A review of the gall midges (Diptera: Cecidomyiidae) of Indonesia: taxonomy, biology and adult key to genera

Abstract

Forty-seven species in 25 genera of gall midges (Diptera: Cecidomyiidae: Cecidomyiinae) known from Indonesia are reviewed. Available historic types were examined, and taxa are revised. Leefmansiella Kolesik & Gagné gen. nov. is erected for Trishormomyia pandani Felt. Actilasioptera falcaria (Felt) is found to be a senior synonym of Actilasioptera tumidifolium Gagné. Apoasphondylia orientalis Felt is transferred to Bruggmanniella Tavares, new combination; Asphondylia bursaria Felt is changed to Bruggmanniella, new combination; Contarinia eragrostidis Felt to Stenodiplosis Reuter, new combination; the Australian Contarinia brevipalpis Harris to Stenodiplosis, new combination; Dasyneura tetrastigma Felt to Gephyraulus Rübsaamen, new combination; and the Philippine Itonida paederiae Felt to Eucalyptodiplosis Kolesik, new combination. For each species, type data, a description, the basic biology and the known geographical distribution are given. A key to Cecidomyiidae genera based on adults, with references to described species, is provided. Species of economic importance, as pests of cultivated plants or biological control agents of weeds and pest arthropods, are listed.

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https://doi.org/10.11646/zootaxa.4847.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:1F8E3DED-6EA9-4D8A-8DA9-CD8C0CC9147F
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4847
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Magnolia Press
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Copyright © 2020 Magnolia Press Monograph
A review of the gall midges (Diptera: Cecidomyiidae) of Indonesia:
taxonomy, biology and adult key to genera
PETER KOLESIK1 & RAYMOND J. GAGNÉ2
1South Australian Museum, Department of Terrestrial Invertebrates, Adelaide SA 5000, Australia.
�
pkolesik@outlook.com.au; https://orcid.org/0000-0001-5569-6330
2Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, c/o Smithsonian Institution
MRC-168, P.O. Box 37012, Washington, DC20013-7012, USA.
�
raymond.gagne@ars.usda.gov; https://orcid.org/0000-0002-3464-757X

KOLESIK & GAGNÉ
2 · Zootaxa 4847 (1) © 2020 Magnolia Press
PETER KOLESIK & RAYMOND J. GAGNÉ
A review of the gall midges (Diptera: Cecidomyiidae) of Indonesia: taxonomy, biology and adult key to
genera
(Zootaxa 4847)
82 pp.; 30 cm.
7 Sept. 2020
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ISBN 978-1-77688-043-0 (Online edition)
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GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 3
Table of Contents
Abstract ...................................................................................................4
Introduction ................................................................................................4
Material and methods ........................................................................................5
Gall midges of Indonesia .....................................................................................8
Actilasioptera falcaria (Felt) ..................................................................................8
Actilasioptera sp. ..........................................................................................10
Asphondylia Loew in Indonesia ...............................................................................10
Asphondylia callicarpae Felt .................................................................................11
Asphondylia capsicicola Uechi, Yukawa & Tokuda ................................................................12
Asphondylia ixora Felt ......................................................................................13
Asphondylia leeae Felt ......................................................................................14
Asphondylia litseae Felt .....................................................................................16
Asphondylia strobilanthi Felt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Asphondylia vitea Felt ......................................................................................18
Asphondylia viticola Kieffer & Docters van Leeuwen-Reijnvaan .....................................................19
Asphondylia yushimai Yukawa & Uechi ........................................................................20
Asphoxenomyia Felt ........................................................................................21
Asphoxenomyia smilacis Felt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Bruggmanniella bursaria (Felt). New combination. ...............................................................23
Bruggmanniella orientalis (Felt). New combination. ..............................................................24
Dasineura elatostemmae Felt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Diadiplosis duni (Harris) ....................................................................................26
Diadiplosis smithi Felt ......................................................................................27
Feltiella acarivora (Zehntner) ................................................................................29
Gephyraulus tetrastigma (Felt). New combination ................................................................31
Gynodiplosis Felt ..........................................................................................32
Gynodiplosis humata Felt, 1927 ..............................................................................32
Hypodiplosis Kieffer ........................................................................................34
Hypodiplosis paederiae (Kieffer & Docters van Leeuwen-Reijnvaan) .................................................35
Lasioptera javanica Kieffer & Docters van Leeuwen-Reijnvaan .....................................................35
Lasioptera manilensis Felt ...................................................................................36
Leefmansiella Kolesik & Gagné. New Genus ....................................................................38
Leefmansiella pandani (Felt, 1921) ............................................................................39
Lestodiplosis lacciferi (Barnes) ...............................................................................40
Lestodiplosis oomeni Harris ..................................................................................42
Lestodiplosis syringopais (Hering) .............................................................................44
Megommata leefmansi (Nijveldt) .............................................................................45
Oligoxenomyia Felt ........................................................................................47
Oligoxenomyia radicis Felt ..................................................................................48
Orseolia Kieffer & Massalongo in Indonesia .....................................................................48
Orseolia apludae (Felt) ......................................................................................49
Orseolia caulicola Gagné ...................................................................................49
Orseolia graminicola (Kieffer & Docters van Leeuwen-Reijnvaan) ...................................................50
Orseolia graminis (Kieffer & Docters van Leeuwen-Reijnvaan) .....................................................51
Orseolia javanica Kieffer & Docters van Leeuwen-Reijnvaan .......................................................52
Orseolia orientalis (Felt) ....................................................................................53
Orseolia oryzae (Wood-Mason) ...............................................................................55
Orseolia paspali (Felt) ......................................................................................56
Procontarinia matteiana Kieffer & Cecconi .....................................................................57
Procontarinia robusta Li, Bu & Zhang ........................................................................59
Psephodiplosis rubi Kolesik ..................................................................................60
Schizomyia Kieffer in Indonesia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Schizomyia laporteae Felt ...................................................................................62
Schizomyia nodosa Felt .....................................................................................63
Schizomyia villebrunneae Felt ................................................................................65
Stenodiplosis eragrostidis Felt, new combination .................................................................66
Thorodiplosis Felt ..........................................................................................69
Thorodiplosis impatientis Felt ................................................................................69
Species unplaced to genus ...................................................................................71
“Calopedila” polyalthiae Felt ................................................................................71
“Gnesiodiplosis” garcinia Felt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

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4 · Zootaxa 4847 (1) © 2020 Magnolia Press
“Stefaniella” orientalis Felt ..................................................................................73
Addendum ...............................................................................................74
Eucalyptodiplosis paederiae (Felt). New combination. .............................................................74
Key to adults in genera of Cecidomyiidae currently known from Indonesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Acknowledgments ..........................................................................................78
References ................................................................................................78
Abstract
Forty-seven species in 25 genera of gall midges (Diptera: Cecidomyiidae: Cecidomyiinae) known from Indonesia are
reviewed. Available historic types were examined, and taxa are revised. Leefmansiella Kolesik & Gagné gen. nov. is
erected for Trishormomyia pandani Felt. Actilasioptera falcaria (Felt) is found to be a senior synonym of Actilasioptera
tumidifolium Gagné. Apoasphondylia orientalis Felt is transferred to Bruggmanniella Tavares, new combination;
Asphondylia bursaria Felt is changed to Bruggmanniella, new combination; Contarinia eragrostidis Felt to Stenodiplosis
Reuter, new combination; the Australian Contarinia brevipalpis Harris to Stenodiplosis, new combination; Dasyneura
tetrastigma Felt to Gephyraulus Rübsaamen, new combination; and the Philippine Itonida paederiae Felt to
Eucalyptodiplosis Kolesik, new combination. For each species, type data, a description, the basic biology and the known
geographical distribution are given. A key to Cecidomyiidae genera based on adults, with references to described species,
is provided. Species of economic importance, as pests of cultivated plants or biological control agents of weeds and pest
arthropods, are listed.
Key words: gall midges, Cecidomyiidae, Cecidomyiinae, fauna, Indonesia, crop pests, biological control, weed, insects,
mites
Introduction
Cecidomyiidae are a dipteran family of presently 6590 known species formed of six subfamilies: Catotrichinae,
Lestremiinae, Micromyinae, Winnertziinae, Porricondylinae and Cecidomyiinae (Gagné & Jaschhof 2017). Larvae
of Catotrichinae, Lestremiinae, Micromyinae, Winnertziinae and Porricondylinae feed on detritus or fungi and are
not treated here. Larvae of most Cecidomyiinae, by far the largest subfamily, feed on plants, some feed on fungi and
a certain number are predators of insects or arachnids. Many of the plant-feeders induce galls, hence the common
name “gall midges”. Galls are species-specific malformations that may be simple or complex and develop on buds,
flowers, fruit, leaves, stems and sometimes even roots. In simple terms they are tumors induced by plant growth
hormones produced and injected by the larva into the plant tissue. The gall develops around the larva which then
feeds on plant cells with its piercing-sucking apparatus. Most plant-feeding gall midges are monophagous or closely
oligophagous and some have one or two unrelated seasonal alternate hosts.
The subfamily contains many species of economic importance, as pests of cultivated plants or biological control
agents of weeds and insects. Agricultural and horticultural pest gall midges known from Indonesia include those
that attack plant reproductive organs, including Asphondylia capsicicola Uechi, Yukawa & Tokuda and Asphondylia
yushimai Yukawa & Uechi that deform fruit of chili peppers and soybeans, respectively, and Schizomyia nodosa
Felt that damages flowers of musk basil. The rice stem gall midge, Oseolia oryzae (Wood-Mason), causes a gall
that replaces the main flowering stem. Species that gall leaves, e.g. Procontarinia spp. on mango, may occur in
large enough numbers to cause a nutrient drain and, additionally, may instigate secondary infection by fungi. The
presence of other species impairs the attractiveness of plants grown as ornamentals for local and international trade.
These include Asphondylia callicarpae deforming leaves of beautyberry Callicarpa longifolia, Asphondylia ixora
malforming flowers of ixora Ixora timorensis and Thorodiplosis impatientis Felt deforming leaves of orange-flow-
ered impatiens Impatiens platypetala. Some species infest unwanted plants and may be considered biological weed
control agents: Orseolia javanica malforms leaves of cogon grass Imperata cylindrica, Orseolia graminis malforms
leaves of slender panic grass Ottochloa nodosa and Orseolia graminicola causes leaf malformation on Bermuda
grass Cynodon dactylon. Several predators of insects and mites are known whose beneficial nature is not yet fully
appreciated. These include Diadiplosis duni Harris, Diadiplosis smithi Felt, Lestodiplosis lacciferi (Barnes) and Me-
gommata leefmansi (Nijveldt) feeding on scale insects, Feltiella acarivora (Zehntner) on spider mites, and Lesto-
diplosis oomeni Harris on tea mites.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 5
Cecidomyiinae new to science are continually being found with the ultimate number of species currently ines-
timable, mainly due to the poor knowledge of tropical faunas. Our understanding of Indonesian Cecidomyiinae is
rather atypical in the sense that the galls of many species are known but the insects themselves remain undescribed
and unnamed. In 1926, the Dutch couple Jenny Docters van Leeuwen-Reijnvaan and Willem Marius Docters van
Leeuwen (Fig. 1) published “The Zoocecidia of the Netherlands East Indies”, a brilliant book of 600 pages contain-
ing some 1100 ink drawings of galls by the Javanese artist Raden Sastrasaputra. The book provides descriptions and
geographical information for over 1500 plant galls induced by insects, mites and nematodes occurring in Indonesia.
Of these, over 600 are caused by gall midges. The authors used mostly the larval stage to diagnose the gall-inducing
agent. In the case of Cecidomyiinae, larvae readily enable family-level identification. These are small, often less
than 3 mm long, legless, spindle form, white or bright coloured (red, pink, orange, yellow), with tiny, conical head
capsules. In addition, full-grown larvae usually possess on the first thoracic segment the sternal spatula - a hard,
fork-shaped dermal structure unique to gall midges (Figs 13c, 34d, 37c, 38d). Host plant identification to the species
level, exquisite illustrations of the galls and meticulously recorded data for geographical distribution make “The
Zoocecidia of the Netherlands East Indies” an invaluable reference to plant galls even now, more than 90 years after
its publication. Despite the wealth of information on the galls these insects cause, progress in the taxonomy of In-
donesian Cecidomyiinae has been slow and only 47 species were recorded from the Indonesian archipelago prior to
this review. Thirty-nine species were described before 1936, the remaining eight after 1954, by which time descrip-
tion standards had greatly improved. Jean-Jacques Kieffer with silent partners Giacomo Cecconi and Jenny Docters
van Leeuwen-Reijnvaan in 1906 and 1910, respectively, described a total of seven Indonesian species whose types
are lost. Ephraim Porter Felt described 29 species between 1915 and 1927. Types of nearly all of his Indonesian
species survive, but most were slide-mounted directly into Canada balsam without benefit of first clearing them. We
remounted a few slides to practical effect, but in doing so one needs to weigh perceived benefits against consequent
loss of parts, especially of antennae and legs, in the process.
FIGURE 1. Authors of “The Zoocecidia of the Netherlands East Indies” published in 1926. Left: Jenny Docters van Leeuwen-
Reijnvaan (1880–1963) in 1907. Middle: Willem Marius Docters van Leeuwen (1880–1960) in 1926. Right: Mt Gede, collec-
tion locality of many Indonesian Cecidomyiidae, 1871 oil painting “River Valley in Java with Mt. Gede and Mt. Pangrango in
the Background” by Raden Saleh (Collection of the Tan Family, National Gallery Singapore).
Our purpose here was to review the currently known Indonesian Cecidomyiinae as a basis for further work,
taxonomic to be sure, but also to help ecologists and plant protection workers in their identification and biological
assessments. Notes on the systematics of these taxa are given, many with redescriptions and copious illustrations.
Little can be done now about missing or partial types but the best thing we can say for the named Indonesian species
is that the hosts and niches of all of them are known. This means that they can all be found again and redescribed in
a way better than we have been able to because of our lack of perfect specimens and all stages of most species. Re-
cently, Javanese gall midges have been studied by Purnama Hidayat and his students at the IPB University, Bogor,
and we are thankful to him for providing biological observations and photographs of some of the species. Indonesia
is a large tropical country with rich biodiversity, vibrant agriculture, extensive forest industry and advancing bio-
logical research, and we expect that many gall midges treated here will attract interest in years to come.
Material and methods
We examined and redescribe here the available extant types of Indonesian gall midge taxa. If we deemed the original

KOLESIK & GAGNÉ
6 · Zootaxa 4847 (1) © 2020 Magnolia Press
description satisfactory, we refer to the original paper or alternatively, recite the major morphological characters,
including figures, mainly from older papers that might not be easily available to some readers. This review includes
the taxa known to us through December 31, 2019. The greater part of the Indonesian gall midge species was de-
scribed by E.P. Felt but were almost all mounted directly in Canada balsam on slides without benefit of initial clear-
ing. In 1917, when Felt received these specimens from Docters van Leeuwen, he no longer enjoyed the services
of technicians who had previously served him very well with effectively cleared and dressed specimens on slide
mounts in Canada balsam (Gagné 1994). Some specimens were cleared and remounted by us in Canada balsam but
we decided against this in most cases because the more fragile parts such as antennae and legs are too often lost in
the process. The proper way to mount gall midges in that medium to ensure good clearing and permanence is out-
lined in Gagné (1994). Some specimens we examined were mounted by more recent authors in Hoyer’s medium.
Although we could observe what we needed to see on these particular specimens, the medium is gradually crystal-
lizing and the specimens will eventually become unrecognizable. Hoyer’s medium is not permanent, even when
ringed as these specimens are, so should never be used for type series. We have used Hoyer’s medium to good effect
in the one-step clearing and mounting of first instar larvae that easily become lost in the mounting process but these
can always be remounted later if need be.
Type material of taxa we reviewed are lodged in the following institutions.
BMNH Natural History Museum, London, UK.
NYSM New York State Museum, Albany, New York, USA. Types of all of the Indonesian taxa described
by Felt (1915–1927) are the property of the NYSM in Albany and are on indefinite loan to the
National Museum of Natural History (USNM).
RMNH Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands.
USNM National Museum of Natural History, Smithsonian Institution, Washington DC, USA. A few of the
Felt specimens from elsewhere than Indonesia that we studied are the property of the USNM. Al-
though they are Felt types, these were initially provided by the USNM to Felt for study and descrip-
tion and subsequently remitted to the USNM where they were given USNM specimen numbers.
Because of the length of the names for two often-cited authors, (Willem) Docters van Leeuwen and (Jenny)
Docters van Leeuwen-Reijnvaan, we cite them in the text as DvL and DvLR. We have also abbreviated text cita-
tions for Docters van Leewen-Reinvaan, Docters van Leeuwen, and Docters van Leewen-Reinvaan & Docters van
Leeuwen as DvLR, DvL, and DvLR & DvL, respectively. Drawings were made using a compound microscope with
the aid of a drawing tube, length of body parts was measured with a microscope imaging system. Figure numbers
in round brackets refer to figures in the original papers, square brackets are used for figures presented in this paper.
Figures reproduced from the original papers are acknowledged in the text, in the figure captions and the Acknowl-
edgements. Our comments within a text cited from the original papers are placed in square brackets. We use the geo-
graphical names in their present forms which, in the instances below, differ from those used in the original papers.
Name used in the original publication Present name
Amsterdam island, near Batavia, Java Untung Jawa island, near Jakarta, Java,
Mt Andoeng, Java Mt Andong, Java
Bandarbaroe, near Medan, Sumatra Bandar Baru, near Medan, Sumatra
Bandoeng, Java Bandung, Java
Batavia, Java Jakarta, Java
Buitenzorg, Java Bogor, Java
Dépok, Java Depok, Java
Djerakah, near Samarang, Java Jrakah, near Semarang, Java
Garoet, Java Garut, Java
Mt Gedé (or Gedeh), Tjibodas, Java Mt Gede, Cibodas, Java
Mt Gede, Tjibeureum, Java Mt Gede, Cibeureum, Java
Mt Gegerbintang, Java Mt Putri, near Bandung, Java
Island Kajoeadi, Salajar group Kayuadi Island, Selayar archipelago
Kamal, Madoera Island Kamal, Madura Island

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 7
Kamodjan, near Garoet, Java Kamojang, near Garut, Java
Mt Keloed, Java Mt Kelud, Java
Keravat, New Britain, Papua New Guinea Kerevat, New Britain, Papua New Guinea
Island Klein-Kombius, north of Batavia Bokor island, north of Jakarta
Koeripan, Pekalongan, Java Kuripan, near Pekalongan, Java
Desa Kubang, Kec Guguk, West Sumatra Kubang, Guguak district, West Sumatra
Kurruckpore, Monghyr, India Kurruckpore, Munger, Bihar, India
Mt Maquling, Luzon, the Philippines Mt Makiling, Luzon, the Philippines
Mt Mandalagiri, near Garoet, Java Mt Mandalagiri, near Garut, Java
Mt Manglatang, near Bandoeng, Java Mt Manglayang, near Bandung, Java
Mt Moerja, Java Mt Muria, Java
Ngarengan, near Samarang, Java Ngarengan, near Semarang, Java
Oengaran Gebirge, Java Mt Ungaran, Java
Mt Oengaran, Java Mt Ungaran, Java
Mt Pantjar, near Buitenzorg, Java Mt Pancar, near Bogor, Java
Paree (or Paré), near Kediri, Java Pare, near Kediri, Java
Pase, near Bandung, Java Paseh, near Bandung, Java
Pasir Nanka, Indonesia Pasir Nangka, Cianjur Regency, Java
Pengalengan, near Bandoeng, Java Pangalengan, near Bandung, Java
Pesewahan, south of Tjibadak, Java Pesewahan, south of Cibadak, near Bogor, Java
Plaboean, near Weliri, Java Plabuan, near Weleri, Java
Mt Ratè, Wai Lima Telokbetong, Sumatra Mt Rate, Way Lima, Telokbetung, Sumatra
Rimboepengadang, Sumatra Rimbo Pengadang, Sumatra
Sajoong, near Semarang, Java Sayung, near Semarang, Java
Mt Salak, Goenong Boender, Java Mt Salak, Gunung Bunder, Java
Samarang, Java Semarang, Java
Seketjer, near Weliri, Java Sekecer, near Weliri, Java
Mt Slamat, Java Mt Slamet, Java
Soekamangli, near Weliri, Java Sukamangli, near Weleri, Java
Soember Pitoe, near Weliri, Java Sumber Pitu, near Weleri, Java
Tambakan, Tankuban Prahu, Indonesia Tambakan, Tangkuban Perahu, Java
Mt Tangkoeban Prahoe, near Bandoeng Mt Tangkuban Perahu, near Bandung, Java
Tegal Waroe, near Krawang Tegalwaru, near Krawang
Tjadas, Melang, near Tjibeber, Java Cadas, Malang, near Cibeber, Java
Mt Tjampea, near Buitenzorg, Java Mt Ciampea, near Bogor, Java
Tjandi, near Samarang, Java Candi, near Semarang, Java
Tjianten, near Buitenzorg Cianten, near Bogor
Tjiastana, near Tjisalak, Mt Halimoen, Java Ciastana, near Cisalak, Mt Halimun, Java
Mt Tjibodas, Tjampea, near Bogor Mt Cibodas, Ciampea, near Bogor
Tjigombong, near Buitenzorg Cigombong, near Bogor
Tjiharoem, near Tjibeber, Java Ciharum, near Cibeber, Java
Tjikadongdong, near Bandoeng, Java Cikadongdong, near Bandung, Java
Tjikampek, Java Cikampek, Java
Mt Tjikorai, Java Mt Cikuray, Java
Tjinjiroean, near Bandoeng Cinyiruan, near Bandung
Tjipanas, near Tjisolok, Java Cipanas, near Cisolok, Java
Tjisaroewa, near Tjimahi, Java Cisarua, near Cimahi, Java
Tjisokan, near Tjibeber, Java Cisokan, near Cibeber, Java
Mt Toegoe, near Tjibeber, Java Mt Tugu, near Cibeber, Java
Wanajasa, near Mt Boerangrang, Java Wanayasa, near Purwakarta, Java
Wanajasa, near Poerwakarta, Java Wanayasa, near Purwakarta, Java
Mt Welirang, Trètes, Java Mt Welirang, Java

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8 · Zootaxa 4847 (1) © 2020 Magnolia Press
Weltevreden, Java Jakarta, Java
Weltevreden, Pegangsaän, Java Pegangsaan, Jakarta, Java
Currently, genera from two subfamilies of Cecidomyiidae are reported from Indonesia: Porricondylinae (spe-
cies Colomyia cortici) that is not treated here (while is included in the key) and Cecidomyiinae (the remaining
species). Treated taxa are listed in alphabetical order, starting with genera and species placed to genus followed by
species currently unplaced to genera.
Gall midges of Indonesia
Actilasioptera falcaria (Felt)
[Figs 2a–j]
Stefaniella falcaria Felt, 1921b: 141 (in part); Gagné (in Gagné & Law 1999): 26, new combination.
Actilasioptera tumidifolium Gagné (in Gagné & Law 1999): 25. New junior synonym.
Material examined of types of names under this taxon. Actilasioptera falcaria (Felt): lectotype, female, desig-
nated by Gagné & Law (1999), and three female and one male paralectotypes, from “Avicennia officinalis L” [cor-
rected to Avicennia marina (Forssk.) Vierh. var. intermedia (Griff.) Bakh.], Semarang, Java, Indonesia, 27-iv-1914,
W. Docters van Leeuwen, Felt #a3089. The lectotype female, cleared and remounted by RJG in 1970, has the thorax
and abdomen mounted laterally, the head frontally, and all body parts well-preserved except for missing wings. One
female paralectotype, cleared and remounted by RJG in 1970, has the head mounted frontally, thorax and abdomen
with ovipositor mounted laterally and all body parts-well preserved except for missing antennae and wings. The
two remaining female paralectotypes are mounted laterally, whole and uncleared, both with the ovipositor retracted
inside the abdomen and not visible, one missing both wings, the other with one damaged wing. The male lectotype
is mounted laterally, whole and uncleared, with all antennal segments present but shrunken and both wings folded.
Actilasioptera tumidifolium Gagné (in Gagné & Law 1999): holotype, male, from leaf gall on Avicennia marina,
Sandgate, Queensland, Australia, 23-VIII-1996, L. Law, deposited in Australian National Collection, Canberra
(ANIC).
Description. The male, female, pupa and larva were described and illustrated in Gagné & Law (1999) as A.
tumidifolium. Illustrated here in Fig. 2e–j are male and female structures of Felt’s series of A. falcaria to validate our
synonymy of the two names. See elsewhere in Gagné & Law (1999) for descriptions and key to all six known species
of Actilasioptera and their galls.
Remarks. In the light of the correction by DvL (1921) of the host’s name from Avicennia officinalis to Av.
marina and the lack of apparent differences between adults of the two species, we consider A. falcaria and A. tu-
midifolium to be synonymous. Felt (1921b) did not describe the immature stages, and differences could possibly still
be found to separate these two species, but there is no reason to consider them as distinct at this time. Part of Felt’s
(1921b) original type series is a single paralectotype female that does not correspond to A. falcaria that causes the
large leaf gall and is treated below as Actilasioptera sp.
Biology. This species causes large irregular leaf galls on Avicennia marina (Acanthaceae). According to DvLR
& DvL (1910a) the gall (No. 96, Figs 51a, b [Fig. 2a]) is a 1 cm large, irregular swelling situated on or very close to
the leaf midrib, equally developed on both sides of the leaf, the upper side yellowish green and shiny, the underside
grey and matte. The affected leaf blade is undeveloped, with ruffled edges. The gall contains several chambers, each
with a single white larva. The host plant name was originally given as “Avicennia officinalis L.” (DvLR & DvL
1910a; Felt 1921b), but DvL (1921) corrected it to Avicennia marina (Forssk.) Vierh. var. intermedia (Griff.) Bakh.,
reiterated the basic gall morphology, and provided a detailed illustration of the gall in his Fig. 2 [Fig. 2b]. Additional
illustrations and description of the gall were given by DvLR & DvL (1926, gall No. 1283 Fig. 921 [Fig. 2c]), with
the host plant name in agreement with the 1921 correction, A. marina var intermedia.
Geographical distribution. This species is known from Indonesia (DvLR & DvL 1926; Felt 1921b) and Aus-
tralia (Gagné & Law 1999) from the following localities: Semarang, Java, ix.1909, 29-iv-1914; Jrakah, near Sema-
rang, Java, iii.1912, iii.1913; Untung Jawa island, near Jakarta, Java, iv.1919; Sebesi island, iv.1921; Sandgate,
Queensland, 23-viii-1996; Donnybrook, Queensland, 16-iv-1994; Toorbul, Queensland, 20-iii-1994; Brighton,
Queensland, 4-ix-1995; and Saunders Beach, Townsville, Queensland, 8-vii-1995. Avicennia marina occurs also
along the coast and on islands of east Africa, south-west, south and south-east Asia, Australia, New Guinea, Solo-

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 9
mon Islands, New Caledonia and north of New Zealand (Duke 1991), so Actilasioptera falcaria may have a wider
distribution than currently known.
FIGURES 2a–j. Actilasioptera falcaria: a–d, galls on Avicennia marina: a, leaf upper side on top, gall cross section on bottom
(Figs 51a, b of DvLR & DvL 1910); b (Fig. 2 of DvL 1921); c (Fig 921 of DvLR & DvL (1926)); d, leaf underside (Fig. 2 of
Gagné & Law 1999); e–f, male: e, tarsal claw with empodium and pulvillus; f, terminalia (somewhat misshapen in the prepara-
tion), from left to right: cerci (ce), mesobasal lobe (ml), aedeagus (ae), hypoproct (hy), gonocoxite (gc), gonostylus (gs); g–k,
female: g, end of abdomen (lateral); h, palpi; i, sixth flagellomere (ventral); j, same (dorsal). Figs 2k-n. Actilasioptera sp.: k,
palpi; l, wing; m, galls on Avicennia marina, drawn to same scale as Fig. 2a, leaf underside on left, upper side on right (Fig. 3
of DvL (1921); n, gall cross section (Fig. 51c of DvLR & DvL 1910).

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Actilasioptera sp.
[Figs 2k–n]
Stefaniella falcaria Felt, 1921b: 141 (in part).
Material examined. Female paralectotype of Actilasioptera falcaria (Felt), designated by Gagné (in Gagné &
Law 1999), “Avicennia officinalis L.” [corrected to Avicennia marina (Forssk.) Vierh. var intermedia (Griff.) Bakh.
by DvL (1921)], Semarang, Java, Indonesia, 29-iv-1914, W. Docters van Leeuwen, Felt #a3090. The specimen is
mounted whole, uncleared, with head and thorax mounted laterally, the abdomen dorsoventrally, the ovipositor
retracted inside the abdomen.
Description. Female. Wing 1.7 mm long, 0.8 mm wide; R5 two-thirds wing length, straight. Antennae with 12
flagellomeres, as in A. falcaria. Palpus [Fig. 2k] 2-segmented; first segment twice as long as wide, constricted at
basal third, with several setae; second segment one-third width and length of first, with single terminal seta; palpi-
ger absent. Ovipositor retracted inside abdomen, so extent of its dorsoventral curvature, a character differentiating
Actilasioptera spp. (Gagné & Law 1999), is not visible. Tarsal claws, empodia, pulvilli as in A. falcaria.
Male, pupa, larva unknown.
Remarks. This specimen was reared from the same host as Actilasioptera falcaria but from a different leaf
gall. While the gall of A. falcaria is a large, irregular swelling [Figs 2a–d], the one associated with this female is a
small, round pustule [Figs 2m, n]. The female differs from that of A. falcaria in the size of the second palpal seg-
ment that in this species is substantially smaller than the first and bears a single seta [Fig. 2k], while in A. falcaria it
is only slightly smaller than the second and bears several setae [Fig. 2h]. At the time of the description, DvL (1921)
believed that the two gall types were caused by two different species but Felt (1921b) wrote that with little question
this female was referable to A. falcaria. The palpus of the specimen differs also from that of the remaining conge-
ners in which the second palpal segment is larger than the first and bears several setae (see Gagné & Law 1999).
We did not name this segregate in as much as the available female is in poor condition and the male, pupa and larva
are lacking.
Biology. This species causes small pustule-like leaf galls on Avicennia marina (Acanthaceae). According to
DvLR & DvL (1910a, gall No. 97, Fig. 51c [Fig. 2n]) and DvLR & DvL (1926, gall No. 1284) the gall is a round leaf
pustule, 3 mm wide and 1 mm or less thick, yellowish green on the upper side, yellow on the underside, containing
two larval chambers. Following adult emergence, the pupal exuviae remain wedged in the exit holes on the under-
side of the gall. While the host was given as “Avicennia officinalis L.” in DvLR & DvL (1910a) and Felt (1921b),
DvL (1921) corrected the species to Avicennia marina (Forssk.) Vierh. var. intermedia (Griff.) Bakh. and provided
a new, detailed illustration of the gall in Fig. 3 [Fig. 2m].
Geographical distribution. This segregate is currently known only from Semarang, Java where it was col-
lected 29-iv-1914 (Felt 1921b) and also from Jrakah, near Semarang, Java, iii.1912; Brebes, Java, i-1919; Untung
Jawa island, near Jakarta, Java, iv.1919; Sebesi island, iv.1921 (DvLR & DvL 1926).
Asphondylia Loew in Indonesia
Nine species of this genus are so far known from Indonesia. Two were initially well-described from Japan, but the
types of four of the remaining species are in poor condition and those of the other three are lost. The taxonomy of
this group depends equally on males, females, pupae and larvae, but three of the species here lack one or the other
sex, two the pupa and six the larva. Nonetheless, the variety manifested by the few Asphondylia species known from
Indonesia is intriguing, if only from what we can appreciate of the tarsal claws, whether thick or thin, some lobed,
others anisomorphic within a species or sex. Interestingly, most of the known pupae share a triplet of lower frontal
horns that form an equilateral triangle. We have refrained from making further comparisons among species because
available specimens are not fully comparable.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 11
Asphondylia callicarpae Felt
[Figs 3a–o]
Asphondylia callicarpae Felt, 1918: 285.
Material examined. Syntypes, male, 3 females, and a pupal exuviae [fragment], Felt #a2843, reared from leaf
galls on Callicarpa erioclona Schauer (Verbenaceae [now Lamiaceae]), Mt Makiling, Laguna Province, Luzon,
the Philippines, collected at altitude 200–300 m viii-1917 and reared by L.B. Uichanco 3-ix-1917. One slide bears
a male missing an antenna, another slide with a complete female except with its head inside an anterior fragment
of a pupal exuviae, and the two other females are still pinned and well-preserved. Three years after the original de-
scription, Felt (1921d) described one female and two pupae reared by DvL from a leaf gall on Callicarpa longifolia
Lam. collected at Mt Ungaran, Java, iv-1914, and concluded that they belonged to this species. We examined those
specimens also: an uncleared female and pupa and a pupal exuviae, are on a single slide (Felt #a3102).
Description. Male. Wing length 2.3 mm, width 1.0 mm. Antenna: scape 2x longer than wide, pedicel slightly
wider than long, flagellomeres slightly progressively shorter [Fig. 3g]. Palpus 3-segmented, division between sec-
ond and third may be indistinct, segments progressively longer [Fig. 3d]. First tarsal segment with apicoventral spur
short, robust, curved at basal third [Fig. 3e]. Tarsal claws robust, curved at midlength, empodia as long as claws,
pulvilli minute [Fig. 3f]. Gonostylus as wide as long in posterior view, one tooth long and pointed, one short and
blunt [Fig. 3h].
Female. Wing length 2.6 mm, width 1.0 mm, vein R5 joining C at wing apex, R1 2/3 wing length [Fig. 3j].
Flagellomeres 1–9 progressively slightly shorter, 10–12 progressively much shorter, first flagellomere 5x longer
than wide [Fig. 3k]. Palpus 3-segmented, segments progressively longer [Fig. 3i]. Needle-like protrusible part of
ovipositor 2.4x longer than seventh sternite [Fig. 3l].
Pupa. The anterior end of the pupal exuviae fragment collected in 1917 in the Philippines and mounted by Felt
has large antennal horns with finely serrated inner edges, one large anterior frontal horn and three smaller posterior
frontal horns placed in the shape of an isosceles triangle, the posteriormost, situated adjacent to the labella, recurved
and rugose [Fig. 3m].
Larva unknown.
Remarks. The pupa has serrated inner edges of the antennal horns, and the posteriormost of the three lower
frontal horns is somewhat removed from the other two and uniquely rugose. We note that the frontal horns of the
Javan specimens [Figs 3m, n] are noticeably larger than those seen on the Philippine pupa [Fig. 3m]. We defer to
future study, possibly until larvae can be compared, whether that difference is a specific one.
Biology. Uichanco (1919) reared the type specimens in the Philippines from leaf galls on Callicarpa erioclona
Schauer (gall No. 18147, Plate X Fig. 2 [Fig. 3a]) and described the gall as about 30 mm, width 15 mm, thickness 16
mm, polythalamous, consisting of an enlargement of the midrib and forming with the atrophied leaf lamina a single
mass of succulent tissue. The enlarged portion is tomentose, concave above and convex on the nether surface. The
petiole, usually the leaf apex, and a small portion at the base are sometimes unaffected. The hair covering is long,
dark brown and concolorous with the normal short pubescence of the plant. Larval chambers are subellipsoid, their
size variable and arranged irregularly in close proximity to the upper, concave surface. The deformation is confined
to younger, subterminal leaves. Galls were abundant at the type locality and present most of the year (Uichanco
1919). DvL (1921, Fig. 8 [Fig. 3c]) and DvLR & DvL (1926, gall No. 1291, Fig. 927 [Fig. 3b]) reported a similar
gall on Callicarpa longifolia from several localities in Indonesia. The insects reared from the Indonesian galls were
examined by Felt who considered them conspecific with the insects from the Philippine galls (Felt 1921b; DvLR
& DvL 1926).
Geographical distribution. This species is known from its type locality at Mt Makiling, Laguna Province,
Luzon, in the Philippines where it was reared from Callicarpa erioclona viii-1917 (Felt 1918) and from several
localities in Indonesia (DvLR & DvL 1926) where the galls were found on Callicarpa longifolia on Java at Mt
Muria, alt. 500 m, x-1912; Ciharum, near Cibeber,, alt. 1000 m, vi-1916; Cadas Malang, near Cibeber, alt. 1000 m,
iv-1917; Mt Gede, Cibodas, alt. 1500 m, ix-1918; Pesewahan, south of Cibadak, near Bogor, alt. 600 m, viii-1918;
Mt Ungaran, alt. 1200 m, xi-1919; Wanayasa, near Purwakarta, alt. 1000 m, vii-1920 and in Sumatra at Rimbo
Pengadang vi-1916.

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FIGURES 3a–o. Asphondylia callicarpae: a, gall on Callicarpa erioclona from the Philippines (Fig. X-2 of Uichanco (1919));
b, gall on Callicarpa longifolia from Java (Fig. 927 of DvLR & DvL (1926)); c, gall on Callicarpa longifolia from Java (Fig. 8
of DvL 1921); d, male palpi; e, male first tarsal segment; f, male last tarsal segment with claw, empodium and pulvillus; g, male
antenna; h, gonostylus in posterodorsal view; i, female palpi; j, female wing; k, female antenna (within pupal exuviae); l, end of
ovipositor in ventral view; m–o, anterior part of pupa: m, o, ventral; n, lateral. Insect drawings d–m are based on types reared
from C. erioclona in the Philippines, n–o from C. longifolia in Java.
Asphondylia capsicicola Uechi, Yukawa & Tokuda
[Fig. 4a]
Asphondylia capsicicola Uechi, Yukawa & Tokuda, 2016 in Uechi et al. (2016: 4).
Type series. Holotype male (slide No CPS001), paratypes (male, 7 females, pupa, 3 pupal exuviae, 2 larvae,
CPS002–CP0015) and other slide-mounted and ethanol-stored specimens are deposited in the collection of the
Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan and in the collection of the
[Bogor] Zoological Museum, Cibinong, Indonesia (Uechi et al. 2016). We did not find it necessary to examine the
types of this species. Uechi et al. (2016) described the morphology and infestation symptoms, and sequenced COI
mitochondrial gene fragment (GenBank accession numbers LC164724–LC164733, LC167140–LC167146).
Morphology. Both sexes, pupae and larvae of this species are described in Uechi et al. (2016). The authors
compare A. capsicicola to other Asphondylia occurring on Solanum elsewhere in the world.
DNA. The COI sequence of A. capsicicola (Uechi et al. 2016) shows 0.91–2.52 % intraspecific divergence and

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 13
>13.62 % divergence to A. yushimai, the closest species with published sequence (GenBank, accessed 14-xii-2019,
>95% query cover used).
Biology and geographical distribution. Asphondylia capsicicola causes deformation of the fruit pods of culti-
vated chili peppers, Capsicum annuum L. and Capsicum frutescens L. (Solanaceae), in Indonesia and Vietnam (Uechi
et al. 2016, Fig. 1 [Fig. 4a]). Usually one, occasionally two or more larvae live in the gall (Busniah 2014 [as A. capsici],
Uechi et al. 2016). The life cycle lasts 25 days (Busniah 2014), suggesting a possibility of more than one generation
per year. In Indonesia, A. capsicicola has been found at the following localities: Cisarua, near Bogor, Java, alt. 1000
m, on C. annuum (16-iii-2004, 14-i-2016) and C. frutescens (16-iii-2003, 16-iii-2004, 29-xii-2015, 5 & 14-i-2016);
Gianyar, Bali on C. annuum (18-iii-2004); Kubang, Guguak district, West Sumatra on C. annuum (1-x-2001) (Uechi
et al. 2016). Plant infestation by A. capsicicola in Bogor area, Indonesia, reached 41% in 2005, suggesting that a
potential outbreak of this species could cause severe crop damage (Maryana et al. (2006), as Asphondylia sp.).
FIGURE 4a. Asphondylia capsicicola: a, gall on Capsicum sp. in South East Asia (Fig. 1 of Uechi et al. (2016). Figs 4b–c.
Asphondylia viticola: b, young gall on Causonis trifolia (Fig. 48 of DvLR 1909b); c, mature gall on C. trifolia (Fig. 637 of
DvLR & DvL (1926)). Figs 4d–e. Asphondylia yushimai (photos by Atsushi Kikuchi, Hiroshima, Japan, October 2007): d,
pupal exuviae in infested pod of Glycine max; e, damage of pod of G. max.
Asphondylia ixora Felt
[Figs 5a–g]
Asphondylia ixora Felt, 1927b: 388.
Material examined. Syntypes, 2 females reared by DvL 20-v-1923 from flower galls on Ixora timorensis Decne.
(Rubiaceae), Bokor Island, north of Jakarta, Felt #a3389), Both are in fair condition on a single slide, but their ovi-
positors are retracted inside the abdomen.
Description. Female. Wing length 2.6 mm (2.4–2.8), width 0.9 mm (0.8–1.0). Flagellomeres 1–9 progressively
slightly shorter, 10–12 progressively much shorter, first flagellomere 6x longer than wide [Fig. 5c]. Palpus 3-seg-
mented, segments progressively longer [Fig. 5b]. First tarsal segment with apicoventral spur slightly curved [Fig.
5d]. Tarsal claws strongly curved, anisomorphic, thinnest on forelegs, stout and widened at midlength on mid and
hindlegs, empodia about as long as claws [Figs 5e–g]. Needle-like protrusible part of ovipositor 2.0x longer than
seventh sternite.
Male, pupa, larva unknown.
Remarks. The anisomorphic tarsal claws are unique among known Indonesian Asphondylia.
Biology. Asphondylia ixora causes the flower gall on Ixora timorensis described and illustrated by DvLR &
DvL (1926, gall No. 1433 Fig. 1016 [Fig. 5a]). The ovary is swollen into a sphere about 4 mm across, the surface is
green and glossy, and inside is a spacious larval chamber surrounded by a succulent wall.
Geographical distribution. This species is known only from the type locality on Bokor Island, north of Ja-
karta, where it was collected 20-v-1923.

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FIGURES 5a–g. Asphondylia ixora: a: gall on Ixora timorensis (Fig. 1016 of DvLR & DvL (1926); arrows indicate gall (ga)
and normally developed ovary (ov); b-g, female: b, palpus; c, antenna; d, first tarsal segment; e, last tarsal segment of foreleg
with claw and empodium; f, same of midleg; g, same of hindleg.
Asphondylia leeae Felt
[Figs 6a–m]
Asphondylia leeae Felt, 1921b: 148.
Material examined. Syntypes, male, 2 females, and 2 pupae reared from fruit galls on Leea sambucina (L.)
Willd. [now junior synonym of Leea indica (Burm. f.) Merr.] (Vitaceae), collected in Bogor, Java, 24-ix-1918, Felt
#a3097). One of Felt’s two slides bearing this series contains whole mounts of one male and two females, the other,
two pupae. The male has a well-preserved abdomen, shriveled wings, all flagellomeres missing except the basal
three of one antenna, and most legs shriveled; one female has one antenna preserved, shriveled wings, the tip of the
needle-like protrusible part of ovipositor broken off, and most legs well-preserved, the other has a well-preserved
ovipositor, shriveled wings, and is missing all flagellomeres; one pupa is mounted laterally, containing a female
insect inside, the other is mounted dorsoventrally and also contains a female inside. The date written on both of
Felt’s slides is “24/IX’14” which is presumably an error because in the 1921b paper he states “24/9, ‘18”, which is
consistent with the date in DvLR & DvL (1926).
Description. Male. Antenna: first flagellomere 4x longer than wide [Fig. 6e]. Palpus 3-segmented, segments
progressively longer [Fig. 6d). Apicoventral spur on first tarsal segment robust [Fig. 6b]. Tarsal claws robust, strong-
ly curved at distal third, empodia longer than claws, pulvilli about ¼ empodial length [Fig. 6c]. Gonostylus round in
posterior view, teeth of equal length, one narrow and curved, one wide and straight [Fig. 6f].

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 15
FIGURES 6a–m. Asphondylia leeae: a, gall on Leea indica, two normal fruit on left, gall on right (Fig. 203 of DvLR & DvL
1914); b–f, male: b, first tarsal segment; c, last tarsal segment with claw and empodium; d, palpus; e, first three flagellomeres;
f, gonostylus (posterior); g–i, female: g, posterior end of abdomen (ventral); h, end of needle-like protrusible part of ovipositor
(ventral); i, antenna, 6th flagellomere shriveled; j–m, pupa: j, head (ventral); k, head (ventrolateral); l, last abdominal segments
(ventrolateral); m, prothoracic spiracle. Fig. 6n. Asphondylia litseae: n, pupal antennal horns (dorsal, Fig. 5 of Felt 1921b).
Female. Flagellomeres 1–9 progressively slightly shorter, 10–12 progressively much shorter, first flagellomere
4.5x longer than wide [Fig. 6i]. Needle-like protrusible part of ovipositor about 2x longer than seventh sternite, not
widened at terminus in ventral view [Fig. 6g–h],
Pupa. Antennal horns long, narrow, smooth [Fig. 6j]. Anterior frontal horn robust, sharply pointed [Fig. 6j,
k]. Three posterior frontal horns, small, sharply pointed, subequal in size, the two upper posterior horns slightly
closer to each other than to lower horn [Fig. 6j, k]. Cephalic papillar setae about 1/4 length antennal horns [Fig. 6j].
Prothoracic spiracle short, gradually tapering, rugged at distal third, trachea bulging, ending at basal 2/5 [Fig. 6m].
Abdominal tergal spines simple, gradually larger from anterior to posterior rows [Fig. 6l].
Larva unknown.
Remarks. A fruit gall on Leea sambucina Willd. in Gabon, Africa was identified by Houard (1923) as caused
by A. leeae, but his accompanying figures 1107 and 1108 show irregular, lumpy galls that appear different from
the regular, smooth gall of Felt’s type series from Indonesia [Fig. 6a]. Resolving this identification should depend

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16 · Zootaxa 4847 (1) © 2020 Magnolia Press
on comparison between the gallmakers, so we are not including Gabon in the geographical distribution range of A.
leeae. In Indonesia, undescribed gall midges were found feeding on Leea aequata L. and L. indica. These cause
flower galls that are different from the fruit gall of Asphondylia leeae in their irregular ball-shape, swollen calyx and
corolla with still visible lobes and forming a cup-like structure (DvLR & DvL 1926, Nos 861, 862).
Biology. This species causes a fruit gall on Leea indica. DvLR & DvL (1914) describe and illustrate the gall
(No. 443, Fig. 203 [Fig. 6a]). Normal fruit are depressed-spheroidal, about 10 mm wide and 7 mm high while galled
fruits are substantially larger, pear-shaped to round, 15 mm wide and 20 mm long, the surface smooth and grey-
green. The inside is composed of parenchymatic tissue and an irregular larval chamber in the center.
Geographical distribution. This species is known from Java, where it was found at Pare, near Kediri, alt. 200
m, v-1912, Bogor, alt. 250 m, 24-ix-1918 and Depok, alt. 100 m, viii-1918 (DvLR & DvL 1914; Felt 1921b; DvLR
& DvL 1926).
Asphondylia litseae Felt
[Fig. 6n]
Asphondylia litseae Felt, 1921b: 149.
Type series. Syntypes, female and 2 pupae, reared from stem galls on Litsea sp. (Lauraceae) collected in Ungaran
Mts, Java, alt. 1400 m, 21-iv-1914, Felt #a3096). This series could not be found in the Felt slide collection.
Description based on Felt (1921b), some superficial characters omitted. Female. Length 2.5 mm. Flagello-
meres 12 in number, fifth 2.5x longer than wide, terminal reduced, globose. Circumfila coarse, bowed, resembling
Schizomyia. Palpus 2-segmented, first segment subquadrate, second broadly oval, acute apically. Tarsal claws stout,
strongly curved, simple; empodia about 2/3 claw length. Pseudocerci at base of eversible part of ovipositor rudi-
mentary.
Pupa (based on Felt (1921b)). Length 2.5 mm. Antennal horns unusually long, narrowly triangular [Fig. 6n
(from Felt 1921b, Fig. 5)]. Prothoracic spiracle slender, strongly curved.
Male, larva unknown.
Remarks. Felt (1921b) only tentatively placed this species in Asphondylia and wrote that it might belong to
Schizomyia due to the heavy [dense] and relatively high circumfila and the absence of well-developed lobes at the
base of the ovipositor. Only further collecting can solve this problem.
Biology. This species was reared from a stem gall on Litsea sp. (Lauraceae) 21-iv-1914 by DvL (1921). He
stated that the gall specimen became lost before it could be illustrated and described it only as a swelling at the top
of the stem with several larval chambers.
Geographical distribution. This species is known only from the type locality in Ungaran Mts, Java, where it
was collected at 1400 m altitude, 21-iv-1914 (Felt 1921b; DvL 1921).
Asphondylia strobilanthi Felt
[Figs 7a–n]
Asphondylia strobilanthi Felt, 1921b: 150.
Material examined. Syntypes, male, pupa [with female inside] and 2 larvae, reared from galls on aerial root of Stro-
bilanthes cernua Bl. (Acanthaceae) collected at Mt Gede, Cibodas, Java, alt. 1800 m, xii-1918, Felt #a3097. One
slide bears the male, the other the remainder of the series. The male is well-preserved, all legs entire, wings slightly
folded, one antenna with remaining 7, the other with 6 flagellomeres, and terminalia in posterior position. The pupa
and the female inside it are well-preserved. One of the larvae is cleared, the other uncleared.
Description. Male. Length 3 mm, wing length 3.8 mm, width 1.4 mm. Antenna: first flagellomere 5.5x longer
than wide, sixth 4.5x [Fig. 7e]. palpus 2-segmented, first segment short, irregular, second broadly oval. Apicoventral
spur on first tarsal segment long, slightly bent [Fig. 7c]. Tarsal claws strongly curved at distal third, empodia as long
as claws, pulvilli about ¼ empodial length [Fig. 7d]. Gonostylus 2x longer than wide in posterior view, teeth blunt,
of slightly unequal length [Fig. 7b].

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FIGURES 7a–n. Asphondylia strobilanthi: a, galls on aerial root of Strobilanthes cernua, cross section of single gall at bottom
right (Fig. 9 of DvL (1921); b–e, male: b, gonostylus (posterior); c, first tarsal segment; d, last tarsal segment with claw, empo-
dium and pulvillus; e, sixth and seventh flagellomeres; f–h, female: f, last tarsal segment with claw, empodium and pulvillus; g,
last flagellomeres; h, end of needle-like protrusible part of ovipositor; i–k, larva: i, head (dorsal); j, sternal spatula with adjacent
papillae; k, end of body (dorsal); l–n, pupa: l, prothoracic spiracle; m, anterior segments (ventral); n, last three abdominal seg-
ments (dorsal).
Female. Description based on specimen still inside its pupa. Flagellomeres 1–9 progressively slightly shorter,
10–12 progressively much shorter [Fig. 7g], first flagellomere 5.5x longer than wide. Needle-like protrusible part of
ovipositor slightly widened at terminus in ventral view [Fig. 7h].
Pupa. Length 5 mm, thorax dark, abdomen dark yellow (Felt 1921b). Antennal horns long, triangular, serrate
ventrally [Fig. 7m]. Frontal anterior horn robust, bilobed, lobes pointed; posterior horns absent [Fig. 7m]. Cephalic

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papillae minute [Fig. 7m] Prothoracic spiracle unusually long, narrow, of constant width, trachea bulging, ending
at basal fifth [Fig. 7l]. Abdominal dorsal spines simple, relatively small, arranged in sparse anterior field and dense
posterior row, size of spines increasing posteriad [Fig. 7n].
Larva. Length 2.5 mm. Yellowish orange. Head rounded, trapezoid, wider than long, antennae 1.5x longer
than wide at base, posterolateral apodemes absent [Fig. 7i]. Spatula broad, short-shafted; with two long, narrowly
triangular teeth separated by distance 3x greater than their basal width; on either side with 2 pairs of lateral setose
papillae; no pigmented area around shaft [Fig. 7j]. Terminal segment with anus positioned ventrocaudally and no
terminal papillae visible on available specimens [Fig. 7k].
Remarks. Asphondylia strobilanthi is unique among its Indonesian congeners for the narrow, widely separated
lobes of the larval sternal spatula [Fig. 7j], and, in the pupa, the long, tube-like prothoracic spiracle [Fig. 7l], the
two-lobed anterior frontal horn, the lack of posterior frontal horns [Fig. 7m], and the ventrally serrated antennal
horns [Fig. 7m]. It can be separated from Indonesian congeners whose males are known by the prolonged gonosty-
lus [Fig. 7b].
Biology. DvL (1921, Fig. 9 [Fig. 7a]) described the gall on the aerial roots of Strobilanthes cernua, a plant that
grows gregariously in virgin forests on mountain sides [of Mt Gede, Cibodas, Java]. Parts of the root develop above
ground, which is where these were found, partly hidden under remains of leaves and patches of moss. They are galls
of the bark, whose shape is either pyriform or more globular, stout and round at the base, and gradually narrowing
towards the apex. The largest galls measured 10 mm in length, with a thickness of 6 mm. The usual size is smaller,
being about 4 to 7 mm. The rind of the bark is thick and the chamber wide. The surface is densely pilose with long,
white, unbranched hairs. Galls are found both singly and gregariously.
Geographical distribution. This species is known from Mt Gede, Cibodas, Java, where it was collected at
1800 m altitude, xii-1918 (Felt 1921b), and at 1600 m, xii-1913 and x.1925 (DvLR & DvL 1926).
Asphondylia vitea Felt
[Figs 8a–i]
Asphondylia vitea Felt, 1918: 284.
Material examined. Lectotype of Asphondylia vitea Felt here designated, male, reared from stem galls on Cissus
trifolia (L.) K. Sch. [now Cayratia trifolia (L.) Domin (Vitaceae)], Manila, the Philippines, 1907, Bureau of Science
accession No. 6650, W. Schultze, Felt #a2839; paralectotypes, one male, one female, same data as lectotype except
Felt #a2840. All the specimens are whole-mounted, uncleared, with all characters preserved. Felt (1918) originally
noted also that collections were made in 1905 by C.S. Banks and in 1910 by E.D. Merrill, but we have seen no
specimens corresponding to those collection dates.
Description. Male. Wing [Fig. 8h] 2.5 mm long, 0.9 mm wide. Flagellomeres cylindrical with short necks,
progressively longer, 6th segment 3.5x longer than wide, 12th 5x longer than wide; circumfila comprising four wavy
longitudinal bands, (likely) connected by loops at anterior and posterior end [Fig. 8g]. Palpus 3-segmented, second
segment 3x longer than first, third 1.5x longer than second [Fig. 8d]. Apicoventral tarsal spur robust, curved slightly
upwards [Fig. 8e]. Tarsal claws strong, curved at distal third [Fig. 8i], those of the foreleg similarly shaped but not so
robust as those of the mid and hindleg. Terminalia [Fig. 8f]: gonocoxite robust, gonocoxal apodemes long, separated
distally; gonostylus twice as long as wide in posterior view, with merged, equally large distal teeth; cerci ovoid;
lobes of hypoproct triangular, divided by shallow, narrow incision; aedeagus long, narrow, rounded distally.
Female. Ovipositor with moderately large dorsal pair of pseudocerci basally; needle-like protrusible part of
ovipositor short, 1.6x as long as seventh sternite.
Pupa, larva unknown.
Remarks. Felt (1918) stated that no gall description was associated with the type material of this species from
Manila, but Uichano (1919, Figs 4 & 6 in Tab. VIII [Figs 8a, b]) described and photographed the gall of this species
collected at Laguna, Luzon between viii and ix-1917. DvLR & DvL (1926, No 891, Fig. 636 [Fig. 8c]) reported and
illustrated a petiole and stem gall from Cayratia trifolia in Java which they deemed “probably identical” to the gall
described by Uichano (1919) and attributed to Asphondylia vitea Felt. Despite there being no insects available from
this Javanese collection, we are including A. vitea in the Indonesian fauna because the same kind of gall occurs on

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 19
the same host there. In Indonesia, Asphondylia viticola also occurs on the same host, but its galls are lopsided [see
Fig. 4c], unlike those of A. vitea that are symmetrically developed around petioles and stems [Figs 8a–c].
Biology. DvLR & DvL (1926, No 891, Fig. 636 [Fig. 8c]) describe the gall on Cayratia trifolia as oval or almost
cylindrical outgrowths of the stem which is distended in all directions. All the tissues of the stem take part in the
gall-formation. The surface is green, glabrous or pubescent, the wall succulent. The galls are 10–50 mm long and
about 7 mm across, with larger specimens containing more larval chambers than smaller ones.
Geographical distribution. This species is known from the Philippines and Indonesia. In Indonesia it was
found at Jrakah, near Semarang, Java, ii-1012 (DvLR & DvL 1926).
FIGURES 8a–i. Asphondylia vitea: a–b, galls on Cayratia trifolia from the Philippines (Figs VIII-4 & VIII-6 of Uichanco
(1919)); c, gall on C. trifolia from Indonesia (Fig. 636 of DvLR & DvL (1926)); d, palpus; e, first tarsomere; f, male terminalia
(dorsoposterior); g, 6th male flagellomere (only part of circumfila visible in type specimen); h, wing; i, tarsal claw of front leg
with empodium. Insect drawings made from types reared in the Philippines.
Asphondylia viticola Kieffer & Docters van Leeuwen-Reijnvaan
[Figs 4b–c]
Asphondylia viticola Kieffer & DvLR, 1910: 124.
Material. Syntypes, male, female and pupal specimens reared from stem and leaf stalk galls on Vitis trifolia L. [now
Causonis trifolia (L.) Mabb. & J.Wen, previously also known as Cayratia trifolia (L.) Domin] (Vitaceae) collected
at Salatiga, Java, [before] x-1910 (Kieffer & DvLR 1910; DvLR & DvL 1909b). The types are presumed lost.

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Description (based on Kieffer & DvLR (1910), some insignificant characters omitted). Adult. Palpus 3-segmented,
first and second segments as wide as long, third twice as long as first and second together. Flagellomeres cylindrical;
in male about 5x longer than wide, with looped circumfila typical for Asphondylia; in female progressively shorter,
first flagellomere about 6x longer than wide, tenth 2.5x, eleventh 1.5x, twelfth round.
Pupa. Antennal and frontal horns exceptionally strong, antennal horns long and pointed, one anterior frontal
horn, two closely placed upper posterior horns and one lower posterior horn. Cephalic papillae with setae half the
length of antennal horns. Prothoracic spiracles slightly shorter than posterior horns, thin, curved.
Larva unknown.
Remarks. The types are presumed lost (Gagné & Jaschhof 2017) so any distinction between this species and A.
vitea, both gallformers on the same host, cannot be ascertained. DvLR & DvL (1941) found a similar stem gall on
Causonis japonica (Thunb.) Raf. in Java at Cipanas, near Cisolok, alt. 50 m, xi-1927.
Biology. Galls were described by Kieffer & DvLR (1910), DvLR & DvL (1909b, Fig. 48 [Fig. 4b], young gall]
and DvLR & DvL (1926, Fig. 637 [Fig. 4c], mature gall). Asphondylia viticola causes galls on stems and sometimes
on leaf stalks of Causonis trifolia. Galls are brown, lopsidedly developed, 10–50 mm long and 6–12 mm wide, the
infested part of the stem is often curved, inside are one to several larval chambers. The gall is succulent and covered
with a red-brown corky layer. Adults emerged at the end of October at Salatiga, Java [before 1910].
Geographical distribution. Asphondylia viticola was found in Java: Salatiga (in October of an unrevealed year
as stated in 1910 publication), Sumber Pitu, near Weleri; Sukamangli, near Weleri, alt. 500 m, ii-1909; Madura is-
land, v-1910; Kuripan, near Pekalongan, vii-1911, Semarang, i-1912; Jrakah, near Semarang, iii-1913; Pegangsaan,
Jakarta; v-1915; Depok, x-1918 (Kieffer & DvLR 1910, DvLR & DvL 1909b, 1926).
Asphondylia yushimai Yukawa & Uechi
[Figs 4d–e]
Asphondylia yushimai Yukawa & Uechi, 2003 in Yukawa et al. (2003a: 77), nomen nudum.
Asphondylia yushimai Yukawa & Uechi, 2003 in Yukawa et al. (2003b: 265).
Type series. Holotype male (slide Cecid. No. B2461) from malformed pod of soybean, Glycine max (L.) Merrill
(Fabaceae), collected at Kamifukumoto, Kagoshima City, Japan in mid x-1979, emerged 17-x-1979; paratypes: 35
males, 35 females, 14 larvae, 26 pupae, collected and reared from G. max (slides Cecid. Nos. B2401–2413, B2417–
2436, B2451–2455, B2462–2479) and Prunus zippeliana Miquel (slides Cecid. Nos. C0501–0514, C0521–0530,
C0544–0565, C0571–0578) in Japan at various dates (Yukawa et al. 2003a). Types are lodged in the Entomological
Laboratory, Faculty of Agriculture, Kyushu University, Japan. We did not find it necessary to examine the types of
this species.
DNA. Yukawa et al. (2003a) described the morphology and sequenced COI mitochondrial gene fragment from
populations on G. max (GenBank accession numbers AB085776–AB085787, AB085856–AB085859, AB085863–
AB085866, AB085868, AB085870, AB085871, AB086429) and on P. zippeliana in Japan (AB085860–AB085862,
AB085867, AB085869). Uechi et al. (2005) sequenced COI from population on Osmanthus heterophyllus (G. Don)
P. S. Green in Japan (AB194469–AB194477). Uechi & Yukawa (2006) sequenced COI from population on G.
max in Japan (AB197981–AB197984). Uechi et al. (2018) sequenced COI from population on G. max in Japan
(LC348697–LC348701) and South Korea (LC348702–LC348706). GenBank (accessed 15-xii-2019) currently con-
tains 50 sequences of COI, 6 sequences of 12S ribosomal gene and 4 of other genes, with the intraspecific divergence
in COI <1.59%.
Morphology. This species can be distinguished from its congeners with known larvae and pupae by the combi-
nation of the following characters (Yukawa et al. 2003a). Larval sternal spatula with long shaft; four anterior lobes,
pointed apically, outer lobes longer than inner lobes; adjacent three inner and two outer lateral papillae, all setose.
Pupal antennal horn long, finely serrated on medial edge; anterior frontal horn simple, pointed; three posterior fron-
tal horns forming a roughly equilateral triangle.
Biology. Asphondylia yushimai causes malformation of the pods of soybean, Glycine max (Fabaceae) [Figs 4d,
e], and fruit malformation on Prunus zippeliana (Rosaceae) and Osmanthus heterophyllus (Oleaceae) Uechi et al.
(2018). This insect is one of the major pests of soybeans in Japan (see references in Yukawa et al. 2003a). It is one of
several Asphondylia spp. with alternate hosts (Uechi et al. 2004). In Japan, A. yushimai utilizes G. max as the summer
host, and P. zippeliana and O. heterophyllus as the autumn-to-spring hosts (Uechi et al. 2018).

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 21
Geographical distribution. Asphondylia yushimai was found feeding on Glycine max, Prunus zippeliana and
Osmanthus heterophyllus in Japan, and on G. max in South Korea, China and Indonesia (Nakayama 1982 [as As-
phondylia sp.]; Yukawa et al. 2003a; Uechi et al. 2018). The extent of this insect’s distribution and the damage to
soybean production in Indonesia have yet to be assessed.
Asphoxenomyia Felt
[Figs 9a–l]
Asphoxenomyia Felt, 1927a: 382.
Type species: Asphoxenomyia smilacis Felt, by original designation.
Description. Adults. Wing with R5 bent distally, joining C at wing apex; C broken at juncture with R5; Rs not pres-
ent; M4 and Cu1 forming fork [Fig. 9a]. First tarsal segment without apicoventral spur [Fig. 9f], last tarsal segment
unusually long [Fig. 9g], tarsal claws strongly bent at basal third, pectinate, empodia reaching bend in claws, pulvilli
minute [Fig. 9h]. Occipital protuberance absent. Palpus 1-segmented [Fig. 9b]. Number of flagellomeres unknown
as terminal ones missing for all antennae of available specimens but presumably 12.
Male. Flagellomeres cylindrical, similar in size; circumfila anastomosing, irregular, dense [Figs 9c, d]. Termi-
nalia (mounted laterally and compressed on both available males): gonocoxite short, stout, broadly rounded; gono-
stylus short, unidentate; aedeagus robust; hypoproct deeply and triangularly emarginate; cerci elliptical [Fig. 9e].
Female. Abdominal seventh sternite appreciably longer than sixth. Eleventh and twelfth flagellomeres missing
[Fig. 9j]; circumfila anastomosing, irregular, simpler than in male [Fig. 9k]. Abdominal seventh sternite appreciably
longer than sixth. Terminalia: partially obscured, only distal parts clearly visible, without apparent lobes at posterior
end of eighth tergite: cerci subglobular, separated, covered with setae and microtrichia [Fig. 9i].
Pupa, larva unknown.
Remarks. This monotypic genus belongs to the tribe Asphondyliini, characterized by the female seventh ster-
nite being much longer than the sixth and a foreshortened gonostylus situated dorsocaudally on the gonoxocite. It
belongs most likely to the subtribe Schizomyiina, based on the lack of an apicoventral spur on the first tarsomere
and the apparent lack of large lobes at the posterior end of the female eighth tergite. It can be distinguished from
other genera of Schizomyiina in the triangular shape of the apical gonostylar tooth (except Luzonomyia Felt) [Fig.
9e] and the strongly curved, pectinate tarsal claws [Fig. 9h]. Asphoxenomyia shares with the monospecific Lu-
zonomyia, a triangular gonostylar tooth, a robust aedeagus (see Gagné 1969) and a short ovipositor (see Elsayed
et al. 2018a) but the relationship between these two genera can be further clarified only on properly prepared and
mounted specimens, once both species have been collected again.
Asphoxenomyia smilacis Felt
[Figs 9a–l]
Asphoxenomyia smilacis Felt, 1927a: 382.
Material examined. Syntypes, 2 males, female, reared from leaf galls on Smilax modesta DC., collected at Mt
Gede, Cibodas, Java, alt. 1600 m, 14-i-1926, Felt #a3449, One slide bears a male, the other a male and a female,
all mounted laterally and uncleared. One male has six and seven flagellomeres remaining with well-preserved cir-
cumfila, one fully extended wing, the other folded, an entire midleg, the remaining legs missing the distal tarsal
segments, and the substantially compressed terminalia. The other male has two and three flagellomeres remaining,
circumfila and wings well-preserved, two mid and one hindleg present, remaining legs missing the distal tarsal
segments, terminalia heavily compressed, mostly filled with bubbles containing black matter. The female bears 10
flagellomeres on each antenna, one well-preserved wing, the other folded, an entire midleg present with claws in-
side a clear bubble, remaining legs missing their tarsi, and terminalia with only distal parts visible, remainder with
bubbles containing black matter.

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FIGURES 9a–l. Asphoxenomyia smilacis: a–h, male: a, wing; b, palpus; c, basal part of antenna; d, 5th flagellomere; e, termi-
nalia (lateral; arrows indicate aedeagus (ae), cercus (ce), gonocoxite (gc), gonostylus, (gs), hypoproct (hy)); f, first tarsomere; g,
last tarsomere; h, tarsal claw of midleg with pulvillus and empodium; i–k, female: i, terminalia (lateral); j, basal part of antenna;
k, 9th flagellomere; l, galls on Smilax modesta (Fig. 59 of DvLR & DvL (1926)).
Description. Male. Length 3 mm (Felt 1927a), wing length 3.3 mm (3.3–3.4), width 1.4 mm (1.3–1.5). An-
tenna: scape slightly longer than wide, pedicel 1.5x wider than long, fifth flagellomere 2.5x longer than wide [Fig.
9c]. Palpus as in Fig. 9b. Last tarsal segment 10x longer than wide at midlength [Fig. 9g]; tarsal claws pectinate,
bent at acute angle at basal third; empodia as long as bend in claws; pulvilli one-third length of empodia [Fig. 9h].

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 23
Terminalia with wide aedeagus bearing small apical process with a few ventral asetose sensoria; hypoproct with
deeply incised lobes each bearing long seta, cerci setose apically.
Female. Length 3 mm (Felt 1927a). Wing length 3.9 mm, width 1.3 mm. Flagellomeres [Figs 9j, k]: first 3x
longer than wide, second to tenth equal in length, 2.5x longer than wide, eleventh and twelfth missing. Ovipositor
with subglobular cerci, setae dorsally.
Larva, pupa unknown.
Biology. This species induces a mass of small, round leaf galls on Smilax modesta (Smilacaceae), described and
illustrated by DvLR & DvL (1926, gall No. 103 Fig. 59 [Fig. 9l]). Affected leaves are entirely deformed except the
apices that remain normal. Individual galls are small, approximately spherical and hard. They occur closely together
in groups of 40–50, causing the leaf to thicken and bend. Galled leaves are only about 20 mm long while the normal
leaves are about 100 mm in length.
Geographical distribution. This species is known from two localities on Java: Mt Putri, near Bandung, alt.
1400 m, ii-1916; Mt Gede, Cibodas, alt. 1600 m, 14-i-1926 (DvLR & DvL 1926; Felt 1927a)
Bruggmanniella bursaria (Felt). New combination.
[Figs 10f–m]
Asphondylia bursaria Felt, 1927b: 387.
Material examined. Syntypes, 5 males, 9 females and pupa(e) (NYSM #a3393), on two slides, reared from stem
galls on Symplocos fasciculata Zoll. collected at Mt Gede, Cibodas, near Bogor, Java, Indonesia, altitude 1500 m,
ii-1925. The adults are on two slides, one with 2 males and 5 females, the other, 3 males and 4 females, all uncleared
and mounted laterally. No pupa appears on either slide so is evidently lost.
Description. Male. Length 2 mm (Felt 1927b). Wing length 2.5 mm, width 1.2 mm (1.2–1.3). Head. Antenna
with scape 1.5x longer than wide, 2x length pedicel; pedicel wider than long; flagellomeres 12 in number, evenly
cylindrical, 3x longer than wide, slightly constricted at basal third [Fig. 10i]; circumfila comprising four wavy lon-
gitudinal bands connected by high loops anteriorly and low loops posteriorly. Palpus 2-segmented, second segment
slightly shorter and narrower than first [Fig. 10j]. Wing with C ending near wing apex, R1 ending slightly anteriorly
of wing midlength. Tarsal claws simple, narrow, bent at distal third, empodia reaching bend in claws [Fig. 10g];
pulvilli minute. Apicoventral spur on first tarsomere robust, short, bent at right angle [Fig. 10h]. Gonostylus with
two separate teeth, one more pointed than the other [Fig. 10f].
Female. Length 2.5 mm (Felt 1927b). Wing length 2.6 mm (2.5–2.7), width 1.4 mm (1.3–1.4). Flagellomeres
10–12 progressively shortened [Fig. 10m], circumfila sparse, comprising three wavy longitudinal bands connected
by low loops at either end. Needle-like protrusible part of ovipositor about 2x longer than seventh sternite [Figs
10k, l].
Pupa, larva unknown.
Remarks. Known only from the adults, B. orientalis could fit in either Bruggmanniella or Pseudasphondylia,
genera defined on the basis of larval and pupal characters (Gagné 1994; Elsayed, pers. comm.). We tentatively com-
bine this species with the older generic name, Bruggmanniella.
Biology. Bruggmanniella bursaria induces a stem gall on Symplocos fasciculata (Symplocaceae), described
and illustrated by DvLR & DvL (1926, gall No. 1216 Fig. 877 [Fig. 10n]). On young stems, irregular outgrowths
develop that are about 3 mm across, each containing an oval larval chamber lined with fungal tissue. The galls are
often arranged gregariously, giving an appearance of a complex gall 50–100 mm in length. Affected stems are often
bent or curved. Pupation takes place in the gall.
Geographical distribution. The currently known range comprises three localities in the Bogor area, Java
(DvLR & DvL 1926): Mt Gede, Cibodas, altitude 1500 m, collected by M. E. Jacobson ii-1916, ix-1918, xii-1918
and ii-1925; Mt Panisian, near Bogor, altitude 700 m, xii-1923; Mt Salak, Gunung Bunder, altitude 1000 m, viii-
1925.

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FIGURES 10a–e. Bruggmanniella orientalis: a, gall on Phoebe declinata (Fig. 331 of DvLR & DvL (1926)); b, male last tarsal
segment with claw and empodium; c, male first tarsal segment; d, male palpus; e, gonostylus (posterior). Figs 10f–m. Brug-
gmanniella bursaria: f, gonostylus (dorsal); g, male last tarsal segment with claw and empodium; h, male first tarsal segment; i,
male last 6 flagellomeres; j, female palpus; k, ovipositor (lateral); l, apex of ovipositor (lateral); m, female last six flagellomeres;
n, gall on Symplocos fasciculata (Fig. 877 of DvLR & DvL (1926)). Arrows indicate hypertrophic bark tissue (hb), apex tube
containing larval chamber (tl), single gall (sg).
Bruggmanniella orientalis (Felt). New combination.
[Figs 10a–e]
Diceromyia orientalis Felt, 1927b: 388.
Apoasphondylia orientalis (Felt): Gagné 1973: 493, new combination.
Material examined: Holotype male reared by DvL, 3-ii-1924 from Phoebe declinata Nees. (Lauraceae) collected
in virgin forest, near Sibolangit (Medan area), Sumatra, Indonesia, Felt #a3387). The specimen was “badly broken”
when Felt (1927b) received it. The Canada balsam slide preparation contains the uncleared body with flagellomeres
missing, wings collapsed and only two entire legs present, one fore- and one hindleg. The terminalia are viewable
caudally, only the gonostyli are visible.
Description. Male. Length 2 mm. Palpus 1-segmented, twice as long as wide, with long setae [Fig. 10d]. First
tarsal segment with strong, slightly curved, pointed, ventroapical spur [Fig. 10c]. Tarsal claws rather narrow, bent at
distal third, empodia reaching bend in claws [Fig. 10b], pulvilli tiny. Gonostylus as long as wide at midlength, with
two separate, pointed, distal teeth, 3x as long as basal width, one of them slightly shorter than the other [Fig. 10e].
Female, pupa, and larva unknown.
Remarks. This species, previously assigned to Apoasphondylia Gagné, a replacement name for the preoccupied
Diceromyia Felt, shows no close affinity to the type species A. vernoniae (Felt) that causes an undescribed gall on

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 25
Vernonia lancifolia Merr. (Asteraceae) in the Philippines (Felt 1918). That species has a single, bifurcate gonostylar
tooth with two greatly uneven points, one of them twice the length of the other (Gagné 1969, Fig. 4). Known only
from the male, B. orientalis could fit in either Bruggmanniella or Pseudasphondylia, genera defined on the basis of
larval and pupal characters (Gagné 1994; Elsayed, pers. comm.). We elect to place A. orientalis in Bruggmanniella,
the older name of the two possibilities.
Biology. This species induces a complex stem gall on Phoebe declinata (Fig. 10a), described and illustrated by
DvLR & DvL (1926, gall No. 468, Fig. 331). The gall consists of a large, irregular, amorphous swelling made of
hypertrophic bark tissue from which several thin, cylindrical, pointed tubes protrude. The basal swelling is grey-
brown, about 30 mm long and 10–15 mm wide, and does not alter the woody tissue inside the bark. The protruding
tubes, green with red bases, narrow, pointed, hard-walled and 10–20 mm long in total, are embedded within the
swelling. Each tube contains a larval chamber that stretches along the length of the tube and is occupied by a single
larva.
Geographical distribution. This species is currently known only from the type locality near Sibolangit, Medan
area, on the east coast of Sumatra where it was collected from Phoebe declinata 3-ii-1924.
Dasineura elatostemmae Felt
[Figs 11a–g]
Dasyneura elatostemmae Felt, 1921b: 140.
Material examined. Syntypes, male, female, reared from petiole galls on Elatostema sp. [specified as Elatostema
paludosum Miq. by DvLR & DvL (1926), see Remarks below] at Mt Ungaran, alt. 1000 m, 1910, Felt #a3095). Felt
(1921b) gives the year 1914, likely an error (see DvL 1921). Felt’s single slide bears both type specimens. The male
is mounted laterally, the abdomen uncleared, wings entirely shriveled, head and palpi partly shriveled, antennae
with all 12 flagellomeres present, slightly shriveled, two legs with all segments, parts of remaining legs missing,
terminalia partially misaligned and slightly shriveled. The female is uncleared, with wings entirely shriveled, head
and palpi partially shriveled, antennae with all 12 flagellomeres present, slightly shriveled, all legs entire, cerci
well-preserved.
Description. Male. Length 1.25 mm (Felt 1921b). Palpus 4-segmented, progressively longer (Fig. 11f). Flag-
ellomeres with nodes twice as long as wide, necks as long as nodes (Fig. 11c). Tarsal claws toothed, as long as
empodia (Fig. 11d). Terminalia: mesobasal lobes sheathing aedeagus with 4 short-setose distal papillae, gonoxocal
apodemes merged basally, cerci large, tapered to apex, hypoproct divided into two narrow lobes, each with setae
apically, gonostylus long, tapered, sparsely setose, setulose at basal 1/7 dorsally and 3/5 ventrally (Fig. 11b).
Female. Length 1.5 mm (Felt 1921b). Flagellomeres with neck 1/10 length nodes (Fig. 11g). Fully extended
ovipositor as long as remainder of abdomen; fused cerci 5x longer than wide, cylindrical, setulose, setose, these
denser apically, several of them thickened; hypoproct about 1/5 length of cerci (Fig. 11e).
Pupa unknown.
Larva. White (DvL (1921).
Remarks. At the time of the type series collection at Mt Ungaran on 7-v-1910, the host plant was not in bloom
and its identification to species level was not possible (DvL 1921). The plant was identified later as Elatostema
paludosum Miq. by DvLR & DvL (1926), evidently based on the second collection of this species in xi-1919.
Biology. This species causes [single-chambered] stem and petiole galls on Elatostema paludosum Miq. (Urti-
caceae). The gall was described by DvL (1921, Fig. 1 [Fig. 11a]) as follows. They are mostly spheroid or oblong
swellings of the stem and the midrib. The petiole and the flower stalk can also be infested. The surface is smooth
and light green. Inside is a small larval chamber with a white larva.
Geographical distribution. This species is known from its type locality at Mt Ungaran, Java, alt. 1000–1200
m, where it was found 7-v-1910 and xi-1919 (DvL 1921; DvLR & DvL 1926).

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FIGURES 11a–g. Dasineura elatostemmae: a, galls on Elatostema paludosum, one in cross section at bottom right (Fig. 1 of
DvL 1921); b, male terminalia (dorsal), top left: gonostylus (ventral), gonocoxites shriveled, cerci and hypoproct artifactually
flipped ventrally; c, outline of male 6th flagellomere, partially shriveled; d, last tarsomere with claw and empodium; e, female
terminalia (lateral); f, male palpus; g, outline of female 6th flagellomere, partially shriveled.
Diadiplosis duni (Harris)
[Figs 12a–g]
Nipponodiplosis duni Harris, 1968: 460.
Diadiplosis duni (Harris): Gagné 2004: 144, new combination.
Type series. Holotype: male (No 16198), 2 paratypes males (Nos 16199–16200), paratype female (No 16211),
Kerevat, New Britain, Papua New Guinea, ex Planococcus sp. on cacao pods, i-1965; 4 paratype males (Nos
16613–16615, 16634), 3 paratype females (Nos 16635, 16636, 16663), same locality as holotype, ex Planococcus
citri (Risso), vii-1965. The entire type series is deposited in the BMNH.
Description. Wing with R5 joining C at wing apex [Fig. 12a]. Palpus 3-segmented. Occipital protuberance ab-
sent. Flagellomeres 12. Male third flagellomere [Fig. 12b]: basal node as wide as long, internode half-length basal
node, distal node slightly constricted, about 1.5x longer than wide, neck half-length distal node, circumfilar loops
of basal node reaching base of distal node, basal loops of distal node reaching node midhalf, distal loops reaching
neck’s end. Female flagellomeres [Fig. 12e] twice as long as wide, circumfila consisting of two transverse and two
longitudinal, interconnected bands. Tarsal claws toothed on all legs. Male terminalia [Figs 12c, d]: aedeagus sub-
stantially longer than cerci and hypoproct, gonostylus widest at basal third, gonocoxal apodemes not fused anteri-

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orly, shorter than distance between them. Female terminalia [Figs 12f, g] with cerci rounded, 2x longer than wide.
Pupa, larva unknown.
Remarks. Diadiplosis duni differs from D. smithi in the tarsal claws being toothed on all legs as opposed to
only on forelegs (Harris 1968); the aedeagus being substantially longer than the hypoproct and cerci as opposed to
as long as hypoproct and cerci; and gonocoxal apodemes being shorter [Fig. 12d] as opposed to longer than distance
between them [Fig. 13g].
Biology. Larvae of this species were found preying on mealybugs Planococcus sp. (Hemiptera: Pseudococci-
dae): P. citri on citrus in Papua New Guinea and P. lilacinus on cacao pods in Indonesia (Harris 1968). On all three
occasions where they were found, Diadiplosis smithi was also present (Harris 1968).
Geographical distribution. This species has been recorded in Papua New Guinea from Kerevat on the island
of New Britain and in Indonesia from Bogor, Java, 2-iv-1937 (Harris 1968).
FIGURES 12a–g. Diadiplosis duni: a, wing; b, third male flagellomere; c, gonostylus (dorsal); d, male terminalia (dorsal); e,
first and second female flagellomeres; f, female terminalia (lateral); g, same (dorsal). Figures from Harris (1968).
Diadiplosis smithi Felt
[Figs 13a–j]
Diadiplosis smithi Felt, 1915: 178.
Coccodiplosis smithi (Felt): Harris (1968: 430), new combination, redescription.
Coccodiplosis pseudococci Meijere, 1917: 239; Harris (1968: 430), junior synonym.
Types of names in this taxon. D. smithi. Reared from larvae feeding on Planococcus sp. (Hemiptera: Coccidae)
[Pulvinaria sp. in Felt’s original description, corrected to Pseudococcus sp by Harris (1968) based on slide label] on
citrus in Manila, the Philippines, on or before 1915, Felt #a2495a. Not seen, presently on loan to Netta Dorchin, Tel
Aviv University, Israel. We examined Felt’s topotypes, a male and a female (Felt #a2496), reared from larvae feed-

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ing on Planococcus (as Pseudococcus) citri Risso (Hemiptera: Pseudococcidae) presumably on citrus in Manila, the
Philippines, 25-ii-1914. The specimens are mounted on separate slides as whole insects; the male is cleared, with
distal flagellomeres missing, terminalia slightly misaligned; the female is uncleared, with all body parts partially
shriveled. The morphology of these topotypes fits Harris’s (1968) description of Felt’s types except the cerci being
strictly as long as opposed to slightly shorter than aedeagus [Fig. 13g]. We assume the slight difference in the rela-
tive length of cerci is caused by the angle of mounting, a common artifact of Cecidomyiidae slide preparation.
Coccodiplosis pseudococci. Found feeding on the coffee mealybug Planococcus lilacinus (as Pseudococcus
crotonis) in Salatiga, Java (Meijere 1917). We have not seen Meijere’s types (assumedly deposited in Zoological
Museum, Amsterdam, now merged into Naturalis Biodiversity Center, Leiden The Netherlands). Meijere’s descrip-
tion fits Harris’s except that the male cerci are slightly longer (Meijere 1917: Fig. 1d) as opposed to slightly shorter
[Fig. 13g] than the aedeagus. As stated above, the relative length of the cerci is often subject to the position of the
terminalia on the slide mount.
FIGURES 13a–j. Diadiplosis smithi: a, wing; b, third male flagellomere; c, larval spatula; d, head; e, palpus; f, gonostylus
(dorsal); g, male terminalia (dorsal); h, first and second female flagellomeres; i, female terminalia (dorsal); j, (lateral). Figures
from Harris (1968).
Description. Length: male 0.8 mm, female 2 mm (Felt 1915). Wing with R
5
joining C at wing apex [Fig. 13a].
Palpus 3-segmented. Occipital protuberance absent. Flagellomeres 12. Male third flagellomere [Fig. 13b]: basal node
slightly longer than wide, internode half-length basal node, distal node round, slightly longer than wide, neck half-
length distal node, circumfilar loops of basal node reaching node end, basal loops of distal node reaching node basal
third, distal loops reaching neck midlength. Female flagellomeres [Fig. 13h]: nodes twice as long as wide, necks
short, circumfila consisting of two longitudinal and two longitudinal, interconnected bands. Tarsal claws toothed on
fore and simple on mid and hindlegs. Male terminalia [Figs 13f, g]: aedeagus as long as cerci and slightly longer than

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 29
hypoproct, gonostylus widest at midlength, gonocoxal apodemes longer than distance between them. Female termi-
nalia [Figs 13i, j] with cerci tapered in dorsal view, rounded in lateral view, 1.5x longer than wide at base.
Pupa unknown.
Larva. Harris (1968) tentatively (due to the presence of another species of Cecidomyiidae at collection site)
described the larva as bearing on the terminal segment two pairs of dorsal and one ventral papillae, and a sternal
spatula with two triangular lobes divided by deep V-shaped incision. Harris’s (1968) spatula of D. smithi [Fig. 13c]
is similar to that depicted for this species by Meijere (1917) (Fig. 1f) who describes the larva as orange-red, 1.5 mm
in length.
Remarks. We also examined slides of a male and a female reared by L. Kikuchi from larvae feeding on Ni-
paecoccus viridis (as vastator) (Newstead), likely to have been infesting the commercially used tree Leucaena
leucocephala (Lam.) de Wit [see Guam Agricultural Station (1979)] at Yigo, Guam, vii-1981. The specimens are
mounted on separate slides as whole, cleared insects. These specimens fit Harris’s (1968) description of Felt’s types
except the hypoproct is strictly blunt as opposed to slightly concave [Fig. 13g]. We consider this slight difference
part of the variability of the species.
Biology. Diadiplosis smithi is a predator of polyphagous plant-feeding mealybugs Planococcus citri and P.
lilacinus (Hemiptera: Pseudococcidae).
Geographical distribution. The Philippines (Felt 1915; Harris 1968), Papua New Guinea (New Britain) (Har-
ris 1968) and Indonesia where it was found feeding on P. lilacinus in Java at Salatiga in or before 1917 (Meijere
1917) and at Buitenzorg 2&12.iv.1937 (Harris 1968).
Feltiella acarivora (Zehntner)
[Figs 14a–k]
Diplosis acarivora Zehntner, 1901: 203.
Feltiella acarivora (Zehntner): Gagné (1995: 25), new combination.
Material examined. Zehntner’s description was based on the larva, pupa, male and female in association with the
spider mite Tetranychus exciccator Zehntner (Acari: Tetranychidae) feeding on sugar cane in Java. No collection
data was provided, and the study specimens are presumed lost, but Zehntner included drawings that have allowed
identification of his species. This was proven by Ganaha-Kikumura et al. (2012) who collected and reared it from
populations preying on several spider mite species feeding on various plants in Okinawa Prefecture, Japan. Their
slides are stored in the Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan. We
did not find it necessary to examine them.
Morphology. This description is based on illustrations by Zehntner (1901) and the redescription by Ganaha-
Kikumura et al. (2012). Male. Wing length 0.9–1.2 mm, R5 joining C slightly anteriad of wing apex [Fig. 14a].
Palpus 3-segmented, second and third segments equally long, 2.5x longer than first [Fig. 14b]. Occipital protuber-
ance short. Flagellomeres with circumfilar loops of basal whorl reaching midlength of internode, loops of median
whorl reaching midlength of distal node, loops of distal whorl reaching midlength of neck [Fig. 14d]. Tarsal claws
bowed near midlength, toothed on forelegs, simple on mid and hindlegs [Fig. 14c]. Male terminalia [Figs 14e, f]:
gonocoxite long, narrow, with mesobasal bulge; gonostylus elongate, narrow, cylindrical, setulose near base, cari-
nate beyond; aedeagus broad, evenly cylindrical, slightly concave apically; cerci short, rounded apically; hypoproct
about as wide as and nearly as long as aedeagus, parallel-sided, posterior margin squared with median ligule and
minute setae.
Female. Flagellomere nodes 1.5x longer than wide, necks only 1/5 as long as nodes [Fig. 14i]. Cerci ovoid [Figs
14g, h].
Pupa. Length about 1 mm. Antennal bases rounded, cephalic papillae with long setae [Fig. 14k].
Larva. Length 1.10–1.25 mm. Antennae long. Terminal papillae: two outer pairs with long setae, one inner pair
with short setae, a possible additional pair with very short setae unspecified but present in other species [Fig. 14j].
Remarks. In the revision by Gagné (1995) this species was placed in Feltiella based on Zehntner’s original
drawings. This placement was confirmed by Ganaha-Kikumura et al. (2012) who provided a photo of the male
terminalia, an ink drawing of the aedeagus and hypoproct, a redescription of the species based on newly reared
specimens, and a comparison of the species with the nearly cosmopolitan Feltiella acarisuga (Vallot).

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DNA. Ganaha-Kikumura et al. (2012) sequenced the COI mitochondrial gene for nine individuals reared from
spider mites feeding and various host plants (814 bp, GenBank accession numbers AB698995–AB69900). The in-
terspecific similarity of these sequences was 0–0.12%, while the similarity to the closest species, Feltiella acarisuga
(Vallot), was >7.51% (GenBank, access 16-xii-2019).
FIGURES 14a–k. Feltiella acarivora: a, male; b, palpi; c, tarsal claw with empodium (of mid or hindleg; that of foreleg is
toothed); d, two male flagellomeres; e, male terminalia (ventral); f, distal part of aedeagus and hypoproct (ventral); g, end of
female abdomen (ventral); h, same (lateral); i, two female flagellomeres; j, larva (dorsal); k, pupa (ventral); Figs 15a–e, g–k
from Zehntner (1901), Fig. 14f from Ganaha-Kikumura et al. 2010.
Biology. Feltiella acarivora preys on several spider mites (Acari: Tetranychidae) that feed on various plants.
Zehntner (1901) wrote that full-grown larvae made cocoons on the plant surface. Most records are based on the
identification of the gall midge and the mite rather than the direct observation of the predatory behavior. Mites
reported as prey are Tetranychus exsiccator in Indonesia, and Eutetranychus africanus (Tucker), Oligonychus bi-
harensis (Hirst), T. kanzawai Kishida, T. neocaledonicus Andre, T. okinawanus Ehara, T. parakanzawai Ehara, T.
pueraricola Ehara, and T. urticae Koch (green form) in Japan (Ganaha-Kikumura et al. 2012). Feltiella acarivora
was found co-occurring with spider mites feeding on the following plants: Saccharum officinarum L. in Indonesia
(Zehntner 1901), and Annona atemoya Mabb. (Annonaceae), Bauhinia sp. (Caesalpiniaceae), Canavalia lineata

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(Thunb.) DC. (Fabaceae), Carica papaya L. (Caricaceae), Clerodendrum trochotomum Thunb. (Verbenaceae), Mal-
lotus japonicus (L.f.) Müll.Arg. (Euphorbiaceae), Melanolepis multiglandulosa (Reinw. ex Blume) Rchb.f. & Zoll.
(Euphorbiaceae), Morus australis Poir. (Moraceae), Mucuna macrocarpa Wall. (Fabaceae), Passiflora edulis Sims
(Passifloraceae), Pueraria montana (Lour.) Merr. (Fabaceae) and Vigna angularis (Willd.) Ohwi & H. Ohashi (Fa-
baceae) in Japan (Ganaha-Kikumura et al. 2012).
Geographical distribution. Japan (Iheya, Noho, Okinawa, Irabu, Ishigaki and Iriomote Islands, all Okinawa
Prefecture) (Ganaha-Kikumura et al. 2012), Indonesia (Java, localities unspecified) (Zehntner 1901).
Gephyraulus tetrastigma (Felt). New combination
[Figs 15a–h]
Dasyneura tetrastigma Felt, 1927b: 385.
Material examined. Syntypes, 18 females, labelled as reared from stem gall [but as “leaf gall” in DvLR & DvL
(1926)] on Tetrastigma pergamaceum (Blume) Planch. collected at Mt Gede, Cibodas, near Bogor, Java, Indonesia,
altitude 1500 m, 30-xii-1924, Felt #a3392. Nine whole, uncleared females are mounted on each of two Felt slides.
The specimens are relatively well-preserved, except most of the abdomens are uncleared and shrunken, only one
having the ovipositor outside of the abdomen and observable.
FIGURES 15a–h. Gephyraulus tetrastigma: a, galls on Tetrastigma pergamaceum (Fig. 633 of DvLR & DvL (1926)); b,
contour of female abdomen (partially shriveled, lateral); c, wing; d, last tarsomere with claw and empodium; e, female 8th flagel-
lomere; f, end of ovipositor (ventral); g, end of ovipositor (lateral); h, palpus.

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Description. Female. Wing length 1.9 mm (1.7–2.0), width 0.7 mm (0.6–0.8). Body covered with large scales.
Palpus 4-segmented, segments equally wide, first as wide as long, second to fourth 2.5x longer than wide [Fig.
15h]. Antenna: flagellomeres 16 in most specimens, some with 16th flagellomere extra-long, some with 17; first and
second fused; 6th with node 1.5x longer than wide, neck 1/10 node length [Fig. 15e]. Occipital protuberance absent.
Wing with R5 joining C anteriad of wing apex [Fig. 15c]. Tarsal claws toothed, empodia as long as claws [Fig. 15d].
Abdominal sclerites not recognizable on available specimens. Ovipositor [Fig. 15b] extrusible, 3–4x longer than
8th tergite when fully extended, setulose, sparsely covered with short setae; cerci [Figs 15f, g] fused, bilaterally
compressed, tapering to rounded apex in lateral view, to pointed apex in ventral view [Fig. 15f], glabrous, without
microtrichia, with short setae apically, one pair of setae on dorsum thickened and more prominent; hypoproct nar-
row in lateral view, with two short apical setae [Fig. 15g].
Male, pupa, larva unknown.
Remarks. The species is transferred to Gephyraulus Rübsaamen based on the female cerci that are bilaterally
flattened and glabrous and bear among the sparse short apical setae one noticeably thicker pair. Gephyraulus is a ge-
nus of 14 species of the Palearctic and Oriental Regions, mainly on Brassicaceae (Gagné & Jaschhof 2017; Elsayed
et al. 2017; Jiao et al. accepted for publication). This is the first record of the genus from Vitaceae.
Biology. This species causes a leaf gall on Tetrastigma pergamaceum (Vitaceae) described by DvLR & DvL
(1926, Fig. 633 [Fig. 15a]). The whole leaf is changed into a cylindrical, often curved gall, 20–40 mm long and 4–8
mm across, gradually narrowing towards both ends and tapering into a long point. The surface is pink and glabrous.
Inside is a single, spacious larval chamber covered with a great many cylindrical papillae.
Geographical distribution. This species is known only from the type locality at Mt Gede, Cibodas, near Bo-
gor, Java, Indonesia, altitude 1500 m, 30-xii-1924 and v-1925 (Felt 1927b; DvLR & DvL 1926).
Gynodiplosis Felt
[Figs 16a–i]
Gynodiplosis Felt, 1927a: 383.
Type species: Gynodiplosis humata Felt, by original designation.
Description. Adult. Antennae in both sexes with 12 binodal flagellomeres, basal node with one, distal with two
loops of circumfila [Figs 16f, i]. Occipital protuberance absent. Palpus 1-segmented [Fig. 16g]. Wing with R5
curved distally, joining C beyond wing apex; C broken at juncture with R5; R1 half-length of wing; Rs not present,
M4 and Cu1 forming fork [Fig. 16b]. Tarsal claws with two teeth on all legs [Figs 16d, e].
Male. Terminalia (seen in lateral view only )[Fig. 16c]: gonocoxite with bulbous, setulose mesobasal lobe;
gonostylus long, cylindrical, gradually tapered, with many setae on distal two-thirds, setulose basally, carinate be-
yond; aedeagus longer than gonocoxite, cylindrical, curved dorsally, rounded apically; cerci ovoid with many setae;
hypoproct incised apically, lobes narrow, each with single seta.
Female. Terminalia in lateral view [Fig. 16h]: ovipositor barely protrusible; cerci separate, ovoid, microsetose,
with many setae, an apical pair thickened and blunt.
Pupa, larva unknown.
Remarks. This monotypic genus belongs to the supertribe Cecidomyiidi and is notable for the one-segmented
palpus, the curved, two-toothed tarsal claws and the female flagellomeres with two nodes and three circumfila,
identical to that of the male. It is unplaced to tribe.
Gynodiplosis humata Felt, 1927
[Figs 16a–i]
Gynodiplosis humata Felt, 1927a: 383.
Material examined. Syntypes, male, female, reared from Davallia repens (L.fil.) Kuhn [as Humata alpina Moore
in Felt (1927a) and H. repens Diels in DvLR & DvL (1941)], collected at Mt Gede, Cibodas, Java, alt. 1500 m, 13-
i-1926, Felt #a3448). Felt’s types are mounted in lateral position as whole, uncleared, on separate slides.

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FIGURES 16a–i. Gynodiplosis humata: a, gall on Davallia repens, at top left cross section of gall containing pupa (Fig. 8 of
DvLR & DvL (1926)); b–f, male: b, wing; c, terminalia (lateral); d–e, last tarsal segment of midlegs of same specimen, with
claw, empodium and pulvillus; f, 4th flagellomere; g–i: female, g, palpi; h, end of ovipositor (lateral); i, 4th flagellomere.
Description. Male. Length 3 mm (Felt 1927a). Antenna: scape widened distally, as long as wide distally;
pedicel oval, longer than wide; flagellomeres: [Fig. 16f] first and second not fused, basal node round, as wide as
long, internode half-length basal node, distal node slightly constricted at mid length, twice as long as wide, neck
as long as internode, length of circumfilar loops closely embracing flagellomere: basal loops reaching mid length
of internode, middle loops reaching base of distal loops, distal loops reaching end of neck; nodes densely covered
with microtrichia, internode with few microtrichia. Palpus 1-segmented, 1.5x longer than wide [as in Fig. 16g].
Wing length 3.5 mm, width 1.4 mm [Fig. 16b]. Tarsal claws with two basal teeth, the distal tooth about half size of
claw, the proximal variable in size, ranging from half to same size of distal tooth, even on same leg [Figs 16d, e].
Terminalia as for genus [Fig. 16c].
Female. Length 4 mm (Felt 1927a). Wing length 4.0 mm, width 1.5 mm. Flagellomere [Fig. 16i] similar to
male. Terminalia as for genus [Fig. 16h].
Pupa, larva unknown.
Biology. This species causes a leaf gall on the fern Davallia repens (L.fil.) Kuhn (Davalliaceae) described by
DvLR & DvL (1926, gall No. 14, Fig. 8 [Fig. 16a]). The spherical gall is attached by a short pedicel to the upper
side of a secondary leaf vein. Leaflets surrounding the gall are distorted and may be shaped into a kind of calyx sur-
rounding the base of the gall. The gall is glabrous, thick and solid and surrounds a spacious larval chamber.
Geographical distribution. Gynodiplosis humata is known from Java where it was found at Mt Tangkuban
Perahu, near Bandung, Java, alt. 1600 m, vi-1915 and Mt Gede, Cibodas, alt. 1400 m, i-1925 (DvLR & DvL
1926).

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Hypodiplosis Kieffer
[Figs 17m –o]
Hypodiplosis Kieffer 1913: 187.
Type species: Clinodiplosis paederiae Kieffer & DvLR, by monotypy.
This monotypic genus was erected by Kieffer (1913) for Clinodiplosis paederiae that induces a leaf-roll gall on
Paederia foetida L. from Java (Kieffer & DvLR 1910). The type series is presumably lost (Gagné & Jaschhof 2017),
but the original description includes a drawing of the male terminalia that allows us to place the genus in the tribe
Clinodiplosini for the first time.
Description (adapted from Kieffer & DvLR (1910: 129) and Kieffer (1913: 187). Male. Flagellomeres binodal,
the first node spherical with a single circumfilum, the second cylindrical with two circumfila, the loops regular, none
reaching to base of next distal circumfilum [Fig. 17n]. Palpus 4-segmented. Wing with R5 wing vein reaching C well
beyond wing apex, M4 and Cu1 forming fork. Tarsal claws simple, empodia a bit shorter than claws. Terminalia with
cerci concave apically; hypoproct longer than cerci, slender at base, widening and ovoid distally; aedeagus narrow,
longer than hypoproct; gonostylus elongate, narrow and bowed [Fig. 17o].
FIGURES 17a–l. Eucalyptodiplosis paederiae: a–c: galls on Paederia foetida, a, the Philippines (cross section, magnified 2x
when compared to b, Fig. I-1 of Uichanco (1919)); b, the Philippines (longitudinal section, Fig. I-6 of Uichanco (1919)); c, the
Philippines (overall view, Fig. VIII-5 of Uichanco (1919)); d–f male: d, wing; e, 3rd flagellomere; f, terminalia (dorsal); g–l,
female: g, head (ventral, arrow indicates occipital protuberance); h, terminalia (ventral); i, 3rd flagellomere; j, tarsal claw on
foreleg; k, tarsal claw on midleg; l, tarsal claw on hindleg. Figs 17m–o. Hypodiplosis paederiae: m, galls on Paederia foetida:
Fig. 1012 of DvLR & DvL (1926)); n, last two male flagellomeres (Fig. 8 of Kieffer & DvLR (1910)); o, terminalia (dorsal, Fig.
9 of Kieffer & DvLR (1910)).

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Female. Flagellomeres cylindrical, circumfila closely appressed. Abdomen not protrusible, cerci elongate-
oval.
Pupa. Antennal bases not modified. Prothoracic spiracles long. Dorsal spines present, simple.
Remarks. This monotypic genus belongs to the supertribe Cecidomyiidi. It is placed for the first time in the
tribe Clinodiplosini because of the concave caudal edge of the male cerci. The single unique character that separates
Hypodiplosis from other Clinodiplosini is the tapered end of the hypoproct. All else we know of the genus conforms
to Clinodiplosis in the broad sense.
Hypodiplosis paederiae (Kieffer & Docters van Leeuwen-Reijnvaan)
[Figs 17m–o]
Clinodiplosis paederiae Kieffer & DvLR, 1910: 129.
Hypodiplosis paederiae: Kieffer (1913: 187), new combination.
Type series. Syntypes, male, female and pupa, reared from leaf-roll galls on Paederia foetida L. collected at Salati-
ga, Java, [likely in] i-1909 (Kieffer & DvLR 1910; DvLR & DvL 1926). The types are presumed lost (Gagné &
Jaschhof 2017).
Description. As for the genus above. Male length, 1.5 mm, female, 2.0 mm.
Biology. This species causes a leaf roll gall on Paederia foetida (Rubiaceae). The gall was collected in Java
and described by Kieffer & DvLR (1910), DvLR & DvL (1909a, Fig. 12) and DvLR & DvL (1926, Fig. 1012 [Fig.
17m]).
Geographical distribution. This species was found in Java: Salatiga, alt. 600 m, i-1909; Mt Welirang, alt. 800
m, vi-1911; Pare, near Kediri, alt. 200 m, v-1912; Srondol, near Semarang, alt. 200 m, iii-1913; Bandung, alt. 800,
vii-1915 and vi-1918; Cikampek, alt. 75 m, vi-1924 and on Sebesi Island, alt. 400 m, iv-1921 (DvLR & DvL (1926,
gall Nos 1426, 1428).
Remarks. We cannot ignore here that a similar gall was found in the Philippines (Uichanco (1919, gall No.
18165, Figs I-1, I-6, VIII-5 [Figs 17a–c]) and described as Itonida paederiae Felt (1919: 293), an unplaced species
of Cecidomyiinae (Gagné & Jaschhof 2017). See Addendum at the end of this paper for the newly combined Euca-
lytodiplosis paederiae (Felt) and its redescription.
Lasioptera javanica Kieffer & Docters van Leeuwen-Reijnvaan
[Figs 18a–b]
Lasioptera javanica Kieffer & DvLR, 1910: 123.
Type series. Kieffer & DvLR (1910) collected larvae and reared the type male and female from a stem gall [Fig.
18a] on Zehneria maysorensis (Wight & Arn.) Arn. (as Melothria perpusilla Cogn.) in Java [likely at Salatiga]. The
types are presumed lost (Gagné & Jaschhof 2017).
Description (from Kieffer & DvLR 1910 (changed to fit current terminology)). Adult. Palpus 4-segmented,
first segment as long as wide, second slightly longer than wide, third and fourth about 2.5x longer than wide. An-
tenna with 24 flagellomeres, flagellomeres without constriction. Wing with vein R5 longer than half-length of wing,
nearly 2x as long as R1, M4 and Cu1 forming fork, the branches longer than base; anterior wing edge, R1, R5 covered
with black scales, distal end of R5 with white spot. Tarsal claws toothed, as long as empodia.
Male. Flagellomeres slightly shorter than wide. Terminalia: hypoproct not appreciably longer than cerci, nar-
row, rounded terminally, slightly shorter than aedeagus, aedeagus shorter than gonocoxites.
Female. Length 1.5 mm. Flagellomeres without necks. Ovipositor long, protractible.
Pupa unknown.
Larva. Length 2 mm. Orangey yellow. Spatula dark brown; anterior part as long as wide, with two short blunt
teeth; shaft 3x longer than anterior part.
Remarks. The species description is essentially generic and cannot be improved upon until galls are found
again.

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FIGURES 18a–b. Lasioptera javanica: a, galls on Zehneria maysorensis (Fig. 8 of DvLR & DvL 1909a); b, same (Fig. 1061
of DvLR & DvL (1926)). Arrows indicate galled tissue.
Biology. This species causes a stem gall on Zehneria maysorensis (Wight. & Arn.) Arn. (Cucurbitaceae). The
gall is a several centimeters-long swelling of the young stem, petiole and occasionally leaf midrib (DvLR & DvL
(1909a, Fig. 8 [Fig. 18a]), DvLR & DvL (1926, gall No. 1497, Fig. 1061 [Fig. 18b]). Inside are several larval cham-
bers, each containing fungal mycelium and a single larva that pupates in the gall. Adults emerge in July.
Remarks. Additionally, DvLR & DvL (1926, gall No. 1496, Fig. 1060) described a globular flower gall, con-
taining orange larvae, from Java (Salatiga, alt. 600 m, i-1909; Bandung, alt. 700 m, v-1918, Telaga Warna, near Mt
Gede, alt. 1400 m, ix-1923) and Kayuadi Island, Selayar archipelago (v-1913), which they attributed to this species
(as Prolasioptera javanica Kieffer & Docters van Leeuwen-Reijnvaan). Adults need to be reared to confirm this
latter species’ identity, and we are not associating the gall with L. javanica here.
Geographical distribution. This species is known from Java and Selayar Island, South Sulawesi province
(DvLR & DvL (1926). Java: Salatiga, alt. 600 m, i-1909; Cinyiruan, near Bandung, alt. 1600 m, vi-1910; Mt Unga-
ran, alt. 1000 m, x-1910; Plabuan, near Weleri, alt. 2 m, viii-1912; Mt Muria, alt. 300 m, x-1912; Kamojang, near
Garut, alt. 1500, xii-1915; Mt Slamet, alt. 2000 m, v-1918. Selayar Island: Bitombang, alt. 400 m, v-1913.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 37
Lasioptera manilensis Felt
[Figs 19a–j]
Lasioptera manilensis Felt, 1918: 288.
Material examined. Holotype female (NYSM #a2851, labelled “Type”), reared by L. B. Uichanco from leaf gall
on Leea manillensis Walp. (Vitaceae), Los Baños, Laguna, Luzon, the Philippines, 2-viii-1917, type gall No 18318.
Paratype: female, No. 18318 (Uichanco), Mt Makiling, Laguna, Luzon, the Philippines, 21-viii-1917. We exam-
ined the female holotype and the female paratype. A male, originally listed in Felt (1918) as a paratype (under No.
18318), was not described and is not in the Felt collection. The holotype is a whole mount, cleared, with all char-
acters present, but most of the ovipositor is inserted inside the dorsolaterally mounted abdomen. The paratype is a
cleared remount (by RJG), bearing an additional label reading “C F Baker collection 1927”, and consists of a wing
and a laterally mounted abdomen, the two parts are in good condition and mounted under separate coverslips.
Description. Female. Wing 2.1 mm long, 0.9 mm wide; Rs 2/3 wing length, C interrupted at juncture with Rs,
R1 third wing length [Fig. 19e]. Flagellomeres 21 on both antennae in the holotype, slightly longer than wide, necks
minute, circumfila consisting of two horizonal and one vertical, interconnected bands [Fig. 19h]. palpus 3-seg-
mented, first segment as long as wide, second 2x, third 3x as long as first [Fig. 19i]. Tarsal claws with strong tooth,
empodia as long as claws [Fig. 19f]. Abdominal eighth tergite membranous, with pair of anterior trichoid sensilla
and, beyond on each side, a lateral group of dense spines [Fig. 19g]. Ovipositor: protrusible part nearly bare dorsally,
setose ventrally; cerci: fused, cylindrical, broadly rounded apically; basal, glabrous two-thirds with long, curved,
spoon-like setae dorsally and shorter, apically broadened setae laterally; distal third setulose, with closely spaced,
short, thick setae dorsoapically and scattered, thinner setae elsewhere; pair of discrete, large, setose and setulose,
dorsolateral lamellae [Fig. 19j].
Remarks. The ovipositor of L. manilensis differs from other congeners in the presence of a pair of dorsolateral
lamellae, resembling the lateral lamellae of the Australian Dactylasioptera (Kolesik & Veenstra-Quah 2008; Veen-
stra et al. 2018).
Male, pupa, larva unknown.
Biology. Felt (1918) stated that the types were reared from a leaf gall on L. manillensis and referred to Uichan-
co’s description (1919, gall No. 18318, Figs II-3, IX-1, IX-3 [Figs 19a–c]) which we paraphrase as follows. Mono-
thalamous leaf gall; irregularly subspherical and appearing on both sides of the leaf, the superior lobe smaller than
inferior and sometimes flushed with carmine, otherwise uniformly lettuce green throughout, smooth, non-pubes-
cent. The original puncture of the ovipositor is usually marked by a small, funnel-shaped, eccentric depression. The
inferior lobe is generally carmine, with a thin lettuce green border, the general contour more or less wavy in outline,
smooth and non-pubescent. The leaf lamina immediately adjoining the gall is convoluted circumferentially. The
wall is hard but brittle, the interior is fleshy, light green and semitranslucent. The larval chamber is located in the
center along the longitudinal axis of the gall and curved downward at one end. When the adult is ready to emerge,
a circular flap is pushed outwards from the inferior lobe. The superior lobe is 7.5 to 9.5 mm long and 5 mm wide,
the inferior lobe 7 to 9.5 mm long and 5.5 to 7 mm wide, and 4 to 5.5 mm thick. DvLR & DvL (1926, gall No. 867,
Fig. 622 [Fig. 19d]) found the galls to be common on Leea indica (Burm. f.) Merr. in Indonesia. Their description
agrees in all particulars with that of Uichanco (1919).

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FIGURES 19a–j. Lasioptera manilensis: a–c, galls on Leea manillensis in the Philippines (Figs. IX-1, IX-2, II-3 of Uichanco
(1919)); d, galls on L. indica in Indonesia (Fig. 622 of DvLR & DvL (1926)); e, wing; f, tarsal claw with empodium; g, female
postabdomen from tergite 6 (lateral); h, 3rd female flagellomere; i, palpus; j, end of ovipositor (lateral). Arrow indicates dorso-
lateral lamella.
There are two other types of globular leaf galls induced by Cecidomyiidae on L. indica in Indonesia (DvLR &
DvL (1926) – a round, hairy gall on the leaf’s upper side (gall No. 868, Fig. 623) and a fusiform, glabrous leaf rib-
gall visible on both sides of the leaf (gall No. 869, Fig. 624), but the gall makers were not reared.
Geographical distribution. This species occurs in the Philippines (Uichanco 1919) and Indonesia where it was
found in Java and Sumatra (DvLR & DvL, 1926). Java: Tempuran, near Semarang, alt. 100m, v-1909; Mt Ungaran,
alt. 1000 m, x-1910; Tretes, near Mt Welirang, alt. 600 m, vi-1911; Pare, near Kediri, alt. 200 m, v-1912; Kuripan,
near Pekalongan, i-1911; Plabuan, near Weleri, ix-1912, Mt Muria, alt. 400 m, ix-1912; Nusa Kambangan island,
x-1913; Cisokan, near Cibeber, alt. 700 m, xii-1917; Sawahan, near Mt Wilis, alt. 500, iv-1918; Depok, alt. 100 m,
v-1911, vii-1918, x-1919; Mt Salak, alt. 1000 m, vi-1918, ix-1919, vii-1920; Bogor, alt. 250 m, xii-1918, viii-1919;
Mt Cibodas, Ciampea, near Bogor, alt. 200 m, iv-1919, Wanayasa, near Purwakarta, Java, alt. 1000 m, vii-1920;
Tegalwaru, near Krawang, xii-1923. Sumatra: Padang, ix-1913; Mt Rate, Way Lima, Telokbetung, alt. 400 m, xi-
1921.

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Leefmansiella Kolesik & Gagné. New Genus
[Figs 20a–s]
Type species: Trishormomyia pandani Felt, by present designation.
Description. Body robust [Fig. 20p]. Head without occipital protuberance, frons with setae. Flagellomeres 12, first
and second not fused, binodal in both sexes, bearing one whorl of looped circumfila on basal and two whorls on dis-
tal node, loops longer in male than in female on which the two circumfila on distal node connected by single vertical
band [Figs 20m, o, q, s]. Palpus 3-segmented [Fig. 20l]. Wing: R5 joining C posteriad of wing apex; C weakened
but not interrupted at juncture with R5; R1 slightly shorter than half wing length; Rs as stub on R5, closer to arculus
than end of R1; M4 and Cu1 forming fork [Fig. 20k]. Tarsal claws simple, bent apically, empodia slightly shorter than
claws [Fig. 20j]. Tergites with posterior and lateral setae, without scales. Male terminalia [Fig. 20n]: gonocoxite
cylindrical, without mesobasal lobes, densely covered with setae; gonocoxal apodomes merged basally; gonostylus
with many long setae and setulose basally, with short setae and carinate distally; cerci triangular, very short; lobes
of hypoproct broad, incised apically, with rounded lobes, no longer than cerci,; aedeagus broad, columnar, nearly as
long as gonocoxite, blunt apically with many peg-like setae. Ovipositor short; cerci broad, rounded apically, covered
with dense, simple setae and microtrichia; hypoproct wide, short, setulose, covered with sparse setae [Fig. 20r].
Terminal segment of larva with pair of large lateral lobes [Figs 20g–i].
Etymology. This genus is named after the discoverer of the type species, Dutch entomologist Salomon Leef-
mans (1884–1954).
Remarks. The new genus belongs to the supertribe Cecidomyiidi. The adults fit the tribe Cecidomyiini, but the
larval terminus is highly modified into a bilobed structure and different from most Cecidomyiini which have one
pair of corniform papillae and three pairs with short setae. The type species was tentatively accommodated by Felt
(1921c) in Trishormomyia Kieffer, now a synonym of Planetella Westwood. That genus contains over 50 Holarc-
tic species, with no apparent morphological connection with Leefmansiella (see Gagné 2018). All known hosts of
Planetella belong to Cyperaceae (Gagné & Jaschhof 2017).
Leefmansiella pandani (Felt, 1921)
[Figs 20a–s]
Trishormomyia pandani Felt, 1921c: 270.
Material examined. Syntypes, one male, one female reared from galls on Pandanus nitidus (Miq.) Kurz at Cibodas,
Java, vii-1919 (see Leefmans 1921) (NYSM a 3107). The types on Felt’s slides are whole-mounted, uncleared but
most characters are well observable, all tarsi missing except one in the female.
Description (additional to generic characters). Male. Wing 4.8 mm long, 1.8 mm wide. Flagellomeres: third 3x
longer than wide, segments cylindrical at basal part of antenna becoming binodal distally Fig. 20m]; all segments
with three, separate whorls of circumfilar loops, none reaching next distal circumfilum [Fig. 20o]. Palpus: first seg-
ment as long as wide, second 1.5x longer than wide, third 2x longer than wide [Fig. 20l]. Terminalia as for genus
[Fig. 20n].
Female. Wing 5.0 mm long, 1.9 mm wide. Flagellomeres somewhat binodal, constriction between nodes bare.
Pupa [Figs 20d–f]. Length 4.5 mm, width 1.5 mm. Head, thoracic segments, wing sheaths, terminalia dark
green to black, leg sheaths orange, abdomen carrot-red; antennal horns pointed (Leefmans 1921).
Larva [Figs 20g–i]. Length about 5 mm, width 1.3 mm, carrot-red, anal segment with two lateral sack-like
processes, colourless distally, spatula absent (Leefmans 1921).
Biology. This species causes leaf galls on Pandanus nitidus Kurz (Pandanaceae) (host plant identified by C. A.
Backer, the botanist of the Java Flora, see Felt (1921c) and Leefmans (1921). DvLR & DvL (1926, gall No. 49, Fig.
27 [Fig. 20b]), found the same gall but corrected the host plant name to Pandanus furcatus Roxb. While both plant
names are valid, the main distribution of P. nitidus is confined to Java whereas P. furcatus occurs between Thai-
land, Sri Lanka and Nepal, and we consider the original diagnosis as the likely correct one. Leefmans (1921, Plates
IX– XI, [Fig. 20a]) describes the gall and its development as follows. Galls are oval, up to 5 cm long and 1.5 cm
wide, occurring on both sides of the leaf, with a circular [emergence] hole, 1.5–2.5 mm in diameter, at the underside

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40 · Zootaxa 4847 (1) © 2020 Magnolia Press
of the leaf. The egg is laid on the underside of a young leaf at the top of the plant. The young larva is overgrown
by swollen plant tissue that forms the gall, leaving a narrow tunnel connecting the larval chamber with the surface.
Shortly before the pupation in the gall, the larva closes the opening of the tunnel by spinning a thin membrane and
additionally a second membrane inside the tunnel, just above its body. Before adult emergence, both membranes are
penetrated by the pupa. The pupae were found to be parasitized by a chalcid (Leefmans 1921, Plate XII Figs 6, 7).
Geographical distribution. This species is known from Java and Sumatra. Java: Cibodas, Mt Gede, alt. 1450
m, vii-1919 (Leefmans 1921, host P. nitidus); Mt Ungaran, alt. 1000 m, xii-1909; Mt Salak, alt. 1000 m, vii-1920;
Mt Gede, near Cibodas, alt. 1600 m, x-1923; Mt Mandalagiri, near Garut, alt. 1500 m, iv-1929; Mt Manglayang,
near Bandung, alt. 1400 m, iii-1924; Sumatra: Bandar Baru, near Medan, alt. 1200 m, ii-1917; (all DvLR & DvL
1926, host P. furcatus). Additionally, the galls were found and larvae, pupae and an adult were collected by Melinda
Agustina and Purnama Hidayat (IPB University (pers. comm., [Fig. 20c])) at Mt Gede and Mt Salak, Java, between
February and April 2018.
Lestodiplosis lacciferi (Barnes)
[Figs 21a–c, Fig. 21d (reference)]
Dentifibula lacciferi Barnes, 1935: 525.
Lestodiplosis lacciferi (Barnes): Harris 1968: 453, new combination.
Material. Syntypes (all in BMNH): males (Cecid. 2443–45), females (Cecid. 2446–48); reared from [larvae preda-
ceous on] scale insect Kerria javana (Chamberlin) (Coccoidea: Kerriidae) [as Laccifer javanus Chamberlin], Bukit
Badong, Malaysia, 7-ii-1933. Other material: larva (Cecid. 2451), same data as cotypes but Kuala Lumpur, Malay-
sia. Other material: females (Cecid. 2449, 2450), reared from larvae predaceous on Kerria sp. [as Tachardia sp.],
Kuala, Selangor Road, Malaysia, 15-ix-1932, which Harris (1968) examined and noted that they were fragmentary
specimens and exhibited insufficient evidence either to confirm or disprove their conspecificity with the syntypes.
Other material: reared from [larvae predaceous on] scale insect Kerria javana (Chamberlin) (Coccoidea: Kerriidae)
[as Laccifer javanus Chamberlin], Namkum [Jharkhand state], India ex Java, no date (Harris 1968). We have not
seen the types of this species.
Description. Based on Barnes (1935) and Harris (1968). Male. Length about 1 mm. Flagellomeres 12, binodal,
first and second fused, basal node about as long as distal, internode and neck about as long as nodes, circumfila
looped, bifilar, with third, posterior, whorl reduced to single, very short loop [Fig. 21b]. Palpus 4-segmented; first
segment approximately quadrate; second longer, about 2.5x longer than wide; third and fourth slightly longer than
second. Wing [Fig. 21a]: R5 reaching C slightly anteriad of wing apex; C interrupted at junction with R5; R1 about
third wing length; Rs in form of stump at crook of R5. Tarsal claws simple, strongly curved. Terminalia: gonocoxite
slender, with angular, setulose mesobasal lobe; gonostylus long, narrow; cerci ovoid, lobes rounded, with a few
setae; hypoproct ovoid in dorsoventral view, slightly concave with two setae apically, with definite ventral bulge;
aedeagus about as long as gonocoxites, cylindrical, flattened at apex. Harris (1968) noted that the structure of the
male genitalia of Lestodiplosis lacciferi and L. aonidiellae (described from South Africa) [Fig. 21d] are similar.
Female. Flagellomeres cylindrical, node 2x longer than wide, neck slightly shorter than node, circumfila of 2
interconnected horizontal bands, proximal band straight, the distal bowed [Fig. 21c].
Larva and Pupa. Unknown.
Remarks. Any collector searching for predaceous cecidomyiids should find in a short time a multitude of spe-
cies in Indonesia. Known to date are only three species whose males, at least, are easily differentiated: Lestodiplosis
lacciferi has bifilar, instead of trifilar, male flagellomeres, unique in the genus, a wide, angular, mesally-setulose,
mesobasal gonocoxal lobe, and flattened aedeagal apex [as in Fig. 21d]; Lestodiplosis oomeni has a merely low,
convex mesobasal gonocoxal lobe, a gonostylus thinnest at midlength and an apically bulbous aedeagus [Fig. 22c];
and Lestodiplosis syringopais has an acutely triangular gonocoxal lobe, a thin, tapered gonostylus and slender,
cylindrical, apically-rounded aedeagus [Fig. 23j]. Females of these three species were described well enough to dif-
ferentiate only L. oomeni. Larvae are unknown for L. lacciferi.
Biology. Larvae of this species prey on the scale insect Kerria javana (Barnes 1935; Harris 1968).
Geographical distribution. This species was found 7-ii-1933 in Malaysia at Bukit Badong and Kuala Lumpur
(Barnes 1935), and in Java at an unknown locality and date (Harris 1968).

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 41
FIGURES 20a–s. Leefmansiella pandani: a–c: galls on Pandanus nitidus in Java: a (Plate IX of Leemans 1921); b (Fig. 27 of
DvLR & DvL (1926)); c, whole plant; d–f, pupa: d–e (ventral); f (lateral); g–i: larva: g–h, (ventral); i, terminal segments; j, tarsal
claw and empodium (female); k–o: male: k, wing; l, palpus; m, antenna; n, terminalia, gonostyli (detached, dorsal); o, 9th flagel-
lomere; p–s, female: p, whole insect (dorsal); q, 9th flagellomere; r, cerci and hypoproct (dorsolateral); s, antenna (second flagel-
lomere misshapen). Figs a, e–g, i, p from Leefmans (1921), Figs c, d, h by Purnama Hidayat and Melinda Agustina, 2018.

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FIGURES 21a–c. Lestodiplosis lacciferi: a, wing; b, 3rd male flagellomere; c, 3rd female flagellomere. Fig. 21d. Lestodiplosis
aonidiellae (from South Africa): d, male terminalia and gonostylus (dorsal). Figs from Harris (1968).
Lestodiplosis oomeni Harris
[Figs 22a–o]
Lestodiplosis oomeni Harris, 1982: 20.
Material. Holotype male, 5 paratypes males, 9 females and 10 larvae (deposited in BMNH ((E)251546–59), USNM
(a19250–2) and RMNH) were collected and reared from on Calacarus carinatus (Green) (Acarina: Eriophyidae)
feeding on Camellia sinensis (L.) in Indonesia (Harris 1982). We examined the USNM paratypes: Nos 19250 (male,
collected 16-xii-1977), 19251 (female, 11-xi-1977) and 19252 (larva, 16-xii-1977). Those specimens were mounted
in Hoyer’s medium. Although we could observe what we needed to see on these specimens, this impermanent me-
dium is gradually crystallizing, and the specimens will eventually become unrecognizable.
Description. Adult. Wing length (arculus to apex) 0.8 mm in male, 1.0 mm in female; R5 nearly straight, reach-
ing wing apex; M4 absent; Cu1 evident [Fig. 22b]. Head [Fig. 22a] with 11-14 frontoclypeal setae, axillary palpus
3-segmented, occipital protuberance short, eye bridge 4-5 facets long, facets sparse opposite antennal bases. Anten-
nae with 12 flagellomeres.
Male. Flagellomere with three separate whorls of looped circumfila [Figs 22d, e]. Terminalia [Fig. 22c]: gono-
coxite narrow, mesobasal lobes low, rounded, setulose; gonostylus short, narrowest near midlength; aedeagus flared
near midlength, prominently narrowed below globular tip; cerci ovoid; hypoproct bulging and conspicuously setu-
lose on ventral half, broadly convex apically, with two apical setae.
Female. Flagellomeres: circumfila consisting of two horizontal whorls of short-looped bands, longitudinally
linked with two simple vertical bands [Figs 22f, g]. Terminalia: cerci broadly ovoid, setose along length with field
of about 15 closely approximated short and thick distal setae [Fig. 22h].
Pupa unknown.
Larva. Length 1–1.5 mm long. Head [Fig. 22i]: head capsule triangular, longer than wide, posterolateral
apodemes nearly as long as capsule; antennae thin, tapered, curved forwards, longer than capsule width. Sternal
spatula absent. Meso- and metathorax each with two pseudopods, abdominal segments 1–7 with three pseudopods,
pseudopods with sparse setulae, greatly narrowed before tip [Figs 22j, k, m, o]. Papillae as in Figs 22i–o, dorsal
papillae with all six setae of same length. Terminal segment with 8 terminal papillae with long setae, a number of
additional socketed setae and many setulae [Figs 22n, o].
Remarks. To differentiate males from its two known congeners from Indonesia, see under L. lacciferi above.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 43
Larvae of L. oomeni and L. syringopais were both described. In L. oomeni the head is acutely triangular with anten-
nae that are longer than head width, all dorsal papillae bear setae of similar length, and the terminal segment has
setae additional to those of the eight terminal papillae. In L. syringopais the head is acutely triangular with antennae
shorter than head width, the outer pair of dorsal papillae have setae that are much longer than the two inner pairs,
and the terminal segment has six papillae with or without setae [possibly broken off] and no additional vestiture
[Fig. 23q].
FIGURES 22a–o. Lestodiplosis oomeni: a–e, male: a, head; b, wing; c, terminalia (dorsal); d, third flagellomere; e, last flagel-
lomere; f–h, female: f, third flagellomere; g, last two flagellomeres; h, cerci (ventral); i–o, larva: i, head through mesothorax
(dorsal); j, abdominal (lateral); k, head, thorax (lateral); l, prothorax (ventral); m, last three abdominal segments (lateral); n,
same (dorsal); o, same (ventral). Figures from Harris (1982).
Biology. Type specimens were collected and reared from a population preying on the purple tea mite Calacarus
carinatus (Green) (Acarina: Eriophyidae) that was feeding on the tea plant Camellia sinensis (L.) in Java (Harris

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1982). Additionally, the larvae were observed preying on the pink tea mite Acaphylla theae (Watt) (Acarina: Erio-
phyidae) and the scarlet or false tea mite Brevipalpus phoenicis (Geijskes) (Acarina, Tenuipalpidae) (Harris 1982).
Geographical distribution. This species is currently known from Java: Gambung, near Bandung, 16 &
22.xii.1977; Pasir Nangka, Cianjur Regency, south-west of Bandung, 11.xi.1977; Tambakan, Tangkuban Perahu,
north of Bandung, 18.vi.1977 (Harris 1982).
Lestodiplosis syringopais (Hering)
[Figs 23a–q]
Coprodiplosis syringopais Hering 1931: 59.
Lestodiplosis syringopais (Hering): Gagné 1973: 504, new combination.
Type series. Hering (1931) described the male, female, pupa and larva from a population co-occurring with a popu-
lation of the mining moth Phyllocnistis nepenthae Hering feeding on the pitcher plant Nepenthes tobaica Danser,
collected along a road between Balige and Siborong-Borong, Lake Toba, Sumatra, 17-iv-1929. The types are not
present in the Museum für Naturkunde, Humboldt-Universität, Berlin (ZMHB, Jenny Pohl, pers. comm. 11-v-2017)
and might be lost.
Description (based on description and illustrations in Hering (1931)). Adult. Flagellomeres 12. Palpus 3-seg-
mented, first segment half-length of second, third slightly longer than second [Fig. 23b]. Wing with R5 evenly
bowed, reaching beyond wing apex; R1 about half wing length; M4 and Cu1 forming fork; RS positioned less than
halfway between arculus and end of R1 [Fig. 23a]. Tarsal claws thin, simple, empodia nearly as long as claws [Figs
23d, e].
Male. Size 1.2 mm. Flagellomeres with spheroid proximal and ovoid distal node; proximal node with single
whorl of looped circumfila, distal with two, proximal loops reaching mid length of internode, middle loops barely
reaching bases of distal loops, distal loops reaching end of neck [Figs 23f–h]. Terminalia [Figs 23i–k]: gonocoxite
narrow, mesobasal lobe long, acutely pointed; gonostylus narrow, tapering; cerci and hypoproct blunt apically; ae-
deagus evenly cylindrical, sinuous in lateral view.
Female. Flagellomeres constricted at mid length, with two whorls of short-looped, bowed, longitudinally linked
circumfila [Figs 23l, m]. Cerci ovoid, hypoproct about one-third cercal length [Fig. 23k].
Pupa. Little in a diagnostic way is described or drawn of this stage in Hering (1931) except that the abdominal
spiracles of the fourth through seventh segment appear long and tube-like.
Larva. Little is diagnostic for the larva, also, although the drawing of the head appears remarkably more ac-
curate than any other part of the body. [Fig. 23n]. Other than that, characters that may be diagnostic are: outer pair
of dorsal thoracic papillae have long setae and two middle pairs much shorter [Figs 23o, p]. The terminal segment
shows 6 or 8 papillae but without the usual setae, which could have broken off [Fig. 23q].
Remarks. See comparison among adults of congeners occurring in Indonesia under L. lactifera and between
this species and L. oomeni under the latter species.
Biology. The life cycle of this species as the predator of a mining moth needs to be confirmed. Larvae of the
type population were found on pitcher plants Nepenthes tobaica (Nepenthaceae) with a dense, superficial network
of mines on their pitchers caused by caterpillars of the mining moth Phyllocnistis nepenthae (Lepidoptera: Gracil-
lariidae: Phyllocnistinae) at Lake Toba, Sumatra (Hering 1931). Gall midge larvae pupated in a silky cocoon on the
pitcher. The assumption that L. syringopais is a predator of P. nepenthae was based on a large abundance of both
species, the gall midge and the mining moth (Thienemann pers. comm. in Hering (1931)), rather than an observation
of predation. On the other hand, Beaver (1979) found in Penang, Malaysia, several larvae that he identified as L.
syringopais in pitchers of N. albomarginata and N. gracilis that had no association with any leaf miner. The larvae
were seen to attack and feed on small larvae of Endonepenthia schuitemakeri (Diptera: Phoridae) in the liquid at the
bottom of the pitchers. When mature, the larvae of the gall midge were seen to crawl up the side of the pitcher and
construct silken cocoon-like coverings in which they pupated. We have not seen Beaver’s specimens.
Geographical distribution. The confirmed occurrence of this species is currently confined to its type locality
at Lake Toba, Sumatra, 17-iv-1929.

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FIGURES 23a–q (from Hering (1931)). Lestodiplosis syringopais: a–c, adult: a, wing; b, palpus; c, tarsus of midleg; d–j, male:
d, last tarsal segment (lateral); e, last tarsal segment (ventral); f scape, pedicel and first flagellomere; g, flagellomere (with neck
of previous member); h, last flagellomere; i, terminalia (lateral); j, terminalia (dorsal); k–m, female: k, postabdomen (lateral); l,
flagellomere; m, scape to first flagellomere; n–q, larva: n, head; o, second thoracic segment; p, third thoracic segment; q, larval
terminal segment.
Megommata leefmansi (Nijveldt)
[Figs 24a–j]
Coccomyza leefmansi Nijveldt in Barnes & Nijveldt (1954: 92).
Megommata leefmansi (Nijveldt): Harris (1968: 455), new combination.
Type series. Holotype male, reared by Tjoa Tjien Mo from larva feeding on eggs of scale insect Pulvinaria polygo-
nata Cockerell (Hemiptera: Coccidae) on Citrus sp., Bogor, Indonesia, x-1952, Cecid. 525 of Nijveldt collection

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(Wageningen, The Netherlands). Paratype males (Cecid. 522–524), paratype females (Cecid. 525–534), same data
as holotype. Other material: males (Cecid. 8494–8498), females (Cecid. 8499–8500), Barnes collection (BMNH),
same data as holotype. We have not seen this material.
FIGURES 24a–j. Megommata leefmansi: a, male head; b, tarsal claw of foreleg; c, tarsal claw of mid or hind leg; d, palpus; e,
wing; f, female terminalia (lateral); g, 3rd male flagellomere; h, gonostylus (dorsal); i, male terminalia (dorsal); j, 1st and 2nd
female flagellomeres. Figs a, d–j from Harris (1968), Figs b, c from Nijveldt (1954).
Description. slightly modified from original description by Nijveldt (1954) and by Harris (1968) who exam-
ined types deposited in BMNH. Male. Length about 1.2 mm. Abdomen bright red, thorax dark ochreous. Eyes not
quite divided laterally, still contiguous, occipital protuberance absent [Fig. 24a]. Flagellomeres 12, first and second
fused, gynecoid: consisting of barrel-shaped node and short neck, circumfila simple [Fig. 24g]. Palpus 4-segmented
[Fig. 24d]. Wing: R1 joining C slightly anterior to half-length of wing, Rs absent, R5 joining C slightly posterior to
wing apex, M fold apparent, M4 and Cu1 forming fork [Fig. 24e]. Tarsal claws bent at right angle near distal third,
those of forelegs bearing strong tooth [Fig. 24b], remainder simple [Fig. 24c]. Terminalia [Figs 24h, i]: gonocoxite
stout, gonocoxal apodemes separated basally; gonostylus tapered, completely setulose; cerci rounded apically; hy-
poproct entire, broadly rounded; aedeagus tapered, as long as long as gonocoxite.
Female. Length about 1.2 mm. Flagellomeres 12, first and second fused, consisting of cylindrical node and
short neck, slightly shorter than in male [Fig. 24j]. Ovipositor: short, barely protrusible; cerci separate, ovoid in
lateral view; hypoproct 2/3 length cerci; 10th sternite in form of large, setose lobe, pointed distally in lateral view
[Fig. 24f].
Pupa. Brown, length 1.3–1.9 mm.
Larva. White with yellow dorsal stripe. Length 1.5–1.7 mm.
Biology. Larvae were found feeding in ovisacs of the scale insect Pulvinaria polygonata.
Geographical distribution. This species has been found thus far only at its type locality in Bogor, Java, x-
1952.

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Oligoxenomyia Felt
[Figs 25a–n]
Oligoxenomyia Felt, 1927b: 386.
Type species: Oligoxenomyia radicis Felt, by original designation.
Diagnosis. This monotypic genus belongs to the supertribe Cecidomyiidi and is notable for the large, weakly sclero-
tized gonostylar tooth that is set perpendicularly on the gonostylus. It remains at present unassigned to tribe.
Description. Adult. Wing with R5 straight, joining C slightly behind wing apex; C broken at juncture with R5;
Rs not present; M4 and Cu1 forming fork [Fig. 25a]. Flagellomeres 12; in male foreshortened with two indistinct
nodes, proximal node with one, distal with two whorls of circumfila, all with loops shorter than distance between
them, internode partially setulose [Fig. 25b]; in female cylindrical, circumfila of two horizontal and two vertical
appressed bands [Fig. 25i]. Palpus 3-segmented [Figs 25g, h]. Tarsal claws simple, strongly bent near basal third,
empodia shorter than claws [Fig. 25f]. Male terminalia [Figs 25c–e]: gonostylus broad, basal 2/3 asetose and setu-
lose, distal third with sparse, short setae and no setulae, tooth set at strong angle to shaft, large, weakly sclerotized,
carinate, with scattered short setae; aedeagus narrow in lateral view, about as long as cerci and hypoproct, recurved
dorsally, rounded apically, with apical asetose sensilla.
FIGURES 25a–n. Oligoxenomyia radicis: a, wing; b–g, male: b, 3rd flagellomere; c, terminalia (mesolateral); d, gonostylus
(ventral); e, gonostylus (dorsal); f, claw and empodium; g, palpus; h–k, female: h, palpus; i, 3rd flagellomere; j, ovipositor (lat-
eral); k, cercus and hypoproct (lateral); l–m, pupa: l, antennal horns and cervical papillae (ventral); m, prothoracic spiracle; n,
aerial root gall on Freycinetia imbricate (Fig. 24 of DvLR & DvL (1926)).

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Ovipositor [Figs, 25j, k] retractable, cerci round in lateral view, setose, setulose, with pair of prominent dor-
soapical, thick, blunt setae much longer than surrounding setae; hypoproct about third length of cerci, rounded api-
cally with pair of distal setae. Pupa with large antennal horns [Fig. 25l], prothoracic spiracle long [Fig. 25m].
Oligoxenomyia radicis Felt
[Figs 25a–n]
Oligoxenomyia radicis Felt, 1927b: 386.
Material examined. Syntypes: 6 males, 10 females, 2 pupae reared from aerial root galls on Freycinetia sumatrana
Hemsl. (as F. valida Ridl.) (Pandanaceae), gall No 45 in DvLR & DvL (1926) collected at Bolang, near Bogor, Java,
iii-1924. Felt #a3391). Felt’s types are mounted on 4 slides, all uncleared, whole specimens, adults in lateral posi-
tion, pupae in dorso-ventral. All characters preserved in at least one specimen.
Description. Adult. Palpal segments 3, segments progressively longer [Figs 25g, h]. Tarsal claws bent at right
angle close to basal third, tip pointed, bowed, empodia reaching bend of claw [Fig. 25f].
Male. Wing length 2.7 mm (2.5–2.9), width 1.0 mm (0.9–1.2) [Fig. 25a]. Flagellomeres [Fig. 25b]. Terminalia
[Figs 25c–e].
Female. Wing length 3.5 mm (3.3–3.7), width 1.4 mm (1.3–1.4). Flagellomeres slightly constricted at mid-
length [Fig. 25i]. Terminalia [Figs 25j, k].
Pupa. Antennal horns long, narrow, broadly rounded in ventral view [Fig. 25l]. Prothoracic spiracle long, ta-
pered, bowed at distal third, with small spikes distally, trachea ending below distal end [Fig. 25m]; spiracles on
abdominal segments sessile. Abdominal segments dorsally with dense field of large, simple spikes.
Larva unknown.
Biology. This species causes an aerial root gall on Freycinetia sumatrana Hemsl. (Pandanaceae) described by
DvLR & DvL (1926, gall No 45, Fig 24. [Fig. 25n]). Irregular swellings are partly coalescent and attached to the
root with broad bases. The surface is covered with numerous short, brown hair-like scales. The interior contains
many long and tubular larval chambers, crossing the longitudinal axis of the root.
Geographical distribution. This species is known from Java: Mt Ciampea, near Bogor, alt. 200 m, 1920; Mt
Burangrang, alt. 1100 m, vii-1921; Bolang, near Bogor, alt. 950 m, iii-1924 (DvLR & DvL 1926).
Orseolia Kieffer & Massalongo in Indonesia
This is a genus of 28 known species mainly of tropical Asia, but three from tropical Africa, one from the Mediter-
ranean Region and one from Japan and Korea. One of the more important species is O. oryzae, a pest of rice in
Indonesia and elsewhere. All form leaf sheath galls on grasses (Poaceae). One remarkable aspect of these galls is
that they continue to lengthen after the larva turns into a pupa. DvLR & DvL (1926) attribute this to the continued
absorption of water by the plant. Seven species are known from Indonesia and all were part of a generic revision
by Gagné (1985). They are fairly large gall midges with a wing length of 2.3-4.5 mm, the R5 vein bowed to join C
beyond the wing apex [Fig. 31h]. Heads lack an occipital protuberance. The last flagellomere usually has a conical
extension at its apex [Figs. 27f, 31j]. Male flagellomeres have two nodes, the proximal spherical, with one whorl
of looped circumfila, the distal prolonged, slightly constricted at midlength, with two whorls of looped circumfila.
Tarsal claws are simple, narrow, bent at distal third, and are as long as the empodia [Fig. 31g]. Female cerci are
large, ovate and covered with setae among which is a pair of thick, blunt, apical setae [Fig. 31e]. Some of the best
taxonomic characters in the genus are differences in male terminalia and pupal antennal horns. Larvae are known for
only two species but they do have a spatula. We do not repeat below all details of descriptions from Gagné (1985)
but present additional, new morphological drawings as well as illustrations of galls from DvLR and DvL (1926).

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 49
Orseolia apludae (Felt)
[Fig. 26a–b]
Orseoliella apludae Felt, 1920: 8
Orseolia apludae Felt: Gagné (1973: 506), new combination; Gagné (1985: 137), revision.
Material examined. Lectotype male, designated by Gagné (1985: 138) and paralectotype female, reared from a
leaf sheath gall on Apluda mutica (as A. varia) at Coimbatore, Tamil Nadu, India, 1-i-1917, Felt type #a2974. Male,
female, pupal exuviae, same locality as types, collected 1921.
Description. Male terminalia as in Fig. 26b.
Pupa, larva unknown.
Biology. This species causes a gall on Apluda mutica L. (Poaceae) described simply as a leaf sheath gall by
Felt (1920) from India, but later extended by DvLR & Dvl (1926, gall No 52, Fig. 28 [Fig. 26a]) who found it on
the same host in Indonesia. The gall protrudes from the sheath of the youngest leaf. The leaf blade remains unde-
veloped. The sheath is shaped into a tube enclosing and supporting the growth tip. The gall grows out of the older
leaves and buds which develop after the infection and are long and thin, with a slender apex.
Geographical distribution. Orseolia apludae was collected and reared at Coimbatore, Tamil Nadu, India,
1-i-1917 (Felt 1920). The galls were found in Indonesia in Salatiga, Java, alt. 600 m, vi-1909 and Kamal, Madura
Island, v-1910 (DvLR & DvL 1926).
FIGURES 26a–b. Orseolia apludae: a, gall on Apluda mutica (arrows indicate galls; Fig. 28 of DvLR & DvL (1926)); b, male
terminalia (dorsal, cerci omitted, from Gagné 1985). Figs 26c–d. Orseolia graminis: c, gall on Ottochloa nodosa (arrow indi-
cates gall; Fig. 38 of DvLR & DvL (1926)); d, last two female flagellomeres (from Kieffer & DvLR (1910)).
Orseolia caulicola Gagné
[Figs 27a–g]
Parallelodiplosis javanica Felt, 1921d: 91.
Orseolia caulicola Gagné, 1973: 507, new name for P. javanica, (junior secondary homonymy); Gagné 1985: 139, revision.
Material examined. Lectotype male designated by Gagné (1985: 139), and male and 2 female paralectotypes,
reared from leaf sheath gall on Sacciolepis indica (L.) Chase (as Panicum indicum L.) (Poaceae) in Bogor, Java,
Indonesia, vi-1920, Felt type #a3150. We examined the males which are mounted as whole, cleared insects on a
single slide, with all characters preserved except the barely observable circumfila.
Description. Adult. Wing as in Fig. 27b. Palpus 4-segmented [Fig. 27c]. Tarsal claws simple, narrow, bent at
distal third, as long as empodia [Fig. 27g].

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Male. Flagellomeres as for genus [Fig. 27e], last flagellomere with apical extension [Fig. 27f]. Terminalia [Fig.
27d]: gonostylus short, strongly tapered, setulose except immediately next to distal tooth where carinate; cerci
triangular, wide; hypoproct conical, slightly tapered; aedeagus narrow, as long as hypoproct, with several asetose
sensilla on distal half.
Female. Tergum 10 with numerous short lateral setae, more similar to O. paspali than to less densely setose in
O. oryzae [Fig. 31f], cerci as in O. oryzae [Fig. 31e] (Gagné 1985).
Pupa, larva unknown.
Remarks. Orseolia caulicola is morphologically similar to O. oryzae and its close relatives O. eragrostisae
(Mani), O. similis (Mani), O. difficilis Gagné and O paspali (Gagné 1985).
Biology. This gall midge causes a leaf sheath gall on Sacciolepis indica (L.) Chase described by DvLR & DvL
(1926) as Hymenachne indica Buese. (= Panicum indicum L), gall No 59, Fig 36. [Fig. 27a]). The slender white
or greenish gall tubes develop chiefly in the inflorescences. Leaf buds grow between the flowers in the axils of the
leaves of the flower stalks and the leaf ends of these buds change into galls. The galls occur also in axillary buds on
the main stems and side branches.
Geographical distribution. This species is known from its type locality in Bogor, Java, where it was collected
vi-1920 (DvLR & DvL 1926).
FIGURES 27a–g. Orseolia caulicola: a, gall on Sacciolepis indica (arrows indicate galls; Fig. 36 of DvLR & DvL (1926),
gall No 59); b–g, male: b, distal part of wing; c, palpus; d, terminalia (dorsal); e, 3rd flagellomere (circumfila partly obscured on
specimen); f, 12th flagellomere; g, claw and empodium.
Orseolia graminicola (Kieffer & Docters van Leeuwen-Reijnvaan)
[Figs 28a–e]
Clinodiplosis graminicola Kieffer & DvLR, 1910: 127.
Pachydiplosis graminicola (Kieffer & DvLR): Kieffer (1913: 224), new combination.
Orseolia graminicola (Kieffer & DvLR): Gagné (1973: 507), new combination; Gagné (1985: 139), revision.
Type series. Syntypes, male(s) and female(s), reared from Cynodon dactylon (L.) Pers. (Poaceae) in Salatiga, Java,
iv-1909. The types are likely lost (Gagné & Jaschhof 2017).
Description. The relevant character states and figures are from Kieffer & DvLR (1910).
Adult. Length 2.5 mm in male, 3.5 mm in female.
Male. Flagellomeres [Fig. 28c] and terminalia [Fig. 29e], generally as for genus.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 51
Female. Flagellomere [Fig. 28d] as for genus.
Pupa, larva unknown.
Remarks. This species appears similar to O. oryzae. It differs from O. difficilis Gagné, which causes similar galls
on the same host plant in India, in the more strongly tapered gonostylus and the narrower hypoproct (Gagné 1985).
Biology. This species causes a leaf sheath gall on Cynodon dactylon (Poaceae) described by DvLR &DvL
(1926, gall No 52, Figs 29, 30) [Figs 28b, d]. Females deposit eggs on the upper surface of the leaves at night. Newly
hatched larvae penetrate the young leaves until they have reached the upper end of the growth tip where they live
for some time. A number of axillary buds develop which grow into small stems. Leaf sheaths become enlarged and
forced outward by the great number of buds, until a fusiform gall is produced, which is often somewhat laterally
compressed [Fig. 28b]. A leaf then develops from the primary axillary buds like a cylindrical case that encloses the
larva, in this way forming a gall chamber. The leaf sheaths open at the top and the young twigs and galls emerge
[Fig. 28d]. An aperture is formed near the top of the gall from which the adult gall midge emerges. After some time,
when the larvae have pupated, all the axillary buds, both those in which a larva lives and the uninfected ones, begin
to grow.
Geographical distribution. This species is currently known from Java: Salatiga, alt. 600 m, iv-1909; Sema-
rang, i-1910; Candi, near Semarang, alt. 100 m, viii-1912; Roban, near Pekalongan, vi-1913.
FIGURES 28a–e. Orseolia graminicola: a, male flagellomere; b, young gall on Cynodon dactylon; c, female flagellomere; d,
mature gall on Cynodon dactylon; e, male terminalia (ventral). Figs a, c, e from Kieffer & DvLR 1910), Figs b and d from DvLR
& DvL (1926, Figs 29, 30). Arrows indicate galls.
Orseolia graminis (Kieffer & Docters van Leeuwen-Reijnvaan)
[Figs 26c–d]
Courteia graminis Kieffer & DvLR, 1910: 131.
Orseolia graminis (Kieffer & DvLR): Gagné (1973: 507), new combination; Gagné (1985: 143) (revision).
Type series. Type female, now likely lost, was reared from Ottochloa nodosa (Kunth) Dandy (as Panicum nodosum
Kunth) (Poaceae) in Salatiga, Java, 10-iv-1909.
Description. Female. The only relevant descriptive characters given in Kieffer & DvLR (1910) are that the fly
is large, 3.5 mm long, and has a 2-segmented palpus. Their single drawing of the two last flagellomeres is shown
here [Fig. 26d].

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Male, pupa, larva unknown.
Biology. Kieffer & DvLR (1910) and DvLR & DvL (1926, gall No 64, Fig. 38) [Figs 26c] describe the gall as
a hairy, 8 mm long and 3 mm wide, leaf sheath on Ottochloa nodosa (as Panicum trigonum Retz.) (Poaceae). The
sheath does not grow into a long tube, but remains short, oval at base and pointed towards the tip. The leaf blade is
often rudimentary at the top of the gall or totally missing [Fig. 26c]. Galls develop at the end of the stem or out of a
side bud. The type female emerged 10-iv.
Geographical distribution. This species is known from Java: Salatiga, alt. 600 m, i-1909; Mt Ungaran, alt.
1000 m, v-1910; Bogor, Botanic Gardens, alt. 250 m, v-1920; Ngarengan, near Semarang, alt. 40 m, ii-1920; Cigom-
bong, near Bogor, alt. 400 m., iv-1925.
Orseolia javanica Kieffer & Docters van Leeuwen-Reijnvaan
[Figs 29a–g]
Orseolia javanica Kieffer & DvLR, 1910: 125; Gagné (1985: 145), revision.
Type series. Syntypes, male(s) and female(s), were reared from Imperata cylindrica Beauv. (L.) P.Beauv. (Poaceae)
in Salatiga, Java in or before 1909. The types are likely lost. Gagné (1985) examined 2 specimens, possibly of this
species, reared from I. cylindrica in Bogor, Java, 16-iv-1973, and deposited in BMNH.
Description. Relevant characters from Kieffer & DvLR (1910) are given here followed by Gagné’s (1985)
comments on the BMNH specimens (see Remarks). Adult. Size: male 4 mm, female 6 mm. Palpus 4-segmented or
3-segmented in which case last segment made up of two fused segments.
Male. Terminalia [Fig. 29b]: generally agrees with Bogor specimens illustrated by Gagné [Fig. 29g].
Female. Flagellomeres cylindrical, circumfila positioned in distal 2/3 of segment, consisting of some five trans-
verse bands connected by longitudinal bands [Figs 29d, e].
Pupa, larva unknown.
Remarks. The BMNH specimens fit the original description of O. javanica in palpi, male flagellomeres and
male terminalia [Fig. 29g] but differ in the female circumfila not being reticulate [Fig. 29f], a difference which can-
not be resolved here. Felt (1917) identified Indian specimens from the same host plant as O. javanica, but the male
terminalia as he described them have a short and much broader hypoproct exceeded in length by the aedeagus and
thus probably represent a different species (Gagné 1985).
DNA. Purnama Hidayat and Ayu Lestari, IPB University, sequenced COI of one adult specimen of O. javanica
from Cianjur, Java, and two of O. oryzae from Bogor and Cianjur – the O. oryzae sequences showed an intraspecific
similarity of 99.58% while the interspecific similarity of each of them to O. javanica was 90.31% (unpublished,
pers. comm. 8-iv-2019; we compared these three, 717bp-long sequences, in GenBank’s MegaBlast 8-iv-2019). The
low level of the interspecific similarity suggests that the two species are indeed discrete. Biology of the population
which included the sequenced adult specimen of O. javanica was described in Hidayat et al. (2020a).
Biology. The leaf gall on cogon grass I. cylindrica was described in detail by DvLR & DvL (1926, gall No
60, Fig. II-5 [Fig. 29a]) and DvLR & DvL (1910a). The first stage of gall development while larvae are growing is
superficially not obvious, but after pupation occurs the gall emerges from the center of the surrounding leaves and
develops into a long, often curved, tube which is pointed at the tip. The galls are 4–20 cm long and about 3 mm
in diameter, mostly red, seldom white. It is a common and conspicuous gall. The larva lives within a chamber that
takes up the whole length of a gall. Pupation takes place at the bottom of the chamber after which the pupa then mi-
grates to the upper part of the gall. Following adult emergence, the pupal exuviae remain attached to the emergence
hole. Cogon grass is a weed in several countries of Asia, Africa and USA (Overholt et al. 2016). The search for a
host-specific pathogen triggered an interest in biology of O. javanica, which has been studied extensively (Loan et
al. 2002; Syah & Hidayat 2019; Arini et al. 2017; Hidayat et al. 2020a).
Geographical distribution. This species occurs in West and Central Java (DvLR & DvL 1926; Loan et al.
2002; Hidayat et al. 2020b).

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FIGURES 29a–g. Orseolia javanica: a, gall on Imperata cylindrica (arrow indicates one of two galls; Fig. II-5 of DvLR &
DvL (1926)); b, male terminalia; c, 12th male flagellomere; d, 3rd female flagellomere; e, 12th female flagellomere; f, 3rd female
flagellomere; g, male terminalia (dorsal). Figs b–e from Kieffer & DvLR (1910) depicting types (now lost), Figs. f, g from
Gagné (1985) of specimens in BMNH.
Orseolia orientalis (Felt)
[Figs 30a–o]
Orseoliella orientalis Felt, 1921d: 90.
Orseolia orientalis (Felt): Gagné (1973: 507), new combination; Gagné (1985: 147), revision.
Material examined. Lectotype male, designated by Gagné (1985: 148), paralectotypes female and pupa, reared
from leaf sheath gall on Oplismenus compositus P. Beauv., Bogor, Java, vi-1920, Felt type #a3152. The types are
whole, cleared mounts on two separate slides, remounted by RJG, entire except for most circumfila being dissolved
and mostly unrecognizable, and the 2 and 3 terminal male flagellomeres are missing.

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FIGURES 30a–o. Orseolia orientalis: a–b, galls on Oplismenus compositus (Figs 32, 33 of DvLR & DvL (1926)): a, young gall
(ga = gall); b, mature gall (gb = gall breaking point); c, wing apex; d, tarsal claw and empodium; e, palpus; f–i, male: f, termina-
lia (dorsal); g, 3rd flagellomere; h, 6th flagellomere; i, 12th flagellomere; j-k, female: j, 3rd flagellomere; k, 10th tergum and cerci
(dorsolateral: arrow indicates pair of thick setae on cercus); l–o, pupa: l, dorsal spines on 3rd abdominal tergum; m, prothoracic
spiracle (lateral); n, prothoracic spiracle (ventral); o, anterior segments (lateral). Figs j–l from Gagné (1985).
Description. Adult. Wing length 3.1 mm in male, 3.5 in female (Gagné 1985). Vein R1 slightly bowed, joining
C posteriad of wing apex [Fig. 30c]. Palpus 4-segmented [Fig. 30e]. Tarsal claws as for genus [Fig. 30d].
Male. Flagellomeres [Figs 30g, h, i] with short circumfilar loops. Terminalia [Fig. 30f]: gonocoxite narrow
beyond bulbous base; gonostylus long, cylindrical, widened just proximad of tooth, setulose at basal 1/4, carinate
beyond; cerci tapered caudally; hypoproct much longer than cerci, convex apically; aedeagus longer than and nearly
as wide as hypoproct, conical apically, with asetose sensilla terminally.
Female. Flagellomere nodes narrowed at basal third, circumfila slightly irregular [Fig. 30j]. Tergum 10 with
setae along caudal margin, cerci ovoid, setae narrower towards apex, thick sensory setae slightly longer than adja-
cent setae [Fig. 30k].
Pupa. Antennal horns bifid, one part longer, wide and serrated apically, the other shorter, narrow, pointed and
smooth [Fig. 30o]. Prothoracic spiracle long, slightly tapered and bowed, widened and serrate at apex, trachea
reaching apex [Figs 30m, n]. Tergal spines short and broad [Fig. 30l].
Larva unknown.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 55
Remarks. This distinct species differs from the other congeners in the long, narrow gonostylus and wide ae-
deagus and shape of the pupal antennal horns. Two other, differently shaped, leaf sheath galls on O. compositus by
unknown Cecidomyiidae. occur in Indonesia (DvLR & DvL 1926, galls No 56, 57).
Biology. This species causes a leaf sheath gall on Oplismenus compositus (Poaceae), described by DvLR &
DvL (1926, gall No 55, Figs 32, 33) [Figs 30a, b]. The leaf sheath of the youngest leaf is changed into a tube emerg-
ing among other, unchanged leaves [Fig. 30a]. The leaf blade is developed though not quite normal at the end of the
tube, and in this case the occurrence of the leaf blade is a feature of the gall, although that feature occurs occasion-
ally with leaf sheath galls of other species. When older, the gall tube grows very quickly. There is a weak spot about
a third of the total length from the tip of the gall where the gall breaks in two, allowing the gall midge to emerge
[Fig. 30b].
Geographical distribution. This species is currently known only form its type locality in Java: Bogor, alt. 250
m, v and vi-1920 (Felt 1921d; DvLR & DvL 1926).
Orseolia oryzae (Wood-Mason)
[Figs 31a–j]
Cecidomyia oryzae Riley, 1881: 149, nomen nudum.
Cecidomyia oryzae Wood-Mason (in Cotes 1889: 103).
Pachydiplosis oryzae (Wood-Mason): Felt (1921a: 16), new combination.
Orseolia oryzae (Wood-Mason): Gagné (1973: 507), new combination; Gagné (1985: 148), revision.
Material. Syntypes, female, pupae, pupal exuviae (originally deposited in the National Zoological Collection, Cal-
cutta, India, now presumed lost (Gagné 1985)), were collected and reared from rice at Kurruckpore, Munger, Bihar,
India, x-1880. Previously Gagné (1985) examined specimens from India, China, Thailand and Indonesia.
Description. Both sexes, the pupa and larva were described in Harris & Gagné (1982) and further in Gagné
(1985). Some of their drawings are reproduced here. Wing [Fig. 31h]: length 2.9–3.3 in male, 3.0–4.1 in female.
Palpus usually 4- but occasionally 3-segmented [Fig. 31i]. Tarsal claws as in Fig. 31g. Male flagellomeres as in Fig.
31c. Female flagellomeres [Figs 31d, i, j] cylindrical. Male terminalia [Fig. 31b]: gonostylus short, tapered, nar-
rowest subapically, almost entirely setulose; hypoproct long, narrow, rounded apically; aedeagus narrow, tapered,
conical apically, longer than hypoproct; cerci tapered caudally. Female tergum 10 with few to many lateral setae but
absent mesally; cerci ovoid [Fig. 31e].
Remarks. The female 10th tergite in O. oryzae has several lateral setae, that are more numerous in the Indian
[Fig. 31f] than in the Indonesian population [Fig. 31e] (Gagné 1985). The difference between the Indian and the
South East Asian (Indonesian and Thai) populations is evident also in the COI sequence (see below) and it is pos-
sible that there are more than one species of Orseolia spp. feeding on rice in Asia. Orseolia oryzae, as defined cur-
rently, is the only Orseolia that feeds on rice (Gagné 1985). It is morphologically similar to O. caulicola, O. difficilis
Gagné, O. eragrostisae (Mani), O. paspali (Gagné 1985) and O. similis (Mani).
DNA. Fragments of several genes of O. oryzae have been sequenced, including the taxonomically important
COI of which 13 sequences are available from Thailand (GenBank accession numbers KX447431–KX447443) and
two verified sequences from India (KM888183, KC506565) (GenBank, accessed 27-iii-2019). The pairwise dif-
ference in COI between the Indian (internal divergence of 0.37%) and the Thai (internal divergence of 0–3.25 %)
sequences is 6.22–7.05%, suggesting an existence of two separate species (Janique et al. 2017). Our comparison of
two unpublished Indonesian O. oryzae sequences (see more information under O. javanica) (internal divergence of
0.42%) showed a difference of 0.91–3.57% to the Thailand population and 6.49–7.78% to the Indian population,
suggesting that the Indonesian and Thai populations belong to the same species which is different to the nominal
Indian species.
Biology. This species is a serious pest of rice Oryza sativa L. (Poaceae), causing a leaf sheath gall called a “sil-
ver shoot” [Fig. 31a] which emerges from the leaves and is white, 5–15 cm long and 2–3 mm in diameter (DvLR &
DvL (1926, gall No 61).
Geographical distribution. The range comprises Pakistan, India, Nepal, Sri Lanka, Bangladesh, Myanmar,
Thailand, Laos, Vietnam, Cambodia, Malaysia, Indonesia, China and the Philippines (Plantwise Knowledge Bank
2019). In Indonesia, the last comprehensive report found O. oryzae to be common on all main islands (Java, Su-

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56 · Zootaxa 4847 (1) © 2020 Magnolia Press
matra, Kalimantan, Sulawesi) except Maluku and Irian Jaya [now provinces of West Papua and Papua] (Hidaka &
Budiyanto 1984).
FIGURES 31a–j. Orseolia oryzae: a, gall on Oryza sativa in India (arrow indicates gall; photo from NICRA, 2011); b, male
terminalia (dorsal); c, 3rd male flagellomere; d, 3rd female flagellomere; e, female 10th tergite and cerci (dorsolateral; arrow indi-
cates pair of thick setae on cercus; Karawang, Java, Indonesia); f, female 10th tergite (Cuttack, Odisha, India); g, claw and empo-
dium; h, wing; i, female head; j, female flagellomeres 3–12. Fig. b from Harris & Gagné (1982), Figs c–f from Gagné (1985).
Orseolia paspali (Felt)
[Figs 32a–k]
Parallelodiplosis paspali Felt, 1921d: 92.
Orseolia paspali (Felt): Gagné 1973: 507, new combination.
Material examined. We examined Felt’s male lectotype, des. by Gagné (1985: 150) and 1 male, 2 female and 1
pupal paralectotypes reared from a leaf sheath gall on Paspalum scrobiculatum L. (as P. scrobiculatum Hook.) in
Bogor, Java, Indonesia, vi-1920, Felt type #a3151. The lectotype is mounted whole, cleared, with terminal flagel-
lomeres missing, gonocoxites partially folded, gonostylus mounted at angle, wings folded, legs intact, head partially
shriveled. The male paralectotype, mounted with the lectotype on the same slide, has all flagellomeres preserved
and all tarsi missing.
Description. Wing length 2.5–2.9 mm in male, 2.8–3.5 mm in female Gagné (1985). Palpus 4-segmented,
palpiger present [Fig. 32k]. Tarsal claws [Fig. 32c]. Male flagellomeres [Fig. 32b, d]. Male terminalia [Figs 32e–j]:

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 57
gonostylus wide, distal edge constricted at midlength, setulose on basal half, carinate beyond, tooth large; hypoproct
long, wide; aedeagus narrow, as long as hypoproct; cerci triangular.
Remarks. Orseolia paspali is similar to O. oryzae from which it can be separated by the gonostylus that is con-
stricted at midlength and setulose on its basal half [Figs 32e, f], as opposed to gradually tapered and almost entirely
setulose [Fig. 31b]. The female tergum 10 has more numerous setae than O. oryzae [Fig. 31f] (Gagné 1985).
Biology. This species causes a 10–15 cm long and 2 mm wide, tubular leaf sheath gall, similar to Orseolia
oryzae, on Paspalum scrobiculatum L. (Poaceae) (DvLR & DvL (1926, gall No 66, Fig. 40 [Fig. 32a]) and P. con-
jugatum P.J.Bergius (Gagné 1985).
Geographical distribution. Orseolia paspali occurs in Indonesia and India. In Indonesia it was found only
in the type locality Bogor, alt. 250 m, v-1920, on P. scrobiculatum (DvLR & DvL 1926). In India it is common,
with the following confirmed localities (all from Gagné 1985): Kannur, Kerala, 20–30-viii-1928, on P. scrobicula-
tum; Allahabad, Uttar Pradesh, 1973, as laboratory colony on P. scrobiculatum; Orissa, Cuttack, Odisha, 1979, on
Paspalum sp.; Jabalpur, Madhya Pradesh, 1948, on P. scrobiculatum; Raipur, Bhopal, Madhya Pradesh, 1981, on
Paspalum sp.; Goa, 1981, on Paspalum sp.
FIGURES 32a–k. Orseolia paspali: a, gall on Paspalum scrobiculatum (arrow indicates gall; Fig. 40 of DvLR & DvL (1926));
b–k, male: b, 12th flagellomere; c, claw and empodium; d, 6th flagellomere; e, gonostylus (dorsal); f, gonostylus (ventral); g, cerci
(dorsal); h, hypoproct (dorsal); i, aedeagus (dorsal); j, terminalia (dorsal); k, palpus including palpiger.
Procontarinia matteiana Kieffer & Cecconi
[Figs 33a–i]
Procontarinia matteiana Kieffer & Cecconi, 1906: 135
Material. Male, female, pupal and larval syntypes were reared from pustulate leaf galls on Mangifera indica L. in
the Botanical Gardens, Palermo, Sicily, Italy, v-1906 (Kieffer & Cecconi 1906). The syntypes are presumed lost
(Gagné & Jaschhof 2017). One of us (PK) visited the Botanical Gardens in Palermo, ix-2018, but found only young,
recently planted, mango trees with no galls on their leaves. Harris (1966) described adults from an undisclosed
country and locality, possibly India. The presence of P. matteiana in Indonesia is based on the presence of galls
recorded by DvLR & DvL (1926, gall No. 801).
Description (based on Kieffer & Cecconi (1906) and Harris (1966). Adult: Wing: 1.5–2.0 mm long, R1 joining
C at basal half, R5 slightly curved, joining at near wing apex [Fig. 33i]. palpus 4-segmented [Fig. 33g]. Antenna
with 12 flagellomeres; in male binodal, internode very short, each node with whorl of looped circumfila [Fig. 33f];

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in female nodes cylindrical, not longer than wide, neck shorter than wide, circumfila simple. Tarsal claws according
to Harris (1966) toothed on some, possibly all legs. Kieffer & Cecconi (1906) recorded simple claws, but teeth in
other Procontarinia spp. are very thin so conceivably were overlooked by them. Male terminalia: gonocoxite long,
narrow, cylindrical except for pronounced obtusely triangular mesobasal lobe; gonostylus narrow, curved medially;
aedeagus long, evenly broad, bearing many asetose papillae; cerci ovate, well-separated; hypoproct slightly wider
and much shorter than aedeagus, deeply bilobed [Figs 33d, h]. Ovipositor barely protrusible, cerci stout [Fig. 33e].
Pupa without antennal horns.
Larva unknown.
DNA. Three COI sequences (GenBank numbers JQ823235–JQ823237) are available for the Réunion Island
population (Amouroux et al. 2013).
Remarks. The genus Procontarinia contains 16 described (Gagné & Jaschhof 2017, Jiao et al. 2018) and over
10 undescribed (Kolesik et al. 2009, 2017) species, all restricted to mango where they feed on leaves, young stems,
flowers or fruit. While only two species are currently reported from Indonesia (P. matteiana and P. robusta) it is
likely that many more occur there. Procontarinia matteiana differs from P. robusta in several characters of the male:
flagellomeral internode bare, narrow gonocoxite bearing acute mesobasal lobe, long and narrow gonostylus, blunt
aedeagus of even width, deeply divided hypoproct and triangular cerci in P. matteiana as opposed to flagellomeral
internode setulose, robust gonocoxite without mesobasal lobe, short and proximally wide gonostylus, tapering ae-
deagus, shallowly divided hypoproct and shallowly divided round cerci in P. robusta.
FIGURES 33a–i. Procontarinia matteiana: a–c, galls on Mangifera indica: a, Sicily, Italy (Kieffer & Cecconi 1906); b, Indo-
nesia (Fig. 577 of DvLR & DvL (1926)); c, South Africa (Augustyn et al. 2013); d, male terminalia; e, female cerci; f, 5th flagel-
lomere; g, palpus; h, male terminalia (dorsal); i, wing. Figs d–e from Kieffer & Cecconi (1906), f–i from Harris (1966).
Biology. This species causes leaf galls on Mangifera indica (Anacardiaceae) (Kieffer & Cecconi (1906, Fig.
3 [Fig. 33a]), DvLR & DvL (1926, gall No. 801, Fig. 577 [Fig. 33b])). Tiny circular galls, about 1.5–2 mm across
and 1 mm high, developed on both sides of the leaf or on young twigs. Galls are pale green at first [Fig. 33c], later
becoming pink and purple, finally black. Inside, there is a single larval chamber. The larva pupates in the gall (Au-
gustyn et al. 2013). Procontarinia matteiana is a pest not only on the Indian subcontinent, where mango was first
domesticated, but also in other countries of the world where it is cultivated now (Kolesik et al. 2017). Resistant
mango varieties (Githure et al. 1998) and the presence of natural enemies (Sankaran 1988) are currently the major
control strategies.
Geographical distribution. Procontarinia matteiana has been confirmed in Italy [Fig. 33a], India, Indonesia,
Mauritius, Réunion, Kenya and South Africa [Fig. 33c] (Gagné & Jaschhof 2017), and reported also from Oman,
Trinidad and Iran (Kolesik et al. 2017). In Indonesia it was found in Java at the following localities (DvLR & DvL
1926): Mangkang, near Semarang, vii-1910; Candi, near Semarang, alt. 50 m, iii-1911; Mt Muria, alt. 500 m, ix-
1912; Bandung, alt. 600 m, vi-1916 & vi-1918; Jakarta, v-1917; Bogor, alt. 250 m, ix-1918; Mt Pancar, near Bogor,
alt. 500 m, xii-1923.

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 59
Procontarinia robusta Li, Bu & Zhang
[Figs 34a–j]
Procontarinia robusta Li, Bu & Zhang, 2003: 148; Cai et al. 2014, as junior synonym of P. matteiana but unsubstantiated.
Material. Type male, 16 paratype males and 23 paratype females were reared from bell-shaped leaf galls on Man-
gifera indica L. in Xiamen, Fujian Province, China, 12- 16-ix-2001 (deposited in the Department of Biology, Nan-
kai University, Tianjin, China). Cai et al. (2013) mounted specimens (deposited in the South Australian Museum,
Adelaide) reared at the type locality 1.vii.2010 (6 larvae, 6 pupae, 6 males, 4 females) and in Oecusse district, East
Timor 27.x.2004. We examined specimens by Cai et al. (2013). The presence of P. robusta in Indonesia is based on
gall records by DvLR & DvL (1926, gall No. 802) and by observation of one of us (PK).
FIGURES 34a–j. Procontarinia robusta: a–b, galls on Mangifera indica: a, Java (Fig. 578 of DvLR & DvL (1926)); b, Bali,
Indonesia, 2013; c, wing; d, larval spatula; e, male 3rd flagellomere; f, female 3rd flagellomere; g, male last tarsomere; h, female
terminalia (ventral); i, male terminalia (cerci omitted; dorsal); j, male cerci and hypoproct (dorsal). Figs c, e–j from Li et al.
(2003), d from Cai et al. (2013).

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60 · Zootaxa 4847 (1) © 2020 Magnolia Press
Description (based on Li et al. (2003) and Cai et al. (2013)). Adult. Wing: length 1.4–1.5 in male, 1.7–1.9
in female, R5 straight, joining C at wing apex, RS fading towards R1, slightly closer to arculus than end of R1, M4
and Cu1 forming fork, wing fold visible [Fig. 34c]. Head: occipital protuberance present, short; palpus mostly 3-,
sometimes 4-segmented; antennae with 12 flagellomeres. Tarsal claws toothed, curved at basal third, about as long
as empodia [Fig. 34g].
Male. Flagellomeres binodal, first and second not fused, internode very short, setose, neck shorter than nodes,
asetose; each node with whorl of looped circumfila, equal in length, proximal loops reaching midlength of distal
node, distal loops reaching end of neck [Fig. 34e]. Terminalia [Figs 34i, j]: gonocoxite robust, without mesobasal
lobe; gonostylus evenly tapered from wide base to very narrow apex, tooth small; cerci nearly fused mesally; hypo-
proct longer than cerci, shallowly incised, lobes with 2–3 distal setae; aedeagus long, wide, slightly tapered, a small
notch apically, with several asetose papillae along length.
Female. Antennal flagellomeres with cylindrical nodes, necks 1/5 node length, circumfila of two vertical and
two horizontal, slightly arched bands [Fig. 34f]. Terminalia short; cerci stout, short, fused, crescent-shaped in dorsal
view; hypoproct rounded in ventral view, with pair of setae [Fig. 34h].
Pupa. Length 1.4–1.8 mm. Antennal horns large, triangular, serrated along outer edge; prothoracic spiracle
short, as long as wide, papillae on vertex without setae.
Larva. Yellowish white. Head capsule conical ventrally, hemispherical dorsally, posterolateral apodemes as
long as head capsule. Sternal spatula long, narrow, anterior part slightly wider than shaft, trilobed, central lobe sub-
stantially larger than side lobes [Fig. 34d]. Terminal segment with anus terminal, no papillae visible.
Remarks. See morphological comparison under P. matteiana.
DNA. Three COI sequences are available from the type locality in Xiamen, China (GenBank accession
JX110976–8) and one from Oecusee, East Timor (JX110979) (Cai et al. 2013).
Biology. This species causes a conical leaf gall on Mangifera indica (Anacardiaceae), described by DvLR &
DvL (1926, gall No. 802, Fig. 578 [Fig. 34a]), Li et al. (2003) and Cai et al. (2013). The gall is cylindrical to ob-
tusely conical, glabrous, reddish brown, 1 mm in diameter at the base and 1–2 mm in height. It occurs on the leaf
upper side with the necrotic leaf epidermis creating a brown wreath around the gall base. Following pupation, the
gall is left with a circular hole at its top, sometimes with the pupal skin still attached to the rim of the hole. See de-
tailed biology in Li et al. (2003) and Cai et al. (2013).
Geographical distribution. This species occurs in China, India, Indonesia and East Timor (Cai et al. 2013).
In Indonesia the galls have been found in Java, Sumatra, Sebesi Island and Bali but the species is likely to occur
through the whole Indonesian archipelago. Java: Salatiga, iii-1909; Semarang, viii-1912; Jakarta, v-1917; Cikam-
pek, vi.1924; Mt Raté, Way Lima, Lampung; Sumatra, xi.1921; Sebesi Island, i.1922; (all by DvLR & DvL 1926);
Ubud, Bali, 10-vi-2013 (Peter Kolesik, new record, [Fig. 34b]).
Psephodiplosis rubi Kolesik
[Figs 35a–j]
Psephodiplosis rubi Kolesik, 2017, in Kolesik et al. (2017: 2).
Material. Holotype, male (deposited in South Australian Museum, Adelaide (SAMA) 29–004698), reared from
leaf galls on Rubus nebulosus A.R.Bean collected in Border Range National Park, New South Wales, Australia, 18-
ix-2014 & 26-iii-2013. Other material: males, females and pupae (same data as holotype); males, females, larvae,
SAMA) reared from similar galls on Rubus moorei F.Muell. collected at Springbrook, Queensland, 28-ii & 8-xi-
2007 and Lamington National Park, Queensland, 18.iv.2013. The presence of this species in Indonesia is speculated
based on the occurrence of similar galls on the related Rubus moluccanus L. recorded by DvLR & DvL (1926, gall
No. 495, Fig. 346 [Fig. 35a]) (Kolesik et al. 2017). Rubus moluccanus occurs in South East Asia but also in Austra-
lia where its distribution range overlaps with those of R. nebulosus and R. moorei, the other hosts (Atlas of Living
Australia 2019.
Description. Adult (from Kolesik et al. 2017). Occipital protuberance absent. Palpus 1-segmented. Wing with
R5 curved to join C beyond wing apex, C with break at juncture with R5, R apparent. Tarsal claws toothed, strongly
bent at basal third; empodia reaching bend in claws [Fig. 35i].

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 61
Male. Flagellomeres binodal, with 1 circumfilum on basal node, 2 on distal [Fig. 35g]. Terminalia [Fig. 35j]:
gonocoxite with small mesobasal lobe, apodemes separated at base; gonostylus setulose basally, carinate beyond,
with comb-like tooth; aedeagus short, no longer than hypoproct, robust, with numerous asetose papillae; cerci
widely separated, ovoid; hypoproct bilobed, with many setae.
Female. Flagellomeres cylindrical, circumfila bowed, with numerous connections along vertical strands [Fig.
35h]. Cerci ovoid, with long and short setae basally, blunt, thickened setae at midlength and much shorter, pointed
setae apically [Fig. 35f]; hypoproct hemispherical, with several setae [Fig. 35f].
FIGURES 35a–j. Psephodiplosis rubi: a, galls on Rubus moluccanus in Indonesia (Fig. 346 of DvLR & DvL (1926)), b, galls
on Rubus moorei, c, galls on Rubus nebulosus, d, larval head through 1st thoracic segment, e, pupal antennal horns (dorsal), f,
end of ovipositor (ventrolateral), g, 6th male flagellomere, h, 6th female flagellomere, i, male tarsal claw with empodium, j, male
terminalia (dorsal). Galls and insects in b–j from Australia. Photos in Figs b, c by Casey R. Hall, Figs d–j from Kolesik et al.
(2017).

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Pupa. Vertex with pair of short setae cervical papillae. Antennae greatly produced anteriorly, mesal edge un-
evenly serrate, with field of asetose papillae mesobasally [Fig. 35e].
Larva. Spatular tooth triangular, shaft undeveloped [Fig. 35d]. Terminal segment with 6 short-setose papillae.
Biology. This species causes hairy spherical galls on Rubus nebulosus (Rosaceae) [Fig. 35c], R. moorei [Fig.
35b], R. moluccanus and R. alceifolius (Kolesik et al. 2017). Their appearance on R. moluccanus is as follows
(DvLR & DvL (1926, gall No. 495, Fig. 346 [Fig. 35a]). Galls are produced on the leaf underside, are spheroid,
2–6 mm across and covered with a thick yellow-brown or gray hair. Their presence on the upperside of the leaf is
marked only by the same brown hairs. On R. alceifolius galls appear not only on leaves but on leaf stalks (Kolesik
et al. (2017).
Geographical distribution. This species is known from Australia on R. nebulosus and R. moorei; its presence
in Indonesia (on R. moluccanus) and Thailand (on R. alceifolius Poir., is based on evidence of galls similar to those
found in Australia (Kolesik et al. 2017). In Indonesia it was found in Java and Sumba by DvLR & DvL (1926) at
the following localities. Java: Mt Ungaran, alt. 800 m, xii-1909; alt. 1000 m, xi-1919; Depok, alt. 100 m, viii-1918;
Mt Tugu, near Cibeber, alt. 1000 m, vi-1923; Mt Panisian, near Bogor, alt. 700 m, xii-1923; Mt Salak, alt. 1000 m,
ii-1925. Sumba: Kananggar, alt. 700 m, v-1925.
Schizomyia Kieffer in Indonesia
Only three species have been described from Indonesia of this moderately large cosmopolitan genus of 64 species
(Gagné & Jaschhof 2017; Elsayed et al. 2018b; Elsayed et al. 2019a; Elsayed et al. 2019b; da Silva dos Santos et al.
2019). The three show remarkable diversity and their generic placement will need to be evaluated when the genus as
a whole undergoes restudy. Several structures of some of these three species are odd for the genus: female cerci of
S. laporteae are no longer distinct but incorporated into the needle-like protrusible part of ovipositor; male cerci of
S. villebrunneae are incised caudally instead of being evenly convex; and gonostylar teeth are solid in S. laporteae,
missing in S. nodosa, and spiny and narrow in S. villebrunneae.
Schizomyia laporteae Felt
[Figs 36a–l]
Schizomyia laporteae Felt, 1921b: 143.
Material examined. Syntypes, female, male and pupa, the male enclosed within the pupa, reared from a large swell-
ing of the petiole or leaf base on Laportea stimulans Miq. (DvLR & DvL 1910b, gall No. 180, Fig. 74 [Fig. 36a]) at
Mt. Ungaran, Java, 10-iv-1914, Felt type #a3088. The series is on a single slide. The female is nearly complete but
palpi are lacking and the ovipositor is hidden because retracted into the end of the abdomen. The only observable
male characters are part of the head including the palpi and the distal part of one gonopod.
Description. Adult. Wing with R5 bowed at distal third, joining C at wing apex, R1 shorter than half wing length,
RS rudimentary, M4 and Cu1 forming fork. Palpus 3-segmented [Fig. 36h]. First tarsomere with small ventroapical
lobe [Fig. 36g]. Tarsal claws simple, robust, bent at distal third, longer than empodia [Fig. 36i].
Male [specimen inside pupa]. Gonocoxite with triangular ventroapical lobe, bearing long setae at apex; gono-
stylus twice as long as broad in dorsal view, slightly curved, tapered distally, uniformly setulose, with solid, trian-
gular tooth covering half width of gonostylus [Fig. 36c].
Female. Wing length 2.5 mm, width 1.0 mm. Flagellomeres cylindrical with short necks, progressively shorter,
the last abruptly so, first and second not fused; circumfila weakly bowed [Figs 36, l]. Postabdomen long, dorsobasal
lobes not visible in type; tip of needle-like apex of ovipositor completely incorporating indistinguishable cerci, with
4–5 pairs of setae of mixed length [Fig. 36j].
Pupa [Figs 36d–f]. Length 2.5 mm. Antennal horns large, serrate along anterior edge, with basal appendix. Face
with triangular horn. Prothoracic spiracle elongate, trachea ending at apex. Abdominal segments evenly covered
with short spicules except anterodorsally with large spines.
Larva unknown.

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FIGURES 36a–l. Schizomyia laporteae: a, gall on Laportea stimulans (Fig. 74 of DvLR & DvL 1910b); b, same (Fig. 221 of
DvLR & DvL (1926)); c, apex of gonopod (ventral); d–f: pupa: d, head; e, thoracic spiracle; f, abdominal tergum 6 (lateral);
g, female first tarsomere; h, male palpus; i–l, female: i, tarsal claw and empodium; j, ovipositor apex; k, 3rd flagellomere; l,
antenna.
Remarks. Schizomyia laporteae differs from S. nodosa in the palpus being 3-segmented, the ovipositor with
4–5 pairs of setae and undefinable cerci, and the halberd-shaped pupal antennal horns. In S. nodosa the palpus is
4-segmented, female cerci are discrete, pupal antennal horns are not pronounced and a frontal horn is absent. Schizo-
myia villebrunneae differs from both those species by the pronounced pupal antennal horns that are bifid apically
and the male gonostylus with a spiny tooth. The female of S. villebrunneae is unknown.
Biology. This species causes a swelling on petioles and the underside of leaf bases of Laportea stimulans (Ur-
ticaceae) (Felt 1921b; DvLR & DvL 1910b, gall No. 180, Fig. 74 [Fig. 36a]; DvLR & DvL 1926, gall No. 314, Fig.
221 [Fig. 36b]). The gall is irregularly spheroid, 30–50 mm long and 5–12 mm thick on the petioles but only 8–10
mm long and 5–7 mm thick when on the midrib. The surface is light green and often granulate.
Geographical distribution. This species was found in Java (DvLR & DvL 1910b; Felt 1921b; DvLR & DvL
1926) at the following localities. Mt Ungaran, alt. 600 m, xii-1909; alt. 1400 m, 10-iv-1914; alt. 1000 m, xi-1919;
Malabar, near Bandung, alt. 1600 m, ix-1911; Pateunteung, near Garut, alt. 1500 m, xi-1918; Mt Gede, Cibeureum,
alt. 1700 m, ii-1916; Mt Gede, Cibodas, alt. 1500 m, vii-1920; Mt Burangrang, alt. 1000 m., vii-1920; Mt Beser,
near Cibeber, alt. 1000 m., vi-1923.
Schizomyia nodosa Felt
[Figs 37a–p]
Schizomyia nodosa Felt, 1921b: 144.
Material examined. Syntypes, male, two females, one pupa and three larvae were reared from Basilicum polys-
tachyon (L.) Moench (as Moschosma polystachum [error for polystachyum] Benth). collected iv-1914 at Semarang,
Java, Felt #a2099. The series is on three slides and uncleared. The male is entire, mounted laterally, with shrunken
terminalia; the slide contains additionally a cecidomyiine male with tricircumfilar antennae, ignored by both Felt
and us here. The females are missing some wings and tarsi, antennae are shrunken but appear entire, ovipositors are

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extended and visible. The pupa is partially damaged. Of the three larvae, one is entire with the terminal segment
filled with debris, one consisting only of a head and a spatula, and one is parasitized with only the spatula observ-
able.
Description. Adult. Wing R5 bowed at distal third, joining C at wing apex, R1 shorter than wing half, RS rudi-
mentary, M4 and Cu1 forming fork. Palpus 4-segmented [Fig. 37m]. First tarsomere with small, straight ventro-api-
cal lobe [Fig. 37j]. Tarsal claws simple, thin, bent at distal third, longer than empodia [Fig. 37i].
Male. Wing length 1.5 mm, width 0.6 mm. Flagellomeres 12, vaguely trinodal, circumfila strongly bowed [Fig.
37k], gonocoxite with triangular ventroapical lobe, bearing long setae at apex, gonostylus twice as long as wide,
bent and blunt distally, uniformly setulose, tooth on gonostylus not properly visible on available specimen [Fig.
37h].
FIGURES 37a–p. Schizomyia nodosa: a, gall on Basilicum polystachyon (Fig. 947 of DvLR & DvL (1926)); b–d, larva: b,
head (ventral); c, spatula with adjacent papillae; d, cuticle of 7th abdominal segment (ventrolateral); e–g, pupa: e, anterior seg-
ments (ventrolateral); f, prothoracic spiracle; g, 2nd abdominal tergum (dorsolateral); h–k, male: h, terminalia (lateral); i, tarsal
claw and empodium; j, first tarsomere; k, 3rd flagellomere; l–p, female: l, postabdomen (lateral, arrow marks dorsobasal lobe);
m, palpus; n, 3rd flagellomere; o. cerci; p, antenna.
Female. Wing length 1.7 mm (1.7–1.8), width 0.6 mm. Flagellomeres 11 in both antennae of both available
females, first and second fused, all with short necks, progressively shortened, with last flagellomere abruptly so,
circumfila appressed to node [Figs 37n, p]. Ovipositor long, thin, flexible, with small dorsobasal lobe [Fig. 37l];
needle-like part about 6x longer than seventh sternite, bearing minute, medially appressed cerci visible, each with
a single lateral seta [Fig. 37o].
Pupa. Length 1.8 mm. Antennal bases not produced, face without horns [Fig. 37e]; prothoracic spiracle long,
about 10x longer than basal width, trachea ending at apex [Fig. 37f]; abdominal segments anterodorsally with large
spiny spicules, otherwise mostly evenly covered with short spicules, on each side of terga a pair of closely set ase-
tose papillae [Fig. 37g].
Larva. Length 2.1 mm. Spatula bilobed, lateral papillae comprising one pair of setose papillae and one triplet of
two setose and one asetose papillae [Fig. 37c]; head with antennae 1.5x longer than wide at base, apodemes longer
than head capsule [Fig. 37b]; cuticle of rounded verrucae throughout [Fig. 37d].
Remarks. The reduced number of flagellomeres to 11 in both females of S. nodosa is odd for this genus so

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might only be an aberration in this series. See under the heading of Schizomyia and under S. laporteae for characters
that separate the three species of Schizomyia in Indonesia.
Biology. This species causes a flower gall on musk basil, Basilicum polystachyon (Lamiaceae), described by
DvLR & DvL (1926, gall No. 1324, Fig. 947 [Fig. 37a]). The calyx and the corolla are inflated, spheroid, about 5
mm across and have a rough surface. Anthers are atrophied but the pistil is swollen, inside of which a larva lives in
a small cavity. Often all the flowers of an inflorescence are infected.
Geographical distribution. This species was found in Java (Felt 1921b; DvLR & DvL 1926) at the following
localities. Kemantran, near Tegal, ix-1909; Semarang, vii-1910; iv-1914; Tirto, near Pekalongan, vi-1911; Kuripan,
near Pekalongan, vii-1911; Jrakah, near Semarang, viii-1912; Sayung, near Semarang, xi-1912.
Schizomyia villebrunneae Felt
[Figs 38a, d–q, Figs 38b–c (references)]
Schizomyia villebrunneae Felt, 1921b: 146.
Material examined. Lectotype male, here designated from a syntypic series of two males, two pupae and a larva
taken from a medium-sized, oval, thickly haired leaf gall on Oreocnide rubescens (Blume) Miq. (as Villebrunnea
rubescens Bl. [note misspelling of Villebrunea]) (Urticaceae) [Figs. 38a–c], collected at Mt Ungaran, Java, alt. 1000
m, 13-iv-1914 by DvL, Felt #a3092. The types are mounted whole, uncleared, on two slides. The first slide contains
the two males, with all body parts observable except for shrunken wings in both specimens, and only nine and three
flagellomeres present in the lectotype. The second slide holds two pupae with all characters preserved, one contain-
ing a male, the other undifferentiated matter, and a larva mounted in lateral position with only spatula and posterior
part of the head recognizable. Galls were illustrated in DvLR 1911, gall No. 247; DvLR & DvL (1926), gall No.
331, Fig. 237, [Fig. 38a]). Felt (1921b) suggested that this species was also responsible for two other leaf galls on
the same host (see under remarks), for one of which he described a series of two females (#a3091). Felt did not label
them as types and only tentatively assigned them to this species. We show that the females do not appear to belong
to this species.
Description. Male. Palpus 4-segmented, first segment as long as second, third and fourth progressively longer
[Fig. 38g]. Tarsus long, narrow, first tarsomere 1.3x, second 5x, third 2x, fourth 1.3x length last tarsomere [Fig.
38e]. Tarsal claws slender, evenly narrowing towards tip, slightly bent at distal third, longer than empodia [Fig.
38h]. Flagellomeres vaguely trinodal, circumifila irregular, anastomosing, strongly bowed [Fig. 38f]. Terminalia
[Fig. 38i]: gonocoxite with large triangular ventroapical lobe; mesobasal lobe ovoid, small, 2x longer than wide;
cerci irregularly incised on caudal margin; hypoproct as long as cerci, shallowly incised at apex, resulting lobes with
single seta; aedeagus longer than hypoproct, tapered and pointed; gonostylus in dorsal view as long as wide, distal
end covered about two-fifths width with spinose tooth, the remainder blunt, short-setose.
Female unknown.
Pupa [Figs 38j, k]. Antennal horns elongate, bifid at apex, resulting lobes of equal length and minutely serrate
apically. Prothoracic spiracle tapering, about 10x longer than basal width, trachea not quite reaching apex. Face
with pair of setae and no horns. Abdominal segments uniformly covered with spiculae except for rows of spines on
anterior third of terga.
Larva. Sternal spatula bilobed, lobes triangular, incision as long as lobes, with long shaft.
Remarks. To distinguish between this species and S. laporteae and S. nodosa, the two other species of the
genus from Indonesia, see under the headings of Schizomyia and S. laporteae. The two females that Felt (1921b;
specimens #a3091) only provisionally assigned to S. villebrunneae do not appear to fit this species because they
have shorter tarsomeres and more robust and more strongly curved tarsal claws [Fig. 38p]. For the record we show
a palpus, a flagellomere, the end of the ovipositor and a tarsus [Figs 38m–q] of these females. Another reason we
consider the females a separate Schizomyia species from S. villebrunneae is that they came from a different leaf gall
[Fig. 38b].
Felt (1921b) originally received from DvL three series of insects from three distinct kinds of gall on Oreocnide
rubescens (Blume) Miq. (as Villebrunnea rubescens Bl. [note misspelling of Villebrunea]) [Figs 38a–c]. The first
kind, from which the syntype series #a3092 was reared, is an oval, thickly haired gall [Fig. 38a], 7–9 mm long,

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4–5 mm broad and about 4 mm high, attached by a short pedicel to veins on both sides of the leaf. The second kind
[Fig. 38b] is a large, spherical, thinly haired gall, 5–12 mm in diameter, attached by a very short pedicel to the leaf
surface, and covered with thin, white, transparent, often pink hairs (DvLR & DvL 1926, gall No. 330, Fig. 236). The
third kind is a small, oval, glabrous gall, 4 mm long and 2.5 mm broad, situated on veins of the upper side of the
leaf (DvLR & DvL 1926, gall No. 332, Fig. 238). Felt (1921b) considered all three kinds to be made by the same
species, yet he noticed a difference between males of the first kind and females from the second, causing him to only
provisionally assign the females to S. villebrunneae. DvL (1921) was not convinced that the three distinct galls were
caused by the same gall midge, but DvLR & DvL (1926) listed all three kinds of galls as made by S. villebrunneae.
We consider only the first kind of gall [Fig. 38a] to be made by S. villebrunneae.
Biology. As noted above, galls of S. villebrunneae [Fig. 38a] develop on the veins of both sides of the leaf. Galls
are oval, the surface is whitish, yellowish, but mostly dark red and covered with white hairlets. The single larval
chamber is spherical and centered at the middle of each gall (DvLR & DvL 1926).
Geographical distribution. The known occurrence of all three gall types is confined to Java (Felt 1921b;
DvLR & DvL 1926). We expect that each of the three galls will one day be proven to be caused by a different gall
midge, so we list here their geographical distribution separately. The first kind from which the syntypes came [Fig.
38a] was found at the following localities. Mt Ungaran, alt. 1000 m, xii-1909; 13-iv-1914; Sekecer, near Weliri, alt.
100 m, x-1910; Cikadongdong, near Bandung, alt. 45 m, v-1915; Cisokan, near Cibeber, alt. 900 m, xii-1917; Mt
Cibodas, Ciampea, near Bogor, alt. 300 m, iv-1919; Cianten, near Buitenzorg, alt. 400 m, viii-1924. Additionally,
this gall was found by Purnama Hidayat and Mahindra Dewi Nur Aisyah (IPB University (pers. comm.) at Naring-
gul, near Cianjur (7°21’6.15”S 107°7’8.87”E), alt. 252 m, 12-iv-2018. The second kind [Fig. 38b] was found at the
following localities. Mt Ungaran, alt. 600 m, xii-1909 & v-1910; alt. 1000 m, 13-iv-1914; Mt Andong, alt. 1200 m,
x-1911; Mt Kelud, alt. 600 m, v-1912; Pateungteung, near Garut, alt. 1500 m, xi-1918; Mt Gede, Cibodas, alt. 1300
m, xii-1914; ii-1916; ix-1918; i-1919; Pangalengan, near Bandung, alt. 1400 m, vi-1921. Additionally, this gall was
found at Mt Gede Pangrango National Park, Cianjur (6°44‘52“S, 106°59‘31“E), alt. 1591 m, 17-iv-2018 (Hidayat
& Nur Aisyah, pers. comm.). The third kind [Fig. 38c] was found at the following localities. Mt Ungaran, alt. 1000
m, ix-1910; alt. 1400 m, 14-iv-1914; Cadas, Malang, near Cibeber, alt. 1200 m, ix-1917; Mt Salak, alt. 1200 m,
vii-1920.
Stenodiplosis eragrostidis Felt, new combination
[Figs 39a–g]
Contarinia eragrostidis Felt, 1927b: 381.
Material examined. Types: 9 males, 11 females (NYSM #a3440), reared from seed heads of Eragrostis unioloi-
des (Retz.) Nees ex Steud., collected by C.A. Backer at Bogor, Java [see DvLR & DvL (1941) for this locality],
viii-1922. The type series is mounted on two slides, one with 4 males and 5 females, the other with 5 males and 6
females, all whole and uncleared. Most characters are well-preserved in at least one of the specimens, except for the
male terminalia that are mostly shriveled in all the males.
Description. Male. Length 1 mm (Felt 1927b), wing length 1.2 mm (1.1–1.4), width 0.4 mm (0.4–0.5). Palpus
1-segmented [as in Fig. 39e]. Antenna: 12 flagellomeres, first and second fused; flagellomeres binodal, each node
with one whorl of looped circumfila, basal node slightly wider than long, distal slightly longer than wide, internode
one third length of basal node, neck half-length distal node; circumfilar loops subequal, on basal node reaching end
of internode, on distal node reaching end of neck [Fig. 39c]. Wing with R5 slightly bowed at distal ¼, joining C
slightly beyond wing apex [Fig. 39a]. Tarsal claws simple, curved beyond midlength, widened beyond bend, em-
podia as long as claws [Fig. 39b]. Abdominal tergites lacking lateral setae. Terminalia: gonostylus stout, strongly
tapering from wide base towards tooth, completely setulose [Fig. 39d].
Female. Length 1.5 mm (Felt 1927b), wing length 1.2 mm (1.1–1.4), width 0.4 mm (0.4–0.5). Antenna: flagel-
lomeres about 2.5x longer than wide, neck 1/7 total length, node with shallow, bare constriction at midlength, each
half of flagellomere with separate whorl of looped circumfila, loops on basal half reaching constriction, loops on
distal half nearly reaching end of neck [Fig. 39f]. Terminalia: ovipositor as long as abdomen; cerci [Fig. 39g] closely
appressed, tapered, about 2.5 longer than basal width, without microtrichia and with setae of various sizes.

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FIGURES 38a–c. Galls on Oreocnide rubescens: a, galls of Schizomyia villebrunneae type series (Fig. 237 of DvLR & DvL
(1926)); b, galls of Schizomyia sp. (Fig. 236 of DvLR & DvL (1926)), c, galls of undetermined cecidomyiid (Fig. 238 of DvLR
& DvL (1926)). Figs 38d–k. Schizomyia villebrunneae: d, larval spatula with associated papillae; e-i, male: e, tarsus; f, 3rd
flagellomere (damaged); g, palpus; h, last tarsomere; i, terminalia (dorsal); j, pupal anterior segments (lateral); k, same, detail
(ventral). Figs 38l–q. Schizomyia sp.: female: l, last tarsomere; m, palpus; n, antenna; o, tarsus; p, 2nd flagellomere (damaged);
q, ovipositor apex.

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Pupa, larva, unknown.
Remarks. We transfer this species to Stenodiplosis Reuter, a Holarctic group presently comprising 12 species,
all of them associated with grass seeds, some of considerable economic importance (Gagné & Jaschhof 2017). The
most important relevant adult character for this generic placement is the lack of lateral setae on the abdominal ter-
gites (Gagné 2018). Many Stenodiplosis spp. exhibit a reduction in palpal segments, so S. eragrostis also fits that
criterion with its single segment. Not an exclusive character, but most Stenodiplosis species have fully setulose
gonostyli, as does this species. Larval characters also define the genus, including the loss of the larval spatula, the
miniscule terminal segment and terminally situated hind spiracles. The larva of S. eragrostidis is unknown, but we
expect that it shares these character states.
Felt (1927b) stated that the male palpi were indistinguishable in the mounted specimens and that the female had
a four-segmented palpus. We found that the palpi are one-segmented in both sexes, as did Harris (1979) who also
examined the type series. When Harris (1979) described Contarinia brevipalpis from the inflorescence of Eragrostis
brownii in tropical Australia, he noted that it shared with C. eragrostidis the one-segmented palpus, and the binodal
female flagellomeres that bear two whorls or circumfila. Additionally, these two species share the shape of the
gonostylus and female cerci, also the relative length of internodes and necks in both male and female flagellomeres.
They can be separated by the tarsal claws that are shorter and less curved in C. brevipalpis (Harris 1979, Fig. 44)
than in C. eragrostidis [Fig. 39b] and by the setation of female cerci that bear apically two pairs of long setae in C.
brevipalpis (Harris 1979, Fig. 36) and one pair in C. eragrostidis [Fig. 39g]. These differences may turn out to be
very minor but, without Harris’s types and additional specimens before us, we cannot make a decision at this time.
We take the occasion to place C. brevipalpis in Stenodiplosis also, as a new combination.
Biology. Contarinia eragrostidis causes slight swelling and developmental retardation of seed heads of
Eragrostis unioloides (Poaceae), with the gall approximately 5 mm long and 2 mm in diameter (Felt 1927b).
Geographical distribution. This species is known from Java where it was found at the type locality in Bogor,
alt. 250 m, viii-1922 (Felt 1927b; DvLR & DvL 1941). Additionally, it was found on Andaman Islands (India):
Port Blair, 27-xi-1981; Dhanikhari, 11-xii-1981; Manarghat, 13-xii-1982; Wandoor 24-ix-1983. Several males and
females were reared and deposited in the collection of the museum of Zoological Survey of India at Port Blair
(Sharma 1984) but we have not seen these specimens.
FIGURES 39a–g. Stenodiplosis eragrostidis: a–d, male: a, wing; b, last tarsal segment of hind leg with claw and empodium; c,
6th flagellomere; d, gonostylus (dorsal); e–g, female: e, palpi; f, 6th flagellomere; g, cerci (dorsal).

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Thorodiplosis Felt
[Figs 40a–j]
Thorodiplosis Felt, 1921d: 89.
Type species: Thorodiplosis impatientis Felt, by original designation.
Description. This monotypic genus was erected for two males and two females based on rather broad characteris-
tics: very long and narrow wings, elongate antennae, simple tarsal claws curved at almost right angle, short oviposi-
tor with bilobed cercus, and 4-segmented palpus (Felt 1921d). The types are uncleared and partially shriveled but all
characters are recognizable, except for the male circumfila that were missing already when Felt (1921b) examined
them. Wing [Fig. 40g] with R5 bent distally, joining C posteriad of wing apex; C broken at juncture with R5; M4 and
Cu1 forming fork; Rs not present. Head with occipital protuberance present. Palpi shriveled on all specimens but
evidently four-segmented. Antennae: 12 flagellomeres, progressively shorter, necks bare of microtrichia; binodal in
male, presumably trifilar, with long necks [Fig. 40i]; cylindrical in female, all except last with long necks, proximal
flagellomeres with nodes 3x as long as wide, constricted at basal third, distal flagellomeres with nodes 2x as long
as wide, progressively losing basal constriction, circumfila consisting of one uneven horizontal and two more distal
vertical bands [Figs 40c, d]. Tarsal claws simple, narrow, bent at right angle at midlength, as long as empodia [Fig.
40e]. Male terminalia [Figs 40h, j]: gonocoxite narrow, with low, wide mesobasal lobe; gonostylus very long, nar-
row, widest at distal fourth, straight and setulose basally, curved and carinate beyond, with small, solid, tooth; cerci
broad and concave apically, the mesal lobe of the concavity with single seta; hypoproct longer than cerci, narrow,
incised apically, resulting lobes each with single seta apically; aedeagus longer than hypoproct, tapered to narrowly
rounded apex, with asetose papillae distally. Female terminalia [Fig. 40f]: ovipositor short, cerci large, ovoid, setose
mainly distally, each with pair of long, blunt apical sensory hairs; hypoproct with pair of setae.
Remarks. We place Thorodiplosis in the tribe Clinodiplosini because of the conformation of the male termina-
lia, particularly the apically concave cerci.
Thorodiplosis impatientis Felt
[Figs 40a–j]
Thorodiplosis impatientis Felt, 1921d: 89.
Material examined. Type male and female reared from a marginal roll on the upper leaf surface of Impatiens
platypetala Lindl. (Fig. 599 [Fig. 40a] in DvL & DvLR (1918)), at Mt Gede, Cibodas, Java, 15-vii-1920 (NYSM
#a3153). Felt’s two slides contain two males and two females, all mounted whole and uncleared, with most body
parts present but partially shriveled.
Description. Male. Wing length 1.6 mm, width 0.6 mm. Antenna: basal node slightly longer than wide, inter-
node as long as basal node, proximal node 1.5x length basal node, neck 2x length basal node [Fig. 40i]. Terminalia:
gonocoxite splayed with low mesobasal lobe, gonostylus as long as gonocoxite, curved at distal third, setulose at
basal third, carinate and setose beyond [Figs 40h j].
Female. Wing length 2.2 mm, width 0.8 mm. Antenna as in Figs 40c, d. Terminalia: cerci setulose, sparsely
setose, with pair of long, blunt, closely set sensory hairs at apex [Fig. 40f].
Pupa, larva unknown.
Biology. This species causes a marginal roll on the leaf of Impatiens platypetala (Balsaminaceae) [Figs 40a, b].
The gall is figured in DvL & DvLR (1918, gall No. 599, Fig 599 [Fig. 40a]) and described by DvLR & DvL (1926)
as follows. The margin of the leaf is rolled up for a small part of its length. The infected parts are white or yellowish
and considerably thickened. The gall can be common locally.
Geographical distribution. This species was reported from Java (DvLR & DvL 1914, 1918; Felt 1921; DvLR
& DvL 1926, 1941) from the following localities: Pagergunung, alt. 900 m, ix-1911; Paseh, near Bandung, alt. 1000
m, xii-1915; Ciharum, near Cibeber, alt. 1200 m, vi-1916; Mt Slamet, alt 800 m, v-1918; Cisarua, near Cimahi, alt.
1300 m, v-1918; Mt Gede, Cibeureum, alt. 1700 m, ii-1916; Mt Gede, Cibodas, alt. 1400 m, ix-1918; alt. 1500 m,
15-vii-1920; alt. 2000 m, vii-1928; Mt Pangrango, alt. 2300 m, vi-1920; Mt Salak, alt. 800 m, vi-1918; vii-1920; x-
1921; Mt Cikuray, alt. 1600 m, v-1925; Maseng, near Bogor, alt. 600 m, vi-1925. Additionally, Purnama Hidayat

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FIGURES 40a–j. Thorodiplosis impatientis: a–b, galls on Impatiens platypetala: a, galls indicated with arrows (Fig. 599 of
DvL & DvLR 1918); b, Cianjur near Mt Gede (photo by Purnama Hidayat and Mahindra Dewi Nur Aisyah, 2018); c–g, female:
c, 1st and 2nd flagellomeres; d, antenna; e, tarsal claw and empodium; f, cerci and hypoproct (lateral); g, posterior part of wing; h–
j, male: h, terminalia (dorsal); i, 6th flagellomere (circumfila not visible on types); j, aedeagus of both type specimens (dorsal).

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and Mahindra Dewi Nur Aisyah (IPB University) (pers. comm., Fig. 40b) found the galls at Mount Gede Pangrango
National Park, Cianjur (6°52’33”S, 106°58’04”E), alt. 1407 m, 17-iv-2018; Kertajaya, Pasirkuda, near Cianjur
(7°14’38”S, 107°10’24”E), alt. 504 m, 10-iv-2018; Pagelaran, near Cianjur (7°17’45”S, 107°7’7”E, alt. 967 m, 13-
iv-2018; Mt Sari, Pamijahan, Bogor (near Mt. Bunder) (6°41’23”S, 107°12’01”E), alt. 967 m, 17-iii-2018.
Species unplaced to genus
The following three species are too incompletely known to be placed in existing genera or to describe new genera
for them. Their types were reared from distinctive galls on known plants, therefore more specimens should someday
be found on which to base proper descriptions.
“Calopedila” polyalthiae Felt
[Figs 41a–h]
Calopedila polyalthiae Felt 1927b: 385; Gagné 1973: 288, unplaced to genus.
Material examined. Syntype series, 8 males, 17 females, pupa, reared from fruit gall on Polyalthia subcordata
(Blume) Blume, Mt Gede, Cibodas, Java, xi-1923, Felt #a3386. The types are uncleared and mounted on two slides,
the first with 7 males, 6 females and 1 pupa, all abdomens filled with black, non-transparent matter and male termi-
nalia mounted laterally and badly shriveled, the second slide with 1 male, 9 females, all abdomens filled with black
matter except for part of the terminalia on some specimens.
Description. Adult. Occipital protuberance absent. Wing: R5 nearly straight, joining C anteriad of wing apex,
C broken at juncture with R5, Rs not present [Fig. 41e] Irregular number of flagellar segments, first and second
flagellomeres not connate, 17–18 (n=3 in male, cylindrical and short-necked [Fig. 41h], and 19 (n=1) flagellomeres
in female with very short necks [Fig. 41g]. Palpus 3-segmented, palpiger present [Fig. 41c]. Tarsal claws bent at
midlength, robust, toothed, empodia as long as claws [Fig. 41d].
Male. Wing 2.4 mm (2.4–2.5) long, 1.1 mm (1.0–1.1) wide. Terminalia not properly observable on available
specimens; gonostylus completely setulose.
Female. Wing 3.0 mm (2.9–3.0) long, 1.2 mm wide. Terminalia: elongate, protractible, cerci fused, tapered,
more than 2x longer than basal width, hypoproct narrow in lateral view, 1/3 length of cerci [Fig. 41f].
FIGURES 41a–h. “Calopedila” polyalthiae: a, galls on Polyalthia subcordata (Fig. 293 of DvLR & DvL (1926)); b, pupal
vertex and antennal bases; c–g, female: c, palpus; d, tarsal claw and empodium; e, wing; f, postabdomen (lateral); g, 10th flagel-
lomere; h, male 7th flagellomere.

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Pupa [Fig. 41b]. Cervical setae short, less than half-length of antennal horns. Antennal horns elongate, in ven-
tral view narrowed by half at distal third, resulting in an acutely pointed lateral lobe and a terminal, rounded apical
lobe. Prothoracic spiracle short, narrow.
Remarks. Felt (1927b) placed this species only tentatively in Calopedila, a genus erected for Rhopalomyia
herbsti (Kieffer 1903) that causes a branch gall on Baccharis rosmarinifolia (Asteraceae) in Chile. The Chilean spe-
cies has tarsal claws with small teeth and empodia that are nearly twice the length of the claws, male flagellomeres
without necks, and female cerci that are no longer than broad. From these characters Calopedila appears to have
an affinity to Rhopalomyia Rübsaamen and is treated as a junior synonym of Rhopalomyia in Gagné & Jaschhof
(2017). “Calopedila” polyalthiae is shown here to have tarsal claws with large teeth and empodia that are only as
long as the claws [Fig. 41d], male flagellomeres with necks [Fig. 41h], and female cerci that are elongate and narrow
[Fig. 41f], all of which show that the species was initially misplaced to genus. The species belongs to the supertribe
Lasiopteridi, but we cannot guess where this species might be assigned below that level given the poor specimens
available and the fact that so little is known of SE Asian Lasiopteridi generally. We might feel justified in erecting a
new genus for “C.” polyalthiae once better material is found.
Biology. This species causes a fruit gall on Polyalthia subcordata (Annonaceae) described by DvLR & DvL
1926, gall No. 408, Fig. 293 [Fig. 41a]). Individual galls are spheroid, about 3 mm across and covered with short
brown hairs. They develop on the surface of the fruit, occurring mostly gregariously and arranged in thick irregular
clumps. The larval chamber is small and spherical.
Geographical distribution. This species is known only from its type locality at Mt Gede, Cibodas, Java, alt.
1400 m, vi-1920; alt. 1500 m, xi-1923 (DvLR & DvL 1926).
“Gnesiodiplosis” garcinia Felt
[Figs 42a–h]
Gnesiodiplosis garcinia Felt, 1927b: 388; Gagné 1973: 514, unplaced to genus.
Material examined. Syntypes, 17 males, 10 females, 2 pupae, reared from a leaf gall on Garcinia dioica Blume at
Depok, near Jakarta, viii-1922, Felt #a3388. The types are mounted on four slides (on slide labels, Depok is mis-
spelled as “Depote”): one with 2 cleared (by RJG males without distal leg segments; a second with 3 uncleared
females mounted laterally with most body parts retained except most distal leg segments; a third with 10 males and
2 females, uncleared, mounted laterally, most body parts recognizable except terminalia that are mostly shriveled
and distal leg segments that are missing; and a fourth with 5 males, 5 females, 2 pupae, all uncleared, most body
parts barely recognizable.
Description. Adult. Occipital protuberance absent; flagellomeres 12, first and second separated. Palpus 1-seg-
mented, palpiger present [Fig. 42g]. Wing with R5 joining C slightly beyond wing apex, C with break at juncture
with R5, Rs rudimentary [Fig. 42e]. Tarsal claws simple, strongly bent at midlength, empodia substantially shorter
than claws [Fig. 42h].
Male. Wing 2.6 mm (2.6–2.7) long, 1.0 mm wide. Flagellomeres binodal, proximal node as long as wide, distal
twice as long as wide, internode half the length of proximal node, neck as long as proximal node, circumfila not pre-
served on available specimens [Fig. 42b]. Terminalia: gonocoxite robust; gonostylus long, tapered, setulose basally,
carinate beyond; cerci apparently partly conjoined; hypoproct broad, incised, the lobes rounded; aedeagus as long
as gonocoxite, broad at base, tapered to acute tip [Fig. 42c].
Female. Wing 2.1 mm long, 0.7 mm (0.6–0.8) wide. Flagellomeres with simple, slightly bowed circumfila con-
sisting of two horizontal and one vertical, interconnected bands. Terminalia: ovipositor short; cerci not clearly seen,
apparently folded with massed short setae covering caudal surface; hypoproct, narrow in lateral view [Fig. 42f].
Pupa. Length 2.4 (2.3–2.4) mm. Antennal base with minute angular horns. No abdominal dorsal spines pres-
ent.
Larva unknown.
Remarks. Felt (1927b) only provisionally placed this species in Gnesiodiplosis. That genus is based on the
monotypic Gnesiodiplosis itaparicae Tavares, a Brazilian species that causes rosette bud galls on an unidentified
species of Rubiaceae in Brazil. The terminalia of both species are quite distinct, the male of G. itaparicae having an

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 73
evenly cylindrical aedeagus and long parallel-sided and blunt-ended hypoproct and the female elongate-ovoid cerci.
In contrast, the aedeagus of “G.” garciniae is robust and tapered, and the hypoproct is short with two rounded api-
cal lobes, and the female cerci are shorter and blunted apically. There is no apparent reason why these two species
should be considered congeneric and we can offer no suggestion as to where the present species belongs. In due
course, with fresh specimens, a new genus could be erected for “G.” garciniae.
Biology. This species causes a leaf gall on Garcinia dioica (Clusiaceae) described by DvLR & DvL (1926, gall
No. 993, Fig. 708 [Fig. 42a]). The midrib is swollen into irregular, green or brown, oval or spheroid galls that show
on both sides of the leaf blade. In cases of a strong infection the midrib may be zigzagged and the leaf blade more
or less wrinkled and folded. Inside each gall is a tiny larval chamber surrounded by a succulent thick parenchymatic
tissue Average size of a single gall is 5 mm across.
Geographical distribution. This species is known from Java where it was found at Depok, near Jakarta, alt.
100 m, v-1911, iv-1919, viii-1920, additionally Felt (1927) gives viii-1922, and at Ciastana near Cisalak, Mt Hali-
mun, alt. 550 m, xii-1920.
FIGURES 42a–h. “Gnesiodiplosis” garcinia: a, galls on Garcinia dioica (Fig. 708 of DvLR & DvL (1926)); b–c, male: b, 4th
flagellomere (circumfila missing); c, terminalia (dorsal); d–h, female: d, 3rd flagellomere; e, wing; f, terminalia (dorsolateral); g,
palpus and palpiger; h, tarsal claw and empodium.
“Stefaniella” orientalis Felt
[Figs 43a–e]
Stefaniella orientalis Felt, 1921b: 142; Gagné 1973: 491, unplaced to genus.
Material examined. Holotype male, reared from a petiole gall on Lepidagathis javanica Blume at Mt Ungaran,
Java, altitude 1200 m. 18-iv-1912, Felt #a3098. The type is mounted uncleared with all body parts present but folded
or shriveled.
Description. Male. Wing 1.5 mm long, 0.6 mm wide; R5 2/3 wing length, C interrupted at juncture with R5.
Flagellomeres 16 on both antennae, first and second fused, trapezoid, necks minute [Fig. 43e]. palpus 2-segmented,
first segment 1.5x longer than wide, third 3x longer than wide [Fig. 43d]. Tarsal claws thin, bent at midlength with
long, thin tooth, empodia slightly shorter than claws [Fig. 43c]. Terminalia: gonostylus bulbous basally, tapered
distally, setulose at basal third and narrow, carinate beyond; cerci triangular; hypoproct bilobed, setulose with few

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74 · Zootaxa 4847 (1) © 2020 Magnolia Press
setae apically; mesobasal lobes closely sheathing aedeagus, covered with dense setulae, three setose papillae on
each lobe apically; aedeagus triangular in lateral view, without sensoria [Fig. 43b].
Female, pupa, larva unknown.
FIGURES 43a–e. “Stefaniella” orientalis: a, gall on Lepidagathis javanica (Fig. 976 of DvLR & DvL (1926)); b, male termi-
nalia (distorted; aedeagus (ae), cercus (ce), gonocoxite (gc), gonostylus (gs), hypoproct (hy), mesobasal lobe (ml); c, tarsal claw
and empodium; d, palpus; e, male antenna.
Remarks. Felt (1921b) tentatively placed the male in Stefaniella, a genus of Palearctic Lasiopterini of 10 spe-
cies that feed on Chenopodiaceae (Gagné & Jaschhof 2017). The male does belong to Lasiopterini but one needs
females to assign species to genera.
Biology. This species causes a petiole gall on Lepidagathis javanica (Acanthaceae) described by DvLR & DvL
(1926, gall No. 1377, Fig. 976 [Fig. 43a]). It is an aggregate, uneven swelling about 10 mm long and 6 mm across.
Inside each gall is a small, longitudinal. Hard-walled larval chamber.
Geographical distribution. This species is currently known only from Mt Ungaran, Java where it was found
at alt. 1000 m, ix-1910 and 1200 m, 18-iv-1912 (DvLR & DvL 1926; Felt 1921b).
Addendum
Eucalyptodiplosis paederiae (Felt). New combination.
[Figs 17a–l]
Itonida paederiae Felt 1919: 293; Gagné 1975: 515, as unplaced species of Cecidomyiidi.
This Philippine species was mentioned under the Indonesian Hypodiplosis paederiae Kieffer & DvLR that we
treated above because it forms a similar leaf roll on the same host, Paederia foetida. We established that the two
species are distinct, which we show here with the redescription of I. paederiae and its consequent reassignment to
Eucalyptodiplosis.
Material examined. Syntypes, a male and female, were collected in Luzon, Laguna Province, Los Baños, the
Philippines, 13-x-1917 Felt #a2882. The specimens are mounted separately as whole insects. They are cleared and
well-preserved.
Description. Adult. Occipital protuberance short, as long as wide [Fig. 17g]. Antenna with 12 flagellomeres,
first and second fused. Palpus 4-segmented, palpiger present [Fig. 17g]. Wing: RS reduced to slight thickening on
R5 closer to the arculus than to end of R1; R5 slightly curved, joining C posteriad of wing apex; C broken at juncture
with R5 [Fig. 17d] Tarsal claws nearly as long as empodia, bent at nearly 90˚ at midlength, widened at distal third,
pointed apically, with narrow curved tooth on foreleg only [Figs 17j, k, l].
Male. Wing 4.8 mm long, 1.8 mm wide. Flagellomeres binodal, basal node with one whorl of circumfila, distal
with two, the loops regular, not reaching next distal circumfilum [Fig. 17e]. Terminalia [Fig. 17f]: cerci tapered,

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 75
slightly concave mesally with seta at basal end of concavity, acutely triangular and setose apically; hypoproct nar-
row, longer than cerci, deeply and widely incised, lobes thin, each bearing single apical seta; aedeagus long, thin,
evenly cylindrical, longer than hypoproct and gonocoxite, rounded apically and bearing asetose sensilla; gonocoxite
narrow, with obtuse mesobasal lobe; gonostylus long, narrow, setulose basally, carinate beyond, with many setae in
carinate area, dense apically.
Female. Wing 5.1 mm long, 1.9 mm wide. Flagellomeres cylindrical, necks slightly less than half as long as
nodes [Fig. 17i]. Ovipositor short; cerci elongate-ovoid, densely setulose, setose, apically with pair of thick blunt
setae; hypoproct setulose, with pair of thick apical setae [Fig. 17h].
Biology. Uichanco’s (1919, gall No. 18165, Figs I-1, I-6, VIII-5 [Figs 17a–c]) description of the gall is as fol-
lows but slightly edited. The gall consists of a superior longitudinal infolding of both margins of leaf blade, with a
consequent upward curvature of the affected leaf, and extends from the base to about one-sixth of the leaf length
from apex. The margins touch each other tangentially at sides along midrib. Apical and subapical portions of the leaf
lamina remain normal. The chambers formed inside the roll are thickly lined with long pubescence. The midrib and
lateral veins are abnormally pubescent on the undersurface of the leaf. Adults make their exit at either end.
Remarks. We assign I. paederiae to Eucalyptodiplosis Kolesik (2002) because its male terminalia best fit that
genus, which is known from three Australiasian species, two of them on Eucalyptus (Myrtaceae) and one on Chi-
onochloa (Poaceae). Male cerci are tapered so that they appear nearly triangular but are slightly concave mesally
with a seta at the basal end of the concavity. That concavity is usually situated across the caudal end of the cerci of
the tribe Clinodiplosini. In Eucalyptodiplosis the mesal lobe of the concavity is skewed basad of the lateral lobe, but
the single seta on the lower lobe is the tell-tale sign that the genus also belongs to the Clinodiplosini. We should note
that E. paederiae has toothed tarsal claws only on the forelegs while the claws of all legs are toothed in the three
Australiasian species of the genus (Kolesik et al. 2002; Kolesik et al. 2007). Considering that Clinodiplosis itself
harbors species that may have toothed and untoothed claws and of different shapes (Gagné 1994), the difference in
that structure should not disqualify E. paederiae from its present assignment to Eucalyptodiplosis.
Key to adults in genera of Cecidomyiidae currently known from Indonesia
This key is by no means comprehensive for the gall midges to be found in Indonesia, it rather constitutes an initial
step in creating a framework for future discoveries. When finding new taxa, keys to genera of neighboring faunas
will also be of assistance: Kolesik (2015) for Australia and Papua New Guinea, that was expanded by Kolesik &
Gagné (2016) to include additional genera, and Yukawa (1971) for Japan, which can be used to good effect together
with descriptions of new genera since published by him, Makoto Tokuda, Nami Uechi and Ayman K. Elsayed. Of
considerable help will be the comprehensive key to genera of North America by Gagné (2018).
All but one genus of Cecidomyiidae known from Indonesia belong to the subfamily Cecidomyiinae, which
contains all of the plant-feeding and predaceous forms. The exception is Colomyia that belongs to Porricondylinae,
one of five primitive, fungus-feeding subfamilies that are barely known in the Oriental region. To learn more of
Porricondylinae and its sister group Winnertziinae one should see Jaschhof & Jaschhof (2013); for Lestremiinae and
Micromyinae Jaschhof & Jaschhof (2009a); and for Catotrichinae Jaschhof & Jaschhof (2008).
In the key, each genus is immediately followed by world distribution and a list of Indonesian species and their
hosts. For further information on the genera in other geographical regions see Gagné & Jaschhof (2017). We were
able to key two species, “Gnesiodiplosis” garcinia and “Calopedila” polyalthiae, that are unplaced to genus. Al-
though not strictly genera, the two taxa are distinct entities that may one day attain generic status. We could not
place a third, “Stefaniella” orientalis Felt. It is known only from a male so does not have enough cues to be distin-
guishable among genera in the tribe Lasiopterini to which it belongs. We did not include Eucalyptodiplosis, which
contains the Philippine E. paederiae treated here, but it would run to Feltiella from which it differs in the male
hypoproct being bilobed as opposed to entire. We did include Colomyia cortici (Porricondylinae) which effectively
makes this a key to genera of the whole family Cecidomyiidae currently known from Indonesia.
1 Rs as strong as other wing veins. Male gonocoxites fused ventrally. Female cerci two-segmented. . . . . . . . . . . . . . . . . . . . . . .
..........................................................Colomyia Kieffer (Porricondylinae, not treated here)
Palearctic and Oriental

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76 · Zootaxa 4847 (1) © 2020 Magnolia Press
Colomyia cortici feeding on fungus Corticum sp. (Agaricomycetes: Corticiaceae) infesting tea plant Camellia sinensis (Thea-
ceae) (Edwards 1923)
- Rs weak, reduced to stub [Figs 31h] or missing [Figs 39a]. Male gonocoxites separated ventrally [Fig. 27d]. Female cerci one-
segmented, sometimes fused or otherwise modified [Figs 2g, 3l, 19j, 34h, 39g]. . . . . . . . . . . . . . . . . . . . . . . . .Cecidomyiinae 2
2 R5 as long [Figs 3j] or nearly as long [Fig. 15c] as wing. Antennal flagellomeres usually clearly sexually dimorphic [Figs 17e,
i] or binodal in both sexes [Figs 16i, f]. ....................................................................3
- R5 half to two-thirds length of wing, closely adjacent and nearly parallel to R1 and C [Fig. 19e]. Antennal flagellomeres similar
between sexes [Figs 2i, j]. .............................................................................23
3 Gonostylus situated on dorsum of gonocoxite [Fig. 37h]. Female abdominal seventh sternite appreciably longer than sixth. . 4
- Gonostylus situated on apex of gonocoxite [Fig. 17f]. Female abdominal seventh sternite not appreciably longer than sixth. . .
...................................................................................................7
4 First tarsal segment with large ventrodistal spur [Fig. 3e]. .....................................................5
- First tarsal segment without spur [Fig. 9f] or with minute ventrodistal lobe [Fig. 36g]. ...............................6
5 Gonostylus with single wide tooth [Figs 3h, 7b]. ...............................................Asphondylia Loew
Cosmopolitan
Asphondylia callicarpae causing leaf gall on Callicarpa erioclona and Callicarpa longifolia (Lamiaceae)
Asphondylia capsicicola causing pod deformation on Capsicum annum and C. frutescens (Solanaceae)
Asphondylia ixora causing flower gall on Ixora timorensis (Rubiaceae)
Asphondylia leeae causing fruit gall on Leea indica (Vitaceae)
Asphondylia litseae causing stem gall on Litsea sp. (Lauraceae)
Asphondylia strobilanthi causing aerial root gall on Strobilanthes cernua (Acanthaceae)
Asphondylia viticola and A. vitea causing stem and petiole galls on Cayratia trifolia (Vitaceae)
Asphondylia yushimai causing fruit malformation on soybean Glycine max (Fabaceae), Laurocerasus zippeliana (Rosaceae)
and Osmanthus heterophyllus (Oleaceae)
- Gonostylus with two separate teeth [Figs 10e, f]. .........................................Bruggmanniella Monzen
Palearctic, Oriental and Oceanian
Bruggmanniella bursaria causing stem gall on Symplocos fasciculata (Symplocaceae)
Bruggmanniella orientalis causing stem gall on Phoebe declinata (Lauraceae)
6 Tarsal claws simple, curved at distal third [Fig. 36i].. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizomyia Kieffer
Cosmopolitan
Schizomyia laportae causing leaf gall on Laportea stimulans (Urticaceae)
Schizomyia nodosa causing flower gall on Basilicum polystachyon (Lamiaceae)
Schizomyia villebrunneae causing leaf gall(s) on Oreocnide rubescens (Urticaceae)
- Tarsal claws pectinate, strongly bent at basal third [Fig. 9h]. ....................................Asphoxenomyia Felt
Oriental
Asphoxenomyia orientalis causing leaf gall on Smilax modesta (Smilacaceae)
7 Antenna with 12 flagellomeres, binodal in male [Fig. 35g] except in Megommata [Fig. 24g] and then eyes nearly divided at
mid-height [Fig. 24a]. Male terminalia with no mesobasal lobes [Fig. 40h] or with lobes not sheathing aedeagus [Fig. 33h].
Female cerci discrete [Fig. 12g] but may be closely appressed [Fig. 39g]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Antenna with irregular number of flagellomeres within species, seldom 12. Male and female flagellomeres with single bar-
rel-shaped node, those of male with longer necks than in female [Figs 11c, g]. Male terminalia with mesobasal lobes tightly
sheathing aedeagus [Fig. 11b]. Female cerci fused [Figs 11e; 15f, g]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
8 Foreclaws toothed, mid- and hindclaws with or without teeth [Figs 17j–l]. ........................................9
- Claws without teeth. ..................................................................................14
9 Only foreclaws toothed. ...............................................................................10
- All claws toothed. ....................................................................................12
10 Eyes nearly divided at midheight [Fig. 24a]. Male flagellomeres with single node as in female [Fig. 24g]. ................
....................................................................................Megommata Barnes
Afrotropical and Oriental
Megammota leefmansi preying on eggs of Pulvinaria polygonata (Hemiptera: Coccidae)
- Eyes not divided at midheight. Male flagellomeres binodal [Fig. 32d]. ..........................................11
11 Gonostylus setulose only basally. Female tenth tergum covered with setae. .........................Feltiella Rübsaamen
Cosmopolitan
Feltiella acarivora preying on spider mites (Acari: Tetranychidae)
- Gonostylus completely setulose. Female tenth tergum with only two long setae [Fig. 13i]. ................Diadiplosis Felt
Cosmopolitan
Diadiplosi duni and D. smithi preying on mealybugs, Planococcus spp. (Hemiptera: Pseudococcidae)
12 Tarsal claws each with two basal teeth [Fig. 16d]. ...............................................Gynodiplosis Felt
Oriental
Gynodiplosis humata causing leaf gall on Davallia repens (Davalliaceae)
- Tarsal claws each with single basal tooth. .................................................................13
13 Male flagellomeres each with 3 whorls of looped circumfila [Fig. 35g]. Female vertical horizontal circumfilar strands each with
about 10 connections to node. .........................................................Psephodiplosis Kolesik
Australasian and Oriental

GALL MIDGES OF INDONESIA Zootaxa 4847 (1) © 2020 Magnolia Press · 77
Psephodiplosis rubi causing leaf gall on Rubus moorei, R. moluccanus and R. alceifolius (Rosaceae)
- Male flagellomeres each with 2 whorls of looped circumfila [Fig. 34e]. Female vertical and horizontal circumfilar strands each
with about 5 connections to node. ........................................................Procontarinia Kieffer
Oriental and Neotropical (introduced)
Procontarinia matteiana and P. robusta causing leaf galls on mango Mangifera indica (Anacardiaceae)
14 Palpus 1-segmented. ..................................................................................15
- Palpus 3 or 4-segmented. ..............................................................................16
15 Male flagellomeres with 2 whorls of looped circumfila, the two nodes nearly equal in size [Fig. 39c]. Ovipositor nearly as long
as abdomen, the cerci long, narrow, closely appressed [Fig. 39g]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenodiplosis Rondani
Cosmopolitan
Stenodiplosis eragrostidis causing seed head swelling on Eragrostis unioloides (Poaceae)
- Male flagellomeres with 3 whorls of looped circumfila, the two nodes unequal in size [Fig. 42b]. Ovipositor short, barely pro-
trusible [Fig. 42f]. .............................................................Gnesiodiplosis” garcinia (Felt)
This species causes leaf gall on Garcinia dioica (Clusiaceae)
16 Tarsal claws robust, strongly bent at basal third [Fig. 25f]. ..................................... Oligoxenomyia Felt
Oriental
Oligoxenomyia radicis causing aerial root gall on Freycinetia sumatrana (Pandanaceae)
- Tarsal claws otherwise, curved or bent near or beyond midlength. ..............................................17
17 Aedeagus with setae [Fig. 20n]. Female flagellomeres with 3 nearly discrete, looped horizontal bands of circumfila [Fig.
20q]. ...................................................................... Leefmansiella Kolesik & Gagné
Oriental
Leefmansiella pandani causing leaf gall on Pandanus nitidus (Pandanaceae)
- Aedeagus without setae. Female flagellomeres with one [Fig. 40c] or two [Fig. 31d] horizontal bands of circumfila. ......18
18 Abdomen with purplish blob inside. Male hypoproct with ventrobasal lobe [Fig. 22c]. Female cerci with field of tiny setae
apicoventrally [Fig. 22h]. ............................................................... Lestodiplosis Kieffer
Cosmopolitan
Lestodiplosis lacciferi predator of scale insect Kerria javana (Coccoidea: Kerriidae)
Lestodiplosis oomeni preying on tea mite Calacarus carinatus (Acarina: Eriophyidae) tea plant Camellia sinensis (Theaceae)
Lestodiplosis syringopais, a likely predator associated with pitcher plants Nepenthes (Nepenthaceae)
- Abdomen without conspicuous purplish blob inside. Male hypoproct dorsoventrally flattened [Fig. 31b]. Female cerci without
field of tiny setae apicoventrally [Fig. 31e]. ................................................................19
19 Male cerci convex apically [Fig. 31b]. Female with definite, setose tenth tergum [Fig. 31e]. ..............Orseolia Kieffer
Palearctic, Afrotropical and Oriental
Orseolia apludae causing leaf gall on Apluda mutica (Poaceae)
Orseolia caulicola causing leaf gall on Sacciolepis indica (Poaceae)
Orseolia graminicola causing leaf gall on Cynodon dactylon (Poaceae)
Orseolia graminis causing leaf gall on Ottochloa nodosa (Poaceae)
Orseolia javanica causing leaf gall on cogon grass Imperata cylinbdrtica (Poaceae)
Orseolia orientalis causing leaf gall on Oplismenus compositus (Poaceae)
Orseolia oryzae causing leaf gall on rice Oryza sativa (Poaceae)
Orseolia paspali causing leaf gall on Paspalum scrobiculatum (Poaceae)
- Male cerci terminally or obliquely concave, mesal lobe formed by concavity with single seta. Female without definite, setose
tenth tergum [Fig. 40f].. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Male hypoproct bilobed apically [Fig. 40h]. Female flagellomeres constricted basally [Fig. 40c–d]. . . . . . . Thorodiplosis Felt
Oriental
Thorodiplosis impatiensis causing leaf gall on Impatiens platypetala (Balsaminaceae)
- Male hypoproct not incised apically [Fig. 17o]. Female flagellomeres with evenly cylindrical nodes. ......Hypodiplosis Felt
Oriental
Hypodiplosis paederiae causing leaf gall Paederia foetida (Rubiaceae)
21 Palpus 3-segmented [Fig. 41c]. ................................................... “Calopedila” polyalthiae Felt
This species causes fruit gall on Polyalthia subcordata (Annonaceae)
- Palpus 4-segmented [Fig. 11f]. ..........................................................................22
22 Ovipositor flattened bilaterally, cerci with pair of small horn-like sensoria dorsoapically and sparse setae elsewhere [Fig.
15g]. ............................................................................Gephyraulus Rübsaamen
Palearctic, Oriental, introduced to Neotropics
Gephyraulus tetrastigma causing leaf gall on Tetrastigma pergamaceum (Vitaceae)
- Ovipositor cylindrical, cerci with a few thickened setae apically and many thin setae scattered elsewhere [Fig. 11e]. ........
.................................................................................... Dasineura Rondani
Cosmopolitan
Dasineura elatostemmae causing stem and petiole gall on Elatostema paludosum (Urticaceae)
23 Pulvilli not unusually large, about 1/3 length of empodia. Ovipositor cylindrical, cerci partially setulose, with some scoop-
tipped setae [Fig. 19j]. ...................................................................Lasioptera Meigen
Cosmopolitan
Lasioptera javanica causing stem gall on Zehneria maysorensis (Cucurbitaceae)

KOLESIK & GAGNÉ
78 · Zootaxa 4847 (1) © 2020 Magnolia Press
Lasioptera manilensis causing leaf gall on Leea manilensis and L. indica (Vitaceae)
- Pulvilli large, about 2/3 length of empodia [Fig. 2e]. Entire protrusible part of ovipositor bilaterally flattened, glabrous, cerci
indistinguishable [Fig. 2g]. ..............................................................Actilasioptera Gagné
Oriental, Australasian
Actilasioptera falcaria and Actilsiosptera sp. causing leaf galls on mangrove Avicennia marina (Acanthaceae)
Acknowledgments
We thank Nami Uechi, National Agriculture & Food Research Organization, Tsukuba, Japan, for translation of
the Nakayama (1982) report on Asphondylia yushimai; Willy de Prins, Royal Belgian Institute of Natural Sci-
ences, Brussels, for providing a copy of Hering (1931) paper; the University of Padua, Italy, for the photograph of
Jenny Docters van Leeuwen-Reijnvaan; the Museum of the Tropics Amsterdam, Netherlands, for the photograph
of Willem Marius Docters van Leeuwen; Atsushi Kikuchi, National Agriculture and Food Research Organization,
Fukuyama, Japan, for photographs of Asphondylia yushimai galls [Figs 4d, e]; and Renée Staal, National Gallery
Singapore, and the family Tan, Singapore, for permission to reproduce the oil painting “River Valley in Java with
Mt. Gede and Mt. Pangrango in the Background” by Raden Saleh. Purnama Hidayat, IPB University, Bogor, Indo-
nesia, kindly provided photographs of Leefmansiella pandani [Figs 20c, d, h], the gall of Thorodiplosis impatientis
[Fig. 40b], the previously unpublished data on geographical distribution of L. pandani, S. villebrunneae and T. im-
patientis, and DNA sequences of Orseolia javanica and O. oryzae.
We thank the following colleagues and journals for granting permissions to reproduce previously published
figures: Nami Uechi for the photograph of Asphondylia capsicicola infestation [Fig. 4a] from Uechi et al. (2016,
Applied Entomology and Zoology); Keith M. Harris, Ripley, Surrey, UK, for figures of Procontarinia matteiana
[Figs 33f–i] from Harris (1966, Transactions of the Royal Entomological Society of London); of Diadiplosis duni
[Figs 12a–g], Diadiplosis smithi [Figs 13a–j], Lestodiplosis aonidiellae [Fig. 21d], Megommata leefmansi [Figs
24a, d–j], all from Harris (1968, Transactions of the Royal Entomological Society of London); and of Lestodiplosis
oomeni [Figs 22a–o] from Harris (1982, Entomologische Berichten); Wilma A. Augustyn, Tshwane University of
Technology, Arcadia, South Africa, for the photograph of Procontarinia matteiana galls [Fig. 33c] from Augustyn et
al. (2013, African Entomology); National Innovations on Climate Resilient Agriculture, New Delhi & Hyderabad,
India, for the photograph of Orseolia oryza galls [Fig. 31a] from Manual for Rice Pest Surveillance (2011); Wen-Jun
Bu, Nankai University Tianjin, China, for figures of Procontarinia robusta [Figs 34c, e–j] from Li et al. (2003, Acta
Zootaxonomica Sinica); Tomoko Ganaha-Kikumura, Prefectural Agricultural Research Center, Okinawa, Japan, for
the figure of Feltiella acarivora [Fig. 14f] from Ganaha-Kikumura et al. (2012, Applied Entomology and Zoology);
Philippine Journal of Science for figures of Asphondylia callicarpae [Fig. 3a], Asphondylia vitea [Figs 8a–b], Eu-
calyptodiplosis paederiae [Figs 17a–c] and Lasioptera manilensis [Figs 19a–c] from Uichanco (1919); Treubia for
figures of Leefmansiella pandani [Figs 20a, e–g, i, p] from Leefmans (1921); Entomologische Berichten for figures
of Megommata leefmansi [Figs 24b–c] from Barnes & Nijveldt (1954).
We are thankful to Ayman K. Elsayed (Botanical Gardens, University of Tokyo, Japan), Mathias Jaschhof
(Station Linné, Öland, Sweden) and Kai Heller (Heikendorf, Germany) for commenting on an early draft of the
manuscript.
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