Klyveria 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 : 409-410

Klyveria Burckhardt, Serbina and Malenovský gen. nov.

Zoobank registration: urn:lsid:zoobank.org:act:568C5A77-9010-4F66-B83E-575317BA95A5 .

Type species: Paurocephala crassiflagellata Burckhardt, 1996 , by present designation.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1H ); in dorsal view slightly wider than thorax, very short and wide. Vertex rhomboidal, moderately to strongly transverse ( Fig. 2B ); covered in finely granular 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 ( Fig. 3G ); median ocellus visible in perpendicular view to vertex; compound eyes, in dorsal view, subspherical, collared. Clypeus pear-shaped, large, flattened ventrally, hardly visible in lateral view as it is hidden by the genae. Antenna 10-segmented, distinctly longer than head width; flagellum with simple setae; segment 3 thickened, longest, about as long as segments 4–5 together; segments 3, 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 narrowly subrectangular, divided by perpendicular suture into subequal epimeron and episternum. Metapostnotum with blunt tubercle. Mesosternum ( Fig. 5C ) narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave with median hump; pleurosternal suture not visible; basisternum triangular; katepisternum small antero-laterally, not bent dorsad laterally; angle between arms of precoxale right or acute. Pro- and mesotibiae cylindrical. Metacoxa with subacute horn-shaped meracanthus. Metafemur with the three ventral sense organs in medial position ( Fig. 6K ); apex with a group of stout, long setae. Metatibia longer than metafemur, hardly widened apically; bearing 12 or 13 densely spaced apical, weakly sclerotized spurs, without posterior peg-likw or thorn-like setae ( Fig. 7I ). Both metatarsal segments relatively short, subequal in length. Forewing oval or oblong-oval, widest in apical third; 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 narrow or wide, entirely membranous; nodal line developed; veins R and M + Cu subequal; vein Rs weakly convex relative to costal margin or sinuous; vein M longer than M 1 + 2; vein Cu 1a weakly or 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 spinules fine, spaced, present in all cells. Hindwing slightly shorter than forewing; with two or three costal setae proximal and five to eight setae distal to costal break; vein R + M + Cu bifurcating into R and M + Cu. Abdominal base with a sclerotized area on either side covered in spines. Male proctiger simple ( Fig. 9A ). Aedeagus with simple proximal portion bearing many weak folds subapically; distal portion moderately to strongly expanded basally, apex subdivided from stem ( Fig. 9G ). Female subgenital plate bearing apical process.

Last instar immature. Antenna 10 segments; bearing sectasetae or lanceolate setae on antennal flagellum. Mid- and hindlegs without massive peg-like setae. Tarsal arolium small, pedicel with lateral wings ( Fig. 9L ). Dorsal body surface bearing minute clavate setae. Precaudal abdominal tergites lacking densely spaced simple setae or sectasetae. Anus in terminal position; with additional pore fields developed.

Etymology: Dedicated to the US entomologist Frederick Detlev Klyver for his exemplary contributions to psyllid taxonomy.

Comments: Monophyly strongly supported in all analyses ( Figs 10 , 11 ; Supporting Information, File S4). Klyveria currently includes two described species: K. crassiflagellata ( Burckhardt 1996: 79) , comb. nov. (from Paurocephala ; Diclidophlebia, Burckhardt and Mifsud 2003: 13 ) and K. setinervis ( Burckhardt 1996: 78) , comb. nov. (from Paurocephala ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ). Their 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 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 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).