Camarotoscena Haupt, 1935

Aléné Geonho Cho, Daniel Burckhardt Liliya Š. Serbina Igor Malenovský Dalva L. Queiroz Désirée C. & Percy, Diana M., 2024, Phylogeny and classification of jumping plant lice of the subfamily Liviinae (Hemiptera: Psylloidea: Liviidae) based on molecular and morphological data, Zoological Journal of the Linnean Society 201 (2), pp. 387-421 : 406

Camarotoscena Haupt, 1935

Camarotoscena Haupt 1935: 228 .

Type species: Rhinocola speciosa Flor, 1861 , by original designation.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1C ); in dorsal view, about as wide as thorax, moderately transverse. Vertex rhomboidal; covered in areolate-rugose microsculpture; passing smoothly into genae anteriorly; coronal suture fully developed; genae weakly produced ventrally but not enlarged into processes; frons elongate, parallel-sided to narrowly triangular ( Fig. 3E ); median ocellus not visible in perpendicular view to vertex; subgenae not differentiated into separate sclerites ( Fig. 3E ); compound eyes, in dorsal view, hemispherical, adpressed to head. Clypeus pear-shaped, moderate-sized, rounded ventrally, hardly visible in lateral view as it is hidden by genae. Antenna slightly longer than head width; flagellum with simple setae; segment 3 longest, as long as segments 4–5 together; segments 4, 6, 8, and 9 bearing each a subapical rhinarium lacking marginal spines. Thorax moderately slender; dorsal outline, in lateral view, weakly curved. Pronotum, in dorsal view, weakly curved posteriad laterally; propleurites subrectangular, divided by perpendicular suture into subequal epimeron and episternum. Metapostnotum with blunt tubercle. Mesosternum ( Fig. 5B ) narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave; pleurosternal suture hardly visible; basisternum triangular; katepisternum small antero-laterally, not bent dorsad laterally; angle of arms of precoxale obtuse. Pro- and mesotibiae cylindrical. Metacoxa with blunt, moderately long, horn-shaped meracanthus. Metafemur with the three ventral sense organs in the middle; apex with few stout long setae ( Fig. 6O ). Metatibia longer than metafemur, distinctly widened apically; bearing 9–12 slightly irregularly spaced apical sclerotized spurs and two to four peg-like setae adjacent to inner spurs ( Fig. 7C ). Both metatarsal segments relatively short, subequal in length. Forewing slightly subtrapezoidal; costal and anal margins subparallel or widening towards apex, 2.2–2.3 times as long as wide, membranous; vein C + Sc weakly, evenly convex, slender, distinctly delimited from cell; costal break developed, close to apex of vein R 1; pterostigma wide, entirely membranous; nodal line developed; veins R and M + Cu subequal; vein Rs relatively straight; vein M about as long as M 1 + 2; vein Cu 1a almost straight or weakly curved towards anal margin; veins M 1 + 2 and M 3 + 4 oblique to wing margin apically; anal break adjacent to apex of vein Cu 1b; surface spinules fine or coarse, spaced or dense, present in all cells. Hindwing slightly shorter than forewing; with one or two costal setae proximal to costal break and two clearly separated groups of two to four proximal and two to four distal setae, distal to costal break; vein R + M + Cu indistinctly trifurcating, base of vein Cu indistinct. Abdominal base with a sclerotized area on either side covered in spines. Aedeagus with simple proximal portion bearing many weak folds subapically; apex of distal portion not differentiated from stem. Female subgenital plate lacking apical process.

Last instar immature. Antenna seven segments; lacking sectasetae or lanceolate setae on antennal flagellum. Mid- and hindlegs without massive peg-like setae. Dorsal body surface lacking minute clavate setae. Precaudal abdominal tergites bearing densely spaced simple setae and sectasetae. Anus in ventral position; with additional pore fields developed.

Comments: Monophyletic with strong molecular support, morphologically supported by two homoplastic characters that we consider non-unique synapomorphies ( Figs 10 , 11 ; Supporting Information, File S4). The circumscription of the genus by Burckhardt and Mifsud (2003) is supported here. Included available species, distribution, and host plants are summarized in Table 3 and Supporting Information, File S3.

Figures

Figure 1. Habitus images illustrating generic diversity within Liviinae: A, Anomoterga africana (Loginova); B, Aphorma lichenoides (Puton); C, Camarotoscena speciosa (Flor); D, Livia junci (Schrank); E, Syntomoza magna (Kuwayama); F, Diclidophlebia eastopi Vondráček; G, Haplaphalara dahli (Rübsaamen); H, Klyveria setinervis (Burckhardt); I, Liella cf. insolita (Mifsud and Burckhardt); J, Melanastera smithi (Burckhardt et al.); K, Paurocephala chonchaiensis Boselli; L, Woldaia nebulosa Brown and Hodkinson. Scales 1.0 mm.

Figure 3. Head of adult Liviidae: A, Diclidophlebia xuaniMessi; B, Syntomoza magna (Kuwayama); C, Anomoterga tahuata Klyver; D, Aphorma lichenoides (Puton); E, Camarotoscena speciosa (Flor); F, Livia junci (Schrank); G, Klyveria setinervis (Burckhardt); H, Paurocephala dayak Mifsud and Burckhardt.A, B, dorsal view; C–H, ventral view. Scales A, B = 0.1 mm; C–H = 0.1 mm.

Figure 5. Mesosternum of adult Liviinae: A, Anomoterga scolopiae (Yang); B, Camarotoscena speciosa (Flor); C, Klyveria setinervis (Burckhardt); D,Livia junci (Schrank); E, Paurocephala robusta Mifsud and Burckhardt; F, Syntomoza magna (Kuwayama).Abbreviations:bas, basisternum; kat, katepisternum; pcx, precoxale; pss, pleurosternal suture; scs, sternocostal suture. Scales 0.1 mm.

Figure 6. Hindlegs of adult Liviidae: A, Anomoterga africana (Loginova); B, Syntomoza magna (Kuwayama); C, Liella cf. insolita (Mifsud and Burckhardt); D, N, Paurocephala bifasciata Kuwayama; E, Psyllopsis fraxinicola (Foerster); F, Diclidophlebia oceanica (Crawford); G, D. excetrodendri (Li and Yang); H, D.xuani (Messi); I, Haplaphalara dahli (Rübsaamen); J, H. irvingiae (Burckhardt et al.); K, Klyveria setinervis (Burckhardt); L, Liella lanceomedia (Brown and Hodkinson); M, Melanastera lucens (Burckhardt et al.); O, Camarotoscena speciosa (Flor). A–D, metacoxa; E–N, metafemur, base left; O, apex of metafemur. Scales A–D = 0.2 mm; E–N = 0.1 mm; O = 0.05 mm.

Figure 7. Metatibial apex of adult Liviidae:A, Strophingia cinereae Hodkinson; B, Aphorma lichenoides (Puton); C, Camarotoscena speciosa (Flor); D, Livia junci (Schrank); E, Diclidophlebia oceanica (Crawford); F, D.xuani (Messi); G, Haplaphalara dahli (Rübsaamen); H, Liella lanceomedia (Brown and Hodkinson); I, Klyveria setinervis (Burckhardt); J, Melanastera lucens (Burckhardt et al.); K, M. maculipennis (Brown and Hodkinson); L, Paurocephala sauteri Enderlein.Abbreviations:a, anterior view, l, lateral view, m, medial, p, posterior view. Scales 0.05 mm.

Figure 10. Maximum likelihood (ML-mix) tree resulting from a mixture (heterotachous) model (–m GTR+H4 option in IQ TREE) based on the molecular data of the Liviidae and showing the three subfamilies Euphyllurinae, Neophyllurinae, and Liviinae, and two tribes Liviini and Paurocephalini. The numbers below the branches represent the bootstrap values of ≥50. Nodes with boostrap values of <50 are shown in interrupted lines.Light blue and yellow bars indicate the non-monophyletic genera, Diclidophlebia and Haplaphalara.

Figure 11. Phylogeny of the Liviidae showing strict consensus tree of the most parsimonious trees based on morphological data (character matrix in Supporting Information, File S2) and obtained with ‘Traditional search’ in TNT using equal weights.‘Unambiguous changes only’ in WINCLADA was used to optimize the character states on the cladogram. Black circles represent synapomorphies, light circles homoplasies; numbers above circles refer to characters numbers, number below to the character states (Table 2). Host plants of Liviini: Malpigiales, Poa, Poales, Ran, Ranunculales; of Paurocephalini:representation of members of Malvales. Distribution: Afrotropics (Afr), Australasia (Aus), Indomalaya (Ind), Nearctic (Nea), Neotropics (Neo), Oceania (Oce), Palaearctic (Pal).