Melanastera Serbina, Malenovský, Queiroz and Burckhardt, 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 : 411

Melanastera Serbina, Malenovský, Queiroz and Burckhardt gen.nov.

Zoobank registration: urn:lsid:zoobank.org:act:4201F6CF-FBF3-4BBD-BA1E-B9EBF39F9137 .

Type species: Diclidophlebia lucens Burckhardt et al., 2005 , by present designation.

Diagnosis: Adult. Head in lateral view, deflexed 45–80° from longitudinal axis of body ( Fig. 1J ); 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 ( Fig. 2C ); 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, hardly visible in lateral view as it is hidden by genae. Antenna 10 segments, slightly to distinctly 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 usually surrounded by a wreath of spines (partly lacking in Melanastera Venezuela). 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 larger epimeron and smaller episternum. Metapostnotum ( Fig. 4C ) with laterally compressed tooth. Mesosternum narrower than head, forming transverse band more than three times as wide as long laterally; anterior margin weakly concave with median hump; pleurosternal suture well visible; basisternum triangular; katepisternum large 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 position ( Fig. 6M ); apex with a group of stout long setae. Metatibia longer than metafemur, slightly widened apically; bearing 4–11 grouped apical sclerotized spurs separated by one to seven unsclerotized bristle-like setae anteriorly or antero-laterally (rarely lacking), without posterior peg-like or thorn-like setae ( Fig. 7J, K ). Both metatarsal segments relatively short, subequal in length. Forewing oval or oblong-oval, widest in the middle or in apical third; 1.9–2.7 times as long as wide, membranous; vein C + Sc weakly or strongly convex, slender, distinctly delimited from cell; costal break developed, close to apex of vein R 1; pterostigma narrow to wide, entirely membranous ( Fig. 8C ); 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. 8G ) fine or coarse, spaced or dense, present in all cells or sometimes partially reduced. Hindwing slightly shorter than forewing; with two to four costal setae proximal to costal break and two distinct groups distal to costal break, with three to six dense setae proximally and three to five spaced setae distally; vein R + M + Cu bifurcating into R and M + Cu. Abdominal base with a sclerotized area on either side covered in spines. Aedeagus with proximal portion bearing many weak folds subapically or strongly subdivided subapically; apex of distal portion differentiated from stem ( Fig. 9I ). Female subgenital plate lacking apical process.—Last instar immature. Antenna 10-segmented; bearing sectasetae or lanceolate setae on antennal flagellum. Mid- and hindlegs without massive peg-like setae. Tarsal arolium small ( Fig. 9N ). 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: Melanastera is a pun using the first syllables of some host families: Mel-astomataceae, An-nonaceae, and Astera-ceae, and is a tribute to the British musician David Bowie in reference to his final studio album ‘Blackstar’ (from Ancient Greek μέλας = black and Latin aster = star).

Comments: The monophyly of Melanastera is strongly supported in all molecular analyses if the single taxon, M. Venezuela is excluded. With the inclusion of the latter taxon the genus is not (ML-mix, Fig. 11 ), or only weakly (ML-part; Supporting Information, File S4A) or moderately (BI; Supporting Information, File S4B) supported. In the morphological analysis, the monophyly of Melanastera (including M. Venezuela) is supported by one synapomorphy (and five homoplasies) ( 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: Melanastera fava ( Brown and Hodkinson 1988: 44) , comb. nov. (from Haplaphalara ; Diclidophlebia, BurckhardtandMifsud 2003: 14 ); M. heterotrichi ( Caldwell and Martorell 1952: 605) , comb. nov. ( Paurocephala ; Haplaphalara, Brown and Hodkinson 1988: 40 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); M. longitarsata ( Brown and Hodkinson 1988: 42) , comb.nov. ( Haplaphalara ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); M. lucens ( Burckhardt, Hanson and Madrigal 2005: 742) ; comb. nov. ( Diclidophlebia ); M. maculipennis ( Brown and Hodkinson 1988: 47) , comb. nov. ( Haplaphalara ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); M. paucipunctata ( Brown and Hodkinson 1988: 40) , comb. nov. ( Haplaphalara ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ); M. pilosa (Burckhardt et al., 2006: 384) , comb. nov. ( Diclidophlebia ); M. smithi (Burckhardt et al., 2006: 242) , comb. nov. ( Diclidophlebia ); M. tuxtlaensis ( Conconi 1972: 51) , comb. nov. ( Paurocephala ; Haplaphalara, Brown and Hodkinson 1988: 40 ; Diclidophlebia, Burckhardt and Mifsud 2003: 14 ).

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 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 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 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).