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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by B. Sinclair: 8 Feb. 2022; published: 11 Mar. 2022 131
Zootaxa 5115 (1): 131–145
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5115.1.9
http://zoobank.org/urn:lsid:zoobank.org:pub:5FF52BDB-1928-4C4A-ACA2-47B5B869A611
Tipula (Vestiplex) levisoni sp. nov., a new cranefly (Diptera, Tipulidae)
from the Western Himalayas
PAVEL STARKEVICH1,4, SIGITAS PODĖNAS1,2 & DUNCAN SIVELL3
1Nature Research Centre, Akademijos str. 2, LT-08412 Vilnius, Lithuania
2Life Sciences Centre of Vilnius University, Sauletekio str. 7, LT-10257 Vilnius, Lithuania
�
sigitas.podenas@gamtc.lt; https://orcid.org/0000-0002-2597-566X
3Natural History Museum, Cromwell Road, London, SW7 5BD, United Kingdom
�
d.sivell@nhm.ac.uk
4Corresponding author:
�
pavel.starkevic@gmail.com; https://orcid.org/0000-0002-0746-0398
Abstract
A new cranefly, Tipula (Vestiplex) levisoni sp. nov., is described and illustrated based on males and females collected
in India and Nepal. The illustrations of male and female genitalia for the most closely related regional species, T. (V.)
tanycera Alexander, 1961 and T. (V.) mitchelli Edwards, 1927 are also provided. The female of T. (V.) mitchelli is described
for the first time.
Key words: Kashmir, hypopygium, ovipositor, taxonomy, nematoceran, Tipuloidea
Introduction
The world fauna of the subgenus Tipula (Vestiplex) Bezzi, 1924 currently contains 177 described species distributed
throughout the Nearctic, Palaearctic and Oriental regions (Oosterbroek 2021). The majority of the species are asso-
ciated with mountain ranges (Pyrenees, Alps, Caucasus and Himalayas) where adults are found at altitudes reaching
4300 m in the Himalayas (Savchenko 1960, 1964). The highest species diversity is observed in the East Palaearctic
(81 species) and Oriental (80 species) regions with four species known in the Western Himalayas (Oosterbroek
2021).
The subgenus can be recognized by females possessing powerfully constructed and heavily sclerotised cerci
with the outer margin usually serrated. The male genitalia typically have the epandrium distally forming a sclero-
tised plate or divided mid-dorsally (Alexander 1935, 1965). The subgeneric characters are also summarized in Men
et al. (2021) and Starkevich et al. (2020).
The aim of this study is to describe and illustrate the new species, provide illustrations of local, closely related
species and construct an identification key for all five regional species of T. (Vestiplex).
Material and methods
The new species was detected while working with the T. (Vestiplex) cranefly collection at the Natural History Mu-
seum, London (BMNH). A total of two males and two females were found in a separate unit tray labeled as “Leuco-
procta group (Sav.) sp.nov.” with additional labels mounted with corresponding specimens with the following data
“Tipula (Vestiplex) sp. nov. det. R.I. Vane-Wright 1962”. Abbreviations for institutional collections used herein:
BMNH = Natural History Museum, London, United Kingdom; NRC = Nature Research Centre, Vilnius, Lithuania;
SEM = Snow Entomological Museum, University of Kansas, Lawrence, KS, USA; USNM = Smithsonian, National
Museum of Natural History, Washington, DC, USA; ZIN = Zoological Museum of the Zoological Institute of the
Russian Academy of Sciences, St. Petersburg, Russia; another abbreviation: PS = slide Pavel Starkevich.
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132 · Zootaxa 5115 (1) © 2022 Magnolia Press
The pinned specimens were studied with a Nikon SMZ800 stereomicroscope. Photographs were taken with an
INFINITY–1 camera mounted on a Nikon Eclipse 200 stereomicroscope and a Canon EOS 80D camera mounted on
an Olympus SZX10 dissecting microscope. Dry specimens were photographed with a Canon EOS 80D at the Natu-
ral History Museum, London and the Nature Research Centre, Vilnius. Digital photos were processed and layers
were stacked using the program HeliconFocus (http://www.heliconsoft.com/heliconsoft-products/helicon-focus/).
Genitalia were studied after boiling them in 10% NaOH for 5–10 minutes, neutralized by acetic acid and then
preserving them in microvials filled with glycerol. An identification key is provided for T. (Vestiplex) species known
from the Western Himalayas including Kashmir and the northernmost geographical region of the Indian subconti-
nent (includes Indian, Pakistani and Chinese administered territories).
Descriptive terminology follows that of Cumming & Wood (2017). The term gonocoxal fragment (= sclerites
sp1 and sp2 (Neumann 1958); = genital bridge (Dobrotworsky 1968)) for the inner structure covered by the epan-
drium is adopted from Brodo (2018).
The following specimens were studied for comparative purposes and to generate the identificiation key to spe-
cies:
Tipula (Vestiplex) avicularia Edwards, 1928: HOLOTYPE ♂, labelled: CHINA, Tibet, Yatung, 10000 ft. [3048
m], 16.iv.1924, Maj. R. W. G. Hingston; Everest Exp. Brit.Mus. 1924-386; Tipula avicularia F. W. Edwards det.
1928; Holotype Tipula avicularia Edwards, det. J. E. Chainey, 1995; BMNH(E)#246051 (BMNH). PARATYPES:
Tibet, Yatung, 4500 ft. [1372 m], A. E. Hobson [wing and genitalia on slide mounted] (1 ♂, USNM); INDIA, Sik-
kim: Lachen R., 8000 ft. [2438 m], 25.iv.1924, Maj. R.W.G. Hingston; Paratype Tipula avicularia Edwards, det. J.
E. Chainey, 1995; Everest Exp. Brit.Mus. 1924-386; BMNH(E)#246052 (1 ♂, BMNH); INDIA, Kashmir, Kaj-Nap
Range, 9000 ft. [2743 m], 3.vi.1934, V. Hutchinson [wing and genitalia on slide mounted] (1 ♂, USNM).
Tipula (Vestiplex) opilionimorpha aligera Savchenko, 1956: LECTOTYPE ♂, labelled: INDIA, Kashmir, La-
dakh, near Pass Kounsi-La, upper basin of Indus River, 19 VII 1910, Trubeckoj (ZIN). PARALECTOTYPES:
same data as lectotype (1 ♂, ZIN); near Pass Kounsi-La, settlement Chipuscot, 25–30.VII.1910, Trubeckoj (1 ♂,
ZIN); near Pass Kounsi-La, Sumda, alt. 11000 ft. [3353 m], 20.VII.1910, Trubeckoj (1 ♀, ZIN).
List of Tipula (Vestiplex) species from the Western Himalayas
Tipula (Vestiplex) avicularia Edwards, 1928
Tipula (Vestiplex) levisoni sp. nov.
Tipula (Vestiplex) mitchelli Edwards, 1927
Tipula (Vestiplex) opilionimorpha aligera Savchenko, 1956
Tipula (Vestiplex) tanycera Alexander, 1961
Key to Tipula (Vestiplex) species from the Western Himalayas
Males
1 Femora with yellow preapical ring. Abdominal segments 1–4 and base of segment 5 yellow, remainder black. Hypopygium
with epandrium divided at midline by pale membrane. Appendages of hypandrium present … Tipula (Vestiplex) avicularia
- Femora darkened at tip, without preapical yellow ring. Abdominal segments with hypopygium of similar colour or slightly
darker (Figs 1, 21, 38). Hypopygium with epandrium entire, not divided by membrane medially (Figs 7, 24, 43). Appendages
of hypandrium absent .................................................................................. 2
2 Antenna with flagellar segments nearly cylindrical. Epandrium forming sclerotised plate with middorsal furrow, anterior border
not elevated ........................................................ Tipula (Vestiplex) opilionimorpha aligera
- Antenna with flagellar segments distinctly enlarged at base (Fig. 2). Epandrium forming dark brown, polished, sclerotised
saucer-shaped plate (Figs 7, 24, 43) ....................................................................... 3
3. Thorax blackish, heavily dusted with grey pruinescence (Fig. 22). Epandrium rectangular in outline (Fig. 24). Inner gonostylus
without middorsal tooth (Fig. 27) ....................................................Tipula (Vestiplex) mitchelli
- Thorax brownish, thinly dusted with grey pruinescence (Figs 2, 39). Epandrium crescent-shaped (Figs 7, 43). Inner gonostylus
with middorsal tooth (Figs 10, 46) ........................................................................ 4
4 Antenna reaching base of abdomen when bent backwards. Epandrium with large posterolateral tooth; posterior margin with
deep median incision (Figs 5, 7). Outer gonostylus flattened, leaf-shaped (Fig. 9) . . . . . . . Tipula (Vestiplex) levisoni sp. nov.
- Antenna almost reaching middle of the abdomen when bent backwards. Epandrium with distinctly small posterolateral tooth;
NEW VESTIPLEX CRANEFLY FROM HIMALAYAS Zootaxa 5115 (1) © 2022 Magnolia Press · 133
posterior margin with shallow median emargination (Figs 41, 43). Outer gonostylus nearly oval (Fig. 45) . . . . . . . . . . . . . . . .
..............................................................................Tipula (Vestiplex) tanycera
Females
(Note: Females of T. (V.) avicularia remain undescribed)
1 Wing well-developed, extending beyond middle of abdomen (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Wing reduced, brachypterous (Fig. 23) .................................................................... 2
2 Cercus distally slightly expanded with deep, wide serrations. Hypovalva comprising long filament, its length reaching 2/3 of
cercus .............................................................Tipula (Vestiplex) opilionimorpha aligera
- Cercus nearly slender, with gentle serrations (Fig. 32). Hypovalva comprising short filament (Fig. 34), its length reaching base
of cercus .......................................................................Tipula (Vestiplex) mitchelli
3 Body length exceeds 30 mm (33.8–38.2 mm). Cercus with small, indistinct serrations, tip broadly rounded (Fig. 15) ........
....................................................................... Tipula (Vestiplex) levisoni sp. nov.
- Body length less than 20 mm (13.5–15.9 mm). Cercus with distinct serrations on distal end, tip narrowed (Fig. 51) .........
............................................................................. Tipula (Vestiplex) tanycera
Taxonomy
Tipula (Vestiplex) levisoni Starkevich & Podenas sp. nov.
(Figs 1–20)
Type material. HOLOTYPE ♂, labelled: INDIA, Kashmir, Gulmarg, 29.vi.1931, Fletcher coll., B.M.1932.13
[larval exuvium on same pin, male genitalia slide No. PS0375m] (BMNH). PARATYPES: INDIA, same data as
holotype, 26.vii.1931 [genitalia slide No. PS0376f] (1 ♀, BMNH); same data as holotype, 9.vii.1931 (1 ♀, BMNH).
NEPAL, Baitadi, Tinkar Khola, 13,000 ft. [3962 m], 5.vii.1953. J.B. Tyson; W. Nepal Expedition, B.M.1953-592
(1 ♂, BMNH).
Diagnosis. Tipula (V.) levisoni sp. nov. can be recognized by its brownish-yellow body, elongated antenna
which reaches the base of the abdomen and the epandrium forming a large, crescent in outline, saucer-shaped plate
with deep posterior U-shaped notch bordered by denticles on either side. The gonocoxite is unarmed with a rectan-
gular ventromesal part. Females are large with an elongated abdomen and a body length longer than 30 mm. Cercus
slender with upturned tip and ventral margin distally with small, distinct serrations. The hypovalva comprises a
pale, elongated, slender filament. The median incision between the hypovalvae is deeper than the posterior margin
of sternite 8, with the lateral incision deep and narrow.
Description. Male. Body length 18.9–21.0 mm, wing length 20.6–21.2 mm (N=2). General body colour brown-
ish yellow (Figs 1, 2).
Head. Vertex and occiput grey with narrow brown median line (Fig. 2). Rostrum brownish yellow, dorsally
thinly dusted with grey pruinescence. Nasus distinct. Antenna 13-segmented, when bent backward almost reaching
base of abdomen. Scape and pedicel yellow, first flagellar segment yellow, brown at tip, remaining flagellomeres
brown. Each flagellomere, except first, with basal enlargement with small incision and narrowing apex; below nar-
rowing, with additional narrowing at apex, making each segment slightly sinuous. Apical flagellomere small. Lon-
gest verticils approximately same length as their corresponding segments. Palpus with basal segment pale, second
segment pale with brown surface, last segments dark brown.
Thorax. Pronotum yellowish brown with brown median line. Prescutum and presutural scutum with four grey-
ish brown stripes narrowly bordered with brown (Fig. 2). Medial pair of stripes fused in basal half, distally narrowly
separated. Interspaces yellowish grey, anteriorly with small brown dots. Postsutural scutum with lobes each hav-
ing greenish-brown nearly oval spot bordered with brown. Scutellum yellowish brown, mediotergite brown, thinly
dusted, both sclerites with brown medial line. Pleura yellowish brown, thinly dusted with grey pruinescence. Coxae
yellow, thinly dusted, trochanter yellow, femora and tibiae yellow, narrowly darkened at tip; first tarsal segment yel-
low, darkened at tip, remaining tarsal segments dark brown. Tarsal claw without tooth. Wing patterned with brown
(Fig. 4). Halter pale with brown knob.
Abdomen. Brownish yellow, terminal segments brown. Tergites with broad brown median and lateral stripes.
Hypopygium (Figs 5–14). Brown, broader than abdomen. Epandrium forming large, concave, sclerotised sau-
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cer-shaped plate, crescent-shaped in outline (Figs 5–7). Main body of plate brown with blackened rim. Posterior
margin of plate broadly emarginated medially with deep U-shaped notch, laterally bordered by black denticles on
either side; rest of margin toothed. Lateral angle of plate broadly obtuse. Anterior and lateral portions of plate raised
into narrow sclerotised border, posterolaterally terminating in large, black and obtuse teeth. Gonocoxite unarmed,
ventromesal part rectangular in shape (Fig. 8). Outer gonostylus flattened, leaf-shaped (Fig. 9). Inner gonostylus,
large, curved, claw-shaped with single black, large mid-dorsal tooth. Beak extended into blackened, narrowed
rostrum with indistinctly split tip (Fig. 10). Aedeagal guide inshape of narrow, elongated tube, base with dorsal
flange narrow and elongated, membranous at base, ventral flange flattened; distal margin bent inward forming two
microscopic hooks (Fig. 11). Gonocoxal fragment large with lateral and medial sclerites well-developed (Fig. 12).
Medial sclerites fused, anterior apodeme distinct, flattened and narrowed in middle; posterior part broadly flattened,
expanded at base, distal parts flattened and arched. Lateral sclerites U-shaped. Sperm pump with small and flattened
central vesicle (Fig. 13). Compressor apodeme with broad median emargination. Posterior immovable apodeme
longer than compressor apodeme, large and extended laterally into broad plate, medially with darker ridge. Anterior
immovable apodeme nearly triangular, narrowed at lateral margin. Aedeagus with distal part ventrally membranous
(Fig. 14).
Female. Body length 33.8–38.2 mm, wing length 25.9–27.4 mm (N=2). Generally similar to male, but abdo-
men more elongate (Fig. 3).
FIGURES 1–4. Tipula (Vestiplex) levisoni sp. nov. 1, holotype male with larval exuvium, lateral view; 2, head and thorax,
holotype, dorsal view; 3, female, paratype, dorsal view; 4, wing, male, paratype. Scale of 1, 3, 4 = 0.5 mm; 2 = 1 mm.
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FIGURES 5–14. Male terminalia of Tipula (Vestiplex) levisoni sp. nov. 5, hypopygium, dorsal view; 6, hypopygium, lateral
view; 7, epandrium, dorsal view; 8, left gonocoxite, lateral view; 9, left outer gonostylus; 10, left inner gonostylus, lateral view;
11, aedeagal guide, lateral view; 12, gonocoxal fragment, dorsal view; 13, sperm pump, dorsal view; 14, distal part of intromit-
tent organ, lateral view. Abbreviations: aia, anterior immovable apodeme; ca, compressor apodeme; ep, epandrium; gcx, gono-
coxite; ig, inner gonostylus; ls, lateral sclerite of gonocoxal fragment; ms, medial sclerite of gonocoxal fragment; mt, mid-dorsal
tooth of inner gonostylus; og, outer gonostylus; pia, posterior immovable apodeme; pt, posterolateral tooth of epandrium. Scale
a: 5, 6 = 0.5 mm; b: 7–13 = 0.5 mm; c: 14 = 0.25 mm.
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FIGURES 15–20. Female terminalia of Tipula (Vestiplex) levisoni sp. nov. 15, ovipositor, left lateral view; 16, sternite 8 with
hypovalvae, ventral view; 17, sternite 9, dorsal view; 18, furca, dorsal view; 19, bursa copulatrix, dorsal view; 20, spermatheca.
Abbreviations: bp sd, basal part of spermathecal duct; cds, cul-de-sac of bursa copulatrix; c, cerci; h, hypovalvae; s8, sternite 8;
t10, tergite 10. Scale a: 15 = 1 mm; b: 16–20 = 0.5 mm.
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Female terminalia (Figs 15–20). Tergite 10 shiny brown. Cercus brownish yellow, about as long as tergite 10,
slender with up-turned tip (Fig. 15). Ventral margin distally with small, distinct serrations. Hypovalva pale, elongat-
ed, filamentous, with short trichia at tip (Fig. 15). Median incision between hypovalvae deeper than posterior margin
of sternite 8. Lateral incision deep and narrow, about three times as deep as maximum width. Posterolateral angle
of sternite 8 rounded. Sternite 9 with posterior part shovel-shaped, margin with lateral incisions, medially forming
groove; anterior parts flattened (Fig. 17). Furca flattened, base narrow (Fig. 18). Bursa copulatrix with spermathecal
ducts sclerotised at the base, in shape of curved, slender, dark brown sticks (Fig. 19). Cul-de-sac of bursa copulatrix
curved. Spermatheca pear-shaped (Fig. 20).
Etymology. The new species is named after Levison James Wood (1982–), a British army officer, explorer and
writer, best known for his remarkable expeditions in the Himalayas, Central America, the Arabian Peninsula and
walking the entire length of the River Nile.
Distribution. India (Jammu and Kashmir), Nepal (Sudurpashchim Province).
Remarks. Tipula (V.) levisoni sp. nov. is assigned here as a member of T. (V.) leucoprocta species group pro-
posed by Savchenko (1960, 1964) and as discussed by Starkevich et al. (2020). Males of the leucoprocta species
group are distinguished by an unarmed gonocoxite, the epandrium having a large sclerotised plate with an extensive
saucer with a raised border, posterolaterally terminating in teeth and a remarkably large gonocoxal fragment. In fe-
males, the bursa copulatrix and basal part of the spermathecal ducts are sclerotised. Two members of the leucoprocta
species group are known from the Western Himalayas, T. (V.) tanycera (Pakistan) and T. (V.) mitchelli (India).
Tipula (V.) levisoni sp. nov. is closest to T. (V.) tanycera based on body colouration and the shape of the hy-
popygium. Both species are characterized by a crescent-shaped epandrium, inner gonostylus with middorsal tooth,
slender aedeagal guide and the shape of the gonocoxal fragment. Females are characterized by sternite 8 with deep
lateral incisions.
Males can be separated by their antennae which reaches the base of the abdomen in T. (V.) levisoni sp. nov. while
in T. (V.) tanycera the antenna is longer,almost reaching the middle of the abdomen. They can also be separated by
the following details of the epandrium: T. (V.) levisoni sp. nov. has the posterior margin with a deep median notch
and a large posterolateral tooth (Fig. 7), while in T. (V.) tanycera the median notch is shallower and has a distinctly
smaller posterolateral tooth (Fig. 43). The outer gonostylus of T. (V.) levisoni sp. nov. is flattened, leaf-shaped (Fig.
9), while in T. (V.) tanycera it is nearly oval (Fig. 45). The inner gonostylus bears a middorsal tooth with a broadly
expanded base in T. (V.) levisoni sp. nov. (Fig. 10). Tipula (V.) tanycera has a middorsal tooth with a distinctly less
expanded base (Fig. 46).
Females can be separated by their body size which exceeds 30 mm in T. (V.) levisoni sp. nov. and is less than 20
mm in T. (V.) tanycera. The cercus has small serrations and a nearly rounded tip in T. (V.) levisoni sp. nov. (Fig. 15),
while in T. (V.) tanycera the serrations are distinct on the distal end and the cercus tip is pointed (Fig. 51).
Another regional species related to T. (V.) levisoni sp. nov. and T. (V.) tanycera is T. (V.) mitchelli, which can be
separated by a rectangular epandrium and the inner gonostylus lacking a middorsal tooth.
Tipula (Vestiplex) mitchelli Edwards
(Figs 21–37)
Tipula mitchelli Edwards, 1927: 229–230.
Tipula (Vestiplex) mitchelli: Alexander, 1942: 39; 1953: 342; Savchenko, 1960: 212; 1964: 202–203; Alexander & Alexander,
1973: 64.
Material examined. INDIA, Kashmir, Sheeshnag, 14,000 ft. [4267 m], 14.vii.1960, S.W. Batra (4 ♂, SEM); Kash-
mir, Sheeshnag, 16,000 ft. [4877 m], 14.vii.1960, S.W. Batra [female genitalia slide No. PS0379f] (1 ♀, SEM);
Kashmir, Killarmarg, 10,000 ft. [3048 m], 19.vii.1923, Fletcher; India. Pres.by T.B. Fletcher. B.M.1925-546, male
genitalia slide No. PS0373m (1 ♂, BMNH); Kashmir, Killarmarg, 10,000 ft. [3048 m], 21.vii.1923, Fletcher; India.
Pres. by T.B. Fletcher. B.M.1925-546 (1 ♂, BMNH).
Redescription. Male. Body length 15.9–16.2 mm, wing length 18.1–18.5 mm. General body colour yellow
(Figs 21, 22).
Head. Vertex and occiput grey with dark brown median line (Fig. 22). Rostrum yellow, dorsally thinly dusted
with grey pruinescence. Nasus distinct. Antenna 13-segmented, when bent backwards extends beyond base of abdo-
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men. Scape and pedicel yellow, flagellum brown. Each flagellomere, except first, with basal enlargement and small
incision. Apical flagellomere small. Distal flagellar segments sinuous. Longest verticils about three times shorter
than corresponding segments. Palpus brown.
Thorax. Pronotum yellow with medial brown spot or line. Prescutum and presutural scutum with four bluish
grey stripes, narrowly bordered with brown. Brown borders of medial stripes fused (Fig. 22). Interspaces grey. Post-
sutural scutum grey, each lobe with two bluish grey, spots without borders. Scutellum yellow, mediotergite yellow,
thinly dusted, both sclerites with brown line. Pleura brownish yellow. Coxae yellowish, thinly dusted, trochanters
yellow, femora and tibiae yellow, narrowly darkened at tip; first two tarsal segments yellow, distally darker with
blackened tips, last segments dark brown. Tarsal claw without tooth. Wing patterned with brown. Halter pale with
brown knob.
Abdomen. Abdominal segments yellow with distinct brown dorsal and lateral stripes.
Hypopygium (Figs 24–31). Yellowish brown. Epandrium distally forming rectangular, saucer-shaped plate (Fig.
24). Plate with straight posterior margin, medially with rounded projection, small denticles on either side, and ob-
tuse lateral corners. Anterior margin elevated into blackened border terminating in blackened obtuse angles. Gono-
coxite unarmed, ventromesal part short, nearly triangular in shape (Fig. 25). Outer gonostylus narrowed at base and
flattened in middle (Fig. 26). Inner gonostylus in form of curved plate, with triangular beak blackened at tip and
right angle to rest of sclerite (Fig. 27). Aedeagal guide in shape of slender tube, base with dorsal flange narrow,
membranous at base, ventral flange flattened (Fig. 28). Gonocoxal fragment large with lateral and medial sclerites
well-developed (Fig. 29). Medial sclerites fused, anterior apodeme distinct, slightly flattened, posterior part broadly
flattened and arched distally. Lateral sclerites broadly U-shaped, expanded at base. Sperm pump with flattened cen-
tral vesicle (Fig. 30). Compressor apodeme with median emargination. Posterior immovable apodeme longer than
compressor apodeme, large and extended laterally into broad plate, medially with darker ridge. Anterior immovable
apodeme widely rounded. Aedeagus with distal part ventrally membranous and nearly scoop-shaped (Fig. 31).
Female. Body length 26.9 mm, wing length 4.7 mm. Generally similar to male (Fig. 23). Tergites and sternites
with pale margins. Wing greatly reduced, brachypterous.
FIGURES 21–23. Tipula (Vestiplex) mitchelli. 21, male, lateral view; 22, male, head and thorax, dorsal view; 23, female, lateral
view. Scale of 21, 23 = 0.5 mm; 22 = 1 mm.
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FIGURES 24–31. Male terminalia of Tipula (Vestiplex) mitchelli. 24, epandrium, dorsal view 25, left gonocoxite, lateral view;
26, left outer gonostylus; 27, right inner gonostylus, lateral view; 28, aedeagal guide, lateral view; 29, gonocoxal fragment,
dorsal view; 30, sperm pump, dorsal view; 31, distal part of intromittent organ, lateral view. Scale a: 24–30 = 0.5 mm; b: 31 =
0.25 mm.
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FIGURES 32–37. Female terminalia of Tipula (Vestiplex) mitchelli. 32, tergite 10 with cercus, left lateral view; 33, distal part
of cercus, lateral view; 34, sternite 8 with hypovalvae, ventral view; 35, sternite 9 with furca, dorsal view; 36, bursa copulatrix,
dorsal view; 37, spermatheca. Abbreviation: cds, cul-de-sac of bursa copulatrix. Scale a: 32 = 1 mm; b: 33–37 = 0.5 mm.
Female terminalia (Figs 32–37). Tergite 10 shiny brown. Cercus brown, slender, with distal part strongly up-
curved (Fig. 32). Ventral margin distally with small serrations (Figs 32, 33). Hypovalva short, filamentous (Fig. 34).
Median incision between hypovalvae deeper than posterior margin of sternite 8. Lateral angle of sternite 8 rounded.
Sternite 9 with lateral parts parallel-sided, posterior part broad and flattened (Fig. 35). Furca posteriorly flattened,
anteriorly narrowed (Fig. 35). Bursa copulatrix with spermathecal ducts sclerotised at base; cul-de-sac of bursa
copulatrix curved (Fig. 36). Spermatheca broadened at base, nearly pear-shaped (Fig. 37).
Distribution. India (Jammu and Kashmir).
Remarks. Tipula mitchelli was described by Edwards in 1927 based on two males collected by F.J. Mitchell
from Kashmir, although the exact locality is unknown (Edwards 1927). Unfortunately, the type specimens of T.
mitchelli were not found in BMNH.
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The association of the female specimen with this species are based on the similar general body coloration; the
female belongs to the leucoprocta species complex which can be easily distinguished by the specific bursa copula-
trix; specimens were collected at the same locality, same day with differences in 600 meters at the altitudes lying in
the zone of subalpine meadows.
Tipula (Vestiplex) tanycera Alexander
(Figs 38–56)
Tipula (Vestiplex) tanycera Alexander, 1961: 140–142.
Tipula (Vestiplex) tanycera: Alexander and Alexander, 1973: 65.
Type material examined. HOLOTYPE ♂, labelled: Salf-ul-Maluk, Northwest Frontier Province [Pakistan],
11,000 ft. [3350 m], along lake margin, 1953.vii.2, Schmid (USNM). PARATYPES: same data as holotype (4 ♂,
1 ♀, USNM) [among them 1 ♂ and 1 ♀ (female labeled as allotype) pinned together on same pin, both dissected for
present study, male genitalia number No. PS0377m, female genitalia number No. PS0378f and other male with an-
tenna, wing and genitalia on slide mounted by C.P. Alexander]; Kalarian Baihk Northwest Frontier Province, 11,270
ft. [3435 m], along lake delta, 1953.vii.13, Schmid (5 ♂, USNM) [among them one ♂ with antenna, leg, wing and
genitalia on slide mounted by C.P. Alexander].
FIGURES 38–40. Tipula (Vestiplex) tanycera. 38, male and female, paratypes; 39, male, thorax, dorsal view; 40, wing. Scale
of 38, 40 = 0.5 mm; 39 = 1 mm.
STARKEVICH ET AL.
142 · Zootaxa 5115 (1) © 2022 Magnolia Press
FIGURES 41–50. Male terminalia of Tipula (Vestiplex) tanycera. 41, hypopygium, dorsal view; 42, hypopygium, lateral view;
43, epandrium, dorsal view; 44, left gonocoxite, lateral view; 45, left outer gonostylus; 46, left inner gonostylus, lateral view; 47,
aedeagal guide, lateral view; 48, gonocoxal fragment, dorsal view; 49, sperm pump, dorsal view; 50, distal part of intromittent
organ, lateral view. Abbreviations: mt, mid-dorsal tooth of inner gonostylus; pt, posterolateral tooth of epandrium. Scale a: 41,
42 = 0.5 mm; b: 43–49 = 0.5 mm; c: 50 = 0.25 mm.
NEW VESTIPLEX CRANEFLY FROM HIMALAYAS Zootaxa 5115 (1) © 2022 Magnolia Press · 143
FIGURES 51–56. Female terminalia of Tipula (Vestiplex) tanycera. 51, ovipositor, left lateral view; 52, sternite 8 with hy-
povalvae, ventral view; 53, sternite 9, dorsal view; 54, furca, dorsal view; 55, bursa copulatrix, dorsal view; 56, spermatheca.
Abbreviation: cds, cul-de-sac of bursa copulatrix. Scale a: 50 = 1 mm; b: 51–54 = 0.5 mm.
STARKEVICH ET AL.
144 · Zootaxa 5115 (1) © 2022 Magnolia Press
Redescription. Male. Body length 17.3–20.0 mm, wing length 19.4–21.0 mm. General body colouration
brownish yellow (Fig. 38).
Head. Vertex and occiput yellowish grey pruinescent with brown median line. Rostrum brown, dorsally thinly
dusted with grey pruinescence. Nasus short and distinct. Antenna 13-segmented, when bent backwards extends
beyond wing base. Scape and pedicel yellow, flagellum brown. Each flagellomere, except first, with weak basal
enlargement and incision, and additional apical incision; shape of segments slightly sinuous. Apical flagellomere
small. Longest verticils shorter than corresponding segments. Palpus with first segment yellowish, second brownish
yellow, other brownish black.
Thorax. Whole thorax brown, dusted with grey pruinescence (Fig. 39). Pronotum brown, with median brown
line. Prescutum and presutural scutum with four brown, grey dusted stripes, bordered by brown. Medial stripes
fused in basal half. Interspaces grey. Postsutural scutum grey, each lobe with two brown spots, bordered by brown.
Scutellum and mediotergite brown, dusted with grey, with brown median line. Pleura brown. Coxae yellow, dusted
with grey, trochanters yellow, femora and tibiae yellow, darkened at tip; first two tarsal segments yellow, distally
brown, rest of segments brown. Tarsal claw without tooth. Wing patterned with brown (Fig. 40). Halter yellowish
brown with brown knob.
Abdomen. Brownish yellow, with brown dorsal and lateral stripes, terminal segments brown.
Hypopygium (Figs 41–50). Brown, broader than rest of abdomen. Epandrium forming large concave, sclero-
tized, crescent in outline saucer-shaped plate (Figs 41–43). Main body of plate brownish yellow with blackened rim.
Posterior margin of plate broadly emarginated, medially with shallow U-shaped emargination, laterally bordered
with obtuse projections or denticles (Alexander 1961, pl. 4, fig. 33; Starkevich 2012, fig. 142) and additional small
denticles on either side. Lateral angle of plate broad and obtuse. Anterior and lateral portions of plate raised into
sclerotised rim, posterolaterally terminating in black obtuse teeth. Gonocoxite unarmed (Fig. 44). Outer gonosty-
lus flattened, nearly oval (Fig. 45). Inner gonostylus large, claw-shaped curved plate with single, large mid-dorsal
tooth (Fig. 46); beak extends into blackened rostrum. Aedeagal guide in shape of narrow, elongated tube, base with
dorsal flange handle-shaped, membranous at base, ventral flange flattened; distal margin bent inward forming two
microscopic hooks (Fig. 47). Gonocoxal fragment large with lateral and medial sclerites well-developed (Fig. 48).
Medial sclerites fused, anterior apodeme distinct, slightly flattened, posterior part broadly flattened and expanded
at base and arched distally. Lateral sclerites nearly V-shaped. Sperm pump with small and flattened central vesicle
(Fig. 49). Compressor apodeme with median shallow emargination. Posterior immovable apodeme longer than
compressor apodeme, large and extended laterally into broad plate, medially with darker ridge, anterior margin
bent inwards. Anterior immovable apodeme nearly rounded, narrowed at lateral margin. Aedeagus with distal part
ventrally membranous (Fig. 50).
Female. Body length 23.4 mm, wing length 19.4 mm. Generally similar to male in body colour (Fig. 38).
Female terminalia (Figs 51–56). Tergite 10 shiny brown. Cercus brownish yellow, about as long as tergite 10,
slender, narrowing towards upcurved tip (Fig. 51). Ventral margin distally with distinct serrations. Hypovalva in
shape of pale, elongated slender filament, with short trichia at tip (Fig. 52). Median incision between hypovalvae
deeper than posterior margin of sternite 8. Lateral incision relatively deep and broad, about 1.5 times as deep as
maximal width. Posterolateral angle of sternite 8 rounded. Sternite 9 with posterior part shovel-shaped, margin
rounded, medially forming groove; anterior parts flattened at base (Fig. 53). Furca anteriorly narrow but posteriorly
broad (Fig. 54). Bursa copulatrix with spermathecal ducts sclerotised at base, in shape of slightly curved, slender,
dark brown sticks (Fig. 55). Cul-de-sac of bursa copulatrix curved. Spermatheca nearly spherical (Fig. 56).
Distribution. Pakistan (Khyber Pakhtunkhwa [= North-West Frontier Province]) (Alexander 1961).
Remarks. The species is known only from the type series collected near Salf ul Maluk Lake, a mountainous
lake located at the northern end of the Kaghan Valley, in Saiful Muluk National Park, northern Pakistan.
Acknowledgements
We are grateful for the following Museum curators for their help and assistance for obtaining specimens: J.C. Thom-
as (SEM), Dr. F. Shockley and Dr. T. Dikow (USNM), Dr. J.K. Gelhaus (Academy of Natural Sciences of Drexel
University, USA) for the possibility to use specimens from the USNM collections and Dr. N. Paramonov (ZIN).
Special thanks to Dr. R.I. Vane-Wright for his kind support and communication while working on this manuscript.
NEW VESTIPLEX CRANEFLY FROM HIMALAYAS Zootaxa 5115 (1) © 2022 Magnolia Press · 145
The visit of Dr. P. Starkevich to BMNH was supported from the SYNTHESYS Project http://www.synthesys.info/,
financed by European Community Research Action under the FP7 Integrating Activities Programme. The visit of
Dr. S. Podėnas to BMNH was supported from the SYNTHESYS+ Project http://www.synthesys.info/ financed by
European Community Research Infrastructure Action under the H2020 Integrating Activities Programme, Project
number 823827.
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