characters of the external female genitalia which can … · elena mădălina morariu are for...
TRANSCRIPT
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
CHARACTERS OF THE EXTERNAL FEMALE GENITALIA
WHICH CAN BE USED IN THE SYSTEMATICS OF
PENTATOMINAE (HETEROPTERA: PENTATOMIDAE)
Elena Mădălina MORARIU
Faculty of Biology, Alexandru Ioan Cuza University of Iași, B-dul Carol I, no. 20A, 700505 Iași, Romania;
Abstract. In this paper we analyse the external female genitalia of 10 species (Aelia acuminata (Linné, 1758);
Eysarcoris aeneus (Scopoli, 1763); Stagonomus amoenus (Brullé, 1832); Rubiconia intermedia (Wolff, 1811);
Holcosthetus vernalis (Wolff, 1804); Carpocoris purpureipennis (De Geer, 1773); Dolycoris baccarum (Linné,
1758); Piezodorus lituratus (Fabricius, 1794); Pentatoma rufipes (Linné, 1758); Eurydema ornata (Linné, 1758))
belonging to genera of the subfamily Pentatominae. The external female genitalia of Stagonomus amoenus are
considerably distinctive from the other female genitalia of the subfamily Pentatominae.
Keywords: external female genitalia, laterotergites, gonocoxites, spermatheca, Pentatomidae.
Rezumat. Caractere ale genitaliei externe la femelă, utile în sistematica subfamiliei Pentatominae
(Heteroptera: Pentatomidae). În această lucrare analizăm genitalia externă la femele aparținând la 10 specii din
(Aelia acuminata (Linné, 1758); Eysarcoris aeneus (Scopoli, 1763); Stagonomus amoenus (Brullé, 1832);
Rubiconia intermedia (Wolff, 1811); Holcosthetus vernalis (Wolff, 1804); Carpocoris purpureipennis (De Geer,
1773); Dolycoris baccarum (Linné, 1758); Piezodorus lituratus (Fabricius, 1794); Pentatoma rufipes (Linné,
1758); Eurydema ornata (Linné, 1758)) genuri ale subfamiliei Pentatominae. Genitalia femelă, în cazul speciei
Stagonomus amoenus este evident diferită de ale altor specii ale aceleași subfamilii.
Cuvinte cheie: gentalia externă la femelă, laterotergite, gonocoxite, spermateca, Pentatomidae.
Introduction Pentatomidae is a family of bugs with the body covered in a strong cuticula (Kis,
1984; Schuh & Slater, 1995). The antennae are made up of five joints. They present 2 ocelli
(Kis, 1984; Derjanschi & Péricart, 2005; Rider, 2006). Many of the species feed on plants,
some of them damaging the croplands (Panizzi et al., 2000; Mihalache et al., 1969; Kis,
1984; Schuh & Slater, 1995). As a particular feature of the subfamily Pentatominae,
scutellum is triangular, shorter than the abdomen and it does not cover the hemielytra (Kis,
1984; Derjanschi & Péricart, 2005).
Some of the first authors who started analysing external genitalia on Pentaomoidea
were Crampton (1920, 1922) and Dupuis (1951, 1952, 1953, 1970). They described both
male and female genitalia. External male genitalia are easier to be analysed since
noninvasive methods of research can be used (Dupuis, 1951, 1953; Štys, 1961; Tay, 1969;
Schaefer, 1968; Gapon, 2001). Dupuis described the external female genitalia, particularly at species belonging to genus Eurydema (Dupuis, 1951, 1953, 1970). He analysed the
structures resulted from modifications of the segments eight and nine. McDonald (1966)
started to study the external genitalia of the North American genera of Pentatomoidea in
order to clarify the relationships between them. He made a key using only features of the
external male genitalia. The external female genitalia are used for describe difficult genera
- 31 -
Elena Mădălina Morariu
are for reviews. Derjanschi and Péricart in „Faune de France” use female genitalia only for
difficult genera (Aelia, Sciocoris) (Derzhansky & Péricart, 2005).
The objective of this paper is to examine the female genitalia of Pentatominae and
to establish the characteristics useful for systematics.
Material and Methods
The external female genitalia of 10 species in 10 genera were studied (Tabel 1).
Almost all the species were collected with the entomological net from two Romanian (RO)
counties: Iasi (IS) (Botanical Garden, Ciric area, Bârnova forest) and Vrancea (VN)
(Paltin). They were kept in alcohol 70% until they were mounted on cards. The mounting
was made in such a manner that the taxonomical features were easy to notice. This way, the bugs are better preserved (Millar et al., 2000).
When studying the external female genitalia, we first detached the last segments of
the abdomen (sixth-tenth segments) and then boiled them in NaOH 10% for about 15-20
minutes. Afterwards, we washed them with alcohol, and easily detached the external female
genitalia. We embedded them in Faure solution and drew them with a Reichart drawing
tube.
Morphological terms used in text. The female genitalia of Pentatomidae are the
plate-shaped type, situated on abdominal segments eight and nine. The tenth and eleventh
segments represent the anal segments (Dupuis, 1951, 1955, 1970; Štys, 1961; McDonald,
1966; Schaefer, 1968, 1998; Kis, 1984; Derjanschi & Péricart, 2005). There are two
categories of components of the external genitalia which can be used in taxonomy: the structures resulted from modification of the eighth and ninth segments and receptaculum
seminis (spermatheca) (Dupuis, 1970).
The last visible tergite is the eighth one. It is folded ventrally on each side and
forms the laterotergites. Each laterotergite bears an articulated gonocoxite (Dupuis, 1970;
Derjanschi and Péricart, 2005), also called valvifer (Snodgrass, 1935; Schaefer, 1968).
Gonocoxites carry the gonapophyses (Dupuis, 1970; Derjanschi and Péricart, 2005), also
called valvulae (Snodgrass, 1935; Schaefer, 1968). As compared to some other groups of
bugs, gonapophyses are strongly reduced, determining the gonocoxites and laterotergites to
become the most important parts of the ovipositor. Their main role is to protect the genital
aperture (Dupuis, 1951, 1970).
The parts of the first pair of valvifers are generally separated from each other. The
second pair of valvifers is fused on the median line. The first pair of valvulae is fused in one piece, triangulum, and the second pair of valvulae is fused in one piece, arcus (Dupuis,
1970; Schaefer, 1968).
Receptaculum seminis (spermatheca) presents the following parts: an orificium
receptaculi, ductus receptaculi (spermathecal duct), a pars intermedialis (spermathecal
pump) and capsula seminalis (Snodgrass, 1935; Dupuis, 1970; Derjanschi & Péricart,
2005).
Results
Aelia acuminata (Linné, 1758) (Plate 5, Fig. 2)
The first pair of valvifers, laterotergites 8, the second pair of valvifers and
laterotergites 9 are covered up with long setae, symmetrically distributed. Capsula seminalis is globose, with two lateral long expansions which surpass the anterior annular
- 32 -
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
creste. The posterior annular creste is well developed as the anterior annular creste. The
latter is narrow.
Table 1. List of specimens examined.
Species Number of specimens
Data labels
Aelia acuminata 3 Ciric area (IS; RO), 28.06.2007 3 Botanical Garden (IS; RO), 9.07.2007 5 Bârnova (IS; RO), 18.07.2007 2 Bârnova (IS; RO),18.08.2007 2 Botanical Garden (IS; RO), 28.09.2007 3 Botanical Garden (IS; RO), 7.05.2008 2 Ciric area (IS; RO), 10.05.2008 1 Botanical Garden (IS; RO), 30.06.2008 6 Botanical Garden (IS; RO), 29.07.2008
Eysarcoris aeneus 1 Botanical Garden (IS; RO), 9.07.2007 1 Bârnova (IS; RO), 18.07.2007 1 Paltin (VN; RO), 8.07.2009
Stagonomus amoenus 1 Botanical Garden (IS; RO), 9.07.2008
Rubiconia intermedia 1 Botanical Garden (IS; RO), 9.07.2007 5 Bârnova (IS; RO), 18.07.2007 1 Botanical Garden (IS; RO), 28.09.2007
Holcosthetus vernalis 1 Bârnova (IS; RO), 18.07.2007 ♀, 2 Paltin (VN; RO) (leg. Chiriac N.), 8.07.2009 2 Paltin (VN; RO), 10.07.2009
Carpocoris purpureipennis 1 Ciric (IS; RO), 28.06.2007 6 Bârnova (IS; RO), 18.07.2007 1 Botanical Garden (IS; RO), 28.09.2007 1 Botanical Garden (IS; RO), 9.07.2008 2 Botanical Garden (IS; RO), 29.07.2008 1 Botanical Garden (IS; RO), 12.08.2008
Dolycoris baccarum 1 Ciric (IS; RO), 28.06.2007 1 Botanical Garden (IS; RO), 9.07.2007 2 Bârnova (IS; RO), 18.07.2007 1 Bârnova (IS; RO), 18.08.2007 1 Botanical Garden (IS; RO), 28.09.2007 1 Ciric (IS; RO), 10.05.2008 1 Bârnova (IS; RO), 27.06.2009 1 Bârnova (IS; RO), 5.07.2009 3 Paltin (VN; RO), 8.07.2009
Pentatoma rufipes 10 Bârnova (IS; RO), 18.07.2007 2 Bârnova (IS;RO), 18.08.2008 1 Bârnova (IS; RO), 22.08.2009
Piezodorus lituratus 1 Botanical Garden (IS; RO), 29.07.2008 1 Paltin (VN; RO), 8.07.2009
Eurydema ornata 2 Bârnova (IS; RO), 18.07.2007
Eysarcoris aeneus (Scopoli, 1763) (Plate 3, Fig. 1)
The first pair of valvifers, laterotergites 8 and the second pair of valvifers,
laterotergites 9 and the tenth segment of the abdomen are covered with both long and short
setae. They are symmetrically distributed. Capsula seminalis is globose, without
expansions. The posterior annular creste well developed. The anterior annular creste is narrow.
Stagonomus amoenus (Brullé, 1832) (Plate 3, Fig. 2)
The first pair of valvifers, valvulae and laterotergites 8 are merged. The tenth
segment of the abdomen and the laterogites 9 are sharp at the posterior border. The anterior
- 33 -
Elena Mădălina Morariu
border of the tenth segment of the abdomen is concave. Capsula seminalis is ovoid. The
proportion between length and width of capsula seminalis is 1.6. The anterior annular creste
is narrow. The posterior annular creste is difficult to notice.
Rubiconia intermedia (Wolff, 1811) (Plate 4, Fig. 1)
The first pair of valvifers, laterotergites 8, the second pair of valvifers,
laterotergites 9 and the tenth segment of the abdomen are covered with long setae. Capsula
seminalis is globose, with three spur like expansions. The anterior annular creste is narrow.
The posterior annular creste is difficult to notice.
Holcosthetus vernalis (Wolff, 1804) (Plate 4, Fig. 2)
The first pair of gonocoxites, laterotergites and the second pair of laterotergites
and gonocoxites are covered up and the tenth segment of the abdomen are covered with setae. The anterior border of the gonocoxite 9 is concave. Capsula seminalis is ovoid; the
proportion between length and width of capsula seminalis is 1.8. Capsula seminalis has one
sleazy expansion.
Carpocoris purpureipennis (De Geer, 1773) (Plate 3, Fig. 3)
The first pair of gonocoxites is covered with setae only on the posterior and
internal borders. Gonocoxites 9, laterotergites 8 and 9 and the tenth segment of the
abdomen are covered with setae regularly distributed. Capsula seminalis is globose without
any expansions. The posterior annular creste is well developed. The anterior annular creste
is very well developed.
Dolycoris baccarum (Linné, 1758) (Plate 4, Fig. 3)
Gonocoxites 8, laterotergites 8 and 9 and the tenth segment of the abdomen are covered with setae. The anterior annular creste is broad. The posterior annular creste is well
developed. Capsula seminalis is globose without any expansions.
Pentatoma rufipes (Linné, 1758) (Plate 5, Fig. 1)
Gonocoxites 8 are covered with numerous long setae. Gonocoxites 9, laterotergites
8 and 9 and the tenth segment of the abdomen is covered with long setae. The
gonapophyses 9 is strongly concave. Capsula seminalis is globose with two long
expansions which surpass the anterior annular creste. The posterior annular creste is
conspicous.
Piezodorus lituratus (Fabricius 1794) (Plate 5, Fig. 3)
Gonocoxites 8 are covered with setae only on the posterior and internal borders.
Gonocoxites 8, laterotergites 8 and 9 and the tenth segment of the abdomen are covered up
with few setae. The anterior border of the gonapophyses 9 is somehow convex. The diameter of the anterior annular creste is smaller than the diameter of capsula seminalis.
Capsula seminalis has a particular form (Plate 5, Fig. 4).
Eurydema ornata (Linné, 1758) (Plates 1, 2)
Gonocoxites 8, laterotergites 8 and 9 and the tenth segment of the abdomen are
covered up with setae. The setae of gonocoxites 8 are denser on the posterior and internal
borders. The tenth segment of the abdomen is easily concave. The anterior and posterior
annular creste are well developed. The proportion between the length and the maximum
width of the capsula seminalis is 1.6. The apical half of capsula seminalis is distended.
Discussion and Conclusions
The female genitalia of Stagonomus amoenus is considerably distinctive from the other female genitalia of the subfamily Pentatominae. The tenth segment of the abdomen
- 34 -
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
and the laterogites 9 are sharpened on the posterior border and, therefore, the apex of the
abdomen is sharp too. Another distinctive feature is the mergence of the laterotergites, the
gonocoxites and gonapophyses 8.
The distribution of the setae on the gonocoxites and the laterotergites is an
important criterion. For example, the gonocoxites 8 of Carpocoris purpureipennis and
Piezodorus lituratus are covered up with setae only on the internal and posterior sides.
The shape of the anterior edge of the tenth segment is another feature useful for
systematics. The anterior edge can be concave (Pentatoma rufipes, Holcostethus vernalis,
Stagonomus amoenus), convex (Piezodorus lituratus, Eysarcoris aeneus) or almost straight.
Two other important criteria are the form of capsula seminalis and the presence or
absence of expansions at this level. At the species Aelia acuminata, Eysarcoris aeneus, Carpocoris purpureipennis, Dolycoris baccarum, Pentatoma rufipes and Rubiconia
intermedia, capsula seminalis is globose.
A particular form of capsula seminalis is encountered at Piezodorus lituratus.
Capsula seminalis is ovoid at the species Stagonomus amoenus, Holcostethus vernalis and
Eurydema ornata.
The expansions of capsula seminalis are absent at the species Stagonomus
amoenus, Piezodorus lituratus, Eysarcoris aeneus, Carpocoris purpureipennis, Dolycoris
baccarum and Eurydema ornata. Rubiconia intermedia has three spur like expansions.
Aelia acuminata and Pentatoma rufipes have two long expansions. Holcostethus vernalis
has just one expansion of capsula seminalis.
The anterior annular creste can be narrow as in most of the species or broad as in Carpocoris purpureipennis and Dolycoris baccarum.
McDonald (1966) described spermatheca of the species Eysarcoris aeneus as
follows: “Spermathecal dilation constricted proximally forming a large distal chamber and
a smaller more elongate proximal chamber, spermatheca otherwise normal” (Plate 6). In
our research McDonald's findings are confirmed as it can be seen from the drawings (Plate
3, Fig. 1; Plate 6): there is a constriction in spermathecal dilation and capsula seminalis
lacks expansions.
Candan et al. (2010) described the morphology of Dolycoris baccarum’s
spermatheca, using electronic microscopy. In our research we used optical microscopy and
the findings were similar to those presented by Candan et al. (2010).
The main similarities and differences found by us between female external
genitalia are summarized in the following key: 1 Gonocoxites and laterotergites 8 merged; posterior edge of tenth segment and
laterotergites 9 sharp (Plate 3, Fig. 2) Stagonomus amoenus
- Gonocoxites and laterotergites 8 not merged; posterior border of tenth segment and
laterotergites 9 unsharpened 2
2 Capsula seminalis globose 3
- Capsula seminalis of different form 8 3 Capsula seminalis with three spur-like expansions (Plate 4, Fig. 1)
Rubiconia intermedia
- Capsula seminalis with two long expansions or without 4
4 Capsula seminalis without expansions 5
- Capsula seminalis with two expansions 7
- 35 -
Elena Mădălina Morariu
5 Gonocoxites 8 covered with setae only on posterior and internal edges (Plate 3, Fig. 3)
Carpocoris purpureipennis
- Gonocoxites 8 covered with setae on the whole surface 6
6 Anterior annular crest broad (Plate 4, Fig. 3) Dolycoris baccarum
- Anterior annular crest narrow (Plate 3, Fig. 1) Eysarcoris aeneus
7 Gonocoxites 9 strongly concave (Plate 5, Fig. 1) Pentatoma rufipes
- Gonocoxites 9 almost straight (Plate 5, Fig. 2) Aelia acuminata
8 Capsula seminalis of particular form; proportion between length and smallest width 3.2-
3.4 (Plate 5, Figs. 3, 4) Piezodorus lituratus
- Capsula seminalis ovoid; proportion between length and smallest width 1.6-1.8 9
9 Capsula seminalis with long expansion (Plate 4, Fig. 2) Holcostethus vernalis - Capsula seminalis without expansions (Plates 1, 2) Eurydema ornata
Plate 1. Generalized female genitalia in pentatomides – Eurydema ornata (Linné, 1758).
median dilation
capsula seminalis (spermathecal bulb)
pars intermedialis (spermathecal pomp)
posterior annular crest (distal flanges)
ductus receptaculis (distal spermathecal duct)
anterior annular crest (proximal flanges)
sclerotized median dilation
thickening of vaginal intima
proximal spermathecal duct
gonapophysis 8 (valva I, triangulin)
orificium receptaculi
gonocoxite 8 (valvifer I)
gonapophysis 9 (valva II, arcus)
gonocoxite 9 (valvifer II)
paratergite (laterosternite) 9
paratergite (laterosternite) 8
tenth segment
- 36 -
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
Plate 2 (Figs. 1-2). Female genitalia of Eurydema ornata (Linné, 1758) 1. Segment 8; 2. Segments 9
and 10.
- 37 -
Elena Mădălina Morariu
Plate 3 (Figs. 1-3). Female genitalia: 1. Eysarcoris aeneus (Scopoli, 1763); 2. Stagonomus amoenus (Brullé, 1832); 3. Carpocoris purpureipennis (De Geer, 1773).
1 2
3
- 38 -
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
Plate 4 (Figs. 1-3). Female genitalia: 1. Rubiconia intermedia (Wolff, 1811); 2. Holcostethus vernalis (Wolff, 1804); 3. Dolycoris baccarum (Linné, 1758).
- 39 -
Elena Mădălina Morariu
Plate 5 (Figs. 1-4). Female genitalia: 1. Pentatoma rufipes (Linné, 1758); 2. Aelia acuminata (Linné,
1758); 3. Piezodorus lituratus (Fabricius, 1794); 4. P. lituratus, detail of capsula seminalis.
1
2
3 4
- 40 -
Analele Științifice ale Universității „Alexandru Ioan Cuza” din Iași, s. Biologie animală, Tom LVIII, 2012
Plate 6. Spermatheca of Eysarcoris aeneus (after McDonald, 1966).
The analysis of the female genitalia of pentomines shows the existence of
distinctive characters, at least between genera. The most important characteristics that may be used in systematics are: the chaetotaxy, the shape of the ninth gonocoxite’s anterior
edge, the shape of capsula seminalis and the number of its expansions.
Aknowledgements
I would like to thank Dr. Petr Kment for providing references, Dr. Ovidiu Popovici
for assisstance and guidance in my research and Dorina Pascu for her kindness in helping
me translates the paper.
References
Candan, S., Erbey, M., Sümeyye Yilmaz, F., 2010. Surface Morphology of the Spermatheca of Dolycoris
baccarum (Linnaeus, 1758) (Heteroptera: Pentatomidae). Entomological News, 121(4): 334-341.
Crampton, G.C., 1920. Remarks on the basic plan of the terminal abdominal structures of the males of winged
insects. The Canadian EntomologistI, 52(8): 178-183.
Crampton, G.C., 1922. The genitalia of the males of certain hemiptera (Heteroptera) and homoptera. Bulletin of
Brooklyn Entomological Society, XVII: 46-57.
Derjanschi, V., Péricart, J., 2005. Hémiptères Pentatomoidea Euro-méditerrnéens vol. I Généralités Systématique:
Première Partie, Faune de France (France et régions limitrophes). Fédération des Sociétés des
Sciences Naturelles, 494 pp.
Dupuis, C., 1951. Les espèces françaises du genre Eurydema Laporte (Hemiptera, Pentatomoidea, subfam.
Pentatominae). Révision systematique avec une introduction à l’étude morphologique des organes
génitaux externes des Pentatomoidea. Annales de la Société Éntomologique de France, 118: 1-29.
Dupuis, C., 1952. Donné sur la morphogenese des genitalia males des insectes. Leur importance pour une
nomenclature rationelle de ces structures. Transsactions of the Ninth International Congress of
Entomology, 1: 151-154.
- 41 -
Elena Mădălina Morariu
Dupuis, C., 1953. Les génitalia mâles des hémiptères hétéroptères leur utilisation en systématique. Transsactions
of the Ninth International Congress of Entomology, 2: 7-10.
Dupuis, C., 1955. Les genitalia des hemipteres Heteropteres (Genitalia externes des deux sexes; Voies
ectodermiques femalles). Revue de la morphologie. Lexique de la nomenclature. Index bibliographique
analytique. Memoires du Museum d'Histoire Naturelle, 6: 183-278.
Dupuis, C., 1970. Heteroptera, pp. 190-208. In : S.L. Tuxen (Editor). Taxonomist´s glossary of insects.
Copenhagen, Munksgaard, Denmark 359 pp.
Gapon, D., 2001. Inflation of heteropteran aedeagi using microcapillaries (Heteroptera: Pentatomidae).
Zoosystematica Rossica, 9(1): 157-160.
Kis, B., 1984. Heteroptera – partea generală și suprafamilia Pentatomoidea, Fauna Republicii Socialiste
România, vol. VIII, fasc. 8. Editura Academiei Republicii Socialiste România, București, 216 pp.
McDonald, F.J.D., 1966. The genitalia of North American Pentatomoidea (Hemiptera: Heteroptera). Quaestiones
Entomologicae 2: 7-150.
Mihalache, C., Săvescu, A., Buguleanu, Gh., Paulian, Fl., Balaj, D., Pașol, P.,1969. Entomologie agricolă. Editura
Agrosilvică, București, 564 pp.
Millar, I.M., Uys, V.M., Urban, V.P. (eds.), 2000. Collecting and preserving insects and arachnids. R. P., Plant
Protection Research Institute, Pretoria, South Africa.
Panizzi, A.R., McPherson, J. E., James, D.G., Javahery, M., McPherson, R.M., 2000. Stink Bugs (Pentatomidae).
In Schaefer, C.W., Panizzi A.R. (eds.) Heteroptera of economic importance. CRC Press LLC: 421-474.
Rider, D.A., 2006. Family Pentatomidae Leach, 1815. In Aukema, B., Rieger, Ch. (eds) Catalogue of the
Heteroptera of the Palaearctic Region. Volume 5. Pentatomomorpha II. The Netherlands
Entomological Society, Amsterdam: 233-402.
Schaefer, C.W., 1968. The Homologies of the Female Genitalia in the Pentatomoidea (Hemiptera-Heteroptera).
New York Entomol. Society, 76: 87-91.
Schaefer, C.W., 1998. Phylogeny, Systematics, and Practical Entomology: The Heteroptera (Hemiptera). Anais da
Sociedad Entomológica do Brasil, 27(4): 499-511.
Schuh, R.T., Slater, J.A., 1995. True bugs of the world (Hemiptere: Heteroptera). Cornell University Press, 337
pp.
Snodgrass, R.E., 1935. Principles of insect morphology. McGraw-Hill Book Co., Inc., New York & London, 667
pp.
Štys, P., 1961. Morphology of the abdomen and female ectodermal genitalia of the trichophorus Heteroptera and
bearing on their classification. 11th International Congress of Entomomolgy, Wien, 17-25.VIII.1960.
Verhandl., Bd. I, sect. I-IV: 37-43.
Tay, E.B., 1969. Morphology of the external genitalia of some pentatominae (Heteroptera). Papers of Department
of Entomology of University of Queensland, 2: 75-84.
- 42 -