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Accepted by D. Rider: 4 May 2016; published: 5 Jul. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4136 (1): 141
–
154
http://www.mapress.com/j/zt/
Article
141
http://doi.org/10.11646/zootaxa.4136.1.6
http://zoobank.org/urn:lsid:zoobank.org:pub:BF1CF1FC-F21C-41E1-9227-D164B830B7CE
The identity of the Brachyplatys species recently introduced to Panama,
with a review of bionomics (Hemiptera: Heteroptera: Plataspidae)
DÁVID RÉDEI
1,2
1
Institute of Entomology, College of Life Sciences, Nankai University, Weijin Road 94, 300071 Tianjin, China.
E-mail: david.redei@gmail.com
2
Department of Zoology, Hungarian Natural History Museum, H-1088 Budapest, Baross u. 13, Hungary
Abstract
A recent report of a population of Brachyplatys vahlii (Fabricius, 1787) (Hemiptera: Heteroptera: Plataspidae) introduced
to Panama is considered as misidentification, the species in concern is recognized as B. subaeneus (Westwood, 1837). Syn-
types of B. subaeneus and diagnostic characters of the species are illustrated, published information on its distribution,
bionomics and economic importance is reviewed. Syntypes of B. vahlii are illustrated, taxonomic problems in connection
with the species are highlighted.
Key words: Plataspidae, Brachyplatys subaeneus, B. vahlii, misidentification, invasive species, distribution, bionomics,
economic importance
Introduction
Plataspidae is an endemic Old World family of true bugs (Hemiptera: Heteroptera). An Indomalayan species,
Megacopta cribraria (Fabricius, 1798) has recently been introduced to the USA (Eger et al. 2010) where it started
a rapid expansion and soon became a significant pest of soybean and other legumes (Ruberson et al. 2013).
Another plataspid was reported recently to be introduced to Panama (Aiello et al. 2016). The authors identified
the species in concern as Brachyplatys vahlii (Fabricius, 1787), a species widely distributed in the Indomalaya.
Photos of a male (Aiello et al. 2016: 9, figs. 2B–C), however, indicate that the attribution of this species to B. vahlii
is unsound. The purpose of the present note is to clarify the identity of the invasive species, and provide a short
review of its diagnostic characters, distribution, and bionomics.
Material and methods
External structures were examined using a stereoscopic microscope (Zeiss Discovery V8). Drawings were made
with the aid of a camera lucida. Male genitalia were dissected after careful heating in hypertonic KOH solution,
stained with Chlorazole Black E if necessary, and examined using stereoscopic microscope (Zeiss Discovery V8).
Label data are cited verbatim, lines on a single label are divided by a backslash (\), comments on label data are
provided in square brackets. Printed text (pr) is not indicated except if the preceding text is handwritten (hw). Type
depositories and localities verified by us are marked with exclamatory point (!). Morphological terminology
follows Tsai et al. (2011).
Abbreviations of depositories:
NKUM Institute of Entomology, Nankai University, Tianjin, China;
OXUM Oxford University Museum, Hope Entomological Collections, Oxford, United Kingdom;
ZMUC Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark.
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Taxonomy
Brachyplatys Boisduval, 1835
Platycephala Laporte, 1833: 73 (as subgenus of Coptosoma Laporte, 1833). Type species by subsequent designation (Kuhlgatz
1901: 205): Platycephala metallica Laporte, 1833. Junior homonym of Platycephala Fallén, 1820 (Diptera). Synonymized
by White (1839: 539).
Brachyplatys Boisduval, 1835: 627. Type species by monotypy: Brachyplatys vanikorensis Boisduval, 1835.
Plataspis Westwood, 1837: 16. Unnecessary new name for Platycephala Laporte, 1833.
Brachyplatys subaeneus (Westwood, 1837)
(Figs. 1–19)
Plataspis subaenea Westwood, 1837: 17. Syntype(s): Bengalia [= India or Bangladesh]; OXUM!
Brachyplatys vahlii (non Fabricius, 1787): Aiello et al. (2016: 8). Misidentification.
A complete list of synonyms was provided by Davidová-Vilímová (2006).
Selected bibliography
Platycephala subaenea: Westwood (1837: 5) (listed, distribution).
Brachyplatys subaenea: Dallas (1851: 70) (records), Walker (1867: 100) (records), Stål (1876: 8) (in key, catalogue,
distribution), Atkinson (1887: 28) (English translation of original description, distribution), Kirby (1891: 79) (listed),
Lethierry & Severin (1893: 5) (catalogue, distribution), Montandon (1894: 124) (records), Montandon (1897: 444)
(records), Distant (1899: 213, 227) (synonymy), Breddin (1900: 275) (intraspecific variability, records), Distant (1900:
820) (generic placement), Breddin (1901: 7, 139) (record, distribution), Distant (1902: 11) (diagnostic characters, records,
distribution), Noualhier & Martin (1904: 168) (record), Breddin (1905: 208) (record, distribution), Breddin (1909: 250)
(record), Kirkaldy (1909: 325, 380) (catalogue, distribution, host plant), Maxwell-Lefroy (1909a: 302) (distribution),
Maxwell-Lefroy (1909b: 671) (habitus, host plants, economic importance), Kershaw & Kirkaldy (1910: 69) (descriptions
and figures of egg, egg batch, all larval instars and adult, distribution, host plant, life history), Kirkaldy (1910: 111)
(record), Bainbrigge Fletcher (1920: 249) (record, host plant, economic importance), Esaki (1926: 143) (records, host
plants), Dammerman (1929: 219) (diagnostic characters, host plants, economic importance), Hoffmann (1931: 138) (host
plants), Miller (1931: 7) (records, distribution, descriptions and figures of egg batch, all larval instars and adult,
oviposition, development, host plants, economic importance), Hoffmann (1932: 1014) (distribution, host plants, egg batch,
aggregation), Miller (1934: 504) (figure of egg burster), Yang (1934: 145) (in key), 151) (redescription, figures,
distribution, diagnostic characters), Hoffmann (1935: 10, 156, 205) (catalogue, distribution), Wu (1935: 255) (catalogue,
distribution), Hafiz & Ribeiro (1941: 77) (records, distribution), Manna (1951: 5, 32) (karyology, figures of
chromosomes), Chandra (1953: 97) (records), Banerjee (1958: 11, 17) (record, host plant, karyotype), Vayssière (1958:
799) (diagnosis, habitus, host plants, bionomics), Hsiao & Jen (1977: 77) (in key, redescription, photo, figures,
distribution), Ueshima (1979: 75) (karyotype), Datta et al. (1985: 16) (diagnosis, figure, distribution), Chen (1985: 32)
(redescription and figures of egg batch, first, third and fifth larval instars and adult, distribution, host plants, phenology,
economic importance), Zhang & Sie (1986: 97) (records), Ren (1987: 105) (in key, figures, distribution), Ahmad &
Moizuddin (1992: 6) (in key, redescription, habitus, figures, record, distribution), Johnson (1996: 283) (parasitoid), Easton
& Pun (1997: 579) (record, host plants), Li et al. (1997: 20, 81, 118) (distribution, host plant, economic importance), Lin &
Zhang (1998: 5) (catalogue, distribution, host plants), Hayashi (2002: 146) (distribution), Wu & Xu (2002: 29) (listed, host
plant), Tsai & Yang (2005: 103) (photo), Davidová-Vilímová (2006: 152) (catalogue, distribution), Liu & Zheng (2006:
920) (listed, distribution), Pol et al. (2010: 50) (diagnosis, photo, host plant, economic importance), Yamashita & Kikuhara
(2012: 439) (redescription, photos, records, distribution), Yiu & Yip (2012: 184) (redescription, photos), Bao et al. (2013:
723) (listed), Zheng & Lin (2013: 114) (redescription, photo), Nagashima & Tanaka (2014: 51) (diagnosis, photo,
comparison with B. vahlii).
Brachyplatys subaeneous [incorrect subsequent spelling]: Mitsuhashi (1915: 484) (listed, distribution).
Brachyplatus [incorrect subsequent spelling] subaeneus: Wall (1928: 231) (parasitoid), Wall (1931: 633) (records, distribution,
descriptions and figures of egg batch, all larval instars and adult, oviposition, development, moulting, host plants,
economic importance).
Brachyplatys vahlii (misidentification): Aiello et al. (2016: 8) (photos of adults and egg batch, records, host plants).
Type material examined. Syntype #1: ♀, “Type” [circle with red margin], “Bengal” [hw], “Brachyplatys \
subaeneus \ Westw.” [W.L. Distant’s hw], “TYPE \ == \
WESTW
. (
HOPE
) \ C.Hemipt.1837 \ Part I, page [pr] 17 [hw]
\ Distant, P.Z.S., \ 1900, p.807-825” [with hw red frame], “TYPE [pr] HEM [hw] : [pr] No 7 1/2 [hw] \ PLATASPIS
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[hw] \ SUBAENEA [hw] \ WESTWOOD [hw] \ HOPE DEPT. OXFORD”; pinned, intact (OXUM) (Figs. 1–3).
Syntype #2: ♀, “Bengal” [hw], “TYPE [pr] HEM [hw] : [pr] No 7 2/2 [hw] \ PLATASPIS [hw] \ SUBAENEA [hw]
\ WESTWOOD [hw] \ HOPE DEPT. OXFORD”; pinned, wings in abduction, right hind tarsus lacking (OXUM)
(Figs. 4–6).
Diagnosis. A relatively small species of Brachyplatys (body length: 4.5–5.8 mm) recognized among other
Indomalayan congeners based on the following combination of characters: disk of pronotum and base of scutellum
devoid of pairs of small, rounded, pale patches (Figs. 1, 4); yellow fasciae on lateral margins of abdominal ventrites
narrow, those on ventrites VI–VII restricted to not more than one tenth of total width of the respective ventrites
(Figs. 2, 5, 7–8). The external male (Figs. 9–17) and female genitalia (Figs. 18–19) are diagnostic and they are
FIGURES 1–6. Two syntypes of Plataspis subaenea Westwood, 1837 and their labels. Fig. 1, syntype #1, dorsal view; Fig.
2, same, ventral view; Fig. 3, same, labels; Fig. 4, syntype #2, dorsal view; Fig. 5, same, ventral view; Fig. 6, same, labels.
Scales in mm, Figs. 3 and 6 not to scale (© OXUM).
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FIGURES 7–14. Brachyplatys subaeneus (Westwood, 1837). Fig. 7, left half of abdominal venter of male; Fig. 8, same of
female; Fig. 9, genital capsule, posterior view (left paramere removed); Fig. 10, same, posteroventral view (pilosity omitted);
Fig. 11, same, lateral view, with outline of phallus; Fig. 12–14, left paramere, three different aspects. Abbreviations: at = anal
tube; cs = cuplike sclerite; dif = dorsal infolding of genital capsule; lp = left paramere; rp = right paramere; vif = ventral
infolding of genital capsule. Scales in mm.
described in detail below. Photos of living specimens were provided by Pol et al. (2010), Yamashita & Kikuhara
(2012), Yiu & Yip (2012), and Zheng & Lin (2013).
Description of external male genitalia. Genital capsule (Figs. 9–11) black, infolding of dorsal rim (Fig. 9:
dif) broadly yellow, with a broad, deep central impression around posterior aperture associated with a pair of large,
obtuse, denticle-like projections laterally (Fig. 10: arrow); with dense, very fine and short pubescence on dorsal
infolding, with longer and stronger setae on the obtuse projections and on ventral infolding; cuplike sclerite (Figs.
10–11: cs) protruding as a transverse, shallowly emarginate plate bordering posterior aperture ventrally, completely
fused with infolding of ventral rim (Fig. 10: vif), their boundary indistinct. Parameres (Fig. 10: rp, Fig. 11: lp, Figs.
12–14) with a relatively short stem and a broadly oval crown produced into a long, narrow, arched, apically obtuse
projection; oval body of crown provided with long, densely distributed setae; body of crown directed obliquely, its
projection dorsally and slightly laterally in rest (Fig. 9). Phallus (Figs. 15–17) elongate; articulatory apparatus
small; phallotheca voluminous, elongate, largely membranous, its wall provided with a pair of sclerites laterally;
hinge (Fig. 16: hi) enlarged, occupying about half of total length of phallotheca; conjunctiva (Figs. 15–16: con)
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elongate, tubular, provided with two pairs of processes: with a pair of lateral processes close to base that are
scerotized and pigmented, relatively short, oval, apically tapering and pointed (Fig. 15: cp), of uncertain homology,
and with a pair of lateral processes at apex that are membranous, elongate, tubular, and provided with a more
narrow finger-like projection subapically (Fig. 15: cp-II?), probably homologous with cp-II in other Pentatomoidea
(cf. Tsai et al. 2011); aedeagus (Fig. 16: aed) elongate, its proximal third membranous and relatively broad, its
distal two-thirds sclerotized, pigmented, narrow, tubular, gradually tapering to apex, abruptly curved
posteroventrad around its distal third to fourth, provided with a large, obtuse, strongly pigmented denticle at base of
sclerotized portion ventrally (Figs. 16–17: arrow); endophallic reservoir (Fig. 15: res) small, more or less globular;
endophallic tube (Fig. 15: end) simple, sclerotized.
Description of external female genitalia. Terminalia as in Fig. 18; surface of valvifers VIII shallowly
excavate. Gynatrium elongate, membranous, dorsal surface with processes of unknown homology as shown in Fig.
19, no distinct ring sclerites could not be traced in the examined specimens; proximal extremity of spermathecal
duct relatively broad (for reception of distal portion of aedeagus), then strongly narrowed and continued in a long
and thin portion (Fig. 19: sthd), intermediate part short, relatively narrow, with a smaller proximal and a larger
distal flange, apical receptacle large, globular.
Intraspecific variability. The marginal fasciae of the abdominal venter are sexually dimorphic: in females
each of ventrites III–VII are provided with distinct subtriangular patches (Fig. 8), whilst in males the markings of
ventrites V–VII are strongly reduced, present merely as narrow yellow markings along the extreme lateral margins
of the respective ventrites (Fig. 7). The yellow markings on the dorsum of the head are somewhat variable in both
sexes, it is frequently reduced, with the transverse marginal and interocular fasciae broken up to separate patches
(cf. Breddin 1900) (Figs. 1, 4). The aedeagus is also somewhat variable, most examined specimens were as in Fig.
17, but a specimen as in Fig. 16 have been examined as well; the latter one was part of a long series of B. subaeneus
specimens and apparently there is no reason to recognize it as different from B. subaeneus.
Immatures. Egg batch and all larval instars were illustrated by Kershaw & Kirkaldy (1910), Miller (1931) and
Wall (1931); egg batch and selected larval instars by Chen (1985).
Bionomics and economic importance. Brachyplatys subaeneus shows a clear preference to species of the
family Fabaceae. The following leguminous crops were recorded as host plants: India: the lablab, Dolichos lablab
L. (Maxwell-Lefroy 1909b); China and Indochina: various cultivars of the common bean, Phaseolus vulgaris L.,
lima bean, Ph. lunatus L., yardlong bean, Vigna unguiculata Walp. var. sesquipedalis (L.) H.Ohashi (= V.
sesquipedalis (L.) Fruwirth), mung bean, V. r a di at a (L.) R.Wilczek, and soybean, Glycine max (L.) Merr.
(= Glycine hispida Maxim.) (Hoffmann 1931, 1932; Wall 1931; Wu & Xu 2002; Pol et al. 2010); Malay Peninsula:
soybean, jack bean, Canavalia ensiformis (L.) DC., quickstick, Gliricidia sepium (Jacq.) Kunth (= G. maculata
(Kunth) Kunth), and catjang, V. unguiculata (L.) Walp. subsp. cylindrica (L.) Verdc. (= Vigna catjang Walp.)
(Miller 1931, Hoffmann 1932); Sumatra: tropical kudzu, Pueraria phaseoloides Benth. (= P. javanica Benth.) and
comon bean (Vayssière 1958). In India it was reported from the invasive needle bush, Acacia (now Vachellia)
farnesiana (L.) Wight et Arn. (Banerjee 1958); in Macau it is common on the kudzu species Pueraria lobata
(Willd.) Ohwi (= P. thunbergiana Benth.) (Kirkaldy 1909, Kershaw & Kirkaldy 1910, Yiu & Yip 2012) and
probably also on other Pueraria species. The host plant records from the pigeon pea, Cajanus cajan (L.) Millsp.
(= C. indicus Spreng.) and Indigofera suffruticosa Mill. (= I. anil L.) by Hoffmann (1932) are based on unknown
sources, and perhaps they were taken from literature concerning other species of Brachyplatys.
The following host plant records pertaining to members of other plant families were published: Cannabaceae:
the hemp, Cannabis sativa L. in India (Bainbrigge Fletcher 1920, Hoffmann 1932); Solanaceae: potato, Solanum
tuberosum L. in the Malay Archipelago (Dammerman 1929, Hoffmann 1932); Convolvulaceae: sweet potato,
Ipomoea batatas (L.) Lam. in Indochina and the Malay Archipelago (Dammerman 1929, Hoffmann 1932, Pol et al.
2010); Asteraceae: bitter wine, Mikania micrantha Kunth in Macau (Yiu & Yip 2012); “jute plants” (apparently
Corchorus sp., Sparrmanniaceae) in India (Maxwell-Lefroy 1909b, Hoffmann 1932); Poaceae: sugarcane,
Saccharum sp. (probably S. officinarum L.), rice, Oryza sativa L. (Pol et al. 2010), and unspecified “grasses”
(Ahmad & Moizuddin 1992).
The introduced population in Panama was reported from pigeon pea and an endemic New World fabacean
species, Leptolobium panamense (Benth.) Sch.Rodr. & A.M.G.Azevedo, and also from two species belonging to
other families, the peach palm, Bactris gasipaes Kunth (Arecaceae) and the umbrella tree or octopus tree,
Schefflera actinophylla (Endl.) Harms (Araliaceae) (Aiello et al. 2016).
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FIGURES 15–18. Brachyplatys subaeneus (Westwood, 1837). Fig. 15, phallus, dorsal view; Fig. 16, same, lateral view; Fig.
17, aedeagus of another individual from the same population, lateral view; Fig. 18, female terminalia, posteroventral view; Fig.
19, gynatrium and spermatheca, dorsal view after removal of pregenital tergum (basal portion of spermathecal duct lifted in the
direction shown by arrow to allow better visualisation of structures). Abbreviations: aed = aedaeagus; con = conjunctiva; cp =
conjunctival process; cp-II? = second pair of conjunctival process?; end = endophallic duct; hi = hinge; res = endophallic
reservoir; lt
9
= laterotergite IX; phth = phallotheca; sthd = spermathecal duct; vf
8
= valvifer VIII. Scales in mm.
The life history of the species was studied in southern China (Wall 1931) and the Malay Peninsula (Miller
1931) in detail; the data provided by Chen (1985) are apparently taken from the work by Wall (1931). Copulation
proceeds in an end-to-end position (cf. Pol et al. 2010, Yiu & Yip 2012) and usually lasts for about 2–3 hours (Wall
1931). Eggs are rarely encountered in the field and they were reported to be deposited on the leaves or stems of the
host plants or other plants (Kershaw & Kirkaldy 1910, Miller 1931) or probably in crevices or other inaccessible
narrow spaces around the host plants (Wall 1931). The egg batch is composed of 20–40 eggs, arranged in two
parallel rows, each egg lying on its side, the opercula pointing obliquely outwards (Miller 1931: 8, fig. 1; Wall
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1931: plate 29, fig. 1; Yiu & Yip 2012: 184, photos). The oviposition process and the hatching was described by
Wall (1931). Females deposit approximately 300–400 eggs over a period of two months or longer. Egg hatch after
an incubation period of 5–6 days (Kershaw & Kirkaldy 1910, Miller 1931, Wall 1931). First instar larvae aggregate
around the empty egg shells, later instars disperse. Larvae feed on young shoots and stems of the host plants and
almost never on leaves (Wall 1931); they are, just like the adults, capable of rapid movement. Larval development
takes about 35–44 days; the longevity of the adults is about two months in average but it can be as long as 179 days
(Miller 1931, Wall 1931). Individuals frequently aggregate on the stem or under leaves of the host plants.
In southern China later instar larvae and adults overwinter. They leave their refugia and the larvae develop to
adults in April. The species is trivoltine, with mass appearance of young adults in mid June, late July, and mid
September, respectively. Hibernation begins from October (Wall 1931).
Brachyplatys subaeneus is harmful to leguminous plants both in southern China and the Malay Peninsula, but
its significance is minor except for heavy infestations (Miller 1931, Wall 1931). Its feeding results in deformation
of the pods or stunting of the host plants. Foreign varieties are more infested than local cultivars (Wall 1931).
Damage inflicted on jute plants (Maxwell-Lefroy 1909b) and hemp (Bainbrigge Fletcher 1920) were reported, but
these cases are certainly exceptional. Its significance on crops belonging to other plant families is probably
insignificant.
Paratelenomus saccharalis (Dodd, 1914) (Hymenoptera: Platygastridae), misidentified as Dissolcus (now
Paratelenomus) tetartus Crawford, 1911 (Scelionidae) by Wall (1931) (cf. Johnson 1996), parasitizing eggs of
Megacopta cribraria (Fabricius, 1798) in Guangdong, southern China, was reported to develop equally well in
eggs of B. subaeneus under laboratory conditions, but no field observations are available on its significance (Wall
1928, 1931).
Distribution. The species is widely distributed and abundant all over the Indomalaya, apparently restricted to
the tropical and subtropical moist broadleaf forest zone. Several specimens have been seen in various collections in
Europe, most of them correctly identified, therefore the vast majority of the previous literature records are probably
correct, with at least the following exceptions:
1. The redescription (p. 54) and illustration (plate IV, fig. 8) of B. subaeneus provided by Vollenhoven (1863)
leave no doubt that the species in concern is B. vahlii.
2. The photo of B. subaeneus provided by Hosokawa et al. (2006: 1842, fig. 1C) is based on misidentification and
pertains to B. vahlii.
3. The colour painting of B. subaeneus provided by Cai & Li (2015: 154) is based on misidentification and shows
a female of Ponsilasia montana (Distant, 1901).
4. The record from Ternate Island of the Malay Archipelago (Yang 1934) is based on an unknown source and
therefore it is considered as in need of confirmation.
5. Li et al. (1997) listed the species from several provinces of China, but the records of Jiangsu, Anhui, Zhejiang,
Jiangxi, Hubei, Hunan and Sichuan are based on unknown sources; since no other Chinese authors recorded
the species from any of the above provinces, these are considered as doubtful.
The examined specimens are not listed in detail except of the following specimens which represent the first
records for the Tibet Autonomous Region: Mêdog, No. 3 bridge—Baibung [= Bepung], 780–1100 m, 11.viii.2003,
leg. H.J. Xue (4 ♂♂ 3 ♀♀ NKUM). The specimens used for illustrating the diagnostic characters of the species
(Figs. 7–18) are all part of a long series from southwestern China, Yunnan, Dehong Dai and Jingpo Autonomous
Prefecture, Yingjiang County, Tongbiguan Township, on the way to Jinshanliao, 318–1160 m, 15.v.2009, leg. B.
Cai (NKUM). The geographic distribution of the species might be summarized as follows.
IND O M A LAYA. INDIA. Uttarakhand: Dehra Dun [= Dehradun] (Chandra 1953); Bihar: Pusa (Maxwell-
Lefroy 1909a, Chandra 1953), Katihar (Hafiz & Ribeiro 1942); West Bengal: Calcutta [= Kolkata], Mungphu
[= Mungpoo] (Distant 1902, Maxwell-Lefroy 1909a), Maldah [= Malda Distr.] (Hafiz & Ribeiro 1942); Assam:
Sadiya (Chandra 1953); Odisha: Balugaon (Hafiz & Ribeiro 1942); Andhra Pradesh: Mahanandi (Hafiz &
Ribeiro 1942); Karnataka: Mysore [= Mysuru], Bangalore [= Bengaluru] (Distant 1902); Andaman Islands
(Distant 1902, Hafiz & Ribeiro 1942).—SRI LANKA (Kirby 1891, Distant 1902, Breddin 1909, Hafiz & Ribeiro
1942).—BANGLADESH. Rangpur (Bainbrigge Fletcher 1920), Chittagong Hill Tracts: Maini Mukh (Hafiz &
Ribeiro 1942), unspecified locality (Ahmad & Moizuddin 1992).—BURMA (MYANMAR). Bhamo, Teinzo,
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Metanja (Montandon 1894, Distant 1902); Tanintharyi Region: Meetan, Kawkareet (Montandon 1894, Distant
1902).—THAILAND. Siam [= Thailand] (Distant 1902).—CHINA. Guizhou (Liu & Zheng 2006); Yunnan:
Jinghong (Hsiao & Jen 1977); Guangxi: Longzhou, Youyi Guan [= Friendship Pass] (Hsiao & Jen 1977); Hainan
(Hsiao & Jen 1977); Guangdong: Canton [= Guangzhou] (Wall 1931, Hsiao & Jen 1977), Shenzhen (Bao et al.
2013); Fujian: Fuzhou, Zhangping (Zhang & Sie 1986, Wu & Xu 2002); Hong Kong (Dallas 1851); Macau
(Montandon 1897, Kirkaldy 1910, Easton & Pun 1997); Tibet (new record).—JAPAN. Ryukyu Archipelago:
Amami Islands: Okinoerabu-jima; Okinawa Islands: Okinawa Is.; Kerama Islands: Geruma-Jima; Daito
Islands: Kitadaito-jima; Miyako Islands; Yaeyama Islands: Ishigaki-jima, Iriomote-jima, Hateruma-jima,
Yonaguni-jima; Senkaku Islands (Hayashi 2002, Yamashita & Kikuhara 2012).—TAIWAN (Esaki 1926, Zheng
& Lin 2013).—VIETNAM. Tonkin [= northern Vietnam]; Tuyen Quang Prov.: Montagnes du H[au]t Song-Chaï
[= mounts of the upper Chay River (= Song Chay)] (Montandon 1897).—CAMBODIA (Noualhier & Martin
1904, Montandon 1897, Pol et al. 2010).—PHILIPPINES. Balabac (Breddin 1900).—MALAYSIA. Malay
Peninsula: Neg[e]ri Sembilan, Serdang, Pahang, Sepatak (Miller 1931); Sabah: Mons Kina Balu [= Mt. Kinabalu]
(Breddin 1900); Banggi Is.: Banguey [= Banggi Is.] (Breddin 1900, 1905).—SINGAPORE (Walker 1867,
Montandon 1897).—INDONESIA. Java (Walker 1867, Breddin 1900); Sumatra: Soekaranda [= Sukaranda]
(Breddin 1900); Sulawesi: Mapane (Breddin 1901); Buru: Bouru [= Buru] (Walker 1867).—NEOTROPICS.
PAN A M A . Several localities of the Canal area (introduced) (Aiello et al. 2016).
Brachyplatys vahlii (Fabricius, 1787)
(Figs. 20–25)
Cimex vahlii Fabricius, 1787: 283. Syntypes: India orientalis [= East Indies]; ZMUC!
Brachyplatys subaeneus (non Westwood, 1837): authors. Misidentification.
A complete list of synonyms was provided by Davidová-Vilímová (2006).
Brachyplatys subaeneus (misidentification): Vollenhoven 1863: 54 (redescription, figure, record), Tomokuni 1993: 213 (photos
of adult and immatures, record, distribution), Hosokawa et al. 2006: 1842 (photo, gut symbionts), Kikuchi et al. 2008: 45,
49, 51 (gut symbionts).
Type material examined. Syntype #1: ♀, without labels; pinned, segments IV of both antennae and left hind leg
lacking (ZMUC, no. 00102796) (Figs. 20–21). Syntype #2: ♀, “Vahlii” [hw]; pinned, head, right mid and hind legs
and terminalia lacking, right thoracic pleuron damaged by museum beetles (ZMUC, no. 00102797) (Figs. 22–24).
The following labels are pinned in the drawer next to the syntypes: “63 \ Brachyplatys \ vahlii F” [hw, with double
black pr frame]; “Tetyra Vahlii F. \ Syst. Rhyng. p. 142.69” [typewritten] (Fig. 25).
Remarks. Photographs of two syntypes deposited in ZMUC (cf. Zimsen 1964) were examined in connection
with the present study (Figs. 20–25).
Other than of B. vahlii, only one valid species of Brachyplatys with similar extensive yellow fasciae on the
abdominal venter is recognized in Indomalaya, namely B. picturifrons Walker, 1867 (Sulawesi). Based on
topotypic specimens examined it certainly differs from B. vahlii at species level. Brachyplatys vahlii, however, is
apparently a complex of morphologically similar species readily recognizable based on the genital capsules of the
males. Based on the reexamination of the type material of several species currently recognized as junior synonyms
of B. vahlii (cf. Kirkaldy 1909, Davidová-Vilímová 2006) some of these species represent valid species. The
complex is therefore in need of revision. The situation is complicated further by the fact that the type material of B.
vahlii consists of two females from an unspecified locality (“India orientalis” [= East Indies], incorporating usually
the whole Indomalaya or at least the Malay Archipelago and Indochina). This problem is not examined further in
the present study, the main goal of treating the species here is to call the attention on its differences from B.
subaeneus.
Distribution. Brachyplatys vahlii is widely distributed throughout the Indomalaya, and it was also recorded
from Wallacea and New Guinea. Several specimens were seen from the Indian Subcontinent, Indochina, and the
Malay Archipelago. Because of the broad recognition of this species, some of the old records, however, might need
correction. The following remarks are made:
1. The redescription (p. 54) and illustration (plate IV, fig. 8) of B. subaeneus provided by Vollenhoven (1863)
leave no doubt that the species in concern is B. vahlii.
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FIGURES 20–25. Two syntypes of Cimex vahlii Fabricius, 1787 and their labels. Fig. 20, syntype #1, dorsal view; Fig. 21,
same, ventral view; Fig. 22, syntype #2, dorsal view; Fig. 23, same, ventral view; Fig. 24, same, label; Fig. 25, labels pinned in
the bottom of the drawer next to the specimens. Scales in mm, Figs. 24–25 not to scale (© ZMUC).
2. The photos of B. subaeneus provided by Tomokuni (1993: plate 83 figs. 252a–b) are also based on
misidentification and pertain to B. vahlii. Japanese records of B. subaeneus after Tomokuni (1993) are
potentially based on this work and therefore might represent misidentifications too; at least the record of
Hosokawa et al. (2006: 1842, fig. 1C) pertains to B. vahlii. Yamashita & Kikuhara (2012) and Nagashima &
Tanaka (2014) identified both species correctly.
Discussion
An introduced population of Brachyplatys vahlii was reported from Panama by Aiello et al. (2016). The photos of
a male (Aiello et al. 2016: 9, figs. 2B–C), however, conflict with this identification: in B. vahlii the disk of
pronotum is invariably provided with a pair, the basal margin of scutellum with two pairs of small, rounded, yellow
to orange patches (cf. Figs. 20, 22), and the lateral margins of the abdominal ventrites are invariably decorated with
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yellow transverse fasciae occupying at least the lateral thirds of at least ventrites II–IV (cf. Figs. 21–23). The
photos of a Panaman specimen readily allow an identification of the species as B. subaeneus.
The identification of the Panaman specimens was based on comparisons of mitochondrial COI and 16S rDNA
sequences with sequences obtained from GenBank. Aiello et al. (2016) claimed that the 16S rDNA sequence of
their species showed 99% similarity with the corresponding sequence of Brachyplatys vahlii (GenBank accession
number AB240167), therefore they identified their species as B. vahlii. Further, they stated that their sample
showed only an 85% similarity to B. subaeneus (GenBank accession number AB240167 [sic]). Both of the above
sequences originate from Hosokawa et al. (2006). The following remarks are made:
1. The GenBank accession number of the sequence of B. subaeneus is AB240166; the number appearing in the
paper of Aiello et al. (2016) is apparently incorrect.
2. The photograph provided by Hosokawa et al. (2006: 1842, fig. 1C) leaves no doubt that the species identified as
B. subaeneus by these authors (GenBank accession number AB240166) is B. vahlii.
3. The identity of B. vahlii sensu Hosokawa et al. (2006) (GenBank accession number AB240167) is uncertain.
Supposing that it is not conspecific with B. subaeneus sensu Hosokawa et al. (2006) (= B. vahlii), and
considering that it shows high similarity with members of the population introduced to Panama readily
identifiable as B. subaeneus based on the photos accompanying the record (Aiello et al. 2016: figs. 2B–C), it
probably pertains to B. subaeneus.
4. The sequences of B. vahlii with GenBank accession numbers KT447151 and KT447152 (Aiello et al. 2016)
pertain to B. subaeneus.
5. Secondary literature citing the results of Hosokawa et al. (2006), e.g. Barton et al. (2007), Kikuchi et al. (2008,
2009) and probably also other related papers based on the same taxa, e.g. Kikuchi & Fukatsu (2003), contain the
same misidentification. Sequences of the symbiont Ishikawaella capsulata extracted from the gut of B.
subaeneus sensu Hosokawa et al. (2006) (AB240159) and B. vahlii sensu Hosokawa et al. (2006) (AB240160,
AB244767) are also associated with this misidentification, therefore the study of Prado & Zucchi (2012) partly
based on these data probably also needs correction. In different pages of the paper by Kikuchi et al. (2008) the
sequence AB244767 is stated to refer to the symbiont extracted from B. vahlii (p. 6) or B. subaeneus (p. 13).
Problems caused by the proliferation of misidentifications of Heteroptera in the NIH genetic sequence
database (GenBank) and the online barcode database (BOLD) were stressed by Lis & Lis (2011) and Lis et al.
(2016). Their criticism is greatly agreed. It is stressed that the situation in GenBank is much worse, since this portal
does not include images of voucher specimens and does not allow corrections or comments from third-party users.
As barcode-based identifications are usually done by users without proper taxonomic background, the data are
frequently applied without appropriate quality control, as it happened in the above discussed case of B. subaeneus.
Misidentifications might rapidly spread in subsequent papers, and their multiplication, together with the frequent
lack of voucher specimens, might make future corrections difficult. Barcoding is already very popular, but future
improvement of the technology and growth of the databases will certainly make barcode-based identifications an
everyday routine. Without necessary control measures misidentifications might, as a worst-case scenario, even
develop to an alternative barcode-based nomenclature in several animal groups.
According to Aiello et al. (2016) B. subaeneus was first detected in Panama in 2012. Lukáš Sekerka (pers.
comm.) has, however, informed me that he encountered it frequently in the southern half of the Canal (from
Gamboa to Panama City) already in 2007. There is no doubt that the species has already established in Central
America.
In its native area B. subaeneus is restricted to the tropical and subtropical moist broadleaf forest zone
(extending in the Sub-Himalayan Belt, southern China, Indochina, and all over the Malay Archipelago), but it does
not enter to the temperate and mixed broadleaf forests of the North China Plain. Accordingly, its expansion to
temperate North America is unlikely, but further expansion to northern part of South America is highly expected.
The species is already common and frequently abundant all over the Canal area of Panama (L. Sekerka pers.
comm.). The available literature records and also the broad native area of the species suggest that it is broadly
oligophagous and it is able to colonize a wide variety of legumes. The available observations (Aiello et al. 2016, L.
Sekerka pers. comm.) indicate that it readily feeds on a variety of native Neotropical fabaceans. As a summary, the
species has the potential to become a widely distributed and significant pest of legume crops in tropical and
subtropical parts of Central and South America.
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Acknowledgements
Amoret Spooner and Katherine Child (OXUM) provided photos of the syntypes of B. subaeneus, Henrik Enghoff
and Mikkel Høegh Post (ZMUC) of syntypes of B. vahlii reproduced in this paper. The comments of Petr Kment,
Lukáš Sekerka (National Museum, Prague) and Jing-Fu Tsai (National Museum of Nature and Science, Taichung)
and the reviews of Joe E. Eger (Dow AgroSciences, Tampa) and Jerzy A. Lis (University of Opole, Opole)
significantly improved the manuscript. The cooperation of all of these colleagues is sincerely appreciated. This
study received financial support from the National Natural Science Foundation of China (grant no. 31472024) and
the One Hundred Young Academic Leaders Program of Nankai University.
References
Ahmad, I. & Moizuddin, M. (1992) Plataspidae Dallas (Hemiptera: Pentatomoidea) from Pakistan and Bangladesh with keys
including Indian taxa. Annotationes Zoologicae et Botanicae, 208, 1–38.
Aiello, A., Saltonstall, K. & Young, V. (2016) Brachyplatys vahlii (Fabricius, 1787), an introduced bug from Asia: first report
in the Western Hemisphere (Hemiptera: Plataspidae: Brachyplatidinae). BioInvasion Records, 5 (1), 7–12.
http://dx.doi.org/10.3391/bir.2016.5.1.02
Atkinson, E.T. (1887) Notes on Indian Rhynchota, Heteroptera, No. 1. Journal of the Asiatic Society of Bengal, 56 (2), 22–61.
Bainbrigge Fletcher, T. (1920) Annotated list of Indian crop-pests. In: Bainbrigge Fletcher, T. (Ed.), Report of the proceedings
of the Third Entomological Meeting held at Pusa on the 3rd to 15th February 1919, Volume III. Superintendent
Government Printing, Calcutta, pp. 33–314.
Banerjee, M.R. (1958) A study of the chromosomes during meiosis in twenty-eight species of Hemiptera (Heteroptera,
Homoptera). Proceedings of the Zoological Society, Calcutta, 11, 9–31.
Bao, Q., Chen, X.Q., Xu, H.L. & Jiang, S.H. (2013) A survey of insect fauna and community of Futian mangrove, Shenzhen.
Journal of Environmental Entomology, 35 (6), 720–727. [in Chinese, English summary]
Barton, N.H., Briggs, D.E.G., Eisen, J.A., Goldstein, D.B. & Patel, N.H. (2007) Evolution. Cold Spring Harbor Laboratory
Press, Cold Spring Harbor, New York, xiv+833 pp.
Boisduval, J.B.A.D. (1835) Voyage de découvertes de l’Astrolabe exécuté par ordre du Roi, pendant les années 1826-1827-
1828-1829 sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l'Océan Pacifique, avec
l’illustration des insectes nouveaux recueillis pendant le voyage. Deuxième partie. Coléoptères et autres ordres. Tastu,
Paris, vii+716 pp. [plates VI–XII issued separately]
Breddin, G. (1900) Hemiptera Sumatrana collecta a dom. Henrico Dohrn. Pars I. Stettiner Entomologische Zeitung, 61, 275–336.
Breddin, G. (1901) Die Hemipteren von Celebes. Ein Beitrag zur Faunistik der Insel. Abhandlungen der Naturforschenden
Gesellschaft zu Halle, 24, 1–213.
Breddin, G. (1905) Versuch einer Rhynchotenfauna der malayischen Insel Banguey. Mitteilungen aus dem Naturhistorischen
Museum in Hamburg, 22, 201–206.
Breddin, G. (1909) Rhynchoten von Ceylon, gesammelt von Dr. Walter Horn. Annales de la Société Entomologique de
Belgique, 53 (6), 250–309.
http://dx.doi.org/10.5962/bhl.part.21874
Cai, W.Z. & Li, H. (2015) [Illustrated insects of China.] Shanxi Science and Technology Press, Taiyuan, [12]+307 pp. [in Chinese]
Chandra, A. (1953) Order Hemiptera (in part). Family Pentatomidae. In: Roonwal, M.L., Mathur, R.N., Bhasin, G.D., Singh,
B., Pant, G.D., Chandra, A., Katiyar, K.N. & Mathur, R.K. A systematic catalogue of the main identified entomological
collection at the Forest Research Institute, Dehra Dun. Parts 9–21. Orders Isoptera, Embioptera, Psocoptera, Anoplura,
Ephemeroptera, Odonata, Thysanoptera and Hemiptera. Government of India Press, Calcutta, pp. 83–102.
Chen, Z.Y. (1985) Brachyplatys subaeneus (Westwood). In: Zhang, S.M. (Ed.), Hemiptera (1). Economic insect fauna of China,
vol. 31. Science Press, Beijing, pp. 32–33. [in Chinese]
Dallas, W.S. (1851) List of the specimens of hemipterous insects in the collection of the British Museum. Part 1. Trustees of the
British Museum, London, 368 pp., XI plates.
Dammerman, K.W. (1929) The agricultural zoology of the Malay Archipelago. The animals injurious and beneficial to
agriculture, horticulture and forestry in the Malay Peninsula, the Dutch East Indies and the Philippines. J.H. De Bussy,
Amsterdam, XI+473 pp., 40 plates.
Datta, B., Ghosh, L.K. & Dhar, M. (1985) Study on Indian Pentatomoidea (Heteroptera: Insecta). Records of the Zoological
Survey of India, Miscellaneous Publication, Occasional Paper, 80, 1–43.
Davidová-Vilímová, J. (2006) Family Plataspidae Dallas, 1851. In: Aukema, B. & Rieger, Ch. (Eds.), Catalogue of the
Heteroptera of the Palaearctic Region. Volume 5. Pentatomomorpha II. The Netherlands Entomological Society, Amsterdam,
pp. 150–165.
Distant, W.L. (1899) Rhynchotal notes.—Heteroptera: Plataspinae, Thyreocorinae, and Cydninae. Annals and Magazine of
Natural History, Series 7, 4 (21), 213–227.
RÉDEI
152
·
Zootaxa 4136 (1) © 2016 Magnolia Press
Distant, W.L. (1900) Revision of the Rhynchota belonging to the family Pentatomidae in the Hope Collection at Oxford.
Proceedings of the Zoological Society of London, 1900, 807–825, plates LII–LIII.
Distant, W.L. (1902) The fauna of British India, including Ceylon and Burma. Rhynchota, vol. 1. Heteroptera. Tay lor &
Francis, London, xxxviii+438 pp.
Easton, E.R. & Pun, W.W. (1997) Observations on some Hemiptera/Heteroptera of Macau, Southeast Asia. Proceedings of the
Entomological Society of Washington, 99 (3), 574–582.
Eger, J.E. Jr., Ames, L.M., Suiter, D.R., Jenkins, T.M., Rider, D.A. & Halbert, S.E. (2010) Occurrence of the Old World bug
Megacopta cribraria (Fabricius) (Heteroptera: Plataspidae) in Georgia: A serious home invader and potential legume pest.
Insecta Mundi, 121, 1–11.
Esaki, T. (1926) Verzeichniss der Hemiptera-Heteroptera der Insel Formosa. Annales Musei Nationalis Hungarici, 24, 136–189.
Fabricius, J.C. (1787) Mantissa insectorum sistens species nuper detectas adiectis synonymis, observationibus, descriptionibus,
emendationibus. Tom. II. Proft, Hafniae, 382 pp.
Hafiz, H.A. & Ribeiro, S. (1941) Notes on Indian Hemiptera in the Zoological Survey of India. I. On the subfamily Plataspinae
of the family Pentatomidae. Records of the Indian Museum, 43 (1), 73–85.
Hayashi, M. (2002) Heteroptera. In: Azuma, S., Yafuso, M., Kinjo, M., Hayashi, M., Kohama, T., Sasaki, T., Kimura, M. &
Kawamura, F. (Eds.), Check list of the insects of the Ryukyu Islands. Second edition. The Biological Society of Okinawa,
Nishihara, pp. 125–149. [in Japanese]
Hoffmann, W.E. (1931) Notes on Hemiptera and Homoptera at Canton, Kwangtung Province, Southern China, 1924–1929, 1.
Insect Pest Survey Bulletin, 11 (3), 138–151.
Hoffmann, W.E. (1932) Notes on the bionomics of some oriental Pentatomidae (Hemiptera). Archivio Zoologico Italiano, 16
(3–4), 1010–1027.
Hoffmann, W.E. (1935) An abridged catalogue of certain Scutelleroidea (Plataspidae, Scutelleridae, and Pentatomidae) of
China, Chosen, Indo-China, and Taiwan. Lignan University Science Bulletin, 7, 1–294.
Hosokawa, T., Kikuchi, Y. Nikoh, N., Shimada, M. & Fukatsu, T. (2006) Strict host-symbiont cospeciation and reductive
genome evolution in insect gut bacteria. PloS Biology, 4 (10), 1841–1851.
http://dx.doi.org/10.1371/journal.pbio.0040337
Hsiao, T.Y. & Jen, S.C. (1977) Plataspidae. In: Hsiao, T.Y., Jen, S.C., Cheng, L.I., Liu, S.L. & Ching, H.L. A handbook for the
determination of the Chinese Hemiptera-Heteroptera. Vol. I. Science Press, Beijing, pp. 14–38, 292–295, plates 1–3. [In
Chinese, English summary]
Johnson, N.F. (1996) Revision of World species of Paratelenomus Dodd (Hymenoptera: Scelionidae). Canadian Entomologist,
128, 273–291.
http://dx.doi.org/10.4039/Ent128273-2
Kershaw, J.C.W. & Kirkaldy, G.W. (1910) On the metamorphoses of two coptosomine Hemiptera from Macao. Annales de la
Société Entomologique de Belgique, 54, 69–73.
Kikuchi, Y. & Fukatsu, T. (2003) Diversity of Wolbachia endosymbionts in heteropteran bugs. Applied and Environmental
Microbiology, 69 (10), 6082–6090.
http://dx.doi.org/10.1128/AEM.69.10.6082-6090.2003
Kikuchi, Y., Hosokawa, T. & Fukatsu, T. (2008) Diversity of bacterial symbiosis in stinkbugs. In: Dijk, Th. van (Ed.) Microbial
ecology research trends. Nova Science Publishers, New York, pp. 39–63.
Kikuchi, Y., Hosokawa, T., Nikoh, N., Meng, X.Y., Kamagata, Y. & Fukatsu, T. (2009) Host-symbiont co-speciation and
reductive genome evolution in gut symbiotic bacteria of acanthosomatid stinkbugs. BMC Biology, 7 (2), 1–22.
http://dx.doi.org/10.1186/1741-7007-7-2
Kirby, W.F. (1891) Catalogue of the described Hemiptera Heteroptera and Homoptera of Ceylon, based on the collection
formed (chiefly at Pundaloya) by Mr. E. Ernest Green. Journal of the Linnean Society of Zoology, 24, 72–176, plates 4–6.
http://dx.doi.org/10.1111/j.1096-3642.1891.tb02479.x
Kirkaldy, G.W. (1909) Catalogue of the Hemiptera (Heteroptera) with biological and anatomical references, lists of foodplants
and parasites, etc. Vol. I. Cimicidae. Berlin, xl+392 pp.
Kirkaldy, G.W. (1910) A list of the Hemiptera of Oriental China. Part II. Annales de la Société Entomologique de Belgique, 54
(4), 103–112.
Kuhlgatz, Th. (1901) Uebersicht über die indo-australischen Coptosoma-Arten aus der Verwandtschaft von Coptosoma
cinctum (Eschz.). Archiv für Naturgeschichte, 67, 205–264, plates VIII–IX.
Laporte, F.L. de (1832–1833) Essai d’une classification systématique de l’ordre des Hémiptères (Hémiptères Hétéroptères,
Latr.). Magasin de Zoologie, 2 (suppl.), 1–88, plates 51–55. [1832, 1–16, 1833, 17–88]
Lethierry, L. & Severin, G. (1893) Catalogue général des Hémiptères. Tome I. Hétéroptères, Pentatomidae. F. Hayez,
Bruxelles, x+286 pp.
Li, L.Y, Wang, R. & Waterhouse, D.F. (1997) The distribution and importance of arthropod pests and weeds of
agriculture and forestry plantations in Southern China. Chinese Academy of Agricultural Sciences, Guangdong
Entomological Institute, and Australian Centre for International Agricultural Research, Beijing, Guangzhou and
Canberra, x+185 pp.
Lin, Y.J. & Zhang, S.M. (1998) [A checklist of Plataspidae (Hemiptera: Pentatomoidea) of China (I)]. Jiangxi Plant Protection,
21 (3), 4–8. [in Chinese]
Zootaxa 4136 (1) © 2016 Magnolia Press
·
153
BRACHYPLATY S INTRODUCED TO PANAMA
Lis, J.A. & Lis, B. (2011) Is accurate taxon identification important for molecular studies? Several cases of faux pas in
pentatomoid bugs (Hemiptera: Heteroptera: Pentatomoidea). Zootaxa, 2932, 47–50.
Lis, J.A., Lis, B. & Ziaja, D.J. (2016) In BOLD we trust? A commentary on the reliability of specimen identification for DNA
barcoding: a case study on burrower bugs (Hemiptera: Heteroptera: Cydnidae). Zootaxa, 4114 (1), 83–86.
Liu, G.Q. & Zheng, L.Y. (2006) Checklist of the Chinese Plataspidae (Heteroptera, Pentatomoidea). In: Rabitsch, W. (Ed.), Hug
the bug—For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia, 19, 275–398.
Manna, G.K. (1951) A study of the chromosomes during meiosis in forty-three species of Indian Heteroptera. Proceedings of
the Zoological Society of Bengal, 4 (1–2), 1–116.
Maxwell-Lefroy, H. (1909a) The insect fauna of Tirhut. I.–Rhynchota Heteroptera. Records of the Indian Museum, 3, 301–338.
Maxwell-Lefroy, H. (1909b) Indian Insect Life. A manual of the insects of the plains (tropical India). Thacker and Spink,
Calcutta, xii+786 pp., LXXXIV plates.
Miller, N.C.E. (1931) The bionomics of some Malayan Rhynchota (Hemiptera-Heteroptera). Department of Agriculture Straits
Settlements and Federated Malay States, Scientific Series, 5, 1–142.
Miller, N.C.E. (1934) The developmental stages of some Malayan Rhynchota. Journal of the Federated Malay States Museum,
17 (3), 502–525.
Mitsuhashi, S. (1915) A list of Pentatomidae of Japan [part 1]. Insect World, 19 (219), 481–484. [in Japanese]
Montandon, A.L. (1894) Viaggio di Leonardo Fea in Birmania e regioni vicine. LVIII. Hémiptères de la s. fam. des
Plataspidinae récoltés par M. Leonardo Fea en Birmanie et régions voisines. Annali del Museo Civico di Storia Naturale di
Genova, Serie 2
a
, 14, 119–144.
Montandon, A. L. (1897) Les Plataspidines du Muséum d’histoire naturelle de Paris. Annales de la Société Entomologique de
France, 65 [1896], 436–464.
Nagashima, S & Tanaka, Y. (2014) A new distributional record of Brachyplatys vahlii (Heteroptera, Plataspidae) from Ishigaki-
jima Island of the Ryukyus, Japan. Rostria, 56, 51–52. [in Japanese]
Noualhier, M. & Martin, J. (1904) Hémiptères recueillis par M.A. Pavie. In: Pavie, A. (Ed.), Mission Pavie Indo-Chine 1879–
1895. Études diverses III. Recherches sur l’histoire naturelle de l’Indo-Chine orientale. Ernest Leroux, Paris, pp. 165–185,
plates X–XI.
Pol, Ch., Belfield, S. & Martin, R. (2010) Insects of upland crops in Cambodia. ACIAR Monograph No. 143. Australian Centre
for International Agricultural Research, Canberra, 132 pp.
Prado, S.S. & Zucchi, T.D. (2012) Host-symbiont interactions for potentially managing heteropteran pests. Psyche, 2012,
Article ID 269473, 1–9.
http://dx.doi.org/10.1155/2012/269473
Ren, S.Z. (1987) Plataspidae. In: Huang, F.S. (Ed.), Forest insects of Yunnan. Yunnan Science and Technology Press, Kunming,
pp. 104–109. [in Chinese]
Ruberson, J.R., Takasu, K., Buntin, G.D., Eger, J.E. Jr., Gardner, W.A., Greene, J.K., Jenkins, T.M., Jones, W.A., Olson, D.M.,
Roberts, P.M., Suiter, D.R. & Toews, M.D. (2013) From Asian curiosity to eruptive American pest: Megacopta cribraria
(Hemiptera: Plataspidae) and prospects for its biological control. Applied Entomology and Zoology, 48 (1), 3–13.
http://dx.doi.org/10.1007/s13355-012-0146-2
Stål, C. (1876) Enumeratio Hemipterorum. Bidrag till en förteckning öfver alla hittills kända Hemiptera, jemte systematiska
meddelanden. 5. Kongliga Svenska Vetenskaps-Akademiens Handlingar (N.F.), 14 (4), 1–162.
Tomokuni, M. (Ed.) (1993) A field guide to Japanese bugs. Terrestrial heteropterans. Zenkoku Noson Kyoiku Kyokai, Tokyo,
380 pp. [in Japanese]
Tsai, J.F., Rédei, D., Yeh, G.F. & Yang, M.M. (2011) Jewel bugs of Taiwan (Heteroptera: Scutelleridae). National Chung Hsing
University, Taichung, 309 pp.
Tsai, J.F. & Yang, M.M. (2005) [The primary identification of phytophagous and predaceous bugs (Hemiptera: Heteroptera).] In:
Lu, K.H. (Ed.), [Workshop on the identification and diagnosis of important pests of plant health inspection and quarantine
(special issue)], 5, 81–111. Bureau of Animal and Plant Health Inspection and Quarantine, Council of Agriculture, Executive
Yuan, Taipei, and Department of Entomology, National Chung Hsing University, Taichung. [in Chinese]
Ueshima, N. (1979) Hemiptera II: Heteroptera. In: John, B. (Ed.), Animal cytogenetics, 3, 1–117. Gebrüder Borntraeger, Berlin,
Stuttgart.
Vayssière, P. (1958) Entomologie malaise (Lépidoptères, Coléoptères, Hémiptères) (suite et fin). Journal d’Agriculture
Tropicale et de Botanique Appliquée, 5 (12), 789–808.
http://dx.doi.org/10.3406/jatba.1958.2516
Vollenhoven, S.C.S. van (1863) Essai d’une faune entomologique de l’Archipel Indo-Néerlandais. Première monographie:
Famille des Scutellérides. Martinus Nijhoff, La Haye, 60 pp., IV plates.
Walker, F. (1867) Catalogue of the specimens of Heteropterous-Hemiptera in the collection of the British Museum. Part I.
Scutata. Trustees of the British Museum, London, 240 pp.
Wall, R.E. (1928) A comparative study of a chalcid egg parasite in three species of Plataspidinae. Lingnan Science Journal, 6
(3), 231–239.
Wall, R.E. (1931) A study of Brachyplatus subaëneus Westw. Lingnan Science Journal, 7 [1929], 633–648, 1 plate.
Westwood, J.O. (1837) In: Hope, F.W.(Ed.), A catalogue of Hemiptera in the collection of the Rev. F.W. Hope, M.A., with short
Latin diagnoses of the new species. Bridgewater, London, 46 pp.
RÉDEI
154
·
Zootaxa 4136 (1) © 2016 Magnolia Press
White, A. (1839) Description of two hemipterous insects. Magazine of Natural History, New Series, 3 (35), 537–543.
Wu, C.F. (1935) Catalogus insectorum sinensium. Volume II. Fan Memorial Institute of Biology, Peiping, 634 pp. [Family
Pentatomidae (pp. 275–382) by T.H. Tang]
Wu, M.X. & Xu, K.T. (2002) Preliminary studies on the pests of green soybean in Fuzhou suburbs. Wuyi Science Journal, 18,
27–32. [in Chinese, English summary]
Yamashita, I. & Kikuhara, Y. (2012) Family Plataspidae Dallas, 1851. In: Ishikawa, T., Takai, M. & Yasunaga, T. (Eds.), A field
guide to Japanese bugs—Terrestrial heteropterans. Vol. 3. Zenkoku Noson Kyoiku Kyokai Publishing Co., Tokyo, pp.
439–444, plates 102–103. [in Japanese]
Yang, W.I. (1934) Revision of Chinese Plataspidae. Bulletin of the Fan Memorial Institute of Biology, Zoology, 5 (3), 137–235.
Yiu, V. & Yip, C.H. (2012) A photographic guide to Hong Kong true bugs. Hong Kong Entomological Society, Hong Kong,
263 pp. [in Chinese]
Zhang, S.M. & Sie, R.F. (1986) New records and faunal structure of plataspid, acanthosomatid and urostylid bugs (Hemiptera)
in Fujian Province, China. Wuyi Science Journal, 6, 97–101. [in Chinese, English summary]
Zheng, S.Z. & Lin, Y.X. (2013) Stinkbug. In: Xu, H.Y. (Ed.), [Taiwan Nature Identification Series], 29, 1–381. Morning Star
Publishing, Taipei. [in Chinese]
Zimsen, E. (1964) The type material of I.C. Fabricius. Munksgaard, Copenhagen, 656 pp.
