taxonID	type	description	language	source
25053808FF80FF8CB918FBC75E30F97B.taxon	description	The monophyly of Bothriomirini is not questioned (see node 5), which confirms the results of the study by Namyatova et al. (2019). Most of the apomorphies listed by Namyatova et al. (2019) and the present work are contradicted. However, we discovered that the microsculpture under the eye is present in different genera of this tribe and might be one of the main diagnostic characters for this group (Namyatova et al. 2019: fig. 5 F). In the analysis by Wolski (2021), Bothriomirini are a sister group to the clade comprising Cylapini and Vanniini. In the paper by Oh et al. (2023), Bothriomirini form a clade with Eccritotarsini, although with low support. Our analyses did not confirm those relationships, because Bothriomirini form sister-group relationships with some representatives of Cylapini (node 8; see discussion of Cylapini). In the future, a comprehensive phylogenetic study of the tribe using molecular data is needed to test the position of the Bothriomirini genera relative to each other. It is important to include Afrobothriomiris Gorczyca, 2000, which is the only African representative of this tribe (Gorczyca 2000), in the subsequent analyses.	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
25053808FF80FF8CBAEEFDBB598CFB89.taxon	description	In the previous studies, the monophyly of Cylapinae was considered to be doubtful, because there were no apomorphies for this group (Namyatova and Cassis 2019 a, b). Many representatives of this subfamily share a similar set of characters in the tarsus and pretarsus, i. e. slender tarsi, setiform and asymmetric parempodia, and three rows of tiles on the unguitractor, with those in the middle row acute. However, this set of characters is not unique for Cylapinae, because Psallopinae and Isometopinae have similar structures (e. g. Schuh and Schwartz 1984, Namyatova and Cassis, 2016, 2019 b, 2022, Namyatova et al. 2016). There was an argument that those subfamilies are related (e. g. Schuh and Schwartz 1984, Wolski and Henry 2015), and Wolski and Henry (2015) synonymized Psallopinae with Cylapinae, which was not supported by Namyatova and Cassis (2019 b). In the previous molecular-based analyses, representatives of the different Cylapinae groups did not form a monophyletic clade (Schuh et al. 2009, Namyatova and Cassis 2019 a, Oh et al. 2023). In the study by Schuh et al. (2009), only three species from three tribes were sampled (Cylapini, Fulviini, and Vanniini). In the study by Namyatova and Cassis (2019), all tribes were included in the phylogeny; however, in that work only representatives of Psallop s (Psallopinae) and Myiomma (Isopmetopinae) were added as outgtoup taxa, and this was not sufficient to test the monophyly of Cylapinae. The phylogenetic study of Oh et al. (2023) provided the most comprehensive analysis of the relationships within Miridae based on the molecular data to date, and they included 11 genera from all Cylapinae tribes, including nominotypical genera. However, Psallops was not included in this analysis. The tribes of Cylapinae in the study by Oh et al. (2023) did not form sister-group relationships with each other. Therefore, in our analysis we expected that some of the taxa placed into Cylapinae might form clades with other mirids. The results of the present analysis agree with the previous studies and suggest non-monophyly of Cylapinae. In the total-evidence phylogeny, Bothriomirini, Cylapini, and Vanniini form a clade with Bryocorinae, Isometopinae, Mirinae, Orthotylinae, Phylinae, and Deraeocoris ater (Deraeocorinae) (PPTE = 98; see node 14). This clade does not appear in other phylogenies. In the morphology-based phylogeny a similar clade is present, but Deraeocoris ater is not included there. In the tree resulting from the Bayesian analysis with 65 taxa, Bothriomirini, Cylapini, and Vanniini form a clade with Mirinae, Orthotylinae, Phylinae, and Bryocorinae without Nesidiocoris sp. However, Psallops, Rhinomiris, Phyllofulvius, Psallofulvius, and the undescribed Fulviini genus from Australia are also included in this clade (PP 65 = 90). Those clades do not correspond to the results of Oh et al. (2023). The position of Psallops remains uncertain. Gorczyca (2000) considered Cylapinae, Isometopinae, and Psallopinae as closely related. Wolski and Henry (2015) reported that the last-instar nymphs of Psallops are similar to those of Fulvius. Additionally, Psallofulvius and Phyllofulvius have the labial segment IV subdivided, similar to Psallops (Namyatova 2022, Namyatova and Cassis 2022). In our phylogeny, Psallops never forms well-supported sister-group relationships with any representatives of Cylapinae. The position of Isometopinae relative to cylapines is also uncertain. In the morphology-based phylogeny, this subfamily forms sister-group relationships with the clade comprising Bothriomirini, Carvalhoma parvum, Dariella rubrocuneata, Cylapinus minusculus, and Schizopteromiris lordhowensis, although with low support. In the total-evidence phylogeny, Isometopinae form a well-supported clade with Bothriomini, Cylapini, Vanniini, and Eccritotarsus cf. nigrocruciatus (PPTE = 98; see node 11). A similar clade does not appear in the phylogenies based only on the molecular data; however, in most of those analyses Isometopinae form sister-group relationships with Ecrritotarsus cf. nigrocruciatus (see node 10). A close phylogenetic position of Isometopinae and Eccritotarsini has never been proposed before. It could also be a result of insufficient sampling of eccritotarsines, which is a highly diverse group (Konstantinov et al. 2018).	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
25053808FF80FF8DB930F97C5B08FB2F.taxon	description	Gorczyca placed the Rhinomiris complex, the Rhinocylapus complex, and Rhinomiriella Gorczyca, 2001 (junior synonym of Ceratofulvius Reuter, 1902) in Rhinomirini (Gorczyca 2000, 2001). Gorczyca (2000) suggested that this is a sister group to Cylapini. Superficially, Rhinomirini are more similar to Fulviini, because their representatives have a more or less prognathous head and long labium, and this tribe forms a clade with Fulviini in the study by Wolski (2021). Namyatova and Cassis (2019 a) showed that Rhinomirini sensu Gorczyca (2000) are non-monophyletic based on molecular and morphological data. In that work, the Rhinomiris complex did not form well-supported sister-group relationships with any other group. The Rhinocylapus complex and Ceratofulvius formed well-supported clades with representatives of Fulviini. Therefore, Rhinomirini were restricted to the Rhinomiris complex, comprising only four genera: Rhinomiris, Rhinomiridius Poppius, 1909, Lundbladiolla Carvalho, 1955, and Pararhinomiris Gorczyca, 2003 (Namyatova and Cassis 2019 a, Tyts et al. 2022). Those results were confirmed in the present study. In the paper by Oh et al. (2023), only Rhinomiris camelus Poppius, 1909 was included in the analysis, and it did not form well-supported sister-group relationships with other taxa. In this analysis, we included the molecular data for Rhinomiris only, and we also included morphological data for Rhinomiridius dentatus. These two genera form a well-supported clade, and this confirms the previous results (Namyatova and Cassis 2019 a, Tyts et al. 2022). In most analyses, Rhinomirini form poorly supported clades with other groups. The exception is the Bayesian analysis with 65 taxa, where this tribe is in the clade with Bothriomirini, Cylapini, Vanniini, Mirinae, Orthotylinae, Phylinae, and Bryocorinae without Nesidiocoris tenuis (PPTE = 90; see discussion of Cylapinae). We did not have specimens of Lundbladiolla and Pararhinomiris for examination. The next step in studying Rhinomirini sensu Namyatova and Cassis (2019 a) should involve testing their monophyly with morphological and molecular data. The position of Rhinomirini within Miridae remains uncertain, and most probably, genomic data are needed to clarify it.	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
25053808FF81FF8DBA9AFB215F5EFBE8.taxon	description	In the previous work on the morphology-based phylogeny of Cylapini, it was shown that this tribe is paraphyletic and includes Vannini (Wolski 2021). Our results show that Cylapini are polyphyletic. The genera Amapacylapus Carvalho & Fontes, 1968, Cylapinus Carvalho, 1986, Cylapus, Peltidocylapus, and Valdasus form the first clade (node 1), which either includes Vanniini or forms sister-group relationships with them (node 3). In the analysis by Oh et al. (2023), this clade was not recovered; however, only a single representative of Vanniini, Vanniopsis lordhowensis, was included in that study. Carvalhoma Slater & Gross, 1977, Cylapoides Carvalho, 1952, Dariella Namyatova & Cassis, 2021, Labriella Namyatova & Cassis, 2021, and Schizopteromiris form a clade with Bothriomirini (node 8). The position of Cylapomorpha spp. is uncertain. In most analyses, it does not form sister-group relationships with any taxa. However, in the Bayesian analysis with 78 taxa, it is a sister group to a clade that includes Phyllofulvius spp., Psallofulvius spp., and the undescribed genus from Australia (PP 78 = 99). In the total-evidence phylogeny, it forms a well-supported clade with Cylapus complex and Vanniini (node 4). Wolski (2021) provided a diagnosis for the Cylapus complex, in which he included four genera: Amapacylapus, Cylapus, Peltidocylapus, and Valdasus. In the work of Wolski (2021), Cylapinus was not included in this complex; however, it possesses many salient features of this group, i. e. it has a punctate body (Wolski 2021: fig. 11 A), its vertex has a distinctly depressed midline (Wolski 2021: fig. 8 A), and the antennal fossa is situated well above the suture between mandibular and maxillary plates (Wolski 2021: figs 8 A, 11 A). According to Wolski (2021), the eyes are strongly pedunculated in the Cylapus complex, whereas in Cylapinus they are slightly pedunculated. However, the differences between those two states are unclear; for example, at least in Peltidocylapus, the eyes are slightly raised above the vertex (Wolski 2021: fig. 8 F, H). In our analyses, Cylapinus minusculus forms a clade with representatives of the Cylapus complex, and we include this species in this group. The close relationships of Schizopteromiris lordhowensis and Labriella fusca with Bothrimirini are well supported in all analyses. We did not have molecular data for Dariella rubrocuneata, but it is likely to be close to Schizopteromiris (see Namyatova and Cassis 2021, 2022). The positions of Carvalhoma parvum and Cylapoides unicolor need further testing with molecular data, because they are not very similar morphologically to any other genera. Although Cylapoides unicolor and Labriella fusca form a clade in the present analysis and the study by Wolski (2021), they live on different continents, and the convergence of some characters is possible. Although we included both types of data for Cylapomorpha, its position remains uncertain, and it is likely that only genomic data will help us to understand the position of this genus within Miridae. In the future, the position of Cylapini genera not included in this analysis, i. e. Corcovadocola Carvalho, 1948, Mangalcoris Murphy & Polhemus, 2012, and Phyllocylapus Poppius, 1913, should be tested with both types of data.	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
25053808FF81FF8AB9D5FBE75AF8FB4D.taxon	description	Vanniini are rendered as paraphyletic in the analyses based only on the molecular data, and the species from this tribe form a monophyletic group in the total-evidence analysis. The results of the molecular-only analyses are unexpected, because the monophyly of Vanniini has never been questioned in morphological assessments (Gorczyca 1997, Cassis et al. 2003). The genera of this tribe did not form a clade in the study byNamyatova and Cassis (2019 a); however, only two genera of Vanniini and a single genus of Cylapini were included there. This group is absent in the morphology-based analysis. We had only morphological data for Palaucoris, and it forms sister-group relationships with the rest of Vanniini in the total-evidence analysis (node 2), which also confirms the results provided by Wolski (2021). This clade is supported by a single contradicted apomorphy, which is the pretarsus with flattened parempodia, as opposed to all other Cylapinae tribes, which have setiform parempodia (e. g. Cassis et al. 2003). The Vanniini species with the pretarsus available for examination also have a rounded middle row of tiles on the unguitractor, whereas in other Cylapinae it is acute. However, the middle row of tiles is absent in the examined Palaucoris species. Further studies should be performed to test whether the structure of the unguitractor is uniform within Vanniini. Judging from previous studies, male genitalia are very different in Vanniini, e. g. the vesica is strap-like in Vannius (Gorczyca 1997), and it is more or less voluminous in other genera (Gorczyca and Konstantinov 2001, Cassis et al. 2003). However, Palaucoris does not have vesica. This genus has other significant morphological differences, e. g. its scent gland metathoracic evaporative area is wide, the body is punctate, the suture between the meso- and metapleura does not extend behind metathoracic spiracle, and the tarsi are moderately widened (Namyatova et al. 2016). To answer the question on the monophyly of Vanniini, both molecular and morphological data should be studied for all the genera of this tribe. In the paper by Oh et al. (2023), the Cylapus complex does not form well-supported sister-group relationships with other taxa, but Wolski (2021) suggested that there is a strong argument for synonymizing Vanniini with Cylapini. The results of our analyses support those of Wolski (2021); however, the Сylapini genera outside of the Cylapus complex are not close to this group. Before suggesting any taxonomic changes regarding Vanniini, we need to test the monophyly of this tribe, in addition to the position of the Cylapini genera outside of the Cylapus complex, using more molecular data. Moreover, we did not recover any apomorphies for the group comprising the Cylapus complex and Vanniini. In the study by Woski (2021), there are four strict apomorphies for this clade: their head in the anterior view is about twice higher than wide, the base of clypeus is distinctly below the ventral margin of eyes, the antennal fossa is strongly removed from the suture between the mandibular and maxillary plate in the dorsal direction, and the ventral margin of the eye is strongly removed from the ventral margin of the head. The shape of the head and the position of the clypeus base, antennal fossa, and eye vary strongly within this clade and other mirids. We did not include the characters on the head height-to-width ratio and the position of the ventral margin of the eye in relationship to the ventral margin of the head. Two other characters were coded differently, and their states do not appear as apomorphies for this group. However, we agree that the combination of characters in the head can be diagnostic for the Vanniini + Cylapus complex group.	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
25053808FF86FF8BBA89FA835A81FD47.taxon	description	In the previous works, Fulviini were not monophyletic (Namyatova and Cassis 2019 a, Wolski 2021). In the total-evidence analysis by Namyatova and Cassis (2019 a), Fulviini were polyphyletic and Phyllofulvius was not related to other members of this tribe. In the morphology-based phylogenies (Namyatova and Cassis 2019 a, Wolski 2021), Fulviini are paraphyletic and include Rhinomirini sensu Namyatova and Cassis (2019 a). Our analyses with molecular data never place Rhinomirini within or close to any genera of Fulviini. Fulviini are the largest tribe within Cylapinae, comprising 66 recent genera, and only 30 are included in this study. The results show that there are at least two clades in Fulviini, which might not be closely related (see discussion for Cylapinae). In the phylogenies based only on the molecular data, most Fulviini genera form a clade with medium to high support (node 21), and a similar clade was found by Oh et al. (2023). The results of the total-evidence and morphology-based analyses neither contradict nor confirm this topology. This clade is likely to exist, and the morphology could not support it owing to the numerous cases of convergences and missing data, because many species were coded from a single specimen. In some cases, it is possible to assign a genus to this clade based solely on the morphological descriptions, even if molecular data are unknown for the taxon. For example, among the genera in our study, Bironiella Poppius, 1909, Callitropisca, Cylapofulvius Poppius, 1909, Fulvidius Poppius, 1909, Schmitzofulvius Gorczyca, 1998, and Sulawesifulvius Gorczyca et al., 2004 are most probably in this clade, because they form well-supported relationships with the genera from this group. Among the genera not included in our analysis, Cassisotropis, Carvalhofulvius Stonedahl & Kovac, 1995, Comefulvius, Euchilofulviella Gorczyca, 1999, Infernotropis Taszakowski, Masłowski, Wolski & Gorczyca, 2022, Henryfulvius Wolski, 2014, Rewafulvius Carvalho, 1972, and Xenocylapoides Carpintero & Chérot, 2014 most probably also belong to clade 21. The second Fulviini clade includes Phyllofulvius, Psallofulvius, and the undescribed Fulviini genus from Australia, which is recovered in all analyses with molecular data (node 26). Namyatova (2022) suggested that Laetifulvius morganensis is also related to Phyllofulvius and Psallofulvius, and this is confirmed by the total-evidence and morphological phylogenies. Fulviella Carvalho, 1991, which is not included in this analysis, most probably belongs to this group too, because it shows affinities to Phyllofulvius (Namyatova and Cassis 2022). Namyatova (2022) also hypothesized that Psallops might be close to this group, and this is recovered in the morphology-based analysis, but the analyses with molecular data do not confirm this. The monophyly of Rhinocylapus complex (node 22) is confirmed again (Namyatova and Cassis 2019 a, Tyts et al. 2022); however, the morphology-based and total-evidence phylogenies show that it might include more genera, i. e. Bironiella, Cylapofulvius, and Fulvidius. Additionally, it was hypothesized that Teratofulvioides Carvalho & Lorenzato, 1978 might be close to or even synonymous with Punctifulvius (Namyatova and Cassis 2022). The sister-group relationships of Rhinocylapus complex with Fulvius are confirmed in the Bayesian inference analyses, and the RAXML tree with 78 taxa does not contradict those relationships. Fulvius sp. from Australia forms a clade with other Fulvius species only in the Bayesian inference analysis with 78 taxa and in the total-evidence phylogeny. Australian species might represent another genus; however, an analysis with more specimens from Australia should be performed to confirm this idea. Another well-supported clade includes Australian genera Callitropisca, Ceratofulvius, Lygaeoscytus Reuter, 1893, Micanitropis, and Xenocylapidius, in addition to African Schmitzofulvius bigibber and South American Xenocylapus tenuis (node 20). Only morphological data were included for Callitropisca florentine; however, this taxon is very similar to Micanitropis and Xenocylapidius morphologically (Namyatova and Cassis 2021), and their close relationships are very possible. We did not have molecular data for S chmitzofulvius bigibber and Xenocylapus tenuis, and their position within this clade should be tested in the future. Euchilofulvius antennatus and Peritropisca bituberculata form a clade together (node 15), and the similarity of those two genera has been mentioned before (Wolski and Gorczyca 2014 b, Namyatova and Cassis 2022; Masłowski et al. 2023). These taxa might form a clade with Peritropis and the genus near Peritropella (node 17). Morphological data also suggest that Peritropis forms a clade with Sulawesifulvius cf. thailandicus. Gorczyca (1998, 1999), and Maslowski et al. (2023) hypothesized that Schmitzofuvius might be close to Euchilofulvius and Peritropisca; however, this is not supported by our results. Future studies of Fulviini should involve more genera included in this group. It is important to test using molecular data the position of the genera that cannot be assigned to any of the clades based on morphology. In our analysis, such genera are Cylapocoris, Howefulvius Schmitz & Štys, 1973, and Trynocoris Herring, 1976. There are many Fulviini genera not included in this analysis, and their relationships with other taxa are unclear, e. g. Hemiophthalmocoris Poppius, 1912, Gulacylapus Carvalho, 1986, Rhinophrus Hsiao, 1944, Rhyparochromomiris Henry & Paula, 2004, and Tucuruisca Carvalho, 1986. Adding such genera into the phylogeny will help us to understand the borders of Fulviini and the relationships of their subgroupings with other mirids.	en	Namyatova, Anna A., Tyts, Veronica D. (2025): Total-evidence phylogeny of the subfamily Cylapinae and the divergence dates for its subgroupings (Insecta: Heteroptera: Miridae). Zoological Journal of the Linnean Society 203 (1): 1-34, DOI: 10.1093/zoolinnean/zlae008, URL: https://doi.org/10.1093/zoolinnean/zlae008
