Fulviini

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), pp. 1-34 : 29-30

Monophyly of Fulviini and relationships of their subgroupings

In the previous works, Fulviini were not monophyletic ( Namyatova and Cassis 2019 a, Wolski 2021). In the total-evidence analysis by Namyatova and Cassis (2019a), 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 (2019a) . 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 2014b, 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.