ie oye if ti), ofatelidelngs ephetalath tay rite HAND per Metals COS eOtatee ca ere ate Sst aS a Sys + BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 1972 [* war ” | 22 MAYI975 ESENTED 4 @ < LOcy yee! BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1975 DATES OF PUBLICATION OF THE PARTS No. I 24 October 1972 No. 2 18 October 1972 No. 3 8 December 1972 No. 4 18 October 1972 No. 5 29 December 1972 No. 6 3 November 1972 No. 7 4 October 1972 No. 8 4 October 1972 No. 9 14 December 1972 No. 10 28 December 1972 No. 11 29 December 1972 ISSN: 0007-1498 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU No. I No. 2 No. 3 No. 4 No. 5 No. 6 No. 7 No. 8 No. 9 No. r0. No. Ir. CONTENTS f 22 May i970 ZOOLOGY VOLUME 23 Noe THE GUNONG BENOM EXPEDITION 1967 1. Introduction. By Lorp MEepway (4 Pls.) . z. An outline description of the forest zones on north-east Gunong Benom. By T. C. Wuitmor: (8 Pls.) . 3. Bryophytes and filmy ferns from Gunong Benom. By ANNE JOHNSON 4. New records of Malayan bats, with taxonomic notes and the description of a new Pipistrellus. By J. E. H1ty 5. Reptiles and amphibians of Gunong Benom with a description of a new species of Macrocalamus. By ALICE G. C. GRANDISON 6. The distribution and altitudinal zonation of birds and mammals on Gunong Benom. By Lorp MEDWaAy 7. The systematic status of Malayan Rattus rajah and Rattus surifer. By H.S. Yon (x Pl.) , 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. By H. Hoocstraat, B. L. Lim, M. NADCHATRAM and G. ANASTOS g. A collection of chiggers from Gunong Benom (Prostigmata : Trombiculidae). By M. NADCHATRAM io. Phthiraptera (Insecta) chewing and sucking lice. By THERESA CLAY : 11r. Notes on zoogeography, convergent evolution and taxonomy of fleas (Siphonaptera), based on collections from Gunong Benom and elsewhere in South-East Asia. I. New taxa eee Pygiopsyllinae). By R. TRAuB (58 Pls.) . 12. Notes on zoogeography, convergent evolution a Core of fleas (Siphonaptera), based on collections from Gunong Benom and elsewhere in South-East Asia. II. Convergent evolution. By R. TRAvB (20 Pls.) : b ; 13. Notes on zoogeography, convergent evolution and taxonomy of fleas (Siphonaptera), based on collections from Gunong Benom and elsewhere in South-East Asia. III. Zoogeography. By R. TRAUB = : : 5 ; : : 6 Index to Volume 23 199 201 307 389 451 1 E GUNONG BENOM EXPEDITION ee 1967 1. INTRODUCTION - LORD MEDWAY MUSEUM ( ea. HISTORY) Vol. 23 No. 1 LONDON: 1972 + Be paket ya) mh oes THE GUNONG BENOM EXPEDITION 1967 1. INTRODUCTION \ BY LORD MEDWAY School of Biological Sciences University of Malaya Pp. 1-7; 4 Plates, 2 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 1 LONDON : 1972 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 a separate supplementary series of longer papers was instituted, numbered serially for each Depariment. This paper is Vol. 23 No. 1 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 d TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) F Issued 24 October 1972 Price 75p THE GUNONG BENOM EXPEDITION 1067 1. INTRODUCTION By LORD MEDWAY Apart from relatively narrow coastal plains of Quaternary or Recent alluvium on both flanks of the peninsula, the topography of Malaya (= West Malaysia) is generally hilly or mountainous. About 40 per cent of the total land area rises above the 500 ft contour, and 23 per cent above Iooo ft (Panton, 1965). The principal mountain formations occur as a series of approximately parallel ranges striking north-northwest/south-southeast (Fig. 1). The largest of these, generally called the Main Range, forms a continuous ridge extending from the border of Perak state with Thailand to southern Negri Sembilan. In its northern sector several peaks rise to over 7000 ft, the highest being Gunong Korbu (7160 ft). Further south the elevations attained become progressively lower. The highest mountain in Malaya, Gunong Tahan (7186 ft), is not part of the Main Range, but rises from rugged hill country further to the east in an isolated position on the border of the states of Pahang and Kelantan. Still further eastward an irregular group of lower mountains collectively form the Eastern Range (Fig. I). In the centre of the country between the Main Range and the Eastern Range, situated on the right bank of the River Pahang just south of 4°N, lies the compact montane formation named after its highest peak, Gunong Benom (6916 ft) (Plate 1). Here the total extent of connected highland terrain exceeding 3000 ft in elevation amounts to about 75 square miles. These uplands are separated from neighbouring areas of comparable elevation by intervening lowlands. Southwards, the Benom formation is limited by the extensive lowlands of central Pahang. To the west it is divided from the highlands of the Main Range, about 20 miles distant at the nearest point, by the valleys of two tributaries of the River Pahang, one south- flowing and the other north-flowing, between which the watershed lies a little below 500 ft elevation. To the north and east, lower and more extensive valleys of the River Pahang itself and its major tributaries intervene between highland elevations on Gunong Tahan, 47 miles distant, and the Eastern Range, 52 miles distant at the nearest point. On its south-eastern quarter, a large segment of Gunong Benom extending from the lowlands to the peak is incorporated into the Kerau Game Reserve. Here, and in fact continuously over a much more extensive area of the mountain, the natural vegetation has survived inviolate and the native fauna has been largely protected from hunting and other human disturbance. Benom thus offers the rare opportunity to work in virgin forest along an altitudinal transect from the lowlands to a mountain peak that is among the highest in Malaya. |GUNONG BENOM 4 LORD MEDWAY or os rs — —— MAXWELL'S iLL —— GUNONG TAHAN CAMERON HIGHLANOS — FRASER'S HILL — ELEVATION EXCEEDING . = ; 5.000 Fr t 7] 4000 Fr \ = = \ } ones ° 20 40 60 ————— —_ = ae a 101 10s" Fic. 1. Malaya (= West Malaysia), showing the principal mountain formations. INTRODUCTION 5 Accounts of earlier ascents of Benom have been published by Barnes (1903), Evans (1924), Henderson (1927), and Strugnell and Willbourn (1931). On some of these expeditions animals were collected, and the resultant specimens have been listed or described by several authors, ¢.g., Chasen & Kloss (1928), Gibson-Hill (1949). Despite this, no previous expedition was primarily zoological in purpose. Accordingly, in early 1967, a trip was planned with the principal aim of establishing sites for zoological collection and observation along a transect from the base to the summit of the mountain. The expedition proper was in the field from 1 February to 15 April 1967. Later return trips to the area were made by several participants resident in Malaysia, as detailed in individual reports (below). Teams from the following organizations participated: British Museum (Natural History), comprising Miss A. G. C. Grandison assisted by Miss F. V. Slade; University of Malaya, Department of Zoology, com- prising Lord Medway, H. S. Yong, P. H. Soo, L. C. Ratnam, Miss S. E. Ang, Miss M. C. Leong and D. Labang; University of Malaya, Department of Parasitology, C. P. Ramachandran assisted by K. C. Lim and Poopala Chelvam; University of Malaya, Mosquitoes of Malaysia Project, O. Sulaiman and S. W. James; the Arbo- virus Research Unit of the Hooper Foundation, University of California Inter- national Center for Medical Research and Training, N. J. Marchette, R. Garcia, and D. W. MacVean, assisted by J. Jeffery, H. Lee, Miss S. K. Teh, A. H. Ahmad, Nagiah Vangitasamy, S. W. Chan, T. K. Tee and R. de Silva; Institute for Medical Research, Kuala Lumpur, B. L. Lim and M. Nadchatram, assisted by P. Ramachand- ran, K. Krishnaswamy, A. Mottan, B. Ahmad, M. Sharif, W. Mohamed, R. S. Ratnam and I. Yusof; Forest Research Institute, Kepong, Selangor, T. C. Whitmore assisted by Chelliah, I. Rahim and Y. Zahir. THE ENVIRONMENT Base Camp was established just within the Kerau Game Reserve, on the bank of a south-flowing tributary of the River Kerau, at 102°11’25"E, 3°51'30’N, at 700 ft above sea level (Grid reference 097 667 on sheet 79, Malaya 1:63,360, series L7or10). Access from the Damak-Ulu Cheka road was by an abandoned logging track. This track ran through State Forest Reserve, consisting of tall forest recently disturbed by selective commercial felling, which had been carried out as far as the camp site but had not extended across the stream (Plate 2). From Base camp the route up the mountain followed the succession of ridges that form the north-east boundary of the Kerau Game Reserve (Fig. 2). Camps and collecting areas were established along this route at the following sites: camp 2 at grid reference 081 667, 1700 ft a.s.l. (Plate 3); camp 24 at 063 666, 2500 ft; camp 3 at 055 661, 3500 ft; camp 4 at 035 653, 5000 ft; and camp 5 at 015 645, 5900 ft. The parent rock of Gunong Benom itself is an intrusive hornblende-granite, with exposures of syenite, pyroxene-granite prophyry, and diorite (Scrivenor, 1931). On the north-east approach followed by the expedition, sedimentary rocks of Triassic 6 LORD MEDWAY age, with some admixture of Quaternary volcanic effusives, extend to over 1000 ft (Alexander, 1965). At higher elevations, as far as is known, the geology of the mountain is relatively uniform and the main factors affecting the vegetation are therefore not geological but related to local variations in elevation, aspect and climate. The altitudinal zonation of the forest in which the expedition worked on the north-east slopes of Benom is described and discussed by Whitmore (1972). The route to camp 2 was a relatively easy slope following an abandoned logging track running between the trunks of standing forest giants on a broad ridge. Except after a lot of rain, it could be negotiated by Landrover, sometimes requiring the aid of chains. From camp 2, the path immediately climbed steeply to a long undulating ridge at 2400-2500 ft. At the time of the expedition, logging had not extended above 2000 ft on this slope. Camp 24 was sited at the further end of the ridge, at the foot of a second steep climb to 3500 ft and camp 3. From camp 3 the track climbed by a series of narrow undulating ridges. Camp 4 was located on such a ridge, with unstable, boulder-strewn slopes dropping away precipitously on each side. Above about 3500 ft, many large granite boulders were exposed on the hillsides. Between these, the ground surface was very irregular and in many places damp tunnels and rock shelters were formed. Camp 5 was sited under the overhang of an exceptionally large boulder, at the north-western end of an extended ridge rising to 6300 ft. Beyond this ridge, the path dropped to below 6000 ft before climbing again to the summit ridge proper. The first peak on the summit ridge rose to 6760 ft. Between this peak and the true summit (6916 ft) the path wound round, up and down a series of subsidiary crests and knolls (Plate 4). On the summit itself a survey beacon had been erected. The surrounding vegetation had been felled and, at time of our visit, the regrowth had recently been cut back. Climatically, Benom lies within the West Rainfall Region (Dale, 1959), at the northern extreme of the central south dry belt. The mean annual rainfall is inferred to be in the range 80-90 inches, distributed in two maxima (October/November and April) and two minima (July and February) each year. Rainfall records were not kept during the expedition, and nothing is known of local variations in precipita- tion on the mountain. During most of the month of March 1967, we experienced a spell of fine dry weather, which was probably not untypical. At higher elevations surface water was scarce, but streams flowing largely or completely underground were found up to 5750 ft. Air temperatures were not recorded on Benom during the expedition, and there is no meteorological station on the mountain. The equatorial position of Malaya coupled with a generally humid climate prevent wide variations in surface tempera- ture. Annual mean temperatures recorded at 58 stations in the lowlands fall within the range 77-83°F, and diurnal variations rarely exceed 22°F (Dale, 1963). At most stations there is a discernible pattern of seasonal variation in monthly mean temperature, but the annual range of this variation is small, not exceeding 3-8°F at lowland stations (4-1°F at Fraser's Hill). With increasing altitude there is a 1,000 Contour ---- Game Reserve ( elevation Boundary in feet asl) boulders np 5 was 1-westem _ dropped first peak > summit nd knolls rounding ntly been )), at the s inferred ‘ovember ords were precipita- perienced levations 1000 Contour {elevation in feet asi) ~ Track — Game Reserve Boundary KAMPONG KAMPONG © ULU CHERA KEI GAME RESERVE the expedition camp sites and route of ascent. to JERANTUT INTRODUCTION 7 regular decrease in temperature, but freezing level in the atmosphere, at 13,500— 15,500 ft, is as high again as the highest mountain, so that very low temperatures nowhere occur (Dale, 1963). At most hill stations, the range of diurnal variation is low. Most members of the expedition felt distinctly cold at camp 5, and here and at camp 4 we burned fires at night for warmth. ACKNOWLEDGEMENTS Overhead costs of the University of Malaya, Department of Zoology team, including preliminary reconnaissance and the establishment of camps and working sites, were met in chief by a grant from the British Museum (Natural History). Permission to operate within the bounds of the Kerau Game Reserve was kindly given by the State Game Warden, Pahang, who also suggested the site for Base camp, and provided generous assistance in the early stages of setting up this camp. The participation of the Arbovirus Research Unit, UC-ICMRT, was partially supported by U.S. Public Health Service Grant No. TW oo144 of the Office of International Research, U.S. National Institutes of Health and by the U.S. Army Medical Research and Development Command under Contract No. DA-49193-MD- 2931. REFERENCES ALEXANDER, J. B. 1965. Geological Map of Malaya. 6th edn. (1963). Geological Survey H.Q., Ipoh, Malaysia. Barnes, W. D. 1903. Notes on a trip to Gunong Benom in Pahang. J. Straits By., Roy. Asiatic Soc. 39 : 1-18. Cuasen, F. N. & Kioss, C. B. 1928. Birds, from Mt. Benom, Pahang; the Kledang Hills, Perak; and the islands of Penang, Tioman and Aor. J. Malayan Br., Roy. Asiatic Soc, 6 (3) : 70-75. Dare, W.L. 1959. The rainfall of Malaya: Part I. J. trop. Geogr. 13 : 23-37. 1963. Surface temperatures in Malaya. J. trop. Geogr. 17 : 57-71. Evans, I. H. 1924. An expedition to Gunong Benom. J. Fed. Malay States Mus. 12 : 1-6. Grpson-Hitr, C. A. 1949. A handlist of the birds of Malaya. Bull. Raffles Mus. 20. HENDERSON, M. R. 1927. Ona collection of plants from Gunong Benom. J. Fed. Malay States Mus. 13 : 217-227. Panton, W. P. 1965. Topography, geology and soils. Malayan Forest Records 23, pt. II, ch 2): I-21: ScCRIVENOR, J. B. 1931. The Geology of Malaya. London, Macmillan & Co. STRUGNELL, E. J. & WitLBourN, S. 1931. An ascent of Gunong Benom from Raub. J. Malayan Br., Roy. Asiatic Soc. 9 : 15-27. Wuitmore, T.C. 1972. The Gunong Benom Expedition 1967. 2. An outline description of the forest zones on north-east Gunong Benom. Bull. Br. Mus. nat. Hist. (Zool.) 23 : 9-15. Wyatt-SmitH, J. 1965 (ed.). Manual of Malayan silviculture for inland forest. Malayan Forest Records 23. THE Lorp MEDWAY GREAT GLEMHAM HousE SAXMUNDHAM, SUFFOLK, U.K. Photographs by Lord Medway PLATE, x The Damak—Ulu Cheka road, with a distant view of Gunong Benom from the north-east. PLATE 1 I 23, Zool.) ( \ nat. Hist. Mus. Bull. Br. Pe AE 2) Base camp. Note the dense stand of Mallotus paniculatus and other secondary vegetation on the edge of the clearing. Bull. Br. Mus. nat. Hist. (Zool.) 23, 1 PLATE 2 PLATE 3 The site of camp 2 at 1700 ft. Very little disturbance of the natural vegetation had occurred > , “is es 7 Bull. Br. Mus. nat. Hist. (Zool.) 23, 1 PLATE 3 PLATE 4 The summit ridge of Gunong Benom, looking west-southwest from 6760 ft towards the true summit (centre background). 4 Bull. By. Mus. nat. Hist. (Zool.) 23, 1 NAT. Hist. \ PLATE 4 =a6 pan) ¢ A | 4, ei >} SW Tn"a ; ae , aguyy { ‘4 « f aay y t ,) : A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Atison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. ; WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. Taytor, J. D., KennEDy, W. J. & Hart, A. The Shell Structure and Mineralogy — of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, — 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) — 1962-1964. (In press). , E sik a fas ie Dy WAT. HIST, a = 8 MOV [972 SO PRESENTED 4 ce ‘S Oby Liss Be . BCE Printed in England by Staples Printers Limited at their Kettering, Northants, establishment y E GUNONG BENOM EXPEDITION 1967 OUTLINE DESCRIPTION OF THE FOREST ES ON NORTH-EAST GUNONG BENOM T. C. WHITMORE BRYOPHYTES AND FILMY FERNS FROM _ GUNONG BENOM ANNE JOHNSON LP mass.” \ | ~ BNOV 972 SUPREEEAITED & Boa 9 BULLETIN OF | ISH MUSEUM (NATURAL HISTORY) VARS tee Vol. 23 No. 2 LONDON: 1972 THE GUNONG BENOM EXPEDITION 10967 2. AN OUTLINE DESCRIPTION OF THE FOREST ZONES ON NORTH-EAST GUNONG BENOM BY TIMOTHY CHARLES WHITMORE Forest Research Institute, Kepong, Malaysia 3. BRYOPHYTES AND FILMY FERNS FROM GUNONG BENOM BY ANNE JOHNSON Nanyang University, Singapore > Pp. 9--20; 8 Plates tie WAGs we NAT..HIST. —\ - 8 NOY a) S PRESENTED 4 Yo, a Ser BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No.2 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, ts 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 to 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. 23, No. 2 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 18 October, 1972 Price £1.15 THE GUNONG BENOM EXPEDITION 1967 2. AN OUTLINE DESCRIPTION OF THE FOREST ZONES ON NORTH-EAST GUNONG BENOM By T. C. WHITMORE THE north-east ridge of Gunong Benom provides in a single majestic sweep a fine locus, perhaps the best in Malaya, for the study of the change of forests with elevation. Above about rooo ft the rock is uniformly granite and the ridge is broad and more or less continuous. Therefore there are no complications intro- duced by changes in geology or by broken relief. In 1967 Kepong botanists spent about four weeks plant collecting on Benom, part of the time with the University of Malaya/British Museum (Natural History) Expedition. I myself also made an outline survey of the vegetation types. This I describe here to set the picture for the more particular studies made by the varied groups of zoologists on the Expedition. I hope that publication of this background will stimulate a forest ecologist to visit Benom to make a detailed study of the phenomenon of the altitudinal zonation of tropical forest, for which this mountain is ideally suited. Tropical rain forest altitudinal zones may be classified in two ways. Firstly we may distinguish Rain Forest Formation Types, defined on structure and physiog- TABLE I ALTITUDINAL FOREST ZONES IN MALAYA (AFTER SYMINGTON, 1943) World-wide structural and Malayan floristic zones Elevation physiognomic zones Lowland Dipterocarp c. 1000 ft Lowland Rain forest Hill Dipterocarp c. 2500 ft Upper Dipterocarp c. 4000 ft Lower Montane Rain forest Montane Oak —_—_—_—_—————— _ ¢. 5000 ft Montane Ericaceous Upper Montane Rain forest Bull. Br. Mus. nat. Hist. (Zool.) 23, 2. At 12 T. C. WHITMORE nomy, and recognizable throughout the humid tropics. Secondly we may define zones on flora; these are consequently restricted to a particular country or region. These two classifications run together, as set out in Table 1. Further description of the Formation Types is given in Grubb ef al. (1963). The Malayan floristic zones were described and defined by Symington (1943), and have been redescribed and amplified several times by Wyatt-Smith (notably Wyatt-Smith, 1964). The forest at about 7oo ft around Base camp is Lowland Rain forest, and floristically is very rich Lowland Dipterocarp forest. There had been selective logging of the most valuable timber, including many of the dipterocarps, a few months before we visited Benom, so it is no longer easy to give a floristic account of the area. Logging did not extend west of the river Kerau. The area east of the Kerau had relict tall trees isolated or in groups, and the main and lower parts of the forest canopy was variously broken or disturbed by extraction tracks etc. Before disturbance the original forest had been magnificent, as witnessed by the still standing trees of Koompassia excelsa (Leguminosae), with its fluorescent green, smooth trunk and cauliflower-shaped crown, reaching 150 to 200 ft or even taller (Plate 5). Here we felled a relict 150 ft tree which proved to be Kibatalia arborea, only otherwise known in Malaya from a few trees at Kepong and one in Ulu Langat. Further details of the flora of this zone can be found through Appendix il In the open places left by timber extraction, for instance around the camp, a dense stand of Mallotus paniculatus (Euphorbiaceae) had grown up with some Sapium (Euphorbiaceae) and some Macaranga gigantea (Euphorbiaceae). In other places dense thickets of gingers and wild banana had sprung up along extrac- tion tracks (Plate 6). The unfelled forest across the stream was typical of many valley bottoms, some high forest with riverine species, e.g., Dillenia reticulata, and some scrub and climber thickets. There was little bamboo. An old Rafflesia hasseltii flower was found in one scrub thicket. South of the camp were patches to an acre or so in extent of the scrambling fern Gleichenia (s.l.) with secondary forest trees growing through; these areas are probably sites of former aborigine cultivation. Although all human settlements were moved a few years ago to Kuala Lompat, outside the Game Reserve, the map given by Ogilvie (1949) shows both Negrito and Jah Chong people on north-east Benom, Similar Gleizchenia patches and rattan tangles on the steep west slope of the Kerau valley have probably resulted spontaneously from landslips. Felling had continued, but less intensively, on the hill slope up to camp 2 at 1700 ft. Larger patches of original forest remained, and the track from Base camp ran amongst standing forest giants (Plate 7). Along this track and above camp 2 felling had been restricted to the north side, outside the Game Reserve. Since the Expedition was in the field, extensive felling in 1968 has covered a large area here, up to and abutting on the Game Reserve boundary along the main north-east ridge, and there has been some encroachment. The undisturbed forest begins almost at once above camp 2. The ridge to camp 2} THE FOREST ZONES ON NORTH-EAST GUNONG BENOM 13 (2500 ft) carries very fine Hill and Upper Dipterocarp forest. Shorea curtisit (Dipterocarpaceae) is prominent, associated with Shorea maxwelliana and a Swintonia (Anacardiaceae). Isolated big trees of the conifer Agathis dammara were scen. Eugeissona tristis, a big undergrowth palm commonly associated with S. curtisiz, is absent. Between 2500 ft and 3500 ft a sharp and almost total change in flora occurs. This is associated with a change to Lower Montane Rain forest, within which camp 3 lay. The change in forest type can be clearly seen on aerial photographs. At camp 3 the forest was described as follows* (and following the scheme in Table 10 in Grubb e¢ al., 1963): 3500 ft Broad ridgetop Canopy: 70-80 ft, the biggest trees 8-9 ft girth, with tall straight boles, amongst them Adinandra (Theaceae), Santiria rubiginosa (Burseraceae) and Tristania (Myrtaceae) prominent (Plate 8). Climbers: Photophytes: none seen. Skiophytes: Freycinetia javanica (Pandanaceae) and another Freycinetia sp. frequent. Epiphytes: Vascular: infrequent; non-vascular: 10% thin cover on tree trunks; 20%, or rarely up to 90%, cover up to 6 ft on butts. Soil: Continuous leaf litter, over a spongy 6 in. peat layer itself overlying a coarse yellow sandy clay. The presence of a peat over the mineral soil at an elevation as low as this is noteworthy. Peat was noticed on G. Benom from here to the summit. At camp 4, 5000 ft, it was about 1 ft thick. The boundary to Lower Montane Rain forest is sharp on G. Benom. With the Lower Montane formation there are local differences in flora. Thus about } mile above camp 3, at about 3800 ft where the ridge swings southwards and becomes narrower, the species composition changes and many of the species one sees on the Main Range, e.g. at Fraser’s Hill come in. Prominent in the canopy here due to its velvety brown leaves is the new and beautiful Palaquiwm regina-montium (Sapota- ceae), an endemic Malayan species now known from several mountains at about this elevation. The conifer Dacrydium comes in, as do Symingtonia populnea (Hama- melidaceae) and Weinmannia blumei (Cunoniaceae). The giant mountain rattan Plectocomia griffithii, was seen only in this zone. From about 4000 ft upwards the tallest forest (canopy about 50 ft) with the biggest trees is on broad ridge-crests and on saddles. Narrow exposed ridges carry low Upper Montane Rain forest with gnarled trees. Lower Montane Rain forest persists on gentle slopes and protected valleys to high elevations. In a broad valley about } mile south-west of camp 4 at about 4800 ft the flora contained several lowland elements, including new altitude records by a few hundred feet in Malaya for Knema oblongifolia v. monticola (Myristicaceae) and Payena lucida (Sapotaceae). This valley has a grove of the rare Malayan Maple Acer caesium. The boundary between Lower and Upper * No species list is given, to avoid giving a misleading impression of completeness. Species found at this elevation can be ascertained through appendix 1. 14 T. C. WHITMORE Montane forest, with the former growing high up valleys, is clearly visible on aerial photographs. Most valley sides and ridge slopes are not gentle, but precipitous and unstable with evidence of land slips. In some places huge boulder piles cover the slopes with caves between and under them. Trees are few and rather small and the under- growth is very dense with many palm and gingers (Plate 9). Fallen trees are abundant. Many characteristic trees are virtually exclusive to the ridges and do not grow on the unstable boulder littered slopes. Examples are Ternstroemia FRI 3320, Dacrydium comosum and D. elatum and Podocarpus imbricatus (Coniferae) and amongst the smaller trees the very lovely pink-flowered Rhododendron wrayt. With increasing elevation one passes through less Lower Montane forest; nearly everywhere is clothed in the Upper Montane formation whose facies and probably species composition varies from place to place, notably with relief. In places Baeckia frutescens (Myrtaceae) and Leptospermum flavescens (Myrtaceae) (Plate 10) are abundant. The deep gully at c. 6000 ft below and to the south-west of the first peak on the summit ridge has vestiges of Lower Montane forest. It is the last station of the common undergrowth montane palm Pinanga polymorpha and of the rattans (a slender Calamus). The first peak, 6760 ft, has an open stunted Upper Montane forest, (Plate 5) almost a pure stand of 15-20 ft Leptospermum flavescens (Plate 11) with a little Dacrydium over a grassy, bushy undergrowth in which the trunked sedge Gahnia baniensis and Rhododendron obscurum are conspicuous (Plate 12). Open turfy places carry Gentiana malayana (one of the North Temperate—Himalayan plants which reach into Malesia on high mountains), and the small sedges Lepidosperma chinense and Oreobolus kukenthallii. The ground is covered by thick peat in which the bogmoss Sphagnum junghunianum is prominent. © The flora is much less rich than the similar formation on G. Tahan, with which (and probably other high peaks) it shares most of its species. Upper Montane Rain forest is characterized inter alia by abundant epiphytes. These are far less well developed on Benom than is common. Orchids are un- common. Above c. 3500 ft tree boles do have a bryophyte layer but it is only c. 1 in. thick. Filmy ferns are inconspicuous. There are no great festoons from the trees even in the moist gullies. The exposed first summit peak has very little bryophyte development indeed. Slight development of epiphytes and skiophytic climbers as compared with other humid tropical countries seems to be a widespread feature of montane forests of Malaya (see Whitmore & Burnham, 1969, describing G. Ulu Kali in the Main Range). It may reflect the lower rainfall here than elsewhere. G. Benom, lying in the rain shadows of both the Main Range and the east coast mountains, is in a dry part of Malaya and this probably is an important factor limiting the epiphyte flora. REFERENCES Grup, P. J., Ltoyp, J. R., Pentncron, T. D. & WuitmorE, T. C. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy and floristics. J. Ecol. 51 : 567-601. THE FOREST ZONES ON NORTH-EAST GUNONG BENOM 15 OaitveE!I, C.S. 1949. Che Wong word list and notes. Bull. Raffles Mus. (Series B) 4 : 11-43. SyMINGTON, C. F. 1943. Foresters’ manual of dipterocarps. Malay. For. Rec. 16. Wuitmore, T. C. & BuRNHAM, C. P. 1969. The altitudinal sequence of forests and soils on granite near Kuala Lumpur. Malay. Nat. J. 22, 99-118. Wyatt-SmITH, J. 1964. A preliminary vegetation map of Malaya with descriptions of the vegetation types. /. trop. Geogr. 18 : 200-213. APPENDIX x Previous expeditions to Benom have either been by a botanist or with a botanist, and several species lists have been published with the accounts of the ascents to which Medway has referred in his introduction. Our collections, 408 numbers, are the first from the north-east side of the massif, and in total are probably the biggest made on the mountain. We do not propose to publish a list of the species collected. They are being named in the normal course of our research work, and duplicates have been distributed to our usual correspondent herbaria: Arnold Arboretum, Kew, Leiden, and Singa- pore. We keep a herbarium register at Kepong from which at any time the identities of the plants collected may be ascertained. It is thus sufficient to quote the numbers of the plants which were collected on the mountain, and interested persons can then discover identifications from the register. FRI 3126-3143 FRI 3150-3445 Coll. T. C. Whitmore FRI 3997-4000 KEP 97801-97850 > KEP 99783- Coll. Rahim bin Ismail KEP LOOLOI-10012g KEP 99101-99150 Coll. Zahir bin Yusof KEP —__104301-104313 alc KEP 104382-104450 Sf Coll. Chelliah Dr. T. C. WHITMORE, F.L.S. ForREST RESEARCH INSTITUTE KeEponc, MALaysiIa April, 1968. dua THE GUNONG BENOM EXPEDITION 1967 3. BRYOPHYTES AND FILMY FERNS FROM GUNONG BENOM By ANNE JOHNSON THE following is a short account of bryophytes and filmy ferns collected on Gunong Benom from 7-9 April 1967. Apart from three collections by J. A. Bullock at 6000 ft, all specimens were collected between 800 and 3800 ft, from the ground, rock surfaces, and lower tree trunks, where possible up to about 6 feet above ground. All specimens have been lodged in the Cryptogamic Herbarium, School of Biological Sciences, University of Malaya. The accession number is given for each collection. The material confirms to a certain extent Whitmore’s general remarks on epiph- ytes (1972). The extreme poverty of the bryophyte flora in the lowland forest at 800 ft reflects the effect of rain shadow. There is a sharp change in the bryophyte flora at about 2500 ft, especially in valleys. Above this elevation there is a marked increase in both number of species and their abundance, with up to 100% cover in wet places near the ground. There is little distinction between ground and tree bryophytes. Peat found at 3800 ft is not sphagnum peat, but is composed of dead bryophytes other than sphagnum and vascular plant debris. Sphagnum junghuh- mianum occurs at higher elevations (Whitmore, 1972). The flora is similar to that of lowland forest and lower montane forest in other parts of Malaya but is noteable for the paucity of members of the Leucobryaceae and Calymperaceae which are far better developed in other areas. SYSTEMATIC LIST Dilaenaceae Pallavicinia indica Schffn. On small stones near the stream, Base camp, 800 ft (B 0964). Pallavicinia leveiri Schffn. On tree trunks in valley, 2800 ft (B 0950). Metzgeriaceae Metzgeria hamata Lindb. On fallen trees in valley, 2800 ft (B 0957). Riccardiaceae Riccardia pinguis (L). Gray. In mossy forest, 3800 ft (B 0959). Riccardia parvula Schffn. On palm inflorescence in valley, 2900 ft (B 0888). Bull. Br. Mus. nat. Hist. (Zool.) 23, 2 17 18 ANNE JOHNSON Jungermaniales Acrogynae Ptilidiaceae Trichocolea tomentella Huds. On ground, Base camp, 800 ft (B 0go2). Cephaloziellaceae Cephaloziella sp. Abundant, in mossy forest, 3800 ft (B 0923, B 0939, B og4o, B 0941). Bazzaniaceae Bazzania distans (Nees.) Trev. With seedlings of Podocarpus, 6000 ft, collected by J. A. Bullock (B 0899). Bazzania longicaulis (Lac.) Schffn. In mossy forest, 3800 ft (B 0916, B 0929). Lejeuneaceae Schistochila sp. In mossy forest, 3800 ft (B 0923, B 0927, B 0956). Stictolejeunea sp. At Base camp, 800 ft (B 0962). Musci Dicranaceae Dicranoloma blumii (Nees.) Par. In mossy forest, 3800 ft (B 0915). Leucobryaceae Leucobryum candidum (P. Beauv.) Wils. In mossy forest, 3800 ft (B og12, B 0955). Leucophanes octoblepharoides Brid. On trees at Base camp, 800 ft (B 0984); in valley, 2800 ft (B 0890, B 0896). Syrrhopodontaceae Syrrhopodon croceus Mitt. In mossy forest, 3800 ft (B 0924). Syrrhopodon schiffnerianus (Fleisch.) Par. In mossy forest, 3800 ft (B 0931, B 0945). Syrrhopodon spiculosus Hook. at Grev. In mossy forest, 3800 ft (B 0887, . B 0910, B og14, B 0922, B 0928, B 0958). Syrrhopodon treubit Fleisch. In valley, 2800 ft (B 0893, B 0898); in mossy forest, 3800 ft (B 0921). Thyridium undulatum (Broth. et Geh.) Fleisch. In valley, hanging from the trees, 2800 ft (B 0880). BRYOPHYTES AND FILMY FERNS FROM GUNONG BENOM 19 Calymperaceae Calymperes hampei Doz. et Molk. On trees, Base camp, 800 ft. (B 08809). Calymperes longifolium Mitt. In isolated patches on trees in the valley, 2800 ft (B 0909). Orthotrichaceae Groutiella goniorrhyncha (Doz. et Molk.) Wijk. et Marj. On trees at Base camp, 800 ft (B 0906). Rhizogoniaceae Rhizogonium spiniforme (Hedw.) Bruch. On ground in valley, 800 ft (B 0946). Meteoriaceae Aerobryopsis longissima (Doz. et Molk.) Fleisch. In valley, 2800 ft (B ogor); in mossy forest, 3800 ft (B 0933). Neckeraceae Thamnium ellipticum (v.d.B. et Lac.) Kindb. On tree trunks at Base camp, 800 ft (B 0984). Thuidiaceae Thuidium meyenianum (Hpe.) v.b.B. In the valley, 2800 ft (B 0895). Hookeriaceae Callicostella pappillata (Mont.) Mitt. On tree bark, Base camp, 800 ft (B og11); valley on trees, 2800 ft (B 0952). Distichophyllum mittenii Bosch. et Lac. In mossy forest, 3800 ft (B 0944). Sematophyllaceae Clastobryophilum bogoricum (Bosch. et Lac.) Fleisch. Very abundant in mossy forest, 3800 ft (B 0947, B 0951, B 0953, B 0966). Acroporium diminutum (Brid.) Fleisch. On trees in the valley, 2800 ft (B 0892). Sematophyllum saproxylopilum (C. Muell.) Fleisch. In the valley, 2800 ft (B 0897). Trichosteleum mammosum (C. Muell.) Jaeg. In the valley, 2800 ft (B 0961). Trismegista lancifolia (Harv.) Broth. In mossy forest, 3800 ft (B 0886, B 0930, B 0935). Hypnaceae Ectropothecium intorquatum (Doz. et Molk.) Jaeg. In the valley, 2800 ft (B 0963). 20 ANNE JOHNSON Hypnodendraceae Hypnodendron reinwardtii (Reinw. et Hornsch.) Lindb. In mossy forest, 3800 ft (B og19, B 0954, B 0960). Filicinae Hymenophyllaceae Hymenophyllum acanthoides (v.d.B.) Rosenst. In peaty area, 6000 ft, collected by J. A. Bullock (B 0903). Trichomanes meifolium Bory. On exposed slope, 6000 ft, collected by J. A. Bullock (B ogo0). REFERENCE WuirmoreE, T. C. 1972. An outline description of the forest zones on north-east Gunong Benom. Bull. Bry. Mus. nat. Hist. (Zool.). 23: 9-15. Professor ANNE JOHNSON Department of Biology NANYANG UNIVERSITY SINGAPORE 22 Photographs by Lord Medway PLATE 1 < A fine tualang (Koompassia excelsa), temporarily deciduous, standing alongside the track to” Base camp. I ATE PL Mus. nat. Hist. (Zool.) 23, 2 By. Bull. PLATE & A disused extraction track at rooo ft. Note the height of standing timber (compare the figure of a man, arrowed), and the dense thickets of young vegetation springing up on both sides. ATE Pie > 235 nat. Hist. (Zool.) Mus Bull. Br. i sane) NG. PLATE 3 : The track to camp 2, at 1600 ft, running among relatively undisturbed forest vegetation. The vertical scale is indicated by the figure of a man (arrowed). Bull. Br. Mus. Nat. Hist. (Zool.) 23 PLATE 4 Lower Montane Rain forest near camp 3. n Oa) a N = x 4 oe nt ging Page Mus Bull. Br. PLATE 5 Ridge vegetation at 4700 ft. Note the figure of a man (arrowed). Bull. Br. Mus. nat. Hist. (Zool.) 23, 2 PLATE 6 Blossoms of Leptospermum flavescens, photographed near the summit of Benom, a a a N = <= ~~ 5 Bull. Bi PLATE 7 A stand of Leptospermum flavescens on the north-east end of the summit ridge, at 6760 ft. > 235) Mus. nat. Hist. (Zool.) Br. Bull. IPILJNINIS, &) The undergrowth of sedge and low bushes between Leptospermwm trees at 6760 ft. Bull. By. Mus. nat. Hist. (Zool.) 23, 2 = NAT. Hist; - 8 NOV 972 PLATE 8 } A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Artson. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier eri Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. TayLor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In Press). ia se ‘ ¢ » NAT. HIST. - 8 NOV 1972 si, PRESENTED & sy ogy wast al Pe Sat Printed in England by Staples Printers Limited at their Kettering, Northants, establishment , GUNONG BENOM EXPEDITION ey 1967 { RECORDS OF MALAYAN BATS, WITH JOMIC NOTES AND THE DESCRIPTION BULLETIN OF SH MUSEUM (NATURAL HISTORY) ee Vol. 23 No. 3 - LONDON: 1972 Fy THE GUNONG BENOM EXPEDITION 1967 4. NEW RECORDS OF MALAYAN BATS, WITH TAXONOMIC NOTES AND THE DESCRIPTION OF A NEW PIPISTRELLUS BY JOHN EDWARDS HILL Pp. 21-42 2 ; NAT. Hist, ~ \ (2 1 cee it \ : t 2 | 3 PRESENTED my 0 ny L0G, pst BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 3 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), tistituted 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 ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 23, No. 3 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 8 December, 1972 Price 85p THE GUNONG BENOM EXPEDITION 1967 4. NEW RECORDS OF MALAYAN BATS, WITH TAXONOMIC NOTES AND THE DESCRIPTION OF A NEW PIPISTRELLUS By J. E. HILL THE 1967 Expedition to Gunong Benom, Pahang, 1965 resulted in the accession of a number of specimens of bats of especial taxonomic interest, including one example that has proved to represent an undescribed species of Pipistrellus. Addi- tionally, during recent years, specimens of similar interest or in some cases represent- ing species new to Malaya have been received from Lord Medway, formerly of the Unit of Zoology, School of Biological Sciences, at the University of Malaya, Kuala Lumpur. All measurements in this paper are in millimetres : where these refer to a series of specimens the minimum and maximum values are given, followed by the mean in parentheses. Rhinolophus malayanus Bonhote, 1903 2° B.M. 68.812 (in alcohol, skull extracted). Wang Tangga, Kaki Bukit, Perlis, 6° 39’ N, 100° 12'E. 2 January 1968. 6 B.M. 68.816, 92 B.M. 68.813-815 (all in alcohol, skulls extracted). Kisap Forest Reserve, Pulau Langkawi, Kedah, 6° 23’ N, 99°52’ E. 31 December 1967. Rhinolophus malayanus is not listed from the States of Malaya by Chasen (1940 : 36) and does not appear to have been subsequently recorded. Described originally from Biserat, Patani, southern Thailand, its presence in Perlis and on Pulau Lang- kawi is not unexpected. The species is recorded from Koh Lak, on the east coast of southern Thailand rather to the north of Patani by Gyldenstolpe (1916 : 13), from the Laos Mountains by Andersen (1905a : 89) and from Laos by Tate and Archbold (1939 : 6) and by Phillips (1967 : 634), while Osgood (1932 : 218) records it provisionally from Muong Moun, Tonkin (North Vietnam). These specimens from Perlis and Langkawi agree closely with the type specimen, having the lancet with slightly concave lateral margins and an elongate tip and with the lateral margins of the sella almost parallel. The connecting process is rounded and rises slightly above the tip of the sella: its point of insertion on the 24 JE. oiew posterior leaf is a little above the corresponding insertion on the back of the sella. The small anterior premolar (pm2) is in the toothrow but much reduced: the second lower premolar (pms) is slightly extruded or is external to the toothrow, with pme and pmg sometimes almost in contact. MEASUREMENTS (B.M. 68.812, 68.816, 68.813-815, in that order) : length of forearm 40-3, 39°6, 42:2, 41-2, 40-7; width of sella, 2-2, 2-0, 2-1, 2-0, 1-9; greatest length of skull 18-2, 18-5, 18-0, 17-9, 18-0; condylocanine length 15-3, 15-5, 15:2, 15°2, 15‘; supraorbital length (distance from junction of supraorbital ridges to median anterior point of nasal swellings) —, 4-9, 4:9, —, 4:8; width of nasal swellings —, 4-9, 5-1, 5:1, 5:0; zygomatic width 8-8, 8-9, 8-8, 8:5, 8-8; least interorbital width —, 2-2, 2°3, 2-3, 2:4, width of braincase 7-4, 7:4, 7:4, 7:4, 7:4; mastoid width 8-4, 8-4, 8-2, 8:3, 8-3; clcl 4:4, 4°5, 4-4, 4-2, 4-3; m3m8 6-7, 6:5, 6:5, 6-4, 6-3; c-m 68, 6:8, 6-6, 6-5, 6-6; length of mandible 11-3, —, —, —, 11-4; c—mg 6-9, 7:2, 6:9, 6°8, 68. Rhinolophus malayanus is very similar to R. borneensis, differing chiefly in the size and shape of the median anterior nasal swellings, which in malayanus are more inflated, higher, wider and more clearly demarcated than in borneensis, extending laterally down the sides of the rostrum and thereby reducing the area of the lateral swellings as compared with that species. Robinson and Kloss (1915 : 116) recorded as R. borneenstis (2?) a specimen from Khao Nawng, Bandon, southern Thailand but this subsequently became the type specimen of FR. vobinsoni Andersen (1918 : 375). An examination of the type specimen of R. chasent Sanborn (1939 : 38) from Con Son Island (Pulo Condore), off the southeast coast of Indochina indicates that chasent should be considered a subspecies of R. borneensis rather than related to R. malayanus as was suggested by its describer or conspecific with it as thought by Phillips (1967 : 634) from the published description. Rhinolophus robinsoni Andersen, 1918 A single specimen from Fraser’s Hill, Pahang (see below) resembles the nominal species robinsoni and klossi in its major features and although in some respects it differs slightly from these, in others it connects them. They have never been described in detail and rest on brief diagnoses by Andersen in a key published (on behalf of Andersen by Oldfield Thomas) in 1918. The examination of a limited number of specimens suggests that they are conspecific: vobinsoni has line priority and becomes the specific name. The species is relatively small with large, bluntly pointed ears; prominent anti- tragal lobe; large anterior leaf covering the muzzle; lancet cuneate; sella wide, its central part (excluding the basal lobes) expanded laterally so that its width at this point nearly equals or is greater than the width at its base, but sharply constricted a little above its central point; upper margin of sella slightly convex; connecting process low and rounded, its insertion on the posterior leaf level with or beneath its MALAYAN BATS AND A NEW PIPISTRELLUS 25 insertion on the back of the sella; third, fourth and fifth metacarpals subequal; second phalanx of third digit short, its length less than one and one half times the length of the first phalanx; braincase elongate with low sagittal crest ; supraorbital region relatively long, the supraorbital length (from junction of supraorbital crests to median anterior point of nasal swellings) exceeding the width across the lateral anterior nasal swellings; a clearly defined frontal depression; prominent nasal swellings, the median anterior pair high and globular; mesopterygoid fossa wide with deep mesopterygoid pit; anterior upper premolar (pm?) in the toothrow, usually jn contact with canine and posterior premolar (pm4); second lower premolar (pms) small, partially or wholly extruded, with pmg and pmy sometimes in contact or nearly so. Measurements appear in Tables 1 and 2. The external and cranial features of R. vobinsoni refer it to the ferrumequinum group of Tate and Archbold (1939 : 2) (the s¢mplex group of Andersen (1905a : 75, 1905b : 648) or megaphyllus group of that author (1918 : 374). Its wing structure places it in either the s¢mplex (more properly the megaphyllus) or borneensis sub- groups of Tate and Archbold! and not in the affinis subgroup as these authors defined it and in which they listed both vobinsoni and klossit. The long supraorbital region of vobinsoni is characteristic of the simplex subgroup but its rounded sella, short palatal bridge and extruded second lower premolar ally it more closely to the borneensis subgroup and the species is evidently closely related to R. borneensis which in many respects it resembles. However, in R. borneensis the sella is narrower with its lateral margins parallel or nearly so and the connecting process is inserted on the posterior leaf at a point above its insertion on the back of the sella. The supraorbital region is shorter and the median anterior nasal swellings in borneensis less globular and wider, extending laterally to the sides of the rostrum. Rhinolophus robinsoni robinsonit Andersen, 1918 Rh{inolophus] vobinsoni Andersen, 1918 : 375. Khao Nawng, Bandon, Lower Thailand. Type specimen B.M. 18.8.2.1 (in alcohol (in bad condition), skull extracted). Larger than R.7. klossi with slightly narrower lancet; width of sella as in klossz, not greater as stated by Andersen; palate wider; mesopterygoid fossa wider anter- iorly ; sphenoidal bridge very slightly wider; second lower premolar (pmg) completely extruded with pmz and pm, in contact. Rhinolophus robinsoni klossi Andersen, 1918 Rh{inolophus] klossi Andersen, 1918 : 375. Pulau Pemanggil, Johore Archipelago, off east coast of Malaya. Type specimen B.M. 18.8.5.5 (in alcohol, skull extracted). June 1915. (?) ¢ B.M. 67.214 (in alcohol, skull extracted). Pine Tree Hill, Fraser’s Hill, Pahang, 3° 43’ N, 101° 42’ E, at 4500 ft. 10 October 1966. _ + Tate and Archbold mistakenly noted (p. 5) that the second phalanx of the third digit is unshortened in the simplex and borneensis subgroups and shortened in the affinis and ferrumequinum subgroups: in fact, the reverse is true as indicated by Andersen (19052). 26 Yo 1D NABI Smaller than R.v. vobinsoni with lancet slightly wider at its base; palate narrower ; mesopterygoid fossa narrower anteriorly; sphenoidal bridge narrower; second premolar (pms) partially or wholly extruded, pmg and pmg separated by a narrow space or in contact. One example (B.M. 55.1479, from Pulau Pemanggil) has the right pmg completely extruded but the left pmg has been lost and pmg in this ramus remains wholly in the toothrow. Specimens from Pulau Pemanggil and Pulau Aor have relatively small skulls: one specimen from Pulau Tioman has a slightly longer forearm and a larger skull approaching the skull of vobimsont in size but nevertheless this example retains the narrow palate, narrow mesopterygoid fossa and narrow sphenoidal bridge of Aloss?. The specimen from Pahang is referred to klossi with some reservation. Externally, it almost exactly resembles this subspecies, but although its skull is similar to that of klossi, in size it approaches or in some respects exceeds the skull of robinsont and the supraorbital region is longer and the mesopterygoid fossa narrower than in either of these. The material available, however, is totally inadequate for any attempt at evaluating the status of the northern and southern populations of R. robinsoni on the Malay Peninsula. TABLE I EXTERNAL MEASUREMENTS OF Rhinolophus vobinsont Ry. klossi —_— o o Sie SS v= =! pe ae eee ee : we na ag osg 4 aa > Sve of TRS ke 2 Se S8q 2 oO HO 1 oO HO ii 2 Sod Suess Soe Sa oS ene -s mas! si os ss i 8 Sie VA52 25 AS 23 Ss Se eee yon cay ma cofet ma aes mane Length of forearm 45°3 41°8 40°9 —_— 42°2 43°6 41°8 IlIrd. metacarpal 34°0 Suey 30°6 kaon D 31-0 32°8 31°3 Iii 14°5 12°8 1795} itg}O9/ 12°8 13°3 HAO) Ii12 19°7 1/7/40 16°8 18-3 18-1 I9°I 18-3 IVth. metacarpal 33°8 32°0 -- 33°1 32°1 34°1 32°9 Ivt II‘4 9°5 Q°2 10-0 9°3 10'3 O°5 Iv2 12:1 10°3 Io'l DT Il"4 Ir°5 TD2 Vth. metacarpal 33°8 31°8 31°1 B21 BO” 34°2 32°0 vi II‘5 10-0 9°8 10°7 o'7 1074 9°8 v2 I2°5 10-8 10-2 nue} 10°9 II‘2 II°4 Width of sella 2°7 2°5 2:8 2-7 3°0 2:9 2:8 Length of tibia 19°4 16°8 16°3 16-8 17°5 17°6 18-3 MALAYAN BATS AND A NEW PIPISTRELLUS 27 TABLE 2 CRANIAL MEASUREMENTS OF Rhinolophus robinsoni Rv. klossi _— asses ae Be _ I t= Os Scie Bae 5 Sires peugae Ord ued) | GS auy se SUE Or ee ats Gy, Re esis Be 2o Ho Ho HO Gh} SSG ete se) eee eee oS aos Sf dl & - BI ers fat} a) Bes See Sc ce Se se Se Brie eon Mh Me ma Me MA RAMs Greatest length of skull = = = LS25 LO) ETOes 20°3 Greatest length of skull to canine 19:2 — ae 18-0 17°5 19:0 I9*2 Condylocanine length 17-2 —_— —_— 16-0 15°5 16°9 17'0 Length of palatal bridge 2:0 2-0 2-0 2-0 18 290 2-1 Supraorbital length 5°3 5:2 Ges 5°3 5°2 5°5 6:0 Width of nasal swellings 572 5'0 4°9 5:0 5°0 5'2 5°5 Zygomatic width OF 9°3 g°2 9°4 9-1 @)°5)} C7 Least interorbital width 2:6 2°5 2°4 2:13 2:6 2°5 2:6 Width of braincase — — — — 7°4 79 8-4 Mastoid width Qr2 — — — 8°5 8-9 9:2 Width of mesopterygoid fossa Bie2 2:8 2:8 2:9 2:9 2°9 2°77 Width of sphenoidal bridge 1605) I-5 I°*4 I°5 I°4 1-4 I°5 cicl 4°6 Ape a4 ome AOL 43 44 4°6 m3—m3 6:9 6°5 6°5 6:6 6°4 6:6 6:8 c-m3 Gf. 6-8 6-6 6°8 6-6 722 FOG} Length of mandible I2°5 — II°5 — — 12°3 I12°8 c-m3 7°6 7°3 Tee Wa3 GO Yoo 7°8 Rhinolophus macrotis dohrni Andersen, 1907 6 B.M. 67.1595 (in alcohol, skull extracted), 9 B.M. 67.1596 (skin and skull). Base Camp, Gunong Benom, Pahang, 3° 51’ N, 102° 11’ E, 700 ft. 20-23 February 1967. : 6 B.M. 67.1597 (in alcohol). Above Camp 2, Gunong Benom, 2500 ft. 7 April 1967. 33 B.M. 67.1598-1598A (in alcohol). Camp 3, Gunong Benom, 3500 ft. 7-8 March 1967. 3d PM 45, PM 47-48 (skeletons), B.M. 67.1599 (in alcohol), PM 43 (in alcohol, returned to the University of Malaya). Camp 3, Gunong Benom, 3600 ft. Io-11 March 1967. Rhinolophus macrotis has not previously been recorded from the States of Malaya although known from Sumatra (R.m. dohrni) and, following Tate (1943 : 2), from Thailand (R.m. siamensis), Fukien (R.m. caldwelli), Szechuan (R.m. episcopus), Nepal (R.m. macrotis) to the Philippines. However, according to Osgood (1932 : 219), 28 Yo De Tues, caldwelli and siamensis are apparently sympatric in Tonkin (North Vietnam) but the specimens he refers to siamensis are intermediate in size between caldwelli and the very much smaller siamensis. Specimens from Gunong Benom agree very closely with R.m. dohrni but are very slightly larger: among Malayan Rhinolophus the species may be readily recognized by its cuneate lancet, very broad, tongue-shaped sella which has a light covering of hairs on its anterior face, and low connecting process which originates below the apex of the sella. Cranially it is distinguished by its long palatal bridge and by the anterior upper premolar (pm?), which has a prominent, inwardly deflected cusp and which stands in the toothrow and is divided from the canine and posterior premolar (pm4) by small interspaces. Measurements of seven specimens in alcohol and five skulls (except where stated): length of forearm 44-5-45°3 (44-9); greatest length of skull 19-1-19-5 (19:4); greatest length of skull to front of canine 18-5-18-7 (18-6); condylocanine length 16-3-16-7 (16-5); basilar length to front of canine 14-6-14-9 (14:8); width of nasal swellings 5-0-5-2 (5-1); zygomatic width 8-3-8-6 (8-5); least interorbital width (four skulls 2-4-2-6 (2:5); width of braincase 7-5-7-8 (7-6); mastoid width 8-8—g-2 (8-9); cl-cl 4-0-4°3 (4:1); m3-m3 5-7-5-9 (5-8); c-m® 6-6-6°8 (6-7); length of mandible (two skulls) 11-8; c-mg 6-8-7-0 (6:9). Hipposideros lylei Thomas, 1913 32 B.M. 64.831-832 (in alcohol, skulls extracted). Wang Tangga, Kaki Bukit, Perlis, 6° 39'N, 100° 12’ E. 7 March 1964. Chasen (1940: 42) listed H. lylei from the States of Malay, apparently on the basis of two skins in the collection of the National Museum, Singapore from Bukit Chint- amani, Pahang (Medway, in litt.). Described originally from Thailand, its occurrence in the northern part of the States of Malaya is not unexpected. Measurements (B.M. 64.831-832): length of forearm 78-2, 75-0 greatest length of skull 30-2, 29:0; condylobasal length 26-8, 25-5; condylocanine length 26-3, 25-3; rostral width 9-4, 9:3; zygomatic width 15-9, 15-9; least interorbital width 4-4, 4-2; width of braincase 12-0, 11-9; mastoid width 14:2, 14-1; cl-cl 7-3, 7:1; m$-m$ II-0, 10-7; cm? 11-3, 11-0; length of mandible 19-9, 18-9; c-mg 12-4, 12:0. Coelops frithii Blyth, 1848 gd B.M. 67.215 (in alcohol, skull extracted but fragmentary). 16th. mile, Ulu Gombak, Selangor, 3° 18’ N, ror° 43’ E. 27 October 1966. This specimen constitutes the first definitive record of the species from the States of Malaya. Bats of the genus Coelops are rare in collections and its taxonomy seems uncertain: Tate (1941a : 5) reviewed it briefly and summarized its history but no subsequent work has appeared. The discovery of C. frithii in Malaya has led to an MALAYAN BATS AND A NEW PIPISTRELLUS 29 examination of the material of Coelops in the collections of the British Museum (Natural History), which, although quite inadequate for any general revisionary treatment, has sufficed to show that two distinct species exist in that country and which has suggested the likely allocation of most of the remaining named forms. The Malayan specimen agrees closely with C. frithii which may be characterized by: elongate, narrow outline of the lappets projecting from the supplementary leaflets flanking the anterior leaf; narial part of the anterior leaf not especially depressed and not sharply demarcated from the intermediate part of the leaf, the intervening ridges extending from the central process or ‘sella’ toward the lateral margins of the leaf low and indefinite; posterior part of the upper surface of the rostrum sloping anteriorly; rostral inflations low, separated by a shallow sulcus, in profile forming only a slight convexity ; a definite pocket or recess within the maxillary root of the zygoma, the upper edge of which is shallowly curved anteriorly to ter- minate on the rostrum above the anterior face of the second molar ; upper tooth- tows convergent anteriorly; posterior lower premolar (pma) elongate and narrow; lower molars with prominent, horizontal external cingula. Named forms apparently conspecific with frithit are bernsteini Peters, inflatus Miller, sinicus G.M. Allen, and formosanus Horikawa. The range of the species extends from Bengal ( frithiz) to southern China (sinicus, inflatus), Formosa ( formosanus) to Java and Bali (bern- stem: Tate (1941a : 7, 11) records specimens from Bali). Osgood (1932 : 226) records inflatus from Tonkin and Annam (North V: ietnam) and there are specimens from Laos and Thailand in the British Museum (Natural History) which must be referred to C. frithii, previously recorded from Thailand by Gyldenstolpe (1916 : 15). The Malayan example thus fills a distributional hiatus but insufficient specimens are available to justify any attempt at elucidating the validity of subspecies or to allocate it at this level. Measurements of this specimen : length of forearm 43-8; of tibia 17-0; maxillary toothrow (c—m3) 5-7. The type specimen of C. vobinsoni Bonhote (1908 : 4) from the foot of Gunong Tahan, Pahang differs so markedly from C. frithii as to justify specific separation although Ellerman and Morrison-Scott (1951 : 132) considered the possibility that these might be conspecific. Coelops robinsoni may be distinguished from C. frithii by: rounded, wide outline of the lappets projecting from the supplementary leaflets; narial part of the leaf depressed and sharply demarcated from the intermediate leaf by prominent ridges extending from the central process or ‘sella’ toward the lateral margins of the leaf; posterior part of the upper surface of the rostrum nearly horizontal and only slightly inclined anteriorly ; rostral elevations large, separated by a prominent sulcus, in profile forming a distinct convexity; a small foramen but no definite pocket or recess within the maxillary root of the zygoma, the upper edge of which is sharply curved anteriorly to terminate on the rostrum above the posterior face of the second molar; upper toothrows nearly parallel; posterior lower premolar (pm) wide; lower molars with low, upwardly curving cingula. The genus Chilophylla was described by Miller (1gr0 : 395) for the species Chilo- phylla hirsuta, described concurrently from the Alag River, Mindoro, Philippines, from a skin without skull. Later, this author (Igt2 : 117) referred a specimen 30 ifs 13g SSN HCL, (United states National Museum 175000) from Port Swettenham, Malaya to jirsuta and described the cranial characters of the species from its skull. Subsequently, Miller (1928 : 85) agreed with an opinion originally advanced by Andersen (in litt.) that Chilophylla should be referred to the synonymy of Coelops, but the name has nevertheless persisted, Taylor (1934 : 247) using it in his account of the Philippine land mammals and Chasen (1940 : 48) including it in the Malaysian fauna, admittedly with the reservation that it seemed extremely close to Coelops. There can be no doubt from the meticulous descriptions of the skull of U.S.N.M. 175000 by Miller (Ig12 : 117) and Tate (1941a : 5) together with the plate in Howell (19209 : pl. 2, fig. 2e) which presumably depicts this skull (as C. robinsonz) that this specimen from Port Swettenham represents C. robinsoni. The status of C. jirsuta must remain uncertain until the features of its skull are known but its small size (length of forearm 33:8), the presence of a curved ridge extending laterally above each nostril towards the margin of the horseshoe and the statement by Miller (ibid.) that he could detect no external peculiarities that seemed of specific importance between hirsuta and the Port Swettenham specimen suggest affinity with C. robinsont. Coelops robinsoni is recorded from Teratau Island, Thailand by Chasen (1940 : 47) and from Khao Nawng, Bandon, southern Thailand by Robinson and Kloss (1915 : 116). The two specimens recorded by Robinson and Kloss are now in the British Museum (Natural History) (B.M. 68.605-606, formerly Federated Malay States Museum 531/12, 532/12): both are subadult and their skulls cannot now be found. From their external features both are referable to C. frithii, the length of the fore- arm in B.M. 68.605 being 39-4 and in B.M. 68.606 38-2 while the measurements of the skulls as quoted by Robinson and Kloss are in close agreement with those of that species. Myotis siligorensis (Horsfield, 1855) 6 B.M. 67.219 (skin and skull). Cheras cave, Panching, near Kuantan, Pahang, 3°53’ N, 103° 09’ E. 23 January 1967. This is the first record of Myotis siligorensis from the States of Malaya. Although there are few specimens in collections the species has been recorded from Kumaon, Nepal and Sikkim (M.s. siligorensis), Fukien (M.s. sowerbyi Howell), Tonkin (North Vietnam) (M.s. alticraniatus Osgood) and Thailand (M.s. thainus Shamel). From descriptions this specimen is similar to alticraniatus or thaianus but in the absence of comparative material no definitive subspecific determination can be made. The dorsal surface of the specimen is dark brown and the ventral surface of similar colour but slightly more buffy. Measurements: length of forearm (from collector) 29°6; greatest length of skull 12:2; condylobasal length 11-1; condylocanine length 10-4; zygomatic width 7-0; least interorbital width 3-0; width of braincase 6-0; depth of braincase 4:5; mastoid width 6-3; m®-m% 4-6; length of entire upper toothrow 5:1; c-m3 4-4. Bats of this species may be recognized by their small MALAYAN BATS AND A NEW PIPISTRELLUS 31 size, high, domed braincase and much reduced canines, the lower tooth about equal in height to the posterior lower premolar (pma), with the second lower premolar (pms) in the axis of the toothrow. Myotis horsfieldii horsfieldii (Temminck, 1840) 63 B.M. 65.320 (skin and skull), B.M. 65.321 (in alcohol, skull extracted). Ampang Reservoir, Kuala Lumpur, Selangor 3° 11’ N, ro1° 49’E. 15 October 1964. Vespertilio horsfieldii Temminck was considered a synonym of Myotis adversus (Horsfield, 1824) by Dobson (1878 : 291 et seq) and by Tate (1941b : 551, 558). However, Thomas (1910 : 385) identified a series of specimens from Java (which he (with Wroughton) had formerly (Igoga : 381) determnied as M. hasseltit) as M.horsfieldit, pointing out that this species could bedistinguished from M. adversus by the attachment of the wing membrane to the metatarsus instead of to the end of the tibia asin adversus. It is clear from a drawing now preserved in the British Museum (Natural History) copy of Monographies de Mammalogie at Temminck’s description of horsfieldii that Thomas had submitted illustrations of this feature in his “‘adversus’’ from Java and in the specimens of “‘hasseltit”’ which he had from the same collection to F. A. Jentink at the Rijksmuseum van Natuurlijke Historie, Leiden, wherein are Temminck’s type specimens. Jentink indicates on this drawing that the insertion of the wing membrane in one of the “cotypes” of horsfieldii (‘c’ of his catalogues (1887 : 280, 1888 : 186)) is identical with the specimens which Thomas considered representative of hasseltii and which he subsequently recognised as horsfieldiv. Myotis horsfieldit is a relatively small species, the dorsal surface being blackish and the ventral surface blackish grey, with narrow, uninflated braincase, narrow inter- orbital region, interdental palate terminating anteriorly to a line joining the posterior faces of m®-3, long post-palatal extension, the median post-palatal spine supported by bony laminae. The second upper premolar (pm®) is large, one quarter to one third the crown area of pm2, usually but not always completely intruded from the toothrow so that pm? and pm‘ are almost in contact, with pm? not compressed in the row; second lower premolar (pms) one third or a little more the crown area of pms, separating pmg and pmy but usually slightly intruded from the toothrow. The species may be distinguished from M. adversus and M. hasseltii by its small size, blackish coloration and narrow braincase. These specimens (and one other, 9 B.M. 62.2133, also from Ampang Reservoir, obtained originally by the Institute for Medi- cal Research, Kuala Lumpur) agree closely with M.h. horsfieldii and are rather larger than M. deignani Shamel, 1942, from Chiengmai, Thailand, which seems likely to be a subspecies of M. horsfieldit. MEASUREMENTS (B.M. 65.320-321, 62.2133 (forearm only),in that order) : length of forearm 38-7, 37-8, 36-4, 36-9; greatest length of skull 15-7,15-9, 15-2; condylobasal length 14-3, 14-2, 13-7; condylocanine length 13-6, 13-6, 13:1; zygomatic width —, 32 Yo 1. NIE. 9-3, —; least interorbital width 3-6, 3:5, 3:4; width of braincase 7-3, 7-3, 6:9; mastoid width 7-9, 7-9, 7-5; cl-c! 4-3, 4:2, —; m3-m 6-1, 6-1, 5-7; c-m3 5-7, 5-9, 5:6. Myotis hasseltii (Temminck, 1840) @ B.M. 65.322 (skin and skull). Kuala Gula, Perak, 4° 52’ N, 100° 32’ E. 8 November 1964. 3 B.M. 68.842, 2 B.M. 68.843 (both in alcohol, skulls extracted). Kuah, Pulau Langkawi, Kedah 6° 16’ N, 99°50’ E. 1 January 1968. Q subadult B.M. 68.844 (in alcohol). Kangar, Perlis, 6°27’N, 100° 12’ E. I January 1968. These specimens agree closely with those from Lekop, Karimon Island, Riau Archipelago, Indonesia recorded by Thomas and Wroughton (1909b : 110) as ‘Myotis adversus (Horsf.) (?)’ and subsequently identified by Thomas (1916 : 4) as Leuconoe hasseltii, and also with two specimens from Java labelled as originating from the Rijksmuseum van Natuurlijke Historie, Leiden prior to 1844 and thereby possibly examined by Temminck. They may be distinguished from specimens attributed to the otherwise similar species adversus by their short and not woolly pelage, short post-palatal extension lacking thin bony laminae supporting the post-palatal spine and in having a minute second upper premolar (pm3) with the second lower premolar (pms) very small, usually intruded from the toothrow some- times to the extent that pmg and pmy are in contact. There exist apparently two colour variants: two specimens from Java (both old) are brownish above and below as is one from Karimon Island but two others from that island and four from Malaya are greyish brown dorsally and paler greyish white or greyish buff ventrally. MEASUREMENTS (B.M. 65.322, B.M. 68.842-843); length of forearm 40-2, 40-7, 41-2; length of foot 8-5, 10.0, 9-5; greatest length of skull —, 16-3, 16-1; condylo- basal length 14-2, 15:0, 14-8; condylocanine length 13-5, 14:3, 14:0; zygomatic width —, 10-3, —, least interorbital width 3-8, 4:2, 3-9; width of braincase 7-6, 8-0, 8.0; mastoid width 8-0, 8-7, 8:6; cl-cl 4:1, 4:6, 4:5; m3m3 6-1, 6-6, 6-6; c-m 5-7, 6-1, 6:0. The named forms of large-footed Myotis (sometimes referred to a subgenus, Leuconoe) from southeastern Asia need detailed revision but unfortunately the majority seem represented by no more than a few isolated specimens or by a series from no more than one or a few localities. There seems little doubt that horsfieldii is a distinct species (Medway, 1965 : 60) despite the views of Dobson (1876 : 127, 1878 : 292) and Tate (19g4Ib : 551, 598) that it is a synonym of adversus. The relationship between hasseltii and adversus as here understood is obscure and it may be that the advent of further Javan specimens and an examination of the type material in Leiden might well alter their taxonomy. Meanwhile, hasseltii is used for those bats in which pmg is very small and usually intruded or fully intruded from the toothrow, adversus for those in which this tooth is larger and stands within the MALAYAN BATS AND A NEW PIPISTRELLUS 33 toothrow or is only slightly intruded, and which have rather long, woolly pelage and a long post-palatal extension, its median spine supported by thin bony laminae. Pipistrellus javanicus javanicus (Gray, 1838) 6 (skin and skull) B.M. 71.813). Camp 2, Gunong Benom, Pahang 3° 51’ N, too° 11’ E, 700 ft. 31 January 1968. The small anterior upper premolar is sometimes lacking from the toothrows of Pipistrellus (Kuzyakin, 1944 : I0I, 1950 : 338 et seq., 1965 : 103); on the other hand, it may sometimes be present in those of Eptesicus (Hayman, 1954 : 289, 290; Hill, 1966 : 303). However, this specimen is noteworthy for the presence of a small supernumerary premolar in the right toothrow, situated internally in a shallow angle formed by the postero-internal face of the canine and the antero-internal face of the anterior upper premolar normal to this species, which is situated more nearly in the line of the toothrow than the corresponding tooth in the other row. The supernumerary tooth is small, with a narrow cingulum and a slender conical cusp slightly exceeding the normal anterior premolar in height and just visible externally. It is tightly in contact with the canine and the normal anterior pre- molar, with a basal area of one half that of this tooth. There are no pecularities in the mandibular dentition. Pipistrellus ridleyi Thomas, 18981 6 B.M. 67.1604 (skin and skull). Base Camp, Gunong Benom, Pahang, 3° 51’ N, 102° 11’ E, 700 ft. 16 February 1967. This species has been represented hitherto in the collections of the British Museum (Natural History) only by the type and paratype specimens, B.M. 98.3.15.5—6, from Selangor. The teeth of the example from Gunong Benom are less worn than are those of the type specimen and agree closely with those of the paratype, the skull of which has now been prepared from the specimen in alcohol. The description of the incisors by Thomas is based solely on the type specimen and an examination of specimens in which wear is less evident enables some additional features to be noted. The inner upper incisor (i?) is low, short and wide, narrow anteriorly but wider posteriorly, its length at the cingulum only slightly exceeding its greatest width. The principal cusp is projected forward beyond the premaxilla and is directed slightly inwardly rather than anteriorly: in profile its anterior surface is strongly convex and its posterior face slightly concave so that it is hooked posteriorly. There are low, paired posterior accessory cusps; the labial cusp is smaller than the lingual cusp and in the type specimen is almost worn away so that the inner cusp becomes readily visible from the side. The outer upper incisor (13) is short and wide, its width exceeding its length, and has a principal cusp hooked posteriorly so that the tooth has a caniniform appearance. The principal cusp is equal in height to the principal cusp of i? and is flanked by two lateral accessory cingulum cusps: 1 Further study of this species indicates that it should be transferred to Myotis (Hill & Topal, in press). 34 Vo Ide isnonie the lingual cusp is well-developed and prominent but the labial cusp is smaller, less obvious and in the type specimen has been worn away. The anterior upper premolar (pm2) is large, in the toothrow and separated from pm* by a narrow diastema. The lower incisors are tricuspid and stand in the line of the toothrow: as remarked by Thomas, the third is considerably more massive than the other two. The anterior lower premolar (pmg) is not greatly reduced and is not extruded from the toothrow. Measurements of B.M. 67.1604, followed by those of the type and paratype specimens: length of forearm 28-8, 28-5, 29-1; greatest length of skull 12-3, 12-7 12:3; condylobasal length 11-6, 11-8, 11-5; condylocanine length 10-9, 11-2, III; width across anteorbital foramina 3-6, 3:5, 3:7; zygomatic width —, 8-0, —; least interorbital width 2-9, 3-0, 3:3; width of braincase 6-2, 6-3, 6-2; mastoid width 6-7, 6-7, 6-8; cl-cl 3-4, 3-3, 3-4; m3-m3 5-4; 5:3, 5-3; c-m8 4-4, 4°5, 4-4; length of mandible —, 8-7, 8-7; c—mg 4-7, 4'8, 4-7. Pipistrellus societatis sp. nov. TYPE AND ONLY SPECIMEN: ¢ B.M. 67.1605 (in alcohol, skull extracted). Base Camp, Gunong Benom, Pahang, Malaya, 3°51’ N, 102° 11’ E, 800 ft. Collected 15 March 1967 by Lim Boo Liat and Yong Hoi Sen for the Gunong Benom Expedi- tion, 1967. Original number BM 225. Diacnosis: Allied to and externally closely resembling Pipistrellus circumdatus (Temminck), but differing in smaller size; in greater inflation of the anterior part of the braincase; in its shorter, narrower rostrum which is more inflated laterally; in the absence of well-defined supraorbital ridges; in its more abruptly curved supraorbital margin; in the absence of a deeply excavated frontal depression; narrower narial emargination which is not constricted posteriorly; in having a shorter post-palatal extension with the pterygoid region widened posteriorly; in shallow basial pits which are not sharply excised into the basisphenoid; in shorter toothrows, less massive dentition and much reduced third upper molar, in which the metacone and third commissure have been all but lost. DescriPTION: Externally almost exactly like P. circumdatus but with slightly shorter forearm. Dorsal pelage predominantly blackish brown, heavily overlaid by orange or bronze tipping, the individual hairs basally brown, blackish brown for most of their length and profusely but irregularly tipped with orange or bronze. Crown pelage orange tipped but with a small area of straw-coloured, dark brown tipped hairs just anterior to the junction of the aural anterior margin with the head, a feature not so far noted in civcwmdatus. Ventral pelage dark brown, the hairs tipped with greyish white: there is a small wart bearing a few longer, darker hairs medianly situated on the throat just anterior to a line joining the angles of the mouth. Fur extending a little on to the uropatagium but tibia not especially hairy, the digits with a sparse cover of moderate hairs. Ears large and bluntly rounded, their anterior margin strongly convex in the proximal half, with prominent, posteriorly directed basal lobule. Distal half of MALAYAN BATS AND A NEW PIPISTRELLUS 35 anterior margin almost straight, upper part of posterior margin similarly nearly straight but becoming convex with an emargination opposite the base of the tragus and terminating in a wide, quadrangular antitragal lobe, its anterior edge standing squarely to the margin of the ear and to the cheek, inserted a little behind and below the angle of the mouth, forming a small downwardly directed pendent lobule similar to that of Chalinolobus gouldi. Tragus large, its sharply concave anterior margin terminating in a rounded, anteriorly directed point: posterior margin strongly convex, its upper part almost horizontal, terminating basally in a large, rounded lobule. Ear margins, especially anteriorly, yellowish white, a feature remarked by Temminck in the original account of P. civcumdatus and evident to some extent in dried specimens. Skull short, wide, with rounded, rather globular braincase, inflated and elevated in the frontal region, the lambdoid crests not especially prominent and the sagittal crest lacking. Interorbital region wide, the supraorbital crests only faintly defined, terminating in small tubercles. Frontal depression shallow, with weakly defined boundaries extending anteriorly as a very shallow median sulcus, bordered by wide, low lateral swellings which contrast with the higher, more ridge-like lateral swellings of circumdatus. Rostrum short and wide, inflated laterally above anteorbital foramen, the rostrum in frontal aspect having a rounded rather than angular profile as in that species. Supraorbital ridges curving abruptly from the interorbital region to join the anterior margin of the orbit which forms a flange at the side of the rostrum. Narial emargination parallel-sided, almost square posteriorly. Palate short with a narrow anterior emargination a little wider than the distance between the inner faces of i2-2, extending posteriorly to a line joining the back of the canines. Post-palatal extension short, with apparently narrow post-palatal spine, the pterygoid wings divergent posteriorly. Basial pits shallow, not sharply excised into the basisphenoid. Teeth of type specimen heavily worn: inner upper incisor (i2) massive, faintly bifid at tip, with small posterior cingulum cusp, outer upper incisor (i) approxi- mately one third basal area of i? with large central cusp flanked by smaller lateral cusps, its tip reaching slightly above cingulum of i?, separated from canine by a narrow diastema and pushed forward so that the upper incisors lie almost in a straight line. Anterior upper premolar (pm2) very small, completely intruded from toothrow: in the type specimen pm? is lacking from the right toothrow but a small alveolus can be discerned. Third upper molar (m*) reduced, narrow, platelet-like: in the type specimen the complete loss of the third commissure is clearly due to wear but it is obvious that this and the metacone are much reduced. Inner (i;) and second (ig) lower incisors incipiently four-cusped; ig shorter and wider than these, tricuspid. Anterior lower premolar (pmg) one fourth crown area of pm4: hypoconid and entoconid of mg slightly reduced. Measurements of P. societatis and P. circum- datus appear in Table 3. Remarks: This new species is clearly very closely related to P. circumdatus, a poorly known species apparently so far represented only by the female type specimen from Tapos, Java in the Rijksmuseum van Natuurlijke Historie, Leiden (Jentink, 36 J. E. HILL 1887 : 277, 1888 : 178); a specimen (B.M. 7.1.1.401) labelled “Java” from the Tomes Collection, now in the British Museum (Natural History) and possibly one of the original specimens; an example (B.M. 61.12.10.1) without skull labelled “Tndia’’ and also in the British Museum (Natural History); and from a specimen from Pyepat, Upper Burma, recorded by Anthony (1941 : 81) and Tate (1942 : 250) and now part of the collection of the American Museum of Natural History.1 Through the courtesy of the authorities of the Rijksmuseum van Natuurlijke Historie I have been able to examine the type specimen and I am indebted to Dr. K. F. Koopman who has arranged the loan of the Burmese specimen from the American Museum of Natural History. Both P. circumdatus and P. societatis may be readily recognized by their striking coloration, the blackish dorsal pelage having characteristic orange or bronze tips. The skull of c’vcumdatus, however, does not seem to have been described except in so far as Tate (1942 : 250) based his group definition on the skull of A.M.N.H. 114850 from Burma. Cranially, civcwmdatus is more massive than societatis, its braincase less inflated frontally and with a low sagittal crest. The supraorbital ridges are prominent, terminating in small tubercles, enclosing a deeply excavated frontal depression which extends anteriorly as a shallow rostral sulcus, bound laterally by low longitudinal inflations which converge to merge anteriorly. Rostrum longer and broader than in societatis, the area above the anteorbital foramen uninflated but instead sloping more abruptly from the inflations bordering the rostral sulcus to give therostrumin frontalaspect anangularratherthanroundedoutline. Supraorbital ridges curving less abruptly from the interorbital region than in societatis, the anterior margin of the orbit forming a flange as in that species. Narial emargination nar- rowed posteriorly with a rounded apex, the anterior palatal emargination equal in width to the distance between the inner faces of i2-2 or slightly exceeding it and extending posteriorly almost to a line joining the posterior faces of the canines. Post-palatal extension slightly longer than in societatis, with narrow post-palatal spine, the pterygoid wings parallel: basial pits prominent, deep, sharply excised into the basisphenoid. Dentition as described for P. societatis but teeth larger and more massive, m? not reduced but with well-developed metacone and clearly evident third commissure, while m’ has the hyopconid and entoconid unreduced. The skull of the Burmese example differs from the skulls of the type and second Javan specimens in slightly smaller size, less prominent cranial crests, its supraorbital ridges terminating in slight tubercles which are barely evident in the Javan examples, in a shallower frontal depression, shorter post-palatal extension and narrower post- palatal spine, and slightly less massive dentition. It is possible therefore that adequate series of specimens may show the mainland and Javan populations to be subspecifically distinct.1 The name of the new species is selected in allusion to the combined effort of the several members of differing nationalities and institutions who constituted the Gunong Benom Expedition, 1967. 1 Lord Medway has now obtained (1971) a specimen of P. circumdatus from Fraser’s Hill, Pahang. It agrees closely with the Burmese example. MALAYAN BATS AND A NEW PIPISTRELLUS 37 TABLE 3 MEASUREMENTS OF Pipistrellus societatis AND P. civcumdatus g 2 2 an) S S 8 Ue ewe 3 Sam ete S of 3 8 86 Ss; 28 ge aes ae S oe R ¢ iS = 3 eS 3 qe ester Oy Bag eee Sam S35 SAR Be G RAs YH yYAare Aa Length of forearm 37:6 42* 43°6 41°6 Greatest length of skull 15°3 — — 15°5 Condylobasal length 14°4 a 15°7 14°9 Condylocanine length 14°! -= 15°4 14°7 Palatal length (excluding post-palatal spine) 6:6 — 8-4 7°6 Length palatal bridge (excluding post-palatal spine) 5:0 — 6-6 6-1 Length orbit — gnathion 3°6 4°2 4°2 4°3 Width across anteorbital foramina EjOn 5°5 5°6 5°3 Lachrimal width 6°7 7/373 TA GpO8} Width across supraorbital tubercles 6-0 6:7 6-6 6°7 Zygomatic width 10-4 — — 10°8 Least interorbital width 4°3 4:12 4°4 4°4 Width of braincase 7°9 _— — 7°9 Mastoid width 8-5 — — 8-5 cl-cl 4°6 5:2 5°2 4-9** m3—m3 6°7 a 7°5 7-1 c-m3 5°2 6-2 6°4 6-0 Length of mandible 10-4 -- 12-1 — c—m3 5:6 6°5 6-7 6-4 * From Tate (1949 : 292 ** Right upper canine damaged Hesperoptenus blanfordi (Dobson, 1877) 2 B.M. 65.345 (in alcohol, skull extracted). Jenka, Temerloh, Pahang, 3° 37’ N, 102° 30’ E. Collected and presented by the Earl of Cranbrook. Described originally from Tenasserim, this species is relatively rare in collections but is recorded by Anderson (1881 : 133) from Johore and by Robinson and Kloss (1915 : 116) from Khao Nawng, Bandon, Lower Thailand, while Thomas (1916 : 2) lists specimens from the Semangkko Pass, below Fraser’s Hill, on the Selangor- Pahang boundary; Gunong Tampin, Negri Sembilan; Telok Bahang (= Rahang), Penang, and from Kuala Lumpur, Selangor. The specimen listed from Khao Nawng by this author is doubtless that previously recorded by Robinson and Kloss. The species is easily recognized by its small size (length of forearm 26-28), the 38 Vo 25 1UnKE presence of a broad, well-developed pad at the base of the thumb extending to the base of the second metacarpal, and by the small size and extreme displacement of the outer upper incisor (i8) which is intruded from the toothrow to the extent that it lies directly behind the large inner upper incisor (i?), in contact with the posterior face of that tooth and with the lingual face of the canine. Davis (1962 : 42) recorded a specimen from the Sapagaya Forest Reserve, Sanda- kan, Sabah (5° 37’ N, 118° 04’ E) as Hesperoptenus doriae. Through the kindness of Dr. J. C. Moore of the Field Museum of Natural History, Chicago, I have been able to borrow and examine this specimen, F.M. 77025. As Davis points out, it is a juvenile male. However, it is much larger than the adult type specimen of Hesper- optenus doriae and agrees very closely with the type specimen of H. tomesi, described by Thomas (1905 : 575) from Malacca, to which it must be referred. Hesperoptenus tomesi has been known hitherto only from the type specimen and the example from Sabah is therefore the first record for Borneo. It may be recognized by its large size (length of forearm c. 50) which is similar to that of the Indian H. tickelli, by its rich, dark brown colour, by its displaced outer upper incisor and by its strongly imbricated lower incisors. Hesperoptenus doviae is smaller (length of forearm 38-0) and is (from specimen in spirit) pale brown, with the outer upper incisor not displaced from the toothrow and the lower incisors not much imbricated. Murina huttoni (Peters, 1872) ¢ B.M. 67.1606 (in alcohol, skull extracted). Near Camp 4, Gunong Benom, Pahang, 3°51’ N, 102° 11’ E, 4800 ft. 17 March 1967. This species establishes the presence of M. huttoni in Malaya, whence M. suilla, M. cyclotis and the striking M. aenea have already been recorded (Hill, 1964). Although very similar to M. cyclotis peninsularis in external appearance, this species may be readily distinguished by the features of its molar teeth, the mesostyles of ml-2 being more developed than in M.c. peninsularis with the paracones and proto- cones less reduced, and especially by the unreduced hypoconids and entoconids of m,_2, which are separated from the protoconids and metaconids by a wide trough. These features indubitably refer the specimen to huttoni: in cyclotis the mesostyles, paracones and protocones of m1~2 are reduced and in m_2 the hypoconids and ento- conids are much reduced, sometimes barely evident, and are only narrowly separated from the protoconids and metaconids. The degree of reduction is less apparent in mg. Subspecific determination of the specimen is less certain: in dorsal coloration it resembles M.h. rubella from Fukien, the hairs with bright golden brown sub- terminal annulations as in that subspecies. MEASUREMENTS: length of forearm 34:1; greatest length of skull 17-5; condylo- basal length 15-6; palatal length 8-5; length orbit-gnathion 4:5; rostral width 5-5; zygomatic width 9-4; least interorbital width 4-3; width of braincase 7-5; height of braincase 6-1; mastoid width 8-0; cl-cl 4-3; m3—m3 5-5, c-m3 5-8. MALAYAN BATS AND A NEW PIPISTRELLUS 39 Murina cyclotis peninsularis Hill, 1964 2 B.M. 67.1607 (skin and skull). Base Camp, Gunong Benom, Pahang, 3° 51’ N, to2° 11’ E, 700 ft. 20 February 1967. 3 B.M. 68.845 (in alcohol, skull extracted). Waterworks Road, Batu Pahat, Kangar, Perlis, c. 6° 27’N, 100° 12'E. 4 January 1068. These specimens are the first examples of this subspecies to be recorded since its description (Hill, 1964). Although in some dimensions the specimen from Perlis is similar to M.c. cyclotis, it agrees with M.c. peninsularis in having the rostrum wide anteriorly, with toothrows which are not especially convergent, and in massive canines, premolars and molars. Measurements (B.M. 67.1607, B.M. 68.845): length of forearm 38-8, 34-7; greatest length of skull 18-6, 17-4; condylobasal length 16-7, 15-4; length orbit-gnathion 4-5, 4-3; palatal length 9-2, 8-2; rostral width 6-0, 5-8; zygomatic width 10-9, 9-9; least interorbital width 4-5, 4-3; width of brain- case 8-1; 8-0; height of braincase 6-7, 6-7; mastoid width 9-1, 8-5; cl—cl 5-1, 4°5 m3—m3 6-0, 5-6; c-m3 6:2, 5-5. Shamel (1942 : 327) recorded two specimens from northern Thailand as M .toxoper, described by Thomas (1923 : 254) from the island of Buru in the Moluccas. From the description and measurements given by Shamel it seems more probable that these specimens represent M. cyclotis. Miniopterus schreibersii blepotis (Temminck, 1840) 33 B.M. 67.216-217 (skins and skulls). Pine Tree Hill, Fraser’s Hill, Pahang, 3° 43'N, to1° 42’ E, 4250 ft. 12 October 1966. 3 B.M. 67.218 (in alcohol). Fraser's Hill, 4750 ft. 22 October 1966. Although known to occur from mainland localities as far south as Koh Lak in southern Thailand (specimen in British Museum (Natural History)) and from Sumatra, Java and Borneo, there seems to be no published record of M. schretbersii from Malaya, whence these specimens now establish it. Measurements of B.M. 67.216-217: length of forearm 44-8, 50-8; condylobasal length 15-0, 16-3; condylo- canine length 14-1, 15:5; zygomatic width 9-0, 9-4; least interorbital width 4-0, 4-2; width of braincase 8-1, 8-5; m3-m3 6-6, 7-4; c-m3 6:0, 6:5. The length of forearm in B.M. 67.218 is 49-3. The large example (B.M. 67.217) agrees almost exactly with a similarly large specimen from Kok Lak, further north on the east coast of Lower Thailand. Small specimens of M. schreibersii are difficult to distinguish from large examples of the sympatric species M. medius. However, apart from its usually longer fore- arm, M. schreibersii differs from medius in longer skull and wider palate: specimens from Java, Malaya and Indochina have a length of forearm of 43-6-50-8, condylo- basal length 14-7-16-3, condylocanine length 13-8-15-5 and m3-m3 6-5-7°4 while specimens of medius from Java and Pulau Kaban (off the east coast of Johore, States of Malaya) have a length of forearm of 40-8-43-1, condylobasal length 13-8- 14-3, condylocanine length 13-1-13-5 and m3-m3 5-8-6-3. 40 J. E. HILL Miniopterus medius Thomas and Wroughton, 1909 © B.M. 67.1610 (skin and skull). Bukit Cheras, Panching, near Kuantan, Pahang, 3° 53’ N, 103° 09’ E. 17 February 1967. This is a further Malayan record for the species, hitherto known on the mainland only from Gunong Pondok, Perak (Chasen, 1926 : 156) but also recorded from Pulau Kaban, off the east coast of Johore, and from Pulau Terutau, southern Thailand by Thomas (1916: 4). Measurements: length of forearm 42-2; greatest length of skull 14-4; condylobasal length 13-8; condylocanine length 13-0; zygo- matic width —; least interorbital width 3-7; width of braincase 7-7; mastoid width 8-1; cl-cl 4-0; mm 5-9; c-m3 5-4. Measurements of other specimens from eastern Pahang (supplied by Lord Medway) ($¢g2 Medway 66.46—48) length of fore- arm (Medway 66.47-48 only) 43; condylobasal length 13-8-14-0; condylocanine length 12-9-13-2; m3—m3 5-8-5-9. SUMMARY Taxonomic notes on the more unusual species of bats obtained by the 1967 Expedition to Gunong Benom are presented in this paper, with similar accounts of other specimens of taxonomic interest obtained in recent years in Malaya by Lord Medway and his associates. The genus Coelops is reviewed so far as available material permits, with the recognition of two valid species, C. frithii and C. robinsont. A new species, Pipistrellus societatis, is described from Gunong Benom: Rhinolophus malayanus, R. robinsoni (2?) klossi, R. macrotis dohrni, Coelops frithit, Myotis sili- gorensis and Murina huttoni are added to the bat fauna of the States of Malaya while additional specimens of Hipposideros lylet, Myotis horsfieldi1 horsfieldit, Myotis hasseltii, Pipistrellus ridleyi, Hesperoptenus blanfordi, Murina cyclotis peninsularis, Miniopterus schreibersu blepotis and M. medius are recorded. Hesper- optenus tomesi is recorded for the first time from Borneo from a specimen hitherto thought to be H. doriae and an example of anomalous dentition in Pipistrellus javanicus is described. ACKNOWLEDGEMENTS I am indebted to the authorities of the Rijksmuseum van Natuurlijke Historie, Leiden, the American Museum of Natural History, New York and the Field Museum of Natural History, Chicago, for the loan of specimens essential to this study. REFERENCES ANDERSEN, K. 1905a. On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty six new forms. Pvoc. zool. Soc. Lond. 2 : 75-145, 22 figs, 2 pls. 1905b. A list of the subspecies of the genus Rhinolophus, with some notes on their geo- graphical distribution. dnn. Mag. nat. Hist. (7), 16 : 648-662. MALAYAN BATS AND A NEW PIPISTRELLUS 41 ANDERSEN, K.1918. Diagnoses of new bats of the families Rhinolophidae and Megadermatidae. Ann. Mag. nat. Hist. (9), 2 : 374-384. ANDERSON, J. 1881. Catalogue of the Mammalia in the Indian Museum, Calcutta. I. Pri- mates, Prosimiae, Chiroptera, and Insectivorva. Calcutta. AntHony, H. E. 1941. Mammals collected by the Vernay —Cutting Burma Expedition. Publs Field Mus. nat. Hist. (Zool. Ser.) 27 : 37-123, 4 pls., map. Bonuote, J. L. 1908. Report on the Gunong Tahan Expedition, May—September, 1905. I. Reportonthemammals. J. fed. Malay States Mus.3:1-11,1 pl. (Field notes by H.C. Robinson). Cuasen, F. N. 1926. Miniopterus medius in the Malay Peninsula. J. Malay. Brch R. Asiat. Soc. 4 : 156-157., 2 tabs. 1940. A Handlist of Malaysianmammals. Bull. Raffles Mus. No. 15 : i-xx, 1-209, map. Davis, D. D. 1962. Mammals of the lowland rain-forest of North Borneo. Bull. natn. Mus., Singapore No. 31 : 1-129, 20 figs., 23 pls. Dogson, G. E. 1876. Monograph of the Asiatic Chiroptera and catalogue of the species of bats im the collection of the Indian Museum, Calcutta. London. 1878. Catalogue of the Chiroptera in the collection of the British Museum. London. ELLERMAN, J. R. & Morrison-Scott, T. C. 5S. 1951. Checklist of Palaearctic and Indian mammals, 1758-1946. 1st ed., London. GYLDENSTOLPE, N. 1916. Zoological Results of the Swedish Zoological Expeditions to Siam IQlI-1912 & 1914-1915. V. Mammals. II. K. svenska Vetensk-Akad. Handl. 57, 2 :1-59, 6 pls. Hayman, R. W. 1954. Notes on African bats, mainly from the Belgian Congo. Revue Zool. Bot. afr. 50 : 277-295. Hi1, J. E. 1964. Notes on some tube-nosed bats, genus Murina, from southeastern Asia, with descriptions of a new species and a new subspecies. Fedn. Mus. J. (N.S.) (1963), 8 : 48-59, 4 pls. 1966. The status of Pipistrellus vegulus Thomas (Chiroptera, Vespertilionidae). Mam- malia, 30 : 302-307. — &Topat,G. The affinities of Pipistvellus ridleyi Thomas, 1898 and Glischropus rosseti Oey, 1951 (Chiroptera: Vespertilionidae). (In press.) Howe tt, A. B. 1929. Mammals from China in the collections of the United States National Museum. Proc. U.S. natn. Mus. 75 : 1-82, to pls. JEnTINK, F. A. 1887. Catalogue Ostéologique des mammiféres. TomeIX. Leiden. 1888. Catalogue Systématique des mammiféres (Rongeurs, Insectivores, Cheivoptéeres, Edentés et Marsupiaux). Tome XII. Leiden. Kuzyakin, A. P. 1944. In Bobrinskii, N. A., Kuznetzov, B. A. & Kuzyakin, A. P. Mam- mals of the U.S.S.R. Moscow. i —— 1950. Letucie Myshi. (Bats. (Systematics, life history and utility for agriculture and forestry)]. Moscow. 1965. In Bobrinskii, N. A., Kuznetzov, B. A. & Kuzyakin, A. P. Mammals of the U.S.S.R. (New ed.). Moscow. Mepway, Lorp. 1965. Mammals of Borneo. Field Keys and an annotated Checklist. J. Malay. Brch R. Asiat. Soc. (1963), 36, 3 (No. 203): i-xiv, 1-193, 9 figs, 34 pls, 5 tabs, map. Mitter, G. S. 1910. Descriptions of two new genera and sixteen new species from the Philippine Islands. Pyvoc. U.S. natn. Mus. 38 : 391—404, 3 pls. 1912. The cranial and dental characters of Chilophylla. Proc. biol. Soc. Wash. 25 : 117. 1928. A new bat of the genus Coelops. Proc. biol. Soc. Wash. 41 : 85-86. Oscoop, W. 1932. Mammals of the Kelley — Roosevelts and Delacour Asiatic Expeditions. Publs Field Mus. nat. Hist. (Zool. ser.) 18 : 193-339, 30 figs, 2 pls. Puituirs, C. J. 1967. A collection of bats from Laos. J. Mammal. 48 : 633-636, 1 tab. Rosginson, H. C. & Ktoss, C. B. 1915. Ona collection of mammals from the Siamese Prov- ince of Bandon, N.E. Malay Peninsula. J. fed. Malay St. Mus. 5 : 111-127. 42 J. E. HILL SANBORN, C. C. 1939. Eight new bats of the genus Rhinolophus. Publs Field Mus. nat. Hist. (Zool. Ser.) 24 : 37-43. SHaMEL, H.H. 1942. A collection of bats from Thailand (Siam). J. Mammal. 23 : 317-328. Tate, G. H. H. 1941a. Results of the Archbold Expeditions. No. 36. Remarks on some Old World leaf-nosed bats. Am. Mus. Novit. 1140 : I-11. 1941b. Results of the Archbold Expeditions. No. 39. Review of Myotis of Eurasia. Bull. Am. Mus. nat. Hist. 78 : 537-565, 2 figs. 1942. Results of the Archbold Expeditions. No. 47. Review of the Vespertilionine bats, with special attention to genera and species of the Archbold Collections. Bull. Am. Mus. nat. Hist. 80 : 221-297, 3 figs. 1943. Results of the Archbold Expeditions. No. 49. Further notes on the Rhinolophus philippinensis group (Chiroptera). Am. Mus. Novit. 1219 : 1-7, 2 tabs. —— & ARCHBOLD, R. 1939. Results of the Archbold Expeditions. No.24. Oriental Rhino- lophus, with special reference to material from the Archbold Collections. Am. Mus. Novit. 1036 : I-12, 1 tab. Taytor, E. H. 1934. Philippine land mammals. Monogr. Philipp. Bureau of Science Manila, 30 : 1-548, 17 figs, 24 pls. Tuomas, O. 1905. A new genus and two new species of bats. Ann. Mag. nat. Hist. (7), 16 : 572. 1910. New mammals from the E. Indian Archipelago. Ann. Mag. nat. Hist. (8), 5 : 383- 387. 1916. List of Microchiroptera other than leaf-nose bats in the collections of the Federated Malay States Museums. J]. fed. Malay States Mus. 7 : 1-6. 1923. On some small mammals, chiefly bats, from the East Indian Archipelago. Ann. Mag. nat. Hist. (9), 11 : 250-255. —— & Wroucuton, R.C. 1909a. Ona collection of mammals from Western Java presented to the National Museum by Mr. W. E. Balston. Pyoc. zool. Soc. Lond. 371-392. tgogb. On mammals from the Rhio Archipelago and Malay Peninsula collected by Messrs. H. C. Robinson, C. Boden Kloss and E. Seimund and presented to the National Museum by the Government of the Federated Malay States. J. fed. Malay St. Mus. 4: 99-129. J. E. Hitt Department of Zoology British Museum (Naturat History) CROMWELL Roap Lonpon. S.W.7 2 i ee lj A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF tea THE BRITISH MUSEUM (NATURAL HISTORY) BS : Kay, E. Attson. Marine Molluscs in ihe Cuming Collection British Museum _ (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. a WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier ana it Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. “7 Taytor, J. D., Kennepy, W. J. & Hart, A. The Shell Structure and Mineralogy _ of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, — 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In press.) ul Sut Printed in England by Staples Printers Limited at their Kettering, Northants, establishment A. G, C. GRANDISON BULLETIN OF MUSEUM (NATURAL HISTORY) BBP Vol. 23 No.4 _ - LONDON: 1972 | THE GUNONG BENOM EXPEDITION 1967 5. REPTILES AND AMPHIBIANS OF GUNONG BENOM WITH A DESCRIPTION OF A NEW SPECIES OF MACROCALAMUS BY ALICE G. C. GRANDISON - 8 NOV al Va PRESERTED ES . Loreal region vertical, lips not flaring. to. Iris vermilion in life. RANGE: Peninsular Thailand and Malaya. Megophrys monticola nasuta (Schlegel) Ceratophryne nasuta Schlegel 1858 : 57. MATERIAL: BM 1967.2318-24 (1 9, 5 g, I juvenile). Hasirat: Most of the examples were found at night on the forest floor among dead leaves in areas close to streams, between 600 and 1650 ft. One adult male was found at night sitting upright in the stream by fallen leaves, between boulders, just below the small waterfall at 1650 ft in the valley below camp 3. COLOUR IN LIFE: Above tan or chocolate brown, the middle of the trunk from behind the orbits to the vent a darker shade, this area being bordered anteriorly by a wide angled ‘V’. One or two dark brown transverse stripes across the top of the head. A dark brown, white bordered loreal patch and an oblique dark chocolate brown, sometimes white-edged stripe from below the posterior half of the eye to the upper lip were constantly present. A few, small, round, conical, 64 A. G. C. GRANDISON almost black tubercles irregularly scattered over the back. Across the vent a dark brown area with median constriction. Throat and chest usually chocolate brown or grey-brown mottled with darker brown and with a dark, median band; belly marbled brown and cream. A dark brown spot under forearm and dark patches under wrist and on knee; under surfaces of tarsus and metatarsus dark chocolate brown. Iris medium brown. REMARKS: Inger (1954) has given an excellent and detailed description of monticola and an account on geographic variation and the characteristics of monticola nasuta. See also remarks under M. aceras. RANGE: Malay Peninsula, Sumatra, Natuna and Borneo (Inger 1954). Family RANIDAE Amolops larutensis (Boulenger) Rana larutensis Boulenger 1899 : 273. MATERIAL: BM 1967.2328-57, 1967.2489-91 (7 2, 17 5, 9 immature). HABITAT: 700-5000 ft in disturbed rainforest and in primary forest. The species was common at night on large, wet boulders in the fast-flowing stream at Base camp but became progressively rarer at higher altitudes, only two examples being found at 3400 ft and one at 5000 ft, but these occupying a similar niche to those in the lowlands. Rance: W. Malaysia. Rana blythi (Boulenger) Rana macrodon var. blythi Boulenger 1920 : 43. MATERIAL: BM 1967.2423-34, 1967.2435, 1967.2500-01 (6 9, 5 3, 4 juveniles). HABITAT: 500-1600 ft in both narrow and wide, clear streams among virgin forest, generally on stones or on the banks. REMARKS: Females vary in body size from 71-7 to 147-5 mm, five that have well developed ovarian eggs and convoluted oviducts ranging from 71-7 to 81-6 mm. The five males measure from 83-7 to 98-2 mm. In head proportions and in tibia length, as well as in other characters, they agree closely with the proportions given by Inger (1966) for a series of Malayan blythi. In the G. Benom specimens the head width to snout—vent ratio is 0:337—0°374 (29), 0-366-0-386 ($g), head length to snout— vent 0:383-0:421 (92), 0-443-0°477 ($d) and the tibia to snout—vent ratio is 0-546— 0°582 (22), 0:521-0:557 (gd). They further agree with Inger’s Malayan sample in lacking a dark, horizontal loreal stripe. Four individuals have a broad mid- dorsal light band, one has only a thin, light line. Inger states that male blythi lack vocal sacs, nuptial pads and any other distinctive (secondary sex) structures but in the G. Benom sample the bony mandibular projections are considerably better developed in males. RancGE: W. Malaysia, Singapore, Sumatra and Borneo. REPTILES AND AMPHIBIANS 65 Rana chalconota raniceps (Peters) Polypedetes vaniceps Peters 1871 : 580. MATERIAL: BM 1967.2436-59 (20 3, 4 9). Hapitat: The females were found on stones and boulders splashed by spray in the stream at Base camp (700 ft) and also among bank-side vegetation. All the males were around the quiet backwater to the stream, either on the muddy bank or sitting on branches of trees overhanging the water (see also Rana signata). CoLouR IN LIFE: Dorsum either grass green or yellow-green and usually with brown spots. Posterior aspect of thighs pale buff coloured. Upper lip creamy yellow, especially on posterior half, the light colour extending to the edge of the lip, as is usual in Bornean and Malayan populations (Inger 1966). Remarks: The entire series was collected between the 18th and 25th February when females contain enlarged pigmented ova and males were calling. In Borneo the breeding period extends at least from April to September (Inger 1966). In eight of the adult males the nuptial pad covers the medio-dorsal area of the base of the first finger, in the remainder the pad is less well developed and is partially or wholly divided into two. Rance: The subspecies vaniceps occurs in W. Malaysia and Borneo, and possibly also Sumatra and peninsular Thailand (Inger 1966). Rana hosii Boulenger Rana hosii Boulenger 1891 : 290. MATERIAL: BM 1967.2460-80, 1967.2481-88 (7 9, 21 J). Hapirat: All the specimens were obtained from boulders in the clear, fast flowing stream at Base camp, 700 ft, or from its banks and overhanging branches. COLOUR IN LIFE: The colour was very variable, the dorsum and flanks of some specimens being uniform grass green, of others olive green or green spotted with brown or brown spotted with green. The backs of the thighs were marbled dark green and brown; the tympanic area and the dorso-lateral fold dark brown. The upper lip and the spot at the jaw angle were yellow and the iris golden yellow. RancE: Thailand, W. Malaysia; Sumatra, Java and Borneo. Rana laticeps Boulenger Rana laticeps Boulenger 1882 : 20. MaTERIAL: BM 1967.2390-2420 (males, females, halfgrown & juveniles. Hasirat: All specimens were obtained in aquatic situations under forest from 600-3400 ft. Examples would invariably be found at night in quiet shallow pools at the edges of streams, lurking between stones or under wet fallen leaves and twigs. 66 A. G. C. GRANDISON Occasional individuals, usually females or immature specimens, were taken from crevices in rotten logs, from the clay banks of streams or from the gravelly stream bed. When disturbed Jaticeps either rapidly swims away or remains motionless, if in water lying with only the eyes protruding from the surface. The wide, fast- flowing stream at Base camp harboured this species only in a restricted area where the current slackened at the margins and shallow (less than 3” deep) gravel-based pools formed. At higher elevations examples were found at night in rock pools in a steep and exceedingly wet area after prolonged rains, about 20 yds from a fall of water in a narrow stream in the valley below camp 2. CaLL: The male calls from water, its head and upper jaw above the surface. The gular skin inflates and pulsates as the soft, melodious gargling sound, which goes up the scale, is uttered. Since no vocal sacs are present in this species it is presumed that when the male calls air that is being shunted to and fro to the buccal cavity causes distention of the loose plicate lining to the floor of the mouth and thus inflation of the gular skin. The mating call was recorded and males responded readily to the play back. COLOUR IN LIFE: The corrugated upper surfaces were generally orange-brown with darker brown interorbital bar, paired blotches on the middle of the back, lip spots, diagonal temporal band and crossbands on the limbs. Gular region mottled brown and white in females and immature individuals, paler and more greyish in males; belly grey/white. Undersurfaces of hind limbs in all but juvenile frogs vivid orange. None of the examples had yellow on the chest, belly or ventral surfaces of the legs as reported by Inger (1966) as occurring in Bornean examples. REMARKS: Eleven females with enlarged ova range in size from 36-9-42-0 mm (M = 39:7). Males in which the testes are apparently ripe and the mandibular odontoids well developed and sharp vary from 34-7—45-4 (M = 41-9 Ng); none has any trace of a nuptial thumb pad. RANGE: Assam to Thailand, W. Malaysia, Borneo (Sarawak). Rana limnocharis limnocharis Wiegmann Rana limnocharis Wiegmann in Meyen 1835 : 255. MaTERIAL: BM 1967.2492-99, 1967.2578 (3 9, 6 2). Hapitat: This species was found only in the cleared areas at Base camp, 700 ft. RANGE: From Ceylon and India to Taiwan and Japan, also W. Malaysia, Singapore, Sumatra, Java, Borneo and the Lesser Sundas as far as Flores. Rana luctuosa (Peters) Limnodytes luctuosus Peters 1871 : 579. MATERIAL: BM 1967.2371-77 (4 6, 3 gravid 9, 1 larva). Hasirat: All the examples were taken in or near flooded ruts on the logging tracks in disturbed forest between 700 and 1950 ft. REPTILES AND AMPHIBIANS 67 CoLour IN LIFE: Head and back rich chocolate brown bordered by narrow cream dorso-lateral line which extends from tip of snout along canthus, edge of upper eyelid to above the vent. Sides of body black. Limbs crossbarred pale grey and black; back of thighs black spotted with white. Ventral surfaces light grey. Flower’s (1896) colour plate of this species closely resembles the Benom specimens. Remarks: The size of the males (S~V 39:3-42:9 mm) and the absence of stripes on the undersurfaces of the limbs of this Benom series confirm Inger’s (1966) state- ment that Malayan examples of Juctuosa are not only smaller but lack the limb pattern of Bornean populations. The four males appear to be sexually mature and have well developed humeral glands which vary in length from 2:9-3-9 mm. The females with enlarged unpigmented ova measure 46:5, 47-9 and 51-7 mm. 575 1-1: III. The division of the outermost rows on the upper lip may be due to damage. The dental formula in the larva, which has hind limbs well developed, is Rance: W. Malaysia (Penang, Perak, Pahang and Selangor States), Borneo (Sarawak and W. Sabah). Rana miopus Boulenger Rana miopus Boulenger 1918 : 11. MaTERIAL: BM 1967.2421-2 (2 gravid 9). Hasirat: Both frogs were found on the main logging track within 4 mile of Base camp at a point where there is considerable secondary growth and wild bananas flanking the track at ca 750 ft. CoLouR IN LIFE: Upper surfaces orange-red, the dorso-lateral folds somewhat lighter. The post tympanic diagonal band, the spots in the groin as well as the streaks on the haunches adjacent to the dorso-lateral folds black. Back of thighs marbled black and grey. Legs with grey brown crossbars. Diagonal streaks across back also grey-brown. RanGE: Known only from the type locality in the southern part of peninsular Thailand, from one specimen obtained in the Chikus Forest Reserve, Perak and from individuals taken in Kelantan and in north, central and southeast Pahang, W. Malaysia. Rana nicobariensis (Stoliczka) Hylovana nicobariensis Stoliczka 1870 : 150. MatTERIAL: BM 1967.2725 (1 2), BM 1967.2389 (x juvenile). Hasitat: The female was collected at Base camp, 700 ft, in the large camp clearing in disturbed rainforest, while the juvenile was found on the wide track leading to the camp. CoLouR IN LIFE: No colour notes were made on the adult. Since at the time the frog was collected it was assumed to be another Rana chalconota it is likely that the 68 A. G. C. GRANDISON colour in life was similar in the two species. The upper lip of the juvenile pale green; iris reddish brown. RemARKS: Although this species superficially resembles Rana chalconota raniceps, in Benom individuals there is a marked difference in the size and proportions of gravid females, the nicobariensis specimen measuring only 46-8 mm in body length while the four adult females of chalconota (obtained within 200 yds of the nico- bariensis) varied from 49:6 to 55°7 mm, average 52:9 mm. Inger (1966) however reports no size difference between the species in Bornean populations, the size ranges for long series of females being 46-5-53:2 mm in nicobariensis and 46-0-59-4 mm in vaniceps with means 50-11 + 0-44 and 50-68 + 0-51 mm respectively. With only one example of nicobariensis it is unwise to draw any definite conclusions on Pahang populations. The most reliable distinguishing feature between the two species is the considerably less webbing in nicobariensis (see figures in Inger 1966). RANGE: Peninsular parts of Burma and Thailand, W. Malaysia, Sumatra, Java, Borneo to Bali and Philippines (Palawan). Rana plicatella Stoliczka Rana plicatella Stoliczka 1873 : 116. MATERIAL: BM 1967.2362-70 (2 3, 7 juveniles). Hapirat: One of the adult males was found on the bank above a soft muddy area with standing water, close to a rivulet that had little current and joined the left bank of the River Kerau tributary 200 yards further on. The other male was said to have come from the same area, about 300 yds downstream from Base camp and at approximately 700 ft. COLOUR IN LIFE: Upper surfaces mid-chestnut brown with darker brown glandular ridges and with dark crossbars on the limbs. One adult with a cream vertebral band. Throat white, chest and belly sulphur yellow, undersurfaces of hind limbs yellow speckled with brown. RANGE: Thailand south of Yala Province, W. Malaysia. Rana signata signata (Ginther) Polypedetes signata Giinther 1872 : 600. MATERIAL: BM 1967.2378-88 (10 3g, I 9). Hapirat: On Benom Rana signata seems to be limited to the lowland rainforest at 700 ft where it occupies a very restricted niche by secondary growth close to the fast flowing tributary of the River Kerau. A distinct preference was shown for a steep mud bank above a still backwater. The backwater, approximately 20 x 12 ft with clear water and a soft mud base, was about 3 ft deep and overhung by branches. At one end where there were large boulders sprayed by the force of the stream Amolops lavutensis occurred and at the base of the boulders, close to the water, REPTILES AND AMPHIBIANS 69 Bufo asper wascommon. The males of s7gnata were always on the bank 6-12” above the water calling from the ground but exceedingly difficult to locate by their call as they seem to throw their voices. The call is like a laugh, ha-ha-ha-ha-ha repeated quickly and quietly. Higher on the bank and closer to the forest litter and bases of trees and bushes or on branches of the trees overhanging the backwater males of Rana chalconota occurred and I agree with Inger (1969) that chalconota has stronger arboreal tendencies than signata for nowhere did I witness any evidence of signata having even partial arboreal habits. The only adult female that was obtained was collected by a porter at night from the ‘stream area at Base camp’; no further data are available. CoLouR IN LIFE: Dark brown or blackish on upper surfaces with pale apple green or cream spots and blotches on the dorsum and backs of thighs. The limbs with orange coloured cross bars, the bars often broken up into spots. From the tip of the snout, along the canthus and the edge of the upper eyelid an orange line extended dorso-laterally to the groin. In all the examples there is at some point a break in the dorso-lateral line. SECONDARY SEX CHARACTERS: Similar to those described by Inger (1966) for Bornean populations and no geographical variation in size seems to occur. RANGE: Peninsular parts of Thailand, W. Malaysia (Pahang and Kelantan), Sumatra and Borneo. Rana tweediei Smith Rana tweediei Smith 1935 : 62. MATERIAL: BM 1967.2358-61 (29, 1 halfgrown Q, I 3). Hapsitat: All were obtained in muddy areas under forest close to small streams, between 700 and 2500 ft. CoLouR IN LIFE: Dorsum grey and dark brown, the grey being more or less con- fined to a broad band between the front halves of the upper eyelids, to paravertebral longitudinal streaks, to the sides of the body, to limb cross bars and to vertical streaks on the upper lips. The rest of the upper surfaces were brown, the dorso- lateral glandular folds and interorbital bar rather darker brown. Belly lemon, undersurfaces of thighs and tibia vivid orange, throat white, mottled with brown. A vertebral line similar to that present in the type occurs in the male but is absent in all the females. In life this line was whitish, not yellow as in Smith’s (1935) description of the type. The sexes do not differ in their colours in life. RemarRKS. The two adult 99, S-V 39-9 and 41:8 mm, have enlarged ova with heavily pigmented poles. They are apparently the first females of this species to have been collected. The sexually mature J, S-V 41-1 mm, has, like the type, no secondary sexual characters. No size difference in the sexes occurs and the only external feature that may distinguish them is the male’s vertebral band, the fine white line on the hind limbs which in the Benom specimen is restricted to the heel and tarsus, and a greater number of tubercles on the back and limbs (see below). 70 A. G. C. GRANDISON The series has been compared with the holotype and paratypes and agreement is close. Although Smith mentioned the presence of tubercles on the upper eyelids of the holotype he omitted any reference to similar white tipped tubercles above the vent, on the haunches and especially on the hind limbs; such tubercles occur on these areas in both the type series and in all the Benom material but are less well developed in the females. Additional characters omitted by Smith but present in all examples are a feeble transverse groove on the head, immediately behind the dark interorbital bar, and one to three white spots along under-surface of forearm. RANGE: Previously known only from the type series (near R. Yum, Headwaters of R. Plus, E. Perak, W. Malaysia). Family RHACOPHORIDAE Philautus aurifasciatus (Schlegel) Hyla aurifasciatus Schlegel 1837 : 27. MATERIAL: BM 1967.2620-57 (10 9, 25 g, 3 juveniles). Hapsirat: This species was seen and heard only between 3400 and 5000 ft. Speci- mens were collected mostly at night. They were never more than 5 ft from the forest floor, generally on the leaves of small palms or other low plants and bushes. In the valley below camp 3 it was the most common species especially on and at the sides of the wide boulder strewn stream bed. COLOUR IN LIFE: There was marked individual variation both in pattern and in the shades of brown on the dorsum. A dark chocolate brown interorbital bar was generally evident. Some individuals have a bold dark brown H-shaped mark on the back and on each side of the cross bar of the H a roundish dark brown blotch. Other individuals have a khaki coloured mid dorsal band which extends from the tip of the snout, covers the entire upper surface of the head, narrows behind the eyes and passes back to the vent. A thin, dark vertebral line or a broader, dark brown vertebral stripe may also be present. Yet other individuals have only remnants of the H-shaped dorsal pattern, usually a pair of dorsolateral dark streaks and occasionally in addition the anterior dark blotch. The sides of the body and head were reddish brown with light spots on the haunches, groin, front and back of the thighs and on the undersurface of the tibia. Upper surfaces of the hind limbs crossbarred in dark brown. Ventral surfaces mottled greyish brown, especially on the throat. REMARKS. These specimens have been compared with the syntypes of Ixalus petersi Boulenger, considered by Inger (1966) to be conspecific with aurifasciatus, and with the syntypes of Ixalus larutensis Boulenger which Smith (1930) placed in the synonymy of J. petersi. It is not clear from Inger’s account of aurifasciatus whether he accepted Smith’s 1930 concept of Philautus peterst. Inger (1966) concluded that a conically tipped snout in awrifasciatus is a highly variable character that may or may not be developed in material from Borneo and Java, even in speci- mens from the same locality. In the G. Benom examples, the conical tip is more REPTILES AND AMPHIBIANS 71 often present and better developed in mature females than in males. A lingual papilla is generally present in both sexes. The outer edge of the fifth metatarsal and of the fifth toe bear a narrow crenulated fringe. SECONDARY SEX CHARACTERS: The Malayan examples are slightly larger at sexual maturity than the Bornean populations reported on by Inger (1966). The seven gravid females in the G. Benom material vary in body length 31-0-36-7 mm (average 33-6) and 25 males that have a nuptial pad developed measure 20-6—26-2 (average 23°8). The means of the body lengths of Bornean adults are 29°85 (92) and 21-78 (33). RANGE: From Thailand and Cambodia to Java, Borneo and the Philippines (Inger 1966). Philautus vermiculatus Boulenger Ixalus vermiculatus Boulenger 1900 : 187. MATERIAL: BM 1967.2658-61 (4 3). Hapsirat: On rattan and branches of trees in neighbourhood of camp 4 at 5000 ft. This species calls only from high up in the trees. The call crack-crack- crack-crack was heard at several points but the frogs were difficult to locate and collect. CoLour IN LIFE: The dorsum of two adult specimens was brown and fawn, the forearm, belly, thighs and flanks golden yellow or orange. The thighs of one individual were spotted yellow and brown. The smallest specimen had a green dorsum with an irregular brown hour-glass pattern and a yellowish throat. Iris silver-grey. REMARKS: This species was described as having a smooth dorsum (Boulenger Igoob) but examination of the type and of the type of Ixvalus brevipes Blgr. with which it was synomymized shows that both have small warts scattered over the upper surfaces, especially on top of the snout, above the supratympanic folds and on the heels. The G. Benom specimens also have warts, these being especially prominent on the upper eyelids, shoulders and tibias. Additional distinguishing features are the strongly oblique lores and flaring lips, the curved canthus, and the internarial distance greater than the distance between the naris and the anterior border of the eye and between two and three times the tympanic diameter (in adults twice). In the type as well as in the G. Benom examples the diameter of the third finger disk is consistently greater than the tympanic diameter and not subequal as stated by Boulenger (1goob). The four Benom examples vary in pattern from the symmetrical markings typical of brevipes (Boulenger 1908) to close vermiculations. A light line between the anterior halves of the upper eyelids and two oblique light lines from below the eye to the upper lip are constant. Pigmentation of the groin and of the front of the thighs is absent. RancGE: Thailand and W. Malaysia. 72 A. G. C. GRANDISON Rhacophorus appendiculatus (Giinther) Polypedetes appendiculatus Ginther 1858 : 79. MATERIAL: BM 1967.2503 (I 4). Hapitat: The specimen was found at night 6 ft from the forest floor on a tree in an area of secondary growth at the side of a track close to Base camp at 700 ft. COLOUR IN LIFE: Dorsum mottled light brown. Thighs pinkish orange on both anterior and posterior surfaces. Throat and belly yellow. Remarks: In all characters except snout shape this individual closely resembles the type of Rhacophorus chaseni (Smith 1924a) which Inger (1966) considered a synonym of R. appendiculatus. Like chaseni it has below the vent a pronounced transverse dermal flap with crenulated border interrupted on the midline, and from the heel to the 5th toe and also along the forearm to the 4th finger con- tinuous scalloped fringes. There is a scalloped seam too along the lower edge of the lower jaw. But while there is a small dermal projection at the tip of the snout it is very much less pronounced than in the gravid female holotype of chasent. Additional Malayan examples may show that there is a sexual difference in the development of this appendage. The webbing on the toes of the holotype of chasent does not extend to the disks, as described and shown in pl. I fig. 1 (Smith 1924a) ; instead, on the inner aspect of the second, third and fourth toes the web reaches only the distal subarticular tubercles and in this respect agrees with the G. Benom example. RANGE: W. Malaysia, Sumatra, Borneo, Philippines. Rhacophorus colletti Boulenger Rhacophorus colletti Boulenger 1890 ; 36. MaTERIAL: BM 1967.2504 (1 gravid 9). Hasitat: The specimen was collected at night on a leaf on the ridge E.S.E. of Base camp at about 800 ft. A Cyrtodactylus marmoratus was obtained on the same tree. COLOUR IN LIFE: Dorsum uniform pinkish brown. ReMArRKS: The specimen agrees well with Inger’s (1966) excellent description of the species except for the absence on the dorsum of the dark hour glass-shaped figure which apparently is usually present in Bornean and Thai examples. RANGE: Extreme southern States of Thailand, W. Malaysia (Pahang), Sumatra, Borneo and islands in the South China Sea. Rhacophorus leucomystax leucomystax (Boie) Hyla leucomystax Boie in Gravenhorst 1829 : 26. MATERIAL: BM 1967.2534-77, 1967.2579-89, 1967.2590-619 (27 2, 57 dg, I im- mature). REPTILES AND AMPHIBIANS 73 HasiraT: 600-1950 ft in both primary and secondary forest. The species was particularly common on stems of plants up to 7 ft from the ground, in flooded ditches and in temporary pools between 600 and 1700 ft around clearings and in other areas of disturbance associated with logging. Above 1700 ft specimens were found only on bushes flanking the track. RANGE: The nominate form occurs from Tenasserim through Thailand to Hainan and northern Vietnam and in W. Malaysia, Singapore, Sumatra, Java, Borneo and the Philippines. Rhacophorus macrotis Boulenger Rhacophorus macrotis Boulenger 1891 : 282. MATERIAL: BM 1967.2533 (1 3). Hasirat: No information other than ‘Base camp area, 700 ft’. CoLour IN LIFE: The ground colour of the upper surfaces was brick red, with a pair of darker spots between the eyes forming an interorbital bar interrupted medially; other median dark spots on the back. From behind the eye to the flanks a striking dark, almost black horizontal band which covers the tympanum and tapers slightly behind the arm; the band is bordered below by a cream line. A fine white line extends along the outer edge of the forearm and hand. A fine, white line above the vent; two white tubercles below the vent. Flanks marbled. Hind limbs with dark cross bars above. A prominent dark heel patch. Venter whitish, the belly finely speckled. RemaRKs: In the field this specimen was mistaken for Rhacophorus leucomystax, two leucomystax having a similar brick red dorsum and a somewhat similar appear- ance, but a closer examination revealed not only that this specimen is much larger than male leuwcomystax (S-V 60:5 mm) but that it has a sharper canthus, more vertical loreal region and more widely flaring lips, less finger webbing and more extensive toe webbing and in other characters too closely resembles R. macrotis which was not known to occur in the Malayan peninsula. Rance: W. Malaysia, Sumatra, Borneo, Natuna Is., Philippines (Palawan and Sulu Archipelago). Rhacophorus nigropalmatus Boulenger Rhacophorus nigropalmatus Boulenger 1895 : 170. MaTERIAL: BM 1967.2505-21 (3 9, 14d). Hasitat: On Gunong Benom the habitat and altitudinal distribution of this species closely paralleled those of Rhacophorus prominanus and R. veinwardtii. All these species were obtained from flooded, muddy ruts and other temporary pools on the main track, as well as from branches overhanging the pools, in forest between 700 and 1650 ft approximately, but R. nigropalmatus showed a preference for higher trees, bushes and palms and seem called from greater heights. 74 A. G. C. GRANDISON Catt: A loud, clear, teuk-teuk-teuk-teuk which somewhat resembles the tapping of a woodpecker. COLOUR IN LIFE: Grass green granular upper surfaces with minute white tubercles and occasionally also large, white blotches on the head and hind limbs. Flanks, inside of thighs and undersurfaces yellow. Finger and toe web jet black except for the distal margins which were yellow finely streaked with black. Upper surfaces of toes 1-4 yellow. Iris white. RANGE: Peninsular parts of Thailand, W. Malaysia, Sumatra, Borneo. Rhacophorus prominanus Smith Rhacophorus prominanus Smith 1924 : 185. MATERIAL: BM 1967.2530-32 (3 g), BM 1967.2662—4 (3 larvae). HasitaT: On branch overhanging puddle in track and in flooded ruts in the track at 800 ft, 1300 ft and 1650 ft. CoLouR IN LIFE: Upper surfaces of the body and limbs a soft blue-green with small brown spots; belly yellowish. Web between the fingers yellowish, toe web blood red between the 3rd and 4th and 4th and 5th toes. Iris pale buff, heavily streaked with red. LarvAE: Three premetamorphic tadpoles at stages XI—XVII, obtained from a pool on a logging track at ca goo ft are tentatively assigned to this species. Their size, very broad feet with full web extending to the toe disks suggests that they belong to the nigropalmatus, prominanus, reinwardtii group. Their dental formula, however, does not agree with that given for nigropalmatus (Inger 1966) and the unpigmented webbing on the toes makes assignment to reinwardti unlikely. A composite description of the three G. Benom tadpoles is as follows: body ovate, flattened above and below; eyes dorsal, not visible from below; diameter of eye subequal to eye-nostril and 4/5 internarial distance, 2/3-1/2 the interorbital dis- tance; spiracle sinistral, not tubular, below line connecting eye to root of hind limb, slightly nearer eye than root of hind limb; anus dextral, opening not reaching margin of ventral fin. Mouth ventral, subterminal; beaks black-edged, finely serrate, upper one a smooth arc; papillae small and homogeneous, continuous along lower lip and in 3—4 rows, confined to lateral corners of upper lip; dental formula I: 6-6/ 1-1: II; very narrow gaps between the two outermost divided rows of upper lip and the divided row of lower lip. Tail 0:54-0-6 of total length, weakly convex, deeper than body at centre, abruptly tapering in last quarter to slender, rather blunt tip; dorsal fin only very slightly deeper than ventral fin and both narrower than caudal muscle at centre. Colour in alcohol of head and body pale greyish brown above, white below, without pattern; tail fins pale grey, caudal muscle yel- lowish, without pattern. RANGE: Peninsular parts of Thailand, W. Malaysia, Nias Is., ? Sumatra. REPTILES AND AMPHIBIANS 75 Rhacophorus reinwardtii (Wagler) Hypsiboas veinwardtii Wagler 1830 : 200. MATERIAL: BM 1967.2522-29 (7 d, I 9). HapsitaT: On Gunong Benom this species occurs in similar areas and in somewhat similar habitats to Rhacophorus nigropalmatus and R. prominanus. It is a lowland species and was never seen or heard above 1600 ft yet the G. Benom examples appear to be the first record of the species from Malaya. Specimens were found on the wide logging track in depressions caused by tyre tracks and flooded by rains and in other temporary pools by secondary growth in disturbed forest and near to banana stands, as well as up to 12 ft from the ground on banana fronds, bamboo and bushes that overhung pools. CALL: R. reinwardtii calls from bushes, often bamboo, and from a lower level than R. nigropalmatus. Its call is a rat-tat-tat resembling a rattle or a wood- pecker’s tapping and is softer and quieter than the call of R. nigropalmatus. CoLour IN LIFE: Dorsum and upper parts of the limbs a soft apple green, sides of body brick red with large, black areas, marbling and spots all of which were outlined with sky blue. Inner surfaces of the arm, of the tibia and to a lesser extent of the tarsus black with pale blue spots. Posterior aspect of thighs grey with very fine black reticulations. Undersurfaces of throat and belly pale yellowish grey or chalky white with black marbling. Iris white. Web between all the toes and all the fingers from base to distal margin intense black with sky blue spots and streaks on both the upper and lower surfaces. Except for the outermost, the fingers and toes were similarly coloured. REMARKS: While these specimens agree with reinwardtii and differ from b1- maculatus Blgr. in the extent of the web on both fingers and toes, in the tympanic and finger disk sizes and proportions, in head proportions and in the development of dermal appendages (Inger 1966) they exhibit marked differences in size from the Javan and Sumatran populations of reinwardtii, and in colour pattern are unlike Javan rveinwardtii although they show certain resemblances to the var Jateralis described and figured by Werner (1900) from a single individual obtained at Batu Bahru, Sumatra (Basel Museum 1192). The G. Benom males vary in body length from 59-66 mm (average 63 mm) and the female measures 80:3 mm. Inger (1966) gave a range of 46-55 mm for mature male reinwardtii while Wolf (1936) recorded 56 mm for males and 76 mm for females. Five examples of reinwardti obtained in Java and in the British Museum collections have body lengths of 52 mm (3 33) and 70 and 71 mm for the two females. As can be seen from the description of colour in life, in the G. Benom sample black pigmentation was much more extensive than in Javan populations of reinwardtii (Schlegel 1844 plate 30, fig. 4, Van Kampen 1923, fig. 28) and in none of the Malayan examples is the black pigment on the finger and toe web restricted to large black spots between the second and third and third and fourth fingers and between all the toes except the first and second, as occurs in Javan reinwardtit. RanGE: W. Malaysia (Pahang), Sumatra and Java. c 76 A. G. C. GRANDISON SAURIA Family AGAMIDAE Aphaniotis fuscus Peters Otocryptis (Aphaniotis) fusca Peters 1864 : 385. MATERIAL: BM 1967.2837-39. Hapitat: The specimens were found at 7oo ft and 2700 ft. No further infor- mation on their habitat is available. RanGE: Peninsular parts of Thailand, W. Malaysia, Borneo, Natuna Is. Goniocephalus borneensis (Schlegel) Lophyurus borneensis Schlegel 1848 : 6. MaTERIAL: BM 1967.2840-41 (2 3). HasitatT: The smaller specimen was found on a bush in disturbed forest near Base camp, 700 ft, the larger was caught by a dog near the track in the forest at 1900 ft. COLOUR IN LIFE: The throat pouch had on each side a vivid pink patch surrounded by bright ultramarine blue. The base of the pouch was yellow green. The trunk was greyish green dorsally with brown edged ocelli on the flanks. Tail black and cream banded. RanGE: Peninsular parts of Thailand, W. Malaysia, Borneo. Draco fimbriatus fimbriatus Kuhl Draco fimbriatus Kuhl 1820 : tor. MaTERIAL: BM 1967.2798-800 (I 9, 2 3). Hapitat: The larger of the two males (107-3 mm) was found lying apparently stunned on the verandah of a hut at 700 ft in the forest clearing in early morning. The other male (104-8 mm) was obtained at ca 1900 ft on the ridge southwest of camp 2. The female (101-8 mm) was shot on a tree at 35 ft from the ground in disturbed forest. The tree, 18” in diameter and at the side of the logging track between Base camp and Kampong Damak at ca 600 ft, had Dyaco volans and D. punctatus on it as well, but at a lower height from the forest floor. COLOUR IN LIFE: The general appearance and colouration of the upper surfaces of this species were of rough bark in shades of grey, black, brown and reddish brown. The wing membranes were indistinctly longitudinally striped black and greyish brown above; their lower surfaces were orange with black spots (9) or light grey with dark brown spots (g). In the female the belly, undersurfaces of the base of the tail and of the thighs were orange, the throat pale grey mottled with darker grey and the gular appendage and wattles coral pink. The males had similarly coloured dewlaps and wattles and there were also coral spots on the cheek, chin, REPTILES AND AMPHIBIANS 77 and sides of the head. Rather duller orange marks were present on the vertebral region of the large male. RANGE: Southern provinces of peninsular Thailand, W. Malaysia, Singapore, Gt. Natunas, Sumatra, Java, Borneo. Draco formosus formosus Boulenger Draco formosus Boulenger 1900 : 190. MATERIAL: BM 1967.2801-26 (5 9, 19 g, 2 juveniles). Hasirat: The altitudinal range of this species on G. Benom is 700-2500 ft but above camp 2 (1700 ft) it was very rarely seen and only five of the series were ob- tained from above 1800 ft. All specimens were on trees at heights varying from 8-20 ft from the forest floor. D. formosus was more closely associated with the trees skirting the camp clearing and the wide logging tracks than other Dracos and was the commonest species of Draco in the area. CoLouR IN LIFE: Head and body brown above with obscure darker markings. A dark brown transverse band across the upper eyelids and a median dark brown interorbital spot present in both sexes. A pair of jet black spots on the occiput and a pair of more widely separated dark spots farther back on the neck constantly present in the males. Wing membranes mostly yellowish above with five black, somewhat wavy transverse lines, the bands narrower than the interspaces, the distal portion of the wing maroon. Lower wing surfaces similar in colouration but the bands often indistinct. Gular pouch thin and translucent, a whitish grey. Throat and undersurfaces of the wattles blood red. Chin grey with dark reticula- tions. Remarks: The females which contain three or four, usually four, large, broadly oval eggs vary in snout to vent length 74:8-87-6 (average 80-0 mm). Males vary in length from 80-4-102-7 (average 93-7 mm). RANGE: The nominate form is said to be restricted to S. Thailand and W. Malaysia (Hennig, 1936, Taylor 1963). Draco maximus maximus Boulenger Draco maximus Boulenger 1893 : 522. MaTERIAL: BM 1967.2828 (1 3). Hasirat: On tree flanking logging track at ca 800 ft at junction of the track between Base camp and K. Damak and the path leading to camp 2. RANGE: W. Malaysia, Gt. Natunas, Sumatra, Borneo. c* A. G. C. GRANDISON “1 ios} Draco melanopogon Boulenger Draco melanopogon Boulenger 1887 : 492. MATERIAL: BM 1967.2830-34 (2 9, 3 3). Hasirat: All specimens were shot in trees among secondary growth flanking the logging tracks and the trail to the stream below Base camp and between 600 and tooo ft. COLOUR IN LIFE: Head and vertebral region brown becoming green on the sides of the trunk. Upper surfaces of limbs greenish brown. Some indistinct dark brown cross bars and spots across head and back. Wing membranes black with numerous yellow spots on both upper and lower surfaces. Gular pouch yellowish green in the female, jet black in the male; wattles white in both sexes. Throat, chest and under- surface of forelimbs orange in the male; undersurface of tail similarly coloured. REMARKS: The two females captured 23/24th February are gravid. One measur- ing 82 mm snout-vent contains only one oviducal egg, 8-0 mm long, the other individual, 79 mm, has the normal melanopogon complement of two, each 5-5 mm in length (Inger & Greenberg 1966). The eggs are almost spherical and not distinctly pointed as described by Hendrickson (1966). RANGE:- Thailand south of the Isthmus of Kra, W. Malaysia, Gt. Natunas, Sumatra, Borneo. Draco punctatus Boulenger Dyvaco punctatus Boulenger 1900 : 189. MATERIAL: BM 1967.2828 (x 3). Hasirat: The specimen was taken 25-35 ft from the forest floor on a tree near the main logging track at ca 600 ft. In the same tree simultaneously were two other species of Draco, fimbriatus and volans. COLOUR IN LIFE: The upper surfaces of the head, body and limbs grey and brown with scattered dark brown speckling. Wings reddish brown above, longitudinally streaked with light grey. Gular pouch pale grey at base, brilliant yellow towards apex, chin yellow spotted grey. Wattles intense lemon yellow. Undersurfaces of the trunk grey-green, of the wings greyish yellow with scattered black spots. RANGE: Southern provinces of peninsular Thailand, W. Malaysia, Sarawak. Draco quinquefasciatus quinquefasciatus Hardwicke & Gray Draco quinquefasciatus Hardwicke & Gray 1827 : 219. MATERIAL: BM 1967.2829 (x 9). Hasitat: Shot on 6” diameter tree, to ft from the forest floor, on the ridge at ca 800 ft NNE of Base camp at noon on 25 February. The individual is gravid and contains four well matured oval eggs, the largest 15-7 mm long. REPTILES AND AMPHIBIANS 79 CoLour IN LIFE: Ground colour of body yellow-green with broad greyish black cross bands. Wings above orange-red with five broad black cross bands, each with a single row of small white spots confluent with those of the trunk, the orange be- coming yellowish medially. Throat green, speckled brown, the wattles black at their bases with a white band, yellowish anteriorly. Gular appendage yellowish orange, belly cream. Undersurfaces of wings yellow becoming orange laterally and posteriorly, with five broad black bands each of which bordered by greyish green, the most posterior band short and not extending to the lateral wing margins. RANGE: Extreme southern provinces of peninsular Thailand, W. Malaysia. Draco volans volans Linnaeus Draco volans Linnaeus 1758 : 199. MATERIAL: BM 1967.2835-36 (I 9, Id). HasitatT: The female which contains three large (13-4 mm) eggs was seen on the ground in the kitchen quarters of Base camp 700 ft. When disturbed it ran up a low leafy tree and was shot. The male was found 25-35 ft from the forest floor on a tree near the main logging track at ca 600 ft. On the same tree and at the same time D. f. fimbriatus occurred but at a greater height, and nearer the ground an example of D. punctatus was obtained. COLOUR IN LIFE: Above, the head, body and wings in both sexes were brown and grey, the grey on the back forming discreet transverse series of spots and lines which extended to the wings. There was a dark brown interorbital spot and in the female a dark medial nuchal spot. The gular pouch in the female was a vivid turquoise blue, in the male brilliant yellow, speckled black at its base. The inner surfaces of the wattles were greyish white. The eyelids in the male were sky blue. The undersurfaces of the trunk and limbs were pale greenish blue medially and yellow- green on the sides (2) or powder blue (g). In the female the undersurfaces of the wings were vivid yellow with black interrupted transverse lines and spots, while those of the male were powder blue with black interrupted transverse wavy bands. RANGE: Peninsular parts of Thailand, W. Malaysia, Singapore, Sumatra, Java, Borneo, Natunas, Philippines (Palawan). Family VARANIDAE Varanus bengalensis nebulosus (Gray) Monitor nebulosus Gray in Cuvier 1831 : 27. MATERIAL: BM 1967.2842 (1 juvenile). Hasitat: The example was found during late afternoon at the entrance to an elliptical hole 18 ft from the forest floor in a tree by the trail, downstream from Base camp, 700 ft. RANGE: The subspecies nebulosus occurs from S. Burma to S. Vietnam and south- wards throughout W. Malaysia and Java. 80 A. G. C. GRANDISON Varanus salvator salvator (Laurenti) Stellio saluatoy Laurenti 1768 : 56. MATERIAL: BM 1967.2843 (juvenile 9). HapsiratT: This specimen, which weighed 4 lbs and had a total length of 1450 mm, was obtained from the banks of the stream about 4 mile below Base camp and at about 650 ft. Its stomach contained a mouse deer. RANGE: The nominate subspecies occurs from Ceylon and India to Thailand, Vietnam and Hainan, W. Malaysia and islands to the south and east as far as northern Australia. Family GEKKONIDAE Cnemaspis affinis (Stoliczka) Cyrtodactylus affinis Stoliczka 1870 : 167. MATERIAL: BM 1967.2781 (x juvenile). Hapsitat: Camp 2 clearing, 1700 ft. RANGE: Southernmost provinces of Thailand, W. Malaysia. Cnemaspis kendallii (Gray) Hetevonota kendallit Gray 1845 : 174. MaTERIAL: BM 1967.2782 (1 9). Hasirat: On kitchen table at Base camp during afternoon, 700 ft. RANGE: Thailand, W. Malaysia, Singapore, Gt. Natuna Is., Borneo. Cyrtodactylus ?brevipalmatus (Smith) Gymnodactylus brevipalmatus Smith 1923 : 48. MATERIAL: BM 1967.2783 (I 3). Hasirat: At night on leaf litter of forest floor near Base camp clearing and stream at 700 ft. REMARKS: The G. Benom example differs from the holotype, from a juvenile also obtained in Nakhon Si Thammarat and from a female collected in Raheng, N. Thailand 2500 ft, in having fewer trunk tubercles and in their being disposed in approximately three longitudinal paravertebral rows that do not extend to the flanks. It further differs from the type in lacking femoral pores and in having only eight preanal pores. The tail is not cylindrical but is noticeably depressed and has almost vertical sides so that in section it is squarish. Tail denticulation appears to be less pronounced in the type but this may be artifact and due to preservation although the pointed tubercles along the dorsolateral edges are certainly shorter REPTILES AND AMPHIBIANS 81 than in the G. Benom example. The toe web, the number of ventral scales at midbody between the lateral folds (44) and the number of the digital lamellae agree with the type and the other examples, but dilation of the basal phalanges appears to be greater. Smith’s figure of the toes of brevipalmatus (1923, pl. 5) is inaccurate in omitting the plates under the distal portion of the distal halves. RANGE: Known only from specimens obtained in the Nakhon Si Thammarat Mts., Peninsular Thailand, from Raheng, N. Thailand (Smith 1935), and from the Gunong Benom individual. Cyrtodactylus consobrinus (Peters) Gymnodactylus consobyinus Peters 1871 : 569. MATERIAL: BM 1967.2784 (1 juvenile). Hasirat: The specimen was smoked out of a hole in a live tree in the valley below camp 2, at about 1600 ft. RANGE: W. Malaysia and Borneo. Cyrtodactylus marmoratus (Kuhl) Phyllurus marmoratus Kuhl in Fitzinger 1826 : 47. MATERIAL: BM 1967.2785-95 (6 9, 43). HapitatT: The series was obtained in forest between 700 and 3500 ft. Most examples were found at night on branches and trunks of trees, one being on the undersurface of a large, rotten tree trunk that had fallen across the boulder strewn stream bed in the valley below camp 3, another inhabited a hole between the roots of a 6” diameter tree close to the camp 3 huts and emerged from its hole soon after dusk to crawl up the trunk. At the lower elevations a few individuals were found on the high bank at the side of the logging track. REMARKS: These specimens agree well with all the diagnostic characters of marmoratus except for the position and number of pores in the males. The male C. marmoratus is said to be characterized by both preanal and femoral pores and a longitudinal preanal groove, the preanal pores numbering 12 or 13, the femoral pores 4-6 on each side (Boulenger 1912) but while these G. Benom specimens have a row of enlarged femoral scales none of the scales has a pit or pore and there are, at the most, only five preanal pores in a V-shaped line. They vary in body size from 55:4-61:3 mm. RANGE: Thailand, W. Malaysia, Sumatra, Java, Borneo, Celebes, New Guinea, Christmas Island. 82 A. G. C. GRANDISON Gehyra mutilatus (Wiegmann) Hemidactylus mutilatus Wiegmann in Meyen 1835 : 238. MATERIAL: BM 1967.2796-97 (I 2, I 3). HasiraT: Both specimens were smoked out of live trees in the forest near camp 2 at 1700 ft. RANGE: Widely distributed from Ceylon and S. Burma to S. China, Japan, Thailand, W. Malaysia, Indo-Australian Archipelago, Oceania and islands in the Indian Ocean. Family SCINCIDAE Lygosoma (Scincella) vittigerum vittigerum Boulenger Lygosoma vittigerum Boulenger 1894 : 615. MATERIAL: BM 1967.2854 (x juvenile). Hapitat: The specimen, which has a body length of only 17-5 mm, was found during the morning on the bark of a tree close to camp 2 at 1700 ft. RANGE: Peninsular parts of Burma and Thailand, W. Malaysia, Sumatra, Men- tawei Is., Borneo. Mabuya multifasciata (Kuhl) Scincus multifasciatus Kuhl 1820 : 126. MATERIAL: BM 1967.2844-53 (4 adults, 6 juveniles). Hapirat: This species was the most common lizard in and around camp at 700 ft, invariably frequenting secondary growth. Individuals were obtained from the ground, on and among fallen leaves, on piles of felled branches, on a pile of sand at the edge of the track, as well as apparently asleep on the trunk of a tree. COLOUR IN LIFE: Taylor’s description (1963) of the colour pattern of this species applies equally well to the G. Benom examples but on the chin, throat and chest of one adult male was mustard yellow, the rest of the ventral surfaces being a greyish green. From the tympanum to the flanks this individual was a reddish orange. The lateral ocelli may or may not be clearly marked and the dark brown lateral stripe may or may not be bordered above by a light, longitudinal dorsolateral stripe 14 scales wide. ReMARKS: The midbody count in these examples is 34(7) and 32(3). The first loreal is usually lower than the second, rarely of similar height. In each specimen the frontonasal forms a suture with the nasal and generally a broad suture. Both the head shields and the dorsal trunk scales have dark streaks at their edges, usually at the posterior borders of the head shields and at the lateral edges of the body scales. REPTILES AND AMPHIBIANS 83 RANGE: From N.E. India to S. China, Thailand, W. Malaysia, Singapore, Sumatra, Java, northwards to the Philippines and eastwards to New Guinea. Sphenomorphus indicus indicus (Gray) Hinulia indica Gray 1853 : 388. MaTERIAL: BM 1967.2855 (1 juvenile). Hapsirat: On fallen tree trunk and among dead leaves on peaty ground at 3650 ft during the afternoon. RemArKS: This individual, and two juveniles obtained at Bukit Fraser which were reported on by Smith (1922b & 1930), differ somewhat from the nominate form of indicus. They have a higher midbody count (36) than in both Tennasserim examples (30-32: 32 in the types of zebratum) and the average N.W. Thailand speci- men (34:3: N 17) (Taylor 1963) and the number of subdigital lamellae on the fourth toe is slightly greater (22-24) than in the adult zebratuwm syntype, in N.W. Thailand examples (18-19) and in the indicus types (19 : 20, 18 : 20). Both Smith (1935a) and Taylor (1963) err in stating that the frontal is as long as—or longer (Smith)— than the combined parietals for while this is so in the syntypes of indicus and in the adult zebratuwm syntype, in the juvenile zebratwm syntype and in both the Gunong and Bukit Fraser examples the frontal is 3/4 the distance from its posterior border to the posterior border of the parietals, measured along the midline. RANGE: From India to central China and southwards to W. Malaysia. SERPENTES Family TYPHLOPIDAE Typhlops diardi mulleri Schlegel Typhlops Miilleri Schlegel 1839 : 32. MaTERIAL: BM 1967.2267. Hasitat: Among leaf litter at base of tree in Base camp clearing, 700 ft. RanGE: Tenasserim through Thailand to S. Vietnam, also W. Malaysia, Singapore, Sumatra and Borneo. Family NATRICIDAE Natrix trianguligera (Boie) Tropidonotus trianguligerus Boie 1827 : 535. MaTERIAL: BM 1967.2273 (I 9). RemARKS: The specimen was found at night swimming in a still backwater of the stream at Base camp, 700 ft. In life the sides of the anterior half of the trunk were brick red, the upper surface of the head dark olive, the iris light olive. Upper labials and mental pale buff-coloured, venter pale yellowish. 84 A. G. C. GRANDISON Scale rows. ; : 5 a 20) Ventrals é : : ‘ . 139 +2 Sub caudals . : ; a OR RANGE: Peninsular parts of Burma and Thailand, S. Vietnam, W. Malaysia, Singapore, Sumatra, Java, Mentawei Is., Borneo, Celebes. Amphiesma sarawacensis (Ginther) Tropidonotus savawacensis Giinther 1872 : 596. MATERIAL: BM 1967.2274-5 (2 3). REMARKS: The two specimens appeared within five minutes of each other on the soft peaty soil behind the log fire in camp 4 (5000 ft) at 1300 hrs. Their colouration was as described by Tweedie (1954) for this species. 2274 2275 Scale rows 5 d : 2 Ly 17 Ventrals ; ; ; . 139 +2 140 +2 Subcaudals.. : : a 32+ The gall bladder is situated deep to ventral 50. RANGE: W. Malaysia, Borneo. Macropisthodon rhodomelas (Boie) Tropidonotus yhodomelas Boie 1827 : 535- MATERIAL: BM 1967.2282 (I 3). REMARKS: Found midmorning on the ground among forest litter at the timber stack at junction between main path from Base camp and track to camp 2. Scale rows. : : 5 5 iG) Ventrals : : : s . 132 +2 Subcaudals . : : : 54 RANGE: Peninsular Thailand, W. Malaysia, Singapore, Sumatra, Java, Borneo. Family COLUBRIDAE Elaphe flavolineata (Schlegel) Coluber flavolineata Schlegel 1837 : 141. MATERIAL: BM 1967.2287 (1 9). Hasitat: On track above camp 2 at approximately 1800 ft. COLOUR IN LIFE: Body green above, the dorsal scales edged with black, with indistinct yellowish transverse bands on the mid body that became orange farther back and more clearly defined. Ventrals yellow and narrowly edged with black, REPTILES AND AMPHIBIANS 85 underside of tail with broad orange bands, 6-7 scales in length on a greenish ground. Upper labials vivid yellow. A black temporal blotch. Scale rows. 5 2K) Ventrals , 2236 Subcaudals_. . 116+ (tip of tail missing) Upper labials : 9g 5th and 6th entering eye Temporals. : 2+2 Rance: Andamans, Peninsular Burma and Thailand, W. Malaysia, Singapore, Sumatra, Java, Mentawei Is., Borneo, Celebes. Gonyophis margaritatus (Peters) Gonyosoma margaritatum Peters 1871 : 578. MATERIAL: BM 1067.2278 (1 3). REMARKS: This specimen was found at about 2750 ft crossing the track between camps 2 and 3. In life it was most beautifully coloured, the upper parts being pale green with the dorsals heavily margined in black and with yellowish, broad cross bands on the posterior two-thirds of the body and tail that became brick red towards the tail. On the undersurfaces of the snake the mental and first and second labials and sides of the snout were orange, the rest of the labials and the throat yellow, the ventrals as far as V34 were also yellow and farther back an orange midventral line developed that became deep orange with blackish cross bars. There was a deep black temporal band. Scale rows ; : : 2) x9 Ventrals : ; : . 238 +2 Subcaudals .. ? 2 9 AS) Totallength . j . - 570mm RANGE: W. Malaysia (Perak, Pahang, Kelantan), Borneo. Liopeltis longicauda (Peters) Ablabes longicaudus Peters 1871 : 574. MATERIAL: BM 1967.2268-70 (2 3, I 9). REMARKS: 750 ft. Two of the specimens were found in coitus on the main track leading from Base camp at midday on 21st February. The third specimen was obtained in the same area. All three differ from Tweedie’s description (1957) of the species in having the venter from the throat to the tail tip yellow. The chin was white in life and the head and nape markings white. 2268 2269 2270 Sex . . : 3 Q 3 Scale rows . A ers 13 13 Ventrals.. 5 26300) II4 116 Subcaudals . I2+ 24 +1 54+ (tail damaged) 86 A. G. C. GRANDISON The gall bladder is situated deep to ventral 74 in the male and deep to ventral 79 in the female. Liopeltis baliodeirus (Boie) Coronella baliodeiva Boie 1827 : 539. MATERIAL: BM 1967.2271 (1 d). REMARKS: Found during late afternoon at about 1000 ft on the track between Base camp and camp 2. The undersurfaces of the posterior half of the body and all of the tail were orange in life. Scale rows . ‘ A : els Ventrals . : ; : peer 2 Subcaudals P : 2 OF RANGE: Peninsular Thailand, W. Malaysia, Singapore, Sumatra, Java, Natuna Is., Borneo. Pseudorhabdion longiceps (Cantor) Calamaria longiceps Cantor 1847 : 63. MATERIAL: BM 1967.2272 (1 9). REMARKS: 700 ft. The specimen was dug out of a rotting hollow log in the forest near Base camp. In life the head was a dull blood red, the collar yellowish and the trunk dark brownish red. Scale rows . : : F . 2 wg Ventrals. : ; : 2) LS) Subcaudals : : : : 5 8) RANGE: Southernmost provinces of Thailand, W. Malaysia, Singapore, Sumatra, Borneo, Celebes and ? Philippines (Taylor 1965). Family DIPSADIDAE Calamaria lumbricoidea Boie Calamaria lumbricoidea Boie 1827 : 540. MATERIAL: BM 1967.2279 (1 3). REMARKS: The individual was found at 21 00 hrs. crossing the earth floor of the kitchen at Base camp (700 ft). The dorsum except for the upper labials is brown and without stripes and most of the body scales have a darker brown apical spot. The lips are cream. The venter has black bars, each bar covering two adjacent ventral scales and separated from the next by a varying number of yellowish scales but usually by 3, 4 or 5. The absence of stripes in this specimen, coupled with a ventral count of 172 and four gulars between the posterior chin shields and the first ventral, agree better with individuals obtained from states on the western side of the peninsula than with Pahang Jwmbricoidea (Inger 1965). REPTILES AND AMPHIBIANS 87 The hemipenis is forked and papillate. Scale rows . : : j - 13 Ventrals : : ; . 172 +1 Subcaudals : : F 5 oh Supralabials : : : ‘ 6 (3-4) Temporals . . : i : o+1 RANGE: Southern Thailand to Java, Borneo and Leyte (Inger 1965). Lepturophis borneensis Boulenger Lepturophis borneensis Boulenger 1900 : 183. MaTERIAL: BM 1967.2294 (I 3). Remarks: The G. Benom individual appears to constitute the first record of the occurrence of Lepturophis in the Malayan Peninsula. Previously this monotypic genus was known only from the type obtained in Sarawak and from a subsequent series of twelve in the Field Museum Chicago, from Sabah and Sarawak. Fic. 2. Lepturophis borneensis (B.M. 1967.2294). Boulenger (1g00a) likened the teeth of Lepturophis to those of Lycodon and gave the formula of the type specimen as 6 +6. The type specimen has not been examined by me; it is probably in the Kuching museum, Sarawak, but con- firmation has not been received. The G. Benom example has been compared with the Field Museum series assigned to Lepturophis which in scale counts and in denti- tion agree favourably with the G. Benom specimen. See table A, p. 47. The only resemblance between the teeth of Lycodon and those specimens examined here is in the large fang-like anterior maxillary tooth followed by a diastema then a series of small teeth. In the G. Benom individual the maxillary formula is 6 + 6 + 3, the first six gradually increasing in size, the second group of six being small and of more or less equal size while the posterior group are again fang-like. The left maxilla of FMNH 158650 which was dissected out appears to have a formula of 6 + 3 +3 but FMNH 148895-6 have series of six small teeth in the centre of the maxillae. These enlarged D 88 A. G. C. GRANDISON rear teeth are of a curious shape; their medial surfaces are flat but on the lateral sur- faces flattening is restricted to the anterior and posterior surfaces of the teeth with the central portion of each tooth rounded externally and thus semicircular in transverse section. The flat flanges on the posterior and anterior borders of the tapering teeth give them sharp, cutting edges. Similar lateral compression of the anterior Fic. 3. Maxilla and maxillary teeth of Lepturophis borneensis (B.M. 1967.2294). and posterior surfaces of the maxillary teeth occurs in Xenopeltis (Thomson 1913) but in that genus the teeth have blunt tips and a blade-like appearance and moreover are set at an oblique angle to the long axis of the jaw. Lateral compression of the teeth is also developed in [gwanognathus which however has spatulate, chisel-shaped teeth with broad tips and of quite a different type. The notched and grooved ventrals, coupled with the extremely slender, elongate body (total length of G. Benom example 167-6 cms) suggest that Lepturophis is an arboreal snake, although the Malayan example was found on the ground at night on the main track near Base camp at 700 ft and the Field Museum series was collected in or on the banks of forest streams. In life the snake was a uniform dark grey dorsally with a whitish venter. RanGE: W. Malaysia (Pahang), Borneo (Sabah, Sarawak). Macrocalamus jasoni sp. nov. HoLotyPe: BM 1967.2283, a gravid female from leaf litter on track below camp 5 at about 5800 ft, Gunong Benom, C. Pahang, W. Malaysia, collected by John A. Bullock, April gth 1967. PARATYPES: BM 1967.2284, female, from leaf litter on forest floor of summit knoll above camp 5 Gunong Benom, 6500 ft collected by Lord Medway, March 27th 1967. BM 1967.2285, female, locality as holotype collected by Bah Chung, April oth 1967. DESCRIPTION OF HOLOTYPE: Midbody scales in 15 rows, ventrals 131, subcaudals 17. Loreal precisely twice as long as deep, one preocular, one postocular, 1 + 2 temporals, 8 upper and 7 lower labials, the fourth and fifth upper labials entering the eye, the fourth making contact with the loreal. Gall bladder deep to ventral 96, maxillary teeth 11, palatine teeth 11, pterygoid teeth 19. The colour in life was iridescent black on the dorsum with a conspicuous pair of dorsolateral yellow ochre 89 REPTILES AND AMPHIBIANS €+ze+2 Nn ++ Nn + a Nn a ++ is] + + a a + NNAA AN +++ ++++4++4 NAAN AN anaanaaammnmm J T Nn Nn z+z Jol|/€ + z2+2z z+7z+2 z+z syerodura WH a ee ()6 (SE) (*) 8 I z 4(6) 6 (S-€) g I z (S) 6 (S-€) g I Zz (S) 6 (S-€) g I Zz (S) 6 (S-€) g I Zz (S) 6 (S-€) g I z (S) 6 (S-€) g I z (S) 6 (S—€) g I z (S) 6 (S-€) g I z (S) 6 4 (S-€) 6 I z (S) 6 (S-€) g I z (S) 6 (S-€) g 9-£) 8 I tc (S) 6 §(S—£) I z (S) (S-€) g reynooaig = Te[N90}sog s[eIqey s[erqey IOMO'T aoddq + Sgr s+ Slr S + 9L1 S + 90z s+ 141 S + £0oz tZor S + blr S + 041 S + 6g1 + €11 s + zgr s+ 641 £61 2) z+ Lhe z+ 1€z z+ 6zz z + €bz z + o€z z+ €bz Zz + gzz z+ of z+ Szz z+ bbz z+ o€z z+ Lez z+ gtz z+ 1bz A susuaausog swqgoanjdaT Ut s}unod a[eoS VY alavy “Suisstu Z ynoqy |, ‘yerodwi9} raddn puz oy} ut yee1q e Aq pasnes oq Aeur yerodu193 raddn 443 pry || *soyozeros Aq poinosqo seiqe] 1oddn 49] yI9 pue 43S usaM}0q OIN4Ns Jo UrSIIO TesIOq § “Sursstur g 10 § ynoqy t “yynoul ey} WO pepnyoxe st pif pue puz oy} udemjoq ,,JeIqe] Joddn,, 1oy}0ue sapis y}0q UG } “posny + 2 € Opls HOT * Li Lt b6zz'Lo61 wousg * © 19) Wa rad “ Td 1SggSt & « “ oSogSt 2 « “ 6bogStr 5 o “ L6gghx ‘2 % FS gogghr 6 ‘aia S6gghr fo} “ef “e b6gghr 5 “ o €6gghr 2 us a 6o0LSbr & ‘aAiq pre yemeres Loggtr rs) g6SEo 5 “4SIq LO6SE9 ueyepues :yeqes HNWH é yemeres Hd AL oS) AyyesoT = “ON uInesnyL go A. G. C. GRANDISON stripes which extend from the temporals to the tip of the tail. These stripes which are two scales wide and are separated from each other by three scales are bordered below by black, the black extending from the fourth longitudinal dorsal row of scales to the lateral fifths of the ventral scales. The ventrals, apart from their black lateral margins and medial black flecks on ventrals 1-8 are bright yellow. The subcaudals are yellow except for occasional black speckling at their medial margins. Fic. 4. Macrocalamus jasoni Holotype. The head is brownish yellow with blackish areas on the prefrontals, loreal, pre- and postoculars, anterior temporal and labials, except for the sixth upper labial which is entirely yellow. From the last upper labial to the last lower labial the black coloura- tion forms a broad but rather ill-defined vertical bar. The mental and first lower labials are black and the black extends on to the anterior chin shields where it surrounds a yellowish oval patch on each shield. Total length 760 mm, midbody girth 237 mm. VARIATION: The paratypes are similar to the holotype in dorsal colouration and in their stocky build but are considerably smaller in total length (380 mm and 550 mm). Apart from black tips to the outer margins of the ventral scales the undersurfaces of 2285 are a uniform yellow but in 2284 most ventral and subcaudal scales have a medial black patch or streak in addition to the lateral black marks In the paratypes the ventral and subcaudal counts are 132, 134 and 22, 20, and the arrangement of the head shields corresponds closely with that of the holotype. The gall bladder was situated deep to ventral 96 in BM 1967.2285 and V88 in BM 1967.2284. RELATIONSHIPS AND ECOLOGY: In its pattern and in its considerably larger size and much more robust proportions M. jasoni differs strikingly from the two other species of Macrocalamus that are known. It is further distinguished from M. lateralis Giinther by its higher number of pterygoid teeth and from M. tweediet Lim by its lower ventral count. Neither lateralis nor tweediei has dorsolateral stripes, lateralis commonly being a uniform reddish brown above but occasionally havinglight chevrons on the head and nape while tweediei, known only from the holotype and paratype, is said to be uniform black above with a black and yellow chequered REPTILES AND AMPHIBIANS gI venter (Lim 1963). None of the jasoni specimens has the thin pale longitudinal line along the lowermost row of dorsal scales that is formed from a whitish spot on each scale and is characteristic of M. lateralis, nor the bright coral red venter of M. lateralis (Boulenger 1912). But the most remarkable distinguishing feature of jasont is its size and bulk at sexual maturity which are at least twice those of either Jateralis or tweediei. Gravid female lateralis vary in body length from 220 to 280 mm, whereas the gravid holotype of jasoni measures 700 mm from snout to vent. The female paratype of tweediet, FMNH 109868, which according to Inger (pers. comm.) seems to be sexually mature is stated by Lim to be 348 mm, but this measurement may include the broken tail. No differences in external meristic characters could be found between M. jasoni and M. lateralis although the ventral count of jasoni (131-134) is in the upper part of the wide range of variation noted in female /ateralis (116-136, N 16). Dental counts which were made on a number of British Museum and University of Malaya examples of M. lateralis, on the paratype of M. tweediei and on the three M. jasoni were found to be of value in demonstrating further differences between the three taxa. While the number of maxillary and palatine teeth in jasoni (M 11-13, P11) fall within the range of the counts for ten examples of Jateralis (M 10-14, Pg-12) the number of pterygoid teeth is non-overlapping. M. jasoni has 19, 20 and 21 in the three type specimens while 10 examples of Jateralis from which pterygoids were dissected out varied from 13-18 (mean 16) teeth, only the type having 18. Dr Inger, who kindly examined the paratype of tweediei reports that the right maxilla has 18 teeth (subequal), right palatine has 13 and the right pterygoid 23. In each of these dental counts tweediei is outside of the range of both Jateralis and jasom. An interesting comparison can be made by analysing these figures by the provenance of the individuals. The pterygoids of seven Jateralis obtained in the Larut Hills and Maxwell Hill vary from 14-16 while those lacking a locality have 16, 17 and 18, the highest count being in the type. The number of maxillary teeth in the type are also higher (14) than in the other lateralis that were examined (10-12) and this raises some doubt about the origin of the type specimen and of whether dental counts in Macrocalamus may vary geographically. No precise locality is available for the type of lateralis but Lim (1963) restricts it to Cameron Highlands. The series of Jateralis collected on the Cameron Highlands (Lim 1963 & 1967) has not been seen by me but I would venture to suggest that an examination of their pterygoid teeth and a comparison of counts with those made on Larut Hills material would reveal whether or not dental formulae in Macrocalamus are subject to geo- graphical variation, and if they are, whether the restriction of the type locality of lateralis to Cameron Highlands is appropriate. The higher number of pterygoid teeth in jasont presumably has some dietary significance although there is no evidence of jasoni having different feeding habits from lateralis. The stomach of the holotype of jasoni is empty. All three species of Macrocalamus occupy similar ecological niches. The jasoni series were obtained in a damp habitat among leaf litter or crossing the track. Lim (1963) records lateralis and tweediei from under logs in a damp environment at 6000 ft on G. Bringchang. The highest altitude from which M. lateralis has 92 A. G. C. GRANDISON been obtained is 6300 ft (Lim 1967). All other published records of the vertical distribution of this species indicate that it occurs between 3500 and 5700 ft (Bou- lenger 1912, Flower 1899, Smith 1922) in the Larut and Maxwell Hills, Perak, Cameron Highlands and G. Tahan, Pahang. M. tweediei has so far been found only on G. Brinchang, Cameron Highlands, at 5000 and 6000 ft (Lim 1963) where at the higher elevation it was in close association with M. lateralis. Lim (1967) described lateralis as being (with Pareas vertebralis) the commonest snakes on Gunong Brinchang and often found on the road basking in the early morning sun. Both Tweedie (1954) and Lim (1967) stated that Jateralis eats insects and their larvae. The specific name is appropriate in view of the striking golden ventral markings of the species; it is also cordially dedicated to the Hon. J. Jason Gathorne-Hardy, son of Lord and Lady Medway. Pareas vertebralis (Boulenger) Amblycephalus vertebralis Boulenger 1900 : 307. MATERIAL: BM 1967.2277 (1 9). REMARKS: This snake was found at night on a branch of a tree about 200 yards from camp 4 (5000 ft) on the trail leading to the water hole. The snake’s eyes reflected a bright red in torchlight. Scale rows . ; : é at] Ventrals . : : . 202 +1 Subcaudals ‘ : : 5, 66 Supralabials : : ’ : 7 (3-5) Loreal , : : F : iC Postoculars : : : ; I Temporals . , : ; : 2+2 Rance: W. Malaysia (mountains of Perak and Pahang). Pareas malaccanus (Peters) Asthenodipsas malaccana Peters 1864 : 273. MATERIAL: BM 10967.2276 (x 9). RemaRKS: Collected on the high bank at the side of the main track from Base camp, at about 800 ft. Ground colour black with irregular brownish grey cross bands bordered with white. Head silvery white. Scale rows . : : : te 5 Ventrals : z ; 5 . 158 +1 Subcaudals . : 5 Sy Rance: S. Thailand (Yala Prov.), W. Malaysia, Sumatra, Borneo (Taylor 1965). REPTILES AND AMPHIBIANS 93 Family ELAPIDAE Maticora intestinalis (Laurenti) Aspis intestinalis Laurenti 1768 : 106. MATERIAL: BM 1967.2280-81 (2 9). REMARKS: Both examples were found at night on steep slopes between 2000 and 3500 ft. When disturbed, the snake made no attempt to raise its tail and expose the vivid red undersurface, contrary to the normal behaviour of this species. In life the vertebral line from behind the frontal to the tail tip was red. Flanking the line were dark brown longitudinal bands which below were edged with a white line. The undersurface of the body had broad black and white bands of equal width. 2280 2281 Scale rows. : : 7 13 13 Ventrals : : é . 239 +1 244 +1 Subcaudals . F : 20 24 Supralabials : ; : 6 (3-4) 6 (3-4) Preoculars . : : F I I Postoculars . : j : 2 2 Temporals . : : é 1+2 I+2 RancE: Thailand, W. Malaysia, Singapore, Java, Borneo. Bungarus flaviceps (Reinhardt) Bungarus flaviceps Reinhardt 1843 : 267. MATERIAL: BM 1967.2286 (1 juvenile). Remarks: This specimen was found in an area between the disturbed forest around Base camp and the plantations near K. Damak. It is in poor condition. RANGE: Peninsular Burma, Thailand, S. Vietnam, W. Malaysia, Sumatra, Java, Borneo. Trimeresurus hageni (Lidth de Jeude) Bothrops hageni Lidth de Jeude 1886 : 54. MATERIAL: BM 1967.2290-91 (1G, I 9). Hapirat: One of the individuals (2291) was found in the late afternoon among ferns four to five inches from the forest floor by a narrow stream in the valley below camp 2, at about 1600 ft. The other example was obtained in the area of Base camp at approximately 700 ft. Remarks: Brongersma (1933) distinguished hageni from sumatranus on the basis of colour pattern, the number of upper labials in contact with the subocular and the juxtaposition of the supranasals. Both of the G. Benom individuals have the hageni pattern, that is the trunk scales are narrowly edged with black but have 94 A. G. C. GRANDISON no black crossbands, and a light lateral line along the first and second rows of dorsals and bordered below by a series of dark spots is present. A light streak (pink in life) from behind the eye to above the last supralabial is also present. Along the sides of the body from the neck to the vent is a series of light spots which in life are pinkish in colour. The spots continue on to the proximal half of the tail where they coalesce to form pinkish crossbands. The distal half of the tail is entirely pinkish in life. In head squamation however the two specimens differ, one agrees with Brongersma’s diagnosis of hageni in having contiguous supranasals and in two of the upper labials, the third and fourth, making contact with the subocular while the other (2290) has a small scale separating the supranasals and none of the upper labials is in contact with the subocular. The ventral and subcaudal counts for the female are 181 and 74, for the male 182 and 85. In the British Museum collections are two additional specimens that Dr A. R. Hoge has referred to T. hageni. One obtained by Malcolm Smith at Kuala Teku, Pahang (BM 1936.9.12.5) is a g and has 185 ventrals and 79 subcaudals. Its supranasals are in contact and on the right side of the head the third upper labial meets the subocular and on the left side the third and fourth. Its pattern closely resembles that of the G. Benom individuals. The other specimen identified as hageni (BM 8o.9.10.7), an adult 9, was collected in Singapore. Its counts are V185 C69 and it not only lacks the spots of the other examples but has indistinct black crossbanding on the trunk. Its supranasals are separated by a large internasal and the third supralabial (LHS) or the third and fourth (RHS) make contact with the subocular. RanGE: I believe these specimens constitute the first published record of the occurrence of T. hageni in Malaya. Brongersma (1933) examined examples of the species from Sumatra, Banka, Simalur, Batu Is., Nias and Mentawei Is. Trimeresurus popeorum Smith Trimeresurus popeorum Smith 1937 : 730. MATERIAL: BM 1967.2289 (1 9). Hasirat: Found during the day lying motionless at the side of the narrow trail above camp 3 at about 3700 ft. : COLOUR IN LIFE: Upper surface of body uniform green, venter paler green. Tail wine red, green below. Ventral 165 Subcaudals 62 Body length 680 mm Tail length 133-8 mm RANGE: E. Himalayas to Cambodia, W. Malaysia, Sumatra and Borneo. SNVIGIHd NV Caudacaecilia nigroflava Cacophryne borbonica Ansonia leptopus Rhacophorus macrotis Rana chalconota raniceps Rana hosii Rana 1. limnocharis Rana nicobariensis Rana miopus Rana plicatella Ranas. signata Rhacophorus appendiculatus Rhacophorus colletti Kalophrynus p. pleurostigma Microhyla berdmorei Microhyla butleri Microhyla heymonsi Phrynella pulchva Bufo asper Bufo parvus Pelophryne brevipes Megophrys monticola nasuta Amolops larutensis Rana laticeps Rana luctuosa Rana blythi Rana tweediei Rhacophorus 1. leucomystax Rhacophorus nigropalmatus Rhacophorus prominanus Rhacophorus reinwardtii Leptobrachium heteropus Chaperina fusca Ansonia ?tiomanica Megophrys aceras Philautus aurifasciatus Leptobrachium gracile Microhyla annectans Philautus vermiculatus Metaphrynella pollicaris REPTILES AND AMPHIBIANS 3 8 3 3 3 3 3 So So o So Oo So o o o o oO Oo So 1 4 4 4 —L —— i = - = = = = - . -= = - = = 95 yooy Ul opnzwyTy sueiqrydme wousg Suouny jo vornqiysrp [eurpnynty I alavy SGUVZIT SHMVNS 96 [ Cnemaspis kendalli Cyrtodactylus brevipalmatus Draco f. fimbriatus Draco f. formosus Draco maximus Draco melanopogon Draco punctatus Draco quinquefasciatus Draco v. volans Aphaniotis fuscus Goniocephalus borneensis Varanus bengalensis nebulosus Varanus salvator Mabuya multifasciata Cnemaspis affinis Cyrtodactylus consobrinus Cyrtodactylus marmoratus Gehyra mutilatus Scincella v. vittigerum Sphenomorphus 1. indicus Typhlops diardi mulleri Natrix trianguligera Macropisthodon rhodomelas Liopeltis longicauda Pseudorhabidon longiceps Calamaria lumbricoidea Lepturophis borneensis Pareas malaccanus Bungarus flaviceps Trimeresurus hageni Elaphe flavolineata Liopeltis baliodeirus Trimevesurus puniceus Gonyophis margaritatus Maticora intestinalis Trimeresurus popeorum Amphiesma savawacensis Paveas vertebralis | Macrocalamus jasoni A. G. C. GRANDISON uw 8 8 3 3 3 8 8s 8 8 8 8 s ++ + 4 4 se a a ——-—, —EEEEE__ Ld] = a a = —— i s s a a ———SSS a 1] s s s a s Ld] a a t — a a s a ——EE a a —— joo} UI epnzyTy sajiqde1 Wousg Suouny jo uorNqLysSIp [eUIpnynLy REPTILES AND AMPHIBIANS 97 Trimeresurus puniceus (Boie) Cophias punicea Boie 1827 : 561. MaTERIAL: BM 1967.2292-3 (I 9, 1 4). HasitaT: One specimen was found on a tree at 1700 ft, the other on a fallen tree between Base camp and camp 2. Ventrals 162 (g), 158 (9) Subcaudals 54 (8), 49 (9) RanGE: Peninsular Thailand, W. Malaysia, Sumatra, Mentawei Is., Java, Natuna Is., Borneo. ACKNOWLEDGEMENTS Tribute must be paid to the other members of the herpetology team, Félice V. Slade, Ben Ensol and Inche Sipang. Without Félice Slade’s enthusiasm, energy and keen eyesight the results of the team’s field work would undoubtedly have been very much poorer. The sagacious Ben Ensol guarded and guided ‘the ladies’ and provided not only much material but innumerable aids to camp comfort. The unobtrusive, conscientious Inche Sipang also made notable contributions in his own inimitable way. Members of the other teams and especially the porters, in particular the industrious Kam Mee Chow and the agile Berim bin Uda, also helped enormously both by collecting as time and opportunity allowed and by making expedition life so much more pleasant and profitable. I also wish to thank the following persons for the privilege of examining specimens in their care: Dr R. F. Inger and Mr H. Marx, Field Museum of Natural History, Chicago; Mrs P. Y. Berry, School of Biological Sciences, University of Malaya, Kuala Lumpur. This work has further benefitted from the advice of Dr G. L. Underwood, City of London Polytechnic, and from the assistance given by Mr A. F. Stimson who provided the scale counts on the snakes. I acknowledge my gratitude to them all. REFERENCES Bargour, T. 1938. Notes on ‘Nectophryne’. Pyoc. biol. Soc. Wash. 51 : 191-195, 1 text-fig. Berry, P. Y. & Henprickson, J. R. 1963. Leptobrachium nigvops, a new Pelobatid frog from the Malay Peninsula, with remarks on the genus Leptobyachium in Southeastern Asia. Copeia no. 4 : 643-648, 4 text-figs. Berry, P. Y. 1964. The breeding patterns of seven species of Singapore Anura. J. Anim. Ecol. 33 : 227-243, 5 text-figs. 1965. The diet of some Singapore Anura (Amphibia). Pyoc. zool. Soc. Lond. 144 : 163- 174, 1 text-fig. 1966. The food and feeding habits of the Torrent frog, Amolops lavutensis. J. Zool. Lond. 149 : 204-214, 3 text-figs. 98 A. G. C. GRANDISON Biytu, E. 1856. Proceedings of the Asiatic Society of Bengal, for October, 1855: report. J. Asiat. Soc. Beng. 24 : 711-725. Bore, F. 1827. Bemerkungen iiber Merrem’s Versuch eines Systems der Amphibien. tte Lieferung: Ophidier. Jsis Jena 20 : 508-566. Boutencer, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. London ed. 2, iv+503 pp. 29 pls. 1885. Catalogue of the lizards in the British Museum (Natural History). London ed. 2, vol. I, xii + 436 pp. 32 pls. 1887a. On new batrachians from Malacca. Ann. Mag. nat. Hist. (5) 19 : 345-348, 1 pl. 1887b. Catalogue of the lizards in the British Museum (Natural History). London ed. 2 vol. 3, vi+ 575 pp. 40 pls. 1890. List of the reptiles, batrachians, and freshwater fishes collected by Professor Moesch and Mr. Iversen in the district of Deli, Sumatra. Proc. zool. Soc. Lond. : 31-40. 1891. Descriptions of new oriental reptiles and batrachians. Ann. Mag. nat. Hist. (6) 7 : 279-283. 1893. Descriptions of new reptiles and batrachians obtained in Borneo by Mr. A. Everett and Mr. C. Hose. Proc. zool. Soc. Lond. : 522-528, 3 pls. —— 1894. A list of the reptiles and batrachians collected by Dr. E. Modigliani on Sereinu (Sipora), Mentawei Islands. Annali Mus. civ. Stor. nat. Giacomo Doria 14 : 613-618. 1895. Descriptions of four new batrachians discovered by Mr. Charles Hose in Borneo. Ann. Mag. nat. Hist. (6) 16 : 169-171. 1900a. Descriptions of new reptiles and batrachians from Borneo. Proc. zool. Soc. Lond. : 182-187, 4 pls. 1900b. Description of new batrachians and reptiles from the Larut Hills, Perak. Ann. Mag. nat. Hist. (7), 6 : 186-193. Igooc. Descriptions of new reptiles from Perak, Malay Peninsula. Ann. Mag. nat. Hist. (7) 5 : 306-308. 1903. Report on the batrachians and reptiles. Fasc. malayenses 1 : 131-176, 3 text-figs, 6 pls. 1908. Report on the Gunong Tahan expedition, May—September, 1905. III. Report on the fishes, batrachians and reptiles. J. fed. Malay St. Mus. 3 : 61-69, 2 pls. 1912. A Vertebrate Fauna of the Malay Peninsula from the Isthmus of Kva to Singapore including the adjacent Islands. London. xiii + 294 pp. 79 text-figs, 1 map. — 1918. Description of a new frog (Rana miopus) from Siam. J. nat. Hist. Soc. Siam 3; 11-12. 1g20a. Results of an expedition to Korinchi Peak, Sumatra. J. fed. Malay St. Mus. 8 : 82-285, 1 pl. 1920b. A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Rec. Indian Mus. 20 : 1-226. BroNGERSMA, L. D. 1933. 1.—Herpetological notes I-IX. Zool. Meded. Leiden 16 : 1-29. Buttock, J. A. 1966. Observations on the fauna of Pulau Tioman and Pulau Tulai. 7. The food of the amphibians and reptiles. Bull. natn. Mus. St. Singapore no. 34 : 85-96, tables 1-4. Butter, A. L. 1904. A list of the batrachians known to inhabit the Malay peninsula, with some remarks on their habits, distribution, etc. J. Bombay nat. Hist. Soc. 15 : 193-205 & 387-402. Cantor, T. 1847. Catalogue of reptiles inhabiting the Malayan Peninsula and Islands. Cal- cutta. 157 pp., 2 pls. Corsert, E.H. 1967. Adaptations for gliding in Dvaco. Am. Mus. Novit. no. 2283 : 1-20. Daupin, F. M. 1802. Histoive naturelle des vainettes, des grenouilles et des crapauds. Paris. 108 pp. 38 pls. Davis, D. D. 1965. Wallace’s flying frog. Malay. Nat. J. 19 : 149-151, 1 pl. FirzincEr, L. I. 1826. Neue Classification dey Reptilien nach ihren natiivlichen Verwandt- schaften. Vienna. viii + 66 pp. REPTILES AND AMPHIBIANS 99 FLowER, S. S. 1896. Notes on a collection of reptiles and batrachians made in the Malay peninsula in 1895-96; with a list of the species recorded from that region. Proc. zool. Soc. Lond. : 856-914, 3 pls. 1899a. 1. Notes on a second collection of reptiles made in the Malay Peninsula and Siam, from November 1896 to September 1898, with a list of the species recorded from those countries. Proc. zool. Soc. Lond. : 600-696. 1899b. Notes on a second collection of Batrachians made in the Malay Peninsula and Siam, from November 1896 to September 1898, with a list of the species recorded from those countries. Proc. zool. Soc. Lond. : 885-916, 2 pls. GRAVENHORST, J. L. C. 1829. Deliciae Musei Zoologici Vvatislaviensis. Reptilia Musei Zoologici Vratislaviensis vecensita et descripta... Fasciculus primus continens Chelonios et Batrachia. Lipsiae xiv-+106 pp. 17 pls. Gray, J. E. 1831. In Cuvier, edit. GrirFitH; The Animal Kingdom. Reptilia. A synopsis of the species of the class veptilia. London. 110 pp- 1845. Catalogue of the specimens of lizards in the collection of the British Museum. London. Xxvili + 289 pp. 1853. Descriptions of some undescribed species of reptiles collected by Dr. Joseph Hooker in the Khassia Mountains, East Bengal, and Sikkim Himalaya. Ann. Mag. nat. Hist. (2) 12 : 386-392. GtnTHer, A. 1858. Catalogue of the Batrachia Salientia in the collection of the British Museum. London. vi+160 pp., 12 pls. 1872. On the reptiles and amphibians of Borneo. Proc. zool. Soc. Lond. : 586-600, 5 text-figs, 6 pls. Haire, N. S. 1958. The snakes of Borneo, with a key to the species. Savawak Mus. J. 8 : 743-771, 2 pls, 2 text-figs. Hairston, N. G. 1957. Observations on the behaviour of Dyaco volans in the Philippines. Copeia no. 4 : 262-265, 1 pl. HaARDWICKE, Major-General & Gray, J. E. 1827. A synopsis of the species of saurian reptiles, collected in India by Major-General Hardwicke. Zool. J. Lond. 3 : 213-2209. HENDRICKSON, J. R. 1966. Observations on the Fauna of Pulau Tioman and Pulau Tulai. Bull. natn. Mus. St. Singapore no. 34 : 53-84, 2 text-figs, 14 pls. HEnnic, W. 1936. Revision der Gattung Draco (Agamidae). Temminckia 1:1 53-220, 11 text-figs, 14 maps. Incer, R. F. 1954. Systematics and zoogeography of Philippine amphibia. Fieldiana Zool. 33 : 183-531. 98 text-figs, 50 tables. 1960a. Notes on toads of the genus Pelophryne. Fieldiana Zool. 39 : 415-418, 1 text-fig. —— I960b. A review of the oriental toads of the genus Ansonia Stoliczka. Fieldiana Zool. 39, 473-503, II text-figs. —— 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana Zool. 52 : 1-402, 71 text-figs, 50 tables. 1969. Organization of communities of frogs along small rain forest streams in Sarawak. J. Anim. Ecol. 38 : 123-148. InceErR, R. F. & GREENBERG) B. 1966. Annual reproductive patterns of lizards from a Bornean rain forest. Ecology 47 : 1007-1021, 13 text-figs. Incer, R. F. & Marx, H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana Zool. 49 : 1-304, 73 text-figs. Kuni, H. 1820. Beitrdge zur Zoologie und vergleichenden Anatomie. Frankfurt am Main. vol. I, viii + 152 pp. Laurent, J.N. 1768. Austriaci Viennensis Specimen Medicum exhibens Synopsin reptilium Vienna. 214 pp., 4 pls. LiptH DE JEuDE, Tu. W. van. 1886. Note X. On Cophias wagleri, Boie and Coluber sumatranus, Raffles. Notes Leyden Mus. 8 : 43-54, 1 pl. 100 A. G. C. GRANDISON Lim, B. L. 1955. Snakes collected near Kuala Lumpur. Malay. Nat. J. 9 : 122-125. —— 1963. Macrocalamus tweediei, a new species of reed snake from Malaya. Bull. natn. Mus. St. Singapore no. 32 : 99-102, 1 text-fig., 2 pls. 1964. Comments on some rare snakes. Fed. Mus. J., 8 : 60-64. 1967. Snakes collected in Gunong Brinchang, Cameron Highlands, Pahang. Malay. Nat. J. 20 : 121-127. 1968. Further comments on rare snakes. Fed. Mus. J. 12 : 123-126. LinnaEus, C. 1758. Systema naturae per vegna tria naturae, secundum classes, ordines, genera, species cum characteribus differentiis, synonymis, locis. Stockholm. Ed. to, vol. 1, ii + 824 pp. Lioyp, M., IncEr, R. F. & Kine, F. W. 1968. On the diversity of reptile and amphibian species in a Bornean rain forest. Am. Nat. 102 : 497-515. MocguarD, F. 1892. Description de deux ophidiens et d’un batracien d’espéces nouvelles. Naturaliste (2) 6 : 35. Otiver, J. A. 1951. ‘Gliding’ in amphibians and reptiles, with a remark on an arboreal adaptation in the lizard, Anolis carolinensis carolinensis Voigt. Am. Nat. 85 : 171-176. ParkER, H. W. 1928. The Brevicipitid frogs of the genus Microhyla. Ann. Mag. nat. Hist. (10) 2: 473-499. 1934. A Monograph of the Frogs of the Family Microhylidae. British Museum London. vill + 208 pp. Peters, W. 1864a. Uber neue Amphibien (Typhloscincus, Typhlops, Asthenodipsas, Ogmodon). Mber. k. preuss Akad. Wiss : 271-276, 1 pl. 1864b. Uber einige neue Saugthiere ..., Amphibien... und Fische.... Mber. k. preuss Akad. Wiss : 381-399. 1867. Herpetologische Notizen. Mber. k. preuss. Akad. Wiss : 13-37. 1871. Uber neue Reptilien aus Ostafrica und Sarawak (Borneo), vorziiglich aus der Sammlung des Hrn. Marquis J. Doria zu Genua. Mber. k. preuss Akad. Wiss. : 566-581. REINHARDT, I. TH. 1843. Beskrivelse af nogle nye slangearter. K. dansk. Vidensk. Selsk. Skr. (4) 10 : 235-279, 3 pls. SCHLEGEL, H. 1837-1844. Abbildungen neuer oder unvollstindig bekannter Amphibien, nach dey Natur oder dem Leben entworfen, herausgegeben und mit einem erlduternden Texte begleitet von H. Schlegel. Diisseldorf. xiv + 141 pp, 50 pls. 1848. Descriptions de plusieurs espéces nouvelles du genre Lophyrus. Bijdy. Dierk. 1 : 4-6, 3 pls. 1858. Handleiding tot de beoefening dey Dierkunde. Breda. Vol. 2, xx+628 pp. SmitH, M. A. 1916. On a collection of reptiles and batrachians from Peninsular Siam. J. nat. Hist. Soc. Siam 2 : 148-171. 1917. On tadpoles from Siam. J. nat. Hist. Soc. Siam 2 : 261-278, 2 pls. 1922a. ‘The frogs allied to Rana doriae. J. nat. Hist. Soc. Siam 4 : 215-229, 1 text-fig., I pl. 1922b. Ona collection of reptiles and batrachians from the mountains of Pahang, Malay Peninsula. J. fed. Malay St. Mus. 10 : 263-282. 1923. Notes on reptiles and batrachians from Siam and Indo-China (No. 2). J. nat. Hist. Soc. Siam 6 : 47-53, I pl. — 1924a. New tree-frogs from Indo-China and the Malay Peninsula. Proc. zool. Soc. Lond. : 225-233, 3 pls. —— 1924b. VIII. Two lizards and a new tree frog from the Malay Peninsula. J. fed. Malay St. Mus. 11 : 183-186, 2 figs. — 1925. III. Onacollection of reptiles and amphibians from Mt. Murud, Borneo. Sarawak Mus. J.3 : 5-14, 1 pl. —— 1926. The functions of the ‘funnel’ mouth of the tadpoles of Megalophrys, with a note on M. acervas Boulenger. Proc. zool. Soc. Lond. : 983-988. —— 1930. The Reptilia and Amphibia of the Malay Peninsula. Bull. Raffles Mus. no. 3: I-149, 13 text-figs. REPTILES AND AMPHIBIANS 101 SmitH, M. A. 1935a. The fauna of British India including Ceylon and Burma. Reptilia and Amphibia. Vol. II—Sauria. London. xiii + 440 pp., 1 pl., 2 maps, text-figs. 1935b. Ona collection of reptiles and amphibians from Perak, Malay Peninsula. Bull. Raffles Mus. no. to : 61-63, 1 text-fig., 1 pl. 1937a. A review of the genus Lygosoma (Scincidae: Reptilia) and its allies. Rec. Indian Mus. 39 : 213-234, 5 text-figs. 1937b. Draco blandfordi and its allies. Bull. Raffles Mus. no. 13 : 75-76, 1 pl. 1937c. The names of two Indian vipers. J. Bombay nat. Hist. Soc. : 730-731. StoLiczKa, F. 1870. Observations on some Indian and Malayan Amphibia and reptiles. J. Asiat. Soc. Beng. 39 : 134-228, 3 pls. 1873. Notes on some species of Malayan Amphibia and Reptilia. J. Asiat. Soc. Beng. 42 : 111-126, I pl. Taytor, E. H. 1960. On the caecilian species Ichthyophis monochrous and Ichthyophis glutinosus and related species. Kans. Univ. Sci. Bull. 40 : 37-120, 38 text-figs. 1962. The amphibian fauna of Thailand. Kans. Univ. Sci. Bull. 43 : 265-599, 107 text- figs. 1963. The lizards of Thailand. Kans. Univ. Sci. Bull. 44 : 687-1077, 99 text-figs. — 1965. The serpents of Thailand and adjacent waters. Kans. Univ. Sci. Bull. 45 : 609- 1096, 125 text-figs. Tuomson, J. C. 1913. Contributions to the anatomy of Ophidia. Proc. zool. Soc. Lond. : 414-425, 2 text-figs. TscHupI, J. J. 1838. Classification dey Batrachier, mit Berucksichtigung der fossilen Thiere. Neuchatel. 100 pp., 6 pls. (Reissued in Mem. Soc. Sci. nat. Neuchatel 2 : 1-100, 6 pls. 1839 [1840].) Tweepiz, M. W. F. 1954. The Snakes of Malaya. Government Printing Office, Singapore. 139 pp., 12 pls., 27 text-figs. Van Kampen, P. N. 1922. The Amphibia of the Indo-Australian Archipelago. Leiden. xii + 304 pp. Wacter, J. 1830. Natiivliches System der Amphibien mit vorvangehender Classification dev Sdugthieve und Vogel. Miinchen, Stuttgart and Tiibingen. vi + 354 pp., 9 pls. WERNER, F. 1900. Reptilien und Batrachier aus Sumatra. Zool. Jb. Ab Syst. 13 : 32-508, 5 pls. WiecMann, A. F. A. 1835. In MEyYEN,F.J. F. Beitrage zur Zoologie gesammelt auf einer Reise um die Erde. Nova Acta physico-med.17 : 185-268. Wo tr, S. 1936. Revision der Untergattung Rhacophorus. Bull. Raffles Mus. no. 12 : 137- 217, 8 text-figs. Wray, L. 1890. Journal of a collecting expedition to the mountain of Batang Padang, Perak. J. Straits Brch. R. Asiat. Soc. no. 21 : 123-165. ' - 8 NOVi992 Miss A. G. C. GRANDISON, B.Sc. Department of Zoology British Museum (NaTuRAL History) CROMWELL Roap Lonpon SW7 5BD si, PRESENTED O% ah/ OGy We A LIST OF SUPPLEMENTS 2am TO THE ZOOLOGICAL SERIES ies OF THE BULLETIN OF ee THE BRITISH MUSEUM (NATURAL HISTORY) : Kay, E. Attson. Marine Molluscs in the Cuming Collection British Museum — (Natural History) described by William Harper Pease. Pp. 96; 14 Plates 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. ; Taytor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure and Mineralogy _ of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates - 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Guaises of the R.V. Antur ye 1962-1964. (In Press). iz Bc Printed in England by Staples Printers Limited at their Kettering, Northants, establishment bee at < 29 Re 1973 6000 Total Trap-nights 3380 1716 155 200 122 104 100 5777 Echinosovex gymnurus I Tupaia minor I Callosciurus notatus 6 I 5 Callosciurus caniceps Callosciurus nigrovittatus Sundasciurus tenuis I Sundasciurus lowit I Lariscus insignis 2 Rhinosciurus laticaudatus 4 Iomys horsfieldii 4 Rattus tiomanicus I — — od Rattus muelleri 6 Rattus bowersit 3 Rattus cremoriventer 3 Rattus surifer — — T Rattus vajah 4 I Rattus inas — Rattus whiteheadi 3 I I Rattus sabanus 32 Rattus edwardsi 2 4 2 Len! NOK HWW AHR HRW NHN HH OHH mW HN Total 77 13 4 4 5 4 Catch/1oo trap nights 23 0-76 26 2:0 4'1 3°8 Total species 16 5 4 3 2 2 2°0 20 ee ee | H H on 147 BIRDS AND MAMMALS ON GUNONG BENOM Jay}O IO syIeI} = 4 ‘ploosal jYSIS = s ‘pozoa]joo usutroeds = x :s[eArozUI 7F OO SurjUesarder sjoquiAs Aq wears ore oSuvI [eUIPN4][L $I JO SPU] PeArasqo ay} UTYZIM satoads Yowa OF sp1099yy WONHY ONONNA*S) NO SIVWWYW LSHXOd AO SGUOOTA IWNIGNLILTY b aTavy RS snaiuvanl snyjaysidig ‘20 3 vuaporp sosapisoddiy 61 * sngisaqvs soaapisodqiy gt 2s 4010919 sosapisodqiyy “LI *s snqoyofiay snydopouryay “9 t= <— === == BOSSI OOS BOOS AIOOC ES syodrvu snydojouryy St PASI III III ICIP EIS xxx suasnfaa snydojouryy *¥1 Mis Sie = = TeXsiss ae XS = = ~9X=X ouayss snydojourya ~€1 ES III IY, SFI IASI PD 9 swuyfy snydojouryy zi BSS = DIOINUOU DANUOYYQuUT “II xx SNUUM SNSSOPSOAIVDIA OI x svywpnogsa sdoaavsapy °6 opsayy sdojpyjapy Sep SOD eee Soto SSS SEIS ICO SCC SoBeSee x snyoydasounjau xouoay) °L Xececes===x vIDINIVU SiAaItuoyvg 9 x ispany Aojayjuag °S eX syodyavaq snaajqoudg + xx siywsarava snyvydasouky © s (é) vsoursynf vanpr904g Zz x \ snAnumaAs xadosoulyoy “1 ee ‘ON 4 € z T (34 000,) epnyaTy satoeds *poiloyur soueseid = - ‘Sunp o1OATUIvO UT [elI9}eUI = 4 ‘sodvI} LORD MEDWAY 148 KaXeXe wn XX Xa ee eg x x-xXXxX XeewaX-Xeon nm X-- - x-xXxXxX S--------------~--~~-~--sgg-----x X---------x x S$ ----------------- ~~ - eg eg --5 §-------------- --------s------ s----- ----s $---------- s x $--------------------------- -------------------- -s-------- s X---s----- xXsSX=----x--x x x 26 S------------X-------s X=-------x sinuay snanvsypung Sb snandqiy snanwsopung “bh SNADYLNOASIU SNANIOSOIIVI EY snuvminny snaniasoyvy “zh sdaavupa snanwsoyvg “1b ee jsonaag snaniasoywvy oF syMjou snAniasojjvy “6E swuify vfnoy “gt 4010919 vfnyoy LE vouvavl siuvpy «= E smdgoppuds sayoqojApyy SE Av] sayoqojdpy FE DULASAaMaU DIVIDE “EE Suunno.asvf{ vIvIV], “Tz soydojojau sydqsa4q “1€ Ss (3 000,) epnznTy vanasgo Sydqsadg “of Suvanos snqaayotn 67 UMO] SNIAAIOINT Bz dour vivdny ‘Lz sys viwdny, 9z vsoyidog vjnonuay Sz qwuoyny vuiunyl “hz syoj9h9 vuianyy Ez Yyojargos snyadjsidig ‘tz thalpia Snppaajsidig “1z ‘on setoads (‘pjuoo) F aqey Suoanurg syoyoap ol z x Suvsuy uopouoig ‘69 ------84------- ? SnUDADIDU SOJIADIAET “39 * snanosrnut snanaayyp °LO X--------- X-X-------X---xXX ispavmpa snywvy *99 Soest eteene at X-X--XxX-XXX : snungns snywy So X---------- ----xX-X s ipvayaziym snywy ‘'9 (o) Xa = =e Xooeesos= Saee=ux z, spur snyoy “£9 Be | ee Bi aD TS SSS a a a * * ae yoloa snyoy "ZO > a Aafians snywy “19 {e) XX Aajuanraowmadd SNYIDY O09 a 5 x dsaamog snyny “OS 3 xx ) F: ‘ ° 7 wad ‘ P>0-02), mean tail length (0-or>P>0-oo1), and mean hind foot length (PP>0:-2). TABLE I Bopy DIMENSIONS OF RATTUS RAJAH anv RATTUS SURIFER Head & Body Length Tail Length Mean + S.E Cot V Obs. Mean + S.E. C of V Obs. Species of Mean (%) Range of Mean (%) Range R. vajah 174°29 + 3°54 9°30 150-210 165°25 + 3°88 10°51 132-200 R. surifer 187-32 + 4°24 9°86 155-212 185-69 + 4°33 9°32 157-215 Hind Foot Length Ear Length Mean + S.E. Cof V Obs. Mean + S.E. C of V Obs. Species of Mean %) Range of Mean (%) Range R. vajah 37°86 + 0°42 5°03 34-40 21°95 40°71 4°66 20-23 R. suvifer 42°21 + 0-42 4°30 39-45 23°11 + 0°68 4°05 22-25 DR Hie Buys) Mean + S.E. C of V Obs. Species of Mean (%) Range R. vajah 94°91 + 1-30 6-13 82-11-105'81 R. suvifer 100-18 + 1°57 6:26 9Qf-51-112-12 S.E. = standard error C of V = coefficient of variation All measurements in millimetres MALAYAN RATTUS RAJAH AND RATTUS SURIFER 159 The pelage and tail colouration have been described in detail by Harrison (1957, 1966), Hill (1960) and Medway (1969). The two species are most readily distin- guished by the presence in R. surifer of a sharp demarcating stripe on the flanks, formed by orange-tipped spines, and by the general body colouration. SKULL ANATOMY. Various skull dimensions are given in Tables 2, 3 and 4. Of these only the mean length of anterior palatine foramina and the mean length of upper molar series between these rats differ significantly (P cs “ NAT. Hist. - § NOV1972 , e BH Printed in England by Staples Printers Limited at their Kettering, Northants, establishment ye? Jp 5 at aT at THE 3 GUNONG BENOM EXPEDITION : 1967 CKS (IXODIDAE) OF GUNONG BENOM ‘THEIR ALTITUDINAL DISTRIBUTION, STS AND MEDICAL RELATIONSHIPS ~ BULLETIN OF MUSEUM (NATURAL HISTORY) ore cs Vol. 23 No. 7 ~ LONDON: 1972 THE GUNONG BENOM EXPEDITION 1967 8. TICKS (IXODIDAE) OF GUNONG BENOM AND THEIR ALTITUDINAL DISTRIBUTION, HOSTS AND MEDICAL RELATIONSHIPS BY HARRY HOOGSTRAAL U.S. Naval Medical Research Unit, Cairo. LIM BOO LIAT Institute for Medical Research, Kuala Lumpur. M. NADCHATRAM Institute for Medical Research, Kuala Lumpur. GEORGE ANASTOS University of Maryland. Pp. 165-186 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 7 LONDON: 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), ‘instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 23, No. 7 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull, Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 4 October, 1972 Price 75p THE GUNONG BENOM EXPEDITION 1967 8. TICKS (IXODIDAE) OF GUNONG BENOM AND THEIR ALTITUDINAL DISTRIBUTION, HOSTS AND MEDICAL RELATIONSHIPS By H. HOOGSTRAAL, B. L. LIM, M. NADCHATRAM & G, ANASTOS SYNOPSIS Ticks collected on Gunong (Mt.) Benom, an undisturbed forest environment particularly favourable for studying altitudinal distribution (700 to 6916 ft) of tick species occurring in the Malay Peninsula, were Ixodes granulatus, I. simplex, Haemaphysalis caluus, H. bispinosa, H. c. cornigera, H. hylobatis, H. nadchatrami, H. semermis, H. traguli, H. vidua, H. wellingtoni, H. asiatica, H. doenitzi, H. koningsbergeri, Dermacentor atrosignatus, D. compactus, Amblyomma testudinarium, and A. geomeydae. T.simplexand H.calvus were not previously recorded from the Malay Peninsula. Haemaphysalis centropi Kohls, 1949, is a synonym of the earlier described H. doenitzi Warburton and Nuttall, 1909. The species of Deymacentor in Malaya are reported for the first time. No ticks were collected in mossy forests (5300-5500 ft). One found on a person at 6000 ft may have attached at a lower level. Deymacentor atrosignatus comprised 48% of the collections from vegetation between 500 and 2500 ft altitudes and 41% of those from 2600 to 5200 ft. D. compactus comprised 16% of collections from vegetation up to 2500 ft, but only 0:2% of those from higher altitudes. H. nadchatvami and H. semermis comprised 6 and 11%, respect- ively, of collections from vegetation between 500 and 2500 ft but 54% and 2% respectively, of those from 2600 to 5200 ft. Data are provided for tick parasitism of 28 mammal and bird species on Gunong Benom, for samples obtained by sweeping forest vegetation at different altitudes, and for vertebrates that were not tick-infested. Species biting human beings were H. nadchatrami, H. semermis, H. c. corvnigera, Dermacentor atrosignatus, and A. testudinarium. This mountain area is eminently suitable for investigating “primitive’’ cycles of tickborne viruses and rickettsiae in nature and many of the tick species listed are important candidates for epidemiological research. Elsewhere in Malaya, I. granulatus has been found to be infected by Langat and Lanjan viruses, Devmacentoy sp. and H. semermis by Lanjan virus, and H. nadchatrami by Q fever (Coxiella burneti). Both haemaphy- salid species, Deymacentoy sp. and I. granulatus have been implicated in the natural cycle of tick typhus (Rickettsia sp.) in Malaya. INTRODUCTION Mount (Gunong) BENom offers an exceptional opportunity for investigating the effects of altitude on ecological zonation in Malaya (Whitmore, 1972). The highest peak (6916 ft) in a compact montane formation, Gunong Benom is situated at approximately 4°N in the middle of Malaya between the Main and Eastern Ranges. Forest extending from 700 ft to the peak has been disturbed by logging operations only near the base and in scattered patches up to about 1700 ft. Ecologically, lowland forest merges with montane forest at 2500 ft and montane mossy forest begins at 5300 ft. 168 H. HOOGSTRAAL AND OTHERS During the current Malayan Tick Survey!, we have obtained considerable data concerning the distribution and host relationships of the ticks inhabitating low- lands and highlands of the Malay Peninsula. However, prior to the Gunong Benom Expedition (Medway, 1972), we had not had the opportunity to compare systemati- cally the effects on tick distribution and incidence of altitudinal factors in a single unbroken, undisturbed Malayan transect such as this. Results from the Gunong TABLE I Ticks from vegetation on Gunong Benom, Pahang Tick species Number 3 i] Nymph Larva Total (%) 500-2500 Feet altitude (Lowland forests) Dermacentor atrosignatus 103 158 fo) fo) 261 (47°8) Dermacentor compactus 40 40 fo) (0) 86 (15°8) Haemaphysalis nadchatrami 19 12 ° ° 31 (5°7) Haemaphysalis semeyrmis 25 37 fo) fe) 62 (11-3) Haemaphysalis koningsbergeri 3 3 ° fo) 6 (1-1) Haemaphysalis hylobatis I 3 fo) ° 4 (0-7) Haemaphysalis caluvus 16 18 fo) fo) 34 (6-2) Haemaphysalis c. cornigera 18 21 3 fo) 42 (7:7) Amblyomma testudinarium 7 5 2 fo) 14 (2-6) Amblyomma geoemydae fo) fo) 6 fo) 6 (1-1) Total 232 303 1G fo) 546 2600-5200 Feet altitude (mountain forests) Dermacentor atrosignatus 63 109 to) fo) 172 (41-4) Dermacentor compactus* ° I fo) fo) I (0-2) Haemaphysalis nadchatrami 110 113 fo) fe) 223 (53:6) Haemaphysalis semermis 3 7 fo) ° 10 (2-4) Haemaphysalis hylobatis fo) I fo) fo) I (0:2) Amblyomma testudinarium 5 4 I ° 9 (2-2) Total 181 234 I to) 416 5300—5500 Feet altitude (Mountain mossy forests) (No ticks) ** fo) to) fo) fo) fo) (0-00) *Collected at 2700 ft. **5 persons sweeping 24 March (1500-1800 hrs), 25 March (1000-1500 hrs), 26 March (1100-1400 hrs), 14 January (1400-1800 hrs), 15 January (0900-1300 hrs). 1The Malayan Tick Survey, which has been in operation since mid-1966, is a collaborative study between the Institute for Medical Research, Kuala Lumpur, the Zoology Department, University of Maryland, and NAMRU-3, Cairo, supported in part by U.S. Army Research and Development Command grant number DADA-17-67-C-7003. TICKS (IXODIDAE) OF GUNONG BENOM 169 Benom study have been particularly useful in providing material and data for the first author’s revisional study of the Dermacentor ticks of Southeast Asia and of the biology and distribution of Haemaphysalis ticks of the world.? The University of Malaya— British Museum (Natural History) — sponsored Gunong Benom Expedition, headed by Lord Medway, who has described (1972) previous exploration of the area, expedition aims and operations, and participating organizations and personnel, was in the field from 1 February to 15 April 1967. Members of the George W. Hooper Foundation of the University of California Medical Center, San Francisco, then attached to the Faculty of Medicine, University of Malaya, participated in the expedition and later revisited the mountain for additional collecting (Marchette, 1967; Dr. Richard Garcia, personal communication). Certain members of the Malayan Tick Survey also collaborated with the expedition and worked in the area prior to and following the period the expedition was on Gunong Benom. The names of persons responsible for the excellent tick collections and data reported here are listed at the end of this paper. The outline description of forest zones on northeast Gunong Benom by Whitmore (1972) should be consulted by readers who are particularly concerned with the ecological aspects of this study. The term Malaya, as used in this paper, refers only to the Malay Peninsula from the Southern border of Thailand to Singapore. In discussions of Borneo, political divisions are not mentioned. MATERIALS AND METHODS Vertebrate animals collected by members of the Gunong Benom expedition or the Malayan Tick Survey were routinely inspected for ectoparasites. For ticks, bath towels attached to a stick were also used to flag vegetation at intervals from 700 to 5500 ft. Owing to the absence of ticks between 5300 and 5500 ft, and the cold, wet weather above this altitude, no effort was made to collect material between 5500 ft and the peak at 6916 ft. Actual dates of collection of each specimen are available and will be furnished on request. RESULTS The ticks of Malaya have been reviewed by Kohls (1957) and by Audy et al. (1960). Subsequent changes in the taxonomic status of certain Haemaphysalis species are mentioned below. The reader is referred to these papers for taxonomic details, which are omitted in this report. Results are presented in tabular form for ticks taken by flagging vegetation at various altitudes (Table 1) and for those from hosts (Table 2), together with a list of vertebrate species that were apparently not tick- infested. 2The Dermacentor and Haemaphysalis studies are supported in part by contract number 03-005-o1 between the U.S. Public Health Service National Institute of Allergy and Infectious Diseases, Bethesda, Maryland, and NAMRU-3, Cairo. 170 H. HOOGSTRAAL AND OTHERS TABLE 2 Ticks from vertebrates on Gunong Benom, Pahang* 500-2500 Feet altitude (Lowland forests) Vertebrate species INSECTIVORA Echinosorex gymnurus CHIROPTERA Penthetor lucast Rhinolophus stheno Rhinolophus affinis RopDENTIA Ratufa affinis Callosciurus caniceps Sundasciurus tenuis Rhinosciurus laticaudatus Lariscus insignis Rattus sabanus Rattus muelleri Rattus bowersit Rattus vajah Rodent burrow CARNIVORA Felis bengalensis Arctiogalidia trivirgata Arctictis binturong Hemigalus derbyanus Pavadoxurus hermaphroditus Domestic dog (Lord Medway) Domestic dogs (aborigine village) Number examined nN 40 15 Number infested H Tick species (sex/stage) Ixodes granulatus (42, 3N, 4L) Amblyomma sp. (1L) Ixodes simplex (1L) Ixodes simplex (62, 2N, 5L) Ixodes simplex (39, 2N, 4L) Dermacentor sp. (1N) Dermacentor atrosignatus (2N) Haemaphysalis nadchatrami (2N) Haemaphysalis nadchatrami (11N) Haemaphysalis ?koningsbergert (2N) Ixodes granulatus (19) Haemaphysalis nadchatrami (3N, 25L) Dermacentor atrosignatus (4N) Dermacentor sp. (2N, 29L) Ixodes granulatus (1g, 19) Amblyomma sp. (1L) Dermacentor atrosignatus (1N) Dermacentor sp. (16L) Haemaphysalis nadchatvami (1N) Haemaphysalis semermis (3N, 1L) Dermacentor sp. (1L) Haemaphysalis nadchatrami (1N) Ixodes granulatus (12L) Haemaphysalis asiatica (13) Haemaphysalis koningsbergeri (143) Haemaphysalis semermis (13) Haemaphysalis koningsbergeri (23) Haemaphysalis koningsbergeri (263, 29) Haemaphysalis semermis (13) Haemaphysalis hylobatis (23) Haemaphysalis koningsbergeri (13) Haemaophysalis vidua (3N, 26L) Haemaphysalis honingsbergeri (19) Dermacentor sp. (4L) Haemaphysalis koninigsbergeri (73, 89) | Haemaphysalis semermis (33) : Haemaphysalis bispinosa (83, 52, 4N) | Haemaphysalis koningsbergeri (33, 99) | TICKS (IXODIDAE) OF GUNONG BENOM 171 Vertebrate species Number Number examined infested ARTIODACTYLA Tragulus javanicus 4 4 Tragulus napu I I Sus scrofa 2 2 AVES Centropus sinensis eurycercus 2 2 Dendrocopos m. moluccensis Argusianus argus Lophura evythrophthalma Anthvacoceros malayanus HUMAN BEINGS (biting) ** Ln | Lo HUMAN BEINGS (crawling on) 2600-6000 Feet altitude (Mountain RODENTIA Rattus inas (3600 ft) 4 I HuMAN BEING (biting, 6000 ft)** HUMAN BEING (crawling on, 3000 ft) Tick species (sex/stage) Dermacentor sp. (IN) Haemaphysalis nadchatrami (4N) Haemaphysalis semermis (1g, 2N, 1L) Haemaphysalis traguli (13, 29, 5N, 5L) Haemaphysalis ?nadchatrami (1L) Amblyomma testudinarium (iN) Dermacentor atrosignatus (273, 92) Dermacentor compactus (113, 5) Haemaphysalis nadchatrami (183, 52) Haemaphysalis semermis (23, 19) Amblyomma sp. (IN) Amblyomma geomeydae (1N) Haemaphysalis doenitzi (13) Haemaphysalis doenitzi (2L) Haemaphysalis wellingtoni (2N, 1L) Haemaphysalis wellingtoni (23, 10L) Amblyomma sp. (1N) Amblyomma testudinarium (1N) Dermacentor atrosignatus (1N) Haemaphysalis c. cornigeva (19) Haemaphysalis nadchatvamt (29) Haemaphysalis semermis (1g, 2N) Amblyomma testudinarium (1g, 22) Dermacentor atrosignatus (13, IN) Haemaphysalis nadchatrvami (29) Haemaphysalis semermis (22) forests) Haemaphysalis nadchatrami (1N) Haemaphysalis nadchatrami (19) Haemaphysalis hylobatis (19) *The 243 vertebrates (representing 53 forms) inspected and not found to be tick infested, and their numbers (in parentheses), were: 500-2500 Feet altitude (Lowland forests) Primates. Tupaia minor (1), T. glis (2), Ptilocercus lowii (2), Nycticebus coucang (1), Macaca fascincularis (1), Presbytis obscurus (5). CHIROPTERA. Cynopterus brachyotis (11), C. horsfieldi (22), Eonycteris spelaea (18), Chivonax melanocephalus (1), Macroglossus lagochilus (6), Balionycteris maculata (3), Rhinolophus macrotus (2), R. vefulgens (12), Emballonuva monticola (12), Hipposideros diadema (2), H. galevitus (2). RopeEnt1a. Hylopetes spadiceus (4), Pteromyscus pulverulentus (1), Petaurista petaurista (11), P. elegans (4), Aeromys tephromelas (4), Iomys horsfieldi (5), Sundasciurus lowi (4), S. tenuis (8), S. hippurus (2), Callosciurus notatus (14), C. nigrovittatus (4), C. caniceps (1), C. prevosti (1), Rhinosciurus laticaudatus (2), Lariscus insignis (1), Ratufa bicolor (6), R. affinis (6), Rattus cremoriventer (7), R. sabanus (15), R. bowersit (1), R. vajah (6), R. inas (3), R. canus (2), R. whiteheadi (6), Chivopodomys gliroides (1). Reprizia. Varanus salvator (1), V. rudicollis (1), Gonocephalus boreensis (1). Aves. Otus rufescens (1), Chrysocolaptes validus (1), Avachnothera longirostra (1), Anorrhinus galeritus carinatus (1). 2600-5300 Feet altitude (Mountain forests) RopEnt1A. Sundasciurus tenuis tahan (3), Callosciurus flavimanus (1), Rattus edwardsi (7), R. tiomanicus jalorensis (3). ** Also see remarks in text on larvae (presumably 4. testudinarium), infesting persons at 3000 to 4000 foot altitudes during the wet season but not during the dry month of February. 172 H. HOOGSTRAAL AND OTHERS Ixodes granulatus Supino I. granulatus was collected at 700 and 800 ft from the single moonrat (Insectivora, Erinaceidae, Echinosorex gymnurus) and from 2 rodents, Rattus sabanus and R.muellevi. The moonrat, trappedon28 February beside astream, was parasitized by 4 female I. granulatus on the dorsum and 3 nymphs and 4 larvae on the thighs and belly. One male and 1 female infested 1 of 5 R. muelleri (3 March) and 1 female infested x of 25 R. sabanus (12 January). Twelve larvae resting on the roof of a rodent burrow dropped to a black plate (used for collecting chiggers) a few moments after the plate was inserted by Nadchatram into 1 of 40 burrows on Gunong Benom, Notably, the infestation rate of this tick species on the mountain at the times of collection was low; squirrels and several other rodents were not infested. I. granulatus commonly parasitizes small mammals, and sometimes birds, living on the ground or in trees in forests from Southeast Asia (southern China, Vietnam, Okinawa, Borneo, Indonesia) to eastern India (Assam). The life cycle of this 3-host tick under laboratory conditions was described by Nadchatram (1960). This is the only Malayan tick species in which each developmental stage parasitizes rodents. The nidicolous behaviour of J. gvanulatus has been confirmed by Nadchatram in several areas of Malaya (unpublished data.) I. granulatus from Rattus muelleri and R. sabanus in the Ulu Langat Forest Reserve, Selangor, was the original source of the virus causing Langat encephalitis of humans (Smith, 1956; Hoogstraal, 1966). The natural history of Langat virus remains to be studied. J. granulatus is also involved in the cycles of tick typhus (Rickettsia sp.) and Q fever (Coxiella burnett) in climax forests of Malaya (Marchette, 1966). Specimens from Trengganu have yielded Lanjan virus (N. J. Marchette, personal communication). Only a single specimen of J. granulatus has been recorded biting man in Malaya. Thus, this species plays no important direct role in the epi- demiology of human disease. However, many other ticks, among them species that more or less frequently infest human beings, feed on the same hosts and may acquire pathogens circulated in a granulatus — rodent — granulatus cycle and transmit these disease agents to other hosts during feeding in subsequent developmental stages. The potential indirect epidemiological role of J. granulatus should be espec- ially interesting to investigate. Ixodes simplex Neumann Bats parasitized by I. simplex were Penthetor lucasi (1 larva from 1 examined, 3600 ft), Rhinolophus stheno (6 females, 2 nymphs, 5 larvae, from 11 of 24 examined between 700 and goo ft), and R. affinis (3 females, 2 nymphs, 4 larvae, from 6 of 6 examined at 1500 ft). Eleven other bat species (gt specimens) between 500 and 2500 ft were free of ticks. These are the first published records of J. simplex from Malaya and from the genus Penthetor. I. simplex infests many microchiropteran bats, chiefly in the genera Rhinolophus, Myotis, and Miniopterus,in Africa, Europe, Asia and Australia. Other (unpublished) TICKS (IXODIDAE) OF GUNONG BENOM 173 Malayan records are from R. stheno and Miniopterus medius at Karak and Fraser’s Hill, Pahang. The disease relationships of J. simplex have not been investigated. Haemaphysalis calvus Nuttall and Warburton Sixteen male and 18 female H. caluvus were found on lowland forest vegetation in Kuala Kerau and Kuala Lompat Forest Reserves. This species is known only from the original collection from a buffalo in northern Borneo, and from our collections (unpublished) from forest vegetation in Kedah, Pahang, and Selangor, from a tame sambar deer (Cervus unicolor) and a tiger [British Museum (Natural History)j in Pahang, and from a bear near Bangkok, Thailand. Disease relationships have not been studied. Haemaphysalis bispinosa Neumann Eight males, 5 females, and 4 nymphs were taken from 6 of 15 domestic dogs examined in Kuala Kerau Forest Reserve in September 1966. These dogs belonged to aborigine villagers and should be recognized as a source distinct from Lord Medway’s pet dog, which accompanied Gunong Benom Expedition members and yielded ticks characteristic of wild forest-dwelling carnivores. In Malaya, H. bispinosa shows all the biological features of an introduced parasite, being associated entirely with domestic animals (Hoogstraal et al., 1969). In its homeland, Ceylon, India, southern Nepal, and western Burma, H. bispinosa com- monly parasitizes wild and domestic birds and mammals. The long list of disease relationships attributed to this species in much literature from Asia, Australia, New Zealand, Japan, and Pacific islands is completely erroneous, having been assembled from studies of misidentified ticks (Hoogstraal et al., 1968). The true disease relationships of H. bispinosa remain to be determined. Haemaphysalis cornigera cornigera Neumann Specimens collected in Kuala Kerau and Kuala Lompat Forest Reserves at the southeast base of Gunong Benom by the Malayan Tick Survey were: (September 1966) 16 males, 13 females, 3 nymphs from forest vegetation, 1 female, biting person; (April 1968) ; 2 males, 8 females from forest vegetation. H. c. cornigera is widely but apparently erratically distributed in Malayan forests, where adults parasitize various kinds of deer, and wild pigs and carnivores. Nymphs and larvae feed on forest rodents. This parasite is common in Borneo and Indonesian islands. Its northern distributional limits and disease relationships remain to be investigated. 174 H. HOOGSTRAAL AND OTHERS Haemaphysalis hylobatis Schulze Eight specimens were taken, 2 males from the bear cat, Arctictis binturong, at 700 ft, 1 female crawling on a man at 3000 ft, 1 male and 3 females from lowland forest vegetation, and 1 female from mountain forest vegetation (2700 ft). H. hylobatis is an infrequently encountered tick of Malaya and Sumatra, and the 8 from Gunong Benom comprise the largest known number from any single locality. This deep-forest parasite, infests monkeys, man, and carnivores, including domestic dogs, in Pahang, Selangor, and Kedah, Malaya, and in Sumatra, Indonesia. A female reared from a nymph infesting Rattus surifer in Selangor (Audy et al., 1960) is the only previously published record of this species in Malaya, and the only available data for immature stage hosts. The disease relationships of H. hylobatis have not been studied. Haemaphysalis nadchatrami Hoogstraal, Trapido and Kohls Collections from sweeping forest vegetation and from hosts (examined/infested) at various altitudes are as follows: Rattus sabanus (40/5), 700 to 950 ft, 3 nymphs, 25 larvae Rattus bowersi (2/1), 800 ft, I nymph Rattus vajah (7/1), 1700 ft, 1 nymph Rattus inas (4/1), 3600 ft, 1 nymph Sundasciurus tenuis (9/1)., lowlands, 2 nymphs Rhinosciurus laticaudatus (3/1), 800 ft, 1x nymphs Sus scrofa, (2/2) 800 and 1400 ft, 183, 52 Tragulus napu (1/1), lowlands, 1 larva (tentative identification) Tragulus javanicus (4/2), lowlands, 4 nymphs Domestic dogs (aborigine village) (15/2), lowlands, 1g, 12 Man (biting), 750 and 6000 ft, 29 Man (crawling on), 750 ft, 19 Vegetation, 500 to 2500 ft, 234, 22° Vegetation, 2600 to 5200 ft, 110g, 1139 H. nadchatrami larvae and nymphs feed on forest rodents and other small mammals Usually rather few are collected from hosts, presumably owing to especially short periods of parasitism. Adults feed on wild pigs, deer, carnivores, and man. As usual, the 2 wild pigs (Sws scrofa) examined were infested by a number of H. nad- chatrami adults. Sweeping forest vegetation in all parts of Malaya almost always yields questing adults of this species. Notably, adults were much more numerous on vegetation on Gunong Benom at 2600 ft and above than below this elevation (Table 1). All samples, particularly those from higher elevations, were at the upper range of robustness for this species. The Gunong Benom forest environment is obviously ideal for H. nadchatrami. The comparative incidence in Malaya of this species and of H. semermis is discussed in the section on the latter species. ———————— ee eee ee a a TICKS (IXODIDAE) OF GUNONG BENOM 175 One of the 2 females biting humans was attached to the ankle, under the sock; a large patch of host skin remains surrounding the hypostome. The second dis- covered on a person near camp 5 at 6000 ft, may have attached at a lower altitude. H. nadchatrami, a common forest dweller from central Thailand through the Malay Peninsula to Borneo, Java, Sumatra, and adjacent islands of Indonesia (Hoogstraal, Trapido, and Kohls, 1965), also occurs in the Phillipines on Palawan Island near Borneo (Hoogstraal, Kohls, and Parrish, 1968). In Malaya, adults have been found the year around. While there is probably a seasonal peak in population numbers, this has not yet been demonstrated. Owing to its ubiquity, wide range of immature and adult stage hosts including man, and year-round feeding activity, H. nadchatrami is highly suspect as a reservoir and vector of viruses and rickettsiae. The ‘‘Haemaphysalis spp.’’ in Marchette’s (1966) study of Malayan tick typhus obviously consisted of many H. nadchatrami (Hoog- straal, 1967). Specimens of this species from Trengganu have yielded Coxiella burnett, the agent of Q fever (N. J. Marchette, personal communication). During the Malayan Tick Survey, 7 males and 4 females, in addition to those listed here, were collected while biting man. Haemaphysalis semermis Neumann Collections from sweeping forest vegetation and from hosts (examined/infested) at various altitudes are as follows: Rattus bowersit (2/2), 800 ft, 3 nymphs, 1 larva Felis bengalensis (1/1), 800 ft, 12 Arctictis binturong (1/1), 700 ft, 1g Domestic dog (Lord Medway’s), 700 ft, 3g Domestic dogs (aborigine village) (15/6), lowlands, 8g, 119, 5 nymphs Sus scrofa (2/2), 800 and 1400 ft, 2g, 19 Tragulus javanicus (4/1), lowlands, 1g, 2 nymphs, 1 larva Man (biting), 800 ft, 14, 2 nymphs Man (crawling on), 700 and 1000 ft, 39 Vegetation, 500 to 2500 ft, 47g, 532 Vegetation, 2600 to 5200 ft, 3g, 72 The host range of H. semermis and H. nadchatrami is the same, though preferred hosts appear to be carnivores for semermis and wild pigs for nadchatrami. The 2 species are practically always sympatric in Malaya, and often on the same hosts. Of the 2259 adults of these 2 species in our Malayan collections, 1589 (70°) are nadchatrami and 670 (30%) are semeymis. Of the 427 adults of these 2 species from Gunong Benom, 286 (67%) are nadchatrami and 141 (33%) are semermis. The distribution of H. semermis, from southern Thailand through the Malay Peninsula and adjacent islands to Borneo and Sumatra, is almost the same as that of H. nadchatrami (Hoogstraal, Trapido, and Kohls, 1965). 176 H. HOOGSTRAAL AND OTHERS The remarks on potential medical relationships of H. nadchatrami apply equally to H. semermis. Specimens ot H. semermis from Trengganu have yielded Lanjan virus (N. J. Marchette, personal communication). Haemaphysalis traguli Oudemans The 4 larger mouse-deer, Tvagulus javanicus, examined in the lowland forests of Gunong Benom were infested by 1 male, 2 females, 5 nymphs, and 5 larvae of H. traguli. (The single smaller mouse-deer, T. napu, was parasitized by only a larva of H. ?nadchatrami.) Adults and immature stages of H. traguli commonly feed on Tvagulus javanicus and 7. napu in forests of Malaya, and also in Indonesia, Borneo, Thailand, and Burma (Hoogstraal, 1966). The subgeneric position of this and the related H. vidua is uncertain. The disease relationships of H. traguli have not been investigated. Haemaphysalis vidua Warburton and Nuttall Three nymphs and 26 larvae of H. vidwa were taken on a banded palm civet, Henugalus derbyanus, at goo ft. This rare tick is known only from various species of civets (Viverridae) in Pahang, Malaya, and near Kalabakan, Borneo (Hoogstraal, 1962, 1964). Disease relation- ships have not been studied. Haemaphysalis wellingtoni Nuttall and Warburton Two nymphs and 1 larva were taken from an argus pheasant, Argusianus argus, in lowland Kuala Lompat Forest Reserve on 12 March 1966. Two males and 10 larvae were found among the ear coverts of a crestless fire-back pheasant, Lophura erythropthhalma, in the same locality the same week. H. wellingtoni ranges from Ceylon, Andaman Islands, and India through southern Nepal, Burma, Thailand, Vietnam, and Malaya, to Borneo and Indonesia (reports from New Guinea are incorrect). All stages typically parasitize wild and domestic gallinaceous birds. Other ground-feeding birds, and sometimes small mammals, are less frequently infested. The migratory habits of some hosts undoubtedly account for the extraordinarily wide distribution of H. wellingtoni and specimens have been found in Japan on a migrant gray thrush, Twrdus c. cardts. The disease relationships of H. wellingtont unfortunately have not been studied This species is an especially interesting candidate for arbovirus investigation. TICKS (IXODIDAE) OF GUNONG BENOM 177 Haemaphysalis asiatica (Supino) A single male of this uncommon species was taken on a leopard cat, Felis bengalen- sis, at 750 ft. H. asiatica [= H. dentipalpis Warburton and Nuttall] occurs in forests of Burma, southern China, Vietnam, Thailand, peninsular Malaya, and Borneo, where its chief hosts are civets (Viverridae) and, less often, other carnivores (Hoogstraal and Trapido, 1966). The disease relationships of this species have not been studied. Haemaphysalis doenitzi Warburton and Nuttall Two of the 9 birds examined on Gunong Benom were infested by H. doenitzt. These were a coucal, Centropus sinesis, by a male (7oo ft), and a short-tailed babbler, Trichastoma malaccense, by 2 larvae (1500 ft). H. doenitzi adults and immature stages infest birds in forests from Ceylon and southern India to Borneo, Indonesia, New Guinea, southern China, Laos, Vietman, Taiwan, and southern Japan (Hoogstraal, unpublished data). Coucals, gallin- aceous birds, and other birds that feed on or close to the ground are frequent hosts. Some records from Thailand and Laos are from mammals (leporids and domestic dogs.) H. centropi Kohls (1949) is a synonym of H. doenitzi Warburton and Nuttall (r909) (Hoogstraal, unpublished). The disease relationships of this widely-ranging species unfortunately have not been investigated. Haemaphysalis koningsbergeri Warburton and Nuttall Collections from sweeping forest vegetation and from hosts (examined/infested) are as follows: Lariscus insignis (3/2), 700 and 800 ft, 2 nymphs (tentative identification) Felis bengalensis (1/1); 800 ft, 144 Arctiogalidia trivirgata (1/1), 750 ft, 2g Arctictis binturong (1/1), 700 ft, 263, 22 Hemigalus derbyanus (1/1), goo ft, 1g Paradoxurus hermaphroditus (1/1), lowlands, 12 Domestic dog (Lord Medway’s), 750 and 800 ft, 73, 89 Domestic dogs (aborigine village) (15/6), lowlands, 33, 99 Vegetation, 500 to 2500 ft, 3g, 32 Adults of H. koningsbergeri are common parasites of all wild carnivores and of domestic dogs in Malayan forests. Exceptional hosts (unpublished) are the mouse- deer (Tvagulus javanicus and T. napu), wild pig (Sus scrofa), pangolin (Manis javanica), and domestic goat. A single record, possibly erroneous, is from domestic 178 H. HOOGSTRAAL AND OTHERS fowl. Among our numerous records of this species, none is from man. The immat- ure stages have not been reared in the laboratory and identification of nymphs and larvae, from squirrels and Rattus, is tentative. Adults are frequently obtained by sweeping forest vegetation, but in comparat- ively small numbers that appear not to indicate the true incidence of H. konings- bergert in Malayan forests. The questing position and location of adults on vegetat- ion, possibly closer to the ground than usually reached in sweeping, should be investigated. The range of this species includes Borneo, Java, Sumatra, and nearby islands. It would be interesting to know the capacity of H. koningsbergert to transmit Babesia spp. among carnivores and to serve as a vector of viruses and rickettsiae. “Dermacentor auratus group” The true D. auratus Supino is rare in Malaya. We have records of D. auratus only from Johore (Bekok Forest Reserve, Gunong Sumalayang, and Gunong Berem- ban), Trengganu (Bukit Kedap Forest Reserve and Bukit Bintang Besut), Perak (Sungei Klah Forest Reserve), and Pahang (Petoh Forest Reserve). The “D. auratus’ or ““D. auratus group’’ ticks earlier reported in literature on tickborne pathogens in Malaya refer in fact to D. astrosignatus or D. compactus, or to both, but not to D. auratus (sensu strictu). Dermacentor ticks are involved in the cycles of tick typhus (Rickettsia sp.) and of Q fever (Coxiella burnett) in Malayan climax forests (Marchette, 1966). Immature Dermacentor from rodents from Bukit Lanjan, 7 miles from Kuala Lumpur, provided the first isolation of Lanjan virus (Smith et al., 1967). The following specimens from Gunong Benom cannot be identified to species and are considered as ‘‘Deymacentor sp.” Ratufa affinis (7/1), 1200 ft, 1 nymph Rattus sabanus (40/11), 700 to 2200 ft, 2 nymphs, 29 larvae Rattus rvajah (7/1), 1700 ft, t larva Rattus bowersii (2/1), 800 ft, 16 larvae Tragulus javanicus (4/1), lowlands, 1 nymph Domestic dog (Lord Medway’s), 700 ft, 4 larvae Dermacentor atrosignatus Neumann Collections from sweeping forest vegetation and from hosts (examined/infested) are as follows: Callosciurus caniceps (3/1), 700 ft, 2 nymphs Rattus sabanus (40/11), 700 to 2200 ft, 4 nymphs Rattus bowersit (2/1), 800 ft, I nymph TICKS (IXODIDAE) OF GUNONG BENOM 179 Sus scrofa (2/2), 800 and 1400 ft, 273, 92 Man (biting), 1700 ft, 1 nymph Man (crawling on), 750 and 2500 ft, 1g, I nymph Vegetation, 500 to 2500 ft, 161g, 243° Vegetation, 2600 to 5200 ft, 63g, 1092 These 613 adult specimens comprise 84% of the 731 adult Dermacentor ticks taken on Gunong Benom (see also Tables 1-3). The wild pig, Sus scrofa, appears to be the chief host of all adult Dermacentor in Malaya, and probably of most if not all Dermacentor species of tropical Asia. Sus scrofa is the most common and generally distributed large mammal in Malayan forests. In the absence of wild pigs, Dermacentor ticks would probably be rare in this environment. The 2 wild pigs examined on Gunong Benom yielded 52 adult Dermacentor, of which 36 (69%) were D. atrosignatus and 16 (31%) D. compactus. The 530 wild pigs examined throughout Malaya (Table 3) yielded 8294 adult Derma- centor, of which 5498 (66%) were D. astrosignatus and 2796 (34%) D. compactus. More males (61%) than females (39%) were found on wild pigs owing to the male’s longer attachment period and the female’s shorter feeding period. In collections of questing adults from Malayan forest vegetation, females (58%) outnumbered males (42%) (Table 3), as they did on Gunong Benom. TABLE 3 Adult Deymacentor atrosignatus and D. compactus specimens from all Malayan sources (Hoogstraal collection, 20 April 1969) Source D. atrosignatus D. compactus Total 3 & 9 Total 3 g 3 2 3 2 adults Vegetation 3,725 4,750 1,631 2,650 5,350 7,406 12,762 Sus scrofa (530) 3,305 2,133 1,700 1,096 5,005 3,229 8,294 Other hosts (27) 22 43 7 14 29 57 86 Human beings (8) crawling on 3 2 fo) I 3 3 6 biting fo) I I fo) I I 2 Total 7,115 6,935 3,339 3,761 10,454 10,696 21,150 % adults 66 34 Immature-stage hosts of Dermacentor ticks in Malaya are various small size mammals such as insectivores, squirrels and other rodents, carnivores, muntjac, mouse-deer, domestic goats and dogs, and man. Our data for about 800 Malayan nymphs and larvae are not sufficiently extensive or representative to permit dis- cussion of comparative importance of different mammal species as immature-stage hosts. Human beings are seldom recorded as hosts in Malaya in our data and by 180 H. HOOGSTRAAL AND OTHERS Audy e¢ al. (1960). This is in marked contrast to the incidence of human infestation by the true D. auratus (Supino) in India and the terai (lowlands) of Nepal (Hoogstraal, 1970). Most Dermacentor specimens in our Malayan tick collections are from altitudes below 2500 ft. On Gunong Benom, the 2500 ft altitudinal level appears to be the upper limit of distribution of D. compactus, except for an occasional straggler to 2700 ft. On this mountain, tick collections from vegetation up to 2500 ft consisted of 261 (75%) D. atrosignatus and 86 (25%) D. compactus (Table 1). Above this level, however, these percentages were 41 and 0-24, respectively. The only D. compactus specimen taken in the montane forest range was a female at 2700 ft. Different altitudinal distribution results for Dermacentor ticks were obtained on Mt. Ophir, (= Gunong Ledang), Johore, about 125 miles south of Gunong Benom (Malayan Tick Survey data, September 1968). On Mt. Ophir, D. compactus appears to be more numerous in relation to D. atrosignatus than in any other Malayan locality known to us. Of 1253 ticks of both species taken from Mt. Ophir forest vegetation, 698 (56%) were D. atrosignatus and 555 (44%) were D. compactus. The altitudinal distribution of these samples was as follows. altitude number number number total (ft) collections D. atrosignatus (%) D. compactus (%) 1000 3 98 (50%) 97 (50%) 195 1500 4 155 (64%) 86 (36%) 241 2000 5 217 (68%) 103 (32%) 320 2500 3 73 (38%) 119 (62%) 192 3000 I 8 (42%) II (58%) 19 3500 4 114 (54% 99 (46%) 213 4000 2 33 (45%) 40 (55%) 73 total 22 698 (56%) 555 (44%) 1253 The 46 to 58 percentages recorded for D. compactus in relation to D. atrosignatus at altitudes above 2500 ft on Mt. Ophir are in sharp contrast to the data from Gunong Benom. A comparative environmental study of altitudinal forest zones on Gunong Benom and Mt. Ophir might yield data showing the basic ecological re- quirements of each species and reasons for the disproportionate numbers of D. compactus at all levels and especially at upper elevations of Mt. Ophir. Dermacentor compactus Neumann Collections from sweeping forest vegetation and from hosts (examined/infested) are as follows: Sus scrofa (2/2), 800 and 1400 ft, 11g, 59 Vegetation, 500 to 2500 ft, 45g, 562 Vegetation, 2600 to 5200 ft, 12 For comments on these collections, see text on Dermacentor auratus group and D. atrosignatus, and also Tables 1-3. TICKS (IXODIDAE) OF GUNONG BENOM 181 Amblyomma testudinarium Koch Collections from sweeping forest vegetation and from hosts (examined/infested) at various altitudes are as follows: Sus scrofa (2/1), 800 ft, 1 nmyph Man (biting), 2500 ft, 1 nymph Man (crawling on), 700 and 1500 ft, 1g, 29 Vegetation, 500 to 2500 ft, 25g, 209, 2 nymphs Vegetation, 2600 to 5200 ft, 5g, 32, I nymph A. testudinarium occurs practically everywhere in Malayan forests, though usually only in moderate numbers. The incidence on Gunong Benom is either lower than in most other forests of Malaya or collections were made here at periods when adults and other stages were relatively inactive. Forty-seven specimens were taken from vegetation up to 2500 ft, and g at higher altitudes. Only 1 nymph was found on the 2 Sus scrofa examined. Sus scrofa is the chief host of adult A. testudinariwm in Malaya and elsewhere in Asia. The incidence of infestation of Malayan wild pigs is usually almost as high as that of Dermacentor ticks but the numbers of adult A. testudinarium on each pig are almost invariably much lower (unpublished). Other hosts of adults in Malaya are the tapir, tiger, sambar deer, pangolin, and domestic buffalo. We have records of 5 males, 5 females, and 1 nymph parasitizing human beings in Malaya. Adults also feed occasionally on the tortoise, Testudo emys, monitor lizard, Varanus salvator, and python, Python reticulatus, in Malaya (unpublished). Immature A. testudinarium are the most catholic in host preference of all Malayan ticks. They often occur on domestic chickens, goats, dogs, sheep, and buffalo. While no larvae are recorded from human beings by the Malayan Tick Survey, these tiny ticks are easily overlooked and we suspect that infestation of people may be fairly frequent. A nymph was taken biting a person at 2500 ft.on Gunong Benom. The Hooper Foundation group working with the Gunong Benom Expedition exper- ienced heavy infestation of several members by seed ticks (larvae), presumably A. testudinarium, at 3000 to 4000 ft during the wet March-April period but not during the dry month of February (N. J. Marchette, personal communication). Other Malayan hosts of immature stages are (INSECTIVORA), the common treeshrew (Tupaia glis); (CARNIVORA), the Malay bear (Helarctos malayanus), weasel (Mustela nudipes), Malay civet (Viverra tangalunga), common palm civet (Paradoxurus hermaphroditus) (frequent), small-toothed palm civet (Arctogalidia trivirgata), banded palm civet (Hemigalus derbyanus), tiger (Panthera tigris), leopard cat (Felis bengalensis) (frequent), black giant squirrel (Ratufa bicolor), three-striped ground squirrel (Lariscus insignis), common porcupine (Hystrix brachyura) (frequent), wild pig (Sus scrofa) (infrequent), mouse-deer (Tvagulus javanicus) (frequent), cobra (Naja naja), monitor lizard (Varanus salvator), jungle fowl (Gallus gallus) (frequent), and crested green wood-partridge or roulroul (Rollulus roulroul). Aside 182 H. HOOGSTRAAL AND OTHERS from immature stage hosts reported from Vietnam (Hoogstraal et al., 1968), these are almost the only records of immature-stage A. testudinarium. Engorged A. testudinarium reach the largest size (25 x 20 mm) of any tick species in Malaya and a single female may deposit 10,500 eggs. One of us (M. N.) has collected several hundred larvae from forest vegetation a few inches to 3 meters above the ground in the vicinity of a pig wallow in Malaya. In view of the height to which questing larvae climb on plants, it is suprising that this stage is so seldom represented in tick collections from Malayan forest vegetation. A. testudinarium inhabits forests from Ceylon and India to Borneo, Indonesia, Thailand, Vietnam, the Philippines, Taiwan, and southern islands of Japan. It has not been associated with diseases of man and lower animals. Amblyomma geoemydae (Cantor) Six ornate nymphs were collected from lowland forest vegetation and 1 ornate nymph from a coucal, Centropus sinensis, at 1000 ft. A. geomydae parasitizes chiefly reptiles, especially tortoises, Testudo emys, T. spp., and monitor lizards, Varanus spp., in Borneo, Java, Malaya, and the Philippines. Taxonomic problems concerning this species were discussed by Kohls (1957). Disease relationships have not yet been studied. Amblyomma spp. Unidentified Amblyomma specimens from Gunong Benom are 1 larva from Echinosorex gymnurus, t larva from Rattus bowersii, 1 nymph from Centropus sinensis, and 1 nymph from Anthracoceros malayanus. DISCUSSION AND CONCLUSIONS Ticks from sweeping vegetation Sweeping (flagging) yielded 546 ticks in lowland forests, 416 in mountain forests, and none in mountain mossy forests (Table 1). Of 5 tick species usually taken in greatest numbers by sweeping vegetation in Malayan forests [Dermacentor atro- signatus, D. compactus, Haemaphysalis nadchatrami, H. semermis, and Amblyomma testudinartum|], each is represented in varying proportions in both altitudinal regions in which ticks were found. The reduction in numbers of D. compactus from 16% in lowland collections to 0-24 °% in mountain forests, was compensated in these zones by arise in H, nadchatrami numbers, from 6% to 54%. As usual in collections from Malayan forest vegetation, D. atrosignatus comprised at least 40% of the specimens. No Dermacentor were found above 4000 ft. Only H. nadchatrami and H. semermis were taken in the 4000 to 5200 ft range and, as already stated, no ticks were found above the latter level. TICKS (IXODIDAE) OF GUNONG BENOM 183 H. c. cornigera adults and nymphs were also collected from lowland vegetation. Reasons for the apparently erratic distribution of cornigera in Malayan forests are not known. The collection of nymphs of A. geomydae from vegetation is unusual in our experience. The presence of 2 exceptionally rare species, H. hylobatis and H. calvus, among ticks from vegetation is notable. Reasons for the paucity of H. koningsbergeri in collections from Malayan forest vegetation, in areas where this species is believed to be relatively common on small carnivores, remain to be investigated. Ixodes granulatus, a medically important parasite of small, ground- feeding animals, is unrepresented in tick samples from forest vegetation; it is a nidicolous species that rarely if ever quests from vegetation. Ticks from vertebrate hosts Almost all vertebrates examined for ticks on Gunong Benom were in lowland forests (Table 2). Bats of the genera Penthetor and Rhinolophus were infested by larvae, nymphs, and females of Ixodes simplex, which has not previously been recorded from Malaya. The single insectivore, a moonrat, carried J. granulatus adults and immature stages, and an Amblyomma larva. Certain rodents yielded a few larvae and nymphs of H. nadchatrami, H. semermis, H. koningsbergeri (tentative identification), D. atrosignatus, D. sp., and Amblyomma sp., and adults of J. granulatus. Many rodents were not tick-infested at the time of examination (see footnote, Table 2). The feeding period length of many tropical forest larvae and nymphs should be studied in detail to determine whether com- parative rarity of finding these stages on small mammals may be explained by rapid feeding and early dropping from the host. Each of the 5 wild carnivore species examined was parasitized by adults of H. koningsbergeri; a leopard cat was also infested by adults of H. asiatica and H. semermis, a binturong by adults of H. hylobatis and H. semermis, and a banded palm civet by nymphs and larvae of H. vidua. Lord Medway’s dog yielded adult H. koningsbergeri and H. sémermis and larval D. sp. Dogs of aborigine villagers bore adults of H. koningsbergeri and adults and nymphs of H. bispinosa, a parasite that has been introduced into Malaya and seldom infests wild forest animals in this country. The common large mouse-deer, Tvagulus javanicus, was parasitized by nymphs of D. sp. and H. nadchatrami, adults, nymphs, and a larva of H. semermis, and, as usual, by all stages of H. traguli. One smaller mouse-deer, T. napu, yielded only a single larva of H. ?nadchatrami (tentative determination). The comparatively heavy infestation of the 2 wild pigs, Sus scrofa, by adults of D. atrosignatus, D. compactus, H. nadchatrami, H. semermis, and A. testudinarium (in this exceptional case only by a nymph) is typical of Malayan collections from this host. Six of the 9 birds examined on Gunong Benom were infested by specific bird parasites, H. doenitzi and H. wellingtoni, and by a few Amblyomma nymphs, 184 H. HOOGSTRAAL AND OTHERS including 1 of A. geoemydae. A more extensive study of bird-tick interrelationships on Gunong Benom would be certain to be rewarding. Ticks biting human beings in lowland forests were adult H. nadchatrami and H.c. cornigera, adult and nymphal H. semermis, and nymphal A. testudinarium and D. atrosignatus. Adults of these species (except H. c. cormigera) and a nymphal D. atrosignatus were also collected while crawling on people. At 6000 ft, an adult H. nadchatrami was removed while biting and at 3000 ft an adult H. hylobatis was crawling on an expedition member. Numerous larvae, presumably A. testudinarium, infested persons working at the 3000 to 4000 ft level in the rainy season but not during the dry month of February. Medical relationships Owing to the relatively undisturbed forest environment of Gunong Benom and to the continuity of forest from lowlands to altitudinal zones above those inhabited by most Malayan ticks, this mountain area is eminently suitable for investigating the natural history of “primitive’’ cycles of tickborne pathogens infecting man and lower animals. Among the tick speciesfound here, Jvodes granulatus is the original source of the virus (Russian spring-summer encephalitis complex) causing Langat encephalitis. This virus can be transmitted experimentally by other tick species, a finding /that enhances interest in its more widespread presence in nature (see Hoogstraal, 1966). I. granulatus, Dermacentor spp., and Haemaphysalis spp. [including H. nadchatrami and H. semermis| have been shown by Marchette (1966) to be involved in the cycles of tick typhus (Rickettsia sp.) and Q fever (Coxiella burneti) in climax forests of Malaya. Lanjan virus, first recovered from Dermacentor sp. from Bukit Lanjan near Kuala Lumpur, has recently been isolated from Malayan J. granulatus and H. semermis (N. J. Marchette, personal communication). Other tick species that should be studied epidemiologically are H. bispinosa, owing to its close association /with domestic animals, H. koningsbergeri in relation to Babesia infections of carnivores, H. doenitzi and H. wellingtoni to determine their virus-vector potential among wild and domestic birds, and H. traguli in relation to infections in the mouse-deer. The question of virus infections in wild pigs and their chief tick parasites, D. atro- signatus, D. compactus, H. nadchatrami, H. semermis, and A. testudinarium, should also be investigated. ACKNOWLEDGEMENTS Persons from the Institute for Medical Research (Kuala Lumpur) (Malayan Tick Survey) who participated in tick and host collecting during the Gunong Benom Expedition, and who also worked in this area before and afterwards, are Messrs R. S. Ratnam, P. Ramachandran, K. Krishnasamy, A. Mottan, Ahmad bin Bachik, Sharif bin Mansoor, Mohamed bin Wahid, and Yusof bin Idris. Personnel of the TICKS (IXODIDAE) OF GUNONG BENOM 185 Hooper Foundation Arbovirus Research Team, Drs. Nyven J. Marchette, Richard Garcia, and Duncan W. MacVean, assisted by Miss Teh Siew Khim and Messrs John Jeffrey, Harry Lee, Ahmad bin Abdul Hamid, Nagiah Vengitasamy, Chan Wah San, Tee Kem Tho, and Regis de Silva, also participated in ectoparasite collecting and made subsequent trips to the area for additional material and data. Lord Medway kindly reviewed the preliminary manuscript and Dr. Marchette generously pro- vided notes on human infestation by larval ticks and on viruses recently isolated from some of the tick species reported here from Gunong Benom. To each of these persons we express our gratitude for assistance. REFERENCES Aupy, J. R., Napcuatram, M. & Lim, B.-L. 1960. Malaysian parasites XLIX. Host distribution of Malayan ticks (Ixodoidea). Stud. Inst. med. Res. Malaya, (29) : 225-246. HoocstraaL, H. 1962. Rediscovery, rediscription, and relationships of Haemaphysalis vidua Warburton and Nuttall, 1909 (Ixodoidea, Ixodidae) from Malaya. J. Parasit., 48 : 719-722, 14 text-figs. 1964. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodoidea). Re- description, hosts, and distribution of H. tvaguli Oudemans. The larva and nymph of H. vidua W. and N. Identity of H. papuana toxopei Warburton (n. comb.). J. Parasit., 50 : 765-782, 65 text-figs. 1966. Ticks in relation to human diseases caused by viruses. Amn. Rev. Ent., 11 : 261-308. 1967. Ticks in relation to human diseases caused by Rickettsia species. Ann. Fev. Ent., 12 : 377-420. 1970. Human infestation by ticks (Ixodidae) in the Himalaya. Jn: H. D. Srivastava commemoration volume, Singh, K. S. & Tandan, B. K. (Eds.). Div. Parasit. Indian Vet. Res. Inst., Izatnagar; India, pp. 75-89. HoocstraaL, H. & Trapipo, H. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Species described by Supino in 1897 from Burma, with special reference to H. (Rhipistoma) asiaticus (= H. dentipalpis Warburton and Nuttall.) J. Parasit., 56 : 1172-1187, 62 text-figs. Hoocstraat, H., Kouts, G. M. & Parrisu, D. W. 1968. Studies on Southeast Asian Haem- aphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) susphilippensis sp. n., a parasite of Luzon and Mindanao boars. J. Parasit., 54 : 616-621, 20 text-figs. HoocstraaL, H., Lim, B.-L. & Anastos, G. 1969. Haemaphysalis (Kaiseriana) bispinosa Neumann: evidence for consideration as an introduced species in the Malay Peninsula and Borneo. J. Parasit., 55 : 1075-1077. Hooestraat, H., Traprpo, H. & Konts, G. M. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) papuana nadchatrami ssp. n. and rediscription of H. (K.) semermis Neumann. J. Parasit., 51 : 433-451, 56 text-figs. Hooecstraat, H., Roperts, F. H. S., Kouts, G. M. & Tipton, V. J. 1968. Review of Haem- aphysalis (Kaiseviana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, and northeastern China and U.S.S.R., and its partheno- genetic and bisexual populations (Ixodoidea, Ixodidae). J. Parasit., 54 : 1197-1213. 36 text-figs. Hoocstraat, H., Santana, F. J. & VAN PEENEN, P. F. D. 1968. Ticks (Ixodoidea) of Mt. Sontra, Danang, Republic of Vietnam. Ann. ent. Soc. Am., 61 : 722-729, 1 text-fig. Kouts, G.M. 1949. Haemaphysalis centropi, a new species of tick from birds in the Far East. J. Pavasit., 35 : 388-390, 1 text-fig. 186 H. HOOGSTRAAL AND OTHERS Kouts, G. M. 1957. Malaysian parasites XVIII. Ticks (Ixodoidea) of Borneo and Malaya. Stud. Inst. med. Res. Malaya, (28) : 65-94. MarcHetteE, N. J. 1966. Rickettsioses (tick typhus, Q fever, urban typhus) in Malaya. J. med. ent., Honolulu., 2 : 339-371. — 1967. Anexpedition to Gunong Benom. Hooper Newsvector, 1 : 2-4. Mepway, Lorp. 1972. The Gunong Benom Expedition, 1967, 1. Introduction. Bull. Brit. Mus. nat. Hist. (Zool.) 23: 1-7. NapDcHATRAM, M. 1960. Malaysian parasites XLVIII. Notes on rearing Malayan ixodid ticks (Acarina, Ixodidae) with special reference to Ixodes granulatus Supino and Amblyomma geoemydae (Cantor). Stud. Inst. med. Res. Malaya, (29) : 217-224. SmitH, C. E.G. 1956. A virus resembling Russian spring-summer encephalitis virus from an ixodid tick in Malaya. Nature, Lond., 178 : 581-582. Situ, C. E. G., McManon, D. A. & Bowen, E. T. W. 1967. Lanjan virus, a new agent isolated from Dermacentor auratus in Malaya. Nature, Lond., 214 : 1154-1155. Warsurton, C. & Nuttatt, G. H. F. 1909. On new species of Ixodidae, with a note on abnormalities observed in ticks. Parasitology, 2 : 57-76, 26 text-figs. WuitmoreE, T.C. 1972. The Gunong Benom Expedition, 1967, 2. An outline description of the forest zones on northeast Gunong Benom. Bull. Brit. Mus. nat. Hist. (Zool.) 23: 9-15. a “ AS NAT. HIST. - 8 NOV 1972 =, PRESERTED Py O% cay Ocy i> Dr. H. HooGstraaL NAMRU-3 AMERICAN INTERESTS SECTION SPANISH EMBASSY Cairo, EGypt B. L. Lim INSTITUTE FOR MEDICAL RESEARCH Kuara Lumpur, MALAYSIA M. NADCHATRAM INSTITUTE FOR MEDICAL RESEARCH Kuarta Lumpur, MALAYSIA G. ANASTOS Zoology Department UNIVERSITY OF MARYLAND COLLEGE PARK MarYLanp, U.S.A. A LIST ‘OF. SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. {4. TAyLor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. A 1962-1964. (In Press). pee Bee Printed in England by Staples Printers Limited at their Kettering, Northants, establishment TERS HE GUNONG BENOM EXPEDITION ee! 1967 . A COLLECTION OF CHIGGERS FROM --- ~ GUNONG BENOM _ (PROSTIGMATA: TROMBICULIDAE) © -—s M, NADCHATRAM HIRAPTERA (INSECTA) CHEWING AND SUCKING LICE THERESA CLAY ‘BULLETIN OF | MUSEUM (NATURAL HISTORY) bs ve LOND ON: 1972 ety oe 5°. Weliay Wawa ee THE GUNONG BENOM EXPEDITION 1967 9. A COLLECTION OF CHIGGERS FROM GUNONG BENOM (PROSTIGMATA: TROMBICULIDAE) M. NADCHATRAM Bishop Museum, Honolulu to. PHTHIRAPTERA (INSECTA) CHEWING AND SUCKING LICE THERESA CLAY Pp. 187-200 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 8 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 18 issued in five series, corresponding to the Departments of the Museum, and an Historical serves. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 23, No. 8 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 4 October 1972 Price 65p THE GUNONG BENOM EXPEDITION 1967 9. A COLLECTION OF CHIGGERS FROM GUNONG BENOM (PROSTIGMATA: TROMBICULIDAE) By M. NADCHATRAM TuIs report summarizes the trombiculid mites collected on Gunong Benom from 17 February through 8 April 1967. Topography and vegetation are described by Medway (1972a) and Whitmore (1972), and the taxonomy and distribution of hosts by Grandison (1972), Hill (1972) and Medway (1972b). Although the Institute for Medical Research in the past has made intensive as well as extensive collections of trombiculid mites as part of an investigation of the epidemiology of scrub typhus, this is the first time that these and other ectoparasites have been collected on Gunong Benom. The knowledge of the mite fauna of this isolated mountain adds to the information being gathered for zoogeographical and ecological studies. Furthermore, it will enhance our understanding of the eco- logical relationships of scrub typhus vectors to topography and vegetation. Mites of the family Trombiculidae (sub-class Acari) are parasitic only in the larval stage (when they are known as chiggers), yet they are amongst the most abundant, both in numbers and species, of all ectoparasites of mammals. They are soft-bodied and range from 0-2 to 2 mm in length. The post-larval stages, i.e. the nymph and adult, are non-parasitic. The colour of chiggers ranges from white or yellow to light orange or red. _ Chig- gers are essentially habitat-specific, and their colour is to a great extent dependent on their biotope. They may be separated into two broad ecological groups— ground-surface dwellers and nidicolous species. Ground-surface dwellers include the scrub-itch mites and the vectors of scrub typhus, and are coloured orange to red. Nidicolous species, with a few exceptions, are white to yellow, and represent almost 83 per cent of the 146 species of chiggers known in Malaya. Of these 70 to 80 per cent are cavernicolous in habitat, i.e. live in the ground-burrows of small mammals. Nadchatram (1970) has proposed seven ecological groups for the species of chiggers known in Malaya (Ecological Groups I to VII) on the basis of (a) the colour of the chigger prior to attachment to a host, (b) association of the chiggers with habitat and (c) their association with the favoured hosts. Under each species reported below the appropriate ecological group number is given. The chiggers of medical importance in Malaya known thus far are of two categories. Those that transmit the agent of scrub typhus are Leptotrombidium akamushi (Brumpt) mostly found in grassland, L. deliense (Walch) mostly found in plantations, forest and the forest fringe and, probably, L. arenicola Traub restricted to seacoast Bull. Br. Mus. nat. Hist. (Zool.) 23, 8 190 M. NADCHATRAM vegetation. The other category of chiggers causes dermatitis in man They are Eutrombicula wichmanni (Oudemans), Blankaartia acuscutellaris (Walch), and Schoengastia psorakart Nadchatram and Gentry. Out of 177 examined, 87 reptiles, birds and mammals were infested with chiggers. Over 3,000 chigger specimens from animal hosts and from ground-holes were col- lected, representing 35 species in 14 genera. L. deliense was found only in small numbers, but a very closely related species, L. bodense (Gunther), was abundant. No other of the medically important species listed above were found, although the genus Leptotrombidium, which is the important genus from the viewpoint of epidemi- ology of scrub typhus, is represented by 5 species. The collection yielded a new species of Myotrombicula from Rhinolophus stheno (Nadchatram and Lakshumy 1969), and a new record of chiggers for Malaya, Neoschoengastia solitus Nadchatram, from Pitta sordida. The infestation data is presented in three parts. The first is a list of chiggers found in Gunong Benom with relevant notes on host and habitat distribution. The second part is a list of the infested animals and the species of chiggers found on them, and the third a list of unattached chiggers found in holes. OBSERVATIONS 1. Chiggers found on Benom Audyana thompsoni Womersley, 1954 14 ex I Heterometrus longimanus, 750 ft. This larval species is not a trombiculid, but belongs to a closely allied family, Trombidiidae; it is often found together with a trombiculid species, Eltonella eltoni. Colour pallid. Ecological Group VII. Odontacarus audyi (Radford, 1946) Two specimens collected, with Toritrombidula densipiliata, from Pitta caerulea, 800 ft. Orange chiggers parasitic exclusively on birds and widely distributed in Southeast Asia. Indications are that this species is a ground-surface dweller. Ecological Group I. Eltonella tweediei Audy, 1956 29 specimens ex 8 Dyaco spp., 750 to 800 ft. Because of concurrent habitat- specificity of host and parasite, this species has never been found on other animals. Often found with Neoschoengastia riversi. An arboreal species, bright orange to red. Ecological Group III. Siseca rara (Walch, 1923) One ex Pitta sordida, goo ft and 2 ex 1 Callosciurus caniceps, 900 ft. This is a common species found on reptiles, birds and mammals, including man (in Sumatra). Unfed larvae were recovered from ground-surface and nests of squirrels. Orange to red. Ecological Groups I and III. CHIGGERS FROM GUNONG BENOM 191 Leptotrombidium bodense (Gunther, 1940) A very common species in Malayan forests, it is represented in the Gunong Benom collection as follows: 6 ex 1 Napothera macrodactyla, 700 ft; 3 ex 1 Echino- sorex gymnurus, 700 ft; 24 ex 6 Rattus sabanus, 700 to 1000 ft; 8 ex 1 R. muelleri, 800 ft; 36 ex 1 R. bowersi, 800 ft; 6 ex I R. cremoriventer, 800 ft; 124 ex 4 R. ed- wardsi, 3500 to 4000 ft. Unfed larvae of L. bodense have been found on ground- surface and in ground-burrows. Ground-surface chiggers are orange and hole chiggers pallid. Nadchatram & Upham (1966) synonymized L. langati Audy & Womersley with L. bodense. Ecological Groups I and II. Leptotrombidium deliense (Walch, 1922) A well-known vector of scrub typhus, this species is poorly represented in the Gunong Benom collection, as follows: 1 ex 1 Pitta sordida, goo ft; 1 ex I R. sabanus, 800 ft; 16 ex 1 R. muelleri, 800 ft; 13 ex 3 R. edwardsi, 3600 to 4000 ft. This species, however, is common in plantations and the forest fringe. It has been recorded frequently from ground-surface, ground-burrows, and occasionally in nests of rats and squirrels on trees. It is believed that L. deliense evolved from mites of a sylvatic origin and a cavernicolous habitat. Like L. bodense, hole chiggers are pallid and ground-surface dwellers orange. Ecological Groups I and II. Leptotrombidium gentryi Nadchatram & Upham, 1966 This species was first described from unfed larvae collected in ground-holes. The Gunong Benom collection includes 2 specimens collected on a ‘bakelite’ plate in two separate ground-burrows in a bank alongside a disused logging track, 23 ft above the ground, at 1000 ft. The host is still unknown. Pallid. Ecological Group II. Leptotrombidium kundini Nadchatram & Upham, 1966 This species was also taken by plate collections. 13 specimens found in 4 ground- burrows in the bank of the logging track at 1000 to 1200 ft. A pallid species, in Ecological Group II. Leptotrombidium sylvestre (Audy & Traub, 1950) 39 specimens ex 3 R. edwardsi, 3500 to 4000 ft. This is also an uncommon sylvatic species and appears to be associated more with R. edwardsi than other rodent hosts. This collection further confirms that L. sylvestre is a denizen of montane and sub-montane forests. Pallid chiggers in Ecological Group IT. Toritrombicula densipiliata (Walch, 1922) 12 ex I Pitta caerulea, 800 ft; 1 ex Pitta sordida, goo ft; and 3 ex 1 Philentoma velata, 1800 ft. This species is restricted to birds and widely distributed throughout Southeast Asia, possibly as a result of the migratory habit of some of its hosts (including P. sordida). Orange chiggers in Ecological Group I. 192 M. NADCHATRAM Myotrombicula medwayi Nadchatram & Lakshumy, 1969 Gunong Benom is the type locality for this species. A single specimen ex Rhino- lophus stheno, 700 ft. It is only the second species of the genus known in Malaya. Orange chiggers in Ecological Group V. Chiroptella sandoshami Nadchatram, 1966 23 ex 15 Rhinolophus stheno, 700 to 3600 ft; one ex R. affinis, 2500 ft; 2 ex I R. macrotis, 750 ft. The 24 species of chiggers known from bats in Malaya are apparently exclusive to these mammals. Whether species of chiggers are specific to any species or genus of bats is not yet clear. C. sandoshami is a common parasite of Rhinolophus, but the apparent host-specificity may be due to ecological factors. Yellow chiggers in Ecological Group V. Neoschoengastia riversi Wharton & Hardcastle, 1946 18 ex 1 Draco sp., 800 ft. Members of the genus Neoschoengastia are mostly parasites of birds. In Malaya, however, N. riversi is frequently found on flying lizards, although the type host of the species is a bird. Orange chiggers in Eco- logical Group III. Neoschoengastia solitus Nadchatram, 1967 4 ex I P. sordida, goo ft. This is the first record of its occurrence in Malaya. The host is a migratory bird*. Orange chiggers in Ecological Group IV. Walchiella impar (Gunther, 1939) 50 ex I Echinosorex gymnurus, 700 ft; 16 ex 1 Tupaia minor, 700 ft; 20 ex I Ptilocercus lowii, 1700 ft; and 5 ex 1 R. edwardsi, 4000 ft. This species is closely related to W. oudemanszi, but restricted in habitat. Pallid chiggers in Ecological Group II. Walchiella lacunosa (Gater, 1932) 149 ex 4 R. edwardsi, 3500 to 4000 ft. This is a sub-montane species commonly parasitizing R. edwardsi. Yellow chiggers in Ecological Group II. Walchiella oudemansi (Walch, 1923) 2ex 1 E. gymnurus, 700 ft; 127 ex 6 R. sabanus, 700 to 800 ft; ror ex 2 R. muelleri, 700 to 800 ft; 5 ex 1 R. bowersi, 800 ft; 13 ex 1 Sundasciurus lowii, 800 ft; 2 ex 2 Callosciurus notatus, 1000 ft; 26 ex 2 Lariscus insignis, 700 to 800 ft; 122 ex I *A new species of Tovitvombicula was described from the same species of host in the Cameron Highlands (Nadchatram, 1967). The species, Tori. uphami, was found with Tori. densipiliata and Odon. audyt. While the two latter species have been found both on migratory and resident birds in Malaya, it would seem that the host of T. wphami was a recent arrival in Malaya. CHIGGERS FROM GUNONG BENOM 193 Rhinosciurus laticaudatus, 700 ft; and 2 ex 1 Hemigalus derbyanus, goo ft. This common species was found on a variety of ground-dwelling and, to a lesser degree, tree-dwelling small mammals, and has been found on man in Indonesia. The range of habitat is also broad. Ground-hole chiggers are pallid and ground-surface dwellers orange. Ecological Groups I, II, and III. Ascoschoengastia audyi (Womersley, 1952) 7 ex I P. lowii, 1700 ft; 5 ex 2 R. sabanus, 700 ft; 42 ex 2 C. nigrovittatus, 700 ft; 164 ex 5 C. notatus, 700 to 1000 ft; and Io ex 1 Iomys horsfieldi, 800 ft. R. sabanus is an unusual host; this chigger is a common arboreal species frequently associated with tree squirrel in forests and plantations. Traub et al, (1950) recovered Rick- ettsia tsutsugamushi from A. audyi. Pink to orange. Ecological Group III. Ascoschoengastia ctenacarus Domrow, 1962 3 ex I I. horsfieldii, 800 ft. This is an uncommon arboreal species. Orange chiggers. Ecological Group III. Ascoschoengastia indica (Hirst, 1915) 2 ex I R. edwardsi, 4000 ft; and 2 ex 1 C. nigrovittatus, 800 ft. The rarity of this species in Gunong Benom is expected since it is essentially associated with commensal rats, especially the house rat Rattus (r.) diardii. It has an extensive geographical, ecological and host distribution. Gispon (1950) recovered the infective agent of murine fleaborne typhus, Rickettsia mooseri, from A. indica collected from R. (r.) diavdii and R. norvegicus in Indonesia. White to orange chiggers. Ecological Groups I, II and IIII. Ascoschoengastia roluis (Traub & Audy, 1954) 4 ex I Ptilocercus lowit, 1700 ft; 4 ex 1 Iomys horsfieldi1, 800 ft. This is an uncom- mon species. White chiggers. Ecological Group III. Helenicula mutabilis (Gater, 1932) 2 ex I Rattus sabanus, 800 ft; 3 ex I R. muellerz, 800 ft; 6 ex R. bowerst, 800 ft; and 20 ex 1 Hemigalus derbyanus, goo ft. This species is essentially a ground- surface dweller most frequently found in grassland and is therefore presumed to be a secondary parasite of these forest rats. Orange chiggers. Ecological Group I. Doloisia brachypus (Audy & Nadchatram, 1957) 2 ex I Sundasciurus lowii, 800 ft. This squirrel is an unusual host for the species. The genus Doloisia represented by Io species in Malaya, is exclusively intranasal and previously known only from murids. Pallid chiggers. Ecological Group II. 194 M. NADCHATRAM Gahrliepia (Gahrliepia) cetrata Gater, 1932 2 ex 2 R. sabanus, 900 to 950 ft; 1 ex R. edwardsi, 4000 ft. An uncommon, pallid chigger. Ecological Group II. Gahrliepia (Gahrliepia) fletcheri Gater, 1932 4 ex I R. sabanus, 800 ft. This is one of the commonest species of Gahrliepia in the Malayan forests. Its replacement by G. neteralla, a very closely related species, in Gunong Benom is interesting. Pallid chiggers. Ecological Group II. Gahrliepia (Gahrliepia) insigne Womersley, 1952 A single specimen ex R. sabanus, 700 ft. Yellow chiggers. Ecological Group II. Gahrliepia (Gahrliepia) neteralla Traub & Morrow, 1955 I ex Ptilocercus lowii, 1700 ft; and 644 ex 23 Rattus sabanus, 700 to 1500 ft. This is the second commonest chigger species taken on Gunong Benom. The distribution of this species is spotty, with localized dense populations in some areas, few or none in other. Our records show a similar dense infestation pattern in the Ampang Forest Reserve, Selangor, where the common host was also R. sabanus. In most other areas investigated, although R. sabanus was caught frequently, G. neteralla was either absent or occurred in fewer numbers. Pallid chiggers. Ecological Group II. Gahrliepia (Gahrliepia) picta Traub & Morrow, 1955 1 ex R. sabanus, 950 ft; 1 ex R. bowersi, 800 ft. This is a rare species. Unfed larvae were recovered from ground-burrows elsewhere, Pallid chiggers. Ecological Group II. Gahrliepia (Gahrliepia) rutila Gater, 1932 g ex 2 R. sabanus, 700 to 1000 ft; 4 ex 1 R. edwardsi, 4000 ft. This is a rare species; unfed larvae are known from ground-burrows. Pallid chiggers. Eco- logical Group II. Gahrliepia (Gahrliepia) sp. 9 ex 6 R. sabanus, 700 to 1000 ft. This indeterminate species is intermediate between G. fletcheri and G. neteralla. Pallid chiggers. Ecological Group II. Gahrliepia (Schoengastiella) argalea Traub & Morrow, 1957 A single unfed specimen recovered from ground-burrow in the bank of the old logging track, goo ft. The favoured host of this species is R. sabanus. Pallid chiggers. Ecological Group II. CHIGGERS FROM GUNONG BENOM 195 Gahrliepia (Walchia) alpestris Traub & Morrow, 1957 A single specimen ex Ptilocercus lowii, 1700 ft. A rare species. Pallid chiggers. Ecological Group IT. Gahrliepia (Walchia) disparunguis pingue (Gater, 1932) 66 ex 2 Rattus whiteheadi, 800 ft; 36 ex I Rattus inas, 3600 ft; and 13 ex I Rattus tiomanicus, 3600 ft. R. whiteheadi is the favoured host of this species. Pallid chiggers. Ecological Group II. Gahrliepia (Walchia) rustica Gater, 1932 14 ex 2 R. sabanus, 800 ft; 156 ex 14 ground-burrows in bank of forest path. Pallid chiggers. Ecological Group II. Gahrliepia (Walchia) turmalis Gater, 1932 2 ex 1 Echinosorex gymnurus, 700 ft; 831 ex 20 Rattus sabanus, 700 to 1500 ft; 46 ex I R. bowersi, 800 ft; 6 ex 1 R. edwardsi, 4000 ft; and 5 ex 1 ground-burrow in bank of forest path. This species is close to G. rustica, and the commonest species taken in Gunong Benom. Pallid chiggers. Ecological Group II. 2. Hosts List of numbers infested (number examined in parentheses) and species of chiggers. REPTILES Draco spp. 8 infested (of 16): Eltonella tweediei, Neoschoengastia riverst BirpDs Pitta caerulea i (1): Odontacarus audyt, Toritrombicula densipiliata Pitta sordida 1 (1): Leptotrombidium deliense, Tori. densipiliata, Siseca rara, Neoschoengastia solitus Napothera macrodactyla x (1): L. bodense Philentoma velata i (1): Tort. densipiliata MAMMALS Echinosorex. gymnurus 1 (1): L. bodense, Walchiella impar, W. oudemansi, Gahrliepia (Walchia) turmalis Rhinolophus stheno 16 (54): Myotrombicula medwayi, Chiroptella sandoshami Rhinolophus affinis i (21): C. sandoshanu Rhinolophus macrotis i (1): C. sandoshami Tupaia minor I (1): W. oudemansi Ptilocercus lowii 1 (1): Ascoschoengastia audyi, A. roluis, W. impar, G. (W.) alpestris 196 M. NADCHATRAM Rattus sabanus 24 (33): L. deliense, L. bodense, A. audyt, W. oudemansi, Helent- cula mutabilis, G. (W,) rustica, G. (W.) turmalis, G. (G.) cetrata, G. (G.) insigne, G. (G.) neteralla, G. (G.) picta, G. (G.) rutila, G. (G.) fletcheri, G. (G.) sp. Rattus muellert 2 (3): L. deliense, L. bodense, W. oudemanst, H. mutabilis Rattus whiteheadi 2 (5): G. (W.) disparunguis pingue Rattus bowersti 2 (2): L. deliense, W. oudemansi, H. mutabilis, G. (W.) turmalis, G. (G.) picta Rattus cremoriventer 1 (2): L. bodense Rattus edwardst 4 (4): L. deliense, L. bodense, L. sylvestre, A. indica, W. impar, W. oudemansi, G. (W.) turmalis, G. (G). cetrata, G. (G.) rutila Rattus inas i (2): G. (W.) disparunguis pingue Sundasciurus lowit 2 (4): W. oudemansi, Doloisia brachypus Callosciurus nigrovittaus 2 (6): A. audyi, A. indica Callosciurus notatus 5 (5): A. audyi, W. oudemanst Callosciurus caniceps i (1): Siscea vara Lariscus insignis 2 (3): W. oudemansi Rhinosciurus laticaudatus 1 (3): W. oudemansi Iomys horsfieldit 1 (3): A. audyi, A. ctenacarus, A. indica Hemigalus derbyanus 1 (1): W. oudemansi, H. mutabilis Tragulus javanicus i (1): L. bodense Heterometrus longimanus i (1): Audyana thompsoni 3. Unattached chiggers Aside from the examination of animals for chiggers and other ectoparasites, ground-holes, shallow cavities in the ground, depression under over-hanging roots, and leaf-litter on the forest floor were examined with the aid of strips or plates of black “bakelite’. Chiggers were found only in tunnels which are presumed to be rat-burrows. Of approximately 40 holes sampled 16 were positive for chiggers, which were represented by the following four species: Leptotrombidium gentry, L. kundini, Gahiliepia (W.) rustica and G. (W.) turmalis. G. rustica was the com- monest species of hole chiggers. One of the holes found positive for chiggers also yielded several larvae of the common rodent tick, Ixodes granulatus. DISCUSSION With 35 species, the chigger fauna recorded from Gunong Benom is considered rich considering the short duration of the survey. While this collection fills a gap in our knowledge of the overall distributional pattern of Malayan chiggers, the mat- erial reported above is typical of other forests of the Malayan mainland. The survey supports the ecological findings of the author, and in particular the ecological grouping of Malayan species (Nadchatram, 1970). The forest is the reservoir of parasites and, possibly, infections. The constant environment that the forest provides, plus the microclimatic conditions of the subterranean nests of the host animals—ideal for the existence of forest species—, serve to explain why approxi- mately 80 per cent of the 146 Malayan species are endemic in forest. The ease with —— CHIGGERS FROM GUNONG BENOM 197 which the newly hatched larvae of cavernicolous species attach to the host animal (by dropping from the roof of the burrow) is perhaps one of the major contributing factors for the vigorous propagation of the trombiculid fauna. Several species, i.e. 17 percent of 146 species, proven to be adaptable to fluctuating environmental conditions, are believed to have evolved from a sylvatic origin and perhaps a cavernicolous habitat. These species are found on ground-surface, stems of grass, and leaf-litter on forest floor, and include all species which have been indicted with causing scrub-typhus and scrub itch. Because of the habitats they occupy, these are the species which have the best opportunities of coming into contact with man (Ecological Group I). The writer’s observations support the well-known belief that scrub typhus is a man-made malady. When the forest environment is disturbed by man, artificial environments are created which disrupt the natural conditions in which chiggers occur. Some sylvatic species are unable to tolerate the artificial conditions and perish; some that have the opportunity with- draw to their natural retreats, while a few others become adapted to the new situ- ation created by deforestation and flourish. In this new ecological situation other groups of hosts are also encouraged, i.e. ground birds, reptiles, and mammals. Some chigger species, including Leptotrombidium deliense the well-known vector of scrub typhus, are readily dispersed by birds and rats. Rattus tiomanicus jalorensis, a wide-ranging rat, is a common host of L. deliense and most numerous in oil-palm, coconut, and rubber plantations. It also has been collected in sub-montane forest and the fringes of primary and secondary forests. The 35 Gunong Benom species are placed in six ecological groups. The eight species which fall in Ecological Group I were all collected around the base camp between 700 and goo ft, where the vegetation is mostly secondary as a result of previous logging. ACKNOWLEDGEMENTS I am indebted to Lord Medway, formerly of the School of Biological Sciences, University of Malaya, Kuala Lumpur, leader of the expedition, for inviting me and other members of the Acarology Staff, I.M.R. to participate in this expedition, and for other courtesies; to the Director, I.M.R., for approving our participation; the Hooper Foundation for meeting part of the field expenses from U.S.P.H.S. Grant TW-o00144-05 to the University of California (I.C.M.R.T.); to Dr. J. L. Gressitt, Chairman, Department of Entomology, Bishop Museum, Honolulu, for allowing me to complete this study under N.I.H. Grant Al-07575 during my term of service in his department; and to Messrs Lim Boo Liat, R. S. Ratnam and Wan Mohamed for their assistance in collecting the ectoparasites. I am especially grateful to Mr. R. S. Ratnam and Miss G. Thanalakshumy for extracting infestation data from the I.M.R. records and other technical assistance during my absence from the Institute. REFERENCES Aupy, J. R. 1956. Trombiculid mites infesting birds, reptiles and arthropods in Malaya, with a taxonomic revision, and descriptions of a new genus, two new subgenera, and six new species. Bull. Raffles Mus. 28: 27-80. 198 M. NADCHATRAM Aupy, J. R. & Traus, R. 1950. Tvombicula sylvestris sp. nov. (Acarina, Trombiculidae) from Malaya. Bull. Raffles Mus. 23 : 235-237. Aupy, J. R. & Napcuatram, M. 1957. Malaysian Parasites—XXVI. New intranasal species of Tvaubacarus n. gen. (Acarina, Trombiculidae). Stud. Inst. med. Res., Malaya 28: 187-230. Domrow, R. 1962. Four new species of Laurentella from S.E. Asia, (Acarina, Trombiculidae). Acarologia 4 : 586-594. GisPEN, R. 1950. The virus of murine typhus in mites (Schoengastia indica, fam. Trombi- culidae). Docum. néerl. indon. hortus trop. 3: 225-230. Granpison, A. G. C, 1972. Reptiles and Amphibians of Gunong Benom with a description of a new species of Macrocalamus. Bull. Br. Mus. nat. Hist. (Zool.) 23 : 43-101. NapcHATRAM, M. 1966. Revision of the bat-infesting chiggers of Chivoptella Vercammen- Grandjean (Acarina, Trombiculidae), with descriptions of two new larval species and a nymph. J. med. Ent. 3 : 19-28. 1967. Notes on the genus Torvitvombicula Sasa et al. with designation of neotype, des- cription of nymph and redescription of larva of Tori. densipiliata (Walch), and descriptions of two new species from Southeast Asia (Acarina, Trombiculidae). J. med. Ent. 4: 401- 415. 1967. Descriptions and re-descriptions of five species of Neoschoengastia (Acarina, Trom- biculidae) from the Oriental-Australian region. Acavologia 9 : 141-151. (1970). Notes on the habitat of chiggers, correlation of environment and chigger coloration, and its potential significance in the epidemiology of scrub typhus in Malaya (Prostigmata, Trombiculidae). J. med. Ent. 7: 131-144. Napcuatram, M. & UpHam, R. W. JR. 1966. Six new species of sylvatic Leptotrombidium Nagayo et al. (Acarina, Trombiculidae) from Malaysia, with some ecological notes. J. med. Ent. 3 : 101-124. NapcuHatraM, M. & LaxsHumy, G. T. 1969. A new species and a new record of chiggers from West Malaysia (Prostigmata : Trombiculidae). J. med. Ent. 6 : 283-285. Mepway, Lorp. 1972a. Gunong Benom : Introduction. Bull. Br. Mus. nat. Hist., (Zool). 23 : 1-7. 1972b. Distribution and altitudinal zonation of birds and mammals on Gunong Benom. Bull. Br. Mus. nat. Hist., (Zool.) 23 : 103-153. Travus, R., Frick, L. P., & Diercxs, F. H. 1950. Observations on the occurrence of Rick- ettsia tsutsugamushi in rats and mites in the Malayan jungle. Am. J. Hyg. 51 : 269-273. Traus, R. & Aupy, J. R. 1954. Malaysian Parasites—V. Species of Euschoengastia sensu lato (Acarina, Trombiculidae) from Borneo. Stud. Inst. med. Res., Malaya 26 : 77-88. Traus, R. & Morrow, M. L. 1955. A revision of the chiggers of the subgenus Gahrliepia (Acarina, Trombiculidae). Smithsonian Misc. Coll. 128 : 1-89. Traus, R. & Evans, T. M. 1957. Malaysian Parasites— XXVIII. Records and descriptions of chiggers of the subgenus Walchia Ewing, 1931, from Southeast Asia (Acarina, Trom- biculidae). Stud. Inst. med. Res., Malaya 28 : 297-358. WuitmoreE, T.C. 1972. An outline description of forest zones on north-east Gunong Benom. Bull. Brit. Mus. nat. Hist., (Zool.) etc. WomersLeEy, H. 1952. The scrub-typhus and scrub-itch mites (Trombiculidae, Acarina) of the Asiatic-Pacific region. Rec. S. Aust. Mus. 10 : 1-673. — 1954. Malaysian Parasites—VII. New genera and species, apparently of Apoloiinae (Acarina, Leeuwenhoekiidae), from the Asiatic-Pacific region. Stud, Inst. med. Res., Malaya (26) : 108-119. M. NapcHATRAM Institute for Medical Research Kuara Lumpur, MALAYSIA. THE GUNONG BENOM EXPEDITION 1967 to. PHTHIRAPTERA (INSECTA) : CHEWING AND SUCKING LICE By THERESA CLAY MALtopuaGa (Chewing lice) were collected from 26 species of birds and Anoplura (Sucking lice) from two species of mammals. Many of the Mallophaga belong to genera (marked with *) parasitic on the Passeriformes which need revising before accurate determinations can be given; this material will be valuable for such revis- ions. MALLOPHAGA PARASITE HOST Menoponidae Cuculiphilus snodgrassi (Kellogg & Kuwana, 1902) Phaenicophaeus diardi. (BN. 122) Bucerocolpocephalum degnani Elbel, 1967 Anorrhinus galeritus BB. 09 Garrulax erythrocephalus. BN. 372 Garrulax mitvatus BN. 381 | eri viridis. BN. 116 i * SUSIE: pees leucotis BN. 432 Heterophasis picaoides BA. 31 Hypsipetes viridescens BA. 08 Menacanthus sp.* Arachnothera magna BN. 319 Laemobothriidae Laemobothrion maximum (Scopoli, 1763) Ictinaetus malayensis BN. 141 Ricinidae Ricinus tinctus, Harrison, 1916 Crinigery ochraceus BN. 457 Philopteridae Craspedorrhynchus sp. Ictinaetus malayensis BN. 141 Phaenicophaeus curvirostvis BN. 166 Phaenicophaeus javanicus BN. 160 Rallicola unguiculatus (Piaget, 1880) Centropus sinesis BN. 170; BNN. 6 Cuculicola sp.* Bull. Br. Mus. nat. Hist. (Zool.) 23, 8 200 tf. CLAY MALLOPHAGA (Contd.) PARASITE HOST Harpactes diavdi. BN. 452 Pitta caerulea BN. 7 Terpsiphone paradisi BN. 461 ’ Philopterus sp.* Rhipidura albicollis BN. 332, 337 Hypsipetes flavulus BN. 397 Hypsipetes viridescens BM. 08 | Dicrurus remifer BN. 299, 303; BA. 50 Sturnidoecus sp. Nymphs only. Indicator archipelagus BB. 05 Megalaima henrict NB. 171 Megalaima corti BN. 342, 326 Pomatorhinus hypoleucos BA. 40, 20 eae erythrocephalus BN. 372; BA. 33 Brueelia sp.* Garrulax mitratus BN. 381 J Megalaima henrici BN. 17 j * EA Hig ODS \ Psolopogon pyrolophus BA. 37 ANOPLURA PARASITE HOST Neohaematopinus callosciuri Johnson, 1959 Callosciurus notatus Neohaematopinus batuanae Ferris, 1923 Petaurista petaurista Dr THERESA CLAY Department of Entomology British MusEuM (NaTuRAL History) CROMWELL Roap Lonpon, SW7 5BD bal Y - } 5 ¢ f 1 5 ve ha 5 é Hes ¥ 4 P a : J ¥ 7 om » - z ih A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Atison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. {4. Taytor, J. D., KENNEDY, W. J. & Haiti, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates — 7 77 Text-figures. 1969. £4.50. : Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In Press). ‘ q Bt Printed in England by Staples Printers Limited at their Kettering, Northants, establishment ar E GUNON G BENOM EXPEDITION | 1967 _(SIPHONAPTERA), BASED ON COLLECTIONS FRO} MC GUNONG BENOM AND ELSEWHERE IN | 3 _ SOUTH-EAST ASIA -_-R. TRAUB BULLETIN OF - MUSEUM (NATURAL HISTORY) ges 2.) Mol 2gceNoe : _ LONDON 1072 hak THE GUNONG BENOM EXPEDITION 1967 11. NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA 1. NEW TAXA (PYGIOPSYLLIDAE, PYGIOPSYLLINAE) P5\ [es Wo, PRE SENITED 4 Bog, LIBS — ROBERT TRAUB University of Maryland Pp. 201-305; 58 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 9 LONDON : 1972 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 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 23, No. 9 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 14 December, 1972 Price £7.15 THE GUNONG BENOM EXPEDITION 1967 NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA NEW TAXA (PYGIOPSYLLIDAE, PYGIOPSYLLINAE) By ROBERT TRAUB CONTENTS Page SyNopsIs ‘ : ‘ : : : ; 5 c : 3 204 INTRODUCTION 5 é : : : 3 : ¢ : 204 MATERIALS AND Mxetuops : . - é : 5 205 Methods of Collection and Handling : : 0 : : é 205 Major Collection Areas . : ; : 207 Introductory Remarks Conceiae the Descriptions 3 : , 208 Format and Keys . : : 208 Landmarks and Points of Reference Used in Taxonomy 0 : 209 A. Medwayella gen. nov.~ : : : 5 é é : 211 1. WM. dryadosa sp. nov. 215 Host-Relationships and Distribution ard Other Commtens 226 M. arcuata sp. nov. . a 9 0 5 : 0 5 227 . M. angustata sp. nov. : 0 é 228 4. M. robinsoni (Rothschild, 1905) See eambmeron : a 230 a. WM. robinsoni robinsoni (Rothschild, 1905) 0 a a 233 b. M. robinsoni peregrinata subsp. nov. . ¢ : : 234 c. M. robinsoni bogora subsp. nov. . ‘ é : 9 235 d. M. robinsoni tiomanica le nov. : : 3 é 236 5. M. phangi sp. nov. 6 0 5 : 7 237 a. M. phangi phangi eubSp. nov. . 240 Host-Relationships and Distribution and Other Comments 240 b. M. phangi tana subsp. nov. . 2 . a a 242 6. M. limi sp. nov. : 242 Host-Relationships and Paseeibation aad Other Gonmenis 244 7. M. thurmani sp. nov. ¢ . : : - . 3 244 8. M. calcavata sp. nov. ¢ c 6 2 c é 247 9. M. batibacula sp. nov. : : é : : o : 250 10. M. veruta sp.nov. . . ° : 252 11. M. loncha (Jordan, 1926) New Combination a 5 : 255 12. M. javana (Jordan, 1933) New Combination ci : 2 256 13. M. rhaeba (Jordan, 1926) New Combination o 257 14. Geographic and Host Distribution of Medwayella gen. nov. in Malaya . 258 15. Key to the Known Species and Subspecies BE Medwayella pen nov. . : 5 0 : ° 9 265 204 ROBERT TRAUB Page B. Lentistivalius gen. nov. , : : : 4 : : 5 269 1. L.vomerus sp.nov. . 272 Host-Relationships al Dt fan and Other Gomnenes 280 2. Key to the Known Species of Lentistivalius gen. nov. . 5 280 C. Stivalius Jordan & Rothschild, 1922, Sensu Strictu ; : 0 281 1. S.cognatus Jordan & Rothschild, 1922 c : é 285 a. S. cognatus cognatus Jordan & Rothschild, ages é 286 b. S. cognatus spivamus Jordan, 1926, New Combination . 287 c. S. cognatus bamus subsp. nov. . é : : ° 287 2. Comments on Stivalius s. str. : 290 Host-Relationships, Distribution ond Potential Vectorship of Disease 6 é 290 The Specialized Aedeagas aad Baa Goatees é c 291 Evolutionary Status of Stivalius s. str. . 292 3. Key to the Known Species and Subspecies of Stivalius Jordan & Rothschild, 1922, s. str. . 0 5 : a 5 : 293 ACKNOWLEDGEMENTS ° ¢ : D 3 ; ° p 204 REFERENCES . F : 6 é é ot Rs " : 2096 List OF HOST-NAMES F 5 : : : 5 208 List OF LANDMARKS AND POINTS OF REFERENCE . c 0 : ; 2098 LisT OF ABBREVIATIONS . c : : ° 5 : S : 300 LIST OF ILLUSTRATIONS). o : . 5 9 : : ° 302 SYNOPSIS Stivalius robinsoni (Rothschild, 1905) and its allies constitute the dominant group of fleas infesting squirrels and tupaiids in the forested foothills of much of the “Indo-Malaysian” region ranging from Indo-China (and perhaps eastern India) through Malaya, to Indonesia and the Philippines. The taxon is reviewed and a new genus is proposed and named to encompass the 4 previously known species and 9 new species. Descriptions, diagnoses and detailed illustrations are provided for these 13 species and 4 new subspecies. The members of the Stiwalius ferinus- group are also discussed and placed in a separate genus, which is named, described and illustrated, along with a new species from North Borneo and some of the species heretofore known. For purposes of comparison, Stivalius Jordan & Rothschild, 1922 s. str. is redescribed and figured, and a new subspecies of S. cognatus Jordan & Rothschild, 1922 is used to illustrate the diagnostic features of the generic level and those for species and subspecies. Notes on the host-relationships and distribution of these fleas are presented, as applicable. Keys to the known species and subspecies are included for all the genera treated. INTRODUCTION THE genus Stivalius Jordan & Rothschild, 1922 (s. Jat.) includes some of the most prevalent and characteristic fleas parasitizing rats, squirrels, tupaiids or other small mammals over much of southeastern Asia and the Indo-Australian Archipelago, and hence throughout this vast area Stivalius s. lat, must be considered as of potential importance regarding vectorship of any infection harbored by rodents and other such mammals. In fact, species of Stivalius have been found infected with plague in India and Java (Pollitzer, 1954), as is pointed out below. The fleas collected by Lord Medway’s team on Gunong Benom in Pahang, Malaya (W. Malaysia) amply indicate the predominance of Stivalius s. lat. in the relatively untouched dipterocarp forests of that country, but the value of the collection far exceeds that point, or that of the new species represented therein. The specimens NEW TAXA OF FLEAS 205 well illustrate certain major points concerning the systematics, zoogeography and evolution of fleas, and contribute to our understanding of the current and past distribution of host-mammals. The findings emphasized the need for the revision of the classification of the St:valius robinsoni-group of fleas, and of certain other mem- bers of Stivalius s. lat. For these reasons, it was decided to regard the Medway collections as the founda- tion and catalyst for a study dealing successively, in three articles, with taxonomy, convergent evolution and zoogeography of certain groups of Stivalius. This first paper in the series includes: (1) the erection of a new genus for the S. robinsoni-group, with descriptions and illustrations of nine new species and four new subspecies, along with keys for the identification of the 13 known species and four subspecies. The hitherto unknown female of S. loncha Jordan, 1926 is described. (2) A discussion of the hosts and distribution of the Malayan peninsular species of the S. robinsoni- group. (3) A new genus for the S. ferinus-group, with descriptions and figures of one new species and a keyto the five known species. (4) A redefinition of Stzvalius s.str., for purposes of comparison with the new genera, and based upon a new subspecies of S. cognatus Jordan & Rothschild, 1922, along with keys to the described forms (four species and three subspecies), including the hitherto unknown male of S. c. spiramus Jordan, 1926. Dorsal or ventral views of the aedeagus are presented to elucidate some of the major points in systematics, comparative morphology or convergent evolution. The second and third articles in the series follow immediately after this one. MATERIALS AND METHODS Methods of Collection and Handling It is emphasized that throughout the course of the field-projects in Malaya, Borneo, Thailand and other countries, which resulted in the specimens treated in this article, the collectors were primarily engaged in other types of studies. The investigators were actually working on scrub typhus, leptospirosis and other infections, or doing research on land-leeches, or engaged in malaria-control, or conducting survey-type operations, and the small number of specimens of fleas obtained at times, and the limitations of the data on infection-rates, reflect the necessarily secondary nature of the collecting-programs. The bulk of the specimens were collected by the writer while serving as Command- ing Officer of the U.S. Army Medical Research Unit (Malaya) during varying periods in Malaya from 1948 to 1959, or by teams of the Colonial Office Medical Research Unit (later Division of Virus Research and Medical Zoology) of the Institute for Medical Research, under the direction of Dr J. R. Audy. Both of these Units were based on the grounds of the Institute for Medical Research (I.M.R.) at Kuala Lumpur and worked closely together. Other important collections were made by the Department of Zoology, University of Malaya (UM), by teams led by Lord Medway or J. R. Bullock. The U.S. Army Medical Research Unit (USAMRU) required live animals for its research on infections, while the British (later Malaysian) Unit (henceforth indicated as DMZ-IMR) was collecting living animals in connection with studies on 206 ROBERT TRAUB ecology. Lord Medway also frequently needed the animals alive for his investiga- tions. For these reasons the vast majority of rats, tree-shrews and ground-squirrels, and even many tree-squirrels, examined for fleas in Malaya had been trapped in various types of live-traps, predominantly in the “‘wire basket- trap,’ a modification of a trap of presumed Chinese origin and commonly used for catching rats throughout southeastern Asia. In this type of device, constructed of thin chicken-wire, the front door is closed by a spring when the mammal tugs at the bait on a hook and thereby releases the mechanism holding the door open. A folding- trap of heavy wire, of U.S. design, was also employed and in this type, the door was released by a treadle on the floor. Closed traps of galvanized metal, operating in a similar way, were also used. Another kind was prepared by fastening a large can to an ordinary large snap-trap and placing a screen on the movable arm of the trap. When released, the arm would hurl the rat into the can and close the aperture. Sundry modifications of these and other live-traps were employed, but regardless of the model, there were always inherent disadvantages insofar as concerns the collecting of fleas. Certain species tend to leave the host soon after feeding to repletion, others do so when the animal becomes excited, and its fur gets ruffled, as happens when the mammal realizes it is trapped. Since the traps were generally examined but once daily (and the animals became agitated anew at the approach of the collector), many fleas were undoubtedly lost before the host could be popped into a cloth bag for subsequent examination. In traps made of mesh, the mammals would become drenched during showers of heavy rain, which usually are a daily occurrence in or near the tropical rain-forests. In traps of solid metal, the host often became too hot, and hyperactive, or died as a result, while at other times, as at night, they became wet and cold with equally disastrous results. These consequences all affected the degree of flea-infestation at the time of collection, as did the attack on the hosts by myriads of ants, both large and small, which was a fairly frequent occurrence. On the other hand, utilization of traps which killed the victims, i.e., snap-traps of sundry sizes, or dead-falls, had other and more serious disadvantages, even though some species of fleas linger for hours on a dead host. Within minutes, dead rats almost invariably attracted swarms of ants, whose movements through the fur unquestionably hastened the departure of many fleas. The removal of innumerable chunks of skin by the ants also would effectively depilate the carcass within an hour or two, and only exceedingly rarely was a flea ever noted on the body of an animal infested by ants. Even when the traps were examined every hour it was impossible to cope with the depredations of the ants in some areas. Nevertheless, the use of snap-traps was a useful adjunct, especially in the mountains, and certain “‘rare”’ fleas were collected by this means. The bait used in the trapping operations consisted of fruit such as bananas or durian, burnt coconut, nuts, peanut butter, sweet potato or other yams, dried fish, bacon, a mixture of oatmeal, peanut butter and bacon, etc. Baits that were sweet (e.g., raisins) or oily or fat tended to attract so many ants that they were soon eaten without trace. NEW TAXA OF FLEAS 207 Trapping was supplemented by shooting, and most of the tree-squirrels and some of the ground-squirrels and Tupaia were collected in this way. Carnivores, in particular, were taken by shooting, usually at night, when their eyes would glow in the light from powerful flashlights. All hosts, including live animals, were immediately placed in individual cloth bags on collection and the bags were sealed pending examination in the laboratory. The bags of dead specimens were treated with chloroform and sprayed with aerosol insecticide to stupefy or kill the fleas; while live animals destined for release after checking were exposed to ether long enough for anesthesia, whereas those required for microbiological study were chloroformed. (Chloroform is more effective but ether is less toxic, hence the use of the latter.) If living chiggers or other ectoparasites were required for inoculation, rearing or study, the host was asphyxiated after exposure to the anesthetic; the host was then brushed vigorously while it was held over a tray, and the stupefied fleas which fell in the process were collected via forceps and preserved in 70% ethyl alcohol. The brushing was repeated several times because of the difficulty in collecting all the fleas hidden in the fur. The fleas were prepared for definitive examination and study by placing them in 10% KOH for 24-48 hours or until the internal tissues had dissolved, after which they were washed in water, dehydrated by exposure to 70%, 95% and 100% ethyl alcohol in series, cleared in oil of wintergreen and mounted in Canada balsam on microscope-slides. In the case of mammals collected by USAMRU, records were kept as to the num- bers of hosts which were not infested with fleas. This was not possible routinely for the collections by our colleagues, but the difficulties mentioned above in collecting fleas in the tropics serve to demonstrate inherent limitations in obtaining infestation- Tates in such regions. When discussing the numbers of fleas taken from various hosts, as in Table 2 (p. 259), the entry ‘minimum number exam’d.”’ therefore refers only to the mammals collected by USAMRU. Major Collection Areas Specimens of fleas of the S. robinsoni-group were never taken in fields of grass or waste land, but only where there was forest, at least in remnant form. Accordingly, the major collecting areas on the Malayan peninsula were in the dipterocarp forest in the lowland or foothills, particularly within 20 miles of Kuala Lumpur, i.e., the Forest Reserves at Gombak, Ulu Langat, Bukit Lagong and Kepong, and also at Klang Gates (Selangor), where trees such as Shorea, Dipterocarpus, Swintonia and Balanocarpus and Hopea dominate the forest. Most of the collections were made between 800 and 2000 ft elevation (mainly at 1500 ft) in areas which were still largely “primary jungle” in 1948-49 but which were thereafter fairly extensively but selec- tively thinned for timber so that by 1956, bamboos and other features of secondary forest were evident in scattered patches throughout. The findings in the dipterocarp forest are treated below as ‘““Kuala Lumpur Forest” (Table 1, p. 216, et seq.). Another well studied site was the Subang Forest Reserve west of Kuala Lumpur, about 8 miles from Batu Tiga on the Batu Tiga-Subang road. Here the forest was of a much more secondary nature and at a lower elevation (at about 200-600 ft) than in the wooded habitats mentioned above. 208 ROBERT TRAUB Field work was undertaken in the limestone hills west of Ipoh, Perak, as a result of an outbreak of scrub typhus in British forces operating deep in the (secondary) forest. This was in the Kledan Saiong Forest Reserve, 8 miles west of Ipoh and again 10 miles northwest of Ipoh, each trip including about 4-5 days intensive collection at eleva- tions of about 800-1100 ft in September—October 1958. The habitat at Gunong Benom, the scene of operations for Lord Medway’s expedition, has been discussed elsewhere in this volume. Limited collections were also made by USAMRU in the coastal or adjacent forests of the East Coast, i.e., at Kuantan, Pahang, and at Bukit Beseiin Trengganu. Another area of dense lowland forest briefly examined was that in Kelantan. Over the years, about one month was spent by USAMRU (and an equivalent period by DMZ-IMR) collecting in the montane forests in the vicinity of Fraser’s Hill (3500-4500 ft elev.), the Cameron Highlands (4500-6600 ft), and about 2-3 weeks at Maxwell's Hill (3900-4200 ft) and Kedah Peak (Gunong Hijau, G. Jerai) (3500— 3900 ft). These areas, in general, have essentially the same mammalian, siphona- pteran and trombiculid fauna and hence the collections are consolidated under the caption “Mountains” in the discussion of host-relationships below (but an exception is noted in the case of Kedah Peak). An interesting set of specimens was taken in Kedah by a group from the Department of Zoology of the University of Malaya (UM) under the direction of J. R. Bullock. This was at the Bukit Wang Forest Reserve at Jibra and the results are treated separately. Other valuable material was collected by UM teams on Tioman Island (Pulau Tioman) off the East Coast of Malaga. Although three joint teams of USAMRU and DMZ-IMR operated on Mt. Kinabalu, North Borneo (Sabah), based at 5000 ft elevation at Tenompak for periods of about 4-6 weeks annually, in 1951-1953, and Lord Medway’s group also collected there subsequently, as have other naturalists interested in ectoparasites, no specimens of the S. robinsoni-group have ever been taken at such altitudes (or higher) in Borneo. However, we did find such fleas on flying-squirrels and in squirrel nests, etc., in the vicinity of Ranau, at the base of this mountain, as indicated in the description of a new species below. Other areas where members of the S. vobinsoni-group were taken, in Sarawak, Thailand and Indonesia, etc., are indicated, along with the collectors, in the descrip- tions that follow. The individuals and institutions participating in all of these studies are mentioned in the paragraph on acknowledgements. Introductory Remarks Concerning the Descriptions Format and Keys The S. vobinsoni-complex of fleasis relatively rich innumbers of species, and some of these are superficially similar in appearance (although differing abundantly in critical details—at least in the males). Moreover, as many as three species have been collected on the same individual ground-squirrel. Accordingly, in order to facilitate identification and to provide an adequate background for the evaluation of dif- ferences at the generic-, species- and subspecies-levels and for use in discussions on evolution and zoogeography, the various species are diagnosed and illustrated in detail. NEW TAXA OF FLEAS 209 Further, the first new species is described in full so as to serve as the basis for com- parison for the forms that follow. Major distinctions alone are therefore stressed in diagnoses, and only significant morphological differences are cited in the descriptions following such analyses. In the keys for identification of species and subspecies, frequently more than one set of diagnostic characters is cited in the couplets. This was done to facilitate the use of the keys, so that if there is indecision about one point, an alternate is available. It is stressed that each feature is diagnostic, i.e., only one need apply, and hence the couplets do not deal with combination of characters, but with alternates. Landmarks and Points of Reference Used in Taxonomy The most important features for classifying the members of the S. robinsoni-group (and other Stivalius s. lat.) are in the male sternum 9g and the aedeagus, and in the internal genitalia of the female, including characters unused or un-noted in the past. These are all labeled and indicated in the text and figured and defined, if new. However, in addition there are morphological structures and “landmarks” that are useful in taxonomy but which are difficult to describe in a few words, or to locate precisely because they are on a curved surface, or else are not oriented in the usual fashion (i.e., the “ventral arm”’ of the male sternum g is often caudal or vertical in position). For clarification and simplification, therefore, a system of reference-points, utiliz- ing lower case letters, has béen employed in some instances (head, claspers, male sternum 8 and sternum g) to designate the specific level or site being described or compared. Claspers, Sterna 8 and g. The need for a system of this type becomes apparent by a glance at figs 28 and 29, the male sterna 8 (8 S.) of two allied fleas. It is obvious that the outlines of the sterna differ markedly, and in the case of the former (fig. 28) the temptation is to say the “dorsal margin is flattened.” However, a question arises as to the “dorsal margin”’ in the latter, for what appear to be the anterior and posterior margins curve together imperceptibly. Moreover, study of the cutaneous marking or striae indicates that the dorsal margin really extends down to the first (uppermost) large bristle on the caudal margin. The structure herein termed the “lJumacaudate process” (LUM.), on 8 S., empha- sizes the problem, for while there are noteworthy differences in proportion, there are inadequate landmarks to use in comparison. Thus, in the case of fig. 29, the length is fairly easy to determine because the dorsal margin is outlined, but this is not so in the other species. Accordingly, the following reference-points have been selected as guide-marks. In the case of the claspers and sternum 9, they are illustrated only in figs 1 and 3 which were specially prepared for this purpose, although the abbreviations are frequently used in the text. It will therefore be necessary to refer to these figures, or to the List of Abbreviations (Landmarks and Points of Reference on p. 298) below, in order to properly utilize this system. However, the text has been so prepared that the sundry structures are generally described in the usual manner and hence are self- explanatory, while the reference-points are cited to serve as supplemental guides. 210 ROBERT TRAUB The exceptions are instances used in comparisons, and these immediately follow such descriptive phrases, and therefore should not be confusing. Chaetotaxy and Dimensions of the Preantennal Region. Members of Stivalius s. lat. are liberally endowed with bristles, and the number and shape of the sundry pre- antennal bristles are often important in taxonomy. Unfortunately, the bristles (except for those in the first row) are usually not arranged in definite rows, unlike some other groups of fleas. Moreover, when relatively large numbers are present, there is some variation in both arrangement and actual number, in accordance with a well known principle in insects. In order to facilitate discussion and indication of the major preantennal bristles, an arbitrary system has been used to designate “tows” of bristles, as shown in fig. 2 (I-IV). No logical pattern could be discerned which would encompass all the large bristles, and the one or more which do not fall in the rowa are indicated as ‘‘supernumerary”’ (SY.). It should be noted that not all the members of ‘‘Stivalius’’ have preantennal chaetotaxy that jibes with even this arbitrary system, but at least it is possible in this way to more readily categorize and denote the “‘exceptions.”’ In the species illustrated in fig. 2, the bases of the anteriormost row of bristles are linked by a cuticular line and hence this row, designated as I, is unmistakable. The next row, of four subvertical large bristles, is Il. The third row, III, distinguished by the eye-bristle (E.B.), which is above and anterior to eye, may be termed the eye- row (E.R.B.). It consists of three sub-horizontal long bristles, and commences in line with the ventralmost of II. Row IV is likewise sub-horizontal and is termed the genal row of bristles (G.R.B.), ranging along ventral margin, from the base of the maxillary lobe (MX.) half-way towards the ventral part of the eye. In addition there is one large bristle out of line, between II and III; it is designated as a super- numerary (SY.). The shape of the head is likewise of significance in the classification of pygiopsyllids, but here, too, difficulties arise. Not only do the sexes differ (as in other fleas), and there also are the usual problems of indicating unmarked points on curved surfaces, but variations occur in the position of certain ‘landmarks,’ depending upon the taxon. Thus, in one new genus herein described, the maximum length of the pre- antennal region is at the level of the median sensillary crater (fig. 2, M.S.C.), but in another, the crater is more dorsal in position, where the frons is narrower, In some instances the labral or anteroventral angle of the head is in line with eye- bristle, in others, not. Those “‘lengths’’ are therefore not comparable. Another problem is that structures that would appear to be useful as landmarks are either movable, and hence subject to individual variability (i.e., the palpi and the maxillary lobe), while others are at times impossible to see clearly (i.e., the bases of the palpi). In order to make valid comparisons, therefore, guide-marks are indicated in fig. 2 as follows: The height of the preantennal regions is measured from the level of the vertex at the falx, viz., }3-i3, to the true ventral margin of the head (f4-g4). The length may be measured at various levels, according to genus and sex, but always should be along lines parallel to the longitudinal axis of the flea, and directed to the margin of the antennal groove, i.e.: (1) immediately below the middle sensory crater (M.S.C.) NEW TAXA OF FLEAS 211 (x3-y3) ; (2) the dorsalmost long bristle of row I (z3-a4) ; (3) 7bid. for row II (a4—b4); (4) the eye-bristle, which at times is in line with the labral angle and then would be d4—e4; and (5) the most anterior point on the arc of the frontal margin, which in fig. 2 is the same as level b4-c4. Other aspects of this system are cited in the List of Landmarks (p. 298). A. Medwayella gen. nov. Diacnosis. ¢ instantly recognizable by the ventral spinose “‘lumacaudate’’ process on sternum 8 (figs 29, 12, LUM.). Shape and structure of ¢ sternum g (figs I, 18, 20) and of sclerites of aedeagal endchamber (figs 10, 22-25) are characteristic and are described below. Spermatheca (fig. 15, SP.) shaped somewhat like a ground-nut or peanut and with basal portion of duct of spermatheca (figs 15, 27, D.SP.) bearing internal sclerotized rings. 2 basal sternum with lateral patch of bristles restricted to proximal portion of segment. 2 tergum 8 bearing a mesal tanned zone (figs 27, 31, M.R.8) anteriorly marked by a vertical ridge and a contigu- ous dark ovoid area. GENERIC DESCRIPTION. Caput integrecipit (figs 4, 9; 6, 9). Anterior margin of head evenly ovate, upper half more rounded than lower. Front part of gj head about thrice as high (from vertex at falx, to ventral margin) as long (at level of dorsalmost bristle of row II, viz., b4—-c4 in fig. 2). Preantennal region with 4 irregular rows of bristles, the lowest ventromarginal. Eye reniform, well developed, inserted far back, above base of procoxa. Antennal segment 2 with bristles short in both sexes. Antennal groove not extending on to propleuron. Postantennal region with 3 complete rows of bristles. Labial palp (L.P.) 5-segmented (excluding palpiger) and generally reaching near apex of procoxa; occasionally longer (fig. 104). Prosternosome lacking distinct sinus for receiving first vinculum (VC.1.). Dorsal margin of pronotum subequal to, or slightly longer than, length of adjacent spines of comb. Mesonotum (fig. 9, MSN.) generally with 1 pseudoseta (PS.S.); subdorsal. Pleural arch (figs 6, 9, PL.A.) well developed. Metanotum (MTN.) lacking apical spinelets. Metacoxa lacking mesal spiniforms, but with a few mesal thin bristles. Profemur lacking a group of mesal bristles but with scattered lateral ones. Meso- and metafemora lacking median and submedian lateral bristles. Dorsolateral bristles in notches on posterior (outer) margin of tibiae largely paired. First metatarsal seg- ment about 1-6 times length of II and nearly 2-5 times that of mesotarsusI. Third metatarsal segment longer than fifth. Except for fringe on first segment of protar- sus, none of tarsal bristles extending well beyond middle of following segment. Tarsal segment V with 6 pairs of stout lateral plantar bristles, of which first pair displaced towards midline on pro- and mesotarsi (fig. 7), and third pair somewhat so displaced on all legs (fig. 8, metatarsus). Fourth vinculum (fig. 9, VC.4) usually upright and not projecting forward as a short rod. Some abdominal terga with 1 subdorsal apical spinelet. Unmodified terga very broad, extending ventrad to lower third or fourth of sterna; with 2 virtually complete rows of bristles in g, 3 in 2 but incomplete. Basal abdominal sternum of 9 with lateral patch of small bristles, but these near anterior margin; nude in g. Fossae of representative abdominal spiracles sagittate. Both sexes with 2 antepygidial bristles per side (figs. 11, 3; Zi) ROBERT TRAUB 26, 9, A.B.). Tergum 7 of 9 with some bristles modified to resemble antepygidials, i.e., I dorsomarginal (U.M.B.), one immediately below plate of A.B. (L.M.B.) and 1 (L.B.M.-2) in ventrocaudal corner of group of bristles of 7 T. Tergum 7 produced into a short lobe (U.L.7) above A.B. plate and, in 9, a longer, pointed lobe (L.L.7) immediately below it. Sensilium (pygidium) higher than long in g (figs 11, 13, SN.) ; not as arched in 9, where dorsal-ventral axis slightly exceeds anterior-posterior axis (fig. 26, SN.). Mate. Tergum 8 (fig. 11, 8 T.) very small, extending only slightly ventrocaudad of its spiracular fossa (8 SPC.). Sternum 8 (8 S. and fig. 28) correspondingly very large, extending dorsad to near base of ventral anal lobe (V.A.L.) and cephalad to about level of A.B.; “lumacaudate”’ in bearing a conspicuous, spinose, mesal ventral bulbous process (LUM. and fig. 12) arising near middle of ventral margin. “‘Luma- caudate process’’ attached to 8 S. by an apical fold or seam (S.LUM.); inner surface armed with close-set spiniforms from near basal fourth or fifth, to apex; spiniforms directed mesad. Ventral anal lobe (fig. 13, V.A.L.) with ventral margin with a conical process at middle; this bearing 1 long apical bristle, hence structure somewhat resembling anal stylet of 9 (fig. 27, A.S.). Subanal sclerite (S.S.) relatively large and well tanned. Manubrium (MB.) broad to near apex; ventral margin with apical third quite straight but usually biconvex near middle; dorsal margin somewhat convex; its bay (B.MB.) extending cephalad to near apex. Immovable process of clasper (figs 14, 17, P.) dorsally somewhat truncate except for caudomarginal bulge around longish apical bristle; with a smaller dorsal bristle. Conical process (C.P.) of P. usually more than 5 times as long (cf. fig. 1, e-h)! as broad at middle (f-g). Mov- able finger (F.) of clasper with stiva (STV.) (distocaudal extension) too short to resemble a plough-handle and base far too broad to look like a plough-shaft (in con- trast to the condition in Stivalius s. str., which suggested to Jordan the coining of the generic name based upon the Latin word, stiva, for plough-handle). Caudal margin of F. usually sigmoid so that F. is narrowed subapically and bulges caudad proxi- mally. Distal fringe (D.FR.) generally of about 4 stout bristles; subapical, at level of sensilla-group (S.G.) of 3 short bristles; the bases of bristles of the fringe mesal and slightly anterior to caudal margin of F. (vide fig. 87). Fulcral sclerite (F.S. and fig. 65) of F. about 1-3 times as long (high) (fig. 1, ww/xx—zz/a3) as broad at expanded base (ww-xx). Tergal apodeme of segment g (T.A.P.9) long and narrow. Proximal arm of sternum 9 (P.A.9 and fig. 63) apically massive, and base rapidly broadening so that subapical breadth (cc-ee) is about 2-5 times that of base (gg—hh). Distal arm of sternum 9 (figs 18, 20, D.A.g) subequal to P.A.g in length (fig. 1, li-qq; li-are aa/bb) and characterized as follows: (1) Fairly long and narrow, some- what broadening at level of transverse sclerotization (T.S.) which represents dorsal limit of internal fusion of the two distal arms. (2) With subapical group of 4-6 caudomarginal (ventromarginal) short spiniforms (S.G.SPN.). (3) Caudomarginal submedian group of 2-3 stout bristles (CM.G.B.) above T.S. (4) A median dense 1 For methods used in measurements, see figs 1-3 and text pp. 209-211. The small letters indicate the points in question; letters connected by a dash denote the two points or axes being measured, while a fraction signifies an imaginary line connecting those points. Thus, ww/xx-zz denotes the distance between zz and an imaginary line connecting ww and xx. NEW TAXA OF FLEAS 213 group of microspinules in clear area near tip of D.A. g herein termed the subapical patch of microspinules (MSP.P.) and presumably sensory in function. (5) With subapical lobate extension (SUB.L.) on anterior (dorsal) margin; this bearing a group of fine bristles near margin. (6) Often with a distinct subapical dorsal (ante- rior) notch (fig. 20, NCH.) or broad sinus (fig. 33, ARC.) resulting in a characteristic apical lobe (AP.L.). If sinus or arc absent, then margin between apex and SUB.L. quite homolate (“‘uniform-sided,” viz., flat and unmodified) (fig. 67, HOM.). (7) With a laterad-directed, lightly tanned, flap herein termed the supramedial flap (SUP.FL.) overlapping more mesal patch of microspinules (MSP.P.) ; its basal margin extending from near level of SUB.L. to, or near, apex of D.A.9, its ventral (caudal) margin delineated by a median arc of thin bristles, the supramedial group (SUP.G.). Base of SUP.FL. associated with a sclerotized thickening (THK.), at least for upper portion. (SUP.FL. appressed to surface of D.A.g in mounted specimens and hence appearing as contiguous with it and not at all flap-like.) (8) With a convex semi- membranous microtufted or microspiculose area in a cavity of anterior (dorsal) margin at level of T.S. herein named the mid-microspiculate area (M.MSP.), and of presumed sensory function. (9) In some species, heavily sclerotized and ventral margin appearing as an apical spur-like extension (fig. 111, EXT.) because of semi- membranous nature of distal portion of dorsal margin. Aedeagal apodeme (fig. 10, AE.A.) resembling manubrium (MB.) for most of its length; lacking apical appendage. Middle lamina (M.LAM.) with its bay (B.M.L.) extending nearly as far cephalad as anterior margin of bay of manubrium (B.MB.) when the two apices are contiguous. With an ovoid, subdorsal cavity in lateral laminae containing a spiculose body, the entire structure herein termed the caverna spiculosa (fig. 22, CAV.SPIC.). Aedeagal pouch (AE.P.) well tanned; thickened ventral walls (figs 23, 42, AE.P.-V.) extending cephalad to level of F.S. and distad to level of base of sclerotized inner tube (S.I.T.), near base of phylax (PHY.); lateral walls (AE.P.-L.) continuing along sides of the crochet process (CR.P.) and blending with it, appearing at least as the basal ventral portion of the ‘‘crochet”’ (fig. 22) and at times as the entire ventral part (fig. 71); the unified structure, whose components are at times difficult to delimit, herein termed the quasi-crochet (Q.C.). _ Hood (HD.) sinuate to ventral margin and then projecting anterodorsad as the lateral ‘‘deltoid flap” (DEL.FL.), which covers much of the endchamber except for base of phylax (PHY.) and base of S.I.T. Deltoid flap dorsocaudally terminating at dorsal apex of body of crochet (B.CR.) ; its own apex subdorsal, at level of aedeagal fulcrum (AE.F.). Lateral lobes (L.L.) reduced; primarily ventral in position and mesad of DEL.FL.; extending from apex of thickened ventral margin of pouch wall over base of phylax of crochet process and here merging with caudolateral region of pouch wall (AE.P.-L.). Sclerotized inner tube (S.I.T.) relatively unmodified, fairly long, horizontal and sinuate and with ventral margin extending somewhat more distad than dorsal margin; distal third ridged, often with a dorsal spur-like projection (figs 23, 96). Ford’s sclerite (F.SC.) (formerly termed apicomedian sclerite) very well developed; apex of its alpha-portion (ALPH.) thumb-like (THM.) or else forming a groove-like structure (figs 71, 115, GRV.) by paralleling part of the thickened basal margin of the securifer (SEC.). Upper arm (U.A.) of securifer narrowed distally ; lower arm (L.A.) broad to 214 ROBERT TRAUB at least middle and then narrowing somewhat. Ford’s sclerite bifid to midline in dorsal or ventral aspect (figs 24, 25). Phylax (PHY.) relatively broad (lateral aspect); about twice as high as broad at base; subligulate, with ventral margin straight; arising near apex of pouch wall at fusion with lateral lobe (L.L.) and extending to apex of pivotal ridge (PIV.R.) of aedeagus. With a short, tanned “pivotal chord” (PIV.CD.) linking apicocaudal angle of PHY. with base of U.A., arising at apex of PIV.R. Crochet boomerang-shaped, its body or basal sclerite (B.CR.) vertical, narrow except for expanded truncate apex which is contiguous with base of Ford’s sclerite (F.SC.) and junction with DEL.FL.; crochet process (CR.P.) extending caudad as a horizontal digitoid arm and ventrally indistinguish- ably fused with AE.P.-L. to form the quasi-crochet (Q.C.). Often with a semimem- branous spiculose lobe (fig. 22, SPIC.L.) (which is probably the ventral lamella of Hopkins & Rothschild 1966), apparently arising from base of phylax and intimately associated with ventral region of Q.C. Aedeagal fulcrum (AE.F.) quite narrow. Cresc- ent sclerite (C.S.) long. Satellite sclerite (SAT.S.) very short. Central sclerite (CEN.S.) very close to fulcral medial lobe (FUL.M.L.), the two together resembling upper portion of an arrow-head. Lateral shafts of capsule (L.S.C.) well tanned. Y-sclerite (Y.S.) large. Vesicle (V.) well developed. Penis rods (P.R.) short and thick. Third apodemal rod (AP.R.) of endophallus nearly as long and stout as penis rods. FEMALE. Spermatheca (fig. 27, SP., and fig. 15) with bulga (B.) about twice as long as broad, medially somewhat constricted dorsally and with basal (caudal) part of bulga slightly broader than apical; hilla (H.) short, basal portion internal and entire length only slightly exceeding maximum girth of bulga; with an apical papilla (PAP.). Duct of spermatheca (D.SP.) with portion near bursa copulatrix greatly dilated (figs 27, 77, DIL.P.) and with long internal slightly sclerotized annulae; middle portion narrower and bearing a ladder-like series of internal sclerotic rings; portion nearest spermatheca still narrower and also with dark annulae. Blind duct of bursa copulatrix inapparent. Bursa copulatrix (B.C.) with its perula (P.B.C.) somewhat ovate (fig. 31) or broadly vermiform (fig. 77). With a large sac (SAC.) associated with DIL.P. and dorsal region of perula. Duct of bursa copulatrix (D.B.C.) lightly tanned and sinuate (degree depending upon relative position). Lura of bursa copulatrix not specialized. With an accessory ‘‘gland’’? or fissure anterior to glandula vaginalis (G.VG.) and at times, one caudad. With a paired, internal, semimembra- nous structure lying between bursa copulatrix and ventral part of sternum 9 (9 S.), immediately above G.VG. and caudoventrally associated with dorsal wall of vagina at genital chamber, and herein termed the paragenital morion (PG.M.). When well developed, PG.M. occupies most of this space (i.e., fig. 77) but it may be greatly reduced (fig. 31). It is of unknown homology and function but may have something to do with the muscles controlling the vaginal aperture and/or with the bursa copulatrix. Sternum 7 (7 S.) with a ventral lobe. Ventral anal lobe (figs 16, 27, 73, V.A.L.) with base short; apical margin long and sinuate, with spaces between groups of long bristles. Dorsal anal lobe (D.A.L.) with 1 long bristle above, and another imme- 2 These “glands,” like the glandula vaginalis and duplicatura vaginalis, presumably house various hooks or lobes of the male in copulation, and are probably not glands in the usual sense of the word, although they may be lined with cells that provide lubrication, etc. NEW TAXA OF FLEAS 215 diately below base of anal stylet (A.S.), which is quite straight, long and narrow. Tergum 8 (fig. 26, 8 T.) with caudal margin bearing a long, shallow sinus extending from near apex to lower fourth; usually with a protruding lobe at ventrocaudal angle; with a pair of mesal, long, thin marginal setae above lobe and another pair of such genitalic bristles at level of V.A.L. With mesal tanned genitalic ridge (M.R. 8) marked by anterior, often sinuate or crescentic, vertical thickening, the dorsal or median portion of which is buttressed by a small ovoid sclerotized area. Eighth spiracular fossa (8 SPC.) relatively small, vertical portion ovate, about twice as high as long. Sternum 8 (figs 27, 32, 8 S.) flask-shaped, with dorsoapical short bristles. The type of the genus is M. dryadosa sp. nov. described and figured below. Here also belong the members of the “‘Stivalius robinsoni-group” of Smit (1958), viz., vobinsoni (Rothschild, 1905), rhaebus Jordan, 1926, lonchus Jordan, 1926, javanus Jordan, 1933 (to use the original spelling, instead of the feminine endings, as hence- forth) and the additional new species described below. CoMMENT. The genus is named for Lord Medway, both for the present holder of the title and for the former, now the Earl of Cranbrook, in token recognition of the many contributions these scientists have made to the study of the natural history of mammals and birds in the Indo-Malaysian area. It is also pertinent and significant that the Earl of Cranbrook’s collections of insectivores in remote regions of north- east Burma and those of his son Gathorne, the current Lord Medway, in Malaya and Borneo, led to the discovery of new and little-known fleas and added considerably to our knowledge of zoogeography. Science is also indebted to these investigators for the encouragement given to other workers to undertake such faunal behavioural and distributional studies. Additional comments on the genus follow the descriptions of new species below. 1. Medwayella dryadosa sp. nov. TYPE MATERIAL. Holotype male (B-45320-1) ex Rhinosciurus laticaudatus; MALAYA: Selangor, Gombak Forest Reserve, 16 mi N of Kuala Lumpur; 300 m elev.; Coll. R. Traub for U.S. Army Medical Research Unit (Malaya); 22.VII.1956. Allotype female (B-47809) ex R. laticaudatus; MALAYA: Selangor, Ampang Reser- voir, Ampang Forest Reserve; 1.V.1958; Coll. R. Traub. The data for 80 g and 51 2 paratypes are summarized in Table 1, which cites the hosts and localities represented. Holotype (U.S.N.M. number 71598), allotype and paratypes deposited in the U.S. National Museum, Washington, D.C. Paratypes deposited in the collections of the British Museum (Natural History), Bernice P. Bishop Museum (Honolulu), Field Museum of Natural History (Chicago), the Canadian National Collection (Ottawa), the Division of Medical Zoology, Institute for Medical Research (Kuala Lumpur), the Parasitological Laboratory (Stavropol, Caucasus, USSR), the Rijks- museum van Natuurlijke Historie (Leiden, The Netherlands), Robert E. Lewis, E. W. Jameson, the author, etc. Diacnosis. Agrees with Medwayella robinsoni in possessing a distinct subapical notch near apex of dorsal (anterior) margin of distal arm of f sternum 9 (figs 20, 21, UWI NUN VAVIVWN ONUW -VSsSNn AWW Aempey a) = VPs UWI S\ 10 pi0T SE PS1 - - - - d1 PE dS PS PI £1 jr PF es e1 = = = pz - - - ae PF - ax £1 - - = Qz a = = z E : : b i ‘aa = = = a = i) & % or Fe = y fz = = = mck a (ea) - PI = = = ca) — — — — — ie} jaa — — =— — — &S Per - - - - &1 PI eye (2k - PI - ge - - - - stag 10 suIe} 4s01044 3s01044 woueg yepey -unojY uPjUrjay yseoD *y suouns 8 “ZL ‘9 ¢ + VAVIVJ{ NI ‘AON ‘dS JDNFHd IDNF Hd wuoyd ‘dW = ‘dW “AW WWI OWN “OYUN VPS Spits Sir Pez Slr P11 d9 P 39 £9 = Se gz dF £9 df PF = 39 PS = J PI dr £9 dS Pr = bz 64 Pz1 S11 Pr = dr PI STITH 4se104 yody sueqns xg oe WWI ates ao i\ -wsa $$ P&E ‘dW dof PPS CW = ‘dW = ‘aw éF ‘dW dz Pik CW = ‘aw of ek GW ~ ‘aw = ‘aw £1 ‘dW = ‘aw = ‘dw or fr ‘aw = ‘dW or P€ ‘dW = ‘dW = ‘aw = ‘dw = ‘aw $1 ‘dW ov Pe 8-aW = ‘dw £1 ‘aw 61 fr ‘dW = ‘aw = ‘dW = ‘aw yseioy “CAIN indun’y jo ereny ‘dds a vsopvtap “Ww = "aw $10}99]]09, [e301 10] SNIABIOINT snqopnvsyv} SNANIISOUIY Y SUUBISUL SNISLADT siuasyna SAMOUaAT sisuasoqol snaMDmMoY snyDY taayjanwm snyoy yoloa snyway is4amog SND AOU DIDGN T sya pwn { rpunpjajz9vM sdoump sinua] SNANIISDPUNS snyojou snan19s0jjv5 ysOP] a Sey EL ei te ‘W Al al = foe) < VITHA VFMAANW INV “AON “dS PSOCVFAUC FTTAZAVMCAW AO SAdALVUVd AO SNOILOATIOD HOA VLVC AO ANVWWAS 216 ‘I alaVy NEW TAXA OF FLEAS 217 NCH.) and thereby separable from other previously described species in which this arm bears a broad sinus instead of a notch, viz., M. javana (fig. 134, ARC.) or else lacks a sinus or notch and instead is merely flattened subapically, viz., M. loncha (fig. 127) and M. rhaeba. Near M. robinsoni but separable as follows: (1) Re distal arm of sternum 9 (fig. 20, M. dryadosa; fig. 21, M. robinsoni; and cf. fig. 1 for abbre- viations of reference points). (a) Apical lobe (fig. 20, AP.L.) (00-qq) above sub- apical notch (NCH.) squared or truncate across entire breadth in new species, instead of AP.L. being somewhat rounded dorsally (pp—qq) and usually slightly concave ventrally as in fig. 21 (oo—pp) (AP.L.), or angled (fig. 54). In M. vobinsont, even if oo-pp is straight and subvertical, pp-qq is convex (fig. 55). (b) Notch (NCH.) symmetrical, shaped like a rounded ““U”’, instead of lower margin (mm-rr) being more oblique and hence with axis of notch (nn-rr) facing more ventrad, and with lower margin (mm-tr) usually distinctly longer than upper (oo-1r). (c) Transverse sclerotization (T.S.) with lower margin quite straight; not sinuate as in fig. 48 (M. vobinsont). (2) Re aedeagus (figs 22, 42, M. dryadosa; figs 23, 44, M. robinsont). (a) Thumb-like distal region (THM.) of alpha-portion (ALPH.) of Ford’s sclerite (F.SC.) with anterior margin ovate and apex subtruncate; about thrice as long (tall) as broad; whereas in M. rvobinsoni “‘thumb”’ (THM.) is more rounded and apex ovate; distal margin about 1-2 times as long as thumb is tall. (b) Lower arm (L.A.) of securifer acuminate subapically, not broad to near apex. (c) Ventral region of phylax (PHY.) scarcely broader than at level near S.I.T. instead of being very much broader. (d) Apex of S.I.T. with ventral margin strongly upcurved and extending well distad of dorsal margin instead of only slightly so. (3) Re g sternum 8. (a) Sternum proportionately taller, viz., only 1-4 times as long (fig. 29, p3-t3) as high (i3-r3) instead of 1-7 times as in M. robinson: (fig. 28). (b) Distance between longest bristle on caudal margin and ventral margin (m3-t3) proportionately greater in M. dryadosa, 1.e., length of sternum at level of that bristle (13-m3) only 2:2 times distance m3-t3, whereas in M. robinsoni itis 2-7 times. (c) Lumacaudate process (figs 20, 29) (LUM.) relatively shorter and broader and apex less ovate, viz., length (q3-s3) along ventral margin only 1-6-1-8 times height at middle (level n3~03), instead of 2-8 times. (4) Re sternum 7 of 9 (7 S.; figs 26, 27, M. dryadosa; fig. 49, M. robinsont). (a) Sinus on caudal margin with upper margin sloping at angle of 50° in new species, instead of usually about 30°; an imaginary line connecting lobes of this sinus there- fore more than thrice height of ventromarginal lobe below sinus, instead of being subequal. (b) Dorsal margin of sinus sinuate ; resulting lower sinus much taller than, but not nearly as long as ventral lobe. This margin evenly convex above sinus in M, robinsont. (c) Dorsal margin of sinus does not parallel ventral margin at all but immediately arches dorsad in new species. In M. robinsoni the sinus is really a notch. (d) Dorsal third or fourth of this segment broader than in M. vobinsoni in that caudal margin here sloping at angle of less than 60° instead of nearly perpendicular. (5) Spermatheca lacking dorsal peak on caudal bulge of bulga (figs 15, 31, B.); in M. robinsont, peak (figs 32, 49, PK.) visible if organ is in proper perspective. (6) Perula of bursa copulatrix (figs 27, 31, P.B.C.) about twice as high as broad instead of three or more times (figs 32, 59, P.B.C.). (7) Mesal tanned genitalic area (figs 26, 31, M.R.8) with ovoid sclerotization near middle of vertical ridge, instead of near upper B 218 ROBERT TRAUB limit of vertical ridge (fig. 32). (8) Lacking characteristic pair of tanned chords (representing ventral portion of paragenital morion) found just caudad of apex of glandula vaginalis (G.VG.) in M. vobinsoni (fig. 32). (9) Lacking sclerotized bead- like thickening(s) representing accessory glands in dorsal wall of apical portion of vagina of M. robinsoni. DescrirTIon. Head. (figs 4, 5; 6, 9). Anterior margin evenly curved from vertex to ventral margin, arc making a broad semi-ellipse. Front portion of head of J, measured horizontally from level of eye-bristle (E.B.) and labral angle (d4—e4 in fig. 2), and vertically from vertex at falx (v3—-w3) to ventral margin (f4-g4) of head, about twice as high as long and thrice as long at level of uppermost bristle of row I (z3-a4); in 9 the corresponding figures are: twice; 2-4 times. With medium and large preantennal bristles arrangeable in 4 rows (the last 3 arbitrary) as shown in fig.2(g). Ind first 2 (I and II) subvertical, of 6 and 4-6 bristles respectively ; last 2 (III and IV) almost horizontal. Row III, commencing with eye-bristle (E.B.), and Row IV (genal row, G.R.B.) with 3 bristles. With 1 median “supernumerary” bristle (SY.) between rows II and III. 9 similar but first (uppermost) and fifth bristles of I small; II with 3-4 bristles and “‘supernumerary” lacking. Eye well developed, reniform, ventrally excised slightly near middle; about 1-4 times as long as high at ventral third; length of eye subequal to that of third segment of maxillary palpus (M.P.). Genal process unspecialized. Maxillary lobe nearly reaching to apex of second segment of labial palpus (L.P.) and to near middle of ultimate segment of M.P. Labial palpus with apical segment nearly 1-5 times length of fourth (penultimate) and extending to near apex of procoxa. Antennal segment 2 with apical bristles scarcely reaching apex of third joint of club. Postantennal region with 3 rows of bristles arranged 5—5(6)—6 in 3; in 2: 5 to 7—5 to 7—5 (6) but with a large gap between bristle near lower corner of occiput and second bristle in row, especially in 9 where space equals length of remainder of row; with an additional long bristle out of line, near antennal groove, below lower placoid. Thorax. Pronotum dorsally slightly longer than adjacent spines of comb (in 2 about 20% longer); with 2 rows of bristles, but first row quite short, generally not reaching middle. Pronotal spines about 20 in number (total); base of comb not reaching to vinculum (VC.2); axis of comb very slightly convex, height of arch at lower fourth; lower 3 of spines shallowly concave dorsally and ventral margins paralleling dorsal for proximal half and then narrowing gradually, apices pointed, but at about angle of 45° and thus spines fairly broad to near tip; fourth spine (from bot- tom) the broadest; about 5-3 times as long as broad at middle; most spines with longitudinal axis somewhat oblique from horizontal, i.e., 4 middle spines, slope about 45° near base. Mesonotum (fig. 9, MSN.) with 6 rows of bristles, including the anteromarginal and submarginal short rows of very small bristles; the rows covering entire length of notum (but flange bare) ; last row consisting of very long bristles (with tiny intercalaries) and this curving ventrally so that lowest bristle appears in line with penultimate row; last 2 rows terminating near level of overlapping dorsum of mese- pisternum (MPS.); second and third rows shorter, and of smaller bristles; with 1 pseudoseta per side and that subdorsal. Mesepisternum (MPS.) with 1 long bristle near ventrocaudal angle and this preceded by an arc of 3 small bristles; with lobe at NEW TAXA OF FLEAS 219 cephaloventral corner sinuate. Mesepimere (MPM.) with bristles arranged 2-3-1; last contiguous with spiracular fossa. Metanotum (MTN.) with 3} rows of bristles, of which the first is the half rowrepresented by2 or 3 members; second extending just below lucodisc. Third vinculum (VC.3) with dorsal surface quite flat except for apical upcurve to point of articulation; axis almost horizontal. Lateral metanotal area (L.M. and fig. 5) with external measurements longer than high at maxima; with internal measurements somewhat so; with r long bristle, flanked by 2 small ones, near dorsocaudal angle. Metepisternum (MTS.) with 1 long bristle in upper rear quadrant, in line with long, narrow squamulum (SQ.), which is thrice as long as broad at middle and is inclined ventrocaudad about 20° from horizontal. Metepimere (MTM.) usually with bristles arranged 4-3-3 in 3, 4-4-3 in 9, excluding small intercalaries in last row, I of which extends above and below row, and which may be double above ventralmost long bristle; with caudal margin evenly biconvex below level of spira- cular fossa, the upper lobe 1-5 times height of lower bulge. Spiracular fossa sagittate but with base broad. Ventral margin quite straight and horizontal, dorsal margin sloping from middle to apex so that both margins parallel for basal half; about 1-8 times as long as broad at maximum level. Legs. Metacoxa (COX. 3) with lateral bristles on apical ? and these submarginal or nearly so; with a group of 8 thin mesal bristles in 2 rows of 4 on ventral fifth, plus a few small, adjacent but proximal, mesal submarginals. With about 9 small, thin lateral non-marginal bristles scattered over profemur (FM.1) in 2 irregular rows along long axis. Meso- and metafemora with a group of small, widely spaced dorsal submarginal bristles along apical third and 3-4 ventromarginal ones along apical fifth, of which distal 2 quite long; with 2 bristles adjacent to spiniform member of apical pair of stout bristles. Protibia and mesotibia with apical group of stout bristles paired; stout dorsomarginals paired except for fourth pair (from apex). Metatibia (fig. 19) similar but dorsomarginal notches much more pronounced and with 1 stout bristle contiguous to apical group (A.G.) of 2. Fringe of bristles on dorsal (caudal) margin of first protarsal segment extending slightly beyond middle of II. Apical caudal bristle of mesotarsus I and II reaching to middle of II and III respectively; that of metatarsus II nearly reaching middle of III. Measurements (in micra) of tibiae and tarsal segments (petiolate base excluded) for holotype: Tarsal Segments Leg Tibia I Il Ill IV Wi Pro- 223 84 84 66 42 IOI Meso- 336 166 105 72 46 113 Meta- 483 407 253 138 76 125 Tarsal segment V with first pair of stout lateral plantar bristles slightly displaced towards midline on protarsus (fig. 7) and mesotarsus; not on metatarsus (fig. 8). Third pair of plantars not nearly as displaced medially on metatarsus V as on protar- sus. With 2 pairs of thin mesal bristles near first lateral plantars on metatarsus V and I such pair by second and fifth lateral plantars; with a stouter, median apical 220 ROBERT TRAUB pair; 2 such pairs subapical (of these, I median); pair of pre-apical plantar bristles near midline; pre-apical plantar hairs well developed. Abdomen. Tergum 1 (fig. 9, 1 T.) shorter than preceding notum with its flange; with 3 rows of bristles, preceded by 1-3 subdorsals. Basal sternum of 2 with a patch of about 14-16 small lateral bristles commencing near vinculum (VC.4) and extending in 2 irregular subvertical rows to near lower third of sternum. Terga 2—5 in g and 2-4 in 9 with 1 subdorsal apical spinelet per side. Unmodified terga in § with first row not reaching level of spiracular fossa and with ventralmost bristle of second row just below fossa. Caudalmost 2 rows in Q similar (i.e., fig. 30, 6 T.); first row ? length of second and preceded by a rudimentary row of 2-4 small bristles. Abdomi- nal spiracular fossae (fig. 11, 7 SPC., 3; fig. 26, ibid., 9) sagittate but narrow, ranging from 2-7 times as long as broad, on tergum 6, to 1-7 times, on 2 T.; at times dorsally sinuate. Typical sterna in g (fig. 11, 7S., 7 T.) with a subventral row of 3-4 bristles of which ventral 2 long and submarginal; these preceded by 6-8 small ventromarginals in 2 irregular rows. In Q (figs 26, 30, 6 S.), these sterna with 3-4 bristles in first row, 5-7 in second and 5-6 in last; at least 4 ventralmost in last row long, as often is ventromarginal of second row. Upper antepygidial bristle in ¢ (fig. 11, A.B.) slightly more than half length of lower; slightly shorter in 9 (fig. 26, A.B.). 7 T. of 2 with modified bristles, resembling A.B., as follows: upper one (U.M.B.) submarginal; not quite as dark, nor as stout nor as long as upper A.B. Dorsalmost of lower modified bristles (L.M.B.) virtually contiguous with plate of A.B. and longer than and as stout as upper A.B. and superficially appearing as an antepygidial bristle; ventral one (L.M.B.-2) similar, but ventrocaudad of L.M.B., separated by a distance equal to height of plate of A.B. Lobe above antepygidial bases (U.L. 7) shorter than antepygidial plate in J; subequal in 9. Lower lobe of 27 T. (L.L. 7) about twice length of A.B. plate; fairly acutely pointed, but margin turning sharply ventrad at level of L.M.B.-2. Modified Abdominal Segments—Male. Tergum 8 (fig. 11, 8 T.) slightly more than twice as high (along anterior margin) as long at level of base of spiracular fossa; caudal margin extending ventrad only as far as level of subanal sclerite; i.e., scarcely below spiracular fossa; anterior margin terminating at level of ventralmost bristle of 7 T.; with 2-5 small subdorsal bristles near fossa. Spiracular fossa 8 (8 SPC.) right- angled; its ventral arm more than 5 times as long (high) (including region below horizontal or oblique portion) as broad at middle; horizontal section twice as long as broad, and twice as long as middle of upright arm is broad, gradually and slightly broadening from apex (ventral) to base; ventral extension with height equivalent to somewhat less than half diameter of horizontal section. Sternum 8 (8 S. and fig. 29) very large; about 1-3 times as long (p3-t3) as broad (high) (q3413) at maxima; extending dorsad above ventral margin of 8 T. and, ventrally, further cephalad than line of antepygidial plate; somewhat trapezoidal, but apparent dorsal margin (h3-3) convex; and then remainder of true dorsal margin sloping ventrocaudad at angle of 45° to dorsocaudal corner at m3; ventral margin slightly concave at caudal third (q3-s3) or half (and thereafter microdenticulate) ; caudal margin (m3-t3) shallowly convex or almost straight: anterior margin (p3-l3) curving dorsocaudad at about angle of 60° and then (13-h3) mainly at 45°. Lumacaudate process (LUM. and fig. NEW TAXA OF FLEAS 221 12) arising at middle portion of ventral margin; with truly spinose area nearly twice as long (ventrally, q3-s3) as high (level of 03) at middle; apically subovate or broadly rounded; length of apical lateral seam or fold (S. LUM.) equal to } of spinose area; ventral spiniforms in general the stoutest; rest of apical and subapical heavier than median and anterodorsals; the bristles usually in 7-8 rows and about 34-42 in number. 8 S. with a curving suture (SU.8) delimiting a more heavily sclerotized mesal portion; the suture arising near dorsal margin of LUM.; curving dorsocephalad and, at dorsal third, recurving towards summit of dorsal convexity of sternum so that less tanned area commences at base of LUM. and narrows gradually. Considering only bristles caudad of suture and above and behind LUM.: 8 S. with about 18-20 bristles in 4 rows of approximately 5-6-5-4 bristles, those of the last row longer than most, and these well separated between dorsalmost (m3), which is longest, subdorsal and caudomarginal, and a submarginal group of 3 at lower third and fourth; upper- most of third row at caudodorsal corner, near m3; uppermost of second row slightly more dorsal than that of third and anterior by a distance of about 4 its length. Remaining bristles as follows: 1 on LUM. median; a close-set group of 3-4 ventro- marginals at base of LUM. ; 3 oblique, irregular rows, sloping ventrocaudad, of 5-5-3, which first is anteriormost ; these rows preceded by about 4 subventral small bristles. Nude area on 8 S. comprising anteroventral fourth and anterodorsal fifth. Immo- vable process (figs 11, 14, P.) with dorsal margin fairly straight or shallowly concave near dorsal bristle; with apical bristle more than twice length of dorsal; with caudal (ventral) margin quite straight to marked sinus at boundary with manubrium (MB.). Conical process (C.P.) of P. about 5 times as long (cf. fig. 1, e-h) as broad at middle (fg); caudal margin fairly straight. Movable finger (F.) with portion distad of P. (s-n) slightly more than twice as long (s-n) as broad at maximum breadth near apex of P. (i-q) and nearly 4 times as long as broad at middle of subapical constriction (j-k); with broadest portion (i-q) about twice breadth of narrowest (j-k). F. with anterior margin quite convex at level of C.P. but soon straightening and then be- coming mildly sinuate subapically for arc at bases of sensilla-group (S.G.) ; posterior margin markedly convex at apex of P. (rq) but sigmoid so that concavity of upper half equals bulge of lower, with maximum depth of trough (k) at apical fourth. Stiva (STV.) short, the distocaudal expansion (measured from k—o, which denotes a projection of caudal margin if there were no stiva, to point p) slightly less than § of j-k and about 4 length of apex of F. (Ip). F. with a sensilla-group (S.G.) stouter but subequal in length to bristles along middle of apical margin. Distal fringe (D.FR.) with lowermost of 4 bristles thinner than its mates; with bases of lower 3 bristles recessed so that they appear somewhat submarginal and, in lower 2 bristles, distance to level where transversed by caudal margin slightly exceeding diameter of bristles at that point. With a widely-spaced group of 3 fairly long, thin bristles near beginning of sinus of caudal margin; with an oblique submedian row of scattered, thin, lateral bristles starting near pair of anterior group of mesal marginals (AN.GP.) above apex of C.P., and continuing distad to level of S.G.; middle portion of these accompanied by a few smaller mesal bristles. Fulcral sclerite (F.S.) shaped like an axe-head with a broad edge; latter (dorsal) (xx—-ww) more than twice breadth of apex (zz-a3). Manubrium (MB.) broad, only slightly narrowed from middle to near apex; nearly 222 ROBERT TRAUB twice as long (y to level of a/b) (from ventral margin of tergal apodeme of segment g, T.AP. 9, to apex) as broad at anterior ventral bulge (at w); apex (y) broad and only slightly upturned ; dorsal margin slightly convex; ventral margin somewhat biconvex near base, proximal bulge (w) near levelof F.S. T.AP.9 2-5-3 times as long (a/c—b/d) as broad near apex (a-c); longitudinal axis apically somewhat curving towards 8 SPC.; somewhat narrowed at middle. Sternum 9g with proximal arm (P.A.g) with margins diverging from base (gg) at trough of “U.” Anterior (ventral) margin (hh— aa) quite straight and oblique (dorsocephalad) to above basal third (dd), and then curving fairly dorsad (dd—aa); upper half shallowly biconcave but with dorsal (anterior) sinus (aa—cc) longer than lower (cc-dd). P.A.g with caudal (dorsal) margin curving dorsocaudad from trough of ““U”’ (gg) but quite flat or mildly sinuate and in essence almost paralleling dd—aa so that upper # of P.A.g (bounded by aa, bb, ff, dd) broadly rhomboidal; and with long axis of rhombus 1-6 x cc-ee, and dimen- sions of arc aa—bb somewhat exceeding those of dd—ff. Distance aa—cc equals cc-ee. Distal arm of sternum 9 (figs 18, 20, D.A. 9) about 6-8 times as long (ii-qq) as broad at level (uu—jj) of transverse sclerotization (T.S.), which is narrow, straight, except for slight convexity on upper (dorsal) half of anterior (proximal) margin. Subapical notch (NCH.) on anterior (dorsal) margin parallel-sided, its depth (nn—-rr) subequal to its height (mm-oo). Apical lobe (AP.L.) somewhat higher (oo—qq) than long (pp to level of rr). Thickening (THK.) at base of supramedial flap (SUP.FL.) well represented along trough of notch and slightly above and below it. Subapical lobe (SUB.L.) below notch extending slightly more dorsad than AP.L. (mm higher than pp); with a small group of thin submarginal bristles below SUB.L. Subapical group of short spiniforms (S.G.SPN.) 4 to 5 in number, with distalmost the smallest and at level of base of NCH. and basalmost at level of SUB.L. Subapical patch of ‘“‘sensory”’ microspicules (MSP.P.) extending from near upper margin of NCH. to supramedial group (SUP.G.) of 5 thin bristles. Caudomarginal submedian group (CM.G.B.) consisting of 3 stout bristles, of which bases of upper 2 frequently overlap; with 2 much smaller, thinner marginal bristles distad of these. Mid-microspiculate area (M.MSP.) elongate-ovate, equivalent to } length of D.A.g and its cavity extending into arm for about } depth of latter, its apex on anterior (dorsal margin) (kk) well separated from proximalmost bristle of group below SUB.L., at point ll. Aedeagus—Lateral Aspect. Aedeagal apodeme (fig. 10, AE.A.) with middle la- mina (M.LAM. and fig. 22) more tanned than lateral laminae (L.LAM.). Middle lamina about 4 times as long (from anterior edge of crescent sclerite, figs 23, 93 (C.S.) at base of aedeagal fulcrum, AE.F. to apex) as broad at anterior margin of bay of middle lamina (B.M.L.) ; anterior 3 gradually narrowing. Apex ofapodeme upturned, and plates fusing to a point just below tip, which is nearly thrice as long as broad. Apodeme somewhat constricted at level of caudal portion of caverna spiculosa (CAV.SPIC.), basad of AE.F. Lateral laminae with ventral and anterior margins primarily paralleling that of M.LAM. Bay of middle lamina (B.M.L.) extending cephalad 2 of length of apodeme (from AE.F.). Median dorsal lobe (M.D.L.) (and fig. 42) straight but oblique at angle of about 15° commencing at constriction of AE.A. to level of middle of sclerotized inner tube (S.I.T.); here becoming markedly rounded but curving ventrad only as far as line of basal portion, at origin of hood NEW TAXA OF FLEAS 223 (HD.). Aedeagal pouch with girdle (G.) well sclerotized but narrow; ventral walls (AE.P.—-V.) more tanned than girdle and lateral portions (AE.P.—L.) continuing to level of base of S.I.T.; ventrocaudal region marked by a sclerotized ovate area extending to near middle of lower margin. Hood (HD.) with a deep sinus on lower half of caudal margin; sinus reaching to near apex of sclerotized inner tube (S.I.T.) and its depth somewhat less than that of resulting lobe beneath it; this ventral lobe subtruncate dorsally and evenly and strongly convex below as margin of hood continues anterodorsad as triangular deltoid flap (DEL.FL.). Margins of DEL.FL. forming angle of 45° at junction above level of AE.F. and apex somewhat extended, acute; dorsal edge of flap blending caudally with upper apex of body of crochet (B.CR.) at base of alpha-portion (ALPH.) of Ford’s sclerite (F.SC.). Ventral margin of DEL.FL. an arc whose trough is in line with bottom of phylax (PHY.) and covers region of ‘“‘crochet.” Lateral lobes (L.L.) extending caudad from ventral margin of AE.P. as a shallow sinuate structure and then arching to blend with lateral walls of aedeagal pouch (AE.P.—L.) and with crochet-process (CR.P.) to form quasi-crochet (Q.C.), all of which are mesal to overlapping deltoid flap (DEL.FL.). Ford’s sclerite (F.SC.) largest of structures in endchamber; its alpha-portion (ALPH.) basally rectangular; its anterodorsal margin fairly straight to curve of thumblike distal section (THM.), which is short, apically truncate; curved axis of thumb about twice as long as tube-like arch is broad. Securifer (SEC.) of F.SC. chelate (in appearance) apically; dorsally biconvex, i.e., base of upper arm (U.A.) commencing as ovate structure arising near thumb of ALPH., becoming concave at caudal fourth and then with free portion of U.A. extending apicad as a narrow arched rod which is almost claw-like or falcate; with proximal axis of free section nearly horizontal and essen- tially an extension of that of main body of upper arm, i.e., not angling dorsally. Lower arm (L.A.) of securifer sinuate dorsally and acuminate at apex as the opposite member of the ‘‘chela.’’ Pivotal ridge (PIV.R.) relatively lightly sclerotized; rod-like to apex at summit of phylax (PHY.) and base of F.SC. With pivotal chord (PIV.CD.) fairly straight close to basal wall of ALPH. and terminating at a root-like extension from base of U.A. Sclerotized inner tube (S.I.T. and fig. 43) lacking distinctive armature; about 9 times as long as broad at straight portion of middle; very broad basally but rapidly narrowing to a constriction at level of anterior margin of PHY., where it bulges dorsally and ventrally (with curved margins); length of dilated portion shorter than diameter; tube only slightly convex to a short dorsal spur at distal fourth and then becoming somewhat upcurved to apex; with a mem- branous connection between the spur and apex of tube; ventral margin extending distad of apex asa short, sinuate rod-like projection ; distal fourth bearing sclerotized Ting-like ridges. Crescent sclerite (C.S.) shallowly convex; about 2-5 times as long as S.I.T. is broad at middle. Satellite sclerite (SAT.S.) very small, scarcely exceeding depth of C.S. Fulcral lateral lobes (FUL.L.L.) narrow; relatively lightly tanned. Central sclerite (CEN.S.) closely approximating fulcral medial lobe (FUL.M.L.). Y-sclerite (Y.S.) conspicuous as a diamond-shaped structure below fulcrum and with a narrow branch (in lateral aspect) extending distad as floor of capsule. Dorsal virga (D.V.) lightly tanned. Lateral shafts of capsule (L.S.C.) well sclerotized from level of C.S. to ventral margin of FUL.L.L., their tendons (T.L.S.) lightly tanned. Phylax 224 ROBERT TRAUB (PHY.) conspicuous; its altitude (vertical length) about twice breadth of base; proximal half only slightly narrowing; curving dorsocaudad at upper level of S.1.T. Body (basal sclerite) of crochet (B.CR.) as long as crochet process (CR.P.); length of expanded apex equivalent to half of height of B.CR., apparently constituting major or entire component of dorsal, longitudinal portion of quasi-crochet (Q.C.) ; this tanned element directed caudad as an arm subequal in girth to that of B.CR.; about 6 times as long as broad at middle; slightly broadened subapically; apex broadly pointed ; dorsal margin almost straight and ventral margin corresponding. Spiculose lobe (SPIC.L.) (apparent homologue of median lamella) extending below Q.C. for distance equal to half breadth of latter. Vesicle (V.) fairly well developed. Penis rods (P.R.) not extending beyond up-curve of AE.A. Apodemal rod of endophallus (third aedeagal rod) (A.P.R.) fairly well tanned to apex, near that of P.R. Ventral virga (V.V.) well developed and tanned far beyond (cephalad) level of girdle (G.). Aedeagus—Dorsal and Ventral Aspects. (fig. 24, dorsal; fig. 25, ventral). Ford’s sclerite (F.SC.) basally fused along midline. Apical half (U.A.) bifid, the cleavage extending far proximad for L.A. and ALPH. Apical lobes of U.A. broadly pointed; rather spatulate; acuminate in L.A. Aedeagal pouch not completely enclosed ven- trally, its thickened ventral walls (AE.P.—V.) V-shaped, with divergence terminating at base of phylax (PHY.) and origin of lateral lobes (L.L.), which distally blend with lateral margins of aedeagal pouch (AE.P.-L.). Wall of aedeagal pouch merging with sides of crochet process (CR.P.) to form quasi-crochet of lateral aspects. Phylax (PHY.) sloping towards midline from base to apex. Spiculose lobe (SPIC.L.) arising at base of phylax and extending caudad to near apex of CR.P. Deltoid flap (DEL. FL.) triangular at caudal end as well as cephalic; covering lateral thirds of dorsal surface from origin at apex of basal sclerites of crochets (B.CR.) to level of fulcrum; likewise ensheathing sides of ventral portion of endchamber as a development of hood (HD.). Ring-like transverse sclerotizations on apical portion of S.I.T. not completely encircling tube, i.e., are ridged arcs. Crescent sclerite (C.S.) narrowed caudally as it approaches midline. Satellite sclerite (SAT.S.) along dorsal midline, immediately distal to apices of crescent sclerites and directed into chamber of sclero- tized inner tube (S.I.T.). Lateral shafts of capsule (L.S.C.) appearing as long, triangular structures between lateral lobes of fulerum (FUL.L.L.) and fulcral median lobes (FUL.M.L.) and extending to near apex of C.S. Tendons of lateral shafts (T.L.S.) arising from their bases and continuing far cephalad. Y-sclerite (Y.S.) serving as floor for capsule, with one fork or fold terminating near apex of C.S.; the other broader, butterfly-shaped, and the length of FUL.L.L. Central sclerite (CEN.S.) a bell-shaped structure superimposed upon FUL.M.L. Ventral virga (V.V.) bifid at apex. Caverna spiculosa (CAV.SPIC.) elongate, subdorsal ovate sinuses, extending from near base of bay of middle lamina cephalad towards base of aedeagal pouch. Female. (figs 26, 27, 30). Sternum 7 (7 S.) about 1-6 times as high as long at maximum diameters; anterior margin nearly vertical save for dorsal slope towards short dorsal margin; caudal margin with upper half shallowly sinuate and lower half with a large sinus whose lower margin slopes ventrocaudad at a 45° angle and upper margin is slightly biconcave (the component arcs subequal in shape and size) and NEW TAXA OF FLEAS 225 slopes dorsocaudad at 40°; upper lobe of sinus broadly rounded; lower lobe truncate and about twice as long as broad at middle. Sternum 7 with 3 rows of bristles following a group of 2-4 small ventromarginals; first irregular and with about 10 small bristles, second with about 9, slightly longer and of these, 2 ventromarginal; third with 6 long bristles of which 1 is submedian, at level of anal stylet (A.S.), 2 near middle of upper lobe of sinus and 3 at base of lower sinus, including 1 ventromarginal and I sub-ventromarginal; of latter 3 on ventral lobe, anteriormost much closer to ventral member of preceding pair than to dorsal one. Tergum 8 (8 T.) with caudal margin shallowly concave to level of A.S., then quite straight and oblique to a fairly narrow lobe at ventrocaudal angle; this lobe about twice as long as broad at middle. 8 T. with an irregular row of 5 short bristles immediately in front of spiracular fossa (8 SPC.) and this preceded by 1-2 similar bristles, at times by a more anterior, smaller, dorsomarginal; vental pair of mesal marginal genitalic bristles immediately above ventrocaudal lobe twice as long as those at base of ventral anal lobe (V.A.L.); with an oblique, horizontal row of 5 bristles near ventral margin, commencing near middle of margin and terminating at level of upper base of lobe at anteroventral angle, but anterior to it; last 3 of this row the longest ; with another 2 similar rows slightly dor- sad to sub-ventromarginal one, but these bristles smaller, although terminal ones get progressively longer; with a fourth such row represented merely by 1-3 bristles dorsal to these. Mesal genitalic ridge of 8 T. (M.R.8) with anterior, vertical margin slightly sinuate or almost straight; with a crescentic (quarter moon) sclerotization buttressing this, commencing at midline and continuing to subdorsal region; below the crescent, an indiscrete or Y-shaped tanned area. Dorsal anal lobe (D.A.L. and fig. 16) with anterior dorsal half and ventral two-thirds nude; remainder of surface clothed with 2 or 3 rows of bristles, of which only last complete and consisting of long bristles, especially the one above, and, particularly that below, base of anal stylet (A.S.); with an additional very long bristle at dorsocaudal angle. Anal stylet broadest near base; quite straight and cylindrical thereafter, but at times slightly narrowed at apical fourth (especially ventrally); about 4 times as long as broad at middle; with a very short dorsal bristle at base of long apical bristles; at times with a tiny ventral 1 there as well. Ventral anal lobe (V.A.L.) with distal margin bearing a pair of bristles at basal angle, viz., 1 large and 1 small; a pair of long ones near basal fifth; a very long one beyond midpoint and an adjacent smaller one (at apical 3) and, at times, a virtually contiguous small one further distad. Spermatheca (SP. and figs 15, 27, 31) shaped like a peanut in which the caudal lobe is foreshortened® and broader than the other, i.e., bulga (B.) dorsally somewhat constricted at basal third; proximal portion somewhat more convex than anterior (but lacking a distinct dorsal peak) and caudal margin somewhat flattened. Bulga about twice as long (from anteroventral angle) as broad at narrowed area; ventral margin a long, shallow arc. Hilla (H.) with nearly half its length internal; slightly more than 4 times as long (total length but excluding apical papilla, PAP.) as broad at point of entry (ignoring thickenings of wall). Apical papilla well developed, 3 It is too bad that there is no such term as ‘‘hind-shortened,” for that would be more pertinent and descriptive. 226 ROBERT TRAUB longer than high. Duct of spermatheca (D.SP. and figs 27, 31) with section entering bursa copulatrix dilated (DIL.P.) as compared to remaining ? of length; sclerotized internal rings best developed in region anterior to dilated section. Bursa copulatrix (B.C.) with perula (P.B.C.) ovate-oblate; anterior margin straight and vertical; caudal margin somewhat convex; twice as high as broad. Sac of bursa copulatrix (SAC.) semimembranous and dilating at level of middle of perula and extending above it for distance of 1-5-2 times breadth of P.B.C. Duct of bursa copulatrix (D.B.C.) below perula biconcave, with the arch usually quite pronounced. Duplica- tura vaginalis (D.VG.) marked by an oblique sclerotization and entering at undifferen- tiated lura of bursa copulatrix. Glandula vaginalis (G.VG.) appearing a short, oblique slit at the termination of the virtually horizontal, somewhat thickened and muscled roof of the vagina (VAG.) near genital chamber. With an accessory “‘gland”’ (fissure) immediately anterior to G.VG. Paragenital morion (PG.M.) reduced (in mounted specimens), usually represented on each side as a narrow, short, constricted horizontal “‘U’’ near P.B.C. and whose dorsal and ventral branches are virtually contiguous and whose open end is caudad, the lumen between branches at times tanned and appearing asa short rod. Occasionally dorsal loop of PG.M. visible as an indiscrete longitudinal, membranous area extending towards ventral portion of sternum g (9 S.). Eighth sternum (8 S.) usually with 1 or 2 dorsal short bristles, a longer dorsal subapical one and a terminal one; apically narrow, viz., portion distad of level of most proximal bristle equal to twice breadth (height) of sternum at this level; longest bristle about 4 times this breadth. ILLUSTRATIONS 4. Head and prothorax (3) 19. Metatibia (3) 5. Lateral metanotal area (3) 20. Distal arm of sternum 9 6. Head (9?) 22. Apical region of aedeagus (lateral aspect) 7. Apical segments of protarsus 24. Apex of aedeagus (dorsal aspect) 8. Segment 5 of ¢ metatarsus 25. Apex of aedeagus (ventral aspect) 9g. Meso- and metathorax 26. Modified abdominal segments (2) 10. Aedeagus 27. Spermatheca, genitalia and anal seg- 11. Modified abdominal segments (3) ments 12. Lumacaudate process 29. Sternum 8 (3) 13. Dorsal and ventral anal lobes (3) 30. Segments 7 (2) 14. Processes of clasper 31. Genitalia (9) 15. Spermatheca 42. Apex of aedeagus 16. Anal lobes and stylet (2) 43. Sclerotized inner tube 18. Clasper and sternum 9 93. Apical region of aedeagus CoMMENT. It is noteworthy that although the great majority of the long series of M. dryadosa were from squirrels, all of the actual hosts were species that spend all or most of the time on the ground, i.e., there were no true tree-squirrels represented. Thus, out of 1421 squirrels (excluding flying-squirrels) and tupaiids examined per- sonally by the writer or the staff of the U.S. Army Medical Research Unit (Malaya), 149 were ground-squirrels of the genera Laviscus (52 specimens examined), Rhino- scturus (23) and Dremomys (74), as shown in Table 2 (p. 259), which summarizes the host-relationships of 3 Malayan Medwayella. These rodents nest amongst the roots of trees, etc., rather than high on the branches of the tall trees characteristic of the NEW TAXA OF FLEAS 227 Malayan forest. The other hosts occasionally reported were either ground-dwelling rats like Rattus rajah-group or R. mueller or, like Tupaia (237 examined), were abundant on the ground-surface, although scansorial. In contrast, although 624 truly arboreal squirrels, such as Callosciurus nigrovittatus (110), C. notatus (259), C. caniceps (184), C. prevosti (18) and Sundasciurus hippurus were carefully checked for fleas, no M. dryadosa were ever found on them even though they inhabited the same localities and habitats as the ground-squirrels, and even though they at times carried other species of fleas. Even the 348 specimens of the small species of Swndasciurus (S. tenuis and S. low?) were found to be uninfested with this new species of Medwayella, despite the fact that these squirrels often descend to the forest floor (but nest in trees). It also is significant that M. robinsoni, in contrast, could be found on any of these hosts, regardless of where they nested. The name dryadosa reflects the host-relationships of this new species and derives from the Greek words drys or dryas for tree (or even from dryados, meaning tree- inhabitant) and osoy, or hater. The reason for the “‘antipathy’’ of M. dryadosa for tree-squirrels is unknown, but it seems likely that two factors are involved, viz.: (1) Conditions in the arboreal nests of these squirrels may not be suitable for the immature stages of the flea. (2) Tree-squirrels do not spend enough time on the ground to acquire significant numbers of “‘strays’’ of M. dryadosa. Data concerning the 4 other species of Medwayella known to occur in Malaya (new species, described below) lend support to this hypothesis. Thus, all of these parasitize ground-squirrels and 2 of them have virtually never been taken on truly arboreal squirrels like Callosciurus. It is odd that while the fleas of ground-squirrels rarely, if ever, infest tree-squirrels, a species of flea (M. robinsont) relatively common on the arboreal mammals should be reported from all the ground-dwelling species, and, indeed, at times be as prevalent on these hosts as are their “‘characteristic’”’ fleas. This point is discussed further below (p. 233). Among the ground-squirrels, Rhinosciurus laticaudatus was the main host for M. dryadosa, accounting for 65% of the records even though fewer Rhinosciurus were examined than any other species of small mammals listed (except for C. prevostt). Further consideration of the host-relationships is deferred until the end of the descriptive portion dealing with Medwayella in this paper (p. 258). M. dryadosa was collected in all the non-coastal forested areas surveyed in Malaya by USAMRU, by the DMZ-IMR and by the University of Malaya, provided ground- squirrels were examined in sufficient numbers. The records include areas as far north as Kelantan and Gunong Benom. Four specimens (3 from Lariscus) were taken on Tioman Island, off the southeast coast of Malaya, suggesting that the species probably ranges to the forests near the east coast (where inadequate collec- tions have been made). Altitudinally, the species is known to range from the lowland dipterocarp forest of Subang (800 ft) and Gunong Benom (700 ft) to montane forest near the summit of Mt. Brinchang at 6600 ft elevation. 2. Medwayella arcuata sp. nov. TYPE MATERIAL. Holotype g ex Tupaia sp.; INDONESIA: Sumatra, Suban Ajam, Redjang, Bengkoelen (Benkoelen or Bengkulu) ; July, 1916; Coll. E. Jacobson 228 ROBERT TRAUB (in the mountains 20 km from Benkoelen town, approximately 3° 15'S, 102° 25’ E, fide F. G. A. M. Smit, in litt.). No other specimen known. Holotype deposited in Rijksmuseum van Natuurlijke Historie (Leiden). DiacGnosis. Separable from all described species except M. javana in that dorsal margin of distal arm of sternum 9g (fig. 33) is deeply arcuate subapically and hence lacking the true notch (NCH.) of M. dryadosa (fig. 20) and M. robinsoni (fig. 21) or flattened apicodorsal margin of M. loncha (figs 3, e3-qq; 127) and the new species next described (fig. 68, HOM.) and their allies. Near M. javana but separable as follows: (1) Movable finger (fig. 34, F.) much broader throughout, viz., portion distad of apex of immovable process P. about twice as long (cf. fig. 1, r-n)* as F. is broad at level of apex of conical process of P. (C.P.) (i-q) instead of about 4 times as in M. yavana (fig. 133, F.) while length (rn) is about 4 times breadth at level of middle of subapical constriction (j-k), not 5 times. (2) Due to curve of arc of bases of bristles of distal fringe (D.FR.), distance from uppermost bristle to apex of F. equals approxi- mately } of that to most dorsal part of curve at group of sensilla (S.G.) (1), instead of being subequal. (3) Subapical dorsal sinus (fig. 33, ARC.) deeply and almost evenly arcuate, nearly thrice as long (chord between tips of SUB.L. and AP.L., e.g., mm—pp) as deep at trough, not 5 times because of shallow depth (fig. 134, ARC.), nor with apical lobe (AP.L.) extending more dorsad than subapical lobe (SUB.L.). (4) Lower arm of securifer (fig. 98, L.A.) with caudal margin slightly sinuate and subacute at ventrocaudal angle, instead of being oblique and truncate (fig.95). (5) Tanned por- tion of quasi-crochet (Q.C.) tantamount to crochet-process (CR.P.), longer and Narrower in new species, 1.e., nearly 4 times as long as broad at middle, instead of 2-5 times. DESCRIPTIVE NOTES. Otherwise essentially as in M. dvyadosa, except as follows: Distal arm of sternum g with bristles between those of caudomedial group (fig. 33, CM.G.B.) and subapical group of spiniforms (S.G.SPN.) quite stout. Thumb (fig. 98, THM.) of Ford’s sclerite resembling M. vobinsoni in being short and curved. Dorsal margin of upper arm of securifer (U.A.) quite sinuate, the resulting anterior lobe subequal to length and curve of claw-like extension of U.A. Base of phylax with a pronounced caudal bulge. ILLUSTRATIONS 33. Distal arm of sternum 9 34. Processes of clasper 98. Apical region of aedeagus CoMMENT. The name of the species is based upon the characteristic shape of the upper, anterior portion of the distal arm of sternum 9g and is the Latin adjective aycuata meaning bowed or curved. 3. Medwayella angustata sp. nov. TYPE MATERIAL. Holotype g and allotype 9 ex “Sciurus nigrovittatus and Lariscus niobe”’ (= Callosciurus nigrovittatus and L. insignis niobe); INDONESIA: 4 Compare with fig. for abbreviations of reference points used in measurements. NEW TAXA OF FLEAS 229 W. Sumatra: Sipora Island; Nov. 1924; Coll. C. Boden Kloss. Paratypes: 1 g, ibid. ; Ig, I Q, ibid., but ex C. nigrovittatus (all from N. C. Rothschild Collection at British Museum (Natural History)). Holotype, allotype and 1 g paratype deposited in collection at Tring. 1 pair of paratypes in collection of R. Traub, with ultimate deposition in the U.S. National Museum. Diacnosis. Near M. dryadosa regarding truncate margin of apical lobe (fig. 36, AP.L.) of distal arm of sternum 9, etc., but separable as follows: (1) Alpha-portion (fig. 45, ALPH.) not terminating in a “‘thumb”’ (cf. figs 22, 42, 93, THM.) but, instead, with tanned portion apically rod-like and with a semi-membranous truncate exten- sion joining base of upper arm (U.A.). In M. dvyadosa, ALPH. terminates in a sub- quadrate thumb-like process thrice as long (tall) as broad. (2) Securifer with base of upper arm (U.A.) less than 1-5 times length of extended snout-like or digitoid por- tion, instead of being more than twice its length (measured from edge of internal vertical root. (3) Distal 2? of extended portion of lower arm (L.A.) acuminate, fairly straight ; instead of with subapical portion narrowed significantly, and this portion up-curved. (4) Phylax (PHY.) relatively broad, i.e., only about 1-6 times as long as broad at base instead of twice as long. (5) Base of PHY. with caudal margin more convex than anterior margin which is quite straight ventrally; instead of vice versa. (6) Tanned portion of quasi-crochet (Q.C.) relatively broader, i.e., length (measured from caudal margin of phylax) only about 4-4 times girth at middle instead of 5 times. (7) Anterior ventral bulge of manubrium (cf. fig. 1, w) much more pro- nounced than bulge (u) caudad of sinus (v) instead of both being subequal and shallow. (8) Distal arm of sternum g (fig. 36, D.A.g) with notch (NCH.) asymmetri- cal in that lower margin (mm-—nn) is longer than upper one (nn—o0), instead of notch being symmetrical, the upper and lower margins subequal (fig. 20). (9) Notch comparatively shallow, its depth (measured along edge of apical lobe, AP.L.) at middle (nn-rr) hardly more than } breadth of AP.L.; in M. dryadosa it is more than 3. (10) Upper margin of AP.L. convex before truncate apex instead of being quite flat to squared apex. (11) Female sternum 7 (figs 37, 38, 41, 7 S.) with dorsal margin of subventral sinus paralleling ventral margin for at least basal half instead of diverging almost immediately (figs 26, 30, 31). (12) With a well developed vaginal “‘gland”’ caudad to glandula vaginalis (G.VG.). This is lacking in M. dryadosa. (13) Para- genital morion weakly tanned, represented only by two pairs of short sclerotized parallel lines; one set at apex of G.VG. and one somewhat more anterior (fig. 41); lacking the longer rod-like sclerotization near perula of M. dryadosa (fig. 31, PG.M.). Also resembling M. vobinsoni but readily distinguishable as follows: (1) Alpha- portion (fig. 45, ALPH.) not terminating in a ‘‘thumb’”’ (cf. fig. 23, THM.) but instead with tanned portion narrowing apically, rod-like. (2) Snout of upper arm (U.A.) relatively long and narrow, fairly straight or slightly bowed, i.e., digitoid, but very gradually narrowing from base to apex; about 5 times as long as broad at middle instead of being only 2-5-3-5 times as long as broad, more rapidly narrowing from base, and distally fairly broad. (3) Tanned dorsal margin of basal 3 of U.A. (the non-digitoid portion) of almost uniform breadth throughout, instead of being much broader proximally than distally. (4) Distal arm of sternum g (fig. 36, D.A.g) with apical lobe (AP.L.) truncate nearly its entire breadth, with only upper corner rounded 230 ROBERT TRAUB instead of with upper } or 4 convex (figs 21, 47, cf. also fig. I, pp—qq) or at least oblique (fig. 54). (5) Notch (NCH.) in D.A.g9 relatively shallow; its depth measured from level of inner edge of AP.L. (nn-rr) is only } breadth of AP.L., instead of exceeding 3 of its breadth. (6) Male 8 S. with apparent dorsal margin much longer than in M. robinson, i.e., not curving appreciably ventrocaudad until well beyond apex of immovable process P. instead of curving well proximad of apex of P. (fig. 28). (7) Lumacaudate process (fig. 39) relatively broader, i.e., the portion distad of inter- nal seam (S.LUM.) only about 67-75 % of breadth of LUM., instead of being subequal to it (fig. 28, LUM.). (8) Anterior (ventral margin) of proximal arm of g S. with apical sinus (aa—cc) well developed and ventral one (cc—dd) inapparent, while both are visible in M. rvobinsoni (figs I, 47) but aa—cc is hardly larger than cc—dd. (g) Sinus in 27S. fairly symmetrical, i.e., dorsal margin only slightly diverging from degree of curve of ventral margin (figs 37, 41) instead of clearly doing so at apical half or two- thirds (fig. 49). (zo) With a relatively tall and distinct “‘gland’”’ at dorsal wall of vagina caudad of glandula vaginalis (G.VG.) (although smaller in paratype than in figure 41, which is of allotype). In M. vobinsoni there is a small, bead-like gland here. (11) Mesal genital ridge of 2 8 T. (figs 37, 38, 41, M.R. 8) with dorsal, horizontal tanned area convex ventrally, not sinuate (figs 32, 49) and ventral tail-like portion shorter and stouter, about 1-5 times as long as broad, not 3-5 times. (12) Lobe at ventrocaudal angle of 8 T. ovate, not acute. DESCRIPTIVE NOTES. Essentially like M. dryadosa except as indicated above, and as follows. Tergal apodeme of segment 9g with ventral margin evenly convex instead of sinuate. Tergum 7 of 9 with caudal margin relatively flat, not curving caudad to or beyond level of apex of lobe (L.L.7) below antepygidial bristles. Ventral margin of spiracular fossa 8 of 9 straight. ILLUSTRATIONS 35. Processes of clasper 39. Lumacaudate process 36. Distal arm of sternum 9 41. Genitalia (9) 37. Modified abdominal segments (?) 45. Apex of aedeagus 38. Spermatheca, genitalia and anal segments CoMMENT. The name angustata is derived from the Latin term for narrow, and refers to the appearance of the apex of the alpha-portion and of the upper arm of the securifer. 4. Medwayella robinsoni (Rothschild, 1905) NEW COMBINATION Ceratophyllus vobinsont Rothschild, 1905, Novit. zool. 12:483, fig. 6. Pygiopsylla vobinsoni, Rothschild, 1906, Ent. mon. Mag. (2) 17:221; Rothschild, 1908, Proc. zool. Soc. Lond. 50:617; Oudemans, 1909, Notes Leyden Mus. 31:195, fig. 1; Rothschild, rgrg9, J. Fed. Malay States Mus. 8(3):5, pl. 2, figs 4, 5; Jordan & Rothschild, 1922, Ectoparasites, 1:230, figs 223, 224. Stivalius vobinsoni, Jordan & Rothschild, 1922, Ectoparasites, 1:259, figs 223, 224; Costa Lima & Hathaway, 1946, Pulgas: 324; Traub, 1950, Proc. ent. Soc. Wash. 52(3):121; Traub, 1951, Proc. biol. Soc. Wash. 64:11, pls. 2, 4; Smit, 1958, Bull. Brit. Mus. (Nat. Hist.) Ent. 7(2):41. Diacnosis. The features cited in the comparison of M. dryadosa with M. robinsont serve as a diagnosis, but since M. vobinsoni is a widespread species, ranging at least as NEW TAXA OF FLEAS 231 far as Borneo and Indonesia, and subspecies exist, salient characters at the species- level are now cited. (1) Thumb (figs 23, 44, THM.) of alpha-portion (fig. 46, ALPH.) of Ford’s sclerite short and in the ‘“‘cocked”’ position; apically flattened and with rounded corners; somewhat longer (broader) at apex than stem is tall; anterior Margin convex, posterior margin concave. (2) Upper arm (U.A.) of securifer with base quite flat dorsally and slightly exceeding THM. in height; apex digitoid but broader at base and slightly arched. (3) Lower arm (L.A.) almost as high as long at base; caudal margin oblique (75°) and quite straight; with a short, narrow lobe at ventrocaudal angle. (4) Phylax (PHY.) about 1-7 times as high as broad at base and somewhat expanded subventrally. (5) Tanned portion of quasi-crochet (Q.C.) straight ; at least thrice as long as broad at middle; apical margin angled subdorsally. (6) Sclerotized inner tube long and narrow; broadest near base and then slightly arched for 4 of length; straight thereafter; with long, oblique dorsal spur. (7) Hood (HD.) with sinus fairly deep, extending to apex of S.I.T. (8) Distal arm of sternum g (figs 21, 53-55) with conspicuous notch (NCH.) which is subequal—lower margin (mm-tr in fig. I, g.v.) longer than upper (00-11), and usually conspicuously so. (9) Apical lobe (AP.L.) of D.A.g with margin (qq-oo) sinuate; upper half (qq—pp) convex; lower half (pp—oo) straight or slightly concave. (10) Subapical group of spiniforms (S.G.SPN.) about 4 in number, short and fairly evenly spaced. (11) Transverse sclerotization (fig. 48, T.S.) somewhat sinuate. (12) D.A.g lacking a ventro-apical rod-like extension (unlike M. loncha and allies, figs 3, 111, EXT.9). (13) Proximal arm of sternum g (P.A.9 and fig. 63) with upper half (aa—dd) of anterior (ventral) margin shallowly sinuate; upper concavity (aa—cc) taller than lower (cc—dd). (14) Posterior (dorsal) margin of P.A.g with upright part (ff-bb) quite straight. (15) Movable finger (figs 17, 47, F.) fairly broad; region (n-s) distad of immovable process P. slightly less than 4 times as long as broad (j-k) at subapical constriction and only scarcely more than twice as long (n-s) as broad at maximum dimensions (i-q) near level of apex of conical process (C.P.). (16) Stiva short and bluntly ovate; apex of F. (l-p) about 2-2 times as long as stiva (p-o/k). (17) Sternum 8 (fig. 28) with apparent dorsal margin (h3-j3) relatively flat and very short due to slope of anterior and “posterior’’ margins (but actually extending to ventral third—to m3), region h3-j3 only about } length of ventral margin. (18) 8 S. relatively long, its ventral length (p3-t3) about 1-5 times maximum height (i3-r3). (19) Lumacaudate process (LUM.) fairly long and narrow, nearly thrice as long ventrally (q3-s3) as broad at level of 03. (20) Female sternum 7 (figs 32, 49, 7 S.) with subventral sinus with dorsal margin usually paralleling ventral margin for about 4-4 length of sinus, notch therefore generally fairly symmetrical basally. (21) Lobe below 7 S. slightly broader than sinus, of same length, and subtruncate. (22) Spermatheca (figs 32, 52) with bulga (B.) about twice as long as broad in somewhat constricted middle; with a short dorsal peak (PK.) on caudal bulge; inner dorsal margin flat on apex of anterior bulge. (23) Hilla (H.) with anterior margin entering into bulga for nearly } its length, of uniform diameter except for distinct apical papilla. (24) Perula of bursa copulatrix (P.B.C.) narrowly ovate but for flattened anterior margin. (25) Sclero- tized sectors of paragenital morion appearing (per side) as a single long rod above glandula vaginalis (G.VG.) and a pair of short rods near top of G.VG. (26) With an 232 ROBERT TRAUB accessory gland accompanying well developed G.VG. and nearly as large; with 2 or 3 characteristic bead-like glands lining upper wall of vagina near genital chamber, viz.: I or 2near G.VG. and, at times, I at apical fourth. (27) Mesal genitalic ridge of 8 S. (M.R.8) resembling a “‘T” on edge, and with lower arm quite long and narrow; darkest at and above cross-bar, which is curved and sinuate. (28) Third vinculum (fig. 50, VC.3) with dorsal surface quite even. (29) Squamulum (SQ.) parallel to long axis of body; about 3-2 times as long as broad at middle. CoMMENT. WM. robinsoni was described from a single female specimen collected from Callosciurus nigrovittatus in southern Thailand at Nawng Chik, in Pattani, one of the regions formerly called a “‘Siamese Malay State,’”’ and at 6° 53’ N, ror° 15’ E (fide F. G. A. M. Smit, in litt.). In 1919 Rothschild described the male “‘Pygiopsylla vobinsont”’ on the bases of a pair from Tupaia tana, from Korinchi, West Sumatra, apparently overlooking the treatment and illustration of the male by Oudemans, Igog, which referred to two pairs from a nest of Callosciwrus notatus in Batavia. Rothschild also alluded to a pair from Java, also from C. notatus. In 1922, Jordan & Rothschild mentioned additional Sumatran records and illustrated variations in the spermatheca. Later in 1922 these authors summed up the records, including Oudeman’s, and pointed out the “g is easily recognized by the spinulose process which lies on each side beneath the ninth segment.’’ This presumably was con- sidered so diagnostic that no further description was given at the time, nor was the male ever adequately figured by these workers. This is unfortunate, since the Indonesian specimens represent different subspecies from the mainland robinsont, as is shown below. It should be noted, however, that Jordan & Rothschild were not referring to the specimen herein described as aycuata when mentioning material from Korinchi, Sumatra, even though the slide of the unique holotype had been identified as vobinsoni at the Rothschild Museum at Tring. The Korinchi specimens from “Benkulen, Sumatra,’ mentioned in 1922 were from “Padang Bovenlanden,”’ not from the type locality of avcwata, although they were likewise collected by E. Jacob- son in July, 1916. It seems to me that the omission of this specimen from the cited records was deliberate and that Jordan & Rothschild realized that it was not really vobinsont. M. robinsoni has not been fully described or illustrated to date, although Traub (1951) figured the spermatheca and aedeagus and referred to it when diagnosing another species of Stivalius s. lat. The species is somewhat variable throughout its range in the Malayan peninsula, although the differences are sufficiently minor to be considered as individual or artificial, e.g., due to the position of the spermatheca or the movable finger, etc., in the mounted specimen. There seems to be no valid evidence of subspeciation in the material extant, ranging from northern Malaya to Johore. The type specimen of robinsont, unfortunately a female, is indistinguishable from material from Selangor. For these reasons, I feel that the Malayan slides represent the nominate form even though males from the ““Thai Malay States” are unknown. Further, zoogeographic data also indicate that the major faunal differences between Thailand and Malaya, at least insofar as concerns rodents and fleas, occur substantially further north. More- over, certain Malayan rat-fleas, such as S. klossi (Jordan & Rothschild, 1922) and NEW TAXA OF FLEAS 233 Neopsylla dispar Jordan, 1932, range much further north without apparent subspe- ciation (although distinct subspecies of S. klosst occur in the Indo-China region and in southern China). However, there are significant differences between males from peninsular M. robinsoni and those on Java, Sarawak and even Tioman Island, off the east coast of Malaya, and such subspecies are described below. There is insufficient material at hand to treat Sumatran forms, although I believe that at least one subspecies occurs on that island as well. New subspecies are described after the diagnosis of the nominate form. 4a. Medwayella robinsoni robinsoni (Rothschild, 1905) DiacGnosis. (1) Thumb (figs 23, 44, THM.) of alpha-portion (ALPH.) with apical margin only slightly longer than height of structure. (2) Upper arm (U.A.) of securifer with free ‘‘snout’’ about 3-3 times as long (measured from vertical internal root) as broad at middle. (3) Lower.arm (L.A.) relatively broad; about 1-2 times as long (measured from level of apex of pivotal chord, PIV.CD.) as broad at middle; dorsal margin plunging ventrocaudad at basal 4 or } at an angle of about 60°, but margin mildly sinuate and distally slight recurved, apex blunt, not acuminate. (4) Quasi-crochet (Q.C.) with tanned portion about 4 times as long as broad. (5) Distal arm of sternum 9 (figs 1, 47, D.A.g; fig. 21) with apical lobe (AP.L.) fairly deeply sinuate, its upper half (qq—pp) quite convex; lower lobe (pp-oo) somewhat sinuate. (6) Notch (NCH.) of D.A.g asymmetrical, lower margin (mm—nn) longer than upper margin (oo-nn); axis of notch (nn-rr) sloping ventrocephalad. (7) Female sternum 7 (fig. 49) with dorsal lobe (above ventral sinus) flattened or very shallowly concave, but lacking distinct sinus. (8) Ventral lobe of 7 S. with caudal (single) bristle usually equidistant from pair immediately cephalad, or with caudal bristle at most only somewhat closer to ventral member than dorsal member. (9) Dorsal wall of vagina (fig. 32, VAG.) caudad to glandula vaginalis (G.VG.) with 1 or 2 small knot-like or bead-like glands, 1 at about proximal } and second usually near midpoint ; wall fairly well tanned to near apex, at genital aperture. ILLUSTRATIONS 1. Clasper and sternum 9 46. Apical region of aedeagus 2. Preantennal region (3) 47. Clasper and sternum 9 17. Processes of clasper 48. Transverse suture, D.A.9 21. Distal arm of sternum 9 49. Sternum 7 (2) 23. Apical region of aedeagus 50. Lateral metanotal area (3) 28. Sternum 8 (3) 51. Anal segments (9) 32. Internal genitalia (9) 52. Spermatheca 44. Apex of aedeagus 59. Genitalia and anal lobes (9) Comment. As shown in Table 3, the nominate, mainland form of M. robinsoni may be found on virtually any kind of squirrel, rat or tree-shrew in its forest-habitat. The majority of the specimens are from squirrels—Callosciurus or Sundasciurus, while Tupaia was also commonly infested. In general, then, the bulk of the specimens came from mammals that spend much or most of their time in trees, rather than from ground-squirrels. As indicated above, M. 7. robinsoni is broadly distributed c 234 ROBERT TRAUB throughout Malaya, and we collected it in all surveyed forested areas which were not coastal, ranging from elevations as low as roo ft up to as high as 6000 ft. The species was rare above 5000 ft and I believe that the existing records represent recent intro- ductions of C. caniceps into regions cleared of forest by man, as at the top of Mt. Brinchang or along roadsides to such high elevations. Thus, we failed to collect either C. caniceps or M. r. robinsoni on Mt. Brinchang in 1948 and 1950, before a road was cut to the summit and a radio station constructed there. When we next collected there, in the period 1955 and thereafter, C. caniceps was fairly common in secondary forest and on the fringes. However, more data are required to clarity this point. It is worthy of note that M. vobinsoni was never collected in the coastal areas of Malaya nor on the islands off the west coast, i.e., Pangkor, Sembilan I. and Langkawi I.—and neither was any other species of indigenous flea. Over 50 specimens of Callosciurus and hundreds of rats were examined from the coastal mainland or Langkawi (no squirrels were found on Pangkor or Sembilan I.). The only squirrel collected on Langkawi was C. caniceps, and specimens were shot at elevations as high at 2000 ft, and no fleas were ever noted. However, at such altitudes, the samples checked were too few to warrant conclusions on the presence or absence of Medwayella and other fleas. These points are discussed further, below, after the description of new species, etc. 4b. Medwayella robinsoni peregrinata subsp. nov. TYPE MATERIAL. Holotype ¢ (B-48232) ex Callosciurus notatus; EAST MALAY- SIA: W. Sarawak, Pueh; 3.VI.1958; Coll. T. C. Maa. Allotype 2 (B-49372) ex C. notatus; Sarawak, (3rd Division), Bruit Isle, Rajang Delta; 22.VII.1958; Coll. T. C. Maa. Paratypes as follows: 1 g, 1 2 with same data as allotype; 2 g ex C. notatus; S.W. Sarawak, Tapuh; 9.VII.1958; Coll. T. C. Maa. Holotype, allotype in U.S. National Museum, with holotype bearing number U.S.N.M. number 71599. Paratypes in collection of B. P. Bishop Museum (Honolulu) and that of author. Diacnosis. (1) Labial palpus extending to just beyond base of pro-trochanter instead of to (or near) apex of procoxa as in other subspecies. (2) Distal arm of sternum 9 (fig. 55) with apical lobe (AP.L.) appearing at first glance to be largely truncate, but actually upper portion (fig. 1, pp-qq) convex, and only lower, longer portion (oo—pp) quite straight. In the nominate form (fig. 47, D.A.g and fig. 21) the 2 portions are subequal and oo-pp is concave. (3) Lower arm (fig. 57, L.A.) of securifer generally somewhat acute apically, primarily due to the fact that ventral margin is straight to near apex, instead of being sinuate and upcurved subapically (figs 23, 44) so that the tip is broadly angled. (4) Tanned portion of quasi-crochet (Q.C.) about 3-5 times as long as broad at middle, not 4 times. Similar to the nominate form in other respects, including F., and even the details of the internal female genitalia. ILLUSTRATIONS 55. Distal arm of sternum 9 60. Spermatheca and sternum 7 57. Apex of aedeagus 61. Genitalia (2) NEW TAXA OF FLEAS 235 ComMENT. It may be significant that there are no records of M. vobinsoni from North Borneo, where 4 other species of Medwayella are known to occur, and where members of the type-host, C. notatus, were examined for fleas, in one locality, namely Mt. Kinabalu at 5000 ft elevation or higher. However, even where present, M. vobinsoni is rare at such altitudes (at least in Malaya). There have been too few collections from the foot-hills and coastal areas of North Borneo to provide adequate data in regard to the possible occurrence of M. robinsoni. Since this species is so widely distributed in Sarawak, one would expect a similar distribution in N. Borneo, if it were present at all, but as mentioned below in the section on zoogeography, the flea fauna of North Borneo may be significantly different from that of at least southern Sarawak. The subspecies name is derived from the Latin word peregrinor, meaning to travel abroad, and was selected to emphasize the distance between the mainland nominate form and the Bornean subspecies in terms of space, if not geologic time. However, it is not necessarily implied that the spread of M. vobinsoni or of its hosts, through the eons, was from Malaya to the islands. This point is discussed in the third paper in this series and which deals with zoogeography. 4c. Medwayella robinsoni bogora subsp. nov. TYPE MATERIAL. Holotype 4, allotype 9 (B-78347-1) ex Callosciurus notatus ; INDONESIA: Java, Bogor; elev. 260 m; 15.VII.1965; Coll. J. M.Stusak. Paratypes as follows: 16 3, 17 2 with same data as holotype, but collected 21.VI.1965-—2.I1.1966. 23,1 Qex C. nigrovittatus; loc. cit.; 21.VI.1965. 1 Q ex Petaurista elegans ; Tjibodas; 2.X.1965. Id, 4 2 ex Lariscus insignis; loc. cit.; 3.X.1965. All collected by J. M. Stusak. Holotype and allotype deposited in U.S. National Museum, with holotype bearing U.S.N.M. number 71600. Paratypes in collections of B. P. Bishop Museum (Honolulu), that of author, and other institutions listed for M. dryadosa above. Diacnosis. Immediately separable from the nominate form and the new sub- species herein described in that notch in distal arm of sternum g (fig. 54, NCH.) is nearly symmetrical, i.e., lower margin (cf. fig. 1, mm-tr) is only slightly longer than upper (oo-1r) instead of being distinctly shorter (fig. 21). Equally characteristic is the fact that axis of notch (nn-1r) parallels the long axis of the bodyinstead of inclin- ing ventrocephalad. Further distinguishable from M. r. robinsoni as follows: (1) Height of thumb (fig. 58, THM.) of alpha-portion of Ford’s sclerite about twice as broad (long) apically as high and thus not nearly as tall as in nominate form (figs 23, 44), where THM. is nearly as high as long. (2) Upper arm (U.A.) of securifer with free ‘‘snout”’ relatively shorter and stouter, about 2-5 times as long (measured from vertical internal root) as broad at middle, instead of 3:3 times. (3) Apical lobe (AP.L.) of D.A.g with upper portion (cf. fig. I, pp-qq) oblique, not convex. (4) Lacking bead-like sclerotizations on dorsal wall of vagina (fig. 62, VAG.), distad of glandula vaginalis (G.VG.). (5) With only 2 bristles on ventral lobe of sternum 7 (7 S.), not 3. (6) Female often with a “‘supernumerary”’ bristle between rows II and III of head; this is unusual in mainland M. robinsoni (the nominate form). Very similar to the nominate form in other essential respects. 236 ROBERT TRAUB ILLUSTRATIONS 54. Distal arm of sternum 9 62. Spermatheca, genitalia and sternum 7 58. Apex of aedeagus 63. Proximal arm of sternum 9 CoMMENT. The known specimens were primarily all from Callosciurus notatus, but it is anticipated that this subspecies also commonly occurs on C. nigrovittatus. The data, although inadequate, also suggest that, like the nominate form, M. 7. bogora readily infests other squirrels, etc., in its habitat. The name bogora should be regarded as a coined term, although suggested by the type-locality, Bogor. 4d. Medwayella robinsoni tiomanica subsp. nov. TYPE MATERIAL. Holotype ¢ (B-55344-1) ex Callosciurus nigrovittatus; WEST MALAYSIA: Pahang, Pulau Tioman, on Kampong Tekek-Kampong Juara Track; elev. 500 ft; 30.III.1962; Coll. Ng Cheong Kee et al. Allotype 2 (B-55335) ex C. notatus; loc. cit., but Kampong Tekek; 18.III.1962. Paratypes (all collected by Ng Cheong Kee et al. in 1962, unless otherwise indicated) as follows: 2 § with same data as holotype. 2 g, 3 9 ex Sundasciurus tenuis; loc. cit. but 20.1II. 34, 3 2 ex Tupaia glis; loc. cit. but elev. 150-500 ft; 20-23.III. 2 g ex Lariscus insignis; loc. cit. but elev. 150 ft; 22.11]. 3 92 with same data as allotype but 18-19. III. Ig, 2 9 ex C. nigrovittatus; loc. cit. 3 g, 1 9 ex T. glis; loc. cit. but 2.V.1966; Coll. M. Nadchatram. 1 9 ex L. insignis; loc. cit. but 1.V.1966; Coll. M. Nadchatram. 73, 3 2 ex Iomys horsfieldi; Kampong Juara; elev. 150 ft; 25-31.III. 2d, 2 2 ex S. tenuis; loc. cit. but 11.1V. 10 g, 4 9 ex T. glis; near camp; elev. 50-1000 ft; 8-9.1V. 1 ¢ ex Rattus t. tiomanicus; loc. cit. but elev. 500 ft; 9.1V. 564, 4 9 ex T. glis; Ulu Sungei Ayer Besar; elev. 800-900 ft; 9-11.1V. 1d ex L. insignis; loc. cit. but elev. goo ft; 10.1V. 14,1 Qex R. sabanus; loc cit. but 11.1V. 24,1 Qex T. glis; Kampong Mokut; 9.IV. 1 9 ex L. insignis; Ulu Lelang, 17.1V. 1 92 ex R. surifer; loc. cit. but 16.1V. 1g ex C. notatus; Pulau Tulai, 6 mi NW of Pulau Tioman; 4.1V. 1 g, I 2 ex C. notatus nest; loc. cit. Holotype (U.S.N.M. No. 71601) and allotype deposited in U.S. National Museum. Paratypes in that insti- tution, the British Museum (Natural History), B.P. Bishop Museum (Honolulu), University of Malaya and Institute for Medical Research (both Kuala Lumpur) and the author’s collection. Dracnosis. (1) Apical lobe (fig. 53, AP.L.) with upper portion (cf. fig. 1, pp—qq) rather oblique and straigt instead of being convex as in the nominate form (fig. 21, 47); not nearly as vertical as in M. r. peregrinata. (2) Thumb (fig. 56, THM.) of alpha-portion of Ford’s sclerite with stem shorter than nominate form, the untanned vertical portion definitely shorter than distal breadth, instead of only slightly so. (3) Lower arm (L.A.) of securifer with free portion proportionately longer and narrower than in mainland form e.g., length (measured from level of apex of pivotal chord, PIV.CD.) about 1-5 times as long as high at that level, not merely 1-2 times. Relative narrowness of extended part of L.A. applies to apical excised region as well : as proximal section. (4) Lacking bead-like sclerotizations on dorsal wall of vagina ; (fig. 64, VAG.) distad to glandula vaginalis (G.VG.) which are characteristic of the NEW TAXA OF FLEAS 237 nominate form. (5) Ventral lobe of sternum 7 (7 S.) subequal to lobe above it, instead of being definitely larger.® ILLUSTRATIONS 53. Distal arm of sternum 9 56. Apex of aedeagus 64. Spermatheca, genitalia and sternum 7 ComMEntT. WM. +r. tiomanica infests a variety of squirrels and small mammals of similar habit and habitats in the forest on the hills of Tioman Island, primarily Tuvbaia and tree-squirrels, thereby paralleling the mainland form. 5. Medwayella phangi sp. nov. TYPE MATERIAL. Holotype gf and allotype 2 (B-46948-1) ex Sundasciurus tenuts; WEST MALAYSIA: Selangor, Subang; elev. 500 ft; 30.X.1957; Coll. R. Traub. Paratypes (48 g, 19 2) as summarized in Table 1 (p. 216). Holotype (U.S.N.M. number 71602) and allotype in collections of U.S. National Museum. Paratypes distributed as for M. dryadosa. Diacnosis. Male instantly separable from M. dryadosa and M. robinsoni in that dorsal margin of distal arm of $ sternum g is apically homolate (uniformly flat) (fig. 68, HOM.) terminating in an apical nubbin or short claw, instead of bearing a distinct subapical notch (fig. 20, NCH.) and an apical truncate or sinuate lobe (fig. 21, AP.L.). Further distinguishable from these species in that :(1) Quasi-crochet (fig. 71, Q.C.) is narrowed, elongate, its length, measured along entire ventral margin, more than 2-5 times length of base of phylax (PHY.) and upcurved; instead of being of quite uniform breadth, straight and short (at most only somewhat longer than base of PHY.), (figs 22, 23, Q.C.). (2) Dorsal region of Ford’s sclerite characterized by a groove-like process (fig. 71, GRV.) formed by the close appression of the inner faces of the anterior portions of alpha-portion (ALPH.) and elongated base of upper arm (U.A.) of securifer, the tanned and parallel margins creating the effect of an open tube. M. dryadosa and M. robinsoni with Ford’s sclerite lacking narrow groove; and instead apex of alpha-portion is thumblike (figs 22, 23, THM.). (3) Deltoid flap (DEL.FL.) very narrow due to great depth and height of sinus in hood, which extends far cephalad and hence exposing apex of phylax (PHY.) and body of crochet (B.CR.), instead of being broad and covering all of apical sclerites except tips of securifer (U.A. and L.A.) and crochet process (CR.P.). (4) Pivotal chord (PIV.CD.) well tanned and upcurved caudally instead of being lightly sclerotized and straight. (5) Movable finger (figs 65, 66, F.) with distal fringe of bristles (D.FR.) with bases of upper 4 in a straight line, and caudal margin also straight here, instead of line of bases of D.FR. and adjacent caudal margin each forming an arc (figs 14, 17). (6) Margins of F. below D.FR. fairly strongly curved and subparallel to near apex of immovable process P. (level r-s in fig. 1, g.v. and cf.) not with a slight curve to F. and Margins diverging at midline so that both are rather convex at level of conical process _ 5 If the specimen has been distorted in mounting, one side may appear larger than the other. Both sides should be similar in appearance before interpretations are made. 238 ROBERT TRAUB (C.P.) (i-q). (7) Male sternum 8 (fig. 69) with distinct microtufts on caudal 4 or } of ventral margin; not microdenticulate (fig. 29, s3-t3). (8) Upright portion (bb-ff) of dorsal (proximal) arm of sternum g mainly convex; not flat as in M. dryadosa (fig. 18, P.A.g). (9) Upper lobe of 9 sternum 7 (figs 75, 76, 7 S.) slightly concave above subventral sinus, not convex (figs 26, 49). (10) Perula of bursa copulatrix (fig. 77, P.B.C.) long and mainly vertical, e.g., vermiform, its caudal margin well tanned and resembling a shallow, straight arc; sac of B.C. (SAC.) dilated and long (sausage- shaped) paralleling it; instead of P.B.C. being short and oblate-ovate and SAC. quadrate above and behind it (figs 31, 32). (11) Paragenital morion (PG.M.) well developed, several times longer and broader than glandula vaginalis (G.VG.) ; not merely subequal to it. Near M. rhaeba (figs 138, 139) from Mt. Dulit, Sarawak, and known only from 1 pair and agreeing with it as follows: (1) With a groove-like process (fig. 71, GRV., M. phangi) on Ford’s sclerite. (2) Upper arm (U.A.) of securifer (SEC.) relatively long and narrow. (3) Quasi-crochet (Q.C.) long, subapically narrowed and upturned. (4) Distal arm of sternum 9g (fig. 69) lacking a subapical notch; dorsal (anterior) margin rather flat (HOM.) between apex and subapical lobe (SUB.L.). (5) posterior region of ventral margin of g sternum 8 microtufted (fig. 69, 8 S.). (6) Female sternum 7 (fig. 76) shallowly concave above subventral sinus. Markedly differing from M. rhaeba as follows: (1) in new species, constricted portion of movable finger (fig. 66, F.) below distal fringe (D.FR.) (e.g., region marked j-k in fig. 1) almost as broad as region above it, whereas in M. rhaeba (fig. 138) j-k is 3 width of apex (I-p). (2) Caudal margin of F. only somewhat convex at level of apex of P. (r), its breadth here equal to less than } length of F. above P. (n-s); not so grossly convex that breadth equals more than } of n-s. (3) F. so little curved that caudal margin below D.FR. is shallowly concave, instead of being so strongly arched that dorsocaudal angle (tip of stiva) (p) is in a plane with ventrocaudal tip of fulcral sclerite (F.S.) (a3). (4) The pair of longish thin bristles on caudal margin of F. well distad of the middle of sinus, closer to D.FR. than to apex of P., instead of being closer to P. than to D.FR. asin M. rhaeba. (5) Apical margin of distal arm of 3 sternum 9 (fig. 68) flattened (qq-ss) and tip (qq) extending slightly as a short, straight nubbin, whereas in M. rhaeba the top is somewhat convex and bears a distinct tooth-like projection (qq) which is somewhat curved (fig. 138). (6) Upeurved, narrowed por- tion of quasi-crochet (fig. 71, Q.C.) shorter, its length (measured along ventral margin) much less than distance from base of process to ventral angle, instead of slightly exceeding it. (7) Upper arm (U.A.) of securifer slightly arched in new species, not straight. (8) Sinus near ventral margin of ? sternum 7 (fig. 76, 7 S.) more acute than in M. rhaeba, making an angle of about 23° with horizontal, not 30° (fig. 139). (9) Tergum 8 of 9 (fig. 77, 8 T.) with a lobe at ventrocaudal angle; this gies lacking in M. rhaeba. Readily separable from M. javana and M. arcuata by the characters cited as numbers 1-4 above in comparison with M. dryadosa and M. robinsont. Further distinguishable from M. javana by the following: (1) F. fairly broad subapically (fig. 66), breadth immediately below D.FR. (j-k) being almost # of length from this level to apex (j/k-n) instead of being narrow, less than } of length above it, as in M. javana NEW TAXA OF FLEAS 239 (fig. 133). (2) Quasi-crochet (fig. 71, Q.C.) more than twice as long as base of phylax (PHY.) instead of only slightly longer (fig. 95). Distingiushable from M. loncha at a glance, viz., (I) D.A.g (fig. 67) lacking conspicuous spur (mm) at middle of anterior (dorsal) margin (fig. 127). (2) Quasi-crochet (fig. 71, Q.C.) long and narrow, about thrice as long as broad at base instead of being short and broad, only about twice as long as broad (fig. 128). DEscripTIon. Essentially asin M. dryadosa except as noted in diagnosis above and as follows: Mesepisternum with lobe at cephaloventral corner virtually straight. Male. Lumacaudate process (fig. 69, LUM. and fig. 70) not as well developed— viz., lateral apical seam or flap inapparent; process shorter, apically more narrowed and not as heavily armed with spiniforms, either in number or regarding modification, i.e., with only 5-6 transverse rows of short, stout, slightly curved bristles of subspini- forms which are well separated from one another, instead of 12-16 rows of close-set modified bristles of which apical and ventral ones, especially, are true spiniforms (i.e., short, dark, basally widened, curved and broad to near pointed apex, etc., as in fig. 12); LUM. subapically only about # as broad as near base instead of being sub- equal. Male with upper antepygidial bristle only 3 length of lower one. Distal arm of § sternum g with supramedial flap (fig. 69, SUP.FL.) possessing fairly short and weakly sclerotized thickening (THK.) on base, and that is apical. SUP.FL. extend- ing to apex of D.A.9. Dorsal (upper) margin of securifer, i.e., basal 3 of upper arm (fig. 71, U.A.) concave so that entire structure is biconcave dorsally; free portion of U.A. with basal axis somewhat upcurved (about 40° from axis of base). Lower arm (L.A.) apically truncate but lower angle rather acuminate and with tip usually recurved and hence often obscured. Ventral margin of base of S.I.T. broadly convex to level of dorsal triangular thickening (which is not spur-like) at apical third. Apex of S.I.T. with ventral margin extended caudad. Phylax (PHY.) nearly twice as high as broad at base. Quasi-crochet (Q.C.) somewhat boomerang-shaped ; ventral angle rounded; proximal portion slightly exceeding distal; broadest at angle; upturned distal portion with ventral margin shallowly concave; upper margin curving at distal } so that apex is ovate dorsally. (Note—Caudomarginal submedian group of bristles, CM.G.B., on D.A.9, usually with 3 stout bristles, not 2 as in specimen drawn in figure 68.) Female. Sternum 7 (figs 76, 77, 7 S.) with ventrocaudal lobe somewhat too ‘ong and broad to be an inverted, reversed image of sinus above it. Mesal genitalic ridge of 8 T. (figs 72, 77, M.R.8) consisting of a broadly crescentic sclerotization with short, broad, sinuate tail extending caudad and an indefinite tanned stalk arising from ventral base of crescent. Sternum 8 relatively broad, length distad to level of basal (most anterior) bristle subequal to breadth of segment at that level (or shorter), with about 4-6 bristles of which all but x of apical ones are dorsal and subapical; lower apical bristle the longest, but this shorter than twice breadth of 8 S. at level of basal bristle. Spermatheca (figs 74, 77) with dorsal constriction nearer to middle than to caudal }; bulga (B.) about 2-4 times as long (middle longitudinal axis) as broad at constriction ; dorsal peak on caudal bulge short, inapparent unless spermatheca seen in fulllateral aspect. With internal sector of hilla (H.) shorter than external portion. 240 ROBERT TRAUB Sac of bursa copulatrix (SAC.) an indefinite, long, tubular, dilated, sausage-like structure about 3-4 times as broad as diameter of B.C. near orifice to vagina. Spermathecal duct (D.SP.) with dilated portion (DIL.P.) equal to diameter of SAC. and then gradually narrowing so that at middle its diameter is } that of DIL.P. and anterior 1, about 4. Perula (P.B.C.) usually subvertical and quite straight except for cephalad gradual curve of extremities, or else resembling a flattened arc with concavity anterior. Glandula vaginalis (G.VG.) a straight but oblique tanned slit, about thrice height of small accessory “‘gland” between it and unmodified orifice of B.C. Dorsal wall of vagina well developed caudad of G.VG. and here with marginal striae or bands. _ Paragenital morion (PG M.) highly developed, consisting on each side of a tube-like coil surrounding all but the rear of an ovoid lightly tanned core, and occupying most of the region between P.B.C. and lower part of sternum 9 (9 S.). ILLUSTRATIONS 66. Processes of clasper 72. Tergum 8 (9) 68. Distal arm of sternum 9 73. Anal lobes and stylet (2) 69. Sternum 8 (3) 74. Spermatheca 7o. Lumacaudate process 76. Sternum 7 (9) 71. Apical region of aedeagus 77. Spermatheca, genitalia and anal segments ComMENT. The species is named for Phang Ong Wah, who has faithfully served USAMRU (Malaya) as a collector and careful observer of small mammals and ecto- parasites in Malaya, North Borneo and Sarawak since 1948, despite the arduous labor and physical danger (from both disease and hostile guerillas) that the field-work often entailed. Mr Phang was responsible for the collection of many of the specimens and new species of fleas and trombiculid mites studied by the writer, and also contributed significantly to our investigations on the chemoprophylaxis, treatment, control and ecology of scrub typhus. A distinctive subspecies of M. phangi occurs in Sarawak and the nominate form will therefore be characterized before the description of the Bornean one, and before a discussion of the host-relationships and distribution of the Malayan form. 5a. Medwayella phangi phangi DiaGnosis. (1) Movable finger (fig. 66, F.) with distance between basal bristle of distal fringe (D.FR.) and apex of F. subequal to breadth of F. at subapical constric- tion. (2) Concavity of caudal margin of F. shallow; this arc with its chord about 9 times its height. (3) D.A.g (fig. 68) with homolate margin (HOM.) relatively long in that the slope of the subapical lobe (SUB.L.) is fairly abrupt, terminating at basal + of distance between apex of SUB.L. and tip of D.A.g (mm-—qqin fig. 1). (4) Distance mm -qq scarcely exceeding that between apex of SUB.L. and distal margin of median microspiculose area (M.MSP.) (mm-kk), (5) Sinus above ventral lobe of 9 sternum 7 (fig. 76, 7 S.) subequal to lobe beneath it. (6) Paragenital morion (fig. 77, PG.M.) with a thin transverse sclerotization across anterior portion of core. HOST-RELATIONSHIPS AND DISTRIBUTION AND OTHER COMMENTS. It is note- worthy that more than 90% of Malayan M. p. phangi (the nominate form) were from NEW TAXA OF FLEAS 241 Tupaia glis or Sundasciurus tenuis or S. lowi and that there were only 2 records from any of the truly arboreal squirrels (Table 2, p. 259), and only 1 specimen was found on a rat, even though the Callosciurus and Rattus constituted the largest categories of hosts examined. The absence of M. p. phangi from Dremomys and Rhinosciurus and the dearth of records from Lariscus is also striking. The geographic and ecological distribution of this flea in Malaya is unusual and merits further study, but available data are summarized in Table 1 (p. 216). Al- though over the years since 1947, both USAMRU and DMZ-IMR collected mammals and fleas more intensively in the dipterocarp forest in the foothills around Kuala Lumpur (i.e., the forest reserves at Gombak, Ampang, Ulu Langat and Bukit Lagong areas) than in any other region or habitat, only 4 specimens of M. p. phangi were ever found there. In contrast, 27% of the specimens (and 30% of the numbers of collections, i.e., the hosts infested with M. p. phangi) were made from the secondary forests of Subang, which is also in Selangor and equidistant from Kuala Lumpur with the other areas, but at slightly lower elevation (viz., 400-700 ft versus I000— 2000 ft). Further, 40% of the specimens (35 °% of the collections) were from the foot- hills of Perak, 8-10 miles from Ipoh at elevations of about I100 ft. This would at first glance suggest that the distribution might be an altitudinal one, a possibility that is reinforced by the absence of records from montane hosts like Dremomys and Tamiops mceclellandi. However, M. p. phangi is not really a lowland species, because 15 % of the specimens came from Kedah Peak, at elevations often above 3000 ft, an area where relatively little collecting of ectoparasites was done by our teams and colleagues. One likely explanation is that M. p. phangz is essentially a flea of secondary or disturbed forest, rather than primary forest, for all of the sites where it has been taken in reasonable numbers are regions where there has been much cutting of timber over the years or where there is an abundance of limestone formation and a dearth of the huge dipterocarps associated with primary forest in Malaya. The forest reserves of Selangor, on the other hand, have been much less altered by man, and the areas surveyed were not in limestone terrain. Another factor worth checking is that of rainfall, for the Perak and Kedah Peak collections happened to have been made during periods of relative drought. The one-to three-week dry-spells noted annually or semi-annually in most of the southern two-thirds of Malaya do not occur at fixed periods of the year, and no special effort was made to collect Tupaia and S. tenuis in the dipterocarp forest during those irregular periods. Hence data are lacking whereby this point could be determined regarding the forested belts in Selangor. However, the distribution of M. p. phangi may prove to be due to zoogeographic factors, for like another new species described below and known only from west- central Malaya, it occurs in Sarawak as a distinct subspecies. Definite faunal rela- tionships may exist between these two regions, as mentioned anon, and as discussed in the third paper in this series. Further discussion on the host-relationships of M. p. phangi will be found in the comments following the descriptive sections on Medwayella. M. p. phangi also occurs in extreme southern Thailand (Nakhon Si Thammarat), where a single female was collected on an unidentified squirrel by Dr Boonsong 242 ROBERT TRAUB Lekagul and kindly sent to us by Dr R. E. Elbel. It seems significant that no additional specimens of M. phangi turned up in the rather extensive collections of fleas made in Thailand by Dr Elbel, for the vast majority of Elbel’s Thai specimens were taken north of the Malayan peninsula. In these ““non-Malayan”’ provinces of Thailand, a new species of Medwayella, described later in this paper, was commonly found on Menetes berdmorei, a ground-squirrel, and it is of interest that this new species was also taken at Nakhon Si Thammarat. 5.b Medwayella phangi tana subsp. nov. TYPE MATERIAL. Holotype ¢ (B-48275-1) and allotype 2 (B-48275-3) ex Tupaia tana; EAST MALAYSIA: W. Sarawak, Pueh; 11.VI.1958; Coll. T. C. Maa. Para- types as follows: 5 3, 5 2 with same data as holotype. 1 ¢ ex Callosciurus; loc. cit. 5 3, 6 9 ex T. tana; Sarawak, Kuching, 5th mile, Stapok Rd.; [X.1951; Coll. T. Harrison. Holotype (USNM number 71603) and allotype in collection of U.S. National Museum. Paratypes in B. P. Bishop Museum (Honolulu), the British Museum (Natural History) and the author’s collection. DracGnosis. Close to nominate form, and differentiated as follows: (1) Movable finger (fig. 65, F.) narrower and more arched below distal fringe (D.FR.), viz., fringe set on a low “‘stalk”’ in the Sarawak form—breadth of F. at constriction (indicated as j-k in fig. 1) much shorter than distance between basal bristle of fringe and apex of F. instead of being subequal; the arc formed by caudal margin of F.,e.g., between D.FR. and where margin crosses apex of immovable process P. (k—r) quite deep, e.g., with chord equal to 4-5 times of altitude, instead of about 9-fold. (2) Apex of D.A.9 (fig. 67) with homolate margin (HOM.) comparatively short because slope of subapical lobe (SUB.L.) commences at basal 4 of distance between tip of SUB.L. and apex of D.A.g (mm-qq), instead of basal } (fig. 68). Distance mm—qq nearly twice that between apex of SUB.L. and distal margin of median microspiculose area (M.MSP.) (mm-kk) instead of scarcely exceeding it. (4) Sinus above ventral lobe of 9 sternum 7 (fig. 75, 7 S.) definitely shorter and narrower than lobe below it, instead of being subequal (fig. 76, 7 S.). Paragenital morion with a tanned chord along ventral margin of inner core, near its apex. In nominate form there is instead a thin trans- verse sclerotization curving towards middle of or dorsal part of core (fig. 77, PG.M..). ILLUSTRATIONS 65. Processes of clasper 67. Distal arm of sternum 9 75. Sternum 7 (9) CoMMENT. The name of the subspecies denotes a common host, Tupaia tana, as per existing records. However, collections in Sarawak have been too limited in general to determine whether the Bornean subspecies regularly infests ground- squirrels or other hosts. 6. Medwayella limi sp. nov. TYPE MATERIAL. Holotype ¢ (B-45230) ex Rhinosciurus laticaudatus; WEST MALAYSIA: Selangor, Kepong, Bukit Lagong; 25.VII.1956; Coll. R. Traub. NEW TAXA OF FLEAS 243 Allotype 9 (B-52520) ex R. laticaudatus; Selangor, Ampang Reservoir; 13.1.1960; Coll. J. R. Audy. Paratypes as follows: 4 J with same data as holotype. 2 ibid., but 31.X.1956. 1 with same data as allotype. 2 ibid., but 16. VIII.1960; Coll. J. R. Audy. 104, 2 2 Selangor, Ulu Langat Forest Reserve; 21.I—18.II.1965; Coll. Lim Boo Liat. 1 ¢ ibid., but 15.1.1957; Coll. R. Traub. 3 g, 1 9 Selangor, Ulu Gombak Forest Reserve, 16-17 mi N of Kuala Lumpur; 22.VIII, 4.X, 16.X1I.1956; (R.T.). 1d ibid., but 30.1X.1968; (L.B.L.). 24 Selangor, Pahang Road, 16 mi N of Kuala Lumpur; 5.VII.1948, 7.111.1956; (R.T.). 4, 1 2 Trengganu, Sungei Tong Forest Reserve; 26.VIII.1965; (L.B.L.). 2 g ex Lariscus insignis; Selangor, Ulu Gombak, 15-16 mi N of Kuala Lumpur; 1500 ft elev.; 22.VII.1957, 11.VI.1958; (R.T.). 1g Trengganu, Bukit Besi; 5.VIII.1958; (R.T.). 1g ex Tupaia glis; loc cit. I 9 ex Rattus sabanus ; Selangor, Ampang Reservoir; 28.VII.1960; (R.T.). Holotype (U.S.N.M. number 71604) and allotype deposited in U.S. National Museum. Paratypes distributed as for M. dryadosa. Diacnosis. Very close to M. phangi but distinguishable as follows: (1) Groove- like process of Ford’s sclerite (fig. 82, GRV.) incomplete apically due to fact that thickened ventral margin of base of upper arm (U.A.) does not extend as far dorsally as does the opposite member of the furrow, and instead terminates well before apex. In phangi (fig. 71) the edges of the groove parallel each other all the way to apex. (2) Apex of anterior member of GRY. extending over and beyond (dorsad) base of U.A. as a broad, truncate and squared flap whose caudal margin parallels its anterior one, and which crosses base of U.A. at anterior }. In phangz, this flap does not reach dorsal margin of base of U.A. and its caudal margin is convex. (3) Free extension of U.A. (snout) more upright, making an angle of about 45° with longitudinal axis of base of U.A. instead of only about 20-25°. (4) Free portion of lower arm (L.A.) quite narrow subapically due to slope of 60° of upper (dorsal) margin and emargina- tion of ventral one. In M. phangi, upper margin sloping at 80° and ventral margin straight to near apex. (5) Tongue-like apical portion of quasi-crochet (Q.C.) shorter, due to Q.C. being angled at apical 4 instead of near midpoint (fig. 71). (6) Phylax (PHY.) with base much broader; its height only less than 1-4 times length of ventral margin instead of nearly twice as high. (7) Subapical lobe (fig. 81, SUB.L.) of D.A.g with its height equal to twice that of nubbin at apex (indicated as qq in fig. 1) and with homolate margin (HOM.) correspondingly shortened, e.g., flat surface between qq and mm shorter than breadth of D.A.9 at level of apex of M.MSP. (kk) instead of with SUB.L. about subequal to apical nubbin (figs 67, 68) and HOM. longer than D.A.g is broad at level of kk. (8) Movable finger (fig. 80, F.) with bases of distal fringe of stout bristles (D.FR.) making an oblique line with longitudinal axis of F. (n-s), not a subparallel one (fig. 66). (9) Mesal genital ridge of 9 8 T. (fig. 79, M.R.8) with ventral arm longer, thinner and more discrete than in M. phangi (fig. 77), e-g., ventral tanned area clearly exceeding in length the more dorsal oblique sclerotization ; and that of the median ovate ventral bulge. (10) Perula of bursa copulatrix (P.B.C.) with sclerotized caudal margin essentially straight or slightly concave (e.g., with sinus caudal) instead of being arched (convex dorsally) or with at least upper and lower fourths curving cephalad. (11) Paragenital morion (PG.M.) with a subventral sclerotization rather than a transverse one approaching dorsal margin. (12) Ventral 244 ROBERT TRAUB lobe of sternum 7 (7 S.) taller than long, instead of with subequal dimensions. (13) Glandula vaginalis (G.VG.) 4 or more times length (height) of accessory ‘“‘gland”’ (cleft) preceding it, not merely thrice as long. In other respects M. limi is essentially like M. phangi. The diagnostic features cited under M. phangi serve to separate M. limi from M. dryadosa, M. robinsoni, M. arcuata, M. loncha, etc. ILLUSTRATIONS 78. Spermatheca (variation in aspect) 82. Apex of aedeagus 79. Spermatheca and genitalia 83. Lumacaudate process 80. Processes of clasper 84. Spermatheca and genitalia 81. Distal arm of sternum 9 85. Sterna 8 and 9 (9) CoMMENT. The species is named in honour of Mr Lim Boo Liat of the Division of Virus Research and Medical Zoology of the Institute for Medical Research, Kuala Lumpur, as partial acknowledgement of the tremendous assistance he has rendered the author and other members of USAMRU in their research on vectors and reservoirs of infection in Malaya and Borneo. Ever since 1948, Mr Lim not only collected mammals and ectoparasites with us or for us, but provided identification of hosts and served as major-domo, interpreter or liaison officer on innumerable field-trips and often under very difficult circumstances. Mr Lim also participated in Lord Medway’s expedition to Gunong Benom, and, also, in his own right has contributed a great deal to the knowledge of the role of ectoparasites and small mammals in infections in Southeast Asia and in popularizing the study of natural history in that part of the world. M. limi is a sibling species of M. phangi and it is of interest that the two species appear to differ significantly regarding host and habitat. Virtually all the records of M. limi are from the ground-squirrel Rhinosciurus laticaudatus, while the great majority of its sister-species were collected from Sundasciurus tenuis and Tupaia, and it was never found on Rhinosciurus. While nearly all of the M. limi were taken in the dipterocarp forests in the foothills near Kuala Lumpur, it probably is distributed in similar habitats throughout Malaya, since it was also collected in such terrain in Trengganu. There are no records from the lowland forest at Subang nor from the limestone hills of Ipoh, which were the main source of M. phangi. Both host-wise and with respect to micro-environment, M. limi resembles M. dryadosa, a point that is further discussed after the taxonomic portions of this article (p. 258 et seq.). 7. Medwayella thurmani sp. nov. TYPE MATERIAL. Holotype g, allotype Q (B-15288) ex Menetes berdmoret; THAILAND: Nakhon Ratchasima, Si Khiu, Pak Chong; 23.1X.1952; Coll. R. E. Elbel. Paratypes, all from Thailand, as follows: oe °° Host 15 8 I -— Rattus vajah group I 1 _ Rattus vattus 5 3. Menetes berdmorei rt 1 Menetes berdmorei - 5 £4xTupaia glis - 1 Tupaia glis 3. 1 £Menetes berdmorei 2 — Menetes berdmorei I — Callosciurus finlaysoni 3. 5 £Menetes berdmoret Tupaia Menetes berdmorei Menetes berdmorei I -— Tupaia glis Rattus vattus Menetes berdmorei Tupaia glis — Menetes berdmorei Menetes berdmorei Menetes berdmorei Callosciurus notatus miniatus Tupaia glis NEW TAXA OF FLEAS Changwat & Locality Menetes berdmorei Nakhon Ratchasima, Si Khiu, Pak Chong Nakhon Ratchasima, Si Khiu, Chan Thuk, Pongekeng Nakhon Ratchasima, Pak Thong Chai Nan, Pang Nam Un Nan, Sar Dist., 14 km S of Sarat Pahang Nan, Ban Pha Hang Loei, Dan Sai, Kok Sathon, Phu Lom Lo Mt., 2100 m Loei, same as above Loei, same as above Loei, same as above Loei, Dan Sai, Na Haeo, Ban Na Muang, approx. 1780 m Khon Khaen, 112-138 km on Khon Khaen—Loei Rd., 3-6 mi S of Loei Border Khon Khaen, 119 km on Khon Khaen—Loei Rd., 3 mi S of Loei Border Chanthaburi, Klong Ta Kong, A Pong Nan Ron Chanthaburi, same as above Chanthaburi, Wad Bom Phu, A Tha Mai Chanthaburi, Khlung, Tapon, Khao Sabap Mt. Chiengmai, B. Nong Prue Kanchanaburi, Trakhanum Hinlaem Chon Buri, Si Racha, Khroa Wing Sara Buri, Kaeng Khoi, Muak Lek, Dan Kak Sat Nakhon Si Thammarat, Chawang, Chang Klang, Ban Na Khao Yai National Park, Khorat 245 Date Collector IX.1952 & R. E. Elbel IX.1953 15.11.1953 R. E. Elbel 20.1.1967 J. F. Reinert 19-21.1.1953 KR. E. Elbel & H. G. Deignan 9. XI1.1961 Phang Ong Wah 15-19.III.1961 K. Thonglongya 23.11.1954 R. E. Elbel & Boonsong Lekagul 29.III-2.1V. R. E. Elbel & 1954 Boonsong Lekagul 15.11.1955 R. E. Elbel 17.11.1955 R. E. Elbel 9-27.X.1954 R.E. Elbel 26.X1.-2.XII. R. Traub & 1962 J. E. Scanlon 28.X1.1962 R. Traub & J. E. Scanlon 22.11.1963 J. E. Scanlon 22.11.1963 J. E. Scanlon 21.11.1963 J. E. Scanlon 26. XI11.1953 Boonsong Lekagul 15.X1.1965 D. Gould g-25.X1.1952 R.E. Elbel & H. G. Deignan 10. VII.1953 Boonsong Lekagul 8.1X.1953 R. E. Elbel 28.11.1954 Boonsong Lekagul 17-24.1X.1963 J. E. Scanlon Holotype (U.S.N.M. number 71605) and allotype deposited in U.S. National Museum. Paratypes distributed as for M. dryadosa. 246 ROBERT TRAUB Dracnosts. Distinctive in that apical portion of dorsal margin of distal arm of g sternum g (fig. gr) lacks a distinct notch (fig. 47, NCH.) or a long and deep arc (fig. 33, ARC.) and yet is not evenly homolate (fig. 67, HOM.) from above subapical lobe (SUB.L.) to apex. Instead it bears a small, shallow arc (fig. 91, ARC.) proximally and a scarcely differentiated, longer apical lobe (AP.L.). Also separable from known species in the hyperdevelopment of sclerotized thickening (THK.) at base of supra- medial flap (SUP.FL.) of D.A.g, which here extends from ventral region of ARC. to near apex of arm. Near M. dryadosa, and like it in possessing a truncate thumb (figs 90, 94, THM.), as well as an acuminate extension of the lower arm (L.A.) on Ford's sclerite (cf. fig. 22, M. dvyadosa). Further distinguishable from M. dryadosa as follows: (1) Subapical lobe (fig. g1, SUB.L.) not really lobate, its proximal (anterior) margin sloping gradually to mid- microspiculate area (M.MSP.) instead of markedly angling to form a distinct pro- jection (fig. 20). (2) Transverse sclerotization (T.S.) of D.A.g slightly arched proxi- mad (concavely facing base of arm) instead of being sinuate, and with concave margin facing distally. (3) Ford’s sclerite with dorsal margin of base of upper arm (figs 94, 97, U.A.) quite flat instead of convex (fig. 22). (4) Lower arm (L.A.) with a dorsal spur or tooth-like projection, instead of being rounded. (5) Fusion of lateral walls of aedeagal pouch (AE.P.-L.) with crochet process (CR.P.) well tanned, resulting quasi-crochet (Q.C.) well sclerotized through- out; the darker, crochet-like structure of Q.C. less than thrice as long (measured along dorsal margin, from caudal margin of phylax) as broad at middle in M. thurmani. In M. dryadosa, comparable tanned area of Q.C. much narrower, viz., 4 times as long as broad. (6) Caudal sinus of deltoid flap (DEL.FL.) much larger, e.g., almost equal in area to apex of hood (HD.) and median dorsal lobe (M.D.L.) above it, instead of being less than } that size. (7) Caudal (subvertical) margin of g sternum 8 (fig. Lor, m3-t3) shorter; height to level of longer bristle at dorsocaudal angle (m3) subequal to length of bristle, not 25% greater than length of bristle (fig. 29), due to greater slope of dorsocaudal margin in M. thurmani. (8) Lumacaudate process (LUM.) narrowing from middle to apex, instead of being broadly subovate (fig. 12). (9) Female tergum 7 (fig. 99, 7 T.) with lobe below antepygidial base (L.L.7) with angle rounded off; not acute (fig. 26). (10) Anal stylet (fig. 89, S.A.) with a subapical ventral bristle. This missing in M. dryadosa (fig. 16). (11) Ventral lobe of caudal margin of 9 7S. (figs 99, 102) almost as high as long, instead of apex about } as high as long (fig. 27). (12) Subventral sinus of 2 7 S. with basal 4 symmetrical, evenly curved instead of with dorsal margin sloping dorsocaudad almost immediately. DESCRIPTIVE NOTES. Essentially otherwise like M. dryadosa (cf., fig. 86, head g with fig. 4) except as follows. Dorsal margin of immovable process (fig. 92, P.) sinuate, but mainly convex. Mesal genitalic ridge of 2 tergum 8 (fig. 102, M.R.8) resembling an inverted ““Y’’ with rounded arms and a wide cleft. Spermatheca (figs 88, 102) with internal margin of bulga (B.) at base of hilla (H.) less oblique, about 35°-40°. Perula of bursa copulatrix (P.B.C.) elongate-ovoid, nearly as long as broad in middle, more dilated ventrally than dorsally; anterior margin straight and subvertical; caudal margin somewhat convex. Sac (SAC.) of B.C. small, its volume scarcely exceeding that of P.B.C., the portion immediately dorsal to perula with NEW TAXA OF FLEAS 247 height only equal to 4 that of perula. Glandula vaginalis (G.VG.) feebly sclerotized ; at times inapparent; sinuate. Paragenital morion (PG.M.) weakly tanned; indis- crete, presented as a narrow, ovoid structure lying across or near base of P.B.C. and perpendicular to it. ILLUSTRATIONS 86. Head and prothorax (3) 94. Apex of aedeagus 87. Movable finger 97. Apical region of aedeagus 88. Spermatheca 99. Sternum 7 (9) 89. Sensilium and anal segments (9) 100. Lateral metanotal area (¢) go. Apical region of aedeagus tor. Sternum 8 (3) gt. Distal arm of sternum 9 102. Spermatheca and genitalia 92. Processes of clasper ComMEnts. The species is named for the late Deed C. Thurman in token ackno- ledgement of the many and significant contributions on the ecology, control and taxonomy of arthropod-vectors of disease in Thailand which Dr Thurman managed to make before his tragic death from cerebral malaria in that country at the height of his career. M. thurmani is predominantly a flea of the ground-squirrel Menetes berdmoret, with occasional specimens occurring on Tupaia, as can be seen from the tabulation on page 245, which summarizes the data for paratypes. M. thurmanz is evidently a widespread species in Thailand ranging, with its main host, over most of that country, and hence I suspect that it will be found on that ground-squirrel in northern Malaya and Indo-China. 8. Medwayella calcarata sp. nov. TYPE MATERIAL. Holotype (B-49517) ex Lariscus insignis; WEST MALAYSIA: Perak, 8 mi W of Ipoh, Kledan Saiong Forest Reserve; elev. 1300 ft; I1.X.1958; Coll. R. Traub. Allotype 9 (B-48637) ex L. insignis; Perak, Maxwell Hill; elev. 3000 ft; 25.VI.1958; Coll. R. Traub. Paratypes as follows: 3 g with same data as holotype; 2 g ibid. but elev. r100 ft. 2 4 10 mi NW of Ipoh; 29-30. VIII.1958; Coll. R. Traub. 1 g, 1 9 Perak, Maxwell Hill, Gunong Hijau; elev. 4200 ft; 21.III.1958; Coll. R. Traub. Holotype (U.S.N.M. number 71606), allotype and I paratype deposited in U.S. National Museum. Paratype (I g) in the British Museum and remainder in the author’s collection. DiaGnosis. Separable from all described species by virtue of the modified fea- tures of the head (fig. 104, g), viz.: (1) Labial palpus extending to apex of pro- trochanter instead of only to near tip of procoxa. (2) Distal segment of maxillary palpus also unusually long—x1-75 times that of segment 3 instead of about 1-5 times. (3) With additional ‘‘supernumerary”’ bristles (SY.) on the preantennal region,e.g., 3 instead of 1 between rows II and III, with an extra 1 in row II, inf making a total of 19 long or medium-sized bristles instead of the usual 16-17. Female with 1 “supernumerary,” which is absent in most ? members of the genus. 248 ROBERT TRAUB Agrees with M. loncha (figs 118, 126-132) (and the 2 new species next described) in the following respects (and thereby differing from M. dvyadosa and allies): (1) Re distal arm of f sternum 9 (fig. 111): (a) Subapical lobe (SUB.L.) long and sharp- pointed, like a spur, instead of being lobate (figs. 20, 68). (b) Apical 4 of ventral (caudal) margin (qq—u3 in fig. 3; cf., also fig. 1) much more heavily sclerotized than anterior margin e3-qq). (c) Apex of ventral margin (g3-qq) appearing to project distad of arm as a rod-like extension (EXT.g) instead of curving to form an apical lobe (figs 20, 21, AP.L.). (d) With a caudal row of about 6 wide-set, fairly long, straight spiniforms instead of a subapical group (S.G.SPN.) of close-set, fairly small ones. (e) Tanned portion of dorsal margin distad of SUB.L. (c3-f3) extending only midway to apex, e.g., to distal end of supramedial flap (SUP.FL.) and then becoming semimembranous and curving toward ventral margin, thereby accounting for the apical rod-like extension. (2) Ford’s sclerite of the grooved type (fig. 113, GRV.). Readily separable from M. loncha as follows: (1) Movable finger (fig. 108, F.) not nearly as short and broad, viz., length distad from apex of immovable process (P.) (cf., fig. 1, n-s) 4 times breadth of F. at narrowest level (j-k), instead of scarcely more than thrice as long as broad there (fig. 126 ,F.). (2) Bases of setae of D.FR. forming a line tangential to main axis of F. (n-s) but almost subparallel to longitudinal axis of body of flea, instead of vice versa. (3) Sinus on apical (dorsal) margin of immo- vable process of clasper (P.) close to caudal margin and hence resulting in a dorso- caudal lobe, instead of bearing the sinus near the anterior margin and lacking a dorsocaudal lobe. (4) Manubrium with ventral margin lacking a distinct bulge (w) immediately anterior to sinus near posterior } (v) instead of being definitely convex there. (5) Distal arm of sternum 9g (fig. 111) relatively narrow at level of subapical lobe (SUB.L.) so that breadth (cf. fig. 3) (g3-d3) is only 1-3 times length of spur (d3-mm) instead of more than twice d3-mm (fig. 127). (6) Axis of EXT.9 (g3-qq) upcurved, directed more dorsad than caudad, instead of being straight, paralleling axis of D.A.g. (7) Tanned portion of quasi-crochet (fig. 113, Q.C.) about 1-7 times as long as broad, with dorsal margin slightly concave and nearly paralleling ventral one; anterior and posterior margins slightly convex. In contrast, in M. loncha (fig. 128) Q.C. only 1-3 times as long as broad; dorsal margin broadly convex; ventral margin angled at middle, with anterior } straight, horizontal, and posterior } concave, curving dorsocaudad; anterior and posterior margins straight, vertical. (8) Ford’s sclerite with groove (GRV.) broad and with margins equidistant and essentially parallel throughout; not narrowed at area of contact with dorsal expanded apex of body of crochet (B.CR.) and again at dorsal apex of groove. (9) Upper arm (U.A.) of securifer lacking a sclerotized nubbin at apical (caudal) 4 where side of lower arm (L.A.) moves dorsad to link with U.A. (10) Distance between dorsocaudal angle of apex of B.CR. and nearby dorsocaudal angle of U.A. subequal to height of slightly concave or straight apical (caudal) margin of U.A. instead of being nearly double the height of arcuate apical margin of U.A. (xz) Phylax much broader, viz., only slightly more than twice as long as broad at maximum girth (near ventral margin) instead of more than thrice. (12) Dorsal spur at apical 4 of sclerotized inner tube (S.I.T.) curved back over tube so that for most of its length it parallels S.I.T. and is close to it instead of being oblique to it. (13) S.I.T. broad proximad of dorsal spur NEW TAXA OF FLEAS 249 but much narrowed distad (and 4 its diameter) instead of only being somewhat narrowed. (14) Angled base of subventral sinus on 2 sternum 7 (fig. 103, 7 S.) quite diagnostic; in the other known female Medwayella from Malaya and Borneo the margin of the sinus is rounded (fig. 102). (15) Inner walls of sclerotized duct of bursa copulatrix (fig. 103, D.B.C.) in their entirety more tanned than bulk of perula, includ- ing anterior margin. In M. loncha only part of D.B.C. (fig. 118) well tanned, as is, at best, lower rear part of perula. (16) Dorsal anal lobe (fig. 103, D.A.L.) with g non- marginal bristles in 3 vertical rows preceding anal stylet, not with but 6 non-marginals in 2 irregular vertical rows (fig. 129). (17) Anterior margin of hilla (H.) of sperma- theca protruding into bulga (B.) (internal margins) for } length of hilla, instead of 4 (fig. 132). (18) Anal stylet (A.S.) about 5-5 times as long as broad at middle, and there somewhat constricted, instead of 4-6 times and gradually narrowing from base towards apex and very slightly constricted subapically (fig. 129). DESCRIPTIVE NOTES. The diagnosis above serves to cite differences from M. dryadosa as well (q.v.) but M. calcarata is otherwise quite similar, except as follows. Male. About Ig large or medium preantennal bristles in 5 irregular rows (fig. 104). Squamulum horizontal. Sternum 8 with caudal margin straight except for rounded corners; almost entire (scarcely microdenticulate) near ventrocaudal corner; long bristle near dorsocaudal angle with 2 or 3 marginal or submarginal, medium bristles above it. Lumacaudate process (fig. 107, LUM.) with dorsal margin virtually horizontal and straight; ventral margin of lobate portion oblique and usually tri- convex (trebly convex); apex broadly rounded; as measured from base of ventral margin, only about 1-1 times as long as broad at base, 1-6 times as long as broad at middle bulge and 2-5 times that of subapical bulge. Movable finger (fig. 108, F.) with caudal margin quite straight at level of apex of conical process (C.P.) (cf., fig. I, q), which in turn is only about 4-5 times as long (e-h) as broad at middle (f-g). Distal arm of sternum 9 (fig. 111) about 8 times as long (1i-qq) as broad at level (jj) of transverse sclerotization (T.S.). Distance between bulge (cc) at upper } of anterior margin of proximal arm of sternum g and apex of arm (aa) subequal to that from bulge (cc) to height of convexity of posterior margin (ee). Sclerotized inner tube (fig. 113, S.I.T.) with ventral margin bearing only a very short and low bulge, hence most of margin straight and paralleling dorsal one to level of dorsal spur. Female. Caudalmost large bristle on ventral lobe of sternum 7 (fig. 103, 7 S.) approximately equidistant from dorsal and ventral members of pair preceding it. Lobe at ventrocaudal angle of 8 T. relatively long and acute; about 2-3 times as long as broad at middle. Glandula vaginalis (G.VG.) long and narrow, single (not preceded by a companion-‘‘gland’’). Mesal genital ridge of tergum 8 (M.R.8) with anterior Margin sinuate at point of branch of subhorizontal arm; portion beneath fork convex, so is that of upper members of fork. Anal stylet (A.S.) long and narrow; nearly 6 times as long as broad at apical }; slightly constricted at middle; but margins essenti- ally subparallel. ILLUSTRATIONS 103. Spermatheca, genitalia and anal 107. Lumacaudate process segments 108. Processes of clasper D 250 ROBERT TRAUB 104. Head and prothorax (3) 111. Distal arm of sternum 9 105. Lateral metanotal area (3) 113. Apex of aedeagus ComMMENT. This species, known in Malaya from 11 specimens, was only found in two localities, both in Perak. Eight males were taken at the first area, viz., the limestone forest a few miles west of Ipoh at 1100-1300 ft elevation. The species was collected here only on 4 occasions, always from Lariscus insignis, and at times accompanied by M. dvyadosa, which was more common on these ground-squirrels in the area than was M. calcavata. The second area was on Maxwell Hill, Gunong Hijau, 3000-4200 ft elevation, where 3 specimens (1 g, 2 9) were taken from 2 Lariscus. It seems likely that Laviscus will prove to be the most common host of this flea. The lack of records of M. calcavata in the well studied dipterocarp- forested hills near Kuala Lumpur is notable. It is of interest that this species apparently occurs in Sarawak as well. Mr Lim Boo Liat collected 19 fleas from Lariscus 18 miles south of Kuching, of which 17 were males of the new species next described, 1 9 was M. phangi tana and I 9 was inseparable from Malayan calcarvata at the species level, including the long labial palpi and terminal maxillary palpal segment and the internal genitalia, etc. The name calcarata is derived from the Latin term calcar, meaning a spur, and refers to the conspicuous spiniforms on the male sternum 9. g. Medwayella batibacula sp. nov. TYPE MATERIAL. Holotype g (B-84199) ex Laviscus insignis; EAST MALAYSIA: Sarawak, Kuching, Bukit Stigang, Kampong Pangkalan Kuap; 27.XII.1968; Coll. Lim Boo Liat. Paratypes as follows: 1x g with same data as holotype, but 26-28.XI1.1968. 4 Sarawak, Tijirak, 19 mi S of Kuching; 7-12.1.1969; Coll. Lim Boo Liat. 1 ¢ Sarawak, Pueh; 7.VI.1958; Coll. T. C. Maa. Holotype (U.S.N.M. number 71607) and 1 paratype in U.S. National Museum. Paratypes in collections of Bishop Museum, British Museum (Natural History), the Canadian National Collection (Ottawa) and the author. Diacnosis. Near M. calcavata and agreeing with M. loncha in same respects as those cited in the previous description (number 8). Separable from M. loncha by at least diagnostic features numbers 1-3 and 5-9 cited for M. calcavata. Distinguish- able from M. calcarata as follows: (1) Labial palpus not extending beyond apex of procoxa instead of reaching tip of trochanter. (2) Segment 4 of maxillary palpus only 1-6 times as long as third, not 1-75 times. (3) Inner dimensions of lateral metanotal area (fig. 106, L.M.) virtually as high as long instead of being definitely longer than high as in M. calcarata (fig. 105, L.M.) and M. dryadosa (fig. 5). (4) Third vinculum (V.C.3) with dorsal surface sinuate and strongly upcurved at apical 4, the link-plate usually lying at an angle of about 45° except for distal curve into point of articulation in latera metanotal area. In M. calcarata, V.C.3 is quite flat dorsally for about 4 its length before curving towards L.M. and its axis is nearly horizontal, much as in M. drvyadosa. (5) Margin of lobe at cephaloventral corner of mesepister- num virtually truncate instead of definitely sinuate as in the other 2 species (fig. 9, MPS.). (6) With a distinct bulge on ventral margin of manubrium (w in fig. I, g.v. & NEW TAXA OF FLEAS 251 cf.) immediately anterior to sinus (v) near basal } instead of that region (w) being quite flat (even though sinus may be deep) asin M. calcarata and M. dryadosa (fig. 18, MB.). (7) Movable finger (fig. 10g, F.) with caudal margin somewhat convex proxi- mally, e.g., at point q, at level of apex of conical process (C.P.), instead of quite straight as in M. calcarata (fig. 108). (8) Dorsal margin of process of P. quite straight, lacking subapical sinus. (9) D.A.g (fig. 112) with ventral (caudal) margin (cf. fig. 3, qq-u3) curving dorso-apicad at an angle of 45°, commencing just above point q3, somewhat distad of level of subapical lobe (SUB.L.), and resembling a flattened arc. Tip (qq) of apical extension (EXT.9) only slightly upturned. In M. calcarata (fig. 111) the dorso-apical curve commences more distad, at level of third spiniform from tip, and apex of EXT. is so upcurved that its tip (qq) is at right angles to longitudi- nal axis of arm, (10) Semimembranous portion of dorsal margin of D.A.g9, above apex of supramedial flap (SUP.FL.) arcuate and joining EXT.9 at apex instead of being distally excised. (11) Sub-rhomboidal tanned portion of quasi-crochet (fig. 115, Q.C.) proportionately much longer and narrower, viz., more than twice as long as broad instead of 1:7 times. (12) Apicoventral corner of Q.C. produced into a short nubbin instead of heing squared. (13) Body of crochet (B.CR.) much broader sub- dorsally, its breadth there subequal to maximum breadth of phylax (PHY.) instead of being only } that of PHY. (14) Securifer with groove (GRV.) broader at depth of curve than subapically (both dorsally and ventrally); breadth of groove never approaching that of base of upper arm (U.A.) instead of exceeding it throughout. (15) U.A. broad for most its length; length of basal 3 only equal to 2:5 times its breadth instead of more than 4 times. (16) Lower arm (L.A.) of securifer with apical margin angulate, not slightly concave. (17) Phylax (PHY.) proportionately narrow- ef, viz., about 3-4 times as long as broad at maximum level (near base) instead of 2-3 times; its axis sinuate instead of straight for basal 2. (18) Sclerotized inner tube (S.I.T.) with dorsal spur arching dorsad but recurved towards axis of tube; its arc removed from $.I.T by a distance equal to 4-5 times breadth of base of spur. In M. calcarata the spur parallels tube for most its length and is close to it, the distance separating them less than twice basal diameter of spur. (19) S.I.T. subapically dilated instead of being of uniform diameter to apex. (Female unknown) DESCRIPTIVE NOTES. Differences from M. dryadosa are also included in the above diagnoses, and additional ones follow. Squamulum horizontal. Sternum 8 with cuadal margin slightly concave except for well rounded corners. Lumacaudate process (fig. 116, LUM.) neither well developed nor demarcated ; lobate portion coni- cal, with ventral margin commencing at level of 2 relatively long, stout ventro- marginal bristles; length from here twice breadth at midline; with only about 16 close-set bristles, none of which truly spiniform, in 5-6 irregular rows. Movable finger (F.) with a mesal suture running from apex of stiva to middle of tip of F. (fig. 110 and approximating p-—n in fig. I); another mesal seam extending from base of distal fringe (D.FR.) transversely across F. to apical } of anterior margin. Distal arm of sternum g about 8 times as long (ii-qq) as broad at level (jj-uu) of transverse sclerotization (fig. 112, T.S.). Upper portion (bb-ff) of posterior margin of proximal arm of sternum g flat, subparallel to that of anterior margin (aa—dd). 252 ROBERT TRAUB ILLUSTRATIONS 106, Lateral metanotal area (3) 112. Distal arm of sternum 9 10g. Processes of clasper 115. Apex of aedeagus 110. Mesal aspect of F. 116. Lumacaudate process ComMENT. All of the known specimens, 17 males representing 9 collections, were from Lariscus insignis in the same general area and habitat in Sarawak. (Usually female fleas are more common than males, at least in collections, and it is unprece- dented to obtain such a large series of males in the absence of females.) | The superfi- cial similarity to M. calcarata from Perak in Malaya, and which might also be a Lariscus flea, is quite striking. The differences apparent on critical study, however, cited above, are too great for subspecies and indicate the two are sibling species. This belief is supported by the fact that a female virtually indistinguishable from Malayan M. calcarata was collected from the same host and locality as the type series of M. batibacula (along with a single female of M. phangi). It is highly unlikely that the ength of the labial palp and all the other significant characters noted would vary only in the Sarawak male, while the female remained unchanged in the course of evolution. It seems most probable that M. calcavata occurs both in Sarawak and Malaya, parelleling the distribution of M. phangi and M. robinsoni. Further collec- tions are urgently required to clarify this point and the many other questions posed by the chronic shortage of specimens of Siphonaptera for study from sundry hosts and habitats throughout most of the Asiatic-Pacific area. The name batibacula is derived from the Latin words for thorn or spine (batus) and staff or rod (baculus), and the “‘thorny staff” refers to the distal arm of sternum 9. 10. Medwayella veruta sp. nov. TYPE MATERIAL. Holotype 3g, allotype 2 (B-19165-1) ex squirrel nest; EAST MALAYSIA; Sabah (N. Borneo), Mt. Kinabalu, Ranau; elev. 1500 ft; 31. VI1.1953; Coll. R. Traub. Paratypes as follows: 5 g, 3 2 with same data as holotype. 24,19 ex Sundasciurus low; loc. cit., but 13. VII, 6.1X.1953. 1 9 ex Tupaia sp.; loc. cit., but 2.VIII.1953. 1d ex Ptilocercus low; loc. cit., but 13. VII.1953; Coll. R. Traub & H. D. Newson. 2 ex Tupaia sp.; 18 mi S of Tenom Kg Bara Jumpa; 2.I1.1963; Coll. J. A. Bullock. Holotype (U.S.N.M. number 71608) and allotype in U.S. National Museum. Paratypes in British Museum (Natural History) (1 pair) and in author’s Collection. Diacnosis. Agrees with M. loncha in the same respects as those listed for M. calacrata (q.v. number 8) and thereby instantly separable from M. dryadosa and allies, while the characteristic shape of D.A.g of M. veruta (fig. 120), M. batibacula, etc., readily distinguishes this group from M. phangi (fig. 68) and its relatives. Readily separable from M. loncha as follows: (1) Movable finger (fig. 119, F.) not nearly as foreshortened and broad subapically, viz., about 3-75 times as long (s—n in fig. I, g.v. & ef.) as broad (j-k) below distal fringe (D.FR.); not merely thrice (fig. 126). (2) Bristles of D.FR. with bases forming an oblique line instead of a straight line paralleling longitudinal axis (s-n) of F. (3) F. with convexity of caudal margin (q) near P. exceeding that of cephalic margin (i) near conical process (C.P.) instead of NEW TAXA OF FLEAS 253) being flatter than that of opposite margin. (4) Posterior margin of P. (t-u) somewhat convex below level of subapical long bristle instead of being straight to near sinus (v) near fulcral sclerite (F.S.). (5) Anterior (dorsal) margin of D.A.9 (fig. 120) with portion distad and proximad of spur (SUB.L.) (cf. fig. 3, e3-f3 & c3-kk) lying in different planes, D.A.g being much broader proximad to SUB.L. than distad; instead of anterior margin forming a straight line (kk-qq) interrupted by spur. (6) Distance (e3-qq) from SUB.L. to apex of segment (EXT.g9) shorter than in M. loncha, e.g., distance (b3-qq) from dorsal (anterior) apex of transverse sclerotization (T.S.) to tip of arm is 3-7 times distance (e3-qq) from upper margin of base of SUB.L. to apex of EXT.g, instead of only 2-7 times. (7) SUB.L. proportionately closer to upper limit of mid-microspiculate area (M.MSP.), viz., distance (c3-kk) between them subequal to length of bristles of caudomarginal submedian group (CM.G.B.) instead of clearly exceeding it. (8) Anterior margin of D.A.g (e3-qq) extending to apex of arm even though lightly tanned distally, instead of curving caudad subapically (f3). (9) Deltoid flap of aedeagus (fig. 114, DEL.FL.) only twice as tall as long instead of 2-4 times. (10) Tanned portion of quasi-crochet (Q.C.) proportionately longer and narrower, thrice as long as broad at middle instead of 1-3 times. (11) Unique in that phylax (PHY.) is curved caudad apically so that there is a short, broad lobe at dorsocaudal corner. In other Medwayella the caudal margin is straight for the upper 3 and hence there is no lobe. (12) Phylax narrower than in M. loncha and more constricted subventrally, although both are broadest near middle. Length 3-2 times midgirth, and 4 times that at narrowest point, instead of 2-8 and 3-5 times. (13) Groove (GRV.) of Ford’s sclerite widest at middle and narrowest near dorsal apex, instead of vice versa. (14) Upper arm (U.A.) of securifer with dorsal margin sinuate —the trough proximal and height of convexity just apicad of middle, whereas in M. loncha, dorsal margin is evenly concave to near apex. (15) Lower arm (L.A.) with upper margin convex or peaked near apex, not fairly straight (though microserrate). (16) Dorsal spur of sclerotized inner tube (S.I.T.) large, sloping cephalodorsad at angle of 60°, then recurved over tube and apex connected with it; resembling a triangle with a hollow center, whose base equals diameter of S.1.T. and whose altitude is about 3 that diameter. In M. loncha the spur commences more vertically, is apically recurved and presumably lacks elements returning anteriorly to S.I.T. (17) Subventral sinus of 9 sternum 7 (figs 117, 124) open dorsally, there being no true notch, whereas in M. loncha (figs 118, 130) caudal margin above ventral lobe is inclined to an angle of about 60° until near apex of lobe, so that there is a notch. (18) Caudal margin of 7 S. (fig. 124) with a short but distinct median lobe where it recurves cephalodorsad. In M. loncha (fig. 130) curve is effected without production of a lobe. (19) Perula of bursa copulatrix (P.B.C.) relatively shorter and broader, viz., only about 2-2 times as long as broad at lower }, instead of 4 times. (20) Witha distinct “gland” between glandula vaginalis (G.VG.) and orifice of vagina; this lacking in M. loncha. DESCRIPTIVE NOTES. The characters cited above also differentiate M. veruta from M. dryadosa and additional points follow. Lateral metanotal area with quadrate untanned area taller than long. Squamulum horizontal and more than 5 times as long as broad at middle. Spiracular fossa of metepimere sagittate but with ventral 254 ROBERT TRAUB margin straight and sloping somewhat dorsocaudad; dorsal margin slightly inclined from horizontal for first } and then sloping ventrocaudad towards apex; about 1-3 times as long as tall at maximum level. Spiracular fossa on 2 T. 1-6 times as long as broad; 2-3 timeson6T. Lower antepygidial bristle about 2-5 times length of upper one in male; 2-4 times in female. With dorsal lower modified bristle (fig. 124, L.M.B.) a smaller, thinner version of upper antepygidial bristle; L.M.B.-2 resembling long bristles of adjacent row. Lower lobe of 7 T. (L.L.7) below A.B. blunt. Male. Ventral extension of 8 SPC. with height subequal to or exceeding diameter of horizontal section. Lumacaudate process (LUM.) with densely bristled area about 2-4 times as long as high; apically broadly or obtusely rounded. Conical process (fig. 119, C.P.) about 5 times as long (e-h) as broad (f-g). Dorsocaudal corner of F. scarcely produced into a snout, e.g., stiva (STV.) poorly developed, in that distance (l-p) from anterior margin near apex (1) to tip of stiva (p) is 6-5 times length of stiva (p-o/k). Distal fringe (D.FR.) consisting of 4 or 5 bristles; lowermost the thinnest. Sinus of ventral margin of manubrium (v) long and shallow; the bulges (u and w) flanking it correspondingly short. Upper sinus (aa—cc) of anterior margin of P.A.g much deeper than lower sinus (cc—dd), which is virtually flat. D.A.g about 8 times as long (ii-qq) as tanned region is broad (uu—jj). With about 5 or 6 marginal spiniforms on caudal margin of D.A.g9, of which lowest 4 are usually longest and stoutest, and distalmost of long ones generally at level of SUB.L. (d3-q3). Mid- microspiculate area (M.MSP.) highly convex and apically bulging above D.A.g nearly as much as SUB.L., but cavity of receptacle with D.A.9 correspondingly fairly shallow. Lateral laminae of aedeagus and middle lamina relatively broadly yoked at tip of apex of apodeme, margins of the plates parallel until summit. Sinus of hood (HD.) deeply excised, reaching to near apex of B.CR. Upper margin of deltoid flap (DEL.FL.) evenly but shallowly concave. Vertical portion of phylax (PHY.) slightly sinuate. Female. Spermatheca (fig. 123) with bulga (B.) bearing a short but distinct peak. Ventral lobe of sternum 7 short and broad; scarcely longer than tall. Anal stylet (fig. 122, A.S.) scarcely narrowed from base to near apex; about 4-6 times as long as broad at middle. Tergum 8 (fig. 125, 8 T.) broadly ovate at ventrocaudal angle, lacking a distinct lobe. ILLUSTRATIONS 114. Apex of aedeagus 122. Anal lobes and stylet (9) 117. Spermatheca and genitalia 123. Spermatheca 119. Processes of clasper 124. Sternum 7 (2) 120. Distal arm of sternum 9 125. Genitalia (9) 121. Sternum 8 (3) ComMENT. As can be seen from the type data, M.verutawas found on Sundasciurus lowi, on Tupaia sp., and in the nests of squirrels on the foothills at the base of Mt. Kinabalu. In addition, the author and H. D. Newson collected a male from a nest, occupied by a pen-tailed shrew (Ptilocercus low?), in a branch ina tree in the same area. It is worthy of note that this species was also collected on Tupaia tana in Sarawak, in Division III at Nanga Balleh, near Kapit (R.T.). This constitutes only the second NEW TAXA OF FLEAS 255 known instance of a North Bornean indigenous flea occurring in Sarawak (and apparently represents a distinct subspecies). (The other species is Stivalius mjobergi Jordan, 1926, described from Mt. Murud, which is just south of the border of N. Borneo.) However, the differences in the Siphonapteran fauna of Sarawak as compared to North Borneo, which superficially appear to be striking, may prove to be more apparent than real when the mountains of Sarawak are studied, and as more of the lower elevations in North Borneo are surveyed. This is discussed further below (Section A.14), and in the third article in the series. The name veruta is a Latin term meaning armed with a spear or javelin and refers to the rod-like extension of the spined distal arm of sternum 9. 11. Medwayella loncha (Jordan, 1926) NEW COMBINATION Stivalius lonchus Jordan, 1926. Novit. zool. 33:390, figs 11, 12; Costa Lima & Hathaway, 1946, Pulgas: 327; Smit, 1958, Bull. Brit. Mus. (Nat. Hist.) Ent. 7(2) :3. To date this species has been known only from the single specimen collected by Dr E. Mjéberg ex Tupaza sp. in Sarawak, Mt. Poi, 5000 ft elev. described by Jordan in but 4 lines and illustrated by one figure showing the upper portions of the movable finger and distal arm of sternum g. The published material is inadequate for recog- nition of the species, and in fact is misleading, for instead of the movable finger F. being ’’not essentially different,” we now are aware of a sufficient number of species in the group to realize that F. zs distinctive. The collection of M. loncha and the telated new species, M. batibacula, by Lim Boo Liat in Sarawak, and the discovery of another allied new species, calcarata, plus access to the type specimen, have ren- dered clarification of the characteristics of M. loncha possible, as well as description of the hitherto unknown female. Diagnosis. The salient features of M. loncha have been listed in the comparisons made above in the descriptions of the new species M. calcarata and M. veruta, q.v. It is readily recognizable in the g by the relatively short, broad F. (fig. 126), which is scarcely more than thrice as long (from apex of immovable process P., cf. fig. 1, ns) as broad at narrowest level (immediately below level of distal fringe, D.FR., or j-k). It is further characterized by a D.A.g (fig. 127) bearing a distal, vertical “‘spur’’ (EXT.9) instead of being curved to form an apical lobe (cf. fig. 20, AP.L.) etc., and by an aedeagus (fig. 128) with a groove (GRV.) between alpha portion (ALPH.) and base of upper arm (U.A.) of securifer. The short, broad quasi-crochet (Q.C.) is diagnostic—region distad of phylax (PHY.) is only 1-3 times as long as broad because of markedly convex dorsal margin (but upper portion lightly tanned). Female distinguishable by the following combination of characters: (1) Sinus of sternum 7 (figs 118, 130, 7 S.) very broad apically, and already diverging markedly from ventral margin at base. (2) Perula of bursa copulatrix (P.B.C.) elongate-ovate, but with anterior margin quite straight. (3) Paragenital morion so lightly tanned as to be virtually inapparent. (4) Lacking a ‘‘gland”’ caudad to glandula vaginalis (G.VG.). (5) Mesal genitalic ridge of tergum 8 (M.R.8) with anterior margin sinuate; sub- horizontal fork aborted; anterior branch of fork short and narrow. (6) Ventrocaudal corner of 8 T. with a short, broad lobe. 256 ROBERT TRAUB ILLUSTRATIONS 3. Distal arm of sternum 9 129. Sensilium and anal segments (9) 118. Spermatheca and genitalia 130. Sternum 7 (@) 126. Processes of clasper 131. Sternum 8 (3) 127. Distal arm of sternum 9 132. Spermatheca 128. Apex of aedeagus REcoRDS. 2d, I 9 ex Sundasciurus lowi; EAST MALAYSIA: Sarawak, Tijirak, Ig mi S of Kuching; 28-30. XII.1968; Coll. Lim Boo Liat. 1, 1 2 ex Tupaia tana; loc. cit.; 26.XII.1968. CoMMENT. It seems significant that this species was collected only on Sundasciu- vus and Tupaia but not on Laviscus in the same area, and that the latter were para- sitized by M. batibacula. The material herein described and figured as M. loncha agrees closely with the type specimen in the British Museum. 12. Medwayella javana (Jordan, 1933) NEW COMBINATION Stivalius javanus Jordan, 1933, Novit. zool. 38:355-356, figs 76, 77; Costa Lima & Hathaway, 1946, Pulgas: 327; Smit, 1958, Bull. Brit. Mus. (Nat. Hist.) Ent. 7(2): 3. Another species known heretofore only from a single g (and 3 2), M. javana was described from specimens collected from Tupaia javanica occidentalis (holotype 8), “Rattus bukit temmincki” (probably R. fulvescens temmincht) (2 2), and Callosciurus n. nigrovittatus (I 9) at ‘‘Tjiboeni, Bandong, W. Java’ (today known as Tjibuni, Bandung). The present descriptive notes and illustrations are based upon a pair collected from Tupaia javanica by J. M. Stusak of the B. P. Bishop Museum, 30.1X. 1965 at Tjibodas, Java, 5200 ft elev., and which agree quite closely with the original description and photographs of the type material. Dracnosis. Some cardinal characters have already been noted in the description of M. arcuata, q.v. Salient points are: (1) Distinctive amongst known Medwayella in that the movable finger (fig. 133, F.) is long and narrow, viz., portion distad of apex of immovable process P. (cf. fig. 1, rn) is about 4 times as broad at level of apex of conical process (i-q) and 5 times as long as broad at middle of subapical constriction (j-k). (2) The only other species besides M. avcuata in which dorsal margin of distal arm of sternum 9g (fig. 124) is deeply arcuate subapically (ARC.) instead of either bearing a true notch (figs 20, 21) or being homolate (figs 68, 81). In M. javana, ARC. is fairly shallow, i.e., 5 times as long (cf. fig. 1, mm—pp) as deep at trough. (3) Apical lobe (AP.L.) extending more dorsad (cephalad) than subapical lobe (SUB.L.). (4) Aedeagus (figs 95, 96) of the M. robinsoni type in that thumb (THM.)is in the “cocked’’ position and lower arm (L.A.) of securifer is broad and apically blunt, not acuminate. (5) Height of THM. slightly less than breadth of apex. (6) Upper arm (U.A.) of securifer distally acute but broad to basal half. (7) L.A. with lower margin about 1-8 times as long (from edge of body of crochet, B.CR.) as broad at middle; dorsal margin angulate and sloping from middle towards ventrocaudal corner at angle of about 45°. (8) Tanned portion of quasi-crochet (Q.C.) fairly short and broad, about 2-6 times as long (from edge of phylax, PHY.) as broad at middle; subapically somewhat dilated and apex obtuse. (9) Sclerotized inner tube (S.I.T.) well upcurved NEW TAXA OF FLEAS 257 beyond level of long, straight dorsal spur. (10) Lumacaudate process (fig. 135) well defined ; narrow—with only 1 spiniform bristle at apex and no more than 3 or 4 per row in breadth; length (from anterior spiniform) about 1-75 times breadth at middle. (rr) Sternum 7 of 9 (fig. 40, 7 S.) unusual in that ventrocaudal lobe is sinuate apically. (12) Ventrocaudal sinus of 7 S. short, dilated apically. (13) Perula of bursa copula- trix (P.B.C.) pyriform, with expanded portion ventral, but anterior margin quite flat. (14) Paragenital morion (PG.M.) with a conspicuous tanned rod-like element but- tressing P.B.C. (15) Mesal genital ridge of tergum 8 (M.R.8) with anterior and medi- an margins sinuate; lower branch below fork short, thin and lightly tanned. (16) Caudoventral lobe of 8 T. fairly short, broad at base and with an angle of about 35° at tip. (17) With glandula vaginalis (G.VG.) long and thin; no other “glands” caudad of it. ILLUSTRATIONS 40. Genitalia (9) 134. Distal arm of sternum 9 95. Apex of aedeagus 135. Lumacaudate process 96. Sclerotized inner tube 136. Sternum 7 () 133. Processes of clasper 137. Spermatheca, genitalia and anal seg- ments 13. Medwayella rhaeba (Jordan, 1926) NEW COMBINATION Stivalius yhaebus Jordan, 1926, Novit. zool. 33:389, figs 9, 10; Jordan & Rothschild, 1926, Sarawak Mus. Jour. 3 (part III) (10): 290, figs 6, 7; Costa Lima & Hathaway, 1946, Pulgas : 327; Smit, 1958, Bull. Brit. Mus. (Nat. Hist.) Ent. 7(2) : 3. This species is still known only from the single pair from “‘Sciwrus brookst” (lapsus for the squirrel now called Sundasciurus brooket) on Mt. Dulit (Sarawak) and from the abbreviated descriptions and figures by Jordan in 1926 and Jordan & Rothschild in1926. The diagnostic features mentioned in the present article are based primarily upon photographs of the holotype (g) and allotype in the British Museum. Diacnosis. Apparently unlike any of the other described species in the shape of the movable finger (fig. 138, F.). At level of base of crochet process (CR.P.), breadth of F. exceeds that at apex, yet narrowed to half that breadth at apical third (measuring from base of CR.P. or summit of immovable process P.). FF. rapidly broadening above that level, largely due to oblique caudal margin slanting at 45° to tip of stiva. Distal fringe (D.FR.) of 3 stout bristles inserted immediately below apex of stiva. Distal arm of sternum g (D.A.g) diagnostic by virtue of denticulate tip or nubbin at anterodorsal angle. Other characteristics are mentioned in compari- sons made with M. phangi (number 5 above). The heavily sclerotized, arched, horizontal section of the mesal genitalic ridge of 9 sternum 8 (fig. 139, M.R.8) seems to be unique amongst known Medwayella. M. rhaeba agrees with M. phangi in being a species with a grooved Ford’s sclerite (GRV.), with a narrow, upturned quasi-crochet (Q.C.) and with a homolate D.A.g (cf. figs 68, 71, M. phangi). However, the differences are striking, as has been pointed out above. 258 ROBERT TRAUB ILLUSTRATIONS 138. Modified abdominal segments (3) 139. Modified abdominal segments (2) 14. GEOGRAPHIC AND HOST-DISTRIBUTION OF MEDWAYELLA In Mataya A consideration of the ecological, geographic and host-distribution of Medwayella is worthwhile even though virtually nothing is known about the flea-fauna of the eastern districts of the Indian subcontinent (such as Assam, Manipur, the Chittagong Hills, etc.) and of the bulk of the Indonesian Archipelago (i.e., except Sumatra and Java, for which we have sparse information) and despite the fact that data are mini- mal for the northern and eastern parts of the Indo-China subregion. Thus, even though collections from the peripheral regions of the range for Medwayella are sorely needed, it is possible to make some generalizations which seem significant. The zoogeographical observations and implications in the main are discussed in the third article in the series, and what follows here will largely deal with the types of hosts infested and where they were found. Thirteen species of Medwayella are mentioned in this paper. Six of these are known from Sarawak, five from Malaya, two from Sumatra, two from Thailand and one each from Sabah (N. Borneo), Java and Indo-China. Of the 13, M. robinsont is known from four of the areas listed (Malaya, Java, Sarawak and Thailand), and two species (M. calcavata and M. phangi) occur in both Sarawak and Malaya. (The fact that some are found in several countries, of course, meant that the numbers just cited will add up to more than 13.) This recorded distribution, to a great extent, reflects the degree of collecting done per area, but even so, certain trends suggested by the data are valid. Thus, while as yet undescribed species undoubtedly exist in Indo- nesia, and two such are known from the Philippines, and the genus probably occurs in eastern India and Burma, sufficient field-work has been done in Indo-China and Thailand to show that the number of endemic species of Medwayella is well below that for Malaya and Borneo, which seem to be the center of development of the genus. Of the countries mentioned, the only one for which there are sufficient data for detailed discussion of hosts and distribution in Malaya, and the available records are summarized in Tables 1 to 6. (It is reiterated that the numbers of hosts cited in Table 2 as having been examined includes only our own collections, and thus these do not represent the hosts trapped or shot by our colleagues in Kuala Lumpur. The numbers of fleas, however, do include all available specimens.) The sciurids listed as tree-squirrels (T) in Table 2 are essentially arboreal and nest high in the trees, but do come to the ground on occasion. The ground-squirrels and ground-dwelling forest-rats (marked G) spend most of their time on the forest-floor 6 It should be noted that the geographic names and expressions used in this article are those that havea fairly specific connotation in zoogeography, history or usage. Thus, there are no political considerations in the term “Indo-China,” which is an established name for a zoogeographical subregion. Similarly, toa naturalist, ‘‘Malaysian” includes lower Burma and some of the Indonesian Islands, and hence this term has been avoided when speaking of the country formerly called Malaya, or of the part of Borneo previously under British rule. The expression ‘‘Malayan”’ as used here, refers to the Peninsula and does not have the zoogeographic connotation of a Malayan (or Malaysian) subregion. 259 LAS FLE TAXA OF NEW 19z1 s[eqoL, %oorF olor %oor ir %oor zh Y%oor 69 %oo1rF of: %I cr Mr ZI e 6 e 9 %1 II DAS ze e 6 %61 £61 e b onIZ Zz DES ze % Bz ° + %oz goz br 6€1 e 9 %6 z6 % ‘ON 1UOSUIGOAL %I wey} sso] = @ Suljeamp-punoiy = 5 = - %rg gf = = FAS) %oo1 ing GAYE € %9 + %SI o/ - - fot /O = Gat %S - %6€ Lz ° o. . . y’? . Yo ON % ON % ON % Suyjenp-se1l = J I ‘ON p,wrexq DIDADIIVI my asunyd vsopoaAap soroads nyjadvmpapy jo saseyusoied pue siaquinN—seo[q zol €z 6Sz OIL ge ber Joquinyy UWINWIIUTTY o + +++ + YOO Oo usin 15! WS) OIG) ts SUNUA] SNANVISDPUNGS (L) + tal Et eS syiqeH stuadyna shmowmasqr S[eI0L (7oq90) ° SnjopnvIyvy snanwsouryy * SIUSISUL SNISUADT * (punorg) S}EY 4S9IOT - (petzosueos) s7eX] 4S010,T Adour niogn I sys viwdn I upunjaj99vue sdovmp [ 40] “S 9 (JSeN [ermbs Jeer0qry) snandqry snanvasppuns ysonarg SNANVISONJDI snyojou snaniasoyyy 7 SNIDIPULOATU SNANLISONVI snavayjhsa snanvgsoyjv7Z sdanvupa SNANIISOYDI auieN s}soP] VIOSNINGg NVAVIVJ! HL NO SVITA PTTAA PMCAW FAM AO NOLLNAIALSIG-LSOH Z@ ATGVL Li gI Cr “FI €r 6 260 ROBERT TRAUB and nest below the surface, as amongst the roots of trees, but at times climb on tree- trunks and low trees and shrubs. Twpaia are active on the ground and are also at home on trees, at least on lower limbs. Their common name, tree-shrew, therefore is at least partially correct (since most mammalogists no longer regard them as shrews). The Tupaia and the scansorial forest-rats and the squirrels like Sundasciu- rus tenuis,S.lowi and Tamiops maclellandi, which can be seen and/or collected both on trees and the forest-floor, are designated as ““T +G,” for “tree and ground.” Flying- squirrels, e.g., Petaurista and the giant tree-squirrel, Ratwfa, were collected too seldom for discussion, but no characteristic species of fleas have been found on the 15-20 specimens of each examined by me in Malaya, while “strays” like M. vobinsoni were rare. (It is interesting that in the Himalayas, Petaurista and Hylopetes flying-squirrels are infested with several fairly specific genera and species of fleas of Indo-Malaysian affinity and that in Formosa Petaurista regularly bears fleas, but of a palearctic species. An endemic genus has also been found on Hylopetes in N. Borneo.) Several points are readily apparent from the data in Table 2. For example, of the five species listed, only M. vobinsoni has been taken with any frequency on Callosciu- TABLE 3 HosT-DISTRIBUTION OF FIVE SPECIES OF MEDWAYELLA FLEAS ON THE MALAYAN PENINSULA (SUMMARY) SPECIES OF MEDWAYELLA HOST DRYADOSA PHANGI LIMI CALCARATA |ROBINSONT I. ARBOREAL SQUIRRELS II. SEMI-ARBOREAL AND GROUND -SQU TRRELS III, TUPATA IV. FOREST-RATS TOTAL % 100% 100% t1 00% 100% too TOTAL NUMBER OF FLEAS 129 68 42 11 963 NEW TAXA OF FLEAS 261 rus, a genus of tree-squirrels. In fact, of the remaining four species, only M. phangi and M. limi were ever collected on an arboreal squirrel and then only once each, i.e., two of the former from C. notatus and one from C. caniceps for the latter. In contrast, 65 % of the records for M. dvyadosa and 84% of those for M. limi were from Rhinosciu- rus laticaudatus, a ground-squirrel. Nearly half of the M. p. phangi were collected on the tree-shrew, Tupaia glis, but 39 °% were from the small Sundasciurus squirrels, which are frequently on the forest-floor. The second most prevalent host for M. dryadosa was the ground-squirrel, Lariscus insignis, accounting for 15°% of those collected. All of the known Malayan M. calcavata likewise were from that host. However, the numbers involved (11 specimens) are too low for certainty regarding host-specificity of M. calcarata, although they represent seven different collections. The apparent dearth of Medwayella reported from the ground-squirrel Dremomys seems noteworthy, but is misleading in a sense, since this squirrel is montane and the Medwayella being discussed are rare or absent above 4000-5000 ft. The host-range of M. robinsonz is obviously far greater than for other members of the genus, e.g., it was collected at least once on each type of host. The highest percentages, however, are for the arboreal and the partially arboreal squirrels, rats and tree-shrews. These data are summarized in Table 3 below, which compares the numbers and percentages of these fleas which were from arboreal squirrels with those from squirrels that divide their time between trees and the ground, and with those from Tupaia and rats. From this table, it can be seen that 100% of M. dryadosa were from non- arboreal species (accounting for the name of “‘hater of tree-dwellers’’), and that this same tendency was exhibited for M. p. phangi and M. limi (97°% and 98% from ground-dwelling or semi-arboreal hosts respectively). The tree-shrews, Tupaia, were “good”’ hosts for at least three of the five species, particularly M. phangi and M.robinsoni. Various species of forest-rats, such as the scansorial Rattus (Lenothrix) canus and R. (Maxomys) cremoriventer and the ground-dwelling R. (Lenothrix) rajah, R. (Leopoldamys) sabanus and R. (Stenomys) bowersi at times carried one or two Medwayella fleas, generally M. vobinsoni, but occasionally M. dryadosa, and rarely one of the others. When discussing host-relationships, it is always important to consider infestation- rates. Unfortunately, the data here have limitations, even though for our own collections we have records of the numbers of animals which had no fleas at the time of examination. As mentioned above, since the collections were made to obtain living mammals for studies on scrub typhus, etc., many of the rats had been in the traps for hours, or even more than a day, before being placed in a bag for subsequent examina- tion. Such excited hosts, perhaps drenched by rain, soon lose their fleas, while the ubiquitous and innumerable ants promptly ruin mammals killed by other types of traps. Under such condition, it is impossible to obtain reliable “‘flea-indices’’ or infestation-rates, and it is not surprising that the average number of Medwayella fleas per mammal in the tables above at times would be less than one, even if the “minimum number examined”’ were the actual total number. However, it is impor- tant to note that nearly all of the tree-squirrels and many of the Tupaia and ground- squirrels had been collected by shooting and the specimens were then immediately bagged. Even so, the over-all index was frequently below one flea per host, though 262 ROBERT TRAUB an occasional squirrel would have as many as six or even ten Medwayella. No pattern could be determined for the marked variations in numbers noted for squirrels shot in the same locality and on the same day, although I suspect that factors such as sex (lactating females versus males, for example), age, season, etc., all play a role. More data are required in this regard. Some idea of the infestation-rate can be obtained from the “‘collection rate” or the number of times the particular fleas were collected from a specific type of host (e.g., the numbers of hosts ‘‘positive’’ for that species of flea). These data are shown, where significant or relevant, for the four new species of Medwayella and for M. robinsont (Table 4). These figures also are valuable because they offset the skewing effect of a single unusually large collection, which, while unrepresentative, may exert considerable influence on the mean or total number reported. TABLE 4 THE NUMBERS OF SPECIMENS OF CERTAIN SPECIES OF MEDWAYELLA AND THE NUMBER OF TIMES THESE SPECIES WERE COLLECTED FROM SPECIFIC HOSTS IN MALAYA Species of Medwayella Host dryadosa phangt limi calcarata vobinsont Fleas Coll’ns Fleas Coll’ns Fleas Coll’ns Fleas Coll’ns Fleas Coll’ns Sundasciurus tenuis & S. lowt I I 27 13 - - - - 202i 003) Lariscus insignis 20 16 4 4 3 3 II 7 9 6 Rhinosciurus laticaudatus 85 26 - - 36 15 - - 12 9 Tupaia glis 12 8 33 18 I I ~ - 193 82 The data suggest that the indicated association between M. phangi and Sundasciu- rus tenuis and S. lowi is a fairly close one, since this species was collected from those squirrels on 13 different occasions, even though the total number (27) is small. The same seems true for M. vobinsoni on these hosts. M. dvyadosa was relatively fre- quently taken on L. insignis and Tupaia, but in small numbers, whereas in the case of Rhinosciurus, the average infestation rate was over three (26 collections totalling 85 fleas). The facts that M. calcarata was collected from Lariscus on seven different occasions (with a total of 11 fleas) and was never taken on another host, suggest an intimate relationship, but additional data are required for verification of this point. Thus far we have been dealing with Medwayella fleas and reported collections from sundry hosts. It is also worthwhile to consider the data from the converse view, viz., the major hosts and the relative numbers of Medwayella with which each kind was infested. This is shown in Table 5, whence it can be seen that virtually all the fleas taken on each of the three species of Callosciurus were M. robinsont, and that 80% or more of the fleas from Tupaia and forest rats belonged to this species. Lariscus was most apt to be infested with M. dryadosa, although all five species were found on this host at one time or another. Of the 133 fleas reported for Rhinosciurus, 64% were M. dryadosa and 27°% M. limi. The absence of M. phangi on this host is quite striking, particularly in view of the fact (mentioned below) that M. phangi was collected in areas where this ground-squirrel occurred. NEW TAXA OF FLEAS 263 TABLE 5 THE MAIN HOSTS OF MEDWAYELLA FLEAS IN MALAYA AND THE RELATIVE PERCENTAGES PER SPECIES WITH WHICH THEY WERE INFESTED ay | [Basee CALLOSCIURUS SCIURUS SPECIES OF MEDWAYELLA NIGRO- TUPAIA FOREST | LARISCUS LATI- CANICEPS VITTATUS NOTATUS GLIS RATS INSIGNIS CAUDATUS 15 64% 4, 3% DRYADOSA . Re LIMI ROBINSONI ‘ TOTAL % TOTAL NO, OF FLEAS e = less than 1% The major collecting-localities in Malaya are shown in Table 6, while comments on the habitats in the area have been presented above (p. 207). It is apparent at a glance that M. robinsoni was found in all of the regions surveyed, and common in those wherein sufficient material had been collected. In all probability, this species will prove to be abundant in the little-studied Kelantan area and all other inland habitats below 4500 ft elevation where Callosciurus occurs. As previously mentioned, we have never collected indigenous fleas along the coastal areas of Malaya (nor along the coast of any other humid tropical country), despite fairly intensive search in some regions, and the presence of species of hosts like C. caniceps, Tupaia, R. sabanus, etc., which carry fleas when farther inland. Indigenous fleas have not been collected on the Sembilan Islands, Pangkor Island and the Lang- kawi Islands. However, the rodent fauna varies on those islands—only rats occur on the first-named, and rats and tree-shrews are present on Pangkor, while on Lang- kawi C. caniceps was the only squirrel noted, but tree-shrews and sundry forest-rats were found to be plentiful. It is my belief that it is too warm and humid for fleas (or, 264 ROBERT TRAUB TABLE 6 RELATIVE ABUNDANCE OF SPECIES OF MEDWAYELLA IN MAJOR COLLECTING-AREAS IN MALAYAN PENINSULA I. 2. a 4. 5. 6. mp 8. Kuala iE. Kelan- Species of Lumpur Subang Ipoh Gunong Coast tan Moun- Medwayella Forest Forest Hills Benom Forest Forest tains Kedah dryadosa Bahr is * ae @ ® = ig phangi R ae ae - - - @, *% limi ata = - - ba - - - calcarata - - * - - - - - vobinsoni **K ** ** ** ** * * ** #** — Prevalent ** — Fairly Common * = Occasionally Collected @ = Present, but data inadequate = Rarely Taken @ = Collected on Kedah Peak probably, their larvae) in such habitats, but the apparent absence of fleas on Lang- kawi at 2000 ft elevation is somewhat surprising, and further investigations are desirable to obtain better data in this regard, since we only examined a few rats taken at that height. The fact that M. vobinsoni and M. dryadosa occur on Tioman Island off the east coast, is probably due to the presence of a monsoon season there, which may create favorable conditions for fleas in the dry spell. If so, then M. vobinsoni may also prove to be present in the northeast coastal region of Malaya, where the climate is also somewhat different from that associated with the typical rain- forests of that country. As for the altitudinal range of M. robinsonz, it has been mentioned above that we have a few records of this species occurring (on C. caniceps) as high as 5000 ft, at the edge of montane forest, but their presence at such climes seems to be exceptional. Other species of fleas, especially the genus Macrostylophora Ewing, 1929, seem to be more characteristic of the higher altitudes, where Callosciurus erythraeus is the most abundant tree-squirrel and Dremomys the prevalent ground-squirrel. At the other extreme, M. vobinsoni has been found on squirrels in inland forest only 50 ft above sea-level. M. dryadosa is another species that was also taken in all of the localities listed in Table 6, although it never was collected as frequently as M. robinson, probably because its main host, viz., ground-squirrels like Rhinosciurus and Lariscus, are more difficult to trap or shoot than are Tupaia and Callosciurus. It seems to be most prevalent in relatively undisturbed dipterocarp forest in the foothills, as at Ulu Gombak, Ulu Langat, etc., in Selangor and at Gunong Benom, rather than in the limestone hills of Perak, the montane forest, or in the lowland, highly modified forest of Subang. The distribution of MW. phangi is quite different, although it too ranges widely over Malaya. It was rarely noted in the dipterocarp forest in the foothills of Selangor, and then only at the extreme edges, which has been secondary forest for several decades. In contrast, it was quite common in the highly disturbed forest at Subang (400-700 ft elev.), in the limestone hills of Perak (150-250 ft elev.), at Kedah Peak and at Bukit Wang Forest Reserve in Kedah. The occurrence of M. phangi on NEW TAXA OF FLEAS 265 Kedah Peak, in contrast to the other mountains studied, is of interest and is commen- tedon below. The fact that 50% of the specimens came from Tupaia and 39% from Sundasciurus tenuis and S. low is in accord with this belief that M. phangi is associa- ted with secondary forest. The absence of records of M. phangi from Rhinosciurus is noteworthy, and suggests that this ground-squirrel is primarily a denizen of un- disturbed forest, an impression that is heightened by the numbers of collections of M. limi and M. dryadosa from this host. In this regard it is worth noting that the only area besides the dipterocarp forest near Kuala Lumpur where M. limi was found was in the forests of Trengganu, at Bukit Besi, near the east coast. Although the zoogeography of Medwayella and other fleas will be discussed in the third paper in this series, attention is now called to the odd distributional pattern exhibited by sowie species of Medwayella. Three of the species, M. phangi, M. calcarata and M. robinsont, are known from both Malaya and Borneo, but, in each case, only from Sarawak, not North Borneo (Sabah). Further, two of these have been found only in the western part of Malaya and are common in the Perak-Kedah regions. “15. KEY TO THE KNOWN SPECIES AND SUBSPECIES OF MEDWAYELLA? 1 Labial palpus long, extending to apex of fore-trochanter (fig. 104). Distal segment of maxillary palpus unusually long, 1-75 times length of third segment. 4 with 3 “supernumerary” bristles (SY.) between rows II and III on head; 9 with 1 such calcarata sp. nov. (p. 247) — Labial palpus shorter, extending generally only to or near apex of procoxa (fig. 4), rarely to middle of fore-trochanter. Distal segments of maxillary palpus only 1-5 times length of third segment. 4 with 1 supernumerary bristle (SY. and fig. 2); Q generally with none . . ¢ é : 0 a c 2 2 Males : ; : : : ‘ : 6 : : : 0 3 — Females : d : : : g : : : : : a 0 18 3 Distal arm of sternum 9g (figs 20, 21, D.A.9) with a distinct subapical notch (NCH.) on anterior (dorsal) margin . : : : ; ; : ; , 0 4 — D.A.g9 lacking distinct subapical notch; instead, anterior margin flat between apex and subapical lobe (figs 67, 112, SUB.L.) or else arcuate (fig. 33, ARC.) 5 : 9 4 Ford’s sclerite of aedeagus with a thumb-like process (figs 42, 44, THM.) on alpha- portion (ALPH.) which extends well above base of upper arm (U.A.) of securifer. Upright portion of ALPH. much broader for ventral 3 than subapically — Ford’s sclerite lacking thumb-like process of alpha-portion (fig. 45, ALPH.); apex of ALPH. in line with base of upper arm (U.A.) of securifer. Upright portion of ALPH. rod-like . : : 6 6 . . . @ngustata sp. nov (p. 227) 5 Margins of apical lobe of D.A.9 (fig. 20, AP.L.) truncate. Thumb (figs 22, 42, THM.) about thrice as long (tall) as broad 4 2 : . adryadosa sp. nov. (p. 215) — Margin of apical lobe of sternum 9 (figs 53-55, AP.L.) angled or with upper half convex. Thumb (figs 56-58, THM.) only 1-2 times (or less) as tall as broad robinsoni subspp. 6 6 Upper 4 of margin of apical lobe (figs 21, 55, AP.L.) of D.A.9 distinctly convex. Notch (NCH.) with lower margin 1-5-2 times as long as upper margin : 0 7 * Only one of the alternates cited need apply; these are not combined characters. 8 The females of M. yhaeba (Jordan, 1926) and M. batibacula sp. nov. are unknown, E 266 10 II 12 ROBERT TRAUB Upper 4 of margin of apical lobe (figs 53-54, AP.L.) of D.A.g9 quite straight or only slightly convex. Notch (NCH.) with lower margin only slgihtly longer than upper margin : . r 3 : 5 c 8 Lower $4 of apical lobe (fe 20. AP. le ) concave. Lower arm of securifer (figs 23, 44, L.A.) with apex not relatively acuminate; its caudal margin quite straight to near apex. Tanned portion of quasi-crochet (Q.C.) about 4 times as long as broad at middle. (Malayan peninsula) : . robinsoni robinsoni (Roths., 1905) (p. 233) Lower 4 of apical lobe (fig. 55, AP.L.) almost straight. Lower arm of securifer (fig. 57, L.A.) with apex somewhat acuminate due to subventral curve of caudal margin. Tanned portion of quasi-crochet (Q.C.) 3-5 times as long as broad at middle. (Sarawak) . robinsoni peregrinata subsp. nov. (p. 234) Upper arm of securifer (Gg. 58, U.A. ) with apical snout relatively broad, about 2-5 times as long (from vertical internal root) as broad at middle; diameter at middle nearly as broad as subapical portion of lumen of sclerotized inner tube (S.I.T.). Margin of apical lobe (fig. 54, AP.L.) of D.A.9 quite flat and evenly angled at middle. (Java) . 2 3 4 : . robinsoni bogora subsp. nov. (p. 235) Upper arm of securifer (fig. 56, U.A.) with snout comparatively narrow, about 3-4 times as long as broad near middle; its diameter at middle much narrower than subapical lumen of S.I.T. Apical lobe (fig. 53, AP.L.) of D.A.g with lower } of margin slightly concave; the angle at middle blunt. (Tioman Island, Malaya) robinsoni tiomanica subsp. nov. (p. 236) Ford’s sclerite of aedeagus with a distinct groove (figs 71, 113, GRV.) between alpha- portion (ALPH.) and securifer; but lacking a thumb-like apex to ALPH. Anterior margin of D.A.9 flat immediately above base of subapical lobe (figs 67—68, 111-112, SUB) % 10 Ford's sclerite lacking a deeaer groove reaee Nae 97- 98); ena (THM. yell devel oped instead. Anterior margin of D.A.g arcuate (figs 33, 91, 134, ARC.) above base of subapical lobe (SUB.L.), the arc extending for at least 4 length of margin above SUB:L, = P A F : 5 7 : : a ; 16 Movable finger (fig. 138, F.) with breadth of subapical constricted area less than $ that of very broad base and equally broad apex. D.A.9 with a delimited nubbin at anterodorsal angle . : c : rhaeba (Jordan, 1926) (p. 257) Movable finger (figs 67, 109, 119, F.) oath constricted area not as narrowed; always much broader than 4 breadth of apex and generally so for base. Lacking a distinct nubbin at anterodorsal angle of D.A.9, although apex may be Geaikert ee 67, III, E27) II Caudal margin of D. A.g heavily sclerotized and apically appearing as a made like extension (figs 112, 120, EXT.9); opposite margin lightly tanned. Spiniforms of D.A.9 relatively long and narrow, well spaced. Subapical lobe (SUB.L.) of D.A.9 spur-like. Distal 4 of quasi-crochet not appreciably narrower than basal part (figs 114, 128, Q.C.) 0 12 Caudal margin of D.A.g9 curving subapically, (figs 67, 81) and hence not rod- like; not heavily sclerotized; anterior margin tanned to apex. Spiniforms of D.A.g short and at times close-set. Subapical lobe (SUB.L.) broadly triangular and apex blunt. Distal 4 of quasi-crochet much narrower than basal meh A appears like a tongue-like extension (figs 71, 82, Q.C.) 14 Movable finger (fig. 126, F.) short and broad, soneee or more ae Paice as long (above apex of immovable process, P.) as broad at narrowest level (immediately below distal fringe (D.FR.)). Relatively tanned portion of quasi-crochet (fig. 128, Q.C.) short and broad, ony 1-3 times as pee as broad because of markedly convex rea Margin . 5 loncha (Jordan, 1926) (p. 255) Movable finger (figs 109, 119, F) at least - 75 eves as long as broad at narrowest level. Tanned portion of quasi-crochet (figs 114, 115, Q.C.) at least twice as long as broad. j 3 ;: Pi 3 5 : : A z : F 13 13 14 15 16 07 18 NEW TAXA OF FLEAS 267 Movable finger (fig. 109, F.) relatively broad throughout, about 3-75 times as long as broad below distal fringe (D.FR.). Tanned portion of quasi-crochet (fig. 115, Q.C.) nearly straight, not angled upwards (but slightly curved) batibacula sp. nov. (p. 250) Movable finger (fig. 119, F.) much broader basally than at constricted area below D.FR.; about 5 times as long as broad at constriction. Quasi-crochet (fig. 114, Q.C.) angled at middle so that distal 4 is at an angle of about 40° with proximal veruta sp. nov. (p. 252) Movable finger (figs 65, 66, F.) with caudal margin somewhat excised immediately below distal fringe (D.FR.) so that apical portion of F. appears almost stalked. Bases of D.FR. forming a line that is almost perpendicular. Groove of Ford’s sclerite extending dorsad together, of equal length (fig. 71, GRV.) phangi sp. nov. & subsp. nov. 15 Movable finger (fig. 80, F.) with caudal margin evenly and slightly concave below distal fringe (D.FR.); apical portion of F. not differentiated. Bases of D.FR. forming a markedly oblique line. Groove of Ford’s sclerite (fig. 82, GRV.) incomplete dorsally, margin of alpha-portion (ALPH.) extending more distad (dorsad) than opposite member and then bea a characteristic, broad truncate flap . . limisp. nov. (p. 242) Movable fneee (fig. a F. ) peels narrow yaad areted so that distal fringe (D.FR.) is on a short stalk. Breadth of F. at constriction much shorter than distance between proximal-most bristle of D.FR. and apex of F. Tip of subapical lobe of D.A.9 (fig. 67, SUB.L.) much closer to dorsal margin of median microspiculose area (M.MSP.) than to apex of D.A.9. (Sarawak) phangi tana subsp. nov. (p. 242) Movable finger (fig. 66, F.) with distal fringe (D.FR.) not ona short stalk. Breadth of F. at constriction subequal to distance between basal bristle of D.FR. and apex of F. Tip of SUB.L. further removed from apex of median microspiculose area (M.MSP.), distance only slightly less than that between SUB.L. and apex of D.A.9 phangi phangi sp. nov. (p. 240) Movable finger long and evenly narrow (fig. 133, F.) about 5 times as long (distad of apex of immovable process, P.) as broad immediately below distal fringe (D.FR.); caudal margin straight near base so that F. is 4 times as long as broad at level of apex of conical process (C.P.). Quasi-crochet with sclerotized portion (fig. 95, Q.C.) relatively short and broad, about 2-6 times as long as broad at middle javana (Jordan, 1933) (p. 256) Movable finger (figs. 34, 92, F.) much broader in proportion and caudal margin markedly bulging near level of conical process (C.P.); at most, scarcely more than 4 times as long as broad below distal fringe (D.FR.) and 2-4 times as long as broad at level of apex of C.P. Tanned, dark portion of 2S long and narrow, at least thrice as long as broad at middle (figs 97, 98) . 17 D.A.9 (fig. 33) with a deep, symmetrical arcuate sinus (ARC. ) ereenaite om tip of subapical lobe (SUB.L.) to apex of apical lobe (AP.L.). Ford’s sclerite hae thumb (fig. 98, THM.) short in height, broader than high. Lower arm (L.A.) 0 securifer terminating in a short, angulate tip . 5 arcuata sp. nov. e 227) D.A.9 (fig. 91) with arc (ARC.) above subapical lobe short and shallow, almost indistinguishably merging with apical lobe (AP.L.) so that entire apical region truncate,not lobate, on anterior margin. Ford’s sclerite with thumb (fig.97, THM.) about twice as high as long. Lower arm (L.A.) of securifer with apex acuminate thurmani sp. nov. (p. 244) Mesal genitalic ridge of tergum 8 (fig. 139, M.R.8) consisting primarily of a conspi- cuous, short, horizontal, shallowly arched sclerotization; with anterior vertical portion represented only by a faint dorsal arm and ventral arm inapparent rhaeba (Jordan, 1926) (p. 257) 268 19 20 21 22 23 24 25 ROBERT TRAUB Mesal genitalic ridge of tergum 8 (figs 27, 32, 84, M.R.8) with a well developed vertical branch and this usually more sclerotized than horizontal fork; lacking the con- spicuous arched, tanned ridge : 19 Sternum 7 with subventral sinus not semis a sees, even Be base) (figs 26, a 124, 7 S.), its dorsal margin spine away from ventral at an angle ee more than 75° . 20 Sternum 7 with peneaicel sinus moet ie, at rane at Pe (figs. 32, ae 41,7 S.), its dorsal margin sloping at angle less than 60° and often paralleling ventral one for at least 4 length of latter . : 21 Sternum 7 (figs 26, 30, 7 S.) with dorsal portion of caudal maxgin (at level of spiracular fossa) sloping ventrad at an angle of about 60° from vertical. Caudal margin fairly evenly rounded (not produced into a lobe) at junction with upper margin of ventral sinus. Tergum 8 (fig. 31, 8 T.) with a ventrocaudal lobe dryadosa sp. nov. (p. 215) Sternum 7 (fig. 124, 7S.) with caudal margin nearly perpendicular at level of spiracu- lar fossa. Witha caudal, short submedian lobe. Tergum 8 (fig. 117, 8 T.) lacking a ventrocaudal lobe. ; : veruta sp. nov. (p. 252) Sternum 7 with dorsal margin of eee sinus Sante dorsocaudad from near base of sinus instead of paralleling ventral margin for 4 or more ee of latter (figs 40, 130, 7 S.) 22 Sternum 7 with a true seisnenteal aah (ies 41, ve I 18, Osh ), dorcel margin parallel ing ventral for 4 or more of its length . 23 Perula of bursa copulatrix (figs 40, 137, P.B.C.) oni ventral } dilated so that oes 4 twice as broad as dorsal 4. Paragenital morion (fig. 40, PG.M.) represented by a turned, narrow, U-shaped are near P.B.C. Glandula vaginalis (G.VG.) single. Ventrocaudal lobe of tergum 8 (8 T.) somewhat longer than broad at base javana (Jordan, 1933) (p. 256) Perula of bursa copulatrix (fig. 118, P.B.C.) elongate, scarcely broader subventrally thansubdorsally. Paragenitalmorionnear P.B.C.nottanned. Glandula vaginalis (G.VG.) preceded by an accessory “‘gland”’ (fissure) or a part of paragenital morion. Ventrocaudal lobe of 8 T. short, broad and apically blunt; its length less than breadth of base ; loncha (Jordan, 1926) (p. 255) Perula of bursa anprleee wereuiionn ‘(figs 78, 84, P.B.C.), not appreciably broader at middle than nearends. Paragenital morion (PG.M.) large, appearing as a broad, U- shaped loop of stout coils extending from ventral aes of P.B.C. to lower portion of sternum 9 (9 S.) é C a ° 0 6 24 Perula of bursa copulatrix oveid (figs 32, 4t, 102, P. B. C.), more dilated at middle than at ends. Paragenital morion (PG.M.) reduced, not represented by a broad loop of stout coils : : : : - : 5 - 26 Upright branch of mesal penital ridge of 8 T. (figs 79, 85, M.R.8) in oblique position, and caudal fork sloping ventrocaudad at angle of 45°. Caudal margin of P.B.C. flattened . . limisp. nov. (p. 242) Upright branch of mesal genitalic ridge of 8 ik (fig. 78, M. R. 8) subvertical and caudal fork nearly horizontal. Caudal margin of P.B.C. shallowly convex phangi sp. nov. & subsp. nov. 25 Sinus above ventral lobe of sternum 7 (fig. 75, 7 S.) definitely shorter and narrower than lobe below it. 7 S. scarcely concave above submedian lobe. Paragenital morion with a tanned chord along ventral margin of inner core, near anterior margin of loop, but lacking a transverse rod-like sclerotization in the area Phangi tana subsp. nov. (p. 242) Sinus above ventral lobe of 7 S. (fig. 76) subequal to lobe below it. 7 S. fairly deeply concave above submedian lobe. Paragenital morion (fig. 78, PG.M.) with a trans- verse rod-like sclerotization near anterior margin of loop, instead of a ventral one phangi phangi sp. nov. (p. 240) NEW TAXA OF FLEAS 269 26 Ventral, vertical fork of mesal genitalic ridge of tergum 8 (figs 32, 60, 62, M.R.8) with tanned portion as long as, or longer than, dorsal branch. Horizontal branch equally long. With a rod-like, tanned, narrow loop of paragenital morion (fig. 32, PG.M.) in line with and just caudad to base of perula of bursa copulatrix (P.B.C.) robinsoni subspp. 27 — Ventral, vertical fork of M.R. 8 (figs 41, 102) definitely shorter than dorsal branch, and horizontal branch equally short. Paragenital morion (PG.M.) lacking a rod- like, narrow loop near base of P.B.C._.. : : : . 5 30 27 Labial palpus extending beyond base of fore- trochanter. (Sarawak) robinsoni peregrinata subsp. nov. (p. 235) — Labial palpus extending at most to apex of procoxa . 28 28 Dorsal wall of vagina (figs 32, 59, VAG.) with 1-3 bead- like thickenings distad e glandula vaginalis (G.VG.) (Malayan peninsula) robinsoni robinsoni (Roths., 1905) (p. 233) — Dorsal wall of vagina (figs 62, 64, VAG.) lacking bead-like thickenings distad of G.VG. 29 29 Ventral lobe of 7 S. (fig. 62, 7 S.) definitely longer than sinus above it®; with only 2 bristles on the lobe ( but 1 or 2 subventrals immediately preceding it). (Java) robinsoni bogora subsp. nov. (p. 235) — Ventral lobe of 7 S. (fig. 64, 7 S.) subequal to lobe above it; with 3 bristles on lobe. (Tioman Island) . robinsoni tiomanica subsp. nov. (p. 236) 30 With an accessory “gland” or rearaad in dorsal wall of vagina, distad of glandula vaginalis (fig. 41, G.VG.). With a pair of parallel, short, tanned arcs or chords at dorsocaudal Bar of duct of bursa copulatrix (D.B.C.) and another above tip of G.VG. : c angustata sp. nov. (p. 227) — Lacking both an accession “gland” derd of piaedala varinalis (fig. 102, G.VG.) and the sets of tanned chords near D.B.C.andG.VG. . thurmani sp. nov. (p. 244) B. Lentistivalius gen. nov. Diacnosis. Relatively unmodified species with: (1) Labial palpus (excluding palpiger) (fig. 140) 5-segmented. (2) First preantennal row of bristles quite or actually near frons. (3) Metatibia (figs 149, J; 151, 2) with at most 2 stout dorso- marginal bristles in notches and with subdorsals unmodified and thin. (4) Movable finger (fig. 143, F.) long and narrow, with a well developed stiva (STV.). (5) Sternum 9 with distal arm (figs. 144, 145) clavate and with dilated subapical region armed with marginal spiniforms. (6) Ford’s sclerite of aedeagus (fig. 148, F.SC.) with alpha- portion (figs 152, 154, ALPH.) highly developed, apically pointed and often with medial portion approximating the letter ““M.” (7) Crochet with body (B.CR.) and process (CR.P.) together suggesting a broad letter ‘‘U”’ in appearance. (8) Crochet with only narrow ventral elements linking sclerite with aedeagal pouch and hence lacking a conspicuous quasi-crochet. (9) Phylax (PHY.) slender. (10) Sclerotized inner tube (S.I.T.) of moderate length and lacking specialized armature or spurs. (11) With hood (HD.) and its deltoid flap (DEL.FL.) covering most of lateral surfaces ofendchamber. (12) Female tergum 7 with lobe below antepygidial bristles (fig. 156, L.L.7) relatively short, not acuminate. (13) Spermatheca (SP. and fig. 158) at most with a very short dorsocaudal peak. (14) Mesal genital ridge of tergum 8 not ® Care should be taken that the ventral lobes are the same in appearance on both upper and lower sides of the specimen, for that indicates that the specimen was not distorted in mounting. If, on the other hand, one side is smaller than the other, the possible discrepancies should be considered in interpreting this couplet. 270 ROBERT TRAUB developed. (15) Tanned portions of paragenital morion somewhat reduced (fig. 160, PG.M.) rather indistinct. GENERIC DEscriIPTION.!® Anterior margin of head (figs 140, g; 141, 9) only slightly curved below level of median sensory crater; head relatively short and high, about thrice (or somewhat more) as high (from vertex at falx to ventral margin) as long at level of uppermost bristle of row II in J; about 2:5-3-0 times in 2. Ventral margin 0: head below eye biconvex, the 2 lobes overlapping slightly at this level. Generally with 4 rows of tairly narrow bristles and with a few “supernumerary” bristles. Labial palpus with fifth (ultimate) segment not reaching beyond procoxae. Dorsal margin of pronotum short, generally about 4 length of dorsal spines of comb (subequal to spines in aestivalis Jameson & Sakaguti, 1954). Mesonotum (fig. 142, MSN.) with 1 or 2 pseudosetae per side, usually subdorsal. First segment of metatarsus about I-45 times length of second and nearly thrice length of third (which is subequal to fifth); about 2-4 times length of first segment of protarsus. Fourth vinculum (fig. 159, VC.4) usually projecting forward as a short rod. Terga 2-4 or 5 with a short subdorsal spiniform. Unmodified abdominal terga with 2 rows of bristles, preceded by a group of a few small subdorsals. Spiracular fossae of these segments broadly sagittate. Basal abdominal sternum with a few proximal sub- median bristles in g, in 2 with additional bristles reaching near middle of segment (fig. 159). Female with modified bristles of tergum 7 (fig. 156, U.M.B., L.M.B.) not closely resembling antepygidials (A.B.); only 1 such lower modified bristle and this not strongly displaced. Tergum 7 lacking an appreciable dorsal lobe in ¢ (fig. 147, 7 T.) ; in 9, lower lobe (L.L.7) quite short, and upper one (U.L.7) particularly so. Male. Sternum 8 (fig. 147, 8 S.) very large, nearly 1-5 times as long (near ventral margin) as high; unmodified, and lacking close-set or spiniform ventral bristles. Manubrium (figs 143, 146, MB.) broad; with ventral margin biconvex near middle. Bay of manubrium (B.MB.) extending to near apex. Immovable process of clasper (P.) with apex short and broad, subrounded or blunt. Conical process (C.P.) of clasper fairly broad, 3-5 times as tall as broad at middle. Movable finger (F.) long and thin; about 6~7 times as long (from level of apical margin of P.) as broad at middle of distal fringe (D.FR.). Stiva (STV.) long and narrow. D.FR. with about 4-5 well separated, long and relatively thin bristles; wholly or mainly on upright portion of shaft of F., proximad to curve of the stiva. Fulcral sclerite (F.S.) higher (longer) than broad; subvertical; broadest at dorsal end. Proximal arm of sternum g (P.A.g) in subvertical position ; gradually broadening from base to near apex, where it may dilate considerably. Distal arm of sternum 9 (D.A.g) nearly length of P.A.g; widening at apical fourth to sixth, at least caudally (ventrally); with that margin bearing a subapical group of spiniforms or short, stout bristles of decreasing lengths (towards apex). Clava of D.A.g with a lateral supramedial flap (SUP.FL.) bearing small marginal bristles, recalling Medwayella (but D.A.g lacking other specialized features of Medwayella). Aedeagal apodeme (fig. 148, AE.A.) resembling manubrium for most its length 10 The genus is compared with Medwayella gen.nov., q.v., and the differences are stressed; similarities are generally omitted. NEW TAXA OF FLEAS 271 though perhaps slightly narrower; lacking an apical appendage. Lateral laminae (L.LAM.) well sclerotized only anteriorly; ventral and caudal portions often very lightly tanned, indistinct. Aedeagal pouch well tanned ventrally (fig. 152, AE.P.- V.), but less so laterally (AE.P.-L.) ; unsclerotized at base and girdle hence not clearly demarcated; lacking obvious sclerotized connections with crochet process (CR.P.) except for a tail-like ventral process at the base of CR.P. and narrow ventral elements (cf. fig. 153, ventral aspect) and therefore lacking a quasi-crochet (cf. figs 22, 23, Q.C.). Hood (HD.) without notch or sinus, although at times (fig. 155) concave ventrally; covering most of endchamber; its deltoid flap (DEL.FL.) not produced dorsally into an apical acuminate process. Anterior margin of DEL.FL. well caudad of fulcrum (AE.F.); caudodorsal origin near cephalodorsal angle of Ford’s sclerite (F.SC.) rather than upper margin of body of crochet (B.CR.). Lateral lobes (L.L.) greatly reduced; inapparent. Sclerotized inner tube (S.I.T.) sub-horizontal; about 4-6 times as long as broad at middle; lacking dorsal or ventral spurs or other special- ized armature. Ford’s sclerite (F.SC.) very large and constituting dominant sclerite in endchamber, and with alpha (ALPH.) and securifer (SEC.) components fused and distinctions apparent only at margins (cf. fig. 153, ventral aspect). ALPH. a major component of F.SC., and its dorsal, ventral and apical margins with characteristic shapes in various species. Securifer (SEC.) comparatively reduced (in lateral aspect) ; often produced into an apical spur. Phylax (PHY.) fairly long and narrow, some- what crescentic in shape, extending from near base of CR.P. to near ventral margin of F.SC. and apex of pivotal ridge (PIV.R.), which is lightly tanned. Pivotal chord (PIV.CD.) weakly sclerotized. Crochet boomerang-shaped or broadly U-shaped; body of crochet (B.CR.) subvertical and crochet process (CR.P.) sub-horizontal. Aedeagal fulcrum (AE.F.) quite narrow. Crescent sclerite (C.S.) moderately long; satellite sclerite (SAT.S.) relatively so; latter overlying base of S.I.T., i.e., appearing as if “within” it. Vesicle (V.) lightly tanned. Third apodemal rod (AP.R.) of endophallus not particularly well developed. Caverna spiculosa (CAV.SPIC.) long, commencing subdorsally but at about level of base of fulcrum. Female. Spermatheca (figs 156, SP. and 158, 161) longer than broad; bulga (B.) with internal striae; lacking a truly distinct peak; but at times with a short dorso- caudal bulge. Hilla (H.) with an apical papilla (PAP.); base inserted fairly deeply into bulga, protruding element therefore appearing relatively broad in relation to length; internal portion with striae. Duct of spermatheca (D.SP.) bearing internal sclerotized rings at least near dilated portion (DIL.P.). Dorsal region of perula of bursa copulatrix (P.B.C.) associated with 2 sclerotized rod-like or dot-like bodies. Duct of bursa copulatrix (D.B.C.) lightly tanned. Paragenital morion (fig. 160, PG.M.) somewhat reduced ; inconspicuous and rather ventral in position, near apical region of bursa copulatrix. Sternum 7 (figs 156, 160, 7 S.) with a subventral sinus. Ventral anal lobe (V.A.L. and fig. 157) with base short; apical margin fairly long, somewhat sinuate; with a gap between submesal and subapical groups of bristles. Anal stylet (A.S.) fairly long and narrow and lacking long subapical bristles. Tergum 8 (8 T.) lacking a conspicuous lobe at ventrocaudal corner. Mesal genitalic ridge of tergum 8 undeveloped. Spiracular fossa 8 (8 SPC.) medium-sized; upright portion ovate. Sternum 8 (8 S.) flask-shaped and with short apical bristles. 272 ROBERT TRAUB The type of the genus is L. vomerus sp.nov., described below. Here also belong the “Stivalius ferinus-group” of Smit (1958), viz., ferinus (Rothschild, 1908), a parasite of shrews on the Indian subcontinent, Ceylon and South China; imsolla (Traub, 1950), a flea of birds in the mountains of Malaya; aestivalis (Jameson & Sakaguti, 1954) infesting Apodemus speciosus, a wood-mouse in Japan; and alienus (Smit, 1958) stated by the author to be a parasite of rodents, and occurring in East and Central Africa. ComMENT. The generic name is derived from the Latin term Jentes, meaning flexible, and refers to the remarkable facility of this genus to adapt to a broad variety of hosts, in widely separated areas, as indicated above. This point is discussed further below in the second article in the series, dealing with convergent evolution, and in the third, which concerns zoogeography. 1. Lentistivalius vomerus sp. nov. TYPE MATERIAL. Holotype 4, allotype 2 (B-19249) ex Tupaia montana; NORTH BORNEO (EAST MALAYSIA, Sabah): Mt. Kinabalu, Tenompak; elev. 4500 ft; 16.VIII.1953; Coll. R. Traub for U.S. Army Medical Research Unit (Malaya). Paratypes, all from North Borneo (East Malaysia, Sabah) as follows: oo 8 Host Locality, Date and Collector 99 75 Tupaia montana Mt. Kinabalu, vicinity of Tenompak; elev. 4500-5500 ft; July 1, 1951 R. Traub August, 1953 R. Traub May, 1952 J. R. Audy Feb.—Mar. 1964 Lord Medway & Cambridge Univ. Exped. Dec. 1965 Lim Boo Liat & D. Heyneman for G. W. Hooper Foundation 3 I oe a Mt. Kinabalu, vicinity of Mari Parei; elev. 4000-5100 ft; 24—25.VIII.53 Rob I - An 7" Mt. Kinabalu, Lumu Lumu; elev. 6300 ft 23.VII.1951 eles - 3 7 of Mt. Kinabalu, Kamborangah; elev. 7200 ft 3.VI.1952 J.R.A. 6 4 % Mt. Trus Madi, Pampang, Ulu Kaingaran; elev. 4000 ft 19.VII-7.VIII.53 J.R.A. = * of Mt. Trus Madi; elev. 7500 ft 30.VIII.1956 Cambridge U. Exped. 5 3 Dendrogale melanuva Mt. Kinabalu, Lumu Lumu; elev. 6300 ft 21.VII.1951 Rea = I si _ Mt. Kinabalu, Mari Parei; elev. 5100 ft 25.VIII.1953 Ra. —) OK 7 rr Mt. Kinabalu, Tenompak; elev. 4500 ft 30. VIII.53 RAL: NEW TAXA OF FLEAS 273 Host Locality, Date and Collector (continued) PF Mt. Trus Madi, Pampang, Ulu Kaingaran; elev. 4000 ft 1.VIII.1953 JEReAS a Fy Mt. Trus Madi, Kidokarok; elev. 5000 ft 3.1X%.1956 Cambridge U. Exped. Dremomys everetti Mt. Kinabalu, Tenompak; elev. 4500 ft 16.VII.51, 31. VIII.53 Re. =A " Mt. Kinabalu, Tenompak; elev. 9800 ft Dec. 1965 L.B.L. , i Mt. Kinabalu, Lumu Lumu; elev. 6300 ft ead Os . ee Mt. Trus Madi Aug.—Sept. 56 Cambridge U. Exped. Mt. Trus Madi, Pampang, Ulu Kain- garan; elev. 4000 ft July-Aug. 53 J.R.A. Hylomys suillus Mt. Kinabalu, Tenompak; elev. 4500— 5000 ft 16.VII.59, 12-20. VIII.53 Tee 10 os ae Mt. Kinabalu, Mari Parei; elev. 5100 ft 25-VIII1.53 Re Rattus alticola Mt. Kinabalu, Tenompak; elev. 5500 ft 16.VII.1951 R.T. a 7 Mt. Kinabalu, Mesilau Base Camp 10.11.1964 L.M. Rattus whitehead Mt. Kinabalu, Tenompak; elev. 5500 ft 19. VII.1951 1 ceed De Rattus sp. Mt. Kinabalu, Bundu Tuhan; elev. 4000 ft 26.V.1952 J-R.A. - a Mt. Kinabalu, Kamborangah; elev. 7200 ft 5.VI.1952 J-R.A. Haematortyx Mt. Trus Madi, Pampang, Ulu Kain- sanguiniceps garan; elev. 4000 ft 22.VII.1953 J.R.A. I Shrike Mt. Kinabalu, Lumu Lumu, elev. 6300 ft. 23.VIII.1951 R.T. Holotype (U.S.N.M. number 71609), allotype and five pairs of paratypes deposited in U.S. National Museum. Remaining paratypes distributed as for M. dryadosa. Diacnosis. Close to L. insolli (Traub, 1950) (new combination) and even agrees with it in that the dorsal and submedian tanned margins of the alpha-portion of Ford’s sclerite (figs 152, 154, ALPH.) are shaped somewhat like the letter “M.” Instantly separable in that the new species lacks modifications of the pronotal comb of L. insolli which have been cited (Traub, 1969) as being characteristic of bird-fleas, viz.: (t) A large number (about 28 in L. insolli) of narrow, quite straight, mainly horizontal, parallel spines in a comb which does not descend over third vinculum. (2) Axis of the bases of the majority of spines in comb slopes anteroventrad. Instead, in the new species, there are only about 20 spines in the pronotal comb (fig 140) and the spines are broader, i.e., no significant gaps between them and yet height of comb is the same—extending to a level just above VC.3; middle 6 spines fairly concave, and axis of the bases of the main spines largely subvertical, not oblique. It has been pointed out (Traub, 1966) that in bird-fleas (and presumably other ectoparasitic 274 ROBERT TRAUB insects parasitizing birds) the bristles of the body are longer and thinner than in relatives infesting mammals, and this is also true for L. insolli versus L. vomerus,e.g., caudoventral bristle of mesepimere 31 times as long as broad at base in the former species and only 23 times in the latter (fig. 142, MPM.). Further separable from L. insolli as follows: (1) movable finger (fig. 143, F.) proportionately much broader in new species, i.e., only 4-4 times as long (from apex of P.) as broad immediately below stiva (STV.), instead of more than 7-6 times (fig. 146). (2) Distal arm of g sternum 9g (D.A.9 and fig. 144) with apical margin ovate, not oblate (fig. 145); margin above subapical spur curved and slanting, not straight. (3) Spermatheca relatively longer and narrower, i.e., about 2-2 times as long as broad at maximum diameters (figs 158, 160) when undistorted, instead of 1-7 times. Near L. aestivalis but instantly separable as follows: (1) Deltoid flap (fig 152, DEL.FL.) virtually squarely meeting ventral margin of hood (HD.) at anteroventral corner instead of being produced into a long, acuminate, curved structure (fig. 155). (2) Crochet process (CR.P.) narrowly ovate, nearly 5 times as long (to middle of base B.CR.) as broad at middle, instead of being trapezoidal and broader, viz., 2:5 times as long as broad. (3) Lobe above subventral sinus of 9 sternum 7 (fig. 156, 7 S.) at most with a shallow sinus instead of one that virtually equals that below it, so that this region is practically evenly biconcave. Differences from other species of Lenti- stivalius are indicated in the key which follows the description below. DESCRIPTIVE NOTES.!! Head (figs 140, J; 141, 2). With preantennal portion of head of 3 about 2-4 times as high as long at level of eye-bristle and slightly more than thrice as high as long at level of uppermost bristle of row II; in 9 comparable figures are 2:3 and 3. With preantennal bristles in 4 rows except for a fair-sized super- numerary between rows II and III. Labial palpus with apical segment 1-7 times length of penultimate (fourth) ; extending to about apex of procoxa. Thorax. Pronotum (fig. 140) narrow; dorsally with spines of comb about 2:5 times length of notum; with 2 rows of bristles, but first row highly abbreviated. Pronotal comb with a total of about 20 spines, of which, in g, numbers 2 — 8 per side (counting dorsalmost as number 1) are slightly convex, narrowing gradually from base to apex; bluntly pointed or subovate at apex; bases of middle spines inclined ventrocaudad at an angle of about 20°; spine number 7 the broadest, but only slightly exceeding its mates, about 5 times as long as broad at middle. Mesonotum (fig. 142, MSN.) with 4 rows of bristles; first row abbreviated ; those of first 2 rows short; with 1 subdorsal pseudoseta per side. Mesepisternum (MPS.) with oblique row of 4 bristles; dorsalmost (first) of these near middle of caudal margin; slope anteroventrad; first and third bristles long; lowest subventral in position. Mesepimere (MPM.) with 2 rows of 3 long bristles; that near ventrocaudal angle longest. Third vinculum (VC.3) with dorsal margin basally quite flat, distal } concave; axis about 35°-45° from horizontal. Lateral metanotal area (L.M.) with external measurements slightly longer dorsally than high, but internal, relatively untanned region somewhat higher than long. Metepimere (MTM.) with 3 rows of 3-4 bristles each and those of last 11 The species is compared with M. dryadosa, q.v., and the differences are stressed; similarities are generally omitted. NEW TAXA OF FLEAS 275 row with intercalaries (usually 2 such between ventral pair of long ones); caudal margin becoming shallowly convex near level of spiracular fossa. Legs. Metacoxa with a group of about 14-16 short, thin, mesal bristles on apical i, anterior to internal rod. With 5-8 submedian lateral bristles in 2 irregular longi- tudinal rows of profemur. Meso- and metafemora with 4-5 small subdorsal bristles on apical 4; 3 ventromarginals on distal } and 2 submedians near spiniform member of apical pair of stout bristles. Protibia in $ with first (apical) and fifth members of dorsomarginal pair of bristles much shorter and somewhat longer than their mates, hence appearing as “‘single” bristles, in 9, with 3 such single bristles between third and fifth pairs, and since first pair is similarly modified, this sex here bears an incipi- ent comb of false spines. Other legs with dorsomarginals paired in usual manner (metatibia, figs 149, ; 151, 9), and only D.M.4 single; $ with 3 non-marginal rows of lateral bristles and only 1-2 supernumerary bristles; 2 with 4 such rows. An apical bristle of mesotarsus I-III and of metatarsus II and III, reaching to middle of suc- ceeding segment. Proportions of tarsi essentially as that cited for L. imsolli (Traub, 1950). Metatarsus with apical segment with 6 pairs of lateral plantar bristles, but third displaced towards midline (fig. 150). Abdomen. Basal sternum of 2 with group of about 7-10 small, thin, lateral bristles commencing near vinculum and extending to about anterior } and ventral }; in g with at least the bases for about 6 such hair-like bristles, but group nearer to antero- dorsal region. Terga 2-5 usually with 1 subdorsal apical spinelet. Spiracular fossae in unmodified terga of § symmetrical and narrowly sagittate (e.g., fig. 147, 7 SPC.); in 9 shorter and broader and dorsal margin more convex than ventral (fig. 156, 7 SPC.). Representative sterna of § with ventral group of 3 long bristles preceded by a group of 3 smaller ones and r more anterior ventromarginal one (e.g., 7S.). In @ (fig. 156, 6 S.) such sterna with 4 bristles in caudal row, but, except for 6 S., uppermost usually small; middle subventral group of 4-6 small ones; anterior- most, 3 or 4. _ Uppermost antepygidial bristle (A.B.) in each sex somewhat less than half of lower one. Female with only 2 adjacent bristles modified so as to suggest A.B., and only somewhat so, viz., dorsomarginal one (U.M.B.) rather dark and stout, but not displaced; lower one (L.M.B.) fairly close to A.B. but not as long, dark and stout as upper A.B. Dorsal lobe of tergum 7 (U.L.7) not produced caudad above and between bases of A.B.; margin below A.B. (L.L.7) scarcely extending beyond A.B., viz., not truly lobate. Modified Abdominal Segments—Male. Tergum 8 (fig. 147, 8 T.) slightly more than twice as high as long at middle; somewhat dilated ventrally; not extending below level of base of ventral anal lobe (V.A.L.). Vertical arm of spiracular fossa 8 (8 SPC.) rather more than thrice as high as broad at middle; dilated horizontal section only 1-25 times longer than high or than vertical arm is broad at middle. Sternum 8 (8 S.) nearly twice as long (along ventral border) as high at middle (level of first dorsal bristle); ventral margin sloping towards middle much more than upper, but quite straight for most its length; cephalic margin shallowly concave; caudal margin quite straight. 8 S. with bristles approximately as follows: a dorsomarginal row of 4, commencing just beyond midpoint, where margin starts to curve caudoventrad, and terminating at caudal } ,the last bristle by far the longest ; with 5 subdorsal bristles, 276 ROBERT TRAUB arranged 3-2, the first ventral row near midline, the second row oblique and in line with first of dorsomarginals; a horizontal row of 3 along midline, starting at anterior 4 and terminating near margin; another horizontal row of 3 just below this, but bases somewhat more cephalad and otherwise symmetrical; a group of 2 subventrals, sub- median, in a horizontal row; with 8-10 ventromarginals, or contiguous short bristles, arranged in twos and threes, starting near midline and terminating subcaudally; none of the above bristles anterior to cephalic }, and most caudal of midline. Immo- vable process of clasper (fig. 143, P.) dorsally very broadly and shallowly rounded ; with a short dorsomarginal bristle near middle of dorsal margin and another, longer, subapical one near apex of P.; caudal margin very shallowly sinuate distad of marked sinus at base of manubrium (MB.). Conical process (C.P.) about 3 -5—4 times as long a broad at middle; caudal margin straight. Movable finger (F.) with portion distad of apex of P. about 5-4 times as broad at level just proximad of curve of stiva (STV.) (cf. fig. I, }-k) and 3-7 times as broad at level of apex of C.P.; the difference represent- ing the degree of gradual narrowing of F. from base towards apex. F. with anterior and posterior margins otherwise appearing parallel; anterior margin somewhat convex but becoming slightly concave between proximal and third sensilla of group of 4 (S.G.). Stiva (STV.) fairly long and quite narrow; length (cf. fig. 1, measured from k-o to p) about 1-3 times subapical width of F.(j-k) and about ;% of total length of apex of F. (l-p); about thrice breadth of middle of stiva. Distal fringe (D.FR.) of 4 widely spaced, relatively thin, long bristles, commencing at level of middle of F. (above P.) and terminating at level of middle of STV. Fulcral sclerite (F.S.) essen- tially a right triangle with base and hypotenuse slightly sinuate and with altitude concave and apex blunt; length about 1-7 times breadth at dorsal margin, thrice that at middle and about 7 times that at apex. Manubrium (MB.) fairly broad; length (as measured from base of dorsal margin, where base of F. angles ventrad) about 1-9 times breadth of MB. at anterior ventral bulge; dorsal margin slightly convex to near apex, where it becomes sinuate and fairly straight subapically ; ventral margin parallel to dorsal near apex, somewhat sinuate to anterior bulge and here bi- convex but sinus between bulges short; caudal bulge longer than anterior; apex slightly upturned; anterior margin, at tip, relatively straight. Tergal apodeme of ninth segment (T.AP.9) relatively narrow; length (to base of F.) nearly 6 times breadth at middle. Proximal arm of sternum 9 (P.A.g) about 6-5 times as long (¢f. fig. 1, from aa—bb to level of ii) as broad at base (gg—hh); length thrice breadth at bulge of anterior margin at distal } (dd) ; 2-2 times breadth at apex (aa—bb). P.A.g with dorsal (posterior) margin slightly biconcave; anterior margin fairly concave at distal 4 (aa—dd) and thereafter quite straight ; dorsal margin shallowly sinuate except at corners. Distal arm of sternum 9 (D.A.9 and fig. 144) nearly 6 times as long (ii-qq) as broad at middle; margins sinuate but parallel to above apical 4, where caudal margin becomes oblately convex, so that distal portion resembles the head of a mace with posterior and apical margins oblate. Armature of mace-head consisting of marginal spiniforms, as follows: a stout sub-proximal one, above which are 4-5 smaller, paler spiniforms, commencing at middle of distal margin, and below which is 1 fairly long spiniform with a longish subspiniform at base of dilated portion. D.A.g with a subapical nubbin or short, stout, spur on anterior margin, above a short notch NEW TAXA OF FLEAS 277 with well tanned margins. Dilated or club-like apex of D.A.g about 1-4 times as long (from basal subspiniform) as broad at level immediately below subapical notch. Aedeagus—Lateral Aspect (figs 148, 152,172). Aedeagal apodeme (fig. 148, AE.A.) with middle lamina (M.LAM.) more tanned than lateral plates (L.LAM.); latter only well sclerotized near anterior end, and ventral and caudal portions semimembranous, indiscrete, with ventral region appearing confused with penis rods. M.LAM. about 4-4 times as long (from anterior edge of crescent sclerite, (C.S.), at base of aedeagal fulcrum, (AE.F.)) as broad at anterior margin of bay of middle lamina (B.M.L.). AE.A. significantly narrowing only at level of anterior edge of B.M.L.; rather resembling MB. in shape but not as broad; apex somewhat upturned. B.M.L. extending to apical }. Median dorsal lobe (M.D.L.) sloping dorsocaudad at angle of about 45° commencing at level of AE.F.; straight to even, broad curve at level of Ford’s sclerite (F.SC.). Aedeagal pouch (AE.P.) with base very lightly tanned and hence not representing a typical girdle, but with usual elements proceeding dorsad to top of apodeme, near anterior end, and en route blending with extensions of L.LAM. which project ventrad towards fulcrum and which join indistinct ventral margins of L.LAM. from anterior end of apodeme. Ventral wall of pouch (fig. 152, AE.P.-V.) well tanned. Hood (HD.) covering endchamber as a symmetrical cowl whose ventral margin is fairly straight and extends from tip of apices of Ford’s sclerite (F.SC.) to apex of crochet process (CR.P.); extended dorsad as the deltoid flap (DEL.FL.) and terminating acutely in subdorsal region at level of basal 4 of sclerotized inner tube (S.I.T.); dorsal and ventral margins in the main at right angles to cephalic margin. Ford’s sclerite (F.SC.) largest (in mass) of sclerites in endchamber; shaped like a broad, low arch with longitudinal axis horizontal. Most conspicuous part of alpha-portion (ALPH.) an M-shaped ridge or thickening whose one leg is long, sub- dorsal and subhorizontal; opposite leg only about half as long and with cross-bar in the form of a shallow ‘‘V”’, and apical. Distad of the ‘“M,” with a bell-shaped dome which is longer than high and terminates apically in an eccentric (ventral) short stalk, contiguous with a caudal acuminate process of the securifer (SEC.) so that tip of F.SC. is bifid. Securifer semi-circular, its lower (anterior) leg a short, fairly blunt lobe. Pivotal ridge (PIV.R.) lightly sclerotized and its chord (PIV.CD.) short, narrow and straight. Sclerotized inner tube (S.I.T.) nearly 7 times as long (from caudal apex of Y-sclerite (Y.S.) to middle of apex) as broad at level of caudal margin of crochet process (CR.P.) and 3-3 times as broad at sub-basal thickening (i.e., at short ventral bulge) ; axis very slightly arched to short up-curve at apex, where dorsal margin bears a short peak. Crescent sclerite (C.S.) fairly long and narrow; borders at times in- discrete because of unusual degree of tanning of lateral shafts of capsule (L.S.C.) and proximity of relatively large satellite sclerite (SAT.S.). Caudal extremity of C.S., and all of SAT.S., overlying base of S.I.T. Central sclerite (CEN.S.) closely appressed to fulcral medial lobe (FUL.M.L.) ; the two together appearing as the upper part of an arrowhead. Y-sclerite (Y.S.) conspicuous as a large, sub-triangular sclerite as broad as base of fulcrum and ventral to it, and with a narrow, rod-like fork extending dorsocaudad to above mid-line of S.I.T. at level of apex of C.S. Dorsal virga (D.V.) indistinct. Phylax (PHY.) a crescent whose chord is about 8 times its breadth at middle; lying across middle of S.I.T. so that upper 4 is above S.I.T., with apex near 278 ROBERT TRAUB ventral tip (base) of F.SC.; lower } of PHY. below S.I.T. and contiguous with crochet. Body of crochet (B.CR.) terminating at base of F.SC.; clavate, broadest subapically; ventral portion about 4} as broad as upper, as sclerite curves caudad. Crochet process (CR.P.) digitoid, directed caudad and proceeding slightly beyond level of apex of B.CR. With a short spur pointing cephalad and arising from anteroventral angle of crochet and merging with ventral walls of aedeagal pouch. Upper member of penis rods (P.R.) extending to near apex of apodeme; lower members shorter; both paralleling ventral margin of L.LLAM. Ventral virga (V.V.) stout; sclerotized to level of about base of AE.P. Third apodemal rod (AP.R.) well tanned to base of AE.P., but apparent for most its length, which equals P.R. Aedeagus—V entral Aspect (fig. 153). Hood (HD.) bifid apically. Ford’s sclerite with distal } consisting of broad, sub-quadrate, right and left components fused near base, at midline, but well separated distad; alpha (ALPH.) and securifer (SEC.) elements forming blended and integral parts of each 4 (and hence here constituting merely topographic terms useful in taxonomic distinctions). Basal 4 of F.SC. semicircular. Deltoid flap (DEL.FL.) covering lateral 3 of ventral surface. Phylax (PHY.) with ventral portion of each straddling sclerotized inner tube (S.I.T.) and sloping towards midline dorsally so that apices are virtually contiguous, above S.I.T. and here connected by a membrane; dorsal region stouter (from lateral surface towards mesal) than ventral. Wall of aedeagal pouch with ventral extensions (P.W.EX.) leading to floor of base of CR.P. Lateral lobes so ventral in position they appear like mesal structures. S.I.T. dilated subapically. Satellite sclerite (SAT.S.) along midline; well distad of base of S.I.T. and mesad of apices of crescent sclerites (C.S.). Lateral shafts of capsule (L.S.C.) relatively broad as well as long. Vesicle (V.) broad. Y-sclerite with ventral components broadly triangular with winged angles at base, serving as floor of capsule; more dorsal fork with symmetrical, broad, ovoid projections extending more mesad. Fulcral lateroventral lobes (FUL.L.L.) short, fairly widely separated. Caverna spiculosa (CAV.SPIC.) very long, somewhat fusiform, and occupying much of breadth of apodeme. Female. Sternum 7 (fig. 156, 7 S.) about 1-5 times as high as long at maximum diameter (i.e., level of maximum curvature of median lobe on caudal margin); anterior margin very shallowly convex; dorsal margin oblate; caudal margin with a subventral short sinus which is scarcely more than 4 of height of lobe beneath it. 7 S. with lobe above this sinus somewhat concave (at times more so than in figure) but soon straightening and sloping antero-dorsad from level of apex of ventral anal lobe (V.A.L.); margin rather concave from level of dorsal part of sensilium (SN.) to upper part of spiracular fossa 8 (8 SPC.). 7 S. with vertical caudal row of 4 long bristles, of which 1 is ventral and 1 subventral, about at level of middle of ventral lobe; 1 slightly above level of sinus and uppermost above third by distance equal to mouth of sinus; this row removed from caudal margin by distance equivalent to 4 length of lowest bristle. This row preceded by a vertical group of about 10 smaller bristles, most of which are in a line but some subventrals displaced caudad, as is a fairly long ventromarginal. Latter group preceded by about to bristles in 2 short, irregular, subventral rows of 3-4 and 4-5, plus I or 2 marginals or submedians out of line. Tergum 8 (8 T.) with caudal margin straight and oblique to short lobe at NEW TAXA OF FLEAS 279 ventrocaudal angle; with 2 rows of small bristles, usually 2-5, immediately before vertical portion of 8 SPC.; with a subventral, horizontal row of 3 fairly long bristles, preceded by 1 small one and with the caudalmost inserted before apex of sternum 8 (8 S.) ; with an oblique subvertical row of 3 bristles at level of base of V.A.L., of which uppermost is at level of upper member of ventral genital pair at caudal margin and lowest in line with ventrocaudal lobe; with 2 vertical rows of more anterior bristles; the first (anteriormost) of 3 small ones in line with ventral base of 8 S. and lowest of which was cited above as preceding subventral horizontal row; the second, of 3 small submedians, of which ventralmost is in line with first member of subventral horizontal row; third row of 4 submedians in line with second large member of subventral horizontal row. 8 SPC. with vertical portion broadly ovate, about twice as high (including section overlapped by horizontal arm) as long at middle; horizontal portion with tanned area about 2-2 times as long as broad at base. Dorsal anal lobe (D.A.L.) with a dorsomarginal row of 3 or 4 bristles; and 1-2 tiny ones contiguous with base of sensilium (SN.) ; caudalmost dorsomarginal very long; with a subdorsal row of 3, of which caudalmost is long, in line with last dorsomarginal and above base of anal stylet (A.S.) ; with a pair of subventrals immediately preceding long bristle ventrad to base of A.S.; with 2-3 small, scattered submedians. A.S. about 4 times as long as broad at middle; with a very small subapical bristle dorsal and ventral to base of apical long bristle. Ventral anal lobe (V.A.L.) angulate, with anterior margin less than 3 length of ventral margin; with 3 groups of ventromarginal bristles—a subequal but longish pair at anteroventral angle (at times represented by a single bristle); a long one at anterior fourth (at times paired here, e.g., when first is single), and a long pair on a short mesal flap at caudal third or fourth. Spermatheca with outlines variable, depending upon position in slide-preparation. Typically (fig. 156, SP. and fig. 158), as when seen in lateral aspect, with bulga elnogate-oval, approximately twice (or 2-2 times) as long (maximum diameter) as broad in middle. Atypical appearance shown in figs 160 and 161, and latter particularly misleading, for bulga seems only 1-7 times as long as broad. Bulga typically with ventral margin fairly flat except for anterior up-curve at level of apex of lower margin of internal portion of hilla (H.); dorsal margin anteriorly quite straight but with a subtruncate, short posterior bulge; cawdal margin convex. Hilla (H.) inserted into lumen of bulga for about 4 its length; external portion recurved at middle and apex rather flat, oblique below the apical papilla (PAP.); internal portion longer than breadth (inner measurements) at middle; of fairly uniform breadth to near apex. Bursa copulatrix with a pair of characteristic tanned, short rods or ill-defined thickenings flanking apex of perula (P.B.C.), one anterior and one posterior, at base of sac of bursa (SAC.). P.B.C. narrow, unmodified, scarcely differentiated from duct (D.B.C.) below it. Dilated portion of bursa (DIL.P.) weakly tanned and subequal in length to dorsal part of SAC. Duct of spermatheca with internal striae usually visible near entrance into SP. and near DIL.P. Paragenital morion (PG.M.) reduced to an indefinite semimembranous area near upper wall of vagina (VAG.) at genital orifice and overlying ventral portion of bursa; with elements lying anterior to latter. Glan- dula vaginalis (G.VG.) short, broad and indistinct, partially underlying expanded base of apical wall of bursa. Duplicatura vaginalis (D.VG.) indiscrete, at base of 280 ROBERT TRAUB vertical portion of SAC., which broadens at level of vagina, anterior to orifice of bursa copulatrix. HosT-RELATIONSHIPS AND DISTRIBUTION AND OTHER COMMENTS. The name vomerus is derived from the Latin word for plough-share and was selected not only because the shape of the movable finger resembles a plough-handle, but because Ford’s sclerite, as seen in dorsal or ventral aspect, suggests the share of the plough, with the elongate crochet-processes serving as handles, while, in lateral aspect, the crochet-processes themselves suggest the share as seen from the side. Lentistivalius vomerus sp. nov. is obviously common in the mountains of North Borneo, especially at elevations of about 4000-6300 ft, where it parasitizes primarily Tupaia montana but apparently also infests other small mammals in the same habitat, e.g., Dendrogale (a tupaiad), ground-squirrels and rats, on or near the forest-floor. It was collected on occasion at higher elevations, even above gooo ft. The absence of records from tree-squirrels on Mt. Kinabalu and Mt. Trus Madi is significant, since I examined at least 50 such Callosciurus in the former area, and Lim Boo Liat, M. Nadchatram, J. R. Audy and others of our colleagues also collected fleas from tree-squirrels in both regions. ILLUSTRATIONS 140. Head and prothorax (3) 152. Apical region of aedeagus (lateral) 141. Head (9) 153. Apical region of aedeagus (ventral) 142. Meso- and metathorax (3) 156. Modified abdominal segments (9) 143. Clasper and segment 9 157. Anal lobes and stylet (9) 144. Distal arm of sternum 9 158. Spermatheca 147. Modified abdominal segments (3) 159. Abdominal segments 2 (2) 148. Aedeagus 160. Spermatheca and genitalia 149. Metatibia (3) 161. Spermatheca (variation) 150. Segment 5 of metatarsus (3) 172. Apical region of aedeagus 151. Metatibia (2) 2. KEY TO THE KNOWN SPECIES OF LENTISTIVALIUS gen. nov. 1 Pronotal comb with a total of about 26-28 spines, close-set, their bases contiguous; spines narrow, about 6-5 times as long as broad at middle; most of spines parallel and similar in shape. (Malayan bird-fleas)!?_. . insolli (Traub, 1950) — Pronotal comb (fig. 140) with about 18-20 spines; spines eanetly, separated at bases; generally broad—about 5-4 times (or less) as long as broad at middle (narrow in L. fevinus, but not close-set) ; spines variable in shape and inclination, at least regarding dorsal versus ventral ones. (From mammals; in other areas) . 2 2 First preantennal row of bristles spiniform in shape. With 4-5 sinitsdne bordestae frontal margin of head. Pronotal comb!8 with spines mainly concave dorsally and with apices broad and rounded. (On shrews; Indian subcontinent and southern China) ; . ferinus (Rothschild, 1908) — First preantennal row of bristles ‘unmodified: Gay ventrall2 2 bristles bordering frontal margin (fig. 140). Pronotal comb with spines fairly straight, apices narrowed, pointed (fig. 140) or angled.'4_ (Generally on tupaiads or rodents; Africa or Asiatic- Pacific islands) é ; . A < = 3 5 5 c 3 c 3 12 See next article in series for illustrations of pronotal combs of other Lentistivalius, viz., fig. 18 for mmsollr. 13 See fig. 15 in next paper. 14 See fig. 16 in next paper. NEW TAXA OF FLEAS 281 3 With only 1 row of bristles on pronotum. Distal arm of ¢ sternum 9 lacking a subapical spur on anterior margin. Q sternum 7 with subventral sinus near middle of lower } of caudal margin. @ spiracular fossa 8 with vertical portion relatively broad, viz., less than twice as long as broad at middle. (Africa). alienus (Smit, 1958) — With 2 rows of bristles on pronotum (fig. 140). Distal arm of g sternum 9 with an anterior subapical spur (figs 144, 145). Q sternum 7 with subventral sinus much closer to ventral margin than to middle of caudal margin (fig. 156, 7S.). 9 spiracu- lar fossa 8 with vertical portion more than twice as long as broad at middle. (Borneo and Japan) . c : : : é 0 . . o é . 4 Deltoid flap of aedeagus with a ventrocaudad-directed acuminate process arising at anteroventral corner (fig. 155, DEL.FL.). Crochet-process (CR.P.) only about 2°5 times as long as broad at middle. 9 sternum 7 definitely biconcave, upper sinus on caudal 4 taller than and nearly as deep as subventral one. (Japan) aestivalis (Jameson & Sakaguti, 1954) — Deltoid flap of aedeagus relatively squared at anteroventral corner, lacking an acuminate projection (fig. 152, DEL.FL.). Crochet-process (CR.P.) nearly 5 times as long as broad at middle. 9 sternum 7 with only one distinct sinus, and that subventral (fig. 156, 7 S.). (N. Borneo) . 6 c c vomerus sp. nov, (p. 272) C. Stivalius Jordan & Rothschild, 1922, Sensw Strictu Stivalius Jordan & Rothschild, 1922. Ectoparasites 1 : 249-250. Type of genus: S. ahalae (Rothschild, 1904). Stivalius, Holland, 1969, Mem. ent. Soc. Canad. No. 61 : 14. A major contribution to our understanding of Stivalius s. lat. was made by Holland (1969) when he created seven new genera based upon New Guinean species and described and illustrated new species in these taxa. In this connection, Holland aptly and capably defined Stivalius s. str., thereby presenting a standard by which his new genera could be compared, and easing the work of those who followed him. In order to facilitate evaluation and characterization of the new genera described above in the present opus, Stivalius Jordan & Rothschild s. str. is now redescribed, citing aedeagal and some other characters not previously mentioned, so that direct com- parison with the new genera in the present paper is feasible. A new subspecies of S. cognatus Jordan & Rothschild, 1922, is described to illustrate the typical features of the taxon. e Diacnosis. Instantly separable from related taxa by: (1) the elongated sclero- tized inner tube (figs 170, 173, 174, S.I.T.) which is 9-30 times as long as broad at middle and arched at least for apical 4, and (2) the bursa copulatrix, which appears either grossly swollen throughout and coiled for most its length (fig. 176, B.C.) or else the apical 4 is folded upon itself (fig. 162) (S. phoberus Jordan & Rothschild, 1922). Also characteristic as follows: Movable finger (fig. 165, F.) long and narrowed with well developed stiva (STV.). Conical process (C.P.) broad. Ford’s sclerite (figs 170, 173-174, F.SC.) large—longer than adjacent sclerites, narrowed subapically and terminating in 1 or 2 hook-like projections. (This fact, and elongated S.I.T., probably account for the noteworthy absence of ventral paramere-like processes, €.g., the crochet lacks a caudad-directed arm.) S.I.T. bearing a distinctive basal sclerite (L.W.AR.). Distal arm of sternum 9 (figs 167-169, D.A.g) apically with a short lobe (A.LB.9) bearing bristles, and a mesal apicocaudal flap or expansion (AC.F.g) bearing 282 ROBERT TRAUB marginal subspiniforms. Labial palpus 5-segmented (excluding palpiger). Tibiae with some stout subdorsal setae next to, or incorporated within, dorsomarginals so that some of notches appear to contain 3 bristles. Full-length false tibial combs lacking and rarely with even the appearance of one on apical }. GENERIC DESCRIPTION. Head in general resembling Medwayella and Lentistivalius but somewhat longer and bearing more bristles. Head rounded and somewhat recurved in ¢ (fig. 164); “frontal’’ region here somewhat less than thrice as high (vertex to ventral margin) as long at level of uppermost bristle of row II, and about twice as long at level of eye bristle; in 2, corresponding figures are about 2-5 and 1 -6— 2 times, respectively. With head-bristles slender; preantennals in 4 rows but with several supernumeraries, which may appear like a fifth row. Eye large, reniform, with sinus ventral, subcaudal and subventral in position, just above apex of procoxa. Eye bristle (E.B.) inserted above level of eye, but in g, with caudalmost bristle of row IV just in front of eye. Ultimate (fifth) segment of labial palpus reaching apex of procoxa, or nearly so. With 3 rows of postantennal bristles, plus the usual 1 midway between last 2 rows, near antennal groove. Pronotum dorsally subequal to, or slightly exceeding, length of adjacent spines of comb; with 2 rows of bristles, first row incomplete; comb with a total of 22-24 spines, and these narrow, somewhat oblique and bluntly pointed. Mesonotum with 4 regular rows of bristles but anteriorly crowded with smaller bristles that may consti- tute 1-2 additional rows; the greater number occurring in 9, with 1 subdorsal pseu- doseta. Metanotum with 3 regular rows of bristles plus a more anterior incomplete row. Mesepisternum with about 4-5 bristles; mesepimere 5-6. Lateral metanotal area somewhat higher than long; with a dorsocaudal long bristle. Metepisternum with a dorsocaudal bristle. Squamulum short and broad in some species; long and thin in others. Metepimere usually with 12-13 bristles in 3 rows, and some of last row accompanied by 1-2 small intercalaries. Tibiae with 7 groups of stout dorso- marginal bristles (including apical one) ; in pro- and mesotibiae, usually with 4 groups containing 3 stout bristleseach. Metatibia with at least 5 distinct notches containing dorsomarginals and these generally with 2 stout bristles and with 1~2 of subspiniform type below the notch; lacking full-length false combs, and in only one species (S. aporus Jordan & Rothschild, 1922) does even apical 4 appear comb-like. Fifth tarsal segment with 6 pairs of plantar bristles, of which fourth pair (from base) somewhat displaced towards midline; in pro- and mesotarsi, first pair may also be slightly displaced. None of tarsi with apical bristles extending beyond middle of next segment (except perhaps for distalmost of fringe of segment 1 of protarsus in both sexes). Proportions of tarsal segments essentially as that noted for Medwayella dryadosa. First abdominal tergum with 4 rows of bristles, remainder of unmodified abdominal terga with 2 4 rows in g, 3 in 9; ventralmost bristle of caudal row below spiracular fossa. Terga 2-5 with I apical spinelet on each side. Basal abdominal sternum without setae in g, although these often represented by 6-9 microsetae or their bases; in 2 with about 16-24 small bristles in 2 or 3 highly irregular, oblique or sub- vertical rows, commencing subdorsally near anterior margin and extending sub- medially to about ventral 4, at caudal 4; ventralmost bristles accordingly the most NEW TAXA OF FLEAS 283 caudal. Representative sterna in J with a caudal row of 3 long subventral bristles, at times surmounted by a small submedian one; with an additional 2 short rows of small subventrals preceding this, plus an anterior 1-2 small ventromarginals making a total of 11-16 in all; in 9 with about 18~20 bristles in same general pattern but more bristles per row, generally including 5 long bristles in caudal row. Tergum 7 in ¢ with a short dorsal lobe (fig. 165, U.L.7) extending between the sets of antepygidial setae (A.B.); in 2, both upper lobe (fig 166, U.L.7) and lower lobe (L.L.7) well deve- loped and acute. Three bristles of 2 7 T. modified so as to somewhat resemble A.B. and somewhat displaced thereto ; uppermost (U.M.B.) at times quite similar to upper A.B., first of lower such (L.M.B.) often contiguous to plate of A.B. and closely resembling upper A.B., generally more so than second (L.M.B.-2). Dorsal A.B. about 4 length of ventral one in J; usually slightly longer than 4 in 9. Male. Tergum 8 reduced; resembling that of Medwayella. Sternum 8 relatively unmodified, lacking a group of close-set bristles or spiniforms; very large, enclosing most of genitalia; extending dorsad to base of anal segments and cephalad to level of base of A.B.; about 1-4 times as long (near ventral margin) as high at maximum level (at cephalodorsal angle). Dorsal margin of 8 T. fairly straight to caudal } or beyond; and in some species horizontal; in others, oblique at about angle of 30°; ventral margin varying inversely regarding slope so that narrowing effect is the same, viz., caudal margin about } height of cephalic. Manubrium (fig. 165, MB.) broad all the way to near apex; ventral margin biconvex near middle; with a short dorsal distal tubercle. Immovable process of clasper (P.) broadly rounded apically. Conical process (C.P.) fairly broad, at times about 5 times as tall as broad at middle; but in some taxa only about 3 times. Movable finger (F.) very long and thin, about 6-5-7 times as long (from apex of P.) as broad near base of distal fringe (D.FR.), narrowed for most its length; apical margin quite flat. Stiva (STV.) fairly long and narrow. Distal fringe (D.FR.) of about 6-8 bristles commencing on shaft of F. and proceeding onto STV. Sensilla-group (S.G.) consisting of 3 short bristles. Fulcral sclerite (F.S.) higher (longer) than broad; subvertical; broadest at dorsal end; natrowest subventrally. Proximal arm of sternum 9g subvertical; with basal 3 at angle of about 60°; uppermost nearly perpendicular; upper } of caudal margin bi- concave. Distal (“‘horizontal’ or “‘ventral’’) arm of sternum 9 (figs. 167-169, D.A. 9) broad; angled caudad dorsally by virtue of a large mesal flap or extension, the apico- caudal flap (AC.F.9) which is fringed with subspiniforms; apex or subapical region of straight upright portion lobate (A.LB.g) and clothed with thin marginal bristles; with a short apical or subapical, anterodirected spur. D.A.g apparently with ventral margins fused to about apical third at level of marginal transverse sclerotization (T.S.). Aedeagal apodeme resembling manubrium for most its length, though broader at apex and lacking an apical tubercle or spur. Its middle lamina (figs 170, 174, M.LAM.) about 3-3-3-6 times as long (to near anterior end of crescent sclerite (C.S.)), or caudal margin of lateral shafts of capsule (L.S.C.) as broad at anterior margin of its bay (B.M.L.). Lateral laminae (L.LAM.) lightly tanned and most noticeable at anterior end, where each extends ventrad of M.LAM.; with ventrocaudad extensions arising from dorsum, near anterior end of B.M.L., and crossing bay but scarcely Fe 284 ROBERT TRAUB discernible until underlying proximal arm of sternum 9 (P.A.g), and here appearing as a well tanned line (blending with extensions of girdle (G.) of wall of aedeagal pouch) which continues across M.LAM. to base of Y-sclerite (Y.S.), where it meets the virtu- ally inapparent ventral margin. Ventral wall of aedeagal pouch (AE.P.-V.) well sclerotized, extending to base of body of crochet (B.CR.). Median dorsal lobe (M.D.L.) evenly curved; gradually arching dorsad from level of base of sclerotized inner tube (S.I.T.) ; height of convexity near level of base of Ford’s sclerite (F.SC.) and here only slightly arched. Hood (HD.) relatively tall (long) and narrow; its ventral margin continuing cephalad as the deltoid flap (DEL.FL.) to about level of middle of B.CR.; here DEL.FL. curving dorsad (at times recurving dorsocaudad) to level of pivotal ridge (PIV.R.), in line with base of phylax (PHY.). Anterodorsal angle of DEL.FL. not highly acuminate. Lateral lobes greatly reduced; inapparent. Sclerotized inner tube (S.I.T.) generally so greatly elongated as to be vermiform; apical $ or } arched to form a semi-ellipse; at least 7-12 times as long as broad at middle, and a minimum of 5 times as long as broad at base of phylax; usually 11 times as long as broad at phylax and in some instances, more than 20 times as long as broad. S.I.T. characterized by a sclerite which arises ventrally near its base and loops around the sides of the tube, and which is herein termed lateral wings of ventral armature (L.W.AR.). S.I.T. witha short protrusion from apex (figs 170, 171, FIS.) equivalent to girth of lumen of S.I.T. and dorsally somewhat tanned, appearing as if an extension of a tube within a tube and regarded as comparable to the “‘fistula’”’ (Peus, 1956), or ‘outer part of inner tube’ (Holland, 1955) or “band of inner tube” (Traub, 1950) associated with ceratophyllids, etc., although much shorter in Stivalius. Ford’s sclerite (F.SC.) massive, the largest structure in the endchamber; usually flask-shaped or at least constricted subapically ; with 1 or 2 claw-like apices (in lateral aspect); not differentiated into securifer and alpha-portions. Phylax (PHY.) well developed as a large sigmoid or bowed sclerite crossing over S.I.T. near base and extending to level of lightly tanned pivotal ridge (PIV.R.), but well anterior to base of F.SC. Pivotal chord (PIV.CD.) feebly sclerotized. Crochet notable in virtually complete reduction of ventral arm, and hence lacking a crochet-process or paramere- like structure. Body of crochet (B.CR.), the upright arm of crochet of other related taxa, well represented, however; relatively long and narrow; base extending from ventral portion of endchamber; apex flattened and expanded, at base of F.SC.; at times (e.g., S. phoberus) with a ventral, short, caudad-directed spur, probably repre- senting a vestige of crochet-process. Crescent sclerite (C.S.) large; apex within base of S.I.T. and hence satellite sclerite (SAT.S.) wholly within. Fulcral lateroventral lobes (FUL.L.L.) short. Central sclerite (CEN.S.) appressed to fulcral medial lobes (FUL.M.L.). Lateral shafts of capsule (L.S.C.) well tanned. Y-sclerite (Y.S.) very large; both components visible in mounted specimens. Vesicle (V.) and ventral virga (V.V.) well developed. Penis rods short, scarcely reaching anterior end of apodeme when in situ. Third apodemal rod (AP.R.) of endophallus and dorsal virga (D.V.) lightly tanned, especially the latter. Caverna spiculosa (CAV.SPIC.) sub- median in position and anterior to level of fulcrum. Female. Spermatheca (figs 176, SP., 163, 177) with bulga (B.) longer than hilla (H.); bulga broader (taller) caudally than anteriorly and at times constricted in NEW TAXA OF FLEAS 285 middle ; hilla with an apical papilla and somewhat inserted into lumen of bulga. Duct of spermatheca (figs 163, 176-179, D.SP.) with internal striae in dilated portion (DIL.P.) near junction with bursa copulatrix (B.C.), which in turn is thick-walled, very broad most of its length, and generally (fig. 163) markedly involute, or at least coiled or folded upon itself (fig 162), apically, if not elsewhere as well. B.C. with perula not clearly differentiated; with part of inner wall of lumen at times well sclerotized and appearing like a curved band. Wall of roof of oviduct (fig. 163, OV.) well tanned as it (or paired members thereof) approaches vagina (VAG.). Parageni- tal morion reduced, inapparent ; apparently represented by indiscrete lines on each side of B.C., near vagina. Sternum 7 (fig. 175, 7 S.) with ventral 3 of caudal margin biconcave, but upper sinus at times quite small (as in figure). Ventral anal lobe (fig. 176, V.A.L.) with base short; apical margin long; anterior 4+ of margin somewhat convex and bearing 4-5 close-set, long bristles ; remainder of margin sinuate, bearing 2 long bristles at apical }or}. Analstylet (A.S.) long and narrow, parallel-sided and lacking long subapical bristles. Mesal genitalic ridge of tergum 8 undeveloped. Tergum 8 (8 T.) with a short lobe at ventrocaudal angle. Spiracular fossa 8 (fig. 166, 8 SPC.) medium-sized ; upright portion ovate, not much longer than broad. Sternum 8 (8 S.) flask-shaped and with short apical bristles. CoMMENT. Species included: (1) S. ahalae (Rothschild, 1904), the type of the genus; from rats, south India; (2) S. aporus Jordan & Rothschild, 1922, from Rattus, Millardia and other murines from Ceylon, India and Nepal. (A subspecies of aporus, or a closely related new species, occurs in Thailand and Vietnam on rats.) (3) S. phoberus Jordan & Rothschild, 1922, from rats in Ceylon. (4) S. c. cognatus Jordan & Rothschild, 1922, from Rattus (Rattus) in Java; S. c. spiramus Jordan, 1926 (new status), from R. (Rattus) in the mountains of Luzon, Philippines, and the new subspecies next described, from FR. (Rattus) in the mountains of Mindanao, Philippines. (5) S. rectodigitus Li & Wang, 1958. Even though the authors state this is near S. klossi Jordan & Rothschild, 1922, and compared it with that species, it definitely is close to S. aporus (and may be a subspecies thereof) and hence is a member of Stivalius s. str«-while S. klosst belongs to a separate and yet un-named genus. The status of S. rectodigitus is clear, as indicated by Li & Wang’s figures of the bursa copulatrix and of the distal arm of the ninth sternum, etc. Further, those authors point out the aedeagal similarities with S. aporus. Further discussion of the genus is presented after the treatment of S. cognatus below. 1. Stivalius cognatus Jordan & Rothschild, 1922 Stivalius cognatus Jordan & Rothschild, 1922. Ectoparasites I : 253, 264, figs 243, 245; Jordan, 1933, Novit. zool. 38 : 355; Costa Lima & Hathaway, 1946, Pulgas: 326; Traub, 1951, Proc. biol. Soc. Wash. 64: 13; Smit, 1958, Bull. Brit. Mus. (Nat. Hist.) Ent. 2(2): 41. Diacnosis. Immovable process (fig. 165, P.) very short and broadly rounded at apex ; with a small subapical caudomarginal bristle. Movable finger (F.) with portion above apex of P. about 6-3 times as long as broad at level of ventralmost bristles of distal fringe (D.FR.); slightly narrowed near basal portion of D.FR., otherwise Margins quite straight and parallel; apex subtruncate. Stiva (STV.) of F. well 286 ROBERT TRAUB developed; longer than unexpanded portion of apex and about 2-7 times as long as broad. Sclerotized inner tube (figs 170, 173, S.I.T.) elongated, but not nearly as long as in S. ahalae and S. aporus, i.e., extending only slightly beyond apex of F.SC., instead of for more than } its length; with its are paralleling that of median dorsal lobe (M.D.L.) instead of being much more complex. Ford’s sclerite (F.SC.) narrowed near middle of sclerite, flask-like ; with basal portion (proximad of dorsal spur or peak) subequa’ to portion distad, instead of being 3 length of neck of flask. Basal portion of F.SC. only 1-7 times as long as broad at level of dorsal spur instead of 3-5 times. Distal arm of sternum 9 (figs 167-169, D.A.g) with only one spur on anterior (dorsal) margin, occurring at base of apical rounded lobe (A.LB.g), which in turn is not as high as long, instead of vice versa. Sinus above ventral lobe of 9 sternum 7 (figs 163, 176, 7 S.) somewhat similar in size and shape to that of lobe below it. Bursa copula- trix (figs 163, 177) with convolute portion quite symmetrical in size and shape. Spermatheca (fig. 176, SP.) with bulga (fig. 177, B.) somewhat constricted near middle; caudal } more convex dorsally than cephalic portion; slightly more than twice as long as high immediately anterior to constriction (maximum internal mea- surements); caudal and ventral margins externally quite straight, only slightly convex. Hilla (H.) with about basal } inserted in lumen of bulga; somewhat squarely recurved at midpoint and slightly dilated above this curve; about thrice as long as broad near bulge; apex broadly rounded. Apical papil'a (PAP.) about 4-3} diameter of H.; somewhat longer than broad. For further details of aedeagus, see section I.c. below, on new subspecies. Comments. S. cognatus, like other members of the genus (s. stv.), is a typical para- site of members of the subgenus Rattus. The original description was based upon material from Java, but Traub (1951) quoted Karl Jordan, F.R.S., as expressing the belief (in litt.) that S. spivamus Jordan, 1926, from Luzon, known only in the female, was the same species. Jordan’s perspicacity was demonstrated when the writer collected topotypic material of S. sbivamus in 1961 and found that the specimens were indeed cognatus, but of subspecific rank. This subspecies is described below, along with a new one from Mindanao. Undoubtedly other subspecies, and perhaps related species, occur elsewhere in the Indonesian and Philippine Archipelagoes, and our understanding of zoogeography would be enhanced if such material became available for study. 1. a. Stivalius cognatus cognatus Jordan & Rothschild, 1922 DiaGnosts. Characterized by the following features: (1) Distal arm of sternum 9 (fig. 168, D.A.9) with apical lobe (A.LB.g) about 2-4 times as long as high. (2) Api- cocaudal flap (AC.F.9) of D.A.g about 1-6 times as long (from level of margin of transverse sclerotization, T.S.) as broad at middle. (3) Phylax of aedeagus fairly straight and narrow, about 5 times as long as broad at maximum (at middle); scarcely broader at middle than subapically. (4) Ford’s sclerite with total length 1-9 times length of basal section (portion anterior to dorsal angle or nubbin) and about 2-5 times breadth at apex of basal section. (5) Bursa copulatrix (fig. 179, B.C.) with coil oblate, rather flattened dorsally. (6) Vertical diameter of involute portion NEW TAXA OF FLEAS 287 exceeding distance from bottom of loop to vagina (VAG.). (7) Lateral metanotal area usually longer than high (internal measurements). ComMENT. The only published records for the nominate form are those in the original description, viz., from Rattus (R.) rattus (cited as Epimys rattus), from Malang, Java. The specimens studied and the female here illustrated, are from an unrecorded area in Java, coll. M. V. Kuhlavain, and determined by the late Karl Jordan, F.R.S., and received through his kind offices. ILLUSTRATIONS 168. Distal arm of sternum 9 179. Spermatheca and bursa copulatrix I. b. Stivalius cognatus spiramus Jordan, 1926, NEW COMBINATION Stivalius spiramus Jordan, 1926. Novit. zool. 33 : 391, fig. 12; Costa Lima & Hathaway, 1946, Pulgas: 327; Traub, 1951, Proc. biol. Soc. Wash. 64 : 13. Diacnosis. (1) Distal arm of sternum 9 (fig. 169, D.A.g) with apical lobe (A.LB.9) about twice as long ashigh. (2) Apicocaudal flap (AC.F.g) about 1-7 times as long as broad at middle. (3) Phylax (fig. 173, PHY.) symmetrically sigmoid, narrow; about 5 times as long as broad at maximum near middle. (4) Ford’s sclerite (F.SC.) 1-6 times as long as its basal portion and 1-9 times as long as broad at apex of proximal region. (5) Bursa copulatrix (fig. 178, B.C.) with involute port on quite circular in outline. (6) Vertical diameter of loop exceeding distance from bottom of coil to level of vagina (VAG.). (7) Lateral metanotal area usually longer than high (internal measurements). ComMENT. S. spiramus was described from, and heretofore known only from, a single female collected by Professor E. H. Taylor at Baguio Bengue (Luzon) from a rat cited by Jordan as ‘‘Rattus guereci.”” (This name probably was a lapsus for querceti Hollister, 1911, a Luzon rat which is placed by Ellerman (1941) in the Rattus (R.) concolor group but which is treated by Schwarz & Schwarz (1967) as a synonym of Rattus rattus negrinus Thomas, 1898.) The present description and figures are based upon specimens which I collected at Baguio, at 3600-4500 ft elev. in October, 1961, ex Rattus (R.) vattus ssp., and which can be considered as topotypic. The allotype 3g is therefore selected as follows: Allotype g¢ (B-55028) ex Rattus exulans; PHILIPPINES: Luzon, Baquio; elev. 3600 ft; 24.X.1961; Coll. R. Traub; deposited in the U.S. National Museum. ILLUSTRATIONS 169. Distal arm of sternum 9 175. Sternum 7 (2) 173. Apical region of aedeagus 178. Bursa copulatrix I. c. Stivalius cognatus bamus subsp. nov. TYPE MATERIAL. Holotype 4, allotype 2 (B-56383) ex Rattus vatius mindanensis ; PHILIPPINES: Mindanao, Zamboanga del Norte, Mt Malindang, Masawan, Mutia; elev. 4500-5000 ft; 29.XII.1962; Coll. D.S. Rabor. Paratypes as follows: Collected 288 ROBERT TRAUB by D. S. Rabor, 24.XI1.1962/14-16.1.1963; same locality as holotype; 27 g, 36 9 ex Rattus vattus mindanensis ; 2 9 ex R. rabori; 2 2 ex R. exulans todayensis; I 3, I 9 ex R. pantarensis; 4 3, 4 9 ex Apomys insignis bardus; 1 3, I 9 ex Rattus sp., Ibid. but elev. 5200-7000 ft; 2g, 3 2 ex R. r. mindanensis; Mt Malindang, Canon, Mutia; elev. 3200 ft; 22-24.XI1.1961. Holotype (U.S.N.M. number 71610), allotype and 3 pairs of paratypes deposited in the U.S. National Museum. The remainder distributed as for M. dvyadosa. Diacnosis. (1) Distal arm of male sternum g (fig. 167, D.A.g) with apical lobe (A.LB.9) about 2-6 times as long as broad. (2) Apicocaudal flap (AC.F’.9) relatively short and broad, viz., about 1-3 times as long (from level of caudal margin of trans- verse sc'erotization, T.S.) as broad at middle. (3) Phylax (figs 170, 174, PHY.) eccentrically sigmoid; upper 4 narrower than lower } and much more curved; rela- tively broad, 1.e., only 3-4 times as long as broad at maximum breadth (below middle). (4) Ford’s sclerite (F.SC.) with basal portion (to level of dorsal nubbin) only 4 total length of F.SC. and about 2-6 times breadth of sclerite at level of dorsal peak. (5) Bursa copulatrix (figs 163, 176, 177, B.C.) with upper portion evenly involute; symmetrical. (6) B.C. relatively long, so that vertical diameter of loop is merely equal in length to lower portion of bursa, 1.e., from ventral margin of coil to vagina (VAG.), instead of exceeding it. (7) Lateral metanotal area higher than long (inter- nal measurements). DESCRIPTIVE Notes. Since the aedeagus of this species has never been described before, further notes are included here, including comments on this organ as seen from the ventral aspect. It should be borne in mind, however, that most of the points (save for those cited in the diagnosis) are on the species level. Aedeagus—Lateral Aspect (figs 170, 174). Aedeagal apodeme with middle lamina (M.LAM.) about 3:6 times as long (to level of caudal margin of lateral shafts of capsule (L.S.C.) as broad at anterior margin of its bay (B.M.L.). Hood (HD.) with ventral margin shallowly concave save for curves at anterior and posterior margins; lightly reticulated apically; apparently laterally reinforced by transverse parallel light striae, especially over caudal 4, which is the deltoid flap (DEL.FL.). HD. and DEL.FL. extending caudad to base of phylax (PHY.) and body of crochet (B.CR.). Median dorsal lobe sloping at about angle of 30°commencing at level of slightly acute anterodorsal angle of DEL.FL.; maximum height of its gentle, broad arc is at level somewhat apicad of Ford’s sclerite (F.SC.) and thereafter sweep of curve paralleling that of dorsal margin of F.SC. (if not pivoted out of position). Aedeagal pouch (AE.P.) with ventral border well tanned; but anterior wall (girdle, G.) less so, and with its cephalodorsad extensions soon meeting the ventrocaudal prolongations of lateral laminae (LLLAM.). Ford’s sclerite (F.SC.) with length almost equal to that of body of crochet (B.CR.) and quite broad for proximal }, which resembles base of a flask; basal } with dorsal and ventral margins fairly straight and parallel; narrowed portion of flask slightly more than } length; lower margin sinuate. F.SC. with breadth of middle region of anterior } equal to that of basal part of sclerotized inner tube NEW TAXA OF FLEAS 289 (S.I.T.); apex oblique and subtruncate, terminating in a ventrocaudal projection. S.LT. with apical 3 curved in are almost paralleling that of M.D.L. but extending more distad, to a level slightly beyond apex of F.SC.; approximate breadth of middle of tube slightly more than 3 of that of subapical region of F.SC.; about 10-14 times as long as broad at level immediately proximad of B.CR. Lateral wings of ventral armature (L.W.AR.) somewhat ovate, but narrower dorsally than ventrally and reaching to near dorsal margin of S.I.T. Crescent sclerite fairly long; with a pre- apical anterior hump; narrowed caudally and with caudal 4 penetrating S.1.T. where it is virtually contiguous with angled, relatively long satellite sclerite (SAT-S.). Central sclerite (CEN.S.) closely appressed to fulcral medial lobes (FUL.M.L.) and resembling acute wing of an arrowhead. Y-sclerite (Y.S.) with one fork narrow, horizontal and extending to level of base of SAT.S.; larger fork very conspicuous as a diamond-shaped sclerite below fulcrum. Dorsal virga (D.V.) indistinct. Phylax (PHY.) a fairly broad, bowed sclerite which is upcurved below midpoint and some- what narrowing apically, about 2-3 times as long as broad at maximum girth (below middle) (distinctly narrower in other subspecies). Body of crochet (B.CR.) ribbon- like except for flattened, expanded apex; axis flattened except for somewhat concave extremities; ventrally projecting somewhat below S.I.T. as a narrowed, acute rod, this portion presumably representing a vestige of the crochet-process (paramere) of other genera. Lacking discernible tanned lateral connections between AE.P. and B.CR., with what is apparently a small seta on apical margin of AE.P., on each side, at base of B.CR. and of PHY. With the upper penis rod (P.R.) extending to near level of anterior end of aedeagal apodeme when 7m situ; lower rod much shorter. Ventral virga stout; sclerotized for entire length, viz., to near level of base of bay of middle lamina. Third apodemal rod (AP.R.) lightly tanned once leaving AE.P. Caverna spiculosa (CAV.SPIC.) irregularly ovoid; about twice as long as broad; sloping cephaloventrad at an angle of about 60°. Aedeagus—Ventral Aspect (fig. 171). Hood with anterior margin entire, lightly tanned. Ford’s sclerite shaped like a tulip flower, about 1-3 times as long as broad near base; without any caudad-directed paramere-like structures; the hook-like apex of the lateral aspect actually representing a deep structure extending all the way across the sclerite which is seen on edge. Deltoid flap (DEL.FL.) covering only sides of ventral margin. Phylax (PHY.) actually with base (ventral) fairly narrow and constricted; straddling S.I.T. and apices meeting dorsally above it; therefore projection of PHY. in ventral aspect is that of a trapezoid with thickened sides. Body of crochet (B.CR.) dorsally buttressed against base of F.SC. S.I.T. appearing as a long, narrow cylinder whose only modifications are an apex that is slightly recurved. ventrad, viz., the fistula (FIS.), and a narrow, basal belt, viz., the lateral wings of the armature (L.W.A.R). Wall of aedeagal pouch well tanned both laterally (AE.P.-L.) and ventrally (AE.P.-V.); lacking sclerotized connections with body of crochet (unlike the condition in pygiopsyllids with ventral crochet-processes). Vesicle (V.) quite large, and ventral virga (V.V.) broad. Y-sclerite (Y.S.) with bifid anterior base and broad caudal stem flooring middle of capsule, which is quite broad, distance between fulcral lateroventral lobes (FUL.L.L.) nearly equal to length of fulcral area. Caverna spiculosa (CAV.SPIC.) lateral in position. 290 ROBERT TRAUB CoMMENT. (1) The vast majority of records of S. c. bamus are from Rattus rattus mindanensis, with a few from the related murine Apomys, all from Mt Malindang at elevations between 3200-5000 ft. As yet material from other parts of Mindanao is unavailable for study, so that the true range of the subspecies is unknown, and also it cannot be stated that this is the only subspecies of S. cognatus occurring on the island. (2) The aedeagus of S. cognatus. The structure of the aedeagus as seen in the ventral aspect emphasizes and clarifies some of the points made above, in the consideration of the genus. Thus, the lack of overt connections between the walls of the aedeagal pouch and the crochet is quite striking and is in marked contrast to related taxa. The difference seems to be associated with the presence of a well developed quasi- crochet or ventral crochet-process: if these are present, as in Medwayella and Lentistivalius, there are definite links between the crochet and the pouch wall; if absent, as in Stivalius s. sty., there are no discernible ties. It also is apparent that Ford’s sclerite does not bear a pair of narrow hooks or prongs, as one might infer from the lateral aspect. Instead, the apex bears a downward directed flange or lip which extends from one side to the other, and which presumably acts like a prying lever when F.SC. is pivoted ventrad. Bristles or bristle-lke structures are virtually unknown on the aedeagus, and the few isolated examples known to me were regarded as atavisms or freaks. However, the fine seta on each side near the caudoventral margin of the pouch wall in S. cognatus can be seen in most specimens and even seems to arise from an alveolus. Since many of the components of the phallosome of fleas are paired, it seems logical to assume the organ is derived from an ancestral bilaterally symmetrical structure, and that this had borne bristles. If so, a setal vestige is not surprising and is in accord with my belief that pygiopsyllids are in general primitive fleas despite certain adap- tive specializations, which at times are quite marked. (3) Abnormal structures of fleas are often of interest in that they may represent atavisms or anomalies that suggest the origin of the normal organ. A freak bristle is therefore labelled FRK. in fig. 165, although no further comments can be made at this time on its possible significance. ILLUSTRATIONS 163. Spermatheca and genitalia (2) 170. Apical region of aedeagus (lateral) 164. Head and prothorax (3) 171. Apical region of aedeagus (ventral) 165. Processes of clasper 174. Apical region of aedeagus 166. Antepygidial region (9) 176. Spermatheca, genitalia and anal seg- 167. Distal arm of sternum 9 ments 177. Spermatheca and bursa copulatrix 2. COMMENTS ON Stivalius s. str. Host-Relationships, Distribution and Potential Vectorship of Disease. Throughout its known range (Ceylon, Indian Subcontinent, S.E. Asia, Indonesia and the Philippines) the genus Stivalius s. str. is primarily associated with murine rodents, especially the subgenus Rattus. Blood-sucking ectoparasites of murines are of special interest regarding transmission of disease, but the role of Stivalzus s. str. NEW TAXA OF FLEAS 291 in this regard is in need of clarification. Stivalius cognatus, reported as Pygiopsylla ahalae in Java according to Pollitzer & Meyer (1961), and, Stivalius species ‘‘have been found capable of transmitting plague in Java and south India”’ respectively (Pollitzer, 1954). Inasmuch as the studies referred to by Pollitzer concern Stivalius from rats, the species in question are almost undoubtedly members of Stivalius s.str., e.g., S. ahalae, S. cognatus and S. aporus (or S. phoberus, if it occurs on the mainland and not merely Ceylon), since the only other Indian “‘Stivalius’’ is the species referred to above, in the present paper, as Lentistivalius ferinus, and is a parasite of shrews. The role of Stivalius in the ecology of both human and murine plague needs elabora- tion especially since this disease is still endemic in areas where plague exists today in Indonesia, Thailand and Vietnam and where members of this genus are known to, or may be expected to infest Rattus (Rattus). The possible involvement of Stivalius s. sty, in the ecology of murine typhus should be investigated, for this flea-borne disease is likewise associated with the subgenus Rattus and is endemic, or presumed to occur, in regions where Stivalius s. str. infests commensal rats. The Specialized Aedeagus and Bursa Copulatrix Stivalius s. str. exhibits some remarkable genitalic modifications that are not seen in related taxa, many of which have developed other complexities in the sexual apparatus. The marked elongation of the sclerotized inner tube (figs 170-171, 173- 174, S.I.T.) is quite striking as compared to Lentistivalius, for example (fig. 172). Moreover, the apex of S.I.T. is further specialized in Stivalius s. str. by the apically protruding, somewhat tanned fistula (fig. 170, FIS.). There are scores of species referable to Stivalius s. lat., and it can be no coincidence that females in which the bursa copulatrix is broad, sausage-like and coiled (and no others) are the mates of males with snake-like aedeagal tubes. Such parallel developments in the two sexes of Sizvalius s. str. in the course of evolution strongly suggest a functional and funda- mental relationship, namely that, in this taxon, unlike the case in most fleas, S.I.T. itself, and not merely the penis rods,15 enters well into the bursa. This then would explain the thickened walls of the bursa and the sclerotized rod-like lining of one side of the lumen of the bursal duct seen in this genus. Two points are mentioned here as rendering support to this hypothesis. The characteristic absence of ventral ‘‘parameres’’ in Stivalius s. str. indicates that the function of the caudad-directed, finger-like or rectangular crochet-process (CR.P.) of Telated taxa has been taken over by some other structure. The crochets of cerato- phyllids, leptopsyllids and some other fleas serve to secure and maintain a hold on See 15 Holland (1955) reiterated the view of Snodgrass (1946) that the penis rods “. . . are the only elements of the intromittant apparatus that are capable of being protracted from the aedeagus . . .” When Holland (p. 245) speaks of the “long outer part of the aedeagal inner tube’’ (not to be confused with sclerotized inner tube) as being “‘inserted well within the bursa copulatrix,’’ and other writers make similar statements, they are referring to a structure that is regarded as already being outside of S.1.T. Thus, there is no contradiction, for the outer tube is not being protracted from S.I.T. This outer tube is the structure referred to above as the fistula (FIS.) Moreover, it should be borne in mind that Snodgrass, Traub (1950), Holland, Goncharov (1964 a & b) and others, when discussing the copulatory apparatus or copulation in Siphonaptera, were referring to specific taxa of fleas and their observations do not necessarily apply to other fleas. Pygiopsyllids and thopalopsyllids in particular have not been studied in this regard. G 292 ROBERT TRAUB the female genitalic apparatus and to open some of the components to the male organs (Holland, 1955, Goncharov, 1964). Presumably the unusually well developed Ford’s sclerite (F.SC.) of Stivalius s. sty. assumes some of these functions in the absence of the crochet-process, as is suggested by the hook-like appearance of its apex (fig. 170), which, as can be seen in ventral aspect (fig 171) is really a down-projecting flange and which presumably acts as a pry when the apex of F.SC. is tilted ventrad. It also seems logical to believe that the hypertrophied and specialized S.I.T. takes over other functions, especially since there are instances within the Order wherein the aedeagal tube apparently is the true penetrating copulatory organ, and not the penis rods. Thus, Barnes and Radovsky (1969) cite impressive morphological evidence for this belief, in the case of Tunga Jarocki, 1838, and Suter’s figures (1964) of copulating Tunga seem to bear this out. The second point suggesting that these modifications in the male and female of Stivalius s. sty. are correlated, is theillustrative case of S. phoberus Jordan & Rothschild, 1922. Here S.1.T. is not as narrowed and elongated as in other species of the taxon, i.e., it is only 5 times as long as broad at level of phylax, instead of 9-20 times. In this species (fig. 162) the bursa is not as modified either: it is less involute and not quite as broad throughout (cf. fig. 163, S. cognatus banws). In Medwayella and Lentistivalius, which possess well developed caudad-directed ventral crochet-processes, there are obvious tanned connections between the wall of the aedeagal pouch and the crochets. It is noteworthy that in Stivalius s. str., which lack ventral ‘‘parameres”’ and crochet-processes, such bonds are inapparent, even in the ventral aspect (fig. 171), as is pointed out below. A test of this hypothesis may be provided by another group of pygiopsyllid fleas in which the basal half of the bursa copulatrix is very broad and thickened, namely the three species of Acanthopsylla Jordan & Rothschild, 1922, in which there is but 1 (not 2 or 4) spiniforms on the frontal region of the head, viz. A. praxilla M. Rothschild, 1934, A. guba Smit, 1953 and A. richardsoni Smit, 1953. In the other members of the genus (which includes a total of 16 species), the ventral portion of the bursa is not conspicuously dilated and lacks ring-like striations. In these 13 other species, the sclerotized inner tube of the aedeagus is not elongate, corresponding to the relative size typical for the bulk of the pygiopsyllids. In the A. praxilla- group the male is known only for one species (A. vichardsoni), which I have not seen, and its aedeagus has never been figured. However, the shape of the bursa copulatrix in this group of fleas is so suggestive of the condition in Stivalius s. sty. that I expect that here too the sclerotized inner tube of the male must be elongate, if not ophidian in shape, and that ventral crochet- like processes are lacking. Evolutionary Status of STIVALIUS S. STR. It is noteworthy that this taxon, which is the most highly specialized of this group of pygiopsyllids with respect to modifications of the aedeagus and the co-functional bursa copulatrix, consists of characteristic parasites of the subgenus Rattus, the most modern and highly developed of the murines. This suggests that Stivalius s. str. is likewise at the pinnacle of the evolutionary development of this complex of genera to date. These points are discussed further in the two following articles in the series. I 2 NEW TAXA OF FLEAS 3. Kery1!6 To THE KNOWN SPECIES AND SUBSPECIES OF STIVALIUS JORDAN & ROTHSCHILD, 1922, S. str. Male with scolerotized inner tube of aedeagus (figs 170, 173, S.I.T.) at least 9 or more times as long (from base) as broad at base of phylax (PHY.) and 20-30 times as long as broad at middle. Female with bursa copulatrix (figs 163, 178, B.C.) markedly involute, the coil(s) quite symmetrical and not overlying vertical (ventral) portion of tube; often large, involving most of length of bursa. (Indian Subcontinent to Philippines) . Sclerotized inner tube only Bbont 5 times as lone as sibroad at base of phylax Ail about g times as long as broad at middle. Bursa copulatrix (fig. 162, B.C.) folded upon itself at upper 4, the loop asymmetrical. (Ceylon). : phoberus J.& R Arc of S.I.T. (figs 170, 173) shallow, equivalent to curve of median dorsal lobe (M.D.L.), extending only slightly beyond apex of Ford’s sclerite (F.SC.), i.e., only for a distance equivalent to less than 4its totallength. Basal portion of F.SC.,i.e., region proximad to subapical constriction and below nubbin (peak or spur) on dorsal margin, equals about } length of F.SC.; this section only 1-7 times (or less) as long as broad at level of spur. Distal arm of sternum 9 (figs 167-169, D.A.g) with only one spur on anterior (dorsal margin, and that at base of apical rounded lobe (A.LB.9)). Subventral sinus of 9 sternum 7 (figs 163, 176, 7 S.) almost an inverted and reversed image of ventral lobe, its dorsal margin continuing caudad to or beyond level of margin of ventral lobe before turning dorsad. Ventral lobe of 7 S. narrower than basal part of sinus above it. (Indonesia and Philippines) cognatus subsp. S.1.T. with arc much more convex than M.D.L. and equivalent to } a broad ellipse, extending beyond apex of F.SC. by more than }its length. Basal portion of F.SC. at least # length of F.SC. and at least 3-5 times as long as broad at level of nubbin (peak or spur) on dorsal margin. D.A.g9 with 2 spurs on anterior (dorsal) margin; the additional one proximad of that at base of apical rounded lobe. Subventral sinus of 27 S. incomplete, not shaped like ventral lobe due to its dorsal margin being more oblique than ventral one or else sharply curving dorsad near base of sinus. Ventral lobe of 7 S. broader (taller) than basal part of sinus above it. Unden Subcontinent and Ceylon er Yunnan or Southeastern Asia) Dorsolateral bristles of hind-tibia with bristles of groups dissimilar in size, he coups fairly widely separated from one another so that there is no resemblance to a comb. S.LT. extremely long and narrow, at least 18 times as long as broad at base of phylax. F.SC. with only 1 apical prong. Phylax greatly dilated at middle; both margins here convex; only about 2-4 times as long as broad at middle. 97S. with upper angle of dorsal sinus acute and projecting as a short thumb ahalae (Roths., 1908) Dorsolateral bristles of hind-tibia with inner members of groups often similar in size and girth and so arranged that apical } of tibia bears a nearly perfect comb. S.1.T. elongate, but not so markedly, about 15 times as long as broad at base of phylax. F.SC. with 2 apical prongs. PHY. with only anterior margin convex at middle; about thrice as long as broad at middle. 9 7 S. with upper angle of dorsal oe) sinus short, not projecting asa thumb . Distal portion of F.SC. (apicad of nubbin) relatively aneadl, only, about fritee as long as broad. Notch at apex of F.SC. with prongs converging somewhat at tip. Seventh sternum of female with lobe above subventral sinus extending more 293 ., 1922 16 Only one character per couplet need apply, since these are not combinations of characters. alternates are cited to ease identification. Several 204 ROBERT TRAUB caudad than does ventral lobe; this median lobe smaller than ventral lobe. (Yunnan, Indo-China, Thailand). . rectodigitus (Li & Wang, 1958)}? — Distal portion of F.SC. narrow, about thrice as long as broad. Notch at bifid apex of F.SC. with prongs diverging at tip or, at most, parallel. Seventh sternum of female with median lobe subequal in size and extent to subventral one (Indian subcontinent and Ceylon) : : aporus Jordan & Rothschild, 1922 5 PHY. (figs 170, 174) relatively broad, Pinout 2-3 times as long as broad at maximum level. Apicocaudal flap (AC.F.9) of D.A.g relatively short and broad, viz., length (from level of caudal margin of transverse sclerotization, T.S.) only about 1-3 times breadth at middle. B.C. (fig. 177) relatively long, viz., vertical diameter of involute portion equal to distance from bottom of loop to vagina (VAG.). The portion of wall of B.C. which is anterior to tanned lining of lumen, broader than wall of opposite side of duct. Lateral metanotal area with internal dimensions taller than long at subdorsal level. (Mindanao) cognatus bamus subsp. n. (p. 287) — PHY. (fig. 173) narrow; about 5 times as long as broad at maximum level. AC.F.9 (figs 168, 169) comparatively longer and narrower, at least 1.6 times as long as broad at middle. B.C. (figs 178, 179) with vertical diameter of involute portion exceeding distance from bottom of loop to vagina (VAG.). Portion of wall of B.C. which is anterior to tanned lining of lumen, narrower than opposite wall. Lateral metanotal area generally longer than high (internal measurements). He and Luzon) . 6 F-SC. (fig. 173) proportionately short and broad: total length only About 1-6 times as long as its basal portion (to level of dorsal nubbin) and 1-9 times as long as broad at level of dorsal spur. PHY. symmetrically sigmoid. Apical lobe (fig. 169, A.LB.9) of D.A.9 relatively tall, only twice as long as high. B.C. (fig. 178) with involute portion quite circular in outline. (Luzon) cognatus spiramus Jordan, 1926 (p. 287) — FSC. relatively long and narrow; about 1-9 times length of its basal portion and about 2-5 times as long as broad at apex of basal section. PHY. quite straight, definitely not sigmoid. A.LB.9 (fig. 168) proportionately short, about 2-4 times as long as high. B.C. (fig. 179) with involute portion oblate, rather flattened dorsally. (Java) . 5 2 0 : cognatus cognatus J. & R., 1922 (p. 286) ACKNOWLEDGEMENTS This study was supported by Grant No. Ar-04242 of the National Institutes of Health, Bethesda, Maryland, U.S.A., with the Department of Microbiology, Univer- sity of Maryland School of Medicine, Baltimore, Maryland. The large number of specimens that were available for this study was only obtained through the generous co-operation and assistance of friends and colleagues. Much of the material resulted from joint projects undertaken with the U.S. Army Medical Research Unit (USAMRU), by the Division of Virus Research and Medical Zoology of the Institute for Medical Research (DMZ-IMR), or its predecessors; by the Univer- sity of Malaya (UM) or by the International Center for Medical Research and Train- ing, of the George Williams Hooper Foundation of the University of California Medical Center at San Francisco (HF). Personnel who served with the writer in 17S. rvectodigitus was described from specimens from Yunnan. I have seen only specimens from Thailand, Vietnam and Laos, which closely resemble the figures and description of Li & Wang, but which I had regarded as subspecies of S. aporus. Further study, including specimens from Yunnan, are required to clarify the status of S. vectodigitus, but I am inclined to consider it a subspecies of S. aporus. Also, the key-characters cited above for the separation of the females may prove unreliable, but females of related forms of Stivalius s. lat. are often difficult or impossible to identify in the absence of associated males. NEW TAXA OF FLEAS 295 North Borneo as members of USAMRU in 1951 or 1953 included Capt. C. L. Wisse- man, Jr., M.C.,18 David H. Johnson, Ph.D. (Smithsonian Institution), Charles Wharton, Capt. H. D. Newson, M.S.C., and Capt. B. C. Walton, M.S.C. Scientists from DMZ-IMR on those or other trips to Borneo (i.e., 1953, at Mt. Trus Madi) included Dr. J. R. Audy and Dr. J. L. Harrison, who also were responsible for the DMZ-IMR collections from 1947 onwards, for 10 or more years. B. L. Elisberg, M.D., H. E. McClure, Ph.D., Lt. V. J. Tipton, M.S.C., participated in the USAMRU studies in Malaya at various times. Integral members of the field-teams in Malaya, N. Borneo and Sarawak (1958) were Ben Ensoll and Phang Ong Wah (USAMRU) and Lim Boo Liat and M. Nadchatram (DMZ-IMR). Yoon Yuen Fat and Ng Cheong Kee served on some of the USAMRU trips in Malaya and the latter worked on Tioman Island with UM. The USAMRU teams were all field-units of the Walter Reed Army Medical Center, Washington, D.C. The accomplishments of Lord Medway’s expedition to Gunong Benom are amply attested throughout this volume, but Lord Medway also helped us obtain valuable specimens from other parts of Malaya and North Borneo as well. Dr. J. A. Bullock (UM) was most helpful in providing fleas from northern Malaya and Tioman Island. The Bernice P. Bishop Muesum, via J. L. Gressitt, Ph.D., sent for study, specimens from Java collected by J. M. Stusak. R. E. Elbel, who served with Dr. Audy in N. Borneo in 1953, also collected the bulk of the Thai specimens, while with the U.S. Operational Mission to Thailand. Additional Thai material was provided by Lt.Col. J. E. Scanlon, M.S.C., and other members of the U.S. Component of the SEATO Medical Research Unit in Bangkok. Dr. Boonsong Lekagul was extremely helpful during all the studies in Thailand, as was Kitti Thonglongya. Special thanks are due to Dr. T. C. Maa of the staff of the Bishop Museum for the great efforts he expended, under difficult conditions, in collecting fleas in Indonesia, Sarawak, Malaya, New Guinea and elsewhere, while simultaneously making superb collections of other kinds of insects. F. G. A. M. Smit of the British Museum (Natural History), rendered great assistance by lending me paratypes and unidentified specimens from the Rothschild Collection of Fleas, thereby providing us with a new species for description, while another new species was received from the Director of the Rijksmuseum van Natuur- lijke Historie, Leiden, The Netherlands. Mr. Smit was also very helpful in deter- mining some of the precise localities in Malaya, Java and Sumatra where specimens in the N. C. Rothschild Collection had been collected. The illustrations in this series of articles were prepared by members of our Depart- ment at Baltimore. Of the photographs 93 are the work of J. Navarro, while Mrs. Suzanne L. Shipley prepared 33 photographs or drawings, including the difficult figures of the dorsal or ventral aspects of the aedeagus. Thirty-three of the illustra- tions were drawn by Mrs. Avonne Green. The bulk of the figures of L. vomerus were prepared by Miss Ermona McGoodwin, but those of the male genitalia are composites made by her and R. Traub. The dissections were prepared by the latter and mounted by Mrs. Phunthong Malikul. Miss Helle Starcke and T. M. Evans rendered considerable editorial assistance. 18 The military rank cited in each instance is that borne at the time the work was done. 296 ROBERT TRAUB The mammals were identified by J. L. Harrison and Lim Boo Liat (for Malayan and Bornean mammals), Lord Medway (Malaya), D. H. Johnson (N. Borneo and Thailand), Kitti Thonglongya (Thailand for SEATO) and R. E. Elbel (some Thai hosts). With respect to names of hosts, Lord Medway’s book on Bornean mammals was followed (1963), as well as Harrison’s works on the Malayan (1966) and Bornean (1964) mammals. The opus by Ellerman and Morrison-Scott (1951) was utilized for palaearctic mammals, as were the volumes by Ellerman (1940, 1941, and 1949) in some other instances. The scheme of classification followed is essentially that of Simpson (1945) and that of Anderson and Jones (1967), but with modification for the classification of higher taxa of New World mammals as per Hershkovitz (1966, 1969). In instances where there was a conflict in generic names, Walker et al. (1964) was followed, because he generally was the latest reviser. A complete list of relevant host names is appended. My thanks are extended to all of these associates for their cheerful and cheering help. REFERENCES ANDERSON, S. & Knox Jones, J., JR., ed. 1967. Recent mammals of the world. A synopsis of families. 453 pp., New York. Barnes, A. M. & Rapovsky, F. J. 1969. A new Tunga (Siphonaptera) from the Nearctic Region with description of all stages. J. med. Ent. 6 : 19-36, illus., refs. Costa Lima, A. pa & HatHaway,C. R. 1946. Pulgas. 522 pp., Rio de Janeiro. ELLERMAN, J. R. 1940. The families and geneva of living rodents. Vol. 1. 689 pp., London. 1941. The families and genera of living rodents. Vol. II. 690 pp., London. 1949. The families and genera of living vodents. Vol. III. 210 pp., London. ELLERMAN, J. R. & Morrison-Scott, T. C. S. 1951. Checklist of Palaearctic and Indian mammals. 1758 to 1946. 810 pp., London. Goncuarov, A. I. 1964a. On the study of functions of modified segments and some parts of aedeagus important in flea diagnosis. Tyudy armyansk. Protivoch. Stn. 3 : 437-457- 1964b. On coupling (linkage) of male sternite IX of flea with aedeagus. Trudy armyansk. Protivoch. Stn. 3 : 459-471. Harrison, J. L. 1964. An introduction to the mammals of Sabah. x + 244 pp., Singapore. —— 1966. An introduction to mammals of Singapore and Malaya. Malay. Nat. Soc. (Singapore Branch), 340 pp. HeERsHKoviTz, P. 1966. Mice, land bridges and Latin American faunal interchange. pp. 725-751. In WENZEL, R. L. & Tipton, V. J., ed., Ectoparasites of Panama. 861 pp., Chicago. 1969. The evolution of mammals on southern continents. VI. The recent mammals of the Neotropical Region: A zoogeographic and ecological review. Quart. Rev. Biol. 44 : 1-70. Hotranp,G.P. 1955. Primary and secondary sexual characteristics of some Ceratophyllinae, with notes on the mechanism of copulation (Siphonaptera). Tyvans. voy. ent. Soc. Lond. 107 : 233-248, illus., refs. 1969. Contribution towards a monograph of the fleas of New Guinea. Mem. ent. Soc. Canad. (61), 77 pp., illus., refs. : Hopkins, G. H. E. & Rotuscuitp,M. 1966. An illustrated catalogue of the Rothschild collection of fleas (Siphonaptera) in the British Museum. Vol. 4, 549 pp., London. Jorpan, K. 1926. New Siphonaptera. Novit. zool. 33 : 385-394, illus. —— 1933. Fleas collected by Dr Max Bartels in Java. WNovit. zool. 38 : 352-357, illus. Jorpan, K. & Roruscuitp, N.C. 1922a. The Siphonaptera collected by Mr E. Jacobson on Sumatra. Ectoparasites 1 : 223-230, illus. NEW TAXA OF FLEAS 297 Jorpan, K. & RoruscuiLp, N.C. 1922b. On Pygiopsylia and the allied genera of Siphonaptera. Ectoparasites 1 : 231-265, illus. Li, K.-C. & Wane, D.-C. 1958. The fleas of the genus Stivalius J. et R. 1922 (Siphonaptera) from China. Acta ent. Sinica 8 (1) : 67-75, illus., refs. Mepway, Lorp. 1963. Mammals of Borneo. J. Malay. Brch R. Asiat. Soc. 36 (3) : 1-193. Oupemans, A. C. 1909. Pygiopsylla robinsoni (Rothschild) g. Notes (XI) Leyden Mus. 31 : 195-200, illus. Peus, F. 1956. Siphonaptera. pp. 122-131. In Tuxen, S. L., ed., Taxonomist’s glossary of genitalia in insects. 284 pp., illus., Copenhagen. PoLLiTzeR, R. 1954. Plague. WHO Monog. Ser. No. 22, 698 pp., Geneve. Poriitzer, R. & MEYER, K.F. 1961. The ecology of plague, pp. 433-501, 582-590. In May, J., ed., Studies in disease ecology (Vol. 2 in series Studies in medical geography.) 613 pp., New York. RotuHscHILp, N. C. 1905. Some new Siphonaptera. Novtt. zool. 12 : 479-491, illus. 1906. Some new exotic fleas. Ent. mon. Mag. (2) 17 : 221-224, illus. — 1908. New Siphonaptera. Proc. zool. Soc. Lond. 50 : 617-629, illus. 1919. Siphonaptera collected in Korinchi, West Sumatra by Messrs H. C. Robinson and C. Boden Kloss. J. fed. Malay St. Mus. 8(3) : 1-6, illus. Scuwarz, E. & ScHwarz,H. K. 1965 (1967). A monograph of the Rattus vattus group. An. Esc. nac. Cienc. biol., Mex. 14 : 80-178, illus., refs. Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bull. Am. Mus. nat. Hist. 85, 350 pp. Smit, F.G. A.M. 1958. The African species of Stivalius, a genus of Siphonaptera. Bull. Br. Mus. (Nat. Hist.) Ent. 7 : 41-76, illus., refs. SnopGrass, R. E. 1946. The skeletal anatomy of fleas (Siphonaptera). Smithson misc. Collns. 104, 89 pp., illus. Suter, P.R. 1964. Biologie von Echidnophaga gallinacea (Westw.) und Vergleich mit andern Verhaltenstypen bei Fléhen. Acta trop. 21 : 193-238, illus., refs. Traus, R. 1950a. Siphonaptera from Central America and Mexico. A morphological study of the aedeagus with descriptions of new generaand species. Fieldiana Zool. Mem.1 : 1-127, illus., refs. 1950b. Notes on Indo-Malayan fleas, with descriptions of new species (Siphonaptera). Proc. ent. Soc. Wash. 52 : 169-143, illus., refs. —— 1952. Fleas collected by the Chicago Natural History Museum Expedition to the Philippines, 1946-1947. Proc. biol. Soc. Wash. 64 : 1-21, illus., refs. —— 1966. Some examples of convergent evolution in Siphonaptera. (Abstract of paper presented at meeting of Society, Dec. 1966: Proc. ry. ent. Soc. Lond. (C) 31(7) : 37-38.) Reported in Ibid. 31 : 46-47 (1966-1967) with errata in Ibid. 31 : 79 (1966-1967). 1969. Muesebeckella, a new genus of flea from New Guinea (Siphonaptera, Pygiopsyllidae). Proc. ent. Soc. Wash. 71 : 374-396, illus., refs. Watker, E. P., Warnicu, F., et al., 1968. Mammals of the world. Vols. I-II : 1-646, 647- 1500, Baltimore. Colonel Ropert Travs, U.S. Army (Ret.), Ph.D., Professor, Department of Microbiology, UNIVERSITY OF MARYLAND SCHOOL oF MEDICINE, BaLtimore, MARYLAND 21201, U.S.A. 298 ROBERT TRAUB LIST OF HOST-NAMES!9 Apodemus speciosus Temminck, 1845 Apomys Mearns, 1905 Callosciurus Gray, 1867 Callosciurus caniceps (Gray, 1842) Callosciurus evythvaeus (Pallas, 1779) Callosciurus finlaysoni (Horsfield, 1824) Callosciurus n. nigrovittatus (Horsfield, 1824) Callosciurus notatus (Boddaert, 1785) Callosciurus notatus miniatus (Miller, 1900) Callosciurus prevosti (Desmarest, 1822) Dremomys Heude, 1898 Dremomys vufigenis (Blanford, 1878) Epimys vattus (Linnaeus, 1758) Hylopetes Thomas, 1908 Iomys horsfieldi (Waterhouse, 1837) Laviscus Thomas & Wroughton, 1909 Lariscus insignis (Cuvier, 1821) Menetes berdmorei (Blyth, 1849) Millardia Thomas, 1911 Petaurista Link, 1795 Petaurista elegans (Miller, 1839) Ptilocercus lowi (Gray, 1848) Rattus Fischer, 1803 Rattus (Lenothrix) canus (Miller, 1903) Rattus (Lenothrix) vajah (Thomas, 1894) Rattus (Lenothrix) surifer (Miller, 1900) Rattus (Leopoldamys) sabanus (Thomas, 1887) Rattus (Maxomys) cremoriventey (Miller, 1900) ?Rattus (Maxomys) fulvescens temminckt Kloss, 1921 (for ‘Rattus bukit temmincki’’) Rattus (Rattus) Fischer, 1803 Rattus (Rattus) concolor (Blyth, 1859) Rattus (Rattus) querceti Hollister, 1911 Rattus (Rattus) vattus (Linnaeus, 1758) Rattus (Rattus) vattus mindanensis (Mearns, 1905) Rattus (Rattus) vattus negyinus Thomas, 1898 Rattus (Rattus) tiomanicus (Miller, 1900) Rattus (Rattus) tiomanicus jalorensis (Bonhote, 1903) Rattus (Stenomys) boweysi (Anderson, 1879) Rattus (Stenomys) muelleri (Jentink, 1879) Ratufa Gray, 1867 Rhinosciurus Blyth, 1855 Rhinosciurus laticaudatus (Miller, 1844) Sundasciurus Moore, 1958 Sundasciurus brookei (Thomas, 1892) Sundasciurus hippurus (Geoffroy, 1832) Sundasciurus lowi (Thomas, 1892) Sundasciurus tenuis (Horsfield, 1824) Tamiops mcclellandi (Horsfield, 1839) Tupaia Raffles, 1821 Tupaia glis (Diard, 1820) Tupaia javanica Horsfield, 1822 Tupaia javanica occidentalis Robinson & Kloss, 1918 Tupaia minor Giinther, 1876 Tupaia tana Raffles, 1821 LIST OF LANDMARKS AND POINTS OF REFERENCE (Figs 1-3, 28 and 29) Dorsal sinus of T.AP.9 and base of anterior margin of conical process (C.P.). Point of maximum convexity on anterior margin of movable finger (F.). Level for measuring length of dilated portion of apex of F.; anterior margin. Fic. 1 a Anteroventral angle of tergal apodeme of 3 segment 9 (T.AP.9). b Ventral sinus of T.AP.9. c Anterodorsal angle of T.AP.g. d @ Apex of C.P. ff Level of middle of C.P.; anterior margin. g Level of middle of C.P.; posterior margin. h Middle of base of C.P. i j Level of narrowest subapical portion of F.; anterior margin. k Level of narrowest subapical portion of F.; posterior margin. 1 m Apicodorsal angle of F. n Midpoint of apical margin of F.; for measuring height of F. o Level of base of stiva (STV.); a projection of posterior margin of F. at narrowest level (k). Ba 2 a ee EE Ee eee 19 Not all of these names are necessarily regarded by mammalogists as valid today. NEW TAXA OF FLEAS 299 Apex of stiva. Point of maximum convexity on posterior margin of F. Level of most apical portion of immovable process of clasper (P.). Level of r, projected to longitudinal axis of F. Level of base of bulge at apex of P. Point of maximum convexity on upper or dorsal bulge on posterior margin of manu- brium (MB.). Point of maximum concavity on sinus on posterior margin of MB. Point of maximum convexity on lower or apical bulge of MB. Base of apical cap of MB. (posterior margin). Tip of apical cap of MB. Base of apical cap of MB. (anterior margin). Apex of anterior (ventral) margin of proximal arm of sternum 9 (P.A.9). Apex of posterior (dorsal) margin of P.A.9. Subapical bulge (convexity) on anterior margin of P.A.9. Posterior or ventral bulge on anterior margin of P.A.9. Point on posterior margin of P.A.g that is opposite cc (paralleling chord on arc aa—bb). Point on posterior margin of P.A.g that is opposite dd. Lowest point on trough or ‘‘U”’ on dorsal margin at base of P.A. 9. Point on ventral margin of P.A.g opposite gg. Base of distal arm of sternum 9 (D.A.9); anterior margin. Anterior (dorsal) end of transverse sclerotization (T.S.) of D.A.g and level of base of ventral (caudal) margin of mid-microspiculate area (M.MSP.). Base of sclerotized part of apical half of D.A.g; distad of mid-microspiculate area (M.MSP.) and level of proximal-most of caudomedial group of spiniform bristles (CM.G.B.) (see also fig. 3). Proximal marginal point of base of subapical lobe (SUB.L.), at level of proximal-most bristle of SUB.L. Apex of SUB.L. (see also fig. 3). Midpoint of imaginary chord connecting edge of subapical notch (NCH.). Apicoventral angle of apical lobe AP.L. of D.A.9. Midpoint on margin of AP.L. Apicodorsal angle (or point in line with 00) on AP.L. (see also fig. 3). Lowest point in subapical notch (NCH.) on D.A.9. Level of distalmost member of subapical group of spiniforms (SG.SPN.). Level of proximal-most of CM.G.B. Posterior (ventral) margin of T.S. Base of D.A.9 (posterior margin). Anteroventral angle of upper margin of fulcral sclerite (F.S.). Caudodorsal angle of upper margin of fulcral sclerite (F.C.) Dorsal extension of F.S. curving to ventrocaudal angle of F. Anteroventral angle of lower or ventrocaudal margin of F.S. Upper angle of ventrocaudal margin of F.S., opposite zz. Dorsal point of transverse sclerotization (T.S.) of D.A.g in M. loncha and allies (but not ventral margin of M.MSP.). Ventral or proximal base of spur-like subapical lobe (SUB.L.) of D.A.9 in M. loncha and allies. Thickening at base of SUB.L. Dorsal or apical base of SUB.L. Semimembranous margin of D.A.g distad of e3. Level of base of spinform opposite d3 at base of SUB.L. ROBERT TRAUB 300 Fics 28, 29 h3 Dorsal margin of 8 S.—projection of upper portion (anterior fork) of SU.8. i3 Dorsalmost point of 8 S. j3 Dorsal margin—in line with middle of dorsal margin (03) of LUM. and dorsalmost median large bristle (k3). k3 Dorsalmost large median bristle (in line with middle of LUM.). 13 Point on anterior margin, near middle, in line with largest caudomarginal bristle (m3). m3 Longest and secondmost dorsal, caudomarginal bristle. n3 Anteriormost point on tanned dorsal margin of LUM. 03 Middle of LUM. as located on dorsal margin. p3 Ventrocephalad corner of 8 S. q3 + Anteriormost point on ventral margin of LUM. indicated by limit of dense group of rows of spiniforms. r3 +Point on ventral margin directly below highest point (i3). s3 Caudal apex of LUM. t3 Ventrocaudal corner of 8 S. Fic. 3 u3 ~—«~ Projection of axis of T.S. to ventral margin of D.A.g. Fic. 2 v3 __ Level of dorsal margin of head at falx. w3 Falx. x3 Point along antennal fossa in line with bottom of median sensory crater (M.S.C.). y3 Point along frons in line with x3. z3 Point along frons in line with base of uppermost bristle of row I. a4 Point along antennal fossa in line with z3. b4 Point along frons in line with base of uppermost bristle of row II. c4 Point along antennal fossa in line with labral bristles (d4) and base of eye-bristle (E.B.). dq Level of labral bristles. e4 Point along antennal fossa in line with labral bristles and E.B. f4 Point on ventral margin of gena in line with w3 (falx) and height of convexity of first 7 ventral arc of gena. } g4 Point of height of convexity of second ventral arc of gena. | LIST OF ABBREVIATIONS A.B. Antepygidial bristles. ARC. Broad sinus or arc on D.A. 9. A.G. Apical group of dorsomarginal B. Bulga (head) of spermatheca. bristles on tibia. B.€. Bursa copulatrix. A.LB.g Short apical lobe on D.A.9. B.CR. Body of crochet. } AGS: Anal stylet. B.M.L. Bay of middle lamina. AC.F.9 Apicocaudal flap of D.A.9. B.MB. Bay of manubrium. AE.A Aedeagal apodeme. CP: Conical process on anterior por- AE.F Aedeagal fulcrum. tion of P. AE.P. Aedeagal pouch. Cis: Crescent sclerite of aedeagus. AE.P.-L Lateral walls of AE.P. CAV.SPIC. Caverna spiculosa. AE.P.-V. Thickened ventral wallsof AE.P. CEN.S. Central sclerite of aedeagus. ALPH Alpha-portion of Ford’s sclerite. CM.G.B. Caudomarginal group of stout AN.GP. Anterior group of marginal bristles on D.A.9. bristles on F. COX.1 Procoxa. AP.L Apical lobe of D.A.9. COX.3 Metacoxa. AP.R Third apodemal rod of endo- CR.P. Crochet process. phallus. D.A.L. Dorsal anal lobe of proctiger. NEW TAXA OF FLEAS Distal arm of male sternum 9. Duct of bursa copulatrix. Distal fringe of long bristles on caudal margin of F. Dorsal sensillary crater of pre- antennal region. Duct of spermatheca. Dorsal virga. Duplicatura vaginalis. Deltoid flap of hood of aedeagus. Dilated portion of duct of spermatheca. Groups of dorsolateral bristles arising from marginal notches on tibia. Large bristle in front of or near eye. Eye-row of bristles. Spur-like extension of ventral apical margin of D.A.9. Movable finger or process of clasper. Fulcral sclerite at base of attach- ment of F. Ford’s sclerite. Fistula. Profemur. Freak—grossly abnormal bristle. Fulcral lateral lobes of aedeagus. Fulcral medial lobe of aedeagus. Girdle of aedeagal pouch. Genal row of bristles. Glandula vaginalis. Groove-like structure of Ford’s sclerite. Hilla (tail) of spermatheca. Hood of aedeagus. Homolate margin proximad of apex of D.A.9. Lower arm of securifer. Lateral lobe of aedeagus. Lower lobe of tergum 7, below plate of antepygidial bristles. Lateral laminae of aedeagal apodeme. Lateral metanotal area of meta- thorax. Lower modified bristle(s) of 7 T. Second or ventral L.M.B. Labial palpus. Lateral shafts of capsule of aedeagus. Lateral wings of ventral arma- ture of S.I.T. LUM. M.D.L. M.LAM. M.MSP. M.P. M.R.8 M.S.C. MB. MPM. MPS. 301 Lumacaudate process. Median dorsal lobe of aedeagus. Middle lamina of aedeagal apo- deme. Mid-microspiculate area of D.A.9. Maxillary palpus. Mesal tanned genitalic ridge of 9 8T. Median sensillary crater of pre- antennal region. Manubrium. Mesepimere. Mesepisternum. Mesonotum. Patch of microspicules near apex of D.A.g. Metepimere. Metanotum. Metepisternum. Maxillary lobe. Subapical notch on anterior margin of D.A.9. Oviduct. Immovable process of clasper. Proximal arm of g$ sternum 9. Perula—dilated portion of bursa copulatrix. Penis rods. Extension of aedeagal pouch wall. Apical papilla on bulga. Paragenital morion. Phylax. Pivotal chord linking phylax and Ford’s sclerite. Pivotal ridge of phylax. Peak of bulga. Pleural arch of metathorax. Pseudosetae. Quasi-crochet. Group of sensilla near antero- distal angle of F. Subapical group of short spini- forms on caudal margin of D.A.9. Sclerotized inner tube of aedea- gus. Apical fold or seam of LUM. Subanal sclerite of male. Large sac associated with perula. Satellite sclerite of aedeagus. Securifer—ventrocaudal or api- cal member of Ford’s sclerite. Sensilium. ROBERT TRAUB 302 SP. Spermatheca. SPIC.L Ventral lamella or semimem- branous spiculose lobe arising from base of PHY. SQ. Squamulum. STV. Stiva—caudodistal expansion of F., resembling a plow-handle in shape. SU.8. Suture delimiting more heavily sclerotized mesal area of § 8S. SUB.L Subapical lobe on anterior mar- gin of D.A.9. SUP.FL. Supramedial flap of D.A.g. SUP.G. Supramedial group of thin bristles on D.A.g. SY: “Supernumerary’’ bristles be- tween rows on head. T.AP.9 Tergal apodeme of segment 9. ATTICS Tendons of lateral shafts of aedeagus. es Transverse submedian sclero- tization of D.A.9. THK. Sclerotized thickening associa- ted with SUP.FL. THM. Thumb-like apex of alpha-por- tion of F.SC. Fig. 1. Clasper and sternum 9 2. Preantennal region (3) 3. Distal arm of sternum 9 4. Head and prothorax (3) 5. Lateral metanotal area (3) 6. Head (2) 7. Apical segments of protarsus 8. Segment 5 of f metatarsus 9. Meso- and metathorax to. Aedeagus 11. Modified abdominal segments (3) 12. Lumacaudate process 13. Dorsal and ventral anal lobes (¢) 14. Processes of clasper 15. Spermatheca 16. Anal lobes and stylet (9) 17. Processes of clasper 18. Clasper and sternum 9 19. Metatibia (3) 20. Distal arm of sternum 9 21. Distal arm of sternum 9 22. Apical region of aedeagus (lateral aspect) 23. Apical region of aedeagus 24. Apex of aedeagus (dorsal aspect) U.A. U7 Upper arm of securifer. Upper lobe of tergum 7, above plate of antepygidial bristles. Upper modified bristle of 7 T. Vesicle of aedeagus. Ventral anal lobe of proctiger. Ventral sensillary crater of pre- antennal region. Ventral virga. Vagina. Vinculum 1—4. Y-sclerite of aedeagus. First tergum. Second sternum. Second tergum. Sixth sternum. Sixth tergum. Seventh sternum. Spiracle (fossa) of seventh seg- ment. Seventh tergum. Eighth sternum. Spiracle (fossa) of eighth seg- ment. Eighth tergum. Ninth sternum. LIST OF ILLUSTRATIONS Medwayella rv. yvobinsoni (Roths., 1905) loncha (Jordan, 1926) dvyadosa sp. nov. ” vy. vobinsont dvyadosa sp. nov. ” n y. yobinsont dvyadosa sp. nov. y. vobinsont dryadosa sp. nov. —? NEW TAXA OF FLEAS Apex of aedeagus (ventral aspect) Modified abdominal segments () Spermatheca, genitalia and anal segments Sternum 8 (3) Sternum 8 (3) Segment 7 (9) Genitalia (9) Genitalia (9) Distal arm of sternum 9 Processes of clasper Processes of clasper Distal arm of sternum 9 Modified abdominal segments (9) Spermatheca, genitalia and anal segments Lumacaudate process Genitalia (9) Genitalia (9) Apex of aedeagus Sclerotized inner tube Apex of aedeagus Apex of aedeagus Apical region of aedeagus Clasper and sternum 9 Transverse suture, D.A.g Sternum 7 (9) Lateral metanotal area (3) Anal segments (9) Spermatheca Distal arm of sternum 9 Distal arm of sternum 9 Distal arm of sternum 9 Apex of aedeagus Apex of aedeagus Apex of aedeagus Genitalia and anal lobes (?) Spermatheca and sternum 7 Genitalia (9) Spermatheca, genitalia and sternum 7 Proximal arm of sternum 9 Spermatheca, genitalia and sternum 7 Processes of clasper Processes of clasper Distal arm of sternum 9 Distal arm of sternum 9 Sternum 8 (3) Lumacaudate process Apical region of aedeagus Tergum 8 (2) Anal lobes and stylet (9) Spermatheca Sternum 7 (2) Sternum 7 (2) a7 Spermatheca, genitalia and anal segments Spermatheca (variation in aspect) ” ” ” y. vobinsont dryadosa sp. nov. ” » vy. vobinsont aycuata sp. nov. ” angustata sp. nov. javana (Jordan, 1933) angustata sp. nov. dryadosa sp. nov. y. vobinsont angustata sp. nov. vy. vobinsont ” . tiomanica subsp. nov. . bogova subsp. nov. . tiomanica subsp. nov. . bogova subsp. nov. . vobinsont St Ye ” y. bogova subsp. nov. y, tiomanica subsp. nov. phangi tana subsp. nov. phangi phangi sp. nov. p. tana subsp. nov. p. phangi sp. nov. p. tana subsp. nov. p. phangi sp. nov. limi sp. nov. . pevegrinata subsp. nov. . pevegryinata subsp. nov. . peregvinata subsp. nov. 393 ROBERT TRAUB Spermatheca and genitalia Processes of clasper Distal arm of sternum 9 Apex of aedeagus Lumacaudate process Spermatheca and genitalia Sterna 8 and 9 (9) Head and prothorax (3) Movable finger Spermatheca Sensilium and anal segments () Apical region of aedeagus Distal arm of sternum 9 Processes of clasper Apical region of aedeagus Apex of aedeagus Apex of aedeagus Sclerotized inner tube Apical region of aedeagus Apical region of aedeagus Sternum 7 (9) Lateral metanotal area (3) Sternum 8 (3) Spermatheca and genitalia Spermatheca, genitalia and anal segments Head and prothorax (3) Lateral metanotal area (3) Lateral metanotal area (3) Lumacaudate process Processes of clasper Processes of clasper Mesal aspect of F. Distal arm of sternum 9 Distal arm of sternum 9 Apex of aedeagus Apex of aedeagus Apex of aedeagus Lumacaudate process Spermatheca and genitalia Spermatheca and genitalia Processes of clasper Distal arm of sternum 9 Sternum 8 (3) Anal lobes and stylet (9) Spermatheca Sternum 7 () Genitalia (9) Processes of clasper Distal arm of sternum 9 Apex of aedeagus Sensilium and anal segments (2) Sternum 7 (?) Sternum 8 (3) Spermatheca » dryadosa sp. nov. thurmani sp. nov. javana thurmani sp. nov. arcuata sp. nov. thuymani sp. nov. ” ” ” calcavata sp. nov. batibacula sp. nov. calcarata sp. nov. batibacula sp. nov. calcavata sp. nov. batibacula sp. nov. calcavata sp. nov. veruta sp. Nov. batibacula sp. nov. veruta sp. nov. loncha veruta sp. Nov. loncha 133- 134. 135. 136. 137: 138. 139. 140. 141. 142. 143. 144. 145. 146. 147- 148. 149. 150. 151. 152. 153: 154. 155: 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. E72. 173. 174. 175. 176. E77 - 178. 179. NEW TAXA OF FLEAS 305 Processes of clasper Distal arm of sternum 9 Lumacaudate process Sternum 7 (2) Spermatheca, genitalia and anal segments Modified abdominal segments (3) Modified abdominal segments (9) Head and prothorax (¢) Head (9) Meso- and metathorax (3) Clasper and segment 9 Distal arm of sternum 9 Distal arm of sternum 9 Clasper and segment 9 Modified abdominal segments (3) Aedeagus Metatibia (3) Segment 5 of metatarsus (3) Metatibia (2) Apical region of aedeagus (lateral) Apical region of aedeagus (ventral) Apical region of aedeagus (lateral) Apical region of aedeagus (lateral) Modified abdominal segments (9) Anal segments and stylet (9) Spermatheca Abdominal segments 2 (9) Spermatheca and genitalia Spermatheca (variation) Bursa copulatrix Spermatheca and genitalia Head and prothorax (3) Processes of clasper Antepygidial region (9) Distal arm of sternum 9 Distal arm of sternum 9 Distal arm of sternum 9 Apical region of aedeagus (lateral) Apical region of aedeagus (ventral) Apical region of aedeagus Apical region of aedeagus Apical region of aedeagus Sternum 7 (2) Spermatheca, genitalia and anal segments Spermatheca and bursa copulatrix Bursa copulatrix Spermatheca and bursa copulatrix ” javana vhaeba (Jordan, 1926) Lentistivalius vomerus sp. nov. insollt (Traub, 1950) vomerus Sp. NOV. insolli aestivalis (Jameson & Sakaguti, 1954) vomerus sp. nov. Stivalius phoberus J. & R., 1922 cognatus bamus subsp. nov. ” ” c. cognatus J. & R., 1922 c. spivamus Jordan, 1926 c. bamus subsp. nov. ” Lentistivalius vomerus Sp. Nov. Stivalius c. spivamus c. bamus subsp. nov. c. spivamus c. bamus subsp. nov. ” c. spivamus c. cognatus » - { . 2 Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PEATE a Ww maa A GREEN SS. loncha Jordan, 1926 Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 2 FIGS. 4-8 M. DRYADOSA N.SP J AVARRO Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 3 FIGS. 9-10 M. DRYADOSA N.SP Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 SN. 8 SPC , = hg ~~ ee ——— D.A.L ——__ Wa V.A.L FIGS. II-13 M. DRYADOSA N. SP. PLATE 4 Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 5 4 FIAVARRO J FIGS. 14-16 f M. DRYADOSA N.SP. i FIG. 17 M.ROBINSONI ROBINSONI (ROTHSCHILD, 1905) Bull. By Mus. nat. Hist. (Zool.) 23, 9 4 FYRYRERO 1 PIAVARRO PLATE 6 FIGS. 18-20 M. DRYADOSA N.SP. su Ship FIG. 21 M. ROBINSON] ROBINSONI (ROTHSCHILD, 1905) Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 7 M. dryadosa n sp BML PIVR CAV SPIC. CENS DeEL-FL. DW. AE.P-V. YS M.robinsoni robinsoni (Rothschild, 1905) \ PIV.R. TLS, FULM.L. cENS LSC. SATS. cs OEL FL SE Rast ores teveney evn ies ¥S. PHY. SUT ae a DV. Vv. DELFL A.GREEN AE.P-L. AE.P-L ac, Mus. nat. Hist. (Zool.) 23, 9 PLATE 8 Bull. Br. ({oajua@n) WY A-d 30 joadso j0JjUan joadso josiop | bd ‘ds u psoppKap pjjahompaw DIidS AVO q-d 3Vv 17704 os AHd 74 °730 (modo) 11S vo Hd1V vn Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 9 FIGS. 26-27 M. DRYADOSA N.SP 4. TAVARRO 10 PLATE 9 \Wus. nat. Hist. (Zool.) Bull. Br. dS N VSOGVAYNO W O£€-62 SOI CXELAL Za (SO6I ‘QTIHOSHLOY) INOSNISOY INOSNI8OY W ‘ 8¢ Oils , y Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 11 M. dryadosa n.sp. A. GREEN Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 12 SUB “hk XY y N\ AON Bull. Br. Mus. nat. Hist. (Zool.) 23. 9 FIGS. 35-36 M. ANGUSTATA N.SP 4. VAYARRO PAs 13 4 PLATE (Zool.) 23, 9 Vus. nat. Hist Bull. By oN HAG dS N VLVLSNONV W 6¢€-2e'SOl4 Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 15 M. javana (Jordan, 1933) 41 Ue M.angustata n.sp. A.GREEN BEBscs Bull. By. Mus. nat. Hist. (Zool.) 23, 9 UA DY Fics 42-43 LOD M DRYADOSA NSP FIG. 44 t M. ROBINSON! ROBINSON! (ROTHSCHILD, 1905) FIG. 45 M. ANGUSTATA N.SP | NAVARRO Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PAA En a7, 4 WAVRRRO FIGS. 46-48 M. ROBINSONI ROBINSONI (ROTHSCHILD, 1905) Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 18 a FIGS. 49-52 M. ROBINSONI ROBINSONI (ROTHSCHILD, 1905) a a. NAvRRRO aaa s Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 109 FIG.53 M.R. TIOMANICA NSP FIG. 54 M. ROBINSONI BOGORA N.SSP. 8LShiple a ¥ y. Fig.55 7.» M.ROBINSONI PEREGRINATA N SSP Bull. By. Mus. nat. Hist (Zool.) 23, 9 FIG 56 *_ M.ROBINSONI TIOMANICA NSSP FIG 58 M ROBINSONI BOGORA N.SSP. FIG. 57 M. ROBINSONI PEREGRINATA N SSP Bull, Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 2 FIG.S9 M. ROBINSONI ROBINSONI (ROTHSCHILD, 1905) FIGS. 60-6] M. ROBINSON PEREGRINATA N.SSP Bull. By. Mus. nat. Hist. (Zool.) 23, 9 i nee 1 . nal v. VRVARRO FIGS. 62-63 M. ROBINSONI BOGORA N.SSP. PLATE 22 FIG. 64 M.R. TIOMANICA N.SSP | Bull. By. Mus. nat. Hist. (Zool.) 23, 9 f M.phangi phangi n.sp M.MSP FSS \ cal A GREEN PLATE 23 Bull. By. Mus. nat. Hist. (Zool.) 23, 9 (ok VV. V. AE.P-V. S.1.T. PHY. FUL.LL, AE.P-L. DEL.FL. ae TS val A GREEN Medwayella phangi phangi n. sp. Se Qc. PLATE 25 M phangi phangi n sp. M. phongi phangi n sp. M.phang! tana n ssp. “Bull. By. Mus. nat. Hist. (Zool.) 23, 9 Bull. By. Mus. nat. Hist. (Zool.)23, 9 variation (aspect) A. GREEN M. phang! pnang! n.sp. PASTE: 20 at Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE FIGS. 80-83 MEDWAYELLA LIMI N.SP Bull. By. Mus. nat. Hist. (Zool.) 23, 9 FIGS. 84-85 MEDWAYELLA LIMI NSP PLATE 9.4. Shipley 28 Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 FIGS. 86-89 M. THURMANI N.SP 4 RURRRO PLATE 29 Bull. By. Mus. nat. Hist (Zool.) 23, 9 PLATE 30 FIGS. 90-92 ‘ 3 tas M.THURMANI NSP ; m oh Mae Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 31 s P FIGS. 95-96 | = nae he

variation (aspect) S.phoberus Jordan & Rothschild, 1922 S. cognatus bamus n.ssp. 163 VAG. A. GREEN Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE S ai - 53 FIGS. 164-166 STIVALIUS COGNATUS BAMUS N.SSP. \ ¢y Soe Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 54 FIG. 167 a STIVALIUS COGNATUS BAMUS N.SSPR™ - . Z + FIG. 169 Se ae STIVALIUS COGNATUS SPIRAMUS "a= ; <, JORDAN, 1926 —— ‘ CL niav geo Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 | (49adso jo2jUaA) (\oadso |04240)) WITW oL| Dds AVO wv AHd os4 Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 56 FIG. 172 LENTISTIVALIUS VOMERUS N.SP. FIG. 173 STIVALIUS COGNATUS SPIRAMUS JORDAN, 1926 * ove FIG. 174 STIVALIUS COGNATUS@ BAMUS NSSP <9 ee 174 7 AR. y ~ " an : : = ~S NAVARRO Bull. Br. Mus. nat. Hist. (Zool.) 23, 9 PLATE 57 FIG. 175 ~—STIVALIUS COGNATUS SPIRAMUS JORDAN, 1926 JFIG. 176 =STIVALIUS COGNATUS BAMUS N.SSP 176 41 WAVARRS Bull. By. Mus. nat. Hist. (Zool.) 23, 9 PLATE 58 D.SP— FIG. 177 STIVALIUS COGNATUS BAMUS N.SSP FIG. 178 STIVALIUS COGNATUS SPIRAMUS JORDAN, 1926 * FIG) 79 \9STIVALIUS COGNATUS COGNATUS 4 3 PResey,. JORDAN & ROTHSCHLD, 1922 SS ENTEp <0 Sy ;;pRe~ Yn HiSST. ‘he A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier an¢ Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. Taytor, J. D., KENNEpy, W. J.& Hatt, A. The Shell Structure and Mineralog’ of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plat 77 ee 1969. £4.50. 1962-1964. (I n press.) ZSE ED NAT. tis r ?\ ae eee yi Hy PRege J Sut Printed in England by Staples Printers Limited at their Kettering, Northants, establishment Cl ath = THE GUNONG BENOM EXPEDITION a 7 OTES ON ZOOGEOGRAPHY, CONVERGENT OLUTION AND TAXONOMY OF FLEAS HONAPTERA), BASED ON COLLECTIONS M GUNONG BENOM AND ELSEWHERE | IN SOUTH-EAST ASIA. IL CONVERGENT EVOLUTION R. TRAUB | BULLETIN OF : ISH MUSEUM (NATURAL HISTORY) va ‘ Vol. 23 No. 10 LONDON: 1972 THE GUNONG BENOM EXPEDITION 1967 2. NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON ~ COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA. Il. CONVERGENT EVOLUTION BY ROBERT TRAUB University of Maryland Pp. 307-387; 20 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 10 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 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 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper 1s Vol. 23, No. Xo of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 28 December, 1972 Price £4.45 | | | | | / | | Page 354, third line from end of second paragraph: for ‘areas) or are mane-fleas. The archurieopsyllines include . . . ’ The archaeopsyllines include. . . ’, ILUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON yead ‘areas) or are murine-fleas. ERRATUM 967 COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA. II. CONVERGENT EVOLUTION By ROBERT TRAUB CONTENTS INTRODUCTION . BAacKGROUND DATA AND OBSERVATIONS . A. THE PRONOTAL COMB B. THE PLEURAL ARCH Discussion A. THE FUNCTION OF THE PRONOTAL COMB 1. General Comments 2. Special Kinds of Combs on ES. 5 3. The Pronotal Comb of Fleas oe Non-Arboreal Nests B. NEST-FLEAS IN GENERAL . 3 1. Introduction 2. Hystrichopsyllid Nest- Fleas 3. Pygiopsyllid Nest-Fleas 4. Cevatophyllid Nest-Fleas . 5. Leptopsyllid Nest-Fleas 6. Pulicid Nest-Fleas . : 7. Chimaeropsyllid Nest-Fleas 8. Rhopalopsyllid Nest-Fleas g. Differences in Degree of the Nest-Flea Habitus C. THE PLEURAL ARCH OF THE METATHORAX 1. Review of Taxa Lacking a Pleural Arch 2. The Adaptive Value of the Loss of the Pleural Arch 3. Hyperdevelopment of Combs in Bird-Fleas, Bat-Fleas, ete., Lacking the Pleural Arch 0 4. Hyperdevelopment of the Byenotal Gore apEbine Squirrel Fleas, ete. 5. Convergence in Vestiture in Unrelated Fleas 6. Morphological Changes in the Thorax Associated with peed Jumping Ability in Fleas D. THE DEGREE OF DEVELOPMENT OF THE EYE IN VARIOUS FLEAS I. Comparative Size of the Eye 2. Vision in Fleas 3. The Types of Thorax in Eieas Any Reduced. Eyes E. THE EVOLUTION OF THE CHARACTERISTIC PRONOTAL COMB ASCRIBED TO MOST NEST-FLEAS F. DiIstTRIBUTION OF THE NEST-FLEA PHENOMENON IN SIPHONAPTERA . G. THE ASSOCIATION BETWEEN THE MODE OF EGG-LAYING AND THE NEST-FLEA HABIT . . . . 4 Wo wee wp peer we er rene reeeen Uwe Wwe Urey Cewure veady. Volumes will ey 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. 23, No. 10 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 28 December, 1972 Price £4.45 THE GUNONG BENOM EXPEDITION 1967 12, NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA. II. CONVERGENT EVOLUTION By ROBERT TRAUB CONTENTS INTRODUCTION . BacKGROUND 5 DATA AND OBSERVATIONS . A. THE PRONOTAL COMB B. THE PLEURAL ARCH Discussion A. THE FUNCTION OF THE PRONOTAL COMB B. On I. General Comments 2. Special Kinds of Combs on niarpelsire: 3. The Pronotal Comb of Fleas aig Non-Arboreal Nests NEST-FLEAS IN GENERAL . c : 1. Introduction Hystrichopsyllid Mest leas Pygiopsyllid Nest-Fleas Ceratophyllid Nest-Fleas . Leptopsyllid Nest-Fleas Pulicid Nest-Fleas . Chimaeropsyllid Nest-Fleas Rhopalopsyllid Nest-Fleas Differences in Degree of the Nest-Flea avis THE PLEURAL ARCH OF THE METATHORAX Review of Taxa Lacking a Pleuval Arch . 2. The Adaptive Value of the Loss of the Pleural Arch Hyperdevelopment of Combs in Bird-Fleas, Bat-Fleas, an, aekine g the Pleural Arch 4. Hy perdevelopment of the Pronatal Gon OREbAne: ES aainel Fleas, ete. 5. Convergence in Vestiture in Unrelated Fleas 6. Morphological Changes in the Thorax Associated with Lessened Jumping Ability in Fleas THE DEGREE OF DEVELOPMENT OF THE EYE IN VARIOUS FLEAS 1. Comparative Size of the Eye 2. Vision in Fleas 5 3. The Types of Thorax in leas with Reduced Eyes THE EVOLUTION OF THE CHARACTERISTIC PRONOTAL COMB ASCRIBED TO MOST NEST-FLEAS DISTRIBUTION OF THE NEST-FLEA PHENOMENON IN SIPHONAPTERA . THE ASSOCIATION BETWEEN THE MODE OF EGG-LAYING AND THE NEST-FLEA HABIT c : : : C 5 a c 5 6 COI aupwrn H Page 311 312 313 313 317 318 318 318 318 321 322 322 324 325 326 327 328 328 329 329 333 333 334 338 339 341 344 346 346 348 350 351 353 358 310 R. TRAUB H. REDUCTION OF COMBS, ETC. IN FLEAS OTHER THAN NEST-FLEAS) . 359 I. THE POSSIBLE RELATION BETWEEN THE PRONOTAL COMB AND A FRIGID ENVIRONMENT 2 361 J. STRUCTURAL AND BEHAVIOURAL VARIETY WITHIN A UNIFORM ORDER OF INSECTS ° é . c : ° 362 K. THE PRINCIPLE OF STRUCTURAL COMPENSATION . 3 6 é 366 V. CONCLUSIONS . < : : c : : ° : c : : 368 VI. ACKNOWLEDGEMENTS < c - : f ¢ 0 5 c o 369 VII. List OF TAXA CITED IN TEXT f a : 5 c a 0 5 369 VIII. List oF Hosts : : 3 5 é 3 : : 5 5 : 377 IX. REFERENCES . : 3 é 4 : : 5 . : o 377 X. INDEX . c ; : c : F . 3 5 2 a ; 383 ABSTRACT In the Order Siphonaptera there are many examples of the moulding effect of the environment —be it the fur or feathers of the host or its habitat—upon the physiognomy of the flea, resulting in a uniformity of appearance that may reflect convergent evolution, and not phylogeny or even homology. Additional data are presented suggesting that the pattern and shape of the spines of the pronotal comb of fleas may be more frequently adaptive than had been generally supposed, as indicated by the fact that unrelated fleas often have the same type of comb (or vestiture in general), while, in contrast, a variety of patterns may be exhibited by a single genus of flea, depending upon the host. Thus, in the squirrel-fleas in general, the second or third spines of the comb (counting from the bottom) tend to be relatively broad, and often slightly overlapping near the base. This has been noted in 13 genera and 3 families of fleas, and in all instances these spines are proportionately broader and shorter than in the case of members of the same taxon which are ‘“‘fur-fleas’’ of murids, cricetines, insectivores, etc. The trend is particularly well developed in fleas of diurnal tree-squirrels and has occurred on all four continents where these rodents occur. It is believed that modifications of this sort are directly associated with the diameter and shape of the hairs of the host, and it seems likely that the overlapping second and third spines of the comb of fleas of tree-squirrels would grasp the hairs in a scissors-like grip if the flea were pulled backwards by the host in attempts at dislodgement. Precise data on this point are lacking insofar as concerns fleas of squirrels. The modifications of the pronotal comb are by no means always correlated with the vestiture of the host, as is indicated by the fact that, regardless of host, 17 genera of fleas, representing 3 families and 5 subfamilies, exhibit the same unusual type of ctenidium, in which the spines are exceptionally or notably short and broad and relatively few in number. These are all “nest- fleas” in that they are much more prevalent in the nest than on the host, etc. and show reduction in general chaetotaxy, modifications of the thorax and other specialized features. The type of comb in these fleas is accordingly believed to represent a general trend towards reduction which is a feature in the evolutionary development of the nest-habit, which is discussed in detail. Fleas of volant and gliding hosts such as bats, birds and flying-squirrels have solved the dilemma of finding such a bird or mammal without a disastrous fall in the process and yet managing to remain affixed to the host despite its active flight or aerial gymnastics. One common way this has been accomplished is by becoming crawlers, rather than jumpers, and developing supernumerary spines in the pronotal comb, and/or additional combs, long bristles, etc. or by anchoring mouthparts. Such fleas often possess a thorax in which the pleural arch, under which is housed the elastic resilin which provides the power for leaping, has become completely or partially reduced. This thoracic modification is also seen in “‘nest-fleas’’ inhabiting tall trees, banks and caves, etc., where a fall would be hazardous, even if survived, because of the small chance of finding a suitable host. It likewise occurs in many nest-fleas of terrestrial or underground nests, and in other habitats, and reasons are advanced to indicate that the condition 1 CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 311 (in all cases) is adaptive. This modification of the thorax has been achieved by a variety of morphological changes, emphasizing that these are examples of convergent evolution. The relative degree of the development of the eye in fleas is also associated with the habits of the flea, or its host or environment. In many instances, reduction of the eye accompanies that of the metathorax and loss of the pleural arch, but notable exceptions are pointed out. The behaviour of various kinds of fleas in finding and feeding on their hosts is discussed. I. INTRODUCTION FLEAS are highly modified morphologically for an ectoparasitic existence (Rothschild, 1917; loff, 1929; Traub, 1966; Traub & Evans, 1967), and the specializations exhibi- ted include patterns of ctenidia (combs of spines) and chaetotaxy that somehow reflect the environment provided by the host (Traub & Barrera, 1966; Humphries, 1966, 1967; Traub, 1968, 1969), rather than merely the phylogeny or taxonomic position of the flea. The modifications in some instances are obviously adaptive and are concerned with the flea maintaining a grip on the host and escaping capture. Others are difficult or impossible to explain by referring to the vestiture of the infested mammal or bird, but can be correlated with the haunts or habits of the host. Some of the fleas discussed in the first article in this series (Traub, I972a) illustrate such examples of Convergent Evolution, and these and other relevant taxa will be treated in the present paper and new points will be presented along such lines. That the nature of the pronotal comb is adaptive is indicated by two generaliza- tions: (1) The same pattern of comb is frequently exhibited by unrelated fleas para- sitizing the same host, and (2) Within a genus there may be a variety of types of combs, depending upon the host. Therefore, by way of background, variations of the type of pronotal comb seen in a single taxon will be mentioned and associated with certain kinds of hosts. The characteristic features of the class of pronotal comb will be noted, and tabled data and illustrations will be presented for various genera of fleas, and for various kinds of fleas, such as squirrel-fleas, murid-fleas, nest- fleas, etc. In the section on discussion, it is pointed out that while in many instances the precise configuration of the comb can be explained by the nature of the hairs of the host, this does not apply to the type of comb seen in fleas which are characteristic inhabitants of underground nests, and which are modified accordingly, even though a variety of taxa, often unrelated, are represented. In order to demonstrate that this is so, and to offer an explanation, the phenomenon of nest-fleas is discussed in some detail and this leads to a consideration and review of two features commonly seen in nest-fleas, viz., the marked reduction of the eye and loss of the pleural arch. It is pointed out that in some nest-fleas, however, the eye is unusually large and this is shown to be adaptive, as is the loss of the pleural arch in bat-fleas, which are not nest-fleas. Thus, the morphological modifications are shown to be connected with | the pattern of life of the flea, and often with the habits of the host. It should be noted that no attempt is made in this article to review all of the various modifications in the pattern of pronotal combs seen in Siphonaptera. For example, the remarkable and well-known convergence exhibited by combed bird- fleas, in which the pronotal comb always bears a significantly greater number of spines than do allied forms infesting mammals, and which are otherwise specialized, will be treated in detail in a subsequent article. This phenomenon has now been 312 Re eo ReAGS noted in nine genera, representing three families (including a new genus and new records known to the writer). The type of comb deemed characteristic of fleas infesting nocturnal arboreal hosts has been mentioned previously (Traub, 1969), while Traub & Evans (1967) referred to some ctenidia of insectivore-fleas and desert- fleas, etc. and Traub & Barrera (1966) discussed ctenidia in shrew-fleas. Other points about ctenidia were treated by Traub in 1968, and some important considera- tions remain for future treatment. Instead, the present paper will primarily deal with the pronotal comb of fleas infesting arboreal squirrels and with that of nest- fleas in general, with passing reference to murid-fleas, etc., while, the resulting discussion leads to the points mentioned above. II. BACKGROUND The pronotal comb of Medwayella Traub, 1972 (fig. 6),1 a genus which generally infests arboreal and semi-arboreal sciurids, resembles that seen in genera of fleas which are characteristic parasites of tree-squirrels (figs 1-14). However, before considering that subject further, it is instructive to review the features of the pronotal comb in some other fleas, particularly Lentistivalius Traub, 1972, in which the 5 known species are closely allied, as indicated in the article on Medwayella, but which vary considerably in the kind of host infested and their geographical location. In this group of fleas it has been possible to glance at the spines of the comb and correctly state whether the host is a bird, rodent or shrew. Although no bird-fleas had been known in the family Pygiopsyllidae prior to 1950, the author of L. insolli (Traub, 1950b) suggested that the species infested birds because the pronotal comb (vide fig. 18) consisted of 26-28 narrow, fairly straight, closely set spines and the comb did not extend far enough ventrad to cover the third vinculum, thereby _ paralleling the condition in bird-fleas of the families Ceratophyllidae and Leptopsyl- lidae. (This belief about the avian hosts of L. insolli turned out to be correct.) In each instance, the mammal-infesting fleas related to all of these bird-fleas bear combs of 18-24 spines (generally around 20), while the spines are broader, differently shaped, i.e. concave or pointed, or else, if there is a larger number of spines present, the comb extends further ventrad, over the vinculum (Traub, 1969). On the other hand, when Smit (1958) described the flea now called L. alienus, he pointed out that the appearance of the comb (vide fig. 16) led him to state that the host was a rodent, because the spines were ‘‘fairly straight,” unlike L. ferinus (Rothschild, 1908) which is a shrew-flea, where “‘the spines are blunt, distinctly curved and longer than the pronotum,” as shown in fig. 15. A comb of the latter type is characteristic of many shrew-fleas (Smit, 1958; Traub & Barrera, 1966; Traub & Evans, 1967), and is in marked contrast to the fairly straight, pointed spines of L. aestivalis (Jameson & Sakaguti, 1954) (fig. 19). The spines of the comb of L. vomerus Traub, 1972, which is primarily a tupaiid-flea,? are longer and narrower (fig. 17) than in L. ferinus. 1 Unless otherwise indicated, the figures of combs herein are of male specimens. 2 In modern schemes of classification, the tree-shrews (Tupaiidae) are either placed with the primates or as a sub-order of Insectivora (along with macroscelids), distinct from ordinary shrews and moles. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 313 It seems surprising that a ‘simple’ structure such as the pronotal comb should be so flexible or adaptive in evolution, and hence it is worth emphasizing that there are other genera of Siphonaptera where several kinds of pronotal combs are found within the one taxon, depending upon the host-affinities of the species concerned. For example, this is the case in Ctenophthalmus Kolenati, 1856, where the great majority of the approximately 100 species parasitize microtine and murid rodents, and have pronotal combs of the type shown in figs'22 and 26, viz., the spines are quite straight and broadly coming to a point apically. In contrast, in the Ctenophthalmus which are shrew-fleas (e.g., figs 21 and 25), the spines are generally concave and bluntly rounded apically, while mole-fleas have spines which are straight, very narrow and acutely pointed (fig. 24, and similar modifications in mole-fleas of other genera are cited by Traub & Evans, 1967). The combs of Spalax-infesting members of this genus (fig. 28) tend to be quite straight and fairly narrow, but not as stiletto-like as the talpid-fleas. All of the groups of Ctenophthalmus mentioned thus far possess combs in which the majority of the spines are definitely longer than the pronotum, but this is not the case for the few species that parasitize sciurids (Citellus ground- squirrels) (viz. C. pollex Wagner & Ioff, 1926, fig. 23). In the latter instance, the spines are proportionately shorter and broader than in the foregoing species, while the third spine from bottom (herein designated as spine number 3) is somewhat broader than the others. This last point is significant because, oddly enough, this spine is generally broad (especially at the base) in all fleas of diurnal arboreal squirrels, as will be shown. This spine is relatively broad even if those above the more dorsal ones are somewhat narrower, but usually number 2 or number 4 spine is nearly as stout as number 3.3 III. DATA AND OBSERVATIONS A. THE PRONOTAL ComB There are other instances of taxa in which the pronotal combs of the members vary in configuration according to the host, and the number of examples that can be cited indicated that this is true convergence and is based on an actual principle. Thus, in the Holarctic ceratophyllid genus Monopsyllus Kolenati, 1857 (iii.1),4 which infests a variety of cricetid and sciurid hosts, it can be seen that the spines, particularly spine number 3, are broader (especially at the base) and shorter, and fewer in number in the species illustrated in figs 62, 64 and 66, which are true fleas of diurnal tree-squirrels, than in figs 55, 57, 59 and 61, which pertain to fleas of non-sciurids. The spines of the comb of the dormouse-flea (fig. 60) and of the flying-squirrel flea (fig. 58) are also longer and narrower than those of the fleas of diurnal squirrels, and exceed them in number. 3 The hyperdevelopment of a single spine of the pronotal comb reaches a bizarre zenith in the pygiop- syllid genus Tvaubia Smit, 1953a. Smit described this genus on the basis of one species, and cited as a generic character the fact that the number 2 spine was much longer than its mates. Today there are 9 species of Traubia known to me, and all possess this odd feature, which seems to be unique in the Order. This development, however, is in a single taxon, unlike the case of the stout spines in the squirrel-fleas. 4 The families and subfamilies are designated by Roman and Arabic numbers, e.g. iii.1, and a list of the names for these taxa are appended. 314 R. TRAUB Other examples of this trend are provided in the genera Orchopeas Jordan, 1933 (iii.1), Opisodasys Jordan, 1933 (iii.1) and Neopsylla Wagner, 1903 (1.7), some of which parasitize arboreal sciurids while others infest other kinds of hosts. In Orchopeas, the spines of the squirrel-fleas (figs 29, 31, 33 and 35) are significantly broader and shorter than are those of O. sexdentatus (Baker, 1904) (figs 30 and 34) and O. leucopus (Baker, 1904), which are fleas of peromyscines. Similarly, in Opisodasys keen (Baker, 1896) (fig. 43) and O. nesiotus Augustson, 1941 (fig. 45) which infest Pero- myscus, the pronotal spines are noticeably longer and narrower, and the comb extends further down, than in the case of fleas of diurnal tree-squirrels (figs 44, 46 and 48). The comb of O. pseudarctomys (Baker, 1904) (fig. 47) is worthy of special note. Its spines are more numerous and narrower than in the case of the fleas from the diurnal squirrels, and as noted below (Section C.4), combs of this type are characteristic of fleas parasitizing nocturnal, arboreal species. In Neopsylla, in which the majority of the 34 species known to me infest murines or cricetines, the combs of fleas of chipmunks or ground-squirrels (figs 37, 39 and 41) bear spines that are proportionately definitely shorter and broader than in the murid-fleas (figs 38, 40 and 42). The second principle, namely that the combs of sundry fleas of a particular kind of host tend to converge to a uniform configuration, is well shown in the fleas of arboreal or semi-arboreal squirrels. Thus, figs 1-14 illustrate the pronotal combs of 14 species of fleas of such squirrels, representing 13 genera and 3 families of fleas, and it can be seen that in all of these, the spines in general, and at least numbers 2—4 in particular, are broader and proportionately shorter than those of fleas infesting insectivores (figs 21, 24 and 25), murids (figs 49-54, 38, 40, etc.), and the peromyscine cricetids Peromyscus (vide figs 36 and 43), and Neotoma (fig. 32). For example, spine number 3 of the pronotal comb is 4:2 times as long as broad near the base in the case of Aenigmopsylla grodekovi Sychevsky, 1950 (fig. 9); 4-1 times in Macrostylo- phora h. hastata (Jordan & Rothschild, 1921) (fig. 14) and 3-1 for Libyastus infestus (Rothschild, 1908) (fig. 13), all of which are ceratophyllid squirrel-fleas, and 4-6 in Medwayella robinsoni (Rothschild, 1905), a pygiopsyllid from squirrels. In contrast, the ratio of the dimensions of this spine is 5-5 in Lentistivalius ferinus (Rothschild, 1908) (fig. 16), and 6-2 in Ctenophthalmus (Alloctenus) cryptotis Traub & Barrera, 1966 (fig. 20) both of which are shrew-fleas; 5-5 in Neopsylla dispar Jordan, 1932 (fig. 40) and 5-4 in Stivalius cognatus bamus Traub, 1972, both of which infest rats. Attention is directed to the fact that in fleas of arboreal or semi-arboreal squirrels in general, and especially in those of diurnal species, pronotal spines numbers 2 and 3 are particularly broad near the base, and number 2 tends to overlap number I proxi- mally, while the base of number 3 is usually very close to that of number 2 or overlaps it. In this regard it is important to note that the greater breadth of the spines in pro- portion to length, which is here indicated as being characteristic of squirrel-fleas, is a relative feature, not an absolute one that can be expressed in an “‘average”’ ratio of length to width. Thus, while it is a general rule that the spines of the combs of squirrel-fleas are shorter and broader than those of fleas infesting non-sciurid hosts, it is also necessary to consider the norm for the taxon in question, for any modifica- tion of a pronotal comb must be incorporated upon an existing framework. It is 315 AND TAXONOMY OF FLEAS CONVERGENT EVOLUTION “uInjOU JO UISIVU [esIOp JO Y4Sua]/[es1opqns [e}UOZTIOY JO YSus, = wWinJOUCIG/qUIOD :O17PY » snastumosag oS61 ‘qnely, Z+I 1mosayjvU SYaDYI0aT snyvy Oot “BI (ZZOI ““syyOYT ZI yy uepsof) wssojy snyvays snypy gt “BLT $C61 ‘qnery, =+-1 puny) vjydsdoan spidjey +z ‘sty €ogr ‘WeusOy S-I $nqojWapojzI0S2q snuqoyzyqgougy pumojoan ze ‘siz €F61 ‘uosjsnsny +-1 apmojzoau svadoyI4Q, snashmosad 9€ ‘SUT (Fo6r ‘1ayeg) €-1 sndoana) svadoyr4Q snaskmodag €F ‘31 (96g1 ‘10yeq) 1-1 qaay skspposidaQ syey OF “Bry (COO pyryosyzoy) E+ siswaimipuoy “7 snqdsqoson stmoystug SS -8rq (2€6r ‘uepiof) z-1 syixa snqAsdouopy ~ et snashmosag ‘BIq «(6z61 ‘ueprof) 1-1 snquivyy snpjAsqouo pr SNJOAILTAT (So6r ‘pyryosyjoyy) I+ 1 quramb srayzoqvsa fy ySOF] uinjou PHNIOS gich ral -u0N ROT /quio9g x OTFEY z.G z-9 b.v WyPIM. /q38ue7] autds £€ ON SLSOH YAHLO 40 SVATA AO ASOHL g-€ $.¢ c.€ UIPIM. /y38u0T aurds € ‘on O-1 zI-Stg gS6r ‘e1oleg MUOAAQOS SYADYIOIT 8-0 9 “BIT (SO6r “pryosyy0 yy) WOSUIQOA KYadvMpa TT Del 6€ B17 (S161 ‘pyryosyzoyy) vuridour vydsdoan I-I €2 “SY QZOI ‘OT » 1ouse A xayjod snuypyjyqdouay O-1 6z ‘S81 (So6r ‘pyryosyz0y) sodau ‘wu svadoyotQ ZI zB (S6gr ‘1axeq) tpapmoy svaqoyIAQ L.o bb 317 LY61 ‘qneiry upunjjoy skspposidc $.1 €C6r 41ws sisuauojiaa snqjAsqoson 6.0 +9 ‘31yy (Fo6r ‘10yxeg) uosia snydsdouopy I (S061 ‘pyryosyqoy) idjowna a snyjAsqouo py 6-0 (Sz6xr ‘uepi0f) snqaaaw siAyjzoqnsa Ay uinjou -O1g /quiog ea xOTFEY yortmbs-ae1y, sjarinbs-punoiry essoT (je1ummbs-puno1y) snqqanvg (jermmmbs-puno1y) SEL ELD) yotambs-aory, Jorunbs-ae1, yermnbs-aery, yorummbs-wyeg yermbs-ae1], yunurdry) yunudry5 4sOH prog HIIM GauVdWNOO SV SVATJ-1TaNNINGS AO SANIdS ITVLONOUd NIVLYAD AO SNOTLNOdOUd AAILVIAY I alavy 316 R. TRAUB patently absurd to expect a combless flea parasitizing an African ground-squirrel to develop broad spines, especially if descended from combed ancestors, for according to Dollo’s Law, structures lost in the course of evolution cannot be resurrected per se, as discussed by Jordan in 1947 and by Traub in 1968 and 1969. Similarly, ina taxon where the pronotal spines are in general fairly narrow, the response to parasitism of squirrels could not be validly expected to equal that in Ovchopeas, where the spines tend to be fairly broad in all species, regardless of host. In Nosopsyllus Jordan, 1933 the spines are quite broad, as a rule, and hence in that genus an ‘‘average”’ index of length to breadth would exceed the standard for Ctenophthalmus. The best way to examine this hypothesis, therefore, is to compare various members of genera which parasitize both sciurid and non-sciurid hosts and see if there is a significant difference. Such a comparison is presented in Table 1.° From these data it can be seen that in each case, the squirrel-infesting species have pronotal spines that are significantly broader in proportion than those of its allies on other hosts. Moreover, as shown by the comparison with the length of the pronotum, which is also presented in the table, the spines of the comb of squirrel-fleas are also definitely shorter. There definitely is an inverse correlation between the length of the spines and that of the pronotum, as per what has been designated as the Principle of Totality of Mass (Traub, 1969). Of the fleas reported from tree-squirrels, the one with the proportionately narrowest spines is the Central American ceratophyllid Kohlsia graphis (Rothschild, 1909). Here the number 3 spine is 4-3 times as long as broad near the base, and the dorsal spines are about 1-2 times as long as the dorsal margin of the pronotum. However, the comparable figures for the related K. gammonsi Traub, 1950 are 4:8 and 1-4 respectively, so there is a significantly greater relative breadth in K. graphis. How- ever, it should be noted that it is not certain that the latter is truly a squirrel-flea. Apparently there are only about 24 specimens known (in all collections extant), and although virtually all of these are from squirrels (Nicaragua, Costa Rica, Panama, etc.), Tipton & Méndez (1967) could only find 16 on a total of 60 Sciwrus granatensis chiriquensis in Panama, all of which had been carefully examined for fleas. It may be that the true host has not yet been discovered, but my belief has been that K. graphis has only relatively recently transferred to squirrels. Tipton & Méndez (1967) have independently come to a similar conclusion regarding Kohlsia Traub, 1950 in general in Panama, and state that it ‘““may be that some of the true hosts no longer occur in this area and certain of the present hosts have been ‘adopted’.”’ It should also be borne in mind that these differences are consistent within the taxon, i.e. the samples are representative of the comb of squirrel-fleas whether only a few members infest arboreal or semi-arboreal squirrels, e.g. 1 of 24 for Kohlsia, 1 or 2 of 23 for Pleochaetis Jordan, 1933, 1 of 12 Paraceras Wagner, 1916, 6 of 46 Nosopsyllus; or whether the majority of the species do, viz. all 28 species of Macro- stylophora Ewing, 1929, 12 or 13 of 13 species of Medwayella, 8 of 11 Orchopeas and 16 of 22 Monopsyllus. An idea of the comparative differences in the proportions that ® Inasmuch as no Medwayella are known to consistently infest non-sciurids, the representative species in the table is compared with a member of a related genus. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 317 are involved may be gained by reviewing the illustrations of sundry Orchopeas (figs 29-36), Opisodasys (figs 43-48), Monopsyllus (figs 55-66) and Neopsylla (figs 37-42). A further illustration of the principle that the fleas of a particular kind of host tend to disport the same type of pronotal comb is provided by Neopsylla. It is to be noted that in the murid-species (figs 38, 40 and 42), the pronotal spines are relatively long, narrow and straight. This is characteristic of murid-fleas in general (Traub, 1968) and examples are shown in figs 49-54, depicting combs ot murid-fleas from 3 families, infesting such hosts as Mus, Apodemus and subgenera of Rattus. Additional examples may be seen in figs 1g and 20. B. THE PLEURAL ARCH For an Order that is often thought of as “homogeneous,” there is a surprising amount of variation on the structure of the Siphonapteran thorax, and to a consider- able extent the differences represent adaptive changes and not phylogeny, as will be noted in the section on Discussion. One of the major elements involved is the so- called pleural arch, which is present and well developed in some species and reduced or absent in others, and the two extremes may at times be seen within one genus, viz, well developed in Opisodasys hollandi Traub, 1947 (iii.1) (fig. 87, PL.A.) and Cerato- phyllis styx Rothschild, 1900 (ii1.1) (fig. 91, PL.A.) but absent in O. vesperalis°6 (Jordan, 1929) (fig. 88) and C. arcuegens°® Holland, 1952 (fig. 90). It is fairly well developed in Xenopsylla vexabilis Jordan, 1925 (ix.1) (fig. 93, PL.A.) but so reduced as to be virtually absent in X. papuensis® (Jordan, 1933) (fig. 92, PL.A.(?)). In each of the above instances where the pleural arch is well developed, the meta- thoracic region may be described as follows: There is a stout pleural rod (PL.R.) appearing as the margin between the metepisternum (MTS.) and metepimere (MTM.), arising from the dorsal portion of the base of the metacoxa and terminating at or about the ventrocaudal angle of the subquadrate lateral metanotal area (L.M.). The latter sclerite is quite well developed, e.g. longer than high, and as long as the third vinculum (VC.3) which links into it and at least several times as tall (high) as VC.3. Above the expanded dorsal apex of the pleural rod, which is at level of VC.3, or slightly lower, there is a conspicuous gap surmounted by the semi-ovate pleural arch (PL.A.). In O. hollandi the arch is at level of the middle of L.M. Above the pleural arch, the dorsal and caudal margins of L.M. merge and continue dorsad as the caudal margin of the metanotum (MTN.) and the base of the metanotal flange (MT.F.). The metepimere (MTM.) flanks the pleural rod and continues dorsad laterad to L.M. and then turns dorsocaudad and runs to the anterodorsal region of the first abdominal tergum (1 T.). The short thickening known as the squamulum (SQ.) serves as a good landmark for the anterodorsal region of the metepisternum. In the two illustrated ceratophyllids which lack the pleural arch (0. vesperalis®, fig. 88 and C. arcuegens°, fig. 90), the pleural rod (PL.R.) terminates at the ventro- caudal angle of the lateral metanotal area (L.M.). In each case L.M. is narrower than in the species possessing PL.A., and in O. vesperalis°, L.M. bears heavily tanned inter- nal margins and is quite ovate, while the pleural rod (PL.R.) is quite convex. In the 6 ° denotes species in which the pleural arch is absent, or essentially so, and these are indicated thusly henceforth. 318 R. TRAUB pulicid, X. papuensis® (fig. 92), the pleural arch at first glance seems to be missing completely, but actually is a very narrow structure (PL.A.(?)) virtually hugging the apex of the pleural rod (PL.R.) near the ventrocaudal corner of the lateral metanotal area (L.M.); the cap is almost contiguous to the rod on the sides as well as on top. L.M. is fairly large and quadrate, but smaller than in X. vexabilis (fig. 93). The metepisternum (MTS.) extends higher and is longer than in other members of the genus. As can be seen in fig. 89, in the bat-flea Nycteridopsylla eusarca® (Dampf, 1908) (v.1), there is no sign of the pleural arch, and the lateral metanotal area (L.M.) is very narrow and longer than high. The pleural rod (PL.R.) is flat, and serves as the margin between the metepisternum (MTS.) and the metepimere (MTM.), which is not the case in some other ischnopsyllids, e.g. Myodopsylla Jordan & Rothschild, rg11, in which the caudal border of MTS. overlaps MIM. The two hystrichopsyllids shown in figs 85 and 86 likewise lack the pleural arch, but they present a thorax which is very different from the types described above (and differ from one another) but these are treated later, in the Discussion. IV. DISCUSSION A. THE FUNCTION OF THE PRONOTAL COMB I. GENERAL COMMENTS. The modifications of the pattern of the pronotal comb consistently exhibited by shrew-fleas, bird-fleas, etc. should be adaptive, i.e. of some significant value to the insect, and the most obvious function for the specialized comb would be to serve as a device to prevent backward motion amongst the hairs or feathers of the particular host, for, as N. C. Rothschild pointed out in 1917, even non- combed fleas are unable to move backwards, thanks to the firm caudad-directed bristles of the body. Thus, the mere presence of a comb would facilitate ready forward passage through the hairs, but the inclination of the spines and the gaps between them would have to jibe with the structure and density of the hairs of the host before they could effectively prevent backward motion induced by teeth, beak or claws of the host intent on destroying or removing the flea. The various referen- ces cited in the Introduction include data and discussions on these points, and Humphries’ articles (1966, 1967) include specific measurements showing how, in designated instances, the combs of a variety of fleas are so framed that they effec- tively entrap the hairs of the host in backward locomotion. Ioff, as early as 1929 tried to account for the unusual, short, pale, bluntly pointed or rounded genal spines in Ctenophthalmus (Neoctenophthalmus) dilatatus Wagner, 1928 in terms of the “extremely thick soft silky fur of the hosts of this flea’ (the burrowing mole-rat, Myospalax).? 2. SPECIAL KINDS OF ComBS ON FuR-FLEAS. In speaking of fleas which move about fairly freely in the pelage of the host, viz., the species which Toff (1929) called ? Toff (1929) also speculated that the type of genal comb in C. dilatatus represented an acceleration or development of the trend in this group of fleas towards variation in number and reduction in size of the spines of the genal comb. He thought this evolutionary trend might be due to the frequency of overt damage to ctenidial spines seen in these fleas. (Such broken spines are often seen in fleas of other fosso- rial hosts as well—R.T.) CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 319 “fur-fleas,’’ Traub (1968) pointed out that fleas with stiletto-like pronotal spines are characteristic parasites of peramelid marsupials (bandicoots) while their close relatives on rats have bluntly pointed spines and that these ctenidia correlate well with the nature of the fur of the host. Holland (1969) also independently observed the convergence of the pointed spines seen in the pronotal comb of most bandicoot- fleas. Recent collections in New Guinea provide new data reinforcing this hypo- thesis. Thus, on the basis of the six species then known, Holland (1969), when describing Papuapsylla, understandably concluded that the pronotal spines were characteristically bluntly pointed or rounded at the apex. These species infest sundry rats, and the same is true for the minimum of eight new species of Papua- psylla known to me which possess a similar type of comb of broad, subrounded, slightly upcurved spines. However, two undescribed Papuapsylla from bandicoots have combs with spines that are straight and relatively acutely pointed, again indicating the moulding-influence of the vestiture of the host upon the flea. Humphries (1966) well demonstrated how the pronotal spines of certain fleas (which were not squirrel-parasites) ‘“‘appeared to be structured in relation to the host’s hair,’’ and there can be little doubt that the processes of natural selection have tailored the pronotal combs to conform with the host-pelage in many instances. It seems likely that this may also be the case in the fur-fleas of diurnal arboreal and semi-arboreal squirrels, for, as shown above, all the combed species exhibit essentially the same type of ctenidium, even though a variety of taxa of fleas and many genera of sciurids and four continents are involved. Not only do these combs bear relatively broad spines subventrally, but, the bases in particular are broad, and the gaps be- tween spines number I and number 2 and spines number 2 and number 3 are not only acute proximally, but often overlap at the base. As a result, any hair slipping between the teeth would be grasped in a scissors-like vise if the flea were pulled backwards. Moreover, this modification seems to be at the precise height to best grasp the axil of the hairs in their normal position. However, this hypothesis needs testing by experimentation and careful measurements, for we lack such data, and further, the theory implies that the sundry squirrels have a basically similar type of vestiture, and we do not know if this isso. Nevertheless, the similarity of the hairs should not be expected to be precise, for the combs of squirrel-fleas are not fully uniform, viz, the spines in the fleas of semi-arboreal species like Dremomys, Rhino- sciurus, Lariscus etc. are often somewhat narrower than those of fleas of truly arboreal squirrels. This can be seen in Macrostylophora borneensis (Jordan, 1926) (fig. 96), a flea of Dremomys, as compared to M. hastata (Jordan & Rothschild, 1921), a flea of tree-dwelling Callosciurus. This in turn suggests that the hairs of Dremomys are coarser than C. evythraeus. Simple comparative measurements alone will not suffice, however, for heed must be paid to the precise parts of the body of the squirrels where particular species of fleas are most apt to be found, because the sizes of the hairs vary, and so do the habits of different fleas (Traub & Evans, 1967). As those authors indicated, apparently there is limited mobility in the comb itself, so that it can be either somewhat appressed to the body or slightly removed from it, and this is another factor to consider. Certainly, however, the present writer can advance no 320 R. TRAUB other hypothesis but the matching of hairs and spines to account for the convergence in the pronotal combs of the fleas of diurnal, arboreal squirrels. Holland (1964) reported that “‘Archaeopsylla spp. (Pulicidae) and Bradiopsylla echidnae (Denny, 1843) (Pygiopsyllidae) share a structural modification of the pronotal comb not otherwise known in either family,”’ referring to the fact that the ctenidium consists of only a very few spines (dorsal in position) and that these are fairly well separated, very short and are broad at the base and apically pointed. Holland then went on to say that “it may be significant that the former infest hedgehogs (Evinaceus) which are spiny insectivores while the latter lives on echidnas (Tachyglossus) which are spiny monotremes’’. It seems to me that this noteworthy observation really does pertain to an example of true convergence in that the nature of the comb can be explained by considering the spinose vestiture of the host. That is, the broad gap between the pronotal spines would appear to correspond with the diameter of the spines of the host, and the short pronotal spines could successfully span the mammalian spines, fitting snugly and securely against them. In contrast, a comb of long spines could not do so unless the spines were extremely divergent apically—far more so than is known in any flea, and, in fact, the apices of the spines would tend to overlap and cross one another. Moreover, the divergence between the short spines of these fleas of spiny animals presumably also accounts for the short height and dorsal position of the comb. A long comb is incompatible with subdorsal spines that are markedly divergent, for these spines would have to overlap and hence not function properly. That there is an intimate association between a short pronotal comb of small, widely spaced spines, and infestation of spiny animals is further suggested by the fact that the monotypic pulicid genus Centetipsylla Jordan, 1925, a parasite of Madagascar ‘‘hedgehogs”’ (tenrecs), has a comb of this type, while related genera, infesting non-spiny hosts, do not. However, instances have also been cited where the relationship between comb of the flea and vestiture of the host cannot be so simple (Traub & Evans, 1967). An example of this is the ‘“‘flared comb”’ of some shrew-fleas and macroscelid-fleas which arches back over the mesonotum, as described by these authors. Here the gap between some of the spines is inclined at nearly right angles to the longitudinal axis of the body, and hence it would be difficult for such a comb to grasp and retain hairs if the flea were in the usual position most fleas assume when feeding. However, if these particular fleas act like those noted in New Guinea which have marginal rows of spiniform bristles on the head (and/or flattened pronotal margins), and incline the body head-downwards and thus hook the hairs of the host (Traub, 1968, 1969), the flared comb may then be an effective mechanism against dislodgement. This hypo- thesis can be verified only by direct observation, however. Another example of a type of comb which presumably is not directly associated with host-pelage is that characteristic of fleas found in the fur of mammals in desert areas, in which the teeth of the comb are long and narrow, are close-set at the base and point somewhat dorsad or laterad so that the apices are as remote as possible from the mesonotum (Traub & Evans, 1967). The Orchopeas of desert-Neotoma have combs of this pattern, viz, a subspecies of O. sexdentatus (fig. 34) and O. neotomae CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 321 Augustson, 1943 (fig. 32), in contrast to O. sexdentatus agilis (Rothschild, 1905) (fig. 30), from a mesic habitat. 3. THE PRONOTAL CoMB OF FLEAS OF Non-ARBOREAL Nests. The type of pro- notal comb which is most difficult to associate with host-pelage is that exhibited by certain mammal-fleas which presumably characteristically infest the nests of their hosts (usually underground or on the surface) and which are seldom found on the mammals themselves, apparently feeding while the hosts are in the nests (Traub & Evans, 1967). Examples of the combs of fleas believed to be of this type are shown in figs 67-84, 86 and 98, and it will be noted that in these the spines are much broader and generally much shorter than those indicated for squirrel-fleas (and hence far more so than in shrew- and murid-fleas, etc.). In such combs there are only 6 or 7 spines per side, instead of the usual 8-10 or more, and the spines are only 3-5—-4-5 times as long as broad, instead of 5~7 times or more. Some of the number 2 spines are so stout that they are scarcely thrice as long as broad. (Amongst fleas in general there is a good correlation between a low number of spines in the comb and unusually broad girth, as compared to large numbers of spines and narrow breadths, provided the comb is of full size, i.e. extends down to near the vinculum.) It is remarkable how large a variety of fleas, from a large assortment of hosts and from widely separated parts of the world, present essentially this type of pronotal comb. Included in the illustrations are representatives from 17 genera from 3 different families of fleas and a total of 5 subfamilies. Geographically, North America, Central America and Asia are listed, while 12 genera of hosts are cited, and these refer to 2 orders; 2 families of 1 of the orders; 3 subfamilies of cricetids and 2 tribes of cricetines. The question immediately arises as to why fleas of such diverse backgrounds should uniformly present a single morphological feature. It seems extremely unlikely that such a variety of hosts should converge to the same type of pelage and that hence the pattern of pronotal combs represents parallel development of spines adapted to fit the hairs of the host. Thus, if the type of pronotal comb were determined purely by the type of fur of the host, then there should be no difference between the pattern of ctenidium exhibited by a flea which feeds while in the fur of an active host and that of one feeding on that host in the nest. However, frequently the nest-flea has a markedly different type of pronotal comb than the flea living in the fur of precisely the same individual mammal, e.g. the nest-flea Conorhinopsyllanidicola® Jellison, 1945 (i.2) (fig. 86) versus the fur-flea Orchopeas sexdentatus (iii.1) (figs 30, 34) on Neotoma; Phalacropsylla paradisea Rothschild, 1915 (i.7) (fig. 78), versus O. neotomae Augustson, 1943) (iii.1) (fig. 32), also on Neotoma; or nest-fleas like Catallagia charlottensis (Baker, 1898) (i.7) (fig. 67) and Delotelis telegoni (Rothschild, 1905) (i.7) (fig. 74) in contrast to fur-fleas like Monopsyllus wagneri (Baker, 1904) (iii.1) (fig. 59) and Opisodasys keeni (iii.1) (fig. 43), which infest Peromyscus. Most of these genera of nest-fleas do not include fleas which also infest the fur of the hosts, but Pleochaetis is an instructive exception. P. paramundus Traub, 1950 (fig. 62) is the only representative which is believed to be a nest-flea and it has relatively short and broad pronotal spines. Amongst the other 22 species, the one with the broadest spines in the comb is the 322 R. TRAUB squirrel-flea, P. soberoni Barrera, 1958 (fig. 12) and, of course, its spines do not com- pare in breadth with those of the former species. In order to explain the phenomenon of convergence of the pronotal comb to this type, it is necessary to digress and first discuss nest-fleas in general, as well as such topics as the pleural arch of the metathorax and the varying degrees of the develop- ment of the eye seen in various fleas, since morphological modification of these structures are marked in nest-fleas. B. NeEsT-FLEAS IN GENERAL 1. INTRODUCTION. Various students of Siphonaptera have noted that certain groups of fleas are relatively seldom found on the bodies of their hosts but are never- theless common within the nests of the mammal or bird concerned, and that these fleas are morphologically modified in various ways that reflect their pattern of life, viz., the tendency to remain in nest-litter or to crawl into cracks and crevices in the nest and to feed on their hosts while the latter are in their nests, perhaps while they are sleeping. Presumably they are found on the host only if the mammal or bird leaves the nest prematurely, as when frightened or during the breeding season, when the hosts rarely linger in the burrows, and thus the flea is interrupted in its feeding. As indicated below, the true nest-fleas differ substantially in appearance from ‘“‘fur- fleas’ which spend most of their time on the pelage of the host, even when active out of the nest, and are modified accordingly. However, before proceeding, it is necessary to point out that the term “‘nest-flea’’ has been used in several different senses. off (1929) used it to refer to all species which were more common in the nest than on the host, even though they may have often been collected on the host, and even though they were adapted for living in the fur. Hubbard (1947) also noted that it was far easier to collect certain fleas in the nest than on the mammal. In 1941, loff pointed out that ‘“‘fur-fleas” need frequent blood meals and apparently cannot survive if off the host for long periods, whereas the “‘nest-fleas’’ were only occasionally found on the hosts and could remain away from the mammals long enough to be able to hibernate in the nest in its absence. off thus made distinctions that were physio- logical and ecological and of undoubted importance in the epidemiology of plague. Nevertheless, no further definition was given, and, indeed, probably would have been impossible because of the variations in degree that occur in the two groups of fleas, and because the habits of the fleas may vary with the season, etc. As will be shown, some bird-fleas feed avidly while the young are in the nest and then undergo prolonged fasting. Sazonova (1962) cited other examples of species “that feed more frequently than nest-fleas but are also able to survive for a long time in empty nests,” and alludes to “such typical nest species as Neopsylla setosa [that] spend therefore a longer time on the host than usual’ when the female fleas are maturing their eggs. For these reasons Zhovtyi (1960) validly advised against placing fleas in such ecolo- gical groups until such time as more information had been obtained about their habits, bionomics, etc. Zhovtyi later (1963) developed this theme further, and objected anew to classifying fleas into “‘nest’’ and “‘fur’’ categories, pointing out that in his view the fleas in question (Citellophilus tesquorum (Wagner, 1898), Neopsylla setosa (Wagner, 1898), Ctenophthalmus pollex and several species of CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 323 Xenopsylla) were all ‘‘more adapted to the nest than to the fur of their hosts and also spend more time in the nests than on the hosts themselves’. An additional com- plicating factor was that the same species were found to be more prevalent in the nest than on the host at certain times or seasons, whereas at other times the reverse was true. It should be borne in mind that Zhovtyi and the other Soviet authors were dis- cussing differences in behaviour that are physiological in basis, depending upon the life cycles of the fleas (which in turn may prove to be attuned to the hormonal or physiological state of the host itself, as at times of hibernation or breeding) or, as Zhovtyi (1963) indicated, upon ecological conditions such as temperature or atmospheric humidity. In the present paper, however, we are discussing other types (and other taxa) of fleas, e.g. those which are morphologically highly specialized for feeding upon their hosts when they are in repose in the nest, and which rarely accompany the mammal or bird out of the nest. These specially adapted fleas are herein referred to as nest- fleas because the term is convenient, has been used in the past in this sense, and because, for our purposes, it can be defined in terms of structural modifications. In this case too, it will be shown that there are varying degrees of specialization, but the concept of nest-fleas, is an important and useful one. The species called nest-fleas in the present paper are characterized by possessing several or many of the following features: (1) An eye which is rudimentary, vestigial or absent. (2) Reduction in chaetotaxy, including the numbers of rows of bristles and the numbers of bristles on various parts of the body, and in diameter of the bristles. Even the antepygidial bristles may be exceedingly few in number and/or be very thin, and even at times be present in females and absent in males, i.e. most Rhadinopsyllini. (3) Reduction in numbers of spines in combs, or the numbers of the ctenidia, or the height and length of the comb; or else the elimination of combs altogether. (4) A similar, but even more pronounced reduction in or elimination of the apical spinelets, viz., the so-called ‘‘minor combs.”’ (5) Reduction of the thorax, including shortening of sclerites such as the lateral metanotal area; consolidation of parts; diminution in size of the pleural arch, or its loss altogether; or reduction in the degree of sclerotiza- tion of the metanotal flange, or in its size; and loss of its apical spinelets. (6) Legs telatively long and slender. (7) Lessening of the ability to jump, or concomitant increase in the facility of crawling and entering tight cracks. (8) Hyperdevelop- ment of certain structures, e.g. lengthening of stylets and labial palpi (at times with an increase in the number of joints), development of broadened sections of antennal segments which enclose other segments; or of lobes on the coxae, or of straight or angled anterior margins on some thoracic sterna; elongation of legs at times accom- panied by development of a fringe of long bristles on the male hind tarsus. (9) Loss in degree of sclerotization of caudal margins of abdominal segments. This may be accompanied by hyper-sclerotization of the antero-dorsal portions. (10) Diminution in size. At least some of these points were mentioned by Holland, 1949, 1952, 1965; Hopkins & Rothschild, 1962; Lewis, 1969; M. Rothschild & Clay, 1952; Traub, 1950a, 1950b, 1952, 1953a, 1953b; Traub & Tipton, 1951; Traub & Barrera, 1966; Traub & Evans, 1967; and M. Rothschild, 1969. It is also apparent from the B 324 R. TRAUB comments by N. C. Rothschild (1917) and Jordan (1937) on Convergent Evolution, as well as from their descriptive notes on Phalacropsylla Rothschild, 1915, Anomio- psyllus? Baker, 1904 and Callistopsyllus° Jordan & Rothschild, 1915 (vide N. C. Rothschild, 1915; Jordan & Rothschild, 1g15) that these scientists were fully cognizant of such modifications and the correlation with nest-inhabitation, although they usually did not refer to the association by name. The known acme of specialization as nest-fleas are in forms like Anomiopsyllus° and Wenzella° Traub, 1953, which lack combs, eyes, and the pleural arch entirely, and which have long slender legs and mouthparts. Even the apical spinelets, possessed by most fleas and believed to represent vestigial combs (Traub & Evans, 1967), are missing. In the former, the species are often virtually nude, or the few bristles present are short and thin. In Wenzella°, the bristles are very thin, while the inter- calary bristles of the abdominal rows are virtually only vestiges. In Jordanopsylla Traub & Tipton, 1951, this process, which has been termed “evolution by loss’’ or “specialization by simplification’ by Hopkins & Rothschild (1962) has proceeded to the point of reduction of the eye and pleural arch, but not to their complete elimina- tion, although the combs are absent, the mouthparts elongate, legs lengthened, etc., etc. In Megarthroglossus° Jordan & Rothschild, 1915 the modifications include extremely long mouthparts, coupled with reduction in the eye and vestiture, loss of pleural arch, etc. and while there is a pronotal comb, it consists of only 7-8 short, broad spines per side. Other genera possess these attributes to varying degrees, and on these grounds, as well as the fact that the species involved have rarely been collected on reported hosts, the following are cited as examples of nest-fleas. (In many instances the nests of the hosts have been insufficiently studied to permit one to state that these fleas are actually more numerous in the nest than on the birds or mammals themselves.) 2. HysTRICHOPSYLLID NEstT-FLEAS. Amongst subfamilies of Hystrichopsyllidae, 7 of the 8 Anomiopsyllinae genera are deemed nest-fleas, viz.: Anomiopsyllus”, Conorhinopsylla® Stewart, 1930, Callistopsyllus°®, Eopsylla® Argyropulo, 1946, Jordanopsylla, Megarthroglossus°, and Wagnerina loff & Argyropulo, 1934. The exception, Stenistomera® Rothschild, 1915, is discussed later asa special case. (From the ° mark, it can be seen that 6 of these genera lack the pleural arch.) Of the 7 known Rhadinopsyllinae, all of which have no pleural arch, Rhadinopsylla°® Jordan & Rothschild, 1912, Stenischia® Jordan, 1932, and Wenzella° are regarded as nest-fleas. Trichopsylloides’ Ewing, 1938, is probably a nest-flea, and is discussed further in Section IV.C.5. Of the 11 Neopsyllinae, at least 8 fall in the category of nest-fleas, namely, Catallagia Rothschild, 1915, Delotelis Jordan, 1937, Epitedia Jordan, 1938, Genoneopsylla Wu, Wu & Liu, 1966,8 Phalacropsylla, Paraneopsylla Tiflov, 1937, Strepsylla Traub, 1950 and Tamiophila Jordan, 1938. (The exceptions are Meringis Jordan, 1937, Neopsylla, Rothschildiana Smit, 1952.) One of the 6 genera of 8 New Synonymy. Evansipsylla Traub, 1968 is a synonym of Genoneopsylla Wu, Wu & Liu, 1966, the description of which escaped my notice when the Chinese ceased filling subscriptions to their journals. The type-species proposed, thysanota Traub, 1968 is presumably different from the type of Genoneopsylla, namely longisetosa Wu, Wu & Liu, 1966, but one cannot be certain because of the lack of details in the sketches that illustrate longisetosa. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 325 Ctenophthalminae seem to be a nest-flea, viz. the South American Agastopsylla° Jordan & Rothschild, 1923, as indicated by the marked reduction of the thorax, and in chaetotaxy, etc. It has been pointed out that the short pale spines in the genal comb, and their frequent variation in number (even on the two sides of the same specimen), all are suggestive of “an evolutionary tendency in the genus towards complete reduction of the comb’’ (Traub, 1952), while morphological similarities with anomiopsyllines were therein deemed examples of convergent evolution in unrelated nest-fleas. It is possible that one of the 5 Doratopsyllinae may prove to be a nest-flea, viz. Idilla° Smit, 1957, an Australian marsupial-flea, in which there is rather great reduction in chaetotaxy and loss of the pleural arch. However, there seems to be little else to suggest modifications along the lines we have been discussing. The well developed genal comb and pronotal comb, plus the fact that some of the few bristles on the thorax and one on the head are unusually short and stout, lead me to believe it is specialized in another direction, as indicated in Section B.g below. Of unusual interest are hystrichopsyllids which possess some of the typical attributes of nest-fleas but which nevertheless bear genal and ctenidial combs, e.g. Stenischia®. Thus, these fleas are characterized by: reduction of the eye, thorax (including loss of the pleural arch), and chaetotaxy; long, slender legs; development of heavy incrassations on the dorsal margins of the abdominal terga and ventral margins of the sterna, coupled with unusually lightly tanned median portions of these segments—all features associated with nest-fleas. Here too, in my opinion, the ctenidia are reduced, but the term is a relative one, in that what appears to be well developed combs in reality probably represents a marked diminution in the size and number of spines borne by their ancestors, or by related taxa such as Nearcto- psylla Rothschild, 1915, which have much more pronounced combs. This point is considered further in Sections IV.C.2 and IV.E below. 3. PyctopsyLiip Nest-FLEAs. Amongst the pygiopsyllids, there are only very few instances where fleas seem to have assumed the nest-habit. One is Choristopsylla tristis® (Rothschild, 1900), which, even more than other members of its genus, has reduced chaetotaxy (essentially only 1 row of bristles on abdominal terga and those very slender) ; long slender legs, with metatarsus I particularly long; the pleural arch so reduced as to be essentially absent; lateral metanotal area small, etc. Remark- able parallel developments between this flea and those of fleas of Nearctic flying- squirrels (and their hosts too) are mentioned later. Two other pygiopsyllids which are likely candidates, despite the presence of a pleural arch, are the monotypic Lycopsylla Rothschild, 1904 (Australia and Tasmania, on wombats) and Notiopsylla Jordan & Rothschild, 1914 (2 species, I ex procellariiform sea-birds; the other apparently from a parakeet that spends most of its time on the ground; both on sub- antarctic islands). Both of these genera are combless, a highly unusual condition amongst pygiopsyllids, and the lateral metanotal area is narrow. However, Notiopsylla is clothed with many fine hairs, suggesting it may spend some time on the host as well as in the burrows. The scanty records indicate this is so for NV. kergue- 326 R. TRAUB lensis (Taschenberg, 1880), but N. enciari Smit, 1957 is apparently known only from a very few specimens and hence data are lacking. Lycopsylla shows marked reduction of chaetotaxy (both in numbers and size of bristles), but the pleural arch is better developed than in known nest-fleas. As will be shown, the size of the pleural arch is correlated with jumping ability, and since fleas of large mammals tend to be excellent jumpers (e.g. Clenocephalides Stiles & Collins, 1930), and the wombat grows to a size of about 1200 mm in length and about 800 mm in height (more than 20 inches tall), it is not surprising that this flea does not show much reduction of the thorax 4. CERATOPHYLLID NeEsT-FLEAs. Nest-fleas occur in the Ceratophyllidae too, although they are of much less frequent occurrence than in the Hystochopsyllidae. Among the classic examples are a relatively few species of the large genus Cerato- phyllus Curtis, 1832, such as the Nearctic C. arcuegens° (fig. 90), and the Palaearctic C. rusticus® Wagner, 1903, C. caliotes° Jordan, 1937 and C. orites° Jordan, 1937, in which there is reduced chaetotaxy, loss of the pleural arch, development of slender legs, etc. (Holland, 1952; M. Rothschild, 1969). C. delichont° Nordberg, 1935, also belongs in this category, as can be seen from Smit’s excellent re-description (1956). These fleas parasitize martins and cliff swallows, but there are also other species of Ceratophyllus that infest these particular birds, e.g. C. styx (fig. 91), and yet which lack most or all the morphological modifications of the C. aycuegens-group, as can be seen by comparing figs 90 and gt. This is discussed below, when it is pointed out that these bird-nest fleas have well developed eyes, unlike the case in fleas of mammal- nests. Perhaps the most remarkable bird-flea is the Antarctic Glaciopsyllus antarcticus® Smit & Dunnet, 1962, which lives in the nest of the snow-petrel, often under many feet of snow (Murray et al., 1967). This flea is so specialized that it has not only lost the pleural arch but the pronotal comb as well (and is thereby unique in the family), and shows the reduced chaetotaxy, slender legs and other nest-flea features, but here again the eye is large. Amongst ceratophyllids of mammals that seem to be nest- fleas, may be mentioned Syngenopsyllus° Traub, 1950 (Indo-Malaysian), Libyastus° Jordan, 1936 (African), Tarsopsylla® Wagner, 1927 (Holarctic), all of which are fleas of tree-squirrels (or their nests) and Hollandipsylla° Traub, 1953, from a Bornean flying-squirrel. I believe that to a considerable extent the following ceratophyllid fleas spend much of their time in the nest rather than on the host, viz., the 2 species of Opisodasys which parasitize flying-squirrels, O. pseudarctomys° and O. vesperalis® (but not the rest of the species of Opisodasys, which infest tree-squirrels and Peromyscus) ; the dormouse-flea, Myoxopsylla° Wagner, 1927; the Mexican Pleochaetis paramundus, which is a Neotomodon-flea (but not the remaining 22 species of Pleochaetis, which infest primarily Peromyscus); and the Palaearctic marmot-flea Callopsylla dolabris (Jordan & Rothschild, rg1r). The ceratophyllid nest-fleas, especially those of mammals, are not as modified morphologically as are the bulk of the hystrichopsyllids, etc. However, all of those listed, save P. paramundus and C. dolabris, lack the pleural arch, but in the last 2, the arch is smaller than in their relatives, and the latter especially show other thoracic CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 327 changes, such as a narrowing and deepening of the lateral metanotal area. All tend to have reduced eyes and have fewer and thinner bristles; and more slender and elongate mouthparts and legs; and a narrower metepimere (except for P. paramundus), than do their allies which infest the fur of their hosts. The more specialized thoracic differences may be seen by comparing fig. 87 (O. hollandt, a fur-flea of diurnal tree- squirrels) and fig. 88 (O. vesperalis°). The maximum degree of relevant differentia- tion in the family may be seen by contrasting the head and thorax of C. arcuegens° (fig. 90) with that of C. styx (fig. gt). There is an odd species ascribed to the Cerato- phyllidae which may prove to be a nest-flea, namely Brevictenidia mikulini (Schwarz, 1960), the sole representative of the genus (and which I have not seen). This is characterized by a pronotal comb of “‘vestigial’’ widely-spaced, acuminate spines which are less than } the length of the pronotum. The elongate palpi, the very small eye, and reduced chaetotaxy shown in the published figures (Liu & Li, 1965) parallel the modifications indicated above. There is some doubt about whether this really is a ceratophyllid or leptopsyllid. 5. LeptopsyLLip NEst-FLEas. There are few nest-fleas known in the Lepto- psyllidae, but these exhibit the same tendencies. Thus, Brachyctenonotus® Wagner, 1928, a monotypic genus parasitizing mole-rats in south central Asia, not only lacks the pleural arch and has a very reduced eye, but its pronotal comb consists of extremely short and broad spines. Allied to Brachyctenonotus°® is the monotypic genus Calceopsylla° Liu, Wu & Wang, 1965, which judging from the figures and English summary of the original description (Liu, Wu & Wang, 1965), is a typical nest-flea since the eye is absent, the palpi elongate, the pronotal comb markedly reduced, the pleural arch missing, the abdominal spinelets absent, etc. The spines of the comb are narrow, very short and widely spaced, as in the former genus. Phaenopsylla Jordan, 1944, Hopkinsipsylla Traub, 1963 and Paradoxopsyllus microphthalmus off, 1946, although possessing the pleural arch, conform to the picture that is now familiar through repetition. An instructive stage in the development towards the nest-flea habitus is seen in Amphipsylla marikovskii loft & Tiflov, 1939. In ordinary species of Amphipsylla Wagner, 1909, the eye is quite small; there are about 20 spines (total) in the pronotal comb, the pleural arch is well developed, the lateral metanotal area is relatively broad, and the false combs of bristles on the pro- and metatibiae are generally prominent. In A. montana Argyropulo, 1946, for example, it may be noted that spines numbers 2—4 of the pronotal comb (fig. 105) are fairly straight and number 3 is more than 5 times as long as broad; the tanned margins of the pleural arch (PL.A.) extends well below the apex of the pleural rod (PL.R.) ; the gap between the arch and the tip of the rod exceeds the girth of the dorsal part of the rod; the lateral metanotal area (L.M.) is only about 1-3 times as long as tall and scarcely extends beyond the pleural rod; there is a false comb of 8 short, stiff, straight, somewhat spiniform bristles on the protibia (fig. 103) and one of about 6-8 such modified bristles on the metatibia (fig. 104). In contrast, in A. marikovskii, the eye is almost vestigial; there are only about 16 spines in the pronotal comb (fig. 100), spines numbers 3-5 are dorsally concave and relatively broad, e.g. number 3 is about 3-8 times as long as broad at the 328 R. TRAUB base; the pleural arch (PL.A.) is greatly reduced, viz., it is very small and compact, scarcely broader than the apex of the pleural rod (PL.R.) and almost contiguous with it; the lateral metanotal area is somewhat narrowed, viz., about 1-5 times as long and extends well caudad of the rod; the false comb on the protibia (fig. 101) consists of only 4 straight bristles and that of the metatibia (fig. 102) of but 3 bristles and these are inclined more ventrad than in unmodified members of the genus. An earlier step in this evolutionary trend is illustrated by fig. 94, showing the head and thorax of Ophthalmopsylla (Evemedosa) celata Traub, 1965 compared with that of O. jettmart Jordan, 1929, which is not a nest-flea (fig. 95). It is apparent that in O. celata, the eye is greatly reduced, the bristles are far fewer in number and are signifi- cantly more slender; the lateral metanotal area (L.M.) and metepimere (MTM.) narrower, the pleural arch (PL.A.) smaller and narrower, etc. There are fewer spines in the pronotal comb, but here O. celata has not progressed as far as Phaenopsylla (vide fig. 80), which has also become more specialized in other respects, viz., the eye is vestigial. 6. Puricip NEsT-FLEAs. Nest-fleas are rare in the Pulicidae, and it seems highly significant that the only such mammal-flea known to me, Xenopsylla papuensis°, a parasite of Pogonomys in New Guinea, conforms closely to the pattern and may be unique amongst the 70 described species in the genus in lacking a true pleural arch. Actually, the pleural arch is present in vestigial form (PL.A.(?)), arising from the ventrocaudal angle of the lateral metanotal area (L.M.), but it is extremely short and narrow, and is virtually contiguous with the top of the pleural rod, especially in the male. Like all Xenopsylla Glinkiewicz, 1907, it is combless, but as can be seen by comparing fig. 92 with fig. 93 (a New Guinean specimen of the related X. vexabilis, a fur-parasite of rats), the eye is absent in the former species, instead of being quite well developed; there are fewer bristles on the head and on the metepimere (MTM.), e.g. 2-4 instead of 5-5; the bristles in general are distinctly narrower; and MTM. is relatively narrow. In the California pulicid flea of sea-birds, Actenopsylla suavis Jordan & Rothschild, 1923, there is a very small pleural arch and the lateral metanotal area is narrow, recalling the condition in some other fleas of sea-birds, and the accompanying reduc- tion of the eye (fig. 99) and the chaetotaxy suggest that the flea spends much of its time in the nest-habitat of its host (Ptychorhamphus), which is in burrows in the ground. It seems significant that this flea, and its ally, Ornithopsylla laetitiae Rothschild, 1908, the only bird-fleas in the group, are likewise the only members of the tribe lacking genal and pronotal ctenidia. The latter is also a flea of sea-birds (Puffinus), and has been found in their nests as well as on the bodies of the hosts, and is modified along somewhat similar morphological lines as Actenopsylla Jordan & Rothschild, 1923. However, the straight frontal margin of the head of Ornithopsylla Rothschild, 1g08 and the somewhat larger eye, lead me to believe that it is not as much of a nest-inhabitant as the former. 7. CHIMAEROPSYLLID NEST-FLEAS. The small family Chimaeropsyllidae is limi- ted to the Ethiopian Region and primarily to South Africa. The majority of the CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 329 species of the largest genus, Chzastopsylla Rothschild, rgr1o (x1.2) are definitely stated to be nest-fleas, e.g. all members of the nwmae-group and especially C. nwmae (Roths- child, 1904) and C. quadrisetis de Meillon, 1930, and at least 5 of the 8 species in the mulleri-group, namely those found in the nests cf Otomys unisulcatus, the Karoo rat (de Meillon, Davis & Hardy, 1961; Haeselbarth, 1966). Chzastopsylla reminds one of Anomiopsyllus° because of the diminutive size of the species and the marked reduction in chaetotaxy. However, unlike the latter, Chiastopsylla does have combs, both genal and pronotal. Significantly, though, the pronotal comb is greatly reduced and usually consists of only 5~7 short spines per side, and the genal comb at most consists of 2 spines, and at times only 1, and may be missing altogether, as in C. prtchfordi Ingram, 1927. Such variability of numbers of spines suggest the structures are in the process of elimination in the course of evolution. The maxillary palpi are relatively long and thin, and in C. pitchfordi extend to near the apex of the trochan- ters, and the eye is vestigial. The pleural arch is present, but is very small. The known ultimate in this line of development in the family is in the monotypic genus Praopsylla Ingram, 1927, which lacks combs entirely, and in which the eye is so reduced that it is represented by a mere small tanned patch. The head has an unusual subconical appearance at the frontal angle due to the anterior excision of the ventral margin, a configuration that can be seen in someother nest-fleas, e.g. Eopsylla° (i.2) and Conorhinopsylla® (i.2), (fig. 86) while the mouthparts are elongate, extending beyond the apex of the fore trochanters, the pleural arch is very small, and there is but a single row of bristles, and even the bristles on the hind tarsi are unusually short. Superficially Cryptopsylla de Meillon, 1949 resembles these nest-fleas, but considera- tion of this is best deferred. 8. RHOPALOPSYLLID NEST-FLEAS. Members of the family Rhopalopsyllidae are essentially Neotropical rodent-fleas, with a few exceptional members occurring in the subantarctic region or in southern U.S. and with some infesting birds or other hosts. Many new species, and perhaps undescribed genera, undoubtedly await discovery, for most of South America is still unexplored concerning fleas. None of the species bear true ctenidia (Traub, 1968) and one cannot discuss such modifications when considering nest-fleas. However, a number of forms exhibit reduced chaetotaxy (regarding numbers of rows, diameters), lengthened palpi and an unusually large (and variable) number of palpal segments, e.g. Ectinorus polymerus Jordan, 1942 and Dysmicus barrerai (Jordan, 1939). It therefore seems significant that Jordan (1942) cites these as being found in the nests of their hosts (Octomys), or, in the case of the latter species, flourishing there. The features of other related taxa suggest they too may be nest-inhabitants to a degree, and for example, de Meillon (1952) states that ‘“Parapsyllus .. . is essentially a nest-flea’’, but on the whole too little is known about the fauna, and their habits, for further discussion. 9. DIFFERENCES IN DEGREE OF THE NEsT-FLEA Habitus. It is therefore clear that the development of the nest-flea habitus and habit has occurred independently many times within the Order, viz., in at least 7 families, and in 5 subfamilies of one of these, the hystrichopsyllids. However, it is emphasized that the degree of con- 330 R. TRAUB vergence or differentiation varies. Jordanopsylla lacks combs but has a pleural arch; some have a short comb but no pleural arch; neopsyllids are quite small in size while the squirrel-fleas are of fair size, etc. It seems that some species, like P. pavamundus (fig. 68), are relatively late entries in this line of development, and still may be found in fair number on the bodies of the host, and these tend to show a lesser degree of specialization by reduction, e.g. the eye is definitely smaller in P. paramundus than in other Pleochaetis, but it is not vestigial, and the pronotal spines are short as compared with the pronotum and are relatively quite broad, but the comb is not as modified as in Strepsylla (fig. 79) and Epitedia (fig. 77), etc. Other species of genera of nest-fleas have been extremely difficult to collect unless and until one has the opportunity to examine the nests of the hosts, as was the case with Hopkinsipsylla (Traub, 1963, 1965). This has also been the experience with Tamio- phila, Phalacropsylla etc., while others have remained “‘rare’’ despite intensive collec- tion of mammals in the endemic habitat, presumably because it has not been possible to seek or find the nest of the host, as happened with Jordanopsylla and the subgenus Evemedosa. Certain genera of Siphonaptera possess some of the attributes of nest-fleas but have other characteristics which suggest that these fleas are not merely specialized to crawl about in the nest and feed on sleeping hosts. It appears that some of these species are progressing towards the nest-habit, while others may be evolving away from it, and these are herein referred to as partial nest-fleas. Such evolutionary manifestations are not surprising, since it seems logical to believe that the Order arose from a non-parasitic ancestor which lived in the nests of mammals or proto- mammals, feeding on organic debris, and later, perhaps, on scurf on the bodies of the residents, and gradually acquiring the facility to feed on the blood of what were by then their hosts. Development of the ectoparasitic habit was accompanied by the adaptations of making it more and more successful, e.g. flattening of the body, presence of rows of stiff backward projecting bristles or combs of spines; develop- ment of jumping legs (N. C. Rothschild, 1917; loff, 1929; M. Rothschild & Clay, 1952; Traub, 1966); modification of the flight mechanism into a device for leaping (Neville & M. Rothschild, 1967; M. Rothschild, 1969). It is believed that most (but not all) of the early fleas possessed many combs of spines and that most of these ctenidia have become vestigial or lost (at times with scarcely any trace) as the fleas developed other means of remaining safely on the host despite its exertions in running, or flying, or special efforts to destroy the flea by teeth, beak or claw (Traub & Barrera, 1966; Traub & Evans, 1967; Traub, 1968, 1969). Some of these new mechanisms include “false combs”’ of bristles to replace the true combs which had been lost (Traub, 1968). It therefore appears that the “specialization by simplification” (Hopkins & Rothschild, 1962) of nest-fleas is indeed just that, a series of modifications enabling the flea to return to a nest-existence, but in a manner superior to that of its forebears. However, in some instances, superimposed upon the model characterized by reduction of the combs and bristles, etc., are later or concurrent adaptations which enable the flea to remain on the host when it is active in the nest or burrow, or even outside, while running, leaping, gliding or flying. | CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 331 An example of a taxon of fleas with such a “dual-purpose” set of modifications is the anomiopsylline hystrichopsyllid Stenistomera°, known from 3 species in the Nearctic deserts. Here the eye is vestigial and marked modification of the thorax accompanied the loss of the pleural arch (i.e. absence of the lateral metanotal area) and there is reduction in abdominal chaetotaxy (but one row of tergal and sternal bristles, and with intercalaries vestigial), while the legs are long and slender. How- ever, this picture of a typical nest-flea is confused by features illustrating another evolutionary trend, which is exhibited in non-nest fleas representing such diverse taxa as some (but not all) species of Jellisonia Traub, 1944 and Kohlsia, which are ceratophyllids, and by the leptosyllids Amphipsylla, Caenopsylla Rothschild, 1909, Ctenophyllus Wagner, 1927, and Peromyscopsylla I. Fox, 1939, and by pygiopsyllids like certain Metastivalius Holland, 1969. These are the combination of: (1) A narrowed frontal region on the head (i.e. it is taller than long and often bullet- shaped); (2) A frontal margin that is somewhat flattened for the ventral half. (3) A row of frontal bristles bordering the anterior margin. (4) Tendency for bristles of the first row to be thickened near the base, or, somewhat spiniform. (5) Presence of a “false comb’, i.e. a vertical row of single, short or stiff bristles, on the tibiae. (6) A full pronotal comb of many, narrow spines, with the lowest spine (or additional ones) extending over the vinculum. Modifications of the frontal margin of the head in this manner is deemed by me to be characteristic of fleas which become temporarily rather sedentary and cling to the host by hooking the marginal row of subspiniforms or thickened bristles onto the hairs while the flea abuts its head against the skin (Traub, 1968, 1969). It is noteworthy that in Stenistomera® only S. alpina® (Baker, 1895) exhibits all of these “secondary” features. In S. macrodactyla° Good, 1942, the first row of frontal bristles is not so marginal nor subspiniform; the tibial combs not so well developed, etc., while S. hubbardi° Egoscue, 1968 is intermediate. It would seem that S. alpina®° is adapted both to feed on the host while it is in the nest and to cling successfully to the fur while the Neotoma is outside, where a fall from the host might mean death for a nest-flea, as stressed below (Section IV.C.2). In the category of a flea apparently following two lines of evolution simultaneously, viz. specialization by reduction complemented by elaboration of existing structures, is the South African monotypic genus Cryptopsylla (xi.2). It superficially resembles the related Praopsylla, which presumably is a true nest-flea, in that there is no sign of a comb, but the eye is even more reduced—it is completely absent. However, there are significant differences in that the mouthparts are much shorter, there is a false comb of modified bristles on all the tibiae, and while there is but one row of abdominal bristles, these are unusually long. This is a flea of a burrowing rodent with mole-like habits, the bathyergid Cryptomys, and I believe the loss of the eye and development of unusually long bristles is characteristic of fleas of such hosts (i.e. Dactylopsylla Jordan, 1929 (iii.2) of Nearctic pocket-gophers; Dinopsyllus (Crypto- ctenopsyllus) ingens (Rothschild, tgoo) (i.4), a flea of the mole-rat Bathyergus). Such a coat of long thin bristles (coupled with marked reduction of the eye) is also seen in the 4 genera of fleas parasitizing Aplodontia, a North American rodent which spends much of its time burrowing. It seems likely that vestiture of this type not 332 R. TRAUB only may help the flea stay on its host while the latter brushes against the walls of its tunnels, and that the close proximity, great length and overlapping condition of the bristles also offer protection against the grains of soil that would otherwise dust and adhere to the flea in such an environment. Since Cryptopsylla not only has chaetotaxy of this pattern, but also is adorned with tibial combs, I believe it spends time on its host as well as in the nest. Other examples of fleas which have some of the attributes of nest-fleas, but not many, and which have been collected in rodent nests are Xiphiopsylla Jordan & Rothschild, 1913 and Coptopsylla Jordan & Rothschild, 1908. Xziphiopsylla, a genus infesting primarily rodents in the Ethiopian Region, is sufficiently distinctive to be placed in a family by itself, and the features relevant to this discussion include a vestigial eye, long thin maxillary palpi, a short pronotal comb of short spines (and hence pronotum greatly exceeding length of spines) ; a reduced pleural arch (virtually vestigial in X. hyparetes Jordan & Rothschild, 1913) ; a narrow metepimere and fairly slender legs. However, there are 2 or more full rows of abdominal bristles on the terga, and the long bristles are remarkable in being sword-shaped, i.e. virtually straight from base to near apex instead of tapering in the usual way. The body surface is heavily sclerotized, reticulated, sculptured or microspiculose, and there often are dorsal incrassations on the terga. The body ornamentation and the densely tanned nota may be regarded as adaptations for protection against being crushed or caught by the host, but likewise would be of value when squeezing into crevices in the nest of the host, and the stiff bristles and reticulated surfaces would facilitate crawling into tight cracks and offer resistance to being pulled out therefrom. Similar incrassations are seen in some hystrichopsyllids like Xenodaeria Jordan, 1932 (Doratopsyllinae), Rothschildiana (Neopsyllinae), the rhadinopsylline Stenischia® Jordan, 1932 and Corypsylla° C. Fox, 1908, and are examples of convergence. Rothschildiana also bears comblike bristles on the sterna, and the body is rigid, with the surface spiculose. These specializations suggest it likewise hides in narrow apertures in the nest. Coptopsylla likewise is the sole representative of a special family, and is a desert flea in the southwestern subregion of the Palaearctic Region. It is strongly remini- scent of true nest-fleas in the absence of any combs, in the extreme length of the palpi and stylets; the tall, narrow lateral metanotal area, and the presence of but a single row of abdominal bristles. However, there is a well developed pleural arch, a large eye and the bristles are long, and these indicate a flea that spends at least part of the time on an active host, and as we shall see, is a good jumper and is active in the daylight. Additional points about fleas which might be nest-fleas to a degree will be deferred until a discussion of the absence of the pleural arch and the reduction of the eyes in Siphonaptera. Inasmuch as so many of the nest-fleas have been indicated as having lost the pleural arch and possessing reduced or vestigial eyes, these points are discussed below, since other types of fleas are so modified as well. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 333 C. THE PLEURAL ARCH OF THE METATHORAX I. REVIEW OF TAXA LACKING A PLEURAL ARCH. The absence of the pleural arch of the metathorax in certain fleas (e.g. figs 85, 86, 90, 92) is of interest and significance and constitutes another good example of Convergent Evolution in fleas, since the condition occurs in a fairly large number of genera which are often unrelated, while close allies have normal pleural arches (figs 87, 91, 93, PL.A.) (Traub, 1953b). At times, only one or two members of a large genus are affected, e.g. Xenopsylla, and loss or marked reduction of this structure is known in at least 6 families (Hystrichopsylli- dae, Pygiopsyllidae, Ceratophyllidae, Leptopsyllidae, Ischnopsyllidae and Pulicidae) although it is quite rare in the pygiopsyllids, leptopsyllids and pulicids. Further, fleas with this trait are found in a variety of habitats in widely separated geographic areas. Ceratophyllid squirrel-fleas with the thorax modified in this manner were listed by Traub (1950, 1953), viz.: Tarsopsylla°9 (Holarctic), Syngenopsyllus° (Indo- Malaysian), Libyastus° (Ethiopian), the 2 species of Opisodasys Jordan, 1933 which parasitize Nearctic flying-squirrels, viz. O. pseudarctomys°® (fig. 47) and O. vesperalis® (fig. 88) and the monotypic Bornean flying-squirrel flea Hollandipsylla°. A small minority of ceratophyllid bird-fleas lack the pleural arch (Holland, 1952; M. Roths- child, 1969) i.e. Ceratophyllus arcuegens°® (fig. 90) and the 3 other martin and bank- swallow fleas mentioned above, as well as the notable Antarctic flea, Glaciopsyllus antarcticus°. The arch is likewise missing in the ceratophyllid dormouse-fleas Myoxopsylla° and in the leptopsyllids Brachyctenonotus° and Calceopsylla°. A lepto- psyllid in which the arch is greatly reduced but still discernable is Amphipsylla marikovskit. Among hystrichopsyllids, most anomiopsyllines (e.g. Anomiopsyllus°, Megarthroglossus°, Conorhinopsylla°, Callistopsyllus°, Stenistomera® and Eopsylla°) lack the pleural arch (Traub & Tipton, 1951) as do nearly all rhadinopsyllines (e.g. Rhadinopsylla°, Corypsylla°, Trichopsylloides° Ewing, 1938, Paratyphloceras® Ewing, 1940, Stenischia° and Wenzella°). When present in this subfamily (Nearctopsylla Rothschild, 1915), it is very small and the associated pleural rod is abbreviated. In neopsyllines, the pleural arch is definitely reduced in Phalacropsylla, Genoneopsylla, Delotelis, Epitedia faceta (Rothschild, 1915), Tamiophila and somewhat less as in Strepsylla, Meringis, Catallagia and Paraneopsylla as compared to Neopsylla. Amongst ctenophthalmines, the arch is absent in Agastopsylla°. Among the dora- topsyllines, [dilla°, a monotypic genus known from dasyurine and peramelid marsu- pials, is similarly modified. The arch is absent in many ischnopsyllids (bat-fleas) i.e. Thawmapsylla° Rothschild, 1907, Lagaropsylla° Jordan & Rothschild, 1921, Avaeopsylla° Jordan & Rothschild, 1g21, Nycteridopsylla® Oudemans, 1906 (fig. 89), etc. and is greatly reduced in Ischnopsyllus Westwood, 1833, Oxyparius Jordan, 1936 and others. Amongst pulicids, X. papuensis° has a thorax essentially of this type (fig. 92), thereby differing from its many allies (e.g. fig. 93). This structure is much reduced in Rhynchopsyllus Haller, 1880 and Hectopsylla Frauenfeld, 1860, which are sticktight fleas infesting birds or bats, and is also very small in male Tunga Jarocki, 1838 and virtually absent infemale Tunga. These particular sticktight and burrowing fleas are at times placed ® It is reiterated that the species which are marked ° in this article, lack the pleural arch. 334 R. TRAUB in the family Tungidae but are regarded by some as a subfamily of Pulicidae. Choristopsylla tristis® is one of the few pygiopsyllids known to lack the pleural arch. Another is Pygiopsylla celebensis° Ewing, 1924, represented by but one specimen extant and with unknown affinities and host relationships. Although the above review does not purport to be complete, it is clear that reduc- tion of the thorax and loss of the pleural arch has occurred repeatedly and often sporadically throughout the Order, and without much regard for taxonomy. The important question is, what is the significance of this evolutionary development? Unless one wants to believe that an Order may have a built-in mechanism for the haphazard appearance of a single-type mutation or atavism, there ought to be a common factor or basis for such repeated but natural selection of an unusual trait. Moreover, the first possibility is ruled out since the loss of the pleural arch is accom- plished in several different ways, as will be shown. In short, this trait should be adaptive or at least fit with known evolutionary principles, and in what follows an attempt will be made to demonstrate that this is indeed the case. 2. THE ADAPTIVE VALUE OF THE Loss OF THE PLEURAL ARCH. A review of the list of fleas with this type of thorax, made in an attempt to determine the possible value of the modification, immediately leads to some generalizations. One is that virtually all have reduced eyes. However, the relatively few exceptions have unusually large eyes, so apparently that phenomenon is complex, and hence it will be treated later, separately, in Section IV.D.1. It is also evident that most of the names are familiar in that they were mentioned above as nest-fleas. Nevertheless, there is one glaring addition—the bat-fleas, and these provide a clue as to the function of the reduced arch, as shown below. Various workers were impressed by the high incidence of the reduction of loss of the pleural arch in nest-fleas, and postulated that it was associated with the lessening or loss of the ability to jump well, or with sedentary habits (Traub, 1953). Apparent- ly, the first direct statement to this effect was made by Holland (1952) when describ- ing his new bird-flea, Ceratophyllus arcuegens°. He noted that, like 3 Palaearctic members of the genus (out of the many Ceratophyllus known), this species lacked a pleural arch (fig. go), and went on to state that because of the reduction of the legs and thorax, and as indicated by the “rather large and deep abdomen,” this was “evidently a rather sedentary species... ; a crawler rather than a jumper.” Earlier, without specifically mentioning the loss of the pleural arch, Holland (1949), in referring to Tarsopsylla°, stated that “‘in spite of its long legs, [it] appears to be a rather sluggish flea.” Smit (1953b) made similar observations regarding Tarsopsylla°, but again without reference to the arch, declaring “It seems to be a general rule in fleas that the ones with long and slender legs are walkers and climbers rather than jumpers, while those with stout and relatively short legs seem to be the jumping champions”. Johnson (1957) discussed various modifications of the lateral meta- notal area and the pleural arch, and stated she believed the latter was “mainly an aid in jumping”. Johnson also indicated that “prior loss of the pleural arch is somehow connected with incorporation of the lateral metanotal area into the notum or to its complete loss”. Smit & Dunnet (1962) expressed the view that G. antare- CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 335 ticus® was a crawling flea, a point that was verified later by field observations (Murray et al., 1967). Traub & Evans (1967) regarded the modifications of nest- fleas, including reduction in jumping ability, as adaptations for feeding on a quiescent or sleeping host, but the true significance of the loss of the pleural arch became clear only after the brilliant work of M. Rothschild and colleagues. The pleural arch of fleas was shown to be the site of a cap-shaped mass of resilin, ‘‘an elastic protein which stores and releases energy more efficiently than any known rubber, and which can deliver power faster than most efficiently contracting muscle.’’ (M. Rothschild, 1969.) The pleural arch of fleas is homologous with the hinge-ligament of the wing of flying insects, which is situated on the top of the pleural ridge of scorpion-flies, locusts, etc., and consists of resilin (Neville & Rothschild, 1967). The latter authors referred to fleas as ‘insects that fly with their legs’’ since the jump is effected by a modified flight-mechanism. Reduction or loss of the pleural arch, then, can only be associated with lessened ability to jump. What advantage from loss of the arch results to fleas like Megarthroglossus°, Stenistomera® or Conorhinopsylla° in a nest of Neotoma on the ground orinarock-ledge, or Callistopsyllus° in an underground nest of Peromyscus? As already indicated, these fleas are seldom found on rodents away from the nest, presumably not only because they are morphologically and physiologically adapted for feeding on the host while it is resting or sleeping, but because they are so ill-fitted to escape falling off an active host or being killed by it. It seems eminently logical to believe that they are not equipped to survive the external environment outside the nest, and probably neither are the eggs nor larvae. Further, most of the species cited are desert-species, and it probably is no coincidence that a large proportion of nest-fleas are species infesting mammals living in xeric areas. Bearing in mind that nest-fleas are essentially blind and keyed to living in the dark confines of the nest, it is clear that a nest-flea which was a good jumper might, in one leap, end up in an extremely hostile environment just a few inches away from the nest. Moreover, such a flea would be unlikely to find its way back. Darskaya (1955) noted that the desert-flea Xenopsylla gerbilli caspica loff, 1950 (ix.1) dies quickly if it misses in an attempt to attach to a passing host and fails to return to the burrow promptly. Since this is not a nest-flea, and is adapted to conditions at the burrow-entrance, it is no doubt hardier regarding environmental factors than a species attuned to the microclimate deep in the rodent nest. Another disadvantage to a nest-flea of being a good jumper would be the injuries sustained in leaping about blindly in closely confined quarters. A jumper would not be very likely to crawl within the relatively favourable runway of the host if it fell off the rodent, and instead would leap into an alien world. Thus, survival in the nest-flea would be most favoured in the poor jumper and natural selection would in time lead to the reduction and loss of the pleural arch so characteristic of nest-fleas. There is then, much more to the evolution of a thorax without a pleural arch than the mere “‘lack of need’’ for effective leaps in a species of flea living in underground nests. The adaptive value of this development is further indicated by the fact that the loss of the arch is commonly seen in nest-fleas inhabiting nests or roosts at high altitudes, | €.g. squirrel-nests on tall trees, bird-nests on cliffs or in banks, or in caves, or crannies | 336 R. TRAUB in a tall cave where bats cling, or in holes where bats roost. As Dr Miriam Rothschild has pointed out in the lecture summarized by Neville & M. Rothschild (1967) a leap of 10-12”, the type readily accomplished by a flea with a well developed pleural arch, could carry an aerial nest-flea 100 feet down and away from the host it is dependent upon for the blood-meal necessary for nourishment and to prime egg- laying. The selective survival value of the modification of the thorax in such a nest-flea is apparent, particularly if it is faced with the problem of locating and attaching to its host in the dark, as in the case of bat-fleas and flying-squirrel fleas. However, fleas of volant and gliding hosts, or mammals living far above the level of the ground face a dilemma, because coupled with the need for restricted mobility there is the exacerbated problem of safely remaining attached to the host at such altitude (especially if it is volant). It is therefore not surprising that such fleas, lacking a pleural arch, have not followed the evolutionary path of fleas of underground nests and effected reduction of the combs, but instead, on the contrary, have developed supernumerary spines or combs. This is discussed below, in Sections C.3 and C.4. The loss or reduction of the pleural arch and the resulting inability to jump well is also adaptive in other categories of fleas, e.g. the Antarctic flea G. antarcticus*. This flea lives in the nests of the silver-grey fulmar and the snow-petrel, amongst the rocks on the ground (for the former host) or in crevices (for the latter) (Murray et al., 1967). In the spring, the fulmar burrows to its former nest-site through the snow before it has melted, and the petrel also returns to its own individual nest-site. Incubation of eggs and rearing chicks takes 4 months, a period of warmth which is provided by the bird and which is obviously suitable, with respect to time and environment, for the life-cycle of the flea, because all stages have been found in the nest. An Antarctic flea of marine birds that was a good jumper could find itself in the sea, or exposed to extremely cold temperatures without a host. It is worth recalling that other fleas of marine birds have poorly developed pleural arches and are apparently poor jumpers Of special interest are the hystrichopsyllids which seem to be nest-fleas by virtue ~ of the reduction of the eye, thorax, and the number of bristles; loss of the pleural arch, etc. but which nevertheless have well developed ctenidia. In this group are insectivore-fleas like Corypsylla° (1.8) which possess a vertical comb near the frontal margin of the head which presumably hooks onto the hairs of the host as the flea abuts its head against the skin, as discussed by Traub (1968, 1969). The flattened frons and the vertical genal comb of the related Stenischia° suggest a similar habit. The question arises as to why such fleas, particularly Corypsylla°, whose ctenidia would seem to belie a true nest-existence, have lost the pleural arch and are such poor jumpers. In the first place, as is mentioned in Section IV.J below for Cory- psylla’, and has been noted above in IV.B.2, for Stenischia’, there is reason to believe that such ctenidia are literally reduced as compared to those of relatives, and that they thus represent evolutionary tendencies shown by nest-fleas in general. Secondly, it may be that the peculiar vestiture of the head is an adaptation to stay affixed to the fur of a host while it is asleep in its home, and that these fleas are semi- sessile for hours and are indeed nest-fleas. In this connection, it should be noted CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 337 that the “banded” abdomen of these fleas, with its incrassate and rigid margins alternating with a softer middle, coupled with marked reduction of obtrusive abdominal combs and bristles, is ideally fitted to enable the insect to squeeze into, and out of, tiny cracks in the nest without being hurt, just as the reinforced seg- mental margins compensate for the loss of protection offered by the bristles of ordinary fleas. (It is significant that nest-fleas of birds are also “‘banded”’ but the alternating tanned and relatively soft sections of the abdomen extend from front to rear.) It therefore appears that these rhadinopsyllines are nest-fleas, at least in part, but there seems to be a third factor in the loss of the pleural arch, and that is the adaptive benefit to fleas which spend much ‘of their time in narrow burrows, such as in the case of Corypsylla°, which infest moles and shrews. The question arises as to why such fleas have lost the pleural arch. It is, of course, possible for such fleas to utilize their specialized vestiture while feeding on a host that is in the nest asleep, and that development of this habit led to a semi-sessile existence and concomitant loss of the arch. However, other explanations for the Corypsylla-type of habitus seem more plausible. Fleas that jump well (e.g. Ctenocephalides) can leap heights of 8-10 inches (Traub, 1964), while moderately good jumpers like X. cheopis (Rothschild, 1903) may ascend to heights of 3-4 inches (Traub, 1964). Moreover, fleas usually make a series of leaps in succession. It is obvious that not only is there no need for such ability in fleas that may hatch out in, or frequent burrows that are only about 14 inches in height, but that the resulting forcible and repeated collision with the roof of the burrow could be disadvantageous to the species as well as to the individual. Shrews and other insectivores tend to establish runways above ground as well as below the surface, and limited ability to jump would restrict loose fleas to these particular sites, whereas otherwise they may end up in a micro- environment that is injurious or remote from any suitable host. Support for this last-mentioned hypothesis is provided by a new species of Stenischia° from the Himalayas of W. Pakistan. Very few specimens of any of the other species of this genus are known, and hence their true hosts are in doubt. However, the Pakistani flea is a parasite of Hyperacrius, a small burrowing vole that only very rarely emerges from the underground. It seems logical that such a flea would be a poor jumper, just as it is not surprising that the Palaeopsylla of this mole- like vole bears ctenidia resembling those of Palaeopsylla of moles rather than those of shrews (Traub & Evans, 1967). The presence or absence of a well developed pleural arch is also adaptive in the hypodermal, sticktight and semi-sessile fleas. Those, such as the pulicine Echidno- phaga and Spilopsyllus cuniculi (Dale, 1878), which have to seek and infest an active, large host, and which are sessile only part of their lives, have well developed pleural arches, and are thereby fit for the vital ‘‘great leap forward” onto their host. The tungine burrowing and sessile forms, e.g. Tunga, in contrast, presumably have very little difficulty in finding and infesting their hosts at ground-level and remain affixed for life and have greatly reduced pleural arches. In the tungines, the evolutionary path has been towards suppression of the locomotory parts of the body (and even much of the limbs become resorbed after attachment or penetration), while the feeding and reproductive ends are hyperdeveloped. Marked reduction of 338 R. TRAUB the entire thorax has therefore reduced the size of the target of the host bent on destroying the stationary flea and, in hypodermal species, has diminished the volume of body that must penetrate the skin of the host. In the case of Echidnophaga, etc., however, the advantage of being a good jumper outweighed the possible assets of loss of the pleural arch. 3. HyPERDEVELOPMENT OF COMBS IN BiRD-FLEAS, BAT-FLEAS, ETC. LACKING THE PLeuRAL ArcH. It has been stressed above that the loss of the pleural arch is apparently adaptive, whether in fleas of bats, birds, flying-squirrels, squirrels nesting in tall trees, or of hosts nesting underground or on the surface. It has also been noted that instead of showing a tendency towards the reduction of combs exhibited by fleas of underground nests, the fleas of bats, birds, flying-squirrels (and of nocturnal arboreal animals in general) display, on the contrary, a trend towards a significantly greater number of spines in the pronotal comb, and, in the bat-fleas, development of a series of true combs (vide fig. 98) or of false combs (Traub, 1968). These modifica- tions likewise can be explained in terms of adaptations. As in the case of the bird- fleas in burrows on the faces of cliffs, and the squirrel-fleas in nests atop huge trees, bat-fleas face the hazards associated with heights, and substituting the facility to crawl and climb, for the ability to leap is advantageous to the individual and to the species. This is true especially in the case of eyeless bat-fleas and other fleas of nocturnal hosts, which would have less chance of locating their prey by a blind leap in the dark. However, once onanactive, agile, volant, gliding orleaping host, these fleas must solve the severe problem of maintaining their hold, regardless of whether the bird ormammal is perched, moving or trying to kill or remove them. A fall from such a height might not be fatal, or even injurious to a flea, but there would be little chance of finding a new and suitable volant or arboreal host on the ground, even if it were day-time and the flea could see. It is therefore to be expected that bird-fleas and bat-fleas have “supernumerary”’ spines in the pronotal comb as a result of natural selection (note figs. 89-91, and compare the bird-Lentistivalius, fig. 18 with the murid-one, fig. 16), and that many bat-fleas have fairly well-developed abdominal combs, or even ctenidia of false spines on the metathorax, or dorsal stiff, spine-like bristles forming a series of comb-like projections (Traub, 1968). Humphries (1966) independently came to the same conclusions about the adaptive value of the combs of bat-fleas. This same principle, it seems to me, also applies to the nest-fleas of arboreal squirrels, and this is why, although they lack the pleural arch, Tarsopsylla° (fig. 8), Syngeno- psyllus® (fig. 3), Libyastus® (fig. 13) (all ceratophyllids) bear a full complement of spines, instead of having a reduced comb. Although precise data on the habits of some of these squirrel-fleas are lacking, the scarcity of records on the host and their demonstrated presence in nests lead me to believe that the last two are nest-fleas and/or infest nocturnal or crepuscular hosts, as mentioned below (p. 340). In the case of Tarsopsylla° (ii.1), Holland (1949), Smit (1953b) and Sakaguti & Jameson (1962) all cited it as a probable nest-flea, and its long, thin legs are also highly suggestive ofa flea that crawls rather thanleaps. It is noteworthy that the pronotal spines of these particular squirrel-fleas are somewhat narrower and relatively longer CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 339 than in the active fur-fleas of diurnal arboreal squirrels like some Orchopeas (figs 29, 31), Opisodasys (figs 46, 48) and Monopsyllus (figs 62, 64). This is presumably an additional step in the line of development, leading toward the adaptations of fleas of the flying-squirrels next discussed. 4. HyPERDEVELOPMENT OF THE PRONOTAL COMB IN FLYING-SQUIRREL FLEAS, ETc. As indicated above, the fleas of flying-squirrels tend to have a greater number of spines in the pronotal comb than do other members of the same genus which infest diurnal, arboreal squirrels or other hosts, and the spines are narrower and relatively longer, e.g. the Nearctic Opisodasys (111.1) of flying-squirrels (fig. 47,0. pseudarctomys° and fig. 88, O. vesperalis°) versus Opisodasys from diurnal tree-squirrels (figs 44, 87, O. hollandi and fig. 48, O. enoplus (Rothschild, 1909)). The Bornean flying-squirrel flea, Hollandipsylla neali° Traub, 1953 (ii.1) also has a comb of this type (about 22 narrow spines). These particular flying-squirrel fleas also lack the pleural arch, unlike their allies, while in another (Himalayan) ceratophyllid of flying-squirrels, Macrostylophora fimbriata (Jordan & Rothschild, 1921) (fig. 97), the pleural arch is reduced as compared to allies on ground-squirrels, e.g. M. borneensis (fig. 96) and the pronotal comb likewise has a fuller complement than that seen in fleas of diurnal tree-squirrels (cf. fig. 7 versus fig. 14). A similar condition can be seen in comparing the ctenidium of a Monopsyllus from flying-squirrels (fig. 58) (i1.1) with one from ordinary tree-squirrels (figs 62,64). The pleural arch of M. argus (Rothschild, 1908), from flying-squirrels, is also significantly shorter and narrower than in the Mono- psyllus from tree-squirrels like M. vison (Baker, 1904). The Mexican Orchopeas bolivari Barrera, 1955 (New Combination) (iii.1) is believed to be a flea of flying- squirrels, and also shows this tendency (especially in the female), but to a lesser degree, perhaps because it is a recent association. The same trend is seen in Myoxopsylla°, which are parasites of dormice, and al- though there are no known members of the genus occurring on other hosts which could be used for comparison, the comb extends more ventrad and has more spines than in related taxa, and the spines are also narrower. In Monopsyllus sciurorum (Schrank, 1803), which, despite its name, is a dormouse-flea, this same tendency is evident regarding a fuller comb and spines that are narrower in breadth and greater in number, as can be seen by comparing fig. 60 with the Monopsyllus of tree-squirrels shown in figs 62 and 64. It has been pointed out (Traub, 1969) that the fleas of nocturnal arboreal animals characteristically have a relatively large number of spines and that the comb descends sufficiently ventrad over the pronotum to reach or extend below the level of the third vinculum, while the lower spines are oblique in inclination, e.g. Muesebeckella Traub, 1969 (ii.1), Acanthopsylla Jordan & Rothschild, 1922 (ii.1), Pleochaetis ponsi Barrera, 1955 (iii.1), Kohlsia keenani Tipton & Mendez, 1961 (iii.t). Tarsopsylla coloradensis® (Baker, 1895) has a comb of this type, and from the figures cited, it can be seen that the comb of the Opisodasys of flying-squirrels and the other fleas of such crepuscular hosts approach this condition. In the case of fleas of arboreal, nocturnal hosts, as with bat-fleas and fleas of nests atop tall trees, the pressures of natural selection would aid fleas that were crawlers, not good jumpers (hence the loss or reduction of the pleural arch), and/or which had c 340 R. TRAUB an elaborate pronotal comb that would help them latch onto and retain a hold on a gliding or highly active host operating in the night, usually at heights that would offer little chance of survival for a flea that was dislodged. It therefore is not sur- prising that the end-result in such fleas is a pronotal comb which superficially resem- bles that of bird-fleas because of ‘‘supernumerary’’ spines, but which differs in essential details as noted above and elsewhere (Traub, 1969). Of special interest and relevance in this discussion is the remarkable monotypic genus Scolopsyllus Méndez, 1968, the only member of the family Rhopalopsyllidae having a pronotal comb. The enormously significant feature about this unique pronotal comb is that it consists wholly of false spines, paralleling the condition of the “false helmet” on the head of Smitella Traub, 1968. In these instances the spine-like structures arise from definite setal bases, instead of directly from the integument as in the case of true spines, and similar false combs have been noted as occurring on other parts of the body in other taxa (Traub, 1968). That author, writing before the discovery of Scolopsyllus, pointed out that such false combs function as regular ctenidia but suggested that they are not substitute mechanisms for true combs which have been lost in the course of evolution, since they occur on parts of the body which apparently have never borne true combs in the forebears of those particular taxa. Accordingly, he predicted that a rhopalopsyllid might be found with a well developed comb of modified bristles on the pronotum, and went on to state that “if a pronotal comb of specialized bristles were found to occur, it would be in a flea secondarily adapted to bank- or tree-nesting birds or else in a parasite of a nocturnal arboreal mammal.’’ Now that a rhopalopsyllid has been discovered with such a ctenidium, it may be significant that the false comb extends down over the vinculum, analagous to the case of fleas known to infest arboreal nocturnal hosts as mentioned above. The true host of Scolopsyllus 1s not yet known, the seven existing specimens having been collected from three species of Oryzomys or from Reithrodontomys mexicanus. Méndez (1968) believed that ‘Factors related to the micro-habitat of the flea on the host-pelage rather than the habits of the rodent or climatological conditions, are perhaps responsible for such modifications.” It seems to me, however, that the habits of the host account for the evolutionary development in Scolopsyllus, and that this comb is another instance of the adaptive mechanisms mentioned above as being typical in fleas having a special aeed for a device to help them fix on to a very active nocturnal or volant host. The only fleas of arboreal squirrels which show a marked tendency towards a reduction in the number of spines in the pronotal comb, accompanied by an increase in their breadth, are those which are highly modified as true nest-fleas like some Megarthroglossus°, Conorhinopsylla stanfordi° Stewart, 1930 and Epitedia faceta. It seems significant that these are fleas of diurnal squirrels that nest in fairly short trees or else live in tree-holes near the ground surface. It may also be relevant that each of these genera include species which are nest-fleas of ground-dwelling Peromyscini. Hence the adaptation to squirrel-nests may be a secondary or later development. The nest-fleas like Libyastus°® and Syngenopsyllus° which have a fuller complement of spines, as mentioned above, infest squirrels living in nests in very tall trees. More- over, it may turn out that the true hosts of these little-known fleas are nocturnal | | CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 341 or crepuscular, especially in the case of Syngenopsyllus°. (It may be that S. calceatus° (Rothschild, 1905) is really a flea of flying-squirrels.) Regardless, the coupling of the comb of this type with the loss of the pleural arch is adaptive, as indicated. Similarly, even if Tarsopsylla° were a nest-flea of Martes (the fisher and martens) rather than of tree-squirrels, as records for at least the North American form suggest is possible, the argument is still valid, for the various species of Martes are nocturnal and largely arboreal. The possibility also exists that Tarsopsylla° is a nest-flea of flying-squirrels and if true, this would reinforce the above theories. 5. CONVERGENCE IN VESTITURE IN UNRELATED FLeEas. That the above hypothesis about the evolutionary changes in the pronotal comb of fleas of flying- squirrels is correct is indicated by the modifications shown by the Australian Choristopsylla tristis°, one of the very few pygiopsyllids which lacks a pleural arch. Here we have the extraordinary case in which both parasite and host resemble, in important features, Nearctic species which are utterly distant phylogenetically and geographically. C. tristis° is a flea of Petawrus, a sugar-glider, a marsupial that looks and acts like a flying-squirrel (and, of course, there are no sciurids in the Australian— New Guinean area). Moreover, the flea exhibits reduced chaetotaxy, not only as compared with allied genera but also regarding C. ochi (Rothschild, 1904) (which has a very small pleural arch). The pronotal comb of C. tristis° bears about Io narrow spines per side, resembling that of the ceratophyllid Opisodasys vesperalis° (fig. 88), a Nearctic nest-flea of flying-squirrels, but there are further and surprising parallels with nest-fleas of arboreal sciurids. For example, in C. éristis°® there are 4 pairs of stout lateral plantar bristles on the last tarsal segment, with a mesal (ventral) pair in between the first pair and fully in line with them. (The mesal pair is in reality the true basal pair which has become displaced apicad and shifted towards the mid-line.) Thus, there is a total of 5 pairs of stout plantars in all. Ceratophyllids nearly always have 5 pairs of plantar bristles, and squirrel-fleas (and many others in the family) tend to have the first pair displaced somewhat mesad. However, the great majority of pygiopsyllids have 6 pairs, and the first pair is proximad of the second and only slightly moved toward the mid-line, e.g. Acanthopsylla. In Bradiopsylla Jordan & Rothschild, 1922, which is more closely related to Choristopsylla Jordan & Rothschild, 1922, there are only 5 pairs, but all are fully lateral. Even in C. och, which agrees with C. tristis° regarding displacement of the first pair, the third pair is somewhat shifted mesad, as in most members of the family, and there is also the customary dorsal pair of thin bristles on this segment, which C. tristis° lacks. C. tristis° is highly different from the general pygiopsyllids in another significant respect, and thereby also agrees with the Opisodasys of flying-squirrels in a trait that is highly unusual in the Ceratophyllidae—the first segment of the metatarsus is exceptionally long, being subequal to the tibia in length. In fact, the length of that segment approaches the condition in Tarsopsylla° (Holarctic squirrels or nests), which is almost unique in that the length of this segment I exceeds that of II-IV combined. (Liu & Li (1965) report that Brevictenidia, a monotypic genus of squirrel-fleas, resembles Tarsopsylla° in this respect.) Moreover, the relative length of metatarsus I in C. tristis° is signi- ficantly greater than that even in C. ochi, indicating specialization. The eye of the 342 R. TRAUB former species is somewhat larger and better developed than in the latter (although still small), but it is equivalent to that of the Opzsodasys of flying-squirrels (though ventrally excised instead of being entire). Such parallels as those cited for the tarsi strain the credulity as being bona fide examples of convergence, but the precedent for such an extraordinary development exists in the Order, namely the shifting towards the mid-line of the third pair of plantar bristles of the apical tarsal segment, which occurs in 2 unrelated genera of bird-fleas, Dasypsyllus Baker, 1905 (iii.t) and Ornithophaga Mikulin, 1956 (iv.2) (Holland & Loshbaugh, 1958). Similarly, in the pygiopsyllid bird-flea, Lentistivalius insolli, the third pair of plantars is displaced to a greater degree than in the mammal- infesting species of the genus. The same tendency is seen in some other (not all) pygiopsyllid bird-fleas known to me. An analogous shifting in the tarsal plantar bristles of bird-fleas has been noted earlier by Jordan amongst pulicids (1926) and cited as remarkable examples of convergence. In the tribe Spilopsyllini there are 3 genera (10 species) infesting mammals and all are combed, while 2 combless, monotypic genera parasitize sea- birds. In the mammal spilopsyllines the fifth tarsal segment bears 4 pairs of lateral plantar bristles of which the third pair is proximad of the middle or at the mid-line. However, in the bird-fleas, in Ovnithopsylla, the third pair is shifted distad, whereas a similar trend is shown in the male of Actenopsylla suavis (shifted distad in mesotar- sus and absent in the metatarsus) and the female has only 3 pairs on alllegs. The case in the Xenopsyllini is more thought-provoking, for a similar development is seen only in the two Xenopsylla which occur on birds instead of rodents. X. tispinus Waterston, Ig11 which infests Petrochelidon swallows in South Africa, is unique in that the third pair of lateral bristles on each side of segment is much nearer to the fourth pair than to the second. In X. gratiosa Jordan & Rothschild, 1923, a flea of Puffinus, a petrel, the third pair on the metatarsus gives the impression of being absent since one member is generally missing and the other displaced apicad and simulating a pre-apical plantar bristle (which are smaller and thinner than the laterals). On the other legs, in this species, both members of the third pair are shifted apicad and towards the mid-line. Thus, these bird-Xenopsylla exhibit a shift in the position of the third pair of tarsal bristles which Jordan (1926) alluded to as “crowding towards the apex,” or else have eliminated these bristles. In all the mammal-Xenopsylla there are 4 pairs of lateral plantar bristles and the third pair is equidistant from the second and fourth. It is of interest that, as M. Rothschild & Clay (1952) have pointed out, the kittiwake flea, Mioctenopsylla arctica Rothschild, 1922 (iii.1) although a member of another family, also shows a tendency towards “a weakening and loss of tarsal bristles—at least in the hind-tarsus.”’ Holland & Loshbaugh (1958) in remarking on analogous tarsal modifications in Ornithophaga (iv.2) and Dasypsyllus (iii.1) state: “If this is a case of adaptive evo- lution it is surely a remarkable one, and the survival advantage of such a character is not apparent to the authors.’ My feelings are similar, but since we are now learn- ing to appreciate that so many seemingly insignificant structural modifications of fleas do have meaning or function, perhaps the tarsal changes likewise do. However, Jordan (1926) offered another explanation for the tarsal modification in the bird- CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 343 pulicids. He pointed out that the “normal number of plantar bristles in Siphona- ptera is 5 pairs; that is evidently the ancestral number, some genera having acquired more such bristles, many others having lost some of them.’’ In the Spilopsyllini the trend has been to 4 pairs and hence “‘the tendency towards reduction already exists in the tribe.’ Jordan suggests that a “‘strong change in habit (or environment)” such as adaptation to birds as hosts “‘affects the growth and division of the chromo- somes, rendering them unstable’ and, in effect, this enhances the already existing genetic tendency towards modification of the plantar bristles. In short, the tarsal changes are unrelated side-effects that accompany the switch to a bird-host. Per- haps this is so, but certainly the many other modifications herein noted for bird-fleas and nest-fleas, etc. are apparently adaptive. The fleas characteristic of Aplodontia, the sewellel or “mountain beaver’ of the Pacific Coastal areas of North America, are instructive concerning several of the points that have been discussed, ranging from parallels in vestiture, to modifications suggestive of nest-fleas, shown by one species in particular. Three of the 4 Aplodontia-fleas: Paratyphloceras oregonensis Ewing, 1940, Trichopsylloides oregon- ensis® Ewing, 1938 and Hystrichopsylla scheffert Chapin, Igtg, are hystrichopsyllids, and the first two belong to the Rhadinopsyllinae and the third is in the nominate subfamily. The fourth species, Dolichopsyllus stylosus (Baker, 1904) is a lepto- psyllid in a unique monotypic tribe. In many respects the 4 species are as diverse as their generic names and taxonomic positions suggest, but they all agree in being very large, eyeless fleas, heavily clothed with a large number of very long, thin setae, and hence are remarkably similar in configuration (and thereby also resemble other fleas infesting burrowing mammals, as we have seen). The combed fleas of sub- terranean mammals like pocket-gophers and Bathyergus and Aplodontia (which not only digs well but spends most of its time in burrows) share another feature, and that is that the spines of the pronotal comb are unusually long and narrow, closely appressed and greater in number as compared to the condition in their relatives. The same tendency towards supernumerary, long, thin bristles aad spines is seen in mole-fleas as compared to shrew-fleas, viz., the genus Palaeopsylla (Traub & Evans, 1967). It does not seem to have been noticed before, but there seems to be a definite correlation between the possession of a pronotal comb with many, narrow, close-fitting spines and chaetotaxy characterized by a plethora ot long fine bristles. Thus, this is true not only for the fleas of burrowing hosts, but also for all the combed bird-fleas, their highly polyphyletic origin not withstanding. (As has been pointed out by Traub (1966) the trait of unusually fine setae is found not only in bird-fleas as compared to mammal-fleas, but also in Mallophaga, cimicid Hemiptera and hippo- boscid Diptera which infest both kinds of hosts.) This suggests that the ctenidial and chaetotaxic modifications may be adaptive and are associated with the pelage of the hosts at the site of infestation. Moles, pocket-gophers, Aplodontia, etc. have very fine, dense fur, at least on much of their bodies and the pinnules of certain feathers are also delicate and it therefore seems likely that the vestiture of these fleas has been tailored by the processes of selection to fit that of their hosts, but direct or quantitative observations have not been made insofar as concerns these 344 R. TRAUB particular fleas and animals. (Humphries (1966) in his fine paper on the function of combs, mentioned a bird-flea, but limited his studies to fur.) In the case of the fleas of burrowing mammals there is another factor that merits further consideration concerning this condition of close-knit, lengthy fine hairs or spines which extend from one segment well onto the succeeding one. Such fleas infest hosts which are covered with earth and dust, and vestiture of this type must be useful in preventing such minute debris from slipping under the extensions of the terga and sterna. Fleas of other genera, when collected from mice covered with dirt in a trap, not infrequently have grains of soil imbedded under the overlapping segments. Many kinds of birds deliberately and frequently dust themselves in an action interpreted as aimed at reducing the itching caused by lice and other ecto- parasites, and hence the same selective factor may be at work here in producing fleas with such modified vestiture. We lack sufficient data on the habits of birds and the infestation-rates of fleas to evaluate this hypothesis. One of the Aplodontia-fleas, Trichopsylloides® has some of the characteristics of nest-fleas. Thus, it is essentially a Rhadinopsylla° which has lost its genal comb; and the pronotal comb does not extend down as far ventrad as in the other Aplo- dontia-fleas, and the spines are somewhat broader; there is no trace of a pleural arch; the legs are long and thin, etc. The only feature that suggests other habits is the chaetotaxy and the combination indicates that Tvichopsylloides° is another instance of a nest-flea that can remain on the host when it leaves its nest. 6. MORPHOLOGICAL CHANGES IN THE THORAX ASSOCIATED WITH LESSENED JumpinG Apitity In FLEAS. In the Discussion thus far, reduced ability to jump has been shown to have adaptive value in many groups of fleas, ranging from bird- fleas and fleas of arboreal, nocturnal mammals, to fleas in nests underground. Methods of accomplishing this result are now considered. Although the pleural arch has been reduced or lost in many instances and in a variety of taxa throughout the Order, a variety of mechanisms have been employed to achieve the conversion from a good jumper to a poor jumper or crawler, demonstrat- ing that the term Convergent Evolution is an especially apt one in this case. In fact, the polyphyletic origin of the arch-less metathorax is apparent at a glance at figs 85-93, without recourse to the taxonomic characters used in systematics which placed the illustrated fleas in 5 families. This is not to say that representatives of each family modified the thorax in a manner that is characteristic of the family, for there are many variations on the thoracic theme in any one taxon (compare figs 85 and 86), and even in one genus there may be bewildering assortments (i.e. Rhadi- nopsylla). Fleas with a well developed pleural arch (PL.A.), and hence which are good jumpers, are shown in fig. 87 (Opisodasys hollandi) (iii.1) and fig. 91 (Cerato- phyllus styx) (iii.1) and one with a fairly well developed one (Xenopsylla vexabilis) (ix.1) is illustrated in fig. 93. In such fleas, the resilin which provides the power- motif for the leap, is housed under the arch, above the apex of the pleural rod (PL.R.). Fleas lacking the pleural arch have no housing for resilin and hence are poor jumpers. Two members of the first two genera are in this category, and are illustrated in fig. 88 (O. vesperalis°) and fig. 90 (C. arcuegens°). X. papuensis® (fig. 92) has virtually no CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 345 pleural arch (PL.A.(?)) and no doubt likewise leaps poorly. These thoraces have been described earlier in the section on Data and Observations, along with that of a bat-flea (Nycteridopsylla eusarca°) which also completely lacks the pleural arch, and it will be recalled, and can be seen, that there are some significant differences in the metathorax of the 4 depicted species which are inferior jumpers. However, these variations pale in comparison with those of 2 hystrichopsyllid nest-fleas, Conorhino- psylla stanford? (i.2) (fig. 85) and Wenzella obscura® Traub, 1953 (i.8) (fig. 86) which also lack the pleural arch. In the former species, an anomiopsylline, the pleural rod (PL.R.) is very thin and elongate, extending dorsad of the level of the third vinculum (VC.3), and here joins the dorsal margin of the metepisternum (MTS.) as the latter continues dorsocaudad, beyond what is generally the anterodorsal limit of the mete- pimere (MTM.). (The tanned portion of the dorsal margin of MTM. in this species disappears just anterior to the spiracle.) A semi-membranous region which is the vestigial lateral metanotal area (L.M.(?)) is represented as the vertical, narrow, dorsal section of MTS., near the apex of VC.3. The rhadinopsylline Wenzella° is obviously built on an utterly different plan, as witness the large, downward projecting triangle of the metanotum (fig. 86, MTN.) which fits between the metepisternum (MTS.) and the metepimere (MTM.). The metepisternum (MTS.) is reduced and terminates below the third vinculum (VC.3); it is overlapped by the large mesepimere (MPM.), and is caudally flanked by the pleural rod for its entire height. There is no sign of a lateral metanotal area, and even the squamulumis undeveloped. Themesepisternum (MPS.) is unusually straight and angled along the anteroventral corner, and the phragma of the nota (PH.2 and PH.3) exceptionally pronounced. The variety of paths used to reach the same end emphasize the adaptive nature of the response. However, sometimes the final products, superficially similar in appearance, may be achieved for different reasons, as shown by Johnson (1957), who presented an informative and interesting discussion of modifications of the lateral metanotal area, citing and illustrating additional examples than those just mentioned. She made the significant point that although Pulex and allied forms, are good jumpers, the lateral metanotal area has ventrally merged with the metepi- sternum, a condition which is also seen in poor jumpers like Anomiopsyllus’, etc. Johnson explained the paradox by stating that in the former “‘the metathorax has become heightened and narrowed” and indicating that ‘‘consolidation of the metepi- sternum and the lateral metanotal area would lead to greater strength”’ (for facilitat- ing leaping). Moreover, Pulex, etc. possess a pleural arch. “In Anomiopsyllus® and allies (nest fleas and poor jumpers), on the other hand, the pleural arch is missing and there are few strong internal sclerotized rods left which might serve to strengthen the metathorax so far as jumping is concerned.” There is at least one other thoracic feature besides the loss or reduction of the pleural arch and the lateral metanotal area, etc., that is often associated with the lessened ability to jump and the concomitant facility of crawling into cracks and crevices, i.e., the reduction or loss of the chitinized dorsal margin of the metanotal flange (or diminution of this ‘“‘collar’’), along with the loss of the metanotal spinelets. This is seen in many swallow-fleas, viz., those which occupy nests used perennially (and not just for one season) and which hence are not faced with the problem of seeking a new 346 R. TRAUB host elsewhere. The adaptive behaviour and physiological modifications of this first type of flea, which Darskaya (1959, 1964a) calls “settled’’ species, and of the “nomadic” fleas of the second category, will be discussed below (Section J). Here only the morphological features of the two types will be mentioned. In the “settled” group are the typical nest-fleas of the C. arcuegens° (fig. 90) and C. delichoni® pattern, which lack the pleural arch, but even those which retain the arch, are crawling, rather than jumping, species because the membranous or reduced metanotal flange permits a great deal of flexibility in the area of the junction of the thorax and abdo- men, ‘‘while at the same time it deprives the flea’s body of the fusion and compactness necessary for leaping” (Darskaya, 1959). Thus both C. hirundinis (Curtis, 1826), in which the metanotal flange is of this type, while the pleural arch is small and nearly contiguous with the top of the pleural rod, and C. scopulorwm Holland, 1952, which has a fairly well developed pleural arch and a metanotal flange that is only somewhat modified, are adapted to crawl into crevices in the nest. In the typical nest-fleas particularly, but also in fleas like C. hirundinis etc., the caudal margins of the abdominal segments are more lightly tanned, permitting marked expansion of the abdomen when the flea engorges with blood prior to the long months of fasting when there is no bird in the nest, as is mentioned below. In contrast, fleas like C. garet, which have to leave their birthplace and seek out and/or leap on a host, have well developed pleural arches, “normal’’ metanota and are well tanned throughout. Reduction of the posterior margin of the metanotum, including loss of apical spine- lets occurs in some other bird-fleas, such as Oynithophaga, as noted by Holland and Loshbaugh (1958). This is presumably a flea of woodpecker-nests, or nests of tits, which also nest in tree-holes (Rosicky & Smit, 1965). The notable reduction in the number and girth of bristles; the narrow, feeble legs, the over-all light sclerotization, all suggest that the species of Ornithophaga are primarily “settled” or nest-fleas. D. THE DEGREE OF DEVELOPMENT OF THE EYE IN VARIOUS FLEAS I. COMPARATIVE SIZE OF THE EyE. The degree of development of the eye of fleas varies tremendously amongst sundry groups of Siphonaptera, and sometimes even within a genus. Since some nest-fleas lack eyes, while in many they are reduced, and in some, are unusually large, it is necessary to discuss this topic before proceeding with subjects like the distribution of the nest-flea phenomenon in the Order. It is well known that reduced or vestigial eyes occur in many unrelated fleas which parasitize subterranean or nocturnal hosts (loff, 1941; M. Rothschild & Clay, 1952; Traub, 1953; Traub & Barrera, 1966). Further, the allies of such modified fleas may have eyes that are much better developed, especially if they infest diurnal hosts. Amongst eyeless fleas of burrowing mammals are Dactylopsylla (iii.2) and Foxella Wagner, 1929 (iii.2), which infest Nearctic pocket gophers; Pulex sinoculus Traub, 1950 (ix.1), the only eyeless member of the genus and a parasite of Central American pocket gophers; (Cvyptoctenopsyllus) ingens (i.4) a flea of the mole-like Bathyergus, which is essentially a blind and combless member of Dinopsyllus Jordan & Roths- child, 1913 (i.4) and hence its subgeneric status ; and the monotypic Cryptopsylla (x1.2) which infests a South African bathyergid, (Cryptomys) with similar habits. Another such flea is the Angolan species for which C. Fox created the name Rooseveltiella in ~ CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 347 1914, and which likewise parasitizes Cryptomys. Since this species is merely an eyeless Xenopsylla (ix.1), it is now called X. georychi. These hosts spend most of their life underground in dark burrows, and the eyes are minute in all of them. It is of interest that their fleas have followed a similar course of evolution. Although many species of Ctenophthalmus (i.3) have eyes that are only somewhat reduced, e.g. C. (Ethioctenophthalmus) calceatus Waterston, 1912, some species, such as C. (Spala- coctenophthalmus) spalacis Jordan & Rothschild, 1911, lack even a vestige. The latter is a flea of Spalax, a mole-rat whose eyes are so reduced that there isn’t even an opening for the eye in the skin. The analogy continues with respect to hosts like moles (talpids) and burrowing voles (Hyperacrius), which spend much of their time underground—their fleas have greatly reduced eyes, as in the examples cited and illustrated by Traub & Evans (1967) and in Hystrichopsylla talpae (Curtis, 1826) (i.r), while burrow-frequenting or noctural animals like shrew-moles and shrews have small eyes, and again the condition is paralleled by their fleas, e.g. Palaeopsylla Wagner, 1903 (i.3), Corrodopsylla Wagner, 1929 (i.5) and Doratopsylla Jordan & Rothschild, Ig12 (i.5), etc., in which the eyes are quite reduced. Nest-fleas of mammals which inhabit underground or enclosed nests (in hollow trees, etc.) and hence are in the dark virtually all the time, tend to have reduced or even vestigial eyes (figs 85, 86 and 92). Further, there is a direct correlation between the relative size of the eye of the flea and the amount of time the flea of underground nests spends on the host, away from its regular habitat. Thus, the less specialized neopsylline nest-fleas like Epitedia are not infrequently collected on the body of the rodent away from home, and in these fleas the eye is somewhat less reduced (fig. 98) than in the true nest- fleas (e.g. figs 85, 86, 92). Mammal nests made of leaves and perched high in a tree probably admit more light than an underground one, but at any rate, squirrel-fleas like Syngenopsyllus° inhabiting such nests have eyes resembling those of flying-squirrels (fig. 97), i.e. somewhat smaller than those of diurnal fur-fleas like Macrostylophora (as in fig. 96) (which may live on diurnal squirrels residing in the same tree, or even the same nest, as Syngeno- psyllus°). Amongst nocturnal mammals with fleas with poorly developed eyes are bats in general; the New Guinean murines and marsupials infested with many genera of pygiopsyllids; rats in the Pacific Islands with Sigmactenus Traub, 1950 (iv.1) and Neopsylla (i.7), microtines with Ctenophthalmus (i.3) or Amphipsylla (iv.2), etc. However, some nocturnal or crepuscular animals have greatly enlarged eyes which enable them to utilize what little light may be available, e.g. lemurs, lorises, jerboas (Allactaga), flying-squirrels, etc. and it is of interest that a few fleas have developed along parallel lines, e.g. jerboa- fleas like Mesopsylla Dampf, 1910 (iv.2) and cricetid fleas like some Ophthalmopsylla Wagner & Ioff, 1926 (iv.2) (fig. 95), while some desert-X enopsylla also have unusually large eyes (but others in the same habitat have very small ones) (Ioff, 1929; Traub, 1953). Mammals like flying-squirrels and dor- mice which are crepuscular or fairly nocturnal, and some species of rats with similar habits, have fleas with eyes that are well formed but small and oval (fig. 97). In contrast, fleas of diurnal mammals, such as squirrels, gophers, marmots, some 348 R. TRAUB carnivores, pigs, etc. (except in the case of fleas specialized for living in underground or dark nests), have very large and well developed eyes (fig. 96). It should be noted that virtually all known bird-fleas (other than the sticktight fleas, which have followed another evolutionary path) have relatively well developed eyes, whether specific nest-forms like Ceratophyllus arcuegens° (i1.1) (fig. 90) or a good jumper like C. styx (fig. 91). In fact, the eyes of fleas of terrestrial birds (whether ceratophyllids, leptopsyllids or pygiopsyllids) are generally characteristically larger than those of mammal-fleas, even if closely related. (This is one of the intriguing parallels that have accompanied this convergence to the bird-flea habitus, along with presence of many long and fine setae, etc.) All of these bird-fleas parasitize diurnal birds, however, and since the fur-fleas of diurnal mammals have well developed eyes, in contrast to the crepuscular and nocturnal forms, it is not surprising that the eye of the active types of bird-fleas is large. It is unfortunate that there does not seem to be any specific flea of a nocturnal bird, because it would be instructive to examine its eyes. However, the tendency for relative reduction of the once-hypertrophied eye does exist in nest-forms amongst bird-fleas. In C. delichoni°, the eye is rather small, symptomatic of “inhabitants of an ill-lit nest cavity” (Darskaya, 1959). The trend is also exhibited by some fleas of sea-birds which nest in burrows amongst rocks on the ground of small islands, and hence where the fleas presumably spend much time in the dark. For example, in Actenopsylla suavis (ix.1) the eye is quite small (fig. 99), more so than in related pulicids of terrestrial, diurnal mammals. The host of Notiopsylla kerguelensis (ii.1) also nests in such burrows, in the subantarc- tic (Smit, 1957), although the fleas also occur on the bodies of the birds, and it is worthy of note that the eye of this species is conspicuously smaller and more irregular than that of Hoogstraalia Traub, 1950, another pygiopsyllid, but one which infests the bodies of diurnal birds. 2. VISIONIN FLEAS. It is obvious, then, that the size and degree of development of the eye in the fleas is associated with the amount of light in the environment in which the flea evolved. The corollary would seem to be that the ability of the eye to function is equally correlated with this light-factor. Thus, it would be expected that squirrel- and bird-fleas could “‘see’’ better than vole- or mole-fleas, but there seems to be little experimental data in this regard. Rothschild & Clay (1952) state that the eyes of fleas “‘are in fact displaced dorsal ocelli. These relatively simple organs probably do little more than enable their owners to perceive the difference between light and darkness and thus would only assist them in finding a host if it were in their immediate vicinity.’’ Bates (1962) reported that sand-martin fleas (which have a well developed pleural arch) clustered about a burrow-opening on a cliff could hop onto a bird hovering by the aperture for just a few seconds. However, the stimuli causing this reaction are not known, nor has it been determined if the excited fleas merely leaped at random and just a few were successful in alighting on the bird. Bates showed that these martin-fleas, over-wintering as adults in pupal cases inside old burrows occupied the previous year, emerged in the spring and moved towards the opening of the burrow and there remained motionless, in clusters, with the heads pointing towards the aperture. It seems likely that the movement CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 349 towards the opening, where the fleas would be in a position to leap onto a returning migrant that hovered bythe hole, is a response tolight, but this isnot definitely known. However, these fleas were also shown to possess the ability to disperse from old burrows and emigrate as much as 33 meters and enter new burrows. While entrances to new burrows apparently were not detected by vision, the means to do so could not be determined, although it was learned that the fleas somehow recognize horizontal floors of burrows (as distinguished from the flat top of the bank) and tend to congre- gate there. M. Rothschild (1969) reported that many fleas are negatively phototropic and jump away from a bright source of light (and the ones that she mentioned have relatively well developed eyes). Such fleas would generally move towards the shade, if given a choice between light intensities, but a few individuals would move towards the light. Humphries has rendered a great service studying in detail such long neglected subjects as the behaviour of fleas, and among his important contributions are observations on Ceratophyllus gallinae (Schrank, 1803), the hen-flea (1968). Humphries noted that C. gallinae, after emerging as adults in the nest, are generally negatively phototropic for about 2 days, and as a result, stay in the nest rather than dispersing before mating. (The nests usually remain deserted by the hosts unless they return within that period.) Thereafter, the fleas exhibit a positive phototaxis and negative geotaxis, leave the nest and move upwards on the tree harbouring the nest. During the ascent, the fleas frequently pause for long periods and then resume climbing upwards if a host is not encountered. When in the immobile phase which Humphries termed a “‘stance’’, the fleas would immediately leap towards a small object that suddenly reduced the intensity of light. Humphries concluded that these responses were connected with locating a host and showed that fleas would readily leap upon an object the size of a small bird which moved about and paused on the branches. If, upon leaping, the C. gallinae missed Humphries’ imitation-bird, they would alight on the ground and then climb up on the vegetation and start anew. As a result, even if they were only on grass, they would be in a position to leap upon a ground-bird casting a shadow. Thus, while we really do not yet know how well fleas can see, it is apparent that they do respond to light and that those with eyes tend at times to move definitely or precipitously towards shadows. It therefore seems that regardless of other existing stimuli (vibrations, odor, carbon-dioxide, etc.), the sudden appearance of a host by the nest or by the burrow-aperture would cause a distinct change in light- intensity that would attract or spur fleas to leap upon their hosts. Contrariwise, such a shadow could not be thrown in a dark burrow underground, where the fleas lack the eyes to appreciate such changes anyhow. Darskaya (1954a) noted that Citellophilus tesquorum (iii.t) (which has well developed eyes) often cluster in great numbers about the burrow-opening of the suslik, Citellus dauricus (which is active during the day) and readily infest any ground-squirrel that puts its head in the hole or lingers nearby. The facts that fur-fleas of diurnal, large mammals have well developed eyes, e.g. Pulex Linnaeus, 1758 (ix.1), Ctenocephalides (ix.1), Pariodontis Jordan & Roths- child, 1908 (ix.1), Paraceras (iii.t), Oropsylla Wagner & Ioff, 1926 (iii.1), Thrassis 359 R. TRAUB Jordan, 1933 (iii.1), Opisocrostis Jordan, 1933 (iii.x), sundry vermipsyllids (x), infesting man, pigs, some carnivores, Old World porcupines, badgers, marmots, or prairie dogs, etc., and that these all seem to be excellent jumpers, lead me to believe that the large eye is of use to these fleas in finding their hosts. The fleas of the small rodents which are diurnal, likewise have well developed eyes (unless, of course, they are nest-forms) and the same line of reasoning applies regard- ing the function of such eyes is further supported by Darskaya’s detailed and well executed studies on X. gerbilli caspica and its desert-host, the gerbilline Rhombomys (1955). She states the “daily activity of these fleas is to a certain degree tied to the regime of light and daily activities of their hosts.” It is of interest that Darskaya reported that X. g. caspica (which has a fairly well developed eye) will cluster at the mouth of the Rhombomys burrows, particularly if they had hatched out in a burrow- system that was no longer occupied (1955). | Such hungry fleas have been known to instantly attack virtually any mammal near the opening and have been noted to follow such prospective prey for at least 2-5 meters. If they fail in their efforts to alight on the animal, they attempt to return to the burrow-openings. The association between the size of the eye of the flea and the degree of development and utilization of vision in the host is so profound, that I was long disturbed over the eyeless condition in X. papuensis®, for I know of no truly burrowing mammals in New Guinea, and existing records were from Pogonomys, a somewhat scansorial rat. However, after collecting some of these fleas in wnderground nests of Pogonomys on several occasions, it became clear that this species was merely exhibiting the same trend noted in other nest-fleas elsewhere, even to the extreme reduction of the pleural arch. In general, fleas possessing a genal ctenidium have a reduced eye, and the few exceptions are clearly adaptive. Thus, Hoogstraalia (ii.1) has a well developed genal comb and a large eye but it is a bird-flea, and as we have seen, an eye of that type is characteristic of such fleas. Ctenocephalides (ix.1) is similarly adorned, and since these fleas may hatch out anywhere in the haunts of the host, as indicated below, they may have to seek out and infest their hosts (large mammals) during daylight. As originally noted by Karl Jordan (1950) and discussed by Traub (1969), in fleas in which there is an evolutionary trend towards assumption of a vertical comb on the head, the vestigial eye is progressively displaced dorsad as the number of spines in the comb increases (e.g. Peromyscopsylla, Leptopsylla Jordan & Rothschild, 1911, Sigmactenus, etc.), indicating that the head-ctenidium in most cases is genal in nature (unlike the in helmet-fleas and in Smitella Traub, 1968 (ii.1)). 3. THE Types oF THORAX IN FLEAS WITH REDUCED Eyes. Inasmuch as many characteristic fleas of bird-nests have well developed eyes but lack the pleural arch, while many mammal-nest fleas with such a thorax have very small or vestigial eyes, it is clear that reduction of the eye has proceeded independently of loss of the pleural arch, the former associated with the amount of light in the environment, the latter with lessening of the ability to jump. The two phenomena often coincide because conditions under which the environment is invariably in darkness are frequently those in which reduction in jumping-ability is also a useful attribute. Examples CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 351 have been cited in which the eye is large but the pleural arch is missing (e.g. many bird-fleas and nest-fleas of diurnal arboreal squirrels). In many species besides true nest-fleas the eye is so reduced as to be indistinct, and this topic merits consideration with respect to (1) species in which the pleural arch is present, and (2) species lacking the pleural arch but simultaneously showing adaptations for remaining on the fur of the host. In the first category are fleas of mammals like shrews, voles and other hosts that are primarily nocturnal and/or spend much time in burrows, but in which the fleas are adapted for attachment to pelage by means of combs of spines, dense or plural rows of bristles, or false combs of modified bristles. Examples like Dorato- psylla (1.5) have already been cited. Here also belong other hystrichopsyllids such as Neopsylla (i.7), Rothschildiana (1.7), Stenoponia Jordan & Rothschild, r911 (i.9), Hystrichopsylla Taschenberg, 1880 (i.1), etc., as well as leptopsyllids like Peromysco- psylla (iv.1), Acropsylla Rothschild, 1gr1t (iv.2), Paractenopsyllus Wagner, 1938 (iv.1), etc. and many chimaeropsyllids (xi.)._ It is noteworthy that eyes of this type are apparently unreported in ceratophyllids, although the more “degenerate” condition of virtual total absence occurs in the pocket-gopher fleas, which are members of this family. The condition of a reduced eye is approached, however, in Malaraeus Jordan, 1933 (ui.1) which is the name ascribed to a polyphyletic and somewhat anomalous assemblage of fleas characterized by the common character of a small but ovaleye. Certain Jellisonia (i1i.1) and Kohlsva (iii.1) also have an eye of this type. Very few pygiopsyllids have a well developed eye, but in most species the eye is fairly large and well pigmented, although ventrally excised. In the South American and Central American genus Ctenidiosomus Jordan, 1931 (ii.1), however, the eye is reduced and fragmentary, and the species have multiple combs of spines. In the second group, ‘‘blind” fleas which lack the pleural arch but which bear special or well developed pronotal or even genal combs, or are otherwise modified for host-attach- ment, are the insectivore-fleas like Corypsylla discussed in Section LV. C.2 above. E. THE EVOLUTION OF THE CHARACTERISTIC PRONOTAL CoMB ASCRIBED TO MOST NEsT-FLEAS It has been shown above that throughout the Order, a variety of fleas have deve- loped the habit of feeding on their hosts while the latter are in the nests, and have become modified accordingly. A wide assortment of hosts are involved, and their habits have affected the path of development of the adaptive responses of the fleas, as host and parasite evolved together in close association. As we have seen, the prono- tal comb of nest-fleas infesting aerial hosts (bats and birds) (figs 89, 18, 90, 91) or those that parasitize arboreal, nocturnal hosts (figs 47, 88) tend to have an unusually large number of spines in the pronotal comb, as compared to allied taxa (figs 17, 46). However, nest-fleas of hosts nesting below ground or at, or near, the surface, tend to have pronotal combs in which there are fewer spines than “‘normal’’ and the spines are frequently short and broad (figs 67-84), and as previously indicated, such conver- gence cannot be ascribed to parallel proclivities in pelage. Instead, it seems that the pattern of the pronotal comb exhibited by fleas of this last category can only repre- sent evolutionary stages in the general reduction of spines and bristles so characteris- tic of these nest-fleas, and which accompany such “‘modifications by reduction or loss”’ 352 R. TRAUB as has been noted for the eye, pleural arch, metanotum, size of metepimere, girth of legs, etc. Examples have been cited above repeatedly showing how nest-fleas have lost or reduced these structures or a combination thereof, in approaching the configuration of the virtually nude species of Anomiopsyllus° and the combless, eyeless, slender Wenzella°. That reduction in the number of spines in the combs has accompanied the loss of bristles is not only indicated by the general trend, but by the fact that the genera with the fewest, and stoutest spines, e.g. Megarthroglossus® (1.2) (fig. 75) and Conorhinopsylla® (i.2) (figs 73, 86) are those with the most marked loss of setae. The setae seem to become narrower as they approach extinction in the course of evolution, but in the case of the pronotum the spines frequently become shorter and broader as in the nest-fleas. Phalacropsylla (i.7) approaches the last two genera with respect to the points mentioned. Brachyctenonotus° (iv.2) has a comb of extremely short spines, but these are fairly broad and the comb is almost full length (in height). Calceopsylla® (iv.2) duplicates the condition in Brachyctenonotus® regarding tiny spines, but the spines may be somewhat more slender and the comb is not quite as short. The pronotal ctenidium in many of these nest-fleas extends ventrad to near the vinculum, but it is definitely shortened in Rhadinopsylla (i.8) (fig. 72) and most neopsyllines (i.7) (figs 67, 70, 78, 76). In other instances, impending loss of the comb is indicated by the reduction in the vertical extent of the comb and in shortening of the spines, as in the chimaeropsyllids and in Agastopsylla® (1.3), or by loss of pigmenta- tion accompanying reduction in length, as in Mioctenopsylla Rothschild, 1922 (ii1.1) and Brevictenidia (iii.1), or as in both genal and pronotal combs in Agastopsylla. The loss of abdominal ctenidia by marked reduction of the number and size of the spines and by the shortening of the height of the comb as has been discussed by Traub & Evans (1967) and is also illustrated in the bat-flea Nycteridopsylla eusarca (v.1) (fig. 89). A different mechanism for the loss of pronotal spines can be seen in Corypsylla oynata® (i.8) (which may not be a true nest-flea in that it is simultaneously adapted to remain on the host) and involves sexual dimorphism. Here the male has a full length comb of spines, of which the upper fourth are long and stiletto-like, and the emainder becoming progressively shorter and less acutely pointed. The shortening in length is accomplished mainly by the loss of the base of the spines, and not merely by abbreviation of the apices; the proximal ends of the spines forming a sigmoid line whose lower 3 is markedly convex. The original full length of the spines is indicated by pale outlines and caniculi. In the female, however, the dorsal 3 of the comb consists of stiletto-like spines, which also become progressively smaller, but the lower 2 of the comb is represented only by pale blunt nubbins of spines which are more like denticles, with long caniculi proceeding to the former base of the comb. The two known genera of malacopsyllids (xiv) (both monotypic) are not nest-fleas, although they have followed another path leading toward reduction and loss of combs, namely the development of the “‘sticktight’”’ habit. It is instructive that Malacopsylla Weyenbergh, 1881 has no ctenidia at all, but that in Phthiropsylla Wagner, 1939, the pronotal comb is represented by but 3 very short, broad spines or “teeth” which are widely spaced. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 353 The tendency towards loss of ctenidia is general throughout nest-fleas in the Order, as shown by the fact that combless members amongst such fleas occur in 5 of the families which include nest-species, while the trend is also demonstrated in the sixth (Leptopsyllidae). There can be little doubt that intermediate steps in the evolu- tionary path, and not the structural peculiarity of the pelage, account for the conver- gence of patterns of pronotal combs seen in these nest-fleas inhabiting underground nests, etc. F. DistTRIBUTION OF THE NEST-FLEA PHENOMENON IN SIPHONAPTERA Although nest-fleas occur throughout the Order, it is apparent from what has been written above, that the phenomenon is much more common in certain groups of fleas than in others, and that the degree of specialization exhibited varies considerably. Both of these points can presumably be explained by the following factors: (1) The duration and extent of the association between the fleas and their respective hosts. (2) The phylogeny of the fleas and of their hosts. (3) The ecology of the area concerned. (4) The habits of the host. (5) The habits of the fleas. The relative geological ages of the various groups of fleas and some of their hosts will be treated at some length in a subsequent article (Traub, in preparation) but a brief discussion of certain elements is warranted at this time, while considering the other points. The vast majority of the nest-fleas are members of the Hystrichopsyllidae (referred to as family i), where representatives occur in 4 of the subfamilies. There were several examples cited for the Ceratophyllidae (iii), but relatively few for the Puli- cidae (ix) and Leptopsyllidae (iv). The proportion of nest-fleas is fairly high in the Chimaeropsyllidae (xi) but the family is a small one and only a few species are actually involved. The condition is rare amongst known Pygiopsyllidae (ii) but the dearth may be more apparent than real, for the following reasons. (1) The family is pri- marily Australian—New Guinean in distribution, with relatively few species occurring in the Asiatic—Pacific islands, the Indo-Malayan region and Central Africa. Throughout much of their range therefore, the pygiopsyllids are found in regions of heavy rainfall and hence where mammals make few underground nests because of the high water-table and flooding, and where relatively dry, hollow cavities in logs, stumps and trees are rare or absent. In such an environment there has been little opportunity for development of the nest-habit in fleas, except perhaps for those infesting arboreal hosts living in tree-holes or making some sort of aerial nest that adequately sheds water. (2) Relatively few mammal-nests have been examined for fleas throughout the entire range of the pygiopsyllids, and even the fur-fleas of indigenous Australian rodents and marsupials are inadequately known. The same is true tor body-fleas in the other areas, save New Guinea. It seems likely that pygiopsyllid nest-fleas await discovery, perhaps in xeric areas of Australia or in arboreal species elsewhere. Very little has been written about the phylogeny of the various families and other groups of fleas (Traub, 1968), and it is beyond the scope of this article to do more than point out that the families Hystrichopsyllidae and Pygiopsyllidae are regarded by me as being ancient and primitive in many respects (but specialized in others), while the Pulicidae is considered to be a very old, but yet more advanced family than 354 R. TRAUB the first two, and the Leptopsyllidae, Ischnopsyllidae and Ceratophyllidae are deemed younger. Reasons for these views are cited in another article (Traub, in preparation), but are based on three considerations: (1) Morphology, including general skeletal structure and comparative anatomy of the aedeagus. (2) Host-relationships. In general, the more primitive and oldest fleas are associated with the more primitive hosts, and the more specialized fleas with the more advanced (and younger) mammals. (3) Zoogeography. Fleas essentially restricted to two or three of the southern conti- nents are believed to have spread amongst those areas when island-hopping links still existed between them, well before Australia and South America became isolated. In general, the hystrichopsyllids infest mainly insectivores, cricetids, and sciurids, while the ceratophyllids, on the whole, parasitize mainly cricetids and sciurids, and some murids. Relatively few hystichopsyllids (e.g. some Neopsylla) and ceratophyl- lids (e.g. certain Nosopsyllus) infest the genus Rattus and it is significant that there are no known nest-fleas specifically associated with Rattws, which is believed to be a recently evolved group. Leptopsyllids are rarely specific to insectivores except perhaps for tenrecids in Madagascar, and instead are found on murids and, particu- larly in the Old World, on cricetids. Some leptopsyllids are associated with dipodids and gerbillines and a few with ground-dwelling Palaearctic sciurids and lagomorphs. The pygiopsyllids are, on the whole, either fleas of marsupials or murids. Pulicids are found to parasitize a broad range of hosts when considered as a family, but some specialization is shown by certain of the subgroups. Many are sticktight fleas or “burrowing’’ species like Tunga, as has been indicated, and these need not be con- sidered further. The spilopsyllines include rabbit-fleas which tend toward semi- sessile habits, and the two genera of nest-fleas of marine birds treated elsewhere. The xenopsyllines are primarily fleas of gerbillines (and hence occur in rather xeric areas) or are mane-fleas. The archurieopsyllines include carnivore-fleas like Cteno- cephalides, hedgehog-fleas, and one odd rabbit-flea, a Madagascar tenrec-flea and the African Aphropsylla, whose true hosts are unknown. The questions arise as to why the bulk of the nest-fleas occur in the hystrichopsyl- lids, why the phenomenon exists in so many subfamilies and tribes thereof (e.g. 7 of 8 Anomiopsyllinae genera and 8 of 11 genera of Neopsyllinae), and why the examples include some of the most highly specialized nest-fleas, e.g. Anomiopsyllus° and Wen- zella®. Inasmuch as the host-associations and morphology of the hystrichopsyllids suggest that the family is an ancient one, the obvious answer is that hystrichopsyllids and their hosts have been associated intimately for eons, long enough for the nest- habit of fleas to have evolved and progressed to the point of extreme specialization by reduction. Nevertheless, it must be pointed out that, for unknown reasons, very few instances of the relationship involve insectivores, which are believed to be low on the phylogenetic scale. Virtually the only examples are some species of Rhadt- nopsylla, and these are not highly specialized, and may not be specific. Perhaps if more were known about the fleas of nests and shrews in North America and northern South America, some such true nest-fleas would be discovered, but it may be that the nests of shrews are too transient for such fleas to have evolved, and this is why virtually all shrew- and mole-fleas are adapted for remaining on the host. Regard- less, hystrichopsyllids are found on all continents, but have an extremely discon- CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 355 tinuous distribution, e.g. huge gaps over most of South America, and known endemi- city in a restricted portion of Australia, with the geographically closest members of the family occurring in Borneo and Indonesia. Such a range also bespeaks of antiquity, as does the existence of two endemic subfamilies in Africa. Nest-fleas occur in virtually all of these areas, perhaps also including Australia. The highly modified Anomiopsyllus° and Wenzella® are parasites of heteromyids, a group that arose in the early Oligocene, with some modern genera occurring at least in the Miocene (Simpson, 1945). A large proportion of these hystrichopsyllid nest-fleas (e.g. Megarthroglossus°, Stenistomera°, Callistopsyllus°, etc.) are (like Anomiopsyllus°) associated with the fauna of the deserts of North America, another phenomenon that was Oligocene in origin. For these reasons, the extreme age of the family and its associations seem to have been an important factor in the evolution of the nest-habit. The sciurids and leporids, however, are older than the rodents mentioned above, and yet the family of fleas most closely connected with squirrels, the ceratophyllids, include only a few nest-fleas. Here we must differentiate between the geologic age of a group of mammals and the duration of the association of the fleas with such animals, for they may be very different. For example, although birds arose far back in geological time, their parasitism by fleas must be relatively recent, for the sundry bird-fleas, representing several different families, are all supposed to be derived from mammal-fleas (Rothschild & Clay, 1952; Holland, 1964; Traub & Barrera, 1966). That this was so, was well known to Jordan (1937) and N. C. Rothschild (1917), and loff (928) also referred to this phenomenon in pointing out that bird-fleas of the Ceratophyllus gallinae group are related to mammal-fleas now referred to as Monopsyl- lus, e.g. M. sciurorum and M. tamias (Wagner, 1927). Ioff further noted the close affinities of the rodent-infesting Frontopsylla with those on birds (but which are today called Orfrontia Ioff, 1946). Moreover, the bird-infesting genus Ceratophyllus itself is believed to be of polyphyletic origin. Another point is that the particular genus, tribe or subfamily of host may be relatively recent in origin even though the stock is ancient. One feature that is quite striking is that there ave quite a few nest-fleas parasitizing sciurids, but most of these not only are hystrichopsyllids, e.g. Conorhino- psylla°, Megarthroglossus°, Rhadinopsylla°, Epitedia, Tamiophila, etc., but genera which also includes species which are nest-fleas of peromyscines and other cricetids, and hence may represent recent transfers to sciurids. Actually, true squirrels are known from the Miocene, while the groups of tree-squirrels may be much younger, but the fossil record is scanty (Simpson, 1945). It may very well be the relative youth of the tree-squirrels that accounts for the comparatively small number of nest-fleas known amongst such hosts, and for the low degree of specialization shown by such ceratophyllids, despite the selective pressure exerted, for such an evolution, by the hazards of existence encountered by fleas in the high trees. It is noteworthy, however, that the bulk of the ceratophyllid nest-fleas of mammals is found amongst the parasites of squirrels nesting in high trees or amongst flying-squirrels. Another factor for the apparent dearth of nest-fleas in this family is the presumed modernity of ceratophyllids as compared to the hystrichopsyllids—there hasn’t yet been time | for the marked reduction in head, thorax, etc. to occur. > 356 R. TRAUB Offhand it seems strange that so few nest-fleas are known to parasitize the Marmo- tini, the tribe including the northern ground-squirrels (Citellus, etc.), marmots (Marmota), prairie-dogs (Cynomys), chipmunks (Tamzas, etc.), and which is believed to have arisen in the Miocene (although all records save for Citellus are Pliocene or later). A wealth of genera and species of ceratophyllid fleas such as Citellophilus Wagner, 1934, Callopsylla Wagner, 1934, Thrassis, Opisocrostis, Oropsylla, Diamanus Jordan, 1933, are associated with these rodents that essentially are ground-squirrels but which vary in size from small to huge, but only the occasional species like Callopsylla dolabris (fig. 84) seem to be developing the nest-habit. (It should be noted that the fur-fleas of Citellus and other true ground-squirrels, such as Citellophilus, Opisocrostis, Thrassis and Diamanus, do not exhibit the tendency towards broadened subventral pronotal spines of the fleas of the arboreal and semi-arboreal squirrels.) The other nest-fleas occasionally encountered on chipmunks and Citellus, etc., are hystrichopsyllids, e.g.,!amiophila, as has been pointed out to be the case for the nest- fleas of tree-squirrels. The marmotines burrow and nest underground in habitats that should favour the production of nest-fleas, for they abound with other types of fleas, some of which have long palpi (like Diamanus, etc.), indicating feeding at leisure. One contributory factor may be that these hosts are all diurnal, meaning that any flea which lingered on the host would be exposed to light considerably, especially if it fell off—or developed from an egg that had been hatched out on the ground surface. Fleas with reduced eyes would be at a selective disadvantage under such conditions, and it is reiterated that virtually all of the nest-fleas of mammals have hosts that are crepuscular or nocturnal. Many, if not most, of the ceratophyllids of burrowing rodents are well known for swarming about the opening of the burrow (Darskaya, 1954a, 1955), and this habit no doubt is all associated with the presence of a well-developed eye. In this way, the fleas are able to leap upon rodents that approach or enter the burrow, or even pursue them fora while (Darskaya, 1955). In all probability, these fleas have hatched out in the nest or in various depths of the burrow and move toward the light source at the aperture, as Bates (1962) observed for the perambulatory type of sand-martin - flea. Thus, the evolutionary trend of the ceratophyllids has in general been in direc- tions opposed to the formation of true nest-fleas, although the time factor of length of association between host and flea may also be important. Leptopsyllids on the whole have followed much of the route of the ceratophyllids insofar as concerns evolution of habits of infestation, but have detoured with respect to host-range and have become delayed regarding distance traveled. Thus, very few genera and species occur in North America (where all native forms except two are associated with lagomorphs), whereas the ceratophyllids are the dominant members of the Siphonapteran fauna in that part of the world. Leptopsyllids not only infest cricetids but to a great extent parasitize murines, one species having even accom- panied Rattus to New Guinea, whereas there are no native murids in the New World. (The last-mentioned point no doubt helps account for the dearth of leptopsyllids in North America.) There has been little association of leptopsyllids with sciurids and virtually none with tree-squirrels in particular. Except for fleas of murines (which are nocturnal), nearly all leptopsyllids have fairly well developed eyes, while the CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 357 bird-infesting species have large eyes. In general, they seem to be good jumpers and rodent-species may be common around burrow apertures, while nearly all are modified for remaining on the body of the host. The few known species that seem to be nest-forms have converged along the familiar lines indicated above. One relevant and possibly important factor in this discussion is that there still is a great deal to be learned about the nest-inhabitants of desert rodents throughout much of Asia (excluding well studied areas of the U.S.S.R.). The majority of North American nest-fleas are found in xeric areas, and very few leptopsyllids are represented in the fauna, whereas members of this family are common and wide-spread in the Asian counterpart terrain. The African nest-fleas are also inhabitants of deserts and semi- deserts, in the main. The equivalent fauna in Asia may therefore be better deve- loped than current data indicate. The answer to the question as to why nest-fleas are poorly represented amongst pulicids lies in the evolutionary sequences observed in that family, viz., development of sticktight fleas or else active species with well developed eyes, which are good jumpers and which are so highly adapted that they can feed and thrive on an excep- tionally broad variety of hosts. It should be noted that just because a flea has stick- tight habits, it does not mean that it is zpso facto a poor jumper, for the behaviour of a flea after it finds and attaches to its host may be quite different than it was before- hand. Some sticktight fleas like Echidnophaga Olliff, 1886 (ix.1), which are essen- tially sessile after attaching, are excellent jumpers. Thus, E. gallinacea (Westwood, 1875) can leap 200 times its own length (M. Rothschild, 1969), a fact that is useful when it encounters its usual prey of medium or large size, such as rabbits, carnivores, pigs, chickens, etc. The catholicity of pulicids regarding hosts is illustrated by the so-called cat-flea Ctenocephalides felis (Bouché, 1835) which has not only been collected in numbers on all sorts of carnivores, including dogs, foxes, wolves, jackals, tigers, Felis bengalensis and other “wild cats,” racoons, coati-mundi, weasels, mongoose and other civets like the binturong (Arctictis bintuwrong), but on man, opposum, cattle, horses, sundry rodents, lagomorphs, hedgehogs, bats and the armadillo, etc. (Traub, 1950; Hopkins & Rothschild, 1953; Hoogstraal & Traub, 1963a, b, 1966; and unpublished data). It has also been known to feed on lizards (Fox, Fox & Bayona, 1966) and the related C. cams (Curtis, 1826) has been reported sucking blood from a lizard (Jath, 1952). There are not many records of C. felis from rodents, as compared to the large mam- mals, but this, I believe, reflects its superior ability as a jumper, and not to any inherent predilection or aversion, since it ordinarily vaults too high to strike a rat- sized animal. The plague flea, X. cheopis is notorious for feeding on man as well as tats, but has a very broad host-range in general, from a variety of murines and other rodents, to shrews, lagomorphs and bats, as cited by the sources mentioned above. In this case, the vast majority of records are from small hosts, for X. cheopis does not leap as high as Ctenocephalides. X. gerbilli caspica has been reported to feed on lizards (Darskaya & Besedina, 1961; Kulakova, 1964). The majority of the pulicids are compact, i.e. relatively short and broad, a type of build that is often seen in fleas that are good jumpers. The species, whose life-his- tories are known, tend to lay eggs loosely among the hairs of the host, as in Cteno- 358 R. TRAUB cephalides, so that the larvae and adults are apt to emerge in the haunts of the host, rather than in the nest. Larvae of the desert-species X. conformis (Wagner, 1903) have been found to be abundant in the rodent-burrow, at the entrance, indicating that the eggs can develop wherever they fall, if conditions are favourable (Darskaya et al., 1962). In the case of large mammals (e.g. Pulex and pigs) this can be almost anywhere. This particular trait of egg-laying is associated with fleas with large eyes and a well developed pleural arch, as has been noted. Forthese reasons, development of the nest-habit in the Pulicidae has occurred only under special circumstances, as in some of the fleas associated with burrows of marine birds and in underground nests of Pogonomys, etc. A few Xenopsylla associated with the desert have small eyes, as befitting ectoparasites of nocturnal hosts that wouldn’t emerge from their burrows during the heat of the day. The evolution of the nest-habit virtually by definition is limited to fleas of hosts that (1) have a specific nest or burrow and (2) reside there continuously or else return to live or breed there the following year or season. It is probably for this reason that no true nest-fleas are known amongst the rabbits and hares (leporids), and that instead, the rabbit-fleas often act somewhat like sticktight species. It is noteworthy that the rabbit-warrens often do swarm with fleas when the young rabbits are born, but this is presumably due to transfer of fleas from the adult rabbits, as part of the synchronization of the fleas with the hormones of the host, as has been clearly shown for the European rabbit-flea (M. Rothschild, 1965a, b). The splendid fleas of the Mexican volcano-rabbit, Romerolagus described by Barrera in 1967, defied discovery, despite intensive search of adult Romerolagus over a period of years, until the nestlings were located and examined. G. Tue AssociATION BETWEEN THE MODE oF EGG-LAYING AND THE NEST-FLEA HABIT Intimately associated with the pattern of behaviour exhibited by various kinds of fleas in locating a host, is the factor of the particular site where the flea-larvae trans- form into pupae and hence where the adults emerge. In this regard it 1s important to note that not all fleas deposit their eggs loosely amongst the hairs of the host as in the case of the cat- and dog-fleas (Ctenocephalides) (ix.1) so that the eggs may drop off anywhere in the haunts of the host. Some fleas, such as Orchopeas howardi (Baker, 1895) (iii.1) leave the host and glue the eggs to bits of debris in the nest and then return to the mammal. In the latter case, the immature stages are to be found in the nest, and that is where the newly emerged adults are to be found. This is why some species of fleas are common in the nests, especially at certain times of the year, even though they are not true nest-fleas in the sense we are using the term. I believe that Stenoponia and Hystrichopsylla are fleas that lay their eggs in the nest and hence are in this category, but real data are lacking. Stenoponia females are often noted with two huge eggs in the abdomen (as so beautifully depicted in a colour photograph by Sakaguti, 1962), in contrast to ceratophyllids, which may have 8-12 smaller, but fully developed ones. It would be surprising if Stenoponia could spew out such huge eggs while on the fur of an active host. Moreover, Stenoponia is a winter-flea, according to our observations in Pakistan and as indicated by available CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 359 and published records in the U.S.A. and Mexico, and adults are collected only during the cold season. The larvae apparently take nearly a year to develop, and the un- mistakably large larvae can be seen in the rodent-nests in the summer, if not earlier. All these points suggest that the Stenoponia larvae could not find suitable conditions just anywhere, and that, instead, the life-cycle has become attuned for nest-develop- ment. In my view, this is also the case for other fleas which are frequently found in nests but which are modified along the lines of a fur-flea (and which Ioff (1929, 1941) has referred to as “‘nest fleas,’ along with the specialized fleas for which we used the term). Vaschenok (1967), in fine detailed studies on the ecology and life-history of the pulicid Echidnophaga oschanini Wagner, 1930, has provided relevant observations along these lines, which also emphasize the physiological attunement of the flea to its host. This species lays its eggs only when in darkness, and since its host (Ochotona) is diurnal, the result is that the eggs are deposited only when the pika is inits burrow. Further, since the eggs are dry and non-adhesive, they immediately fall to the floor of the nest or lair, where there is an environment suitable for development of the larvae. It is also clear that the egg-laying habits of the squirrel-infesting Orchopeas, mentioned above, are also adaptive, for adults emerging in the nest are ensured of food, especially during the breeding season, whereas larvae developing from eggs dropped indiscriminately from the fur of an active arboreal host may have a difficult time finding a suitable environment for development, as might the newly emerged adult flea in locating a squirrel. Thus it would seem that just as there are nest-fleas, so there may be nest-larvae, although so little is yet known about the larvae of most kinds of fleas that it is premature to discuss possible morphological or physiological differences between the free-living larvae and the nest-bound ones. The possible significance of the specific requirements for a nest environment for some species has been indicated in the first paper of this series (Traub, 1972a) when it was pointed out that the species of Medwayella infesting ground-squirrels had never (or extremely rarely) been collected on arboreal squirrels, and it is believed that the conditions in the nest account for these differences. H. REDUCTION OF COMBS, ETC. IN FLEAS OTHER THAN NEST-FLEAS Inasmuch as the absence of combs and reduction in chaetotaxy is characteristic of many nest-fleas, it is desirable to refer to 3 families in which all members bear these features but are not nest-fleas in the ordinary sense, viz., the Vermipsyllidae (x), Coptopsyllidae (xiii) and Malacopsyllidae (xiv). The majority of species of Vermi- psyllidae belong to the genus Chaetopsylla Kohaut, 1903, and all of which are parasites of carnivores, and are characterized by the following: large eyes; frequently fairly short mouthparts (but at times they are quite long, reaching the fore-trochanters) ; a well developed pleural arch and leathery, flexible abdominal segments, permitting great expansion. This unusual combination of characters can be explained by their habits—they are primarily den-fleas, specialized to feed for long periods on hibernat- ing hosts during the winter months and early spring and/or when the young carni- vores are born in the den. Presumably the eggs are laid loose among the fur of the 360 R. TRAUB host (as in the case of the vermipsyllids mentioned below) and hence can fall off at any time and develop into larvae wherever the micro-environment is suitable, al- though many of these will be in the den. Many of the carnivores are diurnal, but the large eyes of the flea no doubt are also useful in enabling the newly hatched fleas to find their host, particularly if out-of-doors. As is to be expected, in the case of fleas infesting large mammals, especially if prone to hatch out on the ground, the vermipsyllids are good jumpers when unfed. Virtually all the records are from the cold parts of the year, or else are from alpine or subarctic heights in the mountains, indicating the species spend the rest of the year as eggs, larvae or pupae. The dense fur of the hosts enable these large fleas to hide easily if on the body, and to crawl with dispatch when grossly engorged. The other two genera of vermipsyllids, Vermipsylla Schimkewitsch, 1885 and Dorcadia loff, 1946 are extreme in their adaptations for existence as “‘alakurts’, grub-like fleas infesting the dense wool or thick patches of hair on the bodies of various ungulates in the high mountains of Central Asia. Very few fleas parasitize ungulates, probably because such hosts do not have dens or nests where the larvae have a good chance to develop, and the fleas an opportunity for a steady diet. Moreover, the hosts of the alakurt frequent forbidding terrain, and all in all, very few other kinds of fleas could possibly complete their life-cycles under such condi- tions. However, the alakurts emerge in the alpine fields and bogs at the onset of cold weather, when the horses, sheep, deer, etc. have dense coats, and the fleas, which have large eyes and are excellent jumpers, actively seek and leap upon their prey in daylight. (At night it is too cold for the fleas to move about, even if they were able to locate their hosts.) Their success, at least in the case of domesticated animals, is indicated by Ioff’s report (1950) of 7000 specimens on a single sheep. In D. dorcadia (Rothschild, 1912), the size of the newly emerged female is stated to increase more than 250-fold as it engorges on its host and as the eggs mature (Loff, 1950). The males remain active on the host and do not feed much, but the females remain 7 situ on the host for very long periods (as expected in fleas with such long mouthparts). Grossly-swollen females can crawl maggot-like, or by a wiggle induced by peristalsis-like waves, since they are too fat to use their legs. Dorcadia have been observed to remain on a single host for 3 months although moving about from time to time. The eggs are deposited loosely amongst the hairs or wool, and eventually fall to the ground, where the eggs may remain viable but unchanged all winter. The larvae emerge in the spring, feed on organic debris, often in clumps of dried sod, and pupate in the late summer or fall. It is of interest that these larvae, which may hatch out in the open, are believed to have a rudiment of an eye (off, 1950) unlike other known flea-larvae, which are generally in the dark. It is important to note that the Vermipsyllidae are believed to have always been without combs of spines (at least on the pronotum), unlike the families of fleas we have been discussing heretofore (Jordan, 1947; Holland, 1949; Traub & Evans, 1967),1° and thus only superficially resemble the nest-fleas which have lost the combs by reduction. As can be seen from the above outline, the vermipsyllids differ likewise in other details of structure, habits and hosts—one notable development is the labial palpus of 25 joints in Dorcadia, instead of the 5 segments seen in most fleas. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 361 The Coptopsyllidae are another family that may have lacked combed ancestors since none of the species possess even apical spinelets on the abdomen, and there is no trace of genal or pronotal combs. One species even lacks the mesonotal pseuso- setae which is supposed to represent a rudimentary (incipient) ctenidium—a condition that is rare in most families of the Order. Here again the low number of abdominal bristles and the presence of very long mouthparts are suggestive of nest-fleas, but the large eye and frequent presence of extremely long tibial bristles, and the supernume- rary plantar tarsal bristles are not. Moreover, the abdominal bristles are often long. The marked length of the tibial bristles is suggestive of other desert-fleas, e.g. some Xenopsylla, where Ioff (1941) believed they were used in burrowing into the sand to enable the flea to escape the sun and heat. The pleural arch is large and hence although the lateral metanotal area is higher than long, I believe they may be fair jumpers. As Lewis (1964) points out, a fairly large number of species have been described, but only a few specimens have ever been collected, in most instances. Too little seems to be known about their habits to warrent further discussion. The Malacopsyllidae (xiv) include 2 South American monotypic genera, Malaco- psylla and Phthiropsylla. These, although combless or essentially so, and with re- duced vestiture are obviously modified as “‘sticktight”’ fleas, i.e. to stay attached to the host for long periods and feed until swollen with blood (hence the specific name M. grossiventris proposed by Weyenbergh, 1879). Thus, the cuticle is leathery and bears longitudinal pleats, permitting great expansion, while the tibial bristles are extremely short and stout, as are some of the tarsal bristles, and even the plantar bristles are stout or even subspiniform. The eye is large and the pleural arch well developed, indicating these fleas seek out their prey, leap upon them and catch and hold on by means of the specialized tibiae and tarsi, and therefore are not nest-fleas. These conclusions are the same as those reached by Johnson (1957). The sticktight habit tecalls the case in the Pulicidae (ix), where the hectopsyllines and many of the Pulicinae are combless, have large eyes, are excellent jumpers and are fixed or semi- sessile on the host (but which stay attached by means of anchoring mouthparts). I. THE PossiBLE RELATION BETWEEN THE PRONOTAL COMB AND A FRIGID ENVIRONMENT Smit & Dunnet (1962) in remarking on the facts that in the Antarctic Glaciopsyllus antarcticus°®, the pronotal comb is represented only by pseudosetae (or by apical spinelets), and in the Arctic Mioctenopsylla is greatly reduced (i.e. the spines are extremely small and pale), stated that ‘‘there may conceivably be a partial correla- 10 It is not clear as to whether Jordan (1947) meant that the vermipsyllids never had any combs on any of the segments or if he had only the pronotum in mind. Combed fleas always seem to have at least the pronotal comb, and Jordan believed that the pseudosetae represent incipient combs. It would therefore seem that the presence of pseudosetae on the pronotum, which is the case in vermipsyllids, rules out the possibility that the ancestor of that flea had a pronotal comb, and it would then follow that the ancestral flea lacked other combs as well. This may very well be the case, but the Antactic flea Glaciopsyllus antaycticus° is unique in the family Ceratophyllidae. in lacking a pronotal comb, and presumably has pronotal pseudosetae, yet apparently all students of Siphonaptera believe it is descended from combed fleas. However, the prothorax seem to bear seta-like apical spinelets as well (Traub & Evans, 1967), and the relationship between these and true pseudosetae, or the structure resembling pseudosetae in this species, requires further study. If there are no true psuedosetae on the pronotum and the spinelets Tepresent vestigial spines, Jordon may be correct. 362 R. TRAUB tion between temperature and development of ctenidia’. This is an interesting possibility and more information is needed before it can be evaluated. However, data at hand lead me to question it. None of the fleas collected by us in Arctic terrain at 12000 ft elevation in the Himalayas of West Pakistan, where the ground is frozen 9 months of the year and where it may snow in summers (Traub & Evans, 1967), showed any such tendency. Thus, in species like Ctenophthalmus golovi loff & Tiflov, 1930 (i.3) and Frontopsylla elata (Jordan & Rothschild, 1915) (iv.2), the combs are similar to allied forms, instead of being reduced, although it is true that these species are also found in less frigid climes and hence may not be valid examples. However, Monopsyllus thambus (Jordan, 1929) (iii.1) is a flea of the far north and of subartic heights in the Rocky Mountains, and it can be seen by comparing fig. 57 with that of fig. 59 (M. wagneri, another Peromyscus-flea, but a temperate one), that there is no reduction in the comb. Alaskan and other northern fleas show no loss regarding number or size of spines unless they are nest-fleas. Perhaps there is more reason to believe that the reduction of the comb seen in Mioctenopsylla and Glaciopsyllus° Smit & Dunnet, 1962 is correlated with the fact that other fleas of sea-birds lack combs, viz. Notiopsylla (ii.1), Actenopsylla (ix.1), Ornithopsylla (ix.1), 3 species of Xenopsylla (ix.t) and Parapsyllus Enderlein, 1903 (iv.2).1 Further, Notiopsylla is highly exceptional amongst pygiopsyllids in not bearing ctenidia, while Actenopsylla and Ornithopsylla, spillopsyllines, are the only members of their subfamily with this trait. However, no Xenopsylla bear combs and no rhopalopsyllid (to which family Pavapsyllus belongs) has a true ctenidium either. As indicated above, I believe the reduction of the comb seen in these bird-fleas is associated with their being primarily nest-fleas. J. STRUCTURAL AND BEHAVIOURAL VARIETY WITHIN A UNIFORM ORDER OF INSECTS The Order Siphonaptera is so distinctive and basically uniform that Snodgrass (1946) pointed out that even anysmall piece of the exoskeleton of a flea can be prompt- ly recognized without question. Yet, within this overall plan, believed to be of undoubted monophyletic origin (Jordan, 1947), much more diversity exists in behaviour and structure than is generally appreciated. There are the hypodermal Tunginae like Tunga, which are so specialized that the tiny females penetrate into the superficial layers of the skin of the host and ultimately lie almost completely within a cyst, with just the apex of the abdomen protruding. Here the flea may remain for weeks, engorging on blood repeatedly, and as the abdomen fills with dozens of eggs, the females may reach the size of a pea, extruding eggs to the outside world intermittently. The sticktight fleas (other Tunginae and the pulicine Echid- nophaga) act in a somewhat similar way but remain external, anchored by long serrate mouthparts. In both groups combs of spines are absent, and the vestiture 11Pavapsyllus is a member of the Rhopalopsyllidae, all of which lack true combs, and hence the ab- sence of a comb in this genus may have nothing to do with the parasitism of sea-birds. On the whole, the family infests rodents and is essentially Neotropical, but one other genus may also be associated with sea-birds, namely the Argentinian Listronius Jordan, 1942, which includes 3 species all of which are known only from 1 or 2 specimens. One of these species, L. robertsianus (Jordan, 1939), has been found only in the Falkland Islands, on two occasions, once in the burrow of sea-birds and once on an ornithologist. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 363 reduced in general; the thorax is extremely narrow and telescoped; the head very large and angulate, so that it may be pressed against the host as the flea attaches; while the abdomen is capable of marked extension. Others like the pulicine rabbit-fleas are sessile only for certain periods and move about occasionally to a new site or to move onto the young of the host. The vermi- psyllids and malacopsyllids, like the tungines, have a leathery integument and hence are capable of gross enlargement, but they are external ectoparasites and not as fixed. Other attributes of these fleas have already been mentioned. Many kinds of fleas are active on the host, and move about and feed frequently, like the familiar cat-fleas. These are fur-fleas, and possess combs of spines and rows of stiff backward projecting bristles to enable them to remain safely on the host. Variants on this theme, occur- ring throughout the Order, possess spiniforms or other modified types of bristles along or near the front margin of the head so that they can hook onto hairs while feeding or resting (Traub, 1968). Such “‘thorny-headed’’ fleas are further adapted in having the head flattened and in possessing stout procoxae, modifications which enable them to appress their heads against the body of the host while hooked onto the bases of hairs (Traub, 1968, 1969). Most of the above types of fleas have a well developed pleural arch and are good jumpers, enabling them to, at least, find a host before becoming fixed, or in the case of some of the fur-fleas, to locate a second one after leaving the first to lay eggs, etc. In the case of hypodermal or sticktight fleas with ready access to hosts, to which they remain affixed for life, the pleural arch is greatly reduced. Those of diurnal hosts have large eyes. In contrast, are the nest- fleas, marked by reduction of eyes, combs, chaetotaxy, and thorax, as we have seen, and which presumably glue their eggs to debris in the nest rather than deposit them loosely among the hairs of the host. It should be borne in mind that fleas which are closely related taxonomically may vary considerably with respect to habit (and, accordingly, in habitus, to some degree). This is at times true of species placed in the same genus, e.g. Chimaeropsylla Rothschild, 191 (xi.1) and also Ceratophyllus (i.t). As N. C. Rothschild (1g11) and Smit (1952) pointed out, Chimaeropsylla potis Rothschild, 1911 is modified as a sticktight flea, as indicated by the presence of broad, coarsely serrate laciniae and modifications in the head, in which the fronto- clypeal region is shortened and the genal region lengthened. With a head of this type, the anchoring mouthparts may be swung in position as to be almost horizontal when embedded in the host, and be remote from the maxillary palpi. In contrast, in Chim. haddowi Smit, 1952, as that author has shown, the head is rounded, the genal-frontoclypeal regions are of comparable size and the stylets are narrow, suggesting this species is not truly a sticktight flea. The amplitude of swing of the mouthparts of C. haddowi is too great for a normal free-living flea, however, and hence Smit believes that species is tending towards “a sedentary life’. M. Rothschild & Hinton (1968) provide interesting support for Smit’s belief that C. potis is a sticktight flea but C. haddowi is not, and this evidence is based upon their _ demonstration of a significant correlation between special adhesive discs on the antenna of male fleas and the degree of fixation of the female. During copulation, males of most species grasp the abdomen of the female by means of their elevated, _ long antennae, and Rothschild & Hinton believe that the discs help restrain the 364 R. TRAUB female during the act. These adhesive holding-structures have been noted in all but one of the recognized families of fleas (but not necessarily in all representatives thereof), and they are 4pparently well developed in species in which the females are active and mobile at the time of pairing. In the sessile fleas whose mating-habits have been observed, either the male antennae cannot be employed to grasp the female because of their structure and position, or else they lack the adhesive discs. Thus, in the European rabbit-flea, Spilopsyllus cuniculi (ix.t) the males grasp the females by their antennae while mating, but they lack the adhesive discs, as do the related species of the New World rabbit-fleas of the genus Cediopsylla Jordan, 1925. It is noteworthy that these female rabbit-fleas are essentially sessile at the time of mating. What is particularly relevant to the case of Chimaeropsylla is that the adhesive discs are lacking in C. potis but present in C. haddowi, as Rothchild & Hinton point out. As M. Rothschild has stated (im litt.) the convergence between Spilopsyllus (and Cediopsylla as well) and C. potis is quite striking. In all three taxa, the frons is angled, the laciniae are stout and markedly serrate and the cuticula is dorsally incrassate and at times leathery elsewhere. All of these features are often seen in sticktight fleas. The 3 genera also agree in that both genal and prothoracic ctenidia are present, the eye is relatively large and the pleural arch is well developed. The last two features are characteristic of fleas of active diurnal hosts, and the chunky, stout body seen in these genera also suggests species that must seek such a host, just as the ctenidia recall the need to remain thereon once it is found, before the fleas become semi-sessile. Fleas of the genus Ceratophyllus are particularly instructive in that marked behavioural and structural differences may occur even amongst species parasitizing the same kind of host, and hence the swallow-fleas of this genus are discussed in some detail. Darskaya (1954b, 1959, I964a) in a series of interesting articles on the subject, has classified the Palaearctic ceratophyllid bird-fleas into two ecological groups on the basis of the correlation of the behaviour of the fleas with the habits of the particular birds they parasitize. On the one hand are the “‘settled’’ fleas, which have a rela- tively pronounced host-specificity and can survive long periods of starvation in the nest after the birds have reared their young in the spring and have all departed. These are epitomized by the swallow-nest fleas which are poor jumpers and do not emigrate. Since the hosts of these fleas seldom alight, and even feed while flying, the inability of these fleas to leap well, and their disinterest in leaving the nest burrows, are adaptive advantages. If fleas of this class ever did fall to the ground, away from the nest-site, they would have virtually no chance of finding their specific hosts nor of returning to the nest. In the spring, however, some of these fleas may be carried by the birds from one nest or colony to another, before the birds settle down for nesting, but this particular kind of flea does not remain on the host for long periods. During the spring and summer, the “settled” fleas generally produce only one generation. Possession of a very large fat-body, with stored nutriment, enables them to survive the long period (as much as 10 months) while they are nest-bound in the absence of the birds. During the winter, their body-functions are slowed as the CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 305 fleas remain inactive in cracks in the nest. With the onset of warm weather in the spring, and the annual return of the birds, the fleas become very active, suck blood from the host avidly and quickly and rapidly mate and produce eggs. The larval stage is accelerated so the life-cycle is completed while the birds are still in the nest, giving the new crop of adults time to engorge on blood before the long siege begins. Inasmuch as the young birds likewise develop rapidly, and these birds have only one brood a year, an abbreviated span for the reproductive and developmental stages is a necessity. Morphologically, the “‘settled” fleas are light in pigmentation or exhibit the characteristic banded appearance reported by Holland (1952) for C. arcuegens?. This is due to the anterior portions of the abdominal segments (both terga and sterna) being well sclerotized, or even especially so, while the posterior regions are lightly tanned. The former attribute offers protection to the body when the flea crawls into tight crevices in the nest; the latter permits gross enlargement of the abdomen as the fat-body enlarges. The alternating brown and light bands on the abdomen often enables one to recognize the specimens as bird-fleas even with the naked eye. This group of fleas is also characterized by relatively small (but never- theless well developed) eyes, reduction or loss of sclerotization of the metanotal flange or “‘collar’’ (permitting flexibility of the abdomen in crawling, and scope in enlarging) and reduction or loss of the pleural arch, with concomitant lessened ability to jump and increased facility to crawl into crevices. Many of these attri- butes have been shown above to be characteristic in general of nest-fleas, and as indicated by Holland (1952) and Darskaya (1964a, b). Examples of fleas of this type are C. delichoni°, which lacks the pleural arch and C. hirwndinis, in which it is present. In contrast to the “‘settled’” fleas are the bird-fleas which Darskaya calls “nomadic” or “migrating”’ (1954a, 1959, 1964a). (Perhaps in English, ‘““‘perambula- tory’ may be a better term, avoiding the connotation of a fixed direction or a return, and yet indicate a purpose for the movement.) In this category are C. garet Rothschild, 1902 and C. gallinae (Schrank, 1803) which usually parasitize birds which do not return to utilize nests of the previous year, and hence any fleas which had hatched out in such a nest are faced with the problem of finding a bird to infest. This is solved by leaving the nest and actively seeking a host. C. styx (whose host, the sand-martin, generally does return to its old nest-site in the spring) leave the burrows in which they were born and may travel as much as 50 feet to another and new nest-site, “recognize’’ it, and settle there (Bates, 1962; M. Rothschild, 1969). This species, which is an excellent jumper and can span 200 mm (M. Rothschild, 1969), has the facility of immediately leaping onto a bird which hovers for a few seconds near the nest, and it seems likely that other bird-fleas can act similarly. Perambulatory fleas usually have the ability to feed on a variety of hosts, enabling them to effectively parasitize nearly any bird they find. It is not surprising that fleas which have to be highly mobile to survive are: 1. excellent jumpers and climbers and fast walkers; 2. have large eyes; 3. well developed thoraces, including a large pleural arch; and 4. are almost black in colour due to the heavy tanning of the cuticle. The ‘nomadic’ fleas overwinter in the cocoons and must feed soon 366 R. TRAUB after emergence or starve. The fat-body is small and is rapidly consumed during the summer, and the fleas feed frequently and quickly. Several generations are produced a year, since the hosts also tend to raise more than one litter annually. The flea-population in such nests can build up to enormous proportions. Darskaya’s notable observations were necessarily based upon study of a limited number of species of Ceratophyllus in the U.S.S.R. and dealt with a variety of complex and little-known phenomena. Even so, her generalizations, in the main, apply well to bird-fleas trom other parts of the world and to other taxa, as will be shown shortly. As is to be expected, however, some species that have certain of the attributes of “nomadic’”’ fleas exhibit a pattern of behaviour somewhat different from that of C. garei, etc. For example, they conform with respect to the morpho- logical features noted above and in that these fleas appear to spend little or no time in the nests as adults, and actively seek their hosts. These fleas, however, apparently spend most of their adult lives on the birds themselves and hence, as pointed out by M. Rothschild (im ltt.) are analagous to the “‘body-fleas” or “fur- fleas’ of mammals. Such fleas presumably can accompany a bird on migrations of thousands of miles. Thus, the Holarctic Dasypsyllus gallinulae (Dale, 1878) (11.1) has not only been found in North America and Europe but in such extremes as Panama and the Far North, and in regions as distant as the Philippines (Traub, 1950b), North Borneo and Malaya. I was surprised to collect the “arctic’’ species Dasypsyllus stejnegeri (Jordan, 1929) in central Mexico, but Smit (1961) reported it from the Falkland Islands! The Panama records of D. gallinulae were based upon specimens Dr. H. C. Matthes collected from a sudden infestation on his own body, while walking in the forest of Mt. Volcan. It therefore seems likely that this is a perambulatory species. In all probability, other bird-fleas, whose habits have not yet received the type of intensive study undertaken by Darskaya for Palaearctic Ceratophyllus, could be divided into similar ecologic groups. The Antarctic flea, Glaciopsyllus° and the spilopsylline fleas of sea-birds are in the category of “‘settled” fleas, as indicated above. Lentistivalius insolli (ii.1) with its brown and white banded appearance, relatively small eye and records of collections from nests rather than birds, presum- ably is in this category. Fleas of the genus Hoogstraalia, which possess a genal ctenidium, and are darker in colour, etc., may be “perambulatory”’ fleas. Although the ecology of rodent-fleas has not been studied as well as that of bird- fleas, because of the inherent difficulties, a great deal of fine work has been done in the Soviet Union on this subject. It is apparent that the behaviour of the rodent-fleas is as specialized and complex, and as inter-related with the habits of the host as the bird-fleas mentioned above. For example, several species of fleas with well devel- oped eyes have been noted to swarm to and about the burrow-opening, pursue prospective hosts or ‘“‘emigrate’’ into new burrows if their birth-places had been abandoned by the rodents (Darskaya, 1954a, 1955, 1964b; Loff, 1941). K. THE PRINCIPLE OF STRUCTURAL COMPENSATION It has been pointed out that in Siphonaptera the total mass of two or more structures that are associated in a common function generally is a theoretical con- CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 367 stant (Traub, 1969). For example, the claspers, which operate as a unit in grasping the female in copulation, may be of approximately equal size, or else the movable finger (F.) may be enlarged while the immovable process (P.) is small, or thirdly, F. may be reduced and P. huge. It was also shown that the relative size of Ford’s sclerite varies inversely with the comparative degree of development of the crochets or of ventral paramere-like processes. In other words, although the size of the individual components may vary, they do so inversely, in a correlated manner, and, in effect, the total amount of tissue-substance involved is the same, and the over-all functional result is apparently not impaired. The specimens collected by Lord Medway’s team on Gunong Benom, and the other material discussed in the previous article in this series (Traub, 1972a), also provideinteresting examples of this principle. Particularly relevant are the unusual specializations cited and illustrated for each sex of Stivalius cognatus Jordan & Rothschild, 1922 (a member of Stivalius Jordan & Rothschild, 1922 s. stv.). It will be recalled that in the previous article it was emphasized how the sclerotized inner tube (S.I.T.) is characteristically elongated in that taxon and that Ford’s sclerite (F.SC.) is greatly enlarged, while, in contrast, the paramere-like ventral process of the crochet is vestigial or totally absent. The situation is very different in most members of Stivalius s. lat., which have well developed ventral crochet-processes (CR.P.), and in which S.1.T. is short and broad in comparison, i.e. Medwayella and Lentistivalius, as figured in that article. It was therefore suggested that the hypertrophied Ford’s sclerite and highly elongate S.I.T. serve to replace, in function, the ventral “‘parameres,’’ which have been lost in the course of evolution. That the reduction of CR.P. and concomitant modifications of S.LT. and of F.SC. are in accord with this principle of structural compensation is further suggested by developments in other members of Strvalius s. lat., viz. S. jacobsoni Jordan & Rothschild, 1922 and Parastivalius novaeguineae (Rothschild, 1904). These two are in the minority of ‘‘Stivalius’’ in which the aedeagus lacks ven- tral caudad-directed, finger-like or rectangular processes, and in each case there is a unique alternate device that appears as an apparent substitute. Thus, S. jacobsoni is one of the few species in which the very apex of the dorsal wall of the end-chamber of the aedeagus is highly modified, in this case being produced into 2 highly acum- inate fang-like projections. InP. novaeguineae, the ventral armature of S.1.T. is not only displaced far caudad, but it is hook-like suggesting a crochet in appearance. If the principle of structural compensation is a sound one, then it would be expected that there would be intermediate species in which the cooperating members of the functional device were only partially modified, e.g. there would be varying degrees of development of alternate mechanisms. S. phoberus (Rothschild, 1904) is an example of such an intermediate form, since, as has been shown in the previous article in this series, S.I.T. is not nearly as elongate as in other members of the genus (Stivalius s. str.), and the vestige of the crochet-process (CR.P.) is longer than in its allies. What then compensates for the loss of the functional part of CR.P., the ventral caudad-directed “‘parameres’’? It is noteworthy in this regard that several | of the associated sclerites seem more highly developed in S. phoberus than in related fleas with the ophidian-like S.I.T. Thus: (1) F.SC. bears 2 apical hooks per side, notI. (2) S.I.T. has a longer and broader apical fistula (FIS.) which is tanned to a 368 R. TRAUB greater degree than in S. cognatus, etc. FIS. in S. phoberus also appears to have more discreet tanned borders within S.1.T. (and may prove to be protrusible, at least here, if not throughout the genus). Another relevant example of the principle of compensation is provided by the metasternal furca, as has been indicated by M. Rothschild (cn Jitt.). In fleas which have lost the pleural arch, the furca is hypertrophied, as in the case of Wenzella® (fig. 85, I.F.-3) and Conorhinopsylla°® (fig. 86). This tendency is even exhibited in members of the same genus which differ with respect to the development of the pleural arch and ridge, e.g. Opisodasys and Ceratophyllus, and is presumably asso- ciated with the attachment of muscles which ordinarily fasten to the pleural ridge when the arch is present. The principle of compensation actually seems to apply to more than adjacent or cooperating structures or organs, for the extreme modifications of the hypodermal and sessile fleas (Tunginae) presumably fit the concept. Here the head and mouth- parts have become enlarged or greatly strengthened, and the abdomen capable of enlarging enormously, while the thorax has become so tremendously compressed that it is difficult to discern its components. The Principle of Structural Compensation is a corollary of the Principle of Totality of Mass (Traub, 1969), which states that while the components of a structure may vary in size or mass, the over-all size is a relative “constant”. Thus, in some species the pronotal spines are long, while the pronotum is narrow, as in figs 18 and 21; in others, the reverse is true (fig. 63). The latter principle is well illustrated by a species treated above as an example of a flea undergoing reduction of ctenidia, — namely Tvichopsylloides oregonensis®. Here the genal ctenidium is usually entirely absent, or else is represented by but 1 or 2 tiny pale spines. It is noteworthy that the gena itself is unusually large and overlaps the base of the procoxa. It thus encompasses the same area as does the fully developed comb of the related genus Rhadinopsylla°. V. CONCLUSIONS Fleas are far more specialized for their particular mode of life than had been rea- lized a few years ago, not only with respect to their behaviour and physiological responses, but also concerning their morphology. All of these phenomena are | intimately related to corresponding features of their host and bespeak of an ancient | and intimate association between parasite and host, even to the point of evolving together, as witness the fit of the pronotal combs of some fleas to the hairs of the mammal they infest. In other instances, the pronotal comb and structure of the thorax and degree of development of the eye, reflect the environment and habits of the host and the corresponding modifications of the flea thereto. | The adaptive responses of the Siphonapteran pronotal comb to the pattern of the hairs of the host, and the association between the structure of the metathorax and the advantages or disadvantages of the ability to leap, are examples of the moulding- influence of the environment, which tends to result in uniformity of physiognomy, | despite the fact that different mechanisms may have been employed to achieve _ these ends. Such modifications in fleas therefore parallel the remarkable conver- CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 369 gences seen almost everywhere in nature, viz., the amazing similarity in appearance of carnivorous marsupials and the placentate Carnivora even though the two represent two independent lines of evolution. It is apparent that there still is a great deal to be learned about some of the prin- ciples involved in the examples of convergent evolution and adaptations mentioned, to say nothing of the details. Some of the many unanswered questions could be resolved if more were known about the flea-fauna of South America, Australia and the Australo-Indonesian Archipelago, and about host-relationships in general. Even more, we require data on the ecology of fleas and mammals and birds, and study of living material. VI. ACKNOWLEDGEMENTS This study was supported by Grant No. AlI-04242 of the National Institutes of Health, Bethesda, Maryland, U.S.A., with the Department of Microbiology, University of Maryland School of Medicine, Baltimore, Maryland. Various portions of this work were discussed with Dr. Miriam Rothschild, Oxford, England, to whom I express thanks for worthy suggestions, inspiration and en- couragement. Dr. Rothschild and Dr. George P. Holland of the Entomology Research Institute, Department of Agriculture, Ottawa, Canada, kindly reviewed the manuscript and made useful comments, which I greatly appreciate. I am also indebted to F. G. A. M. Smit of the British Museum (Natural History) at Tring for preparing and sending me a sketch of the pleural arch of Praopsylla powelli Ingram, 1927, an important species which was not available for study here. The bulk of the photographs, including all the figures of heads and thoraces, were prepared by Suzanne F. Shipley, of our Department, who also touched up the preliminary photographs of some of the pronotal combs made by Julio Navarro, formerly of our staff. I am indebted to those artists for their painstaking work. Considerable editorial assistance was rendered by Helle Starcke, also of our Department. VII. LIST OF TAXA CITED IN TEXT Code Family Code Subfamily Te Hystrichopsyllidae I Hystrichopsyllinae Anomiopsyllinae Ctenophthalminae Dinopsyllinae Doratopsyllinae Listropsyllinae Neopsyllinae Rhadinopsyllinae Stenoponiinae Pygiopsyllinae Lycopsyllinae Ceratophyllinae Foxellinae ii. Pygiopsyllidae iii. Ceratophyllidae NHN HO ON AUNHRW DN R. TRAUB Leptopsyllidae I Leptopsyllinae 2 Amphipsyllinae Ischnopsyllidae I Ischnopsyllinae 2 Thaumapsyllinae Rhopalopsyllidae I Rhopalopsyllinae 2 Parapsyllinae Pulicidae it Pulicinae 2 Tunginae Vermipsyllidae —— Chimaeropsyllidae I Chimaeropsyllinae 2 Chiastopsyllinae Coptopsyllidae a= Malacopsyllidae -— Xiphiopsyllidae = Genus or Species Acanthopsylla Jordan & Rothschild, 1922 . Acropsylla Rothschild, 1911 - girshami Traub, 1950 Actenopsylla Jordan & Rothschild, 1923 3 A suavis Jordan & Rothschild, 1923 . Aengmopsylla grodekovt Sychevsky, 1950 . °Agastopsylla Jordan & Rothschild, 1923 Amphipsylla Wagner, 1909. 5 marikovskit loft & Tiflov, 1939 rr montana Argyropulo, 1946 °“Anomiopsyllus Baker, 1g04. Aphropsylla Jordan, 1932 : °Araeopsylla Jordan & Rothschild, 1921 Archaeopsylla Dampf, 1908 °Brachyctenonotus Wagner, 1928 ! Bradiopsylla Jordan & Rothschild, 1922 9 echidnae (Denny, 1843) . Brevictenidia Liu & Li, 1965 : aS mikulint (Schwarz, 1960) Caenopsylla Rothschild, 1909 9 laptevi Mikul n & Zagniborodova, 1958 °Calceopsylla Liu, Wu & Wang, 1965 . : °Callistopsyllus Jordan & Rothschild, 1915 . Callopsylla Wagner, 1934 i dolabris (Jordan & Rothschild, ‘TorI) Catallagia Rothschild, 1915 : : +i charlottensis (Baker, 1898) a striata Scalon, 1950 . Page 339, 342 351 2 F50 328, 362 "328, 342, 348, Fo9 314, F9 325, 333, 352 327, 331, 347 327, 333, F1oo, Froz, F1ro2 327, F103, Fro4, Fr05 324, 329, 333, 345, 352, 354, 355 354 333 : 320 327, 333, 352 341 320 341, 352 327 331 ‘ 3 F71 327, 333, 352 324, 333, 335 j i 356 326, 356, F84 324, 333 321, F67 F8r CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 371 Cediopsylla Jordan, 1925. ‘ : ; : : ‘ : 364 Centetipsylla Jordan, 1925 ; : : 320 Ceratophyllus Curtis, 1832 : , 326, 334, 355, 363, 364, 366, 368 x arcuegens Holland, 1952 317, 326, 327, 333, 334, 346, 348, 365, Foo g caliotes Jordan, 1937. : ; ; 326 a delichoni Nordberg, 1935. ; 326, 346, 348, 365 gallinae (Schrank, 1803) : ; ; 349, 355, 365 garet Rothschild, 1902 . : : : 346, 365, 366 ; hirundinus (Curtis, 1826) : ; : 346, 365 a orites Jordan, 1937 x. 5 : : ‘ , 326 vusticus Wagner, 1903 . : ; : : ‘ 326 ; scopulorum Holland, 1952. 3 346 5s styx Rothschild, tg00 . 317, 326, 327, 344, "348, 365, For Chaetopsylla Kohaut, 1903 ; : : : 359 Chiastopsylla Rothschild, 1910 . 2 : : , 3 : 329 rf numae (Rothschild, 1904). F ; : : 329 Py pitchfordi Ingram, 1927 : ‘ : ; ; 329 quadrisetis de Meillon, 1930 . ; ; ; 3 320 C himaeropsylla Rothschild, 1911 : ; é : : 363, 364 haddowit Smit, 1952 . : : : ; 363, 364 potis Rothschild, ro11z ¢ : : : 363, 364 Choristopsylla Jordan & Rothschild, 1922 . 2 ‘ : : 341 s ocht (Rothschild, 1904) ; : : : : 341 a if tristis (Rothschild, 1900) : : 325, 334, 341 Citellophilus Wagner, 1934 ; ; : ; 3 ; ; 356 5p tesquorum (Wagner, 1898) . : F 322, 349 “Conorhinopsylla Stewart, 1930 . : 324, 329, 333, 334, 352, 355, 368 5 nidicola Jellison, 1945 : ; : 4 321, F86 3 stanfordi Stewart, 1930. : d 340, 345, F73 Coptopsylla Jordan & Rothschild, 1908 : : : é 332 Corrodopsylla Wagner, 1929. : , : : 347 “Corypstila C. Fox, 1908. ; 3 : 332, 333, 336, 337, 351 i ornata C. Fox, 1908. : : : ; 3 352 Cryptopsylla de Meillon, 1949 . : : : : 329, 331, 332 Ctenidiosomus Jordan, 1931. F 351 Ctenocephalides Stiles & Collins, 1930 326, 337, 340, 350, 354, 357, 358 nt canis (Curtis, 1826) . : : 357 felis (Bouché, 1835) . : : : : 357 Ctenophthalmus Kolenati, 1856. E "333, 316, 347 3 (Ctenophthalmus) a. agyrtes (Heller, 1896) : é F22 is bisoctodentatus Kolenati, 1863 F24, Tx 5p (Ethioctenophthalmus) calceatus Waterston, 1912 . 347 s (Alloctenus) cryptotis Traub & Barrera, 1966 : F25 3) (Neoctenophthalmus) dilatatus Wagner, 1928 ; 318 ix.I v1.2 1) 1.7 187) eg) ey 11.2 1V.2 1V.2 ey) i.7 1.7 ili.I 1.1 1X.2 iil. ii. R: TRAUB Ctenophthalmus (Nearctoctenophthalmus) expansus myodosus Traub & Barrera, 1966. ; : F2r (Medioctenophthalmus) golovi loft & Tidey 1930 : 362 (Nearctoctenophthalmus) haagi Traub, 1950 . ; F26 (Euctenophthalmus) 1. levanticus Lewis, 1964 j F28 . pollex Wagner & loff, 1926 3135322, Baa alee (Palaeoctenophthalmus) rettigi Rothschild, 1908 _ . B27 (Spalacotenophthalmus) spalacis Jordan & Rothschild, ro1r_ . : : : : 347 Ctenophyllus Wagner, 1927. : ; ‘ ‘ ‘ F 331 Dactylopsylla Jordan, 1929 : : F : : ‘ 331, 346 Dasypsyllus Baker, 1905 . : ‘ : : : : 342 gallinulae (Dale, 1878) : ‘ 2 : : F 366 steynegeri (Jordan, 1929) : : A : : 366 Delotelis Jordan, 1937. : : : : ; 324, 333 telegont (Rothschild, 1905) j : ‘ : : 321, F74 Diamanus Jordan, 1933 . : ; F : ; 356 Dinopsyllus Jordan & Rothschild, 1913 ‘ : : 340 ” (Cryptocten psyllus) ingens (Rothschild, 1900) . 331, 346 Dolichopsyllus stylosus (Baker, 1904). , : : : 343 Doratopsylla Jordan & Rothschild, 1912. : ‘ : 347, 351 Dorcadia Ioff, 1946. ; ‘ : : ; ‘ : 360 dorcadia (Rothschild, 1912) ‘ : : : ‘ 360 Dysmicus barrerai (Jordan, 1939). F : : : 329 Echidnophaga Oliff, 1866 . ; : : , 337, 338, 357, 362 ; gallinacea (Westwood, 1875) ‘ : : : 357 ro oschanini Wagner, 1930 F : : : : 359 Ectinorus polymerus Jordan, 1942. : - 5 : ‘ 329 *Eopsylla Argyropulo, 1946 ; : ‘ : : 324, 329, 333 Epitedia Jordan, 1938. : ; “ ¢ - 324, 330, 347, 355 cavernicola Traub, 1957. F . ‘ : : F69 faceta (Rothschild, 1915). : , ; 333, 340, F77 stanfordi Traub, 1944. : : ; : : ; F98 Foxella Wagner, 1929. : : : : : : 346 Frontopsylla (Orfrontia) loff, 1946 ih (Frontopsylla) elata (Jordan & Rothschild, 1915) F 362 Genoneopsylla Wu, Wu & Liu, 1966 . : : : ; 324, 333 longisetosa Wu, Wu & Liu, 1966 . ; ; : 324 & thysanota (Traub, 1968) ‘ F : ; 324, F70 Glasopsyiius Smit & Dunnet, 1962 . : j ; 362, 306 bi antarcticus Smit & Dunnet, 1962 . 326, 333, 334, 330, 361 Hectopsylla Frauenfeld, 1860. ; ; : 5 : 333 Eolas sipli Traub, 1953. ; . : : ; 326, 333 7" neali Traub, 1953S ; ; : : ; 339 lv.2 CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 373 Hoogstraalia Traub, 1950 : ; ; , ‘ 348, 350, 366 Hopkinsipsylla Traub, 1963. ‘ é : 6 3 327, 330 Hys'vichopsylla Taschenberg, 1880 . ; : ; : 351, 358 5 schefferi Chapin, 1919 ; : : : , 343 talpae (Curtis, ha : : ; : , 347 °Idilla cats 1957. : : : ‘ j : 325, 333 Ischnopsyllus Westwood, 1833 . 5 : : : : ; 333 Jellisonia Traub, 1944. : ‘ : : : 331, 351 Jordanopsylla Traub & Tipton, 1951. ; : : : 324, 330 Kohlsia Traub, 1950 : : : : : : 316, 331, 351 » gammonst Traub, 1950 ; \ : : : ; 316 » graphis (Rothschild, 1909) . ; : : ; 316, Fro » _ keenani Tipton & Mendez, 1961. : : : : 339 “Lagaropsylla Jordan & Rothschild, TO2T ls ; : : ; 333 Leptopsylla Jordan & Rothschild, TOI é : : 350 * (Pectinoctenus) pamirensis (Ioff, 1946) : 5 3 Fo3 Lentistivalius Traub, 1972 : ‘ 367, 338, 312 is aestivalis (Jameson & Sakari 1954) ‘ : 312, Fig 33 alienus (Smit, 1958). . ‘ ‘ 312, 314, F16 3 ferinus (Rothschild, 1908). : . 312, Fr5 - insolli (Traub, 1950). 4 ‘ : here, 342, 366, F18 5 vomerus Traub, 1972 . : ‘ ‘ : 312, Fx7 Libya tus Jordan, 1936 . : 3 - 326, 332, 338, 340 is infestus (Rothschild, 1908) : : i ‘ 314, F13 Listronius Jordan, 1942 . : 3 ‘ 3 : 5 362 55 vobertsianus (Jordan, 1939) : ; F ; é 362 Lycopsylla Rothschild, 1904. : ‘ 5 : 5 325, 326 Macrostylophora Ewing, 1929 . 5 ; : : 316, 347 3 borneensis (Jordan, 1926) . : 319, 339, F96 PF Jimbriata (Jordan & Rothschild, 1921) - 339, F7, Fo7 h. hastata (Jordan & Rothschild, 1921) 314, 319, F1r4 Malacopsylla Weyenbergh, 188r , : : 5 352, 361 a gvossiventris AUeuesiet 1870) : 3 : ‘ 361 Malaraeus Jordan, 1933 . 3 ‘ : : 351 Medwayella Traub, 1972 . ‘ . : 312, 316, 359, 367 % robinsont (Rothschild, 1905) A : eSr4, Onsen. Megabothris acerbus (Jordan, 1925) . : ; : : : Ise ~ quirint (Rothschild, 1905) : ; : alia “Megarthroglossus Jordan & Rothschild, I9QI5 324, 333, 335, 340, 352, 355 5s divisus exsecatus Wagner, 1936 : i F4 4 procus Jordan & Rothschild, 1915. : : F75 Meringis Jordan, 1937. é ; ‘ 324, 333 Mesopsylla Dampf, r9I0 . : F ‘ ; 3 5 é 347 Metastivalius Holland, 1969. : : : : 5 331 Mioctenopsylla Rothschild, 1922 F ; ‘ ; " 352, 361, 362 R. TRAUB Mioctenopsylla arctica Rothschild, 1922 Monopsyllus Kolenati, 1857 : e anisus (Rothschild, 1907) A argus (Rothschild, 1908) 3 ciliatus protinus (Jordan, 1929) 5 e. eumolpi (Rothschild, 1905) . 5 eutamiadis Augustson, 1941 . is extlis (Jordan, 1937) 3 fornacis Jordan, 1937 5 indages (Rothschild, 1908) 3 sciurorum (Schrank, 1803) 9 tamias (Wagner, 1927) . 53 thambus (Jordan, 1920) . tolli (Wagner, I90T) 3 vison (Baker, 1904) wagnert (Baker, 1904) Muesebeckella Traub, 1969 : Myodopsylla Jordan & Rothschild, Torr °“Myoxopsylla Wagner, 1927. Nearctopsylla Rothschild, 1915. Neopsylla Wagner, 1903 . ; - 3 bidentatiformis (Wagner, 1893) si dispar Jordan, 1932 . . hissarica Toff & Sosnina, 1950 eo inopina Rothschild, 1915 . i luma Traub, 1954 setosa (Wagner, 1898) Nosopsyllus Jordan, 1933 5 ceylonensis Smit, 1953 5 (Nosopsyllus) 1. londiniensis (Rothschild, 1903) . Notiopsylla Jordan & Rothschild, 1914 29 enciart Smit, 1957 . i kerguelensis (ieecheabere! 1880) DE SES Oudemans, 1906 eusarca (Dampf, 1908) Ophithalmopsylla Wagner & Ioff, 1926 eS (Evemedosa) celata Traub, 1965 . = (Cystipsylla) jettmari Jordan, 1929 Optsocrostis Jordan, 1933 Opisodasys Jordan, 1933 . 3 enoplus (Rothschild, 1909) 5 hollandi Traub, 1947 is keent (Baker, 1806). ii nesiotus Augustson, 1941. 5 3 pseudarctomys (Baker, 1904) 314, 317, 324, 333) 347, 351, 354 ‘ 342 313, 3 316, 317, 339, 355 ; For 339, F58 3 F62 Abe : F65 F55, Tr F66 2 F56 339, 355, F6o . : 355 . 362, F57, Dx ; F63 330, Fi, F64, Tr 321, 362, F59 339 : ; 318 326, 333, 339 325, 333 ; F4r 314, F4o : F42 F309, Tr F38, Tr 322, F37 316, 354 Pry ie F49, Tr 325, 302). : 326 325, 348 . : 333 : 318, 345, 352, F89 . . . 347 328, 330, Fo4 328, F95 350, 356 314, 317, 326, 332, 339, 341, 342, 368 339, F48 317, 327, 330, 344, F44, F87, Tx 314, 321, F43, Tr . Ee 314, 326, 333, 339, 47 Thbkae iii.I iii. iv.2 ix.I fixe: iv.I iv.2 iv.2 ix.I iv. iv.2 iv.2 iii.I CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 375 °Opisodasys robustus (Jordan, 1925) F5, F46 si vesperalis (Jordan, 1929) 317, 326, 327, 333, 339, 341, 344, F88 Orchopeas Jordan, 1933 na bolivart Barrera, 1955 5 caedens (Jordan, 1925) fullert Traub, 1950 . Pe howardi (Baker, 1895) oa leucopus (Baker, 1904) 5 neotomae Augustson, 1943. 5 n. nepos (Rothschild, 1905) a sexdentatus (Baker, 1904) . sexdentatus agilis (Rothschild, 1905) Biaihophasa Mikulin, 1956 Ornithopsylla Rothschild, 1908 . me laetitiae Rothschild, 1908 Oropsylla Wagner & Ioff, 1926. Oxyparius Jordan, 1936 . Palaeopsylla Wagner, 1903 Papuapsylla Holland, 1969 Paraceras Wagner, 1916 . Paractenopsyllus Wagner, 1938. : Paradoxopsyllus microphthalmus Loff, 1946. Paraneopsylla Tiflov, 1937 5 Parapsyllus Enderlein, 1903 Parastivalius novaeguineae (Rothschild, 1904) faa loceras Ewing, 1940 oregonensis Ewing, 1940 Pariodontis Jordan & Rothschild, 1908 Peromyscopsylla I. Fox, 1939 Phaenopsylla Jordan, 1944 : m kopetdag loff, 1946 Phalacropsylla Rothschild, 1915 paradisea Rothschild, 1915 . Phthiropsylla Wagner, 1939 : Pleochaetis Jordan, 1933 . i mathesoni Traub, 1950 ‘ paramundus Traub, 1950. 3 ponst Barrera, 1955 5 soberont Barrera, 1958 Praopsylla Ingram, 1927 . 45 powelli Ingram, 1927 Pulex Linnaeus, 1758 » s-noculus Traub, 1950 °Pygiopsylla celebensis Ewing, 1924 °Rhadinopsylla Jordan & Rothschild, 1912 324, 333, 344, 352, 354, 355, 308 ie 317, 320, 339, 359 . 339, F35 = 833 ‘ F31 358, F2, Tx ~ 344, 36; ir 320, 321, F32, Tr : B29; 0r 314, 320, 321, F34 321, F30 342, 346 328, 362 328 349, 356 . 333 - 337) 343, 347 319 316, 349 : 351 327, F82 324, 333 329, 362 367 333 343 . : 349 - 331, 350, 351 327, 328 : , F8o0 - 324, 330, 333, 352 321, F78 352, 361 316, 321 : ‘dae 321, 326, 327, 330, F68 : : 339 3 Sey, Ie, Wa 329, 331 : 369 - 345, 349, 358 346 334 R. TRAUB ' °Rhadinopsylla mexicana (Barrera, 1952) . 3 F ‘ ; F72 Rhynchopsyllus Haller, 1880. : : , : é : 333 Rothschildiana Smit, 1952 F ; ; F : - 324, 332, 35 Scolopsyllus Méndez, 1968 é é ‘ : : é , 340 Sigmactenus Traub, 1950 . : ‘ ; ; 5 ; 347, 350 7" alticola Traub, 1954 ; , : . : ; F54 Smitella Traub, 1968 : 3 : : : ; j 340, 350 Spilopsyllus Baker, 1905 . : : ; 5 : 3 364 sh cuniculi (Dale, 1878) ; : ‘ 337, 364 °Stenischia Jordan, 1932 . f : , 324, 325, 332, 333, 336, 337 °Stenistomera Rothschild, 1915 . ; : » 9324; 331,333) 335 0ee : Ff alpina (Baker, 1895). ; : , : 331 S 5 hubbardi Egoscue, 1968 . : : : : 2 331 2 *, macrodactyla Good, 1942 é : : : ; 331 Stenoponia Jordan & Rothschild, rgrtr : ; ‘ - 351, 358, 359 Stivalius Jordan & Rothschild, 1922 . : ‘ ; 367 i c. cognatus Jordan & Rothschild, 922 . : F 367, 368 a c. bamus Traub, 1972 . ; : ; ; ; i 314 % c. spivamus Jordan, 1926. é : : 2 F52 5 (Gryphopsylla) hopkinsi Traub. 1957 : : ; é F51 », jacobsont Jordan & Rothschild, 1922 . ; : : 367 5 klossi (Jordan & Rothschild, 1922) : : ‘ F20, Ta phoberus (Rothschild, 1904) ; : ; , 367, 368 Strepsylla Traub, 1950. F ; : : j - 324, 330, 333 a mina Traub, 1950. : : F F79 ene en opis Traub, 1950. : : 326, 332, 338, 340, 341, 347 st calceatus (Rothschild, ie) F : : 341, F3 Tamiophila Jordan, 1938. ; . 324, 330, 333, 355anu 5 grandis (Rothschild, 1902) ; ; ; : F760 TOSSES Wagner, 1927 : : 5 326, 332, 334, 338, 342 mf coloradensis (Baker, 1895) ; ‘ : : 339 - 3 5 octodecimdentata (Kolenati, 1863). ; : : F8 °Thaumapsylla Rothschild, 1907 : : z ; : 4 333 Thrassis Jordan, 1933. ; : 2 : : : 349, 350 Traubia Smit, 1953. : 3 : : : : : : 313 | EIA Ewing, 1938 ‘ ; j 5‘ : . 324, 333, 344 oregonensis Ewing, 1938 . E : : 343, 368 Tunga Tecee 1838 : : : j . 333, 337) 354, 362 | Vermipsylla Schimkewitsch, 1885 : ; : ; 6 : 360 Wagnerina loft and Argyropulo, 1934 : : : g 324 | % schelkovnikowi loft & Argyropulo, 1934 : : : F83 °Wenzella Traub, 1953. ‘ ; 324, 333, 345, 352, 354, 355» 308 | ae: obscura Traub, 1953 . ‘ 5 , 345, F85 Xenodaeria Jordan, 1932. ; : 332 Xenopsylla Glinkiewicz, 1907. 323, 328, 333, 342, 347, 358, 361, 362 | ae CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 377 bead Xenopsylla cheopis (Rothschild, 1903) 337, 357 xe 7 conformis (Wagner, 1903) 358 x 7% georycht (C. Fox, 1914) : : 347 1x. 3 gerbilli caspica loff, 1950. : : . 335, 350, 357 rea 59 gratiosa Jordan & Rothschild, 1923 . ; 342 ier | ° 5 papuensis (Jordan, 1933) 317, 318, 328, 333, 344, 350, Fo2 1X.I . trispinus Waterston, IgII ; 342 bene vexabtlis Jordan, 1925 317, 308, 328, 344, Fo3 Xv. Xi iphiopsylla Jordan & Rothschild, 1913 332 XV. 5p hyparetes Jordan & Rothschild, 1913 332 ° The °, if applicable, precedes the name in this List for convenience in scanning, whereas in the text it follows the name. VIII. Allactaga Cuvier, 1836 Aplodontia Richardson, 1829 Apodemus Kaup, 1829 Arctictis binturong (Raffles, 1821) Avrvicanthis Lesson, 1842 Bathyergus Mliger, 1811 Callosciurus Gray, 1867 mn erythraeus (Pallas, 1778) Calomyscus Thomas, 1905 Citellus Oken, 1816 » dauricus Brandt, 1843 Cryptomys Gray, 1864 Cynomys Rafinesque, 1817 Dremomys Heude, 1898 Byrinaceus Linnaeus, 1758 Eutamias Trouessart, 1769 Felis bengalensis Kerr, 1792 Heteromys Desmarest, 1817 Hylopetes Thomas, 1908 Hyperacrius Miller, 1896 Lariscus Thomas & Wroughton, 1867 Marymota Blumenbach, 1779 Marites Pinel, 1792 Meriones Wliger, 1811 Microtus Schrank, 1798 Mus Linnaeus, 1758 LIST OF HOSTS Neotoma Say & Ord, 1825 Neotomodon Merriam, 1898 Ochotona Link, 1795 Octomys Thomas, 1920 Onychomys Baird, 1857 Oryzomys Baird, 1857 Otomys unisulcatus Cuvier & Geoffroy, 1829 Peromyscus Gloger, 1841 Petauwrus Shaw & Nodder, 1791 Petrochelidon Cabanis, 1850 Pogonomys Milne-Edwards, 1877 Ptychorhamphus M. Brandt, 1837 Puffinus Brisson, 1760 Rattus Fischer, 1803 Rattus (Rattus) Fischer, 1803 Reithrodontomys mexicanus Saussure, 1860 Rhinosciurvus Gray, 1843 Rhombomys Wagner, 1841 Romerolagus Merriam, 1896 Sciurus Linnaeus, 1758 » gvanatensis chiviquensis Bangs, 1902 Spalax Guldenstaedt, 1770 Tachyglossus Uliger, 1811 Tamias, Illiger, 1811 Tamiasciuvus Trouessart, 1880 Tupaia Raffles, 1821 Myospalax Laxmann, 1769 ’ IX. REFERENCES 1967. Redefinicién de Cediopsylla Jordan y Hoplopsyllus Baker. Nuevas especies, comentarios sobre el concepto de relicto y un caso de evolucién convergente. Rev. Soc. méx. Hist. nat. 27 : 67-88, illus., refs. Bates, J. K. 1962. Field studies on the behaviour of bird fleas. I. Behaviour of the adults of three species of bird fleas in the field. Parasitology 52 : 113-132, illus., refs. BARRERA, A. 378 R. TRAUB Darskaya, N. F. 1953. On the fauna and the ecology of fleas of the Insectivora in the central strip of the European part of RSFSR. Questions of marginal, general, experimental para- sitology and medical zoology. Vop. kvaev. obshch. éksp. Parazit. med. Zool. 8 : 164-174. 1954a. The fleas of the Daursk suslik (Citellus dauricus Brandt). Fleas at hole entrance. Report 2. Izv. Ivkutsk. gos. nauchno-issled. protivoch. Inst. Sibivi dal’n. Vost. 12 : 245-250. 1954b. A note on the ecology of bird fleas of the genus Cevatophyllus. Tez. Dokl. 3 ekol. Konf., Kiev (4) : 100-103. 1955. Oncounting fleas at the entrances to burrows of Rhombomys opimus in northern and western Kyzul-Kum. Vop. kraev. obshch. éksp. Parazit. med. Zool. 9 : 87-95, refs. 1957. Fleas of the Dauria marmot (Ochotona daurica Pall.). Mater. Pozn. Fauny Flory SSSR, n.s. Otdel. Zool. 37 (52) : 163-170, refs. 1959. A preliminary study of the ecology of bird fleas of the genus Cevatophyllus Curtis 1832. Vop. Ekol. (Questions on Ecology) 3 : 35-46, refs. 1964a. On the comparative ecology of bird fleas of the genus Cevatophyllus Curt. 1832. In Ectoparasites. Fauna, Biology and Practical Significance. Vol. 4. Proc. Study Fauna and Flora of the USSR, sect. Zool, issue 39 (54) : 31-180, refs. 1964b. Fleas (Suctoria). pp. 54-67. In Petrishcheva, P, A. & N. G. Olsufjeva (eds.) Methods for the study of natural foci of human diseases. Meditsina, Moskva, refs. 1965. On the methods of study of the yearly cycle of fleas. In Mazurmovich, B. N. Parasites and parasitism in man and animals. Akad. Nauk. Ukvain. SSR, Repub. Mezh- vedom. Sbron., Ser. Probl. Parazit. pp. 363-385, refs. Darskaya, N. F., BakeeEv, N. N. & Kapatskaya, K. P. 1962. On the annual cycle of Xenopsylla conformis Wagn. in the Azerbaidjan Republic. Medskaya Parazit., Moskva 31 (3) : 342-346, refs. Darskava, N. F. & Besepina, K. P. 1961. On the possibility of fleas (Suctoria) feeding on reptiles. Tvudy nauchno-issled. protiv. Inst. Kavk. Zakavk. 5 : 33-39, refs. Fox, I., Fox, R. I. & Bayona, I. G. 1966. Fleas feed on lizards in the laboratory in Puerto Rico. J. med. Ent. 2 : 395-396, illus., refs. HAESELBARTH, E. 1966. V. Order: Siphonaptera. pp. 117-212. Jn Haeselbarth, E., Segerman, J. & Zumpt, F. The arthropod parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Vol. III. (Insecta excl. Phthivaptera). 283 pp., illus., refs. Johan- nesburg. Horranp, G. P. 1949. The Siphonaptera of Canada. Publ. Dep. Agric. Can. (817), 306 pp., illus, refs. Ottawa. 1952. Descriptions of fleas from northern Canada (Siphonaptera). Canad. Ent. 84 : 297— 308, illus, refs. 1964. Evolution, classification, and host relationships of Siphonaptera. Ann. Rev. Ent. 9 : 123-146, illus, refs. 1965. New species and subspecies of Anomiopsyllus Baker from Mexico (Siphonaptera: Hystrichopsyllidae). Canad. Ent. 97 : 1051~1058, illus, refs. 1969. Contribution towards a monograph of the fleas of New Guinea. Mem. ent. Soc. Canad. (61), 77 pp., illus, refs. Hotianp, G. P. & LosHBAuGH, G. JR. 1958. Two new species of fleas from Utah, with notes on the genus Oynithophaga Mikulin (Siphonaptera). Canad. Ent. 90 : 486-493, illus. HoocstraaLt, H. & Traup, R. 1963. The fleas (Siphonaptera) of Egypt. Host-parasite relationships of Insectivora and Chiroptera. J. Egypt. publ. Hith Ass. 38 : 111-130, refs. 1963. The fleas (Siphonaptera) of Egypt. MHost-parasite relationships of Lago- morpha (Hares). J. Egypt. publ. Hlth Ass. 38 : 243-246, refs. 1966. The fleas (Siphonaptera) of Egypt. Host-parasite relationships of Carnivora. J. Egypt. publ. Hith Ass. 41 : 75-91, refs. Hopkins, G. H. E. & Rotuscuitp, M. 1953. Az illustrated catalogue of the Rothschild collection of fleas (Siphonaptera) in the British Museum. Vol. I, 361 pp., illus, London. CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS 379 Hopkins, G. H. E. & Rotuscuitp, M. 1962. An illustrated catalogue of the Rothschild collec- tion of fleas (Siphonaptera) in the British Museum. Vol. III, 559 pp., illus. London. Hupparp,C.A. 1947. Fleas of western North America. Their relation to the public health. 533 pp., illus., refs. Ames, Iowa. Humpnriss, D. A. 1966. The function of combs in fleas. Ent. mon. Mag. 102 : 232-236, illus, refs. 1967. Function of combs in ectoparasites. Nature 215 : 3109. 1968. The host-finding behaviour of the hen flea, Cevatophyllus gallinae (Schrank) (Siphonaptera). Parasitol. 58 : 403-414, illus, refs. Iorr,I.G. 1928. Results of investigations on the flea fauna of S.E. USSR. Trudy 1. Veseoiuz. Protivoch. Soveshchan. 1927 : 204-241. 1929. Material for the study of the ectoparasite fauna of southeast USSR. VI. Fleas of mole-rats (Spalacidae). Izv. gosud. mikvobiol. Inst. Rostove (8) : 29-43, German summary pp- 56-59, illus, refs. 1941. Problems in the ecology of fleas in relation to theiy epidemiological importance. Ordzhonikidze Regional Publishing House, Piatigorsk, pp. 1-116. 1950. Thealakurt. Mater. Pozn. Fauny Flory SSSR., Ectoparasites, n.s. Zool. Sec. No. 15 (30) 2 : 4-29, illus, refs. JAtH, H. 1952. Beobachtungen zur lebensweise der Fléhe (Insecta, Siphonaptera). Z. hyg. Zool. 40 : 133-137. Jounson, P.T. 1957. A classification of the Siphonaptera of South America with descriptions of new species. Mem. ent. Soc. Wash. No. 5, 299 pp., illus, refs. Jorpan, K. 1926. On Xenopsylla and allied genera of Siphonaptera. 3 Int. ent. Kongr. Verhandl. (1925) 2 : 593-627, illus. 1937. Three new bird-fleas from Kashmir. Novit. zool. 40 : 299-306, illus. 1942. On Parapsyllus and some closely related genera of Siphonaptera. ‘‘Eos’’, Rev. Esp. Ent. 18 : 7-29, illus. 1947. On some phylogenetic problems within the order of Siphonaptera (= Suctoria). Tijdschr. Ent. 88 : 79-93, illus. 1950. On characteristics common to all known species of Suctoria and some trends of evolution in this order of insects. 8 Int. Congr. Ent., pp. 1-9, illus. Jorpan, K. & Rotuscuitp, N.C. 1915. Contribution to our knowledge of American Siphon- aptera. LEctoparasites 1 : 45-60, illus. Kurakova, Z.G. 1964. Feeding habits of the flea, Xenopsylla gerbilli caspica loff and some other fleas. Ectoparasites. Fauna, Biology and Practical Significance. Moscow State University 4 : 205-220, illus, refs. Lewis, R.E. 1964. Anewspecies of Coptopsylla Jordan and Rothschild, 1908, from Northern Saudi Arabia with comments and a key to the genus (Siphonaptera: Coptopsyllidae). Proc. biol. Soc. Wash. 77 : 199-214, illus, refs. 1969. A new Stenischia Jordan, 1932 (Siphonaptera: Hystrichopsyllidae) from the Sikkim large-clawed shrew. J. Parasit. 55 (4) : 872-876, illus, refs. Liu, C.-Y., Wu, H.-Y. & Wana, T.-C. 1965. A new genus and species of flea, Calceopsylla aduncata gen. et sp. nov., from Tsinghai, China. Acta. zootax. sin. 2 : 57-62, illus, refs. Liu, T.-S. & Li, K.-C. 1965. The discovery of Aenigmopsylla mikulini Schwartz in China and the establishment of Brevictenidia gen. nov. Acta zool. sin. 17 : 414-420, illus. DE MEILLon, B. 1952. The fleas of sea birds in the Southern Ocean. Aust. nat. Antarctic Res. Exp., Rep. Ser. B, 1 Zool., pp. 1-11, illus. DE MEILLoN, B., Davis, H.S. & Harpy, F. 1961. Plague in southern Africa. Vol. I. The Siphonaptera (excluding Ischnopsyllidae). 280 pp., illus, refs. Pretoria. MENDEZ, E. 1968. Scolopsyllus columbianus, new genus and species of the family Rhopalo- psyllidae (Siphonaptera) from Columbia. J. med. Ent. 5 (3) : 405-410, illus, refs. 380 R. TRAUB ' Murray, M. D., Orton, M. N. & Cameron, A. S. 1967. The antarctic flea Glaciopsyllus antarcticus Smit and Dunnet. Jn Gressitt, J. L. (ed.) “Entomology of Antarctica.” Antarctica Res. Ser. 10 : 393-395, illus, refs. NEvILLE, C. & Roruscuitp, M. 1967. Fleas—insects which fly with their legs. Proc. R. ent. Soc. Lond. (C) 32 (3) : 9-10. Rosicky, B. & Smit, F. G. A. M. 1965. Two new species of Ornithophaga (Siphonaptera: Leptopsyllidae). Cslkd. Pavasit. 12 : 245-253, illus, refs. RotuscuiLp, M. 1965a. Fleas. Sci. Amer. 213 : 44-53, illus, refs. 1965b. The rabbit fleaand hormones. Endeavour 24 : 162-168, illus, refs. 1969. Notes on fleas. Proc. Brit. ent. Nat. Hist. Soc. 1(1) : 1-8, illus, refs. Rortuscuitp, M. & Cray, T. 1952. Fleas, flukes and cuckoos, a study of bird parasites. 304 pp., illus, refs. London. Rotuscuitp, M. & Hinton, H. E. 1968. Holding organs on the antennae of male fleas. Proc. R. ent. Soc. Lond. (A) 43 (7-9) : 105-107, illus, refs. RotuscuHiLp, N. C. 1911. On a new genus and species of Siphonaptera from Nyasaland. Bull. ent. Res. 2 : 269-272, illus. 1915. On Neopsylla and some allied genera of Siphonaptera. Ectoparasites 1 : 30-44, illus. 1917. Convergent development among certain ectoparasites. (Presidential Address.) Proc. ent. Soc. Lond. 1916 : 141-156. SakacuTl, K. 1962. a = 2 V5 PSS 4 73 (1.2) CONORHINOPSYLLA STANFORDI EASTERN U.S.A.- SQUIRRELS Bo . —— = 74 (L7) DELOTELIS TELEGONI (ROTHSCHILD, 1905) WESTERN NO. AMERICA — PEROMYSCUS fe Pop 7 175 (1.2)MEGARTHROGLOSSUS PROCUS J. GR., |915 76 (1.7) TAMIOPHILA GRANDIS (ROTHSCHILD,1902) WESTERN NO. AMERICA — SQUIRRELS EASTERN NO. AMERICA - CHIPMUNK ll 77 (1.7) EPITEDIA FACETA (ROTHSCHILD, 1915) _ 78 (1.7) PHALACROPSYLLA PARADISEA ROTHS., 1915 ___EASTERN. U.S/A.- SQUIRRELS a SOiWESTERN U.S. - NEOTOMA PLATE 13) Pronotal Combs—Fleas of Underground Nests Fic. 79. (i.7) Strepsylla mina Traub, 1950 Mexico—Neotomodon Fic. 80. (iv.2) Phaenopsylla kopetdag loff, 1946 S.W. Asia—Calomyscus Fic. 81. (i.7) Catallagia striata Scalon, 1950 E. Palaearctic—Voles Fic. 82. (iv.2) Pavadoxopsyllus microphthalmus loff, 1946 S.W. Asia—Calomyscus Fic. 83. (i.2) Wagnerina schelkovnikovi loft & Argyropulo, 1934 S.W. Asia—Meriones Fie. 84. (iii.1) Callopsylla dolabris (J. & R., 1911) S.E. Palaearctic—Marmota Bull. Br. Mus. nat. Hist. (Zool.) 23, 10 PLATE 19 PRONOTAL COMBS—FLEAS OF UNDERGROUND NESTS (1.7) STREPSYLLA MINA TRAUB, 1950 SER 80 (IV 2) PHAENOPSYLLA KOPETDAG IOFF, 1946 MEXICO—NEOTOMODON SOUTH WESTERN ASIA—CALOMYSCUS 7) CATALLAGIA STRIATA SCALON, 1950. 82 (IZ. 2) PARADOXOPSYLLUS MICROPHTHALMUS EAST PALAEARCTIC— VOLES IOFF,1946 SOUTH WESTERN ASIA—CALOMYSCUS 84 (IIT.|) CALLOPSYLLA DOLABRIS (J. &R,, 1911) | SOUTH EASTERN PALAEARCTIC-MARMOTA Balle SCHELKOVNIKOVI .2) Wi tonne ARGYROPULO, 1934 SOUTH WESTERN ASIA— MERIONES PLATE 14 Head and Thorax of Some Nest-Fleas Fic. 85. (1.8) Wenzella obscura Traub, 1953 Central America—Helervomys Fic. 86. (1.2) Conorhinopsylla nidicola Jellison, 1945 Nearctic—Neotoma % Bull. By. Mus. nat. Hist. (Zool.) 23, 10 PLATE 85 (I 8) WENZELLA OBSCURA TRAUB, ‘1953 CENTRAL AMERICA—HETEROMYS |86 (12) CONORHINOPSYLLA NIDICOLA JELLISON, 1945 NEARCTIC — NEOTOMA z “HEAD AND THORAX OF SOME NEST- PLEAS PLATE 355 Some Examples of the Thorax of Fleas Fic. 87. (iii.t) Opisodasys hollandi Traub, 1947 Mexico—Tree-Squirrels Fic. 88. (iii.1) Opisodasys vesperalis (Jordan, 1929) Nearctic—Flying-Squirrel Fic. 89. (v.1) Nycteridopsylla eusarca (Dampf, 1908) Palaearctic—Bats Bull. By. Mus. nat. Hist. (Zool.) 23, 10 ALI NAMID,, G0 ~ ~ es 87 (I.1) OPISODASYS HOLLANDI TRAUB, 1947 MEXICO—TREE-SQUIRREL 88 (ZI. 1) OPISODASYS VESPERALIS (JORDAN, 1929) NEARCTIC—FLYING- SQUIRREL 89 (WZ.1) NYCTERIDOPSYLLA EUSARCA (DAMPF, 1908) PALAEARCTIC— BATS “ae a THe ma Ym ~ - | aN SOME EXAMPLES OF HORAX OF FLEAS PLATE x6 Head and Thorax of Two Ceratophyllus From Cliff-Swallows Fic. 90. (111.1) Cevatophyllus arcuegens Holland, 1952 Palaearctic Fic. 91. (iii.1) Cevatophyllus styx Rothschild, 1900 Palaearctic Bull. By. Mus. nat. Hist. (Zool.) 23, 10 PLATE 16 | ; | i | mo 4 - ied Nie f. is, : 90(II.1) CERATOPHYLLUS ARCUEGENS HOLLAND, 1952 NEARCTIC | | 4 INS . _ ; SL.Shipt “9| (II. 1) CERATOPHYLLUS STYX ROTHSCHILD, 1900 PALAEARCTIC ’ s a : TPN EP aX Sa — HEAD AND THORAX OF TWO CERATOPHYLLUS FROM CLIFF —SWALLOWS PLATE 17 Head and Thorax of Two Xenopsylla from New Guinea Fic. 92. (ix.1) Xenopsylla papuensis (Jordan, 1933) Pogonomys Fic. 93. (ix.1) Xenopsylla vevabilis Jordan, 1925 Rattus (Rattus) Bull. Br. Mus. nat. Hist. (Zool.) 23, 10 PLATE 17 Ce 2? ZAIN a al 92 (IX.!1) XENOPSYLLA PAPUENSIS (JORDAN, 1933) ae POGONOMYS ‘ —_——— ‘ | oe > - : é ge ee: Wig, eee % SL. Shipley ¥ 93 (IX 1) XENOPSYLLA VEXABILIS JORDAN, 1925. oi’ hy! RATTUS (RATTUS) EP ORES. 2 TORE aE ae HEAD AND THORAX OF TWO XENOPSYLLA FROM NEW GUINEA PEATE 18 Head and Thorax of Two Ophthalmopsylla Fic. 94. (iv.2) Ophthalmopsylla (Evemedosa) celata Traub, 1965 S.E. Palaearctic—A podemus, etc. Fic. 95. (iv.2) Ophthalmopsylla jettmari Jordan, 1929 E. Palaearctic—Citellus, etc. Bull. Br. Mus. nat. Hist. (Zool.) 23, 10 PLATE 18 Yo (IW. 2) OPHTHALMOPSYLLA (EREMEDOSA) CELATA | TRAUB, 1965 SOUTH EASTERN PALAEARCTIG—APODEMUS, ETC. — OPHTHALMOPSYLLA “JETTMARI JORDAN, 1929 EASTERN PALAEARCTIC— CITELLUS, ETC. Ee HEAD AND THORAX OF TWO OPHTHALMOPSYLLA Fic Fic Fic Fic . 96. OTe . 98. . 99. (iii.1 (111.1 (1.7 (ix.1 Pe AIEEE KG Contrasts in Head and Prothorax of Some Fleas Macrostylophora borneensis (Jordan, 1926) Borneo—Ground-Squirrels Macrostylophora fimbriata (J. & R., 1921) Himalayan—Flying-Squirrels Epitedia stanfordi Traub, 1944 S.W. U.S.A.—Peromyscus Actenopsyllus suavis J. & R., 1923 California—Sea-Birds Bull. By. Mus. nat. Hist. (Zool.) 23, 10 PLAGE 19 CONTRASTS IN HEAD AND PROTHORAX OF SOME FLEAS 96 (IIT. 1) MACROSTYLOPHORA BORNEENSIS (JORDAN, 1926) 97 (II. 1) MACROSTYLOPHORA FIMBRIATA (J. &R., 1921) S HIMALAYAN-FLYING—SQUIRRELS BORNEO-GROUND—SQUIRREL | : . gel 4~ (1.7) EPITEDIA STANFORDI TRAUB, 1944 , 99(TX.1) ACTENOPSYLLUS SUAVIS J.&R , 1928 q SOUTH WESTERN U.S.A.—PEROMYSCUS CALIFORNIA—SEA-BIRDS Fic. Fic. Fic. Fic. Fic. Fic. 100. 101. 102. 103. 104. 105. PEATE 20 Thorax, Pro- and Metatibia of Two Amphipsylla (iv.2) Amphipsylla marikovskvi Loft & Tiflov, 1939 Holarctic—Voles Amphipsylla marikovskit loft & Tiflov, 1939 Holarctic—Voles Amphipsylla marikouskit Lofft & Tiflov, 1939 Holarctic—Voles Amphipsylla montana Argyropulo, 1946 Palaearctic—Voles Amphipsylla montana Argyropulo, 1946 Palaearctic—Voles Amphipsylla montana Argyropulo, 1946 Palaearctic—Voles Bull. By. Mus. nat. Hist. (Zool.) 23, 10 See aN MEO HOLARCTIC - NioltS. se Me AMPHIPSYLLA MONTANA ARGYROPULO, 1946 “ PALAEARCTIC'— VOLES j Zante * ‘ > THORAX, PRO- & METATIBIA OF TWO AMPHIPSYLLA (IW.2) eee:ssscs.. ” \ - 2 MAY | a7 | Ay PRESERTED 2 / >. & A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF . THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Attson. Marine Molluscs in the Cuming Collection British Muset (Natural History) described by William Harper Pease. Pp. 96; 14 Plate 1965. (Out of Print.) £3.75. hl WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. ei. Taytor, J. D., KENNEDY, W. J. & Hart, A. The Shell Structure and Minera! of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 | 77 Text-figures. 1969. £4.50. . Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R. Vv j 1962-1964. (In press.) ei id jut Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 1967 13. NOTES ON ZOOGEOGRAPHY, VERGENT EVOLUTION AND TAXONOMY _ ELSEWHERE IN SOUTH-EAST ASIA Fah Ill ZOOGEOGRAPHY R. TRAUB Frc > DAT. BIST. i > MAY 1973 fp PRESENT. ED & Kis vy ey ib BULLETIN OF RITISH MUSEUM (NATURAL HISTORY) | Vol. 23 No. 11 LONDON : 1972 THE GUNONG BENOM EXPEDITION 1967 : 13. NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA Ill ZOOGEOGRAPHY BY ROBERT TRAUB Pp. 389-450 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 23 No. 11 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), ‘stituted in 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 yeady. 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. 23, No. 11 of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.) © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 29 December, 1972 Price £2.40 THE GUNONG BENOM EXPEDITION 1967 13. NOTES ON ZOOGEOGRAPHY, CONVERGENT EVOLUTION AND TAXONOMY OF FLEAS (SIPHONAPTERA), BASED ON COLLECTIONS FROM GUNONG BENOM AND ELSEWHERE IN SOUTH-EAST ASIA Il ZOOGEOGRAPHY By ROBERT TRAUB CONTENTS Page SYNOPSIS 392 I. INTRODUCTION 392 II. Data AND OBSERVATIONS : i 395 A. DISTRIBUTION OF RELEVANT MAMMALS IN PERTINENT AREAS 395 . Introduction 395 2. Squirrels and Murids Occuring in Malaya patil Borneo 396 3. Tree-Shrews (Gapaiidae) 3909 4. Additional Data on Mammals 399 B. Data ON DISTRIBUTION OR HOSTS OF SIPHONAPTERA 401 1. Pygiopsyllids in General 401 2. Callosciurine Fleas 403 3. Fleas of Tupaiids, Rats and Insectivores 406 4. Leptopsylla and Allies 408 5. Neopsylla and Allies . 408 6. Ceratophyllid Rat-Fleas 409 7. Insectivore-Fleas 409 8. Squirrel-Fleas in General 410 C. NOTES ON THE FAUNA OF SUMATRA AND JAVA A: AS COMPARED WITH MALaya AND BORNEO 415 III. Discussion : 419 A. INTRODUCTION 419 B. CALLOSCIURINES AS THE TRUE HOSTS OF Weatmayella’ 419 C. ORIGIN AND AFFINITIES OF Medwayella . 420 D. GEOGRAPHIC ORIGINS OF CALLOSCIURINE SQUIRRELS AND THEIR FLEAS 423 . The Callosciurine Bless 423 2. Origins of Callosciurines . 425 E. GEOGRAPHIC ORIGINS OF Rattus-FLEAS AND THEIR HOSTS . 430 1. Rattus-Fleas 430 2. Geographic Origins Of the. Australo- Asian Murids and Rattus : n : : 434 392 ROBERT TRAUB FE’. THE ORIGINS OF INSECTIVORE-FLEAS AND INSECTIVORES OF THE INSULAR MALAYAN AREA. , : ‘ 5 436 . Shrews and Shrew-Fleas_. : : ‘ a 436 Erinaceids and Their Fleas a 437 eHiese AND THEIR HOSTS IN THE INSULAR MALAYAN AREA. 438 1. Zoogeographic Inter-Relationships of Malaya, Sarawak, and Sabah + i 438 2. The Relative Poverty of the Javanese icaneachete Fauna 439 3. Zoogeographic ieee Relarossiipe of “Eanes aad the Philippines 6 440 4. Some Unusual Features alee the Sie tiwigon of Callosciurine Fleas < . : 5 : 441 ITV. CONCLUSIONS .« : ; : : 3 : : : 6 442 V. SOURCES 5 : 0 f . : : 5 443 VI. ACKNOWLEDGEMENTS é : : . c : ¢ é 444 VII. REFERENCES . - ¢ 2 5 : : 445 VIII. List oF SIPHONAPTERA CITED IN TEXT é 5 “ : ¢ 447 IX. LisT OF MAMMALS CITED IN TEXT : ; c . . 5 448 X. List oF TABLES : . . : c 0 : c 0 449 SYNOPSIS The Siphonapteran genus Medwayella, like other pygiopsyllids, is believed to have descended from fleas originating in the Australian Region, but it is a comparatively youthful taxon, with its centre of development in Borneo. Throughout its range, which is primarily in the Malayan Subregion, it is especially associated with callosciurine squirrels in the foothills and lower moun- tains. Other Oriental squirrel-fleas are of definite Palaearctic affinity, as are those of the insecti- vores and as is true for certain local murid-fleas. The data on Siphonaptera suggest that the squirrels and insectivores occurring in what is here termed the Insular Malayan Area, had their origin on the Asian Mainland. The route of entry to Borneo, however, seems to have been from Malaya through Sumatra or Java, rather than across the China Sea from Indo-China to northern Borneo. It is also clear that some members of the genus Rattus, and especially the subgenus Rattus, through the eons, must have moved back and forth between the Malayan Peninsula and Indonesia and Borneo, etc., since some of their fleas are of definite Palaearctic ancestry, and indeed one such genus occurs as far east as New Guinea. These rats, along with their fleas of Asian descent, are presumed to be recent entries, geologically speaking. How--) ever, the fleas of the more ancient rats in New Guinea and the so-called ‘Malfilindo Archipela- | goes’ are all pygiopsyllids and are believed to have had their roots in the Australian Region, and the forebears of these murids themselves may have originated there as well. Malaya, Sumatra, Java and Borneo share many faunal features, although the Javanese | fauna is relatively impoverished, while the mammals and fleas of Sarawak resemble those of | Malaya more closely than do those of Sabah. Palawan seems to have obtained its squirrels | and tree-shrews and some rats and related fleas from northeast Borneo, while much of the Mindanao fauna apparently entered from southeastern Sabah. Mindanao, however, also has” faunal elements from Celebes and the New Guinean Subregion. The data also provide support for the theory of Continental Drift in suggesting that there were ancient faunal connections, involving ‘island-hopping’ mammals and their fleas, between the Southern Continents. I. INTRODUCTION ae Stupy of the distribution and hosts of the genera of Siphonaptera treated in the first article in this series not only contributes to our understanding of the zoogeog- raphy of fleas and of their hosts, particularly as pertains to Southeast Asia and the ZOOGEOGRAPHY OF FLEAS 393 islands of the Oriental Region, but also appears to be of significance in subjects as far afield as the theory of Continental Drift (Traub, in prep.) and medical geography (Traub et al., in prep.). Accordingly, an analysis of the geographic and host relations is herewith presented in some detail, along lines listed in the Table of Contents, and the data suggest that the forebears of Medwayella Traub, 1972, and other pygiopsyllids originated in the Australian Region or Wallacea, not the Asian mainland, but that other Oriental squirrel-fleas (and the squirrels) and the non- pygiopsyllid fleas of the subgenus Rattus Fischer, 1803, were derived from Palae- arctic stocks. The data also indicate that Java has had a different faunal history than has Sumatra, and that the rodents and fleas of Sarawak have significantly more in common with Malaya than do those of North Borneo (Sabah). Evidence suggesting that there were tenuous, aboriginal faunal connections between the southern continents, as by ‘island-hopping’ mammals, is mentioned only briefly and occasionally, since the subject of Continental Drift will be treated in detail elsewhere (Traub, in prep.). In order to facilitate the discussion, much of the basic data and observations on pertinent Siphonaptera and their hosts are presented first, in a special section which in- cludes a review of squirrel-fleas from various parts of the world. Generalizations and deductions based upon such material are presented in the subsequent section on Dis- cussion, where highlights of the data are reviewed, and where new information (citing sources) is given as required. The section on Conclusions summarizes the results. In general, the Zoogeographic Regions and Subregions cited here are those used by Bartholomew, Clarke & Grimshaw (1911), but with certain new qualifications regarding Subregions, based upon unpublished notes (Traub & Hoogstraal, in litt.). Thus, it has been recognized that a map-reading alone is insufficient for denoting the limits of a Subregion, since the ecology of a particular area, depending upon the altitude or rainfall, may be an over-riding factor. Hopkins & Rothschild (1966) therefore refer to ‘the occurrence of “islands” which belong faunistically to one region in a ‘‘sea’’ of another’ and cite, as an example, the fact that the mountains of Guatemala and southern Mexico evidently have a Nearctic flea-fauna though that of the lowlands at their foot is Neotropical. In the present article, this concept has been extended. Thus, the mesic elements of the Himalayas and adjacent mountains, and the mountains of Formosa (Taiwan) are deemed as being in the ‘Eastern Palaearctic Subregion’. Instead of being placed in the Oriental Region as heretofore, the xeric parts of western Pakistan and India, etc., are here considered as being in the ‘Southeastern Palaearctic Subregion’ (along with the arid side of the Himalayas), since their faunal affinities are with the deserts of Soviet Central Asia and Iran. The Oriental Region is herein divided into the following Subregions: (x) The Indian Subregion, as used in this paper, is limited to the plains and monsoon areas of the subcontinent, western Burma, Ceylon, the terai of Nepal, etc. (2) What is here termed the ‘Burmo-Chinese Subregion’ encompasses not only the continent from Assam east to Indo-China, but south to northern Malaya, and including the foothills and lowlands of Formosa. (3) The ‘Malayan Subregion’ as used here in- cludes Malaya south of Perlis, Borneo, Sumatra, Java, Bali and Palawan (Philip- pines). The other islands of the Philippines are a zoogeographic problem and have 394 ROBERT TRAUB been assigned to various Subregions (Darlington, 1963) or ignored. Tentatively, I am placing Luzon in the Southeastern Oriental Subregion, and the remaining islands in the Malayan Subregion. Unpublished data suggest that Mindanao has some faunal similarities with Celebes, which is here placed in the “Wallacean Sub- region’ (Wallacean Region) along with Timor and the Lesser Sundas. It is impos- sible for me to evaluate such observations without adequate information about the Siphonaptera and mammals of the lowlands, foothills and mountains of Mindanao and of Celebes. The Moluccas, Ceram, etc., are placed in the Moluccan Subregion, but we have no data on Siphonaptera from those islands. In accordance with this concept of using ecological factors and faunal affinities in denoting zoogeographical areas, the Nearctic Region is regarded as extending along the upper regions of the mountain ranges into northern South America. These montane areas commencing south of the deserts of northern Mexico are termed the ‘Temperate Middle American Subregion’, while the Neotropical Region is viewed as extending north along the coasts of Mexico. From the geographic or cartographic point of view this system has drawbacks. Thus, it is awkward to have the Oriental Region rent by the vast mountain-ranges of the Himalayan Chain, so that, for example, much of Burma and south Yunnan are both in the Oriental Region, but between them are massive Palaearctic ‘islands’ of the Himalayas. However, faunistically and geologically, this makes sense. The Himalayas are actually a young group of mountains, and it appears that the tremendous ecological changes wrought by the relatively recent upthrusts (e.g. creation of deserts in the rain-shadows) have obliterated or drastically modified florae and faunae that formerly were distributed continuously across much of southern Asia. As a result, despite the extensive patches of desert, there exist ‘ecological islands’ scattered amongst the mountains throughout the region, where these same groups of mammals, ectoparasites, associated microbial infections and plant-habitats may be found as relics of past eons (Traub & Evans, 1967; Traub & Wisseman, 1968). Allusion was made, in the first article in this series, to the difficulties posed by the multiple meanings ascribed to geographical names, especially when the terms acquire political connotations which vary with the passage of time. Good (1964), for example, referred to the confusion associated with the name Malaya and its allied forms, and therefore defined and limited such usage in his opus (p. 20) as follows: (t) Malay Peninsula—‘that part of continental Asia south of the isthmus of Kra’; (2) Malayan Archipelago—'the islands thence as far as and including New Guinea’ and (3) Malaysia—both together’. This is a useful terminology, but the current use of the name Malaysia to denote the countries of Malaya and the former British colonies in Borneo has added a new source of confusion. Also, in zoogeog- raphy, apparently in contrast to plant-geography, it is desirable to deal with New Guinea and adjacent islands as an entity rather than something submerged in a ‘Malayan Archipelago’. A different terminology is required, and valid objections can be, and have been, raised to all proposals. Nevertheless, for purposes of | convenience, utility and uniformity, it seems desirable to select and specify names | for the main geographic areas used in this paper. Accordingly, in addition to the ZOOGEOGRAPHY OF FLEAS 395 Regions and Subregions as defined above, and the standard names of countries and islands, etc., the following are employed: (1) ‘Malayan Peninsula’ for the continental area south of the Isthmus of Kra. (2) The acronym ‘Malfilindo Archipelagoes’ for the entire groups of islands of Indonesia, Philippines and Borneo but excluding the islands of the New Guinean Subregion. (3) ‘Southeast Asian Mainland’ for the entire continental land-mass encompassed in the Oriental Region with the exclusion of the Indian Subregion. The Palaearctic portions of the Himalayas are therefore included. (4) The ‘Oriental Mainland of Southeast Asia’ refers to that same area in the zoogeographic sense, i.e. limited purely to the Oriental aspects and therefore excluding the Himalayas. (5) ‘Insular Malayan Area’ for the islands of the Malayan Subregion, i.e. Borneo, Java, Sumatra and the Philippines (excluding Luzon). (6) Australo-Asian Archipelagoes for the entire group of islands, from Sumatra and the Philippines to, and including, New Guinea and Australia. Two other points are mentioned for clarification and emphasis. Throughout, the term ‘Oriental’ has a zoogeographic connotation, meaning the Oriental Region, and it is not used in the geographic, political, ethnological or popular sense denoting China and Japan. Secondly, unless otherwise indicated, all comments about numbers of taxa of Siphonaptera refer to the fleas of non-volant hosts only, and not necessarily to all the known genera or species. This approach is desirable in zoo- geography because bats and birds may fly enormous distances, and the fleas found on them in one area may really be more representative of another, having accom- panied the host on its journey. The sources for the scheme of taxonomy and nomenclature of mammals are presented near the end of the article (p. 443), but scientific references are cited when unusual or controversial names are used. The data presented below, in essence, include all the known fleas from Malaya, Sarawak, Sabah, Thailand, Indo-China, the Philippines, etc., with the exception of commensal species, bat-fleas and bird-fleas. II. DATA AND OBSERVATIONS A. Distribution of Relevant Mammals in Pertinent Areas 1. Introduction In discussing the zoogeography of fleas, it is necessary to consider that of the hosts, and accordingly data are first presented on the distribution of the major hosts for which we have records on mammal-fleas, in the Burmo-Chinese and Malayan Sub- regions of the Oriental Region. Data on Siphonaptera will be presented thereafter. It should be borne in mind that no effort is made to treat all the rats and squirrels that occur within the geographical areas cited, since in some important instances data on Siphonaptera are lacking. Even with these limitations, however, certain generalizations appear warranted and zoogeographic trends can be noted which appear valid, as will be shown later. Members of the sciurid tribe Callosciurini of the subfamily Sciurinae are particu- larly well represented in the Malayan and Burmo-Chinese Subregions, and since these tree-squirrels and ‘ground-squirrels’ have a relatively rich Siphonapteran 396 ROBERT TRAUB fauna, and apparently constitute the main hosts of the genus Medwayella, they are treated in some detail. The Petauristinae, or flying-squirrels, are also wealthy in numbers of genera and species in this Subregion, certainly far more so than in the Palaearctic and Nearctic Regions. However, since relatively little is yet known about the fleas of Oriental flying-squirrels, these mammals are not included in the charts listing geographic areas, and are mentioned only briefly throughout. 2. Squirrels and Murids Occurring in Malaya and Borneo Malaya and Borneo have a rich fauna of murids and squirrels, especially callo- sciurines, both as genera and as species, as indicated in Table 1. It is noteworthy that a significant number of the genera, and even the species, are found in both areas. Thus, Malaya and Borneo share 6 of 12 genera of squirrels and 3 of 7 murids, while 9 out of 25 species of squirrels, and 12 of 30 non-commensal species of murids are found in both areas. Further, some of the species treated as distinctive are actually closely allied, so that, for example, the Malayan Rattus (Stenomys) bowerst (Anderson, 1879) may be regarded as a sibling species of the Bornean R. (S.) infra- luteus (Thomas, 1888) (just as their respective fleas are siblings), and the Malayan R. (Lenothrix) inas Bonhote, 1906, deemed the twin species of Bornean R. (L.) alticola (Thomas, 1888). However, it is also pointed out that Borneo exhibits an important degree of endemicity, especially with respect to sciurids. Thus, Rhezthrosciurus Gray, 1867, and Glyphotes Thomas, 1898, are found nowhere else, while Exiliscrurus Moore, 1958, is known elsewhere only from the Philippines, and Nannosciurus Trouessart, 1880, from Java and Sumatra. Haeronvys Thomas, 1911, occurs only in Celebes and Borneo. The Malayan taxa not listed for Borneo are generally northern forms reaching the limit of their ranges in Malaya (e.g. Hapalomys Blyth, 1859, Bandicota Gray, 1873, and Menetes Thomas, 1908), or else are restricted to Malaya, Sumatra and Java, such as Pithecheir Cuvier, 1838. Of particular relevance is the distribution of non-commensal murids and squirrels within Borneo, especially insofar as concerns Malayan species, and this is treated in Table 2. (Since little is known about the fleas of Indonesian Borneo or Brunei, _ those regions are not considered herein.) Of the mammals listed, ro of the murids and 7 of the squirrels are found in all 3 areas. Six of the rats and 9 of the squirrels are absent from Malaya. Three of the rodents are limited to Malaya and Sarawak, viz., Rattus (Lenothrix) canus (Miller, 1903), Lariscus (L.) insignis (F. Cuvier, 1821) and Sundasciurus (S.) tenuis (Horsfield, 1824). Of the Bornean rats, 3 are found only in North Borneo (Sabah), and there is 1 species of squirrel which is known only from Sarawak. Glyphotes (G.) simus Thomas, 1898 and Lariscus (Paralariscus) hosei (Thomas, 1892) are reported only from northern Sarawak and from Sabah, while Rheithrosciurus macrotus (Gray, 1856) apparently does not occur farther north than the south of Sabah. It therefore appears that there are rather significant faunal differences between Sabah and Sarawak, even though a large proportion of these rodents occur in both areas, and even Malaya as well. It is noteworthy that none of the rodents common to both Sabah and Malaya are absent from Sarawak, suggesting that the path of migration was via Sarawak. ZOOGEOGRAPHY OF FLEAS 397 TABLE I Species of squirrels (excluding flying-squirrels) and murids occuring in Malaya and Borneo Murids Malaya Borneo Squirrels Malaya Borneo _Bandicota bengalensis = Callosciurus albescens + Bandicota indica ae Callosciurus caniceps + Chiropodomys glivoides os 4 Callosciurus erythvaeus oS Chiropodomys major + Callosciurus nigrovittatus + + Chivopodomys muroides + Callosciurus notatus + + Haeromys margarettae 4 Callosciurus prevosti + + _Haeromys pusilius a Dremomys everetti + Hapalomys longicaudatus + Dremomys vufigenis + Mus musculus Je -|- Exilisciurus exilis + Pithecheiy melanurus + Exilisciurus whiteheadi 4 Rattus (Leno.) alticola + Glyphotes (G.) simus - ‘atius (Leno.) baeodon ao Glyphotes (Hess.) canaluus a ‘Rattus (Leno.) canus — + Lariscus (L.) insignis + + ttus (Leno.) inas + Lariscus (Para.) hosei = Rattus (Leno.) ochraceiventer a Menetes berdmorei o Rattus (Leno.) vajah = + Nannosciurus melanotis Rattus (Leno.) surifer + + Ratufa affinis +E + -) whiteheadi + + Rheithrosciurus macrotis + tattus (Leop.) edwardsi + Rhinosciurus laticaudatus + + attus (Leop.) sabanus ote a6 Sundasciurus (Alte.) hippurus + + Rattus (Maxomys) Sundasciurus (S.) brookei + cremoriventer + + Sundasciurus (5.) jentinki + ttus (Maxomys) fuluescens + + Sundasciurus (S.) lowi + a tattus (Maxomys) niviventer + Sundasciurus (S.) tenuis + + ttus (Rattus) annandalei + Tamiops macclellandi + Rattus (Rattus) argentiventer + oe Rattus (Rattus) baluensis oh tus (Rattus) exulans aia + ttus (Rattus) norvegicus + + Rattus (Rattus) vattus + + Rattus (Rattus) tiomanicus + + Rattus (Stenomys) bowersi + ‘attus (Stenomys) infraluteus + tus (Stenomys) muelleri + = Summary—Numbers of taxa Squirrels Murids Squirrels Murids IT. Species 12 7 Total 25 33 Number in common 6 3 Number commensal fo) 3 Absent from Malaya 4 T No. of others in common 9 I2 Absent from Borneo 2 3 Absent from Malaya II 9 Absent from Borneo 5 9 + + sinuat (S$) Snaniasppuns a + + uo, (*S) snanvasppuns a + 1yuyual (*S) snaniasppuns ar + + tdayjanw (skumouass) snjway f + 1ayooaq (S$) SNANVSDPUNS + snamywafur (skwmouass) snzyway + + + snandgqry ('a1, 7) snaniasppunsg { f = snoupumoy (snyny) snqqwar 4 + + SIYMPNYIYDY) SNANIISOUIY Y + sisuanjng (snyoy) snywy (yynos) ++ SYOAIDUL SNANIISOAYIVAY + + + Aaquanyuasan (sngyoy) snyyway + + + ae stun vfnqgory + + + suassaan{ (sAmoxn py) snyyway ia AP SYOUD]IUL SNANIISOUUD NT se Se + AajuanAowmadd (SKmoxv I) SNYIDAT B + (y3I0N) 1asoy (‘DADG) SNISIUDT = + + snunqnos (goat) snywy io ar be) + + StUsISUL (7) SHISUDT a + + ipoayazvym (‘ouaT) snyoy is + snapouna (‘ssay) sazoydapy + + + dafiAns (oUuaT) SNBWwAT ty =+- (qqI0N) snus (*5) sazoydayy = + + yoloa (ouaT) snyoy jaa) + + + AajuanraavAygo (-ouaT) SnyIwAT q =| + (so1oads z) snanisyixa = p ‘ 5 Siey/ annectans, Dicrurus 114, 142 annectans, Microhyla 40, 57-59, 95 Anomiopsyllus RDS 345, 352, 354, 355 Anoplura. - 200 Anorrhinus TT, 730: 171, 199 Ansonia 49-52, 62,95 antarcticus, Glaciopsyllu 326, 333, 334, 336, 361 Anthracoceros 154, 171, 182 Anthreptes 117, 134, 145,154 Aphaniotis . : : 76, 96 Aphropsylla . A “ é 5 - 354 Aplodontia a E 331, 343, 344 Apodemus . 272, 208, 317, Pl. 3, Pl. 8; PL 18) 408, 431 Apodidae. é 5 j = Apomys 288, 290, 298 285, 286, 291, 294, Tab. 7 (opp. p. 406), 407 appendiculatus, Polypedetes . : ar appendiculatus, Rhacophorus . : 72,95 aporus, Stivalius 452 INDEX apsidata, Palaeopsylla Tab. 7 (opp. p- 406), 416 Apus . 3 c 5 30G) Arachnothera 117, 134, eG 145, 146, 154, 171, 199 Araeopsylla . a 6 é : 5 5%} arborea, Kibatalia ; < : 6 12 Archaeopsylla 2 c eeazo: archipelagicus, Indicator’ 112, 140, 200 arctica, Mioctenopsylla . : ees 42 Arctictis 122, 149, 170, 174, 175, 177, 357 Arctogalidia . 122, 136, 150, 170, 177, 181 arcuata, Medwayella . 227-228, 232, 238, 244, 256, 267, 303, 304, Pl. 12, Pl. 32, Tab. 5 (opp. p. 402), Tab. 7 (opp. p. 406) arcuegens, Ceratophyllus 317, 326, 327, 333, 334, 344, 346, 348, 365, Pl. 16 arenicola, Leptotrombidium . 2 - 189 argalea, Gahrliepia = - 5 . 194 argalea, Schoengastiella . - “ - 194 argentaurus, Leiothrix . 6 - 25 argentiventer, Rattus argus, Argusianus . argus, Monopsyllus 153, 397, 398 108, 138, 171, 176 339, Pl. 9, 412, 429 Argusianus 108, 138, 171, 176 armiger, Hipposideros_ . = ‘ 56553 Arvicanthis . : : a : Ps Ascoschoengastia 2 193, 195, 196 asiatica, Haemaphysalis . 167, 170, 177, 183 asper, Bufo 52, 62, 69, 95 Aspis . 5 0 “ 3 1 493) Asthenodipsas “ c “ : oH Astivalius . 4 a 2 ; + 441 Atherurus . , 3 5 122, 149 atriceps, Pycnonotus 5 c - 154 atrogularis, Orthotomus . 116, 144, 154 atrogularis, Prinia . A LS E25 767; 168, 170, 171, 178-180, 182, 183, 184 Audyana 5 & : 190, 196 audyi, Agcoschoenpastia | 193, 195, 196 audyi, Cratynius Tab. 7 (opp. p. 406), 416 audyi, Odontacarus F 190, 192, 195 atrosignatus, Dermacentor auratus, Dermacentor 178, 180 “auratus-group, Dermacentor” “ - 178 aurifasciatus, Hyla - 70 aurifasciatus, Philautus . australis, Megalaima avida, Neopsylla - 46, 62, 70-71, 95 III, 132, 140 Tab. 7 (opp. p. 406) azurea, Hypothymis 117, 134, 145 azurea, Sitta 114, 142 Babesia a 9 5 A 5 ee key badia, Ducula 109g, 132, 138 badius, Phodilus 3 ‘ I10, 132, 139 Baeckia 5 4 4 a . 14 baeodon, Tenothrix 397, 398 baeodon, Rattus 6 . 397, 398 bailwardi, Calomyscus_ . : 5 + 442 bakkamoena, Otus II0, 132, 139 baliodeira, Coronella c c : ree ete baliodeirus, Liopeltis . ¢ : 86, 96 Balionycteris - I19, 136, 147, 153, 171 baluensis, Rattus 397, 398, Tab. 7 (opp. p. 406), 497, 421, 443 287-290, 292, 294, 305, Pls. 52-58, 314 bamus, Stivalius cognatus Bandicota 396, 397 baniensis, Gahnia . 5 : A 9 14 barbatus, Sus : 3 : 5 22% bardus, Apomys insignis. . ( - 288 barrerai, Dysmicus 329 bartelsi, Cratynius Tab. 7 (opp: p- 406), 416 bartelsi, Maxomys : - 418 bartelsi, Rattus. : 5 . 418 Bathyergus 431, 343, 340 batibacula, Medwayella 250-2 52, 255, 256, 265, 267, 304, Pls. 35-37, Tab. 5 (opp. p. 402), Tab. © (opp. P- 404), 416 bats. 21-42 batuanae, Neohaematopinus 5 5 + 200 Bazzania 3 5 a zi XY 5 18 Bazzaniaceae 5 : ; 0 . 18 bengalensis, Bandicota . a ~ 397; bengalensis, Felis 122, 150, ey 175, 177, 181, 357 bengalensis nebulosus, Varanus : 79, 96 berdmorei, Bullimus : A : 7 433 berdmorei, Engystoma . : 59 » 242,244, 245, 247, 298, 397, 399, Tab. 6 (opp. p. 404) berdmorei, Menetes berdmorei, Microhyla_ . 3 5 59,95 berdmorei, Rattus 407, 433 Berenicornis . > O III, 139 bernsteini, Coelops 2 e . 2g) Berylmys . 2 5 f - 407 bicolor, Hipposidexos 120, 147 bicolor, Ratufa 6 136, 148, 153, 171, 181 bicolor, Trichastoma = 115, 133, 142 bidentatiformis, Neopsylla_. : -. PL6 bimaculatus, Rhacophorus ¢ 5 ee 75 binturong, Arctictis 122, 149, 170, 174, 175, 177, 357 birds. 5 : 103-154 bisoctodentatus, Ctenoplithalmus 315, Pl. 4 bispinosa, Haemaphysalis 167, 173, 183, 184 blanfordi, Hesperoptenus 37-38, 40 Blankaartia . : ; . 190 blepotis, Miniopterus schreibersii ’ 39, 40 blumei, Weinmannia , ° : 9 13 blumii, Dicranoloma 5 3 F : 18 blythi, Rana ‘ ‘ : ; -64, 95 blythi, Rana macrodon . ‘ : ay» (64 Blythipicus : II2, 132, 140 bodense, ep fae eee 190, 191, 195, 196 bogora, Medwayella robinsoni 235~—236, 266, 269, 303, Pls. 19-20, Pl. 22 bogoricum, Clastobryophilum . 5 i 9) Boiga . : 2 Vas bolivari, Orchopeas 339, Pl. 5, 412 bolivari, Orchopeas howardi_ . “<4E2 INDEX 453 borbonica, Cacophryne borbonica, Hylaplesia . . A oeel53 borealis, Phylloscopus 116, 144 borneensis, Goniocephalus 76, 96, 171 borneensis, Lepturophis . 87- 88, 89, 96 borneensis, Lophyurus . 0 4 76 borneensis, Macrostylophora 319, 339, Pl. 19, Tab. 5 (opp. p. 402), Tab. 6 (opp. p. 404), Tab. 7 (opp. p. 406), 416 borneensis, Rhinolophus a D 24, 25 borneensis- eke Rhinolophus_ . é 25 Bos 9 9 a 6 5 rel 5S Bothrops 5 ci z 5 ; os Bovidae 3 a a0 123 bowersii, Rattus 122; hae: 146, 149; 153, 170, 171, 174, 175, 178, 182, 191-196, 216, 261, 298, 396, 397, 401, Tab. 7 (opp. p. 406), 409, 433 bowersii, Stenomys 261, 298, 396, 397, 401, Tab. 7 (opp. p. 406), 409, 433 Brachyctenonotus . 327, 333, 352 brachyotis, Cynopterus . 119, 1 He, 147, 153, 171 Brachypteryx 4 3 D 25 brachypus, Doloisia 193, 196 brachyura, Hystrix E 5 : A adshe brachyurus, Micropternus 5 : 554 Bradiopsylla ° 320, 341 bres, Criniger 113, 133, 141 brevicaudata, Napothera 115, 133, 142 Brevictenidia 327, 341, 352, 411 ?brevipalmatus, Cyrtodactylus 80-81, 96 brevipalmatus, Gymnodactylus ; - 80 brevipes, Hylaplesia ° b : 4 brevipes, Ixalus_. “ : 5 ee 7 brevipes, Pelophryne . ; 0 54,95 brodiei, Glaucidium brookei, Sundasciurus I10, 132, 139 +257, 298, 397, 398 brooksi, Sciurus. . : A 257. Brueelia © c 5 3 + 200 brunneata, Rivnomiyias 5 0 - 154 brunneicauda, Alcippe . 115, 134, 135, 143, 146 brunneus, Pycnonotus I13, 141 Bryophytes 5 17-20, 8 Pls. Bucerocolpocephalum . 5 5 ekg) Buceros 0 5 a III, 139 Bucerotidae . 5 SLs Bufo . c 5 : ae 52- 53, 62, 69, 95 Bufonidae . 6 3 é 49-54 bukit temmincki, Rattus ; - 256 Bullimus : é 407, 421 Bungarus. c t ¢ 93, 96 burneti, Coxiella 5 167, 172, 175, 178, 184 butleri, Lycodon . 3 AT, butleri, Microhyla . a 6 6 57,95 Cacomantis . : c A eet 54: Cacophryne . : 53-5495 caedens, Orchopeas o 1 Caenopsylla . 5 Ee elas: caerulea, Pitta I12, 133, 140, 154, 190, I9I, 195, 200 caesium, Acer a a 6 a 13 Calamaria 40; re 86-87, 96 Calamus 5 14 calcarata, Medwayelia 247-250, ast, 252, 255, 258, 259, 260, 261, 262, 263, 264, 265, 304, Pls. 34-37, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. p. 404), 406, 416, 420 calceatus, Ctenophthalmus a S47, calceatus, Ethioctenophthalmus 5 S47, calceatus, Syngenopsyllus =») 341,,lon, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. p. 404), 405, 416 Calceopsylla . 327, 333, 352 caldwelli, Rhinolophus macrotis = 272. caliotes, Ceratophyllus_ . ; =g20) Callicostella . : 5 a 19 Callistopsyllus 324, 333. 335, 355 Callopsylla 326, 356, Pl. 13 callosciuri, Neohaematopinus . = (200 Callosciurus . 121, 124, 136, 146, 148, 153, 179, 171, 178, 190, 192, 193, 196, 200, 216, 227,228, 229, 232, 233, 234, 235, 236, 241, 242, 245, 256, 259, 260, 261, 262, 263, 264, 280, 298, 319, 397. 398, 399, 401, 404, Tab. 6 (opp. p. 404), 405, 4 415, 429, 439, 441, 442 Calomyscus . Pl. 13, 408, 442 Calorhamphus a III, 132, 140 Calotes s fs 40 calvus, Haemaphysalis 16741 168, 173, 183 Calymperaceae é ae lG: Calymperes . . 5 5 5 Deak) Calyptomena 112, 133, 135, 140, 199 Campephagidae . 6 : z of ETE} canalvus, Glyphotes : 397, 398 canalvus, Hessonoglyphotes_ . : - 397 candidum, Leucobryum . 18 caniceps, Callosciurus. 121, 124, 146; 148, 153, 170, 171, 178, 190, 196, 227, 234, 259, 261, 263, 264, 298, 397, 399, Tab. 6 (opp. p. 404) canis, Ctenocephalides_ . : 5 ey canus, Lenothrix 261, AS 396, 397, 398, 415 canus, Rattus 171, 261, 298, Ee 397, 398, 415 capellei, Treron . 9 53 capistratum, Pellorneum_ 115, 133, 135, 142 Capitonidae . ILI-112 Capricornis 3 ‘ 123, 150 cardis, Turdus ead 5 a a a te7{9} cardis cardis, Turdus 5 , ili) carinatus, Anorthinus galeritus 6 7D caroliniensis, Sciurus r 4 6 age caspica, Motacilla . 6 Geis caspica, Xenopsylla gerbilli 335, 350, 357 castaneceps, Alcippe 115, 134, 135, 143, 146 castaniceps, Seicercus 116, 144 Catallagia 321, 32 a 333, blir, blag Caudacaecilai A A 48,95 caudolineata, Abaetulla 5 6 a7, cavernicola, Epitedia . A Perr Cediopsylla . : : , 364 celata, Eremedosa . o 328, Pl. 18 454 INDEX celata, Ophthalmopsylla . 328, Pl. 18 celebensis, Pygiopsylla_ . a é esse. celebensis-group, Stivalius 402, 407 Centetipsylla c o F 320 centropi, Haemaphysalis) 167, 177 Centropus 109, 154, I7I, 177, 182, 199 Cephaloziella : 2 : : 18 Cephaloziellaceae . . : 4 18 Ceratophryne c . : : 63 Ceratophyllidae 411, 413 Ceratophyllus 230, 317, 326, 327, 333, 334 344, 346, 348, 349, 355, 363, 364, 365, 366, 368, Pl. 16, 423 Cercopithecidae. : . “ ec ZO Cervidae ; 5 F 6 : 5 see) Cervus . 153, 173 cetrata, Galaliepias 194, 196 ceylonensis, Culicicapa 116, 134, 144 ceylonensis, met a ‘ 315, Pl. 2 Ceyx - . 2 III, 132, 139 Chaetopsylla o : ; 5 a 5359 Chaetura 4 5 : 4 : I10, 139 chalconota, Rana . 67, 68, 69 chalconota raniceps, Rana 65, 68, 95 Chalcophaps . = 109, 132, 138 Chalinolobus ; é : Fi Z 35 Chaperina : 2 54-55, 95 charlottae, Hy psipetes . 114, 133, 141, 154 charlottensis, Catallagia . 6 Seng ells aii chaseni, Rhacophorus” . . 5 : 72 chaseni, Rhinolophus Z . 7 24 cheela, Spilornis 108, 138 cheopis, Xenopsylla 337) 357 433 chewing lice . 6 a 5 199-200 Chiastopsylla : < 5 f or) chiggers 5 ; : : 187-198 Chilophylla_ . : : 7 29, 30 Chimaeropsylla_. 9 + 363, 364 Chimarrogale ; ; - 2 - 437 chinense, Lepidosperma . - : c 14 chinensis, Kitta. . E25 chiriquensis, Sciurus pravateneis : 316 Chironax 119, 126, 136, sag 17I Chiropodomys 122, 149, 171, 397, 398, 441 Chiroptella i 192, 195 Chiroptera. : z 21-42 chlorolophus, Picus 112, 140 chlorophaeus, Phaenicophaeus’ 109, 132, 139 Chloropsis . b - 113, 133, 141, 154 Choristopsylla : < 325, 334, 341 chrysarga, Natrix . 6 2 ° ied Wf chrysarga, Rhabdophis . A See ut chrysea, Stachyris . : 1s 134, 143 Chrysococcyx ° d : 109, 138, 154 Chrysocolaptes 5 3 112, 133, 140, 171 Chrysopelea . D = 2 E 47 chrysopogon, Megalaima III, 132, 139 Ciconia 6 3 53 ciliatus protinus, Monopsyllus 5 Pl. 10 cinereum, Malacopteron . I15, 135, 142 cinnamomeus, Ixobrychus ; 3 see SS cinnamomeus, Pericrocotus 113, 141 circumdatus, Pipistrellus és 2 34-37 Citellophilus . 322, 349. 356 Citellus 313, 315) 349, aside Play Pls gt as ‘ a 19 Clay, T. 5 : 3 = . 199-200 Cnemaspis_ . : : ; 80, 96 cochinchinensis, Chloropss 113, 133, 141 Coelops 28-30, 4o cognatus, Stivalius! comm 205, 28%, 285-286, 290, 293, 367, 368, Tab. 7 (opp. p. 406), 407, 416, 430 cognatus, Stivalius cognatus 285, 286-287, 294, 305, Pl. 54, Pl. 58 cognatus bamus, Stivalius 287-290, 292, 294, 305, Pls. 52-58, 314 cognatus cognatus, Stivalius 285, 286-287, 294, 305, Pl. 54, Pl. 58 cognatus spiramus, Stivalius 205, 285, 287, 294, 305, Pl. 54, Pl. oe Pl. 8 collaris, Polydontophis_ . 2 40 collaris, Sibynophis : 5 F : 40 colletti, Rhacophorus’. ; ; 72,95 Collocalia. : : : = EEO Collorhabdion 2 ; 47 coloradensis, Tarsopsylla. 5 2 389) Coluber : 5 : : 7 84 Colubridae_. ‘5 “| : ; 84-86 Columbidae . F ‘ : + To9 comata, Hemiprocne ° é F 110, 139 comatus, Berenicornis. . s III, 139 comosum, Dacrydium . E é a 14 compactus, Dermacentor 167, 168, 171, ee 179, 180, 182, 183, 184 concolor, Rattus . 5 . 287, 208 concreta, Halcyon . III, 132, 139 concretus, Hemicircus 112, 140, 154 conformis, Xenopsylla_ . . 5 - 358 Conorhinopsylla 321, 324, 329, 333, 335, 349, 345, 352, 355, 368, Pl. 12, Pl. 14, 411 consobrinus, Cyrtodactylus’. . 81, 96 consobrinus, Bilge ge be: : : = Tod Cophias : : Sl. Copsychus “116, ea 135, 143, 144 Coptopsylla . - 2 SgB2 Coracina 113, 133, 140, 154 cornigera, Haemaphysalis cornigera 167, 168, 171, 173. 183, 184 cornigera cornigera, Haemaphysalis 167, 168, 171, 173, 183, 184 coronatus, Anthracocerus - 154 coronatus, Phylloscopus . 0 116, 124, 144 Coronella 5 : : : 7) 86 Corrodopsylla . 2 . - 347, 441 Corvidae 3 4 - 3 2 Pa i | Corvus é a : : 114, 133, 142 Corydon ‘ 4 154 Corypsylla 332, "333. 336, 337. 351, 352 coucang, Nycticebus 120, 148, 171 INDEX 455 Coxiella 167, 172, 175, 178, 184 Craspedorrhynchus 5 - 199 Cratynius Tab. 7 (opp. p. #66); co 416, 417, 418, 419, 431, 437 cremoriventer, Maxomys 261, 298, 397, 398, 418 cremoriventer, Rattus 122, 136, 146, 149, 153, 171, 191, 196, 261, 298, 397, 398, 418 Criniger 113-114, 126, 133, 135, 141, 199 criniger, Hypsipetes . II4, 133, 141 crispis, Paraceras Tab. 5 (opp. p. 402) croceus, Syrrhopodon c 18 118-119, Ta 401, Tab. 7 (opp. P- 406), 436, 437, 438 cruentus, Oriolus o =) 204, 126, 542 crypticus, Cratynius Crocidura Tab. 7 (opp. p. 406), 416 Cryptoctenopsyllus 331, 346 Cryptomys : te 331, 346, 347 Cryptopsylla : E 329, 331, 332, 346 cryptotis, Alloctenus 5 6 6 aesi4 crypototis, Ctenophthalmus 314, Pl. 4 ctenacarus, Ascoschoengastia . 193, 196 Ctenidiosomus : ; < : = 352 Ctenocephalides 326, 337, 349, 350, 354, 357, 358 Ctenophthalmus 313, 314, 315, 316, 318, 322, 347, 362, Pl. 4 Ctenophyllus. , ss cuculatus, Orthotomus 116, 134, 144 Cuculicola. 5 a e 199 Cuculidae. eS : z F - 109 Cuculiphilus . f , ; 199 Cuculus : i noo! Ha 153 cuiae, Macrostylophora Tab. 5 (opp. p. 402) Culicicapa 116, 134, 144 cuniculi, Spilopsyllus 337, 364 curtisii, Shorea 2 5 3 ¢ 5 13 curvirostra, Treron 5 3 109, 132, 138 curvirostris, Phaenicophaeus 109, 132, 138, 199 Cutia . : 5 : 5 115, 134, 143 cyane, Erithacus . 5 4 116, 124, 143 cyaniventris, Pycnonotus 113, 133, 141, 154 cyanopogon, Chloropsis : LES; 1SS 40 cyanouroptera, Minla_. F 0 PD 5 cyanurus, Psittinus 2 5 5 Pae5 cyclotis, Murina 38, 39, 120, 136, 148, 153 cyclotis, Murina cyclotis : 5 5 3) cyclotis cyclotis, Murina 6 : =. _ Sis) cyclotis peninsularis, Murina 38, 39, 40 Cylindrophis a 2 P 3 47 Cymbirhynchus . TZ) 033 e135) 7403 154 Cynocephalidae_ . : 4 4 Bibs) Cynocephalus ei a . it, 136, 147 Cynomys P : 5 5 350. Cynopterus 119, 136, 147, 153, 171 Cyrtodactylus 47, 72, 80-81, 96 Dacrydium . : = . USreia Dactylopsylla 331, 346 dammara, Agathis 3 z - : 13 Dasypsyllus . 3 342, 366, 423 dasypus, Delichon . 5 113, 135, 141 dauricus, Citellus . ,. : - 349 degnani, Bucerocolpocephalum 2 - 199 deignani, Myotis. . : 31 Delichon ae 135, 141 delichoni, Ceratophyllus . deliense, Leptotrombidium 189, 190, 191, 195, 196, 197 Delotelis ; 320,324; 333; El xr Dendrocopos 0 oeelT Dendrogale 272 , 280, 309, 400, 406, Tab. 7 (opp. p. 406), 420, 439 densipiliata, Toritrombicula 190, 191, 192, 195 dentipalpis, Haemaphysalis_ . 5 a derbyanus, Hemigalus 122, 150, 170, 176, 177, 181, 193, 196 Dermacentor 167, 168, 169, 170, 171, 178-180, 181, 182, 183, 184 diadema, Hipposideros 120, 136, 147, 153, 171 Diamanus_. » 9356 diardi, Phaenicophaeus “z09, a2, 138, 154, 199 diardi mulleri, Typhlops 47, 83, 96 diardii, Harpactes . I10, 132, 139, 200 326, 346, 348, 365 diardii, Rattus E 3 ‘ 3 Os diardii, Rattus rattus. * : =, 193 Dicaeidae . 3 . ‘ GB . wiKy Dicaeum : 6 117, 145, 154 Dicranaceae . : 2 a a 6 18 Dicranoloma , : 5 5 . 18 Dicruridae . : =e ee! Dicrurus “114, 133, 142, 154, 200 Dilaenaceae . 0 17 dilatatus, Giedophthalmnus 9 a ESS, dilatatus, Neoctenophthalmus z “Sts Dillenia ¢ . c : 12 diminutum, Acroporium : é c 19 Dinopium . a ¢ : 0 54! Dinopsyllus . A 331, 346 Dipsadidae . . 86-92 dispar, Neopsylla "233, aie Pl. 6, Tab. 7 (opp. P- 406), 409, 416 disparunguis pingue, Gahrliepia 195, 196 disparunguis pingue, Walchia . 195, 196 distans, Bazzania . 5 E x P 18 Distichophyllum_ . G 19 divisus exsecatus, Megarthroglossus op hls doenitzi, Haemaphysalis 167, 171, 177, 183, 184 dohrni, Rhinolophus macrotis 27-28, 40 dolabris, Callopsylla 326, 356, Pl. 13 dolens quitanus, Pleochaetis . 5 + 414 Dolichopsyllus 0 a 0 : Ees43s Doloisia a 193, 196 Doratopsylla > 347, 351 Dorcadia : 5 5 : - 360 dorcadia, Dorcadia : : ; - 360 doriae, Hesperoptenus . 3 5 38, 40 dorsalis, Tupaia_. - 400 Draco . A : 76-79, 96, 190, 192, 195 456 INDEX Dremomys 125, 216, 226, 241, 259, 261, 264, 273, 298, 319, 397, 399. 401, Tab. 5 (opp. Pp. 402), Tab. 6 (opp. p. 404), 405, 415, 439 dryadosa, Medwayella 215-227, 228, 229, 230, 235, 237, 238, 239, 243, 244, 245, 246, 248, 249, 250, 251, 252, 253, 259, 260, 261, 262, 263, 264, 265, 268, 273, 274, 282, 288, 302-304, Pls, 2-11, Pl. 16, Pl. 31, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. p. 404), 406, Tab. 7 (opp. p- a8, 416, 420 Dryocopus . ; B - 154 Ducula 5 4 é 109; 132, 138 dumetoria, Muscicapa : 5 116, 134, 144 duvaucelii, Harpactes . ° I10, 132, 139 Dysmicus - o 9 A é - 329 ecaudatus, Megaerops” . : 119, 136, 147 echidnae, Bradiopsylla . : = Sie) Echidnophaga 337, 338, 357, 359, 362, 433 Echinosorex 119, 146, 147, 170, 172, 182, 191, 192, 195, 415, 437 Ectinorus. 5 2 : 2 329) Ectropothecium. é 4 5 19 edwardsi, Leopoldamys 307, 433 edwardsi, Rattus 122, 136, 146, 149, 171, 191, 192, 193, 194, 195, 196, 397, 433 Elaphe 5 47, 84-85, 96 Elapidae c “ r : 93-97 elata, Frontopsylla 4 5 z ee OZ elatum, Dacrydium F : = Sra elegans, Petaurista T2001, ao) 171, 235, 298 Elephantidae 5 : 123 Elephas : 5 c 123, 150, 153 ellioti, Anathana . a é 400 ellipticum, Thamnium . : - ee) Eltonella : 6 5 : 190, 195 eltoni, Eltonella . j - 190 Emballonura. 5 6 119, 147, 171 Emballonuridae . : c 119 emys, Testudo 5 4 " 6 en 182 enca, Corvus c n é II4, 133, 142 enciari, Notiopsylla - 5 : = 326) Engystoma . F 59 Enicurus oe 134, 135; 1445 154 enoplus, Opisodasys 339, PL 7 Eonycteris . 6 5 A 5 ashy oui Eopsylla * 324, 329, 333 epilepidota, Napothera I15, 126, 135, 142 Epimys . 287, 298 episcopus, Rhinolophus macrotis 5 a7 Epitedia = 324, 330, 333, 349, 347, 355, Pl. 11, Pl. 12, Pl. 19, 411, 412 Eptesicus. 4 A c 533 Eremedosa 338, 330, Pl. 18 Erinaceidae . i 5 é 2 IT9 Erinaceus’. c 3 9 sz0 Erithacus 116, 124, 143 erithacus, Ceyx III, 132 erythraeus, Callosciurus 259, 264, 298, 319, 397, 429 erythrocephalus, Garrulax IT5, 134, 135, 143, 199, 200 erythrocephalus, Harpactes 111, 126, 132, 139 erythrophthalma, Lophura_. 153, 171, 176 erythroptera, Stachyris . 115, 134, 135, 143 erythropthalmos, Pycnonotus 113, 133, 141, 154 esculenta, Collocalia 0 : é A ente} Ethioctenophthalmus) . c 3 S47, etruscus, Suncus . : : - - 118 Eugeissona . : 13 eumolpi, Monopsyllus enmolpi 5 Cor 33 eumolpi eumolpi, Monepeyales : BES: Eurostopodus ‘i . 5 - 154 eurycercus, Centropus sinensis - 7 Eurylaimidae 5 c 112 Eurylaimus . " 112, 133, 135, 140 eusarca, Nycteridopsylla 318, 345, 352, Pl. 15 eutamiadis, Monopsyllus : 2 Pl. 10 Eutamias. Pl. 10 euteles, Macrostylophora Tab. 5 (opp. p. 402) eutilotus, Pycnonotus . - Oe tGy Eutrombicula : 2 : . - 190 Evansipsylla 4 ; » 324 everetti, Dremomys 273) 397, 398 everetti, Helictis orientalis 5 433 everetti, Melogale orientalis. A rA33 everetti, Urogale . : 3 i + 400 everetti, Zosterops excelsa, Koompassia exilia, Macrostylophora 117, 145 piel ed: res aint, Tab. 5 (opp. p. 402) exilis, Exilisciurus . 397, 401 exilis, Monopsyllus 315, Pl.g Exilisciurus . 306, 397, 398, 401 expansus myodosus, Ctenophthalmus sabe! exsecatus, Megarthroglossus divisus 5 Jai exulans, Rattus 153, 287, 397 exulans todayensis, Rattos : . 288 faceta, Epitedia 333, 340, Pl. 12 Falconidae . 5 : “ - 108 fasciatus, Nosopsyilus G > - 416 fascicularis, Macaca 120, ee 153, 171 Felidae. 2 kez iRelisiy ees 150, 170, 175; seh 181, 357 felis, Ctenocephalides. 5 . 357. ferinus, Lentistivalius 272, 280, 291, 312, 314, Pl. 3, 401 ferinus, Stivalius . A : e772 ferinus-group, Stivalius . 204, 205, 272 ferruginea, Muscicapa 116, 144 ferrumequinum-gvoup, Rhinolophus ay Filicinae = a enzo filmy ferns yaaa 8 Pls. fimbriata, Coracina : 113, 140, 154 fimbriata, Macrostylophora 339, Pl. 1, Pl. 19, Tab. 5 (opp. p- 492), 403, 412, 424, 429 fimbriatus, Draco . A ; E 76, 78 INDEX fimbriatus, Draco fimbriatus 76-77, 79, 96 fimbriatus fimbriatus, Draco 76-77, 79, 96 finlaysoni, Callosciurus . ° + 245, 298 finlaysoni, Pycnonotus . a 2 - 154 finschii, Criniger_ . 4 é 113, 133, 141 flammeus, Pericrocotus . 3 113, 133, 141 flavala, Hypsipetes - 114, 126, 133, 142, 200 flavescens, Leptospermum 14, Pls. 6-7 flaviceps, Bungarus 93, 96 flavimanus, Callosciurus ae 22 148, I71 flavinucha, Picus 125 flaviscapis, Pteruthius 115, 143 flavolineata, Coluber : 84 flavolineata, Elaphe 84-85, 96 fleas . 201-450, 78 Pls. fletcheri, Gahrliepia 194, 196 floweri, Calotes O E46 forest zones . 9-15, 8 Pls. formosanus, Coelops 29 formosus, Draco i 77 formosus, Draco formosus 77, 96 formosus formosus, Draco 77, 96 fornacis, Monopsyllus Pl. 10 Foxella 346 franklinii, Megalaima 112, 132, 140 fraseri, Calamaria . 47 Freycinetia 2 13 fringillarius, Microhierax. 108, I 35, 138, 154 Fringillidae 3 eel, frithii, Coelops 28-30, 40 frontalis, Sitta 114, 142 Frontopsylla ee 362, 408, 431 frutescens, Baeckia 6 14 fuliginea, Ansonia . E50) fuliginosa, Crocidura I 18-1 19, 147 fuliginosus, Calorhamphus III, 132, 140 fulleri, Orchopeas . PL5: fulvescens, Maxomys 397, 308, 401, Tab. 7 (Opp. P. 406), 407, 409, 415, 438 fulvescens, Rattus 397, 398, 401, Tab. 7 (opp. P- 406), 407, 409, 415, 438 fulvescens temmincki, Maxomys 9 - 298 fulvescens temmincki, Rattus . 404, 412, 415, 423, 424, 426 Funambulus fusca, Aphaniotis . fusca, Chaperina fusca, Gerygone fusca, Otocryptis fuscus, Aphaniotis Gahnia Gahrliepia galericulatus, Platylophus galeritus, Anorrhinus galeritus, Hipposideros 256, 298 2 14 194-195, 196 114, 133, 142 III, 139, 199 + 120, 136, 147, 153, 171 galeritus carinatus, Anorrhinus 2 5 aeepe galgulus, Loriculus gallinacea, Echidnophaga gallinae, Ceratophyllus 109, 138 = 9357 349, 355, 365 457 gallinulae, ep Ege 366, 423 Gallus . 181 gallus, Gallus 181 gammonsi, Kohlsia 7) 306) garei, Ceratophyllus ; 246; 365, 366 Garrulax 5 “pS = 135, 143, 199, 200 gaurus, Bos . 5 4 253 Gehyra . si a 6 : 82, 96 Gekkonidae . ¢ 0 5 a 80-82 Genoneopsylla 324, 333, Pl. 11 Gentiana a 3 “eens gentryi, Leptotrombidium 5 A 191, 196 geomydae, Amblyomma 167, 168, 171, 182, 183, 184 georychi, Xenopsylla_. 7 - 347 gerbilli caspica, anos? Bee 350, 357 Gerygone . 4 - 116,144 gigantea, Macaranga = é : : 12 gimletti, Calamaria é 4 3 ae ea gimletti, Calamaria lowi 6 oud: girshami, Acropsylla : Pl. 8 Glaciopsyllus ie 333, 334, a6: 362, 362, 366 Glaucidium 110, 132, 139 Gleichenia . O 12 gliroides, Chiropodomys . 122, irae. int, 397, 398 glis, Tupaia . 120, 148, 153, 171, 181, 216, 236, 241, 243, 245, 259, 261, 262, 263, 298, 399, 400, 415, 438 Glyphotes . cee 397, 398, 405 golovi, Ctenophthalmus : = = 9362 Goniocephalus : A - 46, 76, 96 goniorrhyncha, Groutiella 5 : : 19 Gonocephalus 5 f : : Sati Gonyophis . ; 5 : -85, 96 Gonyosoma . 6 0 : oS gouldi, Chalinolobus : 35 gracile, Leptobrachium . 5 i 57. 60-62, 95 gracilis, Tupaia.. : 0 400 Gracula c : - 154 granatensis chiriquensis, Sciurus : - 316 granatina, Pitta . 5 154 grandis, Muscicapa 5 a 117, 134, 144 grandis, Tamiophila B 4 e Pl. 12 Grandison, Alice G.C. . 43-101 granulatus, Ixodes ee: 170, 172, 183, 184, 196 graphis, Kohlsia_ . é = GLO bia gratiosa, Xenopsylla Ps 5 E 700342 griffithii, Plectocomia . Qo 0 13 grodekovi, Aenigmopsylla 6 6 314, Pl. 2 grossiventris, Malacopsylla_. n - 361 Groutiella. 2 - : 19 Gryphopsylla Pl. 8, 402, Tab. 7 (opp. Pp. 406), soy ae: 417, 418, 421, 431 guajana, Pitta : 2 - 154 guba, Acanthopsylla 5 6 j - 292 guereci, Rattus. 5 5 2 ezoy, gularis, Macronus . : 115, 143 Gunong Benom Expedition 1067 1-450, 91 Pls. Gymnodactylus. 5 - 47, 80-81 Gymnophiona = 5 5 ‘ ae Oe) 458 gymnurus, Echinosorex 119, 146, 147, 170, 172, 182, 191, 192, 195, 415 haagi, Ctenophthalmus . : 6 © Elta haddowi, Chimaeropsylla 363, 364 Haemaphysalis 167, 168, 169, 170, 171, 173-178, 182, 183, 184 Haematortyx 5 5 a é e775 Haeromys “ 2 396, 397, 398 hageni, Bothrops . : 3 : BE ag3 hageni, Trimeresurus 93-94, 96 Halcyon 0 III, 132, 139, 154 hamata, Metzgeria 2 ‘ 17 hamatum, Paraceras Tab. 5 (opp. p- 402) hampei, Calymperes 2 19 hanitschi, Ansonia 5 “ 5 Hapalomys 396, 397 hardwickii, Chlorospis 113, 141 Harpactes 110, 726) 132, 139, 154, 200 hascheana, Rana . é 5 : 40 hasselti, Leptobrachium * = Or hasseltii, Leuconoe 5 : 32 hasseltii, Myotis ant 32-33, 40 hasseltii, Rafflesia . : 2 6 5 12 hastata, Macrostylophora 319, Pl. 2, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. p. 404), 405, 416, 442 hastata, Macrostylophora hastata_ . + 314 hastata hastata, Macrostylophora_ . = Ta: Hectopsylla . . 9333 heinrichi, Macrostylophora Tab. 5 (eae. p- 402) Helarctos 122, 149, 181 Helenicula i 193, 196 Helictis 5 , 5 : = 436 Hemicircus f I12, 140, 154 Hemidactylus 5 . 82 Hemigalus . 122, 150, 170, 177, ae 193, 196 Hemiprocne . I10, 139, 154 Hemiprocnidae . : = kG: Hemipus 113, 141 henricii, Megalaima III, 132, 140, 200 hermaphroditus, Paradoxurus 153, 170,177, 181 Hesperoptenus 5 37-38, 40 Hessonoglyphotes . z 2 c = 397 Heterometrus 3 190, 196 Heteromys . 7 5 = : Pl. 14 Heteronota . 3 2 : 80 Heterophasia : “rs, 134, 143, 199 heteropus, Leptobrachium 55, 61, 62,95 heymonsi, Microhyla 59-60, 95 Hieraaetus 5 5 . 108, 138 EATS feneen e 3 6 P 21-42 himalaica, Peromyscopeylla 4 - 408 Hinulia ; E A ee t's} Hipposideros 28, oe 120, "136, 147, 153, 171 hippurus, Aletesciurus . - 398 hippurus, Callosciurus Tab. 6 (opp. P- 404), 415 hippurus, Sundasciurus . 121, 148, 153, 171, 227, 259, 298, 397, 398 NDEX hippurus-group, Callosciurus . . - 40% hippurus perse, Callosciurus . » QOL hirsuta, Chilophylla 9 A 29, 30 hirundinaceus, Hemipus F 113, 141 Hirundinidae 5 bv : ~~ GDS hirundinis, Ceratophyllus : - 346, 365 Hirundo 5 5 d Q = ITZ hissarica, Neopsylla 2 2 : 7 ELXG hodgsoni, Muscicapa hollandi, Opisodasys 315, 317, 327, 339, 344, Pl. 7, Pl. 15 Hollandipsylla 326, 333, 339, 411, 412, 423, 424, 428, 429 Hoogstraal, H. 6 é 165-186 Hoogstraalia 348, 350, 366 Hookeriaceae 5 : 5 : 19 hopkinsi, Gryphopsylla Pl. 8, Tab. 7 (opp. p. 406), 416 a 3 a Fee tS) 1I7, 145 hopkinsi, Stivalius Hopkinsipsylla horsfieldi, Cynopterus horsfieldii, lomys 106, 121, 136, 146, 149, 171, 193, 196, 236, 298 327, 330 153,171 horsfieldii, Myotis . < a 31, 32 horsfieldii, Myotis horsfieldii 31-32, 40 horsfieldii, Vespertilio . F z si BY horsfieldii horsfieldii, ee 31-32, 40 hosei, Lariscus 396, 397, 398 hosei, Paralariscus 5 396, 397, 398 hosii, Rana . a Fs q ei 65,95 howardi, Orchopeas c 315, 358, 441 howardi, Orchopeas howardi . Fig see howardi bolivari, Orchopeas_ . P ip a2 howardi howardi, Orchopeas_ . : ple) a hubbardi, Stenistomera . C a8 133% huttoni, Murina c 38, 40, 120, 148 huttoni rubella, Murina . 5 c : 38 Hyla . c : g ‘ ‘ 70, 72 Hylaplesia . a - 5 F 53-54 Hylobates ; : 120, 148, 153 Hylobatidae . : F e = /£20 hylobatis, Haemaphysalis 167, 168, 170, 174, 183, 184 Hylomys - 125, 273, 401, Tab. 7 (opp. p. 406), 409, 415, 437, 438 Hylopetes 121, 149, 171, 260, — 412, 429, 440 Hylorana 0 6 67 Hymenophyliaceas: , 5 ‘ : 20 Hymenophyllum . : : . <0 S20 hyparetes, Xiphiopsylla . : 2 o5ge2 Hyperacrius . : A 337, 347 hyperythra, Muscicapa 117, 134, 144 Hypnaceae . : o 5 19 Hypnodendraceae . 3 : F az) Hypnodendron é . 3 4 2 820) Hypogramma 117, 134, 145 hypogrammicum, Hy pogrammna 117, 134, 145 hypoleucos, Pomatorhinus 115, 133, 142, 200 Hypothymis . 117, 134, 145 Hypsiboas_ . : : 2 oS INDEX 459 Hypsipetes 114, 126, 133, 141, 142, 154, 199, 200 Hystrichopsylla -343, 347, 351, 358 Hystrichopsyllidae 411, 413 Hystricidae . - 5 3 6 22 Hystrix : r 2 5 LOE Ichthyophiidae .. : “ on 7k) Ichthyophis . 5 2 ~ 748 Ictinaetus 9 108, 132, 738, 154, 199 Idilla . : 325, 333 idoneus, Macrostylophora Tab. 5 (opp. p. 402) igneus, Pericrocotus c a - 4154 ignipectus, Dicaeum 5 117, 145 Iguanognathus 3 a 9 0, BS imbricatus, Podocarpus . 7 : 14 impar, Walchiella . 192, 195, 196 inas, Lenothrix ‘ : : ae 390 inas, Rattus 122, 136, 146, 149, 171, 174, 195, 196, 396 indages, Monopsyllus_~. 7 PING indica, Ascoschoengastia 193, 196 indica, Bandicota . ; 2 - 1397, indica, Chalcophaps 109, 132, 138 indica, Hinulia a ci : A = 83 indica, Pallavicinia $ 3 P 17 Indicator : F 112, 140, 200 Indicatoridae 5 R 5 7 lee indicus, Sphenomorphus | 5 “ Pass indicus, Sphenomorphus indicus = 83, 96 indicus, Tapirus 5 123, 150 indicus indicus, Sphenomorphus é 83, 96 indicus zebratum, Sphenomorphus . MARES 3 infestus, Libyastus B14 ble inflatus, Coelops. . c é : ‘i 29 infraluteus, Rattus 396, 397, 398, 401, Tab. 7 (opp. p. 406), 409 396, 397, 398, 401, Tab. 7 (opp. p. 406), 409 ingens, Cryptoctenopsyllus 331, 346 ingens, Dinopsyllus ‘ = ; earSge infraluteus, Stenomys inopina, Neopsylla 315, Pl. 6 inopinatum, ee, 108, 138 Insecta 187-450, 78 pls insigne, Gahrliepia_ 194, 196 insignis, Lariscus 121, 136, ane, 149, 170, 171, 177, 181, 192, 196, 216, 235, 236, 243, 247, 250, 252, 259, 261, 262, 263, 298, 396, 397, 398, Tab. 6 (opp. p. vee 405, 415 insignis bardus, Apomys ci = hele) insignis niobe, Lariscus . - 228 insolli, Lentistivalius . 272, Aye, A, 275, 280, 305, Pl. 47, Pl. 50, 312, 342, 366, Pl. 3, 416, 418 insolli, Stivalius . ‘ hi P e272 intestinalis, Aspis . s 5 5 - 93 intestinalis, Maticora 5 é 3 93, 96 intorquatum, Ectropothecium ‘ 5 19 Iomys 106, 121, 136, 146, 149, 171, 193, 196, 236, 298 Irena . : S . 113, 133, 141 Ischnopsyltus 0 : x D 6 333 Ixalus . 5 z : 5 70, 71 Ixobrychus . Bad, St5S Ixodes . 167, 170, 172 ee 183, 184, 196 Ixodidae a é 165-186 jacobsoni, Stivalius 367, Tab. 7 (opp. p. 406), 407, = 418, 440 jacobsoni-group, Stivalius 2, 403, oy jalorensis, Rattus tiomanicus 171, oe. 216, 2 jambu, Ptilinopus : cl 5 se japonica, Rhadinopsylla fi - 412 jasoni, Macrocalamus 88-92, 96 jaspidea, Boiga : 47 javana, Medwayella_. 217, 2 228, Pay 256-257, 267, 268, 303-305, PI. 15, Pl. 31, Pls. 43-44, Tab. 5 (opp. p. 402), Tab. 6 (opp. p. 404), Tab. 7 7 (ep. p- 406), 416 javanica, Freycinetia. f 13 javanica, Manis -121, 148, 153,177 javanica, Nycteris . : 4 2 eE53 javanica, Tupaia H 256, 298, 400 javanica occidentalis, Tupaia . 256, 298 javanicum, Paraceras Tab. 5 (opp. p. 402), 416 javanicus, Eurylaimus 112, 133, 135, 140 javanicus, Phaenicophaeus 109, 132, 139, 199 javanicus, Pipistrellus 40, 120, 147 javanicus, Pipistrellus javanicus D -. 33 javanicus, Tragulus 123, 136, 150, 153, 171, 174, 175, 176, 177, 178, 181, 183, 196 javanicus javanicus, Pipistrellus 6 > | 65) javanus, Medwayella . A é ~ (205 javanus, Stivalius . 215, 256 javensis, Dryocopus A - a Spel 5 4 Jellisonia H 5 391,35 jentinki, Gundasciurus é i 397, 398 jerdoni-group, Rattus. : : 157 jettmari, aeboad Sue 21328; PL 18 Johnson, Anne 17-20, 8 Pls. Jordanopsylla 324, 330 Jungermaniales Acrogynae 2 18 junghunianum, Sphagnum c 0 14,17 Kalophrynus : 2 5 3 55,95 kasumba, Harpactes 6 i : - 154 keenani, Kohlsia_ . z : 2339) keeni, Opisodasys . 314, 315, 321, Pl. 7 kendallii, Cnemaspis - iS 80, 96 kendallii, Heteronota. P : eso Kenopia 5 4 5 II5, 143 kerguelensis, Notiopsylla : 325, 348 Kerivoula . . - 2 3 120, 148 Kibatalia ° " 5 2 5 12 kienerii, Hieraaetus : F : 108, 138 Kitta . : : 5 FideettIe4 klossi, Rhinclophus , : 0 24,2 klossi, Rhinolophus robinsoni . - 25-27, 40 460 INDEX klossi, Stivalius . 232, 233, 285, 315, Pl. 3, 403, 406, Tab. 7 (opp. p. ely, 407, 416, 418 Knema o 5 z 5 ° 13 Kohlsia 316, 331, 339, 351, Pl. 2, 411, 412, 414, 425 koningsbergeri, Haemaphysalis 167, 168, 170, 177-178, 183, 184 Koompassia . 12, Plan err kopetdag, Phaenopsylla . ‘ = Pl. 13 kopsteini, Rothschildiana Tab. 7 (opp. p. 406), 416 kukenthallii, Oreobolus . : 14 kundini, Leptotrombidium 191, 196 lacunosa, Walchiella C 2 j A pd Laemobothriidae . q - - - 199 Laemobothrion 4 ; ; 3 [LOO laetitiae, Ornithopsylla . . é ses lafresnayei, Aegithina 113, 141, 154 Lagaropsylla A : 6 sss lagochilus, Macroglossus : 5 iE lancifolia, Trismegista. 5 : : 19 langati, Leptotrombidium ° = LOE Lanius . 3 5 4 4 - 154 laptevi, Caenopsylla : 5 3 Pl. 11 lar, Hylobates 120, 148, 153 Lariscus 121, 136, AG 149, 170, 171, 177, 181, 192, 196, 216, 226, 227, 228, 235, 236, 241, 243, 247, 250, 252, 256, 259, 261, 262, 263, 264, 298, 319, 396, 397, 398, Tab. 6 (opp. p. 404), 405, 415 larutensis, Amolops 5 Aone 62, 64, 68, 95 larutensis, Ixalus . é 5 5 7o larutensis, Lygosoma - 5 - 46 larutensis, Rana. . : 2 46, 64 larutensis, Siaphos 5 5 r 5 46 lateralis, Macrocalamus . 47, 90-92 lateralis, Rhacophorus reinwardtii . 5 75 laticaudatus, Rhinosciurus 121, 136, 146, 149, 153, 170, 171, 174, 193, 196, 215, 216, 227, 242, 243, 244, 259, 261, 262, 263, 298, 397, 398, Tab. 6 > (eRe: P: 404), 415 laticeps, Rana 65-66, 95 latidisca, Ansonia . 6 : 50 latirostris, Musicapa : 116, 124, 144 laxata, Palaeopsylla Tab. 7 (opp. p. 406), 416 Leiothrix : : : 5 ae Lejeuneaceae . c 18 Lenothrix 160, 261, 208, 306, 397, 401, Tab. 7 (opp. p- 406), 415, 432 Lentistivalius 269-281, 282, 290, 291, 292, 305, Pls. 46-52, 312, 314, 338, 342, 366, Pl. 3, 401, 402, 403, Tab. 6 (opp. p. 404), 406, Tab. 7 (opp. P- 406) 410, 411, 416, 417, 418, 420, 421, 438, 442 Leopoldamys 261, naar 397, 398, 415, 433 Lepidosperma , : 14 Leptobrachium ve 55> sa, 60-62, 95 Leptopsylla . - 350, Pl. 8, 408, 431, 432, 437 leptopus, Ansonia . 5 ¢ 49, 5 leptopus, Bufo - B Leptospermum Si Pls. os Leptotrombidium . 189, 190, I9I, 195, 196, 197 Lepturophis . 87-88, 89, 96 leschenaulti, Enicurus 116, 134, 144 Leucobryaceae c c : an eat Leucobryum 2 : c “ A 18 leucomystax, Hyla é A . aad a7 2 leucomystax, Rhacophorus. . O73 leucomystax, Rhacophorus leneounpeeoe 72-73, 95 leucomystax leucomystax, Rhacophorus 72-73, 95 Leuconoe < 0 : : 5 kz Leucophanes c 4 2 18 leucophrys, Brachypteryx . 5 L25 leucopterus, Platysmurus = (154 leucopus, Orchopeas 404, 315, Pl. 5, 441 leucopygialis, Chaetura . 110, 139 leucotis, Stachyris. 115, eat 135, 143, 154, 199 leucura, Myiomela : 5 +, $125 levanticus, Ctenophthalmus levanticus o 1A levanticus levanticus, Ctenophthalmus . Pl. 4 leveiri, Pallavicinia 5 A 17 levis, Macrostylophora Tab. 5 (opp. Pp. 402), Tab. 6 (opp. p. 404), 406, 416 liae, Macrostylophora Tab. 5 (opp. p. 402), 404, 429, 442 Libyastus 314, 326, 333, 338, 340, Pl. 2, 411, 412, 416, 424, 425 lice 5 ; . D c - 199-200 Lim, B. L. 165-186 limi, Medwayella 242-244, 259, 260, 261, 262, 263, 264, 265, 267, 268, 303, Pls. 26-28, Tab. 5 (opp. p. 402), Ae: 416, 420 limnocharis, Rana . 66 limnocharis, Rana liunochacis, : ‘66; 95 limnocharis limnocharis, Rana ° -66, 95 Limnodytes . R 5 2 ¢ om, 166) linsang, Priondodon 122, 149 Liopeltis c . 85-86, 96 Listronius . ee gO2 loncha, Medwayella " ax7, 228, 231, 239, 244, 248, 249, 250, 252, 253, 255-256, 266, 268, 302, 304, Pl. 1, Pl. 38, Pls. 41-42, Tab. 5 (opp. p. 402), Tab. 6 > (OPP. P- 404), 416 lonchus, Medwayella_ . 2 2 eRe) lonchus, Stivalius . 215, 255 londiniensis, Nosopsyllus Pl. 8, 416 londiniensis, Nosopsyllus londiniensis ion 35 londiniensis londiniensis, Nosopsyllus apes longicauda, Liopeltis c 85-86, 96 longicaudatus, Hapalomys < : 3 - 397 longicaudaus, Ablabes_ . 3 : Bie} longicaulis, Bazzania 5 : : < 18 longiceps, Calamaria z < é <4.) 86 longiceps, Pseudorhabdion Ss : 86, 96 longidigita, Ansonia : : : - 49 longifolium, Calymperes : S oe, 29 INDEX 461 longimanus, Heterometrus 190, 196 longipennis, Hemiprocne : - - 154 longirostra, Arachnothera : oA longirostris, Arachnothera 117, 134, I 1355 145, 146 longirostris, Rhizothera 109, 138, 154 longisetosa, Genoneopsylla 4 6 24 longissima, Aerobryopsis 6 d 19 Lophura 153, 171, ve Lophyurus 5 : cs Lorentzimys . A : : = ie loricata, Napothera 115, 142 Loriculus 0 5 7 109, 138 Lorisidae . A : a 59) lowi gimletti, Calamaria_ 3 F 5 eG lowii, Ptilocercus 120, 148, 171, 192, 193, 194, 195, 216, 252, 254, 298, 400, 415 lowii, Sundasciurus 121, 136, 146, 149, 153, 171, 192, 193, 196, 227, 241, 252, 254, 256, 259, 260, 262, 265, 298, 397, 398, Tab. 6 (opp. p. 404), 415 lucasi, Penthetor - IIQ, 147, 153, 170, 172 lucida, Payena 5 5 : ews luctuosa, Rana : 66-67, 95 luctuosus, Limnodytes_ . 6 5 = 166, luctus, Rhinolophus 0 0 ; Ets S lugubris, Garrulax 115, 143 lugubris, Surniculus 109, 132, 138 luma, Neopsylla 315, Pl. 6, Tab. 7 (opp. p. 406) 409, 416 47, 86-87, 96 lumbricoidea, Calamaria : Tab. 5 (opp. p. 402) lupata, — oo Lycodon 4 0 ‘ 47, 87 Lycopsylla 9 325, 326 Lygosoma . C 5 ° 46, 82 lylei, Hipposideros. P 3 28, 40 Mabuya : 82-83, 96 Macaca 9 +120, 43 153,171 Macaranga . f 12 macclellandii, Hypsipetes I14, 133, 141 macclellandii, Tamiops . 121, 136, 149, 216, 241, 259, 260, 208, SOS TG Macrocalamus 2 2 43-101 macrodactyla, Napothera I 15, 133, 142, 191, 195 macrodon blythi, Rana . 9 m1) 104 Macroglossus - I19, 136, 147, 153,171 Macronus ¢ 3 5 115, 133, 143 Macropisthodon . 4 5 a 84, 96 macrops, Pseudoxenodon g = Ray Macropygia . 109, 124, 132, 135, 138, 154 macrorhynchos, Cymbirhynchus 112, 133, 135, 140, 154 Macrostylophora 264, 314, 316, 319, 339, 347, Pl. 1, Pl. 2, Pl. 19, Tab. 5 (opp. p. 402), 403, 404, Tab. 6 (opp. p. 404), 405, 406, Tab. 7 (opp. p. 406) 41, 412, 414, 416, 422, 423, 424, 425, 428, 420, 439, 440, 442 macrotis, Rhacophorus . c . 73,95 macrotis, Rheithrosciurus 396, 397, 398 macrotis, Rhinolophus 27, 120, 136, 147, 171, 192, 195 macrotis, Rhinolophus macrotis 4 an R27) macrotis caldwelli, Rhinolophus 0 27, 28 macrotis dohrni, Rhinolophus 27-28, 40 macrotis episcopus, Rhinolophus_—. a2 7, macrotis macrotis, Rhinolophus 5 J 27 macrotis siamensis, Rhinolophus’~ . 27, 28 macrourus, Atherurus 5 122, 149 maculata, Balionycteris 119, 136, 147, 153, I71 maculata, Stachyris 115, 133, 143 maculatus, Prionochilus . I17, 134,145 magna, Arachnothera magnirostre, Malacopteron - 117, 134, 145, 199 - 115, 133, 135, 142 magnum, Malacopteron . 115, 133, 142 major, Chiropodomys_ . 4 a a3 97; malabaricus, Copsychus . 116, 134, 144 malaccana, Asthenodipsas “ “ een OZ malaccanus, Pareas a 5 a 92,96 malaccense, Trichastoma 115, 133, 142, 177 malaccensis, Hypsipetes f : oe LA: malacensis, Anthreptes . c E E54. Malacopsylla - 352, 361 Malacopteron : + 115, 133, 135, 142, 154 Malaraeus 8 351, 441 malayana, Ansonia 5 & 5 50-52 malayana, Gentiana os : ay malayanus, Anthracoceros 154, 171, 182 malayanus, Chrysococcyx 9 a oed54. malayanus, Helarctos 122, 149, 181 malayanus, Rhinolophus 23-24, 40 malayensis, Ictinaetus . 108, 132, 138, 154, 199 Mallophaga F 199-200 Mallotus “ 12 mammals 2I- np, 103- 154 nee 163, I pl. mammosum, Trichosteleum 2 19 maniculatus, Peromyscus : : . 441 Manidae 5 d ° eens Manis . a ‘ 121, 148, 153,177 margarettae, Hacromys 5 397, 398 margaritatum, Gonyosoma_ . fs at 185 margaritatus, Gonyophis é 3 85, 96 marikovskii, Amphipsylla 327, 333, Pl. 20 marmoratus, Cyrtodactylus 72, 81, 96 marmoratus, Phyllurus . “ : ee Marmota : 2 - 356, Pl. 13 Martes . . : - B n34r mathesoni, Pleochaetis 5 6 : #3315 Maticora . 5 : s 93,96 maxima, Collocalia . 6 : = DLO) maximum, Laemobothrion 5 a LOG maximus, Draco . . fs 77,96 maximus, Draco maximus maximus, Elephas 123, 150, 153 maximus maximus, Draco 77 Maxomys . . 261, 298, 307, ae 401, Tab. 7 (opp. Pp. 406), IBD ce 432, 438 maxwelliana, Shorea Q : 13 - 39, 40, 173 1-7, 4 Pls., 103-154 medius, Miniopterus Medway, Lerd 462 INDEX Medwayella 211-270, 273, 282-283, 290, 292, 302-305, Pls. 1-45, 312, 314, 315, 316, 359, 367, Pl. 1, 392, 393, 396, 399, 401, 402, Tab. 5 (opp. p. 402), 403, 404, Tab. 6 (opp. p. 404) 405, 406, Tab. 7 (opp. p. 406), 410, 411, 412, 414, 416, 418, 419, 420, 421, 422, 423, 424, 428, 438, 440, 442, 443 medwayi, Myotrombicula 5 192, 195 Megabothris . oi : J 2) 305 Megaerops 119, 136, 147 Megalaima III-I12, 126, 132, 140, 200 Megalophrys . Hee d62 megaphyllus-subgroup, Rhinolophus Se ee25 Megarthroglossus . 324, 333, 335, 34 352, 355, Pl. 1, Pl. 12, 411, 412 Megophrys 4 62-64, 95 meifolium, Trichomanes > 5 20 Meiglyptes : 112, 132, 140, 154 melalophos, Presbytis 120, 153 melanocephalus, Chironax 119, 126, 136, 147, 171 Melanochlora 114, 133, 142 melanoleucos, Pyenonetus 114, 154 melanopogon, Draco 5 : 2 78, 96 melanotis, Nannosciurus 397, 398, 401 melanotis, Pteruthius 115, 143 melanura, Dendrogale - 272, 400, Tab. 7 (Opp: p- 406) melanurus, Pithecheir 397, 415 Tab. 5 (Opp. Pp. 402) Tab. 5 (opp. p. 402) melinum, Paraceras melis, Paraceras Melogale " e 3 P ~ 2433 Menacanthus : - 199 Menetes 242, 244, We, 247, 208) 306, 397, 399, Tab. 6 (opp. p. 404) Menoponidae : ; : 5 S199 mentalis, Picus 112, 140 Meringis 5 9 324, 333 Meriones 7 5 5 Z di TAL iG} Meropidae . c : 7 )EDE Merops : < III, 135, 139 Mesopsylla_ . a , : ag 47, Metaphrynella 45: 49, 47, 55-57, 95 Metastivalius . 5 - q 331, 430 Meteoriaceae 5 3 A : 19 Metzgeria. . a 2 5 F 17 Metzgeriaceae a fs ¢ ‘ 17 mexicana, Rhadinopsylla 5 : Pl. 11 mexicanus, Reithrodontomys . 3 «340 meyenianum, Thuidium . : 19 Microhierax . 108, TA5) 138, 154 Microhyla ° 40, 57-60, 95 Microhylidae 5 : 54-60 microphthalmus, Paradoxopsyllus 327 Plans Micropternus : - : é - 154 micropterus, Cuculus 109, 138 Microtus z 0 : 2 S SUe3r5 Microura c : 115, 143 mikulini, Brevictenidia z . P ps 27; Millardia a F 5 0 5 285, 298 mina, Strepsylla . Y : < Pl. 13 287, 288, 290, 298 245, 298 mindanensis, Rattus rattus miniatus, Callosciurus notatus minimus, Macroglossus 119, 136, 147, 153 Miniopertus . 2 39-40, 172, 173 Minla . 115, 125, 134, 143 minor, Tupaia 120, 146; 148, 153, I71, 192, 195, 216, 259, 298, 399, 400, 415 Mioctenopsylla - 342, 352, 361, 362 miopus, Rana 3 z 5 67,95 mitratus, Garrulax ris, 134, 143, 199, 200 mittenii, Distichophyllum : 6 19 mjoebergi, Stivalivs 255, 406, Tab. 7 (opp. p. 406) 416, 418, 420 mjoebergi-group, Stivalius 402, 403, 406 modestum, Piprisoma_. 2 SoG) moluccensis, Dendrocopos moluccensis + EF moluccensis moluccensis, Dendrocopos . 171 Monitor ¢ 3 79 Monopsyllus . 313, 315, 316, 317, 321, 339, 355, 362, Pl. 1, Pl. 9, Pl. 10, 403, 410, 411, 412, 413, 414, 416, 423, 425, 429 montana, Amphipsylla 1327,.b1 20 montana, Tupaia 272, 280, 400 montana aceras, Megalophrys . ¢ aeG2 montanus, Pomatorhinus I15, 133, 142, 154 monticola, Emballonura I1Q, 147, 171 monticola, Trimeresurus i ‘ buoy 474 monticola nasuta, Megophrys . 63-64, 95 montis, Seicercus , 116, 134, 144 mooseri, Rickettsia 5 : 193 Motacilla : , 5 r PeeLrS Motacillidae . c II3 muelleri, Rattus 122, , 124, 136, 146, 149, 153, 170, 172, 191, 192, 193, 196, 216, 227, 298, 397, 398, 415 muelleri, Stenomys a 397, 398, 415 Muelleripicus 5 : : - 154 Muesebeckella a a : - 339 Miilleri, Typhlops . 7 : ef ot 83 mulleri, Typhlops diardi - 47, 83,96 mulleri-group, Chiastopsylla . < ees20) multifasciata, Mabuya . ‘ - 82-83, 96 multifasciatus, Scincus . E A 82 Muntiacus 123, 150, 153 muntjak, Muntiacus 123, 150, 153 Muridae : é » HZZ Murina . 38- 30, rey 120, 136, 148, 153 murina, Dendrogale 5 399, 400 muroides, Chiropodomys 4 ; = 397 Mus. 5 : : : Bia Pl. 8, 2) Musci : Muscicapa n16-117, 124) 134, vn 145, os Muscicapidae 116-117 musculus, Mus c : G = =) 9397 Mustela : “ c a SE mutabilis, efeleniGnies 193, 196 mutilatus, Gehyra . $ ; : 82, 06 mutilatus, Hemidactylus “ é Phe (2175 Myiomela. 4 ¢ “ . = R25 Myodopsylla : 7 ; R - 318 INDEX 463 myodosus, Ctenophthalmus expansus Pla Myospalax . a : a “9 328 Myotis . 30-33, 40, 153, 172 Myotrombicula 190, 192, 195 Myoxopsylla 326, 333, 339 Myrsidea ; 5 5 5 7 299 mystacalis, Kethopyea 117, 145, 154 mystacinus, Myotis , c ‘ eet53 mystacophanos, Megalaima III, 140 Nadchatram, M. 165-186, 187-198 nadchatrami, Haemaphysalis 167, 168, 170, 171, getles oe 182, 183, 184 Naja . ; o S. titehs naja, Naja . 5 d eereT nakagawai, Frontopsylla c . 408 Nannosciurus - 396, 397. 308, 401, 440 Napothera 115, 126, 133, 135, 142, I9I, 195 napu, Tragulus - 171,174, 176, 177, 183 nasuta, Ceratophryne . i 3 ; 63 nasuta, Megophrys monticola . - 63-64,95 Natricidae. . é e : 83-84 Natrix . 47, 83-84, 96 neali, Hollandipsylla 339, 424 Nearctopsylla 5 325, 333 nebulosus, Monitor 9 z f 79 nebulosus, Varanus bengalensis : 79, 96 Neckeraceae . 3 - é 4 P 19 Nectarinia . D 9 0 7 7 154) Nectariniidae 5 6 : 4 gts Nectophryne 5 a 3 - ees 4 negrinus, Rattus rattus . 287, 298 nemestrina, Macaca 120, 148, 153 Neoctenophthalmus - . 5 Spas} Neohaematopinus . 2 sae200) Neopsylla 233, 314, 315, 317, 322, 32 4, 333, 347, 351, 354, Pl. 6, Tab. 7 (opp. p. 406), 408, 409, 416, 432, 436, 439 Neoschoengastia 190, 192, 195 Neotoma 314, 315, 320, 321, 335, Pl. 5, Pl. 11, Pl. 12, Pl. 14, 411, 412 neotomae Orchopeas 315, 320, 321, Pl. 5 Neotomodon : = 326; Pl. 13 nepos, Orchopeas nepos . 315, Pl. 5 nepos nepos, Orchopeas . 315, Pl. 5 nesiotes, Xenopsylla : 3 “ >) 433 nesiotus, Opisodasys 314, Pl.7 neteralla, Gahrliepia 194, 196 nicobaria, Tupaia . 5 6 “ + 399 nicobariensis, Hylorana . 5 b oleh nicobariensis, Rana : a - 67-68, 95 nidicola, Conorhinopsylla 0 2320, Pina nigriceps, Stachyris - 115, 133, 143, 154 nigricollis, Stachyris - 115, 133, 143, 154 nigroflava, Caudacaecilia : i 48,95 nigroflavus, Ichthyophis “ = 48 nigropalmatus, Rhacophorus . 59, 73-74; 75,95 nigrovittatus, Callosciurus 121, 136, 146, 148, 070, 193, 196, 227, 228, 2209, 232, 235, 236, 259, 263, 397, 398, Tab. 6 (opp. p. 404), 405, 415, 441, 442 nigrovittatus, Callosciurus nigrovittatus 256, 298 nigrovittatus, Sciurus. 228 nigrovittatus nigrovittatus, Callosciurus 256, 298 Ninox . 5 9 3 S 2) 0054 niobe, Lariscus dl 3 5 ° e220 niobe, Lariscus insignis . 6 228 nipalensis, Cutia 115, 134, 143 nipalensis, Pyrrhula 117, 134, 135, 145 niviventer, Maxomys F : = 397 niviventer, Rattus - 7 : 397 norvegicus, Rattus 193, 397 Nosopsyllus ane aye 354, PL 2, PL 8, 409, 411, 2, 413, 415, 416, 423, 424, 425 notatus, @hitessniras 121, 136, 146, 148, 153, 171, 192, 193, 196, 200, 216, 227, 232, 234, 235, 236, 259, 261, 263, 298, 397, 398, 399, 404, Tab. 6 (opp. p- 404), 405, 415, 441, 442 notatus miniatus, Callosciurus 245, 298 Notiopsylla c 325, 348, 362 nouhuysi, Lorentzimys . 5 5 - 442 novaeguineae, Parastivalius . D a 3a7/ novaehollandiae, Coracina 113, 133, 140 nudipes, Mustela. 5 c ~ 8x numae, Chiastopsylla_. 3 5 - 329 numae-group, Chiastopsylla_. S29 Nycteridopsylla om: 333, Baa: 352, Pl. 15 Nycteris 0 : SB tiG ie} Nycticebus 120, 148, 171 Nyctiornis III, 132, 139, 154 oblongifolia monticola, Knema I 13 obscura, Presbytis 120, 136, 148, 153 obscura, Wenzella . 5 - 345, Pl. 14 obscurum, Rhododendron 5 : : 14 obscurus, Presbytis 3 4 3 eel L occidentalis, Tupaia javanica . 256, 298 ocellata, Rheinardia E 5 2 - 109 ochi, Choristopsylla a ; gt Ochotona AGG, PL. 11, 410 ochraceiventer, enoehrie 397, 398 ochraceiventer, Rattus 397, 398 ochraceus, Criniger - I14, 126, 133, 41, 199 ochromalus, Eurylaimus 112, 140 octoblepharoides, Leucophanes : : 18 octodecimdentata, Rae Pl. 1, 410, 441 Octomys , bs : - 329 Odontacarus . 190, 192, 195 olax, Treron . - a a 5 53 Onychomys . 315, Pl. 9 oorti, Megalaima Lis, "126, 132, 140, 200 Ophthalmopsylla 328, 347, Pl. 18 Opisocrostis . 350, 356 Opisodasys 314, 315, 317, 321, 326, 327, 333, 339, 341, 342, 344, 368, Pl. 1, Pl. 7, Pl. 15, 411, 412 414 Orchopeas 314, 315, 316, 317, 320, 321, 339, 358, 359, Pl. 1, Pl. 5, 411, 412, 414, 441 oregonensis, Paratyphloceras . a - 343 oregonensis, Trichopsylloides . 343, 368 Oreobolus” . f é 6 5 " 14 464 INDEX oreskios, Harpactes 4 110, 139 Orfrontia A d 2355 orientalis everetti, Helictis F 5 »* 433 orientalis everetti, Melogale . : 2 8433 Oriolidae 2 c : S 7 en) Oriolus 5 II4, 126, 135, 142 orites, Ceratophyllus 4 3 5 e320 ornata, Ansonia. : : C + 50 ornata, Corypsylla “ 5 : es 52 Ornithophaga . 342, 346 Ornithopsylla 328, 342, 362 Oropsylla 0 5 349, 356 orrhophaeus, Harpactes . 3 - 154 Orthotomus . 2 : 116, 134, 144, 154 Orthotrichaceae . : : 2 19 Oryzomys . 5 2 340, 414 oschanini, Echidnophaga A . 2359 Otocryptis . : : : . 4 76 Otomys 5 5 A > 0 29) Otis); 2 II0, 132, 139, 171 oudemansi, Walchiella 192-193, 195, 196 Oxyparius . 0 3 9 : 2 6333 pacificus, Apus 5 - 110 Palaeopsylla = 337, 343. 347. Tab. 7 (opp. Pp. ele 409, Be 417, 5 432, 437 Palaquium . c 13 Pallavicinia . : : 2 2 17 pamirensis, Leptopsylla 3 : Pl. 8 paniculatus, Mallotus” . 3 9 ~) ei2 pantarensis, Rattus . : - . 288 Panthera 123, 150, 181 papillosa, Kerivoula : 120, 148 pappillata, Callicostella . : 2 5 19 Papuapsylla . : : 3 - 319 papuensis, Xenopsylla 317, 318, 328, 333, 344, 350, Pl. 17, 408 Paraceras 316, 349, Tab. 5 (opp. p. 402), 404, Tab. 7 (opp. p. 406), 409, 411, 415, 416, 423, 424, 428, 433 Paractenopsyllus . 5 5 Ceo paradisea, Phalacropsylla 321, Pl. 12 paradiseus, Dicrurus 114, 133, 142 paradisi, Chrysopelea. : men gy paradisi, Terpsiphone 117, 134, 145, 154, 200 Paradoxopsyllus ° 327, Pl. 13 Paradoxurus 153, 170, 177, 181 Paralariscus . : . 396 paramundus, Pleochaetis 321, 326, 327, 330, Pl. 11 Paraneopsylla 324, 333 Parapsyllus . : 329, 362 Parastivalius. a : 5 : Bes 07; Paratyphloceras 333, 343 pardus, Panthera . 123, 150 Pareas . 46, 92, 96 Paridae 2 ; 0 F c eer) Pariodontis . 9 A : 4 + 349 parvula, Riccardia b 5 5 5 17 parvus, Bufo. < “ : : 53, 95 pavimentata, Calamaria 5 é a ets) Payena 5 c i 0 0 - 13 Pectinoctenus 408, 432 pellax, Rattus rajah : 157, 158, 161 Pellorneum < 115, 133, 135, 142 Pelobatidae . 5 é 60-64 pelodytoides, Leptobrachium e : “a 26x Pelophryne . a F cs 54, 95 penangensis, Ansonia. » 49 pendleburyi, Paraceras Tab. 5 (opp. Pp- 402), 416 Penenirmus . z + 200 penicilliger, Malaraeus : B : « 44% peninsularis, Murina cyclotis 38, 39, 40 Penthetor 119, 147, 153, 170, 172, 183 peracensis, Alcippe 115, 134, 143, 146 peregrinata, Medwayella robinsoni 234-235, 236, 266, 269, 303, Pls. 19-21 Pericrocotus . f 113, 133, 141, 154 perlata, ae 116, 134, 144, 154 Pernis ‘i a) e053 Peromyscopsylla 331, 350, 351, 408, 431 Peromyscus 314, 315, 321, 326, 335, 362, Pl. 5, Pepe) bl-mr, Pl 12; Piro) 40Snamrraie. 441 perse, Callosciurus hippurus . é - 401 Petaurista I2I, 136, 149, 153, I71I, 200, 235, 260, 298, 412, 429 petaurista, Petaurista 121, 136, fie 153, 171, 200 Petaurus > a : : + 342 petersi, Ixalus c c a c me) 70 petersi, Philautus . 5 5 F Pre Petinomys . 3 5 6 + 440 Petrochelidon P 342 Phaenicophaeus 109, 132, 138-139, 154, 199 Phaenopsylla 327, 328, Pl. 13, 442 phaeocephalus, Criniger . 113, 133, 135 Phalacropsylla 321, 324, 330, 333, 352, Pl. 12 phangi, Medwayella 237-240, 242, 243, 244, 252, 257, 258, 259, 260, 261, 262, 263, 264, 265, 267, 268, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. p. 404), Tab. 7 (opp. p. 406), 416, 420 phangi, Medwayella phangi 216, 240-242, 261, 267, 268, 303, Pls. 23-26 phangi phangi, Medwayella 216, 240-242, 261, 267, 268, 303, Pls. 23-26 phangi tana, Medwayella 242, 250, Be 268, 303, Pl. 23, Pl. 25 Phasianidae . 108-109 Philautus Les 47, 62, 70-71, 95 Philentoma 117, 134, 145, I9I, 195 philippinus, Merops : ~ FOr phillipsi, os wgbeaea Tab. 5 (opp. p. 402) Philopteridae 5 5 2/599. Philopterus . my 2zO0) phoberus, Stivalius 281, 284, 285, 291, 292, 293, 305, Pl. 52, 367, 368 Phodilus E 110, 132, 139 Phoenicophaeus” . c 0 Be ay Phrynella 55, 60, 95 INDEX 465 Phthiraptera F 5 199-200 Phthiropsylla ; : ~ 3525-301 Phylloscopus. : 116, as 134, 144 Phyllurus”. 81 picaoides, Heterophasia : I 115, 1345 143, 199 picatus, Hemipus . 0 - 113, 141 Picidae F 2 : . =) Dt2 picta, Gahrliepia . a : - 194, 196 picta, Tupaia : 5 c 2 + 400 Picus . é 5 112, 125, 140 pilata, Macrostylophora Tab. 5 (opp. p. ye Tab. 6 (opp. p. 404) pileata, Halcyon . 0 : neeli5 4: Pinanga 6 " 14 Pingue, Gahrliepia disparunguis 195, 196 Pingue, Walchia disparunguis . 195, 196 pinguis, Riccardia . 4 B 17 Pipistrellus 2 oe 120, 147, 148 Piprisoma . : meee?) pitchfordi, Chiastopsyla. 0 - 329 Pithecheir a 306, 397, 415 Pitta 112, 133, 140, 1545 190, I9I, 192, 193, 195, 200 Pittidae 3 5 5 2 < eer T2 Platylophus . 0 114, 133, 142 Platysmurus 3 é : : - 154 platysoma, Ansonia ° 2 ~ 50 Plectocomia . a a 0 0 3 13 Pleochaetis 315, 316, 321, 326, 327, 330, 339, Pl. 2, Pl. rz, 411, 412, 414 pleurostigma, Kalophrynus_. 55 pleurostigma, Kalophrynus pleurostigma 55, 95 pleurostigma oa eae 55, 95 Plicatella, Rana. : 68, > Podocarpus . Pogonomys poliocephala, Stachyris . pollex, Ctenophthalmus pollicaris, Metaphrynella 328, 350, 358, Pl. 17, 408 115, 133, 143 313, 315, 322, Pl. 4 2, 46, 47, 55-57, 95 pollicaris, Phrynella 2 3 SS Polydontophis 4 6 . ° - 46 polymerus, Ectinorus . 9 “0 S20) polymorpha, Pinanga cS Polypedetes . 65, 68, 72 Polyplectron 108, 138 Pomatorhinus “115, 133, 142, 154, 200 pomerantzi, Stivalius Tab. 7 (opp. p. 406), 407 pomerantzi-group, Stivalius 402, 407 ponsi, Pleochaetis . f G 5 + 339 popeorum, Trimeresurus. 2 47, 94, 96 populnea, Symingtonia . 6 0 : 13 porphyracea, Elaphe 5 : 5 a7, potis, Chimaeropsylla 363, 364 powelli, Praopsylla 0 A c = 369 Praopsylla a . 329, 331, 369 praxilla, Acanthopsylla . a er zO2 Presbytis 120, 136, 148, 153, 171 prevostii, Callosciurus 121, 148, 153, 171, 227, 259, 298, 397, 398, 399, 401, 404, Tab. 6 (Opp. Pp. 404), 415 Prinia . ; : : : + 125, 154 Prionochilus . 117, 134, 145 Prionodon . : - 122, 149 probata, Macrostylophora Tab. 5 (opp. p. 402), Tab. 6 (opp. P- 404) procus, Megarthroglossus a Pipa2 prominanus, Rhacophorus 7: 74-75, 95 Prosciurillus . 4 0 > 8 + 404 Prostigmata . 187-198 protinus, Monopsyllus ciliatus > Pl to Psammodynastes . c : ae4g6 pseudarctomys, Opisodasys 314, 333, 339, Pl. 7, 412 Pseudorhabdion . 9 86, 96 Pseudoxenodon. : era, Psilopogon LI2,'r 32, 139, 200 Psittacidae . 5 : é 3 + 109 Psittinus 5 5 9 f ee 5S psorakari, Schoengastia . d - 190 Pteromyscus a 121, 149, 171 Pteropodidae 6 : P B + MELO Pteruthius 5 4 115, 143 Ptilidiaceae . 3 5 6 A a 18 Ptilinopus. eak53 Ptilocercus 120, 148, 17m Sep. 193, 194, 195 216, 252, 254, 298, 399, 400, 415, 420, 439 ptilorhynchus, Pernis . 5 rs s » 153 ptilosus, Macronus 5 ; 115, 133, 143 Ptychorhamphus . “ a 6 eeszs puella, Irena : 113, 133, 141 Puffinus 9 328, 342 pulchellus, Gyrtodactyins : : cer pulchellus, Gymnodactylus_. " ehA7 pulchra, Phrynella a : 60, 95 Pulex . : 345, 346, 349, 358 pulverulentus, Muelleripicus 6 5 nS 4 pulverulentus, Psammodynastes A - 46 pulverulentus, Pteromyscus 121, 149, 171 punctatus, Draco 76, 78, 79, 96 punicea, Cophias . ? 0 F 3 97 puniceus, Picus 3 3 II2, 140 puniceus, Trimeresurus . F 3 96, 97 pusilla, Microura 3 F 115, 143 pusillus, mea 5 : =) 397 Pycnonotus . ae 114, 133, 154 Pygiopsylla 230, 232, 291, 334, 401, 402, Tab. 7 (Opp. p. 406), 407, 416, 417, 418, 420, 421 Pygiopsyllidae 201-305, 58 Pls., 411, 413 Pygiopsyllinae 201-305, 58 Pls. pyrhoptera, Philentoma . 117, 134, 145 pyrolophus, Psilopogon . 112, 132, 139, 200 pyrtrhotis, Blythipicus I12, 140 Pyrrhula rey 134, 135, 145 pyrropyga, Copsychus 3 116, 134, 135, 143 Python 6 9 3 oe 28r quadrisetis, Chiastopsylla 6 é - 329 querceti, Rattus “ 5 287, 298 466 INDEX quinquefasciatus, Draco 2 ‘ 78, 96 quinquefasciatus, Draco quinquefasciatus 78-79 quinquefasciatus quinquefasciatus, Draco 78-79 quirini, Megabothris 5 5 i s 315 quitanus, Pleochaetis dolens . 6 a chil rabori, Rattus 3 0 : 2 - 288 Rafflesia 5 c 3 : ‘ 12 rafflesii, Dinopiam | - 154 rajah, Lenothrix 261, 268, 307; BS) 401, Tab. 7 (opp. Pp. 406), 415 rajah, Rattus 122, 136, 146, 149, 153, 155-163, Pl. 1, 170, 171, 174, 178, 216, 261, 298, 397, 398, 401, Tab. 7 (opp. p. 406), 415 rajah-group, Lenothrix . . - 409 rajah-group, Rattus 157. 227, 245, 409 rajah pellax, Rattus 157, 158, 161 rajah subgroup, Rattus . 6 57 Rallicola ; 4 : . 199 Rana . : 5 AG 64-70, 95 raniceps, Polypedetes < 5 : : 65 raniceps, Rana chalconota : 65, 68, 95 Ranidae - E - 5 64-70 rara, Siseca 190, 195, 196 Rattus 118, 124, 136, 146, 149, 153, 155-163, Ply ry x70} L7T, L72) 174, WSLS. Lo2 cou, 192, 193, 194, 195, 196, 197, 216, 227, 236, 241, 243, 245, 256, 261, 263, 273, 285, 286, 287, 288, 290, 291, 292, 298, 315, 317, 354, Pl. 3, Pl. 6, Pl. 8, Pl. 17, 392, 393, 396, 397, 398, 401, 403, Tab. 7 (opp. p- 406), 406, 407, 408, 409, 415, 418, 419, 421, 430-437, 438, 442, 443 rattus, Epimys 287, 298 rattus, Rattus 245, 287, 298, 397, 407, 421, 431, 436, 443 rattus diardu, Rattus. c = £93 rattus mindanensis, Rattus 287, 288, 290, 298 rattus negrinus, Rattus . 287, 298 Ratufa 136, 148, 153, 170, 171, 178, 181, 260, 298, 397, oe 404, 426, 427 rectodigitus, Stivalius 285, 294 rectungis, Centropus : + 054 refulgens, Rhinolophus 119, 136, Ta 153, 171 regina-montium, Palaquium . 5 ' 13 reinwardtii, Hypnodendron . : AZO) reinwardtii, Hypsiboas_ . é , 5 75 reinwardtii, Rhacophorus, 5 73-74, 75, 95 reinwardtii lateralis, Rhacophorus . 75 Reithrodontomys . 5 m 2 340 religiosa, Gracula . 5 - 154 remifer, Dicrurus ry 133, 142, 200 reptiles : 43-101 reticulata, Dillenia. : a : : 12 reticulatus, Python : 9 . 7 LOT rettigi, Ctenophthalmus . é 5 Pl. 4 Rhabdophis . 4 ' F 9 “47 Rhacophoridae 0 5 6 : 79-75 Rhacophorus Rhadinopsylla : 59, 72-75, 95 324, 333. 344, 352, 354, 355, 368, Pl. 11, 410, 411, 412, 425 thaeba, Medwayella 217, 238, 257-258, 265, 266, 267, 305, Pl. 45, Tab. 5 (opp. p. 402), 416 rhaebus, Medwayella_ . 2 205 rhaebus, Stivalius . : 215, 257 Rheinardia . 5 =) LOO) Rheithrosciurus 396, Aoy, 26h 404, 4206 rhinoceros, Buceros : III, 139 Rhinolophidae 4 2) DEG Rhinolophus 23- 28, 40, Tg! 736, 147, 153, 170, 171, 172, 183, 190, 192, 195 Rhinomyias . 117, 134, 145, 154 Rhinoplax . = he pee cls) Rhinosciurus 121, 136, 146, 149; 153, 170, 171, 174, 193, 196, 215, 216, 226, 227, 241, 242, 243, 244, 259, 261, 262, 263, 264, 265, 298, 319, 397, 398, 401, Tab. 6 (opp. p. 404), 405, 415 Rhipidura 116, 117, pn 144, 154, 200 Rhizogoniaceae - 5 $ 19 Rhizogonium : 5 3 3 - 19 Rhizothera . 5 “ 6 109, 138, 154 Rhododendron c 2 : 14 rhodomelas, Macropisthodon : E 84, 96 thodomelas, Tropidonotus s 5 a Ot Rhombomys 2 : 5 + 350 Rhynchopsyllus”. . ¢ Paes | Rhyticeros 5 0 é III, 139, 154 Riccardia.. ‘ ° 4 : 5 1074 Riccardiaceae 0 S © 17 richardsoni, Acanthopsylla ‘ i OZ Ricinidae " a ; . 2 ies) Ricinus : 5 f 5 - 199 Rickettsia 167, 172, 178, 184, 193 ridleyi, Pipistrellus 33-34, 40, 120, 148 riversi, Neoschoengastia : 190, 192, 195 robertsianus, Listronius . : 0 sees O2 robinsoni, Ceratophyllus 5 c = 4230 robinsoni, Coelops . . 29, 30, 40 robinsoni, Goniocephalus 5 5 146 robinsoni, Medwayella 215, 217, 218, 227, 228, 229, 230-233, 234, 235, 238, 244, 252, 256, 258, 259, 260, 261, 262, 263, 264, 265, 260, 314, 315, Pl. 1, Tab. 5 (opp. p. 402), Tab. 6 (opp. P- 404), 405, Tab. 7 (opp. p. 406), 416, 438, 449, 442, 443 robinsoni, Medwayella robinsoni 233-234, 235, 237, 266, 269, 302, 303, Pls. 5-7, Pls. 10—- 11, Pls. 16-18, Pl. 21, robinsoni, Pygiopsylla 230, 232 robinsoni, Rhinolophus . : 24-27 robinsoni, Rhinolophus robinsoni —. 25-27) robinsoni, Stivalius 4 a robinsoni bogora, Medwayella 235-236, 266, 269, 303, Pls. 19-20, Pl. 22 robinsoni-group, Stivalius 204, 205, 207, 208, 209, 215 robinsoni klossi, Rhinolophus 25-27, 40 204, 215, 230 INDEX 467 robinsoni peregrinata, Medwayella 234-235, 236, 266, 269, 303, Pls. 19-21 robinsoni robinsoni, Medwayella 233-234, 235, 266, 269, 302, 303, Pl. 1, Pls. 5-7, Pls. 10— 11, Pls. 16-18, Pl. 21 robinsoni robinsoni, Rhinolophus . 25-27 robinsoni tiomanica, Medwayella 236-237, 266, 269, 303, Pls. 19-20, Pl. 22 robusta, Arachnothera 117, 145, 154 robustus, Opisodasys Is Jel 9) Rollulus 108, 138, 154, 181 roluis, Ascoschoengastia 193, 195 Romerolagus. : ° , : 358 Rooseveltiella : 3 : 0 - 346 roseus, Pericrocotus a 4 : aay rostratum, Trichastoma . 154 Rothschildiana 324, 332, ast, Tab. 7 (opp. P. 406), 408, 409, 416, 417, 418, 432, 436 roulroul, Rollulus . 108, 138, 154, 181 rubeculoides, Muscicapa 116, 124, 144 rubella, Murina huttoni . 4 : o ays rubiginosa, Santiria : 5 9 Q 13 rubiginosus, Blythipicus 112, 132, 140 Tudicollis, Varanus. : : eye rufescens, Otus 110, 132, 139, 171 rufescens, Prinia . F 6 6 By! ruficapillus, Enicurus 116, 134, 135, 144, 154 ruficeps, Macropygia 109, ep 132, 135, 138, 154 ruficeps, Orthotomus. 154 tufidorsus, Ceyx . d IIl, 132, 139 rufigenis, Dremomys es 216, 259, 298, 397 rufus, Cylindrophis 47 rustica, Gahrliepia 3 é . 195, 196 rustica, Hirundo . é i : 5 aes} rustica, Walchia_ . 5 5 195, 196 Tusticus, Ceratophyllus . & é 7320) rutila, Gahrliepia . : 5 F 194, 196 sabanus, Hipposideros 153 sabanus, Leopoldamys Wen. Bas 307, Ah 415 sabanus, Rattus 122, 136, 146, 149, 153, 170, 171, 172, 174, 178, 191, 192, 193, 194, 195, 196, 236, 243, 261, ES: lay 397, 398, 415 Salientia : : 49-75 salvator, Stellio . : : = 80) salvator, Varanus . : 96. 171, 181 salvator, Varanus salvator : 80 salvator salvator, Varanus i : , 80 sandoshami, Chiroptella . 192, 195 sanguinea, Amphiesma . 4 c | 47 sanguinea, Natrix . : : 5 nay, sanguiniceps, Haematortyx . : e273 Santiria = c é 2 7 : 13 Sapium : 5 9 12 saproxylopilum, Sematophyllum o 19 sarawacensis, Amphiesma 47, 84, 96 sarawacensis, Tropidonotus . cod! sarawakensis, Natrix ‘ : F a YP Sasia . : ; “ 112, 132, 140 saturata, Aethopyga 117, 145 Sauria 5 : 76-83 sauteri, Paraceras Tab. 5 ‘opp: P- 402), 415 schefferi, Hystrichopsylla : 8343 schelkovnikovi, Wagnerina. : Plh x3 schiffnerianus, Syrrhopodon . : Otis} Schistochila . : 6 : . 18 Schoengastia : é 5 5 TroO Schoengastiella . 3 3 : - 194 schreibersii, Miniopterus . . 4 caf) schreibersii blepotis, Miniopterus 39, 40 Scincella 5 4 : : : 82, 96 Scincidae 5 : ° 3 ¢ 82-83 Scincus ps ; ; SG ‘ sees2 Sciuridae : : 5 : ena sciurorum, Monopsyllus . 350) 355, Pl.g Sciurus. a 5 228, 257, 316, 441 Scolopsyllus . af : - 340 scopulorum, C eratophyllus : Q 346) scrofa, Sus 123, 153, 171, 174, 175, 177, 179, 180, 181, 183 scutulata, Ninox . “ 5 E54 Seicercus : F 116, 134, 144 Sematophyllaceae . : ; : 5) Sematophyllum_. ° 19 semermis, Haemaphysalis 167, 168, 170, 171, 174, 175-176, 182, 183, 184 sepiarium, Trichastoma 115, 133, 142, 154 sericeus, Orthotomus 116, 144 Serpentes . E e 83-97 setosa, Neopsylla 2 322, Pl. 6 setosa-group, Neopsylla ‘j - 409 sexdentatus, Orchopeas 314, 320, 321, Pl. 5 sexdentatus agilis, Orchopeas . B2T pb les Shorea . ; ; a 13 siamensis, Rhinolophus macrotis 0 27, 28 Siaphos c 5 C 46 sibirica, Muscicapa 116, 124, 144 Sibynophis 46 347. 350, PL 8, “406, Tab. 7 (opp. Sigmactenus P- 406), 408, 431, 432, 436, 443, 444 signata, Nectophryne . : a Gy signata, Polypedetes a : a 268 signata, Rana 4 : 65, 68, 69 signata, Rana signata 68-69, 95 signata signata, Rana 68-69, 95 siligorensis, Myotis . 30-31, 40 siligorensis, Myotis siligorensis. c 5 63 siligorensis alticraniatus, Myotis 6 aie) siligorensis siligorensis, Myotis : : 30 siligorensis sowerbyi, Myotis . ° coe 50) siligorensis thainus, Myotis. - ) 30 117, 134, 145 simplex, Anthreptes 3 167, fo 172-173, 183 simplex, Ixodes simplex, Pycnonotus a 5 113, 133, 141 simplex-group, Rhinolophus_ . : aes simplex-subgroup, Rhinolophus 5 Sh simus, Glyphotes 396, 397, 398 sinensis, Centropus 109, 177, 182, 199 sinensis eurycercus, Centropus A yes singalensis, Anthreptes 117, 145, 154 468 INDEX sinicus, Coelops’ . 6 0 o 5 29 sinoculus, Pulex . a C : 9346 Siphonaptera 201-450, 78 Pls. Siseca . ‘ : 5 : 190, 195, 196 Sitta . z 5 5 : je ngage Sittidae < - : c =) enig Smitella 340, 350 smiti, Rothschildiana Tab. 7 ‘opp. P. 406), 416 snodgrassi, Cuculiphilus . 5 - 199 soberoni, Pleochaetis societatis, Pipistrellus sodalis, Macrostylophora : 325) 322, Pliz2 34-37, 40, 120, 148 Tab. 5 (opp. p. 402) solaris, Pericrocotus 113, 133, 141 solitaris, Muscicapa 117, 144 solitus, Neoschoengastia 190, 192, 195 sondaica, Neopsylla Tab. 7 (opp. p. 406), 409, 416 sonnerati, Chloropsis I13, 141, 154 sordida, Pitta 112, me 154, 190, I9I, 192, 195 Sorex . 0 5 - 441 Soricidae 5 é : eLLO sowerbyi, Myotis siligorensis : : She spadiceus, Hylopetes I2I, 149, 171 spalacis, Ctenophthalmus Q S47. spalacis, Spalacoctenophthalmus_ . a S347. Spalacoctenophthalmus . a ; 0347 Spalax 4 347, Pl. 4 sparverioides, Gueulus 109, 138 speciosus, Apodemus 272, 298 spelaea, Eonycteris 153, 171 sperata, Nectarinia ; - 5 - 4154 Sphagnum . 4 : : 2 14, 17 Sphenomorphus~ . : : : 83, 96 spiculosus, Syrrhopodon : 5 co 18 spilocephalus, Otus 110, 132, 139 Spilopsyllus . 337, 364 Spilornis “| 108, 138 spiniforme, Rhizogonium 5 a - 19 spiramus, Stivalius 286, 287 spiramus, Stivalius cognatus 205, 285, 287, 294, 305, Pl. ~ Pls. ee? Pl. 8 Spizaetus. 4 wESS splendidula, Tupaia 2 + 400 squamatus, Pycnonotus 113, 133, I4I, 154 Stachyris 115, 133-134, 135, 143, 154, 199 stanfordi, Conorhinopsylla 340, 345, Pl. 12 stanfordi, Epitedia 5 p Pl. 19 stejnegeri, Pesyyeyve = = 11030057423 Stellio ree tet?) Stenischia 324, 325, “332, 333, 336, 337, 432 Stenistomera. 324, 331, 333, 335, 355 Stenomys 261, 298, 396, 401, Tab. 7 (opp. P- 496), 409, 415, 433 Stenoponia 5 351, 358, 359 stevensi-group, Neopsylia : : - 409 stheno, Rhinolophus 119, 136, 147, 153, 170, 172, 173, 190, 192, 195 Stictolejeunea ‘i ES. Stivalius 204, 205, 207, 208, 209, 210, 212, 215, 230, 232, 233, 255, 256, 257, 272, 281-294, 305, Pls. 52-58, 314, 315, 367, Pl. 3, Pl. 8, 401, 402, 403, 406, Tab. 7 (opp. p. 406), 407, 416, 417, 418, 419, 420, 421, 430, 431, 436, 439, 440, 443, 444 stormi, Ciconia é 4 es3 Strepsylla 0 324, 330, 333, Pl. 13 striata, Catallagia ; : Pivrg striata, Coracina . ; a : - 154 striata, Kenopia 115, 143 Strigidae 109-110 strigula, Minla 115, 134, 143 Sturnidoecus a : i . 200 stylosus, Dolichopsyllus f ; : - 343 styx, Ceratophyllus 317, 327, 344, 348, 365, Pl. 16 suavis, Actenopsylla 328, 342, 348, Pl. 19 sucking lice ; é 199-200 Suidae . - : 4 ¢ 3 A) eG) suilla, Murina 4 38, 153 suillus, Hylomys 125, 273, "401, Tab. 7 (opp. P- 406), 409, 415, 438 sultanea, Melanochlora 114, 133, 142 sumatraensis, Capricornis : eek ir chra tat sumatrana, Corydon - 9 2 . 154 sumatranus, Phoenicophaeus . : - 154 sumatranus, Trimeresurus 2 93 Suncus 118, 456: 437 Sundasciurus 121, a6; ee 148, 149, 153, 170, 171, 174, 192, 193, 196, 216, 227, 233, 236, 237, 241, 244, 252, 254, 256, 257, 259, 260, 261, 262, 265, 298, 396, 397, 398, 401, Tab. 6 (opp. P- 404), 415 surifer, Lenothrix . 298, 397, 398, 401 surifer, Rattus 122, 146, 149, 153, 155-163, Pl. 1, 174, 236, 298, 397, 398, 401, Tab. 7 (opp. p. 406), 415 surifer, Rattus surifer 157, 158, 161 surifer surifer, Rattus 157, 158, 161 Surniculus 109, 132, 138 Sus 123, 150, 153, 171, 174, 175, 177, 179, 180, 181, 183 sutorius, Orthotomus . ; ; Ay swinhoei, Tamiops 404, 429 Swintonia. : 5 13 sylvestre, Leptotrombidium O F 191, 196 Sylviidae 5 : : 0 . “126 Symingtonia i 5 13 syndactylus, Hylobates 120, 148, 153 Syngenopsyllus 326, 333, 338, 340 341, 347, Pl. 1, Tab. 5 (opp. p. 402), 404, Tab. 6 (opp. P- 404), 411, 412, 416, maker seh 424, 428 Syrrhopodon 18 Syrrhopodontaceae 9 : - Z 18 Tachyglossus : : ; * 320 tahan, Sundasciurus eons 121, 171 tahitica, Hirundo . b : ; eens) talpae, Hystrichopsylla . - 5 34 Tamias. : : - 3506 tamias, Monopsyllus - a n - 355 Tamiasciurus : : . : Pl. 10 INDEX 469 Tamiophila 324, 330, 333, 355, 356, Pl. 12 Tamiops 121, 136, 149, 216, 241, 259, 260, 298, 397, 399, 404, Tab. 6 (opp. p. 404), 426, 429, 442 Tana . : 3 - 439 tana, Medwayella phangi 242, 250, 267, 268, 303, Pl. 23, Pl. 25 tana, Tupaia 232, 242, 254, cae aa 400, 415 tangalunga, Viverra : seats Tapiridae . 5 : : - e273 Tapirus 123, 150 Tarsopsylla 326, 333, 334, 338, 339, 341, Pl. 1, 410, 411, 412, 414, 425, 441 telegoni, Delotelis . BY ah ef by pir) temmincki, Maxomys fulvescens ° - 2098 temmincki, Rattus bukit 6 a . 256 temmincki, Rattus fulvescens . 256, 298 temminckii, Eurostopodus , eer! tenuis, Sundasciurus 121, 136, 146, 148, 153, 170, 171, 174, 216, 227, 236, 237, 241, 244, 259, 260, 262, 265, 298, 396, 397, 398, Tab. 6 (opp. Pp. 404), 415 tenuis tahan, Sundasciurus 121, 171 tephromelas, Aeromys . F 12%, 149, 171 Terpsiphone 117, 134, 145, 154, 200 tesquorum, Citellophilus 322, 349 testudinarium, Amblyomma 167, 168, 171, 181-182, 183, 184 Testudo “ a) 1S 5rS2 thainus, Myotis siligorensis 4 é 30 thambus, ar aaa 315, 362, Pl. 9g Thamnium . 3 5 9 & 19 Thaumapsylla a . x 5 333 Thomasomys D ° a 5 - 414 thompsoni, Audyana 190, 196 Thrassis 349, 356 Thuidiaceae F 6 i : : 19 Thuidium . é a : 19 thurmani, Medwayella 244-247, 267, 260, 304, Pls. 29-34, Tab. 5 (opp. p. 402), 404° Tab. 6 (opp. p. 404), Tab. 7 (opp. Pp. 406) Thyridium . 18 thysanota, Genoneopsylla 324, Pl. II tickelli, Hesperoptenus . 9 0 é 38 tickelli, Trichastoma 5 3 r Eeb25 ticks . : : 7 5 165-186 tigrinus, Lanius b 5 9 c - 154 tigris, Panthera 123, 150, 181 Timaliidae : II4-116 tinctus, Ricinus. 5 é . 199 ?tiomanica, Ansonia 49-52, 62, 95 tiomanica, Medwayella robinsoni 236-237, 266, 269, 303, Pls. 19—20, Pl. 22 tiomanicus, Rattus 122, 136, 146, 149, 195, 298, 397, 398 tiomanicus, Rattus tiomanicus 5 - 236 tiomanicus jalorensis, Rattus 171, 197, 216, 298 tiomanicus tiomanicus, Rattus : 230 tiptoni, Pygiopsylla Tab. 7 (opp. p. 406), 407, 416, 420, 421 todayensis, Rattus exulans . 5 Serzco tolli, Monopsyllus . 5 5 Pl. 10, 410 tomentella, Trichocolea . : : 18 tomesi, Hesperoptenus . 5 38, 40 Toritrombicula FG IQI, 192, 195 toxopei, Murina. é 30 toxopeusi, Sigmactenus . - 408, 444 traguli, apa 167, 171, 176, 183, 184 Tragulidae . Fi prs Tragulus 123, 136, 150, 53) 171, 174, 175, 176, 177, 178, 181, 183, 196 Traub oR. _ 201-450, 78 Pls. Traubia : e : 5 B&G} Treron . 5 109, 132, 138, 153 treubii, Syrrhopodon : ets trianguligera, Natrix 83-84, 96 trianguligerus, Tropidonotus . : 83 tricata, Neopsylla . Tab. 7 (opp. p. 406) Trichastoma 115, 125, 133, 142, 154, 177 Trichocolea . 7 : 18 Trichomanes ; - 20 Trichopsylloides tam 333, aaah 344, 368 Trichosteleum 0 5 19 trifoliatus, Rhinolophus . trigonostigma, Dicaeum . 120, 136, 147, 153 117, 145, 154 Trimeresurus 47, 93-94, 90-97 Trismegista . i 19 trispinosa, Macrostylophora Tab. 5 (opp. p. 402) trispinus, Xenopsylla_. ¢ = sd2 Tristania 0 3 - 2 13 tristis, Choristopsylla . 9 325, 334, 341 tristis, Eugeissona . A é . ‘ 13 tristis, Meiglyptes trivirgata, Arctogalidia 122, 136, 150, 170, 177, 181 I12, 140 trivirgatus, Phylloscopus a 116, 134, 144 Trogonidae : : - IIO-III Trombiculidae 6 5 - 187-198 Tropidonotus : : a 83-84 tsutsugamushi, Rickettsia b R . 193 tukki, Meiglyptes .- 112, 132, 140, 154 Tunga . : : 292, 333, 337, 354, 362 Tupaia 120, 146, 148, 153, 171, 181, 192, 195, 207, 216, 227, 232, 233, 236, 237, 241, 242, 243, 244, 245, 247, 252, 254, 255, 250, 259, 260, 261, 262, 263, 264, 265, 272, 280, 298, Pl. 3, 399, 400, 406, Tab. 7 (opp. p. 406), 411, 415, 420, so 439, 440 Tupaiidae . c c : 120 Turdidae 6 4 : 5 4 : D6 Turdus : : 4 6 et70 turmalis, Ganniepes 3 , P 195, 196 turmalis, Walchia 7 5 . 195, 196 tweediei, Eltonella 5 3 - 190, 195 tweediei, Macrocalamus . A - 47, 90-92 tweediei, Rana : 6 c . 69-70, 95 Typhlopidae . - z ‘ 5 ees Typhlops 47, 83, 96 Tytonidae . a a a SELILO) 470 INDEX umbratilis, Rhinomyias - uncinalis, Macrostylophora 117, 134, 145 Tab. 5 (opp. p- 402) undulatum, Thyridium : : A 18 undulatus, Rhyticeros III, 139, 154 unguiculatus, Rallicola . F 4 199 unicolor, Cervus ; 153, 173 unicolor, Muscicapa 116, ie 154 unisulcatus, Otomys . : 329 uphami, Toritrombicula . : bs a key] Urogale ; : 399, 400, 420 Ursidae 0 c ‘ é neaz2 vagans, Cuculus. . , . - 153 validus, ela aan < DE2, 133; 140/070 Varanidae . Z 79-80 Varanus Toes) 96, 171, 181, 182 variegatus, Cynocephalus 119, 136, 147 variolosus, Cacomantis . ; é aeeet54) velatum, Philentoma 117, 134, 145, I9I, 195 vermiculatus, Ixalus : 2 71 vermiculatus, Philautus . 46, 47, 71, 95 Vermipsylla . : z . 360 vertebralis, Amblycephalus : OZ vertebralis, Pareas 0463 92, 96 veruta, Medwayella 252-255, 267, 268, 304, Pls. 38-40, Tab. 5 (opp. p. 402), Tab. 6 (opp. p- 404), 406, Tab. 7 (opp. p. 406), 416 vesperalis, Opisodasys 317, 327, 333, 339, 341, 344, Pl. 15, 412 Vespertilio . 0 ees, Vespertilionidae . 120 vexabilis, Xenopsylla 317, 318, 328, 344, Pl 17, Tab. 7 (opp. p. 406), 407, 433, 434, 436 vexabilis-group, Xenopsylla_. 9 + 443 vidua, Haemaphysalis 167, 170, 176, 183 vigil, Rhinoplax . ¢ S =) tango viridescens, Hypsipetes 114, 133, 141, 199, 200 viridis, Calyptomena II2 133 135, 140, 199 viridis, Merops . z 3 III, 135, 139 vison, Monopsyllus 315, 339, Pl. 1, Pl. 10 vittigerum, Lygosoma_ . é 82 vittigerum, Lygosoma vittigerum a oy vittigerum, Scincella vittigerum 5 82, 96 vittigerum vittigerum, Lygosoma_ . At} vittigerum vittigerum, Scincella - 82, 96 Viverra : : : zi é = Ler Viverridae . 5 0 a 6 - 122 volans, Draco 9 5 = 76.78.70 volans, Draco volans : . : 79, 96 volans volans, Draco 5 . 79, 96 vomerus, Lentistivalius 272-281, 305, Pls. 46-49, Pls. 51-52, 312, Pl. 3, Tab. 6 (opp. P- 404), 406, Tab. 7 (opp. p. 406), 416, 420 wagneri, Monopsyllus 321, 362, Pl.g Wagnerina +324) Pls Walchia 195, 196 Walchiella ee 195, 196 Weinmannia : : f 13 wellingtoni, Haemaphysalis 167, 171, 176, 183, 184 Wenzella 324, 333, 345, 352, 354, 355, 368, Pl. 14 werneri, Sigmactenus Tab. 7 (opp. p. 406), 408 westermanni, Muscicapa 117, 134, 144 whiteheadi, Exilisciurus . 2 5 SGT whiteheadi, Lenothrix 397, 398, 401, Tab. 7 (opp. p. 406), 415 whiteheadi, Rattus 118, 122, 136, 146, 149, 153, 171, 195, 196, 273, 397, 398, 401, Tab. 7 (Opp. p. 406), 415 Whitmore, T. C. 7 9-15, 8 Pls. wichmanni, Eutrombicula a ‘ : 190 williamsoni, Collorhabdion ‘ E ree i wrayi, Rhododendron. “ 5 14 xanthonotus, Oriolus 114, 135, 142 xanthorhynchus, Chrysococcyx tog, 138 xanthurus, Bullimus : A se pes xanthurus, Rattus 407, 421 Xenodaeria . d : c > te G2 Xenopeltis . z 88 Xenopsylla 317, 318, 323, 328, 333, 335, 337, 342, 344, 347, 35° 357, 358, 361, 362, Pl. 17, Tab. 7 (opp. p- 406), 407, 408, 433, 434, 436, 443 Xerus . j - ri - 410, 4II Xiphiopsylla | : - o 2 382 Yong, H. S. 155-163, 1 Pl. Yuhina I15, 143 zantholeuca, Yuhina 115, 143 zebratum, Sphenomorphus indicus - « BOR zeylanicus, Pycnonotus . “ a - 154 Zosteropidae . : 5 é Ee. Zosterops 117, 145 Ash Mis < = WAT. HIST 2 2 MAY 1975 wa PRESENTED o/ Neer 0?