Cladistics and revision of Alitocoris with considerations on the phylogeny of the Herrichella clade (Hemiptera, Pentatomidae, Discocephalinae, Ochlerini) Author Garbelotto, Thereza De A. Departamento Zoologia, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves 9500, Prédio 43435, Porto Alegre, RS 91501 - 970, Brasil therezagarbelotto@hotmail.com Author Campos, Luiz A. Departamento Zoologia, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves 9500, Prédio 43435, Porto Alegre, RS 91501 - 970, Brasil Author Grazia, Jocelia Departamento Zoologia, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves 9500, Prédio 43435, Porto Alegre, RS 91501 - 970, Brasil text Zoological Journal of the Linnean Society 2013 2019-12-17 168 452 472 journal article 24586 10.1111/zoj.12032 0e835bd9-b158-4858-a55a-fc5a32cdfe68 3585663 9AFF707D-E3FB-4176-A12B-82F3757A772F ALITOCORIS SCHRADERI SAILER, 1950 ( FIGS 3A , 4A, B , 5A–G , 6A, E, 7A, B) Alitocoris schraderi Sailer, 1950: 71–72 , figures 1, 2, 12, 13, 17 ; Schrader, 1960a: 105–110 ; Schrader, 1960b: 499–500 , figure 4 ; Arismendi & Thomas, 2003: 226 . Holotype : GUATEMALA , 1 male NMNH Type Cat. no. 59342 ( Sailer, 1950 ) . Allotype : GUATEMALA , 1 female ( Sailer, 1950 ) . Figure 3. Species of Alitocoris , dorsal view. A, Alitocoris schraderi (male). B, Alitocoris grandis sp. nov. (holotype female). C, Alitocoris lateralis sp. nov. (holotype female). D, Alitocoris ornatus sp. nov. (holotype male). Scale bars = 1 mm Paratypes : COSTA RICA , Cartago : Turrialba , 1 m and 1 f ; San Jose : San Isidro Del General , 1 m and 2 f ( Sailer, 1950 ) . . Material examined: HONDURAS , Santa Barbara : ( 13 km south-east of El Mochito ), 1 f , 22.vii.1977 , OBrien ( DARC ) . COSTA RICA , ( Hamburg Farm ), 1 f , 31.iii , C. P. Dodge ( AMNH ) ; Heredia : ( Estación Biológica La Selva ), 1 f and 1 m , 10–12.vi.1986 , B. C. Ratcliffe & party ( DBTC ) ; (Estación Biológica La Selva 2 km south of Puerto Viejo ), 1 f , 15–22.iii.1987 : J. Negrón ( DARC ) ; ( Estación Biológica La Selva 150 ma .s.l. ), 1 f , vii.-viii.1998 , FIT, Nico Franz ( JEE ) ; ( Finca La Selve Verde , 12 km south of Puerto Viejo, 125.4 ma .s.l. ), 1 f , 23–26.ix.1986 , J. E. Eger , [ 10.4667 ; -84.0167 ] ( JEE ) ; Guanacaste : ( Estación Pitilla , 9 km south of Sta. Cecilia, 700 m a.s.l .), 1 f ( Costa Rica INBio CRI 001 008706 ), vii.1988 , GNP Biodiversity Survey [ 11.1000 ; -85.4167 ], 10°59 ′ N, 85°25 ′ W ( INBio ) ; 1 m ( Costa Rica Inbio CRI 000154808 ) and 1 f ( Costa Rica Inbio CRI 000154862 ), ii.1990 , P. Ríos, C. Morgada & R. Blanco , [ 11.1000 ; -85.4167 ], 330200 , 380200 ( INBio ) ; ( Est. Pitilla , 9 km south of Sta Cecilia, P. N. Guanacaste , 700 m a.s.l. ), 1 f ( Costa Rica Inbio CRI 000808667 ), v.1992 , P. Ríos , [ 11.1000 ; -85.4167 ], L-N 330200, 380200 ( INBio ) ; 1 f ( Costa Rica CRI 000 819068 ), 10–17.vi.1992 , C. Moraga , [ 11.1000 ; -854167 ], L-N 330200 , 380200 ( INBio ) ; 1 f and 1 m ( Costa Rica Inbio CRI 001818072 ), i.1994 , P. Ríos , [ 11.1000 ; -854167 ], L-N 330200 _ 380200 ( INBio ) ; 1 f ( Costa Rica Inbio CRI 002 048807 ), vii.1994 , P. Ríos , [ 11.1000 ; -85.4167 ], L-N 330200_380200 # 3140 ( INBio ) ; ( Est. Biol. Cacao , A. C. Guanacaste , 1100 ma .s.l. ), 1 m ( Costa Rica Inbio CRI 002113078 ), xii.1993 – ix.1994 , Fam. Guadamuz Chavarría, [ 10.9333 ; -85.4500 ], L-N 323300_375700 # 3318 ( INBio ) ; 1 f ( Costa Rica Inbio CRI 002113079 ), xii.1993 – i.1994 , Fam. Guadamuz Chavarría , [ 10.9333 ; -85.4500 ], L-N 323300_375700 # 3318, ( INBio ) ; ( Estación Cacao , south-west side Volcan Cacao , 1000–1400 ma .s.l. ), 1 m ( Costa Rica Inbio CRI 000147453 ), 1 m ( Costa Rica Inbio CRI 000147468 ) and 1 f ( Costa Rica Inbio CRI 000 147462 ), xi.– xii.1989 , URCG, R. Blanco & C. Chavez , [ 10.9333 ; -85.4500 ], 323300, 375700 ( INBio ) ; Chilamate (Selva Verde Lodge) , 1 m , 11–13.viii.1995 , J. E. Eger , [ 10.4500 ; -84.0667 ], Costa Rica collecting permit no. 00113412 ( JEE ) ; ( Est. Magasasay, P. N. Braulio Carrillo , 200 m a.s.l. ), 1 m ( Costa Rica Inbio CRI 000298374 ), i.1991 , M. Barrelier , [10.1599; -839744], L-N 264600, 531100, ( INBio ) ; Cartago : Turrialba , 1 m , 20.v.1948 , F. Schrader 445 [ 9.9000 ; -83.6833 ] ( AMNH ) ; 1 f , 23.vi.1948 , F. Schrader 597, [ 9.9000 ; -83.6833 ] ( AMNH ) ; Limon : ( Sector Cerro Cocori, Fca de E. Rojas , 150 m a.s.l .), 1 f ( Costa Rica Inbio CRI 000794714 ), 26.iii.–24.iv.1992 , F. A. Quesada , L-N 286000, 567500 ( INBio ) ; Cerro Tortuguero ( P. N. Tortuguero , 100 ma .s.l. ), 1 f ( Costa Rica Inbio CRI 000088898 ), iv.1989 , R. Aguilar & J. Solano , [ 10.5333 ; -83.5167 ], 285000, 588000 ( INBio ) ; Amubri ( 70 m , Talamanca ), 1 f ( Costa Rica Inbio CRI 000 965969 ), 12–30.ix.1992 , G. Gallardo , [ 9.5167 ; -82.9667 ] L-S 385500, 578050 ( INBio ) . PANAMA , Bocas del Toro : Corriente Grande ( 100 m a.s.l. ), 1 m , 25.iv.1980 , Henk Wolda , [ 9.5167 ; -82.8667 ], 9°17 ′ N, 82°32 ′ W ( DBTC ) . Figure 4. Species of Alitocoris , pygophores. A, B, Alitocoris schraderi . C, D, Alitocoris ornatus sp. nov. A, C. dorsal view. B, D. ventral view. Abbreviations: dr, dorsal rim; pa, posterolateral angle; par, paramere; pil, projection of inferior layer of ventral rim; vr, ventral rim; X, segment X. Scale bar = 1 mm. Diagnosis: Dark brown to black, punctures deep and concolorous. Legs not contrasting with body colour. Head black, punctures not confluent at apical third of juga. Vertex slightly tumid. Antennae dark brown to black, except distal half of fourth segment and entire fifth segment light brown; dorsal pale blot on first segment, sometimes inconspicuous; third segment sometimes light brown. Proportional length of antennal segments: I < II < III > IV < V; fifth the longest. Bucculae lacking anterior tooth. Labium yellowish, reaching to posterior margin of urosternite V; proportinal length of labial segments: I < II > III > IV. Figure 5. Alitocoris schraderi , phallus and right paramere. A–D, phallus: A, dorsal; B, anterior; C, posterior; D, lateral views. E–G, paramere: E, dorsal; F, ventral; G, lateral views. Abbreviations: bv, bend of vesica; dp, dorsal process of phallotheca; ds, ductus seminis distalis; lph, lateral projection of phallotheca; ph, phallotheca; v, vesica. Scale bars = 0.5 mm . Pronotum bearing median transverse discontinuous line, lighter anteriorly, sometimes inconspicuous; punctures shallower between anterior margin of pronotum and cicatrices. Cicatrices flat. Anterolateral angles parallel to posterior margin of eyes, exceeding lateral limit of eyes for at most the diameter of ocellus. Anterolateral margins slightly sinuous. Scutellum reaching the level of middle of fifth segment of connexivum; punctures confluent on disc, forming wrinkles; 1 + 1 minute yellowish spots close to foveae. Corium with few pale spots; apex little surpassing anterior margin of sixth segment of connexivum. Yellowish spot at apex of radial vein equivalent to the diameter of ocellus. Membrane with nine veins, reaching or little surpassing apex of abdomen. Pro-, meso-, and metapleura black. Peritreme exceeding half the width of evaporatorium. Coxae and trochanters yellowish with coppery areas; femora and tibiae dark brown; tarsi yellowish. Connexivum black. Abdomen velvety, sparsely pubescent; shallow punctures inconspicuous on disc; tubercle of trichobothria brown. Anteromedian margin of spiracles yellowish, not elevated. Male . Measurements ( N = 9): head length 1.76 ± 0.07 (1.64–1.83); width 2.42 ± 0.06 (2.33–2.52); pronotum length 2.32 ± 0.11 (2.08–2.46); width 5.97 ± 0.19 (5.73–6.30); scutellum length 4.03 ± 0.14 (3.78–4.22); width 3.59 ± 0.13 (3.34–3.78); length of antennal segments: I 0.73 ± 0.03 (0.69–0.76); II 0.68 ± 0.04 (0.63–0.76); III 1.63 ± 0.07 (1.51–1.70); IV 1.38 ± 0.06 (1.26–1.45); V 1.72 ± 0.13 (1.51–1.89); length of labial segments: I 1.23 ± 0.05 (1.13–1.26); II 2.18 ± 0.06 (2.08–2.27); III 1.76 ± 0.12 (1.58–1.89); IV 1.55 ± 0.12 (1.26–1.64); total length 10.61 ± 0.15 (10.40–10.80); width of abdomen 6.32 ± 0.14 (6.10– 6.50). Figure 6. Species of Alitocoris , female. A–D, genital plates, ventroposterior view. E, F, receptaculum seminis and ausenwand, ventral view. A, E, Alitocoris schraderi . B, Alitocoris grandis sp. nov. C, Alitocoris lateralis sp. nov. D, F, Alitocoris ornatus sp. nov. Abbreviations: ch, chitinellipsen; cm, conical microsculptures; cs, capsula seminalis; d9, basal depression of laterotergites 9; dr, ductus receptaculi; gc8, gonocoxites 8; gc9, gonocoxites 9; id, inner duct of vesicular area; la8, laterotergites 8; la9, laterotergites 9; md, median duct of vesicular area; od, outer duct of vesicular area; pi, pars intermedialis; tg9, thickening of gonapophyses 9; vi, thickening of vaginal intima; VII, urosternite VII. Scale bars = 1 mm . Genitalia. Dorsal rim widely excavated with 1 + 1 spine projections directed towards the base of segment X. Inferior layer of ventral rim projecting lateral to segment Xover genital cup, perpendicular to the frontal plane of pygophore; projections spatulate, in dorsal view with an elevation midway between apex of projection and posterolateral angle of pygophore. Superior layer of ventral rim bent beneath the projection of inferior layer, forming a translucent, less sclerotized lobe, and looking almost membranous ( Figs 4A , 7B ). Median third of ventral rim concave, emarginate ( Fig. 4B ). Lateral margins of segment X sinuous; 1 + 1 lateral tuft of short setae; apex covered with fringe of long setae and narrowed from the base ( Fig. 4A ). Parameres sinuous, slender, and lacking projections; head spatulate with obtuse apex, slightly curved ventrally; fringe of setae on lateral and apex of head; base flattened, with dense, short setae ( Fig. 5E– G ). Parameres placed ventral to segment X, projecting over and little surpassing the ventral rim of pygophore ( Fig. 4B, D ). Phallus. Phallotheca globose, lateral projection wide; ventral and dorsal apical processes short, little projected. Vesica long, pseudojointed and bent backward, surpassing the base of phalotheca; dorsal expansion at basal half of vesica ( Fig. 5A–D ). Female . Measurements ( N = 15): head length 2.00 ± 0.07 (1.89–2.14); width 2.58 ± 0.07 (2.46–2.65); pronotum length 2.76 ± 0.11 (2.52–2.96); width 6.66 ± 0.26 (6.3–7.18); scutellum length 4.65 ± 0.2 (4.35–5.04); width 3.97 ± 0.17 (3.65–4.28); length of antennal segments: I 0.78 ± 0.04 (0.69–0.82); II 0.81 ± 0.07 (0.69–0.95); III 1.61 ± 0.09 (1.45–1.76); IV 1.49 ± 0.06 (1.39–1.58); V 1.82 ± 0.13 (1.58–1.95); length of labial segments: I 1.33 ± 0.07 (1.26–1.45); II 2.27 ± 0.11 (2.08–2.46); III 1.91 ± 0.16 (1.39–2.08); IV 1.70 ± 0.09 (1.58–1.83); total length 12.23 ± 0.45 (11.7–12.9); width of abdomen 7.11 ± 0.25 (6.7–7.5). Figure 7. Scanning electron microscopy of genitalia of Alitocoris schraderi . A, female, arrow indicates conical microsculptures of gonapophyses 9. B, male, superior layer of pygophore ventral rim, detail of lobular expansion (arrow). Genitalia. Gonocoxites 8 wider than long; posterior margins sinuous, concave medially, projected over the base of laterotergites 9; disc with median callus. Spiracles of laterotergites 8 on low callus, placed lateral to lateral angle of gonocoxites 8 ( Fig. 6A ). Gonocoxites 9 with longitudinal suture; anterior margin subrectilinear ( Fig. 6A, E ). Laterotergites 9 not surpassing the band connecting laterotergites 8; median margins on a higher plane than lateral margins; lateral basal angle flat; anterior margin slightly sinuous ( Fig. 6A, E ). Gonapophyses 9 with conical microsculptures occupying lateral portion of posterior folds ( Figs 6E , 7A ). Thickening of gonapophyses 9 in wide ‘V’. Chitinellipsen lateral to thickening of vaginal intima. Ductus receptaculi anteriorly to vesicular area wider than median duct of vesicular area. Length of vesicular area twice the combined length of pars intermedialis and capsula seminalis; base of median duct clavate; inner duct uniformly cylindrical. Tubular portion of ductus receptaculi posterior to vesicular area short, not attaining half length of vesicular area. Pars intermedialis barrelshaped; at least four times longer than capsula seminalis; annular ridges convergent. Capsula seminalis hemispherical; projections long, slender, and twisted, sometimes bifid at apex, placed at base of capsula seminalis ( Fig. 6E ). Comments: Alitocoris schraderi is in a clade with Ali. grandis sp. nov. and Ali. lateralis sp. nov. ( Fig. 1 ), and can be distinguished by the rostrum reaching the posterior margin of urosternite V, and antennal segment Vbeing the longest. Gonocoxites 8 of Ali. schraderi bears a more tumid callus, and lateral angle more projected over laterotergites 9. The material examined in this study extends the distribution of the species to Honduras and Panama.