Haplaphalara Uichanco, 1912

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 : 408-409

Haplaphalara Uichanco, 1912 , stat. rev.

Haplaphalara Uichanco 1921: 260 .

Type species: Aphalara dahli Rübsaamen, 1905 , by original designation.

= Aconopsylla Tuthill and Taylor 1955: 247 .

Type species: Psylla sterculiae Froggatt 1901 , by original designation. Syn. nov.

Diagnosis: Adult. Head, in lateral view, deflexed 45–90° from longitudinal axis of body ( Fig. 1G ); in dorsal view about as wide as or slightly narrower than thorax, moderately transverse. Vertex rhomboidal; covered in areolate-rugose or imbricate microsculpture, sometimes much reduced on disc; passing smoothly into genae anteriorly; coronal suture fully developed; genaeweaklyproducedventrallybutnotenlargedintoprocesses; 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- to large-sized, flattened ventrally, not or hardly visible in lateral view as it is hidden by genae. Antenna 10-segmented, distinctly longer than head width; flagellum with simple setae; segment 3 longest, longer than segment 4 and shorter than 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, weakly curved. Pronotum, in dorsal view, weakly curved posteriad laterally; propleurites subrectangular, divided by oblique suture into larger epimeron and smaller episternum. Metapostnotum with blunt tubercle or shallow longitudinal ridge. Mesosternum narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave; pleurosternal suture not or hardly visible; basisternum indistinct, but large, oval in H. irvingiae and H. Laos; katepisternum small antero-laterally, not bent dorsad laterally; angle between arms of precoxale obtuse. Pro- and mesotibiae cylindrical. Metacoxa with blunt or subacute horn-shaped meracanthus. Metafemur with the three ventral sense organs in medial or submedial position ( Fig. 6I, J ); apex with a group of stout long setae. Metatibia longer than metafemur, hardly to distinctly widened apically; bearing 6–10 evenly spaced to distinctly grouped apical sclerotized spurs that may be on raised processes, without unsclerotized bristle-like setae ( Fig. 7G ). Both metatarsal segments relatively short, subequal in length. Forewing oblong-oval, widest in the middle or in apical third; 2.2–2.4 times as long as wide, membranous; vein C + Sc weakly, evenly convex, slender, distinctly delimited from cell; costal break developed, close apex of vein R 1; pterostigma wide, entirely membranous; nodal line developed in some species; vein R as long as or shorter than M + Cu; vein Rs weakly or moderately 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 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 almost completely lacking. Hindwing slightly shorter than forewing; with one or two costal setae proximal to costal break and two distinct groups one to four proximal and two to five distal setae, distal to costal break; vein R + M + Cu indistinctly trifurcating, base of vein Cu indistinct. Abdominal base usually with a sclerotized area on either side covered in spines. Aedeagus with proximal portion simple or subdivided subapically; apex of distal portion differentiated from stem, though sometimes indistinctly ( Fig. 9E ). Female subgenital plate with or without 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 lacking densely spaced simple setae or sectasetae. Anus in terminal position; with additional pore fields developed.

Comments: In all analyses, Haplaphalara as redefined here is an artificial, paraphyletic assemblage ( Figs 10 , 11 ; Supporting Information, File S4). It is morphologically similar to Diclidophlebia and Melanastera from which it differs in the ungrouped apical metatibial spurs and the apex of the metatibia lacking unsclerotized antero-apical bristle-like setae. Except for the strong support in the molecular analyses for H. irvingiae + H. Laos as sister-taxa, there are no consistently supported groupings warranting generic rank within this assemblage. We, therefore, keep it as an artificial, paraphyletic genus awaiting further analyses. Included species, distribution, and host plants are summarized in Table 3 and Supporting Information, File S3. The following new or revived combinations are proposed here: Haplaphalara adelaidae ( Braza and Calilung 1981: 344) , comb. rev. (from Paurocephala ; Haplaphalara, Navasero and Calilung 1998: 14 ; Diclidophlebia, Burckhardt and Mifsud 2003: 13 ); H. dahli ( Rübsaamen 1905: 23) , comb. rev. ( Aphalara ; Haplaphalara, Uichanco 1921: 261 ; Strophingia, Crawford 1925: 40: 40 ; Diclidophlebia, Burckhardt and Mifsud 2003: 13 ); H. dombeyae (Burckhardt et al., 2006: 369) , comb. nov. ( Diclidophlebia ); H. grewiae ( Kandasamy 1986: 61) , comb. nov. ( Paurocephala ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); H. irvingiae (Burckhardt et al., 2006: 376) , comb. nov. ( Diclidophlebia ); H. maculata ( Crawford 1919: 151) , comb. rev. ( Paurocephala ; Haplaphalara, Loginova 1972: 841 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); H. menoni ( Mathur 1975: 50) , comb. rev. ( Paurocephala ; Haplaphalara, Hollis 1984: 28 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); H. parinari (Burckhardt et al., 2006: 382) , comb. nov. ( Diclidophlebia ); H. sterculiae ( Froggatt 1901: 255) , comb. nov. ( Psylla ; Aconopsylla, Tuthill and Taylor 1955: 247 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); H. trimaculata ( Mathur 1975: 69) , comb. rev. ( Paurocephala ; Haplaphalara, Hollis 1984: 28 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); H. tuberculata (Burckhardt et al., 2006: 385) , comb. nov. ( Diclidophlebia ); H. wagneri (Burckhardt et al., 2006: 386) , comb. nov. ( Diclidophlebia ).

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 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).