Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species
Author
Tan, Ming Kai
2F544BE6-7986-4905-AE51-CF16B44D5251
Institut de Systématique, Evolution et Biodiversité (ISYEB), Muséum national dʼhistoire naturelle, CNRS, SU, EPHE, UA, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France.
mingkai@gmail.com
Author
Wahab, Rodzay bin Haji Abdul
50868E6A-34E4-471A-B941-C504CC9E895A
Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, Jalan Universiti, BE 1410, Brunei Darussalam.
rodzay.wahab@ubd.edu.bn
Author
Japir, Razy
B05E507D-FBA4-437F-AB03-94E349912EA5
Forest Research Centre (Sepilok), Sabah Forestry Department, P. O. Box 1407, 90715 Sandakan, Sabah, Malaysia.
razy.Japir@sabah.gov.my
Author
Chung, Authur Y. C.
6E7371D6-6FF9-42DF-8745-4A8304CB0978
Forest Research Centre (Sepilok), Sabah Forestry Department, P. O. Box 1407, 90715 Sandakan, Sabah, Malaysia.
arthur.Chung@sabah.gov.my
Author
Robillard, Tony
19BBE007-85D2-4325-8396-A1F4BAF5775C
Institut de Systématique, Evolution et Biodiversité (ISYEB), Muséum national dʼhistoire naturelle, CNRS, SU, EPHE, UA, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France.
tony.robillard@mnhn.fr
text
European Journal of Taxonomy
2021
2021-07-28
761
1
1
75
http://dx.doi.org/10.5852/ejt.2021.761.1449
journal article
10.5852/ejt.2021.761.1449
bf24ff1f-569a-4222-afe1-8527c83d8003
2118-9773
5156259
519E2F56-40E1-4431-BC90-8484E308D16E
Genus
Nisitrus
Saussure, 1878
Nisitra
Walker, 1869: 91
.
Nisitra
–
Chopard 1940: 199
.
Nisitrus
–
Saussure 1878: 511
, 522 (nomen novum for
Nisitra
Walker
). —
Chopard 1968: 352
. —
Desutter-Grandcolas 1990: 239
. —
Otte 1994: 67
. —
Preston-Mafham 2000: 2241
(behaviour). —
Robillard & Desutter-Grandcolas 2004a: 276
;
2004b: 578
;
2004c: 304
;
2006: 644
;
2008: 67
(
Nisitrini
tribe); 2011: 637. —
Robillard
et al.
2007: 1265
. —
Desutter-Grandcolas
et al.
2010: 616
. —
Nattier
et al.
2011: 2201
(phylogeny and molecular dating). —
Robillard & Tan 2013: 707
(redescription and diagnosis). —
Vicente
et al.
2017: 2203
(phylogeny, historical biogeography). —
Cigliano
et al.
2020
(
Orthoptera
Species File Online).
Type
species
Nisitra vittatus
(
Haan, 1844
)
.
Remarks on
type
species
Kirby (1906)
and
Chopard (1968)
listed
N. vittata
as the
type
species of
Nisitra
and treated the original
type
species
N. marginata
Walker, 1869
as a synonym of
N. vittata
. Subsequent authors listed
N. vittatus
as the
type
species of the replacement name
Nisitrus
, although
Otte (1994)
reinstated
N. marginatus
as a valid species (Braun
2013 in
Cigliano
et al.
2020
). The present revision revalidates the synonymy of
N. marginatus
under
N. vittatus
, making
N. vittatus
the valid
type
species of the genus.
Diagnosis
(emended from
Robillard & Tan 2013
)
Among genera of
Eneopterinae
,
Nisitrus
is characterised by an elegant, wasp-like, slender and colourful body (
Figs 7–9
). Size average for subfamily. Colouration vivid, dominantly yellow, whitish and black, with species-specific patterns, particularly in the head (
Figs 10–11
), pronotum (
Figs 11–12
) and FWs (
Figs 12–14
). Characteristic head shape with narrow fastigium and large prominent eyes with thin dorso-ventral black stripes. Pronotum setose, square-shaped to slightly trapezoidal, usually bisinuated posteriorly. Metanotum with large glandular structures in males, made of a wide pit on scutum and a pair of posterior pits on scutellum, with glandular pores mostly organised by groups of 2 or 3 (
Fig. 15
). FWs more or less as long as abdomen. Male
Nisitrus
show modified FW venation and characteristic glossy transparency. Hindwings longer than FWs, bicoloured in most species, their base transparent with a wide dark posterior margin. Legs long and thin, FIII with a characteristic narrowed area before knee. Other general traits: TI with two tympana, inner one slit-like, covered by a swollen cuticular expansion; outer tympanum oval, its membrane transversally plicate in dorsal half. TI with three apical spurs; outer dorsal spur missing. TII with four apical spurs, inner longest. TIII with three inner and three outer apical spurs, median longest on each side; four pairs of subapical spurs, inner and outer spurs almost straight, their apex hook-like; TIII serrulate over whole length, slightly furrowed dorsally. Tarsomeres III-1 with one row of dorsal spines on external edge in addition to apical spines. Apical claws of legs slightly indented. Both males and females show characteristic genitalia.
Fig. 7.
Habitus of ♂♂ in dorsal view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. hughtani
Robillard & Tan
sp. nov.
D
.
N. insignis
Saussure, 1878
.
E
.
N. malaya
Robillard & Tan
sp. nov.
F
.
N. musicus
Ingrisch, 1987
.
G
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 10 mm.
Fig. 8.
Habitus of ♀♀ in dorsal view.
A–B
.
Nisitrus brunnerianus
Saussure, 1878
.
C
.
N. crucius
Robillard & Tan
sp. nov.
D
.
N. danum
Robillard & Tan
sp. nov.
E
.
N. hughtani
Robillard & Tan
sp. nov.
F
.
N. insignis
Saussure, 1878
. Scale bar = 10 mm.
Fig. 9.
Habitus of ♀♀ in dorsal view.
A
.
Nisitrus malaya
Robillard & Tan
sp. nov.
B
.
N. musicus
Ingrisch, 1987
.
C–D
.
N. rindu
Robillard & Tan
sp. nov.
E
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 10 mm.
Fig. 10.
Face in anterior view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. danum
Robillard & Tan
sp. nov.
D
.
N. hughtani
Robillard & Tan
sp. nov.
E
.
N. insignis
Saussure, 1878
.
F
.
N. malaya
Robillard & Tan
sp. nov.
G
.
N. musicus
Ingrisch, 1987
.
H
.
N. rindu
Robillard & Tan
sp. nov.
I
.
N. vittatu
s (
Haan, 1844
). Scale bar = 2 mm.
Fig. 11.
Head and pronotum in dorsal view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. danum
Robillard & Tan
sp. nov.
D
.
N. hughtani
Robillard & Tan
sp. nov.
E
.
N. insignis
Saussure, 1878
.
F
.
N. malaya
Robillard & Tan
sp. nov.
G
.
N. musicus
Ingrisch, 1987
.
H
.
N. rindu
Robillard & Tan
sp. nov.
I
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 2 mm.
Fig. 12.
Body in lateral view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. danum
Robillard & Tan
sp. nov.
D
.
N. hughtani
Robillard & Tan
sp. nov.
E
.
N. insignis
Saussure, 1878
.
F
.
N. malaya
Robillard & Tan
sp. nov.
G
.
N. musicus
Ingrisch, 1987
.
H
.
N. rindu
Robillard & Tan
sp. nov.
I
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 5 mm.
Fig. 13.
Male tegmina in dorsal view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. hughtani
Robillard & Tan
sp. nov.
D
.
N. insignis
Saussure, 1878
.
E
.
N. malaya
Robillard & Tan
sp. nov.
F
.
N. musicus
Ingrisch, 1987
.
G
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 2 mm.
Fig. 14.
Female tegmina in dorsal view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. crucius
Robillard & Tan
sp. nov.
C
.
N. danum
Robillard & Tan
sp. nov.
D
.
N. hughtani
Robillard & Tan
sp. nov.
E
.
N. insignis
Saussure, 1878
.
F
.
N. malaya
Robillard & Tan
sp. nov.
G
.
N. musicus
Ingrisch, 1987
.
H
.
N. rindu
Robillard & Tan
sp. nov.
I
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 2 mm.
Male genitalia (
Fig. 16
) with membranous and setose lophi well individualised; pseudepiphallic parameres including two membranous posterior lobes and a sclerotized anterior lobe oriented anteriorly toward median axis of genitalia and carrying a secondary preapical sclerotized lobe; ectophallic arc transverse, incomplete due to a little-sclerotized median area; ectophallic fold narrowed preapically, its apex narrow, located between anterior apex of pseudepiphallic parameres; dorsal part of fold sclerotized, forming two elongated sclerites connected to the median area of arc; ventral part of ectophallic fold membranous, with two strong rounded lateral sclerites of variable shapes. Membrane of endophallic cavity smooth. Female copulatory papilla (
Fig. 17
) conical, apex generally sclerotized and pointed. Ovipositor typically longer than FIII; apex thin with both dorsal and ventral edges smooth.
Phylogenetic relationships
Based on previous phylogenetic studies (
Robillard & Desutter-Grandcolas 2004c
;
Nattier
et al.
2011
;
Vicente
et al.
2017
),
Nisitrus
is the sister genus of
Paranisitra
Chopard, 1925
(revision in
Gorochov 2009
and
Baroga-Barbecho
et al.
2019
). The two genera form the
Nisitrini
tribe (
Robillard & Desutter-Grandcolas 2008
) and are very close morphologically;
Paranisitra
is mostly characterised by a lack of wings in both sexes, but except for this, it shares with
Nisitrus
its general body shape, long thin legs with indented claws, colouration with yellow and black, head shape with large prominent eyes and narrow fastigium, and the general structure of male and female genitalia.
Nisitrini
are less easy to relate to other clades of
Eneopterinae
.
Natural history
Species tend to be found at forest edges and gaps in the forest where the vegetation is more exposed. They are also found among shrubs and herbaceous plants (but not grasses).
Nisitrus
are observed to be generally diurnal, during which the males can be heard calling and individuals are found foraging in the vegetation. It seems as
Nisitrus
are mostly herbivorous, feeding on leaves, but also on flowers (
Tan
et al.
2017
).
Nisitrus
are also observed to congregate with one another on the same plant, sometimes making up of more than 20 individuals of both adults and nymphs. They can be considered a pest in some parts of Southeast Asia (
Willemse 2001
).
Included species
Nisitrus brunnerianus
from Borneo and
Java
;
N. crucius
sp. nov.
and
N. danum
sp. nov.
(both from Borneo);
N. hughtani
sp. nov.
from
Sumatra
;
N. hyalinus
from Borneo;
N. insignis
from
Sumatra
and Malay Peninsula;
N. malaya
sp. nov.
from Malay Peninsula;
N. musicus
and
N. rindu
sp. nov.
from Borneo; and
N. vittatus
from
Sumatra
and Borneo.
Fig. 15.
Metanotal gland.
A
.
Nisitrus hughtani
Robillard & Tan
sp. nov.
B
.
N. brunnerianus
Saussure, 1878
. Scale bar = 2 mm.
Fig. 16.
Male genitalia in ventral view.
A
.
Nisitrus brunnerianus
Saussure, 1878
.
B
.
N. cruciu
s Robillard & Tan sp. nov.
C
.
N. hughtani
Robillard & Tan
sp. nov.
D
.
N. insignis
Saussure, 1878
.
E
.
N. malaya
Robillard & Tan
sp. nov.
F
.
N. musicus
Ingrisch, 1987
.
G
.
N. vittatus
(
Haan, 1844
)
. Scale bar = 1 mm.
Fig. 17.
Female genitalia in lateral (A, C, E, F, H, J, L, N, P) and dorsal (B, D, G, I, K, M, O, Q) views.
A–B
.
Nisitrus brunnerianus
Saussure, 1878
.
C–D
.
N. crucius
Robillard & Tan
sp. nov.
E
.
N. danum
Robillard & Tan
sp. nov.
F–G
.
N. hughtani
Robillard & Tan
sp. nov.
H–I
.
N. insignis
Saussure, 1878
.
J–K
.
N. malaya
Robillard & Tan
sp. nov.
L–M
.
N. musicus
Ingrisch, 1987
.
N–O
.
N. rindu
Robillard & Tan
sp. nov.
P–Q
.
N. vittatus
(
Haan, 1844
)
. Scale bar =
0.5 mm
.
Distribution
Borneo (
Sabah
and
Sarawak
States of East
Malaysia
,
Brunei Darussalam
, and
Kalimantan State
of
Indonesia
),
Java
, Malay Peninsula (Peninsular
Malaysia
and
Singapore
) and
Sumatra
(
Indonesia
) (
Fig. 5
).
Key to the species of
Nisitrus
Saussure, 1878
1. Frons mostly black, or with large dark markings (e.g., cross-shaped pattern); gena usually black, although may have yellow band ventral of eye ................................................................................ 2
– Frons mostly yellow, sometimes with small dots but without large dark markings; gena usually yellow ............................................................................................................................................... 5
2. FW with longitudinal white stripe around region of CuA, M and R with a distinct margin from black colouration on dorsal field; lateral field usually infumated black ventrally. Distribution:
Sumatra
...................................................................................
N. hughtani
Robillard & Tan
sp. nov.
– FW with longitudinal white stripe around region of CuA, M and R darkens progressively towards the dorsal field without a distinct margin from black colouration on dorsal field; lateral field usually light coloured or transparent ventrally. Distribution: Borneo and
Java
............................................ 3
3. Smaller in size, female FWL typically <
11.3 mm
. Frons with a characteristic black cross marking ....................................................................................................
N. crucius
Robillard & Tan
sp. nov.
– Larger in size, female FWL typically>
11.8 mm
. Frons completely black; without a characteristic black cross marking .......................................................................................................................... 4
4. Dorsum of head completely black. Pronotal lateral lobe either with ventral half yellow or with an oval yellow spot at the anterior end ................................................
N. brunnerianus
Saussure, 1878
– Dorsum of head pale with some dark markings. Pronotal lateral lobe with a characteristic dorsal yellow band that extends ventrally ...............................................
N. rindu
Robillard & Tan
sp. nov.
5. Frons with black band between eyes, mouthparts mostly yellow. Body darker; female FW typically with infumated black cells, with lateral parts of dorsal field usually with an infumated yellow longitudinal stripe. Distribution: Borneo (
Sabah
) ......................
N. danum
Robillard & Tan
sp. nov.
– Frons without black band between eyes, mouthparts black. Body usually with more yellow colouration; male FW with some black and white infumation, but generally transparent. Distribution: Borneo, Malay Peninsula and
Sumatra
.......................................................................................................... 6
6. Legs usually pale in colouration; FIII with at least one black longitudinal stripe on external face. Male FW with distinct infumated white spots and infumated black pattern near base, sometimes with infumated white patterns around chords, but otherwise transparent, harp very small, with one distinct straight oblique vein; female FW almost always with yellow/white veins. Distribution:
Sumatra
....................................................................................................
N. insignis
Saussure, 1878
– Legs usually brown in colouration; FIII with faint or without black longitudinal stripe on external face. Male FW with infumated black pattern near base, sometimes with infumated white spots, but otherwise transparent, harp larger, with 3 oblique veins; female FW sometimes with yellow/white veins. Distribution:
Sumatra
, Borneo, Malay Peninsula ................................................................... 7
7. Eyes dark when alive. Frons yellow, face part of fastigium (including scapes) and mouthparts black, maxillary palpi yellow; dorsal part of legs black. Male pseudepiphallus slender. Distribution: Borneo, near Kinabalu area ....................................................................................
N. musicus
Ingrisch, 1987
– Eyes green when alive. Frons yellow to brown, mouthparts sometimes yellow or brown; legs typically homogenous brown. Pseudepiphallus more robust, broad. Distribution:
Sumatra
, Borneo, Malay Peninsula ........................................................................................................................................... 8
8. Lighter in colouration. Lateral field of FWs usually light in colouration. Juveniles of pale white colouration with numerous dark stripes all around the body. Female copulatory papilla without truncated lobule at the ventro-anterior end. Distribution: Borneo,
Sumatra
...................................... ......................................................................................................................
N. vittatus
(
Haan, 1844
)
– Darker in colouration. Lateral field of FWs usually dark in colouration. Juveniles of darker colouration, usually grey, their pronotum without longitudinal stripes but dorsal disk sometimes with a large velvety black spot. Female copulatory papilla with a truncated lobule at the ventro-anterior end. Distribution: Malay Peninsula ...................................................
N. malaya
Robillard & Tan
sp. nov.