HELEOCORINI, Sites, 2022

Sites, Robert W., 2022, Phylogeny and revised classification of the saucer bugs (Hemiptera: Nepomorpha: Naucoridae), Zoological Journal of the Linnean Society 195, pp. 1245-1286 : 1265-1266

TRIBE HELEOCORINI TRIB. NOV.

FIGS 4B , 5A, B , 12 , 14A, C , 15B , 21

Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 8CBB9ABB-857E-4540-8213-1B1648FDF868

Diagnosis: This group includes some taxa with extreme habitat specializations, thus a simple characterization of the tribe is not possible. A feature characteristic for all species except Pogonocaudina indica Sites & Zettel, 2011 and Decarloa darlingtoni La Rivers, 1969 is that the spinose setae of the posteroventral row on the mesofemur are tightly spaced such that adjacent setae or their sockets are in contact or nearly so ( Fig. 5a ) and number approximately 60–90. In addition, in all except the clade containing Decarloa , Diaphorocoris and Interocoris , the anteroventral seta row on the metafemur is noticeably arcuate or with a posteromesal direction change in the basal third as it extends basally ( Fig. 5b ).

Comments: Because the clades have a high degree of fidelity with respect to geography and the type species is from Mumbai, the Heleocoris clade from India remains unchanged as the genus Heleocoris (see Comments section above) and the containing group is the new tribe Heleocorini . Representing this clade in the analyses are Heleocoris elongatus Montandon, 1897b , Heleocoris rotundatus Montandon, 1908 and Heleocoris vicinus Montandon, 1910b . Species in the clades containing Heleocoris from Madagascar and Southeast Asia are here transferred to the newly erected genera Tsingala gen. nov. and Heleolaccocoris gen. nov., respectively (described below). Tsingala was recovered with its sister genus Temnocoris , both endemic to Madagascar (red-coloured clades in Fig. 13 ), in the tribe Laccocorini . Species of Laccocoris from Borneo, Sumatra, the Philippines and Laccocoris dissidens Montandon, 1910 , described from Burma, are transferred to Heleolaccocoris along with the Southeast Asian species of Heleocoris (blue-coloured clades in Fig. 13 ).

Among the Indian waterfall-inhabiting species, Pogonocaudina indica was recovered as a derived member of a clade of Diaphorocoris . Although it exhibits morphological features supporting its distinction as a genus-level taxon, including singlesegmented fore tarsi in both sexes, its evolutionary relationship with species of Diaphorocoris require that the genus be synonymized. As such, the monotypic Pogonocaudina is a junior synonym of Diaphorocoris with the species becoming Diaphorocoris indicus ( Sites & Zettel, 2011) comb. nov. Nested in this group of Indian taxa are Interocoris mexicanus ( Usinger, 1935) and Decarloa darlingtoni , both from northern tropical America (Mesoamerica and Hispaniola, respectively). These two monotypic genera have long been enigmatic because there is little representation of this subfamily in the New World. That they are most closely related to taxa from India raises new questions about their origin.

Included taxa: Heleocoris and Diaphorocoris occur in India; Heleolaccocoris occurs from the easternmost Indian states and Indochina through the Sunda Islands to the Philippines. Decarloa and Interocoris are American monotypic genera.

Figures

Figure 4. Foretibiae and foretarsi of Laccocorinae: Ctenipocoris asiaticus (a), Heleolaccocoris ovatus (b), Namtokocoris dalanta (c).

Figure 5. Heleocoris elongatus mesofemur (a), metafemur (b), Tsingala gen. nov. sp. C mesofemur (c), metafemur (d). White arrows indicate posteroventral spine rows, black arrows indicate anteroventral spine rows.

Figure 12. Mesotibia and mesotarsus of male and female Heleolaccocoris ovatus showing a sexually dimorphic ventral tomentose pad of hairs.

Figure 13. Detail of Most Probable Phylogeny dendogram of Laccocorinae.The red-coloured clade includes the Madagascar genera Temnocoris and Tsingala gen. nov.(formerly Heleocoris). The blue-coloured clade represents Heleolaccocoris gen.nov., which comprises species of the former genera Heleocoris and Laccocoris. All species of Tsingala gen. nov. and Heleolaccocoris gen. nov. are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure.

Figure 14. Revised generic representation of the former genus Heleocoris. Dorsal habitus of Heleocoris vicinus (a), Laccocoris minisculus (b), Heleolaccocoris ovatus (c), Tsingala humeralis (d).

Figure 15. Revised generic representation of the former genus Laccocoris. Dorsal habitus of Laccocoris spurcus (a), Heleolaccocoris horvathi (b).

Figure 21. Heleolaccocoris horvathi male: dorsal habitus (a), genital capsule with terga IX and X removed (b).