Livia Latreille, 1802

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 : 406-407

Livia Latreille, 1802

Livia Latreille 1802: 266 .

Type species: Psylla juncorum Latreille, 1798 , by monotypy.

= Diraphia Illiger 1803: 284 .

Type species: Chermes junci Schrank, 1789 , by monotypy. Synonymized with Livia by Latreille 1807: 170.

= Diraphia Waga 1842: 275 , nec Illiger 1803.

Type species: Diraphia limbata Waga, 1842 , by monotypy.

= Neolivia Hedicke 1920: 71 ; replacement name for Diraphia Waga nec Illiger. Synonymized with Livia by Loginova 1974: 862.

= Vailakiella Bliven 1955: 13 .

Type species: Vailakiella eos Bliven, 1955 , by original designation and monotypy. Synonymized with Livia by Hodkinson and Bird 2000: 4.

Diagnosis: Adult. Head, in lateral view, directed in longitudinal axis of body ( Fig. 1D ); in dorsal view slightly narrower than thorax, about as long as wide ( Fig. 2A ). Vertex elongate, forming a lobe on either side of midline, continuing to ventral face; covered in irregularly imbricate, partly reduced microsculpture; clearly separated from genae by transverse suture ( Fig. 3F ); coronal suture fully developed; genae flattened, not enlarged into processes; frons narrowly triangular ( Fig. 3F ); median ocellus not visible in perpendicular view to vertex; subgenae differentiated into separate sclerites ( Fig. 3F ); compound eyes, in dorsal view, elongate, strongly adpressed to head. Clypeus pear-shaped, moderately large, flattened ventrally, hardly visible in lateral view as it is hidden by genae. Antenna about as long as head width; flagellum with simple setae; segment 2 longest, segment 3 shorter 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, flattened. Pronotum in dorsal view, almost straight, subrectagular; propleurites subrectangular, divided by perpendicular suture into subequal epimeron and episternum. Metapostnotum ( Fig. 4B ) forming backwards directed tooth. Mesosternum ( Fig. 5D ) relatively long; pleurosternal suture and basisternum well developed; mesosternum narrower than head, subrectangular less than twice as wide as long laterally; anterior margin with medial angle or notch; pleurosternal suture well visible; basisternum large, oval to rhomboidal; katepisternum large antero-laterally, not bent dorsad laterally; angle between arms of precoxale acute of right. Pro- and mesotibiae cylindrical. Metacoxa with long, subacute, horn-shaped meracanthus. Metafemur with three ventral sense organs positioned at midlength; apex lacking stout long setae. Metatibia distinctly widened apically; with three to eight strongly sclerotized apical spurs that are relatively evenly spaced usually arranged to form a posteriorly open crown, laterally on short processes; with several irregularly spaced thorn-like setae postero-apically and many stout spines ( Fig. 7D ). Both metatarsal segments relatively short, subequal in length. Forewing oblong-oval to ovoid; widest in the middle or in apical third, 1.6–2.3 times as long as wide, rugose, opaque; vein C + Sc relatively evenly convex, slender, distinctly delimited from cell; costal break absent; pterostigma absent or small; nodal line not developed; vein R shorter than M + Cu; vein Rs relatively straight or slightly sinuous, turned towards costal margin apically; vein M longer than its branches; vein Cu 1a strongly curved towards anal margin; veins M 1 + 2 and M 3 + 4 relatively perpendicular to wing margin apically; anal break adjacent to apex of vein Cu 1b; surface spinules coarse, often restricted to wing margins. Hindwing slightly shorter than forewing; with two to three costal setae proximal to costal break and two indistinct groups of four to five dense proximal and three to four spaced distal setae, distal to costal break; vein R + M + Cu bifurcating into R and M + Cu. Abdominal base lacking 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 to 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 ventral or subterminal position; with or without additional pore fields developed.

Comments: Monophyletic with strong molecular support ( Fig. 10 ; Supporting Information, File S4) and morphologically supported by four synapomorphies ( Fig. 11 ). The generic concept of Hodkinson and Bird (2000) 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 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 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 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).