(PDF) Stink Bugs (Pentatomidae)

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

e-mail: jocelia@ufrgs.br; carolinegreve@gmail.com; luiz.campos@ufrgs.br

A. R. Panizzi

Laboratório de Entomologia , Embrapa Trigo , Caixa Postal 3081 ,

Passo Fundo , RS 99001-970 , Brazil

e-mail: antonio.panizzi@embrapa.br

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

e-mail: schwertner@unifesp.br

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

e-mail: therezagarbelotto@hotmail.com

J. A. M. Fernandes

Instituto de Ciências Biológicas , Universidade Federal do Pará ,

Rua Augusto Corrêa n°1 , Belém , PA 66075-110 , Brazil

e-mail: joseamf@ufpa.br

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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

Subfamily/tribe

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

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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

Argentina.

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 ).

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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

Slater

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

Schwertner)

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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,

BOL, CU, JAM, CR, HON, VEN

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,

NIC, ECU,

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

(continued)

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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,

BR, BOL, NIC

Heteroscelis servillei Laporte BR, PER, BOL, ARG, PAR, VEN, GUY, FG,

SUR, TRI

Heteroscelis lepida (Stål) USA, MEX, GUA, PAN, CO, VEN, FG, ECU,

HON

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,

SUR, PAR

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,

PER, ARG, BR, FG, PAR

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

(continued)

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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,

CR, PAN, VEN, JAM, GRE, DRE, HAI, DOR,

PUR, TRI, Curacao, CU, BAH

Podisus nigrispinus (Dallas) BR, PAN, CR, BOL, ARG, PER, ECU, SUR,

COL, GUY, PAR

Podisus distinctus (Stål) ECU, BOL, COL, BR, VEN, FG, PAR, ARG,

PER

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 &

Serville)

BRA, SUR, BOL, PAR, URU, ARG, PER

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

(continued)

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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

(Figs.

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

(Figs.

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)

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6. Base of the abdomen with anteriorly directed tubercle or spinous process

(Fig.

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

7 ′. Rostral segment II shorter or subequal in length to III and IV combined ......... 8

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.

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Fig. 22.2 Stiretrus

anchorago (F.), dorsal view

(Courtesy of G. Fauske)

Fig. 22.3 Perillus bioculatus

(F.), dorsal view (Courtesy of

G. Fauske)

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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

(Thomas

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

Schwertner)

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Fig. 22.6 Supputius

cincticeps (Stål), ventral

view showing detail of the

evaporatorium (Photo by CF

Schwertner)

Fig. 22.7 Alcaeorrhynchus

grandis (Dallas), ventral view

showing detail of the

evaporatorium (Photo by CF

Schwertner)

Fig. 22.5 ( a ) Oplomus catena (Drury), dorsal view (Photo by CF Schwertner). ( b ) Oplomus

dichrous (Burmeister), dorsal view (Courtesy of G. Fauske)

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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)

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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)

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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

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Fig. 22.13 Apateticus

lineolatus (Herrich-

Schaeffer), head in dorsal

view (Photo by CF

Schwertner)

Fig. 22.14 Apateticus

lineolatus (Herrich-

Schaeffer), dorsal view

(Photo by CF Schwertner)

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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

Schwertner)

Fig. 22.16 Podisus

nigrispinus (Dallas), dorsal

view (Photo by CF

Schwertner)

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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)

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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.

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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

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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

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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

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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)

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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

ﬁ 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

ﬁ 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.

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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 ;

Rolston

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 )

(Rolston

1981 ).

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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

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(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

Albuquerque

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)

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Stictochilus

Marghita

Janeirona

Brochymena

Dalpada

Discocephala

Adoxoplatys

Neoadoxoplatys

Pseudadoxoplatys

Ocellatocoris

Orbatina

Coranda

Schraderiellus

Ochlerus b

Ochlerus a

Stalius trisinuatus

Parastalius

Moncus

Ochlerus handlirsch

i

Eritrachys

Phereclus

Herrichella

Paralincus

Lincus

Similiforstona

Tetrochlerus

Alitocoris a

Parochlerus

Alathetus

Alitocoris parvus

Barola

Catulona

Cromata

Schaefferella

Miopygium

Uvaldus

Brachelytron

Forstona

Clypona

Macropygium

Fig. 22.23 Phylogeny of Ochlerini depicting the relationship between genera; strict consensus

tree after successive weighting (Redrawn from Campos and Grazia (

2006 ) )

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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

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

literature)

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

Acanthocephalonotum

Petrulevičius & Popov

a

A. martinsnetoi Petrulevičius & Popov – ARG

a

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

(continued)

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(continued)

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;

PER; SUR; VEN;

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;

SUR; VEN

A. mimeticus Ruckes – BRA; PER

A. mixtus (F.) (type) – ARG; BOL; BRA; GUF; GUY;

SUR; TTO; VEN

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)

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Table 22.3 (continued)

Genera Species

A. tripterus (F.) – ARG; BOL; BRA; COL; ECU; PAN;

PRY; TTO; VEN

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;

GUF

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;

VEN

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;

SUR; VEN

D . ( P .) nigrodecoratus Becker & Grazia – BRA

D . ( P .) prolineatus Becker & Grazia – ARG; BRA; PRY;

URY

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

(continued)

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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

(continued)

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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;

GUY; PAN; PER; SUR; VEN

Priapismus Distant P. costaricensis Rolston – CRI

P. ecuadorensis Rolston – ECU

(continued)

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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

inquirenda

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

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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

(Becker

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

(Fig.

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

(Ruckes

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)

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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

Grazia

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

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)

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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

(continued)

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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;

PER; BRA

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

b

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

(continued)

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(continued)

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, &

Campos

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)

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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

(Fig.

22.19b ), and the later includes the remaining 20 genera known by that time

(Fig.

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,

BRA, ECU

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

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721

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)

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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

s

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 ) )

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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)

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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

Carvalho-Filho)

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Fig. 22.32 Edessa elaphus Breddin (Photo by F Carvalho-Filho)

Fig. 22.33 Edessa

dolichocera Burmeister,

lateral view (Photo by F

Carvalho-Filho)

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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.

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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)

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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

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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.

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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

Pentatomidae.

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

McDonald

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 ).

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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

Barros)

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Fig. 22.38 Euschistus

irroratus Bunde, Grazia, &

Mendonça Jr. (Photo by J

Grazia)

Fig. 22.39 Pellaea stictica

(Dallas) (Courtesy of

CSRibeiro-Costa)

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Fig. 22.40 Nezara viridula

(L.) (Photo by A Ferrari)

Fig. 22.41 Phalaecus sp.

(Photo by J Grazia)

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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

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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

Panizzi

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)

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Fig. 22.46 Oebalus

ypsilongriseus (De Geer)

(Courtesy of H. Prando)

Fig. 22.47 Tibraca

limbativentris Stål (Courtesy

of H. Prando)

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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)

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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

literature)

Carpocorini

Acledra Signoret BRA, CHI, ARG, URU

Agroecus Dallas PER, ECU, BOL, GUY, FGU, COL, VEZ, BRA, PER, BOL,

ARG, PAR

Amauromelpia Fernandes &

Grazia

CR, FGU, BRA, BOL, COL, PER

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,

URU

Epipedus Spinola BRA

Euschistus Dallas USA, MEX, NIC, GTM, ELS, HON, CR, PAN, COL, VEZ,

W.I. (ALL), GUY, BRA, ECU, BOL, FGU, PER, CUB, JAM,

CURAÇAO, BLZ, ARG, URU,

Galedanta Amyot &

Serville

CR, GUY, COL, BRA

Glyphepomis Berg BRA, ARG, URU

Hypanthracos Grazia &

Campos

BRA, URU

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

BRA

Mathiolus Distant GTM

Mecocephala Dallas BRA, ARG, URU

Menudo Thomas PUR

Mormidea Amyot & Serville CAN, USA, MEX, NIC, PUR, CUB, HIS, DRE, COL, GUY,

FGU, PER, BOL, SUR, BRA, ECU, TRI, PAN, ARG, URU,

VEZ, JAM, PAR, GRE, TRI

(continued)

J. Grazia et al.

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Table 22.5 (continued)

Carpocorini

Oebalus Stål USA, MEX, W.I. (ALL), SUR, BRA, PER, PAR, ARG, URU,

TRI, COL, GUY, ECU, BOL, HIS, PUR, CUB

Oenopiella Bergroth BRA, CHI, ARG, URU

Ogmocoris Mayr BRA

Padaeus Stål USA, MEX, GTM, NIC, HON, CR, ECU, COL?

Parahypatropis Grazia &

Fernandes

BRA, ARG

Paramecocephala Benvegnú SUR, GUY, GUF, BRA, ECU, ARG, URU

Paratibraca Campos &

Grazia

PAN, TTO, SUR, COL, BRA, PER, BOL

Parentheca Berg BRA, PAR, ARG, URU

Pedinonotus Fernandes &

Grazia

BRA

Pentatomiana Grazia &

Barcellos

BRA (RJ, SP, SC)

Poriptus Stål TTO, COL, BRA, BOL, ARG

Prionotocoris Kormilev BOL, ARG

Proxys Spinola USA, MEX, DRE, CUB, GTM, NIC, FGU, BLZ, CR, GRE,

PAN, COL, VEZ, BRA, ARG, URU

Sibaria Stål MEX, ALL CENTRAL AM., COL, VEZ, FGU, GUY, SUR,

TTO, BRA, ECU, PER, BOL, PAR, ARG

Spinalanx Rolston & Rider COL, PER, ECU, BOL

Stysiana Grazia, Fernandes

& Schwertner

COL, SUR, TTO, GUY, BRA, PER, ARG, PAR

Tibraca Stål DRE, CUB, CR, COL, VEZ, BRA, PER, BOL, ARG, URU

Trichopepla Stål CAN, USA, MEX

Catacanthini

Arocera Spinola MEX, BLZ, HON, CR, PAN, CUB, JAM, PUR, DRE, TTO,

VEZ, FGU, GTM, CR, COL, GUY, SUR, FGU, BRA, ECU,

BRA, BOL, ARG

Boea Walker CR, BRA

Rhyssocephala Rider MEX, BLZ, GTM, CR, PAN, JAM, COL, VEZ, ECU, BRA,

PER, BOL

Runibia Stål USA, JAM, Virgen Is, COL, VEZ, ECU, BRA, PER, BOL,

GUF, ARG, PAR, URU

Vulsirea Spinola USA, MEX, HON, HAI, JAM, CR, CUB, PAN, COL, VEZ,

BRA, BOL

Halyini

Brochymena Amyot &

Serville

USA, MEX, HON, CR, BAH, CUB

Parabrochymena Larivière USA, MEX, GTM, CR

Mecideini

Mecidea Dallas USA, MEX, NIC, HON, CR, PUR, ANTIGUA, ARG

(continued)

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740

Table 22.5 (continued)

Carpocorini

Menidini

Elanela Rolston SUR, BRA, PER

Rio Kirkaldy MEX, GTM, ELS, CR, PAN, VEZ, SUR, BRA, ARG

Nezarini

Chinavia Orian CAN, USA, MEX, BLZ, CR, CUB, DRE, PUR, JAM, Grenada,

Guadeloupe, Dominica, St. Croix, VIR IS., GTM, NIC, HAI,

HON, BAH, COL, VEZ, SUR, FGU, GUY, BRA, ECU, PER,

BOL, CHI, PAR, ARG, URU

Chlorochroa Stål CAN, USA, MEX

Nezara Amyot & Serville MEX, NIC, HON, CUB, DRE, PAN, VEZ, BRA, CHI, ARG,

URU

Roferta Rolston BLZ, PUR, PAN, TRI, CUB, BRA, ARG

Pentatomini

Adevoplitus Grazia &

Becker

MEX, GTM, CR, PAN, VEZ

Arvelius Spinola DRE, GUP, MAR, BAR, SVI, USA, MEX, ELS, GTM, HON,

CUB, DRE, PUR, VIR, ANT, BAH, NIC, JAM, CR, PAN,

COL, VEZ, SUR, GUY, TTO, HAI, ECU, BRA, BOL, PAR,

ARG, URU, GUY

Banasa Stål USA, MEX, ALL CENTRAL AM., ALL W.I., COL, ECU,

VEZ, PER, BOL, BRA, PAR, SUR, GUY, GUF, ARG, URU,

Brasilania Jensen-Haarup BRA

Chlorocoris Spinola USA, MEX, BLZ, NIC, HON, GTM, JAM, CR, PAN, COL,

VEZ, SUR, TRI, ECU, BRA, PER, BOL, PAR, ARG

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

Lopadusa Stål PAN, VEZ, GUY, BRA, ECU, PER, BOL, PAR, ARG

Loxa Amyot & Serville USA, MEX, CUB, DRE, BAH, PUR, DOMINICA,

GUADALOUPE, CURAÇAO, GRENADINIES, ST. LUCIA,

JAM, HON, NIC, PAN, COL, GUY, VEZ, SUR, FGU, BRA,

PER, ARG

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

(continued)

J. Grazia et al.

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Table 22.5 (continued)

Carpocorini

Neopharnus Van Duzee EUA, CUB

Neotibilis Grazia &

Barcellos

MEX, CR, PAN, COL, VEZ, GUY, GUF, BRA, ECU, PER,

BOL

Nocheta Rolston SUR, BRA

Pallantia Stål MEX, GTM, PAN, VEZ, PER, BRA, PAR, ARG

Paratibilis Ruckes MEX, PER

Pellaea Stål USA, MEX, NIC, HON, CR, PAN, COL, VEZ, GUY, BRA,

ECU, ARG, URU

Phalaecus Stål GUY, SUR, GUF, BRA, COL

Pharnus Stål CUB

Pharypia Stål MEX, HON, GTM, CR, PAN, COL, VEZ, FGU, BRA, ARG

Placocoris Mayr BRA, ARG

Platistocoris Rider BRA

Praepharnus Barber &

Bruner

CUB

Pseudevoplitus Ruckes VEZ, BRA, PER, BOL, ARG

Ramosiana Kormilev MEX, CR, PAN, PER, BOL

Rhyncholepta Bergroth PAN, VEZ, GUF, VEZ, BRA, BOL

Rideriana Grazia &

Frey-da-Silva

BRA, PER, BOL

Serdia Stål CR, PAN, VEZ, BRA, PER, BOL, PAR, ARG

Similliserdia Fortes &

Grazia

BRA

Stictochilus Bergroth ARG, BRA

Taurocerus Amyot &

Serville

MEX, GTM, CR, PAN, COL, GUY, BRA, PER, BOL

Tibilis Stål BRA, ECU, PER, BOL

Vidada Rolston PER

Piezodorini

Piezodorus Fieber USA, TRI, PUR, GTM, HON, NIC, JAM, CUB, DRE, St.

Vincent, Grenada, PAN, VEZ, BRA, ECU, PAR, ARG

Procleticini

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

(continued)

22 Stink Bugs (Pentatomidae)

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Table 22.5 (continued)

Carpocorini

Strachiini

Murgantia Stål USA, MEX, GTM, ANT, CR, PAN, BRA, PER, BOL

Sciocorini

Trincavellius Distant ECU, CHI, PER

Unplaced

Capivaccius Distant MEX, HON, PAN, BRA, ARG

Cyptocephala Berg USA, CENTRAL AM. (ALL), BAH, CUB, DRE, JAM, VIR

IS., DRE, PUR, BRA, PER, BOL, ARG, URU

Patanius Rolston BRA

Senectius Rolston PER

Tepa Rolston & McDonald USA, CURAÇAO

Thyanta Stål USA, MEX, GTM, BLZ, ELS, HON, NIC, CR, PAN, BAH,

CUB, JAM, DRE, HAI, PUR, VIR IS., BARBADO, BRITISH

W.I., FRENCH W.I., CURAÇAO, TTO, VEZ, COL, SUR,

FGU, BRA, PER, ECU, CHI, BOL, PAR, ARG, URU

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

(Table

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.

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743

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

(Grazia

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”

(Atkinson

1888 ).

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744

The tribe Procleticini was described to include two monotypic genera: Lobepomis

Berg and Procleticus Berg (Pennington

1920 ). Currently it includes 11 genera

(Table

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

Grazia

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

monotypic.

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.

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745

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 ﬁ 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 ).

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746

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

(Schaefer

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.

joseamf@ufpa.br

747

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

(Fig.

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

Schwertner)

22 Stink Bugs (Pentatomidae)

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748

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

access to the collection at the NDSU and providing most of the Asopinae specimens that illustrate

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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Stink Bugs (Pentatomidae)

Abstract and Figures