Taxonomic revision of Montina (Hemiptera: Heteroptera: Reduviidae) from Colombia with description of three new species Author Mejía-Soto, Andrés Grupo de Entomología Universidad de Antioquia (GEUA), Universidad de Antioquia, Medellín, Colombia; e-mails: andres. mejias 300 @ gmail. com, martha. wolff @ udea. edu. co Author Forero, Dimitri Laboratorio de Entomología, UNESIS, Departamento de Biología, Pontificia Universidad Javeriana, Bogotá, Colombia; and Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia (present address); e-mail: iforerof @ unal. edu. co Author Wolff, Marta Grupo de Entomología Universidad de Antioquia (GEUA), Universidad de Antioquia, Medellín, Colombia; e-mails: andres. mejias 300 @ gmail. com, martha. wolff @ udea. edu. co text Acta Entomologica Musei Nationalis Pragae 2022 2022-12-16 62 2 325 381 http://dx.doi.org/10.37520/aemnp.2022.019 journal article 10.37520/aemnp.2022.019 1804-6487 7617332 Montina Amyot & Serville, 1843 Montina Amyot & Serville, 1843: 363 (description). Type species: Reduvius sinuosus Lepeletier & Serville, 1825 , by monotypy. Montina : STÂL (1859) : 197 (new species); STÂL (1865) : 48 (key); STÂL (1872) : 73 (list of species); WALKER (1873) : 91 (list of species); LETHIERRY & SEVERIN (1896) : 195 (catalog); CHAMPION (1899): 286 (list, diagnosis); PUTSHKOV & PUTSHKOV (1988) : 115 (catalog); MALDONADO (1990) : 234 (catalog); ZHANG & WEIRAUCH (2014) : 341 (phylogenetic placement); ZHANG et al. (2016) : 540 (phylogenetic placement); GIL- SANTANA (2019): 516 (new records). Ploeogaster (nec Amyot & Serville, 1843 ): STÂL (1859) : 197 (new species); STÂL (1872) : 73 (synonym). Aristippus Stål, 1865: 48 (description in key). Type species: Aristippus fenestratus Stål, 1867 , by subsequent designation. Aristippus : STÂL (1867) : 48 (description, no species assigned); STÂL (1867) : 299 (new species); STÂL (1868) : 99 (key); STÂL (1872) : 74 (list, synonym of Montina as subgenus); WALKER (1873) : 93 (list of species, as synonym of Ploeogaster ); LETHIERRY & SEVERIN (1896) : 195 (catalog, as subgenus); PUTSHKOV et al. (1987) : 103 (as valid subgenus); PUTSHKOV & PUTSHKOV (1988) : 115 (catalog, as valid subgenus); MALDONADO (1990) : 234 (catalog). Diagnosis. Distinguished among all Neotropical Harpactorini genera by the following combination of characters: head elongate, length/width ratio 2.3; disc of the anterior lobe of the pronotum with a pair of discal tubercles ( Figs 2A–B ; 5E–F ; 7E ; 9E–F ; 11F ), each posteriorly connected to the submedial longitudinal carinae on the posterior pronotal lobe; submedial carina on posterior lobe rises prominently on the discal area forming a laterally compressed elevation ( Figs 2B ; 15E ), which have its dorsal area truncated or rounded; the posterior margin of the pronotum have a pair of paramedial lobes ( Figs 2A ; 13D ; 19B ), each with its lateral margin entire and curved, and the margin between the lobes (in front of the scutellum) from slightly to strongly curved; the humeral angle of the pronotum is rounded, and anterior to it there is a laterally directed process, the posterolateral process ( Fig. 2A ); the mesopleura has a tubercle (the “plica” of authors); the connexivum is strongly protruding laterally on the abdomen ( Figs 7B, D ; 15B, D ), the connexival margins range from being almost straight ( Figs 7C ; 9C ) to strongly rounded lobes ( Figs 15A, C ). Males have the medial process of the pygophore ( mpp ) almost cylindrical in cross section, short, directed posteriorly at about 45 degrees, slightly to greatly widened basally, and with a small subapical hook-like posteriad projection ( Figs 3A–B ); the parameres ( pa ) are long, reaching the medial process, narrow, slightly enlarged apically and curved basally ( Figs 6A, C ; 8A, C ; 10A, C ); the phallosoma has the dorsal phallothecal sclerite ( dps ) ovoid with its margin broadly rounded ( Figs 10F ; 14F ); the endosoma has a pair of elongated basal dorsolateral lobes ( bdl ), a small distal dorsal lobe ( ddl ) beset with microtrichia, a pair of small distal lateral lobes ( dll ), a bilobed distal ventral lobe ( dvl ), and a pair of lateral lobes ( ll ) ( Figs 3C–E ). Females have the gonocoxa 8 with an emargination on the posterior margin ( Fig. 24A , red arrow); the bursa copulatrix has long and wide projecting lateral lobes ( lbs ) which are as long as the length of the bursa ( Figs 26 ; 27 ), anterior dorsal region with a dorsal semi-sclerotized fold and a ventral pair of folds, usually with a U-shaped sclerotization ( usp ) ( Fig. 28A ). Fig. 1. Live specimens of Montina . A–B – M. confusa ( Stål, 1859 ) , female, Madre de Dios, Peru (A – lateral view, B – dorsal view) (photo by Sandra Lamberts). C – M. lobata Stål, 1859 , Orellana, Ecuador (photo by Felipe Campos). D – M. scutellaris Stål, 1859 , female, Chocó, Colombia (photo by Kritzzia Copete Murillo). E – M. gladiator sp. nov. , pair in copula, Yolombo, Colombia (photo by Julian Vallejo-Sosa). F – M. calarca sp. nov. , pair in copula, Risaralda, Colombia (photo by Fernando Lopez). Biology. Individuals of Montina are generally found on shrubs or low vegetation ( Fig. 1 ) and are common in agricultural areas. All examined specimens were collected manually or with an entomological net, which suggests that other types of collecting methods may not be very effective. The biology and natural history of the species are poorly known, only being documented aspects of the predatory capacity, life cycle, and description of immatures of M. confusa ( BUENO & BERTI FILHO 1984a , b, 1987; DELLAPḖ et al. 2002; FREITAS 1994 , 1995 ). Montina has also been reported as a natural predator of agricultural pests. Montina confusa is a common predator of Lepidoptera larvae in Eucalyptus plantations ( ZANUNCIO et al. 1993 , 1994 ) and of other pests in different crops in Brazil (e.g., TREVISAN & MENEGUETTI 2012 ). Similarly, in Colombia unidentified species of Montina has been documented as predators of insect pests of soybean, corn, and other crops (e.g., AYALA et al. 2013 , GUEVARA & JIMḖNEZ 2018, LEṒN MARTÍNEZ & GUEVARA AGUDELO 2006 ). Thus, it is very important to adequately document the species to help plan future pest management programs that are willing to include Montina species in their strategies. Differential diagnosis. The original description of Montina is short and not very detailed ( AMYOT & SERVILLE 1843 ); however, it mentions two important characters, the laterally protruding abdomen, and a pair of tubercles on the anterior lobe of the pronotum connected with a submedial longitudinal carina. These characters are shared with Ploeogaster . Given the extreme similarity between these two taxa we have based our hypothesis of the generic limits of Montina on the study of photographs of the lectotype and paralectotype of Reduvius sinuosus Lepeletier & Serville, 1825 ( Fig. 38 ) (NHMW) (see below), and numerous Colombian specimens of both Ploeogaster and Montina . We thus propose novel characters to help delimit the two genera ( Table 1 ). The presence of a pair of prominent tubercles on the anterior pronotal lobe that are continued posteriorly by a longitudinal submedial carina is a common feature in both genera. The tubercles of the anterior lobe of the pronotum have no marked differences in both genera, as they can be small acute tubercles or subcylindrical tubercles with a rounded apex. The structure of the posterior pronotal lobe between the two genera, on the other hand, shows a marked difference that has not been documented before. In Montina , the longitudinal submedial carina is slightly compressed laterally, and has an elevated portion on the discal area of the posterior lobe, this elevation is dorsally truncated or slightly rounded ( Figs 21E ; 23C ); whereas in Ploeogaster the longitudinal submedial carina is less compressed, and the elevated portion on the discal area has a tubercle with a rounded apex ( Fig. 4B ). The posterior margin of the pronotum presents in Montina a pair of paramedial lobes, which have an entire lateral margin ( Figs 2A ; 7B ; 13D ), whereas in Ploeogaster these lobes have the lateral margin deeply emarginated giving the impression of being almost bilobed ( Fig. 4A ). In both genera the shape of the posterior margin connecting the paramedial lobes range from being almost straight to strongly curved, so we consider this a variable character between the genera which might be species specific. In Montina , the posterolateral process of the pronotum, which is located anterior to the humeral angles, is small and no longer than its base width ( STÂL 1867 ), sometimes being a blunt tubercle ( Figs 5D ; 13D ). In Ploeogaster , the posterolateral process is a conspicuous acute projection directed laterally and always longer than its base, giving the appearance of a sharp process ( Fig. 4A ). Another unexplored character is the structure of the fore legs. In Montina , the fore femur is cylindrical on its entire length with its ventral margin nearly straight, and the fore tibia is also straight ( Figs 5A, C ). Ploeogaster , on the other hand, has the fore femur wider near the middle with its ventral margin slightly curved ventrally, being thus the fore tibia similarly curved ( Fig. 4C ). It also seems that the profemur in Ploeogaster is much wider than the mesofemur ( Fig. 4A ) when compared to the width ratio between the pro- and mesofemur of Montina ( Figs 5B, D ), but this must be tested with accurate measurements in several species. Table 1. Main differential morphological characters between Montina Amyot & Serville, 1843 and Ploeogaster Amyot & Serville, 1843 .

Montina	Ploeogaster
Longitudinal submedial carina of posterior pronotal lobe	Compressed laterally; medial elevated portion truncated or slightly rounded (Figs 21E; 23C).	Less compressed laterally; medial elevated portion produced as a rounded tubercle-like process (Fig. 4B).
Lateral margin of paramedial lobes of posterior margin of pronotum	Entire (Figs 2A; 7B; 13D).	Deeply emarginated (Fig. 4A).
Posterolateral process of the pro- notum	Small, not longer than its base width, sometimes a blunt tubercle (Figs 5D; 13D).	A sharp process, projected laterally, always longer than its base (Fig. 4A).
Profemur	Cylindrical, ventral margin straight (Figs 5A, C).	Wider in the middle, ventral margin slightly curved (Fig. 4C).
Protibia	Straight.	Slightly curved.
Medial process of the pygophore	Nearly cylindrical, directed posteriorly at about 45 degrees, short and narrow, widened at the base, and with a small hook-like projection directed posteriad (Figs 3A–B; 8A–C).	Flattened antero-posteriorly, nearly laminar, directed almost vertically, longer, of subparal- lel margins (Figs 4D, E).
Parameres	Narrow, slightly enlarged apically, gently curved basally (Figs 8A; 10A; 16A).	Wide, enlarged medially, strongly L-shaped (Fig. 4D).

The male genitalia are also useful to delimit these genera. In Montina the medial process of the pygophore ( mpp ) is almost cylindrical, reclined, directed posteriorly at about 45 degrees, short and narrow, sometimes widened at the base, and with a small hook-like projection directed posteriad ( Figs 3A–B ; 8A–C ). In Ploeogaster , on the other hand, the mpp is flattened antero-posteriorly, having thus a laminar form, being much longer than in Montina , of subparallel margins, and directed almost vertically ( Figs 4D, E ). Furthermore, the parameres in Montina are narrow, slightly enlarged apically, and curved basally ( Figs 8A ; 10A ; 16A ), whereas in Ploeogaster they are usually wide and enlarged medially ( Fig. 4D ). Fig. 2. General external morphology of Montina Amyot & Serville, 1843 . A – dorsal view. B – lateral view. Fig. 3. Pygophore and phallus morphology of Montina Amyot & Serville, 1843 . A – pygophore, dorsal view; B – pygophore, lateral view; C – phallus with extended endosoma, ventral view; D – non-extended phallus, ventral view; E – phallus with extended endosoma, dorsal view; F – phallus, dorsal view, detail. Abbreviations: aed – aedeagus, ao – anterior opening of the pygophore, apt – articulatory apparatus, bdl – basal dorsolateral lobes of the endosoma, br – transversal bridge of the pyphohore, ddl – distal dorsal lobe of the endosoma, dll – distal lateral lobes of the endosoma, dps – dorsal phallothecal sclerite, dvl – distal ventral lobe of the endosoma, end – endosoma, ll – lateral lobes of the endosoma, lpp – lateral sclerotization of the phallosoma, mpp – medial process of pygophore, pa – paramere, pha – phallosoma, po – posterior opening of the pygophore, str – struts, vsp – ventral sclerotization of the phallosoma. Scale bars: 1 mm. The structure of the male genitalia is usually useful for species delimitation in Harpactorini (e.g., FORERO et al. 2008 ; ZHANG et al. 2016 ). In Montina , male genitalia are rather uniform, only with small differences between species. Particularly, the male aedeagus has only slight differences among species. Nonetheless, the structure of the medial process of the pygophore ( mpp ) is clearly different among species (e.g., Figs 6A–C ; 10A–C ; 14A–C ). Other characters such as the shape of the margin of the connexivum, color patterns, and the vestiture of the ventral laterotergites and sternites of the abdomen are more conspicuous and are used here as the primary characters to delimit the species. A character that has been used to separate Montina and Ploeogaster is the structure of the connexivum ( AMYOT & SERVILLE 1843 , STÂL 1867 ). Traditionally has been indicated that each of the connexival segments are forming distinct rounded lobes in Montina in opposition to straight margins in Ploeogaster . However, this is not always the case, because in some Montina species the margin of the connexivum is almost straight ( Figs 7A, C ; 9A, C ; 11C ; 19C ; 21A, C ; 23A ) and it could be nearly lobate in some Ploeogaster species. Therefore, the shape of the margin of the connexivum seems an unreliable character to separate these two genera. Despite this, having a lobate margin of the connexivum could help to recognize certain Montina species. Fig. 4. Morphological characters of Ploeogaster Amyot & Serville, 1843 . Ploeogaster sp. , male from Florencia, Caquetá, Colombia. A – dorsal view; B – detail of pronotum; C – lateral view; D – pygophore, dorsal view; E – pygophore, lateral view; F – phallus, ventral view; G – phallus, dorsal view. Scale bar: 1 mm. The value of the aforementioned characters to delimit the two genera is congruent with the results of the phylogenetic analysis of ZHANG & WEIRAUCH (2014) . Among the Neotropical clades recovered in their analysis, there is one monophyletic group containing species that we positively ascribe to Montina , and another one which contains species that we identify as Ploeogaster . This assessment was based on the examination of images of their voucher specimens (http://research.amnh.org/pbi/heteropteraspeciespage/, searched for each taxon and the RCW numbers used in ZHANG & WEIRAUCH 2014 ) and verified against the proposed characters mentioned above to distinguish Montina from Ploeogaster . Although these two groups are not sister lineages in their final topology, as would be the expectation based on the very similar morphology of the pronotum and abdomen, the low support values indicate that the relationships presented must be viewed as unresolved at best, therefore allowing the possibility that these two clades might be sister groups. Furthermore, the specimen UCR_ENT 00001516 (RCW_ 636 in their analysis) that is nested within Ploeogaster species correspond to what has been described as Cidoria Amyot & Serville, 1843 , a monotypic genus known from French Guiana (MALDONADO 1990). In Cidoria , the pronotum exhibit the same pronotal structure as species of Ploeogaster , particularly the presence of a pair of discal tubercles on the posterior lobe of the pronotum and the strongly emarginate paramedial lobes on its posterior margin. The only remarkable feature of Cidoria is the strongly curved medial posterior margin of the pronotum between the paramedial lobes, thus covering almost completely the scutellum in dorsal view, but as indicated above, the shape of the posterior pronotal margin is a variable feature in both Ploeogaster and Montina . Future studies might conclude that Cidoria is congeneric with Ploeogaster . Distribution. Costa Rica , Panama , French Guiana, Guyana , Brazil , Ecuador , and Peru (CHAMPION 189; GIL- SANTANA 2019; MALDONADO 1990 ; STÂL 1859 , 1865 , 1867 , 1872 ). Newly recorded from Colombia . Key to the known species of Montina (for Spanish translation see the Appendix) 1 Connexival margin, at least segments 4 and 5, noticeably lobed ( Figs 13A, C ; 15C ; 19A ); if slightly lobed ( Figs 19C ; 17A ) then general coloration orange with head and legs black or brown ( Figs 19A–D ), or general coloration pale brown with ventral laterotergites dark with a pale-yellow oblique band on the posterior margin of segments 2–6 ( Figs 17A, C ). .......................... 2 – Connexival margin straight or at most slightly lobed ( Figs 7A ; 9A ; 11D ; 21A, C ; 23A ); if segments 4 and 5 are slightly lobed ( Fig. 11A ) then general coloration is brown, with ventral laterotergites dark without pale contrasting areas ( Figs 11A, B ). ............................... 8 2 Ventral laterotergites dark without any pale contrasting areas ( Figs 38A–B ). ..................................................... ................ M. sinuosa ( Lepeletier & Serville, 1825 ) – Ventral laterotergites dark with red or yellow contrasting areas. .................................................................. 3 3 Ventral laterotergites dark with contrasting paleyellow areas, either an oblique band on the posterior margin of segments 2–6 ( Figs 15A, C ; 17A, C ), or on the dorsal margin of segments 2–7 ( Fig. 35A ). ........ 4 – Ventral laterotergites with a red band on the dorsal margin of each segment, sometimes not very conspicuous ( Figs 5A, C ; 13A, C ; 19A, C ), but contrasting areas never yellow. ................................................... 6 4 Connexival margin not deeply lobed, with a short, acute process on the posterior half of segments 2–6, more acute in males ( Fig. 17A ); general coloration pale brown ( Figs 17B, D ). .......... M. ruficornis ( Fabricius, 1803 ) – Connexival margin deeply lobed on segments 4–5 ( Figs 15A, C ; 38B ); general coloration dark to black. ........ 5 5 Pronotum pale-yellow, densely setose ( Fig. 15E ); pale--yellow bands present on the posterior margin of each connexival segment ( Figs 15A–D ). ............................. ................................................. M. lobata Stål, 1859 – Pronotum dark red, with sparse setae ( Fig. 35B ); dorsal margin of connexivum with a pale-yellow, narrow band ( Figs 15A–D ). ................ M. nigripes Stål, 1859 6 Connexival margin with segments 4–6 deeply lobed (both males and females), without a conspicuous acute process on each segment ( Figs 5A, C ); discal tubercles of the anterior lobe of the pronotum not well developed ( Figs 5E–F ). .......................................................... ............... M. calarca Mejía-Soto & Forero sp. nov. – Connexival margin only with segments 4 and 5 lobed, posterior margin of each segment oblique, with a short, acute process on the posterior half ( Figs 13A, C ; 19A, C ); discal tubercles of the anterior lobe of the pronotum subconical and well developed ( Figs 13E, F ; 19E, F ). ............................................................................. 7 7 Head red ( Figs 13B, D, G ); posterior margin of pronotum red; proximal portion of the corium red ( Figs 13B, D ). .......... M. gladiator Mejía-Soto & Forero sp. nov. – Head black ( Figs 19B, D ); posterior margin of the pronotum usually with a transverse dark band connecting the bases of the paramedial lobes; proximal portion of the corium dark ( Figs 19B, D ). .................................... .......................................... M. scutellaris Stål, 1859 8 Forewing membrane with a conspicuous hyaline medial area ( Fig. 32B ). .......... M. fenestrata ( Stål, 1867 ) – Forewing membrane with uniform coloration ( Figs 21B ; 23B ). ................................................................ 9 9 Ventral laterotergites uniformly black, with scattered black, erect setae ( Figs 23E–F ); abdominal sternites entirely black, with decumbent, silver setae ( Fig. 23F ). .......... M. tikuna Mejía-Soto & Forero sp. nov. – Ventral laterotergites with contrasting coloration ( Figs 7A, C ; 9A, C ), if apparently with homogenous coloration ( Figs 11A, C ; 21A ) with numerous decumbent setae and no erect setae; abdominal sternites not entirely black, with silver or black erect setae. .................... 10 10 Overall dorsal coloration and legs red ( Figs 7B, D ; 21B, D ). .................................................................. 11 – Overall dorsal coloration and legs brown to pale brown ( Figs 9B, D ; 11B, D ). ............................................. 12 11 Dorsal laterotergites with segments 2–4 mostly black, 5–7 black with a broad and conspicuous yellow or orange dorsal band increasing in size toward the posterior segments ( Figs 7B, D ), tubercle of the anterior pronotal lobe straight, constricted near the middle and markedly globose apically ( Fig. 7E ). .......................... ........................................... M. confusa ( Stål, 1859 ) – Dorsal laterotergites with segments 3–7 black with a narrow and uniform orange band on their lateral margin ( Figs 21B, D ), tubercle of anterior pronotal lobe slightly curved anteriad, not constricted near the middle ( Fig. 21E ). ...................... M. testacea ( Stål, 1859 ) 12 Ventral laterotergites mostly dark brown ( Figs 11A, C ). ......................................... M. fumosa ( Stål, 1867 ) – Ventral laterotergites with a dorsal, broad, pale-yellow band contrasting with the black ventral area (males on segments 4–5 with black reaching dorsal margin) ( Figs 9A, C ). ................................ M. distincta ( Stål, 1859 )