ZooKeys | 06! : 23-55 (202 | ) A peer-reviewed open-access journal
doi: 10.3897/zookeys.1061.49999 RESEARCH ARTICLE Z00Ke y S

https:/ / ZOO keys. pensoft.net Launched to accelerate biodiversity research

Tipula (Vestiplex) crane flies
(Diptera, Tipulidae) of Korea

Pavel Starkevich', Sigitas Podénas'’, Virginija Podéniené?’,

Sun-Jae Park?, A-Young Kim?

| Nature Research Centre, Akademijos str. 2, LT-08412 Vilnius, Lithuania 2. Life Sciences Centre of Vilnius
University, Sauletekio str. 7, LT-10257 Vilnius, Lithuania 3 Animal Resources Division, National Institute of
Biological Resources, Incheon 22689, South Korea

Corresponding author: Pavel Starkevich (pavel.starkevic@gmail.com)

Academic editor: C. Borkent | Received 9 January 2020 | Accepted 18 September 2020 | Published 30 September 2021
http://zoobank.org/ FFA31901-68EF-457D-A4BF-DC807033F6BF

Citation: Starkevich P, Podénas S, Podéniené V, Park S-J, Kim A-Y (2021) Tipula (Vestiplex) crane flies (Diptera,
Tipulidae) of Korea. ZooKeys 1061: 23-55. https://doi.org/10.3897/zookeys. 1061.49999

Abstract

The Korean species of Tipula (Vestiplex) Bezzi, 1924 crane flies are taxonomically revised. Five species are
recognized. Tipula (V.) coquillettiana Alexander, 1924, 7. (V.) kuwayamai Alexander, 1921, T’ (V.) tchukchi
Alexander, 1934, and 7) (V.) verecunda Alexander, 1924 are newly recorded from the Korean Peninsula,
and 7 (V.) serricauda Alexander, 1914 was previously recorded. The larva of 7’ (V) serricauda is described
and illustrated, and the larvae of the subgenus 7’ ( Vestiplex) are divided into four groups based on spiracu-
lar lobe morphology. An identification key, redescriptions, and illustrations of Korean 7. (Vestiplex) adults

and grouping of known larvae are presented.

Keywords

distribution, hypopygium, larva, new record, North Korea, ovipositor, South Korea, taxonomy

Introduction

Tipula Linnaeus, 1758 is the largest genus in the family Tipulidae with a worldwide
distribution, and it is divided into 41 subgenera. The subgenus 7. (Véestiplex) Bezzi,
1924 is a terrestrial group represented by 177 species and subspecies recorded from
Holarctic and Oriental regions (Oosterbroek, 2019). The highest diversity of this

Copyright Pavel Starkevich et al. 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.

24 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

group is documented in the Eastern Palaearctic (76 species) and Oriental (74 species)
regions (Oosterbroek 2019).

The first 7. (Vestiplex) crane flies from the Korean Peninsula were collected by A.M.
Yankovsky in 1938-1940. He lived and worked in the northern part of Korea. Only
one species, 7’ (V.) serricauda Alexander, 1914, had been recorded from the Korean
Peninsula (Starkevich et al. 2015).

The aim of this study was to document, redescribe, illustrate, and prepare keys for

all Korean 7’ (Vestiplex) species.

Material and methods

The specimen material examined in this paper (Table 1) was obtained from: the United
States National Museum (USNM), Smithsonian Institution, Washington, DC, USA;
the Snow Entomological Museum, University of Kansas (SEM), Lawrence, KS, USA;
the National Institute of Biological Resources (NIBR), Incheon, South Korea and the
Korea University (KU), Seoul, South Korea; the Academy of Natural Sciences of Drex-
el University (ANSP), Philadelphia, PA, USA; Zoological Museum of the Zoological
Institute of the Russian Academy of Sciences, St. Petersburg, Russia (ZIN), and the
Nature Research Centre (NRC), Vilnius, Lithuania.

Adult crane flies were collected by insect net and at lights. Some specimens were
preserved dry in envelopes in the field and later mounted in the laboratory on their
side on a paper point with legs generally surrounding the insect pin. The specimens are
pinned except when noted otherwise.

Adult specimens were studied with a Nikon SMZ800 stereomicroscope. Photo-
graphs were taken with an INFINITY-1 camera mounted on a Nikon Eclipse 200 ster-
eomicroscope and with a Canon EOS 80D camera mounted on an Olympus SZX10
dissecting microscope. Genitalia were studied after heating them in 10 percent NaOH
solution for 5—10 minutes and then preserved in microvials filled with glycerol at-
tached to the pin. All redescriptions and illustrations are based only on Korean mate-
rial, except when otherwise mentioned.

Two identical last instar larvae were collected by hand and one of them was left for rear-
ing. A female of 7’ (V.) serricauda emerged after 36 days. ‘The larva is preserved in 70% etha-
nol though the head capsule was slide-mounted in glycerin jelly with corresponding label.
The larva was studied with an Olympus SZX10 dissecting microscope with photographs
taken with a Canon EOS 80D digital camera fitted with a Canon MP-E 65 mm macro lens.

Collecting localities with approximate coordinates are summarized in Table 1, and
this was used to generate the geographical distribution maps (Figs 86-90). The identi-
fication key is based on morphological characters primarily observed in Korean speci-
mens, but in cases when females are unknown from Korea, characters were observed
from other specimens collected in other Asian countries.

Descriptive terminology of adults generally follows that of Cumming and Wood
(2017). The term appendage of ninth sternite is adopted from Mannheims (1963),

25

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26 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

the terms ventral lobe and dorsal lobe of appendage of ninth sternite are adopted from
Gelhaus (2005), the term gonocoxal fragment (= sclerites sp/ and sp2 (Neumann,
1958), = genital bridge (Dobrotworsky 1968)) for inner structure covered by ninth
tergite is adopted from Brodo (2017). Descriptive terminology of larva generally fol-
lows that of Gelhaus (1986) and Neugart et al. (2009).

The overall world distribution of species is given according to Oosterbroek (2019).

Taxonomy

Tipula (Vestiplex) Bezzi, 1924

Tipula (Vestiplex) Bezzi 1924: 230; Edwards 1931: 79; Alexander 1934: 396; 1935:
117; 1965: 355; Mannheims 1953: 116; Savchenko 1964: 132.

Type species. Tipula cisalpina Riedel, 1913.

Vestiplex was first proposed by Bezzi (1924) as a subgenus of the genus Tipula for
the type species 7. cisalpina Riedel, 1913, which was recorded from the Western Pal-
aearctic (Italy and Switzerland). No fossil species of 7! (Vestiplex) are described so far
and only Matthews and Telka (1997) mentioned ovipositors of possibly 7’ (Vestiplex)
females from Cape Deceit Formation in Western Alaska (1.8 MY old).

The world fauna of the subgenus 7! (Vestiplex) includes 177 recent species and
subspecies, which are distributed throughout the Nearctic, Palaearctic, and Oriental
regions (Oosterbroek 2019). The majority of the species are associated with mountain
systems (Pyrenees, Alps, Caucasus, and Himalayas) where adults are commonly found
at altitudes ranging from 700 to 2500 m and rarely up to 4500 m (Savchenko 1960).

Females belonging to subgenus 7! (Véestiplex) are characterized by the ovipositor
having powerful cerci, which are heavily sclerotized, and serrated along outer margin
(but smooth in several Asiatic species), and small to rudimentary hypovalva (Alexander
1935, 1965; Alexander and Byers 1981). The male genitalia are extremely polymorphic
(Savchenko 1964), typically with the ninth tergite forming a shallowly concave and
sclerotized saucer, with other species having their ninth tergite completely divided lon-
gitudinally by a pale membrane (Alexander 1935; Alexander and Byers 1981).

Just seven species have described larvae. Two are North American species, 7. (V)
arctica Curtis, 1835 and T. (V) platymera Walker, 1856 (Alexander 1920a, Gelhaus
1986), and five are European species, 7. (V.) excisa excisa Schummel, 1833, T. (V.) hor-
torum Linnaeus, 1758, T. (V.) nubeculosa Meigen, 1804, T. (V.) semivittata Savchenko,
1960, and 7. (V.) scripta Meigen, 1830 (Chiswell 1956; Theowald 1965; Savchenko
1986; Podeniene 2003; Lantsov 2003).

The immature stages develop in terrestrial habitats such as the uppermost layer of
soil under leaf or needle litter, or under mosses (Chiswell 1956; Rogers 1942; Theowald
1967; Teale and Gelhaus 1984; Lantsov 2003; Podeniene 2003). Larvae of T’ (Vestiplex)

are easily recognized because of a brown band separating the anus from the anal papillae.

Vestiplex crane flies of Korea 27

The lobes surrounding spiracular field are subconical with the lateral lobe situated
dorsolaterally. The sclerotization of the dorsal lobe varies depending on species, with some
species bearing a sclerite only on the basal part of the posterior surface, while in other
species the entire anterior and posterior surfaces are sclerotized. In this case the apex of the
dorsal lobe is sclerotized, pointed, and directed anteriorly. The lateral lobe may possess a
narrow and vertical sclerite, but it may be entirely absent in some species. The ventral lobe
is the smallest and trianglar. It varies from extensively sclerotized to possessing only a small
sclerite. Larvae have two pairs of short, blunt anal papillae. The length and macrosetal
arrangement is consistent on the dorsum and tergum among all known species. Short
microscopic hairs are arranged in transverse rows and cover most of the abdomen.

List of Korean Tipula (Vestiplex) crane flies

Tipula (Vestiplex) coquillettiana Alexander, 1924
Tipula (Vestiplex) kuwayamai Alexander, 1921
Tipula ( Vestiplex) serricauda Alexander, 1914
Tipula (Vestiplex) tchukchi Alexander, 1934
Tipula ( Vestiplex) verecunda Alexander, 1924

Key to Korean Tipula (Vestiplex) crane flies

Males
it Gonocoxite armed with a powerful black spine or bifurcate horn (Figs 3, 32)...2
= Gonocoutersimn ple wunarin edi (Pigs 16,71 oh sich s.e Ravers i Selals cossause Seb lban's 3
2 Flagellum bicolored with inconspicuous basal enlargement. Gonocoxite
with a strong black spine (Fig. 32). Ninth tergite divided by pale midline
CE eS cringe sac teiisetate te cear tea ietaese tos Neltteser Tipula (Vestiplex) serricauda
_ Flagellum dark brown with weak basal enlargement. Gonocoxite horn-shaped
with bifurcate margin (Fig. 3). Ninth tergite forming narrow sclerotised sau-
SERUM a secs that Soesteaths wahranttnenag ies. Tipula (Vestiplex) coquillettiana
6) Eighth sternite with long setae (Figs 67, 70 ). Ninth sternite ventrally with
median tubercle (Fig. 67). Appendage of ninth sternite present, finger-shaped
(Figs 74, 75). Ninth tergite divided by pale midline (Fig. 68) wee eee
a ers Re A Din Sib rly th RO ER Tipula (Vestiplex) verecunda
= Eighth sternite without long setae (Fig. 16). Ninth sternite without ventral
tubercle (Fig. 16). Appendage of ninth sternite absent. Ninth tergite forming
a Scletotised:saucert( Bigs: 1h 9G) cea waneseraetareracrninoterhad sot ete lt iSvi bagstall it 4
4 Size relatively small (body length 16.8 mm, wing length 17.1 mm). Wing
pattern indistinct, only weak darkening along vein CuA. Abdomen with me-
Pee V EES Ga bf Sree RRO Ae IR ome eee 2h, Fe aeely Tipula (Vestiplex) tchukchi
= Size large (body length 17.8-19.7 mm, wing length 19.8—22.9 mm). Wing
distinctly marbled. Abdomen without median stripe.........eeeeeeeeeseeeeeeeeeees

ts Ru Bel ik Be, Cece en Re GES Tipula (Vestiplex) kuwayamai

28 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

1 Wing well developed, extends beyond middle of abdomen... ee 2
= Wing reduced, not reaching middle of abdomen... eee eeeeseceeeeeeeeeeeees
LS Dia LAW Aeon doh te Ni Tipula (Vestiplex) coquillettiana
Os Wing pattern indistinct, only weak darkening along vein CuA. Cercus api-
cally with incision in addition to serrated border (Fig. 65) ......eeeeeeeeeseeeeeeees
Tepe SUES A ori de Set Nien Rho ee Tipula (Vestiplex) tchukchi
= Wing pattern distinctly marbled. Cercus without apical incision (Figs 11, 27,

<=) Antennal flagellum brownish-black. Cercus with smooth ventral margin (Fig. 80).
Hypovalva long, blade-shaped (Figs 81, 82).......... Tipula (Vestiplex) verecunda
- Antennal flagellum yellow or bicolored. Cercus with serrate ventral margin
(Fig. 27, 38). Hypovalva short, filamentous or plate-shaped (Figs 28, 39) ...4
4 Large species with wing length 24.5—26.1 mm. Antennal flagellum bicolored.
Hypovalva filamentous (Fig. 28)... Tipula (Vestiplex) kuwayamai
- Smaller species with wing length16.4-17.1 mm. Antennal flagellum var-
ies from yellow to bicolored. Hypovalva in the shape of a dark brown plate

Cite S9) Mosstunlas tices ached. ttgaee Main cataauce Tipula (Vestiplex) serricauda

Tipula (Vestiplex) coquillettiana Alexander, 1924
Figs 1-15, 86

Tipula coquillettiana Alexander 1924: 605; 1925: 91.
Tipula (Vestiplex) coquillettiana: Alexander 1934: 405; 1935: 118; Savchenko 1960:
172; 1964: 180; Oosterbroek and Theowald 1992: 154.

Type material examined. Holotype, male, RUSSIA, Odasam [Southern Sakhalin], 5
August 1922, Esaki; paratype, male (USNM).

Other examined material (Fig. 86). NORTH KOREA, 1 male, Seren Mts, alt.
3000 ft, 25 June 1938, Yankovsky (USNM); 2 males, alt. 3000 ft, 26 June 1938,
Yankovsky (USNM); 2 males, alt. 3000-4000 ft, 29-30 June 1938, Yankovsky
(USNM); 1 male, alt. 3500-5500 ft, 29-30 June 1938, Yankovsky (USNM); 3 males,
alt. 4000-5000 ft, 29-30 June 1938, Yankovsky (USNM); 1 male, alt. 5000 ft, 29-30
June 1938, Yankovsky (USNM); 2 females, alt. 4000 ft, 30 June 1938, Yankovsky
(USNM); 2 males, alt. 3500 ft, 5 July 1938, Yankovsky (USNM); 3 males, alt. 4500 ft,
5—6 July 1938, Yankovsky (USNM); 1 male, alt. 4000 ft, 10-11 July 1938, Yankovsky
(USNM); 1 male, alt. 5500 ft, 18 July 1938, Yankovsky (USNM); 1 male, Kankyo
Nando Puksu Pyaksan, alt. 6000 ft, 23 June 1939, Yankovsky (USNM); 1 male, alt.
6000 ft, 31 July 1939, Yankovsky (USNM); 1 male, alt. 5000 ft, 2 August 1939, Yank-
ovsky (USNM); 2 males, Chonsani, alt. 4000 ft, 2 June 1940, Yankovsky (USNM);
1 male, alt. 3000 ft, 8 June 1940, Yankovsky (USNM); 4 males, alt. 4500 ft, 20 June
1940, Yankovsky (USNM); 1 male, alt. 4000 ft, 23 June 1940, Yankovsky (USNM).

Vestiplex crane flies of Korea 29)

Diagnosis. Tipula (V.) coquillettiana can be easily recognized by the ninth tergite,
which forms a narrow, saucer-shaped plate, and the horn-shaped gonocoxite. The tip
of the gonocoxite is bifurcate. The female of this species has a gray, elongated abdomen
and greatly reduced wing. The cercus is straight with its tip narrowed and the ventral
margin and apical part of the dorsal margin distinctly serrated. The hypovalva is in the
shape of an elongated filament.

Male. Body length 17.1-21.3 mm, wing length 18.9—22.8 mm. General body
coloration brownish yellow.

Head. Vertex and occiput gray with dark median line. Rostrum yellowish, thinly
dusted with gray dorsally. Nasus distinct. Antenna 13-segmented, if bent backward
extending beyond wing base. Scape and pedicel yellow; first flagellar segment basally
yellow, distally brownish black; succeeding flagellar segments brownish black. Each
flagellomere, except first, with basal enlargement and small incision. Apical flagel-
lomere small, reduced. Verticils approximately as long as corresponding segments.
Palpus with first segment yellowish, second brownish yellow, and other segment
brownish black.

Thorax. Pronotum yellowish, thinly dusted with gray. Prescutum and presutural
scutum gray; stripes bluish gray bordered by brown. Postsutural scutum, scutellum,
and postnotum gray with bluish shade; all sclerites with dark, median line. Pleura
brownish, dusted with gray. Coxa gray; trochanter yellowish; femur brownish yel-
low with tip broadly darkened; tibia brownish yellow; tarsal segments dark brown.
Tarsal claw without tooth. Wing patterned with brown. Halter brownish yellow with
brown knob.

Abdomen. Yellow. First abdominal segment dusted with gray. Abdominal segments
2 and 3 yellow, 4 and 5 brownish-yellow, and remaining segments dark brown. Lateral
margin of tergites pale yellow. Dorsal median stripe pale, broadly interrupted.

Hypopygium. Brown (Fig. 1). Ninth tergite distally forming a narrow saucer-shaped
plate (Figs 1, 2). Tergal saucer posteriorly with broad median notch; lateral comer with
serrated and blackened margin, provided with setae. Anterior margin elevated into
narrow, blackened rim reaching lateral part of tergal saucer and terminating into acute
projection. Gonocoxite apically extended, with tip split into blackened beak pointed
caudad and rounded projection (Figs 1, 3). Outer gonostylus slightly curved, finger-
shaped with tip narrowed (Fig. 4). Inner gonostylus in the shape of a curved plate;
beak blackened, triangular; dorsal margin claw-shaped; middorsal edge extended and
serrated (Fig. 5). Gonocoxal fragment with flattened medial sclerites, fused into broad,
nearly triangular base (Fig. 6). Lateral sclerite flattened and bilobed. Adminiculum
boat-shaped, with tip and dorsal edge apically blackened (Fig. 7). Semen pump with
flattened central vesicle (Figs 8, 9). Compressor apodeme with round median inci-
sion, forming a 30° angle with posterior immovable apodeme. Posterior immovable
apodeme much longer than compressor apodeme, flattened, rounded apically. Anterior
immovable apodeme flattened, rounded. Intromittent organ tube-shaped, about twice
as long as semen pump, brownish-black. Distal part with preapical yellow membrane,
apex with pale median incision, lateral parts split, filament-shaped (Fig. 10).

30 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

9 10

Figures 1-10. Male terminalia of 7’ (Vestiplex) coquillettiana \ hypopygium, lateral view 2 ninth tergite,
dorsal view 3 left gonocoxite, lateral view 4 left outer gonostylus 5 left inner gonostylus, lateral view
6 gonocoxal fragment, dorsal view 7 adminiculum, lateral view 8 semen pump, dorsal view 9 semen pump
and intromittent organ, lateral view 10 distal part of intromittent organ, lateral view. Abbreviations: aia,
anterior immovable apodeme; ca, compressor apodeme; gcx, gonocoxite; ig, inner gonostylus; ls, lateral
sclerite of gonocoxal fragment; ms, medial sclerite of gonocoxal fragment; og, outer gonostylus; pia, poste-
rior immovable apodeme; t9, ninth tergite. Scale bars: 0.7 mm (1); 0.5 mm (2-9); 0.25 mm (10).

Vestiplex crane flies of Korea 31

_—t10

Figures | 1-15. Female terminalia of 77 (Vestiplex) coquillettiana \\ ovipositor, left lateral view 12 eighth
sternite with hypovalvae, ventral view 13 ninth sternite with furca, dorsal view 14 bursa copulatrix, dorsal
view 15 spermatheca, lateral view. Abbreviations: ap bc, anterior part of bursa copulatrix; bp sd, basal part
of spermathecal duct; c, cerci; f, furca; h, hypovalvae; s8, eighth sternite; s9, ninth sternite; t10, tenth
tergite. Scale bars: 0.8 mm (I 1), 0.5 mm (12-14).

Female. Body length 26.9-30.2 mm, wing length 4.7—-5.7 mm. Generally similar
to male, but with elongated and gray abdomen. Tergites and sternites with pale mar-
gins. Wing greatly reduced.

Female terminalia. Tenth tergite shining dark brown. Cercus brown, straight, as
long as tenth tergite, with tip narrowed; ventral margin and apical part of dorsal mar-
gin distinctly serrated (Fig. 11). Hypovalva elongated and filamentous (Fig. 12). Me-
dian incision between hypovalvae deeper than posterior margin of eighth sternite. Lat-

32 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

eral angle of eighth sternite sloping. Ninth sternite with lateral parts straight (Fig. 13).
Furca anteriorly narrowed, shaped posteriorly as broad membranous plate (Fig. 13).
Bursa copulatrix with spermathecal duct sclerotized at base, in shape of thickened,
curved stick (Fig. 14). Spermatheca broadened at base, pear-shaped (Fig. 15).

Known distribution. Russia, Kazakhstan, and Japan (Oosterbroek 2019). Re-
corded here for the first time from North Korea.

Tipula (Vestiplex) kuwayamai Alexander, 1921
Figs 16-29, 87

Tipula kuwayamai Alexander 1921: 130; 1925: 93.
Tipula (Vestiplex) kuwayamai: Alexander 1934: 405; 1935: 118; Savchenko 1964: 179;
Oosterbroek and Theowald 1992: 156.

Type material examined. Holotype, male, JAPAN, Maruyama, Sapporo, 1 June 1919,
Kuwayama (USNM).

Other examined material (Fig. 87). NORTH KOREA, 4 males, Ompo, 23 May
1937, Yankovsky (USNM); 1 female, alt. 500 ft, 2 May 1938, Yankovsky (USNM);
1 male, alt. 400 ft, 10 May 1938, Yankovsky (USNM); 1 male, Chonsani, alt. 4900
ft, 2 June 1940, Yankovsky (USNM); 1 female, alt. 3500 ft, 13 June 1940, Yank-
ovsky (USNM). SOUTH KOREA, 1 male, #8, Central National Forest, 18 miles
NE of Seoul, 28 May 1954, G.W. Byers (SEM); 1 male, Gyeongi-do, Pocheon-si,
Soheul-eup, Gwangneung Forest, 30 May 1961, Gyeong-suk Jeon (KU); 1 female,
Chungcheonnam-do, Danyang-gun, Danyang-eup, Mt Sobaeksan, 6 June 1981, K-S
Lee (KU); 1 female, Gyeonggi-do, Seongnam-si, Sangjeok-dong, Mt Cheongyesan, 4
May1984, In-suk Hyeon (KU); 1 female, Gyeonggi-do, Namyangju-si, Hwado-eup,
Mt Cheonmasan, 20 May 1984, Yeong-cheol Heo (KU); 1 male, 3 females, Geo-
chang-gun, Gajo-myeon, Suwol-ri, Mt Bigye, Gogyeonsa, 6~7 May 2000, S.B. Jeong,
IN0000297019, IN0000297023, IN0000296935, IN0000297021 (NIBR); 1 female,
S.H. Baek, IN0000296936 (NIBR); 1 female, J.S. Choi, IN0000297020 (NIBR); 1
female, Ulju-gun, Sangbuk-myeon, Doekhyeon-ri, Mt Gaji, Helkijang, 18-19 May
2001, Y.S. Kim, IN0000226477 (NIBR). CHINA, 1 male, Heilongjiang Province,
Hsiaoling, 20 May 1938, leg. Weymarn (USNM); 6 males, 2 females, Heilongjiang
Province, Maoershan, 8 June 1941, [collector not designated] (USNM); 1 female,
11 June 1941 (USNM); 1 male, 13 June 1941 (USNM); 1 female, 14 June 1941
(USNM); 1 female, 16 June 1941 (USNM); 1 male, Jilin Province, Yablonia Station
[Yabuli], 26 May 1939, [collector not designated] (USNM).

Diagnosis. Tipula (V.) kuwaymai can be recognized by the unarmed gonocoxite
and by the ninth tergite forming a sclerotized, oval saucer which has an elevated edge
anteriorly and is yellow posteriorly with the posterolateral angle blade-shaped. The
wing is distinctly patterned with brown. The female has the cercus brownish yellow
with the tip narrowed and upturned, and the ventral margin has distinct serration. The

Vestiplex crane flies of Korea 33

hypovalva is filamentous. The median incision between hypovalvae is slightly deeper
than posterior margin of eighth sternite.

Male. Body length 17.8-19.7 mm, wing length 19.8—22.9 mm. General body
coloration brownish yellow.

Head. Vertex and occiput gray with brown median line. Rostrum dark brown,
dorsally dusted with gray. Nasus small, almost lacking. Antenna 13-segmented, if bent
backward extending beyond the wing base. Scape and pedicel reddish yellow, flagel-
lum dark brown. Each flagellomere except first one with distinct basal enlargement
and incision. Apical flagellomere small, reduced. Verticils longer than corresponding
segments. Palpus dark brown.

Thorax. Pronotum gray with brown median line. Prescutum and presutural scutum
gray. Median stripes anteriorly gray, posteriorly brown, bordered by darker brown,
fused at base. Lateral stripes blackish gray, bordered by brown. Postsutural scutum
blackish gray; each lobe with light brown spot bordered by brown. Scutellum and
postnotum brownish, dusted with gray, each with brown median line. Pleura brown-
ish, dusted with gray. Wing distinctly patterned with brown. Halter pale with brown
knob. Coxa gray; trochanter yellow; femur brownish yellow with tip dark brown; tibia
and tarsal segments brown. ‘Tarsal claw with tooth.

Abdomen. Brownish yellow. First tergite laterally brown, dorsally yellow, tergites
2—5 yellow with pale and interrupted dorsal stripe. Tergites 6—9 brown, without me-
dian stripe. Lateral stripe distinct. First sternite yellow, sternites 2-6 yellow; remaining
sternites dark brown.

Hypopygium. Dark brown (Fig. 16). Ninth tergite forming concave sclerotized sau-
cer (Fig. 17). Central part of tergal saucer brownish, in the shape of an oval, concave,
transverse plate, anteriorly with elevated edge, laterally and posteriorly with black-
ened, medially interrupted rim. Posterior part of tergal saucer yellow, posterolateral
angle blade-shaped, with blackened ridge connecting with central part of tergal saucer.
Gonocoxite narrow, unarmed (Fig. 18). Outer gonostylus club-shaped (Fig. 19). In-
ner gonostylus in the shape of a curved plate, bidentate at apex, dorsally with acute
tooth, beak claw-shaped (Fig. 20). Gonocoxal fragment with medial sclerites slender,
fused at base; lateral parts apically flattened (Fig. 46). Lateral sclerite relatively small,
nearly V-shaped. Adminiculum semi-open, nearly triangular in lateral view (Fig. 22),
flattened and broadened at base in ventral view (Fig. 23). Semen pump with swollen
central vesicle (Figs 24, 25). Compressor apodeme with broad and round median inci-
sion, forming a 75° angle with posterior immovable apodeme. Posterior immovable
apodeme much longer than compressor apodeme, flattened, with acute tip. Anterior
immovable apodeme narrow, rounded. Intromittent organ tube-shaped, about five
times as long as semen pump, basally and medially brown but passing into brownish
yellow towards apex. Distal part yellow, funnel-shaped (Fig. 26).

Female. Body length 27.4—34.4 mm, wing length 24.5—26.1 mm. Generally similar
to male. Antenna with four basal segments yellow; remaining flagellomeres bicolored.

Female terminalia. Tenth tergite shining-brownish. Cercus brownish yellow, as long
as tenth tergite, with tip narrowed and upturned, ventral margin with distinct serra-

34 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Figures 16-26. Male terminalia of 7! (Vestiplex) kuwayamai \6 hypopygium, lateral view 17 ninth ter-
gite, dorsal view 18 left gonocoxite, lateral view 19 left outer gonostylus 20 left inner gonostylus, lateral
view 21 gonocoxal fragment, dorsal view 22 adminiculum, lateral view 23 adminiculum, ventral view
24 semen pump, dorsal view 25 semen pump and intromittent organ, lateral view 26 distal part of intro-
mittent organ, lateral view. Abbreviations: ig, inner gonostylus; Is, lateral sclerite of gonocoxal fragment;
ms, medial sclerite of gonocoxal fragment; og, outer gonostylus; t9, ninth tergite. Scale bars: 0.7 mm (16);

0.5 mm (17=25); 0.25 mm (26).

Vestiplex crane flies of Korea 35

Figures 27-29. Female terminalia of 7) (Vestiplex) kuwayamai 27 ovipositor, left lateral view 28 eighth
sternite with hypovalvae, ventral view 29 ninth sternite with furca, bursa copulatrix and spermathecae,

dorsal view. Abbreviations: bc, bursa copulatrix; f, furca; s9, ninth sternite; sp, spermatheca. Scale bars:

0.8 mm (27); 0.5 mm (28=29).

tion (Fig. 27). Hypovalva filamentous, distally pale, with short trichia at tip (Fig. 28).
Median incision between hypovalvae slightly deeper than posterior margin of eighth
sternite. Lateral incision scarcely outlined, posterior margin with fine additional projec-

36 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

tion. Lateral angle of eighth sternite rectangular. Ninth sternite posteriorly with split tip
(Fig. 29). Furca in the shape of narrow stripe (Fig. 29). Spermatheca spherical (Fig. 29).

Known distribution. Russia, Japan China (Oosterbroek, 2019) and North and
South Korea. Recorded here for the first time from the Korean Peninsula.

Tipula (Vestiplex) serricauda Alexander, 1914
Figs 30-55, 88

Tipula serricauda Alexander 1914: 237; 1920b: 18.

Tipula asio Alexander 1918: 68 (synonymy after Alexander 1953: 156).

Tipula (Vestiplex) asio: Alexander 1935: 118.

Tipula (Vestiplex) serricauda: Alexander 1935: 118; Oosterbroek and Theowald 1992: 159.

Type material examined. Holotype, female, JAPAN, Tokyo, August 1912 (USNM).

Other examined material (Fig. 88). SOUTH KOREA, 2 males, #14, Oho-ri,
east coast, 10-50 ft, 128°30'E, 38°20'N, 11 June 1954, G.W. Byers (SEM); 1 fe-
male, Geongsanbuk-do, Chilgok-gun, Jicheon-myeon, Mt Hwanghaksan, 4 June
1978, Seon-hui Lee (KU); 1 male, 1 female, Jeollanam-do, Suncheon-si, Songgwang-
myeon, Mt Jogyesan, 23 May 1988, Dokgo (KU); 1 male, Hadong-gun, Okjong-
myeon, Wolhoeng-ri, 24 May 1990, M.J. Gang, IN0000296230 (NIBR); 1 male,
Chungcheongnam-do, Gongju-si, Gyeryong-myeon, 5—7 June 1997, Yeong Lee (KU);
1 male, Minjeong Kim (KU); 1 male, Hadong-gun, Geumseong-myeon, Gadoek-ri,
Hwaryeokbonbu, 19-20 September 1998, J.S. Jeon, IN0000298240 (NIBR); 3 males,
1 female, Goseong-gun, Sangri-myeon, Osan-ri, Mt Odu, 10-11 September 1999,
J.S. Jeon, IN0000298933, IN0000298927, IN0000298197, IN0000297995 (NIBR);
3 females, J.S. Park, IN0000298194, IN00000298192, IN0000298188 (NIBR); 2
males, 1 female, S.Y. Lee, IN0000298932, IN0000298935, IN0000298193 (NIBR);
2 females, G.H.Gang, IN0000298196, IN0000298187(NIBR); 2 females, Hadong-
gun, Geumseong-myeon, Gadoek-ri, Hwaryeokbonbu, 22—23 September 2000, J.S.
Park, NIBR IN0000333964 (NIBR); 1 male, Gwangyang-si, Junggun-dong, Mt Gaya,
Hanseokgwangwangnongwon, 22—23 September 2000, J.H. Son, IN0000333965; 2
females, K.L. Han, IN0000333962, IN0000333956 (NIBR); 1 male, Gwangyang-si,
Junggun-dong, Mt Gaya, Hanseokgwangwangnongwon, 26-27 May 2001, J.S. Park,
IN0000323569 (NIBR); 2 females, Gwangyang-si, Junggun-dong, Mt Gaya, Gun-
jangijae, 31 May—1 June 2003, T.H. An, IN0000298937, IN0000299044 (NIBR);
1 male, Changpyeong-ri, Bongsung-myeon, Bonghwa-gun, Gyeongsangbuk-do,
36°55.12'N, 128°48.65'E, 2014.05.05, H.-W. Byun (NIBR); 1 male, Jeollanam-do,
Gurye-gun, Toji-myeon, Naeseo-ri, Jirisan National Park, Piagol valley, 35°15.95'N,
127°34.85'E, alt. 450 m, 24 August 2014, S. Podenas (NIBR); 1 female, 1 larva, Piagol
valley, 35°16.47'N, 127°33.82'E, alt. 600 m, 1 May 2015, female emerged 5 June 2015,
V. Podeniene (NIBR); 1 male, 2 females, Piagol valley, 35°15.95'N, 127°34.85’E, alt.
450 m, 2 June 2016, S. Podenas (NIBR); 1 male, 6 females, Piagol valley, 35°15.95'N,

Vestiplex crane flies of Korea 37

127°34. 85'E, alt. 450 m, 3 June 2016, S. Podenas (NIBR); 16 males, 4 females, Piagol
valley, 35°15.95'N, 127°34.85'E, alt. 450 m, 3 June 2016, S. Podenas (NIBR); 1 male,
2 females, Piagol valley, 35°16.31'N, 127°33.29'E, alt. 500 m, 3 June 2016, S. Podenas
(NIBR); 2 males, 3 females, Piagol valley, 35°16.4'N, 127°34.15’E, alt. 550 m, 3 June
2016, S. Podenas (NIBR); 8 males (in alcohol), Piagol valley, 35°15.95'N, 127°34.85'E,
alt. 450 m, 24 June 2016 S. Podenas, at light (NIBR); 4 males (in alcohol), 26 June
2016, S. Podenas, at light (NIBR); 3 males (in alcohol), 2 males, 2 females, Jeju-do,
Jeju Island, Seogwipo, Sanghyo-dong, 33°18.51'N, 126°33.57'E, alt. 500 m, 22 May
2017, S. Podenas (NIBR); 2 males, Jeju-do, Jeju Island, Cheju, Jochon-eup, Seonheul-
ri, 33°30.59'N, 126°42.92'E, alt. 1500 m, 23 May 2017, at light, S. Podenas (NIBR);
1 male, Jeju-do, Jeju Island, Jeju-si, Yonggang-dong, 33°25.82'N, 126°35.84'E, alt. 600
m, 24 May 2017, at light, S. Podenas, V. Podeniene (NIBR); 1 male, Gyeonggi-do,
Gunpo-si, Suri-dong, 37°21.03'N, 126°54.93’E, alt. 140 m, 27 May 2017, S. Podenas,
at light, (NIBR); 1 male, 1 female (in alcohol), Jeju-do, Jeju Island, Seogwipo, Saekdal-
dong, 33°21.45'N, 126°27.85'E, alt. 1100 m, 19 June 2019, S. Podenas (NIBR); 1
male, 18 June 2019, H.-Y. Seo (NIBR). Also material listed by Starkevich et al. (2015).

Diagnosis. Tipula (V.) serricauda can be recognized by the gonocoxite being armed
with a black spine and the ninth tergite divided by pale membrane with ventral portion
yellow and bearing a pair of blackened lobes. ‘The body is yellowish, with short antenna
reaching the pronotum if bent backward. Female can be recognized by the short, plate-
shaped hypovalvae.

Male. Body length 12.9-17.3 mm, wing length 15.6-20.2 mm. General body
coloration yellowish.

Head. Yellowish dusted with gray, vertex and occiput yellowish, with dark brown
median line. Rostrum yellowish with conspicuous nasus. Antenna 12-segmented, if
bent backward reaching pronotum. Scape, pedicel, the first and second flagellar seg-
ments yellow; flagellar segments 3-10 darkened at base and yellow apically; remaining
segments dark brown. Each flagellomere, except first one, with small, inconspicuous
enlargement. Apical flagellomere very small, reduced, distinctly shorter than preceding
segment. Verticils longer than corresponding segments. Palpus dark brown.

Thorax. Brownish yellow. Pronotum gray with brown median line. Prescutum and
presutural scutum with four longitudinal, grayish-brown stripes bordered by darker
brown. Intermediate pair brownish, fused anteriorly and posteriorly, separated in the
first third. Lateral stripe grayish. Interspace between median and lateral stripes light
brown. Postsutural scutum yellowish, dusted with light gray; scutal lobe with two yel-
lowish spots. Scutellum yellowish; postnotum yellowish, lightly dusted with gray with
brown median line. Pleura brownish yellow, lightly dusted with gray. Coxa yellowish;
trochanter yellow; femur yellow, distally brown; tibia yellowish brown; tarsal segments
dark brown; claw with tooth. Wing distinctly patterned with brown. Halter yellow,
knob brown with distal part pale yellow.

Abdomen. Brownish yellow, tergites 1 and 2 with brown spots; tergites 3—5 with
dorsal stripe which varies from pale to brown; tergites 6—9 with dorsal stripe distinct,
dark brown. Lateral stripe distinct, dark brown. Posterior and lateral margins of ter-

38 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

gites pale. First sternite yellowish, sternites 2-4 reddish yellow, remaining sternites
passing into brown.

Hypopygium. Brownish (Fig. 30). Ninth tergite divided by pale membrane; dorsal
portion brown medially, provided with long setae; ventral portion yellow with a pair
of blackened, microscopically roughened lobes on either side of midline (Fig. 31). Tip
of lobe obliquely truncated; anal plate shaped as a brown, oblong sclerite. Gonocox-
ite apically produced into a strong, black spine; ventromesal portion a black point
(Fig. 32). Outer gonostylus club-shaped (Fig. 33). Inner gonostylus yellow; beak slen-
der and blackened; lower beak small, nearly triangular (Fig. 34). Dorsal margin with
distinct incision; dorso-lateral crest rounded. Dorsal surface medially with blackened
area. Adminiculum triangular in ventral view; apex acute and split (Figs 30, 35), fused
medially forming a distinct sclerite with raised base. Ventral lobe of appendage of
ninth sternite blackened, roughly rounded, and provided with setae; dorsal lobe re-
duced into small, rounded sclerite (Fig. 35). Semen pump with central vesicle swollen
(Figs 36, 37). Compressor apodeme with median incision forming a 65° angle with
posterior immovable apodeme. Posterior immovable apodeme longer than compressor
apodeme, basally narrow, apically flattened. Anterior immovable rounded. Intromit-
tent organ tube-shaped, about three times as long as semen pump, basally and medially
brownish-yellow, passing into yellow towards acute apex.

Female. Body length 20.2—21.1 mm, wing length 16.4—17.1 mm. Generally simi-
lar to male. Antenna, if bent backward, reaching pronotum; scape, pedicel, and two
basal flagellar segments yellow; remaining flagellomeres vary from yellow to bicolored.
Abdomen trivittate, with broad dorsal stripe.

Female terminalia. Tenth tergite shining brown. Cercus reddish brown, nearly
straight with tip narrowed and upturned; ventral margin with small, distinct serra-
tion; dorsal margin distally also with small serration (Fig. 38). Eighth sternite brown,
apically darker brown (Fig. 39). Hypovalva short, dark brown, in the shape of an
obliquely truncated plate. Lateral angle of eighth sternite obtuse, with small and dis-
tinct incision. Median incision between hypovalvae with serrated edge and provided
with setae. Ninth sternite with lateral sclerites nearly straight, posteriorly with incision;
surface covered by short setae (Fig. 40). Furca long and narrow (Fig. 40). Spermathecae
nearly oval (Fig. 41).

Larva (N = 1). Length 29 mm, width 4 mm. Body light brown (Figs 42-44).

Head capsule. Length 2.3 mm, width 1.2 mm. Head capsule prognathous, hemi-
cephalic, oval, slightly depressed dorsoventrally, and heavily sclerotized (Figs 45, 46).
Internolateralia and externolateralia separated by incisions which reach almost middle
of head capsule. Externolateralia widely separated ventrally (Fig. 46). Premaxillary su-
ture separates side plate from rest of head capsule. Side plate wide, elongated, with two
sensory pits and two long setae anteromedially; a short seta located posteromedially
(Fig. 47). Hypostomium asymmetrical, basally fused with ventral margins of genae and
side plates; bearing eight sharp teeth: four on left side, most prominent tooth in mid-
dle, and three on right side (Fig. 48). Prementum visible from below, bearing five large,
sharp teeth on anterior margin; sides of prementum strongly sclerotized (Fig. 49). La-

Vestiplex crane flies of Korea 39

36 3/

Figures 30-37. Male terminalia of 7’ (Vestiplex) serricauda 30 hypopygium, lateral view 31 ninth ter-
gite, dorsal view 32 left gonocoxite, lateral view 33 left outer gonostylus 34 left inner gonostylus, lateral
view 35 ninth sternite, ventral view (ninth tergite, outer and inner gonostyles removed) 36 semen pump
and intromittent organ, lateral view 37 semen pump, dorsal view. Abbreviations: adm, adminiculum; ap,
anal plate; dl, dorsal lobe of appendage of ninth sternite; dp, dorsal portion of ninth tergite; gcx, gonocox-
ite; ig, inner gonostylus; og, outer gonostylus; t9, ninth tergite; vp, ventral portion of ninth tergite. Scale

bars: 0.7 mm (30); 0.5 mm (31=37).

bial area entirely covered with firm bristles and bearing a pair of cone-shaped palpes.
Prementum dorsally fused with hemispherical and membranous hypopharynx which
is covered with numerous short hairs. Lateral arm of hypopharynx curved and strongly
sclerotized. Frontoclypeus fused with internolateralia. Clypeal part of frontoclypeus
membranous, frontal part sclerotized. One long seta and three sensory pits on anterior

40 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Figures 38-41. Female terminalia of 7’ (Vestiplex) serricauda 38 ovipositor, left lateral view 39 eighth
sternite with hypovalvae, ventral view 40 ninth sternite with furca, dorsal view 41 spermatheca, dorsal
view. Abbreviations: f, furca; s9, ninth sternite. Scale bars: 0.25 mm (38); 0.5 mm (39=41).

part of clypeus; three short setae near inner margin of antenna (Fig. 50). Clypeolabral
suture obscure. Dorsal ecdysial sutures (frontal sutures) present, meeting each other
posteriorly and forming a short median coronal suture. Ecdysial sutures enclose V-
shaped frontoclypeus and extend anteriorly only to base of clypeus. Labrum trapezoidal
and composed of two triangular plates separated by membranous area (Fig. 50). Apical
part of labrum and epipharynx covered with numerous, short hairs. Membranous part
of labrum with a pair of medium-long setae in middle. Labral plates sclerotized only
posteriorly and bearing numerous long firm spines on outer margin. Each plate bears

Vestiplex crane flies of Korea 4]

49

1mm

Figures 42-46. Larva of 7’ (Vestiplex) serricauda 42. general view, dorsal aspect 43 general view, lateral
aspect 44 general view ventral aspect 45 general view of head capsule, dorsal aspect 46 general view of

head capsule, ventral aspect. Scale bars: 5 mm (42-44); 1 mm (45-46).

42 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

52 0.1 mm

Figures 47-53. Larva of 7) (Vestiplex) serricauda 47 side plates 48 hypostoma 49 hypopharynx and
prementum 50 frontoclypeus 51 labrum and antennae 52 right mandible, dorsal view 53 left maxilla,

ventral view. Scale bars: 0.1 mm (47, 49, 51—53); 0.5 mm (48, 50).

Vestiplex crane flies of Korea 43

a long seta, one long and two very short papillae on anterior part, one long seta almost
at middle, and a sensory pit on postero-lateral part (Fig. 51). Antenna elongated, cylin-
drical. It has just one cylindrical segment, which is three times as long as wide at base
(Fig. 51). Apically it has one small cone-shaped and several (exact number difficult to
establish) small, peg-like sensillae; dorsally it has a sensory pit near middle. Mandible
l-segmented and more sclerotized than rest of head capsule, armed with four teeth
(Fig. 52). Apical tooth is the most prominent; first dorsal and ventral teeth smaller
than apical; second dorsal tooth smallest. Prostheca or lacinia mobilis present on dorsal
side of mesal mandibular base; prostheca sclerotized, distinctly widening distally, and
set with numerous hairs. Lateral margin of mandible with two long setae near base; a
sensory pit present at base of dorsal side of mandible. Mandibles operate in horizontal
plane. Conspicuous larval eye spot present below base of mandible. Maxilla consists
of cardo and outer and inner lobes. Cardo wedge-shaped, bearing two long setae near
distal end and a long seta near its base (Fig. 53). Outer lobe (stipes) sclerotized, ex-
cept apex, which bears prominent, cylindrical palpus with several sensory structures.
Short sensory structure and several long, sclerotized spines (exact number difficult to
establish) on inner margin of stipes. Inner lobe (galea fused with lacinia) ventrally
bears elongated, narrow sclerite extending around inner margin onto its dorsal surface;
diamond-shaped sclerite present dorsally at base (Fig. 53); apical part with numerous
short hairs, three long setae, and prominent sensory structure. Lacinia armed with sev-
eral stout bristles and bears a ridge with sclerotized spines on outer margin.

Thorax. All thoracic segments wider than long. Anterior part of second and third
segments covered with much denser pubescence than posterior (Figs 42-44).

Abdomen. First abdominal segment almost twice as long as wide. Abdominal seg-
ments II—VII almost as long as wide. All abdominal segments except last one covered
by short microscopic hairs arranged into transverse rows, which are interrupted by pu-
bescence on ventral and dorsal sides. Most macrosetae dark brown except L2 and L3.
Dorsal setae D2 and D3 longest; seta D1 only slightly shorter than D2 and D3; setae
D4—D6 short and appressed, more than three times shorter than D2 and D3. Setae D2
+ D3 and D5 + D6 close to each other and separate from others. Lateral setae L2 and
L3 very short and pale; L1 and L4 long and almost equal in length; L2 dorsolateral to
L1. Setae L1, L4 more than four times as long as L2 and L3. V2 almost equal to V3
and both the longest of ventral setae. Setae V4 and V5 slightly shorter than V2 and V3.
Seta V1 very short, more than five times shorter than V2 and V3.

Anal division. Dorsal and lateral lobes of spiracular disc subconical, lateral lobes in
dorsolateral position (closer to dorsal than to ventral lobes). Dorsal and lateral lobes
similar in length, twice as long as wide at base (Fig. 54). Ventral lobe very small, trian-
gular, its length almost equal to width at base. Ventral lobe almost five times as short
as dorsal or lateral lobe. Dorsal lobe completely sclerotized, with apex extended into
acute, anteriorly directed point (Fig. 55). Lateral lobe with long, narrow, curved, dark
sclerite that starts near base of spiracle and extends to mid-length of lobe; a long seta
present in middle of outer edge of lobe. Ventral lobe with three small, dark spots at
base; outermost spot most prominent. Distal half of lobe sclerotized, with long apical

44 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

ah its ae" age

bey...

1mm

Figures 54-55. Larva of 7! (Vestiplex) serricauda 54 spiracular field 55 anal division, lateral view. Scale

bar: 1 mm.

seta. Spiracle subcircular, inner circle black, outer ring brown; distance between spira-
cles almost twice diameter of a spiracle. Remaining area around spiracles white and
glabrous. Four white, fleshy anal papillae arranged into anterior and posterior pairs.
Anterior papilla broadly rounded; posterior papilla more elongated (Fig. 54). A brown
band separates anus and anal papillae; this band connected to marginal band.

Habitat. Larvae were found under leaf litter and woody debris accumulated on
boulders. Two identical last instar larvae were collected and one of them was kept for
rearing. A female emerged after 36 days and identified as 7’ (V) serricauda.

Known distribution. China, Japan, and South Korea (Starkevich et al. 2015;
Oosterbroek 2019).

Remarks. A single male belonging to 7’ (V.) serricauda was erroneously identified
and published as 7’ (Mediotipula) anatoliensis Theowald, 1978 by Baek and Bae (2016)
based on Korean material. The hypopygium, with broken apical part of gonocoxite
(Baek and Bae 2016: fig. 2E), is identical to that of 77 (V) serricauda and can be easily
recognized by the shape of inner gonostylus.

New data received from the larvae of 7. (V.) serricauda once again confirm that the
most important synapomorphy in the subgenus T° (Vestiplex) is a brown band sepa-
rating the anus from the anal papillae, which is a unique character for this subgenus.
According to the type of sclerotization of the spiracular field, larvae of the subgenus
LT. (Vestiplex) (based on T. (V.) nubeculosa Meigen, 1804, T. (V.) ortorum Linnaeus,

Vestiplex crane flies of Korea 45

1758, T. (V.) montana Curtis, 1834, T. (V.) excisa Schummel, 1833, T. (V.) scripta
Meigen, 1830, 7: (V.) platymera Walker, 1856, T. (V.) arctica Curtis, 1835, T. (V.)
semivittata Savchenko, 1960, and 7. (V.) serricauda) can be divided into four groups.
The first group includes larvae of such species as T. (V.) scripta, T. (V.) platymera, and
T. (V.) semivittata. They possess sclerotized dorsal lobes with sclerotization encom-
passing the apices of the lobes and forming an acute, slightly anteriorly directed point.
In addition, each lateral lobe has a thin, straight, and more or less vertical sclerite. The
ventral lobes of this first group have sclerotized apices with several short setae and a
narrow sclerotized band (with three dark spots inside) extending across the base. The
second group includes 7. (V.) arctica, T. (V.) nubeculosa, and T. (V.) hortorum. This
group possesses sclerotized dorsal lobes but the sclerotization does not reach the apex
of the lobe. Lateral and ventral lobes are very similar to that of the first group. The
third group includes T: (V.) excisa and T. (V.) montana. This group of species possesses
sclerotized dorsal lobes, with the sclerotization reaching the apex, but never forming
anteriorly directed points. The lateral and ventral lobes are very similar to that of the
first and second groups. The fourth group consists of only 7. (V.) serricauda. Each
dorsal lobe of this species is sclerotized both anteriorly and posteriorly, with each
apex forming an acute, strongly anteriorly directed point. Each lateral lobe has a long,
narrow, curved, dark sclerite, extending from near the base of a spiracle to the mid-
length of each lobe. Each ventral lobe has a sclerotized distal part with a long apical
seta, the base of each lobe with three small dark spots, with the narrow band missing.
Sclerotization of the spiracular field of the fourth group most resembles that of larvae
of T. (Triplicitipula) but not as seen in other groups of the subgenus 7’ (Véestiplex).
The macrosetal arrangement on the dorsum and venter of the abdomen appears to
be consistent in the subgenus, but the arrangement of the abdominal lateral setae is
species-specific. Head capsules have never been comparatively studied in detail for
the genus 7ipula or the subgenus 7. (Vestiplex); thus, comparison has been impossible
among species and subgenera.

Tipula (Vestiplex) tchukchi Alexander, 1934
Figs 56-66, 89

Tipula (Vestiplex) tchukchi Alexander 1934: 408.

Tipula (Vestiplex) tchukchi obtusidens Savchenko 1964: 205 (synonymy after Starkevich
and Paramonov 2016).

Tipula (Vestiplex) bo Mannheims 1967: 148 (synonymy after Mannheims and
Savchenko 1973).

Type material examined. Holotype, male, RUSSIA, Chukchi Peninsula, Chukotka
Autonomous Okrug, Markovo township near Anadyr town, 6 July 1896, Gondatti
(ZIN); paratype, female, topotypic (ZIN); paratype, male, Kamchatka Kray, mouth of
Kichiga River, 27 June 1910, Skorikov (ZIN).

46 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Other examined material (Fig. 89). NORTH KOREA, 1 male, Seren Mts, alt.
3500 ft, 25 June 25 1938, Yankovsky (USNM); 1 female, MONGOLIA, Tov Aimag,
Erdene Soum, Gorkhi Terelj National Park, unnamed tributary of Tuul River on its
west side, 1.6 km upstream from Daichin crossing, 48.21780°N, 107.90392°E, alt.
1600 m, 9 July 2003, SRP#03070902, coll. O. Yadamsuren (ANSP).

Diagnosis. 7ipula (V.) thukchi can be recognized by the unarmed gonocoxite and
the ninth tergite forming a concave, roughly rectangular, sclerotized saucer. The body
coloration is blackish yellow, and the wing pattern is indistinct. The female has the cer-
cus with an apical incision and outer margin rough and distinctly serrated. The eight
sternite has a distinct lateral incision, and the hypovalvae are filamentous.

Male. Body length 16.8 mm, wing length 17.1 mm. General body coloration
blackish yellow.

Head. Gray, vertex and occiput gray with brown median line. Rostrum brown,
dorsally dusted with gray. Nasus short. Antenna 13-segmented, if bent backward ex-
tending beyond the wing base. Scape and pedicel yellowish; first flagellar segment
brownish; subsequent flagellar segments dark brown. Each flagellomere except first one
with basal enlargement and moderately incised. Apical flagellomere small, reduced.
Verticils shorter than corresponding segments. Palpus dark brown.

Thorax. Brown, dusted with grey. Pronotum blackish, gray dusted, with brown
median line. Prescutum and presutural scutum brown, grey pruinose with four longi-
tudinal stripes bordered by brown. Intermediate pair fused into brown median line.
Interspace between median and lateral stripes light gray. Postsutural scutum blackish,
gray pruinose with median line. Scutal lobe with two spots bordered by brown. Scutel-
lum brown, postnotum brown, dusted with gray-brown; both sclerites with darker
median line. Pleura brown, dusted with gray. Coxa brown, grey pruinose. Trochanter,
femur, and tibia yellowish. Tarsal segments brown. Distal part of femur and tibia dark
brown. Tarsal claws toothed. Wing pattern indistinct, only weak darkening along vein
CuA. Halter yellowish, with brown knob.

Abdomen. Yellow. Abdominal segments 1-4 yellow, subsequent segments passing
into dark brown. Tergites with lateral margins narrowly pale; dorsal stripe broad; lat-
eral stripe pale.

Hypopygium. Brownish black. Ninth tergite forming a large, concave, roughly
rectangular sclerotized saucer. Main body of tergal saucer brown and rim blackened
(Fig. 56). Posterior margin of tergal saucer toothed with small denticles, broadly emar-
ginated, with deep median U-shaped notch. Lateral angles of tergal saucer obtuse,
broadly truncated. Anterior and lateral portions of tergal saucer raised into sclerotized
border; border laterally produced into obtuse point directed caudad and situated un-
der lateral angle of tergal saucer so that ninth tergite with two teeth in lateral view.
Gonocoxite unarmed (Fig. 57). Outer gonostylus flattened, slightly curved, with apex
rounded (Fig. 58). Inner gonostylus in the shape of a curved plate, terminating dor-
sally with obtuse tooth; beak claw-shaped (Fig. 59). Gonocoxal fragment large, with
lateral and medial sclerites well developed (Fig. 60). Medial sclerites fused anteriorly

Vestiplex crane flies of Korea 47

63

Figures 56-64. Male terminalia of T (Vestiplex) tchukchi 56 ninth tergite, dorsal view 57 left gonocoxite,
lateral view 58 left outer gonostylus 59 left inner gonostylus, lateral view 60 gonocoxal fragment, dorsal
view 61 adminiculum, lateral view 62 semen pump, dorsal view 63 semen pump and intromittent organ,
lateral view 64 distal part of intromittent organ, lateral view. Abbreviations: Is, lateral sclerite of gonocoxal

fragment; ms, medial sclerite of gonocoxal fragment. Scale bars: 0.5 mm (56-63); 0.25 mm (64).

48 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Figures 65-66. Female terminalia of 7’ (Vestiplex) tchukchi (Mongolia). 65 tenth tergite with cercus, left

lateral view 66 eighth sternite with hypovalvae, ventral view. Scale bar: 0.5 mm.

into long, narrow apodeme; posteriorly with rounded apodeme; lateral parts broad-
ened and arched. Lateral sclerite large and bilobed, expanded at base. Adminiculum
canoe-shaped (Fig. 61). Semen pump with central vesicle relatively small and flattened
(Figs 62, 63). Compressor apodeme with broad median incision, forming a 50° angle
with posterior immovable apodeme. Posterior immovable apodeme approximately as
long as compressor apodeme. Anterior immovable apodeme rounded. Intromittent or-
gan tube-shaped, about four times as long as semen pump, basally and medially brown,
passing into yellow towards apex. Distal part ventrally truncated, shovel-shaped, with
rough edge and two stripped fragments (Fig. 65).

Female. Female not known from Korean Peninsula, but can be recognized by cer-
cus having apical incision and rough and distinctly serrated outer margin (Fig. 65).
Hypovalva filamentous, flattened, broadened at base, distally pale, with short trichia
at tip (Fig. 66). Median incision between hypovalvae deeper than posterior margin of
eighth sternite; lateral incisions distinct.

Known distribution. Finland, Sweden, Russia, and Mongolia (Oosterbroek
2019). Recorded here for the first time from the Korean Peninsula.

Tipula (Vestiplex) verecunda Alexander, 1924
Figs 67-85, 90

Tipula verecunda Alexander 1924: 606.
Tipula (Vestiplex) verecunda: Alexander 1935: 118; Savchenko 1964: 152; Oosterbroek
and Theowald 1992: 160.

Type material examined. Holotype, male, RUSSIA, [Sakhalin Island] Toyohara
[Yuzhno-Sakhalinsk], July16, 1922, Esaki (USNM); paratypes, 3 males, 1 female,

Vestiplex crane flies of Korea 49

13-14 July 1922, Esaki, topotypic (USNM); paratype, female, Shimizu [Southern
Sakhalin], 27 July 1922, Esaki (USNM); paratype, female, Odasam [Southern Sakha-
lin], 31 July 1922, Esaki (USNM).

Other examined material (Fig. 90). SOUTH KOREA, 5 males, 3 females, Jeju-do,
Jeju Island, Seogwipo, Saekdal-dong, 33°21.45'N, 126°27.85'E, alt. 1100 m, 18 June
2019, H.-Y. Seo (NIBR); 6 males, Seogwipo, Saekdal-dong, 35°21.62'N, 126°27.76'E,
alt. 1100 m, 18 June 2019, S. Podenas (NIBR). CHINA, 1 male, Shaanxi, Qinling
Mts, Hauzherza vill., 33°52.42'N, 107°48.77'E, alt. 1600 m, 2—3 June 2009, leg. Sal-
daitis & Floriani (NRC).

Diagnosis. Tipula (V.) verecunda can be easily recognized by the eighth sternite
laterally having long setae and the ninth sternite being ventrally produced into a small
tubercle. The ninth tergite has a U-shaped notch posteriorly, and anterior to the notch,
the tergite is divided by a pale membrane. ‘The ventral portion of the ninth tergite is
yellow, with a pair of blackened narrow plates, and the gonocoxite is unarmed. The
thorax is grey, with four darker grey stripes narrowly bordered by brown, and the wing
is distinctly patterned with dark brown. The female can be recognized by the cercus
having a smooth margin and a long, blade-shaped hypovalvae.

Male. Body length 17.4—21.6 mm, wing length 20.0-24.5 mm. General body
coloration brownish yellow.

Head. Vertex and occiput ochraceous yellow, with dark median line. Rostrum dark
brown, dorsally narrowly ochraceous yellow. Nasus distinct. Antenna 13-segmented, if
bent backward extending beyond the wing base. Scape and pedicel yellow; flagellum
brownish black. Each flagellomere, except for first one, with weak basal enlargement.
Apical flagellomere small, reduced. Verticils longer than corresponding segments. Palpus
dark brown.

Thorax. Pronotum ochraceous yellow, with dark median spot. Prescutum and
presutural scutum grey, with four darker grey stripes, narrowly bordered by brown. In-
termediate pair fused anteriorly. Interspace ochraceous light yellow. Postsutural scutal
lobe with two ochraceous brownish yellow spots bordered by brown. Scutellum brown,
with broad, dark brown median line. Postnotum brownish, with pale median line, and
dusted with gray. Pleura yellowish. Wing distinctly patterned with dark brown. Halter
pale, with brownish-black knob. Coxa yellowish; trochanter yellow; femur basally yel-
low, passing into brown towards dark brown tip. Tibia and tarsal segments brownish
black. Tarsal claw with tooth.

Abdomen. Abdominal segments 1—4 yellow; segment 5 blackish, yellowish laterally;
remaining segments black. Dorsal stripe on first tergite broad, on tergites 2-4 pale, on
fifth tergite black. Lateral abdominal stripe distinct.

Hypopygium. Black (Fig. 67). Ninth tergite posteriorly with deep U-shaped
notch, anteriorly to notch divided by pale membrane, dorsal portion dark brown,
posterior margin provided with setae, additional short projection on either side of
midline (Figs 68, 69). Ventral portion yellow with a pair of blackened and micro-
scopically serrated narrow plates; tip of plate pointed outward. Anal plate shaped as
a small brown sclerite (Fig. 68). Gonocoxite unarmed, irregular in outline (Fig. 71).

50 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Figures 67-79. Male terminalia of T’ (Vestiplex) verecunda 67 hypopygium, lateral view 68 ninth ter-
gite, dorsal view 69 ninth tergite, caudal view 70 eighth sternite, lateral view 71 left gonocoxite, lateral
view 72 left outer gonostylus 73 left inner gonostylus, lateral view 74 ninth sternite, ventral view (ninth
tergite, outer and inner gonostyles removed) 75 right dorsal lobe of appendage of ninth sternite 76 right
ventral lobe of appendage of ninth sternite 77 adminiculum, lateral view 78 semen pump, dorsal view
79 semen pump and intromittent organ, lateral view. Abbreviations: adm, adminiculum; ap, anal plate;
dl, dorsal lobe of appendage of ninth sternite; dp, dorsal portion of ninth tergite; gcx, gonocoxite; gcx,
gonocoxite; ig, inner gonostylus; og, outer gonostylus; s8, eighth sternite; vl, ventral lobe of appendage of
ninth sternite; vp, ventral portion of ninth tergite; vt, ventral tubercle of ninth sternite. Scale bars: 0.8 mm

(67); 0.5 mm (68-79).

Outer gonostylus nearly oval, with basal part narrowed (Fig. 72). Inner gonostylus in
the shape of rounded sclerite, terminating into a short upper beak with a small lower
beak; beaks separated by round incision (Fig. 73). Dorsal crest with yellow setae and
short, black spines grouped on dorsal surface, edge basally bent outward. Adminicu-
lum nearly parallel-sided in ventral view, fused medially forming a distinct sclerite

Vestiplex crane flies of Korea 51

Figures 80-85. Female terminalia of 7 (Vestiplex) verecunda 80 ovipositor, left lateral view 81 eighth
sternite with hypovalvae, ventral view 82 distal part of eighth sternite with hypovalvae, ventral view
83 ninth sternite, dorsal view 84 furca, dorsal view 85 spermatheca, dorsal view. Scale bars: 0.7 mm

(80, 82); 0.5 mm (81, 83-85).

(Fig. 74). Basal part of adminiculum broadened and raised; apex funnel-shaped, with
preapical incision in ventro-lateral view (Fig 77). Ninth sternite ventrally produced
into small tubercle (Figs 67, 74). Appendage of ninth sternite with ventral lobe ob-
long, tip narrowed, surface with setae (Figs 74, 76). Dorsal lobe in the shape of flat-
tened, curved plate; tip on inner surface provided with setae (Figs 74, 75). Eighth

52 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Figures 86-90. Collecting sites of 7’ (Vestiplex) in Korean Peninsula 86 T (V) coquillettiana 87 T. (V.)
kuwayamai 88 T. (V.) serricauda 89 T: (V.) tchukchi 90 T: (V.) verecunda.

sternite posteriorly with pale median area; laterally provided with long setae reaching
1.1 mm long. Semen pump with central vesicle swollen (Figs 78, 79). Compressor
apodeme flattened forming a 50° angle with posterior immovable apodeme. Posterior
immovable apodeme shorter than compressor apodeme, narrowed. Anterior immov-
able apodeme narrow. Intromittent organ tube-shaped, about three times as long

Vestiplex crane flies of Korea 53

as semen pump, basally and medially brownish yellow, passing into yellow towards
acute apex.

Female. Body length 27.3—28.7 mm, wing length 23.0—24.5 mm. Generally simi-
lar to male. Thorax brown. Abdomen with distinct median stripe. Tergites and sternites
with lateral margin pale.

Female terminalia. Tenth tergite basally shining brown but on other two-thirds
shining black. Cercus reddish brown, nearly straight, with tip narrowed; ventral margin
smooth (Fig. 80). Eighth sternite with hypovalva long, blade-shaped, basally with setae
(Figs 80-82). Lateral angle of eighth sternite flattened and slightly extended. Ninth
sternite with posterior half funnel-shaped, rounded apically (Fig. 83). Furca in the
shape of a pale stripe narrowed anteriorly (Fig. 84). Spermatheca spherical (Fig. 85).

Known distribution. Russia, Japan, and China (Oosterbroek 2019). Recorded
here for the first time from South Korea.

Acknowledgements

We give our warmest thanks to all our Korean friends and colleagues who helped us
during our visits to South Korea. We are very grateful for professor Y.J. Bae for the
specimens from the Korea University, Seoul, Republic of Korea; J.C. Thomas (SEM) for
the help with Korean specimens from the University of Kansas, USA.; Dr E Shockley
and Dr T. Dikow (USNM), Dr J.K. Gelhaus (ANSP) for use of the USNM specimens;
Dr N. Paramonov (ZIN), St. Petersburg, Russia; and A. Saldaitis (NRC), Vilnius,
Lithuania. We thank J. Kim (KU) for the help with Korean labels.

We are grateful to Dr F. Brodo (Canadian National Collection of Insects, Arach-
nids and Nematodes, Ottawa, Canada) and Dr J. Gelhaus (ANSP) for revising and
improving the manuscript.

This work was supported by a grant from the National Institute of Biological Re-
sources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of
Korea (NIBR201902108).

References

Alexander CP (1914) Report on a collection of Japanese crane-flies (Tipulidae, Diptera) [concl.].
Canadian Entomologist 46: 236-242. https://doi.org/10.4039/Ent46236-7

Alexander CP (1918) New species of tipuline crane-flies from eastern Asia (Tipulidae, Diptera).
Journal of the New York Entomological Society 26: 66-75.

Alexander CP (1920a) The crane-flies of New York. Part II. Biology and phylogeny. Mem-
oirs, Cornell University Agricultural Experiment Station 38: 691-1133. https://doi.
org/10.5962/bhI.title.33641

Alexander CP (1920b) New or little-known crane-flies from Japan (Tipulidae, Diptera). Trans-
actions of the American Entomological Society 46: 1-26.

54 Pavel Starkevich et al. / ZooKeys 1061: 23-55 (2021)

Alexander CP (1921) Undescribed species of Japanese crane-flies (Tipulidae, Diptera). Part II.
Annals of the Entomological Society of America 14: 111-134. https://doi.org/10.1093/
aesa/14.2.111

Alexander CP (1924) New or little-known crane flies from northern Japan (Tipulidae, Dip-
tera). Philippine Journal of Science 24: 531-611.

Alexander CP (1925) New or little-known crane-flies. Part I. Encyclopedie Entomologique, (B
II), Diptera 2: 87-93.

Alexander CP (1934) New or little-known Tipulidae from eastern Asia (Diptera). XVII. Philippine
Journal of Science 52: 395-442.

Alexander CP (1935) New or little-known Tipulidae from eastern Asia (Diptera). XXV. Philippine
Journal of Science 57: 81-148.

Alexander CP (1953) Records and descriptions of Japanese Tipulidae (Diptera). Part II. The
crane-flies of Shikoku. II. Philippine Journal of Science 82: 141-179.

Alexander CP (1965) New subgenera and species of crane-flies from California (Diptera:
Tipulidae). Pacific Insects 7: 333-386.

Bezzi M (1924) Una nuova Tipula delle Alpi con ali ridotte anche nel maschio (Dipt.). Annali
del Museo Civico di Storia naturale di Genova 51: 228-233.

Baek HM, Bae YJ (2016) New records of the genus 7ipu/a (Diptera: Tipulidae: Tipulinae) in
Korea. Entomological Research Bulletin 32: 161-164.

Brodo F (2017) Taxonomic review of Angarotipula Savchenko, 1961 (Diptera: Tipulidae) in
North America. Canadian Entomologist 150: 12—34. https://doi.org/10.4039/tce.2017.43

Chiswell JR (1956) A taxonomic account of the last instar larvae of some British Tipulinae
(Diptera: Tipulidae). Transactions of the Royal Entomological Society of London 108:
409-484. https://doi.org/10.1111/j.1365-2311.1956.tb01273.x

Cumming JM, Wood DM (2017) Adult morphology and terminology. In: Kirk-Spriggs AH,
Sinclair BJ (Eds) Manual of Afrotropical Diptera, Volume 1: Introductory Chapters and
Keys to Diptera Families. Suricata 4. South African National Biodiversity Institute, Preto-
ria, 89-133.

Dobrotworsky NV (1968) The Tipulidae (Diptera) of Australia. I. A review of the genera of the
subfamily Tipulinae. Australian Journal of Zoology 16: 459-494. https://doi.org/10.1071/
ZO9680459

Edwards FW (1931) Some suggestions on the classification of the genus Tipula (Dip-
tera, Tipulidae). Annals and Magazine of Natural History 8(10): 73-82. https://doi.
org/10.1080/00222933108673359

Gelhaus JK (1986) Larvae of the crane fly genus Tipula in North America (Diptera: Tipulidae).
The University of Kansas Science Bulletin 53: 121-182.

Gelhaus JK (2005) Systematics and biogeography of the desert crane fly subgenus Tipula (Ere-
motipula) Alexander (Diptera: Tipulidae). Memoirs of the American Entomological Society
46: 1-235.

Lantsov VI (2003) Biology, ecology and preimaginal stages of the crane fly Tipula semivittata
semivittata (Diptera, Tipulidae). Entomological Review 83: 906-913.

Mannheims B (1953) 15. Tipulidae. In: Lindner E (Ed.) Die Fliegen der palaearktischen Region,
3(5)1, Lief 173: 113-136.

Vestiplex crane flies of Korea »»

Mannheims B (1963) 15. Tipulidae. In: Lindner E (Ed.) Die Fliegen der palaearktischen Re-
gion, 3(5)1, Lief 238: 137-176.

Mannheims B (1967) Tipula (Vestiplex) bo sp. n. und andere Tipuliden aus Fennoskandien
(Diptera). Notulae Entomologicae 47: 147-156.

Mannheims B, Savchenko EN (1973) Ergebnisse der zoologischen Forschungen von Dr. Z.
Kaszab in der Mongolei, 303. Tipulidae (Diptera), II. Folia Entomologica Hungarica,
(N.S.), Supplement 26: 157-186.

Neugart C, Schneeberg K, Beutel RG (2009) The morphology of the larval head of Tipuli-
dae (Diptera, Insecta), the dipteran groundplan and evolutionary trends. Zoologischer
Anzeiger 248: 213-235. https://doi.org/10.1016/j.jcz.2009.10.001

Neumann H (1958) Der Bau und die Funktion der mannlichen Genitalapparate von Trichocera
annulata Meig. und Tipula paludosa Meig. (Dipt. Nematocera). Deutsche Entomologische
Zeitschrift (N.F.) 5: 235-298.

Oosterbroek P (2019) Catalogue of the Craneflies of the World (CCW), 12 Nov. 2019. http://
nlbif-eti.uva.nl/ccw/index.php [Accessed on: 29 Nov. 2019]

Oosterbroek P, Theowald B. (1992) Family Tipulidae. In: Sods A, Papp L (Eds) Catalogue of
Palaearctic Diptera 1. Hungarian Natural History Museum, Budapest, 56-178.

Podeniene V (2003) Morphology and ecology of the last instar larvae of the crane flies (Diptera,
Tipulomorpha) of Lithuania. Doctoral dissertation, Vilnius University, 1-295.

Rogers JS (1942) The crane flies (Tipulidae) of the George Reserve, Michigan. Miscellaneous
Publications, Museum of Zoology, University of Michigan 53: 1-128.

Savchenko EN (1960) A contribution to the taxonomy of crane-flies (Diptera, Tipulidae) of the
subgenus Vestiplex Bezzi of the genus Tipula L. Horae Societatis Entomologicae Rossicae
Unionis Soveticae 47: 143-216. [In Russian]

Savchenko EN (1964) Crane-flies (Diptera, Tipulidae), Subfam. Tipulinae, Genus Tipula L., 2.
Fauna USSR, N.S. 89, Nasekomye Dvukrylye [Insecta Diptera] 2(4): 1-503. [In Russian]

Starkevich P, Paramonov NM (2016) New records of Tipula (Vestiplex) crane flies from the
Palaearctic [Diptera: Tipulidae] with a new synonym. Journal of the Kansas Entomological
Society 89: 80-84. https://doi.org/10.2317/151024.1

Starkevich P, Podenas S, Young CW (2015) New distribution records for crane flies of Tipula
(Vestiplex) Bezzi (Diptera, Tipulidae). Journal of the Kansas Entomological Society 88:
121-123. https://doi.org/10.2317/JKES1303.31.1

Teale SA, Gelhaus JK (1984) Natural history and synonymy of Tipula (Vestiplex) platymera
Walker (Diptera: Tipulidae) with description of the larvae and pupae. Journal of the Kansas
Entomological Society 57: 423-429.

Theowald B (1967) Familie Tipulidae (Diptera, Nematocera). Larven und Puppen. Bestim-

mungsbucher zur Bodenfauna Europas 7: 1-100.