Zoologischer Anzeiger 253 (2014) 236–258
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Zoologischer Anzeiger
journal homepage: www.elsevier.com/locate/jcz
External thoracic scent efferent system of Scutelleridae
(Hemiptera: Heteroptera)
Shama Parveen
a,*
, Kamil Usmani
b
, Sucheta Khokhar
c
,
Vilayanoor Venkataraman Ramamurthy
a
, Jitka Vilímová
d
a
Division of Entomology, Indian Agricultural Research Institute, New Delhi 110012, India
b
Department of Zoology, Aligarh Muslim University, Aligarh 202002, India
c
Department of Entomology, C.C.S.H.U, Hisar 125004, India
d
Faculty of Science, Department of Zoology, Charles University, Vinicna 7, 128 44 Praha 2, Czech Republic
article info
Article history:
Received 22 August 2013
Received in revised form
14 November 2013
Accepted 19 November 2013
Available online 28 December 2013
Corresponding Editor: Sven Bradler.
Keywords:
Scutelleridae
Metathoracic scent glands
Ostiole
Peritreme
Evaporatorium
Microsculptures
abstract
The metathoracic scent glands in the Heteroptera produce defence secretions which are spread out-
side the body through and by using the thoracic external scent efferent system. That complex system
was studied in 18 species from 11 genera of four subfamilies, Elvisurinae, Eurygastrinae, Hoteinae and
Scutellerinae of the family Scutelleridae (Pentatomoidea). The results have been compared with pub-
lished data. The pattern of that system is more consistent at the level of genus, mostly very similar in
the congeneric species, but mostly variable within higher taxonomic levels, tribes and subfamilies. Five
types of the external scent efferent system are recognized within the family Scutelleridae, basic two of
them in studied species: (i) peritreme well developed, covering large part of metapleuron, evaporatorium
small, developed only on metapleuron, (ii) evaporatorium large, more conspicuous than moderate-sized
to small peritreme, extending to mesopleuron as large structure. The results do not support a hypothesis
that the system of structures associated externally with metathoracic scent glands is in correlation with
type of a habitat. However, these structures are well usable as diagnostic characters for scutellerid genera
(e.g. Cantao, Hyperonchus, Scutellera and Solenosthedium).
© 2013 Elsevier GmbH. All rights reserved.
1. Introduction
A specific pattern of scent glands including their unique efferent
systems is considered autapomorphic feature for the Heteroptera
within the Insecta (e.g. Wheeler et al., 1993; Schuh and Slater, 1995;
Weirauch, 2006). These glands are primarily significant in a defence
(Remold, 1962, 1963; Cobben, 1978; Staddon, 1979; Aldrich, 1988).
Dorsoabdominal scent glands are basically developed in larvae
and often persist to adulthood (e.g. summary in Weirauch, 2006 and
Vilímová and Kutalová, 2012), however, metathoracic/metapleural
scent glands (hereinafter just MTGs) are exclusively adult struc-
tures (e.g. Staddon, 1979). The MTGs were first mentioned by
Dufour (1833) as a source of repelent secretion in the heteropteran
adults. Subsequently, Künckel d’Herculais (1866) described other
scent glands in a quite different position, dorsally in the abdomen
in larvae. Brindley (1930) was the first who use the structural
details of the scent apparatus for the Heteroptera classification,
*
Corresponding author. Tel.: +91 011 25842042.
E-mail addresses: shama parven@yahoo.co.in (S. Parveen),
usmanikamil94@gmail.com (K. Usmani), suchetakhokhar@gmail.com (S. Khokhar),
vvrento@gmail.com (V.V. Ramamurthy), vilim@natur.cuni.cz (J. Vilímová).
and also tried to establish a phylogenetic relationships in some
families. Cobben (1978) emphasized the importance of MTGs in a
phylogenetic hypothesis of the Heteroptera, while Schaefer (1972)
suggested that level of development of the external structures asso-
ciated with MTGs is in a correlation with type of habit within
trichophoran Heteroptera.
The MTGs occupy a ventral position in the hind part of the
metathorax, concealed dorsally by the abdominal nerves and gut.
The glands themselves consist of differentiated epithelial cells
forming units function as secretory structures. A secretion of
them is emptying into a reservoir lined with a cuticular intima.
Sometimes, also an accessory gland attached to the reservoir is
developed, which apparently serves to deliver enzymatic prod-
ucts important in a production of the final scent gland substances
(Noirot and Quennedey, 1991). The metathoracic scent apparatus
is divided to internal parts, i.e. proper glands with ductuli, reser-
voir, lateral duct and its valvular apparatus and to thoracic scent
efferent system composed from internal scent efferent system and
external scent efferent system (details and summary of the ter-
minology in Kment and Vilímová, 2010a). The external cuticular
structures associated with the MTGs help in a rapid evaporation
of the secretion along with protection of the individual itself from
its toxicity (Remold, 1962, 1963; Staddon, 1979; Carver, 1990). The
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http://dx.doi.org/10.1016/j.jcz.2013.11.003