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Hath Hats aie sa RESHRSaiREEs iit atti sts rete ait ota . teh - ae, itinerant 3 Sere patyrar teeta . if Hee ‘ nt 4 ore it Haale anette H Siigsit " iat it ttt Meindl Mitte a ia Ha ;$. Pthes attests? sai 1) + ; itt si a38t3 ttt ies tite f RRO aesstatassteaie atte ‘Shane iti tr ith Hatta Heat ‘| i - He +3 $3 ni i - > _ _* i wet. oe — _ BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY VOEE ES 1967 BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1967 DATES OF PUBLICATION OF THE PARTS No. I Z oo OS Oa wD 16 January 1967 28 February 1967 . 18 April 1967 . 18 April 1967 2 June 1967 2 June 1967 PRINTED IN GREAT BRITAIN BY ADLARD AND SON LIMITED DORKING, SURREY CONTENTS ENTOMOLOGY VOLUME XIx The Phlebotomine sand-flies of West Pakistan (Diptera: Psychodidae). By D. J. LEwis A revision of the genus Palorus (sens. lat.) (Coleoptera: Tenebrionidae). By D. G. H. HALsTEAD A survey of the extra-Ethiopian Oretinae (Lepidoptera: Drepanidae). By A. WATSON Collecting in Turkey 1959, 1960 and 1962. By K. M. GUICHARD & D. H. HARvEY Hymenoptera from Turkey. Sphecidae, I. By J. DE BEAUMONT Appendix Sphex Linné, Subgenus Palmodes Kohl. By P. RotH Hymenoptera from Turkey. Sphecidae, II (Genera Astata Latreille and Tachysphex Kohl). By W. J. PULAWSKI Index to Volume XIX 149 223 253 383 411 tate THE PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN (DIPTERA : PSYCHODIDAE) D. J. LEWIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 1 LONDON : 1967 THE PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN (DIPTERA : PSYCHODIDAE) BY D. J. LEWIS | Scientific Staff, Medical Research Council, Lgndon, c/o British Museum (Nat. Hist.) Pp. 1-57, 136 Text-figs., 1 Map BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 1 LONDON : 1967 THE PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN (DIPTERA : PSYCHODIDAE) BY D. J. LEWIS Scientific Staff, Medical Research Council, Lgndon, c/o British Museum (Nat. Hist.) Pp. 1-57, 136 Text-figs., 1 Map BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 1 LONDON : 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), imstituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical serves. 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. 19, No. i of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation: Bull. Br. Mus. nat. Hist. Entomology. © Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 16 January, 1967 Price {1 3s. THE PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN! (DIPTERA : PSYCHODIDAE) By D. J. LEWIS CONTENTS Page INTRODUCTION . : p . ; : : : : 3 : 3 METHODS ‘ 4 , ‘ ‘ d : , : A : 4 CLASSIFICATION : ; . ; : ; : : 3 : 6 KEYS TO SPECIES KNOWN FROM WEST PAKISTAN. ; ; ; : 8 TAXONOMY AND DISTRIBUTION OF THE SPECIES ‘ PF : , : 12 ZOOGEOGRAPHY . F : ‘ ‘ Z ‘ ‘ ; ; ; 46 BIOLOGY . é : é : ‘ : : : : : : 49 RELATION TO DISEASE ; : 4 F : : ‘ ‘ : 51 ACKNOWLEDGMENTS . i : , : : d ; F ; 52 REFERENCES. : : . ‘ ‘ ; : ‘ : 4 53 INDEX TO SPECIES AND SUBSPECIES , : é , P : d 57 SYNOPSIS Some aspects of classification are discussed, and keys to the known species are given. Most of the species are described or redescribed, and distribution lists are provided. Zoogeography is discussed with special reference to the 29 species and one local subspecies found hitherto. The interpenetration of eastern and western faunas has been increased by the Himalayan mountain system. Notes are given on aspects of biology and relation to human disease. INTRODUCTION In May and June 1959 Professor H. C. Barnett collected many thousands of sand- flies during studies on sand-fly fever and other viruses (Barnett & Suyemoto, 1961) and kept as samples 9,900 which he asked me to identify, together with specimens collected in the Keris area by Lieut. Col. M. I. Burney and Lieut. Col. J. E. Scanlon. Professor Barnett arranged with the Pakistan Medical Research Centre for me to make a survey from 13th May to 30th June, 1963, and I visited Lahore, Rawalpindi, Taxila, Peshawar, Saidu Sharif, Bahrein, Kalam, Abbottabad, Nathia Gali, Gilgit, Chilas, Skardu, Keris, Karachi and neighbouring areas. Mr. W. A. McDonald contributed sand-flies from Lahore, Gujrat and Mir Muhammad. All the specimens numbered 11,100, and I have also examined many in the British Museum collected by the late Brigadier J. A. Sinton during his pioneer studies of the Phlebotominae. 1 This investigation was supported in part by U.S. Public Health Service Research Grants TW-—oo01 42-06 and A1—05589—03 from the Division of General Medical Sciences and the National Institute of Allergy and Infectious Diseases, National Institutes of Health, respectively. The work is based largely on collections made by Professor Herbert C. Barnett, now Director of the Institute of International Medicine, University of Maryland, Baltimore, U.S.A., and on a survey made by the author under the auspices of the Pakistan Medical Research Centre, Lahore. ENTOM. IQ, I, I 4 D. J. LEWIS West Pakistan (Text-fig. 1), including northern Kashmir, comprises the great alluvial plain of the Indus and its tributaries in Sind and West Punjab, the hills and plateaux south-west of Peshawar, and the high mountains and valleys of the north. The northern mountains radiate from the Pamirs and comprise parts of the Hindu Kush, Karakoram and Himalayan ranges. South of the Himalayas the climate is tropical and is dominated by the rhythm of the monsoons, but the rainfall is low and much of the area is cold in winter. North of the Punjab there are intermittent wet days in summer, and forest begins at about 1400 metres. The Himalayas stand in the path of the monsoon, so that the Karakoram is part of the central Asian desert. In the Gilgit and Skardu areas the general arid conditions contrast with narrow valleys of streams fed by vast areas of melting snow. Weak depressions from the Mediterranean (Bharadwaj, 1961) cause light winter rains in the north and some of the Himalayan snowfall. The area is described in the Imperial Gazetteer of India (1907) and briefly by Schmid (1958). Prater (1965 : 21) indicates the plant zones, and Mani (1962), Lorimer (1939) and Maraini (1961) describe the western Himalayas and the country around Gilgit and Skardu respectively. METHODS Collecting. Sand-flies resting in houses were usually collected with a suction catcher, and some were taken by Mr. McDonald around Mir Muhammad in routine pyrethrum-spray catches of mosquitoes and on cattle. Most outdoor catches in 1963 were made with sticky traps. They consisted of pieces of paper about I0 X 16:5 cm. smeared with castor oil and supported by cleft sticks about 28 cm. long, and were placed before dusk near animal burrows, rock crevices and other likely places. Two hundred and fifty ml. of oil sufficed for about roo traps. Five hundred and forty were set, on 25 nights, and yielded 419 flies (from o to 2°6 per trap per locality—average 0-8), whereas the maximum on a single trap in Iran in June, 1960 (Lewis e¢ al., 1961) was 132. The low numbers in Pakistan were probably due partly to the preceding cold winter and, near Karachi, to wind. Some sand-flies were driven from termite hills with tobacco smoke and caught in a fine-mesh black net, and a few were taken in a Damasceno-type trap, but this is more suitable for forest country where it can be slung from trees. Preservation. Many specimens were preserved dry in vials or cardboard boxes, with tissue paper or cotton-wool to prevent shaking. Others were stranded in plastic vials and wetted with a few drops of gum-chloral mounting medium. Mounting. Dry specimens were put in tap water containing one per cent of domestic detergent (to wet them), which was heated to about 85° C. for four.minutes to remove bubbles. Specimens were roughly sorted according to external characters, including size, colour and, in the case of male Sergentomyza, the shape of the abdomen. Most of the flies were mounted in gum-chloral medium composed of distilled water (20 ml.), gum acacia (16 gm.), chloral hydrate (140 gm.), glycerol (12-6 gm.) and acetic acid (6:3 gm.), beneath circular cover-glasses I cm. in diameter and o-r mm. thick. The mounts were allowed to dry partially for a few months, or kept over silica gel PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 4 ORD a aE: RIFE: OETA ER 7 CET, 7. Keris ““Gwadi “. KAR AKORAM~% ordu * Parkuta_”’,, 77, ERAT ¢ : uO. °7 s LET G7, LOLIG EAL LPL CO Ue Rtg earn sre o t dee) tT. 4144 SRGLF CUED GMOS 2 GIG CONE dE se cob av OR xan / TOUS THT L, SUS ; ies D A Coen we. ./ AN s Breton ae Lon ey du Sharit. 7? \. ©, Khaira Gali a hee . : we: ie ie “ \.Murree > .). + Said . . Nowshera -| : joxila Rawalpindi Peshawar * : Jhelur - ‘ Landi Kotal - ore : +s Kohate Seis ee <~“Hangue” .. . )- pet LILA, Py eer 7 rf 104457 METRES, |7777 Ad / V6 ay a fi Pp / NA ade 4 APPROX.N/U 7 7 Tonk Cade oar a Lee reg 04 / ty Jandola, * Lelany “sit Arai Khirai deg tigh cule Vyt a : rg Ct Saad 2000 -—~+ 7 AAMAS IRISTAN . ee / Meas, AS, Sa, r Lipa, 200 aX LEG rt ° ifatere KILOMETRES 1 * Kashmore 1 si x Kandhkot* eA) ePano Aqil : Shikarpur, (sukkur H A, Rohri 1 x ° *Khairpur \ Larkana e MS Qambar® Ss!tno Mirpur Khas \y * Jamesabad : i Ne i SLY = Yoe oN *Digri See te 3 _ Hyderabad \, Jando Muhammad Khan 3 Tando Bago Go aes el t cones a \ Ne KILOMETRES ° 1000 \ N Fic. 1. Showing places mentioned in the text. 6 D. J. LEWIS for a few days, and ringed with Euparal. An oven was not used because it involves the risk of over-drying or overheating. Three beads, each Imm. high, were attached with Euparal to each slide to protect the cover glasses, so that many slides could be stored in drawers without slots. In mounts of this kind the specimens remain soft and flexible, and suitable for reorientation of particular structures. The soft parts become invisible without being destroyed, as they are by caustic potash, so that a specimen keeps its shape and can even be sectioned to show the position and general structure of internal organs. The cibarium and pharynx were examined 7m situ in ventral view. The sper- mathecal ducts of many previously mounted specimens were examined. For this purpose they were extracted by wetting, and mounted temporarily in an equal mixture of water and gum-choral to give a suitable refractive index. In the case of some very delicate species tergite 9 and sternite 8 were removed. In damp climates where gum-chloral mounts may require too much attention fairly good results can be obtained by mounting in Euparal after dehydration in alcohol up to 95 percent. Various methods of staining after maceration give a very clear picture of certain chitinous structures but they hide colour variation and the natural appearance of the insect as seen in freshly dissected flies. CLASSIFICATION The term sand-fly is used here for species of the tribe Phlebotomini, as defined by Fairchild (1955), other than Hertigia and Warileya. The classification of the sand-flies, the names of certain structures, and various taxonomic expressions are discussed by Kirk and Lewis (1951), Fairchild (1955), Theodor (1958), Quate (1964), Abonnenc & Minter (1965) and others. At present some authors place all sand- flies, of both Old and New Worlds, in one genus Phlebotomus, and others divide them into several genera. The system of Theodor (1948, 1958) is used here. It is often hard to delimit species and subspecies of Phlebotominae. Cross- mating is difficult, the early stages of most species are unknown, and knowledge of distribution is limited by the need for special collecting methods. Furthermore, some characters are less precise than they seem. For instance, when the cibarial teeth are counted 7m situ in the normal way some lateral ones may be hidden. If the head is flattened the specimen is damaged and the lateral teeth may be found to merge imperceptibly into spicules and become impossible to count exactly. Measure- ments of the pharynx are often used, but it may be soft and flexible and its hind end is difficult to define. Various difficulties are illustrated by the recent treatment of the Sergentomyia squamipleuris complex by experienced entomologists. Theodor and Mesghali (1964) recognized four species and one subspecies, and Quate (1962a, 1964) placed all forms in one species. Fortunately, however, the small size of sand- flies and the practice of mounting them all on slides make it possible to amass large but compact collections for the study of variation and distribution, and the cibarial teeth provide useful characters for most Old World species. Morphology. This is described in several standard works, and the following notes refer to particular features. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 7] All the scales of at least one species have petioles (Nicoli, 1956), and many of the petioles in two species (P. kandelakii and P. chinensis) are distinctive. There is some variation in the cibaria of Phlebotomus species, particularly the degree of visibility of the spicules or teeth, the hind margin of the ventral plate, and the chitinous arch. The cibarial teeth of most species of Sergentomyia are clearly seen but those of S. batlyi are much less conspicuous than those of P. colabaensis. The punctiform teeth, denticles, or nodules in front of the main cibarial teeth of Sergentomyia are often called vertical teeth but this term is seldom used here because they have also been termed horizontal teeth. The posterior bulge (Text-fig. 9) in the dorsal wall of the cibarium of most American phlebotomines has been used by Theodor (1965a) as an important character for distinguishing them from Old World sand-flies. This seems to be a useful character but should be used with care because some Pakistan and other Old World species, particularly in the genus Phlebotomus, have a somewhat similar bulge (Text-figs. 3, 6, 8, 35, 84, 85 etc.). In most of these species the bulge is usually asymmetrical and evidently flexible, so that its appearance depends on method of preparation and angle of view. It tends to be less pronounced than in American species, and dome- shaped rather than mitre-shaped when seen in lateral view after removal of the sides of the head. In Phlebotomus there is an additional bulge in the region of the chitinous arch. The length of the labrum was measured from the tip of the clypeus as seen in ventral view. This measurement, which is the approximate length of the structure, is often expressed as the length of the epipharynx, but this terminology is not correct because the epipharynx of insects is the ventral surface of both labrum and clypeus (Matsuda, 1965). In male sand-flies the tip of the labrum is surmounted by a crest, as it may be called, which Grassi (1907) described as a laminetta in P. papatast. Christophers et al. (1926) referred to it as a “ peculiar flattish pad ’”’ in P. argentipes, which arises from the median chitinisation. They pointed out that near the base of the labrum this chitinisation lies between the two dilated parts of the lateral pieces, which form the basal bulge. Perfil’ev (1937) observed that the crest was rounded in a species of the subgenus Sergentomyia and terminally spiculated in P. chinensis. The crest often rises to a point which may be termed the summit in distinction from the terminal apex. In females the crest is lacking (Text-fig. 118) and would presumably prevent the labrum from piercing skin. There is a rather indefinite relation between types of crest and certain subgenera or groups of subgenera. In the genus Phleboto- mus the summit hardly ever projects forward, and spicules are easy to see, partly owing to the size of the species; in subgenera Phlebotomus and Paraphlebotomus the summit is well defined. In the subgenus Servgentomyia the crest is rather similar but the spicules are usually inconspicuous. In most species of Parrotomyia the crest tapers and lacks a definite summit. In the two Pakistan species of Grassomyia and Rondanomyia, and in most Sintonius, the summit is rounded and projects forward. There are minor specific features in some cases, and certain species are characterized by other aspects of the labrum, the size and degree of 8 D. J. LEWIS pubescence of the basal bulge, and the width of the lateral flanges (as seen in lateral view). It is interesting to note that unusual types of labrum occur in the rather unusual species S. sguamipleuris, S. pawlowskyi and S. bailyi. In the palpal ratio the first number indicates the combined lengths of segments 1 and 2, according to Quate’s suggestion, and segment 5 is omitted in some species. Text-fig. 29 shows the junctions of the segments. Palpal segments were not always measured very accurately because this would have involved remounting or flattening many specimens. Wing lengths were measured from the proximal end of the hairy basal bulge of the costa (Text-fig. 73). According to Quate’s (1964) suggestion the terms Ry, R5,3, and R, tip are used instead of alpha, beta and delta. In the males of most African species of Sergentomyia, other than subgenus Sintonius, the abdomen tapers toward the hind end. In a number of Pakistan species, however, tergite 6 is more or less enlarged so that the terminalia spring from a truncated end. This enlargement is often associated with absence of true hairs and the presence of large microtrichia. The hair-like structures at the apex of the spermatheca are here termed gland ducts. As suggested by Quate, the terms “ basistyle ’’ and “ dististyle ’’ (discussed by Van Emden and Hennig, 1956, and Prasad and Grover, 1963) are used for “ coxite ’’ and “ style ’’, and “‘ surstyle ”’ for “‘ lateral lobe ”’. The name “ sand-fly’’. According to Murray (1888) this term was published for the first time by Walter (1748) who used it for insects which attacked members of Anson’s expedition on the island of Santa Catarina, Brazil, in 1740. These were probably Culicoides, breeding in water but biting near sandy beaches. The name evidently did not originate from the Portuguese word for these flies or from the vernacular word discussed by Lane (1942) in his comments on the work of G. Marcgravius which was first published in 1648. The name “‘sand-fly ’’ came to be used in several parts of the world, including India (Adams, 1867, p. 59 ; Annandale, Ig10), for various small biting flies. It became adopted as the usual English word for phlebotomines in some of the dry parts of Pakistan, India and the Mediterranean area, probably because the dusty conditions made the name seem appropriate, man-biting midges and simuliids were generally scarce or absent, and the extensive researches on leishmaniasis in these areas focussed attention on the phlebotomines. In Iran they are called pasheh khaki, “ earth-coloured mosquitoes ’’, but the English name for these inconspicuous flies does not seem to be due to their colour. KEYS TO THE SPECIES KNOWN FROM WEST PAKISTAN The following keys are based partly on Theodor’s (1958) keys to the Palaearctic species, and should be used with the knowledge that additional species may be found in West Pakistan (Table I). KEY TO GENERA Cibarium unarmed or with scattered spicules, without pigment patch. Hairs on hind ends of abdominal tergites 2-6 erect, their sockets as large as on 1. Dististyle with 4 or 5 spines. : : : : ; ‘ : ‘ : : PHLEBOTOMUS PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN Cibarium with I or more rows of teeth (scarcely visible in S. bailyi), pigment patch usually present. Hairs on hind ends of abdominal tergites 2-6 recumbent, sockets much smaller than on 1, except in a few species which have a few erect hairs. Dististyle with 4 large spines and an accessory ventral seta TABLE I. Some Records of Species which May Occur in West Pakistan. Distance from Species Pakistan, Reference km. Phlebotomus bergeroti Parrot 163 Theodor & Mesghali, 1964 salehi Mesghali, 1965 380 Mesghali, 1965 jacusieli Theodor 527 Mesghali, 1965 mofidit Theodor & Mesghali 740 Theodor & Mesghali, 1964 mongolensis Sinton 373 Dolmatova et al., 1962 eleanorae Sinton 260 Sinton, 1931a eleanorae Sinton 4605 Mesghali, 1965 newsteadi Sinton 212 Sinton, 1926 Sergentomyia sintoni Pringle 315 Mesghali, 1965 p. pawlowskyi Perfil’ev 515 Mesghali, 1965 zeylanica Annandale 22 Sinton, 19285 KEY TO FEMALES OF PHLEBOTOMUS (except P. nuri) Pharyngeal teeth small and punctiform, in rows or curved groups Pharyngeal teeth large, spermathecae without long process . ‘ Spermathecae with 30-35 segments, very long. Median pharyngeal teeth larger than lateral ones . : : : ; ‘ kandelakii burneyi (p. Spermathecae with 8—21 segments Spermathecae nearly cylindrical, with 12-16 seyments. , : major (p. Spermathecae with about 18-21 segments, narrowing at one or both ends : Spermathecae with about 18 segments, narrowing towards duct and broad at other end. Hind part of pharynx with small anterior toothed scales and posterior small punctiform teeth . ; : : keshishiani (p. Spermathecae with about 21 segments, narrowing at both ends. ; sp. A (p. Spermathecae incompletely segmented, with indistinct transverse striations chinensis longiductus (p. Spermathecae segmented . Pharynx armed with finely toothed scales 0 ora network of lines , . papatasi (p. Pharynx armed partly or wholly with strong backwardly directed teeth or scales Pharyngeal armature with an anterior median group of strong scales or spines, and some scales forming concentric lines é Pharyngeal armature composed of large backwardly directed teeth ' Spermathecae with terminal segment much larger than the rest, ducts about 3 dimes as long as spermathecae : ; : , ; argentipes (p. Spermathecal segments equal, ducts very long ‘ : colabaensis (p. Antennal segment 3 short (0-12—0-16 mm.), 0-5—0-6 length of labrum alexandri (p. Antennal segment 3 long (0:23-0:33 mm.), 0-7—1 length of labrum . sergenti (p. SERGENTOMYIA Io PO) cod [| w f{ SWS hoe FS Ve Pa | D. J. LEWIS Key To Mates or PHLEBOTOMUS (except P. sp. A) Basistyle with a hairy process near its base. Genital filaments short, I-3—2-3 times length of pump : Basistyle without such process . : : ; ; ; : i ‘ Basistyle long (0-37-0-63 mm.); basal process small, with few hairs. Dististyle long and cylindrical with 5 spatulate spines, 3 of them terminal. Paramere with 2 long dorsal processes. Surstyles with terminal spines . : . papatasi (p. Basistyle short (0-2—0-33 mm.); basal process large or very large with many hairs. Paramere simple without upper process, with flat elliptical upper surface Basal process of basistyle very large and thick, with many hairs on its distal third nuri (p. Basal process of basistyle small and thin, with hairs only at the end Antennal segment 3 short (0-12—0-16 mm.), 0-7—0-9 length of labrum. Genital pump short (0-12 mm.), with small basal plate . : : alexandri (p. Antennal segment 3 long (0-25—0-34 mm.), I-1-4 times length of labrum. Genital pump large (0-17—0-2 mm.), with broad basal ae or funnel . . sergenti (p. Dististyle with 4 long spines ‘ ; colabaensis (p. Dististyle with 5 long spines, 2 at the end and 3 in the middle. Genital filaments long, 3-11 times length of pump Paramere with 2 ventral lobes; 2 long slender spines o on each side of the aedeagus ae (p. Paramere without ventral lobes Mid-ventral surface of aedeagus finely serrated, aedeagus tapering gradually toa point through which the genital filaments emerge . . kandelakii burneyi (p. Aedeagus smooth : ‘ : ; ; ; ; Genital filaments 3—5 anes length of pump ; : : ; : major (p. Genital filaments 6-11 times length of pump ; : ; . : ‘ Aedeagus with subterminal spike or tooth . ‘ P chinensis longiductus (p. Aedeagus without subterminal spike, narrow ; : ; ; keshishiani (p. Kry To FEMALES OF SERGENTOM YIA Spermathecae segmented, ducts long and narrow . Spermathecae not segmented but may be indistinctly striated, ducts not long and narrow Cibarium with about 50 teeth or more ‘ : : : : ; hospitii (p. Cibarium with about 18 teeth or less. Cibarium with about 5 widely spaced teeth ; : ‘ . Christophersi (p. Cibarium with about 12-15 teeth close together . ‘ ‘ , : : Cibarium with 12—15 long equal pointed teeth in a straight line. Pharynx with thick walls and an abrupt constriction behind the bulge. 1-4 papillae on antennal: segment 3, I-20n4 . ‘ clydei (p. Cibarium with 12 teeth, their ‘points directed upward and usually hidden, outer teeth sloping toward centre. Pharynx thin-walled, narrowing gradually behind bulge. 1 papilla on antennal segments 3 and 4 3 : ; tiberiadis (p. Spermathecae with rounded capsules and external spicules. Cibarium with 33-36 parallel teeth in a comb-like row convex posteriorly. Pharynx bulging posteriorly, with well-developed teeth. Antennal segment 3 with no ascoid, I ascoid on segments 4-15. : : ; ; ‘ - squamipleuris indica (p. Spermathecae smooth Spermathecae with capsules Spermathecae tubular : Spermathecal capsules about twice as Jong ; as wide 1 oO | © | ON fOr of A: ff Ge fk PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN Spermathecal capsules nearly spherical Spermathecal capsules narrow towards tip . Spermathecal capsules nearly elliptical Cibarial teeth scarcely visible . ; 4 bailyi (p. Cibarium with distinct teeth and lateral groups of denticles; piedient patch with postero-dorsal process . ‘ i ; ‘ : ; : montana (p. Cibarium with 2 teeth. : ; : ? . : : . sp. B (p. Cibarium with ro or more teeth , : Cibarium with notch in hind end of ventral plate, and 10-30 teeth in a row concave posteriorly . : Cibarium without notch in 1 hind ‘end of ventral plate er about 4 ee 50 nearly equal teeth in a comb-like row : F , africana asiatica (p. Cibarium with about 10-14 teeth, noth sheila : : ; . Sshorttii (p. Cibarium with about 16-30 teeth; notch deep : F P " : Cibarium with about 16—18 teeth : , : : : ; baghdadis (p. Cibarium with about 30 teeth . ‘ ; : : : ; é babu (p. Pharynx with well developed teeth . , : ; . palestinensis (p. Pharynx with transverse folds or irregular scales ; . grekovi (p. Spermathecae with transverse striations. Pharynx with almost invisible spicules Spermathecae smooth. Pharynx with numerous teeth Pharynx funnel-shaped, 1-5—2 times as long as hind width Pharynx conical, 2-5-3 times as long as hind width Posterior edge of pharynx sharply defined and deeply indented . punjabensis (p. Posterior edge of pharynx ill-defined and not deeply indented ‘ . theodori (p. Cibarium with 4 very large lateral teeth on each side of small central ones, and well- developed vertical teeth : ‘. : 4 dentata dentata (p. Cibarium usually with 5 lateral teeth, which are not as large as in dentata, vertical teeth small . : : . : : : ‘ dentata arpaklensis (p. KrEy TO MALES OF SERGENTOMYIA (except sp. B) Genital filaments widened at tips. Cibarium with internal lateral projections in front of teeth. No ascoid on antennal segment 3. Parameres with rounded or truncated ends. Aedeagus much shorter than parameres and narrowing before tip like short sword. Abdominal tergite 6 narrower and longer than 5 squamipleuris indica (p. Genital filaments not widened at tips Aedeagus thick and finger-shaped Aedeagus conical and ne, eae Parameres hooked . : : : : . punjabensis (p. Parameres with rounded ends ; ‘ : ‘ Aedeagus bluntly pointed ; , , : dentata dentata (p. Aedeagus with rounded end . : . OE arpaklensis (p. 2 7 theodori (p. Aedeagus with a sharp point Aedeagus with a blunt point Pigment patch conspicuous and very well defined. Cibarium aan about 34 ‘teeth. Large species : ; . hospitii (p. Pigment patch inconspicuous. ‘Cibariam with 30 teeth or fewer ; , Palpal formula 1, 2, 3, 4, 5. Aedeagus jeer ot tapering uniformly from the base, long . : : africana asiatica (p. Palpal formula 1, 2, 4, 3, 5. Aedeagus narrowing abruptly behind the base, very narrow apically , pawlowskyi hodgsoni (p. Ww Ro N NS CRs ines ieee 12 D. J. LEWIS 8 Cibarium with 8 or fewer widely spaced teeth . ‘ ; . Christophersi (p. 40) — Cibarium with 12 or more teeth, close together or in groups. : . ; : 9 g Cibarium with 20—22 small pointed teeth, most in groups of 2-4. Surstyles distinctly longer than parameres. Papilla formula 1-3/3, 1-2/4. : clydei (p. 42) — Cibarium with 12~14 wide teeth. li only slightly longer than parameres. Papilla formula 1/4. : : tiberiadis (p. 44) 10 Paramere with a spine-bearing process on “its lower side at the base of the neck. Genital filaments with marked transverse striations . pps hodgsoni (p. 37) — Paramere without such a process : II 11 Cibarium with a patch of teeth on each side ofa ‘central row : montana (p. 39) Cibarial teeth not differentiated thus . . ‘ ‘ ‘ : 12 12 Cibarium with well developed teeth in a curve or a straight line ; 4 13 — Cibarium with 1 or 2 rows of pointed teeth, often scarcely visible, in a posteriorly concave row : ‘ ‘ . ‘ 14 13 Cibarium with 8-12 teeth i ina straight line : : ; : palestinensis (p. 34) — Cibarium with 14-16 teeth in a posteriorly concave row ‘ . grekovi (p. 32) 14 Cibarial teeth very indistinct; hind end of cibarium very broad; centre of hind end of ventral plate straight when visible; chitinous arch ee Abdominal tergite 6 much narrower than5 . : ‘ bailyi (p. 38) — Cibarial teeth often indistinct; hind end of cibarium not very broad; centre of hind end of ventral plate concave; chitinous arch not prominent. Abdominal tergite 6 slightly narrower than5 . babu (p. 28), baghdadis (p. 30), shorttii (p. 31) TAXONOMY AND DISTRIBUTION OF THE SPECIES The references in the taxonomic citations have been selected from a large number which may be found in the papers of Sinton (1932, 1933c, d), Theodor (1948, 1958), Kirk & Lewis (1951), Quate (1964) and Theodor and Mesghali (1964), and in papers mentioned by them. Descriptions of some species have been published by Theodor (1958), and many were figured by Sinton (1932, 1933d). Some descriptions given below refer to Pakistan forms of variable species, and some others are given here because existing descriptions are old or not readily available. Some descriptions of cibaria and pharynges differ from those of Sinton because he removed them from the head, after maceration and staining. The numbers of ascoids on antennal segments, and numbers of papillae on proximal segments (Parrot, 1953) are omitted unless they are unusual. Hair sockets of females are shown in dorsal views of segment 3, and those of males in lateral views of the hind end of segments 5 and 6. Certain of the structures of minor taxonomic importance, such as the furca, are omitted from descriptions for the sake of brevity. In the sections on distribution, place names from various sources are omitted if they have been listed from earlier sources. Records from places which could not be certainly identified have been omitted, together with some records published before modern methods of identification came into use. Some localities are taken from notes deposited by Sinton in the British Museum. Many specimens were taken in a group of localities and are recorded from the principal one, as follows. Gujrat: Mangowol. Mir Muhammad: Mianwali, Sadhana, Shahzada. Peshawar: Ahmad Khel, Badbher, Bahadur, Bazid Khel, PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 13 TABLE II. Percentages of Species of Phlebotomus (Males) and Sergentomyia (Females) in Certain Areas. Italics Indicate Actual Numbers. The Proportions of the Genera and Sexes are Largely Due to Selection After Capture. Houses and Collections Houses sticky Light traps traps Area Lahore R’pindi Peshawar Keris Lahore R’pindi Peshawar Phlebotomus papatasi 84 40°1 g2'I — — — 2 alexandri — — — 0-6 _ — — nurt —- 2:0 a “= — —- — sergentt 16 51°I 79 39°0 — I — ka. burneyti — — — 9°8 — — =: keshishianti — 16 —- 1-2 — — — major a 2-7 — _ a — — sp. A — orl — — —_ — — c. longiductus == 2°2 - 49°4 _ — — argentipes —_— orl — — -— -— — Males 94 853 5363 164 — Ig 2 Total I59 IOoIg 5712 189 f I 4 Sergentomyia d. arpaklensis —- 0-4 2:2 2 o— — $°% theodort == O"4 4°5 — 0°3 12-1 punjabensis 28-9 I°5 2:2 — 0-3 0-3 orl babu 22°4 39°1 — me 2°2 — baghdadis 30°3 17:0 89-9 — I‘o 2°2 0:8 shorttit = 0-4 — —- —- — — sp. B — — 0-6 — — — —_ palestinensis — — —_— —_ — _— orl sq. indica I°3 7:0 — -- 82-9 42°7 72°5 paw. hodgsoni 1*3 2°6 — —_— bare) 0:8 bailyi — 25°1 — 0-3 10°4 orl montana —_— O°-7 —_ 2 — — christophersi 15°8 _ a -= _— _ clydet — 0-4 — — 15°4 40°7 10°4 hospitit os 5°5 — —- -— — — tiberiadis —_ _ 0-6 _- a — — Females 76 271 178 4 298 316 662 Total 309 1376 359 7 454 458 816 Shahb Khel, Sheikh Muhammadi. Rawalpindi: Bakra Mandi. The areas in Table II include the places mentioned and the following. Keris: Gol, Gwadi (or Guari), Parkuta. Lahore: Mir Muhammad. Peshawar: nearby villages. Rawalpindi: Said Pur, Taxila. The locations of certain type specimens are abbreviated as follows. British 14 D. J. LEWIS Museum (Natural History): ‘B.M.(N.H.)’’. Department of Parasitology, Hadassah—Hebrew University Medical School, Jerusalem: “‘ Jerusalem ”’. PHLEBOTOMUS Rondani Phlebotomus (Phlebotomus) papatasi (Scopoli) (Text-figs. 2-4) Bibio papatasi Scopoli, 1786, Deliciae Florae et Faunae insubricae 1:55. Ticini. Phlebotomus papatasii (Gmelin) ; Grassi, 1907; Newstead & Sinton, 1921 : 104 [surstyles] ; Sinton, 1925a: 468 [surstyles]. Phlebotomus papatasi (Scopoli); Sacca, 1950, Rc. Ist. sup. Sanita 13: 684 [early stages] ; Quate, 1964 ; Schmidt & Schmidt, 1962, 1963. The following notes on Pakistan specimens include particulars for comparison with specimens from other areas. Q. Cibarium with a few scattered minute ventral teeth and some lateral spicules; hind margin of ventral plate easily seen; chitinous arch prominent; dorsal wall with 2 bulges. Labrum 0°35 (0°30—-0°40) mm. long. Antennal segment 3 is 0°26 (0°24-0°30) mm. long, 1°04 (o-g9-1'09) length of 4+5, 0°75 (0°67-0°86) length of labrum; ascoid 0°43 (0°39-0°53) length of segment 4. Palpal segment 3 longer than 4; average ratio 10 : 10°0: 7°3 : 18-9. Wing length 2-25 (2-01-2:57) mm., width 0°61 (0°53-0°69) mm.; Re is 1°5 (1°3-1°6) times length of Re,3; Ri apex is 0-3 (0°2—-0'4) length of Re. g. Cibarium very like that of female. Labrum 0:25 (0:22-0'28) mm. long, crest 0-09 as high as length of labrum, sloping upward to summit, truncated or projecting very slightly forward, pubescent near apex. Antennal segment 3 is 0°30 (0°26-0°335) mm. long, 1-0 (o-g-1'1) times length of 4+5, 1:2 (1:0-1°3) times length of labrum; ascoid is 0:23 (0°20-0°25) length of segment 4. Palpal ratio 10: 10°4: 8-4: 19:0. Wing length 2:14 (1°78—2-40) mm., width 0°52 (0°45-0°63) mm.; Reg is 1°4 (1'2-1°5) times length of Ra,3; Ri apex is 0-30 (o-17-0°41) length of Re. Genital filament about 1°6 times pump length. Surstyles each with 2 terminal spines except in 1 fly with 2 and 3. Comparison of the above figures with those of the extensive analysis of Egyptian specimens by Schmidt and Schmidt (1963) does not indicate any marked regional variation in this species. The Pakistan figures do show rather long ascoids and wings (easily measurable structures) in the males, but very similar results were ob- tained with ro males taken in winter at Delgo and Saras in the northern Sudan : ascoid 0:25 (0:22-0:29) length of segment 4; wing length 2:20 (2:04-2:34) mm., width 0:56 (0:51-0:62)mm. The difference between Pakistan and Egyptian specimens may be due to individual variation and to seasonal or other local ecological conditions. There are 6 spines on one dististyle of a male from Kashmore, 3 along the shaft instead of 2. Specimens measured. 109 and 10 ¢ from the Rawalpindi area. Distribution. Newstead & Sinton (1921): Bannu, Dera Ismail Khan (iii, iv, v, ix, x), Idak, Tank. Sinton (1924): Kohat, Lahore, Miramshah, Nowshera, Quetta, Rawalpindi, Sinton (1927a): Jandola, Khirgi, Landi Kotal, Peshawar. Sinton (1932): scattered all over the plains of the Indo-Pakistan subcontinent, PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 15 more especially in hot dry areas ; has been found as far east as Calcutta and as far south as Madras City, but is most common in the north-west. Munir (1963): Chilas, Gilgit. B.M. (N.H.): Digri, Jhelum, Kandhkot, Kashmore, Larkana, Mirpur Khas, Shikarpur. Sinton’s notes: Chaman, Hyderabad, Jamesabad, Jamrud, Karachi, Khairpur, Pano Aqil, Tando Bago. Present survey: Abbottabad, Mir Muhammad, Said Pur, Taxila. Phlebotomus (Paraphlebotomus) alexandri Sinton (Text-fig. 5) Phlebotomus sergenti var. alexandri Sinton, 19280 : 308. In the one male taken in the present survey the cibarium is very like that of P. papatasi but the teeth are more delicate and the hind margin of the ventral plate is indefinite. The labral crest is obviously pubescent and rises rather abruptly to its rounded non-projecting summit. Antennal segment 3 is 0-154 mm. long, 0-66 length of labrum. The genital pump is 0-13 mm. long. Distribution. Sinton (19284): Waziristan. Sinton (1932): western frontier. B.M. (N.H.): Kambhar [probably Qambar], north-west frontier. Sinton’s notes: Dera Ismail Khan, Hyderabad, Kandhkot, Larkana, Shikarpur, Tank. Present survey: Parkuta. This species is widely distributed around the Mediterranean and is apparently always rare (Theodor & Mesghali, 1964), and the same is probably true of West Pakistan. Phlebotomus (Paraphlebotomus) nuri sp. n. (Text-figs. 6, 7) 2. Unknown. g. Cibarium with numerous delicate lateral spicules which merge into a group of about 10 small pointed scattered ventral denticles, and about 40 minute spicules in front of them ; chitinous arch well developed ; pigment patch absent ; dorsal wall of cibarium with posterior bulge. Pharynx armed with transverse scale-like ridges and posterior rows of minute denticles. Labrum 0-30 (0:28-0:34) mm. long, crest pubescent and non-projecting as in P. sergenti. Antennal segment 3 is 0-33 (0:31-0-37) mm. long, about 1-2 times length of 4+5, about 1-1 times length of labrum ; 2 ascoids on segments 3-15, that on 4 is about 0-25 length of segment; a papilla on 3-5. Palpal ratio 10: 10:0: 6:8: 20-2 ; segment 3 with inward bend at 0-7. Wing length 2-30 (2-08 2°52) mm., width 0-62 (0-56-0-69) mm.; R, is 1°5 (1-3-1°7) times length of Rois; Ry apex is 0-2 (0-1—0-3) length of R,. Basistyle 0-30 (0-28-0-34) mm. long; with basal process about 0-08 mm. long and 0-03 mm. wide, bearing on its distal third a thick brush of long downward-curving hairs, the most proximal ones being ventral. Dististyle 0-17 (0-16—0-19) mm. long, more than half length of basistyle and about 4 to 6 times as long as its own width ; witha thick terminal spine, another at 0-8, and a thick and a thin spine at 0-4. Paramere with flat elliptical upper surface. Aedeagus short and conical, with blunt end. Surstyle 0-35(0-32—0-39) mm. long. Genital pump about 0-17 mm. long, filaments about 1-5 times this length. Holotype 3. WeEsT PAKISTAN: Said Pur, 21.v.1959 (H. C. Barnett), in B.M. (N.H.). Paratypes: WEST PAKISTAN: Rawalpindi and Said Pur, 9 g, in B.M. (N.H.) ; Said Pur, r g, in U.S. National Museum ; r gin Jerusalem. 16 D. J. LEWIS = We — O-l MM 2, base of hair from first coxa; 3, ci- 4, labrum. FP. alexandri, g. 5, labrum. P. nuri, g. 6, cibarium; 7, 8, semidiagrammatic optical section of cibarium. Lut- Fics. 2-9. Phlebotomus papatasi, 3, 2, 2, 4, 3. barium ; terminalia. P. sergenti, 9. zomyia panamensis (Shannon) (American), 9. 9, the same for comparison. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 17 Distribution. Present survey: Rawalpindi and Said Pur. This species has a longer dististyle than that of P. caucasicus and is related to P. andrejevi Shakirsjanova (discussed by Theodor & Mesghali, 1964) from which it differs in the relation of antennal segment 3 to 4+5, the relatively short palpal segment 4, the arrangement, length and curvature of hairs on the process of the basistyle, and the long narrow dististyle with a slightly different arrangement of spines. Specimens examined. 11 ¢ from Rawalpindi and Said Pur (10 measured). This species is named in honour of Lieut. Col. Nur Ahmad. Phlebotomus (Paraphlebotomus) sergenti Parrot (Text-fig. 8) Phlebotomus sergenti Parrot, 1917, Bull. Soc. Path. exot. 10: 564. The cibarium of both sexes is very like that of P. papatasi but the teeth are smaller and less scattered. The labral crest of the male is like that of P. alexandri. Distribution. Newstead & Sinton (1921): Dera Ismail Khan. Sinton (1922) : Lahore. Sinton (1924b): rare and localized; very localized at Lahore; Quetta. Sinton (1927a): Chitral, Landi Kotal. Sinton (1929): Sukkur. Sinton (1932) : in the Indo-Pakistan subcontinent it is apparently confined to the plains north- west of a line between Bombay and Simla, and it and P. papatasi are found under similar conditions. Sinton’s notes: Cherat, Jhelum, Shikarpur, Tank. Present survey: Chilas, Gilgit, Gol, Gwadi, Keris, Mir Muhammad, Parkuta, Peshawar, Rawalpindi, Said Pur, Taxila. Phlebotomus (Larroussius) kandelakii burneyi subsp. n. (Text-figs. 10-14) Phlebotomus kandelakiit Shurenkova ; Barnett & Suyemoto, 1961: 616; Nasir, 1964: 26. 9. The petioles of many of the scales are distinctive owing to their transparency in contrast to the rest of the scale. Cibarium without denticles, distinct chitinous arch, or a visible hind margin of the ventral plate; dorsal wall with 2 folds. Pharyngeal armature occupying the hind 0-28 of the pharynx, anterior teeth scale-like and bearing spicules, the rest are small denticles. Labrum 0-28 (0:27-0:29) mm. long. Antennal segment 3 is 0-265 (0:24-0:30) mm. long, I-2 (1-:2—1-3) times length of 4+5, same length (o0-8—1-0) as labrum ; 2 ascoids on segments 3-15, that on 4 being 0-35 (0-3—0-4) length of segment ; 1 papilla on segments 3-5. Palpal ratio about 10:8:7:21. Wing length 2-35 (2:29-2:46) mm., width 0-68 (0:63-0:72) mm.; R, is 2-0 (1-8—-2-3) times length of R,,;; R, apex is 0-3 (0-3-0°4) length of R,. Spermatheca narrow at ends and with long neck and about 30-35 segments which merge gradually into ill-defined rings on the duct ; duct wide and thin-walled at its hind end ; posterior edges of furca meeting at an acute angle. 6. Scale petioles and cibarium as in female. Pharynx with smaller armature. Labrum 0-22 (0:20-0:24) mm. long; transparent part of crest short and low, with an ill-defined curved summit and a few distal spicules. Antennal segment 3 is 0-31 (0-26—0-33) mm. long, 1-1 times length of 4+ 5, 1°4 (1-3-1°5) times length of labrum ; ascoids paired on segments 3-7, single on ENTOM. 19, I. 2 18 D. J. LEWIS 8-15, that on 4 being 0-2 (0-2-0-3) length of segment. Palpal ratio 10: 8-5:8:21. Wing length 2-02 (1-81-2-07) mm., width 0-53 (0-45-0°57) mm. ; RR, is 1-8 (1-3—2-1) times length of Ry3; Ry, apex is 0-3 (0-2-0-4) length of R,. Basistyle about 0-31 mm. long, with about 20, not very long, ventral non-deciduous hairs. Dististyle about 0-16 mm. long, about half length of basistyle, with 2 terminal spines, 1 spine at 0-5, and 2 ato-75. Paramere light brown, with a short sub-basal ventral process bearing about 7 spines, a narrow neck and, on the distal half, a grey nearly smooth ventral flange ; depth of distal part about 0-32 of distance from its tip to tip of process (about 0-22 in P. k. kandelakii). Aedeagus long, with a blunt transparent tip and a row of very fine ventral teeth which are mainly on the basal half. Genital pump about 0-15 mm. long, filaments about 3-6 times as long. Surstyle about 0-34 mm. long. Holotype ¢g. West PAKISTAN: Gwadi, 25.vi.1959 (M. I. Burney), in B.M. (N.H.). Paratypes: WeEsT PAKISTAN: Gwadi, Kalam, Keris and Parkuta, 69, 13 3 in B.M. (N.H.): Keris, 1 9, 3 g, in U. S. National Museum, Washington. Distribution. Present survey: Gwadi, Kalam (2,070 metres), Keris (house in evening), Parkuta. ait 0:05 MM Fics. 10-14. Philebotomus kandelakii burneyi, 11, 12, 9, 10, 13, 14, ¢. 10, base of hair from first coxa ; 11, cibarium; 12, spermatheca ; 13, labrum: 14, terminalia. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 19 This form differs from P. k. kandelaki in the deeper distal part of the paramere. Further study is needed to show if slight differences in the pharyngeal armature, the ascoid distribution in the male, and the position of teeth on the aedeagus are significant. On the average, the labrum of P. k. burneyi is relatively short, palpal segment 5 is long, R, is relatively long in the female, and non-deciduous hairs of the style are more numerous than in P. k. kandelakw. The tip of the aedeagus resembles that of a male taken by Shurenkova at Tiflis in 1930. Specimens measured. 69 and 10 ¢ from Gwadi and Keris. This form is named in honour of Lieut. Col. M. I. Burney. Phlebotomus (Larroussius) keshishiani Shurenkova (Text-figs. 15-19) Phlebotomus keshishiani Shurenkova, 1936, Med. Parasit. 5: 922 ; Theodor, 1958: 24; Theodor & Mesghali, 1964 : 291. g. A very large pale sand-fly. Cibarium with delicate lateral spicules which merge into a small ventral group of minute spicules ; chitinous arch absent ; ventral plate without definite hind margin ; 2 dorsal bulges present. Pharyngeal armature composed of anterior scales and posterior punctiform teeth. Labrum 0-38 mm. long. Antennal segment 3 is 0-34 mm. long, 1:3 times length of 4+-5, o-9 length of labrum ; ascoid 0-4 length of segment 4. Palpal ratio 10:8:7. Wing length 2-90 mm., width 0-85 mm.; R, is 1-7 times length of R,,,; Ry, apex is o-2 length of R,. Spermatheca with about 18 segments, conical, narrowing towards duct ; end process relatively short, about 3 times as long as wide. 6. Cibarium very like that of female. Pharynx armed with punctiform teeth and a few scales in front of them. Labrum 0-34 (0-30—0-38) mm. long, crest with rounded non-projecting summit, terminal spicules, and a small basal bulge. Antennal segment 3 is 0-47 (0:40-0°57) mm. long, 1°3 (1-I-1°5) times length of 4+-5, 1-4 (1:3-1-6) times length of labrum ; ascoids paired on 3-8, single on 9, that on 4 being 0-3 (0-3-0-4) length of the segment. Palpal ratio about 10:9:7:21. Wing with rather straight front margin, length 2-89 (2:42-3:29) mm., width 0°82 (0:70-0:97) mm.; R, is 1:8 (1-4—2-2) times length of R,,,;; R, apex is 0-2 (0-2—0-4) length of R,. Basistyle about 0-44 mm. long, with about 30 non-deciduous hairs. Dististyle about 0-22 mm. long, with 2 spines terminal, 2 at 0-7, and one ato-5. Aedeagus conical with extremely narrow rounded tip. Genital pump about 0-16 mm. long, filaments about 8 times as long. Paramere with scattered ventral hairs beyond basal bulge, distal part slightly deeper than neck and grey along its lower surface. Surstyle about 0-43 mm. long. Professor O. Theodor has identified this form by comparing it with Russian specimens. The above description tallies fairly well with his, but the Pakistan speci- mens are relatively large and have a different ascoid distribution, and the relative lengths of the palpal segments may indicate some variation. P. keshishiani differs from P. smirnovi Perfil’ev (1941) in having long spermathecal ducts and genital filaments, a large area of pharyngeal teeth, and other features. In Theodor’s (1958) key to the males of Palaearctic Phlebotomus, P. keshishiani would run to couplet 16, differing from the other species indicated in the combination of blunt aedeagus, without lateral teeth, and long genital filaments. Specimens examined, 12 and 20 g (10 ¢ measured). Distribution. Present survey: Gilgit, Parkuta, Rawalpindi, Said Pur. 20 Fics. 15-19. Phlebotomus keshishiani, 15, 16, 9, 17-19, 4. 15, cibarium ; mathecae; 17, labrum; 18, wing; I9, terminalia. 16, sperm- PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 21 Phlebotomus (Larroussius) major major Annandale (Text-fig. 20) Phlebotomus major Annandale, 1910, Rec. Ind. Mus. 4: 46; Quate, 1962b; Theodor, 1958 ; Theodor & Mesghali, 1964: 281. 6. No females were seen. Some particulars of males are as follows (10 from Said Pur measured). Cibarium with scarcely visible lateral spicules, 2 dorsal bulges, and no visible chitinous arch or hind margin to ventral plate. Labrum 0-28 (0-25—0-31) mm. long; crest with small basal bulge, slightly rounded non-projecting summit, and terminal spicules. Antennal segment 3 is 0-40 (0-35—0-48) mm. long, 1-2 (1-I-1°3) times length of 4+5, 1-5 (1-4-1°5) times length of labrum; ascoids paired on segments 3-8, single on 9, that on 3 being about 0-3 (o-3-0-4) length of segment. Palpal ratio about 10:9:7:18; segment 3 is 1-2 (I-o—1-3) times length of 2. Wing length 2-44 (2:23-2:81) mm., width 0-70 (0-63—0-86) mm. ; R, is 1-9 (1°5—2:2) times length of R,,,; R, apex is 0-2 (0-2—0°3) length of R,. Basistyle about 0-38 mm. long. Dististyle about 0-19 mm. long. Genital pump about 0-15 mm. long, filaments some 4:3 times this length. Surstyle about 0-38 mm. long. The relative lengths of palpal segments 2 and 3 suggest that the West Pakistan form has some affinity to P. m. syriacus Adler and Theodor, but it is treated as P.m. major in view of the degree of variation. P.m. major and P. chinensis vary in colour (Sinton, 19285). Type. <¢ lectotype, Naini Tal, India; in Zoological Survey of India (Quate, 19620). Distribution. Sinton (1932): it seems to be essentially a hill species in the Indo-Pakistan subcontinent where it occurs in areas some I,500 to 2,100 metres up where there is marked rainfall in summer, apparently existing all along the Himalayan foothills. Present survey: Abbottabad, Rawalpindi, Said Pur. Phlebotomus (Larroussius) sp. A (Text-figs. 21, 22) Q. Cibarium with rows of lateral spicules but no ventral ones, chitinous arch absent, ventral plate without definite hind margin, 2 dorsal bulges present. Pharynx very like that of P. m. major. Labrum o-43mm. long. Antennae, palps and wings missing. Spermatheca narrow at each end, with about 21 segments and a rather long thick-walled neck which is very narrow where it joins the terminal segment ; ducts very long and narrow. The one available specimen was taken in a house collection at Said Pur on 21.v.1959 (H. C. Barnett) together with a male of P. m. major, but it does not seem to belong to that species. Phlebotomus (Adlerius) chinensis longiductus Parrot (Text-figs. 23-33) Phlebotomus chinensis ; Sinton, 19286 : 306, [in part] ; Adler & Theodor, 1929, [in part]. Phlebotomus major var. longiductus Parrot, 1928, Archs Inst. Pasteur Algér. 6: 29. Phlebotomus chinensis var. longiductus ; Theodor, 1948 : 107 ; Theodor & Mesghali, 1964 : 293. Phlebotomus chinensis hindustanicus Theodor, 1958 : 29, syn. n. Phlebotomus chinensis longiductus ; Theodor, 1958 : 29. 22 D. J. LEWIS O'2 MM Q:1MM__, — 1 O:2 MM 24 0°OS MM = 1 , 4 Fics. 20-35. Phlebotomus major, 6. 20, aedeagus. P. sp. A, 9. 21, abdominal sternite 2; 22, spermatheca. P. chinensis longiductus, 23, 9, 24-33, ¢. 23, sperm- matheca; 24, labrum; 25-28, antennal segments 3-6; 29, palp; 30-33, aedeagus and tip from various angles. P. argentipes, g. 34, 35, cibarium. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 23 Q. Petioles of many scales pale, as in P. k. burneyi, but this feature is less marked owing to the pallor of the shafts of the scales. The insect is very pale. Cibarium with scarcely visible lateral internal spicules, a short internal flange on each side, and 2 bulges in the dorsal wall ; without a chitinous arch or a visible hind border to the ventral plate. Pharyngeal teeth occupy- ing a third of the length of the pharynx, anterior ones forming a median group, posterior ones very broad and scale-like. Labrum 0-37 (0:32-0:39) mm. long. Antennal segment 3 is 0-32 (0-27—-0°35) mm. long, 1-3 (1-2-1-4) times length of 4+ 5, 0-9 (0-8-1-o) length of labrum ; ascoid on 4 is 0-4 (0-3—-0°4) length of segment. Average palpal ratio 10: 7-4: 6-2: 16-9. Wing length 2°54 (2:23-2°72) mm., width 0-74 (0:65-0:80) mm.; R, is 1-8 (1-4-2-0) times length of R,,;; R, apex is 0-2 (0:2—0°3) length of R,. Spermatheca narrow at each end, particularly toward the “head ’’; with ill-defined compartments of different sizes, some of them packed together like cells ; ducts long and wrinkled, each with a short posterior thick-walled section which joins the other duct at the outlet. ¢. Cibarium much like that of female. Pharynx with smaller armature. Labrum 0:27 (0:24-0-30) mm. long with very small basal bulge and a shallow crest with rounded summit and terminal spicules. Antennal segment 3 is 0-34 (0-28—0-39) mm. long, 1-2 (1-1-1-3) times length of 4+5, 1-3 (1-1-1-4) times length of labrum; ascoids paired on segments 3-5 and single on 6, except in one fly with an ascoid and a vestige of one on 6; ascoid on 4 is 0-3 (0-2-0-3) length of segment. Average palpal ratio is 10: 8-0:7:0: 19:2. Wing length 2-27 (2-01-2°54) mm., width 0-63 (0:54-0:77) mm. ; R, is 1-8 (1-6—2-0) times length of R,,,; R, apex is 0-2 (o-1-0°3) length of R,. Basistyle about 0-38 mm. long, with a not very dense patch of about 60 non-deciduous hairs. Dististyle about 0-20 mm. long, 2 of spines terminal, one at 05, and 2 at abouto-6. Paramere very like that of P. major. Each plate of aedeagus tapering to a rounded tip, with a lateral subapical tooth and a transparent subterminal flange extending downward and slightly inward. In lateral view the tooth is scarcely visible, but slight changes in orientation can make it conspicuous and sometimes appear to be dorsal (in some descriptions of P. chinensis it is stated to be ventral) ; this tooth is 0-016 (0-015—0-018) mm. from the tip, and 0-015 (0-014—0:020) in 10 flies from Said Pur. Genital pump 0-13 (0:12-0:15) mm. long. filaments about 8-9 times this length. Surstyle about 0-41 mm. long. Theodor (1958) pointed out that the form Aindustanicus differed from P. c. longiductus in the position of the subterminal tooth of the aedeagus and that, when more specimens were available from the area between Turkestan and north- west India, the two forms might possibly be found to be the same. P. chinensis from West Pakistan proves to be intermediate in this respect, and form hindustanicus is therefore treated here as a synonym of P. c. longiductus. In Pakistan specimens antennal segment 3 is shorter than in those from India described by Theodor, and the ascoid distribution differs from that in Turkestan and India. Specimens measured. 6 9 from Gol, Gwadi, Keris and Parkuta ; 10 ¢ from Gol. Distribution. Sinton (1932) : similar, in the Indo-Pakistan subcontinent, to that of P. major. Mitra (1959) and Jacob and Kalra (1951) : Punch (Kashmir). Nasir (1964) : Gilgit. Present survey: Gol, Gwadi, Keris, Parkuta, Said Pur. Phlebotomus (Euphlebotomus) argentipes Annandale & Brunetti (Text-figs. 34, 35) Phlebotomus argentipes Annandale & Brunetti, 1908, Rec. Indian Mus. 2: 101; Sinton, 1925¢ ; Christophers, Shortt & Barraud, 1926; Theodor, 1948: 108; Keilin & Tate, 1937, Para- 24 D. J. LEWIS sitology 29: 254 [larva]; Quate, 1962a@:254; 1962b; Quate & Fairchild, 1961, Pacif. Insects 3: 211. Phlebotomus argentipes var. glaucus Mitra & Roy, 1953), syn. n. 6. The following are some particulars of West Pakistan specimens. Cibarium with a few small punctiform teeth, the anterior ones in a row; chitinous arch not visible; hind margin of ventral plate scarcely visible ; 2 variable dorsal bulges present ; sides very dark. Labrum 0:20 (o-19-0'21) mm. long, with small basal bulge; crest with low rounded non-projecting pubescent summit. Antennal segment 3 is 0-24 (0:22-0:25) mm. long, I-1I (1-1I-1-2) times length of 4+-5, 1:2 (1-I-1'3) times length of labrum ; ascoids paired on segments 3-10, single on 11-15, that on 4 being 0-4 (0-4-0-5) length of segment; papilla on 3 and 4. Average palpal ratio 10: 9:0: 4-8: 13:8. Most of mesonotum and scutellum dark brown, pleura pale. Wing length 1-99 (1:98-2:02) mm., width 0-59 (0-55—0-62) mm.; WR, is 1-8 (1-6—2-1) times length of Roi3; R, apex iso-1lengthofR,. Basistyle about 0-27 mm. long. Dististyle about 0-18 mm. long. Genital pump dark brown, 0-15 mm. long, filament about 2-3 times this length. P.a. var. glaucus appears, from its description, to be a minor variant of this species. Specimens measured. 6 J. Type. 3 lectotype, Calcutta, India ; Zoological Survey of India (Quate, 19620). Distribution. Nasir (1964): Lahore. Present survey ( ¢): Mir Muhammad (10.iv.63, W. A. McDonald, i in house), Shahzada (May, 1963, W. A. M., 3 on cattle), Taxila (29.v.59, H. C. Barnett, r in house). Phlebotomus (Anaphlebotomus) colabaensis Young & Chalam (Text-figs. 36, 37) Phlebotomus colabaensis Young & Chalam, 1927, Indian J. med. Res. 14: 859; Sinton, 1932 : 59; 1933¢: 226; 1933d: 418; Theodor, 1958: 108. 2. The following are some features of the one available specimen. Cibarium with no teeth in middle line, but an irregular row of about 5 spiculated teeth with broad delicate bases on each side; small spicules in front of these teeth; chitinous arch brown and well defined ; ventral plate with visible hind margin; dorsal wall with 2 bulges. Labrum o-26 mm. long. Antennal segment 3 is 0-28 mm. long, 1-1 times length of 4+5, 1-1 times length of labrum ; ascoids paired on segments 3-15, that on 4 being 0°55 length of segment ; papilla on segments 3-5. Palpal ratio 10:9:5:... Thorax pale grey. Wing length 1-96mm., width 0-56 mm. ; R, is 1-2 times length of R,,,; R, apex is 0-4 lengthof R,. Spermatheca with about 20 narrow segments which merge imperceptibly into wrinkles of the duct, terminal segment small and head oblong; ducts about Io times length of spermathecae, uniting in short thick-walled common duct. This may be a new species but is provisionally classed as P. colabaensis in the absence of a male. Sinton stated that the spermathecal ducts were four times as long as the spermathecae but the variation may be due to different methods of mounting. Distribution. Sinton (1932): India (the small peninsula of Colaba at Bombay, and Vishakhapatnam). Present survey: Lahore, sticky trap in Davipur orchard, 16.v.1963 (D. J. Lewis). PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 25 SERGENTOMYIA Franca & Parrot Sergentomyia (Sergentomyia) dentata dentata (Sinton) Phlebotomus dentatus Sinton, 1933a: 869; 1933¢ : 227; 1933d: 421. Sergentomyia dentata (Sinton) ; Theodor, 1958 [in part] ; Theodor & Mesghali, 1964 : 293. 9. Cibarium with 4 very large pointed teeth on each side and 6 small central ones, and a patch of small vertical teeth on each side ; chitinous arch present. Pharynx about 1-4 times as long as hind width, its armature comprising numerous long slender anterior teeth and a narrow group of small posterior teeth. Antennae missing. Palpal formula apparently 1—2-3-4-5 ; ratio 10: 11:9:16. Wing length 1-68 mm., width 0-31 mm.; R, 0-6 length of R,,,; R, apex about 0-4 of R,. Spermathecae tubular with wide ducts. 6. Cibarium with about 15 large pointed teeth in a row which is markedly concave back- wards. Pharynx with several rows of pointed teeth, about 6 in each anterior row and about 3 in each posterior one. Antennae and wings missing. Dististyle with 4 apical spines, and seta at 0-8. Aedeagus finger-shaped and almost straight, with bluntly pointed end ; parameres blunt ; genital pump not visible. This description is based on Sinton’s. More specimens are needed for study. LECTOTYPE 9°. West PAKISTAN: “ Quetta co-type 9 1”’, in B.M. (N.H.), by present designation. Paralectotypes: 9 from Quetta labelled “‘co-type’’, 3g from Quetta labelled “ type’, in B.M. (N.H.), by present designation. Distribution. Sinton (1933a), Quetta. Sergentomyia (Sergentomyia) dentata arpaklensis (Perfil’ev) (Text-figs. 38, 39) Phlebotomus minutus ; Sinton, 1932 : 61 [in part]; 1933d: 42 [in part]. Phlebotomus minutus var. arpaklensis Perfil’ev, 1933, Zool. Anz. 101: 226; Adler, 1946, Bull. ent. Res. 36 : 506. Sergentomyia dentata var. mediensis Pringle, 1953, Bull. ent. Res. 43: 714. Synonymy after Theodor & Mesghali, 1964. Sergentomyia dentata : Theodor, 1958 [in part] ; Lewis, Mesghali & Djanbakhsh, 1961 [in part]. Sergentomyia dentata arpaklensis (Perfil’ev) ; Theodor & Mesghali, 1964. The labral crest of the male is truncated, and in the male of this form or S. theodort abdominal tergite 6 is large. Variation in S. d. arpaklensis is mentioned in the section on S. theodom. A related form in Sinkiang has been described as P. minutus var. sinkiangensis by Ting & Ho (1962). Types. 12 syntypes, from the Kara-Kala area, Turkmeniya; in the Russian Academy of Military Medicine. Distribution. Present survey: Gwadi, Landi Kotal, Peshawar, Rawalpindi. 26 D. J. LEWIS Fics. 36-49. Phlebotomus colabaensis, 2. 36, cibarium ; 37, spermatheca. Sergentomyia dentata or theodori, g. 38, labrum; 39, abdomen. S. punjabensis, 40-43, 2, 44-48, 3. 40, head; 41, cibarium; 42, pharynx; 43, spermatheca; 44, cibarium; 45, pharynx ; 46, labrum; 47, abdomen; 48, terminalia. S. africana africana (Ghana), ¢. 49, labrum. ‘ PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 75, Sergentomyia (Sergentomyia) theodori (Parrot) Phlebotomus minutus : Adler & Theodor, 1926, Bull. ent. Res. 16 : 403 ; Sinton, 1932 : 61, 73 [in part] ; 1933d@: 421, 422 [in part]. Phlebotomus theodori Parrot, 1942, Archs Inst. Pasteur Algér. 20 : 332. Sergentomyia theodorit (Parrot); Theodor, 1958; Lewis, Mesghali & Djanbakhsh, 1961 ; Theodor & Mesghali, 1964. Some features of 10 9 from the Peshawar area are as follows. Labrum 0-15 (o:r4-0-16) mm. long. Antennal segment 3 is 0-II (0-10—0-11) mm. long, shorter than 4-++5, 0-7 (0-7—-0-9) length of labrum; ascoid about 0-4 length of segment 4. Palpal formula 1-—2-4-3-5 or I-2-(3-4)-5; ratio about 10:12:11; Newstead spines on 3 conspicuous. Wing length 1-50 (1-43-1°57) mm., width 0-30 (0:27-0:32) mm. ; R, is 0-6 (0-4-0°8) length of R,,,; R, apex is —0-2 to +0-3 length of Rg. The character given by Theodor and Mesghali for separating females of arpaklensis and theodori, length of pharynx divided by its hind width, is very variable in Pakistan specimens, as it is in S. pumjabensis. I am treating them here as species, however, because the specimens examined were few and from a limited area. The numbers of each form were estimated by regarding as theodori all females in which the above-mentioned fraction was 2:25 or less. The labral crest of the male is truncated. Distribution. B. M. (N.H.): Dera Ismail Khan, Kashmore, Larkana. Nasir (1958) : Peshawar. Present survey: Landi Kotal, Rawalpindi. Sergentomyia (Sergentomyia) punjabensis (Sinton) (Text-figs. 40-48) Phlebotomus minutus var. antennatus ; Sinton, 1932: 61, 73. Phlebotomus antennatus ; Sinton, 1933d : 421 ; Qutubuddin, 1952: 79. Phlebotomus punjabensis Sinton, 1933d: 421. Sergentomyia punjabensis (Sinton) ; Theodor, 1948 : 109 ; Qutubuddin, 1951: 36. Phlebotomus antennatus var. deccanensis Qutubuddin, 1952: 79, syn .n. Q. Cibarium with about 30 nearly uniform teeth on a posteriorly concave line, the outer ones visibly pointed ; about 10 small punctiform teeth present; pigment patch broad, short and very dark; chitinous arch absent; lateral walls of narrow part of cibarium dark, with irregular inward surfaces. Pharynx 1-3-1-8 times as long as hind width, with sharply defined deeply notched posterior outline. Labrum 0-14 (0-13-0-15) mm. long. Antennal segment 3 is 0:09 (0-08—o-09) mm. long, shorter than 4+ 5, 0-6 (0-6-0-7) length of labrum ; ascoid about 0-4 length of segment 4. Palpal formula 1-2-(3-4)—5 or 1-2-3-4-5; ratio about 10: 12: 13. Wing length 1-40 (1:21-1:50) mm., width 0-31 (0:28-0:33) mm.; R, 0-6 (0-4-0-9) length of Roei3; Ry, apex o-2 (o-1-0-4) length of R,. Spermathecae tubular with delicate ducts. $. Cibarium with about 20 nearly equal pointed teeth, which appear blunt in their normal position, and a few small punctiform teeth ; pigment patch variable, usually short, broad and rather pale; chitinous arch absent. Pharynx with faint scaly sculpturing. Labrum 0-13 (o-12—0-14) mm. long, with some spicules on its basal bulge, and a pubescent truncated crest. Antennal segment 3 is 0-10 (0-0g—o-11) mm. long, shorter than 4+5, 0-8 (o-7—0-8) length of labrum ; ascoid about 0-3 length of segment 4. Wing length 1-35 (1:28-1:47) mm., width 0-27 (0:25-0:29) mm. Abdominal tergite 6 very large. Dististyle with 4 apical spines, and seta at 28 D. J. LEWIS about 0-7. Aedeagus thick and curved, with rounded end; genital filaments about 3-5 times pump length ; paramere with beak-like apex. Specimens examined. Many from Dera Ismail Khan, Lahore, Peshawar and Rawalpindi areas, 10 2 and 10 ¢ measured. Form deccanensis appears to be a synonym, because the pharynx of the female is like that of West Pakistan specimens, the greater length of antennal segment 3 is largely due to differences in body size, and the relative length of the dististyle is within the range of variation seen in West Pakistan. Distribution. Qutubuddin (1951): Kohat area. B.M.(N.H.): Dera Ismail Khan, Jhelum, Khanki, Lahore, Peshawar. Present survey: Mangowol, Said Pur Shahzada. Sergentomyia (Parrotomyia) africana asiatica (Theodor) Phlebotomus africanus ; Sinton, 1932 : 61, 71 [Sind] ; 1933d: 422. Phlebotomus africanus var. asiaticus Theodor, 1933, Bull. ent. Res. 24: 541 [Israel]; 1952, Istanb. Univ. Fen Fak. Mecm. (B) 17 : 116 [relation to Israel form unknown]. Sergentomyia africana var. asiatica (Theodor), 1948: 110 [Israel, N.-W. India]; 1958: 43. Sergentomyia africana asiatica (Theodor, 1958). Specimens presented to the British Museum (Nat. Hist.) by Sinton probably came from West Pakistan and accord with Theodor’s descriptions. He differentiated S. a. asiatica from the African form now known as S. a. magna by the absence in aszatica of punctiform cibarial teeth and the presence of more (45-50) horizontal teeth and few, long, pharyngeal teeth. Sinton’s drawings show 38 and 48 horizontal teeth, respectively, in the two forms. Examination of S. a. magna from several areas shows that many specimens have about 50 horizontal teeth and no definite vertical ones, but S. a. asiatica can still be distinguished by its pharyngeal teeth. The labrum of the male in S. a. africana Newstead (Text-fig. 49) and S. a. asiatica has a tapering crest. Types. Types or syntypes in Jerusalem. Distribution. Sinton’s notes: Rhedia (near Larkana), Kandhkot, Shikarpur, all between 1930 and 1932. Sergentomyia (Parrotomyia) babu (Annandale) (Text-figs. 50-59) Phlebotomus babu Annandale, 1910, Rec. Indian Mus. 4: 49; Sinton, 1932 : 60 ; 1933d: 422; Theodor, 1938: 261; Quate, 1962b: 157. Phlebotomus minutus var. niger ; Sinton, 1927, Indian J. med. Res. 15: 31. 9. Cibarium with about 30 nearly equal pointed teeth, the outer 12 or 13 on each side nearly parallel to each other and pointing slightly inward ; ventral plate often dark grey posteriorly, with a deep median notch of variable shape ; on each side of this notch the margin of the ventral plate is divided into a lower surface, with one or more protrusions, and an upper surface, PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 29 Z 0:05 MM : q O:| MM = Fics. 50-64. Sergentomyia babu, 50-54, 9, 55-59, 6. 50, head; 51, cibarium; 52, pharynx ; 53, spermatheca ; 54, spermathecae in end view ; 55, cibarium ; 56, pharynx ; 57, abdomen ; 58, terminalia; 59, genital filament. S. baghdadis, 60-62, 9, 63, 64, 6. 60, cibarium ; 61, pharynx; 62, spermathecae; 63, cibarium; 64, pharynx. 30 D. J. LEWIS above the roots of the teeth ; chitinous arch ill-defined ; pigment patch somewhat obscured by ventral plate, broad with its hind edge in line with the tips of the cibarial teeth. Pharynx narrowing to a varying extent behind the main bulge, teeth pointed. Labrum 0-16 (0-14-0-17) mm. long. Antennal segment 3 is 0-18 (0-17—0-21) mm. long, 1-0 (o-9-1-0) times length of 4+5, 0:8-o-9 length of labrum ; segments 4+5 are 0-18 (0-16—0-20) mm. long; ascoids about o-3 length of segment 4. Palpal formula 1-—2—3~4-5, ratio about 10:12:13. Wing length 1-64 (1:49-1°87) mm., width 0-37 (0:31-0:45) mm. ; R, is 0-7 (0-4-0-9) length of R,,,; R, apex is 0-3 (o-1-0-5) of R,. Spermatheca elliptical, greatest width about 3 times diameter at base of collar; thick-walled, junction with ducts partly flattened so that in some views the duct appears as a thin-walled extension of the spermathecal capsule ; gland ducts and collar delicate, without distinct knob ; spermathecal ducts short and thin-walled. g. Cibarial teeth often scarcely visible ; about 10 in the centre are pointed, and about 5 on each side are often ill-defined ; hind end of ventral plate markedly concave, the apex of the curve sometimes acute but seldom indented ; chitinous arch ill-defined; pigment patch faint or absent. Pharynx with faint transverse ridges. Labrum o-15 (0-14-0-16) mm. long, crest tapering as in S. africana. Antennal segment 3 is 0-16 (0-14—0-17) mm. long, shorter than 4+5, almost same length (0-9-1-1) as labrum ; ascoid about o-2 length of segment 4. Wing length 1°33 (1°30-1°39) mm., width 0-27 (0-24-0-32)mm. Dististyle with 2 apical and 2 subapical spines, and seta at about 0-65. Aedeagus mainly dark brown, curving slightly upward and tapering to a rounded colourless tip; genital filaments with conspicuous transverse ridges, about 2:2 times pump length. Parameres with truncated ends and lower corners extending downwards. Specimens examined. Many from West Pakistan; 1092 from Rawalpindi area and ro ¢ from Lahore measured. Sinton recorded a dark variant from Pusa, India, and Mitra and Roy (1952) described, as P. thapari, a form from Poona which may be a variant of S. babu. Types. Lectotype ¢. Calcutta, India; Zoological Survey of India (Quate, 19620). Distribution. Sinton (1932) : S. babu seems to have a wide distribution over the plains and foothills of India and Pakistan. Sinton (1933d): occurs between the areas of S. baghdadis and S. shorttti. Sinton’s notes: Cherat, Lahore. Present survey : Gilgit, Landi Kotal, Mir Muhammad, Rawalpindi, Said Pur, Taxila. Sergentomyia (Parrotomyia) baghdadis (Adler & Theodor) (Text-figs. 60-64) Phlebotomus baghdadis Adler & Theodor, 1929 : 281 ; Sinton, 1932 : 60; 1932d: 422. 9. Cibarium with about 18 pointed teeth, comprising about 4 small ones in the centre and about 7 large ones on each side; ventral plate with a deep median notch of varying shape, chitinous arch ill-defined ; pigment patch broad, usually hidden by ventral plate. Pharynx with faint transverse ridges, and a few spicules at its hind end. Labrum o-15 (o-14-0-17) mm. long. Antennal segment 3 is 0-16 (0:15-0:17) mm. long, same length (1-0-1°1) as 4+ 5, I-O-I°I times length of labrum; segments 4+5 are 0-16 (0:15-0:17) mm. long; ascoids about 0-3 length of segment 4. Palpal formula 1-—2-3-4-5, ratio about 10:12:13. Wing length 1-62 (I°51I-1'72) mm., width 0-35 (0:32—0°38) mm. ; R, is 0-6 (o-3-0°8) length of R,,,; R, apex is 0-3 (—o-r to 0-5) of R,. Spermatheca (like that of S. babu) with partly flattened outlet to duct ; gland ducts and collar very delicate and lacking a distinct knob; spermathecal ducts short. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 31 g. Cibarial teeth usually scarcely visible, sometimes comprising about 6 central pointed teeth and a number of lateral spicules ; hind end of ventral plate markedly concave ; chitinous arch ill-defined; pigment patch faint or absent. Pharynx with a few transverse ridges. Labrum 0-15 (0:14-0:16) mm. long, with tapering crest. Antennal segment 3 is 0-17 (o-14-— 0-19) mm. long, slightly shorter than 4+ 5, 1-1 (1-o—1-2) times length of labrum ; ascoid about 0-2 length of segment 4. Wing length 1-51 (1-39—1-64) mm., width 0-32 (0-28—0-36) mm. Abdominal tergite 6 nearly as wide as 5. Dististyle with 2 apical and 2 subapical spines, and seta at about 0-6. Aedeagus mainly dark brown, curving slightly upward and tapering to a colourless rounded tip; genital filaments about 2-5 times pump length. Parameres with truncated ends, lower corner directed downward. Specimens examined. Many from West Pakistan, 10 2 and 10 ¢ from Peshawar area measured. Sinton (1932) distinguished females of S. baghdadis from those of S. babu by the numbers of teeth given in the key, and considered that they and S. shorttii were closely related, but regarded them as species rather than varieties on account of their morphology and distribution. He (1933d) found the males difficult to distin- guish without taking their distribution into account. Differences in the cibarial notch and the relative lengths of antennal segments 3 and 4+5 of certain in- dividuals of S. babu and S. baghdadis (Adler & Theodor, 1929) do not hold for all specimens. Types. 129 and 12 9 syntypes, from Baghdad and Basra; in Jerusalem ; 131 2 and r g were examined (O. Theodor, personal communication). Distribution. Sinton (1932) : in the west and north-west of the Indo—Pakistan subcontinent. B.M.(N.H.): Dera Ismail Khan, Jhelum, Kandhkot, Lahore, Rohri, Shikarpur, Sukkur. Sinton’s notes: Bannu, Gujranwala, Jandola, Kashmore, Larkana, Lyallpur, Pano Agqil, Peshawar, Sarghoda, Tank. Present survey: Landi Kotal, Mir Muhammad, Rawalpindi, Said Pur, Taxila. Sergentomyia (Parrotomyia) shorttii (Adler & Theodor) (Text-figs. 65-67) Phlebotomus shorttii Adler & Theodor, 1927 : 65 ; Sinton, 1928b : 317; 1932: 60; 1933d: 422. 9. The one available specimen is damaged and was found far from the known area of S. shorttii, therefore the record requires confirmation. Some features are as follows. Pharynx robust, with few longitudinal folds, slightly pigmented in anterior 3/4, most of teeth short and pointed, some in groups. Labrum o-17mm. long. Antennae missing. Palpal formula I-2-3-4-5 ; ratio 10:10:12. Wing length 1-57 mm., width 0-37 mm.; R, is 0-6 length of R.,3; Ry, apex is 0-4 of R,. Spermatheca elliptical, rigid part of capsule about the same width at each end. Types. 8 2 and 8 ¢ syntypes from Golaghat, India; in Jerusalem. Distribution. Sinton (1932) : north-eastern India, and Burma. Sinton (19334) : eastern frontier of India, andin Burma. Qutubuddin (1944) : Hyderabad (Deccan, India). Present survey: Taxila, 29.v.1959 (H. C. Barnett), in house, I 9. 32 D. J. LEWIS Fias. 65-70. Sergentomyia shoritii, 9. 65, cibarium; 66, pharynx; 67, spermatheca. S. sp. B, 9. 68, cibarium; 69, pharynx; 70, spermatheca. Sergentomyia (Parrotomyia) sp. B (Text-figs. 68-71) 2. The single specimen lacks appendages. The cibarium has two teeth, the pharyngeal armature comprises ridges and some minute posterior spicules, and the spermathecae are like those of S. babu. This may be a new species related to S. shorttii but differing from it in the number of cibarial teeth and the nature of the pharyngeal armature. Distribution. Present survey: Bahadur (near Peshawar), 12.vi.1959 (H. C. Barnett). Sergentomyia (Parrotomyia) grekovi (Khodukin) (Text-figs. 71-78) Phiebotomus grekovt Khodukin, 1929, Medits. Mysl’ Uzbekist. Turkmenist. Suppl. : 101. Phlebotomus graecovi Perfil’ev, 1937 : 127. Sergentomyia grekovi (Khodukin) ; Theodor, 1958. Q. Cibarium with knob-like postero-lateral margin and 18-24 parallel pointed teeth on an arc which is slightly concave posteriorly ; a few punctiform teeth on each side; chitinous arch well defined at sides, postero-lateral edges of cibarium knob-like ; pigment patch with an irregularly truncated forward extension and an irregular hind margin. Pharynx with thick walls and well marked posterior scales. Labrum o-17-0-18mm. long. Antennal segment 3 is PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 33 0:05 MM pe O:|_ MM Pare 0-05 MM, Fics. 71-79. Sergentomyia grekovi, 71-74, 2, 75-78, ¢d. 71. cibarium; 72, pharynx ; 73, wing; 74, spermatheca. 75, cibarium; 76, pharynx; 77, abdomen; 78, term- inalia. S. palestinensis, 9. 79, cibarium. ENTOM. IQ, I. 3 34 D. J. LEWIS 0:20-0:22 mm. long, shorter than 4+5, 1:2 times length of labrum; ascoids 0-3 length of segment 4. Palpal formula 1-2-3-4-5; ratio 10:11:11. Wings long and narrow, length 1°84-1:96 mm., width 0-36—0-38 mm. ; R, short, 0-3-0°5 length of R,,,; R, apex is o or —o-7 in relation to R,. Spermatheca nearly spherical, with short collar, and wrinkles on inner surface of wall. ¢é. Cibarium with about 12 parallel, pointed teeth on an arc slightly concave posteriorly, and punctiform teeth at their bases ; chitinous arch well defined, postero-lateral edges of cibar- ium knob-like ; pigment patch vestigial. Pharynx with scales and transverse ridges. Labrum o-17 mm. long, crest truncated and pubescent as in S. d. arpaklensis. Antennal segment 3 is o:21 mm. long, shorter than 4+ 5, 1-2 times length labrum; ascoid 0-2 length of segment 4. Wing length 1-86 mm., width 0-33 mm. Dististyle with 2 apical and 2 subapical spines, and seta ato-7. Aedeagus slightly upturned in distal half, with a blunt point, brown colouring gradually reduced from base to tip; filaments short, 2-5 times pump length. Parameres with beak-like down-turned ends. Specimens examined. 292 and 1 ¢ from Gilgit. Theodor notes that variation in the number of cibarial teeth may indicate the existence of different forms, and it is therefore interesting to find considerable variation in the two Pakistan females. Distribution. Present survey: Gilgit, rr.vi.1963, sticky trap. Sergentomyia (Parrotomyia) palestinensis (Adler & Theodor) (Text-fig. 79) Phlebotomus sp. Adler & Theodor, 1926: 404. Phlebotomus palestinensis Adler & Theodor, 1927: 64; Theodor, 1947, Bull. ent. Res. 38: 96; Lewis, 1957, Ann. Mag. nat. Hist. 10 : 691 ; Perfil’ev, 1960. Sergentomyia palestinensis (Adler & Theodor) ; Pringle, 1953, Bull. ent. Res. 43 : 719 ; Theodor & Mesghali, 1964 : 295. Q. In the one damaged female available there are 16 teeth, each with a nodular thickening near its centre; the chitinous arch and postero-lateral edges of the cibarium are prominent. The labrum is broad, as shown in the original figure, and 0-14 mm. long. Wing length 1-46 mm., width 0-32 mm.; R, is 0-4 length of R,,,;; R, apex is —o-2 in relation to Rg. g. Inamale from Iran the labral crest is tapering. The number and form of the cibarial teeth of the female support the conclusion of Theodor and Mesghali that this species may be very variable. Type. Holotype 9, from Jericho, Israel; in Jerusalem. Distribution. Present survey: Peshawar, 25.vi.59 (H. C. Barnett), light trap, 1 9. Sergentomyia (Grassomyia) squamipleuris indica (Theodor) (Text-figs. 80-83) Phlebotomus squamipleuris var. indicus Theodor, 1931, Bull. ent. Res. 22 : 470. Phlebotomus squamipleuris ; Sinton, 1923, Indian. J. med. Res. 11:65; 1932:60; Perfil’ev, 1939 [in part]; Quate, 1962a@ : 259. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 35 Sergentomyia squamipleuris var. indica (Theodor) ; Theodor, 1948 : 112. Sergentomyia squamipleuris ssp. indica (Theodor) ; Theodor, 1958 : 47 [in part]. Sergentomyia indica (Theodor) ; Theodor & Mesghali, 1964 : 295. 9. Cibarium with about 34-48 teeth in a posteriorly convex row, a line of punctiform teeth at their bases; chitinous arch weakly developed ; wall of cibarium with inward projections behind teeth ; pigment patch broad and dark. Pharynx brown, with convex outline behind the main bulge; many teeth present on the dorsal plate, partly hidden by spine-like teeth on ventral plates. Labrum 0-15 (0-14—0-17) mm. long. Antennal segment 3 is 0-14 (0-13—0-15) mm. long, shorter than 4+ 5, 0-9-1'1 length of labrum ; ascoids single on segments 3-15, about 0-3 or 0-4 length of segment 4. Palpal formula 1-2-3-4-5 ; ratio about 10:11:13; Newstead scales on 2 (difficult to see) and 3. Femora without a row of spines. Wing length 1-65 (1°52-1°74) mm., width 0-38 (0-35—0-42) mm. ; R, is 0-9 (0-7-1-1) length of R,,;; R, apex is 0°3 (o:I-0'5) of R,. Scales or their bases are to be seen on the pleura. Abdominal tergites dark brown; 3 with about 2-5 erect hairs on each side ; in 3 out of Io flies 2 hairs form the rudiment of a vertical line. Spermathecae with a compact mass of minute gland-ducts and a membrane, delicate or thick, surrounding their bases ; surface of spermatheca smooth near its apex, otherwise marked with brown or colourless, small or very small, spicules, and often with some longitudinal lines ; individual ducts of spermathecae sometimes about 4 times length of spermathecae but often contracted. é. Cibarium with about 22 teeth on a line slightly convex posteriorly ; chitinous arch not very definite ; lateral walls of cibarium with inward projections; pigment patch present. Pharynx with distinct but delicate teeth. Labrum 0-13 (0-13-0-14) mm. long, with wide trans- parent side pieces (as seen in lateral view), basal bulge pubescent, summit of crest rounded and projecting slightly forward. Antennal segment 3 is 0-15 (0-14—0-16) mm. long, shorter than 4+5, (1:I-1'2) times length of labrum ; ascoid about 0-3 length of segment 4. Wing length 1-49 (1-42-1-61) mm., width 0-31 (0:29-0:34) mm. Abdominal tergites dark brown, tergite 6 slightly longer than 5 and curved dorsally. Dististyle with 2 apical and 2 subapical spines, and seta at about 0-75. Aedeagus curved upward and gradually narrowing except at the tip where it narrows abruptly to a rounded end. Genital filaments about 4 times pump length, with expanded ends. Parameres with rounded or truncated ends ; the flattened inward surface of each faces slightly downward, and the curvature of the outward surface gives a beak-like appearance at certain angles of view. Specimens examined. Many from West Pakistan ; ro 9 and 10 ¢ from Peshawar area measured. The tip of the paramere in this complex has been variously described as “rounded ”’, a “‘ bluntly-rounded point ’’, a “truncate ... blunt beak-like tip ’’, and “‘ hooked ’’, to mention only a few terms ; the discrepancies are probably due to a slightly variable structure being viewed from different angles. S.s. indica differs from S. s. sguamipleuris (Newstead) in the shape of the pharynx and from S. s. dreyfussi (Parrot) in the absence of femoral spines, but is treated here as a subspecies because the differences are small. Quate (1964) states that the spermathecal ducts of S. s. squamipleuris are short, but they are very like those of zmdica and any apparent difference may be due to contraction or the effects of different mounting media. The variety poonaensis, first described and then named by Mitra & Roy (19534, 1954) was not seen in Pakistan. Specimens examined. Many from West Pakistan, 102 and Io ¢ measured. Types. Syntypes. InpiaA: Karnal, 19, I g, 24.vii.1928; Saharanpur, 36 D. J. LEWIS 83 SANS ---( WW 2.0 © o @ =p aay 93 Oo 7 0:05 MM L al Fics. 80-94. Sergentomyia squamipleuris indica, 80, 81, 9, 82, 83, g. 80, pharynx; 81, spermatheca; 82, labrum; 83, abdomen. S. pawlowskyi hodgsoni, 84-89, 9, go-94, ¢. 84, 85, cibarium and semidiagrammatic optical section; 86, denticles of another fly ; 87, pharynx; 88, hair scars on abdominal segment 3; 89, spermatheca ; go, cibarium; 91, labrum; 92, abdomen; 93, 94, some hairs or scars on abdominal segments 5 and 6. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 37 32,3 3, 29.vii.1929. WEST PAKISTAN: Jhelum,1 4, 24.iv.1930. All collected by Sinton, now in Jerusalem. (O. Theodor, personal communication). Distribution. Sinton (1927a): Lahore, Peshawar. Sinton (1932): it has a very wide distribution all over the Indo-Pakistan subcontinent, both on the plains and in the foothills up to 1,800 metres; a “ garden species’’. B.M.(N.H.): Dera Ismail Khan, Jhelum, Khanki, Tank. Sinton’s notes: Cherat. Present survey : Gujrat, Rawalpindi, Said Pur, Saidu Sharif (960 metres), Taxila. Sergentomyia (Rondanomyia) pawlowskyi hodgsoni (Sinton) stat. nov. (Text-figs. 84-94) Phlebotomus hodgsoni Sinton, 19330: 874; 1933d: 419. Sergentomyia hodgsoni (Sinton) ; Theodor, 1948 ; Theodor & Mesghali, 1964 : 296. 9. Cibarium with about 40-60 long contiguous teeth in a nearly straight row, with a line of punctiform teeth at their bases; in front of these are about 4 additional punctiform teeth (o-15 in 21 cibaria examined) ; chitinous arch scarcely visible, postero-lateral parts of cibarium directed almost straight backward ; each side of cibarium with large inward extension in front of chitinous arch; pigment patch broad and reddish brown with a conspicuous pale forward extension and a sharply defined hind edge formed by a posterior bulge in the dorsal wall of the cibarium. Pharynx with stout walls, a few transverse ridges, and some fine posterior spicules. Labrum o-18 (0:17-0:19) mm. long. Antennal segment 3 is 0-14 (0:12-0:15) mm. long, shorter than 4+5, 0°8 (0-7-0°8) length of labrum; ascoid about 0-45 length of segment 4. Palpal formula 1-2-3-4-5 ; ratio1I0: 14:10:24. Bodyvery pale. Wing length 1-78 (1-63—1-93) mm., width 0-42 (0:39-0:46) mm. ; R, is 0-7 (0-4—1-0) length of R,,,; R, apex is 0-4 (0-2-0'5) length of R,. Abdominal tergites 2-6 with several large sockets of vertical hairs near hind margins. Spermatheca tubular with a distinct knob, no collar, and numerous fine transverse striations ; each duct broad and similarly marked, joining a broad, thin-walled, wrinkled common duct. é. Cibarium with about 32 teeth in a nearly straight row, and a few denticles ; chitinous arch almost invisible ; pigment patch distinct but pale, with a hyaline bulge behind it. Pharynx with strong walls and posterior transverse lines. Labrum 0-16 (0:15-0:16) mm. long; crest with long dorsal thickening, and scarcely projecting rounded summit. Antennal segment 3 is 0°15 (0°14—0'15) mm. long, shorter than 4+ 5, same (o-9-1-1) length as labrum ; ascoid about 0-3 length of segment 4. Wing length 1-64 (1:49-1:71) mm., width 0-39 (0°35-0-41) mm. Abdominal tergite 6 much longer than 5, with no hairs except large microtrichia. Dististyle with 2 of spines subapical, and seta on proximal half, at about 0-4. Aedeagus tapering to a blunt end ; filaments about 3-6 times pump length. Paramere with beaked end and about 8 short spines on a small distinct ventral lobe. The above description is based on that of Sinton, supplemented by notes by Theodor & Mesghali and by myself. Sinton drew attention to the abdominal hairs of this species and thought it might belong to Sintonius. It hasa striking resemblance to some members of this subgenus through the combination of cibarial characters, shape and texture of pharynx, body colour, abdominal hairs of female and of segment 6 of male, large abdominal tergite 6 of male and beaked parameres. The reduction or loss of medium-size hairs on tergite 6 of male sand-flies seems to be associated with increasing size of the tergite (and possibly a concentration of fat- body) which is a feature of most species of Sintonius. 38 D. J. LEWIS S. p. hodgsoni differs from S. p. pawlowskyi in having broad individual and common spermathecal ducts and, on the average, fewer denticles in front of the main row of cibarial denticles. The anterior denticles are difficult to count if the pigment patch is dark, as in Said Pur flies, and are not always in definite rows. LECTOTYPE 9. WEsT PAKISTAN: labelled “ Cherat, r.vii.32, P. 263/B”’, in B.M. (N.H.), by present designation. Paralectotype 3. Locality as for lectotype, vi. 32, P. 267/B, in B.M. (N.H.), by present designation. Distribution. Sinton (19330): Cherat, Jandola, Landi Kotal. Present survey : Gwadi, Parkuta, Peshawar, Rawalpindi, Said Pur, Taxila. Sergentomyia bailyi (Sinton) (Group nicnic) (Text-figs. 95-103) Phlebotomus bailyi Sinton, 1931b : 822; Quate, 1962a@: 260, 262. Phlebotomus bailyi var. campester Sinton, 1931b : 823 ; Theodor, 1938: 268. Synonymy after Quate, 19624 : 262. Sergentomyia bailyi (Sinton) ; Theodor, 1948. 9. Cibarium with scarcely visible teeth arranged in a posteriorly concave row and small lateral groups ; hind end expanded laterally ; pigment patch small or absent ; chitinous arch well developed. Pharynx with rows of minute spicules. Labrum is 0-16 (0-14-0-17) mm. long. Antennal segment 3 is 0-20 (0-18—o-22) mm. long 1-2-1-3 times length of labrum, equalling or longer than 4+5; ascoids about o-5 length of segment 4. Palpal formula 1—2—3-4-5; ratio 10:10:11:24. Pleura very pale. Wing length 1-90 (1:63-2:20) mm., width 0-46 (0-40— 0:50) mm.; R, about o-go length of R,,,; R, apex o-3-0'5 length of R,. Spermathecae in the form of smooth capsules narrowing toward the apices, with very delicate ducts. $. Cibarium with scarcely visible pointed teeth in a posteriorly concave row; hind end expanded laterally ; chitinous arch well developed ; pigment patch small or absent. Pharynx with minute spicules. Labrum 0-15 mm. long, dorsal part shallow at centre, crest rising to a rounded summit and tapering distally. Antennal segment 3 is 0:25 (0:23-0:28) mm. long, 1-4 (1-3-1-4) times as long as labrum (10 from Taxila measured for this ratio), about equal to 4+5; ascoid about o-2 length of segment 4. Pleura very pale. Wing length 1-93 (1-79- 2°14) mm., width 0-39 (0:35-0:44) mm. Abdominal tergite 6 distinctly narrower than 5. Disti- style with 2 apical and 2 subapical spines, seta at 0-7, nearly level with subapical spines. Basistyle with many non-deciduous hairs on inner face. Aedeagus with colourless, bluntly- pointed, slightly up-turned tip; genital filaments 4 times pump length ; paramere with beak- like apex. This description is based on Sinton’s and, apart from measurements, on examina- tion of many specimens from West Pakistan. The length of antennal segment 3, given in Sinton’s (1933d) key to males, does not serve to separate all S. bailyi from S. babu. This species and S. montana do not belong to any of the existing subgenera but it seems inadvisable to create new ones until related species (Theodor, 1948) are better known. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 39 Distribution. Sinton (1931b): Dera Ismail Khan, Jandola, Lahore, Larkana area, Pano Aqil, Tando Muhammad Khan; very widely distributed in the Indo- Pakistan subcontinent, from sea level to 1,830 metres, much rarer than S. babu and usually forming not more than three per cent of Sergentomyza collections from the plains ; common in hills where it is found with P. major, P. chinensis and S. montana. Qutubuddin (1951) : Kohat-Hangu valley. Sinton’s notes: Peshawar, Shikarpur, Tank. Present survey: Mir Muhammad, Rawalpindi, Taxila. O-1MM Fics. 95-103. Sergentomyia bailyi, 95-98, 9, 99-103, o. 95, head, 96, cibarium ; 97, pharynx; 98, spermatheca; 99, cibarium; 100, pharynx; to1, labrum; tIo02, abdomen ; 103, terminalia. Sergentomyia montana (Sinton) (Grouping uncertain) (Text-figs. 104—11T) Phlebotomus minutus var. montanus Sinton, 1924a: 809; 1927¢: 26. Phlebotomus montanus Sinton, 1927, Indian J]. med. Res. 15:30; 1929:174; 1932: 61; 1933d : 422. 40 D. J. LEWIS Q. Cibarium with about 22 teeth in posterior row, the central 12 knob-like, small and con- tiguous, the outer 5 on each side widely spaced, pointed and inclined towards centre ; in front of lateral teeth is a group of denticles on each side; chitinuous arch well developed, with a brown strip behind it on each side; pigment patch with posterior upward projection, which looks like a dark brown refractive area, and a long wide anterior projection. Pharynx with a posterior network of fine ridges and a few spicules. Labrum o-18 (0-15—0-19) mm. long. Antennal segment 3 is 0:23 (0-22—0-25) mm. long, longer than 4+ 5, 1-3 times length of labrum ; ascoids about o-3 length of segment 4. Palpal formula 1-2—3-4-5; ratio 10:11:12. Wing length 2-10 (1:18-2:29) mm., width 0-49 (0:36-0'54) mm.; RR, long, 1-1 (o-9-1-3) times length of Ry,3; R, apex is 0-6 (0-5-0-8) length of R,. Spermatheca a capsule with thick nearly- straight smooth walls, narrower at apex ; knob small and collar well marked. ¢. Cibarium with about 14 teeth in posterior row and a group of about 4 denticles on each side; chitinous arch well-defined ; pigment patch with postero-dorsal projection, usually appearing as a refractive area, and a long thin anterior process. Pharynx with a faint posterior network of fine lines. Head pale, so that the antennal sockets look like white patches. Labrum 0-17 (0:16-0:18) mm. long, crest truncated and pubescent, very like that of S. theodori. Antennal segment 3 is 0-30 (0:29—0-32) mm. long, slightly longer than 4+ 5, 1-8 (1-6—1-9) times length of labrum; ascoid about 0-15 length of segment 4. Wing length 1-98 (1:84-2:11) mm., width 0:39 (0-35-0'44) mm. Abdominal tergite 6 slightly smaller than 5. Dististyle with 2 apical spines, 1 subapical dorsal, and 1 inward-pointing spine behind it; and seta at about 0-4. Aedeagus tapering to a rounded point; filaments about 4-3 times length of pump which is narrow anteriorly. Paramere with beak-like apex and markedly down-turned point. Specimens examined. 2 2 from Bahrein, 3 2 from Parkuta, and 1 (small) 2 from Said Pur; 10 ¢ from Bahrein. Distribution. Sinton (1924a): Murree. Sinton (1927a): Khaira Gali. Sinton (1932) : occurs in the foothills of the western Himalayas at about 1,830 metres. B.M. (N.H.): Chitral road. Present survey: Bahrein (1,390 metres), Gilgit, Parkuta, Rawalpindi, Said Pur, Taxila. Sergentomyia (Sintonius) christophersi (Sinton) (Text-figs. 112-115) Phlebotomus christophersi Sinton, 1927¢ : 22, 24 ; 1927d : 33 ; 1927, Indian J. med. Res. 15: 30. 9. Cibarium with 4 or 5 widely spaced pointed teeth; chitinous arch pale, with lateral flanges ; sides of cibarium are nearly straight and converge towards the hind end, postero- lateral corners divergent; pigment patch small and carrot-shaped. Pharynx not sharply contracted behind bulge, with a posterior network of fine lines. Labrum long, length 0-24 (0:22-0:25) mm. (in 5 flies). Antennal segment 3 is 0:13 (o-11-0-14) mm. long, slightly shorter than 4+5, 0-6 length of labrum (in 3 flies) ; ascoids about 0-4 length of segment 4. Palpal formula 1-2-4-3-5; ratio 10:14:9:21. Wing length 1-66 (1-57-1-73) mm., width o-40 (0-35-0°43) mm.; R, is 0-6 (o-5—0-8) length of R,,,; AR, apex is 0-4 (0-3—0-5) length of Rg. Abdomen pale, with some erect hairs on hind margins of tergites. Spermatheca small, with 6—9 segments, no collar, a distinct knob, and a very long narrow duct with transverse thickenings. 6. Cibarium with 2 or 3 teeth and several denticles ; chitinous arch pale; postero-lateral corners of cibarium divergent; pigment patch carrot-shaped. Pharynx with transverse lines. Labrum 0-17 (0-16—0-18) mm. long (in 5 from Lahore), summit of crest projecting upward and forward asin S. clydei. Antennal segment 3 is 0-15 (0-14—0-16) mm. long, shorter than 4+ 5, 0-8 (o-7—1-0) length of labrum (in 5 from Lahore) ; ascoid about 0-3 length of segment 4. Wing length 1°55 (1:50-1-63) mm., width 0-37 (0:36-0:37) mm. Abdominal tergites 2-5 with some PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 41 O-l| MM 0:5 MM ee Fics. 104-115. Sergentomyia montana, 104-107, 2, 108-111, g. 104, cibarium; 105, pharynx; 106, hair scars on tergite 3; 107, spermatheca; 108, adbomen; 109, hairs and scars on tergite 6; 110, terminalia; 111, genital pump and filament. S. christophersi, 112-114, 9, 115, g. 112, cibarium; 113, pharynx; 114, spermatheca; 115, abdomen. 42 D. J. LEWIS suberect hairs on hind margins, tergite 6 very large. Dististyle with 2 of spines subapical, and seta at about 0-6. Aedeagus tapering to a point, filaments about 3:8 times pump length. Paramere beaked. The above description is based on Sinton’s supplemented by my observations. Distribution. Sinton (1927c) : Lahore (about three per cent of sand-flies caught in jail in August and September; one elsewhere in May). Theodor & Mesghali (1964): Iran. Sinton’s notes: Jhelum. Sergentomyia (Sintonius) clydei (Sinton) . (Text-figs. 116-124) Phlebotomus clydei Sinton, 1928a:179; Perfil’ev, 1941 [pigment-patch variation]. Sergentomyia clydei (Sinton); Theodor, 1958; Lewis & Minter, 1960; Lewis & McMillan, 1961 ; Theodor & Mesghali, 1964; 297. Sergentomyia clydei latiterga Theodor, 1958. Synonymy after Theodor & Mesghali, 1964. Q. Cibarium with about 12 pointed teeth on a line which is slightly convex posteriorly at the centre ; with a zigzag line of denticles ; chitinous arch faint in the centre but with prominent lateral flanges ; postero-lateral corners of cibarium not prominent ; pigment patch broad and well defined. Pharynx narrowing considerably behind the bulge, with a few posterior lines and some rows of minute spicules. Labrum 0-22 (0:20-0:25) mm. long. Antennal segment 3 is 0-15 (0:14—-0'17) mm. long, 0-9 (0-9—1-0) length of 4+5, 0-7 length of labrum ; ascoid about 0-6 length of segment 4 ; segment 3 with 1-4 papillae, segment 4 with 1 ; palpal formula 1-2—4-—3-5 ; ratio 10:12:10. Wing length 1°75 (1:60-1:93) mm., width 0-43 (0:39-0-48) mm.; R, is 0-7 (o-5-o-9) length of R,,,; R, apex is 0-3 (0-2-0-4) length of R,. Abdomen pale, with suberect hairs on hind margins of tergites 2-6. Spermatheca with about 9 segments, and long narrow duct and common duct (0-65 and o-12 mm. long respectively in a fly with wing 1-77 mm. long). $. Cibarium with about 16-26 pointed teeth arranged in groups of 2—4, and an irregular row of denticles ; chitinous arch faint; pigment patch about o-4 width of cibarium. Pharynx with fine transverse lines. Labrum o-1g (0:17-0-23) mm. long, summit of crest projecting forward and upward. Antennal segment 3 is 0-16 (0:15-0:18) mm. long, shorter than 4+5, 0-8 (o-8—1-0) length of labrum ; ascoid about 0-45 length of segment 4; segment 3 with 1-3 papillae. Wing length 1-65 (1-55-1-74) mm., width 0-38 (0-35-0-41) mm. A few large hairs on abdominal tergites 2-5, tergite 6 with large microtrichia, wider than 5. Dististyle with 2 of spines subapical, and seta at about 0-7. Aedeagus short and tapering to a point, filaments about 4°5 times pump length. Paramere beaked. The above description is based on 10 2 and 10 ¢ from Rawalpindi. In Africa many of the males have a large sixth abdominal tergite. In ten males from Rawalpindi the length of this tergite divided by that of the fifth ranged from 1-0 to 1-2 and averaged I-05. LECTOTYPE 3g. West Pakistan: Jandola, labelled “type 3’’, in B.M. (N.H.), by present designation. Distribution. Sinton (1928a): Jandola, Khirgi. Sinton (1932): recorded widely from the plains of the Indo-Pakistan subcontinent. Perfil’ev (1941) : Central Asia. Pringle (1956): Iraq. Mesghali (1963): Iran. Theodor & Mesghali PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 43 (1964): Iran. Sinton’s notes: Peshawar, Tando Muhammad Khan, Kairpur, Kandhkot. Present survey: Karachi, Lahore, Mir Muhammad, Rawalpindi, Taxila. ’ 2 °o 7 ab aes aD A rf tv : 123 uy ee aig 124 ay eS ® © ate’ owe : = = zs Ds aed) @ : Fics. 116-124. Sergentomyia clydei, 116-120, 9, 121-124, 6. 116, cibarium; 117, pharynx; 118, labrum; 119, hair scars on abdominal tergite 3; 120, spermatheca ; 121, labrum ; 122, abdomen; 123, 124, hairs or scars on abdominal segments 5 and 6. Sergentomyia (Sintonius) hospitii (Sinton) (Text-figs. 125-130) Phlebotomus simillimus var. hospitit Sinton, 1924¢ : 261; 1927¢:22; 1927d: 30. Phlebotomus hospitit Sinton, 1929: 174; 1932: 60; 1933d: 420. Sergentomyia hospitii (Sinton) ; Theodor, 1948. A large species with long antennal segment 3 and, in the 4, a large tergite 6. 9. Cibarium with about 70-80 long contiguous teeth on a posteriorly convex arc ; chitinous arch well developed, with a brown area in front of it and 2 lateral blackish areas behind it ; pigment patch very broad, mainly black except for the broad brown anterior extension. 44 D. J. LEWIS Pharynx markedly restricted beyond bulge, with thick walls, a few transverse lines, and minute posterior spicules. Labrum 0-20 (0:18-0:21) mm. long. Antennal segment 3 is 0-28 (0-24—0-31) mm. long, I-I (1-1-1-2) times length of 4+ 5, 1-4 (1-2—-1°5) times length of labrum ; ascoid about o-3 length of segment 4. Palpal formula 1-2-3-4-5; ratio 10:11:13. Wing length 2-18 (1:95-2:40) mm., width 0:56 (0-48—0-63) mm.; RF, is 1-2 (o-9—1-4) length of R,,,; R, apex is 06 (o°5—0-7) length of R,. Femur 1 with about 6-8 short stout spines and 1 or more short thin ones. Abdominal tergites 2-5 with posterior patches of large suberect hairs. Spermatheca with about 9 segments and a long narrow duct. 6. Cibarium with about 30 teeth on a posteriorly convex arc; chitinous arch well de- veloped at sides, with a brown area in front of it and 2 lateral blackish areas behind it ; pigment patch broad and blackish, without forward extension. Pharynx very like that of female. Labrum 0-20 (0-17—0-22) mm. long, crest well developed with non-projecting summit. Antennal segment 3 is 0-34 (0:31—0-38) mm. long, longer than 4+ 5, 1-7 (1-6—1-8) times length of labrum ; ascoid about o-2 length of segment 4. Wing length 2-30 (1:96-2:54) mm., width 0-59 (0-48— 0:70)mm. Femur 1 with about 8-11 short spines of varying thickness, femur 3 with several long spines. Abdominal tergite 2 with a few large suberect hairs, tergite 6 very large, with microtrichia and no hairs. Dististyle with 2 of spines subapical, and seta at about 0-6. Aedeagus tapering to a point, filaments about 4:2 times pump length. Paramere beaked and rather dark. The above descriptions are based on ro 9 and Io ¢. The femoral spines are difficult to count because some are rather like hairs and usually several are missing. LECTOTYPE 3g. Kasumir: Dulai, 21.ix.23, presented by Sinton in 1950 and labelled “type g of Kr 2” (Kr 2 referring in his notes to locality and date), in B.M. (N.H.), by present designation. Paralectotype 2. Locality and date as for lectotype, labelled “‘ co-type 2 Kr 2”’, B.M. (N.H.), by present designation. Distribution. Sinton (1924c): Dulai. Sinton (1932): recorded only from the western foothills of the Himalayas. Present survey: Abbottabad area, Chilas, Rawalpindi, Said Pur. Sergentomyia (Sintonius) tiberiadis Adler, Theodor & Lourie (Text-figs. 131-133) Phlebotomus sp. near clydei Adler & Theodor, 1929 : 284. Phlebotomus tiberiadis Adler, Theodor & Lourie, 1930, Bull. ent. Res. 21:537; Parrot & Clastrier, 1960. Phlebotomus subtilis Parrot & Martin, 1939, Archs Inst. Pasteur Algéy. 17: 151. Synonymy after Theodor, 1953, J. Washington Acad. Sci. 43: 121. Sergentomyia tiberiadis ; Theodor, 1958; Theodor & Mesghali, 1964 : 297. 9. Cibarium with 12 teeth, their points directed upward and often hidden; 1 or 2 rows of large denticles present, and often small denticles in front of them, which may be in a row or in central and lateral groups; chitinous arch indistinct; postero-lateral corners of cibarium prominent ; pigment patch distinct, nearly as broad as armature, with long forward extension and posterior bulge ; sclerites at bases of stipites conspicuous. Pharynx thin-walled, narrowing gradually behind bulge, almost smooth posteriorly. Labrum o-1g (0:17-0:20) mm. long. Antennal segment 3 is 0-17 (0:16-0:18) mm. long, o-9 length of 4+5, 0-9 (o-8-o-9) length of PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN Fics. 125-136. Sergentomyia hospitit, 125-128, 9, 129, 130, ¢. 125, cibarium; 126, pharynx ; 127, femur1; 128, spermatheca; 129, labrum; 130,abdomen. S. tiberiadis, Q. 131, cibarium; 132, pharynx; 133, spermathecae. Nematode parasites. 134, 135, from P. papatasi ; 136, from S. clvdet. 46 D. J. LEWIS labrum ; ascoid about 0-4 length of segment 4. Palpal formula 1-2—4-3-5 ; ratio 10:14: 9. Wing length 1-50 (1:36-1:61) mm., width 0-32 (0:30-0:33) mm.; BR, is 0:5 (o-1-0-7) length of Ry,3; Ry, apex is —o-7 (—2-o0to1-o) in relation to R,. Abdominal tergites 2-6 with a few large suberect hairs. Spermatheca small, narrowing towards apex, with about 9 faintly discernible segments, the knob ending in a refractive tip from which the gland ducts extend ; duct long and narrow, leading to short common duct. g. One male from Rawalpindi is probably this species. Its labral crest is like that of S. clydet. Specimens examined. 4 9 (palps of 2 and wings of 3). The Pakistan form differs from that described by Theodor in the shortness of R,, the shape of the spermathecae, and the union of their ducts. The spermathecal characters are difficult to evaluate from the examination of a few preserved females from one area, and I am therefore not naming the Pakistan form as a species or subspecies at present. Types. 22and4 dg syntypes, Tiberias, Israel ; in Jerusalem. Distribution. Present survey: Ahmad Khel, Landi Kotal, Peshawar. ZOOGEOGRAPHY Faunal subregions represented in West Pakistan. In Wallace’s zoogeographical map of the world (Bartholomew ¢é al., 1911) West Pakistan includes parts of the Mediterranean and Siberian subregions of the Palaearctic region, and part of the Indian subregion of the Oriental region. The Ethiopian region must also be con- sidered here because, although far away, it has close relations with the Oriental (Gressitt, 1958). These two Palaearctic subregions include the Eremian zone of Uvarov (1938) which extends from the western Sahara to Central Asia and has sometimes been treated as a distinct region. It has a particular significance for eremic sand-flies, for which it may serve as a link between regions instead of a barrier. The Turkestan desert is part of the great Palaearctic desert zone and some groups in its fauna include immigrants from Africa (Heptner, 1938). Kryzhanowsky regards Middle Asia as the most important centre of origin of the desert fauna of the northern hemisphere (Uvarov, 1965). Faunal boundaries used in and near West Pakistan vary according to the animals studied ; Wallace includes Peshawar in the Mediterranean subregion, Gilgit in the Siberian, and Karachi and Lahore in the Indian; and Maxwell-Lefroy (rg09) adopts a rather similar system. His ecological zones, with approximate heights in metres, are: tropical (0-600), subtropical (600—1,800) and temperate (with Palae- arctic fauna) (above 1,800). The Himalayas, according to Darlington (1957), are only a partial zoogeographical barrier and less important than climatic differences. The western Himalayas have brought the Palaearctic boundary southward (Mani, 1962) ; it has been placed at 2,000 and 3,000 metres but varies from place to place and is partly determined by latitude and the intrusion of southern forms along river valleys. The West Pakistan Himalayas lie well within the western Himalayan botanical province (Stearn, 1960). HLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 47 Th West Pakistan fauna in general. Reference to a few zoogeographical works will indicate the general nature of the fauna which is a blend of the Palaearctic and Oriental faunas (Munroe, 1965). Darlington (1957) points out that the fauna is impoverished and that the Himalayas permit some Oriental species to spread northward. The north-western Himalayas show a marked dominance of Palaearctic and endemic animals (Mani, 1962), and endemic Hemiptera have been discussed by Hutchinson (1934). The Hemiptera of north-west Pakistan have substantial Palaearctic affinities but include Oriental elements (Ghauri, 1965). Some 70 per cent of the West Pakistan Trichoptera are Oriental and some species are widely distributed, but this order is poorly represented, perhaps because some main centres of dispersal are far away (Schmid, 1958). Christophers (1933) found that most typically Oriental anopheline mosquitoes tended to become scarce before reaching their western limits but that some alpine forms, which occur from about g00 to 2,400 metres up, extended to the Hindu Kush in Chitral and perhaps beyond. Qutubuddin (19604, b) noted a mixture of Palaearctic and Ethiopian mosquitoes in the north and north-west of West Pakistan, and Lundblad (1934) found a poor hydracarine fauna in the western Himalayas. The phlebotomine fauna. The forms known in West Pakistan are those listed in Table II and P. colabaensis, S. d. dentata, S. a. asiatica and S. grekovi. They com- prise some 29 species and one local subspecies, and ten or more other species may be expected to occur. Thus the number of forms is considerable, owing to the above- mentioned desert link, the mingling of regional faunas, and the fact that many sand-flies flourish in dry conditions. The distribution of many of the species in the Palaearctic region has been discussed by Theodor (1952, 1964). Some Pakistan species are too widely distributed in the world to be described as elements of a particular region or subregion, and some forms, although confined to one subregion, are closely related to species of another sub- region. Therefore, in discussing the Pakistan species according to their general distribution and taxonomic relationships, it is only possible to group them very loosely. Species with a largely Mediterranean distribution are P. papatasi, P. alexandri, P. sergenti, P. major, S. dentata and S. theodort, and they reflect the importance of the genus Phlebotomus in that area. S. papatasi, P. sergenti, S. d. arpaklensis and S. theodort are widely distributed in West Pakistan but P. papatasi was noticeably absent in the Keris area. P. major seems to be associated with the northern hills. S. africana, S. christophersi, S. clydei and S. tiberiadis are among the species with Ethiopian affinities. S. christophersi seems to have a very sporadic distribution and one of its few known localities is in the Palaearctic (Iran). Of these four species, only S. clydet appears to be widely distributed in West Pakistan. Species with central Asian affinities are P. nurt, P. kandelaki, P. keshishianz, S. grekovt, S. palestinensis and S. pawlowskyi. P. chinensis may conveniently be included although it extends from the eastern Mediterranean to China and south to Arabia. All these species occur in or near the northern hills, and Phlebotomus predominates, for example around Keris, where 96 per cent of specimens collected belonged to that genus. 48 D. J. LEWIS Oriental species are P. argentipes, P. colabaensis, S. punjabensis, S. babu, S. shorttit, S. batly1, S. montana and S. hospiti, and S. baghdadis is closely related to two Oriental species. S. sg. indica may be mentioned here but it belongs to a very unusual species, or possibly complex, which extends from West Africa to China (Leng, 1964) and Malaya. The rarity of P. argentipes, P. colabaensis and P. shorttiu at their western limits was to be expected. S. montana is perhaps comparable with the above-mentioned hill anophelines which extend far to the west. S. hospitii also occurs among or near hills and is localized but not rare. S. punjabensis, S. babu, S. baghdadis and S. bailyi are widely distributed, and Table II illustrates the easterly distribution of S. babu, which was pointed out by Sinton. The abund- ance of this species and S. baghdadis is comparable to that of the related S. africana in Africa. On the basis of present knowledge it is possible to summarize some general features of the sand-fly fauna. Most of the Palaearctic species belong to the genus Phlebotomus. The fauna somewhat resembles the northern Ethiopian sand-fly fauna; several species occur in both areas, including the common P. papatasi, S. squamipleuris and S. clydei, and in each case one or more abundant species belong to the subgenus Parrotomyia. In West Pakistan most species with Mediterranean or Oriental affinities occur in the “ tropical’’ zone, and Central Asian forms are found in or near the northern hills. The composition of the fauna is comparable to that of some other groups of animals but the number of species is quite large. P. argentipes and some other rare species. Of the sand-flies known in West Pakistan few species have been found in large numbers, two forms were not encountered at all in the present survey, and eight were represented by single specimens. It is a common experience to find a sand-fly species only after examining hundreds or even thousands of specimens, and this is sometimes due to selection of some species by the special methods of collecting which are necessary for these insects. P. argentipes, being the main vector of kala-azar in eastern India, is of particular interest. Sinton (1925c) reported that it occurred in Ceylon and seemed to be widespread along the coast of India at heights below 460 metres. He stated (19270) that it was known far inland but not west of a line joining Delhi and Bombay, and occurred at Saharanpur, north of Delhi and Sanawar above 1,200 metres in the Simla Hills. He had previously examined thousands of sand-flies from the Sanawar area during seven years (1927e), and he commented on the localization of the species. Sinton (1928) reported it from Burma and expected it to exist in Malaya where it is now known, and (1932) remarked that it was favoured by a moist climate. Qutubuddin (1944) found P. argentipes at Hyderabad, India. Mitra (1954a) pointed out that it had previously been known from warm damp country below 1,220 metres, and reported it from cool dry country up to 1,520 metres in Bombay State and from one place in the Kathiawar Peninsula which is near Pakistan. The species also occurs in Indo-China. An L-shaped line drawn through Kathiawar, Delhi, Lahore and Rawalpindi approximates to the 600 mm. isohyet (Imperial Gazetteer, 1931). The northern arm of this line may also represent a discontinuous extension of P. argentipes parallel to PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 49 _ the northern foothills, where it possibly occurs in certain irrigated areas with dense shade. Near Taxila the orchard of Nikra is a contrast to the barren country around, and in the Lahore area this species has been found near trees (Nasir, 1964). The work of Mitra (1956, p. 237) and Smith (1959) suggests that searches might profitably be made in cattle sheds and houses, particularly in dark corners and crevices near the floor. BIOLOGY Seasonal prevalence. Sinton (19246) reported that sand-flies might vanish for several months in very cold winters, and that they appeared about the end of March and were very abundant by the end of April, their numbers diminishing in the hot dry summer months of June and July and increasing again in the damp weather of August. My visit in 1963 followed an unusually cold winter and sand-flies were not numerous around Lahore in early May or Rawalpindi in late May, but were easy to find in the Keris area in late June. Habitat. There is much to be learnt about the natural habitats in which most species of sand-flies live. Lupascu e¢ al. (1965) in Rumania found that species of Paraphlebotomus, Larroussius and Adlerius were wild sand-flies and might act as vectors of disease in the natural environment. P. major in particular occurred in natural shelters (Lupascu, Dancescu & Cheles, 1965) and seldom approached human settlements (Lupascu ef al., 1965a). In the Soviet Union species of Larroussius, which have large spiracles, predominate in damp areas (Dolmatova, 1962), and in Transcaucasia the hygrophilous P. kandelakw occurs in gardens rather than natural habitats, but S. d. arpaklensis is xerophilous and rarely enters villages (Dergacheva, 1959). The common endophilic sand-flies of West Pakistan include P. papatasi, P. sergentt, S. babu and S. baghdadis. The figures in Table II indicate absence of P. caucasicus Marzinowsky and scarcity of Larroussius, as compared with collections near Tehran in Iran (Lewis et al., 1961). During May, 1963 few P. papatasi or P. sergenti were found in Pakistan houses except bura or underground dwellings near Taxila. During June in the Chilas and Keris areas it was difficult to find sand-flies in houses, probably because people tend to sleep on the flat roofs in summer. Residual spraying might, therefore, be ineffective there at this season, as in the case of Anopheles sergenti in parts of Israel and Jordan (Chang, 1965). With regard to types of outdoor habitat, Dolmatova & Dergacheva (1961) and Dolmatova e¢ al. (1962) found that P. papatasi, S. d. arpaklensis and S. grekovi wei? common in burrows of the large gerbil which appeared to be the natural reservo:t of cutaneous leishmaniasis around Karshi in Uzbekistan. In East Africa kala-azar is associated with certain termite hills in which the vector rests. Other species have this habit, however, in Kenya, Brazil (Martins et al., 1964) and Ghana. During the present survey S. babu and S. bailyi were caught in two termite hills in Lahore, and it seems that these structures harbour many sand flies in places with few other natural shelters. Blood feeding. P. papatasi and P. sergenti are probably the most widespread and numerous man-biting endophilic species, and some other species of Phlebotomus ENTOM. 19, I. 4 50 D. J. LEWIS doubtless bite man in some areas. Several species of Sergentomyia, including S. clydei, sometimes bite man in Africa. A female of S. baghdadis was included in a collection of P. papatasi taken on man at Mir Muhammad, suggesting that further catches on human and animal bait might yield new information. Sand-flies from Shahzada labelled “ biting cattle ’’ included male and female P. papatasi and males of P. sergenti, P. argentipes and S. punjabensis. It might be thought that the presence of males was due to an unusual chance, but this is not necessarily so, for in Central America males of at least one species sometimes accompany females when they are biting man. Some features of sand-flies in the Keris area. In this kala-azar area sand-flies were collected at Gol, Gwadi (2,560 metres), Keris (2,530 metres) and Parkuta. The average number of flies per trap per village was 0-1 to 24, and most of the specimens were males. Sand-flies caught in houses in 1960 (most in June and very few in September) comprised 7 P. sergenti, 20 P. k. burneyi, 5 P. c. longiductus and 1 S. montana. All the others were taken in June, 1963 on sticky traps, more being found near houses and rocks than among cultivation. The complete total for both sexes comprised 1 P. alexandri, 74 P. sergenti, 23 P. k. burneyt, 2 P. keshishianz, 89 P. c. longiductus, 2 S. d. arpakensis, 2 S. p. hodgsoni and 3 S. montana. The high proportion of P. k. burneyi in the 1960 collection suggests it is somewhat endophilic. Perfil’ev (1941) describes P. chinensis and related sand-flies in the U.S.S.R. as “cave species ’’, found mainly out of doors where they attack man. Col. Burney informs me that he was once bitten by many sand-flies in a tent at night in the Keris area but found few in the morning. Many of the P. c. longiductus in this area probably come from rocks near villages. In view of the possible existence of a wild-animal reservoir of leishmaniasis, 18 traps were set about 3 km. from Parkuta up the Indus valley. Three flies were caught, P. c. longiductus and S. p. hodgsoni in the desert and P. keshishiani near the river. At a lower altitude, 1,490 metres around Gilgit, sticky traps in 1963 caught 9 P. sergenti, 5 P. c. longiductus, 1 P. keshishiam, 4 S. babu, 3 S. grekovi and I S. montana. Some parasites. In 1963 single specimens of a relatively large nematode about 3-7 mm. long (Text-figs. 134-136) were found in the abdomen of a teneral female P. papatasi from Taxila, a teneral male P. sergenti from Gilgit, and a female S. clydei from near Karachi. Each worm was folded at two points and was almost full of oval or nearly spherical brown eggs about 0-027 to 0:03 mm. long. The P. papatasz also contained the empty skin of another worm. This parasite can evidently attack quite different species in widely separated areas and seems likely to kill its host and liberate eggs in the soil. It may be that many larvae and pupae are infected but few reach the adult stage. A worm infesting P. papatasi and P. sergent: in Iraq (Adler & Theodor, 1929) may be the same species, but the nematode found in Indian P. papatasi by Mitra (1956) is evidently different. A few mites have been recorded by Jenkins (1964) and others on sand-flies in various countries. Some were found during the present survey and have been examined by Mr. D. Macfarlane, together with J. A. Sinton’s specimens from the PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 51 London School of Hygiene and Tropical Medicine put at our disposal by Dr. B. R. Laurence. The following notes show that several species occur on various phleboto- mines in the subcontinent. Further study may show how specific the true parasites (Stigmaeus, Ledermuelleria and trombidiids) are, and what the sand-flies are doing when they pick up the passenger species. Typhlodromus is one of the genera of Mesostigmata and the others are Prostigmata. Typhlodromus sp. (Phytoseiidae), 2 on P. chinensis from Kasauli. Stigmaeus smithi (Mitra & Mitra, 1953) (= Raphignathus smitht), type specimen on P. papatasi near Poona (Mitra, 1956). Stigmaeus young (Hirst, 1926) (= R. youngt) on 2 P. papatasi at Jamesabad, 3 P. papatasi at Mir Muhammad, ? P. papatasi at Peshawar, and several S. dentata or related form at Lahore. Young ef al. (1926) found that these grey mites, together with less common red trombidiid larvae, infested about four per cent of P. papatasi, mainly females, at Peshawar, and Lewis and Minter (1960) reported a species of Stigmaeus from Kenya. Ledermuelleria sp. (Stigmaeidae), a grey mite, on 2 P. papatasi at Jandola, several P. papatasi at Lahore, 2 P. papatasi at Peshawar, 2 P. papatasi at Sadhana (several nymphs, associated with brown abdominal scars of the host), and Q and ¢ P. major at Kasauli. Most stigmaeid mites are apparently free-living, and all the parasitic ones have been found on sand-flies, namely the two Stigmaeus mentioned above and Ledermuelleria from Central America (Chaudhri, 1965) and West Pakistan. This association is doubtless partly due to the fact that sand-flies, unlike most blood- sucking flies, breed in soil. Trombidium hindustanicum Hirst, 1926, type larva on P. papatasi at Peshawar. Trombidiid larvae of various species on several 2 P. papatasi at Lahore, P. papatasi at Mir Muhammad, several P. papatasi at Peshawar, ¢ Sergentomyia sp. at Karnal, several S. babu at Dehra Dun, and 2 9 S. s. indica at Mangowol. Erythraeid sp. on P. argentipes in India. RELATION “LO DISEASE Several of the sand-fly-borne diseases are zoonoses. The search for vector species therefore involves not only house-frequenting and possibly endophilic species but also wild species which may bite animal reservoirs and possibly man. Irritation. Sinton (19246) found that sand-flies were an intolerable pest on the north-west frontier and in the Punjab, and that scratching led to secondary in- fections and the invaliding of troops. Sand-fly fever. This is caused by at least two types of virus and may have an animal reservoir (Barnett & Suyemoto, Ig61). The disease and its only proved vector, P. papatasi, are widespread in West Pakistan (Nasir, 1964). Barnett & Suyemoto isolated 39 strains of virus of several types from sand-flies around Peshawar and Rawalpindi, and the number of sand-fly species reported in the present paper shows that further study of vectors is necessary. ENTOM. 19, I. 4§ 52 D.jJ. LEWIS Cutaneous leishmaniasis. Oriental sore, due to Leishmania tropica, is extremely common in West Pakistan (Nasir, 1964) where it has practically ceased to be a zoonosis (Garnham, 1965). The survey of this subject by Adler & Theodor (1957) suggests that P. sergenti is the main vector while other species may attack any animal reservoir that exists. S. dentata and S. grekovi were found infected with flagellates in burrows of the rodent reservoir in Uzbekistan (Dergacheva & Dolmatova, 1962) but further study of the flagellates is necessary (Theodor, 19650). Kala-azar. This disease, due to L. donovani, has been found in Baltistan between 2,350 and 2,560 metres above sea level, and its epidemiological pattern seems akin to that of Chinese kala-azar (Ahmad e¢ al., 1960 ; Ahmad & Burney, 1962 ; Barnett & Suyemoto, 1961). Barnett & Suyemoto suggested that P. kandelakiit and P. chinensis might be vectors, and the latter has proved more numerous in collections. Mitra (1959) found a few P. chinensis in the hilly kala-azar district of Riasi E.S.E. of Rawalpindi (Jacob & Kalra, 1951), but, as in the Chilas and Keris areas, indoor catches were small because people slept out of doors. The occurrence of kala-azar in dry uplands accords with the presence of Palaearctic sand-fly vectors, but is a contrast to most of the kala-azar of India, and Napier (1953) suggested that the disease in Riasi might have been due to an indoor micro-focus. Kala-azar has been reported from a few lowland areas (Mitra, 1954) ; Nasir, 1958) but does not seem to have become established. Sinton (1924) ; 19250), writing of the Indian kala-azar then known, reported that its western limit, like that of P. argentipes, seemed to be the Delhi-Bombay line, and he suggested (1927b) that the disease should be sought wherever this sand-fly was discovered. Indian kala-azar associated with P. argentipes occurs in very damp climates with an equable temperature (Napier, 1946; American Geographical Society, 1954; Mitra, 1956, p. 237) and this sand-fly seems unlikely to be a vector in West Pakistan under normal conditions. ACKNOWLEDGMENTS I am grateful to many people for their kind help. Professor H. C. Barnett fostered the work by putting his collections at my disposal and initiating my tour in Pakistan. Dr. E. C. Gangarosa, former Director of the Pakistan Medical Research Centre, was responsible for the tour. Its outcome was largely due to the experience and advice of Lieut. Col. Nur Ahmad, Associate Director, who has done much to further the study of leishmaniasis in Pakistan. Mr. W. A. McDonald, Mr. V. McCarthy, Mr. Z. B. Mirza, Mr. M.S. U. Qadri, Mr. Sana Ullah and Mr. J. Muhammad, then of the same Centre, gave valuable cooperation during the tour. His Highness the Wali of Swat afforded facilities in the Swat Valley. Lieut. Col. A. Rashid and Lieut. Col. M. A. Mannan, Agency Surgeons of Gilgit and Skardu Agencies, organized expeditions in those areas. Surgeon Commander M. Ahram provided an assistant at Karachi. Lieut. Col. M. I. Burney gave me the benefit of his experience of kala-azar in Baltistan, and he and Mr. McDonald and Lieut. Col. J. E. Scanlon kindly contri- buted specimens. Professor O. Theodor gave valuable advice on many taxonomic questions, Professor P. P. Perfil’ev presented specimens of P. smirnovi, Dr. B. R. Laurence provided some mites, and Mr. D. Macfarlane examined them. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 53 REFERENCES Many early papers dealing with sand-flies in West Pakistan are listed in Sinton’s early papers. ABONNENC, E. & MINTER, D. M. 1965. Bilingual keys for the identification of the sandflies of the Ethiopian region. Cahiers O.R.S.T.O.M., Ent. Méd. 5. Apams, A. L. 1867. Wanderings of a naturalist in India .... Edinburgh: Edmonston & Douglas. ADLER, S. & THEODOR, O. 1926. On the minutus group of the genus Phlebotomus in Palestine. Bull. ent. Res. 16 : 399-405. 1927. Ona collection of Phlebotomus sp. of the minutus group. Ann. trop. Med. Parasit. 21 : 61-68. 1929. The distribution of sandflies and leishmaniasis in Palestine, Syria and Mesopotamia. Ibid. 23 : 269-306. 1957. Transmission of disease agents by phlebotomine sand flies. A. Rev. Ent. 2 : 203-226. AumaD, N. & BurNEy, M. I. 1962. Leishmaniasis in northern areas of Pakistan (Baltistan). Pakist. armed Forces med. J. 12 (1) : 1-11. & Wazir, Y. 1960. A preliminary report on the study of kala-azar in Baltistan (West Pakistan). Ibid. 10 (3) : 1-10. AMERICAN GEOGRAPHICAL SOCIETY. 1954. World distribution of leishmaniasis. Atlas of Diseases. Plate 14. ANNANDALE, N. tg10. The Indian species of papataci fly (Phlebotomus). Rec. Indian Mus. 4 : 35-52. Barnett, H. C. & SuyEMoTO, W. 1961. Field studies on sand-fly fever and kala-azar in Pakistan, in Iran, and in Baltistan (Little Tibet) Kashmir. Tvans. N. Y. Acad. Sci. 23 : 609-617. BARTHOLOMEW, J. G., CLARKE, W. E. & GRimsHAw, P. H. trg11. Allas of zoogeography. Edinburgh: Bartholomew. BuHARADWAJ, O. P. 1961. The arid zone of India and Pakistan. Jn L. D. Stamp (Ed.), A history of land use in arid regions. U.N.E.S.C.O. Pp. 143-174. CuanG, L. T. 1965. Resistance status of Anopheles sergenti and its operational importance. World Health Organization Document no. Mal/482.65: 37-43. CHAUDHRI, W. M. 1965. New mites of the genus Ledermuelleria. Acarologia 7 : 467-486. CHRISTOPHERS, S. R. 1933. Fauna of British India, Diptera, IV: Anophelini. London: Taylor & Francis. SHorTT, H. E. & Barraup, P. J. 1926. The anatomy of the sandfly Phlebotomus argentipes, Ann. and Brun. (Diptera). I. The head and mouth parts of the imago. Indian med. Res. Mem. no. 4 : 177-204. DARLINGTON, P. J. 1957. Zoogeography. London: Chapman & Hall. DERGACHEVA, T. I. 1959. On the problem of adaptation of sandflies to various humidities. [In Russian]. Dokl. Akad. Nauk. S.S.S.R. 129 : 431-434. —— & Dormatova, A. V. 1962. On the epidemiology and epizootology of cutaneous leishmaniasis of the rural type in the Karshi Oasis, Uzbekistan. Communication IV. ... [In Russian]. Medskaya Parazit. 31 : 206-211. Dormatova, A. V. 1962. Differences in ecological requirements of some species of Phleboto- minae in the U.S.S.R. Rep. Sympos. 7oth Anniv. V. N. Beklemishev, Moscow. 456-472. —— & DereacHEva, T. I. 1961. On the epidemiology and epizootology of cutaneous leish- maniasis of the rural type in the Karshi oasis, Uzbekistan, I. ... Medskaya Parazit. 30 : 584-591. & EisEEv, L.N. 1962. On the epidemiology and epizootology of cutaneous leishmaniasis of the rural type in the Karshi oasis of the Uzbek S.S.R. Revita Inst. Med. trop. S. Paulo 4: 65-78. FAIRCHILD, G. B. 1955. The relationships and classification of the Phlebotominae. Ayn, ent. Soc. Am, 48 : 182-196, 54 D. J. LEWIS GaRNHAM, P. C. C. 1965. The leishmanias, with special reference to the role of animal reservoirs. Am. Zool. 5: 141-151. Guaur!, M. S. K. 1965. Notes on Hemiptera from Pakistan and adjoining areas. Ann. Mag. nat. Hist. (13) 7: 673-688. Grassi, B. 1907. Ricerche sui flebotomi. Memorie Soc. ital. Sci. nat. (3) 14 : 353-394. GrEssITT, J. L. 1958. Zoogeography of insects. A. Rev. Ent. 3 : 207-230. HEPTNER, V. G. 1938. Origine de la faune désertique du Turkestan et ses particularités zoogeographiques. Byull. mosk. Obshch. Ispyt Prir., N. S. 47 : 338-342. Hirst, S. H. 1926. Report on the acari found on or associated with sandflies in India. Indian J]. med. Res. 13 : 1023-1026. Hutcuinson, G. E. 1934. Report on terrestrial families of Hemiptera Heteroptera. [North India]. Mem. Conn. Acad. Arts Sci. 10: 119-146. IMPERIAL GAZETTEER OF INDIA. 1907. 1. 1931. 26. Oxford: Clarendon Press. Jacos, V. P.& Karra,S.L. 1951. Kala-azarin Kashmir. Indian J. med. Res. 39 : 323-327. Jenkins, D.W. 1964. Pathogens, parasites and predators of medically important arthropods. Bull. Wid Hlth Org. 30: Suppl. Kirk, R. & Lewis, D. J. 1951. The Phlebotominae of the Ethiopian region. Tvans. R. ent. Soc. Lond. 102 : 383-510. LaNE, F. 1942. Comentarios sobre o livro VII de Marcgrave (insetos). Im Jorge Marcgrave : Historia natural do Brasil. Sad Paulo: Museo Paulista. Pp. LXXXVIII-LXXXIX. LenG, Y.-C. 1964. Some new records of Phlebotomus from Hainan ... Acta ent. sinica 13: 118-128. Lewis, D. J. & McMILLAN, B. 1961. The Phlebotominae of Nigeria. Proc. R. ent. Soc. Lond. (B) 30: 29-37. —— Mescuatl, A. & DJANBAKHSH, B. 1961. Observations on phlebotomine sandflies in Tran. Bull. Wid Hith Org. 25 : 203-208. —— & Minter, D. M. 1960. Internal structural changes in some African Phlebotominae. Ann. trop. Med. Parasit. 54 : 351-365. Lorimer, E.O. 1939. Language hunting in the Kavakoram. London: Allen & Unwin. LUNDBLAD, O. 1934. Report on Hydracarina. [North India]. Mem. Conn. Acad. Arts Sci. 10 : 85-118. Lupascu, Gh., Dancesco, P. & CHELES, N. 1965. Contribution a l’étude des espéces de phlébotomes (Diptera, Psychodidae) existants en Roumanie. Archs roum. Path. exp. Microbiol. 24 : 187-194. —— Duport, M., Dancesco, P. & CristEsco, A. 1965a. Recherches sur les espéces de phlébotomes sauvages de Roumanie. Ibid. 24: 195-202. 1965). Recherches sur les espéces de phlébotomes existants dans la nature en Roumanie. Proc. XIIth Internat. Congr. Ent.: 778. Mani, M.S. 1962. High altitude entomology. London : Methuen. MaraInI, F. 1961. Kavrakovam. London: Hutchinson. Martins, A. V., FatcaAo, A. L. and Da Sitva, J. E. 1964. Estudos sébre os flebotomos do Estado de Minais Gerais. VI.... Revta bras. Biol. 24 : 309-315. MatsupDA, R. 1965. The morphology and evolution of the insect head. Mem. Am. ent. Inst. no. 4: 334 pp. MAxwELL-LeFrroy, H. 1909. Indian insect life. Calcutta: Thacker, Spink & Co. MESGHALI, A. 1965. Phlebotominae of Bandar Abbas and Jask area. Bull. Soc. Path. exot. 58 : 259-274. Mitra, R. D. 1954a. Bemerkungen iiber Sandfliegen (Phlebotomen) ... Z. Tvropenmed. Parasit. 5 : 109-113. 19546. Die medizinische Bedeutung der Phlebotomen. Ibid. 5 : 307-317. 1956. Notes on sandflies. Sandflies of the Poona district. Ibid. 7 : 229-240. 1959. Notes on sandflies, Sandflies of Punch and Riasi districts of Kashmir, Ibid, 10 : 56-66, PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 55 Mitra, R. D. & Mirra, S. D. 1953. A new species of Raphignathus (Acarina) associated with Phlebotomus in India. Z. ParasitKde 15 : 429-432. — & Roy, D.N. 1952. Notes on sandflies. Part II. Phlebotomus thapari n. sp. Indian med. Gaz. 87 : 188-193. 1953a. Notes on sandflies, part III. Jbid.: 88: 324-326. 1953b. Notes on sandflies. Part IV. ... Ibid. 88 : 369-372. 1954. Phlebotomus squamipleuris var. poonaensis nov. var. Z. ParasitKde 16 : 191- 194. Munir, A. H. 1963. Observations on the arthropods of medical importance in Gilgit valley. Pakist. armed Forces med. J. 13 : 59-63. MunroE, E. 1965. Zoogeography of insects and allied groups. A. Rev. Ent. 10 : 325-344. Murray, J. A. H. 1888. <A new English dictionary. Oxford ; Clarendon Press. Napier, L.E. 1946. The principles and practice of tropical medicine. New York : Macmillan. 1953. .Kala-azar in Kashmir [review]. Trop. Dis. Bull. 50 : 96-97. Nasir, A. S. 1958. Sandfly fauna in West Pakistan. Pakist. J. Hlth 8: 21-22. 1964. Sandflies as vectors of human disease in West Pakistan. Ibid. 14 : 26-30. NEwsTEAD, R. & Sinton, J. A. 1921. Ona collection of pappataci flies (Phlebotomus) from India. Ann. trop. Med. Parasit. 15 : 103-106. Nicoit, R. M. 1956. Sur la vestiture des Phlebotomidae [DIPT. NEMATOCERA]. Bull. Soc. ent. Fr. 61 : 110-113. Parrot, L. 1942. Notes sur les phlébotomes. XXXIX. ... Phlebotomus fallax. Archs Inst. Pasteur Algér. 20 : 322-335. 1953. Notes... LXVII.—Les “ papilles ’’ des antennes. Ibid. 31: 110-118. & CLASTRIER, J. 1960. Notes...L XXIII. Phlébotomes du Tassili. . . Jbid. 38 : 70-78. PerRFIL’Ev, P.P. 1937.2 Fauna S.S.S.R. Dipterous insects, III, no. 2. Psychodidae (Phleboto- minae). [In Russian]. Moscow: Inst. Zool. Acad. Sci. U.S.S.R. 1939. The sand-fly fauna of the U.S.S.R. [In Russian]. Trudy voenno-med. Akad. RK KA 19: 75-95. 1941. Data on the sand-fly fauna of the U.S.S.R. [In Russian]. Ibid. 25 : 272-283. 1960. Phlebotomus minutus R. and the species of the group minutus (Sergentomyia) in the Crimea and Caucasus [In Russian]. Medskaya Parazit. 29 : 40-48. PRASAD, S. N. & GROVER, P. 1963. The nomenclature of the male genitalia of the cecidomyids (Diptera—Nematocera). Marcellia 31: 45-58. PRATER, S. H. 1965. The book of Indian mammals. Bombay: Bombay nat. Hist. Soc. PRINGLE, G. 1956. Kala azar in Iraq... Bull. endem. Dis. 1 : 235-294. QuaTE, L. W. 1962a. A review of the Indo-Chinese Phlebotominae (Diptera : Psychodidae). Pacif. Insects 4: 251-267. 1962b. Psychodidae (Diptera) at the Zoological Survey of India. Proc. Hawazti. ent. Soc. 18 : 155-188. 1964. Phlebotomus sandflies of the Paloich area in the Sudan. J. med. Ent. 1: 213-268. QUTUBUDDIN, M. 1944. A report on the sandflies of Hyderabad-Deccan (City) with a short note on a new species. Indian J. Ent. 5: 207-211. 1951. The sandfly fauna of the Kohat Valley, N.W. F. P., Pakistan. Pakist. J. Hlth 1 : 34-36. 1952. A comparative study of the two forms of Phlebotomus antennatus Newstead (Diptera, Psychodidae) from Hyderabad, India, and N. W. F. P., respectively. Proc. R. ent. Soc. Lond. (B) 21 : 79-82. 1960a. Mosquito studies in the Indian subregion. Part I....Pacif. Insects 2 : 133-147. 19606. The mosquito fauna of Kohat-Hangu Valley, West Pakistan. Mosquito News 20 : 355-361. SCHMID, F. 1958. Trichoptéres du Pakistan. Tijdschr. Ent. 101 : 181-221, 2 The 1966 revision was received when this paper was in the press, 56 D. J. LEWIS ScumipT, M. L. & Scumipt, J. R. 1962. Variations in antennal ascoid/segment ratio in Phlebotomus papatasi Scopoli (Diptera: Psychodidae). Amn. ent. Soc. Am. 55 : 722-723. 1963. A morphologic study of Phlebotomus papatasi from Egypt (Dipera : Psycho- didae). Ibid. 56 : 567-573. Sinton, J. A. 1922. Entomological notes on field service in Waziristan. Indian J. med. Res. 9: 575-585. 1924a. Notes on some Indian species of the genus Phlebotomus. Part III. Provisional diagnostic table of the males... Indian J. med. Res. 11 : 807-815. — 1924b. Notes ... VIII. Records of the geographical distribution and the seasonal prevalence ... Jbid. 11 : 1035-1049. 1924c. Notes...I1X. Phlebotomus simillimus var. hospitii nov. var. Ibid. 12 : 261-271. —— 1925a. Notes...X. Abnormalities in the appendages... Ibid. 12 : 467-460. 1925b. Notes ... XI. The réle of ... Phlebotomus as carriers of disease ... Ibid. 12 : 701-729. 1925c. Notes... XII. Phlebotomus argentipes ... Ibid. 12 : 789-799. 1926. Notes... XV. Phlebotomus newsteadi n. sp. Ibid. 13 : 559-563. 1927a. Notes ... XVII. Further records of the geographical distribution. Jbid. 14 : 941-945. —— 1927b. Notes... XVIII. Miscellaneous notes. Ibid. 14: 947-953. —— 1927c. Notes... XIX. The value of the female genitalia ... Ibid. 15 : 21-26. —— 1927d. Notes... XXI. Phlebotomus christophersi n. sp. Ibid. 15 : 33-309. —— 1927e. Kala-azar at high altitudes. Indian med. Gaz. 62 : 723. —— 1928a. Notes... XXIII. Phlebotomus clydei n. sp. Indian J. Med. Res. 16 : 179-186. —— 1928). The synonymy of the Asiatic species of Phlebotomus. Ibid. 16 : 297-324. —— 1929. The identification and classification of the species of the genus Phlebotomus, with some remarks on their geographical distribution in relation to disease. Tvans. 7th Congr. far east. Assoc. trop. Med. 3 : 172-193. —— 1931a. Notes... XXVI. Phlebotomus eleanorae n. sp. Indian J. med. Res. 18 : 817. — 1931b. Notes... XXVI. Phlebotomus bailyi n. sp. Ibid. 18 : 821-829. — 1932. Notes...XXxX. Diagnostic table for the females... Ibid. 20 : 55-74. —— 1933a. Notes... XXXII. Phlebotomus dentatus n. sp. Ibid. 20 : 869-872. —— 1933b. Notes... XXXIII. Phlebotomus hodgsoni n. sp. Ibid. 20 : 873-878. —— 1933c. Notes... XXXV. Additions and alterations to the diagnostic table ... [bid. 21 : 225-228. 1933d. Notes... XXXVI. Diagnostic table for the males... Jbid. 21 : 417-428. SMITH, R. O. A. 1959. Bionomics of Phlebotomus argentipes. Bull. Calcutta Sch. trop. Med. Hyg. 7: 17-21. STEARN, W. T. 1960. Allium and Milula in the Central and Eastern Himalaya. Bull. Br. Mus. nat. Hist., Botany, 2: 161-191. THEODOR, O. 1938. On sandflies (Phlebotomus) from Ceylon, Siam, and Malay. Indian J. med. Res. 26 : 261-269. 1948. Classification of the Old World species of the subfamily Phlebotominae (Diptera, Psychodidae). Bull. ent. Res. 39: 85-115. 1952. On the zoogeography of some groups of Diptera in the Middle East. Jstanb. Univ. Fen. Fak. Mecm. 17: 107-119. 1958. Psychodidae-Phlebotominae. Jn Lindner, Fliegen palaearkt. Reg. 9c: 1-55. Stuttgart. 1965a. On the classification of American Phlebotominae. J. med. Ent. 2 : 171-197. 1965). Recent research on transmission of leishmaniasis. Proc. XIIth internat. Congr. Ent. : 771-772. & MeEsGHALI, A. 1964. On the Phlebotominae of Iran. Jbid. 1 : 285-300. TinG, S.-T. & Ho, K.T. 1962. Investigations of the Chinese species of the genus Phlebotomus, Part IX. ... [in Chinese]. Acta ent, sin, 11 : 388-393. PHLEBOTOMINE SAND-FLIES OF WEST PAKISTAN 57 Uvarov, B. P. 1938. Ecological and biogeographical relations of Eremian Acrididae. Mém. Soc. Biogéogr. 6 : 231-273. 1966. Biogeography of Soviet Asia. Nature, Lond. 210: 149. Van EmDEN, F. & HENNIG, W. 1956. Diptera. In S. L. Tuxen Taxonomist’s glossary of genitalia in insects. Copenhagen : Munksgaard. WALTER, R. 1748. A voyage round the world ... by George Anson ... London: J. & P. Knapton (printers). Youne, T. C. M., Ricumonp, A. E. & BrRENpIsH, G. R. 1926. Sandflies and sandfly fever in Peshawar district. Indian J]. med. Res. 13 : 961-1021. [INDEX TO. RECRES AN DsS:U BSPECTES (Synonyms in italics) sp. A, 21 africana, 28 alexandri, 15 antennatus, 27 argentipes, 23 arpaklensis, 25 asiatica, 28 sp. B, 32 babu, 28 baghdadis, 30 bailyi, 38 burneyi, 17 campester, 38 chinensis, 21 christophersi, 40 clydei, 42 colabaensis, 24 deccanensis, 27 dentata, 25 glaucus, 24 grekovi, 32 hindustanicus, 21 hodgsoni, 37 hospitii, 43 indica, 34 kandelaki, 17 keshishiani, 19 latiterga, 42 longiductus, 21 major, 21 major, 21 mediensis, 25 minutus, 25, 27, 28, 39 montana, 39 niger, 28 nuri, 15 palestinensis, 34 papatasi, 14 pawlowskyi, 37 punjabensis, 27 sergenti, 17 sergenti, 15 shorttii, 31 simillimus, 43 squamipleuris, 34 subtilis, 44 theodori, 27 tiberiadis, 44 oh WUs Orr ae SS Z 12 JAN1967 < x % $ eau we 4 Tea eiaishoaaee “9 bins heared do bisanarrenie 4 ae te hae NE ena eat deh leet) oun are é a - Ra pies Tt - dan i een i wy pweigt#l vsgt 4) eee. wa ni s Sestoneanaaésaes est a a ae “eaballwtoiaieeia Vv? & “ae iM Oh Were tate A a oye y c — ¥ aera h © ( S9ir 0 TO Prosper eet @ ; ee 2 = eal! Fo a Pee al | T ewe t dnl, & Vere, GP. | Ast f ii | hea) ocean: iy ae = aed im — lin ‘ ‘ * he) ; i 7 ; ’ @ é ,*o wi ] ' Ne? t 4 } v. tL § a ie a a om. .S #9 *%> ¢ i =) es jv ciabehach Bpowus a ees = b r wit a a . eh r: i fo — ~ i ‘7. a: ¥ x x tial 4 A LIST OF SUPPLEMENTS TO THE ENTOMOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) i. MAsNER, L. The types of Proctotrupoidea (Hymenoptera) in the British Museum (Natural History) and in the Hope Department of Entomology, Oxford. Pp. 143. February, 1965. £5. 2. Nixon, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6. 3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177; 18 plates, 270 Text-figures. August, 1965. {£4 4s. 4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172; 500 Text-figures. October, 1965. £3 5s. 5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 475 Text-figures. November, 1965. {2 15s. 6. OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. Pp. 129; 328 Text-figures. £3. 47. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera : Coccoidea). In press. 8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera : Geometridae). In press. g. HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). In press. Io. STEMPFFER, H. The Genera of the African Lycaenidae (Leipdoptera : Rhopa- locera). In press. PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING os. = 1 MAR ft Ay #L W A REVISION OF THE GENUS PALORUS (SENS. LAT.) (COLEOPTERA : TENEBRIONIDAE) D. G. H. HALSTEAD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 2 LONDON : 1967 A REVISION OF THE GENUS PALORUS (SENS. LAT.) (COLEOPTERA : TENEBRIONIDAE) BY BiG Ho HALSTEAD: . Agricultural Research Council, ae Pest Infestation Laboratory, Slough, England Pp. 59-148 ; 56 Text-figs., 2 Maps BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 2 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the 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 @ separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 19, No. 2 of the Entomological 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.). © Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 28 February, 1967 Price {1 18s. A REVISION OF THE GENUS PALORUS (SENS. LAT.) (COLEOPTERA : TENEBRIONIDAE) By D. G. H. HALSTEAD CONTENTS Page I. INTRODUCTION : ; ‘ ies : : 61 II. GENERIC AND TRIBAL RELATIONSHIPS : : : ; : 62 III. ZOOGEOGRAPHY . ; ‘ : ‘ : ; ; F 63 IV. THE Genus Palorus : : ; 67 V. Tribolium quadricollis (FAIRMAIRE) A AND SPECIES INQUIRENDAE : 67 VI. SYSTEMATICS, KEY CHARACTERS AND INTRA-SPECIFIC VARIATION . 68 VII. NOTES ON KEYS, DESCRIPTIONS AND LABELLING . : 3 5 69 VIII. KEY TOGENERA . 2 : , 69 IX. DESCRIPTIONS OF GENERA AND SPECIES, KEY TO SPECIES. : 72 Palorus Mulsant. , ; ; ; ‘ : : 72 Pseudeba Blackburn gen. rev . : ‘ : ; 124 Austropalorus gen. n. : : ; : : ? ; 129 Palorinus Blair stat.n. . ; : ; 132 Prolabrus Fairmaire and Aas Fairmaire ‘ : : P37 Coelopalorus Blair stat. n. : ; ‘ ‘ : : 140 X. ACKNOWLEDGMENTS : , : : ; ; ‘ ; 145 XI. REFERENCES ; ; é ‘ , ; , , : 146 XII. SPECIES INDEX : : 3 ; , s : . : 147 SYANOE STS Keys to, and descriptions of, seven genera and fifty species are given. One genus is reinstated and two are raised from subgeneric rank. One new genus and twenty new species are described ; one generic and eleven specific synonymies are made. The zoogeography of the species is discussed. : I. INTRODUCTION BEETLES of the genus Palorus and of the other genera included in this study belong to the family Tenebrionidae, tribe Ulomini. Mulsant (1854) erected Palorus as a subgenus of Hypophloeus to contain Hypo- phloeus depressus Fabricius. Jacquelin du Val (1859-63), in his “‘ Genera des Coléoptéres d’Europe ”’, gave Palorus full generic rank, placing Hypophloeus depressus Fabricius as the type and including Hypophloeus ratzeburgii Wissmann. Thomson (1859) described a new genus, Caenocorse, with Hypophloeus depressus Fabricius as type ; Palorus, however, has priority. Champion (1896) gave brief systematic notes on the eight described species referable to Palorus and described a new one. Fleischer (1900) erected a subgenus, Civcomus, to contain Palorus subdepressus (Wollaston). The only major work on Palorus since Champion is that of Blair ENTOM. IQ, 2. 5 62 DG, Hh. BALS LEAD (1930) who dealt with the Indian species, erecting three new subgenera and describing twelve new species. In the present revision subgenera are not used. Two of the subgenera of Blair, Coelopalorus and Palorinus, are raised to generic rank and one new species of Palorinus is described. The genus Pseudeba Blackburn, 1903, described originally in the Colydiidae and later synonymized with Palorus by Carter & Zeck (1937), is reinstated as a distinct Tenebrionid genus closely related to Palorus. Two new species of Pseudeba are described. The genus Platyotus Gerstaecker, 1871 is placed in synonymy with Palorus. One new genus, Austropalorus, containing two new species is described. The close relationship of the genera Prolabrus Fairmaire and Astalbus Fairmaire with Palorus (see Ardoin, 1959) is confirmed. The genus Palorus (sensu stricto of the present revision) currently contains 32 described species; 10 of these are here placed in synonymy, one species, Palorus quadricollis Fairmaire, is removed to Trvibolium (p. 67) and 15 new species are described. II. GENERIC AND TRIBAL RELATIONSHIPS The seven genera Palorus Mulsant, Pseudeba Blackburn, Austropalorus gen. n., Coelopalorus Blair stat.n., Palorinus Blair stat. n., Prolabrus Fairmaire and A stalbus Fairmaire have the following characters in common which, combined, distinguish them from the other members of the tribe Ulomini : (1) Eyes entire and genae tangential to the eyes. (2) Antennae with poorly differentiated, five-segmented club. (3) Scutellum transverse and shaped as in Text-fig. 2. (4) Metendosternite without lamellae, form similar to that in Text-fig. 4i. (5) Males with ‘deep internal pits on the disc of one or more abdominal sternites. (6) Elytra striate-punctate. (7) Wing venation which could easily be derived from that of Coelopalorus foveicollis (Blair) (Text-fig. 55a) by reduction of the anal area. In addition to these common characters, others are shared by two or more genera (see generic descriptions) and the larvae, where known (in Coelopalorus and Palorus) are almost identical in form. Because of the above similarities and their common general facies, itis suggested that these seven genera are closely related and they are subsequently referred to as “ the Palorus genus group ”’. The general facies of the genera Lyphia, Latheticus, Tribolium and Hypophloeus suggest that, of the seventy-four genera contained in the heterogeneous tribe Ulomini, these four are most closely related to the Palorus genus group. The following discussion is based on a knowledge of certain anatomical features (including genitalia, metendosternite and wing venation) of these four genera and of the external characters of the Ulomini in the collection of the British Museum (Natural History), in which a majority of the genera are represented. The female styli in the genera Austropalorus, Pseudeba, Palorus and Astalbus are quite distinct from those of the other Ulomini seen by me. The styli of 1 Deep pits on the underside of the cuticle, each connecting with the surface by a fine canal (Halstead, 1966). REVISION OF GENUS PALORUS (SENS. LAT.) 63 Latheticus oryzae Waterhouse are basically similar, though very different in shape. The aedeagus, in which the basal piece is much shorter than the paramere tube (except in Coelopalorus fovetcollis), is a distinctive feature, contrasting with the aedeagi of the other Ulomini studied, in which the converse is true. The form of the aedeagus in Latheticus is similar to that in Coelopalorus foveicollis but the 9th pleurites in the male are quite different. The metendosternite of the Palorus genus group differs from that of the other genera studied (except Hypophloeus) in that it lacks lamellae. In Hypophloeus, however, the stem is much longer than in the Palorus genus group. Latheticus has an eye ridge very similar to that found in Tribolium (castaneum species group) and the form of the anterior region of the head is similar to that found in Lyphia. Hinton (1948) regards Lyphia and Tribolium as being very closely related genera. Hafeez & Gardiner (1964), on the basis of studies on internal anatomy, particularly the form of the malpighian tubules, place Latheticus very close to Tribolium. Larvae of Tribolium, Latheticus, Palorus and Coelopalorus have paired urogomphi and the margin of the 9th abdominal segment is without spinules. Paired urogomphi are found in many Ulomini but there are usually associated spinules on the margin of the 9th abdominal segment. The characters postulated by Hinton (1948) for the prototype of Tvibolium (and Lyphia) would serve equally well for a common ancestor of the Palorus genus group and since, as has been described above, the genitalia of the latter indicate affinity with Latheticus, the following very speculative suggestions may be made : (i) that Tribolium, Lyphia, Latheticus (World distribution) and the Palorus genus group (Old World distribution) shared a common ancestor prior to the development of the Atlantic rift at the beginning of the Cretaceous (Wilson, 1963), and (ii) that Latheticus diverged from the main line after Triboliwm and Lyphia and before the divergence of the component genera of the Palorus group. III. ZOOGEOGRAPHY The species of the Palorus genus group are all small beetles, varying from 1°5 to 4:0 mm. in length, which, on the evidence of large wings and the frequent light trap records, are probably able to fly. The majority live under the bark of trees or in the galleries of wood-boring beetles but two species, Palorus subdepressus (Wollaston) and Palorus ratzeburgii (Wissmann), are commonly found in stored products and seven species, Coelopalorus foveicollis (Blair), C. carinatus (Blair), Palorinus humeralis (Gebien), Palorus genalis Blair, P. cerylonoides (Pascoe), P. ficicola (Wollaston) and P. laesicollis (Fairmaire) are found there also, but less frequently. Consequently these insects have three methods of dispersal across zoogeographical barriers. They may be carried on air currents associated with tropical storms, on logs or other vegetable materials floating in ocean currents or they may be transported by man in the course of his commercial activities. The distribution of members of the group in the Pacific area appears to be the result of one or more of these dispersal methods (see below). 64 D. G. H. HALSTEAD The apparent absence of small beetles from a particular region may mean that collections have not been made there and distribution maps may reflect chiefly the activity of collectors. The zoogeography described here is inevitably subject to this limitation. The Palorus genus group appears to be absent from the New World, only cosmo- politan species, those associated with stored food products, occurring there. The distribution of the members of this group is discussed under the four main Old World zoogeographic regions, the limits of which are shown in Map I. 1 Distribution of COELOPALORUS Zoogeographical regions ——_—_- foveicollis eo) : Qo b= regions astcues ame carinatus e gu egio PL Palaearctic OR Oriental ET Ethiopian AU Australian Mart. Distribution of Coelopalorus Localities plotted : foveicollis, Inp1aA: Nilgiri Hills, Bangalore; CzEyLon ; Burma: Toungoo, Tenasserim; Maraya: Penang Is., Malacca; Cocos Kreine Is.; Java: Bantam; N. VIETNAM: Hoah Binh; Formosa: Kuraru; PHILIPPINES: Binalnea; Guam Is.; Oaunu Is. ; Hawall. carinatus, INDIA: Nilgiri Hills; CeyLon: Kandy; Hainan: Tunchan; MALaya: Pahang; Java; Hawati (introduced). I. Palaearctic Region Only the genus Palorus is represented in this region. Excluding the cosmopolitan Palorus subdepressus and P. ratzeburgii, and the widespread African species P. ficicola, only three species have been found in this vast region. One of these, P. euphorbiae (Wollaston), is known only from the Canary Islands. The distribution of the other two, P. depressus (Fabricius) and P, orientalis Fleischer, is plotted on Map 2, REVISION OF GENUS PALORUS (SENS. LAT.) 65 Palorus euphorbiae, which is closely related to P. ratzeburgii, was recorded from four of the Canary Islands (Grand Canary, Teneriffe, Hierro and Lanzarote) by Wollaston (1862, 1864, 1865) but recently (Lindberg, 1962) it has been found only on the uninhabited Alegranza and may no longer exist on the other islands. 2 Distribution of PALORUS c cerylonoides d= depressus e neboissi sp.n. f fiecicola m mahenus n nonus spn oO. orientalis Map 2. Distribution of Palorus Localities plotted: depressus; NORWAY; FINLAND; SWEDEN; POLAND: Warsaw; N. GERMANY; FRANCE; SPAIN: Gibraltar; Siciry; ITALY; YUGOSLAVIA; JOMANIA; Caucasus. orientalis, TRANSCAUCASIA (U.S.S.R.); IRAN. ficicola, Eaypt; Linya ; ALGERIA ; MAURITANIA ; CAPE VERDE Is.; GUINEA; N. NIGERIA; CONGO: Elisabethville ; ANGOLA ; RHODESIA: Salisbury. manus sp. n., GuINEA; ConcGo: Elisabethville; RHODESIA: Salisbury ; REPUBLIC oF SouTH AFRICA: Natal; S. ARABIA: Dhala. cerylonoides, IRAN: Abadan ; Inp1a: Dehra Dun and Central Provinces, Bengal; Assam; Burma; N. VIETNAM: Tonkin; PuHitippines; Damma Is.; NEw GuINEA; Sortomon Is.; Fij1; SAMOAN Is. ; Marguesas Is.; Marianas Is.: Guam, Saipan; MApAGASCAR; SEYCHELLE Is. neboisst sp. n., AUSTRALIA: Clermont (Q.) Brisbane (Q.), Dorringo (N.S.W.), Sydney (N.S.W.), Morgan (S.A.), Prospect (S.A.). mahenus, SEYCHELLES ; MADAGASCAR ; ZAMBIA. Palorus depressus is distributed throughout Europe from the Mediterranean to northern Scandinavia (Map 2). In the southern part of its range it occurs most commonly under bark but in Scandinavia it is always associated with the ant, Formica rufa L. The Scandinavian form is usually smaller (mean length of 35 specimens was 2:7mm.) than the general European form (mean length of 44 specimens was 3:0 mm.). It seems possible that this species was able to extend its range northwards to 62° lat. (limit indicated in Lindroth, 1960) only by developing 66 D.G. H. HALSTEAD an association with Formica. The beetle is seldom myrmecophilous in Southern Europe. The Palorus genus group is not indigenous to Britain. Palorus ratzeburgii, an importation, is established in food stores. 2. Ethiopian Region Three genera, Palorus and the Madagascan genera Prolabrus and Astalbus are represented in the Ethiopian region. Most of the Palorus species occur in the rain forests of west and central Africa, but two are widespread. P. ficicola, which was originally described from the Cape Verde Is., extends into the southern Palaearctic (N. Africa) (Map 2). It is often associated with stored products in Africa and has been found in stored products in Asia. The distribution of P. ficicola in Africa may therefore be, to some extent, artificial. P. manus sp. n. however, although widely distributed (Map 2), has not so far been found on stored products. In Madagascar in addition to the endemic genera (which are related to the oriental genera Coelopalorus and Palorinus) there are two species of Palorus, P. cerylonoides and P. mahenus Gebien, which are also present in the Seychelles (Map 2). The oriental cerylonoides may have been imported by man but mahenus is present in Zambia and is probably more widespread in East Africa than at present known. The Malagasy insect fauna is considered to be derived from that of Africa but oriental affinities have been recorded in the mosquitoes (Mattingly, 1962) and the Laemophloeinae (Coleoptera) (Lefkovitch, 1964). 3. Oriental Region Wallace’s zoogeographical line (and other associated zoogeographical limits) do not appear to have significance for Palorus and its allies. The distribution of the Indian species Palorus sinuaticollis Blair and P. shoreae Blair, and perhaps also that of Coelopalorus foveicollis, indicate that as far as this group is concerned Formosa should be retained in the Oriental region. Gressitt (1958) includes the higher mountains of Formosa in the Palaearctic region. Both species of Coelopalorus, C. foveicollis and C. carinatus, are sometimes found associated with stored products in their native countries and are occasionally imported, alive, into Great Britain. The very wide distribution of these species (Map I), particularly in the islands of the Pacific, may thus have been influenced by the activities of man. In Hawaii Coelopalorus foveicollis is well established, occurring in dead branches of Acacia etc., associated with wood-boring beetles. This species has been imported into Kenya and Trinidad. Palorinus humeralis has a wide oriental distribution, though to date it has not been recorded further north than Ceylon. It extends through Malaya, Sumatra, Borneo and Java to New Guinea. Palorus cerylonoides (Map 2) is sometimes associated with stored products in Japan and has been found on stored products imported into Great Britain from the Orient and, rarely, apparently from Africa, It has been caught in light traps, so REVISION OF GENUS PALORUS (SENS. LAT.) 67 both wind (hurricanes etc.) and man (stored products and possibly silviculture) have probably been responsible for the wide distribution of this species. 4. Australian Region Although north-eastern Australia is normally included in the Oriental region (as shown in the maps), for the present purpose the separation is not recognized. The apparently endemic genera, Pseudeba and Austropalorus, appear to be confined to the northern perimeter of Australia (Derby to Townsville). The genus Palorus is represented in Australia by four known (and probably other undiscovered) endemic species and one, P. laxipunctus Fauvel, which occurs also in New Guinea and New Caledonia. The endemic Australian Palorus are morphologically similar to Palaearctic and Pacific (Oriental) species. It seems probable that P. laxipunctus originated in New Guinea, as Papuan elements have been recognized in the faunas of Australia and New Caledonia. New Zealand apparently lacks representatives of the genus group. IV. THE GENUS PALORUS This genus is the largest of the group with thirty-seven known species and probably many more await discovery. The species cannot be readily grouped on morphological grounds and therefore, although close relationships between certain species are manifest, grouping has not been attempted. Certain Palorus species exhibit sexual dimorphism of the genal margin (not found elsewhere in the genus group), most of the Indo-Malayan species and two (or three, if a unique male specimen is included) of the sixteen African species having more strongly developed genae in the male. The Oriental and Ethiopian regions contain the largest number of species and in each region there is an elongate sub-cylindrical species, hypophloeoides Blair (Oriental) and acutangulus sp. n. (Ethiopian), apparently evolved in association with the habitat provided by the galleries of wood-boring beetles. V. TRIBOLIUM QUADRICOLLIS (FAIRMAIRE) AND SPECIES INQUIRENDAE Tribolium quadricollis (Fairmaire) comb. n. Palorus quadricollis Fairmaire, 1902, Annis Soc. ent. Fr. 71 : 331. Tribolium dolon Hinton, 1948, Bull. ent. Res. 39: 47, syn. n. LECTOTYPE, present designation, 9. MADAGASCAR: Andrahomana, bearing labels as follows: ‘“ Madagascar (Sud) Andrahomana Alluaud 1900 38/Palorus quadricoll. Fm n. sp. [Fairmaire’s MS] TYPE [printed red cap’s]”’ the right hand specimen of the pair, in the Paris Museum. Paralectotype 2 mounted on the same card as the lectotype. This species is a member of the genus T7ibolium and is conspecific with Tribolium dolon Hinton of which it is a senior synonym. The holotype of dolon (length 5:3 mm. breadth 1-7 mm.) is larger than the syntypes of quadricollis (length 4-9 mm., breadth 1-4 mm. (both specimens)). 68 D.G. H. HALSTEAD Palorus delicatulus Reitter, 1877, Mitt. miinch. ent. Ver. 1 : 140. Champion (1896) and Blair (1930) failed to recognize Palorus delicatulus Reitter. Champion noted that the type was represented solely by an abdomen and Blair said “It is doubtful whether it is a Palorus at all...” Reitter described Palorus delicatulus from India and said of the pronotum “ ante basin foveolis punctiformibus minutis quatuor leviter impressis”’. I have not seen an Indian Palorus with this character. Specimens of P. beesoni Blair have been sent to me labelled “ P. delicatulus Reitter ”’. Palorus shikhae Sarup, Chatterji & Menon, 1960, Indian J. Ent. 22 : 239. Sarup, Chatterji & Menon (1960) described a new Indian species, Palorus shikhae and subsequently Chatterji, Sarup & Menon (1961) described secondary sexual dimorphism of the tarsi and hind tibiae in P. shikhae. The type material was unobtainable. The authors figure the whole beetle, antenna, mouth parts, male genitalia and (in the 1961 paper) tarsi and hind tibiae of both sexes. These figures bear no resemblance to the structures as found in the Palorus genus group and therefore I believe that shikhae can not be correctly assigned to the genus Palorus or to the other members of the Palorus group. Platyotus glabratus Gerstaecker, see p. 80. VI. SYSTEMATICS, KEY CHARACTERS AND INTRA-SPECIFIC VARIATION The raising of Blair’s subgenera, Coelopalorus and Palorinus, to generic rank is based primarily on their distinctive genitalia (see Text-figs. 4b-e, 49b-d, 55c-f, 56b-e) and is supported by head, elytral and other external morphological characters (see. generic descriptions). If, however, genitalia alone are considered as being indicative of generic limits, the genera Palorus, Pseudeba and Austropalorus gen. n. form a single genus and the two species placed in Coelopalorus could represent two genera ; a consideration of the external morphology of the species concerned (see key to genera and generic descriptions) clarifies the relationships and the reasons for the present arrangement. Genae and pronotum frequently afford useful specific characters. Dorsal puncturation and micro-reticulation of interspaces are sometimes diagnostic, but intra-specific variation occurs. Ventrally the body is more or less uniform within a genus. Because of intra-specific variation, puncturation of the sclerites is only rarely useful as a key character. In Palorus and Palorinus the genitalia are of little or no use in separating closely related species. Body length, pronotal form and genal development are subject to much intra- specific variation in certain species of Palorus. For example in hypophloeoides the body length varies from 2-1 to 3:2 mm., in ficicola pronotal form varies as in Text- figs. 16a—d and in subdepressus genal development varies as in Text-figs. ga-g. In Palorus carinicollis (Gebien) and Palorus crampeli Pic the males bear genal horns which show striking size variation (see Text-figs. 5 and 6). This variation appears REVISION OF GENUS. PALORUS (SENS. LAT.) 69 to be allometric, being dependent on absolute body size, and in carinicollis is associated with development of a medial horizontal prominence of the pronotal apical margin (as in Text-fig. 5). This variation in genal and pronotal development in the male is similar to that found in the family Scarabaeidae (see Arrow, 1951). Vil. NOTES ON KEYS; DESCRIPTIONS AND LABELLING In descriptions of species “length ”’ is the distance from the anterior margin of the clypeus to the elytral apex, and “ breadth”’ is the maximum elytral breadth. All measurements and ratios are given correct to the first decimal place. In a description of elytral interstitial puncturation “‘ approximating to two rows ’’ means that if the total puncturation is considered the majority of the punctures form that number of irregular longitudinal rows. Micro-reticulation was studied at a magnifica- tion of X180. Mandibles and labrum, although usually visible dorsally, have been omitted from figures except Text-figs. 1 and 2 where structures referred to in keys and descriptions are labelled. All types designated or selected in this study have been labelled with the author’s determination labels which have been given coloured borders as follows: HOLOTYPE, red-bordered; PARATYPE, yellow-bordered ; LECTOTYPE, violet- bordered ; PARALECTOTYPE, violet and yellow-bordered. “ Standard B.M. (Nat. Hist.) type labels ’’’, used in description of labels, means circular red-bordered labels. Abbreviations used for names of institutions etc., are as follows: Ardoin Coll. Collection of Monsieur P. Ardoin, Arcachon (20 Rue du Casino 20, Arcachon, Gironde, France). B.M. (Nat. Hist). British Museum (Natural History), London. Frey Mus. Museum G. Frey, Tiitzing. Hung. Nat. Hist. Mus. Természettudomanyi Muzeum (Hungarian Natural History Museum) Budapest. Paris Mus. or Paris Museum Muséum National d’Histoire Naturelle, Paris. P.I.L. Coll. Collection of the Pest Infestation Laboratory, Slough (London Road, Slough, Bucks, England). S.A. Museum (abbreviation on museum label) South Australian Museum, Adelaide. VIII. KEY To GENERA 1 All elytral interstices carinate (Text-fig. 45a) . ; : ; : ' : 6 Elytral interstices not carinate or only interstice 7 carinate ; é 2 2 Elytra with interstice 7 (humeral interstice) carinate (Text-fig. 54a) Oriental COELOPALORUS Blair (p. 140) — Elytra without carinae . ; 3 3 Elytral interstices distinctly raised above striae (Text- fig. 5 3a) ; head form charaé- teristic (Text-figs. 52a, 53a) Madagascar PROLABRUS Fairmaire and ASTALBUS Fairmaire(p. 137) — Elytral interstices not or slightly raised above striae ; head form not as in Text-figs. 52a, 53a . . ; ; , : : : : : : 4 70 D. Gi. SPALSTEAD 4 Anterior margin of clypeus obtusely angled laterally (Text-figs. 48a, 49, 50) ; head puncturation sometimes longitudinally rugose on vertex. Oriental PALORINUS Blair (p. 132) - Anterior margin of clypeus straight or emarginate, not obtusely angled laterally ; head never with longitudinal rugosity on vertex ‘ : . : ; : 5 ! C3 Al Fics. 1-2. Palorus laesicollis (Fairmaire). (1) ventral side, A 1-5, visible abdominal sternites ; C 1-3, pro-, meso- and metacoxae; ES 2, ES 3, episternites of meso- and metasterna ; E M 2, epimerite of mesosternum ; E P, epipleuron of elytron; F, femur ; GS, gular suture; H, hypomeron; L X, lateral extension of procoxa; M, mentum ; M D, mandible ; M P, maxillary palp ; M X, maxilla; PM, prementum ; P R, prosternal process; S 1-3, prosternum, mesosternum and metasternum; SM, submentum; T, tibia; TS, tarsus. (2) dorsal side, AN, antenna of 11 segments; CL, clypeus; FG, flagellum ; GE, gena; HA, humeral angle of elytron; IT, interstices; LF, lateral fovea; LM, labrum; MD, mandible; MP, maxillary palp; N 1-3, pro-, meso- and metanota; PD, pedicel; SA, scape; SC, scutellum; SO, supra-orbital carina ; SP, spiracle ; S R, striae, 10 present ; S S, scutellary striole ; T S, tarsus ; W, wing base, REVISION OF GENUS PALORUS (SENS. LAT.) 71 3 Fic. 3. Palorus laesicollis (Fairmaire). Mouth parts. (a, b) labrum, (a) ventral, (b) dorsal; (c) left mandible, dorsal; (d) right mandible, ventral; (e) labium, ventral ; (f) maxilla, left ventral. AB, tendon of abductor muscle; AD, tendon of adductor muscle; CA, cardo apodeme ; CD, cardo; CN, condyle; GA, galea; L, lacinia; LG, ligula; L P, labial palp; M, mentum; MO, molar lobe; MP, maxillary palp; PM, prementum; PO, peg organs; PP, palpiger; PR, prostheca; ST, stipes. 72 D. G. H. HALSTEAD 5 Maxillary palps moderately securiform (Text-fig. 46b) ; antennae loosely articulated (Text-fig. 46a) ; head form as in Text-fig. 46. Australian AUSTROPALORUS gen. nov. (p. 129) ~ Maxillary palps elongate, not securiform (Text-fig. 3f) ; antennae compact dai figs. 13, 43a) ; head form rarely similar to that in Text-fig. 46 é 6 6 Dorsal margin of eye lower than side margin of head, not margined dorsally by a supra-orbital carina ; front of head flat ; elytra with interstitial punctures comparatively large, equal to one quarter to two thirds diameter of strial punc- tures, or with longitudinal carinae. Australian PSEUDEBA Blackburn gen. rev. (p. 124) — Dorsal margin of eye level with side margin of head (apical region may be concealed dorsally by gena), usually distinctly margined by a supra-orbital carina ; front of head flat or raised ; elytra with interstitial punctures usually much finer than strial punctures, elytra not carinate. . ‘ ; . PALORUS Mulsant (p. 72) IX. DESCRIPTIONS OF GENERA AND SPECIES, KEYS TO SPECIES PALORUS Mulsant, 1854 Hypophloeus (Palorus) Mulsant, 1854, Hist. nat. Col. France 5, Latigenes : 250. Palorus Mulsant; J. du Val, 1859-63, Geneva Coléopt. Europe 3 : 308. Caenocorse Thomson, 1859, Skand. Coleopt. 1: 117. Eba Pascoe, 1863, J. Ent., Lond. 2: 129. Platyotus Gerstaecker, 1871, Arch. Naturgesch. 37 (1) : 62, syn. n. Type-species : Hypophloeus depressus Fabricius, 1790 (by monotypy). Length 1-9-3-8mm.; body moderately depressed to cylindrical, brownish yellow to dark brown, often with head and pronotum slightly darker than elytra, strongly shining to dull; cuticle with micro-reticulation varying from weak and ill-defined to strong and distinct. Head. Usually moderately densely punctured ; clypeus flat or raised in middle, clypeo-genal sutures more or less distinct; genae tangential to eye, often raised above level of clypeus, usually slightly raised above antennal insertions, often moderately produced antero-dorsally, sometimes forming horns in males ; eye latero-ventral, usually large, not emarginate, separated from dorsal surface of head by a supra-orbital carina and limited postero-laterally by small projection of head (see Text-figs. 23a-b). Antennae 11-segmented, rather compact and tightly articulated, inserted beneath genae ; scape usually concealed dorsally by genae, pedicel slightly longer than first flagellar segment, five apical segments forming very indistinct club. Labrum prominent, with dorsal, transverse ridges bearing rows of setae (Text-fig. 3b) ; mandibles with two apical teeth, slightly larger on right mandible than on left (Text-figs. 3c, d) ; mentum with two lateral ridges ventrally and a median protuberance bearing a puncture (? canal) ; labial palps with apical segment ovoid (Text-fig. 3e) ; lacinia without apical tooth or teeth or specialized setae (present in Alphitobius, Tribolium, etc.) ; maxillary palps with apical segment ovoid (Text-fig. 3f). [Description of mouthparts based on those of /Jaesicollis (Text-figs. 3a—f) which appear to be typical for genus ; detailed examination of all species was not possible. } Thorax. Pronotum transverse to elongate, trapezoid (widest at base or apex), quadrate, rectangular, cordiform or with sides evenly arcuate, margined at base, sides and anteriorly on lateral sixth ; sublaterally with narrow almost vertical lenticular flat region (see Text-fig. 6b), lateral margins rarely narrowly explanate ; puncturation moderately dense, becoming coarser towards sides. Scutellum slightly angled apically, usually strongly transverse but only moderately so in cylindrical species (hypophloeoides and acutangulus). Prosternum with median process elongate, margined laterally and apically. Metendosternite (Text-fig. 4i) with stem short, almost quadrate, base narrow, lamellae absent, with tendons near apices. Procoxae with long, lateral, concealed extensions (Text-fig. 1) ; femora simple ; protibiae (Text-fig. 4h) with REVISION OF GENUS PALORUS (SENS. LAT.) 73 apical external angle forming broad tooth, external margin with irregular row of fine setae ventrally, internal margin bearing row of long setae, internal apical angle with two articulated teeth, one large and one small ; tarsal formula 5—5~4 in both sexes ; tarsi simple, apical segment usually longer than combined basal segments, first segment of pro- and meso-tarsi very small. Fic. 4. Palorus laesicollis (Fairmaire). (a) wing—1A, first anal vein? ; 4A, fourth anal vein? ; Cu, cubitus; Ju, jugum; M, media; R, radius; Rm, radio-medial cross vein. (b) pleurites of ¢ 9th abdominal segment ; (c, d) aedeagus (c) dorsal (d) lateral view ; (e) Q genitalia, stylus enlarged; (f, g) 2nd abdominal sternites (f) 2, (g) ¢ (cleared preparations) ; (h) right tibia, ventral ; (i) metendosternite. "4 D. G. H. HALSTEAD Elytva. Free, covering abdomen, with more or less pronounced humeral angles, each elytron usually with ro punctured striae and short scutellary striole ; interstices with punctures smaller than strial punctures, forming I, 2, 3 or, in fenuipunctatus, 4 irregular rows ; epipleura inclined, tapered to apical eighth then narrow toapex. Wings well developed, venation reduced (Text- fig. 4a). Abdomen. Tergites—6 visible but ill-defined ; sternites—5 visible, in cleared preparations 2nd of female bearing fine and diffuse punctures on disc (Text-fig. 4f), that of male appearing to have large deep punctures (= internal pits) sometimes tending to form two patches (Text- fig. 48). Genitalia. with aedeagus (Text-figs. 4c, d; 15b; 23e; 37e; 38e) moderately sclerotized, composed of short basal piece, tubular at base, and long parameres fused dorsally and closed ventrally by narrow membrane, forming paramere tube, partially articulated with basal piece, and with slit-like opening apically ; paramere tube distinctly tapered to apex (Text-fig. 23e), moderately tapered (Text-figs. 4c, d) or moderately or strongly sinuate (Text-fig. 15b) ; median lobe inconspicuous but short basal struts often visible within, and towards base of, paramere tube ; pleurites of 9th segment (= gth sternite of El-Kifl, 1953, in Tvibolium spp.) forming sclerotized ring surrounding aedeagus, bearing few small setae basally (Text-fig. 4b), base more or less rounded, becoming membraneous apically. 9 with styli (Text-fig. 4e) sclerotized, bearing setae apically, and somewhat triangular in cross section, with dorsal edge carinate. Key TO SPECIES OF PALORUS 1 Genal horns present (Text-figs. 5, 6) or genae petaloid (Text-fig. 8) . ; : 37 — Genae not forming horns, not petaloid . 2 2 Pronotum expanded anteriorly and somewhat globose, lateral margin narrowly explanate from basal to apical sixth (Text-fig. 19) ; antennae appearing very short (Text-fig. 19) ; length 2:2-3-o mm. Ethiopian (Seychelles, Madagascar, Zambia) . : : mahenus Gebien (p. 96) — Pronotum not as above ; antennae not appearing very short 3 Pronotum with lateral margins distinctly explanate (Text-fig. 27) and moderately to weakly rounded from base to apex; eyes large; length 3:1-3-7 mm. Ethiopian : : marginatus sp. n. (p. 104) — Pronotum with lateral margins not distinctly explanate ; ; , ‘ ‘ 4 4 Apical pronotal angles very strongly acute (Text-fig. 30) ; body elongate ; genal breadth half that of clypeus ; eyes large with small facets (Text-fig. 30) ; facies characteristic ; length 2-°8mm. Ethiopian . ‘ ‘ acutangulus sp. n. (p. 108) — Apical pronotal angles not very strongly acute ; other characters not present combined. : : 5 5 Antennal length equal to or greater than pronotal length ; pronotum cordiform or elongate (length 2:2-2-6 mm.) ; 6 — Antennae not as long as estate, if nearly as ; long then pronotum not cordiform or elongate : : 7 6 Pronotum cordiform with a “Gistinct fovea medially near base ; puncturation sparse and fine ; supra-orbital carinae ill-defined ; antennae longer than pronotum (antennal length : pronotal length, 1 £09) ; ; length 2-2-2-4 mm. Ethiopian (Text-fig. 15). ‘ nanus sp. n. (p. 91) — Pronotum elongate, frequently with sides evenly arcuate, ‘without a basal fovea ; puncturation moderately dense ; supra-orbital carinae very strongly developed ; antennae as long as pronotum ; length 2:2-2-6mm. Canary Islands (Text-fig. 18). - : euphorbiae (Wollaston) (p. 94) 7 Head witha median somewhat triangular depression producing two low pyramidal prominences traversing the frons (Text-fig. 29) ; pronotum elongate, apical angles strongly acute (Text-fig. 29) ; length 2-6-2-7 mm. Ethiopian baphiae sp. n. (p. 106) - Head not as above ; if pronotum elongate, apical angles not strongly acute 8 Io II I2 13 14 15 REVISION OF GENUS PALORUS (SENS. LAT.) 75 Body cylindrical and elongate (Text-fig. 34a) ; male with frontal tubercles and genae with small projections at clypeo-genal suture ; female without tubercles or projections (though frons with two feebly raised areas); facies distinctive ; length 2-I-3-2mm. Oriental . : ; : hypophloeoides Blair (p. 114) Facies not as above. 9 Pronotum with deep lateral longitudinal Hoveas (Text- ey 13; 14) ; ; genae ‘(Text- figs. 13, 14) not produced antero- ae. to form a semicircular or somewhat triangular flange (or horns) : 10 Pronotum without deep lateral foveae or, if present, head ite genae produced antero-dorsally to form a semicircular or somewhat triangular flange (or horns) (Text-figs. 5a—f) F : : : ; : : ; ; II Body more elongate and cylindrical ; ‘ pronotal foveae more elongate ; eyes larger, dorsal length equal to or slightly less than breadth of scutellum ; prono- tum elongate to quadrate ; elytra more elongate, breadth : length, 1:2; length 2-2-2-5mm. Ethiopian (Text-fig. 14) : : bobiriensis sp. n. (p. 90) Body less elongate, moderately depressed (Text-fig. 13) ; pronotal foveae usually less elongate (Text-fig. 13) ; eyes small, dorsal length approximate to two- thirds scutellum breadth ; pronotum transverse to quadrate ; elytra less elongate, breadth : length, 1 : 1-8 ; length 2-2-2-9mm. Ethiopian laesicollis (Fairmaire) (p. 89) Pronotum widest near apex, without longitudinal shallow depressions (punctures of lateral third mostly separated by more than a puncture diameter) ; dorsal punctures without fine setae ; eyes very small, dorsal length equal to half breadth of clypeal margin ; supra-orbital carina very strongly developed, distinct from apex to base of eye (Text-fig. 23a) ; length 2-4-3:0 mm. Cosmo- politan in stored products, under bark in Europe . ratzeburgii (Wissmann) (p. 96) Without above characters combined. Supra-orbital carina not very sone developed, may be very distinct apically but not basally , : 12 Pronotum widest near apex, with lateral shallow depressions which are ‘broad apically becoming obsolete at base, apical angles as in Text-fig. 25 ; head and pronotum coarsely and densely punctured, all dorsal punctures bearing fine setae ; elytra with comparatively broad lateral rim from base to apex (Text- fig. 25) ; eyes small, dorsal length equal to two-thirds of clypeal breadth ; length 2-8-2-9 mm. Australian : gZrossi sp. n. (p. 102) If pronotum widest near apex, apical angles not as in Text- -fig.25 . 4 ; 13 Pronotum broadest at or close to base (between basal seks and base) (Text- figs. 10, II, 12) ; body form as in Text-fig. looriI2.. : . 14 Pronotum not broadest at or close to base, or body form not as above ; ‘ 16 Pronotal lateral apices obtuse, not produced to form distinct angle (Text-fig. 12), base strongly sinuate ; moderately dull ; body form as in Text-fig. 12 ; length 26mm. Fiji . : ; : obtusus sp. n. (p. 88) Pronotum with distinct apical angles (Text- figs. 10, 12), base almost straight to moderately sinuate ; shining ; body elongate-ovate, strongly convex (Text- fig. 10). Australian and Oriental (Pacific) . : ; ; ‘ : : 15 Larger species, usually longer than 2:8 mm. ; elytra longer, elytral length : prono- tal length, 2-4—2:5 : 1 ; antennae shorter, antennal length : elytral breadth 1 : 2*I-2'4 (mean 2-3) ; lateral marginal rim of elytron usually much broader basally than at middle (Text-fig. 10). Australian and Oriental. laxipunctus Fauvel (p. 86) Smaller species, length 2:4-2-°8mm.; elytra shorter, elytral length : pronotal length 2:2~-2:3 : 1; antennae longer, antennal length : elytral breadth, 1 : 2 ; lateral marginal rim of elytron only slightly broader basally than at middle ; (pronotal sides less convergent (Text-fig. 11) than in normal laxipunctus (Text- fig. r0)). Oriental . ; ‘ , : , F upoluensis Blair (p. 88) ENTOM. IQ, 2. 6 76 16 17 18 19 20 21 22 25 D.G. H. HALSTEAD Pronotal apex very strongly sinuate (Text-fig. 35) ; large elongate-ovate species ; genae strongly raised above level of clypeus, those of male very prominent (Text-fig. 35a), of female moderately prominent (Text-fig. 35b) ; dull; length 3°3-3°8 mm. Oriental : : ; sinuaticollis Blair (p. Pronotal apex if appearing strongly sinuate then form not as above and par or strongly shining Elytra with very distinct, long pubescence, strial setae ‘twice as long as strial punctures (Text-fig. 36, inset) ; (pronotum distinctly pubescent, setae twice as long as punctures (Text-fig. 36, inset), moderately sinuate apically, apical angles strongly produced) ; length 2-°8mm. Oriental (Philippines) kaszabi sp. n. (p. Elytra without very distinct long pubescence, if pubescent strial setae only slightly longer than strial punctures Elytral interstices 2, 3 and 4 with at least three irregular: rows of punctures, four in some regions (Text-fig. 41) ; pronotal apical angles as in Text-fig. 36, acute ; micro-reticulation between punctures of pronotum and elytra deep and dense (Text-fig. 41) ; dull ; length 2-8-3-omm. Oriental . tenuipunctatus Blair (p. Elytral interstices 2, 3 and 4, if with more than two rows of punctures (i.e. with three in some regions), pronotum not with apical angles as in Text-fig. 36 Large species, length 3:3-3-6mm.; pronotum almost quadrate and strongly convex but with disc depressed, basal rim distinctly lower than disc (Text-fig. 39a) ; distinct scutellary striole of 6-7 punctures ; (genae of male angular (Text-fig. 39a), of female straight to slightly rounded (Text-fig. 39b)). India longifoliae Blair (p. Smaller species or, if as large, without the above characters combined, genae variable Length 3-I-3:4 mm. ; pronotum transverse with very distinct ‘pubescence (setae twice as long as punctures) ; dorsal eye length equal to or slightly greater than breadth of clypeal margin. Oriental . , : ; shoreae Blair (p Length less than 3:1 mm. or, if 3:I mm. or more ‘and pronotum with distinct pubescence, dorsal eye length less than breadth of clypeal margin . Elytra short and broad, elytra : pronotum, 2-0—2-2 : 1, (Text-fig. 42a) ; pronotum darker than elytra ; margin of head above eye distinctly sinuate ; (male genae produced and angular (Text-fig. 42a), female genae with margin strongly thickened (Text-fig. 42b)) ; length 2-o-2-8mm. Oriental . andrewesi Blair (p. Elytra not short and broad, without the above characters combined, genae variable Elytral disc usually distinctly depressed (best seen at low magnification) ; prono- tum with lateral longitudinal shallow depressions which, combined with the lateral vertically flattened regions, produce raised borders (Text-fig. 24) ; head as in Text-fig. 24 ; whole body moderately depressed ; length 2-5—3-2 mm. Australian , . neboissi sp. n. (p. Elytral disc not distinctly depressed : if pronotum with shallow lateral longitudinal depressions, head form (i.e. genae or eye size) not as in Text-fig. 24. Pronotum strongly transverse, breadth : length, 1-4 : 1 and somewhat rectangular (Text-fig. 28) ; vertex with a small median depression ; length 2-9—3-3 mm. Ethiopian : ; ardoini sp. n. (p. Pronotum not strongly transverse, breadth : : length ratio less than 1-4: somewhat rectangular then vertex without a median depression Strongly shining, small species 2:2-2-4mm.; pronotum appearing almost quadrate, apical angles as in Text-fig. 26 (‘‘ straight-obtuse’’). Ethiopian ¥, 4£ camerouniensis sp. n. (p. Larger species oy if small and pronotum almost sarge then not strongly shining and apical angles not as in Text-fig. 26 : : Clypeus raised medially and slightly higher than genae (Text- fig. 16a) (length 115) 17 117) 18 122) 19 122) 20 . 116) 21 124) 22 101) 23 106) 24 104) 25 26 27 28 29 30 3r 32 REVISION OF GENUS PALORUS (SENS. LAT.) 2-228 mm.) or if clypeus not raised medially then length 1-9—2:2 mm. and body moderately cylindrical Clypeus not raised medially or, if slightly raised, not higher than genae, body not moderately cylindrical; length 2-3~3-3 mm. Clypeus raised medially and slightly pigs than genae : body moderately elongate and depressed (length 2:2—2-8 mm. ; breadth o-8-o-g mm.) ; pronotum with sides moderately to strongly convergent to base (often somewhat cordi- form) and usually with a small lateral tooth near base (Text-figs. 16a-d) ; basal margin of pronotum not as in Text-fig. 17. Ethiopian (and associated with 77 26 27 stored products from W. Africa and rarely from Asia) . ficicola (Wollaston) (p. 92) Clypeus not higher than genae ; body moderately cylindrical and elongate (length I-9-2'2 mm., breadth 0-5—o-6mm.) ; pronotum more or less quadrate, sides subparallel or slightly convergent to base ; basal margin of pronotum usually as Text-fig. 17, rarely with indentations ill defined. Oriental, and Madagascan subregions of Ethiopian. (Often associated with stored produce from the Orient—rarely in African produce) Genal margin—(i) produced antero-dorsally forming a triangular (Text-fig. 38a, b) or rounded projection (Text-fig. 40), or (ii) produced anteriorly and angularly (Text-fig. 37b), or (iii) forming a slightly lenticular (Text-fig. 7) to semicircular (Text-fig. 5e) flange with hind margin produced posteriorly and covering a large apical fraction of eye, or (iv) genal horns (triangular to strongly Shay present or genae petaloid (Text-figs. 5a—d, 6a—d, 8) ; Genal margin not produced but strongly (Text-fig. 38d) to slightly thickened or flat ; not triangular, rounded, angular, forming a flange or horns or petaloid Body ‘moderately to strongly shining; lateral pronotal punctures without distinct setae ; pronotum may be produced medially and may have deep lateral longitudinal foveae ; genal margin as in couplet 27. (ili) or (iv) Body moderately shining to dull; pronotum laterally with distinct fine pubescence ; pronotum not produced medially and without deep lateral longitudinal foveae ; genal margin as in couplet 27 (i) or (11) Genal margin distinctly thickened, forming a low arcuate ridge (Text- fig. " 38d) ; clypeus clearly differentiated from the genae, strongly shining and comparatively sparsely punctured ; eye length equal to or (usually) greater than breadth of clypeal margin ; length not greater than 2-6mm. ; pronotum not broadest towards apex, but with distinct fine setae. Oriental (sometimes associated cerylonoides (Pascoe) (p. 108) 28 29 oy EF with stored products) : : genalis Blair (9) (p. 118) Genal margin slightly thickened or ‘flat ; without the above characters combined Facies distinctive (Text-fig. 33) ; pronotum slightly cordiform ; head flat anteriorly ; micro-reticulation deep and very distinct (Text-fig. 33, inset) ; dull ; length of holotype 2-5 mm. Australian. : reticulatus sp. n. Facies not as in Text-fig. 33 ; head may be flat anteriorly ; pronotum not somewhat cordiform ; micro-reticulation variable ; shining or dull Clypeus distinctly differentiated from the genae, strongly shining and sparsely punctured ; eye length less than breadth of clypeal margin ; ezther length 2°3-3'2 mm. and pronotum coarsely and densely punctured laterally or length 3°I-3°3 mm. and elytral base broader than pronotal base. Palaearctic . : Clypeus not distinctly differentiated from the genae, approximately equally shining and only slightly less densely punctured ; eye length greater than or slightly less than breadth of clypeal margin ; pronotum not coarsely and densely punctured ; length 2-3-2-9mm. Not Palaearctic : : ; Larger species, 3:I-3:3mm.; pronotum somewhat quadrate, lateral third moderately densely punctured (punctures mostly separated by about or more than a puncture diameter) strongly shining, apical angles usually more strongly produced anteriorly (Text-fig. 21) ; elytra distinctly raised along sutural 32 33 78 33: 34 35 36 37 38 D.G. H. HALSTEAD margin (at least on middle half); antennae slightly longer and broader ; eyes larger. Transcaucasia and Iran . , : : orientalis Fleischer (p. 100) Smaller species, 2-3-3'2mm.; pronotum usually distinctly transverse (rarely somewhat quadrate), lateral third coarsely and densely punctured (punctures mostly separated by much less than a puncture diameter—sometimes punctura- tion somewhat rugose), moderately shining to dull, apical angles less strongly produced anteriorly (Text-fig. 22) ; antennae shorter and narrower ; eyes smaller ; elytra usually not neue raised along sutural margin. Europe and Fennoscandia ; : . . depressus (Fabricius) (p. 99) Apical pronotal angles less obtuse, more strongly produced anteriorly (Text-fig. 31) ; eyes prominent ; side margin of pronotum (seen from side) very slightly raised from basal third or half (Text-fig. 31, inset) to base or straight from apex to base ; sides subparallel or very slightly convergent to base ; strongly to moderately shining ; length 2:2-2-6mm. Australian . intermedius sp. n. Apical pronotal angles more obtuse, less strongly produced anteriorly (Text-figs. 32, 37a) ; eyes variable ; side margin of pronotum (seen from side) usually moderately (Text-fig. 32 inset) or strongly raised from basal half or third to base (rarely almost straight) ; sides subparallel to distinctly convergent to base ; moderately shining to dull , Pronotum with more or less distinct, very shallow, lateral longitudinal depressions extending from approximately apical to basal fifth (Text-fig. 32) ; apical angles of pronotum obtuse, very weakly produced anteriorly ; eyes not prominent ; setae of lateral pronotal punctures usually not very distinct ; form more elongate (Text-fig. 32) ; length 2-3-2-8mm. Oriental: Pacific (and North W. (p. 110) 34 Australia) : : . austrinus Champion (p. 109) Pronotum without lateral depressions : apical angles of pronotum less obtuse (Text-fig. 37a) ; eyes usually prominent; setae of lateral pronotal punctures usually very distinct ; form less elongate (more as in Text-fig. 31) ; length 2:2-2:9mm. Oriental ‘ ‘ : ; beesoni Blair ($9) (p. 119) Eyes prominent with large facets ; pronotal sides moderately rounded from base to apex (Text-fig. 40) ; male genae rounded (Text-fig. 40), female unknown ; antennae comparatively long; length 2-4 mm. (holotype). Oriental (Malay Peninsula) ; ‘ . auranteus sp. n. If eyes prominent, pronotal sides not moderately rounded from base to apex ; genae not rounded ; antennae comparatively short. Oriental : Genae triangular (Text- fig. 38a, b); other characters as in couplet 29 (sometimes (p. 121) 36 associated with stored products) : : . genalis Blair (3) (p. 118) Genae angled at clypeo-genal suture (Text- fig. 37b) beesoni Blair (3, see also couplet 34) (p. 119) Pronotum transverse, laterally with deep, or well-defined shallow, longitudinal foveae (in male, anterior pronotal margin with median horizontal apical prominence of variable size (Text-figs. 5a—d), or sinuate as in female) ; male genae forming horns of variable size which (seen from side) have distinct angle above eye (Text-fig. 5b-d) ; female genae semi-circular, maximum expansion equal to twice dorsal breadth of eye (Text-fig. 5e) ; usually strongly shining ; Length 2-4-3: mm. Ethiopian : . carinicollis (Gebien) (p. 79) Pronotum laterally without deep longitudinal foveae but ill-defined, shallow, depressed areas may be present ; genal horns, when present, without a distinct angle above eye (Text-figs. 6b—d) : ; : ; ; : : . Metatarsal segment ratio, apical segment: basal segments, not greater than I'5 :1; pronotal apical margin slightly to moderately sinuate : : Metatarsal segment ratio greater than 1°5:1 (i.e. I*7-1°8:1); male with pronotal apical margin produced horizontally and medially ; genae petaloid 38 39 (Text-fig. 8) ; length 2-7mm. Ethiopian . ‘ : longitarsus sp. n. (p. 85) REVISION OF GENUS PALORUS (SENS: LAT.) 79 39 Mesosternum very strongly longitudinally rugose (i.e. medial interspaces raised and confluent forming longitudinal rounded ridges, mostly extending from apex to base of mesosternum) ; pronotum quadrate to slightly transverse ; head with vertex always comparatively high with moderately deep depression at base ; male with genal horns of variable size (Text-figs. 6a—d), margin of horn continuous with supra-orbital carina (i.e. without an angle above eye) ; female genal expansion (maximum) equal to or slightly greater than dorsal breadth of eye (Text-figs. 6e, f) ; length 2-5-3: mm. Ethiopian : crampeli Pic (p. 81) - Mesosternum sometimes with coarse and rugose puncturation but not longitudinally rugose ; pronotum transverse ; head with vertex usually comparatively low but variable (Text-fig. 9a—b), basal depression usually shallow ; genae not sexually dimorphic, never as strongly developed as cvampeli in males but sometimes approaching the form of crampeli in females (Text-fig. 9g) ; length 2-7-3:°0mm. Almost Cosmopolitan (instored products) ; apparently indigenous to Ethiopian region (Text-fig. 7) , ; ; subdepressus (Wollaston) (p. 82) Palorus carinicollis (Gebien) comb. n. (Text-figs. 5a-e) Platyotus carinicollis Gebien, 1907, Mems. R. Soc. esp. Hist. nat. 1: 405. Palorus diversicornis Pic, 1924, Mélang. exot.-ent. 41 : 26, syn. n. ? Platyotus glabratus Gerstaecker (see p. 80). Length 2-4-3: mm.; breadth o-7-1:1mm.; usually dark brown, sometimes with dark pronotum and lighter elytra, usually strongly shining; micro-reticulation variable, usually shallow and often distinct on pronotum. Head. 3. Genae produced into triangular to strongly tapered horns of variable size (apparently dependent on absolute size of individual) (Text-figs. 5a—d), forming distinct angle above eye (seen from side) ; eyes large but apical region covered dorsally by horns to greater or lesser extent and appearing small; a distinct depression on each side of head and at base of vertex forming a somewhat triangular region at front of head. 9. Genae (Text-fig. 5e) well developed, forming rounded semicircular flange-like prominences ; eyes large but apical region covered by genae and appearing small dorsally ; maximum dorsal breadth of eye equal to half or less of maximum genal expansion; with distinct depression at base of vertex but lateral depressions not as deep as in male although usually distinct. Pronotum. Transverse, normally with deep lateral longitudinal foveae but sometimes only with moderately shallow but distinct depressions ; in ¢ anterior margin produced horizontally in midline to a greater or lesser extent dependent on absolute size of individual, rarely only sinuate (Text-figs. 5b, c, and d); in 2 anterior margin sinuate (Text-fig. 5e); anterior angles strongly produced anteriorly. Elytra. Interstitial puncturation rather confused and variable but usually approximating to double rows on interstices 2, 3 and 4. LECTOTYPE of carinicollis Gebien, present designation, g. GABON: Nkogo, bearing labels as follows: “‘ ¢/Congo Francese Nkogo XII 1902 L. Fea [blue label]/ Platyotus carinicollis Geb./Cotype No. 47 [orange label] ’’, Frey Mus. Paralectotype of carinicollis Gebien, g¢ GABON: Fernan Vaz, bearing labels as follows: “2 [but is 3]/Congo Francese Fernand-Vaz IX—X 1902 Fea/Cotype No. 47 [orange label] ”’, Frey Mus. LECTOTYPE of diversicornis Pic, present designation, J. SIERRA LEONE, bearing labels as follows ; ‘‘ Sierra Leone/Palorus diversicornis n. sp. ’’ (Pic’s MS), Paris Mus, 80 D.G. H. HALSTEAD Paralectotypes of diversicornis Pic, two examples, SIERRA LEONE, labels in Pic’s manuscript, Paris Mus. Distribution. Ethiopian. GuINEA: N’Zérékoré and Kindia region. SIERRA LEONE. GHANA: Sunyani. TOGOLAND: Bismarkburg. CAMEROUN: N’Kong- samba. SPANISH GUINEA: N’Kolentangan. GABON (see type designation above). Conco: Haut Uélé; Ifuri; Elizabethville. ANGOLA. Habitat. Dr. G. H. Thompson collected this species in Ghana under the dry loose bark of a dead tree (Ficus exasperata Vahl) heavily attacked by bark beetles (Scolytidae) and with many species of Ambrosia beetle (Platypodidae) in the wood. Platyotus glabratus Gerstaecker, 1871, Arch. Naturgesch. 37 : 62. Platyotus glabratus was described from Ugono (Zanzibar) by Gerstaecker (gen. nov. sp. nov.). Gebien (1907) when describing Platyotus carinicollis, stated that he a od 5 Fic. 5. Palorus carinicollis (Gebien). (a) ¢; (b) head and pronotum side view, pronotal apex dorsal view—all of (a) ; (c, d) g, head side view, pronotal apex side and dorsal view ; (e) 9, head and pronotal apex side and dorsal view, REVISION OF GENUS PALORUS (SENS. LAT,) 81 did not know glabratus but that Gerstaecker did not mention anything suggestive of the characteristic features of the prothorax or the basal impression of the elytral stria such as are found in carinicollis. I believe, however, that the possibility of glabratus and carinicollis being synonymous cannot be ruled out altogether. I have failed to get the type material of glabratus and am therefore unable to place Platyotus glabratus Gerstaecker. I would like to thank Dr. F. Hieke of the Zoological Museum, Humboldt University, for the trouble he has taken in searching through the Gerstaecker collections for me. Palorus crampeli Pic (Text-figs. 6a-f) Palorus crampeli Pic, 1924, Mélang. exot.-ent., 41: 26. 2Palorus crampeli var. bicornutus Pic, 1924. Ibid. Length 2-5-3: mm. ; breadth 0-9-1: mm. ; brown to dark brown, usually strongly shining ; micro-reticulation very shallow and indistinct, absent or almost absent on pronotum. Head. 3. Genae produced into triangular to strongly tapered horns of variable size depen- dent on absolute size of individual (Text-figs. 6a—d), horns continuous with supra-orbital carinae (not angled before meeting carinae) ; supra-orbital carina and outer edge of gena forming a straight line; eyes large, only a small region covered dorsally by genal horn; vertex with shallow but distinct depression on each side and shallow depression at base. @: Genae only moderately produced, shape somewhat variable, dependent on breadth of head (Text-figs. 6e, f) ; shallow but usually distinct depression at base of vertex ; eyes generally large, separated ventrally by 2-4—2-8 diameters ; maximum dorsal breadth of eye equal to or very slightly less than maximum genal expansion. Pronotum. Quadrate to transverse; anterior margin only slightly sinuate in both sexes ; sides almost parallel; towards sides rarely with shallow, ill-defined, longitudinal depressed areas. Mesosternum very strongly longitudinally rugose. Elytra. Slightly more elongate than in carinicollis (Text-figs. 5a and 6a) ; interstitial punc- turation somewhat variable and confused, puncturation of interstice 3 and frequently of 2 approximating to a single row. LECTOTYPE, present designation, J. CENTRAL AFRICAN REPUBLIC : Ft. Crampel, bearing labels as follows : ‘‘ Fort Crampel Congo-Frangais Coll. Le Moult Naturaliste, Paris/Coll. K./Palorus ou voisin/Palorus crampeli n. sp.”’ (the latter two labels in Pic’s MS) Paris Museum. Comparative notes. Antennae slightly longer and pronotum more elongate than in carinicollis. Longitudinal rugosity of mesosternum slightly stronger than in carinicollis. Colour usually darker than in subdepressus. See also key and comparative notes on subdepressus. Distribution. Ethiopian. GUINEA: Fouta Djallon and Kindia regions. SIERRA LEONE. GHANA: Mpraeso. CENTRAL AFRICAN REPUBLIC (lectotype). ETHIOPIA. UGANDA: Budongo. Habitat. A specimen from Uganda was found in Scolytid borings in mahogany. ® The var. bicornutus of Pic is a 3 P. crampeli Pic, 82 D. Gi H: HALSTEAD Palorus subdepressus (Wollaston) (Text-figs. 7, ga—g) Hypophloeus subdepressus Wollaston, 1864, Cat. Col. Ins. Canaries B.M.: 499. Palorus subdepressus (Wollaston) Champion, 1896, Entomologist’s mon. Mag. 32 : 27. Palorus bifoveolatus Baudi (nec Duftschmidt). Syn. teste Champion, 1896, Entomologist’s mon. Mag. 32 : 27. Palorus (Circomus) subdepressus (Wollaston) ; Fleischer, 1900, Wien. ent. Zig 19 ; 236. Palorus subdepressus (Wollaston) was frequently confused with P. depressus (Fabricius) by early European entomologists. Length 2-7-3:0mm.; breadth o-9-1:0omm.; red-brown, moderately shining; micro- reticulation shallow but usually distinct. ad 6 Fic. 6. Palorus crampeli Pic. (a) $; (b) head and pronotum of (a) side view ; (c, d) 3, head and pronotal apex side view ; (e, f) 9, head and pronotal apex dorsal view. REVISION OF GENUS PALORUS (SENS. BAT:) Imm TST 229907 00 eo? Imm Fics. 7, 8. 7, Palorus subdepressus (Wollaston). 8, Palorus longitarsus sp. n., head and pronotum. Head.* Usually with more or less distinct shallow median depression towards base—see Text-figs. 9a-g, profile outline ; genae raised above clypeus, developed to a greater or lesser extent independently of sex or head size (Text-figs. ga—g, dorsal), posteriorly produced back- 3 Text-figs. 9a—g. The form of the head is variable, the depth of the depression at the base of the vertex, the development of the genae and the eye size varying as shown in Text-fig. 9. The figures were drawn from photographs (dorsal and ventral views to the same scale, profile to a larger scale) and are of specimens from the following localities : 9a, 9, from culture P.I.L., Slough (stock from Mersin, S. Turkey) ; 9b, 3, Sarawak, Sarikei, in spillage of rice etc., (P.I.L. Coll.) ; 9c, gf, ‘“‘ Tangier/G.C. Champion Coll. B.M. 1927-409 ” (B.M. (Nat. Hist.)) ; 9d, 9, ‘‘ Exped, Mus. G. Frey Franz. Guinea 1951 W. Afr. leg Bechyne/Région Kindia Seguéia 10.5.51’. (Frey Mus.); ge, 3, N. Nigeria, on Sorghum (P.I.L. Coll.) ; of, ‘‘ Salisbury Mashonald Dec. 1900 GAKM. [under card mount]/Marshall Coll. 1911-263 ”’ (B.M. (Nat. Hist.)) ; 9g, 3, “ Leopoldville Belg. Kongo G. Frey 1.1952’ (Frey Mus.). _ The magnitude of difference between heads a, b and f, g is very great, suggesting specific difference but intermediate forms exist between these (heads c, d, e). 83 84 D. G. H. HALSTEAD wards covering apical region of eyes to a variable extent and continuous in a straight line with supra-orbital carinae ; eyes variable in size, separated ventrally by 2-7—4:1 diameters (Text-figs. ga-g, ventral). Pronotum. Transverse, sides slightly rounded to almost parallel; larger specimens some- times with very shallow, ill-defined, lateral longitudinal depressions ; a very small impunctate Fic. 9. Palorus subdepressus (Wollaston). (a-—g) heads, profile, dorsal and ventral views (latter to show eyes) (see p. 83). REVISION OF GENUS PALORUS (SENS, LAT.) 85 region on either side on basal half is often present. Mesosternum usually with coarse, rarely rugose but not longitudinally rugose, puncturation. Elytra. Very slightly depressed. Holotype in British Museum (Nat. Hist.) The cosmopolitan P. subdepressus of stored-products usually has a head form similar to that illustrated in Text-figs. ga and b, i.e., eyes tend to be small and genae poorly developed. African P. subdepressus from the field and from stored-products in West Africa have head forms as illustrated in Text-figs. gc—g, i.e., eyes tend to be large and genae well developed. In any one series from an African locality a large part of the range can be seen. The holotype, collected under camel dung in the Canary Is., approximates to Text-figs. gb-c in head form. Comparative notes. The form of the genae (i.e. forming a flange covering part of the eye and continuous with the supra-orbital carina) separates this species from all others except females of the African species crampeli and carinicollis. Female carinicollis, however, have genae more strongly developed than in subdepressus and usually distinct lateral pronotal foveae. P. subdepressus is not always easily distinguished from female crampeli on head characters but may be separated on characters of the mesosternum—see key. In addition, subdepressus is usually only moderately shining whereas crampeli and carinicollis are usually strongly shining. Distribution. Cosmopolitan—in stored products. The forms with larger eyes appear to be restricted to Africa. P. subdepressus probably originated in Africa, judging from the close relationship with carinicollis and crampeli and the apparent lack of closely related forms elsewhere in the world. Habitat. In Africa subdepressus has been collected at light in large numbers and a colleague, Mr. J. M. Lyall, has collected it under bark of Parkia? stumps in Nigeria with other Palorus species. Beeson, in Blair (1930), includes subdepressus in a list of species collected in the forest (India), definitely under or on the bark of trees attacked by bark beetles or sapwood borers. The under-bark habitat must be the natural one. The Indian record, above, seemingly represents a return to the natural habitat from stored products. In stored products P. subdepressus is a secondary pest frequently associated with the grain weevil, Sztophilus. Hence it is most frequently recorded on cereals and cereal products, often in spillage. Other commodities on which subdepressus has been found are ginger, groundnuts, copra, illipe nuts etc. Palorus longitarsus sp. n. (Text-fig. 8) 3 (2 unknown). Length 2:7mm.; breadth 1:omm.; dark brown, pronotum strongly shining ; elytra moderately shining ; micro-reticulation shallow and indistinct. Head. Genae petaloid (Text-fig. 8) and elevated above level of clypeus ; clypeus and frons strongly shining, clypeus with a few small punctures ; frons towards vertex with a few large punctures (see Text-fig. 8); vertex with strong sparse puncturation and a large, moderately deep depression ; eyes large. 86 D.G. H. HALSTEAD Pronotum (Text-fig. 8). Somewhat transverse, anterior margin produced horizontally ; puncturation strong and dense, punctures separated by one diameter or less, finer on projection. Mesosternum with large punctures but not longitudinally rugose. Elytra. Strial punctures large and deep (deeper than in the related crampeli and carinicollis) ; interstitial punctures fine and shallow, puncturation somewhat confused, approximate to at most two rows in interstices 2 and 3, and one in 4. Legs. Apical segments of tarsi long, metatarsal apical segment nearly equal to twice length of three basal segments, apical segment : basal segments, 1-8 : 1 (basal segments measured from dorsal basal notch of basal segment to apex of third segment) ; tibiae comparatively short, ratio of length of metatibia and metatarsus, 1-3: I. Holotype ¢ (dissected). CAMEROONS: Victoria, bearing labels as follows : “N.W. Kamerun Holiwe, b. Victoria’”’ in the Hungarian Natural History Museum, Budapest. Comparative notes. The form of the genae and the pronotum readily distinguish this species from other Palorus. This species, however, probably exhibits allometric growth, as seen in crampeli and carinicollis, and these characters may therefore prove to be variable. Palorus laxipunctus Fauvel (Text-fig. ro) Palorus laxipunctus Fauvel, 1904, Revue. Ent. 23 : 176. Acthosus pygmaeus Carter, 1914, Trans R. Soc. S. Aust. 38 : 225, syn. n. Palorus pygmaeus (Carter) Carter, 1926, Aust. Zool. 4: 136. Length 2-8-3-1 mm. (one syntype, see below, 2-6 mm.) ; breadth 1-2~-1-4 mm. ; elongate-ovate (Text-fig. 10) ; red-brown, shining ; micro-reticulation shallow but distinct ; punctures of head and pronotum sometimes with quite distinct setae. Head. Moderately densely punctured, punctures separated by 1-2 diameters ; genae forming more or less distinct prominences, in large males similar to genalis, usually raised above level of clypeus ; eyes large and protuberant ; antennal length : elytral breadth, 1 : 2-1-2-4 (mean 2:3). Pronotum. Widest at base, transverse, length: maximum breadth, 1:1-4-1'5; sides moderately convergent to approximately apical fifth then curved to apex ; apical angles obtuse but sharply defined ; basal margin slightly sinuate, slightly expanded in the middle (less so than in upoluensis) ; transversely very convex. Elytra. Elytral length : pronotal length, 2:4-2:5:1; side margin usually much broader basally than at the middle (seen best with light directed onto side of elytron) ; interstitial puncturation somewhat confused but approximating to two rows in interstices 2—4 ; scutellary striole ill-defined, represented by three to six punctures, often all indistinct ; strial punctures moderately deep (slightly deeper in Australian than in New Caledonian type specimens). LECTOTYPE, present designation, 9. NEw CALEDONIA : Noumea, bearing a label as follows: ‘“‘ Noumea F”’ (Fauvel’s MS), left hand specimen of the pair, in the collection of Fauvel in the Institut Royal des Sciences Naturelles de Belgique, Bruxelles. Paralectotypes, three: the right hand specimen on the same mount as the lecto- type, a specimen without data, and one labelled ‘‘ Baie du Sud”’ (Fauvel’s MS (a type locality for laxipunctus) which does not agree well with Fauvel’s descrip- tion. This specimen is abnormally small for the species (length 2-6 mm.) and has REVISION OF GENUS PALORUS (SENS. LAT.) 87 the head flat anteriorly but I believe that it is conspecific with the rest of the type material. Holotype of pygmaeus (Carter) is in the South Australian Museum, Adelaide. cotypes ce ”) I have not seen this specimen but have seen most of the (paratypes). 10 Fics. 10, 11. 10, Palorus laxipunctus Fauvel. 11, Palorus upoluensis Blair, pronotum. 88 D.G. H. HALSTEAD Mr. Gross of the South Australian Museum has kindly compared paratypes seen by me with the holotype and confirmed their conspecificity. Comparative notes. The wide pronotal base combined with size, convexity and the elongate-oval, compact form readily distinguish this Australian Palorus. The species upoluensis has the same general facies but is smaller and has the middle of the pronotal basal rim more strongly expanded (see Text-figs. 10 and 11, and the key for other diagnostic characters). Distribution. AUSTRALIA: “coastal”? Queensland and N.S. Wales, NEw GUINEA and NEW CALEDONIA. Palorus upoluensis Blair (Text-fig. 11) Palorus upoluensis Blair, 1928, Insects Samoa 4: 75. Length 2-4-2:°8mm.; breadth I-o-1:1mm.; facies, puncturation and micro-reticulation similar to that of laxipunctus ; brown, shining. Head. More or less rounded anteriorly, some specimens have genae slightly prominent anteriorly ; genae slightly raised above level of clypeus; antennal length: elytral breadth, 1: 2 (comparatively longer than in laxipunctus). Pronotum (Text-fig. 11). Widest at base, usually slightly less transverse than in laxipunctus and with sides usually more distinctly convergent to approximately apical fifth then curved to apex; apical margin slightly sinuate; anterior angles obtuse but sharply defined; basal margin sinuate, expanded in the middle. Elytva. Facies as in laxipunctus but shorter, elytral length : pronotal length, 2:2-2:3: 1; side margin not much broader basally than at middle. Holotype in British Museum (Nat. Hist.) Comparative notes. This somewhat variable species is very closely related to laxipunctus—see key and laxipunctus ‘‘ Comparative notes ”’ for separation. Distribution. Oriental (Pacific Area). N. Motuccas, NEw Guinea, Moa Is. (off N. Coast of Australia), NEw BRITAIN, SOLOMON Is. and SAMOAN Is. Habitat. It has been collected under dead bark and at light in Samoa and at light in New Britain. Palorus obtusus sp. n. (Text-fig. 12) $ (2 unknown). Length 2-6mm.; breadth 1-1 mm.; red-brown, moderately dull; micro- reticulation distinct. Head. Moderately densely punctured, punctures separated by one diameter or less and bearing setae; clypeus slightly lower than genae; genae very slightly raised along margin ; eyes large and prominent ; supra-orbital carina distinct. Pronotum. Moderately transverse, length : maximum breadth 1 : 1-3, widest towards base ; moderately densely punctured, towards sides punctures bearing long fine setae (length nearly twice the diameter of the larger punctures) ; apical margin straight ; apical angles very obtuse, somewhat rounded ; sides slightly convergent to apex ; basal margin distinctly sinuate (see Text-fig. 12) slightly broader in the middle. REVISION OF GENUS PALORUS (SENS. LAT.) 89 Elyiva. Scutellary striole not differentiated ; interstitial puncturation confused, approxi- mating to one or two rows. Holotype g. Fij1IsLanps: Lovonivonu, bearing labels as follows: ‘“ Fiji Is. Lovonivonu 7.vi.1924 Dr. H. S. Evans/1672.24”’, B.M. (Nat. Hist.). Paratype g. NEw HEBRIDES: Malekula, bearing labels as follows: ‘‘ New Hebrides: Malekula, Ounua iv.v.1929 Miss L. E. Cheesman: B.M. 1929-37 ”’, B.M. (Nat. Hist.) Comparative notes. The obtuse rounded apical angles of the pronotum, the narrow medial region of the basal margin and its form distinguish this species from the rather similarly shaped upfoluensts. Palorus laesicollis (Fairmaire) (Text-figs. 2a, b, 3a-f, 4a—i, 13) Hypophloeus laesicollis Fairmaire, 1893, Annls Soc. ent. Belg. 37 : 28. Palorus laesicollis (Fairmaire) Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 136. Palorus (Coelopalorus) laesicollis (Fairmaire); Gebien, 1940, Mitt. miinch. ent. Ges. 30 : 766. Length 2-2—-2-9 mm. ; breadth o-8-1-0 mm. ; brown, head and pronotum darker than elytra, moderately shining; micro-reticulation variable (usually more distinct than in bobiriensis sp. n.). Head. Supra-orbital carina moderately pronounced ; eye small, dorsal length approximately equal to two-thirds to one half of scutellum breadth. Pronotum. Quadrate to transverse ; laterally with deep longitudinal foveae, deepest towards or at middle (rarely shallower than illustrated, Text-fig. 13); basal margin weakly sinuate. Elytra. Breadth: length, 1: 1-8; single rows of interstitial punctures. Genitalia as in Text-figs. 4b-e. Comparative notes. By means of the deep pronotal foveae this species is readily separated from all others except the closely related bobiriensis sp. n., which, however, is more elongate and has comparatively larger eyes than Jaesicollis. LECTOTYPE, present designation, sex indet. EtHiop1A: Shoa, bearing labels as follows: ‘‘ Choa [Fairmaire’s MS]/72 [printed|/Hypophloeus laesicollis Fairm 1892 Choa [Fairmaire’s MS] ”’, Paris Mus. Fairmaire’s description in “‘ Note sur les Coléopteres du Choa ”’ does not include a particular type locality nor the number of specimens on which the description was based. Distribution. Ethiopian. EtTHiopiA: Shoa and Djem-Djem Forest; KENyA, widely distributed in the Highlands (6—g000 ft.) on stored produce on farms, Messrs C. W. Coombs and J. A. McFarlane (personal communications). Habitat. Under bark of Mimosa, both dead and living trees (Djem-Djem Forest). In Kenya, where it has been found only in stored produce, daesicollis occurred on kibbled (broken) maize cob core, broken maize, oats (4 years in store) and other detritus, often in very large numbers and usually associated with other stored products beetles, particularly Sztophilus spp. go D. G. H. HALSTEAD Palorus bobiriensis sp. n. (Text-fig. 14) Length 2:2—-2-5mm.; breadth o-7-0-8 mm. ; facies similar to /aesicollis but more elongate and less depressed ; brown, distinctly bicoloured, head and pronotum darker than elytra, shining ; micro-reticulation very sparse and ill-defined, almost absent on pronotum. Head. Eyes with dorsal length equal to or slightly less than breadth of scutellum ; supra- orbital carina well developed ; antennal length 0-5—o-6 mm. Pronotum. Elongate to quadrate; with deep lateral elongate foveae deepening slightly towards base (Text-fig. 14) ; sides subparallel to very slightly rounded for apical two-thirds then slightly convergent to base: base more or less sinuate ; strongly shining. Elytra. Elongate, length : breadth, 2: 1; interstitial puncturation in single rows. Holotype ¢ (dissected, left middle leg absent). Conco: Elisabethville, bearing labels as follows: ‘“‘ Belgian Congo 18m S.W. of Elizabethville 1928/Dr. H. S. Evans/Pres. by Imp. Inst. Ent. Brit. Mus. 1932-147” and, on the card, the pencilled number 817-28, B.M. (Nat. Hist.). Paratypes : four examples, three with same locality data as holotype but collected 13.X1.1927 and with the following pencilled numbers on the card mounts: 559/27, 560/27, 561/27; I ex., GHANA: Kumasi, bearing labels as follows: “ Imperial 2 13 Fics. 12-14. 12, Palorus obtusus sp.n. 13, Palorus laesicollis (Fairmaire). 14, Palorus bobiriensis sp. n., head and pronotum. REVISION OF GENUS PALORUS (SENS. LAT.) gI College Ghana Expdn. 1960 7.8.60 Bobiri Forest, Kumasi, Ashanti ’’, all paratypes in B.M. (Nat. Hist.). Palorus nanus sp. n. (Text-figs. 15a, b, Map 2) Length 2-2-2-4mm.; breadth 0o-7-0-8 mm.; elongate and moderately depressed ; yellow- brown to brown, shining ; micro-reticulation sparse and indistinct. Head. Genae slightly raised above antennal insertions; clypeus raised medially ; eyes protuberant ; supra-orbital carina indistinct, visible only at a magnification of x 180; antennae very long, antennal length : pronotal length, 1 : 0-9. 0-5mm a 15 Fic. 15. Palorus nanus sp.n. (a) $; (b) aedeagus, dorsal view. ENTOM. 19, 2 7 92 D.G. H. HALSTEAD Pronotum. Cordiform, slightly depressed ; puncturation sparse and fine (see Text-fig. 15a) ; a small distinct fovea towards base ; sides sinuate. Elytra. Elongate, interstitial puncturation in single rows; strial punctures larger than pronotal punctures. Aedeagus. Text-fig. 15b. Holotype g. GuINEA: Kindia region, bearing labels as follows: ‘‘ Exped. Mus. G. Frey Franz. Guinea 1951 W. Afr. leg Bechyne/Région Kindia Seguéia 10.5.51”’, Frey Mus. Paratypes: eight examples, three with same data as holotype, Frey Mus., B.M. (Nat. Hist.) and Ardoin Coll.; 1 ex., GUINEA: Fouta Djallon, bearing labels as follows: “Exped Mus. G. Frey Franz Guinea 1951 W. Afr. leg Bechyne/Fouta Djallon Dalaba 1200m 16.6.51”’, Frey Mus. ; 1 ex., CONGO: Elisabethville, bearing labels as follows : “ Belgian Congo 18m S.W. of Elizabethville 17.xi.1927 Dr. H. S. Evans/Pres by Imp. Inst. Ent. 1932-147’ and, on card mount, pencilled number 696-27 ; I ex., RHODESIA: Salisbury, under card mount “ Salisbury Dec. 98 under bark G.A.K.M.’’, data label: “ Salisbury Mashonaland G. A. K. Marshall xii. 1898/ Brit. Mus. 1922-431 ; I ex., REPUBLIC OF SOUTH AFRICA: Natal, bearing labels as follows: ‘‘ Malvern Natal 8.97 8642/Malvern, Natal G. A. K. Marshall/Brit. Mus. 1922-431 ’’, this and the previous two specimens in the B.M. (Nat. Hist.) ; 1 ex., S. ARABIA: Dhala, bearing labels as follows: “‘ Taken at moth-screen near rest- house/W. Aden Prot. Dhala, 4,800 ft. 14.ix.1937/B.M. Exp. to S.W. Arabia, H. Scott & E. B. Britton, B.M. 1938-246’, P.I.L. Coll. Comparative notes. Similar to ficicola but easily separated from this species by the long antennae (longer than in all other species though only slightly longer than in euphorbiae), indistinct supra-orbital carinae, smaller size, form of aedeagus etc. Distribution. Ethiopian. See Map 2. Habitat. Under bark (Salisbury). Palorus ficicola (Wollaston) (Text-figs. 16a—d, Map 2) Hypophloeus ficicola Wollaston, 1867, Col. Hesperidum: 205. Palorus ficicola (Wollaston) Champion, 1896, Entomologist’s mon. Mag. 32: 29. Palorus subfilum Fleischer, 1900, Wien. ent. Zig 19 : 237, syn. n. Palorus deserticola Sahlberg, 1913, Ofvers finska VetenskSoc. Férh. 55 (8) : 51, syn. n. (Andres, 1931, placed subfilum Fleischer, in synonymy with deserticola Sahlberg, but Gebien, 1940, did not follow Andres.) Length 2:2-2:°3mm.; breadth o-8-0-9mm; elongate; yellow-brown to red-brown, moderately shining to dull; micro-reticulation distinct. Head. Genae slightly rounded, not produced anteriorly ; clypeus raised medially, punctured as genae; vertex almost flat, usually with a small apical depression ; dorsal length of eye normally greater than breadth of scutellum, supra-orbital carina distinct ; antennal length : pronotal length, 1 : 1-3, in largest specimen seen antennae were slightly shorter. Pyronotum. Variable in shape (Text-fig. 16a—d), cordiform to trapezoid (widest near apex), typical form Text-fig. 16a, extreme form Text-fig. 16d, extreme form occurs in the largest REVISION OF GENUS PALORUS (SENS. LAT.) 93 specimens ; punctures of disc small, separated by 2~3 diameters, towards sides larger and denser ; moderately depressed and usually with a very shallow ill-defined fovea towards base (see Text- fig. 16a) ; side margin usually with a more or less distinct tooth, sometimes indistinct or absent. Elytra. Parallel-sided (as manus sp. n., Text-fig. 15a) ; interstices with single rows of punc- tures ; scutellary striole ill-defined and variable, represented by 4-7 punctures. LECTOTYPE of ficicola Wollaston, present designation (sex indet.). No locality data, in Wollaston’s Cape Verde collection in the B.M. (Nat. Hist.), a small black paint mark traversing the right hand corner of the card mount—Wollaston’s mark denoting S. Iago as the locality, bearing the following labels : “‘ Type [standard B.M. (Nat. Hist.) type label]/ficicola Woll [Wollaston’s MS] ’’. Paralectotypes of fictcola Wollaston, two examples, without locality data, with a black paint mark as in the lectotype, B.M. (Nat. Hist.) general collection. LECTOTYPE of subfilum Fleischer, present designation, (sex indet.). ALGERIA: Biskra, bearing the following labels: “‘ Biskra [MS]/Dr. Puton [MS]/Typus Palorus subfilum Fleischer [Hungarian Museum type label]/Palorus subfilum Fleischer det. dr. Kaszab’’, mounted on a bent micro-pin, Hung. Nat. Hist. Mus. Fleischer (1g00) in his description of this species makes the following statement ‘In Reitter’s Sammlung zwei Examplars aus Biskra als Ratzeburgi von Dr. Puton determinirt ”’. I have located only one specimen. The lectotype of subfilum has a pronotum of the form illustrated in Text-fig. r6c. LECTOTYPE of deserticola Sahlberg, present designation (sex indet.). EGypr: Cairo, bearing labels as follows: ‘“‘ Cairo/J. Sahlb [printed]/deserticola J. Sahlb. Type Heliopolis [MS]/Typus Palorus deserticola Sahlberg [Hungarian Museum type label] ’’ Hung. Nat. Hist. Mus. Paralectotypes of deserticola Sahlberg, five examples, the same locality and kind of card mount as the lectotype, all bearing the printed labels: “‘ Cairo/J. Sahlb’’, four in Hung. Nat. Hist. Mus. and one in Frey Mus. Comparative notes. Similar to vatzeburgi in head structure but eyes larger, body more elongate and usually smaller. General facies similar also to that of cerylonotdes but ficicola is a larger, more depressed species, often with comparatively smaller strial punctures. The pronotal shape is usually distinctive for ficicola. Distribution. (see Map 2). AFRICA (Ethiopian and Palaearctic of N. Africa) ; Colombo (Ceylon) and W. Pakistan in stored products. This species has previously been recorded from Cape Verde (type locality of ficicola), Egypt (deserticola), Algeria and Morocco (subfilum) but its association with stored-products has not previously been recognized. I have seen specimens from Haiti, WEsT INDIEs, 1899, which must represent an importation. Specimens have been seen from the following additional African localities: Lipya, Kufa Oases ; MAURITANIA, Bafrechie ; GUINEA, Dalba ; NIGERIA, Gwoza, Yandev, Kano, Ibadan; Conco (S.W. of Elisabethville) ; RHODESIA ; ANGOLA. Habitat. Under the bark of various trees—Acacia (Algeria), Ficus (Cape Verde), Jatropa (Cape Verde, Lindberg, 1962), Poinciana (Egypt). It has been recorded from Poinciana attacked by Sinoxylon (Andres, 1931) and was collected under the 94 D. G. H. HALSTEAD bark of a dead Parkia with the Colydiid Cicones sqguamosus Grouv., and the Scolytid Xylophagus ferrugineus (F.) by a colleague, Mr. J. M. Lyall, in Nigeria. Associated with stored products I have seen the following: a long series from threshed sorghum heads, N. Nigeria ; two specimens from stored sorghum, Pokoase, Ghana; one specimen from groundnuts, Kano, Nigeria; two specimens from Senegal—collected in milling machinery by Dr. J. A. Freeman (M.A.F.F.) ; two speci- mens collected in spillage in a rice store, Colombo, Ceylon; ten specimens collected from roller mill hinges, mill at Dacca and five from white Pacific wheat in a warehouse, Chittagong. Both these W. Pakistan collections made by Dr. Freeman. Palorus euphorbiae (Wollaston) (Text-fig. 18) Hypophloeus euphorbiae Wollaston, 1862, Tvans. ent. Soc. Lond. 1 : 186. Palorus euphorbiae (Wollaston) Champion, 1896, Entomologist’s mon. Mag. 32 : 29. Length 2-2-2-6 mm. ; breadth 0-7-0-8 mm. ; elongate; brown, moderately shining to dull ; micro-reticulation shallow but distinct. Head. As in Text-fig. 18; almost flat anteriorly ; clypeo-genal sutures ill-defined, clypeus slightly raised medially ; eyes small; supra-orbital carina well developed, almost as pronounced as in vatzeburgii ; antennae very long, equal to length of pronotum. Pronotum. Elongate ; sides sub-parallel to slightly rounded (Text-fig. 18). Elytra. Strial punctures much smaller than pronotal punctures ; interstices with single rows of punctures. LECTOTYPE, present designation (sex indet.). No locality label, in Wollaston’s Canary Island Coleoptera collection in the B.M. (Nat. Hist.), the card mount of the specimen is without a coloured basal margin—denoting Teneriffe as locality, bearing labels as follows: ‘‘ Hypophloeus euphorbiae Woll. type [Arrow’s MS]/Type [standard B.M. (Nat. Hist.) type label] ” card mount held on a “ gold ”’ pin. Paralectotypes, ten examples as follows: 1, labelled “‘ Hypophloeus euphorbiae Woll. type ’’ (Arrow’s MS—as on the lectotype, also card mount without a coloured basal margin) B.M. (Nat. Hist.) general collection. 1, no data labels, basal margin of card mount red—denoting Lanzarote as locality, mount held on a “ gold’ pin, B.M. (Nat. Hist.) Wollaston collection. 1, as above but with a blue basal margin on the card mount—denoting Grand Canary as the locality. 3, basal margin on card mounts red (Lanzarote) and bearing labels as follows “‘ Canary Is. 99-203 ” not on “ gold”’ pins, B.M. (Nat. Hist.) general collection. 4, no data labels, with- out a coloured basal margin on the card mount (meaning?), on “ gold” pins in the Wollaston collection of Canary Island Coleoptera in the Hope Department of Entomology, Oxford. Comparative notes. The form of the pronotum, head and the long antennae (longer than in other Palorus species except nanus sp. n.) readily distinguish euphorbiae. Distribution. CANARY ISLANDS: Grand Canary, Teneriffe, Hierro, Lanzarote (Wollaston 1862, 1864, 1865) Alegranza (Coll. Gonzalez, 1953—Lindberg, 1962). REVISION OF GENUS PALORUS (SENS. LAT;) 95 Habitat. Wollaston, 1864, said of this species “‘...seems to be peculiar to the rotten stems of various Euphorbias—beneath the loose bark of which I have taken ae Lindberg, 1962, observes that although Wollaston collected this species in four of the Canary Islands (see above) it is rarely collected in the Canaries today. He suggests that this is due to the activity of the inhabitants who collect all dead wood for kindling. Alegranza is a small uninhabited island. Scale lines -0:-5mm 18 19 Fics. 16-19. 16, Palorus ficicola (Wollaston): (a) head and pronotum; (b-d) different forms of pronotal side margin. 17, Palorus cerylonoides (Pascoe), pronotum and head. 18, Palorus euphorbiae (Wollaston), pronotum and head. 19, Palorus mahenus Gebien, pronotum and head, 96 D.G. H. HALSTEAD Palorus mahenus Gebien (Text-fig. 19, Map 2) Palorus mahenus Gebien, 1922, Tvans. Linn. Soc. Lond. 18 : 303. Length 2:2-3-0 mm.; breadth o-8-1:2 mm.; short ovate; brown, sometimes with a darker pronotum, shining to dull; micro-reticulation variable. Head. Flat anteriorly ; clypeus very slightly raised above level of genae; genae slightly produced anteriorly ; supra-orbital carina distinct ; antennae short. Pronotum. Expanded anteriorly and somewhat globose (Text-fig. 19) ; side margin sinuate and distinctly explanate from approximately basal to apical sixth, where explanation tapers off. Elytra. Short, subparallel for basal half, then gradually rounded to apex. LECTOTYPE, present designation (sex indet.). SEYCHELLES : Long Island, bearing labels as follows: “ Mahé 335 [MS, on card mount] Type [standard B.M. (Nat. Hist.) type label]/Mahé rg08-9 Seychelles Exp./Percy Sladen Trust Exped. Brit. Mus. 1926-246/Palorus mahenus Geb type! [MS] H. Gebien det 1920/Long Island, Mahe VII. 1908 found in decayed log together with the ant Pheidole punctulata Mayr. other Coleops., and Lepismatidae [MS, H. Scott] ”’, B.M. (Nat. Hist.). Paralectotypes : two examples, SEYCHELLES: Mahé (Long Island), B.M. (Nat. Hist.) and Frey Mus. Comparative notes. The form of the pronotum and the short antennae readily distinguish this species. Distribution. Ethiopian (including the Malagasy subregion). ZAMBIA: Mwengwa ; SEYCHELLES: Long Island (type); Mapacascar. Previously only recorded from the Seychelles. See Map 2. Habitat. The short antennae and compact form suggest a myrmecophilous association (see ‘‘ Lectotype designation ’’ above). Specimens from Zambia were collected at bark sap. Palorus ratzeburgii (Wissmann)! (Text-figs. 20, 23a, b) > Hypophloeus ratzeburgii Wissmann, 1848, Stettin. ent. Zig9: 77. Hypophloeus depressus Stephens (nec Fabricius), 1832, Illusty. British Ent. (Coleopt.5): 7. Hypophloeus ambiguus Wollaston, 1857, Cat. Coleopt. Ins. Madeiva B.M.: 152. Palorus ratzeburgii (Wissmann) Jacquelin du Val, 1859-63, Gen. Coléopt. Europe 3: 308. Palorus floricola Marseul, 1876, Annis Soc. ent. Fr. 6 (5) : 112. Palorus galilaea Sahlberg, 1913, Ofvers. finska VetenskSoc. Férh. 55: 49, syn. n. * Reversion to the original spelling is required by Article 32 of the International Code of Zoological Nomenclature 2nd Edition, 1964. ° I have failed to locate the type material of Hypophloeus ratzeburgii Wissmann. Through the kindness of Dr. E. Schimitschek, I have seen one specimen in Wissmann’s collection (in the Forstzoologische Institut der Universitat Géttingen, Hann. Miinden) collected in Nordheim and determined as H. vatzeburgit (manuscript label, presumed to be in Wissmann’s MS). This specimen conforms to the current concept of P. ratzeburgii (Wissmann). REVISION OF GENUS PALORUS (SENS; LAT.) 97 Length 2-4-3 mm.; breadth 0-9-1: mm.; brown to dark brown, moderately shining rarely dull ; micro-reticulation distinct. Head. Moderately densely punctured; genae not produced or raised, almost level with clypeus, slightly raised above antennal insertions, puncturation a little finer than that of vertex ; clypeus raised medially, puncturation similar to genal puncturation ; eyes small (Text-fig. 23b), dorsal length equal to or less than breadth of scutellum ; supra-orbital carinae very strongly developed (Text-fig. 23a). Pronotum. Usually widest near apex (see Text-fig. 20) but sometimes with sides more parallel ; punctures not bearing distinct setae, small on disc, separated by 4—5 diameters, coarser laterally, separated by 1-2 diameters (usually much finer and sparser than in depressus) ; usually shining. Elytra. Scutellary striole represented by five or less, ill-defined punctures ; interstices with single rows of punctures. LECTOTYPE of ambiguus Wollaston, present designation (sex indet.). No locality labels, in Wollaston’s Coleoptera collection from MADEIRA in the B.M. (Nat. Hist.), bearing labels as follows: ‘‘ Type [standard B.M. (Nat. Hist.) type label]/ambiguus, Woll [Wollaston’s MS} ”’. Paralectotypes of ambiguus Wollaston, two examples, without data labels, in Wollaston’s Coleoptera collection from MADEIRA, in the Hope Department of Entomology, Oxford. LECTOTYPE of floricola Marseul, present designation (sex indet.). JAPAN, bearing labels as follows: ‘‘ Type H.T. [standard B.M. (Nat. Hist.) holotype label]/Japan C. Lewis Ig10—320/ratzeburgi Wissm [Champion’s MS]/M. Lewis 58 Hypophloeus floricola [MS] ”’, left hand specimen, B.M. (Nat. Hist.). Paralectotype of floricola Marseul, on the same card as the lectotype, right hand specimen of the pair. LECTOTYPE of galilaea Sahlberg, present designation (sex indet.). ISRAEL: Nazareth, bearing labels as follows: ‘‘ Nazareth [printed]/U. Sahlb [printed]/galilea J. Sg [MS] Spec. typ [printed]/Palorus galilaea Sahlberg/Palorus galilaea Sahlb det dr. Kaszab’’, Hung. Nat. Hist. Mus. Paralectotypes of galilaea Sahlberg, two examples, locality as lectotype bearing the printed labels: ‘‘ Nazareth/U. Sahlb”’ and on the same form of card mount as the lectotype, Hung. Nat. Hist. Mus. and Frey Mus. Comparative notes. Closely related and similar to depressus, orientalis and ficicola from which it is easily separated on eye size. The strongly pronounced supra- orbital carina distinguishes this species from other Palorus except P. euphorbiae. Distribution. Cosmopolitan (although described from Europe and found in the field there, it is probably of N. African origin). There is a specimen in the Paris Natural History Museum that was collected by Goudot in Madagascar during 1834. Habitat. In Europe ratzeburgii is sometimes found under bark of beech (Fagus) etc. The under bark habitat is apparently the natural one. Specimens have been collected under bark in N. America and represent a recolonization (as in Europe?) of the natural habitat from stored-products. This species, spread to all parts of the world by commerce, occurs as a secondary 98 D.G. H. HALSTEAD pest in granaries, warehouses, flour mills etc., where it is found in cereals and cereal products (often in spillage), and less commonly in other stored produce. It is associated with other pests, particularly the weevils, Sitophilus species. 24 Fics. 20-23. 20, Palorus rvatzeburgii (Wissmann). 21, Palorus orientalis Fleischer, pronotum and head. 22, Palorus depressus (Fabricius), pronotum. 23a, b, P. vatzeburgti, eye (a) dorsal, (b) lateral view. 23c-e, P. depressus, (c) eye dorsal view ; (d) eye lateral view ; (e) aedeagus, REVISION OF GENUS PALORUS (SENS. LAT.) 99 Palorus depressus (Fabricius) (Text-figs. 22, 23c-e, Map 2). Hypophloeus depressus Fabricius, 1790, Sky. nat. Selsk. 1 (1) : 223. Ips unicolor Olivier, 1790, Entomologie 2 (18): 12. Syn. teste Fabricius, 1790, Ent. Syst. 2 (1) : 501. Hypophloeus (Palorus) depressus Fabricius; Mulsant, 1854, Hist. nat. Col. France 5, Latigenes: 250. Palorus depressus (Fabricius) Jacquelin du Val, 1859-63, Gen. Coléop. Europe 3 : 308. Caenocorse depressus (Fabricius) Thomson, 1859, Skand. Coleop. 1: 117. Palorus melinus auctt. (nec Herbst). Syn. teste Champion, 1896, Entomologist’s mon. Mag. 32 : 29. Palorus depressus var. formiceticola Munster, 1928, Norsk ent. Tidsskr. 2 : 296. Length 2-3-3-2mm.; breadth o-9-1:2mm.: facies similar to vatzeburgii; brown to dark brown, usually moderately dull; micro-reticulation distinct, usually deep. Head. Densely punctured; genae slightly raised above the level of the clypeus, densely punctured ; clypeus flat, sparsely punctured, shining; eyes large (Text-fig. 23d); supra- orbital carinae distinct but not as strongly developed as in vatzeburgii (Text-fig. 23c, ratzeburgii Text-fig. 23a) ; antennae appearing slightly thicker than in most species. Pronotum (Text-fig. 22). Widest towards the apex (usually less so than in vatzeburgii), transverse to somewhat quadrate; usually coarsely and densely punctured, disc less so than sides ; sides sometimes almost rugosely punctured, punctures usually larger than strial punctures and bearing distinct setae; apical angles as in Text-fig. 22, obtuse but moderately sharply defined ; apical margin weakly to moderately sinuate ; base slightly to distinctly (in small specimens) narrower than elytral base. Elytva. Scutellary striole of up to 8 (but usually 6) punctures ; puncturation of interstices 2 and 3 variable, one or two rows, puncturation of interstice 3 usually approximating to two rows basally in large specimens but to one in very small specimens (including the ‘‘ type’ of var. formiceticola) ; very slightly raised at suture. . Aedeagus as in Text-fig. 23e, distinctly tapered from basal half to apex. LECTOTYPE, present designation (sex indet.). No locality labels, labelled “ 2 ”’ (type-written label), right elytron missing, in the Fabrician collection, the University Museum, Copenhagen. In addition to the lectotype there are three specimens, numbered “1”, “3” and “4” (type-written labels) no other data labels, placed above the name Hypophloeus depressus in the Fabrician collection. I have been able to examine two of these, ““r’”’ and “4”. Specimen “4” is Palorus subdepressus (Woll.) and probably does not represent part of the original type series. Specimen “1”’ is conspecific with the lectotype but it is in a very poor state of preservation, consisting of an assortment of parts attached to a pin. I would like to thank Dr. S. G. Larsson for making examination of this material possible. The var. formiceticola was described by Munster (1928) as smaller, lighter in colour and with a single row of punctures in the third interstice, occurring in ants’ nests in coastal areas of Norway. In Fennoscandian depressus the colour varies, as does the puncturation of the third interstice (see above) but specimens from this area are smaller. The length range of 35 depressus from Fennoscandia was 2:3-3‘I mm., mean 2-7 mm. and that of 44 specimens from Europe and the Mediterranean area was 100 D.G. H. HALSTEAD 2°5-3'2mm., mean 3:0mm. The overlap of length studied in these 79 specimens was far too great for subspecific differentiation—suggested by distribution and limited habitat. The distribution, habitat and smaller size distinguish this variety. Comparative notes. Similar to orientalis and ratzeburgii. It is readily separated from ratzeburgit on eye size, supra-orbital carina (Text-fig. 23a—-d) and pronotal puncturation (coarse in depressus). From ortentalis it may be distinguished by size and the form of the pronotal apical angles (Text-figs. 22 and 21, and key (p. 78)). Distribution. Palaearctic. FENNOSCANDIA as far north as 62° lat. (var. formiceticola) EUROPE, MEDITERRANEAN and east to CAucAsuS, Map 2. One specimen was seen from Australia, “Sunshine, Victoria (c. oke)’’ undoubtedly an importation. Kocher (1958) records this species from Forét des Zaer, S. Rabat, Morocco (unfortunately I have been unable to obtain the specimen as it is not in Dr. Kocher’s collection). Habitat. Found in nests of Formica rufa L. in Fennoscandia (var. formiceticola). Elsewhere usually under bark especially of oak (Quercus) but sometimes associated with F. rufa. Palorus orientalis Fleischer (Text-fig. 21, Map 2) Palorus orientalis Fleischer, 1900, Wien. ent. Zig 19 : 237. Length 3-1-3:3 mm.; breadth 1-1-1:2 mm.; facies similar to depressus (to which it is very closely related) but body form appearing slightly more elongate ; brown to dark brown, shining, (pronotum always more strongly shining than in depressus) ; micro-reticulation usually shallow but always distinct. Head (Text-fig. 21). Eyes large, slightly larger than in depressus ; antennae very robust and moderately long (see Text-fig. 21). Pronotum (Text-fig. 21). Somewhat elongate, widest towards apex ; puncturation moderately dense ; punctures at sides equal to or slightly larger than strial punctures, punctures bearing setae (not as distinct as in depressus) ; apical angle as in Text-fig. 21, usually more strongly produced than in depressus ; apical margin weakly sinuate to almost straight ; base distinctly narrower than elytral base. Elytra. Scutellary striole ill-defined or represented by 5—6 punctures ; interstitial punctura- tion somewhat confused, interstices 2 and 3 with punctures approximating to one or two rows ; distinctly raised at suture. LECTOTYPE, present designation (sex indet.). TRANscAUCASIA: Talish Mts, bearing labels as follows: ‘‘ Talyschgebg. Transcaucas, Leder, Reitt [printed]/ orientalis Fleisch typ [MS on blue-bordered type label]/Typus Palorus orientalis Fleisch/Palorus orientalis Fleisch det dr. Kaszab’”’, Hung. Nat. Hist. Mus. Paralectotype (sex indet.) “ Lenkoran, Leder (Reitter)/Typus Palorus orientalis Fleischer [red-bordered label] ’’, Hung. Nat. Hist. Mus. Fleischer, in his original description, makes the following statement: “In Reitter’s Sammlung vier ubereinstimmende Example mit Patria; Transkaukasus, Talysch und Lenkoran ; als Ratzeburgi determinirt.’’ There are a number of specimens in the Hungarian Natural History Museum and in the Frey Museum, Tutzing, labelled (Reitter) or coll. Reitter and bearing the locality data indicated by Fleischer—any of these may represent syntypes. REVISION OF GENUS PALORUS (SENS. LAT.) 101 Distribution. Palaearctic, as for types and one specimen in the Frey Museum from IRAN (Map 2). Comparative notes. See key and depressus, ‘‘ Comparative notes ”’. Palorus neboissi sp. n. (Text-fig. 24, Map 2) Length 2:5-3:2mm.; breadth 1-o-1:Imm.; brown, usually dark, pronotum moderately shining to dull, elytra moderately shining ; micro-reticulation variable, on pronotum deep and distinct (as in holotype) to shallow and indistinct, on elytra shallow but distinct. Head. Rounded anteriorly, moderately densely punctured, punctures with short setae ; genae only slightly raised above level of clypeus, not produced anteriorly ; clypeal and genal region almost flat ; supra-orbital carinae distinct ; eyes small, dorsal length equal to or less than clypeal breadth. Pronotum. Slightly transverse (Text-fig. 24), often widest near apex, moderately depressed (a median narrow impunctate region is often present) ; distinct longitudinal depression at each side of disc which, combined with the lateral vertical flattened region, produces a lateral raised border ; apical angles obtuse, not sharply defined, very slightly produced anteriorly ; lateral margins almost straight, parallel or slightly convergent to base ; basal margin straight. Elytra. Disc usually distinctly depressed, rarely only moderately so ; scutellary striole of 2 or 3 punctures—often indistinct ; interstices with single row of punctures. Holotype g. AUSTRALIA: Queensland, bearing labels as follows: ‘‘ Brisbane : H. Hacker 24.6.18/Queensland Museum/Palorus depressus F Id by A. M. Lea Introduced ’’, in Queensland Museum type No T 6353. Paratypes: 20 examples, AUSTRALIA: Queensland, N.S.W., and South Australia 1, bearing labels as follows: 1, ““ Wyreema O. W. Tiegs/S.A. Museum specimen ”’ ; “Wyreema O. W. Tiegs/Queensland Museum ’’, Queensland Museum type No T 6354; 1, “ Dorrigo N.S.W. Jan 19 [0?] r, National Museum of Victoria Melbourne ”’ ; 2 (same card) “‘ Sydney W. du Boulay/H. J. Carter Coll. P. 20.4.22./National Museum of Victoria Melbourne’; 4 (two cards on same pin) “ Sydney viii 31 N.S. Wales Dr. K. K. Spense/K. K. Spense Collection/Palorus depressus F Id by H. J. Carter ’’ in Australian Museum, Sydney ; 4 (same card) ‘‘ Morgan S. Australia A. M. Lea/Palorus? austrinus Champ det H. J. Carter/S.A. Museum specimen’’; 1, “Windsor N.S.W., Lea/S.A. Museum specimen ”’; 1, ‘‘ Yanco N.S.W. 26 Spt. 26 Oct ’32 K. C. McKeown/K 65681/Palorus ? depressus F. Det. H. J. Carter”; 1, “ Prospect 9.03 H.J.C. K 67.16”’, this and the previous specimen both in the Australian Museum, Sydney ; 1, ‘‘ Clermont X 29 Queensland Dr. K. K. Spense/ Palorus (Acthosus) pygmaeus 11.7.13 Carter’ (label with many pin holes!) ; 1, “ Bogan R. N.S. Wales/31/x,25/pygmaeus ”; I, ‘““Blue Mts N.S. Wales/E. W. Ferguson Collection/Palorus ratzeburgi Wissm (introduced) ”; 1, Wahr’ngs 8-19 W. Du B”, this and the previous three specimens in the National Collection, Canberra. Comparative notes. This species is somewhat similar to austrinus. It is, however, larger, has smaller eyes, comparatively deep lateral foveae, deeper strial punctures and has the elytra distinctly depressed on the disc. 102 D. Gi Hy HALSTEAD Distribution. Australian. See Map 2. Species named after Mr. A. Neboiss, National Museum of Victoria, Melbourne. Palorus grossi sp. n. (Text-fig. 25) Length 2-8-2-9mm.; breadth 1:1mm.; brown, moderately shining; micro-reticulation shallow, more or less distinct. Head. Almost flat anteriorly (clypeus and genae), moderately densely and coarsely punctured (puncturation of paratype slightly coarser than that of holotype), punctures with fine setae ; clypeus very slightly raised in the middle with puncturation very slightly sparser than that of genae ; genae very slightly produced at clypeo-genal suture ; eyes small ; supra-orbital carinae distinct ; vertex moderately high. Pronotum. Transverse, sides with lateral rim broad, moderately densely punctured, punctures with fine setae; apical margin sinuate; lateral margin expanded, rim appearing somewhat serrate due to the presence of coarse punctures (see Text-fig. 25); with a shallow lateral 24 25 Fics. 24, 25. 24, Palorus neboissisp.n. 25, Palorus grossi sp. n. REVISION OF GENUS PALORUS (SENS. LAT.) 103 longitudinal depression at each side of disc, deepest towards apex (deeper in paratype than holotype). Elytra. With a comparatively broad lateral rim (broader than in other Palorus spp., see Text-fig. 25) from base to apex; strial punctures deep ; interstitial puncturation somewhat confused but approximating to single rows. Holotype g. AusTRALIA: Victoria, bearing labels as follows: “ Wallan. Vic. C. Oke/Palorus austrinus Champ/National Museum of Victoria Melbourne’ (Text-fig. 25—limbs are absent as indicated by dotted lines). Paratype: g AUSTRALIA: N.S.W., bearing labels as follows: ‘‘ Gosford H. W. Cox/H. J. Carter Coll. P. 20.4.22/Palorus austrinus Champ Id by H. J. Carter/ National Museum of Victoria Melbourne ’’. Comparative notes. Rather similar to austrinus and neboissi sp. n but readily distinguished by the form of the pronotum and the broad lateral rim of the elytra. Distribution. Australian—as above. Species named after Mr. G. F. Gross, the South Australian Museum, Adelaide. 26 Fic. 26. Palorus camerouniensis sp. 0. 104 D. G. H. HALSTEAD Palorus camerouniensis sp. n. (Text-fig. 26) Length 2:2-2-4 mm.; breadth 0-8 mm. ; a small moderately elongate species ; brown, head and pronotum slightly or distinctly darker than elytra, shining ; micro-reticulation shallow, ill-defined. Head. Moderately densely punctured ; clypeus almost flat, slightly less densely punctured than genae ; genae very slightly raised above level of clypeus, not produced, slightly wider than dorsal length of eye ; vertex with a median depression. Pronotum. Almost quadrate, moderately densely punctured, punctures separated by one or two diameters ; apical margin moderately sinuate; apical angles straight—obtuse (Text-fig. 26), with a very slight emargination posteriorly (see Text-fig. 26) in all specimens except one 9 paratype ; sides subparallel; basal margin almost straight. Elytra. Scutellary striole of 2-4 punctures; interstices with single rows of fine punctures. Holotype g. CAMEROUN: N’Kongsamba, bearing labels as follows: “AVR 1957 N’Kongsamba Cameroun, J. Cantaloube ’’, Paris Mus. Paratypes : 4 (2 3, 2 9) with the same data, three in the Paris Museum, one in PL Coil. Comparative notes. General facies similar to vatzeburgit but smaller, more elongate, with larger eyes and a more quadrate pronotum. Distribution. Ethiopian. CAMEROUN. Palorus marginatus sp. n. (Text-fig. 27) Length 3-:1-3:7 mm. ; breadth 1:2~-1'4 mm. ; a large robust species ; brown, moderately dark, pronotum usually darker than elytra, dull, rarely moderately shining; micro-reticulation usually distinct, especially on head and pronotum. Head. fPunctures separated by one diameter ; clypeus slightly raised above level of genae, puncturation much as that of genae; genae not raised or produced ; eyes large with compara- tively small facets ; supra-orbital carinae distinct ; very slight depression at base of vertex. Pronotum. Transverse ; moderately densely punctured, punctures becoming coarse towards the sides; apical margin slightly sinuate or straight; apical angles obtuse, very slightly produced ; lateral margins distinctly explanate, very slightly crenulate, moderately (Text-fig. 27) to slightly (Text-fig. 27, inset) rounded from base to apex ; basal margin slightly rounded. Elytra. Scutellary striole of 3-6 punctures; interstitial puncturation confused, approxi- mating to two rows ; lateral margin with rim slightly broader than usual. Holotype 3 (dissected). ETHIOPIA: Beica, bearing labels as follows : ‘‘ —ix—1960, Beica Wellega Ethiopiae Leg P. Jolivet’’ (Text-fig. 27) in the Paris Museum. Paratypes : eleven examples, three with the same data as the holotype, two in Paris Mus. (3, 9) one in P.I.L. Coll. (9) ; eight, ErHiopi14, bearing the following data : 4, ‘“ Box 31/Under bark of decaying Mimosa/Abyssinia : Djem-Djem Forest nearly 9,000 ft. .x.1926 Dr. H. Scott’’; three in B.M. (Nat. Hist.) Coll., one in P.I.L. Coll. ; 1, ‘‘ Beaten from grass-thatch of hayricks/Abyssinia : Djem-Djem circa 8,000 ft. 6.x.1926 Dr. H. Scott’’; 1, “ From decaying parts of tree Euphorbia abyssinica Rausch/Abyssinia : Djem-Djem Forest circa 8,000 ft. 5-7.x.1926 Dr. H. Scott” ; 105 REVISION OF GENUS PALORUS (SENS. LAT.) E £ ie he ew iy teat ae atne. _—-. 2 Pe. ? soe nerite ai ieee at 270 eS oo 6 oe]: op ee 9990.0) iar Inset variant of pronotal lateral margin. Fic. 27. Palorus marginatus sp. n. 106 D. G. H. HALSTEAD I, ‘‘ Abyssinia: British Legation Pond No. 1 8,100 ft. 8.ix.1926 J. Omer-Cooper ”, the previous 4 specimens in the B.M. (Nat. Hist.). Comparative notes. The explanate margin of the pronotum and the size readily distinguish this species from other African and Palaearctic species. The large eyes, pronotal shape, etc., separate it from the Australian P. grossi sp. n. which has a broad lateral pronotal rim. Distribution. Ethiopian. ETHIopta. Palorus ardoini sp. n. (Text-fig. 28) N’Kongsamba specimens (incl. holotype). Length 3-2-3-3 mm.; breadth 1-3 mm.; Ibadan specimen—length 2-9 mm. ; breadth 1-1 mm. ; brown, head and pronotum slightly darker than elytra, moderately shining ; micro-reticulation distinct. Head. Moderately densely punctured ; clypeus slightly narrower than genae, almost flat ; genae slightly raised above level of clypeus, slightly produced anteriorly ; eyes large with large facets ; vertex with a median depression. Pronotum. Distinctly transverse, length : breadth, circa 1: 1-4 and somewhat rectangular ; moderately densely punctured ; apical margin slightly sinuate ; apical angles somewhat obtuse, weakly produced ; sides subparallel ; basal margin almost straight. Elytra. Scutellary striole usually of 5 or 6 punctures ; interstitial puncturation somewhat confused but approximating to two rows in interstices 2-4. Holotype g. CAMEROUN: N’Kongsamba, bearing labels as follows: ‘‘ Fevrier 57 N’Kongsamba Cameroun J. Cantaloube ”’ in the Paris Museum. Paratypes: five examples, four with the same data as the holotype but collected on different dates, November 56 (3 dissected) in P.I.L. Coll., AVR 1957 ($ dissected and 2 Q) allin the Paris Museum, and one (a small, less robust specimen) NIGERIA : Ibadan, bearing data as follows: “ Nigeria Ibadan at light 26.i.1956/G. H. Caswell Coll. B.M. 1956-673 ”’ in the British Museum (Nat. Hist.) Comparative notes. The very transverse rectangular pronotum readily distin- guishes this species. Distribution. Ethiopian. CAMEROUN and NIGERIA. Species named after Mons. P. Ardoin, Arcachon, France. Palorus baphiae sp. n. (Text-fig. 29) 3 (2 unknown). Length 2-6-2-7mm.; breadth o-8mm.; elongate; brown, pronotum slightly darker than elytra, somewhat dull to moderately shining ; micro-reticulation strong and distinct. Head. Moderately densely punctured ; clypeus slightly raised in the middle, slightly less densely punctured than genae and more strongly shining; genae very slightly raised, very slightly produced anteriorly ; vertex with a deep, somewhat triangular depression—producing two low pyramidal prominences traversing the frons (Text-fig. 29); eyes large with small facets ; supra-orbital carinae distinct. REVISION OF GENUS PALORUS (SENS. LAT.) 107 Pronotum. Elongate; moderately densely punctured; a shallow depression towards base, apical margin sinuate ; apical angles acute ; sides almost parallel for apical three-quarters, then convergent to base ; basal margin slightly rounded. Elytra. Scutellary striole ill-defined, of approximately three punctures; interstices with single rows of punctures. Holotype 3. GHANA: Mpraeso, bearing labels as follows: “ A.W. 34 1:3: 46 Gold Coast, Mpraeso, 1945-46 G. H. Thompson ’”’ B.M. (Nat. Hist.). Paratypes : two (9) with the same data, in the B.M. (Nat. Hist.). Comparative notes. The form of the head and the elongate body readily distin- guish this species. Distribution. Ethiopian. GHANA. Imm 28 Fics. 28, 29. 28, Palorus ardoini sp.n. 29, Palorus baphiae sp. n. ENTOM. IQ, 2 8 108 D.G. H. HALSTEAD Habitat. Dr. G. H. Thompson has kindly given me the following information about these specimens : “A.W. 34: Baphia pubescens Hook. f. Adults and one pupa were in the wood of a small tree that had been felled 5 months previously in a teak plantation at Kwahu Tafo, Mpraeso Forest District, which is in the semi-deciduous high forest. Ceram- bycid and Bostrychid larvae were also in the wood. ” Palorus acutangulus sp. n. (Text-fig. 30) 3 (Q unknown). Length 2:°8mm.; breadth 0-7mm.; elongate, moderately cylindrical ; yellow-brown, shining ; micro-reticulation ill-defined. Head. Punctures separated by one to two diameters ; clypeus very large, breadth greater than dorsal length of eye, slightly raised medially, slightly raised above level of genae ; genae small, not produced, equal to approximately half breadth of clypeus; genae and clypeus moderately shining, puncturation of approximately equal density ; frontal region with two slightly raised areas ; vertex with a depression towards base; eyes large with small facets ; supra-orbital carinae long and distinct. Pronotum. Elongate, depressed on disc, moderately densely punctured, punctures approximately equal in size to those of elytral striae ; apical margin very slightly bi-arcuate; apical angles very strongly acute ; sides subparallel, slightly convergent towards base ; basal margin very slightly rounded. Elytra. Scutellary striole of 8-10 punctures; interstices with single rows of punctures, punctures only slightly smaller than strial punctures. Holotype ¢ (dissected). CAMEROUN: N’Kongsamba, bearing labels as follows : “AOU 1957 N’Kongsamba Cameroun J. Cantaloube”’, in the Paris Museum. Comparative notes. This species is readily recognised by the very acute apical pronotal angles, the large eyes with small facets and the elongate form. It is some- what similar, in general facies, to certain Hypophloeus (sens. lat.). Palorus cerylonoides (Pascoe) (Text-fig. 17, Map 2) Eba cerylonoides Pascoe, 1863, J. Ent., Lond. 2: 129. Palorus exilis Marseul, 1876, Annls Soc. ent. Fr. 6 (5) : 116. Palorus minor Waterhouse, 1894, Ann. Mag. nat. Hist. 14 (6) : 71. Palorus praslinensis Gebien, 1922, Trans. Linn. Soc. Lond. 18 : 304, syn. n. Palorus cerylonoides (Pascoe) Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 141. Palorus papuanus Kaszab, 1939, Nova Guinea (n.s.) 3: 218, syn. n. Palorus zimmermani Kaszab, 1955, Proc. Hawait. ent. Soc. 15 (3) : 657, syn. n. Length 1-9-2:2mm.; breadth o-5-o-6mm.; elongate, moderately cylindrical; brown to yellow-brown, shining ; micro-reticulation shallow and ill-defined. Head (Text-fig. 17). Moderately densely punctured, punctures bearing fine setae ; anterior margin somewhat angulate ; clypeus broad and flat, broader than dorsal length of eye, sparsely punctured, shining ; genae not raised above level of clypeus, anterior margin straight ; vertex strongly convex. Pronotum. Quadrate to slightly elongate (Text-fig. 17) ; sides subparallel or very slightly convergent to base ; basal margin usually as in Text-fig. 17 rarely with indentations ill-defined ; an ill-defined, shallow, depressed area is sometimes present towards base of disc. REVISION OF GENUS PALORUS (SENS. LAT.) 109 Elytra. Elongate, convex ; strial punctures deep ; interstices with single rows of punctures ; scutellary striole ill-defined, may be represented by 4-7 small punctures. Holotypes of cerylonotdes Pascoe (New Guinea), exilis Marseul (Japan) and praslinensis Gebien (Seychelles) are in the British Museum (Nat. Hist.), those of papuanus Kaszab (New Guinea) and zimmermani Kaszab (Upolu) are in the Hungarian Natural History Museum, Budapest—all have been examined by the author. LECTOTYPE of minor Waterhouse, present designation (sex indet.). DAMMA IsLAND bearing labels as follows: ‘‘ Damma Is. 92—20/5873 ’’, left hand specimen, B.M. (Nat. Hist.) Paralectotype of minor Waterhouse on same card mount as type, right hand specimen of the pair. Comparative notes. General facies similar to ficicola (see ficicola ‘‘ Comparative notes ’’). The small size and sub-cylindrical form combined with the shape of the pronotal basal margin (Text-fig. 17) distinguish this species. Distribution. Oriental and Ethiopian (Malagasy sub-region—Seychelles and Madagascar) illustrated in Map 2, probably of Indo-Malayan origin. The wide distribution of this species in the Oriental region may be partly due to commerce. In the Pacific Island distribution wind (typhoons etc.) may have combined with man. The islands appear to have been used as stepping-stones. I have seen single specimens, presumably representing importations, from Iran, Abadan (Corporaal Coll.) and French W. Africa, Yapo. Habitat. Occurs under bark of a wide variety of trees. In India it is most frequently recorded from sal (Shorea robusta), Blair (1930). It is not infrequently found in produce, illipe nuts, rice etc., from the Orient arriving in Great Britain. In Japan this species is sometimes found in flour mills. One specimen from Assam, identified by Blair, was collected in a cave. Palorus austrinus Champion (Text-fig. 32) Palorus austrinus Champion, 1896, Entomologist’s mon. Mag. 32 : 30. Length 2:3-2:°8mm.; breadth o-g-1-:0mm.; brown, moderately shining to dull; micro- reticulation shallow, frequently distinct. Head. Flat anteriorly (clypeus and genae) ; clypeus with puncturation slightly sparser than genae ; eyes large, not protuberant. Pronotum. Transverse, often slightly broader towards apex, a very narrow (breadth of approx. 3 punctures) longitudinal median impunctate region, from base to apex, is present but often indistinct ; apical angles obtuse, rounded, dorsally (see Text-fig. 32) and ventrally weakly produced ; apical margin almost straight ; lateral margins subparallel or slightly convergent to base, in lateral view moderately (Text-fig. 32 inset) or strongly raised from basal half or third to base (sometimes more so on one side than on the other) ; with a more or less distinct, very shallow depression on each side extending from approximately apical to basal fifth (Text-fig. 32) ; basal margin nearly as wide as base of elytra. 110 D.G. H. HALSTEAD Elytva. Usually elongate (as Text-fig. 32) ; interstices 2 and 3 with puncturation somewhat confused but approximating to single rows. LECTOTYPE, present designation (sex indet.). AUSTRALIA: Roebuck Bay, bearing labels as follows: “‘ TYPE [standard B.M. (Nat. Hist.) type label]/Roebuck Bay N.W. Australia J. J. Walker/G. C. Champion Coll. B.M. 1927-409/Palorus austrinus Ch. [Champion’s MS] ”’ left hand specimen, B.M. (Nat. Hist.). Paralectotypes : nine examples: the right hand specimen on the same card as the lectotype and eight others from Roebuck Bay and Damma Island, seven in the B.M. (Nat. Hist.) and two in the Australian Museum, Sydney. Champion (1896) gives the following localities in his description of austrinus : ‘“N.W. Australia, Roebuck Bay and Port Darwin; Damma Island.’’ I am placing the syntype material from Port Darwin (six specimens) in intermedius sp. n. Comparative notes. PP. austrinus is easily confused with female genalis if the form of the genae is overlooked and on pronotal shape some individuals resemble vatzeburgiu. The larger eyes of austrinus, however, immediately separate it from vatzeburgit. It is also close to intermedius sp. n. and beesoni—see key for separation. Distribution. Oriental. PHILIPPINES, BRUNEI and NoRTH WESTERN AUSTRALIA. Palorus upoluensis, genalis and ratzeburgii have been mis-determined as austrinus in the past® and therefore I am including full details of data on specimens (in addition to syntypes) seen by me. 2, ‘‘ Broome/N.V. Austr. Mjoberg/Museum Frey Tutzing ”’; 1, ““ Roebuck Bay/S.A. Museum specimen ’’ ; 6, “‘ Philippines, Basilan ” (Hung. Nat. Hist. Mus.) ; 1, ‘‘ Davao, Mindafiao Baker ”’ (in Smithsonian Institute, Washington) ; 1, ‘‘ P. Princesa Palawan Baker/Museum Frey, Tutzing”’; 5, ‘ Mt. Makiling, Luzon Baker” (3 in Smithsonian Institute and 2 in Museum Frey) ; 5 “‘ Los Banos, P. I. Baker ”’ (2 in Smithsonian Institute, 2 in Frey Museum and 1 in Australian National Collection, Canberra); 1, “ Borneo, Brunei’? (Hung. Nat. Hist. Mus.). Habitat. Champion (1896) said of austrinus (sens. Champion) occurred in Australia under bark and away from habitations ”’. ce ... the insect Palorus intermedius sp. n. (Text-fig. 31) Length 2-:2—2-6 mm. ; breadth 0-8-o-9 mm. ; brown, usually light, shining ; micro-reticulation shallow and indistinct. Head. Almost flat anteriorly (clypeus and genae); clypeus with puncturation slightly sparser than genal puncturation ; genae very slightly rounded, very slightly raised above level of clypeus ; eyes large, moderately protuberant. Pronotum. Transverse; apical angles obtuse, sharply defined and usually moderately produced (Text-fig. 31) (rarely, in very small specimens, they are only weakly produced), ventrally moderately produced ; apical margin slightly sinuate ; lateral margins sub-parallel 6 Blair (1935) records austrinus from the Marquesas, Hivasa; I have seen two specimens, which I believe form part of the series of five seen by Blair, and these are P. genalis Blair. A specimen labelled by Blair as austrinus from Samoa was also P. genalis Blair (see Blair, 1935). Blair, in the same publica- tion, records austrinus from the Gilbert Islands and from Dehra Dun (India) ; I have not located these specimens. I have not seen the austrinus recorded from New Guinea by Gebien (1920). The specimens recorded as austrinus from New Guinea by Dr. Kaszab (1939) were upoluensis Blair. REVISION OF GENUS PALORUS (SENS. LAT.) IT from approximately apical eighth (apical angle) to base, in lateral view straight to basal half or third then slightly raised to base (Text-fig. 31, inset) (rarely moderately raised) or straight from base to apex ; basal margin straight ; base as wide as base of elytra. Elytra. Usually comparatively short (when compared with austrinus) (see Text-figs. 31, 32) ; interstices 2 and 3 with single rows of punctures usually becoming irregular towards base. Holotype g. AUSTRALIA: Queensland, bearing labels as follows: “ Cairns distr. A. M. Lea/K 67015 ’’, in the Australian Museum, Sydney. Paratypes: fifty-five examples; AUSTRALIA: Northern Territory, Queensland and N.S.W., bearing labels as follows: 1, ‘‘ Cairns distr. A. M. Lea/Attracted to light/K 67015 ’’, (in the Australian Museum, Sydney) ; 3, “ Cairns distr. A. M. Lea ”’ (2, B.M. (Nat. Hist.) and 1, National Collection, Canberra) ; 1, ‘‘ Cairns E. Allen ”’ (National Collection, Canberra); 1, “Cairns distr. J. A. Anderson/Queensland Museum ”’ Queensland Museum type No. T6352; 1, ‘‘ Cairns Hacker/S.A. Museum specimen”’; 1, ‘‘ Clermont vii.28, Queensland Dr. K. K. Spense”’ (Australian Museum, Sydney) ; 1, “ Yeppoon Q. H.J.C. x/24”’; 3 (on same card), “‘ Kuranda N. Q. Hacker”; 1, ‘‘ Qloomba, Queensland F. H. Saylor”’ (this and the previous Imm Fics. 30-32. 30, Palorus acutangulus sp. n. 31, Palorus intermedius sp. n. Inset lateral view of pronotal side margin. 32, Palorus austrinus Champion. Inset lateral view of pronotal side margin, T12 D.G. H. HALSTEAD 4 in the National Collection, Canberra) ; 1, ““ Queensland Rockhampton/Museum Frey, Tutzing’’; 1, “ Queensland P. P. Dodd 1904—27/Townsville’”’ (B.M. (Nat. Hist.)) ; 1, ‘‘ Gladstone Q, Lea/S.A. Museum specimen’”’; 2 (same card) “ Port Darwin, N.W. Australia J. J. Walker/Palorus austrinus Ch [Champion’s MS] ” (= syntypes of austrinus Champion); 2 (same card) “ Port Darwin 92-2/4734” (= syntypes of austrinus Champion) ; ibid. “ /4735’”’ (= syntypes of austrinus Champion); 1, “ Adelaide River 91-49/979’’; 2 (same card) “ Adelaide River g2-20/5605’; 1, ““ Adelaide River N.W. Australia J. J. Walker/Palorus austrinus Ch. [Champion’s MS]”’ (this and the previous nine specimens in the B.M. (Nat. Hist.)) ; 1, ““ Adelaide River N.W. Australia J. J. Walker/S.A. Museum specimen ”’ ; 8 (5 specimens on one pin and 3 on one pin), “‘ Groote Eylandt A. H. Elston Collec- tion’ (in the Australian Museum, Sydney) ; 2 (both on same pin) “ Melville I. W. D. Dodd/S.A. Museum specimen ’”’; 6 (2 on one pin, 4 on another), “‘ Milingimbi, Crocodile Is. N. Austr. C. Barrett/F. E. Wilson Collection/National Museum of Victoria, Melbourne’”’; I, “ Richmond R. N.S. Wales 1909-174”’ (in B.M. (Nat. Hist.)). Comparative notes. This species is very close to austrinus. It may be separated from this species by characters of the pronotum (see key) and is usually shorter than austrinus due to comparatively shorter elytra. In addition certain individuals resemble upoluensis but the pronotum is more parallel sided and its basal rim is not as strongly expanded in the middle as in upoluensis. Distribution. Australian. NORTHERN TERRITORY to NEW SouTH WALES—as above. Palorus reticulatus sp. n. (Text-fig. 33) 9 (g unknown). Length 2-5 mm.; breadth 0-9 mm.; brown, dull; micro-reticulation deep, uniform and distinct over dorsal surface (see Text-fig. 33). Head. Flat anteriorly ; moderately densely punctured, punctures with fine setae; clypeus slightly raised medially ; genae slightly lower than top of clypeus ; eyes comparatively large, not prominent ; supra-orbital carinae distinct. Pronotum. Moderately transverse, slightly cordiform, puncturation as head ; apical margin, excluding apical angles, almost straight ; apical angles sharply defined ; lateral margin gradually rounded and convergent to base ; basal margin almost straight. Elytra. Scutellary striole not differentiated ; strial punctures with fine setae—as pronotal punctures (setae not illustrated) ; interstices with single rows of punctures. Holotype 9. AUSTRALIA: Queensland, bearing labels as follows: ‘‘ Clermont ix 29 Queensland Dr. K. K. Spense/K. K. Spense Collection/Palorus sp. prob. new A. Neboiss, 1959 ”’ in the collection of the Australian Museum, Sydney. Comparative notes. The uniform distinct micro-reticulation and the “ flat ”’ head combined with the pronotal form distinguish this species, REVISION OF GENUS PALORUS (SENS. LAT.) 33 Fic. 33. Palorus reticulatus sp.n. Inset small areas of pronotum and elytron to show micro-reticulation. 113 114 DD. GLB HALSTEAD Palorus hypophloeoides Blair (Text-figs. 34a, b) Palorus (Stenopalorus) hypophloeoides Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 136. Length 2-1-3:2 mm. ; breadth 0-5—o-8 mm. ; very elongate and cylindrical ; brown, pronotum shining, elytra somewhat dull; micro-reticulation very shallow and ill-defined, absent from some regions of pronotum. Head. Moderately densely punctured, supra-orbital carinae well developed ; ¢ with a pair of prominent frontal tubercles and small projections of genae at clypeo-genal sutures (Text-fig. 34a) ; Q (only one specimen seen) with frontal tubercles represented by two very weakly raised areas (Text-fig. 34b). Pronotum. Elongate, disc slightly depressed ; punctures moderately deep, separated by 2-3 diameters ; basal margin straight for approximately median third, then angled to sides, 0-5mm 34 Fic. 34. Palorus hypophloeoides Blair, (a) g; (b) 2 head. REVISION OF GENUS PALORUS (SENS. LAT.) 115 Elytra. With deep strial punctures ; scutellary striole of 4 to 5 punctures—often becoming very shallow at apex ; interstitial puncturation approximating to single rows. LECTOTYPE, present designation, gj. INDIA: Dehra Dun, bearing labels as follows: ‘‘ Type [standard B.M. (Nat. Hist.) type label]/Cantt Road Dehra Dun C. F. C. Beeson 8.x.1926/R.R.D. 499 B.C.R. 163 Cage 290/ex Dalbergia Sissoo/489/ hypophloeoides [MS] ’’, B.M. (Nat. Hist.). Blair (1930) gives two records for the type material (i.e. at least two specimens). In the British Museum (Nat. Hist.) there is only the above specimen with type locality data. Comparative notes. This species is quite distinct from all other Palorus on external dorsal morphology and for this reason Blair placed it in the subgenus Stenopalorus. I have not chosen to place it in a distinct genus and have not used subgenera. P. hypophloeoides has normal genitalia and a series of deep punctures on the 2nd sternite in the male—typical of Palorus. Distribution. Oriental. In addition to the lectotype, from INDIA: Dehra Dun, I have seen specimens from SINGAPORE and INDIA: Kumaun with the following data, 3 J, 1, “ Bi 6/22 [1, 9/22 and 1, 2/22]/Singapore C. J. Saunders B.M. 1933-227 ”’ ; 1, 2, “ R. Sarda Gorge Kumaon U.P. Dec. 1918 H.G.C./2683 ” (B.M. (Nat. Hist.)). Habitat. According to Blair, from Dalbergia sissoo and Pinus longtfolia (specimen from Pinus not seen by me). Palorus sinuaticollis Blair (Text-figs. 35a, b) Palorus sinuaticollis Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 137. Palorus fuhoshoanus Kaszab, 1941, Stettin. ent. Zig 102 : 56, syn. n. Length 3-:3—-3:8 mm. ; breadth 1-3-1-5 mm. ; large robust species ; red-brown to dark brown, dull; micro-reticulation deep and distinct. Head. Anteriorly emarginate ; eyes large and prominent, dorsal length greater than breadth of clypeus. In $ (Text-fig. 35a) with deep median depression ; punctures generally separated by slightly more than one diameter; clypeus ill-defined, clypeal area sparsely punctured and shining ; genae strongly raised above level of clypeus and eye (form characteristic see Text-fig. 35a). Only one large specimen seen, small males probably approach the female in development of genae. In 9 (Text-fig. 35b) with a shallow ill-defined median depression; punctures on vertex separated by one diameter or less; clypeus sparsely punctured and moderately shining in the middle ; genae raised above level of clypeus and moderately prominent. Pronotum. Transverse; moderately densely punctured, punctures on disc fine (punctures without long setae); apical margin strongly sinuate; apical angles not sharply defined ; lateral margin slightly rounded from base to apex ; basal margin almost straight. Elytra. Scutellary striole of 5 or 6 punctures, more or less distinct ; interstices with punctures approximating to two rows, three rows in some areas. LECTOTYPE, present designation, 9. Burma: Pyinmana, bearing labels as follows : ‘“‘ Type [standard B.M. (Nat. Hist.) type label]/For. Zool. Coll. Yananngmyin R. Pyinmana, Burma 17.iv.191g9 C. F. C. Beeson/Tectona grandis/Ex. Coll. Dehra 116 D.G. H.. HALSTEAD Dun B.M. 1924-219 256/P. sinuaticollis Blr Type det K. G. Blair”, B.M. (Nat. Hist.). Paralectotype 9. ToNKIN: Hoabinh, as recorded by Blair (1930), B.M. (Nat. Hist.). The specimen recorded by Blair from Inthabaing Reserve, Insein has not been found. Holotype of fuhoshoanus Kaszab is in the Deutsches Entomologisches Institut, Berlin. Comparative notes. The form of the head and pronotum, combined with size distinguish this species. Distribution. Oriental. INDIA, VIETNAM, FoRMOSA. In addition to the types (from India and Formosa (fuhoshoanus)) I have seen the following : 1, 9, ““ Formosa Polisha ”’ (in the Hung. Nat. Hist. Mus.) ; 1, 3, (specimen figured) ‘“‘Hoah Binh Tonkin 12.1934 A de Cooman/Museum Frey Tutzing”’. Habitat. Recorded from Tectona grandis and Mangifera sp. by Blair (1930). Palorus shoreae Blair Palorus shoreae Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 138. Length 3:1-3-4mm.; breadth 1-:2-1-3mm.; large, moderately elongate species similar to beesoni and kaszabi sp. n.; red-brown to dark brown, somewhat dull; micro-reticulation very deep and distinct to shallow and indistinct. Head. Moderately densely punctured, punctures bearing short indistinct setae; clypeus slightly less densely punctured than genae; eyes large, dorsal length equal to breadth of clypeus ; male with genae rounded, slightly to distinctly raised above level of clypeus and slightly prominent anteriorly (similar to angular form of beesoni Text-fig. 37b) ; female with head anteriorly almost flat or genae very slightly raised above level of clypeus, genae rounded, not prominent anteriorly. Pronotum. ‘Transverse; moderately densely punctured, punctures bearing long fine setae, equal in length to two diameters of largest punctures; apical margin straight to slightly sinuate ; apical angles more or less obtuse ; lateral margin subparallel for apical half or two thirds then slightly rounded and convergent to base ; basal margin almost straight ; sides with very shallow, diffuse depression towards base. Elytra. Somewhat elongate, equal to about 2-25 times the pronotal length; strial and interstitial punctures with setae, not as long as pronotal setae; scutellary striole indistinct, represented by two or three shallow, ill-defined punctures, sometimes absent on one or both elytra; interstitial puncturation variable and confused, third interstice with punctures approximating to two rows. LECTOTYPE, present designation (sex indet.). InpDIA: Manipur, bearing labels as follows: ‘‘ Type [standard B.M. (Nat. Hist.) type label]/Shugnu 3000’ Manipur S. N. Chatterjee 30.iii.1924/Under bark/497/shoreae/Palorus shoreae Blr Type det K. G. Blair’, B.M. (Nat. Hist.). Paralectotypes : nine examples, two in the Hungarian Natural History Museum, Budapest and seven in the British Museum (Nat. Hist.), locality data as in Blair (1930). REVISION OF GENUS PALORUS (SENS.LAT:) 1% Comparative notes. P. shoreae has the general facies of kaszabi sp. n. and beesont and is also somewhat similar to genalis; it is distinguished in the key. Distribution. Oriental. INDIA and VIETNAM. In addition to the types from India I have seen specimens labelled, ‘“‘ Anamalai Hills Cinchona S. India” (gf, 9—g in Frey Mus. @ in Ardoin Coll.) ; ‘ Tonkin Hoabinh Aug rg18 R. V. de Salvaza/Indo. China R. V. de Salvaza rg18. 1”’ (in B.M. (Nat. Hist.)). Palorus kaszabi sp. n. (Text-fig. 36) Length 2:3mm.; breadth 1:1mm.; dark brown, somewhat dull; micro-reticulation distinct (not illustrated). Head. Puncturation moderately dense, coarse on vertex, punctures bearing fine indistinct setae; clypeus slightly raised in the middle, puncturation slightly less dense than that of genae ; genae slightly raised, rounded, not prominent ; eyes large. 36 35 Figs. 35, 36. 35, Palorus sinuaticollis Blair, (a) 3; (b) 9, head and pronotal apex. 36, Palorus kaszabi sp.n. Inset setation of a small region of elytron and pronotum. 118 D.G. H. HALSTEAD Pronotum. Moderately densely punctured, punctures bearing long distinct setae (Text-fig. 36, inset); apical margin sinuate; apical angles sharply defined, slightly more than right angles ; lateral margin slightly rounded dorsally, in lateral view raised to base from approxi- mately basal half; shallow longitudinal depressions at sides of disc; basal margin slightly sinuate due to expansion of basal rim medially (Text-fig. 36). Elyiva. Strial and interstitial punctures with long distinct setae (Text-fig. 36, inset) ; puncturation of interstices somewhat confused but approximating to two or three rows. Holotype 3. PHILIPPINES: Binalnan, bearing label as follows: ‘“ Philippines, Binalnan ”’ Paratype, g, with the same data as the holotype, both specimens in the Hung. Nat. His. Mus. Comparative notes. This species is readily distinguished by the very long fine setae on pronotum and elytra and is separated from the closely related shoreae by pronotal form etc.—as in key. Species named after Dr. Z. Kaszab, Hungarian Natural History Museum, Budapest. Palorus genalis Blair (Text-figs. 38a—d) Palorus genalis Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 140. Palorus saipanensis Kulzer, 1957, Insects Micronesia 17 (3) : 220, syn. n. Length 2:1-2-6mm.; breadth 0-7-0-9mm.; brown, moderately shining to dull; micro- reticulation strong and distinct. Head. Fairly densely punctured ; eyes appearing comparatively flat (when compared with beesoni etc.) dorsal length more than twice dorsal breadth ; maximum expanse of gena equal to (9) or greater than ($) maximum dorsal breadth of eye ; clypeus, and a narrow region at base of genae, sparsely punctured and shining. $ with genae projecting beyond clypeus, produced medially to form a triangular projection, always distinct but developed to a varying degree dependent on absolute size of individual (see Text-figs. 38a and b); vertex high, frons forming an almost vertical somewhat triangular region, moderately densely punctured as the rest of the head. Q with genae very slightly prominent anteriorly, distinctly raised above level of clypeus, margin thickened and forming a low arcuate ridge (Text-fig. 38d) ; vertex not as high as in the male and frons not somewhat triangular. Pronotum. More or less transverse (Text-figs. 38a, 38d), punctures bearing fine short setae, often absent (rubbed off?) on disc; sides subparallel to slightly rounded (Text-fig. 38d) in female; very slightly rounded (Text-fig. 38c), or distinctly so in large individuals (Text-fig. 38a), in males. Elytva. Strial punctures with short fine setae, usually discernible but not very distinct ; interstitial puncturation variable and confused, usually approximating to two rows, at least for part of length of interstices 2-4. Aedeagus as in Text-fig. 38e. LECTOTYPE, present designation, J. CEYLON: Peradeniya, bearing labels as follows: “Type [standard B.M. (Nat. Hist.) type label]/Ceylon E. E. Green 1900-239/From dry paddy Peradeniya [MS]/Palorus genalis Blr. det. K. G. Blair T’’, B.M. (Nat. Hist.). Paralectotypes : three examples, data as the lectotype. REVISION OF GENUS PALORUS (SENS. LAT.) 119 Blair (1930) includes Ceylon, (E. E. Green), North Andaman (B. M. Bhatia) and Philippine Islands (Semper) in his type localities for this species. I have not been able to find the specimen from N. Andaman and there is a very long series of genalis from the Philippines (Semper) whereas Blair said ‘“‘ The short series from the Philippine Islands...’’ I am therefore considering only the Ceylon material, the above four specimens, as syntypes. Holotype of satpanensis Kulzer is in the Chicago Natural History Museum. Comparative notes. The triangular form of the genae readily distinguishes the male from that of other species. The female is somewhat similar to both sexes of beesont (normal form) and austrinus but may be separated on genal form as in the key. Also in both sexes of genalis the elytra are slightly more acuminate than in beesont and austrinus. Distribution. Oriental (also Africa and W. Indies, apparently imported). The type locality for satpanensis Kulzer is Mariana Islands. Specimens have been seen from the following localities: West InpiEs, Trinidad (on Paddy var sughandi, 1958) and Guadeloupe (old specimens 1900?) ; Americas, BRITISH HONDURAS, Rio Temas, 1937; Africa, KENYA, Msambareni (1952 on Sorghum) and Lindi (ex Cassava), WEsT Africa, N’Zérékoré, 1951 ; CELEBES, (collected by Wallace, 1866) ; S1AM; Mataya (Pahang, in stored rice in rice mill 1939) ; MARQUESAS ISLANDs, Atuoua Hiva Oa, 1929. Habitat. The types (genalis) were collected on dry paddy (Ceylon) and the specimen from N. Andaman (not seen) on Myristica andamanica. It is frequently associated with stored products and has been imported into Great Britain in small numbers on cassava root (loaded Zanzibar), cattle food beans (from Mombasa) (see also Kenya records above), illipe nuts and sago flour (from Singapore), tapioca root (from Java), gaplek root (from Borneo), rice and groundnut cake (from Burma) and ginger (from W. Indies). Specimens from Saipan and Guam (satpanensis paratypes) were collected at light. Palorus beesoni Blair (Text-figs. 37a-c) Palorus beesoni Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 140. Length 2-2~-2-9 mm. ; breadth o-8-1-1 mm. ; facies similar to ratzeburgit ; brown, often dark, moderately shining, rarely dull; micro-reticulation usually distinct but shallow. Head. Clypeus with puncturation somewhat sparser than genal puncturation; genae of female and non-angular male form (Text-fig. 37a) almost straight to slightly rounded, usually slightly raised above level of clypeus but sometimes level with clypeus ; angular form male with genae sometimes produced anteriorly forming an angle near clypeo-genal suture (as in Text-fig. 37b); this form grades into non-angular form; maximum expanse of gena, at middle, approxi- mately two thirds or less of maximum dorsal breadth of eye ; vertex not as high as in genalis, frons not somewhat triangular; eye slightly variable in size, large, dorsal length normally approximately twice dorsal breadth (usually more prominent than in genalis). Pronotum. More or less transverse (sometimes nearly quadrate) (Text-fig. 37a), densely and coarsely punctured laterally, punctures bearing fine setae, usually distinct ; apical margin 120 D.G. H. HALSTEAD slightly sinuate ; apical angles moderately sharply defined ; lateral margins, in dorsal view, from approximately basal third slightly convergent to base, in lateral view, raised from basal half or third to base, basal margin slightly narrower than elytral base. Elytra. Strial punctures moderately deep; interstitial puncturation confused, approxi- mating to one or two rows in interstices 2 and 3. Aedeagus as in Text-fig. 37¢. 38 e Scale lines:0-5mm 8 Sd Fics. 37, 38. 37, Palorus beesoni Blair, (a) head and pronotum of normal ¢ (also typical of 9); (b) head of g with angular genae; (c) aedeagus. 38, Palorus genalis Blair, (a) pronotum and head of $ with strongly developed genae; (b) moderately developed 3d gena (outline) ; (c) straight form of pronotal side; (d) 9 head and pronotum ; (e) aedeagus. REVISION OF GENUS PALORUS (SENS. LAT.) 121 LECTOTYPE, present designation (sex indet.). INpDIA: Singhbhum, bearing labels as follows: ‘‘ Type [standard B.M. (Nat. Hist.) type label]/Singhbhum, Bihar & Orissa C. F. C. Beeson 9.i.1921/Ex Ficus religiosa/34/Ex Coll. Dehra Dun B.M. 1924-219/Palorus beesoni Blr. T. det K. G. Blair’, B.M. (Nat. Hist.). Paralectotypes: fourteen examples, data as in Blair (1930), thirteen in B.M. (Nat. Hist.) and one in Hung. Nat. Hist. Mus. Specimens from the localities “ Patri...’ and ‘“‘ North Sinhawa range... (localities given in Blair’s original description) have not been found. a”) ” Comparative notes. This species is similar to female genalis and austrinus and apart from eye size could be confused with vatzeburgi. From genalis females it may be distinguished on genal form (Text-figs. 37a and 38d) and from austrinus on pronotal form (more convergent to base in beesont). The paramere tube is distinct from that of genalis (see Text-figs. 37c, 38e). Distribution. Oriental. Blair (1930) records this species from INDIA and BuRMA ; in addition I have seen specimens from CEYLON, CELEBES, JAVA, N. VIETNAM (Hoah-Binh) and CuHinA (Kwangtung). Habitat. Under bark of various trees attacked by bark-beetles or sapwood- borers (Blair, 1930). Blair records the following genera of trees: Ficus, Butea, Boswellia and Mangifera. One specimen seen was collected at light. Palorus auranteus sp. n. (Text-fig. 40) 3 (2 unknown). Length 2-4mm.; breadth 0-9 mm. ; orange-brown, pronotum moderately shining, elytra dull; micro-reticulation shallow, indistinct on pronotum, distinct on elytra. Head. Moderately densely punctured, punctures bearing fine distinct setae; clypeus flat, sparsely punctured and shining ; genae raised above level of clypeus, densely punctured, round and prominent (Text-fig. 40); eyes large, protuberant, with large facets; antennae long (slightly longer than in beesont) ; vertex high. Pronotum. Transverse; moderately densely punctured, at sides punctures separated by one diameter, punctures bearing fine setae; apical margin almost straight; apical angles not prominent, obtuse, not very sharply defined ; lateral margin in dorsal view rounded from base to apex, in lateral view arcuate; with very shallow, ill-defined, longitudinal depressions at sides of disc (Text-fig. 40). Elytra. Scutellary striole of 4 (or 5?) ill-defined, shallow punctures ; interstitial punctures approximating to two rows in interstices 2 and 3. Holotype, g. MALaya: Fraser’s Hill, bearing labels as follows: “‘ Gap (Fraser’s Hill) Malay Peninsula A. M. Lea & Wife/S.A[ustralian] Museum specimen ”’ (on each elytron there is the remains of a uropodid mite (Acarina) stalk). Comparative notes. This species (J) is very close to beesont and has the general facies of genalis. It may readily be distinguished by the rounded, moderately prominent genae, the large, coarsely faceted eyes, the long antennae and the pronotal shape (Text-fig. 40). 122 D.G. H. HALSTEAD Palorus tenuipunctatus Blair Palorus tenuipunctatus Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 140. Length 2:8-3:0mm.; breadth 1-Imm.; rather convex transversely ; yellowish brown ; micro-reticulation deep and dense (Text-fig. 41) giving an opaque dull appearance. Head. Moderately densely punctured, punctures separated by a diameter or less and bearing distinct short setae (exceeding the longitudinal length of a puncture) ; eyes large and close to front of head, dorsal length greater than genal length; clypeus and genae with puncturation of equal density ; clypeus wide (equal to about 1-2 times the genal length) ; genae small, not distinctly raised or prominent. Pronotum. Transverse, moderately densely punctured, punctures fine on disc, separated by one or two diameters, becoming coarser towards sides, bearing fine, moderately long, distinct setae ; apical margin slightly sinuate ; apical angles obtuse, sharply defined ; lateral margins almost parallel for middle half then slightly rounded to base and apex ; basal margin slightly rounded. Elytra (Text-fig. 41). Scutellary striole not differentiated ; strial and interstitial punctures with short setae (very distinct in some lights—not illustrated in Text-fig. 41); interstitial puncturation confused but tending to form three or four rows. LECTOTYPE, present designation (sex indet.). BurMA: Inthabaing Reserve, bearing labels as follows: “‘ Type [standard B.M. (Nat. Hist.) type label]/Mangifera sp/Inthabaing Res. Insein, Burma. D. J. Atkinson 31.xii.1926/570/Palorus tenuipunctatus Type Blr. det. K. G. Blair ’’, B.M. (Nat. Hist.). Although Blair records two examples, both from the same locality, there is only the above specimen in the British Museum (Nat. Hist.). Comparative notes. The puncturation of the elytra, the micro-reticulation and the narrow genae, combined with size, distinguish this species. Distribution. Oriental. In addition to the lectotype (BURMA) there is a specimen in the British Museum from MIDDLE ANDAMAN with the following data: ‘‘ Middle Andaman, B.M. Bhatia 6.x11.1928/ex Dipterocarpus turbinatus/1177 ”’. Palorus longifoliae Blair (Text-figs. 39a, b) Palorus longifoliae Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 138. Length 3:3-36 mm. ; breadth 1-1-1:2 mm.; very large species, red-brown, shining ; micro- reticulation more distinct on elytra than on pronotum. Head. Eyes large and prominent; clypeus shining, slightly less densely punctured than genae ; genae of male raised above level of clypeus, angular (Text-fig. 39a), of female not angular, straight to slightly rounded (Text-fig. 39b). Pronotum (Text-fig. 39a). Slightly transverse ; disc moderately depressed ; lateral margins almost straight and very slightly convergent to base; apical margins straight; basal margin straight for medial two thirds, slightly raised at sides. Elytra. Elongate; strial punctures deep; scutellary striole of 6-7 comparatively shallow but moderately distinct punctures ; interstitial puncturation approximating to one or two rows. LECTOTYPE, present designation, 9. N. Inp1a: Trisula, bearing labels as follows: “‘ Type [standard B.M. (Nat. Hist.) type label]/ex Pinus longifolia/Ex REVISION OF GENUS PALORUS (SENS. LAT.) 123 Coll. Dehra Dun 1924—219/For. Zool. Col. Trisula Garhwal U.P. 27.i.1919 A. E. Osmaston ’’, B.M. (Nat. Hist.). Paralectotypes: thirteen examples, data as in Blair (1930) (two from Trisula and eleven from W. Almora) B.M. (Nat. Hist.) and one W. Almora specimen in PAL. Coll, Comparative notes. This large species is readily distinguished by the pronotal é . Age F © SY Fics. 39-41. 39, Palorus longifoliae Blair, (a) g¢ head and pronotum; (b) 9 head. 40, Palorus auranteus sp. n., head and pronotum. 41, Palorus tenuipunctatus Blair, small area of pronotum and elytron to show puncturation and micro-reticulation. ENTOM. IQ, 2 9 DOG 124 D. G. H. HALSTEAD form (Text-fig. 39a) and comparatively distinct scutellary striole of 6-7 punctures. It is probably most closely related to beesont. Distribution. N. INDIA. Habitat. From Pinus longifolia (types). Palorus andrewesi Blair (Text-fig. 42) Palorus andrewesi Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 139. Length 2:0-2:°8mm.; breadth o-9g-1:°0mm.; brown, bicoloured—pronotum much darker than elytra, moderately to strongly shining ; micro-reticulation variable, moderately deep and distinct or shallow and more or less distinct. Head. Moderately densely punctured, punctures separated by 1-2 diameters ; clypeus much lower than genae, sparsely punctured, shining ; eyes large and prominent; genae of g angular (Text-fig. 42a), degree of development dependent on absolute size, distinctly raised above level of clypeus at an angle of approximately 45° to it; genae of 9 not angular, raised above level of clypeus, rounded (Text-fig. 42b). Pronotum. ‘Transverse, punctures of disc separated by 2—3 diameters, puncturation coarser and denser towards sides; apical margin sinuate ; apical angles obtuse but sharply defined ; lateral margin slightly rounded from base to apex; basal margin almost straight, slightly expanded in the middle. Elytra. Comparatively short and broad, elytral length : pronotal length, 2-2-2 : 1 ; scutellary striole of 3-5 small shallow punctures. Holotype in the British Museum (Nat. Hist.). Comparative notes. Somewhat similar to upoluensis but with pronotum not as convergent (Text-fig. 11); also andrewesi is bicoloured with elytra shorter, 2-2°2: 1 (2:2-2:3: 1 in upoluensis). In addition, the genae, which are angular in the ¢ and with a more strongly raised margin in the 9, will separate this species from upoluensis. Distribution. Oriental. In addition to the Holotype from INnp1A I have seen specimens from SABAH and SINGAPORE with the following data: 1 ¢ “ Sandakan Borneo Baker/289/Museum Frey Tutzing’”’; 2 j and 1 9 with the same data in the British Museum (Nat. Hist.) “‘ Singapore C. J. Saunders B.M. 1933-27/Scotland 15.10.22 dry bark [Saunders’ MS]”’; 1 ¢ in British Museum “ Singapore C. J. Saunders B.M. 1933-227/04 Orchd. Rd. 9.22 [Saunder’s MS].”’ PSEUDEBA Blackburn, 1903 gen. rev. Pseudeba Blackburn, 1903, Tvans. R. Soc. S. Aust. 27: 119. Type-species : Pseudeba novica Blackburn (by monotypy). (Originally described in the Colydiidae, its true position in the Tenebrionidae was recognised by Carter and Zeck (1937) who synonymized Pseudeba novica Blackburn with Palorus eutermiphilus Lea). Length 2-3-2-°8mm.; body moderately depressed; red-brown, moderately dull; cuticle micro-reticulate. REVISION OF GENUS PALORUS (SENS. LAT.) 125 Head. Flat anteriorly, moderately densely punctured ; clypeus almost flat to slightly raised in the middle, well- or ill-defined ; genae flat, very slightly raised above antennal insertions, continuous with side margin of head; side margin of head higher than dorsal surface of eye (supra-orbital carina absent) ; eyes not emarginate, small or very small, just below or distinctly below dorsal surface of head, postero-laterally limited by small projections of the head. Antennae 11-segmented, inserted beneath genae, similar to Palorus but short in fossor sp. n. and thickened, with shallow sulci, in movica ; mouth parts as in Palorus. Thorax. Pronotum transverse, widest towards apex, sides moderately or strongly rounded to base, margined basally, laterally (may be very narrowly) and apically on apical angles or, Imm A2 Fic. 42. Palorus andrewesi Blair, (a) 3; (b) 2 head and pronotum. 126 DD: G. Ho HALSTEAD apically, margin may extend to median half; sides may have a narrow, somewhat lenticular, almost vertical region as in Palorus; puncturation moderately dense, may be very dense laterally ; punctures oval, slightly reniform or may be confluent. Scutellum transverse, as in Palorus or more rounded (Text-fig. 45—mnovica). Sterna similar to Palorus, prosternum highly modified in novica, being contracted for reception of head (Text-fig. 45e). Metendosternite as in Palorus. Protibiae similar to Palorus but in fossor sp. n. broad and with elongate scaliform spinules dorsally on external margin (spinule form similar to that found in Palorinus and Coelopalorus (Text-fig. 53b)); tarsal formula 5-5-4 in puncticollis sp. n. and fossor sp. n., 4-4-4 in novica; apical tarsal segment long, basal segment (except in novica) very small. Elytva. Free, covering abdomen, humeral angle moderately developed ; scutellary strioles not differentiated ; each elytron bearing 10 single rows of strial punctures (concealed by carinae in novica) ; interstices with single rows of small or large punctures ; interstices slightly raised for whole length or only slightly raised towards the apex but carinate for the greater part of their length in novica ; epipleura as in Palorus. Wings well developed, venation reduced (Text-fig. 44c), similar to Palorus. Abdomen. Sternites, 5 visible, second of g with deep internal pits on disc (absent in novica), of 2 without deep internal pits. Genitalia. 3g aedeagus with basal piece short (Text-figs. 43b, 44b, 45c) and pleurites of 9th sternite similar to Palorus ; 2 (novica) styli sclerotized and similar to Palorus. KEY TO SPECIES oF PSE UDEBA 1 Elytral interstices strongly carinate as in Text-fig. 45a ; tarsi 4-4-4 novica Blackburn (p. 127) — Elytral interstices not strongly carinate ; tarsi 5—5—4 : : ; , ; 2 2 Front tibiae broadly expanded ; eyes very small ; pronotum as in Text-fig. 44a, with oval to reniform punctures, puncturation not very dense at sides fossor sp. n. (p. 126) — Front tibiae not broadly expanded ; eyes somewhat larger ; pronotum as in Text-fig. 43a, without reniform punctures, puncturation dense at sides where punctures become longitudinally confluent . . . puncticollis sp. n. (p. 129) Pseudeba fossor sp. n. (Text-figs. 44a-c) 3d (2 unknown). Length 2-3mm.; breadth o-9mm.; yellow-brown, moderately dull ; micro-reticulation shallow, distinct. Head. Almost flat, emarginate anteriorly (apparently normally so), moderately densely punctured, punctures oval; clypeus ill-defined, clypeal area slightly raised medially; genae flat, ill-defined, continuous with hind side margin of head, produced anteriorly at junction with clypeus, (producing the emargination) ; eyes protuberant, very small, lateral, separated from dorsal surface by approximately half dorsal length, not rounded to base; antennae short, apical segments somewhat flattened and slightly more expanded than usual. Pronotum. Moderately transverse ; puncturation as in Text-fig. 44, with a narrow median impunctate region extending from base to apex, punctures oval to slightly reniform, bearing short indistinct setae ; apical margin almost straight ; apical angles distinctly acute; lateral margins subparallel from apical sixth to basal third, then convergent to base; basal margin slightly rounded. Elytva. Strial punctures comparatively large, round and shallow ; scutellary striole absent ; interstices with large punctures (diameter equal to two thirds that of strial punctures) bearing short setae ; interstices slightly raised (as indicated by dotted lines in Text-fig. 44). Protibiae strongly dilated and bearing indistinct, elongate, scaliform spinules dorsally on external margin, the row of spinules curving inwards at apex. Wing and aedeagus as in Text-fig. 44. REVISIONCOE GEN US PALORUS (SENS: LAT.) E27 Holotype 3 (dissected). N. WESTERN AUSTRALIA: Derby, bearing labels as follows: ‘‘ Derby N.W.A., W. D. Dodd/S.A. Museum specimen ’’, in the South Australian Museum, Adelaide. Comparative notes. This species has slightly raised interstices, showing a tendency towards the carinate interstices of novica. The general facies is distinctive. Distribution. AUSTRALIA—as above. Habitat. Unknown but the form of the antenna and the general morphology suggest a myrmecophilous association. Pseudeba novica Blackburn (Text figs. 45a-f) Pseudeba novica Blackburn, 1903, Trans. R. Soc. S. Aust. 27 : 120. Palorus eutermiphilus Lea, 1921, Mem. Qd Mus. 7 (3) : 216. (Carter and Zeck (1937) synonymized novica with eutermiphilus but Gebien (1940) did not include the species eutermiphilus (or this synonymy) in his catalogue). Length (head retracted as in Text-fig. 45a) 2-4—2-6 mm. ; breadth of elytral base 0-9-1-1 mm. ; 43a Fics. 43, 44. 43; Pseudeba puncticollis sp. n., (a) 3; (b) aedeagus, dorsal. 44, Pseudeba fossor sp. n., (a) 3; (b) aedeagus, dorsal; (c) wing. 128 D. GY Hs HALSTHAD Head. Flat, normally? (at least in museum specimens) retracted into pronotum ; punctura- tion of vertex moderately dense, becoming sparse towards clypeus, with small impunctate region above antennal insertions; punctures oval and bearing short setae; clypeus well- defined, almost flat ; genae very slightly raised above antennal insertions, continuous posteriorly with side margin of head; eyes not prominent, on side of head just below dorsal surface ; antennae with shallow longitudinal grooves. Pronotum. Moderately transverse ; moderately densely punctured, punctures with short, indistinct setae; sides distinctly sinuate to greater or lesser extent; apical margin almost straight ; apical angles not prominent; basal margin distinctly narrower than base of elytra ; two very shallow depressions often present on either side at base (see Text-fig. 45a) ; prosternum very narrow, modified to receive head (Text-fig. 45e). Elytra. Often slightly sinuate laterally ; interstices strongly carinate, each bearing single row of fine punctures (Text-fig. 45f) ; strial punctures not apparent, replaced by striae which are present between carinae extending from base to approximately apical eighth. Tibiae and tarsi with shallow longitudinal grooves ; tarsal formula 4-4-4. Genitalia as in Text-figs. 45b, c. Fic. 45. Pseudeba novica Blackburn, (a) adult ; (b) stylus; (c) aedeagus, dorsal; (d) head ; (e) pronotum, ventral; (f) part of elytron. REVISION OF GENUS PALORUS (SENS. LAT.) 129 LECTOTYPE, present designation (sex indet.). AUSTRALIA: Townsville, bearing labels as follows: ‘‘T 7253 Townsv [MS, on card mount]/Type HT. {standard B.M. (Nat. Hist.) type label]/Australia Blackburn Coll. B.M. rgro-236/ Pseudeba novica Blackb ’’, B.M. (Nat. Hist.). Comparative notes. This species is a highly specialized termitophile and is readily distinguished by the carinate elytral interstices. The 4-4-4 tarsi are rare (if not unique) in the Tenebrionidae. Distribution. AUusTRALIA—Queensland, Townsville and Morven. Habitat. In nests of Nasutitermes exitiosus (Hill) (termite kindly determined for me by Mr. W. A. Sands of the British Museum) in Townsville and of “ Eutermes magnus ’’ [= Nasutitermes magnus (Froggatt)] in Morven (the latter record according to labels on specimens in the Australian Museum, Sydney). Pseudeba puncticollis sp. n. (Text-figs. 43a, b) 3 (Q unknown). Length 2:83mm.; breadth 1-2mm.; red-brown, somewhat dull; micro- reticulation shallow but distinct. Head. Almost flat anteriorly ; moderately densely punctured, punctures large and decreasing in size towards genae and clypeus, bearing short setae; clypeus large, slightly raised in the middle, slightly emarginate anteriorly, puncturation sparse ; genae lower than top of clypeus, continuous posteriorly with side margin of head and with an impunctate region above insertion of antennae, limits of clypeus/frons not defined ; eyes small, on side of head, separated from dorsal surface by approximately one third of dorsal length of eye. Pronotum. Moderately transverse; punctures deep, becoming large, oval and longitudinally confluent towards the sides, bearing short indistinct setae ; pronotum with very narrow median impunctate region from base almost to apex; apical margin almost straight ; apical angles obtuse but sharply defined ; a shallow, ill-defined depressed area present at side of disc towards base ; lateral margins slightly explanate from basal half to apex, rounded and slightly convergent to base ; basal margin straight, slightly thickened medially. Elytra. With deep round strial punctures ; interstices with single rows of smaller punctures, diameter equal to half or less of that of strial punctures ; scutellary striole not differentiated. Aedeagus as in Text-fig. 43b. Holotype 3 (dissected). AUSTRALIA: Queensland, bearing labels as follows: “Cape York N.Q. J. Farr/Palorus austrinus? Champ/National Museum of Victoria Melbourne ’’. Distribution. AUSTRALIA—as above. AUSTROPALORUS gen. n. Type-species : Austropalorus planatus sp. n. Length 3-3:5mm.; moderately depressed; brown, moderately dull or dull; micro- reticulation deep and dense. Head. Cuticle densely punctured ; clypeus slightly raised medially or almost flat ; clypeo- genal sutures distinct; genae flat, not produced anteriorly ; supra-orbital carinae absent ; eye not emarginate, dorsal margin below surface of head (demarzi sp. n.) or level with it (planatus sp. n.). Antennae 11-segmented, loosely articulated, appearing slender in comparison with 130 D. G. H. HALSTEAD Palorus, inserted beneath genae, scape concealed dorsally, first flagellar segment much longer than pedicel; maxillary palps (Text-fig. 46b) with apical segment slightly securiform ; other mouth parts as in Palorus. Thorax. Pronotum transverse, sides explanate, more depressed than in Palorus, base margined, apex finely margined except median half; at sides margin extremely narrow (demarzi) or absent (planatus) on basal two thirds. Scutellum transverse. Prosternal process moderately elongate, margined at sides and apex, as in Palorus ; metendosternite as in Palorus ; legs comparatively slightly longer and thinner than in Palorus ; protibiae as in Palorus (i.e., without spinules) but slightly broader ; tarsal formula 5—5—4. Elytra. Free, completely covering abdomen; humeral angle distinct, nine striae distinct, tenth apparent only at base, concealed for greater part of its length by lateral rim ; scutellary striole indistinct (represented by 2 or 3 punctures in demarzi); interstices with confused puncturation, punctures small, approximating to two rows ; micro-tubercles may be present ; epipleura comparatively broad (compared with Palorus), slightly concave, tapering to apical eighth, then narrow to apex. Wings well developed, venation as in Palorus. Abdomen. Sternites—five visible, disc of 2nd sternite without deep internal pits in 9 (demarzt) ; in 3 (planatus) disc with small triangular area of deep internal pits, apex of area at middle of sternite and base at basal margin. (3, aedeagus and goth pleurites (planatus) as in Palorus ; 9, styli similar to Palorus and Pseudeba (Text-fig. 46c). Key TO SPECIES OF AUSTROPALORUS I Pronotum slightly convergent to base, sides broadly explanate from base to apex (Text-fig. 47) ; elytra with micro-tubercles on interstices ; eyes with dorsal margins level with dorsal surface of head ; length 3:-5mm. __.. planatus sp. n. (p. 130) — Pronotum cordiform, sides explanate only at middle (Text-fig. 46a) ; elytral interstices without micro-tubercles ; eyes with dorsal margins lower than dorsal surface of head : length 3-o mm. . 5 : ‘ , . demarzi sp. n. (p. 131) Austropalorus planatus sp. n. (Text-fig. 47) 3 (2 unknown). Length 3:-5mm.; breadth 1-4mm.; general facies as demarzi sp. 0. ; brown, dull; micro-reticulation very distinct and dense. Head. Similar to that in Text-fig. 46a; punctures separated by one to two diameters, puncturation finer and less dense than in demarzi; clypeal margin shallowly emarginate, clypeus almost level with genae ; eye with greater part of dorsal surface level with dorsal surface of head. Pronotum (Text-fig. 47). Puncturation dense, slightly less so than in demarzi, punctures separated by up to a diameter ; laterally broadly explanate from base to apex, broader at apex than base; lateral rim present only on apical third, very narrow; apical margin slightly sinuate ; basal margin rounded. Elytra. Strial and interstitial punctures difficult to see due to dense micro-reticulation ; interstices with confused puncturation and micro-tubercles ; laterally comparatively broadly explanate, explanation as broad as basal breadth of pronotal explanation. Holotype ¢ (dissected). WESTERN AUSTRALIA: Wyndham, bearing labels as follows: ‘‘ Wyndham, W. Australia/? Gen. nov. aff. Palorus/N.B. Head, Scutellum etc. as in Palorus but antennae loosely articulated; slender” in the National Collection, Canberra. Distribution. AUSTRALIA—as above. REVISION OF GENUS PALORUS (SENS. LAT.) 131 Austropalorus demarzi sp. n. (Text-figs. 46a-c) Q (¢ unknown). Length 3:omm.; breadth 1:1mm.; brown, somewhat dull; micro- reticulation distinct and moderately dense. Head. Clypeus slightly less densely punctured than rest of head, punctures of head moderately coarse and separated by one diameter or less; clypeal margin almost straight, clypeus very slightly raised above level of genae ; eye lower than surface of head. x 29 nN nee ©6292 % 0 OO S16 eae ae het Se Sallie Se" cocee ce eceo mee ‘ miata ‘0% 46a 47 Fics. 46, 47. 46, Austropalorus demarzi gen. n. et sp. n., (a) 2; (b) maxillary palp ; (c) stylus. 47, Austropalorus planatus gen. n. et sp. n., pronotum, 132 D. Gs Hy HALSTEAD Pronotum. Somewhat cordiform ; puncturation dense, punctures separated by less than a diameter and moderately coarse, at sides nearly equal to diameter of an eye facet ; laterally narrowly explanate, depressed at middle, where explanation is broadest ; lateral rim extremely narrow on basal two thirds ; apical margin almost straight ; basal margin rounded. Elytra. Scutellary striole represented by 2 or 3 ill-defined punctures ; interstitial punctura- tion confused, approximating to two rows; laterally narrowly explanate from base to apex. Styli as in Text-fig. 46c. Holotype 2 (dissected—genitalia and mouth parts (partially)). AUSTRALIA: Northern Territory, bearing label as follows: “ Austral. North T. Berry Springs XITI.57 leg H. Demarz”’ in the Frey Museum, Tutzing. Distribution. AUSTRALIA—as above. Species named after the collector, H. Demarz. PALORINUS Blair stat. n. Palorus (Palorinus) Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 135. Type-species : Palorus humeralis Gebien, rgt4. Length 2-2-3-3 ; somewhat elongate, not depressed ; red-brown to yellowish brown, dull to moderately shining ; densely and coarsely punctured. Head. Densely or very densely punctured ; clypeus broad, very slightly raised medially, obtusely angled before clypeo-genal suture, clypeo-genal sutures distinct ; genae tangential to eye, slightly raised above antennal insertions, not produced anteriorly ; eyes not emarginate, latero-ventral (in humeralis approaching the form in Coelopalorus but head behind eye not forming a very distinct, shelf-like prominence); supra-orbital carinae absent. Antennae inserted beneath genae, 11-segmented, five apical segments forming a very poorly differentiated club, apex of scape visible in bicolor and quadraticollis, pedicel either approximately equal in length to (bicolor, quadraticollis), or longer than (humeralis, opticus sp. n.), the first flagellar segment ; mouth parts (studied in detail only in humeralis) similar to those of Palorus but left mandible with a small medial tooth on cutting edge and lacinia with apical setae heavily sclerotized forming lacinial tooth. Thorax. Pronotum transverse to nearly quadrate, margined basally, laterally and apically at sides, without foveae. Scutellum transverse. Prosternal process somewhat elongate, moderately expanded behind coxal cavities, more or less margined laterally and basally. Metendosternite as in Palorus. Protibiae as in Coelopalorus, with row of scaliform spinules (Coelopalorus, Text-fig. 54b) beneath distal margin (spinules more elongate than in Coelopalorus) but differing from Coelopalorus in that apical external angle is developed, forming moderate tooth ; tarsal formula 5-5-4. Elytva. Free, covering abdomen ; each elytron with 10 single rows of deep strial punctures and a scutellary striole of 5 punctures, interstices slightly raised with single rows of fine punctures ; epipleura inclined, abruptly tapered to apical eight, then very narrow to apex. Hind wings well developed—in humeralis anal area reduced (Text-fig. 48e) (not critically examined in the other three species). Abdomen. Ventrally with five visible sternites, in g disc of sternites 2, 3 and 4 (in humeralis) or of all sternites except the apical one (in quadraticollis) with deep internal pits (¢ of bicolor and opticus sp. n. not known) ; in 9 sternites without deep internal pits. Genitalia. 9, (humeralis, bicolor, opticus sp. n.) styli as in Text-fig. 48b, very lightly sclerotized, papillate in shape, with long apical setae; ¢, (humeralis, quadraticollis) aedeagus as in Text-fig. 48c, pleurites of 9th abdominal segment as in Text-fig. 48d. REVISION OF GENUS PALORUS (SENS. LAT.) 133 Key TO SPECIES OF PALORINUS 1 Apical pronotal angles prominent (Text-fig. 48a), basal pronotal angles may form very distinct minute tooth ; length 2:-5-3-omm. . . humeralis (Gebien) (p. 134) — Apical pronotal angles not prominent (Text-figs. 49, 50, 51), basal pronotal angles not forming very distinct minute tooth ; length 2-5-3-o mm. . , 2 2 Larger species, length 3-2-3-3 mm. : brown, head and pronotum much darker than elytra ; eyes smaller, ratio of dorsal length of eye to distance from apical margin of eye to clypeal suture, 110: 140; pronotum slightly narrowed from apical quarter to apex . ‘ ‘ : ‘ ' : . bicolor (Blair) (p. 135) — Smaller species, length 2-2-2: 4 mm. ; unicolorous brown ; eyes larger, ratio of dorsal length of eye to distance from apical margin of eye to clypeal suture, I10 : 80-99 ; pronotum narrowed apically only at apical angles ‘ ; : 3 3 Head as in Text-fig. 49, with shallow longitudinal median depression ; antenna with pedicel approximately equal in length to first flagellar segment; length 2°3-2'7 mm. ‘ : .quadraticollis (Blair) (p. 135) — Head as in Text-fig. 51, without shallow longitudinal median depression ; antenna with pedicel longer than first flagellar segment ; length of holotype 2:2 mm. opticus sp. n. (p. 136) 48 Fic. 48. Palorinus humeralis (Gebien), (a) head and pronotum ; (b) stylus; (c) aedeagus, dorsal ; (d) pleurites of 9th abdominal segment of ¢; (e) wing, anal region. a) 134 D. G. H. HALSTEAD Palorinus humeralis (Gebien) comb. n. (Text-fig. 48) Palorus humeralis Gebien, 1914, Sarawak Mus. J. 2 (5) : 34. Palorus (Palorinus) humeralis Gebien ; Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 142. Length 2-5-3:0mm.; breadth 0o-7-o-9mm.; moderately elongate; yellow-brown to dark red-brown, moderately shining; micro-reticulation shallow and ill-defined where present. Head (Text-fig. 48a). Densely punctured, punctures separated by one diameter or less but not rugose; clypeus broad, slightly raised medially, less densely punctured than vertex ; genae slightly raised above antennal insertions ; eyes small, dorsal length less than distance from apex of eye to clypeo-genal suture (in ratio, 5 : 7) ; antennae rather thick, pedicel slightly longer than first flagellar segment. Pronotum (Text-fig. 48a). Weakly transverse, disc slightly depressed with punctures separated by 1-2 diameters, lateral punctures separated by one half diameter or less; apex margined to medial half ; apical angles prominent, apices rounded ; sides subparallel or slightly convergent to basal half or third then convergent to base; basal angles frequently produced forming minute tooth. Elytra. Striae strongly punctured, scutellary striole of five punctures, often becoming obsolete apically ; interstices with single rows of punctures, frequently with a double row on basal half of interstice 4. Genitalia. 3, aedeagus as Text-fig. 48c, 9th pleurites as Text-fig. 48d; 9, styli as Text-fig. 48b. LECTOTYPE, present designation (sex indet.). SABAH: Kudat, bearing labels as follows: ‘“‘ Type: No. 287 [pink type label]/Br. N. Borneo Kudat [MS]/Palorus humeralis Geb.’’, Frey Mus. ~ Paralectotypes : two examples, locality as lectotype, bearing labels as follows : “ Cotype : 287 [orange label]/Br. N. Borneo Kudat [MS]/Palorus humeralis Geb ’’, Frey Mus. Comparative notes. The form of the apical pronotal angles readily distinguishes this species. Distribution. Oriental. P. humeralis was originally described from N. BoRNEO (SABAH) and later recorded from CEYLON by Blair (1930). I have seen specimens from “‘ E. BorNEo,”’ JAVA, SUMATRA and NEw GUINEA (it was collected in New Guinea in 1866 by Wallace). It has been imported into Britain in produce from Malaya and Singapore where it is probably indigenous. Habitat. In the British Museum (Natural History) there are specimens from Java with the following data. ‘‘In wood with borers and nangas—termites, Buitenzorg 29.vii.1926 Dr. Kalshoven. m. 6 i g [?]”. This species has been imported in nutmegs from Penang and sago flour from Singapore and there are specimens in the British Museum found in a plant collected on Mt. Kinabalu, N. Borneo in 1949. REVISION OF GENUS PALORUS (SENS. LAT.) 135 Palorinus opticus sp. n. (Text-fig. 51) @ ($ unknown). Length 2:2mm.; breadth 0o-7mm.; dark brown, moderately shining ; micro-reticulation shallow and ill-defined. Head. Eyes large, dorsal length slightly greater than distance from apex of eye to clypeo- genal suture, in ratio of 10:9; antenna with pedicel slightly longer and wider than first flagellar segment. Pronotum. Quadrate; densely punctured; apex nearly straight, margined on lateral thirds ; sides subparallel for apical two thirds then very slightly convergent to base; base slightly rounded, basal angles nearly right angles. Elytra. Striae strongly punctured, scutellary striole of five punctures, interstices with single rows of fine punctures. Holotype 9 (right middle and hind legs missing). BORNEO, bearing labels as follows: “ Borneo Doherty {ink on upper surface of card mount Sharp’s MS]/Sharp Coll. 1905-313 ”’ (Text-fig. 51) in British Museum (Nat. Hist.). Comparative notes. Very close to humeralis but may be distinguished by its smaller size, larger eyes and pronotal apical angles. Palorinus bicolor (Blair) comb. n. (Text-fig. 50) Palorus (Palorinus) bicolor Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 143. Length 3:2-3-3 mm.; breadth 1-1-1 mm. ; large species; brown, head and pronotum much darker than elytra, dull; micro-reticulation ill-defined. Head. Puncturation (excluding that of clypeus) dense, longitudinally rugose (Text-fig. 50) ; clypeus broad with puncturation sparser than on rest of head, not rugose ; genae raised above antennal insertions ; vertex with shallow longitudinal median depression ; eyes small, dorsal length less than distance from apex of eye to clypeo-genal suture in ratio of I1 : 14. Pronotum (Text-fig. 50). Densely punctured, base narrower than elytral base, widest towards base ; apical angles obtuse ; apex margined to medial half; sides slightly convergent apically and distinctly convergent basally. Elytva. Striae on disc separated laterally by approximately three times a puncture diameter. LECTOTYPE, present designation (sex indet.). S. INDIA: Nilgiris, bearing labels as follows: “‘ Type [standard B.M. (Nat. Hist.) type label]/Coonor R. Nilgiris, Madras C. F. C. Beeson 7.iv.1924/R.R.D. 198 B.C.R. 136 Cage 201/ex Poinciana elata/471/P (Palorinus) bicolor Blr. T. det K. G. Blair’, B.M. (Nat. Hist.). Paralectotype from the same locality as the lectotype, B.M. (Nat. Hist.). Comparative notes. The slight apical and strong basal convergence of the pronotal sides, the bicoloured body, the body size and the smaller eyes serve to distinguish this species from guadraticollis. Distribution. Inp1A. I have seen only the type material—there are no other records of this species. 136 D.G. H. HALSTEAD Palorinus quadraticollis (Blair) (Text-fig. 49) Palorus (Palorinus) quadraticollis Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 142. Length 2:3-2:7mm.; breadth 0-7-0-8mm.; unicolourous brown; very similar to bicolor but more shining. Head. (Text-fig. 49). Vertex with longitudinal shallow median depression ; eyes larger and more prominent than in bicolor, ratio of dorsal length to distance from apex of eye to clypeo-genal suture, I1: 8. Pronotum. Puncturation slightly denser than in bicolor ; form similar but sides not distinctly convergent apically. Elytra. Striae on disc separated laterally by one puncture diameter or slightly more ; densely punctured. LECTOTYPE, present designation (sex indet.). S. Inp1A: Nilgiri Hills, bearing 49 51 Fics. 49-51. 49, Palorinus quadraticollis (Blair), head. 50, Palorinus bicolor (Blair), head and pronotum. 51, Palorinus opticus sp. n. REVISION OF GENUS PALORUS (SENS. LAT.) 137 labels as follows: ‘“‘ Type [standard B.M. (Nat. Hist.) type label]/1621r/Nilgiri Hills/Andrews Bequest B.M. 1922-22 ’’, B.M. (Nat. Hist.). Paralectotypes : two examples, data as lectotype, B.M. (Nat. Hist.). Distribution. INpDIA—as for types. There are no additional records. PROLABRUS Fairmaire and ASTALBUS Fairmaire Prolabrus Fairmaire, 1897, Annls Soc. ent. Belg. 41: 111. (Text-fig. 52). Prolabrus parallelus, Fairmaire, 1897, loc. cit. Astalbus Fairmaire, 1889, Annls Soc. ent. Fr. 68: 484. (Text-fig. 53). Astalbus scrobicollis Fairmaire, 1899, loc. cit.; Astalbus longicollis Ardoin, 1959, Naturaliste malgache 11:90. A. longicollis may be separated from A. scrobicollis by the short lateral pronotal foveae, which reach only to the basal half. Ardoin (1959) redescribed these closely related Madagascan genera and suggested that they were related to Palorus. A male of Prolabrus parallelus (from Dicly— new locality record) and two females of Astalbus scrobicollis (from Tsaramandroso, Ampijoroa and the forest of Ankarafantsika—new locality records) were examined and characters pertinent to the relationship of these genera with other members of the Palorus genus group were recorded. The specimens have been deposited in the Paris Museum. Most of these characters are common and are as follows: Head. Moderately densely or densely punctured ; clypeus raised medially ; clypeo-genal sutures indistinct ; genae raised above antennal insertions, not distinctly raised or produced anteriorly ; eyes not emarginate, lateral, appearing somewhat lenticular, prominent due to development of side of head ; supra-orbital carinae absent. Antennae, robust, inserted beneath genae, I1-segmented, apical five segments forming a very indistinct club, apical two thirds of scape exposed, pedicel equal to (A. scrobicollis) or slightly shorter than (P. parallelus) first flagellar segment ; mouth parts not studied in detail but apical segment of maxillary palp elongate. Thorax. Pronotum elongate, margined at base, sides and apical angles, with or without lateral foveae ; scutellum transverse. Prosternal process moderately elongate, weakly margined at sides and apex, in Astalbus apex obcuneate, in Prolabrus apex as in Palorus. Metendosternite similar to Palorus but stem comparatively longer. Protibiae (studied in detail only in Astalbus) with small scaliform spines (similar to those in Palorinus), widely separated, inserted beneath external margin; external apical angle produced into tooth, internal apical angle bearing a small and a large articulated tooth (as in Palorus) and internal margin with long setae ; tarsal formula 5-5-4. Elytva. Free, covering abdomen completely ; humeral angle ill-defined ; ten rows of striae present, striae 9 and 10 close together at margin and with scutellary striole of four or five rather small punctures ; interstices slightly raised, most distinctly so towards apex ; apex of elytra appearing slightly sulcate; puncturation—single rows of fine punctures; epipleura as in Coelopalorus. Wings (Astalbus) similar to Palorinus but anal veins 1-4 represented (anal veins I and 2 very lightly sclerotized and rather indistinct). Abdomen. Five visible sternites; in 2 Astalbus no internal pits on disc; in ¢ Prolabrus disc of sternites 1, 2 and 3 with deep internal pits. Genitalia. (Prolabrus) as in Text-figs. 52b, c, aedeagus with parameres longer than basal piece, fused dorsally and joined ventrally by membrane forming a tube, sclerotized struts of 138 FIG. 52. D.G. H. HALSTEAD 0-Imm a 52 Prolabrus parallelus Fairmaire, (a) g; (b) aedeagus ; (c) pleurites of 9th abdominal segment of ¢ (apex shown enlarged). . a. REVISION OL-GENUS PALOR US (SENS. LACT.) 139 median lobe visible in paramere tube ; pleurites of 9th abdominal segment joined apically and of a distinctive form. 9 (Astalbus) styli (Text-fig. 53b) sclerotized, with heavily sclerotized distal margin and long apical setae, elongate. These two genera appear to be most closely related to the oriental genera Coelopalorus and Palorinus. Sie, Fic. 53. Astalbus scrobicollis Fairmaire, (a) 9; (b) stylus. ENTOM. IQ, 2 Io 140 D..G. HH. HALSTEAD COELOPALORUS Blair stat. n. Palorus (Coelopalorus) Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 135. Palorus (Palorinus) partim. Blair, Ibid. Type-species : Palorus (Coelopalorus) foveicollis Blair. Length 1-9-4:°3mm.; body depressed ; dark brown to yellowish brown, shining or dull ; micro-reticulation indistinct or distinct. Head. Moderately densely punctured, anterior margin rounded or slightly emarginate ; clypeus raised medially ; genae tangential to eye, not raised above level of clypeus, not produced anteriorly, slightly raised above antennal insertions ; with a sulcus at inner basal margin of eye (Text-fig. 55b) ; eyes not emarginate, latero-ventral, lateral base resting on a shelf-like prominence of side of head, dorsal surface level with surface of head; supra-orbital carinae absent. Antennae inserted beneath genae, 11-segmented, pedicel slightly longer than first flagellar segment or approximately equal to it, 5 apical segments forming indistinct club ; mouth parts similar to those of Palorus but left mandible with distinct medial tooth on cutting edge. Thorax. Pronotum transverse, margined basally, laterally and apically at apical angles, © with or without deep lateral foveae, with a slight depression at each side near basal margin (see — Text-figs. 54a, 56a) laterally without a distinctly flattened region; scutellum transverse. Prosternal process comparatively broad, broadly expanded behind coxal cavities, not distinctly margined (unlike Palorus). Metendosternite as in Palorus. Protibiae (Text-fig. 54b) with row of scaliform spinules beneath external and apical margins and (in foveicollis) distinct row of spines beneath internal margin (not apparent in carinatus) ; proximal margin laterally with row of setae ; tarsal formula 5-5-4. Elytra. Free, completely covering abdomen, Io single rows of strial punctures, 6 dorsal and 4 lateral (in foveicollis lateral striae becoming obsolete on basal three-quarters), scutellary striole of two to five punctures (usually absent in foveicollis) ; interstice 7 carinate from humeral angle to approximately apical sixth, interstices with single rows of fine punctures; epipleura only slightly inclined, near apical eighth tapering rather abruptly then narrow to apex. Wings well developed, anal veins all represented in foveicollis (Text-fig. 55a) but venation greatly reduced in carinatus (Text-fig. 56f). Abdomen. With five visible sternites, 3 foveicollis with deep internal pits on discs of sternites 2, 3 and 4, 2 foveicollis and both sexes of carinatus without deep internal pits on sternites. Genitalia. 3, aedeagus—parameres fused dorsally but not ventrally, closed only by mem- brane, articulated with the basal piece, sclerotized struts of the median lobe are visible in paramere tube ; pleurites of 9th abdominal segment forming moderately sclerotized, incomplete ring surrounding the aedeagus when at rest and bearing long setae at apices. 92, styli—cylin- drical, lightly sclerotized, with apical surface bearing long setae. KeEy To SPECIES OF COELOPALORUS 1 Larger species, 3:6-4:3 mm. ; pronotum with deep lateral foveae (Text-fig. 54a) foveicollis (Blair) (p. 140) — Smaller species, I-9-2-6mm.; pronotum without lateral foveae (Text-fig. 56a) carinatus (Blair) (p. 143) Coelopalorus foveicollis (Blair) comb. n. (Text-figs. 54a, 55a-f), Map 1 Palorus (Coelopalorus) foveicollis Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 136. Length 3-6-4:3 mm.; breadth 1:2-1-6mm.; depressed; dark red-brown, shining ; micro- reticulation variable, shallow but usually distinct on head and pronotum. REVISION OF GENUS PALORUS (SENS. LAT.) 141 Head. Moderately densely punctured, punctures separated by 1-2 diameters; anterior margin shallowly emarginate ; clypeus broad, slightly raised medially ; genae flat, anterior margin almost straight; eyes as in Text-fig. 55b, vertex with a small shallow median depression. Pronotum. Puncturation variable on disc, usually fine and sparse, lateral puncturation c o oo e200? te oe ee 4 : = lo bo “180808 00 00°80? Imm 54 Fic. 54. Coelopalorus foveicollis (Blair), (a) 3; (b) right tibia, ventral. ENTOM. 19, 2. Los 142 D. G. H. HALSTEAD moderately coarse and dense ; laterally with deep longitudinal foveae extending from approxi- mately apical eighth to basal quarter, deepest towards base ; side margin almost parallel for basal four fifths ; basal margin shallowly biarcuate ; a small depressed area on each side at base (see Text-fig. 54a). Elytva. Depressed, sides (lateral to carina) almost vertical ; striae lateral to carina becoming obsolete basally ; striae separated by four to seven strial puncture diameters; scutellary striole, if differentiated, represented by two to three small punctures ; interstices with single rows of fine punctures, interstice 7 carinate from base (humeral angle) to apical sixth, other interstices becoming slightly raised from apical third to apex (as indicated in Text-fig. 54a) ; lateral margin narrowly explanate with distinct expansion at about apical sixth. Genitalia. Aedeagus (Text-figs. 55c, d) a short broad structure, basal piece longer than parameres ; styli (Text-fig. 55f) short cylindrical with apical surface bearing long setae. Holotype and paratypes. Blair (1930) in his description of foveicollis, habitat data ‘‘ describes ’’ the specimen from Tenasserim as the type (“‘ Tenasserim (type) ’’). I am accepting this specimen as Holotype—it bears the following data: ‘‘ Type [British Museum circular, red-bordered label]/Tenasserim/Heteromeri/foveicollis [Blair’s MS]/Atkinson Coll 92-3”. Specimens accepted as paratypes bear the following data: 1 “‘Captn. Wimberley/Andaman Islands/heteromera/Fry Coll. 1g00.100”’, 1 “‘ Ceylon”’ (circular blue label, 2 “‘H. L. Andrewes Nilgiri Hills/ Andrewes Bequest. B.M. 1922-221’’, r “ Toungoo/Gen? near Palorus/Andrewes Bequest B.M. 1922-221’, 1 “ Penang/Bowring 63-47”, 1 “‘ Cocos-Keeling Is. Direction Id. June—July 1923 W. R. Pennifold. B.M. 1924-5 ’’, 3 “ Casteln/Malacca/ Sharp Coll. 1905-313’, 1 “ Casteln/Malacca/Fry Coll. 1905-100’, 1 ‘‘ Malacca/G. Lewis Coll. 1915-38’. I have labelled these 12 specimens “ Paratype” (British Museum circular, yellow-bordered labels). Distribution. Oriental (imported into Trinidad and E. Africa) (see Map 1). Blair (1930) records this species from Burma (type locality), CEyLon, S. INDIA, Mavacca, the PHILIPPINES and Cocos KEELING Is. In addition to these localities I have seen specimens from Formosa, N. VIETNAM (Hoah Binh), SARAWAK and KENYA (imported). Spilman (1959) records fovetcollis from the HAWAIIAN IsLAND—Oahu, TRINIDAD (associated with stored produce) and U.S.A., Mobile (in grain products) and Kulzer (1957) recorded it from Guam Is. Habitat. Spilman (1959), referring to fovezcollis collected on Oahu by Mr. E. J. Ford, Jr., said that it was collected in light traps (Feb., July, Aug., Sept., Nov.) and in the tunnels and powdery frass of the Lyctid Lyctus curtulus Casey and the Bostrychid Sinoxylon conigerum Gerst. in monkeypod, Samanea saman Merrill. Corbett, Yusope and Hassan (1937) record this species occasionally associated with copra. They illustrate and briefly describe the immature stages and note that adults and larvae feed on copra mould but not copra. This species is occasionally imported into Britain in stored products and has been recorded from stored products in E. Africa and certain oriental countries (where it is indigenous). It has arrived in Britain in illipe nuts and Malayan sago flour (from Singapore), Burmese groundnut cake, and E. African cattle food, 1954 (loaded at Mombasa). In Msambweni, Kenya, 1952, it was found on Cowpeas. Mr. F. N. Wright, a colleague, collected it in spillage (4 yrs. old) in a rice store in Sarawak. REVISION OF GENUS PALORUS (SENS. LAT.) 143 Coelopalorus carinatus (Blair) comb n. (Text-fig. 56, Map r) Palorus (Palorinus) carinatus Blair, 1930, Indian Forest Rec. Ent. Ser. 14 (5) : 143. Length 1-9-2-6mm.; breadth 0-7-0-9mm.; depressed; yellowish brown, dull; micro- reticulation strong. Head (see Text-fig. 56a). Moderately densely punctured, punctures separated by 1-3 diameters ; anterior margin rounded; genae flat; clypeo-genal suture not distinct, clypeus thus ill-defined ; clypeal region with small anterior median area impunctate (indicated in Text- fig. 56a) and slightly raised medially ; vertex with very shallow median depression ; eyes, see Text-fig. 56a and Text-fig. 55b ; antennae longer than pronotum (antennae : pronotum 13: 12). Pronotum (Text-fig. 56a). Transverse, depressed, densely and coarsely punctured, disc usually with finer, sparser puncturation ; apex straight ; base weakly indented on each side of medial third at a small shallow depression. O-Imm J) Fic. 55. Coelopalorus foveicollis (Blair), (a) wing—W, anal cell, other lettering as in Text- fig. 4.; (b) eye; (c, d) aedeagus (c) dorsal and (d) side view; (e) pleurites of 9th abdominal segment of ¢; (f) stylus, 144 D. G. H. HALSTEAD Fic. 56. Coelopalorus carinatus (Blair), (a) pronotum and head ; (b) styli; (c, d) aedeagus (c) side and (d) dorsal view; (e) pleurites of 9th abdominal segment of ¢; (f) wing, anal region, REVISION OF GENUS PALORUS (SENS. LAT.) 145 Elytra. Similar in shape to foveicollis (Text-fig. 54a), depressed, sides lateral to carina almost vertical ; strial punctures large, striae separated by 2-3 strial puncture diameters, scutellary striole ill-defined, 2-5 punctures, often becoming obsolete apically ; interstices with single rows of fine punctures, interstice 7 carinate from humeral angle to apical sixth (as in foveicollis), other interstices slightly raised for whole length, more distinctly so apically. Genitalia. Aedeagus (Text-figs. 56c, d) elongate, parameres longer than basal piece ; styli (Text-fig. 56b) elongate, cylindrical, bearing long setae on apical surface. Holotype and paratypes. Blair (1930), in his description of this species, said “ Habitat : S. India (type), Nilgiri Hills, 3,500’, January and December, under bark of dead Ficus and Grevillea (H. L. Andrewes) ; Kanara (H. E. Andrewes) ; Ceylon, Dikoya, 4,000’, XII.1881 (G. Lewis)’. In the British Museum (Nat. Hist.) there are three specimens (on the same card) labelled ‘‘ Type [British Museum circular, red bordered label]/1509 [red ink MS}/Nilgiri Hills/Andrewes Bequest. B.M. 1922-221/Palorus carinatus Blr. T. det. K. G. Blair’’, one of the specimens is differentiated by the letter T on the card below the specimen ; I am accepting this as the holotype. In addition there are 13 specimens bearing the label ‘‘ Andrews Bequest B.M. 1922-221” including: 4 labelled ‘““H. L. Andrews Nilgiri Hills ’’, 4’ labelled “ Nilgiri Hills/1509 [red ink MS] ’’, as the series including the holotype, 6 (5 on the same card) labelled “ Nilgiri Hills H. L. Andrewes XII.07 3,500 ft. ”’. Also the specimens from Ceylon (Lewis) and Kanara (H. E. Andrews) are present. I am accepting these 17 specimens as paratypes and have labelled them with British Museum circular, yellow-bordered paratype labels. Distribution. Oriental (see Map 1). In addition to material from the type localities, S. INDIA and CEyYLon, I have seen specimens with the following data: “Lenggong Malay Peninsula Lea & Party/S.A. Museum specimen’’, “ Bantam Java, de Vos”’ (in Frey Mus.), ‘‘ Phil Islands Honolulu H.T. 11.15.1933 on rice grain 5542’ (in Smithsonian Institute, Washington), ‘“‘ Hainan Tuchan 14. vi. 1959 ”’ (in Hung. Nat. Hist. Mus.). Habitat. Blair (1930) records this species under bark of dead Ficus and Grevillea. It is sometimes associated with stored products. I have seen specimens collected from Areca catechu L. in a store in Malaya. It appears to have been carried into, Honolulu in rice (see above) and has been collected in the United Kingdom on illipe nuts from Malaya. xX. ACKNOWLEDGMENTS For loan of material, on which this study has been entirely dependent, I wish to thank the following: Mrs. A. Aitken, Ministry of Agriculture, Fisheries and Food, Tolworth, England, Mons. P. Ardoin, Arcachon (Gironde), France, Dr. Bielawski, Academie Polonaise des Sciences, Warszawa, Dr. G. Colas, Muséum National D’Histoire Naturelle, Paris, Dr. A. Collart, Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Dr. J. W. Evans, The Australian Museum, Sydney, Dr. H. Freude, Zoologische Sammlung des Bayerischen Staates, Miinchen, Mr. G. F. Gross, The South Australian Museum, Adelaide, Dr. Z. Kaszab, Hungarian Natural History Museum, Budapest, Dr. K. H. L. Key, C.S.I.R.O., Canberra, Dr. H. Kulzer, 7 1 in the Hungarian Natural History Museum, Budapest. 146 D.G. H. HALSTEAD Museum G. Frey, Tutzing, Dr. L. R. Natvig, Universitetet i Oslo, Mr. A. Neboiss, National Museum of Victoria, Melbourne, The President, Forest Research Institute, Dehra Dun, India, Prof. Dr. H. Sachtleben, Deutsches Entomologisches Institut, Berlin, Mr. T. J. Spilman, Smithsonian Institute, U.S. National Museum, Washington, Dr. S. Stockmann, Helsinki, Prof. G. C. Varley, Hope Department of Entomology, Oxford University, Miss C. M. F. von Hayek, British Museum (Natural History), London, Dr. R. L. Wenzel, Chicago Natural History Museum, Illinois, Mr. J. T. Woods, Queensland Museum, Brisbane. I am particularly indebted to Mons. P. Ardoin who helped me with material on many occasions and to Dr. Kaszab, Dr. Kulzer and the Trustees of the British Museum (Natural History) for the protracted loans of their large Palorus collections. Mr. E. T. Bezant (Ministry of Agriculture, Fisheries and Food) kindly made available copies of publications on Palorus. Finally I wish to thank Mr. J. Balfour-Browne, British Museum (Nat. Hist.) for advice given on various matters, and the following members of this laboratory : Mr. G. E. Woodroffe and Mr. C. W. Coombs for their careful reading of the manuscript and for testing the keys, and Mr. J. H. Hammond for photographic assistance. XI. REFERENCES ANDRES, A. 1931. Catalogue of Egyptian Tenebrionidae. Bull. Soc. ent. Egypte 15: 122. ARDOIN, P. 1959. Contribution a l’étude des Ténébrionides malgaches, genres Astalbus Fairm. et Prolabrus Fairm. Naturaliste malgache, Tananarive 11: 87-93, 4 figs. Arrow, G. J. 1951. Horned beetles. A study of the fantastic in nature. 154 pp.,15 pls. The Hague. Brair, K. G. 1930. The Indian species of Palorus Muls. (Coleoptera: Tenebrionidae) and some associated beetles. Indian Forest Rec. Ent. Ser. 14 (5) : 133-152, 1 pl. 1935. Further new species and other records of Marquesan Coleoptera. Bull. Bernice P. Bishop Mus. 114 : 289-297. CarTER, H. J., & Zeck, E. H. 1937. A monograph of the Australian Colydiidae. Proc. Linn. Soc. N.S.W. 62 : 181-208, 2 pls., 2 figs. CHAMPION, G. C. 1896. Some remarks on the insects belonging to the genus Palorus Muls., with a description of one new species. Entomologist’s mon. Mag. 32 : 26-30. CHATTERJI, S., SARUP, P. & MENoN, R. G. M. 1961. Biological observations on Palorus shikhae Sarup, Chatterji and Menon, a predator of some stored cereal pests. Indian J. Ent. 23 (3) : 241-243, 1 fig. CorBET, G. H., Yusope, M. & Hassan, A. 1937. Insects and fungi and bacteria associated with copra in Malaya. Scient. Ser. Dep. Agric. Straits Settl, F.M.S. 20 (1): 1-91, 12 tables, 9 pls. EL-Kir_, A.H. 1953. Morphology of the adult Tvibolium confusum Duv. and its differentiation from Tribolium (Stene) castaneum Herbst (Coleoptera: Tenebrionidae). Bull. Soc. Fouad I. Ent. 37 : 173-249, 76 figs. FLEISCHER, [A.]. 1900. Uebersichtstabelle der Arten der Coleopteren—Gattung Palorus Duv. Wien. ent. Zig 19 : 236-237. GEBIEN, H. 1920. Coleoptera, Tenebrionidae. Nova Guinea 13 : 213-500, 3 pls. 1940, Katalog der Tenebrioniden II. Mitt. miinch. ent. Ges. 30 : 765-768. GreEssITT, J. L. 1958. Zoogeography of insects. A. Rev. Ent. 3 : 207-230. HaFEEz, M. A. & GARDINER, B.G. 1964. The internal morphology of the adult of Tribolium anaphe Hinton (Coleoptera: Tenebrionidae). Proc. R. ent. Soc. London (A) 39 : 137-145, 8 figs. REVISION OF GENUS PALORUS (SENS. LAT.) 147 HALsTEAD, D. G. H. 1966. <A systematic and biological study of the genus Palorus Mulsant (sensu lato) (Coleoptera: Tenebrionidae). Unpublished Ph.D. dissertation, London University Library. Hinton, H. E. 1948. A synopsis of the genus Trvibolium Macleay, with some remarks on the evolution of its species—groups (Coleoptera, Tenebrionidae). Bull. ent. Res. 39: 13-56, 33 figs. JacguELIN bu VAL, P. N. C. 1859-63. Manuel Entomologique. Genera des Coléoptéres d’Europe, 3. 464 pp., 100 pls. Paris. KaszaB, Z. 1939. Tenebrioniden aus Neu-Guinea. Nova Guinea (n.s.) 3: 185-267, 73 figs. KLEeBs, R. 1910. Uber Bernsteineinschlusse in allgemeinen und die Coleopteren meiner Bernsteinsammlung. Schr. phys.—ékon. Ges. Kénigsb. 51 : 217-42. Kocuer, L. 1958. Catalogue commenté des Coleoptéres du Maroc. Fasc. VI. Tenebrionides. Trav. Inst. scient. chévif. (Zool.) 12 : 1-185. Kuizer, H. 1957. Coleoptera: Tenebrionidae. Insects Micronesia 17 (3) 185-256, 11 figs., I map. LerkovitcH, L. P. 1964. A review of Laemophloeinae (Coleoptera : Cucujidae) from Réunion and Mauritius. Proc. R. ent. Soc. Lond. (B) 33 : 125-130. LINDBERG, H. 1962. Coleoptera insularum Canariensium III. Tenebrionidae. Commentat. biol. 25 (1) : 1-85, 11 pls., 31 maps. LinproTH, C. H. (Ed). 1960. Catalogus coleopterorum Fennoscandiae et Daniae. 478 pp., I map. Lund. MatTTINGLy, P. F. 1962. Towards a zoogeography of the mosquitoes. Publs Syst. Ass. 4: 17-36, 4 maps. MutsanT, M. E. 1854. Historie naturelles des Coléoptéves de France, 5 Latigénes—x + 396 pp. Paris. SaRuP, P., CHATTERJI, S. M. & MENON, M. G. R. 1960. Taxonomic studies on Indian Tenebrionidae. II—A new species of Palorus Mulsant, predaceous on Latheticus oryzae Waterhouse. Indian ]. Ent. 22 (4) : 239-243, 9 figs. SPILMAN, T. J. 1959. Notes on Edrotes, Leichenum, Palorus, Eupsophulus, Adelium and Strongylium (Tenebrionidae). Coleopts Bull. 13 : 58-64. TuHomson, C.G. 1859. Skandinaviens Coleoptera.1. 290 pp. Lund. Witson, J. L. 1963. Continental drift. Scient. Am. 208 (4) : 86—100. Wot aston, T. V. 1862. The Euphorbia infesting Coleoptera of the Canary Islands. Tyvans. ent. Soc. Lond. (ser. 3) 1 : 136-189. 1864. Catalogue of the Coleopterous insects of the Canaries in the collection of the British Museum. xiii + 648 pp. London. 1865. Coleoptera Atlantidum, being an enumeration of the Coleopterous insects of the Madeiras, Salvages and Canaries. xlvii + 526 pp. London. XII SPECIES INDEX (Synonyms in italics) acutangulus, 108 bobiriensis, 90 ambiguus, 96 andrewesi, 124 camerouniensis, 104 ardoini, 106 carinatus, 143 auranteus, I21 carinicollis, 79 austrinus, 109 cerylonoides, 108 crampeli, 81 baphiae, 106 beesoni, 119 delicatulus, 68 bicolor, 135 demarzi, 131 bicornutus var., 81 depressus, 96, 99 bifoveolatus, 82 deserticola, 92 148 diversicornis, 79 dolon, 67 euphorbiae, 94 eutermiphilus, 127 extlis, 108 ficicola, 92 floricola, 96 formiceticola var., 99 fossor, 126 foveicollis, 140 fuhoshoanus, 115 galilaea, 96 genalis, 118 glabratus, 68, 79, 80 grossi, 102 humeralis, 134 hypophloeoides, 114 intermedius, I10 kaszabi, 117 laesicollis, 88 laxipunctus, 86 longicollis, 137 longifoliae, 122 longitarsus, 85 mahenus, 96 marginatus, 104 melinus, 99 minor, 108 1 MAR 1967 INDEX nanus, 9I neboissi, 101 novica, 127 obtusus, 88 opticus, 135 orientalis, 100 parallelus, 137 papuanus, 108 planatus, 130 praslinensis, 108 puncticollis, 129 pygmaeus, 86 quadraticollis, 136 quadricollis, 67 ratzeburgil, 96 reticulatus, 112 saipanensis, 118 scrobicollis, 137 shikhae, 68 shoreae, 116 sinuaticollis, 115 subdepressus, 82 subfilum, 92 tenuipunctatus, 122 unicolor, 99 upoluensis, 88 zimmermani, 108 Se ‘ A LIST OF SUPPLEMENTS TO THE ENTOMOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) 1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British Museum (Natural History) and in the Hope Department of Entomology, Oxford. Pp. 143. February, 1965. £5. 2. Nixon, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera : Braconidae). Pp. 284; 348 Text-figures. August, 1965. £6. 3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177; 18 plates, 270 Text-figures. August, 1965. £4 4s. 4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172; 500 Text-figures. October, 1965. £3 5s. 5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156; 475 Text-figures. November, 1965. {2 I5s. 6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae. Pp. 129; 328 Text-figures. 3. 7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera : Coccoidea). In press. 8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera : Geometridae). In press. g. Hemminc, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). In press. Io. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa- locera). In press. PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, BARTHOLOMEW PRESS, DORKING eet oS 17 APR 19% Mm SURVEY OF THE Sy, EXTRA-ETHIOPIAN ORETINAE (LEPIDOPTERA : DREPANIDAE) A. WATSON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 3 LONDON: 1967 A SURVEY OF THE EXTRA-ETHIOPIAN 17 APR 1967 ORETINAE (LEPIDOPTERA : DREPANIDAE) Cena fe BY, A. WATSON ,, | } Dept. of Entomology, British Museum (Nat. Hist.) Php. 149-221; 92 Text-figs.; 9 Plates. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 19 No. 3 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five sertes corresponding to the Departments of the Musewm, 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. 19, No. 3 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Entom.). © Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 18 April, 1967 Price £2 nm eae ———————<x«x— —— A SURVEY OF THE EX TRA-ETHIOPIAN ORETINAE (LEPIDOPTERA : DREPANIDAE) By A. WATSON CONTENTS Page SYNOPSIS . : é . . : : 3 i ; ; : 151 INTRODUCTION . : : : 2 : j : : : F I51 Acknowledgements : : , : ‘ : , . ore Generic affinities . , ‘ : : ‘ ? : . : 152 Distribution : . : 152 KEY TO GENERA OF ORETINAE, OTHER THAN ETHIOPIAN GENERA ; ' 153 Oreta Walker : , , : : : : eas Key TO SPECIES- Groups ; ; ‘ : : : » «260 SPECIES-GRouP ROSEA . : ‘ : : ; ; 161 SPEcIES-GrouP INSIGNIS ; s , : : : 195 SPECIES-GRrouP EXTENSA ; : : : : 199 SPECIES-GROUP FUSCOPURPUREA : ; : a 208 SPECIES-GrouP CARNEA : ' : ; - #201 SPECIES-GrouP RUBROMARGINA c : oa : : 5 © 204 Urogonodes Warren ‘ ‘ , ; ‘ ; : : 7 - €206 A statochroa Turner : : : ; : : ; : Ee ae 2D Spectroreta Warren . ; ‘ ‘ ‘ : : ; U2 NieEO Cyclura Warren . : : ; ; f ; P , i 212 REFERENCES . ; : : ; 2 ; : ‘ : 1 ee INDEX : : ; : , : Z , : : ; . 220 SYNOPSIS The 54 species of the five genera of Oretinae occurring outside the Ethiopian Region are reviewed. [The ten genera and 62 Ethiopian species have been dealt with by Watson (1965).] Speciation and distribution in Oveta are discussed. The genus Oveta is revised to species-group level, and to species level in the species-groups vosea and fuscopurpurea; the Chinese species of each species-group are treated in detail. Spectvoreta is fully revised. Astatochroa, Urogonodes and Cyclura are diagnosed, the revised synonymy and distribution of the included species given and the type-species of each genus illustrated. INTRODUCTION THE bulk of the material examined during this study is in the British Museum (Natural History) and the Museum Koenig, Bonn. The Bonn material forms part of the important collection made by the late Dr. H. Héne during the 1930’s in central, southern and eastern China. Other material has been borrowed from: Division of Entomology, C.S.I.R.O., Canberra, Australia; Forest Experiment Station, Seoul, Korea ; H. Inoue Collection, Tokyo, Japan; Institut fiir Spezielle Zoologie und Zoologisches Museum der Humboldt-Universitaét, Berlin, D.D.R.; C. Kimball Collection, U.S.A.; Bryant Mather Collection, U.S.A.; Museum of Comparative ENTOM. 19, 3 II 152 A. WATSON Zoology, Cambridge, U.S.A. ; Muséum national d’Histoire naturelle, Paris, France ; South Australian Museum, Adelaide, Australia; United States National Museum, Washington, D.C., U.S.A. The type material of most nominal species has been examined: the exceptions are noted in the text. Lectotypes or neotypes are designated where necessary. Acknowledgements. Numerous people have helped during this study by arranging the loan of valuable material, by answering questions concerning the identity and whereabouts of types and by discussing other problems either personally or in correspondence. Those who have helped include: Dr. F. M. Brown, U.S.A.; Dr. I. F. B. Common, Australia; Prof. P. J. Darlington, U.S.A.; Dr. A. Diakonoff, Netherlands ; Dr. D. Duckworth, U.S.A.; Dr. H. J. Hannemann, D.D.R.; Dr. B. Hanson, Sweden; Dr. H. Inoue, Japan; Mr. C. Kimball, U.S.A.; Dr. A. N. MacFarland, Australia ; Dr. B. Mannheims, Germany ; Mr. Bryant Mather, U.S.A. ; Dr. S. W. Pak, Korea; Dr. U. Roesler, Germany ; Dr. F. Rindge, U.S.A.; Dr. E. Todd, U.S.A.; Dr. P. Viette, France. The line drawings were prepared by Mrs. Janet E. Saunders. The half-tone illustra- tions are based on photographs taken by the Photographic Section of this Museum under the control of Mr. M. G. Sawyers. My colleagues D. S. Fletcher, I. W. B. Nye and P. E. S. Whalley have checked parts of the manuscript and made several helpful suggestions. Mr. D. S. Fletcher generously offered to carry out the final assembly of the text, drawings and photographs. Generic Affinities. A preliminary study of the affinities between the Oriental and Ethiopian genera of Oretinae indicates that two, hitherto unsuspected, fairly close relationships may exist ; firstly between Spectroreta Warren (see p. 210) and Archi- drepana Warren (19024 : 487 ; see Watson, 1965 : 142) and secondly between Cyclura Warren (see p. 212) and Epicampoptera Bryk (1913 : 7; see Watson, 1965: 9). Both Spectroreta and Cyclura are endemic to and widely distributed on the Oriental Region, ranging from the Indian Subregion to the Papuan Subregion, while Archidrepana and Epicampoptera are endemic Ethiopian genera, the former being known only from Madagascar and Comores, the latter common to Madagascar and Africa south of the Sahara. The three remaining genera of non-Ethiopian Oretinae, Oveta (p. 153), Astatochroa (p. 208) and Urogonodes (p. 206) are probably quite closely allied. On distributional evidence it is arguable that the small Papuan genera Astatochroa and Urogonodes are derivatives of Oveta, a large widespread Oriental, eastern Palaearctic and Nearctic genus. Distribution. The subdivisions of the Oriental Region adopted by Gressitt (1956) have been followed throughout this paper. Details of the distribution of the five genera under review are given under each genus. To summarize the generic and species ranges: Oveta is Oriental (33 endemic species), Palaearctic (Manchurian Subregion) (two endemic species) and Nearctic (one species, vosea Walker), with three species shared between the Manchurian Subregion and the Indo-Chinese Sub- region. Oreta is found throughout the Oriental Region, except for the Polynesian Subregion. Uvogonodes and Astatochroa are Oriental (three and two endemic species — VS a 4 oat EEE Src mh EXTRA-ETHIOPIAN ORETINAE 153 respectively) ; neither is found outside the Papuan Subregion. Spectroreta (one species) and Cyclura (nine species) are also solely Oriental. Spectroreta occurs in the Indian, Indo-Chinese, Malayan and Papuan Subregions ; Cyclura in every Subregion except the Polynesian. It may be useful to mention here the extent of the Chinese representation of Oretinae for purposes of comparison with other groups of Lepidoptera which have received attention over the last 30 years as a result of the availability of the extensive Oberthiir and Hone collections. Seventeen species of Oveta are known to occur in China together with the single species of Spectroreta. Cyclura occurs in Formosa (2 species) but not on the mainland of China. KEY TO THE GENERA OF ORETINAE, OTHER THAN ETHIOPIAN GENERA [For the latter see Watson, 1965] t Fore and hind wings with hyaline patch or patches. Proboscis absent. Antenna bipectinate. Gnathos of ¢$ genitalia without posterior processes SPECTRORETA (p. 210) — Fore and hind wings without hyaline patch or patches. Proboscis present, but vestigial. Antenna bipectinate or lamellate. Gnathos of ¢ iaouiacs with or without posterior process or processes. 2 2 Upper surface of fore wing with irregularly shaped dark marking 3 near costal end of postmedial fascia (Pl. 9, fig. 124). Outer margin of hind wing with or without short curved process. Gnathos of ¢ genitalia with pair of forcipulate posterior processes (Text-fig. 88) : , . CYCLURA (p. 212) — Upper surface of fore wing without irregular dark marking near costal end of post- medial fascia. Hind wing without process. Gnathos of $ genitalia with single medial posterior process (Text-fig. 1), or with non-forcipulate paired processes, or without processes 3 Antenna bipectinate : ‘ 5 ; ‘ ‘ ; ORETA (part) (p. 153) — Antenna lamellate . : : : : , ; ; ; : F ; 4 4 Areole present in fore wing : ; : : ; 5 — Areole absent in fore wing ; . : UROGON ODES (p. 206) 5 Dark subterminal spot present on ike on upper surface of fore wing. Mid and hind tibiae without glabrous longitudinal line. 4 genitalia with anterior margin of tegumen emarginate medially. (Text-fig. 84) ‘ . ASTATOCHROA (p. 208) — Upper surface of fore wing without dark subterminal spot on Cua. Mid and hind tibiae with glabrous longitudinal line. ¢ genitalia with anterior margin of tegumen not emarginate medially . , : ’ ‘ ORETA (part) (p. 153) ORETA Walker (Pl. 7, figs. 113-115; Pl. 9, fig. 119) Oveta Walker, 1855 : 1166. Type-species, Oveta extensa Walker, 1855 : 1166, by subsequent designation by Kirby, 1892 : 728. Oveta Walker; Strand, 1911 : 204; Warren, 1923 : 479; Gaede, 1931 : 42 [Partim]; Inoue, 1962 : 37. Dryopteris Grote, 1862a : 360. Type-species, Drepana rosea Walker, 1855 : 1164, by subsequent designation by Grote, 1863 : 345. [Synonymized by Kirby, 1892 : 728.] Hypsomadius Butler, 1877 : 478. Type-species, Hypsomadius insignis Butler, 1877 : 478, by monotypy. syn.n. Hypsomadius Butler; Strand, r911 : 205; Gaede, 1931 : 42; Inoue, 1962 : 41. 154 A. WATSON Holorveta Warren, 1902 : 340. Type-species, Cobanilla jaspidea Warren, 1896a : 335, by original designation. [(Synonymized by Warren, 1923 : 480, by transference of type-species to Oreta.] Orvetella Strand, 1916: 164. Type-species, Oveta (Ovetella) squamulata Strand, 1916 : 164, by monotypy. [Synonymized by Gaede, 1931 : 43.] Psiloveta Warren, 1923 : 485. Type-species, Oveta sanguinea Moore, 1879: 85, by original designation. syn.n. Psiloveta Warren; Gaede, 1931 : 47; Inoue, 1962 : 38. Mimoreta Matsumura, 1927: 46. Type-species, Mimoreta horishana Matsumura, 1927 : 46, by monotypy. syn.n. Rhamphoreta Bryk, 1943 : 25. Type-species, Oveta (Rhamphoreta) eminens Bryk, 1943 : 25, by monotypy. syn.n. 6. Proboscis vestigial. Antenna bipectinate, open-lamellate or closely lamellate (Text-figs. 91, 92). Apex of fore wing variously falcate, outer margin straight, convex or angulate; areole present, vein FR arising from areole or from cell. Outer margin of hind wing convex; sinuous or angulate; vein Sc + R; approximated to Fs for short distance distal to end of cell, but anastomosed with Rs in some specimens of vosea. Wing without hyaline areas. Ground-colour of wings various shades of brown or yellow, often speckled or striate with buff or brown. Upper surface of both wings with antemedial and postmedial fasciae (but poorly marked or absent in species-group rubromarginata) and with or without lustrous white cell-spot or spots. Postmedial fascia extending obliquely across upper surface of fore wing from near apex, except in species- group vubromarginata in which this fascia, where present, is approximately parallel to outer margin of wing; one or two dark spots present or absent at tornus of fore wing. Large dark spot present near outer margin between Mj, and M2 on upper surface of hind wing in most specimens of obtusa and brunnea, absent in remaining species. Tibiae of mid and hind legs each with one pair of terminal spurs, and with glabrous, often dark-coloured, longitudinal line extending along outer surface. 6 genitalia: anterior margin of tegumen not emarginate medially; saccus digitate or entire; valve variously shaped, with or without process or processes; with anellus in species-group fuscopurpurea, without anellus in remaining groups; diaphragma with paired medial sclerities in species-group insignis and in most species of species-group carnea, without medial sclerite in remaining groups; gnathos without medial process, with pair of medial processes or with single posteriorly directed medial process; uncus bifurcate, emarginate or entire posteriorly, invariably with pair of lateral lobes; aedeagus variously shaped, with or without lateral lobes or terminal processes, vesica with or without cornuti or spines; eighth abdominal sternite with pair of lateral apodemes, posterior margin variously shaped, with lateral processes in most species. Q genitalia: ostium with or without ventral and lateral opercular structures; ductus bursae short, at least partly sclerotized; corpus bursae with single signum or without signum; post- ostial segments variously developed. There is little external sexual dimorphism in Oveta except in the shape of the fore wing, the outer margin of which is more strongly convex in the ° than in the ¢ or is weakly convex in the 9 of those species where the outer margin of the ¢ fore wing is straight. A considerable degree of individual variation is present in most species. This may take the form of variation in the ground-colour, or variation in the colour and definition of the markings. In many species there are two colour-forms: one in which the upper surface of both wings is yellow distal to the postmedial fascia (except for a brown marginal band on the fore wing and a brown area at the outer angle of the hind wing), and a second form in which there is no yellow coloration distal to the postmedial fascia. The first of these forms is referred to in the following pages as the EXTRA-ETHIOPIAN ORETINAE 155 yellow-and-brown form and the second as the brown form of the species. Both colour-forms occur, for example, in pulchripes, and have been illustrated in colour by Inoue (1961). Affinities (see p. 152). The monotypic Madagascan genus Oretopsis Watson (1965 : 145) is similar to Oveta in colour-pattern and wing shape, but differences in the venation and the highly characteristic ¢ and 9 genitalia do not indicate close relation- ships between them. Also similar in colour-pattern, but on the evidence of other characters not closely allied to Oveta, are the Ethiopian genera Jsospidia Watson (1965 : 132) and Uvanometra Bryk (1913: 7). Possible closer relationships are suggested by external and genitalic similarities between Oveta and the Papuan genera Urogonodes Warren (19034 : 347) and Astatochroa Turner (1926: 415). Centre of Origin. It seems unlikely that Oveta evolved outside the Oriental Region. Only six species are known to occur beyond the limits of this Region: the Nearctic rosea, for which an Oriental origin is proposed (see p. 159) ; turpis, insignis and fuscopurpurea which are shared between the Oriental Region and the south- eastern part of the Palaearctic Region ; and pulchripes and paki which are respec- tively Russo-Japanese and Korean but are closely allied to species in a predominantly Oriental species-group. Without fossil evidence it is hazardous to suggest a more exact centre of origin but, judging from the relative paucity of the Papuan fauna in terms of species and species-groups, and the absence of the genus in the Polynesian Subregion, it is reasonable to assume that the genus did not originate in the eastern part of the Oriental Region. The presence of only one species of Oveta in Ceylon and southern India eliminates the Indian Subregion as a probable centre of evolution and points to the mainland of south-eastern Asia and its south-eastern archipelago as the probable evolutionary centre of Oreta. Spectes-groups. Six reasonably well-defined species-groups can be distinguished, the most useful diagnostic features of which can be extracted from the key on page 160. General similarities in the genitalia and colour-pattern of both sexes indicate that the groups vosea and insignis seem to be fairly closely allied and that there are similar close affinities between the groups extensa and fuscopurpurea but less close affinities between carnea and rubromarginata. The present morphological evidence does not justify much more than these tentative generalizations concerning the phylogeny of Oreta. An attempt to establish the characters which are primitive relative to Oreta has proved unsuccessful: those characters which occur generally throughout the subfamily and could therefore be regarded as primitive where they occur in Oreta, are not confined to or clustered in any particular species-group. The distribution of the species-groups is equally unhelpful in assessing the phylogeny of Oveta as their ranges either coincide or overlap over a wide area. Distribution of species-groups and species. [Zoogeographical terms are those of Gressitt (1956).] The group vosea (p. 161) includes seventeen species, fourteen of which occur in China. Its range is chiefly Indo-Chinese (12 endemic species) with minor incursions into the Malayan Subregion and the Celebes transitional zone of the Oriental Region (one species shared between Celebes and the Malayan and Indo- Chinese Subregions) and into the Manchurian Subregion of the Palaearctic Region (two endemic species and one shared with the Indo-Chinese Subregion), but includes a 156 A. WATSON single Nearctic representative, O. rosea. The group insignis (p. 195) is best represented in the Papuan Subregion (four endemic species and one shared between the Papuan and Malayan Subregions and Celebes) with a poorer representation in the Malayan Subregion and Celebes transitional zone (two endemic Malayan species and one shared between Celebes and the Malayan and Papuan Subregions). One species, insignis occurs in China and Formosa but also penetrates into the Palaearctic Region (Japan), the only species in this group todo so. Although at least ten species can be included in the group insignis, only eight of these have so far been named. The group extensa (p. 199) extends across the whole of the Oriental Region except for the Polynesian Subregion. One of the four known species, extensa, is shared by the Indo-Chinese and Malayan Subregions and Celebes ; one is restricted to the Malayan Subregion, and one to the Indian Subregion. There is some doubt about the type- locality of the remaining species, adona (see p. 200). O. extensa is the only species of its group to occur in China. The next group, fuscopurpurea (p. 201), has only one species, recorded from both the Indo-Chinese Subregion of the Oriental Region and the Manchurian Subregion of the Palaearctic Region. Five species are united in the species-group carnea (p. 201): one Malayan, one from Celebes, two Papuan and one common to the Indo-Chinese and Malayan Subregions, the latter species (griseotincta) having been recorded in Formosa but not from continental China. The sixth and final group, rubromarginata (p. 204), which includes five species, has been collected only in the Malayan and Philippine Subregions and in the Celebes transitional zone of the Oriental Region. Three species appear to be endemic to the Malayan Sub- region and one to the Philippine Subregion, while the fifth is shared by these two Subregions and Celebes. To summarize the distribution of the species-groups : vosea is Oriental, Palaearctic and Nearctic ; imsignis is Oriental and Palaearctic ; carnea and rubromarginata are entirely Oriental in distribution. The total representation of the 39 described species of Oveta in each of the three zoogeographical regions where the genus occurs is as follows : Palaearctic Region (Manchurian Subregion) : 5 species, including 2 endemics and 3 shared with the Indo-Chinese Subregion of the Oriental Region. Nearctic Region : 1 endemic species. Oriental Region : 36 species, including 33 endemics and 3 shared with the Man- churian Subregion of the Palaearctic Region. Indo-Chinese Subregion: 18 species, including 12 endemics, 3 shared with the Manchurian Subregion of the Palaearctic Region, 1 shared with the Malayan Subregion and 2 shared with the Malayan Subregion and Celebes. Indian Subregion : 1 endemic species. Malayan Subregion: 12 species, including 8 endemics, 1 shared with the Indo- Chinese Subregion, 2 shared with the Indo-Chinese Subregion and Celebes, and 1 shared with the Papuan Subregion and Celebes. Philippine Subregion : I endemic species. Celebes Transition Zone: 4 species, including 1 endemic, 2 shared with the Malayan and Indo-Chinese Subregion and 1 shared between the Malayan and Papuan Subregions, EXTRA-ETHIOPIAN ORETINAE 157 Papuan Subregion : 7 species, including 6 endemics and r shared with the Malayan Subregion and Celebes. Polynesian Subregion : no species is known. The above analysis shows, as far as the limited number of species allows, the relevance to Oreta of the Subregional divisions of the Oriental Region proposed by Gressitt (1956, 1958). Celebes is transitional both between the Malayan and Philippine Subregions and between the Malayan and Papuan Subregions, while the Indo-Chinese, Malayan and Papuan Subregions are reasonably well-defined by the degree of species endemism. The total known representation of Oveta in China is 17 species, or 18 species if Formosa is included with China. Eight of these are at present known only from China or from China, Formosa and the southern Ryukyu Islands combined : angularis, brunnea, flavobrunnea, hoenet, liensis, loochooana, shania and trispina. One species, griseotincta, occurs in Formosa, N. India and the Malayan Subregion. Three species, fuscopurpurea, insignis and turpis, are shared with the Manchurian Subregion, four species (eminens, pavaca, sanguinea and vatama) are shared with all or part of the rest of the Indo-Chinese Subregion, while two species (extensa and obtusa) are shared with the Malayan Subregion, and Celebes, and the whole or part of the rest of the Indo-Chinese Subregion. Further material will doubtless show the need for revision of the range boundaries of some species and species-groups, although the general pattern of distribution will probably be little changed. No material, for example, has been seen from Halmahera or the Lesser Sundas, and little from the Molluccas and the Philippines. It seems likely too that the range of several species known at present only from China will prove to be more extensive when recent collections made by German and Japanese expeditions to Nepal have been studied. A more comprehensive investiga- tion of the B.M. (N.H.) material of the group imsignis will probably reveal the presence of undescribed species and modify present knowledge of the previously described species. Centres of endemism ‘and speciation. The richest areas of endemism in Orveta are apparently in the Indo-Chinese Subregion (especially southern China), New Guinea, and the Malayan Subregion. In the Indo-Chinese Subregion and New Guinea the relatively small taxonomic gaps between species and the existence of geographic variation within species such as vatama, hoenei and pavaca suggest relatively recent and probably continuing species radiation and that the centres of speciation and endemism coincide. The presence of eight species endemic to the Malayan Subregion possibly reflects the favourable conditions for divergence provided by repeated isolations of popula- tions on the islands of Sundaland during Pleistocene fluctuations in sea level. Similar radiation in the Malay Archipelago has been described in detail by Zeuner (1943) for a genus of Rhopalocera and by Gupta (1962) for a genus of Ichneumonidae. In the Nearctic O. rosea however, the origin of which is possibly middle Tertiary and almost certainly post-Eocene and pre-Pliocene (see discussion on p. 159), much slower evolu- tion seems to have taken place, It is closely allied to the remaining species of its 158 A. WATSON group which are Asiatic, and is separated from its close relative pulchripes by a relatively small taxonomic gap. In China there are two regions which appear to be centres of radiation in Oreta. The first of these corresponds reasonably well with the Yunnan Centre of de Lattin (1957) (subsequently modified by Gross, 1961) where the southward and eastward extensions of the Himalayas have presumably provided sufficient geographical and ecological barriers for speciation to occur since the late Pliocene or Pleistocene orogenesis in this region. In south-eastern China the mountainous provinces of Chekiang and Fukien have also provided conditions conducive to species radiation, similar to that which has occurred in the Rhopalocera (Gross, 1961), although this centre of speciation in Oveta coincides more closely in position with the Sinopacific Centre of de Lattin than to Gross’s South Chinese Secondary Centre. If it is assumed that the centre of origin of Oveta was in Sundaland or was con- tinental Asiatic, it follows that the Papuan centre of speciation is possibly relatively recent and probably dates from after the late Cenozoic elevation of the central mountain chain of New Guinea (King, 1962), which would have greatly increased the numbers of available ecological niches and created new topographical barriers conducive to speciation. Dispersal and speciation. Although some over-water dispersal of Oveta may have occurred between the islands of Malaya, Indonesia, the Philippines and the Papuan area (particularly between Celebes and Borneo, and between Celebes, the Moluccas and New Guinea) it is likely that the most important method of dispersal has been across land connections such as those which existed during Pleistocene regressions of the sea in Sundaland. The Oveta fauna of Borneo, for example, shares many more species with Java, Sumatra and Malaya, with which it was connected at times during Pleistocene Glacial stages, than it does with Celebes, which had no Pleistocene connections with Borneo. In fact only four species have so far been taken in Celebes, three of them, singapura, obtusa and extensa, being widely distributed in the Oriental Region. The possibility that these three widespread species may be migratory is discussed later. An example of Pleistocene speciation within Sundaland, probably as a result of island isolation, is shown in roepkei, a species endemic to Java (which is within the range of the closely related extensa), the origin of which can be explained by postula- ting the divergence of voepkei from extensa during an early Interglacial stage, which must have been of sufficient duration to allow speciation and to prohibit genetic swamping by extensa which subsequently reinvaded Java, probably during a late Glacial stage when Java was reconnected to other partsofSundaland. (The evolution of roepket elsewhere, followed by dispersal to Java, and the subsequent extinction of the species outside Java is also a possibility.) The effectiveness of the water barrier to the west and south-west of New Guinea is indicated by the fact that two of the five species-groups present in the Malayan Subregion and Celebes are not represented in New Guinea, presumably as a result of their failure to disperse across water barriers from their supposed evolutionary centre in Sundaland or continental south-east Asia. It is also possible to argue, however, EXTRA-ETHIOPIAN ORETINAE 159 that those species-groups not present in New Guinea have evolved more recently and have not had time to reach this island. The species singapura is one which must have successfully overcome a series of marine barriers. No other species of Oveta is common to both the Malayan and the Papuan Subregions, though several ancestral species may have had similar extensive ranges in the Pleistocene or earlier as the genus spread eastwards before giving rise to new subspecies, as in the present-day O. singapura, and to new species, as in the group extensa. In the latter group, O. extensa, which occurs in India and China and extends through the Malayan archipelago as far as Celebes, is replaced in New Guinea by an undescribed species. The latter species occurs also in Celebes together with O. extensa, having presumably reached there after attaining specific distinction in New Guinea, thus reversing the general pattern of west to east dispersal for Oveta, although the alternative of evolution of the undescribed species in Celebes followed by dispersal eastwards to New Guinea and the reinvasion of Celebes by extensa remains a pos- sibility. It would be instructive to know whether extensa is migratory or not and whether a migratory habit in extensa and in singapura and obtusa (the two other widely distributed species of Oveta) is of sufficient extent to explain their relatively greater success in surmounting water barriers. Distribution and origin of Oreta rosea. The species-group rosea to which O. rosea belongs (see p. 161) has a generally more northerly, temperate distribution than in the remaining groups: it is centred in southern China, but is represented in the Man- churian Subregion of the Palaearctic Region by the three species paki, pulchripes and turpis. As there are seventeen eastern Asiatic species in this group and only one North American species, O. rosea, it is reasonable to propose an Asiatic origin for the group. Regardless of which direction dispersal took, or whether the forerunners of O. rosea and its closest allies occurred in both North America and N.E. Asia before moving southwards during the gradual Cenozoic lowering of temperatures, it is possible to postulate a middle or late Tertiary or an early Pleistocene origin for O. rosea. The degree of similarity between O. rosea and its closest allies (especially pulchripes, a Russo-Japanese species) is not much less than between obtusa and brunnea for which a Pliocene or an early Pleistocene origin could be proposed (see time scale in Zeuner, 1943). O. obtusa and brunnea were probably members of a superspecies in relatively recent times; the two species now having overlapping ranges in China. Assuming similar rates of evolution both in ancestral O. rosea and in obtusa, at the latest an early Pleistocene dispersal from, say, Asia into North America of rosea or its ancestor must be proposed. However, suitable ecological conditions across the Bering Bridge probably did not exist later than the late Miocene, until when the Arcto-Tertiary Geoflora formed a continuous Holarctic belt (Dorf, 1959, 1960, Schwarzbach, rg61). The earliest date for a Bering dispersal, or the isolation of the North American from the Asian elements of the species-group rosea, is probably the Oligocene, at which time Viburnum, the larval food plant of both O. rosea and its close ally pulchripes, is first recorded as a constituent of the flora of northern North America (Dr. Kathleen M. Chesters, personal communication). Many species in other orders of insects and invertebrates of eastern North America 160 A. WATSON have eastern Asian affinities similar to those of O. rosea. Linsley (1963), for example, states that in many respects the Cerambycidae (Coleoptera) of eastern North America (the Alleghenian fauna) are taxonomically closer to those of the Manchurian Sub- region of the Palaearctic than to the Vancouveran fauna of western North America. Schmidt (1946: 150) cites spoonbills, sturgeons, alligators and cryptobranchids in which the fossil evidence points to an early Tertiary origin of the affinities between present-day Alleghenian and eastern Asian faunal elements. Schmidt contrasts this type of distribution with that between the western European and western North American faunas for which a late Tertiary, Pleistocene or Recent dispersal is suggested. The presence of what may prove to be a southern subspecies of vosea in Florida and Mississippi (see p. 164) is further evidence that vosea probably reached the southern United States before, or at the latest, during the Pleistocene. Hubbell (1961) has shown that geographical differentiation of North American species in Florida and the south-eastern coastal plain could have resulted from the isolation of island popula- tions during Pleistocene inundations of the Florida peninsula. Key To SPECIES-GRouUPS (BOTH SEXES) 1 Antennae bipectinate 2 : ; : : ° . . . . ° 2 — Antennae lamellate . 2 Postmedial fascia of upper surface of fore wing extending obliquely across wing from near apex (PI. 6, fig. 110) . 3 — Postmedial fascia of upper surface of ‘ore wing not extending obliquely across 3 wing SPECIES-GRouUP RUBROMARGINATA (p. 204) 3 Upper surface of fore wing with weakly marked postmedial fascia; large speckled area present at base and apex of wing . ; SPECIES-GRouP CARNEA (p. 201) — Upper surface of fore wing with strongly marked postmedial fascia; base and apex of wing as in Pl. 6, fig. 110. : : . SpecrEs-Group INSIGNIS (p. 195) 4 Antennae open-lamellate (Text-fig. 91) , ; : ‘ é : ' , 5 — Antennae closely lamellate (Text-fig. 92) . : : 7 5 Postmedial fascia on upper surface of fore wing arcuate (FA. re fig. 109) ‘(vatama) SpEcIES-GrouP ROSEA (p. 161) — Postmedial fascia on upper surface of fore wing not arcuate . . ‘ ; 6 6 Genitalia: (g$) eighth abdominal sternite with posterolateral processes; anellus present, heavily sclerotized; gnathos with single glabrous medial process; vesica of aedeagus with cornutus (PI. 8, figs. 116, 117); (9) as in Pl. 8, fig. 118 SpEcIES-GrouP FUSCOPURPUREA (p. 201) — Genitalia: (¢) eighth abdominal sternite without posteriorlateral processes; anellus absent; gnathos with one or two short medial processes, or medial part of gnathos absent; vesica of aedeagus without cornutus (PI. 7, figs. 113, 114); () as in Pl. 7, fig. 115 , ; SpEciEs-GrouPp EXTENSA (p. 199) 7 Wing-shape and colour- -pattern as in Pl. 6, fig. 110; diaphragma of ¢ genitalia with paired medial sclerites (Pl. 6, fig. 111). : SpEcIES-Group INSIGNIS (p. 195) — Wing-shape and colour-pattern not as in Pl. 6, fig. 110; diaphragma of ¢ genitalia without sclerites . : . : ‘ ; . SPECIES-GRrouP ROSEA (p. 161) Scope of revision. The species in the groups rosea and fuscopurpurea are dealt with fully in the following account. Those of the remaining groups are treated critically, in that statements concerning affinities, distribution, new synonymy and other Bur EXTRA-ETHIOPIAN ORETINAE 161 nomenclatorial changes are based on a study of type specimens ; but except for Chinese species no descriptive matter, illustrations or detailed lists of material are given. Treatment. In general this follows Watson (1965: 7). Wing measurements are now given in the following form: range of measurements from apex of fore wing to centre of mesoscutum, followed in parentheses by the number of specimens measured. SPECIES-GROUP ROSEA Antenna open-lamellate or closely lamellate. Outer margin of fore wing angulate in angularis, otherwise convex or straight; postmedial fascia of upper surface oblique except in angularis. Outer margin of hind wing angulate in angularis, sinuous or convex in remaining species. Saccus in ¢ genitalia digitate or entire; valve with or without membranous lobe, with one or more spines or processes; without anellus or diaphragmal sclerites; gnathos with single pos- teriorly directed medial process; aegeagus with terminal process or processes, with or without terminal band or group of spines, vesica scobinate or non-scobinate with one, two or no cornuti. Ductus bursae of 9 genitalia sclerotized, corpus bursae with single signum or without signum. Two well-defined complexes of species are distinguishable in this group. The first includes eminens, flavobrunnea, liensis, pavaca, sanguinea and trispina ; the second, hoenet, loochooana, paki, pulchripes, shania and turpis. The remaining species, angularis, vatama, rosea, and the species-pair obtusa and brunnea, are probably not taxonomically distant from either of the above complexes or from each other. The Nearctic rosea is probably closest to the hoenet complex. Of the seventeen described species, six are known only from China (hoenez, shania, flavobrunnea, liensis, trispina, angularis). A further eight species also occur in China: three of these occur in North India or Sikkim (pavaca, sanguinea, vatama) ; one is common to China, Formosa and the Ryukyu archipelago (loochooana) ; one to China and Formosa (brunnea) ; one to China and Burma (eminens) ; one to China and Japan (turpis) ; while obtusa is found in much of the Indo-Chinese and Malayan subregions and in Celebes. One of the remaining three species is Nearctic (rosea) ; one occurs in Japan and S.E. Russia (pulchripes) ; and one is apparently restricted to Korea (paki). KEY TO SPECIES OF SPECIES-GROUP ROSE A MALES 1 Large brown spot present on hind wing between M,; and Mz near outer margin. : 2 — Hind wing without large brown spot between Mi; and M2 . : F : 4 3 2 Genitalia as in Text-figs. 77-79 ; ; ; ¢ : ‘ brunnea (p. 194) — Genitalia as in Text-figs. 73-75 ; : . obtusa (p. 191) 3 Antenna open-lamellate (Text- fig. 91). Postmedial fascia of upper surface of fore wing strongly arcuate as in Pl. 5, fig. 109 ; ; . vatama (p. 187) Antenna closely lamellate (Text-fig. 92). Postmedial fascia of upper surface of fore wing not as strongly arcuate as in Pl. 5, fig. 109 , : 4 4 Outer margin of fore wing angulate at Cua (Pl. 5, fig. 107) . : angularis (p. 187) — Outer margin of fore wing not angulate at Cua . 5 : 5 5 Broad, grey, outer-marginal band present on fore wing between apex and mM 3 sanguinea (p. 181) | 162 A. WATSON — Grey outer-marginal band inert present) on fore wing narrow and not extending posteriorly as far as M3 ; ‘ ‘ ‘ ‘ 6 6 Outer margin of hind wing sinuous (PL 2, figs. 08 99), or most strongly convex at middle é ‘ : : , : : : 7 — Outer margin of hind wing evenly convex . : : : 12 7 Genitalia: saccus with short digitate medial process (Text- fig. 38) . trispina (p. 177) — Genitalia: saccus without medial process . ; ; 8 8 Genitalia as in Text-figs. 6-8; valve process strongly arcuate pulchripes (p p- 164) — Genitalia not as in Text-figs. 6-8; valve not strongly arcuate ° 9 9 Genitalia: valve without membranous lobe (Text-figs. 1, 2) : ‘ rosea (p. 162) — Genitalia: valve with membranous lobe . : ; ; : ‘ 10 10 Genitalia: vesica of aedeagus with single cornutus. ; i . hoenei (p. 172) — Genitalia: vesica of aedeagus without cornutus . ; : , : , II 11 Genitalia as in Text-figs. 16-18 . 3 , : ‘ : . turpis (p. 169) — Genitalia as in Text-figs. 20-22 ; ; ‘ : : paki (p. 170) 12 Fore wing very strongly falcate (Pl. 3, fig. 103) : : : . flavobrunnea ©. 186) — Fore wing less strongly falcate than in PI. 3, fig. 103 . 13 13 Colour-pattern of upper surface poorly marked; apical part of postmedial fascia on fore wing entirely lustrous white, posterior half of wing speckled with lustrous white scales : . pavaca (p. 181) — Colour-pattern of upper surface well marked; apical part of postmedial fascia on fore wing not entirely white, posterior half of wing not speckled with white scales 14 14 Upper surface of wings maculate (Pl. 3, fig. 102) : : : eminens (p. 184) — Upper surface of wings not maculate asin eminens . 3 : : : : 15 15 Colour-pattern of upper surface as in Pl. 2, fig. 97. Genitalia as in Text-figs. 32-34 shania (p. 175) — Colour-pattern of upper surface not as in PI. 2, fig. 97 : : : i F 16 16 Genitalia: saccus with short digitate medial process Sei sa 40) . . liensis (p. 179) — Genitalia: saccus without medial process . ; : : ‘ F 17 17 Genitalia: valve without membranous lobe : : ‘ 7 . rosea (p. 162) — Genitalia: valve with membranous lobe . : ; ; 18 18 Genitalia: vesica with cornutus; valve process robust (Text- figs. 2 3, 24). hoenei (p. 172) - Genitalia: vesica without cornutus; valve process slender (Text-figs. 14, 15) loochooana (p. 166) Oreta rosea (Walker) (Text-figs. I-5) Drepana rosea Walker, [10.xi] 1855 : 1164. Oreta rosea (Walker) Kirby, 1892 : 728. Oveta rosea (Walker); Dyar, 1928 : 632. [Good plate—probably Q.] Oreta rosea (Walker); Gaede, 1931 : 46. Drepana marginata Walker, 1855 : 1165. syn. n. Cilix americana Herrich-Schaffer, [31.xii.]1855: Band 1, pl. 82, fig. 470.[ Good fig.] [Synonym- ized with vosea Walker by Grote, 1863 : 345.] Platypterix formula Grote, 1862 : 60. [Synonymized with vosea Walker by Grote, 1863 : 345.] Dryopteris wvrovata Packard, [1865] : 377. syn.n. The colour-pattern of the upper surface of a specimen of this species has been figured by Dyar, 1928, in Seitz. The 3 and @ genitalia are figured in Text-figs. 1-5. There is considerable variation in the coloration and colour-pattern in this species. A brown form and a yellow-and-brown form occur, as in other species of this group, EXTRA-ETHIOPIAN ORETINAE 163 but there are some intermediates. Packard gave the name irrorata to a pale specimen with strongly marked fasciae. There is also some variation in shape of the valve in the ¢ genitalia (Text-figs. 1, 2). Wing: ¢§ 14:5-18-0 mm. (15) ; 2 15:5-21-0 mm. (15). The closest ally of this species is probably pulchripes, which is difficult to distinguish from rosea in coloration and colour-pattern but is easily separated by the genitalia. Fics. 1-5. Oreta vosea genitalia. 1, 2, 9; 3, g eighth abdominal sternite; 4, aedeagus; 5, 9. 164 A. WATSON An account of the distribution and probable origin of vosea is given on page 159. Compared with material from Canada and the north-eastern United States, most of the examined specimens of this species from Florida and southern Mississippi have a less strongly falcate fore wing and, in the yellow-and-brown form, a narrower yellow band between the postmedial fascia and subterminal fascia on the fore wing. With more material it should be possible to determine whether these differences have a geographical basis. As mentioned previously, Hubbell (1961) has shown that geographical differentiation (p. 160) of numerous North American species exists in south-eastern United States and can be attributed to Pleistocene events. Type material. vosea. Holotype g. Nova Scotia (ex Lt. Redman Coll.) Drepanidae genitalia slide No. 1706. In B.M. (N.H.). marginata. Holotype ¢ [no locality given in description]. Genitalia slide No. 1778. In B.M. (N.H.). americana. {Not seen.| Sex unknown, “ Am. spt.” formula. [Not seen.]| Holotype 9. New York. irrvovata. LECTOTYPE 9, here designated, labelled: 918; D. trrorata Pack. Maine. Packard Coll.; Type 14704; Dryopteris trrorata Pack., one of my type sp. ...!, Mus. Peab. Acad. In the Museum of Comparative Zoology, Cambridge, US:Ac Other material. B.M.(N.H.) CANADA: ex. from Quebec Province, Ontario, Manitoba. U.S.A.: ex. from New Hampshire, New York State, N. Carolina, Florida, Texas. C. Kimball Collection. U.S.A.: 7 ex., Florida; 2 ex. Massa- chusetts. Bryant Mather Collection. U.S.A.: 3 ex. S. Mississippi. U.S.N.M. U.S.A; 1-ex., Plorida. Oreta pulchripes Butler comb.rev. (Text-figs. 6-8) Orveta pulchripes Butler, 1877 : 477. Oveta pulchripes Butler; Gaede, 1931 : 45. Psiloveta pulchripes (Butler) Inoue, 1956 : 370. Psiloreta pulchripes (Butler); Inoue, 1959: 175. [Good figs.] Psiloreta pulchvipes (Butler); Inoue, 1962 : 40. [Good figs. of moth and genitalia.] Oveta calceolaria Butler, 1877 : 477. [Synonymized by Inoue, 1956 : 370.] Oveta auripes Butler, 1879 : 355. [Synonymized by Inoue, 1956 : 370.] Oveta thermidova Hampson, 1914 : 104. [Synonymized by Inoue, 1956 : 370.] Oveta pulchripes chosenoveta Bryk, 1949 : 28. [Synonymized by Inoue, 1956 : 370.] This species has recently been fully discussed and illustrated in colour by Dr. H. Inoue (1956, 1959, 1962), but for purposes of comparison the genitalia are re-illustrated (Text-figs. 6-8). The yellow-and-brown form of pulchripfes is externally similar to the corresponding form of liensis sp.n., whereas the brown form closely resembles the brown form of loochooana Swinhoe. Both liensis and loochooana differ from pulchripes in the less strongly convex outer margin of the fore wing, and loochooana can be distinguished EXTRA-ETHIOPIAN ORETINAE 165 by the non-angulate postmedial fascia on the fore wing. The overall specific affinities of pulchripes are doubtful. The ¢ genitalia of turpis Butler have much in common with those of pulchripes, although the 9 genitalia do not indicate particularly close affinities between these two species. O. rosea is externally very close to pulchripes. Wing. fg 15:0-21:0 mm. (5); ¢ 17-0-24:0 mm. (30). Fics. 6-8. Orveta pulchripes genitalia. 6, g; 7, g eighth abdominal sternite; 8, aedeagus; 9, Q. The ratio of yellow-and-brown to brown specimens in the material examined is Hi7': T21. Distribution. The B.M. (N.H.) collection contains specimens from S.E. Russia, and JAPAN. Inoue (1959) lists S.E. Russia, KorEA, JAPAN (including Amamioshimia) and Cutna. No Chinese specimen of pulchripes has been seen during the present ENTOM. 19, 3 12 166 A. WATSON study but there is good reason to expect at least accidental occurrences of this species inside the Manchurian border of China, especially near Vladivostock where pulchripes is known to occur. The Hone collection at the Museum Koenig, Bonn, contains only Japanese examples. Type material. pulchripes. LECTOTYPE 4, in B.M. (N.H.), here designated, labelled: 77.9 [B.M. registration 1877.9: Japan, Yokohama (Jonas)|; Oreta pulchripes Butler Type ; Drepanidae genitalia slide No. 522. calceolaria. LECTOTYPE 4, here designated, labelled : 77.9 [Yokohama (Jonas) | Japan ; Oreta calceolaria Butler Type. In B.M. (N.H.). auripes. LECTOTYPE Q, here designated, labelled: Japan, 79.48 [Yokohama], 348 ; Oreta auripes Butler Type. In B.M. (N.H.). thermidora. Holotype 9. Japan, Fushiki, vii.1886 (Leech) ; Drepanidae genitalia slide No. 521. In B.M. (N.H.). chosenoreta. Holotype §@. Korea, Shuotsu, 22.vii. In the Naturhistoriska Riksmuseet, Stockholm. Oreta loochooana Swinhoe comb. rev. (Text-figs. 10--15) Both sexes of this species have been illustrated recently in colour by Inoue (1962 : Pl. 3, figs. 59, 60). The colour-pattern of Joochooana is most like that of paki Inoue, but the two discocellular spots on the fore wing are usually more clearly defined than in paki. In wing shape loochooana can be separated from paki by the less strongly falcate apex and the weakly convex outer margin of the fore wing and by the evenly convex, not sinuous, outer margin of the hind wing. The similarity in the shape of the aedeagus and general similarity in the colour-pattern and most parts of the genitalia suggest phyletic affinities between loochooana and hoenet (Text-figs. 23-31) (especially hoenei inangulata) from which loochooana differs externally in the fore wing, by the less strongly falcate apex, the more distinctly marked tornal markings and the lighter yellow postmedial fascia. Two colour-forms exist in this species, corresponding to the yellow-and-brown and brown forms of pulchripes. Two subspecies are known: the nominate subspecies from China (Shantung), Formosa and Japan (Ryukyu Archipelago), and timutia ssp. n. from China (see below). The single example of the nominate subspecies from Shantung calls for confirmation and further investigation, as the Chinese province of Fukien which faces Formosa across the Taiwan Straits forms part of the known range of subspecies timutia and no connecting populations are yet known to occur in Korea which could form a link between the Ryukyu and Shantung elements of the nominate subspecies. a EXTRA-ETHIOPIAN ORETINAE 167 2 we - = 550 oak eres Fics. 10-15. re, 10—11, loochooana timutia. 12—15, loochooana loochooana. aedeagus, 15, ¢ Oreta genitalia. 10, g eighth abdominal sternite; 12, 2; 13, gd eighth abdominal sternite; 14, 168 A. WATSON Oreta loochooana loochooana Swinhoe (Text-figs. 12-15) Oreta loochooana Swinhoe, 1902 : 591. Oreta pulchripes loochooana Swinhoe; Strand, IgII : 205. Oreta pulchripes var. loochooana Swinhoe; Gaede, 1931 : 46. Psiloveta loochooana (Swinhoe) Warren, 1923 : 485. Psiloveta loochooana (Swinhoe); Gaede, 1931 : 48. Psiloreta loochooana (Swinhoe); Inoue, 1962 : 38. [Good figs.] Oreta (Oretella) squamulata Strand, 1916 : 164. syn. n. Psiloveta pulchripes formosicola Matsumura, 1927 : 46. syn. n. There appears to be no external difference between this subspecies and tumutia. In the ¢ genitalia the eighth sternite is differently shaped, and in the 9 the ostial segment differs in the shape of the sclerites. Wing. 3 15:0-20°5 mm. (17); 2 20:5—22-5 mm. (2). Eight ¢ and one @ of the yellow-and-brown form, and nine ¢ and one @Q of the brown form have been examined. Distribution. Formosa, JAPAN (Ryukyu archipelago). Type material. loochooana. LECTOTYPE 4, here designated from the series of four ¢ syntypes in B.M. (N.H.), labelled: Loochoo, 1896, H. Pryer Coll.; Leech Coll. rg00-64 ; Oreta loochooana Swinh. 3 type; Drepanidae genitalia slide No. 616. Paralecto- types. JAPAN: 3 ¢, Loochoo, 1896 (Pryer). squamulata. Holotype g. [Formosa] Kosempo, xi. 1911 (Sauter); Drepanidae genitalia slide No. 828. In the Deutsches Entomologisches Institut, Berlin. formosicola. Holotype 9. Horisha [Formosa] (Takamuku) [Examined for me at — Hokkaido University by Dr. T. Kumata at the request of Dr. H. Inoue]. Other material. Examples from Formosa, JAPAN (Ryukyu Archipelago, Amamio- shima, Okinawa), in the collections of B.M. (N.H.), Dr. H. Inoue and Dr. I. Daniel. One § from CuiNA (Shantung) in B.M. (N.H.), possibly erroneously labelled. Oreta loochooana timutia ssp. n. (Text-figs. 10-11) As stated earlier, external separation of the two subspecies of loochooana is probably not possible. The subspecies can be separated in the 3 by the shape of the eighth terg ite and in the 9 by the structure of the ostial segment. Wing. 16:5-19:0 mm. (46) ; 2 19°6—20°5 mm. (2). The ratio of the number of specimens of the yellow-and-brown form to the number of brown specimens in the material examined is 30: 23. Both females belong to the yellow-and-brown form of the species. Distribution. CHINA (Szechwan, Kwangtung, Hunan, Chekiang, Fukien). Holotype g. Hunan, Hoeng-shan, g00 m., 28.iv.1933 (Héne); Drepanidae genitalia slide No. 1545. In the Museum Koenig, Bonn. ee EXTRA-ETHIOPIAN ORETINAE 169 Paratypes. Museum Koenig, Bonn. CHINA: 3 3g, Kwangtung, Linping, iii- iv.1922 (Héne) ; 15 3, type-locality, 25 .iv—10.1x.1933 (Héne) ; 7 3, Chekiang, East and West Tien-mu-shan, 22. v—26.i1x. 1931, 18 .iv—29.ix.1932 (Héne) ; 1 3, Chekiang, East Tien-mu-shan, near Lingan, 1500 m., 27.v.1931 (Héne); 9 3, 1 9, Fukien, 5 .1v-12.vi.1938, 9. v—-14.vi.1946 (Klapperich) ; 2 3 [? Kiangsi] Kuling, 9.v.1931, 2.V.1934 (Hone). Oreta turpis Butler comb. rev. (Text-figs. 16-19) Oreta turpis Butler, 1877 : 477. Orveta turpis Butler; Gaede, 1931 : 47. Psiloreta turpis (Butler) Inoue, 1959 : 175. [Good figs.] Psiloreta turpis (Butler); Inoue, 1962 : 39. [Good figs.] Oreta calida Butler, 1877 : 477. [Synonymized with turpis by Inoue, 1956 : 370.] Oveta calida Butler; Strand, tg11 : 205. [Fig.] Orveta calida Butler; Gaede, 1931 : 43. A comprehensive account of this species has been given recently by Inoue (1962 : 39). It is externally close to tvispina sp.n. (brown form), pulchripes Butler (brown form) and hoenei tienia ssp. n. ; though it is probably most like the brown form of trispina (Pl. 2, fig. 99) which, however, has more strongly falcate fore wings, a more strongly sinuous outer margin to the hind wing, and a broadly divided cell-patch on the fore wing. On the basis of overall similarity, paki Inoue is the closest ally of turpis. The latter can be distinguished from paki by the absence of spots at the anal angle of the fore wing, the strongly marked transverse fasciae, and by several differ- ences in the genitalia of both sexes (see Text-figs. 16-19). Wing. 4, 9. 13:0-20:0 mm. [teste Inoue, 1962 : 40]. No equivalent of the yellow-and-brown form of, for example, paki is known to occur in turpis. Distribution. JAPAN [see Inoue references above], U.S.S.R. (Sakhalin, Ussuri), Korea [teste Inoue, 1959:175]. A male specimen labelled Tsingtau [Cu1Na, Shantung, Tsingtao], in the Museum Koenig, Bonn, is doubtless conspecific with the type of turpis but differs in the proportions of the valve processes and the aedeagus and may well represent a new subspecies of turpis. Type material. turpis. LECTOTYPE J, here designated, labelled: 77.9 Japan; Oveta turpis Butler Type ; Drepanidae genitalia slide No. 519. The restricted type-locality is Yokohama as indicated in registration No. 1877-9 [77.9] in the records of the Depart- ment of Entomology, B.M. (N.H.). Type in B.M. (N.H.). caida. LECTOTYPE 4, here designated, labelled: 77.9 Japan [Yokohama] ; Oreta calida 3 Butler Type; Drepanidae genitalia slide No. 518. Lectotype, and a 2 paralectotype from the type locality, in B.M. (N.H.). 170 A. WATSON Fics. 16-19. Orveta turpis genitalia. 16, $; 17, ¢ eighth abdominal sternite; 18, aedeagus; 109, 9. Oreta paki (Inoue) comb. n. (Text-figs. 20-22) Psiloreta paki Inoue, 1964 : 3. [Good figs.] As indicated by Inoue (1964: 3-4), this species has close affinities with turpis Butler, from which it can be distinguished by the presence of one or two dark spots near the anal angle of the fore wing, the poorly marked antemedial and postmedial fasciae on the hind wing, and by differences in the 9 genitalia and the shape of the valve processes, uncus, gnathos, aedeagus and eighth sternite in the ¢ genitalia. The affinities between paki, loochooana Swinhoe and hoenei sp.n. are probably less close than between paki and turpis, on the evidence of the total of genitalic and external characters. The external similarity between paki and hoenei (nominate 171 EXTRA-ETHIOPIAN ORETINAE Ro » fe} oO © Ot %O 4 KS oe ets, Tot Ee RR _ N ¢8 i? 9) os. SS wm XO 5 he a 8 oO. ae) Re fo) Ss ‘= : 3 = 8 Ss as peas a v8 422 s RY. 4, wy oa) ss} ae ~ oe o v na: 8 Ss oO * % ial (94 ae a & o8 ite’ “© Oo _ fy 172 A. WATSON subspecies and :nangulata) is close, but the @ genitalia and several features in the aedeagus, eighth sternite and main body of the ¢ genitalia readily distinguish these two species. Wing. g 17:5-21:0 mm. (21). Of the 52 specimens examined by Inoue, 17 belonged to the yellow-and-brown form of the species (Inoue, 1964: figs. 3-4) and the remainder to the brown form. Distribution. Korea. (See Inoue, 1964 : 4.) Type material. Holotype 3. Korea, Seoul, Chungrangri, 7.vi.1g61 (S. W. Pak). In B.M. (N.H.). [Recently kindly presented, together with five paratypes, by Dr. H. Inoue.] Other material. There is a single male from “ Utikongo im Kongosan (Mittel- Korea) ’’ in the Museum Koenig, Bonn, and further examples from Seoul (collected by Pak) in the Daniel Collection, Munich. Oreta hoenei sp. n. (Pl. 1, figs. 93-96; Text-figs. 23-31) 3. Palp and clypeofrons dull scarlet, vertex buff; antennae closely lamellate. Collar buff dorsally; dull scarlet lateral and ventral to eyes. Thorax buff or reddish brown dorsally; chiefly pale buff ventrally, but dull scarlet anteriorly. Legs dull scarlet and buff on front or outer surface, otherwise pale buff. Outer margin in fore wing moderately or strongly convex. Outer margin of hind wing evenly convex in inangulata, weakly sinuous in nominate subspecies, and strongly sinuous in tenia. Coloration of upper surface of wings variable. In the nominate subspecies and inangulata the wings may be uniformly reddish brown with little trace of the medial shade, brown with darker brown medial shade, varying shades of brown with the distal half of the hind wing bright yellow, or less frequently intermediate between these three categories. In hoenei tienia neither uniformly reddish brown nor yellow and brown specimens have been seen. The under surface of the wings corresponds approximately to the colour-pattern and coloration of the upper surface except that the antemedial fascia is absent and that, except for tienia, the postmedial fascia is poorly defined. Abdomen pinkish buff ventrally, laterally and posterodorsally, but similar to colour of base of hind wing anterodorsally. 6 genitalia as in Text-figs. 23, 24, 26, 27, 30, 31. Q. As for male but outer margin of fore wing more strongly convex. ° genitalia (Text-figs. 25, 28, 29) with emarginate ostial plate. The closest allies of hoene: appear to be turpis Butler or paki Inoue. The latter is externally similar in colour-pattern and coloration to the nominate subspecies and hoenet inangulata, whereas turpis approaches hoenet tienia in these respects. The § and 9 genitalia of both turpis and paki indicate close affinities between them and hoenei. There are similarities in the colour-pattern between hoenei and loochooana, another widespread Chinese species, but the genitalia readily separate them. Three subspecies can be distinguished: imangulata (Tibet, Szechwan, Yunnan) ; the nominate subspecies (Shansi, Shensi) and tenia (Chekiang). The species is unknown outside China. EXTRA-ETHIOPIAN ORETINAE 173 Oreta hoenei hoenei ssp. n. (Pl. 1, figs. 93, 94; Text-figs. 23-25) This subspecies differs externally from imangulata in the more strongly convex outer margin of the fore wing and the sinuous outer margin of the hind wing, and from #ienza by the generally less strongly convex outer margin of the fore wing, the weakly sinuous outer margin of the hind wing, and the usually less strongly marked antemedial and postmedial fasciae on the upper surface of both wings. In the ¢ genitalia the shape of the valve process and the terminal part of the aedeagus (Text-figs. 23, 24) distinguish this subspecies from imangulata and tienia. The shape of the 9 ostial sclerites separates the nominate subspecies from tenia and inangulata. Wing. ¢ 18-0—22:0 mm. (73); 9 21-5—23°5 mm. (4). Of the 73 males and four females examined, seven males and one female have clear yellow hind wings distal to the medial shade (yellow-and-brown form) ; the hind wings of the remainder are either uniformly brown or have the distal part of the wing a paler brown (yellowish brown in a few specimens) than the medial shade. Distribution. CHINA (Shansi, Shensi). Holotype 3. S. Shensi, Tapaishan im Tsinling, 1,700 m., 13.vill.1936 (Héne) ; Drepanidae genitalia slide No. 1554. In the Museum Koenig, Bonn. Paratypes. Museum Koenig, Bonn. CHINA: 52 4, I 9, type-locality, 20.vi- 26.1x.1935, I7.V—-17.vill.1936 (Héne); 13 3, Shansi, Mienshan, 2000 m., 3.vi- 12.vili.1937 (Héne). B.M.(N.H.). CHINA: 9 4, 2, type-locality, 16-21 .ix.1935, 7 .Vi- 21.viii.1936 (Héne) ; Shansi, Mienshan, 2,000 m., 2.vii-6.vili.1937 (Héne). Daniel Collection, Munich. CHINA: 2 3, type-locality, 26.ix.1935, IQ. vili.1936 (Héne) ; 3 3, Shansi, Mienshan, 2,000 m., 1. vii-I10. viii. 1937 (Héne). Oreta hoenei inangulata ssp. n. (Pl. 1, fig. 95 ; Text-figs. 26-28) Separable from both the nominate subspecies and tienia by the weakly convex outer margin of the fore wing, the evenly convex, non-sinuous margin of the hind wing, the shape of the valve processes and aedeagus in the 4, and by the shape of the paired processes of the ostial segment in the Q genitalia. Wing. ¢ 18-0-24:0 mm. (26); 9 22-0-22:5 mm. (2). Of the 28 specimens examined, four (including one ¢) represent the yellow-and- brown form of the species (PI. 1, fig. 95), and 23 the brown form in which the hind wing is almost uniformly brown. One specimen in the Museum Koenig is brown but with some yellow scaling on the fore wing. Distribution. CurINnaA (Szechwan, Yunnan). Holotype 3. N. Yunnan, Likiang, 30. viii.1935 (Héne) ; Drepanidae genitalia slide No. 1556. In the Museum Koenig, Bonn. Paratypes. B.M.(N.H.). CHINA: 7 3g, 1 9, Szechwan, Kwanhsien, vii.1924, 1930 (Franck et al.); 1 3, Szechwan, Kia-ting-fu, vi-vii.1890; 2 jg, Szechwan, 174 A. WATSON Suifu ; 1 3, Szechwan, Mt. Omei, Shinkaisi, 4400 ft.; 1 g, Szechwan, Tatsienlou, 1895 ; I g, Szechwan, Kwanhsien, 9.viii.1926 (Franck). YUNNAN: 5 6, type- locality, 7.vili-I.x.1934, 5, 13.ix.1935 (Héne); 1 9, China. Museum Koenig, Bonn. YUNNAN: 7 @, type-locality, 25.vii-6.ix.1935 (Héne). Fics, 26-31. Oveta genitalia. 26-28, hoenei inangulata. 26, J; 27, aedeagus; 28, 2; 29-31, hoenei tienia. 29, 2; 30, aedeagus; 31, g. EXTRA-ETHIOPIAN ORETINAE 175 Oreta hoenei tienia ssp. n. (Pl. 1, fig. 96; Text-figs. 29-31) Separated from angulata and the nominate subspecies by the more strongly convex outer margin of the fore wing, the strongly sinuous outer margin of the hind wing and the usually strongly marked transverse fasciae on both wings. The shape of the valve processes and aedeagus are diagnostic in the ¢ genitalia and in the 9 the shape of the ventral lip of the ostium distinguishes tienia from the nominate subspecies. Externally tienia resembles trispina sp.n. (Pl. 2, figs. 98, 99), except that on the fore wing the postmedial fascia is less well marked, the cell-patch is not interrupted at the distal end of the cell and there is a diffuse brown batch at the tornus. The genitalia of tvispina show, however, that it cannot be placed in the same species- group of hoenezi. Wing. 4 17°5-21:0 mm. (31); 9 20°5—-24-0 mm. (4). No specimen of the yellow-and-brown form of the species has been seen. The ground-colour of the upper surface varies from reddish brown to pale buff. Distribution. Curina (Chekiang). Holotype 3. China, Chekiang, West Tien-mu-shan, 31.v.1932 (Héne) ; Drepanidae genitalia slide No. 1563. In the Museum Koenig, Bonn. Paratypes. Museum Koenig, Bonn. CHINA: 28 4, 2 9, type-locality, 26.v— 7.X.1932 (Hone) ; 2 3, East Tien-mu-shan, 15.vi, 3.1x.1931 (Héne). B.M. (N.H.). CHINA: 6 J, 2 9, type-locality, 2.vi-12.ix.1932 (Héne) ; 1 3, East Tien-mu-shan, 13.vi.1931 (Héne). Daniel Collection, Mumch. CHINA: 5 3, I Q, type-locality, 24.V-13.V1.1932 (Héne). Oreta shania sp. n. (Pl. 2, fig. 97; Text-figs. 32-35) 3. Palp dull scarlet. Clypeofrons orange-brown, but with scarlet tuft anterior to base of each antenna; vertex dull yellow. Antennae closely lamellate, yellow near base, yellowish brown distally. Collar yellow dorsally, dull scarlet laterally and ventrally. Thorax yellow dorsally, palest anteriorly; paler yellow ventrally. Outer surface of fore legs orange-buff and dull scarlet; outer surface of tibia and tarsus of mid and hind legs mainly orange-buff but with pink fringe to tibia and dull scarlet band at distal margin of each tarsal segment. Inner surface of legs pale buff. Wing shape as in PI. 2, fig.97. Rin fore wing arises from distal end of cell (in most specimens) or from the proximal end of the areole. Sc + R; in hind wing approximates to Fs distal to end of cell. Colour-pattern of upper surface of wings as in PI. 2, fig. 97. Pale areas yellow, darker medial or marginal areas reddish brown and usually well-marked; both wings with two white cell-spots ; black or dark brown spots on costa and at anal angle on 1A and Cu, (trace). Both wings, but especially hind wing, lightly speckled with black or dark brown in a large proportion of speci- mens. Under surface of fore wing pale brownish orange, or deep yellow partly diffused with dark pink heavily speckled with dark brown; well-marked, dark brown postmedial fascia, and pale grey area immediately posterior to apex. Anterior margin of under surface of hind wing as for fore wing, but yellow posterior to cell with light speckling of dark brown. Dorsal surface of abdomen as for adjacent surface of hind wing: yellow, transversed by pale reddish brown band. Ventral surface pale buff but with longitudinal orange band on either side along lateral border of sternites. 176 A. WATSON 34 Fics. 32-35. Oveta shania genitalia. 32, g eighth abdominal sternite; 33, aedeagus; 34,5; 35, 9. ¢ genitalia as in Text-figs. 32-34. There is some minor variation in the dentation of the inner margin of the valve processes. 9. As for J but with outer margin of fore wing more strongly convex. 2 genitalia as in Text-fig. 35. Wing. 15:0-20'5 mm. (79); 9 19:5 mm. (1). No polymorphism in upper surface wing coloration is present in the available material, in contrast with the variation found in related species of this species-group (e.g. hoenei and loochooana, both of which occur in China). The precise affinities of shania within its species-group are uncertain. It is externally most like Joochooana Swin., which has a similar distribution in China but differs from shania in the much broader medial shade on both wings. EXTRA-ETHIOPIAN ORETINAE 177 Distribution. CHINA (Chekiang, Fukien, Szechwan). Holotype 3. China, Chekiang, West Tien-mu-Shan, 1. vii.1932 (Héne) ; Drepanidae genitalia slide No. 1548. In the Museum Koenig, Bonn. Paratypes. Museum Koenig, Bonn. CHINA: 42 3, type-locality, 9.vi-8.vii.1932 (Hone) ; 3 3, East Tien-mu-Shan, 4.vi-17.vi.1931 (Héne); 4 3, East Tien-mu- Shan, Lingan, 1500 m., 3I.v—17.vVi-1931 ; I5 9, Fukien, Kuatun, 2300 m., 16.v- 1.vi.1938 (Klapperich), 10.vii—4.vili.1938 (Héne). B.M.(N.H.). CHINA: 6 8, type-locality, 17-23. vi. 1932 (Héne) ; 2 J, East Tien-mu-shan, 10, 13. vi. 1931 (Hone) ; 3 6, Fukien, Kuatun, 2300 m., 13,17.v.1938, 25.v.1946 (Klapperich) ; 12 3, Szechwan, Kwanhsien, I7.vii.I924, 21. vii.1925, vii—vill. 1930(Fvanck and others) ; I g, Szechwan, Tien-tsuen, 1897; 5 3, I 9, Szechwan, Ig00-1902 ; I g, Szechwan, Moupin, 1897. Daniel Collection. CHINA: 5 4, type-locality, 1,600 m., II—23.vi. 1932 (Héne). Oreta trispina sp. n. (Pl. 2, figs.