Insects 2023,14, 173 18 of 21
lateral sclerotizations (parameres) joining wide apart at apex; with large flat apicolateral
parts. Aedeagal apodeme short, aedeagal teeth not detectable.
Spinopygina quadracantha sp. nov. Figures 7and 9G.
LSID urn:lsid:zoobank.org:act: E8ADE6BD-FF9F-4F09-931A-3DAD514FDCEA.
Comparative diagnosis.
In having all gonostylar megasetae long and slightly re-
curved, Spinopygina quadracantha sp. nov. resembles S. acerfalx sp. nov. and S. edura sp.
nov. but differs in having four instead of three megasetae. However, two of the studied
specimens have an additional procurved megaseta at the basal part in one of their gonostyli
(Figure 7B). These gonostyli with 5 megasetae are difficult to distinguish from those of
Spinopygina camura sp. nov. but the tegmen of S. quadracantha is bigger with larger apicolat-
eral flat parts. A large tegmen with convex parameres joining into a short sclerotized rim
apically is fairly similar to that of S. acerfalx and differs distinctly from that of S. edura sp.
nov. See also under the latter.
Etymology.
The name is a Latin noun in apposition, from the words quadrus, four, and
acantha, spine, referring to the four gonostylar megasetae.
Material examined.
Holotype: Canada, British Columbia,
♂
; Vancouver Island; Rocky
Point; 19 March 1995; N. Winchester leg.; Malaise trap; in Euparal; MZH GE.1944.
Paratypes: Canada, British Columbia, 7 ♂♂; same collection data as for holotype but
29 October 1995; MZH GE.1952 to GE.1958. British Columbia, 2
♂♂
; same collection data
as previous; in Euparal, MZH GE.1974, GE.1975. British Columbia, 2
♂♂
; same collection
data as previous but 15 January 1995; in Euparal; MZH GE.1959, GE.1960.
Description.
Male. Head. Face and antenna uniformly brown, maxillary palpus pale
brown. Eye bridge 2 facets wide. Body of 4th antennal flagellomere 2.40–3
×
as long as
subapically wide, the neck longer than wide, the longest setae longer than the width of
flagellomere. Face with 4–10 dark long and short setae. Clypeus with 1 dark seta. Maxillary
palpus with 2 segments; 2nd segment reduced; 1st segment with 1–2 sharp setae, with an
indistinct dorsal patch of sensilla, 2nd segment with 1–3 setae.
Thorax. Brown; setae dark. Anterior pronotum with 2 setae. Prothoracic episternum
with 3 setae.
Wing. Fumose. Length 1.8–2.1 mm. Width/length 0.40–0.45. Anal lobe small. R1/R
0.85–1.0. c/w 0.70–0.90. stM slightly longer, as long as or shorter than M-fork, bM longer
or shorter than r-m, stCuA shortest. bM and r-m non-setose. Halter yellow, long.
Legs. Yellow, long. Fore femur slender. Fore tibial organ with weak vestiture, forming
an indistinct patch of few setae. Fore tibial spur longer than tibial width. Length of fore
basitarsomere/length of fore tibia 0.60.
Abdomen. Pale brown; setae dark, long and strong. Hypopygium (Figure 7) brown, as
abdomen. Intergonocoxal area long, with short setosity. Gonocoxa longer than gonostylus;
setae rather short, at medial margin shorter. Gonostylus elongated, curved, strongly
impressed medially; with a few elongated setae apically and medially; without apical
tooth, with 4 (rarely 5) megasetae, 1 at the very apex, 1 subapically and 2 medially wide
apart, the megasetae long and strong, slightly recurved, with long basal bodies. Tegmen
(Figure 9D) longer than wide, truncate apically, constricted laterally, with parameres joining
to short sclerotized rim apically; with large flat apicolateral parts. Aedeagal apodeme short,
aedeagal teeth not detectable.
Spinopygina uniceps
(Hippa & Vilkamaa, 1994) comb. nov. Figures 2A, 8and 9H.
Camptochaeta uniceps Hippa & Vilkamaa, 1994: 56, Figure 30A,B.
Corynoptera uniceps Mohrig et al. (2013): 195.
Short redescription.
Male. Wing length 1.6–2.2 mm (Figure 2A). 1st segment of max-
illary palpus with 1 seta, 2nd segment of with 2–5 setae. Gonostylus (Figure 8) widest
medially, strongly narrowed towards apex, strongly excavated medially, apex with rather
short apical megaseta; with two medial megasetae at dorsomedial margin, basal megaseta
with long basal body arising from medial excavation. Tegmen (Figure 9H) with flat apico-
lateral parts, parameres joining short sclerotized rim apically.
Insects 2023,14, 173 19 of 21
Comparative diagnosis.
By its form of gonostylus with four megasetae and by
the tegmen with flat apicolateral parts and the parameres joining a short apical rim,
Spinopygina uniceps
(Hippa & Vilkamaa, 1994) most resembles S. quadracantha sp. nov.
but differs in having a shorter apical megaseta of the gonostylus, in having the basal
megaseta distinctly procurved and with a long basal body and placed in the medial exca-
vation instead at the ventromedial margin of the gonostylus. Spinopygina uniceps has the
apex of tegmen narrower and with smaller flat apicolateral parts (Figures 7and 8). See also
under Spinopygina acerfalx.
Material examined.
Holotype: Canada, British Columbia,
♂
; Vancouver Island; Upper
Carmanah Valley; 30 July 1991; N. Winchester leg.; Malaise trap; forest floor; in Euparal;
CNC. Paratypes. Canada, British Columbia, 3
♂♂
; same collection data as for holotype;
in Euparal; MZH GE.1116 to GE.1118. British Columbia, 1
♂
; same collection data as for
holotype; in Euparal; RBCM.
New material.
Canada, British Columbia, 3
♂♂
; Vancouver Island; Upper Carmanah
Valley; 10–29 September 1991; N. Winchester leg; Malaise trap, forest floor; in Euparal; MZH
GE.1961 to GE.1963. British Columbia, 5
♂♂
; same data as previous but
17–26 October 1991
;
in Euparal; MZH GE.1964 to GE.1968. British Columbia, 1
♂
; same collection data as
previous; in ethanol; MZH GE.1969. British Columbia, 1
♂
; same collection data as previous;
in Euparal; GE.1977.
3.5. Phylogeny
In general, the present hypothesis (Figure 10), based on the same gene markers, is
similar to the earlier published phylogenies [
9
,
10
,
12
]. The proposed subfamilies Sciarinae,
Chaetosciarinae, Cratyninae and Megalosphyinae appear with good supports, as well as
the polyphyletic Pseudolycoriella group of genera.
In this study, Spinopygina nested with Claustropyga as its sister group. As in the
analysis of [
12
], Claustropyga in its original sense [
3
] was not monophyletic, C. refrigerata
(Lengersdorf, 1930), in the earlier morphological cladistic analysis [
3
] appearing as the
sister group of all other species of Claustropyga, now appeared in the Megalosphyinae
clade. Moreover, Pseudolycoriella porotaka nested in the Cratyninae clade, although with
low support. The placement of this genus was different also in the previous molecular
analyses [
9
,
12
]. Morphologically, Pseudolycoriella is not very similar to any other genus of
Sciaridae.
The former placement [
13
] of S. uniceps into the Corynoptera spinifera group proposed
by Menzel and Mohrig (2000) [
1
] is not supported, and that group appears as polyphyletic.
Interestingly, the two analyzed species of the group, C. spinifera Tuomikoski, 1960 and
C. verrucifera
(Lengersdorf, 1952), included in Camptochaeta [
36
] and subsequently trans-
ferred into the C. spinifera group [
1
], in the present hypothesis appear in the Camptochaeta
clade with strong support. Moreover, the present hypothesis supports the close relationship
between Keilbachia and Camptochaeta. The consequent changes in the classification as well
as the description of the new species of Camptochaeta with an exceptional tegmen will
be done in another publication (Vilkamaa et al. in prep.) Morphological similarities and
differences in Spinopygina to Claustropyga,Corynoptera spinifera group and other relevant
taxa are discussed under Comparative diagnosis of Spinopygina.
4. Discussion
Spinopygina gen. nov. has the structure of its tegmen as a unique character. By the
combination of this and other characters, although the latter shared with many other genera,
the genus is easy to identify.
Spinopygina shares quite a few morphological characters with many other genera,
and a hypothesis of its phylogenetic position and of the sister group relationship between
Spinopygina and Claustropyga were only possible by analyzing molecular characters. Al-
though the molecular data were available from only two of the eight species, these species
Insects 2023,14, 173 20 of 21
represent two different morphological types of Spinopygina, and the monophyly of the eight
known species is well founded.
Judging from the few collection localities of the material for the present study, it is
highly probable that more species of the new genus will be found in the western Nearctic
region.
Regarding the whole sciarid fauna, the Nearctic region is far less known than the
Palaearctic fauna [
1
,
13
] and it is possible that more new genera remain to be discovered in
the region, especially in the less studied western parts.
Author Contributions:
Conceptualization, P.V. and J.Š.; methodology, P.V., J.Š. and N.B.; software,
N.B.; validation, P.V. and J.Š.; formal analysis, N.B.; investigation, P.V. and J.Š.; resources, P.V. and
J.Š.; data curation, P.V., J.Š. and N.B.; writing—original draft preparation, P.V., J.Š. and N.B.; writing—
review and editing, P.V., J.Š. and N.B.; visualization, P.V., J.Š. and N.B.; supervision, P.V. and J.Š.;
funding acquisition, P.V. and J.Š. All authors have read and agreed to the published version of the
manuscript.
Funding:
The research by J. Ševˇcík was supported by the Ministry of Culture of the Czech Republic
by institutional financing of long-term conceptual development of research institution (the Silesian
Museum, MK000100595).
Data Availability Statement:
The data presented in this study are available on request from the
corresponding author. The data are not publicly available due to ongoing research.
Acknowledgments:
Brian Brown (Los Angeles), Claudia Copley and Neville Winchester (Victoria),
Scott Fitzgerald (Corvallis) and Jukka Salmela (Rovaniemi) are thanked for material. Max Söderholm
(Helsinki) is thanked for Figure 2A and Pentti Halenius (Helsinki) for processing other photographs.
Conflicts of Interest:
The authors declare no conflict of interest. The funders had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or
in the decision to publish the results.
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