Liella Burckhardt, Serbina and Malenovský, 2024

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 : 410-411

Liella Burckhardt, Serbina and Malenovský gen. nov.

Zoobank registration: urn:lsid:zoobank.org:act:F4331451-EC37-4E7D-BC2B-EAD32D6F198A .

Type species: Paurocephala urenae Russell, 1946 , by present designation.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1I ); in dorsal view slightly narrower than thorax, moderately transverse. Vertex rhomboidal; covered in imbricate microsculpture; passing smoothly into genae anteriorly; coronal suture fully developed; genae weakly produced ventrally but not enlarged into processes; frons widely trapezoidal; median ocellus visible or hidden in perpendicular view to vertex; compound eyes, in dorsal view, hemispherical, adpressed to head. Clypeus pear-shaped, medium-sized, rounded ventrally, hardly to well visible in lateral view. Antenna 10-segmented, distinctly shorter to longer than head width; flagellum with simple setae; segment 3 longest, longer than segment 4 and shorter than segments 4–6 together; segments 4, 6, 8, and 9 bearing each a subapical rhinarium lacking marginal spines. Pronotum, in dorsal view, almost straight, subrectagular, weakly curved posteriad laterally; propleurites narrowly subrectangular, divided by perpendicular suture into larger epimeron and smaller episternum. Metapostnotum with subacute tubercle. Mesosternum narrower than head, forming transverse band about three times as wide as long laterally; anterior margin weakly concave with median hump; pleurosternal suture not visible; basisternum large, oval to rhomboidal; katepisternum small antero-laterally, not bent dorsad laterally; angle between arms of precoxale acute or right. Metacoxa with short, tubular, apically blunt meracanthus ( Fig. 6C ). Metafemur with the three ventral sense organs in submedial position ( Fig. 6L ); apex with a group of stout long setae.Metatibia as long as or longer than metafemur, hardly widened apically, without spur-like setae along length of metatibia; bearing seven or eight evenly spaced, weakly sclerotized apical spurs, without posterior peg-like or thorn-like setae ( Fig. 7H ). Both metatarsal segments relatively short; subequal in length or basal segment distinctly longer than apical one. Forewing oblong-oval, widest in the middle or in apical third; 2.2–2.7 times as long as wide, membranous; vein C + Sc weakly convex, slender, distinctly delimited from cell; costal break developed, close to apex of vein R 1; pterostigma narrow or wide, entirely membranous; nodal line developed; vein R as long as or shorter than M + Cu; vein Rs weakly or strongly convex relative to costal margin; vein M longer than M 1 + 2; vein Cu 1a curved towards anal margin; veins M 1 + 2 and M 3 + 4 perpendicular or oblique to wing margin apically; anal break adjacent to apex of vein Cu 1b; surface spinules ( Fig. 8F ) fine or coarse, spaced or dense, present in all cells, or sometimes partially lacking. Hindwing slightly shorter than forewing; with one to four costal setae proximal to costal break and two distinct groups distal to costal break, with two or three setae proximally and three to five setae distally ( Fig. 8J ); vein R + M + Cu indistinctly trifurcating, base of vein Cu indistinct. Abdominal base with a sclerotized area on either side covered in spines. Male proctiger with elongate or small rounded lateral plates posteriorly ( Fig. 9B ). Aedeagus ( Fig. 9H ) with simple, subapically smooth proximal portion; apex of distal portion differentiated from stem. Female subgenital plate bearing apical process.

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

Etymology: Dedicated to the Chinese entomologist Li Fasheng for his fundamental contribution to the taxonomy of Chinese psyllids.

Comments: Monophyly strongly supported in the molecular analyses ( Fig. 10 ; Supporting Information, File S4), and weakly in the morphological analysis by one homoplastic character that we consider a non-unique synapomorphy ( Fig. 11 ). Included species, distribution, and host plants are summarized in Table 3 and Supporting Information, File S3. The following new combinations are proposed here: Liella abutili ( Mifsud and Burckhardt 2002: 1943) , comb. nov. (from Paurocephala ); L. boxi ( Mifsud and Burckhardt 2002: 1949) , comb. nov. ( Paurocephala ); L. gossypii ( Russell 1943: 115) , comb. nov. ( Paurocephala ); L. hollisi ( Mifsud and Burckhardt 2002: 1956) , comb. nov. ( Paurocephala ); L. insolita ( Mifsud and Burckhardt 2002: 1956) , comb. nov. ( Paurocephala ); L. lanceomedia ( Brown and Hodkinson 1988: 37) , comb. nov. ( Paurocephala ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); L. lienhardi ( Mifsud and Burckhardt 2002: 1959) , comb. nov. ( Paurocephala ); L. lucida ( Mifsud and Burckhardt 2002: 1960) , comb. nov. ( Paurocephala ); L. medleri ( Mifsud and Burckhardt 2002: 1964) , comb. nov. ( Paurocephala ); L. paucivena ( Brown and Hodkinson 1988: 39) , comb. nov. ( Paurocephala ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); L. sinuata ( Mifsud and Burckhardt 2002: 1974) , comb. nov. ( Paurocephala ); L. urenae ( Russell 1946: 94) , comb. nov. ( Paurocephala ).

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 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 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 9. Male terminalia and tarsal arolium of immature Liviinae: A, G, L, Klyveria crassiflagellata (Burckhardt); B, H, Liella cf. insolita (Mifsud and Burckhardt); C, Diclidophlebia fremontiae (Klyver); D, Diclidophlebia leptonychiae Burckhardt et al.; E, Haplaphalara dahli (Rübsaamen); F, Haplaphalara irvingiae (Burckhardt et al.); I, Melanastera lucens (Burckhardt et al.); J, Psyllopsis fraxini (Linnaeus); K, Livia junci (Schrank); N, Melanastera Brazil. A, B, male proctiger, in lateral view; C–I, apex of proximal portion and distal portion(s) of aedeagus; J–N, tarsal arolium of fifth instar immature.Scales A, B = 0.05 mm; C–I = 0.05 mm; J–N, = 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).