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The Pentatomidae, representing the fourth largest family within Heteroptera, are one of the most diverse groups with about 800 genera and more than 4,700 species in the world. In the Neotropics, about 230 genera and 1,400 species are included in seven subfamilies of the world’s nine subfamilies; four subfamilies are exclusively of the Neotropics. In this chapter, for each subfamily, a diagnosis, an overview of the classification, and information on life history, ecology, and economic importance are given. Comprehensive keys and diagnosis to the subfamilies, tribes, and genera for the Neotropical Region, including Mexico, Central and South America, and the West Indies, are also given.
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681© Springer Science+Business Media Dordrecht 2015
A.R. Panizzi, J. Grazia (eds.), True Bugs (Heteroptera) of the Neotropics,
Entomology in Focus 2, DOI 10.1007/978-94-017-9861-7_22
Chapter 22
Stink Bugs (Pentatomidae)
Jocélia Grazia , Antônio R. Panizzi , Caroline Greve ,
Cristiano F. Schwertner , Luiz A. Campos , Thereza de A. Garbelotto ,
and José Antônio Marin Fernandes
Abstract The Pentatomidae, representing the fourth largest family within
Heteroptera, are one of the most diverse groups with about 800 genera and more
than 4,700 species in the world. In the Neotropics, about 230 genera and 1,400 spe-
cies are included in seven subfamilies of the world’s nine subfamilies; four subfami-
lies are exclusively of the Neotropics. In this chapter, for each subfamily, a diagnosis,
an overview of the classifi cation, and information on life history, ecology, and eco-
nomic importance are given. Comprehensive keys and diagnosis to the subfamilies,
tribes, and genera for the Neotropical Region, including Mexico, Central and South
America, and the West Indies, are also given.
J. Grazia (*) C. Greve L. A. Campos
Departamento de Zoologia, Instituto de Biociências , Universidade Federal do Rio Grande do
Sul (UFRGS) , Av. Bento Gonçalves 9500, prédio 43435, Bairro Agronomia ,
Porto Alegre , RS 91501-970 , Brazil
A. R. Panizzi
Laboratório de Entomologia , Embrapa Trigo , Caixa Postal 3081 ,
Passo Fundo , RS 99001-970 , Brazil
C. F. Schwertner
Departamento de Ciências Biológicas, Instituto de Ciências Ambientais,
Químicas e Farmacêuticas , Universidade Federal de São Paulo (UNIFESP) ,
Rua São Nicolau, 210 Centro , Diadema , RS 09913-030 , Brazil
T. de A. Garbelotto
Programa de Pós-Graduação em Biologia Animal, Instituto de Biociências ,
Universidade Federal do Rio Grande do Sul (UFRGS) , Av. Bento Gonçalves 9500,
prédio 43435, Bairro Agronomia , Porto Alegre , RS 91501-970 , Brazil
J. A. M. Fernandes
Instituto de Ciências Biológicas , Universidade Federal do Pará ,
Rua Augusto Corrêa n°1 , Belém , PA 66075-110 , Brazil
22.1 Introduction
Pentatomidae is the fourth most numerous family within Heteroptera including
more than 4,700 species in more than 800 genera (Table
22.1 ). Pentatomids are
represented in all zoogeographical regions although the tropical and subtropical
faunas are more diverse; they are commonly known as stink bugs. Rolston and
McDonald (
1979 ), Rolston ( 1981 , 1984 ), and Rolston et al. ( 1980 ) provided keys to
the identifi cation of subfamilies, tribes, and genera of the Western Hemisphere. The
subfamilies Asopinae, Cyrtocorinae, Discocephalinae, Edessinae, and Pentatominae
are widely represented in the Neotropical Region, while Podopinae and Stirotarsinae
only have one species each. The subfamilies Cyrtocorinae, Discocephalinae, and
Stirotarsinae are exclusively of the Neotropics. The Edessinae, with the exception
of two species found in the Southern United States of America (McPherson
1982 ),
occur in the Neotropics.
The asopines have predacious habits and the incrassate rostrum is the morpho-
logical feature common to the group as an adaptation to predatory behavior. They
are found in all zoogeographical regions.
Table 22.1 Number of genera and species in the subfamilies and tribes of Pentatomidae of the
Neotropical region
World Neotropics
Genera Species Genera Species
Asopinae 63 357 23 100
Cyrtocorinae 4 11 4 11
Discocephalinae 76 315 76 315
Discocephalini 43 192 43 192
Ochlerini 33 123 33 123
Edessinae 7 >300 7 >300
Pentatominae 380 >2,803 116 >667
Carpocorini 106 >800 53 >223
Catacanthini 7 53 5 32
Halyini 82 >390 2 13
Mecideini 1 17 1 4
Menidini 27 >100 2 23
Nezarini 20 150 3 >70
Pentatomini 65 >900 30 >200
Piezodorini 4 >25 1 1
Procleticini 11 33 11 23
Sciocorini 11 >150 1 3
Strachiini 14 >100 1 13
Unplaced 32 92 6 53
Podopinae 62 255 1 1
Stirotarsinae 1 1 1 1
Total 228 >1,400
J. Grazia et al.
Cyrtocorinae is a small subfamily, cryptically colored with the tree bark substra-
tum where they live; the scutellum with a median, long, and tough spine is a mor-
phological feature found in all species. They are distributed from Mexico to
Discocephalinae includes two tribes, Discocephalini and Ochlerini, with more
than 300 species almost all medium sized and dark colored. The morphology of the
immatures and the biology within the group are poorly known. Exceptions are
found in Discocephalini, the genus Antiteuchus Dallas which is a natural host of
cacao ( Theobroma cacao ) in the Amazon region and shows maternal care habits,
and Ochlerini, the genera Lincus Stål and Macropygium Spinola of which some spe-
cies are important vectors of Phytomonas spp. (Trypanosomatidae) in palms in
South America.
Edessinae includes the most colored and largest stink bugs, with great diversity
in the Amazon region, and more than 300 known species. They have in common the
strong development of the metasternum. For more than a century, the group was
treated as a tribe of Pentatominae but studies in the last two decades raised the taxon
to subfamily rank and increased the number of genera to seven.
The subfamily Pentatominae is the most diverse within the Pentatomidae and
includes species recorded in all the zoogeographical regions in numbers higher than
3,000 distributed in about 600 genera. About 1/4 of this diversity is present in the
Neotropics. They are all phytophagous and many species are economically impor-
tant as pests of cultivated crops. Pentatominae is not a monophyletic group and does
not have a defi nition based on phylogenetic studies. Different authors consider the
classifi cation within Pentatominae as having from 8 to more than 40 tribes.
Following Rider’s tentative classifi cation, 11 tribes are registered in the Neotropics:
Catacanthini Atkinson, Carpocorini Mulsant & Rey, Halyini Amyot & Serville,
Mecideini Distant, Menidini Atkinson, Nezarini Atkinson, Pentatomini Leach,
Piezodorini Atkinson, Procleticini Pennington, Sciocorini Amyot & Serville, and
Strachiini Mulsant & Rey.
The subfamily Podopinae has worldwide distribution but is poorly represented in
the Neotropics, with only one species. The subfamily Stirotarsinae was recently
erected to include Stirotarsus abnormis Bergroth, only known from the northern
region of South America.
22.2 Subfamily Asopinae
22.2.1 Introduction
Species of the subfamily Asopinae are predaceous, a secondary condition among
pentatomids. More than 350 species and 60 genera are known worldwide (Thomas
1992 , 1994 ). Asopines are easily recognized by the incrassate rostrum (Fig. 22.1 ),
and some species are useful as biocontrol agents (De Clercq
2000 , 2008 ).
22 Stink Bugs (Pentatomidae)
22.2.2 General Characteristics and Diagnosis
The general morphology of the species is similar to other groups of pentatomids.
Diagnostic characteristics include modifi cations of the head associated to the feed-
ing habits: rostrum strongly incrassate, insertion of the labium very close to the base
of labrum, and posterior margins of buccula merged (Gross
1975 ; Gapud 1991 ). The
group also shared the presence of the genital plates of the male pygophore and pres-
ence of thecal shield (Gapud
1991 ; Gapon and Konstantinov 2006 ).
22.2.3 General Biology and Ecology
Asopines predate insects and other small organisms, usually slow and soft-bodied
individuals, i.e., larvae of Lepidoptera, Diptera, and Hymenoptera (Schuh and
1995 ; De Clercq 2008). Most species seems to be generalists, but some may
have a more restricted diet (i.e., oligophagous) or may be related to specifi c habitats
(De Clercq
2000 , 2008). Nymphs and adults are observed sucking plant fl uids and
water, probably to acquire the basic needs during periods of prey scarcity. However,
asopines do not develop unless they have access to nutrients of animal origin,
required from the 2nd instar on. Early nymphs (2nd and 3rd instars) tend to attack
and feed together, while the late instars and adults tend to attack alone; nymphs and
adults may show cannibalism when preys are not available (De Clercq
2000 , 2008).
Life cycle of the asopines is similar to that of the phytophagous pentatomids.
Adults emerge and start to feed and reproduce using different preys; some species
may live up to 3 months. They use visual, chemical, and tactile cues to locate and
recognize their prey. Mating behavior is similar to phytophagous species, with
males starting courtship by antennal movement. Eggs are laid in masses, on differ-
ent parts of plants used by adults or on nearby structures. The number of eggs per
clutch varies between species, between individuals, or between clutches of the same
female. There are fi ve nymphal instars. First instars are not predatory and only need
Fig. 22.1 Apateticus
lineolatus (Herrich-
Schaeffer), lateral showing
detail of the incrassate
rostrum (Photo by CF
J. Grazia et al.
moisture for survival. From the 2nd instar on, they need to feed on prey for survival.
Initial nymphs are gregarious and tend to attack in groups; gregarious behavior
decreases with the development, and nymphs disperse in search for prey. Incubation
period, nymphal development time, prereproductive period, and longevity are quite
variable; infl uence of food and abiotic conditions (temperature and humidity)
impact the expression of these traits. Egg parasitoids (Hymenoptera: Scelionidae)
are recorded as natural enemies.
Although highlighted as potential agents for pest control, the use of asopines in
biological control programs is still limited. Only about 10 % of the species have
been the focus of such studies (De Clercq
2008 ). In the Neotropical Region, species
of the genera Alcaeorrhynchus Bergroth, Apateticus Dallas, Euthyrhynchus Dallas,
Brontocoris Thomas, Perillus Stål, Podisus Herrich-Schäffer, Stiretrus Laporte,
Supputius Distant, and Tylospilus Stål have been the focus of recent studies. Two
species, Podisus maculiventris (Say) and Podisus nigrispinus (Dallas), show great
potential in controlling populations of phytophagous insects (Saini
1994 ; De Clercq
2000 ). P. maculiventris is the only asopine commercially available for augmentative
biological control (De Clerq
2008 ).
22.2.4 Classifi cation and Diversity
In the Neotropical Region, 23 genera and 100 species are known (Table 22.2 ).
Taxonomy of the subfamily Asopinae was revised by Thomas (
1992 ), which
includes identifi cation keys to all genera and species of the Western Hemisphere.
Table 22.2 Checklist of genera and species of Neotropical Asopinae (distribution as available in
the literature)
Species Distribution
Alcaeorrhynchus grandis (Dallas) USA, MEX, TRI, CO, BR, UR, ARG, ECU,
Alcaeorrhynchus phymatophorus (P. de B.) PR, HT, DOR, CU, USA (Florida Keys)
Andrallus spinidens (F.) USA, MEX, NIC, CR, CU, HON, DOR
Apateticus lineolatus (Herrich-Schaeffer) USA, MEX, CR, ESAL, PAN, HON, CO, VEN,
Apateticus marginiventris (Stål) USA, MEX
Apoecilus invarius (Walker) USA, MEX
Brontocoris nigrolimbatus (Spinola) CHI
Brontocoris tabidus (Signoret) CHI, PAR, ARG, BR
Colpothyreus fl avolineatus (Blanchard) BOL
Comperocoris roehneri (Philippi) CHI, ARG
Coryzorhaphis leucocephala Spinola BR
Coryzorhaphis carneolus Erichson GUY, BR, PER, BOL, ECU, FG
Coryzorhaphis cruciata Stål MEX, BEL, HON, PAN, CO, VEN, BR
22 Stink Bugs (Pentatomidae)
Table 22.2 (continued)
Species Distribution
Coryzorhaphis superba Breddin PER, BOL, ECU
Coryzorhaphis egeri Thomas MEX, CR, HON
Coryzorhaphis dollingi Thomas ECU
Discocera caynnensis Laporte BR, PER, SUR, FG
Discocera coccínea (F.) BR, PER, COL, ECU, SUR, PAR, BOL
Euthyrhynchus fl oridanus (L.) USA, MEX, GUA, ESAL, HON, CR, PAN, ECU,
Heteroscelis servillei Laporte BR, PER, BOL, ARG, PAR, VEN, GUY, FG,
Heteroscelis lepida (Stål) USA, MEX, GUA, PAN, CO, VEN, FG, ECU,
Heteroscelis bimaculata (Walker) BOL, BR, PER, PAN
Heteroscelis robustus Thomas BR, PAR
Marmessulus nigricornis Bergroth BR, ARG, URU
Marmessulus brasilianus Schouteden BR
Oplomus dichrous (Burmeister) USA, MEX, HON
Oplomus catena (Drury) BR, FG, ARG, URU, PAR, ECU
Oplomus cruentus (Burmeister) ARG, BR, PER, URU
Oplomus mutabilis Stål MEX, GUA, HON
Oplomus ebulinus (Herrich-Schaeffer) BR, PER, BOL, PAR, ECU
Oplomus salamandra (Burmeister) COL, VEN, SUR, ECU, BOL, PER, BR
Oplomus marginalis (Westwood) BR, PAN, COL, ECU, PER, BOL, ARG, VEN,
Oplomus mundus Stål USA, MEX, GUA, CR, PAN, NIC, HON
Oplomus pulcher Dallas MEX, CR, PAN, HON
Oplomus annotatus Uhler CU
Oplomus punctatus Montandon ARG
Oplomus pulchiventris Horvath ARG
Ornithossoma rivieri Kormilev ARG
Parajalla sanguineosignata (Spinola) CHI
Perillus confl uens (Herrich-Schaeffer) USA, MEX, GUA, ELS, CR, HON
Perillus circumcintus Stål USA, CAN, MEX
Perillus bioculatus (F.) USA, CAN, MEX
Perillus splendidus (Uhler) USA, MEX
Podisus aenescens (Stål) MEX, GUA, HON, CR, PAN, CO, ECU, BOL,
Podisus sculptus Distant CR, PAN, PER, BOL
Podisus formosus Costa Leite BR
Podisus mucronatus Uhler USA (Florida), CU, PUR, DOR
Podisus falcatus Distant GUA, HON, CR
Podisus semialbus (Walker) BR, VEN
Podisus curvispina Bergroth BR
J. Grazia et al.
Table 22.2 (continued)
Species Distribution
Podisus cornutus (Dallas) CO, BR
Podisus gundlachii (Guérin-Ménéville) CU
Podisus borinquensis Barber PUR
Podisus congrex (Stål) MEX, GUA, CR, ESAL, PAN, VEN, ECU, HON
Podisus crassimargo (Stål) BR, ECU, COL, VEN, CHI, PAN
Podisus volxemi Distant BR, ECU
Podisus tinctus (Dallas) COL, ECU
Podisus subferrugineus Barber & Bruner CU, JAM
Podisus sagitta (F.) USA (Texas), MEX, GUA, ESAL, HON, NIC,
PUR, TRI, Curacao, CU, BAH
Podisus nigrispinus (Dallas) BR, PAN, CR, BOL, ARG, PER, ECU, SUR,
Podisus distinctus (Stål) ECU, BOL, COL, BR, VEN, FG, PAR, ARG,
Podisus maculiventris (Say) USA, MEX, CAN, HAI, DRE, BAH
Podisus neglectus Westwood USA, MEX
Podisus affi nis Distant MEX, GUA, HON, PAN
Podisus mexicanus Distant MEX
Podisus nigriventris Distant MEX, GUA, NIC, HON, CR, PAN, COL
Podisus insignis Distant GUA
Podisus trucidatus Thomas MEX, HON, CR, PAN
Podisus ventralis (Dallas) VEN, BR, ECU, PER, ARG
Podisus sordidus (Stål) ECU (Galapagos)
Podisus rostralis (Stål) BR, BOL
Podisus pallipes (Dallas) VEN, BR, ECU, ARG
Podisus mactans Thomas ECU
Podisus serieventris Uhler CAN, USA, MEX
Stiretrus quinquepunctatus Germar HAI, DOR
Stiretrus decemguttatus (Lepeletier &
Stiretrus decastigmus (Herrich-Schaeffer) ARG, BRA, BOL, PAR
Stiretrus erythrocephalus (Lep. & Serville) ARG, BRA, BOL, PAR, URU
Stiretrus anchorago (F.) USA, MEX, GUA, ESAL, CR, HON, PAN, NIC
Stiretrus bifrenatus Breddin COL, ECU, PER, BRA, URU
Stiretrus loratus Germar BR, BOL
Stiretrus cinctellus Germar BR, ARG
Supputius typicus Distant MEX, CR, PAN, VEM, COL, ECU, BR, HON
Supputius cincticeps (Stål) BR, VEN, PR, CHI, BOL, ARG
Supputius pulchricornis (Stål) MEX
Tylospilus armatus Thomas ARG
Tylospilus cloelia (Stål) MEX, HON, PAN, COL, BOL, BR, PAR, ARG
Tylospilus chilensis (Spinola) CHI, ARG
22 Stink Bugs (Pentatomidae)
Since then, studies about morphology, biology, and distribution records of some of
the Neotropical species have been published (see Sects.
2.5 and 2.6 ).
Key to the genera of subfamily Asopinae with species included in biological control
studies in the Neotropical Region (adapted from Thomas
1992 ). Number in super-
script indicates Neotropical genera that are keyed in the same step (see list at the end
of the key).
1. Scutellum greatly enlarged, covering most of the abdomen, apex surpassing coria
in length, almost attaining apex of abdomen (Fig.
22.2 ) ......... Stiretrus Laporte 1,2
1 . Scutellum not covering most of the abdomen, usually shorter than coria
22.3 and 22.4 ) ........................................................................................... 2
2. Apical part of the scutellum enlarged, equal in length or longer than the
frenal margin of the scutellum, and about as wide as the corium
22.3 and 22.5 ) ........................................................................................... 3 3
2 . Apical part of the scutellum small, shorter than the frenal margin and much
narrower than the corium (Fig.
22.4 ) .................................................................. 4
3. Frenal margin of the scutellum shorter than the postfrenal scutellum; protibia
usually expanded ....................................................... Oplomus Spinola (Fig.
22.5 )
3 . Frenal margin of the scutellum longer than postfrenal scutellum; protibia always
prismatic, never expanded................................................ Perillus Stål (Fig.
22.3 )
4. Scent gland with elevated peritreme, surrounded by evaporatoria (Fig.
22.6 ) .... 5
4 . Ostiole of scent gland without elevated peritreme, terminating in
short or long sulcus not surrounded by evaporatoria (Fig.
22.7 )
.................................................................Alcaeorrhynchus Bergroth 4 (Fig. 22.8 )
5. Protibiae dilated, expanded ................................................Euthyrhynchus Dallas 5
5 . Protibiae not dilated ............................................................................................ 6
Species Distribution
Tylospilus nigrobinotatus (Berg) ARG, BRA, URU
Tylospilus acutissimus Stål USA, MEX, NIC, COL, ANT, DOR
Tylospilus megaspilus (Walker) BR
Tylospilus distans Bergroth BR, CHI
Tylospilus peruvianus (Horvath) PERU, SUR, BR
Tynacantha marginata Dallas BR, ARG, BOL, PER, COL, ECU, PAR, URU
Tynacantha splendes Distant PAN, CR
Tyrannocoris rex Thomas BR, VEN
Tyrannocoris nigriceps Thomas BR, ARG
Tyrannocoris rideri Thomas PUR
Tyrannocoris jole (Stål) MEX, HON, CU, HAI, RDO
Zicrona americana Thomas USA, MEX
Table 22.2 (continued)
J. Grazia et al.
6. Base of the abdomen with anteriorly directed tubercle or spinous process
22.9 ) ............................................................................................................. 7
6 . Base of the abdomen may be prominent but without forwardly directed tubercle
or spinous process ..................................Supputius Distant (in part)
6 (Fig. 22.10 )
7. Rostral segment II distinctly longer than III and IV combined
.....................................................................................Supputius Distant (in part)
7 . Rostral segment II shorter or subequal in length to III and IV combined ......... 8
8. Abdominal spine long, extending to mesocoxae; posterior tibiae terete; most or
all of the dorsal surface of the head with pale colorless punctuation .................. 9
8 . Abdominal spine usually short, not surpassing metacoxae; posterior tibia usually
sulcate; dorsal surface of the head densely punctuate ...................................... 10
9. Scent gland peritreme long, reaching more than half way to pleural edge;
anterolateral pronotal margin dentate to spinulose ............... Brontocoris Thomas
9 . Scent gland peritreme short, reaching less than halfway to the pleural
edge; anterolateral pronotal margin rugulose, smooth (Fig.
22.11 )
...................................................................................Tylospilus Stål (Fig.
22.12 )
10. Inner margins of jugae distinctly convergent (Fig.
22.13 ); males with glandular
patches of silk hairs on abdominal venter ...........Apateticus Dallas 9 (Fig. 22.14 )
10. Inner margins of jugae parallel or concave but never convergent (Fig.
22.15 );
males without glandular patches of silk hairs on abdominal venter
.......................................Podisus Herrich-Schäffer (Figs.
22.4 , 22.16 , and 22.17 ).
1, Colpothyreus Stål; 2, Discocera Laporte; 3, Heteroscelis Laporte,
Coryzorhaphis Spinola, Comperocoris Stål, Marmessulus Bergroth; 4, Parajalla
Distant, Rhacognathus Fieber, Zicrona Amyot & Serville; 5, Eocanthecona
Bergroth; 6, Ornithosoma Kormilev, Andrallus Bergroth; 7, Picromerus Amyot &
Serville; 8, Thynacantha Dallas, Tyrannocoris Thomas; 9, Apoecilus Stål
22.2.5 Main Species
De Clercq ( 2000 ) reviewed the most studied species of asopines (main and
secondary) in a world basis and discussed information available for each one. In this
and the next section, we present a brief characterization for some of the common
Neotropical species that have been the focus of recent studies.
22 Stink Bugs (Pentatomidae)
Fig. 22.2 Stiretrus
anchorago (F.), dorsal view
(Courtesy of G. Fauske)
Fig. 22.3 Perillus bioculatus
(F.), dorsal view (Courtesy of
G. Fauske)
J. Grazia et al.
Brontocoris tabidus (Signoret)
The genus Brontocoris included only two species, and B. tabidus differs from
B. nigrolimbatus (Spinola) by the anterolateral pronotal margin crenulate and
connexivum with a black spot in each anterolateral and posterolateral angles
1992 ; Rider 2014 ); in B. nigrolimbatus the pronotal margin is spinulose
and the connexivum uniformly pale yellow. Considering a promising biological
agent in Brazil (Zanuncio et al.
1994 ; De Clercq 2000 ), it has been the focus of
several studies (e.g., Oliveira et al.
2005 ; Zanuncio et al. 2006 ; Lemos et al. 2009 ,
2010 ; DeMenezes et al. 2013 ; Pires et al. 2011 ).
Perillus bioculatus (F.) (Fig.
22.3 )
Among the species included in the genus, P. bioculatus can be recognized by the fol-
lowing suite of characters: presence of an ante-apical spine or tubercle in the profe-
mur, pronotum tumid and without lunate black mark on the anterior half, abdominal
spiracles enclosed within black area, and dorsal punctuation widely spaced (Thomas
1992 ). The two-spotted stink bug is one of the main predators of the Colorado potato
beetle ( Leptinotarsa decemlineata , Chrysomelidae); the biology and other aspects of
Fig. 22.4 Podisus
maculiventris (Say), dorsal
view (Photo by CF
22 Stink Bugs (Pentatomidae)
Fig. 22.6 Supputius
cincticeps (Stål), ventral
view showing detail of the
evaporatorium (Photo by CF
Fig. 22.7 Alcaeorrhynchus
grandis (Dallas), ventral view
showing detail of the
evaporatorium (Photo by CF
Fig. 22.5 ( a ) Oplomus catena (Drury), dorsal view (Photo by CF Schwertner). ( b ) Oplomus
dichrous (Burmeister), dorsal view (Courtesy of G. Fauske)
J. Grazia et al.
Fig. 22.8 Alcaeorrhynchus
grandis (Dallas), dorsal view
(Photo by CF Schwertner)
Fig. 22.9 Apateticus
lineolatus (Herrich-
Schaeffer), ventral view
(Photo by CF Schwertner)
22 Stink Bugs (Pentatomidae)
this asopine have been investigated by several authors (reviewed in De Clercq 2000 ;
see also Adams
2000 , 2001 ; Wittmeyer and Coudron 2001 ; Coudron et al. 2002 ;
Coudron and Kim
2004 ; Matlock 2005 ; Greenstone et al. 2010 ).
Podisus maculiventris (Say) (Figs.
22.4 and 22.15 )
Thomas ( 1992 ) recognized several groups of species within Podisus , including the
maculiventris group of species. Species included in this group can be recognized by
the dorsal surface of pronotum without calli ; anterolateral margins of the pronotum
Fig. 22.10 Supputius
cincticeps (Stål), dorsal view
(Photo by CF Schwertner)
Fig. 22.11 Tylospilus distans
Bergroth, head in dorsal view
(Photo by CF Schwertner)
J. Grazia et al.
Fig. 22.12 ( a ) Tylospilus distans Bergroth, dorsal view (Photo by CF Schwertner). ( b ) Tylospilus
acutissimus Stål, dorsal view (Courtesy of G. Fauske)
not infl ated and dentate, crenulate, or rugulose; and humeral angles simple and
directed laterad, not elongately produced (Thomas
1992 ). Within the group, P. mac-
uliventris is the only species with humeral angles laterally produced in acuminate
angles (Fig.
22.4 ) and with the abdominal spine reaching between metacoxae. It is
the most common species in North America, also known as spined soldier bug, and
extensive literature about its biology, role, and potential use in biological control
programs has been published (De Clercq
2000 , 2008 ).
Podisus nigrispinus (Dallas) (Fig.
22.16 )
Species included in the sagitta group, which is similar to the maculiventris group
(see above diagnosis of P. maculiventris ), however, have the humeral angles dis-
tinctly bifi d or emarginate (Fig.
22.16 ) (Thomas 1992 ). Within the group, P. nigris-
pinus can be recognized by the last rostral segment darker than the others and by the
anterolateral margins of pronotum paler than the pronotal disk. One of the most
common species in the Neotropical Region, P. nigrispinus has been found attacking
several pest insects (De Clercq
2000 ). Several studies has been carried out regarding
22 Stink Bugs (Pentatomidae)
Fig. 22.13 Apateticus
lineolatus (Herrich-
Schaeffer), head in dorsal
view (Photo by CF
Fig. 22.14 Apateticus
lineolatus (Herrich-
Schaeffer), dorsal view
(Photo by CF Schwertner)
J. Grazia et al.
the biology, predation potential in different agroecosystems, and rearing methods of
this species, also known as the Brazilian soldier spined bug (De Clercq
2000 ; Vivan
et al.
2003 ; Lemos et al. 2003 , De Medeiros et al. 2004 ; Sousa-Souto et al. 2006 ;
Torres et al.
2006 ; Neves et al. 2009 ; Rodrigues et al. 2009 ; De Bortoli et al. 2011 ;
Soares et al.
2011 ; Laumann et al. 2013 ; Vacari et al. 2013 ).
Fig. 22.15 Podisus
maculiventris (Say), head in
dorsal view (Photo by CF
Fig. 22.16 Podisus
nigrispinus (Dallas), dorsal
view (Photo by CF
22 Stink Bugs (Pentatomidae)
Supputius cincticeps (Stål) (Figs. 22.6 and 22.10 )
The genus Supputius include three species, and S. cincticeps can be recognized by
rostral segment II as long as III and IV combined, apex of scutellum concolorous
with the disk and abdominal tubercle strongly produced, reaching between meta-
coxae (Thomas
1992 ). Although not so common as P. nigrispinus , this species is
also found attacking several pests in South America (De Clercq
2000 ; Zanuncio
et al.
2004 , 2005a , b ; De Castro et al. 2013 ).
22.2.6 Secondary Species
The species included in this section are far less studied than the species included in
the previous section, and little is known about them besides their distribution and
morphological aspects. However, they represent common Neotropical asopines and
potential control agents in biological control programs.
Alcaeorrhynchus grandis (Dallas) (Figs.
22.7 and 22.8 )
This is the most common and widespread species of Alcaeorrhynchus , the only one
found in the continental Americas (Thomas
1992 ). The genus include only two spe-
cies, which can be identifi ed by the direction of the spined humeral angle: it is bent
Fig. 22.17 Podisus sagitta
(F.), dorsal view (Photo by
CF Schwertner)
J. Grazia et al.
somewhat forward in A. grandis and directed laterad in A. phymatophorus . The
species A. grandis is the largest Neotropical asopine and is associated to several
cultivated ecosystems, attacking lepidopterous larvae (Malaguido and Panizzi
1998a ; De Clercq 2000 ).
Apateticus lineolatus (Herrich-Schaeffer) (Figs.
22.1 , 22.13 , and 22.14 )
The genus Apateticus include two species, and A. lineolatus can be recognized by
the posterior angle of the pronotum without a tooth and the basal abdominal spine
robust, protruding between metacoxae (Thomas
1992 ). There are few studies about
this species (De Clercq
2000 ); in Mexico, A. lineolatus was found feeding in larvae
of the beetles Epilachna varivestis (Coccinellidae) and Leptinotarsa spp.
(Chrysomelidae), pests of important crops as beans and potatoes.
Euthyrhynchus fl oridanus (L.)
This genus is monotypic, despite the high color polymorphism found in E. fl orida-
nus ; Thomas ( 1992 ) mentioned that North and South American populations repre-
sent distinct lineages (forms) of this species. A very conspicuous species, with red
and metallic blue coloration, with extensive literature on bionomics under labora-
tory conditions is available (De Clercq
2000 ; Briceño 2014 ). However, only a few
notes about its life history on the fi eld are known ( Avila-Núñez et al.
2009 ).
Oplomus dichrous (Burmeister) (Fig.
22.5b )
The genus Oplomus includes 12 species, and O. dichrous can be recognized by two
unique characters: prosternum expanded between procoxae and juga longer than
tylus, contiguous and defl exed anteriorly in males (Thomas
1992 ). A common spe-
cies within it distribution range, it seems to be an important predator of L. decem-
lineata Mexico (De Clercq
2000 ), but laboratory and fi eld tests showed that O.
dichrous has little potential as control agent in cool climates.
Podisus spp.
Podisus sagitta (F.) (Fig.
22.17 ) and P. distinctus (Stål)
Both species belong to the sagitta group (Thomas
1992 ), together with P. nigrispi-
nus (see diagnose for P. nigrispinus above). They can be separated by the coloration
of the rostral segments: the last rostral segment darker than the three preceding seg-
ments in P. sagitta and the fi rst rostral segment distinctly paler than the three suc-
ceeding segments in P. distinctus . Besides, both species have the anterolateral
margin of pronotum concolor, which allows separating these species from P.
22 Stink Bugs (Pentatomidae)
nigrispinus . The biology of Podisus spp. is similar, and P. sagitta and P. distinctus
have been studied as control agents of different pests in North and South America
(De Clercq
2000 ; Matos Neto et al. 2004 ).
Podisus rostralis (Stål)
This species is included in the maculiventris group, together with P. maculiventris
(see diagnosis above) and another eight species of Podisus (Thomas
1992 ). The
group is widely distributed in the American continent; however P. rostralis has a
more restricted distribution (Table
22.2 ). It can be recognized among other species
of Podisus by a unique set of characters: dorsum of the head densely punctuated,
hemelytral membrane transparent with a mesial dark blotch, and basal rostral seg-
ment contrastingly pale to three dark succeeding segments. In Brazil, P. rostralis
was found attacking pests of Eucalyptus spp. and information about its biology
under laboratory conditions are available (Molina-Rugama et al.
1998 ; Matos Neto
et al.
1999 ; Lemos et al. 2005 ).
Stiretrus spp.
The genus includes eight species; S. anchorago (F.) (Fig.
22.2 ) and S. decemgutta-
tus (Lepeletier & Serville) can be recognized by the protibia distinctly dilated,
anterolateral margins of the pronotum concavely sinuate to subrectilineal (never
convex), and basal abdominal spine long, protruding between mesocoxae (Thomas
1992 ). The species can be separated by the length of the ostiole: S. anchorago has
the ostiole long, reaching at least halfway to the pleural margin; in S. decemguttatus
it is short, reaching one-third the distance to the pleural margin.
The distribution of both species does not overlap, and S. anchorago is the only
species within the genus that occurs in the north of Panama (all the other species of
Stiretrus are known from South America). More data on the bionomics and life
history of S. anchorago are available (De Clercq
2000 ), but its effectiveness in
agroecosystems still needs more attention. More recently, Paleari (
2013 ) studied
the development, the polymorphism, and some ecological aspects of
S. decemguttatus .
Tylospilus acutissimus (Stål) (Fig.
22.12b )
This species can be recognized within the genus Tylospilus by humeral angle spi-
nose, often slightly directed forward and without a post-apical tooth on the posterior
edge; base of the scutellum with only indistinct calluses at basal angles, never elon-
gated; ostiole extending halfway to the pleural margin; and basal abdominal spine
reaching almost to the mesocoxae (Thomas
1992 ). Although considered a common
J. Grazia et al.
species in some parts of the United States, this species has received minor attention
and only scattered data about its biology is known (De Clercq
2000 ).
22.2.7 Remarks
The subfamily Asopinae includes a diverse group of species, with worldwide distri-
bution and relatively well-known taxonomy (Thomas
1992 , 1994 ). However, studies
in a phylogenetic context are still lacking for the group. The monophyly of the sub-
family and included genera was never questioned, but its position within Pentatomidae
and the phylogenetic relationships within the subfamily are unresolved.
The unique feeding habits within the superfamily Pentatomoidea make the group
an exciting subject of study in many distinct research areas such as evolutionary
biology, comparative physiology, and crop protection.
Knowledge about the Neotropical asopines is mostly restricted to species related
to agroecosystems. Details about species distribution, life history traits, comparative
morphology, and molecular data are still scarce and will be crucial to understand the
diversity and evolution of the group in the Neotropical Region.
22.3 Subfamily Cyrtocorinae
22.3.1 Introduction
This exclusively Neotropical taxon was revised by Packauskas and Schaefer ( 1998 );
it includes four genera and 11 species. Besides the record of Cyrtocoris trigonus
(Germar) from California (Banks
1910 ; Horváth 1916 ; Brailovsky et al. 1988 ),
Packauskas and Schaefer (
1998 ) agreed with Henry and Froeschner ( 1988 ) that the
lack of any subsequent discovery of Cyrtocoris White in the United States makes
Bank’s record suspect. According to Packauskas and Schaefer (
1998 ), Kormilev
1955 ) appears to have been the fi rst author to present evidence for raising Cyrtocorinae
to family rank as distinct from the Pentatomidae allying Cyrtocoridae with Cydnidae.
22.3.2 General Characteristics and Diagnosis
Length 6–10 mm. Body black to brown in ground color, often covered with brown-
tan and/or white scalelike setae, giving the bug a lighter brown color dorsally and
ventrally. Ventral covering usually variegated with areas of deep brown and yellow-
white. Body broadly rectangular in dorsal view, in lateral view convex above, and
22 Stink Bugs (Pentatomidae)
nearly fl at below; trapezoidal to rectangular in posterior view. Head subquadrate,
angles rounded in front of the eyes; mandibular plates developed, fl attened, and
depressed on either side of clypeus. Apex of clypeus at base of rostrum expanded
laterally beyond bucculae. Bucculae extending past eyes. Antennae arising from
below eyes; V segment longest, slightly swollen distally; II shortest (very short).
Rostrum and antennae fi tting into midventral sulcus on thoracic sterna. Pronotum
broadly trapezoidal; humeri greatly expanded and directed downward anterolaterally.
Scutellum broadly U-shaped, extending to end of the abdomen; most of corium and
sometimes part of membrane exposed. Tarsi 2-segmented; 2nd segment 2–4 times as
long as 1st. Metathoracic scent gland opening sustained by an auricle peritreme.
Abdominal segments II–IV exposed dorsally, strongly extended laterally and angled
ventrally. Each abdominal spiracle closer to preceding segment than to lateral edge of
its own segment or to posterior segment; 2nd abdominal spiracle on membranous
anterior of segment. Abdominal sterna III–VII laterally with one pair of trichobothria
side by side, those on anterior sterna lateral to spiracle; trichobothria all of equal size.
22.3.3 General Biology and Ecology
Cyrtocorines are of minor economic importance and they are not commonly found.
As a consequence little is known about their biology. C. trigonus were found feed-
ing on stems of the weed plant arrowleaf sida, Sida rhombifolia L. (Malvaceae), in
Londrina, Paraná State, Brazil. Despite egg deposition in the laboratory, nymphs did
not develop on this plant. Nymphs and adults of Cyrtocoris egeris Packauskas &
Schaefer were observed feeding on immature soybean pods in Argentina
(Reconquista, Santa Fé Province). Under laboratory conditions, nymphs of neither
species could be reared beyond the 3rd instar (Schaefer et al.
2005 ). Adult and
nymphal C. egeris fed gregariously on the stems of Acalypha diversifolia Jacquin
(Euphorbiaceae) and moved to the leaves when these became available; adults could
also feed on Piper sp. (Piperaceae) (Brailovsky et al.
1988 ). Cyrtocoris gibbus (F.)
is reported to feed on stems of other legume plants such as the trees Acacia sp. and
“bracatinga” ( Mimosa scabrella Bentham), in Rio de Janeiro (Costa Lima
1940 ;
Silva et al.
1968 ). Adults of C. trigonus feed on Syngonium podophyllum Schott
(Araceae), Acalypha sp. (Euphorbiaceae), Piper sp., and Pothomorpha umbellatum
(L.) Miquel (Piperaceae) (Brailovsky et al.
1988 , as C. obtusus Horváth).
Not many studies have been published focusing on Cyrtocorinae immatures.
Brailovsky et al. (
1988 ) described the egg and nymphs of C. egeris [as Cyrtocoris
trigonus (Germar)] from Mexico, including notes on the biology of the species.
Schaefer et al. (
1998 ) described the last three instars of C. egeris from Ecuador.
Bianchi et al. (
2011 ) studied under SEM the egg and the 1st and 5th instars of
C. egeris ; their results corroborate the hypothesis of Schaefer et al. (
1998 ) on the
absence of spiracles on urosternite I of nymphs and the trichobothrial arrangement
(1 + 1 trichobothria on 1st instar and 2 + 2 trichobothria on following instars) contrary
to Brailovsky et al. (
1988 ). Also, the placement of the trichobothria relative to the
J. Grazia et al.
spiracle on different sterna, becoming more medial posteriorly, described for the
later instars and adult of C. egeris in Schaefer et al. (
1998 ) were confi rmed by Bianchi
et al. (
2011 ). The presence of three irregular rows of aero-micropylar processes in the
egg stage and the large amount of dorsal abdominal sclerotization in the nymphal
stage, as stated by Schaefer et al. (
1998 ), seem to be distinctive features of
Cyrtocorinae, comparable to the degree of modifi cation of the adult stage. On the
other hand, barrel-shaped eggs and the pattern of the nymphal morphology are shared
by Cyrtocorinae and other subfamilies of the Pentatomidae (Bianchi et al.
2011 ).
22.3.4 Classifi cation and Diversity
Cyrtocorinae includes four genera, three of them monotypic and one with eight spe-
cies: Ceratozigum Horváth, monotypic with C. horridum (Germar) (Fig.
22.18a, b )
registered for Brazil, Panama, Peru, and Venezuela; Cyphothyrea Horváth, mono-
typic, with C. erosa Horváth, from Peru and Brazil; Cyrtocoris White, the most
diverse genus, including eight species – C. andicola Horváth (Peru), C. egeris
Packauskas & Schaefer (Mexico through Brazil), C. gibbus (F.) (widely distrib-
uted), C. montanus Horváth (Peru), C. obtusus Horváth (Brazil), C. paraensis Pirán
(Southern Brazil), C. simplex Horváth (Brazil), and C. trigonus (Germar) (Mexico
through Argentina) – and Pseudocyrtocoris Jensen-Haarup, monotypic with P. lac-
eratus (Herrich-Schäffer) from Argentina and Brazil.
Fig. 22.18 Ceratozigum horridum (Germar): ( a ) female; ( b ) male (Photos by A Ferrari)
22 Stink Bugs (Pentatomidae)
Packauskas and Schaefer ( 1998 ) provided detailed redescriptions of the taxa and
keys to separate the genera and the Cyrtocoris species, beside the intraspecifi c vari-
ability found in this genus; they also discussed the peculiar sexual dimorphism
found in Ceratozigum horridum (Germar). There are two forms of the head in the
males themselves, one form resembling the female mandibular plates (greatly
produced anteriorly) and another form (but in no female) where the mandibular
plates extend far forward.
Kormilev ( 1955 ) appears to have been the fi rst author to present evidence for
raising Cyrtocorinae to family rank as distinct from the Pentatomidae allying
Cyrtocoridae with Cydnidae. Kormilev's arguments persuaded Rolston and
McDonald (
1979 ) to raise Cyrtocorinae, and for the most part Cyrtocoridae has
been recognized since. Gapud (
1991 ) separated Cyrtocorinae (as a pentatomid
subfamily) from the rest of Pentatomidae by the absence of a triangulin, 2nd
valvifers with a distinct median fusion line, and male phallotheca relatively fl exible.
Packauskas and Schaefer (
1998 ) considered the presence of a triangulin, 2nd valvi-
fers completely fused, and a rigid phallotheca as apomorphies of Pentatomidae
minus Cyrtocoridae. Gapud (
1991 ) placed the Cyrtocorinae + Pentatomidae sensu
stricto as the most apical taxa, “strongly separated from the rest of Pentatomoidea”
by the loss of fi rst valvulae, the absence of the gonangulum, the invagination and
dilation on the spermathecal duct, the retention of membranous fl aps of the 2nd
valvulae, and the presence of an anteroposterior pair of basal sclerites on the sper-
mathecal base. In Grazia et al. (
2008 ) the morphological characters that consistently
support the recognition of a broadly conceived Pentatomidae [Cyrtocorinae
(Aphylinae + Pentatomidae sensu stricto)] are the loss of gonapophyses 8 and the
rst rami , gonapophyses 9 reduced and fused to gonocoxites 9, gonangulum absent,
and the ductus receptaculi dilated and invaginated, forming three distinct walls.
22.4 Subfamily Discocephalinae
22.4.1 Introduction
Discocephalinae (Fig. 22.19 ) is a group of phytophagous species (77 genera; 307
species) restricted to the Neotropical Region. This subfamily was proposed by
Fieber ( 1860 ) as the family “Discocephalida.” It was treated as a subfamily for the
rst time by Stål (
1868 ) and considered by Kirkaldy ( 1909 ) as a tribe of Pentatominae.
More recently, Rolston and McDonald (
1979 ) followed Stål and regarded the group
as a subfamily, providing diagnosis and including 23 American genera removed
from Halyini (Pentatominae). These 23 genera were later grouped in the tribe
Ochlerini by Rolston (
1981 ), and the remaining genera were placed within the nom-
inal tribe Discocephalini. This tribe arrangement of Discocephalinae was followed
by subsequent authors.
J. Grazia et al.
Fig. 22.19 Representative species of Discocephalinae; ( a ) Discocephala marmorea Laporte
(Discocephalini); ( b ) Ochlerus signoreti Breddin (Ochlerini). Scale bars = 1 mm (Photos by LA
Campos & TA Garbelotto)
22.4.2 General Characteristics and Diagnosis
This subfamily includes species usually with fl attened body, small to medium sized,
mottled brown, and black or fuscous. They can be diagnosed primarily by the
insertion of the labium, usually arising on or posterior to the anterior limit of
the eyes (Fig.
22.20 ), and the pairs of abdominal trichobothria usually laterad of the
imaginary line connecting the spiracles (Fig.
22.21 ) – in other subfamilies the
trichobothria are usually in line with spiracles (Rolston and McDonald
1979 ;
1981 ).
The two tribes of Discocephalinae, Discocephalini and Ochlerini, can be distin-
guished from one another by the general aspect, color, and the structure of the
dorsal surface of the third tarsal segment of the hind legs of females, convex in
Discocephalini and shallowly depressed or fl attened in Ochlerini (Fig.
22.22 )
1981 ).
22 Stink Bugs (Pentatomidae)
Fig. 22.20 Ventral view of the head of Tetragonotum megacephalum Ruckes, depicting the inser-
tion of the 1st rostral segment ( i ) placed posterior to the anterior limit of eyes ( dashed line ). Scale
bar = 1 mm (Photo by LA Campos & TA Garbelotto)
Fig. 22.21 Ventral view of part of the abdomen of Opophylax extenebratus Bergroth, depicting
the trichobothria ( t ) placed laterad of imaginary line ( dashed line ) connecting spiracles ( s ). Scale
bar = 1 mm (Photo by LA Campos & TA Garbelotto)
22.4.3 General Biology and Ecology
Discocephalinae are all phytophagous, considered to be phloem feeders (Waldkircher
et al.
2004 ; Guerra et al. 2011 ). Some species of Antiteuchus Dallas (Discocephalini)
are natural hosts of cacao ( Theobroma cacao ) in the Amazon region and in the
Brazilian state of Bahia, and some species of Lincus Stål and Macropygium Spinola
J. Grazia et al.
(Ochlerini) are important vectors of Phytomonas staheli (Trypanosomatidae) in
palms in tropical South America (Costa Lima
1940 ; Desmier-De-Chenon 1984 ;
Couturier and Kahn
1989 ; Llosa et al. 1990 ; Dollet et al. 1993 ).
The morphology of the immatures and the biology within the group is poorly
known; a few studies mention immatures without describing them. Maternal care is
recorded in Discocephalini for species of Antiteuchus (Eberhard
1975 ; Santos and
2001a , b ) with female bugs protecting both egg masses and imma-
tures; trophobiosis with ants is recorded for an unidentifi ed species of Lincus ( Stahel
1964 ) and for Eurystethus ( H .) microlobatus Ruckes (Guerra et al. 2011 ). Field
observations by LA Campos and TA Garbelotto indicate that parental care and tro-
phobiosis with ants also occur in Dinocoris rufi tarsus Ruckes and Catulona pensa
Rolston, respectively.
22.4.4 Classifi cation and Diversity
Systematics of Discocephalinae is yet in a very early stage. Although a fi rst cladistic
approach is available for Ochlerini (Fig.
22.23 ) (Campos and Grazia 2006 ), hypoth-
eses of relationship among genera of Discocephalini are based on morphological
interpretation by classic authors (e.g., Ruckes, Rolston), although lacking at all for
several genera. The relationship among Discocephalinae and other pentatomid sub-
families is also obscure. Campos and Grazia (
2006 ) recovered the monophyly of the
subfamily sustained by two exclusive synapomorphies, fi rst segment of labium
reaching the prosternum and the dorsal surface of the male proctiger membranous
Fig. 22.22 Scanning electron micrographs, third tarsomere of female hind legs of representative
species of Discocephalinae, dorsal; ( a ) Antiteuchus tripterus (F.); ( b ) Schraderiellus cinctus
(Ruckes) (Photos by LA Campos & T Roell)
22 Stink Bugs (Pentatomidae)
Ochlerus b
Ochlerus a
Stalius trisinuatus
Ochlerus handlirsch
Alitocoris a
Alitocoris parvus
Fig. 22.23 Phylogeny of Ochlerini depicting the relationship between genera; strict consensus
tree after successive weighting (Redrawn from Campos and Grazia (
2006 ) )
J. Grazia et al.
at basal third and one homoplastic synapomorphy, metasternum with median carina.
But, as mentioned before, the tribe arrangement adopted for Discocephalinae (with
two tribes, Discocephalini and Ochlerini) is the same since Rolston and McDonald
1979 ) and Rolston ( 1981 ).
Discocephalini comprises 43 genera and 192 species (Table
22.3 ), including the
monotypic fossil genus Acantocephalonotum Petrulevičius & Popov. Members of
this tribe are brown, often mottled with black or shiny black (Figs.
22.19a and 22.24 ),
although a formal diagnosis for this tribe has never been published. A phylogeny for
Table 22.3 Checklist of genera and species of Discocephalini (distribution as available in the
Genera Species
Abascantus Stål A. grandis Becker – BRA
A. lobatus Stål (type) – BRA
A. pubescens Becker – PER
Ablaptus Stål A. amazonus Stål (type) – BRA
A. bolivianus Becker & Grazia – BOL
A. brevirostrum Rolston – COL; CRI; PAN
A. costaricensis Grazia & Zwetsch – CRI
A. phoenix Grazia & Zwetsch – CRI
A. simillimus Becker & Grazia – BOL
A. tavakiliani Rolston – GUF
A. varicornis (Walker) – BRA
Petrulevičius & Popov
A. martinsnetoi Petrulevičius & Popov – ARG
Acclivilamna Ruckes A. vicina (Signoret) (type) – BRA
Agaclitus Stål A. australis Becker & Grazia – BRA
A. dromedarius Stål (type) – BOL; BRA; GUF; PER
A. fallenii Stål – BRA?; GUF
Alcippus Stål A. reticulatus (Stål) (type) – BRA
Allinocoris Ruckes A. nubilus (Dallas) (type) – BRA
Alveostethus Ruckes A. latifrons (Dallas) (type) – BRA
A. politus (Signoret) – BRA; COL; PER; VEN
A. pseudopolitus (Ruckes) – PRY
A. regulosus (Fieber) – BRA
Antiteuchus Dallas A. amapensis Fernandes & Grazia – BRA
A. amplus (Walker) – BOL; BRA; ECU; GUF; PER
22 Stink Bugs (Pentatomidae)
Genera Species
A. bartletti Ruckes – GUY
A. beckerae Fernandes & Grazia – BOL; BRA
A. confi nium Ruckes – ECU; PER
A. costaricensis Ruckes – CRI; PAN
A. cuspidatus Ruckes – COL; PAN
A. doesburgi Fernandes & Grazia – BRA; SUR
A. exiguus Fernandes & Grazia –BRA
A. fulvescens Ruckes – PER
A. geometricus Engleman – BRA
A. graziae Engleman – BOL; BRA; COL; ECU; PER
A. guianensis Ruckes – BRA; GUF; GUY
A. innocens Engleman – BLZ; GTM; MEX
A. kerzhneri Rider –PER
A. ledeburi Fernandes & Grazia – BRA
A. macraspis (Petry) – BRA; COL; CRI; GUF; PAN;
A. maculosus Ruckes – ECU
A. marini Fernandes & Grazia – BRA
A. marmoratus (Erichson, 1848) – GUY
A. melanicus Fernandes & Grazia – PER
A. melanoleucus (Westwood) – BOL; BRA; GUY; PER;
A. mimeticus Ruckes – BRA; PER
A. mixtus (F.) (type) – ARG; BOL; BRA; GUF; GUY;
A. nebulosus Ruckes – BRA
A. nigricans Ruckes – COL; ECU
A. pallescens Stål – BRA; GUY; SUR
A. panamensis (Ruckes) – PAN
A. peruensis Ruckes – PER
A. pictus Ruckes – ECU; PER
A. punctissimus Ruckes – BRA
A. radians Ruckes – BRA
A. rideri Rolston – BRA
A. rolstoni Engleman – COL; ECU
A. ruckesi Rolston – PER
A. rufi tasus Rolston – VEN
A. schuhi Engleman – BRA; ECU
A. sepulcralis (F.) – BOL; BRA; SUR; TTO; VEN
A. simulatus Fernandes & Grazia – BRA
A.tatei (Ruckes) ( species inquirenda ) VEN
A. tesselatus (Westwood) – BRA
Table 22.3 (continued)
J. Grazia et al.
Table 22.3 (continued)
Genera Species
A. tripterus (F.) – ARG; BOL; BRA; COL; ECU; PAN;
A. variegatus Dallas – BOL; BRA; PER
Callostethus Ruckes C. guattatopunctatus (F.) (type) – BRA; PER
C. fl avolineatus Fernandes & Grazia – BRA
Cataulax Spinola C. annulicornis Walker ( insertae sedis ) – BRA
C. eximius (Stål) (type) – BRA
C. froeschneri Grazia, Campos & Becker – BRA
C. pudens (Distant) – PAN; VEN
C. punctipes Walker ( insertae sedis ) – BRA
C. radians Grazia, Campos & Becker – PER
C. subtiliterconspersus Grazia, Campos & Becker – BRA;
C. subvittatus Walker ( insertae sedis ) - BRA
Colpocarena Stål C. complanata (Burmeister) (type) – BRA; SUR; VEN
Coriplatus White C. depressus White – BRA; COL; CUB; GUF; GUY;
Dinocoris Burmeister
Dinocoris Burmeister D . ( D .) antennatus (Dallas) – BOL; BRA; PER
D . ( D .) corrosus (Herrich-Schäffer) – ARG; BRA; PAR
D . ( D .) fabricii Becker & Grazia – unknown
D . ( D .) gibbosus (Fallou) – BRA; PAN; VEN
D . ( D .) gibbus (Dallas) – ARG; BRA; PRY VEN
D . ( D .) histrio (L.) – GUY
D . ( D .) maculatus Laporte – BRA; COL; PER; VEN
D . ( D .) nigroantennatus Becker & Grazia – COL; PER
D . ( D .) ramosus (Walker) – MEX
D . ( D .) reticulatus Becker & Grazia – BRA
D . ( D .) rufi tarsus Ruckes – BRA; COL; HND; NIC; PAN
D . ( D .) variolosus (L.) (type) – GUF; PAN; TTO
Praedinocoris Becker & Grazia D . ( P .) lineatus (Dallas) (type) – BRA; GUF; GUY; PER;
D . ( P .) nigrodecoratus Becker & Grazia – BRA
D . ( P .) prolineatus Becker & Grazia – ARG; BRA; PRY;
Discocephala Laporte D. carvalhoi Becker & Grazia – BRA
D. deplanata Walker – BRA
D. marmorea Laporte (type) – BRA
Discocephalessa Kirkaldy D. andina (Breddin) – ECU
D. humilis (Herrich-Schäffer) – COL
D. notulata (Stål) (type) – CRI; MEX
D. sordida (Walker) – BRA; GUF
D. terminalis (Walker) – BOL; BRA
22 Stink Bugs (Pentatomidae)
Table 22.3 (continued)
Genera Species
Dryptocephala Laporte D. asperula Perty – ?BRA
D. brullei Laporte (type) –BRA
D. crenata Ruckes – PER
D. cydnoides (Perty) – ?BRA
D. dentata Fieber – BRA; URY
D. dentifrons (Latreille) –PER
D. integra Walker – BRA
D. latiloba Stål – BRA
D. lipoloba Ruckes – BRA
D. lurida (Erinchson) – ARG; BRA; COL; GUY
D. maculosa Ruckes – BRA
D. nigricornis Ruckes – BRA
D. obtusiceps Stål – BRA; COL; ECU; PER
D. punctata Amyot & Serville – BRA; PER; VEN
D. spinosa Mayr – BRA
Eurystethus Mayr E. deplanatus Becker – BRA
E. goianensis Becker – BRA
Eurystethus Mayr E . ( E .) ellipsoidalis Ruckes – PAN
E . ( E .) macroconus Ruckes – BRA
E . ( E .) nigropunctatus Mayr (type) – BRA
E . ( E .) ornatus Ruckes – BRA
E . ( E .) ovalis Ruckes – BOL; PER
E . ( E .) pallescens Ruckes – SUR
E , ( E .) parvulus Ruckes –BRA
E . ( E .) sordidus Ruckes – GUF
E . ( E .) spurculus Ruckes – GUF
Hispidisoma Ruckes E . ( H .) fulvescens Ruckes – BRA
E . ( H .) microlobatus Ruckes – BRA; SUR
E . ( H .) nigricornis Ruckes – ECU
E . ( H .) nigroviridis Ruckes – BRA
E . ( H .) punctissimus Ruckes – BRA
E . ( H .) sacculatus Ruckes – BRA
E . ( H .) variegatus Ruckes (type) – BOL; BRA
Glyphuchus Stål G. sculpturatus Stål (type) – BRA
Grassatorama Rider G. nigroventris (Ruckes) (type) – CRI
G. reticulatus (Ruckes) – VEN
G. sinuatus (Ruckes) – CRI
Harpogaster Kormilev H. willineri Kormilev (type) – ARG; BRA
Ischnopelta Stål I. luteicornis (Walker) – BRA
I. oblonga (Fieber) – BRA
I. scutellata (Signoret) (type) – BRA; PRY; VEN
Lineostethus Ruckes L. clypeatus (Stål) (type) – MEX
L. graziae Hildebrand & Becker – MEX
J. Grazia et al.
Table 22.3 (continued)
Genera Species
L. marginellus (Stål) – MEX; USA
L. tenebricornis (Ruckes) – MEX; USA
Mecistorhinus Dallas M. amplus (Walker) – BRA
M. complanatus (Distant) – PAN
M. coralium Ruckes – PER
M. guatemalensis (Distant) – GTM
M. josephi (Stål) – GUF
M. obscurus (Dallas) – MEX
M. rufescens Dallas (type) – BRA
M. semilugens Bergroth – GUF
M. tesselatus (Westwood) – BRA
M. tibialis Ruckes – CRI; PAN
M. variegatus Ruckes – ECU
Oncodochilus Fieber O. integer Breddin – BRA
Oncodochilus Fieber O . ( O .) aradiformis (Herrich-Shäffer) (type) – BRA
O . ( O .) patruelis (Stål) –BRA
Oncoeochilus Breddin O . ( O .) taschenbergi (Breddin) (type) – BRA
O . ( O .) cruciatulus Breddin –BOL
Opophylax Bergroth O. extenebratus Bergroth (type) – BRA
O. signoreti (Distant) – PAN
Pandonotum Ruckes P. bergrothi Becker – BRA
P. punctiventris Ruckes (type) – BRA
Paralcippus Becker & Grazia P. dimidiatus (Ruckes) (type) –ECU
Parantiteuchus Ruckes P. hemitholus Ruckes (type) – GUF
Parvamima Ruckes P. bicolor Ruckes (type) – PAN
P. mexicana Rolston – MEX
Patronatus Ruckes P. binotatus Ruckes (type) – COL; PAN
P. fl avierus Ruckes – VEN
P. punctissimus Ruckes – GUY
Pelidnocoris Stål P. haglundi Ruckes – BRA
P. majusculus Ruckes – PAN
P. stalii Haglund (type) – COL; CRI; PAN; MEX
Phineus Stål P. fucopunctatus Stål (type) – MEX
Phoeacia Stål P. erubescens (Distant) – GTM; PAN
P. gibba (Fieber) – BRA
P. lineaticeps (Stål) (type)– BRA
Placidocoris Ruckes P. bivittatus Ruckes (type) – ARG
Platycarenus Fieber P. umbraculatus (Fieber) (type) – BRA; COL; ECU;GUF;
Priapismus Distant P. costaricensis Rolston – CRI
P. ecuadorensis Rolston – ECU
22 Stink Bugs (Pentatomidae)
Fig. 22.24 Representative species of Discocephalini; ( a ) Acclivilamna vicina (Signoret); ( b )
Coriplatus depressus White; ( c ) Antiteuchus rideri Rolston. Scale bars = 1 mm (Photos by LA
Campos & TA Garbelotto)
Genera Species
P. foveatus Distant (type) – CRI; PAN
P. maculatus Distant – PAN
P. pini Rolston – HND
Psorus Bergroth P. cassidiformis Bergroth (type) –BRA; PER; TTO
P. paraensis Fernandes, Grazia & Lobo – BRA
P. manauara Fernandes, Grazia & Lobo – BRA
Ruckesiocoris Rider R. nitidus (Ruckes) (type) – PAN
Sympiezorhincus Spinola S. punctipes Dallas – BRA
S. tristis Spinola (type) – BRA; GUF; PRY
Tetragonotum Ruckes T. megacephalum Ruckes (type) – BRA
Uncinala Ruckes U. tau Ruckes (type) – BRA
Uncicrus Ruckes U. kollarii (Fieber) (type) – BRA
Insertae sedis and species
Empicoris marmoreus Spinola (see Fernandes
and Grazia
2006 )
a Fossil
b The monotypic genera Anhanga Distant and Braunus Distant (previously included in
Discocephalini) are being transfered to Pentatominae (TdeA Garbelotto, personal communication,
and Barão et al.
in press )
Table 22.3 (continued)
this group is being prepared by TdeA Garbelotto, but groups of genera were recog-
nized by some authors based on morphological similarity, as detailed below.
The genera Abascantus Stål, Coriplatus White, Eurystethus Mayr, Paralcippus
Becker & Grazia, and Pelidnocoris Stål shared the form and proportions of the
metasternum and the spatial relationships of the mesocoxae and metacoxae – where
a metasternum broadly hexagonal, somewhat wider than long (Fig.
22.25 ), causes
J. Grazia et al.
the metacoxae to be placed much farther apart from each other than they are distant
from the respective mesocoxae, the shape of pronotum, and the widely exposed
connexivum (last two not in Eurystethus Mayr) (Ruckes
1966a ; Ruckes and Becker
1970 ; Becker 1977 ; Becker and Grazia 1986 ). Also, Abascantus Stål, Coriplatus
White, and Pelidnocoris Stål have in common the shape of scutellum and gonocox-
ites VIII of external female genitalia much enlarged, covering reduced laterotergites
IX. Paralcippus Becker & Grazia and Eurystethus Mayr share a shorter scutellum
1977 ; Becker and Grazia 1986 ). The genus Alcippus Stål shares with
Paralcippus Becker & Grazia and Eurystethus Mayr the presence of the lateroter-
gites IX and the shorter scutellum, and though the general facies of Alcippus is simi-
lar to that of Paralcippus it has the meso and metacoxae mutually equidistant
(Becker and Grazia
1986 , 1989 ). Pandonotum Ruckes was also placed near
Eurystethus by the shape of the head and scutellum, the labium with an intercalary
unit between the two basal segments, and the reticulate venation of hemelytral
membrane (Ruckes
1965 ).
Ablaptus Stål, Agaclitus Stål, Cataulax Spinola, Sympiezorhincus Spinola, and
Uncinala Ruckes are considered related especially because of the sexual dimorphic
hemelytral membrane, with sclerotized areas and a transversal furrow in males
22.26 ), less clearly differentiated in Cataulax . Species of these genera except
Ablaptus also have the apical angle of corium reaching the apex of the abdomen
1965 ; Becker and Ruckes 1969 ; Becker and Grazia 1989 , 1992 , 1995 ;
Grazia et al.
2000 ).
Fig. 22.25 Ventral view of
thoracic sternites of
Paralcippus dimidiatus
(Ruckes). cx1 procoxae, cx2
mesocoxae, cx3 metacoxae,
me mesosternum, mt
metasternum. Scale
bar = 1 mm (Photos by LA
Campos & TA Garbelotto)
22 Stink Bugs (Pentatomidae)
The larger group of genera within Discocephalini is known as “broadheaded
discocephalines”; it includes 14 genera whose species have the interocular width
equal to or greater than the length of the head (Figs.
22.19a and 22.24a ), the basal
segment of the rostrum projecting onto the prosternum, and the base of abdominal
venter unarmed (Ruckes
1966b ; Becker 1977 ; Rolston 1990 ).
Keys for two of these groups of genera are available, one for the broadheaded dis-
cocephalines (Rolston
1990 ) and another to separate Antiteuchus Dallas of similar
genera (Ruckes
1964 ). Keys for species of several genera are also available (Ruckes
1966a for Eurystethus , 1966b for Alveostethus and Lineostethus , 1966c for
Dryptocephala , 1966d for Pelidnocoris ; Rolston 1984 for Priapismus ; Becker and
1989 for Ablaptus , 1992 for Agaclitus ; Grazia et al. 2000 for Cataulax ; Grazia
and Zwetsch
2000 for Ablaptus ; Fernandes and Grazia 2006 for Antiteuchus ), but some
of them (Ruckes
1966a , b for Lineostethus , Rolston 1984 , and Fernandes and Grazia
2006 ) are out to date since new species were described after the publication of the keys.
Ochlerini was raised by Rolston (
1981 ) and presently includes 32 genera and 115
species (Table
22.4 ) (Garbelotto et al. 2013 ; Cervantes-Peredo and Ortega-León
2014 ; Simões and Campos 2014 ). Species of Ochlerini are, with few exceptions,
recognizable by their dull black or fuscous coloration, and primarily by the shal-
lowly excavated or fl attened superior surface of the third tarsal segment of the hind
legs in females (Fig.
22.22b ) and sometimes in males as well (Rolston 1981 ).
Fig. 22.26 Hemelytral membranes of male of Uncinala tau Ruckes. s sclerotized areas, t transver-
sal furrow. Scale bar = 1 mm (Photo LA Campos & TA Garbelotto)
J. Grazia et al.
Table 22.4 Checklist of genera and species of Ochlerini (distribution as available in the literature)
Genera Species
Adoxoplatys Breddin A. brasiliensis Kormilev – BRA
A. bridarollii Kormilev – BOL
A. comis Breddin – PER; BOL
A. gallardoi Kormilev – ARG
A. giaii Kormilev – ARG
A. minax Breddin (type) – PER; BOL
A. panamensis Kormilev – PAN
A. willineri Kormilev – BOL
Alathetus Dallas A. haitiensis Rolston – HAI
A. rufi tarsis Dallas (type) – JAM
Alitocoris Sailer A. grandis Garbelotto & Campos – PAN
A. lateralis Garbelotto & Campos – CRI
A. ornatus Garbelotto & Campos – PAN
A. schraderi Sailer (type) – CRI; GTM; HND; PAN
A. maculosus Sailer ( sedis mutabilis ) a – GTM; HND
A. brunneus Sailer ( sedis mutabilis ) a
Barola Rolston B. farfala Rolston (type) – PAN
Brachelytron Ruckes B. angelicus Ruckes (type) – BRA
Catulona Rolston C. apaga Rolston – BRA
C. lucida Campos & Grazia – BRA
C. pensa Rolston (type) – BRA
Clypona Rolston C. aerata Rolston (type) – ARG
Coranda Rolston C. castana Rolston – ECU
C. picipes (Stål) (type) – COL; PER
Cromata Rolston C. ornata Rolston (type) – BRA
C. graziae Campos – VEN; BRA
Eritrachys Ruckes E. bituberculata Ruckes (type) – CRI; PAN
E. brailovskyi Ortega-León & Thomas – ECU
Forstona Rolston F. speciosa Rolston (type) – BRA
Herrichella Distant H. thoracica Distant (type) – COL
Hondocoris Thomas H. cavei Thomas (type) – HND
Lincus Stål L. anulatus Rolston – PAN; BRA
L. apollo Dolling – FRG
L. armiger Breddin – BOL
L. bipunctatus (Spinola) – GUY; FRG; BRA
L. breddini Rolston – SUR
L. convexus Rolston – PER
L. dentiger Breddin – PER; ECU
L. discessus (Distant) – CRI; GUY
L. fatigus Rolston – SUR
L. hebes Rolston – PER
L. incisus Rolston – SUR
22 Stink Bugs (Pentatomidae)
Table 22.4 (continued)
Genera Species
L. lamelliger Breddin – COL; FRG; SUR
L. laminatus Rolston – PER
L. lethifer Dolling – ECU
L. leviventris Rolston – PER
L. lobuliger Breddin – BRA
L. malevolus Rolston – PER
L. manchus Rolston – BOL
L. modicus Rolston – ECU
L. operosus Rolston – VEN
L. parvulus (Ruckes) – PER; BRA
L. repizcus Rolston – PER
L. rufospilotus (Westwood) (type) – COL; GUY; SUR;
L. securiger Breddin – BOL; PER; BRA
L. singularis Rolston – PER
L. sinuosus Rolston – PER
L. spathuliger Breddin – PER
L. spurcus Rolston – PER
L. styliger Breddin – COL; PER
L. substyliger Rolston – COL
L. subuliger Breddin – COL; VEN
L. tumidifrons Rolston – PAN; TTO; VEN
L. vallis Rolston – PER
L. vandoesburgi Rolston – SUR
L. varius Rolston – PER
Macropygium Spinola M. reticulare (F.) (type) – BRA
Miopygium Breddin M. cyclopeltoides Breddin (type) – BRA
M. grossa Ruckes – BRA
Moncus Stål M. obscurus (Dallas) (type) – BRA
M. monachus Bergroth – FRG
Neoadoxoplatys Kormilev N. haywardi Kormilev – ARG
N. longirostra Ruckes – COL
N. saileri Kormilev (type) – MEX
N. thomasi Cervantes & Ortega – MEX
Ocellatocoris Campos & Grazia O. dasys Campos & Grazia (type) (MNRJ) – BRA
Ochlerus Spinola O. bergrothi Breddin – UNKNOWN
O. bistillatus Breddin – PER; BOL
O. cinctus Spinola (type) – ?BRA
O. coriaceus Herrich-Schäffer – COL; VEN
O. circummaculatus Stål – BRA
O. communis Breddin – UNKNOWN
O. cotylophorus Breddin – COL; ?PER
J. Grazia et al.
Genera Species
O. dentijugis Breddin – PER
O. handlirschi Breddin – MEX; HND
O. incisulus Breddin – VEN
O. lutosus Herrich-Schäffer – BRA
O. notatulus Breddin – BRA
O. profanus Breddin – PER
O. rusticus Breddin – BRA
O. signoreti Breddin – FRG
O. stylulatus Breddin – PER
O. tenuicornis Breddin – VEN
Orbatina Ruckes O. fuligina Ruckes (type) – PAN; COL; BOL
Paralincus Rolston P. bimaculatus (Ruckes) – GUY
P. silvae Rolston – SUR; BRA
P. terminalis (Walker) (type) – BRA
P. sordidus (Herrich-Shäffer) ( nomen dubium ) – BRA
Parastalius Matesco, Grazia, &
P. trisinuatus (Rolston) (type) – CRI; PAN
P. rolstoni Matesco, Grazia & Campos – CRI
Parochlerus Breddin P. latus Breddin (type) – PER; BRA
Phereclus Stål P. pluto Stål (type) – COL
P. punctatus (Dallas) – COL
P. antennatus Distant – COL
Pseudadoxoplatys Rolston P. mendacis Rolston (type) – PER; BOL
Schaefferella Spinola S. fusca Rolston – ECU; PER
S. incisa (Herrich-Schäffer) (type) – BRA
Schraderiellus Rider S. cinctus (Ruckes) (DBTC) – CRI
S. hughesae (Ruckes) (type) – CRI
Similiforstona Campos & Grazia S. bella Campos & Grazia (type) – BRA
Stalius Rolston S. castaneus (Distant) – NIC
S. leonae Ortega – MEX, GUA
S. scutellatus Thomas – CRI
S. tartareus (Stål) (type) – MEX; HND; COL
Tetrochlerus Breddin T. fi ssiceps Breddin (type) – ECU
Uvaldus Rolston U. concolor Rolston (type) – BRA
Xynocoris Garbelotto & Campos X. calosus Garbelotto & Campos – BOL
X. crinitus Garbelotto & Campos – BRA
X. cupreus Garbelotto & Campos – BRA
X. egeri Garbelotto & Campos – CRI, BRA
X. insularis Garbelotto & Campos – TTO
X. lucidus Garbelotto & Campos – BRA
X. meridionalis Garbelotto & Campos – BRA, PAR
Table 22.4 (continued)
22 Stink Bugs (Pentatomidae)
The most comprehensive systematic work for Ocherini was made by Rolston
1992 a), including the description of nine genera, diagnosis for the tribe, and key
and diagnosis for its genera. The phylogeny of genera of Ochlerini and its sister-
group relationship to Discocephalini were recently the theme of a cladistic study
(Campos and Grazia
2006 ) (Fig. 22.23 ). Unambiguous synapomorphies of
Discocephalinae are the long fi rst segment of labium, reaching prosternum, and the
male segment X largely membranous at least on its basal third (Campos and Grazia
2006 , characters 13 and 37). Unambiguous synapomorphy of Ochlerini is the
presence of a dorsal depression of the third tarsal segment of the hind legs of females
(Campos and Grazia
2006 , character 30).
The most basal genera within Ochlerini are Adoxoplatys Breddin, Neoadoxoplatys
Kormilev, and Pseudadoxoplatys Rolston, whose species can be easily distinguished
from other ochlerines by a usually long and fl attened body, light-brown colored and
of somewhat glossy aspect (Fig.
22.27a ). All other genera were grouped into two
clades, named Ocellatocoris and Herrichella clades by Campos and Grazia (
2006 ).
The fi rst includes the type genus Ochlerus Spinola and seven other genera
22.19b ), and the later includes the remaining 20 genera known by that time
22.27b ), except Hondocoris Thomas that was not included in the cladistic
analysis. The Herrichella clade was subject of another cladistic analysis that aimed
to give a phylogeny-based classifi cation of the genus Alitocoris Sailer and related
species (Fig.
22.28 ) (Garbelotto et al. 2013 ).
The checklist for genera and species of Ochlerini provided by Campos and
Grazia (
2006 ) is updated here, with the inclusion of taxonomic information from
several sources (Arismendi and Thomas
2003 ; Matesco et al. 2007 ; Ortega-León
and Thomas
2010 ; Garbelotto et al. 2011 , 2013 ; Simões and Campos 2014 ;
Cervantes-Peredo and Ortega-León
2014 ) (Table 22.4 ).
22.4.5 Remarks
The study of Discocephalinae remains a challenging issue, since species of most
genera are poorly represented in scientifi c collections and their natural history is
virtually unknown. The description of life cycles and morphology of immatures are
Table 22.4 (continued)
Genera Species
X. parvus (Distant) (type) – CRI, PAN, VEN, SUR,
X. recavus Garbelotto & Campos – VEN, BRA
X. tuberculatus Garbelotto & Campos – VEN, BRA
a See Garbelotto et al. ( 2013 )
b There are several undescribed species of Macropygium occurring in different countries, from
Mexico to Argentina
J. Grazia et al.
strongly needed. Behaviors registered for a few species such as parental care and
trophobiosis with ants are interesting research themes, as well as feeding relation-
ship with plants especially of Ochlerini. Phylogeny and systematics of
Discocephalinae are also in need for investigation, focusing the review and modern
taxonomic treatment of genera, use of molecular data in cladistic analyses, and
relationship with other subfamilies of Pentatomidae.
22.5 Subfamily Edessinae
22.5.1 Introduction
Edessinae is one of the largest subfamilies of Pentatomidae with about 300 species
(Schuh and Slater
1995 ; da Silva et al. 2013 ) distributed only in Neotropical Region,
except for two species found in the Southern United States of America [ Edessa
bifi da (Say) and E. fl orida Barber] (McPherson
1982 ). This subfamily includes
seven genera without tribal division. Edessinae was until recently treated as a tribe
of Pentatominae but was raised to subfamily by Rolston and McDonald (
1979 ). This
taxon has well-established limits but they are quite similar to the limits of the genus
Edessa , causing a lot of confusion and turning the latter a depot of species.
Fig. 22.27 Representative species of Ochlerini; ( a ) Adoxoplatys comis Breddin; ( b ) Xynocoris
parvus (Distant). Scale bars = 1 mm (Photos by LA Campos & TA Garbelotto)
22 Stink Bugs (Pentatomidae)
Herrichella thoracica
Tetrochlerus fissiceps
Similiforstona bella
Alitocoris ornatus
Alitocoris schraderi
Alitocoris lateralis
Alitocoris grandis
Brachelytron angelicus
Schaefferella incisa
Forstona speciosa
Miopygium cyclopeltoides
Macropygium reticulare
Clypona aerata
Uvaldus concolor
Alathetus rufitarsis
Alitocoris maculosus
Alitocoris brunneus
Xynocoris crinitus
Xynocoris recavus
Xynocoris tuberculates
Xynocoris parvus
Xynocoris insularis
Xynocoris egeri
Xynocoris calosus
Xynocoris meridionali
Xynocoris lucidus
Xynocoris cupreus
sp. 05
sp. 10
sp. 15
sp. 17
Fig. 22.28 Phylogeny of the Herrichella clade depicting the relationship between species of the
Alitocoris group; strict consensus tree after implied weighting (Redrawn from Garbelotto et al.
2013 ) )
J. Grazia et al.
22.5.2 General Characteristics and Diagnosis
Medium to large size bugs (1–3 cm long). Very attractive bugs with color ranging
from light green to greenish-brown with highly contrasting connexivum and ventral
surface. Head short and wide. Antennae 4 or 5 segmented. Humeral angles ranging
from not to highly developed conical or fl at, apex rounded, or acute. Metasternum
strongly produced anteriorly onto mesosternum (prosternum in Pantochlora Stål)
and usually bifi d except for Pantochlora and Doesburgedessa that show a single
projection. First rostral segment short, reaching anterior notched part. Posterior part
of metasternal process also notched, receiving medial abdominal tubercle (Stål
1872 ; Rolston and McDonald 1979 ; Schuh and Slater 1995 ). Distal angle of the
abdominal segment VII range from before the level of genital plates to surpassing
them by far (Stål
1872 ). Dorsal rim of pygophore with a superior process. Ventral
surface of the body usually lighter than dorsal with dark stripes at least on abdomi-
nal intersegmentar areas (Figs.
22.29 , 22.30 , 22.31 , 22.32 , 22.33 , and 22.34 ).
Female internal genitalia: Description is almost the same of Pentatominae but
capsula seminalis with 3 digitiform processes and a beak-like projection on the
thickening of the vaginal intima (Barcellos and Grazia
2003b ).
Fig. 22.29 Edessa
oxyacantha Breddin (Photo
by F Carvalho-Filho)
22 Stink Bugs (Pentatomidae)
Fig. 22.30 Edessa laticornis Stål: ( a ) adult; ( b ) 5th instar nymph (Photo by F Carvalho-Filho)
Fig. 22.31 Edessa bifi da
(Say) (Photo by F
J. Grazia et al.
Fig. 22.32 Edessa elaphus Breddin (Photo by F Carvalho-Filho)
Fig. 22.33 Edessa
dolichocera Burmeister,
lateral view (Photo by F
22 Stink Bugs (Pentatomidae)
Fig. 22.34 Edessa dolichocera Burmeister, frontal view (Photo by F Carvalho-Filho)
Male internal genitalia: The phallus of Edessinae is quite simple; phalloteca is
strongly esclerotized, tubular to pear-like, and elongated; conjunctiva absent; vesica
small, elliptical in lateral view; and ductus seminis distalis short, not or slightly
surpassing the apex of vesica.
22.5.3 General Biology and Ecology
Edessinae are all plant feeder bugs, and only one species is economically important
as pests of cultivated crops – Edessa meditabunda (F.) (Fig.
22.35 ), particularly on
soybean and on tobacco (Silva et al.
1968 ; Rizzo 1971 ; Lopes et al. 1974 ; Panizzi
et al.
2000 ). Another species is recognized as a secondary pest – Edessa rufomar-
ginata (De Geer) – that feeds on Leguminosae but is more often found on Solanaceae,
causing damage to tomato (Silva et al.
1968 ; Rizzo and Saini 1987 ; Panizzi et al.
2000 ). The species mentioned before are very common throughout South and
Central Americas due to their capability to invade areas modifi ed by man and are
well studied with several registers over time. On the other hand, others may have
local importance and are barely or not studied so far– e.g., a not described species
that is an important pest in camu camu (Myrtaceae) crops in Pucallpa, Peru
(Iannacone et al.
2007 ). Several other species can be found on cultivated or native
plants of economical or ornamental interest but cause no apparent damage: Edessa
scabriventris Stål on guava and Brazilian cherry (Lima et al.
2010 ) and Paraedessa
stolida (L.) on acerola (JAM Fernandes, personal observation).
Only a few works concentrated on economically important species were pro-
duced so far. Rizzo (
1971 ) presented detailed information on the biology, immature
stages, fi eld ecology, and host plants of Edessa meditabunda . This species has been
the subject of studies regarding its association with and damaging potential to soy-
J. Grazia et al.
bean (Silva et al. 2012 ) and sunfl ower (Malaguido and Panizzi 1998b ) and nymph
development and feeding behavior (Panizzi and Machado-Neto
1992 ); also, the
behavior of newly hatched nymphs on the corions has been recently studied in detail
(Calizotti and Panizzi
2014 ).
Rizzo and Saini (
1987 ) presented biology data on Edessa rufomarginata . This
species was subject of another study focused on the description of the immature
stages (Fortes and Grazia
1990 ) under controlled laboratorial conditions. The same
species was observed and studied in the fi eld revealing several interesting ecological
data including trophobiotic interaction between nymphs of E. rufomarginata and
ants (Silva and Oliveira
2010 ). A species not recognized as pest – Edessa scabriven-
tris – had some ecological and biological characteristics revealed in Lima et al.
2010 ). A probably new species considered pest in camu camu crop had its biology,
immature stages, and fi eld ecology studied (Iannacone et al.
2007 ).
22.5.4 Classifi cation and Diversity
The subfamily Edessinae Amyot & Serville has seven genera: Edessa F. (about 280
species), Brachystethus Laporte (10 species), Peromatus Amyot & Serville (seven
species), Olbia Stål (three species), Pantochlora Stål (one species), Doesburgedessa
Fernandes (fi ve species), and Paraedessa Silva & Fernandes (nine species). The
genus Brachystethus is recognized by the short metasternal process and anterior
apex slightly notched. Peromatus has only four antennal segments. Olbia has a pair
of spines on the posterior margin of the pronotum, near the scutellum base.
Fig. 22.35 Edessa meditabunda (F.) (Courtesy of JJ Silva)
22 Stink Bugs (Pentatomidae)
Pantochlora is characterized by the anterior apex of the metasternal process not
bifurcate, projected between procoxae, laterally fl attened, and attached to the ster-
num. Doesburgedessa is recognized by the metasternal process with a single conical
anterior projection which is detached from the mesosternum. Paraedessa has a
reduction of the genital plates of the females plus a remarkable esclerotization of the
gonapophyses 8; males show a lateral expansion of the proctiger and a dark crest
covering base of the paramere. Edessa is composed of several groups of species
with the metasternal process clearly bifurcating anteriorly but variable in many
other characteristics (Fernandes
2010 ). Edessinae is a monophyletic group Barcellos
and Grazia
2003a ) supported by several characteristics, but its composition at supra-
specifi c level is still not clear due to Edessa that is hiding the real diversity of the
subfamily. Studies are revealing different groups of species (e.g., Fernandes and
Van Doesburg
2000 ; Fernandes and Campos 2011 ; da Silva and Fernandes 2012 )
and also new genera Doesburgedessa (Fernandes
2010 ) and Paraedessa (da Silva
et al.
2013 ). The heterogeneity of Edessa can be confi rmed by its confusing taxo-
nomical and nomenclatural history that culminates with Kirkaldy’s catalog (
1909 )
where he listed the subgenera but considered himself unable to include the species
in them (footnote p. 153). Genus Edessa despite its size still has hundreds of new
species to be described. New taxonomical studies probably will split Edessa in dif-
ferent taxa. A phylogenetic study is also important to present a hypothesis of rela-
tionship among taxa and confi rm or not that Edessa is a paraphyletic taxon. In this
book the genera Lopadusa , Mediocampus , Neopharnus , Pharnus , Platistocoris , and
Praepharnus are included in Pentatomini (Pentatominae).
Key to genera of Edessinae and species group of Edessa . The concept of Stål (
1872 )
dividing the genus in species groups is adopted here; genus in italic and group of
species in bold .
1. Body rounded; apex of metasternal process level with mesocoxae, anteriorly
raised, and sulcated but not clearly bifurcated ................... Brachystethus Laporte
Body with different shapes but usually elongated, lateral margins subparallel,
metasternal process not as above .........................................................................2
2. Metasternal process anteriorly clearly bifurcated ................................................ 4
Metasternal process projected anteriorly in a simple process .............................. 3
3. Metasternal process long, reaching mesocoxae, laterally compressed, and
fused with sterna, and apex widely rounded; humeral angle slightly
projected and acute; found in Central America .......................... Pantochlora Stål
Metasternal process short, not reaching procoxae, conical, not in contact with
sterna, apex truncate; humeral angle projected laterally or dorsolaterally,
conical, apex sharp; found in Amazon region .............Doesburgedessa Fernandes
4. Four segmented antennae; second antennal segment very long, at least three
times longer than the fi rst; humeral angle truncate or globose; paramere robust
and large; specimens usually bigger than 2 cm ........Peromatus Amyot & Serville
Five segmented antennae; second antennal segment short, at most twice longer
than the fi rst; humeral angle and paramere strongly variable in shape;
specimens usually smaller than 2 cm ................................................................... 5
J. Grazia et al.
5. Posterior margin of the pronotum with a large pair of spines overlapping
basal angles of scutellum; humeral angles long, sinuous, and laterally
directed; pronotal disk with green and yellow stripes connecting
humeral angles .......................................................................................Olbia Stål
Posterior margin of the pronotum without large spines or other projections;
humeral angles variable in shape; pronotal disk not as above ( Edessa ) .............. 6
6. Humeral angles rounded or sharp but not laterally developed more than the
distance between the eyes ....................................................................................7
Humeral angles variable in shape and laterally developed more than the distance
between the eyes ................................................................................................ 11
7. Species elongate; abdominal segment VII projected posteriorly and contrastingly
dark at least in distal half ..................................................................................... 8
Species elliptical; abdominal segment VII not posteriorly projected and
concolorous with connexivum ...........................................................................10
8. Corium brown with yellow veins; dark rounded spots below the spiracles;
seventh abdominal segment mostly contrastingly dark dorsally, ventrally
concolorous, posterior projection at least 1.5 times longer than wide
at base ........................................................................... Pygoda Amyot & Serville
Corium brown variegate with whitish and blackish spots; without dark
spots below spiracles; abdominal segment VII contrastingly dark dorsally
only in distal half, ventrally with spots or stripes contrastingly dark,
posterior projection at most 1.2 times longer than wide at base .......................... 9
9. Posterior part of proctiger expanded laterally; behind this part bristles short
and sparse; pygophore with a rough ridge on each side of proctiger; gonocoxites
8 slightly to strongly reduced, leaving visible part of gonapophyses 8; gonapophyses 8
strongly developed and sclerotized ....................... Paraedessa Silva & Fernandes
Posterior part of proctiger not expanded laterally at most with a small carina
and dense tuft of long bristles; pygophore without ridge close to
proctiger; gonocoxites 8 not reduced and leaving visible a tiny part of
gonapophyses 8 at most; gonapophyses 8 not developed and membranous
..................................................................................... Hypoxys Amyot & Serville
10. Pronotal disk tumid; costal margin of corium uniformly colored and punctured;
connexivum with punctures dark .........................................................Ascra Say
Pronotal disk not tumid. Costal margin of corium with punctures
and usually color distinct from the rest; connexivum with punctures concolor
.............................................................................. Aceratodes Amyot & Serville
11. Humeral angle long, apex acute. Abdominal segment VII with apex twisted,
strongly projected posteriorly, surpassing considerably the genital plates.
Pygophore with a yellow spot on the base of each genital cup
process ................................................................. Dorypleura Amyot & Serville
Humeral angle long or short; apex acute, truncate, or globose, black or yellow
contrasting with general coloration. Abdominal segment VII with apex not
twisted projected posteriorly but not or slightly surpassing the genital plates.
Pygophore without any spot on the base of genital cup process ........... Edessa F.
22 Stink Bugs (Pentatomidae)
22.5.5 Remarks
Edessinae was treated as a tribe of Pentatominae and was raised to subfamily by
Rolston and McDonald (
1979 ). This taxon has well-established limits; neverthe-
less its concept is quite similar to the genus Edessa . They are well represented in
scientifi c collections, and modern taxonomical studies probably will split Edessa
in different taxa. Recent studies are revealing different groups of species and also
new genera within Edessa , a probably a paraphyletic taxon, deserving a phyloge-
netic study to establish the relationships within the subfamily and among the
22.6 Subfamily Pentatominae
22.6.1 Introduction
Pentatominae is the most diverse subfamily of Pentatomidae and includes species
distributed in all the regions of the world (Schuh and Slater
1995 ; Grazia et al.
1999 ). The exact number of groups (tribes, genera, and species) gathered in
Pentatominae is uncertain, since the defi nition of the subfamily is still under discus-
sion. Nonetheless, it is possible to estimate that between 400 and 600 genera and
2,700 and 3,300 species are included in Pentatominae (Grazia et al.
1999 ; Cassis
and Gross
2002 ; Rider 2014 ). In the Neotropical Region, approximately 120 genera
and about 680 species occur.
22.6.2 General Characteristics and Diagnosis
Extremely variable in color, form, and size. First segment of labium arising anterior
to the line of anterior margin of the eyes, with variable length in relation to the buc-
culae. Humeral angles frequently with process (acute or not); scutellum not reach-
ing the apex of the abdomen, with frenal region extending from 2/5 to 2/3 the length
of the scutellum. Metasternum rarely produced anteriorly. Abdominal segments III
to VII with two pairs of trichobothria, near the spiracular line (Rolston and
1979 ; Schuh and Slater 1995 ) (Figs. 22.36 , 22.37 , 22.38 , 22.39 , 22.40 ,
22.41 , 22.42 , and 22.43 ).
Female internal genitalia: vesicular region of ductus receptaculi membranous,
invaginated into a sclerotized rod; capsula seminalis with anterior and posterior
annular fl anges, bearing or not processes of variable number (Rolston and McDonald
1979 ; Schuh and Slater 1995 ). Male internal genitalia: conjunctiva usually membra-
nous, sometimes absent; vesica usually as a sclerotized tube, with variable length
(Rolston and McDonald
1979 ; Schuh and Slater 1995 ).
J. Grazia et al.
22.6.3 General Biology and Ecology
Pentatominae are all phytophagous and many species are economically important as
pests of cultivated crops (Grazia et al.
1999 ; Panizzi et al. 2000 ). In the Neotropics
the Pentatominae cause damage mainly in Leguminosae and Gramineae crops.
Nezara viridula (L.) (Fig.
22.40 ), Piezodorus guildinii (Westwood) (Fig. 22.44 ), and
Euschistus heros (F.) (Fig.
22.45 ) are important pests of soybean (Panizzi et al. 2000 ),
Fig. 22.36 Oenopiella punctaria Stål (Photo by A Ferrari)
Fig. 22.37 Agroecus
scabricornis Herrich-
Schaeffer (Photo by L
22 Stink Bugs (Pentatomidae)
Fig. 22.38 Euschistus
irroratus Bunde, Grazia, &
Mendonça Jr. (Photo by J
Fig. 22.39 Pellaea stictica
(Dallas) (Courtesy of
J. Grazia et al.
Fig. 22.40 Nezara viridula
(L.) (Photo by A Ferrari)
Fig. 22.41 Phalaecus sp.
(Photo by J Grazia)
22 Stink Bugs (Pentatomidae)
Fig. 22.42 Roferta marginalis (Herrich-Schaeffer) (Photo by JAM Fernandes)
Fig. 22.43 Stictochilus
tripunctatus Bergroth (Photo
by R Lupoli)
and species of Arvelius Spinola, Loxa Amyot & Serville, Agroecus Dallas, Dichelops
Spinola, and Chinavia Orian also occur in this crop, sometimes causing damage
(Panizzi and Slansky
1985 ; Panizzi et al. 2000 ; Schwertner and Grazia 2007 ).
Mormidea v - luteum (Lichtenstein), Oebalus poecilus (Dallas), Oebalus ypsilon-
griseus (De Geer) (Fig.
22.46 ), and Tibraca limbativentris Stål (Fig. 22.47 ) cause
J. Grazia et al.
damage to rice and wheat (Panizzi et al. 2000 ). Also, Glyphepomis Berg species are
rice pests (Campos and Grazia
1998 ; Farias et al. 2012 ) and Dichelops melacanthus
(Dallas) (Fig.
22.48 ) is considered an important problem in wheat (Chocorosqui and
2004 ; Manfredi-Coimbra et al. 2005 ) and corn crops (Ávila and Panizzi
1995 ) in Brazil.
Loxa spp. and Arvelius albopunctatus (De Geer) (Fig.
22.49 ) cause damage in
Solanaceae plants in Brazil (Panizzi et al.
2000 ). The last one is also reported to feed
in tobacco in Brazil (Marques et al.
2006 ).
Pentatominae are, in general, highly polyphagous, and their life history includes
several host plant sequences. For some species this has been investigated in details
in the Neotropics, as illustrated by the studies conducted by Panizzi (
1997 ).
Fig. 22.44 Piezodorus guildinii (Westwood) (Courtesy of JJ Silva)
Fig. 22.45 Euschistus heros (F.) (Courtesy of JJ Silva)
22 Stink Bugs (Pentatomidae)
Fig. 22.46 Oebalus
ypsilongriseus (De Geer)
(Courtesy of H. Prando)
Fig. 22.47 Tibraca
limbativentris Stål (Courtesy
of H. Prando)
J. Grazia et al.
22.6.4 Classifi cation and Diversity
Pentatominae is not a monophyletic group (Gapud 1991 ; Grazia et al. 2008 ) and
does not have a defi nition based on phylogenetic studies. Therefore, authors diverge
about composition of the subfamily (Cassis and Gross
2002 ). Schuh and Slater
1995 ), for example, included eight tribes in Pentatominae, from which only
Pentatomini is registered in the Neotropical Region. Rider (
2014 ) considers 42
tribes in Pentatominae, 11 registered in Neotropics: Catacanthini Atkinson,
Carpocorini Mulsant & Rey, Halyini Amyot & Serville, Mecideini Distant, Menidini
Atkinson, Nezarini Atkinson, Pentatomini Leach, Piezodorini Atkinson, Procleticini
Pennington, Sciocorini Amyot & Serville, and Strachiini Mulsant & Rey. The rela-
tionship among tribes is unknown; also most of the tribes do not have diagnostic
Fig. 22.48 Dichelops melacanthus (Dallas) (Courtesy of JJ Silva)
Fig. 22.49 Arvelius albopunctatus (De Geer) (Courtesy of JJ Silva)
22 Stink Bugs (Pentatomidae)
characteristics to defi ne them, which make the establishment of the exact composi-
tion of these groups diffi cult. Six genera of the Neotropical Region are considered
as unplaced in Rider’s classifi cation (Table
22.5 ).
Table 22.5 Checklist of Neotropical Pentatominae genera (distribution as available in the
Acledra Signoret BRA, CHI, ARG, URU
Agroecus Dallas PER, ECU, BOL, GUY, FGU, COL, VEZ, BRA, PER, BOL,
Amauromelpia Fernandes &
Berecynthus Stål MEX, CR, PAN, COL, VEZ, SUR, BRA, PER
Braunus Distant CR, VEZ, COL, ECU, PER, BOL
Bucerocoris Mayr BRA
Caonabo Rolston BRA
Caribo Rolston BAH, JAM, IS. VIR
Caracia Stål VEZ
Copeocoris Mayr BRA, ARG
Cosmopepla Stål USA, MEX, ELS, CR, VEZ, COL, ECU
Cradia Bergroth BRA
Curatia Stål BRA
Dichelops Spinola PAN, BRA, ECU, PAR, VEZ, BRA, PER, BOL, PAR, ARG,
Epipedus Spinola BRA
Euschistus Dallas USA, MEX, NIC, GTM, ELS, HON, CR, PAN, COL, VEZ,
Galedanta Amyot &
Glyphepomis Berg BRA, ARG, URU
Hypanthracos Grazia &
Hypatropis Bergroth HON, NIC,CR, PAN, TTO, COL, VEZ, SUR, BRA, ARG, URU
Lattinidea Rider & Eger CR, PAN, COL, VEZ, FGU, BRA, ECU, PER
Lattinellica Rider & Eger COL, BRA, ECU, PER
Ladeaschistus Rolston BRA, BOL, PAR, ARG, URU
Luridocimex Grazia,
Fernandes & Schwertner
Mathiolus Distant GTM
Mecocephala Dallas BRA, ARG, URU
Menudo Thomas PUR
Mormidea Amyot & Serville CAN, USA, MEX, NIC, PUR, CUB, HIS, DRE, COL, GUY,
J. Grazia et al.
Table 22.5 (continued)
Oebalus Stål USA, MEX, W.I. (ALL), SUR, BRA, PER, PAR, ARG, URU,
Oenopiella Bergroth BRA, CHI, ARG, URU
Ogmocoris Mayr BRA
Padaeus Stål USA, MEX, GTM, NIC, HON, CR, ECU, COL?
Parahypatropis Grazia &
Paramecocephala Benvegnú SUR, GUY, GUF, BRA, ECU, ARG, URU
Paratibraca Campos &
Parentheca Berg BRA, PAR, ARG, URU
Pedinonotus Fernandes &
Pentatomiana Grazia &
Poriptus Stål TTO, COL, BRA, BOL, ARG
Prionotocoris Kormilev BOL, ARG
Proxys Spinola USA, MEX, DRE, CUB, GTM, NIC, FGU, BLZ, CR, GRE,
Spinalanx Rolston & Rider COL, PER, ECU, BOL
Stysiana Grazia, Fernandes
& Schwertner
Trichopepla Stål CAN, USA, MEX
Arocera Spinola MEX, BLZ, HON, CR, PAN, CUB, JAM, PUR, DRE, TTO,
Boea Walker CR, BRA
Rhyssocephala Rider MEX, BLZ, GTM, CR, PAN, JAM, COL, VEZ, ECU, BRA,
Runibia Stål USA, JAM, Virgen Is, COL, VEZ, ECU, BRA, PER, BOL,
Vulsirea Spinola USA, MEX, HON, HAI, JAM, CR, CUB, PAN, COL, VEZ,
Brochymena Amyot &
Parabrochymena Larivière USA, MEX, GTM, CR
22 Stink Bugs (Pentatomidae)
Table 22.5 (continued)
Elanela Rolston SUR, BRA, PER
Chinavia Orian CAN, USA, MEX, BLZ, CR, CUB, DRE, PUR, JAM, Grenada,
Guadeloupe, Dominica, St. Croix, VIR IS., GTM, NIC, HAI,
Chlorochroa Stål CAN, USA, MEX
Nezara Amyot & Serville MEX, NIC, HON, CUB, DRE, PAN, VEZ, BRA, CHI, ARG,
Roferta Rolston BLZ, PUR, PAN, TRI, CUB, BRA, ARG
Adevoplitus Grazia &
Arvelius Spinola DRE, GUP, MAR, BAR, SVI, USA, MEX, ELS, GTM, HON,
Brasilania Jensen-Haarup BRA
Chlorocoris Spinola USA, MEX, BLZ, NIC, HON, GTM, JAM, CR, PAN, COL,
Chloropepla Stål BRA, BOL, VEZ, CR, PER, GUY, FGU,
Disderia Bergroth MEX, GTM, BLZ
Elsiella Froeschner ECU
Eludocoris Thomas CR
Evoplitus Amyot & Serville BRA
Fecelia Stål PUR, HAI, DRE
Glaucioides Thomas HON, CR, PAN, VEZ, SUR, GUY, BRA
Grazia Rolston MEX, CUB, DRE, PAN, VEZ, BRA, PAR
Janeirona Distant BRA, PER, PAR, ARG
Kermana Rolston USA, MEX, CR, HON, PAN, BRA, ARG, URU
Loxa Amyot & Serville USA, MEX, CUB, DRE, BAH, PUR, DOMINICA,
Marghita Ruckes BRA, ARG
Mayrinia Horváth NIC, CR, COL, VEZ, GUY, BRA, PER, BOL, PAR, ARG
Mediocampus Thomas DOR
Modicia Stål CUB
Myota Spinola BRA
J. Grazia et al.
Table 22.5 (continued)
Neopharnus Van Duzee EUA, CUB
Neotibilis Grazia &
Nocheta Rolston SUR, BRA
Pallantia Stål MEX, GTM, PAN, VEZ, PER, BRA, PAR, ARG
Paratibilis Ruckes MEX, PER
Phalaecus Stål GUY, SUR, GUF, BRA, COL
Pharnus Stål CUB
Placocoris Mayr BRA, ARG
Platistocoris Rider BRA
Praepharnus Barber &
Pseudevoplitus Ruckes VEZ, BRA, PER, BOL, ARG
Ramosiana Kormilev MEX, CR, PAN, PER, BOL
Rhyncholepta Bergroth PAN, VEZ, GUF, VEZ, BRA, BOL
Rideriana Grazia &
Similliserdia Fortes &
Stictochilus Bergroth ARG, BRA
Taurocerus Amyot &
Tibilis Stål BRA, ECU, PER, BOL
Vidada Rolston PER
Piezodorus Fieber USA, TRI, PUR, GTM, HON, NIC, JAM, CUB, DRE, St.
Vincent, Grenada, PAN, VEZ, BRA, ECU, PAR, ARG
Aleixus McDonald BRA
Brepholoxa Van Duzee USA, MEX, W.I.
Dendrocoris Bergroth USA, MEX, GTM,
Lobepomis Berg ARG
Neoderoploa Pennington BOL, ARG
Odmalea Bergroth GUF, TTO, PAN, VEZ, COL, BRA, BOL, ARG, URU
Parodmalea Rider BRA
Procleticus Berg ARG
Pseudobebaeus Distant BRA, PER
Terania Pirán ARG
Thoreyella Spinola BRA, PAR, ARG, URU
22 Stink Bugs (Pentatomidae)
Table 22.5 (continued)
Murgantia Stål USA, MEX, GTM, ANT, CR, PAN, BRA, PER, BOL
Trincavellius Distant ECU, CHI, PER
Capivaccius Distant MEX, HON, PAN, BRA, ARG
Cyptocephala Berg USA, CENTRAL AM. (ALL), BAH, CUB, DRE, JAM, VIR
Patanius Rolston BRA
Senectius Rolston PER
Tepa Rolston & McDonald USA, CURAÇAO
Catacanthini was established by Atkinson ( 1888 ) to include Catacanthus Spinola
(15 spp.) and Chalcocoris Dallas (two spp.), both genera not known in the
Neotropical Region. More fi ve Neotropical genera were added since then
22.5 ). Some of the most colorful species of Pentatominae are included in
Catacanthini. Keys for identifi cation of species are available for Arocera (Rider
1992 ), Rhyssocephala (Rider 1991 ), and Runibia (Zwetsch and Grazia 2001 ). A
study of the relationships within Catacanthini using morphologic characters is
under development (J. Grazia, unpublished).
Carpocorini was fi rst defi ned by Mulsant and Rey (
1866 ), including Carpocoris
Kolenati, Dryocoris Mulsant & Rey (junior synonym of Holcostethus ), and
Peribalus Mulsant & Rey (junior synonym of Holcostethus ). This tribe is one of the
most diverse in the subfamily (Cassis and Gross
2002 ) and includes many of the
species of economic importance as Euschistus heros (F.), Oebalus poecilus (Dallas),
and Tibraca limbativentris Stål. In the Neotropical Region, 53 genera are included
in Carpocorini. A phylogenetic analysis based on morphology and DNA sequences
is under development (J. Grazia, unpublished).
The tribe Halyini was established by Amyot and Serville (
1843 ) to include a
heterogeneous group of 16 genera, which currently included different families of
Pentatomoidea and subfamilies and tribes of Pentatomidae. Dallas (
1851 ) consid-
ered the group as a separate family; however the Halyini as currently circumscribed
share all synapomorphies of the Pentatomidae. The tribe includes 82 genera with
worldwide distribution, with great diversity in the Afrotropical and Australian
regions (Linnavuori
1982 ; Gross 1975 ). Two genera are recorded in the Western
Hemisphere (Table
22.5 ), with species found in the northern Neotropical Region.
Distant ( 1902 ) propose Mecideini to include Aenaria Stål and Mecidea Dallas.
The proposal was questioned by Bergroth (
1905 ), and Mecidea was grouped with
other seven genera today included in the tribe Diemeniini Kirkaldy. Mecideini was
J. Grazia et al.
accepted as a separated tribe (i.e., Rolston and McDonald 1979 ) but remains mono-
typic. The genus Mecidea includes 16 species distributed in all biogeographic
regions except the Australian region (Sailer
1952 ). In the Neotropics, Mecidea has
a disjunct distribution, with 4 described species (Sailer
1952 ; Thomas 2000 ; Grazia
and Schwertner
2008 ).
Menidini was described to include four genera: Cresphontes Stål, Antestia Stål
(currently Antestiini), Apines Dallas, and Menida Motschulsky (Atkinson
1888 ).
The defi nition of the tribe and relationship among the genera, however, are in need
of phylogenetic studies (Rider
1998 ). In the Neotropical Region, only two genera
occur from the 28 actually assigned to the tribe (Table
22.5 ). Keys to the identifi cation
of the species of the Neotropical genera, Rio Kirkaldy and Elanela Rolston are
available (Fortes and Grazia
2000 ; Grazia and Greve 2011 ) and Rio was revised
(Grazia and Fortes
1995 ).
The tribe Nezarini is represented by four genera in the Neotropical Region:
Chinavia , Chlorochroa Stål, Roferta Rolston, and Nezara Amyot & Serville. The
last one is represented only by N. viridula (L.), which has a cosmopolitan distribu-
tion (Ferrari et al.
2010 ). The group was proposed to include species of Nezara and
Acrosternum Fieber (Atkinson
1888 ). Cladistic analyses in the genus and group of
species level were published recently (Ferrari et al.
2010 ; Genevcius et al. 2012 ;
Genevcius and Schwertner
2014 ); however, the tribe lacks a formal defi nition, based
on phylogenetic studies. A pictorial key for the Brazilian species of Chinavia , which
include some secondary pests in South America, is presented in Schwertner and
Grazia (
2007 ).
Pentatomini is the most diverse tribe of Pentatominae. The original description
brings as the only character defi ning the group, the scutellum not covering the wings
or elytra (Leach
1815 ). No further defi nition was elaborated and the scattered phy-
logenetic evidences indicate that Pentatomini is not a monophyletic group (Gapud
1991 ; Hassan and Kitching 1993 ; Campos and Grazia 2006 ). Nonetheless, several
studies using cladistic methodology at genus level for Neotropical groups put some
light on to the relationships, especially among groups of species and genus level
1997 ; Fortes and Grazia 2005 ; Simões et al. 2012 ; Greve et al. 2013 ). A
phylogenetic analysis based in morphology and DNA sequences testing the
Chlorocoris group was recently developed (Greve, personal communication), and
the results derived from the total evidence analysis and the morphological analysis
alone are not in complete agreement. The differences among the two hypotheses of
relationship prevent the proposal of a unifi ed classifi cation for the taxa studied, not
supporting the group as a monophyletic clade.
In the Neotropical Region, Piezodorini is represented only by the genus
Piezodorus Fieber, with the species P. guildinii (Westwood) being a major pest of
soybean in South America (Panizzi et al.
2000 ). The original description was based
in species of Piezodorus and Ambiorix Stål and defi nes the tribe by “tibiae generally
rounded, rarely furnished above with a narrow and obsolete furrow, or fl at and emar-
ginate; venter spinose at the base, spine sometimes extended to the head; apical
angles of the sixth abdominal segment sometimes produced in a large acute tooth”
1888 ).
22 Stink Bugs (Pentatomidae)
The tribe Procleticini was described to include two monotypic genera: Lobepomis
Berg and Procleticus Berg (Pennington
1920 ). Currently it includes 11 genera
22.5 ) almost exclusively Neotropical (Rider 1994 ; Bernardes et al. 2009 ;
Schwertner and Grazia
2012 ). Only Brepholoxa and Dendrocoris have species
occurring in the Nearctic region (Rider
1994 ). The group is considered monophy-
letic and is supported by metasternum sulcate, ventral rim of pygophore with a
medial U-shaped emargination and with 1 + 1 process with different degrees of
development, and gonocoxites 8 smaller than laterotergites 9 (Schwertner and
2012 ).
Amyot and Serville (
1843 ) proposed the group Sciocorides to include genera
currently in different subfamilies of Pentatomidae. A more restricted concept of the
taxon was applied by Stål (
1876 ), who restricted the group to Dyroderes Spinola,
Mennacarus Amyot & Serville, Pododus Amyot & Serville, and Sciocoris Fállen.
The tribe includes currently 11 genera distributed in all biogeographic regions,
although poorly represented in the Neotropics, with only one genus (Table
22.5 ).
The group Strachiaries (Mulsant and Rey (
1866 )) included originally only the
genus Strachia Hahn. Stål (
1876 ) expanded the group to include Agaeus Dallas,
Bagrada Stål, Cinxia Stål, Eurydema Laporte, and Stenozygum Dallas. Currently
the tribe contains 14 genera worldwide, with only the genus Murgantia with repre-
sentatives in the Neotropical Region (Brailovsky and Barrera
1889 ).
Other six genera treated as unplaced in Rider (
2014 ) are registered in the
Neotropical Region ( Capivacius Distant, Cyptocephala Berg, Patanius Rolston,
Senectius Rolston, Tepa Tolston & McDonald, and Thyanta Stål). With the excep-
tion of Cyptocephala (4 spp.) and Thyanta (27 spp.), the remaining genera are
Key to the genera of Pentatominae of economic importance (modifi ed from Rolston
1974 and Rolston and McDonald 1984 ):
1. Third urosternite with medial process .................................................................. 2
1 . Third urosternite without process ....................................................................... 5
2. Specimens not longer than 9 mm, yellowish brown in color Piezodorus Fieber
2 . Specimens with at least 10 mm, green in life, or reddish brown or pale yellow in
dry preserved ....................................................................................................... 3
3. Mandibular plates surpassing clypeus in length; mandibular plates apex acute;
lateral margin of pronotum partially crenulated ..........................Arvelius Spinola
3 . Mandibular plates not surpassing clypeus, rounded at apex; lateral margin of
pronotum not ornamented .................................................................................. 4
4. Wide colored band in the margins of the head, pronotum, or basal third of
hemelytra; spiracle sometimes surrounded by yellow callus, without any
other macula .................................................................................. Chinavia Orian
4 ’. Margins of the head, pronotum, and basal third of hemelytra concolor with the
body or with a thin yellow band; spiracle without callus and green macula
may be present ............................................................ Nezara Amyot & Serville
5. Distal apex of the femora bearing a strong spine; specimens green in vivo
(can be reddish brown or pale yellow in dry preserved specimens)
...........................................................................................Loxa Amyot & Serville
J. Grazia et al.
5 . Distal apex of the femora unarmed, specimens usually brown or dark ..............6
6. Two rows of small tubercles present on inferior surface of femora
..................................................................................................... Agroecus Dallas
6 . Inferior surface of the femora lacking tubercles ................................................. 7
7. Small specimens, at most10 mm length, usually not reaching this length .......... 8
7 . Specimens length over 10 mm .......................................................................... 10
8. Specimens’ dorsum uniform brown to dark in color, without ornamentation
of different colors; insects shorter than 7 mm .......................... Glyphepomis Berg
8 . Pronotum and scutellum frequently with ivory callous; insects around 10 mm
in length .............................................................................................................. 9
9. Apex of rst rostral segment projecting past bucculae
..................................................................................Mormidea Amyot & Serville
9 . First rostral segment lying entirely between bucculae ......................Oebalus Stål
10. Mandibular plates conspicuously longer than clypeus, with apices usually
acute ........................................................................................Dichelops Spinola
10. Mandibular plates nearly equal to clypeus in length, sometimes a little longer or
shorter ............................................................................................................. 11
11. Peritreme long, evanescent ...............................................................Thyanta Stål
11. Peritreme short ................................................................................................ 12
12. First antennal segment reaching and sometimes slightly surpassing the apex
of the head; anterolateral margins of pronotum distinctly undulating,
denticulate, or vertically rugose ............................................... Euschistus Dallas
12. First antennal segment not attaining the apex of the head; anterolateral
margins entire ..................................................................................Tibraca Stål
22.6.5 Remarks
The Pentatominae is the most speciose subfamily within Pentatomidae and are rep-
resented in all geographic regions. New characters should be investigated and ana-
lyzed in all Pentatominae groups, in order to help in the clarifi cation of infra-subfamily
relations. Cladistics studies at genus and species levels in the subfamily are needed.
22.7 Subfamilies Podopinae and Stirotarsinae
22.7.1 Introduction
The subfamilies Podopinae and Stirotarsinae are represented by only one species
each in the Neotropical Region. The subfamily Podopinae is distributed worldwide;
most of its diversity can be found in the Afrotropical, Oriental, and Palearctic
regions (Davidová-Vilímová and Štys
1994 ; Rider 2014 ). The Stirotarsinae was
erected recently (Rider
2000 ) to include Stirotarsus abnormis Bergroth, a species
known only by a female specimen collected in Peru (Fig.
22.50a, b ).
22 Stink Bugs (Pentatomidae)
22.7.2 General Characteristics and Diagnosis
Characteristics of Podopinae include antenniferous tubercle visible in dorsal view;
scutellum usually elongated, surpassing 2/3 the abdominal length; lateral rim of
pygophore well developed, projecting on the ventral rim; ventral wall of pygophore
with a median depression; lateral lobes of the conjunctiva strongly sclerotized
1981 ; Gapud 1991 ; Gapon and Konstantinov 2006 ).
The subfamily Stirotarsinae can be recognized by the antennal segments I, II,
and V distinctly infl ated; rostrum three-segmented, without apparent segmentation
and anterior half enlarged; tarsi two-segmented, enlarged and with a longitudinal
Fig. 22.50 Stirotarsus abnormis Bergroth, dorsal view ( a ) and ventral view ( b ) (From Rider
2000 ), with permission of the Annals of Entomological Society of America)
J. Grazia et al.
carina; tibiae foliate, dorsally concave. Other diagnostic characteristics, like the
general body surface wrinkly, head longer than wide, sternum sulcated, and perit-
reme and evaporatorium reduced, are found in several groups of Pentatomidae and
Pentatomoidea (i.e., Cyrtocorinae and some Podopinae).
22.7.3 Classifi cation and Diversity
The Podopinae was fi rst recognized by Stål ( 1876 ) as a group of genera. Lethierry
and Severin (
1893 ) gave status of subfamily to the group. The current classifi cation
was established by Davidová-Vilímová and Štys (
1994 ), with fi ve tribes:
Brachycerocorini, Deroploini, Graphosomatini, Podopini, and Tarisini (Rider
2014 ). The monotypic genus Neoleprosoma was described by Kormilev and Pirán
1952 ) to include the Neotropical species N. argentinensis Kormilev & Pirán
22.51 ). The authors considered Neoleprosoma related to Leprosoma
Baerensprung and other similar genera of Podopinae from the Palearctic region,
although the classifi cation of the species was never reviewed. Davidová-Vilímová
and Štys (
1994 ) included Neoleprosoma in the tribe Graphosomatini. Distribution
records of N. argentinensis include Argentina (Buenos Aires, Córdoba, and Santa
Fé) and Uruguay (Artigas), in the southern region of South America.
Fig. 22.51 Neoleprosoma
argentinensis Kormilev &
Pirán (Photo by CF
22 Stink Bugs (Pentatomidae)
The monotypic Stirotarsus was proposed by Bergroth ( 1911 ) to include S. abnor-
mis from Peru, but because of its aberrant features, the classifi cation among
Pentatomidae was unsettled (Rider
2000 ). Based on the presence of a ductus recep-
taculi dilated and invaginated with three distinct walls in the female spermatheca
and the unique set of characters found in S. abnormis , Rider (
2000 ) proposed to
place the genus in a separate subfamily. Poorly represented in collections (only one
female), this taxon is probably endemic to the northern region of South America. No
biological information is available; however previous hypothesis placed this taxon
related to the asopines.
22.8 Concluding Remarks
The Pentatomidae, though predominantly herbivorous, includes predacious species
and some economically important species of Heteroptera, such as the green stink
bugs and the brown bugs. The group represents the second largest family of the
infraorder Pentatomomorpha, with nine subfamilies, about 850 genera, and 4.700
species worldwide. The Neotropical fauna treated in this chapter comprises about
230 genera and more than 1,400 species, in seven subfamilies, representing 23 % of
the world’s genera and about 33 % of the species, suggesting that the number of new
taxa awaiting description is immense. We hope this review of the family, which
includes information on hosts, habits, economic importance, classifi cation, check-
lists of species and/or genera, and keys to many of the Neotropical genera, will
provide information to stimulate new interest in this large and diverse group of
taxonomically and ecologically important insects.
Acknowledgments To David Rider (North Dakota State University, Fargo, USA) for allowing
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the group in this chapter. To Gerald Fauske (North Dakota State University, Fargo, USA), who
kindly took the pictures of the asopines from the NDSU collection.
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