Liviidae

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 : 413-415

publication ID	https://doi.org/10.1093/zoolinnean/zlad128
publication LSID	lsid:zoobank.org:pub:35B2566-E5C4-4C18-BCDC-550464F33B1E
DOI	https://doi.org/10.5281/zenodo.13304597
persistent identifier	https://treatment.plazi.org/id/03A58E6A-7441-FFF7-FE0C-05906DD32D66
treatment provided by	Plazi
scientific name	Liviidae
status

Treatment

Key to subfamilies of Liviidae View in CoL View at ENA and genera of Liviinae : adults

1. Three ventral sense organs on metafemur in basal position ( Fig. 6E View Figure 6 ) ......................................................................... Euphyllurinae

- Three ventral sense organs on metafemur in medial or submedial position ( Fig. 6F–N View Figure 6 ) ...............................................................2

2. Metabasitarsus bearing two sclerotized spurs.........................................................................................Neophyllurinae: Neophyllura View in CoL

- Metabasitarsus lacking sclerotized spurs… Liviinae ................................................................................................................................3

3. Head with small preocular sclerite between eye and antennal insertion ( Fig. 3B View Figure 3 ). Apex of metatibia with thorn-like setae posteriorly ( Fig. 7B, D View Figure 7 ) or with one to five peg-like setae laterally adjacent to a spur ( Fig. 7C View Figure 7 ); apical spurs always sclerotized, relatively evenly spaced, forming a posteriorly open crown ( Fig. 7B View Figure 7 )… Liviini ................................................................................4

- Head without preocular sclerite ( Fig. 3A View Figure 3 ). Apex of metatibia lacking thorn-like setae posteriorly or peg-like setae adjacent to apical spurs; unsclerotized bristle-like setae may be present anteriorly between apical spurs ( Fig. 7J, K View Figure 7 ); apical spurs are sometimes grouped ( Fig. 7E, F View Figure 7 ) or weakly sclerotized ( Fig. 7H, I, L View Figure 7 )…Paurocephalini................................................................8

4. Pterostigma of forewing membranous, broad, and long ( Fig. 8C View Figure 8 ). Metafemur with a group of stout, long setae apically ( Fig. 6O View Figure 6 ). Apex of metatibia bearing a row of two to five peg-like setae adjacent to a spur laterally ( Fig. 7C View Figure 7 ) ....................................5

- Pterostigma of forewing leathery, narrow and short or reduced ( Fig. 8A View Figure 8 ). Metafemur lacking stout, long setae apically. Metatibia either with a single peg-like seta adjacent to the most lateral apical spur or with several irregularly spaced thorn-like setae postero-apically ( Fig. 7B, D View Figure 7 ) ......................................................................................................................................................6

5. Frons triangular ( Fig. 3C View Figure 3 ). Antennal segment 3 as long as, or longer than, segments 4–6 together. Forewing with vein M much longer than M 1 + 2 .............................................................................................................................................................. Anomoterga

- Frons elongate, parallel-sided ( Fig. 3E View Figure 3 ). Antennal segment 3 not longer than segments 4–6 together. Forewing with vein M about as long as M 1 + 2 ............................................................................................................................................................ Camarotoscena View in CoL

6. Subgenae not differentiated into separate sclerites. Metatibia with 9 or 10 apical metatibial spurs and one peg-like seta adjacent to inner spurs, lacking thorn-like setae postero-apically............................................................................................... Syntomoza View in CoL

- Subgenae differentiated into a separate sclerite on either side ( Fig. 3D, F View Figure 3 ). Metatibia with three to eight apical metatibial spurs, bearing thorn-like setae postero-apically, lacking peg-like setae adjacent to inner spur ( Fig. 7B, D View Figure 7 )................................7

7. Head with transverse carina between vertex and genae anteriorly ( Fig. 2D View Figure 2 ); vertex lacking anterior lobes. Antennal segment 3 longer than any other segment .................................................................................................................................................. Aphorma View in CoL

- Vertex forming lobes anteriorly ( Fig. 2A View Figure 2 ). Antennal segment 2 longer than any other segment… Livia View in CoL

8. Apical metatibial spurs weakly sclerotized, pale brown..........................................................................................................................9

- Apical metatibial spurs strongly sclerotized, black ............................................................................................................................... 12

9. Metatibia with stout spurs laterally similar to those apically ( Fig. 7L View Figure 7 ).......................................................................... Paurocephala View in CoL

- Metatibia devoid of stout spurs laterally ( Fig. 7H, I View Figure 7 ) ........................................................................................................................... 10

10. Body conspicuously flattened; wings held almost horizontally ( Fig. 1L View Figure 1 ). Forewing widest in basal third. Antenna with a single subapical rhinarium on each of segments 4 and 6–9; segments 3–5 with fine bifid setae ...................................... Woldaia

- Body not conspicuously flattened; wings held roof-like over the body ( Fig. 1H, I View Figure 1 ). Forewing widest in the middle or apical third. Antenna with a single subapical rhinarium on each of segments 4, 6, 8, 9, and sometimes 3; lacking bifid setae ........ 11

11. Male proctiger lacking elongate or small rounded lateral plates posteriorly ( Fig. 9A View Figure 9 ). Distal portion of aedeagus often strongly swollen basally ( Fig. 9G View Figure 9 ). Head often strongly transverse, vertex about a quarter or a third as long as vertex width ( Fig. 2B View Figure 2 ). Clypeus large, truncate anteriorly................................................................................................................................ Klyveria

- Male proctiger with elongate or small rounded lateral plates posteriorly ( Fig. 9B View Figure 9 ). Distal portion of aedeagus hardly swollen basally ( Fig. 9H View Figure 9 ). Head moderately transverse, vertex about a third to half as long as vertex width. Clypeus small or medium-sized, rounded anteriorly....................................................................................................................................................................... Liella

12. Vein C + Sc of forewing widened, not clearly delimited from cell; pterostigma divided into a basal or anterior coriaceous part and an apical or posterior membranous part (sometimes indistinct) ( Fig. 8B View Figure 8 , arrow). Metatibia with distinctly grouped spurs as (1–2) + (3–5) and/or lateral spurs on thumb-like processes longer than spurs ( Fig. 7E, F View Figure 7 ).................. Diclidophlebia View in CoL

- Vein C + Sc of forewing not widened, clearly delimited from cell; pterostigma uniform, never divided into a basal coriaceous and apical membranous part ( Fig. 8C View Figure 8 ). Metatibia with ungrouped or grouped apical spurs, lateral spurs rarely on thumb-like processes ( Fig. 7G, J, K View Figure 7 ) ............................................................................................................................................................................. 13

13. Metapostnotum with small tubercle or shallow longitudinal ridge ( Fig. 4A View Figure 4 ). Mesosternum shallowly concave along anterior margin; pleurosternal suture indistinct; basisternum large oval to rhomboidal or indistinct; katepisternum small antero-laterally. Apex of metatibia with a posteriorly open crown of 7–11, evenly or unevenly spaced spurs of similar size; lacking weakly sclerotized bristle-like setae between spurs ( Fig. 7G View Figure 7 )........................................................................................ Haplaphalara

- Metapostnotum with laterally compressed tooth ( Fig. 4C View Figure 4 ). Mesosternum concave anteriorly with hump in the middle; pleurosternal suture well visible; basisternum large, triangular; katepisternum large antero-laterally. Apex of metatibia with 5–10, sometimes indistinctly, grouped spurs; with antero-lateral unsclerotized bristle-like setae separating apical spurs ( Fig. 7J, K View Figure 7 ) .............................................................................................................................................................................................. Melanastera

Key to genera of Liviinae : immatures

1. Tarsal arolium fan-shaped, sessile ( Fig. 8K View Figure 8 ). Antennal flagellum lacking sectasetae; antenna seven or eight segments… Liviini ................................................................................................................................................................................................................2

- Tarsal arolium petiolate; petiole often with lateral appendages; arolium small relative to pedicel ( Fig. 8L, N View Figure 8 ). Antennal flagellum bearing sectasetae; if absent, then antenna three segments…Paurocephalini .....................................................................6

2. On Salicaceae View in CoL (and perhaps Cunoniaceae View in CoL )...............................................................................................................................................3

- On other host families ...................................................................................................................................................................................5

3. Anus in terminal position............................................................................................................................................................. Syntomoza View in CoL

- Anus in ventral position ................................................................................................................................................................................4

4. Mid- and hindlegs with massive peg-like spurs ..................................................................................................................... Anomoterga

- Mid- and hindlegs lacking massive peg-like spurs .......................................................................................................... Camarotoscena View in CoL

5. Caudal plate bearing dorsal and marginal sectasetae. On Ranunculaceae View in CoL ........................................................................... Aphorma View in CoL

- Caudal plate lacking sectasetae. On Cyperaceae View in CoL and Juncaceae View in CoL .................................................................................................... Livia View in CoL

6. Body usually lacking minute clavate setae dorsally. Abdomen never with pore fields in addition to circumanal ring ..............7

- Body bearing minute clavate setae dorsally. Abdomen usually bearing pore fields in addition to circumanal ring ...................8

7. Antennal flagellum never subdivided. Sectasetae on outer margin of abdomen based upon extended tubercles, if indistinct then antennal flagellum lacking sectasetae ........................................................................................................................................ Liella

- Antennal flagellum usually (indistinctly) subdivided; always with sectasetae. Sectasetae on outer margin of abdomen not based upon extended tubercles.............................................................................................................................................. Paurocephala View in CoL

8. On Luehea spp. ( Malvaceae View in CoL ) ........................................................................................................................................................................9

- On other plant taxa ..................................................................................................................................................................................... 10

9. Legs bearing sectasetae..................................................................................................................................................................... Klyveria

- Legs lacking sectasetae ..................................................................................................................................................................... Woldaia

10. Antenna with 10 segments; if nine segments then on Asteraceae View in CoL ..................................................................................... Melanastera

- Antenna with nine segments. Never on Asteraceae View in CoL ........................................................................ Diclidophlebia View in CoL and Haplaphalara

Species excluded from Liviidae View in CoL

Cockerell (1915) described a fragment of a forewing from the Oligocene of the Isle of Wight as † Necropsylla anglica Cockerell, 1915 . Bekker-Migdisova (1985) transferred the species provisionally to Camarotoscena based on the long, only weakly curved vein Cu 1a and the straight vein Cu 1b. These characters are not diagnostic for Camarotoscena but occur also in † Lapidopsylla Klimaszewski, 1993 ( Aphalaridae , Aphalarinae , †Paleopsylloidini), erected for two species from the Eocene/ Oligocene of the Isle of Wight. † Necropsylla anglica resembles the type species of † Lapidopsylla , † L. thornessbaya Klimaszewski, 1993 , with which it appears congeneric. We suggest the following new combination: † Lapidopsylla anglica ( Cockerell 1915: 487) , comb. nov. from Necropsylla ; Camarotoscena, Bekker-Migdisova 1985: 81 .

Syntomoza lebezia Hodkinson, 1986 View in CoL was described from a single specimen from Belize. Hodkinson (1986) placed this species provisionally in Syntomoza View in CoL as it shares with S. magna View in CoL the lobed male proctiger and the bulbous apex of the distal portion of the aedeagus, but differs in the head, metatarsi, and terminalia. As the species is morphologically quite distinct, Hodkinson (1986) suggested that it ‘will probably deserve generic status in its own right’. Burckhardt and Mifsud (2003) concluded that the species ‘should be excluded from Syntomoza View in CoL and the Paurocephalinae’ and suggested some resemblance in the head to Metapsylla Kuwayama, 1908 View in CoL . However, S. lebezia View in CoL shares with Katacephala View in CoL ( Psyllidae View in CoL : Katacephalinae) the metatibia lacking a genual spine and bearing a posteriorly open crown of apical spurs, the presence of two basimetatarsal spurs, the posteriorly lobed male proctiger, and the host association with Myrtaceae View in CoL (unpublished BMNH data). For this reason, we transfer Syntomoza lebezia Hodkinson View in CoL formally to Katacephala View in CoL , as Katacephala lebezia ( Hodkinson 1986: 149) , comb. nov.

The Afrotropical species P. bicarinata Pettey, 1924 View in CoL and P. hottentotti Pettey, 1933 View in CoL are currently formally included in Paurocephala View in CoL . They possess basimetatarsal spurs and are, therefore, not members of the Liviinae ( Burckhardt and Mifsud 2003).