Cylapini Kirkaldy, 1906

Wolski, Andrzej, 2017, Taxonomic review of the plant bug genera Amapacylapus and Cylapus with descriptions of two new species and a key to the genera of Cylapini (Hemiptera: Heteroptera: Miridae), Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 57 (2), pp. 399-455 : 401-405

Cylapini Kirkaldy, 1906

Cylapini Kirkaldy, 1906: 134 . Type genus Cylapus Say, 1832 .

Diagnosis. Members of the tribe are recognized by the following characters: dorsal surface smooth or punctate, covered with simple setae, without ornamentation composed of tiny and dense tubercles ( Fig. 128); head hypognathous ( Figs 6, 7, 9 , 34, 56–65 , 108 , 123, 148, 150 ); vertex more or less sulcate along midline, anterior portion of vertex perpendicular to the rest of vertex ( Figs 6, 7–9, 28, 33, 34, 56–65 , 91, 92, 95, 107, 108 , 120, 123, 148, 150 ); posterior margin of maxillary and mandibular plates situated at the same line as anterior margin of eye; maxillary plate much narrower than mandibular plate; ventral margin of eye not reaching or barely reaching base of mandibular plate; buccula ringlike; eye projected above plane of vertex ( Figs 9 , 34, 91, 123, 148, 150 ); eye with short and sparse interocular setae and flat ommatidia ( Fig. 124); antenna threadlike, long, usually as long as or longer than body length, with segments III and IV longer than II ( Figs 2, 3, 36 , 42, 45, 46, 52); labial segments I and II not subdivided ( Figs 9 , 95, 125, 126, 150 ); pronotal collar distinct, narrow, dorsally and laterally delimited by the rather deep depression placed anteriorly to the propleural suture ( Figs 8, 9 , 28, 31, 33, 34); mesepimeral spiracle (msp) usually elongate and slitlike, surrounded by mushroom bodies ( Figs 10 , 32, 35, 93, 96, 108 , 129, 146, 149, 151 ); metathoracic scent gland evaporative area (ea) oval, broad, well expanded onto lateral margin of metepisternum; metepisternum with posterior carina (pc) ( Figs 10 , 32, 35, 93, 96, 108 , 129, 146, 149, 151 , arrow); peritreme (per) more or less raised from the evaporative area ( Figs 10 , 32, 35, 93, 96, 108 , 129, 146, 149, 151 , arrow); genital capsule with dorsal wall short, much shorter than ventral wall, opening oriented upward ( Figs 17 , 134); proctiger distinctly developed and strongly sclerotized ( Fig. 134); vestiture on dorsal surface of the paramere body absent (e.g. Figs 14, 16, 20, 22 , 68, 70, 73, 75 ); ductus seminis usually relatively broad and short; secondary gonopore clearly present, well developed ( Figs 12, 18 , 66, 71 , 80, 85 , 97, 102 , 110, 115 , 135, 140 ).

Discussion. The phylogenetic analysis conducted by GORCZYCA (2000) proposed that Rhinomirini is the closest relative of Cylapini with both linked primarily by the long antenna. This analysis, however, did not include the Vannius Distant, 1883 complex, a group of genera with vertical head and long antennae similar to those found in Cylapini , which was postulated to be related to the genus Palaucoris Carvalho, 1956 and transferred to the subfamily Palaucorinae ( GORCZYCA 1997) based on the spatulate parempodia found in both taxa. Subsequently Vannius complex was restored to Cylapinae by CASSIS et al. (2003) and the phylogenetic analyses of the genera of Vannius complex ( CASSIS et al. 2003, CASSIS & MONTEITH 2006) revealed close relationship between Cylapini and Vannius complex based mostly on the vertical head, showing a member of Cylapini as their sister group. CASSIS & SCHUH (2012) recognizing the tribe Vanniini , noted the close similarity between Cylapini and Vanniini with both sharing the vertical head. KONSTANTINOV (2013), supporting the views of GORCZYCA (1997), suggested Palaucoris as a sister group of Vannius complex within Cylapinae paying attention that the vertical head occurring in both taxa being typical to Cylapini . The close relationship of Cylapini and Vanniini was also revealed by NAMYA- TOVA et al. (2016). Their phylogenetic analysis of the subfamily Bryocorinae , including several cylapines as outgroup taxa, showed monophyly of a group containing Cylapini (Amapacylapus) , members of the Vannius complex, and Palaucoris and they proposed to place the latter in the tribe Vanniini . The information obtained from the present study and the literature data indicate that some of the synapomorphies linking Cylapini and Vanniini presented by NAMYATOVA et al. (2016) occur only in the latter tribe and are not found in Cylapini . For example, the gula, oriented vertically in Vannius complex and Palaucoris ( CASSIS et al. 2003: Figs 1A, 2B; KONSTANTINOV 2013: Fig. 4D; NAMYATOVA et al. 2016: Figs 7A, B), in the members of Cylapini is horizontal or subhorizontal ( Figs 9 , 92, 123, 148, 151 ). While in the Cylapini the pronotal collar is rather thin, dorsally and laterally delimited by the rather deep depression placed anteriorly to the propleural suture ( Figs 8, 9 , 28, 31, 33, 34) in Vannius complex and Palaucoris the collar is flat and widely delimited by the shallow depression extending directly to propleural suture at sides ( KONSTANTINOV 2013: Figs 4D, E; NAMYATOVA et al. 2016: Figs 5A, B). Other characters shown as synapomorphies for Cylapini and Vanniini also occur in other cylapine tribes. These include the head with ventral margin of the eye not reaching maxillary plate which is found also in Bothriomirini ( WOLSKI & GORCZYCA 2012: Fig. 20 ; WOLSKI 2012: Fig. 3D) and the tibiae without black spinules that are also present in Bothriomirini , and are mosaically spread among Fulviini and Rhinomirini (Wolski, pers. observ.). Of the characters presented by the authors as synapomorphic for Cylapini + Vanniini only the ringlike buccula, tightly binding labial segment I (NAMYATOVA et al. 2016: Figs 7A, B) and the distinctly elongate mandibular plate (NAMYATOVA etal. 2016: 7A, B) are found exclusively in both groups not being found in other cylapines. More detailed phylogenetic and morphological studies of the subfamily Cylapinae are needed as indicated by NAMYATOVA et al. (2016) to clarify the relationship between Cylapini and Vanniini .

Concerning the characters herein presented as diagnostic for Cylapini the following features are present also in the Vanniini (sensu NAMYATOVA et al. 2016) :

i) Head hypognathous ( Figs 9 , 34, 92, 95, 108 , 123, 148, 150 ; NAMYATOVA et al. 2016:

7A, B). ii) Vertex more or less sulcate along midline ( Figs 8 , 28, 33, 107 , 120; CASSIS et al. 2003:

Fig. 2A; KONSTANTINOV 2013: Fig. 4E; NAMYATOVA et al. 2016: 5A, B). iii) Frons vertical and flat, perpendicular to horizontal vertex. iv) Posterior margin of maxillary and mandibular plates situated at the same line as middle of eye. v) Mandibular plate distinctly elongate ( Figs 9 , 34, 92, 95, 108 , 123, 148, 150 ; NAMYA-

TOVA et al. 2016: 7A, B). vi) Ventral margin of eye not reaching base of mandibular plate. vii) Buccula ringlike, binding labial segment I ( Figs 6, 7, 9 , 34, 56–65 , 108 , 123, 148,

150; CASSIS et al. 2003: Figs 1A, 2 AB; KONSTANTINOV 2013: Figs 4D, E). viii) Eye with short and sparse interocular setae and flat ommatidia ( Fig. 124; NAMYATOVA et al. 2016: Figs 7A, B). ix) Antenna long, threadlike ( Figs 2, 3, 36 , 42, 45, 46, 52; CASSIS et al. 2003: Figs 3A–D;

KONSTANTINOV 2013: Figs 1B, C). x) Labial segments I and II not subdivided ( Figs 9 , 95, 125, 126, 150 ; NAMYATOVA et al.

2016: Figs 9F, G ). xi) Pronotal collar distinct ( Figs 8, 9 , 28, 31, 33, 34; NAMYATOVA et al.: Figs 5A, B). xii) Genital capsule with dorsal wall short, much shorter than ventral wall, opening oriented upward ( Figs 17 , 134; CASSIS et al. 2003: Figs 1H, 2F; KONSTANTINOV 2013: Fig 2M). xiii) Vestiture on dorsal surface of the body of left and right paramere absent ( Figs 14,

16, 20, 22, 68, 70, 73, 75 ; CASSIS et al. 2003: Figs 5A, B, 7A, B; KONSTANTINOV 2013:

Figs 2A–I).

Of these, only the features iii–v and vii seem to be found exclusively in Cylapini and Vanniini .Although hypognathous head is found also in all Bothriomirini , in Psallopini , and several representatives of Fulviini and Rhinomirini , in these taxa the frons is very gently sloping, not being perpendicular to the vertex, the maxillary and mandibular plates are projected forward with their posterior margin being situated at the same line as anterior margin of eye, and the mandibular plate is about the same size as maxillary plate (e.g. WOLSKI 2010: Fig. 5A; WOLSKI & GORCZYCA 2012: Fig. 20 ; WOLSKI & HENRY 2015: 6, 7; NAMYATOVA et al. 2016: Fig. 7C). In most Cylapini and Vanniini , except for such genera as Carvalhoma Slater & Gross, 1977 and Kanakamiris Cassis & Monteith, 2006 the horizontal frons forms right or acute angle with flat and vertical frons ( Figs 9 , 34, 123, 148, 150 ; KONSTANTINOV 2013: Fig. 4D, F; NAMYATOVA et al. 2016: 7A, B). The mandibular and maxillary plates in most Cylapini and Vanniini are situated below eyes, their posterior margin located at the same line as middle of eye ( Figs 9 , 34, 123, 148, 150 ; KONSTANTINOV 2013: Fig. 4D; NAMYATOVA et al. 2016: Figs 7A, B).

Such characters as the eye with ventral margin not reaching base of the mandibular plate and the glabrous body of the parameres is found apart from Cylapini and Vanniini also in Bothriomirini ( WOLSKI 2013a: Figs 3D, 6B, C; WOLSKI & GORCZYCA 2012: Figs 20 , 49, 59, 62, 63 , 81, 82 , 100, 101 ). In members of Fulviini , Psallopini , and Rhinomirini the ventral margin of eye is reaching gula often wrapping around base of antenna (e.g. MOULDS & CASSIS 2006: Figs 1A–D; WOLSKI 2010: Figs 5A, 7A, 16A; WOLSKI & HENRY 2015: 6, 7; NAMYATOVA et al. 2016: Fig. 7C) and the body of both parameres is covered by setae (e.g. STONEDAHL & KOVAC 1995: Figs 10, 11 , 13 ; GORCZYCA 2002: Figs 1–8; WOLSKI 2010: Figs 8C, H ).

The threadlike, long antenna, with segments III and IV longer than II is also found in the representatives of the tribe Rhinomirini ( GORCZYCA 2000; WOLSKI 2010: Figs 1, 2A, D, E–G, Q) and this character was presented by GORCZYCA (2000) as synapomorphy for Cylapini + Rhinomirini . Another character occurring only in Cylapini , Vanniini , and Rhinomirini not found in other tribes is the more or less developed longitudinal sulcus along midline of vertex ( WOLSKI 2010: Fig. 16A ).

The labium with segment I undivided is present also in all Bothriomirini (Wolski, pers. observ.), the Rhinomirini belonging to the Rhinocylapus Poppius, 1909 group (sensu WOL- SKI 2010) (Wolski, pers. observ.), and a few fulviines, such as genera Hemiophthalmocoris Poppius, 1912 , Xenocylapus Bergoth, 1922 , and Henryfulvius Wolski, 2015 (WOLSKI 2015: Figs 34, 35, 66 ; WOLSKI et al. 2016: Fig. 4B). In the Psallopini ( WOLSKI & HENRY 2015: Figs 18, 19 ), most fulviines ( WOLSKI & HENRY 2015: Figs 20–24 ), and Rhinomirini belonging to the Rhinomiris group (sensu GORCZYCA & CHÉROT 1998) and the genera Rhinomiriella Gorczyca, 2001 and Pararhinomiris Gorczyca, 2003 the labial segment I is subdivided ( WOLSKI & HENRY 2015: Fig. 25 ; WOLSKI et al. 2017). The undivided labial segment II was also observed in Bothriomiris Kirklady, 1902 (NAMYATOVA et al. 2016) and other Bothriomirini (Wolski, pers. observ.), rhinomirines belonging to the genus Rhinocylapus (NAMYATOVA et al. 2016) and other representatives of the Rhinocylapus group (sensu WOLSKI 2010) (Wolski, pers. observ.) and some Fulviini . The subdivided labial segment II is common throughout the Fulviini having been observed among others in Xenocylapus (VAN DOESBURG 1985: Fig. 5) and Peritropis Uhler, 1891 (NAMYATOVA et al. 2016: 10A). It is found also in Psallopini (NAMYATOVA et al. 2016: Fig. 10 C ) and the rhinomirines belonging to Rhinomiris group (sensu GORCZYCA & CHÉROT 1998) and the genera Rhinomiriella and Pararhinomiris (Wolski pers. observ.).

The Cylapini are best distinguished from the Vanniini by the setiform parempodia which in Vanniini are flattened, a character that is not found in other cylapines and is discussed among others by GORCZYCA (1997), CASSIS et al. (2003), and NAMYATOVA et al. (2016). NAMYATOVA et al. (2016) paid attention that the unguitractor plate with three narrow columns with acute lamellae of the central column ( Figs 30, 79 ; NAMYATOVA et al. 2016: Fig. 21E ) occurs in all the examined cylapine taxa and are not found in Vannius complex and Palaucoris (NAMYATOVA et al. 20E, 21D). These authors also noticed that of all the psallopine and cylapine taxa they examined, except Vannius complex, possess the asymmetrical parempodia (NAMYATOVA et al. 2016: Figs 21E ). In Cylapinae , however, the asymmetrical parempodia are present only in some, being in other taxa symmetrical, fully developed ( Figs 30, 79 , 133, 147 ), or reduced as shown by WOLSKI (2010: Fig. 16H ).

The Cylapini and Vanniini can be also distinguished from each other based on the gula orientation, the shape of the collar (see above), the metathoracic scent gland evaporative area development (well developed, expanded onto lateral margin of metepisternum in Cylapini , reduced to the ventral portion of metepisternum in Vanniini ) ( Figs 10 , 32, 35, 93, 96, 129, 146, 149, 151 ; CASSIS et al. 2003: Figs 1B, 2C; KONSTANTINOV 2013: Fig. 4G; NAMYATOVA et al. 2016: Figs 15D, E ), and structure of the posterior margin of metepisternum (carinate in Cylapini , ecarinate in Vanniini ) ( Figs 10 , 32, 35, 93, 96, 129, 146, 149, 151 , arrow; CASSIS et al. 2003: Figs 1B, 2C; KONSTANTINOV 2013: Fig. 4G; NAMYATOVA et al. 2016: Figs 15D, E ).

The key provided below does not include three genera described by CARVALHO (1982, 1989), i.e. Duckecylapus Carvalho, 1982 , Microcylapus Carvalho, 1989 , and Valdasoides Carvalho, 1989 as I did not have an access to specimens belonging to these genera. Based on the punctate dorsum ( CARVALHO 1982, 1989) Duckecylapus and Valdasoides would run to the couplet “3” of the key. Duckecylapus with long and erect setae seems to be most similar to Valdasus . Valdasoides is also similar to Valdasus in having the long and erect setae but it also has the raised metathoracic scent gland peritreme ( CARVALHO 1989) which may indicate its close similarity to Cylapus as diagnosed in this present paper. Microcylapus with the impunctate body would best run to the couplet “7” of the key. From other genera with smooth dorsum it can be distinguished by the convex basal portion of the hemelytral radial vein ( CARVALHO 1989).

MURPHY & POLHEMUS (2012) and NAMYATOVA & CASSIS (2016) included the genera Mangalcoris Murphy & Polhemus, 2012 and Carvalhoma in the tribe Cylapini . Their placement was based mostly on the hypognathous head and long antenna ( NAMYATOVA & CASSIS 2016). Both genera significantly differ, however, from other Cylapini by the frons that is very gently sloping, not being perpendicular to the vertex, the maxillary and mandibular plates are projected forward, situated at the same line as anterior margin of eye, and the mandibular plate is about the same size as maxillary plate (see above). The placement of these genera in Cylapini requires further investigations. Nevertheless, both genera are included in the key.

Figures

Figs 8–11. Scanning electron micrographs of Amapacylapus amapariensis Carvalho & Fontes, 1968: 8 – dorsal habitus; 9 – lateral view; 10 – thoracic pleura; 11 – protarsus.Abbreviations: ea = evaporative areas; msp = metathoracic spiracle; pc = posterior carina; per = peritreme.

Figs 56–65. Lateral view of Cylapus species: 56 – C. amazonicus Carvalho, 1989 (J); 57 – C. antennatus Carvalho & Fontes, 1968 (J); 58 – C. citus Bergroth, 1922 (♀); 59 – C. luridus sp. nov. (J); 60 – C. marginicollis (Distant, 1883) (J); 61 – C. ruficeps Bergroth, 1922 (♀); 62 – C. stellatus (Distant, 1883) (♀); 63 – C. striatus Reuter, 1907 (♀); 64 – C. tenuicornis (Say, 1832) (J); 65 – C. tucuruiensis Carvalho, 1989 (J).

Figs 107–109. Scanning electron micrographs of Cylapus striatus Reuter, 1907. 107 – dorsal habitus (♀); 108 – lateral view (♀); 109 – pretarsal structure. Abbreviations: ea = evaporative areas; msp = metathoracic spiracle; pc = posterior carina; per = peritreme.

Figs 146–151. Scanning electron micrographs of Cylapus tucuruiensis Carvalho, 1989 (146, 147), Peltidocylapus scutellaris (Poppius, 1909) (148, 149), and Valdasus sp. (150, 151): 146, 149, 151 – thoracic pleura; 147 – pretarsal structure; 148, 150 – lateral view.Abbreviations: ea = evaporative areas; msp = metathoracic spiracle; pc = posterior carina; per = peritreme.

Figs 36–40. Dorsal habitus photographs of Cylapus species: 36 – C. amazonicus Carvalho, 1989 (J); 37–39 – C. antennatus Carvalho & Fontes, 1968 (37 – J holotype, 38 – J, 39 – ♀); 40 – C. citus Bergroth, 1922 (J).

Figs 12–22. Male genitalia of Amapacylapus amapariensis Carvalho & Fontes, 1968 (12–16) and A. unicolor sp. nov. (17–22): 12, 18 – endosoma; 13, 19 – left paramere (dorsal view); 14, 20 – left paramere (right lateral view); 15, 21 – apical process of left paramere; 16, 22 – right paramere (left lateral view); 17 – genital capsule. Abbreviations: bpr = basal process; es = endosomal sclerite; pb = paramere body; pr = proctiger; ps = paramere socket; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 66–75. Male genitalia of Cylapus amazonicus Carvalho, 1989 (66–70) and Cylapus antennatus Carvalho & Fontes, 1968 (71–75): 66, 71 – endosoma; 67, 72 – left paramere (dorsal view); 68, 73 – left paramere (right lateral view); 69, 74 – apical process of left paramere; 70, 75 – right paramere (left lateral view). Abbreviations: bpr = basal process; dss = sclerotized portion of ductus seminis inside endosoma; es1–4 = endosomal sclerites 1–4; pb = paramere body; sg = secondary gonopore; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 80–89. Male genitalia of Cylapus citus Bergroth, 1922 (80–84) and C. luridus sp. nov. (85–89): 80, 85 – endosoma; 81, 86 – left paramere (dorsal view); 82, 87 – left paramere (right lateral view); 83, 88 – apical process of left paramere; 84, 89 – right paramere (left lateral view). Abbreviations: bpr = basal process; dss = sclerotized portion of ductus seminis inside endosoma; es1–4 = endosomal sclerites 1–4; pb = paramere body; sg = secondary gonopore; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 97–106. Male genitalia of Cylapus marginicollis (Distant, 1883) (97–101) and C. ruficeps Bergroth, 1922 (102–106): 97, 102 – endosoma; 98, 103 – left paramere (dorsal view); 99, 104 – left paramere (right lateral view); 100, 105 – apical process of left paramere; 101, 106 – right paramere (left lateral view). Abbreviations: bpr = basal process; dss = sclerotized portion of ductus seminis inside endosoma; es1–4 = endosomal sclerites 1–4; pb = paramere body; sg = secondary gonopore; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 110–119. Male genitalia of Cylapus stellatus (Distant, 1883) (110–114) and C. striatus Reuter, 1907 (115–119): 110, 115 – endosoma; 111, 116 – left paramere (dorsal view); 112, 117 – left paramere (right lateral view); 113, 118 – apical process of left paramere; 114, 119 – right paramere (left lateral view). Abbreviations: bpr = basal process; dss = sclerotized portion of ductus seminis inside endosoma; es1–4 = endosomal sclerites 1–4; pb = paramere body; sg = secondary gonopore; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 135–144. Male genitalia of Cylapus tenuicornis (Say, 1832) (135–139) and C. tucuruiensis Carvalho, 1989 (140–144): 135, 140 – endosoma; 136, 141 – left paramere (dorsal view); 137, 142 – left paramere (right lateral view); 138, 143 – apical process of left paramere; 139, 144 – right paramere (left lateral view). Abbreviations: bpr = basal process; dss = sclerotized portion of ductus seminis inside endosoma; es1–4 = endosomal sclerites 1–4; pb = paramere body; sg = secondary gonopore; sl = sensory lobe. Scale bars: 0.1 mm.

Figs 23–26. Hemelytron of the species of Amapacylapus Carvalho & Fontes, 1968. 23 – A. amapariensis Carvalho & Fontes, 1968; 24 – A. englemani Carvalho, 1991; 25 – A. labeculosus Bergroth, 1922; 26 – A. unicolor sp. nov. Abbreviations: p1–p7 = yellowish patches; pc = posterior carina; per = peritreme.

Figs 76–79. Scanning electron micrographs of Cylapus antennatus Carvalho & Fontes, 1968 (76–77) and C. citus Bergroth, 1922 (78–79): 76–77 – antennal segment II; 78 – metafemur; 79 – pretarsal structure.