Paurocephala Crawford, 1913

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 : 412

Paurocephala Crawford, 1913

Paurocephala Crawford 1913: 293 .

Type species: Paurocephala psylloptera Crawford, 1913 , by original designation.

= Paurocephala subgenus Thoracocorna Klimaszewski 1970: 427 .

Type species: Paurocephala chonchaiensis Boselli, 1929 , by original designation. Synonymized with Paurocephala by Loginova 1972: 842.

= Marpsylla Navasero and Calilung 2001: 126 .

Type species: Marpsylla baltazarae Navasero and Calilung, 2001 , by original designation. Synonymized with Paurocephala by Burckhardt and Ouvrard 2012: 16.

= Pauroterga Navasero 2010: 20 .

Type species: Pauroterga zamboangensis Navasero, 2010 , by original designation. Synonymized with Paurocephala by Burckhardt and Ouvrard 2012: 16.

Diagnosis: Adult. Head, in lateral view, deflexed 45–80° from longitudinal axis of body ( Fig. 1K ); in dorsal view, about as wide as or slightly narrower than thorax, moderately to strongly 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 elliptical ( Fig. 3H ); median ocellus visible or hidden in perpendicular view to vertex; compound eyes, in dorsal view, hemispherical, adpressed to head. Clypeus pear-shaped, medium-sized to large, rounded ventrally, visible in lateral view. Antenna 8–10 segments, shorter to much longer than head width; flagellum usually with simple setae, rarely with a bifid seta associated the rhinaria; in 10-segmented antenna, segment 3 longest, longer than segments 4–5 together and shorter than segments 4–7 together, and segments 4, 6, 8, and 9 bearing each a subapical rhinarium lacking a wreath of spines. Thorax moderately slender to relatively robust; dorsal outline, in lateral view, weakly curved. Pronotum, in dorsal view, weakly curved posteriad laterally; propleurites subrectangular, divided by perpendicular suture into larger epimeron and smaller episternum. Metapostnotum ( Fig. 4D ) with conical horn. Mesosternum ( Fig. 5E ) narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave; pleurosternal suture not visible; basisternum large, oval to rhomboida; katepisternum large antero-laterally, bent dorsad laterally; angle between arms of precoxale obtuse. Pro- and mesotibiae cylindrical. Metacoxa with slender, tubular, apically blunt meracanthus ( Fig. 6D ). Metafemur with the three ventral sense organs in submedial position ( Fig. 6N ); apex with a group of stout long setae. Metatibia shorter or longer than metafemur, hardly widened apically, with weakly sclerotized spur-like setae along length of metatibia similar to those that are at apex; bearing seven or eight evenly spaced, apical, weakly sclerotized spurs, without posterior peg-like or thorn-like setae ( Fig. 7L ). Basal metatarsal segments as long as or longer than apical segment. Forewing oblong-oval, widest in the middle or in apical third; 2.1–2.5 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 usually as long as but rarely shorter than M + Cu; vein Rs weakly or strongly convex relative to costal margin or sinuous; 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 fine or coarse, spaced or dense, present in all cells, or sometimes partially lacking. Hindwing slightly shorter than forewing; with one to three costal setae proximal to costal break and two distinct groups distal to costal break, with one or two setae proximally and two to five 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, sometimes forming finger-like process ( Fig. 8O, P ). Aedeagus with proximal portion strongly subdivided subapically; apex of distal portion differentiated from stem. Female subgenital plate bearing apical process.

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

Comments: The monophyly of Paurocephala is strongly supported in all molecular analyses ( Fig. 10 ; Supporting Information, File S4) and by four synapomorphies in the morphological analysis ( Fig. 11 ). Included 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 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 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).