ox

JHR 45: 113-123 (2015) JOURNAL OF *r*iercnoscloen
doi: 10.3897/JHR.45.5274 (ME Hymenopter a

http://jhr.pensoft.net The insertional Society of ymenopterists RESEARCH

Clarification of the author and year of publication of
Cotesia chilonis, a species used widely for biological
control of Chilo stem borers

Jose Fernandez-Triana', John S. Noyes’, Andrew Polaszek*, Dicky S. K. Yu!

| Canadian National Collection of Insects, Ottawa, ON, Canada 2 Department of Life Sciences, Natural
History Museum, London, UK

Corresponding author: Jose Fernandez-Triana (jftriana@uoguelph.ca)

Academic editor: Gavin Broad| Received 11 May 2015 | Accepted 20 August 2015 | Published 7 September 2015
Attp:/zoobank.org/D1CA491 C-67FC-4553-823F-A2497 1 1FB69E

Citation: Fernandez-Triana J, Noyes JS, Polaszek A, Yu DSK (2015) Clarification of the author and year of publication
of Cotesia chilonis, a species used widely for biological control of Chilo stem borers. Journal of Hymenoptera Research 45:

113-123. doi: 10.3897/JHR.45.5274

Abstract

Cotesia chilonis (Munakata, 1912) (Hymenoptera: Braconidae, Microgastrinae) is a species used widely
for biological control of several species of Chilo (Lepidoptera: Crambidae) stem borers. The genus name,
species author and year of publication associated with this parasitoid wasp species have varied historically.
Based on the available evidence, a complete account of the history and nomenclatural acts associated with
C. chilonis is presented, the different alternatives are weighed and discussed, and what is considered as the
best alternative is proposed. It is expected that this paper will contribute to clarity and stability in the use

of this species name, author and year.

Keywords

Cotesia chilonis, Microgastrinae, Chilo stem borers, biocontrol, taxonomy, Japan

Copyright Jose Fernandez-Triana et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

114 Jose Fernandez-Triana et al. / Journal of Hymenoptera Research 45: 113-123 (2015)

Introduction

Cotesia chilonis (Munakata, 1912) is a parasitoid wasp (Braconidae: Microgastrinae)
originally described from Japan and widely used in biological control projects against
caterpillar stem borers in the genus Chilo (Lepidoptera: Crambidae), pests of grami-
naceous crops in the Oriental and Afrotropical regions (Kfir et al. 1992, Polaszek and
Walker 1992, Polaszek 1998).

Cotesia chilonis belongs to the Cotesia flavipes species complex, which includes
four species parasitizing stemboring pests associated with sugarcane and cereal crops
(Walker 1994, Muirhead et al. 2008). The monophyly of the complex is well support-
ed by molecular (Smith and Kambhampati 1999; Michel-Salzat and Whitfield 2004;
Muirhead et al. 2006, 2008) and morphological characters, such as a dorsoventrally
compressed mesosoma (Watanabe 1965; Walker 1994, Muirhead et al. 2008).

The correct genus of the species, as well as its author and publication year (= origi-
nal description) have long been in a state of confusion. As a result, the considerable
literature on C. chilonis is replete with errors. The species has been placed either in
Apanteles or in Cotesia, the authorship has been attributed either to Tetsuzo Munakata
or to Shonen Matsumura, and the year of the original description has been cited as
either 1910 or 1912.

Because of the importance of this species in biological control, the generic place-
ment, author and year of description of Cotesia chilonis must be clarified.

Methods

We carefully examined the scientific literature relating to this species, including the
report in which the species was originally described (Anonymous 1912). We made an
effort to study all taxonomic references to Cotesia chilonis that we are aware of, and also
consulted several colleagues who are proficient in Japanese and/or had seen the original
paper (see Acknowledgments).

We estimated the number of citations for the different variants of the species
names, species author, and year of publication by performing searches on Zoological

Record, Google Scholar, and Google.

Results

Cotesia chilonis was originally described in Apanteles (the original description has ‘Apanteles’
misspelled as ‘Apantetus’, see Fig. 1). However, Mason (1981) divided Apanteles into several
genera, either new or resurrected (such as Cotesia Cameron, 1891) and transferred many
known species, but unfortunately not Apanteles chilonis, to his newly proposed genera. Ma-
sons (1981) study was instrumental in highlighting the polyphyly of Apanteles s.1., a finding
that led to the separation of Cotesia and Apanteles s.str. as separate, unrelated genera.

115

Clarification of the author and year of publication of Cotesia chilonis...

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Figure |. Copy of page 69 of the 1912 anonymous report with the original description of Cotesia chilonis

(Munakata, 1912). The red square shows the original species name and attributed author.

116 Jose Fernandez-Triana et al. / Journal of Hymenoptera Research 45: 113-123 (2015)

Austin and Dangerfield (1989: 137) were the first authors to propose une-
quivocally the new combination, and to formally transfer the species to Cotesia
(although they did so briefly in a paper section discussing unrelated species of
Apanteles, and the nomenclatural act was not mentioned elsewhere). One year later
Papp (1990: 117), unaware of that reference, proposed the same new combination
(he did so in an Appendix rather unrelated to the main content of his paper, which
was devoted to reporting Microgastrinae species from Korea). The fact that the
new nomenclatural act was not clearly visible or referenced in either of those two
papers probably contributed to subsequent overlooking of the new generic place-
ment of chilonis. [Both Austin and Dangerfield (1989) and Papp (1990) attributed
the authorship of the species to Matsumura instead of Munakata, see below for
more discussion on that].

Cotesia has been universally recognized and accepted as a valid genus by the taxo-
nomic and biological control research community (e.g., Yu et al. 2012, and references
to the genus cited therein). However, due to the confusing circumstances detailed
above, the species C. chilonis has continued to be variously placed in either Apanteles or
Cotesia. For example, basic searches made in November, 2014 using the names ‘Apan-
teles chilonis or ‘Cotesia chilonis revealed, respectively 35 and 28 citations in Zoologi-
cal Record and, 651 and 714 citations in Google Scholar. At the same time a general
search in Google reveals 4,280 and 6,020 results, respectively.

Searches performed with Zoological Record, Google Scholar, and Google revealed
an equally confusing mix for the author and publication year of C. chilonis (Table 1).
Cotesia chilonis was originally described in a report of the Agriculture Experiment Sta-
tion of Aomori, Japan. The report does not clearly state its author, but apparently it
was written by Tetsuzo Munakata, although the species is mentioned in the paper as
‘Apantetus chilonis Mats.’ (Fig. 1).

Watanabe (1965) provided additional and useful information on the author and
year of publication, and it should be taken as the best information available to settle
the issue. Watanabe (1965: 114, reproduced in Fig. 2) states that “In 1912 Apanteles
chilonis was originally described anonymously as a parasite of the rice stem borer,
Chilo suppressalis, but there are certain evidences that the author is Tetsuzo Munaka-
ta.” The next sentences from Watanabe (see Fig. 2) suggest that Munakata sent the

Table I. Searches performed in November 2014 on three different online databases or search engines,

using two different strings of words: “Apanteles+Cotesia+chilonis+Matsumura” or “Apanteles+Cotesia+

chilonis+Munakata”.
Online Database/ Apanteles/ Cotesia chilonis
Search Engine Matsumura (Total) Munakata (Total)
Zoological Record 9
Google Scholar 49
Google 256

Clarification of the author and year of publication of Cotesia chilonis... 117

174

Okinawa, Mauritius and Australia.

Apanteles chilonis (Munakata)

Apantetus (!) chilonis (Munakata), Extra Report Agr. Exp. Sta. Aomori No. 2:
69, 1912. JAPAN: Aomori. Host: Chilo suppressalis (= C. simplex),

Apanteles (Stenopleura) chilocida Viereck. Proc. U. S. Nat. Mus. 43: 582, 1913.
JAPAN. Host: Chilo “suppressalis.

Apanteles chilocida: Wilkinson, Bull. Ent. Res. 19: 94, 1928,

Apanteles flavipes f. chilonis: Watanabe, Trans. Sapporo Nat. Hist. Soc. 12: 69,
1932.

In 1912 Apanteles chilonis was originally described anonymously as a parasite
of the rice stem borer, Chilo suppressalis, but there are certain evidences that the
author is Tetsuzo Munakata, The type is lost, and yet in the Entomological
Institute, Hokkaido University, are deposited authentic material (4 29, Aomori,
1911, on a block of pith with a determination label —“ Apanteles chilonis n. sp.” —
in Matsumura’s own hand-writing) which was sent by Munakata to Prof. Matsu-
mura for identification. From the series I have selected as neotype of chilonis
the single female which is in the best condition. Just after the » above-mentioned
publication Viereck (1913) described originally Apanteles chilocida as a parasite of
the same host. This name, however, already has been Sumeressen as a synonym
of chilonis by Watanabe (1932).
is Insect 1s So Closely related to A. flavipes that there are two different views
about the taxonomy of the Japanese form: Wilkinson (1928, etc.) treats it as a
full species; on the other hand Watanabe (1932, etc.) considers it a form of
flavipes. Having examined a number of specimens of both the forms I am much
inclined to the opinion that chilonis should be treated as a distinct species.
Differences are noticeable between the two in the punctuation of the prothorax,
mesopleura and hind coxae as well as in the colouration of the hind coxae as
states in the present key. These differences would seem to be more reliable to
separate the two than the prominence of the face and the declivity of the propo-
deum which are characters used by Wilkinson in his key: because in chilonis those
features tend to become inconspicuous, while in flavipes they seem to be very
variable in strength,

In general, this species is darker in colour than flavipes: the antennae are yel-
lowish brown to brown, not reddish yellow as in flavipes. Having examined the
genitalia of both sexes I have found that there are no special differences between
the two species in genital structures. The first abscissa of the radius is not
clearly shorter than the transverse cubitus but as long as or sometimes rather
longer than the latter.

JAPAN: Aomori, 4 29 (including the neotype of chilonis), 1911 (T. Munakata).
Many other specimens examined are collected at the following localities: —Kyu-
shu: Taniyama (S. Fukamachi); Yokkaichi and Fukuoka (lI. Tateishi) ; Futsu-
kaichi (K. Yano); and Miyazaki (H. Sakimura). Honshu: Masuda, ‘Taito, Hikawa,
Ota and Hamada (T. Fujishima) ; Kanazawa (H. Kawase); Shonai and Omagari
(T. Hidaka); Nishiki-mura and Omagari (M. Kikuchi); and Morioka CH. Omori).

Figure 2. Copy of page 114 of Watanabe (1965). The red square shows the most relevant paragraphs.
The words underlined with gray were made by an unknown source to the copy available to us for study,
and are not related to the present paper.

specimens to Shonen Matsumura, who identified the species as “Apanteles chilonis n.
sp. and sent it back to Munakata, who then published the report with the species
description in 1912.

118 Jose Fernandez-Triana et al. / Journal of Hymenoptera Research 45: 113-123 (2015)

From the above account, it seems evident that Matsumura was the one
recognizing the new species, but Munakata was the one publishing it. This opinion
from Watanabe agrees with that of other Japanese entomologists, e.g. the braconid
specialist Kaoru Maeto (personal communication) and the ichneumonid specialist
Jinhaku Sonan (1930: 141). Actually, the 1930 paper from Sonan provides an
interesting fact. Sonan was dealing with another species described in the 1912 report
(the ichneumonid ‘Ophionellus biguttulus currently named ‘Temelucha biguttula’),
and he also considered that, even though the original paper had written ‘Ophionellus
biguttulus Mats.’, Matsumura did not describe the species but it was Munakata
(compare Figs 3 and 4).

Because the author of the species is not clear from the contents of the 1912 report,
the author of the report (Munakata) should also be considered as the author of the
species, as explained in The International Code of Zoological Nomenclature (ICZN).
Article 50.1.1 states “However, if it is clear from the contents that some person other
than an author of the work is alone responsible both for the name or act and for sat-
isfying the criteria of availability other than actual publication, then that other person
is the author of the name or act. “Jf the identity of that other person is not explicit in the
work itself, then the author is deemed to be the person who publishes the work.” The last
sentence (our italics) clearly applies here, and it should be used to recognize Munakata
as the author of the species.

Watanabe, who should also be considered as the First Reviser, sensu Article 24.2
of the ICZN, unambiguously stated the authorship of the species in two papers: Wa-
tanabe (1932a: 69) and Wanatabe (1965: 114). Unfortunately he was not followed
by subsequent authors. Most notably, the latest comprehensive list of world species of
Braconidae (Shenefelt 1972: 468) mentioned Matsumura as the author. Shenefelt was
probably the source of the mistake followed by other important references on Micro-
gastrinae (Table 2) which unintentionally contributed to perpetuating the mistake —at
least partially— until the present.

Last but not least, the year of publication has, at times, been confused by some
sources (Table 2). The most common mistake (1910 instead of 1912) might have
originated from another paper of Shenefelt (1965: 400), which misspelled Munakata
as ‘Munakato’ and incorrectly assigned 1910 as the date for his report on Cotesia chilo-
nis. For clarification about the actual date, we again follow Watanabe (1932a, 1932b,
1965), as he actually saw the report, studied Munakata’s specimens, and even desig-
nated a neotype (Watanabe 1965: 114; see also Fig. 2).

The unfortunate chain of omissions and confusion in publications on Cotesia chilo-
nis has led to this important species to be referred to by several generic names, authors,
and years of publication. For the sake of clarity and stability, it is useful to provide in a
single paper a complete account of the history and nomenclatural acts associated with
this species. Based on the revised evidence, the history of names associated with C.
chilonis is presented below. It is hoped that the correct names and dates (as presently
understood) are adopted by future researchers.

Clarification of the author and year of publication of Cotesia chilonis... 1L9

J. Sonan: A few host-known Ichneumonidae found in Formosa (Hym.) (2). 141

*/s a8 long as the body; mesonotum abruptly sloping apically,
closely punetate and shortly pubescent, notauli obsolete; scutellum_
convex, closely rugose; propodeum rugose, with a rather broad,
but sharrow median furrow, which is t:ansversely striated, external
and lateral areae distinet; spiracles elliptical; first tergite linear,
glabrous, apex fusiform; 4th-9th tergites strongly compressed late-
rally throughout; tarsal clays -pectinate.

Black. Apex of mandibles reddish brown; palpi, tegulae and
legs yellowish brown; antennae, coxae and hind trochanter black;
hind femolae reddish brown, base and apex of tibiae, apex of
metatarsi and 2nd-5th joints dark brown; wings hyaline, stigina
brown, arcolet somewhat large, triangular, with a distinctly petiole;
recurrent nervure straight, emiting from a little beyond the middle
of areolet; basal nervure continuous.

Length: Body 8mm; ovipositor 4mm; fore wing 5mm,

Host. Larvae of Homona imenciana Walker (Author).

‘Hab. Type 9, Shiuchiku, Apr. 4, 1921 (Auhtor): Paratype
9, Heichin (Shinchikn-Shu), May 1, 1930 (Author).

This species is allied to Limuertum forticarinatum Cameron
(1906) and Z. guettaeus Cameron (1906), but differs from them
by the wanting of areola of propodeum, elliptical spiracles, yello-
wish brown legs, aud black apices of hind tibiae.

CREMASTUS, Cravenhorst. Ichn. Europ. iii, p. 7380(1829).

Subgen. TARYTIA Cameron, Jown, Bombay Nat. Hist.
Soo. p. 588 (1907)t *7OY
CREMASTIDEA, Vieveck, Proce. U. 8. Nat. Mus, 48,
O no, 1942 p, 587 (1912).
| | *."| Cremastus (Tarytia) biguttulus (Munskata). _.
Ophioellus biguttulus Matsumura (non desc.), Munakata.
|, Exira Report Agric. Exp. Stat. Aomori, no. 2, p. 67, pl. ii

142 ram. N. H. 8
Trans, N. H. 8. F. Vol. XX. (Jun. 1930)

fig, 19 (1910); Kondo, Extra Rept. Agric. Stat, Naga
a7 15, p. 101 (1917); Nawa, Insect world Gifu
219, p. 456 (1915). (Japan).
2 4S Cremastus (C. reutastidea) chinensis Viereck, Proc, U.S Nat
Mus. 4: 5 aa
a sd 2 1942, p. 587, @ ¢ (1912). (China, Japan).
sane Japonica \Uchida (nee Ashmead), Jour. Fac
ches f Hokk, Imp. Uniy Sapporo, xxi, pt. 2, p» 285, pl
Vi, fig, 16, © (1928) ar

gaki,

xix, no.

Figure 3. Copy of pages 141 and 142 (partially) of Sonan (1930). The red square shows Sonan’s inter-
pretation of the species author for one of the species described by Munakata in 1912. The annotations

added with a pencil (in gray) were made by an unknown source to the copy available to us for study, and

are not related to the present paper.

Jose Fernandez-Triana et al. / Journal of Hymenoptera Research 45: 113-123 (2015)

120

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NK \S ERATED NRBR+ RA ae RR DS

i
i

| SX RX fie to

BIIt+H tu Hi BARA
REMI VRANE HHI ON RRMMaAK AH NK BKR NCE 1] BY RI RA
Bes Ce PLR AMRA ISH KL eNO RBRVS Ihe RH RS
Ri SROVMABAOMMUBI BAIN. OeESRT IBM SAIBrNEaAn&
HN ER WRK ARRAY A HR eR BRARAN RM CARAS BB
Xv RB BXNRBDAN BRENNA Red -+~ BAN RKR( HERE
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Hm rn MRYVEMNMRm er. enna’
H+rileRHWe+Ae¢eBe
I7 AwSTHRHADRA WS (SR 1) BEN |)
Rb ff Ophione!lus biguttulus = Mats. | |

KY?

Figure 4. Copy of page 67 of the 1912 anonymous report. The red square shows the original species

name and attributed author (compare it against Figure 3 above).

Clarification of the author and year of publication of Cotesia chilonis... 121

Table 2. Important published and/or web resources about Microgastrinae with differing accounts for the
species Cotesia chilonis (Munakata, 1912). Comments in square brackets provide additional information

when needed.

a) Sources with Munakata as author of the species, and publication year of 1912 (unless a

different date is indicated in square brackets):

- Taxapad (http://www.taxapad.com/local.
php?&indexstartGS=53 166&taxgroupGS=ichneumonoidea) [As Cotesia chilonis (Munakata,
1910), the year 1910 and the authorship used in Taxapad were based on the following
reference: Shenefelt, R.D. 1965. A contribution towards knowledge of the world literature
regarding Braconidae (Hymenoptera: Braconidae). Beitrige zur Entomologie. 15(3/4):243-
500. On page 400, the Munakata paper is dated as 1910 — and the name Munakata is
misspelled as Munakato].

- Catalog of Life (http://www.catalogueoflife.org/annual-checklist/2014/search/all/key/
Cotesia+chilonis/match/1).

- GBIF (http://www.gbif.org/species/ 1257878).

- Encyclopedia of Life (http://eol.org/pages/ 1206354 1/overview).

- Wikipedia (http://en.wikipedia.org/wiki/List_of_Cotesia_species) [Merely a citation from EOL].

- Microgastrinae Wasps of the World (http://microgastrinae.myspecies.info/microgastrinae/
Cotesia-chilonis).

- Barcode of Life Data Systems (http://www.boldsystems.org/index.php/Public_
Search Terms? query=%22 Cotesia%20chilonis%22|tax]) [The records included in BOLD were
mined from GenBank].

b) Sources with Matsumura as author of the species, and publication year of 1912 (unless a

different date is indicated in square brackets):

- ZooBank (http://zoobank.org/Search?search_term=Apanteles+ chilonis) [As Apanteles chilonis
Matsumura, 1912. ZooBank does not have any record as ‘Cotesia chilonis’).

- Shenefelt 1972 (Braconidae 4. Microgasterinae: Apanteles. Hymenopterorum Catalogus, 1972:
page 468) [This paper represents the last comprehensive list of world species of Braconidae].

- Whitfield Lab website (http://www.life.illinois.edu/whitfield/Cotesia/ Cotesia.htm) [As Cotesia
chilonis Matsumura, 1926; the year is most likely a typographical mistake. This website
contains a significant number of Cotesia species illustrations and is a valuable reference on
the topic].

- Sharkey Lab website (http://www.sharkeylab.org/Cotesia/ Cotesia.cgi) [This website contains a
significant number of Cotesia species illustrations and is a valuable reference on the topic].

Cotesia chilonis (Munakata, 1912)

Apantetus chilonis Munakata, 1912: 69. Original description. Misspelling of the genus
name.

Apanteles chilonis Matsumura in Munakata, 1912: 69. Shenefelt 1972: 468. Incorrect
assignment of author of the species. [Subsequently followed by a substantial num-
ber of authors, websites, and online databases].

Apanteles flavipes £. chilonis Watanabe, 1932: 84. Junior subjective synonym [In the
References below, this paper is cited as “Watanabe 1932b’ to distinguish it from
another work published by Watanabe in 1932 and also cited in the present paper].

Apanteles chilonis Munakata, 1912. Watanabe 1965: 114. Reinstatement as valid species.

122 Jose Fernandez-Triana et al. / Journal of Hymenoptera Research 45: 113-123 (2015)

Cotesia chilonis (Matsumura, 1912). Austin and Dangerfield 1989: 137. New combin-
ation. Incorrect assignment of author of the species.

Cotesia chilonis (Matsumura, 1912). Papp 1990: 117. Unnecessary ‘new combination’.
Incorrect assignment of author of the species.

Cotesia chilonis Munakata, 1910. Yu et al. 2012. Incorrect assignment of date for
species description.

Apanteles chilocida Viereck, 1912: 582. Watanabe 1932: 69. Junior subjective syno-
nym. Incorrect date (1913) assigned to Viereck’s paper. [In the References below,
this paper is cited as “Watanabe 1932a’ to distinguish it from another work pub-
lished by Watanabe in 1932 and also cited in the present paper].

Acknowledgements

The decisions made in this paper about names, authors and dates related to Cotesia
chilonis benefited considerably from fruitful discussion with several colleagues. Dr John
Steel (University of Otago, New Zealand) helped in establishing contact with Japanese
colleagues and kindly sent copies of some of the most important references used for
this paper. Dr Katsuhiko Sayama (Hokkaido Research Center, Forestry and Forest
Products Research Institute, Japan) kindly provided useful comments and copies of
some of the most important references used for this paper. Dr Kaoru Maeto (Kobe
University, Japan) kindly shared his opinions on the names and authors historically
associated with the species. Dr John Huber and Dr Gary Gibson (Canadian National
Collection of Insects, Ottawa, Canada) revised earlier versions of the manuscript and
helped to clarify contents.

References

Anonymous (1912) Extra Report Agric. Exp. Stat. Aomori 2: 67-69.

Austin AD, Dangerfield PC (1989) The taxonomy of New World microgastrine braconids
(Hymenoptera) parasitic on Diatraea spp. (Lepidoptera: Pyralidae). Bulletin of Entomo-
logical Research 79(1): 131-144. doi: 10.1017/S0007485300018642

Kfir R, Overholt WA, Khan Z, Polaszek A (2002) Biology and management of economically
important lepidopterous cereal stem borers in Africa. Annual Review of Entomology 47:
701-731. doi: 10.1146/annurev.ento.47.091201.145254

Mason WRM (1981) The polyphyletic nature of Apanteles Foerster (Hymenoptera: Bra-
conidae): A phylogeny and reclassification of Microgastrinae. Memoirs of the Entomo-
logical Society of Canada, Ottawa, Canada, 147 pp.

Michel-Salzat A, Whitfield JB (2004) Preliminary evolutionary relationships within the para-
sitoid wasp genus Cotesia (Hymenoptera: Braconidae: Microgastrinae): combined analysis

of four genes. Systematic Entomology 29: 1-12. doi: 10.1111/j.0307-6970.2004.00246.x

Clarification of the author and year of publication of Cotesia chilonis... 123

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Papp J (1990) Braconidae (Hymenoptera) from Korea. XII. Acta Zoologica Hungarica 36(1-—2):
87-119.

Polaszek A (Ed.) (1998) African cereal stem borers: Economic importance, taxonomy, natural
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Polaszek A, Walker AK (1992) The Cotesia flavipes species-complex: parasitoids of cereal stem
borers in the tropics. Redia 74: (3) “Appendice” (1991): 335-341.

Shenefelt RD (1965) A contribution towards knowledge of the world literature regarding Bra-
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Shenefelt RD (1972) Braconidae 4. Microgasterinae: Apanteles. Hymenopterorum Catalogus
(nova editio), Pars 7, 429-668.

Smith PT, Kambhampati S (1999) Status of the Cotesia flavipes species complex (Braconidae:
Microgastrinae) based on mitochondrial 168, rRNA and NADH 1 dehydrogenase gene
sequence. Journal of the Kansas Entomological Society 72: 306-314.

Sonan J (1930) A few host-known Ichneumonidae found in Formosa (Hym.) (2). Transactions
of the Natural History Society of Formosa. Taihoku 20: 137-144.

Walker AK (1994) Species of Microgastrinae (Hymenoptera: Braconidae) parasitizing lepidop-
terous cereal stem borers in Africa. Bulletin of Entomological Research 84: 421-434. doi:
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