Anomoterga Klyver, 1932

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 : 395-397

Anomoterga Klyver, 1932 , stat. rev.

Anomoterga Klyver 1932: 93 .

Type species: Anomoterga tahuata Klyver, 1932 , by original designation and monotypy.

= Homalocephala Yang and Li 1986: 54 .

Type species: Homalocephala homali Yang and Li, 1986 , by original designation and monotypy. Syn. nov.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1A ); in dorsal view about as wide as thorax, moderately transverse. Vertex rhomboidal to almost subrectangular; covered in areolate-rugose or imbricate microsculpture, sometimes much reduced on disc; passing smoothly into genae anteriorly; coronal suture fully developed; genae weakly produced ventrally but not enlarged into processes; frons small trapezoidal; median ocellus clearly visible in perpendicular view to vertex; subgenae not differentiated into separate sclerites ( Fig. 3C ); compound eyes, in dorsal view, hemispherical, adpressed to head. Clypeus pear-shaped, moderate-sized, flattened 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 or longer than segments 4–6 together; segments 4, 6, 8, and 9 bearing each a subapical rhinarium lacking marginal spines. Thorax moderately slender; dorsal outline, in lateral view, strongly curved ( Fig. 1A ). Pronotum, in dorsal view, weakly curved posteriad laterally; propleurites subrectangular, divided by perpendicular suture into subequal epimeron and episternum. Metapostnotum medially flattened, or with blunt tubercle or longitudinal ridge ( Fig. 4A ). Mesosternum narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave; pleurosternal suture hardly visible; basisternum indistinct; katepisternum small antero-laterally, not bent dorsad laterally; angle between arms of precoxale obtuse ( Fig. 5A ). Pro- and mesotibiae cylindrical. Metacoxa with blunt or subacute horn-shaped meracanthus ( Fig. 6A ). Metafemur with the three ventral sense organs in the middle; apex with few stout long setae. Metatibia longer than metafemur, slightly widened apically; bearing 9–11 slightly irregularly spaced apical sclerotized spurs and two to five peg-like setae adjacent to inner spurs. Both metatarsal segments relatively short, subequal in length. Forewing oblong-oval or subtrapezoidal; costal and anal margins subparallel or widening towards apex, 2.0–2.4 times as long as wide, membranous or subcoriaceous; vein C + Sc weakly, evenly or irregularly convex, slender, distinctly delimited from cell; costal break developed, sometimes indistinct, close to apex of vein R 1; pterostigma wide, entirely membranous; nodal line developed; veins R and M + Cu subequal; vein Rs almost straight or curved towards costal margin; vein M much longer than its branches; vein Cu 1a almost straight or curved towards anal margin; veins M 1 + 2 and M 3 + 4 perpendicular 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 to three costal setae proximal to costal break and two clearly separated groups distal to costal break, of three to five proximal and three to seven distal setae; vein R + M + Cu bifurcating into R and M + Cu. Abdominal base with a weakly 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 five to seven segments; lacking sectasetae or lanceolate setae on antennal flagellum. Mid- and hindlegs with massive peg-like setae. Dorsal body surface lacking minute clavate setae. Precaudal abdominal tergites lacking densely spaced simple setae or sectasetae. Anus in ventral position; sometimes with additional pore fields developed.

Comments: Supported by one synapomorphy in the morphological analysis ( Fig. 11 ) but paraphyletic in the combined analyses, and paraphyletic (ML-mix) or polyphyletic (ML-part, BI) in the molecular analyses ( Fig. 10 ; Supporting Information, File S4). Burckhardt and Mifsud (2003) synonymized Anomoterga and Homalocephala with Syntomoza but did not include Syntomoza magna , the type species of Syntomoza , in their analysis. Morphology suggests a close relationship of the type species of Anomoterga and Homalocephala with two species originally described in Camarotoscena and transferred by Burckhardt and Mifsud (2003) to Syntomoza . The four species together, however, are only distantly related to, and not congeneric with, S. magna . Similar results are obtained by the molecular and combined analyses, though the type species of Homalocephala was not included. We conclude that Anomoterga and Syntomoza are distinct genera, the latter being monotypic, and Homalocephala is a synonym of Anomoterga . Included available species, distribution, and host plants are summarized in Table 3 and Supporting Information, File S3. The following new or revived combinations are here proposed: Anomoterga africana ( Loginova 1975: 55) , comb. nov. (from Camarotoscena ; Syntomoza, Burckhardt and Mifsud 2003: 16 ); A. homali ( Yang and Li 1986: 54) , comb. nov. ( Homalocephala ; Syntomoza, Burckhardt and Mifsud 2003 16); A. hsenpinensis Fang and Yang 1986: 137 , comb. rev. ( Syntomoza, Burckhardt and Mifsud 2003: 16 ); A. scolopiae ( Yang 1984: 23) , comb. nov. ( Syntomoza ); A. tahuata Klyver 1932: 94 , comb. rev. (from Syntomoza, Burckhardt and Mifsud 2003: 16 ); A. unicolor (Loginova in Loginova and Parfentiev 1958: 99), comb. nov. ( Camarotoscena ; Syntomoza, Burckhardt and Mifsud 2003: 17 ).

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 4. Metapostnotum of adult Liviinae (arrow pointing to tubercle, tooth, ridge or horn): A, Anomoterga hsenpinensis Fang and Yang; B, Livia vernalis Fitch; C, Melanastera smithi (Burckhardt et al.); D, Paurocephala chonchaiensis Boselli. Scales 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 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).