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Review of the neotropical genus Aleixus McDonald (Hemiptera: heteroptera: pentatomidae: procleticini), with description of a new species and cladistic analysis of the tribe procleticini.

Abstract--The genus Aleixus McDonald is reviewed and a new species described. The genus was monotypic and known from a single female from Brazilian Amazonia. The new species Aleixus toby schuhi described from southern Brazil, greatly extends the distribution of A leixus. A cladistic analysis, including 30 morphological characters and 18 taxa, was performed using TNT with the implicit enumeration algorithm, with three weighting schemes: equal (EW), successive approximation (SW), and implied weighting (IW). Analysis under EW resulted in four cladograms (L = 74, CI = 60, RI = 78). Analyses under SW and IW resulted in one most parsimonious cladogram, with fit varying from 20.4 (K = 0.64) to 26.6 (K = 5.80). The cladogram obtained under differential weighting schemes is included in the set of cladograms found under EW. The monophyly of the tribe Procleticini and genus Aleixus were recovered in all analyses. Non-ambiguous synapomorphies for Aleixus include antennal segment II longer than antennal segment I, and humeral angles developed in spatular projections, dorsally bifid. Non-ambiguous synapomorphies of Procleticini are metasternum sulcate, ventral rim of pygophore with a medial U-shaped emargination and with 1 + 1 process with different degree of development, and gonocoxites 8 smaller than laterotergites 9. A hypothesis of relationships among Procleticini genera was possible only under IW analysis. Two main clades were recognized: clade A, including the genera Brepholoxa Van Duzee, Dendrocoris Bergroth, Procleticus Berg, and Thoreyella Spinola; and clade B, including the genera Aleixus, Odmalea Bergroth, Parodmalea Rider, and Pseudobebaeus Distant.

Key words: stink bugs, morphology, Neotropical region, Heteroptera.

INTRODUCTION

The genus Aleixus was described from a female specimen collected in northern Brazil (Rolston and McDonald, 1981). It was included in the Section 2 of the tribe Pentatomini by having a free median tubercle or spine at the base of third urosternite, without the posterior margin of the metasternum produced in apposition to the apex of that tubercle or spine (Rolston et al., 1980; Rolston and McDonald, 1981). Diagnostic characteristics of the genus were the first antennal segment not surpassing apex of head; juga longer than tylus, separated apically; bucculae lobed posteriorly; humeri bearing large dorsal tubercle; peritreme extending more than half the length of the evaporatorium; femora armed with a small spine; and gonocoxites 8 as 1 + 1 small subtriangular sclerites, and obscured under the urosternite 7 (Rolston and McDonald, 1981). Among the genera included in Section 2, bucculae lobed posteriorly was also shared with Brepholoxa Van Duzee, Dendrocoris Bergroth, Odmalea Bergroth, Zorcadium Bergroth (= Pseudobebaeus Distant), Thoreyella Bergroth and Rio Kirkaldy (Rolston and McDonald, 1981).

In the revision of the genus Thoreyella, Rolston (1984) identified these seven American genera of Pentatomini based in abdominal spine projecting beneath the metasternum and bucculae extending as lobes near to or past the distal end of the first rostral segment. The genus Aleixus was diagnosed by the humeral angles bearing a large dorsal tubercle and second antennal segment longer than each succeeding segment (Rolston and McDonald, 1981; Rolston, 1984). Rider (1994) transferred Aleixus and five of above genera to the tribe Procleticini (excluding Rio) because of the typical morphology of male and female genitalia. Among the genera included in Procleticini, Aleixus can be recognized by the scutellum subtriangular, not extending beyond the apices of coria, plus the characters defined by Rolston and McDonald (1981).

The tribe Procleticini was proposed by Pennington (1920) to include the monotypic genera Lobepomis Berg and Procleticus Berg, and diagnosed the tribe by the developed humeral angle, scutellum large and nearly reaching the apex of the abdomen, abdomen very convex below, and rostrum not surpassing hind coxae (Pennington, 1920; Rider, 1994). Kormilev (1955) included in the tribe Neoderoploa Pennington, and Piran (1963) described Terania in Procleticini, both genera sharing the characteristics defined by Pennington for the tribe. Rider (1994) broadened the definition of the tribe to include seven genera, six of them previously placed in the Section 2 of Pentatomini and one new genus (Rolston and McDonald, 1981; Rolston, 1984; Rider and Fischer, 1998): Aleixus McDonald, Brepholoxa VanDuzee, Dendrocoris Bergroth, Odmalea Bergroth, Parodmalea Rider, Pseudobebaeus Distant (as Zorcadium Bergroth), and Thoreyella Spinola. Rider (1994) also defined new unique diagnostic characters for the tribe, all from the morphology of male and female genitalia, which support the hypothesis of the monophyly of the tribe (Rider, 1994; Bernardes et al., 2009).

However, there was no published study with an explicit test of the monophyly of the tribe, and the phylogenetic relationships of the included genera is only partially known (Bernardes et al., 2009). Because the description of Aleixus was based only on one female specimen, any inference about the phylogenetic position of the genus among the Procleticini is difficult. Herein, this present paper provides: 1) the description of a new species of Aleixus, based on specimens of both sexes; 2) the review of the diagnostic characters of the genus; 3) a hypothesis of phylogenetic position of the genus Aleixus among Procleticini, based on the first cladistic analysis of the tribe using morphological characters. The monophyly of the genera included and the position of the tribe among the subfamily Pentatominae are also discussed.

MATERIAL AND METHODS

The description of the new species was based on l0 specimens from southern Brazil. The studied specimens and deposition of types correspond to the following collections (acronyms are according to Evenhuis, 2012): AMNH--The American Museum of Natural History, New York, USA; MAPA--Museu Anchieta, Porto Alegre, Rio Grande do Sul, Brazil; MCNZ--Museu de Ciencias Naturais da Fundacao Zoobotanica do Rio Grande do Sul, RS, Brazil; MZSP--Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, SP, Brazil; NMNH--The National Museum of Natural History, Washington D.C., USA; UFRS--Universidade Federal do Rio Grande do Sul, Departamento de Zoologia, Porto Alegre, RS, Brazil. The type (and only known) specimen of Aleixus virgatus McDonald, deposited in the NMNH, was studied.

Eighteen terminal-taxa and 30 morphological characters were used in the cladistic analysis (Tables 1 and 2). The ingroup includes species from most of the genera of Procleticini, representing the morphological diversity within the tribe (Rider, 1994; Bernardes et al., 2009, 2011). The monotypic genera Lobepomis, Neoderoploa and Terania were excluded from the analysis because they are part of a well-corroborated monophyletic group with the monotypic genus Procleticus, which is the sister-group of the genus Thoreyella (Bernardes et al., 2009). The outgroup includes species from six genera of different tribes of Pentatominae (following Rider, 2012): Arocera acroleuca (Perty) [Catacanthini], Banasa induta Stal [Pentatomini], Euschistus heros (Fabricius) [Carpocorini], Piezodorus guildinii (Westwood) [Piezodorini], Rio pectoralis (Stal) [Menidini] and Thyanta perditor (Fabricius) [Unplaced]. These exemplar-taxa represent genera of Pentatominae that were, at some point and by different authors (i.e., Rolston, 1978; Rolston and McDonald, 1981; Rolston, 1984; Gapon, 2005), considered related to all or part of the genera currently included in Procleticini. The internal genitalia of males and females were studied for all terminal taxa except for Aleixus virgatus, known only by the female holotype (Rolston and McDonald, 1981).

The cladistic analyses were conducted using the parsimony program TNT (Goloboff et al., 2008), under three weighting schemes: equal (EW), successive approximation (SW) and implied weighting (IW) (Farris, 1969; Goloboff, 1993). Search strategies for cladograms used the implicit enumeration algorithm of TNT (branch-andbound). For the definition of the K values in the implied weighting analysis, we followed Mirande (2009); the scripts aaa.run and aab.run provided by the author were adapted to implicit enumeration searches. All characters were considered nonadditive (Fitch, 1971). The resulting trees were rooted between E. heros and the remaining terminal taxa. Cladograms were visualized and printed using Winclada (Nixon, 2002). In the discussion section, characters were cited following the notation "[X.sup.y]'', in which X represents the character and Y represents the state.

Measurements (mean, minimum and maximum) are given in millimeters. In the text, the proportions of the segments were represented with the symbols: "<" minor, ">" greater, "[approximately equal to]" subequal. The terminology of Kment and Vilimova (2010) was adopted for the structure of metatoracical glands; Dupuis (1970) and Schaefer (1977) for the structure of the genitalia. The detailed description of the genitalia followed Bernardes et al. (2009, 2011).

TAXONOMY

ALEIXUS McDONALD, 1981

Aleixus McDonald, in Rolston and McDonald, 1981: 259-260.

TYPE-SPECIES: Aleixus virgatus McDonald, by original designation.

DIAGNOSIS: The addition of a new species in Aleixus requires a rearrangement in the generic diagnosis as given by Rolston and McDonald (1981) and Rider (1994). Head: distinctly declivent at apex; antennal segment I not reaching apex of head; antennal segment II and III cylindrical, segments IV and V somewhat inflated; antennal segment II longer than any other segment; juga surpassing clypeus, obtuse at apex, not contiguous anteriorly. Rostrum reaching metacoxae. Thorax: Pronotum with humeral angles with large tubercles, dorsally bifid and black at apices; tubercles ventral surface bearing a short blunt projection. Scutellum subtriangular, apex broadly rounded not reaching apices of coria. Thoracic sterna nearly flat. Peritreme acuminate apically, extending at least beyond middle of evaporatorium. Legs: with dorsal surface of femora produced distally. Abdomen: Abdominal spine small, not reaching between metacoxae. Characters used as diagnostic to the genus and found to be variable between the two species include length of the humeral angles, development of the tooth placed distally in the femora, length of the peritrema, and shape of tibia (see also comments in the description of the new species).

Aleixus virgatus McDonald, 1981 Fig. 1

Aleixus virgatus in Rolston and McDonald, 1981: 260-262; Rider, 1994: 213-217.

SPECIMENS EXAMINED: HOLOTYPE: Female: Km 8 Est.[rada] do Aleixo, Manaus AM. BV. 26-VI-76. (Type no. 72138 NMNH).

Aleixus toby schuhi, new species Figures 2-5

HOLOTYPE: Male: BRAZIL, Rio Grande do Sul, Canela, Floresta Nacional de Canela, IBDF, 13.VIII.1982, F. Pires leg. Deposited at Museu de Ciencias Naturais, Fundacao Zoobotanica do Rio Grande do Sul, Porto Alegre (MCNZ).

DIAGNOSIS: Coloration of the body without dark brown longitudinal stripes as seen in A. virgatus (Fig. 1); each juga obtuse at apex; rostrum scarcely reaching metacoxae; humeral angles developed into wide and long tubercles.

DESCRIPTION: COLORATION: Dorsal surface yellowish matte to pale brown, with dark brown to black punctures; darker punctures concentrated on anterior and sometimes posterior third of pronotum and base of head. Five patches of dark brown punctures on scutellum and two on hemelytra somewhat delimiting an immaculate area at apex of radial vein (Fig. 2). Connexival segments with one patch of dark brown punctures at middle. Ventral surface yellowish matte with darker and rougher punctures than dorsal side; evaporatorium on meso and metapleuron darkened. Urosternite Vil with a large dark brown to black spot at middle in both sexes. Head: Juga scarcely surpassing clypeus, obtuse at apex. Antenniferous tubercles clearly visible in dorsal view. Antennal segment I almost reaching apex of head; proportion of antennal segments I < II > III < IV [approximately equal to] V. Rostrum scarcely reaching metacoxae. Pronotum: Anterolateral margins concave forming an obtuse angle at middle. Humeral angles developed in wide and long tubercles, bifid and black at apices, ventral surface with a short blunt subapical projection (Fig. 3). Thorax: Scutellum surpassing the middle of connexival segment V but not attaining apex of coria. Peritreme extending 1/3 the distance from ostiole mesial margin to metapleura lateral margin (Fig. 3). Superior surface of femora convex apically (Fig. 3). Fore tibiae entirely cylindrical, II and III scarcely sulcate or flat near apex. Connexivum with posterolateral angles with blunt projections. Abdomen: A pair of trichobothria placed mesially to the line of spiracles. Abdominal spine small, not reaching metacoxae.

Male: Genitalia (Fig. 4): Pygophore almost quadrangular. Posterolateral angles somewhat concave. Ventral rim of pygophore with the medial U-shaped emargination relatively shallow (vre), 1 + 1 flaps slightly bending dorsad into the genital cup ([approximately equal to] Brepholoxa heidemanni Van Duzee); inferior folder of ventral rim developed in a medial bilobed projection (vrp), apparently an autapomorphy of Aleixus. Segment X (x) wider than longer, with flat dorsal surface. Paramere (pa) short stocky, unilobate; base of paramere with an obtuse projection covered with bristles. The segment X and the parameres are placed perpendicularly to the pygophore axis. Superior process of dorsal rim (pdr) well developed, somewhat conical, covering the paramere in dorsal view, obscured by the dorsal rim. Basal plate (bp) of the articulatory apparatus modest developed, about 1/4 of phallotheca length, dorsal connectives (dc) short and processus capitati (pc) almost reaching the middle of phallotheca. Processus phallothecae (pph) present. Conjunctiva with 1 + 1 assymetrical membranous lobes obscuring the slightly sinuous vesica.

Male. Measurements (N = 3): Total length: 5.9 (5.74-6.06); Abdominal width: 4.1 (4.01-4.18); Head length: 1.2 (1.14-1.23). Head width: 1.48 (1.47-1.51); Anteocular length: 0.57; Interocular width: 0.92 (0.90-0.94); Length of antennal segments: I 0.32, II 0.60 (0.57-0.65), III 0.35 (0.32-0.41), IV 0.54 (0.49-0.57), V 0.53 (0.490.57); Pronotum length: 1.53 (1.47-1.64). Pronotum: width 5.70 (5.65-5.82); Scutellum length: 2.48 (2.46-2.54); Scutellum width: 2.54 (2.46-2.62).

Female: Genitalia (Fig. 5): Laterotergites 8 (1a8) very poorly delimited, apparently forming an entire band ventrally, lacking spiracles. Laterotergites 9 (la9) well developed; oblong, not extending over laterotergites 8 and separating the laterotergites 8 of the reduced gonocoxites 8 (gc8). Gonocoxites 8 with posterior margins convex, sutural margins not juxtaposed leaving uncovered most of the sclerotized gonapophyses 8 (g8). Gonocoxites 9 (gc9) reduced, with almost 1/4 the length of segment X (x), anterior and posterior margins concave. Chitinellipsen (ch) present. Ductus receptaculi (dr), before vesicular area (va), almost three times the length of ductus after vesicular area. Pars intermedialis (pi) almost as long as capsula seminalis (cs), the latter semiglobose with a digitiform projection.

MEASUREMENTS. (N = 5). Total length: 6.44 (5.74-7.05). Abdominal width: 4.49 (4.34-4.45). Head length: 1.26 (1.23-1.31). Head width: 1.59 -(1.55-1.64). Anteocular length: 0.63 (0.57-0.73). Interocular width: 0.99 (0.94-1.06). Length of antennal segments: I 0.32, II 0.64 (0.57 0.73), III 0.39 (0.32-0.49), IV 0.54 (0.49-0.65), V 0.52 (0.490.57). Pronotum length: 1.82 (1.64-2.05). Pronotum width: 6.24 (5.98-6.64). Scutellum length: 2.76 (2.54-2.87). Scutellum width: 2.87 (2.70-3.28).

COMMENTS: Rider (1994, p. 213) mentioned that A. virgatus could be recognized by the following generic characters: juga surpassing tylus, but not contiguous anteriorly; lateral jugal margins distinctly sinuous, slightly reflexed; apex of head distinctly declivent when viewed laterally; antenniferous tubercles clearly visible in dorsal view; antennal segment II and III cylindrical segments IV and V somewhat inflated; antennal segment II longer than any other segment; scutellum subtriangular, apex broadly rounded not reaching apices of coria; rostrum reaching metacoxae; each peritreme acuminate apically; basal abdominal spine small. All of these characters are also found in A. tobyschuhi. The humeral angles developed into wide and long tubercles, the peritreme not reaching the middle of metapleura, the apices of femora only convexly projected, and tibia of II and III legs sulcate near apex distinguish A. tobyschuhi from A. virgatus.

ETYMOLOGY: This species is dedicated to Dr. Randall T. Schuh in recognition to his invaluable contributions to the systematic of the heteropterous insects as well as to his permanent and invaluable guidance to the authors.

TYPE LOCALITY: Floresta Nacional de Canela, Canela, Rio Grande do Sul, Brazil.

OTHER SPECIMENS EXAMINED; PARATYPES: BRAZIL, Santa Catarina, Rio Vermelho, lack abdomen, X. 1958, Dirings (MZSP); Rio Grande do Sul, Vila Oliva, 24.X.1957, Pe. Buck leg., 3 [female], 1 [male] (MAPA); 21.II.1954, Pe. Buck leg., l[male] (AMNH); 28.II.1954, Pe. Pio Buck leg, l[female] (AMNH); Porto Alegre, 2 [female] (UFRS).

CLADISTIC ANALYSIS

RESULTS: The analysis under EW resulted in four most parsimonious cladograms, with 74 steps, a consistency index of 60, and a retention index of 78. The strict consensus of these cladograms (Fig. 6) supports the monophyly of the genus Aleixus and of the tribe Procleticini. Analyses under SW and IW (K varying from 0.64 to 5.8) resulted in one most parsimonious cladogram (Fig. 7), which is equal to one of the four most parsimonious cladograms obtained under EW. Under the IW analysis, the fit of the most parsimonious cladogram ranged from 20.4 (K = 0.64) to 26.6 (K = 5.80).

All analyses support the monophyly of the genus Aleixus and the tribe Procleticini (Figs. 6, 7), but the position of the genus Aleixus is defined only under analysis with differential weighting (Fig. 7). A clade including Proeleticus and Thoreyella, is also supported in all analyses. The phylogenetic position of the remaining genera is unresolved under EW. In the cladograms obtained under differential weighting schemes, two main clades are recognized. The genus Aleixus is included in a clade with Odmalea, Parodmalea, and Pseudobebaeus. The monophyletic condition of genera Dendrocoris and Odmalea is not supported by our results.

DISCUSSION. Two unique derived characters included in the analysis are shared by both species of Aleixus, one non-homoplastic and one homoplastic (Fig. 7). The humeral angles developed into large tubercles directed upward and dorsally bifid ([11.sup.2]) are unique to Aleixus, and easily allow its recognition among Procleticini (Rolston and McDonald, 1981; Rolston, 1984; Rider, 1994). The shape of the humeral angles is highly variable within the tribe, but is useful in the recognition of other genera as well (Rider, 1994; Bernardes et al., 2009). The length of antennal segment II longer than antennal segment I ([4.sup.1]), an homoplastic character that supports the monophyly of the genus, is shared with Brepholoxa within the ingroup (Fig. 7).

The original description of Aleixus was based in one female (Rolston and McDonald, 1981) and the relationship with at least some of the Procleticini genera was discussed (Rolston and McDonald, 1981; Rolston, 1984; Rider, 1994). The inclusion of Aleixus in the tribe was justified by gonocoxites 8 small and partially obscured by the last abdominal segment, a characteristic found in all genera within the tribe (Rider, 1994). The description of Aleixus tobyschuhi allows the study of the male genitalia of the genus, and confirms another characteristic proposed by Rider (1994) for Procleticini (ventral wall produced posteriorly and with a distinct medial emargination). In our analyses, these characters appear as non-homoplastic for all species of Procleticini ([22.sup.1] and [29.sup.1]), and together with the presence of processus phalothecae ([24.sup.1]), support the monophyletic condition of tribe (node 1).

Rider (1994) recognized three groups among Procleticini, based in the length of the scutellum. The genus Aleixus was grouped with Brepholoxq Dendrocoris, Odmalea, Parodmalea and Pseudobebaeus, based on all having a short scutellum. Bernardes et al. (2009) found that the elongation of the scutellum among Procleticini is a syapomorphy for the Procleticus group + Thoreyella, which is confirmed in our analyses (Fig. 7, node 8). The presence of short scutellum is symplesiomorphic among Procleticini, and does not support the grouping proposed by Rider (1994).

Our results suggest the genus Aleixus should be included in a monophyletic group that shares three non-ambiguous derived characters (Fig. 7, node 2). The flat dorsal surface of segment X in males ([27.sup.1]) is characteristic to this clade. The pronotum with anterolateral margins not emarginate ([10.sup.1]) and presence of a spine at the apex of the femur ([18.sup.1]) also support the clade, although these characteristics are also shared with Procleticus and Thoreyella (Fig. 7, node 8). The mesosternum flat ([15.sup.2]) is also a homoplastic character that allows the recognition of the members of this clade (see Table 2); however, it is ambiguous concerning the optimization in the cladogram.

Although beyond the scope of this work, the results of the cladistic analysis raised other interesting points that can direct further studies in the systematics of Procleticini. For instance, our analysis did not find any unique derived characters supporting genera Dendrocoris and Odmalea. Shape of the ventral rim laterad to the medial emargination in broadly spatulate processes are shared by all species of the genera Dendrocoris, Procleticus, and Thoreyella (node 6 in Fig. 7), and support the relationship among these genera (Bernardes et al., 2009). Our results also support the relationship among Odmalea and the monotypic genera Parodmalea and Pseudobebaeus (Fig. 7, node 4) by three unique derived characters. The diagnostic characteristics used to define Dendrocoris and Odmalea (i.e., Rolston, 1978; Rider, 1994; Thomas and Brailovsky, 1999) are also present in other Procleticini species. Revisionary studies of these taxa, taking into account the totality of their morphological diversity, will be crucial to defining their identity and their phylogenetic relationships among Procleticini. Particulady for Odmalea, several new species are awaiting a formal description (Rider, 2012; Schwertner and Grazia, personal observation).

Relationships of the genera currently in the tribe have been addressed by some authors in early descriptions (i.e., Bergroth, 1914; Bergroth, 1918; Pennington, 1920; Rolston and McDonald, 1981). However, only after Rider (1994) was the tribe Procleticini definitely identified as a valid group within the subfamily Pentatominae. Relationships with other tribes of Pentatominae were recently discussed. Studying the morphology of the phallus, Gapon (2005) compared Dendrocoris contaminatus (Procleticini) with genera included in other tribes of the subfamily Pentatominae (named Antestini and Aulacentrini in the study) and one genus of the subfamily Podopinae (tribe Bolbocorini). The author suggested a relationship of Dendrocoris with species of the genera Arocera and Thyanta, and to test this hypothesis, we included in our analysis the type-species of both genera.

Based on a number of characters, our results suggest the relationship of the tribe Procleticini with species of Menidini (Rio pectoralis) and Piezodorini (Piezodorus guildinii), mainly based in the shape of bucculae ([7.sup.1] and [8.sup.1]) and the length of rostrum ([9.sup.1] and [9.sup.2]). The weakly developed bucculae, with both anterior and posterior margins low and evanescent, and the elongated rostrum are widespread within Pentatominae, and resulted in plesiomorphic conditions in our analyses. Characters of genitalia do not appear informative under this level of the analysis, although this interpretation may be influenced by difficulty in interpreting them. In addition, the results found here are only preliminary, and a more comprehensive study focusing in the relationship of the tribes within Pentatominae will be necessary to better understand the phylogenetic position of the Procleticini.

ACKNOWLEDGEMENTS

To the curators of the listed collections for the loan of the material studied, especially Thomas J. Henry (NMNH) for providing access and facilities to the study of the holotype of A. virgatus. This work was supported by grants from Conselho Nacional de Pesquisa (CNPq) and Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) to the authors. We also thank two anonymous referees for providing constructive comments and suggestions that improved the manuscript.

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CRISTIANO F. SCHWERTNER [1] AND JOCELIA GRAZIA [2]

[1] Departamento de Ciencias Biologicas and Programa de Pos-graduacao em Ecologia & Evolucao, Instituto de Ciencias Ambientais, Quimicas e Farmaceuticas, Universidade Federal de Sao Paulo, Diadema, SP, Brazil, email: schwertner@unifesp.br

[2] Departamento de Zoologia and Programa de Pos-graduacao em Biologia Animal, Instituto de Biociencias, Universidade Federal do Rio Grande do Sul, Porto Alegre, RS, Brazil, email: jocelia@ufrgs.br

Table 1. Descriptions of the characters and states.

1. Juga contiguous anteriorly: 0) not contiguous; 1) contiguous.

2. Shape of jugae: 0) not spatulate; 1) spatulate.

3. Length of the antennal segment II in relation to the antennal
segment I in males: 0) longer; 1) shorter.

4. Length of the antennal segment II in relation to the antennal
segment III: 0) shorter; 1) longer; 2) subequal.

5. Length of the antennal segment IV in relation to the antennal
segment V: 0) subequal; 1) shorter.

6. Density of hairs in the antennal segment III in relation to the
antennal segment IV and V: 0) lesser; 1) equal.

7. Shape of the anterior teeth of bucculae: 0) in 1 + 1 rounded
lobes; 1) in 1 + 1 acute spines.

8. Bucculae lobed posteriorly: 0) absent; 1) present.

9. Length of rostrum: 0) long, surpassing metacoxae; 1) not reaching
beyond metacoxae; 2) short, not reaching beyond mesocoxae.

10. Shape of the anterolateral margins of pronotum (Bernardes et al.,
2009): 0) strongly depressed dorso-ventrally, emarginated; 1) not
depressed or emarginated.

11. Development and shape of humeral angles: 0) not produced, never
surpassing the base of the adjacent corium, rounded; 1) always
surpassing the base of the adjacent corium, developed into a conical
spine; 2) always surpassing the base of the adjacent corium, developed
into tubercles directed upward, dorsally bifid; 3) slightly surpassing
the base of the adjacent corium, developed into a prominent angulation;
4) always surpassing the base of the adjacent corium, produced into a
cornuted projection.

12. Length of the scutellum in relation to the coria: 0) not reaching
beyond the apices of coria; 1) surpassing the apices of coria.

13. Length of frena in relation to the scutellum: 0) reaching the
middle of the scutellum; 1) reaching the basal third of
the scutellum; 2) reaching the basal fourth of the scutellum.

14. Shape of the lateral margins of scutellum: 0) sinuous; 1) straight.

15. Surface of the mesosternum: 0) carinated; 1) sulcated; 2) flat.

16. Length of the peritreme: 0) not extending more than half the length
of evaporatorium; 1) reaching or extending more than half the length of
evaporatorium.

17. Shape of the evaporatorium on mesopleurum (Bernardes et al., 2009):
0) in a continuous diagonal strip; 1) in a discontinuous diagonal
strip or absent.

18. Presence of the spine at apex of the femur: 0) absent; 1) present.

19. Presence and length of the abdominal spine: 0) absent; 1) present
but inconspicuous, not reaching metacoxae; 2) present and well
developed, reaching or surpassing metacoxae; 3) present and well
developed, surpassing mesocoxae.

20. Shape of the abdominal spine 0) straight; 1) dorsally curved

21. Pygophore, length of the dorsal wall: 0) about 1/2 of pygophore
width; 1) about 1/4 of the pygophore width.

22. Pygophore, ventral wall produced posteriorly, ventral rim with a
distinct medial emargination (Rider, 1994): 0) absent; 1) present.

23. Pygophore, shape of the ventral rim laterad to the medial
emargination: 0) in 1 + 1 enlarged rims, lip-like flaps; 1)
in 1 + 1 keel-like processes; 2) in 1 + 1 broadly spatulate processes
(Rolston, 1978); 3) in 1 + 1 plate-like processes (Nelson, 1955;
Bernades et al., 2009).

24. Processus phallothecae, presence and shape: 0) absent; 1)
rectangular; 2) pyramidal; 3) spherical.

25. Dorsal lobe of conjunctiva (Gapon, 2005): 0) present; 1) absent.

26. Length of vesica in relation to phallotheca: 0) longer than
phallotheca; 1) subequal in length to phallotheca; 2) shorter than
phallotheca.

27. Shape of the dorsal surface of the segment X in males: 0) convex;
1) flat; 2) sulcated;

28. Paramere, apex in one or two lobes: 0) one lobe; 1) bilobed.

29. Length of gonocoxites 8 in relation to the laterotergites 9: 0)
subequal; 1) smaller.

30. Shape of capsula seminalis: 0) globose, without constriction; 1)
digitiform; 2) globose, with a medial or basal constriction; 3)
semiglobose, with an apical process; 4) conical; 5) subquadrangular.

Table 2. Character matrix.

Euschistus heros            -000000000100001000-00-0100000
Thyanta perditor            0000000000100000000-00-0020001
Piezodorus guildinii        0002000120000000003000-0120002
Rio pectoralis              0000001120000000002000-0110002
Banasa induta               0000000000000000001000-0120000
Arocera acroleuca           0000000000000000000-00-0020002
Aleixus virgatus            00?10?11112000200110????????l?
Aleixus tobyschuhi          000100111120002101100101121013
Brepholoxa heidemanni       100100111010001100200101022014
Odmalea concolor            001001111110002001300121011113
Odmalea basalis             001001111110002001200111011013
Paraodmalea rubella         0010011111100020010-0111011114
Pseudobebaeus truncatus     101001112140001001300111021113
Dendrocoris pini            100000111000001000100131010114
Dendrocoris humeralis       110000111030001000100131010114
Thoreyella brasiliensis     110210112111112011311131110113
Thoreyella cornuta          110210112111112011311131110113
Procleticus corniger        110200112111101011101131110111


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Article Details
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Author:Schwertner, Cristiano F.; Grazia, Jocelia
Publication:Entomologica Americana
Article Type:Report
Geographic Code:3BRAZ
Date:Jan 1, 2012
Words:5195
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