Cylapini sensu Gorczyca (2000)

Wolski, Andrzej, 2021, 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, Zootaxa 5074 (1), pp. 1-66 : 18-27

publication ID

https://doi.org/10.11646/zootaxa.5074.1.1

publication LSID

lsid:zoobank.org:pub:7B3C6765-F0D2-4846-BB95-200258ECC0E1

DOI

https://doi.org/10.5281/zenodo.5784429

persistent identifier

https://treatment.plazi.org/id/039587FB-AE75-FF90-FF51-15F9437CFEFC

treatment provided by

Plazi

scientific name

Cylapini sensu Gorczyca (2000)
status

 

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 View TABLE 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).

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 View FIGURE 8 ; 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 View FIGURE 8 , 9a, g View FIGURE 9 ; 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 View FIGURE 9 ; 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 View FIGURE 9 ; 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 View FIGURE 8 , 9a, g View FIGURE 9 ), and the suture behind the mandibular plate can be faint ( Fig. 9b View FIGURE 9 ; Namyatova & Cassis 2016: 2 C) or strongly depressed ( Figs 9a, g View FIGURE 9 ). 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 View FIGURE 9 ; 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 View FIGURE 9 ; 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 View FIGURE 9 , 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 View FIGURE 8 ; 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 View FIGURE 9 ; Namyatova & Cassis 2021: figs 3A, 5A) or are strongly removed in the dorsal direction ( Fig. 9a,c View FIGURE 9 ; Wolski 2017: figs 9, 95). The base of clypeus is situated close to the ventral margin of the eyes ( Fig. 9h View FIGURE 9 , 8a, b, i View FIGURE 8 ; Yasunaga 2000: fig. 2; Namyatova & Cassis 2021: 3 C, 5C) or is strongly removed from the eyes’ ventral margin ( Figs 8c–h View FIGURE 8 , 9g View FIGURE 9 ). The vertex in Cylapini is either carinate posteriorly ( Carvalhoma , Corcovadocola , Cylapoides , Dariella , Labriella ) ( Figs 9h View FIGURE 9 , 11e View FIGURE 11 ; Namyatova & Cassis 2016, 2021: figs 3b, 5b) or devoid of carina ( Cylapinus , Cylapomorpha , Mangalcoris , Cylapus complex) ( Figs 11a, g View FIGURE 11 ; 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 View FIGURE 9 , 11a View FIGURE 11 ; Namyatova & Cassis 2016; Wolski 2017: 33) or strongly developed so the head is V-shaped in anterior view ( Cylapus complex) ( Figs 8c–h View FIGURE 8 , 9g View FIGURE 9 ). 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 View FIGURE 8 , 9h View FIGURE 9 ; Namyatova & Cassis 2016: fig. 2B, 2021: 3B, 5B) or the eyes are pedunculate ( Cylapomorpha , Cylapinus , Cylapus complex) ( Figs 8a, c–i View FIGURE 8 , 9g View FIGURE 9 ). 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 View FIGURE 3 ; 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 View FIGURE 4 ; 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 View FIGURE 9 ; 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 View FIGURE 9 ; 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 View FIGURE 9 , 11e View FIGURE 11 ) or punctate ( Cylapinus , Dariella , Cylapus complex) ( Figs 9a View FIGURE 9 , 10a View FIGURE 10 , 11a, g View FIGURE 11 ). The lateral margin is always ecarinate ( Figs 9a, b View FIGURE 9 , 10a View FIGURE 10 , 11b View FIGURE 11 ). The collar in Cylapini is always present, and in lateral view is situated anteriorly to the propleural suture ( Fig. 9a View FIGURE 9 ).

Thoracic pleura. The mesepimeral spiracle in Cylapini is slit-like ( Figs 10b–e View FIGURE 10 , 11c, f View FIGURE 11 ), in members of the Cylapus complex it is covered with evaporative bodies ( Figs 10b–e View FIGURE 10 ). In most Cylapini taxa except for the Cylapus complex, the metepisternum is narrow and rectangular ( Figs 11c, f View FIGURE 11 ; Namyatova & Cassis 2016: fig. 2K, 2021: figs 3F, 5H). In the Cylapus complex the metepisternum is broad and almost square ( Figs 10b–e View FIGURE 10 ). In most Cylapini genera the metepisternum is carinate posteriorly ( Figs 10b–e View FIGURE 10 , 11c, f, i View FIGURE 11 ). 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 View FIGURE 11 ; 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 View FIGURE 10 ). In many genera of the Cylapini the metepimeron is well exposed, not obscured by second abdominal segment ( Figs 10b–e View FIGURE 10 , 11c, f View FIGURE 11 ). The ostiolar peritreme is usually oval ( Figs 10d–e View FIGURE 10 , 11c, f View FIGURE 11 ), rarely the peritreme is narrow and ear-like ( Amapacylapus ) ( Fig. 10b View FIGURE 10 ) or strongly protruding, thin and arcuate and sharply pointed ( Cylapus ) ( Fig. 10c View FIGURE 10 ).

Hemelytron. The hemelytron in Cylapini is either impunctate ( Corcovadocola , Cylapoides , Labriella , Cylapomorpha ) ( Fig. 11e View FIGURE 11 ; 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 View FIGURE 11 ; 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 View FIGURE 4 , 6a View FIGURE 6 ; 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 View FIGURE 10 , 11d, j View FIGURE 11 ; 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 View FIGURE 10 ; 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 View FIGURE 12 ; 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 View FIGURE 12 , 13m View FIGURE 13 ; 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 View FIGURE 14 ; Wolski 2017: figs 12, 66; Wolski et al. 2020: fig. 48; Namyatova & Cassis 2021: fig. 6) or devoid of spiculi ( Figs 12j, p View FIGURE 12 , 13a, g, m View FIGURE 13 ; Wolski et al. 2020: fig. 40). Sometimes the endosoma is composed of strongly inflated sclerotized lobes ( Fig. 12g View FIGURE 12 ; 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 View FIGURE 21 ; 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 View FIGURE 21 , 22i, j View FIGURE 22 ; 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 View FIGURE 22 ; 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 View FIGURE 20 ), or it possesses a well-developed, elongated sclerite adjacent to the base of the first gonapophyses ( Figs 20a, d–h View FIGURE 20 ).

The bursa copulatrix in Cylapini is voluminous, laterally extending beyond first gonapophyses ( Figs 15a, b, f, h View FIGURE 15 ; 16a, d, g View FIGURE 16 , 17j View FIGURE 17 , 18b View FIGURE 18 , 19a View FIGURE 19 ; 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 View FIGURE 15 ; 16a, d, g View FIGURE 16 , 17j View FIGURE 17 , 18b View FIGURE 18 , 19a View FIGURE 19 ; Wolski et al. 2020: fig. 66). The posterior wall of the genital chamber is simple and membranous ( Figs 19f, g View FIGURE 19 ; 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 View FIGURE 15 , 16g View FIGURE 16 , 18c View FIGURE 18 , 19a, d View FIGURE 19 ) or unpaired ( Figs 15a, b View FIGURE 15 , 16a, d View FIGURE 16 , 17a, d, h, k View FIGURE 17 ). In some genera they can be minute, occupying a small portion of the dorsal wall (roof) of the genital chamber ( Cylapoides , Labriella ) ( Fig. 15g View FIGURE 15 ; 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 View FIGURE 16 , 17a, d, h, k View FIGURE 17 ; Wolski et al. 2020: Figs 57, 63, 65) or occupies most of it ( Amapacylapus , Peltidocylapus , some Valdasus ) ( Figs 15a, b View FIGURE 15 , 18c View FIGURE 18 , 19b, e View FIGURE 19 ). 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 View FIGURE 15 ). The lateral oviducts are either wide ( Cylapus complex) ( Figs 15a View FIGURE 15 , 16d, g View FIGURE 16 , 18a, b View FIGURE 18 , 19b, e View FIGURE 19 ) or thin ( Cylapinus minusculus , Cylapomorpha , Cylapoides unicolor ) ( Figs 15f, g, h View FIGURE 15 ; Yasunaga 2000: Fig. 7 View FIGURE 7 ). They can be short (as in Amapacylapus , Peltidocylapus , and some species of Valdasus ) ( Figs 15a, b View FIGURE 15 , 18c View FIGURE 18 , 19b, e View FIGURE 19 ) or long, reaching beyond lateral or posterior portion of the genital chamber ( Cylapus , some species of Valdasus , Cylapomorpha , Cylapinus , Cylapoides ) ( Figs 15f, g View FIGURE 15 , 16d, g View FIGURE 16 , 17d, k View FIGURE 17 ; Yasunaga 2000: fig. 7). The spermathecal gland opens centrally in the genital chamber, between the lateral oviducts ( Figs 15a, b, f, h View FIGURE 15 , 18a View FIGURE 18 , 19b, e View FIGURE 19 ; 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 View FIGURE 16 , 17d View FIGURE 17 ). 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 View FIGURE 15 ), 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 View FIGURE 16 , 17f, h, k, l View FIGURE 17 ; 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 View FIGURE 12 , 13a, m View FIGURE 13 , 14a, f, e, l View FIGURE 14 ) 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.

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hemiptera

Family

Miridae