Revised classification of the New World Cylapini (Heteroptera: Miridae: Cylapinae): taxonomic review of the genera Cylapinus, Cylapoides and Peltidocylapus and a morphology-based phylogenetic analysis of tribe Cylapini Author Wolski, Andrzej text Zootaxa 2021 2021-12-06 5074 1 1 66 journal article 2938 10.11646/zootaxa.5074.1.1 d7238c35-357f-4920-a5a4-2c89c82c85f9 1175-5326 5760387 7B3C6765-F0D2-4846-BB95-200258ECC0E1 Tribe Cylapini sensu Gorczyca (2000) Diagnosis . For details see Gorczyca (2000) and Wolski (2017) . Remarks . The current phylogenetic analysis rendered the tribe Cylapini paraphyletic with respect to the vanniines. The tribe Vanniini , as currently understood, is composed of the Vannius complex sensu Cassis et al. (2003) and the genus Palaucoris Carvalho ( Cassis & Schuh 2012 ; Namyatova et al. 2016). Distant (1883) first noticed the similarity between Vannius and Valdasus , which are currently classified in the tribes Vanniini and Cylapini , respectively. The taxa of Cylapini and Vanniini were united in one tribe, the Cylapini , by Poppius (1909) , based on the long antenna and by Carvalho (1952a , 1955 , 1957 ) based on the vertical head. Gorczyca (1997) excluded the Vannius complex from Cylapinae and moved it to the controversial subfamily Palaucorinae (with a single genus Palaucoris Carvalho ) based mainly on the flattened parempodia found in both taxa and he erected two tribes: the Vanniini (for Vannius complex) and Palaucorini (for Palaucoris ). Cassis et al. (2003) refuted this concept, and the Vannius complex was assigned again to Cylapinae as incertae sedis . Consequently, Gorczyca (2006a) treated these genera again within Cylapinae but placed them among Cylapini . The Vannius complex was given a tribal status within Cylapinae by Cassis & Schuh (2012) . The present analysis, showing vanniines nested well within Cylapini , provides a strong argument for treatment of vanniines within Cylapini , corroborating the classification of Gorczyca (2006a) . On the other hand, given the uncertainties concerning the results of the current analysis (see discussion above) I believe it is premature to relegate the vanniines to Cylapini , and I suggest retaining the concept of Cylapini in its current composition pending studies with larger sampling and emphasis on the molecular data. Notes on the morphological characters in Cylapini . The Cylapini in its present concept includes the genera listed in the Tab. 2 . Herein, I present the discussion of the main characters found in Cylapini taking into account their diversity within the group and comparison with other tribes. Body shape . In Cylapini body is usually elongate to elongate-oval. In some species of Carvalhoma ( C. ovatum Namyatova & Cassis , C. parvum Namyatova & Cassis or C. taplini Namyatova & Cassis ) and in Schizopteromiris the body is ovate ( Schuh 1986 ; Namyatova & Cassis 2016 ). TABLE 2 . Checklist of Cylapini .

Species	Distribution	Complex
Amapacylapus amapariensis Carvalho & Fontes, 1968	Brazil, Ecuador, French Guiana, Guyana	Cylapus complex
Amapacylapis englemani Carvalho, 1991	Panama	Cylapus complex
Amapacylapus labeculosus ( Bergroth, 1922 )	Brazil	Cylapus complex
Amapacylapus nigricapitis Carvalho, 1986	Brazil	Cylapus complex
Amapacylapus rondoniensis Carvalho, 1986	Brazil	Cylapus complex
Amapacylapus unicolor Wolski, 2017	Ecuador	Cylapus complex
Cylapus amazonicus ( Carvalho, 1989 )	Bolivia, Brazil, Ecuador,	Cylapus complex
Cylapus antennatus ( Carvalho & Fontes, 1968 )	Bolivia, Ecuador	Cylapus complex
Cylapus brasiliensis Carvalho, 1986	Brazil	Cylapus complex
Cylapus citus Bergroth, 1922	Bolivia, Brazil, Guyana, Peru	Cylapus complex
Cylapus clavicornis Poppius, 1909	Peru	Cylapus complex
Cylapus famularis (Stål, 1862)	Mexico	Cylapus complex
Cylapus funebris ( Distant, 1883 )	Panama	Cylapus complex
Cylapus luridus Wolski, 2017	Brazil	Cylapus complex
Cylapus marginicollis ( Distant, 1883 )	Nicaragua, Panama	Cylapus complex
Cylapus nobilis Poppius, 1909	Venezuela	Cylapus complex
Cylapus rondoniensis (Carvalho, 1991)	Brazil	Cylapus complex
Cylapus ruficeps Bergroth, 1922	Brazil, Colombia, Ecuador, Venezuela	Cylapus complex
Cylapus stellatus ( Distant, 1883 )	Guatemala	Cylapus complex
Cylapus striatus Reuter, 1907	Venezuela, Brazil, Peru, Bolivia	Cylapus complex
Cylapus tenuicornis (Say, 1932)	Canada, USA	Cylapus complex
Cylapus tucuruiensis ( Carvalho, 1989 )	Brazil, Venezuela	Cylapus complex
Peltidocylapus calyciformis sp. nov.	Ecuador, Peru	Cylapus complex
Peltidocylapus carmelitanus Carvalho & Fontes, 1968	Brazil	Cylapus complex
Peltidocylapus caudatus sp. nov.	Ecuador, Peru	Cylapus complex
Peltidocylapus cerbereus ( Distant, 1883 )	Panama	Cylapus complex
Peltidocylapus ecuadorensis sp. nov.	Ecuador, Venezuela	Cylapus complex
Peltidocylapus festinabundus ( Bergroth, 1922 ) comb. nov.	Peru	Cylapus complex
Peltidocylapus nubilus ( Distant, 1893 )	Panama	Cylapus complex
Peltidocylapus pallidus sp. nov.	Peru	Cylapus complex
Peltidocylapus parallelus sp. nov.	Ecuador, Guyana	Cylapus complex
Peltidocylapus picatus ( Distant, 1893 )	Panama	Cylapus complex
Peltidocylapus politus ( Poppius, 1909 )	Brazil	Cylapus complex
Peltidocylapus rugosus ( Distant, 1883 )	Costa Rica, Guatemala	Cylapus complex
Peltidocylapus scutellaris Poppius, 1909	Costa Rica, Ecuador	Cylapus complex

...continued on the next page TABLE 2 . (continued)

Species	Distribution	Complex
Peltidocylapus simplex sp. nov. Peltidocylapus spinosus sp. nov. Peltidocylapus tapirapensis Carvalho, 1991	Ecuador French Guiana Brazil	Cylapus complex Cylapus complex Cylapus complex
Valdasus bolivianus Carvalho, 1989 Valdasus erebeus Distant, 1883 Valdasus favrei Wolski, Chérot & Carpintero Valdasus ferrerai Wolski, Chérot & Carpintero	Bolivia Panama Brazil, Ecuador, French Guiana French Guiana	Cylapus complex Cylapus complex Cylapus complex Cylapus complex
Valdasus flavinotum Wolski, Chérot & Carpintero Valdasus henryi Wolski, Chérot & Carpintero Valdasus schoenherri Stål, 1860 Valdasus stygius Distant, 1883	Brazil, Ecuador, French Guiana Ecuador Argentina, Brazil Panama	Cylapus complex Cylapus complex Cylapus complex Cylapus complex
Carvalhoma malcolmae Slater & Gross, 1977 Carvalhoma ovatum Namyatova & Cassis, 2016 Carvalhoma parvum Namyatova & Cassis, 2016 Carvalhoma taplini Slater & Gross, 1977 Carvalhoma weiri Namyatova & Cassis, 2013	Australia Australia Australia Australia Australia	N/A N/A N/A N/A N/A
Corcovadocola hypophylla Carvalho, 1948 Corcovadocola itatiaiana Carvalho, 1980 Corcovadocola pilosa Carvalho, 1948	Brazil Brazil Brazil	N/A N/A N/A
Cylapinus minusculus Carvalho, 1986 Cylapinus yasunagai sp. nov.	Brazil, Ecuador, Guyana Bolivia, French Guiana	N/A
Cylapoides bicolor Carvalho, 1952 Cylapoides unicolor Carvalho, 1952	Belize Brazil, Costa Rica, Ecuador, Panama, Venezuela.	N/A N/A
Cylapomorpha gracilicornis Poppius,1913 Cylapomorpha michikoae Yasunaga	Philippines Japan	N/A
Cylapomorpha migratoria (Distant) Cylapomorpha pacifica Carvalho & Lorenzato, 1978	Seychelles Islands Caroline Island: Palau	N/A N/A
Dariella rubrocuneata Namyatova & Cassis, 2021	Australia	N/A
Duckecylapus duckei Carvalho, 1982	Brazil	N/A
Labriella fusca Namyatova & Cassis, 2021	Australia	N/A
Mangalcoris miniatus Murphy &Polhemus, 2012	Singapore, Thailand	N/A
Microcylapus diminutus Carvalho, 1989	Brazil	N/A
Phyllocylapus lutheri Poppius, 1913	Sri Lanka	N/A
Schizopteromiris carayoni Schuh, 1986	New Caledonia	N/A

...continued on the next page TABLE 2 . (continued)

Species	Distribution	Complex
Schizopteromiris lordhowensis Schuh, 1986 Schizopteromiris monteithi Schuh, 1986 Schizopteromiris queenslandensis Schuh, 1986	Lord Howe Is. Australia Australia	N/A N/A N/A
Valdasoides bahiensis Carvalho, 1989	Brazil	N/A

Head . The head in Cylapini is always hypognathous, varying from weakly wider than high or as high as wide with frons sloping, not perpendicular to vertex ( Carvalhoma , Corcovadocola Carvalho , Cylapinus Carvalho , Cylapoides Carvalho , Dariella Namyatova et Cassis , Labriella Namyatova et Cassis , Mangalcoris Murphy et Polhemus , Schizopteromiris Schuh ) ( Figs 8a, b ; Namyatova & Cassis 2016 : figs 2A, C, 3C, 4A, C, 2021: figs 3A, C, 5A, C) to much higher than wide in anterior view with frons strongly flattened, perpendicular to vertex ( Cylapomorpha , Cylapus complex) ( Figs 8c–i , 9a, g ; Yasunaga 2000 : fig. 2, 3; Wolski 2017 : figs 9, 92; Wolski et al. 2020 : fig. 28). In all Cylapini , Vanniini and Bothriomirini the antennal fossa is placed more or less above the suture between the maxillary and mandibular plates ( Namyatova & Cassis 2016 : figs 2C, 4C, 2021: figs 3A, 5A), and the mandibular plate is separated from the remainder of the head by a suture ( Figs 9a–d ; Namyatova & Cassis 2016 : fig. 2C). In contrast, in all fulviines and rhinomirines the antennal insertion is continuous with the suture between maxillary and mandibular plates ( Namyatova & Cassis 2019 : figs 6G, 10E, 22C) and the mandibular plate is not separated from the remainder of head by any suture ( Figs 9e, f ; Wolski & Henry 2012 : fig. 74; Wolski et al. 2018 : fig. 11). Within Cylapini the antennal fossa is either situated close to the suture between maxillary and mandibular plates (all Cylapini except Cylapus complex) (8a, b, i, 9b; Namyatova & Cassis 2016 : figs 2C, 4C, 2021: figs 3A, 5A) or distinctly removed from it ( Figs 8c–h , 9a, g ), and the suture behind the mandibular plate can be faint ( Fig. 9b ; Namyatova & Cassis 2016: 2 C) or strongly depressed ( Figs 9a, g ). All members of Cylapini , Vanniini , and many bothriomirines have the ventral margin of the eyes removed from the ventral margin of head ( Figs 9a–d ; Namyatova & Cassis 2016 : fig. 2C, 2021: 3A; Namyatova et al. 2019: figs 5C, 10D, 11B) and the base of clypeus is positioned below the ventral margin of the eyes ( Figs 9g –i ; Namyatova & Cassis 2016 : fig. 3A, 2021: figs 3C, 5C; Namyatova et al. 2019: figs 5A, 10A, 11A). In most Fulviini and Rhinomirini the ventral margins of eyes are reaching or almost reaching the ventral margin of the head ( Figs 9e, f , Wolski & Henry 2012: fig. 73; Wolski 2013 : fig. 73) and the base of clypeus is situated above the ventral margin of the eyes ( Figs 8l, m ; Wolski 2013 : figs 33–36; Namyatova et al. 2016: fig. 3H; Namyatova & Cassis 2019 : figs 10A, 13C, 16C). Within Cylapini the eyes are situated either relatively close to the ventral margin of head ( Figs 9b ; Namyatova & Cassis 2021 : figs 3A, 5A) or are strongly removed in the dorsal direction ( Fig. 9a,c ; Wolski 2017 : figs 9, 95). The base of clypeus is situated close to the ventral margin of the eyes ( Fig. 9h , 8a, b, i ; Yasunaga 2000 : fig. 2; Namyatova & Cassis 2021: 3 C, 5C) or is strongly removed from the eyes’ ventral margin ( Figs 8c–h , 9g ). The vertex in Cylapini is either carinate posteriorly ( Carvalhoma , Corcovadocola , Cylapoides , Dariella , Labriella ) ( Figs 9h , 11e ; Namyatova & Cassis 2016 , 2021 : figs 3b, 5b) or devoid of carina ( Cylapinus , Cylapomorpha , Mangalcoris , Cylapus complex) ( Figs 11a, g ; Wolski 2017 : fig. 33). The vertex in most Cylapini possesses a more or less developed depression along the midline. The depression is either faint ( Figs 9h , 11a ; Namyatova & Cassis 2016 ; Wolski 2017: 33 ) or strongly developed so the head is V-shaped in anterior view ( Cylapus complex) ( Figs 8c–h , 9g ). The eyes are either embedded into the head, with the dorsal margin situated at the same plane as vertex ( Carvalhoma , Corcovadocola , Cylapoides , Dariella , Labriella , Mangalcoris , Schizopteromiris ) ( Figs 8b , 9h ; Namyatova & Cassis 2016 : fig. 2B, 2021: 3B, 5B) or the eyes are pedunculate ( Cylapomorpha , Cylapinus , Cylapus complex) ( Figs 8a, c–i , 9g ). The latter condition is unique within Cylapinae . Antennae . The shape of antennae is variable among the genera of Cylapini . In Carvalhoma , Corcovadocola , Cylapinus , Cylapoides , Dariella , Labriella and Schizopteromiris the antenna is relatively short, shorter than body length and the segments III and IV are not thread-like ( Figs 3a, b, d ; Namyatova & Cassis 2016 : fig. 1, 2021: fig. 1). In Mangalcoris , Cylapomorpha and Cylapus complex the antennae are longer than the body length and segments III and IV are thread-like (e.g., Figs 4b, c ; Wolski 2017: 40 , 46, 48, 51; Yasunaga 2000 : fig. 1; Gorczyca 2006a : fig. 5; Murphy & Polhemus 2012 : figs 1A, B, D). The latter type of antenna is also found in Vanniini (e.g., Gorczyca 2006a : fig. 7). Labium . In all Cylapini , Vanniini , and Bothriomirini the labium is stout and rather short, the apex usually reaching the hind coxae or barely beyond it, and the segments I and II are not subdivided ( Figs 9j, k ; Namyatova & Cassis 2016 : figs 5D, 6E, 2021: 3D, 5D; Namyatova et al. 2016: figs 9F, G, 2019: 5E, 9J). In contrast, in Fulviini and Rhinomirini the labium is thin and long, sharply pointed, the apex reaching beyond the middle of the abdomen, and segments I and II are subdivided ( Fig. 9l ; Namyatova et al. 2016: figs 9J, 10A; Namyatova & Cassis 2019 : figs 10 R, 13J, 22D). Pronotum . In Cylapini the pronotum is either impunctate ( Carvalhoma , Corcovadocola , Cylapoides , Cylapomorpha , Labriella , Mangalcoris , Schizopteromiris ) ( Figs 9b , 11e ) or punctate ( Cylapinus , Dariella , Cylapus complex) ( Figs 9a , 10a , 11a, g ). The lateral margin is always ecarinate ( Figs 9a, b , 10a , 11b ). The collar in Cylapini is always present, and in lateral view is situated anteriorly to the propleural suture ( Fig. 9a ). FIGURE 3. Dorsal habitus photographs. a . Cylapinus minusculus (♂, Ecuador); b . Cylapinus yasunagai (holotype); c . Cylapoides bicolor (holotype); d . Cylapoides unicolor (♂, Ecuador). FIGURE 4. Dorsal habitus photographs. a . Peltidocylapus calyciformis (paratype, ♂); b . Peltidocylapus caudatus (holotype); c . Peltidocylapus cerbereus (lectotype); d . Peltidocylapus ecuadorensis (holotype); e . Peltidocylapus festinabundus (♀, Peru). FIGURE 5. Dorsal habitus photographs. a . Peltidocylapus nubilus (lectotype); b . Peltidocylapus pallidus (holotype); c . Peltidocylapus parallelus (holotype); d . Peltidocylapus picatus (holotype); e . Peltidocylapus politus (holotype). FIGURE 6. Dorsal habitus photographs. a . Peltidocylapus rugosus (lectotype); b . Peltidocylapus scutellaris (♂, Costa Rica); c . Peltidocylapus simplex (holotype); d. Peltidocylapus spinosus (holotype). Thoracic pleura . The mesepimeral spiracle in Cylapini is slit-like ( Figs 10b–e , 11c, f ), in members of the Cylapus complex it is covered with evaporative bodies ( Figs 10b–e ). In most Cylapini taxa except for the Cylapus complex, the metepisternum is narrow and rectangular ( Figs 11c, f ; Namyatova & Cassis 2016 : fig. 2K, 2021: figs 3F, 5H). In the Cylapus complex the metepisternum is broad and almost square ( Figs 10b–e ). In most Cylapini genera the metepisternum is carinate posteriorly ( Figs 10b–e , 11c, f, i ). The metepisternal evaporative areas are either relatively narrow, straight posteriorly and not extended onto anterior margin of metepisternum ( Carvalhoma , Corcovadocola , Cylapinus , Cylapoides , Dariella , Labriella , Cylapomorpha ) ( Figs 11c ; Namyatova & Cassis 2016 : fig. 2K, 2021: figs 3F, 5H; Wolski 2017 : fig. 35) or are broad, rounded, extended posteriorly, and extended well onto anterior margin of metepisternum ( Cylapus complex) ( Figs 10b–e ). In many genera of the Cylapini the metepimeron is well exposed, not obscured by second abdominal segment ( Figs 10b–e , 11c, f ). The ostiolar peritreme is usually oval ( Figs 10d–e , 11c, f ), rarely the peritreme is narrow and ear-like ( Amapacylapus ) ( Fig. 10b ) or strongly protruding, thin and arcuate and sharply pointed ( Cylapus ) ( Fig. 10c ). FIGURE 7. Lateral view. a . Cylapinus minusculus (♀); b . Cylapinus yasunagai (paratype); c . Cylapoides unicolor (♀); d . Peltidocylapus calyciformis (paratype); e . Peltidocylapus cerbereus (lectotype). Hemelytron . The hemelytron in Cylapini is either impunctate ( Corcovadocola , Cylapoides , Labriella , Cylapomorpha ) ( Fig. 11e ; Wolski 2017 : fig. 33; Namyatova & Cassis 2021 : fig. 1) or covered with deep and dense punctation ( Carvalhoma , Cylapinus , Dariella , Cylapus complex) ( Figs 11a, g, m ; Namyatova & Cassis 2016 : fig. 1, 2021: fig. 1). Most Cylapini genera are macropterous, and in such genera as Carvalhoma , Corcovadocola and Mangalcoris the hemelytron is further modified (see Wolski & Gorczyca 2014 and Namyatova & Cassis 2019 for detailed discussion on the wing modification in Cylapinae ). Legs . In most Cylapini the legs are not modified; only in Phyllocylapus lutheri Poppius, 1913 the fore tibiae are strongly broadened and leaf-like ( Gorczyca 2000 : fig. 15). Most Cylapini possess moderately elongated legs. In the Cylapus complex the legs, especially hind legs, are strongly elongated ( Figs 4c , 6a ; Wolski 2017 : figs 37, 41, 55). In all Cylapini the tarsus is three-segmented. Tarsal segment I is usually shorter than II and III combined ( Figs 10j , 11d, j ; Namyatova & Cassis 2021 : fig. 3J, 5G), rarely I segment is as long as or longer than segments II and III combined ( Carvalhoma , Cylapus ) ( Fig.10i ; Namyatova & Cassis 2016 : fig. 2I). Male genitalia . The pygophore in Cylapini has no supragenital bridge sensu Konstantinov 2003 , instead the ventral wall is much longer than the dorsal wall, and the genital opening is directed upward ( Yasunaga 2000 : fig. 5; Namyatova & Cassis 2016 : figs 7C, 8C, 2021: figs 4C, 6C; Wolski 2017 : figs 17, 134). The parameres in Cylapini are similar in size, the left paramere is usually C-shaped, and the right paramere is usually sickle-shaped and frequently with more or less shortened apical process ( Figs 12b, c, h, i, m, n ; Yasunaga 2000 : fig. 5; Namyatova & Cassis 2016 : figs 7D, E, 8D, E, 2021: figs 4D–G, 6C–G). The aedeagus is moderately voluminous, not subdivided into the vesica and conjunctiva. The phallotheca is moderately sclerotized. Ductus seminis relatively short and thick ( Figs 12a, g, j , 13m ; Namyatova & Cassis 2016 : figs 7A, B, 8A, B, 2021: figs 4A, B, 6A,B; Wolski 2017 : figs 12, 66; Wolski et al. 2020 : fig. 40, 48). The endosoma in Cylapini is either furnished with sclerites ( Fig. 14r ; Wolski 2017 : figs 12, 66; Wolski et al. 2020 : fig. 48; Namyatova & Cassis 2021 : fig. 6) or devoid of spiculi ( Figs 12j, p , 13a, g, m ; Wolski et al. 2020 : fig. 40). Sometimes the endosoma is composed of strongly inflated sclerotized lobes ( Fig. 12g ; Yasunaga 2000 : fig. 6). Female genitalia . The ovipositor in the Cylapini is reminiscent of that found in Vanniini and Bothriomirini . Its first gonapophyses have the ventral margin more or less arcuate, dorsal margin sinuate, strongly convex subapically ( Figs 21e–i ; Yasunaga & Miyamoto 2006 : fig. 4B; Wolski et al. 2020 : figs 55, 59, 67, 69). The second gonapophyses are ventrally arcuate, with the dorsal margin weakly sinuate and strongly serrate ( Figs 21m –q , 22i, j ; Wolski et al. 2020 : figs 56, 60, 68, 70). In contrast, in most known Fulviini the gonapophyses are rounded, without tooth-like structures or with weakly developed denticles ( Figs 22k, l ; Sadowska-Woda et al. 2006 : figs 2, 3, 4, 12, 13; Kim et al. 2019 : figs 2D, 4D; Gorczyca et al. 2020 : figs 5c, d). The first gonapophyses are often connected by well-developed, membranous structure (e.g., Sadowska-Woda et al. 2006 : figs 2, 3, 12, 13; Gorczyca et al. 2020 : figs 5c, d) and together with the second gonapophyses they form a dilatable ovipositing tube ( Schmitz & Štys 1973 ). The latter authors suggested that such a shape of the ovipositor allow fulviines to deposit eggs into the cavities tree bark where they live, indicating that they are predators. This hypothesis may be supported by the fact that most fulviines (unlike Cylapini , Vanniini , and Bothriomirini ) have long, sharply pointed labium with subdivisions on segments I and II (which may make it highly flexible (van Doesburg 1985)), allowing these bugs for effective hunting. Moreover, direct evidence for predation exists only for fulviines and not for the representatives of the other tribes (see Introduction). The vestibulum in Cylapini is either membranous and without any structures encircling vulva ( Figs 20b, c ), or it possesses a well-developed, elongated sclerite adjacent to the base of the first gonapophyses ( Figs 20a, d–h ). The bursa copulatrix in Cylapini is voluminous, laterally extending beyond first gonapophyses ( Figs 15a, b, f, h ; 16a, d, g , 17j , 18b , 19a ; Namyatova & Cassis 2016: 7 F, 9F; Wolski et al. 2020 : fig. 66), and with the anterior margin strongly removed from the first gonapophyses in many cases ( Cylapus complex) ( Figs 15a, b, f, h ; 16a, d, g , 17j , 18b , 19a ; Wolski et al. 2020 : fig. 66). The posterior wall of the genital chamber is simple and membranous ( Figs 19f, g ; Namyatova & Cassis 2019 : figs 7G, 9G, 2021: fig. 7c; Wolski et al. 2020 : fig. 54). The shape and position of the sclerotized rings are variable within the Cylapini . They can be paired ( Figs 15f, g, h , 16g , 18c , 19a, d ) or unpaired ( Figs 15a, b , 16a, d , 17a, d, h, k ). In some genera they can be minute, occupying a small portion of the dorsal wall (roof) of the genital chamber ( Cylapoides , Labriella ) ( Fig. 15g ; Namyatova & Cassis 2021 : fig. 7A) while in other taxa the rings embrace anterior and lateral portions of the genital chamber ( Cylapus , some species of Valdasus ) ( Figs 16a, d, g , 17a, d, h, k ; Wolski et al. 2020 : Figs 57, 63, 65) or occupies most of it ( Amapacylapus , Peltidocylapus , some Valdasus ) ( Figs 15a, b , 18c , 19b, e ). The lateral oviducts are situated centrally on the roof of the genital chamber roof. They are usually contiguous and, rarely they are separated ( Cylapinus minusculus ) ( Fig. 15f ). The lateral oviducts are either wide ( Cylapus complex) ( Figs 15a , 16d, g , 18a, b , 19b, e ) or thin ( Cylapinus minusculus , Cylapomorpha , Cylapoides unicolor ) ( Figs 15f, g, h ; Yasunaga 2000 : Fig. 7 ). They can be short (as in Amapacylapus , Peltidocylapus , and some species of Valdasus ) ( Figs 15a, b , 18c , 19b, e ) or long, reaching beyond lateral or posterior portion of the genital chamber ( Cylapus , some species of Valdasus , Cylapomorpha , Cylapinus , Cylapoides ) ( Figs 15f, g , 16d, g , 17d, k ; Yasunaga 2000 : fig. 7). The spermathecal gland opens centrally in the genital chamber, between the lateral oviducts ( Figs 15a, b, f, h , 18a , 19b, e ; Wolski et al . 2020: 52 , 57), or it rarely it opens in the posterior area of the bursa copulatrix (some species of Cylapus ) ( Figs 16d , 17d ). The dorsal sac sensu Kullenberg (1947) is absent in most studied Cylapini . The representative of Cylapomorpha investigated herein possesses the medio-longitudinal, U-shaped, membranous, structure ( Fig. 15h ), similar to that observed in Orthotylini (e.g., Pluot-Sigwalt & Matocq 2017 : figs 5, 6). In all investigated species of Cylapus and some Valdasus ( V. henryi Wolski, Chérot et Carpintero , V. schoenherri Stål ) the posterior portion of bursa copulatrix is furnished with a more or less developed, transverse sclerotization connecting the posterior portion(s) of the sclerotized ring(s) ( Figs 16b, e, h , 17f, h, k, l ; Wolski et al. 2020 : figs 63, 64, 65). In anterior view this structure clearly forms a pouch or pocket, and it is reminiscent of the pouch-like structures found in Hallodapini ( Pluot-Sigwalt & Matocq 2017 : fig. 12). It is noteworthy that species investigated here also possess the endosoma furnished with at least three sclerites ( Wolski 2017 : figs 66, 71, 80, 97, 102, 135, 14). At the same time, all studied Cylapus complex species without dorsal sac have the endosoma without sclerites ( Peltidocylapus ) ( Figs 12j , 13a, m , 14a, f, e, l ) or with a single, weakly developed sclerotized appendage ( Amapacylapus ) ( Wolski 2017 : figs 12, 18). This observation may support the hypothesis of Pluot-Sigwalt & Matocq (2017) who suggested the role of the dorsal sac during copulation when some portions of aedeagus and parameres penetrate the gynatrium and probably enter and anchor into the cavity of the dorsal sack sac.