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Sate er « gta bs » welled a ae « HANNS dle ob bicolorata (Samy) (p. 7) The acaciaebaileyanae-group Defined by characters in couplet 1 of key. This group comprises the introduced Australian species. Acizzia acaciaebaileyanae (Froggatt) (Figs 7, 8, 18-20, 294) Psylla acaciae-baileyanae Froggatt, 1901: 257; Tuthill, 1952: 257. Lectotype CO’, AUSTRALIA, designated by Yen, 1977: 15 (ANIC). Arytaina acaciae-baileyanae (Froggatt) Pettey, 1924: 21. Psylla uncata Ferris & Klyver, 1932: 53. Syntypes 0’, 9, NEw ZEALAND (NZAC) [examined]. [Synony- mised by Tuthill, 1952: 91.] Neopsylla unc[t]ata (Ferris & Klyver) Heslop-Harrison, 1949: 162. [Mis-spelling. ] Acizzia acaciaebaileyanae (Froggatt) Capener, 1970: 197; Loginova, 1977: 577; Hodkinson, 1983: 343; Morgan, 1984: 36. Psylla acaciaebaileyanae Froggatt; Yen, 1977: 15. DEscRIPTION. Coloration. General coloration of head and thorax dull orange-yellow with darker yellowish brown markings; abdominal sclerites yellowish brown, intersegmental membranes dull pale yellow; genitalia pale brown: forewing transparent with small irregular pale brown maculations; veins pale yellowish brown; antennae dull whitish yellow with apices of segments 3-7 infuscate and segments 8—10 entirely dark brown; legs pale whitish yellow, often with faint dusky markings on the femora. Structure. Head (Fig. 8) robust, genal cones very short with broadly rounded apices. Forewing (Fig. 7) oblong oval, with a well-developed pterostigma and costal break; spinules present throughout all the cells, much larger in regions where the brown maculate pattern occurs than elsewhere; cell m, elongate; cell cu, high and strongly arched towards the wing base. Metatibia with 4 or 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 18) with a short and broad tubular apical projection and with a rounded posterior lobe bearing a subsidiary finger-like projection; paramere (Fig. 19) simple, broad basally, gradually tapering to rounded apex, with a small posteriorly directed subapical denticle on inner posterior margin; aedeagus (Fig. 20) with a bulbous apex and a long ductus ejaculatorius. Female terminalia (Fig. 294) with proctiger convex throughout dorsal margin, apex bluntly acute; ventral valve wedge shaped; circumanal pore ring simple, about one-third length of proctiger; ovipositor short and broad. Measurements. See Table 1. MATERIAL EXAMINED 12 0, 13 Q, Italy: on Acacia sp. (BMNH). Host PLANT. Acacia baileyana F.v.M. and A. podalyriaefoliae Cunn. PUBLISHED DISTRIBUTION. Australia (Froggatt, 1901; Yen, 1977; Morgan, 1984); introduced to New Zealand (Tuthill, 1952; Ferris & Klyver, 1932), South Africa (Pettey, 1924; Capener, 1970), Italy (Rapisarda, 1985). ComMENTS. This species is native to Australia and has recently been introduced into Italy on ornamental Acacia trees. It is distinguished from its close relative, A. uncatoides, by the shape of the genal cones and details of the male genitalia. Acizzia uncatoides (Ferris & Klyver) (Figs 5, 6, 15—17, 293) Psylla uncatoides Ferris & Klyver, 1932: 53; Heslop-Harrison, 1949: 162; Tuthill, 1952: 89. Syntypes Oo’, Q, NEw ZEALAND (NZAC, BMNH) [examined]. Acizzia uncatoides (Ferris & Klyver) Loginova, 1977: 577; Hodkinson & White, 1981: 492; Hodkinson, 1983: 343. 6 I. D. HODKINSON & D. HOLLIS DEscriPTION. Coloration. General body colour orange throughout with paler markings on dorsum of thorax; forewing membrane pale yellow to pale amber with orange-brown maculations, veins concolorous with membrane but appearing slightly darker; antennal segments 1—3 orange-yellow, the remainder brown; legs slightly paler yellow-brown, often with a slight infuscation of the femora. Structure. Head (Fig. 6) with vertex relatively long, genal cones triangular with rounded apices. Forewing (Fig. 5) oblong-oval, with well-developed pterostigma and costal break; spinules present throughout wing; cell m, elongate; cell cu,, high and strongly arched towards the wing base. Metatibia with 4 or 5 thick black spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 15) with long tubular apical portion and a rounded posterior lobe bearing a subsidiary finger-like projection; paramere (Fig. 16) broad, with apex deflexed posteriorly; aedeagus (Fig. 17) with harpoon-shaped apex. Female terminalia (Fig. 293) short and truncate with a relatively large circumanal pore ring. Measurements. See Table 1. MATERIAL EXAMINED Several hundred OC’, @, Portugal, Israel, Chile (IDH, DB, USNM). Host PLANT. In Europe this species occurs on Acacia floribunda, while in Israel it is found on Acacia saligna. Elsewhere it has been recorded from Albizia lophantha, Acacia verniciflua, A. koaia, A. koa and A. confusa. PUBLISHED DISTRIBUTION. Australia (Hodkinson, 1983); introduced to. France (Bain et al., 1976), Italy (Arzone & Vidano, 1985), U.S.A.: California (Jensen, 1957; Koehler et al. , 1966; Madubunyi & Koehler, 1974; Munro, 1965), Hawaii (Gagne, 1971; Leeper & Beardsley, 1973; 1976), New Zealand (Tuthill, 1952), Mexico (Hodkinson & White, 1983). CoMMENTS. This species is native to Australia and has been introduced into several areas of the world where it has become a minor pest of ornamental or native acacias. Dr A. Yen (Australia) has compared our European material with material from Australia. A. uncatoides is separated from its close relative A. acaciaebaileyanae by the shape of the genal cones, the male proctiger, paramere and aedeagus. The hollisi-group Defined by characters in couplet 1 of key. This group includes the native North African or Middle East species. Acizzia hollisi Burckhardt (Figs 3, 4, 12-14, 262) Acizzia hollisi Burckhardt, 1981: 216. Holotype CO’, Saup1I ARABIA (NMB) [not examined]. DEscrIPTION. Coloration. General body coloration dark reddish brown throughout with paler markings on dorsum of thorax; intersegmental membranes of abdomen lighter orange-yellow; male proctiger often bright red, parameres whitish yellow; female terminalia dark brown; forewing membrane clear with brown colour pattern, veins brown; antennae yellowish brown, darkening towards apex; femora dark brown, remainder of legs pale yellowish brown. Immature specimens light orange-yellow. Structure. Head (Fig. 4) with vertex relatively short; genal cones shorter than vertex, triangular. Forewing (Fig. 3) oblong-oval, with well-developed pterostigma and costal break; spinules largely confined to apical half of wing and around vein Cuj; cell cu;, moderately strongly leaning towards wing base. Metatibia with 4 or 5 thick black spurs, basal metatarsus with 2 black spurs. Male proctiger (Fig. 12) with a broadly triangular posterior projection. Paramere (Fig. 13) with distinct notch in anterior margin; aedeagus (Fig. 14) with bulbous apex. Female terminalia (Fig. 262) relatively long and slender, with a relatively short anal pore ring. Measurements. See Table 1. MATERIAL EXAMINED 3 0,3 Q (paratypes), 2 0’, 1 9, Israel (BMNH, IDH). HostT PLANT. Acacia raddiana. PUBLISHED DISTRIBUTION. Saudi Arabia (Burckhardt, 1981; 1986), Israel (Burckhardt, 1981; Halperin et al., 1982). W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 7 Acizzia bicolorata (Samy) comb. n. (Figs 1, 2, 9-11) Psylla bicolorata Samy, 1973: 451. Holotype O’, Ecyrr (Agr. Coll. Cairo) [not examined]. DEscriPTION. Coloration. General body coloration orange-yellow throughout; forewing membrane pale yellow, veins concolorous; antennae orange-yellow basally, becoming brown apically; legs concolorous with body (based on a single specimen). According to Samy the general colour is bright brown with whitish markings on head and dorsum of the thorax. Structure. Head (Fig. 3) robust, genal cones shorter than vertex, with broadly rounded apices. Forewing (Fig. 2) with somewhat narrowly rounded apex; costal break and pterostigma well-developed; spinules confined to radular areas; cell cu,, high but less strongly displaced towards wing base than in uncatoides. Metatibia with 4 or 5 thick black spurs, basal metatarsus with 2 black spurs. Male proctiger (Fig. 10) with narrowly triangular posterior lobe; paramere (Fig. 11) simple, almost parallel-sided, apex pointing anteriorly; apex of aedeagus (Fig. 12) somewhat reniform. Measurements. See Table 1. MATERIAL EXAMINED 1 OD (same data as holotype), Egypt: Oweinat, xi. 1937 (Kasim) (BMNH). Host PLANT. Unknown. PUBLISHED DISTRIBUTION. Egypt (Samy, 1973). ComMENTSs. A. bicolorata is separated from the other west Palaearctic members of the genus by the lack of a forewing pattern. ARYTAININAE Crawford Arytainini Crawford, 1914: 106, in part; Heslop-Harrison, 1951: 417, in part. Arytaininae Crawford; Vondracéek, 1957: 176; Loginova, 1976a: 589; 1977: 577; White & Hodkinson, 1985: 272. The Arytaininae, as here constituted, is probably paraphyletic and overlaps with the subfamily Psyllinae. The group has been separated previously from the Psyllinae by the more quadrate propleurites, the presence of one, rather than two metatarsal spurs, the absent or rudimentary pterostigma and the genal cones which are usually in the same plane as the vertex, rather than being deflexed downwards. In addition, Arytainine species usually occur on Leguminosae whereas Psyllinae species occur usually on Rosaceae and a variety of other dicotyledonous families. However, Cyamophila, which appears to have a similar form of advanced paramere to Arytaina, has narrower propleurites, a fully developed pterostigma and the 2 metatarsal spurs more typical of the Psyllinae. Together the Arytaininae and Psyllinae are defined by the rounded forewing with the venation pattern in which veins Cu, and M possess a common stem and vein Cu, terminates adjacent to vein Cu;,. The basal metatarsus usually possesses at least one spur, the genae are usually developed into cone-like processes and the 3rd antennal segment is the longest. The hind coxa also bears a well-developed meracanthus, and the male proctiger is simple and unipartite. CYAMOPHILA Loginova Cyamophila Loginova, 1976a: 596; 1977: 582; White & Hodkinson, 1985: 272. Type-species: Psylla fabra Loginova, 1964, by original designation. DiaGnosis. Forewing (Figs 21, 26) with a well-developed pterostigma and costal break, cell cu,, relatively high and arched towards base of wing; male paramere (Figs 24, 29) with apex truncate, formed of a large anteriorly directed denticle; basal metatarsus with 2 spurs; propleurites elongate, divided by a diagonal suture; genal cones (Figs 22, 27) short, in same plane as vertex, enveloping the median ocellus; eyes (Figs 22, 27) less bulbous than in many other genera; antennae relatively stout, usually less than 1-5 x head width. CoMMENTS. Cyamophila is primarily a Palaearctic genus comprised of about 35 species distributed throughout the arid regions of Central Asia but extending into the Mediterranean Basin, China and Afghanistan, and known to feed on legumes within the papilionoid tribes Galegeae, Trifoliae, Loteae and Sophoreae. 8 I. D. HODKINSON & D. HOLLIS Key to species 1 Forewing (Fig. 21) with diffuse reddish brown cloud around apex; radular spinules dense, dark brown, at low magnification appearing as small distinct spots around the wing margin. Cell m +> long, at least as long as the stem of vein M. Vein Cup very strongly curved before its junction with the marginal vein. On Glycyrrhiza species .............0644 glycyrrhizae (Becker) (p. 8) — Forewing (Figs 26, 31, 37) without pattern, radular spinules obvious but not forming distinct brown marks. Cell m,+ , shorter than the stem of vein M. Vein Cu, less strongly curved before junction with the marginal vein. NOtOm GIy CY TAGG1 0 ie oye ee wae epee tc ee Zz 2 Forewing (Fig. 26) with surface spinules occupying all the cells. Male paramere as in Figs 28, 29. ATE IS aos sctts cect ee ent Seirus od dete aoe bee dare eae te ere prohaskai Preisner (p. 9) — Forewing (Figs 31, 37) without surface spinules except in the radular areas. Male paramere as in Pigs 342-40. NOt OD ATE VUEIS od niessinle Oo sehr couc teed tenia en ito iptcinatai ah ing «alga at aeokak 3 3. Male paramere (Figs 33, 34) relatively broad, with an anterior subapical bulge. Turkey, on ASEV ES ALUS. Obits as: aa Secs ROIS a, SAG get a stocklosai (Klimaszewski & Lodos) (p. 10) — Male paramere (Fig. 40) narrower, without an anterior subapical bulge. South-western U.S.S.R.,, On Medicago and Vicia SPCCleS: -- Fein cntna-dcc-uneeco ss eee medicaginis (Andrianova) (p. 9) Cyamophila glycyrrhizae (Becker) (Figs 21—25, 93) Psyllodes glycyrrhizae Becker, 1864: 486. Syntypes CO’, 9, U.S.S.R. [not examined]. Psylla glycyrrhizae (Becker) Low, 1881: 262; 1883: 239; Puton, 1886: 91; Horvath, 1904: 579; Oshanin, 1907: 351; 1912: 127; Aulmann, 1913: 16; Sulc, 1910a: 21; Vondraéek, 1953: 174; Klimaszewski, 1963: 427; 1968a: 230; 1969b: 222; 1973: 208; Loginova, 1964a: 465; 1968: 305; 1972b: 140; 1981: 26; Loginova & Baeva, 1972: 6; Baeva & Kankina, 1971: 84; Gegechkori, 1969a: 222; 1969b: 722; 1978: 20; Halperin et al. , 1982: 32. Cacopsylla glycyrrhizae (Becker) Klimaszewski & Lodos, 1977: 5. Cyamophila glycyrrhizae (Becker) Loginova, 1977: 582; 1981: 26; Klimaszewski & Lodos, 1979: 8; Gegechkori, 1981: 119; 1983b: 148; 1984: 104; Baeva, 1985: 206. Cyamophila odontopyx Loginova, 1978: 86. Holotype O’, U.S.S.R. [not examined]. Syn. n. DEscRIPTION. Coloration. General coloration of head and thorax orange-red with white markings, abdomen and terminalia somewhat paler; forewing pale yellow with diffuse apical reddish brown pattern and dark brown spots in the radular areas; antennal segments 1—8 yellow, segments 3-8 apically darkened, segments 9-10 dark brown. All the specimens to hand are immature and it is likely that fully mature specimens are darker. Structure. Head (Fig. 22) robust, genal cones short and broad, with broadly-rounded apices. Forewing (Fig. 21) with well-developed pterostigma and costal break; spinules confined to radular areas where they form distinct black marginal punctuations; cell m, elongate, cell cu,, high, vein Cu, strongly recurved adjacent to wing margin. Metatibia with 5 thick black spurs, basal metatarsus with 2 black spurs. Male proctiger (Fig. 23) simple; paramere (Fig. 24) parallel-sided with a large inner apical tooth; aedeagus (Fig. 25) with broad hooked apex. Female terminalia (Fig. 93) moderately long. Measurements. See Table 1. MATERIAL EXAMINED 14 0’, 6 Q, Israel and U.S.S.R. (IDH); 10’, 1 9, Iran (BMNH). Host PLANT. Glycyrrhiza glabra L. PUBLISHED DISTRIBUTION. Turkey (Klimaszewski & Lodos, 1977, 1979; Vondracek, 1953), Israel (Halperin etal., 1982), Afghanistan (Loginova, 1972b; Loginova & Baeva, 1972), Iran (Loginova, 1972b; Loginova & Baeva, 1972), Mongolia (Klimaszewski, 19685), U.S.S.R.: S. European part, Ukrainian SSR, Arme- nian SSR, Georgian SSR, Gruz SSR, Azerbaijan SSR, “Transcaucasus’, ‘Pamir’, Kazakh SSR, Uzbek SSR, Turkmen SSR, Tadjhik SSR (Gegechkori, 1984; Baeva, 1985). ComMENTs. Cyamophila odontopyx falls within the range of variation of C. glycyrrhizae. We are unable to separate material of both nominal species, determined by Loginova, and the two are here synonymized. C. glycyrrhizae is separated from other west Palaearctic members of the genus by the presence of a subapical band of infuscation in the forewing. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 9 Cyamophila medicaginis (Andrianova) (Figs 37-42) Psylla medicaginis Andrianova, 1952: 270; Loginova, 1968: 306; 1972b: 141; Gegechkori, 1968: 510; 1969b: 723; 1970: 714; 1976: 65; Baeva & Kankina, 1971: 84; Klimaszewski, 1963: 428; 1969a: 47; 1973: 214; Gegechkori & Djibladze, 1976: 33. Syntypes, 0’, 2, U.S.S.R. [not examined]. Cacopsylla medicaginis (Andrianova) Klimaszewski, 1975: 199. Cyamophila medicaginis (Andrianova) Loginova, 1977: 582; 1981: 26; Gegechkori, 1978: 21; 1981: 121; 1984: 148. DEscrIPTION. Coloration. Specimens to hand had the body, legs and antennae orange-yellow throughout. Slightly darker markings were present on dorsum of thorax. Structure. Forewing (Fig. 37) oblong oval, with spinules confined to radular areas; pterostigma and costal break well-developed. Male proctiger (Fig. 39) simple; paramere (Fig. 40) parallel-sided, expanded apically into forward pointing tooth; apex of aedeagus slightly reniform. Female terminalia as in Fig. 42. Measurements. See Table 1. MATERIAL EXAMINED 20,2 9,U.S.S.R.: Tabagatan, Semipal 6.vii.1962, (Loginova). Host PLANT. Medicago sativa L. and possibly Vicia sp. PUBLISHED DISTRIBUTION. Southern U.S.S.R., “Transcaucasus’ (Gegechkori, 1968; 1969a; 1970; 1976; 1978; 1981; 1983b; 1984; Gegechkori & Djibladze, 1976), Ukrainian SSR (Klimaszewski, 1963), Pamir (Baeva & Kankina, 1971), Kazakh SSR (Andrianova, 1952; Loginova, 1972b), Central Asia (Gegechkori, 1984). CoMMENTS. This species has yet to be recorded from outside the U.S.S.R. but Klimaszewski (1969, 1975) includes it in his key to the Polish fauna on the grounds that as it occurs in the west U.S.S.R. it is likely to occur in Poland. We have followed Klimaszewski. It can be separated from the other clearwing west Palaearctic Cyamophila species by the following combination of characters: the lack of forewing spinules throughout the membrane, the shape of cells cu,;, and m, and the shape of the male paramere and aedeagus. Cyamophila prohaskai (Preisner) (Figs 26-30, 94) Psylla prohaskai Preisner, 1927: 263; Prohaska, 1928: 4; Haupt, 1935: 234; Franz, 1943: 382; Schaefer, 1949a: 53; 1949b: 31; Wagner & Franz, 1961: 167; Kuwayama & Miyatake, 1971: 52; Klimaszewski, 1973: 219. Syntypes CO’ 9, AustTRIA [not examined]. Cyamophila prohaskai (Preisner) Loginova, 1977: 582; Burckhardt, 1983: 54; Conci & Tamanini, 1986: 60. DescriPTIon. Coloration. General body colour of mature specimens orange-red with white markings on dorsum of thorax; forewing membrane clear to pale yellow, veins yellow; antennal segments 1-8 yellow, segments 3-8 infuscate apically, segments 9-10 dark brown; legs orange-yellow. Structure. Head (Fig. 27) with relatively long genal cones. Forewing (Fig. 26) oblong-oval, with well-developed pterostigma and costal break; spinules present throughout all cells. Metatibia with 5 thick black apical spurs; basal metatarsus with 2 spurs. Male proctiger (Fig. 28) simple, paramere (Fig. 29) gradually broadening to a truncate apex, with an anteriorly directed apical tooth; aedeagus (Fig. 30) slender, with a slightly hooked apex. Female terminalia (Fig. 94) long and slender, with dorsal margin of proctiger sinuous; circumanal pore ring small. Measurements. See Table 1. MATERIAL EXAMINED 20,2 2, Switzerland (DB). Host PLANT. Anthyllis vulneraria L. PUBLISHED DISTRIBUTION. Austria (Preisner, 1927; Prohaska, 1928; Haupt, 1935; Franz, 1943; Wagner & Franz, 1961), Italy (Conci & Tamanini, 1986), Switzerland (Schaefer, 1949a, b; Burckhardt, 1983). ComMENTS. The record for China published by Kuwayama & Miyatake (1971) is unlikely to be correct: all other records are from the European Alps. C. prohaskai can be distinguished from the other west Palaearctic clearwing Cyamophila species by the presence of dense spinules on the forewing membrane. 10 I. D. HODKINSON & D. HOLLIS Cyamophila stoklosai Klimaszewski & Lodos (Figs 31-36) Cyamophila stoklosai Klimaszewski & Lodos, 1979: 9. Holotype ©’, Turkey (Lodos, Klimaszewski collection) [not examined]. Material of this species was unavailable and the following description is based entirely on information given in the original description. The drawings are derived from photomicrographs presented by Klimaszewski & Lodos (1979). DEscrIPTION. Head and body sandy yellow with rust-coloured patches and bands; abdominal sclerites brown, intersegmental membranes yellow; genitalia slightly paler brown than rest of abdomen; forewing transparent, slightly yellowish towards apex; antennae yellow except for segments 8—10 and the apices of segments 6—7 which are brown; legs yellow with brownish patches on femur. Structure. Head (Fig. 32) robust, genal cones short, with broadly rounded apices. Forewing (Fig. 31) oblong-oval, with well-developed pterostigma and costal break; spinules confined to the radular areas; cell my, narrow; cell cu,, high, strongly arched. Male proctiger (Fig. 33) simple; paramere (Fig. 35) with bulbous apex. Female terminalia as in Fig. 36. Measurements. See Table 1. Host PLANT. Astragalus sp. PUBLISHED DISTRIBUTION. Turkey (Klimaszewski & Lodos, 1979). Table 1 Measurements of Acizzia and Cyamophila species. Males and females are given separately and all values are in mm. Female Male Male Male Head Antennal Forewing proctiger proctiger paramere aedeagus width length length length length length length Acizzia bicolorata m 0-54 0-93 1-50 — 0-18 0-18 0-16 f a -_ = hee: ek = rae hollisi m 0-57-0-65 1:00-1:14 1-63—1-83 — 0-20-0-23 0-17-0-24 0-15-—0-19 f 0-61-0-65 1-05-1-22 1:92-2:12 0-55—0-88 — — — uncatoides m 0-59-0-63 0-79-0-96 1-67—1-80 — 0-28-0-29 0-21-0-22 0-20—0-21 f 0:58-0:61 0-83-0-88 1-80-1-91 0-35-—0-39 — — — acaciae- m 0-34-0-42 0-55-0-60 1-17-1-24 — 0-12—0-14 0-12-0-13 0-12-0-13 baileyanae f 0-37-0-44 0-60-0-64 1-30-1-61 0-30-0-31 — — — Cyamophila glycyrrhizae m 0-78—-0-80 0-90-0-96 2-20—2-31 — 0-33-0-35 0-29-0-31 0-28—0-29 f 0-79-0-81 1:00-1:10 2-30—2-32 0-70-0-72 — — — medicaginis m not available f prohaskai m 0-75—-0-78 1:07-1:10 2-42—2-44 — 0-31-0-33 0-26-0-27 0-27-—0-29 f 0-81-0-83 1-13-1-16 2-63-2-64 0-86—0-89 — — — stoklosai m 0-62 0-98 1-68 — 37 0-24 — f 0-66 1-10 1-98 0-66 — — ARYTAINA Forster Arytaina Forster, 1848: 69; Pflugfelder, 1941: 76. Type-species: Psylla spartii Hartig, 1841 (= Psylla genistae Latreille, 1804), designated by Oshanin, 1912: 128. Amblyrhina Low, 1879: 599; 1888b: 382; Kieffer, 1905: 165; Heslop-Harrison, 1951: 425; Loginova, 1976a: 598; Hodkinson & White, 1979a: 59. Type-species: Psylla torifrons Flor, 1861, by monotypy. Syn. n. Psyllopa Crawford, 1911: 628. Type-species: Psyllopa magna Crawford, by original designation. [Synony- mised by Crawford, 1914: 122.] W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 11 DiaGnosis. Head weakly deflexed, genal cones short and broad, with broadly rounded apices, in approximately same plane as vertex. Pronotum broad and flat, propleurites quadrate, divided by a vertical suture. Forewing oval to elongate-oval, usually broadest at or before middle; pterostigma and costal break absent; cell cu,, low and rarely slanting towards base of wing. Basal metatarsus with one black spur. Male proctiger without posterior lobe; paramere broad with truncate apex, terminated by a characteristic anteriorly directed tooth; aedeagus with a strongly hooked apex. Associated with legumes of the tribe Genisteae. COMMENTS. Arytfaina is a monophyletic genus of 11 species associated with host-plants of the legume tribe Genisteae within the Mediterranean Basin. It contains the original core species related to the type-species plus most species hitherto referred to Amblyrhina. The synonymy given relates only to references not quoted under the individual species synonymies and to papers that deal specifically with the generic synonymy. Unfortunately Arytaina had become a holding genus for many other species, outside the area under study, which are unrelated to the type-species and it is necessary to reallocate these species, where possible, to other genera (see Hodkinson, 1983; Hodkinson & White, 1981; Jensen, 1957b; Tuthill, 1943; Mathur, 1975; Capener, 1970). The following species are here transferred to other genera as follows. Acizzia obscura (Crawford, 1912; from Psyllopa). Comb. n. = A. spinosa (Mathur, 1975; from Arytaina). Syn. n., Comb. n. A. albizziae (Yang, 1984; from Arytaina). Comb. n. Arytainilla devia (Loginova, 1976; from Arytaina). Comb. n. (see p. 42). A. nubivaga (Loginova, 1976b; from Arytaina). Comb. n. (see p. 42). Ceanothia assimilis (Crawford, 1914; from Arytaina). Comb. n. C. bicolor (Jensen, 1957b; from Arytaina). Comb. n. C. boharti (Jensen, 1957b; from Arytaina). Comb. n. C. essigi (Jensen, 1957b; from Arytaina). Comb. n. C. insolita (Tuthill, 1943; from Arytaina). Comb. n. C. mitella (Jensen, 1957b; from Arytaina). Comb. n. Euphalerus isitis (Cotes, 1893; from Psylla). Comb. n. E. punctinervis (Crawford, 1919; from Arytaina). Comb. n. Euryconus pulchra (Crawford, 1919; from Arytaina). Comb. n. Insnesia brevigena (Crawford, 1919; from Arytaina). Comb. n. I. crawfordi nom. n. = I. pulchra (Crawford, 1920; from Arytaina). Comb. n. (homonym of Arytaina pulchra Crawford, 1919). I. flava (Crawford, 1919; from Arytaina). Comb. n. I. iolani (Crawford, 1919; from Arytaina). Comb. n. I. meridionalis (Crawford, 1919; from Arytaina). Comb. n. I. thakrei (Mathur, 1973; from Arytaina). Comb. n. I. tuberculata (Crawford, 1917; from Arytaina). Comb. n. I. uichancoi (Braza & Calilung, 1981; from Arytaina). Comb. n. I. variabilis (Crawford, 1917; from Arytaina). Comb. n. Peripsyllopsis ramakrishni (Crawford, 1912). Stat. rev., Comb. rev. Retroacizzia mopanei (Pettey, 1924; from Arytaina). comb. n. = R. antennata Heslop-Harrison, 1961b. Syn. n. Spanioneura turkiana (Klimaszewski & Lodos, 1977; from Amblyrhina). Comb. n. S. pechai (Klimaszewski & Lodos, 1977; from Amblyrhina). Comb. n. Arytaina virgina Caldwell, 1944 and A. cornicola Frauenfeld, 1896 are both regarded here as nomina dubia, while A. fasciata Laing, 1930 remains the sole species which cannot be assigned to another genus at present. Key to species 1 Forewing (Fig. 43) with cell m,, elongate, as long as stem of vein M. Cell cu,, narrowly mraneitiar, veins Cuj,and Ci, almost straight ..................ccsescssesetevees longicella sp. n. (p. 16) — Forewing (Figs 45, 51, 57) with cell m,,. much shorter than length of vein M. Cell cu;, not narrowly triangular, veins Cu,, and Cu, more distinctly curved ...................cceeeeeeee eee ee ees 2 YQ 10 11 I. D. HODKINSON & D. HOLLIS Cells of forewing with distinct longitudinal brown or black pattern extending backwards from the apex(Mies 45, 470A), ccpissigec hie ner que Saar Petecniet cerceer oud. « aecjsoients: pero ae Ree Sole ies aie Cells of forewing without distinct longitudinal patterning (in atlasiensis, adenocarpi and angustatipennis (Figs 53, 61, 51) there is occasionally a faint clouding of the wing membrane and in the former this is accompanied by the presence of diffuse spots in the apical cells) ....... Forewing (Fig. 45) with costal margin comparatively straight, surface spinules present through- out cell r,. Genal cones (Fig. 46) broad with rounded apex. Male paramere (Fig. 66) comparatively short and broad. Aedeagus as in Fig. 67 .................006 genistae (Latreille) (p. Forewing (Figs 47, 49) with costal margin more strongly curved; surface spinules absent throughout most of cell r,. Genal cones either broad with an angular apex (Fig. 48) or more slender with rounded apex (Fig. 50). Paramere relatively longer and narrower (Fig. 69, 72). PREMSA GUS AS UT PUSS IU, PO nda es an oe te neta ce cant ae eo en Ect opi Oe oe ace ae eae ee Forewing (Fig. 49) with surface spinules absent from cell c+sc and the base of r,, and confined to narrow bands in the centres of cells r, and m. Genal cones (Fig. 50) slender, with rounded apex. Male paramere and aedeagus as in Figs 71—73. Spain and Sicily maculata (L6w) (Spanish material) (p. Forewing (Fig. 47) with spinules present throughout cells c+sc and r, and broadly distributed throughout cells r, and m. Genal cones (Fig. 48) broader and with a more angular apex. Male paramere and aedeagus as in Figs 68—70. Eastern Europe maculata (Low) (Hungarian material) (p. Forewing (Fig. 51) strongly narrowed to an acute apex. Female proctiger (Fig. 101) long and slender, with a relatively small circumanal pore ring ............ angustatipennis (Loginova) (p. Forewing (Figs 55, 59, 61) less strongly narrowed, apex more broadly rounded. Female proctiger (Figs 92, 99, 100) less slender, relatively shorter or more robust, with a relatively longer circumanal POre TING ................sa- peemedh eters ee Piece enaent ye oe Forewing shaped as in Fig. 53 with cell c+sc somewhat bowed outwards, vein Cu, strongly curved before its junction with the anal margin; nebulous brown spots present on membrane around the apex. (N.B. Several other species have dark spots where the longitudinal veins meet the mareiial yeti). is c..% sacc cape -ton Gene saceaen eee: nea ee ae atlasiensis sp. n. (p. Forewing (Figs 55, 59, 61) of different shape, without nebulous brown spots on membrane around apex (if pattern present as in A. torifrons (Fig. 57) then never consisting of spots); vein Cu, less strongly curved before its junction with the marginal vein .....................0.008- Forewing (Figs 55, 57) opaque amber or yellow-brown throughout or with suffused brown markings along vesaroundapex «. 2... Zagih as! cnt aeense Shenae oft 2k. Bent eRe Forewing (Figs 59, 61) not amber or yellow-brown, and without brown patterning along vein Margins and APOUMEAPER 038) mada ened Mig. hese WemeGn Wei < comet sae eo ee ee Forewing (Fig. 55) elongate oval, amber or yellow-brown throughout. Genal cones (Fig. 56) short with a broad base and a very broadly rounded apex. Male paramere (Fig. 74) less broad. Apex of aedeagus (Fig. 81) more strongly hooked.....................065 putonii (Low) (p. Forewing (Fig. 57) more broadly oval, shorter, not amber or yellow-brown throughout; pattern somewhat variable but usually consisting of suffused brown markings along vein margins and around apex. Genal cones (Fig. 58) slightly longer and less broadly rounded at apex. Male paramere (Fig. 83) broader. Apex of aedeagus (Fig. 80) less strongly hooked torifrons (Flor) (p. Forewing (Fig. 59) broadest in basal third, with a narrowly rounded apex; vein Cu,, weakly arched. Male paramere (Fig. 83) with a subapical lobe on the anterior margin. Aedeagus (Fig. 84) witha weakly curved tip’ 42..5s00 5 eter, eter ee onederene hispanica sp. n. (p. Forewing (Figs 61, 257) broadest towards the centre, with a more broadly rounded apex and with vein Cu,, more strongly arched. Male paramere (Figs 74, 86, 89) without a subapical lobe on the anterior margin. Aedeagus (Figs 87, 90) usually with a strongly recurved tip ....... Male paramere (Fig. 74) almost as long as proctiger, with at most a short anteriorly directed apical tooth. Genal cones (Fig. 258) apparently much shorter. (Figures and description taken from the original, all type material is lost.) Libya ................... africana Heslop-Harrison (p. Male paramere (Figs 86, 89) distinctly shorter than the length of the proctiger, with a large apical tooth. Genal cones (Figs 62, 64) longer, with broadly rounded apices. France, Portal and Spann oid crs «xe sess coeceete cla ac cies a ect ea shih «dt a Forewing (Fig. 63) with dense spinules broadly distributed throughout all cells. Head (Fig. 64) more robust, vertex relatively long. Male paramere (Fig. 89) with a distinct anterior bulge at mid length and with a sinuously truncated apex which is developed anteriorly into a massive 14) 14) 11 W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 13 tooth. Male aedeagus as in Fig. 90. Antennae more than twice as long as head width magnidentata sp. n. (p. 17) — Forewing (Fig. 61) with less dense spinules which are confined to the apical areas of cells r,, r., M, M42, Cj, the base of c+sc and around vein Cuz. Head (Fig. 62) less robust, vertex relatively shorter. Male paramere (Fig. 86) more regularly shaped, curved posteriorly and with a smoothly convex apex which is developed anteriorly into a large but shorter tooth. Male aedeagus as in Fig. 87. Antennae less than twice head width .......... adenocarpi Low (p. 13) Arytaina adenocarpi Low (Figs 61, 62, 85-87, 99) Arytaina adenocarpi Low, 1880: 552; 1883: 230; 1885: 150; Puton, 1886: 92; Aulman 1913: 32; Fuente, 1920: 321; Heslop-Harrison, 1951: 434; Ramirez Gomez, 1956: 75; Klimaszewski, 1973: 191; Loginova, 1976a: 598; Conci & Tamanini, 19845: 262. Lectotype CO’, FRANCE, here designated (NM). Arytaena adenocarpi Low; Oshanin, 1907: 366; 1912: 128. DEscriPTION. Coloration. All the available specimens are immature. General coloration of head, thorax and abdomen and appendages bright green, occasionally orange-yellow; often with orange markings on dorsum of head, thorax and abdomen; forewing clear, occasionally with faint brownish clouds; veins yellow to brown, always much darker at the points where they meet the marginal vein. Structure. Head (Fig. 62) with genal cones swollen, with rounded apices. Forewing (Fig. 61) elongate oval; spinules sparse and confined to wing apex, the base of cell c+ sc and around vein Cu; pterostigma and costal break absent; cell m, of normal size, vein Cu;, strongly curved. Metatibia with 5 thick black spurs; metatarsus with 1 black spur. Male proctiger (Fig. 85) simple; paramere (Fig. 86) parallel-sided, curved posteriorly in apical half, with a large inner apical tooth projecting anteriorly; aedeagus (Fig. 87) with moderately strongly hooked apex. Female terminalia as in Fig. 99. Measurements. See Table 2. MATERIAL EXAMINED Lectotype CO’, France: ‘Gallia, Landes. Type v. Aryt. adenocarpi Low’ (NM). Paralectotype 2, same data as lectotype. 7 0’, 6 9, Portugal and France: ‘Haute Pyrenees’ (BMNH). In addition there is an abundance of material in the BMNH, presumably collected by Heslop-Harrison in Spain or Portugal, which does not bear data labels. Host PLANTS. Adenocarpus complicatus (L.) Gay and Adenocarpus hispanicus (Lam.) D.C. in Lam. & Dc. PUBLISHED DISTRIBUTION. France (Low, 1880; Aulmann, 1913), Spain (Fuente, 1920; Ramirez Gomez, 1956), Italy (Conci & Tamanini, 1984). COMMENTS. This species is closest to A. magnidentata. It can be separated by the form of the forewing spinule pattern and the shape of the male paramere and aedeagus. Arytaina africana Heslop-Harrison (Figs 74, 257, 258) Arytaina africana Heslop-Harrison, 1951: 438; Loginova, 1972a: 16; 1976a; 598. Syntypes C’ and 9, Lipya [not traced]. DESCRIPTION. Coloration. According to Heslop-Harrison this is a ‘greenish or yellowish-brown insect with darker brown markings on the head and thorax, and brown maculations on the forewings; strongly resembling A. genistae in general appearance’. Structure. Head (Fig. 258) with short genal cones. Forewing oval, cells m, and m2 of moderate size, vein Cu, strongly arched; spinules present throughout all cells; pterostigma and costal break absent. Male proctiger (Fig. 74) simple; paramere (Fig. 74) long, parallel-sided, with a truncate apex. Shape of aedeagus and of female terminalia unknown. Measurements. See Table 2. Host PLANT. ‘A yellow flowered broom’ (Heslop-Harrison, 1951). PUBLISHED DISTRIBUTION. Libya (Heslop-Harrison, 1951) and possibly Morocco, although the specimens identified as this species by Loginova (1972) do not fit the original description very well. 14 I. D. HODKINSON & D. HOLLIS ComMMENTs. All type material of this species appears to be lost and we have seen no other authentic material. Our account is based entirely on Heslop-Harrison’s original description. A. africana appears closest to A. genistae but the male paramere is more slender and the genal cones are less massive. Arytaina angustatipennis (Loginova) comb. n. (Figs 51, 52, 81, 101) Amblyrhina angustatipennis Loginova, 1972: 24; Loginova, 1976a: 598. Holotype 0’, Morocco (ZMU) [examined]. DEscrIPTION. Coloration. Dorsal surface of head and thorax brownish yellow with longitudinal reddish brown and orange markings; genal cones dirty yellow; abdominal sclerites reddish brown, darker above, intersegmental membranes yellow; genitalia yellowish brown; forewing membrane clear to very pale yellow; veins yellow basally, becoming dark brown apically; radular spinules prominent, forming dark brown spots at the wing margin; antennae yellow basally, darkening towards the apex; legs brownish ellow. ; Structure. Head (Fig. 52) with vertex very short but broad; genal cones short and very broadly rounded. Forewing (Fig. 51) broadest in basal third, tapered to a narrow acute apex; spinules confined to the radular areas; cells m, and cu, of normal size; vein Cu,, strongly curved; pterostigma and costal break absent. Metatibia with 5 thick black apical spurs; basal metatarsus with 2 spurs. Male proctiger (Fig. 81) simple; paramere (Fig. 81) parallel-sided with a truncate apex. Female terminalia (Fig. 101) long and slender. Measurements. See Table 2. MATERIAL EXAMINED Holotype 0’, Morocco: Atlas Mountains, Reraia, 5—15.vi.1926 (Lindberg) (ZMU). Paratypes, 1 0’, 1 Q, same data as holotype. Host PLANT. Unknown. PUBLISHED DISTRIBUTION. Morocco (Loginova, 1972). CoMMENT. This species is separated from other members of the genus by the apically tapered forewing. Arytaina atlasiensis sp. n. (Figs 53, 54, 91) DEscriPTION. Coloration. Dorsal surface of head and thorax dirty yellow, with orange or reddish brown markings; genal cones yellow. Abdominal sclerites dark brown to black; intersegmental membranes yellow; genitalia dark brown to black; forewing membrane clear to pale yellow, veins concolorous; nebulous small brownish spots present towards wing apex; antennal segments 1—4 yellow; remainder dark brown; legs orange-yellow. Structure. Head (Fig. 54) robust, genal cones short but massive, with broadly rounded apices. Forewing (Fig. 53) broadly oval, with cell c+sc bulging outwards; costal break and pterostigma absent; vein R, reaching wing margin at or about the centre of the wing; cell m, narrow; cell cu,, more strongly arched than normal; vein Cu,, strongly recurved as it approaches wing margin; spinules confined to cells in basal half of wing. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 spur. Male unknown. Female terminalia (Fig. 91) long and robust. Measurements. See Table 2. Holotype 2, Morocco: Middle Atlas, 18.v.1961 (Lawrence) (BMNH). Paratypes. 3 2, same data as holotype. Host PLANT. Unknown. ComMENTSs. This species appears to be closest to A. genistae and A. maculata but can be distinguished by the characteristic shape and pattern of the forewing. Arytaina genistae (Latreille) (Figs 45, 46, 65-67, 96) Psylla genistae Latreille, 1804: 382; Guérin, 1843: 370; Burmeister, 1843: 139; Bolivar & Chicote, 1879: 183; Reiber & Puton, 1880: 75. Syntypes 0’, 2, FRANCE [not traced]. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 15 Psylla ulicis Curtis, 1835: 565 (22a); Low, 1883: 252. Syntypes [incomplete data], GREAT BRITAIN (UMM). [Synonymised by Low, 1877: 126.] Psylla spartii Hartig, 1841: 375; Forster, 1848: 69; Lethierry, 1874: 86; Low, 1877: 126; 1883: 251; Flor, 1861: 358; Szulcezewski, 1927: 199. Type [incomplete data], GERMANY; Berlin. [Synonymised by Low, 1877: 126.] Arytaina spartii (Hartig) Forster, 1848: 69; Low, 1877: 126; Meyer-Dur, 1871: 404. Psylla genistae Fabricius; Lethierry, 1869: 365. [Fabricius reference not traced. | Arytaena ulicis (Curtis) Scott, 1876: 529. Chermes (Ataenia) genistae (Latreille) Thomson, 1877: 828. Chermes genistae (Latreille) Douglas, 1878: 41. Arytaena genistae (Latreille) Low, 1879: 597; Scott, 1880: 132; 1882a: 14; 1882b: 255; Reuter, 1881: 162; Edwards, 1896: 250; Oshanin, 1907: 366; 1912: 128; Heslop-Harrison, J. W., 1915: 401; Van der Goot, 1912: 284; Britten, 1930: 75; Wahlgren, 1934: 89; Haupt, 1935: 241; Murray, 1936: 138; Schaefer, 1949a: 54; 1949b: 31; Vondraéek, 1951b: 128; Smreczynski, 1954: 141; Glowacka & Harisanov, 1983: 64. Arytaina genistae (Latreille) Low, 1883: 239; 1884: 149; 1888a: 19; Puton, 1886: 92; Hueber, 1904: 275; Heslop-Harrison, 1937: 2; 1951: 428; Lauterer, 1971: 197; Loginova, 1976a: 598. (All other references listed under distribution refer to Arytaina genistae unless listed above.) Psyllopa magna Crawford, 1911: 628. Syntypes 0’, 9, U.S.A. [Synonymised by Crawford, 1914: 126.] Arylaina genistae (Latreille) Halbert, 1934: 310. [Mis-spelling. ] DEscrIPTION. Coloration. Immature specimens with body coloration bright green often with orange or brown markings on the dorsum. Mature specimens with dorsal surface of head and thorax yellowish white with extensive longitudinal brown or orange-brown markings; genal cones yellowish white; abdominal sclerites dark brown to black; intersegmental membranes yellow; genitalia dark brown; fore- wing membrane clear with brown or sometimes greyish longitudinal pattern; veins brown; antennal segments 1—5S brownish yellow, remainder dark brown; legs dirty yellow, usually with femora darkened above. Structure. Head (Fig. 46) moderately robust, genal cones short and moderately broadly rounded. Forewing (Fig. 45) oval, pterostigma and costal break absent; spinules present in all cells; cells m, and cu, of average size for genus; vein Cu, strongly curved. Metatarsus with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 65) simple; paramere (Fig. 66) relatively broad, parallel-sided with a truncate apex which is developed into an anteriorly directed tooth; aedeagus (Fig. 67) with strongly hooked apex. Female terminalia as in Fig. 96. Measurements. See Table 2. MATERIAL EXAMINED The material is too extensive to list. It includes all the material in the BMNH, together with material in the collections of IDH and DB. Host pLants. Occurs primarily on Cytisus scoparius (L.) Link but is also recorded in the literature from Chamaecytisus austriacus (L.) Link, Chamaecytisus heuffelii (Wierzb.) Rothm. and Genista tinctoria L. These published records require confirmation as they may reflect the confusion between this species and A. maculata. PUBLISHED DISTRIBUTION. Austria (Léw, 1888a), Bulgaria (Glowacka & Harisanov, 1983), Czechoslovakia (Low, 1888a; Duda, 1892, Sulc, 1905a; Vondraéek, 1957; Lauterer, 1977), Denmark (Jacobsen, 1919), France (Lethierry, 1869; Reiber & Puton, 1880: Carpentier & Dubois, 1892: Dominique, 1902; Lambertie, 1910), Great Britain (Scott, 1880; Edwards, 1896; Britten, 1930; Heslop-Harrison, J. W., 1915; Heslop- Harrison, 1936a,b,c; Murray, 1936; Ing, 1966; 1971; 1974; Watmough, 1968a,b; Hodkinson, 1976; 1978; Hodkinson & White, 1979b; White & Hodkinson, 1982), German D.R. (Lauterer, 1966; Emmrich, 1976; 1978), German F.R. (Férster, 1848), Ireland (Halbert, 1935), Italy (Ferrari, 1888), Netherlands (Van der Goot, 1912; Blote, 1926), Poland (Szulczewski, 1927; Smerczynski, 1954; Klimaszewski, 1961; 1967; 1969a; 1971; 1973; 1975; 1979; Zgardinska, 1976; Glowacka, 1979), Spain (Bolivar & Chicote, 1879; Vondraéek, 1951b), Sweden (Reuter, 1881; Wahlgren, 1934; Ossiannilsson, 1952; 1971), Switzerland (Schafer, 1949a,b; Burckhardt, 1983), Rumania (Dobreanu & Manolache, 1962), U.S.A. (introduced) (Crawford, 1914; Van Duzee, 1917; Tuthill, 1943; Vondraéek, 1953). ComMENntTs. Early records of this species from Hungary have been shown to relate to A. maculata Low (Horvath, 1918). The two species are easily confused and the records given above should be treated with care, particularly those for eastern Europe and Italy where their ranges appear to overlap. The two species can be distinguished by the shape of the forewing and the form of the male genitalia. 16 I. D. HODKINSON & D. HOLLIS Arytaina hispanica sp. n. (Figs 59, 60, 82-84, 92) DESCRIPTION. Coloration. Dorsal surface of head and thorax brownish yellow with darker brown longi- tudinal markings; abdominal sclerites dark brown above, paler beneath; intersegmental membranes dull yellow; genitalia yellow-brown to brown; forewing membrane very pale yellowish orange; veins con- colorous; antennal segments 1—2 yellow; segments 3—4 yellow basally, infuscate apically; segments 5—10 brown; legs brownish yellow. Structure. Head (Fig. 60) with genal cones shorter than vertex and broadly rounded at apex. Forewing (Fig. 59) semi-opaque, broadest towards base, tapering to narrowly rounded apex; veins thick; spinules dense and present in all cells; cell m, of average size; cell cu, small with vein Cu,, weakly curved. Metatibia with 5 thick black apical spurs; metatarsus with 1 black spur. Male proctiger (Fig. 82) simple; paramere (Fig. 83) with distinct subapical bulge on anterior margin; aedeagus (Fig. 84) weakly hooked at apex. Female terminalia as in Fig. 92. Measurements. See Table 2. Holotype CO’, Spain: Sierra Nevada, Loma del Mulhacen, 3200 m, 30.viii.1975 (Barfuss) (ETH). Paratypes. 6 2, same data as holotype. Host PLANT. Undetermined ‘Genista’ species. CoMMENTS. This species has the short, oval wing as found in A. torifrons but in other respects resembles species such as A. adenocarpi. The form of the male genitalia is characteristic. Arytaina longicella sp. n. (Figs 43, 44, 95) DESCRIPTION. Coloration. The type specimen is probably immature. General colour of body and append- ages dull dirty yellow throughout (possibly greenish in life). Forewing membrane very pale yellow, slightly Opaque; veins dull yellow. Structure. Head (Fig. 44) with genal cones shorter than vertex, very broad and truncate at the apex. Forewing (Fig. 43) oval with narrowly rounded apex; lacking spinules except in radular areas and around vein Cup; cell m, very long and narrow; cell cu, triangular, vein Cu, straight; pterostigma and costal break absent. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Female terminalia as in Fig. 95. Male unknown. Measurements. See Table 2. Holotype 9, Spain: Sierra de Guadarrama, vii.1926 (Uvarov) (BMNH). Host PLANT. Unknown. CoMMENTS. This species is based on a single female but the wing venation is so different it should not be confused with other species. Arytaina maculata (L6w) comb. n. (Figs 47—S0, 68-73, 97) Amblyrhina maculata Low, 1886: 157; 1888b: 382; Horvath, 1886: 313; 1918: 58; Puton, 1886: 92; Ragusa, 1907: 237; Oshanin, 1907: 366; 1912: 128; Aulmann, 1913: 32; Haupt, 1935: 241; Vondraéek, 1951b: 128; 1957: 180; Klimaszewski, 1969b: 44; 1970: 422; 1973: 189; 1975: 129; Loginova, 1964: 464; 1976a: 598; Lauterer, 1977: 98; Dobreanu & Manolache, 1962: 147; Andrianova & Klimaszewski, 1983: 38; Glowacka & Harisanov, 1983: 64. Syntypes 0’, @ HunGary (not located). Ambly[r]rhina maculata Low; Low, 1888a: 19. [Arytaina genistae (Latreille), partim; Low, 1888a: 19; Horvath, 1886: 314; 1918: 58. Misidentifications. ] Amblyrhina maculosa Low; Klimaszewski, 1965. [Mis-spelling. ] DEscRIPTION. Coloration. Immature specimens with body colour bright green throughout. Mature specimens with genal cones, vertex and dorsal part of thorax yellowish white with light brown and orange markings; underside of head and thorax yellow with dark brown markings; abdominal sclerites dark brown to black, intersegmental membranes orange-yellow; genitalia orange-yellow to dark brown; forewing membrane clear with brown or sometimes greyish longitudinal pattern; veins brown; antennal segments 1—S yellow, 5—10 dark brown; femora dark brown above, remainder of legs dirty yellow. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 17 Structure. Head (Figs 48, 50) with genal cones shorter than vertex, broad; more slender in the Spanish specimens. Forewing (Figs 47, 49) oval, without pterostigma or costal break; with spinules present throughout all cells except for r;, where they are at most confined to the apex; in the Spanish material they are also absent from cell c+sc; cells m, and cu,, of average size, vein Cu, strongly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 68) simple; paramere (Fig. 69) parallel-sided, truncate, with apical forward-pointing tooth; aedeagus (Fig. 70) strongly hooked. In Spanish males the paramere is more slender (Figs 71—73). Female terminalia as in Fig. 97. Measurements. See Table 2. MATERIAL EXAMINED The only material in the Léw collection comes from a different locality to that stated in the original description. 1 C’, 1 9, Hungary: Szomotor (det. Low) (NM). 4 0’, 8 Q Italy and Spain (BMNH, PL). Host PLants. Chamaecytisus ratisbonensis (Schaeffer) Rothm. and Chamaecytisus borysthenicus (Gruner) A. Klaskova. PUBLISHED DISTRIBUTION. Bulgaria (Klimaszewski, 1965, 1970; Glowacka & Harisanov, 1983), Czechoslo- vakia (Vondraéek, 1957; Lauterer, 1977), Hungary (Low, 1886, 1888a; Horvath, 1886, 1918a; Haupt, 1935), Italy, Sicily (Ragusa, 1907), Rumania (Dobreanu & Manolache, 1962), U.S.S.R.; central and southern European part (Loginova, 1964; Andrianova & Klimaszewski, 1983), Yugoslavia (Horvath, 1918b). ComMENTS. This species is very close to A. genistae with which it has been confused in the past. It is possible that some of the published records for A. genistae refer to this species. The specimens from Sicily have slightly more slender genal cones and parameres than Low’s material from Austria. Similarly the Spanish material listed is included under maculata primarily because of the similarity of the wing pattern. In several other respects such as the shape of the genae and parameres it is close to adenocarpi. This whole group warrants further detailed investigation. The maculata complex is similar in several respects to A. genistae but can be distinguished by the shape of the forewing and the form of the male genitalia. Arytaina magnidentafa sp. n. (Figs 63, 64, 88-90) DESCRIPTION. Coloration. Unknown; the type-material is slide mounted. Forewing clear, without pattern on membrane but veins darkened at the point where they meet the marginal vein. Structure. Head (Fig. 64) robust, genal cones large and broadly rounded at apex, slightly shorter than length of vertex. Forewing (Fig. 63) oval, without pterostigma or costal break; dense spinules present throughout all cells; cells m, and cu, of average size for genus; vein Cu,, strongly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 88) simple; paramere (Fig. 89) with a massive apical tooth; aedeagus (Fig. 90) with a very strongly hooked apex. Female unknown. Measurements. See Table 2. Holotype CO’, Portugal: Rio Mandego, vii.1953 (Heslop-Harrison) (BMNH). Host pLAnt. Unknown. ComMENTs. Although based on a single male this species is quite distinct from other Arytfaina species. It is closest to the other clear elongate-oval winged species, A. adenocarpi, but differs in characters of the forewing and the shape of the male paramere and aedeagus. Arytaina putonii (L6w) comb. n. (Figs 55, 56, 75-77, 100) Amblyrhina putonii Low, 1888b: 381; Aulmann, 1913: 32. Syntypes 0’, 2, FRANCE (missing from Léw collection). Amblyrhina putoni Low; Puton, 1899: 113; Oshanin, 1907: 365; 1912: 128; Lambertie, 1910: 96; Vondraéek, 19516: 128; Klimaszewski, 1973: 190; Loginova, 1976a: 598. Description. Coloration. General body coloration bright green with orange markings on the dorsum of the head, thorax and abdomen; forewing membrane amber coloured, slightly darker towards apex, veins concolorous; antennal segments 1—2 green; segments 3—6 yellowish brown, becoming infuscate towards the apex; segments 7—10 brown; legs greenish yellow. 18 I. D. HODKINSON & D. HOLLIS Structure. Head (Fig. 56) short and broad; genal cones very broad basally and appearing to curve round to give broadly truncate apices. Forewing (Fig. 55) short and oval, of somewhat rugose texture, narrowly rounded at apex; spinules absent except from radular areas; cells m, and cu, of average size, vein Cuj, weakly curved to straight; costal break and pterostigma absent. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 75) simple; paramere (Fig. 76) parallel-sided with large apical anteriorly directed tooth; aedeagus (Fig. 77) with hooked apex. Female terminalia as in Fig. 100. Measurements. See Table 2. MATERIAL EXAMINED 10 0,7 Q, Portugal and Spain (BMNH). Host PLANT. Recorded by Léw (1888) from ‘Cytisus spinosus’ which, in all probability, is Calicotome spinosa (L.) Link. PUBLISHED DISTRIBUTION. France (Léw, 1888b; Lambertie, 1910; Aulmann, 1913; Vondraéek, 19515). CoMmMENTS. This species is closest to A. torifrons but differs in the shape and coloration of the forewing, the shape of the genal cones and the form of the male aedeagus and paramere. Table 2 Measurements of Arytaina species. Males and females are given separately and all values are in mm. Female Male Male Male Head Antennal Forewing proctiger proctiger paramere aedeagus width length length length length length length adenocarpi m 0-77—0-83 1:38-1:59 1-91-2-19 — 0-39-0-42 0-31-0-32 0-28—-0-30 f 0-80-0-89 1-55-1-65 2-14-2-45 0-76—0-85 — — — africana m ,. 0-78 — 2-38 — f (data from original description) angustatipennis m — — DoT — 0-25? 0-20? — f 0-92-0-98 1-65-1-70 2-70-—2-82 0-62—0-80 — — — atlasiensis m unknown f 1:00-1:02 1-56-1:58 2:54-2:56 0-89-0-97 — — — genistae m 0:90-0:96 2-05-—2-20 2-35-—2-75 — 0-39-0-44 0-37-0-39 0-30-0-32 f 0-95-1:05 1-90-2:25 2-40-3-00 0-89-0-99 — — — hispanica m 0-85 — 2-01 — 0-38 0-31 0-31 f 0-86-0-90 1:27-1:29 2-32—2-34 0-85—0-86 — — — longicella m unknown f 0-83 — 2:34 0-96 — — — maculata m 0-86 1-68 2°39 — 0-39 0-33 0-33 f 0-86 1-46 2-35 0-91 _- — — magnidentata m 0-79 1-61 2°12 — 0-37 0-33 0-27 f unknown putonii m 0-81—0-86 1:10-1:14 1-90—2-03 — 0-36-0-38 0-31-0-32 0-30-0-31 f 0-86-0-93 1-15-1-16 2:09-2:20 0-72—0-80 — — _- torifrons m 0-73-0-75 0-85-0-95 1-55-1-59 — 0-34-0-36 0-25-0-27 0-26—0-28 f 0-70-0-83 0-85-1-00 1-68-—2-02 0-64—-0-71 a — — Arytaina torifrons (Flor) comb. n. (Figs 57, 58, 78-80, 102) Psylla torifrons Flor, 1861: 360. LECTOTYPE ©’, France (NM), here designated [examined]. [ Psylla spartiophila Forster sensu Puton, 1871: 437. Misidentification. ] Amblyrhina torifrons (Flor) Low, 1879: 600; 1882b: 259; 1883: 252; 1888b: 382; Puton, 1886: 92; Oshanin, 1907: 365; 1912: 128; Aulmann, 1913: 32; Klimaszewski, 1973: 190; Loginova, 1976a: 598; Hodkinson & White, 1979a: 59. DESCRIPTION. Coloration. Immature specimens pale greenish yellow throughout. Mature specimens with dorsal surface of head and thorax reddish orange; genal cones yellow; abdomen yellowish brown to dark W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 19 brown; genitalia brown; forewing clear to pale yellowish, veins pale brown to dark brown, pigment spreading from the veins onto immediately adjacent areas of the membrane, apical pattern consisting of _ diffuse brown clouds of irregular form; antennal segments 1—6 brownish yellow; segments 7—10 brown; legs brownish yellow. Structure. Head (Fig. 58) relatively broad, with genal cones convergent in front of frons. Forewing (Fig. _ 57) short and broad, oval; without pterostigma or costal break; spinules confined to radular areas; cells m, and cu,, of normal shape; vein Cu,, weakly curved. Metatibia with 5 thick black apical spurs; basal _ metatarsus with 1 black spur. Male proctiger (Fig. 78) simple; paramere (Fig. 79) relatively broad and - appearing swollen along anterior margin, with large apical anteriorly directed tooth; aedeagus (Fig. 80) _ with apex strongly recurved. Female terminalia as in Fig. 102. Measurements. See Table 2. _ MATERIAL EXAMINED Lectotype ©’, France: ‘Gallia, Marseilles, type v. Aryt. torifrons Flor.’ (NM). Paralectotype Q, same data as lectotype. 78 O’, 2, France and Spain (BMNH, PL, IDH). | Host PLANT. Genista hispanica L. PUBLISHED DISTRIBUTION. France (Flor, 1861; Aulmann, 1913; Hodkinson & White, 1979a). _ Comments. This species is closest to A. putonii but differs in the shape and coloration of the forewing, the _ shape of the genal cones and the form of the male paramere and aedeagus. LIVILLA Curtis | Livilla Curtis, 1836: 625; Low, 1863: 106; Puton, 1876: 288; Kieffer, 1905: 163; Oshanin, 1907: 366; 1912: 128; Pflugfelder, 1941: 75; Heslop-Harrison, 1951: 421; Loginova, 1976a: 599. Type-species: Livilla ulicis Curtis, 1836, by monotypy. _ Floria Low, 1879: 590; Kieffer, 1905: 164; Heslop-Harrison, 1951: 421; Oshanin, 1907: 367; 1912: 128; _ Pflugfelder, 1941: 77; Heslop-Harrison, 1951: 421; Loginova, 1976a: 598; Hodkinson & White, 1979a: 55. Type-species: Psylla pyrenaea Mink, 1859, designated by Oshanin, 1912: 128 (not by Haupt, 1935: 241 as indicated by Hodkinson & White, 1979a: 55). Syn. n. _ Alloeoneura Low, 1879: 594; Oshanin, 1907: 368; 1912: 128; Pfleugfelder, 1941: 77; Heslop-Harrison, 1951: 421; Loginova, 1976a: 599. Type-species: Arytaina radiata Forster, 1848, by monotypy. Syn. n. Allaeoneura Low; Puton, 1886: 93; Kieffer, 1905: 163. [Mis-spelling. | Floria (Floriella) Ramirez Gomez, 1956: 87. Type-species:Psylla pyrenaea Mink, 1859, by monotypy. [Objective synonym of Floria.| Syn. n. | | Diacnosis. Head weakly deflexed, in same plane as thorax, eyes hemispherical, antennae usually slender, at least twice as long as head width; genal cones usually elongate and slender, little deflexed from the plane of the vertex. Pronotum broad and flat; propleurites quadrate, divided by a vertical suture. Forewing oval _to oblong-oval, usually membraneous but short and coriaceous in a few species. Costal break present or | absent; pterostigma, if present, short; cell cu, not tall and usually not strongly arched. Basal metatarsus usually with 1 black spur, occasionally with 2 or 0. Male proctiger simple, without posterior process; paramere long and slender, tapered to a hooked apex; aedeagus with apex generally rounded, not strongly hooked as in Arytaina. Head and thorax often covered in abundant long setae. Associated with legumes of _the tribe Genisteae. Comments. As defined above the genus Livilla probably constitutes a monophyletic group of 31 species living on legume hosts of the Genisteae, primarily within the Mediterranean Basin. The single South | African species annosa, which Heslop-Harrison (1961b) designated type-species of the subgenus Brinc- kitia, is excluded from the study. The synonymy given above relates only to references not quoted under the individual species synonymies and to papers which deal specifically with generic synonymy. The genus now contains species previously included under the old generic names Livilla, Floria and Alloeoneura plus one species from Amblyrhina. We could find no characters that warrant the recognition of Floria and Alloeoneura. The general body form and genitalia of all the species is very similar and they differ from one | another primarily in the form of the forewing. We regard the development of short, coriaceous forewings in some species merely as a specific adaptation for ensheathing the body to conserve water in a dry environment rather than as a generic synapomorphy. Based on characteristics of the forewing, six loosely defined groups of species can be recognised: the ulicis-, horvathi-, burckhardti-, radiata- and spartiisuga- groups, with a residual assemblage of heterogeneous species. 20 I. D. HODKINSON & D. HOLLIS Key to species 1 10 11 Forewing (Figs 103, 105, 107) coriaceous and strongly convex, oval, little more than twice as Josh, aS DOA CUMICIS-BL OUND) oo eqns easiest cts capnatte e ga cn eS lam hea ar ge Z Forewing (Figs 109, 115, 125, 139, 153) membraneous, not strongly convex; usually longer and more oblong-oval in shape; if short and/or oval (L. burckhardti (Fig. 165) and L. radiata (Fig. 125)) them with apical OrOwn Paste DMI ooo oe ance on te ee ee 4 Genal cones (Fig. 104) massive, longer than the vertex. Forewing (Fig. 103) dark brown throughout; cell 14> elongate, longer than the stem of vein M,; wing membrane with short distinct furrows arising from and at right angles to the veins. General body colour shining dark browimnte DIAC NTOUSMOWE 23.5. o0e ncn ntti pem ales «> nus on soa doch oad ulicis Curtis (p. 23) Genal cones (Fig. 106, 108) shorter. Forewings (Figs 105, 107) yellowish to amber or brown, with a whitish basal patch; cell m4, shorter than stem of vein M; membrane without distinct furrows. Body either reddish brown with paler markings or dark brown with 2 large white streaks which extend onto the base ofthe wimgsi 7i2.5 22... Skn.0.. ioe eee eee 2 ce a Forewing (Fig. 105) brown with broad white basal patch, costal margin weakly curved in basal third, cell cu,, upright, surface spinules absent from most cells. Genal cones (Fig. 106) with very broadly rounded apices. Male genitalia as in Figs 170-172. Iberian Peninsula bivittata sp. n. (p. 24) Forewing (Fig. 107) yellow to amber throughout, without white basal patch, costal margin strongly curved in basal third, cell c+sc appearing to bulge outwards, cell cu,, leaning towards base of wing, surface spinules obvious in all cells. Genal cones (Fig. 108) acutely tapered. Male genitalia as in Figs 173-175. Austria, Switzerland, N. Italy ..... vicina (L6w) (p. 24) Forewing (Fig. 109) broadest in basal third, acutely tapering towards apex. Genal cones (Fig. 110) very short, extending little beyond the vertex (burckhardti-group) ...._ cognata (L6w) (p. 25) Forewing (Figs 115, 125, 149, 165) broadest at or beyond the middle, not acutely tapering towards apex. Genal cones (Figs 116, 126, 150, 166) much longer ................ 0... ce ceeeeeeeeee ees 5 Forewing (Fig. 165) with subapical brown chevron pattern, wing apex narrowly rounded. Female terminalia (Fig. 270) long and slender. Genal cones shorter than the length of the vettex(barekhardtl-proup) .223....2 7c SA SR burckhardti sp. n. (p. 25) Forewing pattern, if present (Figs 121, 144, 157), not consisting of brown chevrons, wing apex more broadly rounded. Female terminalia (Figs 287—292) more robust. Genal cones (Figs 116,122,426, 132) usually atleastas long.as vertex)! ......020. 0.0. IR Se Forewing (Fig. 153) clear, without colour pattern or suffused apex genistae Ramirez Gomez (p. 38) Forewing (Figs 131, 144, 161) either with a colour pattern or infuscate apically ...................... 7 Forewing (Fig. 125) short and broad, at most 2-1 times as long as broad, with colour pattern as shown in Fig. 125; cell r; short and broad. Genal cones (Fig. 126) very long and slender CR AGTALA-S OUD) eos ncaa, non dda agiaaane ee artes once ee tania AE radiata (Forster) (p. 29) Forewing (Figs 117, 121, 123, 127, 161) longer and narrower, more than 2-1 times as long as broad (except some male L. poggii (Fig. 142)); pattern not as in Fig. 125; cell r; longer and narrower. Genal cones (Figs 118, 122, 124) usually shorter, if long (L. spectabilis (Fig. 162)) PCM WIOTS BOWS Eco) 5 ocr soiree ccs cent eae an see et 8 Forewing pattern (Figs 111, 113, 115, 117, 155) consisting of longitudinal brown streaks (horvathi-group) g Forewing pattern consisting of brown clouds (Fig. 119), small roundish maculations (Fig. 131), infuscation of the wing apex and/or the area adjacent to the anal margin (Figs 159, 163) ora combination of thesé characters (Figs 138, 139) 2222.) Se, Se eee ee 13 Forewing pattern (Fig. 111) consisting of narrow longitudinal streaks immediately adjacent to the veins in the apical half of the wing. Female circumanal pore ring (Fig. 287) small, at most 0-19 timesienstlot proctiger: a3 eee es Oe EL. nervosa sp. n. (p. 27) Forewing pattern (Figs 113-115, 117, 155) consisting of an anterior streak which may or may not be confined to the apex of vein R,, and a posterior streak extending along the anal margin. Female circumanal pore ring (Figs 284, 288, 289, 290) at least 0-21 times length of PROCHIPER vou... den Sk GtSk REN RE BAe RR SU eee 10 Forewing (Figs 113, 115) with anterior streak confined to the apex of vein R,, not extending into the basal half of the wing iv.4). 0). sek tenes. Aw. Wee CE eee eee 11 Forewing (Figs 117, 155) with anterior streak extending across vein R, into the basal half of WHI ore oS Seale on ox hg EARS es «se RRS teil cade Sele L en © JURE Lune Ree See 12 Forewing (Fig. 113) with anterior streak consisting of a small spot around the apex of vein R,; 12 W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 21 costal margin of the wing weakly curved so that apex of cell r, is abruptly truncated by vein R,. Genal cones (Fig. 114) strongly divergent, outer margins strongly concave, apex broadly rounded. Male proctiger (Fig. 182) somewhat lobed posteriorly. Male aedeagus as in Fig. ee hte shee ee, Berge eT ES hn vid cis «

meeting the marginal vein well above the wing apex (radiata-group) .................. 14 Forewing (Figs 131, 132, 135, 157) with the branches of vein M not strongly divergent, vein Mj..>meeting the marginal vein at or below the wing apexX ....................5.ccceecedeeceeeeeee eens 16 Forewing pattern (Fig. 123) consisting of small round brown maculae which become confluent and form an irregular brown pattern in the apical and posteroapical regions of the wing; costal margin strongly curved throughout. Dorsum of thorax uniformly shining chocolate brown. Genal cones (Fig. 124) conical. On Genista fasselata ...............60005 syriaca (L6w) (p. 31) Forewing (Figs 119, 121) without small brown maculae, pattern consisting primarily of brown clouds or bold markings in the apical third of the wing, occasionally with slight infuscation in the basal half; costal margin almost straight between the points where veins R, and R, meet the marginal vein. Dorsum of thorax not uniformly shining chocolate brown. Genal cones re mmoncclongatc. On Other hosts. ....4.;...0026..-0ccas asecndes devens sess eg bences nee Wena 15 Veins in apical third of forewing (Fig. 121) dark brown to black, pattern very dark brown; occasionally with the basal half of the wing weakly infuscate. Genal cones (Fig. 122) divergent. Basal metatarsus with one thick black spur. Male paramere (Fig. 195) with a strongly serrated anterior margin and a massive apical tooth. Male aedeagus (Fig. 196) with thin shaft and relatively large apex. Host plant unknown ....................655 Jautereri sp. n. (p. 30) Veins in the apical third of forewing (Fig. 119) not dark brown to black, pattern more widespread but less intense, mid brown; basal part of wing not infuscate. Genal cones (Fig. 120) not divergent. Basal metatarsus without thick black spurs. Male paramere (Fig. 192) without strongly serrated anterior margin and with a small apical tooth. Male aedeagus (Fig. 193) with a broader shaft and a relatively small apex. On Retama sp. ....... retamae (Puton) (p. 30) Forewing (Fig. 161) long and narrow, of a characteristic shape, pattern consisting of an irregular brown or orange-brown infuscation of the membrane, primarily in the apical third and a darker brown spot around vein Cu;,. Genal cones (Fig. 162) very long, at least 1-3 times length of vertex. Antennae very long, at least 2-85 times head width. On Spartium SS se or spectabilis (Flor) (p. 37) Forewing (Figs 131, 151, 157, 161) relatively shorter and broader, pattern not as above. Genal cones (Figs 132, 152, 158, 164) less than 1-3 times length of vertex. Antennae relatively Semen siess than 2-8) times head width. Omother hosts ..............0.cc.ccceccecceccccceucceccsceces 17 Forewing (Fig. 157) with distinct chocolate brown colour pattern distributed around the apex of vein R,, the apical half and branches of vein M and around veins Cu,, and Cuy, MIME ROI Jr, COTE Tell a wale se aia dtr cul dde coexgvedse dha cd ails vetds pseudoretamae sp. n. (p. 35) Forewing (Figs 131, 144, 145, 159) without distinct chocolate brown patterning in apical third; if similarly shaped pattern to pseudoretamae present then much paler, more diffuse and MEM WHLHSI ALL TOUNG MACUIATIONS (1.0. oe .e. cc. le lies ea ck cece se ecubdencbodsetvescsecudecsces 18 Forewing (Fig. 159) slightly rugose, small maculae absent, yellow-brown to orange-brown infuscation occupying a continuous band extending around the apex and along the posterior wing margin; remainder of membrane lighter; pigment darker along vein margins giving a somewhat furrowed appearance. Male proctiger (Fig. 239) with a slight basal posterior lobe adusta (Low) (p. 39) Forewing (Figs 131, 139) usually with small maculae present and broadly distributed through- 22 19 20 21 22 2 24 25 26 ya | I. D. HODKINSON & D. HOLLIS out the wing, often overlying the areas of infuscation when present. If maculae apparently absent (in some specimens of L. variegata and L. maura they are nebulous and indistinct) then infuscation distributed as in Fig. 169, or forewing membrane uniformly yellow to orange (Fig. 151). Male proctiger (Figs 203, 206, 209, 245) without basal posterior lobe ................. 19 Forewing (Figs 127, 131, 135, 137, 142) with small brown maculae distributed throughout (spartiisuga-g1oup) i. 2..02..6 350s, 4.1 Ak ADO hts AL ge Se 20 Forewing (Figs 151, 163) with maculae absent or very indistinct; when present very nebulous and confined tothe apical half of wing tt a. See. ce AR ee 30 Sexually dimorphic species. Male forewing (Fig. 142) with a pale maculation-free band running transversely across the centre. Female forewing (Fig. 141) maculate throughout; vein M very strongly arched in both sexes, cell cu, very short and high. Apex of male aedeagus as in Fig. 226. On Genisia corsica ip Sarina Jo%o,2) 4 see dd ne 8 poggii (Conci & Tamanini) (p. 36) Forewing (Figs 129, 131, 133, 137) without transverse pale band, occasionally with a pale longitudinal band; vein M less strongly arched and cell cu, flatter and more elongate. Apex of male aedeagus (Figs 217, 220, 223) of different shape. On other host plants .................... Zt Ground coloration of forewing membrane (Figs 127, 129, 131, 133), excluding maculae, uniform, clear to pale yellow throughout of. -)04.0)...0. 22th does ode eee 22 Ground coloration of forewing membrane (Figs 135, 144, 145, 147, 149), excluding maculae, two-tone, clear to pale yellow along anterior half of wing, pale brown to orange-brown along posterior half and/or around the apex 4 2.422, 20.4221... 09.0. Gat eee ee Se 27 Maculae absent from a broad band running along anterior margin of forewing (Fig. 133); wing with a somewhat truncate apex. Male and female genitalia as in Figs 212—214, 274. On | Calicotome spinosa and Genistg pilosa) 02212, <32.0. dbs: ahd take sob e dA pyrenaea (Mink) (p. 34) Maculae present in area along anterior margin of forewing (Figs 129, 131, 137); wing shaped differently, with a less truncate apex. Male and female genitalia as in Figs 206—208, 209-211, 221-223, 271-272, Omother hosts 0... 0 Bois. oka ek ih ee ea 23 Maculae absent from a longitudinal band running the length of cell r, (Fig. 137). Male paramere (Fig. 219) long and curved posteriorly in apical half. Apex of aedeagus (Fig. 220) smoothly GOMOD oo a ccccanuen snictincmanes tag nas tous reaching to just below wing apex; pterostigma and costal break absent. Metatibia with 5 thick black apical spurs; metatarsus with 1 black spur. Male proctiger (Fig. 170) simple; paramere (Fig. 171) without stout basal posterior spines; aedeagus (Fig. 172) with slightly hooked apex. Female terminalia as in Fig. 264. Measurements. See Table 3. Holotype ©’, Portugal: Villa Formosana, 1954 (Heslop-Harrison) (BMNH). Paratypes. 1 0’, 1 9 same data as holotype, 3 0’, 2 2 from unknown locality, but intermixed with specimens from Spain and Portugal (Heslop-Harrison) (BMNH). In addition there is a lot of material of this species without locality labels and in poor condition in the Heslop-Harrison collection (BMNH) which is not included in the type series. Host PLANT. Unknown. ComMENTS. This species is closest to Livilla ulicis, but can be distinguished from it by the shorter genal cones and the shape of cells m, and cu, of the forewing. Livilla vicina (L6w) comb. n. (Figs 107, 108, 173-175, 265) Floria vicina Low, 1886: 159; 1888a: 20, Puton, 1886: 92; Hueber, 1904: 276; Oshanin, 1907: 367; 1912: 128; Aulmann, 1913: 35; Prohaska, 1928: 5; Haupt, 1935: 242; Klimaszewski, 1973: 195; Loginova, 1976a: 599; Burckhardt, 1983: 55. Syntypes 0’, 2, AustTRIA [missing from Léw collection, NM]. Arytaena montana Cerutti, 1939a: 448; 1939b: 583; Schaefer, 1949a: 55; 1949b: 31. Lectotype CO, SWITZERLAND, designated by Burckhardt, 1983 (MZL). [Synonymised by Burckhardt, 1983: 55.] Arytaina montana (Cerutti) Heslop-Harrison, 1951: 435; Tamanini, 1955: 11; 1977: 109; Klimaszewski, 1973: 191, Loginova, 1976a: 598. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 25 DEscrIPTION. Coloration. Genal cones, vertex and thoracic dorsum dirty orange-yellow, with indistinct darker orange markings on thorax; abdominal sclerites brown, intersegmental membranes yellowish orange; genitalia dirty yellow to brown; forewing coriaceous, membrane yellow to orange, veins concolor- ous; antenna orange-yellow basally becoming darker towards apex; legs orange-yellow. Structure. Head (Fig. 108) with genal cones slightly shorter than vertex, proximate, with rounded apices. Forewing (Fig. 107) short and broad, oblong-oval; cell c+sc bulging outwards; costal break and ptero- stigma absent; spinules present throughout all cells; vein Rs sinuous, not strongly curved at apex; cell m4» narrow; vein M,,> reaching margin below wing apex; cell cu, large and leaning towards base of wing; vein Cu, weakly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 173) simple; paramere (Fig. 174) slender, without thickened setae on basal posterior margin; aedeagus (Fig. 175) with strongly recurved apex. Female terminalia as in Fig. 265. Measurements. See Table 3. MATERIAL EXAMINED 90’, 11 9, Switzerland and Italy (BMNH, DB, IDH, MM). Host PLANT. Genista radiata (L.) Scop. PUBLISHED DISTRIBUTION. Austria (LOw, 1886, 1888a; Prohaska, 1928; Haupt, 1935), Italy (Tamanini, 1955, 1977), Switzerland (Cerutti, 1939a,b; Schaefer, 1949a,b; Burckhardt, 1983). ComMENTs. It is a measure of the confusion that has existed within the Arytainini that this species has belonged simultaneously to two separate genera. L. vicina resembles L. ulicis and L. bivittata in possessing short oval coriaceous forewings, but it can be distinguished from both these species by the shape and detailed venation of the forewing and by the shape of the head and genal cones. The burckhardti-group The two members of this group have the forewing apex narrowly rounded, and very short genal cones. Livilla burckhardti sp. n. (Figs 165, 166, 248-250, 270) DEscriPTION. Coloration. Specimens in the type series appear to have been taken while developing their full adult coloration. General body colour (preserved in alcohol) dirty yellow, possibly green in life; sclerites of abdomen beginning to turn brown; forewing clear, with brown colour pattern; veins yellow; antennae and legs concolorous with body. Structure. Head (Fig. 166) with short, broad genal cones. Forewing (Fig. 165) short and oval; pterostigma and costal break absent; spinules present throughout all cells; vein Rs strongly curved towards marginal vein; vein M evenly curved; vein M,,, reaching to just below wing apex; vein Cu,, strongly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 248) simple; paramere (Fig. 249) relatively short and broad; aedeagus as in Fig. 250. Female terminalia (Fig. 270) long and slender with dorsal margin of proctiger shallowly convex. Measurements. See Table 3. Holotype CO’, Spain: Pyrenees, Ordesa National Park, 160 m, on spiny Genista (Lienhard) (ETH). Paratypes. 1 0’, 2 9, same data as holotype. Host PLanT. The type series was collected from a spiny Genista species. ComMENTSs. L. burckhardti appears to be closest to L. cognata but the resemblance may be superficial. It can be separated from all other members of the genus by the chevron patterning in the forewing. Livilla cognata (LOw) comb. n. (Figs 109, 110, 176-178, 266) Amblyrhina cognata Low, 1882b: 258; Oshanin, 1907: 365; 1912: 128; Puton, 1886: 92; Haupt, 1935: 241; Vondraéek, 1951b: 128; 1957: 178; Wagner & Franz, 1961: 162; Lauterer, 1965: 174; 1977: 98; Loginova, 1976a: 598; Klimaszewski, 1970: 420; 1973: 189; Gegechkori, 1984: 147; Tamanini, 1977: 107; Glowacka & Harisanov, 1983: 64. LECTOTYPE ©’, Austria, here designated (NM) [examined]. Ambly[r]rhina cognata Low; Low 1888a: 19. [Mis-spelling. ] DEscriPTION. Coloration. General colour of head, thorax and abdomen bright green, often with orange 26 I. D. HODKINSON & D. HOLLIS markings on dorsum of thorax; forewing membrane clear, occasionally with faint brown clouds in apical half of wing; veins pale, weakly pigmented; antennae green, apical segments infuscate, legs green. Specimens to hand are probably immature and it is likely this species darkens as it becomes older. Structure. Head (Fig. 110) short and broad, with small broadly rounded genal cones. Forewing (Fig. 109) oval, with a narrowly rounded apex; costal break and pterostigma absent; spinules confined to apices of cells; vein R, not strongly curved to wing margin; vein M, evenly curved; vein M,,, reaching to just below wing apex; cell cu, flat; vein Cu,, weakly curved. Metatibia with 5 thick black spurs at apex; basal metatarsus with 1 black spur. Male proctiger (Fig. 176) simple; paramere (Fig. 177) slender, without thickened basal posterior spines; aedeagus (Fig. 178) with a weakly angled apex. Female terminalia as in Fig. 266. Measurements. See Table 3. MATERIAL EXAMINED Lectotype CO’, Austria: ‘Austria inf., Modling, det Low’ (NM). Paralectotype 2, same data as lectotype. 2 0’, 2 2 Austria (BMNH). Host PLANTS. Chaemaecytisus ratisbonensis (Schaeffer) Rothm., Chamaecytisus supinus (L.) Link, Genista germanica L. PUBLISHED DISTRIBUTION. Austria (Léw, 1882b, 1888a; Vondraéek, 1951b; Wagner & Franz, 1961), Bulgaria (Klimaszewski, 1970; Glowacka & Harisanov, 1983), Czechoslovakia (Lauterer, 1965, 1977; Vondraéek, 1957), France (Aulmann, 1913), ‘Germania’ (Haupt, 1935), Italy (Tamanini, 1977), U.S.S.R.: Caucasus (Gegechkori, 1983). CoMMENTS. This species has been separated from those with which it had previously been placed within the genus Amblyrhina. The characters of the genitalia suggest that it belongs in Livilla, even though the genal cones are much shorter than is usual for the genus. It is distinguished further from other Livilla species by the narrowly rounded apex of the forewing. The horvathi-group Members of this group are recognised by the longitudinal brown banding patterns on the forewing membrane. Livilla horvathi (Scott) comb. n. (Figs 115, 116, 185-187, 289) Floria horvathi Scott, 1879: 84; Low, 1883: 240; 1888a: 20; Horvath, 1886: 314; 1918a: 58; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Klimaszewski, 1964: 64; 1967: 14; 1968b: 781; 1969a: 46; 1973: 193; 1975: 130; Loginova, 1964: 464; 1976a: 599; Vondratek, 1957: 192; Lauterer, 1977: 98; Klimaszewski & Lodos, 1979: 7; Glowacka & Harisanov, 1983: 64; Onucar, 1983: 56. Holotype Q, Huneary [not traced]. [Floria vittipennella Reuter sensu Vondraéek, 1951b: 128; Smrezynski, 1954: 141. Misidentifications. ] DEscriPTION. Coloration. Body colour of immature specimens green throughout. Mature specimens with vertex and thoracic dorsum dirty greenish yellow with brown markings; genal cones dirty yellow to pale brown; abdominal sclerites dark brown, intersegmental membranes greenish yellow; genitalia yellow- brown to brown; forewing membrane clear, veins dirty yellow to pale brown; colour pattern orange- brown; antennal segments 1-4 dirty yellow, 2—4 apically darkened, remainder brown; legs brownish yellow, femora occasionally brown above. Structure. Head (Fig. 116) with long divergent genal cones with narrowly rounded apices. Forewing (Fig. 115) elongate oval, without pterostigma or costal break; fine spinules distributed throughout all cells; vein Rs moderately strongly curved towards margin; vein M slightly sinuous; vein M,,, reaching to wing apex; cell cu,, moderately long but not high; vein Cu,, evenly arched. Metatibia with 5 thick black spurs at apex; basal metatarsus with 1 black spur. Male proctiger (Fig. 185) simple; paramere (Fig. 186) without stout setae on basal posterior margin; aedeagus (Fig. 187) with curved bulbous apex. Female terminalia as in Fig. 289. Measurements. See Table 3. MATERIAL EXAMINED 1’, 1 9, ‘Scott collection’ [no further data] (BMNH). 16 o’, 16 9, Greece, Bulgaria and Czechoslo- vakia (DB, MM, IDH). HostT PLANT. Chamaecytisus austriacus (L.) Link. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 27 PUBLISHED DISTRIBUTION. Bulgaria (Klimaszewski, 1967; Glowacka & Harisanov, 1983), Czechoslovakia (Vondracek, 1957; Lauterer, 1977), Greece (Vondraéek, 1951b), Hungary (Low, 1888a; Horvath, 1886, 1918a), Poland (Klimasezewski, 1964, 1967, 19685, 1969a, 1975; Smrezynski, 1954), Rumania (Dobreanu & Manolache, 1962), Turkey (Klimasezewski & Lodos, 1979; Onucar, 1983), European U.S.S.R. (Loginova, 1964). ComMENTS. This appears to be the sister species of L. nigralineata. The two species differ from each other in the distribution of brown markings on the forewing and in the shape of the genal cones and male genitalia, particularly the aedeagus. Livilla nigralineata sp. n. (Figs 113, 114, 182-184, 288) DEscrIPTION. Coloration. The type series comprises specimens which probably have not developed their full adult coloration. General colour pale green throughout with orange markings on the dorsum of the head, thorax and abdomen. Forewing membrane clear, wing pattern reddish brown. Antennae with segments 6—10 dark brown. Structure. Head (Fig. 114) with genal cones widely divergent, longer than vertex, with evenly rounded apices. Forewing (Fig. 113) elongate oval, almost parallel-sided; costal break and pterostigma absent; spinules fine, present in all cells but restricted in the cells along the anterior edge of wing; cell cu, moderately long; vein Cu,, moderately strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 thick black spur. Male proctiger (Fig. 182) simple; paramere (Fig. 183) relatively stout, parallel-sided; aedeagus (Fig. 184) with slightly hooked apex. Female terminalia as in Fig. 288. Measurements. See Table 3. Holotype o’, Algeria: Mtns de Belezma, 1500 m, 5.viii. 1980 (Remane) (BMNH). Paratypes. 1 0’, 2 2, same data as holotype. Host PLANT. Unknown. CoMMENTS. This appears to be the sister species to horvathi. It is separated from the latter by the smaller patch of brown markings around the apex of vein Rs and by the more divergent genal cones. Livilla nervosa sp. n. (Figs 111, 112, 179-181, 287) Description. Coloration. Mature specimens with head, thorax and abdomen pale dirty yellow; dorsum of thorax and vertex with pale orange or pale brown markings; abdomen yellowish or green, turning brownish in older specimens; genitalia yellow to brownish yellow; forewing membrane clear to pale yellow, veins yellow; patterning pale brown; antennal segments 1—2 yellow, segments 3 to 5 darkened apically, segments 6 to 10 dark brown; legs yellow. Structure. Head (Fig. 112) with slender divergent genal cones, slightly longer than vertex. Forewing (Fig. 111) elongate oblong-oval, almost parallel-sided; costal break and pterostigma absent; fine spinules present throughout all cells. Vein Rs strongly curved towards wing margin; vein M long and evenly curved; vein M,,> meeting margin well above wing apex; cell cu,, moderately long. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 179) simple; paramere (Fig. 180) curved posteriorly, with thickened setae present on basal posterior margin; aedeagus (Fig. 181) with angular bulbous apex. Female terminalia (Fig. 287) with very short circumanal pore ring. Measurements. See Table 3. Holotype ©’, Spain: Cordoba, Embalse de Iznajar, nr Iznajar, 12.vi.1977, (Hollis) (BMNH). Paratypes. Spain: 8 0’, 15 9, same data as holotype; 1 9, 1 nymph, Andalusia, Porto de los Alazores, 800 m, 30. vili.1975 (Lienhard) (DB). Host PLANT. Genista radiata (L.) Scop. ComMENTS. The upturned vein M,,, of the forewing also resembles that found in members of the radiata-group; otherwise L. nervosa resembles members of the horvathi-group. It is distinguished by the characteristic forewing pattern. 28 I. D. HODKINSON & D. HOLLIS Livilla vittipennella (Reuter) comb. rev. (Figs 117, 118, 188-190, 290) Psylla vittipennella Reuter, 1875: 333. Syntypes O’, @ YUGOSLAvIA [not located]. Floria vittipennella (Reuter) Low, 1879: 593; 1888a: 20; Puton, 1886: 93; Hueber, 1904: 276; Oshanin, 1907: 367; 1912: 128; Prohaska, 1928: 5; Haupt, 1935: 242; Burckhardt, 1983: 54. Livilla vittipennella (Reuter) Aulmann, 1913: 34. Floria lineata Cerutti, 1939a: 445; 1939b: 583; Schaefer, 1949a: 55; 1949b: 31; Klimaszewski, 1973: 193; Loginova, 1976a: 599; Tamanini, 1977: 111. Lectotype 0’, SwiTzERLAND (MZL) designated by Burc- khardt, 1983. [synonymised by Burckhardt, 1983: 54.] Floria vitipennella (Reuter); Klimaszewski, 1973: 195; Loginova, 1976a: 599. DEscrIPTION. Coloration. Mature specimens with dorsum of thorax and vertex brownish yellow with orange markings; genal cones yellow; remainder of body orange-yellow; forewing membrane clear, veins yellow; colour pattern reddish brown; antennal segments 1 and 2 yellow-orange, remainder pale brownish becoming darker towards antennal apex; legs orange-yellow. In some less mature specimens the body coloration often has a greenish tinge. Structure. Head (Fig. 118) with genal cones long and slender and with narrowly rounded apices. Forewing (Fig. 117) oblong-oval with a somewhat truncate apex; costal break and rudimentary ptero- stigma usually absent; fine spinules present throughout all cells; vein Rs strongly curved towards the margin; vein M evenly curved; vein M4, reaching to wing apex; cell cu,, shortish but tall; vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 188) simple; paramere (Fig. 189) with inner tooth somewhat set back from apex and without thickened setae on basal posterior margin; aedeagus (Fig. 190) with strongly recurved apex. Female terminalia as in Fig. 290. Measurements. See Table 3. MATERIAL EXAMINED 10 0’, 11 9, Switzerland and Italy (IDH, BMNH, DB). Host PLANT. Genista radiata (L.) Scop. PUBLISHED DISTRIBUTION Austria (L6w, 1888a; Prohaska, 1929; Haupt, 1935), Italy (Tamanini, 1977), Switzerland (Cerutti, 1939a,b; Schaefer, 1949a,b; Burckhardt, 1983), Yugoslavia (Reuter, 1875). CoMMENTS. L. vittipennella is distinguished from other members of the genus by the extensive brown band extending along the fore margin of the wing. Livilla klapperichi sp. n. (Figs 155, 156, 230—232, 284) DESCRIPTION. Coloration. Genal cones, vertex and dorsum of thorax brick red with white markings; underside of head and thorax orange-yellow; abdomen brownish yellow; genitalia yellow; forewing membrane clear with reddish brown pattern; veins yellow to brown; antennal segments 1-8 dirty yellow, segments 4—8 apically infuscate; segments 9-10 dark brown; legs orange-yellow. Structure. Head (Fig. 156) with genal cones somewhat divergent, about as long as vertex, with narrowly rounded apices. Forewing (Fig. 155) oblong-oval, broadest at about two-thirds the length; costal break and rudimentary pterostigma present; fine spinules present throughout all cells; cell m, small; vein M,4, reaching to wing apex; cell cu,, relatively long, with vein Cu,, relatively weakly curved; vein M evenly curved; vein Rs moderately strongly curved to wing margin. Metatibia with 4 thick black apical spurs; basal metatarsus with 2 black spurs. Male proctiger (Fig. 230) simple; paramere (Fig. 231) moderately stout, without basal posterior thickened setae; aedeagus (Fig. 232) with large bulbous apex. Female terminalia (Fig. 284) shorter and more wedge-shaped than in most other species. Measurements. See Table 3. Holotype CO (on two slides), Afghanistan: Kamdesch, Bashguital, Nuristan, 2200 m, 17.vii.1952 (Klapperich) (BMNH). Paratypes. 1 oO’, 1 9 and one specimen lacking genitalia, same data as holotype. One female is on the same slide as the holotype, with the wing on a separate slide labelled paratype. Host PLANT. Unknown. —————— W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 29 ComMMENTS. The name klapperichi is a manuscript name used by Vondracéek. This species, which differs from other representatives of the genus in having 2 basal metatarsal spines and only 4 spines at the apex of the tibia. It is somewhat atypical and represents the easternmost record for the genus Livilla. The radiata-group Members of this group can be recognised by the strongly upturned apex of vein Rs and the vein M, > which reaches the wing margin well above the wing apex. Livilla radiata (Forster) comb. n. (Figs 98, 125, 126, 200-202) Arytaina radiata Forster, 1848: 70. Holotype 2, AustRIA [not examined]. Psylla lactea Costa, 1863: 47; Low, 1877: 125; 1883: 241; Puton, 1876: 284. Type [sex not stated], ITALy [not examined]. [Synonymised by Léw, 1877: 125.] Psyllodes cytisi Becker, 1867: 113; Léw, 1879: 595; 1883: 236. Type [incomplete data], U.S.S.R. [not examined]. [Synonymized by Low, 1879: 595. | Psylla radiata (Forster) Low, 1877: 125. Alloeoneura radiata (Forster) Low, 1879: 595; 1882a: 168; 1883: 248; 1884: 150; 1888a: 20; Horvath, 1886: 315; 1918: 59; Duda, 1892: 38; Aulmann, 1913: 35; Oshanin, 1912: 128; Sulc, 1905a: 4; Haupt, 1935: 241; Prohaska, 1928a: 5; Klimaszewski, 1964: 65; 1965: 203; 1967: 13; 1969a: 46; 1973: 189; 1975: 132; Loginova, 1964: 464; 1966: 135; 1976a: 599; Lauterer, 1963: 148; 1977: 98; Vondracek, 19516: 127; 1957: 189; Wagner & Franz, 1961: 162; Dobreanu & Manolache, 1962: 137; Andrianova & Klimaszewski, 1983: 38; Glowacka & Harisanov, 1983: 64. Allaeoneura radiata (Forster); Puton, 1886: 93; Hueber, 1904: 276. Oshanin, 1907: 368. [Mis-spelling. ] Aleuroneura radiata (Forster); Klimaszewski, 1961: 79. [Mis-spelling. ] DEscriPTIon. Coloration. Mature specimens with head and thorax yellowish brown. Abdomen brown with yellow intersegmental membranes; genitalia yellow-brown; forewing membrane translucent white, veins yellow, markings chocolate brown; antennae yellow, apices of segments 3—8 brown, segments 9 and 10 completely brown; legs yellowish brown; apices of tibiae and tarsi dark brown. Structure. Head (Fig. 126) with genal cones long and slender, much longer than the vertex. Forewing (Fig. 125) oblong-oval, very short and broad; costal break and pterostigma usually absent; fine spinules present throughout all cells; vein Rs short, strongly curved to wing margin; vein M short, slightly sinuous; vein M,,> meeting wing margin well above the apex; cell cu;, short and relatively high, vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 200) simple; paramere (Fig. 201) without thickened setae along basal posterior margin; aedeagus (Fig. 202) with slightly reniform apex. Female terminalia as in Fig. 98. Measurements. See Table 3. MATERIAL EXAMINED 20,3 Q, Austria, West Germany and Czechoslovakia (BMNH, DB). Host PLANTS. Chamaecytisus ratisbonensis (Schaeffer) Rothm., Chamaecytisus austriacus (L.) Link, Chamaecytisus borysthenicus (Gruner) A. Klaskova, Lemboptropis nigricans (L.) Griseb. PUBLISHED DISTRIBUTION. Austria (Forster, 1848; Low, 1863, 1888a; Prohaska, 1928a; Haupt, 1935; Wagner & Franz, 1961), Bulgaria (Klimaszewski, 1965; Glowacka & Harisanov, 1983), Czechoslovakia (Léw, 1888a; Duda, 1892; Sulc, 1905a; Haupt, 1935; Vondraéek, 1957; Lauterer, 1977), Hungary (Low, 1888a; Horvath, 1886, 1918), Italy (Aulmann, 1913), Poland (Klimaszewski, 1961, 1964, 1967, 1969, 1975), Rumania (Dobreanu & Manolache, 1962), U.S.S.R., southern European part (Loginova, 1964, 1966; Andrianova & Klimaszewski, 1983), Yugoslavia (Low, 1888a; Aulmann, 1913), ‘S. Germany’ (Haupt, 1935). COMMENTS. This species has been recognised as the type-species of the monotypic genus A//oeoneura Low, 1879 for over 100 years. There is, however, little reason why it should be regarded as a distinct genus. L. radiata is distinguished by the short broad wing, with the characteristic pattern, and the very long genal cones. 30 I. D. HODKINSON & D. HOLLIS Livilla retamae (Puton) comb. n. (Figs 119, 120, 191-193, 291) Psylla retamae Puton, 1878: 134; Bolivar & Chicote, 1879: 184. Syntypes 6 CO’, 12 9, 1 ?sex, Spain (MM, MNHN) [examined]. Floria retamae (Puton) Low, 1883: 248; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Ramirez Gomez, 1956: 85; Loginova, 1976a: 599; Klimaszewski, 1973: 194; Samy, 1973: 452; Halperin et al. , 1982: 34. Floria (Brinckitia) retamae (Puton); Loginova, 1971: 629; 1972a: 27. Alloeoneura retamae (Puton) Vondracek, 19515: 127. DEscrIPTION. Coloration. Immature specimens greenish throughout, developing pale orange markings on dorsum of thorax. Mature specimens with genae, vertex and dorsum of thorax dirty yellow, with orange markings on thorax and vertex; ventral parts of head and thorax pale orange-yellow; dorsal abdominal sclerites brown, intersegmental membranes yellow; underside of abdomen greenish to brown; genitalia dirty yellow to brown; forewing membrane clear, veins clear to pale yellow; pattern brown; antennae dirty yellow, segments 3—6 darkened apically, segments 6—10 dark brown; legs dirty yellow. Structure. Head (Fig. 120) with genal cones slender, longer than the vertex, with narrowly rounded apices. Forewing (Fig. 119) elongate oblong-oval; costal break and rudimentary pterostigma present; spinules absent or thinly scattered in areas of brown patterning; vein Rs curved towards wing margin; vein M sinuous; vein M,,, meeting wing margin well before apex; cell cu,, moderately large, vein Cu,, strongly arched. Metatibia with 5 thick black spurs, basal metatarsus without black spurs. Male proctiger (Fig. 191) simple; paramere (Fig. 192) slightly expanded subapically, inner tooth partially hidden in side view; aedeagus (Fig. 193) with slightly hooked apex. Female terminalia as in Fig. 291. Measurements. See Table 3. MATERIAL EXAMINED 16 CO, 329, Israel, Egypt, Algeria, Morocco, Portugal and Spain (IDH, ZI, ZMU, BMNH, MM, MNHN, DB, PL). Host pLants. Retama sphaerocarpa (L.) Boiss, R. monosperma (L.) Boiss and R. raetam (Forskal) Webb & Berth. PUBLISHED DISTRIBUTION. Algeria (Aulmann, 1913), Egypt (Samy, 1973), Israel (Halperin et al., 1982), Morocco (Loginova, 1972a), Portugal (Aulmann, 1913), Spain (Puton, 1878; Bolivar & Chicote, 1879; Vondracek, 1951b; Ramirez Gomez, 1956). CoMMENTS. A widely distributed species which occurs right across the southern part of the Mediterranean Basin. It is distinguished from other members of the genus by the characteristic pattern and somewhat oblong shape of the forewing. Livilla lautereri sp. n. (Figs 121, 122, 194-196, 292) DEscrIPTION. Coloration. Available specimens do not appear to have developed the full mature coloration. General coloration of head and thorax brownish yellow, abdomen slightly darker above; genitalia brown; forewing membrane clear with very pale yellow clouds and very distinct dark brown markings towards apex; veins yellowish basally becoming dark brown at wing apex; antennae brownish yellow, segments becoming infuscate towards apex; legs yellowish brown. Structure. Head (Fig. 122) with genal cones widely separate and longer than vertex. Forewing (Fig. 121) oblong-oval, almost parallel-sided, with broadly rounded apex; spinules confined to apical cells and around base of vein Cu; costal break and pterostigma absent; vein Rs strongly curved to wing margin; vein M evenly curved; vein M,,, reaching wing margin well above apex; cell cu,;, moderately large, vein Cu, straight in apical half. Metatibia with 4 or 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 194) simple; paramere (Fig. 195) with highly serrated anterior margin and large apical tooth; aedeagus (Fig. 196) very thin with angular apex. Female terminalia as in Fig. 292. Measurements. See Table 3. Holotype 0’, Algeria: Massiv de l’Aures, 6 km Arris, 1300 m, 6.vi.1980 (Remane) (BMNH). Paratype. 1 9, same data as holotype. HostT PLANT. Unknown. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 31 CoMMENTS. The forewing shape is similar to L. retamae but the pattern is quite different and characteristic, separating L. /autereri from other members of the genus. Livilla syriaca (L6w) comb. n. (Figs 123, 124, 197-199, 286) Floria syriaca Low, 1882b: 262; 1883: 252; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 35; Loginova, 1976a: 599; Halperin et al. , 1982: 34. Holotype 9, Syria [not located, missing from Low collection (NM)]. Floria syriac[e]a Low; Klimaszewski, 1973: 194. [Mis-spelling. ] DEscrIPTION. Coloration. Mature specimens with dorsum of thorax shining chocolate brown, remainder of thorax brownish yellow, occasionally with a greenish tinge; vertex and genal cones brownish yellow, occasionally greenish, marked along posterior and eye margins in darker brown; abdomen with dorsal sclerites brown, intersegmental membranes yellowish or greenish; ventral sclerites brown, occasionally greenish; genitalia yellow-brown to brown; forewing membrane clear to pale yellow, veins yellow to dark brown where they are overlaid with dark brown maculations; antennae yellowish brown, apices of segments 3—8 and whole of segments 9 and 10 brown. Structure. Head (Fig. 124) with genal cones triangular, widely divergent, about equal in length to vertex. Forewing (Fig. 123) oblong-oval but becoming broader towards the somewhat truncate apex; costal break and rudimentary pterostigma present; vein Rs strongly curved towards margin; vein M strongly sinous; vein M,,> meeting wing margin well above apex; cell cu,, relatively tall, vein Cu,, strongly curved. Metatibia with 5 thick black spurs; basal metatarsus with 1 spur. Male proctiger (Fig. 197) simple; paramere (Fig. 198) long and slender without thickened setae on basal posterior margin; aedeagus (Fig. 199) bulbous and slightly recurved at apex. Female terminalia as in Fig. 286. Measurements. See Table 3. MATERIAL EXAMINED 20,13 2, Cyprus (BMNH, MM). Host PLANT. Genista fasselata Decne and Genista acanthoclada D.C. PUBLISHED DISTRIBUTION. Syria (L6w, 18825) and possibly Israel (Halperin et al. , 1982). CoMMENTS. The types of this species are missing from the Low collection in the NM, Vienna. We have followed Loginova’s interpretation of the species which is, in all probability, correct. This species is distinguished by the forewing, which is strongly narrowed toward the base, and by the widely divergent genal cones. The spartiisuga-group Members of this group are characterised by the forewing pattern consisting primarily of small round maculations. Livilla spartiisuga (Puton) comb. n. (Figs 137, 138, 218-220, 276) Psylla spartiisuga Puton, 1876: 283. Lectotype CO’, ALGERIA (MNHN), here designated [examined]. Floria spartiisuga (Puton) Low, 1879: 593; 1883: 251; Puton, 1886: 92; Oshanin, 1907: 367; 1912: 128; Aulmann, 1913: 34; Loginova, 1971: 629; 1976a: 599. Description. Coloration. Mature specimens with dorsum of thorax orange-red with brownish markings; head including genal cones orange; abdominal sclerites brown, intersegmental membranes reddish orange; : genitalia orange-brown to dark brown; forewing membrane clear to pale yellow, veins yellow, maculations pale brown; antennae yellow with apices of segments black; legs yellowish orange. _ Structure. Head (Fig. 138) with genal cones slightly longer than vertex, stout and broadly rounded. Forewing (Fig. 137) elongate oblong-oval; with costal break and rudimentary pterostigma present; |spinules present throughout all cells; vein Rs moderately strongly curved towards margin; vein M evenly |curved throughout; vein M,,, reaching to wing apex; cell cu,;, moderately long and flat. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 218) simple; paramere (Fig. 219) long and slender; aedeagus (Fig. 220) with rounded bulbous apex. Female terminalia as in Fig. 276. 32. I. D. HODKINSON & D. HOLLIS Measurements. See Table 3. MATERIAL EXAMINED Lectotype 0’, Algeria: B6ne, with Puton type label (MNHN). 2 CO’, 4 9 paralectotypes, same data as lectotype (MNHN). Host PLANT. Genista sp. PUBLISHED DISTRIBUTION. Algeria (Puton, 1876). CoMMENTS. This species is close to siciliensis and known only from the type series. It can be separated from all other ‘spotted-wing’ species by the absence of maculations from a band stretching primarily across cell r, of the forewing. Livilla siciliensis sp. n. (Figs 131, 132, 209-211, 273) DESCRIPTION. Coloration. Immature specimens pale greenish to yellow throughout. Some of the more mature specimens are more yellowish and are beginning to develop orange markings on the dorsum of the thorax and vertex and the abdominal sclerites are beginning to darken. Genitalia yellow to brown; forewing membrane clear to pale yellowish, veins yellowish to pale brown; maculations brown; antennal segments 1—4 yellow, segments 3—4 infuscate at apex; segments 5—10 dark brown; legs dirty yellow. Structure. Head (Fig. 132) with genal cones rather stout, about as long as vertex, with broadly rounded apices. Forewing (Fig. 131) oblong-oval, almost parallel-sided; costal break and rudimentary pterostigma present; fine spinules present throughout all cells; vein R, curved towards margin; vein M long and very slightly sinuous; cell m4 small. Vein M,,> reaching to wing apex; cell cu, relatively flat. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 209) simple; paramere (Fig. 210) long and slender; aedeagus (Fig. 211) with slightly angular bulbous apex. Female terminalia as in Fig -273. Measurements. See Table 3. Holotype OC’, Italy: Sicily, Palermo c. 750 m, 28.v.1979 (D. & S. Sutton) (BMNH). Paratypes. 10 O’, 18 Y, same data as holotype except that some were collected on 26.v.1979 and 5.vi.1979. Host PLANT. Genista sp. ComMENTs. Morphologically this species is closest to L. maculipennis but it is much larger. Livilla cataloniensis (Hodkinson & White) comb. n. (Figs 127, 128, 203—205, 271) Floria cataloniensis Hodkinson & White, 1979a: 55. Holotype C’, FRANCE (BMNH) [examined]. DEscrIPTION. Coloration. Mature specimens with dorsal surface of head and thorax orange-yellow with paler longitudinal markings; genal cones and underside of thorax brownish yellow; abdominal sclerities brown, often darker above; intersegmental membranes yellow, occasionally with greenish tinge in younger specimens; genitalia yellow to yellow-brown; forewing membrane clear to very pale yellow throughout, maculae brown; antennal segments 1—2 yellow; segments 3—7 yellow with infuscate apex; segments 9-10 dark brown; legs brownish yellow. Structure. Head (Fig. 128) with genal cones conical, slightly longer than the vertex, with narrowly rounded apices. Forewing (Fig. 127) oblong-oval with broadly rounded apex; pterostigma rudimentary; costal break occasionally present; fine spinules present throughout all cells; vein Rs weakly curved towards margin; vein M evenly curved; vein M,,> reaching to wing apex; cell cu,, short and high; vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs at apex; basal metatarsus with 1 black spur. Male proctiger (Fig. 203) simple; paramere (Fig. 204) relatively stout and straight, without thick basal setae along posterior margin; aedeagus (Fig. 205) with bulbous apex. Female terminalia as in Fig. 271. Measurements. See Table 3. MATERIAL EXAMINED CO holotype, France: Pyrénées-Or., Argeles, 16.v.1977 (White) (BMNH). 15 ©’, 21 9 including paratypes, France and Spain (BMNH, IDH). There is also a lot of material from unnamed localities in the Heslop-Harrison collection (BMNH). W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 33 Host PLant. Ulex parviflorus Pourret. PUBLISHED DISTRIBUTION. France and Spain (Hodkinson & White, 1979a). CoMMENTS. This species can be distinguished from other spotted-wing species by the relatively broader forewing. Livilla hodkinsoni (Burckhardt) comb. n. (Figs 139, 140, 221-223, 277) Floria hodkinsoni Burckhardt, 1979: 391. Holotype CO’, GREECE (ETH) [examined]. DEscriPTION. Coloration. Immature specimens bright green throughout with orange markings on dorsum of head and thorax. Mature specimens with dorsal surface of head and thorax brown with yellow markings; underside of head and thorax with dark brown markings; abdominal sclerites dark brown, intersegmental membranes paler; genitalia yellow-brown to brown; forewing membrane clear to pale yellow with brown maculations; antennal segments 1—7 yellow, segments 4—7 infuscate at apex; segments 8—10 dark brown; legs brownish yellow. Structure. Head (Fig. 140) with genal cones longer than vertex, with somewhat angular apices. Forewing (Fig. 139) oblong-oval, with rudimentary pterostigma and costal break present; fine spinules present throughout all cells; vein R, moderately strongly curved towards margin; vein M slightly sinuous; vein M+ reaching to just below the wing apex; cell cu,, moderately long, of characteristic shape. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 221) simple; paramere (Fig. 222) without thickened basal posterior setae; aedeagus (Fig. 223) with bulbous apex. Female terminalia as in Fig. 277. Measurements. See Table 3 MATERIAL EXAMINED CO holotype, Greece: Pelopones, Bilos, 16.v.1979 (Burckhardt) (ETH). 13 0’, 15 9 including paratypes, Greece, including Crete (ETH, DB, BMNH, MM, PL). Host PLANT. Genista acanthoclada D.C., not a Cytisus species as stated in the original description (Burckhardt, pers. comm.). PUBLISHED DISTRIBUTION. Greece (Burckhardt, 1979). ComMENTS. L. hodkinsoni is closest to L. cataloniensis from which it is separated by the shape of cells m, and cu,, of the forewing. Livilla maculipennis sp. n. (Figs 129, 130, 206-208, 272) DEscriPTIONn. Coloration. Body colour of immature specimens bright green to yellow, usually with orange or orange-brown markings on dorsum of vertex and thorax. Mature specimens darker, general colour pale brown with darker brown markings; abdominal sclerites dark brown with yellow intersegmental mem- branes; forewing membrane clear, veins pale yellow, maculations brown; antennal segments brownish yellow basally becoming dark brown towards apex. Structure. Head (Fig. 130) with genal cones divergent, narrowly rounded at apex, slightly longer than vertex. Forewing (Fig. 129) elongate, oblong-oval, almost parallel-sided, pterostigma almost lost, costal break often present; fine spinules present throughout all cells; vein Rs moderately strongly curved towards wing margin; vein M slightly sinuous; vein M,,, reaching to wing apex; cell cu;, very long and low. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 206) simple; paramere (Fig. 207) curved posteriorly beyond middle, with numerous thickened setae on basal posterior margin; aedeagus (Fig. 208) with apex somewhat reniform. Female terminalia as in Fig. 272. Measurements. See Table 3. Holotype ©’, Spain: Huelva, 26 km, S. Almonte, 13.vi.1977 (Hollis) (BMNH). Paratypes. Spain: 11 ©’, 11 9, same data as holotype. Algeria: 7 0’, 3 2, Theniet al Had, Zedernwald, 1700 m, 31.vii.1980 (Remane) (PL); 10 0’, 6 9, N El Gor, 900 m, 28.vii.1980 (Remane) (PL); 7 0’, 11 9, Monte de Tlemcen, 18 km NW. Sebdou, 27.vii.1980 (Remane) (PL). Morocco: 1 0, N Taferaet, 25.vii.1980 (Remane) (PL). _ Host PLant. Genista sp. 34 I. D. HODKINSON & D. HOLLIS ComMENTS. L. maculipennis is separated from the other spotted-wing species by its small size and the elongate cell cu,, of the forewing. Livilla pyrenaea (Mink) comb. n. (Figs 133, 134, 212-214, 274) Psylla pyrenaea Mink, 1859: 430. Syntypes, FRANCE [not located]. Floria pyrenaea (Mink) Low, 1879: 592; 1883: 247; Chicote, 1880: 202; Puton, 1871: 438; 1886: 92; Oshanin, 1907: 367; 1912: 128; Aulmann, 1913: 34; Vondraéek, 1951b: 128; Loginova, 1976a: 599; Hodkinson & White, 1979a: 57. Floria (Floriella) pyrenaea (Mink); Ramirez Gomez, 1956: 87. Floria pyrenea (Mink); Klimaszewski, 1973: 194. [Mis-spelling. | DEscRIPTION. Coloration. Mature specimens with dorsal surface of head and thorax dark orange or orange-brown with paler longitudinal markings; genal cones and underside of thorax brownish yellow; abdominal sclerites dark brown with yellow intersegmental membranes, often with a greenish tinge in less mature specimens; genitalia yellow to brown; forewing membrane clear; maculae brown, absent from area adjacent to wing margin, making leading edge of wing appear white from a distance; antennal segments 1-2 yellow; segments 3-7 yellow with infuscate apex; segments 9-10 brown; legs brownish yellow. Structure. Head (Fig. 134) with genal cones somewhat triangular, divergent, longer than vertex. Forewing (Fig. 133) oblong-oval with a somewhat truncate apex; costal break and pterostigma usually absent; fine spinules present throughout all cells; vein Rs curved towards wing margin; vein M evenly curved; vein M,,> reaching to wing apex; cell cu,, moderately long. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 212) simple; paramere (Fig. 213) relatively short, parallel-sided; aedeagus (Fig. 214) robust, with slightly reniform apex. Female terminalia as in Fig. 274. Measurements. See Table 3. MATERIAL EXAMINED 20 0’, 19 2, France and Spain (BMNH, IDH, MM). Host pLants. Calicotome spinosa (L.) Link. and Genista pilosa L. PUBLISHED DISTRIBUTION. Spain (Chicote, 1880; Ramirez Gomez, 1956; Hodkinson & White, 1979a), France (Mink, 1859; Puton, 1871; Vondraéek, 1951b, Hodkinson & White, 1979a). CoMMENTs. A characteristic species with maculations absent from the leading margin of the forewing. It also differs from other spotted-wing species by having a slightly obliquely truncated wing apex. Livilla bimaculata sp. n. (Figs 149, 150, 254-256, 280) DEscrIPTION. Coloration. Mature specimens with dorsal surface of head and thorax pale reddish brown with paler markings; genal cones and ventral parts of thorax brownish yellow; abdominal sclerites dark brown; intersegmental membranes yellowish; genitalia brown; forewing membrane two-tone, clear and yellow-brown; maculae generally mid brown, those around the apex of vein Rs darker, almost black; antennae with segments 1—4 primarily yellow, segments 6—10 primarily dark brown; legs brownish yellow. Structure. Head (Fig. 150) with genal cones somewhat divergent, as long as vertex. Forewing (Fig. 149) elongate oval, broadest in basal half; fine spinules present throughout all cells; vein Rs strongly curved towards wing margin; vein M sinuous; vein M,,, reaching to wing apex; pterostigma at most rudimentary; costal break occasionally present. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 254) simple; paramere (Fig. 255) slender, with numerous stout spines on basal posterior margin; aedeagus as in Fig. 256. Female terminalia as in Fig. 280. Measurements. See Table 3. Holotype ©’, France: Corsica, Gorge du Ristomica nr Corte, 12.vi.1976, on Genista sp. (J.M.P.) (BMNH). Paratypes. 6 oO’, 10 2, same data as holotype. Italy: 4 0’, 4 9, Sardinia, Genova Borzonasea, Passo, Boco (Conci) 16.vi.1984 (IDH). HOostT PLANT. Genista salzmanii DC. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 35 Comments. This species is separated from other ‘spotted-wing’ Livilla species by the presence of a much darker patch of maculations around vein Rs. Livilla blandula (Horvath) comb. n. (Figs 144-146, 236-238, 278) Floria blandula Horvath, 1905: 277; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Ramirez Gomez, 1956: 84; Klimaszewski, 1973: 193; Loginova, 1976a: 599. LECTOTYPE ©’, Spain (TM) here designated [examined]. | Floria maura Vondraéek sensu Loginova, 1972a: 27. Misidentification. | DeEscriPTION. Coloration. Body colour of immature specimens brownish yellow. Mature specimens with dorsum of thorax and vertex chocolate brown with white markings; genal cones and underside of thorax brownish yellow; dorsum of abdomen brown, remainder pale green to yellow; genitalia greenish yellow to brown; forewing membrane two-tone, clear and yellow-brown; maculae mid to dark brown; antennae with segments 1—5 primarily yellow, segments 6—10 primarily brown. Structure. Head (Fig. 146) with genal cones slightly longer than vertex, slightly divergent, with apices narrowly rounded. Forewing (Figs 144, 145) oblong-oval; costal break and rudimentary pterostigma often present; fine spinules present throughout all cells; vein Rs weakly curved towards wing margin; vein M slightly sinuous; vein M,,, reaching to wing apex. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 236) simple; paramere (Fig. 237) with numerous thick setae on basal posterior margin; aedeagus (Fig. 238) with slightly hooked apex. Female terminalia as in Fig. 278. Measurements. See Table 3. MATERIAL EXAMINED Lectotype ©’, Spain: ‘Hispania. Ciudad Real, det. Horvath’ (TM). 10,4 Q (paralectotypes), same data as lectotype; 19 0’, 27 2, Portugal, Morocco (BMNH, ZMU, ZI). Host PLANT. Genista hirsuta Vahl. PUBLISHED DISTRIBUTION. Spain (Horvath, 1905), Morocco (Loginova, 1972a). ComMENTs. This is the species illustrated as maura by Loginova (1972a). The forewing of maura is relatively broader and with only scattered indistinct maculations. This species shows considerable variation in the wing pattern and the two extremes are illustrated for comparison. Livilla blandula is closest to L. lusitanica and L. pseudoretamae. It can be distinguished from the former by the narrower forewing and from the latter by the lack of a distinct dark brown forewing pattern. Livilla pseudoretamae sp. n. (Figs 157, 158, 233-235, 285) DescriPTIon. Coloration. Vertex, genal cones and dorsum of thorax pale greyish brown, occasionally with a greenish tinge, thorax with chocolate brown markings; abdominal sclerites ranging from green to brown, with pale intersegmental membranes; genitalia dirty yellow to brown; forewing membrane clear, veins yellow to very pale brown; wing pattern light chocolate-brown; antennae dirty yellow basally, segments 5—10 dark brown; legs brownish yellow throughout. The specimens to hand have probably not developed their full coloration. Structure. Head (Fig. 158) with genal cones slightly longer than vertex, with evenly rounded apices. Forewing (Fig. 157) oblong-oval; with costal break and rudimentary pterostigma present; fine spinules present throughout all the cells; vein Rs moderately strongly curved towards margin; vein M moderately sinuous; vein M,,> short but reaching to wing apex; cell cu,;, short and high, vein Cu, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 233) simple; paramere (Fig. 234) straight, with thickened setae present on basal posterior margin; aedeagus (Fig. 235) with a slightly hooked apex. Female terminalia as in Fig. 285. Measurements. See Table 3. Holotype oO’, Algeria: Gran Kabylie, Djurdjura, 3.viii. 1980 (Remane) (BMNH). Paratypes. 2 0’, 5 9, same data as holotype; 3 0’, 9 2, Kabylie, Col de Talmetz (Remane). (BMNH, FL.) Host plant. Unknown. 36 I. D. HODKINSON & D. HOLLIS ComMENTS. This species is very close to L. blandula but the wing pattern, which lacks the characteristic spots, is more distinct and superficially resembles that of Livilla retamae. Livilla lusitanica sp. n. (Figs 147, 148, 251-253, 279) DESCRIPTION. Coloration. Mature specimens with dorsum of head and thorax brown with whitish markings; genal cones and underside of thorax brownish yellow; abdominal sclerites dark brown, intersegmental membranes yellowish brown; genitalia brown; forewing membrane two-tone, clear and yellow-brown; maculae mid to dark brown; antennae with segments 1—5 primarily yellow, segments 6—10 primarily dark brown; legs brownish yellow. Structure. Head (Fig. 148) with genal cones stout, about as long as vertex. Forewing (Fig. 147) broadly oblong-oval, with costal break and rudimentary pterostigma present; fine spinules present throughout all cells; vein Rs moderately strongly curved towards wing margin; vein M slightly sinuous; vein M,,, reaching wing apex; cell cu,, strongly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 251) simple; paramere (Fig. 252) stout, with dense thickened setae on the basal posterior margin; aedeagus (Fig. 253) with apex somewhat reniform. Female terminalia as in Fig. 279. Measurements. See Table 3. Holotype CO, Portugal: 29 km E. Coimbra, 21.vi.1977 (Hollis) (BMNH). Paratypes. 7 O', 10 , same data as holotype; 3 oO’, 3 , Porto, 2—12.viii.1962 (J. Abraham & L. Horascek) (BMNH). HOostT PLANT. Genista triacanthos Brot. CoMMENTS. This species resembles blandula in several respects but the forewing is consistently shorter and broader and the genal cones slightly shorter. Livilla magna sp. n. (Figs 135, 136, 215-217, 275) DEscrIPTION. Coloration. Immature specimens pale green to yellow throughout. Mature specimens with vertex and dorsum of thorax brownish yellow with orange or chocolate brown markings; underside of thorax often with most of sclerites dark brown to black; genal cones brownish yellow; sclerites of abdomen dark brown, occasionally paler along ventral parts; intersegmental membranes brownish yellow; forewing two-tone, clear to pale yellow contrasted with amber; maculations medium brown; antennal segments 1—4 yellow, segments 3—4 infuscate at apex; segments 5—10 dark brown; legs brownish yellow, femora occasionally darkened above. Structure. Head (Fig. 136) with divergent triangular genal cones as long as vertex. Forewing (Fig. 135) elongate oblong oval; rudimentary pterostigma and costal break present; vein Rs moderately curved to margin; fine spinules present in all cells; cells cu,, and m4 of average size for genus; vein M,,> reaching to wing apex. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 215) simple; paramere (Fig. 216) long and thin with a kink in the middle; aedeagus (Fig. 217) long with an angular bulbous apex. Female terminalia as in Fig. 275. Measurements. See Table 3. Holotype CO’, Italy: Sicily, Catania, 1.vi.1979 (D. & S. Sutton) (BMNH). Paratypes. 5 0’, 9 9, same data as holotype; 2 0’, 8 2, same data as holotype except 4.vi.1979; 10,7 9, Sicily, Etna, 1100 m, 16.vii.1977 (D’Iirso) (PL); 1 0’, 3 Q, Sicily, Etna, 1100 m, 19.ix.1977 (Asche) (PL). Host PLANT. Genista aetnensis (Biv.) DG. CoMMENTS. This species is much larger than the other spotted-wing species. Morphologically it is closest to L. siciliensis but it differs in the shape of the forewing and details of the male genitalia. Livilla poggii (Conci & Tamanini) comb. n. (Figs 141-143, 224-226, 281) Floria poggii Conci & Tamanini, 1984a: 43. Holotype O’, ITALy: Sardinia (MCSN) [not examined]. DescriPTIon. Coloration. General coloration dirty yellow to greyish yellow throughout, occasionally with brown markings on the dorsum of the thorax; genitalia yellow to dark brown; forewing membrane clear to W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS i # | yellowish but covered in brown maculations which are often coalescent; in the female the maculations are spread throughout the wing whereas in the male there is a transverse maculation-free band across the middle; also in the latter the apical maculations appear darker than the basal ones; antennal segments 1—8 yellow, segments 3-8 apically infuscate; segments 9-10 dark brown; legs orange-yellow. Structure. Head (Fig. 143) with genal cones weakly divergent, with narrowly rounded apices, about as long as vertex. Forewing (Figs 141, 142) elongate oval, more so in the female; costal break and pterostigma usually absent; fine spinules present in all cells; vein R, strongly curved towards wing margin; vein M strongly sinuous; vein M,,, meeting margin just above wing apex; cell cu,, short and tall; vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 224) simple; paramere (Fig. 225) relatively short and stout; aedeagus (Fig. 226) with an irregularly angled apex. Female terminalia as in Fig. 281. Measurements. See Table 3. MATERIAL EXAMINED 20,3 2 (paratypes), Italy (IDH). Host PLANT. Genista corsica (Loisel.) DC in Lam. & DC. PUBLISHED DISTRIBUTION. Italy, Sardinia (Conci & Tamanini, 1984a). ComMENTs: This is an interesting species which is strongly sexually dimorphic in the wing-pattern, an unusual occurrence in the psyllids. The host plant is endemic to Sardinia and Corsica. Residual assemblage Livilla spectabilis (Flor) comb. n. (Figs 161, 162, 242-244, 268) Psylla spectabilis Flor, 1861: 362. Syntypes 0’, @ FRANCE (not located). Floria spectabilis (Flor) Low, 1879: 594; 1883: 251: 1888a: 20; Chicote, 1880: 202; Ferrari, 1888: 76; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Horvath, 19185: 331; Haupt, 1935: 242; Schaefer, 1949a: 56, 1949b: 31; Vondracek, 1951b: 128; Klimaszewski, 1968b: 781; 1973: 194; Novak & Wagner, 1962: 43; Loginova, 1976a: 599; Burckhardt, 1983: 54. DEscrIPTION. Coloration. Immature specimens pale green to greenish yellow throughout. Mature speci- mens with genal cones, vertex and dorsum of thorax dirty orange-yellow with darker orange and paler streaks; abdominal sclerites dark brown, intersegmental membranes orange-yellow; genitalia orange- yellow to brown; forewing membrane clear basally becoming orange-brown apically, with a dark spot around vein Cu,,; antennae dirty yellow, except segments 3 and 4 apically darkened, segments 5 to 10 brown; legs dirty yellow. Structure. Head (Fig. 162) with massive genal cones which are much longer than the vertex. Forewing (Fig. 161) elongate oval, of characteristic shape, with a strongly curved costal margin throughout; costal break and rudimentary pterostigma normally absent; fine spinules present throughout all cells; vein Rs weakly curved towards margin; vein M evenly curved; vein M,,, reaching to wing apex; cell cu, relatively small; vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 242) simple; paramere (Fig. 243) straight and slender; aedeagus (Fig. 244) with recurved apex. Female terminalia as in Fig. 268. Measurements. See Table 3. MATERIAL EXAMINED 10 0’, 9 Q, Italy, Greece including Crete, Yugoslavia, France (MM, BMNH, IDH, DB, PL). There is 1 O&O, 1 9 inthe ZI, Leningrad labelled Uppsala, Sweden, F. Ossiannilsson. Dr Ossiannilsson informs us that this is a mistake; the species does not occur in Sweden. Host PLanvT. Spartium junceum L. PUBLISHED DISTRIBUTION. France (Puton, 1871), Italy (Ferrari, 1888), Portugal (Aulmann, 1913), Spain (Chicote, 1880), Switzerland (Schaefer, 1949a,b; Burckhardt, 1983). : CoMMENTS. This species has no known close relative within the genus. It is separated from other members _by the narrow forewing of characteristic shape and the very long genal cones. | 38 I. D. HODKINSON & D. HOLLIS Livilla variegata (LOw) comb. n. (Figs 163, 164, 245-247, 269) Floria variegata Low, 1882b: 261; 1883: 253; 1888a: 20; Ferrari, 1888: 76; Puton, 1886: 92; Oshanin, 1907: 367, 1912: 128; Aulmann, 1913: 35; Klimaszewski, 1973: 194; Loginova, 1976a: 599; Hollis, 1978: 149; Hodkinson & Hollis, 1980: 171; Hodkinson & White, 1979a: 63; White & Hodkinson, 1982: 25; Burckhardt, 1983: 54. Syntypes, Oo’ 2, YUGOSLAVIA (MM) [examined]. Floria alpina Cerutti, 1939a: 447; 1939b: 583; Schaefer, 1949a: 55; 1949b: 31; Klimaszewski, 1973: 193; Loginova, 1976a: 599; Tamanini, 1977: 111. Lectotype ©’, SwirzERLAND (MZL) [not examined]. [Synonymised by Burkchardt, 1983: 54. ] DEscriPTION. Coloration. Immature specimens pale green to greenish yellow with orange markings on dorsum of thorax. Mature specimens with dorsum of head and thorax orange-yellow with paler longitudi- nal markings; genal cones orange-yellow; abdominal sclerites dark brown, intersegmental membranes yellow; genitalia orange-brown; forewing membrane clear to pale yellow basally, apical suffusion grey to pale yellowish brown, veins pale yellow to very pale brown; antennae dirty yellow, apices of segments 3 to 6 and whole of segments 7 to 10 dark brown; legs dirty yellow. Structure. Head (Fig. 164) with genal cones slender, slightly longer than vertex, with narrowly rounded apices. Forewing (Fig. 163) oblong-oval; costal break and rudimentary pterostigma present; veins very delicate; dense fine spinules present throughout all cells; vein Rs weakly curved to margin; vein M evenly curved; vein M,,, reaching to wing apex; cell cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 245) simple; paramere (Fig. 246) slender, with thickened setae present on basal posterior margin; aedeagus (Fig. 247) with bulbous apex. Female terminalia as in Fig. 269. Measurements. See Table 3. MATERIAL EXAMINED 27 O&O, 32 9, Yugoslavia, Rumania, Italy, Switzerland, France, Spain, England (MM, IDH, BMNH, DB). Host pLants. Laburnum anagyroides Medicus and Laburnum alpinum (Miller) Berchtold & J. Presl. PUBLISHED DISTRIBUTION. France (Hodkinson & White, 1979a), Great Britain (Hollis, 1978; Hodkinson & Hollis, 1980; White & Hodkinson, 1982), Italy (Ferrari, 1888; Tamanini, 1977), Switzerland (Cerruti, 1939a, b; Schaefer, 1949a,b; Burckhardt, 1983), Yugoslavia (Low, 18825; 1888a). ComMMENTS. Floria alpina was, for a long time, recorded as a separate alpine species but it has now been shown to be synonymous with L. variegata. L. variegata can be separated from other members of the genus by the broad forewing which lacks a distinct pattern and by the veins which are unusually fine and lightly coloured. Livilla genistae Ramirez Gomez (Figs 153, 154, 227-229, 283) Livilla genistae Ramirez Gomez, 1956: 98; Klimaszewski, 1973: 195; Loginova, 1976a: 600. NEOTYPE CO, SPAIN (BMNH) here designated. DEscriPTION. Coloration. Mature specimens with head and thorax yellowish brown with slightly darker reddish brown markings; abdominal sclerites occasionally darker than thorax, terminalia yellow-brown to brown; forewing membrane clear, without pattern; veins yellowish brown; antennal segments 1-2 yellow, segments 3—4 yellow and infuscate at apex; segments 5—10 dark brown; legs brownish yellow. Structure. Head (Fig. 154) with genal cones almost as long as vertex, somewhat divergent with broadly rounded apices. Forewing (Fig. 153) elongate oval without pterostigma or costal break; spinules present throughout all cells; vein Rs strongly curved towards wing margin; vein M evenly curved; vein M42 reaching to just below wing apex; cell cu,, short and tall; vein Cu,, strongly curved. Metatibia with 5 apical thick black spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 227) simple; paramere (Fig. 228) sinuous, with thick setae at base of posterior margin; aedeagus (Fig. 229) with weakly angled apex. Female terminalia (Fig. 283) with very short circumanal pore ring. Measurements. See Table 3. MATERIAL EXAMINED Neotype o’, Spain: Huesca, Barluenga, 1.vi.1976, on ? Genista (Hollis) (BMNH); 8 2, Spain (BMNH). W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 39 Host PLANT. This was omitted from the original description: the name genistae suggests that it feeds on a ‘Genista’ species. The neotype designated above was collected from a plant thought to be a Genista species while 4 of the additional females were collected from Cytisus purgans. PUBLISHED DISTRIBUTION. Spain (Ramirez Gomez, 1956). CoMmMENTS. The original type-material of this species has been destroyed. The above material collected in Spain corresponds closely with the original description. This species is somewhat intermediate between the short coriaceous wing Livilla species (ulicis-group) and the longer wing forms with spotted or longitudi- nally striped wing patterns (spartiisuga- and horvathi-groups). It is recognised by the oval wing and the absence of any wing patterning. Livilla adusta (L6w) comb. n. (Figs 159, 160, 239-241, 267) Floria adusta Low, 1882b: 260; 1883: 230; Puton, 1886: 92; 1889: 113; Oshanin, 1907: 367; 1912: 128; Aulmann, 1913: 34; Klimaszewski, 1973: 193; Loginova, 1976a: 599. ? Holotype 9, SPAIN [missing from Low collection (NM)]. DEscrIPTION. Coloration. Immature specimens greenish yellow throughout, developing pale orange markings on dorsum of thorax. Mature specimens with genal cones, vertex and dorsum of thorax brownish orange with indistinct brown or paler markings; abdominal sclerites dark brown to black, intersegmental membranes orange-brown. Forewing two-tone, clear and orange-brown; veins yellow to yellow-brown; antennae dirty yellow, segments 3—4 darkened apically, segments 5—10 dark brown; legs dirty yellow. Structure. Head (Fig. 160) with genal cones as long as vertex, slightly divergent with narrowly rounded apices; forewing (Fig. 159) elongate oblong-oval, with evenly rounded apex; costal break and pterostigma rudimentary; fine spinules present throughout all cells; veins Rs upturned to wing margin; vein M,,> reaching to wing apex. Metatibia with 5 thick black spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 239) simple but broadest at base; paramere (Fig. 240) long, thin, parallel-sided, with large inner apical tooth, bearing a few stout setae at base of posterior margin; aedeagus as in Fig. 241. Female terminalia (Fig. 267) with a slightly upturned apex to the proctiger. Measurements. See Table 3. MATERIAL EXAMINED 90, 12 9, Algeria and Spain (BMNH, MNHN, MM). Host PLANT. Unknown. Some specimens in BMNH were collected from Cytisus scoparius ? PUBLISHED DISTRIBUTION. Spain (Low, 1882b; Aulmann, 1913). ComMENTSs. This species is not very close to any other member of the genus. It is easily recognised by the distribution of darker pigment along the forewing veins which gives a furrowed appearance. Livilla maura (Vondra¢éek) comb. n. (Figs 151, 152, 282) Floria maura Vondracek, 195la: 119; 1951b: 128; Klimaszewski, 1973: 193; Loginova, 1976a: 599. Syntypes 2, Morocco (MM) [examined]. DEscriPTION. Coloration. Mature specimens with vertex and dorsum of thorax deep brownish orange with darker brown markings; genal cones brown; abdomen dark brown to black, intersegmental membranes brownish orange; genitalia brownish yellow to dark brown; forewing membrane, pale brownish orange throughout, veins concolorous with membrane; apical pattern very diffuse, pale brown; antennae with segments 1—4 dirty yellow, 3—4 apically brown, the remaining segments brown; legs with femora pale brown, the remainder orange-yellow. Structure. Head (Fig. 152) with genal cones about as long as vertex, relatively slender, with narrowly rounded apices. Forewing (Fig. 151) oblong-oval, relatively broad; costal break and rudimentary ptero- stigma present; fine spinules present throughout all cells; vein M evenly curved; vein M,,> reaching to wing apex; cell cu, relatively short with vein Cu,, strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Female terminalia as in Fig. 282. Male unknown. Measurements. See Table 6. 40 MATERIAL EXAMINED I. D. HODKINSON & D. HOLLIS 2 9 (syntypes), Morocco (MM). In addition, there are 2 heads, 5 wings, 2 female genitalia and other oddments on slides labelled 63-69 Floria maura type series (MM). Host PLANT. Unknown. PUBLISHED DISTRIBUTION. Morocco (Vondraéek, 1951a,b). ComMENTs. This species was described from several females. The male is unknown and this makes it difficult to characterize the species. It appears to differ from other members of the genus in possessing a relatively broad forewing of uniform brownish orange coloration. Table 3 Measurements of Livilla species. Males and females are given separately and all values are in mm. adusta bimaculata bivittata blandula burckhardti cataloniensis cognata genistae hodkinsoni horvathi klapperichi lautereri lusitanica maculipennis magna maura nervosa nigralineata poggit pseudoretamae pyrenaea Head width 0-63-0-74 0-67—0-74 0-70—0-73 0-72—0-76 0-82—0-83 0-74—0-76 0-64—0-69 0-69-0-74 0-69—0-79 0-71—0-83 0-54—0-59 0-58—0-63 0-65—0-66 0-71-0-73 0-62—0-78 0-71—0-85 Antennal Forewing length length 1-47-1-66 2-25-—2-50 1-55-1-69 2-28-2-48 1-32-1-37 1-84—1-91 1-25-1-34 2-15-2-24 1:63-1:67 1-91-1-95 1:65-1:68 2-48-2-50 1-14-1-32 1-70-—2-01 1-22-1-40 1-70—2-30 1-14-1-16 1-86-1-90 1-13-1-16 2-18—2-20 1-18-1-34 1-69-2-04 1-19-1-36 1-96—2-33 1-63 2-10 1-76 2:46 1-21 29 1-22-1-24 2-16—2-33 1-42-1-54 1-90-—2-00 1-52—1-68 »2-26=2-38 1-80-2-04 2-51-2-73 1-77—2-04 2-68-—3-06 = 1-50 — 1-86 — ot 1-63 2-2 1-17 0-67—0-85 1-09-1-24 1-84—1-97 1-25-1-40 1-84—2-13 1-16—1-38 2-03—2-24 2°33—2:56 3-09—3-35 2:48-2:67 3-60—3-77 unknown 1:23 1-81-—2-27 1-88-2:06 2-39-2-72 1-80—2-18 2-60—3-15 1-76 233 1-97 2°71 1-06-1-41 1-33-1-57 1-06-1-19 1-76—1-94 1-38-1-40 1-91—1-99 1-40-1-42 2-18-2-29 1-50-1-72 2-02—2-49 1-54-1-82 2-33-2-85 Female proctiger length 0-59—0-63 0-54—0-59 0-82—0-84 0-56-0-62 0-67-0-74 0.93 0-87-0-93 0-71-0-79 0.75-0-89 0-60 0-71 0-53-0-55 0-54—0-59 1-04-1-08 0-55—0-58 1-09-1-15 0-86 0-58-0-59 0-60-0-61 0-76—0-83 Male proctiger length 0-37—0-39 0-35—0-38 0-40-0-41 0.29-0-34 0-32 0-30-0-34 0.38 0-40 0.32-0-33 0-40-0-47 0-24 0.28 0.29--0-32 0-32-0-34 0-44-0-50 Male paramere length 0-35—0-37 0-33-0-36 0-31-0-33 0.22-0-26 0-29 0.25.-0-29 0.35 0-42 0.32-0-33 0.29--0-35 0.23 0.29 0-24--0-26 0.27-0-31 0-44-0-47 male unknown 0-45-0-49 0-38 0-27—0-29 0-31—0-36 0-37—-0-41 0-39-0-41 0-31 0-26—0-27 0-24—0-25 0-26—0-31 Male aedeagus length 0-26—0-28 0-28--0-30 0-31-0-33 0- 190-23 0.29 0.24-0-26 0-31 0.36 0.28-0-29 0.28-0-29 0.20 0.25 0- 17-0-21 0.22-0-23 0-38—0-40 0-35—0-39 0-30 0-19—0-20 0-22—0-23 0-25—0-29 : Table 3 — cont. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 41 Female Male Male Male Head Antennal Forewing proctiger proctiger paramere aedeagus width length length length length length length radiata m 0:73-0:82 1:59-1:78 2-70—2-80 = 0-40-0-41 0-35-0-37 0-30 f 0-78—-0-84 1-82-1-:92 2-60-2-70 0-85 — — -- retamae m 0-84-1-00 2-19-2-75 2-81-3-18 _ 0:46-0:53 0-43-0-47 0-37-0-40 f 0:95-1:09 2:21-2:75 2-96-3-51 1-00-1-08 — _ _ siciliensis m 0:74-0:81 1:72-1:78 2-35-2-50 — 0-37-0-40 0-37-0-39 0-31-0-32 f 0-80-0-85 1-86-1-88 2:69-2:79 0-83—0-87 — — — spartiisuga m 0:73-0:75 1:81-1:83 2-43-2-44 — 0-36-0-38 0-31-0-33 0-33-0-35 f 0-81-0-82 1:94-1:96 2-80—2-86 0-94 — — — spectabilis m 0:94-1:00 2-80-3-06 3-42—3-52 — 0:51-0:54 0-41-0-44 0-35-0-37 f 1:00-1:09 3-29-3-52 3-83-4-14 0-90-0-92 --- —- —— syriaca m 0-67—0-70 1:67-1:88 1-95—1-97 — 0-35-0-42 0-30-0-34 0-24-0-27 f 0-65-0-78 1:89-1:90 2-17-2-30 0-67—0-69 — —_ ulicis m 0:70-0:75 1:37-1:50 1-69-1-81 — 0:36-0:37 0-35-0-36 0-28—0-29 f 0-74—-0-79 1-38-1:60 1-84-2-10 0-78—0-85 — — = variegata m 0:67-0:78 1:78-2:23 2-76—3-06 — 0:40-0:44 0-30—0-34 0-26—0-29 f 0-71-0-87 1:74-2:25 2-82-3-59 0-82-0-84 — — — vicina m 0-82—0-83 1:41-1:50 2-12—2-20 — 0-36-0-45 0-38-0-44 0-33-0-35 f 0-86—-0-87 1-39-1-42 2-37-2-39 0-77-0-83 _- -— —- vittipennella m 0-71-0-76 1:74-1:81 2-35-2-58 — 0:37-0:42 0-37-0-40 0-30—0-33 f 0:67-0:75 1-48-1-79 2-42-—2-66 0-80-0-81 — — — ARYTAINILLA Loginova Spartina Heslop-Harrison, 1951: 443; 1961a: 417. Type-species: Psylla spartii Guérin, by monotypy. [Homonym of Spartina Harris & Burrows, 1891. ] Lindbergia Heslop-Harrison, 1951: fig. 2, a,b. Nomen nudum [no included species] (nec Lindbergia Riedel, 1958). Arytainilla Ramirez Gomez, 1956: 76. Nomen nudum [type-species not designated]. Alloeoneura (Hispaniola) Ramirez Gomez, 1956: 91, partim. Nomen nudum [type-species not desig- nated]. Arytainilla Loginova, 1972a: 17. Type-species: Psylla delarbrei Puton, 1873, designated by Loginova, pezes i]. CoMMENTS. All described species of Arytainilla occur in the west Mediterranean area and the Macarone- sian Islands (Table 4). This is an heterogeneous genus and is almost certainly not a monophyletic group. The type-species, A. delarbrei, is exceptional in having 8-segmented antennae, long genal cones, an oval forewing with a short pterostigma and unusual male parameres. Most other congeners have 10-segmented antennae, short genal cones, an oblong-oval forewing and simple parameres. Arytainilla species are well illustrated by Sulc (1907a; 1910a,b) and Loginova (1972a; 1976b) and there is little point in repeating their descriptions. Further collecting is required to establish relationships between the species and we have confined ourselves to recognising species-groups where possible (Table 4). The spartiophila-group is separated from the rest of Arytainilla by the characteristic upturned and pointed apex of the female proctiger. The dividens-group comprises species that have oblong-oval forewings, short genal cones, moderate to long female terminalia of normal form and male parameres of various shape but most usually slender and with a bulge or projection on the anterior margin. The proboscidea-subgroup is defined by the elongate rostral segments and the spotted patterning along the forewing veins. Members of the dividens- subgroup have parameres with a large bulge on the anterior margin, whereas those in the prognata- subgroup have a smaller ill-defined bulge. The equitans-subgroup consists of species similar to those above but with parameres that lack the anterior bulge. The remaining species tend to have unique characters that set them apart from the rest of the genus. A. delarbrei has already been mentioned; it does, however, have a tendency to the same form of female terminalia as that of the spartiophila-group. A. cytisi has a truncate female ventral valve and a massive Ovipositor. A. umbonata has a more elongate, parallel-sided forewing and a distinct bulge on the ventral valve of the female terminalia. The female of A. egena has long genal cones and short terminalia that are of 42 I. D. HODKINSON & D. HOLLIS a characteristic shape. Arytainilla nubivaga (Loginova) comb. n. and A. devia (Loginova) comb. n., two distinctive Macaronesian species, are transferred to Arytainilla from Arytaina until a more satisfactory position can be found for them within the classification. Nevertheless, they do show some similarity with delarbrei and the remaining species, A. ima, particularly in the longer genal cones and the forewing shape of A. devia. The paramere shape in these two Canary Island species is complex and unique in the genus. Table 4 Species-groups within Arytainilla with information on host plant and distribution. spartiophila-group spartiophila (Forster, 1848) spartiicola (Sulc, 1907) gredi (Ramirez Gomez, 1956) comb. n. dividens-group dividens-subgroup dividens Loginova, 1976b modica Loginova, 1976b hakani Loginova, 1972a cognata Loginova, 1972a pileolata Loginova, 1976b proboscidea-subgroup proboscidea Loginova, 1976b nigralineata Loginova, 1976b prognata-subgroup prognata Loginova, 1976b incuba Loginova, 1976b equitans-subgroup equitans Loginova, 1976b diluta, Loginova, 1976b Residual species delarbrei (Puton, 1873) cytisi (Puton, 1876) ima Loginova, 1972a egena Loginova, 1972a umbonata Loginova, 1976b devia (Loginova, 1976b) comb. n. nubivaga (Loginova, 1976b) comb. n. Host plant Cytisus scoparius Cytisus scoparius Cytisus scoparius? Cytisus proliferus unknown unknown unknown unknown Adenocarpus anagyrus Adenocarpus or Cytisus unknown unknown unknown unknown Genista tinctoria Calicotome spinosa unknown Retama raetam indet. legume Cytisus proliferus Adenocarpus anagyrus Cytisus fragrans Distribution Britain Spain France Netherlands Switzerland Austria Portugal France Switzerland Spain Canaries Canaries Morocco Morocco Canaries Canaries Canaries Canaries Madeira Canaries Canaries France Spain Algeria France Israel (IDH) Italy Spain Yugoslavia Turkey Morocco Morocco, Libya (BMNH) Canaries Canaries W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 43 Discussion The legume-feeding psyllid fauna of the west Palaearctic appears to comprise a number of evolutionary lines that are identified and diagnosed in this paper as separate monophyletic genera or species-groups within the Acizziinae and Arytaininae. Arytaina is defined by characters that include the broad form of the male paramere, the aedeagus with a strongly hooked apex and the lack of a pterostigma and costal break in the forewing. Livilla is defined by the narrow elongate paramere and the long bulbous or weakly hooked apex of the aedeagus. Cyamophila possesses a paramere that resembles that of Arytaina but the propleurites/ pronotum structure is more advanced and the costal break and pterostigma are well-developed in the forewing. We have suggested that Arytainilla is heterogeneous and probably comprises a number of distinct evolutionary lines. Acizzia is separated from the other genera, into the subfamily Acizziinae, by the presence of lateral lobes on the male proctiger. The extent of homoplasy in several of the other characters examined makes it difficult to suggest clear relationships between genera. Thus we have concluded that Arytaina, Livilla, Cyamophila and Arytainilla, together with the monotypic central Asian genus Astragalita, should be retained, for the present, in the subfamily Arytaininae. We recognise, however, that there are no good autapomorphies that define the subfamily and separate it from the Psyllinae. It remains, nevertheless, a convenient and widely recognised grouping. The Acizziinae is a distinct evolutionary line warranting subfamily status. Arytaina, Livilla and some groups within Arytainilla are most diverse in the Mediterranean Basin and are associated with papilionoid legumes of the tribe Genisteae, which itself has a primarily Mediterranean distribution. By contrast, most Cyamophila species occur in the arid regions of central Asia and are associated with other tribes of papilionoid legumes, particularly Galegeae, Trifoleae, Loteae and Sophoreae. The genus is not known from Africa and the one west European species, C. prohaskai, represents the western limit of this genus. Acizzia species are widespread throughout Africa, Arabia, India, South East Asia and Australasia. They are primarily associated with the equally widely distributed mimosoid legume genera Acacia and Albizia. Acizzia appears to be an old genus with a very wide distribution that reaches its natural northern and western limits in the area under consideration. Legume systematics, particularly of the tribe Genisteae, have been the subject of much debate (Bisby, 1981). It is of interest to see if the pattern of host plant relationships for the different psyllid groups can contribute to an understanding of legume classification and vice versa. In a recent classification of the legumes Bisby (1981) split the Genisteae into two subtribes, the Lupininae and the Genistinae. He subdivided the latter into three main groups of genera (Table 5). His Cytisus-group contains the arytainine host genera Cytisus, Laburnum and Chamaecytisus; the Genista-group contains Genista, Ulex and Retama, while the outliers include Adenocarpus, Calicotome and Spartium. Table 5 shows the distribution of psyllids across their host plants in the Genisteae. Each record indicates a known association between a psyllid species and a species of host plant. Species of Livilla are concentrated on hosts within Bisby’s Genista-group, particularly the large genus Genista itself, but also on Ulex and Retama. However, some species also feed on genera in Bisby’s other two groups of Genistinae. Of the two psyllid species known to feed on the outlier group, spectabilis is monophagous on Spartium junceum, whereas pyrenaea is recorded from Calicotome and Genista. Among the three species feeding on the Cytisus-group (radiata, cognata and variegata) cognata also has a known host in the Genista-group, while radiata is oligophagous on several species of Chamaecytisus. The core species of Livilla, the spartiisuga- and horvathi-groups, tend to occur on Genista. The morphologically isolated species, such as variegata and spectabilis , tend to occur on the more taxonomically isolated groups of host plants. By contrast with Livilla, the hosts of Arytaina and Arytainilla are more evenly spread across the Genisteae. Within the Cytisus-group, Arytaina genistae and A. maculata occur on Cytisus and Chamaecytisus, whereas Arytainilla species such as gredi, spartiophila, spartiicola, devia, dividens and nubivaga are concentrated on Cytisus. The Genista-group supports just four species, Arytaina genistae (record requires confirmation), A. torifrons and Arytainilla delarbrei 44 I. D. HODKINSON & D. HOLLIS Table 5 Host plant relationships of Livilla, Arytaina and Arytainilla species across the Genisteae. The number of species in each genus (N) is taken from Bisby (1981) except for Argyrolobium which is taken from Flora Europaea (Tutin et al. , 1964). Number of host records N Livilla Arytaina Arytainilla Total Subtribe Lupininae 0 0 0 0 Subtribe Genistinae Cytisus-group Laburnum 2 pi 0 0 Z Hesperolaburnum 1 0 0 0 0 Podocytisus 1 0 0 0 0 Cytisophyllum 1 0 0 0 0 Petteria 1 0 0 0 0 Argyrocytisus 1 0 0 0 0 Cytisus 33 2 1 6 9 Chaemaecytisus 30 6 5 2 13 Genista-group Retama + 3 0 1 4 Genista 87 18 y 1 21 Echinospartum 3) 0 0 0 0 Stauracanthus Z 0 0 0 0 Ulex 20 pi 0 0 Z Outlier group Argyrolobium 2 0 0 0 0 Adenocarpus 15 0 2 2 4 Calicotome 2 i if 1 3 Erinacea 1 0 0 0 0 Spartium it 1 0 0 1 Gonocytisus 3 0 0 0 0 on Genista, and Arytainilla egena on Retama. The outlier plant group supports Arytaina putonii and Arytainilla cytisi on Calicotome, and Arytaina adenocarpi and Arytainilla devia on Adeno- carpus. Table 5 indicates that the larger the plant genus the more psyllid species that genus is likely to support, with a psyllid feeding record for about every four species of plants. This relationship can be interpreted in two ways. 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Transactions of the Entomological Society of London 24: 525=-569. — 1879. Description of a new species of the family Psyllidae — Floria horvathi n. sp. Entomologist’s Monthly Magazine 16: 84-85. — 1880. Description of the nymph of Arytaina genistae Latr. Entomologist’s Monthly Magazine 17: 132-133. — 1882a. Food plants and times of appearance of the species of Psyllidae found in Great Britain together with others which may be expected to occur here. Entomologist’s Monthly Magazine 19: 13-15. — 1882b. The British Psyllina, with corrections in the synonymy. Entomologist’s Monthly Magazine 18: 253-256. Smreczynski, S. 1954. Materialy do fauny pluskwiakow Polski 3. Psylloidea. Fragmenta Faunistica 7: 1-146. Sulc, K. 1905. Revise Psyll sbirky Dudovy. Casopis Ceskoslovenske Spolecnosti Entomologicke 2: 1-4. 1907. Prispevky ku poznani Psyll. 1. Psylla spartii Guerin-Loew a Psylla spartiicola n. sp. Rozpravy Ceske Akademie Ved, Praha (section 2) 16: 1-8. — 1910a. Uvod do studia, synoptika tabulka a synonymicky katalog druhu rodu Psylla, palaearcticke oblasti. Sitzungsberichte der K. B6hmischen Gesellschaft der Wissenschaften 2 (1909): 1—46. 1910b. Prispevky k poznani II. Rozpravy Ceske Akademie Ved, Praha (section 2) 19: 1-32. Szulczewski, J. W. 1927. Materialien zur Psyllidfauna Grosspolens. Sprawozdania Komisji Fizyograficznej Oraz Materialy do Fiziografii Kraju 61: 197-204. Tamanini, L. 1955. Alcuni nuovi repert di Psyllidi Italiani e Francesci (Homoptera, Psyllina). Bolletino della Societa Entomologica Italiana 85: 10-11. — 1977. Notizie corologiche e morphologiche su alcuni psyllidi poco noti delle prealpi (Homoptera: Psylloidea). Studi Trentini di Scienze Naturali 54: 103-119. Thomson, C. C. 1877. XXIX. Ofversigt af Skandinaviens Chermes-arter. Opuscula Entomologica (Editit C. G. Thomson), Trelleborg 8: 820-841. Tuthill, L. D. 1943. The Psyllids of North America north of Mexico (Psyllidae: Homptera) (Subfamilies Psyllinae and Triozinae). 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PALAEARCTIC LEGUME-FEEDING PSYLLIDS 53 Figs1-8 Acizzia species. 1,2, bicolorata: (1) forewing; (2) head. 3, 4, hollisi: (3) forewing; (4) head. 5, 6, uncatoides: (5) forewing; (6) head. 7, 8, acaciaebaileyanae: (7) forewing; (8) head. 54 I. D. HODKINSON & D. HOLLIS GC genitalia, lateral view, (10) & paramere, outer view; f aedeagus. 12-14, hollisi: (12) CO genitalia, lateral view; (13) O paramere, outer (15) CO’ genitalia, lateral view; (16) o f aedeagus. 18-20, acaciaebaileyanae: (18) oO genitalia, lateral view; (19) CO’ paramere, outer view; (20) apical segment of aedeagus. Figs 9-20 Acizzia species. 9-11, bicolorata: (9) (11) apical segement o view; (14) apical segment of aedeagus. 15-17, uncatoides: paramere, outer view; (17) apical segment o ie W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 55 Figs 21-30 Cyamophila species. 21-25, glycyrrhizae: (21) forewing; (22) head; (23) O genitalia, lateral view; (24) ©” paramere, inner view; (25) apical segment of aedeagus. 26-30, prohaskai: (26) forewing; (27) head; (28) CO genitalia, lateral view; (29) O" paramere, inner view; (30) apical segment of aedeagus. 56 I. D. HODKINSON & D. HOLLIS 6, stoklosai (after Klimaszewski & Lodo head; (33) C’ genitalia, lateral view; (34) CG paramere; (35) apical segmen terminalia, lateral view. 37-42, medicaginis: (37) forewing; (38) head; (39) So Figs 31-42 Cyamophila species. 31-3 (40) o’ paramere; (41) apical segment of aedeagus; s): (31) forewing; (32) t of aedeagus; (36) 9 genitalia, lateral view; (42) 9 terminalia, lateral view. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS ST Figs 43-50 Arytaina species. 43, 44, longicella: (43) forewing; (44) head. 45, 46, genistae: (45) forewing; (46) head. 47, 48, maculata: (47) forewing; (48) head. 49, 50, maculata (Spanish material): (49) forewing; (50) head. 58 I. D. HODKINSON & D. HOLLIS Figs 51-58 Arytaina species. 515°525 forewing; (54) head. 55, 56, putonit; (55) forewing; angustatipennis: (51) forewing; (52) head. 53, 54, atlasiensis: (53) (56) head. 57, 58, torifrons; (57) forewing; (58) head. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 59 Figs 59-64 Arytaina species. 59, 60, hispanica: (59) forewing; (60) head. 61, 62, adenocarpi: (61) forewing; (62) head. 63, 64, magnidentata; (63) forewing; (64) head. 60 I. D. HODKINSON & D. HOLLIS | view; (66) CO’ paramere, inner genitalia, lateral view; (69) o aterial); (71) C 74, africana: O Figs 65-74 Arytaina species. 65-67, genistae: (65) CO genitalia, latera view; (67) apical segment of aedeagus. 68-70, maculata: (68) O paramere, inner view; (70) apical segment of aedeagus. 71-73, maculata (Spanish m genitalia, lateral view, (72) & paramere, inner view; (73) apical segment of aedeagus. genitalia, lateral view (after Heslop-Harrison). W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 61 Figs 75-84 Arytaina species. 75-77, putonii: (75) O genitalia, lateral view; (76) GO’ paramere, inner view; (77) apical segment of aedeagus. 78-80, torifrons: (78) O genitalia, lateral view; (79) CO’ paramere, inner view; (80) apical segment of aedeagus. 81, angustatipennis: CO genitalia, lateral view; 82-84, hispanica: (82) & genitalia, lateral view; (83) 6’ paramere, inner view; (84) apical segment of aedeagus. 62 I. D. HODKINSON & D. HOLLIS Figs 85-94 Arytaina and Cyamophila species. 85-87, A. adenocarpi: (85) CO genitalia, lateral view; (86) GC" paramere, inner view; (87) apical segment of aedeagus. 88-90, A. magnidentata: (88) O genitalia, lateral view; (89) OC’ paramere, inner view; (90) apical segment of aedeagus. 91, A. atlasiensis: terminalia, lateral view. 92, A. hispanica: 9 terminalia, lateral view. 93, C. glycyrrhizae: 2 terminalia, lateral view. 94, C. prohaskai: 9 terminalia, lateral view. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 63 figs 95-102 Arytaina and Livilla ? terminalia, lateral view. 95, A. longicella. 96, A. genistae. 97, A. maculata. 98, L. radiata. 99, A. adenocarpi. 100, A. putonii. 101, A. angustatipennis. 102, A. torifrons. 64 I. D. HODKINSON & D. HOLLIS 109 Figs 103-110 Livilla species. 103, 104, ulicis: (103) forewing; forewing; (106) head. 107, 108, vicina: (107) forewing; (108) h (110) head. (104) head. 105, 106, bivittata: (105) ead. 109, 110, cognata: (109) forewing; W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 65 PN So 3 BS ee TIN af ‘ RES a Figs 111-118 Livilla species. 111, 112, nervosa: (111) forewing; (112) head. 113, 114, nigralineata: (113) forewing; (114) head. 115, 116, horvathi: (115) forewing; (116) head. 117, 118, vittipennella: (117) forewing; (118) head. 66 I. D. HODKINSON & D. HOLLIS Figs 119-126 Livilla species. 119, 120, retamae: (119) forewing; (120) head. 121, 122, lautereri: (121) forewing; (122) head. 123, 124, syriaca; (123) forewing; (124) head. 125, 126, radiata; (125) forewing; (126) head. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS Figs 127-134 Livilla species. 127, 128, cataloniensis: (127) forewing; (128) head. 129, 130, maculipennis: (129) forewing; (130) head. 131, 132, siciliensis: (131) forewing; (132) head. 133, 134, pyrenaea: (133) forewing; (134) head. 67 I. D. HODKINSON & D. HOLLIS 68 Figs 135-143 Livilla species. 135, 136, magna: (135) forewing; (136) head. 137, 138, spartiisuga: (137) forewing; (138) head. 139, 140, hodkinsoni: (139) forewing; (140) head. 141-143, poggii: (141) 2 forewing; (142) CO forewing; (143) head. Figs 144-150 Livilla species. 144-146, blandula: (146) h head. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS (144) forewing of lectotype; (145) forewing (Morocco); ead. 147, 148, lusitanica: (147) forewing; (148) head. 149, 150, bimaculata: (149) forewing; (150) 69 70 I. D. HODKINSON & D. HOLLIS Figs 151-158 Livilla species. 151, 152, maura: (151) forewing; forewing; (154 ) head. 155, 156, klapperichi: forewing; (158) head. (152) head. 153, 154, genistae: (153) (155) forewing; (156) head. 157, 158, pseudoreta mae; (157) W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 71 159 161 163 Figs 159-166 Livilla species. 159, 160, adusta: (159) forewing; (160) head. 161, 162, spectabilis, (161) forewing; (162) head. 163, 164, variegata: (163) forewing; (164) head. 165, 166, burckhardti: (165) forewing; (166) head. Tz I. D. HODKINSON & D. HOLLIS 178 Figs 167-178 Livilla species. 167-169, ulicis: (167) & genitalia, lateral view; (168) O' paramere, outer view; (169) apical segment of aedeagus. 170-172, bivittata: (170) & genitalia, lateral view; (171) o paramere, outer view; (172) apical segment of aedeagus. 173-175, vicina: (173) C genitalia, lateral view; (174) C’ paramere, outer view; (175) apical segment of aedeagus. 176-178, cognata: (176) ej genitalia lateral view; (177) CO’ paramere, outer view, (178) apical segment of aedeagus. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 73 Figs 179-190 = Livilla species. 179-181, nervosa: ( 179) & genitalia, lateral view; (180) O’ paramere, outer view; (181) apical segment of aedeagus. 182-184, nigralineata: (182) CO genitalia, lateral view; (183) CO’ paramere, outer view; (184) apical segment of aedeagus. 185-187, horvathi: (185) genitalia, lateral view; (186) CO’ paramere, outer view; (187) apical segment of aedeagus. 188-190, vittipennella: (188) 0 genitalia, lateral view; (189) CO’ paramere, outer view; (190) apical segment of aedeagus. 74 I. D. HODKINSON & D. HOLLIS Figs 191-202 Livilla species. 191-193, retamae: (191) & genitalia, lateral view; (192) O' paramere, outer view; (193) apical segment of aedeagus. 194-196, lautereri: (194) CO genitalia, lateral view; (195) o paramere, outer view; (196) apical segment of aedeagus. 197-199, syriaca: (197) genitalia, lateral view; (198) CG" paramere, outer view; (199) apical segment of aedeagus. 200-202, radiata: (200) genitalia, lateral view; (201) CO’ paramere, outer view; (202) apical segment of aedeagus. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 75 Figs 203-214 Livilla species. 203-205 , cataloniensis: (203) CO genitalia, lateral view; (204) 0" paramere, outer view; (205) apical segment of aedeagus. 206-208, maculipennis: (206) O genitalia, lateral view; (207) O paramere, outer view: (208) apical segment of aedeagus. 209-211, siciliensis: (209) 0 genitalia, lateral view; (210) O’ paramere, outer view; (211) apical segment of aedeagus. 212-214, pyrenaea: (212) CO genitalia, lateral view; (213) O' paramere, outer view: (214) apical segment of aedeagus. 76 I. D. HODKINSON & D. HOLLIS 221 Figs 215-226 Livilla species. 215-217, magna: (215) & genitalia, lateral view; (216) C& paramere, outer view; (217) apical segment of aedeagus. 218-220, spartiisuga: (218) C genitalia, lateral view; (219) & paramere, outer view; (220) apical segment of aedeagus. 221-223, hodkinsoni: (221) CO genitalia, lateral view; (222) OC’ paramere, outer view; (223) apical segment of aedeagus. 224-226, poggii: (224) oS genitalia, lateral view; (225) O’ paramere, outer view; (226) apical segment of aedeagus. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS TF Figs 227-238 — Livilla species. 227-229, genistae: (227) CO genitalia, lateral view; (228) O" paramere, outer view; (229) apical segment of aedeagus. 230-232, klapperichi: (230) & genitalia, lateral view; (231) 0 paramere, outer view; (232) apical segment of aedeagus. 233-235, pseudoretamae: (233) O& genitalia, lateral view; (234) CO’ paramere, outer view; (235) apical segment of aedeagus. 236-238, blandula: (236) © genitalia, lateral view; (237) O paramere, outer view; (238) apical segment of aedeagus. 78 I. D. HODKINSON & D. HOLLIS 240 246 Figs 239-250 Livilla species. 239-241, adusta: (239) O' genitalia, lateral view; (240) C paramere, outer view; (241) apical segment of aedeagus. 242-244, spectabilis: (242) C genitalia, lateral view; (243) O paramere, outer view; (244) apical segment of aedeagus. 245-247, variegata: (245) oO genitalia, lateral view; (246) CO” paramere, outer view; (247) apical segment of aedeagus. 248-250, burckhardti: (248) S genitalia, lateral view; (249) O’ paramere, outer view; (250) apical segment of aedeagus. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 254 Figs 251-262 Livilla, Arytaina, Acizzia and Cyamophila species. 251-253, L. lustianica: (251) oO genitalia, lateral view; (252) o’ paramere, outer view; (253) apical segment of aedeagus. 254-256, L. bimaculata: (254) CO genitalia, lateral view; (255) CO paramere, outer view: (256) apical segment of aedeagus. 257, 258, Arytaina africana (after Heslop-Harrison): (257) forewing; (258) head. 259, Livilla, head and prothorax. 260, Arytaina, head and prothorax. 261, Cyamophila, head and prothorax. 262, Acizzia hollisi, ? terminalia, lateral view. 80 I. D. HODKINSON & D. HOLLIS tata. 265, vicina. 266, cognata. 267, Figs 263-270 Livilla, 9 terminalia, lateral view. 263, ulicis. 264, bivit adjusta. 268, spectabilis. 269, variegata. 270, burckhardti. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 81 Figs 271-278 Livilla, 274, pyrenaea. 275, magna. 276, spartiisuga. 277, hodkinsoni. 278, blandula. Q terminalia, lateral view. 271, cataloniensis. 272, maculipennis. 273, siciliensis. 82 I. D. HODKINSON & D. HOLLIS Figs 279-286 Livilla, 9 terminalia, lateral view. 279, 283, genistae. 284, klapperichi. 285, pseudoretamae. lusitanica. 280, bimaculata. 281, poggit. 282, maura. 286, syriaca. W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 83 Figs 287-294 Livilla and Acizzia, terminalia, lateral view. 287, L. nervosa. 288, L. nigralienata. 289, L. horvathi. 290, L. vittipennella. 291, L. retamae. 292, L. lautereri. 293, A. uncatoides. 294, A. acaciaebaileyanae. 84 Acacia 2, 3, 4,5, 43 Acacia baileyana 5 confusa 6 floribunda 6 koa 6 koaia 6 podalyriaefoliae 5 raddiana 5, 6 saligna 6 verniciflua 6 Acacieae 4 Adenocarpus 42, 43, 44 Adenocarpus anagyrus 42 complicatus 13 hispanicus 13 Albizia 2, 3, 4, 43 Albizia lophantha 6 Anthyllis 8 Anthyllis vulneraria 9 Argyrocytisus 44 Argyrolobium 44 Astragalus 8 Astragalus sp. 10 Calicotome 43, 44 Calicotome spinosa 18, 22, 34, 42 Chamaecytisus 43, 44 Chamaecytisus austriacus 15, 27, 29 borysthenicus 17, 29 heuffelii 15 ratisbonensis 17, 26, 29 supinus 26 Cytisophyllum 44 Cytisus 42, 43, 44 Cytisus-group 43, 44 I. D. HODKINSON & D. HOLLIS Index to host plants Cytisus fragrans 42 proliferus 42 purgans 39 scoparius 2, 15, 39, 42 Echinospartium 44 Erinacea 44 Galegeae 2, 7, 43 Genista 43, 44 Genista-group 43, 44 Genista acanthoclada 31, 33 aetnensis 23, 36 corsica 22, 37 fasselata 21, 31 germanica 26 hirsuta 23, 35 hispanica 19 pilosa 22, 34 radiata 25, 27, 28 salzmanii 34 spp. 16, 25,32, 33, 39 tinctoria 15, 24, 42 triacanthos 23, 36 Genisteae 2, 3, 11, 19, 43, 44 Genistinae 43, 44 Glycyrrhiza 8 Glycyrrhiza glabra 8 Gonocytisus 44 Hesperolaburnum 44 Ingae 4 Laburnum 43, 44 Index Invalid names are in italics; principal references are in bold acaciae (Psylla) 4 acaciaebaileyanae (Acizzia) 4 (key), 5, 6, 10 acaciaebaileyanae (Psylla) 5 acaciaebaileyanae-group 4 (key), 5 acaciae-baileyanae (Arytaina) 5 acaciae-baileyanae (Psylla) 5 Acizzia 2, 3 (key), 4, 10, 43 Acizziinae 2, 3 (key), 4, 43 adenocarpi (Arytaena) 13 adenocarpi (Arytaina) 12, 13 (key), 16, 17, 18, 44 adusta (Floria) 39 adusta (Livilla) 21 (key), 39, 40 africana 12 (key), 13-14, 18 albizziae (Acizzia) 11 albizziae (Arytaina) 11 Allaeoneura 19 Alloeoneura 2, 19, 29, 41 alpina (Floria) 38 Amblyrhina 2, 10, 11, 19, 26 Amblyrrhina 25 Amorphicola 2 angustatipennis (Amblyrhina) 14 angustatipennis (Arytaina) 12 (key), 14, 18 annosa (Floria (Brinckitia)) 19 antennata (Retroacizzia) 11 Arylaina 15 Arytaena 15, 24 Arytaina 2, 3 (key), 7, 10-11, 17, 18, 24, 42, 43, 44 Arytainilla 2, 4 (key), 41, 42, 43, 44 Arytaininae 2, 3 (key), 4, 7, 43 Arytainini 2, 7 assimilis (Arytaina) 11 Laburnum alpinum 38 anagyroides 38 spp. 23 Leguminosae 7 Lemboptropis nigricans 29 Loteae 2, 7, 43 Lupininae 43, 44 Medicago 8 Medicago sativa 9 Ononis spinosa 24 Petteria 44 Podocytisus 44 Retama 43, 44 Retama monosperma 30 raetam 30, 42 sp. 21 sphaerocarpa 30 Rosaceae 7 Sophoreae 2, 7, 43 Spartium 43, 44 Spartium junceum 21, 37, 43 Stauracanthus 44 Trifoleae 2, 7, 43 Ulex 43, 44 Ulex europaeus 24 parviflorus 33 Vicia 8 Vicia sp. 9 assimilis (Ceanothia) 11 Astragalita 2, 43 Ataenia 15 atlasiensis 12 (key), 14, 18 bicolor (Arytaina) 11 bicolor (Ceanothia) 11 bicolorata (Acizzia) 5 (key), 7, 10 bicolorata (Psylla) 7 bimaculata 22 (key), 34-35, 40 bivittata 3, 4, 20 (key), 24, 25, 40 blandula (Floria) 35 blandula (Livilla) 23 (key), 35, 36, 40 boharti (Arytaina) 11 boharti (Ceanothia) 11 brevigena (Arytaina) 11 brevigena (Insnesia) 11 Brinckitia 19 W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 85 burckhardti 3, 20 (key), 25, 40 burckhardti-group 19, 20 (key), 25 callunae (Psylla) 23 cataloniensis (Floria) 31 cataloniensis (Livilla) 22 (key), 32-33, 40 Ceanothia 11 Chermes 15 Ciriacreminae 2 cognata Léw (Amblyrhina) 25 cognata Low (Amblyrrhina) 25 cognata Loginova (Arytainilla) 42 cognata (Léw) (Livilla) 3, 20 (key), 25-26, 40, 42, 43 coleoptera (Psylla) 23 cornicola (Arytaina) 11 crawfordi (Insnesia) 11 Cyamophila 2, 3 (key), 7, 9, 10, 43 Cyamophilini 2 cytisi Becker (Psyllodes) 29 cytisi (Puton) (Arytainilla) 41, 42, 44 delarbrei (Arytainilla) 41, 42, 43 delarbrei (Psylla) 41 devia (Arytaina) 11 devia (Arytainilla) 4, 11, 42, 43, 44 diluta 42 dividens 42, 43 dividens-group 41, 42 dividens-subgroup 41, 42 egena 41, 42, 44 equitans 42 equitans-subgroup 41, 42 essigi (Arytaina) 11 essigi (Ceanothia) 11 Euphalerus 11 Euryconus 11 fabra (Psylla) 7 fasciata (Arytaina) 11 flava (Arytaina) 11 flava (Insnesia) 11 Floria 2, 19 Floriella 19 genistae Fabricius (Psylla) 15 genistae (Latreille) (Arylaina) 15 genistae (Latreille) (Arytaena) 15 genistae (Latreille) (Arytaina) 2, 12 (key), 14-15, 16, 17, 18, 43 genistae (Latreille) (Chermes) 15 genistae (Latreille) (Chermes (Ataenia)) 15 genistae Latreille (Psylla) 10, 14 genistae Ramirez Gomez (Livilla) 3, 20 (key), 38-39, 40 glycyrrhizae (Cacopsylla) 8 glycyrrhizae (Cyamophila) 8 (key), 10 glycyrrhizae (Psylla) 8 glycyrrhizae (Psyllodes) 8 gredi 42, 43 hakani 42 hispanica 12 (key), 16, 18 Hispaniola 41 hodkinsoni (Floria) 33 hodkinsoni (Livilla) 22 (key), 33, 40 hollisi 5 (key), 6, 10 hollisi-group 4 (key), 6 horvathi (Floria) 26 horvathi (Livilla) 21 (key), 26-27, 40 horvathi-group 19, 20 (key), 26, 27, 39, 43 ima 42 incuba 42 Insnesia 11 insolita (Arytaina) 11 insolita (Ceanothia) 11 iolani (Arytaina) 11 iolani (Insnesia) 11 isitis (Euphalerus) 11 isitis (Psylla) 11 klapperichi 21 (key), 28-29, 40 lactea (Psylla) 29 lautereri 21 (key), 30-31, 40 Lindbergia 41 lineata (Floria) 28 Livilla 2, 4 (key), 19, 26, 29, 35, 39, 40, 43, 44 longicella 11 (key), 16, 18 lusitanica 23 (key), 35, 36, 40 maculata (Amblyrhina) 16 maculata (Amblyrrhina) 16 maculata (Arytaina) 12 (key), 14, 15, 16-17, 18, 43 maculipennis 22 (key), 32, 33-34, 40 maculosa (Amblyrhina) 16 magna Hodkinson & Hollis (Livilla) 23 (key), 36, 40 magna Crawford (Psyllopa) 10, 15 magnidentata 13 (key), 17, 18 maura Loginova (Floria) 35 maura Vondraéek (Floria) 39, 40 maura Vondraéek (Livilla) 22, 23 (key), 35, 39-40 medicaginis (Cacopsylla) 9 medicaginis (Cyamophila) 8 (key), 9,10 medicaginis (Psylla) 9 meridionalis (Arytaina) 11 meridionalis (Insnesia) 11 mitella (Arytaina) 11 mitella (Ceanothia) 11 modica 42 montana (Arytaena) 24 montana (Arytaina) 24 mopanei (Arytaina) 11 mopanei (Retroacizzia) 11 Neopsylla 4 nervosa 20 (key), 27, 40 nigralineata Loginova (Arytainilla) 42 nigralineata Hodkinson & Hollis (Livilla) 21 (key), 27, 40 nubivaga (Arytaina) 11 nubivaga (Arytainilla) 4, 11, 42, 43 obscura (Acizzia) 11 obscura (Psyllopa) 11 odontopyx 8 pechai (Amblyrhina) 11 pechai (Spanioneura) 11 Peripsyllopsis 11 pileolata 42 poggii (Floria) 36 poggii (Livilla) 22 (key), 36-37, 40 proboscidea 42 proboscidea-subgroup 41, 42 prognata 42 prognata-subgroup 41, 42 prohaskai (Cyamophila) 8 (key), 9, 10, 43 prohaskai (Psylla) 9 Pseudacanthopsylla 2 pseudoretamae 21 (key), 35-36, 40 Psylla 11, 15, 19, 23 Psyllidae 2 Psyllinae 2, 4, 7, 43 Psyllopa 10 pulchra Crawford, 1919 (Arytaina) 11 pulchra (Crawford, 1919) (Euryconus) 11 pulchra Crawford, 1920 (Arytaina) 11 pulchra (Crawford, 1920) (Insnesia) 11 punctinervis (Arytaina) 11 punctinervis (Euphalerus) 11 putoni 17 putonii (Amblyrhina) 17 putonii (Arytaina) 12 (key), 17-18, 19, 44 pyrenaea (Floria) 34 pyrenaea (Floria (Floriella)) 34 pyrenaea (Livilla) 22 (key), 34, 40, 43 pyrenaea (Psylla) 19, 34 pyrenea 34 radiata (Aleuroneura) 29 radiata (Allaeoneura) 29 radiata (Alloeoneura) 29 radiata (Arytaina) 19, 29 radiata (Livilla) 20 (key), 29, 41, 43 radiata (Psylla) 29 : radiata-group 19, 20-21 (key), 27, 29 ramarkrishni 11 retamae (Alloeoneura) 30 retamae (Floria) 30 retamae (Floria (Brinckitia)) 30 86 retamae (Livilla) 21 (key), 30, 31, 36, 41 retamae (Psylla) 30 Retroacizzia 11 siciliensis 22 (key), 32, 36, 41 Spanioneura 11 spartii Guérin (Psylla) 41 spartii (Hartig) (Arytaina) 15 spartii Hartig (Psylla) 10, 15 spartiicola 42, 43 spartiisuga (Floria) 31 spartiisuga (Livilla) 22 (key), 31-32, 41 spartiisuga (Psylla) 31 spartiisuga-group 19, 21-22 (key), 31, 39, 43 Spartina 41 spartiophila (Arytainilla) 2, 42, 43 spartiophila (Psylla) 18 spartiophila-group 41, 42 spectabilis (Floria) 37 spectabilis (Livilla) 20, 21 (key), 37, 41, 43 I. D. HODKINSON & D. HOLLIS spectabilis (Psylla) 37 spinosa (Acizzia) 11 spinosa (Arytaina) 11 stocklosai 8 (key), 10 syriaca (Floria) 31 syriaca (Livilla) 21 (key), 31, 41 syriacea 31 thakrei (Arytaina) 11 thakrei (Insnesia) 11 torifrons (Amblryhina) 18 torifrons (Arytaina) 12 (key), 16, 18-19, 43 torifrons (Psylla) 10, 18 tuberculata (Arytaina) 11 tuberculata (Insnesia) 11 turkiana (Amblyrhina) 11 turkiana (Spanioneura) 11 uichancoi (Arytaina) 11 uichancoi (Insnesia) 11 ulicis (Curtis, 1835) (Arytaina) 15 ulicis Curtis, 1835 (Psylla) 15 ulicis (Curtis, 1835) (Arytaena) 15 ulicis Curtis, 1836 (Livilla) 4, 19, 20 (key), 23-24, 25, 41 ulicis-group 19, 20 (key), 23, 39 umbonata 41, 42 uncata (Psylla) 5 uncatoides (Acizzia) 4 (key), 5-6, 10 uncatoides (Psylla) 5 unctata 5 variabilis (Arytaina) 11 variabilis (Insnesia) 11 variegata (Floria) 38 variegata (Livilla) 22, 23 (key), 38, 41, 43 vicina (Floria) 24 vicina (Livilla) 3, 4, 20 (key), 24-25, 41 virgina 11 vitipennella 28 vittipennella (Floria) 26, 28 vittipennella (Livilla) 21 (key), 28, 41 vittipennella (Psylla) 28 sek ; ‘ oy ; ey ap % fae BA, S Be Led aaa it 5 Ty ‘" t ] ¢ ' = ‘ b a ; » & if a a : . Y; a 7 — ae, t- i eee oo tt a a Grapacione’ Vi eggs Spregitherccnd ® 4 wae ea geget Ciera (Pr rey st... sera lactis? Gewtpieay BS md tare hp hep 8. 5 °-- Te png robes 4 ethan) 's ‘Pee Th hag) AA Es, . pe wigs 4) a bain (hav ).. 9243 mre #6 afr : wa Neyer fy A Le tt. @, A? ct ergerd] eae 2 as Ue ‘ae yo ) ~ ores 4 ue vce a = 2 itl ee Aa p's ine sore if ig > pegs vl haat Cy ite Filey Tt } + sc erin (ae waaay’ oy che Ae ar 4 Ra) \ aaah tial, Aes ve Anke ad - ts Cit \ ’ “4 pits Oy \ ap.tt) pai re + ghee: ttre ; ~~ e« a ~ . = ; -. ; a Sut Li = —~ =a >< ae 7 — Oy ot C = an: © Prt | British Museum (Natural History) Milkweed butterflies: their cladistics and biology P.R. Ackery & R. I. Vane-Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of their biology have stimulated far more attention than can be justified by species numbers alone. In recent years, an expansive literature has grown, considering aspects of their courtship and pre-courtship behaviour, migration, larval hostplan associations, mimicry and genetics. The popularity of danaines among biologists can certainly be attributed to this combination, within one small group, of so many of the factors that make butterflies such an interesting group to study. The obvious need to place this wealth of biological data within an acceptable systematic framework provided the impetus for this volume. Started eight years ago within the conventions of evolution by natural selection and Hennig’s phylogenetic systematics, the book is now largely about natural history (what the animals have and do, where they live and how they develop) and natural groups — as revealed by a form of analysis approaching that practised by the new school of ‘transformed cladistics’. The authors have prepared a handbook that will appeal to a wide range of biologists, from museum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. ISBN 0 565 00893 5. 1984. Price £50. Titles to be published in Volume 56 The legume-feeding psyllids (Homoptera) of the west Palaearctic Region By I. D. Hodkinson & D. Hollis A review of the Malvales-feeding psyllid family Carsaridae (Homoptera) By D. Hollis The world genera of Rhadalinae (Coleoptera: Melyridae) By E. R. Peacock Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester Bullet . of the British Museum (Natural History) A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera) BRITISH MUSEUM (NATURAL HISTORY) - 2? NOV 1987 RESENTED ENTOMOLOGY LIBRARY D. Hollis | ntomology series Vol 56 No2 29 October 1987 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of clan carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 SBD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1987 The Entomology series is produced under the editorship of the Keeper of Entomology: Laurence A. Mound = Manager ee W. Gerald Tremewan aw iciieacagi slanting \ ‘ a Fo ig : j <4 J i iv eee i tae ’ 4 as : . . j ,. . Sree ranean wre od Pid i? ; fi. te Pool om, 5 > in broad contact; costal break absent (present in Protyora); radular spinules sharply defined into narrow triangular patches, often absent from cu;, a patch of spinules present in cell c+sc. Ventral sense organs of hind femur in a basal position and not arranged linearly; hind tibia with a well-developed basal spine; hind basitarsus with a single apical spur. Male subgenital plate with a pair of secondary lobes anterior to parameres, these lobes appearing to be sclerotised projections arising from the membrane lining the inner surface of the subgenital plate. Final instar larva elongate, clearly divided into head, thorax and abdomen; antennae elongate, 10-segmented; legs elongate, tarsal arolium sessile and fan-shaped or globular; wing buds small, without humeral lobes; thoracic sclerites poorly differentiated; caudal region of abdomen differentiated and bearing convoluted pore bands, anus terminal or terminodorsal; body setae mainly simple but scattered, small, lanceolate setae present on caudal sclerites marginally and submarginally. HostTpLants. Malvales (Sterculiaceae, Bombacaceae and Malvaceae). Discussion. An historical account of the early conceptual development of the family is given by Hollis (1976: 4). Heslop-Harrison (1958) considered the group as a tribe within the Ciriacreminae including the genera Carsidara, Mastigimas, Mesohomotoma, Protyora, Epicarsa and Diceraopsylla, and separated these genera from another of his Ciriacremine tribes, the Tenaphalarini, containing Tenaphalara, Strogylocephala, Togepsylla and Tyora. Hollis (1984) referred Diceraopsylla to the Aphalaridae and White & Hodkinson (1985) placed Togepsylla in the same family. Vondraéek (1957: fig. 52) considered the Carsidaridae as the sister-group of the Triozidae, differentiat- ing both families from the Ciriacreminae which he placed in the Psyllidae. Klimaszewski (1964: fig. 7) agreed with these concepts. Bekker-Migdisova (1973) recognised seven subfamilies in her interpretation of the Carsidaridae which amounted to a very large assemblage of many quite unrelated genera. White & Hodkinson (1985: 242, 273) considerably restricted their concept of the family to the genera Mastigimas, Tenaphalara, Protyora, Mesohomotoma, Paracarsidara, Carsidara and tentatively Aconop- sylla, Carsidaroida, Nesiope and Tyora. They placed Epicarsa in the Phacopteronidae, considering the anastomosis of Rs and M_,> as the important synapomorphy. For reasons given below (p. 90) Mastigimas is now removed from the family and placed provisionally in the Calophyidae; Carsidaroida and Nesiope are synonyms of Tyora, and Aconopsylla is referred to the Aphalaridae, close to Haplaphalara and Diclidoph- 90 D. HOLLIS lebia which are also Malvales-feeding. Brown (1985) removed Epicarsa from the Phacopteronidae to the Carsidaridae, using the presence of supplementary lobes on the male subgenital plate as a more reliable synapomorphy. This character, and the presence of a non-tracheate rs-m crossvein in the forewing, are the best pair of autapomorphies for the Carsidaridae. A crossvein of this type is unique to the family. The supplementary male subgenital lobes occur elsewhere in-the Psylloidea only in the Oriental Cedrela- feeding genus Trisetipsylla Yang & Li (1985a) which, in other characters, is a member of the family Psyllidae. The male hypovalves and tracheate rs-m crossvein of Ciriacremum (Psyllidae) are not homo- logues (Hollis, 1976). White & Hodkinson diagnosed the family on the following larval characters: anal pore field arranged in convoluted bands; dorsal surface of abdomen lacking distinct sclerites; body margin and dorsal surface without sectasetae or derivable structures. To this character-set can be added that poorly defined lanceolate setae are present marginally and submarginally on the caudal plate. These same authors postulated that the Carsidaridae formed the sister-group of the Triozidae, citing the absence of a costal break and the presence of fewer than six apical spurs on the hind tibia as adult synapomorphies, but they suggested no larval synapomorphies. These two adult characters are probably unreliable as they are subject to homoplasy within the Psylloidea generally. Brown (1985) considered the family to be the sister-group of the Ficus-feeding Homotomidae, citing the ‘cleft head’, weakly sclerotised A> vein and the absence of A; in the hindwing and a patch of spinules in c+sc of the forewing as synapomorphies. This is a more acceptable hypothesis, although A, is present in the hindwing of many homotomids. Further synapomorphies for the two groups are the development of a pair of strong epiphyses on the metapostnotum, and the position and arrangement of the ventral sense organs of the hind femur. The former structures appear to function as ‘tethering posts’ for both pairs of wings when these are in the resting position. Similar but probably non-homologous structures occur on the metas- cutellum in some triozids. The basal position of the ventral sense organs on the hind femur is considered to be primitive (Hollis, 1984: 11) but in homotomids and carsidarids they are not linearly arranged and the proximal sensillum is placed at right-angles to the other two. White & Hodkinson (1985: 274) provisionally placed Mastigimas in the Carsidaridae, presumably on larval characters, and Brown (1985) did not dispute this. However the only carsidarid adult character which Mastigimas displays is the enlarged antennal sockets giving the head a cleft appearance. The apical rhinarium is absent from the third flagellomere, there is no rs-m crossvein, no metapostnotal epiphyses, the forewing radular areas are diffuse, one of the metafemoral sensoria is medial, the basal spine of the hind tibia is absent, there are two apical spurs on the hind basitarsus and there are no supplementary lobes on the male subgenital plate. Furthermore the larva lacks lanceolate setae marginally and submarginally on the caudal plate, and the host plants of all known species are species of Cedrela (Meliaceae). For these reasons Mastigimas is excluded from the Carsidaridae but this raises the difficult question of where to place the genus in White & Hodkinson’s system. A sister-group relationship cannot be argued for it with either the Carsidaridae or the Carsidaridae + Homotomidae and I provisionally place Mastigimas near Apsylla which White & Hodkinson tentatively referred to the Calophyidae. The Carsidaridae, as it is recognised here, contains eight genera: Epicarsa, Carsidara, Tyora, Tenapha- lara, Protyora, Paracarsidara, Mesohomotoma and a new genus, Allocarsidara. These are diagnosed below and a key is given for their identification. Nominal species within each genus are discussed in general terms. Insufficient material and incomplete hostplant data prevent thorough revision to species level at present. Key to genera 1 Forewing widening to a broadly rounded apex, Rs and M,,,in broad contact, cu; almost as high as wide and with a value of less than 1-3 (Fig. 18); antennal flagellum thickened, 1st flagellar segment short and broad, not more than four times longer than its greatest width (Fig. 2) EPICARSA (p. 91) — Forewing narrowing to a subacute apex, Rs and M,,>, notin contact but connected by a false rs-m crossvein (Figs 19-39), cu; much wider than high and with a value of at least 1-6; antennal flagellum elongate, 1st flagellar segment long and narrow, not less than nine times longer than itsercatest width: ..:..c10: 4 sek ERS ee Se Se eee eee 2 2 Pterostigma absent and M+Cu about half as long as Cu stem (Figs 36-39); male proctiger bipartite, with a large, anvil-shaped median posterior lobe in addition to lateral lobes (Fig. 77) MESOHOMOTOMA (p. 106) — Pterostigma present (Figs 19-31, 35), if absent (Figs 32-34) then M+ Cu almost twice as long as Cu stem; male proctiger unipartite, without median posterior lobe, with or without lateral lobes (Figs 40, 435468 S157 665 TS) 77 Poe seek SR ae ee ee MALVALES-FEEDING PSYLLIDS 91 3. M+Cu very short, about one-third as long as R stem and less than half as long as Cu stem (Fig. 19); female terminalia, in profile, rounded dorsally, ventrolateral margins of proctiger with dense fringes of setae, lateral palps ridged (Fig. 79) ...............ccccdecececeeeees CARSIDARA (p. 92) — M+Cu longer, about as long as or longer than R stem and Cu stem (Figs 20-35); female terminalia, in profile conical, proctiger sometimes with a median lobe posterior to anal pore, Bee aaa CART TACVE PIGITCC, (PINS SOO) isco ck , M@+Cu stem slightly longer than Cu stem and about as long as M stem, cu; cell value about 1-0 and without radular spinules, apex of claval suture distant from apex of Cu;,; hindwing with grouped costal setae; apical spurs of hind tibia arranged 1+2(or 3)+1. © proctiger unipartite with strongly expanded lateral lobes, each with a secondary inwardly pointing subapical lobe. Q proctiger abruptly stepped in apical fifth, apical part with a dense covering of short, spine-like setae, apex upcurved (see Ferris, 1928: figs 3a—c); subgenital plate short, similarly stepped apically. Larva. Unknown. HostPLanT GENUS. Ceiba (Bombacaceae). ComMENTs. Although possessing most of the features of the family character-suite, the forewing shape, venation and pattern of Epicarsa (Fig. 18) is quite unlike other carsidarids. These forewing characters are similar to many homotomids, the carsidarid sister-group, but the similarity may be convergent. Further characters shared between Epicarsa and homotomids are the thickened antennal flagellum (Fig. 2) and the bifid lateral lobes of the male proctiger. For these reasons I consider Epicarsa to be the sister-group of the rest of the carsidarids. Brown (1985: 259, fig. 98H) described the endtube of the ductus ejaculatorius of this genus as being 92 D. HOLLIS barbed and, from this, suggested a close relationship to the other South American carsidarid genus Paracarsidara (and therefore, by inference, to Mesohomotoma). I have not examined this character in the holotype of E. corniculata and the only other males known (Ferris, 1928) are whole-mount, permanent, microslide preparations in which the structure is obscured. Included species Epicarsa corniculata Crawford (Figs 2, 18) Epicarsa corniculata Crawford, 1911: 488; 1914: 56; 1925b: 56; Ferris, 1928: 244; Costa Lima, 1942: 103; Tuthill, 1950: 58; Hodkinson & White, 1981: 496; Brown, 1985: 260. Holotype C’, BRazit (USNM) [not examined]. Epiciasa corniculata Caldwell, 1941: 419. [Misspelling. ] Epicrasa corniculata Caldwell, 1941: 421. [Misspelling. ] RECORDED DISTRIBUTION. Brazil (Crawford, 1911), Panama (Brown, 1985), Mexico (Ferris, 1928). All other authors refer to these records. MATERIAL EXAMINED 2 0,2 9, Panama, Mexico (slide-mounted material of Ferris (1928) and Brown (1985)). HostTpLantv. Ceiba sp. CARSIDARA Walker Carsidara Walker, 1869: 329; Scott, 1882: 466; Kieffer, 1906: 387, in part; Enderlein, 1910: 138; Aulmann, 1913: 79, in part; Heslop-Harrison, 1960: 241; Hodkinson, 1983: 344; 1986: 303. Type-species: Carsidara marginalis Walker, by monotypy. Dynopsylla Crawford, 1915: 264, in part. Thysanogyna Crawford, 1919: 157; Wu, 1935: 126. Type-species: Dynopsylla minor Crawford, by original designation and monotypy. [Synonymised by Crawford, 1924: 619.] Eustigmia Enderlein, 1921: 119. Type-species: Dynopsylla minor Crawford, by original designation and monotypy. [Junior objective synonym of Thysanogynaa. | DEscRrIPTION. Integument of head and thorax with dense, moderately long setae. Head (Figs 3, 4) with disc of vertex concave, discal foveae clearly marked as broad longitudinal oblique grooves, frontal margin sharply defined and deeply incised by median suture, lateral and hind margins prominent but obtuse, lateral ocelli placed posteriorly on vertex, anterolateral tubercles well-developed; antennal sockets enlarged, flagellum 2-4—2-8 times longer than head width, Ist flagellomere elongate, more than 9 times longer than its greatest width; genae swollen ventrally on either side of median suture anterior to clypeus, each with a prominent papilla, weak lateroventral tubercles also present; apical rostral segment elongate, more than four times longer than its greatest width. Forewing (Fig. 19) ovate with subacute apex, 2-4—2-9 times longer than wide; costal break absent, pterostigma present, 7)-rs crossvein absent, M+Cu stem very short, about one-third as long as R stem and 0-4—0-5 times as long as Cu stem, cu, cell value 2:33—3-75, without radular spinules, apex of claval suture adjacent to apex of Cu,,; hindwing with grouped costal setae; apical spurs of hind tibia arranged Eto CO proctiger (Fig. 40) unipartite, lateral lobes moderately developed; apex of basal segment of aedeagus expanded and strongly sclerotised, apical segment of aedeagus highly modified (Fig. 42), endtube of ductus ejaculatorius heavily sclerotised. Q proctiger (Fig. 79) without posterodorsal lobe, apex strongly sclerotised and upcurved, posterolateral margins with fringes of long setae; lateral palps heavily ridged. Larva. Unknown. Host PLANT GENERA. Brachychiton and Firmiana (Sterculiaceae). ComMENTSs. Carsidara is distinct from the rest of the family and is diagnosed by its broader forewing, the very short M+Cu stem in the forewing, the modified distal segment of the male aedeagus and the structure of the female genital apparatus. I regard the genus as the sister-group of the rest of the family less Epicarsa. MALVALES-FEEDING PSYLLIDS 93 Included species Carsidara marginalis Walker Carsidara marginalis Walker, 1869: 329; Scott, 1882: 467; Aulmann, 1913: 79; Crawford, 1924: 619; Heslop-Harrison, 1960: 237 et seg.; Hodkinson, 1983: 344; 1986: 304. Holotype 9, ‘CELEBES’ (BMNH) [examined]. Dynopsylla minor Crawford, 1915: 263. Holotype ©’, PHILippINEs (7 USNM) [not examined]. [Synony- mised by Crawford, 1924: 619.] Thysanogyna minor (Crawford) Crawford, 1919: 158; Miyatake, 1981: 37. Eustigmia minor (Crawford) Enderlein, 1921: 119. RECORDED DISTRIBUTION. Philippines, Sulawesi, Tanimbar Is. MATERIAL EXAMINED 1 0,9 Q, from Malaya, Hong Kong (on Brachychiton rupestris), Sabah, Sulawesi. HostTPLant. Brachychiton rupestris. ComMMENTS. Crawford (1919) records slight differences in colour and forewing vein proportions in the Tanimbar Is specimens. The Malayan and Hong Kong material studied differs from Walker’s holotype in that there is no brown pattern on the forewing around the radular spinules or apices of M)45,M3440r Cujg, and M72 is less strongly arched basally. The significance of these differences is not apparent and further material with accurate hostplant data is required. Carsidara limbata (Enderlein) Thysanogyna limbata Enderlein, 1926: 397; Wu, 1935: 126. Syntypes 0’, 2, Cuna [not examined]. Carsidara limbata (Enderlein) Hodkinson, 1986: 303. RECORDED DISTRIBUTION. Only known from the type-series, ‘China. Tsingtau’. HostTpLant. Unknown. ComMENTs. Enderlein separated this species from marginalis (as minor) on its larger size (forewing length 5:25-5:75 mm), the brown pattern on the posterior margin of the forewing and some slight venational differences. Carsidara shikokuensis (Miyatake) comb. n., stat. n. Thysanogyna minor skikokuensis Miyatake, 1981: 37. Holotype C’, JAPAN: Kashima Is (OMNH) [not examined]. RECORDED DISTRIBUTION. Only known from the type series. Hoste ants. Firmiana simplex; also found on Mallotus japonicus. ComMENTS. Miyatake’s figures (1981: figs 1A—G), especially of the male genitalia, show this taxon to be quite distinct from marginalis and it is given full species status. However, there is a strong possibility that it is synonymous with limbata. Carsidara africana sp. n. (Figs 3, 4, 19, 40-42, 79) Body ochraceous to mid brown, mesopraescutum with a dark brown anteromedial patch and mesonotum dark brown laterally; antennal flagellomeres 1—6 dark brown apically, 7-8 entirely dark brown; forewing hyaline, with brown pattern as in Fig. 19. Head and dorsum of thorax with moderately long, dense setae. Antennal flagellum 2-7—3-0 times longer than head width, 1st flagellomere more than nine times longer than its median width and with 6—12 rhinaria in apical third, 2nd flagellomere with 10—20 rhinaria in apical half, 3rd flagellomere with 5—7 rhinaria in apical quarter, 4th flagellomere with 2 subapical rhinaria and up to 5 more in apical third, 5th flagellomere with 0 or 1 subapical rhinarium and up to 3 more in apical third, 6th flagellomere with 2 subapical rhinaria, 7th flagellomere with a single subapical rhinarium; ultimate rostral segment more than 6 times longer than wide. Forewing (Fig. 19) 2-47—2-63 times longer than wide and about 1-6 times longer than hindwing; 94 D. HOLLIS pterostigma short, less than half as long as R;; M+ Cu stem about 0-33 times as long as R stem and about 0-5 times as long as Cu stem, cu, value 2:33-3:39. CO genitalia as in Figs 40—42; proctiger about 1-4 times longer than wide; paramere with an outwardly directed, rounded, posteroapical lobe in addition to posteroapical hook. Q terminalia as in Fig. 79. Measurements (3 OC’, 1 Q). Head width, CO 0-95-1-01, Q 1-01; length of antennal flagellum, OC 2:68-2:82, 9 2-81; length of ultimate rostral segment, C’ 0-43-0-45, 9 0-48; length of forewing, 0 4-98-5-71, 9 5-54; length of hind tibia, C’ 1-08-1-24, 9 1-14. LARVA AND HOSTPLANT. Unknown. Holotype 0’, Angola: 7 mls W. Gabela, 16—18.ii1.1972, at light (D. Hollis) (BMNH); dry-mounted. Paratypes. Ghana: 2 9, Northern Region, Garu, 250 m, N 10-51—W 0-12, 27.x.1971, Nr 510 soil trap (Endrody-Younga). Nigeria: 1 ©’, Ile-Ife, 26.11.1970 (J. T. Medler). Uganda: 2 ©’, Kampala, 7.111.1933 (H. Hargreaves). (BMNH; TM); slide- and dry-mounted. ComMENTs. The male and female genitalia of this species are remarkably similar to marginalis but the two species differ markedly in antennal structure, that of marginalis being simple, with a single subapical rhinarium on flagellomeres 2, 3, 4, 6, and 7. Carsidara sp. Carsidara sp. Heslop-Harrison, 1960: 241. A single damaged female of this genus is in BMNH from Madagascar, Nosy Komba. The antennae are missing and I am unable to identify the specimen. PROTYORA Kieffer Protyora Kieffer, 1906: 390; Aulmann, 1913: 29; Tuthill & Taylor, 1955: 250; White & Hodkinson, 1985: 242, 273. Type-species: Tyora sterculiae Froggatt, by monotypy. Neocarsidara Crawford, 1925a: 32; 1927: 31; Tuthill & Taylor, 1955: 250. Type-species: Tyora sterculiae Froggatt, by monotypy. [Junior objective synonym. | DEscrIPTION. Integument of head and thorax almost glabrose. Head (Figs 5, 6) with disc of vertex weakly concave, foveae present as broad longitudinal grooves, hind and lateral margins of vertex rounded, anterior margin not sharply defined but deeply incised by median suture, lateral ocelli placed posteriorly on vertex, anterolateral tubercules absent or present; antennal sockets enlarged, antennal flagellum about 3-7 times longer than head width, 1st flagellomere more than 10 times longer than wide; genal cones present as a pair of small tubercles on either side of mid ventral suture immediately anterior to clypeus, lateroventral tubercles absent; ultimate rostral segment 2:7—3-8 times longer than wide. Forewing (Figs 27, 28) narrow-elongate, 2-9—3-3 times longer than wide, with a narrowly rounded or subacute apex, costal break present, pterostigma present, r;-rs crossvein absent, M+Cu stem almost as long as or longer than R stem and Cu stem, cu, cell value greater than 2-0, radular spinules absent or very weakly present in cu;, apex of claval suture distant from apex of Cu;,; hindwing with grouped costal setae; apical spures of hind tibia arranged 1+2+1. © proctiger (Fig. 43) unipartite, flask-shaped, with poorly developed lateral lobes; apex of aedeagus (Fig. 45) with enlarged apical lobes, endtube of ductus ejaculatorius simple. 2 proctiger (Fig. 80) short, conical in profile, without posterodorsal lobe. Larva. Caudal plate as in Fig. 90. HOsTPLANT GENUS. Brachychiton (Sterculiaceae). CoMMENTS. The two main features of this genus are the presence of a costal break in the forewing and the swollen and modified apex of the aedeagus. The costal break is either primitively retained or secondarily derived and, as I consider its absence as a groundplan feature of the family, I postulate the latter. The form of the aedeagal apex is also regarded as a derived character but not strictly homologous with that of Carsidara. Apart from the simple, flask-shaped male proctiger, Protyora shares other characters, such as the narrow-elongate forewing and long M+Cu stem vein, with the Tyora + Tenaphalara group and the Allocarsidara + Paracarsidara + Mesohomotoma group; it is regarded as the sister-group of these genera. Until now Protyora was thought to be a purely Australian genus, unique in this respect within the Carsidaridae. The discovery of another species in Indonesia shows the genus to have a closer relationship with the Oriental fauna and to be less isolated. _ ane MALVALES-FEEDING PSYLLIDS 95 Tuthill & Taylor (1955) regarded Neocarsidara as a nomen nudum but under Article 12(b)(5) of The International Code of Zoological Nomenclature (3rd edn) the name is valid. However, it has the same type-species as Protyora and is therefore an objective synonym. Included species Protyora sterculiae (Froggatt) (Figs 27, 90) Tyora sterculiae Froggatt, 1901: 289; 1923: 84. Syntypes Oo’, Q and larvae, AusTRALIA (NSW on Brachychiton populneum, QLD on Brachychiton sp.) (ANIC) [not examined]. Protyora sterculiae (Froggatt) Kieffer, 1906: 390; Aulmann, 1913: 29; Tuthill & Taylor, 1955: 250; Morgan, 1984: 37, 68. Neocarsidara sterculiae (Froggatt) Crawford, 1925a: 32; 1927: 31. [Invalid combination, Tuthill & Taylor, 1955: 250.] HostTpLant. Brachychiton populneum (Sterculiaceae). RECORDED DISTRIBUTION. Australia (QLD, NSW, SA). MATERIAL EXAMINED Adults and larvae from Australia (QLD, ACT). CoMMENTS. Brief accounts of the life-history of this species are given by Froggatt (1923) and Morgan (1984). It differs from the new species described below in that the anterolateral tubercles of the vertex are poorly developed, the 1st flagellomere is relatively shorter (10 times longer than its greatest width), the forewing (Fig. 27) is relatively broader (2-9—3-0 times longer than wide) and has a more rounded apex, the pterostigma is broader, M+Cu stem is relatively longer, the male parameres and aedeagal apex are of a different shape and the female proctiger is more obtuse apically (Morgan, 1984). Protyora wilsoni sp. n. (Figs 5, 6, 28, 43-45, 80) DESCRIPTION. Similar to sterculiae. Differs in that antennal flagellum is 3-28—3-75 times longer than head width and has alternating ochraceous and dark brown rings, !st flagellomere longer and narrower, about 15 times longer than its greatest width; anterolateral tubercles of vertex more strongly developed. Forewing (Fig. 28) 3-14—3-25 times longer than wide and with a more acute apex, pterostigma narrow, M-+Cu stem almost as long as R stem and Cu stem, cu; cell value 2-35-3-0. CO paramere and aedeagal apex as in Figs 44, 45. 2 proctiger (Fig. 80) more acute and slightly upcurved apically. Measurements (2 O’, 2 9). Maximum width of head, Oo’ 0-70-0-71, 2 0:74-0:77; length of antennal flagellum, O' 2:51-2:65, 9 2:52-2:64; length of ultimate rostral segment, CO’ 0-20, 9 0-21; length of forewing, 0 3-67-3-70, 9 4:24—4-28; length of hind tibia, CO’ 0-62, 2 0-66. LARVA AND HOSTPLANT. Unknown. Holotype CO’, Indonesia: Sulawesi Utara, Dumoga Bone NP, Hogs Back camp, 21.vi.1985, swept (M. R. Wilson) (BMNH); dry-mounted. Paratypes. 2 O’, 2 2, same data as holotype; 1 Q, 6.vi.1985; 1 Oo’, 1 9, 25.x.1985 (M. Asche); 1 9, 13.x1.—4.xi1.1985, malaise trap (BMNH); 2 oO, x.1985 (Bosmans & Van Stalle) (IRSNB). Slide- and dry-mounted. TYORA Walker Tyora Walker, 1869: 330; Scott, 1882: 470; Kieffer, 1906: 387, in part; Enderlein, 1910: 138; Crawford, 1911: 483; 1920: 355; 1925a: 32; 1927: 30; Aulmann, 1913: 80, in part; Hodkinson, 1983: 366; Yang & Li, 1985b: 206, 215. Type-species: Tyora congrua Walker, by monotypy. Nesiope Kirkaldy, 1908: 389; Aulmann, 1913: 81; Crawford, 1919: 160; Tuthill, 1964: 356; Hodkinson, 1983: 354; 1986: 319. Type-species: Nesiope ornata Kirkaldy, by monotypy. [Synonymised with Carsidaroida by Crawford, 1919: 161, and with Tyora by Crawford, 1920: 355.] Carsidaroida Crawford, 1917: 164. Type-species: Carsidaroida heterocephala Crawtord, by original designation and monotypy. [Synonymised by Crawford, 1920: 355.] 96 D. HOLLIS DESCRIPTION. Integument of head and thorax very sparsely short haired. Head (Figs 7, 8) with disc of vertex flat or weakly concave, foveae present as shallow circular depressions or furrows, lateral and hind margins of vertex sharply raised, anterior margin sharply defined but weakly incised by median suture, lateral ocellae placed posteriorly on vertex, anterolateral tubercles weak or well-developed; antennal sockets enlarged, flagellum 2-3—3-0 times longer than head width, first flagellomere more than 12 times longer than its median width; genal cones present as a pair of small tubercles on either side of mid ventral line immediately anterior to clypeus, weak lateroventral tubercles also present; occiput with a small tubercle on each side below eye; ultimate rostral segment long, more than 4 times longer than wide. Forewing (Figs 20—24) narrow, about 3 times longer than wide, with subacute apex, costal break absent, pterostigma present, 7;-rs crossvein present in addition to rs-m crossvein, M+Cu stem as long as R stem and Cu stem, cu; cell value 1-6 or greater and with radular spinules, apex of claval suture adjacent to apex of Cu,,; hindwing usually with ungrouped costal setae (grouped in T. striata); apical spurs of hind tibia arranged 1+2+2. CO Proctiger (Figs 46, 48) unipartite, with well-developed lateral lobes; apex of aedeagus narrow, endtube of ductus ejaculatorius slightly swollen (Fig. 50). ° proctiger (Figs 81, 82) triangular in profile, usually without posterodorsal lobe (present in T. striata). Larva. Undescribed (see Miyatake (1965)). HOSTPLANT GENERA. Sterculia and Heritiera (Sterculiaceae). ComMENTs. Like Tenaphalara an additional; r)-rs crossvein is present in Tyora and the apical spurs of the hind tibia are arranged 1+2+2. I regard these two characters as autapomorphies for the two genera. Tyora differs from Tenaphalara in the raised margins of the vertex, swollen antennal sockets, presence of radular spinules in cell cu;, close proximity of the apex of the claval suture to the apex of Cu;,, ungrouped costal setae of the hindwing, well-developed lateral lobes of the male proctiger and the absence of a specialised posterodorsal lobe on the female proctiger, although a simple conical lobe is present in T. striata. The two genera are taken to be the sister-group of the Allocarsidara + Paracarsidara + Mesohomotoma complex. Included species Further collecting with accurate hostplant data is required before the validity of all the taxa presently included in the genus can be adequately assessed. 7. ornata and T. striata appear to be good species but it is possible that the remainder represents a single species with highly variable forewing pattern. Tyora buxtoni Crawford (Figs 22, 46, 47) Tyora buxtoni Crawford, 1927: 30; 1928: 34; Hodkinson, 1983: 366; Yang & Li, 1985b: 207, 215. Syntypes, 2 0, SAMOA [1 CO labelled ‘Type’ in BMNH examined]. HostpLantT. Unknown. RECORDED DISTRIBUTION. Samoa. MATERIAL EXAMINED Further specimens with a similar forewing pattern have been examined from various islands in the Solomons group. ComMENTs. Although the forewing pattern (Fig. 22) is consistently different from T. congrua, the male and female genitalia of buxtoni are morphologically identical to that species. Tyora congrua Walker (Figs 7, 8, 20, 21, 50, 81) Tyora congrua Walker, 1869: 330; Scott, 1882: 471; Aulmann, 1913: 80; Crawford, 1920: 355; Hodkinson, 1983: 366; Yang & Li, 19855: 206, 215. Holotype ©’, Mysot Is [not ‘India: Mysore’ as in Yang & Li (19855)] (BMNH) [examined]. Carsidaroida heterocephala Crawford, 1917: 165; 1920: 355. Syntypes Oo’, 9, PHiLipPpINES (USNM) [examined]. [Synonymised by Crawford, 1920: 355.] Nesiope heterocephala var. intermedia Crawford, 1919: 161. Syntypes OC’, 2 TANimBaR Is, JAva [not examined]. MALVALES-FEEDING PSYLLIDS 97 Hostp.ants. Sterculia ureolata, S. lanceolata (Sterculiaceae) (BMNH). RECORDED DISTRIBUTION. Indonesia (Java, Tanimbar Is, Mysol Is), Philippines. MATERIAL EXAMINED Adults from Hong Kong (on Sterculia lanceolata), West Malaysia (Pahang), Indonesia (Java on S. ureolata, Sulawesi Utara, Mysol Is), Philippines, Papua New Guinea (Morobe Prov., New Britain on Sterculia sp.). ComMENTS. Walker’s holotype male is very badly damaged and only part of the thorax, one hind leg, the abdomen and shreds of wing remain. A microslide preparation of the genitalia, made in 1964, has discoloured badly but sufficient remains visible to say that the genitalia are morphologically very similar to those of all other specimens examined. The forewing pattern (Figs 20, 21) varies in extent and intensity in the material studied, being most dense and extensive in specimens from Sulawesi Utara and least so in the Malayan and Hong Kong material. Tyora guandongana Yang & Li Tyora guandongana Yang & Li, 1985b: 212, 215. Holotype O’,CHINA: Guandong (BAUIC) [not examined]. HosteLant. Unknown. COMMENTS. From the original figures it appears that this species is not distinct from T. congrua but, as there is a dark patch of pattern bordering R+M+Cu stem and R stem (Yang & Li, 19855: fig. 8b) which is not present in all material of congrua examined, formal synonymy is not established. Tyora ornata (Kirkaldy) (Figs 23, 48, 49) Nesiope ornata Kirkaldy, 1908: 390; Aulmann, 1913: 81; Laing, 1922: 555; Tuthill, 1964: 357; Miyatake: 1965: 175; Hodkinson, 1983: 354. Syntypes (? sex), Fri [not examined]. Tyora ornata (Kirkaldy) Crawford, 1920: 355; Caldwell, 1942a: 21; Tuthill, 1951: 273; Yang & Li, 1985b: 206, 215. HosteLant. Heritiera littoralis (Sterculiaceae). RECORDED DISTRIBUTION. Ryukyu Is, N. Borneo, S. Marinana Is, Caroline Is, Fiji. MATERIAL EXAMINED Adults from Fiji (on H. littoralis), East Malaysia (Sabah). CoMMENTS. This species is distinguished from T. congrua by the shape of the male paramere and the female proctiger and the pattern of the forewing (Figs 23, 49). Tyora striata (Crawford) (Figs 24, 82) Tenaphalara striata Crawford, 1919: 166; Miyatake, 1972: 13; Hodkinson, 1983: 360. Syntypes OC’, @, SINGAPORE (USNM) [examined]. Tyora striata (Crawford) Yang & Li, 1985b: 206, 215. Tenaphalara sulcata Crawford, 1919: 167, nomen nudum [? in error for striata]. HosteLant. Unknown. RECORDED DISTRIBUTION. Singapore. MATERIAL EXAMINED Adults from Singapore and Brunei. ComMENTs. The presence of a simple posterodorsal lobe on the female proctiger and the grouped costal setae of the hindwing distinguish this species from its congeners. TENAPHALARA Kuwayama Tenaphalara Kuwayama, 1908: 155; Aulmann, 1913: 75; Crawford, 1919: 163, in part; Mathur, 1975: 165; Hodkinson, 1983: 360, in part; 1986: 331, in part; Yang, 1984: 181; White & Hodkinson, 1985: 240, 242; 98 D. HOLLIS Yang & Li, 1985b: 207, 214. Type-species: Tenaphalara acutipennis Kuwayama, by original designation and monotypy. DESCRIPTION. Integument of head and thorax almost glabrose. Head (Figs 9, 10) with disc of vertex convex, foveae present as very shallow circular depressions, hind and lateral margins of vertex rounded, frontal margin not sharply defined but deeply incised by median suture, lateral ocellae placed posteriorly on vertex, anterolateral tubercles absent; antennal sockets slightly enlarged, antennal flagellum about 2-3 times longer than head width, Ist flagellomere more than 8 times longer than its median width; genal tubercles minute, lateroventral tubercles absent; ultimate rostral segment about 5 times longer than wide. Forewing (Figs 25, 26) elongate narrow, with subacute apex, about 3-5 times longer than wide, costal break absent, pterostigma present, 7)-rs crossvein present in addition to rs-m crossvein, M+ Cu slightly longer than R stem and slightly longer than Cu stem, cu; cell value greater than 2-0 and without radular spinules, apex of claval suture distant from apex of Cu;,; hindwing with grouped costal setae; apical spurs of hind tibia arranged 1+2+2. CO’ proctiger (Figs 51, 54) unipartite, with or without lateral lobes; apex of aedeagus narrow, endtube of ductus ejaculatorius simple (Figs 53, 56). Q proctiger (Fig. 83) with a well-developed dorsal lobe posterior to anal pore. Larva. For descriptions of the larval stages of T. acutipennis see Rahman (1932) and Yang (1984). The Sth instar larva of 7. pseudonervosa is described by Mathur (1973). Wax pore ring arrangement of T. camerunus as 1n Fig. 91. HOsTPLANT GENERA. Bombax, Ceiba (Bombacaceae). ComMENTS. Tenaphalara is considered here to be the sister-group of Tyora but has many more derived features than the latter. The margins of the vertex are rounded, the antennal sockets are much less swollen, the genae bear minute mid ventral tubercles, radular spines are absent from cell cu, of the forewing and the apex of the claval suture is distant from the apex of Cu;,, the costal setae of the hindwing are grouped and the female proctiger bears a well-developed and specialised posterodorsal lobe. The genus has been the most widely misunderstood of the family. The confusion stemming from Crawford’s (1919) misinterpretation when he included species belonging to other carsidarid genera, e.g. malayensis (Crawford) (see p. 101), juliana (Crawford) (see p. 102) and striata (Crawford) (see p. 97) and some non-carsidarid species, e.g. fascipennis (Crawford) (see p. 99) and triozipennis (Crawford) (see p. 100). Yang & Li (19855) were aware of this confusion and removed several species from Tenaphalara but recombined only fascipennis and striata with more appropriate genera. Both included and excluded species are listed below with all the latter recombined. Included species Tenaphalara acutipennis Kuwayama (Figs 25, 51-53) Tenaphalara acutipennis Kuwayama, 1908: 156; Crawford, 1911: 491; 1919: 164 [possible misidentifica- tion]; Aulmann, 1913: 75; Ramakrishna Ayyar, 1924: 623; Boselli, 1930: 188; Kuwayama, 1931: 125; Mathur, 1935: 63; 1975: 166; Beeson, 1941: 595; Heslop-Harrison, 1951: 26; Sohi & Sandhu, 1978: 237; Hodkinson, 1983: 360; 1986: 331; Yang, 1984: 181; Yang & Li, 1985b: 206, 214. Syntypes oO’, 2, JAPAN, ? TAIWAN [not examined]. Tenaphalara elongata Crawford, 1912: 432; 1919: 164; Rahman, 1932: 367; Yang & Li, 1985b: 206, 214. Syntypes CO’, 9, Inp1A: on ‘Silk cotton’ [not examined]. [Synonymised by Crawford, 1919: 164.] Ctenophalara elongata Bhatia & Shaffi, 1933: 545 et seq. [Misspelling. ]. HostTpLants. Bombax ceiba (Bombacaceae). Crawford’s (1919) record on Sterculia foetida and BMNH specimens on Hibiscus esculentus are regarded as doubtful host records. RECORDED DISTRIBUTION. India, Taiwan, Japan, Philippines. MATERIAL EXAMINED Adults and larvae from Pakistan (on Bombax ceiba), India, Bangladesh (on Bombax ceiba and Hibiscus esculentus). ComMENTS. Most authors have accepted Crawford’s synonymy of elongata with acutipennis but Yang & Li (19856), basing their judgement on a combination of Kuwayama’s (1908) and Mathur’s (1975) figures, MALVALES-FEEDING PSYLLIDS 99 regarded acutipennis and elongata as separate species. Evidence from the BMNH collections suggests that only one species is involved. Tenaphalara camerunus (Aulmann) comb. n. (Figs 9, 10, 26, 54-56, 83) Carsidara camerunus Aulmann, 19126: 21; 1913: 79. Holotype C&’, CAMEROON (MNHU). HostTpLants. Ceiba pentandra, Bombax buonopozense and B. sessile (Bombacaceae). MATERIAL EXAMINED Adults and larvae from Senegal, Sierra Leone, Ghana, Nigeria, Cameroon, Zaire and Zanzibar. ComMENTS. Although the holotype has not been examined there are several samples of this species in BMNH collections; a study of these and the original description has shown that camerunus should be placed in Tenaphalara. The species is similar to acutipennis but differs in that the apical segment of the aedeagus is relatively shorter (Fig. 56), the male proctiger has weakly developed lateral lobes (Fig. 54), the paramere is of a different shape (Fig. 55) and the female subgenital plate is trilobed apically (Fig. 83). Tenaphalara gossampini Yang & Li Tenaphalara gossampini Yang & Li, 1985b: 208. Holotype ©’, CuH1nA: on Bombax ceiba |as ‘Gossampinus malabarica’| (BAUIC) [not examined]. CoMMENTS. From the original figures (Yang & Li, 19855) of this and the following three species, it is likely that all four are synonyms of T. acutipennis but formal synonymy is not established without reference to type material. No larvae were mentioned in the original data for all these species and it is improbable that Aphanamixis grandifolia (Meliaceae), Mangifera indica (Anacardiaceae) or Dimocarpus longana (Sapin- daceae) are true hostplants. Tenaphalara aphanmixis Yang & Li Tenaphalara aphanmixis Yang & Li, 19856: 209. Holotype CO’, CHINA: on Aphanamixis grandifolia (BAUIC) [not examined]. Tenaphalara dimocarpi Y ang & Li Tenaphalara dimocarpi Yang & Li, 1985b: 211. Holotype 2, Cutna: on Dimocarpus longana (BAUIC) [not examined]. Tenaphalara mangiferae Yang & Li Tenaphalara mangifera Yang & Li, 1985b: 210. Holotype ©’, CHINA: on Mangifera indica (BAUIC) [not examined]. Tenaphalara pseudonervosa (Mathur) comb. n. Nesiope pseudonervosa Mathur, 1973: 71; Hodkinson, 1986: 319. Holotype oO’, INDIA: on Bombax ceiba (FRI) [not examined]. Tyora pseudonervosa (Mathur) Yang & Li, 1985b: 207, 215. Host PLANT. Bombax ceiba (Bombacaceae). COMMENTS. This new combination is made on the basis of the original description; the type material has not been examined. Mathur’s figures show a convex vertex, absence of radular spinules from cell cu; and the characteristically specialised posterodorsal lobe of the female proctiger. Excluded species Strogylocephala fascipennis Crawford Strogylocephala fascipennis Crawford, 1917: 167; Yang & Li, 1985b: 207, 215. Syntypes 0’, 2, PHILIPPINES (USNM) [examined, all damaged]. Tenaphalara fascipennis (Crawford) Crawford, 1919: 165; Miyatake, 1971: 68. 100 D. HOLLIS ComMENTs. Crawford (1919) placed this Philippines species in Tenaphalara and synonymised Strogy- locephala with Tenaphalara. This synonymy was accepted by some subsequent authors (Yu, 1957; Miyayake, 1971) but Yang & Li (1985b) disputed this and regarded Strogylocephala as a valid genus. Yu (1957) described Tenaphalara confluens from Taiwan, on Palaquium formosanum (Sapotaceae), and suggested it was closely related to fascipennis. Yang & Li (1985b) placed confluens also in Strogylocephala. Chung Tu Yang (1984) redescribed confluens and placed it in a new genus Synaphalara, in the subfamily Carsidarinae. After examining Crawford’s type material of fascipennis and recently collected specimens of the same species from Sulawesi Utara, on Palaquium sp. I agree with Yang & Li (1985b) that Strogylocephala is a distinct genus. It is quite unrelated to the Carsidaridae and I tentatively place it in the Calophyidae. Furthermore I consider Synaphalara to be a junior synonym of Strogylocephala (syn. n.) and it is likely that confluens and fascipennis are synonyms. Placing Strogylocephala in the Calophyidae conflicts with published data on the larva of the type-species which was clearly described as a triozid larva by Chung Tu Yang (1984: 186). However, when the Sulawesi specimens of fascipennis were collected, larvae, which agree generally with Yang’s description, and a single adult triozid were taken at the same time. This triozid is an undescribed Trioza species belonging to the Sapotaceae/Ebenaceae-feeding obsoleta-group (Hollis, 1984). It is therefore possible that the larvae described by Yang belong to this triozid species and are not associated with the Strogylocephala species. Strogylocephala confluens (Yu) Tenaphalara confluens Yu, 1956: 46. Holotype C’, TAIwan [not examined]. Synaphalara confluens (Yu) Chung Tu Yang, 1984: 185. Strogylocephala confluens (Yu) Yang & Li, 1985b: 207, 216. COMMENTS. See under S. fascipennis. Allocarsidara juliana (Crawford) comb. n.; see below (p. 102) Allocarsidara malayensis (Crawford) comb. n.; see below (p. 101) Tenaphalara sulcata Crawford, 1919: 167, nomen nudum; see above (p. 97) Trioza triozipennis (Crawford) comb. n. Tenaphalara triozipennis Crawford, 1919: 167. Holotype ?sex [abdomen missing], SINGAPORE (USNM) [examined]. ? Tenaphalara triozipennis Crawford; Yang & Li, 19855: 207, 216. ComMENTS. The holotype is clearly a triozid, probably related to T. umalii (Miyatake) (see below), and is provisionally placed in Trioza. Trioza umalii (Miyatake) comb. n. Tenaphalara umalii Miyatake, 1972: 11. Holotype 9, PHILIPPINES (OMNH) [not examined]. ? Tenaphalara umalii Miyatake; Yang & Li, 1985b: 207, 216. CoMMENTS. From Miyatake’s description there can be little doubt that this species is a triozid and is probably related to triozipennis (Crawford). The above new combination is therefore proposed. ALLOCARSIDARA gen. n. [Tenaphalara Crawford, 1919: 163, in part; Hodkinson, 1983: 360, in part; 1986: 331, in part; Braza & Calilung, 1981: 354. Misinterpretations. | Type-species: Tenaphalara malayensis Crawford. DEscRIPTION. Integument of head and thorax almost glabrous. Head (Figs 11, 12) with disc of vertex flat or convex, foveae weak, lateral and hind margins of vertex rounded, anterior margin not clearly defined and weakly incised by median suture, lateral ocelli placed medially on vertex, anterolateral tubercles well-developed; antennal sockets enlarged, flagellum 3-26—4-60 times longer than head width, first flagellomere more than 10 times longer than wide; genal cones present as a pair of very weakly developed MALVALES-FEEDING PSYLLIDS 101 humps on either side of mid ventral line anterior to clypeus, genae also with an outer pair of lateroventral tubercles, occiput with a small tubercle on each side below eye; ultimate rostral segment more than four times longer than wide. Forewing (Figs 29-34) narrow elongate, at least 2-8 times longer than wide and with an acute apex, costal break absent, pterostigma present or absent, r)-rs crossvein absent, M+Cu stem as long as R stem and more than 1-7 times longer than Cu stem, cu, cell value 1-76—2-70 and without radular spinules, apex of claval suture distant from apex of Cu,,; hindwing with grouped costal setae; apical spurs of hind tibia arranged 1+2+1. CO proctiger (Figs 57, 60, 63, 66, 69, 72) unipartite, with well-developed lateral lobes that are sometimes subdivided posteroapically, each inner lateral surface with a patch of differentiated spinules; apical segment of aedeagus straight or flexed, aedeagus narrow apically with endtube of ductus ejaculatorius contained within apical lobes (Figs 59, 62, 65, 68, 71, 74). Q proctiger (Figs 84—87) with or without a posterodorsal lobe. Larva. As in Fig. 92. HosTPLANT GENUS. Durio (Bombacaceae). ComMMENTS. The sister-group relationship of Allocarsidara needs to be resolved. The genus is placed with Paracarsidara and Mesohomotoma on the structure of the male proctiger but it differs from these genera in having rounded lateral and posterior margins of the vertex, well-developed anterolateral tubercles on the vertex, a simple endtube of the ductus ejaculatorius that is enclosed within the apical lobes of the aedeagus, and a patch of differentiated spinules on each inner lateral surface of the male proctiger. Some species have lost the pterostigma of the forewing as in Mesohomotoma. Yang & Li (19855) expressed doubt that Tenaphalara malayensis Crawford and T. juliana Crawford belonged in Tenaphalara and this suggestion is confirmed here. Crawford (1919) misinterpreted the genus Tenaphalara (see above, p. 98) and his two species malayensis and juliana are transferred to Allocarsi- dara. A further four new species are described below. Relationships among these species are not at all clear. A variety of characters appear in the group, e.g. the absence of a pterostigma, an apically bifid male paramere, a flexed distal aedeagal segment, the male proctiger with subdivided lateral lobes, and the female proctiger with a posterodorsal lobe. However, the distribution of these characters within the genus (Table 1) gives little indication of sister-group rela- tionships. The six species included in Allocarsidara may be separated from one another by using the character-matrix in Table 1; three are known to develop on Durio zibethinus. Table 1 Allocarsidara spp., character-matrix. Primitive condition given in parenthesis. Posterodorsal lobe of Q apical segment lateral lobes pterostigma proctiger CO’ paramere of aedeagus of CO proctiger absent present truncate flexed divided Character (present) (absent) (conical) (straight) (entire) Species malayensis (-) + le (5) (=) iriana ) ? + —) (=) incognita ) =" ) + += bakeri i= 2 () + (—) juliana + (-) + t b=) elongata a5 st te) SS) ) Included species Allocarsidara malayensis (Crawford) comb. n. (Figs 29, 57-59, 86) Tenaphalara malayensis Crawford, 1919: 165; 1928: 425; Laing, 1930: 39; Hodkinson, 1983: 360; 1986: 332. Syntypes, 0’, 9, MALaya (Penang) and SINGAPoRE (USNM) [examined]. ? Tenaphalara malayensis Crawford; Yang & Li, 1985b: 207, 216. HosteLant. Durio zibethinus (Bombacaceae). 102 D. HOLLIS RECORDED DISTRIBUTION. Malaya, Singapore, Sumatra, Java. Braza & Calilung (1981: 355) record this species from the Philippines but their material represents a different species (see below). MATERIAL EXAMINED Adults and larvae from Thailand, Malaya and Java. Allocarsidara juliana (Crawford) comb. n. (Figs 32, 66-68, 84) Tenaphalara juliana Crawford, 1919: 166; 1920: 355; Hodkinson, 1983: 360. Holotype, ? sex, ‘BORNEO’ (BPBM) [examined]. ? Tenaphalara juliana Crawford; Yang & Li, 1985b: 207, 216. LARVA AND HOSTPLANT. Unknown. RECORDED DISTRIBUTION. Sabah. MATERIAL EXAMINED Holotype (? sex), East Malaysia: Sabah (‘Sandakan, Borneo, Baker’); 1 0’, 1 9, same data as holotype (USNM); 2 9, Sarawak, offshore gas platform (BMNH). ComMMENTS. This species was described from a single specimen of unknown sex (the posterior half of the abdomen was missing) from Sabah, and further recorded (Crawford, 1920) from a single female from an unstated locality but presumably in Borneo as the paper in which the specimen is recorded deals with Bornean psyllids. Apart from the holotype (BPBM), there are three specimens bearing the type data, under the name ‘Tenaphalara juliana’, in USNM; two complete males and a complete female. The two males are not conspecific with each other and neither can be positively identified as the same species as the damaged holotype of juliana. The complete female agrees well with the holotype and I assume that it is the specimen Crawford recorded as Tenaphalara juliana in 1920. As well as the locality label this specimen also bears a handwritten label ‘11734’, which is almost certainly an original Baker collection label. One of the two males also bears a similar label and, therefore, I regard it as having been collected by Baker at the same time as the female. I consider this specimen to be the male of Allocarsidara juliana and figure it below (Figs 32, 66-68). The other male in USNM is described below as A. bakeri sp. n. (p. 103). Allocarsidara juliana differs from malayensis in lacking a pterostigma in the forewing (Fig. 32), lacking a posterodorsal lobe on the female proctiger (Fig. 84) and having a flexed apical aedeagal segment in the male (Fig. 68). It is apparently closest to bakeri (see below) but differs from that species in the shape of the paramere (Figs 67, 70). Allocarsidara incognita sp. n. (Figs 30, 72-74, 85, 92) [ Tenaphalara malayensis Crawford; Braza & Calilung, 1981: 355. Misidentification. | DESCRIPTION (only slide-mounted and spirit-stored material available for study). Antennal flagellum 3-83—4-30 times longer than head width, 1st flagellomere more than 13 times longer than wide; ultimate rostral segment 1-52—2-17 times longer than flagellomeres 7+8 together. Forewing (Fig. 30) 2-99—3-18 times longer than wide, pterostigma present, Cu stem 3 times longer than Cujp, Cu; cell value 1:83-2:62. CO genitalia as in Figs 72—74; lateral lobes of proctiger each with a supplementary apicoposterior lobe; paramere narrow apically and with an inwardly directed spine, apical segment of aedeagus flexed. ° terminalia as in Fig. 85, proctiger with a well-developed posterodorsal lobe. Measurements (7 0’, 7 9). Maximum width of head, Oo 0-67—-0-71, 2 0-71—-0-74; length of antennal flagellum, O 2:67-2:90, 9 2-74-3-06; length of ultimate rostral segment. OC’ 0-26—0-30, 9 0-30—0-31; length of forewing, C’ 3-32—3-53, 9 3-65—3-89; length of hind tibia, O’ 0-68—0-76, 9 0-71-0-74. Larva. Abdominal wax pore rings as in Fig. 92. HostTPLant. Adults and larvae collected from Durio zibethinus (Bombacaceae). MALVALES-FEEDING PSYLLIDS 103 Holotype C’, Indonesia: Sulawesi Utara, Dumoga Bone NP, Bungalows light trap, 21.xi.1985 (RESL Project Wallace) (BMNH); slide-mounted. Paratypes. Indonesia: 1 2, same locality data as holotype, Base Camp light trap, 26.x.1985. Philippines: 1 9, Davao Exp. Stn, ground level trap, xi.1962 (M. R. Gavarra);2 0,2 9, larvae, Davao City, 2.x.1975, Durio zibethinus (M. Gavarra). East Malaysia: 5 CO’, 4 92, Sarawak, vi.1963, Maize (C. R. Wallace) (BMNH). CoMMENTS. This species is very similar to malayensis but can be recognised by the structure of the male genitalia; females of the two species are very difficult to separate from one another. Allocarsidara iriana sp. n. (Figs 11-13, 31, 63-65) DEscrIPTION. Body mid brown dorsally and laterally, ochraceous ventrally; head black but with a median longitudinal pale stripe on vertex, apices of anterolateral tubercles ochraceous; antenna with scape and pedicel ochraceous, flagellum black; forewing hyaline; femora ochraceous, brown anterodorsally, fore and mid tibiae and tarsi dark brown, hind tibia and tarsus ochraceous. Antennal flagellum relatively short, 3-26 times longer than head width, first flagellomere 10 times longer than its greatest width, seventh flagellomere with a very long subapical seta (Fig. 13); ultimate rostral segment 1-46 times longer than flagellomeres 7+8 together. Forewing (Fig. 31) 2-86 times longer than wide, pterostigma present, M+Cu stem just over twice as long as Cu stem, Cu stem twice as long as Cu), cu, value 2-13. CO genitalia as in Figs 63-65; lateral lobes of proctiger short, not subdivided posteroapically; paramere broad apically, with an anteriorly directed spine and an apicoposterior tubercle; apical segment of aedeagus straight. Q unknown. Measurements (1 OC’). Maximum width of head, 0-58; length of antennal flagellum, 1-88; length of ultimate rostral segment, 0-23; length of forewing, 3-00; length of hind tibia, 0-76. LARVA AND HOSPLANT. Unknown. Holotype Oo’, New Guinea: NW, Nabire, S. Goelvink Bay, 7.ix.1962, light trap in jungle (H. Holtman) (BPBM); slide-mounted. ComMENTS. The forewing of this species has a pterostigma, as in malayensis and incognita, but it may be recognised by the shape of the male paramere and aedeagus, the relatively short antennal flagellum and the long subapical seta on flagellomere seven. Allocarsidara bakeri sp. n. (Figs 33, 69-71) DESCRIPTION. Body light brown, head darker; antenna with scape, pedicel and first flagellomere dark brown, rest of flagellum lighter; forewing hyaline; legs ochraceous, fore and mid femora with dark anterodorsal stripes, mid tibia and tarsus darker. Antennal flagellum 4-66 times longer than head width, first flagellomere 12 times longer than its greatest width. Forewing (Fig. 33) 3-45 times longer than wide, pterostigma absent, Cu stem 3-5 times longer than Cup, cu, cell value 2:7. O genitalia as in Figs 69-71; lateral lobes of proctiger long but not subdivided posteroapically; apex of paramere narrow and with an inwardly directed curved spine; apical segment of aedeagus flexed. 2 unknown. Measurements (1 ©’). Maximum width of head, 0-59; length of antennal flagellum, 2-77; length of ultimate rostral segment, 0-24; length of forewing, 3-12; length of hind tibia, 0-57. LARVA AND HOSTPLANT. Unknown. Holotype ©’, East Malaysia: Sabah; ‘Sandakan, Borneo, Baker’ (USNM); slide-mounted. ComMENTS. This species is very similar to juliana but differs in that the lateral lobes of the male proctiger are longer (Figs 66, 69), the paramere is narrow apically (Figs 67, 70), the apex of the aedeagus is of a different shape (Figs 68, 71) and the forewing is relatively narrower (3-10 times longer than wide in juliana). 104 D. HOLLIS Allocarsidara elongata sp. n. (Figs 34, 60-62, 87) DESCRIPTION (only slide-mounted and spirit-preserved material available for study). Antennal flagellum 3-98—4-60 times longer than head width, first flagellomere 16 times longer than its greatest width; ultimate rostral segment relatively long, 2-54—2-89 times longer than flagellomeres 7+8 together. Forewing (Fig. 34) 3-21—3-35 times longer than wide, pterostigma absent, Cu stem about 4 times longer than Cuz), cu; cell very small. © genitalia as in Figs 60-62; lateral lobes of proctiger undivided posteroapically; apex of paramere narrow, with a large and inwardly directed apical tubercle; apical segment of aedeagus straight. Q terminalia as in Fig. 87, proctiger with a well-developed posterodorsal lobe. Measurements (6 0’, 7 9). Maximum width of head, 0” 0-64—-0-73, 2 0-69-0-75; length of antennal flagellum, C 2-83-3-00, 2 3-11—3-26; length of ultimate rostral segment, CO’ and 9 0-31—0-33; length of forewing, O 3-47-3-59, 9 3-81—4-14; length of hind tibia, 0’ 0- 69-0-73, 9 0-71-0-74. HostTpLant. Adults and larvae collected from Durio zibethinus (Bombacaceae). Holotype o', West Malaysia: Malaya, Alor Star, Jitra, 3.xii.1985, Durio zibethinus (N. K. Ho) (BMNH); slide-mounted. Paratypes. 7 0’, 9 2, same data and depository as holotype; slide- and dry-mounted. CoMMENTS. This species resembles juliana and bakeri in lacking a pterostigma in the forewing; it differs from both in having a straight apical aedeagal segment, and from juliana in having an apically narrow paramere and a well-developed posterodorsal lobe on the female proctiger. PARACARSIDARA Heslop-Harrison [Carsidara Walker; Crawford, 1911: 484; 1914: 57; Tuthill & Taylor, 1950: 53; Brues et al., 1954: 156; Morgan, 1984: 19, fig. 15. Misinterpretations. ] Paracarsidara Heslop-Harrison, 1960: 244; Hodkinson & White, 1981: 505; Brown, 1985; 246. Type- species: Carsidara dugesii Low, by original designation. DESCRIPTION. Integument of head and thorax almost glabrose. Head (Figs 14, 15) with disc of vertex concave, foveae present as deep oblique grooves, lateral margins of vertex sharply raised, posterior margin angular, anterior margin sharply defined and deeply incised by median suture, lateral ocellae placed posteriorly on vertex, anterolateral tubercles very weak or absent; antennal sockets swollen, flagellum 2:5—3:-5 times longer than head width, 1st flagellomere more than 10 times longer than its greatest width; genae with a pair of small mid ventral tubercles anterior to clypeus, each gena with a larger lateroventral tubercle; occiput, on each side below, eye with a small tubercle or swelling; ultimate rostral segment more than 6 times longer than wide. Forewing (Fig. 35) narrow-elongate, about 2-8 times longer than wide and with a subacute apex, costal break absent, pterostigma present, 7;-rs crossvein absent, M+ Cu stem almost as long as R stem and Cu stem, cu, cell value greater than 2-7 and without radular spinules, apex of claval suture distant from apex of Cuz», costal setae of hindwing grouped; apical spurs of hind tibia arranged 1+3+1. CO proctiger (Fig. 75) unipartite, expanded basally and with 2 small apicolateral lobes; aedeagus narrow apically, sometimes lobed basoventrally, endtube of ductus ejaculatorius enlarged and with lateral and apical hooks (Fig. 76). @ terminalia (Fig. 88) conical, proctiger more or less ‘stepped’ posteriorly, apical part narrow and bearing short, thick, spiniform setae and sometimes an apical hook. Larva. Caudal plate as in Fig. 93. HOSTPLANT GENERA. Ceiba (Bombacaceae), Malva and Wissadula (Malvaceae). CoMMENTS. On the basis of the narrow-elongate forewing, absence of r)-rs crossvein, long M+Cu stem, presence of lateral lobes on the male proctiger and narrow apex of the aedeagus, Paracarsidara is grouped with Allocarsidara and Mesohomotoma. The modified endtube of Paracarsidara is very similar to that of Mesohomotoma and is used as a synapomorphy for these two genera. Brown (1985) also related Paracarsidara to Mesohomotoma, even doubting their separate identities, but I regard the absence of a pterostigma and the bipartite male proctiger that bears a median posterior lobe in addition to the lateral lobes sufficient to recognise the latter genus. MALVALES-FEEDING PSYLLIDS 105 Included species Paracarsidara dugesii (L6w) (Figs 75, 76, 88, 93) Carsidara dugesii Low, 1886: 160; Crawford, 1911: 486; 1914: 58; Aulmann, 1913: 79; Laing, 1923: 698; Caldwell, 1941: 421, 1944: 58; Tuthill, 1950: 53; Silva et al. , 1968: 200 [?misidentification]. Syntypes O’, @, Mexico: ‘planta Malvacea’ (NM) [not examined]. Paracarsidara dugesii (L6w) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 1985: 250. Carsidara concolor Crawford, 1911: 484; 1914: 58; Caldwell, 1942b: 28; Caldwell & Martorell, 1951: 604. Syntypes CO’, 2, CuBa (USNM). [Synonymised by Heslop-Harrison, 1960: 246. | Paracarsidara concolor (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 505. [ Carsidara gigantea Crawford; Laing, 1923: 698. Misidentification. | Hoste ants. Malva sp., Wissadula perplocifolia (Malvaceae). Silva et al. (1968) record this species from Brazil, on Bombax cyathophorum (Bombacaceae), but this is likely to be a misidentification. RECORDED DISTRIBUTION. Mexico, Guatemala, Cuba, Puerto Rico, Dominica and Jamaica; [?Brazil]. MATERIAL EXAMINED Adults and larvae from Mexico and Jamaica (on Wissadula perplocifolia). Paracarsidara gigantea (Crawford) (Figs 14, 15, 35) Carsidara gigantea Crawford, 1911: 486; 1914: 57; Ferris, 1928: 244; Caldwell, 1941: 421. Syntypes 9°, NICARAGUA (USNM). Paracarsidara gigantea (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 1985;253. HostTp ants. Ceiba sp., C. burchellii (BMNH). RECORDED DISTRIBUTION. Mexico, Guatemala, Belize, Nicaragua, St Kitts, Cuba, Colombia, Brazil. MATERIAL EXAMINED Adults and larvae from Mexico, Panama and Brazil (on Ceiba burchellii). ComMENTS. Tuthill (1950) synonymised this species with dugesii but Brown (1985) considered it to be distinct after examining relevant type material. Paracarsidara rostrata (Crawford) Carsidara rostrata Crawford, 1911: 486; 1914: 58; Tuthill, 1950: 53. Syntypes 2, NicAaRAGUA (USNM). Paracarsidara rostrata (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 1985: 256. HostTeLant. Unknown. RECORDED DISTRIBUTION. Nicaragua, Panama. ComMENTSs. Crawford (1914) regarded this species as a synonym of dugesii and this was followed by Tuthill (1950). Heslop-Harrison raised rostrata from synonymy and Brown (1985) confirmed this after examining type material. Nomen dubium Carsidara mexicana Crawford, 1911: 487; 1914: 58 (as a possible synonym of dugesii). Holotype CO’, MExIco [specimen missing from pin, USNM]. Paracarsidara mexicana (Crawford); Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 248 (as a nomen dubium). Undescribed species Brown (1985) recorded two undescribed species from Panama and there are some poorly preserved specimens of a further undescribed species from Trinidad on Malachra alceifolia (Malvaceae) in BMNH. 106 D. HOLLIS MESOHOMOTOMA Kuwayama Mesohomotoma Kuwayama, 1908: 180; Aulmann, 1913: 36; Crawford, 1920: 356; 1925a: 32; Tuthill & Taylor, 1955: 249; Miyatake, 1971: 58; Mathur, 1975: 144; Braza & Calilung, 1981: 356; Hodkinson, 1983: 352; 1986: 317; Yang, 1984: 178; White & Hodkinson, 1985: 242. Type-species: Mesohomotoma camphorae Kuwayama, by original designation and monotypy. Udamostigma Enderlein, 1910: 138. Type-species: Tyora hibisci Froggatt, by original designation and monotypy. [Synonymised by Crawford, 1920: 356. ] [Tyora Walker; Crawford, 1919: 159. Misinterpretation. | [ Tyor[iJ]a Walker; Uichanco, 1921: 279. Misinterpretation; misspelling. | DESCRIPTION. Integument of head and thorax almost glabrous. Head (Figs 16, 17) with disc of vertex deeply divided by median suture, foveae present as deep oblique grooves, lateral and hind margins of vertex obliquely raised, anterior margin poorly defined but deeply incised by median suture, anterolateral tubercles well-defined, lateral ocellae posteriorly placed; antennal sockets enlarged, flagellum 2-4—3-0 times longer than head width, 1st flagellomere at least 9 times longer than its median width; genae with a small tubercle on either side of mid ventral suture immediately anterior to clypeus, lateral tubercles absent; occiput with a small tubercle on each side below eye; ultimate rostral segment at least 5 times longer than wide. Forewing (Figs 36-39) narrow, elongate, 2-5—3-0 times longer than wide, with subacute apex; costal break, pterostigma and r,-rs absent, M+ Cu stem short, about two-fifths as long as R stem and half as long as Cu stem, cu, cell value almost 2-0 and without radular spinules, Cu ;, strongly arched towards M stem, apex of claval suture distant from apex of Cu ,; costal setae of hindwing grouped; apical spurs of hind tibia arranged 1+3+1. CO proctiger (Fig. 77) bipartite, a strong anvil-shaped median posterior lobe present in addition to well-developed lateral lobes; aedeagus (Fig. 78) narrow apically, apex of ductus ejaculatorius prominent, strongly produced from aedeagal apex and expanded apically. 2 proctiger (Fig. 89), in profile, strongly stepped posteriorly, apical part narrow elongate and bearing short and thickened setae, apex weakly barbed. LarvA. Caudal plate as in Fig. 94. Larval stages of M. camphorae described by Yang (1984). HOSTPLANT GENERA. Hibiscus, Thespesia, Abutilon, Urena (Malvaceae); Cola, Theobroma, Sterculia, Heritiera (Sterculiaceae). ComMENTS. Mesohomotoma differs from all other carsidarids in having a bipartite male proctiger that bears a median posterior lobe in addition to the lateral lobes. Like Carsidara the M+Cu stem is short but this is probably a primitive feature; the pterostigma is absent, resembling some species of Allocarsidara but this is likely to be a parallel development in the two genera; the form of the apical segment of the aedeagus and particularly the endtube of the ductus ejaculatorius is similar to Paracarsidara and this is considered as a synapomorphy for the two genera. Included species A great deal of confusion exists over the identity and validity of the six species currently recognised in this genus. I have not examined original type-material but have seen a large number of specimens from most of the areas from where these species were described. There is considerable variation in size, body coloration and forewing pattern and vein proportions (Figs 36-39) between, but not within, samples, although this variation does not correlate well with hostplant data or geographical distribution. Male and female genitalia and larval morphology are, however, remarkably consistent throughout the geographical and hostplant ranges of the material. The most commonly encountered species is M. hibisci and its hostplant, Hibiscus tiliaceus, is common in strandline and coastal vegetation throughout the Pacific and Indian Ocean regions from Polynesia to the East and South African coast. M. camphorae and M. lineaticollis, both originally described from Taiwan, have been subsequently recorded on Hibiscus tiliaceus and other malvaceous hosts such as Urena lobata and A butilon indicum, in South East Asia. M. lutheri, described from Sri Lanka, is recorded from Hibiscus tiliaceus and Urena lobata in India, Sri Lanka, Réunion and Madagascar. M. africana was described from Natal without host data but there are Mesohomotoma samples in BMNH collected from Hibiscus in eastern Tanzania, Kenya, Uganda and Zaire; M. tessmanni is widespread in Central and West Africa on Cola, Theobroma (cocoa) and other sterculiaceous hosts. I suspect that camphorae, lineaticollis, lutheri and africana are synonyms of hibisci. Furthermore, there appear no consistent morphological characters, either in the adult or larva, that will separate tessmanni MALVALES-FEEDING PSYLLIDS 107 from hibisci but the sterculiaceous hostplant preferences of the West and Central African populations suggest some genetic isolation from the Malvaceae-feeding populations. Hibiscus tiliaceus does occur around the coastal areas of West Africa but it is not known if these bushes support populations of Mesohomotoma. Further collecting here would be useful. Experimental work involving hostplant transfers and crossbreeding techniques is required to clarify the problem. Mesohomotoma hibisci (Froggatt) (Figs 36, 37, 77, 78, 89, 94) Tyora hibisci Froggatt, 1901: 287. Syntypes CO’, 2, AUSTRALIA: Queensland, on Hibiscus tiliaceus (ANIC) [not examined]. Udamostigma hibisci (Froggatt) Enderlein, 1910: 138; Aulmann, 1913: 81. Mesohomotoma hibisci (Froggatt) Crawford, 1920: 356; 1925a: 34; 1927: 30; Klyver, 1933: 25; Caldwell, 1942: 21; Tuthill, 1943: 71; 1951: 273; 1964: 355; Tuthill & Taylor, 1955: 250; Hodkinson, 1983: 353; 1986: 317; Morgan, 1984: 29. HostTeLants. Hibiscus tiliaceus, H. rosasinensis, H. boryanus (Malvaceae). RECORDED DISTRIBUTION. Polynesia (Society, Austral and Rapa Is), Melanesia (Fiji and Solomon Is, New Caledonia), Micronesia (Marianas and Caroline Is), Australia (Queensland). MATERIAL EXAMINED Numerous adults and larvae from: Polynesia (Cook, Society, Tonga, Samoa and Ellice Is), Melanesia (Fiji, Loyalty, New Caledonia, Solomons, Bismark Is and Papua New Guinea), Micronesia (Caroline, Marianas and Gilbert Is), Australia (N.S.W.), Indonesia (Sulawesi), Philippines, Hong Kong, India, Chagos Arch., Seychelles, Réunion, Mauritius, Madagascar, Zaire, Uganda, Kenya, Tanzania, Zimbabwe, South Africa (Natal). Mesohomotoma africana Pettey Mesohomotoma africana Pettey, 1924: 29; 1925: 138; Capener, 1970: 199. Syntypes C’, 9, SouTH AFRICA (Natal) (NCI; BMNH) [1 9 examined]. Hostevant. Hibiscus sp. (Malvaceae). RECORDED DISTRIBUTION. South Africa (Natal). Mesohomotoma camphorae Kuwayama Mesohomotoma camphorae Kuwayama, 1908: 181; Aulmann, 1913: 36; Crawford, 1925a: 33; 1927: 31; 1928: 34; Kuwayama, 1931: 123; Takahashi, 1936: 292; Sasaki, 1954: 32; Miyatake, 1964: 123; 1965: 174; 1971: 58; Hodkinson, 1983: 352; 1986: 317; Yang, 1984: 178. Syntypes Oo’, 2, JAPAN (Osagawara Is) and TAIWAN, on ‘Kampferbaume’ [not examined]. HosteLants. Hibiscus tiliaceus, H. mutabilis, Abutilon indicum, Thespesia populnea, Urena lobata (Malvaceae); Cinnamomum camphora (Lauraceae) but Sasaki (1954) regards the camphor records as erroneous. RECORDED DISTRIBUTION. Samoa, Fiji, Philippines, Japan (Osagawara and Ryukyu Is), Taiwan. Mesohomotoma lineaticollis Enderlein Mesohomotoma lineaticollis Enderlein, 1914: 232; Crawford, 1925a: 35; Boselli, 1930: 188; Kuwayama, 1931: 123; Braza & Calilung, 1981: 356; Hodkinson, 1983: 353. Holotype OC’, ‘Formosa’ [not examined]. HostTpLant. Urena lobata (Malvaceae). Also recorded (probably in error) on Cordia dichotoma (Boragi- naceae). RECORDED DISTRIBUTION. Taiwan, Philippines. Mesohomotoma lutheri (Enderlein) Udamostigma lutheri Enderlein, 1918: 484. Syntypes CO’, 2, Skt LANKA [not examined]. Tyora indica Crawford, 1919: 159. Syntypes 0’, 2, AmBorna and INp1A [not examined]. [Synonymised by Crawford, 192S5a: 34.] 108 D. HOLLIS Tyor[i]a indica Crawford; Uichanco, 1921: 279. [Misspelling. ] Mesohomotoma lutheri (Enderlein) Crawford, 1925a: 34; 1927: 31; Mathur, 1975: 146; Hodkinson, 1983: 353819862 317: Mesohom[a]toma lutheri (Enderlein); Orian, 1972: 2. [Misspelling. | HostTpiants. Hibiscus sp., Urena lobata (Malvaceae). RECORDED DISTRIBUTION. Amboina, Philippines, India, Ceylon, Réunion, Madagascar. Mesohomotoma tessmanni (Aulmann) (Figs 16, 17, 38, 39) Udamostigma tessmanni Aulmann, 1912a: 10; 19126: 101; 1913: 81. Syntypes ©’, ‘SPANISH GUINEA’ [not examined]. Mesohomotoma (Udamostigma) tessmanni (Aulmann) Cotterell, 1927: 109. Mesohomotoma tessmanni (Aulmann); Alibert, 1951: 44; Eastop, 1961: 167; Forsyth, 1966: 73; Leston & Gibbs, 1968: 73; Roberts, 1969: 76; Leston, 1973: 322. Tyora tessmanni (Aulmann) Eastop, 1958: 19; Kaufmann, 1973: 285. Tyora (Mesohomotoma) tessmanni (Aulmann); Wood, 1980: 174. HostTpLants. Cola spp., Theobroma cacao, Sterculia tragacantha (see Forsyth, 1966), Heritiera spp. (BMNH) (Sterculiaceae); Desplatzia lutea (BMNH) (Tiliaceae); Chytranthus talbotii (see Forsyth, 1966) (Sapindaceae); Cleistopholis sp. (BMNH) (Annonaceae) (possible host misidentification). RECORDED DISTRIBUTION. Ghana, Nigeria, Equatorial Guinea. MATERIAL EXAMINED Numerous adults and larvae from Sierra Leone (on Heritiera, Cola, Theobroma), Ivory Coast (on Heritiera), Ghana (on Theobroma, Desplatzia), Nigeria (on Cola, Theobroma), San Tomé (on Theobroma), Cameroon (on Cola, Theobro- ma), Central African Republic (on Cola), Gabon (on Cleistopholis), Uganda (on Cola), Zaire and Angola. CoMMENTS. Cotterell (1927) records this species as a minor pest of cocoa (Theobroma cacao) in West Africa, damaging young shoots and shortening internodes, which causes ‘bunching’. Wood (1980) summarises that, under conditions of drought and high insolation, heavy infestations cause dessication and death of terminal buds. However, Kaufmann (1973) reports that when the insects infest the flower buds there is little effect on pod production and the insects may be involved in flower pollination. Phylogeny, biogeography and hostplants (Summary Fig. 1) The Carsidaridae and Homotomidae are considered here to be sister-groups, the pair being characterised as follows: antennal sockets enlarged and giving the head a cleft appearance in dorsal view; paired metapostnotal epiphyses present; ventral sense organs of hind femora situated basally and not arranged linearly. Hostplants of both families are in the Malviflorae (sensu Thorne, 1983). Homotomids have a characteristic bipartite male proctiger and many have a thickened and densely hirsute antennal flagellum, although the Macrohomotoma-group of genera have a simple filiform flagellum. All homotomids develop on species of Ficus (Urticales) and are distributed mainly in the Old World tropics, but one small genus occurs in Central and northern South America. Two species occur in the Mediterranean Basin on Ficus carica, and a third species extends into eastern South Africa on Ficus petersii. Carsidarids are diagnosed by the character-suite given on p. 89 and are restricted to hostplants in the Malvales; the group is almost entirely pantropical. Within the Carsidaridae the Central American genus Epicarsa has a distinctive forewing and antennal flagellum; the structure of the male proctiger is unique and the genus is considered to be the sister-group of the rest of the family. The single known species is found on Ceiba (Bombacaceae). Apart from Epicarsa and the Central and South American genus Paracarsidara the rest of the family is distributed in the Old World. Carsidara is quite distinct, having highly modified female terminalia and male aedeagal apex; it also retains the primtively broad forewing and very short M+Cu stem and has species in tropical Africa, Madagascar and South East Asia MALVALES-FEEDING PSYLLIDS 109 on sterculiaceous hosts. Of the narrow-winged genera Protyora is unique in retaining the primitive features of a costal break in the forewing and a simple, flask-shaped male proctiger; it is separated from the rest of the group by the derived state of the aedeagal apex. Two species are known, one in Sulawesi Utara and another in Australia on a sterculiaceous host. The remaining genera are difficult to characterise as a whole; they all possess a male proctiger that is enlarged basally and bears small or large lateral lobes but this character is also shared by Carsidara and is not likely to be a parallel development in this genus. Within this assemblage Tyora and Tenaphalara are grouped together, having an additional r;-rs crossvein and hind tibial spurs arranged 1+2+2. Tyora species have retained many primitive features, including the presence of radular spinules in cell cu; and the close proximity of the apex of the calval suture to the apex of Cu;,; they are restricted to sterculiaceous hosts in South East Asia. Tenaphalara has more derived features, its species occur in both Africa and the Oriental Region on hostplants in the Bombacaceae. Mesohomotoma, with species in the Afrotropical, Malagasy, Oriental, Australa- sian and Pacific Regions, has a distinctive bipartite male proctiger that bears a median posterior lobe; it also lacks a pterostigma in the forewing but this feature is shared, probably by convergence, with some species of Allocarsidara. The form of the endtube of the ductus ejaculatorius is similar to that of Paracarsidara and this is considered as a synapomorphy for the two genera. Mesohomotoma species usually develop on malvaceous hosts but one African species is known to live on various Sterculiaceae. Species of the Central and South American genus Paracarsidara develop on both Bombacaceae and Malvaceae. Little is known of the hostplants of the South East Asian genus Allocarsidara but three species develop on Durio zibethinus (Bombacaceae). The sister-group relationship of Allocarsidara is not resolved. It has rounded margins of the vertex, as in Tenaphalara, and some of the species lack a pterostigma, as in Mesohomotoma, but both these characters are likely to be parallel developments. A list of characters is given below and the matrix, from which the phylogeny (Fig. 1) was generated, is given in Table 2. Raven & Axelrod (1974), in their review of angiosperm biogeography, state, with reserva- tions, that the primary radiation of both the Malvales and the Moraceae occurred in Africa and South America in the Upper Cretaceous period. In the Malvales, the Sterculiaceae were the first group to reach Eurasia, followed by the Bombacaceae and then the Malvaceae. This is reasonably congruent with the hostplant preferences of the carsidarid genera. Those displaying more primitive features, e.g. Carsidara, Tyora and Protyora, are on Sterculiaceae, while those with more derived features, e.g. Tenaphalara, Allocarsidara, Paracarsidara and Mesohomoto- ma, are on Bombacaceae and Malvaceae. Mesohomotoma is anomalous as its species develop on both Malvaceae and Sterculiaceae, but the Sterculiaceae-feeding habit is restricted to one African Mesohomotoma species and may well be a secondarily derived condition. If Fig. 1 is a good reflection of carsidarid phylogeny, the subgroups must have been differentiated before the complete separation of Africa from South America (100 m.y.B.P.) as elements of the assemblage showing the most derived characters, e.g. Allocarsidara + Paracar- sidara + Mesohomotoma, are present today in both continents, and this is unlikely to be through subsequent dispersal from West Africa to the Caribbean. However, this is the weakest part of the projected phylogeny as these genera are not diagnosed as a group. List of characters Primitive condition is given in parenthesis 1. Integument of head and thorax sparsely short-setose or almost glabrous. (Integument of head and thorax densely long-setose. ) . Flagellomeres thickened. (Flagellomeres narrow, elongate. ) . Flagellomeres thickened and densely hirsute. (Falgellomeres narrow, elongate.) . Flagellomere 3 with subapical rhinarium. (Rhinarium absent from flagellomere 3.) Forewing angular apically. (Forewing rounded apically. ) Forewing narrow. (Forewing broad. ) . Costal break absent. (Costal break present.) . Pterostigma absent. (Pterostigma present. ) . Rs and M,,> anastomosing. (Rs and M,,, parallel, not in contact.) D. HOLLIS 110 + + + - + + + + + + + + DULOJOWOYOSaW Ae a a Ege oe oe. i 4 Meee DADPISADIDAD [+] + [4+] + + + + + + + + + + DADPISADIOIV + + + + + + + + + + + + + pavjoydoua], [+] + + + + + + + + + + + + AOA], + + + + + + + + + DLOK OAT - + + + + 4+ + + + + DADPISADD n + + + + + + + vsswndy [+] Eales) [+] =) oF SEOUL dnoip ie ice Kim WR WE ocl, BOT ic Ovetce © BOL Gis 6 Sty LO Hh ib RoE Re raqoBIeYD) “SIOQUIOU []e 0} UOWIWIOS jou sojouap [+] “}X9} UT sId}OBILYO JO JST] 0} JOJO sIOQUINU Ia}ORIeYD “OepIWOJOWOPF{ pue dseplepisieD JO xLQeUI-Io}ORIVYD 7 IIQeL 111 MALVALES-FEEDING PSYLLIDS JO S19quoWw [fe Aq poreys JOU 19}DeIeYO sajouap [ ] ‘aeplieptsieD 94) Jo sdnois juL[dj}soy pue uoNNQt}sip ‘AuasoyAyd Jo Arewuins AVUAOTAIATVW ‘(601 °d) 1X9} UT sIOJORIVYO Jo IsIT, 99S SIOQUINU 0} SULIAI S1d}ORIeYO 10,5 ‘dnois AVAOVITNOUALS “AVAOVATVIN PIIOM PIO DWUOJOWOYOSaW AVAOVATVNW ‘AVAOVOVENO PIHOM MON DAD PISADIDID AVAOVOVEWOd BISV “A'S D4DP1S4090]] AVAOVOVAWOd PITOM PIO Davjpydvua J, SLIVATVW AVaADVITAOUaLS BISV “A'S D404 J, AVAOVITNOAALS BIesny “eisy “q's DAOAJOLd AVAOVITNOAALS PIIOM PIO DADPISIDD AVAOVOVEWOE PILIOM MON pDsivid q ‘dds sno14 SATVOILYEN PIIOM MON FY PIO AVGINOLONOH 1z ‘[8] €I ‘Il (ot) ‘(ctl ft See EE [Z1] ‘9 ‘I Ee “9TEz OL ‘Ss 61 6 ‘7 I “sty [L] ‘OZ ‘PI “v 112 D. HOLLIS 10. Non-tracheate rs-m crossvein present. (rs-m crossvein absent. ) 11. Non-tracheate r;-rs crossvein present. (7;-rs crossvein absent.) 12. M+Cu stem long. (M+Cu stem short.) 13. Apical spurs of hind tibia arranged 1+2+2. (Hind tibial spurs arranged 1+2(or 3)+1.) 14. Metabasitarsus with a single apical spur. (Metabasitarsus with two apical spurs.) 15. Male proctiger bipartite, apical section consisting of a simple anal tube. (Male proctiger unipartite, tubular.) 16. Male proctiger bipartite, apical section having a median posterior lobe in addition to anal tube. (Male proctiger unipartite, tubular.) 17. Male proctiger with enlarged basal portion subdivided posteroapically into small lateral lobes. (Male proctiger tubular.) 18. Male proctiger with well-developed, simple lateral lobes. (Male proctiger tubular.) 19. Lateral lobes of male proctiger each with a secondary inward-pointing lobe. (Secondary, inward- pointing lobes absent.) 20. Male subgenital plate with supplementary lobes. (Male subgenital plate without supplementary lobes.) 21. Inner lateral surfaces of male subgenital plate with patches of spinules. (These spinules not differenti- ated.) 22. Apex of aedeagus enlarged, endtube of ductus ejaculatorius simple. (Apex of aedeagus narrow, endtube of ductus ejaculatorius simple.) 23. Apex of aedeagus enlarged, complex, endtube of ductus ejaculatorius laterally compressed and heavily chitinised. (Apex of aedeagus narrow, endtube of ductus ejaculatorius simple.) 24. Ductus ejaculatorius with a complex endtube bearing ridges and hooks. (Endtube of ductus ejacula- torius without ridges or hooks.) 25. Female proctiger with a posterodorsal lobe. (Female proctiger without a posterodorsal lobe.) 26. Lateral margins of female proctiger densely long-setose. (Lateral margins of female proctiger sparsely short-setose. ) 27. Lateral palps of ovipositor heavily ridged. (Ovipositor palps not ridged.) 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The ant mosaic -— tropical tree crops and the limiting of pests and diseases. Pest Articles and News Summaries 19: 311-341. Leston, D. & Gibbs, D. G. 1968. A new Deraeocorine (Hemiptera: Miridae) predacious on Mesohomoto- ma tessmanni (Aulmann) (Hemiptera: Psyllidae) on Cocoa. Proceedings of the Royal Entomological Society of London (B) 37: 57-88. Low, F. 1886. Neue Beitrage zur Kenntniss der Psyllidae. Verhandlungen der kaiser-k6niglichen zoolo- gisch-botanischen Gesellschaft, Wien 36: 149-169. Mathur, R. N. 1975. Psyllidae of the Indian Subcontinent. xii + 429 pp. New Delhi. Miyatake, Y. 1965. Notes on Psyllidae from the Ryukyu Islands (Hemiptera: Homoptera). Kontyii 33: 171-189. — 1971. Studies on the Philippine Psyllidae (Hemiptera: Homoptera). I. Bulletin of the Osaka Museum of Natural History 25: 51-60. 1972. Studies on the Philippine Psyllidae (Hemiptera: Homoptera). II. Bulletin of the Osaka Museum of Natural History 26: 11-33. 1981. Discovery of the tropical psyllid Thysanogyna minor from Japan, with description of a new subspecies (Hemiptera: Psyllidae). Bulletin of the Osaka Museum of Natural History 35: 37-41. Morgan, F. D. 1984. Psylloidea of South Australia. Handbook of the flora and fauna of South Australia. 136 pp. South Australia. Orian, A. 1972. The Psylloidea of Mauritius with a description of Trioza eastopi sp. nov. Insecta — Homoptera. Fauna of Mauritius 1: [vii] + 1-6 + [2]. Pettey, F. W. 1924. South African Psyllids. Entomology Memoirs. Union of South Africa, Department of Agriculture 2: 21-30. 1925. New South African Psyllids. South African Journal of Natural History 5: 125-142. Rahman, K. A. 1932. Observations on the immature stages of some Indian Psyllidae [Homoptera: Rhynchota]. Indian Journal of Agricultural Science 2: 358-377. Raven, P. H. & Axelrod, D. I. 1974. Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden 61: 539-673. Roberts, H. 1969. Forest insects of Nigeria, with notes on their biology and distribution. /nstitute Paper, Commonwealth Forestry Institute, Oxford 44: 206 pp. Sasaki, K. 1954. A list of the known species and their host-plants of the Psyllidae of Japan (Homoptera). Scientific Reports of the Matsuyama Agricultural College 14: 29-39. Scott, J. 1882. On certain genera and species of the group of Psyllidae in the collection of the British Museum. Transactions of the Entomological Society 1882: 449-473. Silva, A. G. d’Araitijo e et al. 1968. Quarto catalogo dos insetos que vivem nas plantas do Brasil. Seus parasitos e predatores. Parte II- 1. Tomo. Insetos, hospedeiros e inimigos naturais. xxiv + [iv] + 622 pp. Rio de Janeiro. MALVALES-FEEDING PSYLLIDS aS Sohi, A. S. & Sandhu, G. S. 1978. Chemical control of Simbal Psylla on silk cotton tree in Punjab. Science and Culture, 44: 237-238. Takahashi, R. 1936. Food habits and new habits of Formosan Psyllidae, with notes on the peculiar food habits of Formosan phytophagous insects. Kontyii 10: 291-296. [In Japanese. | Thorne, R. F. 1983. Proposed new realignments in the angiosperms. Nordic Journal of Botany 3: 85-117. Tuthill, L. D. 1943. Psyllidae from Rapa, the Caroline, Society and Austral Islands (Homoptera). Occasional papers of the Bernice P. Bishop Museum 17: 71-78. 1950. Contributions to the knowledge of the Psyllidae of Mexico (Part II). Journal of the Kansas Entomological Society 23: 52-63. 1951. Records and descriptions of some Micronesian Psyllidae (Homoptera). Pacific Science 5: 273-278. — 1964. Homoptera: Psyllidae. Insects of Micronesia 6: 353-376. Tuthill, L. D. & Taylor, K. L. 1955. Australian genera of the family Psyllidae (Hemiptera, Homoptera). Australian Journal of Zoology 3: 227-257. Uichanco, L. B. 1921. New records and species of Psyllidae from the Philippine Islands, with descriptions of some preadult stages and habits. Philippine Journal of Science 18: 259-288. Vondraéek, K. 1957. Mery — Psylloidea. Fauna CSR 9: 431 pp. Walker, F. 1869. Catalogue of the Homopterous insects collected in the Indian Archipelago by Mr. A. R. Wallace, with descriptions of new species. Journal of the Linnean Society (Zoology) 10: 276-330. White, I. M. & Hodkinson, I. D. 1985. Nymphal taxonomy and systematics of the Psylloidea (Homoptera). Bulletin of the British Museum (Natural History) (Entomology) 50: 153-301. Wood, G. A. R. 1980. Cocoa (3rd end). xii + 292 pp. London & New York. Wu, C. F. 1935. Catalogus Insectorum Sinensium (Catalogue of Chinese Insects). 11. Family Psyllidae: 125-126. Peiping. Yang, Chi-kun & Li Fasheng 1985a. Nine new species and a new genus of psyllids from Yunnan (Homoptera: Psyllidae). Entomotaxonomia 6 (1984): 251-266. [In Chinese; English summary. | — 1985b. Notes on Tenaphalara and Tyora with descriptions of five new species from China (Homop- tera: Psyllidae). Entomotaxonomia 7: 205-216. [In Chinese; English summary. | Yang, Chung Tu 1984. Psyllidae of Tatwan. Taiwan Museum Special Publication 3: 1—305. 116 D. HOLLIS Figs 2-17 Carsidaridae, head and antennal structure. 2, Epicarsa corniculata, head and antenna, dorsal view. 3, 4, Carsidara africana; 3, head, dorsal view; 4, same, lateral view. 5, 6, Protyora wilsoni; 5, head, dorsal view; 6, same, lateral view. 7, 8, Tyora congrua; 7, head, dorsal view; 8, same, lateral view. 9, 10, Tenaphalara camerunus; 9, head, dorsal view; 10, same, lateral view. 11-13, Allocarsidara iriana; 11, head, dorsal view; 12, same, lateral view; 13, flagellomeres 7 and 8. 14, 15, Paracarsidara gigantea; 14, head, dorsal view; 15, same lateral view. 16, 17, Mesohomotoma tessmanni; 16, head, dorsal view; 17, same, lateral view. Scale lines: Fig. 13 0-1 mm, remainder 0-5 mm. Setosity not shown. Figs 3—12, 14-17 drawn from dry-mounted specimens. MALVALES-FEEDING PSYLLIDS ‘7 Figs 18-28 Carsidaridae forewings. 18, Epicarsa corniculata. 19, Carsidara africana. 20, 21, Tyora congrua; 20, from Sulawesi; 21, from Java. 22, T. buxtoni (Solomon Is). 23, T. ornata. 24, T. striata. 25, Tenaphalara acutipennis. 26, T. camerunus. 27, Protyora sterculiae. 28, P. wilsoni. Scale lines, 0-5 mm. Setosity not shown. 118 D. HOLLIS Figs 29-39 Carsidaridae forewings. 29, Allocarsidara malayensis. 30, A. incognita. 31, A. iriana. 32, A. juliana. 33, A. bakeri. 34, A. elongata. 35, Paracarsidara gigantea. 36, 37, Mesohomotoma hibisci; 36, from Sulawesi; 37, from Seychelles. lines, 0-5 mm. Setosity not shown. 38, 39, M. tessmanni; 38, from Angola; 39, from Sierra Leone. Scale MALVALES-FEEDING PSYLLIDS 119 Figs 40-45 Carsidaridae male genitalia. 40-42, Carsidara africana; 40, proctiger and subgenital plate, lateral view; 41, right paramere, inner lateral view; 42, apical segment of aedeagus, lateral view. 43-45, Protyora wilsoni; 43, proctiger and subgenital plate, lateral view; 44, right paramere, inner lateral view; 45, apical segment of aedeagus, lateral view. Scale lines, 0-1 mm. 120 D. HOLLIS Figs 46-56 Carsidaridae male genitalia. 46, 47, Tyora buxtoni (Solomon Is); 46, proctiger and subgenital plate, lateral view; 47, right paramere, anterolateral view. 48, 49, T. ornata (Fiji); 48, proctiger, lateral view; 49, right paramere, anterolateral view. 50, T. congrua (Sulawesi), apical segment of aedeagus, lateral view. 51-53, Tenaphalara acutipennis; 51, proctiger and subgenital plate, lateral view; 52, right paramere, anterolateral view; 53, apical segment of aedeagus, lateral view. 54-56, T. camerunus; 54, proctiger and subgenital plate, lateral view; 55, right paramere, anterolateral view; 56, apical segment of aedeagus, lateral view. Scale lines, 0-1 mm. MALVALES-FEEDING PSYLLIDS 121 Figs 57-65 Carsidaridae male genitalia. 57-59, Allocarsidara malayensis; 57, proctiger and subgenital plate, lateral view; 58, right paramere, inner lateral view; 59, apical segment of aedeagus, lateral view. 60-62, A. elongata; 60, proctiger and subgenital plate, lateral view; 61, right paramere, inner lateral view; 62, apical segment of aedeagus, lateral view. 63-65, A. iriana; 63, proctiger and subgenital plate, lateral view; 64, right paramere, inner lateral view; 65, apical segment of aedeagus, lateral view. Area of differentiated spinules indicated within pecked lines in Figs 57, 60, 63. Scale lines, 0-1 mm. 122 D. HOLLIS 14 Figs 66-74 Carsidaridae male genitalia. 66-68, Allocarsidara juliana; 66, proctiger and subgenital plate, lateral view; 67, right paramere, inner lateral view; 68, apical segment of aedeagus, lateral view. 69-71, A. bakeri; 69, proctiger and subgenital plate, lateral view; 70, right paramere, inner lateral view; 71, apical segment of aedeagus, lateral view. 72-74, A. incognita; 72, proctiger and subgenital plate, lateral view; 73, right paramere, inner lateral view; 74, apical segment of aedeagus, lateral view. Area of differentiated spinules indicated within pecked lines in Figs 66, 69, 72. Scale lines, 0-1 mm. MALVALES-FEEDING PSYLLIDS 123 Figs 75-78 Carsidaridae male genitalia. 75, 76, Paracarsidara dugesii; 75, proctiger and subgenital plate, lateral view; 76, apical segment of aedeagus, lateral view. 77, 78, Mesohomotoma hibisci (Seychelles); 77, proctiger and subgenital plate, lateral view; 78, apical segment of aedeagus, lateral view. Scale lines, 0-1 mm. 124 D. HOLLIS —— if EEL CCH 7 e ( te aC (UC ee AN SS A Figs 79-89 Carsidaridae female terminalia, lateral views. 79, Carsidara africana; 80, Protyora wilsoni; 81, Tyora congrua (Sulawesi); 82, T. striata; 83, Tenaphalara camerunus, 84, Allocarsidara juliana, 85, A. incognita; 86, A. malayensis; 87, A. elongata; 88, Paracarsidara dugesii; 89, Mesohomotoma hibisct (Tanzania). Scale line, 0-1 mm. MALVALES-FEEDING PSYLLIDS 125 Figs 90-94 Carsidaridae larval structure, dorsal (d) and ventral (v) views of caudal area showing arrangement of pore bands. 90, Protyora sterculiae, also showing chaetotaxy; 91, Tenaphalara camer- unus;, 92, Allocarsidara incognita; 93, Paracarsidara dugesii; 94, Mesohomotoma hibisci (Tanzania). Scale lines, 0-1 mm. 126 Abutilon 106 indicum 88, 106, 107 Anacardiaceae 99 Annonaceae 108 Aphanamixis grandifolia 99 Bombacaceae 87, 88, 89, 91, 98, 99, 101, 102,104, 105, 108, 109; 114 Bombax 98 buonopozense 88, 99 ceiba 88, 98, 99 cyathophorum 105 malabaricum 88 sessile 99 Boraginaceae 107 Brachychiton 92, 94 populneum 88, 95 rupestris 93 sp. 95 Cedrela 90 Ceiba 91, 98, 104, 108 burchellii 105 pentrandra 88, 99 sp. 92, 105 Chytranthus talboti 108 Cinnamomum camphora 107 Cleistopholis 108 sp. 108 Cola 106, 108 spp. 108 Cordia dichotoma 107 D. HOLLIS Index to hostplants Invalid names are in italics. Desplatzia 108 lutea 108 Dimocarpus longana 99 Durio 101 zibethinus 88, 101, 102, 103, 104, 109 Ebenaceae 100 Ficus 90, 108 carica 108 peters 108 spp. 111 Firmiana 92 simplex 88, 93 Gossampinus malabarica 99 Heritiera 96, 106, 108 littoralis 88, 97 spp. 108 Hibiscus 106 boryanus 107 esculentus 98 mutabilis 107 rosasinensis 107 sp. 107, 108 tiliaceus 88, 106, 107 Lauraceae 107 Malachra alceifolia 105 Mallotus japonicus 93 Malva 104 sp. 105 Index Malvaceae 87, 88, 89, 104, 105, 106, 107, 108, 109, 111 Malvales 87, 88, 89, 90, 108, 109, et Malviflorae 87, 108, 111 Mangifera indica 99 Meliaceae 90, 99 Moraceae 109 Palaquium formosanum 100 sp. 100 Sapindaceae 99, 108 Sapotaceae 100 Sterculia 96, 106 foetida 98 lanceolata 97 sp. 97 tragacantha 108 ureolata 97 Sterculiaceae 87, 88, 89, 92, 94, 96, 97, 106, 107, 108, 109, 111 Theobroma 106, 108 cacao 88, 108 Thespesia 106 populnea 107 Tiliaceae 108 Urena 106 lobata 106, 107, 108 Urticales 108, 111 Wissadula 104 perplocifolia 105 Invalid names are in italics; principal references are in bold. Aconopsylla 89 acutipennis 88, 98-99 africana, Carsidara 93-94 africana, Mesohomotoma 106, 107 Allocarsidara 87, 90, 91 (key), 94, 96, 100-101, 104, 106, 109, 110, 111 Aphalaridae 89 aphanamixis 99 Apsylla 90 Baccha 88 bakeri 101, 102, 103, 104 buxtoni 96 Calophyidae 87, 89, 90, 100 camerunus, Carsidara 99 camerunus, Tenaphalara 98, 99 camphorae 106, 107 Carsidara 87, 89, 90, 91 (key), 92, 94, 104, 106, 108, 109, 110, 111 Carsidara sp. 93 Carsidaridae 87, 88, 89-90, 94, 100, 108, 110, 111 Carsidarinae 89, 100 Carsidaroida 89, 95 Ciriacreminae 89 Ciriacremum 90 concolor, Carsidara 105 concolor, Paracarsidara 105 confluens, Strogylocephala 100 confluens, Synaphalara 100 confluens, Tenaphalara 100 congrua 95, 96-97 corniculata, Epicarsa 91, 92 corniculata, Epiciasa 92 corniculata, Epicrasa 92 crigi, Deraeocoris 88 Deraeocoris 88 Diceraopsylla 89 Diclidophlebia 89 dimocarpi 99 dugesil, Carsidara 104, 105 dugesii, Paracarsidara 105 Dynopsylla 92 elongata, Allocarsidara 101, 104 elongata, Ctenophalara 98 elongata, Tenaphalara 98, 99 Epicarsa 87, 89, 90 (key), 91-92, 108, 110, 111 Epiciasa 91 Epicrasa 91 Eustigmia 92 fascipennis, Strogylocephala 99-100 fascipennis, Tenaphalara 98, 99, 100 gigantea, Carsidara 105 gigantea, Paracarsidara 105 gossampini 99 guandongana 97 Haplaphalara 89 heterocephala, Carsidaroida 95, 96 heterocephala, Nesiope 96 hibisci, Mesohomotoma 106, 107 hibisci, Tyora 106, 107 hibisci, Udamostigma 107 Homotomidae 87, 90, 91, 108, iO. 111 incognita 101, 102-103 indica, Tyora 107 indica, Tyoria 108 intermedia, Nesiope 96 iriana 101, 103 javana, Sphaerophoria 88 juliana, Allocarsidara 100, 101, 102, 103, 104 juliana, Tenaphalara 98, 101, 102 limbata, Carsidara 93 limbata, Thysanogyna 93 lineaticollis 106, 107 lutheri, Mesohomotoma 106, 107-108 MALVALES-FEEDING PSYLLIDS lutheri, Udamostigma 107 Macrohomotoma 108 malayensis, Allocarsidara 100, 101, 102, 103 malayensis, Tenaphalara 98, 100, 101, 102 mangiferae 99 marginalis 92, 93, 94 Mastigimas 87, 89, 90 Mesohomotoma 87, 88, 89, 90 (key), 92, 94, 96, 101, 104, 106, 107, 109, 110, 111 mexicana, Carsidara 105 mexicana, Paracarsidara 105 minor, Dynopsylla 92, 93 minor, Eustigmia 93 minor, Thysanogyna 93 Neocarsidara 94, 95 Nesiope 89, 95 obsoleta-group, Trioza 100 ornata, Nesiope 95, 97 ornata, Tyora 96, 97 Paracarsidara 87, 89, 90, 91 (key), 92, 94, 96, 101, 104, 106, 108, 109,110, 111 Phacopteronidae 89, 90 picta, Baccha 88 praeusta, Baccha 88 Prionocnemidinae 89 Prionocnemina 89 Prionocneminae 89 Protyora 87, 89, 90, 91 (key), 94-95, 109, 110, 111 pseudonervosa, Nesiope 99 pseudonervosa, Tenaphalara 98, 99 pseudonervosa, Tyora 99 Psyllidae 89, 90 pulchriformis, Baccha 88 rostrata, Carsidara 105 rostrata, Paracarsidara 105 shikokuensis, Carsidara 93 shikokuensis, Thysanogyna 93 sinuata, Baccha 88 Sphaerophoria 88 sterculiae, Neocarsidara 95 sterculiae, Protyora 88, 95 sterculiae, Tyora 94, 95 striata, Tenaphalara 97, 98 striata, Tyora 96, 97 Strogylocephala 87, 89, 100 sulcata, Tenaphalara 97, 100 Synaphalara 87, 100 Tenaphalara 87, 89, 90, 91 (key), 94, 96, 97-98, 99, 100, 101, LOS; 110. 111. Tenaphalarini 89 tessmanni, Mesohomotoma 88, 106, 108 tessmanni, Tyora 108 tessmanni, Udamostigma 108 Thysanogyna 92 Togepsylla 89 Trioza 100 Triozidae 88, 89, 90 triozipennis, Tenaphalara 98, 100 triozipennis, Trioza 100 Trisetipsylla 90 Tyora 87, 89, 90, 91 (key), 94, 95-96, 98, 106, 109, 110, 111 Tyoria 106 Udamostigma 106 umalii, Tenaphalara 100 umalii, Trioza 100 wilsoni 95 127 at ns oo jalne? hoes Of eta better i? wvabia) tanien dusty, aie y= eq poor zal Se" Lol ™ ware FO F he = os ‘aie, Wihvad eo | Qn es ; | est ms ae a * a - eo, a —— Py * ° 4) =ee 2 ov duane “as eee , 7 | Sf ae. Ai 1) a British Museum (Natural History) Milkweed butterflies: their cladistics and biology P.R. Ackery & R. I. Vane-Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of their biology have stimulated far more attention than can be justified by species numbers alone. In recent years, an expansive literature has grown, considering aspects of their courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and genetics. The popularity of danaines among biologists can certainly be attributed to this combination, within one small group, of so many of the factors that make butterflies such an interesting group to study. The obvious need to place this wealth of biological data within an acceptable systematic framework provided the impetus for this volume. Started in 1976 within the conventions of evolution by natural selection and Hennig’s phylogenetic systematics, the book is now largely about natural history (what the animals have and do, where they live and how they develop) and natural groups — as revealed by a form of analysis approaching that practised by the new school of ‘transformed cladistics’. The authors have prepared a handbook that will appeal to a wide range of biologists, from museum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. ISBN 0 565 00893 5. 1984. Price £50. Titles published in Volume 56 The legume-feeding psyllids (Homoptera) of the west Palaearctic Region By I. D. Hodkinson & D. Hollis A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera) By D. Hollis A review of the Rhadalinae (= Aplocneminae) (Coleoptera: Melyridae) By E. R. Peacock Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural History) LO LIE LIE EL OLIN IE NOME PT ae wh tet * : 4 Vt i PaN\/ (scum Vat foyi ; MATEO “er AY (NATURAL HISTGRY) 22 DEC i#37 PRESENTED ENTOMOLOGY LIBRARY A review of the Rhadalinae = Aplocneminae) Coleoptera: Melyridae) Enid R. Peacock Entomology series Vol 56 No 3 17 December 1987 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1987 The Entomology series is produced under the editorship of the Keeper of Entomology: Laurence A. Mound Publications Manager (Entomology): W. Gerald Tremewan gE TT NI een eee f Pe 8 4 y Ss < / Ae, al I f/ f eS ws y é - ry a ae VEL 409 cL ow i I7 DEC / i Hl tea % 3 ‘ 2 x * es ISBN 0 565 06030 9 SE JRAL HIST ISSN 0524-6431 British Museum (Natural History) Cromwell Road London SW7 5BD _ Issued 17 December 1987 Entomology series Vol 56 No 3 pp 129-170 i Tos oe A review of the Rhadalinae (= Aplocneminae) (Coleoptera: Melyridae) Enid R. Peacock Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD Contents Se MMR REEC NRE Pe yat le teeters (hee Re Mca tee radon ee Doi R ln Fcbicek Pu lnsd dalns's anetdoudb ai’ bolded be deeds 129 RIS SS Rs Ae oe ey ee ee ee ee eee ae £29 EE LSC SMLIS IS BE Ee ere ee ne, ee ee a eee oe eee 130 RIDES PALET SOC AUIS er 200 eo. tcise aad) 3s. ewaees Sane bose xoas ces ee alt agen ee, AL hate 130 RICO CONG aries ee he sd art Mees oan 9 Bln se Saba Bo Goa Ma At vinch a ved vbu DN tos ane 131 erm ANG CISCIMOUISMING TEACUTES (7 (26): 2 cock ae nocd vn ne daming ee neon voaets somes sanevens 131 EN SI ry ae PREG alt ales Pa ae wa tks ss st otgusddadseleas rose ge coveds don sanatessnoeens 131 _ TEES ww chese ecentoe angie oli Sate tesa er eee a ae en en 131 (SD TY noo goge odode ea aetig athe me Ase in cli ees tie a ene eee aan een £51 macemisr otthcwenera ol the Rhadalinae’.. 2.25.1. co. feces ccegectectac cer eeenseqasneseene 132 etesion the keyand ChecklistSOfSpecies: .... ee... cee cece cece ecececceccesecetondenoeeees £35 SE VLGL TOIT Sie TA >: BSR ee ee ee er ee ee eet ner eee re terre 135 eye adults of the world'genera of the Rhadalinae .................02.-scscteeteeceereceeeeeees 133 2 EDL EDLS GIS INO OES Es SNnocs Bae 8 Sacer aoe ase a enn rece ek gee 144 me MNNUMLE CLC INTC IIS) eepeette 26 ost oeics Sse entudnons Lome cate 22 laine cn damonet leans ade dae ot Sea acne = ae 162 © LL EPPETLGES La sh hare ao ctl gee 8 i ne Va a ge a a ©. oe ee 162 RE RR ET Ree nt BN 5 WE Loe ad bo Genb chonnine a emmmaes do ogee Phu ubasiaines oeeies 167 Synopsis Thirteen world genera of Rhadalinae (Melyridae) are recognized here, of which three are included in the subfamily for the first time: Antinea Peyerimhoff, Indiodasytes Pic and Hemipleurus gen. n., erected for H. floriger sp. n. from Borneo. A generic key is given together with a diagnosis and type species citation for each genus. Eurelymis Casey (NW. America), from the Melyrinae, and Celsus Lewis (Japan), from the Dasytinae, are regarded as junior synonyms of Semijulistus Schilsky (Holarctic Region), Cymbolus Gorham is synonymized with Rhadalus LeConte and Xamerpus Fairmaire with Malthacodes Waterhouse. The two Indian species of Aplocnemus Stephens, A. indicus Champion and A. moestus Gorham, are transferred to Malthacodes Waterhouse. Malthacodes parvus nom. n. is proposed as a replacement name for M. minutus (Pic, 1931) [originally in Xamerpus] (nec Pic, 1906). Checklists of genera and species are provided. Distinguishing features are given for adult and larval Melyridae and Rhadalinae and comments are made on the biology of the latter. Introduction The Melyridae (sensu Crowson, 1964 and Lawrence, 1982) is the largest family in the superfamily Cleroidea, with at least 200 genera. Crowson (1964) considered it to be divisible into five subfamilies: the Rhadalinae (Aplocneminae), the Melyrinae, the Malachiinae, the Prionocerinae and the Dasytinae. These subfamilies have all, at one time or another, been given family rank and some are currently considered as such by recent authors (e.g. Constantin, 1965; Liberti, 1984; Wittmer, 1984 etc.). In spite of its size the family has been poorly studied in recent years, except for the subfamily Malachiinae which has been the subject of extensive studies by Wittmer (1930 et seq.). At the beginning of the century and before, there were several prolific writers whose unco-ordinated descriptive efforts, mainly at specific level, caused a certain amount of taxonomic confusion in the family. Bull. Br. Mus. nat. Hist. (Ent.) 56 (3): 129-170 Issued 17 December 1987 130 E. R. PEACOCK The first author to create some order out of this confusion was Crowson who, in his major work on the higher classification of Coleoptera (1955) and in subsequent works (1964, 1970), keyed out and defined the superfamilies, families and subfamilies. Since then, Majer (1987) has produced a useful work completely revising the phylogeny and taxonomy of the family. He introduced new taxa, divided the family into subfamilies, supertribes and tribes and discussed in detail the supposed phylogeny of the group. In the course of his study of the Melyridae, Crowson (1964) discovered a distinctive group of genera whose close relationship had been hitherto overlooked and for which he erected a new subfamily, the Aplocneminae. This included genera from the melyrid subfamilies Melyrinae and Dasytinae, the Cleridae and the Rhadalidae (Coleopterorum Catalogus, Pic, 1929, 1937; Corporaal, 1950; Pic, 1926). However, Crowson overlooked the fact that a family group name (Rhadalidae) was already in use for one of his included genera: Rhadalus LeConte. The name Rhadalinae therefore has priority (Article 23(a), International Code of Zoological Nomencla- ture, 1985) and is employed here. These discoveries, along with some additional genera, nomenclatural changes, new generic synonymies and the erection of a new genus, indicated the need for a review of the entire subfamily and hence this study. In the present paper Crowson’s (1964) treatment of the subfamily is reappraised, decisions concerning the status of the included genera being based on the study of their type species (something that Crowson failed to do). As a result 12 of the 25 nominal genera are regarded as valid and one additional genus is described as new. Although the Rhadalinae comprises only 13 (out of 25 nominal) genera, its distribution is almost world-wide. The group seems to be restricted mainly to semi-dry forested, mountainous or hilly areas, although some species have been collected on sea beaches. The adult beetles are found on flowers, shrubs or on the leaves or bark of coniferous or deciduous trees and although most are carnivorous to a certain extent, some are pollinivorous or both carnivorous and pollinivorous. The subfamily is well defined, all of its species exhibiting the unusual feature of the connation of the first two visible abdominal sternites. This feature is found in no other group of Melyridae nor in other cleroid families. Its other unusual and notable feature is the sclerite articulated to the (morphologically) dorsal side of the median lobe of the aedeagus. This sclerite articulates at its basal end with the median lobe in a hinge-like joint, and is freely moveable in all the genera examined except Microjulistus, in which it appears to be fused at the base to the median lobe. The only genus in the group lacking this sclerite (or appendage) is the Indian Indiodasytes. MELYRIDAE Leach, 1815’ Distinguishing features Adults. Adult Melyridae are generally distinguished from related families by the following characters: antennae usually 11-segmented and serrate or filiform, but sometimes pectinate or with some basal segments modified, apical segments not differentiated to form a club; claws either simple, split, toothed or with membranous appendages; front coxal cavities open behind, coxae large and projecting, with exposed trochantins; tarsi 5-segmented, sometimes 4,5,5 in male [exception Anthriboclerus, 4,4,4] and without strongly lobed segments; prothorax usually with distinct side margins; elytra without striae (except in some Melyris spp. which also have costae); sometimes with exsertile vesicles at anterior angles of prothorax and abdominal pleura; abdomen with 6 visible sternites which are usually free but occasionally with the first two connate; aedeagus with undivided tegmen. Larvae. Melyrid larvae can be distinguished from those of all other cleroid families [except Phycosecidae] by the following characters: head with a well-marked median epicranial suture and no endocarina; mouthparts retracted, stipes much longer than cardo; spiracles annuliform (Crowson, 1970); mandible with a long, stiff prosthecal process near the middle or at the base of inner margin (Boving & Craighead, 1930). They differ from Phycosecidae in possessing 1—5 ocelli on each side instead of 6. [The larvae of other cleroid families have the following characters: head rarely with a distinct median cranial suture, but if so, ' Fide Watt (1975: 33). Pic (1937: 3) and Majer (1987: 784) give Olivier, 1790 as author but there is no family group name in that work. REVIEW OF THE RHADALINAE 131 then with mouthparts strongly protracted and stipes not longer than cardo; endocarina usually distinct; spiracles usually bicameral; mandible either without a prosthecal process or with a short one. | RHADALINAE LeConte Rhadalini LeConte, 1861: 191, 194; LeConte & Horn, 1883: 213, 216. ‘Haplocnemates’ Mulsant & Rey, 1868: 181. Rhadalinae LeConte; Casey, 1895: 457; Blaisdell, 1938: 3; Hatch, 1962: 86; Arnett, 1968: 609, 611, 613; Blackwelder, 1975: R67.10. Rhadalidae LeConte; Pic, 1926a: 3; Crowson, 1964: 315, 320. Aplocneminae ([erratim Haplocneminae] Crowson, 1964: 316, 317, 318, 319, 320; Constantin, 1965: 92; Vinson, 1967: 330); Majer, 1983: 387. Syn. n. Aplocnemina; Majer, 1987: 800. Taxonomy and distinguishing features Adults Adult Rhadalinae are distinguished from the other subfamilies of Melyridae by the following characters in combination:- first two visible abdominal sternites connate; apical segment of maxillary palps broadened, securiform or triangular except in some species of Aplocnemus Stephens); median lobe of aedeagus with dorsal appendage (Fig. 7) or ‘lever’ (Majer, 1982). Members of the Rhadalinae are diverse in appearance but all have the three characters mentioned above, except for Aplocnemus (Ischnopalpus) which has a spindle-shaped apical segment to the maxillary palps, and Indiodasytes which lacks the dorsal appendage to the median lobe of the aedeagus. They are also characterized by the following features:- antennae usually serrate, sometimes pectinate; eyes entire or weakly emarginate; head generally broader than long, often retractable under pronotum; body usually densely hairy and strongly convex; tarsi varying from finely elongate and simple to short, broad and weakly lobed, segment 1 normally at least as long as 2; tarsal claws toothed or with free membranous appendages; dorsum either unicolorous or patterned; puncturation on head simple, rimmed, or confused; pronotal punctures entirely simple, or simple on disc but rimmed or tuberculate at sides, or all rimmed or all tuberculate, or confused so that individual punctures are not distinguishable; elytral punctures usually simple, sometimes with a seta emerging from the puncture, but more usually from beside it; epipleura well developed at base; wing venation variable as shown in Figs 48-59. Crowson (1964) erected the subfamily Aplocneminae (as Haplocneminae) [= Rhadalinae] for the following genera: Anthriboclerus Schenkling, [H]/aplocnemus Stephens, Cymbolus Gorham, Diplambe Schilsky, Donaldia Alluaud, Eucymbolus Champion, Ischnopalpus Schilsky, Julistus Kiesenwetter, Malthacodes Waterhouse, Pelecophora Dejean, Rhadalus LeConte, Trichoceble Thomson and Xamerpus Fairmaire. Majer (1983), who regards the group as of lower taxonomic rank than subfamily, added Kubanius Majer, Semijulistus Schilsky and Microjulistus Reitter. In this paper I add: Antinea Peyerimhoff, Celsus Lewis, Eurelymis Casey, Indiodasytes Pic and Hemipleurus gen. n. A complete checklist of included genera is provided below. Larvae. The known rhadaline larvae (Trichoceble, Aplocnemus, and Pelecophora) possess the following characters (Crowson, 1964): head with 2 ocelli on each side, the larger one being anterior to the smaller, so that a line through the 2 would pass through the antennal foramen; urogomphi, on abdominal tergite 9, varying from minute to very long; abdominal tergite 9 either with a membranous appendage in the middle of each side (Pelecophora and Aplocnemus), or with a group of setiferous tubercles (unnamed larva from Chile) or forming a large flat plate with minute, widely separated urogomphi (Trichoceble). Rhadaline and melyrine larvae have glandular openings on the 9th abdominal tergite only (Crowson, 1981) and some Aplocnemus larvae have dark subcuticular patches on abdominal segments 1-8, usually 2 pairs on each segment. Vinson gives a detailed description of a larva assumed to be Pelecophora pikei Vinson (1957). Larvae of the other melyrid subfamilies usually have more ocelli, but if there are only two on each side, then they are one above the other so that a line through them would not pass through the antennal foramen. Tergite 9 is without the above mentioned structures. Biology With the exception of Australasia, species of Rhadalinae are known to occur in all major biogeographic regions, but little is known about their biology. Most adult rhadalines are probably carnivorous to a certain extent. Examination of the gut contents of adult species of Aplocnemus, Trichoceble, Pelecophora and Donaldia |= Malthacodes| has revealed insect 132 E. R. PEACOCK fragments (Vinson, 1946; Crowson, 1964), but in species of Rhadalus and Indiodasytes only pollen grains were found (Crowson, 1964; Peacock, pers. obs.). Denticulation of the cutting edge of the mandible, a character indicating pollinivorous habits, is absent in most Rhadalinae, although it is shown in a figure of a mandible of Pelecophora (Vinson, 1946), indicating perhaps that this genus is both carnivorous and pollinivorous. It is likely that the larvae of this group are carnivorous as are most cleroid larvae, even though the examination of some larval gut contents by Crowson (1964) revealed no identifiable remains. This view is supported by the presence of a pedunculate seta (Vinson called it a ‘lacinia’) on the maxillary mala of a larva of Pelecophora (Vinson, 1957), a character peculiar to all known carnivorous cleroid larvae (Crowson, 1964). There appear to be scant records of habitat preference and the few larvae that have been collected seem to live concealed either on or in the soil or in decaying wood. The following paragraphs give an indication of the habitat preferred by some adults. In Mauritius, Pelecophora and Donaldia |= Malthacodes| have been found during the day on the leaves and branches of shrubs and trees, where they were probably feeding on small insects (Vinson, 1946). Vinson noticed that these seemingly carnivorous beetles are constantly associated with the indigenous vegetation, occurring in their greatest numbers during the hot season from November to March. He also found a Pelecophora larva under a stone in association with an adult crawling out of a pupal cell (1957). Adults of the North American Eurelymis [= Semijulistus| are often found at high altitudes. They are attracted to wood smoke (Leech, 1931) and have been found on Achillea (Hatch, 1961). E. [= S.] atra LeConte has been reared from pupae which were found in bee burrows, in sand, in Alberta, Canada, and E. [= S.] flavipes LeConte was found on Pyrocantha in California [specimens loaned to author]. Cymbolus [= Rhadalus| elongatus Champion was collected in Temascaltepec, Mexico, in large numbers on Mimosa and Spondias spp. and several individuals of C. [= R.] wolcotti Hinton were collected on Pinus pseudostrobus Lindley (Hinton, 1934). In Japan, Celsus [= Semijulistus| is not common and has only been seen in recent years (Nakane, pers. comm.). In Kogoshima, Nakane found adults in abundance on flowers of Viburnum, Rosa?, Prunus? etc. in April and May, but collected only one in July. They were usually captured singly or in pairs except in mid-April when as many as 45, of both sexes, were seen at a time. No larvae were found. In Central Europe Aplocnemus is found on the flowers of pine trees, but has also been collected on deciduous trees, and Trichoceble (widespread but not common) has been found on flowering shrubs and in dry wood under trees (Lohse, 1979). Aplocnemus has been recorded on oak in Spain (Constantin, 1965) and Kubanius elegans Majer (U.S.S.R.) was ‘reared in Picea schrenckiana’ (Majer, 1983). Checklist of the genera of the Rhadalinae ANTHRIBOCLERUS Schenkling, 1922: 328. ANTINEA Peyerimhoff, 1929: 191. APLOCNEMUS Stephens, 1830: 316. Elicopis Stephens, 1829: 136. Haplocnemus Stephens; Agassiz, 1846[7]: 29. Helicopis Stephens; Agassiz, 1846[7]: 29. Subgenus APLOCNEMUS Stephens, 1830: 316. Subgenus DIPLAMBE Schilsky, 1894: 234. Subgenus HOLCOPLEURA Schilsky, 1894: 234. Subgenus ISCHNOPALPUS Schilsky, 1894: 235. Subgenus PSEUDAPHYCTUS Pic, 1896: 47. EUCYMBOLUS Champion, 1913: 129. HEMIPLEURUS gen. n. INDIODASYTES Pic, 1916: 14. KUBANIUS Majer, 1983: 385. MALTHACODES Waterhouse, 1876: 116, Xamerpus Fairmaire, 1886: 41. Syn. n. Donaldia Alluaud, 1898: 102. MICROJULISTUS Reitter, 1889a: 111. Ceralliscus Bourgeois, 1894: 121. PELECOPHORA Deyjean, 1821: 115. Diglobicerus Latreille, 1829: 475. RHADALUS LeConte, 1852: 212. Cymbolus Gorham, 1886: 324. Syn. n. REVIEW OF THE RHADALINAE 133 SEMIJULISTUS Schilsky, 1894: 227. Celsus Lewis, 1895: 118. Syn. n. Eurelymis Casey, 1895: 600. Syn. n. TRICHOCEBLE Thomson, 1859: 109. Julistus Kiesenwetter, 1859: 175. Notes on the key and checklists of species The measurements of approximate body length in the descriptions are taken from the front of the pronotum to the apex of the elytra, since in many genera the head is retractable under the pronotum and hence only partially visible from above. The term ‘free’ when describing the membranous appendages on the claws means that they are attached to the claws at the base only (Fig. 15). In most dasytines possessing these structures, the appendage is attached to the claw along its length almost to the apex, or for at least half its length (Blaisdell, 1938: pl. 2, figs 1-20). A ‘rimmed’ puncture means that there is a circle around the puncture but the area within this is not raised above the surrounding surface (Fig. 1). A ‘tuberculate’ puncture is within a circular area which is raised above the surrounding surface (Figs 21, 22, 24). In the species checklists it should be noted that the species of earlier authors are taken from the Coleopterorum Catalogus (Pic, 1926, 1929, 1937) in order to complete the lists and have not in all cases been examined. However, the references have been checked and many errors have been rectified. The varieties cited in Pic’s catalogue have been listed here in synonymy. The generic names in parentheses denote the genus in which the species was described, if not the current genus. Abbreviations BMNH _ British Museum (Natural History), London. MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. MNHN Muséum national d’Histoire naturelle, Paris. USNM National Museum of Natural History [U.S. National Museum], Smithsonian Institution, Washington, D.C. [Note. See Acknowledgements for full list of institutions from which material was borrowed. | Key to adults of the world genera of the Rhadalinae 1 Epipleura very narrow, usually mat or granular, and deflecting laterad at, or before, level of hind coxae (Figs 3, 4), becoming subvertical. Pubescence of dorsum uniformly short and recumbent, sometimes scale-like (Figs 1, 2). Tarsi elongate and slender (Figs 3, 5), claws with small teeth, without membranous appendages. Antennae fairly short, sometimes with segments 7-10 enlarged, compactly serrate (Fig. 1). Length not more than5 mm. .............. 2 — Epipleura broader, usually shiny, deflecting dorsad either at about level of 2nd or 3rd visible abdominal sternite (Figs 8, 17, 35) or at or before base of 1st visible abdominal sternite (Fig. 9). Pubescence of dorsum not as above, nearly always with some erect or semi-erect setae. Tarsi variable. Antennae elongate, reaching base of pronotum or beyond and usually loosely serrate or pectinate from segments 4—10 (rarely 3—10) (Figs 40—44). Length 2-7 mm........... 3 2 Epipleura evanescent anterior to lst abdominal sternite (Fig. 4). Lateral pronotal margin smooth, or with a few small teeth near base. Pronotum mat, puncturation indistinct, with strong microsculpture, forming an irregular reticulum under pubescence (Fig. 2). Disc of metasternum flat, with distinct median longitudinal groove (Fig. 4). Females without unpunctured patch on apical third of elytron. Length less than 3 mm. (C. Europe & Asia, Peay WADA, WMICCHECTLANEAN) «. ooocecccvacsvenssacdessvacecutsdcaaevecensces MICROJULISTUS(p. 161) — Epipleura narrow and becoming subvertical from level of 1st abdominal sternite, visible in side view from just behind shoulder. Lateral pronotal margin strongly serrate. Pronotum with shallow, rimmed punctures (Fig. 1), sometimes obscured by pubescence. Disc of metaster- num convex, without a distinct median longitudinal groove (Fig. 3). Females usually with shiny unpunctured patch on apical third of each elytron (Fig. 6). Length 3—5 mm. (C. & E. Eimepe, sia, Japan, western U-S.A., Canada): ......2..4.....cicceceveceees SEMIJULISTUS (p. 159) 3 Tarsi elongate and claws appendiculate, membranous appendages either absent or minute. Nea Oe SCOP pore Piel eeu ei de occ saucaeusustccdeseensdebsdeleagueneh ici ate ats aaeennene 4 134 E. R. PEACOCK Figs 1-7 1,6, 7, Semijulistus ater (Canada): (1) pronotum; (6) elytral apices of female; (7) aedeagus of male. 2, 4, Microjulistus fulvus (U.S.S.R., Kazakhstan): (2) pronotum; (4) metasternum. 3, 5, Semijulistus spectabilis (Japan): (3) metasternum; (5) hind tarsus. (Scale line = 0-3 mm.) da = dorsal appendage; ml = median lobe.) REVIEW OF THE RHADALINAE 135 "ods i * j Nile eS alot eae Dee 4 4 bg Vers Ax ee £ 3 r; PGE ; 3 Pe es a) tT AP Ze. ‘ A i Pa a! alas 4, a — A Fi all ae i pe lias : / Pond 1 gh pe eae, Figs 8-15 8, Malthacodes pictus (Rodriguez I.): epipleuron. 9, Hemipleurus floriger (Sarawak [para- type]): epipleuron. 10, 11, Kubanius elegans (U.S.S.R., Kazakhstan [paratype]): (10) hind tarsus; (11) hind claw. 12, 13, Trichoceble floralis (France): (12) hind tarsus; (13) claw. 14, Aplocnemus impressus (England): hind claw. 15, Pelecophora illigeri, (Mauritius): hind claw. (Scale line = 0-1 mm.) ma = membranous appendage; ac = appendiculate claw. 136 E. R. PEACOCK — Tarsi stout or delicate, claws not appendiculate, but with large free membranous appendages (Figs: 14515, 45247) ooo. askew gens ities sce tra cd geal Se Ga nena ical ane eRe ee ee 6 4 Eyes small, oval, not projecting beyond contour of head. Head weakly convex between eyes. Temples long, subparallel. Claws with minute teeth. Apterous, without humeral swellings. (N. Afnica: Salata) 2.0, 25 ot tin occa atte. «eee ae une eee ae eee oe ANTINEA (p. 159) — Eyes large, reniform, prominent, projecting beyond contour of head when seen from above. Head sometimes with a short, longitudinal groove or depression at inner side of antenna and eye. Temples constricted behind eyes. Claws with strong teeth. Winged, elytra with distinct humeral swellitigs! ¢..22:3j0c2) Lacon. ceset oaniecel, incor qeptencteeeiis aeone ste econ ee 5 5 Dorsum without metallic sheen. Hard-bodied. Pronotum usually elliptical, broader than head (including eyes). Elytra and pronotum convex and densely punctured; epipleura long, extending to about 2nd visible abdominal sternite. Hind tarsus not quite as long as tibia. Claws with either pointed (Figs 12, 13) or blunt teeth. (C.,S. & E. Europe, Asia, China.) TRICHOCEBLE(p. 158) — Dorsum with slight metallic sheen. Soft-bodied, malacodermiform. Pronotum subquadrate or elliptical, sometimes flattened on disc, not always broader than head. Elytra flattened on disc, apices abruptly declivous, rather sparsely punctured; epipleura short, not reaching hind coxae. Hind tarsus about as long as tibia (Fig. 10). Claws with large, blunt teeth, sometimes also with small, obscure membranous appendages (Fig. 11). (U.S.S.R. (Kazakhstan), N. Wadia, )c. 0. een cahint haces. ee KUBANIUS(p. 7) 6 Elytra with sublateral carina in basal half, forming a double elytral edge (Fig. 16). ................. — Elytra without sublateral carina im basalNalf (Fig. 17)... <2 22. eso. cc .-+-0-ce0e--sceeuee ete eee ; 7 Pronotal punctures all simple. Apical segment of maxillary palps only weakly securiform (Fig. 31). Form elongate, elytra more than twice as long as wide. Dorsum usually black, sometimes with a metallic sheen. (Greece.) ................+. APLOCNEMUS (DIPLAMBE)(p. 151) — Pronotal punctures tuberculate, at least near sides. Apical segment of maxillary palps triangular. Form elongate and dorsum usually unicolorous brown, or form very broad (length less than twice width), head brown, elytra and pronotum black. .......................008 8 8 Form elongate. Elytra and pronotum brown. Scutellum mat, densely and finely punctate and densely setose (Fig. 18). Sublateral elytral carina close and subparallel to costal margin, forming an abrupt, narrow, horizontal shelf, with a row of very large punctures along inner side (Fig. 16). Disc of pronotum with strongly rimmed or tuberculate punctures, sometimes with raised, shiny puncture-free patches (Figs 21, 22). (Southern U.S.A.,C. & S. America.) RHADALUS(p. 144) — Formvery short and broad (see couplet 7). Elytra and pronotum bluish-black. Scutellum shiny, very sparsely punctate and setose. Sublateral carina oblique to costal margin and less prominent, so scarcely forming a shelf, without such large punctures along inner side. Pronotum with small simple punctures on disc and tuberculate punctures near sides. (C. AMMO TiCa ays oes... emia ieee eek ee Re See EUCYMBOLUS(p. 145) 9 Humeral swellings very prominent and elytra with a pronounced transverse depression across basal third. (Pigs 29. 30) 3 282 ecteeio Re e 10 — Humeral swellings less prominent and elytra without a transverse depression across basal third. sinashacs ne aQnalt met Be eelirb Saute ale arid Sgt ea ena aieto 5 ore OMRON 5 Ri RNG dee eaetce te UR ae 11 10 Pronotum very large and quadrate, about half as long as elytra; shiny, with sparse setae, puncturation inconspicuous (Fig. 27). Head large, shiny and convex with faint diagonal ridges from base of antenna to eye (Fig. 25). Elytra with semi-erect shorter setae between the sparse, very long, erect setae; tufts of dense setae present on weak tubercles, just posterior to transverse elytral depression, and also on larger tubercles at inner side of humeral swellings (Fig. 29); epipleura extending to base of 2nd visible abdominal sternite. (Seychelles. ) ANTHRIBOCLERUS(p. 145) — Pronotum of moderate size, transverse, much less than half as long as elytra; with distinct puncturation (Fig. 28). Head mat, without diagonal ridges (Fig. 26). Elytra with recumbent setae between sparse, long, erect setae (Fig. 30); epipleura not extending further than base of 1st visible abdominal sternite (Fig. 9). (Borneo.) ..................0c000e HEMIPLEURUS(p. 147) 11 Pronotal punctures simple (Fig. 19). Maxillary palps moderately securiform (Fig. 31) or spindle-shaped (Fig. 32). Dorsum usually brown or black, often with a metallic bluish or greenish tinge (some species from the Canary Is are light brown or yellow). Setae on elytra usually of fairly uniform length and colour. (Palaearctic, N. & W. Africa (Angola? & Ivory Coast), Asia Minor, Philippines.) 1.0... <...0.0 deebe ee ee APLOCNEMUS(p. 151) 137 REVIEW OF THE RHADALINAE , “ns vil scutellum. (Scale line = lateral view of body. 18, Rhadalus rufopiceus madurensis (India [holotype]) Figs 16-18 16, Rhadalus rufopiceus (Guatemala [syntype]): lateral view of elytron. 17, Indiodasytes 0-4 mm.) 138 E. R. PEACOCK Figs 19-24 Pronotum of: (19) Aplocnemus impressus (England); (20) Pelecophora illigeri (Mauritius); (21) Rhadalus rufopiceus (Guatemala [syntype]); (22) Rhadalus testaceus (U.S.A.); (23) Trichoceble floralis (France); (24) Indiodasytes madurensis (India [holotype]). (Scale line = 0-5 mm.) REVIEW OF THE RHADALINAE 139 26 mer - Figs 25-30 25, 27, 29, Anthriboclerus scotti (Seychelles [paratype]); (25) part of head; (27) pronotum; (29) basal half of elytra. 26, 28, 30, ditto, Hemipleurus floriger (Sarawak [paratype]). (Scale line = 0-1 mm in Figs 25, 26; = 0-3 mm in Figs 27—30.) d/ = diagonal line. 140 E. R. PEACOCK Swe y A blak og | Figs 31-35 31-33, apical segment of maxillary palp of: (31) Aplocnemus nigricornis (England); (32) Aplocnemus (Ischnopalpus) sanctus (Palestine); (33) Pelecophora illigeri (Mauritius). 34, 35, ventral view of prosternum of: (34) Pelecophora illigeri; (35) Malthacodes vageguttatus (Madagascar). (Scale line = 0-1 mm in Figs 31-33; = 0-5 mm in Figs 34-35.) Pronotal punctures tuberculate or rimmed at sides. Maxillary palps broadly triangular (Fig. 33). Dorsum brown or black or bicolorous and patterned. Elytral setae not usually uniform, sometimes both erect and semi-erect, or with patches of recumbent setae, often bicolorous. iV: Head very large, not retractable under pronotum. Prosternum long in front of coxae; prosternal process well developed (Fig. 34). Pronotal punctures usually simple on disc and tuberculate at sides, sometimes becoming weakly tuberculate on disc also, which then appears rugose (Fig. 20). Elytra usually with short and long, erect or suberect setae, often forming patches. Cuticle or setae bicolorous. (Mauritius, Réunion, E. Africa.) PELECOPHORA (p. 146) Head smaller, somewhat vertical, partly retractable under pronotum. Prosternum shorter in front of coxae; prosternal process small and pointed (Fig. 35). Pronotal punctures rimmed or tuberculate on disc, or not clearly defined, but never simple. Cuticle either bi- or unicolor- ous; setae variable in-lenpth, type and colours es o.oo cee ene ee 13 12 REVIEW OF THE RHADALINAE 141 Figs 36-39 Elytral puncturation of: (36) Rhadalus rufopiceus (Guatemala); (37) Hemipleurus floriger (Sarawak); (38) Semijulistus ater (Canada); (39) Trichoceble floralis (France). (Scale line = 0-1 mm.) 13. Tarsi stout, hind ones longer than half a tibia, dark brown, without very long ventral setae, segments 1—4 subequal, penultimate segment apically truncate dorsally (Fig. 45). Form elongate. Dark brown and/or black. ................. Be ea eee petted eanresctmante ena tooes a 14 — Tarsi delicate, hind ones usually about half tibial length or less, usually pale coloured, segments 1-3 subequal sometimes with very long ventral setae, segment 4 smaller, dorsally apically emarginate (Fig. 46). Form either broad and short or elongate. Unicolorous or bicolorous, often patterned. (Madagascar, Seychelles, Aldabra, Mauritius, Réunion, Rodriguez I., eM Ae PUMMCH A SEP ATCA) oe vos piede,) cca sdcweacaatediceddacecsassss MALTHACODES [in part] (p. 149) 14 Head and pronotum black, elytra dark brown. Pronotum very strongly punctured and with a distinct longitudinal unpunctured area (sometimes raised) in mid line near base. Dorsal pubescence recumbent or semi-erect, usually with a few erect setae laterally (Fig. 24). Wings with strong venation, anal cell either present or obsolete. Aedeagus without dorsal appen- CAS CURED NEL WAATIICA) Gelcbes.dccs Radial cell of fore wing open or closed (Figs 112, 129); metapleural ridges not converging nmacas propodeum, almost parallel (Fig. 130). 2 1....42. 4. savcd.cdaedees 000) bee hau eae SR eee Radial cell of fore wing closed (Fig. 140); metapleural ridges converging towards propodeum PLD LZ. ee See ee ee eee tere ee heterotoma (Thomson) (p. eens SCOTMEME 9 SMOLLED tials (E10, LAA oo ae one cciaew suannen qo ulave mses +cuatvavasesecsevaaqean ss Antennal segments 3 and 4 subequal in length (Fig. 168). Semrellar cup extendm~¢e to apex of scutellar dise (Fig. 127). ..................---- syphax sp. n. (p. Scutellar cup viewed dorsally not extending to apex of scutellar disc, disc reticulate-rugose errr TONOtAl Mate (EL EIS. 1IZ).. 5.x... cenaceccccsonseqacecatecstestaneqesces ap fannius sp. n. (p. Scutellar cup viewed dorsally not extending to apex of disc, with radiating striae; pronotal plate (Fig. 166). Propodeal carinae converging; radial cell of fore wing open. ............... misensus sp. n. (p. Metapleuron without ridges in upper region; propodeal carinae closer together apically (Fig. 153); radial cell of fore wing closed on wing margin. boulardi (Barbotin, Carton & Kelner-Pillault) (p. Metapleuron with ridges in upper region; propodeal carinae variable; radial cell of fore wing Sa mm UM TC URGED UM NMEAE ONE 02050 a6 sana ventas etn agro aan ates shge's ang canac ser avn aappennennemses Scutellar cup oval (cf. Fig. 148); segment 3 of antenna shorter than 4. tT 02 OL Fata Cel MOE INGICALER (PID. LOG). .ctcccsesccceccasneetecsercscavertecsiegenacersaues Scutellar cup elongate; segment 3 of antenna shorter than or subequal to 4. .......................68. Antennal segments 3 and 4 slender, curved (cf. Fig. 133); scutellar disc reticulate-rugose, MPRCRMCERTEMIEY [AROS OM RUD, ADIs - ce cies voscd angne sane sh ¢gnesscunmmeneas tains vanes use faunus sp. n. (p. Antennal segments 3 and 4 robust and swollen (Fig. 133); scutellar disc with radiating strigose SeUIUELe. Scutcliar Cup stial (Cl. Fig. 148), . oc: ccnnscccssavasdenssenctaours mahensis (Kieffer) (p. Antennal segments 3 and 4 subequal in length (Fig. 132) (4 weakly swollen). ..... vesta sp. n.(p. PRM ee SCOIMCIL 5 GISEMICULY SMUERGR CG (PIG. 147), oc .scnswesccesccenesecnshecverneonsuverqucseseasss Radial cell of fore wing with vein Rs + M in form of a distinctive spur (Fig. 146); antennal segment 4 swollen medially and curved, outer side flattened (Fig. 147). victoriae Nordlander (p. 193 14 198) 199) 194 J. QUINLAN — Radial cell of fore wing without a distinctive spur (Fig. 150); antennal segment 4 weakly swollen medially and curved, outer side flattened. Pronotal plate-(FigwiS9)n. 28 2.4 cod BER Be Aes Re Re A eee 13 13 Scutellar disc reticulate-rugose (Fig. 148), rim of scutellar cup thick, not converging apically; metapleural mdgesjpanaliel.c (52.1 307.) BAe 2 POs GA Vee apellasp.n.(p. 194) — Scutellar disc coarsely rugose-reticulate, scutellar cup converging apically (cf. Fig. 124); metapleural ridges weakly converging towards propodeum. .................... pisonis sp. n. (p. 198) Leptopilina apella sp. n. (Figs 119, 148-150, 159) DEscrIPTION. 9 antenna weakly clavate, segments 6—13 with rhinaria, segment 3 shorter than 4 (Fig. 119); CO antenna with segments 3-4 curved (cf. Fig. 147). Eyes large, converging towards clypeal area, closer together than the height of an eye where they converge; malar grooves fine. Pronotal plate with medial bridge narrow, as wide as a fovea (Fig. 159). Lateral bars of scutellum broad-based, triangular; scutellar fovea smooth and shining, shallow, lenticular; scutellar cup small, oval, not extending to apex of disc; scutellar disc reticulate-rugose, apex rounded (Fig. 148). Mesopleural suture converging towards meta- pleuron with two ridges in upper region (cf. Fig. 167); anteroventral cavity pubescent; lateral margins of propodeum densely pubescent on under margins of carinae, weakly sculptured; propodeal carinae parallel (Fig. 149). Segment 1 of gaster in form of short crenulate ring; segment 2 with tuft of pubescence basally on lateral margins; segment 3 partially visible; gaster impunctate. Radial cell of fore wing closed on wing margin, broad apically; vein 3r—m produced; vein M indicated basally, weakly pigmented (Fig. 150). Legs long and slender; hind coxa with tuft of hairs on posterodorsal margin. Colour: antenna yellow basally, segments 6—13 blackish; head blackish; thorax chestnut brown; gaster brownish yellow; legs yellow. MATERIAL EXAMINED ) Holotype 9 , Madagascar: Tam, Perinet, 27.iv.—3.v.1983 (Noyes & Day) (BMNH). Paratypes. Madagascar: 1 2,3 CO’ (BMNH). Zaire: 8 9 (MRAC). Leptopilina atraticeps (Kieffer) (Figs 122, 123, Toa) Ectolyta atraticeps Kieffer, 1911: 312. Holotype 9 , SEYCHELLES (BMNH) [examined]. DESCRIPTION. 9 antenna clavate, segments 7-13 forming a club, rhinaria weakly indicated, each segment more than 2 times as long as broad, segment 3 slightly longer than 4, less than 4+5, 4 longer than 5. Eyes oval, as far apart measured medially as the height of an eye; subantennal sutures absent. Pronotal plate projected forward; anterior part finely sculptured; lateral foveae open on either side of medial bridge, wide (Fig. 151). Mesoscutum smooth and polished, variable surface sculpture indicated through the chitin (Fig. 123); lateral bars of scutellum smooth and polished; scutellar foveae deep, polished; scutellar cup elliptical, widest medially, with a wide rim, with a large apical fovea; scutellar disc reticulate-rugose, rounded apically. Mesopleuron smooth and polished; mesopleural suture distinct, curved; metapleuron smooth and polished with 2 conspicuous ridges; anteroventral cavity obscured; sides of propodeum with dense short pubescence; propodeum almost parallel, very weakly bowed medially. Segment 1 of gaster in form of crenulate ring, wider than long; segment 2 with a dense ring of short hairs basally not completed on dorsal surface; tergites 3 and 4 partially visible. Wings not conspicuously narrow; pubescent with long apical hair fringe; radial cell of fore wing partially open on wing margin; veins Rs + M and M weakly indicated (Fig. 122). Legs slender; hind coxa with a tuft of pubescence postero-dorsally. Colour: antenna yellowish orange, some median segments light brown; head reddish brown; thorax and gaster orange- yellow; legs orange-yellow. CO unknown. MATERIAL EXAMINED Holotype 9, Seychelles: Mahe I., Mare au Cochons (BMNH). Paratype. Nigeria: 1 9 (BMNH). REMARKS. Similar in many respects to faunus, but differs in the shape of the radial cell and the scutellar cup. AFROTROPICAL EUCOILIDAE 195 Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault) (Pages, 12d Loe. has) Charips mahensis Kieffer, 1911: 313. Holotype &’, SEYCHELLES (BMNH) [examined]. [Junior secondary homonym in Leptopilina of Erisphagia mahensis Kieffer (see p. 198).] [Synonymised by Nordlander, 1980: 432. ] Cothonaspis (Cothonaspis) Boulardi Barbotin, Carton & Kelner-Pillault, 1979: 20. Holotype 9, GUADE- LOUPE (MNHN) [not examined]. DEscRIPTION. Q antenna clavate, the 4 or 5 apical segments forming a club, each of club segments with rhinaria, each a little longer than broad, generally darker than the yellow basal segments, segment 3 as long as 4+5, 4 longer than 5, 5 and 6 subequal in length, 7 longer than 6 (Fig. 115); Ch antenna 15-segmented, filiform, segment 3 shorter than 4, 4 curved, swollen, longer than 5. Eyes almost round, further apart than the height of an eye measured medially; subantennal sutures prominent on lower face, sparse hairs present; malar grooves percurrent. Pronotal plate projected forward; lateral foveae long and narrow (Fig. 152). Mesoscutum smooth and polished, notauli absent; lateral bars of scutellum smooth and polished; scutellar foveae shallow; scutellar cup long and narrow, widest medially, apex weakly converging, apex with a large fovea, area in front weakly convex, sculptured; scutellar disc with radiating ridges, apex of disc rounded. Mesopleuron smooth and polished; mesopleural suture distinct, curved; metapleuron with one or two incomplete ridges (Fig. 121); nucha short, weakly pubescent; propodeal carinae closer together in upper half, bowed medially. Segment 1 of gaster crenulate, in form of a crescent; segment 2 widened dorsally, hairy ring at base very thin, only a few hairs present laterally; visible parts of segments 3—5 punctate. Fore wings not conspicuously narrow, pubescent, apical hair fringe long; radial cell of fore wing closed on margin; 2r—m distinct; Rs + M and M weakly indicated. Legs robust; hind coxa without a tuft of hairs postero-dorsally. Colour: apical segments of antenna brownish black, basal segments yellow; head, thorax and gaster brownish black-chestnut; legs yellow. MATERIAL EXAMINED Charips mahensis Kieffer, holotype OC’, Seychelles (Percy Sladen Trust Expedition) (BMNH). Aldabra: 1 2 (BMNH). Guadeloupe: 1 2, 1 o’ (BMNH). Madagascar: 4 9 (BMNH). South Africa: 10 (BMNH). Zaire: 32 2, 33 h (MRAC). Zambia: 4 9 (BMNH). Zimbabwe: 2 9, 1 h (BMNH). REMARKS. Separated from all other species in the genus by the absence of metapleural ridges, form of antenna and scutellar cup. It is a solitary primary internal parasite of Drosophila melanogaster Meigen (see Nordlander, 1980). Leptopilina clavipes (Hartig) (Figs 110, 136) Cothonaspis clavipes Hartig, 1841: 357. REMARKS. This species is included in the key for comparative purposes. Nordlander (1980) gives a description and synonymy. At present clavipes is known to occur in Europe and North America but it has not been found in the Afrotropical region. Leptopilina fannius sp. n. (Figs 120, 143, 144, 154, 155, 156) DescriPTIon. 9 antenna filiform, segments 5—13 with rhinaria, segment 3 shorter than 4, subequal to 5, apical two segments pale (Fig. 120); C’ antenna 15-segmented, filiform, segment 3 shorter than 4, weakly curved (Fig. 144). Eyes oval, as far apart measured medially as the height of an eye; face smooth and polished; malar grooves percurrent. Pronotal plate weakly protruding; medial bridge broad (cf. Fig. 159). Mesoscutum smooth and polished; lateral bars of scutellum broad basally, short; scutellar cup longer than wide, not extending to apex of disc, widest medially, converging apically (Fig. 143); scutellar disc reticulate-rugose laterally, rounded apically (sometimes the margins are striated). Mesopleuron smooth and polished; mesopleural suture almost straight; metapleuron with three distinct ridges, the upper ridge branched (Fig. 154); anteroventral cavity large, hairless; lateral margins of propodeum crenulate, sparsely pubescent; propodeal carinae bowed (cf. Fig. 161). Segment 1 of gaster in form of crenulate ring, wider than long; tergite 2 with a few sparse hairs on lateral margins of base, apical half densely punctate; visible parts of segments 3 and 4 punctate. Fore wing relatively broad, rounded apically, surfaces pubescent, apical margins with a fringe of hairs; radial cell of fore wing closed on margin, weakly pigmented, broad 196 J. QUINLAN apically; vein 2r—m not projecting, merging into vein M; vein Rs + M not indicated (Fig. 155). Legs robust; hind coxa stout, a few hairs present on postero-dorsal margin. Colour: antenna yellow basally, dark medially, apical two segments pale yellow; head and thorax black, except metapleuron orange-brown; legs orange-yellow. MATERIAL EXAMINED Holotype 9, Zaire: Massif Ruwenzori, Kyandolire, 1700 m, Camp des Gardes, 7—15.x.1952 (Vanschuyt- broeck & Kekenbosch) (MRAC). Paratypes. Zaire: 1 9,6 C (MRAC). REMARKS. Separated from vesta by the relative proportions of the antennal segments and the form of the scutellar cup. Leptopilina faunus sp. n. (Figs 157, 158) DEscrIPTION. 9 antenna clavate, segments 7—13 with rhinaria, segment 3 longer than 4, 5 and 6 subequal, each shorter than 4 (cf. Fig. 156); O’ antenna 15-segmented, segment 3 shorter than 4, both slender and curved (cf. Fig. 137). Eyes almost round, further apart measured medially than the height of an eye; malar grooves percurrent; face smooth and polished, a few sparse hairs present in clypeal region. Pronotal plate weakly protruding; lateral foveae open (but not readily appreciated); medial bridge broad (cf. Fig. 151). Mesoscutum smooth and polished; lateral bars of scutellum broad basally, longer than wide; scutellar foveae lenticular, smooth and shallow; scutellar cup elongated, not extending to apex of disc, widest medially, tapering apically, apical fovea large; scutellar disc reticulate-rugose, rounded apically. Meso- pleuron smooth and polished; mesopleural suture weakly curved; metapleuron with two strong ridges in upper quarter converging towards margins of propodeum (Fig. 157); anteroventral cavity pubescent; lateral margins of propodeum sculptured, partially covered with pubescence; propodeal carinae widely spaced, weakly bowed. Segment 1 of gaster in form of crenulate ring; tergite 2 with a few hairs at lateral margins of base, apical margins sparsely punctate; segments 3 and 4 partially visible, punctate. Fore wing broad, wing surfaces pubescent with apical hair fringe; radial cell of fore wing partially open on wing margin; vein 2r—m weakly projecting; veins M and Rs + M weakly indicated by pigmentation (Fig. 158). Legs slender; hind coxa with a few hairs on posterodorsal margin. Colour: antenna orange-yellow basally, brownish yellow apically, apical segment lighter than rest; head and thorax brown, metapleuron orange- brown; gaster chestnut brown; legs orange-yellow. MATERIAL EXAMINED Holotype 9, Zaire: Nyasheke (Volc. Nyamuragira), 1820 m, 14—26.vi.1935 (de Witte) (MRAC). Paratypes. Zaire: 3 2,5 CO’ (MRAC). RemMARKS. Distinguished from atraticeps by the shape of the scutellar cup and radial cell. Leptopilina fimbriata (Kieffer) (Figs 111, 116, 135, 137, 138) Eucoela fimbriata Kieffer, 1901: 175. REMARKS. This species is included in the key for comparative purposes. Nordlander (1980) gives a full synonymy, description and bionomic information. It has not been found outside Europe. Leptopilina heterotoma (Thomson) (Figs 124, 126, 140, 160) Eucoila heterotoma Thomson, 1862: 403. Holotype 2 , SWEDEN (ZI) [examined]. Ganaspis monilicornis Kieffer, 1904: 622. Lectotype 2, FRANCE (MA) [not examined]. [Synonymised by Nordlander, 1980: 430. ] Ganaspis subnuda Kieffer, 1904: 64. Holotype ’, Canary Is (not located). [Synonymised by Nordlander, 1980: 430.] Erisphagia philippinensis Kieffer, 1916: 282. Lectotype 9 , Los BANos (MNHN) [not examined]. [Synony- mised by Nordlander, 1980: 430.] Pseudeucoila bochei Weld, 1944: 65. Holotype 9, U.S.A. (USNM) [examined]. [Synonymised by Nordlander, 1980: 461.] AFROTROPICAL EUCOILIDAE 197 DESCRIPTION. @ antenna weakly clavate, segments 3 and 4 subequal in length, 5 shorter than 4, apical 7 segments with rhinaria forming a club (Fig. 160); G& antenna 15-segmented, filiform, segment 3 shorter than 4, 4 twisted and swollen medially (Fig. 137). Face smooth and polished with sparse hairs; anterior tentorial pits minute; malar grooves finely indicated; head viewed dorsally as wide as thorax; eyes large; occipital carinae distinct. Pronotal plate projected forward; medial bridge broad; anterior plate sculp- tured; pronotum either side of plate pubescent. Mesoscutum smooth and polished without trace of notauli; lateral bars of scutellum smooth and polished; scutellar fovea deep, lenticular; scutellar disc smooth to reticulate-rugose with scattered hairs, rounded apically; scutellar cup smooth and polished, convex in front of apical pit, long, widest above medial area, converging apically and narrow (Fig. 124). Mesopleuron smooth and polished; mesopleural suture distinct, weakly curved; metapleuron depressed above ridges 1 and 3, ridges 1, 2 and 3 entire (Fig. 126); lateral margins of propodeum below metanotal plate pubescent; propodeal carinae sub-parallel, wide (cf. Fig. 150). Segment 1 of gaster in form of a narrow ring, sculptured, weakly inclined posteriorly; tergite 2 with ring of dense pubescence ventrally, not complete on dorsal surface; tergite 3 partially visible; apical margins of tergite 2 and whole of visible parts of 3 punctate (cf. Fig. 108). Wing surfaces puhescent, apical margins with fringe of hairs; radial cell of fore wing closed on margin; vein Rs as long as vein 27; vein M indicated, not pigmented (Fig. 140). Legs slender; hind coxa with distinct tuft of hairs on posterodorsal surface. Colour: antenna yellow basally, darker apically; head black; thorax and gaster dark brown-black. MATERIAL EXAMINED Eucoila heterotoma Thomson, holotype 2, Sweden: Skane, Baggeboda (ZI). Madagascar: 1 2 (MRAC). Palestine: 1 9, 1 &’ (BMNH). Zaire: 1 9 (MRAC). REMARKS. The specimens from Madagascar and Palestine are only tentatively placed as this species because the apical segments of the antenna are white. L. heterotoma is world-wide in distribution (Nordlander, 1980). The species is a solitary primary internal parasitoid of Drosophila larvae and it has been widely studied in Europe and the U.S.A. Leptopilina itys sp. n. (Figs 118, 161, 162) DESCRIPTION. 9 antenna weakly clavate, segments 7-13 with rhinaria, 4 times as long as broad, segment 3 shorter than 4, 4—6 subequal in length (Fig. 118). Eyes large, almost round, further apart measured medially than the height of an eye; face smooth and polished with sparse scattered hairs; malar grooves very fine, percurrent. Pronotal plate protruding; medial bridge as wide as a lateral fovea. Mesoscutum smooth and polished; lateral bars of scutellum broad-based, triangular; scutellar fovea wider than long, shallow; scutellar cup elongate, not extending to apex of disc, with a large apical fovea; scutellar disc finely reticulate-rugose, apex rounded (Fig. 162). Mesopleuron smooth and polished; mesopleural suture almost straight; metapleuron with two ridges in upper region, parallel, widely spaced (cf. Fig. 131); anteroventral cavity large, pubescent; lateral margins of propodeum sculptured, partially concealed by pubescence; propodeal carinae closer together anteriorly (Fig. 161). Segment 1 of gaster in form of a crescent-shaped crenulate ring, partially visible; lateral margins of base of tergite 2 with a ring of hairs, thin dorsally, thicker ventrally; tergites 3-5 partially visible in lateral view, sparsely punctate. Fore wing relatively narrow compared with other species in the genus, wing surfaces pubescent with apical hair fringe; radial cell of fore wing closed on wing margin; vein 2r—m weakly protruding; veins M and Rs + M not indicated. Legs long and slender; hind coxa with a few short hairs on posterior dorsal margin. Colour: antenna yellow basally, apical 5—6 darker; head, thorax and gaster dark brownish black; legs yellow. CO’ unknown. MATERIAL EXAMINED Holotype 2, Zimbabwe: Salisbury, Chishawasha, iii. 1980 (Watsham) (BMNH). Paratype. Zaire: 1 9 (MRAC). REMARKS. The length of the antennal segments, shape of the scutellar cup and the sculpture of the scutellar disc separate this species from thetus. Leptopilina longipes (Hartig) (Figs 106-109, 113, 139) Cothonaspis longipes Hartig, 1841: 356. REMARKS. This European species is included in the key for comparative purposes. It is the type species of the genus. Nordlander (1980) gives a description. 198 J. QUINLAN Leptopilina mahensis (Kieffer) (Figs 133, 163, 164) Erisphagia mahensis Kieffer, 1911: 312. Holotype C’, SEYCHELLES (BMNH) [examined]. Description. 9 unknown. CO antenna 15-segmented, filiform, segment 3 shorter than 4, weakly swollen apically, 4 curved and swollen medially (Fig. 133). Face smooth and polished; eyes large, protruding; anterior tentorial pits distinct; malar grooves percurrent but finely indicated; a few sparse hairs present; head viewed dorsally as wide as thorax; occipital carinae pronounced. Pronotal plate projected forward: posterior plate angled; medial bridge narrow; lateral foveae partially open (cf. Fig. 149). Mesoscutum smooth and polished, without trace of notauli or hairs in their place; lateral bars of scutellum polished; scutellar foveae large, deep and smooth; scutellar disc rounded apically, surface reticulate-rugose; scutellar cup almost round apically, narrower basally with a pronounced r m, apical third with a large pit or fovea (cf. Fig. 148). Mesopleuron smooth and polished; mesopleural suture percurrent, almost straight; metapleuron polished and strongly ridged in anterodorsal corner; anteroventral cavity sculptured; propodeum not elongated in lateral view; nucha distinct; viewed dorsally propodeal carinae almost parallel, weakly bowed medially (Fig. 163). Segment 1 of gaster in form of ring or collar, crenulate, widened posteriorly; tergite 2 occupying the whole area of gaster in lateral view with a few hairs basally on lateral margins. Wing surfaces pubescent with apical hair fringe long; radial cell of fore wing closed, longer than wide; veins Rs + M and M absent (Fig. 164). Legs of normal shape and proportions; coxae with sparse hairs, hind coxa without a tuft of hairs. Colour: antenna orange-yellow; head and thorax light brown; gaster chestnut red; legs yellow. MATERIAL EXAMINED Holotype ©’, Seychelles: Mahe I., Mare Cochons district, 1000—2000 ft, 26.i1.—2.11.1909 (BMNH). REMARKS. Because the female is unknown the affinities of this species are uncertain. The male is separated from faunus by antennal, scutellar disc and radial cell characters. Leptopilina misensus sp. n. (Figs 114, 141, 165-167) DEscriPTION. 9 antenna filiform, segments 5—13 with weak rhinaria, segment 3 shorter than 4, 4 subequal to 5, 6-13 subequal, apical segments lighter than rest of antenna (Fig. 114); CG’ antenna 15-segmented, filiform, segment 3 shorter than 4, 4 weakly swollen medially, longer than 5 (Fig. 165). Eyes almost round, further apart than the height of an eye measured medially; face smooth and polished; malar grooves percurrent. Pronotal plate weakly projecting; lateral foveae long and narrow (Fig. 166). Mesoscutum smooth and polished, without trace of notauli; lateral bars of scutellum smooth and polished; scutellar foveae broad basally; scutellar cup longer than wide with apical fovea widest medially, tapering or converging apically; scutellar disc with aberrant reticulate-rugose sculpture, rounded apically. Meso- pleuron smooth and polished; mesopleural suture distinct, weakly curved; metapleuron with 2 ridges, almost parallel; anteroventral cavity large, hairless (Fig. 167); lateral margins of propodeum crenulate; propodeal carinae parallel (cf. Fig. 163). Segment 1 of gaster in form of crescent-shaped crenulate ring; tergite 2 occupying almost the whole visible area in lateral view, with a few hairs on the lateral margins basally; tergites 3 and 4 partially visible; apex of tergite 2 and visible parts of 3 and 4 punctate. Fore wing relatively broad, apical margins rounded, with a fringe of hairs, wing surfaces pubescent; radial cell of fore wing open on wing margin, broad apically; vein 2--m weakly projecting; Rs + M not indicated (Fig. 141). Legs slender; hind coxa without a tuft of hairs posterodorsally. Colour: antenna dark yellow basally, brownish medially, apical segments sometimes lighter; head, thorax and gaster brownish yellow. MATERIAL EXAMINED Holotype 9, Zaire: Nyasheke (volc. Nyamuragira), 1820 m, 14—26.[?month].1935 (de Witte) (MRAC). Paratypes. Zaire: 3 9, 18 O&' (MRAC). Uganda: 2 O (BMNH). REMARKS. This species has characters in common with heterotoma in the scutellar cup, but the antennal characters and open radial cell enable it to be recognised easily. Leptopilina pisonis sp. n. (Figs 129, 130) DESCRIPTION. 9 antenna clavate, segments 7—13 with rhinaria, swollen medially, segment 3 longer than 4, 4 longer than 5, 5 and 6 subequal (cf. Fig. 156); CG antenna filiform, segment 3 shorter than 4, 4 swollen AFROTROPICAL EUCOILIDAE 199 medially (cf. 137). Eyes oval, further apart measured medially than the height of an eye; face smooth and polished; malar grooves percurrent. Pronotal plate weakly protruding; medial bridge as wide as a lateral fovea (cf. Fig. 159). Mesoscutum smooth and polished, without trace of notauli; lateral bars broad, short and triangular; scutellar foveae lenticular, shallow; scutellar cup elongate, broadest medially, weakly converging apically, not extending to apex of disc; scutellar disc with large reticulate-rugose sculpturing, apex rounded (cf. Fig. 124). Mesopleuron smooth and polished; mesopleural suture weakly curved; metapleuron weakly ridged in upper quarter (Fig. 130); anteroventral cavity pubescent; lateral margins of propodeum partially obscured by dense pubescence, visible part crenulate; propodeal carinae weakly bowed (cf. Fig. 163). Segment 1 of gaster in form of crescent-shaped crenulate ring; lateral margins of base of tergite 2 with dense pubescence ventrally, thin dorsally; tergites 3-6 partially visible in lateral view; apical margin of tergite 2 and whole of visible parts of 3—6 punctate. Fore wing moderately broad, wing surfaces pubescent with apical hair fringe; radial cell of fore wing partially open on wing margin; vein 2r—m not projecting; veins M and Rs + M not indicated (Fig. 129). Legs long and slender; hind coxa with a tuft of hairs on posterodorsal margin. Colour: antenna pale yellow basally, darkish medially, pale yellow apically; head, thorax and gaster orange-brown; legs pale yellow. MATERIAL EXAMINED Holotype @, Zaire: Eala, v.1935 (Ghesquiere) (MRAC). Paratypes. Zaire: 30 9,8 CO (MRAC). REMARKS. Similar to syphax but differing in the shape and size of the scutellar cup, the radial cell and the colour pattern. Leptopilina syphax sp. n. (Figs 127, 128, 134, 142, 168) DEscrIPTION. Q antenna clavate, segments 7-13 subequal in length with rhinaria, 3 and 4 subequal in length, 5 shorter than 4, 6 shorter than 5, apical three segments lighter than median segments (Fig. 168); antenna 15-segmented, filiform, 3—5 subequal in length (Fig. 142). Eyes oval, as far apart as the height of an eye measured medially; face smooth and polished; malar grooves percurrent. Pronotal plate projecting; medial bridge broad. Mesoscutum smooth and polished, without trace of notauli; lateral bars of scutellum long, broad basally; scutellar cup longer than wide, extending to apex of disc, widest medially, converging apically (Fig. 127), apical fovea large; scutellar disc areolate-reticulate-rugose, rounded apically (Fig. 127). Mesopleuron smooth and polished; mesopleural suture almost straight; metapleuron with two ridges, weakly converging towards the propodeum; anteroventral cavity large, hairless (Fig. 134); lateral margins of propodeum canaliculate, weakly pubescent; propodeal carinae obscured by pubescence. Segment 1 of gaster in form of a crenulate ring, wider than long; tergite 2 with a few hairs on lateral margins basally; tergites 3—5 partially visible; apex of tergite 2 and visible parts of 3—5 punctate. Fore wing relatively broad, rounded apically, wing surfaces pubescent with apical hair fringe; radial cell of fore wing closed on wing margin; vein 2r—m weakly projecting; vein M complete but faintly represented; vein Rs + M not indicated. Legs slender; hind coxa without a tuft of hairs posterodorsally. Colour: basal segments of antenna orange-yellow, medial segments brownish, apical three segments light yellow; head and thorax blackish; metapleuron and gaster orange-brown; legs orange-yellow. MATERIAL EXAMINED Holotype @, Zaire: R. Kilalamatambo (affl. Lusinga), 17.vii.1945 (de Witte) (MRAC). Paratypes. Zaire: 12 9,24’ (MRAC). REMARKS. This species is closely related to pisonis but characteristics in the radial cell, scutellar cup and colour pattern enable them to be separated. Leptopilina thetus sp. n. (Figs 112, 117, 169) DESCRIPTION. 2 antenna weakly clavate, segment 3 shorter than 4, longer than 5, 5 and 6 subequal in length, 7-13 three times as long as broad, with rhinaria (Fig. 117). Eyes large, round, as far apart as the height of an eye measured medially; face smooth and polished; malar groove not visible; a few hairs present in clypeal area. Pronotal plate weakly projecting; medial bridge narrow, not as wide as a lateral fovea (cf. Fig. 159). Mesoscutum smooth and polished; lateral bars of scutellum broad basally, wide apically; scutellar foveae lenticular, smooth and shallow; scutellar cup extending almost to apex of scutellar disc, longer than wide, widest medially, converging apically (Fig. 169), apical fovea large, rim of scutellar cup 200 J. QUINLAN thick; scutellar disc with radiating reticulate-rugose sculpture, apex rounded. Mesopleuron smooth and polished; mesopleural suture weakly curved; metapleuron with two widely spaced parallel ridges in upper region; anteroventral cavity large, a few setae present; lateral margins of propodeum obscured by dense pubescence; propodeal carinae parallel. Segment 1 of gaster in form of crenulate ring, wider than long; tergite 2 with a tuft of hairs on lateral margins, thinner on dorsal surface; tergites 3—5 partially visible; apex of segment 2 and visible parts of 3—5 with sparse punctures. Fore wing moderately broad, wing surfaces pubescent with apical hair fringe; radial cell of fore wing closed on wing margin; vein 2r—m weakly projecting; veins M and Rs + M not indicated (Fig. 112). Legs short; hind coxa with a few hairs on posterodorsal surface. Colour: antenna pale yellow; head, thorax and gaster pale chestnut brown; legs orange-yellow. CO’ unknown. MATERIAL EXAMINED Holotype 9, South Africa: Port St John, Pondoland, 10—31.vii.1923 (Turner) (BMNH). Paratypes. South Africa: 2 9 (BMNH). Zaire: 3 9 (MRAC). REMARKS. This species is separated from closely related species by the form of the antenna and the elongated scutellar cup. Leptopilina vesta sp. n. (Figs 132, 145, 170) DEscRIPTION. Q antenna filiform, segments 3—13 with rhinaria, segment 3 shorter than 4, 4 and 5 subequal in length (Fig. 132); O& antenna 15-segmented, filiform, segments 3 and 4 subequal in length, 4 weakly swollen (Fig. 145). Eyes large, round, as far apart measured medially as height of an eye; face smooth and polished; malar grooves finely indicated. Pronotal plate protruding; medial bridge as broad as a lateral fovea (cf. Fig. 159). Mesoscutum smooth and polished; lateral bars of scutellum triangular, short; scutellar foveae lenticular; scutellar cup longer than wide, not extending to apex of disc; scutellar disc reticulate- rugose, apex rounded (cf. Fig. 169). Mesopleuron smooth and polished; mesopleural suture converging towards metapleuron; metapleuron with three distinct ridges in upper region (cf. Fig. 134); anteroventral cavity pubescent (cf. Fig. 134); lateral margins of propodeum partially obscured by dense pubescence; propodeal carinae bowed (cf. Fig. 163). Segment 1 of gaster in form of narrow ring, weakly crenulate; tergite 2 with a tuft of pubescence basally on lateral margins; apex of tergite 2 and visible part of tergite 3 finely punctate. Fore wing broad, apex rounded, wing surfaces pubescent with apical fringe of hairs; radial cell of fore wing closed on wing margin, narrow apically; vein 2r—m protruding; veins M and Rs + M weakly indicated, not pigmented (Fig. 170). Legs long and slender; hind coxa without tuft of hairs on posterodorsal margin. Colour: antenna yellowish basally, merging to brownish yellow apically; head and thorax dorsally blackish; mesopleuron and gaster brownish orange; legs yellow. MATERIAL EXAMINED Holotype 2, Cameroun: Nkoemvon, 1980 (Jackson) (BMNH). Paratypes. Cameroun: 1 O' (BMNH). Zaire: 2 9,8 Oh (MRAC). REMARKS. The filiform antenna separates this species from all others in this genus. Leptopilina victoriae Nordlander (Figs 125, 131, 146, 147) Leptopilina victoriae Nordlander, 1980: 447. Holotype 9 , SEYCHELLES (Nordlander coll.) [not examined]. DEscriPTION. antenna clavate, segments 7-13 with rhinaria forming a club, segment 3 longer than 4, 5 and 6 subequal, each shorter than 4 (cf. Fig. 156); O& antenna 15-segmented, segment 3 shorter than 4, 4 swollen medially, bent, outer surface flattened (Fig. 147). Eyes large, almost round, further apart measured medially than the height of an eye; face smooth and polished with sparse hairs around clypeal area; malar grooves fine, percurrent. Pronotal plate protruding; lateral foveae round, narrow; medial bridge slightly broader than a fovea. Mesoscutum smooth and polished; lateral bars of scutellum short, broad based; scutellar cup elongate, weakly converging apically with a large apical fovea; scutellar disc rounded apically, surface reticulate-rugose (Fig. 125). Mesopleural suture directed strongly down towards margin of anteroventral cavity; metapleuron with three ridges (Fig. 131); anteroventral cavity hairless; lateral margins of propodeum crenulate, partially obscured by dense pubescence; propodeal carinae very weakly bowed, almost parallel (cf. Fig. 149). Segment 1 of gaster in form of a crescent, crenulate; lateral AFROTROPICAL EUCOILIDAE 201 margins of base of tergite 2 with a thin ring of hairs and a few longer hairs behind them; tergites 3-5 partially visible, impunctate. Fore wing moderately broad apically, wing surfaces pubescent, apical margins with a hair fringe; radial cell of fore wing closed on wing margin, broad apically; vein 2r—m projected; vein Rs + M not indicated; vein M weakly indicated by pigmentation (Fig. 146). Legs slender; hind coxa swollen, with a tuft of hairs on posterodorsal margin. Colour: antenna yellow basally, apical segments darker; head, thorax and gaster dark chestnut brown; legs orange-yellow. MATERIAL EXAMINED Seychelles: 14 9, 46 CO’ (including 1 2, 1 C’ paratypes of L. victoriae Nordlander, labelled ES 551 and 552) (BMNH). Uganda: 2 2 (BMNH). Zaire: 32 9, 18 Oh (MRAC). South Africa: 1 9 (BMNH). REMARKS. Separated from vesta by antennal, pronotal plate and scutellar cup characters. Nordlander (1980) considers victoriae probably to be a parasitoid of Drosophila malerkotliana Parshad & Paika. Some of the specimens from Zaire are paler and a few have pale apical antennal segments. SIRENES gen. n. Type species: Sirenes sinis sp. n. Diacnosis. 9 antenna 13-segmented, filiform-subclavate with from 3-8 segments with rhinaria, basal segments finely granulate; O’ antenna 15-segmented, filiform-flagellate apically, segment 3 shorter or subequal to 4. Head viewed frontally longer than broad, eyes measured medially as far or further apart as height of an eye; weakly coriaceous in region of antennal sockets and eye margins; supra-clypeal area raised; face with sparse hairs; malar grooves weakly indicated with weak striations on either side; viewed dorsally ocellar area weakly coriaceous (Fig. 172), extending to occipital carina. Pronotal plate viewed dorsally rectangular (cf. Fig. 191), lateral fovea on either side weak and open; pronotum either side of pronotal plate densely pubescent. Mesoscutum polished medially, sometimes strongly coriaceous on lateral margins; notauli not indicated, parapsidal furrows or lateral lines present apically (Fig. 189). Mesopleuron polished, sometimes very weakly coriaceous, mesopleural suture or carina absent; sub- pleural area densely pubescent, antero-ventral cavity obscured by dense woolly pubescence enveloping the lateral margins of propodeum; metapleuron smooth and polished. Lateral bars of scutellum strongly striate; scutellar fovea large; scutellar disc coriaceous to weakly broken striate, almost square apically; scutellar cup (scutellar plate) long and narrow, rarely oval, not reaching apical margin of scutellar disc, apical fovea long and narrow. Propodeum completely covered dorsally and laterally with dense woolly pubescence. Segment 1 of gaster almost completely obscured by ring of dense woolly pubescence at base of tergite 2; tergites 2—5 (or less often 2—4) visible in lateral view, tergite 2 occupying most of visible area; apex of tergite 2 and visible parts of tergites 3—4 sculptured. Wing surfaces pubescent, margins ciliate, longest apically; fore wing with radial cell open or closed. Legs long, with scattered pubescence. DistrIBuTION. Afrotropical Region. Discussion. Sirenes is most closely related to Glauraspidia but the sculpture of the head, mesoscutum and mesopleuron together with the usually closed radial cell of the fore wing and the form of antenna in both male and female distinguish it. Key to the species of Sirenes gen. n. Females 1 Antenna filiform, rhinaria present on the 8—9 apical segments, segment 3 shorter than 4 (4-13 subequal in length) (Fig. 174); scutellar cup sharply declined apically (Fig. 175). spiosp. n.(p. 204) — Antenna weakly clavate, rhinaria present on a lesser number of apical segments, segments 3 and 4 subequal in length; scutellar cup either weakly or sharply declined (Fig. 175, 176). ............. 2 2 Strong rhinaria present on the 3 apical segments of antenna, segment 10 with weak rhinaria (Fig. 177); scutellar cup tear-shaped, sharply declined (Fig. 176); gaster finely punctate apically (Fig. 193); mesopleuron weakly coriaceous; radial cell of fore wing open on margin AT riche lad NED ore Aba cgi dm cag tctt ox EET. KG bv Coie ARE ew dws NRE REES silenus sp. n. (p. 203) — Strong rhinaria present on 4 or more apical segments of antenna; scutellar cup long and narrow or weakly oval apically; gaster either punctate or sculptured; mesopleuron sculptured or pelished zadialcelloffore sing Close, :.. teaecnsxavitav tins seonnaiinn anes ds walle Sve LZ ae a a /) L) RAY) Figs 1-7 1,2, frontal view of head of (1) Angustacorpa sp.; (2) Trichoplasta sp. 3, basal segments of male antenna of Angustacorpa sp. 4, mesonotum of Angustacorpa apsus. 5, gaster of A. apsus. 6, 7, female antenna of (6) A. apsus; (7) A. triton. AFROTROPICAL EUCOILIDAE 209 Figs 8-16 8,9, female antenna of (8) Angustacorpa prodicus; (9) A. persa. 10, mesonotum of A. persa. 11, pronotal plate of A. persa. 12, gaster of A. triton. 13, 14, radial cell of (13) A. triton; (14) A. apsus. 15, 16, Bothrochacis erythropoda (15) female antenna; (16) scutellum. J. QUINLAN 210 ~ S 17 “Seana es om female fore 9 (18) female gaster. 19-21 (21) Emargo sp. b) reed dotoma sp is erythropoda (17) pronotal plate ; (20) Klei Bothrochac 9 18 f (19) B. erythropoda y) Figs 17-21 17 wing oO AFROTROPICAL EUCOILIDAE pA4'a | Figs 22-31 22,23, female gaster of (22) Kleidotoma psiloides; (23) Emargo micipsa. 24, 25, radial cell of (24) Glauraspidia sp.; (25) Sirenes sp. 26-30, Bothrochacis serratepilosa (26) female antenna; (27) pronotal plate; (28) hind tibia; (29) radial cell; (30) scutellar cup. 31, basal segments of female antenna of B. rufiventris. 212 J. QUINLAN Figs 32-40 32,33, Bothrochacis serratepilosa (32) scutellum; (33) female gaster. 34, 35, B. rufiventris (34) lateral view of scutellum; (35) gaster. 36, 37, Diglyphosema latesulcatum (36) basal segment of male antenna; (37) pronotal plate. 38, radial cell of D. eupatorii. 39, 40, D. utica (39) gaster; (40) mesonotum. AFROTROPICAL EUCOILIDAE 213 Figs 41-49 41, 42, radial cell of (41) Diglyphosema utica; (42) D. latesulcatum. 43, antenna of D. utica. 44—46, D. eupatorii (44) pronotal plate; (45) basal segment of gaster; (46) mesonotum. 47—49. D. latesulcatum (47) female antenna; (48) mesonotum; (49) basal segments of gaster. 214 J. QUINLAN Figs 50-57 50, female antenna of Emargo sp. 51, 52, Emargo ascia (51) mesonotum; (52) female gaster. 53-57, female antenna of (53) E. laverna; (54) E. pexus; (55) E. capito; (56) E. cantus; (57) E. ascia. AFROTROPICAL EUCOILIDAE 215 Figs 58-66 58, female antenna of Emargo matius. 59, scutellum of EF. palloris. 60, mesonotum of E. numa. 61-64, female antenna of (61) E. themis; (62) E. vacuna; (63) E. micipsa; (64) E. peleus. 65, radial cell of E. ascia. 66, mesonotum of E. cantus. | 216 J. QUINLAN CLE AGE = aes cnn 75 Figs 67-75 67, radial cell of Emargo cantus. 68, 69, mesonotum of (68) E. capito; (69) E. matius. 70, 71, E. numa (70) female antenna; (71) mesonotum. 72, female antenna of E. palloris. 73, gaster of E. peleus. 74, 75, mesonotum of (74) E. pexus; (75) E. eciton. AFROTROPICAL EUCOILIDAE 217 77 Figs 76-83 76-80, Eucoila crassinerva (76) female antenna; (77) head; (78) basal segments of male antenna; (79) mesonotum; (80) radial cell. 81, fore wing of Trybliographa rapae. 82, 83, female antenna of (82) Eucoila marina; (83) E. erinna. 218 J. QUINLAN Figs 84-91 84, 85, female antenna of (84) Eucoila bantia; (85) E. erinna. 86, 87, E. bantia (86) pronotal plate; (87) mesonotum. 88-90, E. marina (88) mesopleuron; (89) pronotal plate; (90) radial cell. 91, scutellum of E. erinna. AFROTROPICAL EUCOILIDAE 219 EES Se ogee et a = 93 100 99 Figs 92-102 92, radial cell of Eucoila erinna. 93-96, Glauraspidia microptera (93) female antenna; (94) male antenna; (95) pronotal plate; (96) radial cell. 97, radial cell of G. scyphus. 98-102, G. casca (98) female antenna; (99) male antenna; (100) pronotal plate; (101) scutellum; (102) gaster. 220 J. QUINLAN OAS ee, =, te CL e q Ae OCR Yo) ey eens” Figs 103-11 103-105, Glauraspidia scyphus (103) female antenna; (104) basal segments of male antenna; (105) female gaster. 106-109, Leptopilina longipes (106) basal segments of male antenna; (107) mesonotum; (108) gaster; (109) hind coxae. 110, 111, mesonotum of (110) L. clavipes; (111) L. fimbriata. AFROTROPICAL EUCOILIDAE 221 cronies, = n Figs 112-119 112, radial cell of Leptopilina thetus. 113-119, female antenna of (113) L. longipes; (114) L. misensus; (115) L. boulardi; (116) L. fimbriata; (117) L. thetus; (118) L. itys; (119) L. apella. pp) J. QUINLAN Figs 120-126 120, female antenna of Leptopilina fannius. 121, mesopleuron of L. boulardi. 122, radial cell of L. atraticeps. 123-125, scutellum of (123) L. atraticeps; (124) L. heterotoma; (125) L. victoriae. 126, mesopleuron of L. heterotoma. ~ i; \ 4 N ‘ AFROTROPICAL EUCOILIDAE 223 Yh oy: eo oN <4, ware AG ae) ™ arene: —— ae 132 134 Figs 127-135 127, 128, Leptopilina syphax (127) scutellum; (128) radial cell. 129, 130, L. pisonis (129) radial cell; (130) mesopleuron. 131, mesopleuron of L. victoriae. 132, female antenna of L. vesta. 133, male antenna of L. mahensis. 134, mesopleuron of L. syphax. 135, scutellum of L. fimbriata. 224 J. QUINLAN 147 145 ee ae ne \ \ vl ly 139 137 Figs 136-147 136, scutellum of Leptopilina clavipes. 137, 138, L. fimbriata (137) male antenna; (138) fore wing. 139, male antenna of L. longipes. 140, 141, radial cell of (140) L. heterotoma; (141) L. misensus. 142, male antenna of L. syphax. 143, scutellum of L. fannius. 144, 145, male antenna of (144) L. fannius; (145) L. vesta. 146, 147, L. victoriae (146) radial cell; (147) male antenna. AFROTROPICAL EUCOILIDAE 225 159 Figs 148-159 148-150, Leptopilina apella (148) scutellum; (149) propodeum; (150) radial cell. 151, 152, pronotal plate of (151) L. atraticeps; (152) L. boulardi. 153, propodeum of L. boulardi. 154-156, L. fannius (154) metapleuron; (155) radial cell; (156) female antenna. 157, 158, L. faunus (157) meta- pleuron; (158) radial cell. 159, pronotal plate of L. apella. 226 J. QUINLAN Figs 160-170 160, female antenna of Leptopilina heterotoma. 161, 162, L. itys (161) propodeum; (162) scutellum. 163, 164, L. mahensis (163) propodeum; (164) radial cell. 165-167, L. misensus (165) male antenna; (166) pronotal plate; (167) metapleuron. 168, antenna of L. syphax. 169, scutellum of L. thetus. 170, radial cell of L. vesta. AFROTROPICAL EUCOILIDAE a2] Figs 171-180 171-173, Sirenes sinis (171) female antenna; (172) vertex; (173) scutellum. 174, 175, S. spio (174) female antenna; (175) scutellum. 176, 177, S. silenus (176) scutellum; (177) female antenna. 178-180, female antenna of (178) S. steropes; (179) S. orbilus; (180) S. syrinx. 228 J. QUINLAN ial Reeren way rl yet ass ia: £ aN MY 4 Nn OE EET ea : =< SS BARRE NACE NSN 187 eRe rexel SS Figs 181-189 181, 182, Sirenes syrtes (181) female antenna; (182) mesopleuron. 183, 184, male antenna of (183) S. syrinx; (184) S. syrtes. 185, mesopleuron of S. spio. 186-188, S. floccus (186) female antenna; (187) gaster; (188) radial cell. 189, mesonotum of S. orbilus. AFROTROPICAL EUCOILIDAE 229 f Lee “¢: Mee: She ek " Figs 190-199 190, radial cell of Sirenes orbilus. 191-194, S. silenus (191) pronotal plate; (192) mesono- tum; (193) gaster; (194) radial cell. 195, gaster of S. sinis. 196, pronotal plate of S$. spio. 197, 198, radial cell of (197) S. syrinx; (198) S. spio. 199, vertex of head of S. steropes. British Museum (Natural History) Milkweed butterflies: their cladistics and biology P.R. Ackery & R. I. Vane-Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of their biology have stimulated far more attention than can be justified by species numbers alone. In recent years, an expansive literature has grown, considering aspects of their courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and genetics. The popularity of danaines among biologists can certainly be attributed to this combination, within one small group, of so many of the factors that make butterflies such an interesting group to study. The obvious need to place this wealth of biological data within an acceptable systematic framework provided the impetus for this volume. Started in 1976 within the conventions of evolution by natural selection and Hennig’s phylogenetic systematics, the book is now largely about natural history (what the animals have and do, where they live and how they develop) and natural groups — as revealed by a form of analysis approaching that practised by the new school of ‘transformed cladistics’. The authors have prepared a handbook that will appeal to a wide range of biologists, from museum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. ISBN 0 565 00893 5. 1984. Price £50. Titles to be published in Volume 56 The legume-feeding psyllids (Homoptera) of the west Palaearctic Region By I. D. Hodkinson & D. Hollis A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera) By D. Hollis aA The world genera of Rhadalinae (= Aplocneminae) (Coleoptera: Melyridae) By E. R. Peacock A revision of some Afrotropical genera of Eucoilidae (Hymenoptera) By J. Quinlan Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester ean es: = ras i, uy i yh Py oe “oe fare Serene reeset as ae ow veers cs - - Lares rerersaees vente ree The he ia as ae ae pasar ee A feeme lente we Ae Peer r Rasy Stet NER ey to ceed ary rawr) * y v eA * tees at vai ae Lo " co : eons aes ree ot ee eer Los Be pcre ee poe ta an RR NARS ND Rafe eae eee nen ea C sa : S Peatipe pic bok che conv) ; : ere sigetes ont anh 3 aan oS Sapenrenn ue Rerree tate ? Tae ewe? | rs