Review of the Neotropical Genus Aleixus Mcdonald (Hemiptera: Heteroptera: Pentatomidae: Procleticini), with Description of a New Species and Cladistic Analysis of the Tribe Procleticini

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Review of the Neotropical Genus Aleixus Mcdonald(Hemiptera: Heteroptera: Pentatomidae: Procleticini), withDescription of a New Species and Cladistic Analysis of the TribeProcleticiniAuthor(s): Cristiano F. Schwertner and Jocelia GraziaSource: Entomologica Americana, 118(1):252-262. 2012.Published By: The New York Entomological SocietyDOI: http://dx.doi.org/10.1664/12-RA-034.1URL: http://www.bioone.org/doi/full/10.1664/12-RA-034.1

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REVIEW OF THE NEOTROPICAL GENUS ALEIXUS MCDONALD (HEMIPTERA:

HETEROPTERA: PENTATOMIDAE: PROCLETICINI), WITH DESCRIPTION OF ANEW SPECIES AND CLADISTIC ANALYSIS OF THE TRIBE PROCLETICINI

CRISTIANO F. SCHWERTNER1

AND JOCELIA GRAZIA2

1Departamento de Ciencias Biologicas and Programa de Pos-graduacao em Ecologia & Evolucao,

Instituto de Ciencias Ambientais, Quımicas e Farmaceuticas, Universidade Federal de Sao Paulo,

Diadema, SP, Brazil, email: [email protected] 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: [email protected]

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 tobyschuhi described from southern

Brazil, greatly extends the distribution of Aleixus. 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 5

74, CI 5 60, RI 5 78). Analyses under SW and IW resulted in one most parsimonious cladogram, with fit varying

from 20.4 (K 5 0.64) to 26.6 (K 5 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 char-

acteristics 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 sub-

triangular 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 Ber-

groth, Zorcadium Bergroth (5Pseudobebaeus Dis-

tant), 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 McDon-

ald, 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

Entomologica Americana 118(1–4):252–262, 2012

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 Pen-

nington (1920) to include the monotypic genera

Lobepomis Berg and Procleticus Berg, and diag-

nosed 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 Ber-

groth, 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

10 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 Muse-

um 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, repre-

senting 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-and-

bound). For the definition of the K values in the

implied weighting analysis, we followed Mirande

2012 REVIEW OF NEOTROPICAL ALEIXUS 253

(2009); the scripts aaa.run and aab.run provided

by the author were adapted to implicit enumera-

tion 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 ‘‘Xy’’, in which X represents the

character and Y represents the state.

Measurements (mean, minimum and maxi-

mum) are given in millimeters. In the text, the

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 K 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.

254 ENTOMOLOGICA AMERICANA Vol. 118(1–4)

proportions of the segments were represented with

the symbols: ‘‘,’’ minor, ‘‘.’’ greater, ‘‘<’’

subequal. The terminology of Kment and Vili-

mova (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 sub-

triangular, apex broadly rounded not reaching

apices of coria. Thoracic sterna nearly flat. Peri-

treme 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 tobyschuhi, 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

Table 2. Character matrix.

Euschistus heros -000000000100001000-00-0100000Thyanta perditor 0000000000100000000-00-0020001Piezodorus guildinii 0002000120000000003000-0120002Rio pectoralis 0000001120000000002000-0110002Banasa induta 0000000000000000001000-0120000Arocera acroleuca 0000000000000000000-00-0020002Aleixus virgatus 00?10?11112000200110????????1?Aleixus tobyschuhi 000100111120002101100101121013Brepholoxa heidemanni 100100111010001100200101022014Odmalea concolor 001001111110002001300121011113Odmalea basalis 001001111110002001200111011013Paraodmalea rubella 0010011111100020010-0111011114Pseudobebaeus truncatus 101001112140001001300111021113Dendrocoris pini 100000111000001000100131010114Dendrocoris humeralis 110000111030001000100131010114Thoreyella brasiliensis 110210112111112011311131110113Thoreyella cornuta 110210112111112011311131110113Procleticus corniger 110200112111101011101131110111

2012 REVIEW OF NEOTROPICAL ALEIXUS 255

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 dark-

ened. Urosternite VII 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 < V. Rostrum scarcely reaching

metacoxae. Pronotum: Anterolateral margins con-

cave 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: Scu-

tellum surpassing the middle of connexival

segment V but not attaining apex of coria.

Peritreme extending 1/3 the distance from ostiole

Fig. 1. Aleixus virgatus McDonald, holotype female. Views: A. Dorsal. B. Lateral. C. Frontal. Scale bar 5 1 mm.

Fig. 2. Aleixus tobyschuhi new species, holotype

male, dorsal view. Scale bar 5 1 mm.

256 ENTOMOLOGICA AMERICANA Vol. 118(1–4)

Fig. 3. Aleixus tobyschuhi new species. A. Head and pronotum in frontal view. B. right metapleuron showing

peritreme. C. Apex of femur. Scale bar 5 1 mm (A, B) and 0.5 mm (C).

Fig. 4. Aleixus tobyschuhi new species, male. A. Pygophore, dorsal view. B. Pygophore, posterior view. C.

Pygophore ventral view. D-F. Phallus, dorsal, lateral and ventral views. G. Right paramere, lateral view.

Abbreviations: bp 5 basal plates of articulatory apparatus, cj 5 conjunctiva, dc 5 dorsal connectives, dr 5 dorsal

rim, par 5 paramere, pc 5 processus capitati, spdr 5 superior process of dorsal rim, ph 5 phallotheca, pph 5

processus phallothecae, sg 5 secondary gonopore, v 5 vesica, vre 5 ventral rim emargination, vrp 5 ventral rim

projection, x 5 segment X. Scale bar 5 1 mm.

2012 REVIEW OF NEOTROPICAL ALEIXUS 257

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 (<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 J of phallotheca length, dorsal

connectives (dc) short and processus capitati (pc)

almost reaching the middle of phallotheca.

Processus phallothecae (pph) present. Conjuncti-

va with 1 + 1 assymetrical membranous lobes

obscuring the slightly sinuous vesica.

Male. Measurements (N 5 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 (0.49–

0.57); Pronotum length: 1.53 (1.47–1.64). Prono-

tum: 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 (la8)

very poorly delimited, apparently forming an

entire band ventrally, lacking spiracles. Latero-

tergites 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 semiglo-

bose with a digitiform projection.

MEASUREMENTS. (N 5 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

Fig. 5. Aleixus tobyschuhi new species, female. A.

genital plates; B. laterotergites 8 and 9, gonocoxites 9,

gonapophyses 9, and ectodermical genital ducts (aaf 5

anterior annular flange, chi 5 chitinellipsen, cs 5

capsula seminalis, dre 5 ductus receptaculi, g8 5

gonapophyses 8, g9 5 gonapophyses 9, gc8 5 gonocox-

ites 8, gc9 5 gonocoxites 9, la8 5 laterotergites 8, la9 5

laterotergites 9, paf 5 posterior annular flange, pco 5

pars communis, pi 5 pars intermedialis, tvi 5 thickening

of the vaginal intima, va 5 vesicular area, x 5 segment

X). Scale bar 5 1 mm.

258 ENTOMOLOGICA AMERICANA Vol. 118(1–4)

(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 (0.49–

0.57). Pronotum length: 1.82 (1.64–2.05). Prono-

tum 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 dis-

tinctly sinuous, slightly reflexed; apex of head

distinctly declivent when viewed laterally; anten-

niferous 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 sub-

triangular, 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 heteropter-

ous 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: BRA-

ZIL, Santa Catarina, Rio Vermelho, lack abdo-

men, X.1958, Dirings (MZSP); Rio Grande do Sul,

Vila Oliva, 24.X.1957, Pe. Buck leg., 3 R, 1 =(MAPA); 21.II.1954, Pe. Buck leg., 1= (AMNH);

28.II.1954, Pe. Pio Buck leg, 1R (AMNH); PortoAlegre, 2 R (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 5 0.64) to 26.6 (K 5 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 Procleticus and Thor-

eyella, is also supported in all analyses. The

phylogenetic position of the remaining genera is

unresolved under EW. In the cladograms ob-

tained 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 homo-

plastic (Fig. 7). The humeral angles developed

into large tubercles directed upward and dorsally

bifid (112) 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

Fig. 6. Strict consensus of four most parsimonious

cladograms found in the analysis with equal weights

(L 5 74, CI 5 60, RI 5 78).

2012 REVIEW OF NEOTROPICAL ALEIXUS 259

than antennal segment I (41), 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-homo-

plastic for all species of Procleticini (221 and 291),

and together with the presence of processus

phalothecae (241), 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 Brepholoxa

Dendrocoris, Odmalea, Parodmalea and Pseudobe-

baeus, based on all having a short scutellum.

Bernardes et al. (2009) found that the elongation of

the scutellum among Procleticini is a syapomorphy

Fig. 7. Cladogram resulted from analyses under successive and implied weighting (Fit of 21.53, K 5 1), showing

optimization of unambiguous characters. Black circles represent nonhomoplastic derived states; white circles

represent homoplastic derived states. Numbers above circles represent characters; numbers below circles represent

character state. Number at nodes represent major clades (see Discussion).

260 ENTOMOLOGICA AMERICANA Vol. 118(1–4)

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 (271) is characteristic to this clade. The

pronotum with anterolateral margins not emar-

ginate (101) and presence of a spine at the apex of

the femur (181) also support the clade, although

these characteristics are also shared with Procle-

ticus and Thoreyella (Fig. 7, node 8). The meso-

sternum flat (152) 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 mono-

typic 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. Revision-

ary studies of these taxa, taking into account the

totality of their morphological diversity, will be

crucial to defining their identity and their phylo-

genetic relationships among Procleticini. Particu-

larly 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 au-

thor 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 (71 and 81) and the length

of rostrum (91 and 92). The weakly developed

bucculae, with both anterior and posterior mar-

gins 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 rela-

tionship 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 theloan of the material studied, especially Thomas J.Henry (NMNH) for providing access and facilities tothe study of the holotype of A. virgatus. This workwas supported by grants from Conselho Nacional dePesquisa (CNPq) and Coordenacao de Aperfeicoa-mento de Pessoal de Nivel Superior (CAPES) to theauthors. We also thank two anonymous referees forproviding constructive comments and suggestionsthat improved the manuscript.

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