ArticlePDF Available

A review of the fruit fly tribe Pliomelaenini (Diptera: Tephritidae: Tephritinae) in the Indo-Australian Region

Authors:
  • Independent researcher - retired from Government Service

Abstract

Five genera and 17 species o f Indo-Australian Tephritinae are placed in the tribe Pliomelaenini. Quadrimelaena gen. n. is described to include Q. quadrimaculata (Agarwal & Kapoor), comb, n. from India, Q. sonani (Shiraki), comb. n. from Taiwan [type species] and Q. translúcida (Hering), comb. n. from Sri Lanka [all transferred from Pliomelaena Bezzi].
A REVIEW OF THE FRUIT FLY TRIBE PLIOMELAENINI
(DIPTERA: TEPHRITIDAE : TEPHRITINAE)
IN THE INDO-AUSTRALIAN REGION
D.L. HANCOCK
PO Box 2464, Cairns, Qld 4870
Abstract
Five genera and 17 species of Indo-Australian Tephritinae are placed in the tribe Pliomelaenini.
Quadrimelaena gen. n. is described to include Q. quadrimaculata (Agarwal & Kapoor), comb,
n. from India, Q. sonani (Shiraki), comb. n. from Taiwan [type species] and Q. translúcida
(Hering), comb. n. from Sri Lanka [all transferred from Pliomelaena Bezzi].
Introduction
The tribe Pliomelaenini is an Afrotropical and Indo-Australian group of
Tephritinae that breeds in the flowerheads of Acanthaceae such as Asystasia,
Dicliptera, Hypoestes, Justicia and Lepidagathis (Hancock et al. 2003).
Previously known as tribe Platensinini, that name was transferred to the
Dithrycini as a subtribe and redefined by Hancock (2001). All species have 3
pairs of frontal, 2 pairs of orbital and 4 scutellar setae, mostly of uniform
coloration although the upper pair of orbitals are sometimes paler.
Five genera occur in the Indo-Australian region: Elaphromyia Bigot,
Pliomelaena Bezzi, Pseudafreutreta Hering, Quadrimelaena gen. n. and
Sundaresta Hering. The first three also occur in Africa. The 17 Indo-
Australian species are poorly understood, with most known from very few
specimens. Three species occur in Indonesia and two in Papua New Guinea
but the tribe has not yet been recorded from Australia.
One currently included species is excluded from the tribe; others were noted
by Hancock (2001). 4Elaphromyia magna Hardy, from Java (Hardy 1988),
has a relatively short wing, bare arista, subshining black spot between eye and
antennal base, dense scale-like setae on the scutum and a densely tomentose
abdomen and does not belong in Elaphromyia. Its taxonomic position is
uncertain but it appears to belong near Afreutreta Bezzi in tribe Eutretini.
Elaphromyia Bigot
This genus is characterised by the elongate, almost parallel-sided wings with
numerous subhyaline or diffusely yellow spots on a brown pattern. Vein Rj
lacks a bare, non-setose dorsal area below the end of vein Sc. Six Indo-
Australian species are known.
Elaphromyia hardyi Wang, from southwestern China (Sichuan), differs from
all other species in having a basomedial longitudinal hyaline band on the
wing. It was illustrated by Wang (1998).
Elaphromyia multisetosa Shiraki, from Taiwan, is distinguished from E.
pterocallaeformis by the palpi, which have numerous stout, black setae on
their ventral margins. It was illustrated by Shiraki (1933).
Elaphromyia pterocallaeformis (Bezzi) is widespread in south and southeast
Asia, including Indonesia (Hering 1941). It lacks a medial longitudinal
hyaline band and the palpi have only a few yellowish setae on their ventral
margins. It was illustrated by Hardy (1974) and Wang (1998). E. incompleta
Shiraki and E. i. punctata Shiraki were placed as synonyms by Wang (1998).
Elaphromyia siva Frey is a small species known only from Sri Lanka.
Elaphromyia transversa Hardy occurs in Papua New Guinea. It resembles E.
pterocallaeformis but the subhyaline spots on the wing are arranged in
transverse rather than longitudinal rows. It was illustrated by Hardy (1988).
Elaphromyia yunnanensis Wang is known from southwestern China
(Sichuan, Yunnan). It differs from E. pterocallaeformis in the larger and more
extensive hyaline wing spots along the anterior and posterior margins and the
shorter oviscape. It was illustrated by Wang (1998).
Pliomelaena Bezzi
Indo-Australian species generally have the head and thoracic setae yellow
with a brownish tint, except for the postvertical, lateral vertical and postocular
setae, which are whitish. The wing has three hyaline spots in cells X\ and r2 +3
forming a more or less distinct ‘V (as in all Afrotropical species) and three
hyaline indentations in cell m. Protephritis Shiraki and Indaresta Hering are
synonyms (Hardy 1988). Five species are included.
Pliomelaena callista (Hering) is known from Indonesia and Papua New
Guinea. The wing pattern is a little variable but it has three hyaline
indentations in cell r! and no hyaline spot in cell br before R-M crossvein. It
was illustrated by Hering (1941) and Hardy (1988).
Pliomelaena luzonica Hardy is known only from Luzon in the Philippines. It
is similar to P. callista but has only two hyaline indentations in cell r¡. It was
illustrated by Hardy (1974).
Pliomelaena sauteri (Enderlein) is known from Taiwan and southeast China
(Hainan). It is very similar to P. callista but has the hyaline spot at the base of
the pterostigma well developed, rather than vestigial or absent. It was
illustrated by Enderlein (1911) and Wang (1998).
Pliomelaena udhampurensis Agarwal & Kapoor is known only from
northwestern India (Jammu and Kashmir). It is similar to P. luzonica but has
a hyaline spot in cell br before R-M crossvein and the abdomen is almost
entirely brown to black. The yellowish head and thoracic setae are a darker
brown than in other species. It was illustrated by Agarwal and Kapoor (1988).
Pliomelaena zonogastra (Bezzi) is known from India (including Nicobar
Islands) and southwestern China (Yunnan). It is similar to P. udhampurensis
but the abdomen is largely reddish-yellow, rather than brown to black. It was
illustrated by Wang (1998).
Pseudafreutreta Hering
This largely Afrotropical genus differs from Pliomelaena in the absence of a
row of setae along the sides of the epistome and the darker wings, the hyaline
spots and indentations being small or largely absent. Vein Ri lacks a bare,
non-setose dorsal area below the end of vein Sc (present in Pliomelaena).
Pseudafreutreta nigrifacies (Wang) occurs in southwestern China (Yunnan)
and northern Thailand. It was transferred from Platensina Enderlein by
Hancock (2001) and illustrated by Hancock and McGuire (2002). A male has
been collected on Strobilanthes imbricatus (Acanthaceae) in Thailand
(Hancock and McGuire 2002).
Quadrimelaena gen. n.
Type species Protephritis sonani Shiraki, 1933, by present designation.
Quadrimelaena closely resembles Pliomelaena in most characters, including
the presence of a row of setae along the sides of the epistome, but these setae
are weaker, the wings are narrower and more elongate, there are two hyaline
spots (one anterior and one posterior, rather than one medial) in the outer half
of cell dm beyond the level of the R-M crossvein and four (rather than three)
marginal/submarginal spots or indentations in cell m, the inner, extra
indentation above the apex of vein Cui. The hyaline spots in cells and r2 +3
do not form a ‘V’ and several small posterior spots (absent in Pliomelaena)
are present in cell ri along vein R2+3. It further differs from Asian
Pliomelaena species in the more rounded third antennal segment (rather than
slightly concave dorsally), longer and almost parallel ocellar setae and the
brown rather than yellowish head and thoracic setae, except for the whitish
postvertical, lateral vertical and postocular setae. For a more detailed
description see that of the type species (Shiraki 1933).
Three species are included, all transferred from Pliomelaena. Korneyev
(1999) suggested that Ictericodes cashmerensis (Hendel) might also belong
here but its head and thoracic setae are mostly yellow-brown with the upper
orbitals paler, there is a brown spot between the eye and antennal base, vein
R4 +5 is extensively setose above and below and cell bcu has a longer apical
extension.
Quadrimelaena quadrimaculata (Agarwal & Kapoor), comb. n. is known
only from northwestern India (Himachal Pradesh). It differs from Q. sonani in
the fewer small hyaline discal spots and was illustrated by Agarwal and
Kapoor (1989).
Quadrimelaena sonani (Shiraki), comb. n. occurs in Taiwan. It was originally
placed in Protephritis and illustrated by Shiraki (1933).
Quadrimelaena translúcida (Hering), comb. n. occurs in Sri Lanka. It differs
from the other species in the more extensive hyaline discal areas and lack of a
hyaline apical spot on the wing. It was illustrated by Hering (1942).
Sundaresta Hering
This genus differs from Pliomelaena primarily in the presence of two (rather
than one) distinct marginal hyaline indentations in cell r2+3. The oviscape is
elongate, as long as or longer than the abdomen.
Sundaresta hilaris Hering is known only from Java and Sumba in Indonesia.
It was illustrated by Hardy (1988).
Sundaresta malaisei (Hering) is widespread in India, Burma, southwestern
China (Yunnan), Thailand and Laos. It differs from S. hilaris in the more
numerous small hyaline spots on the wing. It was transferred from Xyphosia
Robineau-Desvoidy and illustrated by Hancock and McGuire (2002). Both
sexes have been collected on Strobilanthes imbricatus (Acanthaceae) in
Thailand (Hancock and McGuire 2002).
References
AGARWAL, M.L. and KAPOOR, V.C. 1988. Four new species of fruit flies (Diptera:
Tephritidae: Tephritini) together with redescription of Trupanea inaequabilis Hering and their
distribution in India. Journal o f Entomological Research (New Delhi) 12: 117-128.
AGARWAL, M.L. and KAPOOR, V.C. 1989. New Tephritidae (Diptera) from India.
Entom ologist’s Monthly Magazine 125: 31-35.
ENDERLEIN, G. 1911. Trypetiden-Studien. Zoologische Jahrbücher. Abteilung fiir Systematik,
Oekologie und Geographie der Tierre 31: 407-460.
HANCOCK, D.L. 2001. Systematic notes on the genera of Australian and some non-Australian
Tephritinae (Diptera: Tephritidae). Australian Entomologist 28: 111-116.
HANCOCK, D.L. and McGUIRE, D.J. 2002. New species and records of non-dacine fruit flies
(Diptera: Tephritidae) from south and southeast Asia. Steenstrupia 27: 1-17.
HANCOCK, D.L., KIRK-SPRIGGS, A.H. and MARAIS, E. 2003. New records of Namibian
Tephritidae (Diptera: Schizophora), with notes on the classification of subfamily Tephritinae.
Cimbebasia 18: 49-70.
HARDY, D.E. 1974. The fruit flies o f the Philippines (Diptera: Tephritidae). Pacific Insects
Monograph 32: 1-266.
HARDY, D.E. 1988. The Tephritinae of Indonesia, New Guinea, the Bismarck and Solomon
Islands (Diptera: Tephritidae). Bishop Museum Bulletin in Entomology 1: i-vii, 1-92.
HERING, E.M. 1941. Dipteren von den Kleinen Sunda-Inseln. ü. Trypetidae. Arbeiten über
morphologische und taxonomische Entomologie aus Berlin-Dahlem 8: 24-45.
HERING, E.M. 1942. Neue Gattungen und Arten palaearktischer und exotischer Fruchtfliegen.
Siruna Seva 4:1-31.
KORNEYEV, V.A. 1999. Phylogeny of the subfamily Tephritinae: relationships of the tribes
and subtribes. Pp 549-580, in: Aluja, M. and Norrbom, A.L. (eds), Fruit flies (Tephritidae):
phylogeny and evolution o f behavior. CRC Press, Boca Raton; 944 pp.
SHIRAKI, T. 1933. A systematic study o f Trypetidae in the Japanese Empire. Memoirs of the
Faculty o f Science and Agriculture, Taihoku Imperial University 8(Entomology 2): 1-509.
WANG, X.-j. 1998. The fruit flies (Diptera: Tephritidae) of the East Asian Region. Acta
Zootaxonomica Sinica 21(Supplement 1996): i-viii, 1-378, pis 1-41.
... Elaphromyia Bigot is a small genus of tephritid flies in the tribe Pliomelaenini of the subfamily Tephritinae, with 10 previously described species from the Oriental, Afrotropical, Palearctic and Australasian Regions , one of which, the Indonesian E. magna Hardy, was transferred to Afreutreta Bezzi in tribe Eutretini by Hancock (2004Hancock ( , 2007. Species of this genus are characterized by elongate, parallel-sided, predominantly dark brown wings with a series of subhyaline spots and the dorsal setulae on vein R 1 without a gap posterior to the end of vein Sc. Hancock (2004) provided taxonomic notes on Elaphromyia species then known from the Indo-Australian Region but, apart from illustrations of E. pterocallaeformis (Bezzi) and E. transversa Hardy provided by Hardy (1973Hardy ( , 1974Hardy ( , 1988, postabdominal structures of most species had not been studied previously. ...
... Elaphromyia Bigot is a small genus of tephritid flies in the tribe Pliomelaenini of the subfamily Tephritinae, with 10 previously described species from the Oriental, Afrotropical, Palearctic and Australasian Regions , one of which, the Indonesian E. magna Hardy, was transferred to Afreutreta Bezzi in tribe Eutretini by Hancock (2004Hancock ( , 2007. Species of this genus are characterized by elongate, parallel-sided, predominantly dark brown wings with a series of subhyaline spots and the dorsal setulae on vein R 1 without a gap posterior to the end of vein Sc. Hancock (2004) provided taxonomic notes on Elaphromyia species then known from the Indo-Australian Region but, apart from illustrations of E. pterocallaeformis (Bezzi) and E. transversa Hardy provided by Hardy (1973Hardy ( , 1974Hardy ( , 1988, postabdominal structures of most species had not been studied previously. Wang (1998) studied Elaphromyia species from East and South-East Asia and provided wing illustrations of E. hardyi Wang, E. yunnanensis Wang and E. pterocallaeformis. ...
Article
A new species of Elaphromyia, E. juncta David, Hancock & Sachin, sp. n. is described from India. It can be differentiated from the morphologically similar E. siva Frey and E. pterocallaeformis (Bezzi) by the wing pattern, epandrial characters, morphology of spicules on the eversible membrane and spermathecal shape. Elaphromyia siva Frey and E. yunnanensisWang are recorded for the first time from India. Records of E. pterocallaeformis (Bezzi) from southern India are regarded as misidentifications. A key to the 7 known non-African species is included.
... Hardy (1973) listed 211 species in the Tenasserim Division of Myanmar, Thailand, Cambodia, Laos, Vietnam, Peninsular Malaysia and Singapore. This list has not been updated greatly although Hancock (1999Hancock ( , 2004Hancock ( , 2011Hancock ( , 2012, Hancock & Drew (1994a,b, 1999, 2004, Hancock & McGuire (2002) and Chua (2010) added some new species and records and synonymised or transferred many other species from peninsular Southeast Asia. Among these, only 10 species were previously recorded in Cambodia by Hardy (1973), based on a small collection made by N. R. Spencer at 700 m asl (above sea level) in Kirirom National Park (Kampong Speu Province) in April 1961. ...
... Hardy (1973) listed 211 species in the Tenasserim Division of Myanmar, Thailand, Cambodia, Laos, Vietnam, Peninsular Malaysia and Singapore. This list has not been updated greatly although Hancock (1999Hancock ( , 2004Hancock ( , 2011Hancock ( , 2012, Hancock & Drew (1994a,b, 1999, 2004, Hancock & McGuire (2002) and Chua (2010) added some new species and records and synonymised or transferred many other species from peninsular Southeast Asia. Among these, only 10 species were previously recorded in Cambodia by Hardy (1973), based on a small collection made by N. R. Spencer at 700 m asl (above sea level) in Kirirom National Park (Kampong Speu Province) in April 1961. ...
... The fruit fly tribe Pliomelaenini contains several Indo-Australian species associated with the flowers of Acanthaceae. They were reviewed recently by Hancock (2004) but continued investigation has revealed a new synonym and a new combination within the group. ...
... This species is known from NE Afghanistan and NW India. For further discussion see Hancock (2004). ...
Article
Full-text available
Pliomelaena udhampurensis Agarwal & Kapoor is placed as a new synonym o f P. spathuliniforma (Dirlbek & Dirlbek), comb. n. [transferred from M etasphenisca Hendel]. The species is known from NW India and NE Afghanistan. Introduction The fruit fly tribe Pliomelaenini contains several Indo-Australian species associated with the flowers of Acanthaceae. They were reviewed recently by Hancock (2004) but continued investigation has revealed a new synonym and a new combination within the group.
... Studies of the diversity in South-East Asia were provided by several researchers, but knowledge about Tephritidae diversity in this area remains incomplete. In addition, knowledge of fruit fl ies in this area was signifi cantly expanded by Hancock (1999Hancock ( , 2004Hancock ( , 2011Hancock ( , 2012Hancock ( , 2013Hancock ( , 2014, a, b, 1999, Hancock & McGuire (2002), and Chua (2010). Th e fauna of Dacinae was recently documented in Bangladesh by Leblanc et al. (2021). ...
Article
Full-text available
Based on a field survey from 2017, twenty-three species are reported from Bhutan for the first time: Acroceratitis ceratitina, A. hardyi, Anomoia approximata, Bactrocera connecta, B. latifrons, B. nigrifacia, B. syzygii, Campiglossa sororcula, Cecidochares connexa, Dacus jacobi, Gastrozona fasciventris, Hoplandromyia antelopa, Lenitovena ultima, Ptilona confinis, Rioxoptilona dunlopi, R. formosana, R. vaga, Spathulina acroleuca, Themara yunnana, Trypeta indica, Zeugodacus apiciflavus, Z. diversus, and Z. fereuncinatus. Four species of the genera Cornutrypeta, Hemilea, Morinowotome, and Vidalia are also recorded for the first time, but the precise determination to species needs additional study and material. As the result, 71 species is listed from Bhutan by far. Their taxonomic position and key characters are discussed. Illustrations for most of the newly recorded species are given.
... Hardy (1973) listed 211 species in the Tenasserim Division of Myanmar, Thailand, Cambodia, Laos, Vietnam, Peninsular Malaysia and Singapore. This list has not been updated greatly although Hancock (1999Hancock ( , 2004Hancock ( , 2011Hancock ( , 2012, Hancock & Drew (1994a,b, 1999, Hancock & McGuire (2002) and Chua (2010) added some new species and records and synonymised or transferred many other species from peninsular Southeast Asia. Among these, only 10 species were previously recorded in Cambodia by Hardy (1973), based on a small collection made by N. R. Spencer at 700 m asl (above sea level) in Kirirom National Park (Kampong Speu Province) in April 1961. ...
Article
Full-text available
The true fruit flies (Tephritidae) of Cambodia are poorly known: only 10 species were recorded by Hardy (1973) and 27 species (all of the genus Bactrocera) by Leblanc et al. (2015). Based on additional material collected in June 2017, we document eight additional species for the fi rst time in Cambodia. We also provide a list of 44 species of tephritid fl ies arranged in 12 genera which are currently known to occur in Cambodia, with notes on their distribution and biology.
... Afreutreta magna (Hardy), comb. n. is transferred from Elaphromyia Bigot; this placement was initially suggested by Hancock (2004). It is known only from Java in Indonesia. ...
Article
Full-text available
Two genera and species of Indo-Australian Tephritidae are placed in the tribe Eutretini, viz. Afreutreta magna (Hardy), comb. n. and Perirhithrum longiseta (Hering).
... It is interesting that Hancock (2001) classified Oedaspis in the Platensinina (as a subtribe of Dithrycini), which was previously included in the Tephrellini. Hancock et al. (2004) later treated the Platensinina as the separate tribe Pliomelaenini. ...
Article
Full-text available
The phylogeny of the subfamily Tephritinae (Diptera: Tephritidae) was reconstructed from mitochondrial 16S ribosomal RNA gene sequences using 53 species representing 11 currently recognized tribes of the Tephritinae and 10 outgroup species. The minimum evolution and Bayesian trees suggested the following phylogenetic relationships: (1) monophyly of the Tephritinae was strongly supported; (2) a sister group relationship between the Tephritinae and Plioreocepta was supported by the Bayesian tree; (3) the tribes Tephrellini, Myopitini, and Terelliini (excluding Neaspilota) were supported as monophyletic groups; (4) the non-monophyletic nature of the tribes Dithrycini, Eutretini, Noeetini, Tephritini, Cecidocharini, and Xyphosiini; and (5) recognition of 10 putative tribal groups, most of which were supported strongly by the statistical tests of the interior branches. Our results, therefore, convincingly suggest that an extensive rearrangement of the tribal classification of the Tephritinae is necessary. Since our sampling of taxa heavily relied on the current accepted classification, some lineages identified by the present study were severely under-sampled and other possible major lineages of the Tephritinae were probably not even represented in our dataset. We believe that our results provide baseline information for a more rigorous sampling of additional taxa representing all possible major lineages of the subfamily, which is essential for a comprehensive revision of the tephritine tribal classification.
Article
An updated checklist for 335 species of fruit flies of India belonging to 97 genera in 22 tribes and 5 subfamilies is provided. A new species of Oedaspis Loew, O. freidbergi David & Singh, sp. nov. is described along with five new records namely Bactrocera gombokensis Drew & Hancock, Chaetellipsis alternata (Zia), Goniurellia persignata Freidberg, Actinoptera reticulata Ito and Terellia serratulae (Linnaeus).
Article
Full-text available
The Australian genera of Tephritinae are assigned to tribes, subtribes and genus-groups. The Platensina group is redefined to contain 6 genera: Australasinia gen. nov., Bezzina Munro, Collessomyia Hardy & Drew, Euthauma Munro, Malaisinia Hering and Platensina Enderlein. It is placed with the Oedaspis group in subtribe Platensinina (= Oedaspidina), the larvae o f which form stem galls on Asteraceae, Goodeniaceae and Onagraceae. The African Chipingomyia nigrapex (Munro), comb. nov. is transferred from Bezzina. The following 14 new combinations for Indo-Australian species are proposed: Australasinia sexincisa (Malloch), comb, nov., Collessomyia heringi (Hardy), comb, nov., C. trimaculata (Hardy), comb, nov., Malaisinia variegata (Radhakrishnan), comb. nov. and Oedaspis australis (Malloch), comb. nov. (all transferred from Hendrella Munro); Bezzina assimilis (Shiraki), comb, nov., B. shirouzui (Ito), comb. nov. and Malaisinia biseta (Wang), comb. nov. (transferred from Pliomelaena Bezzi); Bezzina quadrula (Hardy), comb, nov., B. trimaculata (Hardy & Drew), comb. nov. and Pseudafreutreta nigrifacies (Wang), comb. nov. (transferred from Platensina)', Liepana apiciclara (Hardy & Drew), comb. nov. (transferred from Oedaspis Loew); Scedella infrequens (Hardy & Drew), comb. nov. and S. orientalis (de Meijere), comb. nov. (transferred from Campiglossa Rondani [= Paroxyna Hendel]). Previous Australian records o f Trupanea amoena (Frauenfeld) are referred to T. opprimata Hering and the introduced Procecidochares alani Steyskal is added to the Australian list.
Article
Full-text available
Information on distribution, plant association or taxonomy is provided for 82 species of fruit fflies from south and southeast Asia, including 13 new country records for Thailand and 5 for India.Ceratitella inthanona sp. n. is described from Thailand. Sundaresta malaisei (Hering) is transferred from Xyphosia and Perirhithrum longiseta (Hering) is transferred from Dictyotrypeta. Aciura kashmirica Zaka-ur-Rab is placed as a new synonym of Aciura afghana (Hering).
Trypetiden-Studien. Zoologische Jahrbücher. Abteilung fiir Systematik
  • G Enderlein
ENDERLEIN, G. 1911. Trypetiden-Studien. Zoologische Jahrbücher. Abteilung fiir Systematik, Oekologie und Geographie der Tierre 31: 407-460.
  • D L Hancock
  • A H Kirk-Spriggs
  • E Marais
HANCOCK, D.L., KIRK-SPRIGGS, A.H. and MARAIS, E. 2003. New records of Namibian Tephritidae (Diptera: Schizophora), with notes on the classification of subfamily Tephritinae. Cimbebasia 18: 49-70.
  • D E Hardy
HARDY, D.E. 1974. The fruit flies of the Philippines (Diptera: Tephritidae). Pacific Insects Monograph 32: 1-266.
The Tephritinae of Indonesia, New Guinea, the Bismarck and Solomon Islands (Diptera: Tephritidae)
  • D E Hardy
HARDY, D.E. 1988. The Tephritinae of Indonesia, New Guinea, the Bismarck and Solomon Islands (Diptera: Tephritidae). Bishop Museum Bulletin in Entomology 1: i-vii, 1-92.
Neue Gattungen und Arten palaearktischer und exotischer Fruchtfliegen
  • E M Hering
HERING, E.M. 1942. Neue Gattungen und Arten palaearktischer und exotischer Fruchtfliegen. Siruna Seva 4:1-31.
A systematic study o f Trypetidae in the Japanese Empire. Memoirs o f the Faculty o f Science and Agriculture
  • T Shiraki
SHIRAKI, T. 1933. A systematic study o f Trypetidae in the Japanese Empire. Memoirs o f the Faculty o f Science and Agriculture, Taihoku Imperial University 8(Entomology 2): 1-509.
The fruit flies (Diptera: Tephritidae) of the East Asian Region
  • X Wang
WANG, X.-j. 1998. The fruit flies (Diptera: Tephritidae) of the East Asian Region. Acta Zootaxonomica Sinica 21(Supplement 1996): i-viii, 1-378, pis 1-41.