* seen ou ae) seer Lt AT eve ae , KOC? ehh etre recta tes o. \) \“\ i Cas re 7 7 ‘i Rk Wag oe i i : Lind 7 avy tora BPUPEETIN OF ori bRItish MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 BRITISH MUSEUM (NATURAL HISTORY) LONDON : 1976 sg Printed in England by v 5 Unwin Brothers Limited CONTENTS ENTOMOLOGY VOLUME 34 Jumping plant lice of the tribe Ciriacremini (Homoptera : Psylloidea) in the Ethiopian Region. By D. HoLtiis A taxonomic revision of the genus Ornativalua Gozmany, 1955 (Lepidoptera : Gelechiidae). By K. SATTLER The taxonomy of the genus Heteropelma Wesmael (Hymenoptera: Ichneumonidae). By I. D. GAULD A revision of the Indian species of Haplothrips and related genera (Thysanoptera: Phlaeothripidae). By B. R. Pirk1n The ant tribe Tetramoriini (Hymenoptera : Formicidae). Constituent genera, review of smaller genera and revision of Tviglyphothrix Forel. By B. BoLtton Ae a_i _ — h \ JUMPING PLANT LICE/ °7,, OF THE TRIBE CIRIACRE INT’ ; (HOMOPTERA : PSYLLOIDEA) IN THE ETHIOPIAN REGION D. HOLLIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 1 LONDON : 1976 . Mt) : j > / 0 Pe = JUMPING PLANT LICE OF ‘THE TRIBE, . ting 1% CIRIACREMINI (HOMOPTERA : PSYLLOIDEA) IN THE ETHIOPIAN REGION BY DAVID HOLLIS Pp. 1-83; 191 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 1 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five serves corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 34 No. 1 of the Entomology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) ISSN 0524-6431 © Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 27 May, 1976 Price £6.50 JUMPING PLANT LICE OF THE TRIBE CIRIACREMINI (HOMOPTERA : PSYLLOIDEA) IN THE ETHIOPIAN REGION By D. HOLLIS CONTENTS Page SYNOPSIS . 3 ; é : : 2 ; F ‘ ‘ : 3 INTRODUCTION . 3 < : 5 : - , é : - 3 HISTORICAL : , F : : : : : ; : : 4 MATERIAL, METHODS AND TERMINOLOGY , : ; : : : 6 ACKNOWLEDGEMENTS , : : ? : : : ; : 10 TAXONOMIC TREATMENT j : 3 : ‘ : : : : 10 Tribe Ciriacremini . , : : ; / ° : : : 5 fe) Key to genera of Ciriacremini : : : ; : ‘ - 13 Palmapenna gen. n. . ~ : : : é : ‘ : 14 Kleiniella Aulmann, 1912 = : : : ; : * : 15 Key to species of Kleiniella . - : - . : : : 18 Civiacremum Enderlein, 1910 . ; P . , : E ; 24 Key to species of Civiacremum : : , : . : : 26 REFERENCES. ; : : A , ; : * - - 53 INDEX é - : ; : F 2 3 ; : ; : 82 SYNOPSIS An historical account of the taxonomy and nomenclature of the group called here the tribe Ciriacremini is given. The tribe Ciriacremini is redefined, its internal phylogeny briefly discussed and a key is given to the five genera included. In the Ethiopian region 30 species in three genera are recognized and keys are given to all these taxa. One genus and 21 species are described as new and the remaining taxa are redefined. Three new generic synonyms and three new combinations are proposed, and four species are replaced in their original cpmbinations. INTRODUCTION THE psyllid faunas of the tropics, subtropics and the south temperate region are poorly known and that of the African continent is no exception. Currently 92 described species are known from Africa, south of the Sahara, but recent collecting in the region would suggest that this figure represents only a small proportion of the existing fauna. For example, prior to the author’s recent field trip to East Africa, nine described species of Psylloidea had been recorded in the literature from the area, yet on this visit approximately 90 species were collected and most of these appear to be undescribed. 4 D. HOLLIS Psylloidea can transmit plant viruses, are pests of fruit, ornamental and timber trees, and one species is known to have developed resistance to several insecticides (McMullen & Jong, 1971). Research on the systematics of the group is of increasing interest to foresters and economic entomologists and the need for some alpha- taxonomic studies on tropical faunas is abundantly clear. Some such work was begun by Karel Vondracek in the last decade shortly before his death. Capener (1968; 1970a; 1970); 1973) produced some useful work on the South African fauna but he has now retired leaving a large collection of unworked material. About 15% of the material recently collected from Africa consists of specimens in which the fore wing bears a clearly developed vm crossvein, suggesting they are related to Enderlein’s genus Civiacremum. As much of this material represents undescribed taxa, a taxonomic study of the African Ciriacremini is given below. Prior to this work the group was thought to be a tribe in the subfamily Psyllinae containing, worldwide, five monobasic genera, a genus with four species and a genus with two species. The current work, based on a study of over 2,250 specimens, agrees with the systematic position of the group but rearranges the Ethiopian constituents into one new monobasic genus, a genus with six species, two of which are new, and a genus with 23 species, 18 of which are new. The New World component of the group, three species in three genera, is not considered here to any depth because only five specimens were available for study. HISTORICAL Enderlein (19100) first used the name Ciriacreminae as a replacement name for Prionocnemidae Scott, 1882, pointing out that Scott’s name was invalid as it was not derived from a generic name in the group to which it was applied. Scott included two genera, Carsidara and Tyora, in his group, and Prionocnemidae was used in various derivations by Léw (1886), Froggatt (1901), Kieffer (1906) and Kuwayama (1907). None of these authors widened the scope of the group. Kieffer (loc. cit.) erected the subfamily Phacoseminae for two genera Phacosema and Phacopteron, and Kuwayama (loc. cit.) placed the genus Anomoneura into the Phacoseminae. Enderlein considered this group to be a tribe in the Ciriacreminae. Enderlein’s principal character defining the Ciriacreminae is the presence of an vm crossvein between the radial sector (Rs) and the upper branch of the medial vein (M,,,) or the punctiform contact between Rs and M,,,. He divided the subfamily into two tribes, the Ciriacremini and the Phacosemini, on the presence or absence of a basal spine on the hind tibia. In the former tribe he placed the genera Ciriacremum, Panisopelma, Tyora, Carsidara and Udamostigma, and in the latter tribe he placed Phacosema and Phacopteron. These two latter genera were tentatively synonymized by Crawford (1912) and Laing (1930) confirmed the synonymy. Aulmann (1912b) added Kleiniella to the Ciriacremini and, in his psyllid catalogue (Aulmann, 1913), included the following genera in the Ciriacre- minae: Carsidara, Kleiniella, Tyora, Geyerolyma, Panisopelma, Ciriacremum, Phaco- sema, Phacopteron, Anomoneura, Udamostigma and Nesiope. Sulc (1914) erected ee ee A ae ee ee CIRIACREMINI OF THE ETHIOPIAN REGION 5 Connectopelma as a subgenus of Panisopelma and Enderlein (1927) further added the genus Desmiostigma to the group. . Crawford (1911) seems to have partly misunderstood Enderlein’s work for, although agreeing that Prionocnemidae Scott was an invalid name, he also rejected Ciriacreminae Enderlein for a spurious nomenclatural reason. However, he quite rightly pointed out that Enderlein’s subfamily was a rather heterogeneous mixture of unrelated genera and went on to define a subfamily, the Carsidarinae, contain- ing the following genera: Carsidara, Tyora, Mesohomotoma, Tenaphalara, Macrohomotoma, Epicarsa, Freysuwila, Homotoma, Bactericera and Rhinopsylla. Later (Crawford, 1914) he partially accepted Enderlein’s subfamily, using the name ‘Ceriacreminae’ for two New World species. Also in that paper Crawford suggested the ‘Ceriacreminae’ were probably most closely related to the Psyllinae. Heslop-Harrison (1958), in his reorganization of the classification of the Psylloidea, went to considerable lengths to sort out the nomenclatural complications revolving around the Ciriacreminae but then proceeded to introduce several taxonomic complications. He used the Ciriacreminae in a very broad sense to include the old Ciriacreminae of Enderlein and the Carsidarinae of Crawford, and many other groups of genera in what amounted to a ragbag subfamily for those genera he could not place in the other subfamilies, i.e. Liviinae, Aphalarinae, Psyllinae, Spondyliaspinae and Triozinae. He split the subfamily into seven tribes including the Ciriacremini and Carsidarini. In the former tribe he placed the genera Ciriacremum, Bunoparia, Kleiniella, Panisopelma, Connectopelma and Anomalopsylla. At no point in this paper did Heslop-Harrison clearly define the Ciriacreminae but in the key to tribes the Ciriacremini were stated to have ‘. . . venation not triozine, antennae long and slender. Fore wing with a distinct tracheate ym crossvein or with Rs and M,,, closely approximating. Head not cleft in front’. In a later paper (Heslop-Harrison, 1960) he mentions that members of both the Carsidarini and the Ciriacremini have a bipartite male proctiger, secondary terminal epiphyses (‘Sternalzapfen’ of Enderlein, ‘Hypovalves’ auctt.) in the male, and an rm crossvein or with Rs and M,., closely approximating. He distinguished the latter group by the non-cleft head, the ‘usual’ development of genae and the ‘normal’ insertion of the antennae. Vondracek (1957) in his figure 52 obviously considered the Ciriacreminae in a more restricted sense, possibly that of Crawford, and placed the group close to the Psyllinae and Arytaininae. Later (Vondracek, 1963) he described the monobasic African genus Syndesmophlebia, placed it in the Ciriacreminae and differentiated it from Civiacremum, Anomoneura, Bunoparia, Panisopelma and Connectopelma. Also in that paper (Vondracek, loc. cit.) he erected the tribe Anomalopsyllini within the Spondyliaspinae for the Australian genus Anomalopsylla. Klimaszewski (1963) erected a separate tribe Anomoneurini for the Asian genus Anomoneura because of the multiple branching of Rs in the fore wing of the two known species. Both Klimaszewski (1964) and Bekker-Migdisova (1973) in their classifications of the Psylloidea placed the Ciriacreminae in the family Psyllidae. The latter author separates the group from other members of the family Psyllidae on the presence of an ym crossvein or contact between Rs and M,,., a bipartite male proctiger 6 D. HOLLIS and the presence, in some species, of rudimentary genal cones. She further separates the group into two tribes, the Ciriacremini and the Anomoneurini, and distinguishes the former by the absence of a pectinate Rs and the presence of more or less developed hypovalves on the male subgenital plate. MATERIAL, METHODS AND TERMINOLOGY Most of the material studied in this work was collected in the last twenty years by a number of field entomologists working in various parts of Africa. Principals among these are Drs José Passos de Carvalho and Tony van Harten in Angola, Laurie Capener in South Africa, Professor J. T. Medler in Nigeria, Dr V. F. Eastop in West and East Africa, and the present author in South-western, Central and East Africa. The bulk of this material is now incorporated into the collections of the British Museum (Natural History). The collections of the Central African Museum, Tervuren, and the National Collection of Insects, Pretoria, provided valuable additional material. Type-depositories are given in the text below in abbreviated form as follows. MNHU, Berlin Museum fiir Naturkunde der Humboldt-Universitat, Berlin. SAM, Cape Town South African Museum, Cape Town. IPK, Eberswalde Institut fiir Pflanzenschutzforschung Kleinmachnof, Eberswalde. ZI, Leningrad Zoological Institute, Academy of Sciences of USSR, Leningrad. BMNH, London British Museum (Natural History), London. NCI, Pretoria [National Collection of Insects] Plant Protection Research Institute, Pretoria. NM, Nairobi National Museum, Nairobi. NR, Stockholm Naturhistoriska Riksmuseum, Stockholm. MRAC, Tervuren Musée Royal de 1’Afrique Centrale, Tervuren. IZPAN, Warsaw Instytut Zoologiczny, Polska Akademia Nauk, Warsaw. USNM, Washington National Museum of Natural History, Washington. The material is stored dry (on card points or micropins), in 80° ethanol, or as permanent microscopical preparations on glass slides in a modified Berlese mountant. It was found necessary to make these preparations in order to study antennal structure, wing chaetotaxy and genital structures. All measurements quoted below were made from slide mounted material. The reference points within which measurements were taken of the various structures are shown in Text-figs 2-4, 9, 14 and Ig. Structural terminology (Text-figs 1-19) used follows that of Vondracek (1957) and Dobreanu & Manolache (1962). One point where I differ from previous authors, e.g. Crawford and Vondracek, is in the use of the terms ‘spur’ and ‘spine’, particularly in relation to the hind tibia. In this paper the term ‘spine’ is used for a prolongation of the cuticle which does not have an articulated base (Text-fig. 14), and ‘spur’ is used for a structure with an articulated base and is a derived seta (Text-figs 13, 14). Thus the structure at the base of the hind tibia is called a spine and those at the apex are spurs, exactly the opposite to the previous nomenclature of these structures in Psylloidea. CIRIACREMINI OF THE ETHIOPIAN REGION a Fics 1-8. Ciriacremini species, morphological features. Palmapenna hymenostegioides, I, head and prothorax, lateral view. Civiacvemum bicaudatum, 2, head, dorsal view; 6, seventh antennal segment; 7, head and thorax, dorsal view. Kleiniella jassina, 3, rostrum; 5, seventh antennal segment. C. filiverpatum, 4, rostrum. C. africanum, 8, head and thorax, dorsal view. (aol—anteoccipital lobe; as — antennal socket; gc — genal cone; lo — lateral ocellus; lurs — length of ultimate rostral segment; mo — median ocellus; ms— median suture; mwh-—maximum width of head; pem — proepimeron; pes — proepisternum; pn — pronotum; v — vertex.) Scale line represents 0:1 mm. 11 12 Fics 9-14. Ciriacremini species, morphological features. Czviacremum bicaudatum, 9, fore wing; 10, hind wing; 14, hind tibia. Kleiniella jassina, 11, fore tibia. C. pervatum, 12, fore tibia. K. superba, 13, apex of hind tibia. (A —anal vein; ab-—anal break; asr — apical spurs; bsp — basal spine; C — costa; c— costal cell; cb —costal break; cls — claval suture; cs — costal setae; Cu —cubital vein; Cu,, —first cubital branch; Cuy — second cubital branch; cu, — first cubital cell; cu,—second cubital cell; lfw — length of fore wing ; Iht — length of hind tibia; Isr — lateral spurs; M—medial vein; M,, ,—combined first and second medial branch; M,, ,-combined third and fourth medial branch; m, — first medial cell; m,—second medial cell; mwfw-— maximum width of fore wing; R—- radial vein; R, — first radial vein; Rs —radial sector; 7,, 75, 73 — first, second and third radial cells; vm-—radio-medial cross-vein; rt—retinaculum; vsr—ventral spurs; Sr Meo CIRIACREMINI OF THE ETHIOPIAN REGION iat Fics 15-19. Ciriacvemum species, morphological features. C. bicaudatum, 15, male genitalia, lateral view; 16, male hypovalve, lateral view of inner surface; 17, male paramere, lateral view of inner surface; 18, distal segment of aedeagus. C. africanum, 19, Ovipositor, lateral view. (dsa-—distal segment of aedeagus; hv —hypovalve; iat —inner apical tooth; ipvs — inner postero-vertical setae; Ipgr — length of proctiger; p-—paramere; pbs —postero-basal setae; pgr—proctiger; pmr-— postero-medial ridge; pms — postero-medial setae; sgp—subgenital plate; tlt—terminal tube.) Scale line represents o°I mm. Io D. HOLLIS ACKNOWLEDGEMENTS I would like to thank Dr V. F. Eastop, of the British Museum (Natural History), for showing sustained interest in the preparation of this work, and, together with Mr P. M. Hammond of the same institution, for offering constructive criticism of the manuscript. Dr N. K. B. Robson, of the British Museum (Natural History), and Dr J. B. Gillett, of the East African Herbarium, Nairobi, among others, provided much valuable expertise in the identification of host plants. Sheila H. Halsey, of the British Museum (Natural History), kindly helped with some of the micro- scopical preparations, and Tony Sutton, of the same institution produced the fine illustrations of the larvae. The following colleagues are acknowledged for their help with loans of type-, and other, material for study: A. L. Capener, formerly of the Plant Proctection Research Institute, Pretoria; Dr José Passos de Carvalho and Dr A. van Harten of the Instituto de Investigacao Agronémica de Angola; Dr U. Gdéllner-Scheiding of the Museum fiir Naturkunde der Humboldt-Universitat, Berlin; E. Holm and E. Hartwig of the Plant Protection Research Institute, Pretoria; Paul D. Hurd Jr, of the National Museum of Natural History, Washington; Dr T. Kronestedt of the Naturhistoriska Riksmuseet, Stockholm; Professor J. T. Medler, formerly of the University of Ile-Ife, Nigeria; Dr J. Nast, of the Polish Academy of Sciences, Warsaw; Miss L. M. Russell, formerly of the United States Department of Agri- culture, Maryland; and Dr G. Schmitz of the Central African Museum, Tervuren. TAXONOMIC TREATMENT Tribe CIRIACREMINI Ciriacremini Enderlein, 1910) : 137, partim. Type-genus: Civiacremum Enderlein. Ciriacreminae Enderlein; Aulmann, 1913 : 79, partim. Ceriacreminae Enderlein; Crawford, 1914 : 62. Ciriacremini Enderlein; Heslop-Harrison, 1958 : 563, partim. Ciriacremini Enderlein; Vondracek, 1963 : 273. Ciriacremini Enderlein; Bekker-Migdisova, 1973 : III. Small to large size, head width 0:52-1:46 mm. Body robust or somewhat elongate. Head as broad as thorax and distinctly separated from thorax, in profile slightly depressed from axis of body; vertex wider than long with complete median suture, mostly clearly demarked from genae by transverse sutures, anteoccipital lobes mostly present; genae covering frons dorsally and either slightly bulbous or developed into cones. Antenna filiform to elongate filiform, rhinaria only present on segments four, six, eight and nine. Thorax convex from above, arched in profile, pronotum mostly narrower than head; propleurites mostly subequal and both touching pronotum dorsally. Fore wing parallelo- grammatic or ellipsoid; costal and anal breaks present, pterostigma mostly present, vm crossvein present or there is punctiform contact between Rs and M,,., M and Cu with common stem. Hind coxa with clearly developed meracanthus, hind tibia mostly with a well developed basal spine, hind basitarsus with a pair of apical spurs. Male proctiger shaped to give broad basal part and narrow apical part which may be long or short, subgenital plate often with a pair of apical appendages (hypovalves). Female proctiger and subgenital plate subconical, sometimes elongate, former often longer than latter, CIRIACREMINI OF THE ETHIOPIAN REGION II DISTRIBUTION, HOST PLANTS AND BIOLOGY. The bulk of the known species are restricted to the Ethiopian region but three species apparently belonging to the tribe are recorded from Central and South America. Much of the material studied was collected at light and only four species are known from larval stages as well as adults. Of the 30 known species in the Ethiopian region only nine have any recorded host data but these species were all found on trees of the leguminous family Caesalpiniaceae. There are no host records for the six species placed in the genus Kleiniella but all are known from both lowland and montane forest regions of Africa (Map 2). Cuiriacremum species seem to have successfully exploited both forest and woodland areas, their typical habitat being the Brachystegia woodland zone in Central Africa (Maps 3-7). Recorded host genera for Ciriacremum are: Brachystegia, Hymenostegia, Cynometra, Cassia, Julbernardia and Schotia and the distribution of these genera in Africa is summarized in Map I. From personal observations it seems that only a few species of Psylloidea are attracted to light and their response to artificial light sources seems to vary considerably with their state of maturation. However, members of the Ciriacremini are frequently caught in light traps and Dr Passos de Carvalho, working in Angola, informs me in a personal communication ‘I never saw so many Psyllidae attracted to light; here in Nova Lisboa thousands of these examples (Ciriacremum) came through my bathroom window and the bottom of the bath tub was almost covered with them’. Table I is a summary of the numbers of specimens of six apparently sympatric species occurring in Central Africa in Brachystegia woodland. The material was collected in a variety of ways; on host plants, as vagrants, by yellow trays and light traps. From this table one can see the main period of adult activity takes place between August and December, roughly coincidental with the end of the dry season and the onset of the rains, with adults and larvae found on host plants between September and March. Although these data are very incomplete and the sampling not at all random it does suggest that Ciriacremum species in Central Africa have only one generation per year with a dry season diapause probably ina larval stage. It is of interest to note that from over 250 microscopical preparations of females, eggs were found in only five specimens. The known larvae (Text-figs 24-27) are of the psylloid type, free living on the leaves and shoots of their host plants. They do not produce wax filaments and have little apparent affect on their hosts. Larvae of Civiacremum harteni sp. n., occurring in large numbers on regenerating Brachystegia tamarindoides, were causing some leaf atrophy on young shoots. Discussion. As mentioned earlier (p. 5) Bekker-Migdisova suggests that the Ciriacremini is the sister group of the Anomoneurini, a tribe with two species occurring in the Oriental region and probably developing on trees of the family Moraceae. As I have no evidence to the contrary this suggested phylogeny within the Ciriacreminae is accepted here but with reservations. The presence of an rm crossvein or the punctiform contact of Rs and M,., is considered here to be a derived feature in the modern Psylloidea and has developed independently several D. HOLLIS I2 ’ aT *(1r ‘d) 3x0} 00s UoMeURdxe J9q}INj 10,4 “AeI} MOTAA = 34 “QueIdeA = A ‘soy UO = Yo ‘eArIe] = u ‘dex} 4q9t = 3] *MOI [E}UOZLIOY seq UI USATS oe Bore O[OYM 94} IOJ SeInsy [Teyurex ATqyUoM ues, $ Zz FE Seem tad S1r £11 gir (uru) Treyureyy 34 SSS (PS yo 34 ‘dds maajshysvag uo fu‘Sz‘Pr} ‘sdbxr ‘Por | qo! Spr ‘Pr 34 Sr ‘Po 1uayAvy 44 SSr ‘Po ysoq ¢ uy tdzo ‘PLb] ap SShe “Pre If Srr ‘Pez wnunnafosaur A ‘Go ‘Px yh SSr1 (Pr A ‘SSbr (PS | ap SHrx “Per 34 S&o PS yo ‘ds vissy9 uo widse‘Por| ‘der ‘Per] ar‘do ‘Ps ununnsfo A‘SSE ‘Pr 44 S&o ‘Pr WSS “Poe}] 44 ‘So ‘Pr yo yo ‘dds wypavusaqinf[ uo fu ‘Sr ‘Pe| fu‘gs “pe saprovpsvusaqnt 44 SS “Po qsoy ¢ qidor Por} asde ‘Pej aisrzr ‘Prr| wide ‘Pe wnyopnong (sddo q}IM) yo ‘dg ‘Po (s8do yyIM) at i$ ‘Po 44 Hex ‘Pr widr ‘Po! wsd4‘Pse] asder Ose} arsdr ‘Po a‘ SEo “PL “ald :a es ie en oe +(eIsapomy 0} a1eZ) eoLIpY TeI}WAD UI wnmasIv1419 Jo soloads o11;eduIAS XIS Jo pouTUIeXe SUsUTTOads Jo SIaquINuU pUe ejep [eoTso[oOIg Jo AreUIUUNS “ds visajstyavag uo wnjoduanyy 1 'hz ‘Po u a < aunt AeW “Ie “‘qad uel salodds I AAV] CIRIACREMINI OF THE ETHIOPIAN REGION 13 times in the evolutionary history of the group. In the genera Phacopteron and Pseudophacopteron Rs and M,,, make punctiform contact but on the structure of the head and thorax these genera are naturally placed near the Pauropsyllini in the Carsidaridae (Loginova, 1972). Also several other genera in the Carsidaridae, e.g. Carsidara and Mesohomotoma, possess an ym crossvein but in these genera it is of a different nature to that found in the Ciriacreminae. An vm crossvein is present in the New Zealand genus Anomalopsylla which is currently placed in the Spondy- liaspinae (Vondracek, 1963). These examples are all from outside the family Psyllidae. Where the 7m crossvein occurs within the Psyllidae its presence is used here to indicate monophyly and all species possessing this feature are placed in the Ciriacreminae. This usage must remain speculative until the functional significance of the presence of an 7m crossvein is understood and other corroborative syna- pomorphic characters are found. Other authors (Klimaszewski, 1963; Bekker- Migdisova, 1973) use the apparently bipartite male proctiger to diagnose the group further but the latter author considers this feature to be primitive. If this is the case then it should not be given much weight when considering phylogeny. In most Ciriacremini the male proctiger is subdivided into a broader basal part, which sometimes bears lateral lobes or expansions, and a narrower apical part which is often relatively long. It is not clearly bisegmented, as in Phytolyma or Mesohomotoma, and may not be a primitive feature. This kind of proctiger structure occurs widely in the Psylloidea but is not often as clearly developed as in the Ciriacremini. Apart from the Ethiopian species, which are considered here, three other species are currently placed in the Ciriacremini and are distributed in the New World. Pamisopelma quadrigibbiceps Enderlein, 1910a, was described from east-central Argentina, P. (Connectopelma) conifrons Sulc, 1914, from the other side of the Andes in central Chile, and Ceriacremum setosum Crawford, 1914, from Central America. In Panisopelma and Connectopelma the anteoccipital lobes are absent but the vertex, on either side of the median suture, is swollen to form a pair of lobes or humps, the basal spine of the hind tibia is very weak or absent and the male subgenital plate lacks hypovalves. If these two species share a common ancestry with the Ethiopian Ciriacremini, and this is open to doubt, their divergence from the Old World forms was early in the history of the group. The third New World species, Ceriacremum setosum, more closely resembles the Old World forms; the anteoccipital lobes are present and the vertex is unmodified, the basal spine of the hind tibia is strongly developed and the form of the male proctiger is similar to that of Kleiniella medleri sp. n. and Ciriacremum capense Enderlein. KEY TO GENERA OF CIRIACREMINI I Vertex on either side of median suture swollen to form two clearly protruding lobes or humps (Text-fig. 28), tanteoccipital lobes absent; basal spine of hind tibia very weak or absent. South America. PANISOPELMA and CONNECTOPELMA — Vertex not or hardly swollen on either side of median suture, anteoccipital lobes present (Text-figs 29-33); basal spine of hind tibia well developed. Ethiopian Region, Central America ; : : : : : : - 5 14 D.. HOLLIS 2 Fore wing without a pterostigma, Cu curves forwards to touch M (Text-fig. 59) PALMAPENNA @. 14) — Fore wing with a pterostigma, Cu not touching M (Text-figs 51-56) ; ; 3 3 Veins of fore wing with hairs which are more than twice as long as width of veins (Text-figs 51-58); hairs on antennal flagellar segments longer than width of pedicel (Text-fig. 5); fore wing usually patterned (Text-figs 51-58), pterostigma usually sessile but if pedunculate then wings clearly pigmented (Text-figs 57-58); male subgenital plate without hypovalves (Text-figs 96, 102, 108) KLEINIELLA (p. 15) — Hairs on veins of fore wing usually short (Text-fig. 60), rarely longer than width of veins; hairs on antennal flagellar segments not longer than width of pedical (Text-fig. 6); fore wing usually hyaline (Text-fig. 61), pterostigma usually pedunculate (Text-fig. 62) but if sessile then wing hyaline; male subgenital plate usually with hypovalves (Text-figs 114, 126, 138) ; ; CIRIACREMUM (p. 24) PALMAPENNA &en. n. Type-species: Palmapenna hymenostegioides sp. n. DEscRIPTION. Medium size, head width 0-64-0-783mm. Integument shiny, with a moderately dense covering of long hairs. Head wider than pronotum, with vertex and genal cones in same plane as axis of body; vertex almost flat; genal cones not distinctly separated from vertex, completely covering frons dorsally; antenna filiform, flagellum two to two and a half times longer than maximum width of head, sparsely covered with short hairs; ultimate rostral segment slightly shorter than third antennal segment. Thorax, in profile, weakly convex, mesoscutellum raised medially into a blunt tubercle; propleurites subequal, both in contact with pronotum dorsally (Text-fig. 1). Fore wing rounded triangular, broadest distally; pterostigma absent, costal and anal breaks present, veins sparsely covered with long hairs on dorsal surface; vm crossvein present, long, M stem also in contact with Rs proximally and with Cw, distally, points where veins make contact or branch often marked by a sharp callus dorsally; costal setae of hind wing divided into two groups. Fore femur with a few longer setae antero-ventrally, fore tibia with four to five pairs of longer setae antero-dorsally; apical spurs of hind tibia not arranged to form a distinct ventral group. Male subgenital plate without hypovalves; proctiger with short, narrow apical part and broad basal part which bears a pair of backwardly pointing lateral lobes. Ovipositor short, simple, conical. Discussion. The form of the fore wing in this genus is highly derived and it is very difficult to assess its ancestral type. The pterostigma is completely lacking, M curves in an unusual manner to make contact with both Rs and Cu,, and the overall wing shape is unique among known psyllids. However, the vm crossvein is present and well developed and the veins bear long hairs similar to species of Kleiniella. The genal cones are not completely separated from the vertex by the transverse sutures but in most other respects Palmapenna resembles other members of the Ciriacremini and the genus is provisionally placed in this tribe. Palmapenna hymenostegioides sp. n. (Text-figs I, 29, 59, 89, 93-95, 183, 184) DESCRIPTION. Colouration. Overall body colour brown with ochraceous and orange mottling. Head ochraceous above, brown below; antenna with scape and pedicel brown, segments three to nine ochraceous with darkened tips, segment ten brown; dorsum of thorax with broad longitudinal ochraceous band, mesoscutellum brown antero-medially, pro- and CIRIACREMINI OF THE ETHIOPIAN REGION 15 mesopleurae brown, metapleurites ochraceous-green; fore wing hyaline with heavy brown pattern in distal third and in posterior half giving wing a paddle-like appearance (Text-fig. 59), veins alternately ochraceous and brown; fore leg brown with ochraceous mottling, middle and hind legs ochraceous with orange mottling; abdomen dark brown above, mottled ochraceous- orange ventrally. Structure. g. Head about one and a half times longer than wide, sparsely haired dorsally, densely haired ventrally; post-orbital ridges absent, anteoccipital lobes reduced to small tubercles; genal cones well developed (Text-fig. 29), conical; antenna moderately long, flagellum 2:08-2:28 times longer than width of head, relative lengths of flagellar segments from base to apex —1'0 : 1-3 :1°4:1'7:2°2:2°4:1°'6:0°8; ultimate rostral segment 0o-85—0-96 times as long as third antennal segment. Pronotum short, narrower than maximum width of head; propleural suture diagonal, episternum narrow above (Text-fig. 1), epimeron narrow below; fore wing venation and spinule arrangement as in Text-fig. 59; hind tibia with four or five spurs apically not arranged to form a ventral group. Proctiger (Text-fig. 93) with very short and narrow apical part, basal part broad with a pair of backwardly directed lateral expansions; paramere (Text-figs 93, 94) elongate conical, slightly longer than proctiger, inner surface evenly and densely covered with short curved setae and with a short thick spine postero-medially; subgenital plate (Text-fig. 93) without hypovalves. ®. Antennal flagellum 2:10-2:43 times longer than width of head, relative lengths of flagellar segments from base to apex—I-0: 1-4 :1°3:1°6:2*1 :2°3:1°5:0°7; ultimate rostral segment 0-81—0-93 times as long as third antennal segment. Ovipositor (Text-fig. 183) short, proctiger 0-69—0-75 times as long as hind tibia. Measurements (+0-01 mm, based on 5 g and 7 9). Maximum width of head, ¢ 0-64-0:72, 9 0:63-0°78; length of ultimate rostral segment, J 0-11, 9 o-11-0-13; length of antennal flagellum, 3 1°46-1°51, 9 1-46-1°71; length of fore wing, J 2:00-2:22, 2 2-16~2-50; length of hind tibia, 3g 0°69-0-80, 9 0-71-0°85; length of 2 proctiger, 0-52-0-61. Host PLANT. Hymenostegia laxiflora (Bentham) Harms. A long series of adults was collected from a group of young seedlings of the host plant in an old coffee forest that was reverting to its natural state. Holotype g, ANGora: Salazar, I.I.A.A., 9-15.iii.1972, beaten from Hymenostegia laxiflora (BM Southern Africa Expedition) (BMNH, London), dry mounted. Paratypes. ANGOLA: 125 J, 1549, same data as holotype; dry and slide mounted, in 80% ethanol; deposited in BMNH, London; MNHU, Berlin; MRAC, Tervuren; USNM, Washington; ZI, Leningrad. KLEINIELLA Aulmann, 1912 Kleiniella Aulmann, 1912b : 100. Type-species: Kleiniella superba Aulmann, by monotypy. Desmiostigma Enderlein, 1927 :240. Type-species: Desmiostigma jassina Enderlein, by monotypy. Syn. n. Syndesmophlebia Vondracek, 1963 : 272. Type-species: Syndesmophlebia oblongata Vondracek, by monotypy. Syn. n. DEscripTion. Large to medium size, head width 0:79-1:46 mm. Integument matt with a sparse covering of long hairs. Head as wide as or wider than pronotum and slightly depressed from axis of body; each lateral half of vertex with a median concavity; genal cones poorly or well developed, distinctly separated from vertex by transverse sutures; anteoccipital lobes present; antenna filiform, 2-0—4-5 times longer than width of head, flagellar segments bearing long hairs; ultimate rostral segment short, 0:25-0:89 times as long as third antennal segment and 1-2—1-8 times longer than tenth antennalsegment. Thorax, in profile, convex; propleurites usually subequal and both in contact with pronotum dorsally. Fore wing parallelogrammatic or ellipsoid, normally hyaline with a clearly developed brown pattern, rarely uniformly 16 D. HOLLIS brown; pterostigma sessile or pedunculate; veins bearing long hairs on dorsal surface, at least in proximal half of wing; costal and anal breaks present; Rs in punctiform contact with M,,, or there is a short vm crossvein; costal setae of hind wing tending not to be arranged into groups. Fore femur with a few longer setae antero-ventrally, fore tibia with four to six pairs of longer setae dorsally; hind tibia with apical spurs usually arranged evenly around apex, rarely grouped to form a ventral comb. Male subgenital plate without hypovalves; proctiger with broad basal part and short, narrow apical part. Female ovipositor conical, proctiger o-81—1-80 times longer than hind tibia. Host PLANTS. Unknown. DISTRIBUTION. Congo basin and montane forest regions of tropical Africa from Tanzania to Sierra Leone (Map 2). Discussion. It is suggested here that the genus Kleiniella contains the following six species: K. superba Aulmann, Desmiostigma jassina Enderlein, K. congoensis sp. n., Phacosema guineensis Aulmann, Syndesmophlebia oblongata Vondracek and K. medleri sp. n. According to Enderlein (1927) D. jassina differs from K. superba because of its smaller genal cones, the absence of hairs on the veins of the fore wing and the overall shape of the fore wing. In fact the wing veins in jassina do bear a double row of hairs on the dorsal surface and, in the proximal half of the wing, these hairs are long and similar to those found in superba. The wing shape in jassina is more like the parallelogrammatic wing of superba than the ellipsoid wing of S. oblongata and K. medlert. On this basis Desmiostigma is synonymized with Kleiniella. K. guineensis (Aulmann), originally placed in the genus Phacosema but more naturally placed here, has an intermediate pterostigma condition for the genus. In oblongata and medleri the pterostigma is triangular and pedunculate, in guineensis it is triangular and sessile, and in superba, jassina and congoensis it is trapezoid and sessile. The synonymy of Syndesmophlebia with Kleiniella is more difficult to accept. Together ‘with medleri, oblongata is rather intermediate between Kleiniella and Ciriacremum. The wing veins, antennal segments, body and legs bear long hairs, the ym crossvein is punctiform or short, and the male subgenital plate lacks hypovalves, as in Kleiniella. On the other hand the pterostigma is pedunculate, the fore wing is ellipsoid and, in medleri, the apical spurs of the hind tibia are arranged to form a distinct ventral group, features these two species have in common with Civiacremum species. The states of expression and functional significance of these characters are not at all clear. However, in Text-fig. 20, using nine characters, a highly speculative phylogeny displaying the relationships among K/leiniella species and between Kleiniella and Ciriacremum is given. The nine characters used and their states of expression are as follows. 1. Wing pattern: absent — plesiomorphic, present —- apomorphic; Text-figs 58, 60. 2. Wing vein setae: short — plesiomorphic, long — apomorphic; Text-figs 57, 61. 3. Antennal setae: short — plesiomorphic, long — apomorphic; Text-figs 5, 6. 4. Pterostigma: pedunculate — plesiomorphic, sessile - apomorphic; Text-figs 52, 62. CIRIACREMINI OF THE ETHIOPIAN REGION 17 - la Ciriacremum a 2 wn en“ = .o) 0 £2905 2 mae 6 a2oonr Oo 4 4 6 5 9 8 5 3 2 | Character states: [] plesiomorphic @ apomorphic © HBR et edlcr © ancestral species Fic. 20. Dendrogram of projected phylogeny within the Ciriacremini (for further explanation see text, p. 16). 18 D. HOLLIS 5. Costal setae of hind wing: ungrouped — plesiomorphic, grouped — apomorphic; Text-figs 84, go. 6. Hind tibial spurs: ungrouped ventrally — plesiomorphic, grouped ventrally — apomorphic; Text-figs 13, 14. Body and leg setae: long — plesiomorphic, short - apomorphic; Text-figs 11, 12. ym crossvein: punctiform — plesiomorphic, long — apomorphic; Text figs 58, 60. g. Male hypovalves: absent — plesiomorphic, present —- apomorphic; Text figs 96, 138. eo It is postulated that the primitive Ciriacremini stock had all nine characters in the plesiomorphic condition. In the present day fauna where the characters are expressed in the apomorphic condition they are regarded as having been uniquely derived, although there are exceptions, e.g. C. funestum sp. n. has long hairs on the veins of the fore wing and a sessile pterostigma (see discussion of this species on P- 37). Key TO SPECIES OF KLEINIELLA I Genal cones poorly developed, only scones tie by weakly swollen areas of genae (Text-fig. 31) . ‘ 2 — Genal cones well develgeed: aed or Getanely She (Text- figs 30, 32, 3 i 2 Pterostigma broadly sessile, trapezoidal (Text-fig. 52); male subgenital plate wrichaue hypovalves (Text-figs 96, 99) . : 3 — Pterostigma narrowly sessile, triangular (Text: a. 82): male subeenital plate with well developed hypovalves (Text-fig. 129) Ciriacremum funestum sp. n. (p. 37) 3 Larger species, head width 0.96-1.12 mm; fore wing narrower, 2.40—2.72 times longer than wide, without spinules in costal cell or in proximal part of second radial cell (Text-fig. 52); apex of aedeagus as in Text-fig. 98; 2 proctiger longer than hind tibia with downcurved apex (Text-fig. 187) : : . jassina (Enderlein) (p. 19) — Smaller species, head with 0-85—0-93 mm; fore wing broader, 1-9—2-2 times longer than wide, spinules clearly present in costal cell and in proximal part of second radial cell (Text-fig. 53); apex of aedeagus as in Text-fig. 101; 2 proctiger shorter than hind tibia, apical part not downcurved (Text-fig. 188) congoensis sp. n. (p. 20) 4 Pterostigma of fore wing pedunculate (Text-figs 57, 58) : : : : : 5 — Pterostigma of fore wing sessile (Text-figs 51, 54-56) . : : : 6 5 Pterostigma short, about two and a half times longer than wide (Text: fig. 57); genal cones rounded rectangular (Text-fig. 32) . é oblongata (Vondracek) (p. 22) — Pterostigma elongate, about eleven times longer than wide (Text-fig. 58); genal cones obtusely conical (Text-fig. 33) : . medleri sp. n. (p. 23) 6 Pterostigma of fore wing broadly Papeete (ext. -fig. 51) superba Aulmann (p. 18) — Pterostigma of fore wing triangular or very narrowly trapezoidal (Text-figs 54-56) guineensis (Aulmann) (p. 21) Kleiniella superba Aulmann, 1912 (Text figs 13, 51, 83, 185) Kleiniella superba Aulmann, 19120 : rot, figs 1-4. LECTOTYPE 9, Tanzania: ‘D. O. Afrika, Mkulumuri b. Amani, xii. 05, Dr Schréders. G’ (MNHU, Berlin), here designated [examined]. Kleiniella superba Aulmann; Aulmann, 1913 : 79. DEscRIPTiON. Colouration. Overall body colour olive-green; fore wing pattern as in Aulmann’s original figure, CIRIACREMINI OF THE ETHIOPIAN REGION 19 Structure. 4 unknown. Q. Head as wide as pronotum; genal cones well developed, obtusely conical. Fore wing (Text-fig. 51 and original figure) parallelogrammatic, pterostigma broadly trapezoidal; Rs and M,,.2 in punctiform contact, first cubital cell elongate; costal setae of hind wing forming a continuous row (Text-fig. 83); apex of hind tibia with three inner and two outer spurs. Ovipositor conical (Text-fig. 185), proctiger relatively short, 1-30 times longer than hind tibia. Measurements (+0-01 mm, based on single incomplete 9). Maximum width of head, 9 1-18; length of ultimate rostral segment, 9 0-13; length of hind tibia, 9 0-99; length of 9 proctiger, 1°29. Host PLANT. Unknown. MATERIAL EXAMINED. Lectotype and one paralectotype ?, same data and depository as lectotype. Discussion. Aulmann’s type-series of two females are the only specimens of this species available for study and both are in a poor state of preservation. A permanent preparation has been made of the remnants of the lectotype. The antennae are missing (they were not mentioned in the original description) and only part of one fore wing remains (Text-fig. 51). However, from this remnant material and Aulmann’s original description it is clear that superba is distinct from other species in the genus, having the most derived form of fore wing shape and venation. Kleiniella jassina (Enderlein, 1927) comb. n. (Text-figs 3, 5, II, 31, 52, 86, 96-98, 187) Desmiostigma jassina Enderlein, 1927 : 241, fig. 1. Holotype 9, CAMEROUN: ‘Kamerun’ (IPK, Eberswalde) [unavailable for study]. DESCRIPTION. Colouration. Overall body colour light green with orange-brown markings. Lateral ocelli orange; thoracic tergites brown medially, mesopleurites brown; legs green with brown tarsal segments; fore wing pattern as in Text-fig. 52; abdomen orange-brown dorsally and ventrally. Structure. g. Head wider than pronotum; genal cones very poorly developed (Text-fig. 31) ; antennal flagellum 2:09-2:21 times longer than maximum width of head, relative lengths of flagellar segments from base to apex—1I-0:0°9: I'l :1°5:1°9:1°9: 1:1 :0°5; ultimate rostral segment 0-74—0-81 times as long as third antennal segment. Fore wing parallelogram- matic, 2:40-2:52 times longer than wide, spinule arrangement as in Text-fig. 52; pterostigma broadly trapezoidal, dorsal surface of veins with a double row of long hairs in proximal half of wing, in distal half the hairs are shorter; Rs makes punctiform contact with M,,. or there is a very short ym crossvein; Cu stem about two and a half times longer than M+Cuwu stem; first cubital cell elongate, about twice as long as wide; apex of hind tibia with a pair of spurs on each side and a single ventral spur. Proctiger (Text-fig. 96) with oval lateral expansions in basal part and a short narrow apical part; paramere (Text-figs 96, 97) thumb-like, on inner surface with a large group of downwardly directed setae in apical part and a small median ridge in basal part; distal segment of aedeagus as in Text-fig. 98. 9. Antennal flagellum 2:07-2:09 times longer than width of head, relative lengths of flagellar _ segments from base to apex — I'0 : 1:0 : I°2 : I°4 : 1°9 : I°9: 1:0: 0'5; ultimate rostral segment 0°82 times as long as third antennal segment. Fore wing 2:49-2:65 times longer than wide. Ovipositor elongate conical (Text-fig. 187), proctiger 1:16-1:29 times longer than hind tibia, with downcurved apex. 20 D. HOLLIS Measurements (-+0-01 mm, based on 6 g and 3 9). Maximum width of head, ¢ 0:96-1:12, Q 1-02-1-12; length of antennal flagellum, $ 2:06-2:30, 2 2-13-2-26; length of ultimate rostral segment, ¢ 0-17-0°18, 9 0-18-0-19; length of fore wing, J 3:00-3:63, 2 3-63-4:07; maximum width of fore wing, g 1:25-1:46, 9 1:37-1:63; length of hind tibia, J 1-08—1-23, 9 1-:14-1-36; length of 2 proctiger, 1-48-1-60. Host PLANT. Unknown. MATERIAL EXAMINED. ZAIRE: I 4, Stanleyville, iv. 1928 (A. Collart); 3 3, 3 9, Yangambi, 0-47N—24:24E, x. 1956 (N. L. H. Krauss); 1 9, Elizabethville, xi. 1959, at light (Ch. Seydel). ANGOLA: 36,12, Dundo, 4—29.vill.1953, light trap (Luna). Discussion. K. jassina is closely related to the following species and the two are regarded here as sister species. As both species have long setae on the antennal flagellum and the veins of the fore wing, a parallelogrammatic fore wing and a broadly trapezoid pterostigma, they are regarded here as more closely related to K. superba than the other three species of Kleiniella. K.jassina and K. congoensis clearly differ from K. superba in having poorly developed genal cones. A number of cecidomyid fly larvae, probably of the genus Endopsylla (identified by K. M. Harris of the Commonwealth Institute of Entomology, London) were obtained from the abdomens of the series of specimens from Yangambi; Zaire. Kleiniella congoensis sp. n. (Text-figs 53, 87, gg-1o1, 188) DESCRIPTION. Colourvation. Similar to jassina but with darker orange markings; fore wing pattern as in Text-fig. 53. Structure. . Similar to jassina but smaller. Antenna missing from holotype; ultimate rostral segment short. Fore wing broader than in jassina, 1:97 times longer than wide, with a moderately dense covering of spinules (Text-fig. 53), costal cell and proximal part of second radial cell with spinules; Cu stem about twice as long as M-+Cuw stem, first cubital cell shorter than in jassina, only one and a half times longer than wide. Genitalia similar to jassina; paramere (Text-figs 99, 100) thumb-like, on inner surface with short conical setae in apical two-thirds, without tubercle or ridges; distal segment of aedeagus relatively simple (Text- fig. Ior). Q. Slightly larger than g. Antennal flagellum 2-03 times longer than width of head, relative lengths of flagellar segments from base to apex —1:'0 ! 1:0 : I'l :1°3 :1°9:2°0:1'53: 0:6; ultimate rostral segment 0-89 times as long as third antennal segment and 1-45 times longer than tenth antennal segment. Fore wing 2:11 times longer than wide. Ovipositor short (Text-fig. 188), proctiger with straight apex, 0-81 times as long as hind tibia. Measurements (+0-o1 mm, based on 1 gf and1Q). Maximum width of head, g 0°85, 9 0°93; length of antennal flagellum, 2 1-88; length of ultimate rostral segment, ¢ 0:15, 9 0-16; length of fore wing, f 2:76, 9 3:11; maximum width of fore wing, g 1-40, 9 1:48; length of hind tibia, 3 0°97, 2 1:03; length of 9 proctiger, 0-84. Host PLANT. Unknown. Holotype g, ZAIRE: Yangambi, 0-47N—24:24E, x. 1956 (N. L. H. Krauss) (BMNH, London); slide mounted. CIRIACREMINI OF THE ETHIOPIAN REGION 21 Paratypes. ZAIRE: 2, same data as holotype. NIGERIA: 1 9, SE. State, Oban, 7.iv.1975 (J. IT. Medler) (BMNH, London), slide and dry mounted. Discussion. Obviously K. congoensis sp. n. is closely related to K. jassina but shows significant differences in the structure of the fore wing and genitalia of both sexes. The type-series was apparently collected together with specimens of jassina. Unfortunately no indication of host plant data was given by the collector and host preference may be the isolating factor between the two species. Kleiniella guineensis (Aulmann, 1912) comb. n. (Text-figs 30, 54, 85, 102-104) Phacosema guineensis Aulmann, 1912@ : 35, figs 1-6. Holotype g, EQUATORIAL GUINEA: ‘Span. Guinea, Nkolentangan, xi. 07—v. 08, G. Tessmann S.G.’ (MNHU, Berlin) [examined]. Udamostigma guineensis (Aulmann) Aulmann, 1913 : 81. DESCRIPTION. Colouvation. Overall body colour brown with ochraceous markings; vertex brown medially, ochraceous laterally; genal cones brown; dorsum of thorax brown, pronotum ochraceous laterally, mesoscutum with poorly developed longitudinal ochraceous markings, meso- and metascutellum brown medially and ochraceous laterally, propleurites ochraceous, mesopleurites brown; fore wing hyaline with brown pattern as in Text-fig. 54; legs mainly ochraceous, apex of fore femur, dorsally, and fore tarsal segments brown, mid femur and tarsus brown, hind femur, basally, and hind tarsus brown; abdomen brown dorsally, ochraceous ventrally. Structure. g. Head wider than pronotum; genal cones well developed (Text-fig. 30), obtusely conical. Fore wing ellipsoid, 2-45 times longer than wide, spinule arrangement as in Text-fig. 54; pterostigma triangular; Cu stem slightly more than one and a half times longer than M-+-Cu stem, first cubital cell twice as long as wide. Apex of hind tibia with a pair of spurs on each side and a single ventral spur. Proctiger (Text-fig. 102) with oval lateral expansions in basal part and a short narrow apical part; paramere (Text-figs 102, 103) thumb- like, on inner surface with a few downwardly directed setae in apical part and a subapical tooth; distal segment of aedeagus as in Text-fig. 104. Unknown. Measurements (+0-01 mm, based on1 g). Maximum width of head, J 0-79; length of ultimate rostral segment, ¢ 0-12; length of fore wing, g 3:21; maximum width of fore wing, 3 1°31; length of hind tibia, ¢ 0-81. MATERIAL EXAMINED. Holotype and one further g specimen from ‘Westafrika, Uelleburg, vi—viii. 08’ Tessmann S. G.) (MNHU, Berlin). Discussion. K. guineensis differs from both K. superba and K. jassina in having an ellipsoid fore wing and a triangular pterostigma. The sessile form of the latter also separates guineensis from K. oblongata and K. medleri. Apart from the material mentioned above two female specimens have~ been examined which show slight but obvious differences from the type-series. As I have no clear idea of the extent to which K. guineensis varies I cannot assign these specimens to that or any other named species. A brief indication of their main characteristics is given below. 22 D. HOLLIS Kleiniella sp. 1, near guineensis (Text-fig. 55) Q. Similar to guineensis. Antennal flagellum 2-43 times longer than width of head, relative lengths of flagellar segments from base to apex—1-:0 : 1-0: 1-1 : 1:4 :1°8: 1:8 : 1:0 :0°5; ultimate rostral segment short, 0-57 times as long as third antennal segment. Fore wing (Text-fig. 55) 2:28 times longer than wide, with less well developed pattern than guineensis and broader pterostigma; Cu stem two and a half times as long as M+Cu stem, Cu, more steeply parabolic and first cubital cell slightly less than twice as long as wide. Proctiger relatively long, 1:77 times longer than hind tibia. Gg unknown. Host plant unknown. ~ Measurements (-+-0-01 mm, based on 1 9). Maximum width of head, 9 0-86; length of antennal flagellum, 9 2:09; length of ultimate rostral segment, 2 0-12; length of fore wing, 9 3-15; maximum width of fore wing, 9 1-38; length of hind tibia, 9 0-74; length of 9 proctiger, 1-31. MATERIAL EXAMINED. NIGERIA: I 9, SE. State, Obudu CR, 21.iii.1971 (J. T. Medler) (BMNH, London). Kleiniella sp. 2, near guineensis (Text-fig. 56) Q. Similar to guineensis and preceding species. Genal cones rather more robust (Text- fig. 56); antennal flagellum 2-43 times longer than width of head, relative lengths of flagellar segments from base to apex—1:0:1:0:1'0:1'2:1'6:1°'5 ;0°8:0°3; ultimate rostral segment shorter, 0-45 times as long as third antennal segment. Fore wing slightly more parallelogrammatic (Text-fig. 56), 2°55 times longer than wide, pterostigma broadly triangular, Cu stem almost three times longer than M-+Cuw stem, first cubital cell about twice as long as wide. Proctiger shorter, 1-48 times longer than hind tibia. g unknown. Host plant unknown. Measurements (+0:01 mm, based on 1). Maximum width of head, 9 1-02; length of antennal flagellum, 9 2:48; length of ultimate rostral segment, 9 0-13; length of fore wing, 9 3-83; maximum width of fore wing, 9 1-38; length of hind tibia, 9 1-00; length of 2 proctiger, 1-48. MATERIAL EXAMINED. UGANDA: I 9, Ruwenzori Range, Bwamba Pass (west side), xii. 1934-1. 1935, 5500’—7500’ [1680-2290 m] (Ff. W. Edwards) (BMNH, London). Kleiniella oblongata (Vondracek, 1963) comb. n. (Text-figs 32, 57, 84, 105-107) Syndesmophlebia oblongata Vondracek, 1963 : 273, figs 31-45. Holotype g, ZarRE: ‘Rutshuru, 27.V.1936 (L. Lippens)’ (MRAC, Tervuren) [examined]. DESCRIPTION (for more details see original description). ¢. Head wider than pronotum; genal cones well developed, rounded rectangular from above (Text-fig. 32); ultimate rostral segment short. Fore wing ellipsoid, 2:54 times longer than wide, spinule arrangement as in Text-fig. 57; pterostigma pedunculate, short; Rs and M,,, in punctiform contact; first cubital cell elongate, more than twice as long as wide; Cu stem about three times longer than M-+Cu stem; costal setae of hind wing forming a continuous row (Text-fig. 84); apex of hind tibia with three inner and two outerspurs. Proctiger (Text-fig. 105) with oval lateral expansions CIRIACREMINI OF THE ETHIOPIAN REGION 23 in basal part and a short narrow apical part; paramere (Text-fig. 106) thumb-like, on inner surface with a subapical tooth and a number of downwardly directed setae in apical half, in basal half with a group of anteriorly directed setae; distal segment of aedeagus as in Text-fig. 107. Q. Antennal flagellum 2-30 times longer than width of head, relative lengths of flagellar segments from base to apex — 1-0 : 1:0 : II : 1-3 : 1-8 : 1°8 : 0-6 : 0-3; ultimate rostral segment 0°58 times as long as third antennal segment. Fore wing 2-57 times longer than wide. Proctiger relatively long, 1-80 times longer than hind tibia. Measurements (+0-01 mm, based on 1 gf and 1Q). Maximum width of head, 3 1-36, 9 1-46; length of antennal flagellum, 9 3-37; length of ultimate rostral segment, J 0-20, 2 0-21; length of fore wing, ¢ 5:10, 9 5:70; maximum width of fore wing, ¢ 2-00, 9 2-22; length of hind tibia, 3 1°13, 9 1°24; length of 2 proctiger, 2-22. Host PLANT. Unknown. MATERIAL EXAMINED. Holotype and 1 9 paratype from ZAIRE: Luebo, viii. 1921 (Lt. Ghesquiére) (MRAC, Tervuren). Both specimens had been dissected and mounted on several slides; some remounts were necessary. Discussion. Unlike the three previous species K. oblongata has a pedunculate pterostigma and an ellipsoid fore wing, features it has in common with K. medleri sp. n. and most species of Civiacremum. It differs from medleri in the shape of the genal cones, the short pterostigma and the form of the male genitalia. It is placed in Kleiniella because of the long hairs on the veins of the fore wing, antennal flagellum and fore tibia, the absence of a ventral group of spurs at the apex of the hind tibia and the absence of hypovalves on the male subgenital plate. Kleiniella medleri sp. n. (Text-figs 33, 58, 88, 108-110, 186) DESCRIPTION. Colouvation. Overall body colour mid brown with light green or ochraceous markings; vertex green with central brown area and orange ocelli; genal cones green with reddish brown apices; antenna ochraceous with darkened area at joints of segments, segments nine and ten brown; pronotum brown anteriorly, green posteriorly and laterally; propleurites green; mesopraescutum brown with green lateral and hind margins and median longitudinal ochraceous stripe; mesoscutum brown with median greenish area, with a proximo-lateral ochraceous stripe and a disto-lateral green patch on each side; mesoscutellum brown with green antero-lateral areas; metascutellum green; parapteron brown dorsally, ochraceous-green ventrally; fore wing pattern as in Text-fig. 58; legs brown, hind tibia ochraceous; abdomen reddish brown dorsally, green dorso-laterally, dark brown ventro-laterally and yellow ventrally; 6 proctiger with reddish apical part. Structure. g. Head wider than pronotum; genal cones well developed, conical with rounded apices (Text-fig. 33); antennal flagellum relatively long, 4:23-4:48 times longer than width of head, relative lengths of flagellar segments from base to apex — 1°0 :0°9:1°0:1°4:1°4:1°8:0°5: 0-2; ultimate rostral segment relatively short, 0-28—o-31 times.as long as third antennal segment. Fore wing ellipsoid (Text-fig. 58), 2:36-2:48 times longer than wide; pterostigma pedunculate and very long, continuing almost to point where Rs meets costa; Cu stem almost twice as long as M-+Cu stem; Cw, steeply parabolic and first cubital cell about 1-3 times longer than wide; costal setae of hind wing arranged into two groups (Text-fig. 88); apex of hind tibia with one inner and one outer spur and a ventral group of two or three spurs. Proctiger (Text-fig. 108) with triangular wing-like expansions on basal part and a short narrow apical part; paramere 24 D. HOLLIS (Text-figs 108, 109) thumb-like, on inner surface with a postero-apical ridge which ends in an apical tooth, apical part with a vertical row of five antero-ventrally directed setae, basal part with a large group of posteriorly directed setae; distal segment of aedeagus as in Text-fig. 110. Q. Larger than g. Antennal flagellum 4:37-4:43 times longer than width of head, relative lengths of flagellar segments from base to apex—I-0 : 0-9 :1°0 :1°3 :1'4 : 1:8 :0°5 :0°2; ultimate rostral segment 0-25-0-30 times as long as third antennal segment. Fore wing 2°36-2°43 times longer than wide. Proctiger moderately long (Text-fig. 186), 1:49-1:60 times longer than hind tibia. Measurements (+0-01 mm, based on 7 gf and 6 9). Maximum width of head, ¢ 1:01-1:19, Q 1:03-1:26; length of antennal flagellum, $ 4-41-5-28, 9 4:55-5:58; length of ultimate rostral segment, gf 0:15-0:18; 2 0-16—0-18; length of fore wing, g 2:92-3:42, 9 3:12-3:69; maximum width of fore wing, g 1:18-1:40 9 1-32-1'52; length of hind tibia, ¢ 0-78-0-95, 2 0-81-1-00; length of 9 proctiger, 1-23-1-49. Host PLANT. Unknown. Holotype g, NicEriA: W. State, Ile-Ife, 25.iii.1969 (J. T. Medler) (BMNH, London). Dry mounted. Paratypes. NIGERIA: 2 g, 4 9, same data as holotype; 1 g, 28.ii.1970; I d, 19.11.1970; I 9, 30.Vi. 1970; 2\¢, 20.X11.1970; 4 gi, 26.11.1071; Ig, 9-1.1073; Dae Akure, x. 1974; 1 9 NW. State, Badeggi RRS, 19.ili.1972; 2 g, 14.xii.1974; I 6, Abuja, 20.11i1.1972; I 5, 1 9, MW. State, Benin, 8.iv.1973; 4 3, 3 Q, I.iv.1975; I 9, nr Siluko, ii. 1973; 3 3d, 3 2, Udo FR, I1.iv.1975; 1 g, 1 9, W. State, Agbabu, 3.i11.1973; 1 ¢, BP. State, Takuni, 19.11.1973; 1 g, 1 9, SE. State, Ikom, 4. 1v.1075; 2 g, 1 9, Ibadan, 15.11.1969 (J. T. Medler); 8 3, 3 2, Ibadan, Moor Plantation, 23. ix.-2.x.1960, light trap (L. A. Mound); 1 g, 1 2, Zaria, Samaru, II.ix.1974, to light (J. C. Deeming). GHANA: 2 3, 2 2, Tafo, Cocoa Res. Inst., 1973, light trap (N. A. K. Akotoye). SIERRA LEONE: I 9, Niala, 15.xi.—8.xii.1930, at light (E. Hargreaves). (BMNH, London; ZI, Leningrad; NCI, Pretoria). Slide and dry mounted. Discussion. It is difficult to assign this species to a genus as it has features in common with both Kleiniella and Ciriacremum species. K. medleri sp. n. has long setae on the antennal flagellum, wing veins and legs, a short vm crossvein, and lacks hypovalves in the male, as in Kleiniella; but has a pedunculate pterostigma, a ventral group of spurs at the apex of the hind tibia, and grouped costal setae on the hind wing, as in Ciriacremum. On the basis that medleri sp. n. shares more derived features with Kleiniella species than Ciriacremum species it is arbitrarily placed in the former genus. CIRIACREMUM Enderlein, 1910 Civiacremum Enderlein, 1910b : 139. Type-species: Civiacremum filiverpatum Enderlein, by original designation. Ciriacvemum Enderlein; Aulmann, 1913 : 80. Ceriacremum Enderlein; Crawford, 1914 : 63, partim. [Incorrect subsequent spelling. ] Bunoparia Enderlein, 1926 : 397. Type-species: Civiacremum capillicorne Enderlein, by original designation. Syn. n. DESCRIPTION. Small to large size, head width 0:52-1:37 mm. Integument matt, with a moderate covering of short hairs. Head wider than pronotum and depressed from axis of body; each lateral half of vertex with a median concavity; genae from slightly swollen to having well CIRIACREMINI OF THE ETHIOPIAN REGION 25 developed cones, distinctly separated from vertex by transverse sutures; anteoccipital lobes present; antenna filiform, short or long, flagellum 1-2 to more than 5-5 times longer than width of head, flagellar segments bearing short hairs; ultimate rostralsegment short orlong. Thorax, in profile, convex; propleurites subequal and both in contact with pronotum dorsally; fore wing ellipsoid, sometimes a little elongated, usually hyaline, rarely with a marginal infuscation distally, sometimes with small darkened areas in the marginal cells distally; pterostigma usually pedunculate, rarely sessile; veins usually bear short hairs on dorsal surface, rarely these hairs are longer than the width of veins; long ym crossvein present; costal setae of hind wing clearly arranged into two groups; fore tibia without long setae or with only a single seta dorsally ; hind tibia with apical spurs arranged into lateral and ventral groups. Male subgenital plate mostly with a clearly developed pair of hypovalves, rarely fused to form a single structure, rarely without hypovalve development; proctiger usually with moderately narrow basal part and narrower apical part which is often as long as or longer than basal part. Female ovipositor conical, proctiger short or long, 1-0—2-7 times longer than hind tibia. Larva free-living psylloid type (Text-figs 24-27). DISTRIBUTION. West, central, east and southern Africa, in both forest and woodland areas (Maps 3-7); Central America. Host PLANTS. Data are known for eight of the 23 known species in the genus and all the host plants are from the family Caesalpiniaceae. Discussion. From the phylogeny postulated above (p. 16) it can be seen that Ciriacremum species are separated from Kleiniella on the basis of common possession in the former of three derived features: short body and leg setae, a long ym crossvein, and the presence of hypovalves on the male subgenital plate. They also differ from most of the species in Kleiniella, medleri being the exception, in having grouped costal setae on the hind wing and apical spurs of the hind tibia forming a distinct ventral group; two further derived features. One of the most interesting of these attributes is the presence of hypovalves on the male subgenital plate. Development of these structures is very rare within the Psylloidea and I know of them occurring only in two other groups, Mesohomotoma and related genera, and an undescribed genus distributed in tropical Africa which may be a member of the Spondyliaspidae. The function of these structures is not known; they could be used by the male to orientate himself appropriately during copulation or to induce the required response in the female by tactile or visual stimulus. A study of these structures in relation to psyllid reproductive behaviour would be very interesting and informative. Three species in the genus, filiverpatuwm capillicorne and capense, lack these _hypovalves and it is not clear whether this condition is primitive or the result of secondary loss. By a comparative study of the male genitalia within the genus some species groups are easily defined but the relationships of a few species cannot be clearly shown so a phylogeny for Ciriacremum is not postulated. Some of the more obvious species groups are: filiverpatum with pervatum, relatum and kleinielloides; bicaudatum with harteni, capeneri, carvalhoi, daubicatum, cabudiatum, tubacadium, jubernardioides and orientale; and africanum with megafricanum, vondraceki and possibly angolense. The type male of C. capillicorne could not be traced and no further material was available for study. As this species is the type-species of Bunoparia the synonymy given above must remain speculative. Enderlein separated Bunoparia from 26 D. HOLLIS Ciriacremum on the basis of the form of the genal cones. These structures are very variable interspecifically in the Ciriacremini and I do not think their form has any great significance when considering generic limits. | |) I2 KeEy TO SPECIES OF CIRIACREMUM Pterostigma sessile (Text-figs 63, 82). ; ‘ : j : : : - 2 Pterostigma pedunculate (Text-figs 64-81) : 3 Larger elongate species, head width 1-13 mm, aatennal aeecliend 4°27 eennes lledeee than width of head; fore wing 2-69 times longer than wide kleinielloides sp. n. (p. 32) Smaller, more robust species, head width o-78—-o-93 mm; antennal flagellum 2°50-3°10 times longer than width of head; fore wing 2-3—-2-5 times longer than wide .. : : funestum sp. n. (p. 37) Hairs on dorsal surface ats veins of fore wing more ‘than twice as long as width of veins (Text-fig. 58); hairs on segments of antennal flagellum longer than width of pedicel (Text-fig. 5) . : . Kleiniella medleri sp. n. (p. 23) Hairs on veins of fore wing Sock, not or hardly longer than width of veins (Text- fig. 60); hairs on antennal flagellum short, not longer than width of pedicel (Text-fig. 6) : : : : : : : : : : 4 Males. : ; : : 5 : : : : E : : : 5 Females . : : : E ; 26 Subgenital plate atea an on pair of eppeedaees ay terminally (Text-figs I11, 132) : 6 Subgenital plate with a pair of ees feria (Reet figs — 126, I 38) : 9 Proctiger with wing-like expansions laterally (Text-fig. 132) : 7 Proctiger without wing-like expansions laterally (Text-fig. 111) filiverpatum Enderlein (p. 29) Antenna very long, more than twice as long as length of body capillicorne Enderlein (p. 35) Antenna shorter, at most as long as body . 8 Genal cones short, from above wider than long (Text. fig. 49): fore wing with ion spinules (Text-fig. 81). Southern Africa : capense Enderlein (p. 35) Genal cones longer than wide; fore wing evenly and densely covered in spinules. Central America F : setosum Crawford Hypovalves fused to form a single eee (ext: figs 168, 170) tea sp. n. (p. 43) Hypovalves paired (Text-figs 114, 126, 138) : < : 10 Hypovalves longer than or at least as long as parameres (Text: figs 129, I 38) ; II Hypovalves shorter than parameres (Text-figs 114, 123, 172) : : 19 Membrane of fore wing densely and evenly covered with spinules (Text- fig. 75) : 12 Membrane of fore wing at least with areas on either side of veins devoid of spinules (Text-figs 68, 78) F : : 13 Antennal flagellum 1-6—2-2 tienes longer gen maximum width of read, julbernardioides sp. n. (p. 45) Antennal flagellum 1-:2-1:5 times longer than maximum width of head orientale sp. n. (p. 46) Antenna short, flagellum 1-9—2-3 times longer than width of head harteni sp. n. (p. 42) Antenna longer, flagellum 2-6—3-4 times longer than width of head ‘ : : 14 Genal cones poorly developed (Text-fig. 49); distal segment of aedeagus relatively short and robust with a long terminal tube (Text-fig. 180) vondraceki sp. n. (p. 51) Genal cones well developed (Text-figs 38-42) ; distal segment of aedeagus relatively long and thin, with a short terminal tube (Text-figs 141, 149) . - c ‘ 15 15 19 20 23 24 25 26 CIRIACREMINI OF THE ETHIOPIAN REGION Hypovalves with a small group of specialized setae subapically on dorsal surface (Text-fig. 161). : é carvalhoi sp. n. (p. Hypovalves without a group of specialized setae subapically but often with such a group medially on dorsal surface (Text-figs 138, 143) Third antennal segment 2-3—2-7 times longer than ultimate rostral segment; - distal segment of aedeagus with broad apex (Text-fig. 149) . cabudiatum sp. n. (p. Third antennal segment less than 2:2 times longer than ultimate rostral segment; apex of distal segment of aedeagus narrower (Text-figs 141, 145) Hypovalves very long and narrow, apical part no wider than apical part of proctiger, median group of specialized setae well developed (Text-fig. 143) daubicatum sp. n. (p. Hypovalves broader, apical part almost twice as wide as apical part of proctiger, median group of specialized setae poorly developed (Text-figs 138, 139, 150) Paramere with long and narrow apical part, on inner surface the postero-medial group of setae are restricted to an area immediately below the tooth (Text-fig. 140) ; hypovalves with inner Se ta row of setae relatively well developed (Text-fig. 16) ; : : ‘ bicaudatum Enderlein (p. Paramere with a very short narrow apical part, on inner surface the postero-medial group of setae are curved and extended to form a compound vertically arranged group (Text-fig. 151); hypovalves with inner postero-vertical row of setae poorly developed (Text-fig. 150) 2 : . tubacadium sp. n. (p. Antennal flagellum at most 2°5 times longer than maximum width of head Antennal flagellum at least 4-0 times longer than width of head . . Paramere broad, shorter than proctiger, with a longitudinally arranged group of sensory pegs on inner surface (Text-fig. 176); antennal flagellum not more than 2-2 times longer than width of head; ultimate rostral segment longer than length of third antennal segment : Paramere narrow, about as long as proctiger, without trace of specialized sensory pegs on inner surface (Text-fig. 136); antennal flagellum not less than 2-3 times longer than width of head; ultimate rostral segment much shorter than third antennal segment : : jilorense sp. n. (p. Ultimate rostral segment shorter, I*00-1°45 times longer than third antennal segment. : ‘ africanum Enderlein (p. Ultimate rostral segment longer, I 535-2: 75 ‘times longer than third antennal segment megafricanum sp. n. (p. Postero-basal patch of sensory hairs on inner surface of paramere present (Text- figs L175, 118, 121, 127) : Postero-basal patch of sensory hairs on inner surface of paramere absent (Text- figs 123, 124) : ; nigeriense sp. n. (p. Genal cones represented by two rounded a areas, often. shiny brown or black ee fig. 34) Genal cones, although small clearly present and ‘not at all shiny (Text- figs 3 5, Sy) hee Ultimate rostral segment long, 2:29 times longer than tenth antennal segment; M-+Cu stem about two-thirds as rg as Cu stem (Text-fig. 61); paramere as in Text-fig. 118 : : relatum sp. n. (p. Ultimate rostral segment shorter, I° 48-1- +88 times longer than tenth antennal segment ; M-+Cu stem at most half as long as Cu stem (Text-fig. 62); paramere as in Text-fig. 115 é pervatum sp. n. (p. Antennal flagellum shorter, 4: 19 mes longer than width of head: Cu, steeply para- bolic, second radial cell devoid of spinules (Text-fig. 64) . angolense sp. n. (p. Antennal flagellum longer, 4:90-5:26 times longer than width of head; Cu, weakly parabolic, second radial cell with spinules at least in distal half (Text-fig. 65) Fore wing densely and evenly covered with spinules (Text-fig. 75) nigripes sp. n. (p. 27 41) 16 40) 17 40) 18 37) 44) 20 22 21 52) 48) 49) 23 33) 24 25 31) 30) 34) 32) 27 28 35 36 39 40 D. HOLLIS Fore wing at least with narrow bands on either side of veins devoid of spinules and often with other areas without spinules (Text-figs 78, 79) : : : : 29 Genal cones well developed, longer than wide. Central America setosum (Crawford) Genal cones moderately developed, wider than long. Ethiopian region ; : 28 Antennal flagellum 1-7—2-3 times longer than width of head julbernardioides sp. n. (p. 45) Antennal flagellum 1-2—1-4 times longer than width of head orientale sp. n. (p. 46) Antennal fagellum not more than 3:3 times longer than width of head : . 34 Antennal flagellum not less than 3-9 times longer than width of head : ‘ 30 Ultimate rostral segment relatively short (Text-fig 3), 1-1-1°6 times longer than tenth antennal segment; genal cones present but poorly developed (Text-figs 35, 30)F %: , ; : ‘ I 31 Ultimate rosea eee seach long (Text: aes Ae 1-7-2'5 times longer than tenth antennal segment; genal areas only slightly swollen and often shiny brown or black . 3 32 Ovipositor relatively aur: — i sear 76 finties iene iba hind ia, vale of ovipositor not modified (Text-fig. 188) : : nigripes sp. n. (p. 32) Ovipositor relatively long, proctiger 2:22-2:49 times longer than hind tibia, valves of ovipositor elongated apically (Text-fig. 189) : : nigeriense sp. n. (p. 33) Antennal flagellum very long, 5:5—6:2 times longer than width of head; larger species, hind tibia 1-o—1-1 mm long : ; relatum sp. n. (p. 31) Antennal flagellum shorter, 3-9—-5:3 times loaeer fen head width; smaller species, hind tibia 0-70-0-90 mm long A 5 33 Ultimate rostral segment shorter, I-7—1-9 times igaaer than qu Ais feantel cues proctiger longer, 2-4—2-7 times longer than hind tibia : pervatum sp. n. (p. 30) Ultimate rostral segment longer, 2:0—2°5 times longer than roth antennal segment; proctiger shorter, 2-0-2-2 times longer than hind tibia filiverpatum Enderlein (p. 29) Antennal flagellum 2-7-3-3 times longer than width of head; genal cones sae well developed (Text-figs 38-40) . 35 Antennal flagellum not more than 2-4 fas longer et ae of hea: oe cones poorly developed (Text-fig. 48) . ; : : E : : : 40 Proctiger less than 1-5 times longer than hind abe : ; : : : : 36 Proctiger more than 1-5 times longer than hind tibia : 38 Smaller species, head width about 0-71 mm; ultimate rostral eeeeneat relatioaly long, 0-58—0-62 times as long as third antennal segment; ovipositor shorter, less than 1-1 times longer than hind tibia. : F vondraceki sp. n. (p. 51) Larger species, head width 0-78—o-94 mm; ultimate record segment relatively short, 0°39-0'52 times as long as third antennal segment; ovipositor longer, 1:28-1-41 times longer than hind tibia : : 37 Fore wing relatively narrow, 3:2-3:6 ates loupe on iain of era Peco small (Text-fig. 70) : 2 . cabudiatum sp. n. (p. 40) Fore wing relatively broad, 2- oe I ee longer than width of head, pterostigma large (Text-fig 81) : : capense Enderlein (p. 35) Third antennal segment not more dicen 2:0 Bois a than ultimate rostral segment. : 39 Third antennal segment oo less than Zoi times longer ese oltmuate rostral oe carvalhoi sp. n. (p. 41) Proctiger 1:6—1°8 times longer than hind tibia . ' . daubicatum sp. n. (p. 40) Proctiger more than 1-9 times longer than hind tibia bicaudatum Enderlein (p. 37) and tubacadium sp. n. (p. 44) Ultimate rostral segment longer than third antennal segment - : : : 41 Ultimate rostral segment shorter than third antennal segment . : : 5 42 » CIRIACREMINI OF THE ETHIOPIAN REGION 29 41 Proctiger relatively long, 1-7-1-9 times longer than hind tibia; ultimate rostral segment relatively long, 1-4-1-8 times longer than third antennal segment megafricanum sp. n. (p. 49) — Proctiger relatively short, 1:3-1-5 times longer than hind tibia; ultimate rostral segment relatively short, 1-o—1-3 times longer than third antennal segment africanum Enderlein (p. 48) 42 Second radial cell of fore wing with no spinules proximally and only a few spinules in distal part (Text-fig. 80) - : jilorense sp. n. (p. 52) — Second radial cell of fore wing more evenly spinuled throughout (Text-figs 72, 74) . 43 43 Fore wing narrow, 2:36-2:69 times longer than wide; proctiger longer, 1-5-1-7 times longer than hind tibia : capeneri sp. n. (p. 43) — Fore wing wider, 2:26-2:40 times longer than wide; proctiger shorter, 1-3—1°4 times longer than hind tibia : : ‘ : : : harteni sp. n. (p. 42) Ciriacremum filiverpatum Enderlein, 1910 (Text-figs 4, 34, 60, 90, III-113, 190) Ciriacremum filiverpatum Enderlein, 1910b : 139, text-fig. A, fig. 1. LECTOTYPE 4g, Tanzania: ‘Deutsches Ost Afrika’ (NR, Stockholm), here designated [examined]. Civiacremum filiverpatum Enderlein; Aulmann, 1912b : 101 Civiacremum filiverpatum Enderlein; Aulmann, 1913 : 80. DeEscrRiIPTION. Colouration. Overall body colour mid brown with ochraceous and black markings; vertex brown, ocelli orange, anteoccipital lobes ochraceous; genae ochraceous, each with a median shiny brown or black area; pronotum dark brown anteriorly and laterally, posterior margin ochraceous; mesopraescutum brown with ochraceous postero-lateral patches; mesoscutum brown, often with broad median longitudinal dark stripe and pale lateral patches; mesoscutellum brown with ochraceous antero-lateral corners; metascutellum ochraceous; tegula and pleurae light brown; wings hyaline; legs ochraceous with darkened tarsal segments; abdomen dark brown dorsally, ochraceous ventrally. Structure. g. Anteoccipital lobes close to lateral ocellar humps; genae rounded, hardly swollen (Text-fig. 34); antenna long, flagellum 4-03—4-69 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 :0°9: 1:0: 1:2 :1'4:1'7:0°4: 0-2; ultimate rostral segment long, 0-40-0-52 times as long as third antennal segment and 2:04-2:51 times longer than roth antennal segment. Fore wing elongate, 2:71-2:84 times longer than wide; Cu stem slightly more than twice as long as M+ Cu stem; first cubital cell elongate, about two and a half times longer than wide; spinule arrangement as in Text-fig. 60. Proctiger (Text-fig. 111) with simple basal part and a moderately long and narrow apical part; paramere (Text-figs 111-112) laterally flattened, pentagonal, on inner surface with a small anterior subapical tooth, a large apical tooth and a postero-apical ridge, in basal part with a large group of posteriorly directed hairs; distal segment of aedeagus as in Text-fig. 113; subgenital plate without hypovalves (Text-fig. 111). Q. Larger than g. Antennal flagellum 3-96—4:55 times longer than width of head, relative lengths of flagellar segments from base to apex—1:'0 :0°:9: 1:0: 1:2: 14: 1:8 +0*4 : 0:2; ultimate rostral segment 0-45-0-52 times as long as third antennal segment and 2:02-2:47 times longer than tenth antennal segment. Fore wing 2:77-2:89 times longer than wide. Ovipositor long (Text-fig. 190), proctiger 2:05-2:17 times longer than hind tibia. Measurements (+0-01 mm, based on 10 fg and 69). Maximum width of head, 3 0-99-1-09, © 1:01-1:12; length of antennal flagellum, J 4:11-4:90, 9 4:32—4-67; length of ultimate rostral segment, ¢ 0:25-0:29, 9 0:27-0:29; length of fore wing, ¢ 3-00-3-43, 2 3:25-3:70; maximum width of fore wing, ¢ 1-06-1-23, 9 1-13-1°33; length of hind tibia, J 0-78-0-89, 2 0-77-0°89; length of 2 proctiger, 1-67—1-92. Host PLANT. Some adults have been collected from Brachystegia spiciformis Bentham in Rhodesia but the bulk of the material examined was collected at light. 30 D. HOLLIS MATERIAL EXAMINED. CENTRAL AFRICAN REPUBLIC: 2 3, 2 9, La Maboke, 7.xi.1966, at light (M. Boulard). ZAIRE: I 9, Katanga, Elizabethville, ix. 1925; 11 g, 4 9, x. 1926; 7 4, 8 9, x. 1936; 12 3,9 9, ix.—xil. 1949, at light; 1 g, x. 1954, at light; 1 J, 1 Qix. 1958- v. 1959, at light; 8 g, 18 9, ix.—x. 1959, at light (Ch. Seydel); 1 g, 29,11. 1940 (H. Bredo); 3 3, 2 9, ix. 1931 (T. D. A. Cockerell); 1 9 Kapema, ix. 1924 (Ch. Seydel); 2 9, Kanzenze, 12.viii.19g31 (G. F. de Witte); 2 9, Kundelungu, riv. Kalumbulwa, 18.x.1951 (G. Marlier); 6 3, 5 2, Kolwezi, 5.x.1953, at light (Mme L. Gilbert); 1 g, I 9, Bianco, 8-11.viii.1931 (J. & L. Ogilvie); 1 9, Sakania, ix. 1931 (A. Mackie); 3 6, 5 9, Kivu, Kavimvira (Uvira), ix.—x. 1954, at light (G. Mazlier). ANGOLA: 2 9, Gangassol, x. 1972, at light; 1 J, 10 9, Nova Lisboa, 23.ix.1968 (M. Umbelina) ; 44, 109, 19-27.vili.1974, at light; 82 9g, 5 9, Chianga, 7—-9.xi.1966, at light; 15 d, 25 9, 17.X.1972, at light; 39 3, 50 9, 26.vill.1974, at light (J. Passos de Carvalho); I 3, 2 9, 5-19.1x.1970, yellow tray (A. van Harten); i 9, 8.ix.1963, at light (H. Cardosa); 16 3, 16 9, Ceilunga, 8.x.1972, at light; 2 g, 2 9, Cela, 27.viii. 1973; 1 g, Salazar, I.1.A.A., 29.xi.1972, at light; 1 9, vill. 1973 (J. Passos de Carvalho); 4 3, 2 8, Dundo, 4-29.viii.1953, light trap (Luna); 2 9, Benguela, Baia Farta (12-36S, 13°12E), vi. 1963, at light (Aldino Victorina); 1 9, Luzamba (9:03S, 18-07E), 22.V.-2.Vi.1971, at light (S. A. Peles); 1 9, Lac Calundo (11-485, 20-52E), 1150 m, 1g.xil.1954, at light (Machado & Luna). TANZANIA: 3 g, 4 9, ‘Deutsches Ost Afrika’ (type-series). RHODESIA: 6 4, 8 9, Salisbury, 21.xi.1967, from Brachystegia spiciformis (J. R. McDaniel); i 3, 29, Bulawayo, iv. 1968, at light (B. M. Gerard); 5 6, 7 2, Gokwe Distr., nr Kandariandzee Pan, vili.—ix. 1974, light trap (G. F. Cockbill). Discussion. C. filiverpatum, together with the following three species, seem to form a natural group within the genus Civiacremum. All these species have a peculiar head shape in which the genal areas are only slightly swollen and are mostly shiny brown or black. Apart from C. kleinielloides, which is only known from the female, they all have similar male genitalia; the paramere is of a similar form, showing slight variations in the form of the apical teeth and ridges (Text-figs 112, II5, 118); the distal segment of the aedeagus is very similar (Text-figs 113, 116, 11g); and the subgenital plate shows a complete progression from having well developed hypovalves in C. pervatum sp. n. (Text-fig. 114), through partly reduced hypovalves in C. rvelatum sp. n. (Text-fig. 117) to their complete reduction in C. filiverpatum. In all four species the wing shape and venation is very similar. Enderlein’s type-series is stored in alcohol and permanent microscopical prepara- tions have been made of the lectotype and male paralectotype. Ciriacremum pervatum sp. n. (Text-figs 12, 62, 114-116) DESCRIPTION. Colouration. Overall body colour varies from ochraceous to dark brown, probably depending on maturity; genae with central shiny brown or black areas, as in C. jilwerpatum. CIRIACREMINI OF THE ETHIOPIAN REGION 31 Structure. ¢. Similar to C. filiverpatum. Antennal flagellum relatively longer, 4-70—-5-36 times longer than width of head, relative lengths of flagellar segments from base to apex — I°O :0°9: 1-0 : 1:2: 1°4 : 18 : 0°5 : 0-2; ultimate rostral segment relatively shorter, 0:32-0:40 times as long as third antennal segment and 1-48—1-98 times longer than tenth antennal segment. Fore wing narrower, 2:89-3:24 times longer than wide; venation similar to C. filiverpatum, spinule arrangement as in Text-fig. 62. Proctiger (Text-fig. 114) and distal segment of aedeagus (Text-fig. 116) similar to C. filiverpatum; paramere (Text-figs 114, 115) on inner surface with more strongly developed apical tooth and posteroapical ridge; subgenital plate with short, broad, but well developed hypovalves (Text-fig. 114). Q. Antennal flagellum 4:72-5:22 times longer than width of head, relative lengths of flagellar segments from base to apex—1:0:0°9: 1I'I :1°3:1°5:1°8:0°4:0°2; ultimate rostral segment 0:33-0:36 times as long as third antennal segment and 1-72-1-88 times longer than tenth antennal segment. Fore wing 2:91-3:06 times longer than wide. Proctiger relatively longer, 2:44-2:70 times longer than hind tibia. Measurements (+0-01 mm, based on 5 g, 4 9). Maximum width of head, g 0-97-1°13, © 1-08-1-23; length of antennal flagellum, 3 4°57—-6-00, 9 5-10-6-30; length of ultimate rostral segment, ¢ 0:20-0:24, 2 0-23-0-26; length of fore wing, g 2-78-3-27, 9 3-23-3°46; maximum width of fore wing, g 0-95-1-06, 9 1-09—1-19; length of hind tibia, J 0-70-0-75, 9 0-71-0°79; length of 2 proctiger, 1-86—2-03. Host PLANT. Unknown. Holotype g, Nicertia: SE. State, Onya, 17.11.1971 (J. T. Medler) (BMNH, London); dry mounted. Paratypes. NIGERIA: I g, 5 9, Obudu CR, 21.iii.1971; 2 g, Ikom, 4.iv.1975; I g, W. State, Owena FR, 15.ili.1970; 2 9, Ile-Ife, 30.x.1968; 1 9, 30.x.1969; I J, Zana70, I 9, 26.11.1970; 3 g, 29.xi1.1970; 2.4, 2 9, 25.x.1971; 2 g, I 9, 30.X11.1971; I 9, Erin-Odo, 20.ii.1970 (J. T. Medler); 1 g, Erin-Odo, Waterfall, 17.11.1972 (EZ. W. Classey) (BMNH, London); slide and dry mounted. Discussion. See preceding species. Ciriacremum relatum sp. n. (Text-figs 61, 117-119) DEscriIPTION. Colouration. Overall body colour and pattern very similar to C. filiverpatum but genal humps hardly darkened. Structure. 3. Larger, more robust and a little more Censely haired than C. filiverpatum. Antennal flagellum 5:79-5:97 times longer than width oi head, relative lengths of flagellar segment from base to apex — 1:0 :0°9 : II : I°3 : 1°62 2*1 :0°5 : 0:2; ultimate rostral segment 0:46-0:49 times as long as third antennal segment and ~-25-2:34 times longer than tenth antennal segment. Fore wing 2:62-2:73 times longer than wide; Cu stem only about one and a half times longer than M-+Cu stem; first cubital cell elongate, more than twice as long as wide; spinule arrangement as in Text-fig. 61. Paramere (Text-figs 117-118) less robust than in C. filiverpatum and the apical teeth and ridge are less well developed; subgenital plate has reduced but clearly developed hypovalves (Text-fig. 117). Q. Larger than fg. Antennal flagellum 5:55-6:12 times longer than width of head, relative lengths of flagellar segments from base to apex —I'0 : 1°0 : I'l : 173 : 1:8 : 2°3 :0°5 10:2; ultimate rostral segment 0:47-0:49 times as long as third antennal segment and 1-92—2'15 times longer than 1toth antennal segment. Fore wing 2:65-2:72 times longer than wide. Ovipositor long, proctiger 2-02—2-08 times longer than hind tibia. Measurements (+0-01 mm, based on 4 ¢ and 3 9). Maximum width of head, ¢ 1-26-1°31, Q 1-30-1-37; length of antennal flagellum, ¢ 7-37-7°60, 9 7°60-7-95; length of ultimate rostral 32 D. HOLLIS segment, 3 0:42-0'43, 9 0-41-0-42; length of fore wing, f 3-42-3°51, 9 3-78-3-95; maximum width of fore wing, J 1:26-1-32. 9 1-43-1°46; length of hind tibia, J 1-02-1-09, 2 1-02-1-08; length of 9 proctiger, 2:08-2:24. Host PLANT. Unknown. Holotype g, NiGERIA: SE. State, Ikom, 4.iv.1975 (J. T. Medler) (BMNH, London); dry mounted. Paratypes. NIGERIA: 7 J, 149, same data as holotype; 2 3, 4 9, Oban, 7.iv.1975; 2 3, MW. State, Udo FR, I1.iv.1975; 1 J, W. State, Owena, 15.ii.1970 (J. T. Medler) (BMNH, London; and ZI, Leningrad); slide and dry mounted. Discussion. See C. filiverpatum Enderlein (p. 30). Ciriacremum kleinielloides sp. n. (Text-fig. 63) DESCRIPTION. 4. Unknown. Q. Similar to C. filiverpatum. Antennal flagellum 4:27 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 :0°9 : I'I !1'5 : I°7 : 2:3 :0°5: 0:3; ultimate rostral segment very long, 0-65 times as long as third antennal segment and 2-6 times longer than tenth antennal segment. Fore wing elongate, 2:69 times longer than wide; pterostigma sessile, broadly trapezoidal; first cubital cell shorter, about one and a half times longer than wide; Cu stem more than three times longer than M+ Cw stem; spinule arrangement as in Text-fig. 63. Ovipositor of moderate length, proctiger 1-89 times longer than hind tibia. Measurements (+0-01 mm, based on it 9). Maximum width of head, 9 1-13; length of antennal flagellum, 9 4:81; length of ultimate rostral segment, 9 0:34; length of fore wing, ° 3:29; maximum width of fore wing, 9 1:23; length of hind tibia, 9 0-87; length of 9 proctiger, 1°63. Host PLANT. Unknown; collected at light by waterfall. Holotype 2, ANGOLA: Cachoeiras, 20 mls [32 km] SW. Gabela, 18—19.iii.1972, at light (BM Southern Africa Expedition) (BMNH, London); slide mounted. Discussion. This species is unusual for the genus Civiacremum as the fore wing pterostigma is sessile, as in the genus Kleiniella. However, other characters suggest it is placed more naturally in Civiacremum. The head and wing shape are similar to C. filiverpatum but C. kleinielloides may be distinguished from that species by the relative proportions of the Cu stem to the M+Cuwu stem, the shape of the first cubital cell and the shorter ovipositor. Ciriacremum nigripes sp. n. (Text-figs 35, 65, 120-122) DESCRIPTION. Colouration. Overall body colour ochraceous and green; ocelli orange; fore wing hyaline; apical segment of fore tarsus brown or black; abdomen creamy white ventrally. Structure. dg. Medium size. Genal cones small but clearly developed (Text-fig. 35); antenna long, flagellum 4:90-5:26 times longer than width of head, relative lengths of flagellar segments from base to apex —I°O0 : I°O : I'l : 1:4 : 1-4 : 2°0 : 0-4 : 0-2; ultimate rostral segment short, 0:29-0-32 times as long as third antennal segment and 1:23-1:36 times longer than CIRIACREMINI OF THE ETHIOPIAN REGION 33 tenth antennal segment. Fore wing 2:49-2:63 times longer than wide, spinule arrangement as in Text-fig. 65; pterostigma elongate; Cu stem almost three times longer than M-+Cu stem; Cu, shallowly parabolic and first cubital cell about twice as long as wide. Proctiger (Text- fig. 120) with simple basal part and a moderately long and narrow apical part; paramere (Text-figs 120, 121) compressed with truncate apex, on inner surface in apical half with a transverse apical ridge, a small group of antero-ventrally directed setae and a large group of postero-ventrally directed setae, basal part with a posterior process bearing a large group of posteriorly directed hairs; distal segment of aedeagus as in Text-fig. 122; subgenital plate with short broad hypovalves (Text-fig. 120). 9. Antennal flagellum 4:95-5:42 times longer than width of head, relative lengths of flagellar segments from base to apex—1I'0:1°0: 1:2: 1°74 :1'5 :2:0:074:0°2; ultimate rostral segment 0-30-0-35 times as long as third antennal segment and 1-27-1-39 times longer than tenth antennal segment. Fore wing 2:53-2:62 times longer than wide. Proctiger moderately long, 1-58-1-76 times longer than hind tibia. Measurements (+0-o1 mm, based on 5 g and 5 9). Maximum width of head, 3 0-77-0-87, Q 0:84-0-91; length of antennal flagellum, gf 4-18-4-52, 2 4-18-4°62; length of ultimate rostral segment, ¢ 0:15-0°16, 9 0-15-0-17; length of fore wing, J 2-75-3°11, 9 2:97-3:28; maximum width of fore wing, gf, 1-08-1-23, 9 1-14-1-30; length of hind tibia, J 0-65-0-67, 2 0:64-0:70; length of 2 proctiger, 1-08-1-19. Host PLANT. Unknown. Holotype g, NicEertA: SE. State, Obudu CR, 13.iv.1973 (J. T. Medler) (BMNH, London); dry mounted. Paratypes. NIGERIA: 2 g, same data as holotype; 12 J, 5 9, 21.11.1971; 3 J, 3 Y, Onya, 17.11.1971; 26 g, Oban, 7.iv.1975; 18 3, 9 9, EC. State, Umuahia, 10.iv.1973; 16 g, 3 2, MW. State, Udo FR, I1.iv.1975; 1 g, W. State, Ile-Ife, 26.ii.1970; 1 J, NW. State, Bida CRH, 1.ix.1970 (J. T. Medler); 1 9, Ibadan, 9.iv.1966, at light trap (Dept. Agric. Res.); 1 9, Lagos, Ikoyi, 7.iii.1975 (M. A. Cornes) (BMNH, London; MRAC, Tervuren; and USNM, Washington); slide and dry mounted. Discussion. C. nigripes sp.n. and the following species have very similar general body form and fore wing shape and venation but both species have distinctive male and female genitalia and are probably not very closely related. In C. nigripes the general form and chaetotaxy of the male paramere is unlike any other species in the genus although it has the postero-basal sensory patch of hairs common in Ciriacremum species. Ciriacremum nigeriense sp. n. (Text-figs 36, 66, 123-125, 189) DEscriPTION. Colouration. Overall body colour very similar to C. nigripes but apical segment of fore tarsus ochraceous. Structure. g. Very similar to C nigripes. Antennal flagellum shorter, 4-0-4:68 times longer than width of head, relative lengths of flagellar segments from base to apex — I-0 : 0°9 : I°2 :1°3 :1°3:1°5 :0°4:0°2; ultimate rostral segment o-29-0-39 times as long as third antennal segment and 1-18-1-5 times longer than tenth antennal segment. Fore wing slightly more elongate, 2:58-2:78 times longer than wide, spinule arrangement as in Text-fig. 66. Proctiger (Text-fig. 123) with relatively longer apical part; paramere (Text-figs 123, 124) thumb-like with subconical apex, inner surface in apical half with a diagonal row of five to seven antero-ventrally directed setae and a small tooth posteriorly; distal segment of aedeagus as in Text-fig. 125; hypovalves short and narrow (Text-fig. 123). 34 D. HOLLIS Q. Larger than g. Antennal flagellum 4-34—4-72 times longer than width of head, relative lengths of flagellar segments from base to apex —I1-:0 : 1:0: I-2 : 1:3 :1°3 :1°5 :0°4 :0:2; ultimate rostral segment 0-33-0-38 times as long as third antennal segment and 1:18-1:53 times longer than tenth antennal segment. Fore wing 2-76—2-91 times longer than wide. Ovipositor (Text-fig. 189) elongated in apical half with the ventral valves thickened, proctiger 2:22-2:49 times longer than hind tibia. Measurements (+0-01 mm, based on 7 g and 6 9). Maximum width of head, $ 0-69-0°81, © 0:77-0'84; length of antennal flagellum, ¢ 2:74-3:82, 2 3:46-3:91; length of ultimate rostral segment, gf 0:13-0°15, 9 o-14-0-16; length of fore wing, ¢ 2:27-2:83, 9 2:84-3:24; maximum width of fore wing, ¢ 0:83-1:03, 2 1-o1-1-14; length of hind tibia, J 0-48-0-64, 9 0:54-0°65; length of 9 proctiger, 1°34-1°45. Host PLANT. Unknown. Holotype 3, NIGERIA: SE. State, Ikom, 4.iv.1975 (J. T. Medler) (BMNH, London); dry mounted. Paratypes. NIGERIA: 22 g, 16 9, same data as holotype; 2 g, Onya, 17.iii.1971; I g, Obudu CR, 13.iv.1973; 22 gf, 2 9, Oban, 7.1v.1975; I g, 2 9, EC. State, Umuahia, 10.iv.1973; 3 g6, MW. State, Udo FR, 11.iv.1975; 1 9, W. State, Ile—Ife, 25.iii.1969 (J. T. Medler) (BMNH, London; MRAC, Tervuren; and USNM, Washington) slide and dry mounted. Discussion. Superficially C. nigeriense sp. n. resembles the preceding species but the male genitalia are of a very different form. The paramere lacks a postero- basal sensory patch of hairs and is a relatively simple, thumb-like structure, much more like that found in the genus Kleiniella. The form of the female ovipositor is unique among the known members of the tribe and suggests that C. nigeriense lays its eggs in an unusual site. A similarly modified ovipositor is found in Psylla lovantht Capener, 1973, and that author mentions that the long ovipositor enables the female of P. loranthi to penetrate the calyx of the Loranthus flower and deposit eggs on the style. Ciriacremum angolense sp. n. (Text-figs 37, 64, gI, 126-128) DESCRIPTION. 4G. Medium size. Genae moderately developed (Text-fig. 37); antenna long, flagellum 4-19 times longer than width of head, relative lengths of flagellar segments from base to apex—1-:0 :0°9 : 1:2 :1°5 :1°'7:1°9:0°5 : 0:2; ultimate rostral segment short, 0°36 times as long as third antennal segment and 1-40 times longer than tenth antennal segment. Fore wing ellipsoid, broad, 2:39 times longer than wide, spinule arrangement as in Text-fig. 64; Cu stem slightly more than twice as long as M-+Cwu stem; Cu, steeply parabolic and first cubital cell hardly longer than wide. Proctiger (Text-fig. 126) with simple basal part and a short narrow apical part; paramere (Text-figs 126, 127) compressed, on inner surface in upper half - with a transverse subapical ridge and a number of strong downwardly directed setae, in basal part with a posterior lobe which bears a large group of short conical sensory pegs; distal segment of aedeagus as in Text-fig. 128; hypovalves (Text-fig. 126) short, narrow, with incurved apices. 9. Unknown. Measurements (+0-01 mm, based on I g). Maximum width of head, ¢ 0-95; length of antennal flagellum, ¢ 3:97; length of ultimate rostral segment, gj 0-16; length of fore wing, 6S 3°16; maximum width of fore wing, ¢ 1-32; length of hind tibia, ¢ 0-71. CIRIACREMINI OF THE ETHIOPIAN REGION 35 Host PLANT. Unknown; the single known specimen was collected at a light trap by a waterfall. Holotype g, ANGOLA: Cachoeiras, 20 mls [32 km] SW. Gabela, 18-19.iii.1972, at light (BM Southern Africa Expedition) (BMNH, London); slide mounted. Discussion. The affinities of this species are obscure. It has a very distinctive fore wing shape and venation and the male genitalia are quite unlike any other known species in the genus. In particular the male paramere has retained the postero-basal sensory patch but the setae are modified to short, thick sense cones. The form of the distal segment of the aedeagus is similar to that found in Ciriacremum capense and Kleiniella medleri. Ciriacremum capillicorne Enderlein, 1918 comb. rev. Ciriacremum capillicorne Enderlein, 1918 : 483, figs D, E. Holotype g, CAMEROUN: ‘Kamerun. Mitte Dezember’ (? depository). Bunoparia capillicornis (Enderlein) Enderlein, 1926 : 397. Description. (Partly translated from original description and interpreted from original figures.) Genae and anterior margin of vertex whitish, anterior half of vertex black, posterior half dirty yellow. Median suture sharp. Eyes almost round, large, black. Antenna more than twice as long as body, very flimsy, with almost complete covering of very short hairs, dark brown, first three segments brown-black. Thorax red-brown, prothorax dark brown, sternum dirty yellow. Abdomen brown-black, dorsally without hairs, ventrally with moderately thick silver-white hairs, those of ninth sternite very short. First tergite with chitinous yellow cone-like hump. Subgenital plate (original fig. D) without hypovalves. Parameres unusually long and slender, dark brown, each one apically with an inwardly directed tooth-like spine. Basal part of proctiger with elongate triangular lateral expansions. Fore wing (original fig. E) hyaline, pterostigma short, Cu stem about twice as long as M-+-Cu stem, Cu, steeply parabolic, first cubital cell slightly longer than wide. Measurements (from original description). ¢@, length of body, 3mm; antenna 6:2 mm; fore wing length, 3 mm. Host PLANT. Unknown. Discussion. C. capillicorne is only known from the male holotype which could not be traced. From the original description and figures it is difficult to relate this species within the genus but it would appear to resemble C. capense as it lacks hypovalves, the basal part of the proctiger has triangular lateral expansions and the paramere is a relatively simple elongate conical structure without a postero- basal sensory patch. It apparently differs from C. capense in its very long antennae. Ciriacremum capense Enderlein, 1923 comb. rev. (Text-figs 47, 81, 132-134) Civiacremum capense Enderlein, 1923 : 543. Holotype 9, SoutH Arrica: ‘Siid-Afrika Capland, East London, Lightfoot, Juli r914’ (SAM, Cape Town). Bunoparia capensis (Enderlein) Enderlein, 1926 : 397. Bunoparia capensis (Enderlein); Capener, 1970@ : 197. DEscRIPTION. Colouvation. Overall body colour mid brown with ochraceous and dark markings; pattern on dorsal surface of head and thorax very similar to C. bicaudatum (Text- 36 D. HOLLIS fig. 7); genae brown apically and ochraceous ventrally; proepisternum ochraceous, proepimeron dark brown, meso- and metapleurae dark brown; fore wing hyaline with brown patch proximal to point where claval suture joins hind margin; legs mainly ochraceous, femora sometimes darkened medially; abdomen brown dorsally, ochraceous ventrally. Structure. 3. Small species. Genal cones (Text-fig. 47) moderately well developed; antenna moderately long, flagellum 2:87-3:25 times longer than width of head, relative lengths of flagellar segments from base to apex—1I-0:0°9 : II :1°4 :1°3:1°5 :0°5 :0°3; ultimate rostral segment short, 0:42—0'52 times as long as third antennal segment and 1:24-1:57 times longer than tenth antennal segment. Fore wing broadly ellipsoid, 2:25—2-37 times longer than wide, spinule arrangements as in Text-fig. 81; pterostigma short and broad; Cw stem about one and two-thirds longer than M-+Cu stem; Cu, steeply parabolic and first cubital cell short, about one and one-third longer than wide. Proctiger (Text-fig. 132) with broad basal part which bears a pair of triangular lateral expansions, and a very short and narrow apical part; paramere (Text-figs 132, 133) thumb-like, on inner surface in apical half with a transverse subapical ridge which has a small tubercle at either end, and a few downwardly directed setae, basal part with a very poorly defined postero-basal group of setae; distal segment of aedeagus as in Text-fig. 134; subgenital plate (Text-fig. 132) without hypovalves. Q. Slightly larger than g. Antennal flagellum 2-77—3-12 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 :0°9 : II :1°3 :1:3:1°5 :0°5: 0°3; ultimate rostral segment 0-42—0-52 times as long as third antennal segment and 1:26-1:91 times longer than tenth antennal segment. Fore wing 2:24-2:41 times longer than wide. Ovipositor short, proctiger, 1:28-1:41 times longer than hind tibia. Measurements (+0:01 mm, based on 7 g and 69). Maximum width of head, ¢ 0-72-0-82, 2 0-78-0-90; length of antennal flagellum, $ 2:24-2°56, 9 2:43-2:80; length of ultimate rostral segment, gf 0-12—-0'14, 9 0-13-0'15; length of fore wing, gf 2-06—2°44, 2 2:26-2:70; maximum width of fore wing, ¢ 0:89-1:08, 9 0:95-1:20; length of hind tibia, g 0-49-0°'57, 9 0:50-0-61; length of 9 proctiger, 0-71-0°85. Host plants. Restricted to native South African species of the genus Schotia: S. speciosa Jacqin, S. latifolia Jacqin and S. brachypetala Sonder. MATERIAL EXAMINED. SoutH Arrica: Transvaal, Boyne, 15-17.xii.1973; 6 g, 7 9, Pretoria, 5.111.1964, Schotia brachypetala (A. L. Capener); 1 3, Kruger NP, Oliphants camp, 7.viii.1974, swept; I g, Skukusa, 8.viii.1974, swept (B. R. Pitkin); 1 3, 3 9, several larvae, Swaziland, Umbduzi R., 3.x.1967, Schotia brachypetala (H. D. Catling); 30 3, 30 8, several larvae, Natal (Zululand), Umfolozi GR, 25.viil.1g71, Schotia brachypetala (H. P. Insley); 1 3, Cape Province, Mossel Bay, i. 1922; 1 J, Somerset East, ix. 1930; 3 9, Katberg, xii. 1932-11. 1933 (R. E. Turner); 4 3, 49, van Stadens Pass, 27.x.1964, Schotia latifolia; 1 3, 1 9, Knysna, 18.x.1964; 1 g, Bathurst, 9.ii.1966; I g, I 9, Port Alfred, 7.ii.1966 (A. L. Capener); 3 3, 2 9, several larvae, 31.1.1972, Schotia afra (V.C. Moran). Discussion. C. capense is the only known species in the whole group to have extended into temperate southern Africa. It displays some distinctive features particularly with respect to wing venation and the male genitalia. The male subgenital plate lacks hypovalves, the paramere is a relatively simple structure and the proctiger has lateral expansions, similar to Kleiniella species. Furthermore the distal segment of the aedeagus is similar to that of K. medleri. The chorological, morphological and host plant evidence suggests that C. capense has been isolated from the bulk of Ciriacremum species for some time. CIRIACREMINI OF THE ETHIOPIAN REGION 37 Ciriacremum funestum sp. n. (Text-figs 46, 82, 92, 129-131) DeEscriPTION. Colouration. Overall body colour green or ochraceous; fore wing hyaline. Structure. g. Mediumsize. Genal cones poorly developed (Text-fig. 46); antennal flagellum moderately long, 2:81-3:07 times longer than width of head, relative lengths of flagellar segments from base to apex—I:0:1'0:1'2:1°5:1:°6:2:0:0:6:0-4; ultimate rostral segment relatively long, 2:09-2:50 times longer than tenth antennal segment, third antennal segment I-08—-1-23 times longer than ultimate rostral segment. Fore wing relatively broad, 2:39-2-49 times longer than wide, pterostigma punctiformly sessile, strongly developed and causing the costa to bulge outwards, Cu stem about one and three-fifths longer than M-+Cwu stem, Cu, steeply parabolic and first cubital cell hardly longer than wide, spinule arrangement as in Text-fig. 82. Proctiger (Text-fig. 129) with simple basal part and moderately long and narrow apical part; paramere (Text-figs 129, 130) laterally flattened with broad basal half, narrow apical half and a postero-basal lobe, inner surface with an apical tooth and a poorly developed medial sensory patch of hairs; distal segment of aedeagus as in Text-fig. 131; hypovalves (Text-fig. 129) simple, narrow, about as long as paramere. ®. Antennal flagellum 2-59-2-98 times longer than width of head, relative lengths of segments from base to apex — 1°0 - 1-0 : 1:2 3 1°5 : 1°7 : 2*I : 0°7 : 0°4; ultimate rostral segment 2°23-2°39 times longer than tenth antennal segment, third antennal segment 1:04-1:22 times longer than ultimate rostral segment. Fore wing 2-35—2-47 times longer than wide, costal bulge at pterostigma less developed than in g. Ovipositor relatively short, proctiger 1-47—1°56 times longer than hind tibia. Measurements (+0-01 mm, based on 7 ¢ and 6 9). Maximum width of head, ¢ 0-78-0-86, © 0°79-0'93; length of antennal flagellum, ¢ 2:32-2:43, 9 2:26-2:40; length of ultimate rostral segment, ¢ 0:21-0°25, 2 0:21-0:25; length of fore wing, ¢ 2:18-2:47, 2 2:17-2:61; maximum width of fore wing, ¢ 0:90-1:02, 9 0:89-1:09; length of hind tibia, J 0-60—0-66, 2 0-58-0-66; length of 2 proctiger, 0:87-1:02. Host PLANT. Unknown. Holotype ¢, NicErIA: SE. State, Ikom, 4.iv.1975 (J. T. Medler) (BMNH, London); dry mounted. Paratypes. NiGERIA: 27 J, 20 9, same data as holotype; 3 3, 1 2, Oban, 7.iv.1975; I 9, Obudu CR, 17.iii.1973; 1 g, 1 9, EC. State, Umuahia, 10.iv.1973 (J. T. Medler) (BMNH, London; NCI, Pretoria; and USNM, Washington); slide and dry mounted. Discussion. C. funestum sp. n. is an interesting species having characters common to both Kleiniella and Ciriacremum. It has the sessile pterostigma and long wing-vein hairs of K/leiniella but the short antennal hairs, grouped hind wing costal setae, grouped hind tibial spurs, and male hypovalves of Civiacremum. The presence of these hypovalves indicates that this species belongs in the latter genus. The male paramere is most unusual in having a clearly developed postero-basal lobe, and the strongly developed pterostigma with a corresponding costal bulge is also remarkable. Ciriacremum bicaudatum Enderlein, 1918 comb. rev. (Text-figs 2, 6, 7, 9, 10, 14-18, 38, 67, 68, 138-141, 191) Ciriacremum bicaudatum Enderlein, 1918 : 482, fig.C. Syntypes gj and Q, Tanzania: ‘Deutsches Ost-Afrika, November’ (? depository). Bunoparia bicaudata (Enderlein) Enderlein, 1926 : 397. DEscRIPTION. Colouvation. Overall body colour brown with dark brown and ochraceous markings; dorsum of head and thorax with pattern as in Text-fig. 7; head dark brown ventrally; 38 D. HOLLIS proepisternum ochraceous, proepimeron dark brown, meso- and metapleurae dark brown; fore wing hyaline, rarely with brown infuscation distally, apices of veins brown, marginal cells with median apical brown spots; femora dark brown, ochraceous basally and apically; tibiae and basitarsi ochraceous, apical tarsal segments brown; abdomen red-brown dorsally, yellow laterally but with a red median longitudinal stripe, dark brown ventrally; ¢ proctiger brown basally and red apically, hypovalves dark brown; ovipositor dark brown. Structure. 4. Mediumsize. Genal cones moderately well developed (Text-fig. 38); antenna moderately long, flagellum 2:61-3:36 times longer than width of head, relative lengths of flagellar segments from base to apex—I:0:1°0:1':2:1°5:1'5 :1°6:0°6:0:3; ultimate rostral segment short, 1:37-1:92 times longer than tenth antennal segment, third antennal segment 1-69—2-14 times longer than ultimate rostral segment. Fore wing ellipsoid, 2-44—-2-65 times longer than wide, spinule arrangement as in Text-figs 67, 68; pterostigma long, Cu stem about twice as long as M-+Cuw stem, first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 138) with simple basal part and a very long narrow apical part; paramere (Text-figs 138-140) compressed, with broad basal part and a long narrow apical part, on inner surface with an apical tooth and a large postero-medial toothed ridge which bears a group of downwardly directed setae, ventral half bears a group of postero-ventrally directed setae; hypovaives long and broad (Text-figs 138, 139) with a poorly defined medial hump bearing a few barely modified setae, on inner posterior surface with a moderately developed vertical row of inwardly directed curved setae; distal segment of aedeagus (Text- fig. 141) with a narrow apex. Q. Slightly larger than g. Antennal flagellum 2-78-3-03 times longer than head width, relative lengths of flagellar segments from base to apex —1'0 : 1:0 : 1:2 :1°5:1°6:1:6:0°6: 03; ultimate rostral segment 1-33-1-67 times longer than tenth antennal segment, third antennal segment 1:67-2:14 times longer than ultimate rostral segment. Fore wing 2:39-2:66 times longer than wide. Ovipositor relatively long (Text-fig. 191), proctiger 1-67—2-14 times longer than hind tibia. Measurements (+0-01 mm, based on 14 g and 89). Maximum width of head, 3 0-72-0:94, 2 0-82-0-94; length of antennal flagellum, ¢ 2-30-2°78, 2 2:33-2:80; length of ultimate rostral segment, g 0-14-0'18, 9 0-14-0°16; length of fore wing, ¢ 2:60-3:10, 9 2:82-3:25; maximum width of fore wing, g 0:99-1:20, 9 1:12-1-31; length of hind tibia, 9 0-65—-0°75, 9 0-67-0°75; length of 9 proctiger, 1:29-1:56. Host PLANT. Unknown. MATERIAL EXAMINED. NIGERIA: EC. State, Umuahia, 1.iv.1973 (J. T. Medler). ZatrE: 10 g, 16 9, Kivu, Kavimvira (Uvira), xii. 1954, at light (G. Marlier); 1 g, Elizabethville, 1953-1955, at light; 1 g, 1 9, xi. 1959, at light (Ch. Seydel). ANGOLA: 2 3, I Q, Salazar, 29.x1.1972, at light; 1 9, vill. 1973; I 9, Cela, 27.viii.1973; 11 gf, 11 9, Chianga, 17.x.1972, at light; 1 g, 26.viii.1974, at light (J. Passos de Carvalho); 3 3, 8.ix.1963, at light (H. Cardosa) ; 2 2, 15—19.1x.1970, yellow trays (A. van Harten); 1 2, Tundavala, 13-16 km NW. Sa da Bandeira, 27—29.iii.1972, general sweeping (BM Southern Africa Expedition). MOZAMBIQUE: 2 g, I 9, Chibuto, 29.1x.1968, on Cashew. Discussion. C. bicaudatum and the following eight species present a very interesting species-group. A study of the biology and systematics of the group could give some useful information to refine species concepts and understand mechanisms of speciation in tropical psyllids. C. bicaudatum, as it is understood here, is a widespread and morphologically variable species. Its geographic range extends from southern Nigeria, through Zaire and Central Africa to Mozambique. The populations examined from either CIRIACREMINI OF THE ETHIOPIAN REGION 39 end of this range differ from much of the Central African material in that the fore wing has a more even covering of spinules on the dorsal surface (Text-figs 67, 68). Also, in the Nigerian specimens, the seventh and eighth antennal segments are relatively longer, the relative lengths of the flagellar segments being, in the Jj - hoon; 1:2: 1-6: 1-9: 2-3: 0-6: 0:4, and in the 9-1-0: 1°0:1°'2:1-6:1-°7: 2:0: 0°6:0-4. The male genitalia are also subject to slight variation, particularly in the length of the hypovalves (Text-figs 138, 139). The other eight species in the group occur both allopatrically and sympatrically with C. bicaudatum. C. daubicatum sp. n. from Ghana and C. cabudiatum sp. n. from the Ivory Coast would appear to be westward replacement species of bicaudatum, and C. capeneri sp. n. from Rhodesia may be a southern replacement, although C. bicaudatwm is known from southern coastal Mozambique. C. harteni sp. n. C. carvalhoi sp. n., C. julbernardioides sp. n. and C. orientale sp. n. are sympatric with C. bicaudatum in Brachystegia woodland but may occur on different host plants. For instance C. carvalhoi feeds on Hymenostegia laxiflora, C. julbernar- dioides on species of Julbernardia, and C. harteni is found on Brachystegia spiciformis and B. tamarindoides. This point will only be verified when the host of C. bicaudatum is known. The male genitalia of the bicawdatum-group conform to a basic plan but show modifications in different structures which reflect the various lines of evolution within the group. Taking C. bicaudatum as a starting point, but not necessarily suggesting this species displays the primitive condition for all attributes concerned, one can trace these patterns of radiation. In C. bicaudatum the paramere (Text-fig. 140) has an elongate apical part and a moderately well developed postero-medial tubercle and sensory patch of setae; the hypovalves (Text-figs 138, 139) are long, broad and paired, with a slight medial hump bearing relatively undifferentiated setae, and the inner postero-vertical setae are moderately developed. The simplest development of the state is found in C. cabudiatwm sp. n. where the apical lobe of the paramere is shortened, the postero-medial tubercle and sensory patch are slightly more developed (Text-fig. 148); and on the hypovalves (Text-fig. 147) the medial hump is a little more developed with slightly more differentiated setae, and the inner postero-vertical row of setae is very much more developed. C. daubicatum represents a more derived condition along this line as the hypovalves have become elongate and slender, the medial hump is well developed and bears strongly differentiated setae, and the postero-vertical row of setae is very well developed (Text-figs 142, 143). In C. carvalhoi the thinner apical part of the hypovalve is lost but the specialized setae are retained in a subapical position (Text-fig. 161); in C. harteni the specialized setae are replaced by a tubercle (Text-fig. 165); and in C. capeneri the hypovalves have fused distally to form a single posterior structure (Text-figs 168, 170) with each half bearing a group of specialized setae on a tubercle. C. tubacadium has evolved along a different line. In this species the hypovalves have a less developed postero-vertical row of setae but on the paramere the apical lobe is reduced and the postero-medial sensory patch of setae is extended vertically along the inner posterior margin so forming an equivalent setal arrangement to the hypovalves of C.. bicaudatum (Text-figs 150, 151). 40 D. HOLLIS C. julbernardioides and C. orientale have similar paramere and hypovalve structures to C. bicaudatum but the apical part of the proctiger is reduced. These two species have heavily spinuled fore wings (Text-figs 75, 76), quite unlike any other Ethiopian species, and apparently breed on Julbernardia species rather than Brachystegia. Ciriacremum daubicatum sp. n. (Text-figs 39, 69, 142-145) DESCRIPTION. g. Very similar to C. bicaudatum. Antennal flagellum 3:00-3:12 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 : 1:0: I'2:1°5 :1°6:1°8 : 0-6 : 03; ultimate rostral segment 1-65 times longer than tenth antennal segment, third antennal segment 1:74-1:94 times longer than ultimate rostral segment. Fore wing 2:52-2:66 times longer than wide, spinule arrangement as in Text-fig. 69; pterostigma long; Cu stem about two and a quarter times longer than M + Cu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 142) with simple basal part and a very long and narrow apical part; paramere (Text-figs 142, 144) compressed, with broad basal part and moderately long and narrow apical part, on inner surface with an apical toothed ridge and well developed postero-medial toothed ridge and sensory patch of strongly developed setae, postero-basal patch of backwardly directed hairs moderately developed; hypovalves (Text-fig. 143) elongate, apical part hardly wider than apical part of proctiger, medial hump well developed and bearing four to five strongly developed curved setae, inner postero-vertical row of curved setae well developed; distal segment of aedeagus (Text-fig. 145) with narrow apex. Q. Slightly larger than g. Ultimate rostral segment 1I-60-1-77 times longer than tenth antennal segment, third antennal segment 1-67—1-87 times longer than ultimate rostral segment. Fore wing 2:5—2:6 times longer than wide. Ovipositor shorter than in C. bicaudatum, proctiger 1:68-1°79 times longer than hind tibia. Measurements (+0-01 mm, based on 4 gf and 69). Maximum width of head, ¢ 0:84-0:92, Q 0:83-0-95; length of antennal flagellum, 3 2-52-2-74, 9 2:50-2:90; length of ultimate rostral segment, gf 0-16-0°17, 9 0-16-0-18; length of fore wing, ¢ 2:80-3:36, 9 2:97-3:34; maximum width of fore wing, g 1:11-1:27, 9 1-17—-1°33; length of hind tibia, ¢ 0-70-0-83, 2 0-70-0:79; length of 9 proctiger, 1-23-1°37. Host pLant. Unknown; all specimens examined were collected at light. Holotype g, GHANA: Tafo, Cocoa Res. Inst., ix. 1973, light trap (M. A. K. Akotoye) (BMNH, London); slide mounted. Paratypes. GHANA: 3 4, 6 9, same data and depository as holotype; all slide mounted. For discussion of this species see under C. bicaudatum (p. 38). Ciriacremum cabudiatum sp. n. (Text-figs 40, 70, 146-140) DEscRIPTION. Colouvation. Overall body colour similar to C. bicaudatum but pattern less well developed. Structure. ¢g. Similar to C. bicaudatum. CGenal cones slightly more strongly developed (Text-fig. 40); antennal flagellum 3:11-3:35 times longer than width of head, relative lengths of flagellar segments from base to apex—1I:0 : 1:0: 1'2 :1°6: 1:6: 1'9 : 0:6: 0:3; ultimate rostral segment very short, 1:22-1:29 times longer than tenth antennal segment, third antennal CIRIACREMINI OF THE ETHIOPIAN REGION 41 segment 2:38—2-75 times longer than ultimate rostral segment. Fore wing 2:45-2:51 times longer than wide, spinule arrangement as in Text-fig. 70; pterostigma shorter than in C. bicaudatum; Cu stem slightly more than twice as long as M+Cuwu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 146) with simple basal part and a long and narrow apical part; paramere (Text-figs 146, 148) compressed, narrow apical part considerably shorter than in C. bicaudatum, on inner surface with an apical toothed ridge and well developed postero-medial toothed ridge and sensory patch of setae, basal part with poorly defined postero-basal sensory patch of hairs; hypovalves (Text-figs 146, 147) with moderately well developed medial hump which bears moderately differentiated setae, and a well developed inner postero-vertical row of curved setae; distal segment of aedeagus (Text-fig. 149) with broad apex. Q. Slightly larger than g. Antennal flagellum 3-01—3-12 times longer than width of head, relative lengths of flagellar segments from base to apex — 1-0 : 1-0: 1:2 :1°6:1'7:1°8:0°6: 0-3; ultimate rostral segment 1-23-1-:26 times longer than tenth antennal segment, third antennal segment 2°31—2°58 times longer than ultimate rostral segment. Fore wing 2:40-2:49 times longer than wide. Ovipositor relatively short, proctiger 1-38-1-40 times longer than hind tibia. Measurements (+0-01 mm, based on 5 ¢ and 3 9). Maximum width of head, 3 0:85-0:93, © 0:92-0:94; length of antennal flagellum, 3 2-65-2-98, 2 2-82-2-88; length of ultimate rostral segment, ¢ 0-12—-0'13, 2 0-12-0'13; length of fore wing, ¢ 2:80-2:94, 9 3-09-3:25; maximum width of fore wing, ¢ 1°13-1-19, 9 1-25-1-33; length of hind tibia, J 0:66-0:75, 2 0-70-0°75; length of 2 proctiger, 0-96-1-03. Host PLANT. Unknown. Holotype g, Ivory Coast: Bingerville, ix. 1962 (J. Decelle) (MRAC, Tervuren); dry mounted. Paratypes. Ivory Coast: 7 3, 8 9, same data as holotype. NIGERIA: I 4, I Q, SE. State, Ikom, 4.iv.1975 (J. TI. Medler) (MRAC, Tervuren; BMNH, London); slide and dry mounted. For discussion of this species see under C. bicaudatwm (p. 38). Ciriacremum carvalhoi sp. n. (Text-figs 44, 73, 160-163) DESCRIPTION. Colouration. Overall body colour ochraceous-green with light or dark brown markings; apices of genal cones red-brown; mesopraescutum with a pair of brown patches antero-medially; mesoscutum with two pairs of broad longitudinal brown bands; fore wing hyaline with a small brown spot proximal to point where claval suture joins hind margin; fore and mid femora dark brown ventrally; abdomen with median longitudinal dorsal brown stripe, proctiger with broad brown spot dorsally near junction between basal and apical parts. Structure. g. Similar to C. bicaudatum. Genal cones (Text-fig. 44) well developed; antennal flagellum 2-81-3-14 times longer than width of head, relative lengths of flagellar segments from base to apex — I'0 : 1°0 : 12: 1°5 : 1°6: 1°8 : 0°6 : 0°3; ultimate rostral segment relatively short, 1-05—-1-42 times longer than tenth antennal segment, third antennal segment 2:18-2:82 times longer than ultimate rostral segment. Fore wing 2°40-2:58 times longer than wide, spinule arrangement as in Text-fig. 73; pterostigma relatively short, Cu stem about one and one-third longer than M-+Cuwu stem; first cubital cell about one and one-third longer than wide. Proctiger (Text-fig. 160) with simple basal part and moderately long and narrow apical part; paramere (Text-figs 160, 162) compressed, with short narrow apical part, inner surface with an apical toothed ridge, a moderately well developed postero-medial toothed ridge and sensory patch of well developed setae, basal part with moderately well defined postero-basal sensory patch of setae; hypovalves (Text-figs 161) relatively short, with a subapical group of 42 D. HOLLIS four to six differentiated setae, postero-vertical row of curved setae moderately developed; distal segment of aedeagus (Text-fig. 163) with narrow apex. Q. Slightly larger than g. Antennal flagellum 2:89-3:07 times longer than width of head, relative lengths of flagellar segments from base to apex—as in g; ultimate rostral segment I-I12-1:21 times longer than tenth antennal segment, third antennal segment 2:25-2:54 times longer than ultimate rostralsegment. Fore wing 2-35-2-49 times longer than wide. Ovipositor relatively long, proctiger 1-83—1-98 times longer than hind tibia. Measurements (+0-01 mm, based on 9 g and 5 9). Maximum width of head, $ 0-77-0°87, Q o-79-0'85; length of antennal flagellum, ¢ 2-16-2-71, 9 2-29.2°53; length of ultimate rostral segment, g 0-oI—0-12, 9 o-10-0-11; length of fore wing, ¢ 2:34-2:73, 9 2:54-2:83; maximum width of fore wing, g 0:92-1:11, 9 1-02-1-18; length of hind tibia, J 0-52-0-60, 2 0-54-0-60; length of 2 proctiger, 1-06—1-14. Host PLANT. Type-series collected from young seedlings of Hymenostegia laxiflora (Bentham) Harms in an old coffee forest that was reverting to the wild state. Holotype g, ANGOLA: Salazar, I.1.A.A., 9-15.111.1972, Hymenostegia laxiflora (BM Southern Africa Expedition) (BMNH, London); dry mounted. Paratypes. ANGOLA: 27 g, 15 9, same data as holotype; 1 g, 2 9, at light (BM Southern Africa Expedition); I 9, 22.1x.1972, light trap (J. Passos de Carvalho); i 9, 5 km SW Salazar, 15.iii.1972, at light (BM Southern Africa Expedition). NIGERIA: I 4, Ile-Ife, 25.11.1969; 1 3, 28.11.1970; 1 9, MW. State, Benin, 20.iii.1972; 1 4, I Q, I.1v.1975; 2 6, 8.iv.1973; 2 9, nr Siluko, iii. 1973 (J. T. Medler); 1 3, 1 9, Sapoba FR, nr Benin, 10-12.ii1.1972 (E. W. Classey). GHANA: I g, I 9, Tafo (B. M. Gerrard). (BMNH, London; MRAC, Tervuren; and ZI, Leningrad); slide and dry mounted. Discussion. (See also under C. bicaudatum, p. 38). C. carvalhoi is closely related to the following two species, forming with them a well defined subgroup within the bicaudatum-group. All three species show similar modifications of the male hypovalves (Text-figs 161, 165, 170) from the bicaudatum-type. Ciriacremum harteni sp. n. (Text-figs 24, 42, 72, 164-167) DESCRIPTION. Colouration. Similar to C. bicaudatum. Femora and tarsal segments yellow; apical part of g proctiger light brown. Structure. ¢. Similar to C. bicaudatum. Genal cones less well developed (Text-fig. 42); antenna shorter, flagellum 1-91-2:23 times longer than width of head, relative lengths of flagellar segment from base to apex—1°0:0°9:1°O0:1'I : 1:2: 11 :0°5:0°5; ultimate rostral segment relatively longer, 1-84—2-14 times longer than tenth antennal segment, third antennal segment 1:06-1:19 times longer than ultimate rostral segment. Fore wing slightly broader, 2:28—2-56 times longer than wide, spinule arrangement as in Text-fig. 72; pterostigma short; Cu stem slightly less than twice as long as M+ Cw stem; first cubital cell one and one- quarter times longer than wide. Proctiger (Text-fig. 164) with relatively short apical part; paramere (Text-figs 164, 166) compressed, with long broad basal part and short narrow apical part, inner surface with an inner toothed ridge and a moderately developed postero-medial lobe, postero-medial and postero-basal sensory patches of hairs relatively undifferentiated ; hypovalves (Text-fig. 165) similar to C. carvalhoi sp. n. but with a subapical tubercle; distal segment of aedeagus (Text-fig. 167) relatively short with a broad apex. CIRIACREMINI OF THE ETHIOPIAN REGION 43 ®. Slightly larger than g. Antennal flagellum 1-91—2-19 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 :0°9 :0°9 : IT°O:1°I:I*1 :0°5 :0°4; ultimate rostral segment 1-74—2-26 times longer than tenth antennal segment, third antennal segment 1:05-1:25 times longer than ultimate rostral segment. Fore wing 2:26-2:40 times longer than wide. Ovipositor shorter than in C. bicaudatum, proctiger 1:32-1:40 times longer than hind tibia. Measurements (+0-01 mm, based on 8 g and 89). Maximum width of head, ¢ 0-63-0°68, @ 0-66-0-75; length of antennal flagellum, J 1-25-1-40, 9 1-34-1°57; length of ultimate rostral segment, ¢ 0-16-0°18, 9 0-16-0-20; Length of fore wing, g 1-91-2-05, 9 2:06-2:45; maximum width of fore wing, g 0-77-0°87, 9 0-88-1-06; length of hind tibia, J 0°52-0-58, 2 0-55-0-64; length of 2 proctiger, 0-77-0-87. Host prants. Adults and larvae have been taken from Brachystegia spiciformis Bentham and B. tamarindoides Welwitsch ex Bentham. Holotype g, ANGOLA: Chianga, 21-24.ii1.1972, Brachystegia tamarindoides (BM Southern Africa Expedition) (BMNH, London); dry mounted. Paratypes. ANGOLA: 20 J, 13 2, same data as holotype; I Q, vi. 1970, yellow tray; IO g, 14 9, 5-14.1x.1970, yellow tray; 5 g, 5 9, x. 1970, yellow tray; 2.4, xii. 1970- i. 1971; 4 4d, 14 9, 31.vill.1970, Brachystegia spiciformis; 4 3, 2 9, several larvae, 10.x.1970; 6 3, 2 9, several larvae, Catata, 17.11.1972, Brachystegia tamarindoides (A. van Harten); 2 3, 4 9, Tundavala, 9 mls [14.5 km] NW. Sa da Bandeira, 23.11.1972, Brachystegia spiciformis; 2 2, 15 mls [24km] N. Sa da Banderia, c. 6500’ [1981 m], 3.iii.1972, Brachystegia spiciformis (BM Southern Africa Expedition) (BMNH, London; MRAC, Tervuren; NCI, Pretoria); slide and dry mounted. Discussion. (See also under C. bicaudatum, p. 38). In C. harteni the male hypovalves are very similar to C. carvalhoi but the subapical setae are replaced by a small tubercle; also the inner postero-vertical row of curved setae is more strongly developed. The paramere has undergone considerable change with the postero- medial tubercle and sensory patch relatively undifferentiated. The aedeagus is unlike any other species in the bicaudatum group, the distal segment being short and thick. Ciriacremum capeneri sp. n. (Text-figs 27, 43, 74, 168-171) DEscRIPTION. Colouvation. Overall body colour and pattern similar to C. bicaudatum; genae completely ochraceous; femora not darkened; apical part of ¢ proctiger brown. * Structure. 3g. Similar to C. bicaudatum. Genal cones (Text-fig. 43) less well developed; antennal flagellum 1-82—2-40 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 : 1°O- 1-2 : I°5 : 1°4 : I*3 : 0°6 : 0-4; ultimate rostral segment 1°35-2-03 times longer than tenth antennal segment, third antennal segment 1:17-1:61 times longer than ultimate rostral segment. Fore wing 2:47-2:68 times longer than wide, spinule arrangement as in Text-fig. 74; pterostigma long; Cu stem about two and a half times longer than M+Cu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 168) with simple basal part and relatively short narrow apical part; paramere (Text- figs 168, 169) compressed, with moderately long and narrow apical part, on inner surface with apical tooth, a well developed medial toothed ridge and moderately well developed sensory patch of setae, basal part with a poorly defined sensory patch of hairs; distal segment of 44 D. HOLLIS aedeagus (Text-fig. 171) with broad apex; subgenital plate (Text-fig. 170) with short hypovalves which are fused along their inner apical surfaces to form a single structure or hypovalvular plate, a subapical tubercle with a few short setae is present on either half of this plate on the dorsal surface, ventro-medially the inner vertical rows of curved setae are developed up to the point where the hypovalves are fused. Q. Slightly larger than g. Antennal flagellum 1-97—2-18 times longer than width of head, relative lengths of flagellar segments from base to apex —1'0 : 1-0: 1:2 :1°5 :1*4:1I'4:0°7: 0°5; ultimate rostral segment 1-49—1-86 times longer than tenth antennal segment, third antennal segment I-2I-1-42 times longer than ultimate rostral segment. Fore wing 2:36-2:69 times longer than wide. Ovipositor of moderate length, proctiger 1-56-1-69 times longer than hind tibia. Measurements (+0-01 mm, based on 12 g and 5 9). Maximum width of head, ¢ 0-64-0°80, © 0-70-0°83; length of antennal flagellum, j 1:17-1:74, 9 1:38-1:74; length of ultimate rostral segment, gf 0-12-0'16, 9 0-12-0'15; length of fore wing, g 1-81—-2:50, 9 1:96-2:64; maximum width of fore wing, g 0°72-0°97, 9 0:83-1-05; length of hind tibia, J 0:42-0:58, 9 0°43-0°57; length of 9 proctiger, 0:67-0:90. Host PLANT. Brachystegia glaucescens Burtt Davy & Hutchinson. Holotype 3, RHODESIA: Rusape, vil. 1970, Brachystegia glaucescens (J. McDaniel) (NCI, Pretoria); slide mounted. Paratypes. RHODESIA: 24 g, 13 9, several larvae, same data as holotype; I 9, Salisbury, 6.vi.1958 (C. E. Taylor); 29 g, 38 9, 1967/1968, sticky traps (J. R. Blowers); 7 3, 3 2, Enterprise, 20.x.1960, on weeds in streak field-maize (Det. Agric.). TANZANIA: I 9, Nachingwea, x. 1953-i1. 1954, yellow tray (V. F. Eastop); I g, no data (probably Kenya) (BMNH, London; NCI, Pretoria); slide mounted and in 80% ethanol. Discussion. (See also under C. bicaudatum, p. 38.) C. capeneri is unlike any other species in the genus in that the male hypovalves are fused apically to form a single structure. On the general form of the male genitalia this species is clearly a member of the bicaudatum group and probably the most derived species of the subgroup including C. carvalhoi and C. hartent. Ciriacremum tubacadium sp. n. (Text-figs 41, 71, 150-152) DESCRIPTION. 4. Very similar to C. bicaudatum. Antennal flagellum 2:86-3:12 times longer than width of head, relative lengths of flagellar segments from base to apex —I°0 : I'I : 12 :1°6:1°7:1:7:0:7:0'4; ultimate rostral segment 1:52-1:58 times longer than tenth antennal segment, third antennal segment 1:71-1:92 times longer than ultimate rostral segment. Fore wing 2:55-2:72 times longer than wide, spinule arrangement as in Text-fig. 71; pterostigma long; Cu stem about twice as long as M+Cu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 150) with simple basal part and a long narrow apical part; paramere (Text-figs 150, 151) compressed with truncate apex, inner surface with apical tubercle, postero-medial tubercle moderately well developed, postero-medial sensory patch of setae extended vertically along inner margin of paramere, postero-lateral sensory patch of setae moderately well developed; distal segment of aedeagus (Text-fig. 152) with narrow apex. Q. Slightly larger than g. Antennal flagellum 2-88-2-99 times longer than width of head, relative lengths of flagellar segments from base to apex —I:0 : 1:0 : 1:3 :1°6:1°7:1°8:0°7: 0-4: ultimate rostral segment 1-40—-1°58 times longer than tenth antennal segment, third antennal CIRIACREMINI OF THE ETHIOPIAN REGION 45 segment 1:36-2:08 times longer than ultimate rostral segment. Fore wing 2-45—2-77 times longer than wide. Ovipositor relatively long, proctiger 2:05-2:18 times longer than hind tibia. Measurements (+0-01 mm, based on 6 ¢ and 5 9). Maximum width of head, $ 0-71-0-77, © 0-68-0-90; length of antennal flagellum, ¢ 2:17—2-31, 2 2:03-2°34; length of ultimate rostral segment, ¢ 0°13-0'14, 9 0:13-0'17; length of fore wing, g 2:63-2:94, 2 2:50-3:16; maximum width of fore wing, ¢ 1:03-1:13, 9 0:96-1:24; length of hind tibia, ¢ 0:60-0°65, 9 0°54-0°71; length of 2 proctiger, 1:14-1°45. Host PLANT. Unknown; type-series collected at light. Holotype 3, ANGoLA: Dundo, 10-13.viil.1953, light trap (Lwna) (BMNH, London); slide mounted. Paratypes. ANGOLA: 5 3, 4 2, same data and depository as holotype; slide mounted. Discussion. (See also under C. bicaudatum, p. 38.) C. tubacadium is unlike any other species in the bicaudatum-group because the vertical row of curved setae on the inner margins of the male hypovalves are much reduced and apparently replaced by the postero-medial sensory setae of the parameres which have become extended vertically along the inner posterior margin of each paramere. Ciriacremum julbernardioides sp. n. (Text-figs 21, 25, 45, 75, 153-156) DEscriPTion. Colouvation. Overall body colour light brown with ochraceous markings; dorsum of head and thorax with pattern similar to C. bicaudatum but not so obvious because the ground colour is much lighter; fore wing with yellowish tinge due to heavy density of spinules, sometimes distal hind margin infuscate; legs ochraceous. Structure. 9. Smaller than C. bicaudatum. Antennal flagellum relatively short, 1-61-2-15 times longer than width of head, relative lengths of flagellar segments from base to apex — IO :0°9 :I°O:I'2:1I°I : 1° :0°6:0°5; ultimate rostral segment 1:6-1-9 times longer than tenth antennal segment, third antennal segment 1-07—1°31 times longer than ultimate rostral segment. Fore wing relatively broad, 2-21-2-39 times longer than wide, evenly and densely covered with spinules on dorsal surface (Text-fig. 75); pterostigma long; length of Cu stem variable, from 1-7—2-4 times longer than M+ Cu stem; first cubital cell about one and a quarter times longer than wide. Proctiger (Text-fig. 153) with simple basal part and relatively short and narrow apical part; paramere (Text-figs 153, 155) compressed, with long narrow apical part and broad basal part, on inner surface with a moderately developed apical ridge, a well developed postero-medial toothed ridge and sensory patch of setae, postero-basal sensory patch of hairs moderately well developed; distal part of aedeagus (Text-fig. 156) with broad apex; hypovalves (Text-fig. 154) shorter than in C. bicaudatum with inner postero-vertical row of setae developed only in basal half. 9. Slightly larger than g. Antennal flagellum 1-77—2-21 times longer than width of head, relative lengths of flagellar segments from base to apex—1'0 :0'°9: I'O : 1:2: I:I :170:0'5: 0-4; ultimate rostral segment 1-6—1-8 times longer than tenth antennal segment, third antennal segment 1:06-1:44 times longer than ultimate rostral segment. Fore wing 2:18-2:37 times longer than wide. Ovipositor relatively short, proctiger 1:t4—1-31 times longer than hind tibia. Measurements (0-01 mm, based on 15 g and 109). Maximum width of head, ¢ 0:69-0-79, © 0-77-0°83; length of antennal flagellum, ¢ 1-14-1-62, 2 1:36-1-79; length of ultimate rostral segment, gf 0-15-0'18, 9 0-16—0-18; length of fore wing, g 2:05-2°54, 9 2:45-2:82; maximum width of fore wing, j 0-89-1-13, 9 1:05-1:29; length of hind tibia, 9 0:52-0:66, 9 0:59-0:69; length of 2 proctiger, 0-71-0-87. 46 D. HOLLIS Host pLants. Larvae and adults have been collected from- Julbernardia paniculata (Bentham) Troupin, and J. globiflora (Bentham) Troupin. Adults only have been collected from Brachystegia spiciformis Bentham. Holotype 3, ANGOLA: Tundavala, 9 mls [15 km] NW. Sa da Bandeira, 23.11.1972, beaten from Brachystegia spiciformis (BM Southern Africa Expedition) (BMNH, London); dry mounted. Paratypes. ANGOLA: 11 J, 7 9, same data as holotype; 1 9, I 9, 27—29.111.1972, general sweeping; I g, I 9, 15 mls [24 km] N. Sa da Bandeira, c. 6500’ [1981 m], 24.11.1972, Brachystegia woodland; 3 J, I Q, 3.iil.1972, Brachystegia spiciformis (BM Southern Africa Expedition); 2 3, 3 2, Chianga, 7—9.1x.1966, at light; 3 3, 5 9, several larvae, 15.1x.1970, Julbernardia paniculata; 1 3, 15-19.1x.1970, yellow tray (A. van Harten) ; 30 3, 43 2, 17.X.1972, at light; 1 J, 2 9, Cela, x. 1972, at light; 1 g, 27.11.1973 (J. Passos de Carvatho); 6 3, 6 9, Ceilunga, 8.x.1972, at light (MW. Umbelina); 3 3, rt 9, several larvae, Nharea, 6.x.1974, Julbernardia paniculata (A. van Harten). ZAIRE: I g, 3 9, Elizabethville, 17.x.1923 (Ch. Seydel). RHODESIA: I g, I larva, Salisbury, 12.1.1958, lucerne, bred on peas; 1 g, 18-19.x.1958, yellow trays; I dg, xi. 1958, yellow tray; 3 g, I 9, v. 1959, yellow tray (C. E. Taylor); 6 3, 6 9, several larvae, i. 1968, Julbernardia globiflora (J. R. Blowers) (BMNH, London; MRAC, Tervuren; NCI, Pretoria; ZI, Leningrad); slide and dry mounted, and stored in 80% ethanol. Discussion. (See also under C. bicaudatum, p. 38.) C. julbernardioides is closely related to the following species and the two are regarded here as sister species. On the structure of the male genitalia they are obviously members of the bicaudatum-group but both are easily distinguished because the membrane of the fore wing is evenly and densely covered with spinules on the dorsal surface. Ciriacremum orientale sp. n. (Text-figs 21, 76, 157-159) DESCRIPTION. ¢. Very similar to C. julbernardioides. Antennal flagellum shorter, 1-27— 1°43 times longer than width of head, relative lengths of flagellar segments from base to apex — I'0 :0°8 : 0:9 : I'0 : 0:9 : 0:9 : 0'5 : 0°4; ultimate rostral segment 1:87-2:50 times longer than tenth antennal segment, third antennal segment 0:93-1:00 times longer than ultimate rostral segment. Fore wing 2:12-2:27 times longer than wide. Proctiger (Text-fig. 157) with shorter narrow apical part; paramere (Text-figs 157, 158) with short narrow apical part and less well developed postero-medial ridge; distal segment of aedeagus (Text-fig. 159) with narrower apex; hypovalves relatively shorter, with inner postero-vertical row of setae not at all developed. Q. Slightly larger than g. Antennal flagellum 1-23-1-36 times longer than width of head, relative lengths of flagellar segments from base to apex — 1:0 : 0:8 : 0:9 : 0:9: 1:0 :0°9 :0°6: 0:6; ultimate rostral segment 1I-50-1-75 times longer than tenth antennal segment, third antennal segment 1:o-1:14 times longer than ultimate rostral segment. Fore wing 2:13-2:29 times longer than wide. Ovipositor shorter than in C. julbernardioides, proctiger 1:00—1-22 times longer than hind tibia. Measurements (+0-01 mm, based on 8 g and 69). Maximum width of head, 3 0:67-0:73, Q o-78-0-80; length of antennal flagellum, ¢ 0-85-1-00, 9 0:96-1:07; length of ultimate rostral segment, ¢ 0°15-0°17, 9 0-14; length of fore wing, ¢ 1:80—2-16, 9 2:34-2:57; maximum width of fore wing ¢ 0:80-0:98, 2 1-06-1-19; length of hind tibia, g 0-52-0:62, 2 0:55-0:59; length of 2 proctiger, 0°59-0-71, CIRIACREMINI OF THE ETHIOPIAN REGION 47 Host PLANT. Unknown. Holotype g, Tanzania: Nachingwea, x. 1953-1. 1954, yellow tray (V. F. Eastop) (BMNH, London); slide mounted. Paratypes. TANZANIA: 5 ¢, 29, same data as holotype. East Arrica [Kenya coast]: 2 4g, Myombo [Swahili for Brachystegia woodland], 13.iii.1914. MOZAMBIQUE: 4 9, 9 mls [14-5 km] NW. Dondo, 11.xi.1967 (H. D. Brown) (BMNH, London; NCI, Pretoria); all slide mounted. Discussion. The genetic relationship between C. julbernardioides and C. orientale presents an interesting problem at the species level that cannot be clearly resolved without further information on the biology and distribution of these two so-called species. julbernardioides - 6, 0;9,+ orientale oy. @19:* 0-90 0-80 0-70 head width, mm —~3> +10 0-20 0:30 length of 3rd antennal segment, mm —-> Fic. 21. Scatter diagram comparing head width against length of third antennal segment in Civiacremum julbernardioides and C. orientale. It is suggested here that C. julbernardioides and C. orientale form a sister pair of species separable, morphologically, on the structure of the male genitalia (Text-figs 153-159) and on the relative proportions of the antennal flagellum to head width (Text-fig. 21). C. julbernardioides is a widespread species occurring from southern Mozambique, through Rhodesia to southern and central Angola, apparently breeding 48 D. HOLLIS on trees of the genus Julbernardia. C. orientale is restricted to the coastal areas of Brachystegia woodland in East Africa. The host plant of C. orientale is not known, the species being described from a small series of both sexes trapped in yellow trays at Nachingwea (southern Tanzania), and two males labelled ‘Myombo [Swahili for Brachystegia woodland] East Africa, 13.iii.1914’. This date would preclude Tanzania so it is thought these are Kenyan specimens probably collected in the coastal woodland somewhere between Mombasa and Malindi. Also in the Nachingwea material there is a male and a female which have the antennal char- acteristics of C. ovientale but the male genital structure of C. julbernardioides. Furthermore I have examined two male and two females collected on the rift escarpment at Muguga, near Nairobi, which are similarly intermediate. Clearly the existence of morphologically intermediate specimens must cast doubt on the specific validity of C. orientale and it may be argued that these differences are due to the effect of the environment on the phenotype. Whatever the explanation, the observed morphological differences are considered here to have some genetic basis and C. julbernardioides and C. orientale are regarded as distinct species. The intermediate specimens are not assigned to either taxon. Ciriacremum africanum Enderlein, 1910 comb. rev. (Text-figs 8, 22, 23, 77, 172-174) Ciriacremum africanum Enderlein, 1910) : 140, figs 2 and B. LECTOTYPE 4, Tanzania: ‘Kilimandjaro’ (NR, Stockholm), here designated [examined]. Bunoparia africana (Enderlein) Enderlein, 1926 : 397. DESCRIPTION. Colouvation. Overall body colour mid or dark brown with ochraceous markings; dorsum of head and thorax with strongly developed pattern as in Text-fig. 8; genal cones ochraceous apically and ventrally; fore wing hyaline, vein endings often darkened; legs ochraceous. Structure. 3. Small species. Genal cones weakly developed (Text-fig. 8); antennal flagellum relatively short, 1:79-1:95 times longer than width of head, relative lengths of flagellar segments from base to apex—1:0:1'0:1'2:1'4:1'°5:1:°6:0:9:0°7; ultimate rostral segment relatively long, 1:02-1:43 times longer than third antennal segment and 1*57-2:03 times longer than tenth antennal segment. Fore wing 2:38-2:53 times longer than wide, spinule arrangement as in Text-fig. 77; pterostigma short; Cu stem two and a half to two and three-quarters times longer than M@+Cuwu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 172) with simple basal part and short narrow apical part; paramere (Text-figs 172, 173) strongly compressed, apical two-thirds broader than basal third, on inner surface with an antero-apical tubercle, posterior margin with a compound vertical row of short peg-like setae, postero-basal sensory patch of hairs moderately well developed; distal segment of aedeagus (Text-fig. 174) with narrow apex. Q. Slightly larger than g. Antennal flagellum 1-9g0-1-95 times longer than width of head, relative lengths of flagellar segments from base to apex—I:0: 1:0: I'l :1°4:1'5:1:°6:0°8: 0-7; ultimate rostral segment 1:00-1:46 times longer than third antennal segment and 1-69-I-91 times longer than tenth antennal segment. Fore wing 2:39-2:56 times longer than wide. Ovipositor short, proctiger 1:32-1:50 times longer than hind tibia. Measurements (+0-01 mm, based on 10 g and g 9). Maximum width of head, j 0:53-0:64, 2 0:52-0:67; length of antennal flagellum, ¢ 0:97-1:20, 9 1-05-1°30, length of ultimate rostral segment, § 0:13-0'15, 2 013-016; length of fore wing, gf 1:64-1:99, 9 1:69-2:24; maximum CIRIACREMINI OF THE ETHIOPIAN REGION 49 width of fore wing, ¢ 0-66-0-82, 9 0:69-0:94; length of hind tibia, J 0-38-0-46, 2 0-37-0-49; length of 2 proctiger, 0-54-0°67. Host PLANT. Among the material examined was a series of adults collected from Cassia sp. No further information is available. MATERIAL EXAMINED. ZAIRE: 3 6, 3 Q, Elizabethville, x.—xi. 1950, at light; I J, 2 9, 1953-1955, at light; I 9, xi. 1959, at light (Ch. Seydel). ANGOLA: I g, Chianga, 6~7.vili.1963, at light; 12 g, 3 2, 7-9.1x.1963, at light (H. Cardosa); 18 3, 31 9, 7-9.1x.1966, at light; 9 3, II 9, 5-14.ix.1970, yellow trays; I 9, xii. 1970-1. 1971, yellow trays (A. van Harten); 22 6, 20 9, 17.x.1972, at light; 6 g, 17 9, Ceilunga, 8.x.1972, at light; 4 3, Nova Lisboa, 19.vili.1974, at light; 5 g, 14 9, 19.x.1974 (J. Passos de Carvalho); 18 2, 22 9, 2.1x.1964, on Cassia sp. (E. de Fonseca). TANZANIA: I 4, I 9, Kilimanjaro (type-series). Discussion. C. africanum and the following species are regarded here as sister species. The form of the male paramere is similar in the two species and quite distinct from others in the genus. In C. vondraceki sp. n. (p. 51) the male paramere is similar in shape but the postero-medial sensory patch is composed of short non- articulated denticles, while that of C. africanum and C. megafricanum sp. n. is composed of peg-like articulated setae. Ciriacremum megafricanum sp. n. (Text-figs 19, 22, 23, 48, 78, 175-177) DESCRIPTION. Colouration. Similar to C. africanum (p. 48). Structure. 3. Slightly larger than C. africanum. Antennal flagellum 1-94—2-14 times longer than width of head, relative lengths of flagellar segments from base to apex —1:0 : 1:0: 1:2: I°5 :1°6:1'7 : 0-9 : 0-6; ultimate rostral segment much longer than in C. africanum, 1:53-2:00 times longer than third antennal segment and 2:19-2:76 times longer than tenth antennal segment. Fore wing 2:41-2:63 times longer than wide, spinule arrangement as in Text-fig. 78. Genitalia similar to C. africanum (Text-figs 175-177), inner surface of paramere with slightly better developed group of short cone-like setae postero-medially. Q. Slightly larger than g. Antennal flagellum 1-96—2-10 times longer than width of head, relative lengths of flagellar segments from base to apex—1°0 : 1:0: 1:2: 1°4:1'6:1:8:0°9: 0-6; ultimate rostral segment 1-51-2-04 times longer than third antennal segment and 2:27-2:94 times longer than tenth antennal segment. Fore wing 2:46-2:67 times longer than wide. Ovipositor (Text-fig. 19) longer than in C. africanum, proctiger 1:74—1-95 times longer than hind tibia. Measurements (0-01 mm, based on 9 g and 7 9). Maximum width of head, 3 0-54-0-72, © 0:56-0:69; length of antennal flagellum, ¢ 1-05—1-41, 9 1-18-1-45; length of ultimate rostral segment, ¢ 0:19-0:24, 2 0-22-0:25; length of fore wing, g 1°64-2°31, 9 1-97-2°34; maximum width of fore wing, g 0:63-0:91, 9 0:77-0-92; length of hind tibia, J 0-45—-0°58, 2 0-48-0°58; length of 2 proctiger, 0:87-1:08. Host pLAnT. Unknown, all known material collected in traps or as vagrants. Holotype g, ANGoraA: Nova Lisboa, 19.viii.1974 (J. Passos de Carvalho) (BMNH, London); slide mounted. 50 D. HOLLIS oy africanum - 5x; 9,0 0-17 : 7 megafricanum - o,+; 9 e@ 15 mm — Q w length of 3rd antennal segment, ° = ° orl2 O18 0-24 length of ultimate rostral segment, mm ——~> 0-6 pe 125 vi / me Z re / G ZA / yf ‘. oh / / # aon / / | 7 / / . , / VA rs / / 7 a uy oa, Vi / / B. May / / : / / ays Vf if WA S if Bs = gh = ose / / . A : / Yé ig 4“ e@ eee , ys 2 / je Za = / ee e 7. co) / / 7 £ / e / 7 = / Fa Ze © / i 7 africanum -e ie) / / oo megafricanum - # oO 1-20 23 length of proctiger, mm —> Fics 22, 23. Scatter diagrams comparing lengths of various characters of Civiacremum africanum and C. megafricanum. 22, length of third antennal segment against length of ultimate rostral segment; 23, length of female hind tibia against length of female proctiger. CIRIACREMINI OF THE ETHIOPIAN REGION 51 Paratypes. ANGOLA: 22 3g, 32 9, same data as holotype; 1 g, 2 9, Dundo, 4-29.Vili.1953, at light (Luna); 3 g, 2 9, Chianga, 2~7.vili.1963, at light; I g, 2 9, 7-9.1x.1963 (H. Cardosa); 47 3, 62 2, 7-9.xi.1966, at light; 24 g, 24 9, 17.x.1972, at light; 6 3, 12 9, 26.viii.1974, at light (J. Passos de Carvalho); 2 3, I 2, 5-14.ix.1970, yellow trays (A. van Harten); 1 9, Tundavala, 8-10 mls [13-16 km] NW. Sa da Bandeira, 27—29.iii.1972, general sweeping (BM Southern Africa Expedition) (BMNH, London; NCI, Pretoria; USNM, Washington); slide mounted and stored in 80% ethanol. Discussion. C. megafricanum is very similar to C. africanum. The two species are distinguished from one another by the relative lengths of their ultimate rostral segments (Text-fig. 22) and female proctiger (Text-fig. 23), both structures being much longer in C. megafricanum. This would suggest either the two species have different host plants or, if not, then they have different feeding and oviposition sites on a common host. The host plant of C. africanum is probably Cassia sp. but C. megafricanum is only known from trapped and vagrant material. Both species occur together in trap catches. Ciriacremum vondraceki sp. n. (Text-figs 49, 79, 178-180) DESCRIPTION. ¢G. Small species. Genal cones poorly developed (Text-fig. 49); antennal flagellum 2:89-3:22 times longer than width of head, relative lengths of flagellar segments from base to apex—1:'0:0:9:1'0: 1:3 :1°4:2:0:0:6:0'4; ultimate rostral segment short, 0°52-0'61 times as long as third antennal segment and 1:46-1:66 times longer than tenth antennal segment. Fore wing 2:43-2:71 times longer than wide, spinule arrangement as in Text-fig. 79; pterostigma short; Cu stem about one and three-quarter times longer than M-+Cu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text- fig. 178) with simple basal part and a relatively short narrow apical part; paramere (Text-figs 178, 179) compressed, with short narrow apical part and broad basal part, on inner surface with a well developed apical tooth and a postero-medial group of large non-articulated denticles, postero-basal setae not at all developed; distal segment of aedeagus (Text-fig. 180) with broad apex and an extraordinary long terminal tube; hypovalves (Text-fig. 178) moderately long and broad. 9. Slightly larger than g. Antennal flagellum 3:05-3:11 times longer than width of head, relative lengths of flagellar segments from base to apex —I:0 :0°9: 1:0: 1°4:1'5:1'°9:0°6: 0-3; ultimate rostral segment 0:58-0:62 times as long as third antennal segment and 1-73-1-76 times longer than tenth antennal segment. Fore wing 2:47-2:52 times longer than wide. Ovipositor relatively short, proctiger I1-o4—1-09 times longer than hind tibia. Measurements (+0:01 mm, based on 3 g and 29). Maximum width of head, ¢ 0:63-0:65, Q o-71; length of antennal flagellum, ¢ 1:81-2:08, 9 2:16-2:20; length of ultimate rostral segment, jf 0:13-0'14, 9 0-15; length of fore wing, g 1:91-2:16, 9 2:39-2:46; maximum width of fore wing, g 0-73-0°89, 9 0:95-1:00; length of hind tibia, ¢ 0:47-0-48, 2 0-52-0°53; length of 2 proctiger, 0:56-0°57. Host pLantT. Unknown, the type-series was collected at light. Holotype g, Ancora: Dundo, ii-iii. 1954, at light (Luna) (BMNH, London); slide mounted. Paratypes. ANGOLA: 2 4, 2 9, same data and depository as holotype; slide mounted. 52 D. HOLLIS Discussion. C. vondraceki sp. n. is superficially similar to C. africanum but may be distinguished by its relatively longer antennae and the form of the male genitalia. The male paramere is very unusual in that on the postero-medial inner surface there is a large patch of apparently non-articulated denticles. Normally this area bears strongly developed articulated setae. Also in C. vondraceki the terminal tube of the aedeagus is very large. In the BMNH collections there are several specimens of both sexes from various localities in Central and West Africa which are similar to the type-series of C. vondraceki but show clear differences in the structure of the distal segment of the aedeagus (Text-figs 181, 182), wing venation and spinule arrangement. In the male specimens the aedeagus bears a large terminal tube, very similar to C. vondraceki, but different from one another in each case. Furthermore the male parameres are similarly adorned with non-articulated spinules on the inner postero- medial surface as in C. vondraceki. I suspect that these forms, together with C. vondraceki, form a complex of closely related species but inadequate material precludes further description at present. Ciriacremum jilorense sp. n. (Text-figs 50, 80, 135-137) DESCRIPTION. Colourvation. Overall body colour from green to mid brown with ochraceous markings; pattern on dorsum of head and thorax similar to C. bicaudatum (Text-fig. 7); genal cones ochraceous apically and dark brown or black ventrally; pleurae ochraceous or light brown; fore wing hyaline with a dark brown spot proximal to point where claval suture meets hind margin; legs mainly ochraceous, femora dark brown medially; abdomen dark along dorsal mid line, pale laterally and ventrally. Structure. @. Small species. Genal cones moderately well developed (Text-fig. 50); antennal flagellum relatively short, 2:35-2-51 times longer than width of head, relative lengths of flagellar segments from base to apex—I:0 : 1:0: I'l : 1-4: 1:6: 1:8 :0°7:0°5; ultimate rostral segment short, 0:55—-0-70 times as long as third antennal segment and 1-26-1-45 times longer than tenth antennal segment. Fore wing 2:26-2:44 times longer than wide, spinule arrangement as in Text-fig. 80; pterostigma short; Cu stem two to two and a half times longer than M+Cu stem; first cubital cell about one and a half times longer than wide. Proctiger (Text-fig. 135) with simple, relatively narrow basal part, and short narrow apical part; paramere (Text-figs 135, 136) relatively large, compressed, on inner surface in apical half with a vertical ridge on anterior margin, and a compound row of differentiated setae on posterior margin, a well developed tubercle medially and with a poorly developed postero-basal sensory patch of setae; distal segment of aedeagus (Text-fig. 137) with narrow apex; hypovalves (Text-fig. 135) short, narrow. Q. Slightly larger than g. Antennal flagellum 2:19-2:27 times longer than width of head, relative lengths of flagellar segments from base to apex—1'0 : 1:0: I-I :1°3:1°5 :1°7:0°7: 0+5; ultimate rostral segment 0:63-0:70 times as long as third antennal segment and 1-32-1°44 times longer than tenth antennal segment. Fore wing 2:22-2:46 times longer than wide. Ovipositor of moderate length, proctiger 1-55—-1-81 times longer than hind tibia. Measurements (-++-o-01 mm, based on 7 g and 5 9). Maximum width of head, ¢ 0:61-0:74, Q 0:68-0:79; length of antennal flagellum, $ 1°51-1°83, 2 1:49-1:76; length of ultimate rostral segment, ¢ 0-11-0°12, 9 0:12-0:13; length of fore wing, ¢ 1:73-2:00, 9 2:00-2:31; maximum width of fore wing, g 0-71-0-88, 9 0-82-1-05; length of hind tibia, J 0:43-0:50, 2 0-45-0°54; length of 9 proctiger, 0:75-0°85. CIRIACREMINI OF THE ETHIOPIAN REGION 53 Host PLANT. The type-series was collected from several specimens of Cynometra webberi Baker in the Kenya coastal Brachystegia woodland; no larvae were found. Holotype g, Kenya: Arabuku-Sokoke Forest, Jilore track, 2-3.vii.1974, from Cynometra webbert (David Hollis) (BMNH, London); dry mounted. Paratypes. KENYA: 9 3, 14 9, same data as holotype (BMNH, London; NM, Nairobi); slide and dry mounted. Discussion. C. silorense sp. n. is difficult to relate to other species in the genus, having quite distinctive male genitalia. The relatively simple paramere is similar to the primitive type found in Kleiniella species and it may be that C. jilorensis is an early offshoot of the main Ciriacremum stock. Chorological and host plant evidence would also suggest this species has been isolated from the bulk of the Central African species for some considerable time. REFERENCES AULMANN, G. I912a. Beitrage zur Kenntnis der afrikanischen Chermiden- (Psylliden-) Fauna. III. Ent. Rdsch. 29 : 35-36, 6 figs. —— 1912). Beitrage zur Kenntnis der afrikanischen Psylliden-Fauna. IV. Ent. Rdsch. 29 : 100-101, 4 figs. 1913. Psyllidavum Catalogus. 92 pp. Berlin. BEKKER-Micpisova, E. E. 1973. On the systematics of the Psyllomorpha and the position of the group within the order Homptera. [In Russian.]. Jn Narchik, E. P. (Ed.), Doklady na dvadzat’ chetvertom eshcheghodnom chtenii pamyati N. A. Kholodovskogo. 1971 Leningrad, 1973 : 90-117. CAPENER, A. L. 1968. A new genus and species of Psyllidae (Homoptera) from South Africa. J. ent. Soc. sth. Afr. 31 : 361-364, 15 figs. 1970a. Southern African Psyllidae (Homoptera) —1: A check-list of species recorded from South Africa, with notes on the Pettey collection. J. ent. Soc. sth. Afr. 33 : 195-200. 1970b. South African Psyllidae (Homoptera) — 2: Some new species of Diaphorina Low. J. ent. Soc. sth. Afr. 33 : 201-226, 102 figs. — 1973. Southern African Psyllidae (Homoptera) - 3: A new genus and new species of South African Psyllidae. J. ent. Soc. sth. Afr. 36 : 37-61, 123 figs. Crawrorp, D.L. 1911. American Psyllidae IV. A partial revision of subfamilies. Pomona Coll. J. Ent. 3 : 480-503, figs 157-160. 1912. Indian Psyllidae. Rec. Indian Mus. 7 : 419-435, 3 pls. 1914. A monograph of the jumping plant-lice or Psyllidae of the New World. Bull. U.S. natn. Mus. 85 : 182 pp., 30 pls. Dosreanu, E. & MANOLACHE, C. 1962. Homoptera Psylloidea. [In Rumanian.] Fauna Repub. pop. vom. Insecta 8 (3) : 1-376, 270 figs. ENDERLEIN, G. 1910a. Panisopelma quadrigibbiceps, eine neue Psyllidengattung aus Argentinien. Zool. Anz. 36 : 280-281, 1 fig. 1g10b. In Sjostedt, Y., Wissenchaftliche Ergebnisse dey Schwedischen Zool. Exped. nach dem Kilimandjavo, dem Meru und den umgebenden Massaisteppen Deutsches-Ostafrikas, 1905-1906. 12 Hemiptera. 8 Psyllidae, 137-144 pp., 2 figs, 1 pl. Stockholm. — 1918. Psyllidologica V. Zool. Jb. 41 : 479-486, 7 figs, 1 pl. — 1923. Psyllidologica VII. Dt. ent. Z. 5 : 543-544. — 1926. Psyllidologica VIII. Ent. Mitt. 15 : 397-401. —— 1927. Psyllidologica IX. Eine neue Psyllidengattung aus Kamerun. Ent. Mitt. 16 : 240-241, 1 fig. Froceatt, W. W. 1901. Australian Psyllidae. Part II. Proc. Linn. Soc. N.S.W. 1901 : 242-208, 3 pls. 54 D. HOLLIS HEsLop-Harrison, G. 1958. Subfamily separation in the homopterous Psyllidae — III (a-c). Ann. Mag. nat. Hist. (13) 1 : 561-579. 1960. Subfamily-separation in the homopterous Psyllidae — III (d-e). Ann. Mag. nat. Hist. (13) 2 : 157-168. KIEFFER, J. J. 1906. Eine neue gallenerzeugende Psyllide aus Vordre-Indien. Z. wiss. InsektBiol. 2 : 387-390, 5 figs. KLIMASZEWSKI, S. M. 1963. Eine neue Art der Unterfamilie Ciriacreminae aus Korea (Homoptera, Psyllidae). Bull. Acad. pol. Sct. Cl. II Sév. Sci. Biol. 11 : 91-94. 1964. Studien tiber Systematik der Unterordnung Psylloidea. [In Polish.] Avnls zool. Warsz. 22 : 81-138, 51 figs. KuwayaMA, S. 1907. Die Psylliden Japans. 1. Tvans. Sapporo nat. Hist. Soc. 2 : 149-189, mpl Lainc, F. 1930. Some records of Indo-Malayan Psyllidae. Indian Forest Rec. (Ent.) 14 : 166-175, 5 figs. Locinova, M. M. 1972. Revision of jumping plant-lice of the tribe Pauropsyllini Crawf. (Homoptera, Psylloidea, Carsidaridae). [In MRussian.] Ent. Obozr. 51 : 837-853, 49 figs, 2 tabs. Léw, F. 1886. Neue Beitrage zur Kenntniss der Psylliden. Verh. zool.-bot. Ges. Wien 36 : 149-170, 1 pl. McMUut ten, R. D. & Jone, C. 1971. Dithiocarbamate fungicides for control of Pear Psylla. J. econ. Ent. 64 : 1266-1270, 5 tabs. Sutc, K. 1914. Eine neue Panisopelma (Connectopelma sbgn. n.) und eine neue Tvioza aus Chile. Sber. K. béhm. Ges. Wiss. Math.-nat. Kl. 1914 (5) : 1-8, 2 figs. VONDRACEK, K. 1957. Fauna CSR. 9. Mery—Psylloidea. 431 pp., 265 figs. Prague. 1963. Jumping plant-lice (Psylloidea - Homoptera) of Central Africa. Part I (Congo). Acta ent. Mus. natn. Prague 35 : 263-290, 84 figs. CIRIACREMINI OF THE ETHIOPIAN REGION 55 —Brachystegia ————Julbernardia ———QCynometra _ a Hymenostegia ++++++Schotia sase===-( QSSIO 10 20 30 Mari. Estimated distribution limits in Ethiopian region of six genera of Caesalpiniaceae recorded as hosts of Ciriacremini. 56 D. HOLLIS 30 10— P hymenostegioides K. superba Wwe ‘ K. jassina “a K.congoensis 20— ae K. guineensis K. sp.1 nr guineensis ih me @®QOournm@dr @ | K. sp. 2 nr guineensis ae K.oblongata 3°— oa = 30 K. medleri 10 20 30 Map 2. Distribution of Palmapenna hymenostegioides and Kleiniella species. CIRIACREMINI OF THE ETHIOPIAN REGION w@ kleinielloides e| i» \ 10— “4 @ es @ filiverpatum 20— / A pervatum v relatum ‘i Map 3. Distribution of Civiacremum species, filiverpatum-group. 57 58 D. HOLLIS fe) as 2 a ; C4 = aS ee ae 30 wn%e La T ov ( we _?? a i O°. y \uomf Ay SSH 6 @ nigripes 2o-— Ss ™@ angolense be VA @ nigeriense a4 y) 4 capense saul Ne oe ine Map 4. Distribution of Ciriacremum species. CIRIACREMINI OF THE ETHIOPIAN REGION @ bicaudatum v daubicatum % cabudiatum © tubacadium @ julbernardioides @ orientale Map 5. Distribution of Civiacremum species, bicaudatum-group, part. 59 60 D. HOLLIS ns oC on g Vl fe) s pe o \ \ dD r) , 10 (9) H 1o— ¢ @ carvalhoi o> @ harteni mM capeneri 2o== ee i 10 20 30 Map 6. Distribution of Civiacremum species, bicaudatum-group, part. CIRIACREMINI OF THE ETHIOPIAN REGION africanum megafricanum vondraceki jilorense funestum Map 7. Distribution of Civiacremum species. 61 62 D. HOLLIS Scale line represents 1:0 mm. 24, C. harteni; 25, C. julbernardioides. Civiacremum species, late instar larvae. Fics 24, 25. 63 CIRIACREMINI OF THE ETHIOPIAN REGION ‘WIUI O.I syusseider OUT, aye9S ‘tuauago2 *D ‘Lz ‘ asuaqgvno *D ‘9z ‘OPAILT IesUL 93R[ ‘soloeds wnmasoviMg Lz « 9z SOL 64 D. HOLLIS Fics 28-37. Ciriacremini species; head, dorsal view. 28, Panisopelma (Connectopelma) conifrons; 29, Palmapenna hymenostegioides; 30, Kleiniella guineensis; 31, K. jassina; 32, K. oblongata; 33, K. medleri; 34, Civiacremum filiverpatum; 35, C. nigripes; 36, C. nigeriense; 37, C. angolense. Scale line represents o-I mm. CIRIACREMINI OF THE ETHIOPIAN REGION 65 Fics 38-50. Ciriacremum species; head, dorsal view. 38, C. bicaudaium; 39, C. daubicatum; 40, C. cabudiatum; 41, C. tubacadium; 42, C. harteni; 43, C. capeneri; 44; C. carvalhoi; 45, C. julbernardioides; 46, C. funestum; 47, C. capense; 48, C. megafricanum; 49, C. vondraceki; 50, C. jilorense. Scale line represents 0-1 mm. 66 D. HOLLIS Fics 51-56. Kleiniella species, fore wings. 51, K. superba; 52, K. jassina; 53, K. congoensis; 54, K. guineensis; 55, Kleiniella species 1 near guineensis; 56, Kleimiella species 2 near guineensis. Scale line represents 1-0 mm. CIRIACREMINI OF THE ETHIOPIAN REGION 67 Fics 57-61. Ciriacremini species, fore wings. 57, Kleiniella oblongata; 58, K. medleri; 59, Palmapenna hymenostegioides; 60, Civiacremum filiverpatum; 61, C. velatum. Scale line represents 1-0 mm. 68 D. HOLLIS Jt Fics 62-66. Ciriacremum species, fore wings. 62, C. pervatum; 63, C. kleinielloides; 64, C. angolense; 65, C. nigripes; 66, C. nigeriense. Scale line represents I-o mm. CIRIACREMINI OF THE ETHIOPIAN REGION 69 Fics 67-74. Civiacremum species, fore wings. 67, C. bicaudatum, from Angola; 68, same, from Nigeria; 69, C. daubicatum; 70, C. cabudiatum; 71, C. tubacadium; 72, C. harteni; 73, C. carvalhoi; 74, C. capeneri. Scale line represents Io mm. 70 D. HOLLIS s Fics 75-82. Ciriacremum species, fore wings. 75, C. julbernardioides; 76, C. orientale; 77, C. africanum; 78, C. megafricanum; 79, C. vondraceki; 80, C. jilorense; 81, C. capense ; 82, C. funestum. Scale line represents 1-0 mm. CIRIACREMINI OF THE ETHIOPIAN REGION 71 92 Fics 83-92. Ciriacremini species, costal margin of hind wing. 83, Kleiniella superba; 84, K. oblongata; 85, K. guineensis; 86, K. jassina; 87, K. congoensis; 88, K. medleri; 89, Palmapenna hymenostegioides; 90, Civiacremum filiverpatum; 91, C. angolense; 92, C. funestum. Scale line represents 0:1 mm. 72 D. HOLLIS 101 Fics 93-101. Ciriacremini species, male genitalia. Palmapenna hymenostegioides, 93, entire genitalia, lateral view; 94, paramere, inner surface; 95, distal segment of aedeagus. Kleiniella jassina, 96, entire genitalia, lateral view; 97, paramere, inner surface; 98, distal segment of aedeagus. K. congoensis, 99, entire genitalia, lateral view; 100, paramere, inner surface; 101, distal segment of aedeagus. Scale line represents 0-1 mm. CIRIACREMINI OF THE ETHIOPIAN REGION 73 105 Fics 102-110. Kleiniella species, male genitalia. K. guineensis, 102, entire genitalia, lateral view; I03, paramere, inner surface; 104, distal segment of aedeagus. K. oblongata, 105, proctiger, lateral view; 106, paramere, inner surface; 107, distal segment of aedeagus. K. medleri, 108, entire genitalia, lateral view; I09, paramere, inner surface; 110, distal segment of aedeagus. Scale line represents o-I mm. 74 D. HOLLIS III, entire Fics I1I-119. C. filiverpatum, genitalia, lateral view; 112, paramere, inner surface; 113, distal segment of aedeagus. C. pervatum, 114, entire genitalia, lateral view; 115, paramere, inner surface; 116, C. velatum, 117, entire genitalia, lateral view; 118, Scale line represents 0-I mm. Ciriacremum species, male genitalia. distal segment of aedeagus. paramere, inner surface; 119, distal segment of aedeagus. CIRIACREMINI OF THE ETHIOPIAN REGION Fics 120-128. lateral view; 121, paramere, Civiacremum species, male genitalia. C. nigvipes, 120, entire genitalia, inner surface; 122, distal segment of aedeagus. C. nigeriense, 123, entire genitalia, lateral view; 124, paramere, inner surface; 125, distal segment of aedeagus. C. angolense, 126, entire genitalia, lateral view; 127, paramere, inner surface; 128, distal segment of aedeagus. Scale line represents o-I mm. 75 76 D. HOLLIS 134 —S — 131 wae! FIGs 129-137. Civiacremum species, male genitalia. C. funestum, 129, entire genitalia, lateral view; 130, paramere, inner surface; 131, distal segment of aedeagus. C. capense, 132, entire genitalia, lateral view; 133, paramere, inner surface; 134, distal segment of aedeagus. C. jilovense, 135, entire genitalia, lateral view; 136, paramere, inner surface; 137, distal segment of aedeagus. Scale line represents o-I mm. CIRIACREMINI OF THE ETHIOPIAN REGION 77 Fics 138-149. Civiacremum species, male genitalia. C. bicaudatum, 138, entire genitalia, lateral view, specimen from Angola; 139, same, specimen from Nigeria; 140, paramere, inner surface; 141, distal segment of aedeagus. C. daubicatum, 142, entire genitalia, lateral view; 143, hypovalve, inner surface; 144, paramere, inner surface; 145, distal segment of aedeagus. C. cabudiatum, 146, entire genitalia, lateral view; 147, hypovalve, inner surface; 148, paramere, inner surface; 149, distal segment of aedeagus. Scale line represents 0-I mm. 78 D. HOLLIS Fiegs 150-159. Civiacremum species, male genitalia. C. tubacadium, 150, entire genitalia, lateral view; 151, paramere, inner surface; 152, distal segment of aedeagus. C. julbernardioides, 153, entire genitalia, lateral view; 154, hypovalve, inner surface; 155, paramere, inner surface; 156, distal segment of aedeagus C. orientale, 157, entire genitalia, lateral view; 158, paramere, inner surface; 159, distal segment of aedeagus. Scale line represents 0-I mm. CIRIACREMINI OF THE ETHIOPIAN REGION 79 Fics 160-171. Civiacremum species, male genitalia. C. cavvalhoi, 160, entire genitalia, lateral view; 161, hypovalve, inner surface; 162, paramere, inner surface; 163, distal segment of aedeagus. C. harteni, 164, entire genitalia, lateral view; 165, hypovalve, inner surface; 166, paramere, inner surface; 167, distal segment of aedeagus. C. capeneri, 168, entire genitalia, lateral view; 169, paramere, inner surface; 170, subgenital plate, anterior view; 171, distal segment of aedeagus. Scale line represents 0-1 mm. 80 D. HOLLIS dad (ast l t ws! 180 , f 181 SS Fics 172-182. Civiacremum species, male genitalia. lateral view; 173, paramere, inner surface; 174, distal segment of aedeagus. C. megafricanum, 175, entire genitalia, lateral view; 176, paramere, inner surface; 177, distal segment of aedeagus. C. africanum, 172, entire genitalia, C. vondraceki, 178, entire genitalia, lateral view; 179, para- mere, inner surface; 180, distal segment of aedeagus. C. species I near vondracekt, from Angola, 181, distal segment of aedeagus. C. species 2 near vondvaceki, from Nigeria, 182, distal segment of aedeagus. Scale line represents 0:1 mm. CIRIACREMINI OF THE ETHIOPIAN REGION 81 ~aig Su steshe SANAS 23S 191 Fics 183-191. Civiacremini species, female genitalia. Palmapenna hymenostegioides, 183, ovipositor, lateral view; 184, proctiger, dorsal view, with detail of anal margin. Kleiniella superba, 185, ovipositor, lateral view. K. medlevi, 186, proctiger, dorsal view. K. jassina, 187, ovipositor, lateral view. K. congoensis, 188, ovipositor, lateral view. Civiacremum nigeriense, 189, ovipositor, lateral view. C. filiverpatum, 190, ovipositor, lateral view. C. bicaudatum, 191, ovipositor, lateral view. Scale line represents 0-I mm. 82 D. HOLLIS INDEX Synonyms are in italics; page references to descriptions are in bold. africana, 48 africanum, 12, 25, 27, 29, 48-49, 50, 51, 52 angolense, 25, 27, 34, 35 Anomalopsylla, 5, 13 Anomalopsyllini, 5 Anomoneura, 4, 5 Anomoneurini, 5, 6, 11 Aphalarinae, 5 Arytaininae, 5 Bactericera, 5 bicaudata, 37 bicaudatum, 12, 25, 27, 28, 35, 37-38, 39, 49, 41, 42, 43, 44, 45, 46, 52 brachypetala, 36 Brachystegia, II, 12, 29, 30, 39, 40, 43, 44, 46, 47, 48, 53 Bunoparia, 5, 24, 25 cabudiatum, 25, 27, 28, 39, 40-41 Caesalpiniaceae, II capeneri, 25, 26, 29, 39, 43-44 capense, 13, 25, 26, 28, 35-36 capensis, 35 capillicorne, 24, 25, 26, 35 capillicornis, 35 Carsidara, 4, 5, 13 Carsidaridae, 13 Carsidarinae, 5 Carsidarini, 5 carvalhoi, 25, 27, 28, 39, 41-42, 43, 44 Cassia, II, 12, 49, 51 Ceriacreminae, 5, 10, II Ceriacremum, 13, 24 Ciriacreminae, 4, 5, IO, II, 13 Ciriacremini, 4, 5, 6, 10, 11, 13, 17, 18, 26 Citiacremum), 4, 5; 10) 11, 22) E4) 16) 18) 23) 24-25, 26, 30, 32, 33, 36, 37, 53 congoensis, 16, 18, 20, 21 conifrons, 13 Connectopelma, 5, 13 Cynometra, II, 53 daubicatum, 25, 27, 28, 39, 40 Desmiostigma, 5, 15, 16, 19 Endopsylla, 20 Epicarsa, 5 filiverpatum, 12, 24, 25, 26, 28, 29, 30, 31, 32 Freysuila, 5 funestum, 18, 26, 37 Geyerolyma, 4 glaucescens, 44 globiflora, 46 guineensis, 16, 18, 21, 22 harteni, II, 12, 25, 26, 29, 39, 42-43, 44 Homotoma, 5 Hymynostegia, II, 15, 39, 42 hymenostegioides, 14-15 jassina, 15, 16, 18, 19-20, 21 jilorense, 27, 29, 52, 53 Julbernardia, 11, 12, 39, 40, 46, 48 julbernardioides, 12, 25, 26, 28, 39, 40, 45, 46, 47 Kleiniella, 4, 5, 11, 14, 15-16, 18, 20, 23, 24, 25, 32, 34, 36, 37, 53 kleinielloides, 25, 26, 30, 32 latifolia, 36 laxiflora, 15, 39, 42 Liviinae, 5 loranthi, 34 Loranthus, 34 Macrohomotoma, 5 medleri, 13, 16, 18, 21, 23-24, 25, 26, 35, 36 megafricanum, I2, 25, 27, 29, 49, 50, 51 Mesohomotoma, 5, 13, 25 Moraceae, II Nesiope, 4 nigeriense, 27, 28, 33-34 nigripes, 27, 28, 32-33 oblongata, 15, 16, 18, 21, 22-23 orientale, 25, 26, 28, 39, 40, 46, 47, 48 Palmapenna, 14 paniculata, 46 Panisopelma, 4, 5, 13 Pauropsyllini, 13 pervatum, 25, 27, 28, 30-31 Phacopteron, 4, 13 Phacosema, 4, 16, 21 Phacoseminae, 4 Phacosemini, 4 Phytolyma, 13 Prionocnemidae, 4, 5 Pseudophacopteron, 13 Psylla, 34 Psyllidae, 5, II, 13 Psyllinae, 4, 5 quadrigibbiceps, 13 relatum, 25, 27, 28, 30, 31-32 Rhinopsylla, 5 Schotia, II, 36 setosum, 13, 26, 28 speciosa, 36 spiciformis, 29, 30, 39, 43, 46 Spondyliaspidae, 25 Dattorris, B.Sc. Department of Entomology BritisH Museum (NATURAL History) CROMWELL ROAD Lonpon SW7 5BD INDEX Spondyliaspinae, 5, 13 superba, 15, 16, 18-19, 20, 21 Syndesmophlebia, 5, 15, 16, 22 tamarindoides, II, 39, 43 Tenaphalara, 5 Triozinae, 5 tubacadium, 25, 27, 28, 39, 44-45 Tyora, 4, 5 Udamostigma, 4, 21 vondraceki, 25, 26, 28, 49, 51, 52 webberi, 53 83 tay A ee wi\ ie ee j ry ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. 3.50. . HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounp, L. A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. . CRosSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 108: 68 plates, 15 text-figures. October, 1971. 12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CrosskEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . von Havex, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CRosskEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £09.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND Bye b+ tes A TAXONOMIC REVISION OF THE GENUS ORNATIVALVA GOZMANY, 1955 (LEPIDOPTERA : GELECHIIDAE) ~ Ni 0 . 7 pe f s /~ generar ” | 29 JUL i976 , LIBRARY X ' & > kK. SATTLER BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 2 LONDON : 1976 A TAXONOMIC REVISION OF THE GENUS ORNATIVALVA GOZMANY, 1055 (LEPIDOPTERA : GELECHIIDAE) Biv KEAUOS SATTLER Pp. 85-152; 27 Plates; 27 Text-figures BULLETIN, OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34. No. 2 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 15 issued in five series corresponding to the Scientufic Departments of the Museum, and an Historical serves. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 34 No. 2 of the Entomology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.). ISSN 0524-6431 © Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 29 July, 1976 Price £9.80 A TAXONOMIC REVISION OF THE GENUS ORNATIVALVA GOZMANY, 10955 (LEPIDOPTERA : GELECHIIDAE) By K. SATTLER CONTENTS Page SYNOPSIS i ; ; : ‘ : ; ‘ : ; ; 87 INTRODUCTION : ; : ; t Z : : : : 87 HEAD PREPARATIONS : : ‘ : : : 90 ABBREVIATIGNS OF MUSEUMS AND INSTITUTIONS ; : : ‘ fore) ACKNOWLEDGEMENTS : : : : : : ; : : 90 Ornativalua GOZMANY, 1955 ‘ : . ‘ , : gI THE SYSTEMATIC POSITION OF Ovnetuelon ; : ‘ ; j ; 95 KEY TO THE SPECIES OF Oynativalva : MALES 3 : : : 4 96 KEY TO THE SPECIES OF Oynativalva: FEMALES. : ; P F 98 CHECK-LIST OF THE SPECIES OF Oynativalva . ; : ; ; : 100 The heluanensis-group : ; : : ; : : ; : IOI The evubescens-group ; : : : : : : : ; 106 The ornatella-group . : : : ‘ : : : : : 116 The plicella-group. : : : : - : : : é 117 The tamariciella-group - : : : : : : : ; 119 The plutelliformis-group. : : - : ; : Cf : 127 The cevostomatella-group . : , F : : : : ‘ 135 BIBLIOGRAPHY : : é ’ : ; : : , : 143 TEXT-FIGURES ‘ ‘ ' , : ; : . , ; 148 INDEX . : ; : : : : : : s : : 151 SYNOPSIS The Old World genus Oynativalua Gozmany is revised. The primary types of all but two of the discussed nominal species have been examined. Forty-three species and one subspecies are regarded as valid. Twelve new species are described and four new synonyms in species rank are established; Nothvis minutella Turati is removed from the synonymy of O. cerostomatella (Walsingham) and transferred to Anarsia Zeller. Keys to the species are provided, head structures with frontal processes are described and illustrated for a number of species, and the biology and distribution are discussed for all species. INTRODUCTION SIncE the publication of a taxonomic revision of the genus Oynativalva Gozmany (Sattler, 1967), further collections have become available, including a number of undescribed species. Particularly valuable material was received from Dr Z. 88 K. SATTLER Kaszab, Természettudomanyi Mtizeum, Budapest, who has made several expeditions to Mongolia. Asa result of Dr Kaszab’s collections in Mongolia many papers were published under the title Evgebnisse der Zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. The present paper is no. 362 in that series. During his first expedition Dr Kaszab discovered the first Mongolian Ornativalva species. Once alerted, he paid special attention to this genus and during subsequent trips collected extensively in localities in which Tamarix, the host-plant of the Ornatialva larvae, was observed. This resulted in a collection of about 200 speci- mens, representing at least 17 species, 7 of which are described as new in this paper. A separate publication on the Mongolian Ornativalva species could not be justified, as taxonomic papers which are based entirely on a limited geographical area and include descriptions of new taxa are generally unsatisfactory. Instead, the earlier publication (Sattler, 1967) is revised to include all additional material which has become available since 1967. In the course of the present work revised keys to the species are provided and each species is discussed. Descriptions and illustrations published previously are not repeated, but are supplemented or corrected where additional material makes this possible. Several species are illustrated in colour (Pl. 1). The colour figures are generally based on one particular specimen; however, where no perfect model was available, the artist was supplied with additional examples to facilitate the reconstruction of parts which are damaged on the model specimen. The measurements at the beginning of each description are those of the fore wing length of the smallest and biggest specimen measured in millimetres from the base to the apex of the wing. Sometimes only inadequate material was available, for example, rubbed specimens without fringes or specimens with curled or damaged apex and distorted wing base; therefore many measurements are necessarily approximate. Where only a limited number of specimens was available the variation in size may, of course, be greater than is indicated by the recorded measurements. A number of Ornativalva species have modified head structures with enlarged scale bases, prominent frons or distinct frontal process. These structures were not described in 1967. Where applicable, descriptions and illustrations of the modified head structures are provided in this paper. The terminology of the genitalia follows Klots (1956). A separate process at the base of the costa for which no suitable term was found is called ‘harpe’ in the present paper following the terminology used previously (Sattler, 1967). Under ‘Distribution’ only the countries are listed; however, if a species is restricted to a limited area of a country or if it is only known from one locality, this is expressed, for example, ‘Italy (Sicily)’ or ‘S. Iran’. All distribution records are based on material examined by myself unless stated otherwise. Each specimen is recorded in my card index and bears my determination label. Material which was considered in 1967 is not listed in the present paper; additional material is recorded under the heading ‘Further material examined’. Kaszab (1963; 1965; 1966; 1968) gave detailed accounts of his journeys in Mongolia. Each collecting site is identified by a number which also appears on the data labels REVISION OF GENUS ORNATIVALVA 89 of the specimens. These numbers are recorded in the present paper under ‘Material examined’ and follow the collector’s name, for example, ‘(Kaszab, no. 840)’. The collecting sites of the British Museum (Natural History) Southern African Expedition are identified by similar numbers on the specimen labels. Photographic records and detailed information on these sites are available in the diaries of the members of the expedition and a summary will be prepared for publication. In a few instances lectotype designations were necessary. When a species is based on an unspecified number of individuals without designation of a ‘type’ and there is now only one original specimen known, this is designated as the lectotype in accordance with the widely accepted interpretation of the International Code of Zoological Nomenclature. The question whether such a unique ‘type’ should be regarded as a holotype or lectotype has been discussed recently by Vane-Wright (1975 : 26-28) whose conclusions seem generally acceptable. If the only existing original specimen or one particular specimen out of a series of syntypes was clearly designated as ‘the type’ or ‘holotype’ by a subsequent author, this is accepted as a valid lectotype designation. Walsingham usually based his new species on a ‘type 3”, ‘type 9’ and ‘paratypes’. Mostly he lists the 3 in the first place, but sometimes the 9. Although this seems to indicate that he considered them as holotype, allotype and paratypes, they are all syntypes under the present Code and thus eligible for consideration as the lectotype. For the sake of consistency and in accordance with Walsingham’s intention it seems, however, advisable to select as the lectotype the specimen which Walsingham lists in first place, unless there are good reasons for doing otherwise. I have examined the holo- or lectotypes of all nominal species included in the present paper with the exception of cimelion Amsel, of which only paratypes were available, and siewersiellus Christoph, type-material of which could not be traced. The literature was considered as completely as possible and over 100 papers dealing with Ornativalva species are recorded. All papers examined are listed in the Bibliography (pp. 143-147). A complete bibliography for each species is recorded in my card index from which detailed information will be made available on request. Bibliographic references in the systematic part of this paper are confined to publica- tions from which unchecked information was extracted for the present study or which contain taxonomic changes, biological information and important illustrations. Misidentifications which have been corrected by re-examination of the material on which they were based are also recorded. Excluded are, in particular, check-lists and faunistic papers unless they fall into one of the above categories. In addition to the 43 species recognized as valid in this paper, 6 doubtful species are also included. In some instances inadequate material, such as poorly preserved specimens, groups in which the sexes are doubtfully associated or in which only one sex is represented, could not be placed with certainty. These doubtful species are excluded from the keys and the check-list; however, they are discussed in the appropriate places in the systematic part as ‘Ornativalva species 1-6’ and their genitalia are illustrated. go K, SATTLER HEAD PREPARATIONS The modified structures on the head of various Ornativalva species I studied and photographed with a ‘Stereoscan’ scanning electron microscope (Pls 5-12). The head of the moth was carefully cleared of scales by brushing with a fine snipe feather. The antennae, proboscis and labial palpi were removed and preserved in a capsule on the pin under the specimen. The denuded head was separated from the moth, mounted on a metal stub in a position that permitted examination and photography of all important aspects and coated with gold. Photographs were taken to show the dorsal, lateral and frontal view of the head. After examination the head was glued back on the specimen with a water-soluble glue. In this way the head can be studied on the moth under a binocular microscope. If required for further ‘Stereoscan’ examination, it can be removed again without damage to the specimen by dissolving the glue with a small drop of water. ABBREVIATIONS OF MUSEUMS AND INSTITUTIONS BMNH British Museum (Natural History), London, U.K. IE Istituto di Entomologia, Bolzano, Italy. LN Landessammlungen fiir Naturkunde, Karlsruhe, West Germany. MINGA Muzeul de Istorie Naturala ‘Grigore Antipa’, Bucharest, Rumania. MNHN Muséum national d’Histoire naturelle, Paris, France. MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin, East Germany. NM Narodni Museum, Prague, Czechoslovakia. NM Naturhistorisches Museum, Vienna, Austria. RSM Royal Scottish Museum, Edinburgh, U.K. SMN Staatliches Museum fiir Naturkunde, Stuttgart, West Germany. AVG) Department of Zoology, Tel-Aviv University, Tel-Aviv, Israel. ™ Transvaal Museum, Pretoria, South Africa. ™ Természettudomanyi Muzeum, Budapest, Hungary. ZSBS Zoologische Sammlung des Bayerischen Staates, Munich, West Germany. ACKNOWLEDGEMENTS I acknowledge gratefully the assistance with material and information received from Mr E. Arenberger, Vienna, Dr W. Dierl, ZSBS, Munich, Dr D. Gerling, TAU, Tel-Aviv, Ing. W. Glaser, Vienna, Dr L. A. Gozmany, TM, Budapest, Dr H. J. Hannemann, MNHU, Berlin, Dr K. W. Harde, SMN, Stuttgart, Count F. Hartig, IE, Bolzano, Mr E. Jackh, Bidingen, West Germany, Dr F. Kasy, NM, Vienna, Dr Z. Kaszab, TM, Budapest, Prof. J. Kugler and Mr S. Limon, TAU, Tel-Aviv, {he J. Moucha, NM, Prague, Mr E. C. Pelham-Clinton, RSM, Edinburgh, Dr A. Popescu- Gorj, MINGA, Bucharest, Prof. Dr R. Remane, Marburg, West Germany, Mr A. Valetta, Malta, Dr L. Vari, TM, Pretoria and Dr P. Viette, MNHN, Paris. I am also indebted to my colleagues at the British Museum (Natural History), particularly D. Hollis, W. G. Tremewan, K. Tuck, R. I. Vane-Wright, A. Watson and P. E. S. Whalley. The colour plate was produced by Mr B. Hargreaves, Orpington, Kent. The photographs of moths and genitalia preparations were produced by the Photographic REVISION OF GENUS ORNATIVALVA gI Unit of the BMNH. During the study of head structures I was assisted and advised by members of the Electron Microscope Unit of the BMNH. ORNATIVALVA Gozmany, 1955 Ornativalva Gozmany, 1955, Annls hist.-nat. Mus. natn. hung., S.N. 6 : 308, 309 [keys], 310. Type-species: Gelechia plutelliformis Staudinger, 1859, by original designation and monotypy. Ornativalvua Gozmany; Gozmany, 1958 : 194, 196 [keys], 233. Ornativalva Gozmany ; Sattler, 1960 : 16 [keys], 59. Pelostola Janse, 1960, Moths S. Afr. 6: 188. Type-species: Pelostola kalahariensis Janse, 1960, by original designation and monotypy. [Synonymized by Sattler, 1967 : 34.] Ornativalva Gozmany; Sattler, 1967 : 34. Ornatiwalva Gozmany; Sattler, 1973 : 231. Pelostola Janse; Sattler, 1973 : 236. [As synonym of Ornativalva.| Head (Pls 5-12) with or without frontal modifications or processes. Ocellus present. Man- dible present as small lobe. Proboscis well developed, squamose at base, apical portion with taste papillae. Maxillary palpus with four short segments. Labial palpus recurved, first segment much shorter than second; second segment with moderate brush of scales below; third segment slender, as long as or slightly longer than second (Sattler, 1967, pl. 1, fig. 3). Antenna without pecten on scape. Metascutum with paired group of narrow, hair-like scales (Text-fig. 2); in cerostomatella-group with broad, ovoid scales (Text-fig. 1). In fore wing veins R, and R; on long common stalk, distance R,-R, about twice distance R,—-R,;. In hind wing vein M, free from cell near RR, from one point with RF or from short common stalk with RR. Frenulum of 9 with three long setae, in some species of cerostomatella-group with only two setae. Humeral field on ventral surface of fore wing densely scaled, without microtrichia (Text-fig. 3). Fore wing 4:0-10:0 mm, often with characteristic pattern. Costal two-thirds of wing dark, dorsal third light between fold and dorsal margin. Light and dark zones sharply separated at fold, border line characteristically W-shaped. Light area sometimes with short dark streak on dorsal margin at base of wing. Fore wing usually smooth, rarely with groups of raised scales. In fresh specimens fore wing sometimes suffused with pink which fades under prolonged influence of daylight. GENITALIA g (Pls 13-20). Eighth tergite and sternite laterally fused, not separated into free flaps. Genitalia withdrawn inside eighth segment. One pair of coremata present at base of genitalia. Uncus poorly developed or reduced; developed and deeply divided in cervostomatella- group. Gnathos absent. Valva divided into two to five branches. Costa long, slender, clavate, always present, sometimes with basal process (termed ‘harpe’ in this paper). Valva (s. str.) always longer than costa and sacculus, usually with short apical spine. Sacculus fused with valva or clearly separated. In oynatella-group additional process arising in angle between valva and sacculus (Text-figs 13, 14). Anellus usually pair of weakly sclerotized lobes, densely set with setae. In some species of cevostomatella-group anellus lobes of characteristic shape. Aedeagus with bulbous base, slightly or strongly bent, apex often sharply pointed. Anterior part of bulbus ejaculatorius sometimes with sclerotization (Pl. 17, fig. 95). GENITALIA 9 (Pls 21-27). Lobus analis rectangular, wider than long, densely set with setae; row of setae with curved tips along posterior margin, in cevostomatella-group setae straight. Apophysis posterior short, about as long as eighth segment (including apophysis anterior). Posterior margin of eighth tergite with irregular row of setae; in cevostomatella-group without setae but with narrow band of scales instead (in genitalia preparations only scale bases remain) (Pl. 27, fig. 122). Apophysis anterior short, approximately one-third to one-half length of apophysis posterior, sometimes much reduced, rod-like, sometimes dilated distally. Sclerotized antrum present in some species. Ductus bursae of varying length, in some species very long, coiled. Ductus seminalis arises near ostium bursae; short section of ductus seminalis dilated near ductus bursae, inner wall of dilated section sometimes with minute spines. Inner surface 92 K. SATTLER of corpus bursae and sometimes anterior section of ductus bursae evenly set with minute spines. In some species (phavaonis, servatisignella, caecigena) spines in corpus bursae concentrated in narrow band. Corpus bursae always with signum. Typical signum with pair of strong spines, teeth or transverse ridges on irregularly shaped sclerotized basal plate; sometimes signum divided into two or signum of completely different shape. RemARKs. On the denuded head of most Ovnativalva species the transfrontal sulcus is externally visible as a narrow band which is free of scale bases and runs in a gentle arc between the antennal pits. (In this paper the transfrontal sulcus is taken as the dividing line between vertex and frons.) Below the transfrontal sulcus the scale bases are evenly distributed over most of the frontal region but more densely concentrated along the margin of the eyes. A narrow band between the tentorial pits is completely free of scale bases. Above the transfrontal sulcus the scale bases are unevenly distributed, with the densest concentration laterally and along the posterior margin of the head. The centre of the vertex is an area of low density and in some species (pulchella, frontella, cerostomatella and others) can be completely free of scale bases. The density of scale bases is not a direct indication of the density of the scale cover above a given point of the head surface. The areas which are free or almost free of scale bases are usually covered by scales arising on neighbouring areas. The bare patches on the vertex are covered by scales which arise laterally and point towards the centre line of the head. In some species of the evubescens-group and all but one of the cerostomatella-group, modifications of the frontal region occur. The scale bases below the transfrontal sulcus and immediately above it can be reduced in numbers and raised to form small knobs or teeth. They can be concentrated immediately below the transfrontal sulcus and form a more or less pronounced process. The scale bases immediately above the transfrontal sulcus can form a regular or irregular arc. The most extreme development of frontal processes is found in some species of the cerostomatella-group (cornifrons and some forms of mixolitha) (Pls 8-12). The frontal modifications are usually hidden under the scale cover of the head and can be easily overlooked unless the head is denuded for examination. Even in species with extreme processes only the tip may be visible among the scales. Nothing appears to be known about the function of such modifications of the frontal region. In some instances fine particles of sand were found among the raised scale bases. This suggests that some species pupate in the ground and that the moth has to bore through a layer of soil after emerging from the pupa. A strengthening of the frons would seem advantageous for such purposes; however, numerous species of moths manage perfectly well under identical conditions without such modifications. Head processes have developed independently in several families of Lepidoptera, for example, Cosmopterigidae, Symmocidae, Pyralidae (Crambinae, Phycitinae), Geometridae, Thyrididae, Noctuidae and Thaumetopoeidae. In the Gelechiidae modifications of the frons occur independently in a number of genera, for example, Lita Treitschke, Rhynchopacha Staudinger, Cerofrontia Janse, Scrobipalpa Janse, Ilseopsis Povolny and others. Species with frontal processes are found in different parts of the world; however, REVISION OF GENUS ORNATIVALVA 93 in the Gelechiidae there is a notable concentration in arid areas of the Palaearctic and Nearctic regions. In the Palaearctic region Gelechiidae with frontal modifica- tions occur from Morocco through North Africa, the Near and Middle East to western India and Central Asia, with a particular concentration in Mongolia. Frontal processes are found in both sexes and no sexual dimorphism was observed in this character. In Ornativalva mixolitha considerable variation of the frontal structures is found. The majority of specimens have no definite process but merely enlarged scale bases which are evenly distributed, whereas a distinct process is developed in specimens from Turkey and Mongolia. In southern Iran specimens with and without process and intermediate stages were found. In most Gelechiidae the metascutum is bare except for a paired group of modified scales near its posterior margin. The long, narrow, hair-like scales point in a postero-medial direction. The scutellum is covered with normal body-covering scales. In the species of the cevostomatella-group the scales on metascutum and scutellum are short, broad, ovoid and arranged in a shingled manner (Text-figs I, 2). The function of this structure, which is equally developed in both sexes, and the significance of its modification in the cevostomatella-group of the genus Ornativalva is unknown. Such modification is also found elsewhere in the Gelechiidae, for example, in Rhynchopacha gussakovskw (Gerasimov), which differs in that character from all other Rhynchopacha species. Like the species of the cerostomatella-group R. gussakovskii has also a frontal process; however, there is no correlation between frontal process and scale structure of the metascutum and there are many species of Gelechiidae with frontal process but hair-like scales on the metascutum. Differences between species are found in the wing shape and some species of the cerostomatella-group (cerostomatella, cornifrons) differ more strongly from the majority of the Ornativalva species (Text-figs 3-12). No sexual differences in the wing shape of Orvnativalva species were observed. For examination of the 9 frenulum it is recommended to look at both pairs of wings as there are specimens which have lost a seta or acquired an additional one, usually on one side only. The humeral field, a basal area on the ventral surface of the fore wing, between costa, subcosta and humeral cross-vein (Text-fig. 3) bears in many species of Lepi- doptera a group of microtrichia. The presence or absence of the microtrichia was used by some authors to separate Gelechiidae and Scythrididae; however, this character is so variable in the Gelechiidae that it has no significance at the family level. Several species of Ornativalva were examined but no microtrichia were found. The ductus bursae can reach considerable length in some species and is then coiled inside the abdomen. When uncoiled and fully extended in a preparation, it can measure up to three times the length of the extended abdomen. There appears to be some correlation between the length of the ductus ejaculatorius of the g and the ductus bursae of the 9. O. rufipuncta and longiductella with an exception- ally long ductus bursae have also an extremely long ductus ejaculatorius. This is not surprising when we consider that the spermatophore may be partially formed in the ductus ejaculatorius during copulation. The spermatophore is probably 94 K SATILER globular with a long coiled stalk that reaches the posterior part of the ductus bursae where the ductus seminalis branches off. Remnants of the spermatophore were found during preparation in the ductus bursae of rufipuncta. BioLtocy. The genus Oynativalva appears to be closely associated with the plant genus Tamarix (Tamaricaceae). The larvae of twelve Orvnativalva species were reared on Tamarix species. Many more species were collected as adults flying among Tamarix, indicating that their larvae can likewise be expected to feed on that plant genus. The larva of O. heluanensis was reared on Tamarix and Frankenia (Frankeniaceae). Frankeniaceae are considered to be closely related to Tamaricaceae. One specimen of O. plutelliformis is labelled ‘Rhus oxyacanthe’, but it is unlikely that Rhus is the host-plant of the larva of any Ornativalva species. Relatively few Ornativalva species were collected in the larval stage and then only in a small number of localities. There is no indication at this stage that Ornativalva species are host- specific within the genus Tamarix. The comparison of the distribution patterns of Tamarix species and Ornativalva species indicates that widely distributed species must have different host-plants in different parts of their range. This is confirmed by the fact that several different species of Tamarix were recorded for O. plutelli- formis. The species of Tamarix are not always easy to identify. A number of species were recorded as host-plants of Ornativalva species; however, some are clearly misidentifications whereas others remain doubtful. The host-plants are discussed under each species in the taxonomic section. The nomenclature of the Tamarix species in this paper follows Baum (1966). Ecc. Unknown. It is possibly deposited singly on the terminal shoots of Tamarix, where the larva is known to feed. Larva. The chaetotaxy of the larvae has never been studied. Descriptions of colour and pattern exist for several species. Two colour forms (green and pink) of the larva of plutelliformis were illustrated by Milliére (1861, (liv.4) pl. 1, figs 1, 2). Most of the descriptions are not comparative and thus of little value. Misidentifica- tions are frequent and authors were sometimes dealing with a mixed series of larvae. According to the literature the larva lives in a silken tube or free on the terminal shoots, leaves or flowers of Tamarix species. O. heluanensis was also recorded as mining the succulent leaves of Frankenia. Pupa. Superficial descriptions exist for a few species. The pupa of plutelliformis was illustrated in colour by Milliére (1861, (liv. 4) pl. 1, fig. 3); however, the figure is too small and shows little. The existing descriptions are unsuitable for comparison and the identification of the species is often doubtful. According to the literature Ornativalva species pupate on or in the ground in a cocoon that is covered with small particles of soil. In areas where hibernation is required this takes place in the pupal stage. AvuLt. Little is known about the habits of the moths. Mann observed adults of tamariciella flying and mating before and after sunset (Zeller, 1850 : 153). The Ornativalva species are readily attracted to light and most species have been collected that way. No observations exist on the number of generations per year, but all indications are that Oynativalva species breed continuously where climatic conditions REVISION OF GENUS ORNATIVALVA 95 are right. The most common and widely distributed heluanensis and plutelliformis were collected as adults in every month of the year. Clear-cut generations probably exist in areas with distinct seasons. DISTRIBUTION. The distribution of the genus Ornativalva coincides almost completely with that of the plant genus Tamarix. Ornativalva species are known from the Cape Verde Islands, Canary Islands and Madeira through southern Europe and North Africa to India and Mongolia. No species are known from China, where Tamarix also occurs. In the south some species are known from the Sudan and Socotra. An isolated species occurs in South Africa. THE SYSTEMATIC POSITION OF ORNATIVALVA Gozmany (1955 : 310; 1958 : 233), in his papers on the Hungarian Gelechiidae, placed the genus Ornativalua between Friseria Busck and Chionodes Hiibner; however, its relationship to other genera was not discussed. The genus Friseria is American in distribution and not yet recorded from the Palaearctic region; the only Hungarian species included by Gozmany belongs to RAynchopacha Staudinger. There is no evidence that Orvnativalva is closely related to either Chionodes or Rhynchopacha. Janse (1960 : 188), who described the genus as Pelostola in his work on the South African Gelechiidae, placed it ‘near to Scrobipalpa and Melitoxoides’. Neither genus is morphologically similar to Oynativalva. The placement of Orvnativalva between Altenia Sattler and Gladiovalva Sattler (Sattler, 1960 : 59) was arbitrary and not based on affinities. The higher classification of the Gelechiidae is still poorly understood. A number of subfamilies have been separated, but most of them have never been defined properly and the subfamily Gelechiinae remains a heterogeneous assemblage of all the genera which could not be assigned with certainty to any other subfamily. 2 Fics 1,2. Thorax of Ovnativalva species. 1, cevostomatella-group. 2, heluanensis-group — plutelliformis-group. 96 K. SATTLER The Gelechiinae are at present subdivided into three tribes, Teleiodini, Gnori- moschemini and Gelechiini, the last comprising a mixture of genera which could not be assigned with certainty to one of the other two tribes. As the tribes are defined at present, Oynativalva can only be included in the Gelechiini. Ornativalva does not appear to have close morphological affinities to any other genus and in particular is not related to the other Tamarix-feeders in the Gelechiidae. For practical purposes Ornativalva should remain next to Chionodes until a more appropriate place can be suggested. KEY 1O DE SPECIES OER IOR NAT VAL VAP NGAICESS (Note. The ¢ of undella is unknown.) I South African species : : ; ‘ : ‘ kalahariensis (p. 127) - Palaearctic species : : : é 2 2 (1) Metascutum with short broad Seale lees fg. I); uncus divided 2 = Metascutum with long narrow scales (Text-fig. 2); uncus not divided 9 3 (2) Fore wing with dark longitudinal band; costa and dorsal margin light 4 Fore wing without clearly defined longitudinal band; costa usually dark, dorsal margin lighter : : : 4 : ; 4 (3) Frons with strong process (PL. 12), 2 ES 78- 80): ground colour of fore wing white cornifrons (p. 138) = Frons weakly or strongly cages but without process; ground colour of fore fon) wing ochreous ; 5 5 (4) Distal half of valva een narrower fen basal ae oe iheweily ee (Pl. 9, figs 60-62) : . cerostomatella (p. 139) - Distal half of valva hardly narrower en asa half: frons strongly prominent (Pl. 10, figs 63-65) ; ‘ ! 3 mongolica (p. 140) 6 (3) Distal portion of valva strongly ered domed : : ; : . : 7 - Distal portion of valva straight F ‘ : : ; 8 7 (6) Frons with short cylindrical process (Pl. 11, figs 69— 7 ; afghana (p. 141) - Frons prominent, without cylindrical process (Pl. 10, figs 66-68) . curvella (p. 142) 8 (6) Base of anellus bulged ‘ : ‘ : : 3 : angulatella (p. 137) - Base of anellus not bulged : A : : mixolitha (p. 135) 9 (2) Sacculus developed, clearly separated from ae ‘ : 3 : : 10 - No separate sacculus developed : : ; : 38 10 (9) Frons with process and eg scale bases (Ps j Tp figs 39- 50) ; 3 , ae - Frons not modified . : P f 14 Ir (10) Fore wing with basal streak (PI. 1, fig. 4; Pl. Be fig. 20) : frontella (p. 114) - Fore wing without basal streak 3 ; : : : : 12 12 (11) Harpe clavate : : : : F . : : lilyella (p. 108) - Harpe digitate : : ‘ ; : : : 13 13 (12) Sacculus broadest near eae (PL. 14, = 85) : : : . aspera (p. 109) - Sacculus broadest distally (Pl. 14, fig. 86) . : ‘ : pulchella (p. 110) 14 (10) Fore wing with basal streak : ‘ é : : : : ; : 15 - Fore wing without basal streak F 22 15 (14) Conspicuous process between valva and Basie (PL 16, ae QZ: Text- figs ae mm ornatella (p. 116) - No process between valva and sacculus : F : é : : : 16 16 (15) Distal half of sacculus separated from valva g : : . : ‘ 17 = Sacculus fused with valva for most of its length : é : : : 20 33 (27) 34 (33) 35 (34) 6 (35) 37 (36) REVISION OF GENUS ORNATIVALVA Sacculus somewhat dilated in distal half Sacculus not dilated in distal half Sacculus broadest near apex (Pl. 19, fig. 99) ; : ; basistriga (p. Sacculus broadest at three-quarters, distal portion narrow (PI. 18, fig. 98) singula (p. Harpe long, digitate. ‘ d c : ; : ‘ . sieversi (p. Harpe short, rounded : : : : : heligmatodes (p. Apex of aedeagus sharply bent... : : : ; : erubescens (p. Apex of aedeagus gently curved : : : : . ‘ F : Harpe clavate : : ; : : . arabica (p. Harpe globular (PI. Ty fig, is ‘ : : - : levifrons (p. Harpe absent ; : : : : : : . ignota (p. Harpe present : . F é : : : : Harpe angulate, with shaxp pout : : ; : : pharaonis (p. Harpe rounded or digitate Aedeagus with rounded or truncate apex Aedeagus pointed Costa clavate . ; ‘ ; é , ; : Costa not clavate (PL. 18, fies 96); fore wing with raised scales (Pl. 1, fig. 7) caecigena (p. Valva strongly bent in middle (Pl. 17, fig. 93) . . : . plicella (p. Valva straight, not bent in middle. - : macrosignella (p. Harpe digitate, at least half length of costa Harpe rounded, short Valva strongly bent in middle (PL. I 5 fig. 8; 7) ; : ochraceofusca (p. Valva not bent in middle . Sacculus about twice as long as wide, . ; : ‘ sesostrella (p. Sacculus much longer than wide : ‘ Distal half of sacculus narrower than basal eal P Distal half of sacculus not narrower than basal half Aedeagus curved, S-shaped F : ‘ ; : : ontipyramis (p Aedeagus curved but not S-shaped . : . triangulella (p Bulbous base of aedeagus as long as narrow apical portion . plutelliformis (p. Bulbous base of aedeagus shorter than narrow apical portions . grisea (p Fore wing with small groups of raised scales ‘ : ; . indica ® Fore wing smooth, without raised scales : : Posterior margin of anellus with circular Seemann (Pl. 1, fie. 94) misma (p. Posterior margin of anellus without emargination : F 3 : Sacculus broad at base (PI. 17, fig. 95) : : : serratisignella (p. Sacculus narrow : . : : C : Sacculus short, hardly eaendae forond hacne ‘ lea. ea (p. Sacculus long, extending well beyond harpe : Narrow distal portion of aedeagus clearly separated a bulbous base heligmatodes (p. Narrow distal portion of aedeagus not clearly separated from bulbous base tamariciella (p. Uncus pointed (Pl. 20, fig. 104) ‘ : : ‘ é acutivalva (p. Uncus more or less truncate Harpe a short process Harpe absent Ductus ejaculatorius about half as long as abdomen : heluanensis (p. Ductus ejaculatorius about as long as abdomen . Apical portion of aedeagus strongly curved (Pl. 13, fig. 83) : rufipuncta (p. 98 42 (39) K. SATTLER Apical portion of aedeagus gently curved . ; : longiductella (p. Apical portion of aedeagus strongly curved, twice length of bulbous base roseosuffusella (p. Apical portion of aedeagus gently curved, not longer than bulbous base zonella (p. KEY TO THE SPECIES OF ORNATIVALVA: FEMALES (Note. The 9 of curvella is unknown.) South African species : : P ‘ : : kalahariensis (p. Palaearctic species : : Metascutum with short broad scales (Text: Ae: 1) 6 Metascutum with long narrow scales (Text-fig. 2) Frenulum with two setae (Text-fig. 12) Frenulum with three setae (Text-figs 3, 5-7, 9) . < . : : : Fore wing white with black longitudinal band (PI. 1, fig. 13; Pl. 4, fig. 29); frons with strong process (Pl. 12, figs 78-80) . : ; casas ape (p. Fore wing ochreous or grey; frons with or without process : Fore wing ochreous with dark longitudinal band (PI. 1, fig. 10; Pl. A; fee 30) mongolica (p. Fore wing without longitudinal band (Pl. 1, fig. 9; Pl. 4, fig. 28) mixolitha (p. Frontal process truncate (Pl. 11, figs 69-74) ‘ : : . afghana (p. No frontal process developed : : ; ‘ : é Fore wing ochreous with broad longitudinal band , cerostomatella (p. Fore wing without distinct longitudinal band : : 2 angulatella (p. Frons with process or enlarged scale bases (Pls 6, 7) Frons without modifications. : : < : : Fore wing with basal streak (PI. 1, fig. A; IPA, i 20) : frontella (p. Fore wing without basal streak : : : : Antrum nearly as long as apophysis anterior : : : lilyella (p. Antrum much shorter than apophysis anterior ; : : Vertex with distinct arc of enlarged scale bases (PI. 6, ie 42— a . aspera (p. Enlarged scale bases on vertex not forming arc (Pl. 7, figs 45-47) pulchella (p. Fore wing with basal streak : : : Fore wing without basal streak : F : : ; : . Antrum longer than apophysis anterior : ‘ c erubescens (p. Antrum shorter than apophysis anterior or est! Signum with pair of strong spines Signum with pair of transverse folds or ridges : : : ; Signum. spines at least twice as long as signum is mile . heligmatodes (p. Signum spines hardly longer than their basal plates are wide Antrum with longitudinal fold; apophysis anterior usually dilated aigeiie (Text-figs 15-17) . : ornatella (p. Antrum, when present, without fourteen fold: noeEyoe anterior distally not dilated . Eighth sternite with pair oe longitudinal § folds; aed a share tube (PL. a6, fig. 119) : ; : . singula (p. Eighth sternite without loneacnelice foldss no bake aneeend . Sieversi (p. Eighth tergite with scale bases (Pl. 27, fig. 125) . : acutivalva (p. Posterior margin of eighth tergite with row of setae; no o Seale bases : 104) 104) 105) REVISION OF GENUS ORNATIVALVA No sclerotized antrum (PI. 26, fig. 120) : : : ‘ basistriga (p. Antrum short, tubular Fore wing between costa and fold aniformiy dark brown : arabica (p. Fore wing between costa and fold with light zone (PI. 1, fig. 1; Pl. 2, fig. 17) levifrons (p. Signum with pair of strong spines; spines more or less circular in cross-section . Signum of different shape or spines not circular in cross-section Signum spines longer than signum is wide (PI. 22, fig. 110) ochraceofusca (p. Signum spines as long as signum is wide or shorter Ductus bursae much longer than abdomen Ductus bursae not longer than abdomen Apophysis anterior rod-like; antrum as long as or enees eae Bpopliyais anterior (Pl. 21, fig. 106) : : rufipuncta (p. Apophysis anterior not rod-like; bere absent or shorter than apophysis anterior Apophysis anterior ceased: siete absent : : ‘roseosuffusella (p. Apophysis anterior broad; antrum shorter than apophysis anterior Antrum short, tubular : : Antrum absent, when present not pupae : Base plates of signum separate or connected by narrow selerauced bridge. Base plate of signum broad, hardly constricted medially . triangulella (p. Antrum more than half length of apophysis anterior (Pl. 23, fig. 112) ignota (p. Antrum at most half length of (a anterior ‘ : sesostrella (p. Apophysis anterior reduced : , : : : . xzonella (p. Apophysis anterior clearly developed | ‘ : : : : : Antrum absent (Pl. 21, fig. 105) : : . : . heluanensis (p. Short antrum present : 3 : : plutelliformis (p. Single signum without pair of ae or processes Two separate signa or signum with paired processes : : : : Signum a narrow transverse band without spines ‘ . tamariciella (p. Signum longitudinal, large, with small spines 5 é macrosignella (p. Posterior third of ductus bursae straight, not coiled, weakly sclerotized Entire ductus bursae coiled, membranous Apophysis anterior almost as long as apophysis een (PL. 25, fie. I 17) misma (p. Apophysis anterior much shorter than apophysis posterior . pharaonis (p. Cervix bursae with one or two sclerotized folds near opening of corpus bursae . Cervix bursae or ductus bursae without sclerotizations near ae of corpus bursae. Antrum longer feta apophysis estos: cervix pee oh one selerouized fold (Pl. 24, fig. 115) . : . undella (p. Antrum absent; cervix bursae mich ine Peleratiaed Ke (Pl. 24, fig. 114) Signum a pair of irregular base plates with transverse folds or ridges Signum of different shape Antrum medially at posterior margin eat wae ef faneeninal fale antipyramis (p. Antrum not developed. : : : : é . grisea (p. Signum a narrow sclerotized pad Signum not band-like Signum with pair of folds 3 : F 3 pseudotamariciella (p. longiductella (p. plicella (p. Too K. SATTLER - Signum with pair of broad terminal teeth . : ; . indica (p. 41 (39) Signum with pair of strong spines; edges of spines ae . heligmatodes (p. - Signum with pair of processes with serrated edges 42 (41) Processes of signum serrated along one edge 2 : 2 caecigena (p. - Processes of signum serrated along both edges. : serratisignella (p. CHECK-LIST OF THE SPECIES OF ORNATIVAEVA ORNATIVALVA Gozmany, 1955 PELOSTOLA Janse, 1960 heluanensis-group heluanensis (Debski, 1913) frankeniivorella Chrétien, 1917 oasicolella Turati, 1924, syn. n. siculella Mariani, 1937, syn. n. longiductella Sattler, 1967 rufipuncta sp. n. roseosuffusella Sattler, 1967 zonella (Chrétien, 1917) cimelion Amsel, 1935, Syn. n. wanella Sattler, 1967, syn. n. erubescens-group levifrons sp. n. erubescens (Walsingham, 1904) lilyella (Lucas, 1944) aspera sp. n. pulchella sp. n. ochraceofusca Sattler, 1967 frontella sp. n. arabica Sattler, 1967 sesostrella (Rebel, 1912) ignota Sattler, 1967 ornatella-group ornatella Sattler, 1967 plicella-group plicella sp. n. undella sp. n. tamariciella-group indica Sattler, 1967 pharaonis Sattler, 1967 misma sp. n. serratisignella Sattler, 1967 caecigena (Meyrick, 1918) macrosignella Sattler, 1967 heligmatodes (Walsingham, 1904) tamariciella (Zeller, 1850) pseudotamariciella Sattler, 1967 kalahariensis (Janse, 1960) plutelliformis-group triangulella Sattler, 1967 antipyramis (Meyrick, 1925) 120) 124) 42 123) 122) REVISION OF GENUS ORNATIVALVA IoI singula Sattler, 1967 basistriga sp. n. plutelliformis (Staudinger, 1859) olbiaella Milliére, 1861 siewersiellus Christoph, 1867 sinuatella Walsingham, 1904 grisea Sattler, 1967 sieversi (Staudinger, 1871) , cerostomatella-group mixolitha mixolitha (Meyrick, 1918) mixolitha bipunctella Sattler, 1967 angulatella (Chrétien, 1915) nigrosubvittatella Lucas, 1933 cornifrons sp. n. cerostomatella (Walsingham, 1904) tvipartitella Mabille, 1907 cevostomella Meyrick, 1925 biclavata Meyrick, 1934 mongolica Sattler, 1967 afghana Sattler, 1967 curvella sp. n. acutivalva sp. n. THE HELUANENSIS-GrRovup (Text-fig. 3) Head without enlarged scale bases or frontal process. Metascutum with paired group of narrow, hair-like scales (Text-fig. 2). Fore wing without basal streak on inner margin. Frenulum of 9 triple. In ¢ genitalia uncus reduced; costa and valva present, harpe short or absent; sacculus not separated from valva. Anellus membranous. Aedeagus of some species with very long ductus ejaculatorius. In @ genitalia posterior margin of eighth tergite with irregular row of long setae. Apophysis anterior short or reduced, in vufipuncta longer. Sclero- tized antrum generally absent, in longiductella short, in rvufipuncta well developed. Ductus bursae extremely long in some species. Signum not divided, with pair of strong spines. BroLtocy. O. heluanensis and zonella have been bred on Tamarix species, heluanensis also on Frankenia. Ornativalva heluanensis (Debski, 1913) (Pl, 13) fig. Su; Pl. ax; figs 105; Text-fig. 3) Teleia heluanensis Debski, 1913, in Andres, Bull. Soc. ent. Egypte 3: 111. Lectotype gj, Ecyprt: Helwan, larva on Tamarix, 6. iv., moth emerged 4.v.1910 (Debski) (genitalia slide no. 3063; NM, Vienna), designated by Sattler (1967 : 88) [examined]. [Teleia tamariciella (Zeller); Rebel, 1912 : go (partim). Misidentification.] [Teleia tamariciella (Zeller); Chrétien, 1917 : 473. Erroneously cited as tamariciella Mann. Misidentification. ] Teleia frankeniivorella Chrétien, 1917, Annls Soc. ent. Fr. 85 :474. Lectotype 9, ALGERIA: Biskra, larva on Frankenia, 11.-12.xi.191I, moth emerged 27.iii.1912 (Chrétien) (genitalia 102 1a sata) INGA BIR slide no. 387a, Sattler; MNHN, Paris), designated by Sattler (1967 : 87) [examined]. [Synonymized by Sattler, 1967 : 39.] Teleja [sic] oasicolella Turati, 1924, Atti Soc. ital. Sci. nat. 63 : 161, pl. 6, fig. 4 [poor colour photograph]. LECTOTYPE 9, Lisya: Cyrenaica, Bengasi, Saluk, 24. iv. (Kviiger) (genitalia slide no. 685, Sattler; coll. Turati, IE, Bolzano), here designated [examined]. Syn. n. Gelechia frankeniivorella (Chrétien) Meyrick, 19254 : 78. Gelechia heluanensis (Debski) Meyrick, 1925a : 84. [Teleia tamaricalis (Zeller); Amsel, 1933 : 125 (partim). Incorrect subsequent spelling of tamariciella Zeller. Misidentification. } [Teleta tamariciella (Zeller); Amsel, 19356 : 263 (partim). Muisidentification. | Telphusa oasicolella (Turati) Gaede, 1937 : 131. Gelechia frankenivorella (Chrétien); Gaede, 1937 : 173. Gelechia heluanensis (Debski); Gaede, 1937 : 177. Lita siculella Mariani, 1937, G. Sci. nat. econ. Palermo 39(3) : 9, pl. 1, fig. 1. LECTOTYPE g, IraLy: Sicily, Prov. Palermo, Casteldaccia, 23.ix.1927 (Mariani) (genitalia slide no. 631, Povolny; MNHU, Berlin), here designated [examined]. Syn. n. [Teleia tamariciella (Zeller); Amsel, 1955a : 28. Misidentification. | [Teleia tamariciella (Zeller); Amsel, 1959 : 32. Misidentification. | [Ornativalva trifasciella (Rebel) Sattler, 1960 : 59. Miusidentification. | Ornativalva frankenivorella (Chrétien) Sattler, 1964 : 577. Ornativalva frankeniivorella (Chrétien); Amsel, 1966 : 128. Ornativalva heluanensis (Debski) Sattler, 1967 : 38, 88, pl. I, fig. 5, pl. 3, figs 15, 16, pl. 9, fig. 45. Ornativalva heluanensis (Debski); Amsel, 1968 : 17. GENITALIA g. PI. 13, fig. 81. GENITALIA 9. PI. 21, fig. 105. REMARKS. The observations of Debski (1913: 110-112) on the larvae are incorrect or unreliable. He observed two different types of larvae, both feeding on Tamarix tetragyna Ehrenberg and 7. nilotica (Ehrenberg) Bunge. The larva described on the specimen labels as ‘larva libera bicornuta’ he attributed to tamari- ciella (Debski, 1913 : 110); however, from his detailed description it is clear that this was not a gelechiid larva but probably that of an A gdistis species (Pterophoridae) (Chrétien, 1917 : 473). The second larval type, described as ‘larva tubicola’, Debski attributed to heluanensis. I have examined five of Debski’s specimens. One ‘ex larva libera bicornuta’ and two ‘ex larva tubicola’ are heluanensis, while one ‘ex larva libera bicornuta’ and one ‘ex larva tubicola’ are macrosignella. Chrétien (1917 : 473, 474) separated two species, tamariciella and frankeniworella, based on different host-plants (Tamarix species and Frankenia pallida Boissier) as well as on certain differences in the colour of the larvae and the wing markings of the moths. I have examined the lectotype 9 of frankeniivorella and a 3 which Chrétien reared on Tamarix; both represent the same species. While the differences in the moths can be explained by the considerable variability of wing markings and colour in heluanensis, not enough is known about the larvae to account for their colour differences; however, considerable variation in colour has been observed in the larvae of O. plutelliformis. Teleja [sic] oasicolella Turati was described from an unspecified number of specimens. The Turati collection contained one g and one 9. The @ bears the label ‘oasicolella H{olo] Type.’, probably in Turati’s handwriting. Although the colour photograph in the original publication is of poor quality, it seems certain that REVISION OF GENUS ORNATIVALVA 103 it illustrates this particular specimen, which is here designated as the lectotype. The ¢ bears (?) Turati’s label ‘oasicolella All{o|-Type.’ and has now been labelled as paralectotype. Lita siculella Mariani was described from one specimen in coll. Mariani and one specimen in MNHU, Berlin. I was unable to obtain the specimen from Mariani’s collection which is still in the hands of his family; however, the specimen from MNHU, Berlin, was available and is here designated as the lectotype. The specimen bears the label ‘Lita siculella Mariani’, in Mariani’s handwriting. Mariani stated that the species was collected in August. On the data label of the lectotype the printed ‘VIII.’ has been altered by hand to ‘23.1 X.’ and it seems likely that Mariani had sent this specimen to Berlin without recording the exact date for his publication. Biotocy. Host-plants: Frankenia pallida Boissier (moth bred by Chrétien); Frankenia species (moth bred by Walsingham); Tamarix tetragyna Ehrenberg and T. nilotica (Ehrenberg) Bunge (moths bred by Debski); unspecified species of Tamarix (moths bred by Chrétien, Dumont, Gerling and Lupo). As far as can be seen from the literature (Debski, 1913 : 111; Chrétien, 1917 : 473, 474) and information from data labels of specimens, the larva lives in a silken tunnel which is covered with the deciduous terminal shoots of Tamarix or other material. The larva feeds on the terminal shoots of Tamarix or mines the succulent leaves of Frankenia which then turn white. Pupation takes place in a cocoon on the ground. The number of generations per year is unknown. There may be two clear-cut _ generations in southern Europe or Central Asia; however, there are indications that heluanensis breeds continuously under favourable conditions. In North Africa larvae have been observed by various collectors in April-June, November and December whereas moths have been collected in all months of the year. DISTRIBUTION. Spain; Italy (Sicily); Malta; Yugoslavia; U.S.S.R. (S. Russia, Turkestan); Turkey; Cape Verde Islands (St. Vincent); Canary Islands (Gran Canaria); Morocco; Algeria; Tunisia; Libya; Egypt; Sudan; Israel; Syria; Saudi Arabia; Iraq; Iran; Afghanistan; Pakistan; Mongolia. FURTHER MATERIAL EXAMINED. SPAIN: 2 3, 15 2, Prov. Almeria, (Cabo de Gata) Mazarulleque, 3.vili.1968 (Sattler & Carter) (BMNH); 1 4, 8 9, Prov. Murcia, Alhama de Murcia, 19.—20.1x.1974 (Glaser) (coll. Glaser, Vienna; BMNH) [first record for Spain]. MALta: 1 9, Gozo - Xlendi, 3.1x.1956 (de Lucca) (coll. de Lucca, Malta); 1 2, Melleha, 9.ix.1956 (de Lucca) (coll. de Lucca, Malta) [first record for Malta]. TuRKEy: 1 g, Esan river, 50 km E. of Fethiye, 20.vili.1972 (Gerling) (TAU, Tel-Aviv); 1 g, 50km W. of Silifke, 17.vili.1972 (Gerling) (TAU, Tel-Aviv) [first record for Turkey]. CANARY ISLANDS: 3 6, 2 9, Gran Canaria, Maspalomas, 26.—27.11.1967 (Kasy) (NM, Vienna; BMNH) [first record for the Canary Islands]. Lipya: 1 g, Cyrenaica, Bengasi, 30.iv.1922 (Kriiger) (IE, Bolzano) (paralectotype of T. oasicolella Turati). ISRAEL: 2 4, 4 8, Jifthk Post, 35 km N. of Jericho, larvae in webs on Tamarix, 3.vi., moths emerged 4.Vil.—15.viil.1971 (Lupo) (TAU, Tel-Aviv); 1 3, Nahal Ze’elim, 12 km S. of Ein Gedi, larva in web on Tamarix, 27.xii.1970, moth emerged 17.1.1971 (Gerling) (TAU, 104 KK. SAd PEER Tel-Aviv); 1 g, Ein-Feshcha, 22.ix.1971 (Kugler) (TAU, Tel-Aviv); 1 g, Nahal David, 26.ix.1g71 (Kugler) (TAU, Tel-Aviv). AFGHANISTAN: I g, Ghorband Valley, N. of Kabul, 1900 m, 30.vii.1965 (Kasy & Vartian) (NM, Vienna). MONGOLIA: 3 dg, 3 9, Bajanchongor aimak, oasis Echin gol, 90 km NE. of borderpost Caganbulag, 950m, 27.-29.vi.1967 (Kaszab, no. 857) (TM, Budapest); 3 g, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (TM, Budapest; BMNH); 1 9, South Gobi aimak, 10 km NNE. of Dalanzadgad, 1450 m, 7.vii.1967 (Kaszab, no. 898) (TM, Budapest) [first record for Mongolia]. Ornativalva longiductella Sattler, 1967 (Pll 13; te: 82) Ornativalva longiductella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26(3) : 40, pl. 3, fig. 17, pl. 9, fig. 46. Holotype 9, ALGERIA: Hammam-es-Salahin, 15.iv.1904 (Walsingham) (genitalia slide no. 6563; BMNH) [examined]. Ornativalva longiductella Sattler, 1964 : 578. Nomen nudum. GENITALIA Gg. PI. 13, fig. 82. GENITALIA Q. Sattler, 1967, pl. 9, fig. 46. BroLocy. MHost-plant unknown. Moths have been taken by various collectors in February—April, June, July and October. DistripuTion. Algeria; Tunisia; Sudan; Iraq; Afghanistan; Mongolia. FURTHER MATERIAL EXAMINED. MONGOLIA: 5 4, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.v1.1967 (Kaszab, no. 834) (TM, Budapest; BMNH); 1 3, Bajanchongor aimak, spring Talyn Bilgech Bulag, 47 km E. of borderpost Caganbulag, 1200 m, 23.V1.1967 (Kaszab, no. 840) (IM, Budapest); 3 g, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (TM, Budapest; BMNBH) [first record for Mongolia}. Ornativalva rufipuncta sp. n. (Pl. 2, ue. 16; Pl 13, he. 82° Pl 25 ie. Foo) 3, Q. 5:0-6:0mm. Head without frontal modifications. Frons pale, whitish grey, scales on vertex mixed with brown. Labial palpus whitish, outer surface of second segment with brown band near base and before apex; third segment with broad brown band around middle, apex brown. Antenna brown with white rings above, white below. Thorax and tegula greyish, apex of tegula whitish. Fore wing greyish with red-brown markings. Two red-brown dots in cell, double dot at end of cell, short streak in fold at base; three red-brown dots on fold, at one-third, two-thirds and end of fold. Narrow red-brown zone along base of fringes. Outer transverse line sharply angulate in middle. Fringes whitish, divided by several grey lines. GENITALIA ¢ (Pl. 13, fig. 83). Tegumen with deep anterior emargination, sclerotized part narrow. Costa as long as valva. Harpe short. Basal two-thirds of valva broad, distal third much narrower, curved, tapering, without apical spine. Basal third of aedeagus oval, apical two-thirds much narrower, strongly curved. GENITALIA 9 (Pl. 21, fig. 106). Apophysis posterior slightly longer than apophysis anterior. REVISION OF GENUS ORNATIVALVA 105 Sclerotized antrum as long as apophysis anterior. Ductus bursae coiled, extended about three times length of abdomen. Corpus bursae small, globular. Signum long, irregular, sclerotized plate with pair of strong spines. RemMARKS. O. rufipuncta differs externally from all other Oynativalva species by the distinct red-brown fore wing markings. Similarities in the 2 genitalia, parti- cularly the extremely long ductus bursae, place rufipuncta near longiductella. BioLtoGy. Host-plant unknown. The type-series was taken in association with other species known to feed on Tamarix. Moths have been collected at light in April. DIsTRIBUTION. S. Iran (Luristan). MATERIAL EXAMINED. Holotype J, Iran: S. Iran, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 15.iv.1972 (Exped. Mus. Vind.) (NM, Vienna). Paratypes. IRAN: 1 J, S. Iran, [Luristan,] 8 km E. of Bandar-Abbas, 23.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 6 3, 12 9, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 10., 15.1v.1972, 21.1v.-17.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 3 6, 1 Q, [Luristan,] 22 km E. of Bandar-Abbas, Ig. iv., 10.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 1 dg, 3 9, [Luristan,] 30 km E. of Bandar-Abbas, 3.1v.1970, 8.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 3 9, [Luristan,] 13 km NE. of Bandar-Abbas, 6.iv.1972, 24.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 3 dg, 2 9, [Luristan,] 22 km N. of Bandar-Abbas, 9.iv.1972, 18., 25.iv.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 1 9, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna). Ornativalva roseosuffusella Sattler, 1967 Ornativalva voseosuffusella Sattler, 1967, Beitr. naturk. Forsch. SiidwDitl. 26(3) : 42, pl. 3, fig. 18, pl. 9, fig. 47. Holotype @, Iran: W. Iran, Khuzistan, Shadegan, 1.-8.iii.1956 (Richter) (genitalia slide no. 298b, Sattler; SMN, Stuttgart) [examined]. GENITALIA g. Sattler, 1967, pl. 3, fig. 18. Genitatia 9. Sattler, 1967, pl. 9, fig. 47. BioLtocy. Host-plant unknown. The type-specimens have been collected in March and April. DistrisuTion. W. Iran (Khuzistan). Ornativalva zonella (Chrétien, 1917) Teleia zonella Chrétien, 1917, Annis Soc. ent. Fr. 85 : 474. Lectotype 9, Tunisia: Gafsa, larva on Tamarix, 21.vi., moth emerged 3.v.1909 (Chrétien) (genitalia slide no. 370c, Sattler; MNHN, Paris), designated by Sattler, 1967 : 88 [examined]. Gelechia zonella (Chrétien) Caradja, 1920 : 99. Gelechia zonella (Chrétien) ; Meyrick, 1925a : 78. 106 Ke SAG Wer Teleia cimelion Amsel, 1935a, Veréff. dt. Kolon. u. Ubersee-Mus. Bremen 1 : 210, pl. 11, figs 14, 15. Type 9, IsRAEL: south end of Dead Sea, 15.—27.1ii.1933 (Aigner) (location of type unknown) [not examined]. Syn. n. ’ Gelechia cimelion (Amsel) Gaede, 1937 : 154. Gelechia zonella (Chrétien) ; Gaede, 1937 : 231. Ornativalva cimelion (Amsel) Sattler, 1960 : 59. Oynativalva ivanella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26(3) : 44, pl. 3, fig. to. Holotype g, IRAN: S. Iran, [Kerman,] Jiroft (“Djiroft’), Anbar-Abad, 21.-30.iv.1956 (Richter) (genitalia slide no. 297b, Sattler; SMN, Stuttgart) [examined]. Syn. n. Oynativalva zonella zonella (Chrétien) Sattler, 1967 : 44, 88, pl. 1, fig. 6, pl. 3, fig. 20, pl. 9, fig. 48. Ornativalva zonella cimelion (Amsel); Sattler, 1967 : 45, pl. 1, fig. 7, pl. ro, fig. 49. GENITALIA g. Sattler, 1967, pl. 3, figs 19, 20. GENITALIA 9. Sattler, 1967, pl. 9, fig. 48, pl. 10, fig. 49. ReMARKS. The examination of fresh material and the previously studied specimens has led‘me to the conclusion that the separation of cimelion as a subspecies of zonella is not justified and that ¢vanella is only a form of zonella. T. cimelion was described from “Typen und Cotypen, 399’. I have been unable to locate a specimen labelled ‘Typus’; however, two ‘Cotypus’ 9 have been examined. BioLtocy. Host-plant: Tamarix species (moths bred by Chrétien). According to Chrétien (1917 : 475) the larva lives in a small silken tube between two or three twigs of Tamarix. It is fully grown in November and pupates on the ground in a cocoon which is covered with particles of soil. For a description of the larva see Chrétien (loc. cit.). Moths have been taken by various collectors between March and June. DIsTRIBUTION. Algeria; Tunisia; Israel; Saudi Arabia; S. Iran (Kerman, Luristan). FURTHER MATERIAL EXAMINED. IRAN: 1 gf, 1,5. Iran, [Luristan,] 17 km E. of Bandar-Abbas, dunes, I0., 15.1v.1972 (Exped. Mus. Vind.) (NM, Vienna); 2 3, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna). THE ERUBESCENS-GRoOuP (Text-fig. 4) Head with or without enlarged scale bases and frontal process. Metascutum with paired group of narrow, hair-like scales (Text-fig. 2). Fore wing with or without basal streak on inner margin. Frenulum of 9 triple. In g genitalia uncus reduced; costa and valva present, harpe digitate or clavate, in ignota absent. Sacculus partially separated from valva. Anellus membranous or anellus lobes weakly sclerotized. In Q genitalia posterior margin of eighth tergite with irregular row of long setae. Apophysis anterior well developed, rod-like. Sclerotized antrum developed. Signum single or divided into two, with pair of strong spines or transverse tooth-like ridges. BroLocy. Several species have been bred on Tamarix. REVISION OF GENUS ORNATIVALVA 107 Ornativalva levifrons sp. n. Semen ae. 1; Fl. 2 tie. 17; PI 14, fig. 84; PI. 21, fig. 107; Text-fig. 22) 3, 2. 6:0-7-omm. Head without enlarged scale bases or frontal process. Vertex pale, without dark longitudinal line. Labial palpus pale, outer surface of second segment dark brown near base, ochreous at apex; third segment light ochreous above, brown below. Thorax ochreous, laterally brown. Tegula brown with light apex. Anterior half of fore wing pale ochreous with brown markings. In fresh specimens lighter area suffused with pink. Posterior half of fore wing ochreous with short, dark brown basal streak, longer streak along basal half of fold, patch behind middle of fold and patch near end of cell. GENITALIA ¢ (Pl. 14, fig. 84). Lateral margins of uncus parallel, posterior margin slightly rounded, with number of setae. Anterior margin of tegumen with deep square emargination. Harpe short, robust, densely set with setae. Sacculus reaches middle of valva. Basal third of aedeagus bulbous, apical two-thirds slender, strongly curved before apex. GENITALIA Q (Pl. 21, fig. 107). Antrum large, funnel-shaped, anterior portion narrow. Ductus bursae three to four times length of apophysis posterior, wider than narrow anterior portion of antrum. Signum in posterior part of corpus bursae, near entrance of ductus bursae, composed of pair of weakly sclerotized basal plates with transverse ridge; basal plates connected by narrow sclerotized bridge. Posterior margin of seventh sternite with shallow median emargination (Text-fig. 22). REMARKS. (O. levifrons is closely related to erubescens and lilyella, differing from erubescens by the dark markings of the fore wing which are not strongly angulate at the fold, the short harpe, the shape of the aedeagus, the antrum and the signum with sclerotized bridge. O. levifrons differs from lilyella by the absence of a frontal process of the head, the presence of a dark basal streak on the fore wing, the short robust harpe, the long ductus bursae and the weaker signum with a narrower sclerotized bridge. O. levifrons differs from arabica by the presence of a light area between costa and fold on the fore wing. In the ¢ genitalia the uncus is shorter, the harpe more globular and the saccus narrower. The 2 genitalia of levifrons are very similar to those of avabica but the tubular part of the antrum is narrower and the sclerotized bridge between both parts of the signum is very narrow; however, the width of the bridge is variable in arabica. The 9 genitalia of levifrons are also very similar to those of lilyella, aspera and pulchella, all of which are distinguished externally by the frontal processes of the head. O. levifrons is the only Mongolian species of the erubescens-group without modified head structures. BrioLtocy. Host-plant unknown. Moths have been taken by Kaszab between 23 June and 5 July. DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Moncoria: Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (TM, Budapest). Paratypes. Moncoira: 2 g, Uburchangaj aimak, 130km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (ITM, Budapest; BMNH); 1 @, Bajanchongor aimak, spring Talyn Bilgech bulag, 47 km E. of borderpost Cagan- bulag, 1200 m, 23.vi.1967 (Kaszab, no. 840) (TM, Budapest); 1 9, Bajanchongor aimak, oasis Dzun mod, 100 km S. of Somon Schine zinst, 1300 m, 29.v1.1967 108 Ke SA ie ik (Kaszab, no. 869) (BMNH); 1 3, Chovd aimak, 10 km SSW. of Somon Bulgan, 1200 m, 5.vii.1966 (Kaszab, no. 633) (IM, Budapest). Ornativalva erubescens (Walsingham, 1904) (Pl. 5, figs 36-38; Text-fig. 4) Gelechia erubescens Walsingham, 1904, Entomologist’s mon. Mag. 40 : 265. Lectotype 9 [not gf as stated by Walsingham], ALGERIA: Biskra, 11.111.1903 (Walsingham) (genitalia slide no. 6145; BMNH), designated by Sattler (1967 : 47) [examined]. Gelechia evubescens Walsingham; Meyrick, 1925a : 79. [Gelechia plutelliformis Staudinger; Amsel, 19350 : 263 (partim). Misidentification.] Gelechia evubescens Walsingham; Gaede, 1937 : I7I. Ornativalva ervubescens (Walsingham) Sattler, 1964 : 578. Ornativalva evubescens (Walsingham); Amsel, 1966 : 128. Ornativalva erubescens (Walsingham) ; Sattler, 1967 : 45, 47, pl. 1, fig. 8, pl. 4, figs 21a—c, 22, pl. Io, fig. 50. Head (PI. 5, figs 36-38) without enlarged scale bases or frontal process. GENITALIA g. Sattler, 1967, pl. 4, figs 21a—c, 22. GENITALIA 9. Sattler, 1967, pl. 10, fig. 50. RemARKS. The lectotype 9 was erroneously recorded by Walsingham as ‘type 3’ (no. 96 595). The ‘type 9’ (no. 96 596) is not conspecific with the lectotype but belongs to O. ignota. Brotocy. Host-plant unknown. Moths have been taken by various collectors in January, May, August and November. DisTRIBUTION. Morocco; Algeria; Tunisia; Libya; Egypt; Sudan; Israel; Saudi Arabia; Iran; Pakistan. FURTHER MATERIAL EXAMINED. PAKISTAN: I 9, 150 km SW. of Quetta, goo m, 13.v.1965 (Kasy & Vartian) (NM, Vienna) [first record for Pakistan]. Ornativalva lilyella (Lucas, 1944) (Pl. 6, figs 39-41) Gelechia lilyella Lucas, 1944, Bull. Soc. ent. Fry. 48 : 135. Holotype g, ALGERIA: El Goléa, 21.11.1939 (Lucas) (genitalia slide no. 581a, Sattler; MNHN, Paris) [examined]. Ornativalva lilyella (Lucas) Sattler, 1967 : 48, pl. 4, fig. 21d, 23, pl. 10, fig. 51. Head (Pl. 6, figs 39-41) with arc of irregularly arranged teeth above transfrontal sulcus. Short thick frontal process truncate, dorsally and laterally smooth, without teeth or scale bases, anterior surface densely set with teeth; some big teeth scattered in area below frontal process. GENITALIA g. Sattler, 1967, pl. 4, figs 21d, 23. GENITALIA 9. Sattler, 1967, pl. 10, fig. 51. REMARKS. The head structure resembles that of aspera; however, the frontal process has a wider diameter than that of aspera and the frons is not so densely REVISION OF GENUS ORNATIVALVA 109 set with teeth. The 9 genitalia are very similar to those of levifrons, aspera and pulchella. BioLtocy. Host-plant unknown. Moths have been collected by Dumont and Lucas in February—April and October. DISTRIBUTION. Algeria. Ornativalva aspera sp. n. (Pipd, fig. 2; Pl. 2, fig. 18; Pl. 6, figs 42-44; Pl. 14, fig. 85; Pl. 22, fig. 108) 3, 2. 5:0-6:5mm. Head (PI. 6, figs 42-44) with arc of irregular teeth above transfrontal sulcus. Frontal process short, truncate, dorsal surface smooth, without teeth or scale bases, frontal and ventral surfaces covered irregularly with enlarged scale bases. rons pale, vertex light brown, dark brown around ocellus. Thorax light brown, along lateral margins dark brown. Tegula dark brown. Labial palpus whitish; outer surface of second segment light brown at base and before apex; third segment brown below. Antenna dark brown, above with light rings, whitish below. Fore wing with basal two-thirds of anterior half dark brown, in places lighter towards costa; posterior half light brown, with short dark basal streak. Dark brown area of wing angulate at fold, with whitish tooth extending deep into area between dark angles; in some places dark angles lined with white scales. Apical third of wing light brown, with dark shadow on costa near apex; whitish patch behind dark dot at end of cell. Base of fringes with indistinct dark line. GENITALIA ¢ (PI. 14, fig. 85). Lateral margins of uncus parallel. Anterior margin of tegumen with square emargination. Harpe digitate, about half length of costa. Apical half of valva curved, pointed, but not ending in distinct spine. Terminal quarter of sacculus free, not fused with valva; inner surface near middle with distinct projections which point towards costa. Dorsal margin of sacculus with long setae. Genitatia Q (Pl. 22, fig. 108). Antrum large, funnel-shaped, anterior portion narrow. Ductus bursae approximately twice length of apophysis posterior, half width of narrow anterior _ portion of antrum. Signum in posterior part of corpus bursae, near entrance of ductus bursae, composed of pair of weakly sclerotized basal plates with transverse ridge; basal plates connected by sclerotized bridge of moderate width. REMARKS. O. aspera is closely related to pulchella but is slightly smaller, the frontal process of the head is slightly shorter and a distinct arc of irregular teeth is present above the transfrontal sulcus. In the fore wing aspera differs from pulchella by the short dark basal streak, the light brown area which does not extend to the apex, the whitish tooth between the angles of the dark anterior markings, and the | more distinct black dot at the end of the cell. The ¢ genitalia differ from those of pulchella by the stronger, less curved, terminal portion of the valva and the sacculus which is widest near its middle. In the Q genitalia the setae with strongly curved tips on the posterior margin of the papilla analis are weaker and more numerous, _the sclerotized part of the eighth tergite is narrower, the ductus bursae is shorter | and narrower and the sclerotized bridge of the signum is wider than in pulchella. As only 1 9 of pulchella was available for examination it is impossible to decide _ which of these differences are of significance. The ° genitalia are also very similar to those of levifrons but differ by the shorter and narrower ductus bursae. O. levifrons is externally distinguished by the absence of frontal modifications of the 110 Ko SADIE R head. O. aspera differs from the closely related Jilyella, which is known only from Algeria, by the smaller size, the large number of teeth around the frontal process of the head, the basal streak and the distinct pattern of the fore wing, the longer, more digitate harpe, the sacculus and the shorter antrum. BioLocy. MHost-plant unknown. Moths have been collected by Kaszab between 22 June and 7 July in localities where Tamarix occurs (Kaszab, 1968 : 22, 31). DISTRIBUTION. Mongoha. MATERIAL EXAMINED. Holotype g, Moncoita: South Gobi aimak, 100km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (IM, Budapest). Paratypes. MONGOLIA: 5 4, 3 2, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (TM, Budapest; BMNH); 1 dg, I 2, South Gobi aimak, 10 km NNE. of the town Dalanzadgad, 1450 m, 7.vii.1967 (Kaszab, no. 898) (TM, Budapest; BMNH); 2 3, 4 9, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (TM, Budapest; BMNH). Ornativalva pulchella sp. n. (PL 1, fig. 3; Pl. ‘2, fig. 19; Pl. 7, figs 45-47; Pl. 14, fig. 86; Pl. 22). fig nog} 3, 2. 6:0-7-omm. Head (Pl. 7, figs 45-47) without distinct arc of irregular teeth above transfrontal sulcus. Frontal process truncate, smooth, frontal surface covered with teeth. Frontal area below transfrontal sulcus and frontal process slightly raised, covered with strong irregular teeth. Head pale ochreous, brown along eye. Thorax light brown, lateral margins dark brown. Tegula dark brown. Labial palpus whitish, outer surface of second segment with dark brown patch at base. Antenna dark brown with narrow light rings above, whitish below. Fore wing without basal streak. Anterior half of wing dark brown, markings angulate at fold, posterior half light brown up to apex. Small dark dot at end of cell, sometimes indistinct. Base of fringes with indistinct dark line. GENITALIA g (Pl. 14, fig. 86). Lateral margins of uncus parallel, posterior margin rounded, with setae. Anterior margin of tegumen with square emargination. Harpe slender, digitate. Apical portion of valva slender, strongly curved, apex with short spine. Sacculus triangular, with acute angle pointing towards base of valva; dorsal margin with long setae. Aedeagus with bulbous base, apical two-thirds curved, tapering. GENITALIA 9 (Pl. 22, fig. tog). Antrum funnel-shaped, ductus bursae approximately three times length of apophysis posterior, same width as narrow anterior portion of antrum. Signum in posterior part of corpus bursae, near entrance of ductus bursae, composed of pair of weakly sclerotized basal plates with transyerse ridge; basal plates connected by very narrow sclerotized bridge. REMARKS. 0. pulchella is closely related to aspera; for differences see there. It is also closely related to the Algerian /ilyella from which it differs by the darker colour and the more distinct markings of the fore wing, the more clearly defined frontal process of the head, the triangular sacculus, and the very narrow sclerotized bridge of the signum. The @ genitalia are very similar to those of levifrons but differ by the wider anterior portion of the antrum and the smaller signum. 0. REVISION OF GENUS ORNATIVALVA III levifrons is externally distinguished by the absence of frontal modifications of the head. BroLtocy. Host-plant unknown. The type-series has been collected by Kaszab on 22 June in a locality where Tamarix occurs (Kaszab, 1968 : 22). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Monco.ta: South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (genitalia slide no. 635d, Sattler; TM, Budapest). Paratypes. MONGOLIA: 2 3, I 2, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (TM, Budapest; BMNH). Ornativalva ochraceofusca Sattler, 1967 (Pl..£; fig-5; PL. 15, tg, 67; PI. 22,. fig. FL) Ornativalva ochraceofusca Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26(3) :77, pl. 18, fig. 74. Holotype g, AFGHANISTAN: W. of Charikar (N. of Kabul), 1.viii.1963 (Kasy & Vartian) (genitalia slide no. 3277; NM, Vienna) [examined]. Head without enlarged scale bases or frontal process. GENITALIA ¢g (Pl. 15, fig. 87). Valva strongly bent at middle, narrow, distally nearly as wide as at separation point of sacculus. Geniratia Q (Pl. 22, fig. 110). Apophysis posterior as long as eighth segment (including apophysis anterior). Short sclerotized antrum with longitudinal fold. Signum a narrow sclerotized plate with pair of long curved spines. Posterior margin of seventh segment with deep median emargination. REMARKS. O. ochraceofusca was previously placed tentatively in the plutelliformis- group (Sattler, 1967 : 78) but is here transferred to the erubescens-group. In the 3 genitalia the length of the harpe and the shape of the sacculus were found to be variable. In the holotype from Afghanistan the distal half of the costa is club- shaped, while in Turkish specimens it is digitate. The 2 genitalia are very similar to those of ornatella. In ornatella the eighth segment is laterally set with dense rows of microtrichia between the base of the apophysis anterior and the posterior margin of the segment. In ochraceofusca there is only a limited area set with very small microtrichia or they are completely absent. In orvnatella the antrum appears to be longer than in ochraceofusca and the apophysis anterior is distally more clearly dilated. Externally ornatella is characterized by a clear basal streak on the fore wing, while ochraceofusca has some dark scales near the fore wing base without developing a clear basal streak. Although the 3 genitalia are quite distinct there appears to be no reliable character for separating the 9 genitalia of ochraceofusca and ornatella. When examined earlier, the holotype bore only a temporary label, the data of which were cited in the original description (Sattler, 1967: 78). The completed and corrected label data are cited above. Te Ke SAwl ieee BroLtocy. Host-plant unknown. Moths have been taken by various collectors in June, July and August in association with other Lepidoptera species known to feed on Tamarix. DISTRIBUTION. Turkey; Afghanistan. FURTHER MATERIAL EXAMINED. TURKEY: 5 ¢, 49, 10 km NW. of Giimiischane, 1000 m, 10.vi.1969 (Kasy; Aven- berger) (NM, Vienna; coll. Arenberger, Vienna; BMNH); 3 dg, 2 9, 40 km SW. of Elazig, 900 m, 18.vi.1969 (Kasy; Arenberger) (NM, Vienna; coll. Arenberger, Vienna; coll. Jackh, Bidingen; BMNH) [first record for Turkey]. AFGHANISTAN: 3 Q,: Ghorband Valley, N. of Kabul, 1900 m, 30.vil.1965 (Kasy & Vartian) (NM, Vienna; BMNH). Ornativalva species I (PL 15, fig.\88; Pl 23, ig. rn Pext-te, 23) 3, @. 4:0-5:°0mm. Most of the moths are in poor condition. Head without enlarged scale bases or frontal process. Dark W-shaped markings of fore wing crossing fold at one-third and two-thirds. No basal streak. Distinct black dots in cell at distal end of W-markings and at end of cell. GENITALIA ¢ (Pl. 15, fig. 88). Distal third of costa twice as wide as proximal two-thirds. Harpe slender. Valva straight, narrow. Sacculus triangular, widest at middle. Genitalia Q (Pl. 23, fig. 111). Eighth segment laterally with large zone of densely set microtrichia. Apophysis anterior half length of apophysis posterior, slightly dilated distally. Antrum short, funnel-shaped, half length of apophysis anterior. Anterior margin of eighth sternite with rounded projection near antrum. Signum pair of irregularly shaped medially fused sclerotized basal plates with sharp spine. Posterior margins of seventh tergite and sternite with deep median emargination (Text-fig. 23). REMARKS. The ¢ genitalia differ from those of ochvaceofusca by the valva which is straight, not strongly bent at middle, and the sacculus which is widest at middle, not in distal half. The 9 genitalia differ from those of ochvaceofusca by the dense area of microtrichia on the eighth segment, the longer antrum, the rounded anterior margin of the eighth sternite, the much weaker base of the signum, the strong signum spines and the distinct median emargination of the seventh sternite. The fore wing base is light, without basal streak; in ochraceofusca it is dark, but no basal streak is separated; in ornatella a distinct basal streak is present. BroLtocy. Host-plant unknown. Moths have been collected by Kaszab in June in localities where Tamarix occurs (Kaszab, 1968 : 22, 23, 26-27). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. MoNnGOLlIaé: I g, South Gobi aimak, Z66l6n ul, 58 km WSW. of Somon Bajandalaj, 1500 m, 16.vi.1967 (Kaszab, no. 807) (TM, Budapest); 2 g, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (TM, Budapest; BMNH); 2 9, Bajanchongor aimak, spring Talyn Bilgech bulag, 47 km REVISION OF GENUS ORNATIVALVA 113 E. of borderpost Caganbulag, 1200 m, 23.vi.1967 (Kaszab, no. 840) (TM, Budapest); 3 6, Bajanchongor aimak, oasis Echin gol, 90 km NE. of borderpost Caganbulag, 950 m, 27.—29.vi.1967 (Kaszab, nos 857, 858) (IM, Budapest); 1 3, 1 9, Bajanchongor aimak, oasis Dzun mod, 100 km S. of Somon Schine zinst, 1300m, 29.vi.1967 (Kaszab, no. 869) (TM, Budapest; BMNH). Ornativalva species 2 (Pl. 15, fig. 89) Head without enlarged scale bases or frontal process. GENITALIA ¢ (Pl. 15, fig. 89). Valva narrow, straight, distally rounded, narrowest at three- quarters. Sacculus narrow, not triangular. GENITALIA 9. Unknown. REMARKS. Externally like species 1; differing in the genitalia by the distally dilated valva and the narrow sacculus. BroLtocy. Host-plant unknown. The moths have been collected by Kaszab in June in localities where Tamarix occurs (Kaszab, 1966 : 591; 1968 : 22). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. MONGOLIA: I g, Gobi Altaj aimak, Zachuj Gobi, 1o km N. of Chatan chajrchan mountains, 1150 m, 27.vi.1966 (Kaszab, no. 594) (TM, Budapest); 1 g, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (IM, Budapest). Ornativalva species 3 (Pl. 16, fig. go) Head without enlarged scale bases or frontal process. GENITALIA ¢ (Pl. 16, fig. 90). Valva broad at base, not strongly bent, distal third narrow. GENITALIA 9. Unknown. REMARKS. Like species 1 and 2 but slightly bigger, fore wing between fold and inner margin distinctly ochreous. In the ¢ genitalia the sacculus is similar to that _ of species 2 but the distal portion of the valva is not dilated. __ Brotocy. Host-plant unknown. The only specimen has been collected by _ Kaszab in June. DISTRIBUTION. Mongolia. MATERIAL EXAMINED. MONGOLIA: I 3, Bajanchongor aimak, Cagan Bogd ul, Tooroin bulag, 13 km E. of borderpost Caganbulag, 1500 m, 25.vi.1967 (Kaszab, no. 849) (TM, Budapest). 114 K. SATILVER Ornativalva frontella sp. n. (Pl. 1, fig. 43 PL. 2, fig. 20; Pl. 7, tigs-48—50; Plax, fie ox) 3, 2. 6:0-7-0mm. Head (Pl. 7, figs 48-50) with dorsal surface of strongly curved arc above transfrontal sulcus almost completely free of scales. Vertex without enlarged scale bases. Strong frontal process below transfrontal sulcus truncate, anterior surface covered with enlarged scale bases. Frons below transfrontal sulcus with numerous enlarged scale bases arranged in irregular rings. Head pale ochreous, brown along eye. Labial palpus whitish, outer surface with scattered light brown scales; third segment brown, lighter above. Antenna dark brown with lighter rings; light below. Thorax light brown, laterally dark brown. Tegula dark brown. Light anterior and dark posterior half of fore wing divided by black longitudinal markings. Anterior half of wing suffused with pink, costa lined with brown scales. Black longitudinal markings lined anteriorly with white, separated by white patch in middle of cell. Markings gradually becoming lighter towards fold. Inner margin light brown up to fold, sometimes suffused with pink, with distinct dark basal streak. GENITALIA ¢ (Pl. 16, fig.91). Uncus short, tubular, posterior margin set with setae. Anterior margin of tegumen with deep angular emargination; sclerotized part of tegumen reduced to narrow frame. Costa slightly shorter than valva, distal third hardly wider than base, not clavate, densely set with short setae. Harpe approximately two-thirds length of costa, bent at right angle near middle. Basal half of valva broad, distal half narrow, curved, apex with short spine. Sacculus triangular, widest at base. Aedeagus with swollen base, tapering, apical third strongly curved. Manica partially sclerotized. GENITALIA 9. Unknown. The only available 9 has lost its abdomen. REMARKS. O. frontella differs from aspera and pulchella by the strongly developed arc above the transfrontal sulcus of the head, the light anterior half of the fore wing and the presence of a distinct basal streak. In the ¢ genitalia frontella differs by the bent harpe, the triangular sacculus which is widest at the base, the aedeagus with strongly curved apical portion and the sclerotized manica. BroLocy. Host-plant unknown. Moths have been collected by Kaszab in June and July in localities where Tamarix occurs (Kaszab, 1966 : 591, 597; 1968 : 22). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Monco ta: South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (genitalia slide no. 637c, Sattler; TM, Budapest). Paratypes. MONGOLIA: I 9, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (TM, Budapest) [abdomen missing]; 2 §, Gobi Altaj aimak, Zachuj Gobi, 1o km N. of Chatan chajrchan mountains, 1150 m, 27.vi.1966 (Kaszab, no. 594) (TM, Budapest; BMNH); 2 g, Chovd aimak, 10 km SSW. of Somon Bulgan, 1200 m, 5.vii.1966 (Kaszab, no. 633) (TM, Budapest; BMNH). Ornativalva arabica Sattler, 1967 Ornativalva avabica Sattler, 1967, Beitr. naturk. Forsch. SiidwDitl. 26(3) : 48, pl. 4, fig. 25 [not 24!], pl. 10, fig. 52. Holotype 3, SaupiI AraBiA: Jeddah, 29.xii.1957 (Diehl) (genitalia slide no. 350a, Sattler; LN, Karlsruhe) [examined]. Head without enlarged scale bases or frontal process. REVISION OF GENUS ORNATIVALVA II5 GENITALIA g. Sattler, 1967, pl. 4, fig. 25 [not 24!]. GENITALIA 2. Sattler, 1967, pl. 10, fig. 52. BroLtocy. Host-plant unknown. Moths have been taken by various collectors in February—April, August, November and December. DISTRIBUTION. Sudan; Saudi Arabia. FURTHER MATERIAL EXAMINED. SAuDI ARABIA: I 3, Mecca, 15.11.1934 (Philby) (BMNH). Ornativalva sesostrella (Rebel, 1912) Gelechia sesostrella Rebel, 1912, Dt. ent. Z. Iris 26 : 88. Holotype 9, Ecypr: Helwan, larva on Tamarix, moth emerged 15.—29.iv.1910 (Debski) (genitalia slide no. 3061; NM, Vienna) {examined}. Gelechia sesostvella Rebel; Meyrick, 1925a : 79. Gelechia sesostrella Rebel; Gaede, 1937 : 212. Ornativalva sesostrella (Rebel) Sattler, 1964 : 578. Ornativalva sesostrella (Rebel); Sattler, 1967 : 50, pl. 4, fig. 24 [mot 25!], pl. 11, fig. 53. Head without enlarged scale bases or frontal process. GENITALIA g. Sattler, 1967, pl. 4, fig. 24 [not 25!]. GENITALIA 9. Sattler, 1967, pl. 11, fig. 53. BioLtocy. Host-plants: Tamarix species (moths bred by Debski and Kasy). Larvae have been collected by Kasy in January and February on Tamarix (2) nilotica (Ehrenberg) Bunge; the moths emerged in February. Moths have been taken by various collectors in January—August, October and November. It appears that sesostrella breeds continuously where conditions permit this. The larva was described by Debski on a specimen label as ‘larva libera fusiformis’; no detailed description is available. DIsTRIBUTION. Algeria; Tunisia; Egypt; Sudan; Saudi Arabia; S. Iran (Kerman); _ Pakistan. FURTHER MATERIAL EXAMINED. IRAN: I dg, S. Iran, [Kerman,] 80 km SE. of Sirjan, 9.1v.1970 (Kasy) (NM, Vienna) [first record for Iran]. PAkisTAN: I 9, 80 km NW. of Quetta, 2100 m, I5.v.1965 (Kasy & Vartian) (NM, Vienna) [first record for Pakistan]. Ornativalva ignota Sattler, 1967 (Pil 23, fig. 112) Ornativalva ignota Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26(3) : 52, pl. 4, fig. 26. Holo- type g, ALGERIA: Hassi el Abiod, 29.ix.1918 (Dumont) (genitalia slide no. 376c, Sattler; MNHN, Paris) [examined]. [Gelechia evubescens Walsingham, 1904 : 265 (partim). Misidentification. ] Head without enlarged scale bases or frontal process. GENITALIA g. Sattler, 1967, pl. 4, fig. 26. 116 Ky SATE E R GENITALIA @ (Pl. 23, fig. 112). Apophysis anterior half length of apophysis posterior. Sclerotized antrum slightly shorter than apophysis anterior. Signum pair of irregular basal plates with one straight slender spine each. REMARKS. The @ genitalia of 7gnota are similar to those of sesostrella but differ by the much longer antrum. Although the holotype and paratype are labelled ‘Tunisie’, their localities (Hassi el Abiod and Laghouat) are situated in Algeria. BioLtocy. Host-plant: Tamarix species (moth bred by Dumont). No informa- tion is available on the larva or larval habits. Moths have been bred or collected by Dumont and Walsingham in March, July and September. DISTRIBUTION. Algeria. FURTHER MATERIAL EXAMINED. ALGERIA: I 9, Biskra, 13.111.1903 (Walsingham, no. 96 596) (genitalia slide no. 6267; BMNH) [paralectotype of Gelechia erubescens Walsingham]. THE ORNATELLA-GrRovupP (Text-fig. 5) Head without enlarged scale bases or frontal process. Metascutum with paired group of narrow hair-like scales (Text-fig. 2). Fore wing with basal streak on inner margin. Frenulum of 9 triple. In g genitalia costa, harpe, valva and sacculus present. Conspicuous lobe arising in angle between valva and sacculus. In Q genitalia posterior margin of eighth tergite with irregular row of long setae; eighth segment laterally set densely with rows of microtrichia. Antrum modified, with longitudinal folds. Signum with pair of strong spines; basal plates connected by wide sclerotized bridge. BrioLtocy. Host-plant unknown. Moths have been collected in localities where Tamarix was observed or in association with other Lepidoptera species known to feed on Tamarix. Ornativalva ornatella Sattler, 1967 (Pl. 16, fig. 92; Pl. 23, fig. 113; Text-figs 5, 13-17, 24) Ornativalva ornatella Sattler, 1967, Beity. naturk. Forsch. SidwDtl. 26(3) : 78, pl. 2, fig. 12, pl. 7, fig. 39, pl. 14, fig. 65, pl. 16, fig. 70. Holotype g, AFGHANISTAN: Herat, 970 m, 5.v.1956 (Amsel) (genitalia slide no. 292a, Sattler; LN, Karlsruhe) [examined]. GENITALIA ¢ (Pl. 16, fig. 92; Text-figs 13, 14). GENITALIA Q (Pl. 23, fig. 113; Text-figs 15-17, 24). Seventh segment modified, tergite and sternite medially emarginate; emargination of sternite much wider than that of tergite (Text- fig. 24). REMARKS. Ina from Rumania the anterior two-thirds of the fore wing up to the fold are almost entirely of a uniform grey, whereas the area between the fold and the inner margin is ochreous. In the 2 paratype from Iran the anterior half of the fore wing is not as dark grey as in the Rumanian specimen and the ochreous REVISION OF GENUS ORNATIVALVA 117 area along the inner margin extends to the apex. The longitudinal streak which extends from the base along the posterior margin of the cell is distinct in the specimens from Iran, Afghanistan and Mongolia but is absent in the Rumanian specimen. The wings of a 2 from Sarepta are too denuded of scales for an external comparison. The Sarepta specimen was originally identified by Christoph as plutelliformis. Walsingham (1908 : 939) recognized it as ‘an allied species distinct from both [plutelliformis and sieverst]’. In the § genitalia of specimens from Mongolia the lobe that arises in the angle between valva and sacculus is straight, not curved, and is distally much wider than at the base (Text-fig. 13). In a g from Rumania the harpe is much shorter than in specimens from Afghanistan and Mongolia and the lobe between valva and sacculus is distinctly larger than the free distal end of the sacculus (Text-fig. 14). In the 9 genitalia of specimens from Mongolia the distal end of the apophysis anterior is wider than in specimens from Iran and Afghanistan (Text-figs 15, 16). Ina ®Q from Sarepta the apophysis anterior is distally not dilated (Text-fig. 17). BrioLocy. Host-plant unknown. Most of the Mongolian specimens have been collected at light in localities where Tamarix occurs (Kaszab, 1968 : 23, 26, 27). Moths have been taken by various collectors in May—July, the Rumanian specimen in August. DIsTRIBUTION. Rumania; U.S.S.R. (S. Russia); Turkey; Iran; Afghanistan; Mongolia. FURTHER MATERIAL EXAMINED. RuMAnIA: I 4, [Dobrogea,] Sfintu Gheorghe, 15.viii.1971 (Popescu-Gor7) (MINGA, Bucharest) [first record for Rumania]. TurRKEy: 1 9, Taurus, 20km S. of Mut, 8.vii.1968 (Avenberger) (coll. Arenberger, Vienna) [first record for Turkey]. Mon- GOLIA: I g, I 9, Bajanchongor aimak, spring Talyn Bilgech bulag, 47 km E. of borderpost Caganbulag, 1200 m, 23.vi.1967 (Kaszab, no. 840) (ITM, Budapest); 2 9, Bajanchongor aimak, Cagan Bogd ul, Tooroin bulag, 13 km E. of borderpost Caganbulag, 1500 m, 25.vi.1967 (Kaszab, no. 849) (TM, Budapest); 2 3, 5 2, Bajan- chongor aimak, oasis Echin gol, 90km NE. of borderpost Caganbulag, 950 m, 27.-29.v1.1967 (Kaszab, nos 857, 858, 859) (IM, Budapest; BMNH); 8 4, 3 9, Bajanchongor aimak, oasis Dzun mod, 100 km S. of Somon Schine zinst, 1300 m, 29.v1.1967 (Kaszab, no. 869) (IM, Budapest; BMNH) [first record for Mongolia]. THe PLICELLA-Group (Text-fig. 6) Head without enlarged scale bases or frontal process. Metascutum with paired group of narrow hair-like scales. Fore wing without basal streak on inner margin. Frenulum of 2 triple. In g genitalia uncus reduced; costa, valva and harpe present. Sacculus separated from valva. Anellus lobes sclerotized. In @ genitalia posterior margin of eighth tergite with irregular row of long setae. Apophysis anterior rod-like, straight, or broad with bent distal 118 Ke, SA Peek third. Sclerotized antrum well developed or absent. Cervix bursae with one or two sclerotized transverse folds. Pair of rounded signa densely set with spines. BroLtocy. Host-plant unknown. Ornativalva plicella sp. n. (Pl. 2, fig. 21-°Pl. 17, fig. 93: Pl 24) fig: 124} Text=ien6) 3, 2. 6:0-7-0mm. Head ochreous. Labial palpus pale ochreous, outer surface of second segment with scattered brown scales, particularly at base; third segment strongly mixed with brown scales. Antenna dark brown with indistinct paler rings above, whitish below. Thorax ochreous, laterally brown, with weak longitudinal brown line. Tegula brown, sometimes with light apex. Fore wing dark, grey-brown, between fold and inner margin ochreous, division line between dark and light area W-shaped at fold. Inner margin dark brown at base but no distinct basal streak separated. Indistinct black dot at end of cell. Indistinct light mark on costa at two-thirds, no distinct outer transverse line. Apex of wing ochreous, indistinct line of short dark streaks at base of fringes. GENITALIA ¢ (Pl. 17, fig. 93). Costa shorter than valva, slender, apical portion clavate. Harpe short, broad. Valva strongly curved at two-thirds, distal third narrow, apex rounded, without terminal spine. Sacculus two-thirds length of valva, narrow. Anellus lobes large, rounded. Aedeagus with big bulbous base and narrow apical half; apex rounded. GENITALIA @ (Pl. 24, fig. 114). Apophysis posterior as long as eighth segment (including apophysis anterior). Distal third of apophysis anterior conspicuously bent. No sclerotized antrum. Ductus bursae shorter than abdomen. Cervix bursae bears two sclerotized folds with serrated edges; folds connected by narrow sclerotization. Corpus bursae globular, with pair of irregularly rounded signa with strong spines. REMARKS. O. flicella differs from uwndella by the larger size, the darker colour of the fore wing and the @ genitalia with the strongly bent apophysis anterior, the two sclerotized folds of the cervix bursae and the absence of a sclerotized antrum. Biorocy. Host-plant unknown. Moths have been collected by Kasy in April. DISTRIBUTION. S. Iran (Luristan). MATERIAL EXAMINED. Holotype 3, Iran: S. Iran, [Luristan,] 100 km N. of Bandar-Abbas, 5.iv.1970 (Exped. Mus. Vind.) (genitalia slide no. 3877; NM, Vienna). Paratypes. IRAN: 2 4, 2 9, S. Iran, [Luristan,] 100 km N. of Bandar-Abbas, 5.1v.1970 (Exped. Mus. Vind.) (NM, Vienna; BMNH). Ornativalva undella sp. n. (PL..1, tig. 255. 2.ciee 22 el 2a hee ras) ®. 5:0mm. Fore wing dark brown between costa and fold, ochreous between fold and inner margin, division line between dark and light area W-shaped at fold. Light area crosses fold near middle and extends to end of cell. Wing ochreous at apex and along outer margin. Light dot on costa at two-thirds; no outer transverse line. Fringes ochreous, basal half mixed with brown. GENITALIA g. Unknown. REVISION OF GENUS ORNATIVALVA 119 GeEniTALia Q (Pl. 24, fig. 115). Apophysis posterior nearly three times length of apophysis anterior. Apophysis anterior straight. Antrum well developed, about twice length of apo- physis anterior. Cervix bursae with sclerotized fold. Corpus bursae globular, with pair of irregularly rounded signa with strong spines. REMARKS. The holotype lacks the head which had to be reconstructed in the colour figure. The head structures are probably similar to those of flicella. O. undella differs from plicella by the smaller size, the lighter fore wing colour, the long antrum, the straight apophysis anterior and the presence of only one sclerotized fold in the cervix bursae. BroLocy. Host-plant unknown. The only specimen has been collected by Kasy in May. DIsTRIBUTION. SE. Iran (Baluchestan). MATERIAL EXAMINED. Holotype 2, IRAN: SE. Iran, [Baluchestan,] 100 km W. of Zahedan (‘Sahidan’), 1150 m, I1.v.1965 (Kasy & Vartian) (genitalia slide no. 3864; NM, Vienna). THe TAMA RICIELLA-GRoupP (Text-figs 7-9) Head without enlarged scale bases or frontal process. Metascutum with paired group of - narrow, hair-like scales (Text-fig. 2). Fore wing in some species with groups of raised scales, without (rarely with) basal streak on inner margin. Frenulum of 9 triple. In ¢ genitalia uncus reduced; costa and valva present; harpe short, rounded or specialized (pharaonis). Sacculus partially separated from valva. Anellus lobes weakly sclerotized. In Q genitalia posterior margin of eighth tergite with irregular row of long setae. Apophysis anterior short, rod-like. Sclerotized antrum absent, posterior part of ductus bursae sometimes weakly sclerotized (phavaonis, misma) or with sclerotized ring at starting point of ductus seminalis (macrosignella). Signum of varied shape. BrioLtocy. Several species have been bred from Tamarix. Ornativalva species 4 (Pl. 25, fig. 116) 9. 5:0-5°5mm. Specimens poorly preserved. GENITALIA g. Unknown. GENITALIA @ (Pl. 25, fig. 116). Apophysis posterior as long as eighth segment (including apophysis anterior). Apophysis anterior short, wide at base, not rod-like, distally bent. No sclerotized antrum. Ductus bursae short and wide, gradually widening into corpus bursae. Sclerotized plate of signum elongate, medially constricted, with pair of transverse ridges. REMARKS. The signum appears somewhat similar to the signa of Jalyella and arabica; however, the structure of the eighth abdominal segment, the short apophysis anterior and the absence of a sclerotized antrum place this species in the tamariciella- group. 120 K. SATTLER BioLtocy. Host-plant unknown. Moths have been collected by Kaszab in June and July in localities where Tamarix occurs (Kaszab, 1966 : 597-598; 1968 : 26-27). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Monco la: I 9, Chovd aimak, 10 km SSW. of Somon Bulgan, 1200 m, 5.vii.1966 Kaszab, no. 633) (TM, Budapest); 1 9, Bajanchongor aimak, oasis Echin gol, 90 km NE. of borderpost Caganbulag, 950 m, 27.-29.vi.1967 (Kaszab, no. 857) (BMNH). Ornativalva indica Sattler, 1967 (Text-fig. 8) Ornativalva indica Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26(3) : 54, pl. 5, fig. 27, pl. 11 fig. 54. Holotype g, Inp1a: Bihar, Pusa, 29.i11.1930 (Fletcher) (genitalia slide no. 6509; BMNH) [examined]. [Gelechia tamaricella Zeller; Maxwell-Lefroy, 1909 : 534. Incorrect subsequent spelling of tamariciella Zeller. Misidentification. ] [Gelechia tamariciella Zeller; Fletcher, 1920 : 82 (partim). Misidentification. ] [Gelechia tamariciella Zeller; Meyrick, 1925a : 78 (partim). Misidentification. ] [Gelechia tamariciella Zeller; Fletcher, 1932 : 41. Misidentification. } GENITALIA g. Sattler, 1967, pl. 5, fig. 27. GENITALIA 9. Sattler, 1967, pl. 11, fig. 54. BioLtocy. Host-plant: Tamarix species (moths bred by Fletcher). The host- plant is possibly Tamarix indica Willdenow. Fletcher (1920: 82; 1932: 42) erroneously identified it as Tamarix gallica Linnaeus, which does not occur in Pakistan and India. According to the literature (Maxwell-Lefroy, 1909 : 82, Fletcher, 1920 : 82; 1932 : 41) the larva ties three or four twigs together into a case which it carries about. It feeds on dry twigs of Tamarix. Larvae have been observed in February, March, May and December. Pupation takes place in a cocoon on the ground. The pupal period lasts from 8 to 15 days. For a description of the larva and pupa see Fletcher (1920 : 82; 1932 : 42). Moths have been bred or collected in May—July. DISTRIBUTION. S. Iran (Luristan); Pakistan (Peshawar); India (Bihar). FURTHER MATERIAL EXAMINED. IRAN: I g, 4, S. Iran, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH) [first record for Iran]. Ornativalva pharaonis Sattler, 1967 Ornativalva phavraonis Sattler, 1967, Beitr. naturk. Forsch. SiidwDitl. 26(3) : 58, pl. 2, fig. 10, pl. 5, fig. 28, pl. 17, fig. 73. Holotype g, SupaN: Blue Nile Province, Wad Medani, 2.viii.1962 (Remane) (genitalia slide no. 19 859; BMNH) [examined]. GENITALIA g. Sattler, 1967, pl. 5, fig. 28. GENITALIA 9. Sattler, 1967, pl. 17, fig. 73. REVISION OF GENUS ORNATIVALVA 120 Biotocy. Host-plant: Tamarix species (moths bred by Wiltshire and Trought); Tamarix aphylla (Linnaeus) Karsten (moths bred by Limon from larvae inhabiting the galls of Eviophyes tlaiae Trabut). Moths have been bred or taken by various collectors in March and June—November. DISTRIBUTION. Tunisia; Libya; Egypt; Sudan; Israel. FURTHER MATERIAL EXAMINED. LipyA: 1 9, Tripoli, larva on Tamarix, 22.xii.1960, moth emerged iii. 1961 (Trought) (BMNH) [first record for Libya]. IsRAEL: 1 3, 1 9, Ashkelon, larvae in galls of Eriophyes tlaiae Trabut: on Tamarix aphylia (Linnaeus) Karsten, moths emerged 26., 27.x.1972 (Limon) (TAU, Tel-Aviv) [first record for Israel]. Ornativalva misma sp. n. (Plt; eb; Pl. 3; fig. 237 Pl. 17, fig. 94; Pl. 25, fig. x27) 3,2. 4:0mm. Head pale ochreous, darker on vertex. Labial palpus whitish, outer surface of basal segment brown; outer surface of second segment with brown transverse bands at base and before apex; third segment with broad dark brown ring around middle and with dark apex. Antenna whitish below, above brown with paler rings. Thorax light brown, laterally dark brown. Tegula dark brown, apex light brown or whitish. Anterior two-thirds of fore wing pale ochreous with brown markings. Oblique brown band near base between costa and fold. Broader, more distinct band from basal third of costa to outer third of fold, darkest in fold; black dot at outer angle of band. Indistinct brown patches on costa at two-thirds and directly before apex. Black dot at end of cell. Posterior third of fore wing between fold and inner margin ochreous or light brown, without basal streak. Fringes light brown with dark brown base; outer dark band distinct below apex, fading towards tornus. GENITALIA ¢ (Pl. 17, fig. 94). Costa clavate, approximately as long as sacculus, distinctly shorter than valva. Harpe short, rounded. Valva tapering, apex with short spine. Sacculus broad, fused with valva for most of its length. Saccus large. Aedeagus with bulbous base; apical two-thirds gently curved, tapering, apex pointed. GenITaLiA 9 (Pl. 25, fig. 117). Apophysis anterior as long as eighth sternite. No sclerotized antrum. Posterior third of ductus bursae almost straight, weakly sclerotized; anterior two- thirds membranous, coiled. Corpus bursae with narrow band of minute spines across middle. Signum an elongate plate with pair of strong teeth. REMARKS. O. misma is externally similar to heluanensis but the fore wing lacks the basal field which is often clearly separated in heluanensis. The fore wing fringes are fairly uniform in colour whereas in heluanensis they are usually disrupted by two or three light narrow longitudinal zones. The ¢ genitalia of misma are similar to those of indica but the apex of the costa is not conspicuously enlarged and the aedeagus has a longer and more pointed apex. Externally indica differs from misma by the presence of some groups of raised scales on the fore wing. The aedeagus of misma resembles that of heligmatodes; however, heligmatodes has a much narrower sacculus. In the 9 genitalia the almost straight posterior portion of the ductus bursae resembles that of phavaonis but the signum and the longer apophysis anterior are distinct. O. heligmatodes with a similar signum has shorter apophyses anteriores and stronger signum teeth. In the only available 2 of misma one signum tooth is much shorter than the other; however, this could be accidental. ! 122 K. SATTLER Brotocy. Host-plant unknown. The type-specimens have been collected by Remane in June in association with other Lepidoptera species known to feed on Tamarix. DISTRIBUTION. Sudan MATERIAL EXAMINED. Holotype g, SUDAN: NE. Sudan, Port Sudan, 23.vi.1962 (Remane) (genitalia slide no. 18 841; BMNH). Paratype. SUDAN: 19, NE. Sudan, Port Sudan, 23.vi.1962 (Remane) (BMNH). Ornativalva serratisignella Sattler, 1967 (PL. 35 higs24e Play yptigng5) Ornativalva servatisignella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 60, pl. 12, fig. 59. Holotype 9, Irag: Baghdad, Abu-Ghraib, 30.ix.1958 (Remane) (genitalia slide no. 19 858; BMNH) [examined]. Ornativalva servatisignella Sattler, 1964 : 578. Nomen nudum. 3,2. 5:0-7-0mm. Head grey. Labial palpus grey, outer surface of second segment with some brown scales near base and before apex; third segment with dark ring below middle, apex dark. Antenna brown with paler rings above, whitish below. Thorax and tegula grey. Fore wing grey, between fold and inner margin lighter than between costa and fold; with indistinct brown markings. W-shaped marking at fold, dark dot at end of cell. Dark shadow between end of cell and outer transverse line extending to tornus. Outer transverse line indistinct, strongly angulate. Incomplete dark line along termen. Fringes grey, on termen and tornus suffused with pink. GENITALIA ¢ (Pl. 17, fig. 95). Costa as long as valva, apex densely set with strong setae. Harpe short, rounded. Valva with short apical spine. Sacculus well separated from valva, at one-third approximately three times as wide as before apex. Aedeagus with big bulbous base, apical portion straight. GENITALIA 9. Sattler, 1967, pl. 12, fig. 59. Remarks. As the type-material was in poor condition the above description of external characters is made from fresh specimens. Some specimens show an indica- tion of raised scales on the fore wing, where the fold is crossed by the dark W- markings and in the tornus. The specimen from Iranshahr, listed below, was available to me at the time of the original description; however, it was accidentially omitted from the type-series. BroLocy. Host-plant unknown. Moths have been taken by various collectors in February, April, May and September in association with other Lepidoptera species known to feed on Tamarix. DISTRIBUTION. Sudan; Iraq; Iran (Luristan, Baluchestan). FURTHER MATERIAL EXAMINED. IRAN: I 4, 3 Y, S. Iran, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 15.iv.1972, 26.iv.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 6 9, [Luristan,] 22 km E. of Bandar-Abbas, 1g. iv., 10.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 3 4, REVISION OF GENUS ORNATIVALVA 123 11 9, [Luristan,] 30 km E. of Bandar-Abbas, 8.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 2 dg, [Luristan,] 13 km NE. of Bandar-Abbas, 24.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); I J, 2 9, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna); 1 9, [Baluchestan,] Iranshahr, 800 m, 22.—30.iv.1954 (Richter & Schduffele) (SMN, Stuttgart) [first record for Iran]. Ornativalva caecigena (Meyrick, 1918) (PEs fig: 7; PL 28, tig."96) Gelechia caecigena Meyrick, 1918, Exot. Microlepidopt. 2:134. Holotype 9, PaAkIsTAN: Peshawar, [Tarnab,] v. 1916 (Fletcher) (genitalia slide no. 8251, Clarke; BMNH) [examined]. Telphusa caecigena (Meyrick) Meyrick, 1925a : 70. Telphusa caecigena (Meyrick); Gaede, 1937 : 122. Ornativalva caecigena (Meyrick) Sattler, 1967 : 67, pl. 15, figs 69, 69a. Telphusa caecigena (Meyrick); Clarke, 1969 : 431, pl. 215, figs 2—2c. GENITALIA ¢ (Pl. 18, fig. 96). Costa almost as long as valva, basal two-thirds with nearly parallel margins, terminal third gently tapering, apex rounded. Harpeshort. Valva tapering, with short terminal spine. Sacculus narrow, distally dilated, near apex approximately twice as wide as near base. Aedeagus with bulbous base, angulate at middle, apex blunt. GENITALIA 9. Sattler, 1967, pl. 15, figs 69, 69a. Remarks. The ¢ genitalia are very similar to those of other species of the tamariciella-group but differ by the sacculus which is much wider near the apex than at the base. They are also distinguished by the blunt apex of the aedeagus. BioLocy. Host-plant unknown. Moths have been taken by various collectors in March—May. DISTRIBUTION. Saudi Arabia; Kuwait; S. Iran (Luristan); Pakistan. FURTHER MATERIAL EXAMINED. SAUDI ARABIA: I 9, Riad, 16.11.1960 (Diehl) (LN, Karlsruhe) [first record for Saudi Arabia]. Kuwait: 1 @, 10.11.1944 (Wiltshire) (BMNH) [first record for Kuwait]. IRAN: 2 g, 1 9, S. Iran, [Luristan,] 8km E. of Bandar-Abbas, dunes, 8., Il.iv.1972 (Exped. Mus. Vind.) (NM, Vienna); 1 g, 1 9, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 2., 15.iv.1972 (Exped. Mus. Vind.) (NM, Vienna); 1 9, [Luristan,] 22 km E. of Bandar-Abbas, 10.v.1974 (Exped. Mus. Vind.) (NM, Vienna); I 9, [Baluchestan,] Iranshahr, 800 m, 1.—10.i11.1954 (Richter & Schduffele) (SMN, Stuttgart) [first record for Iran]. PAKISTAN: 2 9, 150 km SW. of Quetta, goo m, 13.v.1965 (Kasy & Vartian) (NM, Vienna; BMNH). Ornativalva macrosignella Sattler, 1967 Ornativalva macrosignella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 60, pl. 5, fig. 29, pl. 11, fig. 55. Holotype 9, Tunisia: Tozeur, vi. 1910 (Lucas) (genitalia slide no. 446b, Sattler; MNHN, Paris) [examined]. [Teleia tamariciella (Zeller); Rebel, 1912 : 90 (partim). Misidentification.] [Teleia tamariciella (Zeller); Debski, 1913 : 110 (partim). Mlisidentification.] [Teteia heluanensis Debski; Debski, 1913 : 111 (partim). Misidentification.] 124 KK, SATIERR [Teleia tamaricalis (Zeller); Amsel, 1933 :125 (partim). Incorrect subsequent spelling of tamariciella Zeller. Misidentification. ] [Teleia tamariciella (Zeller); Amsel, 19356 : 263 (partim). Misidentification.] GENITALIA g. Sattler, 1967, pl. 5, fig. 29. GENITALIA 9. Sattler, 1967, pl. 11, fig. 55. BioLocy. Host-plants: Tamarvix tetragyna Ehrenberg and T. nilotica (Ehrenberg) Bunge (moths bred by Debski). Larvae have been observed in April and December. Moths have been bred or taken by various collectors in February—July. As Debski (1913 : 111) was dealing with mixed material his description of the larva of ‘heluanensis’ could apply to macrosignella (see also heluanensis, p. 102). DisTRIBUTION. Algeria; Tunisia; Egypt; Sudan; Israel; S. Iran (Luristan). FURTHER MATERIAL EXAMINED. SUDAN: 2 g, I 9, Ed Damer, Hudeiba, 1.-8.v.1962 (Remane) (ZSBS, Munich). IRAN: 2 g, S. Iran, [Luristan,] 30 km E. of Bandar-Abbas, 8.v.1974 (Exped. Mus. Vind.) (NM, Vienna) [first record for Iran]. Ornativalva heligmatodes (Walsingham, 1904) Gelechia heligmatodes Walsingham, 1904, Entomologist’s mon. Mag. 40 : 267. LECTOTYPE J, ALGERIA: E] Kantara, 25.v.1903 (Walsingham) (BMNH), here designated [examined]. Teleia heligmatodes (Walsingham); Caradja, 1920 : 104. Gelechia heligmatodes Walsingham; Meyrick, 1925a : 78. Gelechia heligmatodes Walsingham; Gaede, 1937 : 177. Ornativalva heligmatodes (Walsingham) Sattler, 1960 : 59. Ornativalva heligmatodes (Walsingham) ; Sattler, 1967 : 62, pl. 5, fig. 30, pl. 12, fig. 56. GENITALIA g. Sattler, 1967, pl. 5, fig. 30. GENITALIA 9. Sattler, 1967, pl. 12, fig. 56. REMARKS. Walsingham described heligmatodes from four specimens, including ‘type, d (96 465); 2 (89 225)’. The specimen bearing Walsingham’s number 96 465 and referred to as the ‘type 3” on the specimen label and in the original description is here designated as the lectotype. Caradja (1920 : 104) erroneously considered heligmatodes to be a desert form of tamariciella. Rebel (1907 : 93; 1931 : 123), followed by Meyrick (1925a : 78), recorded heligmatodes from Socotra; however, his material proved to be antipyramis. BioLtocy. Host-plant unknown. Moths have been taken by various collectors in March—June and August. DISTRIBUTION. Algeria; Tunisia. Ornativalva species 5 (Pl. 18, fig? 97; Pi: 25, hig. 118) 3, 9. 4:5-5-omm. Head pale ochreous. Labial palpus whitish, outer surface of second segment with brown scales, particularly at base and before apex; third segment with irregular brown ring around middle and brown apex. Thorax and tegula pale ochreous with scattered REVISION OF GENUS ORNATIVALVA 125 brown scales. Costal half of fore wing pale ochreous to grey. Light patch on costa before apex. Outer transverse line indistinct. W-shaped markings dark at fold. Black dots in middle and at end of cell. Posterior half of wing ochreous between fold and inner margin. Some dark scales on inner margin at base of wing, but no distinct basal streak. GeEniTaLia ¢ (Pl. 18, fig. 97). Costa almost as long as valva, truncate, terminally almost twice as wide as basally. Harpe short, rounded. Apical portion of valva slender, with terminal spine. Sacculus shorter than costa and valva. Aedeagus with bulbous base; apical half straight, with parallel sides, not pointed. GENITALIA Q (PI. 25, fig. 118). Apophysis anterior half length of apophysis posterior. Narrow sclerotized plate below ostium bursae. No sclerotized antrum. Posterior portion of ductus bursae with zone of minute spines near junction of ductus seminalis. Signum a curved basal plate with pair of strong processes. Remarks. The ¢ genitalia resemble those of macrosignella, particularly the aedeagus; they differ by the slender apical portion of the valva. The 9 genitalia are near those of heligmatodes, but differ by the signum. O. heligmatodes is also distinguished externally by the basal streak on the fore wing. BioLtocy. MHost-plant unknown. Moths have been collected by Kasy in April and May in localities where Tamarix occurs. DISTRIBUTION. S. Iran. MATERIAL EXAMINED. IRAN: I g, 5 9, S. Iran, [Luristan,] 22 km N. of Bandar-Abbas, 18., 22.iv.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 1 9, [Baluchestan,] 100 km W. of Zahedan (‘Sahidan’), 11.v.1965 (Kasy & Vartian) (NM, Vienna). Ornativalva tamariciella (Zeller, 1850) (Text-fig. 7) Gelechia tamariciella Zeller, 1850, Stettin. ent. Zig 11: 153. Lectotype 9, Itaty: Livorno, v. 1846 (Mann) (genitalia slide no. 6061; BMNH), designated by Sattler (1967 : 88) [examined]. Gelechia tamariciella Zeller; Herrich-Schaffer, 1854 : 167, pl. 75, fig. 567. Gelechia tamariciella Zeller; Wocke, 1861 : 114. Gelechia tamariciella Zeller; Stainton, 1867 : 20. Gelechia tamariciella Zeller; Stainton, 1869 : 8o. Teleia tamariciella (Zeller); Wocke, 1871 : 295. Telcia tamariciella (Zeller); Mann, 1873 : 128. Incorrect subsequent spelling of Teleza Heine- mann. Teleja tamariciella (Zeller); Curd & Turati, 1882 : 43. Incorrect subsequent spelling of Teleza Heinemann. Teleia tamariciella (Zeller); Constant, 1892 : 67 (partim). Teleia tamariciella (Zeller); Rebel, 1901 : 150 (partim). Gelechia tamariciella Zeller; Meyrick, 1925a : 78 (partim). Gelechia tamariciella Zeller; Gaede, 1937 : 217 (partim). Ornativalva tamariciella (Zeller) Sattler, 1960 : 59. Ornativalva tamariciella (Zeller); Sattler, 1967 : 63, 88, pl. 6, fig. 31, pl. 12, fig. 57. Ornativalva tamariciella (Zeller); Zocchi, 1971 : 60, fig. vii. GENITALIA g. Sattler, 1967, pl. 6, fig. 31. GENITALIA 9. Sattler, 1967, pl. 12, fig. 57. 126 K. SATTLER REMARKS. The name tamariciella has been erroneously attributed to Mann by various authors. Although the name originated from Mann it was made nomen- claturally available by Zeller. Between 1850 and 1967 the name tamariciella was recorded in approximately 40 publications. All of those records were either based directly or indirectly on the specimens which Mann collected in 1846 and 1872 or they were misidentifications. Several authors have erroneously recorded tamariciella from Portugal, Spain, France, Malta, Algeria, Tunisia, Egypt, Israel, Saudi Arabia, Iran and India. Re-examination of most of the material on which those records were based has revealed the following misidentifications: Portugal, Spain, France — pseudotamari- ciella; Malta, Algeria, Iran — heluanensis; Egypt, Israel — heluanensis, macrosignella; India — indica. Records for ‘Syria’ are directly or indirectly attributable to Kalchberg (1898 : 190), who recorded tamariciella from Haifa [Israel]. Kalchberg’s specimens have not been traced. The records for Tunisia and Saudi Arabia could not be checked but probably apply to heluwanensis. Records for S. Italy and Sicily are doubtful and must be checked. The record for Piemonte (Mariani, 1943 : 166) might apply to tamariciella or pseudotamariciella. BioLocy. Host-plants: Tamarix species (moths bred by Amsel); 7. gallica Linnaeus (Zocchi, 1971 : 61). Nothing is known about the larva and its habits. Mann observed the moths in May 1846; they were uncommon on Tamarix trees, flying before and after sunset when mating also took place (Zeller, 1850 : 153). Moths have been bred by Amsel in June 1930. DISTRIBUTION. Italy (Toscana); Yugoslavia (Dalmatia). Ornativalva pseudotamariciella Sattler, 1967 Ornativalva pseudotamariciella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 65, pl. 6, fig. 32, pl. 12, fig. 58. Holotype 9, France: Alpes-Maritimes (? Constant) (genitalia slide no. 364; ZSBS, Munich) [examined]. [Televa tamariciella (Zeller); Milliére, 1876a : 330. Misidentification. ] [Televa tamariciella (Zeller); Stainton, 1881 : 247. Misidentification.] [Teleia tamariciella (Zeller); Constant, 1892 : 67. Misidentification. ] Ornativalva pseudotamariciella Sattler; Agenjo, 1968 : [5]. GENITALIA g. Sattler, 1967, pl. 6, fig. 32. GENITALIA 9. Sattler, 1967, pl. 12, fig. 58. Remarks. All records of tamariciella from Spain and S. France are probably referable to pseudotamariciella. BroLtocy. Host-plants: Tamarix species (moth bred by Constant); 7. gallica Linnaeus (Milliére, 1876a : 330); T. africana Poiret (Constant, 1892 : 67). According to Milliére (1876a : 330) the larva feeds in June and again in September on the leaves and flowers of its host-plant. Moths have been bred or taken by various collectors in May—July and September. REVISION OF GENUS ORNATIVALVA 127 DISTRIBUTION. Portugal (Algarve); Spain (Catalufa, Murcia, Andalucia); France (Loire-Atlantique, Alpes-Maritimes). FURTHER MATERIAL EXAMINED. PorTuGAL: 1 4g, Algarve, 17.v.1880 (Eaton) (BMNH) [first record for Portugal]. SPAIN: I 3, Catalufia, Rosas, salt marshes, 20 m, 14.vi.1964 (Glaser) (coll. Glaser, Vienna); 1 9, Prov. Murcia, Alhama de Murcia, 19.—20.1x.1974 (Glaser) (coll. Glaser, Vienna). Ornativalva kalahariensis (Janse, 1960) (Text-figs 9, 18-20) Pelostola kalahariensis Janse, 1960, Moths S. Afr. 6 : 189, pl. 78, fig. e, pl. 98, fig. b, pl. 99, figs, pl. 116, fig. f, pl. 117, figs f, h, i. Holotype ¢, SoutnH Arrica: SW. Kalahari, Auob, iv. 1933 (van Son) (genitalia slide no. 5800, Janse; TM, Pretoria) [examined]. Stegasta species; Janse, 1949, pl. 31, fig. I. Ornativalva kalahariensis (Janse) Sattler, 1967 : 67, pl. 16, fig. 71, pl. 18, fig. 75. GENITALIA ¢ (Text-figs 18-20). Sattler, 1967, pl. 18, fig. 75. GENITALIA 9. Sattler, 1967, pl. 16, fig. 71. Remarks. The genitalia of the holotype 3 were illustrated by Janse (1960, pl. 78, fig. e) and Sattler (1967, pl. 18, fig. 75, erroneously marked as paratype). Some variation is found in the ¢ genitalia, particularly in the shape of costa and sacculus. In the holotype ¢ the right valva (Text-fig. 19) is wider than the left (Text-fig. 18), although part of the apparent difference may be due to the different position in the slide. BroLtocy. NHost-plant unknown. The specimens collected by the BMNH Southern African Expedition were all taken in localities where Tamarix usneoides E. Meyer ex Bunge occurs (records of members of the expedition). Moths have been collected in January (BMNH Sth. Afr. Exped.), April and November (G. van Son). DISTRIBUTION. South Africa; South West Africa. FURTHER MATERIAL EXAMINED. SoutH WEsT AFRICA: 5 3, 39, Homeb, 10 mls ESE. Gobabeb, at light, 23.-25.i.1972 (BMNH Sith. Afr. Exped., no. 23) (BMNH); 3 3, Swakopmund, at light, 26.-30.1.1972 (BMNH Sth. Afr. Exped., no. 25) (BMNH); 6 g, Kahn River, 5 mls N. Usakos, at light, 30.-31.1.1972 (BMNH Sth. Afr. Exped., no. 29) (BMNH) [first record for South West Africa]. THE PLUTELLIFORMIS-GrovuP (Pl. 5, figs 33-35; Text-fig. 10) Head without enlarged scale bases or frontal process. Metascutum with paired group of narrow, hair-like scales (Text-fig. 2). Fore wing with or without basal streak on inner margin. Frenulum of 9 triple. In ¢ genitalia uncus reduced; costa, valva and harpe present. Sacculus 128 K. SATTLER long, narrow, separated from valva. Anellus lobes weakly sclerotized. In 9 genitalia posterior margin of eighth tergite with irregular row of long setae. Apophysis posterior short. Sclero- tized antrum short. Signum with pair of strong spines or transverse tooth-like ridges, single or divided into two parts. BroLocy. Several species have been bred on Tamarix. Ornativalva triangulella Sattler, 1967 Ornativalva triangulella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 68, pl. 6, fig. 33, pl. 13, figs 60, 60a. Holotype 9, AtcrErRta: Biskra, 6.iv.1907 (Chrétien) (genitalia slide no. 390c, Sattler; MNHN, Paris) [examined]. [Gelechia plutelliformis Staudinger; Amsel, 19555 : 126. Misidentification.] GENITALIA g. Sattler, 1967, pl. 6, fig. 33. GENITALIA Q. Sattler, 1967, pl. 13, figs 60, 60a. REMARKS. The holotype 9 from the Chrétien coll. was labelled ‘plutellif. ? v. auctella’ in Chrétien’s handwriting. The name auctella is a nomen nudum. It originated from Chrétien and was introduced into the literature by Turati (1927 : 338) without becoming nomenclaturally available. Specimens in coll. Turati under the name auctella are antipyramis and plutelliformis. BroLocy. MHost-plant unknown. Moths have been taken by various collectors in March—May and October. DistrisuTion. Algeria; Tunisia; Kuwait; Iraq; SE. Iran (Baluchestan); Afghanistan. FURTHER MATERIAL EXAMINED. Kuwalt: I 9, 30.iv.1943 (Wiltshire) (BMNH) [first record for Kuwait]. IRAN: 14, SE. Iran, [Baluchestan,] 100 km W. of Zahedan (‘Sahidan’), 1150 m, II.v.1965 (Kasy & Vartian) (BMNH) [first record for Iran]. Ornativalva antipyramis (Meyrick, 1925) Gelechia antipyvamis Meyrick, 1925), Bull. Soc. ent. Egypte 9:209. Lectotype g, EGyprT: Ballah, vii. 1916 (Buxton) (genitalia slide no. 6144; BMNH), designated by Sattler (1967 : 87) [examined]. [Telera heligmatodes (Walsingham) Rebel, 1907 : 93. Misidentification.] [Gelechia heligmatodes Walsingham; Meyrick, 1925a : 78 (partim). Misidentification. } [Teleja haligmatodes auctella Turati, 1927 : 338 (partim). Nomennudum. Incorrect subsequent spellings of Teleia Heinemann and heligmatodes Walsingham. Misidentification.] [Teleia heigmatodes (Walsingham); Rebel, 1931 : 123. Misidentification.] Gelechia antipyvamis Meyrick; Gaede, 1937 : 147. [Gelechia lacertella Walsingham; Amsel, 1958 : 80, text-fig. 11. Misidentification.] Ornativalva antipyramis (Meyrick) Sattler, 1964 : 578. Ornativalva antipyramis (Meyrick); Amsel, 1966 : 128. Ornativalva antipyramis (Meyrick) ; Sattler, 1967 : 70, 87, pl. 6, fig. 34, pl. 13, fig. 61. Ornativalva antipyvamis (Meyrick); Amsel, 1968 : 17. REVISION OF GENUS ORNATIVALVA 129 GENITALIA g. Sattler, 1967, pl. 6, fig. 34. GeniTaLia 9. Sattler, 1967, pl. 13, fig. 61. REMARKS. The name auctella (nomen nudum) originated from Chrétien. Turati (1927 : 338) recorded four specimens, Cyrenaica: Giarabub, vii. 1926 (Kriiger), which according to him agreed with a specimen from Biskra, identified as auctella by Chrétien. Two of those specimens, though not the one from Biskra, are now preserved in coll. Turati (IE, Bolzano). A g without hind wings and abdomen is clearly plutelliformis, whereas a 9 without abdomen is antipyramis; an ‘auctella’ specimen in coll. Chrétien is triangulella. BroLtocy. Host-plant: Tamarix species. A @ (locality and collector unknown) in MNHN, Paris, is labelled ‘Tamarix, x.—xi.1908, écl. 13.v.1909’._ Moths have been taken. by various collectors in February—July and October. DISTRIBUTION. Cape Verde Islands (St Vincent); Canary Islands (Gran Canaria); Morocco; Algeria; Tunisia; Libya; Egypt; Sudan; Socotra; Jordan; Saudi Arabia; United Arab Emirates; S. Iran (Luristan); Pakistan. FURTHER MATERIAL EXAMINED. LisyA: I 9, Cyrenaica, vil. 1926 (Kriiger) (IE, Bolzano). JORDAN: 1 g, Azraq ed Druz, pumping station, 22.iv.—9.v.1966 (Fletcher) (BMNH); 5 5, 49, Wadier Ratam, 24.1v.-10.v.1966 (Fletcher) (BMNH) [first record for Jordan}. UNITED ARAB EMIRATES: I 9, Masafi, 22.-24.11.1971 (Gallagher) (BMNH) [first record for United Arab Emirates]. IRAN: 19, S. Iran, [Luristan,] 8 km E. of Bandar-Abbas, 23.1v.1974 (Exped. Mus. Vind.) (NM, Vienna); 7 3, 9 9, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 2.-15.iv.1972, 21.iv.-17.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); Ig, 19, [Luristan,] 22 km E. of Bandar-Abbas, 19.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 4 3, 6 9, [Luristan,] 30 km E. of Bandar-Abbas, 3.iv.1970, 8.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 2 Q, [Luristan,] 22 km N. of Bandar-Abbas, 4., 14.iv.1972 (Exped. Mus. Vind.) (NM, Vienna); 2 3, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna) [first record for Iran]. Ornativalva singula Sattler, 1967 (Pl. 3, fig. 25; Pl. 18, fig. 98; Pl. 26, fig. 119; Text-figs 21, 25) Ornativalva singula Sattler, 1967, Beitr. naturk. Forsch. SiidwDil. 26 (3) : 71, pl. 7, fig. 35. Holotype g, AFGHANISTAN: Herat, 970 m, 5.v.1956 (Amsel) (genitalia slide no. 352a, Sattler; LN, Karlsruhe) [examined]. GENITALIA ¢ (PI. 18, fig. 98; Text-fig. 21). GENITALIA Q (PI. 26, fig. 119; Text-fig. 25). Posterior margin of seventh sternite with pair of projections (Text-fig. 25). Sternopleural region of eighth segment with pair of minutely spined longitudinal folds. Apophyses anteriores short, bases connected by narrow sclerotization along anterior margin of eighth sternite. Sclerotized antrum very short, tubular. Ductus bursae coiled, approximately three times length of apophysis posterior. Corpus bursae globular. Signum with pair of strong spines; basal plates weakly sclerotized, connected by weakly sclerotized bridge. 130 Ke SAMPLER ReMARKS. In the ¢ genitalia of specimens from Mongolia the harpe is more digitate than in the holotype from Afghanistan (Text-fig. 21). The 9 genitalia differ from those of other species by the pair of posterior processes on the seventh sternite (Text-fig. 25) and the pair of spined folds on the eighth sternite. BroLtocy. Host-plant unknown. The Mongolian specimens have been collected at light in localities where Tamarix occurs (Kaszab, 1968 : 22, 23). Moths have been collected by Amsel and Kaszab in May and June. DISTRIBUTION. Afghanistan; Mongolia. FURTHER MATERIAL EXAMINED. MONGOLIA: 10 4, 10 9, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250 m, 22.vi.1967 (Kaszab, no. 834) (IM, Budapest; BMNH); 2 g, 49, Bajanchongor aimak, spring Talyn Bilgech bulag, 47 km E. of borderpost Caganbulag, 1200 m, 23.v1.1967 (Kaszab, no. 840) (TM, Budapest) [first record for Mongolia]. Ornativalva basistriga sp. n. (PI. x, fig. 8; Pl_.3, ixes:26; 27; Pl. x9, fag..99; Pl. 26, te 120) 3, 2. 6:5-8:omm. Head grey. Labial palpus grey, outer surface of second segment mixed with brown scales; basal half and apex of third segment brown. Thorax grey. Fore wing grey, between fold and inner margin ochreous, lighter grey along costa. W-shaped markings darkest along fold. Dark dot at end of cell with short dark streak which extends towards apex. Base of fringes with dark markings, particularly distinct along termen. Distinct dark basal streak on inner margin. GENITALIA ¢ (PI. 19, fig. 99). Costa slender, clavate, as long as sacculus, distinctly shorter than valva. Harpe slender, clavate, reaching dilated apical portion of costa. Valva long, distal half narrow with parallel margins, apex with short spine. Sacculus narrow, distally dilated. Aedeagus with bulbous base, apical two-thirds slender, curved, pointed. GENITALIA @ (Pl. 26, fig. 120). Apophyses anteriores distally somewhat dilated, bases connected by narrow sclerotization along anterior margin of eighth sternite. No sclerotized antrum. Ductus bursae narrow, coiled, approximately twice length of apophysis posterior. Signum with pair of strong teeth; basal plates connected by indistinct, weakly sclerotized bridge. Remarks. An albinistic 9 (Pl. 3, fig. 27) is almost completely white with only a few dark markings along the fold, between the end of the cell and the apex, on the base of the fringes on apex and termen, and on the inner margin near the base. Externally basistriga is similar to fplutelliformis and grisea but differs by the distinct basal streak on the inner margin of the fore wing. O. singula, which also has a basal streak, is much smaller. The ¢ genitalia of basistriga differ from those of all other species of the plutelliformis-group by the distally dilated sacculus. The 2 genitalia differ from those of singula, plutelliformis and sieverst by the strong teeth — not spines — of the signum. The similar signum of grisea is divided, whereas in basistriga the basal plates are connected by a sclerotized bridge. However, as the sclerotization is indistinct and weak, it is possible that in a larger material basistriga specimens with two separate signa willbefound. Equally, grisea specimens with a sclerotized bridge between the basal plates of the signum may be expected. REVISION OF GENUS ORNATIVALVA 131 BroLtocy. Host-plant unknown. Most of the type-specimens have been collected at light in localities where Tamarix occurs (Kaszab, 1966 : 597; 1968 : 18, 22, 31). Moths have been collected by Kaszab in June and July. DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Monco ria: Chovd aimak, 10 km SSW. of Somon Bulgan, 1200 m, 5.vul.1966 (Kaszab, no. 633) (IM, Budapest). Paratypes. MONGOLIA: I g, I 9, Chovd aimak, 10 km SSW. of Somon Bulgan, 1200 m, 4.-6., 5.vii.1966 (Kaszab, nos 632, 633) (IM, Budapest; BMNH); 1 9, South Gobi aimak, SW. edge of salt lake Dund gol (‘old’ Somon Gurban-tes), 1300 m, 18.vi.1967 (Kaszab, no. 817) (TM, Budapest); 1 3, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250m, 22.vi.1967 (Kaszab, no. 834) (IM, Budapest); 1 9, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vul.1967(Kaszab, no. 882) (BMNH). Ornativalva plutelliformis (Staudinger, 1859) (Pl. 5, figs 33-35; Text-fig. 10) Gelechia plutelliformis Staudinger, 1859, Stettin. ent. Zig 20 : 239. Lectotype 9, Spain: Cadiz, Chiclana, bred on Tamarix, 25.vi.1858 (Staudinger) (genitalia slide no. 155b, Sattler; MNHU, Berlin), designated by Sattler (1967 : 88) [examined]. Alucita olbiaella Milliére, 1861, Iconogy. Descr. Chenilles Lépid. inédits 1 : 193, 218, (liv. 4) pl. 1, figs 1-6. LECTOTYPE 9, France: Provence (Milliéve) (genitalia slide no. 382c, Sattler; MNHN, Paris), here designated [examined]. [Synonymized by Stainton, 1867 : 14.] Alucita olbienella Milliére; Wocke, 1861 : 191. Incorrect subsequent spelling of olbiaella Milliére. Hypsolophus siewersiellus Christoph, 1867, Stettin. ent. Zig 28 : 239. Syntypes, U.S.S.R.: S. Russia, Krasnoarmeysk (‘Sarepta’) (Christoph) (depository unknown). [Synonymized by Wocke, 1871 : 290.] Gelechia plutellifoymis Staudinger; Stainton, 1867 : 14. Gelechia plutelliformis Staudinger; Wocke, 1871 : 290. Gelechia plutelliformis Staudinger; Christoph, 1885 : 158. Gelechia sinuatella Walsingham, 1904, Entomologist’s mon. Mag. 40 : 223. LECTOTYPE gd, ALGERIA: Biskra, 9.iii.1895 (Eaton) (BMNH), here designated [examined]. [Synonymized by Sattler, 1967 : 72.] Gelechia plutellifoymis Staudinger; Powell, 1905 : 164. Gelechia plutellifovmis Staudinger; Walsingham, 1908 : 938. Gelechia plutelliformis Staudinger; Chrétien, 1917 : 469. Gelechia sinuatella Walsingham; Caradja, 1920 : 98. Gelechia sinuatella Walsingham; Meyrick, 1925a : 79. Gelechia plutellifoymis Staudinger; Meyrick, 19254 : 79. [Teleja haligmatodes auctella Turati, 1927 : 338 (partim). Nomen nudum. Incorrect sub- sequent spellings of Teleia Heinemann and heligmatodes Walsingham. Misidentification. ] Gelechia plutelliformis Staudinger; Gaede, 1937 : 203. Gelechia sinuatella Walsingham; Gaede, 1937 : 213. Ornativalva plutelliformis (Staudinger) Gozmany, 1955 : 311, text-figs 11, 12. Ornativalva plutelliformis (Staudinger) ; Sattler, 1960 : 59, pl. 17, fig. 72, pl. 30, fig. 134. Ornativalva plutelliformis (Staudinger); Kasy, 1962 : 76. 132 K. SATTLER Ornativalva plutelliformis (Staudinger) ; Sattler, 1967 : 72, 88, pl. 1, figs 3, 4, pl. 7, fig. 36, pl. 13, fig. 62. Ornativalva plutelliformis (Staudinger); Zocchi, 1971 : 61. Head (PI. 5, figs 33-35). GENITALIA g. Sattler, 1967, pl. 7, fig. 36. GENITALIA 9. Sattler, 1967, pl. 13, fig. 62. REMARKS. G. flutelliformis Staudinger was described from 2 2 which Staudinger bred at the end of June 1858, from larvae collected on Tamarix near Chiclana. Both specimens are now preserved in coll. Staudinger (MNHU, Berlin). A. olbiaella Milliére was described from an unspecified number of specimens which Milliére bred on Tamarix gallica Linnaeus or collected as adults in S. France: Provence, Toulon and Hyéres area. The lectotype 2 bears the label ‘Alucita Olbiaella Mill. sp. nov.’ but no further data. H. siewersiellus Christoph was described from an unspecified number of specimens which Christoph bred on Tamarix laxa Willdenow or collected as adults near Sarepta. No type-material has been traced. The synonymy of siewersiellus was discussed in detail by Walsingham (1908 : 938). G. sinuatella Walsingham was described from eleven specimens of both sexes, including ‘type, 3 (8298); 2 (96 594)’. The specimen bearing Walsingham’s number 8298 and referred to as ‘type 3’ in the original description and on the specimen label is here designated as the lectotype. The specimen labelled ‘type 9’ bears Walsingham’s number 96 589, whereas the specimen referred to as ‘type 9 (96 594)’ in the original description is labelled ‘paratype 8/9’. Ten of the eleven original specimens are now preserved in BMNH. A 3 from Italy: Calabria has a short dark basal streak on the inner margin of the fore wing. The absence of the basal streak usually distinguishes plutelliformis from some other similar species. For a discussion of auctella (nomen nudum) see triangulella (p. 128) and antipyramis (p. 129). O. plutelliformis, one of the commonest and most widespread Ornativalva species, is recorded in more than 60 publications. BroLtocy. Host-plants: Tamarix gallica Linnaeus (Milliére, 1861 : 193); T. laxa Willdenow (recorded by Christoph, 1867 : 239, as T. laxa and pallasit. T. pallasw Desvaux is currently considered to be a synonym of T. laxa.); T. canariensis Willdenow or T. africana Poiret (recorded by Walsingham, 1908 : 938, as T. gallica which does not occur in the Canary Islands); T. parviflora de Candolle (Kasy, — 1962 : 76); unspecified species of Tamarix (moths bred by Staudinger, Lucas, Kasy and Gerling). One specimen in MNHN, Paris, bears the label ‘Rhus oxyacanthe’; however, it is unlikely that Rhus is a host-plant of plutelliformis. As far as can be seen from the literature (Staudinger, 1859 : 239; Milliére, 1861 : 193; Christoph, 1867 : 239; 1885: 158; Powell, 1905: 164; Walsingham, 1908 : 938; Chrétien, 1917 : 469; Kasy, 1962: 76; Zocchi, 1971 : 61) the larva lives free on Tamarix species. It has been described as varying in colour from dark green to reddish and brown. Larva and pupa have been illustrated in colour by Milliere REVISION OF GENUS ORNATIVALVA 133 (1861, liv. 4, pl. 1, figs 1-3). Pupation takes place in or on the ground in a light cocoon that is covered with small particles of soil. Hibernation probably takes place in the pupal stage. Milliére concluded from his observations of larvae and adults that plutelliformis is probably single-brooded in southern France; however, Powell observed larvae in the same area in June as well as August and September. There may be one or two clearly defined generations in southern Europe whereas plutelliformis probably breeds continually where the climatic conditions are suitable. Larvae have been observed by the above authors in May and June (Turkmeniya), June, August and September (S. France), October and November (Tunisia), December and January (Canary Islands). Moths have been collected through all months of the year. Dates of capture in North Africa range from February till August. DISTRIBUTION. Spain; S. France; Italy; Yugoslavia; Hungary; Rumania; Turkey; Cyprus; U.S.S.R. (S. Russia); Madeira (Madeira, Porto Santo); Canary Islands (Tenerife, Gran Canaria); Morocco; Algeria; Tunisia; Libya; Egypt (Sinai); Sudan; Israel; Jordan; Lebanon; Syria; Saudi Arabia; Iraq; Iran; Afghanistan; Pakistan. According to the literature also found in U.S.S.R. (Turkmeniya) (Christoph, 1885 : 158). The record for Kuwait (Amsel, 1955): 126) is erroneous and applies to triangulella (see p. 128). No records are available for Portugal, Albania, Greece and Bulgaria, although Plutelliformis must occur there. FURTHER MATERIAL EXAMINED. IrALy: I 3, Calabria, Marcellina (Cosenza), 20.ix.1971 (Pelham-Clinton) (RSM, Edinburgh); 1 9, Calabria, Gizzeria Lido, 1.-15.v.1971 (de Medina Alberich) (BMNH); I 4, [Sicily,] Licata, viii. 1906 (Re) (BMNH). Rumania: 1 9, [Dobrogea,] Vasile- Roaita, 18.vil.1947 (Popescu-Gorj7) (MINGA, Bucharest); 1 3, [Dobrogea,| Sfintu Gheorghe delta, 12.viii.1g71 (Popescu-Gor7) (MINGA, Bucharest); 2 9, [Dobrogea, | Eforie Sud, 9.vii.1947, 24.vili.1948 (Popescu-Gorj]) (MINGA, Bucharest). TURKEY: Ig, 292, 40 km SW. of Elazig, goo m, 18.vi.1969 (Kasy) (NM, Vienna); 2 3, Eceabat, Tamarix, moths emerged 25.viii.1972 (Gerling) (TAU, Tel-Aviv). Cyprus: 1 3,19, Skylloura, 8.ix.1967 (Zahradnik) (NM, Prague) [first record for Cyprus]. LrBya: I g, Cyrenaica, vii. 1926 (Kriiger) (IE, Bolzano). Ecypt: 1 4, [Sinai,] Wadi Feiran (‘Wadi Feran’), 4.iii.1935 (collector unknown) (BMNH). IsrRAEL: 1 J, Tel-Aviv, 24.ix.1971 (Kugler) (TAU, Tel-Aviv); 1 3, Ein el Turaba, larva on Tamarix, moth emerged 28. ix. (collector unknown) (TAU, Tel-Aviv). AFGHANISTAN: 2 3, Ghor- band valley, N. of Kabul, 1900 m, 30.vii.1965 (Kasy & Vartian) (NM, Vienna). Ornativalva grisea Sattler, 1967 Ornativalva grisea Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 73, pl. 7, fig. 37, pl. 14, fig. 63. Holotype g, ArGHanistaNn: N. Afghanistan, Polichomri, 700 m, 5.vi.1956 (Amsel) (genitalia slide no. 294b, Sattler; LN, Karlsruhe) [examined]. GENITALIA g. Sattler, 1967, pl. 7, fig. 37. GENITALIA 9. Sattler, 1967, pl. 14, fig. 63. BioLtocy. MHost-plant unknown. Moths have been collected in May-July. DistTRIBUTION. Afghanistan. 134 K. SATTLER Ornativalva species 6 (BL10,, fig... 100; Pl. 26, Tig. 12x) GENITALIA 4 (Pl. 19, fig. 100). Costa slender, distally somewhat dilated, as long as sacculus. Harpe digitate, slightly shorter than costa. Valva longer than costa, pointed. Sacculus narrow, shorter than valva, distal third not fused with valva, rounded. Aedeagus with bulbous base, tapered, apical third curved. GENITALIA Q (PI. 26, fig. 121). Lateral area of fine wrinkles on eighth segment marked off by curved fold. Apophysis anterior rod-like, distally slightly dilated. Antrum one-half length of apophysis anterior, posterior margin triangularly extended. Signum a pair of strong teeth, basal plates connected by narrow, indistinct, sclerotized bridge. Teeth of signum with dentate margin. Remarks. The moths are externally similar to plutelliformis and grisea. The d§ genitalia agree with those of grisea but the harpe appears to be longer. The Q genitalia are unusual for the plutelliformis-group and resemble more those of the erubescens-group. The males and females were collected in different localities and, although they agree well externally, their association must still be considered as doubtful. BioLocy. Host-plant unknown. The moths have been collected by Kaszab in June in localities where Tamarix occurs (Kaszab, 1966 : 591; 1968 : 22). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. MONGOLIA: 2 g, South Gobi aimak, 100 km W. of borderpost Ovot Chuural, 1250m, 22.vi.1967 (Kaszab, no. 834) (TM, Budapest; BMNH); 1 9, Gobi Altaj aimak, Zachuj Gobi, 10 km N. of Chatan chajrchan mountains, 1150 m, 27.vi.1966 (Kaszab, no. 594) (IM, Budapest). Ornativalva sieversi (Staudinger, 1871) Gelechia sieversi Staudinger, 1871, Berl. ent. Z. 14: 309. Lectotype g, U.S.S.R.: S. Russia, Krasnoarmeysk (‘Sarepta’) (Christoph) (MNHU, Berlin), designated by Sattler (1967 : 88) [examined]. Gelechia sieverst Staudinger; Wocke, 1871 : 290. Gelechia steverst Staudinger; Christoph, 1887 : 120. Gelechia plutelliformis Staudinger ab. szeveysi Staudinger; Rebel, 1901 : 144. Gelechia sieverst Staudinger; Walsingham, 1908 : 938. Gelechia sieversi Staudinger; Caradja, 1920 : 98. Gelechia sieverst Staudinger; Meyrick, 1925a : 79. Gelechia sieversi Staudinger; Gaede, 1937 : 212. Ornativalva sieverst (Staudinger) Sattler, 1960 : 59, pl. 17, fig. 73, pl. 30, fig. 135. Ornativalva sieversi (Staudinger) ; Sattler, 1967 : 75, 88, pl. 2, fig. 11, pl. 7, fig. 38, pl. 14, fig. 64. GENITALIA g. Sattler, 1967, pl. 7, fig. 38. GENITALIA 9. Sattler, 1967, pl. 30, fig. 135. REMARKS. The date of publication is generally cited as 1870; however, according to the original wrapper of ‘Drittes und viertes Vierteljahrsheft’, pages 209-432 were issued in January 1871. REVISION OF GENUS ORNATIVALVA 135 BioLocy. Host-plant: (?) Tamarix. Staudinger stated in the original description that Christoph bred this species on Tamarix ‘...if Iam not mistaken...’. A moth has been collected by Kasy and Vartian in July. Various dates on old specimens are unreliable as they often refer to the month in which a specimen was received from the collector rather than the date of capture. DistTRiBuTION. U.S.S.R. (S. Russia); Iran (Tehran); Afghanistan. According to the literature also found in U.S.S.R. (Turkmeniya) (Christoph, 1887 : 120). FURTHER MATERIAL EXAMINED. AFGHANISTAN: I 3, Ghorband Valley (N. of Kabul), 1900 m, 30.vii.1965 (Kasy & Vartian) (NM, Vienna) [first record for Afghanistan]. THE CEROSTOMATELLA-Grovup (Text-figs 11, 12) Head with enlarged scale bases or distinct frontal process. Metascutum with paired group of short broad scales (Text-fig. 1). Fore wing in some species deviating from typical Ornativalva pattern; inner margin without basal streak. Frenulum of 9 double or triple. In 3 genitalia of most species uncus developed, deeply divided. Costa, harpe and valva present, sacculus not separated from valva. Anellus lobes modified in some species. In 9 genitalia posterior margin of eighth tergite densely set with scales. Apophysis anterior short, wide, not rod-like; bases of apophyses connected by narrow sclerotization along anterior margin of eighth sternite. No modified ostium bursae. Antrum a small sclerotized ring or absent. Signum with pair of trans- verse folds or ridges. REMARKS. O. acutivalva sp. n. is placed in the cerostomatella-group mainly on account of the presence of scales — not setae — on the eighth tergite of the 2 and the absence of a separate sacculus in the 3. It differs from all other species of this group by the absence of modified head structures and by the long hair-like scales on the metascutum. Biotocy. The host-plants of all species in the cevostomatella-group are still unknown; however, some of the species have been collected as adults in localities where Tamarix occurs or in association with other Lepidoptera species known to feed on Tamarix. Ornativalva mixolitha (Meyrick, 1918) fe i, fig. 9; Pl. 4, fig. 28; Pl. 8, figs 51-56; Pl. 27, fig. 122; Text-fig. 12) Phthorimaea mixolitha Meyrick, 1918 : 135. Head (Pl. 8, figs 51-56) with frontal modifications variable, enlarged scale bases more or less regularly distributed over frons and part of vertex. Sometimes weak or strong frontal process developed. GENITALIA g. Clarke, 1969, pl. 153, figs 2a—b; Sattler, 1967, pl. 8, fig. 4o. GeEniITaLiA Q (Pl. 27, fig. 122). 136 K. SA TaEE RR: Remarks. The North African subspecies bipunctella differs from typical mixolitha by the ochreous — not grey —colour of the fore wing. No differences are found in the genitalia. The frenulum of the 9 is double. The 9 genitalia do not differ from those of cerostomatella, angulatella and mongolica. BroLocy. Host-plant unknown. The moths are usually collected in association with other Ornativalva species and it seems likely that the larva feeds on Tamarix. Moths have been taken by various collectors in February—October. DisTRIBUTION. Morocco; Algeria; Tunisia; U.S.S.R. (S. Russia); Turkey; Sudan; Iraq; Iran; Afghanistan; Pakistan; India (Bihar); Mongolia. Ornativalva mixolitha mixolitha (Meyrick, 1918) (Pl. 1, fig. 9; Pl. 4, fig.i28; PL. 8) figs 51-56; PL 27, fie: 122; Texte. 12) Phthovimaea mixolitha Meyrick, 1918, Exot. Microlepidopt. 2 : 135. Holotype g¢, Inp14: Bihar, Pusa, vi. 1911 (Fletcher) (genitalia slide no. 8307, Clarke; BMNH) [examined]. Phthovrimaea mixolitha Meyrick; Meyrick, 1925a : 92. Phthorimaea mixolitha Meyrick; Gaede, 1937 : 273. Ornativalva mixolitha (Meyrick) Amsel, 1966 : 128. Ornativalva mixolitha (Meyrick); Sattler, 1967 : 80. Phthorimaea mixolitha Meyrick; Clarke, 1969 : 307, pl. 153, figs 2—2b. Head (PI. 8, figs 51-56). GENITALIA g. Clarke, 1969, pl. 153, figs 2a—b. GENITALIA Q (PI. 27, fig. 122). Remarks. The specimens from Turkey and a single 9 from Mongolia are a little bigger and more distinctly marked than typical mixolitha. They also differ by their frontal structure. An arc of strongly raised scale bases extends between the antennal pits above the transfrontal sulcus. The strongly developed frontal process is surrounded by three or four irregular rings of enlarged scale bases (PI. 8, figs 54-56). Ina series of specimens from the Sudan the enlarged scale bases are more or less evenly distributed over the frontal area and there is no indication of a frontal process (Pl. 8, figs 51-53). In material from S. Iran specimens with and without a frontal process are found as well as intermediate stages. Specimens from Afghanistan and Pakistan show also some indication of a frontal process. In two specimens from S. Russia: Sarepta the enlarged scale bases are concentrated in the centre of the frons, but no definite process is developed. Brotocy. Host-plant unknown. Moths have been taken by various collectors in February—August and October. DISTRIBUTION. U.S.S.R. (S. Russia); Turkey; Sudan; Iraq; Iran; Afghanistan; Pakistan; India (Bihar); Mongolia. FURTHER MATERIAL EXAMINED. U.S.S.R.: 13,19, S. Russia, Krasnoarmeysk (‘Sarepta’), 5., 6.viii.1870 (Christoph) (BMNH) [first record for the U.S.S.R.]. Turkey: 22 3, 17 9, Taurus, S. of Mut, REVISION OF GENUS ORNATIVALVA 137 23.v.1969 (Avenberger; Kasy) (coll. Arenberger, Vienna; coll. Jackh, Bidingen; NM, Vienna; BMNH); 1 J, 1 9, road Ankara-Kayseri, 1000 m, 26.vii.1965 (Glaser) (coll. Glaser, Vienna); 1 g, K6opriikoy, Kizilirmak, 750m, 20.-22.vi.1969 (Hahn) (coll. Glaser, Vienna) [first record for Turkey]. SupDAN: I g, Wadi Halfa, 20.-25.i.1962 (Nubien Exp. Mus. Vind.) (NM, Vienna); 2 3, 1 9, Faras W, N. of Wadi Halfa, 3.41.1962 (Nubien Exp. Mus. Vind.) (NM, Vienna); 1 3, 1 9, Khor Musa Pascha, S. of Wadi Halfa, 27. i., 10.11.1962 (Nubien Exp. Mus. Vind.) (NM, Vienna). IRAN: r 9, [Luristan,] 8km E. of Bandar-Abbas, 23.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 13 g, 20 9, [Luristan,] 17 km E. of Bandar-Abbas, dunes, 23.iii.—15.1v.1972, 15.-26.iv.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 1 ¢, 2 9, [Luristan,] 22 km E. of Bandar-Abbas, 19.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 3 3, 7 , [Luristan], 30 km E. of Bandar-Abbas, 3.iv.1970, 25.11.1972, 8.v.1974 (Exped. Mus. Vind.) (NM, Vienna; BMNH); 2 9, [Luristan,] 13 km NE. of Bandar-Abbas, 24.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 1 9, [Luristan,] 22 km N. of Bandar-Abbas, 18.iv.1974 (Exped. Mus. Vind.) (NM, Vienna); 2 3, 3 9, [Luristan,] 25 km S. of Minab, 4.v.1974 (Exped. Mus. Vind.) (NM, Vienna); 1 g, Baluchestan, Iranshar, 800 m, II.—21.v.1954 (Richter & Schduffele) (SMN, Stuttgart) [first record for Iran.] AFGHANISTAN: 2 3, E. of Kandahar, 27.vi.1963 (Kasy & Vartian) (NM, Vienna) [first record for Afghanistan]. PAKISTAN: 2 g, 2 9, Karachi, vicinity of airport, 23.11.—9.111.1961 (Vartian) (NM, Vienna) [first record for Pakistan]. MONGOLIA: I 9, Bajanchongor aimak, between Bajangobi and Somon Bajanleg, 26km SE. of Bajanleg, 1450 m, I.vii.1967 (Kaszab, no. 876) (TM, Budapest) [first record for Mongolia]. Ornativalva mixolitha bipunctella Sattler, 1967 Ornativalva mixolitha bipunctella Sattler, 1967, Beitr. naturk. Forsch. SiidwDtl. 26 (3) : 81, pl. 8, fig. 40. Holotype ¢, ALGERIA: Laghouat, 18.—20.viii.1919 (Dumont) (genitalia slide no. 399¢, Sattler; MNHN, Paris) [examined]. Ornativalva mixolitha bipunctella Amsel, 1966 : 128. Nomen nudum. Head as in the nominate subspecies. GENITALIA g. Sattler, 1967, pl. 8, fig. 4o. GeniTaia 2. As in the nominate subspecies. RemaRKS. The frontal structures of bipunctella are variable as in the nominate subspecies; however, no extreme frontal process as in Turkish and Mongolian mixolitha was observed. BroLtocy. Host-plant unknown. Moths have been taken by various collectors in March—September. DISTRIBUTION. Morocco; Algeria; Tunisia. Ornativalva angulatella (Chrétien, 1915) (Pl. 9, figs 57-59) Gelechia angulatella Chrétien, 1915, Annis Soc. ent. Fr. 84 : 318. Lectotype 9, Tunisia: Tozeur, 20.vi.1907 (Chrétien) (MNHN, Paris), designated by Sattler (1967 : 87) [examined]. Gelechia angulatella Chrétien; Meyrick, 1925a : 79. 138 K. SATTLER Gelechia nigvosubvittatella Lucas, 1933, Bull. Soc. ent. Fry. 38: 199. Holotype g, ALGERIA: Guelt-es-Stel, 11.x.1931 (Lucas) (genitalia slide no. 404c, Sattler; MNHN, Paris) [examined]. [Synonymized by Sattler, 1967 : 83.] Gelechia angulatella Chrétien; Gaede, 1937 : 147. Gelechia nigvosubvittatella Lucas; Gaede, 1937 : 193. Ornativalva angulatella (Chrétien) Sattler, 1967 : 81, 87, pl. 8, fig. 41. Head (Pl. 9, figs 57-59) without frontal process. Enlarged scale bases evenly distributed over frons and anterior part of vertex. GENITALIA g. Sattler, 1967, pl. 8, fig. 41. Remarks. The frontal modifications are similar to those of cerostomatella and certain specimens of muixolitha. The enlarged scale bases of cevostomatella are stronger and appear to be fewer. O. mixolitha possesses no enlarged scale bases on the vertex except a single row above the transfrontal sulcus. The frenulum of the @ is triple. The @ genitalia do not differ from those of mixolitha, cerostomatella and mongolica. BroLocy. Host-plant unknown. Moths have been taken by various collectors in March—June and October. DISTRIBUTION. Algeria; Tunisia. Ornativalva cornifrons sp. n. (Pl. 1, fig. 13; Pl. 4, fig. 29; Pl. 12, figs 78-80; Pl. 10, fig. ror; Pl. 27, fig 3,9. 5:0-6:0mm. Head (PI. 12, figs 78-80) with strong frontal process with smaller dorsal and ventral processes at base, surrounded by ring of approximately 20 strongly enlarged scale bases. Head white, narrow stripe along eye brown. Labial palpus white, outer surface of first segment brown; second segment with loose brush of long white and brown scales below. Thorax white, tegula brown. Fore wing with dark brown longitudinal band which reaches costa before apex and is interrupted near apex. Termen with dark line along base of fringes. Fringes with dark dividing line and tips. Frenulum of 2 double. GENITALIA g (Pl. 19, fig. ror). Eighth tergite weakly sclerotized, pair of narrow transverse stronger sclerotizations on anterior margin and pair of larger ones posteriorly near lateral margin. Posterior margin of uncus with number of long setae, medially with small emargina- tion. Anterior margin of tegumen with deep emargination. Saccus long, wide, gradually narrowing, distally rounded. Anellus lobes large, triangular, setose. Costa distinctly longer than valva, narrow, with parallel margins, apical portion not clavate. Harpe very short, with long terminal seta. Valva almost triangular, outer surface near dorsal margin with long stiff seta in group of shorter setae. Sacculus not separated from valva. Aedeagus with bulbous base; curved apical portion strong, apex rounded. GENITALIA @ (Pl. 27, fig. 123). Posterior margin of seventh tergite sinuate. Apophyses anteriores very short, bases connected by narrow sclerotization along anterior margin of eighth sternite. Antrum with short sclerotized ring, opens directly into corpus bursae; no long ductus bursae. Corpus bursae elongate, inner surface minutely spined; cervix bursae and antrum without minute spines. Signum near middle of corpus bursae, transverse, with pair of serrated ridges. REMARKS. 0. cornifrons differs from most Ornativalva species by the conspicuous longitudinal band of the fore wing. 0. cerostomatella with a similar band is generally larger and the ground colour of the fore wing is ochreous, not white. The thorax REVISION OF GENUS ORNATIVALVA 139 of cerostomatella is divided by a dark longitudinal line and the head lacks a frontal process. In the 3 genitalia cornifrons differs from all other species of the cerosto- matella-group (except acutivalva) by the uncus, which is not deeply divided, the short triangular valva and the large saccus. The aedeagus is similar to that of curvella but differs in details of the apex structure. The genitalia are similar to those of afghana. The periostial region is completely membranous whereas in afghana it is partially sclerotized. The posterior margin of the seventh tergite is sinuate in corniforns but straight in afghana. BioLtocy. Host-plant unknown. The type-specimens have been collected by Kaszab in June—July in a locality where Tamarix occurs (Kaszab, 1968 : 31). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Monco.ra: Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (ITM, Budapest). Paratypes. Moncotta: 10 4, 3 9, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150m, 3.vii.1967 (Kaszab, no. 882) (TM, Budapest; BMNH); 1 3, Bajanchongor aimak, oasis Dzun mod, 100 km S. of Somon Schine Zinst, 1300 m, 29.v1.1967 (Kaszab, no. 869) (TM, Budapest). Ornativalva cerostomatella (Walsingham, 1904) (Pl. 9, figs 60-62; Text-fig. 11) Gelechia cerostomatella Walsingham, 1904, Entomologist’s mon. Mag. 40 : 266. Holotype 9 [not 6, as stated by Walsingham], ALGERIA: Biskra, 15.iv.1903 (Walsingham) (BMNH) [examined]. Gelechia tripartitella Mabille, 1907, Bull. Soc. ent. Fy. 1907 : 79. Lectotype 9, Tunisia: Nefta, Djerid, v. 1904 (Lucas) (genitalia slide no. 369c, Sattler; MNHN, Paris), designated by Viette & Fletcher (1968 : 393) [as holotype] [examined]. [Synonymized by Chrétien, 1917 : 469.] Gelechia cerostomatella Walsingham; Chrétien, 1917 : 469. Gelechia cerostomella Walsingham; Caradja, 1920 :99. Incorrect subsequent spelling of cevostomatella Walsingham. Gelechia cevostomella Meyrick, 1925a, Geneva Insect. 184:79. Unjustified emendation of cerostomatella Walsingham. Gelechia biclavata Meyrick, 1934, Exot. Microlepidopt. 4:511. Holotype g, Inp1a: Great Indian Desert, xi. 1930 (Glennie) (genitalia slide no. 8288, Clarke; BMNH) [examined]. [Synonymized by Sattler, 1967 : 83.] Gelechia biclavata Meyrick; Gaede, 1937 : 150. Gelechia cerostomella [sic] Walsingham; Gaede, 1937 : 153. Gelechia tripartitella Mabille; Gaede, 1937 : 223. Gelechia cerostomatella Walsingham; Amsel, 1940 : 48. Gelechia cerostomatella Walsingham; Amsel, 1958 : 80. Ornativalva cerostomatella (Walsingham) Sattler, 1960 : 60, pl. 17, fig. 74. Ornativalva cerostomatella (Walsingham) ; Sattler, 1967 : 83, pl. 2, fig. 13, pl. 8, fig. 42, pl. 15, fig. 66. Gelechia tripartitella Mabille; Viette & Fletcher, 1968 : 393. Gelechia biclavata Meyrick; Clarke, 1969 : 108, pl. 54, figs 1-1b. Head (PI. 9, figs 60-62) without frontal process. Scale bases enlarged to coarse teeth. Gentle are of enlarged scale bases above transfrontal sulcus; some scattered enlarged scale bases on 140 Ie. SAI vertex. Frons below transfrontal sulcus covered with enlarged scale bases, particularly dense concentration in centre of frons. GENITALIA g. Sattler, 1967, pl. 8, fig. 42. GENITALIA 9. Sattler, 1967, pl. 15, fig. 66. Remarks. The head structure is very similar to that of typical mixolitha; it differs by the presence of enlarged scale bases on the vertex in addition to the arc above the transfrontal sulcus. The frenulum of the @ is triple. The 9 genitalia do not differ from those of mixolitha, angulatella and mongolica. Gelechia tripartitella Mabille was described from an unspecified number of specimens. The specimen recorded as the ‘holotype’ by Viette & Fletcher should therefore be considered the lectotype. Nothris minutella Turati, 1929, Boll. Lab. Zool. gen. agr. Portict 23 : 124, fig. 4, was previously synonymized with O. cerostomatella (Walsingham) on the basis of Turati’s description and poor photographic illustration (Sattler, 1967 : 83). Turati based his description on three specimens; however, in his collection there is now only one 9 without abdomen left. This specimen was made available to me by Count F. Hartig (IE, Bolzano) and bears the following labels: ‘Typus’ — ‘Tripolitania Nord-Africa, Sidi Mesri, 21-23 Marzo 1924, Romet’ —‘Nothris minutella Trt, Typ’. It proved to be an Anarsia species (possibly a synonym of Anarsia luticostella Chrétien, 1915) and is here removed from synonymy: Anarsia minutella (Turati, 1929) comb. n., sp. rev. Consequently Libya must be eliminated from the dis- tribution of cerostomatella, although the species will undoubtedly be found there. BroLtocy. Host-plant unknown. Moths have been taken by various collectors in February—August. DISTRIBUTION. Mauritania; Algeria; Tunisia; Egypt; Saudi Arabia; India. FURTHER MATERIAL EXAMINED. MAURITANIA: I 9, between Kiffa and Tidjidja, 1931 (Steele) (BMNH) [first record for Mauritania]. EGypt: 1 9, Romani, 30.vii.1g16 (Boyd) (BMNH) [first record for Egypt]. Ornativalva mongolica Sattler, 1967 (Pl. 1, fig. 10; Pl. 4, fig. 30; Pl. ro, figs 63-65) Ornativalva mongolica Sattler, 1967, Beitr. naturk. Forsch. SiidwDil. 26 (3) : 85, pl. 18, fig. 76. Holotype ¢, Moneoria: East Gobi aimak, Cagan Elis, 30 km ESE. of Zuun-Bajan, 800 m, 22.v1.1963 (Kaszab, no. 21) (TM, Budapest) [examined]. Head (PI. ro, figs 63-65) with frons strongly prominent but without definite process. Scale bases very‘strong, arranged in irregular rings on frons; small group on vertex above transfrontal sulcus. GENITALIA g. Sattler, 1967, pl. 18, fig. 76. REMARKS. The frontal structure of mongolica is similar to that of cerostomatella but the frons is more prominent and the enlarged scale bases are fewer and stronger. REVISION OF GENUS ORNATIVALVA 141 The frenulum of the 2 is double. The Q genitalia do not differ from those of mixolitha, angulatella and cerostomatella. BroLtocy. Host-plant unknown. Moths have been collected by Kaszab in June-July in a locality where Tamarix occurs (Kaszab, 1968 : 31). DISTRIBUTION. Mongolia. FURTHER MATERIAL EXAMINED. Monco iA: 3 3, Uburchangaj aimak, 130 km ESE. of Somon Bajanleg, 1150 m, 3.vii.1967 (Kaszab, no. 882) (IM, Budapest; BMNH); 1 9, Bajanchongor aimak, 8 km ESE. of Somon Bajanleg, 1350 m, 2.vil.1967 (Kaszab, no. 879) (TM, Budapest); 2 9, Bajanchongor aimak, SE. corner of lake Orog nur, 1200 m, 23.vi.1964 (Kaszab, no. 183) (ITM, Budapest). Ornativalva afghana Sattler, 1967 (PI. i, figs 11, 12; Pl. 4, fig. 31; Pl. 11, figs 69-74; Pl. 12, figs 75-77; Pl. 20, fig. 102; Pl 27;,-fig.. £24) Ornativalva afghana Sattler, 1967, Beitr. naturk. Forsch. SiidwDil. 26 (3) : 75, pl. 17, fig. 72, pl. 18, fig. 77; pl. 19, fig. 79. Holotype g, AFGHANISTAN: SE. of Shindan, ca 150 km S. of Herat, 25.vi.1963 (Kasy & Vartian) (genitalia slide no. 3276;-NM, Vienna) [examined]. Head (PI. 11, figs 69-74; Pl. 12, figs 75-77) with enlarged scale bases fused to form strong arc above transfrontal sulcus. Entire frons densely set with enlarged scale bases. Frontal process short, truncate. GENITALIA ¢ (PI. 20, fig. 102). GENITALIA Q (PI. 27, fig. 124). Remarks. The Mongolian specimens differ from the type-series from Afghanistan. On the head of Mongolian specimens the arc above the transfrontal sulcus is more strongly developed, the frontal process is approximately twice the length of that of specimens from Afghanistan and the enlarged scale bases on the frons are fewer and stronger. The fore wings of Mongolian specimens appear narrower and the line that separates the dark costal and the light posterior half of the wing is clearly W-shaped in the fold. In the ¢g genitalia the harpe is narrower in Mongolian specimens. In the 9 genitalia the eighth tergite of specimens from Afghanistan has a distinct dent above the spiracle (this dent is not visible in the illustration given by Sattler, 1967, pl. 17, fig. 72) and the sclerotization on the anterior margin of the eighth sternite is much broader than in the Mongolian specimens and surrounds most of the ostium bursae. The frontal structure of the Mongolian specimens resembles that of mixolitha specimens from Turkey and Mongolia, but the frontal process is clearly truncate in afghana. The frenulum of the 9 is triple. BioLtocy. MHost-plant unknown. The Mongolian specimens have been collected 142 K. SATTLER by Kaszab in a locality where Tamarix occurs (Kaszab, 1966: 591). Moths have been collected by Kasy and Vartian and Kaszab at the end of June. DisTRIBUTION. Afghanistan; Mongolia. FURTHER MATERIAL EXAMINED. Monco ta: 6 4, 1 9, Gobi Altaj aimak, Zachuj Gobi, 10 km N. of Chatan chajrchan mountains, I150m, 27.vi.1966 (Kaszab, no. 594) (IM, Budapest; BMNH) [first record for Mongolia]. Ornativalva curvella sp. n. (Pl. 10, figs 66-68; Pl. 20, fig. 103; Text-figs 26, 27) 3. 770mm. Head (PI. ro, figs 66-68) with enlarged scale bases on vertex fused to form strong arc above transfrontal sulcus. Enlarged scale bases on frons strong, arranged in irregular rings, densely concentrated below transfrontal sulcus; frons irregularly prominent. Labial palpus pale, outer surface of second segment with light brown patches near base and below apex. Costal half of fore wing greyish brown, darkest at fold, with two dark angular projections extending across fold into lighter posterior half of wing. Small dark brown dot in cell and slightly larger dot at end of cell. GENITALIA ¢ (PI. 20, fig. 103; Text-figs 26, 27). Eighth tergite membranous with narrow transverse sclerotized band on anterior margin and pair of longitudinal bands laterally near posterior margin. Eighth sternite rounded, weakly sclerotized. Uncus strongly developed, posteriorly truncate, lateral portion strongly sclerotized, large posterior area of tegumen membranous. Inner surface of uncus set with small number of long setae near posterior margin. Costa nearly as long as valva, narrow, distal two-thirds strongly curved dorsad. Harpe a broad rounded process at base of costa. Distal two-thirds of valva narrow, distal third strongly curved dorsad, apex rounded, without terminal spine. Anellus lobes rounded. Saccus large. Aedeagus with clearly defined bulbous base and narrow, strongly curved apical portion. GENITALIA 9. Unknown. RemaArRKS. The frontal structures of cwrvella are similar to those of mongolica but differ by the clearly defined arc above the transfrontal sulcus and the irregular process below the sulcus. The only specimen of curvella is poorly preserved. The 3 genitalia with the well developed uncus and strongly curved costa and valva do not closely resemble those of any other species. BioLocy. MHost-plant unknown. The holotype has been collected by Kaszab at light in a locality where Tamarix occurs (Kaszab, 1966 : 591). DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype g, Moncoria: Gobi Altaj aimak, Zachuj Gobi, 10 km N. of Chatan chajrchan mountains, 1150 m, 27.vi.1966 (Kaszab, no. 594) (genitalia slide no. 647d, Sattler; TM, Budapest). Ornativalva acutivalva sp. n. (PI. 1, fig. 14; Pl. 4, ng: 32: Pl 20, fig. ro4; Pl 2752. 725) 3,2. 5:0-6:0mm. Head without enlarged scale bases or frontal process. Metascutum with paired group of narrow, hair-like scales (Text-fig. 2). Head ochreous. Labial palpus whitish, REVISION OF GENUS ORNATIVALVA 143 second segment with moderate brush below, only twice as wide as third segment, with brown bands near base and apex with some scattered brown scales in between; apex and broad ring around middle of third segment brown. Scape of antenna brown with white distal ring; flagellum above dark with lighter rings. Thorax ochreous, tegula brown with ochreous apex. Fore wing narrow, costal half mostly dark brown, dorsal half ochreous; dark area distally narrower, not reaching apex, barely reaching fold. Projections of dark area touch fold at one-third and two-thirds without crossing it. Small dark dots in cell and at end of cell in ochreous area but connected with dark area. Some dark scales along termen. GENITALIA ¢ (Pl. 20, fig. 104). Uncus pointed. Costa slender, as long as valva. Harpe short, rounded. Valva broad at base, tapering, apex rounded, without terminal spine. No separate sacculus. Anellus lobes broad at base, bulging, narrowed at distal third. Aedeagus with bulbous base, distal two-thirds strong, bent near middle, apex rounded. GENITALIA Q (PI. 27, fig. 125). Apophysis posterior as long as eighth segment. Apophysis anterior very short. Eighth tergite with irregular rows of scale bases at posterior margin. Ductus bursae with open sclerotized ring near ostium bursae. Ductus bursae short, wide, not coiled, gradually widening into corpus bursae. Signum oval, both ends serrated, with pair of transverse folds. REMARKS. The structure of the anellus and the absence of a separate sacculus in the ¢ genitalia and the presence of scales — not setae — on the eighth tergite, the short apophyses anteriores and the structure of the signum in the 9 genitalia place acutivalva in the cerostomatella-group. O. acutivalva differs from all other species of the cerostomatella-group by the absence of frontal modifications of the head and the presence of long scales on the metascutum. The anellus lobes are similar to those of angulatella and cerostomatella. All three available 2 have a variable number of frenulum setae. Two specimens have three setae on the left hind wing and two on the right whereas the third specimen has two on the left and three on the right. BroLocy. MHost-plant unknown. Moths have been collected by Kaszab in July. DISTRIBUTION. Mongolia. MATERIAL EXAMINED. Holotype 3, Moncotta: South Gobi aimak, 10 km NNE. of the town Dalanzadgad, 1450 m, 7.vii.1967 (Kaszab, no. 898) (TM, Budapest). Paratypes. MONGOLIA: 10 4, 3 9, South Gobi aimak, 10 km NNE. of the town Dalanzadgad, 1450 m, 7.vii.1967 (Kaszab, no. 898) (TM, Budapest; BMNH); 1 3, Middle Gobi aimak, 8 km NW. of Oldoch Chijd, 54 km NNW. of Somon Zogt-Ovoo, 1350 m, 9.vli.1967 (Kaszab, no. 905) (TM, Budapest). BIBLIOGRAPHY AGENJO, R. 1968. Catalogo ordenador de los Lepidépteros en Espana. Gvraellsia 23, Appendix : [1]-[6]. AmseL, H. G. 1933. Die Lepidopteren Palastinas. Eine zoogeographisch-dkologisch- faunistische Studie. Zoogeographica 2 : 1-146, text-figs I, 2. — 1935a. Zur Kenntnis der Microlepidopterenfauna des stidlichen Toten-Meer-Gebietes, nebst Beschreibung neuer palastinensischer Macro- und Microlepidoptera. Vevréff. dt. Kolon. u. Ubersee-Mus. Bremen 1 : 203-221, pls 11, 12. 19350. Weitere Mitteilungen tiber palastinensische Lepidopteren. Verdff. dt. Kolon. u. Ubersee-Mus. Bremen 1 : 223-277. 144 K. SATTLER 1940. Ueber alte und neue Kleinschmetterlinge aus dem Mittelmeer-Gebiet. Verdff. dt. Kolon. u. Ubersee-Mus. Bremen 3 : 37-56, text-figs 1-4, pl. 3. 1955a. Uber mediterrane Microlepidopteren und einige transkaspische Arten. Bull. Inst. vy. Scr. nat. Belg. 31 (83) : 1-64, text-figs 1-5, pls 1-6. 19550. Irakische Kleinschmetterlinge. Betty. naturk. Forsch. SiidwDil. 14 : 119-129, text-figs I-15, pl. 6. 1956. Kleinschmetterlinge vom mittleren Jordantal. Beity. naturk. Forsch. SiidwDil. 15 : 56-62, text-figs 1-6, pl. 4. 1958. Kleinschmetterlinge aus Nordost-Arabien der Ausbeute A. S. Talhouk. Beitr. naturk. Forsch. SiidwDtl. 17 : 61-82, text-figs 1-23, pl. 5. 1959. Microlepidoptera aus Iran. Stuttg. Beitr. Naturk. 28 : 1-47, text-figs 1-3, pls I-5. 1966. Zur Kenntnis der Microlepidopterenfauna von Marokko. WNotul. ent. 46 : 125-130, map I, text-fig. 1. 1968. Zur Kenntnis der Microlepidopterenfauna von Karachi (Pakistan). Stuttg. Beitr. Naturk. 191 : 1-48, pls 1-12. ANDRES, A. 1913. Verzeichnis der bis jetzt in Aegypten beobachteten Schmetterlinge. Bull. Soc. ent. Egypte 3 : 53-114. Baum, B. 1966. Monographic revision of the genus Tamarix. 193 pp., 120 text-figs. Jeru- salem. BODENHEIMER, F. S. 1930. Die Schadlingsfauna Palastinas. Unter besonderer Beriicksich- tigung der Grossschadlinge des Mittelmeergebietes. Monogrn angew. Ent. 10, xv + 438 pp., 139 text-figs, I map. 1935. Animal lifein Palestine. [vi] + 506 pp., 77 text-figs, 70 pls. Jerusalem. Carapja, A. 1920. 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Jn Staudinger, O. & Wocke, M., Catalog dev Lepidopteren des euvopaeischen Faunengebiets. xxxviii + 426 pp. Dresden. ZELLER, P. C. 1850. Verzeichniss der von Herrn Jos. Mann beobachteten Toscanischen Microlepidoptera. (Fortsetzung.). Stettin. ent. Ztg 11 : 139-162. ZERNY, H. 1936. Die Lepidopterenfauna des Grossen Atlas in Marokko und seiner Rand- gebiete. Mém. Soc. Sci. nat. Phys. Maroc 42 : 1-163, text-figs 1-4, pls 1, 2. [Dated *31 Décembre 1935’ on original wrapper and title page. Page [164] marked ‘Achevé d’imprimer le 1 Aout 1936’.] Zoccut, R. 1971. Contributo alla conoscenza dell’entomofauna delle Tamerici in Italia. Redia 52 : 31-129, text-figs 1-22. 148 K. SATTLER Fics 3-8. Wing venation of Ovnativalva species. (H—humeral field.) 3, O. heluanensis (Debski), 9. 4, O. evubescens (Walsingham), g. 5, O. ovnatella Sattler, 9. 6, O. plicella sp.n.,Q. 7, O. tamariciella (Zeller), 9. 8, O. indica Sattler, ¢. REVISION OF GENUS ORNATIVALVA 149 Fics 9-14. 9-12, wing venation of Ovnativalva species. (9) O. kalahariensis (Janse), . (10) O. plutelliformis (Staudinger), §. (11) O. cerostomatella (Walsingham), ee (2) Oe mixolitha (Meyrick), 9. 13-14, O. ornatella Sattler. Outlines of ¢ left valvae. (13) Mongolia (slide no. 14 878; BMNH). (14) Rumania (slide no. 686, Sattler; MINGA, Bucharest). 150 K. SATTI LER Fics 15-27. 15-17, Ovnativalva ornatella Sattler. Outlines of 9 apophyses anteriores. (15) Mongolia (slide no. 14 875; BMNH). (16) paratype, Iran (slide no. 3239; NM, Vienna). (17) S. Russia (slide no. 7288; BMNH). 18-20, O. kalahariensis (Janse). Outlines of g valvae. (18) left valva, holotype, S. Africa (slide no. 5800, Janse; TM, Pretoria). (19) right valva, holotype, S. Africa (slide no. 5800, Janse; TM, Pretoria). (20) right valva, SW. Africa (slide no. 18 891; BMNH). 21, O. singula Sattler. Harpe of g, Mongolia (slide no. 649a, Sattler; TM, Budapest). 22-25, seventh abdominal. segments of Orvnativalva Q. (22) O. levifrons sp. n., paratype, Mongolia (slide no. 15 379; BMNH). (23) O. species 1, Mongolia (slide no. 650, Sattler; TM, Budapest). (24) O. ornatella Sattler, Mongolia (slide no. 14 875; BMNH). (25) O. singula Sattler, Mongolia (slide no. 15 731; BMNH) (sternite left, tergite right). 26-27, O. curvella sp. n., holotype g, Mongolia (slide no. 647d, Sattler; TM, Budapest). (26) eighth abdominal tergite. (27) uncus and tegumen. REVISION OF THE GENUS ORNATIVALVA I51 ' INDEX Synonyms and unavailable names are in ?falics. Page numbers of principal references are in bold. acutivalva sp. n., 97, 98, IOI, 135, 139, 142, 143 afghana Sattler, 96, 98, 1o1, 139, 141 Agdistis Hiibner, 102 Altenia Sattler, 95 Anarsia Zeller, 87, 140 angulatella Chrétien, 96, 98, ror, 136, 137, 138, I40, 141, 143 antipyramis Meyrick, 97, 99, 100, 124, 128, 129, 132 arabica Sattler, 97, 99, 100, 107, 114, 119 aspera sp. n., 96, 98, 100, 107, 108, 109, 110, 114 auctella nomen nudum, 128, 129, 131, 132 basistriga sp. n., 97, 99, 101, 130 biclavata Meyrick, 101, 139 bipunctella Sattler, 101, 136, 137 caecigena Meyrick, 92, 97, 100, 123 Cerofrontia Janse, 92 cerostomatella Walsingham, 87, 92, 93, 96, 98, IOI, 136, 138, 139, 140, 141, 143 cevostomella Meyrick, to1, 139 Chionodes Hiibner, 95, 96 _ cimelion Amsel, 89, 100, 106 cornifrons sp. n., 92, 93, 96, 98, 101, 138, 139 Cosmopterigidae, 92 Crambinae, 92 curvella sp. n., 96, 98, 101, 139, 142 erubescens Walsingham, 97, 98, 100, 107, 108, 115, 116 frankeniworella Chrétien, 100, 101, 102 _ Friseria Busck, 95 frontella sp. n., 92, 96, 98, 100, 114 Gelechiidae, 92, 93, 95 Gelechiinae, 95, 96 Gelechiini, 96 Geometridae, 92 Gladiovalva Sattler, 95 Gnorimoschemini, 96 | grisea Sattler, 97, 99, 101, 130, 132, 134 gussakovskii Gerasimov, 93 haligmatodes misspelling, 128, 131 heligmatodes Walsingham, 97, 98, 100, 121, 124, 125, 128 | | heluanensis Debski, 94, 95, 97, 99, 100, 101, TO2, 10g, 12%, 123, 124, 126 _ ignota Sattler, 97, 99, 100, 106, 108, 115, 116 Ilseopsis Povolny, 92 indica Sattler, 97, 100, 120, 121, 126 ivanella Sattler, 100, 106 kalahariensis Janse, 96, 98, 100, 127 lacertella Walsingham, 128 levifrons sp. n., 97, 99, 100, 107, 109, 110, 111 lilyella Lucas, 96, 98, 100, 107, 108, r10, 119 Lita Treitschke, 92 longiductella Sattler, 93, 98, 99, 100, Io1, 104, 105 luticostella Chrétien, 140 macrosignella Sattler, 97, 99, 100, 102, 119, 123, 124, 125, 126 Melitoxoides Janse, 95 minutella Turati, 87, 140 ‘misma sp. n., 97, 99, 100, 119, 121 mixolitha Meyrick, 92, 93, 96, 98, 1o1, 135, 136, 137, 138, 140, I41 mongolica Sattler, 96, 98, tor, 136, 138, 140, 142 nigvosubvittatella Lucas, 1o1, 138 Noctuidae, 92 oasicolella Turati, 100, 102, 103 ochraceofusca Sattler, 97, 99, 100, 111, 112 olbiaella Milliére, 101, 131, 132 olbienella misspelling, 131 ornatella Sattler, 96, 98, 100, I11, 112, 116 Ornativalva Gozmany, 91, 95 Pelostola Janse, 91, 95, 100 pharaonis Sattler, 92, 97, 99, 100, 119, 120, 121 Phycitinae, 92 plicella sp. n., 97, 99, 100, 118, 119 plutelliformis Staudinger, 94, 95, 97, 99, IOI, 102, 108, 117, 128, £29, 130, 131i) 132; 133, 134 pseudotamariciella Sattler, 97, 99, 100, 126 Pterophoridae, 102 152 INDEX pulchella sp. n., 92, 96, 98, 100, 107, 109, 110, 114 Pyralidae, 92 Rhynchopacha Staudinger, 92, 93, 95 roseosuffusella Sattler, 98, 99, 100, 105 rufipuncta sp. n., 93, 94, 97, 99, 100, IOI, 104, 105 Scrobipalpa Janse, 92, 95 Scythrididae, 93 serratisignella Sattler, 92, 97, 100, 122 sesostrella Rebel, 97, 99, 100, 115, 116 siculella Mariani, 100, 102, 103 sieversi Staudinger, 97, 98, IOI, 117, 1309, 134 stewersiellus Christoph, 89, ror, 131, 132 singula Sattler, 97, 98, 101, 129, 130 K. SATTLER, Dr. rer. nat. Department of Entomology BritisH Museum (NATURAL History) - CROMWELL RoapD Lonpon SW7 5BD sinuatella Walsingham, to1, 131, 132 Symmocidae, 92 tamaricalis misspelling, 102, 124 tamaricella misspelling, 120 tamariciella Zeller, 94, 97, 99, 100, IOI, 102, 120, 123, 124, 125, 126 Teleiodini, 96 Thaumetopoeidae, 92 Thyrididae, 92 triangulella Sattler, 97, 99, 100, 128, 129, 132 trifasciella Rebel, 102 tyipartitella Mabille, ror, 139, 140 undella sp. n., 96, 99, 100, 118, 119 zonella Chrétien, 98, 99, 100, Io1, 105, 106 IPL ACAD I, at Fic. 1. Ovnativalva levifrons sp.n. @, holotype. Mongolia. (IM, Budapest.) Fic. 2. O.asperasp.n. <4, holotype. Mongolia. (IM, Budapest.) Fic. 3. O. pulchellasp.n. 4, holotype. Mongolia. (IM, Budapest.) Fic. 4. O. frontellasp.n. 3, holotype. Mongolia. (IM, Budapest.) Fic. 5. O. ochvaceofusca Sattler. ¢g. Turkey. (NM, Vienna.) Fic. 6. O.mismasp.n. 6, holotype. Sudan. (BMNH.) Fic. 7. O. caecigena (Meyrick). 9. Pakistan. (NM, Vienna.) Fic. 8. O. basistrigasp.n. 4, holotype. Mongolia. (IM, Budapest.) Fic. 9. O. mixolitha (Meyrick). 9. Turkey. (BMNH.) Fic. 10. O. mongolica Sattler. 9, paratype. Mongolia. (BMNH.) Fic. 11. O. afghana Sattler. g. Mongolia. (IM, Budapest.) Fic. 12. O. afghana Sattler. 9, paratype. Afghanistan. (BMNH.) Fic. 13. O.cornifrons sp.n. 4, holotype. Mongolia. (IM, Budapest.) Fic. 14. O. acutivalva sp.n. 4, holotype. Mongolia. (IM, Budapest.) Fic. 15. O.undellasp.n. 9, holotype. Iran. (NM, Vienna.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PEATE + Fic. Fic. Fic. Fic. Fic. Fic. 16. IOs 18. 1); 20. 20. IPILANINIS, 2 Ornativalva vufipuncta sp.n. 4, paratype. Iran. (NM, Vienna.) O. levifrons sp.n. 4, holotype. Mongolia. (IM, Budapest.) O. aspera sp.n. 9, paratype. Mongolia. (ITM, Budapest.) O. pulchellasp.n. 4, paratype. Mongolia. (IM, Budapest.) O. frontellasp.n. 64, holotype. Mongolia. (IM, Budapest.) O. plicellasp.n. 4, paratype. Iran. (NM, Vienna.) Bull. By. Mus. nat. Hist. (Ent.) 3 2 7s PAM 92 Fic. JOE. Fic. Fic. Fic. Fic. Nn NN WN WN ~—I An ew tN iS) Ornativalva undella sp. n. . misma sp.n. 4, holotype. . sevvatisignella Sattler. g. Iran. . singula Sattler. . basistriga sp. n. . basistviga sp. n. SISISISTS PLATE 3 3. Mongolia. 3, holotype. Q, paratype. 9, holotype. Iran. Sudan. (BMNH.) Mongolia. Mongolia. (NM, Vienna.) (BMNH.) (BMNH.) (TM, Budapest.) (ITM, Budapest.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PEATE, 3 Fic. Fic. Fic. FTG Fic. PLAGE Ornativalva mixolitha (Meyrick). 9. Mongolia. (IM, Budapest.) O. cornifrons sp.n. 4, holotype. Mongolia. (IM, Budapest.) O. mongolica Sattler. 9. Mongolia. (BMNH.) O. afghana Sattler. g. Mongolia. (BMNH.) O. aciutivalua sp.n. 4, holotype. Mongolia. (ITM, Budapest.) Bull. By. Mus. nat. Hist. (Ent.) 34, 2 PLATE 4 a the Fics 33-35: Fics 36-38. : be. PLATE 5 Ornativalva heads; frontal, lateral and dorsal view Ornativalva plutelliformis (Staudinger). 9. Sudan. O. evubescens (Walsingham). 9. Sudan. (BE MNH U —— PLATE 5 nt.) 34, 2 (E nat. Hist. Mus. Bull. Br. \ i. PLATE 6 Ornativalva heads; frontal, lateral and dorsal view Fics 39-41. Ornativalva lilyella (Lucas). ¢g. Algeria. Fics 42-44. O.asperasp.n. 3, paratype. Mongolia. Bull. Br. Mus. nat. Hist. (Ent.) 3 IPIVINADIS, Ornativalva heads; frontal, lateral and dorsal view Fics 45-47. Ornativalua pulchella sp.n. g, paratype. Mongolia. | Fics 48-50. O. frontellasp.n. g, paratype. Mongolia. (TM, Bu Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 7 \ cs Rieke Ornativalva heads; frontal, lateral and dorsal view | Fies 51-53. Ornativalva mixolitha (Meyrick). 9. Sudan Fics 54-56. O. mixolitha (Meyrick). 9. Turkey. (B 8 PLATE Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PILINANS, 6) Ornativalva heads; frontal, lateral and dorsal view Fics 57-59. Ovnativalva angulatella (Chrétien). 3. Algeria. (MNH N, Fics 60-62. O. cerostomatella (Walsingham). 9. Tunisia. (BMNH.) Bull. Bry. Mus. nat. Hist. (Ent.) 34, 2 PLATE 9 ~~ PLATE to Ovnativalva heads; frontal, lateral and dorsal view Fics 63-65. Ovnativalua mongolica Sattler. 3. Mongolia. (BMNH.) Fics 66-68. O.curvellasp.n. 3, holotype. Mongolia. (TM, Bu Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE tro \ Ss PLATE oe Oynativalva heads; frontal, lateral and dorsal view Fics 69-71. Ovnativalva afghana Sattler. Q, paratype. Afghanistan. Fics 72-74. O. afghana Sattler. 3. Mongolia. (BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, > PLATE Il Fic. Fic. TG 7 Fic. Fic. Fic. PLATE 22 Oynativalva head structures Oynativalva afghana Sattler. 9, paratype. Afghanistan. (BMNH.) — Frontal process. O. afghana Sattler. 9, paratype. Afghanistan. (BMNH.) — Modified scale of frons. O. afghana Sattler. 9, paratype. Afghanistan. (BMNH.)—Section of modified scale of frons. O. cornifrons sp.n. @, paratype. Mongolia. (BMNH.) — Frontal view. O. cornifrons sp.n. , paratype. Mongolia. (BMNH.) — Lateral view. O. cornifrons sp.n. , paratype. Mongolia. (BMNH.) — Dorsal view. Bull. By. Mus. nat, Hist. (Ent.) 34, 2 PLATE PIL NAD IS es Genitalia 3 Fic. 81. Ovnativalva heluanensis (Debski). Mongolia. (Slide no. 14 881, BMNH.) Fic. 82. O. longiductella Sattler. Mongolia. (Slide no. 14 893, BMNH.) Fic. 83. O. vufipuncta sp. n., paratype. Iran. (Slide no. 19 854, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PAR ACI, ns 81 es, 82 % y ; ’ & ee. th : & ‘i ‘ u7 83 PLATE 14 Genitalia ¢ Fic. 84. Ovnativalva levifrons sp. n., paratype. Mongolia. (Slide no. 15 382, BMNH.) Fic. 85. O. aspevasp.n., paratype. Mongolia. (Slide no. 14 896, BMNH.) Fic. 86. O. pulchella sp. n., holotype. Mongolia. (Slide no. 635d, Sattler; TM, Budapest.) Bull. By. Mus. nat. Hist. (Ent.) 34, 2 PEA 4 PLATE 15 Genitalia ¢ Fic. 87. Ovnativalva ochraceofusca Sattler. Turkey. (Slide no. 3868, NM, Vienna.) Fic. 88. O. species 1. Mongolia. (Slide no. 15 784, BMNH.) = Fic. 89. O. species 2. Mongolia. (Slide no. 656, Sattler; TM, Budapest.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PHATE 15 87 8 89 _. [ a ee 4 = _ \ _ EDs. " _ = - 4 * _ _ > 4 } = i { 4 PVAT E16 Genitalia 3 Fic. 90. Ovnativalva species 3. Mongolia. (Slide no. 636d, Sattler; TM, Budapesay Fic. 91. O. frontella sp. n., paratype. Mongolia. (Slide no. 15 380, BMNH.) Fic. 92. O. ornatella Sattler. Mongolia. (Slide no. 14 878, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 16 90 of 91 92 IPILSN IIS, 16-7) Genitalia ¢ Fic. 93. Ovrnativalva plicella sp. n., paratype. Iran. (Slide no. 19 855, BMNH.) Fic. 94. O. misma sp.n., holotype. Sudan. (Slide no. 19 841, BMNH.) : Fic. 95. O. serrvatisignella Sattler. Iran. (Slide no. 19 850, BMNH.) Bull. By. Mus. nat. Hist. (Ent.) 34, 2 [PUL TNA, ier PEATE 13 Genitalia 3 Fic. 96. Ovnativalva caecigena (Meyrick). Kuwait. (Slide no. 18 890, BMNH.) Fic. 97. O.species 5. Iran. (Slide no. 3882, NM, Vienna.) Fic. 98. O. singula Sattler. Mongolia. (Slide no. 14 889, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 18 NG, Ce). Fic. 100. Fic. ror. PLATE 19 Genitalia ¢ Ornativalva basistriga sp. n., paratype. Mongolia. (Slide no. 14 874, BMNH.) | O. species 6. Mongolia. (Slide no. 14 888, BMNH.) O. cornifrons sp. n., paratype. Mongolia. (Slide no. 15 730, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 19 99 100 101 PEATE Zo Genitalia 3 Fic. 102. Ovnativalva afghana Sattler. Mongolia. (Slide no. 15 378, BMNH.) Fic. 103. O. curvella sp. n., holotype. Mongolia. (Slide no. 647d, Sattler; TM, Budapest.) Fic. 104. O. acutivalva sp. n., paratype. Mongolia. (Slide no. 14 882, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 20 12 103 ga" — : ¢ ' - as — ‘am - SL } 2 Be -o PVAGE 2a Genitalia 2 Fic. 105. Ovrnativaluva heluanensis (Debski). Mongolia. (Slide no. 637d, Sattler; Budapest.) : Fic. 106. O. vufipuncta sp. n., paratype. Iran. (Slide no. 3885, NM, Vienna.) Fic. 107. O. levifrons sp. n., paratype. Mongolia. (Slide no. 15 379, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 AI Bai IRIDANIT I, BZ Genitalia 9 Fic. 108. Ovnativalva aspera sp. n., paratype. Mongolia. (Slide no. 14 877, BMNH.) Fic. 109. O. pulchella sp. n., paratype. Mongolia. (Slide no. 14 869, BMNH.) Fic. 110. O. ochvaceofusca Sattler. Afghanistan. (Slide no. 15 787, BMNH.) Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 PLATE 22 108 109 110 1G, win. IFIG, WUD. Fic. 113. IPIL GAN II, 23} Genitalia 9 Ornativalva species 1. Mongolia. (Slide no. 650, Sattler; TM, Budap O. ignota Sattler. Algeria. (Slide no. 6267, BMNH.) O. ornatella Sattler. Mongolia. (Slide no. 14 875, BMNH.) 2 PLATE 23 PLATE 24 Genitalia 9 Fic. 114. Ovrnativalva plicella sp. n., paratype. Iran. (Slide no. 18 952, BMNH.) Signa and sclerotization of cervix bursae separate. = Fic. 115. O. undella sp. n., holotype. Iran. (Slide no. 3864, NM, Vienna.) Signa and sclerotization of cervix bursae separate. (a PLATE 24 Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 SS Le 415 115 Fic. 116. BiG. 117. UE; WIS}, \ PLATE 25 Genitalia 9 Oynativalva species 4. Mongolia. O. misma sp. n., paratype. Sudan. O. species 5. Iran. (Slide no. 3865, NM, Vienna.) (Slide no. 15 783, BMNH.) (Slide no. 19 842, BMNH.) Bull. By. Mus. nat. Hist. (Ent.) 34, 2 PLATE 25 116 FEZ 118 PLATE 26 Genitalia 9 Fic. 119. Ovrnativalva singula Sattler. Mongolia. (Slide no. 15 731, BMNH.) Fic. 120. O. basistviga sp. n., paratype. Mongolia. (Slide no. 645d, Sattler; TM, Budapeel Fic. 121. O. species 6. Mongolia. (Slide no. 636c, Sattler; TM, Budapest.) PLATE 26 Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 "2 120 121 Fic. Fic. Fic. Fic. WPA. 12S. 124. 125. IGN. 25 Genitalia 9 Ornativaluva mixolitha (Meyrick). Mongolia. (Slide no. 635c, Sattler; TM, Buda- pest.) Ventral and dorsal surface of eighth segment. O. cornifrons sp. n., paratype. Mongolia. (Slide no. 14 891, BMNH.) O. afghana Sattler. Mongolia. (Slide no. 637b, Sattler; TM, Budapest.) = O. acutivalva sp. n., paratype. Mongolia. (Slide no. 14 894, BMNH.) ~ PLATE 27 Bull. Br. Mus. nat. Hist. (Ent.) 34, 2 as — - . 4 ~ i vs a ¥ he : / ‘ / - pe ; H : i ; : ” fy s é; i ; J / : . Zee / | / : / = [ ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp.172: 500 text-figures. September, 7005. 73:25: . OxapDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, g maps. February, 1967. 3.50. . HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8. . Mounp, L.A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . AFiFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. £12. . Sanps, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30 . CrosskEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND NGS ae THE TAXONOMY OF THE GENUS HETEROPELMA WESMAEL (HYMENOPTERA : ICHNEUMONIDAE) i976 é, LIBRARY . CT 4 LD. GAUDLD BULLETIN OF _ THE BRITISH MUSEUM (NATURAL HISTORY) _ ENTOMOLOGY Vol. 34 No. 3 | LONDON : 1976 THE TAXONOMY GF THE GENUS HETEROPELMA WESMAEL (HYMENOPTERA : ICHNEUMONIDAE) BY IAN DAVID GAULD Commonwealth Institute of Entomology Pp. 153-219; 64 Text-figures; 3 Charts; 2 Maps BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 3 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear ut irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 34 No. 3 of the Entomology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.). ISSN 0524-0431 © Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 26 August, 1976 Price £5.60 THE TAXONOMY or THE GENUS HETEROPELMA WESMAEL (HYMENOPTERA : ICHNEUMONIDAE) By I. D. GAULD CONTENTS Page SYNOPSIS : : ; ‘ : : : : : : : 155 INTRODUCTION : : ; : : : z : - : 155 NOMENCLATURAL SUMMARY : : ; . : : : 156 MATERIALS EXAMINED AND METHODS USED : : : : : 157 Heteropelma Wesmael : 3 : : c : : , : 158 Description ; : : ; : : ; 2 , : 158 Immature stages : : - : - : : - 159 Discussion of some characters : : : 3 : : : 160 Systematic position of genus : : : , : . : 162 Key TO SPECIES OF Hetevopelma. : : : ; : : ‘ 162 DESCRIPTIONS OF SPECIES . 5 ‘ : 3 : . : 164 RELATIONSHIPS BETWEEN THE SPECIES : ; ‘ : ; 3 201 Phenetic interrelationships . : - : : t ; 201 Proposed phylogenetic qriemelaadshigs : : : : - 203 ACKNOWLEDGEMENTS : . 2 : : : . : : 208 REFERENCES . : 2 ; - ; ‘ , ‘ . 208 INDEX TO RECORDED HOSTS ; : : - : : . : 218 INDEX TO PARASITES : : : : ; : : : : 218 SYNOPSIS The genus Heteropelma is defined and Tanypelma is included as a synonym of it. Seventeen species are recognized, four of which are new. Fifteen specific names are reduced to synonyms and one new combination is made. Six lectotype designations are made. All species are keyed and redescribed and notes on the distribution and biology, where known, are included. Intraspecific variation is discussed at length for some widely distributed species. The inter- specific relationships are discussed from both phenetic and phylogenetic standpoints. INTRODUCTION Heteropelma is a moderate-sized genus belonging to the tribe Therionini of the subfamily Anomaloninae. Species have been recorded from all regions except the Ethiopian, but the majority of species occur on high ground in the Indo-Papuan region. Although host records are scarce certain species have repeatedly been recorded as parasites of pest species of Lepidoptera and thus several species of Heteropelma are insects of economic importance. As far as is known species of this genus are only parasitic on lepidopterous larvae, especially those of the Noctuoidea, Geometroidea and Sphingoidea. Oviposition is usually into early instar larvae. 156 I. D, GAVED The ichneumonid completes development in the host puparium and the adult invari- ably emerges by biting off the extreme anterior end of the host puparium. Previous studies on this genus are mostly represented by scattered descriptions of new species and less often by keys to species of a particular region. Morley (1913a) included a key to world species represented in the collections of the British Museum (Natural History), but unfortunately he confused Heteropelma with other genera, thereby considerably reducing the value of his work. Schmiedeknecht (1936) and Uchida (1958) included useful keys to western and eastern Palaearctic species respectively. In these and all earlier works species now placed in Heteropelma were divided between two genera, Heteropelma and Schizoloma. Asa result of examining some intermediate species Townes (1971) synonymized Schizoloma with Heteropelma. A number of synonyms have been created because of the wide range of geographi- cal distribution of several species. No attempt has been made by earlier authors to ascertain the range of variation occurring in the several species. The present paper is an attempt to delimit the known species more accurately and to give some insight into the range of variation, distribution and habits of members of this interesting genus. NOMENCLATURAL SUMMARY Townes and co-workers (1951; 1961; 1965) catalogued 27 species as either Hetero- pelma or Schizoloma. At the start of this work 25 were considered to be valid species, two having been reduced to synonyms by Gauld (1974). The nomenclatural changes made in the present work are summarized below. HETEROPELMA Wesmael, 1849 Schizoloma Wesmael, 1849 Tanypelma Townes, 1971 syn. n. calcator Wesmael, 1849 fulvitarse Cameron, 1899 veticulatum Cameron, 1899 syn. n. binghami Cameron, 1907. syn. n. gvossator Shestakov, 1923 syn. n. pervlongum Cushman, 1937 syn. n. panargis Heinrich, 1953 syn. n. elongatum Uchida, 1928 flaviscutellum Uchida, 1928 tavsale Cushman, 1937 Syn. n. perornatum (Cameron), 1902 ocypeta sp. n. celeno sp. n. aello sp. n. scaposum (Morley), 19134 atvichiosoma Morley, 1913a flavitarse (Brullé), 1846 tvichiosomum Cameron, 1906 perniciosum (Turner), 1919 quodi (Vachal), 1907 savatiense (Fullaway), 1940 townesi sp. n. TAXONOMY OF GENUS HETEROPELMA 157 datanae Riley, 1888 flavicorne Brullé, 1846 fulvicorne Townes, 1945 syn. n. nigricorne (Szépligeti), 1906 comb. n. amictum (Fabricius), 1775 capitatum Desvignes, 1856 syn. n. bucephalum Brauns, 1898 fulvicorne Cameron, 1899 syn. n. tinctipenne Cameron, 1899 syn. n. acheron Morley, 1913a@ syn. n. acheron var. scutellatum Morley, 1913b syn. n. sachalinensis Matsumura, 1918 crassicalx Enderlein, 1921 syn.n. coveanum Uchida, 1928 syn. n. amictum var. nigricoxalis Uchida, 1928 amictum var. intermedium Uchida, 1928 Nomen inquirendum orbitale (Morley), 1913 The holotype and only known specimen of this species is deposited at the Museo Civico di Storia Naturale, Genoa. It has not been possible to examine this specimen. From the original description it would seem that this species is a synonym of H. fulvitarse. MATERIALS EXAMINED AND METHODS USED The following abbreviations have been used for depositories containing material which has been examined. ANIC Australian National Insect Collection, Canberra, Australia. BMNH British Museum (Natural History), London, England. BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A. DEI Deutsches Entomologisches Institut, Eberswalde, D.D.R. EIHU Entomological Institute, Hokkaido University, Sapporo, Japan. HKT Townes Collection, Ann Arbor, Michigan, U.S.A. IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium. IZPAN Instytut Zoologiczny, Warsaw, Poland. MNMH Muséum National d’Histoire Naturelle, Paris, France. QM Queensland Museum, Fortitude Valley, Qld., Australia. RSM Royal Scottish Museum, Edinburgh, Scotland. ™ Természettudomanyi Mizeum, Budapest, Hungary. UM University Museum, Oxford, England. USNM Smithsonian Institution, National Museum of Natural History, Washington D.C., WS.A: WAM Western Australian Museum, Perth, W.A., Australia. ZI Zoological Institute, Leningrad, U.S.S.R. For each species discussed a standardized format has been adopted. Whenever possible descriptions are based on type-material examined. Notes on intraspecific variation are given in a separate paragraph. Recorded hosts and synopses of recorded distribution, together with the appropriate references, are listed in separate paragraphs. 158 I. D. GAULD Drawings were made using a Wild M5 binocular microscope with a camera lucida attachment, except for figures of the genitalia which were made using a Watson ‘Bactil’ microscope in conjunction with a squared eyepiece. The method of male genitalia preparation is particularly important as excessive maceration with caustic potash (KOH) causes undue distortion of the aedeagal membranes. The method described by Gauld (1976) for genitalia preparation was adhered to in the present study. A number of indices adopted in an earlier work have been used in the present study. These are: CI (cubital index of forewing) = length of Cu, between 1m-—cu and Cu,, length of Cu, ICI (intercubital index of forewing) = length of 2 + 37m length of M between 2 + 37m and 2m-cu MI (marginal index of forewing) = length of Fs length of Rs + 2r NI (nervellar index of hindwing) = length of Cu, between cu—a and M length of cu—a The measurement of wing veins follows the method outlined by Gauld (1976). The abbreviations IOD and OOD were used respectively for the inter-ocellar and orbito-ocellar distances. Measurements of lower face were made taking the height as the distance between the mid clypeal apex and a mid point on a level with the lower margins of the antennal sockets, and the width as the minimum distance between the internal orbits. Morphological nomenclature in this work follows that of Richards (1956) as inter- preted for the Anomaloninae by Gauld (1976). The naming of genitalia structures follows that proposed by Peck (1937). The naming of microsculpture follows the system proposed by Eady (1968) and the larval terminology follows that of Short (1959). HETEROPELMA Wesmael Heteropelma Wesmael, 1849: 120. Type-species: Hetervopelma calcatoy Wesmael, 1849 : 120, by monotypy. Schizoloma Wesmael, 1849 : 120. Type-species: Ichnewmon amictus Fabricius, 1775 : 341, by monotypy. Schizopoma Foerster, 1868 : 145, 220. [Unjustified emendation. ] Tanypelma Townes, 1971 :157. Type-species: Hetevopelma fulvicovne Townes, 1945 : 129 [= Heteropelma datanae Riley, 1888 : 177], by original designation. Syn. n. DESCRIPTION. Eyes without pubescence, inner margins from subparallel to convergent TAXONOMY OF GENUS HETEROPELMA 159 ventrally; occipital carina from close to posterior ocelli to widely separated from ocelli, rarely dorsally incomplete; frons with a weak to strong interantennal lamella. Antennae of moderate length, scape truncate, 1-3—-1-7 times as long as pedicel; fourth flagellar segment 1-5—2-0 times as long as wide. Clypeus truncate, to rounded, sometimes with a pair of lateral swellings and apical clypeal margin reflexed; mandible bidentate, lower tooth always the shorter; labial palpi 4-segmented, cardo basally lobate. Genal carina reaching base of mandible. Pronotum with a weak transverse furrow dorsally; posterior corner of pronotum partially occluding spiracular sclerite; lower anterior margin abruptly rounded without an apical con- cavity; notauli present, rarely absent. Mesopleuron dorsally usually rugose or coriaceous, lateroventrally punctate below a large speculum. Posterior transverse carina usually complete, rarely centrally absent. Fore coxae smooth. Mid tibia usually bicalcarate, or in a Neotropical species with a single spur. Male hind tarsi swollen, 2nd segment usually with an impressed area ventrally which bears flattened microtrichia, or if rarely without this area then sometimes with microtrichia still present on ventral tarsal surface; hind claws usually geniculate and basally lobate, rarely simply curved. Forewing with Rs weakly sinuate, 2 + 3vm proximal to 2m—cu; 1m-—cu and Cu,, basally separate. Cu, between cu—a and Im-—cu, 0:80-1:05 times as long as 1m-—cu. Hindwing with 12-18 hamuli on vein F,; distal abscissa of Cu, always present. Propodeum reticulate, spiracle 2:50-3:50 times as long as broad; propodeum occasionally with conspicuous lateral protuberances. Gaster elongate, basal segments slender. 0 genitalia. Valvula 3 a little longer than apical depth of gaster; apex of ovipositor strongly constricted, extreme apex decurved; conspicuous sclerotized nodus present (Text-fig. 55). Ovarian egg obovate, dorsally flattened, bearing an obvious mushroom-like protuberance (the sucker of Iwata, 1958) that is basally extended anteriorly to form a thickened ridge or pair of ridges on the outer surface of the egg (Text-figs 56-64). 6 genitalia. Single syntergum present; 9th abdominal sternite from almost quadrate to distinctly transverse; gonosquamae short, usually acute apically. Gonolacinia distally abruptly angled about go°, teeth short to vestigial; basal apodeme angled at about 20° to the axis of the gonolacinia, reaching at least 0-65 of length of basivolsellar strut. Distivolsella slender, basally less than 0-5 times as broad as long, distally more or less swollen; teeth generally arranged diagonally on clasping face often with cluster on upper inner face, sometimes with smaller teeth distributed all over inner face. Aedeagal paramere proximally rounded, weakly spatulate or acute. Aedeagus in profile more or less sinuate, usually apically bilobate with obvious lateral sclerotized region; ventral membranous area bearing spines in specific patterns. IMMATURE STAGES. Only the first instar larva of one species, H. calcator, has been described (Plotnikov, 1914). The characteristic features of this larva are the elongate caudal appendage and presence of a small mouth with minute mandibles. The second instar larva resembles the first but has a much shorter caudal appendage. The final instar larvae of four species, H. amictum, H. calcator, H. scaposum and H. datanae, have been examined. The cephalic capsule of species of this genus is characterized by the broad pleurostoma and epistoma, short hypostoma and by having the labral sensilla arranged in two separate clusters. Most species were found to have the lateral ends of the hypostoma decurved, unlike species of Therion which have the hypostoma laterally straight. However, one species, H. scaposum, was observed to have the hypostoma laterally straight (Text-fig. 37). H. amictum and H. datanae were observed to have the ventral part of the labial sclerite lightly sclerotized. In the larva of H. scaposwm this sclerite is medio-ventrally extremely weakly defined. H. calcatoy was observed to be more variable. Some specimens have been found with the labial sclerite medio-ventrally sclerotized whilst Short 160 I. D. GAULD (1970) observed that H. calcator has the ventral part of the labial sclerite incomplete and not sclerotized (Text-fig. 36). DISCUSSION OF SOME CHARACTERS. In previous work (Gauld, 1976) it was stated that Heteropelma species invariably have the inner margins cf the orbits convergent ventrally and have the posterior transverse carina of the mesosternum complete. Exceptions have now been found to both these generalizations. A Papuan species, described below, was found to have the posterior transverse carina of the mesosternum centrally incomplete and H. nigricorne was observed to have the lower face sub- quadrate with the inner orbits extremely weakly convergent ventrally. This recent advance in the state of knowledge of this group makes it desirable to modify couplet 7 in the key to genera included by Gauld (1976). The amended couplet should read: 7 (6) Posterior transverse carina of mesosternum interrupted before each mid coxa; propodeum simple without lateral protuberances; lower face at narrowest point at least 0:85 times as broad as high; hind tarsal claws simply curved; ¢ with proximal apodeme of gonolacinia angled at 40° to gonolaciniar axis and less than 0.3 times as long as basivolsellar strut. Holarctic, extending into the Neotropics as far as about 20°N and into the Oriental region as far as New Guinea : : : : : : 3 : . THERION Curtis - Posterior transverse carina of mesosternum complete, or if rarely incomplete then propodeum with conspicuous lateral protuberances; lower face less than o-g times as broad as high; hind tarsal claws often geniculate; gj with proximal apodeme of gonolacinia angled at about 20° to gonolaciniar axis and more than 0-6 times as long as basivolsellar strut. All regions except Ethiopian HETEROPELMA Wesmael Couplet 8 is made redundant as, for reasons given below, Tanypelma has now been included as a synonym of Heteropelma. In the work cited above, Tanypelma was included as a separate genus, but it was stated that this genus appeared to have close affinities with Heteropelma, and that its status might need reconsideration. Recently a number of additional species have been studied and as a result the status of Tanypelma has been re-examined. The differences previously used for separating the two genera are tabulated below. HETEROPELMA TANYPELMA Scutellum in profile flat (Text-fig. 24). Scutellum in profile convex (Text-fig. 26). NI less than 0-60. NI greater than 0-80. 2nd hind tarsal segment of ¢ usually with 2nd hind tarsal segment of ¢ without an an impressed area ventrally. impressed area ventrally. Lower face elongate. Lower face subquadrate. Most species of Heteropelma were observed to have the scutellum flattened and more or less longitudinally concave. Species referred to Tanypelma were observed to have the scutellum more or less strongly convex. However, two species, H. quodt and H. savaiiense, were found to have a weakly swollen scutellum intermediate between that of typical Heteropelma and Tanypelma (Text-fig. 25). The differences in the range of variation of NI previously considered useful for effecting generic separation have been invalidated by the examination of additional TAXONOMY OF GENUS HETEROPELMA 161 material. A Brazilian species, H. townesi, resembles Heteropelma in most features (i.e. has the 3 hind tarsi impressed, scutellum flat, etc.) but has NI between 0-70 and 0-90 thus overlapping the range of Tanypelma. H. savaviense was observed to have NI equal too-70. The range of values of NI for Heteropelma must now be accepted to be 0:20-0:90 and it is therefore no longer possible to use this character to separate Heteropelma from Tanypelma. The majority of species of Heteropelma have the 2nd hind tarsal segment of the male ventrally impressed. Two species of Heteropelma do not have such an impres- sion present and thus resemble Tanypelma. Tanypelma species have the lower face subquadrate whereas most species of Heteropelma have the lower face elongate. However, H. nigricorne has been found to have the lower face subquadrate. Bearing in mind the combination of characters exhibited by Tanypelma datanae (the only species referable to the genus), it is apparent that this species differs some- what from the majority of species of Heteropelma. Five other species differ as much, or more, from the majority of Heteropelma species as does T. datanae (Chart 2), but it would not be concomitant with the classificatory criteria adopted for the Therionini as a whole to accord each of these species separate generic status. One may conclude that, because of the presence of intermediate species, and the existence of a much greater range of morphological variation than was presupposed to occur within the genus Heteropelma, the status of Tanypelma as a distinct genus can no longer be considered to be justifiable. It is suggested therefore that it should be included as a synonym of Heteropelma as formally stated above. In earlier works (that is all pre-1970 literature) Heteropelma and Schizoloma were treated as distinct genera on account of differences in the form of the clypeus and ratio of hind basitarsus to 2nd tarsal segment. Numerous intermediate forms between the two extremes of clypeal shape have been found (this is discussed further under the variation occurring in H. amictum). There is a considerable range of variation in the values of the hind tarsal ratios between species. As the ranges of several species overlap, the apparent difference between Heteropelma and Schizoloma is invalidated. The author therefore concurs with the opinion of Townes (1971) who placed Schizoloma as a synonym of Heteropelma. The characteristic form of the ovarian eggs of species of this genus was described by Iwata (1958). It has been possible to examine the ovarian eggs of six species, H. fulvitarse, H. calcator, H. perornatum, H. amictum, H. celeno and H. scaposum (Text-figs 56-64). The characteristic pedicel, called a sucker by Iwata, cannot in fact function as such, as the end is clearly distally convex (Text-fig. 56). The function of the pedicel is apparently to secure the egg to a suitable position within the host (Tothill, 1922). It is suggested that, rather than acting as a sucker, the membranous flange surrounding the distal apex of the pedicel functions as a barb preventing the pedicel from becoming dislodged from the tissue in which it had been embedded. Specific differences were found in the form of the pedicel and its basal insertion. In the majority of species the pedicel was positioned postero-dorsally but in H. scaposum a particularly conspicuous pedicel was positioned medio-dorsally. The 162 I. D. GAULD exact form of the anterior constricted region of the egg was observed to be variable. In immature eggs this region was extended into an elongate spout, whilst in more mature eggs the size of this spout was considerably reduced. The eggs of H. scaposum were not observed to have a spout. SYSTEMATIC POSITION OF THE GENUS. Heteropelma is apparently related to Habronyx (Habronyx), Therion and less closely to Trichomma. All these genera share a number of features in common including specialized eggs, a slender distally swollen distivolsella and broad larval pleurostomae. The latter two genera have like Heteropelma the gonolacinial apodeme formed from a sclerotized portion of the outer margin of the gonolacinia. Gauld (1976) included a full discussion of the possible evolutionary significance of these and other characters but was unable to satisfactorily relate Heteropelma to Therion on account of marked differences in the form of the aedeagus. A Papuan species, H. celeno, was observed to have an aedeagus intermediate in structure between those of typical Heteropelma and Therion. A second and closely related Papuan species, H. aello, was observed to have an incom- plete posterior transverse mesosternal carina, a characteristic of Therion rather than Heteropelma. However, in all other features these species were clearly referable to Heteropelma. It is now considered possible that Heteropelma may have evolved from a common ancestor with Therion, and that these two Papuan species represent the closest known species to this hypothetical ancestor. KEY TO SPECIES OF HETEROPELMA 1 Propodeum with a pair of lateral thorn-like projections (Text-figs 33, 34); pronotum without a well developed hook on lower anterior margin (Text-fig. 29) ; gaster black, thorax predominantly brick red. Papuan species : 2 — Propodeum without thorn-like ere prejeenions: peo with eae well developed hook on lower anterior margin (Text-figs 27, 28); gaster and thorax variously coloured, the latter usually darker than the former . 3 2 Face with elongate black pubescence; lateral projection of Seoedeaal very es (Text-fig. 34); posterior transverse carina of mesosternum centrally absent aello sp. n. (p. 176) — Face with short pale pubescence; lateral projection of propodeum blunt (Text-fig. 33); posterior transverse carina of mesosternum complete . celeno sp. n. (p. 175) 3 Forewing with abscissa of M between 2 + 3vm and 2m-—cu more than 3:0 times as long as 2 + 3vm (Text-fig. 39); an ey brick red species. Samoan Islands : ; savaiiense (Fullaway) (p. 183) — Forewing with abscissa of WM pecneen 2+ 3m eae 2m—cu less than 2-5 times as long as 2 + 3vm (Text-fig. 40) ; insect blackish marked on thorax or hind legs c 4 4 Marginal cell elongate, Rs more than 2:0 times as long as Rs + 2y (Text-fig. 40) ; hie trochanter equal to or longer than trochantellus ventrally; Cu, between Im—cu and Cu,, less than 0-6 times as one as Cu,,; mid tibia unicalcarate. Neotropical species . ‘ ‘ townesi sp. n. (p. 184) — Marginal cell shorter, Rs less than 1-9 ees as Meas” as Rs + 2v; hind trochanter ventrally shorter than trochantellus; Cu, between 1m-—cu and City, more than 0-6 times as long as Cu,,; mid tarsi bicalcarate, rarely with inner spur very reduced . 5 Io qT 12 TAXONOMY OF GENUS HETEROPELMA 163 Scutellum in profile swollen (Text-fig. 26); hind wing with NI greater than 0-70; male without either an impressed area or flattened microtrichia on 2nd hind tarsal segment. Nearctic species datanae Riley (p. 185) Scutellum in profile flat (ieee Fey ae 25); naa ae eal with NI less than 0-65; male with either an impressed area or flattened microtrichia or both on ventral surface of 2nd hind tarsal segment i 6 Hind basitarsus_ bicoloured, eagnieigs black, distally yellowish: Australasian species : ej Hind basitarsus mmicolorons ollon oan earely with migiseoset infuscation at exieme proximal end; Palaearctic or Oriental species . : 10 Hind legs except tarsi almost entirely black; wings ice ietamactes tepulae black: New Caledonia . : : quodi (Vachal) (p. 182) Hind legs with at least femora broadly md marked; wings hyaline or very weakly infumate; tegulae often yellow; Australian species . 8 Hind tarsal claws weakly curved, without a basal lamella (Text: fig. 41): lower fee usually black marked, always with elongate black pubescence; hind coxa ventrally with dense elongate pubescence; hind ocelli separated from occipital carina by a distance about equal to an ocellar diameter and flagellum with 60+ segments. Large species, forewing length 12-16 mm; Tasmania and southern Australia flavitarse (Brullé) (p. 180) Hind tarsal claws geniculate with a strong basal lobe or lamella (Text-fig. 30); lower face usually entirely yellowish with pallid pubescence; hind coxa ventrally with short sparse pubescence; hind ocelli separated from occipital carina by a distance greater than 1-2 times the ocellar diameter or ont with less than 2 ee or both : k 9 Thorax mainly black; dorsum of tergite 3 0-90 tienes: or more as ‘Tong ; as tergite 4; common species . : scaposum (Morley) (p. 177) Thorax almost entirely dash Grown, aay black mented dorsally and ventrally; dorsum of tergite 3 0-80 times or less as long as tergite 4; rare species perniciosum (Turner) (p. 181) Hind basitarsus more than 3-00 times as long as 2nd tarsal segment; clypeus be rounded, in profile not swollen ; II Hind basitarsus less than 2:80 times as long as ond tarsal segment: clypeus with a weakly to very strongly produced pair of lateral swellings, in profile convex or apically reflexed (Text-figs 16—18) : : , : ; : ; : 16 Mid tibia with spurs extremely inequal, the inner spur vestigial, less than 0-25 times as long as the outer (Text-fig. 35); occipital carina separated from posterior ocelli by more than 1-30 times ocellar diameter (Text-fig. 10). Thorax highly polished; Malaysian species . : ; ocypeta sp. n. (p. 174) Mid tibia with subequal to inequal spurs, the inner spur at the very least 0-45 times as long as the outer; occipital carina separated from posterior ocelli by less than 1:20 times ocellar diameter (Text-figs 1-6) ; ‘ : ‘ . : : 12 Mid tibial spurs inequal, the longer about 2-0 times as long as the shorter and the interantennal lamella vestigial (Text-fig. 23); scutellum yellow; flagellum with about 45 segments; male with hind basitarsus and 2nd tarsal segment confluent in outline, the latter without an impressed ventral area (Text-fig. 38) Japan, Taiwan. : 5 ? é : . flaviscutellum Uchida (p. 170) Mid tibial spurs generally subequal, the longer less than 1-5 times as long as the shorter or if very rarely with the longer 1-6—1-8 times as long as the shorter then the interantennal lamella is strongly developed (Text-fig. 20); interantennal lamella moderately to strongly developed (Text-figs 20-22); scutellum usually black; flagellum often with more than 46 segments; male with 2nd hind tarsal 164 13 14 15 16 Heteropelma calcatoy Wesmael, 1849 : 120. Syntypes, BELGium (IRSNB) [not examined]. i DE GAWD segment in outline discrete from basitarsus, and often with a ventral impressed area on the former (Text-fig. 41) : : ; ; : : Head posteriorly very strongly ened (Text-fig. 6); occipital carina medio- dorsally flattened with a more or less distinct median dip; flagellum with about 55 segments; pronotum with postero-dorsal half smooth with scattered punctures (Text-fig. 27). Borneo . . perornatum (Cameron) (p. Head less strongly waerouee Giese fee I- 25 eeciatell carina medio-dorsally usually convex, rarely slightly flattened but then with flagellum with 60+ segments; pronotum with postero-dorsal half coriaceous : : Head when viewed dorsally with genae broad and el, coeearicced (Text-figs 3, 4); flagellum with 48 or less segments; hind coxa usually entirely black, rarely in some east European specimens with reddish marks. Palaearctic; coniferous woodlands : A : calcator Wesmael (p. Head when viewed dorsally with genae not exactly as above, somewhat more posteriorly constricted (Text-figs 1, 2, 5); flagellum with 50+ segments; hind coxa entirely red or with extreme proximal margin black. . F : Hind basitarsus more than 4:8 times as long as 2nd tarsus; male without an impressed area on 2nd hind tarsal segment; propodeum dorsally about as long as broad; interantennal lamella triangular in profile (Text-fig. 22); flagellum with 60+ segments; eastern Palaearctic : : . elongatum Uchida (p. Hind basitarsus less than 4-50 times (usually a5 than 4:30 times) as long as 2nd tarsus; male with an impressed area on 2nd hind tarsal segment; propodeum ye broader than long; interantennal lamella trapezoidal in profile (Text-fig. 20); Oriental and Palaearctic : : fulvitarse Cameron (p. Lower face subquadrate, about 0-80 eae as Seed as high (Text-fig. 14); IOD 0-6 times as long as OOD; vertex immaculately black; flagellum blackish; clypeus with weak lateral Ei a S. India . : nigricorne (Szépligeti) (p. Lower face elongate, less a 0°70 fee as ieee as high (Text-fig. 16); IOD 0-65 or more times as long as OOD; vertex with yellow marks . amictum (Fabricius) (p. DESCRIPTIONS OF SPECIES Heteropelma calcator Wesmael (Text-figs 3, 4, 19, 36, 48, 59, 63) Hetevopelma calcatoyv Wesmael; Morley, 19134 : 69. Heteropelma calcatoy Wesmael; Plotnikov, 1914 : 34. Heteropelma calcatoy Wesmael; Morley, 1915 : 231. Hetevopelma calcatov Wesmael; Uchida, 1928 : 237. Heteropelma calcatoy Wesmael; Meyer, 1935 : 60. Heteropelma calcatoy Wesmael; Schmiedeknecht, 1935 : 62. Heteropelma calcatoy Wesmael; Uchida, 1958 : 94. Heteropelma calcatov Wesmael; Iwata, 1958 : 70. Hetevopelma calcatoy Wesmael; Townes, Momoi & Townes, 1965 : 382. Heteropelma calcatov Wesmael; Constantineanu & Pet¢u, 1969 : 93. Heteropelma calcatoyv Wesmael; Gauld, 1973 : 304. a) 173) 14 164) 5 169) 167) 187) 188) The type-material of this species was being studied by Dr Schnee (August, 1975) who was preparing a work on European Anomaloninae. TAXONOMY OF GENUS HETEROPELMA 165 Description. Lower face 0:60-0:70 times as broad as high; clypeus apically simply convex, in profile almost flat; mandibles twisted about 15°, the upper tooth about 2-00 times as long as lower; IOD 1:10-1:40 times as long as OOD; posterior ocelli separated from occipital carina by about maximum ocellar diameter. Head rather weakly constricted posteriorly, vertex punctate to coriaceous; occipital carina medio-dorsally usually evenly rounded, sometimes flattened. Flagellum of moderate length, 43- to 48-segmented; ist flagellar segment 2-0-2°3 times as long as 2nd; interantennal lamella strong, in profile almost square. Face with long, yellowish brown pubescence. Pronotum coriaceous, anterior hook strong. Mesoscutum coriaceous, laterally punctate; notauli strong, reaching beyond centre of mesoscutum; mesoscutal pubescence long, yellowish brown in colour. Mesopleuron dorsally rugose, lateroventrally punctate; epicnemial carina reaching above centre of mesopleuron, bowed so that upper end reaches anterior pleural margin; sternaulus weak. Scutellum usually longitudinally concave, slightly elongate, coriaceous. Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral projections, ventrally sculptured; propodeum dorsally 0-70—0-80 times as long as broad. Mid tibial spurs subequal, the outer about 1-10 times as long as inner. Hind leg with basitarsus 0:50-0:60 times as long as tibia, 3-40—3-80 times as long as 2nd tarsal segment; 2nd tarsal segment 1-go—2-20 times as long as 3rd; trochanter 0-80—0-g0 times as long as trochantellus ventrally; hind tarsal claw geniculate, basally strongly lobate. 4 with 2nd hind tarsal segment with an impressed ventral area. Wings very weakly infumate; CI = 0:55-0:80; MI = 1-75~-1-90; ICI = o-g0-1-40; NI = 0°45-0'50. Forewing length 8-11 mm. Gaster with tergite 2 2:20-2-40 times as long as tergite 3; 9 with ovipositor about 0-60 times as long as tergite 2. 6 genitalia. Distivolsella with spines arranged in apical cluster and with a short row of double spines on ventral region. Aedeagus in profile apically subequally bilobate; lateral sclerotized region apically dilated, indistinctly delineated. Head black, lower face and orbital marks yellow; flagellum reddish, proximally black. Thorax black, tegulae reddish. Legs reddish brown, coxae usually black, rarely reddish tinged or occasionally entirely red distally; distal 0-3 of hind tibia black, hind tarsi yellow. Gaster reddish, black marked on dorsal margin of tergite 2 and with terminal segments infuscate. VARIATION. This species exhibits less variation than the related species H. fulvitarse although it extends over a large range. Specimens from eastern Europe occasionally have the hind coxae distally red but all such specimens were observed to have the coxae black at extreme proximal ends. Remarks. This is one of the commonest Palaearctic species of Anomaloninae, about which a very considerable body of literature has accumulated. An excellent account of the biology and descriptions of the larval stages were included by Plotnikov (1914). The ovarian egg was described by Iwata (1958). This species is a well known parasite of Bupalus piniaria (L.) and because this moth is an important pest of conifer plantations a number of workers have studied the host-parasite interrela- tionship (Ljungdahl, 1916; Ritzema, 1920; Seitner, 1922; Sitowski, 1922; Barbey, 1927; Gornitz, 1931; Steiner, 1931; Friedrichs, Schaerffenberg & Sturm, 1941; Hedgqvist, 1949 and Malysheva, 1960; 1962). The following brief account of adult behaviour is based on personal observation of two populations, one at Cannock Chase, Staffordshire, England and a second at Craigen Darroch, Grampian, Scotland. H. calcator is generally associated with Pinus woodlands. It seems to prefer 166 tl. D. GAULD areas in which there are many young trees (2-3 m high) or in which taller trees (4-5 m) are widely spaced. Males appear in mid July and specimens of both sexes are most common in mid August. In flight both sexes adopt a very characteristic attitude, flying with antennae outstretched, hind legs fully extended at about 30° to the axis of the body and with the slender gaster elevated at about 15°. A considerable amount of the flight path is in the vertical plane, close to the conifer foliage. At night and during periods of inclement weather both sexes rest on conifer twigs. At rest they elevate the gaster at an angle approximately the same as that subtended by the leaves to the twigs. Thus the insect renders itself very inconspicuous. Males adopt a characteristically rapid flight in clearings, rising vertically and sinking, with the conspicuous hind tarsi outstretched. Two or three males behaving so together have been seen, but no evidence for aggregate male displays exists. Whether this behaviour is some form of pre-copulatory sexual display is not known. The female searches for prospective hosts on the wing, and alights only for the briefest moments to preen the antennae and wings. Having located a suitable host (usually the early instar larvae of B. piniaria) the female alights, reflexes the ovipositor forward and then oviposits, after which she resumes flight. The reflexing of the ovipositor serves to move an egg from the ovipositor base to an expanded chamber immediately proximal to the nodus. During this reflexing the 1st valvulae are retracted relative to the 2nd valvulae, and the valvilli (a pair of minute flanges of tissue positioned on the distal internal faces of the Ist valvulae, vide Gauld, 1976) engage the egg. As the Ist valvulae slide along the 2nd to regain their resting position the egg is forced down the lacuna of the ovipositor by the valvilli. Thus when the ovipositor is inserted into the lepidopterous larva an egg is already positioned near the apex. Observation of acts of oviposition were too infrequent to permit any reliable inferences being made about the position of insertion of the ovipositor into the host. H. calcator is undoubtedly closely related to the Oriental species H. fulvitarse which apparently occupies a similar ecological niche in the Oriental region. The two species are morphologically rather similar but it seems possible to separate the two by differences in head shape (Text-figs 1-4) and usually by differences in coxal coloration. Whether or not these differences will prove to be reliable when the fauna of western and central Asia is better known is a matter of great interest. Host RECORDS. Other than B. piniaria which is referred to above, this species has been recorded as a parasite of Panolis flammea (Denis & Schiffermiiller) (Meyer, 1935; Schmiedeknecht, 1935), Anarta myrtilli (L.) (Schmiedeknecht, 1935), Bena prasinana (L.) (Morley, 1915; Meyer, 1935). DISTRIBUTION (Map 1). This species has been recorded from most European countries (Schmiedeknecht, 1935), but is perhaps absent in the south as Ceballos (1963) failed to find it in Spain. Meyer (1935) recorded this species from many localities in the U.S.S.R. Uchida (1958) recorded this species from eastern U.S.S.R., China, Korea and Japan. TAXONOMY OF GENUS HETEROPELMA 167 MATERIAL EXAMINED. BELGIuM: I 9, no further data (BMNH). GERMANY: 49, 24, no further data (BMNH). Great Britain: 11 9, 14 3, Devon, Lustleigh, vii. 1934 (R. C. L. Perkins) (BMNH); ro 9, 10 g, Grampian ‘Aberdeenshire’, Ballater, Craigen Darroch, vii. 1975 (Gauld) (BMNH); 5 9, 7 3, Hampshire, New Forest, vii—vili. 1927-1936 (Morley) (BMNH); 40 9, 48 g, Staffordshire, Cannock Chase, bred ex B. piniaria on Pinus sylvestris L., vi. 1954 (Forestry Commission) (BMNH); Io 9, 10 g, same locality, on Pinus sp., viii. 1972-1973 (Gauld) (BMNH). Norway: 3 9, 3 4, Brekkesto, viii. 1938 (Chaworth-Musters) (BMNH). Rumania: 2 9, I g, no further data. SWEDEN: I Q, Fijellfota sj6, vii. 1938 (J. F. Perkins) (BMNH). Turkey: I 9, Unye, vii. 1972, no further data (BMNH). Heteropelma fulvitarse Cameron (Text-figs I, 2, 20, 28, 41, 52, 55, 62) Heteropelma fulvitarse Cameron, 1899 : 111. Holotype 9, INp1a (UM) [examined]. Heteropelma reticulatum Cameron, 1899 : 110. Holotype g, Inp1a (UM) [examined]. Syn. n. Anomalon binghami Cameron, 1907: 14. Holotype g, INp1A: Sikkim (BMNH) [examined]. Syn. n. Heteropelma fulvitayse Cameron; Morley, 19134 : 70. [Exochilum acheron Morley, 1913a:79. Partim, 1 g¢ from Shanghai. Misidentification.] Anomalon binghami Cameron; Morley, 19134 : 84. [Heteropelma calcatoy Wesmael; Morley, 1913b : 406. Misidentification. | Heteropelma fulvitarsis Cameron; Morley, 1913b : 407. Anomalon binghami Cameron; Morley, 19130 : 424. Anomalon? binghami Cameron; Morley, 1914 : 325. Heteropelma grossatoy Shestakov, 1923 : 46. Holotype 9, U.S.S.R. (ZI) [examined]. Syn. n. [Heteropelma calcatoy Wesmael; Dutt, 1923 : 18. Misidentification. } Heteropelma grossatoy Shestakov; Meyer, 1935 : 60. Heteropelma perlongum Cushman, 1937 : 297. Holotype 9, Taiwan (DEI) [examined]. Syn. n. Heteropelma panargis Heinrich, 1953:181. Holotype g, GERMANY (WEstT) (HKT) [examined]. Syn. n. Heteropelma perlongum Cushman; Uchida, 1958 : 94. Heteropelma binghami (Cameron) Townes é al., 1961 : 321. Heteropelma fulvitarse Cameron; Townes ét al., 1961 : 321. Heteropelma perlongum Cushman; Townes ef al., 1961 : 321. Heteropelma reticulatum Cameron; Townes ef al., 1961 : 322. Heteropelma grossatoy Shestakov; Townes et al., 1965 : 383. DeEscRIPTION. Lower face 0:60-0:70 times as broad as high; clypeus apically simply convex, in profile rather flat; mandibles twisted about 20°, the upper tooth 2-0-3-o0 times as long as lower. JOD o-go—1°30 times as long as OOD; posterior ocellus separated from occipital carina by less than the greatest ocellar diameter. Head constricted posteriorly, vertex closely punctate to coriaceous; occipital carina evenly rounded or rarely medio-dorsally concave. Flagellum of moderate length, 50- to 55-segmented; 1st flagellar segment 2:10-2:50 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with long pale pubescence. Pronotum coriaceous, anterior hook strong. _Mesoscutum rather coarsely punctate, centrally rugose; notauli moderately strongly to weakly impressed, discernible to centre of scutum; mesoscutal pubescence short, usually ruddy or badious, rarely yellowish red. Mesopleuron dorsally rugose or reticulate, latero-ventrally finely punctate; epicnemial carina usually reaching 168 i DIGAWLD to centre of mesopleuron, bowed and with upper end reaching anterior pleural margin; sternaulus weak. Scutellum longitudinally concave or flat, elongate, rugosely punctate. Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral protu- berances, ventrally strongly sculptured; propodeum dorsally 0-80—0-95 times as long as broad. Mid tibial spurs subequal, the outer about 1-1 times as long as inner. Hind leg with basitarsus 0°55-0°65 times as long as tibia, 3:60-4:20 times as long as 2nd tarsal segment; 2nd tarsal segment 1-70—2-60 times as long as 3rd segment; trochanter 0-75—0-95 times as long as tro- chantellus ventrally; hind tarsal claw geniculate, basally with a strong lamella. 2nd hind tarsal segment of ¢ ventrally impressed. Wings weakly infumate; CI = 0:55-0:85; MI = 1:60-1:90; ICI = 0:70-1:30; NI = 0-40- 0-50. Forewing length 1o-15 mm. Gaster with tergite 2 2-00—2-20 times as long as tergite 3; 2 with ovipositor 0:50-0:60 times as long as tergite 2. $ genitalia. Distivolsella with spines arranged in an apical cluster and with short spines in a double row on ventral margin of distivolsella. Aedeagus in profile apically obliquely truncate; lateral sclerotized area apically acute, ventrally extended (Text-fig. 52). Head black, face and genal marks yellow; antennae reddish or yellowish distally. Thorax black, tegulae reddish yellow. Anterior two pairs of legs yellowish, hind legs reddish, tibia distally darker, tarsi paler; gaster reddish. VARIATION. This species was observed to be morphologically rather variable. There was observed to be a marked clinal variation in the specimens examined, those from the west of the range being gencrally darker and stouter than those from the east. Intermediates between the extremes of variation have been observed. To some extent the island populations, particularly that of Taiwan, are separable from the mainland form. The Taiwanese specimens were found to have the flagella pale yellowish and the scape not at all black marked as is most usual in this species. There are, however, no consistent morphological differences between the island and the mainland Chinese populations. As already mentioned there is a tendency for the specimens in the east of the range to be paler and so it is considered that the Taiwan population is conspecific with the mainland population. A single 9 from Hainan differs from the mainland Chinese population in having the lower tooth of the mandible subequal to the upper. There is quite a considerable variation of the ratio of lengths of the mandibular teeth within any one population and although no other specimens have been found with such subequal teeth it was considered possible that the Hainan specimen may represent an extreme variant of H. fulvitarse. The holotypes of H. grossatoy Shestakov and H. panargis Heinrich were observed to have the head posteriorly strongly constricted and have a centrally depressed occipital carina. In all other features these species are similar to typical H. fulvi- tayse specimens and as head shape and the form of the occipital carina are very variable features H. grossator and H. panargis are included as synonyms of H. fulvitarse. Isolated specimens of H. fulvitarse were observed to have the mid tibial spurs rather inequal, the longer about 1-50 times the length of the shorter. These specimens differ from H. flaviscutellum in having the interantennal lamella strongly developed, the 2nd hind tarsal segment of the j with an impressed ventral area and having a black scutellum. LAN ONOMSY OFGENUS BETEROPELAIA 169 Remarks. This species apparently replaces H. calcator in the Oriental region. No biological information is at present available. Host rEcorps. None available. DISTRIBUTION (Map 1). This species has been recorded from northern India and Sikkim (Cameron, 1899; 1907; Morley, 1913b). The latter author also recorded this species from Burma. It has been recorded from U.S.S.R. (Shestakov, 1923; Meyer, 1935) and Taiwan (Cushman, 1937; Uchida, 1958). MATERIAL EXAMINED. Heteropelma fulvitarse Cameron, holotype 9, INDIA: Assam, Khasia Hills (UM). Heteropelma reticulatum Cameron, holotype 3, INDIA: Assam, Khasia Hills (UM). Anomalon binghami Cameron, holotype 3, India: Sikkim (BMNH). Heteropelma grossator Shestakov, holotype 9, U.S.S.R.: Tomsk, Yelizavetinskiy (ZI). Hetero- pelma perlongum Cushman, holotype 2, TAIwANn (DEI). Heteropelma panargis Hein- rich, holotype 3, GERMANY (WEsT): Steiermark (HKT); paratype, I 3, same data as holotype (HKT). Burma: I g, Maymayo, xii. 1937 (Heinrich) (HKT); 39,53, Mt Victoria, iv. 1938 (Heinrich) (HKT). Cuina: 6 9, Fukien, Shaowu Hsien, iv—v. 1942-1945 (Maa) (HKT); 1 9, Tibet, Yatung, vii. 1924 (Hingston) (BMNH); 1 9, no further data (BMNH). Inp1a: 2 9, Simla, vil. 1918 (Brunetti) (BMNH); 1 2, Assam, Mishmi Hills, xi. 1936 (BMNH); 2 J, no further data (BMNH); 39,4, no further data (UM). Laos: 1 9, Muong Pek, xii. 1918 (Salvaza) (BMNH). NEPAL: 1 9, 1 g, Taplejung, x-xi. 1961 (Coe) (BMNH). Taiwan: 1 Q, Taiheizan, vii. 1934 (Gressitt) (HKT); I 9, Musha, v. 1932 (Gressitt) (HKT); 1 3, Arizan, viii. 1908 (Waleman) (BMNH). Heteropelma elongatum Uchida (Text-figs 5, 22, 50) Hetevopelma calcator var. elongatum Uchida, 1928 : 238. Lectotype, JAPAN (EIHU), designated by Townes e¢ al., 1965 : 383 [examined]. Heteropelma elongatum Uchida; Uchida, 1953 : 127. Heteropelma elongatum Uchida; Uchida, 1958 : 94. Hetevopelma elongatum Uchida; Iwata, 1960 : 168. Heteropelma elongatum Uchida; Townes ef al., 1965 : 383. Description. Lower face 0-60—0-65 times as broad as high; clypeus apically simply convex, in profile rather flat; mandibles twisted about 15°, the upper tooth about 2-00 times as long as the lower. LOD o-go-1-10 times as long as OOD; posterior ocellus separated from occipital carina by less than maximum ocellar diameter. Head constricted posteriorly, vertex coriaceous, laterally grading to coarsely punctate; occipital carina evenly rounded. Flagellum elongate, 61- to 65-segmented; ist flagellar segment 2-20—2-30 times as long as 2nd; interantennal lamella of moderate size, in profile more or less triangular. Face with long pale pubescence. Pronotum coriaceous, anterior hook strong. Mesoscutum centrally coriaceous, laterally punctate; notauli moderately strong, reaching beyond centre of scutum; mesoscutal pubescence short, yellowish red. Mesopleuron dorsally reticulate, latero-ventrally punctate; epicnemial carina strong, reaching to about centre of mesopleuron, usually with upper end remote from anterior pleural margin; sternaulus indistinct. Scutellum longitudinally concave. Posterior 170 I. D. GAULD transverse carina of mesosternum complete. Propodeum reticulate, without lateral protu- berences, ventrally strongly sculptured; propodeum dorsally about 1-00 times as long as broad. Mid tibial spuis subequal, the outer about 1-20 times as long as inner. Hind leg with basitarsus 0:66-0:68 times as long as tibia, 4:90-5:90 times as long as 2nd tarsal segment; 2nd tarsal segment 1-70-1-80 times as long as 3rd tarsal segment; trochanter 0-83—-0-88 times as long as trochantellus ventrally; hind tarsal claw geniculate with a strong basal lamella. $ without an impressed area on 2nd hind tarsal segment. Wings weakly infumate; CI = 0:53-0:68; MI = 1-50-1-80; ICI = 0:60-1:20; NI = 0-50— 0°65. Forewing length 12-17 mm. Gaster with tergite 2 2-10-2-20 times as long as tergite 3; Q with ovipositor 0-45 times as long as tergite 2. 3 genitalia. Distivolsella with spines arranged in apical cluster and with short row of double spines extending down ventral margin. Aedeagus in profile apically subequally bilobate; lateral sclerotized area apically broadened, indistinctly delineated. Head black, lower face marks on genae yellow; antennae reddish. Thorax black, tegulae reddish yellow. Anterior two pairs of legs yellowish, hind legs reddish, tibia distally darker, trochanter ventrally pale. Gaster reddish. VARIATION. Some variation in the coloration was observed. The female from China was found to have the hind coxa somewhat more swollen than is typical for this species. Slight variation in the shape of the interantennal lamella was also observed, a few specimens tending to have the lamella subquadrate in profile. REMARKS. This species is clearly very closely related to H. fulvitarse from which it differs in having a longer flagellum (61-65 segments compared with 50-55), the hind basitarsus extremely elongate (4-g0-5-90 times as long as tarsus 2 compared with 3:60-4:20 times as long as tarsus 2) and having the 2nd hind tarsal segment of the 3 devoid of an impressed area. Host RECORDS. None available. DistTRIBUTION (Map 1). This species has previously been recorded from Japan (Uchida, 1928; 1958). MATERIAL EXAMINED. Heteropelma elongatum Uchida, lectotype g, JAPAN: Honshu, Tokyo, v. 1918 (Teranischt) (ETHU). JAPAN: I ¢ (Teranischit) (HKT). Curna: 1 9, Fukien, Shaowu Hsien, ix. 1943 (Maa) (HKT); 2 3, same locality, v. 1943 (Maa) (HKT); 1 g, Tsin Leong San, vi. 1936 (Kwantung) (HKT). Heteropelma flaviscutellum Uchida (Text-figs 23, 38) Heteropelma calcator var. flaviscutellum Uchida, 1928: 238. Lectotype g, JAPAN (EIHU), designated by Townes et al., 1965 : 383 [examined]. Hetervopelma tarsale Cushman, 1937 : 298. Holotype 9, TArwan (DEI) [examined]. Syn. n. Heteropelma flaviscutellum Uchida; Uchida, 1958 : 95. Hetevopelma tarsale Cushman; Uchida, 1958 : 95. Heteropelma tarsale Cushman; Townes e¢ al., 1961 : 322. Heteropelma flaviscutellum Uchida; Townes ef al., 1965 : 383. TAXONOMY OF GENUS HETEROPELMA som @ FULVITARSE 4 CALCATOR @ PERORNATUM tk OCYPETA © ELONGATUM 171 Map 1. Palaearctic and Oriental zoogeographic regions (Zenithal equal projection; origin 40°N, 95°E) showing distributions of Heteropelma fulvitarse, H. calcator, H. perornatum, H. ocypeta and H. elongatum. 172 iE Dy GAUED DESCRIPTION. Lower face 0-65—0-75 times as broad as high; clypeus apically simply convex, in profile swollen; mandibles twisted about 5°, upper tooth slightly less than 2-00 times as long as lower. JOD equal to OOD; posterior ocellus separated from occipital carina by about 0-90 times maximum ocellar diameter. Head posteriorly moderately constricted, vertex punctate; occipital carina medio-dorsally evenly rounded. Flagellum rather long and slender, 45-segmented, 1st flagellar segment about 1-80 times as long as 2nd; interantennal lamella almost absent. Face with long pallid pubescence. Pronotum coriaceous, anterior hook moderately developed. Mesoscutum coarsely punctate; notauli strong, reaching beyond centre of scutum; mesoscutal pubescence short and pale. Mesopleuron dorsally coriaceous, ventrally sparsely punctate; epicnemial carina reaching above centre of pleuron, bowed so that upper end reaches anterior margin of mesopleuron, sternaulus weak. Scutellum convex, quadrate, striate. Posterior transverse carina of meso- sternum complete. Propodeum reticulate, without lateral protuberance, ventrally strongly sculptured; propodeum dorsally 0-80 times as long as broad. Mid tibial spurs inequal, the outer about 2:00 times as long as the inner. Hind leg with basitarsus 0:62-0:65 times as long as tibia, 4:30-6:10 times as long as 2nd tarsal segment; 2nd tarsal segment 1:30-1:90 times as long as 3rd tarsal segment; trochanter 0:73-0:77 times as long as trochantellus; hind tarsal claws as fulvitavse. Hind tarsus of ¢ without an impressed area. Wings very weakly infumate; Cl = 0-72-0-87; MI = 1-73-1-81; ICI = 1:44-1:60; NI = 0:63-0:70. Forewing length 1o-11 mm. Gaster with tergite 2 about 1-80 times as long as tergite 3; 2 with ovipositor 0-50 times as long as tergite 2. 4 genitalia. Not available for dissection. Head black, lower face and vertical marks yellow; antennae reddish yellow. Thorax black, tegulae reddish, scutellum yellow. Anterior two pairs of legs yellow, hind legs brownish, coxae proximally darker, tibia distally dark brown, tarsi pale yellow. VARIATION. The colour of this species is extremely variable. In some females the thorax was observed to be almost entirely badious with more reddish areas at bases of notauli, on mesopleuron and margins of the varioussclerites. The scutellum, tegulae and sometimes subalar prominences were observed to be yellow. In the lighter specimens the antennae are yellowish, gaster except for the dorsum of tergite 2 orange-red, hind legs except for tarsi, distal 0-30 of tibia and dorsum of trochanter yellowish brown. Intermediates between this extreme and the typical form have been seen. REMARKS. The basitarsus and 2nd tarsal segment of males of this species are unusual in being particularly intimately associated and in outline continuous with each other. This feature enables males to be distinguished from all other species. Host REcoRDsS. None available. DIsTRIBUTION. Previously recorded from Japan and Taiwan (Uchida, 1928; 1958; Cushman, 1937). MATERIAL EXAMINED. Heteropelma flaviscutellum Uchida, lectotype gj, JAPAN: Hokkaido, Jozankeim, vill. 1924 (Uchida) (EIHU). Heteropelma tarsale Cushman, holotype 2, TAIWAN: Kankau, Koshun, v. 1912 (Sauter) (DEI); paratypes, 1 9, same data as holotype, I dg same locality and collector as holotype, vill. 1908 (DEI). TAIWAN: I 9, Kuraru, v. 1934 (Gressitt) (HKT). TAXONOMY OF GENUS HETEROPELMA eS Heteropelma perornatum (Cameron) (Text-figs 6, 21, 24, 27, 44, 58) Anomalon perovnatum Cameron, 1992 :50. LECTOTYPE 9, Borneo: Sarawak (BMNH), here designated [examined]. Anomalon perornatum Cameron; Szépligeti, 1905 : 12. Hetevopelma perovnatum (Cameron); Morley, 19134 : 70. Hetevopelma perornatum (Cameron); Townes e¢ al., 1961 : 321. Description. Lower face 0:60-0:65 times as broad as high; clypeus apically simply convex, in profile weakly swollen; mandibles twisted about 35°, the upper tooth twice as long as the lower. IOD 1-1o—1-40 times as long as OOD; posterior ocelli separated from occipital carina by about o-go times a maximum ocellar diameter. Head strongly constricted, vertex smooth with scattered punctures; occipital carina medio-dorsally usually dipped, rarely simply flattened. Flagellum of moderate length 53- to 55-segmented; 1st flagellar segment 1-g0—2-10 times as long as 2nd; interantennal lamella weak, in profile obtusely triangular. Face with short pale pubescence. Pronotum dorsally rather polished, smooth with obsolescent scattered punctures, ventrally strongly rugose, anterior hook strong. Mesoscutum punctate or rugulose; notauli weakly impressed but nevertheless obvious, extending beyond the centre of the scutum; mesoscutal pubescence short and pale. Mesopleuron dorsally strongly rugose, ventrally smooth with obsolescent punctures; epicnemial carina strong, reaching above centre of mesopleuron, bowed so that upper end reaches anterior pleural margin; sternaulus weak. Scutellum flat, elongate, rugosely striate. Posterior transverse carina of mesosternum very strongly raised, complete. Propodeum reticulate, without lateral protuberances, ventrally strongly sculptured; propodeum rather long, dorsally 0:90-1:10 times as long as broad. Mid tibial spurs subequal, the outer about 1-10 times as long as the inner. Hind leg with basitarsus 0:65-0:70 times as long as tibia, 5:00-5:50 times as long as 2nd tarsal segment; 2nd tarsal segment 1:70-2:00 times as long as 3rd; trochanter 0-80—-0-go0 times as long as tro- chantellus ventrally; hind tarsal claw geniculate, strongly lobate. Hind tarsus 2 of male ventrally impressed on proximal 0-50 only. Wings weakly infumate, CI = 0:75-0:95; MI = 1:65-1:75; ICI = 0:80-1:40; NI = 0:40-0:60. Forewing length 12-15 mm. Gaster with tergite 2 2:00-2:20 times as long as tergite 3; 2 with ovipositor 0:50-0:60 times as long as tergite 2. 6 genitalia. Distivolsella with spines arranged in apical cluster and with short row of paired spines on ventral margin. Aedeagus in profile equally bilobate; lateral sclerotized area apically truncate, lower corner ventrally extended. Head black, lower face and orbits yellow; flagellum black, rarely distally paler. Thorax black, tegulae reddish. Anterior two pairs of legs dark brown, tibia centrally and tarsi pale yellow. Gaster reddish brown. VARIATION. Some variation in colour, especially that of the extent of the yellow on the hind tibia, has been observed. REMARKS. This species is morphologically closely related to H. fulvitarse which presumably it replaces in Borneo. Despite the close relationship the differences in pronotal sculpture and head shape will allow specific separation. Host rEcorbDs. None available. DISTRIBUTION (Map 1). Recorded only from Borneo (Cameron, 1902; Szépligeti, 1905). 174 LD eae MATERIAL EXAMINED. Anomalon perornatum Cameron, lectotype 2, BORNEO: Sarawak, Kuchung, vii. 1899 (BMNH); paralectotype 1 2, BORNEO: xi. 1898 (Shelford) (BMNH). Borneo: I 9, Sarawak, Kuchung, vi. 1902 (BMNH); 3 9, 1 g, ‘Borneo’ x. 1850 (Hewitt) (RSM); 1 3, Kalimantan, Midden, viii. 1925 (Szebers) (HKT). Heteropelma ocypeta sp. n. (Text-figs 10, 35) |Hetevopelma perornatum (Cameron); Gauld, 1976: 28. Misidentification. ] This and the following two new species, H. celeno and H. aello, are named after the Harpyai of classical mythology, in whom are embodied the concepts of wind spirits and predatory ghosts. DESCRIPTION. Lower face 0:60-0:65 times as broad as high; clypeus apically simply convex, in profile flattened; mandibles twisted about 35°, the upper tooth about 2-50 times as long as the lower. IOD about 1-30 times OOD; posterior ocellus separated from occipital carina by 1:30-1:40 times maximum ocellar diameter. Head strongly constricted posteriorly, vertex coriaceous; occipital carina evenly rounded dorsally. Flagellum of moderate length, 49- segmented; rst flagellar segment 2:10-2:40 times as long as 2nd; interantennal lamella weak, in profile obtusely triangular. Face with long badious or rufescent pubescence. Pronotum dorsally punctate, lower 0-40 longitudinally striate, anterior hook strong. Meso- scutum centrally coriaceous, laterally punctate; notauli weakly impressed but nevertheless obvious, extending beyond centre of mesoscutum; mesoscutal pubescence long, badious. Mesopleuron dorsally coriaceous, latero-ventrally sparsely punctate; epicnemial carina reaching above centre of mesopleuron, bowed so that upper end reaches anterior pleural margin; sternaulus weak. Scutellum flat, subquadrate, punctate. Posterior transverse carina of mesosternum strong, complete. Propodeum reticulate, without lateral protuberances, ventrally strongly sculptured; propodeum dorsally 0-85—0-95 times as long as broad. Mid tibial spurs extremely inequal, the outer of normal length, the inner vestigial. Hind leg with basitarsus about 0-60 times as long as tibia, 4:10-5:10 times as long as 2nd tarsal segment; 2nd tarsal segment 1:40-1:70 times as long as 3rd; trochanter 0-g0—0-95 times as long as trochantellus ventrally; hind tarsal claw geniculate, basally strongly lobate. ¢ unknown. Wings weakly infumate, CI = 0:95-1:00; MI = 1:63-1:69; ICI = 0:80-1:00; NI = 0:40-0:60. Forewing length 9-10 mm. Gaster with tergite 2 2-40 times as long as tergite 3; 2 with ovipositor 0-45-0'55 times as long as tergite 2. Head black, lower face and orbits yellow; flagellum reddish, proximally black, distally yellow. Anterior two pairs of legs yellow; hind legs brownish yellow, coxae and distal 0-40 of hind tibia black. Gaster reddish yellow, indistinctly infuscate in irregularly defined areas. VARIATION. The paratype was observed to be somewhat more hirsute than the holotype. Possibly some of the hair on this specimen has been removed in the past by cleaning. REMARKS. This species is similar in general morphology to H. perornatum but differs in having inequal mid tibial spurs, widely interspaced occipital carina and posterior ocelli, and having a more polished thorax. Host REcorRpDS. None available. TAXONOMY OF GENUS HETEROPELMA 175 DISTRIBUTION (Map 1). Malaysia. MATERIAL EXAMINED. Holotype 9, MataysiaA: Pahang, Cameron Highlands, 1700 m, xil. 1939 (Pendle- bury) (BMNH). Paratype. MALAysIA: I 9, Pahang, Cameron Highlands, 1600m, vi. 1935 (Pendlebury) (BMNH). Heteropelma celeno sp. n. (Text-figs II, 29, 32, 33, 47, 56, 57) DESCRIPTION. Lower face 0:68-0:78 times as broad as high; clypeus apically simply convex, in profile more or less flat; mandibles twisted about 20°, the upper tooth about 2-00 times as long as lower. JOD 0-70-0-g0 times as long as OOD; posterior ocellus separated from occipital carina by o-go times maximum ocellar diameter. Head strongly constricted, vertex closely but shallowly punctate; occipital carina dorsally flattened or slightly dipped. Flagellum of moderate length, 61- to 65-segmented; ist flagellar segment 2-50—2-80 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with short pale pubescence. Pronotum punctate to weakly coriaceous, anterior hook absent or vestigial. Mesoscutum coarsely and closely puncto-coriaceous, matt; notauli weak to indistinct ; mesoscutal pubescence short, whitish. Mesopleuron dorsally alutaceous, latero-ventrally with fine even puncturation; epicnemial carina indistinct above lower corner of pronotum; sternaulus weak. Scutellum longitudinally concave, subquadrate, punctate. Posterior transverse carina of mesosternum complete, centrally weak. Propodeum reticulate with a pair of blunt lateral protuberances, ventrally weakly sculptured; propodeum dorsally 0-70—0-83 times as long as broad. Mid tibial spurs subequal, the outer about 1-20 times as long as inner. Hind leg with basitarsus about 0-50 times as long as tibia, 2:40-2:80 times as long as 2nd tarsal segment; 2nd tarsal segment 1:70-2:10 times as long as 3rd; trochanter 0-70—0-90 times as long as tro- chantellus ventrally; hind tarsal claw abruptly curved, slender with a weak basal lobe. with 2nd hind tarsal segment strongly impressed ventrally. Wings infumate, CI = 0-68; MI = 1:70-1:80; ICI = 0:30-0:90; NI = 0:20-0:30. Gaster with tergite 2 about 2:50 times as long as tergite 3; 2 with ovipositor 0-43—0°55 times as long as tergite 2. 6 genitalia. Distivolsella with spines arranged in an apical cluster, some small spines distributed on clasping face. Aedeagus in profile apically very inequally bilobate; lateral sclerotized area apically simply acute. Head, gaster and hind legs except for extreme distal end of basitarsus, tarsi 2-5 and coxae, black; lower face, orbits, extreme distal end of hind basitarsus and hind tarsi 2-5 white. Thorax, propodeum, anterior two pairs of legs and hind coxae reddish brown. VARIATION. There was observed to be slight sexual dimorphism in this species. The males have the lower face immaculately white whereas that of the female has a pair of vertical black stripes extending from the anterior tentorial pits to the bases of the antennae. The hind tarsal claws of the males were observed to be slightly more slender than those of the females. REMARKS. This and the following species, H. aello, are immediately distinguish- able from other species of Heteropelma on account of their colour pattern. Unlike other species which are cryptically patterned these two species have bright warning coloration. This pattern is shared with three further synchronous and sympatric Papuan species, a species of Aphanistes, one of Tvichomma and one of Casinaria 176 is DEG AWD (all are believed to be undescribed). Whilst ichneumonoid mimics of Aculeata are not uncommon (Metopius species mimic Eumenid wasps and many species of Ichneumonidae mimic Pompilid wasps (Evans, 1968)), this mimicry complex apparently involves no aculeates. As H. celeno is the most numerous species in Malaise trap collections (and is thus arguably the commonest species) it is likely to be the model for the mimicry complex. It is probable therefore that this species is unpalatable to predators. It is difficult working with preserved material to ascertain why this species should be so unpalatable, but unlike most other species it was observed to have large slender claws, which were observed to be apically strongly curved and appeared to have a cavity in the distal part. A similar cavity has been observed by Townes (1940) in some Pimplinae, and he suggested that the claws may function as organs of defence. Townes suggested that the readiness with which the live insects grasped a net with their claws might indicate that it is their habit, when attacked, to embed their claws in a prospective predator. Because the cavity structurally weakens the claw, the distal portion of it would break off remaining embedded in the prospective predator. Simultaneously the contents of the cavity (possibly a toxin) would be liberated. Heteropelma celeno may well have evolved a similar defence mechanism. In the material examined a large number of specimens were observed to have the claws incomplete, indicating that claw fractures readily occur. This species is obviously closely related to H. aello but the relationship of these species to the remainder of the genus is not at all clear. Host RECORDS. None available. DISTRIBUTION. New Guinea. MATERIAL EXAMINED. Holotype 2, NEw GuINEA: Wau & Mt Keinde, 2110 m, vi. 1969 (Heinrich) (HKT). Paratypes. NEw GUINEA: 3 9, 8 J, NE., Mt Kaindi, 2350 m, vi. 1971 (Sedlacek & Samuelson) (BPBM); 7 9,5 3, NE., Mt Kaindi, 2350 m, ix. 1966 (Samuelson) (BPBM); 1 3, NE., Bulldog Road, 2405 m, v. 1969 (Sedlacek) (BPBM); 1 9, Papua, Dimifa, 2200 m, x. 1958 (Gressitt) (BPBM); 1 9, 1 g, Papua, Mt Hagen, 2000 m, vi. 1969 (Heinrich) (HKT). Heteropelma aello sp. n. (Text-figs 34, 53) DESCRIPTION. Lower face 0-:70—0-80 times as broad as high; clypeus apically simply convex, in profile weakly swollen; mandibles twisted about 40°, the upper tooth 2-50 times as long as lower. IOD 0-70-0-90 times as long as OOD; posterior ocellus separated from occipital carina by about 0-90 times maximum ocellar diameter. Head constricted, vertex with coarse scattered punctures; occipital carina medio-dorsally flattened. Flagellum of moderate length, 59- to 61-segmented; ist flagellar segment 2:10-2:50 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with elongate black pubescence. Pronotum coriaceous, anterior hook vestigial. Mesoscutum coarsely and very closely punctate, matt; notauli indistinct; mesoscutal pubescence short and dense, in 9 badious, in 6g testaceous. Mesopleuron dorsally coriaceous, latero-ventrally with fine even punctures; TAXONOMY OF GENUS HETEROPELM4A 177 epicnemial carina reaching about 0-40 of way up mesopleuron, its upper end widely separated from anterior pleural margin; sternaulus weak. Scutellum longitudinally concave, elongate, punctate. Posterior transverse carina of mesosternum broadly interrupted centrally. Pro- podeum reticulate with a pair of acute lateral protuberances, ventrally weakly sculptured; propodeum dorsally 0-56—0-66 times as long as broad. Mid tibial spurs subequal, the outer about 1-20 times as long as the inner. Hind leg with basitarsus 0-50 times as long as tibia, 2-20—2-60 times as long as 2nd tarsal segment; 2nd tarsal segment 1-80—2-10 times as long as 3rd tarsal segment; trochanter 0o-80—0-90 times as long as trochantellus ventrally; hind tarsal claw abruptly curved, unusually slender, with a small basal lobe. 4 with 2nd hind tarsal segment ventrally strongly impressed. Wings strongly infumate. CI = 0:80-0:90; MI = 1-50-1-75; ICI = o-go-1-10; NI = 0°45-0°55. Forewing length 13-16 mm. Gaster with tergite 2 2-30 times as long as tergite 3; 2 with ovipositor 0-67 times as long as tergite 2. 3 genitalia. Aedeagus in profile subequally bilobate; lateral sclerotized area apically simply acute. Head, gaster, hind femora and basitarsi except distally, also usually ill defined areas on mesoscutum, pronotum and mesosternum blackish; face centrally usually, distal apex of hind basitarsus and hind tarsi 2-5 stramineous; thorax, propodeum, anterior two pairs of legs and hind coxae and trochanteral segments reddish brown. VARIATION. The areas of black colour on the thorax were observed to be variable in their extent. REMARKS. This species is apparently closely related to H. celeno from which it may be distinguished not only in the characters mentioned in the key but also in having a more strongly developed epicnemial carina, more strongly twisted mandibles and having the NI larger. Host REcoRDsS. None available. MATERIAL EXAMINED. Holotype 2, NEw Guinea: SE., Iongai, 1700-1900 m, 9.xi.1965 (Sedlacek) (BPBM). Paratypes. NEw GuINEA: 1 g, NE., Lake Siruki, 2800-2900 m, 15.vi.1963 (Sedlacek) (BPBM); 1 3, SE., Mt Giluwe, 2500-3300 m, 2.vi.1963 (Sedlacek) (BPBM). Heteropelma scaposum (Morley) (Text-figs 30, 37, 49, 60, 64) Exochilum scaposum Morley, 1913a:75. Lectotype 9, AustTRALIA (BMNH), selected by Morley, designated by Townes ef al., 1961 : 322 [examined]. Exochilum atrichiosoma Morley, 1913a:77. Lectotype 9, Austratia (BMNH), selected by Morley, designated by Townes ef al., 1961 : 320 [examined]. [Synonymized by Gauld, 1974 : 543-] Exochilum scaposum Morley; Turner, 1919 : 555. Exochilum atrichiosoma Morley; Turner, 1919 : 555. Heteropelma atrichiosoma (Morley) Townes ef al., 1961 : 320. Heteropelma scaposum (Morley) Townes e¢ al., 1961 : 322. Hetervopelma scaposum (Morley); Gauld, 1974 : 543. Description. Lower face 0-75—-0°85 times as broad as high; clypeus apically almost truncate, in profile flat; mandibles not twisted, the upper tooth about 2-00 times as long as the lower; 178 i) DU GAUED IOD 1:20-1:40 times as long as OOD; posterior ocellus separated from occipital carina by about 1:20 times maximum ocellar diameter. Head not posteriorly constricted, vertex with deep scattered punctures; occipital carina mediodorsally flat or dipped. Flagellum moderately long, 48- to 58-segmented, Ist flagellar segment 2-20—2-60 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with pubescence short to long, always pale. Propodeum punctate, anterior hook moderately well developed. Mesoscutum usually punctate; notauli weak but reaching beyond the centre of the scutum; mesoscutal pubescence usually short and pale. Mesopleuron dorsally rugose, latero-ventrally punctate; epicnemial carina reaching above centre of mesopleuron, bowed and with upper end close to anterior pleural margin; sternaulus moderately impressed, broad. Scutellum slightly convex in longi- tudinal profile, usually with a shallow concavity posteriorly, subquadrate, punctate. Posterior transverse carina of mesosternum complete, rarely rather weak centrally. Propodeum reticulate without lateral protuberances, ventrally sculptured; propodeum dorsally 0-70-0-9g0 times as long as broad. Mid tibial spurs almost equal in length. Hind leg with basitarsus 0-50-0-60 times as long as tibia, 2:30-2:50 times as long as 2nd tarsal segment; 2nd tarsal segment 2:20-2:80 times as long as 3rd; trochanter o-80—0-90 times as long as trochantellus ventrally; hind tarsal claws geniculate, basally lobate. 4 with 2nd hind tarsal segment impressed ventrally. Wings very weakly infumate. CI = 0:70-0:90; MI = 1°55-1:65; ICI = 0-85-1:20; NI = 0:50-0:60. Forewing length 6-15 mm. Gaster with tergite 2 1-80—1-g90 times as long as tergite 3; tergite 3 0-90 times or more (usually more than 1-oo times) as long as tergite 4; 9 with ovipositor about 0-60 times as long as tergite 2. 6 genitalia. Distivolsella with spines scattered on clasping face. Aedeagus in profile apically somewhat obliquely truncate; lateral sclerotized area broad, apically indistinctly defined. Head blackish; lower face and orbital marks yellow; antennae black to reddish, scape ventrally yellow. Thorax black, tegulae yellow. Anterior two pairs of legs yellow; hind legs red-brown, with dorsum of trochanter, trochantellus, distal apex of tibia and proximal end of basitarsus black; segments 2-4 and distal part of basitarsus yellow. Gaster reddish with dorsum of tergite 2 infuscate. VARIATION. This species was observed to have one of the largest size ranges of any Heteropelma species. Smaller specimens were observed to have smaller values of CI than larger specimens. Larger specimens were also disproportionately more hairy, especially on the thorax. The shape of the head was observed to be rather variable. Examination of a long series of specimens from a number of localities has shown that specimens from coastal regions of the southern half of Australia are generally larger and with darker antennae than are those from Queensland. The type-material of H. atrichiosoma is merely a less yellow marked form of H. scaposum. A number of intermediates have been seen thus confirming the opinion that the former species is synonymous with the latter. REMARKS. The basally dark basitarsus distinguishes the Australian species of Heteropelma from all others except the South American species. H. scaposum is closely related morphologically to H. perniciosum and H. flavitarse. It is distinct from the latter in having the hind tarsal claws geniculate and basally strongly lobate as well as being less hirsute and less extensively dark marked. It differs from H. perniciosum less markedly, indeed the only reliable features permitting separation are the relative lengths of the 3rd and 4th gastral tergites and the coloration. Whether these differences will remain when more material is available is yet to be ascertained. TAXONOMY OF GENUS HETEROPELMA 179 H. scaposum is the commonest Australian Anomalonid and as a common parasite of Heliothis armigera may be an insect of considerable economic importance. As yet no worker has investigated the biology of this species. It has been possible to examine the cephalic capsule of the final instar larva of this species. It was found to be similar to that of H. calcator except that the hypostoma was straighter and the labial sclerite more complete ventrally (Text-figs 36, 37). Host RECORDS. Girault (1925) recorded this species as a parasite of Heliothis armigera (Hiibner). Specimens forwarded to the Commonwealth Institute of Entomology for identification are from: Heliothis zea (Boddie) (=obsoleta F.) (Ballard, xii. 1925) Heliothis armigera (Hiibner) (Department of Agriculture, New South Wales, vi. 1974) Omphaloides australasiae (F.) (Williams, vi. 1973). DISTRIBUTION. Australia. MATERIAL EXAMINED. Exochilum scaposum Morley, lectotype 9, AUSTRALIA: Queensland, Mackay (Turner) (BMNH); paralectotypes, 2 9, 4 g, same data as lectotype (BMNH). [In previous work (Gauld, 1974) only one 2 paralectotype was mentioned; subsequently a further 9 which must also be considered to be a paralectotype was discovered in the accessions of the BMNH. Morley failed to state the number of specimens upon which his description was based so that other paralectotypes may also exist. ] Exochilum atrichiosoma Morley, lectotype 9, AUSTRALIA: Victoria (du Boulay) (BMNH); paralectotypes 2 9, same data as lectotype (BMNH); 1 92 ‘Western Australia’. [This specimen which bears no additional data, was labelled as ‘co-type’ by Morley. It is presumably the specimen or one of the specimens from Champion Bay, Western Australia, to which Morley refers. The specimens collected by French in 1901, which are also paralectotypes, have not been located. ] In addition to material listed by Gauld (1974) the following specimens have been examined: AUSTRALIA: I 9, 4 3, Queensland, Biggenden (H. Frauca) (ANIC); 42, 34, Q., Brisbane, 1911-1913 (H. Hacker) (QM); 49, Q., 50 km west of Brisbane, li. 1961 (Gressitt) (BPBM); 1 9, Q., Gatton, xii. 1925 (Ballard) (BMNH); 1 9, Q., Halifax, vi. 1919 (B. Williams) (BPBM); 4 3, Q., Mt Glorius (Stvaatman) (ANIC); 29,1 4, Q., Mt Glorius, ii. 1961 (Gressitt) (BPBM); 1 g, Q., Mt Glorius, i. 1963 (Brooks) (BPBM); 1 3, New South Wales, Beechcroft, vi. 1973 (Williams) (BMNH); I g, Western Australia, Bejoording, ix. 1950 (McMillan) (WAM); 1 3, W.A., Carnac Is., ix. 1934 (Norris) (HKT); 2 9, 7 J, W.A., Coorow, 1952 (McIntosh & Calaby) (ANIC); 2 9, W.A., Cottesloe, ix. 1948 (Glauret) (WAM); 1 3, W.A., Cottesloe, xi. 1945 (Glauret) (WAM); 1 3, W.A., Dumbleyung, x. 1963 (Udell) (WAM); 5 9, 4.3, W.A., Merredin, x. 1952 (Douglas) (WAM); 2 9, 1 3, W.A., Mogamber, x. 1938 (A. Turner) (QM); 1 9, W.A., Nedlands, ix. 1963 (Fuller) (WAM); 2 9, W.A., Northam, xi. 1963 (Sedlacek) (BPBM); 1 9, W.A., Nukarni, ix. 1948 (Willis) (WAM); 19,64, W.A., Rottnest Is., 1936/7 (Glauret) (WAM); 29, W.A., Yallingup, xi. 1942 (Glauret) (WAM); 2 9°, Tasmania, Hobart, i. 1934 (A. Turner) (QM); 1 9, T., Laun- ceston, li. 1916 (Lutiler) (QM). 180 I. D. GAULD Heteropelma flavitarse (Brullé) (Text-fig. 31) Anomalon flavitavse Brullé, 1846: 171. Holotype g, AUSTRALIA (lost). Anomalon trichiosomum Cameron, 1906 : 182. Holotype 2, AusTRALIA (BMNH). [Synonymized by Gauld, 1974 : 542.] Exochilum trichiosoma (Cameron) Morley, 1913a : 77. Barylypa flavitarsis (Brullé) Morley, 19134 : 81. Heteropelma? flavitarse (Brullé) Townes ef al., 1961 : 321. Heteropelma trichiosomum (Cameron) Townes ef al., 1961 : 322. Hetevopelma fiavitarse (Brullé) Gauld, 1974 : 542. DESCRIPTION. Lower face 0:70-0:80 times as broad as high; clypeus subtruncate apically, in profile flat; mandibles not twisted, upper tooth about 2-50 times as long as lower; IOD 1:20-1:30 times OOD; posterior ocellus separated from occipital carina by a distance about equal to the maximum ocellar diameter. Head not posteriorly constricted, vertex punctate; occipital carina mediodorsally flattened. Flagellum of moderate length, stout, 60- to 65- segmented; ist flagellar segment 2-80—3-00 times as long as 2nd; interantennal lamella strong, in profile almost quadrate. Face with long badious pubescence. Pronotum punctate, anterior hook weak. Mesoscutum punctate; notauli weak, reaching to centre of scutum but not discernible at anterior extremities; mesoscutal pubescence blackish. Mesopleuron dorsally coriaceous, ventrolaterally coarsely punctate; epicnemial carina not reaching above lower 0-4 of mesopleuron, its upper end not close to anterior pleural margin; sternaulus moderately strong. Scutellum moderately convex, subquadrate, rugosely punctate. Posterior transverse carina of mesosternum complete. Propodeum finely reticulate, without lateral projections, ventrally sculptured; propodeum dorsally 0-70—-0-80 times as long as broad. Mid tibial spurs equal. Hind leg with basitarsus 0-50—0-60 times as long as tibia, 2:50-2:80 times as long as 2nd tarsal segment; 2nd tarsal segment 2-20—2-30 times as long as 3rd; trochanter about 0-80 times as long as trochantellus ventrally; hind tarsal claws simply curved with a weak basal lamella. Male with 2nd hind tarsal segment ventrally impressed. Wings moderately infumate; CI = 0-80-0-85; MI = 1:50-1:60; ICI = 1:00-1:20; NI = 0-40-0°50. Forewing length 13-16 mm. Gaster with tergite 2 1-g0—2-00 times as long as tergite 3; tergite 3 about I-Io times as long as tergite 4. Q with ovipositor 0:60-0:65 times as long as tergite 2. 4 genitalia similar to those of H. scaposum. Head black, lower face centrally (rarely entirely) and small mark on vertex yellow; flagellum black, scape indistinctly red ventrally. Thorax black, tegulae red. Legs red-brown, forelegs paler; hind tibiae, except proximally, and hind basitarsus proximally black; hind tarsi 2-4 yellowish. Gaster dark red, dorsally and ventrally infuscate. VARIATION. Some variation in the facial colour was observed. The black areas are typically present as a pair of stripes extending from tentorial pits to antennal bases, but these stripes vary in width until in extreme cases the entire lower face is black except for a small central yellow mark. In other cases the width of the stripes is reduced and in isolated specimens the stripes were not discernible except as small infuscate areas above the anterior tentorial pits and below the antennal bases. Males were generally observed to have the face less black marked than females. REMARKS. The tarsal claws, ratio of hind tarsal lengths, clypeal structure and head shape would seem to indicate that this species is one of the least specialized species of Heteropelma. This observation is discussed further below (p. 206). This species is probably closely related to the other Australian species. TAXONOMY OF GENUS HETEROPELMA 181 Host REcorRDS. None known. DISTRIBUTION (Map 2). This species is apparently restricted to the southern part of Australia and Tasmania. MATERIAL EXAMINED. Anomalon trichiosomum Cameron, holotype 9, AUSTRALIA: no further data (BMNH). In addition to the material listed by Gauld (1974) the following specimens have been examined: AUSTRALIA: I 9, Tasmania, Eaglehawk Neck, 11. 1913 (R. E. Turner) (HKT); 2 3, Western Australia, Guildford, viii. 1962 (McMillan) (WAM); 1 3, W.A., Kings Park, iv. 1974 (Postmus & Wade) (WAM); 1 9, W.A., Norrogin, x. 1936 (Douglas) (WAM); 1 9, W.A., Yallingup, x. 1913 (R. E. Turner) (BMNH). Heteropelma perniciosum (Turner) Exochilum perniciosum Turner, 1919 : 554. Lectotype 9, AusTRALIA (BMNH), designated by Gauld, 1974 : 544 [examined]. Hetervopelma perniciosum (Turner) Townes ef al., 1961 : 321. Heteropelma perniciosum (Turner); Gauld, 1974 : 544. DEscRIPTION. Lower face 0-60—0-65 times as broad as high; clypeus apically simply convex, in profile slightly swollen; mandibles not twisted, upper tooth about 1-80 times as long as lower. LOD 1:20-1:30 times as long as OOD; posterior ocellus separated from occipital carina by about 1-20 times ocellar diameter. Head weakly constricted posteriorly, vertex sparsely but deeply punctate; occipital carina mediodorsally evenly rounded. Flagellum moderately long, 45- to 48-segmented; Ist flagellar segment about 2-00 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with long sparse pubescence. Pronotum coriaceous, anterior hook strong. Mesoscutum coarsely punctate; notauli weak but discernible beyond centre of mesoscutum; mesoscutal pubescence short and pale. Meso- pleuron coriaceous, latero-ventrally closely punctate; epicnemial carina reaching to centre of pleuron, bowed so that upper end approaches anterior pleural margin; sternaulus weak. Scutellum longitudinally concave, elongate, coarsely puncto-striate. Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral projections, ventrally sculptured; propodeum dorsally about 0-80 times as long as broad. Mid tibial spurs equal. Hind leg with basitarsus about 0-55 times as long as tibia, 2-10—2-20 times as long as 2nd tarsal segment; 2nd tarsal segment 2-40 times as long as 3rd; trochanter 0-80 times as long as trochantellus ventrally; hind tarsal claws geniculate, with strong basal lamella. Male unknown. Wings very weakly infumate; CI = 0:80-0:85; MI = 1:50-1:60; ICI = 1:10-1:20; NI = 0:50. Forewing length 8-9 mm. Gaster with tergite 2 about 2-40 times as long as tergite 3; tergite 3 0-70-0-80 times as long as tergite 4. 9 with ovipositor about 0-60 times as long as tergite 2. Head black, lower face, marks on vertex and genae reddish; antennae red, proximally darker. Thorax reddish brown, mesoscutum and mesosternum black marked. Legs red, hind tibia distally and basitarsus proximally black. Gaster unicolorous red. Remarks. The validity of this species must remain questionable for several reasons. Firstly, the two females are very similar to H. scaposum, differing only in colour and relative lengths of tergites 3 and 4. Secondly, although extensive collections of Australian specimens have been examined, no further examples of this species have been seen. However, as the fauna of Western Australia is as yet 182 ils IDs (Ey 1U1 1B) only poorly collected it is conceivable that this species has been overlooked by all collectors since Turner. In the present work, therefore, H. perniciosum is provisionally retained as a distinct species. Host RECORDS. None available. DISTRIBUTION. Australia (Western Australia only). MATERIAL EXAMINED. Exochilum perniciosum Turner, lectotype 9, AUSTRALIA: Western Australia, Yallingup (Turner) (BMNH); paralectotype 1 2, same data as lectotype (BMNH). Heteropelma quodi (Vachal) (Text-figs 7, 45) Anomalon quodi Vachal, 1907 : 120. Holotype g, NEw CaLeponia (?MNHN). Heteropelma quodi (Vachal) Townes é¢ al., 1961 : 321. The Vachal types are all believed to be in the MNHN, Paris. Mme Kelner- Pillault was so kind as to search for the holotype of H. guodi and, although she located the type-material of other species described by Vachal in the same publica- tion, she was unable to locate that of H. quodi. I have before me a pair of specimens of this distinct species, the male of which agrees exactly with the description given by Vachal (1907). The descriptions below are based on these specimens. DEscRIPTION. Lower face 0:62-0:66 times as broad as high; clypeus apically simply convex, in profile almost flat; mandibles twisted about 40°, the upper tooth about 3-00 times as long as the lower. JOD 1-30—1-50 times as long as OOD; posterior ocellus separated from occipital carina by a distance about equal to an oceliar diameter. Head constricted, vertex coarsely but not closely punctate; occipital carina medio-dorsally weakly flattened. Flagellum of moderate length, 51- to 53-segmented; 1st flagellar segment 1:90-2:10 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with moderately long pale pubescence. Pronotum strongly punctate, anterior hook moderately developed. Mesoscutum coarsely punctate, subpolished; notauli strongly impressed, reaching beyond centre of scutum; meso- scutal pubescence short, badious. Mesopleuron dorsally rugose, latero-ventrally with numerous finely regularly interspaced punctures; epicnemial carina strong, its upper end not reaching to centre of pleuron and separated from anterior pleural margin; sternaulus weak. Scutellum weakly convex, somewhat elongate, coarsely punctostriate.- Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral protuberances, ventrally strongly carinate; propodeum dorsally 0-80 times as long as broad. Mid tibial spurs subequal, the outer about 1-20 times as long as the inner. Hind leg with basitarsus 0-50 times as long as tibia, 2-20—2-50 times as long as 2nd tarsal segment; 2nd tarsal segment 1:80-2:30 times as long as 3rd tarsal segment; trochanter ventrally 0-70 times as long as trochantellus; hind tarsal claw geniculate, with moderately developed basal lobe. 4 with 2nd hind tarsal segment ventrally impressed. Wings blackish; CI = 0-80-0-90; MI = 1-45-1-65; ICI = 1:20-4:00; NI = 0-55-0°65. Forewing length 9-10 mm. Gaster with tergite 2 about 2-10 times as long as tergite 3; 9 with ovipositor 0-60 times as long as tergite 2. ¢$ genitalia. Distivolsella with spines arranged in apical cluster and with a short row of double spines on ventral margin. Aedeagus in profile apically subequally bilobate; lateral sclerotized area apically indistinctly defined. ; TAXONOMY OF GENUS HETEROPELMA 183 Insect predominantly black, lower face, marks on genal orbits, anterior two pairs of legs (except for the coxae partially) and hind tarsi (except basitarsus proximally) yellow. VARIATION. The 9 was observed to have the fore coxae darker than the 3. REMARKS. The form of the hind legs and head shape indicate that this species is closely related to the Australian species. It is quite distinct from the Papuan species-group in having a simple propodeum and an obvious pronotal hook. Gupta (1962) found in the Ichneumonid genus Theronia a close relationship between Australian and New Caledonian species and a similar relationship has been observed in species of the genus Agrvypon (Gauld, unpublished data). This data would seem to indicate that although New Caledonia is often placed in the Papuan zoogeographic region at least part of its fauna has been derived, possibly through chance migration, from mainland Australia and not from New Guinea via the Solomon Archipelago. Host RECORDS. None available. DISTRIBUTION. New Caledonia. MATERIAL EXAMINED. NEw CALEDONIA: I Q, Foret de Thi, ii. 1957 (Rageau) (BMNH); 1 3, Mt Koghi, x. 1967 (Sedlacek) (BPBM); 1 3, Bourail, xii. 1930 (Cheesman) (BMNH). Heteropelma savaiiense (Fullaway) (Text-figs 9, 25, 39) Therion savatiense Fullaway, 1940 : 400. Holotype 3, SaMOAN ISLANDS (BPBM) [examined]. Hetevopelma savaiiense (Fullaway) Townes et al., 1961 : 322. DESCRIPTION. Lower face 0-63 times as broad as high; clypeus apically simply convex, in profile weakly swollen; mandibles twisted about 35°, the upper tooth more than 2-00 times as long as the lower. JOD 1-10 times as long as OOD; posterior ocellus separated from occipital carina by 1-10 times maximum ocellar diameter. Head somewhat constricted, vertex coarsely punctate; occipital carina dorsally slightly flattened. Flagellum incomplete, Ist flagellar segment 1-90 times as long as 2nd; interantennal lamella weak, in profile obtusely triangular. Face with short sparse pubescence. Pronotum granulate, anterior hook weak. Mesoscutum very coarsely rugose, matt; notauli strongly impressed, reaching to centre of scutum; mesoscutal pubescence short, white. Meso- pleuron dorsally rugose, latero-ventrally sparsely punctate; epicnemial carina strong, close to anterior margin of pleuron, its upper end reaching pleural margin slightly above the centre of the mesopleuron; sternaulus strongly impressed. Scutellum weakly convex, subquadrate, punctostriate. Posterior transverse carina of mesosternum complete, centrally somewhat faint. Propodeum reticulate, without lateral projections, ventrally with strong carinae; pro- podeum dorsally 0-80 times as long as broad. Mid tibial spurs subequal, the outer 1-30 times as long as the inner. Hind leg with basitarsus 0-50 times as long as tibia, 2:40 times as long as 2nd tarsal segment; 2nd tarsal segment 1-90 times as long as 3rd segment; trochanter 0-90 times as long as trochantellus ventrally; hind tarsal claw geniculate, with weak basal lobe; tarsus 2 ventrally impressed. Wings hyaline: CI = 0:85; MI = 1:95; ICI = 0:25; NI = 0:26. Forewing length 7-5 mm. Gaster with tergite 2 1-60 times as long as tergite 3. Q unknown. 6 genitalia. Not available for dissection. 184 i DS GAWLD Insect entirely orange-red except for the lower face, genae and anterior two pairs of legs, which are yellowish, and the frons and vertex, which are brown. REMARKS. This species is probably closely related to H. quodi. Very little collection of Ichneumonidae has been undertaken in the islands east of New Guinea. Heteropelma species have been found on two groups, New Caledonia and Samoa. It is possible that a number of other species await discovery in this general area. Host RECORDS. None available. DISTRIBUTION. Samoan Islands. MATERIAL EXAMINED. Holotype g, SAMOAN IsSLANDs: Savaii, Safune, 650-1300 m, rain forest, v. 1924 (E. Bryan) (BPBM). Heteropelma townesi sp. n. (Text-figs 8, 40, 43, 51) This species is named after Dr H. K. Townes who was kind enough to inform the author of the existence of this Neotropical specics. DEscRIPTION. Lower face 0-60-0-70 times as broad as high; clypeus apically rather flatly convex, in profile weakly swollen; mandibles twisted about 40°, the upper tooth about 3:00 times as long as the lower. JOD about 1-00 times as long as OOD; posterior ocellus separated from occipital carina by about I-10 times maximum ocellar diameter. Head constricted, vertex with scattered punctures, medio-dorsally rugose; occipital carina incomplete medio- dorsally. Flagellum of moderate length, 45- to 47-segmented; 1st flagellar segment 1-g0—2-50 times as long as 2nd; interantennal lamella moderately strong, in profile more or less triangular. Face with short pale pubescence. Pronotum polished with very fine scattered punctures and with strongly impressed margins; anterior hook moderately developed. Mesoscutum polished, evenly punctate, punctures separated by their own diameters; notauli distinct, nearly reaching to posterior margin of mesoscutum; mesoscutal pubescence short, pale. Mesopleuron dorsally longitudinally striate, latero-ventrally finely punctate; epicnemial carina reaching to centre of mesopleuron, bowed so that upper end reaches anterior pleural margin; sternaulus weak. Scutellum longitudinally concave, subquadrate, punctate. Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral protuberances; propodeum dorsally 0-70—0-75 times as long as broad. Mid tibia unicalcarate. Hind leg with basitarsus 0-50—0-60 times as long as tibia, 1-80—3-80 times as long as 2nd tarsal segment; 2nd tarsal segment 2:10-3:50 times as long as 3rd segment; trochanter 1-00—1-05 times as long as trochantellus ventrally; hind tarsal claw strongly geni- culate, basally lobate; ¢ with hind tarsus 2 with a groove-like ventral impression. Wings infumate; CI = 0:40-0:55; MI = 2:10-2:40; ICI = 0-70-1-:10; NI = 0-70-0-90. Forewing length 10-14 mm. Gaster with tergite 2 about 2-30 times as long as tergite 3; 9 with ovipositor 0-50—-0-60 times as long as tergite 2. 6 genitalia. Aedeagus in profile with dorsal lobe strongly reduced; lateral sclerotized region narrow, apically rounded. Insect almost entirely black, yellow marked on lower face, genae, fore legs (except tarsi distally), mid legs (except coxae proximally, distal 0-50 of femur, tibia on proximal o-1o and distal 0-40 and tarsi 2-4), ventral surface of hind trochantellus and band on proximal third of hind tibia; gaster indistinctly reddish brown in part. TAXONOMY OFIGENUS HETEROPELM A 185 VARIATION. There was observed to be an unusual degree of variation in the ratio of the first two hind tarsal segments of this species. One of the females examined, and subsequently not included as a paratype, was observed to be smaller than the holotype and have a red marked gaster. This specimen is also unusual in having a transverse impression dorsally on each hind femur. Whether this is a teratological aberration, or within the normal range of variation for this species is not known. It is possible that this specimen may represent a second and undescribed Neotropical species. Remarks. This species is morphologically quite distinct in having the mid tibia unicalcarate, occipital carina incomplete, trochanters equal to or longer than the trochantelli, etc. However, when compared with the Therionini as a whole it is obvious from the form of the clypeus, form of the hind tarsi, structure of the ovipositor and shape of the claws that this species is an aberrant species of Hetlero- pelma and does not warrant further taxonomic distinction. The affinities of this species are not clear but the structure of the legs and clypeus would seem to indicate that this species may be allied to the H. calcator species-group. Alternatively it is possible that this species may be closer to some of the Australasian species and have developed the elongate tarsi quite separately from the calcator-group. What- ever its ancestry, this species has obviously been separated from the main group of species for a considerable period. This species is not at all closely related to the Nearctic species. Host rEcorDs. None available. DISTRIBUTION (Map 2). Brazil. MATERIAL EXAMINED. Holotype 2, BraziL: Nova Teutonia, Santa Catarina, xi. 1970 (Plawmann) (HKT). Paratypes. I 3, BraziLt: Nova Teutonia, Santa Catarina, xi. 1968 (Plawmann) (HKT); x 3, Nova Teutonia, x1. 1938 (Plawmann) (BMNH). Material excluded from type-series. BraAziL: 1 9, Nova Teutonia, Santa Catarina, xi. 1970 (Plawmann) (HKT). Heteropelma datanae Riley (Text-figs 26, 54) Anomalon flavicorne Brullé, 1846 : 171. Holotype 9, U.S.A.: Philadelphia (PMNHN). Junior primary homonym of Anomalon flavicorne Say, 1823. Heteropelma flavicorne (Brullé) Norton, 1863 : 360. Heteropelma datanae Riley, 1888 : 177. LECTOTYPE 9, U.S.A. (USNM), by present designa- tion [examined]. Hetervopelma flavicorne (Brullé); Ashmead, 1900 : 581. Heteropelma datanae Riley; Ashmead, 1900 : 581. [Heteropelma fulvicorne (Say); Morley, 1913a : 68. Misidentification and incorrect spelling. | Heteropelma datanae Riley; Morley, 19134 : 69. Hetevopelma flavicorne (Brullé); Viereck, 1917 : 285. Heteropelma flavicorne (Brullé); Schaffner & Griswold, 1934 : 145. Heteropelma datanae Riley; Schaffner & Griswold, 1934 : 145. 186 WS 3D, (GAWD) Hetevopelma datanae Riley; Haseman, 1940 : 12. Heteropelma fulvicorne Townes, 1945 : 729. Weplacement name for flavicorne Brullé. Junior secondary homonym of Heteropelma fulvicoyvne (Cameron), 1899 : 104. Syn.n. Heteropelma fulvicovne Townes; Townes, 1951 : 401. Tanypelma fulvicorvne (Townes) Townes, 1971 : 157. Tanypelma datanae (Riley) Townes, 1971 : 157. DESCRIPTION. Lower face 0-80—0-90 times as broad as high; clypeus apically simply convex, in profile flat; mandibles unusually stout, not twisted, the upper tooth subequal in length to the lower. IOD 0:70-0:80 times as long as OOD; posterior ocellus separated from occipital carina by more than 1°50 times ocellar diameter. Head slightly to strongly buccate posteriorly, vertex finely punctate; occipital carina mediodorsally evenly rounded. Flagellum of moderate length, 46- to 55-segmented; 1st flagellar segment 1-65—1-75 times as long as 2nd; interantennal lamella moderately strong, in profile triangular. Face with short pale pubescence. Pronotum rugosely punctate, anterior hook strong. Mesoscutum punctate, notauli weakly impressed but discernible beyond centre of mesoscutum; mesoscutal pubescence short, pale yellowish. Mesopleuron rather evenly punctate except for smooth postero-dorsal region; epicnemial carina not reaching above lower 0:35 of mesopleuron, not reaching anterior pleural margin; sternaulus weak. Scutellum very convex, rugose. Posterior transverse carina of mesosternum complete. Propodeum reticulate, without lateral projections, ventrally sculp- tured; propodeum dorsally rather short, 0-60—0-75 times as long as broad. Mid tibialspurs subequal. Hind leg with basitarsus 0-70—-0-85 times as long as tibia, 5-00—6-00 times as long as 2nd tarsal segment; 2nd tarsal segment about 2-00 times as long as 3rd; trochanter about 0-85 times as long as trochantellus ventrally; hind tarsal claw geniculate, basally lobate; gj with hind tarsus 2 not ventrally impressed, without broadened macrotrichia but often with densely packed microtrichia. Wings infumate; CI = o-go-1-:05; MI = 1-35-1°55; ICI = 0-60—2-20; NI = 0-75-0-95. Forewing length 12-17 mm. Gaster with tergite 2 1-65—-1-75 times as long as tergite 3; 9 with ovipositor about 0-60 times as long as tergite 2. 3 genitalia. Distivolsella with spines on ventral side of a median ridge. Aedeagus in profile apically subequally bilobate; lateral sclerotized region very broad, apically dilated, indistinctly delineated. COLOUR VARIATION. In the southern U.S.A. and in the south-east to about 42°N specimens are reddish brown with the lower face and legs distally paler, whilst in the north-easterm U.S.A. specimens are almost entirely black with only the flagellum yellowish. REMARKS. Examination of material of this species presented a problem not uncommonly encountered in taxonomy. Specimens from lower latitudes are consistently paler coloured than those from the north. Previously the paler form was called datanae whilst the darker specimens were considered to be a separate species, fulvicorne Townes. Some authors when encountering similar variation in other Hymenoptera have retained the forms as distinct subspecies (Townes, 1957 : 253-250). In the present case the author has not used the subspecies category for the following reasons. 1. Many species of Heteropelma with a wide range of geographical distribution exhibit continuous clinal variation in coloration (see p. 168). 2. Both datanae and fulvicorne are parasites of Datana species; indeed both are recorded as parasitizing D. integerrima Grote & Robinson (Townes, 1951). 3. The geographical ranges of the two broadly overlap. An unpublished manu- TAXONOMY OF GENUS HETEROPELMA 187 script by A. B. Gahan (deposited in the USNM) records fulvicorne from Virginia whilst datanae is recorded as far north as New York. 4. Specimens intermediate between the two colour extremes occur. In the BMNH isa black 9 with a pale brown face and a § (from New York) with the thorax and most of the gaster black, but with the lower face yellow and the central segments of the gaster and hind legs brown. 1 ¢ from Georgia (UM, Oxford) is uniformly dark brown with the hind legs black and the lower face pale brown. Some authors regard intergrades between their subspecies as inevitable when there is no geographical barrier present. For example Betrem & Bradley (1971 : 11) when discussing Ethiopian Scoliidae state ‘if a species is constantly different from that in an adjoining region it deserves, in our opinion, a subspecific name; the occurrence of intergrades in an intermediate region is to be expected and is not contradictory’. Whilst not wishing to comment on the taxonomic judgement of these authors it must be pointed out that because of the existence of the intergrades in an intermediate region, one extreme population (subspecies) does not adjoin the other extreme population (subspecies) but adjoins the intermediate intergrades from which it does not differ constantly. Such is the case with H. datanae. One of the main objects of any taxonomic work must be the production of a workable key to the taxa discussed. If, in the present case, two subspecies of datanae were recognized on colour characters, any given key would not work for the intergrades (some 15% of material examined). These specimens would therefore remain indeterminable. The only way the author considers that this can be avoided is by the use of a single taxon, the species H. datanae, and including within it the complete range of colour variation. Host RECORDS. Townes (1951) records this species as a parasite of Datana contracta Walker, D. integerrima Grote & Robinson, D. ministra (Drury) and D. perspicua Grote & Robinson. Ashmead (1900) records it from Sphinx luscitiosa Clemens. DISTRIBUTION (Map 2). U.S.A. east of 100°W. MATERIAL EXAMINED. Heteropelma datanae Riley, lectotype 9, U.S.A.: Virginia, vii. 1880, ex Datana integerrima (TI. Pergande) (USNM). Paralectotypes, 3 2, 2 J, 1 damaged specimen not sexed, same data as lectotype (USNM); 1 9, same data as lectotype (BMNH). U.S.A.: 1 g, Georgia, no further data (UM); 1 9, New York, Wappinger Falls, x. 1934 (BMNH); 19, 1 9, New York, no further data (BMNH); 19, New Hampshire, Monacknock, 1911 (BMNH); 1 9, 1 3, Rhode Island, Westerley, viii. 1951 (H. & M. Townes) (BMNH); 1 9, 1 g, same data (HKT); 2 3, ‘America’, no further data (BMNH). Heteropelma nigricorne (Szépligeti) comb. n. (Text-figs 26, 54) Schizoloma nigvicorne Szépligeti, 1906 : 125. Lectotype 9, Inp1a (TM), designated by Townes et al., 1961 : 323 [examined]. Schizoloma nigricorne Szépligeti; Townes ef al., 1961 : 323. 188 i Dp. GAUL D DESCRIPTION. Lower face 0:80-0:85 times as broad as high; clypeus angularly rounded, with weak lateral protuberances, in profile swollen; mandibles not twisted, the upper tooth about 2:00 times as long as the lower. LOD 0-60 times as long as OOD; posterior ocellus separated from occipital carina by about 1-00 times maximum ocellar diameter. Head not obviously constricted posteriorly; vertex punctate; occipital carina medio-dorsally evenly rounded, often rather weak. Flagellum elongate, 54- to 55-segmented; 1st flagellar segment 2:30-2-50 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with elongate pale pubescence. Pronotum coriaceous, anterior hook strong. Mesoscutum coarsely punctate; notauli moder- ately impressed, discernible to centre of scutum; mesoscutal pubescence short and pale. Mesopleuron dorsally reticulate, latero-ventrally punctate; epicnemial carina reaching about 0-30 of way up mesopleuron, its upper end distant from anterior pleural margin; sternaulus vestigial. Scutellum longitudinally concave, elongate, punctate. Posterior transverse carina of mesosternum complete. Propodeum coarsely reticulate, without lateral projections, ventrally sculptured; propodeum dorsally about 0-70 times as long as broad. Mid tibial spurs subequal, the outer about 1-10 times as long as the inner. Hind leg with basitarsus 0-50 times as long as tibia, 2:30—2-40 times as long as 2nd tarsal segment; 2nd tarsal segment 2:90-3:00 times as long as 3rd segment; trochanter subequal to trochantellus ventrally; hind tarsal claws moderately curved, with a weak basal lobe. Wings infumate; CI = 0-75-0-85; MI = 1:60-1:70; ICI = 0:50-0:60; NI = 0:45-0:55. Forewing length 12-13 mm. Gaster with tergite 2 2-00-2-10 times as long as tergite 3; 9 with ovipositor 0-70 times as long as tergite 2. o unknown. Head black, lower face and marks on genal orbit yellow; flagellum blackish. Thorax and propodeum black, tegulae reddish brown. Legs yellowish brown; anterior two pairs paler than hind pair; hind two pairs of coxae infuscate, hind tibia distally black. ReEMARKS. This species is similar to some variants of H. amictum, especially those which have the lateral clypeal protuberances reduced. However, a number of characters combined in this species have not been observed either to occur at all or in this combination in H. amictum. The lower face of this species is more quadrate than that of H. amictum. This character combined with the colour of antennae, unusually small value of IOD compared to OOD, complete lack of vertical yellow marks and characteristic clypeal shape serve to distinguish this species from others. HosT RECORDS. None available. DISTRIBUTION (Map 2). India (peninsular). MATERIAL EXAMINED. Lectotype 9, Inp1A: Matheran, 800m, 1902 (Bivo) (TM). Paralectotypes, 2 9, same data as lectotype (TM). Heteropelma amictum (F.) (Text-iigs 12, 19, 16; 17, £8, 42, 46, O61) Ichneumon amictus Fabricius, 1775 : 341. Holotype 9 (no further data) (BMNH) [examined]. Ophion amictus (F.) Fabricius, 1798 : 237. Anomalon amictum (F.) Gravenhorst, 1829 : 650. Anomalon (Schizoloma) amictum (F.) Wesmael, 1849 : 120. TAXONOMY OF GENUS HETEROPELMA 189 Anomalon capitatum Desvignes, 1856: 104. Lectotype g, GREAT Britain: England (BMNH) designated by Fitton (1976 : 324) [examined]. Syn. n. Schizoloma bucephalum Brauns, 1898 : 71. Holotype ¢ (no further data) (?lost). [Synonymized by Schmiedeknecht, 1908 : 1463.] Schizoloma fulvicorne Cameron, 1899 : 104. LECTOTYPE , Inpia (UM), here designated [examined]. Syn. n. Anomalon tinctipenne Cameron, 1899 : 112. LECTOTYPE 9, Inpia (UM), here designated {examined}. Syn. n. [Anomalon perornatum Cameron; Szépligeti, 1905 : 12. Misidentification. ] Schizoloma capitata (Desvignes) Szépligeti, 1905 : 34. Schizoloma amictum (F.); Schmiedeknecht, 1908 : 1462. Schizoloma capitata (Desvignes) ; Schmiedeknecht, 1908 : 1463. Schizoloma amicta (F.); Morley, 1913a : 67. Schizoloma capitata (Desvignes); Morley, 19134 : 67. Schizoloma fulvicornis Cameron; Morley, 19134 : 67. Exochilum acheron Morley, 1913a:79. LECTOTYPE Q, Inpia (BMNH), here designated fexamined]. Syn. n. Anomalon tinctipenne Cameron; Morley, 19134 : 83. Schizoloma fulvicornis Cameron; Morley, 19130 : 403. Schizoloma amictum (F.); Morley, 19136 : 404. Exochilum acheron Morley; Morley, 19130 : 412. Exochilum acheron var. scutellatum Morley, 1913b : 412. Syntypes 9 & g, INpiaA (?Calcutta) {not examined]. Syn. n. Anomalon tinctipenne Cameron; Morley, 1913 : 422. Schizoloma amictum (F.); Morley, 1915 : 276. Schizoloma capitatum (Desvignes); Morley, 1915 : 227. Habronyx sachalinensis Matsumura, 1918 : 116. Holotype 9, SAKHALIN (EIHU) [not examined]. {[Synonymized by Uchida, 1928 : 233.] Schizoloma crassicalx Enderlein, 1921 : 11. Holotype g, Sumatra (IZPAN) [examined]. Syn. n. Schizoloma amictum (F.); Uchida, 1928 : 233. Schizoloma amictum var. nigricoxalis Uchida, 1928 : 234. Lectotype 9, SAKHALIN (EIHU), designated by Townes e¢ al., 1965 : 383 [notexamined]. [Synonymized by Uchida, 1958 : 91.] Schizoloma amictum var. intermedium Uchida, 1928 : 234. Lectotype g, Japan (EIHU), designated by Townes ef al., 1965 : 383. [Synonymized by Uchida, 1958 : 91.] Schizoloma coveanum Uchida, 1928 : 235. LECTOTYPE 3, Korea (EIHU), here designated [examined]. Syn. n. Schizoloma amictum (F.); Schmiedeknecht, 1936 : 63. Schizoloma capitatum (Desvignes) ; Schmiedeknecht, 1936 : 64. Schizoloma amictum (F.); Uchida, 1958 : 90. Schizoloma coveanum Uchida; Uchida, 1958 : 91. Schizoloma amictum (F.); Townes et al., 1961 : 322. Schizoloma capitatum (Desvignes); Townes et al., 1961 : 323. Schizoloma crassicalx Enderlein; Townes ef al., 1961 : 323. Schizoloma fulvicovne Cameron; Townes et al., 1961 : 323. Schizoloma tinctipenne (Cameron) Townes é¢ al., 1961 : 323. Heteropelma acheron (Morley) Townes, Momoi & Townes, 1965 : 382. Schizoloma amictum (F.); Townes, Momoi & Townes, 1965 : 383. Schizoloma capitatum (Desvignes); Townes, Momoi & Townes, 1965 : 384. Schizoloma coveanum Uchida; Townes, Momoi & Townes, 1965 : 385. Heteropelma amictum (F.) Townes, 1971 : 157. DESCRIPTION. Lower face 0:55-0:70 times as broad as high; clypeus apically usually with strong lateral protuberances, rarely with these weak; clypeus in profile swollen to excised; 190 ip DAGAULD mandibles twisted about 10°, the upper tooth usually about 2-00 times as long as lower; IOD 0:65-1:05 times as long as OOD; posterior ocelli closer to occipital carina than maximum ocellar diameter. Head of 9 slightly narrowed posteriorly, ¢ from similar to that of 2 to extremely buccate; vertex punctate; occipital carina medio-dorsally rounded or flattened. Flagellum of moderate length, 52- to 63-segmented; Ist flagellar segment 2:00-2:50 times as long as 2nd; interantennal lamella strong, in profile quadrate. Face with moderately long pale pubescence. Pronotum punctate, anterior hook strong. Mesoscutum usually punctate; notauli vestigial to distinctly impressed; pubescence of mesoscutum yellowish. Mesopleuron dorsally coriaceous, latero-ventrally punctate; epicnemial carina reaching 0:30-0:50 of way up mesopleuron, usually with upper end indistinct and not reaching anterior pleural margin. Scutellum usually longitudinally concave, rarely with concavity weak, elongate to subquadrate, usually coarsely punctate. Posterior transverse carina of mesosternum complete. Propodeum reticulate, ventrally sculptured; dorsally 0-70—-0-95 times as long as broad, without lateral protuberances. Mid tibial spurs approximately equal in length. Hind leg with basitarsus 0-44—0-54 times as long as tibia, 1-60—2-30 (isolated specimens up to 2-80) times as long as 2nd tarsal segment; 2nd tarsal segment 1:60-2:30 times as long as 3rd; trochanter 0:70-0:90 times as long as tro- chantellus ventrally; hind tarsal claw geniculate, rarely less strongly curved and with the normally strong basal lobe weaker, 2nd hind tarsal segment of g with strongly impressed ventral area. Wings infumate weakly; CI = 0-68-1-10; MI = 1-40-2-10; ICI = 0:20-0:90; NI = 0-45- 0-65. Forewing length 7-16 mm. Gaster with tergite 2 2:00-2:30 times as long as tergite 3; 2 with ovipositor 0:50-0:60 times as long as tergite 2. 6 genitalia. Distivolsella with spines arranged in apical cluster and with a short row of double spines on ventral margin. Aedeagus in profile apically subequally bilobate; lateral sclerotized region broad, apically dilated, indistinctly delineated. Head black, lower face, vertical marks and genal orbital marks yellow. Thorax black, tegulae usually reddish, very rarely with tegulae, subalar prominences, and occasionally also scutellum yellowish. Anterior two pairs of legs yellow; hind legs reddish, always with tibia distally and often coxae in part, sometimes also femur dorsally blackish; hind tarsi yellowish. VARIATION. This species previously was divided into a number of separate species which, together with H. nigricorne, were placed in a separate genus, Schizoloma. Townes (1971) treated Schizoloma as a synonym of Heteropelma and Gauld (1976) concurred with this opinion. In the present work all species catalogued by Townes et al. (1961; 1965) as Schizoloma (except H. nigritarse) and one species catalogued as Heteropelma (H. acheron) are treated as synonyms of H. amictum. This lumping of species is a rather different approach from that currently employed by many Ichneumonologists, who favour a very narrow species concept and frequently make use of the subspecies category (Gupta, 1962; Townes & Chiu, 1970). Because of this contravention of established procedure a detailed discussion of character variation is included. Townes and co-workers (1961; 1965) have catalogued six species which here are reduced to synonymy. Comparison of the original descriptions and examination of available keys (Morley, 1913a; 19130; Schmiedeknecht, 1936; Uchida, 1958) has enabled the author to select those characters which have been used to facilitate specific separation. Study of extensive material has enabled the relative taxonomic merits of these characters to be assessed. The following characters have been used to facilitate specific separation or show TAXONOMY OF GENUS HETEROPELMA 191 a great deal of variation and therefore may possibly be of use to effect specific separation: I, gross head shape; 2, structure of clypeo-mandibular region; 3, head colour; 4, antennal colour; 5, microsculpture of vertex; 6, microsculpture of mesonotum; 7, sculpture of propodeum; 8, relative lengths of hind tarsal segments; 9, colour of hind legs; 10, wing venation; 11, scutellar colour; 12, shape of hind tarsal claw; 13, geographical distribution. These characters are individually discussed below. 1. Gross head shape. Sexual dimorphism in head shape is not uncommon amongst species of the Ichneumonidae and is particularly obvious in some Therionini. This discussion is concerned only with the gross head shape of the males, which typically have the head somewhat more buccate than that of the corresponding females. Desvignes (1856) described H. capitatum from a pair of males which he separated from H. amictwm by the much more buccate head (Text-figs 12, 13). Although extensive collecting has been undertaken since, no truly intermediate specimens have been discovered. It is therefore quite feasible to regard the two as distinct species as have all previous authors. There are, however, several reasons which have lead the author to believe that H. capitatwm is conspecific with H. amictum. These are as follows. (a) The females of the species are wholly indistinguishable. (b) The species are synchronous and sympatric. H. capitatum has only been taken occasionally (ca 10°) in areas which support a large population of H. amictum, such as heathlands (in the Anomaloninae closely related species frequently manifest clear differences in habitat preference. For example Agrypon anxiwm (Wesmael) prefers open hedgerows whilst A. varvilarsum (Wesmael) is restricted to shaded woodlands). (c) No consistent differences have been observed in other morphological char- acters, including genitalia structure. (d) Although only limited material is available there is apparently no difference in host preference. (e) The geographical range of the two species almost exactly coincides. Towards the centre of the range of geographical distribution in the Indo-Malayan region, the incidence of ‘buccate headedness’ within a population increases. This variation in incidence of buccate headedness of geographically separated populations is tabulated below. LOCALITY No. EXAMINED % BUCCATE HEADED England: Exmoor Dy 10 Nepal: Tapeljung 9 38 Burma: Nam Tami 9 70 Japan: Kamikochi 7 12 Java: Tjibodas region 16 Fi (some intermediates were observed in the Japanese population) 192 ie 1De (EA OED, Whilst much larger samples must be examined before any reliable inferences can be drawn concerning the variation of incidence of ‘buccate headedness’ between the populations, it is probably reasonable to suggest that there is a considerable variation in the frequency of ‘buccate headedness’ between the several populations. The shape of the ocellar triangle was observed to vary between specimens but this variation was apparently linked to that of the head shape. In specimens with buccate heads the ocellar triangle was approximately equilateral (consequently the anterior angle would be 60°) whilst the specimens without a buccate head were observed to have the ocellar triangle posteriorly broadened (with the anterior angle at or more than go°). It may be concluded that gross head shape and shape of the ocellar triangle are not reliable characters for effecting specific separation, as in most populations studied variations of the characters were observed. It is possible that the difference between the simple and buccate headed forms is a single, or relatively few, gene difference as intermediate forms are so uncommon. In buccate headed specimens there was found to be a much greater development of the adductor muscles of the mandibles. It is possible that having these muscles relatively stronger is a selective advantage in certain areas. As the only apparent function of the Ichneumonid mandibles is to effect escape from the host puparium one might expect that in specimens pupating in large hosts with a strong puparium the possession of exceptionally strong mandibular muscles would be a selective advantage. In other regions where such a host is less common and the principal host is a smaller species with a weaker puparium the possession of a buccate head would not bestow any selective advantage, indeed may even be disadvantageous. Thus one would expect variation in the frequency of buccate headedness between populations. 2. Structure of the clypeo-mandibular region. Morley (1913a) separated acheron from other species placed in Schizoloma on account of the difference in clypeo- mandibular structure. In ‘Schizoloma’ the lateral corners of the clypeus are produced until in extreme cases these lateral protuberances entirely conceal the clypeal margin except for a narrow median cleft. In acheron and a number of other specimens examined the lateral protuberances were observed to be very weakly developed. Examples of such specimens have been taken in Burma, India, South Korea, Japan and Taiwan. There are three obvious interpretations of the available material. (a) All represent variants of H. amictum. (b) All are conspecific and represent a distinct species Heteropelma X. (c) All represent distinct species Heteropelma B, 1, K, J, & T respectively. To enable further evaluation of the above hypotheses the specimen groups B, I, K, J & T were compared with typical H. amictwm specimens from neighbouring localities. These data are presented in Table r. In each case excluding that of the Japanese specimens, the variants were found to resemble H. amictum in all except a single additional feature. The characters tabulated were chosen because they are of use in separating a number of species within this genus. If hypotheses b or c were applicable one would have expected 193 TAXONOMY OF GENUS HETEROPELMA snoipeq pue Sg. oo: poonpoid (uemiey) z MOT[EA YSIPPpary 00-Z —GL. -ol. aaoqe se zG—oS Appeam aL, vujadosajaH Sg. 06. (uemiez) t poy o1r-z—ol-1 —GL. —09g- aAoqe se €9-z¢ [euri0ou UNnjIIVUD * FT cL. ol. peonpoid (e110,01A WW “euINg) b Pex Oz-z-0g-1 —09- —oS. aaoqe se _- Apyeem Aaa , vujadosajay C6. 06. (AayeA tee wey ‘eurmng) Fr pey of€-z—-o0g-1 —oL —Sh. aaoqge se 6S—+S yeuliou WnjIVUMDY * FT C6. cL. peonpoid (e]wIS “erpuy) 9 Poy Og-z—-O1-z —¢L. —c¢. aaoqe se Gc—€¢ Appeom I, vuyadosajayy c6. cL. Jeyrep Ajpewtxoid (e[wIs ‘“erpuy) 8 pey of€-.z-ol.1 —CL. —09- Jared Ajye\stp ‘pary gS-1S yew10u MNJIVUD * FT snotpeq pue Cg. og: peonpoid (ea1037) z MOT[PA YSIPpay =: Oz - Z—-OL - I —0g: —09- aaoqe se oS Ayjayer1a pour OM, vumjadosajay snorpeq (reureyy ‘ear0y7) I pue MOTO A oLl.1 og: 09: MOTIOA YSIPpary cS yewiou WNIIVUDY * FT Gsr[se[q z Ajjerjuea Moya XR = OL. - I-09 - I 06. 06. peonpoid (nyeureuey ‘uedef) og: —0g: aaoqe se gs Apyeam [, vujadosaay MO][EA YSIppet 0} c6. 0g: Joyiep Ajpeurxoid (1ysoytuey ‘uede[) gI yoR[q aqeuieA = O1-z7—-0g9-1 —bL. —S¢. razed Ajyeystp ‘pay Z£o-1¢ yewi0u WNIIUUMD * FT G4HNINVXH VXOO CNIH ISUVL GNIH To) IIN wNoTtoo SINHANSHS SNddaAToO INVIYVA SNHWIOUdS FO WNOTOO puc OL 31 WOTTHOVIA AVTIOV Ts 4O YaEaNON JO OILVaY JO Yad NON mnjimMn vujadosajaF{ JO SJULTIVA JO SaIN}eIJ [eOT}IO Jo uostieduioo W I alavy, 194 i Dy GAULED differences in more characters. If b were applicable one would expect more similarity between B, I, K, etc., than these show to H. amic!um. This is not the case. Only in the case of ‘J’ were differences found in more than one additional! character. As this group was represented by only a pair of male specimens investigations could not be taken further. Examination of the available material has shown that reduction of the clypeal protuberance differs in many of the specimens and it is possible to construct a series showing progressive reduction from normal to extremely weakly produced (Burma: Mt Victoria). As the specimens with the weakly produced clypea were otherwise found to be so similar to typical H. amuictim specimens and differences where observed were almost always found to be within the range of variation expected for the species as a whole, it is concluded that these specimens represent variants of H. amictum (that is accept hypothesis a). The specimens therefore do not warrant separate specific status. Accompanying the variation in the shape of the clypeus is a variation in the shape of the mandibles. Those of the typical H. amictum specimens were observed to be stouter and slightly more twisted than those of the specimens with reduced clypeal protuberances. Variation in clypeal shape amongst the Therionini is invariably accompanied by variation in the mandibular shape. In the present work therefore these are regarded as linked characters so the remarks made con- cerning the clypeal variation are equally applicable to mandibular variation. 3. Head coloration. Morley (1913a@ : 66) separated H. amictum from other species on account of its immaculate vertical orbits. The present author has seen no specimens with immaculate vertical orbits although he has examined Morley’s collection. Typically the head was observed to be black with the lower face yellow. Two yellow marks were invariably present on each orbit, one om the vertex and the second just above the centre of the gena. The extent of these marks was observed to vary in the material examined. In some cases the vertical mark was seen to be confluent with the facial aréa of yellow, but generally the two were discrete. It must therefore be concluded that the differences in head coloration are of no use for separating H. amictum from the ‘species’ included here as synonyms. 4. Antennal coloration. Cameron (1899) described /fulvicorne as having the flagellum more evenly fulvous than that of typical H. amictum. Almost all specimens examined were observed to have the antennae reddish, distally yellow, basally externally darker and internally yellowish. In some specimens from the Indo-Malaysian region the yellow areas were less apparent (vide fulvicorne). Some Japanese specimens have been observed to have the basal segments of the antennae black. Philippino specimens were observed to have the antennae more or less entirely badious whilst those of Indonesian specimens were entirely black except for one specimen with the extreme distal apex reddish and the base internally yellow marked. Occasional specimens from other regions have similarly marked antennae. From these observations it may be deduced that antennal coloration may be TAXONOMY OF GENUS HETEROPELMA 195 of some use in separating populations but that the status of these segregates must remain questionable until more characters are examined. 5. Microsculpture of the vertex. Study of species descriptions would lead one to expect that there might be differences between the vertical sculpture of = of the ‘ species’ catalogued as Schizoloma. Most specimens were observed to have the vertex punctate. Some variation in size and density of the puncturation was observed but it was noted that the larger the individual the proportionally more widely separated the punctures were found to be. Consequently more similarity in vertical sculpture was observed between large individuals of geographically widely separated populations, than between large and.small specimens of the same population. Microsculpture of the vertex was not considered to be a useful character to facilitate specific separation. 6. Microsculpture of the mesoscutum (mesonotum of authors). Uchida (1958 : go) distinguished H. coreanum from H. amictum by differences in mesoscutal punctura- tion. The former was described as having the mesoscutum ‘ganz matt, dicht fein punktiert’ whilst that of the latter was described as ‘grésser und zerstreuter als beim coreanum’. The author has examined the lectotype of coreanwm and observed that the puncturation of the mesoscutum is indeed more dense than is usual in H. amictum. The puncturation of the mesoscutum of the female paralectotype is far more like that of H. amictum. There is no other consistent difference between corveanum and amictum, and as there is a considerable range of variation of density of puncturation of the mesoscutum within H. amictwm it is suggested that the puncturation of H. coreanum is an extreme variation of the range of that of H. amictum and that it is therefore justifiable to include this species as a synonym of H. amictum. 7. Sculpture of the propodeum. Differences in propodeal sculpture have been used by Morley (1913@ : 66; 1915 : 226) and Uchida (1958: 90) to separate H. capitatum from H. amictum. That of the former is described as ‘radiating from the centre’ whilst that of the latter is said to ‘form areae’. A very large range of variation in the complex sculpture patterns of the propodea of Heteropelma species have been observed. Large specimens invariably tend to have extremely coarse reticulations with far more enclosed areae. Propodeal sculpture is therefore not considered to be useful in classifying species of Heteropelma. 8. Relative length of the hind tarsal segments. Reference to relative length of the 1st and 2nd hind tarsal segments in species descriptions are generally inaccurate approximations. Thus it appears that there may be differences in the ratios of tarsal lengths between the various described entities considered here. No differences were observed between males and females (other than the broadening of the male tarsi which in a superficial examination may give the impression that the tarsi are proportionally shorter than those of the female). All specimens examined were found to have the hind basitarsus between 1-60 and 2-80 times as long as the second tarsal segment with a mean of I-92. g. Colour of hind legs. Typically the legs are yellowish red with the coxae and trochanteral segments paler ventrally, coxae proximally and distal 0-3 of tibiae 196 i Dr GAWED blackish. The tarsi are immaculately yellow. This colour pattern is fairly constant throughout the European and Indo-Malayan ranges of distribution, with the latter sometimes having the proximal segments less clearly pale marked ventrally. The coxal colour was observed to be more variable. One group of specimens from Nepal showed a range of colour variation. Some specimens had the coxae red with the extreme proximal margins black, other specimens were intermediate and some extreme specimens were observed to have the coxae entirely black. Similar variation has been observed in Japanese species. In some specimens the coxae were observed to be very dark except for the ventral area which was yellow marked. Such specimens were usually, but not always, observed to have the femora badious dorsally. This colour pattern was most common in specimens from Java, Sumatra and the Philippines, although isolated specimens from other regions also exhibit a similar colour pattern. A single specimen from Sulawesi was observed not to have the hind tibia dark marked distally. From these observations one may conclude that coloration of the hind legs alone is not a suitable character for specific separation, but that this colour patterning may be of use in segregating groups of specimens from different regions. 10. Wing venation. Morley (1913a : 66; 19130 : 403; 1915 : 226) used features of wing venation to facilitate species separation. In the two earlier works he used the position of the internal cubital (i.e. 1m—cw) with respect to its point of contact with 2+ 3v7m/M. In this work this position is quantified by the inter-cubital z 10 9° ma z o = oO Cc > h 2 1-3 1-4 15 16 1-7 18 1-9 2-0 2:1 2:2 ___VALUE OF MI, CHarT 1. Histogram showing the range of values of MI in Hetevopelma amictum. (Class width = o-1; sample number 15 9, 15 3]. TAXONOMY OF GENUS HETEROPELMA 197 index ICI. In the latter work Morley used the shape of the radial cell to distinguish two species. This shape is quantified by the marginal index MI. Values of the marginal index are shown in Chart 1. These values were obtained from the measurement of thirty specimens (15 2, 15 ¢) from the Palaearctic/Indo- Malayan regions. Of the specimens measured 7 3 were clearly referable to capitatum whilst 8 were typical amictum. It can be seen from the chart that there is no indication of a bimodal distribution of values. Such a bimodal distribution would be expected if two species differing in values of MI had been confused. It may be assumed therefore that the value of MI is not a reliable character for the separation of species. The cubital cross vein region of the forewing of Anomaloninae is an area of morphological ‘instability’ (i.e. one often encounters considerable intraspecific variation in this region, see Gauld (1976)). Measurement of the ICI was undertaken for a number of typical specimens of H. amictwm and for a number of buccate headed specimens. The results are given below. No. MEASURED RANGE OF ICI MEAN typical H. amictum 15 0-2-0°8 0°55 buccate headed specimens 15 0*3-0°9 0-60 It may be deduced that there is no significant difference between the values of ICI. This character is therefore of no use in facilitating specific separation. iz. Colour of scutellum. Isolated specimens have been found to have the scutellum yellow. No other consistent morphological differences were observed between these specimens and typical specimens of H. amictum. 12. Shape of hind tarsal claws. In H. amictuwm as in most other species of Heteropelma the hind tarsal claws are typically geniculate and basally lobate. In the south-east of the range of distribution some specimens have been observed to have modified claws, that is with the basal lobe reduced. The Javan population was found to have a high incidence of this modification. Of the 18 specimens examined only 4 were found to have normally lobate claws. However, because there is variation in this character it is alone not suitable for facilitating specific separation. 13. Geographical distribution. From the material available it has been established that this species extends from Britain throughout the Palaearctic region to Japan, along the Himalayas into southern China, Taiwan and the Philippines and also in Burma, Sumatra, Java and Sulawesi (Map 2). Ichneumonidae have not been extensively collected and in the past authors tended to regard specimens collected by expeditions to relatively unworked areas as new. Such specimens are described as species novae and the description serves to distinguish them from the better known (usually Palaearctic) species of the genus. Such is the case in question. Numerous species of ‘Schizoloma’ have been described from very little material and generally each differs slightly from other described entities. The problem presented here is whether or not these entities deserve specific status. Examination of fairly extensive material from the above 198 I We CEN WIL ID) 30 pee e AMICTUM 5 SCAPOSUM ® FLAVITARSE 4 NIGRICORNE © DATANAE x TOWNES! Map 2. The world (Mercator projection) showing distributions of Heteyvopelma amictum, H. scaposum, H. flavitarse, H. nigricorne, H. datanae and H. townest. mentioned countries has indicated that there is a large range of variation in several ‘critical’ characters even within a population and that the extent of this variation broadly overlaps that of populations in neighbouring countries. When considering the range of distribution as a whole, widely separated populations were found to exhibit quite different frequencies of the various character conditions. Such differences within a species of broad geographical distribution are to be expected and such clinal variation has been fully discussed by a number of authors (Huxley, 1939; Crowson, 1970). No standard way of designating such clinal variation has been adopted. Huxley (1939) suggested the naming of only the end forms and using formulae to indicate various intermediates. This method has not been adopted by subsequent workers nor is it covered by the current International Code of Zoological Nomenclature. Additional complications are introduced in the present case as the eastern end of the distribution of H. amictum is apparently bifurcate. Using the variations of clinal gradient to delimit natural subspecies (as suggested by Crowson, 1970) is not possible in the present case as insufficient material is available for study. The present author’s approach to the outlined clinal variation is to refrain from designating any population as a subspecies of H. amictum. The status of one population deserves further consideration. A group of 18 specimens from Java were observed to be iather distinct from the remainder of the species. The Javanese specimens were found to have a low percentage incidence of buccate headedness, to generally have the flagellum black, to have the hind TAXONOMY OF GENUS HETEROPELMA 199 femora black marked and to have a high incidence of weakly lobate hind tarsal claws. If clinal gradient may be expressed as a formula of the form, number of character conditions found in 80% of the population A and observed in less than 20% of population B, divided by the linear distance separating the two populations plotted against the linear distance separating the two populations, then the gradient between the Burmese population and the Javan population would be greater than that between the former, and either the European or Japanese populations. This would seem to indicate that the Javan population may deserve separate subspecific status. However, no specimens from the intermediate areas of the Malay peninsula are available for study and only a single specimen from Sumatra was seen. This Sumatran specimen, whilst having many features of the Java population, was observed to have geniculate claws and a buccate head. For the present, therefore, the Javan population is included within H. amictum and not accorded subspecific status. A similar approach was adopted by Perkins (1953) in his work on the British Ichneumoninae. Although he mentions (p. 117) that there are minor differences between the British examples of a species and conspecific European specimens he did not elevate the British population to separate subspecific status. Unfortunately in this work it has only been possible to study the morphology of the adult insect extensively. Such an approach is not wholly adequate when investigating such complex clinal variation. From consideration of the data presented above it has been suggested that all specimens are better retained within a single species, H. amictwm. It is not inconceivable that there may be marked differences in the habitat preference, host selection, behavioural patterns and larval structure between some of the populations. Such data, if and when they become available, may be found to correlate with geographical and character state frequency differences mentioned above. In such a case it may be necessary to reconsider the classification herein proposed. Host RECORDS. This species has been recorded as a parasite of the following lepidopterous larvae: Bena prasinana (L.) (Meyer, 1935; Schmiedeknecht, 1935); Callimorpha dominula (L.) (Meyer, 1935; Schmiedeknecht, 1935); Dasychira pudt- bunda (L.) (Uchida, 1928; 1958; Meyer, 1935; Schmiedeknecht, 1935); Dendrolumus albolineatus Matsumura (Matsumura, 1926; Uchida, 1928); D. sibivicus Tschetverikov (Matsumura, 1926); D. spectabilis (Butler) and D. superans (Butler) (Uchida, 1958); Eupithecia linariata (Denis & Schiffermiiller) (Meyer, 1935; Schmiedeknecht, 1935); Hyloicus pinastri (L.) (Meyer, 1935); Lacanobia oleracea (L.) (Morley, 1915); Lathoe populi (L.) (Meyer, 1935; Schmiedeknecht, 1935); Lithophane ornitopus (Hufnagel) (Meyer, 1935; Schmiedeknecht, 1935); Macrothylacia rubi (L.) (Meyer, 1935; Schmiedeknecht, 1935); Phalera bucephala (L.) (Meyer, 1935; Schmiedeknecht, 1935); Polia nebulosa (Hufnagel) (Morley, 1915); Thaumatopoea processionea (L.) (Meyer, 1935; Schmiedeknecht, 1935). DistrIBUTION (Map 2). This species has been recorded from most northern European countries (Morley, 1915; Schmiedeknecht, 1935) but is possibly absent from the Mediterranean basin as Ceballos (1963) failed to record it from Spain. 200 i Da GAUL It has been recorded from Japan, Korea, Kuriles, Sakhalin and Formosa (Uchida, 1928; 1958), and widely from the U.S.S.R. (Meyer, 1935). In the Oriental region it has been recorded from India and Burma (Morley, 19130) and from Sumatra (Enderlein, 1921). MATERIAL EXAMINED. Ichneumon amictum Fabricius, holotype 2, no further data (BMNH). Anomalon capitatum Desvignes, lectotype g, GREAT BriTAIN: England, no further data (BMNH); (depository of paralectotypes unknown). Amnomalon tinctipenne Cameron, lectotype 2, Inp1A: Assam, Khasia Hills (UM); paralectotypes 3 2, I J, same locality as lectotype (BMNH); 4 9, 1 4, same locality as lectotype (UM). Schizoloma fulvicorne Cameron, lectotype J, INDIA, no further data (UM); 1 g, ? paralectotype, no data (UM). Exochilum acheron Morley, lectotype 9, INDIA: 1851 (Baly) (BMNH); paralectotypes 2 9, 3 g, InpIA: 1851 (BMNH) [Townes ef al. (1961) state that the type depository of acheron is the UM (Oxford). The author has examined the collections in the UM and has seen no specimen that could have indisputably been before Morley when he described this species. However, there are in the BMNH 3 2, 3 3d labelled in 1910 by Morley as Exochilum acheron and from the appended locality data it was clearly apparent that these specimens were syntypes. An intact female was therefore selected as lectotype.] Schizoloma crassicalx Enderlein, holotype 3, SUMATRA: no further data (IZPAN). Schizoloma coreanum Uchida, lectotype J, KorEA: no further data (EIHU). BELGIUM: I 9, no further data (BMNH); 1 9, 1 g, no further data (IRSNB). BuRMA: 2 9, 2 4g, Mt Victoria xti-iv. 1937-38 (Heimrich) (HKT); 1 9, 1 g, Nam Tamai, xi. 1931 (Cranbrook) (BMNH); 4 9, 8 3, Nam Tamai Valley, viti-ix. 1938 (Kaulback) (BMNH); 1 9, 1 g, Nam Tisong, i. 1931 (Ward) (BMNH). CHINA: 2 9, 2 6, Fukien, Shaowu Hsien, vi-x. 1943-45 (Maa) (HKT); 1 3, Kiangsi, Hong San, vii. 1936 (HKT); 2 3, Szechwan, Kwan, viii. 1938 (HKT). GERMANY: I 9, Berlin, no further data (BMNH); 2 9, 2 J, no data (BMNH); 1 9, no data (IRSNB). GREAT BRITAIN: I 9, Cheshire, Delamere Forest, 1904 (BMNH); 1 9, 1 3, Cumbria, Wither- slack, vil. 1922, ex Polia nebulosa (Hufnagel) (Lowther) (BMNH); 20 9, 27 g, Devon, Exmoor, vill. 1971 (Gauld) (BMNH); 7 9, 5 3, Devon, no further data (BMNH); 2 9, 5 6, Grampian ‘Aberdeenshire’, Glen Muick, vii. 1975 (Gauld) (BMNH); 1 9, 1 g, Highland ‘Inverness-shire’, Aviemore, vil. 1929 (Harwood) (BMNH); 1 g no locality data, ex Cucullia sp. (Bond) (BMNH); 7 9, 5 3, no data (BMNH). Inp1a: 49, 6 3, ‘Northern India’, no further data (UM); 19, 1 g, Simla, vill. 1918 (Brunettt) (BMNH); 1 3g, no data (BMNH). IRELAND: 1 J, Eire, Bantry, viii. 1909 (Fletcher) (BMNH). Japan: 2 g, Hanamaki, Iwate, vi. 1953 (Sato) (HKT); 1 9, 17 g, Kami- kochi, vil. 1954 (Townes family) (HKT); 2 9, 1 g, Mt Norikura, vu. 1954 (Townes family) (HKT); 1 g, Mt Wakasugi, ii. 1927 (Yasamatsu) (HKT); 3 9, Sapporo, vil. 1954 (Townes family) (HKT); 1 9, Yamanischi, vi. 1927 (HKT); 1 g, no data (Matsumura) (HKT). Java: 192,14, Dgampang Mts, ix. 1938 (Bengbreng) (BMNH); 1 g, Digeenoeng, Dgampang Tengah, 1939+ (HKT); 1 g, Gunung, Gedeh, viii. 1937 (Sloe) (BMNH); 1 9, Mt Gede, i. 1935 (HKT); 1 g, Nougkodyadjar, v. 1938 (Walsh) (BMNH); 3 8, 4 3, Tjibodas, Gedeh, 1930-7 (Leftinck) (HKT); 2 3, Tjibodas, TAXONOMY OF GENUS HETEROPELMA 201 viii. 1934 (Handschin) (BMNH); 1 9, 1 3, Tjibodas, ili. 1916 (Robinson) (BMNH); I 9, no further data (IRSNB). Korea: 1 dg, vil. 1931 (Sato) (HKT). NEPAL: 1 9, 9 6, Taplejung, between Sangu and Tamrang, x—xi. 1961 (Coe) (BMNH). Puitip- PINES: I g, Abra, Basiwa, iii. 1953 (M.C. Townes) (HKT); 3 g, Kalinga, Babalasang, iii. 1953 (VM. C. Townes) (HKT). SuLAweEsti: 1 9, Todjambol, vii. 1936 (Toxopeus) (HKT). SwepeEn: 1 9, 1 4, Kivik, vii. 1938 (D. M.S. & J. F. Perkins) (BMNH). TAIWAN: I Q, Arisan, vi. 1931 (Gressitt) (HKT); 1 g, Kuraru, vi. 1934 (Gressitt) (HKT); r g Matsumine, v. 1932 (Gressitt) (HKT); 1 g, Sakashen, vii. 1934 (Gressitt) (HKT). U.S.S.R.: 4 9, 3 g, Kiev, no further data (IRSNB); 1 g, Vladivostok (Tosquinet) (IRSNB). ° RELATIONSHIPS BETWEEN THE SPECIES It is not within the scope of a work such as this to discuss the relative merits and shortcomings of phenetic versus phylogenetic classification. It is the opinion of the author that both approaches may be of some use in expanding our knowledge of various groups. In this work therefore phenetic and phylogenetic classifications of the species of Heteropelma are discussed separately below. PHENETIC INTERRELATIONSHIPS. From the species under consideration a large number of characters were considered and those showing little or no inter-specific variation were eliminated. The remaining forty characters were considered to be representative of the interspecific variation. These characters were formulated so as to allow an insect to be scored as either + or —. This system of scoring is not meant to infer that a + character is in any way more significant than a — one. In a few cases intermediates have been found and these were scored + and were not considered to differ from a + or a — character. In a very few cases full information was not available because of lack of material of one or other sex. In these cases percentage similarities were calculated for the number of scored characters compared. The characters used together with their +- and — alternatives are listed below. CHARACTER 4- —- 1. Clypeal shape flat laterally swollen 2. IOD in relation to OOD greater than equal to or less than 3. No. of flagellar segments 49 or less 50 or more 4. Gross head shape constricted not constricted 5. Occipital carina dorsally concave flat or convex 6. Yellow vertical marks present absent 7. Interantennal lamella quadrate not quadrate 8. Facial pubescence white or yellow black or reddish g. Female facial colour entirely yellow not entirely yellow 10. Width of lower face 0-79 or less 0-80 or more compared with height 11. Pronotal hook strong vestigial 12. Notauli present absent 13. Mesoscutum centrally coriaceous punctate 202 CHARACTER . Height of epicnemial carina in relation to 0-4 of height of mesopleuron . Upper end of epicnemial carina . Sculpture of latero-ventral part of mesopleuron . Sternaulus . Scutellum in profile . Posterior transverse carina mesosternum . Propodeal projections . Propodeum ventrally . Propodeum dorsally . Colour of thorax . Colour of scutellum compared with mesoscutum . Mid tibial spurs . Hind basitarsal length . Hind trochanter length . Shape of hind tarsal claw distally . Shape of hind tarsal claw basally . Impressed area on male hind tarsus 2 . Colour of hind basitarsus . Wing colour » Cll . MI NM 5 EN . Tergite 2 length compared with that of tergite 3 . Tergite 3 length compared with that of tergite 4 . Aedeagal apex . Occipital carina dorsally 1 Des KES UIE D) + at or above reaching anterior pleural margin coarsely punctate strong flat complete present smooth o-g— as long as broad black contrasted subequal 3:00+ times length of tarsus 2 0-91 + times length of trochantellum geniculate strongly lobate present entirely yellow black 0-80— 2*00— 0+50— 0:30- 2:00 + o:80— subequally bilobate complete below not reaching anterior pleural margin finely punctate weak humped incomplete centrally absent sculptured o-91+ as long as broad not black concolorous inequal or one absent 2-99— times length of of tarsus 2 o-go— times length of trochantellus simply curved weakly lobate absent not entirely yellow not black o-8r1+ 2:01 + O55 o-31+ 4 1‘99— o:81 + not equally bilobate incomplete By comparison of the various species scored percentage similarities were calculated and these values were tabulated in the form of a similarity matrix (Table 2). From these results an average linkage dendrogram was produced (Chart 2) using the unweighted paired group method (UPGMA) of analysis. State), d Im 3 (Soa ae Sp, a =F Se, a) where S = similarity and a, b, c, & d represents species. few characters in common with the remainder of the genus. Thus It can be seen from the dendrogram that H. celeno and H. aello have comparatively In the 74-76% TAXONOMY OFIGENUS HETEROPEL MA 203 similarity range the greater part of the genus divides into two species-groups, the H. calcatoyr and H. amictum species-groups. Several species separate singly (H. flavitarse, H. savaiiense, H. datanae and H. townest). - elongatum . calcator . fulvitarse . flaviscutellum . perornatum . ocypeta . Nigricorne . amictum . Scaposum . perniciosum . quodi . flavitarse . Savaiiense . datanae . townesi . celeno . aelio PERCENTAGE SIMILARITY NS CHART 2. Dendrogram showing phenetic interrelationships of species of Heteropelma. PROPOSED PHYLOGENETIC INTERRELATIONSHIPS. In any attempt to elucidate the phylogeny of a group (in the absence of a fossil record) the principal source of error is the separation of ancestral characters from derived ones. For example, in Heteropelma the absence of an inter-antennal lamella could either be regarded as an ancestral condition (by assuming that a particular species diverged from the main stock prior to the development of this lamella) or a derived condition (by assuming that the absence of lamella is the result of secondary reduction of the feature). De GAWED I. 204 ouajag 1SAUM07 avunjop Case Gees, (Ge ZS Ase Grd ¢.z9 asualwans 0-z9 0-6 0-09 ¢.z9 asADjuow yf 0.0L 0-oL 0.0 ¢.z9 0.9 (%-o jsarevau 0} Uaye} SoInst,J) uponb mnsoriUusad 0-29 0-29 o-L9 o- +9 0-89 o-SZ o- fe wmnsodvas ¢.z9Q 0-09 0. SZ o-.GL o- FZ 0-08 0-Lg 0-06 S = a Os €.zg o.G¢ 0-69 0-09 o-GL o0.S9 0-GL S.zg C¢.zZ §.z29 ¢.zg o-o4 0-$g 0-€8 0-7g o-LZ o-2g o-Leg ¢.z6 z~ aATAV vyad X20 0.9S 0.6% o-¢Z 0-Z9 0-92 0-66 o-L9 0. zl o-bZ 0-69 o- +9 O- O- wmnqwusosag wmnpjanasvavy AS ¢.LY o-zZ 0-oL o-zZ 0.¢¢ o-S9 0-69 ¢.LL ra G/L o-L9 0-2 ¢.oQ asapquynf *xtjyeul APLMeTIWIS 98e]UVII9g A0JDIIVI ¢.2zS 0.GS¢ o-L9 0-9 o-b¢l 0-59 o-oL o- 2g 0-S8 0-Sg o-LL o- +g 0-78 o-.SZ z-S6 mnqwsuoja 0-09 0-09 o-zZ Le o-zL ¢.z9 o-Llo 0-99 C.LL 0-Sg o-SZL o-LL 0-S8 o-GSZL ¢.28 0-08 onjav ouajaa 1saUmoy apunjpop asuavwwnws asapjuiwyf tponb wmnsoraudsad wmnsodvas wngovmD QUAOIIAD IU pjadi0 mnqwusosag wnpjanasiow py asapquaynf A0JOIDI TAXONOMY OF GENUS HETEROPELMA 205 Consideration of the genus as a whole and its relationship to other genera has enabled the author to tentatively suggest that within Heteropelma the following may be considered to be ancestral character conditions. I, flat clypeus without lateral projections; 2, absence of a pronotal hook; 3, presence of strongly impressed notauli; 4, possession of a complete mesosternal transcarina; 5, absence of lateral propodeal protuberances; 6, presence of a pair of subequal mid tibial spurs; 7, having the hind basitarsus less than 2-90 times as long as tarsus 2; 8, having the hind tarsal claws simply curved; g. having the hind tarsal claws basally weakly lobate; 10, lacking an impressed area on the male tarsi; I1, having CI greater than 0-81; 12, having MI less than 2-00; 13, having the aedeagus terminally unequally bilobate; 14, having the occipital carina complete. All these character conditions are found to occur frequently amongst the more primitive Therionini, that is amongst genera in which the cephalic capsule of the final instar larvae have discernible traces of hypostomal spurs. In the majority of Therionini the hypostomal spurs are entirely lost. Using the method adopted by Gupta (1962) the index of divergence (i.e. number of derived characters) was calculated for each species of Heteropelma (Table 3). The position of the species was then shown in Chart 3 and the proposed phylogenies indicated by lines. It is suggested that four species-groups exist. The largest of these, the calcator- group contains six species, H. calcator, H. fulvitarse, H. elongatum, H. flaviscutellum, H. perornatum and H. ocypeta. H. townesi is also believed to have been derived from this group, but because it is so widely geographically separated from the other species it has diverged morphologically and evolved a number of character con- ditions not found in other species (unicalcarate mid tibiae, high value of MI). All species in this group may be characterized by the possession of elongate hind basitarsi, simple clypeal structure, having the head weakly sexually dimorphic, etc. Within the group there are several very distinct specializations. H. ocypeta and H. perornatum show a tendency to develop strongly constricted heads. H. elongatum and H. flaviscutellum have apparently lost the impressed area on the male hind tarsi. (The reason for assuming these species have lost the impressed tarsal area whilst later assuming that H. datanae may either not have had or have lost a very long time ago such an impressed area are twofold. Firstly, some males of H. fulvitarse, which is morphologically very similar to H. elongatum and H. flaviscutellum, have a reduced tarsal impression, intermediate in form between those of H. calcator and H. elongatum. Secondly, the impressed area typically bears numerous flattened microtrichia which are often also present on ventro-distal (WOIjIPUOD 19zDeILYS 9}LIPUIISZUI UL SayTUsIS S-O ‘UOIZIpUOD psouURApeL o.S fe) o-F fo) ¢.8 I o-.v fe) o-+ fo) o-€ fe) o-.S fo) a o-+ fo) 4 a fo) =) < 0-9 ce) i Gy ° A o-L fe) a C.9 fo) o-¢ fo) ¢.9 fo) o-Z fo) 0-9 fo) AONADUAAIC za AO XHANI 206 Lal Ll ° ° = H F's (07 NS H 4 OrgO) 0" 46") SC) On OL tO) SOn Os On O ICRC © UF 6.0 6 OonBe Olin Ci OD 10). = ze = io eh ek) ie) es SC Se ey eh Noh IS) (oh 19) © So: © © /O— 0) “0 iO nil Omncr yO" Hoy ao BS OO; OF Oy JO) sO: (OOO Or aC!. 1. sou On Seg O. 10%" (Ot 16) Gi pO! Oe 1 1G. Sie On IO) OO) Omi. (OC Omran oO. ©. so 1D NOILIGNOO YALOVUV HO Q0UISIBAIP JO SIOIPUT € aldvy I {UOT}IPUOD IsyOeIeYO [e1}s90Ue = (CO) onjav 0uajaa 1sauUmog apunjop asuarwnvs asanjuiwyy uponb wnsovusag wmnsoqdvas wnjouMn QUAOIIAD IU pag Azo wmnqywousorag wmnyjanasioy asapquaynf 40409109 wnqwsuoja SHIOddS TAXONOMY OF GENUS HETEROPELMA 207 INDEX OF DIVERGENCE CuaRT 3. Proposed phylogenetic interrelationships of species of Heteropelma. apex of the basitarsus. Flattened microtrichia are found on the tarsi of H. elongatum and H. flaviscutellum but have not been observed on the tarsi of H. datanae.) The second and third species-groups are believed to be more closely interrelated than are either to the previously mentioned species-group. One of these, the amictum-group, contains only a pair of species characterized by the obviously swoilen clypeus, relatively short basitarsus, etc. The second of these two groups, the scaposum-group, contains entirely Australasian species, H. scaposum, H. perni- ciosum, H. quodi, H. savaiiense and H. flavitarse. All these species except H. savauense have a bicoloured hind basitarsus. The group as a whole may be characterized by short basitarsus, simple clypeus, etc. Occasional specimens in this species-group have been observed to have a faint trace of lateral clypeal swellings. This and the similarity in form of the hind legs has been interpreted as indicative of a close phylogenetic relationship between this and the amictwm-group. A fourth species-group contains the two Papuan species, H. celeno and H, aello. These are distinct from all other species in a large number of features including propodeal shape, form of tarsal claws, absence of pronotal hook, etc. The affinities of these species are not clear but they do not appear to be at all closely related to the other Australasian species which all belong in the scaposum-group. Certain features of the celeno-group seem to indicate that it diverged from the main stock of the genus a considerable time ago. The structure of the aedeagus, form of the posterior transcarina of the mesosternum, and shape and coloration of the female face are reminiscent of those of Therion species. These two species may possibly be rather specialized descendants of species which diverged from the main Hetero- pelma stock shortly after Heteropelma and Therion separated. Far more evidence 208 De GAwiED is needed, however, before any reliable statements can be made concerning the position of this species-group. It would be of particular interest to examine the first instar larva of these species as considerable differences are known to exist between the first instar larvae of Therion and Heteropelma (Plotnikov, 1914; Tothill, 1922). The position of H. datanae is unclear at present. ACKNOWLEDGEMENTS I would like to thank the Keeper of Entomology, British Museum (Natural History) for providing study facilities and Mr L. A. Mound and Dr M. G. Fitton for their cooperation. I am grateful to the following persons and institutions for loan of material: Miss J. Cardale, ANIC, Canberra, Australia; Dr R. W. Carlson, USNM, Washington, U.S.A.; Dr E. C. Dahms, QM, Fortitude Valley, Australia; Dr P. Dessart, IRSNB, Brussels, Belgium; Mrs C. N. Higa, BPBM, Honolulu, U.S.A.; Dr D.-R. Kasparyan, ZI, Leningrad, U.S.S.R.; Dr D. Kitchener wea Perth, Australia; Mme S. Kellner-Pillault, MNHN, Paris, France; Dr E. Kierych, IZPAN, Warsaw, Poland; Dr J. Oehlke, DEI, Eberswalde, D.D.R.; Mr C. O’Toole, UM, Oxford; Dr J. Papp, TM, Budapest, Hungary; Mr E. C. Pelham-Clinton, RSM, Edinburgh, Scotland; Dr M. Suwa, EIHU, Hokkaido, Japan and Dr H. K. Townes, Ann Arbor, U.S.A. I would like to thank Dr Townes, Dr Fitton and colleagues on the staff of the CIE and BMNH for their advice and cooperation at various stages and for their many helpful criticisms. I am grateful to Miss P. Mitchell for her assistance at various stages and for typing the manuscript. REFERENCES ASHMEAD, W. H. 1900. Hymenoptera. Jn Smith, J. B., Insects of New Jersey (1899) : 501— 613. Trenton. BarBeEy, A. 1927. La fidonie du pin dans les pineraies d’Alsace et de Lorraine. Fev. Eaux et Foréts Dec. 1926 : 1-7, 6 pls. BeEetrREM, J. G. & BrapLeEy, J. C. 1971. The African Campsomerinae. Monografieén ned. ent. Vereen. 6 : 1-326, 6 pls, 52 figs. Brauns, S. 1898. Die Ophionoiden. Aych. Ver. Freunde Naturg. Mecklenb. 43 : 58-72. BRuLtE, A. 1846. Hymenoptera. In Lepeletier de Saint-Fargeau, A. L. 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Anomalinen und Therionen in der Sammlung des Entomologischen Instituts der Hokkaido Universitaet 1. Insecta matsum. 21 : 85-108. VACHAL, J. 1907. Hyménoptéres de la Nouvelle Calédonie. Revue Ent. 26 : 113-123. VierREcCK, H. L. 1917. Guwide to the Insects of Connecticut; Part 3, The Hymenoptera ov wasp- like insects of Connecticut. 824 pp., 10 pls. Hartford. WESMAEL,C. 1849. Revue des Anomalons de Belgique. Bull. Acad. yr. Belg. (2) 16 : 115-139. 212 I. D. GAULD Figs 1-13, heads, dorsal view. 1, Hetevopelma fulvitavse Cameron, ¢; 2, H. fulvitarse Cameron, 9; 3, H. calcatoy Wesmael, 3; 4, H. calcatoy Wesmael, 9; 5, H. elongatum Uchida, 3; 6, H. perornatum (Cameron), 9; 7, H. quodi (Vachal), 9; 8, H. townesi sp. n., g; 9, H. savatiense (Fullaway), 3; 10, H. ocypeta sp. n., 9; 11, H. celeno sp. n., Q; 12, 13, H. amictum (Fabricius), ¢, (12) normal, (13) ‘buccate-headed’. TAXONOMY OF GENUS HETEROPELMA 213 Fics 14, 15, Heteropelma nigricorne (Szépligeti), 9. 14, lower face, anterior view; 15, head, dorsal view. Fics 16-18, H. amictum (Fabricius), 9. 16, lower face, anterior view (showing weakly laterally protruding clypeus); 17, the same, antero-lateral view; 18, lower face, antero-lateral view (showing strongly laterally protruding clypeus). Fics 19-23, interantennal lamella in profile. 19, H. calcatoy Wesmael, 9; 20, H. fulvitarse Cameron, 9; 21, H. peyornatum (Cameron), 9; 22, H. elongatum Uchida, g; 23, H. flavi- scutellum Uchida, g. Fics 24-26, scutellum in profile. 24, H. perornatum (Cameron), g; 25, H. savaiiense (Fullaway), ¢; 26, H. datanae Riley, 3. 214 [hDACAuLD Fics 27-29, pronotum, lateral view. 27, Hetevopelma perovnatum (Cameron), 9; 28, H. fulvitarse Cameron, 9; 29, H. celeno sp. n., 9. Fics 30-32, @ hind tarsal claw; 30, H. scaposum (Morley) ; 31, H. flavitarse Brullé; 32, H. celenosp.n. Fics 33, 34, Q propodeum, dorsal view. 33, H. celeno sp. n.; 34, H. aellosp.n. Fic. 35, H. ocypeta sp. n., apex of mid tibia, internal view. TAXONOMY OF GENUS HETEROPELMA 215 Fics 36, 37, cephalic capsule of final instar larvae, anterior view. 36, Hetevopelma calcator Wesmael; 37, H. scaposum (Morley). Fic. 38, H. flaviscutellum Uchida, ¢ hind tarsi I-IV, ventral view. Fics 39, 40, left forewings, distal part. 39, H. savaiiense (Fullaway) ; 40, H. townesisp.n. Fics 41-43, ¢ hind tarsi II & III, ventral view (detail of pubescence shown only in part). 41, H. fulvitarse Cameron; 42, H. amictum (Fabricius); 43, H. townesi sp. n. 216 I. D. GAULD Fics 44-54, apices of aedeagi, lateral view. 44, Heteropelma perornatum (Cameron); 45, H. quodi (Vachal); 46, H. amictum (Fabricius) ; 47, H. celeno sp. n.; 48, H. calcatoy Wesmael; 49, H. scaposum (Morley); 50, H. elongatum Uchida; 51, H. townesi sp. n.; 52, H. fulvitarse Cameron; 53, H. aello sp. n.; 54, H. datanae Riley. TAXONOMY OF GENUS HETEROPELMA 217 Fic. 55, apex of valvula 2, lateral view. Hetevopelma fulvitavse Cameron. Fics 56-64, Ovarian eggs; 56, H. celeno, detail of ‘sucker’; 57, H. celeno sp. n., dorsal view; 58, H. perornatum (Cameron), dorsal view; 59, H. calcatoy Wesmael dorsal view; 60, H. scaposum (Morley), dorsal view; 61, H. amictum (Fabricius), dorsal view; 62, H. fulvitayse Cameron, dorsal view; 63, H. calcatoy Wesmael, lateral view; 64, H. scaposum (Morley), lateral view. 218 I. D. GAULD INDEX TO RECORDED HOSTS Anarta myrtilli (L.), 166 Bena prasinana (L.), 166, 199 Bupalus piniaria (L.), 165, 166, 167 Callimorpha dominula (L.), 199 Dasychira pudibunda (L.), 199 Datana contracta Walker, 187 Datana integerrima Grote & Robinson, 186, 187 Datana ministra (Drury), 187 Datana perspicua Grote & Robinson, 187 Dendrolimus albolineatus Matsumura, 199 Dendrolimus sibiricus Tschetverikov, 199 Dendrolimus spectabilis (Butler), 199 Dendrolimus superans (Butler), 199 Eupithecia linariata (Denis & Schiffermiiller), 199 Heliothis armigera (Hiibner), 179 Heliothis zea (Boddie), 179 Hyloicus pinastri (L.), 199 Lacanobia oleracea (L.), 199 Lathoe populi (L.), 199 Lithophane ornithopus (Hufnagel), 199 Macrothylacia rubi (L.), 199 Omphaloides australasiae (F.), 179 Panolis flammea (Denis & Schiffermiiller), 166 Phalera bucephala (L.), 199 Polia nebulosa (Hufnagel), 199 Sphinx luscitiosa Clemens, 187 Thaumatopoea processionea (L.), 199 INDEX TO RARASTIZES Synonyms are in ?falics ; principal references are in bold. achevon Morley, 157, 189 aello sp. n., 156, 162, 174, 175, 202, 207 Agrypon Foerster, 182, 191 amictum F., 157, 158, 159, 161, 164, 188, 203 Aphanistes Foerster, 175 atrvichiosoma Morley, 156, 177 binghanu Cameron, 156, 167 bucephalum Brauns, 157, 189 calcator Wesmael, 156, 158, 159, 160, 161, 164, 169, 185, 203, 205 capitatum Desvignes, 157, 189 Casinaria Holmgren, 175 celeno sp. n., 156, 161, 162, 174, 175, 177, 202, 207 coveanum Uchida, 157, 189 crassicalx Enderlein, 157, 189 datanae Riley, 157, 158, 159, 161, 163, 185, 203, 205, 207, 208 elongatum Uchida, 156, 164, 169, 205, 207 flaviscutellum Uchida, 156, 163, 168, 170, 205, 207 flavicorne Brullé, 157, 185 flavitarse Brullé, 156, 163, 178, 180, 203, 207 fulvicoyne Cameron, 157, 189 fulvicorne Townes, 157, 158, 186 fulvitarse Cameron, 156, 161, 164, 165, 166, 167, 170, 173, 205 grossatoy Shestakov, 156, 167 Habronyx Foerster, 162 intermedium Uchida, 157, 189 Metopius Panzer, 176 nigricorne Szépligeti, 157, 160, 161, 164, 187, 190 nigvicoxalis Uchida, 157, 189 ocypeta sp. n., 156, 163, 174, 205 orbitale Morley, 157 panargis Heinrich, 156, 167 perlongum Cushman, 156, 167 perniciosum Turner, 156, 163, 178, 181, 207 TAXONOMY OF GENUS HETEROPELMA 219 perornatum Cameron, 156, 161, 164, 173, scaposum Morley, 156, 159, 161, 162, 163, 174, 205 77, 180, T8I,207 scutellatum Morley, 157, 189 quodi Vachal, 156, 160, 163, 182, 184, 207 tavsale Cushman, 156, 170 tinctipenne Cameron, 157, 189 Therion Curtis, 160, 162, 207, 208 Theronia Holmgren, 183 veticulatum Cameron, 156, 167 sachalinensis Matsumura, 157, 189 townesi sp. n., 156, 161, 162, 184, 203, 205 savaliense Fullaway, 156, 160, 161, 162, tvichiosomum Cameron, 156, 180 183, 203, 207 Trichomma Wesmael, 162, 175 I. D. Gautp, L.I.Biol. COMMONWEALTH INSTITUTE OF ENTOMOLOGY c/o British Museum (NATURAL History) CROMWELL Roap Lonpon SW7 5BD —e wi. ue : a 7) 4 i eee Mt Ye Oey aAy , ; a ees: Ru 7 san Py ire \eG@twt Vo wore m VN! : idles Ate Maire ise eA = ( 1 eee4 , See ae mn oz mt la Si =) 2 a Ps o@ = * q “TY Des nagueaes =~) = _— ~~ ; —-) Melo bert PF > ? A malty \ rulury eat a —e i \ @e, ~ 7 = i ma bi — a a ; : as x at ar) ws “A ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.40. . Hemminc, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounp, L.A. A review of R. S. Bagnall’s Thysanoptera collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. 5. . AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 1098: 68 plates, 15 text-figures. October, 1971. {12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CRossKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . von HaveK,C.M. F. A_ Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. . CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogeus. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND A\\S0 MU ge be GENERAL | ~ 1 NOV 197 A REVISION OF THE : INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA (THYSANOPTERA : PHLAEOTHRIPIDAE) B. R. PITKIN BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 4 , LONDON : 1976 4 ‘ aE, Se * 4 VES be A “ Laem =. : a) “y : i % % fi iM, a m ibe i A; Miki oe po ” ~ i f ete i mr A te AILS Sate i ? ul he ? tt Aba s ‘ ‘ h 7 f ing ‘ r ey ‘sh Hint | . j a ie y iy 4 ne n ray » ch \ hee ‘ : wile >, i ; We : ' eis ‘ ' i 4 i © : - Ma , \ 7 {aero \ ‘ PS aad . ™ ~ =, ' / ( 2 i . ‘ q y ‘ A REVISION OF THE INDIAN SPECIES \ OF HAPLOTHRIPS AND RELATED GENERA (THYSANOPTERA : PHLAEOTHRIPIDAE) BY BRIAN ROY PITKIN Pp. 221-280; 62 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 Vol. 4 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 15 issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at wregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 34 No. 4 of the Entomology serves. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.). ISSN 0524-6431 © Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 28 October, 1976 Price £6.60 A REVISION OF THE INDIAN SPECIES OF HAPLOTARIPS AND RELATED GENERA (THYSANOPTERA : PHLAEOTHRIPIDAE) By B. R. PITKIN CONTENTS Page SYNOPSIS : P ; ; ; : ; : : F : 223 INTRODUCTION ‘ A ‘ ; ‘ - : . : : 223 ACKNOWLEDGEMENTS : : ; : : ‘ : : ; 224 ABBREVIATIONS : : : : : : : ; : : 225 INDIAN RECORDS REFUTED A : : é ‘ ; ; é 225 CHECKLIST OF THE INDIAN SPECIES OF Haplothvips AND RELATED GENERA 226 KEY TO THE INDIAN SPECIES OF Haplothrips AND RELATED GENERA - 228 GENERA AND SPECIES DISCUSSED ALPHABETICALLY : : : : 231 REFERENCES . : ; 2 , : ‘ ; ; : - 276 INDEX . ; : : : : . : 7 ; : : 279 SYNOPSIS Sixty nominal species and two nominal varieties of Haplothvips and related genera are discussed. Of these, two species are described as new. The Indian records of three species are refuted. One new generic synonymy, II new specific synonymies, and 10 new combinations are established. A key for the identification of the 48 species recorded from India is provided. The characteristic features of each genus and species, a brief discussion of any changes in nomenclature and a list of material examined are given. Lectotypes are designated for I1 species. INTRODUCTION THERE are about 4500 described species of Thysanoptera, although recent work on tropical faunas suggests that there may be more than double this number. Of these about 520 species are recorded from India (Ananthakrishnan, 1969). The Thysanoptera may be divided into two suborders, the Terebrantia and the Tubulifera. The latter comprises a single large family, the Phlaeothripidae, and includes about 2700 described species. Of these about 300 are recorded from India. The Phlaeothripidae may be further divided into two subfamilies, the Idolothripinae and the Phlaeothripinae. The Phlaeothripinae is by far the larger subfamily with about 2300 described species. Of these about 230 species are recorded from India. The Phlaeothripinae may be further subdivided into a number of tribes (Priesner, 1960; Mound, 1972), including the Haplothripini. The tribe Haplothripini comprises a group of Haplothrips-like species with the following diagnostic features: antennae eight-segmented; wings usually present 224 Beak. PPR KEN and fully developed, sometimes reduced or absent, when fully developed with or without duplicated cilia; maxillary stylets usually long and retracted far into the head capsule; maxillary bridge usually present; male genitalia with a well developed aedeagus; usually without glandular areas on abdominal sternites. The tribe has a world-wide distribution and comprises more than 300 species. These are found in flowers, at the base of grasses, on dead leaves and branches and as inquilines of galls, but it should be stressed that none of them form galls or cause leaf-rolling as do many species of the tribe Haplothripini. A total of 87 species in 14 genera are recognized from India. The initial object of this study was to find stable diagnostic characters by which Indian species assigned to the haplothripine genera Haplothrips Amyot & Serville and Xylaplothrips Priesner could be recognized. Some of these species, however, are more closely related to species in other genera. The study was therefore extended to cover all Indian species listed by Ananthakrishnan (1969) in Apterygothrips Priesner, Chiraplothrips Priesner, Haplothribs Amyot & Serville, Karnyothrips Watson, Praepodothrips Priesner & Seshadri, Xenothrips Anantha- krishnan (syn. n. of Antillothrips Stannard) and Xylaplothrips Priesner, and any species described subsequently from India in these genera. This group includes 45 described species and two new species. The haplothripine genera not included in this account but represented in India, with the exception of Podothrips Hood and the monobasic genus Segnothrips Ananthakrishnan, may be readily distinguished from Haplothrips and related genera using the key characters given by Ananthakrishnan (1969). Species of Podothrips may be distinguished from other haplothripines by their praepectal plates, which are longer than broad. Segnothrips trivandvensis Ananthakrishnan is being synonymized elsewhere with a species of Haplothrips by Dr J. S. Bhatti (pers. comm.). Six of the seven genera discussed in this account cannot be readily distinguished from each other due to the occurrence of intermediates. Each of these genera, however, exhibits distinct trends and it is on these trends that the generic classification used here is based. In view of the existence of intermediate species, a workable key to genera would be extremely complex and consequently difficult to use. I have therefore written the key in its present form, rather than as a key to genera followed by keys to the species of each genus. The characters used to distinguish species in different genera are not necessarily significant at the generic level. The characteristics of each genus are discussed separately in the text. ACKNOWLEDGEMENTS This study would not have been possible without the active support of many colleagues. Professor T. N. Ananthakrishnan, Dr J. S. Bhatti, Dr B. V. David, Dr C. F. Jacot-Guillarmod, Miss Kellie O’Neill and Dr R. zur Strassen kindly loaned valuable material. To all these I am very grateful. I should also like to thank Dr J. S. Bhatti and Dr L. A. Mound for their advice and criticism. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 225 ABBREVIATIONS The following abbreviations have been used in the text. BMNH British Museum (Natural History), London LCM Loyola College, Madras SMF Senckenberg Museum, Frankfurt USNM United States National Museum, Washington D.C. TNA Professor T. N. Ananthakrishnan INDIAN RECORDS REFUTED The Indian records of Apterygothrips flavus Faure, Haplothrips flavitibia Williams and Xylaplothrips mimus Priesner are refuted for the reasons given below. The Indian records (Ananthakrishnan, 1969) of Apterygothrips hispanicus (Bagnall), Haplothrips andresi Priesner and Haplothrips bagnalli (Trybom), which are accepted here, require confirmation. Apterygothrips flavus Faure Apterygothrips flavus Faure, 1940 : 163. Holotype 9 aptera, SourH Arrica: Cape Province, Middelburg (National Insect Collection, Pretoria) [not examined]. Ahpterygothrips flavus Faure; zur Strassen, 1966 : 161-175. Faure (1940) described flavus from nine apterous females and six apterous males collected on Cenchrus ciliaris L. [Gramineae] at Middelburg, Cape Province, South Africa. Ananthakrishnan (1967) recorded flavus as new to India. None of the specimens on which this Indian record is based represents flavus. They are being described elsewhere as a new species of A pterygothrips (Dr J. S. Bhatti, pers. comm.). Haplothrips flavitibia Williams Haplothrips flavitibia Williams, 1916 : 283-284. Holotype 9, Great Britain: England, Surrey, Merton (Albany Museum, Grahamstown) [examined]. Williams (1916) described flavitibia from six specimens, of unspecified sex, beaten from a hedge of hawthorn (Crataegus oxyacanthoides Thuillier [Rosaceae]) in Surrey, England in July, 1913 and July, 1914. Priesner (1964a) also records flavitibia from Germany. Ananthakrishnan (1952) described a male from Madras collected on the flowers of Portlandia grandiflora L. [Rubiaceae] which he incorrectly designated as the holotype male of flavitibia. This specimen has the base of the mid and hind tibiae dark brown, whereas flavitibia has the mid and hind tibiae completely yellow. Unfortunately I have not examined this Indian male, but in view of the colour differences referred to above it is certainly not flavitibia. In the absence of any other published records, the Indian record of flavitibia is therefore refuted. 226 Bek eran) SPECIMEN STUDIED. Holotype 9, GREAT BriTaIn: England, Surrey, Merton, on hawthorn (Crataegus oxyacanthoides Thuillier) [Rosaceae], vil. 1913 (C. B. Williams) (Albany Museum, Grahamstown). Xylaplothrips mimus Priesner Xylaplothvips mimus Priesner, 1939 : 172-174. Holotype 9, ZarrRE: Rutshuru (SMF) [not examined]. Priesner (1939) described mimus from two females and one male collected on a coffee bush at Rutshuru, Congo (Kinshasa). These specimens apparently have I +1 and 1 +2 sense cones on antennal segments III and IV respectively. Ananthakrishnan (1966) recorded mimus as new to India. This record was based on four females and 11 males collected on grass at Hubli, 21.xi.1964 (TNA). I have examined two females collected on grass at Hubli, 17.xi.1964 (TNA) and identified by Professor Ananthakrishnan as mimus. These specimens, however, have 1 + 2 and 2 4+ 2** sense cones on antennal segments III and IV respectively and I cannot distinguish them from pictipes Bagnall. CHECKLIST OF THE INDIAN SPECIES OF HAPLOLARILS AND RELATED GENERA An asterisk denotes the designation of a lectotype. Genus ANTILLOTHRIPS Stannard, 1957 Xenothvips Ananthakrishnan, 1965) syn. n. graminellus (Ananthakrishnan & Jagadish, 1969) comb. n. *malabaricus (Ananthakrishnan, 1965)) comb. n. *luteus (Ananthakrishnan & Kudo, 1974) syn. n. micropterus sp. n. nayari (Ananthrakrishnan, 1958) comb. n. *varius (Ananthakrishnan & Jagadish, 1969) comb. n. nefvens (Ananthakrishnan, 1972) syn. n. Genus APTERYGOTHRIPS Priesner, 1933a *fungosus (Ananthakrishnan & Jagadish, 1969) comb. n. hispanicus (Bagnall, 1916) *jogensis (Ananthakrishnan & Jagadish, 1969) comb. n. pellucidus (Ananthakrishnan, 1968) comb. n. pini Ananthakrishnan, 19614 rubiginosus (Ananthakrishnan & Jagadish, 1971) comb. n. Genus CHIRAPLOTHRIPS Priesner, 1930 graminellus (Priesner, 1938) priesnevi (Ananthakrishnan, 19610) Genus HAPLOTHRIPS Amyot & Serville, 1843 Subgenus HAPLOTHRIPS Amyot & Serville, 1843 andresi Priesner, 1928 bicolor (Ananthakrishnan, 1964a) comb. n. ceylonicus Schmutz, 1913 INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 227 ganglbaueri Schmutz, 1913 priesnerianus Bagnall, 1933 syn. n. ceylonicus var. vernoniae Priesner, 1921 syn. n. andhva Ramakrishna, 1928 syn. n. gowdeyi (Franklin, 1908) sovoy Schmutz, I913 syn. n. sovorcula Schmutz, 1913 syn. n. longisetosus Ananthakrishnan, 1955 pirus Bhatti, 1967 reuteri (Karny, 1907) tenuipennis Bagnall, 1918 ceylonicus var. mangiferae Priesner, 1933) Subgenus TRYBOMIELLA Bagnall, 1926 apicalis (Bagnall, 1915) articulosus Bagnall, 1926 dervisoy Priesner, 1935 bagnalli (Trybom, 1910) clarisetis Priesner, 1930 euphorbiae Priesner, 1931 nigricornis Bagnall, 1910 vamakvishnai Karny, 1926 syn. n. talpa Priesner, 1930 tirumalraoi Ramakrishna & Margabandhu, 1931 Genus KARNYOTHRIPS Watson, 1922 alpha sp. n. flavipes (Jones, 1912) melaleucus (Bagnall, 1911) mucidus (Ananthakrishnan & Jagadish, 1971) comb. n. nigriflavus Ramakrishna, 1934 Genus PRAEPODOTHIRPS Priesner, 1928 cymbapogoni Ananthakrishnan, 1956 indicus Priesner & Seshadri, 1952 *nigrocephalus Ananthakrishnan, 1964 priesneri Ananthakrishnan, 1955 Genus XYLAPLOTHRIPS Priesner, 1928 debilis Ananthakrishnan & Jagadish, 1971 emineus Ananthakrishnan & Jagadish, 1971 *flavitibia Ananthakrishnan, 1969 flavus Ananthakrishnan, 1964a *inquilinus (Priesner, 1921) inquilinus Ananthakrishnan, 1966 syn. n. *longus Ananthakrishnan & Jagadish, 1969 syn. n. *orientalis Ananthakrishnan & Jagadish, 1969 syn. n. ligs Ananthakrishnan & Jagadish, 1971 micans Ananthakrishnan & Jagadish, 1971 pictipes (Bagnall, 1919) pusillus Ananthakrishnan & Jagadish, 1969 *tener Ananthakrishnan & Jagadish, 1969 228 B. RK PITicin KEY TO THE INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA (excluding longisetosus Ananthakrishnan (p. 252)) 1 Maxillary stylets short, when at rest not retracted far into the head capsule; maxillary bridge absent; praepectal plates weak or absent (cf. Text-figs 1-4) : — Maxillary stylets longer, when at rest retracted far into the head capsule; eaeaallaee bridge usually present; praepectal plates usually present and well developed (cf. Text-figs 9-15) 2 Body brown. On dry twigs and Aveca [Palmae] sheaths. : Antillothrips varius (p. — Body bicoloured, at least abdominal segment II yellow 3. Pronotal antero-marginal setae well reat y and expanded apically (Dest fe 4). On bamboo : ; : Antillothrips nayari (p. — Pronotal antero-marginal aoe eescel os Text-figs I, 2) 4 Pterothorax mainly yellow, light brown at anterior margin only; oe peouieel III and IV each with 1 + 1 sense cones (Text-fig. 6); micropterous. On bamboo : ; : Antillothrips micropterus (p. — Pterothorax brown; Peace senntans III and IV with 1+1 and 1-4 2+! sense cones respectively ; macropterous 5 Post-ocular and median setae (B,) on pede terpite IX porated: Pes einige with 1-4 duplicated cilia. On bamboo and grass ; Antillothrips malabaricus (p. — Post-ocular and median setae (Bj) on abdominal tergite IX expanded apically; fore wings with 5—7 duplicated cilia. On grass and bamboo Antillothrips graminellus (p. 6 Abdominal tergites III-VII without well developed Means setae (Text-fig. 16); wings usually reduced or absent : — Abdominal tergites III-VII with at least one pair, noecliye oe aes of ell developed wing-retaining setae (Text-figs 17, 18); wings alee fully eas 7 Antennal segment IV with 2 + 2*! sense cones — Antennal segment IV with at most I + 2 sense cones . ; : 4 = 8 Antennal segment III with 1 + 1 sense cones (Text-fig. 25); pronotal mid-lateral setae vestigial (Text-fig. 13). On Lantana sp. [Verbenaceae] twigs Apterygothrips rubiginosus (p. — Antennal segment III without sense cones (Text-fig. 23); pronotal mid-lateral setae well developed and expanded apically (Text-fig. 12). On dry twigs and fungus- infested branches of Flacourtia {[Flacourtiaceae] . Apterygothrips jogensis (p. 9 Body mainly brown; post-ocular setae pointed. Tube yellow with brown apex; antennal segment IV with 1 + 1 sense cones. On pine and dry twigs : : é Apterygothrips ae (p. — Body mainly yellow; post- stealer oe epomded apically ‘ 10 Tube brown; antennal segments III and IV with 1 + 1 and 1+ au sense cones respectively: pronotal antero-marginal setae well developed and expanded apically; median setae (B,) on abdominal tergite IX pointed. Macropterous or apterous. On decaying twigs : : , Apterygothrips fungosus (p. — Tube mainly yellow, extreme apex tinged with brown; antennal segments III and IV with o + 1 and 1 + 2 sense cones respectively; pronotal antero-marginal setae vestigial; median setae (B,) on abdominal tergite IX expanded apically. Apterous. On Sorghum [Gramineae] and grass . Apterygothrips pellucidus (p. 11 Abdominal tergites III-VII with one pair of well developed wing-retaining setae (Text-fig. 17). Bicoloured species. On Cynodon [Gramineae] and other grasses Haplothrips apicalis (p. — Abdominal tergites III-VII with two pairs of well developed wing-retaining setae . 237) 241) 255) I2 INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA Antennal segment IV with 2 + 2 or more Ss Se Seo pe gu eee Ne ~) ee tee ma LG Zao AS io: ee 7 ilies iD) ene = SO a INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 237 SPECIMENS STUDIED. Lectotype 2, paralectotypes 1 2, I g of varius, INDIA: Adur, Kerala on Areca [Palmae] sheaths, 28.vi.1967 (TNA) (LCM). Holotype 9, paratypes 2 9, 2 ¢ of nefrens, INDIA: U.P. Dehra Dun, on dry twigs, 12.x.1971 (TNA) (LCM; USNM). APTERYGOTHRIPS Priesner A pterygothrips Priesner, 1933a: 1. Type-species: Apterygothrips haloxyli Priesner, by original designation. Zur Strassen (1966) recognized eight species of Apterygothrips and provided a key for their identification. Of these, five occur in the Mediterranean region, one in southern Africa, one in southern India, and one apparently in the Mediterranean region and India (cf. p. 239). A further species has been described more recently from Australia (Pitkin, 1973). Four Indian species formerly assigned to Xylaplothrips are here transferred to Apterygothrips. A total of six species are recognized here from India, although the Indian record of hispanicus (Bagnall) requires confirmation. Species of Apterygothrips are associated with grasses and dead wood. GENERIC DEFINITION. Small to medium sized yellow, brown or bicoloured Haplothripini. Maxillary stylets long, when at rest retracted far into the head capsule, maxillary bridge usually present; ocelli sometimes reduced or absent; post ocular setae pointed or expanded apically. Antennal segment III with at most 1 + 1 sense cones. Pronotal antero marginal and mid lateral setae well developed or vestigial. Usually apterous and without well developed wing-retaining setae on abdominal tergites III-VII, sometimes macropterous and with two pairs of well developed wing-retaining setae on each of abdominal tergites III-VII; fore wings, if fully developed, with or without duplicated cilia. Apterygothrips fungosus (Ananthakrishnan & Jagadish) comb. n. (Text-fig. 19) Xylaplothrips fungosus Ananthakrishnan & Jagadish, 1969 : 132-133. LECTOTYPE ®Q macroptera, Inp1A: Madras (LCM), here designated [examined]. Yellow-brown species with antennal segment IV—VIII and tube except base distinctly brown. Antennal segments III and IV with 1 + 1 and 1 + 1*1 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; apterous or macropterous, fore wings when fully developed with 1-5 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from a total of six females and six males. None of these specimens was designated as the holotype although one of the female syntypes examined is labelled ‘HOLOTYPE’. This specimen, contrary to the original description, is macropterous and is here designated as the lectotype. SPECIMENS STUDIED. Lectotype 2 macroptera, paralectotypes 2 9 apterae, INDIA: Madras, decaying twigs, 25.vii.1967 (TNA) (LCM; USNM). 238 B. R. PITKIN FIGS 9-I2. Apterygothrips and Haplothrips species: heads, pronota and left forelegs. 9, H. apicalis, lectotype 9. 10, A. flavus, paratype Q (African specimen). 11, A. fungosus, paratype 9. 12, A. jogensis, 9. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 239 Apterygothrips hispanicus Bagnall Cephalothrips hispanicus Bagnall, 1916 : 409-411. Holotype 9, Spain: Zaragosa (BMNH) [examined]. Malacothrips hispanicus (Bagnall) Bagnall, 1924 : 635. A pterygothrips hispanicus (Bagnall) Priesner, 1961 : 56. Brown spécies; antennal segment III yellow; antennal segments IV—VI yellow tinged with brown; antennal segments VII and VIII pale brown; all femora brown; all tibiae brown at base becoming pale yellow at apex. Antennal segments III and IV each with 1 + 1 sense cones; post-ocular setae pointed; pronotal antero-marginal setae moderately well developed and pointed; mid-lateral setae well developed and blunt to expanded; macropterous, wings often broken; fore wings without duplicated cilia (zur Strassen, 1966); median setae (6,) on abdominal tergite IX pointed. Bagnall (1923) recorded a single male of hispanicus from the flower of Rhododendron at Kulhara, Garhwal. The references by Bagnall (1924), Priesner (1928) and Ananthakrishnan (1969) of hispanicus from India appear to refer to this specimen. As far as I am aware no other published records of hispanicus in India exist. Since it is possible that Bagnall accidentally mis-labelled his specimen, the Indian record of hispanicus requires confirmation. SPECIMENS STUDIED. Holotype 9°, paratype 2, SPAIN: Zaragosa, 8.iv.1913 (Navds) (BMNH). ?Inp1A: Kulhara, Garwal, 11,700 ft, I § on flower of Rhododendron, 5.vii.1970 (A. D. Imms) (BMNHB). Apterygothrips jogensis (Ananthakrishnan & Jagadish) comb. n. (Text-figs 12, 23) Xylaplothrips jogensis Ananthakrishnan & Jagadish, 1969 : 124-125. LECTOTYPE 9 aptera, InpIA: Jog Falls (LCM), here designated [examined]. Brown species with apex of tibiae and tarsi yellow; antennal segment III yellow at base darkening towards apex to pale brown. Antennal segments III and IV with o + 0 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial; mid-lateral setae well developed and expanded apically; apterous; median setae (B,) on abdominal tergite IX pointed or blunt. This species was originally described from four females and two males collected from fungus infested branches of Flacourtia sp. [Flacourtiaceae] at Jog Falls, India. None of these specimens was designated as the holotype. The two females and one male that I have examined are labelled ‘HOLOTYPE’, ‘PARATYPE’, and ‘ALLOTYPE’ respectively. The female labelled as the holotype is here designated lectotype. SPECIMENS STUDIED. Lectotype 2 aptera, paralectotypes 1 9 aptera, 1 g aptera, INDIA: Jog Falls, on fungus infested branches of Flacourtia sp., 22.x.1966 (INA) (LCM; USNM). 240 B. R. PITKIN Fics 13-18. 13-15. Apterygothvrips species: heads, pronota and left forelegs of (13) vubiginosus, 2; (14) pini, Q; (15) pelludicus, paratype Q. 16-18, abdominal tergite IV of (16) Apterygothrips pellucidus paratype g; (17) Haplothvips apicalis, 9; 18, H. ganglbaueri (paralectotype 9 of priesnerianus). INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 241 Apterygothrips pellucidus (Ananthakrishnan) comb. n. (Text-figs 15, 16, 21) Xylaplothrips pellucidus Ananthakrishnan, 1968 : 133. Holotype 9 aptera, Inp1a: Madras (LCM) [not examined]. Mainly yellow species with antennal segments V—VIII brown; mesonotum and extreme apex of tube tinged with brown. Antennal segments III and IV with o + 1 and 1 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and expanded apically; apterous; median setae (B,) on abdominal tergite IX expanded apically. This species was originally described from a holotype female, an allotype male, and 27 female and 17 male paratypes which are all apparently apterous. The specimens examined from Kalahasti, India and Taimergarha, Pakistan had been misidentified as flavus Faure. SPECIMENS STUDIED. Paratypes I 2 aptera, I g aptera, INDIA: Madras, on Sorghum [Gramineae] leaves, 12.xi.1965 (INA) (USNM). ?Paratypes 2 ¢ aptera, INDIA: Madras, on Sorghum leaves, 12.x1.1965 (TNA) (BMNH). Inp1A: Madras, 1 2 aptera on grass, 6.vil.1966 (TNA) (BMNH); Kalahasti, 1 9 aptera on grass, 9.ix.1964 (TNA) (BMNH). Pakistan: Taimergarha, 1 ¢ aptera on Cynodon dactylon (L.) Persoon [Gramineae] 30.1.1964 (C7 BC) (BMNH). Apterygothrips pini Ananthakrishnan (Text-figs 14, 22) A pterygothrips pint Ananthakrishnan, 1961a@ : 574-575. Holotype 9 aptera, Inp1A: Kodaikanal, Bryant’s Park (LCM) [examined]. Apterygothrips pint Ananthakrishnan; zur Strassen, 1966 : 161-175. Pale brown species with bicoloured tube and antennal segment III paler than other segments. Antennal segments III and IV with o + 1 and 1 + 1 sense cones respectively; post-ocular setae pointed; pronotal antero-marginal setae vestigial, mid-lateral setae fairly well developed but pointed; apterous; median setae (B,) on abdominal tergite IX pointed. This species was originally described from three females and two males from pine needles. I have examined further material from dry twigs. SPECIMENS STUDIED. Holotype @ aptera, allotype 3 aptera, INp1A: Kodaikanal, Bryant’s Park, on pine needles, 4.vi.1959 (INA) (LCM). Inpia: Kodaikanal, 7 9 apterae on dry twigs, 2.1.1969 (INA) (BMNH; LCM). 242 B. R. PITKIN Lo N See 5 Sf Sagar es _ ce Gee 87 Fl) + oi ee w pis is 6-2 mS, gs ee Ae Sa S J of o-s bbs AY} Qo So S x a) N 22, A. pini, holotype 9. 21, A. pellucidus, paratype Q. A pterygothrips and Haplothrips species: antennae. 19, A. fungosus, paratype Q. 25, A. rubiginosus, paratype 9. 9 (African specimen). lectotype 9. Fics 19-25. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 243 Apterygothrips rubiginosus (Ananthakrishnan & Jagadish) comb. n. (Text-figs 13, 25) Xylaplothrips rubiginosus Ananthakrishnan & Jagadish, 1971 : 263-264. Holotype @ aptera, Inp1A: Kodaikanal, Lantana sp. twigs (LCM) [examined]. Brown species with apex of tibiae yellow; antennal segment III yellow to grey-brown. Antennal segments III and IV with 1 + 1 and 2 + 2+*1 sense cones respectively; post-ocular setae pointed; pronotal antero-marginal and mid-lateral setae vestigial; apterous; median setae (B,) on abdominal tergite IX pointed. This species was originally described from a holotype female, an allotype male and six female and four male paratypes, all apparently apterous. SPECIMENS STUDIED. Holotype 2 aptera, allotype g aptera, Inp1A: Kodaikanal, Lantana sp. twigs, 1.1.1969 (INA) (LCM). ?Paratype 2 aptera, Inp1A: Kodaikanal, Lantana sp. twigs, 1.1.1969 (TNA) (LCM). CHIRAPLOTHRIPS Priesner Haplothrips (Chivaplothrips) Priesner, 1930 : 271. Type-species Haplothrips (Chivaplothrips) faureanus Priesner, by monotypy. Chiraplothrips Priesner; zur Strassen, 1960 : 347. Priesner (19645) provides a key to the three described species of Chirvaplothrips. All of these apparently have the apical margin of the fore femora ‘somewhat reflexed exteriorly’. This character is, however, difficult to observe and consequently I have avoided its use in the key above. Only one species of Chivaplothrips is recorded from India (Ananthakrishnan, 1969). GENERIC DEFINITION. Medium sized brown to dark brown species of Haplothripini. Maxillary stylets long, when at rest retracted far into the head capsule; maxillary bridge present; post-ocular setae pointed. Antennal segment III with o + 1 sense cones, segment IV with I + 1 sense cones. Pronotal antero-marginal and mid-lateral setae vestigial; praepectal plates present. Macropterous, fore wings with duplicated cilia. Abdominal tergites III-VII each with two pairs of well developed wing-retaining setae. Chiraplothrips graminellus (Priesner) (Text-figs 35, 48) Haplothrips (Chivaplothrips) graminellus Priesner, 1938 : 113-115. Syntypes 9, g, Cyprus: Cherkes, Asomotos; [LEBANON] Syria: Beirut; SupaN: Wad Shair (SMF, BMNH) [1 9, 1 3 examined}. Haplothvips (Chivaplothvips) pvriesnevi Ananthakrishnan, 1961b : 426-428. Holotype Q, Inp1A: Madras (LCM) [mot examined]. [Synonymized by zur Strassen, 1968 : 100.] Chivaplothrips gyvaminellus (Priesner) Priesner, 1964b : 434. Brown species; antennal segment III yellow; segments IV and V brownish yellow; segments VI-VIII pale brown; fore tibiae yellow, brown at base and margins; fore tarsi yellow; all femora and mid and hind tibiae brown; mid and hind tarsi pale brown. Antennal segments III and IV with o + 1 and 1 + 1 sense cones respectively; post ocular 244 B. R. PITKIN setae pointed; pronotal antero-marginal and mid-lateral setae vestigial; fore wings with about 6-7 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. Priesner (1938) described graminellus from an unspecified number of females and males collected on turf and grasses in Cyprus, Lebanon and Sudan. I have examined two syntypes from Cyprus and these are both labelled ‘Paratype’. Ananthakrishnan (1961b) described priesneri from a holotype and an allotype collected on Chloris barbata (L.) Swartz at Madras, i. 1959 (INA). I have not seen either of these specimens. Zur Strassen (1968) synonymized priesneri with graminellus. SPECIMENS STUDIED. Syntypes I 9, 1 g, Cyprus: Cherkes, turf, viii.1933 (G. A. Mavroumoustakis) (BMNH). Inp1A: Madras, I 2 on grass, II.vi.1963, I 9 on grass, I1.vii.1963; Coimbatore, 3 2 on grass, 12.vi.1963; Tinnevelley, 1 2 on grass, 27.v.1965; (INA) (BMNH). HAPLOTARIPS Amyot & Serville Haplothrips Amyot & Serville, 1843 : 640. Type-species: Phloeothrips albipennis Burmeister [= Thrips aculeata Fabricius], by monotypy. The genus Haplothrips has a world-wide distribution and comprises over 200 described species. The majority are flower-living although a number of predatory and mycophagous species have been recorded. Eleven of the 14 Indian species recognized here are flower-living. A further species originally described from galls has subsequently been recorded from flowers. The remaining two species are inquilines, one of psyllid galls and the other of galls and lepidopterous leaf-mines. GENERIC DEFINITION. Medium sized, usually brown Haplothripini, rarely yellow or bicoloured. Maxillary stylets usually long and retracted far into the head capsule; maxillary bridge usually present. Post-ocular setae pointed, blunt, or expanded. Antennal segment III with at most 1 + I sense cones; antennal segment IV with 2 + 2+}! or more rarely 2 + 2 sense cones. Pronotal antero-marginal and mid-lateral setae vestigial to well developed; praepectal plates well developed. Usually macropterous, fore wings with or without duplicated cilia. Abdominal tergites III-VII usually with two pairs, rarely one pair, of well developed wing-retaining setae. Subgenus HAPLOTHRIPS Amyot & Serville Haplothrips Amyot & Serville, 1843 : 640. The subgenus Haplothrips is used for species with duplicated cilia on the distal osterior margin of the fore wing. Haplothrips (Haplothrips) andresi Priesner (Text-fig. 38) Haplothrips andvesi Priesner, 1930 : 270-271. Syntypes 9, g, Eacypr: Mersa, Matrouh (SMF; BMNB#) [1 9 examined]. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 245 Haplothrips andresi Priesner; Ananthakrishnan & Jagadish, 1966 : 257. Dark brown to black species with fore tibiae yellow towards apex, outer margins dark; mid and hind tibiae pale yellow at apex; all tarsi yellow; antennal segment III yellow, segments IV, V and VI pale in basal two-thirds; segment VII pale in basal third or more. Antennal segments III and IV with o + 1 and 2 + 2+! sense cones respectively; post-ocular setae pointed apically; pronotal antero-marginal and mid-lateral setae well developed; fore wings with 10-12 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from an unspecified number of males and females collected in Egypt from grass and vine leaves or inside fig fruits, 12.vi.1930 (A. Andres) (Priesner, 1930). No holotype was designated although the single female syntype I have examined is labelled ‘Paratype’. There is a female labelled as a paratype in the SMF (pers. com. L. A. Mound), but the number and whereabouts of the remaining syntypes are unknown to me. Priesner (1964b) recorded andrvesi from the flowers of Thymelaea hirsuta (L.) Endlicher [Thymelaeaceae] in the north of Egypt. The species is also recorded from Palestine (Rivnay, 1933; Priesner, 1936) and southern India (Ananthakrishnan & Jagadish, 1966). Two Indian females identified by Professor Ananthakrishnan as andresi have been examined during the course of this study and these represent tenuipennis Bagnall. I have not seen Indian material of andresi SPECIMENS STUDIED. Syntype 9, Ecyrt: Mersa, Matrouh, in fig fruits, 12.vi.1930 (A. Andres) (BMNH). Haplothrips (Haplothrips) bagrolis Bhatti Haplothrips bagrolis Bhatti, 1973 : 535-537. Holotype 3, Inp1a: Himachal Pradesh, Bagrol, Kangra District (Hans Raj College, Delhi) [not examined]. Brown species; antennal segment III pale yellowish; segments IV—VI light brown; fore tibiae brown on outer margin; tarsi yellow. Antennal segments III and IV with o + 1 and 2 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae fairly well developed and expanded apically; macropterous, fore wings with 5-9 duplicated cilia. This species was described from a male holotype and an allotype and paratype female collected on wheat at Bagrol, Kangra District, 20.iii.1971 (S. K. Ahluwalia). I have not seen any of these specimens. The above diagnosis is drawn from the original description. There are three males and four associated females from Pakistan in the BMNH which could represent this species, but I have not compared them with the type- material. One male and one associated female were collected on Rubus sp. [Rubiaceae] at Murree, 5.vi.1g61; one male and three associated females were collected from Plantago [Plantaginaceae] flowers at Kwai, 14.v.1963; and one male was collected from the roots of an unidentified plant at Thapla, 15.x.1961. Apart from the bifid aedeagus of the males, these specimens are indistinguishable from ganglbaueri (cf. p. 240). 246 B. R. PITKIN Fics 26-29. Karnyothvips species: heads, pronota and left forelegs. 26, alpha, holotype Q. 27, flavipes, 9 (North American specimen). 28, melaleucus, 9. 29, nigviflavus, paratype Q. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 247 32, 35, Chivaplothrips 31, alpha, holotype 9. 34, Haplothrips bicolor, 9. 30, melaleucus, holotype &. 30-33, Karnyothrips species. 33, flavipes 2 (North American specimen). 36, Haplothrips pirus, syntype 9. Antennae. nigviflavus, paratype @. graminellus, 2. Fics 30-36. 248 Bik. PIDIRCIN Haplothrips (Haplothrips) bicolor (Ananthakrishnan) comb. n. (Text-figs 34, 37) Xylaplothvips bicoloy Ananthakrishnan, 1964a : 53-54. Syntypes 2 9, 2 g, Inp1a: Bapatla (LCM) [not examined]. Bicoloured species; head, thorax, posterior half and median third of abdominal segment VIII, abdominal segment IX and tube brown; abdominal segments II—VII yellow, segments III-VII each with a small median pale brown transverse patch near anterior margin; antennal segment I and much of segment II pale brown; antennal segments III—-VI yellow; segments VII and VIII brown; all legs yellow except outer margin of fore femora which are tinged with brown. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed; macropterous, fore wings with 5—6 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Ananthakrishnan (1964a) described bicolor from two females and two males collected on psyllid galls at Bapatla, 1952 (INA). None of these specimens was designated as holotype and I have not examined any of them. The above diagnosis is based on the original description and a single female identified by Professor Ananthakrishnan. SPECIMEN STUDIED. Inp1A: Courtallam, I 2 on grass, 10.x.1964 (INA) (LCM). Haplothrips (Haplothrips) ceylonicus Schmutz Haplothvips ceylonicus Schmutz, 1913 : 1038-1039. Syntypes 9 g, Srr Lanxa: Peradenya (presumed lost) [not examined]. Brown species; antennal segments III—VI yellow, segments VII and VIII brown; fore tibiae, apical half to one-third of mid and hind tibiae and all tarsi yellow. Antennal segments III and IV with o + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 6-9 duplicated cilia; median setae (B,) on abdominal tergite IX slightly expanded apically. Schmutz (1913) described ceylonicus from an unspecified number of females and males collected at Peradenya on Cassipourea sp. [Rhizophoraceae] and Crotalaria striata Candolle [Leguminosae] by Uzel. These are presumed lost. The above diagnosis is based on three females from India collected on Ficus [Moraceae] and identified by Dr J. S. Bhatti. Priesner (19330) redescribed ceylonicus from material collected in Java, Sumatra and Riouw Archipelago. I have examined some of these specimens, from Wai Lima, Lampongs, Sumatra. Contrary to the redescription these specimens have I + I sense cones on antennal segment III and are identical with tenuipennis Bagnall. I have also examined seven females from India (USNM) and one female from Sri Lanka (SMF) identified by Professor Ananthakrishnan and the late Professor Dr Priesner as ceylonicus. These are indistinguishable from tenuipennis. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 249 SPECIMENS STUDIED. Inv1A: Madras, 3 2 on Ficus sp., viii.1973 (J. S. Bhatti) (Hans Raj College, Delhi). Haplothrips (Haplothrips) ganglbaueri Schmutz Haplothrips Ganglbaueri Schmutz, 1913 : 1034. Syntypes 9, Sri Lanka: Peradenya (presumed lost) [not examined]. Haplothrips ceylonicus var. vernoniae Priesner, 1921 : 4. Syntypes 5 9, 1 g, Java: Semerang (4 2, 1g in SMF) [4 9, 1g examined]. Syn. n. Zygothvips andhva Ramakrishna, 1928 : 290-291. Holotype 9, Inp1A: Guntur, Madras Presidency (Tamil Nadu University College, Coimbatore) [examined]. Syn. n. Haplothrips priesnerianus Bagnall, 1933 : 327-328. Lectotype g, Inpi1a: Allahabad (BMNH), designated by Mound (1968 : 114) [examined]. Syn. n. Brown species; antennal segments I and II brown; segment III brownish yellow; segments IV, V and VI successively darker; fore tibiae yellow with brown margins. Antennal segments III and IV with o + 1 and 2 + 2*! sense cones respectively; post ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 5-8 duplicated cilia; median setae (B,) on abdominal tergite IX blunt to slightly expanded. Schmutz (1913) described ganglbaueri from an unspecified number of females collected at Peradenya, Sri Lanka on rice flowers by Uzel. These are presumed lost. Priesner (1921; 19330) redescribed ganglbaueri from material of both sexes collected in Java, Sumatra and Krakatau. This material is indistinguishable from priesnerianus. The variety vernoniae was described from five females and one male collected at Semerang, Java. I have examined four female and one male syntypes labelled ‘COTYPE’ and cannot distinguish them from the type-series of priesnerianus. Zygothrips andhra was described from a unique female holotype collected at Guntur, Madras Presidency, India. I cannot distinguish this specimen from priesnerianus or bagrolis. Dr Bhatti (pers. com.), however, considers andhra as a synonym of ganglbaueri, a view I have accepted here. Haplothrips priesnerianus was originally described from an unspecified number of females and males collected at Allahabad on flowers of Lantana [Verbenaceae], 26.ili.1910 (A. D. Imms). One of the males, which is labelled ‘TYPE’, was designated as the lectotype by Mound (1968) and is deposited in the BMNH with two females and two males regarded as paralectotypes. The above synonymy is based on comparison of the original description of ganglbaueri (Schmutz, 1913), Priesner’s redescriptions (Priesner, 1921; 19330), the lectotype and paralectotypes of priesnerianus and numerous specimens from India identified by Professor Ananthakrishnan and Dr Bhatti as ganglbaueri. Bhatti (1973) distinguished ganglbaueri from bagrolis on the metanotal sculpture, the length of the pronotal postero-angular setae and the shape of antennal segment III and the male aedeagus. Apart from the shape of the male aedeagus these characters vary from one specimen of ganglbaueri to another and are here considered unreliable. I have not found any characters which could be used to distinguish females of bagrolis and ganglbaueri. The females referred to below under 250 B. R. PITKIN ‘Specimens studied’ have been identified by association with males of ganglbaueri. The unassociated females that I have examined are listed separately. These could represent either ganglbaueri or bagrolis. SPECIMENS STUDIED. Syntypes 4 9, I g of ceylonicus var. vernoniae, JAVA: Semerang, ‘aus Bliimen und verkruppelten, teilweise gerolten Blattern von Vernonia cinerea’, 3.vii.1912 (W. Docters v. Leeuwen) (SMF). Holotype 9 of andhra, INDIA: Guntur, Madras Presidency, on cholam shoots (7. V. Ramakrishna No 152) (Tamil Nadu University College, Coimbatore). Lectotype g, paralectotypes 3 9, 2 3 of priesnerianus, Inp1A4: Allahabad, flowers of Lantana in jungle, 26.i11.1920 (A. D. Imms) (BMNH). Inp1A: Annamalais, I 9, 3 ¢ on grass, 26.ix.1965; Bangalore, 2 9, 1 g¢ on grass, 20.iv.1964; Kodaikanal, 1 g on grass, 25.v.1965; Valparai, I 9, I g on grass, 26.ix.1965 (IT NA) (USNM); Rajasthan, Udaipur, 7 9, 5 4, 4.xi.1964 (CIE) (BMNH). PAKISTAN: Muree, 31 9, 7 ¢ on Arundinella nepalensis Trinius, 16 9, 2 3 on Cymbopogon martini (Roxburgh) Watson, 5.vi.1g61 (CIE); Nowsera, I 9, I gf on Tagetes africana, 3.1.1964 (CIE); Peshawar, 9 2, 4 ¢ on Pennisetum typhoideum L. Richard, 30.1x.1964; (BMNH). BANGLADESH: Dacca, I 9, I 3 on wheat, iv.1969 (Agric. Res. Inst.); Ishardi 6 9, 1 g on Chinese paddy, 4.v.1968 (Ma Hasib Plant Quarantine St.); (BMNH). Sortomon Is.: Guadalcanal, 27 9, 9 4, on rice, xi-xii.1965 (M. McQuillan); Guadalcanal, Kukum Agric. St., 1 9, I g on grass and flowers, 21.x.1965 (P. N. Lawrence); (BMNH). Unassociated females of ganglbaueri or bagrolis. Inp1A: Rawalpindi, 9 2 on Cymbopogon distans (Nees) W. Watson, I1.vii.1960 (CIE); Indore, 1 Q on Sorghum vulgare Persoon, 1.x.1966 (Coll. Agric.); Madras, 3 2 on grass, 17.1v.1963 (INA); Mussoorie, 1 9 on fern, 13.vi.1967 (TNA); New Delhi, r 2 on grass 10.viil.1966 (TNA) (BMNH; USNM). Pakistan: Lyallpur, 4 2 on wheat, 26.v.1968 (Ayub Agric. Res. Inst.) (BMNH). BANGLADESH: Dacca, 4 @ on rice, 15.11.1966 (CIBC); Dacca, 2 2 on water hyacinth, 1961 (Govt. Entomologist) (BMNH). JAva: Buitzenzorg, Medan S.O.K., 1 2 on Imperator exaltata (Roxburgh) Brongniart (L. Fulmek) (BMNH). Soromon Is.: Guadalcanal, iI 9 on ? Brachiaria mutica (Forsskal) Stapf, 17.v.1966 (WM. McQuillan); Guadalcanal, Papananism, 1 2 Oct. 1965 (P. Greenslade) (BMNH). Purippines: Alicia Isabella, 1 Q on rice seedlings, 26.v.1972 (A. D. Pawar) Los Banos, I.R.R.I. Lagina, I 9 on rice panicles, 20.xii.1971 (A. D. Pawar) (BMNH). Haplothrips (Haplothrips) gowdeyi (Franklin) (Text-fig. 39) Anthothvips gowdeyi Franklin, 1908 : 724. Syntypes 31 9, BARBADOos (10 9 in USNM; 21 Q in Massachusetts Agricultural College) [not examined]. Haplothrips sororcula Schmutz, 1913 : 1036-1037. Syntypes 9, g, Sri Lanxa: Peradenya (1 9 in SMF; depository of remainder unknown) [1 9 examined]. Syn. n. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 251 Fics 37-40. Haplothrips species: heads, pronota and left forelegs. 37, bicolor, 9. 38, andvesi, paratype 9 (Egyptian specimen). 39, gowdeyi, 9 (African specimen). 40, reuteri, 9 (European specimen). 252 Bo, PAKUN Haplothrips soroy Schmutz, 1913 : 1036-1037. Syntypes 9, g, Sri Lanka: Peradenya (2 @ in SMF; depository of remainder unknown) [2 2 examined]. Syn. n. Haplothrips gowdeyi (Franklin) Watson, 1921 : 38. Brown species; antennal segments III—V yellow; segment IV yellow-brown; segments VII and VIII brown; mid and hind tibiae brown; all tarsi pale brown. Antennal segments III and 1V with 1 + 1 and 2 + 2 Tt! sense cones respectively ; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4-9 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. Schmutz (1913) described sovorcula from an unspecified number of females and males collected on ‘Blattern des Zimtbaumes und in Canna-Bliten’ and ‘Bliten von Crotalaria striata D.C. und in Reis’; and sovor from an unspecified number of females and males on ‘Bluten von ? ? an den Ufern des Maveli’ and ‘Blattern vom Zimtbaum’. As pointed out above, male syntypes of sororcula were collected with female syntypes of ceylonicus Schmutz; and male syntypes of sovor were collected with female syntypes of ganglbaueri Schmutz. Fortunately, however, some female syntypes of sororcula and soror, labelled as ‘Cotype’ and ‘Cotypen’ respectively, are deposited in the SMF. I have examined these and cannot distinguish them from gowdeyi. The number and whereabouts of the remaining syntypes is unknown to me. Haplothrips gowdeyi is a pantropical species breeding in the flowers of Gramineae. It may, however, be frequently collected in the flowers and on the foliage of numerous other plant families. The female examined from Madras on Amaranthus had been misidentified as vamakrishnai Karny (= nigricornis Bagnall); and the two females from Madras on grass as coloratus Trybom. SPECIMENS STUDIED. Syntypes 2 2 of sovor, Skt LANKA (Ceylon): Mahaveli (Uzel) (SMF). Syntype 2 of sororcula, SR1 LANKA (Ceylon): Peradenya, auf Zimtbaum (Uzel) (SMF). INpIA: Conjeeveram, 4 9 on grass, I.vi.1963; Kodaikanal, 1 9 on grass, I2.vi.1963; Udaipur, i 9 on wild flowers, 30.v.1966; Madras, 1 9 on Amaranthus, 22.vi.1963; Tambaram, I 9, 20.iv.1963; Pollachi, 1 9 on Eleusine, 10.vi.1963; Madras, 2 9 on grass, 18.viil.1965; (INA) (USNM). Haplothrips (Haplothrips) longisetosus Ananthakrishnan Haplothrips longisetosus Ananthakrishnan, 1955 : 610-611. Syntypes 2 9, Inp1a: Malabar (LCM) [not examined]. This species was originally described from two females collected at Malabar from wild flowers. No holotype was designated in the original description and I have not studied either specimen. According to the original description and its accompanying figure antennal segment III has four sense cones and antennal segment IV has two sense cones. If this is correct then longisetosus does not belong in Haplothrips or any of the related genera discussed in this account. However, if this statement and the figure are the result of an error, as is suggested by the INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 253 reference ‘antennae as in H. gowdeyi’, then longisetosus is similar to gowdeyi and tenuipennis. The ‘considerably longér’ major setae distinguish longisetosus from both of these species. Haplothrips (Haplothrips) pirus Bhatti (Text-fig. 36) Haplothrips pirus Bhatti, 1967 : 23. Syntypes 9, gj, Inp1aA: New Delhi (Hans Raj College, Delhi) [1 9, 1 f examined]. Yellow species with tube dark brown in slightly less than distal two-thirds; antennal segment V brown in distal half; segment VI brown in slightly less than distal two-thirds; segments VII and VIII dark brown. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed; macropterous, fore wings with 4-6 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. This species was described from an unspecified number of males and females collected from small brownish leaf-galls and from lepidopterous leaf-mines of Syzigium, 21.vi.1966 and 25.ix.1966, New Delhi (J. S. Bhatti). I have examined a female and male syntype labelled ‘PARATYPE’ and ‘ALLOTY PE’ respectively. Contrary to the original description ‘antennal seg. 2’ does not have ‘2 sense cones’; and antennal segment ‘3’ does not have ‘4 sense cones’. I assume that Bhatti confused antennal segment III and IV with II and III. In fact segment II only has the characteristic companiform sensilla (see Heming, 1975) and segment III has I + I sense cones and segment IV has 2 + 2+! sense cones. SPECIMENS STUDIED. Syntypes I 9, 1 3, INDIA: New Delhi, leaf mines of Jamun (Syzygiwm) 21.vi.1966 25.1x.1966 (J. S. Bhatti) (Hans Raj College, Delhi). Haplothrips (Haplothrips) reuteri Karny (Text-fig. 40) Anthemothvips veutevri Karny, 1907: 51. Syntypes, [?YuGosravia] ‘Osterreich-Ungarn’: Karlopago [? Karlobag] (depository unknown) [not examined]. Haplothrips veuteri (Karny) Priesner, 1921 : 14. Haplothvips reuteri (Karny); Moulton, 1929 : 4. Haplothrips tenuisetosus Bagnall, 1933 : 320-321 [examined]. [Synonymised by Priesner, 1964 : 151.] Haplothrips satanus Bagnall, 1933 : 321-323 [examined]. [Synonymised by Priesner, 1964: I51.] Brown species with antennae brown, segment III slightly paler than segment IJ; fore tibiae paler towards apex: fore tarsi yellow-brown. Antennal segments III and IV with r + 1 and 2 + 2+! sense cones respectively; post-ocular setae pointed; pronotal antero-marginal and mid-lateral setae well developed and pointed; macropterous, fore wings with 4-8 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. 254 B. R. PITKIN Karny (1907) described veuteri from an unspecified number of specimens collected at Karlopago in flowers of Knautia sp. [Dipsaceae] 26.vii.1905. The original description does not make it clear whether these specimens included females as well as males. The above diagnosis is based on the examination of the holotype females of satanus Bagnall and tenwisetosus Bagnall, synonymised with veuteri by Priesner (1964). The only published record of veutert in India appears to be that of Moulton (1929). I have examined 14 of the 20 males and females on which this record is based. SPECIMENS STUDIED. Holotype 2 of satanus, FRANCE: Perpignan, on Centaurea solstitialis L., viii.1926 (R. S. Bagnall) (BMNH). Holotype @ of tenuwisetosus, SUDAN: Wad Medani, Dolichos lablab L., 7.1v.1932 (A. P. G. Michelmore) (BMNH). InpiA: Dehra Dun, 9 9, 5 d (J. C. M. Gardner) (CAS). Haplothrips (Haplothrips) tenuipennis Bagnall (Text-fig. 42) Haplothrips tenuipennis Bagnall, 1918: 210. Lectotype g, Inp1a: Darjeeling District, Rington (BMNH), designated by Mound (1968 : 118) [examined]. Haplothrips ceylonicus var. mangiferae Priesner, 1933) : 359. Syntypes 9g, Java: Semarang (SMF) [not examined]. [Synonymized by Mound, 1968 : 118.]} Brown species with antennal segments III-VI yellow, segments IV—VI slightly tinged with brown at apex; segment II paler towards apex; segments VII and VIII pale brown; fore tibiae and all tarsi yellow. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 6-10 duplicated cilia; median setae (B,) on abdominal tergite IX blunt to slightly expanded. Bagnall (1918) described tenucpennis from an unspecified number of males and females collected on rose and tea bushes. There are a total of eleven females and six males in the BMNH all bearing type data. Of these one female and one male are labelled ‘Type’ and one female and three males are labelled ‘Cotype’. The remaining nine females and two males are not labelled as type-material but are here assumed to be some of Bagnall’s syntypes. The male labelled “Type’ was designated as the lectotype by Mound (1968), the remaining 11 females and five males are regarded here as paralectotypes. ‘Haplothrips ceylonicus var. mangiferae was originally described from ‘929d, Semarang, Java; 25-VIII-1912, in Bluten von Mangifera indica L., leg Docters van Leeuwen.’ Some of these syntypes are labelled as cotypes and Mound (1968) refers to these incorrectly as paratypes. All of the specimens in the USNM of tenuipennis had been misidentified by Professor Ananthakrishnan as follows: from Bhopal on Polygonum, Madras on Zizyphus, and Trichur and Tambaram on grass as ceylonicus; from Madras on INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 255 grass as ganglbauert and coloratus; from Valparai on tea as andrvesi; and from Poona on Mango as mangiferae. SPECIMENS STUDIED. Lectotype 3, paralectotypes 11 9, 5 g, Inp1A: Darjeeling District, Rington T. E. on rose and tea bushes, 14.v1.1916 (E. A. Andrews) (BMNH). Inp1A: Bhopal, 1 Q on Polygonum infl. 18.x.1966; Madras, I @ on grass, 17.iv.1963; Madras, 4 2 on Zizyphus fl., 12.vi.1963; Poona, 7 9, 1 g on Mango infl., 23.1.1965; Trichur, 1 2 on grass, 6.x.1963; Tamboram, I 9 on grass, 20.iv.1963; Valparai, 2 9 on tea fl., 26.ix.1965; (INA) (USNM); Indore, 7 9, 2 § on Mangifera indica L., i 9, I g on Phaseolus lunatus L., 1 9, 1 g on Dolichos lablab L. fl. 14.11.1967 (Coll. Agric.); Assam, Darjeeling, 8 9, 1 g on tea shoots, 1965 (Toklai Expt. St.); Udaipur, 1 2 on flowering plant, 11.v.1966 (Udaipur Univ.); (BMNH). Subgenus TRYBOMIELLA Bagnall Trybomiella Bagnall, 1926 : 548. Type-species: Haplothrips bagnalli Trybom, by original designation. The subgenus Tvybomiella is used for species of Haplothrips without duplicated cilia on the distal posterior margin of the fore wing. Haplothrips (Trybomiella) apicalis Bagnall (Text-figs 9, 17, 24) Hindsiana apicalis Bagnall, 1915 : 323. Lectotype 9 microptera, Inp1a: Almora, Kumaon, (BMNH), designated by Mound (1968 : 109) [examined]. Haplothrips (Hindsiana) apicalis (Bagnall) Priesner, 19336 : 361-363. Haplothrips (Trybomiella) apicalis (Bagnall); Ananthakrishnan, 1962 : 473-475. Haplothrips (Trybomiella) apicalis (Bagnall); Ananthakrishnan, 1969 : 138. Bicoloured species; head, antennal segments I, II and VI-VIII, pterothorax, anterior third of abdominal segments II-VII, abdominal segments VIII—X, and mid and hind tibiae brown; prothorax, fore legs and mid and hind femora yellow-brown; remainder yellow. Antennal segments III and IV with o + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; micropterous, brachypterous, or macropterous, fore wings, even when fully developed, without duplicated cilia; median setae (B,) on abdominal tergite IX pointed. Bagnall (1915) described apicalis from ‘several specimens swept from a jungle plant’ at Almora, India. The micropterous lectotype female and a second micropterous female, assumed to be a paralectotype, are in the BMNH. The number and whereabouts of the remaining specimens of the type-series are unknown to me. Unlike other species of the genus, apicalis has only one pair of well developed wing-retaining setae on each of abdominal tergites III-VII. The fore wings, even when fully developed, lack duplicated cilia, as do species of the subgenus Trybomiella. However, apicalis is not closely related to these species and is being 256 Bok. PIlkKin Fics 41-44. Haplothrips species: heads, pronota and left forelegs. 41, ganglbauert (lectotype Q of priesnerianus). 42, tenuipennis, lectotype 9. (African specimen). 44, euphorbiae, paratype 9. 43, articulosus, 9 INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 257 transferred elsewhere to a new genus by Dr Bhatti (pers. comm.). The reduction of the wing-retaining setae and absence of duplicated cilia may be associated with the grass-living habit. This species is recorded from Java (Priesner, 19330) and India (Ananthakrishnan, 1969). I have studied specimens from India and Pakistan. SPECIMENS STUDIED. Lectotype 2 microptera, paralectotype 2 microptera, Inp1a: Almora, Kumaon, on a jungle plant, 4.vii.1g11 (O. Paiva) (BMNH). Inp1A: Madras, 2 2 micropterae, 2 2 macropterae (wings lost), 3 g micropterae on Cynodon [Gramineae], 21.v.1963 (INA) (BMNH; USNM); Poona, 1 2 microptera, 2 ¢ macropterae on grass, 23.1.1965 (JNA) (USNM); Surat, 1 g macroptera on grass, 28.x.1965 (INA) (USNM); 7 @ micropterae, no further data, Bagnall Reg. 163 (BMNH). Pakistan: Taimergarha, 1 9 microptera, 30.1.1964 (CIBC) (BMNH). Haplothrips (Trybomiella) articulosus Bagnall (Text-fig. 43) Anthothrips bagnalli var. pallicornis Trybom, 1911: 10. Syntypes 9, g, Kenya: Mombasa (Riksmuseum, Stockholm) [not examined]. Haplothrips articulosus Bagnall, 1926 : 548-549. Holotype 9, Tanzanra: Morogoro (BMNH) [examined]. Haplothrips trybomianus Priesner, 1927 : 70. Replacement name for bagnalli var. pallicornis Trybom. [Synonymized by Mound, 1968 : 109.] Haplothrips (Trybomiella) derisoy Priesner, 1935 : 324. Holotype 9, SIERRA LEONE [examined]. [Synonymized by Mound, 1968 : 109.] Brown species; antennal segments III-VI brownish yellow, segments VII—-VIII pale brown; apex of fore tibiae and fore tarsi brownish yellow; mid and hind tibiae brown; mid and hind tarsi pale brown. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae slightly expanded apically; pronotal antero-marginal and mid-lateral setae well developed and slightly expanded apically; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX slightly expanded apically. Bagnall (1926) described articulosus from an unspecified number of specimens of both sexes. There are a total of two females and five males in the BMNH including the holotype female and these are labelled either ‘TYPE’ or ‘COTYPE’. Ananthakrishnan (1969) lists articulosus as occurring in India although I know of no published records of this species outside the African continent. There are, however, two females in the USNM from Hyderabad, India which I cannot distinguish from articulosus. SPECIMENS STUDIED. Holotype 2, paratypes 1 9, 5 ¢ of articulosus, TANZANIA: Morogoro, sunflower blossom, 9.i.1925 (A. H. Ritchie) (BMNH). Holotype 9, paratypes 5 9, 1 ¢ of derisor, SIERRA LEONE: Evigeron sumatrensis Retzius leaves (E. Hargreaves) (BMNRH). Inp1a: Hyderabad, 2 9 on grass, 25.vi.1964 (TNA) (USNM). 258 B. R. PITKIN Haplothrips (Trybomiella) bagnalli (Trybom) Anthothrips bagnalli Trybom, 1910 : 65. Syntypes 374 9, 105g, SouTH AFRICA; SouTH WEST AFrica; Botswana (SMF; ?Riksmuseum, Stockholm) [13 2, 9g ?Syntypes examined]. Haplothrips bagnalli (Trybom) Karny, 1912 : 325. Haplothrips bagnalli (Trybom); Faure, 1955 : 219. Brown species; antennal segment III slightly paler than segment II; fore tibiae yellow brown medially and towards apex; fore tarsi greyish yellow; mid and hind tibiae and tarsi brown. Antennal segments III and IV with 1 + 1 and 2 + 2+1 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae vestigial; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX blunt or slightly expanded. Ananthakrishnan (1969) lists bagnalli as occurring in India although I know of no published records outside southern Africa. I have not seen any specimens of bagnalli from India. Four Indian specimens identified as bagnalli by Professor Ananthakrishnan have been examined. These specimens, from Tambaram on grass, could not be distinguished from the closely related species, nigricornis Bagnall. SPECIMENS STUDIED. ?Syntypes 13 2, 9 dg, SouTH AFRICA (L. Schultze) (SMF). Haplothrips (Trybomiella) clarisetis Priesner Haplothrips clarisetis Priesner, 1930 : 237-238. Syntypes 4 9, SoutTH Arrica & EGyPT: (SMF) [not examined]. Haplothrips clarisetis Priesner; Faure, 1955 : 223-230. Brown species; antennal segment III yellow-brown, paler than segment II; segment IV slightly darker than III, nearly as dark as V; segments V—VIII brown; apex of fore tibiae and tarsi brownish yellow; mid and hind tarsi pale brown. Antennal segments III and IV with 1 + 1 and 2 + 271 sense cones respectively; post-ocular setae slightly expanded apically; pronotal antero-marginal setae well developed and slightly expanded apically, mid-lateral setae vestigial; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from one female collected at Rosslyn, Pretoria, on cabbage, 25.iii.1g1g (J. C. Faure); one female at White River, Transvaal on lemons, 27.viii.1926; and two females at Orman Gardens, Cairo on Chenopodium sp. (H. Priesner). I have not seen any of these specimens, The above diagnosis is based on the excellent redescription given by Faure (1955) and material in the BMNH from South Africa, Angola, Egypt, Israel and Palestine. At present I am unable to distinguish clavisetis from the Australian species robustus Bagnall. The Indian material in the USNM that I have examined had been misidentified by Professor Ananthakrishnan as follows: from Bareilly on Celosia, and Madras and Tambaram on grass as migvicornis; and from Baroda and Madras on Amaranthus, and Coonoor on bamboo as vamakrishnat. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 259 SPECIMENS STUDIED. Inp1A: Bareilly, 1 2 on Celosia, 3.viii.1966; Baroda, 4 9, on Amaranthus, 26.x.1965; Coonoor, 1 2 on bamboo sheath, 8.vi.1963; Madras, 2 9, 1 g¢ on Amaranthus, 22.1v.1963; 2 9 on grass, 22.1.1964; Metupaleyam, I 9, I 3, on Celosia, 23.11.1966; Tambaram, 1 9, 8.vili.1966; (TNA) (USNM). Haplothrips (Trybomiella) euphorbiae Priesner (Text-fig. 44) Haplothrips (?) euphorbiae Priesner, 1931 : 1-4. Holotype 9, Sumatra: Medan (SMF) [examined]. Haplothrips euphorbiae Priesner; Priesner, 19330 : 347. Brown species; apex of fore tibiae and fore tarsi greyish yellow. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae reduced; pronotal antero-marginal and mid-lateral setae vestigial; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Priesner (1931) described euwphorbiae from an unspecified number of females and males, including a type female, collected on galls on Euphorbia hirta at Medan, Sumatra. I have examined the holotype and three paratype females plus further material from Sumatra and India. The three Indian specimens from Waltair on grass in the USNM had been misidentified by Professor Ananthakrishnan as talpa Priesner. SPECIMENS STUDIED. Holotype 9, paratypes 3 9, SUMATRA: Medan, on Euphorbia hirta L., v.1931 (L. Fulmek) (SMF). SumaTRA: Medan, S. O. K., 2 2 on Euphorbia hirta L., xii.1923 (L. Fulmek) (BMNH). Inp1a: Anand, 4 9, 1 g on Euphorbia fis, 27.x.1965; Waltair, 3 2 on grass, 20.1x.1964; (TNA) (USNM). Haplothrips (Trybomiella) nigricornis (Bagnall) Anthothrips nigricornis Bagnall, 1910 : 425-426. Syntypes 12 9, 3 g, SoutTH Arrica: Cape Town (BMNH) [examined]. Haplothrips vamakrishnai Karny, 1926: 218. Holotype 9, Inpra: Coimbatore (SMF) {examined]. Syn. n. Haplothrips nigricornis (Bagnall); Faure, 1955 : 208-218. Brown species; antennal segment III slightly paler than segment IV, segments IV—VIII slightly paler than segments I and II; apex of fore tibiae and fore tarsi yellow-brown. Antennal segments III and IV with 1 + 1 and 2 + 2*! sense cones respectively; post ocular setae slightly expanded apically; pronotal antero-marginal and mid-lateral setae vestigial; macropterous fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX pointed or blunt. Bagnall described nigricornis from ‘numerous specimens’ of both sexes collected in South Africa on flowers of Diplopappus, Europs, Olipterus and Sebaea. There 260 B. R. PITKIN are twelve females and three males in the BMNH mounted on two slides. One slide is labelled ‘TYPES’ and the other ‘COTYPES’. The only other data on the slides are ‘S. AFRICA/R. S. BAGNALL’ and ‘TUBULIFERA/Haplothrips nigricornis Bagn./299g’ and ‘Bagnall Coll./B.M. Reg. No. 1932-339’. Karny (1926) described vamakrishnai from a unique female collected inside Chrysanthemum flowers, 13.vili.1918 (Ramakrishna). I have compared this and four other Indian specimens with the syntypes of nigricornis and further material from South Africa, Angola, Rhodesia, Tanzania, Uganda and Ethiopia in the BMNH. The Indian specimens had been misidentified by Professor Ananthakrish- nan as bagnalli. SPECIMENS STUDIED. Syntypes 12 9, 3 ¢ of mgricornis, SouTH AFRICA (BMNH). Holotype @ of vamakrishnat, INDIA: Coimbatore, Chrysanthemum flowers, 3.v1i.1918 (Ramakrishna) (SMF). Inp1A: Tambaram, 4 9 on grass, 29.1v.1965 (INA) (USNM). Haplothrips (Trybomiella) talpa Priesner Haplothrips talpa Priesner, 1930 : 243. Syntypes9, gf, SoutH AFrica: Transvaal, Potgietersrust (depository unknown) [not examined]. Body brown; antennal segment III yellow-brown, segments IV—VI slightly darker than III but paler than VII and VIII; fore tarsi and sometimes apex of fore tibiae yellow-brown. Antennal segments III and IV with o + 1 and 2 + 27! sense cones respectively; post-ocular setae short and pointed; pronotal antero-marginal and mid-lateral setae vestigial; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX pointed or blunt. The syntypes of talpa were collected at Potgietersrust, South Africa (Faure, 1955) and not Rustenburg (Priesner, 1930). I do not know their whereabouts; they are not in either Priesner’s collection (SMF) or Faure’s collection (National Insect Collection, Pretoria). The Indian specimens examined agree with Faure’s (loc. cit.) excellent redescription. SPECIMENS STUDIED. InpIA: Waltair, 3 9, 3 g, on grass, 27.x.1965 (INA) (USNM). Haplothrips (Trybomiella) tirumalraoi Ramakrishna & Margabandhu Haplothrips (Trybomiella) tivumalvaoi Ramakrishna & Margabandhu, 1931 : 1038. Holotype 6, Inp1a: Northern Circars (presumed lost) [not examined]. Ramakrishna & Margabandhu (1931) described tivumalraoi from a unique male collected on Colocasia (V. Tirumal Rao). This specimen is apparently lost. According to the original description antennal segment III bears one sense cone, the fore wings lack duplicated cilia and the post-ocular setae are blunt. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 261 KARNYOTHRIPS Watson Karynia [sic] Watson, 1922:6. Type-species: Karynia weigeli Watson [= Anthothrips flavipes Jones; synonymized by Hood, 1927: 175]. ([Misspelling for Karnyia; preoccupied by Karnyia Shelford, 1909.] Karnyothrips Watson, 1924 : 23. [Replacement name for Karnyia Watson.] The genus Karynothrips is used here for a group of Haplothrips-like species with forwardly directed fore tarsal claws, 1 + 2*1! or fewer sense cones on antennal segment IV and two pairs of well developed wing-retaining setae on each of abdominal tergites III-VII. In addition the post-ocular setae are expanded apically and the pronotal antero-marginal setae are vestigial. Only two species, flavipes Jones and migriflavus Ramakrishna, are listed by Ananthakrishnan (1969) as occurring in India. However, the so called colour forms of nigriflavus referred to by Ananthakrishnan (1965) are recognized as three distinct species, nigriflavus, melaleucus Bagnall and alpha sp.n. A further species, mucidus (Ananthakrishnan & Jagadish) is here transferred to Karnyothrips from Xvylaplothrips. Of the five species recognized here from India at least two are predatory. One of these and two other species are bicoloured and are associated with grasses. GENERIC DEFINITION. Medium sized brown or bicoloured Haplothripini. Maxillary stylets long and retracted far into the head capsule; maxillary bridge present; post-ocular setae expanded apically. Antennal segment III with o + 1 or rt + 1 sense cones; segment IV with 1 + 2+! or fewer sense cones. Fore tarsi with a forwardly directed claw. Pronotal antero- marginal setae vestigial; mid-lateral setae well developed and expanded apically. Abdominal tergites III-VII each with two pairs of well developed wing-retaining setae. Macropterous, fore wings with or without duplicated cilia. Karnyothrips alpha sp. n. (Text-figs 26, 31) Ananthakrishnan (19650) listed five colour forms of nigriflavus as follows: ‘a) Head alone brown, tube brown, rest yellow b) Head, prothorax, tube brown c) Head, prothorax, very little of mesothorax, abdominal segment IX and tube brown d) Head, prothorax, pterothorax, abdominal segments VIII to X brown e) Almost uniformly brownish individuals.’ I have examined 24 9 and 11 4g identified by Professor Ananthakrishnan as nigriflavus and of these 6 9 and 7 g fall into group a; 10 2 and 3 g fall into group b and 8 9 fall into group d. The remaining 1 ¢ does not fit into any group but comes closest to group e. The group b specimens are identical to the 13 9 paratypes of nigriflavus that I have examined and have 0 + I and I + 2 sense cones on antennal segments III and IV respectively and the median setae (B,) on abdominal tergite IX are expanded apically. However, the group d specimens examined are identical with the holotype of melaleucus Bagnall and have 1 + 1 and 1 + 1*? sense cones 262 Be Re earn! on antennal segments III and IV respectively and the median setae (B,) on abdominal tergite IX are pointed. Moreover the group a specimens examined differ from both migriflavus and melaleucus and have 0 + I and I + 2 sense cones on antennal segments III and IV respectively and the median setae (B,) on abdominal tergite IX are pointed. In addition, none of these group a specimens has duplicated cilia on the distal posterior margin of the fore wing. All specimens of nigriflavus and melaleucus that I have examined have at least one duplicated cilium on one fore wing or the other. The group a specimens are therefore recognized as a distinct species and described here as alpha sp. n. The single anomalous male near group e has 0 + E and 1 + 2*? sense cones on antennal segments III and IV respectively and the median setae (B,) on abdominal tergite IX are expanded apically, as in mucidus. It differs from mucidus in colour, i.e. antennal segment III is yellow not brown, and in my opinion probably represents a further new species. Since this specimen is unique I have not described it as a new species. A total of 6 9 and 7 ¢ of alpha, 22 9 and 2 J of nigriflavus and 38 92 of melaleucus have been examined. Within each species the number of sense cones on antennal segments III and IV, the form of the median setae (B,) on abdominal tergite IX, the range of the number of duplicated cilia on the fore wing and the colour pattern are consistent. Bicoloured species of Kaynyothrips; head, tube, sometimes disal portion of abdominal segment IX, antennal segments I, VII and VIII brown; antennal segment VI usually pale brown in distal half; remainder yellow. Antennal segments III and IV with o + 1 and 1 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and expanded apically; macropterous, fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX pointed. Measurements of holotype female (smallest-largest holotype female) in mm: total length 2280 (2016-2280); head length 222 (204-222); pronotal length 159 (150-159); tube length 112 (109-117); tube width at base 61 (57-61). Measurements of smallest-largest paratype males in mm: total length 1296-1680; head length 174-192; pronotal length 111-132; tube length 96—108; tube width at base 43-50. Holotype 9, Inp1A: Palghat, on bamboo, 28.ix.1965 (TNA) (BMNH). Paratypes. INpIA: 2 9, 2 ¢ collected with holotype; Calicut, 1 2, 1 fg on bamboo, 4.x.1963; Calicut, 1 9 on bamboo, 15.x.1964; Chalakudi, 1 g on bamboo, 5.x1.1963: Perintalmanna, 1 2 on bamboo, 6.x.1963; Omalur, I g on sugar cane, 18.vi.1966; Madras, I fon grass, 25.1.1966; Yercaud, I J on bamboo, 19.vi.1966; (TNA) (BMNH; LCM; USNM). Karnyothrips flavipes (Jones) (Text-figs 27, 33) Anthothrips flavipes Jones, 1912 : 18-19. Holotype 9, U.S.A.: California, San Jose (USNM) {not examined]. Karnyothrips flavipes (Jones) Hood, 1927 : 175. Karnyothrips flavipes (Jones); Ananthakrishnan, 1969 : 139. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 263 Brown species; antennal segment III yellow at base, pale brown at apex, segment IV sometimes pale in basal third; femora brown; tibiae yellow-brown to yellow at apex; all tarsi yellow. Antennal segments III and IV with 1 + 1 and 1 + 2+! sense cones respectively; post ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and expanded apically; macropterous, fore wings with 1-5 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. This species is cosmopolitan (Stannard, 1957; as jonesianus) and predates Coccidae, Aleyrodidae and also Acarina on a wide variety of plants (Priesner, 19646). Ananthakrishnan (1969) recorded flavipes from India. I have not seen any Indian specimens of this species. The BMNH collection contains material from Egypt, Cyprus and U.S.A. (California and Hawaii). Karnyothrips melaleucus (Bagnall) (Text-figs 28, 30) Hindsiana melaleuca Bagnall, 1911 : 61-62. Holotype 9, DENMARK: Copenhagen (BMNH) [examined]. Karnyothrips melaleuca (Bagnall) Hood, 1927 : 176. Bicoloured species; head, thorax, abdominal segments VIII or IX—X brown; pelta and abdominal segments II-VII or VIII yellow, segments III-VII each with a small median transverse pale brown patch near anterior margin; antennal segment I pale brown, segments IL-V yellow, VI yellow-brown, VII and VIII brown; fore femora brown in basal half fading to yellow towards apex; mid and hind femora and all tibiae and tarsi yellow. Antennal segments III and IV with r + 1 and r + 1+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial; mid-lateral setae expanded apically; wings fully developed, fore wings with o—4 duplicated cilia (always at least 1 on one fore wing or the other); median setae (B,) on abdominal tergite IX pointed or blunt. The Indian specimens examined had been misidentified as Karnyothrips nigriflavus by Professor Ananthakrishnan (see discussion under alpha). This species is a pantropical thrips living amongst grasses, sometimes in association with scale insects. The BMNH contains material from Denmark (in a palm house), Madeira, Kenya, U.S.A. (Hawaii), Trinidad, Brazil, Australia, West Malaysia, Java, Vietnam, China and India. Unlike alpha and nigriflavus the males of melaleucus are unknown. SPECIMENS STUDIED. Holotype 9, DENMARK: Copenhagen, on cruciferous flowers in palm house, 30.vi.1g09 (R. S. Bagnall) (BMNH). Inpi1A: Palghat, 2 9, 28.ix.1965 (BMNH); Kulatupuzha, 2 9 on wild grass, 2.ix.1966 (BMNH); Thimbam, 2 2 on bamboo, 4.ix.1967 (I 9 BMNH; 1 2 USNM); Mundumalai, t 2 on bamboo, 24.xii.19g66 (USNM); Chalakudi, 1 2 on bamboo, 5-xi.1963 (USNM); Calicut, 1 2 on bamboo, 4.x.1963 (USNM); all collected by TNA. Cuina: Canton, 1 2 on Bambusa, iii.1938 (BMNH). VIETNAM: I 9 on Vanda, 29.xii.1962 (H. A. Woolford) (USNM). 264 B. R. PITKIN Karnyothrips mucidus (Ananthakrishnan & Jagadish) comb. n. Xylaplothrips mucidus Ananthakrishnan & Jagadish, 1971 : 260-261. Holotype 9, Inp1a: Vyithri (Wynad) (LCM) [examined]. Bicoloured species; head, thorax, posterior half of abdominal segment IX, and tube brown, rest of body yellow; fore femora brown with yellow apex; fore tibiae yellow-brown; fore tarsi yellow; mid and hind legs yellow; antennal segments I brown, segment II mainly yellow, segment III yellow, segments IV—VIII brown. Antennal segments III and IV with o + 1 and 1 + 2*! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial; mid-lateral setae well developed and expanded apically; macropterous, fore wings with about 4 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. This species was originally described from two females and one male collected from dry twigs at Vyithri (Wynad). SPECIMENS STUDIED. Holotype @, allotype 3, paratype 9, Inp1A: Vyithri (Wynad), on dry twigs, 15.vill.1969 (INA) (LCM). Karnyothrips nigriflavus Ramakrishna (Text-figs 29, 32) Karnyothrips nigviflavuus Ramakrishna, 1934 : 496. Holotype 9, Inp1a: Coimbatore (LCM) [not examined]. Bicoloured species; head, prothorax and tube brown; base and apex of antennae pale brown; remainder yellow. Antennal segments III and IV with o + 1 and 1 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae. well developed; macropterous, fore wings with 1-5 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Ramakrishna (1934) described nigriflavus from ‘numerous specimens (T.V.R. No. 26ga)’ collected on bamboo in Coimbatore. He referred to a ‘type female’ but made no mention of any males. Professor Ananthakrishnan kindly loaned a slide of nigriflavus from Ramakrishna’s collection (T.V.R. 269). This unfortunately was broken in transit, although none of the specimens were damaged. The slide originally held 13 females of migriflavus and a single specimen of an unidentified Xylaplothrips sp. I have remounted all of these singly on slides. I regard the colour forms of nigriflavus referred to by Ananthakrishnan (1965a) as at least three distinct species (see discussion under alpha). SPECIMENS STUDIED. Paratypes 13 9, Inp1A: Coimbatore, on bamboo (T. V. Ramakrishna 269) (LCM; BMNH). Inp1A: Palghat, 3 9, 1 ¢ on bamboo, 4.ix.1967 (BMNH); Perintalmanna, I 9 on grass, 5.xi.1963 (BMNH); Aryankavu, I 9, 21.ii.1966 (USNM); Yercaud, 1 2 on INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 265 bamboo, 19.vi.1966 (USNM); Tumkur, 1 2 on bamboo, 5.ix.1967 (USNM); Dindigul, I 9 on bamboo, 28.vii.1967 (USNM); Calicut, 1 9, 1 g on bamboo, 4.x.1963 (USNM); Chalakudi, 1 9 on bamboo, 5.xi.1963 (USNM); all collected by TNA. PRAEPODOTHORIPS Priesner & Seshadri Praepodothrips Priesner & Seshadri, 1952 : 407-410. Type-species: Pyaepodothrips indicus Priesner & Seshadri, 1952 : 408-410, by original designation. Praepodothrips is used here for a group of four large grass-inhabiting haplothripines. Three of these have relatively large heads and small pronota and I + I sense cones on antennal segment IV. The fourth species has the head only about as long as the pronotum and 2 + 2+! sense cones on antennal segment IV. This last-mentioned species possibly warrants a genus of its own but is retained for the present in Praepodothrips. GENERIC DEFINITION. Large brown or bicoloured species of Haplothripini. Head usually relatively large; maxillary stylets retracted far into the head capsule; maxillary bridge present. Antennal segment III with o + 1 to 1 + 2 sense cones; segments IV with 1 + 1 or 2+ 271 sense cones; post ocular setae pointed to expanded. Pronotal antero-marginal and mid lateral setae vestigial. Macropterous, forewings with or without duplicated cilia. Abdominal tergites III-VII each with two pairs of well developed wing-retaining setae. Praepodothrips cymbapogoni Ananthakrishnan (Text-figs 46, 50) Praepodothrips cymbapogoni Ananthakrishnan, 1956 : 136-138. Holotype 9, Inp1A: Simuralai Hills (LCM) [not examined]. Praepodothrips cymbapogont Ananthakrishnan; Ananthakrishnan, 1960 : 574. Brown species; apical half to third of antennal segment II, segments III-VI yellow, remainder of antennae pale brown; apex of fore femora, all tibiae and tarsi yellow. Antennal segments III and IV each with 1 + 1 sense cones; post-ocular setae pointed; pronotal antero-marginal setae apparently vestigial; macropterous, fore wings with 5-7 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from two females collected on Cymbapogon citvatus (Candolle) Stapf from Sirumalai Hills, Madras Province ({NA) (Anan- thakrishnan, 1956). The only specimen examined for the present study, which was identified by Professor Ananthakrishnan, is unfortunately mounted ventral side up and the pronotal setae are not readily discernible. SPECIMEN STUDIED. Inp1A: Coonoor, I 2 on Cymbapogon citratus (Condolle) Stapf, iv.1958 (TNA) (BMNH). 266 B. R. PITKIN Fics 45-48. Heads, pronota and left forelegs of Praepodothrips and Chtvaplothrips species. 45, P. indicus, paratype 9. 46, P. cymbopogoni, 9. 47, P. nigrocephalus, holotype 9. 48, C. graminellus, paratype 9. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 267 Praepodothrips indicus Priesner & Seshadri (Text-figs 45, 49) Praepodothrips indicus Priesner & Seshadri, 1952 : 408-410. Syntypes 9, gj, Inpra: Valparai (Tamil Nadu University, Coimbatore; Zoological Survey, Calcutta; SMF) [2 9 examined]. Bicoloured species; head, thorax, abdominal segments IX—X brown; abdominal segment VIII yellow in anterior third, brown in posterior two-thirds, segments II-VII yellow, segments III-VI tinged brown medially, segment VII also tinged brown in posterior fourth; antennal segments I, basal half and interior distal portion of segment II, segment VIII brown; all legs yellow, margins of fore femora slightly tinged with brown. Antennal segments III and IV each with 1 + 1 sense cones; post-ocular setae pointed, rarely blunt; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and pointed; macropterous, fore wings with 4—8 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from ‘several individuals of both sexes’ from Valparai, south India, collected on an unknown host plant ‘but most likely Gramineae sp’. The two syntypes examined are both labelled ‘Paratype’. SPECIMENS STUDIED. Syntypes 2 9, Inp1A: Valparai, ix.1945 (E. R. G. Menon) (SMF). Inp1A: Pachmarhi, 5 9 on grass, 21.xi.1966; Valparai, I 9, 2 g on lemon grass, 26.ix.1965; Ooty, 1 2 on Imperata grass, 25.11.1966; Yercaud, 1 g, on Imperata grass, 17.vi.1966; Salem, I 9, I g on grass, 19.vii.1963; (INA) (BMNH). Praepodothrips nigrocephalus Ananthakrishnan (Text-figs 47, 51) Praepodothvips nigrocephalus Ananthakrishnan, 1964b : 228. LECTOTYPE Q, Inpzia: Kodaikanal Hills (LCM), here designated [examined]. Bicoloured species; mainly yellow; head, posterior third of abdominal segment IX, tube antennal segment I and most of segment II and segment VIII brown; remainder yellow. Antennal segments III and IV with o + 1 and 1 + 1 sense cones respectively; post-ocular setae pointed; pronotal antero-marginal setae vestigial, mid-lateral setae vestigial; macropterous fore wings without duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from ‘18 females and 3 males’ from grasses, Kodaikanal Hills, India (Ananthakrishnan, 1964b). The female and male syntype examined are labelled ‘Holotype’ and ‘Allotype’ respectively. The female is here designated as the lectotype, the remaining 17 females and 3 males being regarded as paralectotypes. SPECIMENS STUDIED. Lectotype 9, paralectotype g, Inp1A: Kodaikanal Hills, 9.xii.1963 (JNA) (LCM). 268 By KK. PIDIREN Praepodothrips priesneri Ananthakrishnan (Text-fig 52) Praepodothrips priesneyvi Ananthakrishnan, 1955 : 608-612. Holotype 9, Inp1A: S., Madras (LCM) [examined]. Dark brown species; antennal segment III—VII almost entirely yellow; apex of fore femora yellow; fore tibiae yellow at base and apex, tinged brown medially on margins; mid and hind tibiae yellow at base; all tarsi yellow. Antennal segments III and IV with 1 + 2 and 2 + 2+! sense cones respectively ; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae vestigial; macropterous, fore wings with 6-9 duplicated cilia; median setae (B,) on abdominal tergite IX pointed or blunt. Although pviesneri is a fairly large grass-inhabiting haplothripine, as are the other species treated here under Praepodothrips, its head is only about as long as the pronotum and there are 2 + 2+1 sense cones on antennal segment IV. SPECIMENS STUDIED. Holotype 9, allotype 3g, InprA: S., Madras, bamboo sheaths, 12.vi.1954 (TNA) (LCM). XYLAPLOTHRIPS Priesner Haplothrips (Xylaplothrips) Priesner, 1928 :572. Type-species: Haplothrips fuliginosus (Schille), by monotypy. Xylaplothrips Priesner; Priesner, 1964a : 171. The genus Xylaplothrips was erected as a subgenus of Haplothrips by Priesner (1928). He distinguished Xylaplothrips from Haplothrips sensu stricto by the symmetrical antennal segment III bearing two sense cones and also by the fungus- feeding habit. Priesner (1964a) elevated Xylaplothrips to generic rank. Subse- quently Xylaplothrips has been used by authors for fungus-feeding Haplothrips-like species regardless of the number of sense cones on antennal segment III. The 10 species recorded from India and discussed here under Xylaplothrips are probably polyphyletic and moreover none of them are strictly congeneric with fuliginosus, which is European. GENERIC DEFINITION. Small to large species of Haplothripini. Maxillary stylets long, retracted far into the head capsule, maxillary bridge present; post-ocular setae’ expanded apically. Antennal segment III usually with 1 + 2 sense cones, sometimes fewer; segment IV usually with 2 + 2+! sense cones, sometimes fewer. Pronotal antero-marginal setae usually well developed, sometimes vestigial; mid-lateral setae well developed and usually expanded apically. Usually macropterous, rarely apterous; fore wings, when present and fully developed, with duplicated cilia. Abdominal tergites III-VII each with two pairs of well developed wing-retaining setae. Xylaplothrips debilis Ananthakrishnan & Jagadish (Text-fig 56) Xylaplothrips debilis Ananthakrishnan & Jagadish, 1971 : 266-267. Holotype 9, INDIA: Kulatupuzha (Kerala) (LCM) [examined]. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 269 Brownish species; head, thorax and tube brown, rest of body brownish yellow; antero-median regions of abdominal segments IJ—VIII with brownish patches; legs yellow except median portions of femora which are shaded with brown; antennal segments I and V—VIII brown, segment II brown in basal half, yellowish apically, segment III yellow in basal half, brown apically. Antennal segments III and IV each with 1 + 1 sense cones; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4—6 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was described from two females collected at Kulatupuzha and two females collected at Aryankavu. One of the female paratypes from Aryankavu has been examined for the present study but this differs from the holotype female from Kulatupuzha in having 1 + 2 and 2 + 2 sense cones on antennal segments III and IV respectively and cannot be distinguished from pusillus. The holotype female of debilis is very similar to pusillus and it is possible that this female is merely an aberrant specimen of pusillus. The female of debilis in the USNM that I have examined had been misidentified by Professor Ananthakrishnan as tener. This specimen has identical data to a female of Jigs in the USNM and it may be that the two were collected together. SPECIMENS STUDIED. Holotype 9, InprA: Kulatupuzha (Kerala) dry twigs, 7.x.1969 (I.NA); paratype 9, Inp1A: Aryankavu (Kerala) dry twigs, 12.xi.1969 (7 NA) (USNM). Inp1A: Tirupathi, 1 2 on dry twig, 31.vii.1g69 (INA) (USNM). Xylaplothrips emineus Ananthakrishnan & Jagadish (Text-fig. 55) Xylaplothrips emineus Ananthakrishnan & Jagadish, 1971 : 264. Holotype 9, Inpia: Tirupathi, dry twigs, 16.vii.1969 (INA) (LCM). [1 9, 1¢ ?paratypes examined]. Bicoloured species; head, lateral margins of pterothorax, anterior margin of abdominal segment II and tube brown; abdominal tergites III—VIII yellow with median transverse patches near anterior margins; abdominal tergite IX yellow-brown; antennae except segment III brown; segment III yellow. Antennal segments III and IV with 1 + 2 and 2 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4—6 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Neither of the specimens examined for the present study are labelled as type-material although they bear identical data to some of the paratypes. This species was originally described from five females and three males collected at Tirupathi, Courtallam, and Aryankavu on dry twigs. I have examined a specimen in the USNM from Tirupathi identified as tener which I cannot distinguish from debilis. 270 Bree een SPECIMENS STUDIED. ?Paratypes 1 9, 1 4g, InpiA: Aryankavu, Tenmalai (Kerala), dry twigs, 12. xi.1969 (INA) (LCM). Xylaplothrips flavitibia Ananthakrishnan & Jagadish (Text-fig. 62) Xylaplothrips flavitibia Ananthakrishnan & Jagadish, 1969 : 128-129. LECTOTYPEQ, Inpia: Courtallam (INA) (LCM), here designated [examined]. Brown species; antennal segment III yellow, segments IV—VI yellow in basal two-thirds, tinged with brown in apical third, remaining segments brown; fore femora brown with yellow apex; mid and hind femora brown; all tibiae and tarsi yellow. Antennal segments III and IV with r + 2 and 2 + 27! sense cones respectively ; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4-8 duplicated cilia; median setae (B,) on abdominal tergite IX very slightly expanded. Ananthakrishnan & Jagadish (1969) described flavitibia from three females and four males from Courtallam and Pachmarhi but did not designate any of these specimens as the holotype. The female and male syntypes from Courtallam I have examined are labelled ‘HOLOTYPE’ and ‘ALLOTYPE’ respectively. The female labelled ‘HOLOTYPE’ is here designated as the lectotype. The two specimens I have examined from Pachmarhi have identical data to some of the syntypes but are not labelled as type-material. SPECIMENS STUDIED. Lectotype 9, Inpra: Courtallam, wild galls, 20.vii.1963 (TNA) (LCM). ?Paralectotypes I 9, I g, Inp1A: Pachmarhi, wild galls, 21.x1.1966 (TNA) (USNM). Inp1A: Allahabad, 2 @ galling Acacia leucophloeae Wildenow, reared 1.xii.1966 (B. D. Agarwal) (USNM). Xylaplothrips flavus Ananthakrishnan (Text-fig. 54) Xylaplothvips flavus Ananthakrishnan, 1964a : 51-52. Syntypes 2 9, 4 9, INpra: Sirumalai Hills (LCM) [1 9 examined]. Brown species; antennal segment III yellow, segments IV and V yellow at base, pale brown at apex, segment VI pale brown, yellow at extreme base; all femora brown, yellow brown at apex; all tibiae yellow tinged with brown medially; all tarsi yellow. Antennal segments III and IV with 1 + 2 and 2 + 2*! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4-5 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. This species was originally described from two females and four males collected at Sirumalai Hills in palm inflorescences. None of these specimens was designated as the holotype although the single female syntype examined is labelled ‘Paratype’. ~~ Ww ee SS SS INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA Co) ar 60 Bice) 55, X. 50, P. cymbapogoni, 9. 57, X. pusillus (paratype 9 of debilis), segments III and IV. 54, X. flavus, syntype 9. 49, P. indicus, paratype 9. 53, X. inquilinus, lectotype Q. 52, P. priesneri, holotype 9. Antennae of Pyraepodothrips and Xylaplothrips species. nigvocephalus, holotype 9. 56, X. debilis, holotype 9, segments III and IV. 58, X. ligs, holotype 9, segments III and IV. Fies 49-58. emineus, &. 272 B. Re PITKIN The only other specimen I have examined has identical data but is not labelled as a type. SPECIMENS STUDIED. Syntype 9, ?syntype g, Inp1A: S. Sirumalai Hills, inflorescence of wild palm, 1959 (I.NA) (LCM). Xylaplothrips inquilinus (Priesner) comb. n. (Text-fig. 59) Haplothvips inquilinus Priesner, 1921 : 4. LECTOTYPE 9, Java: Semarang (SMF), here designated [examined]. Xylaplothrips inquilinus Ananthakrishnan, 1966: 13. Syntypes 9 9, 3 dg, INnp1a: Perintal- manna (Kerala); Alargarkoil (Madura); and Araku Valley (Andrha) (LCM) [2 9 ?syntypes examined]. Syn. n. Xylaplothrips longus Ananthakrishnan & Jagadish), 1969: 129. LECTOTYPE Q, Inpia: Tirupathi, here designated (LCM) [examined]. Syn. n. Xylaplothrips orientais Ananthakrishnan & Jagadish, 1969 : 129. Syntypes 4 9, 3g, InpDIa: Tirupathi; Tambaram (LCM) [not examined]. Syn. n. Brown species; antennal segments III-VI usually yellow in basal half, pale brown in distal half, rarely almost completely yellow; segments I-II, VII-VIII brown; mid and hind femora brown; mid tibiae yellow at base and in apical half; hind tibiae yellow at base and apex. Antennal segments III and IV with 1 + 2 and 2 + 27+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed; macropterous, fore wings with 5-8 duplicated cilia; median setae (B,) on abdominal tergite IX pointed or blunt. Priesner (1921) described inquilinus from two females and four males collected at Semarang, Java in galls on Ficus benjamina L., 9.111.1912 and Smilax sp., 5.1.1912 (Docters v. Leeuwen). I have-examined all of these specimens which are mounted on two slides. One male and one female are mounted on a single slide with a specimen of ‘Andothrips melastomae’. This female is here designated as the lectotype. I have examined females of ‘inguilinus Anan. sp.n./T. N. ANANTHAKRISHNAN’ from Alargarkoil and Perintalmanna which bear identical data to some of the syntypes of inquilinus Ananthakrishnan. Although these are not labelled as such, I suspect they are syntypes. Xylaplothrips longus was described from two females collected at Tirupathi. Neither of these specimens was designated as the holotype although the female syntype examined is labelled ‘HOLOTYPE’. This female is here designated as the lectotype. The above synonymies are based on examination of the lectotype and paralectotypes of inquilinus Priesner, two ? syntypes of inqguilinus Ananthakrishnan, the lectotype of longus and a specimen of orientalis identified by Professor Ananthakrishnan. In addition to this material I have seen a female of inquilinus Ananthakrishnan identified by Professor Ananthakrishnan. This specimen has identical data to the lectotype and paralectotype of longus and the female and male syntypes of orientalis from Tirupathi. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 273 SPECIMENS STUDIED. Lectotype 9, paralectotype ¢ of inquilinus Priesner, JAVA: Semarang, ‘in Gallen auf Ficus benjamina L.’, 9.iii.19g12 (Docters v. Leewwen) (SMF); paralectotypes 3 2, I 3 of inquilinus Priesner, JAVA: Semarang, ‘in Ballgallen auf Smilax spec.’ 5.1.1912 (Docters v. Leewwen) (SMF). Lectotype 2 of longus, Inp1A: Tirupathi, Pavetta sp. 10.viii.1965 (TNA) (LCM). ?Syntypes 2 of inquilinus Ananthakrishnan, INpIA: I 2 Perintalmanna, Pavetta galls with Teucho. longus, 5.xi.1963 (INA) (LCM); 1 9 Alagarkoil, Pavetta galls with Teucho, 10.1x.1964 (INA) (USNM). Inp1a: Tirupathi, 1 9 on Pavetta sp. galls, 10.vili.1g65 (JNA) (LCM). Xylaplothrips ligs Ananthakrishnan & Jagadish (Text-fig. 58) Xylaplothrips ligs Ananthakrishnan & Jagadish, 1971 : 261-263. Holotype 9, Inpia: Tirumalai, Tirupathi (LCM) [examined]. Brown species; tarsi paler; antennal segment III yellow with yellowish brown apex. Antennal segments III and IV with o + 1 and 1 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 5-6 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. I have examined the holotype female and allotype male of Jigs and a second female with identical data to the paratype female. The male differs from both the females in having I + I sense cones on each of antennal segments III and IV as in debilis. I have suggested above that debilis may be an aberrant form of pusillus and it is possible that ligs is likewise an aberrant form. I have examined material of pusillus and debilis with identical data and material of pusillus and ligs with identical data. SPECIMENS STUDIED. Holotype 9, Inp1a: Tirumalai, Tirupathi, on dry twigs, 19.vil.1g69 (INA) (LCM). InpiA: Tirumalai, Tirupathi, 1 2 on dry twigs, 31.vii.1969 (TNA) (USNM). Xylaplothrips micans Ananthakrishnan & Jagadish Xylaplothrips micans Ananthakrishnan & Jagadish, 1971 : 265-266. Holotype 9, Inp1a: Aryankavu (LCM) [examined]. Mainly yellow species; apex of head, mesothorax, posterior portion of pelta and anterior half of abdominal segment II and tube brown; antennal segments I-III yellow, remaining segments yellow; all legs yellow. Antennal segments III and IV with 1 + 1 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal setae vestigial, mid-lateral setae well developed and expanded apically; macropterous or apterous; fore wings when fully developed with 6—7 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Ananthakrishnan & Jagadish (1971) described micans from a total of three 274 B. R. PITKIN females and three males collected on dry and decaying twigs and bark at Aryankavu, Kodaikanal and Madras. The original description refers only to macropterous females and males. Although the holotype female is macropterous, the female paratype that I have examined is apparently apterous. SPECIMENS STUDIED. Holotype 2 macroptera, Inpra: Aryankavu, dry twigs, 12.xi.1969 (TNA) (LCM); paratype 9 aptera, Kodaikanal, Lantana twigs, 31.xi1.1968 (TNA) (LCM). Xylaplothrips pictipes (Bagnall) (Text-fig. 61) Haplothvips pictipes Bagnall, 1919 : 273-274. Lectotype 9, Inp1A: Talimparamta Malabar (BMNH), designated by Mound (1968 : 139) [examined]. Neoheegeria pictipes (Bagnall) Mound, 1968 : 139. Xylaplothrips pictipes (Bagnall) Ananthakrishnan, 1969 : 122. Brown species; intermediate antennal segments paler; apex of fore femora yellow, tinged with brown medially, yellow at base; tarsi yellow. Antennal segments III and IV with 1 + 2 and 2 + 2+! sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 5-7 duplicated cilia; median setae (B,) on abdominal tergite [X pointed or blunt. The two female specimens in the USNM that I have examined had been misidentified as ‘Xylaplothrips mimus Pr.’ This species differs from pictipes in having I + 1 and 1 + 2 sense cones on antennal segments III and IV respectively. SPECIMENS STUDIED. Lectotype 9, paralectotypes 2 9, Inp1A: Malabar, on diseased pepper berries, ix.1918 (Ramakrishna) (BMNH). Inp1A: Hubli, 2 9 on grass, 17.xi.1964 (INA) (USNM). Xylaplothrips pusillus Ananthakrishnan & Jagadish (Text-fig. 57) Xylaplothrips pusillus Ananthakrishnan & Jagadish, 1969 : 125-126. Syntypes 1 9, 1 ¢, Inpia: Kulatupuzha (LCM) [not examined]. Yellow-brown species; head, thorax and tube brown; abdominalsegments II-IX yellow-brown, segments II-VIII with a brown median transverse patch, segment IX darker posteriorly; antennal segment I brown, segments II and III yellow brown, segment III with extreme apex brown, segments IV—VIII brown, darker than head; femora brownish yellow; all tibiae and tarsi yellow. Antennal segments III and IV with 1 + 2 and 2 + 2 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae well developed and expanded apically; macropterous, fore wings with 4-5 duplicated cilia; median setae (B,) on abdominal tergite IX pointed. INDIAN SPECIES OF HAPLOTHRIPS AND RELATED GENERA 275 Fics 59-62. Xylaplothrips species: heads, pronota and left forelegs. 59, inquilinus, 9. 60, pusillus, 9. 61, pictipes, 2. 62, flavitibia, 9. 276 B. R. PITKIN The above morphological diagnosis is based on a single female from Tirupathi identified by Professor Ananthakrishnan as pusillus. I am unable to distinguish this from the paratype females of debilis from Aryankavu that I have examined. As stated above I suspect that both debilis and ligs are forms of pusillus. SPECIMENS STUDIED. Invi: Tirupathi, 1 9 on dry Smilax twigs, 21.vi.1970 (TNA) (LCM); paratype 9 of debilis, Aryankavu, on dry twigs, 12.vi.1968 (INA) (LCM). Xylaplothrips tener Ananthakrishnan & Jagadish Xylaplothrips tener Ananthakrishnan, 1969 : 126-128. LECTOTYPE 9, Inp1a: Goa-Londa Border (LCM), here designated [examined]. Brown species; antennal segment III yellow in basal half, brownish yellow in apical half; tibiae yellow sometimes tinged with brown medially; all tarsi yellow. Antennal segments III and IV with 1 + 2 and 2 + 271 sense cones respectively; post-ocular setae expanded apically; pronotal antero-marginal and mid-lateral setae expanded apically; macropterous, fore wings with 4—6 duplicated cilia; median setae (B,) on abdominal tergite IX expanded apically. Ananthakrishnan & Jagadish (1969) described tener from two females and two males collected at Goa-Londa Border but did not designate any of these specimens as the holotype. I have examined a syntype female labelled ‘HOLOTYPE’ which is here designated as the lectotype. I have also examined a male specimen labelled ‘ALLOTY PE’ but this is apparently from ‘Kulatupuzha, on Areca sheath, 25.1.1967 (INA)’. These data, however, have been crossed out. The only other specimen identified by Professor Ananthakrishnan as tener that I have seen is in fact debilis. SPECIMENS STUDIED. Lectotype 9, INDIA: Goa-Londa Border, 12.x.1965 (INA) (LCM); paralectotype d, ? collected with lectotype. REFERENCES Amyot, C. J. B. & AUDINET-SERVILLE, J.G. 1843. Histoive naturelle des insectes. Hémiptéres. Ixxvi + 675 pp. Paris. ANANTHAKRISHNAN, T. N. 1953. Notes on some Thysanoptera from South India. Ind. J. Ent. 14 (Dec. 1952) : 197-201. 1955. Studies on some Indian Thysanoptera—1. Ann. Mag. nat. Hist. (12) 8 (Aug. 1955) : 608-612. 1956. Studies on some Indian Thysanoptera III. Zool. Anz. 157 (Oct. 1956) : 130-139. 1957. A study on the forms of Haplothrips (Trybomiella) apicalis Priesner. Proc. R. ent. Soc. Lond (B) 26 (9-10) (Oct. 1957) : 143-148. 1958. 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INDEX Synonyms are in ?falics. alpha, 261 emineus, 269 andhra, 249 euphorbiae, 259 andresi, 244 Antillothrips, 231 flavitibia, Haplothrips, 225 apicalis, 255 flavitibia, Xylaplothrips, 270 Apterygothrips, 237 flavipes, 262 ; articulosus, 257 flavus, Apterygothrips, 225 flavus, Xylaplothrips, 270 bagnalli, 258 fungosus, 237 bagrolis, 245 bicolor, 248 ganglbaueri, 249 gowdeyi, 250 graminellus, Antillothrips, 231 pyenicus, 248 graminellus, Chiraplothrips, 243 Chiraplothrips, 243 clarisetis, 2 58 Haplothrips, 244 cymbapogoni, 265 hispanicus, 239 debilis, 268 indicus, 267 devisoy, 257 inquilinus, 272 280 jogensis, 239 Karnyothrips, 261 ligs, 273 longisetosus, 252 longus ,272 luteus, 232 mangiferae, 254 malabaricus, 232 melaleucus, 263 micans, 273 micropterus, 234 mimus, 226 mucidus, 264 nayari, 235 nefrens, 235 nigricornis, 259 nigriflavus, 264 nigrocephalus, 267 orientalis, 272 pallicornis, 257 pellucidus, 241 pictipes, 274 pini, 241 B. R. PITKIN Department of Entomology BritisH Museum (NaTuRAL History) CROMWELL RoapD Lonpon SW7 5BD B. R. PITKIN pirus, 253 Praepodothrips, 265 priesneri, Chiraplothrips, 243 priesneri, Praepodothrips, 268 priesnerianus, 249 pusillus, 274 vamakrishnai, 259 reuteri, 253 rubiginosus, 243 satanus, 253 SOYOY, 252 sovorcula, 250 talpa, 260 tener, 276 tenuipennis, 254 tenuisetosus, 253 tirumalraoi, 260 trybomianus, 257 Trybomiella, 255 varius, 235 vernoniae, 249 Xenothrips, 231 Xylaplothrips, 268 *. ‘ oe | Se 1, os ma spe — "el y hes A a A d fj u . at a My a > a “ ‘ ‘ v , ite ‘ i oe “. a" ? i) Mes as 7 hy , . + sf ae er ee n i] i} the ip : Sukh ‘i i oy) 4 “ ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp.172:500text-figures. September, L005, £3-25. . Oxapda, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50. . HemmineG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounp, L.A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . AFiFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. {5. . CRoSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Exiot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. {12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. {9.90. . CRosskEy, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . VON HavEK,C.M.F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae. Pp. 309: 17 text-figures. October, 1973. £12.30. . CRossKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND GU22 9LH THE ANT TRIBE "5/5 tae oy (HYMENOPTERA : FORMICIDAE) CONSTITUENT GENERA, REVIEW OF SMALLER GENERA AND REVISION OF TRIGLYPHOTHRIX FOREL B. BOLTON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 5 LONDON : 1976 i i | THE ANT TRIBE TETRAMORIINI (HYMENOPTERA : FORMICIDAE). CONSTITUENT GENERA, REVIEW OF SMALLER GENERA AND REVISION OF TRIGLYPHOTHRIX FOREL BY BARRY BOLTON Pp. 281-379; 73 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 34 No. 5 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), wistituted im 1949, 1s issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Paris will appear at wregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Depariment. This paper is Vol. 34 No. 5 of the Entomology serves. The abbreviated titles of perrodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) ISSN 0524-6431 © Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 28 October 1976 Price £10.20 This issue completes volume 34 THE “ANT TRIBE TETRAMORIINI (HYMENOPTERA : FORMICIDAE) CONSTITUENT GENERA, REVIEW OF SMALLER GENERA AND REVISION OF TRIGLYPHOTHRIX FOREL By B. BOLTON CONTENTS Page SYNOPSIS : : : : : é : : : : : 284 INTRODUCTION. : : < : : : : c : 284 MUSEUMS AND INSTITUTIONS ; : : é : : : < 285 MEASUREMENTS AND INDICES. - : : : 286 GENERIC COMPOSITION AND ORIGINS OF > TETRAMORIINI : : 2 ; 286 TRIBE TETRAMORIINI WHEELER. : P : 3 : ‘ 289 GENERA NEWLY EXCLUDED FROM TETRAMORIINI : : : : - 291 SPECIES NEWLY EXCLUDED FROM TETRAMORIINI . : . : : 293 KEYS TO GENERA OF TETRAMORIINI . é : : - : - 294 Anergates Forel E : : ; : : - : - ; 296 Decamorium Forel . ; ; : : : ; : : : 297 Synonymic list of species : : : : : : : : 298 Summary of the species. : A ; : é : : . 298 Rhoptromyrmex Mayr : : : : : ; : ; : 298 Synonymic list of species : : : : : : : ; 300 Key to species (workers) : : : : : : : : 301 Summary of the species. : ; : : : - ; : 301 Strongylognathus Mayr : : : é : : : - : 304 Synonymic list of species é : é c : : : : 305 Summary of the species. 2 : - - - : . : 305 Teleutomyrmex Kutter : : : : : , : : : 309 Triglyphothrix Forel . ‘ ; - . : : : : 310 Species of the Ethiopian seston : : : : ; : . 312 Synonymic list of species. . ‘ ; : : ; ; é 312 The species-groups . : : : - : , : 5 313 Key to species (workers) : . : ‘ ; : : : 314 Treatment by species . ‘ i 317 Species of the Oriental, Indo- Lawes sy Bolpnesena aon 8) ‘ 339 Synonymic list of species. , ‘ : : ‘ : ; 339 The species-groups . : 5 : : : é - : 340 Key to species (workers) . : : : - : : : 341 Treatment by species . : c : : : 3 : - 344 Tetvamovium Mayr . . s . i 5 F . 359 Generic synonymy of Tetramorium : F ; : 2 : , 360 Xiphomyrmex . : < ; ; f : c : é 360 Atopula . : : : . : ; : : : : 362 Macromischoides : - : : é 363 The subgenera Lobomyrmex and Sipomatiies ; ; : : 364 ACKNOWLEDGEMENTS E : : F ; : : F ; 365 REFERENCES . : ¢ : ; : ; é : : : 365 INDEX . é 5 5 é ‘ , ‘ F ; 5 : Sy) 284 B. BOLTON SYNOPSIS The generic composition of the myrmicine ant tribe Tetramoriini is revised and the tribe . re-defined. Seven genera are recognized as valid, namely Anergates Forel, Decamorium Forel, Rhoptromyrmex Mayr, Strongylognathus Mayr, Teleutomyrmex Kutter, Tetramorium Mayr and Triglyphothrix Forel. The genus Iveneella Donisthorpe is synonymized with Rhoptromyrmex. The genera Macromischoides Wheeler and Xiphomyrmex Forel, and the subgenera Lobomyrmex Kratochvil and Sulcomyrmex Kratochvil are synonymized with Tetvamorium. The genera Eutetramorium Emery, Iveneopone Donisthorpe and Tetvamyrma Forel are excluded from the tribe. Keys to genera are given. Tetvamorium is defined and discussed as a preliminary to a world revision. The smaller genera are reviewed and a new species of Rhoptromyrmex described. Tvriglyphothrix is formally revised, 55 species being recognized as valid, of which 21 arenew. Keys to species are provided for Decamorium, Rhoptromyrmex and Triglyphothrix. The species Tetramorium opacum (F. Smith) is transferred to genus Romblonella Wheeler (tribe Meranoplini), and Xiphomyrmex flavigastey Clark is transferred to genus Chelaney Emery (tribe Solenopsidini). Genus Atopula is transferred from tribe Myrmecinini and synonymized with Tetramorium. Of the former species of Atopula, belti (Forel) is transferred to Brunella Forel, longispina Stitz is transferred to Pavatopula Wheeler, and hortensis Bernard is synonymized with Tetramorium seviceiventve Emery. INTRODUCTION THIS paper is the first part of a projected revision of the ant tribe Tetramoriin and presents a review, with keys where possible, of all the smaller genera of the tribe and also gives a formal revision of the moderately sized genus Triglyphothrix. The genus Tetramorium is very large, larger than the rest of the tribe together, and will be dealt with separately in one or more papers which are now in prepara- tion. The present paper only gives a definition and discussion of the genus and its synonyms. The course of the study at the generic level has led to a number of interesting discoveries which make it possible to define the tribe accurately, but unfortunately it has also led to the conclusion that a number of quite well-known and old generic names have no validity. In this categoty come Xiphomyrmex and Macromischoides, both now relegated to the synonymy of Tetramorium. Other casualties include the spurious subgenera Lobomyrmex and Sulcomyrmex of Tetramorium, which are synonymized, and the monotypic Iveneella, based on a male, is sunk under Rhopiro- myrmex. Three small and very poorly known genera which were formerly placed in the tribe, namely Eutetramorium, Ireneopone and Tetramyrma, are excluded from further consideration of the Tetramoriini and are assigned to different tribes elsewhere in the subfamily Myrmicinae. In distribution most of the genera of the tribe Tetramoriini are primarily of the Old World tropics and subtropics, particularly the Ethiopian, Malagasy and Indo- Australian regions. A few Holarctic species of Tetramorium are known but only a single species is unquestionably endemic in the New World, Tetramorium spinosum (Pergande), which is found in the more arid regions of the southern U.S.A. and Mexico. The Palaearctic region has three endemic genera, Anergates, Teleuto- myrmex and Strongylognathus. The first two of these are monotypic genera and are permanent social parasites without a worker caste. The last named is a dulotic genus with falcate mandibles. All the species of these genera utilize species of the ANT TRIBE TETRAMORIINI 285 Tetramorium caespitum-group as hosts. The small genus Decamorium is restricted to the Ethiopian region but the larger genera Rhoptromyrmex and Triglyphothrix are found throughout the Old World tropics although no endemic species of these are known from either Madagascar or Australia. Finally, the huge genus Tetra- morium is based upon the Ethiopian region but is also very strongly represented throughout the Old World tropics and subtropics. One group of this genus, the T. caespitum-group, has colonized the more temperate and Mediterranean zones of the Palaearctic region and numerous Tetrvamorium species are widely distributed across the Pacific island systems. The tribe also contains some of the world’s more successful tramp species, spreading mainly or solely by means of human commerce. Such species are found in Trighyphothrix (Tr. lanuginosa, Tr. kheperra) and Tetramorium (T. simillimum, T. guineense, T. pacificwm) and are more or less commonly encountered in the temperate zones of the world where they are capable of surviving in hothouses or permanently heated buildings, and over wide areas of the world tropics where they are quite at home in nests away from human habitation or in places where the ground has been disturbed by agriculture. In the free-living genera the vast majority of species are terrestrial or sub- terranean, with very few arboreal forms. Ground nesting species characteristically nest directly in the earth, in rotten twigs, in or under rotten logs or in compressed leaf litter, and foraging is normally carried on at ground level. A few species which nest in the ground are not averse to foraging on the stems and leaves of plants or even on trees. Of the arboreal species the most common nest sites include rot-holes or rotten areas beneath the bark of otherwise healthy trees, but a few species build rough fibrous nests under leaves or in branch forks, such nests being constructed of vegetable fibres, small twigs and pieces of leaf. Colony size is usually moderate, with a few hundred workers, but in some Rhoptromyrmex the nests may be enormous. Most tetramoriines are generalized predators or scayengers but a few are more specialized and tend homopterous insects or visit plant nectaries. A few species have very specialized diets, for example all members of the Tetramorium sericeiventre- group are suspected of feeding entirely on other ants, particularly Phezdole species. Previous studies of the Tetramoriini as a tribe are confined to a few works on a regional basis, such as those of Forel (1902) and Bingham (1903) of the Indian fauna, Arnold (1917; 1926) of the South African fauna, Creighton (1950) of the North American fauna and Mann (1919; 1921) of the species of the Solomon and Fiji Islands. Apart from these there have been a number of reviews or revisions of some of the smaller genera, and these are noted under the discussions of the genera in question. MUSEUMS AND INSTITUTIONS ANIC, Canberra Australian National Insect Collection, Canberra City, Australia. BMNH British Museum (Natural History), London, U.K. CAS, San Francisco California Academy of Sciences, San Francisco, California, U.S.A. IE, Bologna Istituto di Entomolgia del’Universita, Bologna, Italy. MCSN, Genoa Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy. 286 MCSN, Verona MCZ, Cambridge MHN, CdF MHN, Geneva MNHN, Paris MNHU, Berlin MRAC, Tervuren NM, Basle NM, Bulawayo NM, Hamburg NM, Vienna UM, Oxford ZM, Munich Total Length (TL). B. BOLTON Museo Civico di Storia Naturale, Verona, Italy. Museum of Comparative Zoology, Cambridge, Mass., U.S.A. Muséum d’Histoire Naturelle, La Chaux-de-Fonds, Switzerland. Muséum d’Histoire Naturelle, Geneva, Switzerland. Muséum National d’Histoire Naturelle, Paris, France. Museum fiir Naturkunde der Humboldt-Universitat, Berlin, Germany (D.D.R.). Musée Royal de l'Afrique Centrale, Tervuren, Belgium. Naturhistorisches Museum, Basle, Switzerland. National Museum, Bulawayo, Rhodesia. Naturhistorisches Museum, Hamburg, Germany. Naturhistorisches Museum, Vienna, Austria. University Museum, Oxford, U.K. Zoologisches Sammlung des Bayerischen Staates, Munich, Germany. MEASUREMENTS AND INDICES The total outstretched length of the individual, from the mandibular apex to the gastral apex. Head Length (HL). The length of the head proper, measured in a straight line from the anterior clypeal margin to the mid-point of the occipital margin, in full-face view. Head Width (HW). The maximum width of the head behind the eyes, measured in full-face view. Cephalic Index (CI). Scape Length (SL). HW x 1r00 IBUE, The straight-line length of the antennal scape excluding the basal constriction or neck. Scape Index (SI). SLX shoe HW Pronotal Width (PW). The maximum width of the pronotum in dorsal view. Alitrunk Length (AL). The diagonal length of the alitrunk in lateral view from the point at which the pronotum meets the cervical shield to the base of the metapleural lobes or teeth. All measurements are expressed in millimetres. GENERIC COMPOSITION AND ORIGINS OF TETRAMORIINI The earliest attempts at defining a tribe Tetramoriini were quite vague as to its limits and a number of unrelated genera which were excluded later were originally grouped together. ‘The situation as it had been reached in the first decade of this century was summarized by Wheeler (1910) under a tribal name of Tetramorii. Beside the presently-included genera this agglomeration also contained the genera Mayriella Forel, Calyptomyrmex Emery, Meranoplus F. Smith, Wasmannia Forel and Ochetomyrmex Mayr. ANT TRIBE TETRAMORIINI 287 A serious attempt to define the tribe more accurately was made by Emery (1914a) in a paper which gave a preview of his later massive works in the Genera Insectorwm series. He presented a key to the myrmicine tribes as he defined them, and further restricted the Tetramoriini by removing disparate genera such as those noted above. A number of other genera were listed as tetramoriine by Forel (1917) and the genera thus included were more or less stabilized in the dual classifications of Emery and Wheeler, both of which appeared in 1922. The only difference between these two systems as regards the tetramoriines lay in the treatment of Tetrvamorium aculeatum, which was treated under Tetramorium by Emery but which had been removed by Wheeler (1920) to a separate genus, Macromischoides. Later Wheeler (1922) defined this genus and placed it in tribe Leptothoracini where it has been retained by most authors (myself included; Bolton, 1973) despite the proof by Santschi (1924) that Macromischoides belongs in the Tetramoriini. As the Emery-Wheeler classification was emerging, Forel (1922) described Tetra- morium subgenus Cephalomorium, the single species of which was shown later by Santschi (1925) to be a member of genus Pheidole. Despite these works the limits of tribe Tetramoriini were still rather vague, and subsequent to the Emery-Wheeler classification various authors added genera and subgenera to the tribe until by 1950 (after the addition of the last generic name but before any critical studies were undertaken) the generic composition of the tribe was as follows. Decamorium Forel Strongylognathus Mayr Dyomorium Donisthorpe = Myrmus Schenk Eutetvamorium Emery Teleutomyrmex Kutter Iveneella Donisthorpe Tetvamorium Mayr Iveneopone Donisthorpe =Tetrogmus Roger Lundella Emery T. subgen. Lobomyymex Kratochvil Macromischoides Wheeler T. subgen. Sulcomyrmex Kratochvil Rhoptromyrmex Mayr Tetvamyrma Forel R. subgen. Acidomyrmex Emery Triglyphothnix Forel Xiphomyrmex Forel Since 1950 a number of modifications to this system have been suggested. Brown (1953) showed that Lwundella was synonymous with Hylomyrma (tribe Myrmicini) and thus removed the Neotropical region’s only claim to endemic tetramoriines (see also Kempf, 1973) and later Brown (1964) synonymized Acido- myrmex to Rhoptromyrmex. Ettershank (1966) showed that Dyomorium was a synonym of Vollenhovia and thus outside the Tetramoriini and he also pointed out that Anergates should be included in the tribe and not treated as a member of Solenopsidini or of a subtribe thereof, as had previously been the case (Emery, 19144; 1922; Forel, 1917; Wheeler, 1922). At generic level the results of the present study have confirmed the above suggestions and shown that Eutetramorium, Ireneopone and Tetramyrma are not tetramoriines; that [veneella is a synonym of Rhoptromyrmex and that Xiphomyrmex, Macromischoides, Lobomyrmex, Sulcomyrmex and Atopula (from outside the tribe) 288 B. BOLTON are all synonyms of Tetramorium. These results are discussed in detail elsewhere in this study. Thus the generic composition of the tribe is now as follows. Anergates Forel Tetramorium Mayr Decamorium Forel Tetrogmus Roger Rhoptromyrmex Mayr Xiphomyrmex Forel syn. n. Acidomyrmex Emery Atopula Emery syn. n. Iveneella Donisthorpe syn. n. Macromischoides Wheeler syn. n. Strongylognathus Mayr T. subgen. Lobomyrmex Kratochvil syn. n. Myrmus Schenck T. subgen. Sulcomyrmex Kratochvil syn. n. Teleutomyrmex Kutter Triglyphothrix Forel Amongst these seven genera Tetrvamorium is by far the largest, exceeding all the other genera combined in number of species. This is also the genus showing the least number of specialised characteristics and is most probably the stem from which the other genera originally radiated. It is difficult to make generalizations about such highly specialized forms as Anergates and Teleutomyrmex, but the other four genera cluster closely around Tetramorium and are obvious derivations from it, mainly through a process of reduction in characters (lower palp formula and antennomere count, reduced dentition, loss of cephalic and clypeal median carinae etc.) and only more rarely by the development of characters not seen in Tetramorium itself, such as branched hairs in Triglyphothrix and falcate mandibles in Strongy- lognathus. Concerning the age of the tribe, Brown (1973) points out that the Tetramoriini may well be a recent group as it is unrepresented in any mid-Tertiary or earlier fossil deposits, whereas most other large myrmicine tribes, including nearly all those which have numerous living genera and species, are well represented or abundant in these deposits. Also, Tertiary fossil forms of numerous small myrmi- cine tribes are present in the deposits. It can be argued that as the major Tertiary deposits which yield properly identifiable ants (as opposed to fragments or impressions) are in the form of amber, then only arboreal or subarboreal forms would be expected to occur in such locations, and as most tetramoriines are terrestrial or subterranean they would be excluded from such fossiliferous systems. However, it is also true that the number of arboreal/subarboreal species of modern Myrmica, Leptothorax etc. are few when compared with the sizes of the genera as a whole but tree-dwelling forms have been represented throughout the history of these genera, and it is most probable that Tetramorium at least of the tetramoriines has as many arboreal/subarboreal forms as the other genera mentioned at the present time. It is difficult to envisage that if such forms were present in mid-Tertiary times they would not have been attracted to the resin oozing from the trees to the same extent as species from other genera. One is left with two possibilities: either the tetramoriines were scarce or absent at that time or they had not then developed arboreal species. In either case there remains the con- clusion that the vast radiation of this tribe must have occurred since mid-Tertiary times as its members are now numerous or abundant throughout the Old World ANT TRIBE TETRAMORIINI 289 tropics and subtropics in all ecosystems, and are well represented in the southern Palaearctic region. Wheeler (1914) was of the opinion that the members of the extinct Baltic amber genus Nothomyrmica resembled some species of Tetramorium but he did not include it in his section devoted to Tetramoriini, in which he actually placed Paramerano- plus, Stigmomyrmex and Enneamerus, none of which are tetramoriine ants. I examined a Wheeler-determined specimen of Nothomyrmica rudis (Mayr) in MCZ, Cambridge and have reached the conclusion that the resemblances between this genus and the tetramoriines are superficial and do not indicate relationship. All this reinforces my belief that the tetramoriine radiation occurred after the formation of the Baltic amber deposits was concluded (i.e. post-Oligocene) and it is interesting to speculate upon whether the rise of the Tetramoriini was in any way responsible for the disappearance of earlier genera which originally occupied the environments now dominated by Tetramorium and its allies in the Old World tropics. Concerning the relationships and derivation of the tribe as a whole it seems most probable, on grounds of habitus and morphology, that Tetramorium and Myrmica are descended from the same ancestral stock, and the more generalized leptothoracines also show similarities to these forms in construction of the body. Particularly interesting is the ease with which the relatively specialized dentitions of the leptothoracines and tetramoriines can be derived from the relatively generalized dentition of Myrmica in forms in which the general construction of the head and body are basically similar. In Myrmica the mandible has a long series of teeth which decrease in size from the apical to the basal and appear relatively unspecialized. In Tetvamorium the number of teeth is usually less than in Myrmica, but not much less, and specialization has been achieved by reducing the basal series in size whilst maintaining the more apical 2-3 teeth as large. In generalized leptothoracines the trend has been towards a reduction in number of teeth rather than towards specialization of what is available, and the number of teeth has been reduced to a consistent count of five, in a decreasing size series from apex to base of the masticatory margin. I consider the three tribes Myrmicini, Leptothoracini and Tetramoriini to be closely related and ultimately to have been derived from the same ancestral stock, which was most probably a creature closely resembling the modern genus Myrmica, but evidence proving or disproving this opinion is still very much incomplete. Tribe TETRAMORIINI Wheeler Tetramorii Wheeler, 1910 : 141. Type-genus: Tetvamorium Mayr. Tetramoriini Wheeler; Emery, 1914a : 38. The definition of the tribe is rendered difficult by the presence in it of two degenerate, workerless parasitic species which occupy two monotypic genera and exhibit many of the changes listed by Wilson (1971) as the ‘inquiline syndrome’. 290 B. BOLTON Disregarding these two species for the moment the remaining vast majority may be diagnosed by the possession in combination of the characters noted below. Diagnosis of worker and female Tetramoriini. Myrmicine ants in which: I, either dentition of 2-3 teeth apically, followed by a row of 3-7 denticles; never with a graded series of teeth and never with fewer than 6 teeth; or mandibles edentate and falcate; 2, palp formula never exceeding 4, 3; 3, sting with an apical or apicodorsal lamelliform appendage of varying shape; 4, anterolateral portions of clypeus raised into a ridge or shield-wall in front of the antennal insertions. Diagnosis of male Tetramoriini. Myrmicine ants in which: I, antennae with second funicular segment an elongate fusion-segment usually consisting of 3-4 antennomeres; 2, mandible dentate or falcate and edentate. DeriniTIon. The following formal definition includes all members of the tribe. Workers and females. Ants belonging to the subfamily Myrmicinae with thick, usually strongly sculptured integument. Mandibles usually triangular or subtriangular, the masti- catory margin with 2-3 large teeth apically, followed by a series of 3—7 denticles, the most frequent dental arrangement being 3 teeth plus 4 denticles, but never with less than 6 in all. Exceptions to this occur in Stvongylognathus where the mandibles are falcate and edentate, and in the workerless parasitic genera Anergates and Teleutomyrmex where the mandibles are edentate and reduced. Palp formula 4,3 at maximum, the vast majority of the tribe with this PF value but reductions to 4,2; 3,3; and 3,2 are known and in parasitic forms the PF is 1,1. Anterolateral portions of clypeus in front of the antennal insertions raised into an acute ridge or shield-wall which is confluent with the sides of the broad median portion of the clypeus as a ridge or sharp edge. This feature is a modification of the clypeus itself and is not merely the result of the impression of the head immediately posterior to the lateral parts of the clypeus. The ridge is eroded in Decamorium and reduced in Anergates. Antennae Io, II or 12-segmented, the three apical antennomeres forming a club. Antennal scrobes often present, situated above the eyes. Ocelli present in females, absent in workers. Anterior lobes of frontal carinae widely separated, the distance across the lobes in a straight line up to twice the distance from the edge of the lobe to the genal margin in full-face view. Most free-living species with a longitudinal cephalic carina running down the midline of the dorsum of the head and continued on the clypeus as a median clypeal carina. This character is reduced or absent in parasitic or dulotic forms and in some free-living species either the cephalic or the clypeal part of the carina may be absent, but it is rarely completely absent. In workers the alitrunk without dorsal sutures although the metanotal groove is generally impressed (not in Triglyphothrix). Females always winged when virgin, the alitrunk with a full complement of flight sclerites. Wing venation as in males, discussed below. Propodeal spiracle circular or subcircular, the propodeum itself usually armed with a pair of spines or teeth but exceptions to this occur in most genera. Metapleural lobes present, often acute or dentiform but by no means universally so. Legs generally with the femora more or less swollen, and often with a single simple spur on each of the middle and hind tibiae, but these are reduced or absent in many species. Petiole pedunculate, the node either squamiform or nodiform, subsessile only in the worker- less parasitic forms; the node never armed with spines or teeth, never with a massive antero- ventral process. Postpetiole very variable in shape. Sting well developed, with a lamelliform and translucent apical or apicodorsal appendage which may be spatulate, triangular, dentiform ANT TRIBE TETRAMORIINI 291 or pennant-shaped. Absent only in Anergates and Teleutomyrmex where the sting is reduced and non-functional. Males. Amongst the Myrmicinae the tetramoriine males are defined by the following combination of characters. Antennae 9-,10- or 11-segmented, the second funicular an elongate fusion-segment composed of 3-5 segments fused together. The usual antennomere count in the tribe is 10, the fusion- segment thus containing 4. Mandibles dentate except in Strongylognathus (falcate) and the workerless permanent social parasites where they are edentate and generally reduced. Palp formula as in worker/female. Eyes and ocelli present. Alitrunk with flight sclerites, all males winged except in Anérgates where the male is pupoidal. Wing venation is remarkably stable throughout the tribe and marked reduction is shown only in Teleutomyrmex and Anergates female. The usual vein-pattern is illustrated in Text-figs 8, 16, 30, 41 and the reductions in Text-figs 23, 26. In Anergates female and Teleutomyrmex, m-cu cross-veins are variably present, usually being weak, incomplete or absent, but in the former m-cu is generally visible. In most species cu-a has shifted well back along Cu towards the wing-roots, but in Anergates female this shift is not so well marked as elsewhere in the tribe. Mesoscutum and scutellum strongly developed, in profile the former overhanging the pronotum. Notauli usually present, at least with the anterior arms of the Y-shape visible, more rarely the notauli very reduced or absent. Parapsidal grooves present or absent. Genitalia usually partially retractile, more rarely apparently fully retractile. Gonopalpi present. GENERA NEWLY EXCLUDED FROM TETRAMORIINI TETRAMYRMA Forel Tetramyrma Forel, 1912d : 766 [as subgenus of Dilobocondyla}. Type-species: Dilobocondyla (Tetvamyrma) braunsi Forel, op. cit.: 767; by monotypy. Tetvamyrma Forel; Forel, 1913a : 122. [Raised to genus. | Tetvramyrma Forel; Emery, 1914a : 42. [Transferred to Tetramoriini.] There is little doubt that Forel’s original placement of Tetrvamyrma outside the Tetramoriini was more accurate than Emery’s (1914a) later transfer of this genus into the tribe, although it has since been retained there without question by later authors such as Wheeler (1922). On the present evidence the genus is certainly to be excluded from any further consideration of tetramoriine ants as both the included species lack all the basic tetramoriine characteristics. The palp formula is 5,3; the mandibles have five teeth in a decreasing-size series from apex to base; the sting lacks a lamelliform appendage apically and the clypeus is not raised into a ridge in front of the antennal insertions. On the present evidence Tetramyrma shows affinities with the Ethiopian region species of Leptothorax in the construction of the head in general and the clypeus in particular, as this has a projecting, arcuate anterior margin which overlaps the base of the mandibles. On the other hand the metanotal groove is deeply impressed, a feature not encountered in other African Leptothorax. However, in overall appearance T. braunsi bears a striking resemblance to members of the Leptothorax anacanthus-group. I have examined a syntype of L. maximus (in MCZ, Cambridge) and compared it directly to the specimens of T. brawnsz available; the similarities are obvious and it may be necessary to ask whether Tetvamyrma can retain its identity or must be sunk as a synonym of Leptothorax. 292 B. BOLTON In view of the above Tetrvamyrma is transferred for the time being into the tribe Leptothoracini, until its relationships can be studied in more detail. The species affected are as follows. Tetramyrma braunsi (Forel) Dilobocondyla (Tetramyrma) braunsi Forel, 1912d : 767. Holotype worker, SourH AFRICA: Cape Colony, Willowmore (Byauns) (BMNH) [examined]. Tetvamyrma braunsi (Forel); Forel, 1913a : 122 et auctt. Tetramyrma simoni (Emery) Tetvamorium simoni Emery, 18954 : 35, pl. 2 fig. 22. LECTOTYPE worker, SoutH AFrRica: Transvaal, Makapan (E. Simon) (MCSN, Genoa), here designated [examined]. Tetvamyrma simoni (Emery); Emery, 1922 : 291. The two species are very closely related but in simoni the propodeum is armed with a pair of teeth which are absent in braunsi. IRENEOPONE Donisthorpe Iveneopone Donisthorpe, 1946b : 242. Type-species: Iveneopone gibber Donisthorpe, loc. cit; by original designation. One of the seven ant species known to be endemic on the island of Mauritius (Brown, 1974), this odd monotypic genus was placed in the Tetramoriini by Donisthorpe without him giving any reasons for the placement. In fact many of the characters cited in the original description militate against the inclusion of Ireneopone in this tribe. The mandibles have five teeth, regularly decreasing in size from apex to base (not three as stated in the original description; the two basals are concealed by the clypeus in the holotype but are visible in the single surviving paratype). Palp formula 5,3 (as opposed to 4,3 maximum in tetramoriines). Clypeus and head capsule without median longitudinal carinae. Lateral portions of clypeus not raised into a shield in front of the antennal insertions but the area below the insertions locally depressed behind the lateral parts of the clypeus. Sting without a lamelliform appendage apically. Ireneopone gibber Donisthorpe Iveneopone gibbey Donisthorpe, 1946) : 242, figs 1, 2. Holotype and paratype workers, Mauritius: Calebasses Mt., 22.x.1944, no 72 (R. Mamet); and Le Pouce Mt., 7.x1i.1940 (J. Vinson) (BMNH) [examined]. Having shown that Iveneopone does not belong in the Tetramoriini, the problem is now to find its correct tribal placement. For the present I propose to transfer this genus to the tribe Myrmecinini, close to the complex of small genera centring on Terataner. The reason for this is that, besides having the characters given ANT TRIBE TETRAMORIINI 293 above, Iveneopone has the antennal insertions widely separated, the clypeus strongly inserted between them. The mesonotum is delimited by a raised portion in front and an impressed metanotal groove behind. The propodeum is unarmed and rounded metapleural lobes are present. The petiole has a long anterior peduncle and a low, narrow node which is rounded-subconical in profile and with a short posterior peduncle. It is by no means claimed that this is the last word on the placement of this peculiar species; the final decision must await a revisionary study of the myrme- cinine genera. EUTETRAMORIUM Emery Eutetramorium Emery, 1899 : 280. Type-species: Eutetramorium mocquerysi Emery, op. cit.: 281; by subsequent designation of Wheeler, 1911 : 163. Since its description Eutetramorium has apparently always been treated as a bona fide member of tribe Tetramoriini, throughout all the varying concepts of this tribe (Wheeler, 1910; 1922; Emery, 1914a@; 1922; Forel, 1917). The only reason for this appears to be the fact that the clypeus is raised laterally, in front of the antennal insertions. Other tetramoriine characters are lacking excepting the palp formula which has a typically tetramoriine count of 4,3. However, the basal segments of both palpi are very long and appear to be the result of the fusion of several shorter palpomeres (paralectotype of E. mocquerysi dissected). The critical lamelliform sting appendage is lacking, as is the differentiated tetramoriine dentition. I am presently of the opinion that Eutetramorium is better placed in the tribe Myrmicini as, beside the fact that the tibial spurs of the middle and hind legs are minutely barbulate or serrated, the general appearance of the head capsule, alitrunk and pedicel is very close to the rugosa-like species of Myrmica of the Oriental and south-eastern Palaearctic regions, some of which also have the lateral portions of the clypeus raised into a shield in front of the antennal insertions. Both known species of this small genus are from Madagascar. Eutetramorium mocquerysi Emery Eutetvamorium mocqueryst Emery, 1899 : 281, fig. LECTOTYPE and paralectotype workers, Mapacascar: Antongil, 1897-1898 (A. Mocquerys) (MCSN, Genoa), here designated [examined]. Eutetramorium monticellii Emery Eutetramorium monticellii Emery, 1899 : 283. Syntype worker and female, MADAGASCAR: Antongil, 1897-1898 (A. Mocquerys) (MCSN, Genoa) [examined]. SPECIES NEWLY EXCLUDED FROM TETRAMORIINI ROMBLONELLA Wheeler Romblonella Wheeler, 1935 : 5. Type-species: Romblonella grandinodis Wheeler, op. cit.: 7 [= Myrmica opaca F. Smith, 1861 : 47]; by original designation. 204 B. BOLTON A small and somewhat obscure genus of ants belonging to the tribe Meranoplini, which contains some seven species. On original description the genus was mono- typic and, during the course of this study, it was discovered that Tetramorium opacum is in fact the senior synonym of the type-species of Romblonella. What is known of the genus is summarised in M. R. Smith (1953 : 1956), and this remains unchanged except for the following, which is now confirmed. Romblonella opaca (F. Smith) comb. n. Myrmica opaca F. Smith, 1861 : 47. Holotype worker, SuLAweEs1: Tondano (A. R. Wallace) (UM, Oxford) [examined]. Tetvamorium opacum (F, Smith); Donisthorpe, 1932 : 469 Romblonella grandinodis Wheeler, 1935 : 7, fig. 2. Syntype workers, PHILIPPINES: Romblon I., 6.v.1928 (L. Marato) (MCZ, Cambridge) [examined]. Syn. n. CHELANER Emery Chelaner Emery, 19146: 410 [as subgenus of Monomorium]. Type-species: Monomorium (Chelaner) forcipatum Emery, loc. cit.; by subsequent designation of Emery, 1922 : 168. Chelaner Emery; Ettershank, 1966 : 93. [Raised to genus. ] Chelaner flavigaster (Clark) comb. n. Xiphomyrmex flavigaster Clark, 1938 : 366, fig. 5. Syntype workers, AUSTRALIA: Sir Joseph Banks Islands, Reevesby I. (ANIC, Canberra.) From Clark’s original description it is obvious that flavigaster is not a tetramoriine, but its correct generic placement is difficult. For the identification of this species as a member of genus Chelaney I am indebted to Dr R. W. Taylor of ANIC, Canberra, who kindly examined the relevant type-material for me. KEYS TO GENERA OF TETRAMORIINI Workers 1 Mandibles narrow and falcate, edentate or at most with a single minute denticle close to the apex (Text-figs 13, 14) (Palaearctic) : STRONGYLOGNATHUS (p. 304) — Mandibles triangular or subtriangular, dentate, with two or three enlarged teeth apically followed by a series of 3-7 denticles_ . : 2 2 Some or all dorsal surfaces of head and body equipped with peeehed oe ies may be bifid, trifid or quadrifid, simple hairs often also present. Metanotal groove in profile not impressed. (Old World Tropics, but no known endemics in Australia or Madagascar) . : TRIGLYPHOTHRIX (p. 310) — Dorsal surfaces of head andl ody. without bifid, peed or quadrifid hairs; simple hairs usually present but some species with bizarre pilosity and some hairless. Metanotal groove often impressed . é 4 2 : : : é E ; 3 3 Antennae with 10 segments (Ethiopian) : : : : DECAMORIUM (p. 297) — Antennae with 11 or 12 segments ; 4 4 Palp formula 3,2. Head roughly heart- -shaped (Text: figs 1-3) i sides convex and 2 Ww fo) Leal ANT TRIBE TETRAMORIINI occiput concave, the head narrowing anteriorly. Median portion of clypeus with a prominent, arcuate anterior margin which overlaps the basal angle of the mandible. Median clypeal and median cephalic carinae vestigial or absent. (Old World Tropics) : : . RHOPTROMYRMEX (p. Palp formula usnally. 4,3, oeay ely coded (one species known with PF 3,2). Head not heart-shaped. Median portion of clypeus with anterior margin not prominent, not overlapping the basal angle of the mandible. Usually both the median clypeal and median cephalic carinae present, more rarely with one or the other absent, very rarely with both absent. (Worldwide but with no Neotropical endemics, one Nearctic endemic) . : : ’ : : : . TETRAMORIUM (p. Females (Queens) Mandibles dentate, armed with 2-3 enlarged apical teeth followed 96 a row of 3-7 denticles : ; : : : ; Mandibles edentate, either faleate or rattced : Branched hairs present on some or all surfaces of the head, alitranik, pedicel abd gaster, either bifid, trifid or quadrifid : : 2 TRIGLYPHOTHRIX (p. Branched hairs absent, simple hairs usually present although in some species pilosity is bizarre or absent Antennae with 10 segments - : 3 : : . DECAMORIUM (p. Antennae with 11 or 12 segments Palp formula 3,2. Median portion of Glypeus with a " prominent, areuate siderite margin which overlaps the basal angle of the mandible. Median clypeal and median cephalic carinae vestigial or absent - . RHOPTROMYRMEX (p. Palp formula usually 4,3, rarely less. Median portion of clypeus without a prominent, arcuate anterior margin which overlaps the basal angle of the mandible. Median clypeal and median cephalic carinae usually 3 pte sometimes one or the other absent, rarely with both absent : : . TETRAMORIUM (p. Anterior clypeal margin deeply emarginate or notched medially (Text-fig. 19). (Palaearctic, Nearctic) . p : ANERGATES (p. Anterior clypeal margin entire, not deeply emarginate or notched medially Mandibles falcate; palp formula 4,3. Postpetiole not broadly attached to first gastral tergite, gaster not dorsoventrally flattened. Sting present, with an apicodorsal appendage. Worker caste present . 3 : Mandibles blade-like (Text-fig. 24); palp formula 1,1. Postpetiole very broadly attached to first gastral tergite, gaster strongly dorsoventrally flattened. Sting reduced and non-functional, without apicodorsal appendage. Worker caste absent. (Palaearctic) . : : , - : : : TELEUTOMYRMEX (p. Males Mandibles edentate, either reduced or reduced-falcate . Mandibles dentate : : : ‘ : : ; ; : : ; Pupoidal and apterous. Anterior clypeal margin hoa emarginate or notched medially ‘ : -ANERGATES (p. Winged, body fully developed or = somewhat Pupoidal | Anterior clypeal margin not emarginate or notched medially Mandibles short-falcate; palp formula 4, ra Gaster not downcurved, ae reflexed anteriorly at the apex . : STRONGYLOGNATHUS (p. Mandibles blade-like and very remecd: “4 formula 1,1. Gaster downcurved and somewhat reflexed anteriorly at the apex. STRONGYLOGNATHUS (p. 295 298) 359) 298) 359) 296) 6 296) 3 304) TELEUTOMYRMEX (p. 309) 296 B. BOLTON 4 Some or all dorsal surfaces of head, alitrunk, pedicel and gaster with branched bifid, trifid or quadrifid hairs . : 3 : TRIGLYPHOTHRIX (p. 310) — Branched hairs absent; simple hairs eenell, present but bizarre ea present in some species . : 5 5 Antennae basically 9- segmented, the Soe apical gated geemicues (aed or definitely incrassate. Palp formula 3,2 . é . RHOPTROMYRMEX (p. 298) — Antennae Io or 11-segmented, the funicular segments filiform, not incrassate apically. . Palp formula usually 4,3 TETRAMORIUM (p. 359) and DECAMORIUM (p. 297) ANERGATES Forel (Text-figs 19-23) Anergates Forel, 1874 : 32. Type-species: Myrmica atratula Schenck, 1852 : 91; by monotypy. Diacnosis. Female. Mandibles reduced, edentate except for the apical tooth. Palp formula 1,1 (Ettershank, 1966, here confirmed), the palps reduced. Clypeus with its median anterior margin broadly and deeply emarginate. Antennae Io- or 11-segmented with a variable degree of fusion of the second to fourth funicular antennomeres (at arrow in Text-fig. 19). Ocelli present (Text-fig. 19). Alitrunk with full set of flight sclerites, the virgin females winged (Text-fig. 23). Postpetiole strongly expanded, broadly attached to gaster (Text-fig. 21). Gaster in virgin females with a broad and deep longitudinal furrow or impression; mature females strongly physogastric. Sting strongly reduced, non-functional. Male. Pupoidal and apterous. Mandibles lobiform, edentate, not capable of full closure (Text-fig. 20). Palp formula 1,1 as in female. Median anterior clypeal margin broadly and deeply emarginate. Antennae Io- or 11-segmented, the third funicular antennomere variably fused with the fourth (Text-fig. 20). Second funicular segment usually quite long when compared to those just distal to it and apparently is a shortened fusion-segment. Ocelli present. Alitrunk with flight sclerites but wings never developed. Petiole and post-petiole strongly anteroposteriorly compressed, roughly thick-disciform in shape and each very broadly attached to the following segment (Text-fig. 22). Apex of gaster reflexed ventrally. Genitalia large, strongly extruded. The single species belonging to this genus, A. atvatulus, is a degenerate, workerless, permanent social parasite in nests of Tetvamorium caespitum. Emery (1922) placed Anergates in a subtribe Anergatini of tribe Solenopsidini, but Ettershank (1966), in his revision of the solenopsidine genera, excluded Anergates and suggested that the genus should be transferred to Tetramoriini on the grounds of its reduced antennal segmentation, wing venation and close parasitic association with Tetra- morium. I am in agreement with this suggestion and believe that Anergates is developed from a T, caespitum-like ancestor although its origins can probably never be traced with certainty because of the many morphological adaptations made by this species in response to its parasitic lifeway. Wilson (1971) has listed 19 such characteristic adaptations of parasitic ants which he terms the ‘inquiline syndrome’, many of which apply to Anergates. What is known of the biology of Amnergates is summarised in Wheeler (1910), Creighton (1950), Kutter (1969), Wilson (1971) and their included references. Anergates atratulus (Schenck) Myrmica atratula Schenck, 1852: 91. Syntype female, male, GERMANY (WEsT): Nassau distr., NW. of Franfurt (Schenck?) (types possibly in MNHU, Berlin). Anergates atvatulus (Schenck) ; Forel, 1874 : 32. ANT TRIBE TETRAMORIINI 297 Anergates friedlandi Creighton, 1934 : 193. Holotype female, U.S.A.: New Jersey, side of U.S. Highway 9, 3 miles N. of Englewood, 31.v.1932 (S. Friedland) (probably in MCZ, Cambridge, originally in Creighton private coll.). [Synonymy by Creighton, 1950.] A. atratulus is distributed throughout Europe within the range of its host species and is also known from Ukraine and the U.S.A. from Connecticut to N. Virginia. To the best of my knowledge it has never yet been recorded outside the nests of its host although there is little doubt that the alate females fly when searching for new host colonies. DECAMORIUM Forel (Text-figs 27-31) Decamorium Forel, 1913a: 121 [as subgenus of Tetramorium]. Type-species: Tetvamorium (Decamorium) decem Forel, loc. cit.; by monotypy. Decamorium Forel; Emery, 1914a : 42. [Raised to genus.] Diacnosis. Worker. Mandibles armed with three teeth plus a series of 4-5 denticles; alternate members of the denticle series enlarged, in some cases with denticles doubled (Text-fig. 28). Palp formula 4,3 (dissections of decem and welense). Antennae with 10 segments. Median longitudinal carina of clypeus absent, but present on dorsum of head capsule. Lateral raised portions of clypeus eroded and very reduced. Antennal scrobes strongly developed, with a marked dorsal margin but very poorly demarcated ventrally, median carina of scrobes absent (Text-fig. 27). Metanotal groove impressed; propodeum armed with a pair of short spines (Text-fig. 31). Apical lamelliform appendage of sting triangular, situated apicodorsally. Female. As worker but with ocelli and flight sclerites; winged when virgin (Text-fig. 30). Male. As Tetvamorium. This small genus contains only two species, is restricted to the Ethiopian region, and is but dubiously maintained as separate from Tetramorium on the grounds of the reduced antennal segmentation, reduced clypeal shield and the modified dentition in workers and females. In the long run none of these characters may be sufficient to keep Decamorium as a genus, for as can be seen, other genera in this tribe show variation in antennomere count; but at present no linking species between Decamorium and Tetramorium are known. Of the two species, decem is reported by Arnold (1917) as forming quite large colonies in sandy soil, whereas welense nests in twigs in leaf litter and humus and appears to form only small colonies. The habits of decem are not known but I have observed welense in western Nigeria raiding a small subterranean termite colony in open woodland. The workers of welense were leaving their nest (in a rotten twig) early in the morning and progressing in single file to the termite colony which was a few yards away. They moved through the leaf-litter and over a rotten branch in single file, almost nose to tail. The two species of Decamorium are separated in the worker cast as follows. 1 Dorsum of promesonotum strongly and distinctly longitudinally striate, Pung with scattered punctures also. (Ghana, Nigeria, Guinea, Zaire) c : uelense — Dorsum of promesonotum smooth and shining, with scattered punctures. (Rhodesia, Gabon, Ghana) . c - ¢ 0 : : 0 : - : decem 298 B. BOLTON SYNONYMIC LIST OF SPECIES decem (Forel). decem var. ultoy Forel syn. n. uelense Santschi stat. n. decem subsp. nimba Bernard syn. n. SUMMARY OF THE SPECIES Decamorium decem (Forel) (Text-figs 27-31) Tetvamorium (Decamorium) decem Forel, 19134 : 121. Syntype workers, RHopDESIA: Redbank, 7.1v.1912 (G. Avnold) (BMNH; MHN, Geneva) [examined]. Decamorium decem (Forel); Emery, 1914a : 42. [Implied in text.] Tetramorium (Decamorium) decem var. ultoy Forel, 1913b : 217. Syntype workers, RHODESIA: Shiloh, 12.v.1913 (G. Avnold) (BMNH; MHN, Geneva) [examined]. Syn. n. Decamorium uelense (Santschi) stat. n. Tetvamorium (Decamorium) decem st. uelense Santschi, 1923 : 285. Syntype worker, female, ZAIRE: Uelé, Vankerkhovenville (Degveef) (NM, Basle; MRAC, Tervuren) [examined]. Decamorium decem subsp. nimba Bernard, 1952 : 250. Holotype worker, GUINEA: Savane de Kéoulenta, st. 1 (Lamotte) (MNHN, Paris) [examined]. Syn.n RHOPTROMYRMEX Mayr (Text-figs I-12) Rhoptromyrmex Mayr, 1901 : 18. Type-species: Rhoptromyrmex globulinodis Mayr, op. cit.: 20; by subsequent designation of Wheeler, 1911. Acidomyrmex Emery, 19150) : 191 [subgenus of Rhoptvomyrmex]. Type-species: Rhoptromyrmex wroughtonit Forel, 1902a : 231; by original designation. [Synonymy by Brown, 1964.] Iveneella Donisthorpe, 1941 : 175. Type-species: Iveneella papuensis Donisthorpe, loc. cit.; by original designation. Syn. n. DiaGnosis. Worker. Head including mandibles roughly heart-shaped in full-face view (Text-figs 1-3), with the sides strongly convex, the occipital margin concave and the head capsule much broader behind than in front. Anterior margin of median portion of clypeus projecting and broadly arcuate. Antennal scrobes absent. Median longitudinal cephalic carina vestigial (indistinguishable from surrounding sculpture) or absent. Antennae II- or 12-segmented, the apical three forming a club. Mandibles with two apical teeth, a much smaller third tooth and a row of 4-6 denticles. Palp formula 3,2 (dissections of melleus, wroughtonii, critchleyi, globulinodis, opacus, transversinodis; see also Brown, 1964). Metanotal groove impressed, but only very feebly so in some specimens. Propodeum armed with a pair of spines in Oriental and Indo-Australian species, unarmed in those from the Ethiopian region (Text-figs 9, 10) except in critchleyi (Text-fig. 12). Ventral surface of petiole generally longitudinally keel-like and convex. Sting with a triangular or dentiform lamelliform appendage apicodorsally. Female. Adapted for a (presumed) temporary social-parasitic lifeway. Generally details ef antennal segmentation, palp formula, dentition and sting structure as in worker but in one or two species the mandibles modified towards a falcate condition or otherwise reduced. ANT TRIBE TETRAMORIINI 299 Alitrunk usually compressed from side to side, narrow and deep in dorsal view but retaining a full set of flight sclerites, and winged when virgin (Text-figs 4, 5). Male. Antennae basically nine-segmented, the second funicular being an elongate fusion- segment (Text-figs 6, 7). Partial freeing of the apical member of this fusion-series or partial fusion of the next antennomere may give an ambiguous count of 8, 9 or 10 segments in some specimens. Rarely the limits of the fused segments are all visible. The three apical antenno- meres tend to be enlarged and are distinctly incrassate in Oriental and Indo-Australian species. Palp formula 3,2 as in worker (dissections of melleus, globulinodis, tvansversinodis). Notauli weakly developed or absent. Radial cell in forewing open or closed. Rhoptromyrmex is a small genus of tetramoriine ants containing only six species, four of which occur in the Ethiopian region and the other two distributed in the Oriental and Indo-Australian regions. The workers are typically tetramoriine in most of their characters but have developed away from their basal stock (believed to be in Tetramorium) by developing the characteristic head shape seen in this genus and by the accompanying loss of antennal scrobes, reduction of palp segmentation and virtual or complete disappearance of the cephalic median carina. Emery (19150; 1922) separated a subgenus Acidomyrmex to include the two Oriental and Indo-Australian species, solely on the grounds that the propodeum is armed in the workers of these two species, whereas it was unarmed in the then- known workers of Ethiopian’ region species. Brown (1964) pointed out that the size of the propodeal spines was very variable in wroughtonii and that the female of melleus has the propodeum unarmed, whereas in the African species the propodeal armament is strongly developed in females of tvansversinodis (and in workers of the new species, cvitchleyi). To define a subgenus on a character which only the worker shows, and which is demonstrably variable in one species, seemed unrealistic and Acidomyrmex was placed in synonymy. The discovery of critchleyi, a species from Nigeria which has propodeal spines present, further validates Brown’s conclusion. The females of Rhoptromyrmex (Text-figs 4, 5) are either aberrant or very aberrant, and Brown (1964) points out that many of their modifications resemble those shown in other species which are known to be temporary social parasites. Whether this theory is correct still remains to be seen as Rhoptromyrmex plus host colonies remain unknown. The males are again typically tetramoriine, being specialised only in their reduced palp formula and antennal segmentation, which is basically of nine antennomeres here as opposed to ten in the remainder of the tribe (Text-figs 6, 7). The males of five species are now known as the single member of the monotypic genus [veneella appears to be the male of melleus. Donisthorpe (1941) distinguished his genus Jveneella from other tetramoriines on the following characters: I, antennae with nine segments; 2, palp formula 3,2 (confirmed in present study); 3, notauli absent; 4, mesonotum not overhanging pronotum; 5, ocelli smaller; 6, antennae not filiform apically (i.e. they are clavate); 300 B. BOLTON 7, petiole is ‘differently formed’; 8, radial cell in forewing is open. Disregarding those characters which are known to be variable within this tribe (numbers 5, 7, 8), what remains is virtually a characterisation of the males of genus Rhoptromyrmex. Comparison of the types of I. papuensis with males of other Rhoptromyrmex species confirmed the synonymy of Iveneella within Rhoptro- myrmex. Further, the very close resemblance between the males constituting the type-series of I. papuensis and those of wroughtonii (the two are separated only on details of sculpture) leads me to believe that papuensis is in fact the male of melleus, previously unknown. Circumstantial evidence in favour of this assumption and weighing against papuensis being merely a more heavily sculptured male of wroughtonit than was previously known lies in the fact that melleus is known mainly from New Guinea, the locality of papuensis, whilst the range of wroughtonii is not known to extend to that island. I feel sure that the association of papuensis with melleus is valid, and that further collections of worker-associated melleus males will confirm this synonymy. Nests of Rhoptromyrmex species may be very large and are made directly into the soil or under stones. Brown (1964) gives biological notes on some species and states that they are ‘general feeders, the ants collect living and dead arthropods, tend homopterous insects for their honeydew both above and below ground, and feed at plant nectaries’. SYNONYMIC LIST OF SPECIES critchleyi sp. n. globulinodis Mayr globulinodis st. alberti Forel globulinodis var. obscurus Santschi melleus (Emery) papuensis Donisthorpe syn. n. opacus Emery opacus var. esta Forel opacus var. laeviceps Santschi ' opacus subsp. monodi Bernard transversinodis Mayr steint Forel tvansversinodis var. pretoriae Arnold wroughtonii Forel wroughtonit st. vothneyi Forel wroughtonit st. vothneyi var. longi Forel vothneyt var. intermedia Forel vothneyi st. sumatrensis Forel vothneyt subsp. Jeno Viehmeyer vothneyi var. taivanensis Wheeler All the above synonymy, with the exception of melleus = papuensis, is from the recent revision of Rhoptromyrmex by Brown (1964). Besides the above Brown ANT TRIBE TETRAMORIINI 301 also removed a number of species originally described in Rhoptromyrmex and transferred them as follows. Rhoptromyrmex tessmanni Forel, 1910b : 421; transferred to Macromischoides as a _ synonym of M. africanus (Mayr) [now in Tetramorium]. Rhoptromyrmex sollert Forel, 1910b : 430; transferred to Monomorium Mayr. Rhoptromyrmex mayri Forel, 1g12c : 57; transferred to Hagioxenus Forel. KEY TO SPECIES (Workers) (based on Brown, 1964) Propodeum armed with a pair of spines or teeth (Text-figs 11, 12) . Propodeum rounded, unarmed (Text-figs 9, 10) : : ‘ ; Antennae with eleven segments (Nigeria) . critchleyi (p. 303 Antennae with twelve segments (Indo-Australian and Griental regions) . Propodeal spines very long, about twice as long as the distance between the centres of their bases (Text-fig. 11) and just about as long as the maximum width of the pronotum. Sculpture of head predominantly finely and densely reticulate- punctate, longitudinal costulae obsolete or few and weak. (New Guinea; Austra- lia: N. Cape York Peninsula) . : : melleus - Propodeal spines variable in length but less than twice as s long as the distance between the centres of their bases, and shorter than the pronotal width. Head and usually also the alitrunk with fine, close longitudinal costulation prominent in the sculpture. (Nepal, India, Burma, Thailand, S. China, Taiwan, Philippines, Borneo, Malaya, Sumatra, Java, Sumba, Australia: Cape York peninsula) . : : wroughtonii 4 Alitrunk and usually also the head with dense, opaque reticulate-punctate sculpture throughout. (Guinea, Uganda, Cameroun, Zaire) . : opacus Head and alitrunk in large part smooth and shining (mainly southern Africa). : 5 5 Postpetiole subglobular, in dorsal view up to about 1.5 times broader than long (Text-fig. 9); in profile with a prominent rounded ventral protuberance. (Zaire, Rhodesia, South Africa) . ‘ : globulinodis - Postpetiole transversely subrectangular, about twice broader than long (Text-fig. ro) ; in profile without a prominent ventral protuberance. (Rhodesia, South Africa) transversinodis SAN wilndt a Ww SUMMARY OF THE SPECIES As the genus has been fully revised (Brown, 1964) full descriptions are not included here, but a summary of the species, their distribution and synonymy is given. INDO-AUSTRALIAN AND ORIENTAL REGIONS. THE MELLEUS-Grovup Rhoptromyrmex melleus (Emery) (Text-figs 2, II) Tetramorium melleum Emery, 1897 : 586, pl. 15, figs 29, 30. Holotype worker, NEw GUINEA: Belaio Island, near Friedrich-Wilhelmshafen (= Madang) (L. Bivo) (HNM, Budapest). 302 B. BOLTON Rhoptromyrmex (Acidomyrmex) melleus (Emery); Emery, 1922 : 290. Iveneella papuensis Donisthorpe, 1941 : 175. Holotype and paratype males, NEw GUINEA: Papua, Mafula, 4000 ft, xii.1933 (L. E. Cheesman) (BMNH) [examined]. Syn. n. [See under discussion of genus. ] Distribution. New Guinea and Cape York Peninsula, Australia. Rhoptromyrmex wroughtonii Forel (Text-fig. 7) Rhoptromyrmex wroughtoni Forel, 1902a : 231. Syntype workers, male, Inp1a: Kanara (Wroughton) (MHN, Geneva). Rhoptromyrmex wroughtoniu st. vothneyi Forel, 1902a : 232. Syntype workers, Inp1a: Banga- lore (Rothney) (MHN, Geneva). [Synonymy by Brown, 1964.] Rhoptromyrmex wroughtonir st. vothneyi var. longi Forel, 1902a : 232. Inpi1A: Assam, Garo Hills (MHN, Geneva). [Synonymy by Brown, 1964.] Rhoptromyrmex vothneyi var. intermedia Forel, 1913c : 80. Syntype workers, SUMATRA: Beras Tagi, 4500 ft (Buttel-Reepen) (MHN, Geneva). [Synonymy by Brown, 1964.] Rhoptromyrmex vothneyi st. sumatrensis Forel, 1913c : 80, fig. W. Syntype workers, SUMATRA: Kampong Keling, near Beras Tagi, 4500 ft (Buttel-Reepen) (MHN, Geneva). [Synonymy by Brown, 1964.] Rhopthromyrmex [sic] vothneyit subsp. leno Viehmeyer, 1914 : 113. Holotype worker, WEST Mataysia: Perak (E. Streesemann) (location of type not known). [Synonymy by Brown, 1964.] Rhoptromyrmex (Acidomyrmex) rothneyi var. taivanensis Wheeler, 1930 : 103. Syntype workers, Tatwan: Hakumo (R. Takahashi) (MCZ, Cambridge). [Synonymy by Brown, 1964.] Distribution. India, Thailand, S. China, Philippines, Taiwan, Malaya, Indo- nesia to Sumba, Australia: Cape York Peninsula. ETHIOPIAN REGION. THE GLOBULINODIS-Grovup Rhoptromyrmex globulinodis Mayr (Text-figs 1, 4, 6, 9) Rhoptromyrmex globulinodis Mayr, 1901 : 20. Syntypes workers, female, male, SouTH AFRICA: Port Elizabeth (NM, Vienna; BMNH) [examined]. Rhoptromyrmex globulinodis st. alberti Forel, 1916: 419. Syntype workers, ZAIRE (Kohl) (MHN, Geneva). [Synonymy by Brown, 1964.] Rhoptromyrmex globulinodis var. obscurus Santschi, 1932 : 339. Syntype workers, male, RHODESIA: Vumba Mts, Cloudland, 6000 ft, iv. 1923 (G. Avnold) (NM, Basle). [Synonymy by Brown, 1964.] Distribution. Zaire, Rhodesia, South Africa. Rhoptromyrmex opacus Emery Rhoptromyrmex opacus Emery in Forel, 1909 : 59 (footnote). Syntype workers, CAMEROUN (Conradt) (MCSN, Genoa). ANT TRIBE TETRAMORIINI 303 Rhoptromyrmex opacus var. esta Forel, 1909 : 59. Syntype workers, females, males, ZAIRE: Bas Congo (BMNH; MHN, Geneva) [examined]. [Synonymy by Brown, 1964.]. Rhoptromyrmex opacus var. laeviceps Santschi, 1916 : 504. Holotype worker, ZarrE: Boma (Bondroit) (NM, Basle). [Synonymy by Brown, 1964.] Rhoptromyrmex opacus subsp. monodi Bernard, 1952 : 251, fig. 14F. Holotype worker, GUINEA: Mt Nimba, Ziéla, st. F 32, savane (Lamotte) (location of holotype not known). [Synonymy by Brown, 1964.] Distribution, Guinea, Uganda, Cameroun, Zaire. Rhoptromyrmex transversinodis Mayr (Text-figs 5, 8, Io) Rhoptromyrmex transversinodis Mayr, 1901 : 22. Syntype workers, SouTH AFRICA: Port Elizabeth (NM, Vienna; BMNH) [examined]. Rhoptromyrmex steini Forel, 1913a: 122. Syntype workers, SoutH ArFrica: Cape Prov., Ladismith (H. Brauns) (MHN, Geneva). [Synonymy by Brown, 1964.] Rhoptromyrmex transversinodis var. pretoviae Arnold, 1926 : 282. Syntype workers, female, male, SouTH AFRICA: Pretoria, 21.iv.1915 (C. K. Brain) (BMNH) [examined]. [Synonymy by Brown, 1964.] Distribution. Rhodesia, South Africa. Rhoptromyrmex critchleyi sp. n. (Text-figs 3, 12) Holotype worker. TL 2-7, HL 0:66, HW 0:64, CI 97, SL 0-42, SI 66, PW 0-42, AL 0°74. With the following combination of characters within the genus: I, antennae with eleven segments; 2, propodeum armed with a pair of spines; 3, head, alitrunk and pedicel densely reticulate-punctate ; 4, postpetiole in dorsal view subglobular, somewhat broader than long. Paratypes: two workers. As holotype but measuring: TL 2-4-2:6, HL 0:60-0:64, HW 0:-58- 0:64, CI 97-100, SL 0-38-0-40, SI 63-65, PW 0:36-0-42, AL 0:66-0:72 (2 measured). Holotype worker, NicERIA: Ibadan, I.1.T.A. no. 34G, 23-27.xi.1973 (B. R. Critchley) (BMNH). Paratypes. 2 workers, as holotype but collected on 11-16.x.1973 (BMNH; MCZ, Cambridge). Within the genus this species is very easy to distinguish as it is the only known species of the Ethiopian region in which propodeal spines are present in the worker caste, and it is the only member of the genus as a whole in which the antennae have eleven segments (as opposed to twelve in all other species). Apart from its reduced antennomere count, critchleyi is a quite ordinary member of genus Rhoptromyrmex and tends to substantiate the synonymy of Acidomyrmex within this genus (see discussion above). It also shows again that the use of antennal segment number to diagnose the genera of Tetramoriini is unsatisfactory. 304 B. BOLTON As in other genera the number of antennal segments varies in Rhoptromyrmex whilst other characters of diagnostic value at generic level remain constant. This is also seen to be true in Tetvamorium and Triglyphothrix, where counts of II or 12, and 10, II or 12 respectively, have been recorded. STRONGYLOGNATAUS Mayr (Text-figs 13-18) Strongylognathus Mayr, 1853 : 389. Type-species: Eciton testacewm Schenck, 1852 : 117 [= Myymus emarginatus Schenck, 1853]; by monotypy. Myrmus Schenck, 1853 : 188. Type-species: Myyvmus emarginatus Schenck, loc. cit.; by monotypy. [Junior homonym of Myrmus Hahn, 1832 : 81.] Diacnosis. Worker. Mandibles falcate, edentate or with a minute denticle which may occur near the apex in some individuals. Mandibular blades elongate, tapering, and curved from base to apex so that when their tips cross a space is enclosed between their inner margins and the anterior clypeal margin (Text-figs 13, 14). Palp formula 4,3 (dissections of testaceus, afer, hubert, destefanii). Median longitudinal carina of clypeus and head capsule absent. Antennal scrobes absent. Antennae 12-segmented, the three apical antennomeres forming a club. Metanotal groove feebly impressed; propodeum usually with a pair of very small teeth or denticles (Text-figs 17, 18), in some these are so reduced that the propodeum is virtually unarmed. Apicodorsal lamelliform appendage of sting triangular or pennant-shaped. Female. As worker but with alitrunk possessing a full complement of flight sclerites, the queens winged when virgin. Ocelli present. Male. Mandibles falcate and edentate as in worker and female, but relatively much smaller. Antennae 1o-segmented, the second funicular forming an elongate fusion-segment as is usual in the tribe (Text-fig. 15). Amongst the Tetramoriini Stvongylognathus is easily distinguished by its remark- able falcate mandibles. The genus is restricted, as far as is known, to the Palaearctic region, ranging from North Africa to northern U.S.S.R., and from the British Isles to Korea and Japan. All species of the genus live in a dulotic, a rather degenerate obligate dulotic association, or as inquilines, in nests of Tetramorium caespitum (L.) and its close relatives. Information on the biology of the species may be found in Wheeler (1910), Stumper (1950), Kutter (1923; 1969), Wilson (1971), and the references included in these publications. The taxonomy of the huberi-group species of western Europe has recently been worked out by Baroni Urbani (1969), and some species of eastern Europe and U.S.S.R. have been reviewed by Pisarski (1965), but a formal revision of the genus in its entirety has not yet been undertaken. Roughly, the genus falls into two species-groups. The first of these includes only testaceus (Text-figs 13, 17) and kavawajewt, in which the occipital angles are strongly produced, whilst the second includes all the remaining forms described to the present in which the occipital angles are not produced (Text-fig. 14). In the list and summary given below, no infraspecific taxa are recognized as throughout the other genera of this tribe they have proved illusory and in all cases have turned out to be either good species or synonyms of some other named form. In the light of these findings the infraspecific names remaining in Strongylognathus, after ANT TRIBE TETRAMORIINI 305 the partial revisions of Pisarski (1965) and Urbani (1969) have been taken into account, are here all raised provisionally to the rank of species until a formal revision of the entire genus can be undertaken. This action applies to the names foreli, ruzskyi, bulgaricus and italicus, but is not to be taken as the last word on the subject. In fact, the variation shown between series constituting what are presently termed species in the huberi-group are in many cases extremely trivial and no greater than that found between different populations or nests of what are unquestionably the same species elsewhere in the tribe. I think that many of the species-level names in the genus may merely be localized populations and I am convinced that further collections and study will reduce the number of species in Strongylognathus to a much lower figure. SYNONYMIC LIST OF SPECIES testaceus-group karawajewi Pisarski testaceus (Schenck) emarginatus (Schenck) divert Donisthorpe huberi-group afer Emery alboini Finzi alpinus Wheeler bulgaricus Viehmeyer stat. n. kratochvili Silhavy caeciliae Forel cecconii Emery christophi Emery dalmaticus Baroni Urbani destefanii Emery huberi st. cecconit var. kutteyi Santschi emeryi Menozzi foreli Emery stat. n. huberi Forel huberi var. gallica Emery insularis Baroni Urbani italicus Finzi stat. n. kervillei Santschi koreanus Pisarski palaestinensis Menozzi rehbinderi Forel ruzskyi Emery stat. n. silvestrii Menozzi SUMMARY OF THE SPECIES THE TESTACEUS-Group Strongylognathus karawajewi Pisarski Strongylognathus kavawajewi Pisarski, 1965 : 521, figs 40-43. Holotype and paratype workers, 306 B. BOLTON U.S.S.R.: Crimea, Magaratsch 26.iii.1903 (Ryasantev) (MNHU, Berlin; Institute of Zoology, Kiev; Institute of Zoology, Warsaw). Distribution. U.S.S.R.: Crimea. Strongylognathus testaceus (Schenck) (Text-figs 13, 15, 16, 17) Eciton testaceum Schenck, 1852: 117. Syntype worker, female, male, GERMANY (WEST): Nassau distr., NW. of Frankfurt (coll. Schenck?) (location of types not known, perhaps in MNHU, Berlin). Strongylognathus testaceus (Schenck); Mayr, 1853 : 389. Myrmus emarginatus Schenck, 1853 : 188. Syntype workers, GERMANY (WEST): Nassau distr., NW. of Frankfurt (Schenck) (location of types not known, perhaps in MNHU, Berlin). [Synonymy by Mayr, 1853.] Strongylognathus diveri Donisthorpe, 1936 : 113, figs I, 2, 7. Holotype and paratype workers, ENGLAND: Dorset, Studland, Curlew Dyke, 5.v.1935 (C. Diver) (BMNH). [Synonymy by Brown, 1955.] Distribution. Europe from Britain to the Caucasus and Ukraine, and from northern Europe to Italy. THE HUBERI-Grovup Strongylognathus afer Emery Strongylognathus afer Emery, 1884 : 380, footnote. Holotype female, ALGERIA: Daya (Bedel) (probably in MCSN, Genoa). Distribution. Algeria, Tunisia. Strongylognathus alboini Finzi Strongylognathus huberi subsp. alboini Finzi, 1924b : 121. Syntype workers, ITaty: Venezia Giulia, Monte Nanos, 28.v.1923 (Finzi) (MCZ, Cambridge). Strongylognathus alboini Finzi; Urbani, 1969: 141. [Raised to species.] Distribution. Italy, Switzerland. Strongylognathus alpinus Wheeler Strongylognathus huberi siibsp. alpinus Wheeler, 1909 : 178, figs 1c, d. Syntype workers, females, males, SwITZERLAND: Zermatt, Matter Valley, 1,620 m, 13.viii.1909; same loc. 14.viii.1909; Zermatt, bank of Triftbach R., 18.viii.1g0g (W. M. Wheeler) (MCZ, Cambridge). Strongylognathus alpinus Wheeler; Urbani, 1969 : 140. [Raised to species. ] Distribution. Switzerland. Strongylognathus bulgaricus Viehmeyer stat. n. (provisional) Strongylognathus huberi subsp. vehbinderi var. bulgavica Viehmeyer, 1922: 211. Syntype workers, female, male, BULGARIA: Tirnowa (F. Schimmer) (probably in MNHU, Berlin). ANT TRIBE TETRAMORIINI 307 Strongylognathus vehbindert subsp. bulgaricus Viehmeyer; Pisarski, 1965 : 515. [Raised to subspecies. ] Strongylognathus kratochvili Silhavy, 1937: 5, figs 1-6. Syntype workers, female, CzEcHO- SLOVAKIA: W. Moravia, near Mohelno, 21.iii.1936 (V. Silhavy) (location of types not known). [Synonymy by Pisarski, 1965.] Distribution. Bulgaria, Czechoslovakia. Strongylognathus caeciliae Forel Strongylognathus caeciliae Forel, 1897 : 132. Holotype female, Spain: central Spain, Pozuelo (M. Cabrera y Diaz) (probably in MHN, Geneva). [See also Urbani, 1969.] Distribution. Spain. Strongylognathus cecconii Emery Strongylognathus huberi subsp. vehbinderi var. cecconii Emery, 1908 : 24. Syntype workers, Iraty: Isole Tremiti, S. Nicola, Caprara (MCZ, Cambridge; probably also in MCSN, Genoa). Strongylognathus cecconit Emery; Urbani, 1969 ; 153. [Raised to species. ] Distribution. Italy. Strongylognathus christophi Emery Strongylognathus christophi Emery, 1889a : 439. Lectotype female, U.S.S.R.: Volga, Sarepta (MCSN, Genoa), by designation of Pisarski, 1965. Distribution. U.S.S.R.: N. shores of Caspian Sea northwards to Urals and vicinity of Kazan. Strongylognathus dalmaticus Urbani Strongylognathus dalmaticus Urbani, 1969: 154, figs 2F, 3F, 4F. Holotype and paratype workers, YuGcostavia: Isola Busi (Archipelago Dalmata), 24.vii.1968 (C. Baroni Urbant) (NM, Basle; MCSN, Verona). Distribution. Yugoslavia. Strongylognathus destefanii Emery (Text-figs 14, 18) Strongylognathus destefanii Emery, 1915¢ : 263. Holotype female, Sicity: Dintorni di Palermo (De Stefani) (probably in MCSN, Genoa). Strongylognathus huberi st. cecconit var. kutteyi Santschi, 1927 : 58. Holotype female, SiciLy: Segesta (H. Kutter) (probably in NM, Basle). [Synonymy by Urbani, 1969.] Distribution. Sicily. 308 B. BOLTON Strongylognathus emeryi Menozzi Stronglognathus emeryi Menozzi, 1921 : 30, fig. 3. Holotype female, Iraty: Calabria, Sambiase (probably in IE, Bologna). Distribution. Italy; Calabria. Strongylognathus foreli Emery stat. n. (provisional) Strongylognathus huberi var. fore Emery, 1909 : 709, fig. 4. Holotype (?) worker, ALGERIA (probably in MCSN, Genoa). [See also Urbani, 1969 : 138; foveli may be worker of afer.] Distribution. E. Algeria. Strongylognathus huberi Forel Strongylognathus hubert Forel, 1874: 71. Syntype workers, SwiTzZERLAND: Fully (MHN, Geneva; BMNH; MCZ, Cambridge). Strongylognathus huberi var. gallica Emery, 1909 : 709. Syntype workers, FRANCE: Marseilles (probably in MCSN, Genoa). [Synonymy by Urbani, 1962.] Distribution. S. France, Switzerland, N. Italy. Strongylognathus insularis Urbani Strongylognathus insularis Urbani, 1968 : 470, figs 38, 39. Holotype and paratype workers, Matta: Comino I., 24.iv.1965 (C. Bavoni Urbani) (NM, Basle; MCZ, Cambridge; MCSN, Verona). Distribution. Malta. Strongylognathus italicus Finzi stat. n. (provisional) Strongylognathus huberi subsp. italica Finzi, 1924a : 14. Holotype female, Iraty: Isle of Elba, iv. 1921 (Finzi) (MCZ, Cambridge). Distribution. Isle of Elba. Strongylognathus kervillei Santschi Strongylognathus kervillei Santschi, 1921 : 113. Syntype workers, TuRKEy: Angora prov. (H. G. de Kerville) (probably in’ NM, Basle). Distribution. Turkey: Angora prov. Strongylognathus koreanus Pisarski Strongylognathus koveanus Pisarski, 1965 : 519, figs 36-39. Holotype and paratype workers, Korea: Myohyang Mts, 9.viii.1959 (B. Pisarski & J. Proszynski) (Institute of Zoology, Warsaw; MCZ, Cambridge). Distribution. Korea. ANT TRIBE TETRAMORIINI 309 Strongylognathus palaestinensis Menozzi Strongylognathus palaestinensis Menozzi, 1933: 75, fig. 5. Holotype female, ‘PALESTINE’: Gebata (Bodenheimer) (probably in IE, Bologna). Distribution. Palestine. Strongylognathus rehbinderi Forel Strongylognathus christophi var. vehbindevi Forel, 1904: 2. Syntype workers, U.S.S.R.: Caucasus, Neu Athos (= Novyj Afon) (Rehbinder) (MHN, Geneva; MNHU, Berlin). Strongylognathus vehbinderi Forel; Pisarski, 1965 : 512. [Raised to species. ] Distribution. Caucasus, E. shores of Black Sea. Strongylognathus ruzskyi Emery stat. n. (provisional) Strongylognathus huberi subsp. ruzskyi Emery, 1909 : 711, fig. 6. Holotype worker, U.S.S.R.: Ural, Inderskaia-Gebirge (probably in MCSN, Genoa). Distribution. Urals. Strongylognathus silvestrii Menozzi Strongylognathus silvestvii Menozzi, 1936 : 292, fig. 14. Syntype workers, female, GREECE: Isole Rodi (= Rhodes), Mt Attairo, 800m (= Mt Attaviros); Cattavia (probably in IE, Bologna). Distribution. Rhodes I. TELEUTOMYRMEX Kutter (Text-figs 24-26) Teleutomyrmex Kutter, 1950 : 82. Type-species: Teleutomyrmex schneideri Kutter, loc. cit.; by original designation. Diacnosis. Female. Mandibles blade-like, edentate except for the acute apical tooth. Palp formula 1,1; the maxillary palp with a second prominence standing beside it, resembling a second one-segmented palp. Antennae Io- or 11-segmented, the second to fourth funicular segments showing a variable degree of fusion. In specimens where antenna is distinctly to-segmented then the second funicular is an elongate fusion-segment. Clypeus shallowly transversely concave. Ocelli present. Alitrunk with flight sclerites, winged in virgins; the alitrunk compressed from side to side; metapleural glands absent. Petiole sessile, postpetiole very broadly attached to first gastral segment. Gaster very strongly dorsoventrally flattened, the ventral surface concave; mature females physogastric. Sting very reduced, non-functional. Male. Mandibles as female but much smaller. Palp formula 1,1. Antennae 10-segmented, the second funicular antennomere an elongate fusion-segment. Ocelli present; wings present but the remainder of the body appearing pupoidal. Gaster downcurved, somewhat reflexed anteriorly. This genus contains a single species, T. schneideri, a degenerate workerless, permanent social parasite in nests of Tetramoriwm caespitum. The females of 310 B. BOLTON schneideri are to be found riding upon the queen of the host colony, which is not killed. The males, as in Anergates, are pupoidal but unlike that genus the males of schneidert possess wings. As yet the species has not been recorded outside the nests of its host-species. Apart from the original description, which contains a good deal of information, studies dealing with the biology, behaviour and anatomy of T. schneideri include those of Stumper (1950), Gdsswald (1952), Kutter (1969), and a summary in Wilson (1971). Teleutomyrmesx schneideri Kutter Teleutomyrmex schneideri Kutter, 1950 : 82, figs 1-23. Syntype females, males, SwITZERLAND: Saas-Fee, 25.vii.1949 (H. Kutter) (in coll. Kutter; BMNH). The species is known only from the type-locality and from a second collection made near Briangon in the French Alps (Collingwood, 1956). TRIGLYPHOTHRIX Forel (Text-figs 32-73) Triglyphothrix Forel, 1890 : cvi. Type-species: Triglyphothrix walshi Forel, op. cit.: cvii; by monotypy. Diacnosis. Worker. Mandibles with three teeth apically, the third smaller than the second, and followed by a row of three or four small or minute denticles; the arrangement 3 teeth + 4 denticles predominating (Text-fig. 36). Palp formula 4,3 (dissections of lanuginosa, walshi, fulviceps, meshena, kheperra, brevispinosa, arnoldi, constanciae, eminti, gabonensis, vothschildi, inezulae), often reduced to a 3,3 formula in minute species (dissections of cryptica, muscorum, gestroi, furtiva). Antennae with 10, II or 12 segments, the apical three forming a distinct club. The 12-segment count predominates, only three species are known with 11 and only one species (decamera) has to antennal segments, but this is known only in the female caste at present. Eyes present, at or in front of the middle of the sides of the head, usually large, rarely reduced. Frontal carinae very broad, the maximum span of the lobes often being more than twice the distance from the edge of the lobe to the side of the head in a straight line. Clypeus convex or angled in profile, with a median longitudinal carina extending its length (weak in some species). Median carina present along length of cephalic dorsum, a continuation of the median clypeal carina and usually very distinct, only in some heavily sculptured species is its track vague. Antennal scrobes usually present, broad and divided into upper and lower compartments by a longitudinal carina or ridge. In a few species the scrobes are reduced but very rarely are they completely absent (Text-figs 42, 44, 45). Alitrunk fusiform, without sutures; metanotal groove absent, the outline of the alitrunk unbroken in profile (Text-figs 52-72). Propodeum usually armed with a pair of spines or teeth, unarmed only in a very few species. Metapleural lobes present, often acute or dentiform. Petiole pedunculate, without a large ventral process. In species of the Ethiopian region there is a marked tendency for the petiole and postpetiole to be expanded laterally, transverse in dorsal view. Apicodorsal appendage of sting triangular, dentiform or pennant-shaped. A simple tibial spur is generally present on the middle and hind legs in large species, absent in smaller forms. Sculpture is usually coarse on head and alitrunk, the predominant sculpture being a dense rugoreticulum, but this differs in some species. Pilosity dense, at least some part of the head or body bearing regularly branched (bifid, trifid or quadrifid) hairs, either with or without simple hairs also being present (Text-figs 32-35). Branched hairs are absent in only one species, which is suspected of a cleptobiotic lifeway. ANT TRIBE TETRAMORIINI 311 Female. Generally as worker but virgin females fully alate, with flight sclerites and ocelli. Male. Known for only a few species. With the general tetramoriine characters and also with branched hairs as in the workers and females. Antennae 1o-segmented, the second funicular forming an elongate fusion-segment (Text-fig. 40). Palp formula 4,3 in the two species dissected (constanciae, inezulae), as worker. Whether the smaller species show the reduced palp formula of 3,3 in males as well as workers remains to be seen. Triglyphothrix is a moderately sized genus containing some 55 presently recognized species, of which 33 occur in the Ethiopian region and the remaining 22 are distributed throughout the rest of the Old World tropics and subtropics, with the exception of Australia and Madagascar where only lanuginosa is present. This species is well known throughout the world as a very successful tramp species, and on occasion it is reported from hothouses or constantly heated buildings in temperate zones, as well as from localities in the tropics which are outside the range of the genus as a whole. As far as is known, most species are predominantly cryptic foragers of the forest floor, although some have extended their range into savannah or open grass or woodland. One species, arnoldi, is apparently cleptobiotic in nests of large ponerine ants, nesting in the walls of their galleries and tunnels, and microps has also been recorded from a similar locality (Arnold, 1917). Apart from these the remaining species nest either directly into the soil, when a small crater-entrance may be produced, or under stones and logs, or in twigs or rotten wood in the leaf-litter layer. In some species the nests are made in twigs anywhere in the litter, but in others a definite site at the base of a standing tree appears to be preferred. No species are known to be arboreal or subarboreal in habits. Most, if not all, species forage singly and move slowly, the more densely hairy species presenting a furry appearance to the naked eye. They are general pre- dators and scavengers, collecting fresh or dead arthropods, but are not known to tend homopterous insects nor to visit plant nectaries. Previous studies of the genus have dealt only with a limited number of species, on a regional basis. These include the work of Arnold (1917; 1926) on the South African fauna, Emery (1893), Forel (1902) and Bingham (1903) on the Indo- Malayan fauna, and Mann (1919) on the New Guinea and Solomon Islands species. The main outstanding feature of the genus as a whole is the possession of numerous or abundant branched hairs. These may be bifid, trifid or quadrifid and in a few species a mixture of two or more of these may be present. In many species simple hairs are also present on the dorsal surfaces, generally far fewer in number than their branched counterparts and in some species seeming to serve a specialized function as they occur only upon the leading edges of the antennal scapes or the dorsal surfaces of the upper margins of the antennal scrobes in a spaced row. The majority of species, however, have simple hairs upon the mouth- parts and around the gastral apex, although in a few these are absent. Only one species in the genus has lost its branched hairs (avnoldi) and I am sure that in this case it is a secondary development as branched hairs are present in the closest relatives. 312 B. BOLTON SPECIES OF THE ETHIOPIAN REGION SYNONYMIC LIST OF SPECIES paupera-group antrema sp. n. cryptica sp. n. distincta sp. n. menkaura sp. n. minima sp. n. muscorum (Arnold) paupera Santschi evicae (Arnold) syn. n. Osiris-group osiris sp. n. reptana sp.n. yarthiella sp. n. inezulae-group arnoldi Forel desertorum Forel furtiva Arnold gestroi Menozzi imbellis Emery inezulae Forel hepburni (Arnold) syn. n. hepburni subsp. mashonana Arnold syn. n. microps Mayr auvopunctatus Forel syn. n. auvopunctatus var. pallens Forel syn. n. auvopunctatus var. fusciventris Forel syn. n. auvopunctatus var. rhodesiana Forel syn. n. auvopunctatus var. bulawayensis (Arnold) syn. n. talpa sp. n. tenebrosa (Arnold) stat. n. trimeni Emery rothschildi-group rothschildi Forel gabonensis-group agna Santschi baufra sp. n. brevispinosa Stitz stat. n. mion Bernard syn. n. constanciae (Arnold) longispinosa Arnold syn. n. dedefra sp. n. emenii Forel marthae Forel syn. n. marleyt Forel syn. n. marleyi var. akermani (Arnold) syn. n. guillodi Santschi syn. n. guillodi var. mus Santschi syn. n. cinereus Weber syn. n. gabonensis André gabonensis st. soyauxt Forel syn. n. ANT TRIBE TETRAMORIINI 313 mucida Forel syn. n. areolata Stitz syn. n. gabonensis var. boulognei Forel syn. n. gabonensis var. khamerunensis Santschi syn. n. aveolata var. burgeoni Santschi syn. n. monardi Santschi pulcherrima Donisthorpe sericea (Arnold) stat. n. silvestrii Emery thoth sp. n. THE SPECIES-GROUPS THE paupera-GRouP. This group contains seven very small to minute species, all of which possess elongate simple hairs which project in a row along the anterior edges of the scapes and along the upper borders of the frontal carinae which form the dorsal margins of the scrobes. Hairs on the first gastral tergite are predomi- nantly a mixture of simple and bifid, sometimes with a few trifid, hairs present but never with dense trifid or quadrifid pilosity. Three of the species of this group (antrema, cryptica, muscorum) have only II antennomeres and have the palp formula reduced to 3,3 from its normal 4,3 value. The other four species have the normal antennomere count of 12. Most of the species are of west and central African origin, more or less confined to the rain forest zones, but muscoruwm occurs virtually throughout Africa where there is wet or riverine forest, and paupera is known only from Rhodesia. All seven species are denizens of the leaf-litter layer in forests. THE osivis-GRoUP. The three species of this small group show relationship to the paupera-group in that they possess a row of elongate simple hairs both on the leading edges of the antennal scapes and on the upper surfaces of the frontal carinae. They are, however, considerably larger than paupera and its allies and have the first gastral tergite covered with a dense mat of branched hairs which are predominantly or universally quadrifid. The members of the group are only known from their type-localities, but the group as a whole shows a wide distribution, with osivis from Ghana, veptana from Uganda and yarthiella from Zambia, THE inezulae-GRouP. Ten species are known in this group; they are characterised by the lack of long simple hairs on the antennal scapes and their presence in a row on the upper margins of the frontal carinae. Also in this group there is a tendency for the node of the petiole to be strongly compressed antero-posteriorly so that it is transverse in dorsal view. This group is predominantly of eastern and southern Africa, with only a few species extending their range into the central and west African forest zones. A number of complexes of closely related species can be discerned within the group. One complex, consisting of arnoldi, imbellis and microps, has the eyes very reduced, their maximum diameter being distinctly less than the maximum width of the scape (in all other African species the eyes are larger than the maximum scape 314 B. BOLTON width). This complex occurs mainly in southern Africa but imbellis is found in Ethiopia. The species desertorum, inezulae, talpa and trimeni possess sculpture on the basal portion of the first gastral tergite, and these species range from Zaire to South Africa. The two smallest species of the group, furtiva and gestroi, occur in forested areas in deep leaf-litter. THE vothschildi-GRoup. Only a single species, vothschildi, is placed in this group and it is certainly the most aberrant member of the genus yet known from the Ethiopian region. The species is large, antennal scrobes are completely absent, and the anterior clypeal margin has a deep median emargination. T. rothschildi is widely but sporadically distributed in dry or semi-desert localities in Africa and appears to be the only African species in this genus to be adapted to such conditions. THE gabonensis-GRrouP. The largest species-group in the Ethiopian region, with 12 members, it also contains two of the most common species of the region. All members of this group are characterised by the lack of elongate simple hairs both on the scapes and on the upper surfaces of the frontal carinae. The body is usually stocky and abundantly covered with branched trifid or quadrifid hairs, giving the ant af urry appearance to the naked eye. Members of this group occur throughout the Ethiopian region, the preferred nesting sites of the species being in rotten wood or directly into the earth. The species agna and eminii are closely related and have the antennal scrobes reduced and non-functional. Of the remainder baufra, brevispinosa, constanciae and gabonensis are quite large, active, relatively conspicuous species; the rest are smaller and tend to be more cryptic, inhabiting leaf-litter or, in the case of pulcherrima, sandy areas. KEY TO SPECIES (Workers) Note. The worker of desevtorum is not known but the queen is very close to that of inezulae. The worker, when found, would therefore be expected to key out at couplet 17. 1 Antennae with 11 segments . : : : : - c : : : ® — Antennae with 12 segments 2 Dorsum of head and alitrunk finely pac douecth: Pe eeatate aepalese — Dorsum of head with a few fine, scattered longitudinal rugulae with eee Be. spaces, alitrunk with -scattered fine wcisaiiaind rugulae, not reticulate-rugulose (Cameroun) ; . antrema (p. 318) 3 Clypeus with a med Rees carina at the et of the Serene anterior portions of the frontal carinae, the transverse carina as strongly developed as the median longitudinal carina which it crosses. Head and alitrunk yellow or yellow-brown. (Ivory Coast, Ghana, Cameroun, Uganda, Zaire, Gabon, C. African Repub., Ww Angola, S. Africa, Mozambique) . ; muscorum (p. 331) — Clypeus without a transverse carina eee the eter longitudinal carina. Head and alitrunk black or blackish brown. (Ghana) . : . cryptica (p. 321) 4 With the head in profile the antennal scrobe is either very short and shallow, without a sharp dorsal margin and not divided into upper and lower portions by a longitudinal carina, or is completely absent (Text-figs 42, 44, 45). The area n 12 ANT TRIBE TETRAMORIINI 315 of the scrobe is generally as strongly sculptured as the remainder of the head and always has hairs arising within the scrobal area e 5 With the head in profile the antennal scrobe developed, long and ao reaching well beyond the posterior margin of the eye and with a sharp dorsal margin. Scrobe less strongly sculptured than the remainder of the head (except when head itself is unsculptured); without hairs arising within the scrobal area and generally divided into upper and lower portions by a median carina, at least anteriorly (Text-figs 43, 46-51) .. . 3 : : : , F 7 Propodeum unarmed, without spines or teeth, (Zaire) : ; : agna (p. 317) Propodeum armed with a pair of spines or teeth 6 Anterior border of clypeus broadly and deeply emarginate medially (Text. fig. oN Basal portion of first gastral tergite finely, closely and densely longitudinally striate. Very large species, HW > 0-95, SL > 0:60. (Arid zones of Ghana, Ethiopia, Kenya) : - rothschildi (p. 334) Anterior border of clypeus ASE a puareiaate made (Text-fig. 44). Basal portion of first gastral tergite unsculptured or with a few pits from which hairs arise, not densely striate. Smaller species, HW < 0°85, SL < 0-60. (Ethiopia, Sudan, Nigeria, Kenya, Tanzania, Angola, S. Africa) . eminii (p. 324) Head, alitrunk, pedicel and gaster without branched hairs, all hairs simple and short. Dorsum of alitrunk finely and very densely reticulate-punctate, matt. [Presumed cleptobiont in nests of large ponerine ants.] (Rhodesia) . arnoldi (p. 318) At least part of head or body with branched hairs, bifid, trifid or quadrifid; addi- tional simple hairs may be absent or present. Dorsum of alitrunk not densely reticulate-punctate : 8 Dorsal margins of frontal carinae which form the upper borders of the antennal scrobes behind the antennal insertions with a spaced row of elongate simple hairs which are distinctly longer than the surrounding (usually branched) hairs (Text- fig. 37). These long hairs are often erect but in some species are directed laterally; rarely similar si hairs may be pent elsewhere on the dorsum of the head also : : 9 Dorsal margins of frontal carinae which form the mpper borders of the antennal scrobes behind the antennal insertions either without a spaced row of elongate simple hairs or very rarely with a few simple hairs which are very short and do not project above the level of the surrounding branched hairs, not easily seen . 23 Propodeum without spines or teeth, with a pair of minute blunt tubercles which are much shorter than the metapleural lobes (Text-fig. 55) ; 10 Propodeum armed with a pair of spines or acute teeth which are usually longer than the metapleural lobes . II Dorsal surfaces of head and alitrank unsculptured apart from the median carina on the head and a few scattered punctures. (Ghana) : : menkaura (p. 328) Dorsal surfaces of head and alitrunk distinctly rugulose. (Ethiopia) . imbellis (p. 327) Hairs on anterior (leading) edge of antennal scape of varying length, the longest being simple and at least as long as the maximum width of the scape. If the longest hairs are subequal to the maximum scape width then elongate simple hairs are present on the dorso-lateral pronotal margins which are distinctly longer and more conspicuous than the neighbouring branched hairs . : 12 Hairs on anterior (leading) edge of antennal scape of uniform length and eeacrathy branched, all distinctly much shorter than the maximum width of the scape, often less than half the maximum scape width and sometimes closely applied to the surface . 17 Strongly bicoloured spedies: ends ae aut eee, also medical black alee legs, antennae and mandibles clear yellow. (Ghana) F : distincta (p. 323) Either uniformly coloured species or with the gaster somewhat darker than the head and alitrunk, but not bicoloured black and yellow as above 5 : 3 13 316 14 15 16 ny 18 19 20 21 22 B. BOLTON Clypeus with a raised transverse carina at the level of the extreme anterior portions of the frontal carinae, the transverse carina as strongly developed as the median longitudinal clypeal carina which it crosses. (Ivory Coast, Ghana, Nigeria) minima (p. Clypeus without a transverse carina crossing the median longitudinal carina. - First gastral tergite basally with a. broad belt of coarse punctate sculpture which extends approximately over the basal one-third of the sclerite. (Zambia) yarthiella (p. First gastral tergite basally without a belt of punctate sculpture, smooth and shining Longest hairs on tessa of Hiead pod alypeus (discoantiag those on. aipper serobe margins) distinctly shorter than the maximum vertical width of the eye with the head in profile. Small species, HW < 0:50, SL < 0:35. (Rhodesia) paupera (p. Longest hairs on dorsum of head behind clypeus (discounting those on upper scrobe margins) distinctly longer than the maximum vertical width of the eye with the head in profile. Larger species, HW > 0:50, SL > 0°35. Apical halves of mandibles longitudinally striate. First gastral terpite with a mixture of simple and bifid hairs, quadrifid hairs absent. Propodeal spines in profile not downcurved along their length. (Cosmopolitan tramp species) lanuginosa (p. Apical halves of mandibles smooth and shining, with scattered small pits. First gastral tergite with hairs predominantly quadrifid. Propodeal spines in profile narrow and slightly downcurved along their length (Text-fig. 61). (Ghana) Osiris (p. Eyes very small, their maximum diameter distinctly less than the maximum width of the antennal scape (Text-fig. 43). (Angola, Rhodesia, S. Africa) . microps (p Eyes larger, their maximum diameter at least equal to the maximum width of the antennal scape, usually distinctly greater Base of first gastral tergite immediately behind the postpetiole with « a trcinswexse band of dense and quite coarse punctate sculpture; this area distinctly more coarsely sculptured than the remainder of the tergite : Base of first gastral tergite without a band of dense punctate sculpture fee behind the postpetiole. Usually this area as smooth as the remainder of the tergite but if some faint shagreening is prescm then the species is ei with SIU 0-50). : Hairs on first gastral tergite almost eaticely simple, with only one or ia bifid or trifid hairs present. Colour uniform pale yellow-brown. (Zaire) . talpa (p. Hairs on first gastral tergite universally branched, bifid, trifid, without simple hairs present. Colour uniform medium to dark brown : : - : Lower portion of antennal scrobe sculptured above the eye. The longitudinal carina which divides the scrobe into upper and lower portions indistinct, appear- ing only as a boundary to or continuation of the sculptured area beneath it. Smaller species, SL 0-34 or less. (S. Africa) . : : . trimeni (p. Lower portion of antennal scrobe not or only very faintly Sealpenned above the eye, the longitudinal carina dividing the scrobe into upper and lower portions sharp and distinct. Larger cee SL 0-38 or more. pe Tanzania, Rhodesia, S. Africa) . . . inezulae (p. Large species, HW o-80 or more, SL O: =e or more, PW o 60 or more. (Kenya, Mozambique) . : : : tenebrosa “ie Small species, HW < o. es SIL, < O- ae, PW 83. (Solomon Islands) antennata (p. Propodeum with a pair of minute, obtuse tubercles which are much shorter than the metapleural lobes (Text-fig. 69). Antennal scapes relatively shorter, SI < 80. (Java) : 5 . brevidentata (p. Basal portion of first gastral tergite Toabitudinally striate or costulate, sometimes with scattered punctures between the striae : Basal portion of first gastral tergite punctate or eeeeiaes ae ely coarsely so; striation or costulation absent Head bright yellow, contrasting strongly with the Peeece Brows alteonls pedicel and gaster. (New Guinea) . fulviceps (p. Entirety of head and body yellow, the sliecuolc eenerdlly slightly darker in shade than the head and gaster. (Solomon Islands) : : pulchella (p. Smaller species with narrower head and relatively ioe antennal scapes, HW < 0-60, CI ca 87-90, SI in range 77-81. Upper boundary of antennal scrobe without an acute, narrow, flange-like margin formed by the frontal carinae. Median longitudinal carina of scrobe very weak or absent, not more strongly developed than the surrounding sculpture. Colour yellow. (Solomon Islands) ‘ : : . mayri (p Larger species with oe heads md eee peer ‘antennal scapes, HW > 0-60, CI ca 94-97, SI in range 61-68. Upper boundary of antennal scrobe with a distinct, acute, narrow flange-like margin formed by the frontal carinae. Median longitudinal carina of scrobe strongly developed, easily dis- tinguished from the surrounding sculpture. Colour red-brown. Propodeal spines long and narrow, much longer in profile than their Te width (Text-fig. 66). Trifid hairs on disc of first gastral tergite and on head short and closely applied to the surface from which they arise. (Sulawesi) adpressa (p. Propodeal spines short, stout and broadly triangular, in profile about as long as their basal width (Text-fig. 67). Trifid hairs on disc of first gastral tergite and on head long and erect or suberect, not —— snus to the surface from which they arise. (West Malaysia, Java) c : meshena (p. Antennal scapes exceptionally long, SL > 0-75, SI 100 or more; antennal scrobes completely absent.. (Vietnam) . F . indosinensis (p. Antennal scapes much shorter, SL < o. 65, SI < ae antennal scrobes most commonly present, only rarely vestigial or absent Node of petiole in dorsal view strongly antero-posteriorly sq eee ee distinctly broader than long. (India, Sri Lanka, China, Singapore, Philippines) walshi (part) (p. Node of petiole in dorsal view not antero-posteriorly compressed, not transverse, generally as long as broad or very slightly broader than long : Hairs on first gastral tergite universally trifid and forming a dense pelt over the sclerite; simple or bifid hairs completely absent from the tergite Hairs on first gastral tergite universally simple, or bifid, or a mixture of cule and branched hairs . 352) 13 15 18 19 20 21 22 ANT TRIBE TETRAMORIINI 343 With the petiole in profile the tergal portion of the node distinctly mae than high (Text-fig. 70) ; 14 With the petiole in profile the ‘tergal povtion af the node af most as long as hich, usually distinctly higher than long (Text-figs 64, 71) : 15 Dorsal margin of frontal carinae forming the border of the eerobe with a spaced row of long, simple hairs which are very prominent. (Philippines) . katypa (p. 349) Dorsal margin of frontal carinae forming the border of the scrobe with abundant short, branched hairs but without a spaced row of long, PARE hairs. (West Malaysia) . . pnyxis (p. 356) Propodeal spines distuctly loneer than the eeaiegral lobes. "Tins, Vietnam, Hong Kong, Philippines, Borneo, Malaya, Java, Sumba, Sulawesi; introduced in temperate zone with plant products) . : 3 kheperra (p. 349) Propodeal spines shorter than metapleural lobes, at most the spines almost as long as the lobes : 16 Larger species, with relatively shorter antennal scapes, “HW - 0-70, SI << 70. (Thailand) . ; nacta (p. 353) Smaller species with relatively longer antennal scapes, “HW 2 0-65, SI > 7o. (Borneo, Philippines, Taiwan) : : parvispina (p. 355) Dorsal (outer) surface of hind tibiae voect frome j in front or bee with numerous long hairs of varying length, the longest of them at least subequal to the maxi- mum tibial width; these hairs never forming a close mat above the tibial surface (Text-fig. 38) . : 18 Dorsal (outer) surface of hind abi viewed fen in seit or Bébiliia ae or short, curved hairs of approximately uniform length which are much shorter than the maximum tibial width, usually less than half the tibial width; these hairs characteristically forming a close mat above the tibial surface (Text-fig. 39) 20 First gastral tergite with all hairs simple. (India) . - coonoorensis (p. 346) First gastral tergite with at least a few bifid or trifid hairs present : : : 19 Mandibles smooth with scattered punctures, not longitudinally striate. HW in range o-76—-0-82. First gastral tergite with trifid hairs on basal half. (India, Burma) : : ; obesa (p. 354) Mandibles ieeecanely shite! the Sereiaan aioe. meting or absent in small specimens. HW in range 0-52-0-70. First gastral tergite basally with a mixture of simple and bifid hairs, trifid hairs usually completely absent. (Throughout Oriental and Indo-Australian regions; cosmopolitan tramp species) lanuginosa (p. 350) Dorsal surfaces of both petiole and postpetiole reticulate-rugose, the two about equally densely sculptured . 21 Dorsum of postpetiole mostly smooth amd shining ies onky vestigial ‘stulpture in places, distinctly less densely sculptured than the petiole dorsum. (Fiji Islands) pacifica (p. 255) First gastral tergite with hairs universally simple or at most with only 1-2 bifid hairs. Antennal scrobes strongly developed, with an acute dorsal margin . 22 First gastral tergite with a distinct mixture of simple and bifid hairs. Antennal scrobes vestigial, without an acute dorsal margin. (India) : : rossi (Pp. 357) Node of petiole in profile massive, its outline almost square, and with well developed antero- and posterodorsal angles (Text-fig. 72). Larger species with HW in range 0-72—0-76, SL in range 0-56—0-58. (Borneo) . : : chepocha (p. 345) Node of petiole in profile relatively low and convex above, its outline not square, and with the dorsal surface rounding into the posterior face so that there is no defined posteroventral angle (Text-fig. 73). Smaller species with HW in range 0-64-0-66, SL in range 0-46—-0-50. (Solomon Islands) : ; . vombis (p. 358) 344 B. BOLTON TREATMENT BY SPECIES Triglyphothrix adpressa sp. n. (Text-fig. 66) Holotype worker. TL 3:1, HL 0-72, HW 0-70, CI 97, SL 0-44, SI 63, PW 0-52, AL 0-80. Sides of head convex, occipital margin shallowly concave in full-face view. Antennal scrobes strongly developed, with a distinct dorsal margin and a marked median longitudinal carina which is well differentiated from the surrounding sculpture and which reaches back beyond the posterior margin of the eye. Propodeal spines long and acute, quite narrow, distinctly longer than their basal width in profile. Metapleural lobes strongly developed, acute, feebly upcurved (Text-fig. 66). Pedicel in dorsal view with the postpetiole very slightly broader than the petiole (about 0-02 difference) and the petiole node slightly broader than long (about 0:04 difference). Mandibles striate; head, alitrunk, petiole and postpetiole densely punctate with scattered rugulae, base of first gastral tergite densely punctate. Pilosity on dorsum of body entirely of short trifid hairs of approximately equal length, which are closely adpressed on the head and gaster; long simple hairs completely absent from dorsal surfaces and from appendages. Hairs on scapes and tibiae short, distinctly less than half the maximum width of the surfaces on which they arise. Paratype workers. TL 2-8-3-1, HL 0-70-0-74, HW 0-66-0-72, CI 94-97, SL 0:42-0-44, SI 61-64, PW 0:50-0:54, AL 0:74-0:80 (3 measured). As holotype. Holotype worker, SULAWEsI (‘N. Celebes’ on label): SW. slope Mt Klabat, 400-600 m, 13-19.vi.1972, rain forest (W. L. Brown) (MCZ, Cambridge). Paratypes. 3 workers with same data as holotype (MCZ, Cambridge, BMNH). This species is very closely related to meshena and I would not be surprised if further collections from Indonesia were to lead to the conclusion that the two are inseparable. For the present, however, I consider that they should be separated on the grounds that in adpressa the cephalic and gastral hairs are very short and strongly adpressed, the propodeal spines are long, narrow and acute and the metapleural lobes are narrower and upcurved. In meshena the cephalic and gastral hairs are longer and are erect or suberect, the propodeal spines are short and broad and the metapleural lobes are broader and scarcely or not upcurved (compare Text-figs 66 and 67). Other closely related species include parvispina and the widespread kheperrva, but in both these species the basal portion of the first gastral tergite is unsculptured. Triglyphothrix antennata Mann stat. n. Triglyphothnix fulviceps subsp. antennata Mann, 1919 : 350, fig. 30. Syntype workers, SoLtomon IsLanpbs: Santa Cruz, Graciosa Bay, 1916 (W. M. Mann) (MCZ, Cambridge) [examined]. Worker. TL 3:0-3:2, HL 0:66-0:68, HW 0:56-0:60, CI 85-89, SL 0:50-0:52, SI 87-89, PW 0:46, AL 0-76—0-78 (2 measured). Antennal scrobes poorly developed, with a weak dorsal border which is only differentiated to about the level of the posterior margin of the eye when viewed in profile. Behind the level of the eye it tends to merge into the surrounding sculpturation, which is moderately strongly developed on the sides of the head behind the eyes. Median longitudinal carina of scrobe absent or so weakly developed that it is indistinguishable from other longitudinal rugulae above the eye. Antennal scapes long, the SI of 87-89 being approached only by mayrvi and ANT TRIBE TETRAMORIINI 345 pacifica which have a maximum SI of about 80. Mandibles striate, the dorsal surfaces of head, alitrunk and pedicel finely reticulate-rugose, the basal portion of first gastral tergite strongly punctate. Pilosity everywhere predominantly of fine, dense, simple hairs which are of varying length on the head and dorsal alitrunk. On the scapes and legs the hairs are shorter and more or less uniform in length; long, erect projecting hairs are absent. Head and gaster yellow-brown in colour, the alitrunk and pedicel darker. Along with its allies brevidenata, pacifica and vombis this species shows a marked tendency towards the loss of branched hairs. This tendency is obviously better developed in antennata than in the other species as here branched hairs appear to be almost completely absent whilst at least a few are retained in the related species. Despite this antennata is still very densely hairy, the individual hairs fine and generally curved. Identification of the species should be relatively simple by the use of the combination of characters given in the diagnosis above. Triglyphothrix brevidentata Kutter (Text-fig. 69) Triglyphothrix brevidentata Kutter, 1932: 208, figs. Holotype worker, Java: Sarangan, Lawoe-Geb. 1927 (H. Overbeck) (Dresden Museum). Worker. TL 2-6—2-:7, HL 0:62-0:64, HW 0:-54-0°58, CI 88-91, SL 0-40-0-42, SI 72-75, PW 0:44-0°46, AL 0-68-0-70 (4 measured). Antennal scrobes developed but weakly so, the median longitudinal carina scarcely more strongly developed than the surrounding sculpture. Propodeal spines reduced to a pair of minute, obtusely-rounded tubercles above the broad, triangular, acute metapleural lobes. Mandibles striate. Dorsal surfaces of head, alitrunk and pedicel finely reticulate-rugulose. Extreme basal portion of first gastral tergite sparsely punctate, the spaces between punctures usually finely shagreened; remainder of gaster smooth and shining. Branched hairs sparse on head and alitrunk, more numerous on petiole and postpetiole, apparently absent on gaster. Long, simple hairs numerous on dorsal surfaces of head and body but scapes and tibiae with only short, curved hairs. Colour brown to reddish brown, the appendages lighter. Of the species known from the regions at present under consideration only two have the propodeal armament strongly reduced, brevidentata and parvispina, but in the latter the propodeal spines are still present as small, acute teeth, whereas in the former only low, rounded tubercles are present. Also, parvispina has abundant branched hairs upon the gaster whilst in brevidentata all gastral hairs appear to be simple. Finally, the base of the gaster is sculptured in brevidentata and is not in parvispina. Within the obesa-group brevidentata appears closest related to antennata but the latter has strongly developed propodeal spines and relatively much longer antennal scapes. Apparently brevidentata is only known from Java, where it has been collected twice in my knowledge, besides the type-collection. Both these collections were made at Tjibodas by Dammerman, in 1921 and 1922. Triglyphothrix chepocha sp. n. (Text-fig. 72) Holotype worker. TL 3:2, HL 0-76, HW 0-72, CI 95, SL 0:56, SI 77, PW 0°58, AL 0:94. 346 B. BOLTON Antennal scrobes shallow, with very feebly defined posterior and ventral margins. Dorsal margin of scrobe more strongly defined and acute, but not a distinct flange or rim as is seen in most species of this group, rather it is an uneven line where the sculpture of the cephalic dorsum terminates at the frontal carinae. Median carina of scrobe absent or not differentiated from the surrounding sculpture. Propodeal spines long and acute, longer than the large, triangular, upcurved metapleural lobes. Node of petiole in profile blocky and massive, its outline almost square, with strongly developed antero- and postero-dorsal angles. In dorsal view the node is about as broad as long, much broader behind than in front and narrowly rounded anteriorly. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely and quite densely reticulate-rugulose. All dorsal surfaces of head and body with numerous long, simple hairs, universally simple on the first gastral tergite. Elsewhere numerous short bifid or trifid hairs are present amongst the more conspicuous long simple hairs. Long hairs such as are present on the head and alitrunk are absent from the legs and antennae where much shorter simple hairs are present with a few bifid individuals. Colour dark red-brown, the appendages lighter, yellow-brown. Paratype workers. As holotype but one or two specimens with a single bifid hair on the first gastral tergite. TL 3-0-3-4, HL 0:74-0:80, HW 0:72-0:76, CI 95-97, SL 0:56-0'58, SI 76-78, PW 0:56-0:60, AL 0:92-0:98 (9 measured). Holotype worker, BorNEO: Kiduk Arok, Trus Madi Massif, 1500 m, Cambridge N. Borneo Exped. no 75 (P. W. Bryant) (MCZ, Cambridge). Paratypes. 11 workers with same data as holotype (BMNH; MCZ, Cambridge). T. chepocha appears to be most closely related to vombis of the Solomon Islands, but this species is smaller and has a differently shaped petiole node, details of which are given in the key. Triglyphothrix coonoorensis (Forel) comb. n. (Text-fig. 65) Tetvamorium coonoorense Forel, 1902a : 237. Syntype workers, female, Inp1a: Coonoor (Wroughton) (MHN, Geneva) [examined]. Worker. TL 2-7-2:8, HL 0:64-0:66, HW 0:56-0'58, CI 87-89, SL 0:44-0:46, SI 77-79, PW 0:44-0-46, AL 0-72—0'74 (4 measured). Antennal scrobes poorly developed and shallow, the dorsal margin formed by the termination of the cephalic rugosity and not by a projecting, flange-like edge. Median longitudinal carina of scrobe not developed. Dorsal outline of alitrunk regularly convex, the propodeal spines long and quite narrow, only slightly tapered from base to apex. Petiole node in profile low, paniform, longer than high, the anterior peduncle of the petiole meeting the node so that its dorsal surface is continuous with the anterior face of the node and the ventral surface meets the ventral surface of the node through an obtuse angle. Dorsum of head, alitrunk and pedicel reticulate-rugulose, the gaster unsculptured. Sculpture within the scrobe basically a continuation of the rugulation present on the remainder of the head but weaker and with a tendency for it to be replaced by dense puncturation, especially in the upper half of the scrobe. Long, simple hairs present on all dorsal surfaces and on appendages. Branched, bifid hairs sparse everywhere, best developed on dorsal alitrunk and sides of petiole, completely absent from first gastral tergite. Colour light brown to mid-brown, the head usually distinctly darker than the alitrunk. Closely related to obesa and lanuginosa but separated from them by the absence of branched hairs from the first gastral tergite; such hairs are numerous or abundant in the related species. ANT TRIBE TETRAMORIINI 347 Apart from the type-series of three workers and a female the only other material of this species examined comes from a short series collected by Bingham, presumably from India but without further data, and the series noted below. MATERIAL EXAMINED. Inp1A: Nilgiri, Madras, Halical nr Coonoor (Besuchet, Lobl, Mussard). Triglyphothrix decamera Forel Triglyphothrix decamera Forel, 1902a : 240. Holotype female, INDIA: Kanara (Aitken) (MHN, Geneva) [examined]. Holotype female. TL 2-9, HL 0-60, HW 0:54, CI 90, SL 0:34, SI 63, PW 0:56, AL 0°82. Although only known from the type-collection of a single female this species should be easily recognised if found again as the antennae have only 10 segments and the propodeum is completely unarmed, without trace of teeth. The petiole is distinctly narrower than the postpetiole in dorsal view, their respective widths ca 0-20, 0-30. Pilosity everywhere is long, erect, and consists of a mixture of simple and bifid hairs, with the former tending to predomi- nate. Head, alitrunk and pedicel covered with fine, superficial rugosity, the gaster shining, unsculptured. The form of the pilosity and the presence of long, erect simple hairs projecting from the tibiae and antennal scapes relates decamera to obesa and its allies, but in all other species of the obesa-group the female has 12-segmented antennae as in the worker and the propodeum is armed with spines or teeth. The possibility that decamera is a temporary or permanent social parasite should not be overlooked, and if this is the case then it is also possible that the workers (if any) do not share the singular characters of the female, as Brown (1964) has pointed out in his study of Rhoptromyrmex, where the queens are quite different from the workers of the same species and are assumed to be social parasites. Triglyphothrix fulviceps Emery (Text-fig. 62) Triglyphothrix fulvicepbs Emery 1897 : 588. Syntype workers, New GuINEA: Friedrich- Welhelmshafen (L. Bivd) (MCSN, Genoa) [examined]. Note. A number of specimens in MHN, Geneva are labelled as cotypes (= syntypes). These were collected in New Guinea by Biro but in 1898-99 and are thus not types. The localities of these specimens are Huon Golf, Sattelberg and Astrolabe Bai, Stephansort. Worker. TL 2-5-3:0, HL 0:60-0:70, HW 0:52-0:64, CI 84-91, SL 0°34-0°46, SI 63-72, PW 0:42-0:52, AL 0:62-0°80 (15 measured). Antennal scrobes distinct, with a weakly developed median longitudinal carina which in some specimens runs the length of the scrobe. Propodeal spines long and acute, varying in length between individuals but always much longer than the metapleural lobes which are somewhat reduced and rounded. Node of petiole low in profile, its anterior face very short, merging into the peduncle. In dorsal view the node broader than long. Dorsal surfaces of head, alitrunk and pedicel closely and very finely and densely reticulate-rugose; mandibles striate. Basal half or more of first gastral tergite strongly longitudinally striate, the spaces between the striae often with punctures, especially basally. First gastral sternite sculptured. 348 B. BOLTON Entirety of head, body and appendages covered with a dense mat of short, branched hairs, apparently mixed trifid-quadrifid. Simple long hairs absent. Alitrunk, pedicel and gaster light to very dark brown, the head bright yellow, contrasting strongly to the remainder of the body. This very distinctive species is apparently confined to New Guinea but two related species, pulchella and mayri, are known from the Solomon Islands. A third species, and the one most closely resembling fulviceps, is pulcherrima from Uganda, but the apparent similarities in colour, sculpture and body form may be due to convergence as the petiole node is very differently constructed in the Ugandan species, the node being high and narrow in profile with a deep anterior face (compare Text-figs 62 and 59). MATERIAL EXAMINED. NEw GuINEA: Huon Peninsula, Lower Busu R. (E£. O. Wilson); Huon Peninsula, Ngandus (£. O. Wilson); Nadzab (E. O. Wilson); Lae, Didiman Ck. (E. O. Wilson); Bunbok Valley, Boana (E. O. Wilson); Lae (E. Hardy); Cyclops Mts, Mt Lina (L. E. Cheesman); Wau N. on Bulolo Rd (S. Peck); W. Highlands, Baiyer R. (S. Peck): Triglyphothrix indosinensis (Wheeler) comb. n. Tetvamorium indosinense Wheeler, 1927 : 97, fig. 6. Syntype workers, VIETNAM: Klinai Tao, 2.vill.1925 (F. Silvestvt) (MCZ, Cambridge) [examined]. Note. In the original description the type-locality is given as Hanoi, but the specimens determined and labelled as types by Wheeler give the locality as Klinai Tao. As these specimens fit the original description perfectly and have been labelled as types by Wheeler himself it is assumed that the locality of Hanoi is an error. Worker. TL 3:4-3:6, HL 0:88-0:92, HW 0:78-0:80, CI 87-89, SL 0:80-0:82, SI 100-103, PW 0:50-0:52, AL 0-94—0-96 (3 measured). Antennal scrobes absent, frontal carinae reduced, ending at the level of the anterior margin of the eye. Eyes small, maximum diameter ca 0-14, very slightly greater than the maximum width of the scape. Antennal scapes exceptionally long, the longest recorded in the genus to date. Promesonotal dorsum evenly convex, metanotal groove faintly indicated in profile. Propodeal spines elongate and acute. Legs very long, maximum length of hind femur ca 0:88. Petiole in profile with a long anterior peduncle and a high, roughly rectangular node. In dorsal view the node is compressed from side to side, slightly longer than broad and broader behind than in front. Mandibles longitudinally striate-rugulose. Dorsum of head pre- dominantly finely longitudinally rugulose, the rugulae diverging posteriorly and with a few faint cross-meshes which are distinctly weaker than the longitudinal component. Dorsal alitrunk with pronotum reticulate-rugulose, the sculpture becoming much weaker posteriorly. Petiole with vestiges of rugulation but the postpetiole almost completely smooth. Gaster unsculptured. All surfaces of head and body with numerous fine short hairs which are reclinate or closely adpressed to the surface and which are predominantly trifid, except on the dorsum of the first gastral tergite where the majority of hairs are simple. Elongate, erect, simple hairs completely absent. Colour dull yellow-brown. The combination of extremely long antennal scapes, lack of scrobes and very long legs makes this species unmistakable. At present it cannot be assigned to ANT TRIBE TETRAMORIINI 349 any definite species-group as it shows very few similarities with any other known species. Triglyphothrix katypa sp. n. Holotype worker. TL 2-7, HL 0°64, HW 0:58, CI 90, SL 0-44, SI 77, PW 0°44, AL 0°74. Antennal scrobes with a sharp dorsal margin, less strongly defined posteriorly and ventrally, primarily demarcated ventrally by a change in sculpture from a rugoreticulum outside the scrobe to reticulate-punctate inside. Median carina of scrobe distinct to level of posterior margin of eye. Dorsal alitrunk evenly convex, the propodeum with a pair of short, acute spines which are shorter than the long, broadly triangular. metapleural lobes. Petiole in profile with the tergal portion of the node distinctly longer than high, in dorsal view the node slightly broader than long, broadest at about its midlength. All dorsal surfaces of head, alitrunk, petiole and postpetiole finely, closely reticulate-rugulose. Mandibles longitudinally striate; gaster smooth and shining. All dorsal surfaces of head and body with numerous trifid hairs, those on the first gastral tergite universally so. Dorsum of head with numerous elongate, simple hairs and especially with a spaced row of long, simple, quite stout hairs along the dorsal edge of the frontal carinae where they form the upper borders of the scrobes. A few simple long hairs are also present on the dorsal alitrunk but are absent from the first gastral tergite, antennae and legs. Colour light red-brown, the appendages and gaster yellow-brown. Paratypes. As holotype, with dimensions TL 2-6—2:7, HL 0:60-0:64, HW 0:54-0:58, CI 90-93, SL 0:42-0:44, SI 73-77, PW 0:42-0:46, AL 0:70-0:74 (4 measured). Holotype worker, PHILIPPINES: Dumaguete, Lake, 30.iv.1931 (J. W. Chapman) (MCZ, Cambridge). Paratypes. 4 workers with same data as holotype (BMNH; MCZ, Cambridge) This small species is most closely related to pnyxis of West Malaysia but is separable by the lack of elongate simple hairs from the upper scrobe margins in pnyxis, distinct in katypa. Several more series of this species are present in the Chapman collection at MCZ, Cambridge, but they are all dirty and in rather poor condition; they will require cleaning and remounting before they can be of any value to systematics. Triglyphothrix kheperra sp. n. (Text-fig. 71) Holotype worker. TL 2:6, HL 0:66, HW 0:64, CI 97, SL 0:44, SI 69, PW 0:46, AL 0-68. Antennal scrobes strongly developed, with an acute dorsal margin and a distinct median longitudinal carina; the lower margin of the scrobe well defined behind the eye. Propodeal spines acute, longer than the metapleural lobes (which in many specimens tend to be low, broad and rounded). The petiole node in dorsal view is slightly broader than long (ca 0-04—0-06 difference) but in profile it is high and relatively narrow, with front and rear faces roughly parallel. Dorsal surfaces of head, alitrunk and pedicel finely and closely reticulate-rugose, the gaster unsculptured, smooth and shining. All dorsal surfaces of head and body coated with abundant trifid hairs, the cephalic dorsum also with a few longer, simple hairs on the upper margins of the antennal scrobes. (In some specimens, rarely, one or two long, simple hairs are present elsewhere on the dorsum of the head.) Hairs on outer surfaces of middle and hind tibiae short, of equal length, distinctly less than the maximum tibial width. Colour uniform brown, varying in shade from light to mid-brown in the material available. Pavatypes. As holotype, with a size-range TL 2-5-2°8, HL 0-56-0-70, HW 0-50-0:66, CI 92-97, SL 0:34-0°46, SI 67-72, PW 0-40-0:52, AL 0:62-0°76 (10 measured). 350 B. BOLTON Holotype worker, JAVA: Bogor, 21.x.1973 (B. Bolton) (BMNH). Paratypes. 8 workers with same data as holotype; 5 workers as holotype but 24.X.73; I worker 19.x.1973, and 6 workers 29.x.73. (BMNH; MCZ, Cambridge; MHN, Geneva.) This species, although previously undescribed, is widespread in the Oriental and Indo-Australian regions and is capable of acting as a tramp species in human commerce dealing with fresh plants and their products. Specimens have been intercepted at least twice in Britain, once from York (1928) in a consignment of illipic nuts from Borneo (recorded by Donisthorpe, 1933 as lanuginosa), and more recently in Kew Gardens, London (1974) amongst the roots of a plant imported from Assam. No doubt the species has been introduced on other occasions but has gone unnoticed or has been assumed to be lanuginosa (= striatidens), the most common tramp-species in this genus both in Britain and in other temperate-zone countries. The closest relatives of kheperra include adpressa and meshena, which are easily separated as they have the first gastral tergite sculptured basally, and parvispina. This last-named species is close to kheperra but has much shorter propodeal spines. MATERIAL EXAMINED. Inp1A: Assam (intercepted at Kew Gdns, England). ViETNAmM: Van Phu (F. Silvestri). Honc Kone (R. H. Lefevre). PuHrittppines: Mt Magading (C. F. Baker). Borneo: Sarawak (R. Thaxter); Pontianak (H. Britten) (intercepted at York, England). SumBa: Kananggar (Dammerman). PRINSEN I. (Dammerman). SULAWESI: Balampesoang For. (W. L. Brown). Triglyphothrix lanuginosa (Mayr) (Text-fig. 63) Tetvamorium lanuginosum Mayr, 1870 : 972, 976. Holotype worker, Java: Batavia (= Dja- karta) (NM, Vienna) [examined]. Tetvamorium obesum race striatidens Emery, 1889) : 501. Syntype workers, BuRMA: Bhamé, vii. 1886 (L. Fea) (MHN, Geneva) [examined]. Syn. n. Triglhyphothrix lanuginosa (Mayr) Emery, 1891 : 4 (implied in footnote 2). Triglyphothrix obesus race stviatidens (Emery); Emery, 1891 : 4. Triglyphothrix striatidens var. laevidens Forel, 1900 : 284. Syntype workers, Mexico (MHN, Geneva) [examined]. Syn. n. Triglyphothrix striatidens (Emery); Forel, 1902b : 704. [Raised to species.] Triglyphothrix striatidens st. ovissana Forel, 1902a : 239. Syntype workers, Inp1A: Orissa XLIV/17b (Taylor) (MHN, Geneva) [examined]. Syn. n. Triglyphothrix striatidens var. australis Forel, 1902c : 449. Syntype workers, females, AUSTRALIA: Queersland, Mackay (Turner) (MHN, Geneva; MCZ, Cambridge) [examined]. Syn. n. Triglyphothrix striatidens var. felix Forel, 1912b : 160. Syntype workers, SEYCHELLE ISLANDS: Félicité, Silhouette, Mare aux Cochons, 1908 (H. Scott) (MHN, Geneva; BMNH) [examined]. Syn. n. Triglyphothrix ceramensis Stitz, 1912 : 506. Holotype worker, SERAmM (Tauern) (location of holotype not known). Syn. n. Triglyphothrix striatidens var. flavescens Wheeler, 1929 : 55. Syntype workers, SINGAPORE: Johore, 11.ii.25 (F. Silvestvi) (MCZ, Cambridge) [examined]. Syn. n. ANT TRIBE TETRAMORIINI 351 Triglyphothrix mauricei Donisthorpe, 1946a : 778. Holotype and paratype worker, Mauritius: Rose Hill, 1942 (R. Mamet) (BMNH) [examined]. Syn. n. Triglyphothrix tricoloy Donisthorpe, 1948 : 136. Paratype workers, females, NEw GUINEA: Maffin Bay, 17 and 20.vi.1944 (E. S. Ross) (BMNH; CAS, San Francisco) [examined]. Syn. n. Worker. TL 2-2-3:0, HL 0-+56-0-72, HW 0:52-0:70, CI 92-97, SL 0:36-0:50, SI 68-75, PW 0-40-0:50, AL 0:58-0:80 (25 measured). Antennal scrobes well developed with a sharply marked dorsal margin and usually with a defined median longitudinal carina, although in some specimens this may be rather reduced. In all material examined the scrobe ends well behind the posterior margin of the eye. Pro- podeal spines long and acute, longer than the metapleural lobes, and are usually feebly upceurved along their length. Metapleural lobes usually somewhat upcurved and acute. Node of petiole in profile with the dorsal surface curving into the posterior face so that the two are not separated by an angle but by a curvature which is much less acute than the angle separating anterior and dorsal surfaces. In extreme cases the dorsal and posterior faces are united in a single convexity (Text-fig. 63). Petiole more massively built than postpetiole. Mandibles striate except in some small individuals where it may be partially effaced or absent. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely reticulate-rugose. On the head there is a tendency for the transverse rugulae to break down on the anterior portion, leaving this area longitudinally rugose, and on the postpetiole the sculpture may be partially or totally effaced, usually in smaller specimens. First gastral tergite smooth and shining. All dorsal surfaces of head and body with abundant hairs, predominantly a mixture of simple and bifid but sometimes also with a few trifid hairs. On the first gastral tergite trifid hairs are nearly always absent. Numerous elongate, simple hairs present. Leading edges of antennal scapes and outer margins of middle and hind tibiae with numerous simple hairs of varying length, the longest of which are at least as long as the maximum width of the appendage from which they arise. Colour brown, varying in shade from light to dark, often with alitrunk and gaster of different shades. An accomplished tramp-species on a worldwide scale, Januginosa (better known by its junior synonym striatidens), is known or has been reported from all major and a great number of minor land masses on the earth’s surface. Wilson & Taylor (1967) state that the species (recorded as striatidens) ‘is probably native to tropical Asia’, and I concur with this as the most closely related species to lanuginosa (coonoorensis and obesa) are known only from India and Burma. In the tropics the species survives outdoors but known collections of lanuginosa from the temperate zones have all been made in botanical or zoological gardens where the ants have most probably been imported with tropical plants or animal foodstuffs. MATERIAL EXAMINED. Inp1A: Barrakpore (Rothney); Calicut (A. B. Soans); Maharashtra, Edalabad (E. S. Ross & D. Q. Cavagnaro); Assam, Jorhat (A. C. Cole); Assam, Garo Hills, Siju Cave; Kerala State, Kannoth (A. B. Soans & W. L. Brown); Calcutta (ex coll. F. Smith); no. loc. (Wroughton); Poona (Wroughton). NrpaLt: Amlekhganj (E. I. Coher). Sri Lanka: Pollonaruwa (K. L. A. Perera). Burma: Bhamo (Bingham). PHILIPPINES: Mindoro, San Jose (E. S. Ross); Luzon, Mt Makiling (K. Dumont & R. Morse); Dumaguette (J. W. Chapman); Moilo (S. Amaguin); Negros I., Bago (F. del Rosario); Davao (A. Reyes); Dumaguete (D. Empeso); Dumaguete (F. del Rosario); Horns of Negros (J. W. Chapman). Sotomon Is.: Guadalcanal (P. Greenslade); New Georgia (P. Greenslade). Wrst Mataysia: Alor Star (G. H. 352 B. BOLTON Lowe); Sg. Patani (G. H. Lowe); Kuala Lumpur (K. W. Ng). Bornero: Kuching (J. Hewitt), Java: Batavia (= Jakarta); Semarang. FLORES: Nangagete (W. L. Brown). SumBa: Laora (Dammerman); Kambera (Dammerman). AUSTRALIA: Queensland, Townsville (Dodd); Cape York, Silver Plains (Darlington); N. Terr., Howard Springs (Brown); N. Terr., Mt Bundy Rd (Brown); Lankelly Ck. (Darling- ton); Mackay (R. E. Turner); N. Terr. (G. F. Hill). Marianas I.: Guam, Pt Oca (G. E. Bohart & J. L. Gressitt); Pati Point (H. S. Dybas); Amantes Point (H. S. Dybas). CAROLINE Is.: Truk I. (Yasu et Yoshi); Yap I., S. Ramung I. (R. J. Goss). WaLtis Is.: Nuku Tapu I; Nuku Hifala I; Uvea; Nukuione I. (G. Hunt). Samoa: Upolu, Lauli (T. E. Woodward). CuristmMan I. (no data). ALDABRA: South I. (B. Cogan & A. Hutson). LEBANON: Antilias Stream (Christiansen). ENGLAND: London, Kew Gdns (Donisthorpe); Birmingham, Bot. Gdns (Bibikoff); Birmingham, Dudley Zoo (A. L. Mills). Cirpperton I. (C. F. Harbison). GERMANY (WEST): Hamburg, imported from Mexico in orchids (ex coll. Forel). JAPAN: Ogasahara, Chichi-jima (M. Tanaka). Other localities include: Seychelle Is., Seram, Singapore, New Guinea, Mauritius, Mexico—see type-data; U.S.A.: south-eastern United States N. to Carolinas (Creighton, 1950); Tunisia (Emery, 1891); Barbados, Mexico (Kempf, 1972). Triglyphothrix mayri Mann Triglyphothrix mayvi Mann, 1919 : 351. Syntype workers, Socomon IsLanps: San Cristoval, Star Harbor (W. M. Mann) (MCZ, Cambridge) [examined]. Worker. TL 2:5-2:8, HL 0:58-0:62, HW 0:52-0:54, CI 87-90, SL 0-42-0-44, SI 77-81, PW 0-42-0'44, AL 0:68-0:70 (5 measured). Antennal scrobe weakly developed, its upper margin poorly defined, not bounded by a narrow flange-like margin but limited by the sculpturation of the cephalic dorsum. Median longitudinal carina of scrobe poorly developed or indistinguishable from other scrobal sculpture. Sides of head more or less straight in full-face view, only extremely feebly convex; eyes prominent. Propodeal spines about as long as the metapleural lobes. Petiole in dorsal view slightly broader than long, in profile the anterior face confluent with the peduncle. Mandibles mostly smooth, with some very faint, almost effaced striation and a few scattered punctures. Dorsal surfaces of head, alitrunk and pedicel finely and densely reticulate-rugose. Basal half or more of first gastral tergite coarsely punctate. All surfaces of head, body and appendages densely coated with short, branched (trifid) hairs of approximately the same length. Long, simple hairs absent from legs, scapes and body behind the mouthparts. Colour uniform pale yellow. Apparently known only from the type-collection, this species is one of four endemic species known to the present only from the Solomon Islands. Two of these species, antennata and vombis, belong to the obesa-group and have numerous simple long hairs projecting from the cephalic dorsum. T. mayri, along with the remaining Solomon Islands endemic, pulchella, do not have such hairs. These last two may be easily separated by the gastral sculpture, which consists of coarse punctures in mayri and longitudinal striation in pulchella. ANT TRIBE TETRAMORIINI 353 Triglyphothrix meshena sp. n. (Text-figs 51, 67) Holotype worker. TL 3:4, HL 0-74, HW 0-70, CI 94, SL 0-46, SI 66, PW 0:54, AL 0°82. Antennal scrobes strongly developed, with a distinct, sharp dorsal margin and a marked median longitudinal carina. Outline shape of alitrunk as in Text-fig. 67. Propodeal spines short and stout, acute, approximately as long as their basal width and scarcely longer than the metapleural lobes. Petiole in profile massive, with a long anterior face and a short, broad peduncle. Postpetiole also massively built, in profile blocky, subrectangular in shape and higher than long. In dorsal view the petiole node as wide as long. Dorsal surfaces of head, alitrunk, petiole and post-petiole finely and densely reticulate-rugose. Basal portion of first gastral tergite coarsely punctate. Mandibles striate. Dorsal surfaces of head, body and appendages covered with a dense mat of erect or suberect trifid hairs, elongate simple hairs absent. Paratype workers. TL 3:1-3:4, HL 0-70-0-74, HW 0:68-0:70, CI 94-97, SL 0°44-0°46, SI 63-68, PW 0:54-0'56, AL 0-80—0-82 (7 measured). Otherwise as holotype. Holotype worker, West MataysiA: Selangor, Gombak, 7.x.1973 (B. Bolton) (BMNH). Paratypes. 5 workers with same data as holotype; 2 workers, WEST MALAYSIA: Selangor, Ulu Gombak For. Reserve, 24.viii.1967, tree lookout area, ca 450m, hill forest (R. Crozier) (BMNH; MCZ, Cambridge). Most closely related to adpressa; notes on the differences of the two species are given under adpressa. A number of specimens in the MCZ, Cambridge collection are also referable to this species. These include three short series from JAvA collected by Dammerman at Depok, 23.xii.1923 (2 workers and a female); Buitenzorg (= Bogor), 18.xii.1922 (2 workers); Buitenzorg, Tjampea, 19.x.1922 (1 worker). The series from Bogor compare perfectly with the type-series, but the specimens from Depok have less massively built pedicel segments and shorter, more obviously reclinate branched hairs on the head and also to some extent upon the gaster. Whether these specimens represent part of the natural variation of meshena or a link in the direction of adpressa or indeed a separate species, cannot be decided on the few specimens presently available, but I note their existence here to bring them to the attention of any future worker on this group. Triglyphothrix nacta sp. n. Holotype worker. TL 3:0, HL 0-78, HW 0-76, CI 97, SL 0:50, SI 66, PW 0:56, AL 0°82. Antennal scrobes strongly defined dorsally and posteriorly, much more weakly delimited ventrally. Median longitudinal carina of scrobe extending back to level of posterior margin of eye and quite distinct. Eyes relatively small, their maximum diameter ca 0-14, only very slightly greater than the maximum width of the antennal scape. Dorsal alitrunk evenly convex in profile, the propodeal spines short, narrow and acute. Metapleural lobes large, lobate, roughly semicircular and longer than the propodeal spines. Petiole node relatively high in profile, the tergal portion of the node higher than long. Postpetiole rounded above in profile, the sternal portion forming a distinctly projecting, roughly rectangular mass ventrally. In dorsal view the petiole is subglobular, slightly broader than long and broadest at about its midlength. Mandibles coarsely striate. Dorsum of head finely and irregularly longitu- dinally rugose, without reticulations except near the occiput. Dorsal alitrunk, petiole and 354 B. BOLTON postpetiole densely reticulate-rugulose, the gaster smooth and shining. Entire head and body densely clothed with trifid hairs, those of the first gastral tergite universally trifid. Simple hairs absent except for one or two relatively short ones which occur on the upper margin of the frontal carina where it forms the dorsal border of the scrobe, but these hairs are not exceptional and are difficult to distinguish from the surrounding branched pilosity. Colour uniform medium brown. Paratype workers. As holotype, with range of dimensions TL 2-9-3-:1, HL 0-76-0-80, HW 0:72-0:78, CI 96-100, SL 0-48-0-52, SI 63-68, PW 0-52-0°56, AL 0-76-0-84 (15 measured). Holotype worker, THAILAND: Nong Hoi, 19.vii.75, in soil (D. Jackson) (BMNH). Paratypes. 19 workers with same data as holotype (BMNH; MCZ, Cambridge). This species is certainly most closely related to parvispina but is a much larger, more robust form in which the petiole is relatively higher and narrower. Triglyphothrix obesa (André) Tetramorium obesum André, 1887 : 294. Syntype workers, INDIA: Gingi (MCZ, Cambridge; MNHN, Paris) [examined]. Triglhyphothrix obesa (André); Emery, 1891 : 4. Worker. TL 3:1-3°5, HL 0:78-0-84, HW 0:76-0:82, CI 95-98, SL 0°54-0'58, SI 68-76, PW 0:54-0:58, AL 0:86-0:90 (7 measured). Antennal scrobes developed, with a distinct dorsal margin but with the median longitudinal carina weakly developed or in some indistinguishable from the surrounding sculpture. Upper portion of scrobe usually less strongly sculptured than the lower. Propodeal spines long, narrow and acute, generally two or more times longer than the metapleural lobes, which are acute and upcurved. Petiole in profile with the dorsal and posterior faces united in a single convex surface, the petiole more massively developed than the postpetiole. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely reticulate-rugose, the reticulation tending to be more strongly developed on the posterior portion of the cephalic dorsum than the anterior, where often only longitudinal rugulae are present. First gastral tergite unsculptured. Mandibles smooth, with scattered pits but without striation. All dorsal surfaces of head and body with numerous branched (bifid and trifid) hairs, also with numerous elongate, simple hairs. Leading edge of antennal scapes and outer surfaces of middle and hind tibiae with abundant long hairs, the longest of which are at least as long as the maximum width of the appendage upon which they arise. Colour orange-brown to reddish brown. Closest related to Januginosa, obesa may be separated by the characters given in the key. In general the presence of mandibular striae in lanuginosa is sufficient to separate the two, and this character has been used on a number of occasions (Forel, 1902); Bingham, 1903; Mann, Ig1g) to isolate obesa. Unfortunately a number of small individuals of Januginosa are now known in which the mandibular striation is reduced or even absent, but in these the HW range falls considerably below the minimum for obesa and the array of branched hairs on the first gastral tergite lacks trifid members, some of which are invariably present in obesa, at least basally. MATERIAL EXAMINED. Inp1A: Travancore (Ferguson); Kerala State, Perumanna Village (A. B. Soans); Kerala State, Nilambur (A. B. Soans). Burma: Ruby Mines District (Bingham); Pegu Yoma (Bingham). ANT TRIBE TETRAMORIINI 355 Triglyphothrix pacifica Mann Triglyphothrix pacifica Mann, 1921 : 460. Syntype workers, female, F1j1 IsLanps: Viti Levu, Nadarivatu (W. M. Mann) (MCZ, Cambridge) [examined]. Worker. TL 2:5-3-0, HL 0-68-0-70, HW 0:60-0:64, CI 88-91, SL 0-48-0-50, SI 77-80, PW 0:50-0:52, AL 0-76—-0-82 (6 measured). Antennal scrobes weakly developed, with a poorly defined dorsal margin and a very weak, meandering median longitudinal carina which is not more strongly developed than adjacent tugulae. Propodeal spines long and acute, longer than the metapleural lobes. Node of petiole blocky and massive, larger than the postpetiole in profile. Mandibles striate. Dorsal surfaces of head, alitrunk and petiole finely reticulate-rugose, dorsal surface of postpetiole unsculptured, smooth and shining or at most with vestigial traces of sculpture. First gastral tergite unsculptured. Leading edge of antennal scapes and outer margins of middle and hind tibiae with short hairs only which are less than half the maximum width of the appendage from which they arise. Dorsal surfaces of head, alitrunk and pedicel with a mixture of simple and ‘branched hairs and with some elongate, simple hairs on head. Hairs on first gastral tergite either almost universally simple, or with bifid hairs present. Colour light reddish brown to dark brown. Apart from a few small individuals of stviatidens this is the only known species of Triglyphothrix outside the Ethiopian region which has the postpetiole unsculp- tured. This single character serves to separate pacifica from its closest relatives also. It is the only endemic species of this genus known from the Fiji Islands, and the species appearing to be closest related are antennata and vombis, both endemic in the Solomon Islands. MATERIAL EXAMINED. Fiji IsLanps: Nausori Highlands, SW. Fiji (W. L. G D. E. Brown); Viti Levu, Nadala (E. O. Wilson). Triglyphothrix parvispina Emery (Text-fig. 64) Triglyphothrix parvispina Emery, 1893: 214 [diagnosis in key]. Syntype (?) workers, BornEo (?): Poulo Laut (Bedot & Pictet) (MCSN, Genoa). Triglyphothrix parvispina var. formosae Forel, 1912a:52. Syntype workers, Taiwan: ‘Formosa’, Pilam (H. Sauter) (MHN, Geneva) [examined]. Syn. n. Worker. TL 2-4-2:6, HL 0:60-0:66, HW 0:56-0:62, CI 93-97, SL 0-40-0-44, SI 70-75, PW 0:40-0:48, AL 0:64-0:70 (6 measured). Antennal scrobes strongly developed, with a sharply defined, narrow, flange-like dorsal margin and a well-differentiated median longitudinal carina. Propodeal spines short, reduced to a pair of triangular, acute teeth which are at most only as long as the metapleural lobes, often shorter. Metapleural lobes low and rounded, not developed into upcurved teeth. Dorsal surfaces of head, alitrunk and pedicel finely reticulate-rugose, first gastral tergite unsculptured. All dorsal surfaces of body densely clothed with short, predominantly branched (trifid) hairs. Long, erect hairs absent. On the whole this species shows a close resemblance to kheperra and their separation at present rests rather uneasily upon the relative lengths of the propodeal spines. Further collections may show that the two species intergrade. With some reservation I refer to parvispina two specimens in MCZ, Cambridge 356 B. BOLTON collection with data as follows: [CHINA:] Foochow, Peta (F. Silvestri). These may represent a variant population of parvispina or may be a separate species. The main character setting them somewhat apart from my concept of parvispina is the array of hairs on the gaster which in these specimens is primarily a mixture of simple and bifid hairs, whilst in parvispina the gastral hairs are predominantly trifid. Despite this I feel that it would be unwise to separate these specimens from parvispina at the present time as collections are too few to allow a sound judgement to be made. MATERIAL EXAMINED. PHILIPPINES: Dumaguete (J. W. Chapman); Dumaguete (D. Empeso). Triglyphothrix pnyxis sp. n. (Text-fig. 70) Holotype worker. TL 2-9, HL 0-68, HW 0-62, CI 91, SL 0-48, SI 77, PW 0:46, AL 0-76. Upper border of antennal scrobe demarcated by the irregular line at which the cephalic sculpture terminates, without a flange or narrow, acute margin formed by the frontal carinae. Posteriorly and ventrally the scrobe margins are even less well defined than this, being indicated merely by a change in sculpturation. Median carina of scrobe present but very weak. Propodeal spines narrow and acute, longer than the triangular metapleural lobes. Node of petiole in profile long and low, distinctly longer than high. In dorsal view the node longer than broad, broadest posteriorly and quite narrowly rounded in front. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely and closely reticulate-rugulose. Mandibles striate; gaster smooth and shining. All dorsal surfaces of head and body densely covered with trifid hairs; the hairs on the first gastral tergite universally trifid. A few very fine simple hairs are present on the dorsum of the head but the upper margins of the scrobes are without a regular spaced row of elongate simple hairs. Colour dark red-brown, the appendages yellow-brown. Paratype worker. As holotype, differing only in that HW is 0-50 and SI 80. Holotype worker, West Matraysia: Trengganu, Gn. Lawit, summit ridge, 4200 ft, 11.11.1974 (T. Clay) (BMNH). Paratype. A single worker with same data as holotype (MCZ, Cambridge). Amongst the five species making up the kheperra-complex in the walshi-group, two species, pnyxis and katypa, have a petiole node which is elongate and low in profile. The two are quickly separable as katypa has a sharply defined dorsal margin to the antennal scrobe from which projects a row of elongate, simple hairs which are stout and very conspicuous. In puyxis on the other hand the dorsal margin of the scrobe is feebly marked and elongate simple hairs are absent from it. Triglyphothrix pulchella Mann Triglyphothrix pulchella Mann, 1919 : 352, fig. 31. Syntype workers, female, SoLOMON IsLanps: Malaita, Auki (W. M. Mann) (MCZ, Cambridge) [examined]. Worker. TL 2:-4-2:6, HL 0:56-0:62, HW 0:52-0'54, CI 89-93, SL 0-38—-0-42, SI 72-77, PW 0:40-0:42, AL 0:62-0:66 (3 measured). Antennal scrobes shallow, with a poorly defined upper margin and a weak, feebly differ- entiated median longitudinal carina. Propodeal spines long, narrow and acute, much longer than the metapleural lobes. Dorsal surfaces of head, alitrunk and pedicel finely reticulate- rugose. Mandibles striate. Basal one-third or more of first gastral tergite with strong ANT TRIBE TETRAMORIINI 357 longitudinal striation. Dorsal surfaces of head and body densely coated with short branched (trifid) hairs, long, simple hairs absent. Leading edges of antennal scapes and outer margins of tibiae with a dense coat of short hairs of approximately equal length, without elongate, simple hairs. Colour yellow, the head and gaster generally a lighter shade than the alitrunk and pedicel. This distinctive small species is endemic in the Solomon Islands and appears to be closely related to fulviceps of New Guinea. The diagnostic notes above should serve to separate this species from other endemic species found in the Solomon Islands. MATERIAL EXAMINED. Sotomon IsLANDS: Malaita I., Dala (P. Greenslade); Nggela I. (P. Greenslade); Isabel I., Buala (P. Greenslade). Triglyphothrix rossi sp. n. Holotype worker. TL 2-8, HL 0:62, HW 0:56, CI go, SL 0-48, SI 86, PW 0:42, AL 0:72. Antennal scrobes very poorly developed, virtually erased, the extensions of the frontal carinae which form the upper scrobe margins are extremely weak and cannot be distinguished from the other sculpture behind the level of the eyes. Scrobal area not differentiated from rest of the head, without a median carina, the scrobal area almost or quite as strongly sculptured as the cephalic dorsum. Dorsal alitrunk evenly convex in profile, terminating in a pair of Narrow propodeal spines. Metapleural lobes dentiform and directed upwards. Petiole in profile shaped as in lanuginosa (Text-fig. 63) but with the antero-dorsal angle more rounded. In dorsal view the petiole broadly triangular with a broadly rounded, blunt apex anteriorly, broadest posteriorly where its maximum width is equal to the width of the postpetiole. Dorsal surfaces of head, alitrunk and the pedicel segments with a fine rugoreticulum. Man- dibles striate. Gaster smooth and shining, unsculptured. All dorsal surfaces of head and body with a mixture of simple and bifid hairs, very distinctive on the first gastral tergite. Dorsum of head with a row of long, erect, simple hairs along the line of the frontal carinae and with scattered long, simple hairs elsewhere on the head and dorsal alitrunk. Outer surfaces of middle and hind tibiae with numerous suberect to subdecumbent hairs which are all of approximately the same length and are much shorter than the maximum tibial width. Leading edges of antennal scapes without long simple hairs which are greater in length than the maximum scape width. Colour uniform dark brown, the appendages lighter. Paratype worker. As holotype, with size range TL 2:6—2-9, HL 0:60-0:66, HW 0:54-0:58, CI 85-90, SL 0:46-0:52, SI 85-89, PW 0:40-0:46, AL 0°68-0-76 (10 measured). Holotype worker, Inp1A: South, 8 miles NE. Munnar, 6200 ft, 20.iii.1962 (E. S. Ross & D. Q. Cavagnaro) (CAS, San Francisco). Paratypes. 32 workers and 1 female mounted, a number of other workers plus brood in alcohol; with same data as holotype (CAS, San Francisco; BMNH; MCZ, Cambridge). Within the obesa-group rossi is most closely related to coonoorensis. Both species have reduced antennal scrobes and both are endemic in the Indian subcontinent. T. rossi is separated from coonoorensis, and also from the remainder of the obesa- group, by its lack of long simple hairs on the leading edge of the antennal scape. In all its allies these hairs are very long, generally longer than the maximum width of the scape. 358 B. BOLTON Triglyphothrix vombis sp. n. (Text-fig. 73) [Triglyphothrix obesa (André) sensu Mann, 1919 : 349. Misidentification.] Holotype worker. TL 3:1, HL 0-72, HW 0:66, CI 92, SL 0:48, SI 73, PW 0-50, AL 0°80. Antennal scrobes strongly developed, with a sharply defined dorsal margin and a weak but distinct median longitudinal carina. Propodeal spines narrow and acute, slightly longer than the acute metapleural lobes. Petiole in profile with the dorsal and posterior faces united in a single convex surface; the postpetiole about as high as the petiole. Mandibles mostly smooth but with a few scattered pits and vestiges of longitudinal striation in places. Dorsal surfaces of head, alitrunk and pedicel finely and densely reticulate-rugose. First gastral tergite smooth and shining. Head, alitrunk and pedicel densely clothed with fine hairs, the majority of which are simple, relatively few bifid hairs being present. Hairs on first gastral tergite universally simple. Dorsum of head and alitrunk also with a number of long, simple hairs. Outer margins of tibiae of middle and hind legs with a dense coat of short hairs, all of approximately equal length and less than half the maximum width of the tibia upon which they arise. Colour light orange-brown, the gaster yellowish brown. Paratype workers. TL 2-9-3:1, HL 0-70-0-:72, HW 0:64-0:66, CI 91-94, SL 0-46-0'50, SI 70-75, PW 0:-46-0-50, AL 0-76—0-80 (7 measured). As holotype but in some the mandibles smooth, without trace of striation. Holotype worker, Sotomon Is.: Ugi (W. M. Mann) (MCZ, Cambridge). Paratypes. 7 workers, 1 female, same data as holotype (MCZ, Cambridge; BMNH). Originally described by Mann (1919) as obesa, this species is in fact distinct and belongs to a complex containing also antennata, brevidentata, chepocha and pacifica. It is easily distinguished from obesa and its allies by its lack of varying-length simple hairs on the tibiae, having instead a dense coat of short hairs of uniform length. Triglyphothrix walshi Forel (Text-fig. 68) Triglyphothrix walshi Forel, 1890 : cvii. Syntype workers, female, Inp1a: Bengal, Poree (J. Walsh), and Poona (R. C. Wroughton) (MHN, Geneva) [examined]. Triglyphothrix musculus Forel, 1902a : 239. Syntype workers, Inp1a: Coonoor (Wvoughton) (MHN, Geneva) [examined]. Syn. n. Triglyphothrix walshi var. spuria Forel, 1912c : 58. Syntype workers, SINGAPORE and SRI Lanka: Colombo (A. Miiller) (MHN, Geneva) [examined]. Syn. n. Worker. TL 2:3-2°8, HL 0-56-0-68, HW 0-52-0-66, CI 93-98, SL 0-32-0-48, SI 61-67, PW 0:40-0'54, AL 0:54-0:74 (14 measured). Antennal scrobes strongly developed, with a sharply defined dorsal margin and a distinct median longitudinal carina. Alitrunk in dorsal view short and broad, compact, the pronotal angles sharp, giving the species a square-shouldered appearance. Propodeal spines acute, longer than the metapleural lobes. Node of petiole in dorsal view strongly antero-posteriorly compressed, transverse, distinctly broader than long. Dorsal surfaces of head, alitrunk, petiole and postpetiole finely reticulate-rugose, the rugosity sometimes reduced on the post- petiole. First gastral tergite usually with the basal one-third or more finely longitudinally striate, often with punctures between the striae, but in some populations from Sri Lanka this sculpture is reduced or absent. Head, body and appendages uniformly clothed with a dense mat of fine, trifid hairs, the head also with a few elongate simple erect hairs which are ANT TRIBE TETRAMORIINI 359 longer than the neighbouring trifid hairs and which are present at least on the dorsal surface of the upper margins of the scrobes. Colour light to dark brown. In a number of ways this species resembles the Ethiopian region forms related to inezulae more than any other species. The resemblance to thoth is particularly striking and further investigation of their relationship would be interesting when thoth is better known. Suffice to say that the two seem distinct as thoth lacks erect simple hairs on the head, has reduced propodeal spines and has the gaster unsculptured. MATERIAL EXAMINED. Inp1A: Kanara (Wroughton); no loc. (coll. Bingham); Izatnagar (ex C.I.E. coll.); Nilgiris (?); Cochin (Rothney); Kerala State, Silent Valley Reserve (A. B. Soans & W. L. Brown); Kerala State, Peruva (A. B. Soans & W. L. Brown); Irinjalakuda (A. B. Soans); Calicut (A. B. Soans); Madras Presidency, Chilka Lake (?); Wallon? (ex coll. Wasmann). Sri LANKA: Colombo (W. L. Brown); Gilimale (E. O. Wilson); Nuwara Eliya (K. L. A. Perera). CuinA: Foochow (H. H. Chung). PHILIPPINES: Dumaguete (J. W. Chapman); Dumaguete, Silliman University (D. Empeso). TETRAMORIUM Mayr Tetvamorium Mayr, 1855 : 423. Type-species: Formica caespitum L., 1758 : 581; by subse- quent designation of Giraud, 1879 : 1016. Tetrogmus Roger, 1857: 10. Type-species: Tetrogmus caldavius Roger, op. cit.: 12 [= Tetra- morium simillimum (F. Smith), 1851 : 118]; by monotypy. [Synonymy by Mayr, 1863 : 456.] Xiphomyrmex Forel, 1887 : 385 [as subgenus of Tetvamorium]. Type-species: Tetvamorium (Xiphomyrmex) kelleri Forel, loc. cit.; by subsequent designation of Wheeler, 1911 : 175. Syn. n. Xiphomyrmex Forel; Emery, 1896a : 183; 1914a : 42. [Raised to genus.] Atopula Emery, 1912: 104. Type-species: Atopomyrmex nodifer Emery, 1901 : 115; by original designation. Syn. n. Macromischoides Wheeler, 1920 : 53. Type-species: Macromischa aculeata Mayr, 1866 : 507; by original designation. Syn. n. Sulcomyrmex Kratochvil, 1941 : 84 [as subgenus of Tetvamorium]. [Proposed without designa- tion of type-species; name not available. ] Lobomyrmex Kratochvil, 1941 : 84 [as subgenus of Tetvamorium]. Type-species: Tetramorium (Lobomyrmex) ferox silhavyi Kratochvil, loc. cit. [= Tetramorium ferox Ruzsky, 1903 : 309], by monotypy. Syn. n. Diacnosis. Worker. Mandibles with three teeth apically, the third smaller than the second and followed by a row of 3-7 small or minute denticles; the arrangement 3 teeth plus 4 denticles overwhelmingly predominant. Palp formula 4, 3 (47 species dissected), very rarely 4, 2 (I species) or 3, 2 (1 species). (The formula 3,3 has not been detected but probably occurs in some minute species as in Triglyphothrix.) Antennae I1- or 12-segmented, the three apical antennomeres forming a club. Eyes present, usually moderate to large, very rarely reduced or minute. Frontal carinae strongly developed in most species, reaching almost to the occipital margins, but in a few groups (e.g. caespitum-group; sericeiventre-group) and a number of isolated species reduced and petering out before the level of the eyes. Median longitudinal clypeal carina usually distinct, reduced or absent in only a few species. Cephalic median carina generally developed but not uncommonly reduced or absent. Antennal scrobes 360 B. BOLTON showing all stages of development from absent to deep, but in the latter case the scrobe is only very rarely divided into upper and lower compartments by a longitudinal ridge or carina. Alitrunk fusiform, without sutures, but the metanotal groove is impressed in many species. Propodeum usually armed with a pair of spines or teeth, only very rarely unarmed. Meta- pleural lobes present, usually acute or dentiform but sometimes reduced and rounded, some- times minute or vestigial. Petiole pedunculate, without a large ventral process. Lamelliform appendage of sting variable in shape, triangular, dentiform, pennant-shaped or spatulate. Tibial spurs present on middle and hind legs in many large species, usually simple but in a few they may be minutely barbulate. In some smaller and some larger species the spur is absent or indistinguishable from the adjacent pilosity. Pilosity very variable; usually with numerous erect or suberect hairs on all dorsal surfaces but these may be absent from various sclerites, reduced in number and size, appressed, or rarely completely absent. Very rarely the pilosity is bizarre, the hairs being spatulate, short-clavate, palmate, plumose or pinnate, but never with regularly bifid, trifid or quadrifid hairs. Female. As worker but with flight sclerites and ocelli, the virgin queens fully alate. Male. Known for only very few species. Antennae Io to 11-segmented, the second funicular forming an elongate fusion-segment. Mandibles dentate. Palp formula as worker. Pilosity variable as in worker, without the branched hairs characteristic of Tviglyphothrix. As defined above Tetramorium is a large compact genus which, on recent count, included some 465 named forms, of which I suspect that about 200-250 are valid. As was the case with Tviglyphothrix it is apparent that numerous common species of Tetramorium have been described many times and have quite a lot of infraspecific or infrasubspecific names attached to them whilst of the less common forms a great number of valid species remain undescribed. Revision of this great mass of names is at present in progress and I have already completed studies on the species with I1-merous antennae which formerly constituted the genus Xipho- myrmex, Now synonymized (see below). I envisage the revision of Tetramorium as being published in two or three sections, each dealing with a fairly restricted faunal area but it is already obvious that the Ethiopian/Malagasy fauna is by far the largest and seems to present the most problems of a taxonomic nature. The distribution of Yetramorium embraces all of the Old World tropics and subtropics and the Palaearctic and Nearctic Regions. The only region without any endemic species is the Neotropical but even here a few species have been introduced by accident and now appear to be flourishing. The principle nest-sites of this genus include rotten wood, under stones or directly into the earth but a few species are known which are arboreal, nesting either in the wood, under bark, or con- structing carton nests of vegetable fibre and debris. The majority of species are cryptic or retiring forms but some are very distinctive and run actively on the surface of the ground or on trees. GENERIC SYNONYMY OF Tetrvamorium Xiphomyrmex. When Forel (1887) described Xiphomyrmex he made it a sub- genus of Tetramorium, stating that the only difference between them was contained in the number of antennal segments which each possessed, namely eleven for Xiphomyrmex and twelve for Tetramorium. Emery (1896a; 1914a) considered that this difference was sufficient to raise Xiphomyrmex to generic status and this view was endorsed by Wheeler (1922). Arnold (1917; 1926) on the other hand continued ANT TRIBE TETRAMORIINI 361 to treat Xiphomyrmex as a subgenus. In later years, as various new species were described, they were assigned to one of these two groups solely on their antennomere count, without regard for any other characters which they possessed. Whilst studying Tviglyphothrix and Rhoptromyrmex it became obvious that it was possible for antennal segmentation to vary within genera in this tribe without affecting characters of generic significance, which remained stable, and this prompted a search for definitive characters which could be used to separate Tetramorium and Xiphomyrmex. No new characters could be found, but Brown (1973) had noted that the sting appendage in workers and females was ‘perpendicular to the shaft and sharply dentiform or pennant-shaped’ in Tetramorium, whilst in Xiphomyrmex the sting appendage was ‘Rounded-spatulate’. On investigation it was found that the rounded-spatulate appendage, supposedly confined to Xiphomyrmex, also occurred in Tetramorium (as defined by their I2-merous antennae) throughout the gvassii- and squaminode-groups, and that an appendage perpendicular to the long axis of the sting or inclined at an angle to it occurred in the carinatum- and schaufussi-groups of Xiphomyrmex (as defined by their 11-merous antennae). Examination of a large number of species further appeared to show that a straight-spatulate appendage may be derived from a perpendicular-dentiform structure. The main forms of sting appendage and the antennomere counts are tabulated below, with some examples. I. Sting appendage perpendicular or slightly inclined from long axis of shaft, acute apically. Sting appendage perpendicular or slightly inclined from long axis of shaft, blunt apically. Sting appendage continuing long axis of shaft (may be feebly up- curved), blunt apically. Sting appendage continuing long axis of shaft; blunt and upcurved apically. Antennae with 12 segments: camerunense- group, caespitum-group, sericeiventre- group, guineense-group. Antennae with 11 segments: striolatum- group. Antennae with 12 segments: part of grassii-group and of sgwaminode-group. Antennae with I1 segments: carinatum- group. Antennae with 12 segments: part of grassii-group and of sguaminode-group. Antennae with II segments: muralti- group, angulinode-group. Antennae with 11 segments: weitzecker- group, angulinode-group. The species in the first category may be regarded as ‘super-Tetramorium’ and those in the fourth as ‘super-Xiphomyrmex’ when sting structure alone is considered but it is obvious that all possible intergrades exist, regardless of their antennomere count. As there are absolutely no other characters which can be considered it is apparent that Tetramorium is capable of showing the same variation as Triglypho- thrix and Rhoptromyrmex as regards antennal segmentation, and Xiphomyrmex falls into the synonymy of Tetramorium. 362 B. BOLTON Once this has been accepted a number of interesting correspondences become apparent. For instance, the wettzeckeri-group (formerly of Xiphomyrmex) is inseparable from the sguaminode-group (formerly of Tetramorium) in all characters except that of antennal segmentation, and the closest relatives of T. grassii seem to be the former Xiphomyrmex species of the schaufussi-group. In view of the above it seems most probable that the antennomere count of eleven has arisen independently and at different times from a number of species- groups with 12-segmented antennae within Tetramorium. One interesting point arising from the II- versus 12-merous antennae is the fact that Tetramorium in which the antennae are 11-segmented are predominant in Australia and Madagascar, where they outnumber the 12-segmented forms in terms of numbers of endemic species, whereas elsewhere they form a relatively minor part of the fauna. It can be postulated from this relict distribution that the parts of the genus with I1-segmented antennae and relatively unspecialized body-form (this excludes the very specialized weitzeckeri-group) represent the remnants of an earlier radiation of the genus which has since been in large part replaced by their later-radiating 12-segmented congeners. This presupposes that the early loss of an antennomere in some groups somehow gave its owners an early advantage over their relatives which retained the ancestral 12-merous antennae but that a later change of circumstance obviated this advantage and the 12-merous species then overhauled and gradually replaced their 11-merous allies except in peripheral or inaccessible parts of the range. This is of course speculation but it does fit the present facts of distribution as regards II- versus 12-segmented antennae and may even explain why a single endemic 11-segmented species is still found in North America, the only holarctic endemic species with this antennomere count. Atopula. Emery (1912) created the genus Atopula to contain two species which he had described previously (Emery, I1go1) in Atopomyrmex André. These were A. nodifera (made the type-species of Atopula) from Cameroun, and A. ceylonica from Sri Lanka (which Emery referred to as ceylonensis in his 1912 publication). Wheeler (1919) recognized that these two species were different at generic level and, disagreeing with Forel’s (1917) placement of ceylonica in Leptothorax Mayr, erected a new genus Paratopula to hold ceylonica and its infraspecific forms. This procedure meant that Atopula at that time contained only its type-species, nodifera, but later authors added a few more names to the genus. Unfortunately the type-species, Atopula nodifera, is a quite ordinary tetramoriine, with all the diagnostic tribal characters, and furthermore the generic name Atopula is a straight synonym of Tetramorium. (Palp formula of 4,3 confirmed by dissection of BMNH specimen. Syntype worker of nodifera examined at MCZ, Cambridge.) Having established this point, the generic identity of the three remaining species of Atopula must now be clarified. Atopula belti (Forel) was placed in the monotypic genus Brunella Forel (1917), and for the time being I think that be/ti is best returned to Brunella until it can be investigated in detail. It is certainly not a tetramoriine and its affinities are unclear. ANT TRIBE TETRAMORIINI 363 Atopula longispina Stitz (1938) appears to be close to ceylonica and its allies and is transferred to Paratopula; thus its name is now Paratopula longispina (Stitz) comb. n. Atopula hortensis Bernard (1948) is a tetramoriine. In fact it is a direct junior synonym of the common African species Tetramorium sericeiventre Emery, and I am amazed that it was not recognized as such by Bernard as its identity is immediately obvious merely by examining the figures which he gives (Bernard, 1948 : 174, fig. 9). JT. sericeiventre is the most common arid-ground and semi- desert inhabiting species of its genus in the entire Ethiopian region and should be well known to anyone who is acquainted with the ant fauna of the region. Macromischoides. When Wheeler (1920) first introduced the name Macro- mischoides he cited a type-species (Macromischa aculeata Mayr) but did not define the genus. Later he gave a diagnosis and notes on the genus (Wheeler, 1922) and in the catalogue in the same publication he assigned the genus to the tribe Leptothoracini, in which it was retained by numerous later authors despite the fact that Santschi (1924) had demonstrated its tetramoriine character. Santschi’s findings are confirmed here and it is shown that Emery (1896d) was correct in assigning the known species of this group to genus Tetramorium. In his diagnosis of the genus Wheeler (1922) does not cite any characters which differentiate Macromischoides from the mass of Tetramorium except for the fact that the palp formula is reduced to 3,2 from the primitive maximum tetramoriine count of 4,3. Taken as it stands Wheeler’s definition of Macromischoides is really no more than a short description of its type-species, and the characters of the only other species which he included, africanum, are not noted. In fact africanum is summarily dismissed with the comment that it is ‘hardly more than a subspecies of aculeata’. This statement is incorrect and a close examination of africanum reveals that it shows numerous characters intermediate between aculeatum and the less specialized members of Tetramorium. In particular, the palp formula of africanum (4,2) is the main character which destroys the validity of Macromischoides as it is intermediate between the great mass of the genus (PF 4,3) and the specialized aculeatum (PF 3,2). All the other characters given by Wheeler are paralleled elsewhere in Tetramorium, particularly in the carinatum-group which, through adoption of a sub-arboreal lifeway, show similar modifications in body form but retains the palp formula of 4,3. A character cited by Wheeler (1922) for the males of Macromischoides concerns the antennomere count in the male of aculeatum, which Wheeler gives as eleven. He states that ‘Mayr claimed that the male aculeata has 11-jointed antennae, but Emery, after examination of six specimens, maintained that these appendages are 10-jointed’. Santschi (1924) proved that the funicular segmentation of the male showed considerable variation in degree of fusion of the antennomeres and this is confirmed in the present study, which also indicates that such variation is quite common throughout the males of this genus. Wheeler (1922) was of the opinion that Tetramorium became more homogeneous with the removal of aculeatum and africanum to a separate genus. I completely disagree with this statement as dividing up a genus by artificial and arbitrarily 364 B. BOLTON drawn lines often generates rather than removes confusion because the definition of the genus-group names thus produced becomes extremely difficult or impossible. This increased difficulty of definition is implicit in the process of splitting a genus by these means as it is usually accomplished by cutting out one or two species from the main mass of the genus which are said to show derived or specialized characters whilst leaving the closest relatives of the derived forms in the parent genus, from which they cannot be separated. The subgenera Lobomyrmex and Sulcomyrmex. Kratochvil (1941; 1944) attempted to overhaul the Tetramorium species of central Europe in general and Czechoslovakia in particular, and in doing so fell into the strange trap of splitting the few central European species into three subgenera, apparently without realizing that subgeneric names cannot be applied on such a limited basis but, once designated, apply to the entire world fauna and include all species of the genus which possess the characters delimited. As soon as this is given objective consideration his subgenera collapse in a welter of derivative forms and species or whole species- groups which overlap or refuse to comply with his subgeneric limits. The subgenus Sulcomyrmex was said to contain the species simillimum and guineense (both of which are tramp species not endemic in the Palaearctic region) and was defined by the fact that the frontal carinae extend back to the occipital margin. On this character Sulcomyrmex would contain the vast majority of the world fauna but unfortunately for the subgenus some species of the guineense-, pacificum-, simillimum- and many other species-groups exist in which the frontal carinae are reduced or lost in some species but not in others, as was recognized by Arnold as long ago as 1917. Consideration must also be given to the sqguaminode- group, the occidentale-group and the wettzeckeri-group. The members of these groups have strongly developed frontal carinae, which would place them firmly in Sulcomyrmex, but also have a petiolar and postpetiolar structure which would place them equally firmly in Lobomyrmex, the next to be discussed. In subgenus Lobomyrmex the frontal carinae are stated as being short, not extending to the occipital margin, and the pedicel is defined as having the ‘petiole dorsally almost as broad or only slightly narrower than the postpetiole’. In contrast to this the third subgenus, Tetrvamorium sensu stricto, as well as having short frontal carinae has, ‘petiole dorsally always distinctly narrower than post- petiole; if not distinctly then .. .’, and one is already on very unfirm ground, as can be deduced from the ambiguous nature of the couplet. Factually, all that this couplet set out to do was to separate T. ferox and its allies (subgen. Lobomyrmex) from T. caespitum and its allies (subgen. Tetryamorium sensu stricto) which in reality only belong to different complexes of species within a single very compact species- group. Applying the definitions of these subgenera to the rest of the world fauna produces unbelievable results as, in almost all groups in which the frontal carinae are reduced in all species (aculeatum-group, solidum-group, sericewentre-group), as opposed to the many groups in which only one or two species have the carinae reduced, some species fall into one subgenus and some into another, and again the question arises as to where to place the squaminode- and related groups. ANT TRIBE TETRAMORIINI 365 I think that this is sufficient to show the artificiality of these subgenera and the impossibility of dividing up the world fauna solely from the study of less than a dozen species from a restricted area of the Palaearctic region, especially when the main fauna of this genus is strictly tropical in its distribution. ACKNOWLEDGEMENTS I would like to express my thanks and gratitude tothe following persons, who have provided types or other material during the course of this study. Dr P. H. 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Raccolte mirmecologiche dell’Africa orientale conservate nel Museo Civico di Storia Naturale ‘“‘“Giacomo Doria” di Genova. Parte seconda. Formiche dell’Uganda e ANT TRIBE TETRAMORIINI 369 delle Isole Sesse raccolte dal Dr. E. Bayon. Annali Mus. civ. Stor. nat. Giacomo Doria 56 : 93-114, 4 figs. 1933. Le formiche della Palestina. Memorie Soc. ent. ital. 12 : 49-113, 5 figs. 1936. Nuovi contributi alla conoscenza della fauna delle isole italiane dell’Egeo. 6, Hym. Formicidae. Boll. Lab. Zool. gen. agy. Portici 24 : 262-311, 19 figs. PisarskI, B. 1965. Etudes sur les fourmis du genre Strongylognathus Mayr. Annales Zool. Warszawa 23 : 509-523, 44 figs. RoGeErR, J. 1857. Einiges tiber Ameisen. Berl. ent. Z. 1 : 10-20. Ruzsxy, M. 1903. Die Ameisenfauna der Astrachanischen Kirgisensteppe. Tvudy russk. ent. Obshch. 36 : 294-316. SANTSCHI, F. 1916. Descriptions de fourmis nouvelles d’Afrique et d’Amerique. Anmnls Soc. ent. Fr. 84 : 497-513, fig. 1917. Fourmis nouvelles de la Colonie du Cap, du Natal et de Rhodesia. Annls Soc. ent. Fr. 85 (1916) : 279-296. 1921. Notes sur les fourmis paléarctiques. Boln R. Soc. esp. Hist. nat. 21 : 110-116, I fig. 1923. Descriptions de nouveaux formicides éthiopiens et notes diverses 1. Revue Zool. Bot. afr. 11 : 259-295, 5 figs. 1924. Descriptions de nouveaux formicides éthiopiens et notes diverses 2. Revue Zool. Bot. afr. 12 : 195-224, Io figs. 1925. Nouveaux formicides brésiliens et autres. Bull. Annls Soc. ent. Belg. 65 : 221-247. 1927. A propos du Tetvamorium caespitum L. Folia Myrm. et Term. 1 : 52-58. 1932. Formicides Sud-Africains. Soc. ent. Fr. livre du centenaive 1932 : 381-392, 12 figs. 1935. Fourmis du Musée du Congo Belge. Revue Zool. Bot. afy. 27 : 254-285, 40 figs. 19374. Fourmis angolaises. (Résultats de la mission scientifique suisse en Angola, 1932-33.) Revue suisse Zool. 44 : 209-248, 48 figs. 19370. Résultats entomologiques d’un voyage au Cameroun. Formicides récoltés par M. le Dr. F. Zumpt. Mitt. schweiz ent. Ges. 17 : 93-104, 5 figs. ScHENK, C. F. 1852. Beschreibung Nassauischer Ameisenarten. Jb. ver. Naturk. Nassau 8 : 1-149. 1853. Die Nassauischen Ameisen-Species. Stettin. ent. Zig 14 : 185-198. Siruavy, V. 1937. Strongylognathus kratochvili n. sp., une fourmi nouvelle, reliquat du temps préglacial de Moravie, Tchécoslovaquie. Sb. priv. Klubu Tyebici 1 : 1-12, 6 figs. SmiTH, F. 1851. List of the specimens of British animals in the collection of the British Museum. 6 Hymenoptera Aculeata. 134 pp. London. 1861. Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the islands of Ceram, Celebes, Ternate and Gilolo. J. Linn. Soc. (Zool.) 6 : 36—66, pl. 1. SmiTH, M. R. 1953. A revision of the genus Romblonella W. M. Wheeler. Proc. Hawaii ent. Soc. 15 : 75-80, 1 fig. — 1956. A list of the species of Romblonella including two generic transfers. Bull. Brooklyn ent. Soc. 51 : 18. Stitz, H. tg10. Westafrikanische Ameisen 1. Mitt. zool. Mus. Berl. 5 : 125-151, 11 figs. 1912. Ameisen aus Ceram und Neu-Guinea. Sber. Ges. naturf. Freunde Berlin 1912 : 498-514, 14 figs. 1938. Neue Ameisen aus dem indo-malayischen Gebiet. Sber. Ges. naturf. Freunde Berlin 1938 : 99-122, 13 figs. STUMPER, R. 1950. Les associations complexes des fourmis. Commensalisme, symbiose et parasitisme. Bull. Biol. Fr. Belg. 84 : 376-399. URBANI, C. BARONI. 1962. Studi sulla mirmecofauna d’Italia 1. Redia 47 : 129-138, 3 figs. 1968. Studi sulla mirmecofauna d’Italia 4. La fauna mirmecologica delle isole Maltese ed il suo significato ecologico e biogeografico. Annali Mus. civ. Stor. nat. Giacomo Doria 77 : 408-559, 52 figs, 12 pl. 1969. Gli Strongylognathus del grupo huberi nell’Europa occidentale: saggio di una revisione basata sulla casta operaia. Boll. Soc. ent. ital. 99-101 : 132-168, 16 figs. HITT | 370 B. BOLTON VIEHMEYER, H. 1914. Ameisen aus Perak, Bali und Ceram. Ent. Mitt. 3 : 112-116, 2 figs. 1922. Neue Ameisen. Avch. Naturgesch. 88 (A) 7 : 203-220, 2 figs. WEBER, N. A. 1943. Ants of the Imatong Mountains, Anglo-Egyptian Sudan. Bull. Mus. comp. Zool. Harv. 93 : 261-389, 16 pls. WHEELER, W.M. 1909. Observations on some European ants. Jl N. Y. ent. Soc. 17 : 172- 187, 2 figs. 1910. Ants, their structure, development and behaviour. 663 pp., 286 figs. New York. 1911. A list of the type-species of the genera and subgenera of Formicidae. Ann. N. Y. Acad. Sci. 21 : 157-175. 1914. The ants of the Baltic Amber. Schr. phys.-dkon. Ges. Kénigsberg 55 : 1-142, 66 figs. 1919. The ants of Borneo. Bull. Mus. comp. Zool. Harv. 63 : 43-147. 1920. The subfamilies of Formicidae, and other taxonomic notes. Psyche, Camb. 27 : 46-55. 1922. Ants of the Belgian Congo, parts 1-8. Bull. Am. Mus. nat. Hist. 45 : 1-1139, 23 pis, 76 figs. 1927. Ants collected by Professor F. Silvestri in Indochina. Boll. Lab. Zool. gen. agr. Portici 20 : 83-106, 9 figs. 1929. Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Boll. Lab. Zool. gen. agr. Portici 24 : 27-64, 7 figs. 1930. Formosan ants collected by Dr. R. Takahashi. Proc. New. Engl. zool. Club 11 : 93-106, 2 figs. 1935. Two new genera of myrmicine ants from papua and the Philippines. Proc. New Engl. zool. Club 15 : 1-9, 2 figs. Witson, E.O. 1971. The insect societies. 548 pp., 22 figs. Cambridge, Mass. & Taytor, R. W. 1967. The ants of Polynesia. Pacif. Ins. Monogr. 14 : 1-109, 84 figs. ANT TRIBE TETRAMORIINI 71 Wy @ @ Fics 1-12. Rhoptromyrmex species. 1-7. Outline of head capsule of (1) R. globulinodis, worker, (2) R. melleus, worker, (3) R. critchleyi, worker, (4) R. globulinodis, female, (5) R. tvansversinodis, female, (6) R. globulinodis, male, (7) R. wroughtoni, male. 8. Right forewing of R. transversinodis, male. g—12. Outline of alitrunk and pedicel. in worker of (9) R. globulinodis (offset shows pedicel segments in dorsal view), (10) R. tvanversinodis (offset shows pedicel segments in dorsal view), (11) R. melleus (12) R. critchleyt. 372 B. BOLTON Fics 13-18. Strongylognathus species. 13-15. Outline of head capsule of (13) S. testaceus, worker, (14) S. destefaniz, worker, (15) S. testaceus, male. 16. Right forewing of S. testa- ceus, male. 17, 18. Outline of alitrunk and pedicel in worker of (17) S. testaceus, (18) S. destefanii. Fics 19-23. Anergates atvatulus. 19, outline of head of female; 20, same of male; 21, pedicel of female; 22, same of male; 23, right forewing of female. Figs 24-26. Teleutomyrmex schneider. 24, outline of head of female; 25, outline of pedicel of female; 26, right forewing of female. ANT TRIBE TETRAMORIINI pl oe San alliage [~~ 27 ee) ms oe) 6) 2) @>) Fics 27-31. Decamorium decem. 27, outline of head of worker; 28, diagram of worker mandible showing variation in denticles; 29, outline of head of male; 30, right forewing of male; 31, outline of alitrunk and pedicel of worker. Fics 32-41. Tviglyphothrix species. 32-35, forms of hairs encountered in the genus; (32) simple, (33) bifid, (34) trifid, (35) quadrifid. 36, diagram of dentition mostly commonly encountered in the genus. 37, diagram of head to show row of simple long hairs on upper scrobe margins. 38, hind tibial pilosity in T. obesa and allies. 39, hind tibial pilosity in T. pacifica and allies. 40, outline of head of T. inezulae, male. 41, right forewing of T. menkaura, female. 374 B. BOLTON JAY) | Sr 45 D 47 Fics 42-57. Tvriglyphothrix species. 42-51. Outline of heads of workers of (42) T. roths- childi, (43) T. microps, (44) T. eminii, (45) lateral view of T. eminii to show extent of scrobe, (46) IT. menkaura, (47) lateral view of T. menkaurva to show extent of scrobe, (48) T. dedefra, (49) T. baufra, (50) T. brevispinosa, (51) T. meshena. 52-57. Outline of alitrunk and pedicel of workers of (52) T. vothschildi, (53) T. microps, (54) T. eminit, (55) I. menkaura, (56) T. dedefra, (57) T. baufra. : ANT TRIBE TETRAMORIINI 375 64 Figs 58-65. Outline of alitrunk and pedicel in Tviglyphothrix species, workers. 58, T. brevispinosa; 59, T. pulcherrima; 60, T. gabonensis; 61, T. osivis; 62, T. fulviceps; 63, T. lanuginosa; 64, T. parvispina; 65, T. coonoorensis. 376 B. BOLTON oA 2 , | ee Fics 66-73. Outline of alitrunk and pedicel in Tviglyphothrix species, workers. 66, T. adpressa; 67, T. meshena; 68, T. walshi; 69, T. brevidentata; 70, T. pnyxis; 71, T. khe- perva; 72, T. chepocha; 73, T. vombis. ANT TRIBE TETRAMORIINI INDEX Synonyms are in ?ttalics. Acidomyrmex, 298 adpressa, 344 afer, 306 agna, 317 akermani, 324 alberti, 302 alboini, 306 alpinus, 306 Anergates, 296 antennata, 344 antrema, 318 aveolatus, 326 arnoldi, 318 Atopula, 359 atratulus, 296 auvopunctatus, 329 australis, 350 baufra, 319 belti, 362 boulognei, 326 braunsi, 292 brevidentata, 345 brevispinosa, 320 bulawayensis, 329 bulgaricus, 306 burgeoni, 326 caeciliae, 307 cecconii, 307 Cephalomorium, 287 ceramensis, 350 ceylonica, 362 Chelaner, 294 chepocha, 345 christophi, 307 cinereus, 324 constanciae, 321 coonoorensis, 346 critchleyi, 303 cryptica, 321 dalmaticus, 307 decamera, 347 Decamorium, 297 decem, 298 dedefra, 322 desertorum, 323 destefanii, 307 distincta, 323 divert, 306 Dyomorium, 287 emarginatus, 306 emeryi, 308 eminli, 324 evicae, 333 esta, 303 Eutetramorium, 293 felix, 350 flavescens, 350 flavigaster, 294 foreli, 308 formosae, 355 friedlandi, 297 fulviceps, 347 furtiva, 325 fusciventris, 329 gabonensis, 325 gallica, 308 gestroi, 326 gibber, 292 globulinodis, 302 grandinodis, 294 guillodi, 324 hepburni, 328 hortensis, 363 huberi, 308 imbellis, 327 indosinensis, 348 inezulae, 327 insularis, 308 intermedia, 302 Iveneella, 298 Ireneopone, 292 italicus, 308 hamerunensis, 326 karawajewi, 305 katypa, 349 kervillei, 308 kheperra, 349 koreanus, 308 kratochvili, 307 huttert, 307 378 laeviceps, 303 laevidens, 350 lanuginosa, 350 leno, 302 Lobomyrmex, 359 longi, 302 longispina, 363 longispinosa, 321 Lundella, 287 Macromischoides, 359 marley, 324 marthae, 324 mashonana, 328 mauricer, 351 mayri (Triglyphothrix), 352 mayri (Rhoptromyrmex), 301 melleus, 301 menkaura, 328 meshena, 353 microps, 329 minima, 330 mocquerysi, 293 monardi, 331 monodt, 303 monticellii, 293 mucidus, 326 MUS, 324 muscorum, 331 musculus, 358 Myrmus, 304 nacta, 353 nimba, 298 nion, 320 nodifera, 362 obesa, 354 obscurus, 302 opaca (Romblonella), 294 opacus (Rhoptromyrmex), 302 ovissana, 350 osiris, 332 pacifica, 355 palaestinensis, 309 pallens, 329 papuensis, 302 parvispina, 355 paupera, 333 pnyxis, 356 pretoriae, 303 pulchella, 356 INDEX pulcherrima, 333 rehbinderi, 309 reptana, 334 vhodesiana, 329 Rhoptromyrmex, 298 Romblonella, 293 rossi, 357 vothneyt, 302 rothschildi, 334 ruzskyi, 309 schneideri, 310 sericea, 335 silvestrii (Stronglyognathus), 309 silvestrii (Triglyphothrix), 336 simoni, 292 solleri, 301 soyansi, 326 SOVAUXL, 326 spuria, 358 stein, 303 striatidens, 350 Strongylognathus, 304 Sulcomyrmex, 359 sumatrensis, 302 taivanensis, 302 talpa, 336 Teleutomyrmex, 309 tenebrosa, 337 Tetramorium, 359 Tetramyrma, 291 Tetrogmus, 359 tessmannt, 301 testaceus, 306 thoth, 337 transversinodis, 303 tricolor, 351 Triglyphothrix, 310 trimeni, 338 uelense, 298 ultor, 298 vombis, 358 walshi, 358 wroughtonii, 302 Xiphomyrmex, 359 yarthiella, 338 ANT TRIBE TETRAMORIINI 379 B. Botton, B.Sc., A.R.C.S. Department of Entomology BritisH Museum (NaTuraL History) CROMWELL ROAD Lonpon SW7 5BD sh ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxapDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. {3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3-15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50. . HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounp, L.A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . AFiFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Exiot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. {12. . SANDS, W. A. The Soldieriess Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . VON Hayex, C. M. F. A_ Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. 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