Taxonomic revision of the genus Xenogryllus Bolívar, 1890 (Orthoptera, Gryllidae, Eneopterinae, Xenogryllini)
Author
Jaiswara, Ranjana
Author
Dong, Jiajia
Author
Ma, Libin
Author
Yin, Haisheng
Author
Robillard, Tony
text
Zootaxa
2019
2019-01-18
4545
3
301
338
journal article
27632
10.11646/zootaxa.4545.3.1
b4e5a26a-0e84-4210-b2f1-284545cc1b8a
1175-5326
2618876
A6A7B9CE-9905-4DA7-8011-747D4B9325F7
Xenogryllus marmoratus
(
Haan, 1844
)
(
Figs 1
D–G, 3I–J, 4E, 5E, 7H–I, 8G, 9B, 11CD, 12B, 14)
Gryllus
(
Phalangopsis
)
marmoratus
Haan, 1844
: 235
.
Calyptotrypus
marmoratus—
Saussure 1878
: 714
(the figures do not correspond to the species
X. marmoratus
, which is consistent with the surprising distribution “Java, Iles de la Sonde” proposed by Saussure, where the species is not distributed);
Bolívar 1900
[1899]: 805.
Madasumma marmorata
—Chopard 1924: 56:
Kirby 1906
: 93
.
Xenogryllus marmoratus
—
Chopard 1968
: 350
;
Yin & Liu 1995
: 96
;
Oshiro 1995
: 43
;
Walker 2010
: 27
; Robillard & Desutter- Grandcolas 2004a: 578 (calling song); 2004b: 273 (morphological phylogeny); 2006: 644 (molecular phylogeny); 2008: 67; 2011: 637;
Anso
et al.
2016
: 9
(molecular phylogeny);
Chintauan-Marquier
et al.
2016
: 62
(molecular phylogeny);
Vicente
et al.
2017
: 2203
(historical biogeography);
Schneider
et al.
2017
(tympanum morphology)
Jing
et al
. 2018
: 274
.
Xenogryllus marmoratus marmoratus
Gorochov, 1992
, in
Gorochov & Belokobylskij 1992
: 11
(nominal subspecies); Ichikawa
et al.
2000: 276; 2006: 180;
Storozhenko & Paik 2007
;
Storozhenko
et al.
2015
: 145
;
Cigliano
et al.
2018
(
Orthoptera
Species File Online);
Olivero & Robillard 2017
: 1
(mating behavior).
Xenogryllus marmoratus unipartitus
(
Karny, 1915
)
—
Gorochov & Belokobylskij 1992
: 10
(subspecies).
Synonym name:
Heterotrypus unipartitus
Karny, 1915
—
Chopard, 1968
: 350
.
Common names: Pine cricket (English), matsumushi (Japanese = pine insect, waiting insect); bao ta ling (Chinese = pagoda bell).
Type material.
Neotype
,
♂
[new designation],
Japan
: Honshu
, collection Finot, identified
Calyptotrypus
species nova by
A. Finot
(MNHN-EO-ENSIF1592).
Additional
material examined.
Japan
: Honshu
,
Mie
,
ix.1957
,
F. Ohmachi
,
1♂
,
1♀
, identified
Xenogryllus marmoratus
de Haan, Xma Robillard
morpho (MNHN). Japon,
1♀
, Exposition Universelle 1869, #1276-69,
identified
Dionymus marmoratus
by L. Chopard (MNHN-EO-ENSIF1593). Japon [Japan, no precise locality and date],
1♂
,
1♀
, identified
Dionymus marmoratus
by L. Chopard, Xma Robillard morpho;
1♂
,
2♀
(MNHN);
1♀
(NHMW);
1♂
, 20.679, identified
X. marmoratus unipartitus
by Karny (NHMW);
4♀
, identified
Calyptotrypus
species nova
, collection Finot (MNHN). Kobe, H. Fruhstorfer,
1♀
, 24.125 (NHMW). Kyoto, I. Yamashiro,
1♂
, Xma Robillard morpho MEB, MEB Ziegler;
1♀
;
1♀
, molecular sample T. Robillard 2004 (MNHN). Kioto [Kyoto], Y. Hirase,
1♂
, identified
Madasumma marmorata
(Haan)
and
Calyptotrypus marmoratus
(Haan)
by Hebard, 1924 (MNHN). Kanagawa Prefecture, Kanate, Ooi-Cho Ashigara-kami-gun [
35° 23' 00"N
,
139° 08' 00"E
], H. Sakai,
14.viii.2010
,
1♂
,
2♀
,
1 juvenile
, identified
Xenogryllus marmoratus
by A. Ichikawa (MNHN);
1♂
, enregistrement appel TR-male2 [call recording—MNHN-SO-2016-14364], identified
Xenogryllus marmoratus
by A. Ichikawa (MNHN-EO-ENSIF1704);
1♂
, enregistrement appel TR-male1 [call recording, MNHN-SO-2016-14365], identified
Xenogryllus marmoratus
by A. Ichikawa (MNHN-EO-ENSIF1598). Japon, environs de Tokyo, J. Harmand, 1906 (MNHN). Kinki Distr.[ict], Wakayama Pref.[ecture], Hashimoto city,
19.x.1986
, A. Ichikawa,
1♂
, identified
X. marmoratus
by A. Ichikawa (RBINS).
Tsushima
, H. Fruhstorfer, septoct [ix–x], 24.123 (NHMW).
China
: Guanxi
, Jin Xiu,
10.x.1981
,
E110°11’
N24°07’
,
1♀
(
14062635
) (SIPPE). Longzhou,
1995-viii18
/23, Xianwei Liu, Weinian Zhang, Xinbao Jin,
1♀
(
14062635
) (SIPPE).
Chongqing
, Beibei,
E106°23’
N29°48’
,
1.x.2000
, Zhou,
1♂
(14080765) (SIPPE).
Anhui
, Huangshan,
E118°19’
N29°43’
[
Shanghai
market],
viii.2012
, T. Robillard,
1♂
(TR40), enregistrement appel [call recording MNHN-SO-
2018-36
] (MNHN-EO-ENSIF1702);
1♂
(TR2), enregistrement appel [call recording MNHN-SO-
2018-52
], molecular sample
X12
(MNHN-EO-ENSIF1594).
Jiangsu
,
Zhenjiang,
32°12'0.00"N
119°27'0.00"E
[
Shanghai
market],
viii.2012
, T. Robillard,
1♂
(TR41), enregistrement appel [call recording, MNHN-SO-
2018-38
] (MNHN-EO- ENSIF1707);
1♂
(TR3), enregistrement appel [call recording MNHN-SO-2018-], molecular sample
X13
(MNHN- EO-ENSIF3562).
Shanghai
, Padang
Shanghai
,
E121°32'
N31°13'
[
Shanghai
market],
viii.2012
, T. Robillard,
1♂
, molecular sample
X11
, enregistrement appel [call recording, MNHN-SO-
2018-37
] (MNHN-EO-ENSIF1599);
1♂
(MNHN).
Shanghai
, Prov. Klange[?], Musée Meude, O. Piel,
30.viii.
[19]30,
1♀
, #1623,
1♂
,
6.ix.
[19]30 (MNHN).
Guangdong
, Shenzhen Distr., Xichong,
27.ix.2014
, coll. Zhang Tao,
4♂
,
2♀
(SNNU).
Henan
, Xinxian Distr.,
4– 8.ix.2014
, coll. Ma Libin,
2♀
(SNNU).
Hainan
, Wuzhishan Mt.,
12.viii.2010
, coll. Jiang Chaozhong,
1♂
(NWAFU).
Zhejiang
, Tianlongshan Mt.,
8.vii.2009
, coll. He Zhuqing,
1♂
(ECNU).
Shaanxi
, Ankang,
10.viii.2017
, coll. Ma Libin,
1♂
(SNNU).
Taiwan
,
Takao, Formosa, Sauter xi.[19]07,
type
of
Heterotrypus unipartitus
,
det. Karny, identified
Dionymus formosanus
n. sp.
Type
by T. Shiraki, (DEI, Eberswalde), identified
Xenogryllus marmoratus unipartitus
(Karny)
by A. V. Gorochov (DEI) [examined on photo].
Sri Lanka
:
Ceylon, Kandy, H. Rolle Berlin SW.11,
1♂
(MNHN).
South Korea
:
data from website “South Korea: Orthopteroids of Korea” (http://www.jasa.pe.kr/pulmuchi/korthoptera/Xenogryllus-marmoratus.htm).
Distribution.
Japan
,
China
(including
Taiwan
and Hainan),
South Korea
(http://www.jasa.pe.kr/pulmuchi/ korthoptera/Xenogryllus-marmoratus.htm),
India
(specimens observed by
Chopard (1969))
,
Sri Lanka
(
1 male
specimen).
Emmended diagnosis.
Species characterised by its small size (
Fig. 1
E–G), head dorsal coloration with a faint median dark band on vertex, and male genitalia with very long and thin pseudepiphallic lophi (
Fig. 8G
); in male FWs (
Fig. 5E
), mirror wider than long, less rounded than in
X. maichauensis
,
X. ululiu
and
X. transversus
.
Redescription.
Species of small to average size, the smallest for the genus (
Fig. 1
E–G). Coloration almost homogeneously light ochre. Median dark band on head dorsum faint on vertex (
Fig. 4E
), prolonged anteriorly by black fastigium, with three narrow lateral bands on each side. Fastigium straight, not widened anteriorly. Antennae light brown, first segment darker. Face relatively flat in lateral view (
Fig. 3J
). Pronotum dorsal disc light brown, with a median brown band variably marked prolonging head coloration; lateral margins not carinated, with a narrow yellow line; lateral lobes ochre, progressively lighter ventrally.
Male.
FWs light brown, translucent (
Fig. 5E
); dark coloration anterior to 1A sometimes extended on angle of 1A. Transverse part of file almost straight, with 435 stridulatory teeth (n=1) on transverse part of 1A. Harp as wide as long. Fused part of cells c1 and b1 as long as individualised part of c1. Cell d2 wider than in other species. Mirror wider than long, little rounded, its inner limit forming a wide angle. Apical field short, including five cell alignments.
FIGURE 14.
Calling song of
Xenogryllus marmoratus
: (A) oscillogram of 5 echemes; (B) detailed oscillogram (upper panel) and sonogram (lower panel) of 1 echeme; (C) detailed oscillogram of one syllable; (D) frequency spectrum.
Male genitalia
(
Fig. 7
H–I): Pseudepiphallic sclerite longer than rami; pseudepiphallic lophi very long (
Fig. 8G
), narrow and sclerotized, slightly convergent, ended by an apical lamella curved dorsally and forming a small notch. Pseudepiphallic sclerite with wide lateral membranous lobes. Basal reinforcement of pseudepiphallic sclerite strong. Rami short with convergent hook-like apex. Pseudepiphallic parameres with a ventral transverse crest absent in other species; inner apex of of pseudepiphallic parameres sharp, slightly sclerotized apically. Ectophallic apodemes strong, not lamellate. Ectophallic lateral arms strong, fused to lateral arms of ectophallic fold, and with lateral arms of endophallic sclerite, forming together a wide gutter. Ectophallic fold entirely sclerotized ventrally. Endophallic sclerite small, apodeme made of a thin anterior dorsal crest and narrow lateral lamellas.
Female.
FWs dorsal field with 8–9 strong longitudinal veins (m = 8, n = 10) and faint transverse ones. Subgenital plate not indented apically (
Fig. 9B
). Ovipositor longer than FIII, its apex rounded and smooth.
Female genitalia:
Copulatory papilla (
Fig. 11
C–D) thin and conical, widened basally; sclerotization strong basally and apically, its membrane plicate.
Life history traits.
X. marmoratus
lives on low vegetation in bushes and grass in secondary habitats and along roads. The species has a nocturnal activity; males call at night from the vegetation. See
Olivero & Robillard (2017)
for description of peculiar mating behaviours.
Calling song
(
Figs 12B
,
14
). At 26.5°C, the calling song of
X. marmoratus
is made of relatively long syllables, each corresponding to one FW closure. Syllables form a two-part echeme: 1–3 singleton syllables (m = 2, n = 10), followed by a chirp made of three syllables, sometimes duplicated. The amplitude profile of the call shows that the singletons are usually less loud than the chirp. Total echeme duration is 711 ± 33 ms, for an echeme period of 6776 ± 368 ms (echeme duty cycle = 10.5%). The syllables have the following characteristics: 1
st
syllable duration = 51.8 ± 1.5 ms (period = 318.3 ± 35.6 ms); 2
nd
syllable duration = 50.3 ± 1 ms; chirp duration = 183.8 ± 1 ms; chirp syllable duration = 49.5 ± 1.9 ms (period = 68 ± 0.8 ms). The frequency spectrum shows a pure tone fundamental peak corresponding to the dominant frequency at
ca.
5.9 ± 0.2 kHz, with a clear harmonic series, the third harmonic (
ca.
18 kHz) being almost as powerful as the fundamental peak.
Measurements.
See
Table 6
.
Taxonomic discussion.
The types of De Haan, supposed to be in Leiden, were not found in RMNH in 2006 (TR, pers. obs.), with no record mentioning them as loaned (Rob De Vries, curator of the Orthopteran collection in RMNH, pers. com. 2006; confirmation by Luc Willemse, current curator of the Orthopteran collection in RMNH, pers. com. 2018). Specimens possibly belonging to the original type series were also searched for in other European museums, but could not be found. We thus designate a
neotype
from
Japan
that will serve as a reference for future systematic works.
TABLE 6.
Measurements of
Xenogryllus marmoratus
BL |
PronL |
PronW |
FWL |
FWW |
HWT |
FIIIL |
FIIIW |
Neotype male |
19.4 |
2.9 |
4.1 |
15.5 |
6.7 |
3.5 |
15.3 |
2.8 |
Males (n=4) |
18.6–22 |
2.5–3 |
3.9–4.8 |
15.5–17 6.4–6.7 |
3.1–4.3 |
14.2–15.8 |
2.6–3.4 |
(Male mean) |
(20.4) |
(2.9) |
(4.3) |
(16.5) |
(6.5) |
(3.4) |
(15.3) |
(3) |
Females (n=5) |
18.8–20 |
2.1–3.1 |
3.1–4.3 |
11.5–15 2.8–4 |
1.2–6.3 |
11.8–16.3 |
2.2–3.2 |
(Female mean) |
(19.1) |
(2.7) |
(3.7) |
(13.4) |
(3.3) |
(3.4) |
(14.4) |
(2.7) |
TIIIL |
TIIIs |
TaIIIs |
OL |
Ias |
Ibs |
Oas |
Obs |
Ids |
Ods |
Ols |
Neotype male |
11.1 |
9 |
11 |
14 |
16 |
0 |
4 |
5 |
- |
Males (n=4) |
11.1–15.8 |
5–12 |
10–12 |
11–20 |
11–16 |
0–2 |
3–7 |
5–7 |
- |
(Male mean) |
(14.1) |
(9) |
(11) |
(16) |
(5) |
(1) |
(5) |
(5) |
- |
Females (n=5) |
12.1–14.9 |
4–10 |
7–12 |
13–18 |
9–13 |
0–2 |
2–5 |
5–10 |
14.1–19 |
(Female mean) |
(13.3) |
(8) |
(9) |
(16) |
(11) |
(1) |
(3) |
(7) |
(16.6) |
Gorochov (1992)
distinguished two subspecies,
X. m.
marmoratus
(
Haan, 1844
)
and
X. m.
unipartitus
(
Karny, 1915
)
. In his study, he considered
Gryllus
(
Phalangopsis
)
marmoratus
Haan, 1844
(
type
from
Japan
), as the nominal subspecies
Xenogryllus marmoratus marmoratus
, while he established the subspecies
X. m.
unipartitus
based on the species
Heterotrypus unipartitus
Karny, 1915
, which was described from one female from
Taiwan
(Takao, examined on photo). The
type
of
H. unipartitus
clearly corresponds to a female of
Xenogryllus
,
however, given the continuous variation observed across the wide distribution of
X. marmoratus
, and the limited information available based on female morphology, there is no reason to maintain these two subspecies. We thus synonymise the subspecies under
X. marmoratus
,
and
H. unipartitus
becomes a junior synonym of
X. marmoratus
,
as previously proposed by
Chopard (1968)
.