British Museum | ' (Natural History) — . . oe » . ¢ ¥ a re . r yr? J e t 4 ? "% ‘ f *. ih. . © , i. | Fs ; Z ee " * it av fey , F - 26 OCTOBER 1989 58 NUMBER 2 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. The Entomology Series is produced under the editorship of the Keeper of Entomology: Dr L. A. Mound Publications Manager (Entomology): Mr W. G. Tremewan 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 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. A volume contains about 256 pages, made up of two numbers: published Spring and Autumn. 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BROOMFIELD Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD CONTENTS Introduction 131 Material, methods and terminology 132 Moraceae-feeding psyllids 133 Homotomidae Heslop-Harrison 135 Key to genera 140 Dynopsyllinae Bekker-Migdisova 141 Diceraopsylla Crawford 142 Dynopsylla Crawford 143 Austrodynopsylla gen. n. 144 Triozamia Vondraéek 145 Afrodynopsylla gen. n. 146 Macrohomotominae White & Hodkinson 147 Mycopsylla Froggatt 148 Macrohomotoma Kuwayama 151 Pseudoeriopsylla Newstead 153 Homotominae Heslop-Harrison 156 Homotoma Guérin-Méneville 157 Synoza Enderlein 165 References 166 Index 181 Synopsis. The associations between Ficus and various groups of insects are briefly discussed. That between Ficus and the Psylloidea, particularly the family Homotomidae, is considered in more detail. It is concluded that, at the present level of our understanding of the phylogeny of Ficus and that of agaonine wasps, drosophilid flies, nymphalid butterflies and jumping plantlice, stepwise coevolution cannot be inferred in any of these associations. The systematics of the psyllid family Homotomidae is reviewed. The family is diagnosed and considered as the sister-group of the Malvales-feeding family Carsidaridae. The 10 genera comprising the Homotomidae are diagnosed; a key is provided for their identifica- tion and a cladogram is offered, based on an analysis of 35 characters. Full synonymy and hostplant data, where known, are provided for the 72 nominal species recognised; where practical, keys to species are given. Two new genera and 15 new species are described; two family-group names, one genus-group name and one species-group name are synonymised. nae The Moraceae is a pantropical family of INTRODUCTION dicotyledonous angiosperms containing more than 1400 species in 53 genera. Most of the genera ‘The combination of botanical and ento- contain few species but the diverse genus Ficus mological research has now put Ficus in includes more than half of the known moraceous the front rank of evolutionary and phy- species. The family is most closely related to the logenetic studies’ Urticaceae (Berg, 1977), and Thorne (1983) even E. J. H. Corner (1985a) regarded it as a subgroup of the latter. 132 There appears to be a consensus of opinion that the Moraceae is of Gondwanan origin. Raven & Axelrod (1974: 592) considered the family old enough for direct dispersal between Africa and South America, and Gentry (1982: 569) classed it as a Gondwanan element. However, there seems to be considerable argument concerning the pal- aeogeographical origin of the genus Ficus. Croizat (1968) argued for an early evolution in Gond- wana, while Corner (1985b) forcefully maintained his long-held theory of a Laurasian ancestry for the genus. Whatever the merits of these argu- ments there is agreement that the present-day Ficus, with its closed inflorescence or syconium, arose from an ancestor with an open inflorescence in the mid Cretaceous Period, about 100 million years ago (Galil, 1977; Murray, 1985; Boucek, 1988). The symbiotic association between Ficus (figs) and their fig wasp pollinators (Agaoninae) is well- documented (Wiebes, 1979; 1986 for reviews), and Jermy (1984) suggested that this association may be one of the very few true examples of coevolution between plants and insects. Pollinat- ing fig wasps are species-specific to their fig hosts and there is a reasonable correlation between fig wasp generic or species-group classification and the subsections and series classification (Corner, 1965) of Ficus. In fact Ramirez (1977, 1980) sug- gested modifications to Corner’s arrangement of Ficus species, based on the specificity and mor- phology of their pollinating wasps. Wiebes (1982) further compared classifications of Agaoninae and Ficus (from Ramirez, 1980) and concluded that there is no correlation, at a higher level than that noted above, between the two classifications. Corner (1985a) reviewed the modifications sug- gested by Ramirez and Wiebes but rejected them, for the large part, in favour of his original classification. Miller (1987) briefly reviewed the case and considered that more rigorous clado- grams were required for both groups of organisms before stepwise coevolution could be inferred. Hill (1967) listed a further 46 genera of non- agaonine (i.e. non-pollinators sensu Boucek, 1988) fig wasps, known to be associated with fig syconia, in other subfamilies of the Agaonidae and the families Eurytomidae, Ormyridae, Eulophidae and Pteromalidae. The roles played by these wasps appear to be varied but have been little studied. Some are phytophagous, others are hyperparasites, and there is some degree of fig specificity. Lachaise et al. (1982) reported on the associ- ation between fig syconia and two groups of drosophilid flies, Lissocephala and _ the Drosophila fima_ species-group, in Africa. D. HOLLIS & P. S. BROOMFIELD Lissocephala larvae develop during the floral period of the syconium and each species of fig studied harboured a particular combination of Lissocephala species; different species of figs pos- sibly having some Lissocephala species in com- mon. Drosophila fima_ species-group larvae develop in the postsexual phase of the syconium and there appeared to be no fig species preference in this group. Lachaise et al. (1982) postulated that the speciation of Lissocephala was a by-pro- duct of the coevolution of figs and their pollinating wasps. Apart from those insects associated with fig syconia there are other groups of insects known to feed on Ficus species. Two such groups worthy of mention are the larvae of nymphalid butterflies, and the jumping plantlice or Psylloidea (Homop- tera). Ackery (1988 and pers. comm.) has reviewed the hostplants of nymphalid larvae and recognised some Moraceae-feeding ‘themes’: Marpesiine larvae have specialised on Moraceae genera, including Ficus; the Limenitine genus Pseudoneptis feeds on Antiaris and Ficus; and the genus Euploea (Danainae) shows a trend towards Ficus-feeding but plants in the families Apo- cynaceae, Rubiaceae, Ulmaceae, Flacourtiaceae, and Convolvulaceae are also known hosts. The Psylloidea (jumping plantlice) comprise a group of small, phloem-feeding sternorrhynchous bugs. Individual psyllid species show a high degree of hostplant specificity, particularly during the larval stages, and related species tend to develop on related species or groups of dicotyledonous angiosperms. Several groups of psyllids are known to utilise moraceous hostplants and one family in particular, the Homotomidae, is almost exclusively Ficus-feeding. The objectives of this paper are to review the known data on Moraceae-feeding in the Psylloidea; to postulate a phylogeny for the gen- era of the family Homotomidae, for future com- parison with a cladogram of the subgroups of Ficus, should this ever be produced; and to review the taxonomy of the species in the family. MATERIAL, METHODS AND TERMINOLOGY Most of the material studied is deposited in the British Museum (Natural History) (BMNH), with supplementary specimens from the Bernice P. Bishop Museum, Honolulu (BPBM); Muséum d’Histoire Naturelle, Geneva (MHNG); Muséum National d’Histoire Naturelle, Paris (MNHN); FICUS-FEEDING PSYLLIDS Musée Royal de |l’Afrique Centrale, Tervuren (MRAC); Stanford University Natural History Museum, California (SUNHM); and the National Museum of Natural History, Washington (USNM). Type material is deposited in BMNH; BPBM; MHNG; MNHN; MRAC; SUNHM; USNM; Australian National Insect Collection, Canberra (ANIC); Beijing Agricultural Univer- sity Insect Collection, China (BAUIC); Ento- mological Institute, Hokkaido University, Sapporo (EIHU); Forest Research Institute, Dehra Dun, India (FRI); Institut ftir Pflanzenschutzforschung, Eberswalde (IPE); Instytut Zoologiczny, Polish Academy of Sci- ences, Warsaw (IZPAN); National Chung Hsing University, Taiwan (NCHU); National Collection of Insects, Plant Protection Research Institute, Pretoria (NCI); and the Zoological Survey of India, Calcutta (ZSI). All measurements are quoted in millimetres; most were taken from slide-mounted material but overall size was taken from the anterior margin of the head to the tip of the forewing of dry-mounted material in lateral view. Other reference points from which measurements of various structures were taken are given in Hollis (1976, 1984). Apart from the head illustrations, all figures were drawn from slide-mounted material. The inner surface of the right paramere is shown. Structural terminology follows Vondraéek (1957) and Hollis (1984, 1987). ACKNOWLEDGEMENTS. We are grateful to the following colleagues for providing loans and gifts of material: Keith Arakaki and the late Wayne Gagné (BPBM); Daniel Burckhardt (MHNG); J. Etienne, formerly of Institut Sénégalaise Recherches Agricoles, Ziguinchor; and Douglass Miller and Miss Louise Russell, United States Department of Agriculture, Systematic Entomo- logy Laboratory, Beltsville, Maryland. MORACEAE-FEEDING PSYLLIDS Table 1 summarises the available data on psyllid genera associated with moraceous hosts. The genus Paurocephala has 25 species in the Old World tropics, with free-living larvae. There is one species on Artocarpus, one on Morus and three on Ficus. Most of the other species live on hosts in the Malvales (Tiliaceae, Malvaceae and Sterculiaceae) but there are a few species on Clusiaceae (Theales) and Connariaceae (Rutales). The South American species are on Melastomaceae but these are probably not con- 1133 generic with the Old World species. The genus is most closely related to Haplaphalara and Diclidophlebia which also have hosts in the Malvales. Phytolyma is an African genus of four species (Hollis, 1973), three on Milicia and one on Morus. The larvae are gall-forming on leaves and petioles. Its relationships are not clear but White & Hodkinson (1985) placed the genus as the sis- ter-group of the rest of the Aphalarinae. Anomoneura is a monobasic Asian Palaearctic genus living on Morus; it may be related to the legume-feeding genus Epipsylla (White & Hodkinson, 1985). The triozid genus Pauropsylla has 22 species in the Old World tropics and subtropics and all con- firmed hosts are Ficus spp. The larvae usually form pit galls on the host leaves. Ceropsylla fulvida is an Indian species recorded by Mathur (1975) on Ficus microcarpa [as macrocarpa| and F. rumphii. The species is not congeneric with American Ceropsylla which are found on Ocotea (Lauraceae) and Sideroxylon (Ebenaceae). Tri- oza is a large and probably paraphyletic genus of some 600 species (Hollis, 1984). Three species are known to occur on Ficus species. T. buxtoni is a Middle Eastern species on F. carica and the larvae cause severe leaf distortion; T. ficicola is a South African species on Ficus sp.; T. brevigena occurs on Ficus sp. in N. India and its larvae cause leaf- margin rolls. The homotomid genus Triozamia is an interest- ing example of the degree to which psyllids are host specific (Hollis, 1984). The three known spe- cies appear to maintain separate identities on the three recognised African varieties of Antiaris toxi- caria welwitschii (cf. Berg, 1978), 1.e. T. lamborni on var. africana, T. vondraceki on var. welwitschii and T. usambarensis on var. usambarensis. Table 2 summarises the available data on psyllids having Ficus spp. as hostplants. Using Corner’s (1965) infrageneric arrangement of Ficus some patterns can be discerned. Pauropsylla occurs mainly on subgenus Sycomorus, with a few species on section Sycidium of subgenus Ficus. The original records of P. ficicola and P. globuli on Ficus hookeriana (Kieffer, 1905; as F. hookeri) are regarded as dubious; P. ficicola adults and larvae have been collected recently from F. auriculata in N. India (BMNH data). Paurocephala species are restricted to subgenus Ficus. Homotomid species occur mainly on the stran- gler and banyan subgenus Urostigma, with a few species on subgenus Ficus and one genus, Dynop- sylla, whose larvae are gall-forming, on F. nervosa of subgenus Pharmacosycea. Of the groups that D. HOLLIS & P. S. BROOMFIELD 134 pm | pudjojAyd pjvydas0inog SNOW (vsoydosojyD ueolyjy =) pudoiyd DIDI pjjksdouasopnasg vz0uy DWOJOWOL DWOJOWOYOLID pjjdsdoukq pjjksdosaD vzouks pydsdookyy | vpyjdsdovsaoigg | vjjdsdounvg Pilg pjpydasoinvg Sn deplwo}OWOY caw aepryAsd QeDLIOW supyupy BaplojAsg JO elouaD JO BIouaD SEUIWIOJOWOF{ | IEUIWIOJOWTOYOIORWY aeulyAsdoukq “RaployAsg Jo eiauas Aq syurjdjsoy se payiojdxa avaoe1oyy Jo e1oUDH | aquy FICUS-FEEDING PSYLLIDS occur on subgenus Urostigma, the African genus Pseudoeriopsylla is restricted to section Galoglychia while its sister-group, the Oriental genus Macrohomotoma, is on section Conosycea. Mycopsylla is less restricted with the Australasian species on section Malvanthera and the Indian species recorded from sections Urostigma and Conosycea. Homotoma species have an even broader spectrum of Ficus hosts in sections Urostigma, Conosycea and Galoglychia of sub- genus Urostigma, and sections Ficus and Rhi- zocladus of subgenus Ficus. The record of Diceraopsylla on Ficus elastica (subgenus Urostigma, section Stilpnophyllum) is doubtful (see p. 142). Corner’s infrageneric conspectus of Ficus is, to some extent, based on plesiomorphies and does not indicate phylogenetic relationships (Wiebes, 1979). Ramirez (1980: fig. 1) produced a ‘clado- gram’ for Ficus, based on ‘fifty-four morphologi- cal characters and a few physiological characters’. Unfortunately, few of these characters were dis- cussed and the resulting branching diagram can- not be assessed critically. However, Table 3 shows a slightly modified version of Ramirez’ cladogram of Ficus sections, and the homotomid genera that are associated with these sections. Clearly step- wise coevolution cannot be inferred from these data. The cladogram of homotomid genera (Fig. 1) is presented so that, when a more rigorous phy- logeny of Ficus is produced and more reliable hostplant data for the psyllids are gathered, the case may be reconsidered. HOMOTOMIDAE Heslop-Harrison Homotomini Heslop-Harrison, 1958: 578; Log- inova, 1964b: 54. Type genus: Homotoma Guérin-Meéneville. Homotominae Heslop-Harrison; Klimaszewski, 1964: 91; Bekker-Migdisova, 1973: 101. Homotomidae Heslop-Harrison; White & Hodkinson, 1985: 272; Hollis, 1987: 90. Brown & Hodkinson, 1988: 179. Carsidarinae Crawford; Yang, 1984: 168, in part. Diacnosis. A pair of strong tubercles present on metapostnotum; ventral sense organs of hind femur in basal position, proximal organ offset from distal pair; C’ proctiger bipartite (not in Syn- oza); © subgenital plate without laterodorsal appendages; rs—m crossvein absent from forewing. 135 Hostp.ants. Moraceae (Antiaris and Ficus). Discussion. As diagnosed here the family con- tains the following genera: Diceraopsylla, Dynop- sylla, Austrodynopsylla, Triozamia, Afrodynop- sylla, Mycopsylla, Macrohomotoma, Pseudo- eriopsylla, Synoza and Homotoma. Diceraopsylla is reassigned from the Aphalaridae, and Tri- ozamia is transferred from the Triozidae. Austrodynopsylla and Afrodynopsylla are described as new below. The group was erected by Heslop-Harrison as a tribe of his polyphyletic subfamily Ciriacreminae, to contain the genera Homotoma, Psausia, Labobrachia, Metapsausia, Mycopsylla, Sphingo- cladia, Synoza, Crawfordella and Dynopsylla. He placed Macrohomotoma in the Phacopteronini and Diceraopsylla in the Carsidarini. Of these original genera Psausia was synonymised with Homotoma by Kuwayama (1931); Labobrachia and Metapsausia were inferred as synonyms of Homotoma by Miyatake (1975); Sphingocladia and Crawfordella have been, most recently, syn- onymised with Dynopsylla by Crawford (1924) and Yang (1984) respectively. Macrohomotoma and Diceraopsylla were transferred to the Homo- tominae by Bekker-Migdisova (1973) and Diceraopsylla was erroneously transferred to the Aphalaridae by Hollis (1984). Loginova (1964b) and Klimaszewski (1964) both noted the polyphyletic nature of Heslop- Harrison’s Ciriacreminae and regarded homo- tomids as a subgroup of the Carsidaridae, as did Bekker-Migdisova (1973) but her concept of the latter included several non-carsidarid groups (White & Hodkinson, 1985; Hollis, 1987). White & Hodkinson (1985) regarded homotomids as a distinct family containing the genera Homotoma, Synoza, Mycopsylla, Macrohomotoma, Pseudoeriopsylla, and probably Dynopsylla and Sphingocladia. They diagnosed the group on the following ‘derived’ characters (White & Hodkin- son, 1985: 239, clade 6). 1. Adult antenna with rhinaria absent from seg- ments 3, 5 and 7. 2. Larva with dorsal surface of thorax with dis- tinct sclerites. 3. Larva broader than long. 4. Antenna of larva short, narrowed evenly to apex. However, for a number of reasons this diag- nosis is not valid. Character 1 does not occur in many members of the Dynopsyllinae, but is a widespread condition in the Psylloidea; character 2 is a primitive condition, occurring in other famil- ies; character 3 does not occur in Synoza or in D. HOLLIS & P. S. BROOMFIELD 10y|vAdLv9 vyAsdouaopnasg (vpoddyoviq se) vIvAO avduvoojnvD piopopdupnjyo vuojowoyY ‘B8uivi vjj reaching wing margin anterior to wing apex; hind- wing with distinct M+ Cu stem; hind tibia without basal spine, apical spurs arranged 1 + 4, hind basitarsus with 1 apical spur; apical segment of aedeagus undivided. Comments. This subtribe contains two genera, Dynopsylla and Austrodynopsylla, species of which have an Oriental and Melanesian distribution. DYNOPSYLLA Crawford Dynopsylla Crawford, 1913: 295; 1924: 618; Enderlein, 1921: 119; 1926: 399; Kuwayama, 1931: 122; Heslop-Harrison, 1958: 578; 1960: 240; Bekker-Migdisova, 1973: 102; Mathur, 1975: 132; Hodkinson, 1983: 346; 1986: 308; White & Hodkinson, 1985: 274. Type species: Dynopsylla cornuta Crawford, by original des- ignation and monotypy. Sphingocladia Enderlein, 1914: 231; 1918: 482; 1926: 399; Heslop-Harrison, 1958: 578; Bek- ker-Migdisova, 1973: 102; White & Hodkinson, 1985: 274. Type species: Sphingocladia pin- nativena Enderlein, by original designation and monotypy. [Synonymised by Enderlein, 1921: 119; resynonymised by Crawford, 1924: 619; Mathur, 1975: 132.] Crawfordella Enderlein, 1926: 398; Heslop-Harri- son, 1958: 578; Bekker-Migdisova, 1973: 102; Hodkinson & White, 1981: 496. Type species: 143 Dynopsylla grandis Crawford, by original des- ignation and monotypy. [Synonymised by Mathur, 1975: 132; resynonymised by Yang, 1984: 175.] Dynopsilla Crawford; Enderlein, [Misspelling. ]. eis iis DescriPTION. Large psyllids, up to 7.5 mm long. Integument of head and thorax with a moderately dense covering of long setae. Head (Figs 4, 5), from above, narrower than mesoscutum; disc of vertex concave and deeply divided by median suture, foveae moderately defined, frontal margin rounded and deeply incised by median suture, lateral margins rounded, occipital margin obtu- seangular, lateral ocelli on raised tubercles, median ocellus visible from above, anterolateral tubercles strongly developed, antennal sockets enlarged giving head a cleft appearance; antennal scape enlarged and elongate, at least twice as long as pedicel, flagellum elongate filiform, with a sin- gle subapical rhinarium on flagellomeres 1, 2, 4, 6 and 7; ultimate rostral segment 2.5-5.0 times longer than wide. Thorax, in profile, strongly arched (Fig. 41), pronotum narrowly visible from above. Forewing (Fig. 49) ovate, with acute apex, about 2.4 times longer than wide; veins bearing long setae, C+Sc thickened, costal break absent, apex of M/,,> reaching margin anterior to apex of wing, M+ Cu long, Cu stem short or absent, 7, cell value about 1.0, cu, cell value about 1.0, apex of claval suture distant from apex of Cu,,, radular areas clearly defined; hindwing (Fig. 50) broad, M+Cu stem present, Cu unbranched, anal lobe not enlarged. Basal spine of hind tibia absent, apical spurs arranged | + 4; hind basitarsus with | apical spur. CO proctiger bipartite, lateral lobes developed but without inner lobes; aedeagus 2-segmented (Fig. 77), basal segment not expanded. Q genital segment conical, anal pore ring convoluted. Larva. 5th instar larva of D. pinnativena described by Yang (1984). HostTpLants. Ficus (Pharmacosycea) sect. Oreo- sycea; larvae are gall-forming. Comments. The three species known in this genus all occur on Ficus nervosa. They may be separated using the following key: 1 Custem absent; M stem contiguous with basal part of Cu,, and medial part of Rs (see Crawford, 1924: 620, fig. 3); antennal pedicel with large ventral lobe grandis — Short Cu stem present; M stem contiguous with medial part of Cu,, and medial part of Rs (Fig. 49); antennal pedicel without ventral lobe .......... 2 144 2 Antennal scape twice as long as pedicel, flagellum less than 4 times longer than hind tibia; ultimate rostral segment 2.5 times longer than wide pinnativena — Antennal scape 3 times longer than pedicel, flagellum 6 times longer than hind tibia; ultimate rostral segment 5 times longer than wide . cornuta Dynopsylla cornuta Crawford (Figs 4, 5, 41) Dynopsylla cornuta Crawford, 1913: 295; 1924: 619; 19256: 62; Enderlein, 1926: 399; Tak- ahashi, 1936: 293; Heslop-Harrison, 1960: 241; Miyatake, 1971: 58; Mathur, 1973: 71; 1975: 133; Hodkinson, 1983: 346. Syntypes, 20°, 19, PHILIPPINES (USNM) [examined]. HostTeLant. Ficus nervosa; Crawford (1924) stated that the species is gall-forming but did not describe the gall. RECORDED DISTRIBUTION. Philippines (Luzon and Palawan). MATERIAL EXAMINED East Malayasia: 1 ©’, Sarawak. Indonesia: 1 Q, Sulawesi Utara (BMNH); 1 9, Sumatra (MNHN). Philippines: 3 o', 2 9, Los Banos (USNM). Dynopsylla grandis Crawford ‘Undetermined species [psyllid]’ Ramakrishna Ayyar, 1920: 1030. Dynopsylla grandis Crawford, 1924: 619; 1925b: 62; Takahashi, 1936: 293; Costa Lima, 1942: 103; Mathur, 1973: 70; 1975: 134; Hodkinson, 1986: 308. Syntypes, 3 2, INDIA (Kerala), Ficus nervosa, (USNM) [examined]. Dinopsylla grandis Crawford; Mani, 1973: 283, [286, probable misidentification]. [Mis- spelling. | Crawfordella grandis (Crawford) Enderlein, 1926: 398; Hodkinson & White, 1981: 496. HostTpLant. Ficus nervosa; Ramakrishna Ayyar (1920) records this species as forming midrib galls on the upper leaf surface. RECORDED DISTRIBUTION. India (Kerala). The four specimens Crawford (19255) records from Brazil are regarded as bearing incorrect data. MATERIAL EXAMINED India: 3 2 (syntypes, USNM). D. HOLLIS & P. S. BROOMFIELD Dynopsylla pinnativena (Enderlein) (Figs 49, 50, 77) Sphingocladia pinnativena Enderlein, 1914: 231; 1918: 482; 1926: 399; Mathur, 1973: 71. Holo- type O', ‘FORMOSA’ (IPE) [not examined]. Dynopsylla (Sphingocladia) pinnativena (Enderlein) Kuwayama, 1922: 368. Dynopsylla pinnativena (Enderlein) Crawford, 1924: 619; 1925b: 62; Kuwayama, 1931: 122; Takahashi, 1936: 293; Miyataki, 1965b: 226; Mathur, 1975: 133 Hodkinson, 1983: 346; 1986; 308; Yang, 1984: 175. HostpLant. Ficus nervosa; Takahashi (1936) records this species forming leaf-margin rolls. RECORDED DISTRIBUTION. Taiwan. MATERIAL EXAMINED Vietnam: | Oh (MNHN). AUSTRODYNOPSYLLA gen. n. Type species: Austrodynopsylla encala sp. n., here designated. DEscriPTION. Large psyllids, up to 5.5 mm long. Integument of head and thorax with a moderately dense covering of long setae. Head (Figs 6, 7), from above, narrower than mesoscutum; disc of vertex weakly concave with defined foveae, shal- lowly divided by median suture, anterior and lat- eral margins rounded, occipital margin obtuseangular, lateral ocelli on raised tubercles, median ocellus visible from above, anterolateral tubercles absent. Antennal sockets not enlarged and head without cleft appearance; scape not enlarged or elongated; flagellum filiform, 2.4 times longer than head width, with a group of apical rhinaria on flagellomeres 1, 2 and 4, two rhinaria on flagellomere 6 and a double rhinarium on 7; ultimate rostral segment about 3 times longer than wide. Thorax, in profile, moderately arched, pro- notum narrowly visible from above. Forewing (Fig. 51) obovate with rounded apex, about 2.3 times longer than wide, veins bearing long setae. apex of M_,, reaching margin anterior to wing apex, M+Cu stem present, Cu stem virtually absent, m, cell value about 1.0, cu, cell value about 1.25, apex of claval suture distant from apex of Cu;,, radular areas clearly defined. Hindwing (Fig. 52) broad, M+Cu stem present, Cu unbranched. Basal spine of hind tibia absent, api- cal spurs of hind tibia arranged 1 + 4 (Fig. 43), hind basitarsus with a single apical spur. FICUS-FEEDING PSYLLIDS CO proctiger bipartite, lateral lobes developed and expanded dorsally, without inner lobes api- cally; aedeagus 2-segmented, basal segment not expanded. @ terminalia conical, anal pore convoluted. Larva unknown. HostTPLant. Ficus sp. ComMENTS. This genus is regarded as the sister- group of Dynopsylla; it differs in head and anten- nal structure (Figs 4-7, 27) and forewing venation (Figs 49, 51). Austrodynopsylla encala sp. n. (Figs 6, 7, 27, 43, 51, 52, 78, 79) Description. Additional to generic characters above. Antennal flagellum 2.35—2.40 times longer than head width; 1st flagellomere with a group of 4-5 rhinaria apically and a subapical rhinarium, 2nd with 34 apical rhinaria (Fig. 27), 3rd without rhinaria, 4th with 3 apical rhinaria, 5th without rhinaria, 6th with 2 apical rhinaria, 7th with a double apical rhinarium, 8th with one short pointed seta and one very short truncated seta apically. Forewing (Fig. 51) hyaline, with small brown patches along R+M+Cu, at fork of M+ Cu and at apex of Cu,,; 2.13—2.34 times longer than wide, M+Cu stem as long as R stem, Rs short and curved towards M stem, branches of M of equal length; hindwing (Fig. 52) about 0.5 times as long as forewing, with irregularly grouped costal setae; apex of hind tibia as in Fig. 43. CO paramere (Fig. 78) lamellar, rounded api- cally and with a posteroapical tubercle; apical seg- ment of aedeagus (Fig. 79) simple, end-tube of ductus ejaculatorius elongate. Q terminalia conical, apices of proctiger and subgenital plate acute, valvulae without serrations. Measurements (1 0’, 1 2). Maximum width of head, O 0.95, 2 0.80; length of antennal flagellum, ©” 2.28, 2 1.88; length of ultimate ros- tral segment, O 0.19; length of forewing, CO’ 3.85, © 4.55; length of hind tibia, O’ 0.98, 2, 0.80. ( measurements taken from dry specimen.) Hostpiant. Adult (1 2) taken from Ficus sp. Holotype OC’, New Caledonia: Mt Koghi, 400- 600 m, 11.1973 (Krauss) (BPBM); slide- mounted. Paratype. New Caledonia: 1 9, Ile des Pins, 7 km N. Kuto, 3-100 m, 16.viii.1979, on Ficus sp. (Gagné) (BMNH); dry mounted. 145 TRIOZAMIINA Bekker-Migdisova Triozamiini Bekker-Migdisova, 1973: 115. Type genus: Triozamia Vondracek. Triozamini Bekker-Migdisova; Hollis, 1984: 9. [Misspelling. | Triozamiinae Bekker-Migdisova; Hodkinson, 1985: 273. DiaGnosis. Several rhinaria present at base or all over surface of Ist flagellomere, 2nd flagellomere without rhinaria; M+Cu stem absent or very short, apex of M,,, reaching wing margin pos- terior to wing apex, Cu stem present; hindwing with M+Cu stem absent or indistinct; basal spine of hind tibia present, apical spurs arranged 0 + 4-7; apical spurs of hind basitarsus absent; aedeagus 3-segmented. White & ComMENTS. This is the sister-group of Dynop- syllina; it contains two Afrotropical genera, Tri- ozamia and Afrodynopsylla. TRIOZAMIA Vondracek Triozamia Vondraéek, 1963: 266; Bekker-Mig- disova, 1973: 114; Hollis, 1984: 22; White & Hodkinson, 1985: 240. Type species: Rhinop- sylla lamborni Newstead, by original designa- tion and monotypy. DEscriPTION. Robust psyllids, up to 5.5 mm long, somewhat dorsoventrally flattened. Integument of head and thorax densely covered with short setae. Head (Figs 8, 9), from above, slightly nar- rower than mesoscutum; disc of vertex concave, shallowly divided by median suture, foveae dis- tinct, frontal margin rounded and not deeply div- ided by median suture, lateral and occipital margins angular, lateral ocelli not on raised tuber- cles, median ocellus not visible in dorsal view, anterolateral tubercles absent, genae slightly enlarged ventrally and each with a small tubercle below the antennal socket; antennal scape not enlarged or elongate, flagellum filiform, about twice as long as head width, 1st flagellomere bear- ing several rhinaria, 2nd without rhinaria (Fig. 28), a single subapical rhinarium present on flagellomeres 4, 6 and 7; ultimate rostral segment elongate, 4.2—-6.0 times longer than wide. Thorax, in profile, weakly arched; pronotum wide, clearly visible from above; forewing (Fig. 53) elongate ovate with subangular apex, about 2.75 times longer than wide, veins bearing short setae, C+ Sc thickened, costal break absent, apex of M,,. reaching wing margin at apex of wing, M+Cu absent, Cu stem present, m, cell value 146 about 4.5, cu; cell value about 0.5, apex of claval suture adjacent to apex of Cu,,, radular areas clearly defined; hindwing (Fig. 54) narrow, M-+Cu stem not clearly defined, Cu unbranched. Hind tibia with a basal spine, apical spurs arranged 0 + 6-7 (Fig. 44), hind basitarsus with- out apical spurs. CO proctiger bipartite, with well-developed lat- eral lobes that bear inner apical lobes; aedeagus 3-segmented, apical segment elongate (Fig. 81). Q terminalia elongate, conical; anal pore ring convoluted. Larva. 5th instar larva of T. lamborni described by Hollis (1984). HostTpLant. Antiaris toxicaria welwitchii. Comments. Hollis (1984) retained Triozamia in the Triozidae on the basis of the trifurcation of R+M-+Cu and the absence of a costal break and pterostigma in the forewing, but was doubtful of its position within that family. White & Hodkin- son (1985) placed the genus as the sister-group of the rest of the Triozidae minus Neolithus. Given the present diagnosis of the Homotomidae, Tri- ozamia is more naturally placed here, and the presence of wax-producing cells on the posterior tergites of the adult abdomen indicate its relation- ship with other members of the Dynopsyllinae. The presence of several rhinaria on the Ist flagellomere and their absence from the 2nd flagellomere, the reduced M+ Cu stem of the fore- wing, the proximal branching of M+Cu in the hindwing, the absence of apical spurs on the hind basitarsus, and the 3-segmented aedeagus suggest a sister-group relationship with Afrodynopsylla. The two genera may be separated using couplet 4 of the key (p. 141). Three species are included, fully treated by Hollis (1984). Triozamia lamborni (Newstead) (Figs 8, 9, 28, 44, 53, 54, 80, 81) Rhinopsylla lamborni Newstead, 1914: 520; Eastop, 1961: 168. Holotype 0’, NIGERIA [not traced]. Triozamia lamborni (Newstead) Vondracek, 1963: 266, in part; Akanbi, 1981: 113; Hollis, 1984: 23. Triozamia lambourni (Newstead); Roberts, 1969: 78. [Misspelling. | HOstpPLaNtT. Antiaris toxicaria welwitchii var. africana. RECORDED DISTRIBUTION. Senegal, Ivory Coast, Ghana, Nigeria, Zaire and Tanzania. D. HOLLIS & P. S. BROOMFIELD MATERIAL EXAMINED Adults and larvae from Senegal, Guinea, Ivory Coast, Ghana, Nigeria, Zaire and Tanzania (BMNH). Triozamia usambarensis Hollis Triozamia usambarensis Hollis, 1984: 24. Holo- type &', TANZANIA (BMNH) [examined]. HOSTPLANT. Antiaris toxicaria welwitchii var. usambarensis. RECORDED DISTRIBUTION. Tanzania. MATERIAL EXAMINED Tanzania (type series, BMNH). Triozamia vondraceki Hollis [Triozamia lamborni (Newstead); Vondraéek, 1963: 268, in part. Misidentification. | Triozamia vondraceki Hollis, 1984: 24. Holotype oO, UGANDA (BMNH) [examined]. HostTPLant. Antiaris toxicaria welwitchii [? var. welwitchii]. RECORDED DISTRIBUTION. Uganda, Central African Republic. MATERIAL EXAMINED Uganda, Central African Republic (type series, BMNH). AFRODYNOPSYLLA gen. n. Type species: Afrodynopsylla gigantea sp. n., here designated. DescripTIon. Large psyllids, up to 8.0 mm. Integ- ument of head and thorax densely clothed with moderately long setae and with groups of very long setae (Figs 10, 11, 42). Head (Figs 10, 11), from above, narrower than mesoscutum; disc of vertex concave, with clearly defined foveae, frontal margin rounded, lateral margins angular, occipital margin sharply defined, lateral ocelli on raised tubercles, median ocellus visible from above, anterolateral tubercles well-developed; antennal sockets enlarged, giving head cleft appearance, scape enlarged and elongate, pedicel thickened; flagellum filiform, about 2.5 times longer than head width, Ist flagellomere with basal swelling that bears a group of rhinaria (Fig. 29), rhinaria absent from 2nd flagellomere, a sin- gle rhinarium present subapically on flagellomeres 4, 6 and 7; ultimate rostral segment elongate. FICUS-FEEDING PSYLLIDS Thorax, in profile (Fig. 42), strongly arched, pronotum broad and clearly visible from above; forewing (Fig. 55) ovate, with subacute apex, about 2.2 times longer than wide, veins bearing sparse short setae, C+Sc strongly thickened, cos- tal break absent, apex of M,,, reaching wing mar- gin posterior to wing apex, M+Cu stem very short, m, cell value about 2.4, cu, cell value about 1.2, apex of claval suture adjacent to apex of Cuj,, radular spines in small but clearly defined groups; hindwing (Fig. 56) with M+Cu stem absent, Cu unbranched, anal lobe expanded; hind tibia with a basal spine, apical spurs arranged 0 + 4. CO proctiger bipartite, lateral lobes well- developed but without inner apical lobes; aedeagus 3-segmented. Q terminalia conical, anus with a simple, dou- ble ring of wax-producing cells. Larva and hostplant unknown. Afrodynopsylla gigantea sp. n. (Figs 10, 11, 29, 42, 55, 56, 82, 83) DescrireTIon. Antennal flagellum 2.0-2.5 times longer than head width; Ist flagellomere with a basal swelling bearing a group of 15-20 rhinaria mainly on ventral surface (Fig. 29), 8th flagellomere with a long pointed seta and a short truncated seta apically; ultimate rostral segment 4.5 times longer than wide. Dorsum of thorax with tufts of long setae (Fig. 42); forewing (Fig. 55) 2.15-2.35 times longer than wide, membrane hyaline, C+Sc consider- ably thickened, Rs straight; hindwing (Fig. 56) about half as long as forewing; hind tibia with a well-developed basal spine; hind basitarsus much longer than apical tarsal segment. CO paramere as in Fig. 82; aedeagus 3-seg- mented (Fig. 83), apical segment short and strongly swollen apically. @ terminalia short, conical in profile; apices of proctiger and subgenital plate acute; apices of valvulae not serrate. Measurements (2 oO’, 1 2). Maximum width of head, CO 1.02-1.14, 9 1.14; length of antennal flagellum, CO 2.58-2.68, 2 2.21; length of ultimate rostral segment, O” 0.42-0.44, 9 0.42; length of forewing, © 5.16-5.43, 2 5.69; length of hind tibia, Cand 9, 1.14. Holotype oO, Angola: Salazar, I.I.A.A., 9-15.ii1.1972, at light (Hollis) (BMNH); dry- mounted. Paratypes. Angola: 1 2, same data as holotype. Central African Republic: 3 0, 2 2, Lobaye Mbaik, rte Mbale, 12.ii.1969, light trap in forest 147 zone (Boulard). Nigeria: 1 Q, Ile-Ife, 11.i1.1971 (Medler). (BMNH; MNHN); dry- and slide- mounted. MACROHOMOTOMINAE White & Hodkinson Macrohomotominae White & Hodkinson, 1985: 272. Type genus: Macrohomotoma Kuwayama. Dynopsyllini Bekker-Migdisova, 1973: 102, in part. Phacopteronini Heslop-Harrison, 1958: 578, in part. Homotomini Heslop-Harrison, 1958: 578, in part. Diagnosed by the presence of a pterostigma and the absence of a costal break in the forewing. White & Hodkinson (1985) defined the Mac- rohomotominae (as clade 34, fig. 188) on the fol- lowing characters. 1. Male proctiger expanded posteriorly to form caudal lobes. 2. Circum-anal ring [of wax-producing cells] of larva constricted on either side of anus, or broken into three groups. 3. Body margin of larva without sectasetae or derivable structures. 4. Dorsal surface of larva without sectasetae or derivable structures. The first character is diagnostic for the Homo- tomidae and is of no value below this level. The second character is not useful as it occurs also in the Homotominae. The third character is not valid as marginal sectasetae do occur in larvae of Mycopsylla, Macrohomotoma and Pseudoeriop- sylla. The fourth character is not diagnostic as sectasetae are present dorsally on the caudal plate of Pseudoeriopsylla larvae, and sectasetae are absent from the larvae of the species of Dynop- syllinae examined and described. The only adult character found to be useful in diagnosing the Macrohomotominae is the pres- ence of a pterostigma in the forewing, coupled with the absence of a costal break. No diagnostic characters were found in the larvae. Hostplants of the group are in Ficus (Urostigma) sects Urostigma, Conosycea, Mal- vanthera and Galoglychia. Two tribes are recognised. EDENINI Bhanotar, Ghosh & Ghosh. Edenini Bhanotar, Ghosh & Ghosh, 1972: 119. Type genus: Edenus Bhanotar, Ghosh & Ghosh. 148 Dynopsyllini Bekker-Migdisova, 1973: 102, in part. Mycopsyllini White & Hodkinson, 1985: 242, 272. Type genus: Mycopsylla Froggatt. Syn. n. DiaGnosis. Pterostigma of forewing elongate, wedge-shaped; M_,. straight, reaching wing mar- gin anterior to wing apex; basal segment of aedeagus swollen in apical half. A single genus, Mycopsylla, is included, species of which have an Oriental and Australasian distribution. MYCOPSYLLA Froggatt Mycorsylla Froggatt, 1901: 258; Aulmann, 1913: 30; Tuthill & Taylor, 1955: 248; Mathur, 1975: 150; Morgan, 1984: 118; Kandasamy, 1987: 68. Type species: Psylla fici Tryon, by original designation. Edenus Bhanotar, Ghosh & Ghosh, 1972: 119. Type species: Edenus gardenensis Bhanotar, Ghosh & Ghosh, by monotypy. [Synonymised by Hodkinson, 1986: 318. ] Description. Medium- to large-sized psyllids, up to 8.0 mm long. Integument of head and thorax sparsely covered with short setae. Head (Figs 12-17), in dorsal view, almost as wide as meso- scutum; disc of vertex from flat to concave and deeply incised by median suture, foveae weak, anterior and lateral margins rounded, occipital margin angular, lateral ocelli not or on slightly raised tubercles, median ocellus just visible from above, anterolateral tubercles absent or present; antennal sockets weakly enlarged, giving head a weakly cleft appearance, antennal scape not enlarged or elongate; flagellum (Figs 30, 31) fil- iform, 1.0-3.3 times longer than head width, a single subapical rhinarium always present on flagellomeres 2, 4,6 and 7, sometimes also present on 1, 3 and 5; genae swollen ventrally; ultimate rostral segment 1.8-3.4 times longer than wide. Thorax, in profile, moderately arched, meso- praescutum sharply descending anteriorly, pro- notum hardly visible from above; forewing (Figs 57-59) narrowly obovate with acute apex, 2.4-2.7 times longer than wide, C+Sc thickened basally, costal break absent, elongate wedge-shaped pterostigma present, apex of M_,, reaching wing margin anterior to wing apex, M+Cu stem longer or shorter than Cu stem, m, cell value about 1.0, cu, cell value 1.4-1.7, apex of claval suture adja- cent to apex of Cu;,, radular areas diffuse or partly defined; basal spine of hind tibia present, apical spurs forming an incomplete ring with inner spurs D. HOLLIS & P. S. BROOMFIELD larger than outer spurs; hind basitarsus with 2 apical spurs. CO proctiger bipartite; lateral lobes well- developed, either elongate-narrow and without inner apical lobes (Figs 84, 85) or broad-robust with inner apical lobes (Figs 87, 90); basal seg- ment of aedeagus swollen in apical half, apical segment either elongate and simple (Figs 84, 85) or short and modified (Figs 87, 90). Q terminalia either short, with proctiger rounded apically and anal pore ring convoluted (Fig. 86), or conical with proctiger acute apically and anal pore ring simple (Fig. 89). Larva. The larvae of M. fici and M. proxima are described by Froggatt (1901). Dorsal and ventral caudal plates of M. kina and M. obliqua are fig- ured below (Figs 106, 107). Comments. Nine nominal species, including four described below, are included in the genus. How- ever, the three recorded names for the Indian species are probably synonymous. Two species- groups can be recognised. fici group Antennal flagellum elongate, with rhinaria on flagellomeres 2, 4, 6 and 7, and occasionally on 1; Cu stem longer than M+ Cu stem; aedeagus short and robust with modified apical segment; lateral lobes of CO’ proctiger elongate-narrow; 2 termi- nalia short and rounded apically, with convoluted anal pore ring. Seven nominal species are included in this group, distributed in India, Australia, New Guinea and New Caledonia on Ficus (Urostigma) sects Urostigma, Conosycea and Malvanthera. Apart from the three Indian ‘species’ they can be individually identified by the structure of the male genitalia. Mycopsylla gardenensis (Bhanotar, Ghosh & Ghosh) (Figs 12, 13, 57) Edenus gardenensis Bhanotar, Ghosh & Ghosh, 1972: 109. Holotype 2, INDIA: on Psidium guajava (ZSI) [not examined]. Mycopsylla gardenesis (Bhanotor, Ghosh & Ghosh) Rajamohan et al., 1975: 138. [Misspelling. | Mycopsylla gardenensis (Bhanotar, Ghosh & Ghosh); Hodkinson, 1986: 318; Kandasamy, 1987: 69. HostpLants. Ficus mollis, F. tsjahela (or F. vir- ens). F. microcarpa, possibly F. religiosa, FICUS-FEEDING PSYLLIDS doubtfully Psidium guajava. According to Rajamohan et al. (1975) the larvae cause leaf- margin rolls on F. mollis. RECORDED DISTRIBUTION. India: W. Bengal, Tamil Nadu, Orissa. MATERIAL EXAMINED India: Tamil Nadu, adults and larvae, on F. mol- lis; adults and larvae, on F. religiosa; Orissa, adults, on F. tsjahela (or F. virens). Bangladesh: adults. Singapore: larvae on F. microcarpa (BMNH). Mycopsylla indica Mathur Mycopsylla indica Mathur, 1975: 151; Hodkinson, 1986: 319; Kandasamy, 1987: 71, 72, 76. Holo- type 0, INDIA: on Santalum album (FRI) [not examined]. HostpLant. Recorded by Mathur on Santalum album but this is regarded here as a doubtful host record as the type series was collected during the ‘Sandal Insect Survey’ and many of these records are for vagrant specimens. RECORDED DISTRIBUTION. India: Tamil Nadu. No material examined. Comments. This species is probably not distinct from M. gardenensis. Mycopsylla mathuriana Kandasamy Mycopsylla mathuriana Kandasamy, 1987: 71. Holotype 9, INDIA: on Ficus religiosa (ZS1) [not examined]. HostpLant. Ficus religiosa. RECORDED DISTRIBUTION. India: Tamil Nadu. No material examined. Comments. This species is probably not distinct from M. gardenensis. Mycopsylla fici (Tryon) Psylla fici Tryon, 1895: 60. Syntypes adults, larvae and eggs, AUSTRALIA (Qld): on Ficus mac- rophylla (not traced). Mycopsylla fici (Tryon) Froggatt, 1901: 259; 1923: 146; Tuthill & Taylor, 1955: 248; Hodkinson, 1983: 354; Morgan, 1984: 110. Mycopsylla fici Froggatt; Aulmann, 1913: 30. HosteLant. Ficus macrophylla RECORDED DISTRIBUTION. Australia: NSW, QLD. MATERIAL EXAMINED Australia: NSW, QLD, Lord Howe Is. (BMNH). 149 Mycopsylla proxima Froggatt Mycopsylla proxima Froggatt, 1901: 261; Aulmann, 1913: 31; Tuthill & Taylor, 1955: 248; Morgan, 1984: 110. Syntypes larvae and adults, AUSTRALIA (NSW): on Ficus rubiginosa (2? ANIC) [not examined]. HostpLant. Ficus rubiginosa. RECORDED DISTRIBUTION. Australia: NSW. MATERIAL EXAMINED Australia: 2 O&', NSW, ‘on yellow fig’ BMNH). Mycopsylla kina sp. n. (Figs 30, 84, 106) Descrirtion. Moderately large psyllids, up to 6.5 mm. long. Vertex concave, without anterolateral tubercles or ridges. Antennal flagellum long, 3.0— 3.3 times longer than head width, Ist flagellomere (Fig. 30) with about 17 rhinaria ventrally in basal third, 2nd with 1 subapical rhinarium, 4th with 1 or 2, 6th with 1, 7th with 1, 8th with 1 short truncate and | long pointed seta apically; ultimate rostral segment 2.8 times longer than wide. Forewing 2.39-2.54 times longer than wide, M-+Cu stem shorter than Cu stem, m, cell value about 1.0, cu, cell value 1.7, radular areas diffuse, apex of claval suture close but not adjacent to apex of Cu,,; hindwing 0.5 times as long as forewing. CO proctiger (Fig. 84) with narrow, curved, strap-like lateral lobes without inner apical lobes; aedeagus as in Fig. 84; paramere conical (Fig. 84). Q terminalia short, bulbous; proctiger broadly rounded apically, 0.5 times as long as head width, circum-anal pore ring weakly convoluted; sub- genital plate broadly incised apically. Measurements (3 0’, 1 2). Maximum width of head, O 1.04-1.12, 2 1.18; length of antennal flagellum, O' 3.2-3.54, 2 3.52; length of ultimate rostral segment, O' 0.18, 2 0.20; length of fore- wing, O' 4.68-4.88, 9 6.10; length of hind tibia, 0 1.02-1.08, 2 1.10; length of 9 proctiger, 0.62. Larva. Dorsal and ventral caudal plates and circum-anal pore as in Fig. 106, marginal sec- tasetae on strongly raised bases. HostTPLant. Ficus sp., ‘hard fruited’. Holotype OC’, Papua New Guinea: E. Highlands, Aiyura, 5,800’, Ficus, hard fruited, under latex cover, 1958 (Barrett) (BMNH); slide-mounted. Paratypes. Papua New Guinea: 2 0’, 6 Q, lar- vae, same data as holotype (BMNH); slide- and dry-mounted. 150 ComMENTS. This species may be distinguished from other members of the fici group by the group of rhinaria present at the base of the lst flagellomere and the simple, non-bifid form of the male paramere. From the data accompanying the type series it would seem that the larvae of this species live beneath a cover composed of their dried anal exu- date, similar to both the described Australian spe- cies. The larvae of another member of the fici group, M. gardenensis, live within the leaf-margin rolls they induce in their host. This contrasts with the larvae of M. obliqua which are free-living. Mycopsylla tuberculata sp. n. (Figs 14, 15, 85, 86) Description. Medium-sized psyllids, up to 5.0 mm. long. Head (Figs 14, 15) with concave vertex and well-developed anterolateral tubercles. Antennal flagellum 2.5 times longer than head width, a single subapical rhinarium present on flagellomeres 2, 4, 6 and 7, 8th flagellomere with 1 long pointed medial seta and 1 short truncate api- cal seta; genae swollen ventrally; ultimate rostral segment 3.0 times longer than wide. Forewing 2.54-2.62 times longer than wide, M-+Cu stem shorter than Cu stem, m, cell value 1.0, cu, cell value 1.5, radular areas diffuse, apex of claval suture a little distant from apex of Cu,,; hindwing 0.6 times as long as forewing. © proctiger (Fig. 85) with strap-like, curved lateral lobes; aedeagus as in Fig. 85; paramere (Fig. 85) strongly bifid apically. ? terminalia (Fig. 86) short, rounded; proctiger broadly rounded apically, circum-anal pore ring weakly convoluted; subgenital plate broadly incised apically. Measurements (1 0’, 1 2). Maximum width of head, C' 0.8, 2 0.84; length of antennal flagellum, © 2.1; length of ultimate rostral segment, 0’ 0.15, ? 0.16; length of forewing, 3.46, 2 4.16; length of hind tibia, O’ 0.74, 2 0.76; length of 2 proc- tiger, 0.54. Larva and hostplant unknown. Holotype Co’, New Caledonia: Ile des Pins, 7 km N. Kuto, 30-100 m, 16.viti.1979 (Gagné) (BPBM); slide-mounted. Paratypes. New Caledonia: 1 2, Mts des Koghis, 400-600 m, i.1969; 1 9, ii.1978; 1 9, Poindimie, 0-50 m, 1.1969 (Krauss) (BPBM; BMNH),; slide- and dry-mounted. Comments. M. tuberculata differs from other members of the fici group in having well- developed anterolateral tubercles on the vertex and a distinctive aedeagal apex that bears a D. HOLLIS & P. S. BROOMFIELD medioventral spinulose lobe. In M. gardenensis the ventral lobe is convoluted and membraneous, and in fici and proxima the structure is bilobed. obliqua group Antennal flagellum short, with a single apical seta and a single apical flagellum on flagellomeres 1-7; Cu stem shorter than M+Cu stem; C’ aedeagus long and slender; lateral lobes of C’ proctiger broad, with inner apical lobes; 9 terminalia long, conical, acute apically, anal pore oval. Two closely related species are included here, one from New Caledonia on Ficus (Urostigma) Malvanthera; the other from Loyalty Is, with unknown trophic relationships. Mycopsylla obliqua sp. n. (Figs 16, 17, 31, 58, 87-89, 107) DescripTIon. Moderately large psyllids, up to 5.0 mm long. Head (Figs 16, 17) with vertex weakly concave and shallowly incised by median suture, anterolateral tubercles absent. Antennal flagellum (Fig. 31) 1.25 (0) and 1.0 (Q) times longer than head width, a single subapical rhi- narium present on flagellomeres 1-7 and a single, long, subapical seta present on flagellomeres 1-6, 8th flagellomere with 2 long setae apically; ulti- mate rostral segment 2.0 times longer than wide. Forewing (Fig. 58) 2.5-2.7 times longer than wide; R, short, one-third the length of R stem, Rs at least 3 times longer than R stem with its apex at the apex of the pterostigma, M+Cu longer than Cu stem, m, cell value 1.0, cu; cell value 1.4, apex of claval suture distant from apex of Cu,,, radular areas defined; hindwing almost 0.5 times as long as forewing. CO proctiger (Fig. 87) with broad lateral lobes; apex of aedeagus (Fig. 87) narrow; paramere (Fig. 88) thumb-shaped in profile, inner surface with a small median tubercle, an anteroapical ridge anda posteroapical tubercle. Q terminalia (Fig. 89) conical, proctiger 1.06— 1.08 times longer than head width, circum-anal pore ring simple. Measurements (3 CO’, 3 2). Maximum width of head, 0 0.92-0.98, 2 1.02-1.04; length of anten- nal flagellum, 0” 1.16-1.24, 2 1.00—1.04; length of ultimate rostral segment, O' 0.10, 9 0.12; length of forewing, 0’ 3.56-3.88, 2 4.16—4.32; length of hind tibia, O’ and 9, 0.70-0.72; length of 2 proc- tiger, 1.08-1.10. Larva. Dorsal and ventral caudal plates and circum-anal pore ring as in Fig. 107; marginal setae of caudal plates not on raised tubercles. FICUS-FEEDING PSYLLIDS HostPLant. Ficus obligua. Holotype OC, New Caledonia: Noumea, ORSTOM Centre, 20-24.iii.1982, Ficus obliqua (Hollis) (BMNH); dry-mounted. Paratypes. New Caledonia: 6 ©’, 8 Q, larvae, same data as holotype (BMNH; MNHN); slide- and dry-mounted. Mycopsylla propinqua sp. n. (Figs 59, 90, 91) DeEscriPTION. Very similar to M. obliqua but dif- fering in forewing venation (Figs 58, 59), and in the structure of the O’ proctiger (Figs 87, 90), aedeagus (Figs 87, 90) and paramere (Figs 88, 91). Antennal flagellum 1.51 times longer than head width; ultimate rostral segment 1.8 times longer than wide. Forewing (Fig. 59) 2.72 times longer than wide; R_half as long as R stem, Rs just over twice as long as R stem with its apex reaching the posterior margin of the pterostigma proximal to the latter’s apex, m, cell value 1.0, cu, cell value 1.59; hind- wing 0.35 times as long as forewing. CO proctiger (Fig. 90) with lateral lobes narrow basally and broadening apically; aedeagus (Fig. 90) bulbous apically; paramere (Fig. 91) with obliquely truncate apex, inner surface with an anterior tubercle in apical third and 2 parallel ridges anteroapically. ? unknown. Measurements (1 0’). Maximum width of head, 1.06; length of antennal flagellum, 1.60; length of ultimate rostral segment, 0.11; length of forewing, 4.68; length of hind tibia, 0.82. Larva and hostplant unknown. Holotype oO’, Loyalty Is: We, Lifou I., 16-18.1i1.1963, light trap (Yoshimoto) (BPBM); slide-mounted. MACROHOMOTOMINI Macrohomotomini White & Hodkinson, 1985: 242, 272. Type genus: Macrohomotoma Kuwayama. DiAGnosIs. Pterostigma short, ovoid; M,,, curved and reaching wing margin posterior to wing apex; basal segment of aedeagus not swollen in apical half. Two genera are included here, the Oriental Macrohomotoma and the Afrotropical Pseu- doeriopsylla. These genera were synonymi- sed by Crawford (1914) and the synonymy was accepted by Enderlein (1921). White & Hodkinson (1985) treated them as separate gen- era and this is accepted here as both groups can be 151 distinguished on adult and larval characteristics (see key couplet 8, p. 141), and have a clear geographical and hostplant separation. MACROHOMOTOMA Kuwayama Macrohomotoma Kuwayama, 1907: 179; Crawford, 1911: 483; 1914: 62; 1919: 157; 1925b: 36; Aulmann, 1913: 36; Enderlein, 1914: 233; 1921: 119; Ramakrishna Ayyar, 1924: 622; Boselli, 1930: 178; Kuwayama, 1931: 125; Heslop-Harrison, 1958: 578; 1960: 161; Log- inova, 1972: 843; Bekker-Migdisova, 1973: 102; Mathur, 1975: 135; Hodkinson, 1983: 352; 1986: 316; Yang, 1984: 156; Yang & Li 1984b: 370. Type species: Macrohomotoma gladiata Kuwayama, by original designation and monotypy. DEscrIPTION. Large psyllids, up to 9.5 mm long. Integument of head and thorax sparsely covered with very short setae. Head (Figs 18, 19), from above, as wide as mesoscutum; disc of vertex weakly concave, foveae forming transverse depressions, anterior and lateral margins rounded, occipital margin angular, lateral ocelli not on raised tubercles, median ocellus just visible from above, anterolateral tubercles absent; antennal sockets not enlarged and head without cleft appearance; antennal scape not enlarged or elongate; flagellum short, filiform, 0.7—0.9 times as long as head width, rhinaria present on flagellomeres 1-7; genae swollen ventrally; ulti- mate rostral segment about twice as long as wide. Thorax, in profile, strongly arched, pronotum visible from above; forewing (Fig. 60) ovate, with subacute apex, 2.2—2.6 times longer than wide, veins bearing short setae, C+Sc not thickened, costal break absent, ovoid pterostigma present, apex of M,,, reaching wing margin posterior to wing apex, Cu stem about as long as M+ Cu stem, m, cell value about 1.8, cu; cell value about 1.0, apex of claval suture adjacent to apex of Cuj,, radular areas well-defined; basal spine of hind tibia absent, apical spurs arranged 0 + 4, hind basitarsus with 2 apical spurs. CO proctiger bipartite, lateral lobes well- developed and each with an inner apical lobe; aedeagus 2-segmented; basal segment weakly swollen subapically. Q terminalia conical, anal pore ring normally simple oval, rarely convoluted. Larva. Anus and associated wax pores ventral (Fig. 108). Hoste ants. Ficus (Urostigma) sect. Conosycea. 152 ComMENTS. Fourteen species are currently recog- nised in this genus, distributed from India to N. Queensland. However, most of these are poorly described and the genus requires revision. Macrohomotoma apsylloides (Crawford) Pauropsylla apsylloides Crawford, 1919:144. Syn- types CO and @, INDONESIA, EAST MALAYSIA, MACAO, HAWAII (USNM) [not examined]. Macrohomotoma apsylloides (Crawford) Craw- ford, 1920: 353; 1925a: 36; Heslop- Harrison, 1958: 570; 1960: 161; Loginova, 1972: 840; Mathur, 1975: 139; Hodkinson, 1983: 352; 1986: 316. HostTpLant. Unknown. RECORDED DISTRIBUTION: Indonesia: Tanimbar Is. East Malaysia: Sabah. Macao and Hawaii. No material examined. Macrohomotoma geniculata Mathur Macrohomotoma geniculata Mathur, 1975: 136; Hodkinson, 1986: 316. Holotype 0’, INDIA: on Ficus microcarpa (FRI) [not examined]. HostTpLant. Ficus microcarpa. RECORDED DISTRIBUTION. India (Uttar Pradesh, Karnataka). MATERIAL EXAMINED India: Karnataka, adults and larvae on Ficus sp. (BMNH). Macrohomotoma gladiata Kuwayama (Figs 18, 19, 60, 108) Macrohomotoma gladiatum Kuwayama, 1907: 180; Crawford, 1911: 490; 1914: 63; 1925a: 37; [1928: 425, misidentification]; Aulmann, 1913: io JEhacleniein, iGilé'e Bae iil 11) Kuwayama, 1922: 368; 1931: 125; Boselli, 1930: 178; Miyatake, 1965a: 173; Mathur, 1975: 135; Hodkinson, 1983: 352. Holotype 9, TAIWAN (EIHU) [not examined]. Macrohomotoma gladiata Kuwayama; Yang, 1984: 157. [Macrohomotoma striata Crawford; Hill & Cheung, 1978: 51; Hill et al., 1982: 175. Misidentifications. | HostTpLant. Ficus microcarpa (many records as F. retusa). RECORDED DISTRIBUTION. Japan (Ryukyu Is), Tai- wan. Crawford’s (1928) record of this species from D. HOLLIS & P. S. BROOMFIELD Sumatra is erroneous as a male _ bearing Crawford’s determination label in BMNH repre- sents a different species. MATERIAL EXAMINED Hong Kong: adults on Ficus microcarpa; adults and larvae on F. ? benghalensis. (BMNH). Macrohomotoma hylocola Yang & Li Macrohomotoma hylocola Yang & Li, 1984b: 376, 380; Hodkinson, 1986: 316. Holotype ©C’, CHINA: on Ficus sp. (BAUIC) [not examined]. HostTPLant. Ficus sp. RECORDED DISTRIBUTION. China: Yunnan. No material examined. Macrohomotoma maculata Mathur Macrohomotoma maculata Mathur, 1975: 140; Hodkinson, 1986: 316. Holotype 2, INDIA: on Santalum album (FRI) [not examined]. HostpLant. The type series was probably vagrant on the recorded host. RECORDED DISTRIBUTION. India: Tamil Nadu, Karnataka. MATERIAL EXAMINED India: 1 O', 2 2, Tamil Nadu (same locality as holotype, BMNH). Macrohomotoma magna Yang & Li Macrohomotoma magna Yang & Li, 1984b: 375; Hodkinson, 1986: 316. Holotype 9, CHINA: on Ficus sp. (BAUIC) [not examined]. Macrohomotoma magne Yang & Li, 1984b: 380. [Misspelling. ] HostTPLant. Ficus sp. RECORDED DISTRIBUTION. China: Yunnan. No material examined. Macrohomotoma minana Yang & Li Macrohomotoma minana Yang & Li, 1984b: 373, 380; Hodkinson, 1986: 316. Holotype oC, CHINA: on Ficus sp. (BAUIC) [not examined]. HostTpLant. Ficus sp. RECORDED DISTRIBUTION. China: Fujian. No material examined. FICUS-FEEDING PSYLLIDS Macrohomotoma robusta Y ang Macrohomotoma robusta Yang, 1984: 160; Hodkinson, 1986: 316. Holotype Oo’, TAI- WAN: on Ficus benjamina (NCHU) [not examined]. HostpLant. Ficus benjamina. RECORDED DISTRIBUTION. Taiwan. No material examined. Macrohomotoma sandakana Crawford [Pauropsylla apsylloides Crawford, 1919: 144. Misidentification, in part. | Macrohomotoma sandakana Crawford, 1925Sa: 38; Hodkinson, 1983: 352. Holotype 0’, EAST MALAYSIA (USNM) [not examined]. HosteiLant. Unknown. RECORDED DISTRIBUTION. East Malaysia: Sabah. No material examined. Macrohomotoma sinica Yang & Li Macrohomotoma sinica Yang & Li, 1984b: 371, 379; Hodkinson, 1986: 316. Holotype OC, CHINA: on Ficus spp. (BAUIC) [not examined]. Hoste ant. Ficus microcarpa (as F. retusa), Ficus sp., ? F. microphylla [not listed by Corner (1965)]. RECORDED DISTRIBUTION. China: Fujian. No material examined. Macrohomotoma striata Crawford Macrohomotoma striata Crawford; Ramakrishna Ayyar, 1924: 622, nomen nudum. Macrohomotoma striata Crawford, 1925a: 38; [Miyatake, 1965a: 174, ? misidentification]; Mathur, 1975: 144; Hodkinson, 1983: 352; 1986: 316. Holotype 2, INDIA: on Ficus sp. (? USNM) [not examined]. HostTpLant. Ficus sp.; Miyatake’s (1965a) record of this species from the Ryukyu Is on F. micro- carpa (as FF. retusa) is probably a misidentification. RECORDED DISTRIBUTION. India: Karnataka. Japan: Ryukyu Is [possible misidentification by Miyatake (1965a)]. No material examined. 153 Macrohomotoma viridis Yang & Li Macrohomotoma viridis Yang & Li, 1984b: 377, 380; Hodkinson, 1986: 317. Holotype 9, CHINA: on Ficus sp. (BAUIC) [not examined]. HostTpLant. Ficus sp. RECORDED DISTRIBUTION. China; Yunnan. No material examined. Macrohomotoma williamsi Crawford Macrohomotoma williamsi Crawford, 1925a: 37; Hodkinson, 1983: 352. Syntypes, CO’ and Q, PHILIPPINES: on Ficus clementis (USNM) [2 © examined]. HostTeLant. Ficus crassiramea var. clementis. RECORDED DISTRIBUTION. Philippines. MATERIAL EXAMINED Philippines: 2 9 syntypes (USNM). Macrohomotoma yunana Yang & Li Macrohomotoma yunana Yang & Li, 1984b: 374, 380; Hodkinson, 1986: 317. Holotype CO, CHINA: on Ficus sp. (BAUIC) [not examined]. HostTpLant. Ficus sp. RECORDED DISTRIBUTION. China: Yunnan. No material examined. PSEUDOERIOPSYLLA Newstead Pseudoeriopsylla Newstead, 1911: 105; Yang & Li, 1984b: 370 (as a synonym of Mac- rohomotoma); White & Hodkinson, 1985: 242. Type species: Pseudoeriopsylla nyasae News- tead, by monotypy. Pseuderiopsylla Newstead; Crawford, 1914: 62 (as a synonym of Macrohomotoma). [Misspelling. ] DescriPTION. Large psyllids, up to 8.0 mm long. Integument of head and thorax sparsely covered with very short setae. Head (Figs 20, 21), from above, almost as wide as mesoscutum; disc of vertex concave with foveae weakly defined, ante- rior and lateral margins rounded, occipital margin angular, lateral ocelli not on raised tubercles, median ocellus just visible from above, ante- rolateral tubercles absent; antennal sockets not enlarged and head without cleft appearance, antennal scape not enlarged or elongate; flagellum filiform, 1.0—1.7 times longer than head 154 width, rhinaria present on flagellomeres 2-7 (Figs 32-37); (sometimes absent from 5); genae slightly swollen ventrally; ultimate rostral segment short, 2.0-4.0 times longer than wide. Thorax, in profile, moderately arched; pro- notum narrowly visible from above and descend- ing vertically behind occiput; forewing (Figs 61, 62) ovate, with subacute apex, 2.15—2.45 times longer than wide, veins bearing sparse short setae, C+Sc slightly thickened proximally, costal break absent, well-defined subcircular pterostigma pre- sent, apex of M,,, reaching wing margin posterior to wing apex, Cu stem much shorter than M+Cu stem, m, cell value 1.5—1.9, cu, cell value 0.7-1.2, apex of claval suture adjacent to apex of Cuj,, radular areas well-defined; basal spine of hind tibia present, apical spurs arranged 0 + 4, hind basitarsus with 2 apical spurs. CO proctiger bipartite, lateral lobes well- developed, each with a small inner apical lobe; aedeagus 2-segmented, basal segment not swollen. Q terminalia conical, anal pore ring convoluted. Larva. Anus and associated wax pores termi- nodorsal (Figs 109-112). HostpLants. Ficus (Urostigma) sect. Galo- glychia. ComMENTS. Crawford (1914) considered this genus to be synonymous with Macrohomotoma but White & Hodkinson (1985) regarded the two as distinct. Although clearly a sister-pair there are good characters in both larvae and adults that separate the two genera (see key couplet 8, p. 141). Until recently Pseudoeriopsylla was thought to be monobasic but a critical examination of the available material has revealed a complex of at least six species in tropical Africa. These are dif- ferentiated and described below. Key to species of Pseudoeriopsylla 1 Rhinaria present over whole surface of Ist tlagellomenrc)(Eigs 38-35) eens 2 — Rhinaria present only apically or subapically on 1st tlagellomere|(Figsis25.465)57) eee ereeeeee eee 4 2 Rhinaria present over whole lengths of flagellomeres 2-5 (Fig. 35); O’ paramere (Fig. 96) obliquely trun- cate apically; 2 proctiger not less than 1.4 times longer than head width................. carvalhoi — Rhinaria present only subapically on flagellomeres 2-5 (Figs 33, 34); O&' paramere (Figs 93, 95) rounded apically; 2 proctiger not more than 1.3 times longer thantheadiwidth sas antec etna 3 D. HOLLIS & P. S. BROOMFIELD 3 CO paramere (Fig. 95) narrow, thumb-like, anterior tubercle on inner surface hook-like........ medleri — © paramere (Fig. 93) broader, anterior tubercle on INNELSULia Ce Gun CaLCmE san eet ene laingi 4 OC paramere (Fig. 92) conical, with a subacute apical tubercle; 1st and 2nd flagellomeres with several rhi- Natialsubapically (Hig= 32). 4-4. anne nyasae — © paramere thumb-shaped (Figs 97, 98), with two tubercles on inner surface subapically; 1st and 2nd flagellomeres each with a single subapical rhinarium (EIoS;SO 31) nse ceo Sade ge 3d Seo Re oa 5 5 Antennal flagellum not less than 1.4 times longer than head width; C’ paramere broad (Fig. 97) kenyae — Antennal flagellum not more than 1.25 times longer than head width; CO paramere narrow (Fig. 98) etiennei Pseudoeriopsylla nyasae Newstead (Figs 32, 92, 109) Pseudoeriopsylla nyasae Newstead, 1911: 105; Yang & Li, 1984b: 369; White & Hodkinson, 1985: 162. Syntypes 9 and larvae, MALAWI: on Ficus sp. (larvae BMNH, @ not traced) [examined]. Macrohomotoma nyasae (Newstead) Crawford, 1914: 63; 1920: 354; 1925a: 36; Enderlein, 1921: 119: HostTPLant. Subsect. Chlamydodorae, Ficus thon- ningil (some records as F. petersii). The type series was recorded from ‘Kachire’ which, according to Binns (1972), is a vernacular name used for Ficus natalensis, F. thonningii, and F. scassellatii (=F. kirkii). RECORDED DISTRIBUTION. Malawi, Mozambique. MATERIAL EXAMINED Adults and larvae from: South Africa (Tvl), on” petersii’; Malawi, Tanzania, Zaire. Pseudoeriopsylla laingi sp. n. (Figs 20. 21, 33, 61, 93, 94, 110) Description. Antennal flagellum 1.46-1.70 (C’) and 1.19-1.28 (Q) times longer than head width; rhinaria present over most of surface of 1st flagellomere, 3 subapically on 2nd (Fig. 33), 0-2 subapically on 3rd, 1—2 subapically on 4th, none on 5th, 34 subapically on 6th, 2 subapically on 7th, 8th bearing 2 subequal setae apically; ulti- mate rostral segment 2.5-3.0 times longer than wide. Forewing 2.20—2.45 times longer than wide, pattern dimorphic (as in Figs 61, 62); 7m, cell value FICUS-FEEDING PSYLLIDS 1.5-1.7, cu, cell value 0.7-1.1 (higher in W. African populations). CO paramere (Fig. 93) broad in profile, rounded apically, inner surface with 2 subapical tubercles, the anterior one truncate apically; apical segment of aedeagus as in Fig. 94. Q proctiger 1.20-1.27 times longer than head width. Measurements (6 0’, 5 9). Maximum width of head, 0’ 0.97-1.12, 9 1.06-1.20; length of anten- nal flagellum, CO” 1.42-1.88, 9 1.30-1.42; length of ultimate rostral segment, CO’ 0.16-0.19, 9 0.15- 0.20; length of forewing, O& 4.12-5.95, 2 4.80- 6.68; length of hind tibia, O’ 0.88-1.10, 2 0.92- | 1.10; length of 9 proctiger, 1.28-1.52. Larva. Dorsal and ventral caudal plates and _ circum-anal pore ring as in Fig. 110. | HosTpvant. Subsect. Chlamydodorae; Ficus thon- _ ningii, F. natalensis . Holotype ©’, Angola: Chianga, 12.iv.1975, on Ficus thonningii (van Harten) (BMNH); dry- mounted. Paratypes. Angola: 5 ©’, 8 Q, larvae, same data as holotype; 1 @. i.1971 (van Harten); 1 Q, 21-24.iii.1972 (Hollis); 2 oO, 2 9, 27.viii.1973 (Amorim). Kenya: 2 0’, 5 @, larvae, Nairobi, Chiromo, ix.1970, Ficus sp. (Schmutterer). | Uganda: 1 ©’, Kampala, ‘Makawkawe’, 9.iii.1925 (Hamwer); 1 0’, 3 9, larvae, Kawanda Res. Stn, 10.vi.1988, on roots of Ficus natalensis (Pol- aszek). Cameroon: 1 9, Bamenda, ¢ 5000’, 26-31.1.1957, yellow tray; 1 ©’, ii.1957 (Eastop). Nigeria: 1 0’, 5 2, Lagos (Brown). Sierra Leone: } larvae, Freetown, 27.viii.1924; 1 oO, 2 Q, | 20.1.1927;2 0", 2 9, Newton, 10.xi.1928, Ficus sp. | (Hargreaves). Guinea: 1 ©’, Kindia, Fulaya, | 4.i1.1952, “Goyavier’ (Balachowsky). Senegal: lar- vae, Djibelor, 15.xii.1981, Ficus sp.; 1 0’, 1 9, 14.11.1982, beating trees (Etienne). (BMNH; | MNHN). ComMENTS. This species appears to be close to medleri and nyasae and replaces the latter on the same host from Kenya northwards and ' westwards. Dr A. Polaszek (pers. comm.) has reared speci- mens of Psyllaephagus secus Prinsloo (Chal- cidoidea: Encyrtidae) and Dilyta sp. (Cynipoidea: _ Charipidae) from Sth instar larvae of the popula- tion from Kawanda Res. Stn, Uganda, mentioned | in the above paratype series. Pseudoeriopsylla medleri sp. n. (Figs 34, 95) Description. Antennal flagellum 1.32-1.41 (C’) | and 1.14~1.16 (Q) times longer than head width; 155 rhinaria present over most of surface of Ist flagellomere (Fig. 34), 6 subapically on 2nd, 1-3 subapically on 3rd, 1—2 subapically on 4th, 1 api- cally on 5th, 3 subapically on 6th, 2 apically on 7th, 8th bearing 2 subequal setae apically; ultimate rostral segment 2.3-3.1 times longer than wide. Forewing pattern similar to that in Fig. 62, 2.18-2.34 times longer than wide, m, cell value 1.6-1.9, cu, cell value 1.13—1.19; hindwing 0.5 times as long as forewing. CO paramere (Fig. 95) narrow in profile, rounded apically, inner surface with 2 subapical tubercles, the anterior one hook-like. ? proctiger 1.2 times longer than head width. Measurements (4 O’, 2 2). Maximum width of head, CO 0.91-0.98, 9 1.02; length of antennal flagellum, CO 1.20-1.38, 2 1.16-1.18; length of ultimate rostral segment, O' 0.12-0.14, 9 0.13; length of forewing, CO 4.28-4.64, 2 4.72-4.76; length of hind tibia, O’ and 9 0.86-0.88; length of Q proctiger, 1.20-1.22. Larva and hostplant unknown. Holotype CO’, Nigeria: S. E. State, Oban, 7.iv.1975 (Medler) (BMNH); dry-mounted. Paratypes. Nigeria: 13 0’, 4 2, same data as holotype (BMNH); slide- and dry- mounted. Comments. P. medleri appears to be close to laingi but is distinguished by the structure of the paramere. Pseudoeriopsylla carvalhoi sp. n. (Figs 35, 96) Description. Antennal flagellum 1.35-1.67 (C’) and 1.14~-1.41 (Q) times longer than head width; rhinaria present over most of surfaces of flagellomeres 1-5 (Fig. 35), 4-6 subapically on 6th, 2 apically on 7th, 8th with 2 subequal setae apically; ultimate rostral segment 3.5—4.0 times longer than wide. Forewing pattern similar to that in Fig. 61, often with hind margin also brown, 2.20-2.41 times longer than wide, m, cell value 1.7, cu, cell value 0.9; hindwing 0.54—0.58 times as long as forewing. OC paramere (Fig. 96) with obliquely truncate apex, inner surface with 1 hook-like anterodorsal and 1 simple posterodorsal tubercle. Q proctiger long, 1.42-1.77 times longer than head width. Measurements (2 CO’, 2 2). Maximum width of head, o 0.90-0.92, 2 1.00-1.06; length of anten- nal flagellum, O 1.22-1.54, 9 1.14-1.50; length of ultimate rostral segment, oO 0.15-0.16, 9 0.16— 0.17; length of forewing, CO’ 3.80-3.96, 2 4.42- 4.72; length of hind tibia, O’ 0.82-1.02, 2 0.82- 1.14; length of 2 proctiger, 1.42-1.88. Larva unknown. 156 HostTpLant. Subsect. Caulocarpae, Ficus ovata (=F. brachypoda). Holotype co’, Angola: Duque de Braganca Falls, 11-12.111.1972, Ficus brachypoda (Hollis) (BMNH); dry-mounted. Paratypes. Angola: 4 0’, 5 Q, same data as holotype (BMNH). Zaire: 6 0’, 4 9, P.N. G., Miss. H. De Saeger, Iso II/9, 17.vi.1952 (3638); 3 o7,4 9, Mt Tungu (S), 9.vi.1952 (De Saeger 3606) (MRAC; BMNH). Nigeria: 1 9, Zaria, Samaru, 26.11.1966, m.v. trap (Deeming) (BMNH). Slide- and dry-mounted. Comments. P. carvalhoi differs from the other known species of the genus in the extreme pro- liferation of rhinaria on the antennal flagellum and the elongate female proctiger. The host spe- cies also belongs to a different subsection of Sect. Galoglychia. Pseudoeriopsylla kenyae sp. n. (Figs 36, 97, 111) Description. Antennal flagellum 1.53-1.57 (C’) and 1.38 (Q) times longer than head width; a single apical rhinarium present on Ist flagellomere, 1 on 2nd (Fig. 36), 0-1 on 3rd, 1-2 on 4th, none on Sth, 1-2 on 6th, a double rhi- narium on 7th, 8th with 2 subequal setae apically; ultimate rostral segment 3.0—3.6 times longer than wide. Forewing pattern as in Fig. 61, 2.25—2.36 times longer than wide, m, cell value 1.7, cu, cell value 1.2; hindwing 0.55—0.57 times as long as forewing. © paramere (Fig. 97) broad in profile, with rounded apex, inner surface with 1 subapical ante- rior tubercle and 1 posteroapical tubercle. Q proctiger 1.2 times longer than head width. Measurements (2 CO’, 1 2). Maximum width of head, Oo 1.08-1.10, 2 1.10; length of antennal flagellum, ’ 1.68-1.70, 2 1.52; length of ultimate rostral segment, CO’ 0.18-0.19, 9 0.18; length of forewing, O' 4.72-4.80, 9 5.13; length of hind tibia, O’ 1.14-1.20, 9 1.12; length of 2 proctiger, is Larva. Dorsal and ventral caudal plates and circum-anal pore ring as in Fig. 111. HostPLant. Ficus sp. Holotype OC’, Kenya: L. Naivasha, W. shore road, ¢ 6,200’, 21—22.vi.1974, Ficus sp. (Hollis) (BMNH); dry-mounted. Paratypes. Kenya: 1 0’, 3 Q, larvae, same data as holotype; 1 0’, 2 9, Nairobi, Karen, ¢ 5,500’, 7.vil.1974, Clausena anisata (Hollis) (BMNH); slide- and dry- mounted. D. HOLLIS & P. S. BROOMFIELD Comments. P. kenyae and etiennei differ from other species of the genus in the lack of multiple rhinaria on the first two flagellomeres. They differ from one another in the relative lengths of the antennal flagellum and the form of the male paramere. Pseudoeriopsylla etiennei sp. n. (Figs 37, 62, 98, 112) Description. Antennal flagellum 1.17—1.24 (Cc) and 1.0() times longer than head width; a single apical rhinarium on Ist flagellomere, 1 on 2nd (Fig. 37), none on 3rd, 1 on 4th, none on Sth, 1 on 6th, a double rhinarium apically on 7th, 8th with 2 subequal setae apically; ultimate rostral segment short, 2.0 times longer than wide. Forewing (Fig. 62) 2.15—2.31 times longer than wide, m, cell value 1.9, cu, cell value 1.05; hind- wing 0.5 times as long as forewing. CO paramere (Fig. 98) narrow in profile, with rounded apex, inner surface with a bifid tubercle anteroapically. Q proctiger 1.2 times longer than head width. Measurements (2 0’, 2 9). Maximum width of head, oO’ 1.08-1.16, 2 1.20-1.24; length of anten- nal flagellum, CO’ 1.34-1.36, 9 1.20-1.24; length of ultimate rostral segment, O’ and Q , 0.12; length of forewing, O' 4.81-4.89, 2 5.09-5.30; length of hind tibia, 0’ 0.94, 2 0.96-1.02; length of 2 proc- tiger, 1.41.5. Larva. Dorsal and ventral caudal plates and circum-anal pore ring as in Fig. 112. HostpLant. Ficus sp. Holotype ©’, Senegal: Kamobeul-Essyhl, 27.x1.1982, Ficus sp. (Etienne) (MNHN)); slide- mounted. Paratypes. Senegal: 1 0’, 2 9, larvae, same data as holotype (BMNH); slide-mounted. HOMOTOMINAE Heslop-Harrison Homotomini Heslop-Harrison, 1958: 577, in part. Homotominae Bekker-Migdisova, 1973: 101, in part; White & Hodkinson, 1985: 272. Carsidarinae Yang, 1984: 168, in part. Diacnosis. Antennal flagellum thickened and densely setose; in forewing costal break present or at least indicated by a weakly sclerotised area of the cuticle in the break area, pterostigma absent. The subfamily comprises two tribes, the Homo- tomini with an Old World distribution, and the Synozini with a New World distribution. FICUS-FEEDING PSYLLIDS HOMOTOMINI Heslop-Harrison Homotomini Heslop-Harrison, 1958: 578, in part; Loginova, 1964b: 54. Dynopsyllini Bekker-Migdisova, 1973: 102, in part. Psausiini Bekker-Migdisova, 1973: 102. Type genus: Psausia Enderlein. Syn. n. Diacnosis. In forewing M+Cu stem present, sometimes immediately adjacent to R+M+Cu stem and R stem, radular area present in cell mp; hind tibia with only part of apical spur ring thick- ened; CO proctiger bipartite, basal segment of aedeagus strongly expanded in apical half. A single, Old World genus, Homotoma, is recognised, with hostplants in Ficus (Urostigma) sects Urostigma, Conosycea and Galoglychia, and Ficus (Ficus) sects Ficus and Rhizocladus. HOMOTOMA Guérin-Méneville | Homotoma Guérin-Méneville, 1844: 376; Flor, | 1861: 337, 412; Low, 1879: 607; Kieffer, 1905: 161; Kuwayama, 1908: 181; Crawford, 1911: 483; 1915: 262; 1919: 162; Aulmann, 1913: 35; Enderlein, 1921: 118; Boselli, 1929: 218; Kuwayama, 1931: 124; Haupt, 1935: 252; Schaefer, 1949: 56; Ramirez Gomez, 1956: 65; Heslop-Harrison, 1958: 578; Dobreanu & Man- olache, 1962: 364; Miyatake, 1975: 17; Braza & Calilung, 1981: 353; Yang & Li, 1981: 77, 84; 1984a: 201, 217; Hodkinson, 1983: 349; 1986: 312. Type species: ‘Psylla ficus’ Linnaeus, by original designationand monotypy. Anisostropha Foerster, 1848: 92; Frauenfeld, | 1867: 804; Meyer-Diir, 1871: 403. Type species: Chermes ficus Linnaeus, by monotypy. [Objec- tive synonym of Homotoma. | Psausia Enderlein, 1914: 232; 1921: 120; Heslop- Harrison, 1949: 376, 378; 1958: 578; Mathur, 1975: 155; Yang & Li, 1981: 78, 85; 1984a: 217; Hodkinson, 1983: 349; 1986: 312; Yang, 1984: 168; Kandasamy, 1987: 72. Type species: Homotoma radiata Kuwayama, by original des- ignation and monotypy. [Synonymised with | _Homotoma by Kuwayama, 1931: 124.] | Labobrachia Enderlein, 1921: 119; Heslop-Harri- ) son, 1949: 376, 378; 1958: 578; Hodkinson, 1983: 349; 1986: 312; Yang & Li, 1984a: 201, 217. Type species: Homotoma __ pacifica Crawford, by original designation and mono- typy. [Synonymised with Homotoma by Hodkinson, 1983: 349.] | Metapsausia Enderlein, 1921: 120; Heslop-Harri- son, 1949: 376, 378; 1958: 578; Yang & Li, 1981: 79, 85; 1984a: 202; Hodkinson, 1983: 349; 1986: iVSy// 312. Type species: Homotoma _ bakeri Crawford, by original designation. [Syn- onymised with Homotoma by Hodkinson, 1983: 349.] Caenohomotoma Yang & Li, 1981: 78, 85. Type species: Caenohomotoma spiraea Yang & Li, by original designation. [Synonymised with Homotoma by Hodkinson, 1986: 312.] Austrohomotoma Yang & Li, 1981: 78, 85. Type species: Homotoma gressitti Miyatake, by monotypy. [Synonymised with Homotoma by Hodkinson, 1986: 312.] Harrisonella Yang & Li, 1981: 78, 85. Type spe- cies: Homotoma distincta Crawford, by mono- typy. [Synonymised with Homotoma_ by Hodkinson, 1986: 312.] Heterohomotoma Yang & Li, 1981: 78, 85. No type species designated. Nomen nudum. Labobracha Yang & Li, 1984a: 217. [Misspelling. } Psausia Yang & Li, 1984a: 217 (nec Enderlein). Type species: Homotoma distincta Crawford, by original designation and monotypy. [Hom- onym of Psausia Enderlein, objective synonym of Harrisonella, and syn. n.| Australohomotoma Hodkinson, [Misspelling. | LO8G: eS: Description. Medium- to large-sized psyllids, up to 6.5 mm. Integument of head and thorax densely covered with long setae. Head (Figs 22-24), from above, about as wide as mesoscutum; vertex from shallowly to deeply concave, anterior margin rounded and often deeply incised by median suture, lateral margins rounded or obtuseangular, occipital margin angular, median ocellus just visi- ble from above, lateral ocelli on weak swellings, anterolateral tubercles absent, antennal sockets enlarged and giving head a cleft appearance; antennal flagellum (Figs 38—40) swollen and den- sely hirsute, rounded or flattened in cross-section, a single apical rhinarium present on flagellomeres 2,4, 6 and 7, flagellomere 8 very reduced and less than half length of 7; genae normally slightly swollen ventrally, small genal cones rarely developed; ultimate rostral segment short, up to twice as long as wide. Thorax, in profile, from weakly to strongly arched; pronotum broadly visible from above, sometimes with a pair of anteromedial and a pair of anterolateral projections. Forewing (Figs 63-73) of variable shape, veins bearing long setae, C+ Sc usually thickened basally, costal break pre- sent or indicated by weakening of cuticle in break area, M stem often partially fused with basal part of Rs and M,,, and sometimes partially fused with apical part of Rs, apex of M,,, reaching wing margin anterior to wing apex or at wing apex, Cu 158 stem short or long, cells 7m, and cu, of very variable shape, apex of claval suture adjacent to or shortly distant from apex of Cu,,, radular areas from diffuse to sharply defined but always present in cell m,; basal spine of hind tibia rarely present, apical spurs arranged 0 + 4~7; hind basitarsus with 2 or rarely 1 apical spurs. © proctiger bipartite, lateral lobes well- developed and with inner apical lobes; basal seg- ment of aedeagus swollen in apical half, apical segment without ventral spiniform processes api- cally (Figs 99-104). Q terminalia conical, circum-anal pore ring simple. Larva. Body form variable, wider than long in H. ficus, longer than wide in other known species. Antennae short, flagellum not subdivided except for flagellomeres 7 and 8; dorsal thoracic sclerites differentiated, separate; humeral lobes well- developed in ficus but not in other known species; body surface covered with lanceolate setae, in ficus these are mounted on elongate tubercles (White & Hodkinson, 1985), in other species these tubercles are smaller or absent; pointed sec- tasetae present marginally; anus ventral and with a large, medially constricted circum-anal pore ring. [Larvae are described or figured for distincta (Heslop-Harrison, 1949), ficus (Boselli, 1929), galbvittata (Yang & Li, 1984a), indica (Mathur, 1975), maculata (Yang, 1984), radiata (Fang & Yang, 1986) and wulinensis (Yang, 1984).] Comments. The synonymy of Homotoma has become confused since Enderlein erected the gen- era Psausia, Metapsausia and Labobrachia, based on type species previously described in Homo- toma. Kuwayama (1931) synonymised Psausia with Homotoma but Heslop-Harrison (1949) recognised all four genera as distinct, and Mathur (1975) and Yang (1984) regarded Psausia as dis- tinct. Miyatake (1975) included the type species of Enderlein’s genera in his treatment of Homo- toma, but it is likely that he was following Kuwayama in the synonymy of Psausia and was unaware of Enderlein’s (1921) paper erecting Metapsausia and Labobrachia. However, Hodkinson (1983) considered that Miyatake had effectively synonymised Enderlein’s genera with Homotoma. This confusion was further com- pounded by Yang & Li (1981, 1984a) when they erected a further five genus-group names in the Homotoma complex, including one that is both a homonym and an objective synonym, and another that is a nomen nudum. Hodkinson (1986) con- sidered Yang & Li’s names to be invalid and fur- ther suggested that there was no good reason for splitting Homotoma. D. HOLLIS & P. S. BROOMFIELD Enderlein (1914, 1921) based his genera on ven- ational characters and Heslop-Harrison sup- ported these arguments, adding further structural characters to support the separation of Psausia. The various forewing types may be characterised as follows: 1 M,,, reaching wing margin at apex of wing (Figs 64, 65) [Primitive condition for Carsidaridae + Homo- tomidae]) is. eesc eee ‘Labobrachia’ type -— M,,» reaching wing margin anterior to wing apex (Figs 63), 66=73))" 0 oe. se ee oe oa 2 2 M-+Custem entirely separated from R stem, M stem entirely separated from Rs, apex of claval suture adjacent or close to apex of Cu,,, radular areas diffuse (Figs 63, 66-70) ........ ‘Homotoma’ type — M-+Cu stem partly or entirely fused with R stem, M stem partly or entirely fused with Rs, apex of claval suture distant from apex of Cu,,, radular areas defined"(Figs 71=73))\7 sane aeeee et eee es 3 3 Mstem partly fused with Rs, M,,, entirely separate fromuRs) (Bigs) 715,72) orsseertoeicee ‘Psausia’ type — M stem almost or completely fused with Rs, M,,, completely fused with Rs (Fig. 73) ‘Metapsausia’ type Assuming the ‘Labobrachia’ type to be the primi- tive forewing condition, transformation series can be defined through the ‘Homotoma’ type to the Afrotropical species (Figs 63, 66-70) in one series, and to ‘Psausia’ types (Figs 71, 72) and ‘Metap- sausia’ types (Fig. 73) in another series. It is even possible to derive the condition found in the South American genus Synoza (Fig. 74) from the ‘Psausia’ type. One could reasonably argue that the ‘Psausia’ type was uniquely derived from the ‘Homotoma’ type, but the ‘Metapsausia’ type could have arisen several times from the “Psausia’ type. The supporting morphological characters used by Heslop-Harrison (1949) to separate Psausia from Homotoma are unlikely to be useful but the pronotum of those species with the ‘Psausia’ and ‘Metapsausia’ types of forewing, that we have examined, have anterodorsal and anterolateral projections. In those species with the ‘Labobrachia’ and ‘Homotoma’ types the pro- notum is a more simple, strap-like tergite. The male genitalia of all specimens examined show no significant differences that would support Enderlein’s generic concepts. We are, therefore, following Hodkinson (1983, 1986) in recognising a single genus, Homotoma. Thirty-one species are currently recognised, including four newly described below from the Afrotropical Region. All these species are listed FICUS-FEEDING PSYLLIDS below on a regional basis; a key is not provided as many of the descriptions are inadequate and we have been unable to obtain type material. Palaearctic Region Homotoma ficus (Linnaeus) ‘Les faux pucerons du figuier’ Reaumur, 1737: S51) ‘La Psylle du figuier’ Geoffroy, 1762: 484. Chermes ficus Linnaeus, 1758: 455; Fabricius, 1794: 223. | Psylla ficus (Linnaeus) Tigny, 1802: 165; Latreille, 1804: 379; Audinet-Serville, 1825: 229; Dufour, 1833: 232; Amyot & Serville, 1843: 593. Homotoma ficus (Linnaeus) Guérin-Méneville, 1844: 376; Flor, 1861: 413; Girard, 1876: 132; Bolivar & Chicote, 1879: 185; Ferrari, 1888: 76; Lambertie, 1901: 221; Dominique, 1902: 225; Ragusa, 1907: 237; Oshanin, 1907: 369; 1912: 128; Aulmann, 1913: 35; Horvath, 1918: 331; Boselli, 1929: 218; Bodenheimer & Theodor, 1929: 35: Silvestri, 1934: 382; Balachowski & Mesnil, 1935: 586; Haupt, 1935: 252; Heslop- Harrison, 1946: 37; 1949: 375; Schaefer, 1949: 56; Vondracek, 1951: 26; Tamanini, 1955: 11; 1966: 105; Ramirez Gomez, 1956: 66; Wagner & Franz, 1961: 170; Klimaszewski, 1961: 115; 1973: 230; Dobreanu & Manolache, 1962: 364; Loginova, 1964a: 472; Hodkinson & White, 1979: 77; White & Hodkinson, 1982: 45; Halperin et al., 1982: 34; Burckhardt, 1983: 64; Andrianova & Klimaszewski, 1983: 42; Yang & Li, 1984a: 201, 217. _ Anisostropha ficus (Linnaeus) Foerster, 1848: 92; | Frauenfeld, 1867: 801; Meyer-Diir, 1871: 403. ' Hostpcant. Ficus carica; ((Ficus) sect. Ficus). _ RECORDED DISTRIBUTION. Widespread in Mediter- ranean Basin, also from Albania, Austria, France, Iran, U.S. S. R. (Caucasus and Crimea), (England and North America, introduced). MATERIAL EXAMINED Numerous adults and larvae from various localities in the Mediterranean Basin, England (BMNH). | Homotoma viridis Klimaszewski | Homotoma viridis Klimaszewski, 1961: 114; 1973: 231; 1975: 18, 30, 37; Tamanini, 1966: 105; Log- inova, 1968: 326; Halperin et al., 1982: 35; Burckhardt, 1983: 64; 1988: 40; Andrianova & Klimaszewski, 1983: 43; Yang & Li, 1984a: 201, 217. Holotype 0’, ALBANIA (IZPAN) [not examined]. 159 HostTpLant. Ficus carica. RECORDED DISTRIBUTION. Bulgaria, Israel, Italy, Spain, Tunisia, Turkey, U.S.S.R. (Caucasus and Crimea), Yugoslavia. ComMENTsS. The morphological differentiation of this species from H. ficus has a weak basis. Fur- thermore, both have been collected at the same time and locality, from the same host individual, and there is no biological evidence that the species are distinct. Afrotropical Region The four new species described below are the first records of the genus in this region. They are dis- tinguished by their wing shape, venation and coloration and by the structure of the antennal flagellum. Homotoma angolensis sp. n. (Figs 22, 23, 39, 63, 99, 100) Description. Dark brown psyllids with a beetle- like appearance, 3.0-3.5 mm long. Head (Figs 22, 23) with vertex flat, anterior margin weakly incised by median suture; antennal flagellum 1.9— 2.0 (’) and 1.6-1.8 (Q) times longer than head width, laterally flattened and expanded (Fig. 39), Ist flagellomere almost 3 times as long as wide, 8th flagellomere with one long pointed seta and one very short and truncate seta apically; genae with weak, conical swellings ventrally; ultimate rostral segment 2.0-2.5 times longer than wide. Thorax weakly arched, pronotum without ante- rior projections; forewing (Fig. 63) coriaceous, obovate with rounded apex, 2.2—2.4 times longer than wide, veins densely clothed with long sinuous setae, M stem and branches entirely separated from R stem and branches, M stem short, , value 2.0-2.4, Custem about as long as M+ Cu stem, cu, value about 0.9, apex of claval suture adjacent to apex of Cu;,, radular areas diffuse; hindwing rela- tively long, 0.85 times as long as forewing, M unbranched, Cu branched apically; basal spine of hind tibia absent, apical spurs arranged 0 + 5; hind tibia with 2 apical spurs. C proctiger (Fig. 99) with lateral lobes of basal segment well-developed, anal tube of moderate length; aedeagus (Fig. 99) with basal segment weakly expanded in apical half; inner surface of paramere (Fig. 100) with a well-developed ante- romedial pointed tubercle and a diagonal ridge lying anteromedially to posteroapically. Q terminalia short, conical; proctiger 1.1 times longer than head width. 160 Measurements (5 0’, 4 2). Maximum width of head, CO 0.62-0.67, 9 0.70-0.74; length of anten- nal flagellum, oO” 1.20-1.36, 2 1.18-1.26; length of ultimate rostral segment, O 0.80-0.10, 2 0.10- 0.11; forewing length, 0” 2.44-2.52, 9 2.96-3.04; length of hind tibia, o 0.56-0.60, 2 0.60-0.64; length of 9 proctiger, 0.76—0.80. Larva unknown. Hosteiant. Adults collected from Ficus thon- ningit ((Urostigma) sect. Galoglychia subsect. Chlamydodorae); possibly also F. mutandifolia. Holotype 0’, Angola: Chianga, 21—24.111.1972, Ficus thonningii (Hollis) (BMNH); dry-mounted. Paratypes. Angola: 13 0’, 7 9, same data as holotype; 6 9, 7.x.1971; 10’, 14.1.1975, Ficus mutandifolia (van Harten); 11 0, 3 2, 7 mls W. Gabela, 16-18.iii.1972, general sweeping; 1 9, at light (Hollis). (BMNH); dry- and slide-mounted. Comments. This species is recognised by the dis- tinctive form, venation and chaetotaxy of the fore- wing. The obovate shape, and proximal branching of M stem and Cu stem of the wing are considered to be primitive features but the coriaceous nature of the membrane is unique within the genus. The structure of the antennal flagellum is highly derived in that it is short, strongly laterally flat- tened and with expanded flagellomeres. No obvious close relatives are known. Homotoma bamendae sp. n. (Figs 24, 38, 67, 101) DescripTIoN. (Only slide-mounted material available for study.) Antennal flagellum 1.73- 1.97 times longer than head width, flagellomeres laterally flattened and moderately expanded (Fig. 38), 1st flagellomere 4.8 times longer than wide, 8th flagellomere with one long pointed seta and one short truncate seta apically. Vertex flat, ante- rior margin very weakly incised by median suture; genae produced into long thin processes ventrally (Fig. 24); ultimate rostral segment very short, 1.75 times longer than wide. Thorax weakly arched, pronotum without ante- rior tubercles. Forewing obovate with subacute apex, 2.42.5 times longer than wide, membrane hyaline with pattern as in Fig. 67; veins bearing long straight setae, M stem and branches entirely separated from R stem and branches, M stem long, m, cell value about 1.0, Cu stem short, cu, cell value 1.6, apex of claval suture adjacent to apex of Cu,,, radular areas small and poorly defined; hindwing 0.65 times as long as forewing, M unbranched, Cu apparently unbranched; basal spine of hind tibia absent, apical spurs arranged 0 + 4; hind basitarsus with 2 apical spurs. D. HOLLIS & P. S. BROOMFIELD CO proctiger (Fig. 101) with moderately developed lateral lobes, anal tube elongate; aedeagus (Fig. 101) with basal segment expanded in apical half; inner surface of paramere with 2 anteromedial tubercles. Q terminalia short, conical, proctiger about as long as head width. Measurements (1 OC’, 1 Q). Maximum width of head, 0’ 0.60, 9 0.68; length of antennal flagellum, Co’ 1.04, 9 1.34; length of ultimate rostral segment, 0’ and 2 0.07; length of forewing, O' 2.48, 9 3.12; length of hind tibia, 0 0.52, 2 0.62; length of 2 proctiger, 0.71. Larva and hostplant unknown. Holotype CO’, Cameroon: Bamenda, 25-31.i. 1957, yellow trays (Eastop) (BMNH)); slide- mounted. Paratype. 1 Q, same locality as holotype, 21-24.1.1957 (BMNH); slide-mounted. ComMMENTS. The forewing venation of this species suggests a close relationship to H. ficus but the thin, elongate genal processes of H. bamendae appear to be unique in the genus. Homotoma chlamydodora sp. n. (Figs 68, 69, 103, 104) DESCRIPTION. Overall length up to 5.0 mm. Vertex concave, anterior margin deeply incised by medial suture. Antennal flagellum 2.52-3.23 (C’) and 2.35-2.83 (2) times longer than head width, cir- cular in cross-section, 1st flagellomere 5.0—7.0 times longer than wide, 8th flagellomere with one long pointed seta and one short truncate seta apically; genae slightly swollen ventrally; ultimate rostral segment short, 1.5—2.0 times longer than wide. Thorax weakly arched, pronotum broadly visi- ble from above and without anterior tubercles. Forewing elongate-oval, with subacute apex, membrane hyaline and with a dimorphic pattern as in Figs 68, 69, 2.75—3.15 times longer than wide; veins bearing long straight setae, M stem and branches entirely separate from R stem and branches, M stem long, mm, cell value 0.9, Cu stem about as long as M+Cu stem, cu, cell elongate with a value of about 2.0, apex of claval suture adjacent to apex of Cu,,, radular areas diffuse; hindwing 0.75 times as long as forewing, M unbranched, Cu branching apically; hind tibia without a basal spine, apical spurs arranged 0 + 5; hind basitarsus with a single apical spur. CO proctiger (Fig. 103) with moderately developed lateral lobes and short anal tube; aedeagus as in Fig. 103; paramere (Fig. 104) thumb-shaped, inner surface with a single tuber- cle anteriorly in apical third and a posteroapical tubercle. FICUS-FEEDING PSYLLIDS @ terminalia short, conical; proctiger 0.9-1.1 times as long as head width. Measurements (10 0’, 9 2). Maximum width of head, CO 0.68-0.88, 2 0.74-0.88; length of anten- nal flagellum, CO 1.88-2.52, 2 1.74-2.32; length of ultimate rostral segment, OC’ 0.06-0.09, 2 0.07— 0.08; length of forewing, O 3.0-3.62, 2 3.08- 3.88; length of hind tibia, CO’ 0.58-0.82, 2 0.58- 0.84; length of 2 proctiger, 0.68-0.92. Larva unknown. Hostptants. Adults collected from Ficus natalen- sis and F. thonningi; ((Urostigma) sect. Galoglychia subsect. Chlamydodorae). Holotype OC’, Tanzania: Arusha NP, Ngurdoto crater rim, c. 5000’, 8.vi.1974, Ficus natalensis (Hollis) (BMNH); dry-mounted. Paratypes. Tanzania: 3 CO’, 3 2, same data as holotype. South Africa: 2 0’, 3 2, Pondoland, Port St John, ix—xii.1923 (Turner); 80°, 129, Natal, Umtentweni, 14-17.x.1969, Ficus petersii (Capener). Kenya: 1 oO, 1 9, Muguga, vii- viii.1954, trapped (Eastop); 1 CO, 19.xi.1969 (Brown); 10,3 9, Limuru, iv.1955 (Thomas); 1 | O, Kakamega, 1500m, 4.ii.1977 (Deharveng). | Uganda: 6 0’, 2 9, Kampala, 17.iii.1923, on Ficus leaves (Hargreaves). Angola: 1 Q, Chianga, 7.x.1971, Ficus thonningii (van Harten). Burundi: 20, 2 9, Gitega, 20.x.1980 (Pointel); 4 CO, Kisozi, 6—20.iii.1981; 8 9, 5-17.iv.1981; 1 oC, 1-15.v.1981; 2 oO’, 8-21.viii.1981; 10 Oo, 9 Q, 22.iii—4.ix.1981; 4 oO’, 3 9, 6-18.ix.1981; 4 Co, 39, 10.ix.—2.x.1981; 10 oO, 18 9, 3-16.x.1981; 4 oO, 3 Q, 17-30.x.1981 (Aubrique). Cameroon: 10 Oo, 12 2, Bamenda, 24.i.—6.ii.1957, yellow trays (Eastop). Nigeria: 1 O’, K. State, 20 mls W. Lokoja, 21.11.1970 (Medler). (BMNH; MNHN; | NCI; MHN); dry- and slide-mounted, stored in 90% ethanol. | Comments. H. chlamydodora may be recognised by the venation and pattern of the forewing and | the single apical spur on the hind basitarsus. Prob- ably it is most closely related to H. bamendae but has a more derived condition of the cubital vein of | the forewing and the antennae have primitive, uncompressed flagellomeres. Homotoma eastopi sp. n. (Figs 40, 70, 102) Description. (Only slide-mounted material available for study.) Antennal flagellum 2.67 times longer than head width, flagellomeres flat and strongly expanded (Fig. 40), 1st flagellomere not more than twice as long as wide, 8th flagellomere with one long pointed seta and one 161 very short truncate seta apically; vertex deeply divided by median suture; genae with very small, conical swellings ventrally; ultimate rostral seg- ment short, 2.0 times longer than wide. Thorax weakly arched, pronotum broadly visi- ble from above and without anterior projections. Forewing narrow-elongate, with acute apex, 3.1— 3.7 times longer than wide, membrane hyaline with pattern as in Fig. 70; veins bearing long straight setae, M stem and branches entirely sepa- rate from R stem and branches, 7, cell value 0.9, Cu stem about as long as M+Cu stem but 3-4 times longer than Cu;,, cu; cell very elongate with a value of about 15.0, apex of claval suture close but not immediately adjacent to apex of Cu;,, radular areas narrow-elongate and _ poorly defined; hindwing 0.75 times as long as forewing, M unbranched, Cu branching distally; basal spine of hind tibiia absent, apical spurs arranged 0 + 6; hind basitarsus with 2 apical spurs. CO proctiger (Fig. 102) with lobes developed laterodorsally, anal tube long; aedeagus as in Fig. 102; paramere of similar profile to that of H. angolensis [inner surface not visible in holotype]. 2 terminalia conical, proctiger 1.27 times longer than head width. Measurements (1 0’, 1 2). Maximum width of head, CO 0.64, Q 0.66; length of antennal flagellum, oO 1.76; length of ultimate rostral seg- ment, O' 0.09, 9 0.10; length of forewing, O 3.1, ? 3.72; length of hind tibia, Ch and 9, 0.74; length of 2 proctiger, 0.84. Larva and hostplant unknown. Holotype ©’, Cameroon: Bamenda, 5000’, 25-31.1.1957, yellow pan trap (Eastop) (BMNH); slide-mounted. Paratype. 1 9, same locality as holotype, 20-24.1.1957 (BMNH); slide-mounted. ComMENTS. This species is differentiated from others in the genus by the shape, pattern and venation of the forewing, and the structure of the antenna and the male proctiger. It is probably the sister-species of H. chlamydodora. Oriental Region Homotoma altissimae (Yang & Li) Caenohomotoma altissimae Yang & Li, 1984a: 206, 217. Holotype 0’, CHINA: on Ficus alti- ssima (BAUIC) [not examined]. Homotoma altissimae (Yang & Li) Hodkinson, 1986: 312. HostpLant. Ficus altissima; ((Urostigma) sect. Conosycea). 162 RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma annesleae (Yang & Li) Caenohomotoma annesleae Yang & Li, 1984a: 205, 217. Holotype 0’, CHINA: on Anneslea fragrans (BAUIC) [not examined]. Homotoma annesleae (Yang & Li) Hodkinson, 1986: 313. HostTpLant. Two adults recorded from Anneslea fragrans (Theaceae). These specimens were prob- ably vagrants. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma bambusae (Yang & Li) Caenohomotoma (Heterhomotoma) bambusae Yang & Li, 1981: 82, 85, 86; 1984a: 210. Holo- type 0’, CHINA: on Bambusa sp. (BAUIC) [not examined]. Caenohomotoma bambusae Yang & Li, 1984a: 216; 218: Homotoma bambusae (Yang & Li) Hodkinson, 1986: 313. HostTpLant. Two adults collected from Bambusa sp. (Graminae). These specimens were probably vagrants. RECORDED DISTRIBUTION. China (Hainan). No material examined. Homotoma benjaminae (Yang & L1) Caenohomotoma benjaminae Yang & Li, 1984a: 211, 217. Holotype &, CHINA: on Ficus ben- jamina (BAUIC) [not examined]. Homotoma benjaminae (Yang & Li) Hodkinson, 1986: 313. HosTpLant. Three adults collected from Ficus benjamina; ((Urostigma) sect. Conosycea). RECORDED DISTRIBUTION. China (Lincang Prov.). No material examined. Homotoma boheae Yu (Fig. 71) Homotoma boheae Yu, 1957: 45; Miyatake, 1975: 20; Hodkinson, 1986: 313. Holotype CO’, CHINA (?BPBM) [not examined]. Caenohomotoma (Caenohomotoma) boheae (Yu) Yang & Li, 1981: 78, 85; 1984a: 218. HostTpLant. Unknown. D. HOLLIS & P. S. BROOMFIELD RECORDED DISTRIBUTION. China (Fukien). MATERIAL EXAMINED Malaya, | Oo’, 1 2 (BMNH). Homotoma chuanana (Yang & Li) Caenohomotoma (Psausia) chuanana Yang & Li, 1981: 81, 86. Holotype 9, CHINA (BAUIC) [not examined]. Caenohomotoma chuanana Yang & Li; Yang & Li, 1984a: 216, 218. Homotoma chuanana (Yang & Li) Hodkinson, 1986: 313. HostTeLant. Unknown. RECORDED DISTRIBUTION. China (Sichuan). No material examined. Homotoma distincta Crawford Homotoma distincta Crawford, 1912: 433; 1919: 162; Ramakrishna Ayyar, 1924: 622; Boselli, 1929: 219; Miyatake, 1975: 21; Hodkinson, 1986: 313. Holotype 9, INDIA (BMNH) [examined]. Psausia distincta (Crawford) Enderlein, 1921: 120; Heslop-Harrison, 1949: 379; Mathur, 1975: 156; Loginova, 1982: 39; Yang & Li, 1984a: 202, 217. Caenohomotoma (Harrisonella) (Crawford) Yang & Li, 1981: 78, 85. Hostpiant. Ficus religiosa (Heslop-Harrison, 1949); ((Urostigma) sect. Urostigma). distincta RECORDED DISTRIBUTION. India (Bihar, U. P.). MATERIAL EXAMINED India (Bihar, U. P.): holotype 2, 1 ho (BMNH). Homotoma galbvittata (Yang & Li) Caenohomotoma galbvittata Yang & Li, 1984a: 203, 217. Holotype o&’, CHINA: on Ficus sp. (BAUIC) [not examined]. Homotoma galbvittata (Yang & Li) Hodkinson, 1986: 313. HostTpLant. Ficus sp.. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma indica (Mathur) Psausia indica Mathur, 1975: 158. Holotype OC’, INDIA: on ‘Ficus infectoria’ (FRI) [not examined]. FICUS-FEEDING PSYLLIDS Caenohomotoma indica (Mathur) Yang & Li, 1984a: 218. Homotoma indica (Mathur) Hodkinson, 1986: 318 HostTp.Lants. There is some confusion in the origi- nal description concerning the identity of the host. Under hostplant data Mathur states ‘Ficus mac- rocarpa (=F. retusa) and F. lucescens (=F. infectoria)’. Under type data only Ficus infectoria is given, and under biological notes Mathur states ‘Both adults and nymphs are commonly found on young shoots of F. macrocarpa at ... Dehra Dun’. Ficus macrocarpa (=F. retusa) is probably a misspelling of F. micocarpa (see Corner, 1965: 22), although it may refer to F. laevis var. mac- rocarpa (Corner, 1965: 53). F. infectoria may refer to either F. tsjahela or F. virens (Corner, 1965: 7 and 9 respectively). RECORDED DISTRIBUTION. India (UP). No material examined. Homotoma lahui (Yang & Li) Caenohomotoma lahui Yang & Li, 1984a: 212, 217. Holotype 9, CHINA (BAUIC) [not examined]. Homotoma lahui (Yang & Li) Hodkinson, 1986: 813: HosteLant. Unknown. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma maculata Yang Psausia maculata Yang, 1984: 173. Holotype ©’, TAIWAN: on Ficus beechyana (NCHU) [not examined]. Homotoma maculata (Yang) Hodkinson, 1986: al3. Hoste ant. Ficus erecta var. beechyana; ((Ficus) sect. Ficus). RECORDED DISTRIBUTION. Taiwan. No material examined. Homotoma mangiferae (Yang & Li) Caenohomotoma mangiferae Yang & Li, 1984a: 209, 217. Holotype &', CHINA: on Mangifera indica (BAUIC) [not examined]. Homotoma mangiferae (Yang & Li) Hodkinson, 1986: 314. HostpLantT. The single known adult, recorded from Mangifera indica, was probably a vagrant. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma pyriformiscola (Yang & Li) Caenohomotoma pyriformiscola Yang & Li, 1984a: 207, 217. Holotype &, CHINA: on Ficus pyriformis (BAUIC) [not examined]. Homotoma pyriformiscola (Yang & Li) Hodkin- son, 1986: 314. HOSTPLANT. Ficus). Ficus pyriformis; ((Ficus) sect. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma radiata Kuwayama (Fig. 72) Homotoma radiatum Kuwayama, 1908: 181; Crawford, 1911: 491; 1912: 433; 1915: 262; 1919: 162; Aulmann, 1913: 36; Kuwayama. 1931: 124; Sasaki, 1954: 31; Klimaszewski, 1973: 231; Hill et al., 1982: 174. Syntypes, Co and 9, TAIWAN (EIHUV) [not examined]. Psausia radiata (Kuwayama) Enderlein, 1914: 232; Heslop-Harrison, 1949: 375; Yang, 1984: 169; Fang & Yang, 1987: 147. Psausia (Homotoma) — radiata Kuwayama, 1922: 368. Homotoma radiata Kuwayama; Boselli, 1929: 219; Miyatake, 1965a: 173; 1975: 20; 1981: 56; Yu, 1957: 44; Hodkinson, 1983: 349; 1986: 313. Caenohomotoma radiata (Kuwayama) Yang & Li, 1981: 79; 1984a: 216, 218. HostpLants. Recorded from Ficus erecta, F. caulocarpa, F. superba var. japonica and Ficus sp. (Kuwayama) RECORDED DISTRIBUTION. Nepal, Taiwan, Japan and Hong Kong. MATERIAL EXAMINED. Hong Kong: adults on Ficus superba var. japonica (BMNH). Homotoma ruiliana (Yang & Li) Caenohomotoma ruiliana Yang & Li, 1984a: 214, 217. Holotype co, CHINA: on Ficus sp. (BAUIC) [not examined]. Homotoma ruiliana (Yang & Li) Hodkinson, 1986: 314. Hostevant. A single male collected from Ficus sp. RECORDED DISTRIBUTION. China (Yunnan). No material examined. 164 Homotoma shuana (Yang & Li) Caenohomotoma (Heterohomotoma) shuana Yang & Li, 1981: 79, 85, 86; 1984a: 210. Holo- type O', CHINA (BAUIC) [not examined]. Caenohomotoma shuana Yang & Li; Yang & Li, 1984a: 216, 218. Homotoma shuana (Yang & Li) Hodkinson, 1986: 314. HostTeLant. Unknown. RECORDED DISTRIBUTION. China (Sichuan). No material examined. Homotoma spiraea (Yang & Li) Caenohomotoma (Caenohomotoma) _ spiraea Yang & Li, 1981: 79, 85, 86; 1984a: 216, 218. Holotype o&’, CHINA: on ‘Spirae’ sp. (BAUIC) [not examined]. Homotoma spiraca (Yang & Li) Hodkinson, 1986: 314. [Misspelling. ] HostTpPLant. Described from 12 adults collected on ‘Spirae’ sp. |? Spiraea sp.]. RECORDED DISTRIBUTION. China (Zhejiang). No material examined. Homotoma unifasciata Yu Homotoma unifasciata Yu, 1957: 45; Miyatake, 1975: 21; Hodkinson, 1986: 314. Holotype C’, CHINA (? BPBM) [not examined]. Caenohomotoma (Caenohomotoma) unifasciata (Yu) Yang & Li, 1981: 78, 85. Caenohomotoma unifasciata (Yu); Yang & Li, 1984a: 216, 218. HostTeLant. Unknown. RECORDED DISTRIBUTION. China (Fukien); Japan. No material examined. Homotoma wulinensis (Yang) Psausia wulinensis Yang, 1984: 170. Holotype C’, TAIWAN: on Ficus sarmentosa (NCHU) [not examined]. Homotoma wulinensis (Yang) Hodkinson, 1986: 314. HOsTpPLANnT. Rhizocladus). RECORDED DISTRIBUTION. Taiwan. No material examined. Ficus sarmentosa; ((Ficus) sect. D. HOLLIS & P. S. BROOMFIELD Homotoma xishuangana (Yang & Li) Caenohomotoma xishuangana Yang & Li, 1984a: 213, 217. Holotype o’, CHINA: on Ficus sp. (BAUIC) [not examined]. Homotoma xishuangana (Yang & Li) Hodkinson, 1986: 314. HostTpLant. Described from four adults collected on Ficus sp. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Homotoma yunnanica (Yang & Li) Caenohomotoma yunnanica Yang & Li, 1984a: 210, 217. Holotype &’, CHINA: on Ficus sp. (BAUIC) [not examined]. Homotoma yunnanica (Yang & Li) Hodkinson, 1986: 314. HostTpLant. Described from nine adults collected on Ficus sp. RECORDED DISTRIBUTION. China (Yunnan). No material examined. Australo-oriental Region Homotoma bakeri Crawford Fig. 73) Homotoma bakeri Crawford, 1915: 263; 1919: 162; Boselli, 1929: 219; Heslop- Harrison, 1949: 376; Miyatake, 1975: 20; Braza & Calilung, 1981: 353; Hodkinson, 1983: 349. Holotype 9, PHILIPPINES (USNM) [examined]. Metapsausia bakeri (Crawford) Enderlein, 1921: 120. Caenohomotoma (Metapsausia) (Crawford) Yang & Li, 1981: 78, 85. Caenohomotoma bakeri (Crawford) Yang & Li, 1984a: 218. HostpLant. Ficus benjamina var. nuda (=F. com- osa; recorded as F. carnosa by Braza & Calilung); ((Urostigma) sect. Conosycea). bakeri RECORDED DISTRIBUTION. Philippines (Luzon and Mindanao); West Malaysia (Penang). MATERIAL EXAMINED Philippines: &’, Q and larvae, Mt Makiling. West Malaysia: 1 2, Pahang (BMNH). Homotoma bilineata Crawford Homotoma bilineata Crawford, 1917: 164; 1919: 162; Boselli, 1929: 219; Miyatake, 1975: 20. FICUS-FEEDING PSYLLIDS Holotype oO, PHILIPPINES [examined]. Metapsausia bilineata (Crawford) Enderlein, 1921: 120. Homotoma biliniata Crawford; Heslop-Harrison, 1949: 376. [Misspelling. ] Caenohomotoma (Metapsausia) (Crawford) Yang & Li, 1981: 78, 85. Caenohomotoma bilineata (Crawford) Yang & Li, 1984a: 218. Hostecant. Ficus sp. (1 2, BMNH). RECORDED DISTRIBUTION. Philippines (Luzon); Thailand. MATERIAL EXAMINED Philippines: 1 C& (holotype), Luzon (USNM); 1 Q, Palawan (BMNH). ComMENtS. Hodkinson’s (1983, 1986) records of H. bilineata from Sarawak refer to a closely related species, of which there are further speci- mens in BMNH from Sulawesi and Seram. (USNM) bilineata Homotoma gressitti Miyatake Homotoma gressitti Miyatake, 1975: 17; Hodkin- son, 1983: 349. Holotype 9, PAPUA NEW GUINEA (BPBM) [not examined]. Caenohomotoma (Austrohomotoma) (Miyatake) Yang & Li, 1981: 78, 85. Caenohomotoma gressitti (Miyatake) Yang & Li, 1984a: 218. HostTpLant. Unknown. gressitti RECORDED DISTRIBUTION. Papua New Guinea. MATERIAL EXAMINED A series of adults from Papua New Guinea (Mondo), in BMNH, differ from the original description slightly in forewing venation and pattern. Homotoma pacifica Crawford (Fig. 64) Homotoma pacifica Crawford, 1915: 262; 1919: 162; Boselli, 1929: 219; Heslop- Harrison, 1949: 378; Miyatake, 1975: 21; Yang & Li, 1981: 78, 85; Hodkinson, 1983: 349. Holotype co’, PHI- LIPPINES (USNM) [examined]. Labobrachia_ pacifica (Crawford) Enderlein, 1921: 19; Heslop-Harrison, 1949: 376; Yang & ix, 1984a: 201, 217. HosteLant. Unknown. RECORDED DISTRIBUTION. Philippines (Luzon). 165 MATERIAL EXAMINED Philippines: 1 CO’ (holotype) (USNM). Indonesia: 1 2, Sulawesi Utara (BMNH). Comments. A single 2 of a closely related but undescribed species (Fig. 65) is in BMNH, also from Sulawesi Utara. SYNOZINI Bekker-Migdisova Synozini Bekker-Migdisova, 1973: 102. Type genus: Synoza Enderlein. Synoziini White & Hodkinson, [Misspelling. | M9853) 162: DiaGnosis. In forewing M+Cu stem absent, M stem completely fused along its basal two-thirds with Rs, radular area absent from cell mm,; hind tibia with an almost complete ring of apical spurs; CO proctiger unipartite; basal segment of aedeagus not expanded, apical segment with a pair of sub- apical spiniform processes ventrally. A single, New World genus, Synoza, is included here. It comprises three species from Central and South America and recorded hostplants are Ficus spp. SYNOZA Enderlein Synoza Enderlein, 1918: 479; Laing, 1923: 697; Ferris, 1928: 109; Caldwell, 1941: 419; Tuthill, 1950: 58: Bekker-Migdisova, 1973: 102; Hodkinson & White, 1981: 509; White & Hodkinson, 1985: 272; Brown & Hodkinson, 1988: 179. Type species: Synoza cornutiventris Enderlein, by original designation and monotypy. DescripTION. Medium-sized psyllids, up to 5.0 mm. Integument of head and thorax sparsely cov- ered with long setae. Head (Figs 25, 26), from above, about as wide as mesoscutum; vertex con- cave on either side of median suture, anterior and lateral margins rounded, occipital margin angular medially but rounded laterally, anterior margin deeply incised by median suture, lateral ocelli on raised tubercles, anterolateral tubercles absent; antennal bases enlarged, giving head a cleft appearance, flagellum about 4 times longer than head width, flagellomeres cylindrical, a single subapical rhinarium present on flagellomeres 2, 4, 6 and 7; genae not swollen ventrally; ultimate rostral segment short, less than 2.5 times longer than wide. Thorax, in profile, strongly arched, pronotum very narrowly visible from above and descending 166 sharply behind occiput. Forewing (Fig. 74) obo- vate, with rounded apex, about 2.5 times longer than wide, veins bearing long setae, C+Sc not thickened, costal break indicated by weakening of chitin in break area, basal two-thirds of M stem completely fused with Rs, apex of M,,, reaching wing margin anterior to wing apex, M+ Cu absent, m, cell value about 1.0, cu, cell value about 1.0, radular areas sharply defined but absent from cell mM, apex of claval suture distant from apex of Cu,,; basal spine of hind tibia absent, apical spur ring almost complete; hind basitarsus with 2 apical spurs. CO proctiger unipartite, with well-developed lat- eral lobes that do not bear inner apical lobes; basal segment of aedeagus not expanded in apical half, apical lobes of aedeagus with a pair of spiniform lobes ventrally (Fig. 105). Q terminalia conical or elongate conical; cir- cum-anal pore ring simple. Larva. Body longer than wide; antenna long, filiform, not subdivided; wing pads small, humeral lobes not developed; body surface mem- braneous with thoracic sclerites not defined, setae mainly simple but abdominal segments bearing small groups of 1-3 pointed sectasetae sub- marginally on dorsal surface, dorsal caudal plate (Fig. 113) small, bearing groups of pointed sec- tasetae; anus ventral (Fig. 113), with a large, medially constricted circum-anal pore ring; tarsal arolium triangular, not petiolate. Comments. Laing (1923) and Ferris (1928) placed Synoza in the Carsidaridae sensu Crawford (1919); Bekker-Migdisova (1973: fig. 2) con- sidered the genus to be the sister-group of the rest of the homotomids, and White & Hodkinson (1985) regarded it as the sister-group of Homotoma. The three included species are discussed and differentiated by Brown & Hodkinson (1988). Synoza cornutiventris Enderlein (Figs 25, 26, 74, 105) Synoza cornutiventris Enderlein, 1918: 480; Laing, 1923: 697; Crawford, 1925b: 58; Ferris, 1928: 109; Costa Lima, 1942: 105; Hodkinson & White, 1981: 509; Brown & Hodkinson, 1988: 180. Holotype 2, PERU (not traced). HostpL_ant. Ficus sp. (Brown & Hodkinson, 1988). RECORDED DISTRIBUTION. Colombia, Panama and Peru. D. HOLLIS & P. S. BROOMFIELD MATERIAL EXAMINED Colombia: 1 C’, 1 Q, on Ficus sp.; Panama: 3 9 (BMNH). Synoza floccosa Ferris (Fig. 113) Synoza floccosa Ferris, 1928: 109; Klyver, 1930: 175 (as asynonym of S. pulchra Laing); Tuthill, 1950: 59; Hodkinson & White, 1981: 509; Brown & Hodkinson, 1988: 180. Syntypes o, 2, larvae, MEXICO: on Ficus sp. (SUNHM) [examined]. Synoza pulchra Laing; Caldwell, 1941: 421, in part. HostTpLant. Ficus sp. RECORDED DISTRIBUTION. Mexico. MATERIAL EXAMINED Mexico: adults and larvae on Ficus sp. (syntype series, SUNHM). 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Notes on the Psyllidae (Homoptera) from Fukien and Taiwan, China. I. Memoirs of the College of Agriculture, National Taiwan University 4: 43-54. 170 D. HOLLIS & P. S. BROOMFIELD Figs 2-11 Dynopsyllinae, head structure. 2, 3, Diceraopsylla brunettii; 2, dorsal view; 3, lateral view. 4, 5, Dynop- sylla cornuta; 4, dorsal view; 5, lateral view. 6, 7, Austrodynopsylla encala; 6, dorsal view; 7, lateral view. 8, 9, Triozamia lamborni; 8, dorsal view; 9, lateral view. 10, 11, Afrodynopsylla gigantea; 10, dorsal view; 11, lateral view. Scale line: 0.5 mm. Setosity not shown in Figs 2, 3, 6-9. FICUS-FEEDING PSYLLIDS 171 12 Figs 12-26 Macrohomotominae and Homotominae, head structure. 12, 13, Mycopsylla gardenensis; 12, dorsal view; 13, lateral view. 14, 15, M. tuberculata; 14, dorsal view; 15, lateral view. 16, 17, M. obliqua; 16, dorsal view; 17, lateral view. 18, 19, Macrohomotoma gladiata; 18, dorsal view; 19, lateral view. 20, 21, Pseudoeriopsylla laingi; 20, dorsal view; 21, lateral view. 22, 23, Homotoma angolensis; 22, dorsal view; 23, lateral view. 24, H. bamendae, dorsal view. | 25, 26, Synoza cornutiventris; 25, dorsal view; 26, lateral view. Scale lines: 0.5 mm. Setosity not shown in Figs 12-21. Fig. 24 drawn from slide-mounted specimen. 172 D. HOLLIS & P. S. BROOMFIELD 4 A TY J ope ie TAIT SE — We | = | Figs 27-40 Homotomidae, antennal structure. 27, Austrodynopsylla encala, \st and 2nd flagellomeres. 28, Tri- ozamia lamborni, \st and 2nd flagellomeres. 29, Afrodynopsylla gigantea, \st flagellomere. 30, Mycopsylla kina, \st flagellomere. 31, M. obliqua, flagellum. 32, Pseudoeriopsylla nyasae, \st and 2nd flagellomeres. 33, P. /aingi, 1st and 2nd flagellomeres. 34, P. medleri, 1st and 2nd flagellomeres. 35, P. carvalhoi, \st and flagellomeres. 36, P. kenyae, \st and 2nd flagellomeres. 37, P. etiennei, 1st and 2nd flagellomeres. 38, Homotoma bamendae, antenna. 39, H. angolensis, antenna. 40, H. eastopi, antenna. Scale lines: Figs 27-30, 0.2 mm; Figs 31, 38-40, 0.5 mm; Figs 32-37, 0.1 mm. FICUS-FEEDING PSYLLIDS 173 Figs 41-46 Homotomidae, thoracic, leg and abdominal structure. 41, Dynopsylla cornuta, thorax, lateral view. 42, Afrodynopsylla gigantea, thorax, lateral view. 43, Austrodynopsylla encala, apex of hind tibia. 44, Triozamia lamborni, apex of hind tibia. 45, 46, Diceraopsylla brunettii; 45, apex of hind tibia; 46, abdominal wax-producing glands. Scale lines: Figs 41, 42, 0.5 mm; Figs 43— 46, 0.1 mm. Figs 41, 42, drawn from dry-mounted specimens. 174 D. HOLLIS & P. S. BROOMFIELD Figs 47-56 Dynopsyllinae, fore- and hindwings. 47, 48, Diceraopsylla brunettii; 47, forewing; 48, hindwing. 49, 50, Dynopsylla pinnativena; 49, forewing; 50, hindwing. 51,52, Austrodynopsylla encala; 51, forewing; 52, hindwing. 53, 54, Triozamia lamborni; 53, forewing; 54, hindwing. 55, 56, Afrodynopsylla gigantea; 55, forewing; 56, hindwing. Scale lines: 0.5 mm. FICUS-FEEDING PSYLLIDS Ws Figs 57-74 Macrohomotominae and Homotominae, forewings. 57, Mycopsylla gardenensis. 58, M. obliqua. 59, M. propinqua. 60, Macrohomotoma gladiata. 61, Pseudoeriopsylla laingi. 62, P. etiennei. 63, Homotoma angolensis. 64, H. pacifica. 65, Homotoma sp., near pacifica. 66, H. ficus. 67, H. bamendae. 68, 69, H. chlamydodora, 68, specimen from Burundi; 69, specimen from Tanzania. 70, H. eastopi. 71, H. boheae. 72, H. radiata. 73, H. bakeri. 74, Synoza cornutiventris. Scale lines: 0.5 mm. Setosity not shown in Figs 57-62. 176 D. HOLLIS & P. S. BROOMFIELD Figs 75-83 Dynopsyllinae, male genitalia. 75, Diceraopsylla brunettii, external genitalia, lateral view. 76, 77, Dynopsylla pinnativena; 76, paramere; 77, apical segment of aedeagus, lateral view. 78, 79, Austrodynopsylla encala; 78, paramere; 79, apical segment of aedeagus, lateral view. 80, 81, Triozamia lamborni; 80, paramere; 81, apical segments of aedeagus, lateral view. 82, 83, Afrodynopsylla gigantea; 82, paramere; 83, apical segments of aedeagus, lateral view. Scale lines: 0.1 mm. FICUS-FEEDING PSYLLIDS 1 9/7/ | Figs 84-91 Edenini, male and female genitalia. 84, Mycopsylla kina, male external genitalia, lateral view. 85, 86, | M. tuberculata; 85, male external genitalia, lateral view; 86, female external genitalia, lateral view. 87-89, M. obliqua; | 87, male proctiger and aedeagus, lateral view; 88, paramere; 89, female external genitalia, lateral view. 90, 91, M. | propinqua; 90, male proctiger and aedeagus, lateral view; 91, paramere. Scale lines: Figs 86, 89, 0.2 mm; remainder 0.1 mm. 178 D. HOLLIS & P. S. BROOMFIELD x var x See : Aly, - FW sa 95 Y! ih Qa SA ESS JAM ZEW Vlad HO. nS a WN \\ So eg AN\Y \ { x : PX | | / Sa | 7 Ze =e SS Figs 92-98 Pseudoeriopsylla spp., male genitalia. 92, P. nyasae, paramere. 93, 94, P. laingi; 93, paramere; 94, apical segment of aedeagus, lateral view. 95, P. medleri, paramere. 96, P. carvalhoi, paramere. 97, P. kenyae, paramere. 98, P. etiennei, paramere. Scale lines: 0.1 mm. FICUS-FEEDING PSYLLIDS 179 _ Figs 99-105 Homotominae, male genitalia. 99, 100, Homotoma angolensis; 99, proctiger and aedeagus, lateral view; | 100, paramere. 101, H. bamendae, proctiger and aedeagus, lateral view. 102, H. eastopi, proctiger and aedeagus, lateral view. 103, 104, H. chlamydodora; 103, proctiger and aedeagus, lateral view; 104, paramere. 105, Synoza cornutiventris, external genitalia, lateral view. Scale lines: 0.1 mm. 180 D. HOLLIS & P. S. BROOMFIELD UW 7, tyes S IW NS aT \! TAMAS WrXt ! TI NW \ \ N /\! Vly 1 \ 4 \ VAAN Figs 106-113 Homotomidae, Sth instar larvae caudal plates. 106, Mycopsylla kina. 107, M. obliqua. 108, Mac- rohomotoma gladiata. 109, Pseudoeriopsylla nyasae, specimen from South Africa. 110, P. laingi, specimen from Senegal. 111, P. kenyae. 112, P. etiennei. 113, Synoza floccosa. d— dorsal view, v — ventral view. Scale lines: 0.2 mm. FICUS-FEEDING PSYLLIDS 181 INDEX TO HOSTPLANTS Americana (sect.) 138 Anneslea fragrans 162 Antiaris 132, 134, 135, 139 toxicaria welwitschii 133, 138, 142, 146 var. africana 133, 146 var. usambarensis 133, 146 var. welwitschii 133, 146 Apocynaceae 132 Artocarpus 133, 134 Auriculatae (series) 137 Bambusa sp. 162 Benjamina (subsect.) 137 Benjamineae (series) 136 Callophylleae (series) 136 Caricae (series) 137 Caulobotryae (series) 136 Caulocarpae (subsect.) 136, 156 Chlamydodorae (subsect.) 136, 154, 155, 160, 161 Chlorophora 134 Clausena anisata 156 Clusiaceae 133 Congestae (subser.) 137 Connariaceae 133 Conosycea (sect.) 135, 136, 138, 139, 147, 148, 151, 157, 161, 162, 164 Conosycea (subsect.) 136 Convolvulaceae 132 Crassirameae (subser.) 136 Drupaceae (series) 136 Ebenaceae 133 Exasperatae (series) 137 Ficus 131, 132, 133, 134, 135, 136, 138 altissima 136, 161 auriculata 133, 137 benghalensis 136, 152 benjamina 136, 153, 162 var. nuda 164 brachypoda 136, 156 capensis 137 carica 133, 137, 159 carnosa 164 caulocarpa 163 clementis 136, 153 comosa 164 crassiramea 136, 153 elastica 135, 136, 142 erecta 137, 163 var. beechyana 163 exasperata 137 fistulosa 137 foveolata 137 gibbosa 137 gnaphalocarpa 137 gul 137 hispida 137 hookeri 133 hookeriana 133, 136 infectoria 136, 162, 163 kirkii 154 Invalid names are in italics laevis var. macrocarpa 163 lucescens 163 macrocarpa 133, 163 macrophylla 136, 149 microcarpa 133, 136, 148, 149, 152, 153, 163 microphylla 153 minahassae 137 mollis 136, 148, 149 mutandifolia 160 natalensis 136, 154, 155, 161 nervosa 133, 137, 143, 144 nota 137 obliqua 136, 151 odorata 137 ovata 136, 156 petersit 154, 161 pumila 137 pyriformis 137, 163 racemosa 137 religiosa 136, 148, 149, 162 repandifolia 137 retusa 136, 152, 153, 163 roxburghii 137 rubiginosa 136, 149 rumphii 133, 136 sarmentosa 137, 164 scassellatii 154 sp. 133, 139, 142, 145, 149, 152, 153, 154, 155, 156, 162, 163, 164, 165, 166 spp. 133, 137, 138, 139, 153, 165 stipulata 137 superba 136 var. japonica 163 sur 137 sycomorus 137 thonningii 136, 154, 155, 160, 161 tinctoria 137 tomentosa 136 tsjahela 136, 148, 149, 163 ulmifolia 137 variegata 137 virens 136, 148, 149, 163 Ficus (sect.) 135, 137, 138, 139, 157, 159, 163 Ficus (subgen.) 133, 135, 137, 159, 163, 164 Ficus (subsect.) 137 Flacourtaceae 132 Galoglychia (sect.) 135, 136, 138, 139, 147, 154, 156, 157, 160, 161 Hispidae (subser.) 137 Indicae (subser.) 136 Lauraceae 133 Leucogyne (sect.) 136, 138 Malvaceae 133 Malvales 133, 139 Malvanthera (sect.) 135, 136, 138, 139, 147, 148, 150 Malvanthereae (series) 136 Malvanthereae (subser.) 136 Mangifera indica 163 Melastomaceae 133 Milicia 133, 134 Moraceae 131, 132, 133, 138 Morus 133, 134 Neomorphae (sect.) 137 Nervosae (series) 137 Ocotea 133 Oreosycea (sect.) 137, 138, 139, 142, 143 Orthoneurae (series) 136 Palacomorpha (subsect.) 137 Pallidae (series) 137 Phaeopsilosae (series) 137 Pharmacosycea (sect.) 138 Pharmacosycea (subgen.) 133, 137 Plagiostigmaticae (series) 137 Plagiostigmaticae (subser.) 137 Platypodeae (subser.) 136 Podosyceae (series) 137 Podosyceae (subser.) 137 Psidium guajava 148, 149 Pungentes (series) 137 Religiosae (series) 136 Rhizocladus (sect.) 135, 137, 139, 157, 164 Rubiaceae 132 Rutales 133 Santalum album 149, 152 Scabrae (series) 137 Sideroxylon 133 Spirae 164 Spiraea 164 Sterculiaceae 133 Stilpnophyllum (sect.) 135, 136, 138, 139, 142 Superbae (series) 136 Sycidium (sect.) 133, 137 Sycidium (subsect.) 137 Sycocarpus (sect.) 137 Sycocarpus (subsect.) 137 Sycomorus (sect.) 137, 138 Sycomorus (subgen.) 133, 137 Theaceae 162 Theales 133 Tiliaceae 133 Tuberculifasciculatae (series) 137 Tuberculifasciculatae (subser.) 137 Ulmaceae 132 Urostigma (sect.) 135, 136, 138, 139, 147, 148, 157, 161, 162 Urostigma (subgen.) 133, 135, 136, 160, 161, 162, 164 Urticaceae 131 Variegatae (series) 137 Variegatae (subser.) 137 Varinga (subsect.) 137 D. HOLLIS & P. S. BROOMFIELD INDEX Invalid names are in italics; principal references are in bold. Afrodynopsylla 135, 138, 139, 141 (key), 145, 146 Agaonidae 132 Agaoninae 132 altissimae (Caenohomotoma) 161 altissimae (Homotoma) 136, 161 angolensis 136, 159 Anisostropha 157 annesleae (Caenohomotoma) 162 annesleae (Homotoma) 162 Anomoneura 133, 134 Anomoneurinae 134 Aphalaridae 135, 142 Aphalarinae 133, 134 apsylloides (Macrohomotoma) 152 apsylloides (Pauropsylla) 152, 153 Australohomotoma 157 Austrodynopsylla 135, 138, 139, 141 (key), 142, 143, 144 Austrohomotoma 157 bakeri (Caenohomotoma) 164 bakeri (Caenohomotoma sausia)) 164 bakeri (Homotoma) 136, 157, 164 bakeri (Metapsausia) 164 bambusae (Caenohomotoma) 162 bambusae (Caenohomotoma _ (Hetero- homotoma)) 162 bambusae (Homotoma) 162 bamendae 160, 161 benjaminae (Caenohomotoma) 162 benjaminae (Homotoma) 136, 162 bilineata (Caenohomotoma) 165 bilineata (Caenohomotoma sausia)) 165 bilineata (Homotoma) 137, 164 bilineata (Metapsausia) 165 biliniata 165 boheae (Caenohomotoma homotoma)) 162 boheae (Homotoma) 162 brevigena 133 brunettii 136, 142 buxtoni 133, 137 (Metap- (Metap- (Caeno- Caenohomotoma 157 Carsidaridae 135, 138, 140, 166 Carsidarinae 135, 156 Carsidarini 135, 142 carvalhoi 136, 154 (key), 155, 156 Ceropsylla 133, 134 Chalcidoidea 155 Charipidae 155 chlamydodora 136, 160, 161 chuanana (Caenohomotoma) 162 chuanana (Caenohomotoma (Psausia)) 162 chuanana (Homotoma) 162 Ciriacreminae 135, 142 conchaiensis 137 cornuta 137, 143, 144 (key) cornutiventris 137, 165, 166 Crawfordella 135, 143 Cynipoidea 155 Danainae 132 deflexa 137 depressa 137 Diceraeopsylla 142 Diceraopsylla 134, 135, 138, 139, 141 (key), 142 Diceraopsyllini 138, 141 (key), 142 Diclidophlebia 133 Dilyta 155 distincta (Caenohomotoma (Harrison- ella)) 162 distincta (Homotoma) 136, 157, 158, 162 distincta (Psausia) 162 Drosophila 132 Dynopsilla 143 Dynopsylla 133, 134, 135, 138, 139, 141 (key), 142, 143, 145 Dynopsyllina 138, 141 (key), 143, 145 Dynopsyllinae 134, 135, 138, 141, 146, 147 Dynopsyllini 138, 141 (key), 142, 143, 148, 157 eastopi 161 Edenini 138, 141 (key), 147 Edenus 147, 148 encala 137, 144, 145 Encyrtidae 155 Epipsylla 133 etiennei 137, 154 (key), 156 Eulophidae 132 Euploea 132 Eurytomidae 132 fici (Mycopsylla) 136, 148, 149, 150 fici (Psylla) 148, 149 fici species-group 148, 150 ficicola (Pauropsylla) 133, 136, 137 ficicola (Trioza) 133 ficus (Anisostropha) 159 ficus (Chermes) 157, 159 ficus (Homotoma) 137, 158, 159, 160 ficus (Psylla) 157, 159 fima species-group 132 floccosa 137, 166 fulvida 133, 136 galbvittata (Caenohomotoma) 162 galbvittata (Homotoma) 137, 158, 162 gardenensis (Edenus) 148 gardenensis (Mycopsylla) 136, 148, 149, 150 gardenesis 148 geniculata 136, 152 gigantea 146, 147 gladiata 136, 151, 152 gladiatum 152 globuli 133, 136 grandis (Crawfordella) 144 grandis (Dinopsylla) 144 grandis (Dynopsylla) 137, 143 (key), 144 gressitti (Caenohomotoma) 165 gressitti (Caenohomotoma homotoma)) 165 gressitti (Homotoma) 157, 165 (Austro- Haplaphalara 133 Harrisonella 157 Heterohomotoma 157 Homotoma 134, 135, 138, 139, 141 (key), 157, 158 (key), 166 Homotomidae 132, 134, 135, 138, 140, 146, 147 Homotominae 134, 135, 138, 141 (key), 147, 156 Homotomini 135, 138, 140, 141 (key), 147, 156, 157 hylocola 137, 152 indica (Caenohomotoma) 163 indica (Homotoma) 136, 158, 162, 163 indica (Mycopsylla) 149 indica (Psausia) 162 kenyae 137, 154 (key), 156 kina 137, 148, 149 Labobracha 157 Labobrachia 135, 157, 158 (key) lahui (Caenohomotoma) 163 lahui (Homotoma) 163 laingi 136, 154 (key), 155 lamborni (Rhinopsylla) 145, 146 lamborni (Triozamia) 133, 146 lambourni 146 Limenitinae 132 Lissocephala 132 Macrohomotoma 134, 135, 138, 139, 141 (key), 147, 151, 153, 154 Macrohomotominae 134, 138, 141 (key), 147 Macrohomotomini 138, 141 (key), 151 maculata (Homotoma) 137, 158, 163 maculata (Macrohomotoma) 152 maculata (Psausia) 163 magna 137, 152 magne 152 mangiferae (Caenohomotoma) 163 mangiferae (Homotoma) 163 Marpesiinae 132 mathuriana 136, 149 medleri 154 (key), 155 Metapsausia 135, 157, 158 (key) minana 137, 152 Mycopsylla 134, 135, 138, 139, 141 (key), 147, 148 Mycopsyllini 148 Neolithus 146 nyasae (Macrohomotoma) 154 nyasae (Pseudoeriopsylla) 136, 153, 154 (key), 155 Nymphalidae 132 FICUS-FEEDING PSYLLIDS obliqua 136, 148, 150 obliqua species-group 150 Ormyridae 132 pacifica (Homotoma) 157, 165 pacifica (Labobrachia) 165 Paurocephala 133, 134 Paurocephalinae 134 Pauropsylla 133, 134 Phacopteronidae 138 Phacopteronini 135, 147 Phytolyma 133, 134 pinnativena (Dynopsylla) 137, 143, 144 (key) pinnativena (Dynopsylla cladia)) 144 pinnativena (Sphingocladia) 143, 144 propinqua 151 proxima (Mycopsylla) 136, 148, 149, 150 proxima (Pauropsylla) 136 Psausia 135, 157, 158 (key) Psausiini 157 Pseuderiopsylla 153 Pseudoeriopsylla 134, 135, 138, 139, 141 (key), 147, 151, 153, 154 Pseudoneptis 132 Psyllaephagus 155 Psyllidae 134 Psylloidea 132, 134, 135 psylloptera 137 Pteromalidae 132 pulchar 166 pulchra 166 pumilae 137 (Sphingo- purpurescens 137 pyriformiscola (Caenohomotoma) 163 pyriformiscola (Homotoma) 137, 163 radiata (Caenohomotoma) 163 radiata (Homotoma) 136, 137, 157, 158, 163 radiata (Psausia) 163 radiata (Psausia (Homotoma)) 163 radiatum 163 robusta 136, 153 ruiliana (Caenohomotoma) 163 ruiliana (Homotoma) 137, 163 sandakana 153 secus 155 shuana (Caenohomotoma) 164 shuana (Caenohomotoma homotoma)) 164 shuana (Homotoma) 164 sinica 136, 153 Sphingocladia 135, 143 spiraca 164 spiraea (Caenohomotoma) 157 spiraea (Caenohomotoma homotoma)) 164 spiraca (Homotoma) 164 stevensi (Diceraeopsylla) 142 stevensi (Diceraopsylla) 142 stevensi (Pauropsylla) 142 striata 136, 137, 152, 153 Synoza 134, 135, 138, 139, 140, 141 (key), 158, 165, 166 Synoziini 165 Synozini 138, 140, 141 (key), 156, 165 (Hetero- (Caeno- 183 trichaeta 137 Trioza 133, 134 Triozamia 133, 134, 135, 141 (key), 142, 145 Triozamiina 138, 141 (key), 143, 145, 146 Triozamiinae 145 Triozamiini 145 Triozamini 145 Triozidae 134, 135, 146 triozoptera 137 tuberculata 150 138, 139, udei 137 unifasciata (Caenohomotoma) 164 unifasciata (Caenohomotoma_ (Caeno- homotoma)) 164 unifasciata (Homotoma) 164 usambarensis 133, 146 viridis (Homotoma) 137, 159 viridis (Macrohomotoma) 137, 153 vondraceki 133, 146 willcocksi 137 williamsi 136, 153 wulinensis (Homotoma) 137, 158, 164 wulinensis (Psausia) 164 xishuangana (Caenohomotoma) 164 xishuangana (Homotoma) 137, 164 yunana 137, 153 yunnanica (Caenohomotoma) 164 yunnanica (Homotoma) 137, 164 Bull. Br. Mus. nat. Hist. (Ent.) 58(2): 185-226 Issued 26 October 1989 The species of Poecilominettia, Homoeominettia and Floriminettia (Diptera: Lauxaniidae) in Panama ELCY C. BROADHEAD Department of Pure and Applied Biology, University of Leeds, Leeds LS2 9JT CONTENTS IMRGQUUCUONE (f gmce cone ee Sk wo tn ee ee a oe ot LOS FIONOUUCSET GMT. ct oem a te ee ee eee eee em Material ee TART ee ee ee ee Ce ee ee ees LOT avOnonnc characterise “se ee. oe ee eee ee ES SS Genencevaluatonsy es ee ee eo eee ey ee ene” Toy Key to the genera Poecilominettia, Floriminettia and Homoeominettia . . . . 188 RIGHIMAURENIGACCI 0) ee eek... Woe Seis eee Ce hare peg Homoeominettia gen. n. MeL eeae:,. . Ve egw. Gaede cre 190 RoecHoOnUnciiaticndclawiie ines? . na es ee eee nia 2 lO References AlAeoly easy foe et. .« \eaetieteeeirans Deroern 014 ent 210 Indexmiesane Piaectecsl Lk btdecrecek =. .olletecverers cee dw coe ure 226 Synopsis. Fifty-four species and 2 genera obtained by light-trapping and by fogging the high canopy in forests of the Panama region are newly described. Fifty-one of these species belong to the genus Poecilominettia. The character of the labellum is used for the first time in taxonomic study and has proved useful in generic delineation. Species of Floriminettia are not fungal grazers, but those of Poecilominettia and Homoeominettia are. Analysis of the gut content indicates that 43 species living on the island of Barro Colorado form a large guild feeding upon phylloplane fungi. Of these a group of 29 very closely related and hitherto undescribed species are separated by examination of the male genitalia. It is suggested that there may be a mutualistic association of lauxaniid flies with the trees. Keys to the genera and species of Poecilominettia, Homoeominettia and Flo- riminettia are provided. INTRODUCTION In Panama, flies of the family Lauxaniidae are found, often in large numbers, in forest areas where they have a close relationship with the trees. Adults of a great many species in various genera have a much modified labellum, enabling them to feed upon phylloplane fungi (Broadhead, 1984), while their larvae live mainly as miners of fallen leaves. Of these genera, Pseudogripho- neura, Sapromyza, Xenochaetina, and Poecilo- minettia in particular were found to be represented by many undescribed and very closely related species. This study is, however, limited mainly to those of the last-named genus. An inter- esting taxonomic problem was presented on the island of Barro Colorado where a swarm of very closely related species, hitherto undescribed, belonging to the genus Poecilominettia was dis- covered. An investigation of this prompted a study of other material available to me from vari- ous Panamanian sites. This paper comprises descriptions of all the new species together with a key to three genera encountered in the Panama region, as well as keys to the species. The family Lauxaniidae at present consists of 126 genera, few of which are worldwide in their distribution. Major contributions to the tax- onomic background of this family were made by 186 Malloch and Hendel working independently in the 1920s and 30s, by Curran somewhat later on Central American species, by Shewell on North American species, and more recently by Stucken- berg in his valuable monograph of the Old World genera. Stuckenberg (1971) pointed out that useful characters for defining genera had not been forth- coming and recommended that further studies should be done on the post-abdomen. This applies especially to Minettia, a genus worldwide in its distribution, which Stuckenberg regarded as probably not a monophyletic group but rather an aggregation of species having in common an intra- alar bristle. The genus Poecilominettia was cre- ated by Hendel (1932) for certain Central Amer- ican species which differed from those of Minettia in respect of the position of the anterior frontal and of the ocellar bristles, the type species being picticornis. His definition of the genus Minettia was later clarified by Collin (1948), who worked on the British Lauxaniidae, and by Stuckenberg (IGTAD). In the present paper a contribution towards further delineation of both Poecilominettia and Minettia is offered, and as a result of these evalua- tions certain species are removed and placed in a new genus, Floriminettia. A second new genus, Homoeominettia, is erected to accommodate three species, two of which were originally described by Malloch (1926) in Minettia. Sub- sequently (1928) he placed all three in Deutominettia, in spite of the fact that they did not possess the haired scutellar disc characteristic of that genus, but retained them in his key to the species of Minettia (1928). Flies of the genus Minettia s.str. were not represented in the collec- tions studied here. In the key to genera, Minettia s.Str. is therefore not included. The taxonomic characters used in this study are based on the male genitalia and the labellum, along with the hairing of the arista and body mark- ing. The large fleshy labellum of the Lauxaniidae has not hitherto been used for taxonomic differen- tiation. The evolution of the structures on the borders of the pseudo-tracheal canals, along with the increase in diameter of the canals resulting in a reduction of their numbers, has enabled the flies to crop and ingest solid food particles of fungal matter. Such feeding habits are quite unusual in adult Diptera. The labellum of unique specimens was not removed but, since the flies were preserved in alcohol, this was gently opened out and the main outlines of its morphology readily observed (Fig. 13D). The gut content was not removed in these cases either, but the presence of fungal material E. C. BROADHEAD was easily recognised by the very dark gut showing through the yellow integument. BIONOMICS Microscopic examination of the pseudo-tracheal canals and of the gut content of all the species described here, as well as picticornis and zebroides, but with the exception of those species represented by a single individual, revealed that the food consisted of short lengths of fungal hyphae and fungal spores and was virtually indis- tinguishable from one species to another. On the island of Barro Colorado, with an area of 23 square kilometres, at least 43 species of the genus Poecilominettia co-exist, all of which are known to feed upon the same material, namely phylloplane fungi. Within that group there are 29 species particularly closely related, which are dif- ficult to separate except by examination of the male genitalia, the structures of which are remarkably varied. Particles of food found in the pseudo-tracheal canals of the labellum and within the gut are virtually indistinguishable from species to species and consist of short lengths of fungal hyphae and fungal spores. Such species diversity cannot therefore be accounted for in terms of selective pressures operating on food require- ments. Broadhead (1983) found a similar situation in Psocoptera collected in Panama. Curiously enough the diet of the psocids and that of the lauxaniid flies are almost identical. Mound (1977) also found species swarms in fungal-feeding thrips (Phlaeothripinae) in one site in southern Brazil. These insects were feeding upon fungal hyphae associated with leaf decay on the forest floor. There is considerable interest among ecologists at the present time in guilds of species. Shorrocks & Rosewell (1986), working on Drosophila, have concluded that, in laboratory conditions, guild- size centres on an average of seven species, but in the wild, such a guild would be two or three times that size. The Drosophila species are of course utilising various ephemeral resources, in contrast to the lauxaniids which are feeding on self- regenerating food material that is uniformly dis- persed in space and time. The many species of lauxaniid flies in the various genera present in the rain forest of Barro Colorado Island would form an enormous guild of size far greater than 21. A large number of individuals of a hitherto undescribed species of Drosophila found in the light-trap samples from Barro Colorado Island appear to have joined the lauxaniid guild, for the POECILOMINETTIA, HOMOEOMINETTIA & FLORIMINETTIA 187 adults have a labellum modified for fungal grazing and indeed the gut, on microscopical examin- ation, was found to be packed with short lengths of fungal hyphae of the type found in the gut of the lauxaniids. Flies of the family Lauxaniidae are not regarded as being of any economic importance. In fact it would seem that they are behaving in a way bene- ficial to the trees, by removing fungal hyphae that reduce the amount of light penetrating the leaf tissue - hyphae that might also facilitate the intro- duction of plant diseases. There may well be here the development of a mutualistic situation. Although the larval habits of the species under consideration here are not known, it is generally accepted that the majority of larvae do develop within the tissues of fallen leaves. I have bred various species of British lauxaniids from leaf mould in my garden. Odum (1963) thought that several types of mutualism originate from neu- tralistic associations. | MATERIAL _ Material for this study was obtained in Panama in several ways, namely, from pyrethrum fogging (July-August 1979) of the high canopy of forests ' in the Canal area, from light traps at Fortuna _ (mountain valley), Miramar (coastal valley), and | Barro Colorado Island (1976-83), as well as from _ hand collecting. In addition, a large collection of Lauxaniidae _ from Central America, many not determined, was made available for examination when the United States National Museum in Washington kindly _ sent me their specimens on a long term loan. Type specimens and other material are deposited in the British Museum (Natural His- tory), abbreviated to BMNH, or in the National Museum of Natural History, Washington, D.C. (USNM). TAXONOMIC CHARACTERS The taxonomic characters used in this study are based on the male genitalia, the nature of the labellum, and any markings, especially on the face, frons and scutellum. Occasionally the ring sclerite (7th abdominal segment) of the male proved useful. Features of the female post- abdomen were examined, but since no pairs were captured in copula, it was not possible in most cases to assign the females to any particular spe- cies and so they were omitted from detailed study here. This is especially true of the group of 29 very closely related species of Poecilominettia found on the island of Barro Colorado. Steyskal (1971), in his study of species of the Minettia obscura group in North America, found that the only useful characters for separating five closely related spe- cies were those of the male genitalia. Part of the male genitalia was readily examined without dissection, viz. the epandrium (dorsal part of the periandrium), and the pair of articu- lated lobes borne on the periandrium which func- tion as claspers during copulation (variously termed gonopods, telemeres, basimeres or para- meres). It was necessary to mount the aedeagus ona slide, however, in order to examine the inter- nal structures. Usually the aedeagus is strength- ened on the inner wall of its lobe-like structure by sclerotised spines and by long chitinised rods armed with posteriorly pointing spines. Occa- sionally the aedeagus is a delicate inflatable sac, similar to that observed in the family Celyphidae which, according to Griffiths (1972), is grouped with the Lauxaniidae into the superfamily Lauxanioidea. The hypandrium can in some cases be seen without dissection, but usually its outline is best observed on a slide since its structure is delicate. In some species the ring sclerite is distinctive, when its characteristic shape is a useful additional item in defining the species. The labellum is best examined microscopically. This was done by dissecting off the ventral surface and mounting it on a slide. The two halves, when opened out and flattened, present a heart shape which varies slightly from species to species. The distal area bears upstanding structures in rows along the borders of the pseudotracheal canals (Figs 14, 15). These enable the flies to graze upon phylloplane fungi, proof of which is readily obtained by examination of the pseudo-tracheal canals and gut content. This modification of the feeding surface of the labellum has been described for many species in various genera of Laux- aniidae, both in Britain as well as Panama (Broad- head, 1984). Fungal-feeding species of lauxaniid flies have fewer and broader pseudo-tracheal canals than liquid-feeding species (Broadhead, 1984). GENERIC EVALUATIONS Since the establishment of Minettia Robineau- Desvoidy, 1830, with nemorosa (= rivosa) as type 188 species, the large number of species, whose only common feature is the presence of an intra-alar bristle, has been grouped into a number of genera. Minettia s.str. has been defined most clearly and most recently by Stuckenberg (1971) to include those species with the following characters: an intra-alar bristle, frons usually as wide as an eye, face dull, wings rarely patterned, arista only short- haired, ocelliin an equilateral triangle or nearly so and quite widely spaced, ocellar bristles as wide apart as the posterior ocelli and placed forwards so that they are in line or almost in line with the hind edges of the anterior ocellus. For the British species, Collin (1932) noted that the ocellar bris- tles are situated outside an imaginary line con- necting the outer faces of the anterior and posterior ocelli. I propose to add two further characters, viz. the presence of delicate upstanding structures in rows along the borders of the pseudo-tracheal canals on the distal half of the ventral surface of the labellum, and pseudo-tracheal canals wide, well strengthened by broad half hoops in the distal areas, as well as being reduced in number. For the type species rivosa there are 7 on each half of the labellum. Although the broad half hoops of the pseudo-tracheal canals can be readily seen, the structures bordering them are delicate and best shown on scanning electron micrograph (Broad- head, 1984). In Homoeominettia and Poecilominettia the half hoops in the outer area of the labellum are not wider than those of the basal area, and the struc- tures associated with fungal grazing are well scle- rotised and very easily observed. Certain species currently placed in Minettia do not fall within the definition of the genus and are here transferred to other genera. Two species, tinctinervis Malloch, 1926 (comb. n.) and fusciner- vis Malloch, 1926) (comb. n.) are here placed in Floriminettia gen. n., described below. Three other species, geniseta Malloch, 1926, approximata Malloch, 1928 and assimilis Malloch, 1926, were somewhat tentatively placed in Deutominettia by Malloch, although geniseta and assimilis were originally placed by him in Minettia. All three do not have the striking feature of the genus Deutominettia, namely a haired scutellar disc. They have a shining face and long plumes on the arista (features not characteristic of the genus Minettia). For them I propose to erect Homo- eominettia gen. n., also described below. Two further species currently in Minettia, bru- neicosta Malloch 1928, and quadrata Malloch, 1928, do not fall within the definition of the genus Minettia s.str. in respect of the position of the ocellar bristles. In addition, although the labellum E. C. BROADHEAD has a reduced number of pseudo-tracheal canals with upstanding structures on the distal area, the supporting half hoops are not broad as they are in rivosa. I now place these two species (comb. n.) in Poecilominettia along with 51 of the new species described in this paper. Poecilominettia was erected by Hendel (1932) for certain Central American species that differed from Minettia s.str. in that their anterior orbital bristles are closer to the anterior border of the frons than to the second orbital bristles, that the ocelli are arranged in an isosceles triangle with the sockets of the ocellar bristles close together and well behind the anterior ocellus, and that the ocellar bristles are short and divergent. The type species is picticornis Coquillet, 1898. I propose to add two further characters to this diagnosis, namely, the presence of upstanding structures in rows along the borders of the pseudo- tracheal canals on the distal half of the ventral face of the labellum, and pseudo-tracheal canals wide, ranging in number from 9 to 12 on each half of the labellum, the number being constant for each spe- cies (Figs 14, 15). The shape of the ocellar triangle is probably of less significance here than the position of the ocel- lar bristles on it, i.e., in Poecilominettia the ocellar bristles are situated about half-way between the anterior and posterior ocelli, the space between them varying with the shape of the triangle formed by the three ocelli. Hendel (1932) transferred a number of species from Minettia to Poecilominet- tia which do in fact have ocellar triangles varying from isosceles to equilateral, but whose ocellar bristles always have their sockets within an imag- inary line drawn from the outer face of the ante- rior to the posterior ocellus and situated at least half-way between the anterior and posterior ocelli. KEY TO THE GENERA POECILOMINETTIA, FLORIMINETTIA AND HOMOEOMINETTIA 1 Fronto-facial angle about 90°. Anterior orbital bristle equidistant from anterior border of frons and pos- terior orbital bristle. Labellum with 20 narrow pseudo-tracheal canals on each half, which do not bear any upstanding structures . FLORIMINETTIA — Fronto-facial angle obtuse. Anterior orbital bristle nearer to anterior border of frons than to posterior orbital bristle. Labellum with considerably fewer, wide pseudo-tracheal canals on each half, bearing upstanding structures on their borders on distal Area cWSAL, SS SSS eee 2 POECILOMINETTIA, HOMOEOMINETTIA & FLORIMINETTIA 189 2 Ocellar bristles long, almost parallel and directed for- wards. Arista with long plumes. Yellow flies without dark markings on face, frons or scutellum. Body length not less that 5.5 mm_ ..HOMOEOMINETTIA — Ocellar bristles short and divergent. Arista more fre- quently with short hairs, but if with long plumes, then distinct black markings on face, frons and scutellum, and thorax with black stripes. Predominantly yellow flies. Body length 4.5 mm or less. POECILOMINETTIA FLORIMINETTIA gen. n. Type SPECIES: Floriminettia coronata sp. n. DiaGnosis. An intra-alar bristle, anterior part of frons drawn out to form a small peak between antennal bases, face shining centrally with silvery dusting at eye margin, labellum with 20 narrow pseudo-tracheal canals on each half, without any upstanding structures on their borders, wings with veins outlined, arista with hairing of medium length, ocellar bristles placed well behind anterior ocellus and fairly close together. Key to species of Floriminettia 1 Scutellum with black spots at apex. One large triangu- lar spot centrally on Sth abdominal tergite. Wings with costal infuscation and all other veins strongly outlined fuscinervis (Malloch) — Scutellum without marks at apex ............... 2 2 3rd antennal segment oval, twice as long as wide. Two lateral spots on Sth abdominal tergite. Wings with costal infuscation and all other veins strongly outlined tinctinervis (Malloch) _ — 3rd antennal segment tapering, three times as long as _ width at base. No spots on abdominal tergites. Wings without costal infuscation and all veins distinct but PENI UM RR ahd ie: ecg alin sieincte tes coeds sfsigne eign jorene cee 3 Ww Thorax with central brown stripe and broken stripe on either side. Anterior part of frons produced upwards centrally at antennal bases. Body length 6 mm ficulnea sp. n. — Thorax not striped. Anterior part of frons barely pro- duced upwards at antennal bases. Body length 7 mm coronata sp. Nn. Floriminettia coronata sp. n. Large yellow fly. Body length 7 mm. All bristles strong. Head yellow with face keeled centrally. Fronto- facial angle about 90°. Frons wider anteriorly and drawn out to form small peak between antennal bases. Antennal segments 1 and 2 slightly longer than wide, segment 3 long-oval (3 times width) and covered with hairs, the dorsal ones being half width of segment. Aristal hairs medium long. Orbital plates shining, closer together anteriorly than posteriorly. Anterior orbital bristles shorter than posterior. Ocellar plate yellow. Face shining centrally, with silver dusting along eye margin. Eyes red, oval. Palps yellow. Labellum with 20 narrow pseudo-tracheal canals on each half which do not bear any upstanding structures. Thorax unstriped, orange-yellow with scutellum paler and edged with fine dark border. Acrosticals in 8 rows. Prescutellars reaching as far as scutellum apex. Wings slightly smoky yellow, veins dark yellow. Legs yellow. Four postero-ventral bristles, 5 postero-dorsals, 3 nearly true dorsals on femur 1; 6 very sturdy short bristles antero-ventrally on apical half and one posterior bristle apically on femur 2; 4 strong proclinate bristles antero-ven- trally on femur 3. Ring of strong spines apically on tibia 2. Dorsal pre-apical of tibia 2 much stronger than that of other tibiae. Abdomen same colour as thorax without marks dorsally, other than fine darkened borders posteriorly on tergites. Paler ventrally and sparsely haired. Ring sclerite (7th abdominal segment) yellow. Male genitalia (Fig. 13E, F). Clasper small with hooked tip. Hypandrium with long pointed pro- cesses. Aedeagus bilobed at tip and containing very small spines. No mark on epandrium. Female similar to male. Hototyre. O', Barro Colorado Island: iii. 1983, light-trap (BMNH). PARATYPES. Panama: 4 ©’, Panama City end of Panama canal, vii. 1979, fogging high canopy; 2 ©’, Barro Colorado Island, xi. 1982, light-trap, v. 1983; 1 9, Barro Colorado Island, x. 1982, light- trap (all BMNH). Floriminettia ficulnea sp. n. Yellow fly. Body length 6 mm. All bristles strong. Head yellow with face keeled centrally, shining but with silver dusting along eye margin. Fronto- facial angle about 90°. Frons wider anteriorly and produced upwards at antennal bases, and drawn out to form small peak between antennal bases. Antennal segments 1 and 2 slightly longer than wide, segment 3 long-oval and tapering (3 times width at base) and covered with hairs. Arista with short hairs. Anterior orbital bristle shorter than posterior. Orbital plates shining, closer together anteriorly than posteriorly. Ocellar plate yellow. Ocellar bristles short, divergent. Frons orange- 190 yellow. Palps yellow. Labellum with 20 pseudo- tracheal canals on each half which do not bear any upstanding structures. Thorax with a central brown stripe and a broken stripe on either side. Acrosticals in 12 irregular rows. Prescutellars reaching to the apex of the scutellum. Wings faintly yellowish with veins yellow. Legs yellow. Femur 1 with 7 bristles irregularly on row curving from postero-dorsally at base to dorsally at apex and 6 bristles posteriorly; femur 2 with 6 bristles antero-ventrally, one strong posterior bristle, a row of long hair-like bristles on apical half, 6 bristles irregularly antero-ventrally; femur 3 with arow of hair-like proclinate bristles on apical half, and one strong bristle postero-dorsally near apex. Tibia 2 with dorsal pre-apical, ventral apical bris- tle and apical spines all very strong. Abdomen yellow, without marks. Male genitalia not dissected, since holotype somewhat fragile. Female similar to male. Ho.otyre. ©’, Panama; Barro Colorado Island, ex fruit of wild Ficus, x. 1937 (Zetek no. 4421, Lot no. 39 11659) (USNM). ParATyPE. | 2, same data as holotype. These two specimens are labelled in pencil ‘“Nimet- tia immaculata type’ but without any author's name. They do not exhibit the characters of the genus Nimettia and clearly are closely related to the other species of Floriminettia. HOMOEOMINETTIA gen. n. Type species. Deutominettia assimilis Malloch, 1926. DiaGcnosis. An intra-alar bristle, face shining, arista with long plumes, anterior orbital bristle nearer to anterior border of frons than to second orbital bristle, length of third antennal segment at least 2.5 times width, ocellar bristles long, almost parallel and directed forwards, their sockets situ- ated about half-way between anterior and pos- terior ocelli and well separated, ocelli arranged in an isosceles triangle, spines usually present on mid tibia postero-ventrally, labellum with reduced number of pseudo-tracheal canals bearing upstanding structures on distal half of ventral surface. Key to species of Homoeominettia 1 Abdomen with dark markings on tergites. No spines postero-ventrally on mid tibia. Third segment of ante- nna long oval (3 times width). Body length 8 mm woldae sp. n. E. C. BROADHEAD — Abdomen without markings. Spines present postero- dorsally on mid tibia. Third segment of antenna shorter, oval (2.5 times width). Body length about 5.5 2 Three or four long spines on mid tibia postero-ven- tral: - cordata sp. n. POECILOMINETTIA, HOMOEOMINETTIA & FLORIMINETTIA Key to species of Poecilominettia from Panama and neighbouring regions 1 At least segment two of antenna black, thorax dis- tinctly striped — All segments of antenna yellow or yellowish, thorax SIMPECOMMOUSIMIDER 9... .e enc eee e es ons ree 14 i) Two basal segments of antenna black .......... 3 — Only segment two of antenna black, palps yellow, no stripes on sternopleuron or mesopleuron (Barro Colorado Island) ........ fungivora sp. n. Ww Wings with conspicuous markings in addition to clouding over cross-veins . tripuncticeps (Malloch) — Wings unmarked apart from possible clouding over ETI MSRTISMENS ces NEST iG cosa eye s.s.613. ah er eakers ages 4 4 Six thoracic stripes Thoracic stripes distinct Thoracic stripes absent or at most very faint ... 18 Wings with distinct marks in addition to clouding on cross-veins octovittata (Williston) Wings unmarked apart from clouding on cross-veins 16 Pleurae with stripes Pleurae without stripes BA. WORT TR 17 Palps yellow, abdominal tergites with dark posterior borders and central marks. Clasper of male large, wider than long, aedeagal internal spines very con- voluted (Fig. 3A) zebroides Hendel Palps black. Abdominal tergites without dark pos- terior borders, but with central marks. Clasper of BRGME THORACIC SETIPES i siz. .ic vice eid vlele ees ene es 5 male rounder (Fig. 2F), aedeagal spines less con- SUIS ANWATOMONOS DIMES. oc. cne cs ded sna ee eons 6 Se te ee aa bare = ASV RIGUNTUITS (orga) °c) Cie ae eee 7 ee 6 Acrosticals in 10 rows, frons slightly longer than 18 | No marks on scutellum Two black marks at apex of scutellum which may DSRS crate 22D E. C. BROADHEAD ww S0*O O 0 eS a ww S*0 ww S°0 Fig. 12 A, B, clasper of (A) Poecilominettia folleata; (B) P. acuta;C,D, hypandrium of (C) P. folleata; (D) P. acuta; E, F, ring sclerite of (E) P. circularis; (F) P. ungulata. POECILOMINETTIA, HOMOEOMINETTIA & FLORIMINETTIA 223 Fig. 13 A, clasper and aedeagus of Poecilominettia aurita, a = clasper, b = aedeagus; B, aedeagus of P. gatuna; C, clasper of P. gatuna; D, labellum of P. pectinata, a = modified zone, b = pseudo-tracheal canals, c = beak; E, aedeagus and processes of hypandrium of Floriminettia coronata, a = aedeagus, b = process of hypandrium; F, clasper of F. coronata; G, tip of male abdomen of Poecilominettia semilunata, a = epandrial black spot, b = claspers, c = aedeagus, d = hypandrium, e = ringsclerite. E. C. BROADHEAD Fig. 14 A, head of Poecilominettia effossa, \abellum fully extended; B, labellum enlarged to show structures on distal half. POECILOMINETTIA, HOMOEOMINETTIA & FLORIMINETTIA 225 Fig. 15 Labellum of Poecilominettia effossa in lateral view. E. C. BROADHEAD INDEX acuta 193 approximata 188 assimilis 188 aurita 193 biprojecta 194 breviplumata 191 bruneicosta 188 calva 194 chelata 194 chilensis 192 circularis 195 circumtexta 195 cordata 195 cornuta 196 coronata 189 curvata 196 Deutominettia 186 Drosophila 186 effossa 196 effossa group 192 enormis 197 epacra 197 erymna 197 falcata 198 ficulnea 189 fimbriata 198 flavescens 198 Floriminettia 189 foliacea 199 folleata 199 Invalid names are in italics. fornicata 199 fortunae 200 fumida 200 fungivora 200 fuscinervis 188 gatuna 201 geniseta 188 grata 192 Homoeominettia 190 lagenata 201 legnota 201 lineolata 202 maniculata 202 membranosa 202 Minettia 186 nemorosa 187 nigriapica 203 notata 203 obscura group 187 obtusa 203 octovittata 191 papillata 204 parouatia 204 pectinata 204 pedata 205 picticornis 186 plicata 205 Poecilominettia 190 Pseudogriphoneura 185 pygmaea 205 quadrata 188 quadriprojecta 206 remata 206 rivosa 187 _ Sapromyza 185 semilunata 206 sentosa 206 sexiprojecta 207 sexiseriata 191 silbergliedi 207 silvicola 207 spinosa 208 tinctinervis 188 trigona 208 uncata 208 ungulata 209 unicolor 192 valida 191 vibrata 209 virgea 209 woldae 190 Xenochaetina 185 zebroides 186 E. W. CLASSEY Ltd P.O. Box 93 FARINGDON, OXON SN7 7DR, ENGLAND SE ——e Natural History Publishers Booksellers and Library Agents rir Catalogues on request Agents for the publications of the BRITISH MUSEUM (NATURAL HISTORY) 0 Publishers of ENTOMOLOGIST’S GAZETTE An illustrated quarterly journal of Palaearctic Entomology Volume 41 commences January 1990 Current Annual Subscription (past free) £16.00 (US $35.00) pee as: in Panama. Elcy C. Broadhead a é . a “ s bd . ' ~ ee. s 2 "g » Fi 3 x . : ‘ } ova “ a, <> ia Pn at » : a ~ %- a 4 4” ; ay +; + e 4 j re Pm 7 ee ‘ s BN, > - uy . a cof 3 ; ; : - sad = © ” * P é , Py ‘ = ae é & Sd eo ‘ _ eta ty Gaye i J t a. i nae 2. 2p . fot ‘ ‘ LS. ge , - } yy * ] €A ‘ wa ‘ ’ 4 " & . : 4 t “—s vy * " ae n 2 se . ’ ¥ & =? y Bus ; x a, ga" , lj % s > % 4 < bing N - 3 i . 3 4 i**s > — ‘ . ‘ é . . i ¥ 3 , et . c . * vy. « ~ ’ 7 ENTOMOLOGY SERIES Vol. 58, No. 2, October 1989 “ P re ome z , ‘ x * ‘ee ek a? se ‘ bs toe Bae * , ~ +% - ' wae 5 - te ? a: oe . > ui = ‘ -