Aphorma Hodkinson, 1974

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 : 405-406

Aphorma Hodkinson, 1974

Aphorma Heslop-Harrison 1949: 783 , 800; Heslop-Harrison 1952: 965; nomen nudum, no type species designated.

Aphorma Hodkinson 1974: 76 .

Type species: Aphalara bagnalli Laing, 1929 , by original designation.

= Leprostictopsylla Li 2011: 376 .

Type species: Leprostictopsylla jiuzhaiensis Li, 2011 , by original designation and monotypy. Synonymized by Burckhardt et al. 2018: 14.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1B ); in dorsal view slightly narrower than thorax, moderately transverse. Vertex rhomboidal; covered in areolate-rugose microsculpture; separated from genae by transverse carina ( Fig. 2D ); coronal suture fully developed; genae weakly produced ventrally but not enlarged into processes; frons parallel-sided to narrowly triangular; median ocellus not visible in perpendicular view to vertex; subgenae differentiated into separate sclerites ( Fig. 3D ); 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 about as long as head width; flagellum with simple setae; segment 3 longest, shorter than segments 4–5 together; segments 4, 6, 8, and 9 bearing each a subapical rhinarium lacking a wreath of spines. Thorax moderately slender; dorsal outline, in lateral view, weakly curved. Pronotum, in dorsal view, almost straight, subrectagular; propleurites subrectangular, divided by perpendicular suture into subequal epimeron and episternum. Metapostnotum medially with blunt tubercle. Mesosternum narrower than head, subrectangular, less than twice as wide as long laterally; anterior margin weakly concave; pleurosternal suture hardly visible; basisternum large, oval to rhomboidal; katepisternum large antero-laterally, not bent dorsad laterally; angle between arms of precoxale obtuse.Pro- and mesotibiae cylindrical.Metacoxa with long or short subacute horn-shaped meracanthus. Metafemur with the three ventral sense organs in the middle; apex lacking stout long setae. Metatibia longer than metafemur, widened apically; bearing seven or eight slightly irregularly spaced apical sclerotized spurs and a few thorn-like setae postero-apically ( Fig. 7B ). Both metatarsal segments relatively short, subequal in length. Forewing oblong-oval or subtrapezoidal; costal and anal margins subparallel or weakly widening towards apex, 1.7–2.3 times as long as wide, coriaceous; vein C + Sc weakly strongly convex, slender, distinctly delimited from cell; costal break absent; pterostigma reduced ( Fig. 8A ); nodal line not visible; veins R as long as or shorter than M + Cu; vein Rs weakly sinuous; vein M much longer than its branches; vein Cu 1a strongly 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 covered in flattened tubercles forming cellular pattern ( Fig. 8D ). Hindwing about as long as forewing; with one to three costal setae proximal to costal break and two indistinct groups distal to costal break, three to four dense setae proximally, and three to four spaced setae distally; vein R + M + Cu indistinctly trifurcating, base of vein Cu indistinct. Abdominal base with a sclerotized area on either side covered in spines ( Fig. 8L, M ). 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 or eight 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 lacking densely spaced simple setae or sectasetae. Anus in terminal position; with additional pore fields developed.

Comments: Represented by only one species in the molecular analyses ( Fig. 10 ; Supporting Information, File S4); supported by one synapomorphy in the morphological analysis ( Fig. 11 ). The synonymy of Leprostictopsylla with Aphorma by Burckhardt et al. (2018) is confirmed 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 2. Adult Liviinae:A, Livia junci (Schrank); B, Klyveria setinervis (Burckhardt); C, Melanastera smithi (Burckhardt et al.); D, Aphorma lichenoides (Puton); E, Camarotoscena badia Loginova. A–C, head, in dorsal view; D, E, head, in oblique lateral view, with anterior carina (D, arrow), and smooth anteriorly (E). Scales 0.1 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 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 8. Wings and abdomen of Liviinae: A, D, L, Aphorma lichenoides (Puton), male; M, A. lichenoides (Puton), female; B, Diclidophlebia excetrodendri (Li and Yang); C, G, Melanastera lucens (Burckhardt et al.); E, H, Livia junci (Schrank); F, J, Liella cf.insolita (Mifsud and Burckhardt); I, Diclidophlebia oceanica (Crawford); K, Tuthillia myrcianthis Burckhardt et al.; N, D. xuani (Messi); O, Paurocephala dayak Mifsud and Burckhardt, male; P, P.dayak, female.A, reduced pterostigma; B, pterostigma basally leathery, apically membranous (separating line see arrow); C, pterostigma entirely membranous; D–G, surface of forewing membrane in cell r 1: H–J, costa of hindwing; K–P, lateral edge of first visible abdominal tergite, base left.Scales A–C = 0.1 mm; D–G = 0.1 mm; H–J = 0.2 mm; K–P = 0.1 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).