Diclidophlebia Crawford, 1919

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 : 408

Diclidophlebia Crawford, 1919

Heteroneura Crawford 1919: 152 ; nec Fallén 1810: 7, 25.

Type species: Heteroneura oceanica Crawford, 1919 , by original designation and monotypy.

Diclidophlebia Crawford 1920: 355 ; replacement name for Heteroneura Crawford nec Fallén.

=Gyroza Enderlein 1921: 122; replacement name for Heteroneura Crawford nec Fallén. Objective junior synonym of Diclidophlebia .

= Paraphalaroida Loginova 1972: 851 .

Type species: Paurocephala fremontiae Klyver, 1930 , by original designation. Synonymized by Burckhardt and Mifsud 2003: 12.

= Sinuonemopsylla Li and Yang 1991: 11 .

Type species: Sinuonemopsylla excetrodendri Li and Yang, 1991 , by original description and monotypy. Synonymized by Burckhardt and Mifsud 2003: 12.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1F ); in dorsal view, moderately transverse, slightly or much narrower than thorax. Vertex rhomboidal to almost subrectangular; covered in imbricate microsculpture, sometimes much reduced on disc or smooth; passing smoothly into genae anteriorly; coronal suture ranging from fully developed to completely absent; genae weakly produced ventrally but not enlarged into processes; frons widely trapezoidal; median ocellus visible in perpendicular view to vertex; compound eyes, in dorsal view, hemispherical, adpressed to head. Clypeus pear-shaped, large, rounded ventrally, well-visible in lateral view or slightly hidden by genae. Antenna 10-segmented, distinctly longer than 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 marginal spines. Thorax moderately slender to massive; dorsal outline, in lateral view, weakly to strongly curved. Pronotum, in dorsal view, weakly curved posteriad laterally; propleurites narrowly subrectangular, divided by perpendicular suture into larger epimeron and smaller episternum. Metapostnotum with laterally compressed tooth, but small in D. leptonychiae . Mesosternum usually narrower, rarely wider (e.g. D. eastopi ) than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave; pleurosternal suture not visible; basisternum indistinct; katepisternum small antero-laterally, not bent dorsad laterally; trochantins in obtuse angle to each other. Pro- and mesotibiae cylindrical or robust and flattened ( D. eastopi , D. leptonychiae , D. oceanica ). Metacoxa with blunt or subacute horn-shaped meracanthus. Metafemur with the three ventral sense organs in medial or submedial position ( Fig. 6F–H ); apex with a group of stout long setae. Metatibia longer than metafemur, slightly widened apically; bearing 4–11 irregularly spaced to distinctly grouped apical sclerotized spurs that may be on raised processes, without unsclerotized bristle-like setae ( Fig. 7E, F ). Both metatarsal segments relatively short, subequal in length. Forewing oval to oblong-oval; widest in the middle, 1.8–2.5 times as long as wide, membranous or subcoriaceous; vein C + Sc almost straight to strongly convex, widened, indistinctly delimited from cell; costal break developed, close apex of vein R 1; pterostigma wide, coriaceous basally, membranous apically ( Fig. 8B ); nodal line sometimes developed; veins R and M + Cu usually subequal, sometimes vein R slightly shorter than M + Cu; vein Rs almost straight, strongly curved or convoluted; vein M shorter or longer than M 1 + 2; veins M 1 + 2, M 3 + 4 and Cu 1a variable; anal break adjacent to apex of vein Cu 1b; surface spinules fine, spaced or dense, present in all cells. Hindwing slightly shorter than forewing; with two to four costal setae proximal to costal break and 6–18 irregularly spaced setae distal to costal break ( Fig. 8I ); vein R + M + Cu bifurcating into R and M + Cu. Abdominal base with a sclerotized area on either side covered in spines ( Fig. 8N ). Aedeagus with simple proximal portion subapically bearing many weak folds or subdivided; apex of distal portion differentiated from stem ( Fig. 9C ). Female subgenital plate lacking long apical process.

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

Comments: Polyphyletic in the molecular analyses ( Fig. 10 ; Supporting Information, File S4), although the genus as delimited here is recovered in the morphological analysis supported by one synapomorphy ( Fig. 11 ). This concept of Diclidophlebia is much more restricted compared to that of Burckhardt and Mifsud (2003) who included species now referred to Diclidophlebia s.s., Haplaphalara , Klyveria , Melanastera , and Woldaia . 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 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).