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Bulletin of the ( ^
V> tte*A*y
British Museum (Natural Histor^
Entomology series Vol 43 1981
British Museum (Natural History) London 1982
Dates of publication of the parts
No 1 30 July 1981
No 2 24 September 1981
No 3 29 October 1981
No 4 26 November 1981
ISSN 0524-6431
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset
Contents Entomology Volume 43
Page
No 1 A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea) and its allies. N. D. Jago 1
No 2 The Asian, Australasian and Pacific Paraboloponinae (Homoptera : Cicadellidae). A taxonomic revision with a key to all the known genera of the subfamily. M. D. Webb 39
No 3 A revision of Phyciodes Hiibner and related genera, with a review of the classification of the Melitaeinae (Lepidoptera : Nymphalidae). L. G. Higgins 77
No 4 A revision of six minor genera of Myrmicinae (Hymenoptera : Formi- cidae) in the Ethiopian zoogeographical region. Barry Bolton 245
GENER
-4AUG
Bulletin of the
British Museum (Natural History
A revision of the genus Usambilla Sjoste( (Orthoptera: Acridoidea) and its allies
N. D. Jago
Entomology series
Vol 43 No 1 30 July 19:
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series.
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1981
ISSN 0524-643 1 Entomology series
Vol 43 No 1 pp 1-38 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 30 July 1981
A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea) and its allies
N. D. Jago
<f^f
Centre for Overseas Pest Research, College House, Wrights Lane, London W8 5SJ
Contents
Synopsis
Introduction ....
Material
Key to genera in the Lentulidae . Chromousambilla gen. n.
Key to species ....
Males
Females .... Microusambilla gen. n. . Altiusambilla gen. n. Rhainopomma gen. n. .
Key to species . . . .
Males
Females . . . . U sambilla Sjostedt
Key to species and subspecies .
Males
Females ....
References
Index .
1
1
2
3
5
6
6
6
10
12
13
14
14
15
20
22
22
24
37
38
GENER
4AUG
LIBRAI
Synopsis
The genus Usambilla Sjostedt is redefined and fully revised with keys to the species and subspecies. Four new genera, ten new species and three new subspecies are described. Altiusambilla modicicrus (Karsch) is implicated in the defoliation of exotic Pinus plantations in northern Tanzania.
Introduction
The genus Usambilla has been assumed to consist of a homogeneous group of small lentulid grasshoppers. The male genitalia of the group reveal, however, that five genera are involved within the old definition of Usambilla. Chromousambilla has male penis valves of great length (Fig. 12) which enable the diminutive male to remain firmly attached to the female during copulation without standing upon her in the usual acridid manner. The penis valves are curved and fit into the female spermathecal duct, which is of similar shape and length (Fig. 23). Usam- billa sensu stricto has tapered valvulae (Fig. 150) with barbs on each side short of the apex, while Rhainopomma (Fig. 72) has these barbs situated apically. Microusambilla clearly has affinity with Usambilla, while Altiusambilla has affinity with Rhainopomma and Mecostibus.
Reports have recently been published (Whellan, 1975; 1976) of eumasticids and lentulids causing defoliation of exotic Pinus, particularly P. patula. In Malawi defoliation has been caused by Plagiotriptus Karsch (Eumastacidae) and members of the lentulid genera Mecostibus, Nyass- acris and Malawia. In Tanzania attacks so far reported are from W. Kilimanjaro in December, 1975 at Msituni, the species involved being members of the genus Chromothericles Descamps (Eumastacidae) and the lentulid Altiusambilla modicicrus. This last is morphologically much
Bull. Br. Mus. not. Hist. (Ent.) 43(1): 1-38
Issued 30 July 1981
N. D. JAGO
closer to Mecostibus than to the rest of Usambilla sensu stricto. Indeed the close similarity between the genera described in this paper may be due to convergent evolution. The external similarity between Altiusambilla modicicrus and Rhainopomma montanum is particularly striking.
Material
Most of the material used for this study is in the collections of the first two institutions listed below. Museums loaning specimens have their abbreviations listed in the text as follows.
BMNH
COPR, London NR, Stockholm MNHU, Berlin MR AC, Tervuren
British Museum (Natural History), London
Centre for Overseas Pest Research, London
Naturhistoriska Riksmuseet, Stockholm
Museum fur Naturkunde der Humboldt-Universitat, Berlin
Musee Royal de 1'Afrique Centrale, Tervuren
Kenya series were immensely improved by the addition of COPR material collected recently by Dr I. A. D. Robertson and Mrs A. Robertson.
Figs 1-9 Chromousambilla species. 1 , lateral aspect of pronotum of male C. veseyi. 2, dorsal aspect of head of male C. latestriata. 3, apex of male abdomen of C. burtti, lateral aspect. 4, entire male of C. latestriata. 5-9, dorsal aspect of male abdominal tip in (5) C. burtti; (6) C. mweruensis; (7) C. robertsoni; (8) C. latestriata; (9) C. veseyi. Scale lines represent 1 mm, that below Fig. 1 applies to Figs 1-4, that above Fig. 9 applies to Figs 5-9.
REVISION OF USAMBILLA 3
Key to genera in the Lentulidae (modified after Dirsh, 1965)
1 External apical spines of hind tibia absent. Head prognathous, or face vertical, or moderately
sloping backwards 2
External apical spine of hind tibia present. Head conical or acutely conical, with face strongly sloping backwards or vertical 16
2 Coxa of middle leg with large conical process, sometimes with a small tubercle
. MECOSTIBUS Karsch Coxa of middle leg without process, sometimes with a small tubercle ...... 3
3 Ninth abdominal tergite of male with a median dorsal bifurcate appendage .... 4 Bifurcate appendage on mid-dorsal margin on ninth abdominal tergite absent .... 5
4 Frons vertical or sloping forwards, slightly excurved or straight; fastigium of vertex not protrud-
ing. Body small but comparatively robust SHELFORDITES Karny
Frons sloping backwards, slightly incurved, fastigium of vertex and upper part of frons strongly protruding forwards. Body elongate, slender KALAHARICUS Brown
5 Dorsum of pronotum crossed by four deep transverse sulci . . . MECOSTIBOIDES Dirsh Dorsum of pronotum crossed by three or fewer transverse sulci, or sulci not crossing disc at all . 6
6 Frontal ridge deeply sulcate below level of antennal sockets 7
Frontal ridge weakly sulcate to flat below level of an tennal sockets ...... 8
7 Fastigium of vertex horizontal, triangular, seen from above, strongly projecting in front of eyes
. . SYGRUS I. Bolivar
Fastigium of vertex sloping obliquely forwards; as seen from above (Fig. 27) weakly projecting in front of eyes . MICROUSAMBILLA gen. n. (p. 10)
8 Fastigium of vertex moderately or slightly projecting in front of compound eyes, as seen from
above 9
Fastigium of vertex not projecting in front of eyes NYASSACRIS Ramme
9 Hind femur not inflated bilaterally but strongly bilaterally compressed 10
Hind femur often stocky and inflated, not bilaterally compressed 11
10 Interocular distance in males about as wide, in females twice as wide, as antennal scape.
Fastigium of vertex in males, from above, seen to be excised at apex . PARALENTULA Rehn Interocular distance in males about twice, in females about four times, as wide as antennal scape. Fastigium of vertex, as seen from above, widely rounded or truncate . . . LENTULA Stal
1 1 Body moderately to very smooth. Frontal ridge moderately widened in upper part with weak
carinulae and shallow depression. Male supra-anal plate with black tubercles (Figs 35, 50-54,
94-100) 13
Body strongly rugose. Frontal ridge strongly widened in upper half, strongly carinulate. Supra- anal plate lacking tubercles on disc 12
12 Hind femora stocky, length to depth ratio R about 3.2. Compound eyes strongly inflated, width
across eyes to pronotal width ratio 3 : 2. Length of pronotum at dorsal midline shorter than its width. Supra-anal plate and ninth abdominal tergite of male lacking any callosities
KARRUACRIS Dirsh
Hind femur slender, ratio R about 4.0. Compound eyes only moderately inflated, width across eyes rather less than maximum pronotal width as seen from above. Pronotal length at midline equal to its greatest width. Supra-anal plate and ninth abdominal tergite with small marginal callosities MALAWIA Dirsh
13 Hind femora of males yellow or greenish yellow with genicular part black. Eyes protruding, red
in males. Penis valves elongate, flagelliform (Fig. 15). Female ventral ovipositor valves shovel- shaped (Fig. 24) CHROMOUSAMBILLA gen. n. (p. 5)
Hind femora green, brown or yellowish brown with genicular part dark brown, light brown, blue or some other colour other than black. Eyes in males never red. Penis valves barbed (Figs 38, 72, 133). Female ventral ovipositor valves pointed, tapered (Figs 147, 148) .... 14
14 Margin of abdominal tergite 9 in males bearing one or two pairs of black tubercles (Fig. 35).
Penis valves apically attenuate with barbs apical. Anterior penis valves short, auricular (Fig.
37). Posterior part of median pronotal carina raised into a large tubercle (Fig. 36) in females.
ALTWSAMBILLA gen. n. (p. 12)
Rear margin of tergite 9 of abdomen without conical black tubercles. Penis valves tapering,
conical in dorso-posterior view (Figs 72, 133). Anterior penis valves flat, vertically orientated
and racket-shaped (as in Fig. 12) 15
N. D. JAGO
15
16
17
18
19
20
21
Male supra-anal plate with eight to twelve strong black tubercles on the disc (Figs 50, 54). Male
penis valves with apical barbs (Figs 64, 67, 68, 72) . . . RHAINOPOMMA gen. n. (p. 13) Male supra-anal plate with eight to ten strong black tubercles on its disc (rarely as in Figs 97, 99, with pair of weak ones making 12 in all). Male penis valves with pre-apical lateral barbs (Figs 133,135,138,141,142,145,150) . . . USAMBILLA Sjostedt (p. 20)
Fastigium of vertex not projecting in front of eyes, sloping strongly forwards and forming part of face . . ... . EREMIDIUM Karsch
Fastigium of vertex projecting or strongly projecting in front of eyes, horizontal . . . . 17
Basal angles of fastigium of vertex detached from eyes and projecting laterally; body strongly
rugose SWAZIACRIS Dirsh
Basal angles of fastigium of vertex of normal shape, touching eyes. Body smooth or only
moderately rugose 18
Frontal ridge, in profile, more or less strongly projecting in upper half or upper third . . . 19
Frontal ridge, in profile, straight, not projecting in upper part 22
Fastigium of vertex elongate. Antennae ensiform, serrated . . . . DEVYLDERIA Sjostedt
Fastigium of vertex shorter. Antennae phylliform or filiform 20
Frontal ridge in upper projecting part lamelliformly compressed, below shallowly sulcate . . 21
Frontal ridge sulcate along whole length BASUTACR1S Dirsh
Antennae very short, phylliform, compressed and strongly widened, about four times longer than wide. Fastigium of vertex with lateral carinulae. Pronotum shorter than its width. Body
rugose KARRUIA Rehn
Antennae narrow, almost filiform. Fastigium of vertex without carinulae. Pronotum longer than its width. Body smooth GYMNIDWM Karsch
14
16
Figs 10-16 Chromousambilla species, male phallic complex. 10, 14-16, lateral aspect of aedeagal valves of (10) C. robertsoni; (14) C. mweruensis; (15) C. burtti; (16) C. veseyi. 11, epiphallus of C. robertsoni. 12, lateral aspect of right side of entire complex in C. latestriata. 13, ventral aspect of cingular rami and anterior valves of penis of C. latestriata. Scale-lines represent 0.5 mm — that below Fig. 12 applies to 10 and 12-16; that below 11 to that figure. Small arrows on Figs 10 and 15 show equivalent position on each set of valvulae.
REVISION OF USAMBILLA 5
22 Fastigium of vertex comparatively short, angular. Body slender, moderately elongate . . 23 Fastigium of vertex elongate, angular or narrow parabolic, with obtusely angular apex. Body
slender and very elongate 24
23 Fastigium of vertex broadly angular, wider than longest diameter of eye. Frontal ridge sulcate
along whole length. QACHAS1A Dirsh
Fastigium of vertex narrowly angular, much narrower than longest diameter of eye. Frontal ridge in upper fourth compressed, without sulcus, sulcate below . . HEL W1GACRIS Rehn
24 Fastigium of vertex angular. Frontal ridge above base of antennae low, compressed. Antennae
thick, strongly and regularly narrowing towards apex, weakly compressed in basal half. Body smooth BETISCOWES Sjostedt
Fastigium of vertex narrow parabolic. Frontal ridge above base of antennae produced, strongly lamelliformly compressed. Antennae very wide, strongly compressed laterally, only slightly narrowed at apex. Body rugose . . . BACTERACRIS Dirsh
CHROMOUSAMBILLA gen. n.
Type-species: Adolfia latestriata Ramme.
DIFFERENTIAL DIAGNOSIS. Male. Separable from closely related genus Usambilla by great length of aedeagal valves, which lack barbs (Figs 10, 14, 15, 16). (Proximal section of female spermathecal duct (Fig. 23) correspondingly stiffened and elastically thickened for a distance corresponding to length of aedeagal valves distal to their basal attachment below greatly inflated cingular arch.) In life aedeagal valves curled over greatly inflated membranous arch of cingulum (Fig. 12), their tips touching ectophallic membrane posterior to epiphallus. Cingular rami also very unusual, small and parallel, being approximated on mid-line so that they lie between flattened anterior (internal) valvulae of penis (Figs 12, 13). Epiphallus very small and simple, lacking ancorae, function of which taken over by hook-like development of ventro-lateral corners of epiphallus (Fig. 1 1). Whole apparatus housed within enlarged pod-like subgenital plate (Figs 4, 8).
Fastigial depression gently concave, its mid-frontal edge being notched as viewed from above (Fig. 2). Inter-ocular groove well developed, short and wider capitad than caudad. Differs from that of Usambilla sensu stricto, which is in form of an elongated slot (Fig. 92) or a slightly wider parallel-sided groove. Body shape like that of Usambilla (Fig. 4).
Supra-anal plate bears callosities of type found in Usambilla, but proportions of supra-anal plate to sub-genital plate clearly differ (Figs 8, 94-101).
Female. Ventral valves of ovipositor (Fig. 24) straight-edged ; median process of subgenital plate almost level with their tips.
Coloration. General colouring very uniform in genus. Male eyes vermilion, frons blackish; three lateral light yellow or whitish stripes on side of head (Fig. 4). Upper pair of light stripes continued along thorax and abdomen. Subgenital plate yellow. Posterior femora yellow to yellow-green, knees black; posterior tibiae yellow-green, sooty at extreme tip and near knee. Dark body stripes blue to blue-black or brown. Female more variable, but in general (Figs 17, 20, 21, 26) lighter brown to olivaceous dorsally, with light banding like that of male, but with uppermost of light (dorso-lateral) bands weak and lower one on pronotum strong. Broad dark lateral band variable in intensity. Light bands yellow or creamy colour. Posterior leg colour as for male but less intense. Body colour of female C. veseyi exceptional in that body and head olive green dorsally and uppermost dorso-lateral light bands bright yellow. Lateral dark bands very black so that female of this species resembles male more closely than do those of rest of genus.
There is great instability in the callosities of the male supra-anal plate and too much variation for this feature to be reliable diagnostically.
DISCUSSION. The genus is known only from NW. Zambia and Tanzania. C. mweruensis in the Mweru Wa Ntipa and Malagarasi R. drainages, which were once part of the Congo R. drainage basin before the formation of Lake Tanganyika by rifting, may be indicative of the great antiquity of the genus. Each species seems confined to the wettest eco-zones of a separate drainage basin. Some of these, as in the Dodoma area or Mpwapwa basin, are closed internal drainages. Others like the Ruaha R. valley are open systems, but have no ecological bridges suitable for the genus by which they can enter other drainages.
6 N. D. JAGO
Key to species
Males
1 Antennae fully twice length of head and pronotum. Large species with blue/black dark lon- gitudinal stripes on body. Light longitudinal stripes white to cream. Subgenital plate larger in profile (Fig. 3) than other species in genus (Fig. 4) and narrowed at
level of cercus apices as seen from above (Fig. 5) C burtti sp. n. (p. 9)
Antennae clearly less than twice length of head and pronotum. Dark body stripes grey, blue or
Prussian blue to blue-green. Light longitudinal stripes yellow 2
2 Supra-anal plate laterad, at level of cercus apices (Figs 8, 9), black in addition to black callosities 3 Supra-anal plate without black areas in addition to black callosities (Figs 6, 7)
. . . . . . . . . C. mweruensis sp. n. (p. 9)
3 Pronotum and more particularly subgenital plate clothed with sparse hairs (Fig. 8). Black area of
supra-anal plate broadly pigmented to base 4
Pronotum and subgenital plate almost without hairs (Fig. 9). Black areas of supra-anal plate not reaching base of plate C. robertsoni sp. n. (p. 10)
4 Lower light yellow band on pronotum laterad, roughly equal in depth to dark band immediately
above it (Fig. 1) C. veseyi sp. n. (p. 8)
Lower light yellow band on pronotum laterad, half depth of dark band immediately above it (Fig. 3) C. latestriata (Ramme) (p. 6)
Females
1 Median dorsal carinula of pronotum elevated and clear to weak but entire 2
Median dorsal carinula of pronotum weak, flattened (Fig. 22).
Colouring akin to that of males with median dorsal part of body olive brown and dorso- lateral yellow band visible and entire as far back as 8th abdominal tergite (Fig. 19)
. C. veseyi sp. n. (p. 8)
2 Large species (Fig. 21). Pronotal side stripe white, bordered above and below by black. Antennae
1.50 times longer than head and pronotum together . ..... C. burtti sp. n. (p. 9)
Smaller species. Pronotal side stripe less heavily delineated or even absent. Antennae less than 1.45 times longer than head and pronotum together 3
3 Lower outer lobe of knee of hind femur uniformly creamy or light brown in colour. Ratio of
length of antennae to that of head and pronotum (R) about 1.0 or 1.4. Dorso-lateral yellow
pronotal line very weak or absent (Figs. 20 and 26) 4
Lower outer lobe of knees of hind femur with additional black markings. Ratio (R) about 1.2. Dorso-lateral yellow pronotal line narrow but usually clear (Fig. 17) .... C. latestriata (Ramme) (p. 6)
4 Brownish insects (Fig. 26); body length less than 18 mm. Ratio of antennal length to length of
head and pronotum (R) about 1.0 C. robertsoni sp. n. (p. 10)
Olivaceous or brownish insects (Fig. 20); body length more than 18 mm. Ratio (R) 1.4.
C. mweruensis sp. n. (p. 9)
Chromousambilla latestriata (Ramme) comb. n.
(Figs 2, 4, 8, 12, 13, 17, 18,24,25)
Adolfia latestriata Ramme, 1929: 307, fig. 29c. Holotype & TANZANIA: Ukimbu, Nkila, 20-2 l.viii. 1899 (Glauning) (MNHU, Berlin) [examined].
DIFFERENTIAL DIAGNOSIS. Male with supra-anal plate blackened laterally (Fig. 8), thus differing from C. burtti and C. mweruensis. Aedeagal valves longer than those of C. robertsoni or C. veseyi (see Figs 10, 12, 16); in former, pronotum and subgenital plate almost lack hairs, and black areas on supra-anal plate do not extend to its base. In contrast, C. latestriata and C. veseyi are sparsely but clearly hirsute on pronotum and subgenital plate, and black areas of supra-anal plate are as in Figs 8, 9. C. latestriata differs from C. veseyi in proportions of lateral pronotal bands (Figs 1, 3).
Female differs markedly in colouring from C. veseyi (see key) and lacks well-developed median pronotal carina of C. burtti. Lower outer lobe of knee of posterior femora with dark spots, thus differing from uniformly light brownish lobe of C. robertsoni and C. mweruensis.
REVISION OF USAMB1LLA
26
Figs 17-26 Chromousambilla species, females. 17, lateral aspect of head and thorax of C. latestriata. 18, dorsal aspect of head and pronotum of C. latestriata. 19-21, lateral aspect of head and thorax of (19) C. veseyi; (20) C. mweruensis; (21) C. burtti. 22, dorsal aspect of head and pronotum of C. veseyi. 23, dissected display of ventral ovipositor valves (vv) and spermatheca showing point distal to which duct is non-sclerotised (arrow) and apical sac plus preapical diverticulum. 24, ventral valves of ovipositor, from below, of C. latestriata. 25, dorsal aspect of head and pronotum of C. latestriata. 26, lateral aspect head and thorax C. robertsoni. Scale line under Fig. 17 represents 1 mm and applies to all except Figs 23 and 24, to which 1 mm scale line under Fig. 23 applies.
8 N. D. JAGO
MEASUREMENTS
Males Females
Head width (11) 3.4-3.7,3.52 (8) 4.1-4.4,4.26
Antenna length (9) 5.6-6.4,6.72 (7) 6.0-7.2,6.71
Posterior femur lengh (11) 7.1-8.3,7.88 (7) 9.2-10.2,9.73
Posterior femur depth (11) 2.0-2.3,2.11 (7) 2.5-2.9,2.68
Body length (11) 12.0-14.1,13.09 (8) 16.8-18.8,17.81
MATERIAL EXAMINED
Tanzania: 4 & 8 ?, Rukwa, Kapombo Hill, 25.iv.1958 (Vesey-FitzGerald) (BMNH); 1 & 1 ?, Ufipa, Ilemba gap, 12.iii.1959 (Vesey-FitzGerald) (BMNH); 11 rf, 10 ?, Ufipa plateau, Mkundi, 26 km NNW. of Sumba- wanga, 16-27.V.1966 (Jago) (5 (J, 2 9; rest COPR, London); 1 cJ, 2 $, Rukwa valley, Red Locust Camp, Musa, 26-27.ix.1964 (Jago) (BMNH).
DISCUSSION. This species was originally described from 'Ukimbu (S.O.-Tanganyika)'. At that time Tanganyika' was applied to a region embracing parts of what are now Katanga, northern Zambia and SW. Tanzania, and 'Ukimbu' was an area to the east of L. Rukwa. The COPR collections include material from the Rukwa valley and Ufipa plateau west of L. Rukwa. The females from the valley floor have dark colouring with two light yellowish body stripes laterad, the rest of the insect being dark greenish. Females from the plateau lack the upper pair of lighter bands laterad but are lighter insects overall.
A collection was made of the plant association characterising the herb cover of the habitat. This insect occurred together with Usambilla haematogramma and was particularly abundant at the base of old termite hills at the edge of a forestry plantation. The plants were kindly identified in 1967 at the East African Herbarium by S. P. Kibuwa; the species were Bidens steppia (Steetz) Scherff, Erlangea sp. near E. laxa S. Moore (Compositae); Hypoestes verticillaris R. Brown (Acanthaceae); Pycnostachys (?) stuhlmanni Guerke (Labiatae); Pseudarthria hookeri Wight & Arnott (Papilionaceae); Achyranthes aspera L. (Amaranthaceae); Physalis peruviana L. (Sola- naceae).
Like U. haematogramma, C. latestriata seems to occur in a plant community which is structur- ally part of a woodland subclimax. It does not live in the forest or grassland and would therefore seem to be an insect adapted to a transient ecosystem. The Ufipa forests seem to be at a curious senescent stage in which many of the broad-leaved trees are dying, leaving many areas with tree euphorbias as dominants. Local ecologists implicate climatic change and the effects of man in accelerating the decline in this unique forest mosaic.
Chromousambilla veseyi sp. n.
(Figs 1,9, 16, 19,22) Holotype J, Tanzania: Ruaha. Nat. Park, riverine, 15.iii.1966 (Vesey-FitzGerald) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male with supra-anal plate (Fig. 9) blackened laterally, thus differing from C. burtti and C. mweruensis. Like latter, however, in having short aedeagal valvulae (Figs 14, 16). Unlike C. robertsoni in having moderately hirsute pronotum and subgenital plate. Dark body stripes definitely bluish black, not black or dark brown. Light body stripes golden yellow, differing in proportions from C. latestriata (see Figs 1, 3; also key, p. 6).
Female differs from all other species (Fig. 19): generally darker dorsally and olive green with clear dorso-lateral yellow side stripes. Dark lateral stripes black.
MEASUREMENTS
Males Females
Holotype
Head width 3.5 3.6 4.1 4.4
Antenna length 6.9 7.6 7.6 7.9
Posterior femur length 7.8 8.1 9.7 10.3
Posterior femur depth 2.1 2.2 2.6 2.8
Body length 14.1 13.2 16.2 15.9
REVISION OF USAMBILLA 9
MATERIAL EXAMINED Paratypes. Tanzania : 1 <$, 2 $, same data as holotype ( 1 <J, 1 $ in COPR, London ; 1 $ in BMNH).
DISCUSSION. This new species is named after the late L. D. E. F. Vesey-FitzGerald, a close friend and colleague. The species is unique in having the male and female with similar bright colour patterns.
Chromousambilla burtti sp. n.
(Figs 3, 5, 15, 21) Holotype <J, Tanzania: Kikombo, Mpwapwa, 17.iv.1947 (E. Bum) (BMNH).
DIFFERENTIAL DIAGNOSIS. Both sexes distinguished at once by larger size. Antennae in both sexes longer proportionately than other species, twice length head and pronotum in male, 1.5 times in female. Supra-anal plate of male lacking areas of black pigmentation (Fig. 5), thus differing from C. latestriata, C. veseyi and C. robertsoni. Dark body stripes of male dark brown; light body stripes dull cream. Female (Fig. 21) with dark brown side stripes with other darker regions black. Dark side stripe on metathorax and first abdominal tergite strongly arched. Back of female pronotum often green. Male and female with pronounced though slender median dorsal carinula.
Male genitalia (Fig. 15) very long and aedeagus narrowed pre-apically with an oblique tip. Whole complex enclosed in large pod-shaped sub-genital plate (Fig. 3).
MEASUREMENTS
Male holotype Female allotype
Head width 3.9 5.0
Antenna length 10.5 10.3
Posterior femur length 9.6 11.9
Posterior femur depth 2.6 3.3
Body length 15.7 21.2
MATERIAL EXAMINED
Paratypes. Tanzania: 2 £, 1 £ allotype and 2 9, same data as holotype (1 & 1 $ (allotype), 1 $ BMNH; 1 & 1 ? COPR, London); 1 cJ, 1 $, Kikombo, Mpwapwa, 20.iv.1947 (E. Burn) ($, BMNH; rf COPR, London); 1 <J, 2 $, Kikombo, Mpwapwa, 18.iv.1947 (E. Bunt) (BMNH).
DISCUSSION. The species is named after the late Eric Burtt who did so much to improve our knowledge of the acridid fauna of central and eastern Africa.
Chromousambilla mweruensis sp. n.
(Figs 6, 14,20) Holotype £, Zambia: Mporokosa distr., Mweru wa Ntipa, [8-13.]vii.l952 (Uvarov) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male differs from all other species, except C. burtti, in having supra-anal plate free of black shading laterally (see Figs 6, 7). Aedeagal valves resemble C. veseyi (see Figs 14, 16), but this differs on supra-anal plate facies. Male dark body stripes blue-black, those of C. burtti being black. Latter also bigger species with longer antennae (ratio of antennal length to length of head plus pronotum 2.0 in males, 1.5 in females); antenna in C. mweruensis having equivalent ratios for male 1.8 and female 1.0.
MEASUREMENTS
Males Females Holotype
Head width 3.5 3.8 4.5 4.3
Antenna length 6.9 7.7 8.1 9.4
Posterior femur length 8.6 8.2 10.2 9.4
Posterior femur depth 2.3 2.2 2.8 3.0
Body length 16.1 14.5 19.5 21.1
10 N. D. JAGO
MATERIAL EXAMINED
Paratypes. Zambia: 1 *, same data as holotype (BMNH). Tanzania: 6 cJ, 2 ?, 35.4 km W. of Kahama, Mkwemi, 14-29.iii.1947 (£. Burn] (1 J, 1 ? COPR, London; rest BMNH); 4 J, 35.4 km W. of Kahama, Mkwemi, [xii.1946-ii.1947] (£. Burn) (BMNH); 2 & Old Shinyanga, l.ii.1947 (£. Burn) (BMNH);2 9, 16.1 km N. of Ussure, on Msigiri rd, 12.iv.1936 (£. Bum) (BMNH).
DISCUSSION. The species name originates from the Mweru wa Ntipa area of Zambia.
Chromousambilla robertsoni sp. n.
(Figs 7, 10, 11,26)
Holotype <J, Tanzania: 70.8 km N. of Dodoma, nr Meia Meia, [16-18.]vi.l967 (Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Aedeagal valvulae (Fig. 10) of intermediate length, pointed apically like C. la- testriata and C. veseyi, with black patches laterad on supra-anal plate, but these areas not reaching broadly to base of plate. Male colour and body markings like those of C. latestriata, but lower outer lobe of hind knee mostly white with limited dark spots (unlike C. latestriata in which this lobe extensively mottled with black). Pronotum and subgenital plate almost without hairs. Female almost uniform dull brown (Fig. 26) with weakly differentiated creamy diagonal side stripe. Hind femora dull olive to yellow with lower outer knee lobe pale brown, immaculate (unlike C. latestriata which has dark spots on this area). Ratio of antenna length to length of pronotum plus head about 1.8 in males and 1.4 in females; antennae being shorter in proportion than any other species.
MEASUREMENTS
Males Females
Head width (12) 3.2-3.5, 3.41 (2) 4.1 4.1
Antenna length (11) 6.2-7.7,6.78 (2) 6.6 6.3
Posterior femur length (12) 7.4-9.1, 8.03 (2) 9.8 9.4
Posterior femur depth (12) 1.9-2.4,2.16 (2) 2.9 2.7
Body length (12)11.0-13.0,12.20 (2)13.1 17.4
MATERIAL EXAMINED Paratypes. Tanzania: 10 £, 2 9, same data as holotype (9 <J, 1 9, BMNH; 1 £, 1 9, COPR, London).
DISCUSSION. The species is named for Dr I. A. D. Robertson and his wife Ann who are formidable collectors of acridids. The species occurred in rather dry Acacia woodland in the same biotope as Physocrobylus burtti Dirsh (described in Jago, 1978).
MICROUSAMBILLA gen. n.
(Figs 27-33)
Type-species : Usambilla cylindricollis Ramme.
DIFFERENTIAL DIAGNOSIS. Male. Vertex with narrow selliform groove opening forward into disc-shaped fastigium (Fig. 27). Pronotum rounded above, lacking median carina (Fig. 29). Antennae slightly flattened, not widened and about 1.25 times length of head and pronotum. Ninth abdominal tergite deeply excavate (Fig. 28), bearing black tubercles on either side of median emargination. Supra-anal plate simple, triangular, bearing two large submarginal tubercles laterally and up to three pairs of small tubercles on disc. Penis complex like that in Usambilla, with large racket-shaped anterior valves and aedeagus bearing a single pair of pre-apical lateral teeth (Figs 32, 33). Epiphallus simple, ancorae (Fig. 31) weak and forwardly directed, ventro-lateral lophus (Figs 30, 31) at right angles to disc and with single apical hook. Hind femur 3.8 times longer than deep, rather slender.
Female. Head as seen from above very similar to that of male. Pronotum with traces of median dorsal carina. Antenna about 1.25 times length of head and pronotum. Ventral ovipositor valves slender, pointed. Hind femora slender, like those of male.
Coloration. General colour pattern of both sexes similar (Fig. 29). Light brown above, dark brown laterally. Dorso-laterally, pair of faint whitish stripes, side lobe of pronotum with another arcuate band of same colour but twice as deep. Hind femora olive, brown above. Hind tibiae brown, sooty below.
REVISION OF USAMBILLA
11
Figs 27-33 Microusambilla cylindricollis. 27, dorsal aspect of male head. 28, dorsal aspect of male supra-anal plate. 29, dorsal aspect of female pronotum. 30, right side, epiphallus and lateral plate, male. 31, epiphallus of male, right side. 32, entire male phallic complex (epiphallus removed), lateral oblique view, left side. 33, cingular rami and apical valvulae, rear aspect. All scale lines represent 0.5 mm.
Figs 34-40 Altiusambilla modicicrus. 34, dorsal aspect of male head. 35, dorsal aspect of male supra- anal plate. 36, dorsal aspect of female pronotum. 37, entire male phallic complex, apical valvulae, rear aspect. 39, left side of epiphallus and lateral plate, male. 40, right side of epiphallus. Scale line under Fig. 37 represents 0.5 mm and applies to Figs 35, 37-40. Other scale lines represent 0.5 mm.
12 N. D. JAGO
DISCUSSION. The genus is monotypic. It is morphologically close to Sygrus from which it differs in having a flattened sloping fastigium of the vertex and a weaker system of black nodules on the male ninth tergite. Sygrus also has a more definite dorsal pronotal carinula (see key, p. 3). In future, when further material is available, it may be best to unite Sygrus and Microusambilla under one genus.
Microusambilla cylindricollis (Ramme) comb. n.
Usambilla cylindricollis Ramme, 1929: 302, fig. 27a. Holotype <3, ZIMBABWE: Mashonaland, Chirinda forest, 1 150 m, 22.ii.1907 (C. F. M. Swynnerton) (BMNH) [examined].
MEASUREMENTS
Male Female
Head width 3.0 4.9
Posterior femur length 7.7 9.0
Posterior femur depth 2.0 2.0
Body length 12.1 14.2
MATERIAL EXAMINED
Zimbabwe: 2 & 3 ?, Monarch Mine, Umtali, 1213 m, 6.iv.l945 (N. C. E. Miller) (BMNH);6$, same data but 7.iv. 1945 (BMNH).
ALTIUSAMBILLA gen. n.
(Figs 34-40)
Type-species : Lentula modicicrus Karsch.
DIFFERENTIAL DIAGNOSIS. Male. Head and fastigium rugulose (Fig. 34) with transverse carinulae in depressed anterior part of fastigium. Upper part of frons, between antennal bases concave and widened. Groove of vertex ending just behind narrowest approximation of compound eyes dorsally, two oblique ridgelets separating it from the highly pitted occiput. Pronotum smoothly rounded above, median carinula being weak but present along whole length of disc. Dorso-lateral light pronotal stripe very narrow while ventral band on pronotal side lobe very deep, extending from pronotal margin one-third of way up side lobe. Posterior margin of ninth abdominal tergite gently concave, bearing one and sometimes two black tubercles on each side (Fig. 35). Supra-anal plate without marginal callosities of the Rhainopomma type (Figs 50-54) but with at least six pairs of tubercles on the disc and base of distal appendage (Fig. 35). Phallic complex like that in Lentula, with small auricular anterior penis valves (Fig. 37). Penis valves styliform with apical barbs (Fig. 38). Epiphallus with no ancorae and with lophal lobes slender, turned upwards at 45° to epiphallic disc (Fig. 40) and ruggedly hooked at tip (Fig. 39). Length to depth ratio of hind femur 4.
Female. Angular differentiation of frontal groove and fastigium more pronounced than in male, latter being pitted and sloping forward at 45°C to horizontal body axis. Lateral body stripes relatively undifferen- tiated. Dorsal median pronotal carina forming a large hump in the metazona (Fig. 36). Tergites of abdomen sharply carinulate. Tips of ventral ovipositor valves quadrilateral with acutangular apices.
Coloration. Fore and midlegs yellow green. Hind femur yellow green ; knee lenules blackish, rest brown. Hind tibiae pale bluish yellow-green, sooty apically. Generally olivaceous brown or greenish with creamy pronotal stripes in males. Dark brown to blackish side stripe in males and females from behind eye to side of first abdominal segment. On latter segment often black, continuing as broken black zig-zag on each side to abdominal segment 6.
Altiusambilla modicicrus (Karsch) comb. n.
Lentula modicicrus Karsch, 1896: 280. Holotype J, TANZANIA: Madjame, Mt Kilimanjaro (MNHU, Berlin) [examined].
MEASUREMENTS
Males Females
Head width (64) 2.5-3.0, 2.69 (42) 3.0-3.5, 3.24
Posterior femur length (60) 6.0-7.0, 6.55 (39) 7.8-9.3, 8.67
Posterior femur depth (62) 1.5-1.8, 1.68 (39) 2.0-2.4, 2.20
Body length (59)10.9-13.4,12.11 (39)13.1-17.1,15.98
REVISION OF USAMBILLA 13
MATERIAL EXAMINED
Tanzania: 1 J, Moshi, 18.xi.1943 (£. Sum) (BMNH); 3 ?, Kirua Vunja, 28.ix.1952 (Guic/jarrf)(BMNH); 2 $, W. Kilimanjaro, Msituni, defoliating Finns patula, 10.xii.1975 (BMNH); 1 9, W. Kilimanjaro, Moshi, on Pinus patula (BMNH); 23 <$, 5 9, nymphs, Kilimanjaro, S. side nr Mandera hut, 2370-2730 m, 15.xi.1964 (Jago) (BMNH); 30 rf, 29 9, E. of Mt Meru, Ngurdota-Meru N.P., crater lake rim, forest glades, 22.xi.1964 (BMNH); 5 cJ, 1 $, Ngurdota-Meru N.P., Kisari L., 22.xi.1964 (Jago) (BMNH); 3 rf, 3 9, Mt Meru, 1426 m, 25.ii.1967 (£. 5. Brown) (COPR, London); 5 & 7 $, Jekukumia R., 3°14'S, 36°47'E, 2.vi.l972 (Robertson & Robertson) (COPR, London).
DISCUSSION. The genus is monotypic. The new generic name refers to the preference this species has for montane forest and lower heath zones on Kilimanjaro and Meru mountains.
The unusual pronotal morphology of this species is reflected in a less extreme manner in other female Lentulidae, e.g. Usambilla emaliensis, which also have an elevated crest in the pronotal metazona. Live material of A. modicicrus is interesting since its mating display shows a possible behavioural function for the dorsal humps. Males approach females, the female responding by jerking and opening the hind legs sideways, while pedalling the hind tibiae which are folded and unfolded alternately. The males often orientate incorrectly head to tail, the female responding by more violent jerking or kicking. If, however, the male approaches correctly from behind he climbs the female abdomen and nibbles at the female tergites after the fashion of some female cock- roaches. In A. modicicrus the male climbs forward until he nibbles the pronotal bulge, whereupon the female presents her abdominal tip for copulation. A similar function may be involved in the strange bilateral pits on the metazona of many flightless Catantopinae, e.g. Aresceutica and Serpusia species.
Its attack on Pinus patula was preceded by years in which this grasshopper was very abundant in the indigenous natural vegetation. At the higher habitats A. modicicrus lives in a zone prone to unstable weather conditions with irregular rainfall. It is possible that drought conditions, forcing the insect to sample exotic Pinus, were responsible for the initial phase of the adaptation that insects have shown to a new food plant. Forestry involving Pinus patula should be free of attack by this species if trees are planted in areas other than Meru and Kilimanjaro. The very similar Rhainopomma montanum found in the West Usambara and Pare mountains is unlikely to adapt to Pinus species. Care should be taken not to introduce A. modicicrus to other highland areas of East Africa. This could be easily done if young trees were transferred with egg pods hidden in soil at their roots.
RHAINOPOMMA gen. n.
(Figs 4 1-72)
Type-species : Adolfia usambarica Ramme.
DIFFERENTIAL DIAGNOSIS. Male. Differing from all other Lentulidae and particularly the closely similar genus Usambilla in the extreme apical position of the aedeagal barbs (Figs 63, 64). Otherwise form of supra-anal plate (Figs 50-54) and epiphallus as in Usambilla (compare Figs 65, 144). Inter-ocular space very narrow, a feature correlated with great inflation of the compound eyes (Figs 41-45).
Female. Vertex narrower in proportion between compound eyes than in females of Usambilla (compare Figs 49, 83), but this apart identical with that genus.
Coloration. Male more brightly coloured than the female. Hind tibiae blue in some species, a character so far known from only one Usambilla species.
DISCUSSION. Distribution of the genus suggests a long period of evolution in wet lowland to middle altitude montane forests in eastern Africa, east of the Rift Valley. Habitats generally in higher rainfall areas than those of Usambilla. Females only surely identified generically by association with the males.
The genus is distributed in SE. Kenya and the piedmont forest blocks of the Pare, Usambara, Nguru, Uluguru mountains and Pugu hills. It is thus indigenous to forests on ancient uplifted mountain blocks which have not experienced the volcanism characteristic of the central rift. Their speciation suggests isolation in forest islets as a major factor in speciation.
14
N. D. JAGO
Figs 41-49 Rhainopomma species, dorsal aspect of head. 41, R. usambaricum, male. 42, R. montanum, male. 43, R. nguruense, male. 44, R. wapugu, male. 45, R. magnificum, male. 46, R. usambaricum, female. 47, R. nguruense, female. 48, R. wapugu, female. 49, R. magnificum, female. All scale lines represent 0.5 mm, that under Fig. 42 applies only to that figure.
Figs 50-54 Rhainopomma species, male supra-anal plate. 50, R. usambaricum. 51, R. montanum. 52, R. nguruense. 53, R. wapugu. 54, R. magnificum. Scale line represents 0.5 mm.
Key to species
Males
1 Knees of posterior femora blue R. usambaricum (Ramme) (p. 16)
Knees of posterior femora red-brown 2
2 Posterior tibiae sooty brown in apical four-fifths R. wapugu sp. n. (p. 19)
Posterior tibiae uniformly coloured or only with black or brown pigment at extreme tips . . 3
3 Supra-anal plate bearing fewer than 10 black teeth or denticles on disc (Fig. 42). Small insects
without clear dorso-lateral light body stripes R. montanum (Kevan) (p. 17)
Supra-anal plate with at least 14 black teeth or denticles on disc. Larger insects, usually with clear dorso-lateral body stripes .... . .... 4
REVISION OF USAMB1LLA
15
Light body stripes white, upper and lower stripe viewed laterally being clearly delineated above.
Smaller insects, body length under 15mm. Hind tibiae bright blue . . R. nguruense sp. n. (p. 17) Light body stripes creamy to buff in colour with upper lateral stripe more poorly delineated on its
upper margin. Large insects, body length never less than 16 mm. Hind tibiae dull green.
R. magnificum sp. n. (p. 20)
Females
1 Hind tibiae uniform blue, spines black tipped 2
Hind tibiae green with or without blue-black pigment 3
2 Hind knee lunules (Figs 55, 59) dark slatey blue, blacker above. Hind femur otherwise yellowish.
Light dorso-lateral and lateral body stripes golden yellow in colour
R. usambaricum (Ramme) (p. 16)
Hind knee and lunules (Figs 56, 61) light brown. Hind femur dark olivaceous with dorsal part brown. Light dorso-lateral and lateral side stripes light ochreous brown
R. nguruense sp. n. (p. 19)
3 Hind tibiae distally blue-black; proximal quarter green. Pronotum with weak but distinct convex
carina dorsally R. wapugu sp. n. (p. 19)
Hind tibiae light yellow-green. Pronotum dorsally lacking median carina and decumbent, con- cave in profile, or lightly convex, especially at rear 4
4 Hind femur 3.25 times longer than deep. Large insects; body length 18 mm. Upper carina of hind
femur convex (Figs 58, 62) R. magnificum sp. n. (p. 20)
Hind femur over 4.00 times longer than deep. Smaller insects with body length 15 mm. Upper carina of hind femur almost straight R. montanum (Kevan) (p. 17)
Figs 55-62 Rhainopomma species. 55-58, male posterior femora of (55) R. usambaricum; (56) R. nguruense; (57) R. wapugu; (58) R. magnificum. Scale line under Fig. 58 represents 1 mm and applies throughout. 59-62, lateral aspect of female head, thorax and hind femur of (59) R. usambaricum; (60) R. wapugu; (61) R. nguruense; (62) R. magnificum. Scale line under Fig. 61 represents 5 mm and applies throughout.
16 N. D. JAGO
Rhainopomma usambaricum (Ramme) comb. n. (Figs 4 1,46, 50, 55, 59, 63-65)
Adolfia usambarica Ramme, 1929: 305, figs 28b, 29b, 30a. Holotype ._?, TANZANIA: Amani, Usambara, iii. 1906 (Vosseler) (MNHU, Berlin) [examined].
DIFFERENTIAL DIAGNOSIS. Male. Antennal segments 1-3 green, 10 and 13 white, rest black. Head (Fig. 41) with dorsal triangular occipital area black, frons and two stripes behind and from ventro-posterior edge of compound eye, dark brown. Vertex, a band round back edge of compound eye and lower half of genae, white. Pronotum dark brown with pair of dorso-lateral and lateral wide white bands, which are yellow in metazone. White pronotal bands continued along abdomen as yellow bands, abdominal sternites and sub-genital plate being yellow. Supra-anal plate (Fig. 50) green, with two pairs of basal and three pairs of submarginal black tubercles. Hind knees (Fig. 55) blue, knee lunules grey, rest of femur yellow-green. Fore and mid-femora yellow to green. Hind tibiae blue; spines white, black-tipped. Aedeagus (Figs 63-65) and epiphallus; aedeagal sheath pear-shaped from posterior aspect (Fig. 64) (like that of R. nyuruense, R. montanum and R. wapugu), lateral epiphallic sclerites large in proportion to rest of epiphallus (unlike those of R. montanum, Fig. 69), with lower lophal lobe of epiphallus pointed but not hooked (Fig. 65, upper, left).
Female. Dark brown markings on head and pronotum as for male (Fig. 59), but equivalent white areas yellow, while median dorsal pronotal carina delineated with yellow. Fastigium of vertex (Fig. 46) black with yellow margins. Dorso-lateral yellow bands on body clearly marked back to at least abdominal tergite 1. Antennal markings similar to male, but black areas dark brown in this sex. All femora green, hind femora yellow on inner side. Hind knees and posterior tibiae dull blue.
MEASUREMENTS
Males Females
Head width (13) 3.4^4.0,3.64 (20) 4.0-5.0,4.45
Inter-ocular distance (14) 0.10-0.24, (20) 0.34-0.49,0.42
Posterior femur length (14) 7.6-9.5,8.37 (20) 9.3-11.0,10.18
Posterior femur depth (14) 2.0-2.7,2.32 (20) 2.5-3.1,2.85
Total length (14) 13.4-17.3, 14.90 (20) 17.2-21.0, 18.90
MATERIAL EXAMINED
Tanzania: 1 ?, same data as holotype (BMNH); 1 9, Amani, iv.1905 (BMNH); 3^, Sigi nr Amani, 460 m, ll.vi.1937 (£. Burn) (BMNH); 7 J, 1 9, same locality but 24.vi.1937 (E. Bunt) (BMNH); 4^, 4?, same locality but 29.V.1932 (£. Bum) (BMNH); 1 <$, 1 9, same locality but 6.vii.l937 (£. Burtt) (BMNH); 1^, same locality but 4.ii.l937 (£. Burtt) (BMNH); 3 & 5 9, same locality but v.1937 (£. Bunt) (BMNH); 1 rf, Kwamtili plant, iii. 1952 (Phipps) (BMNH); 1 J, 2 9, Ngomeni, Mlingano for., i.1953 (Phipps) (BMNH); 1 9, same locality but vii.1952 (Phipps) (BMNH); 5 rf, 6 9, 1 nymph, E. Usambara Mts, Sigi nr Amani, [2-1 l.]iv. 1964 (N. D. Jago) (COPR, London); 1 <J, 7 ?, 3 nymphs, same locality, [1 8-3 l.]xii. 1965 (N. D. Jago) (BMNH); 1 cJ, 1 9 nymph, W. Usambara Mts, Mkussu for. res., 1 l.xi.1964 (N. D. Jago) (BMNH); 2^, 19, E. Usambara Mts, Kwamkora for. res., 7.iv.l966 (N. D. Jago) (BMNH); 2^, 1 9, E. Usambara Mts, Amani-Sigi for. res., xii.1966 (N. D. Jago) (BMNH); 1 <J, 1 9, E. Usambara Mts, Amani Rest. Ho., 5.iv.l966 (N. D. Jago) (BMNH); 4 J, 4 9, E. Usambara Mts, nr Amani, 9.xi.l964 (N. D. Jago) (BMNH). Kenya: 1 J, 49, Shimba hills, iii. 1941 (VanSomeren) (BMNH).
DISCUSSION. R. usambaricum occupies wet lowland forest in the east Usambara mountains and immediately adjacent hills. Its ecological preference is typical for the other members of the genus, which also prefer wet, warm and sunny lowland forest, in contrast to upland habitats. Chromous- ambilla parallels it in having brightly coloured males, but differs in occupying degraded montane forest at higher altitudes in the hinterland of east Africa. R. montanum (see material studied) penetrates a little higher into the wetter warmer forest areas of the West Usambara massif and converges in appearance on A. modicicrus to such an extent that for many years the two were confused taxonomically.
Rhainopomma montanum (Kevan) comb. n. (Figs 42, 5 1,68, 69)
Usambilla montana Kevan, 1950: 21 1, fig. 3b. Holotype^, KENYA: Teita hills, 1370-1680 m, shrubby bushes and forest clearings, 24.xii.1945 (Kevan) (BMNH) [examined].
REVISION OF USAMBILLA 17
DIFFERENTIAL DIAGNOSIS. Male. Much smaller than R. usambaricum. Antennae with segments 1-3 green but rest dull brown, segments 8 and 12 being light in colour, cream or green. Head (Fig. 42) dark brown with black markings, at upper end of frons, around antennal sockets, a pair of spots just behind posterior end of groove of fastigium of vertex, a pair of dorso-lateral narrow post-ocular stripes and a broader one laterally behind compound eye, all black. Dorso-lateral light brown band on pronotum very narrow (contrasting with R. usambaricum) but lower lateral cream band wide (half depth of dark brown band above it). Rest of thorax and abdomen light brown above narrowly edged with lighter brown; black to dark brown laterally. Abdomen green below; subgenital plate green. Supra-anal plate green with only three pairs of callosities on disc; marginal callosity on each side continuous, unbroken (Fig. 51). Fore and mid-legs dark green; hind femora (like Fig. 58) dark green, knees light brown. Posterior tibiae black at the extreme tip, rest green but light brown adjacent to knee. Spines white with black tips.
Female. Small with proportions of inter-ocular space much like those of/?, magnificum (Fig. 49). Body and head light brown dorsally with only a faint trace of a fine line of lighter colour along each side. Dark brown to black lateral pronotal band similar to that in Fig. 60, but lower lateral light brown to cream band deeper than in R. wapugu (half depth of dark band) and upper edge of dark lateral band on segments 1 and 2 of abdomen forming a smooth curve, not angulate. Lateral parts of tergites 1-5 with dark brown or black markings. Hind femora uniformly red-brown, or yellow on inner and lower sides, green on outer area with knee red brown. Hind tibia green, spines green with black tips; extreme tip of tibiae black, part near to hind knee light brown. Fore and midlegs green or brown.
MEASUREMENTS
Males Females
Head width (27) 2.7-3.0, 2.87 (31) 3.3-3.8, 3.80
Inter-ocular distance (27) 0.18-0.30,0.22 (31) 0.34-0.63,0.48
Posterior femur length (26) 6.1-7.0, 6.40 (31) 7.1-9.7, 8.07
Posterior femur depth (26) 1.6-2.0,1-79 (31) 2.1-2.4,2.22
Total length (26)11.1-13.2,11.74 (29)13.8-20.3,15.65
MATERIAL EXAMINED
Kenya: 4 <$, 59, 1 9 nymph, Taita hills, vi.1948 (Van Somm>n)(BMNH); 19, Kenya colony, 1921 (/I. F. J. Gedye) (BMNH); 26 J, 29 9, 9 nymphs, Taita farmers' training centre, 8 km S. of Wundonyi, 6.V.1975 (/. A. D. & A. Robertson) (COPR, London). Tanzania: 6 J, 7 9, W. Usambara Mts, forest above Mazumbai, 8.vii.l967;3J, 1 9, same locality, nr irrigation canal, 8.vi. 1963; 2 <J, 29, 1 cJ, 29 nymphs, W. Usambara Mts, Amboni estate, below Mazumbai, 8.vii.l967; 6 J, 2 9, Mombo-Same rd, foot of S. Pare Mts, Gonja, wet forest, 13.xi.1964; 3 J, 3 9, W. Usambara Mts, Mazumbai F. Res., vi.1967; 1 <£ W. Usambara Mts, nr Bumbuli, Mazumbai, l.i.1967; 15 J, 1 9, W. Usambara Mts, Sumamagamba F. Res., 12.xi. 1964; 14^,8 9, W. Usambara Mts, Mkussu F. Res., ll.xi.1964; 1 £, W. Usambara Mts, Lushoto-Shume rd, 9.7 km from Lushoto, 1 l.xi.1964; 6 £, 4 9, W. Usambara Mts, Lushoto, 1670 m, 8.vii.l967; 1 & 39, W. Usambara Mts, nr Shume, ll.xi.1964; 1 9, W. Usambara Mts, Lushoto arboretum, ll.xi.1964; 6 J, 89, W. Usambara Mts, Mazumbai, secondary regrowth in quinine plantation, 8.vii.l963. (All (Jago); all in BMNH except last COPR, London.)
DISCUSSION. Recorded eating Commellina species (A. and I. A. D. Robertson, pers. comm.) in its Taita hills habitat in Kenya. Here the partial cultivation of a hill slope at the edge of cultivated land offered a particularly favourable habitat, partly shaded by an Albizzia species and palms. Like its sister genus Usambilla this genus is favoured by the opening up of primary forest and population density may be very high, e.g. 100 + /square metre. This may contribute to the attack by Usambilla on exotics planted in the course of forestry in such areas.
Rhainopomma nguruense sp. n.
(Figs 43, 52, 56, 6 1,66, 67)
Holotype J, Tanzania: E. foot Nguru Mts, Mtibwa Forest Reserve, nr Turiani, 7.xi.l964 (N. D. Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Segments 1-3 of antennae green; other segments black, except segments 10 and 14 which are white. Colour pattern on head as in R. usambaricum, but fastigium between eyes margin- ally narrower (Fig. 43). Light pronotal stripes white but otherwise pattern as in R. usambaricum. Meso- and metathorax dark brown, light body stripes white to pink. Posterior third of abdomen, including sub-genital
18
N. D. JAGO
Figs 63-72 Phallic complex in males of Rhainopomma species. 63, left lateral aspect of R. usambaricum. 64, posterior apical aspect of R. usambaricum. 65, epiphallus of R. usambaricum — upper right half and lateral plate, lower from right side. 66, left lateral aspect of R. nguruense. 67, posterior apical aspect of R. nguruense. 68, posterior apical aspect of R. montanum. 69, epiphallus of R. montanum — upper right half and lateral plate, lower from right side. 70, epiphallus of R. magnificum — upper right half and lateral plate, lower from right side. 71, left lateral aspect of R. magnificum. 72, posterior apical aspect of R. magnificum. Scale line under Fig. 71 represents 0.5 mm and applies throughout.
REVISION OF USAMBILLA 19
plate, pale green. Supra-anal plate (Fig. 52) with two pairs of basal tubercles and two or three pairs on disc (therefore resembling Fig. 53). Posterior femora green with knees red-brown (Fig. 56). Fore and mid-legs entirely green. Posterior tibiae light red-brown near knee and black at extreme tip; rest blue with black tipped white spines. Aedeagal complex (Figs 66, 67) showing aedeagal valves less sharply deflexed forwards than in R. usambaricwn (Fig. 63) but more elongate than those of/?, montanum (Fig. 68). Epiphallus like that of/?, usambaricum (Fig. 65).
Female. Body colour and pattern (Fig. 61) like that of R. usambaricum but light body stripes white, not yellow. Median dorsal pale line weak or absent. Hind femora yellow to green; knees brown. Posterior tibiae pale blue, brown near knee, brown at extreme tip and with red-tipped white spines.
MEASUREMENTS
Males Females
Head width (23) 3.6-4.5, 3.79 (18) 4.2^4.7, 4.54
Interocular distance (23) 0.16-0.30,0.23 (18) 0.46-0.59,0.53
Posterior femur length (23) 7.2-8.5, 7.96 (18) 8.5-10.5, 9.86
Posterior femur depth (23) 2.1-2.6, 2.36 (18) 2.7-3.2, 2.92
Total length (23) 14.0-15.8, 15.03 (18) 17.1-20.0, 18.71
MATERIAL EXAMINED
Paratypes. Tanzania: 16 J, 10 ?, 6 nymphs, same data as holotype but [5-7.]xi.l964 (BMNH); 1 <$, same data as holotype (BMNH); 1 J, same data but 7.xi.l964; 5<$, 5$, 3 nymphs, same data but 5.xi.l964(COPR, London).
DISCUSSION. This species is closely similar to R. usambaricum and was probably derived from common stock. Isolation in the adjacent but ecologically isolated forests of the east Usambara and Nguru mountains would explain the divergence between the two species.
Rhainopomma wapugu sp. n.
(Figs 44, 53, 57, 60) Holotype J, Tanzania : Pugu hills, SW. of Dar es Salaam, 1 1 .iii. 1967 (N. D. Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Antennal segments 1-5 cream to light brown; rest dark brown excepting segments 10 and 14 which are white. Interocular space narrowest for genus (Fig. 44). Pattern similar to that of /?. usambaricum but light areas white to pink, while dark areas are dark red-brown. Fore and mid-legs entirely green. Hind femora (Fig. 57) light green with light brown genicular part. Hind tibiae light brown near knee. Distal three-quarters dark brown, remaining proximal section green. Tip of abdomen pale green. Supra-anal plate (Fig. 53) with two pairs of basal tubercles and three pairs of tubercles on disc; apical tongue rather large, two-fifths length of whole plate when measured on mid-line.
Female. Antennal segments 1-3, apical parts of 4, 5 and 6, whole of 9, 11 and 15 white, rest black. Dorso-lateral light body stripes weak, narrow, thus differing from R. usambaricum and /?. nguruense, but resembling /?. montanum. Dorsal side of head with pair of black triangular marks on either side of dorsal mid-line. Dark brown lateral stripes (Fig. 60) about 3 times deeper than lower light pink side stripe and angulate on tergite 1 of abdomen. Fore and mid-legs green, femora brown dorsally. Hind femora light brown above and internally with large grey spot at base; outer area green; lower outer area yellow; knee entirely red-brown. Colour of hind tibiae as for male.
MEASUREMENTS
Males Female
Holotype Paratype Paratype
Head width 3.4 3.6 4.37
Inter-ocular distance 0.15 0.17 0.48
Posterior femur length 7.2 7.2 9.54
Posterior femur depth 2.1 2.1 2.77
Total length 13.7 13.4 17.72
MATERIAL EXAMINED
Paratypes. Tanzania: 1 $, 1 9, data as holotype (BMNH).
20 N. D. JAGO
Rhainopomma magnificum sp. n.
(Figs 45, 49, 54, 58, 62, 70-72)
Holotype <$, Tanzania: Mombo-Same rd, foot of S. Pare Mts, Gonja, wet forest, 13.xi.1964 (N. D. Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Large species, total length about 17 mm. Antennae green in basal half, darkening to brown distally with light cream tip. Vertex and occiput (Fig. 43) with relatively wide ocular interspace, dark brown dorsally and on frons. Cream area between frons and sub-ocular sulcus; gena dark brown bordered above by very narrow cream stripe from rear of compound eye to back of occiput. Pronotum and body back to tergite 3 of abdomen with dark brown dorsal stripe bordered laterally by bold dull yellow bands equal in width to that on pronotal side lobe (unlike female where lateral pale stripe like that of male but dorsolateral pale band lacking (Fig. 62)). Fore and mid-legs green. Hind femora green with knees light brown (Fig. 58). Hind tibiae light green, light brown at each extremity and with black-tipped green spines. Abdomen, including supra-anal plate, brown above, green below. Sub-genital plate green. Supra-anal plate (Fig. 54) with up to seven pairs of non-marginal tubercles; much wider than long. Base of aedeagus (Figs 71, 72) parallel-sided, not flask-shaped. Epiphallus with well-developed hook on apex of ventral lophi and large anterior projection of upper lophal lobe (Fig. 70).
Female. Head and body light brown above. Antennae like those in male. Dark black band behind compound eye bordered with narrow cream stripe ventrally (Fig. 62). Upper margin of dark brown side stripe with distinct up-curved zone at front of pronotal prozona and above side of thoracic tergite III and abdominal tergite I. Black zone extends from abdominal tergite I along side of tergites to back of segment 4. Ovipositor valvulae greenish. Fore and mid-legs green. Hind femora and tibiae as in male, but femora light brown dorsally. Large insects for the group (see measurements).
MEASUREMENTS
Males Females
Head width (5) 4.1^4.4,4.28 (3) 5.1,5.1,4.4*
Inter-ocular distance (5) 0.29-0.32,0.31 (3) 0.79,0.84,0.69*
Posterior femur length (5) 9.0-9.8,9.28 (3) 11.1,11.7,9.4*
Posterior femur depth (5) 2.6-2.9, 2.74 (3) 3.5, 3.4, 2.8*
Total length (5) 16.0-18.7, 17.57 (3) 21.8, 21.5, 18.2*
* Specimen from slightly higher elevation at Soni.
MATERIAL EXAMINED
Paratypes. Tanzania: 3 ^, 2 ?, 1 nymph, same data as holotype (1 <$, 1$, COPR, London; rest BMNH); 1 cJ, data as holotype but 10.vi.1967; 19, Soni, 17.xi.l 950 (J.PWpps) (BMNH).
USAMBILLA Sjostedt
Usambilla Sjostedt, 1909: 191. Type-species: Usambilla olivacea Sjostedt, by original designation.
Adolfia Rehn, 1914: 147. Type-species: Adolfia insolita Rehn, 1914: 148, by original designation. [Homonym
of Adolfia Guerich, 1909 (Brachiopoda).] Rehnula Uvarov, 1939: 457. [Replacement name for Adolfia Rehn.] [Synonymized by Dirsh, 1956: 152.]
DIFFERENTIAL DIAGNOSIS. Male. Vertex with narrow (Fig. 86) to wide (Fig. 92) inter-ocular sulculus. Frons depressed (Figs 90, 91) with frontal ridge almost flat, hardly elevated. Dorsal pronotal carinula moderately developed to weak or absent. Supra-anal plate, but not ninth abdominal tergite, armed with marginal callosities as well as elevated black nodules on disc (Figs 94-101). Epiphallus simple, lacking true ancorae but with hook-like ventro-lateral lophus (Figs 144, 145, 150, 131, 134, 137, 140) terminating in one, two or four hooks. Penis valves with lateral barbs situated short of apex (Fig. 133).
Female. Vertex proportionately wider than male, but with no special generic facies. Ventral ovipositor valves unspecialised (Figs 148, 149), not shovel-shaped as in Chromousambilla (Fig. 24).
DISCUSSION. The genus is known from predominantly upland areas of eastern Zaire, Ruanda, the forests of Uganda, south-western, central and northern Tanzania and the southern half of Kenya. The species occupy a wide range of ecological niches. Thus U. haematogramma lives in the upper Ufipa plateau forest margins, but U. insolita and U. sagonai in clearings in the wettest tropical forests of Ruanda, Zaire and Uganda. U. turgidicrus lives in upland scrub and woodland in
REVISION OF USAMBILLA
21
90
87
Figs 73-93 Dorsal or frontal aspect of head of Usambilla species. 73, male V. turgidicrus turgidicrus. 74, female, same. 75, male U. oraria. 76, female, same. 77, male U. ajfinis affinis, Morogoro. 78, male V. affinis kikomboensis, Ilonga. 79, female, same. 80, male V. emaliensis. 81, female, same. 82, male U. chlorophrygana. 83, female, same. 84, male U. leptophrygana. 85, female, same. 86, male U. insolita paratype, Kwidschwi I. 87, same, frontal aspect. 88, male U. sagonai sagonai, Rwanda. 89, male U. sagonai fractolineata. 90, same, frontal aspect. 91, female, same. 92, male U. haematogramma. 93, female, same. Scale line under Fig. 92 represents 1.0 mm and applies throughout.
Kenya and northern Tanzania (Combretum, Commiphora and Acacia or Juniperus associations), while U. leptophrygana and U. chlorophrygana occupy drier warmer woodland at lower altitudes. The ecological diversity is matched by subspecific variation in several species, suggesting active speciation in progress.
22
N. D. JAGO
Figs 94-101 Male supra-anal plate of Usambilla species. 94, U. turgidicrus. 95, U. oraria. 96, U. qffinis. 97, U. leptophrygana. 98, U. insolita. 99, U. sagonai sagonai. 100, U. sagonai fractolineata. 101, U. haematogramma. Scale line under Fig. 101 represents 0.5 mm and applies throughout.
Key to species and subspecies
Males
1 Ocular interspace dorsally more than 2.5 times width of first antennal segment (Fig. 92). Lateral
black side stripe narrowly edged above with red pigment (as in Fig. 129,9). Supra-anal plate (Fig. 101) with broad black area confluent with both basal tubercles on each side. Hind femora light olive green, knee light orange-brown . . . .U. haematogramma sp. n. (p. 36) Ocular interspace narrower, almost twice width of basal antennal segment. Never with red stripe delineating upper edge of lateral black stripe. Supra-anal plate with marginal rugosity, but black pigment never confluent with basal tubercles 2
2 Interocular distance dorsally equal to width of basal antennal segment (Figs 75, 86, 87). Supra-
anal plate with no more than a pair of basal tubercles on each side (Figs 95, 98) ... 3 Interocular distance at least 1.5 times width of basal antennal segment. Supra-anal plate with two pairs of basal tubercles (Figs 96, 99, 100, 152) or up to three basal tubercles on each side (Figs 94, 97, 99) 4
3 Hind femur (Fig. 116) more than 3 times longer than deep. (Rwanda and Zaire (?))
U. insolita (Rehn) (p. 34)
Hind femur (Fig. 108) very stocky and thickened from side to side, length to depth ratio about 2.6. Subgenital plate very short (Fig. 95) and folding under parameres which are received into impressions on its upper surface. Subgenital plate protrudes beyond supra-anal plate by about length of that plate as seen from above (in U. insolita projecting by roughly twice length of supra-anal plate). (Coastal Kenya and NE. Tanzania) U. oraria sp. n. (p. 32)
4 Dorso-lateral and light lateral body stripes (Figs 102, 103) bright yellow in life. Hind tibiae blue.
Fore and mid-femora conspicuously hairy above and below. Frons declivate, hardly protrud- ing as seen from above (Figs 88, 89). Median pronotal carinae absent or feeble ... 5 Dorso-lateral light body stripes; if present, dull grey, light brown but never bright yellow. Hind tibiae yellowish or greenish, with or without black pigment distally and ventrad. Frons declivate or decidedly protruding (Figs 73, 82). Fore and mid femora almost without hairs above. Median pronotal carina weakly to strongly developed (Fig. 105) 6
REVISION OF USAMBILLA
23
Yellow genal stripe continuous to back of occiput Yellow genal stripe interrupted between back
10
11
U. sagonai sagonai (Ramme) (p. 35) of gena and occiput. (Figs 102, 103)
U. sagonai fractolineata subsp. n. (p. 36) Hind femora bearing conspicuous black markings (Fig. 115) and of a uniform light brown colour. Frons protruding (Fig. 84) and incised as seen from above. Fastigium of vertex con- cave anteriorly. Posterior tibiae sooty below and towards tip . . U. leptophrygana sp. n. (p. 34) Hind femora immaculate or with light markings dorsally at base (Fig. 107), or with a light pregenicular spot (Figs 109, 110). Frons declivate, protruding or decumbent. Fastigium of
vertex deeply to shallowly concave anteriorly 7
Lateral dark brown side stripe intense, angularly bent in outline dorso-anteriorly on pronotum (Fig. 106). Frons in profile and from above (Fig. 73) seen to be very produced at its upper end, forming two tubercles which form margins of deeply incised fastigium of vertex. Hind femora light brown with or without darker mottling. Median dorsal pronotal carinula and its con- tinuation on tergites of abdomen weak to strong. Posterior tibiae sooty below and towards tip
8
Lateral dark brown side stripe with smooth upper outline (Fig. 104). Frons in profile and from above protruding (Figs 77, 78, 82) to strongly declivate and downwardly sloped (Fig. 90). Hind femora tending to be green with knee orange-brown. Median dorsal carinula of prono- tum weak (Figs 104, 105) to absent. Posterior tibiae unicolorous without black colour below 9 Hind femur very short and plump, length to depth about 3.0. (Vicinity of W. Usambara Mts)
V. turgidicrus olivacea Sjostedt (p. 27)
Hind femur less robust, length to depth ratio 3.3-3.4 . . U. turgidicrus turgidicrus (Karsch) (p. 25) Larger insects, body length just over 15 mm. Length to depth of hind femur (Fig. 113) about
3.40-3.50 U. chlorophrygana sp. n. (p. 32)
Smaller insects, body length just under 13 mm at most. Frons only weakly incised as seen from
above (Figs 77, 78, 80). Femur length to depth ratio 3.32-3.66 10
Pronotum lacking median dorsal carinula except for metazonal tubercle 11
Pronotum with clear median dorsal carinula (Fig. 105) in prozona and metazona. Hind femur slender (Fig. 112) with outer area green and rest tending to be brownish
U. emaliensis sp. n. (p. 30)
Length to depth ratio of hind femur 3.3-3.4 . . . U. affinis affinis Kevan & Knipper (p. 29) Length to depth ratio of hind femur about 3.6 . . . V. affinis kikomboensis subsp. n. (p. 30)
102
104
105
Figs 102-106 Lateral or dorsal aspect head and/or pronotum of male Usambilla species. 102, U. sagonai fractolineata, Uganda, Mpanga (darker form). 103, same, Uganda, Mpanga (lighter form). 104, U. emaliensis. 105, same, from above. 106, U. turgidicrus turgidicrus. Scale line under Fig. 102 represents 1 mm and applies throughout.
24 N I) JA(K)
I .iniil.s
Females of I/, Insoliia were not available for study; females of U, qfflnis offlnin and f. <t//)nfo kikomhoensts ciinnol he differentiated.
1 I aleial hlack side stripe bordered above with crimson (Fig. 129), Fastigium of vertex very broad
and lightly indented . U. kwmatogramma sp. n. (p. 36)
l.aicnil hlack side stripe, if present, never bordered ahovc with red. Fastigium of vertex much narrower and more deeply inoiied . . 2
2 Hind lihiac blue. (Colour very variable. No difference between races.) . . £/. saKonai (Kummc) (p. 35) Hind tibiae light brown, greeniih brown, green or ochrteeoui . . . 3
\ Inieiouilai spaaMloisallv 4 tinuv, width of basalaiitcmi.il segment (Figs 76, 79,81) . 4
Inlerocular space dorsally 5 times width of basal antennal segment (Figs 74, 83, 85) . 6
4 Pi onotal median carina dorsally very pronounced (Fig. 1 25), hind femora very slender, length to
depth ratio about 3, ft . ... U. tmalitnsix sp. n. (p. 30)
Median pronotal carina weak. Mind femur much more robust, length to depth ratio 3.0-3.2 . 5
5 Profile of pronotal carina and thorax curved (Fig. 121). Dark lateral tide stripe with upper edge
markedly bent in pro/ona. Mind lemur length to depth ratio around 3.0, Mind tibiae com-
pletely black vent rally in mature (?) specimens, ycllowish-ochraccous dorsally
. U. oraria sp, n. (p. 32) Piotile ni pionoial caima Halter (Figs. 122, 123, 124). Lateral side stripe, if developed, with
anterior upper border almost straight. Length to depth ratio of hind femur about 3.2. Hind
tibiae uniformly yellowish . U. qfflitis Kevan & Knippcr (p. 28)
(> Fronx markedly produced in profile (Figs 1 19, 120), Pronotal carina moderately strong, whole
disc tcctiform . ..... ...... 7
Frons less strongly produced in profile (Figs 12ft, 127), Median pronotal carina almost undevel-
oped, disc depressed . 8
Mm, i lemur more robust, length to depth ratio roughly 3.0 or less. Pronotum wrinkled and
punctate above (Fig, 1 20) . (/. turxMicrus olivacta Sjostcdt (p. 27)
Mind femur more slender, length to depth ratio about 3.2. Pronotum not wrinkled, finely punc-
tate (I..- 119). . (/. turxitticrus tur nidi cms Karsch (p. 25)
8 Large insects, body length more than 20 mm, Probably lack lateral body stripe. Posterior femora
dull olivaceous with knee light brown (Figs 83, 12ft) . . . U. cMontphrygaita sp. n. (p. 33)
Siu.illiM rugose msivls. boih length about IS mm. latci.il sulc stupes \\i-.ik cuvpi KM lust
abdominal tcrgitc and latero-antcrior part of pronotum (Fig. 127). Posterior femora brown
with darker mottling , , U. Itptopkrynaita sp. n. (p. 34)
I hambilla turffitlicrus (Karsch) (Figs 73, 74, 94, 106, 107, 1 19, 120, 131-133) /.c-Miii/ii lurtiitlicrus Karsch. 1896: 280.
DIIIIKINIIAI DIAONOSIN. Af«i/«', Head uu h deeply grooved fastigium of vertex forming a k notch at its point of contact with upper eiul of Irons (Fig. 73). Mead transversely ungulate at level of back end of median fastigial sulculus, occiput being wrinkled and pitted, Sides of vertex in front of eyes and area of ocellus on each side protruding, Pronotum lightly tcctiform on disc with low median carinula. Upper edge of dark lateral pronotal side stripe showing sharp angular change of direction between sulculi one and two. Dorso- lateral light brown to creamy stripes extending to tcrgite 8. Sides of tergites 1-6 of abdomen shiny black. Supra-anal plate (Fig. 94) with four pairs of strong black nodules on disc, two weaker ones at base. L'piphallic tophus with lour teeth (Fig, 131). Penis valves (Figs 132, 133) directed forwards at about 45° to long axis of body,
Fore and midlcgs light olive brown. Mind femora speckled blackish dorso-anteriorly (Fig. 107), with or without black comma in front of knee lunulc. Hind tibiae light olive brown, spines black-tipped. Hind tibiae sooty black or darker brown vcntrally in apical two-thirds or half. Lateral dark stripe on pronotal side lobe extended boldly along side of meso- and metathorax.
. Stocky (Fig. 1 19) with vertex and frontal ridge which meet in a raised protruding shelf in front of
compound eves 1'ionotum lu'htl\ tecliloim above, \\ilh .1 laUM.il d.uk xuK- stupr uiih .in .iiu'.ul.ii up|vi profile in front of second sulculus .uul i.ulnu-. .il>i upd\ Ivhuul (I iy, I I'M 1 .IUM.I! b.uul on ilioi.u-u- segments oK, iiu-. il.uk band intensified on alHloinm.il M-I-IIUMIIS I o. hl.uk .nul slum on M-J'.IIUMIIN .' c Ml.uk aiea i>n side of abdominal segment I variable in si/e.
Kl VISION 01 US
25
01)
ni|l> ill. in, |c liinwil II I ill VI-
107 118 Posterior femora of male lisunihillti species. 107, (/. nirfiulicrua olhwca, l./D .V(K), Momhi). IOS. (/. (ir<iri«, L/D 2.97. Momhasa. I(W. (/. «//!MM kikomhtwnsis, l./D 3.6, Kikombo. 110. same, L/D 3.36, llongti. I II, (/. aljinix u/finis, L/l> 3,28, Uluguru Mis. 112, (/. cmtilu'nsis, l./D «><>. Mmali range. 1 13, (/. chlorophrygana, L/D 3.46, Kikomho, Mpwapwa. 1 14, (/. chlorophryyana, dark lot in, l./D 3,26, 10 mis N. of Ussure. 115, (A /«7>f«>/)/irv</<//i<i, l./D 3.32. 1 16, (/. i>i.w//r<i l./D 3.45. 1 17, U. sagonalfractollntata, l./D 3.47. 1 18, (/. luu'matofirtimmd, l./D 3.55, Dlipa. Scale line under h'ig, 1 12 represents I mm and applies throughout.
Ventral valves of ovipositor simple, pointed. General colour grey to rugose dark brown with black to blackish pronotal side stripe, bordered below by short curved grey stripe. M-p.u.iti-d from ventral margin of side lobe by a band of dark brown or grey of roughly same depth.
Usambilla turiiMicrus turgidicrus (Karsch)
(Figs 73, 74,94, 106, 1 14>, 131 133) U-ntula luryidicrux Karsch. 18%: 280. Holotype •;. KHNYA: Kitui (MNIIll, Merlin) [exannned|
Dn 1 1 RI N IIAI DiA(iNosis. Male hind femora less robust than in subsp. olhwvti, length to depth ratio 3.3 3.4. Female similarly more elongate with slimmer femora, above ratio 3.2. Pronotum not wrinkled, finely punctate (Hg. 119).
26
N. D. JAGO
5mm
123
dg dull green
y yellow
ob olive brown
o olive
db dark brown
r red
Figs 119-130 Lateral aspect of head, thorax and posterior femur of female Usambilla species. 119, U. t. turgidicrus (S2794). 120, U. t. olivacea, W. Usambara. 121, U. or aria, Kenya, Rabai. 122, U. affmis kikomboensis, Ilonga. 123, same, lighter form. 124, same, intermediate form. 125, U. emaliensis. 126, U. chlorophrygana. 127, U. leptophrygana. 128, U. sagonaifractolineata. 129, U. haematogramma. 130, U. sagonai sagonai. Scale line under Fig. 125 represents 5 mm and applies to Figs 1 19-129. Scale line under Fig. 130 represents 1 mm and applies to that figure.
REVISION OF USAMBILLA 27
MEASUREMENTS
Males Females
Head width (17) 2.9-3.5,3.37 (29) 3.7-4.2,4.01
Posterior femur length (17) 6.3-7.9, 7.23 (28) 7.4-9.4, 8.50
Posterior femur depth (17) 1.8-2.4, 2.20 (28) 2.5-3.0, 2.68
Body length (16) 12.3-15.0, 13.75 (27) 14.8-18.1, 16.69
MATERIAL EXAMINED
Kenya: 1 9, Mtunguni hill nr Tulia, Kitui distr., 150 m, 01°12'S, 38°02'E, hillside with Combretum, Commiphora and Acacia woodland, 28.iv.1975 (Robertson & Robertson) (COPR, London); 1 <$, 19, nr Kibwezi, 5 km SE. of Mbuinzau, 02°24'S, 37°55'E, 940 m, lichens and spiney vegetation on recent lava, 5.V.1975 (Robertson & Robertson); 1 £, 1 $, 3 km N. of Mwatate, Wundangi rd, 03°27'S, 38°22'E, 910 m, fairly dense scrub woodland, 6.V.1975 (Robertson & Robertson) (COPR, London); 1531, 259, Mtunguni hill, 01°12'S, 38°02'E to 01°13'S, 38°02'E, nr Tulia, Kitui distr., 910 m, heavy juniper woodland and plantations, lO.v.1975 (Robertson & Robertson) (5 & 4 9, BMNH; rest COPR, London); 1 9, Athi R. crossing, 25.6 km NNE. of Kibwezi, 430 m, 22.vii.1934 (Vanderbilt Exped. Kenya Africa, Rehn) (BMNH); 1 <J, 1 9, S. Pare Mts, hillside above Gonja, c. 1000 m, [12-16] vi.1974 (Hollis) (BMNH).
DISCUSSION. More material of the nominate subspecies is required before the true junction between the subspecies can be determined.
Usambilla turgidicrus olivacea (Sjostedt) stat. n. (Figs 107, 120)
Usambilla olivacea Sjostedt, 1909: 186, 192. Holotype 9, TANZANIA: Usambara, Mombo, vi. (Sjostedt) (NR, Stockholm) [examined].
DIFFERENTIAL DIAGNOSIS. Male with hind femur length to depth ratio 3. Female with hind femur length to depth ratio 3 or less. Pronotum wrinkled and punctate above (Fig. 120). Subspecies markedly more stocky than nominate race. Male genitalia identical.
MEASUREMENTS
Male Females
Holotype
Head width 3.19 4.3 (4.8) 4.20
Posterior femur length 6.66 3.7 (4.0) 8.68
Posterior femur depth 2.25 9.3 (10.0) 2.90
Body length 12.31 14.7 (17.0) 16.15
The female holotype measurements in parentheses are those published by Sjostedt.
MATERIAL EXAMINED Tanzania: 1 ^, 1 9, Mombo, 9.vii. 1967 (Jago) (BMNH).
DISCUSSION. The dimensions of the holotype published by Sjostedt do not tally with the accurate modern ones. It is possible that his optical technique gave consistent overestimates. The holotype is thus a smaller insect than published measurements would indicate. U. turgidicrus olivacea represents the southernmost outliers of the nominate race.
In life the specimens from Mombo had a light silvery side stripe, the female being light grey in colour. Both have darkened considerably and are now brown.
Usambilla affinis Kevan & Knipper (Figs 77-79, 96, 109-111, 122-124, 134-136) Usambilla affinis Kevan & Knipper, 1961 : 372.
DIFFERENTIAL DIAGNOSIS. Male. Fastigium of vertex (Figs 77, 78) with margins diverging forward, lightly emarginate in front and with deeply to lightly impressed interocular groove (lightest in subsp. affinis to deepest in subsp. kikomboensis). Front end of fastigium pitted and sculptured. Frontal ridge and whole of
28
N. D. JAGO
Figs 131-141 Phallic complex of male Usambilla species. 131, epiphallus of U. turgidicrus turyidicrus — upper right half and lateral plate, lower from right side. 1 32, same, aedeagus, left lateral aspect. 133, same, posterior apical aspect. 134, epiphallus of U. qffinis ajfinis (Uluguru Mts) — upper right half and lateral plate, lower from right side. 135, same, aedeagus, posterior apical aspect. 136, same, left lateral aspect. 137, epiphallus of U. emaliensis (Emali range) — upper right half and lateral plate, lower from right side. 138, same, aedeagus, posterior apical aspect. 139, same, left lateral aspect with spermatophore in situ. 140, epiphallus of I/, or aria — upper right half and lateral plate, lower from right side. 141, same, aedeagus, posterior apical aspect. Scale line under Fig. 138 represents 0.5 mm and applies throughout.
REVISION OF USAMBILLA 29
frons evenly punctate and more protruding than U. oraria, but less so than in U. turgidicrus. Pronotum hardly to weakly tectiform above with pronotal carina absent to moderate. Abdominal tergites lightly carinulated dorsally. Supra-anal plate with marginal callosities (Fig. 96) and four to five pairs of tubercles, two or three pairs being distal. Penis sheath and valves apically vertical (Fig. 136). Epiphallus with lophal process ending in a simple hook (Fig. 134). Ancorae absent.
Female. Fastigium of vertex sloping forwards in a smooth convexity to meet upper part of frontal ridge (Figs 79, 122, 123, 124). Pronotum and abdominal tergites quite sharply carinulate dorsally. Tips of ventral ovipositor valves triangular, pointed.
Coloration of male and female more divergent in the closely allied U. emaliensis than in U. affinis. Male in general with yellowish to creamy markings at border of compound eye dorsally, on front of fastigium of vertex, below compound eye straddling fronto-genal suture. Lateral dark brown stripe variable in intensity, usually extending weakly onto side of abdomen. Light side stripes pale yellowish to creamy white, lower- most band separated from ventral margin of pronotal side lobe by darker pigment (Fig. 104) and extending across epimera and episterna of thoracic segments 2 and 3 as a whitish band. Fore and mid legs light olivaceous green. Hind femora green with ginger-brown knees (Figs 109-111). Very faint brown comma- shaped marking in front of genicular lunule. Female light brown above with darker median dorsal area, side stripe dark brown (but variable, see Figs 122, 123, 124). Hind femora light olivaceous green with dorso-basal brownish spot and weak sigmoid mark in front of outer and inner knee lunule. Knees light ginger-brown. Hind tibiae, as in male, light yellow-green with spines black-tipped.
Note. In both male and female upper profile of lateral dark pronotal side stripe forms a smooth convex curve (Fig. 124) unlike that in U. turgidicrus (Fig. 1 19).
DISCUSSION. This species forms a north-south cline in which the southernmost populations have short stumpy hind femora (subsp. affinis), the northernmost having slender hind femora. Separ- ating the components of the cline is difficult, though zoogeographically the nominate subspecies probably occurs in the Uluguru Mts and southwards, while subsp. kikomboensis probably occurs in scattered pockets from Ilonga through Kikombo to the west Usambaras.
Usambilla affinis affinis Kevan & Knipper (Figs 77, 96, 111,134-136)
Usambilla affinis Kevan & Knipper, 1961: 372, fig. 3, pi. 3, ff. 3-7. Holotype J, TANZANIA: Morogoro, Morningside, 31.vii.l954(Pfa>ps)(BMNH) [examined].
DIFFERENTIAL DIAGNOSIS. Male. Main difference in proportions of hind femur, its length to depth ratio at 3.3-3.4 being less than in subsp. kikomboensis. Males tend to have dark dorso-lateral body lines brown. Penis sheath and aedeagus erect, set at right-angles to long axis of body (Figs 135, 136). Females cannot be differentiated from other subspecies.
MEASUREMENTS
Males Females
Head width (3) 3.45,3.38,3.74 (3) 4.18,4.08,4.48
Posterior femur length (3) 7.24, 6.99, 7.48 (3) 9.14, 8.30, 9.10
Posterior femur depth (3) 2.21, 2.89, 3.12 (3) 3.00, 2.65, 2.95
Body length (3) 12.3, 12.93, 14.79 (3) 16.5, 17.22, 17.24
MATERIAL EXAMINED
Tanzania: 1 ?, Morogoro, 1954 (Phipps) (paratype of Usambilla affinis Kevan & Knipper) (BMNH); 1 <?, Morogoro, 1320 m, 24.xi.1939 (£. Burn) (BMNH); 1 rf, 1 $, Morogoro, xi.1939 (E. Burtt) (BMNH); 2cJ, Morogoro, 8.iii.l955 (Phipps) (BMNH); 1 ?, Morogoro, 4.xii.l939 (£. Burtt); I & Uluguru Mts, W. side, 1.6 km S. of Mgeta, 25.x. 1964 (Jago) (COPR, London); 1 nymph, Uluguru Mts, W. side, Bunduki For. Res., fishing camp, [20-23.]x. 1964 (Jago) (COPR, London).
DISCUSSION. The habitats of this subspecies in the Uluguru Mts are characterised by abundant tree ferns and tree lilies, relatively high rainfall and abundant sunshine.
30 N. D. JAGO
Usambilla affinis kikomboensis subsp. n.
(Figs 78, 79, 109, 110, 122-124) Holotype ^Tanzania: Ilonga, 16.vi.l 967 (Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Hind femora length to depth ratio about 3.6. More brightly marked than nominate subspecies, with dark brown stripes in that subspecies replaced by black and light body stripes golden yellow. Penis sheath and aedeagal valves directed vertically (as in Fig. 136).
Female. Pigmentation variable (Figs 122-124). Frons sloping upwards with a gentle angular change into fastigium of vertex. Face dark brown with black and cream spots or patches. Dark side stripe, when fully developed (Fig. 122), with a gently sinuous upper margin (may be reduced to a dark oblique bar, Fig. 123).
MEASUREMENTS
Males Females
Head width (14) 3.5-3.7, 3.63 (5) 4.1-4.5, 4.25
Posterior femur length (14) 7.4-8.0, 7.85 (5) 8.9-9.3, 9.13
Posterior femur depth (14) 2.3-2.9, 2.41 (5) 2.6-3.0, 2.79
Body length (14) 13.2-15.0, 14.18 (5) 16.9-17.3, 17.13
MATERIAL EXAMINED
Paratypes. Tanzania: 12cJ,5$, same data as holotype (5(J, 3?,COPR, London, rest BMNH); 1$, Mpwapwa, Mt Wilkins, 1460 m, 10.iv.1938 (E. Burn) (BMNH); 1 <J, Kikombo, Mpwapwa, 18.ix.1947 (BMNH).
Usambilla emaliensis sp. n.
(Figs 80, 81, 105, 106, 112, 125, 137-139) Holotype £, Kenya: Emali range, Sultan Hamud, 1210-1790 m, iii.!940(BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Penis sheath bent in a smooth arc capitad as in U. turgidicrus (cf. Figs 1 32, 139). Epiphallic lophus bifurcate at tip (up to four teeth in U. turgidicrus) (Fig. 137). Form of frons and vertex much as in U. turgidicrus but otherwise differing as follows: upper margins of dark lateral body stripe smooth in profile (Fig. 104), not angulate as in Fig. 106; pronotal median carina forming a higher crest (Fig. 105) than in U. turgidicrus, in this respect near to V. affinis; hind femur (Fig. 112) distinctly green in outer and lower outer areas (always light brown with darker mottling in nominate subspecies of U. turgidicrus); length to depth ratio about 3.66 (only about 3.30 in nominate subspecies of U. affinis); frontal ridge (Figs 80, 81) weakly incised, unlike U. turgidicrus but like U. affinis.
Female. With much more pronounced pronotal crest than U. turgidicrus (Fig. 125). Hind femur much more long and slender (Fig. 125) (length to depth ratio 3.6; this ratio in subspecies of U. turgidicrus 3.2 or less).
MEASUREMENTS
Male holotype Female paratype
Head width 3.20 3.75
Posterior femur length 7.36 9.05
Posterior femur depth 2.00 2.56
Body length 13.46 15.78
Figs 142-153 Phallic complex of male Usambilla species. 142, U. chlorophrygana aedeagus, posterior apical aspect. 143, same, left lateral aspect. 144, same, epiphallus — upper right half and lateral plate, lower from right side. 145, epiphallus of U. sagonai sagonai (Kibale forest) — left right half and lateral plate, right — right lateral aspect. 146, same, aedeagus, left lateral aspect. 147, same, posterior apical aspect. 148, 149, U. sagonai sagonai female, ventral ovipositor valves from below and left side respectively. 150-152, U. insolita male (150) epiphallus — upper right half and lateral plate, lower right lateral aspect; (151) aedeagus, posterior apical aspect; (152) aedeagus, left lateral aspect. 153, supra- anal plate of U. chlorophrygana male. Scale line under Fig. 146 represents 0.5 mm and applies throughout.
REVISION OF USAMBILLA
31
145
147
152
32 N. D. JAGO
MATERIAL EXAMINED Paratype. Kenya : 1 9, same data as holotype (in copula) (BMNH).
DISCUSSION. This species may represent an isolated offshoot of U. qffinis with which it shares the form of the vertex and pronotum. The very slender hind femora and high pronotal crest are, however, unique.
Usambilla oraria sp. n.
(Figs 75, 76, 95, 108, 121, 140, 141) Holotype^, Kenya: Mombasa, [3-4.]vii.l939(E. Bum) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Vertex very narrow between compound eyes dorsally (Fig. 75), frontal ridge depressed, hardly protruding. Dark body stripes dark reddish brown; light longitudinal stripes creamy white. Lower lateral light band extending across epimera and episterna of thoracic segments 2 and 3. Hind femora very squatt and inflated, length to depth ratio less than three (Fig. 108). Supra-anal plate (Fig. 95) very simple with four pairs of black callosities on the disc. Subgenital plate very short, folding under parameres for which there are two concave depressions in upper surface of subgenital plate itself. Cingular sheath small, rather abruptly tapered (Fig. 141) compared with U. qffinis (Fig. 135). Epiphallus with bifur- cate lophi (Fig. 140).
Female. Fastigium of vertex shallowly concave; frons hardly produced. Occiput with two large rugose ovoid depressions just behind back end of interocular groove. In lateral aspect with smooth convex dorsal profile giving a hump-backed appearance. Pronotum quite strongly tectiform. Hind femur short and stocky, length to depth ratio less than 3 (Fig. 121). Ventral ovipositor valves pointed.
General coloration light brown and dark brown pattern in both sexes. Hind femora unicolorous light brown (Fig. 108), though in darker females with dark comma anterior to knee lunules and marks on dorso-anterior part. Hind tibiae light ochrous brown, often blackish below, with black-tipped tibial spines.
MEASUREMENTS
Male Females
Head width 3.50 (3) 4.5-5.3,4.79
Posterior femur length 6.95 (3) 8.4^9.0, 8.74
Posterior femur depth 2.35 (3) 2.7-3.1, 2.93
Body length 11.53 (3)15.6-17.1,16.54
MATERIAL EXAMINED
Paratypes. Kenya: 2 ^, Mombasa, [3-4.] vii. 1939 (E. Burn) (BMNH) [compared with holotype of U. turgidicrus by V. M. Dirsh]; 1 9, Mombasa, on Vernonia hildebrandti, 12.iii.1969 (BMNH); 1 ?, Rabai, viii. 1937 (Van Someren) (BMNH); 3 9, Arabuko for., 17 km W. of Malindi, 03°13'S, 39°56'E, 61 m, ll.vi.1975 (Robertson & Robertson) (COPR, London); 1 9, Arabuko-Sokoke For.Res., Jilore track, c.50 m, [2-3.]vi.l974 (Hollis) (BMNH).Tanzania: 2 9, Mlingano, Ngomeni, v.1952 (Phipps) (BMNH); 1 9, W. Usambara, Suma- magamba F. Res., 12.xi. 1964 (Jago) (BMNH).
DISCUSSION. Probably one of the most truncated grasshoppers in proportion to its width known to science. It has a largely coastal forest distribution, with enclaves in piedmont forest in the west Usambara mountains. It belongs to the affinis-group of species.
Usambilla chlorophrygana sp. n.
(Figs 82, 83, 1 13, 1 14, 126, 142-144) Holotype <J, Tanzania: Kikombo, Mpwapwa, 16.iv.1947 (E. Burtt) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Large for the genus (see measurements). Fastigium of vertex strongly declivate, frons only protruding slightly in front of eyes. Anterior part of fastigium heavily pitted. Occiput with two oblique echelons of finely pitted cuticle terminating just behind interocular groove in two flat depressions (Fig. 82). Frons and genae wrinkled and punctate. Pronotum finely punctate with weak or obsolete dorsal carinula. Dorso-lateral and lateral light body stripes dull yellowish brown, upper band weak dorsad. Lateral creamy stripe extending across epimera and episterna of meso- and metathorax. Supra-anal plate with at least 3 pairs of distal black tubercles on disc (Fig. 153). Penis sheath strongly decurved capitad (Fig. 143) so that valves point forward. Epiphallic lophi simple; single hook at apex (Fig. 144).
REVISION OF USAMBILLA
33
Figs 154-161 Phallic complex of male Usambilla species. 154, U. haematogramma aedeagus, posterior apical aspect. 155, same, left lateral aspect. 156, same, epiphallus — upper right side and lateral plate, lower right lateral aspect. 157, U. leptophrygana, aedeagus, posterior apical aspect. 158, same, entire complex less epiphallus from above showing shape of ectophallic sheath and apodemes plus position of ejaculatory sacs (shaded). 159, same, entire complex less epiphallus, left lateral profile, showing broad flat spatulate anterior penis valves. 160, same, aedeagus left lateral view. 161, same, epiphallus — left right half and lateral plate, right lateral aspect. Scale lines represent 0.5 mm. Line under Fig. 154 applies to all except Figs 158, 159, to which scale line under Fig. 158 applies.
Female. Large (Fig. 126) with cuticle finely pitted. Female paratype badly discoloured but probably with light band of shiny cuticle across lower edge of pronotal side lobe.
Fastigium very wide, almost flat and punctate anteriorly (Fig. 83). Ovipositor valves apically pointed as in Fig. 188.
General coloration of male olivaceous brown with creamy orange dorso-lateral and lateral side stripes. Hind femora light green; knees light red-brown (Fig. 113). Hind tibiae light yellow-green to yellow, with black spines.
34 N. D. JAGO
MEASUREMENTS
Male holotype Female allotype
Head width 3.71 4.88
Posterior femur length 7.88 10.92
Posterior femur depth 2.51 3.15
Body length 15.36 20.89
MATERIAL EXAMINED
Paratypes. Tanzania: 2 9, same data as holotype (BMNH); 1 $, data as holotype but 17.iv.1947 (£. Bunt) (BMNH); 1 J, data as holotype but 18.iv.1947 (£. Bunt) (BMNH); 1 & data as holotype but 19.iv.1947 (£. Burn); 2 <$, Mpwapwa, Mt Wilkins, 1213 m, 4.iv.l938 (£. Bunt) (BMNH); 2 rf, 19, 16.1 km N. of Ussure, Msigiri road, 12.iv.1936 (£. Bunt) (1 <$, 1 9, COPR, London; 1 <$, BMNH).
DISCUSSION. The name of this new species is derived from the Greek 'chloros' — green and 'phryganos' — twig or stick.
Usambilla leptophrygana sp. n.
(Figs 84, 85, 97, 1 15, 127, 157-161) Holotype <$, Tanzania: 70.8 km N. of Dodoma, 18.vi.1967 (N. D. Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Males. Distinctive (Fig. 127) with well-marked to faint dark brown side stripe and conspicuous black markings on posterior femora (Fig. 1 15). Frons weakly to strongly incised in front (Fig. 84); fastigium of vertex excavate but much more produced than the similar U. chlorophrygana (Fig. 82). Overall colour light brown with dark markings. Hind femora not green in our series. Supra-anal plate with three basal tubercles on each side (Fig. 97) (see U. chlorophrygana, Fig. 153). Epiphallic lophus with single apical hook (Fig. 161), anterior penis valves (Figs 158, 159) typical for genus and illustrated here as representative, being flattened and vertically orientated. Aedeagal valves sloping strongly capitad (Figs 157, 159,160).
Female. Dark pronotal side band weak, fading behind pronotal metazone and only intensifying just behind tympanum. Hind femur with pre-genicular black spot and dorso-basal spot always present, but row of three dark spots along lower outer carina variably developed (Fig. 127). Overall colour dull brown with darker brown markings. Median pronotal carina present, if weak, throughout.
MEASUREMENTS
Males Females
Head width (3) 3.5-3.6, 3.58 (5) 4.1-4.5, 4.31
Posterior femur length (3) 7.7-8.4, 8.01 (5) 8.4-9.8, 9.19
Posterior femur depth (3) 2.3-2.5, 2.41 (5) 2.8-3.1, 2.95
Body length (3) 14.2-15.2, 14.81 (5) 16.4-18.5, 17.58
MATERIAL EXAMINED
Paratypes. Tanzania: 3 9, same data as holotype but 16.vi.1967 (BMNH); 1 cJ, 2 ?, Old Shinyanga, (l-8.)iii.!947 (£. Bunt) (BMNH); 3 & same data, 7.iii.l947 (BMNH); 3 ^,same data, Hi. 1947 (BMNH); 1 & same data, 12.iv.1947 (BMNH); 1 1 <J, 5 9, same data, early iii.1947 (1 & 1 9, COPR, London; rest BMNH); 7 cJ, Old Shinyanga, 30.iii.1947 (£. Bunt) (BMNH); 1 <J, 1 9, same data 30.iv.1947 (BMNH); 1 <J, Old Shinyanga, block 9, 23.V.1956 (£. Bunt) (BMNH); 2 & same data, [22-24.]iiil948 (£. Bunt); 1 9, same data, 24.iv.1948 (£. Bunt) (COPR, London); 3 J, Mkwemi, 17.7 km W. of Kahama, 29.iii.1947 (£. Bunt) (BMNH); 1 9, Ruaha N. P., 15.iii.l 966 (Vesey-FitzGerald) (BMNH).
DISCUSSION. The name of this species is derived from the Greek 'leptos' — slender, 'phryganos' — a twig.
Usambilla insolita (Rehn)
(Figs 86, 98, 150, 151) Adolfia insolita Rehn, 1914: 148. Holotype^, CONGO: L. Kivu, Kwidschwi I. (MNHU, Berlin).
DIFFERENTIAL DIAGNOSIS. Only male paratype examined (ex alcohol). Very similar to U. sagonai (see Figs 86, 87, 88, 90). Differs in having a much narrower vertex and frontal ridge hardly produced forwards. Zone of dark brown or black pigment in U. sagonai (Fig. 90) apparently absent in U. insolita, but this may be due to
REVISION OF USAMBILLA 35
loss of colour after attempted preservation in alcohol. Supra-anal plate similar to that of U. oraria (Fig. 98) but differing in having subgenital plate extending much further beyond tip of supra-anal plate (see key, couplet 3) and hind femur more slender (see Figs 108, oraria; 116, insolita). Epiphallus (Fig. 150) with stout hook at tip of ventral lophi and penis apex curved in an arc cephalad (Figs 151, 152). General colour pattern unknown.
MEASUREMENTS
Male
Head width 3.32
Posterior femur length 7.51 Posterior femur depth 2.07
Body length 13.53
MATERIAL EXAMINED
Paratype. Congo: 1 <$, Kwidschwi I., Lake Kivu, ix.1907 (Adolf Friedrich Duke of Mecklenberg Expdn) (COPR, London).
DISCUSSION. There is clear confusion in the original descriptions of species allocated to Adolfia by Ramme (1929) so that his expansion of U. insolita to include material from modern Zaire and the Ruwenzori region must be open to doubt. The paratype (topotype) examined here is different in detail from U. sagonai and is probably a species confined to the Kivu area only. It is replaced in Uganda by U. sagonai.
Usambilla sagonai (Ramme) (Figs 88, 89, 90, 99, 100, 102, 103, 117, 128, 130, 145-149)
DIFFERENTIAL DIAGNOSIS. Male. Differing from all other Usambilla species by the following combination of characters: dorso-lateral and light lateral body stripes (Figs 102, 103) bright yellow; hind tibiae blue; frons only very slightly produced forwards as seen from above (Figs 88, 89); hind femora green, knee lunules light reddish brown (Fig. 117); supra-anal plate (Figs 99, 100) open to variation but in general with two pairs of basal black tubercles and two pairs of black tubercles near margin at centre of disc; frontal ridge less coarsely pitted than that of U. insolita (Figs 87, 90); penis valves sharply curved cephalad (Figs 146, 147); ephiphallic lophi with single apical hook (Fig. 145).
Generally brightly coloured species found in wet evergreen forest. Oblique yellow stripe across gena from base of eye complete or broken (subsp.fractolineata, Figs 102, 103).
Female. Ovipositor valvulae unspecialised; bluntly pointed (Figs 148, 149). Brightly coloured in various complex shades of green and brown (Fig. 130). In the darkest forms lateral pronotal lobe and area across thoracic segments II and III plus abdominal tergite 1 bear shining black bands. In more lightly pigmented specimens the black areas may be replaced by brown or dark green (Fig. 128). Side of pronotum, below dark band, bright yellow.
In both sexes pronotum is smoothly arched dorsally with median dorsal carinula weak or absent and cuticle finely punctate. In general appearance they converge on Rhainopomma species but have a wider inter-ocular groove, and, of course, apical barbs of the male aedeagus are preapical.
Usambilla sagonai sagonai Ramme (Figs 88, 99, 130, 145-149)
Adolfia sagonai Ramme, 1929: 305, fig. 28c. Holotype £, ZAIRE; Lakes Region (MRAC, Tervuren)
[examined].
DIFFERENTIAL DIAGNOSIS. Differs from subsp.fractolineata by possessing a yellow genal stripe which is entire. MEASUREMENTS
Males Females
Head width (12) 3.1-3.7, 3.46 (9) 3.8-4.1, 3.96
Posterior femur length (12) 7.0-8.6, 7.95 (9) 8.4-9.4, 8.95
Posterior femur depth (12) 2.0-3.5,2.43 (9) 2.5-2.9,2.60
Body length (12)12.4-15.0,14.18 (9)15.7-16.9,16.44
36 N. D. JAGO
MATERIAL EXAMINED
Zaire: 1 & 2 9, La Chute For., Rutchuru, 7.viii.l949 (E. Burtt) (BMNH). Rwanda: 2 <J, Kisenye, 10.viii.1949 (E. Burtt) (BMNH). Uganda: 1 V, Toro, 1.6 km E. of Bundebugyo, 9.viii.l964 (Jago) (BMNH); 1 ?,Kamanve, 17.ix.33 (Johnston) (BMNH); 1 9, Kilembe, 1 370 m,xii. 1934-1.1935 (F. W. Awards) (BMNH); lc^Gaba,18.x.l931(G.//.£./f0p/uns)(BMNH);l J,Nsagu, 3.ix.l933(Jonnsron)(misdet.as V. insolita Rehn by B. P. Uvarov) (BMNH); 1 <J, Ruwenzori, 1610 m, 1913 (Scott-Elliott) (misdet. Adolfia insolita Rehn by Ramme) (BMNH); 8 <J, 6 9, Toro, SE. of Ft Portal, Kibale for. res., 30°25'E, 0°30'N, [13-16.]viii.l964 (Jago) (2 $, 2 9, COPR, London; rest BMNH); 1 cJ, 1 ?, same data, 30.iv.1967 (Jago) (BMNH); 1 9, Ruwenzori, between road and Nyabitaba hut below 2610 m, 27.vi.1963 (P. & P. Carter) (BMNH). Kenya: 1^, 19, Kakamega For. Statn, c. 1520 m, [18-19.] vii. 1974 (Hollis) (BMNH).
DISCUSSION. The recent discovery of this subspecies in Kenya extends its known distribution to the forests of the east side of Lake Victoria. It is possible that the nominate race occurs south and west of the lake while subsp. fractolineata represents an isolated series of populations on its northern side in the Mabira to Mpanga forest blocks of the Nile drainage and northwards to Lake Albert.
I samhilla sagonai fractolineata subsp. n.
(Figs 89, 90, 100, 102, 103, 117, 128)
Holotype <J, Uganda: Buganda, Mpanga F. Res., km 33.8 Kampala-Masaka road, 32°20'E, 0°15'N, [2-3.] viii. 1964 (Jago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Differing from nominate subspecies only in respect of the broken lateral yellow band across the gena in the male (Figs 102, 103).
MEASUREMENTS
Males Females
Head width (24) 3.3-4.4, 3.71 (29) 3.8-4.4, 4.09
Posterior femur length (24) 7.4-8.9,8.09 (29) 8.1-9.8,9.18
Posterior femur depth (24) 2.1-2.4,2.24 (29) 2.3-2.9,2.58
Body length (24) 13.6-15.5, 14.65 (29) 15.7-18.9, 17.15
MATERIAL EXAMINED
Paratypes. Uganda: 4 9, 1 nymph, Bunyoro, W. of Masindi, Budongo F. Res., 31°30'E, 1°48'N, [25- 27.]viii.l964 (Jago (BMNH); 6 & 29, Buganda, Mabira F. Res., nr Jinja, 33°0'E, 0°25'N, 27.iv.1967 (Jago) (BMNH); 10 £, 169, 7 nymphs, same data as holotype (COPR, London); 1 cJ, 1 ?, Mpanga F. Res., nr Kampala, 30.viii.1969 (E. S. Brown) (BMNH); 1 9, Mpanga F. Res., 1210 m (BMNH); 1 & Bunyoro, Lake Albert, Butiaba, [26-28.] viii. 1964 (Jago) (BMNH); 1 & 3 9, Bunyoro, Bugoma F. Res., S. of Hoima, 31°0'E, 1°15'N, [29-31.] viii. 1964 (Jago) (BMNH; 4 <$, 2 9, Mubende, Mubende reservoir, 19.viii.1964 (Jago) (BMNH).
I samhilla haematogramma sp. n.
(Figs 92, 93, 101, 118, 129, 154-156)
Holotype <$, Tanzania: Ufipa plateau, 25.8 km NNW. of Sumbawanga, Mkundi plantation, [ 1 6-27.] v. 1966 (./ago) (BMNH).
DIFFERENTIAL DIAGNOSIS. Male. Inter-ocular groove and fastigium of vertex wide (Fig. 92), frontal ridge flat. Antennae longer than head and pronotum. Hind femora (Fig. 118) entirely pale green, except knee which is light brown; hind tibiae light green, dark brown in apical half on inner and lower side, spines black-tipped. Supra-anal plate (Fig. 101) with dark lateral infuscate areas and four pairs of simple tubercles. Colour, as for females (Fig. 129), distinctive — body pale emerald green with median dorsal black line above and broad black line on each side from behind compound eye to back of tergite 8 of abdomen. This lateral line demarcated above by a red line extending from front of pronotum to at least rear of tergite 1 of abdomen; demarcated below by a dull yellow band bordered with black ventrally. Cerci conical with narrow pre-apical black annulus. Subgenital plate pale green. Aedeagal valves (Figs 154, 155) gently curved capitad; epiphallic lophi (Fig. 156) apically bifurcate.
Female. Colour as for male (Fig. 129) and in this respect unusual for the genus. Combination of colours unique. Note that mid-dorsal black line may be very faint or absent.
REVISION OF USAMBILLA
37
|
Males (35) 3.3-3.5, 3.37 (35) 7.5-9.2, 8.29 (35) 2.1-2.7, 2.27 (35) 13.9-17.5, 15.18 |
Females (25) 4.1-4.8,4.20 (24) 9.0-11.9, 10.42 (24) 2.9-3.2, 2.88 (24) 17.8-21.7, 19.45 |
MEASUREMENTS
Head width Posterior femur length Posterior femur depth Body length
MATERIAL EXAMINED
Paratypes. Tanzania: 2 cJ, 3 9, 4 nymphs, Ilemba gap, 12.iii.1959 (Vesey-FitzGerald); 4$, 2$, 1 nymph, Nsangu, 2120 m, 13.iii.1959 (Vesey-FitzGerald); 6^, 3 9, Ufipa escarpment, [6-9.] vii. 1948 (Waloff); 2<$, Malonje, Ufipa, 8.iv.l951 (Vesey-FitzGerald); 1 9, Mpui, 100 km N. of Abercorn, 17.vi.1947 (B. P. Uvarov); 3 cJ, 5 9, Ufipa plateau, 25.8 km NNW. of Sumbawanga, Mkundi, [1 6-27.] v. 1966 (Jago); 2 & 69, E. of Sumbawanga, Mbisi F. Res., [23-29.] v. 1966 (Jago), 9 3, 39, 19.3 km E. of Sumbawanga, Malonje Mt, plateau grassland, disused road to Mpui, [24-28.] v. 1966 (Jago); 2<$, 29, Rukwa valley, 8 km W. of Muse, bottom descent Red Locust road, 26.V.1966 (Jago); 10 & 5 9, Mbisi For. Res., v.1966 (Jago);9J, 29, Ufipa, Nsangu F. Res., Sumbawanga-Mpui road, 28. v.1966 (Jago). (Last series COPR, London; rest BMNH.)
DISCUSSION. The species derives its name from the Greek 'haematos' — bloody, 'grammes' — a line, emphasising the distinctive dorso-lateral orange-red line which delineates the upper margin of the black lateral stripe. The general coloration is very similar to that of members of the genus Lentula but the phallic complex shows that U. haematogramma has a strong affinity with the olivacea-group of lentulids.
References
Dirsh, V. M. 1956. Orthoptera, Acridoidea. In Hanstrom, B., Brinck, P. & Rudebeck, G., S. Afr. anim. Life 3:
121-272, 2 pis, 42 figs. Dirsh, V. M. 1968. A new genus and species of the family Lentulidae (Orthoptera: Acridoidea). Proc. R. ent.
Soc. Lond. (B) 37: 143-145, 2 figs. Jago, N. D. 1978. The systematic position of Physocrobylus burtti Dirsh, 1951 with description of the
previously undescribed male sex. Acrida 7: 79-83, 7 figs.
Karsch, F. 1896. Neue Orthopteren aus dem tropischen Afrika. Stettin, ent. Ztg 57: 242-359, 38 figs. Kevan, D. K. McE. 1950. Orthoptera from the hills of south-east Kenya. Jl E. Africa nat. Hist. Soc. 19:
192-221, pi. 34, 5 figs. Kevan, D. K. McE. & Knipper, H. 1961. Geradflugler aus Ostafrika (Orthopteroidea, Dermapteroidea,
Blattopteroidea). Beitr. Ent. II: 356-413, 9 pis, 12 figs. Ramme, W. 1929. Afrikanische Acrididae. Revisionen und Beschreibungen wenig bekannter und neuer
Gattungen und Arten. Mitt. zoo/. Mus. Berl. 15: 247-492, 16 pis, 106 figs. Rehn, J. A. G. 1914. Orthoptera I. Mantidae, Phasmidae, Acrididae, Tettigoniidae und Gryllidae aus dem
Zentral-Afrikanischengebiet Uganda und dem Ituri-Becken des Kongos. Wiss. Ergebn. dt. Zent Afr.-
Exped. 1907-19085(1): 1-223. Sjostedt, Y. 1909. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedtion nach dem Kili-
mandjaro, dem Mem und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905-1906. 17. Orthoptera. 7.
Acridoidea: 149-199, 1 pi., 2 figs.
Uvarov, B. P. 1939. Twenty four new generic names in Orthoptera. Ann. Mag. nat. Hist. (11)3: 457-459. Whellan, J. A. 1975. The Acridoidea of Malawi: an annotated check list. Acrida 4: 105-122.
- 1976. Notes on the genus Mecostibus Karsch, 1896 (Orthoptera: Lentulidae). Arnoldia, (Rhodesia) 8(1):
1-6, 8 figs.
Index
Synonyms are in italics; principal references are in bold.
Acacia 10, 21, 27
Adolfia 20
affinis 21, 22, 23, 24, 25, 27, 28, 29, 30, 32
Albizzia 17
Altiusambilla 1, 3, 12
Aresceutica 13
aspera, Achyranthes 8
Bacteracris 5
Basutacris 4
Betiscoides 5
burtti, Chromousambilla 2, 4, 6, 7, 8, 9
burtti, Physocrobylus 10
Catantopinae 13
chlorophrygana 21, 23, 24, 25, 26, 30, 31, 32, 34
Chromothericles 1
Chromousambilla 1, 3, 5, 16, 20
Combretum 21, 27
Commelina 17
Commiphora 21, 27
cylindricollis 10, 11, 12
Devylderia 4
emaliensis 13, 21, 23, 24, 25, 26, 28, 29, 30 Eremidium 4 Erlangea 8
fractolineata 21, 22, 23, 25, 26, 35, 36 Gymnidium 4
haematogramma 8, 20, 21, 22, 24, 25, 26, 33, 36 Helwigacris 5 hookeri, Pseudarthria 8
insolita 20, 21, 22, 25, 30, 31, 34, 35
Kalaharicus 3
Karruacris 3
Karruia 4
kikomboensis 21, 23, 25, 26, 27, 29, 30
latestriata 2, 4, 5, 6, 7, 8, 9, 10
Lentula 3, 12, 37
leptophrygana 21, 22, 23, 24, 25, 26, 33, 34
magnificum 14, 15, 17, 18, 20 Malawia 1, 3 Mecostiboides 3 Mecostibus 1, 2, 3 Microusambilla 1,' 3, 10, 12 modicicrus 1, 2, 11, 12, 13, 16 montanum 2, 13, 14, 15, 16, 18, 19 mweruensis 2, 4, 5, 6, 7, 8, 9
nguruense 14, 15, 16, 17, 18 Nyassacris 1, 3
olivacea, Usambilla, 20, 23, 24, 25, 26, 27
olivacea, Lentula 37
oraria 21, 22, 24, 25, 26, 28, 32, 35
Paralentula 3 patula, Pinus 1, 13 peruviana, Physalis 8 Plagiotriptus 1
Qachasia 5
Rhainopomma 1, 4, 12, 35
Rehnula 20
robertsoni 2, 4, 6, 7, 8, 9, 10
sagonai 20, 21, 22, 23, 24, 26, 30, 31, 34, 35
Serpusia 13
Shelfordites 3
steppia, Bidens 8
stuhlmanni, Pycnostachys (?) 8
Swaziacris 4
Sygrus 3, 12
turgidicrus 20, 21, 22, 23, 24, 25, 26, 28, 29, 30
usambaricum 13, 14, 15, 16, 17, 18, 19 Usambilla 1, 2, 4, 5, 10, 13, 17, 20
verticillaris, Hypoestes 8 veseyi 2, 4, 5, 6, 7, 8, 9, 10
wapugu 14, 15, 16, 17, 19
38
British Museum (Natural History) 1881-1981
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Titles to be published in Volume 42
A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea) and its allies.
By N. D. Jago.
The Asian, Australasian and Pacific Paraboloponinae (Homoptera : Cicadellidae). A taxonomic revision with a key to all the known genera of the subfamily.
By M. D. Webb.
A revision ofPhyciodes Hiibner and related genera, with a review of the classification of the Melitaeinae (Lepidoptera : Nymphalidae). By L. G. Higgins.
A revision of six minor genera of Myrmicinae (Hymenoptera : Formicidae) in the Ethiopian zoogeographical region. By Barry Bolton.
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HI Q
LIBRARY
Bulletin of the
W LIBR<
British Museum (Natural History)
The Asian, Australasian and Pacific
Paraboloponinae
(Homoptera : Cicadellidae)
A taxonomic revision with a key to all the known genera of the subfamily
M. D. Webb
Entomology series
Vol 43 No 2 24 September 1 98 1
The Bulletin of the British Museum (Natural History), instituted in 1 949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come.
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1981
ISSN 0524-643 1 Entomology series
Vol 43 No 2 pp 39-76 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 24 September 1 98 1
The Asian, Australasian and Pacific Paraboloponinae (Homoptera : Cicadel lidae)
A taxonomic revision with a key to all the known genera of the subfamily
M. D. Webb
i«*
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD
Contents
Synopsis 39
Introduction and historical review 39
Acknowledgements 40
Abbreviations and depositories 40
Paraboloponinae Ishihara 41
Key to the genera of Paraboloponinae 41
Parabolopona Matsumura 42
Key to species of Parabolopona (males only) 43
Favintiga gen. n 47
Dryadomorpha Kirkaldy . 49
Key to Asian, Australasian and Pacific species of Dryadomorpha (males only) . 50
Rhutelorbus gen. n 56
Parohinka gen. n. .
Key to species of Parohinka Males .... Females Karoseefa gen. n. .
Key to species of Karoseefa Oceanopona Linnavuori
References Index .
57 59 59 59 70 71 73 75 76
Synopsis
The subfamily Paraboloponinae is redefined and a key is provided to the nine genera recognized, of which four are new. The Asian, Australasian and Pacific genera are fully revised for these regions, with keys to the 25 species (18 new) recognized. Four new generic synonymies, ten new specific synonymies and six new combinations are established. Seven lectotypes and one neotype are newly designated.
Introduction and historical review
The family Paraboloponidae was erected by Ishihara in 1953 as a subdivision of the Cicadelloi- dea (now Paraboloponinae and Cicadellidae respectively) for the Japanese genus Parabolopona Matsumura, containing two species, P. guttata (Uhler) and P. camphorae Matsumura. In Ishi- hara's description of the subfamily he distinguished it from others in Japan by the cylindrical form, conically produced vertex and long antennae. In 1960 Linnavuori listed the group as a tribe of the Deltocephalinae and described a new genus and species, Oceanopona croceipennis, from the Caroline Is., and in 1974 Eyles & Linnavuori raised the group to subfamily level and included Calotettix Osborn, containing metrosideri Osborn and metrosideri var. tincta Osborn, from the Marquesas Is., and a new species, lais, from the Cook Is. In 1975 Hamilton added the Holarctic genera Stymphylus Stal, containing rubrolineata Stal and modestus Linnavuori (transferred to the
Bull. Br. Mus. not. Hist. (Ent.) 43 (2): 39-76 Issued 24 September 1981
40 M. D. WEBB
Deltocephalinae by Linnavuori in 1978), Yakunopona Ishihara, with one species, yakushimensis Ishihara, from Japan, and Zizyphoides Distant, containing indicus Distant, fraternus Distant and punctatus Rao from India and quadricornis Linnavuori from Africa. In addition Hamilton drew further attention to the long, dorsally situated antennae as a means of identifying the subfamily. In 1978 Linnavuori revised the Paraboloponinae from the Ethiopian region and redefined the group as having falcate anterior tentorial branches, dorsally situated antennae, deep antennal pits delimited by a relatively distinct dorsal ledge and the ocelli not visible from above. He added Dryadomorpha Kirkaldy, containing pallida Kirkaldy, lotophagorum Kirkaldy and viridia Osborn from Australia, Fiji and the Marquesas Is. respectively, and Stenomiella Evans, containing one species, viridis Evans, from Africa; he also described a new genus Odmiella for Stenomiella falcata Linnavuori, from Africa. In addition, the genus Paganalia Distant with one species, virescens Distant, from the Seychelles, was transferred to the subfamily as a senior synonym of Zizyphoides and Rhombopsana Metcalf, the latter a replacement name for Rhombopsis Haupt and containing virens (Haupt) from Palestine and chatterjeei (Singh-Pruthi) and viridis (Singh-Pruthi) from India. The species Z. fraternus was transferred to Stirellus Osborn & Ball (Deltocephalinae), a new species of Paganalia from Africa (anacryon) was described and Platymetopius antennalis Lind- berg, from the Canary Is., was synonymized with Paganalia virens. The latter species was synony- mized with Paganalia virescens by Webb (1980).
Whilst identifying Paraboloponinae from material in the British Museum (Natural History) and elsewhere I discovered four new genera and 18 new species from Asia, Australasia and the Pacific and found that many previous descriptions were inadequate. In addition, the species Parabolopona camphorae, Dryadomorpha lotophagorum and Muirella longiseta Melichar, the latter previously included in the Coelidiinae, were found to belong to other genera. A number of generic and specific synonymies were also discovered.
The subfamily characters given by previous workers are for the most part confirmed by the present study, although of those listed by Linnavuori in 1978 (see above) the antennal ledge is sometimes absent and the ocelli are often visible from above. The male genitalia of the group are similar to those of the Deltocephalinae with Y-shaped connective, but those of Parabolopona are somewhat distinct from the remaining genera in having a long membranous connection between the connective and the aedeagus, the apex of the connective extended posteriorly and the basal apodeme of the aedeagus, in some species, horizontal and compressed dorsoventrally. The genus Favintiga is unusual within the Cicadellids in having a ventroapical process on the connective.
The aim of the present paper is to revise the known genera of Paraboloponinae, other than Odmiella and Stenomiella from Africa (revised by Linnavuori in 1978), to describe new genera and species from the Asian, Australasian and Pacific regions, to redescribe the subfamily and to provide a key to the known genera.
Acknowledgements
For the loan of material in their care I would like to thank the following: Dr P. H. Arnaud, CAS, San Francisco; Dr M. Boulard, MNHN, Paris; Dr O. B. Chhotoni, ZSI, Calcutta; Dr J. Dlabola, NM, Prague; Dr P. H. van Doesburg, RNH, Leiden; Dr K. G. A. Hamilton, CNC, Ontario; Dr Y. Hirashima, ELKU, Fukuoka; Dr J. P. Kramer, USNM, Washington; Dr M. Meinander, ZMU, Helsinki; Dr G. M. Nishida, BPBM, Honolulu; Dr P. K. Sen-Sarma, FRI, Dehra Dun; Dr S. Takagi, EIHU, Sapporo; Dr T. Weir, ANIC, Canberra; Dr K. A. J. Wise, IM, Auckland. I would also like to thank Dr J. Evans for the loan of material in his private collection, Mrs G. Nakahashi and Professor J. Medler for sorting material from the Bishop Museum collection, Dr T. Ishihara and Dr C. A. Viraktamath for their helpful communications and Dr W. J. Knight for the information contained in his unpublished check list on the Pacific Cicadellidae.
Abbreviations of depositories
The specimens studied in the course of this work are deposited in the various institutions and private collections whose names are abbreviated in the text as follows: The South Australian Museum, Adelaide, South Australia (SAM, Adelaide); Auckland Institute and Museum, Auck-
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 41
land, New Zealand (IM, Auckland); Department of Scientific and Industrial Research, Auckland, New Zealand (DSIR, Auckland); Zoological Survey of India, Calcutta, India (ZSI, Calcutta); [Australian National Insect Collection,] C.S.I. R.O., Canberra, Australia (ANIC, Canberra); The Ohio State University, Columbus, Ohio, U.S.A. (OSU, Columbus); Zoologisk Museum, Copen- hagen, Denmark (ZM, Copenhagen); Forest Research Institute, Dehra Dun (U.P.), India (FRI, Dehra Dun); Entomological Laboratory, Kyushu University, Fukuoka, Japan (ELKU, Fu- kuoka); Zoological Museum of the University, Helsinki, Finland (ZMU, Helsinki); Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A. (BPBM, Honolulu); private collection of Professor Dr H. J. Muller, Jena, D.D.R. (HJM, Jena); Rijkusmuseum van Natuurlijke Historic, Leiden, Netherlands (RNH, Leiden); British Museum (Natural History), London, United Kingdom (BMNH, London); [Canadian National Collection,] Ottawa, Ontario, Canada (CNC, Ontario); Museum National d'Historie Naturelle, Paris, France (MNHN, Paris); Narodni Muzeum, Prague, Czechoslovakia (NM, Prague); California Academy of Sciences, San Francisco, Califor- nia, U.S.A. (CAS, San Francisco); Entomological Institute, Hokkaido University, Sapporo, Japan (EIHU, Sapporo); private collection of Dr J. W. Evans, Sydney, Australia (JWE, Sydney); [U.S. National Museum,] National Museum of Natural History, Washington, U.S.A. (USNM, Washington).
PARABOLOPONINAE Ishihara
Paraboloponidae Ishihara, 1953: 20. Type-genus: Parabolopona Matsumura. Paraboloponini; Linnavuori, 1960: 299. Paraboloponinae; Eyles & Linnavuori, 1974: 39.
Yellow, greenish yellow or brownish yellow, often apex of clavus and claval veins with a small brown spot.
Head as wide or wider than pronotum; anterior margin rounded or rim-like, transversely striate, ocelli on margin distant from eyes, anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced with fine longitudinal striations. Face as wide or wider than long, shagreen ; antennae very long, arising near dorsal corners of eyes; antennal pits deep; antennal ledges slight or absent; clypeus with lateral margins constricted near antennae; clypellus elongate, usually with sides concave, rarely with sides parallel; lora large. Pronotum with sides very short to moderately long, with or without a carina; transversely striate, with anterior region rugose or shagreen. Scutellum shagreen or shagreen and obscurely rugose posteriorly. Forewing with three subapical cells, the first subapical cell open and the second and third closed ; subcostal region usually with a few veinlets near to fifth apical cell. Fore tibia with setal arrangement 1 : 4.
Male genitalia with pygophore lobes long with several long spine-like setae. Xth segment short to long, without processes. Valve triangulate. Subgenital plate elongate, triangulate with short to long fine marginal setae dorsally. Connective Y-shaped with stem short to long, arms short. Style with lateral lobe and apical process short to moderately long, with a few sensory papilla and sometimes setae adjacent preapical lobe; basal apophyses weak to strongly developed. Aedeagus usually closely attached to connective; shaft cylin- drical, narrow, tapered to apex, usually symmetrical, processes usually at or near apex sometimes basal; basal apodeme usually vertical, rarely horizontal.
Female genitalia with second valvulae elongate, usually slightly expanded distally with a short to long dorsal sclerotized region, with or without a dorsal prominence.
BIOLOGY. The few recorded host plants of Paraboloponinae are shrubs and small trees. Favintiga camphorae is found on Cinnamomum camphora Nees & Ebermaier, Dryadomorpha pallida on Zizyphus jujuba Miller and Dryadomorpha metrosideri on Glochidion ramiflorum J. R. & G. Forster, Rapanea sp., Reynoldsia sp. and Weinmannia parviflora G. Forster.
DISTRIBUTION. The subfamily is confined to the Old World where it is found mainly in Asia and Australasia but also extends into the Pacific and the Ethiopian region.
Key to the genera of Paraboloponinae
1 Side margins of pronotum carinate, moderately long (Fig. 1). Setal arrangement at apex of hind
femur 2 + 2+1 or 2 + 2 + 0 2
Side margins of pronotum not carinate, short (Fig. 41). Setal arrangement at apex of hind femur
2+1 + 1 or 2+1+0. 4
42 M. D. WEBB
2 Vertex with medial length approximately twice length next to eye. 12-15 setae in fore femur
series (Fig. 3). Setal arrangement at apex of hind femur 2 + 2+ 1. (Asia as far south as Nepal
to the Philippines: Luzon) 3
Vertex with medial length approximately four times length next to eyes. 3-4 setae in fore femur series. Setal arrangement at apex of hind femur 2 + 2 + 0. (Africa) . . ODMIELLA Linnavuori
3 Dorsum yellow or yellow tinged with green. Fore margin of head rim-like (Fig. 5). Vertex
shagreen PARABOLOPONA Matsumura (p. 42)
Dorsum brownish yellow. Fore margin of head rounded (Fig. 33). Vertex finely longitudinally striate . .FAVINTIGA gen. n. (p. 47)
4 Vertex shagreen and obscurely rugose. Setal arrangement at apex of hind femur 2+1+0
OCEANOPONA Linnavuori (p. 73) Vertex longitudinally striate or rugose. Setal arrangement at apex of hind femur 2+1 + 1 . 5
5 Length approximately 10-0 mm. Male pygophore lobes with a long process; subgenital plates
shorter than pygophore. (Africa) STENOMIELLA Evans
Length not exceeding 8-7 mm. Male pygophore lobes without a process; subgenital plates longer than pygophore (Fig. 53). (Africa and Oriental region) 6
6 Clypellus with sides parallel (Fig. 151). 10 setae in fore femur series . KAROSEEFA gen. n. (p. 70) Clypellus with sides concave (Fig. 42). 3-7 setae in fore femur series 7
7 Vertex and pronotum rugose. Lateral margins of face adjacent to eyes visible dorsally (Fig. 68)
RHUTELORBUS gen. n. (p. 56) Vertex and pronotum longitudinally striate. Lateral margins of face not visible dorsally (Fig. 41) 8
8 Vertex without pale patches. Female genitalia with posterior margin of pregenital sternite with a
small protuberance each side of midline (Fig. 51); dorsal margin of second valvulae with an anterior prominence (Fig. 56). Male genitalia with aedeagus symmetrical
DRYADOMORPHA Kirkaldy (p. 49)
Vertex with or without pale patches. Female genitalia with posterior margin of pregenital sternite without a protuberance each side of midline; dorsal margin of second valvulae without an anterior prominence (Fig. 133). Male genitalia with aedeagus asymmetrical
PAROHINKA gen. n. (p. 51)
PARABOLOPONA Matsumura
Parabolopona Matsumura 1912: 288. Type-species : Parabolocratus guttatus Uhler, by original designation.
Yellow to greenish yellow ; forewings with a small brown spot on apex of clavus, on apex of veins of clavus and apical cells and a variable brown spot on first m-cu cross vein and base of inner vein of second subapical cell.
Head as wide as pronotum; anterior margin rim-like, carinate; ocelli on margin, distant from eyes, visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approximately twice length next to eyes; sides slightly convex or slightly angularly rounded; apex narrowly angularly rounded; shagreen, transversely striate anteriorly. Face slightly wider than long, shagreen; upper margin depressed medially with a few transverse striations; face in profile more or less straight ; clypeus moderately long and narrow, lateral margins constricted near antennae; clypellus elongate, expanded apically ; transclypeal suture visible ; lora large ; antennal pit deep with inner margin more or less angularly rounded to clypeus; antennal ledge slight; antennae very long, when recurved extending to near apex of clavus. Pronotum approximately twice as wide as long, side margins moderately long, carinate; irregularly and transversely striate, shagreen anteriorly. Scutellum approximately equal in length to prono- tum, shagreen, obscurely rugose posteriorly. Fore wing with three subapical cells, first subapical cell open, second and third subapical cells closed. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of 12-14 fine setae distally on anterior surface; hind femur with apical setal formula 2 + 2+1 with the proximal and more dorsal of the middle setae slightly narrower than others.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore straight in dorsal aspect, with or without an apodeme on each side; pygophore lobes with several macrosetae and numerous short fine setae. Xth segment moder- ately long, cylindrical. Valve triangular. Subgenital plate moderately long, triangular, apical region digitate and lightly sclerotized; ventral surface with several short setae; outer margin of dorsal surface with a few moderately long fine setae on basal lobe and usually towards apex of plate. Style moderately long with basal apophysis and lateral lobe prominent; apical process moderately long, curved ventrally and tapered to apex or with apex foot-like, crenulate dorsally; region adjacent preapical lobe with a few sensory papilla ventrally
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 43
and a few short fine setae dorsally. Connective Y-shaped with stem produced posteriorly, not articulated with aedeagus at apex but with a long membranous connection from approximately midlength of stem; arms short. Aedeagus with shaft straight or curved posteroventrally, short with a pair of apical processes; gonopore situated at apex on posterior surface; basal apodeme large, either vertical and compressed anteroposteriorly or horizontal around base of shaft and compressed dorsoventrally.
Female genitalia with second valvulae united at midlength (arrowed in Fig. 14) and slightly expanded distally, without a basal prominence; dorsal teeth very fine, extending approximately one-quarter distance from apex to base of valvulae; dorsal sclerotized region short to long.
REMARKS. This genus is similar to Favintiga with the lateral margins of the pronotum moderately long and carinate and the hind femur with a setal formula 2 + 2 + 1 ; it differs, however, in the more rim-like fore margin of the head and the shagreen rather than longitudinally striate vertex. In the male genitalia the horizontal and dorsoventrally compressed basal apodeme of the aedeagus in some species, the produced stem of the connective and the distant relationship of the aedeagus and connective to each other are unique within the subfamily.
DISTRIBUTION. Asia as far south as Nepal to the Philippines (Luzon).
Key to species of Parabolopona (males only)
1 Aedeagus with a pair of apical processes; stem of connective without setae 2
Aedeagus with a pair of basal processes; stem of connective with setae. (Philippines)
luzonensis sp. n. (p. 46)
2 Aedeagus with apical processes directed away from base of shaft in lateral aspect, gonopore small 3 Aedeagus with apical processes directed towards base of shaft in lateral aspect; gonopore large
(Figs 16, 17) ishihari sp. n. (p. 45)
3 Aedeagal shaft strongly curved (Fig. 8); connective with apex upturned and expanded laterally
(Figs 7, 13). (Japan and Taiwan) guttata (Uhler) (p. 43)
Aedeagal shaft weakly curved (Fig. 20); connective with apex straight and narrow. (China)
chinensis sp. n. (p. 45)
Parabolopona guttata (Uhler) (Figs 1-14)
Parabolocratus guttatus Uhler, 1896: 291. LECTOTYPE & JAPAN (USNM, Washington), here designated
[examined]. Parabolopona guttata (Uhler) Matsumura, 1912: 288.
Length: cJ, 6-6-7-0 mm, mean 6-8 mm; $, 7-0-8-0 mm, mean 7-5 mm.
Colour and external characters as in generic description.
Male genitalia with pygophore lobes angularly rounded posteriorly; anterior margin of pygophore with a prominent apodeme on each side. Connective with apex upturned and expanded laterally. Style with apical process tapered to apex. Aedeagus with shaft elongate, curved posteroventrally and continued ventrally as a pair of elongate diverging processes; gonopore small; basal apodeme horizontal, compressed dorsoventrally around base of shaft (Fig. 12).
Female genitalia with posterior margin of pregenital sternite sinuate; second valvulae with dorsal scler- otized region short.
REMARKS. The male genitalia of guttata are similar to those of chinensis but the pygophore lobes are broader, the apex of the connective is upturned and expanded laterally and the aedeagal shaft is strongly curved with the apical processes expanded subapically without a small lateral pro- tuberance basally.
DISTRIBUTION. Japan and Taiwan.
MATERIAL EXAMINED
Parabolocratus guttatus Uhler, lectotype cJ, Japan: Gifuyama, 7.vii.l888 [in Japanese] (USNM, Wash- ington).
Japan: numerous specimens from Honshu and Kyushu (BMNH, London; EIHU, Sapporo; ELKU, Fukuoka); 1 $, Gifuyama, 7.vii.l888 [in Japanese] (USNM, Washington) (paralectotypes of Parabolocratus guttatus Uhler). Taiwan: 1 <J, 1 ?, Tattaka, 16, 19.viii.1921; 3 9, Hassenzan, Taichu-shu, Reimei, 13- 14.vii.1932 (ELKU, Fukuoka).
44
M. D. WEBB
Figs 1-14 Parabolopona guttata. 1, head and thorax, dorsal view; 2, face; 3, left fore leg, anterior view; 4, left anterior tentorial branch, lateral view; 5, head and thorax, lateral view; 6, $ pregenital segments, ventral view; 7, connective, lateral view; 8, aedeagus, lateral view; 9, left style, ventral view; 10, cJ genital capsule, lateral view; 11, aedeagus, dorsal view; 12, left subgenital plate and style and connective, dorsal view; 13, connective, dorsal view; 14, second valvulae, lateral view.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
45
Parabolopona ishihari sp. n.
(Figs 15-19)
Length: ^,6-6 mm; $,7-5 mm.
Colour and external characters as in generic description.
Male genitalia with pygophore lobes acute posteriorly; anterior margin without an apodeme on each side. Connective with basal stem straight and narrow throughout length. Style with apical process tapered to apex. Aedeagus with shaft short and robust, more or less straight, terminating in a pair of moderately long ventrally directed processes; gonopore large; basal apodeme horizontal, compressed dorsoventrally around base of shaft (Figs 16,17).
Female genitalia as in guttata.
REMARKS. The male genitalia of ishihari are similar to those of guttata but the pygophore lobes are more acute posteriorly, the apex of the connective is straight and narrow and the aedeagus has the processes directed towards the base of the shaft in lateral aspect and the gonopore large.
DISTRIBUTION. Japan.
MATERIAL EXAMINED
Holotype J, Japan: Northern Honshu, Towada, vii.1905 (EIHU, Sapporo). Paratypes. Japan: 1 <$, 1 $, same data as holotype (BMNH, London; EIHU, Sapporo).
Parabolopona chinensis sp. n.
(Figs 20-21)
Length : <J, 6-4 mm.
Colour and external characters as in generic description.
Male genitalia with pygophore and connective similar to ishihari (Figs 15, 18) but pygophore with an apodeme on each side anteriorly and posterior lobes less acute; remaining structures similar to guttata but
15
20
Figs 15-21 Parabolopona species. 15-19. P. ishihari. (15) ^ pygophore, lateral view; (16, 17) aedeagus, lateral and posterior views; (18, 19) connective, dorsal and lateral views. 20, 21, P. chinensis. (20) aedeagus, lateral view; (21) apex of aedeagus, dorsal view.
46
M. D. WEBB
connective straight and narrow apically and aedeagal shaft only slightly curved with apical processes evenly tapered from base to apex with a small lateral protuberance basally. Female genitalia unknown.
REMARKS. This species is similar to guttata but differs in the male genitalia as noted above. From ishihari with a similarly shaped pygophore and connective (see above), chinensis differs in having the aedeagal processes directed away from the shaft in lateral aspect.
DISTRIBUTION. Central China.
MATERIAL EXAMINED
Holotype cJ, China: Hubei-Sichuan border, trail between Mo-Tai-Chi and Sang-Hou-Ken, 19.vii.1948 (Gressitt & Djou) (CAS, San Francisco).
Parabolopona luzonensis sp. n.
(Figs 22-27) [Parabolocratus guttatus Uhler; Merino, 1936: 364. Misidentification.]
Length : (J, 6-7-7-3 mm, mean 7-0 mm; 9, 7-3-8-2 mm, mean 8-0 mm.
Colour and external characters as in generic description.
Male genitalia with pygophore lobes narrowly rounded posteriorly; anterior margin of pygophore with- out apodemes. Connective with stem nearly straight, tapered to acute apex; distal region with numerous short stout setae dorsally. Style with apical process foot-like apically in lateral aspect. Aedeagus with shaft short and robust, curved dorsally, apex with a slight lamellate expansion arising from anterior margin on each side; a pair of moderately long, dorsally directed processes, arising basally from posterior margin; gonopore moderately large ; basal apodeme vertical, compressed anteroposteriorly.
Female genitalia with posterior margin of pregenital sternite produced medially; second valvulae with dorsal sclerotized region elongate.
23
27
Figs 22-27 Parabolopona luzonensis. 22, $ genital capsule, lateral view; 23, 9 pregenital sternite; 24, connective, dorsal view; 25, 26, aedeagus, lateral and posterior views; 27, apex of style, lateral view.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 47
REMARKS. This species can be distinguished from other members of the genus by the foot-like apex of the style and the aedeagus with the processes arising basally rather than apically and the basal apodeme being vertical rather than horizontal. The female genitalia can be distinguished by the medially produced posterior margin of the pregenital sternite and the elongate dorsal scler- otized region of the second valvulae.
DISTRIBUTION. Philippines (Luzon).
MATERIAL EXAMINED
Holotype J, Philippines: Luzon, Baguio, Benguet (Baker) (USNM, Washington).
Paratypes. Philippines: 10 £, 19 9, same data as holotype (1 (J, 1 $ in BMNH, London; remainder in USNM, Washington); 1 <J, Luzon, Heights Plane (OSU, Columbus).
FA VINT1GA gen. n.
Type-species: Parabolopona camphor -ae Matsumura, 1912.
Brownish yellow dorsally, pale yellow ventrally; fore wings with a small brown spot near mid length of subcostal region, on apex of clavus and on apex of veins of clavus, apical cells and additional vein in subcostal region.
Head as wide as pronotum; anterior margin angularly rounded in profile, transversely striate, becoming carinate and rim-like medially; ocelli on margin, distant from eyes, visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approximately twice length next to eyes, sides slightly convex; apex moderately broadly rounded; finely longitudinally striate, transversely striate anteriorly. Face slightly wider than long, more or less straight in profile; shagreen; clypeus moderately long, narrow, lateral margins constricted near antennae; clypellus elongate, expanded apically; transclypeal suture visible; lora large; antennal pit deep with inner margin obliquely inclined to clypeus; antennal ledge slight; antennae very long, extending to near apex of clavus. Pronotum approxi- mately twice as wide as long, sides moderately long, carinate ; finely and transversely striate, shagreen and more or less smooth anteriorly, Scutellum approximately equal in length to pronotum, shagreen, obscurely rugose posteriorly. Fore wings with three subapical cells, first subapical cell open, second and third subapi- cal cells closed; an additional veinlet in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of 15 fine setae distally on anterior surface; hind femur with apical setal formula 2+1 + 1 with the proximal and more dorsal of the middle setae slightly narrower than others.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes with several macrosetae and short fine setae. Xth segment moderately long, cylindrical. Valve triangulate. Subgenital plates moderately long, distal half digitate and lightly sclerotized; dorsal surface with a few moderatly long fine setae on basal lobe and a few very short setae distally, Style moderately long with basal apophyses and lateral lobe prominent; apical process moderately long, curved ventrally and tapered api- cally, crenulate medially; a few sensory papilla medially adjacent lateral lobe. Connective Y-shaped; stem long, lateral margins keel-like dorsally; a bifurcate process distally on ventral surface; arms short. Aedeagus large, laterally compressed with an elongate preatrium; shaft curved dorsally and tapered apically with a pair of basal processes; gonopore situated at apex on posterior surface, elongate; basal apodeme elongate.
Female genitalia with second valvulae united at first dorsal tooth (arrowed in Fig. 38), narrow throughout length in lateral aspect with a slight dorsoanterior prominence; dorsal teeth very fine, extended over approximately distal third of valvulae; dorsal sclerified region elongate.
REMARKS. This genus is similar externally to Parabolopona (see remarks under that genus) but the long preatrium of the aedeagus and the processes on the connective are unique within the subfamily.
DISTRIBUTION. Japan and Amama-Oshima I. (south of Japan).
48
M. D. WEBB
Figs 28-40 Favintiga camphor ae. 28, head and thorax, dorsal view; 29, face; 30, 9 pregenital segments, ventral view; 31, fore wing; 32, left style, ventral view; 33, head and thorax, lateral view; 34, left subgenital plate and apex of left style, dorsal view; 35, 36, aedeagus, lateral and posterior views; 31,3 genital capsule, lateral view; 38, second valvulae, lateral view; 39, 40, connective, lateral and dorsal views.
Favintiga camphorae (Matsumura) comb. n. (Figs 28^0)
Parabolopona camphorae Matsumura, 1912: 288. LECTOTYPE 9, JAPAN (EIHU, Sapporo), here designated [examined].
Length: <$, 6 mm; 9, 6-3-7-0 mm, mean 6-6 mm.
Colour and external characters as in generic description.
Male genitalia as in generic description with apical process of style with a small subapical tooth on medial surface and connective with each branch of ventral process elongate, tapered to apex and curved ventrome- dially. Aedeagus with shaft elongate; processes arising against posterior margin on each side, extended dorsally close to shaft and terminating a little before its apex, evenly tapered from base to apex.
Female genitalia with posterior margin of pregenital sternite convex; second valvulae as in generic description.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 49
REMARKS. This species is distinguishable by its brownish yellow colour dorsally and its distinctive male genitalia as noted above.
DISTRIBUTION. Japan and Amami-Oshima I. (south of Japan).
MATERIAL EXAMINED
Parabolopona camphorae Matsumura, lectotype 9, Japan: S. Kyushu, Kagoshima, 10.vii.1903 (EIHU, Sapporo).
Japan: 1 $, same data as lectotype (paralectotype of Parabolopona camphorae Matsumura); !<£,!$, Honshu, Gifuyama, 7.vii.l888 (USNM, Washington); 3 9, Amami-Oshima I., Yakkachi, Sumiyo-mura, 17-18.vii.1933 (Esaki & Yasumatsu) (ELKU, Fukuoka, BMNH, London).
DRYADOMORPHA Kirkaldy
Dryadomorpha Kirkaldy, 1906: 335. Type-species: Dryadomorpha pallida Kirkaldy, by monotypy.
Paganalia Distant, 1917: 314. Type-species : Paganalia virescens Distant, by monotypy. Syn. n.
Zizyphoides Distant, 1918: 73. Type-species: Zizyphoides indicus Distant, by original designation. [Synony- mized by Linnavuori 1978: 459.]
Rhombopsis Haupt, 1927: 22. Type-species: Rhombopsis virens Haupt, by monotypy. [Homonym of Rhom- bopsis Gardner, 1916: 456.]
Calotettix Osborn, 1934: 247. Type-species: Calotettix metrosideri Osborn, by original designation. [Hom- onym of Calotettix Bruner, 1908: 309.]
Yakunopona Ishihara, 1954: 12. Type-species: Yakunopona yakushimensis Ishihara, by original designation. Syn. n.
Rhombopsana Metcalf, 1967: 229. [Replacement name for Rhombopsis Haupt.] [Synonymized by Linna- vuori, 1978: 459.]
Osbornitettix Metcalf, 1967: 229. [Replacement name for Calotettix Osborn.] Syn. n.
Khamiria Dlabola, 1979: 252. Type-species: Khamiria mangrovecola Dlabola, by original designation. Syn. n.
Yellow, greenish yellow or stramineous; fore wings with either a small brown spot at apex of both clavus and claval veins or inner margin of clavus bordered with brown. Legs spotted with brown.
Head wider than pronotum; anterior margin angularly or acutely rounded in profile, transversely striate, sometimes becoming carinate and rim-like in longer headed forms; ocelli on margin distant from eyes, not or slightly visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangu- larly produced, medial length 1-5-3-0 times length next to eyes, sides slightly convex to concave, apex fairly broadly angularly rounded to acute in longer headed forms; with a longitudinal depression; finely longitudi- nally striate, transversely striate anteriorly. Face elongate to only slightly longer than wide, shagreen; upper margin slightly depressed each side of mid line in longer headed forms forming a faintly striate longitudinal keel medially; face in profile convex to more or less straight, concave anteriorly in longer headed forms; clypeus elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; trans- clypeal suture distinct or indistinct; lora large; antennal pit deep with inner margin angularly rounded to clypeus, sometimes faintly rim-like and nearly carinate; antennal ledge very slight; antennae very long, extending to beyond apex of clavus when recurved. Pronotum approximately twice as wide as long, sides very short, without a carina; finely and transversely striate, obscurely rugose anteriorly. Scutellum approxi- mately equal in length to pronotum, shagreen with posterior region obscurely rugose. Fore wing with three subapical cells, first subapical cell open, second and third subapical cells closed; one or two additional veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of seven setae distally on anterior surface ; hind femur with apical setal formula 2+1-1-1 with first proximal seta slender.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes with an oblique internal ledge terminating at ventroposterior margin with a darkly pigmented area, pygo- phore lobes with several macrosetae and short to moderately long fine setae. Xth segment moderately long, compressed dorsoventrally. Valve triangulate. Subgenital plates elongate, triangular; ventral surface of lateral lobe with a more heavily sclerotized region apically; outer margin with numerous long fine setae on dorsal and ventral surfaces; apex with a few short stout setae. Style moderately long with basal apophyses prominent ; apical process moderately long, curved ventrally and tapered to acute or rounded apex, crenu- late distally; a few sensory papilla ventrally, adjacent lateral lobe. Connective Y-shaped, stem short to moderately long with lateral margins keel-like dorsally; arms short. Aedeagus with shaft elongate, curved
50 M. D. WEBB
dorsally and tapered to apex with two or four apical, dorsally directed processes ; gonopore small, apical on posterior surface; basal apodeme moderately long and narrow.
Female genitalia with posterior margin of pregenital sternite with a small protuberance each side of mid line; second valvulae united at first dorsal tooth (arrowed in Fig. 56), slightly expanded distally, fairly robust with a dorsoanterior prominence; dorsal teeth robust, unaligned, extending to near mid length of valvulae; dorsal sclerotized region moderately long.
REMARKS. This genus is represented in the southern Palaearctic region by Dryadomorpha pallida Kirkaldy (also present in Africa, Asia and Australasia and described below), and D. mangrovecola (Dlabola) comb. n. Also present in Africa are D. anacryon (Linnavuori) comb. n. and D. quad- ricornis (Linnavuori) comb. n. (both species adequately described by Linnavuori, 1978: 462). Two species incorrectly described in Zizyphoides ( = Dryadomorpha) are Z. fraternus Distant belonging to the genus Stirellus Osborn & Ball (Linnavuori, 1978: 460) and Z. punctatus Rao syn. n. of Mahalana lugubris Distant.
Externally Dryadomorpha is almost identical to Rhutelorbus and Parohinka but differs in lacking the pale patches on the head (usually present in Parohinka and sometimes in Rhutel- orbus), the striate vertex and pronotum (rugose in Rhutelorbus), the lateral margins of the face adjacent to the eyes not being visible dorsally (just visible in Rhutelorbus) and the posterior margin of the female pregenital sternite being transverse (posterior corners produced in Rhutel- orbus and Parohinka or broadly V- or U-shaped in Parohinka). The male genitalia are similar to Karoseefa but have the anterior margin of the pygophore straight dorsally and without apo- demes. In the female genitalia the second valvulae are similar to those of Parohinka but with a dorsoanterior prominence present.
DISTRIBUTION. Ethiopian and southern Palaearctic region, Asia, Australasia and the Pacific.
Key to Asian, Australasian and Pacific species of Dryadomorpha (males only)
1 Aedeagus with apical processes strongly divergent (Fig. 52) 2
Aedeagus with apical processes weakly divergent (Fig. 62) 3
2 Aedeagal processes slightly expanded at mid length in posterior aspect (Fig. 52); fore wings with a
small brown spot at apex of both clavus and claval veins (Fig. 47). (Ethiopian, southern Palaearctic and Oriental regions as far south as Singapore and Java and northern and
north-eastern Australia) pallida Kirkaldy (p. 50)
Aedeagal processes narrow throughout length in posterior aspect (Fig. 60); fore wings without a small brown spot at apex of both clavus and claval veins, sometimes inner margin of clavus brown (Fig. 58). (Marquesas Is., Cook Is. and northern Australia) . . metrosideri (Osborn) (p. 53)
3 Aedeagal shaft robust (Fig. 63); style with apical process tapered to apex (Fig. 61). (Malaya,
Borneo (Sarawak and Sabah)) robustipenis sp. n. (p. 54)
Aedeagal shaft slender (Fig. 66); style with apical process rounded or foot-like apically (Figs 64,
65). (Borneo (Sarawak and Sabah)), New Guinea and Vanuatu . . . pacifica sp. n. (p. 55)
Dryadomorpha pallida Kirkaldy (Figs 41-56)
Dryadomorpha pallida Kirkaldy, 1904: 336. Holotype?, AUSTRALIA, (BPBM, Honolulu) [examined].
Paganalia virescens Distant, 1917: 314. Lectotype $, SEYCHELLES (BMNH, London), designated by Webb (1980: 848) [examined]. Syn. n.
Zizyphoides indicus Distant, 1918: 73. LECTOTYPE ?, INDIA (BMNH, London), here designated [exam- ined]. Syn. n.
Rhombopsis virens Haupt, 1927: 23. Lectotype cJ, ISRAEL (HIM, Jena), designated by Webb (1980: 848) [examined]. Syn. n.
Rhombopsis viridis Singh-Pruthi, 1930: 34. LECTOTYPE <$, INDIA (ZSI, Calcutta), here designated [exam- ined]. Syn. n.
Rhombopsis chatterjeei Singh-Pruthi, 1934: 26. LECTOTYPE <3, INDIA (FRI, Dehra Dun), here designated [examined]. Syn. n.
Platymetopius antennalis Lindberg, 1958: 181. Holotype <$, CAPE VERDE Is. (ZMU, Helsinki) [examined]. Syn. n.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
51
Figs 41-56 Dryadomorpha pallida. 41, head and thorax, dorsal view; 42, face; 43, head and thorax, lateral view; 44, head and pronotum, dorsal view; 45, head and thorax, lateral view; 46, head and pronotum, dorsal view; 47, fore wing; 48, face; 49, aedeagus, lateral view; 50, left style, ventral view; 51, apex of $ abdomen, ventral view; 52, aedeagus, posterior view; 53, <$ genital capsule, lateral view; 54, 55, connective, lateral and dorsal views; 56, second valvulae, lateral view.
52 M. D. WEBB
Yakunopona yakushimensis Ishihara, 1954: 13. Holotype 9, JAPAN (ELKU.Fukuoka) [examined]. Syn. n. Platymetopius australis Evans, 1966: 247. Holotype £, AUSTRALIA (ANIC, Canberra) [examined]. Syn. n.
Length : J, 4-5-5-6 mm, mean 5- 1 mm ; 9, 5-3-6-6 mm, mean, 6-0 mm.
Colour as in generic description with rostrum and sometimes apex of clypellus scarlet; head and thorax rarely with testaceous marking (see 'Remarks' below), sometimes marked with stramineous or orange at apex of vertex and on each side at base of vertex, on lateral margins of clypeus and on anterior region of pronotum. Fore wing variably tinged with brown distally; apex of both clavus and claval veins with a small brown spot.
External characters as in generic description with vertex short to long, medial length 1-7-3-5 times length next to eyes; apex acute, narrowly rounded or fairly broadly angularly rounded, sides concave to slightly convex (see 'Remarks' below).
Male genitalia as in generic description with pygophore variable in shape (see 'Remarks' below). Style with apical process evenly tapered to acute apex, lateral lobe fairly strongly developed. Connective short. Aedeagus with shaft slender, apical processes slightly expanded at mid length, strongly divergent.
Female genitalia as in generic description.
REMARKS. Some specimens from India have the markings on the head and pronotum orange (see description) while one specimen from Australia has the head and thorax with testaceous mar- kings. Considerable variation occurs in the shape of the head; usually the side margins of the vertex are nearly straight but a few specimens from India have the side margins considerably concave (Fig. 46). The face in profile is usually slightly convex to nearly straight but in one specimen from Australia the face is strongly concave. In the male genitalia slight variation occurs in the pygophore in the shape of the posterior lobes, the distal darkly pigmented region of the internal ledge and in the number of macrosetae (compare Fig. 53 from Java with Webb, 1980: 849, fig. 115 from Aldabra).
DISTRIBUTION. Afrotropical, southern Palaearctic and Oriental regions as far south as Singapore and Java (excluding Malaya and Sumatra) and from northern and north-eastern Australia.
MATERIAL EXAMINED
Dryadomorpha pallida Kirkaldy, holotype 9, Australia: Queensland, Bundaberg, ix-xii.1904 (Koebele) (BPBM, Honolulu). Paganalia virescens Distant, lectotype 9, Seychelles: Silhouette, 1908 (BMNH, London). Zizyphoides indicus Distant, lectotype 9, India: Calcutta, on Zizyphus jujuba Miller, 23.vii.1912 (BMNH, London). Rhombopsis virens Haupt, lectotype <$, Israel: 'Palestine', Ben-Shemen, 17- 18.x. 1925 (HIM, Jena). Rhombopsis viridis Singh-Pruthi, lectotype £, India: Punjab, Lyallpur, x.1929 (Rahman) (ZSI, Calcutta). Rhombopsis chatterjeei Singh-Pruthi, lectotype $, India: North Salem, Uduparani, on unspiked sandel, 2911930 (Chatterjee) (FRI, Dehra Dun). Platymetopius antennalis Lindberg, holotype^, Cape Verde Is.: Fogo I., supra Fte Aleixo, 19.ii.1954 (Lindberg) (ZMU, Helsinki). Yakunopona yakushimensis Ishihara, holotype 9, Japan: Yakushima, Anbo, 25.viii.1952 (Takeya & Hiroshima} (ELKU, Fukuoka). Platymetopius australis Evans, holotype <$, Australia: NW., Kimberley Research Station, via Wyndham, 23.viii. 1956 (Langfield) (ANIC, Canberra).
India: l<$, 1 9, Punjab, Lyallpur, x.1929, 7.X.1929 (ZSI, Calcutta) (paralectotypes of Rhombopsis viridis Singh-Pruthi); 1 £, North Salem, Jawalagiri, 22.xii. 1930 (ZSI, Calcutta) (paralectotypes of Rhombopsis chatterjeei Singh-Pruthi); 1 9, Coorg, 9.H.1930 (ZSI, Calcutta) (paralectotype of Rhombopsis chaterjeei Singh-Pruthi); 3 <J, 3 9, Dehli, Indian Agricultural Research Institute, at light; 1 <J, Bihar, Pusa, 14.vi.1931 ; 1 <J, 6 9, Ludhiana, 25.xi.1976 (all in BMNH, London). Nepal: 3 $, 3 9, near Simra, Abhabbar, 180 m, 25-27.viii.1967; 1 9, near Birganj, Lothor, 135 m, 3.ix.l967 (all in CNC, Ontario); 1 9, Kathmandu, 1300-1400 m, 7-12.V.1966 (BPBM, Honolulu); 2 9, Arun Valley, below Tumlingtar, river Sabhaya, west shore, 460 m, 22.xii.1961 (BMNH, London). Sri Lanka: 1 <J, Amiradhapura, at light, 22.iii.1953 (BMNH, London). Bangladesh: 1 <£, Lawa Chera forest, Srimangal, 110 m, 27.ix.1961 (CAS, San Francisco). China: 1 9, Macao, i.1907; 1 & 39, Fujian, Yungan, 4.viii-24.ix.l940(all in BPBM, Honolulu). Japan: 1 & 29, Tokara I., Nakanoshima, 25.v-13.vi.1953; 1 <$, Kyushu, Satamisaki, Osumi, 30.viii.1951 (all in ELKU, Fukuoka). Hong Kong: 1 ^, Lantau I., Shek Pik Reservoir area, 21.vii.1964; numerous examples, Sai Kung Station, 30.i.l965; 9^, 89, Taipokau, 20.vi.-6.vii.1964 (all in BPBM, Honolulu). Taiwan: 1 & 19, Taipei Hsien Santiaoling, 100-400 m, 19.xi.1957 (BPBM, Honolulu). Thailand: 1 9, South Banna, Nakhon, 108 m, 5-10.V.1958; 1 <J, Bangkok, at light, 4.xii.l957; 1 9, Chiangmai, Fang, 12-19.iv.1958 (all in BPBM, Hono- lulu). Laos: 1 9, Vientiane, at light, S.v.1965 (BPBM, Honolulu). Philippines: 2 £, 19, Busuanga I., 4 km north of San Nicolas, at light, 21-24.V.1962, 1 £, Palawan, 3 km north-east of Tinabog, at light, 14.V.1962; 1 9, Palawan, Tarumpitao Point, at light, 26.V.1958; 1 9, Culion I., 6 km west of Culion, at light, 7.vi.l962 (all
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
53
in BPBM, Honolulu). Singapore: 4 $, 1 ?, Mandai, mangrove, at light (BMNH, London). Java: 1$, Bogor, at light, v.1954 (BMNH, London). Australia: 1 9, Western Australia, Kimberley Research Station, via Wyndham, 23.viii.1956 (ANIC, Canberra) (paratype of Platymetopius australis Evans); 1 9, New South Wales, Sydney (BMNH, London).
Dryadomorpha metrosideri (Osborn) comb. n. (Figs 57-60)
Calotettix metrosideri Osborn, 1934: 247. Holotype9, MARQUESAS Is. (BPBM, Honolulu) [examined]. Calotettix metrosideri var. tincta Osborn, 1934: 248. LECTOTYPE $, MARQUESAS Is. (BPBM, Honolulu),
here designated [examined]. Syn. n. Calotettix lais Eyles & Linnavuori, 1974: 40. Holotype^, COOK Is. (IM, Auckland) [examined]. Syn. n.
Length: $, Marquesas Is., 6-3-6-7 mm, mean 6-5 mm; Cook Is., 6-8-7-4 mm, mean 7-2 mm; Australia, 5-5 mm; 9, Marquesas Is., 7-0-7-7 mm, mean 7-2 mm; Cook Is., 7-8-8-5 mm, mean 8-1 mm.
Yellow or yellow tinged with green or orange; darker specimens with additional reddish and dark brown markings on pronotum and scutellum ; some specimens (see 'Remarks' below) have face reddish, fore wings either marked with brown on inner margin of clavus and with a brown spot at base of fourth apical cell, or marked with brown bordering veins of clavus and veins cu and m and with a brown patch at apex of wing, and female pygophore with a dark brown patch posteriorly on each side.
Figs 57-67 Dryadomorpha species. 57-60. D. metrosideri. (57) head and thorax, dorsal view; (58) fore wing; (59, 60) aedeagus, lateral and posterior views. 61-63. D. robustipenis. (61) apex of left style, ventral view; (62, 63) aedeagus, posterior and lateral views. 64-67. D. pacifica. (64) apex of the left style, ventral view, Sarawak; (65) same, New Guinea; (66) aedeagus, lateral view; (67) connective, dorsal view.
54 M. D. WEBB
External characters as in generic description with vertex short, approximately 1-5 times as long medially as next to eyes, sides slightly convex, apex fairly broadly angularly rounded. Male genitalia as in pallida but aedeagal processes in posterior aspect narrow throughout length. Female genitalia as in pallida.
REMARKS. In addition to the differences in size noted above specimens from different localities vary in the following ways: the single specimen from Australia has a medial longitudinal keel dorsally on the face (absent in other specimens), the clypeus reddish, some veins of the fore wing bordered with brown, and a brown patch at the apex of the fore wing; specimens from the Marquesas Is. have the inner margin of the fore wing variably marked with brown (Fig. 58) and the face sometimes marked with red, the thorax marked with red or brown and the female pygophore marked with brown posteriorly; the specimens from the Cook Is. are without mar- kings.
This species can be distinguished from other members of the genus by its short head. The male genitalia are almost identical to those of pallida, differing only in the shape of the aedeagal processes as noted above.
DISTRIBUTION. Marquesas Is., Cook Is. and northern Australia.
MATERIAL EXAMINED
Calotettix metrosideri Osborn, holotype 9, Marquesas Is.: Hiva Oa, Kopoa faa, miscel. sweeping, 831 m, 3.viii.l929 (Mwnford & Adamson) (BPBM, Honolulu). Calotettix metrosideri var. tincta Osborn, lectotype$, Marquesas Is.: Hiva Oa, Feani Ridge, 1 140 m, 22.U932 (Le Bronnec) (OSU, Columbus). Calotettix lais Eyles & Linnavuori, holotype <$, Cook Is.: Rarotonga, 15.xii.1937 (IM, Auckland).
Marquesas Is.: numerous specimens from Hiva Oa, 630-1140 m, on Weinmannia parviflora G. Forster, Reynoldsia sp., Glochidion ramiflorwn G. Forster or Rapanea sp., I.viii-5.i.l932 (Le Bronnec or Mumford & Adamson) (paratypes of Calotettix metrosideri Osborn and paralectotypes of Calotettix metrosideri var. tincta Osborn) (BPBM, Honolulu; OSU, Columbus). Cook Is. 3 & 19, Rarotonga I., Papua Creek, Vaimaanga, at light, 1 3.x. 1975; 7 c?, 39, Rarotonga I., Avatu Valley, at light, x. 1975; 2^, 1 9, Rarotonga I., Totokoitu Ridge, 210 m, 19.x. 1975; 1 £, Rarotonga I., Totokoitu, at light, 14.x. 1975 (all in DSIR, Auckland and BMNH, London). Australia : 1 <J, Northern Territory, Darwin (JWE, Sydney).
Dryadomorpha viridia Osborn Dryadomorpha viridia Osborn, 1934: 244. Holotype?, MARQUESAS Is. (BPBM, Honolulu) [examined].
Length: 9, 6- 7 mm.
Colour as in generic description with inner margin of clavus brown.
External characters as in generic description with vertex moderately long, medial length approximately twice as long as length next to eyes, apex acutely rounded, sides slightly convex.
Male genitalia unknown.
Female genitalia as in generic description.
REMARKS. The single known specimen of this species is tentatively regarded as distinct from metrosideri, differing only in its longer head.
DISTRIBUTION. Marquesas Is.
MATERIAL EXAMINED
Dryadomorpha viridia Osborn, holotype 9, Marquesas Is.: Nuku Hiva I., Toovii, 750 m, 4.viii.l931, beating Metrosideros collina (J. R. & G. Forster) (Le Bronnec & Tauraa) (BPBM, Honolulu).
Dryadomorpha robustipenis sp. n.
(Figs 6 1-63)
Length : $, 5-2-6-0 mm, mean 5-5 mm; 9, 6-0-6-3 mm, mean 6- 1 mm.
Colour as in generic description with a small brown spot at apex of clavus and claval veins of forewing.
External characters as in generic description with vertex moderately long, medial length 2-5-3-0 times as long as length next to eyes, apex acutely rounded, sides slightly convex to slightly concave.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 55
Male genitalia similar to pallida but connective moderately long and aedeagus more robust with apical processes shorter and only slightly divergent. Female genitalia as in generic description.
REMARKS. This species closely resembles pacifica and is known to be sympatric with this species over part of its range in Sarawak and Sabah. It can be distinguished from pacifica in the male genitalia by the narrower apical process of the style, the more pronounced lateral lobe of the style and the more robust shaft of the aedeagus. From the similar but more northerly and southerly distributed pallida it differs in the male genitalia as noted above.
DISTRIBUTION. Malaya and Borneo (Sarawak, Sabah).
MATERIAL EXAMINED
Holotype <?, Sarawak: foot of Mount Dulit, junction of rivers Tinjar and Lejok, at light, l.x.1932 (Hobby & Moore) (BMNH, London).
Paratypes. Malaya: 1 <J, Kuala Lumpur, Klang gates, 31.xii.1958 (Quate) (BPBM, Honolulu); 1 9, Kuala Lumpur, 13.iii.1939 (Pendlebury) (BMNH, London). Borneo: 3 <J, 2 ?, Sarawak, same data as holotype, 26.viii.-25.ix.1932; 1 & Sarawak, Paya, Palah, at light, 4.xi.l964 (Rothschild) (all in BMNH, London); 1 <J, Sarawak, Bau, lake area, 30.viii.1958 (Mao) (BPBM, Honolulu); 1 £, Sabah, Tawau, Quoin Hill, at light, 3-7. vii. 1962 (Holtman); 1 <J, Sabah, Tawau, Quoin Hill, Cocao Research Station, 22.viii.1962 (Hiroshima) (all in BPBM, Honolulu).
Dryadomorpha pacifica sp. n.
(Figs 64-67)
Length: <$, 5-2-6-5 mm, mean 5-6 mm; $, 5-5-6-4 mm, mean 5-9 mm.
Colour and external characters as in robustipenis.
Male genitalia similar to robustipenis but apical process of style broader apically and foot-like, or with a subapical prominence, and lateral lobe ol style less pronounced and aedeagus with shaft narrower and tapered more at midlength to apex.
Female genitalia as in generic description.
REMARKS. In this species slight variation occurs in the shape of the apical process of the style and the processes of the aedeagus. In the specimens from Sarawak and Sabah the apical process of the style is foot-like and the aedeagal processes are narrow and nearly parallel, while specimens from New Guinea have the apical process of the style with a slight subapical prominence and the processes of the aedeagus stouter and more divergent. The significance of these differences cannot be ascertained at the present time.
This species closely resembles robustipenis and is known to be sympatric with this species over part of its range in Sarawak and Sabah. It can be distinguished from robustipenis in the shape of the style and aedeagus as noted above. It differs from the similar but more northerly and southerly distributed pallida in the longer connective and the more robust aedeagus with the apical processes shorter and only slightly divergent.
DISTRIBUTION. Borneo (Sarawak, Sabah), New Guinea and Vanuatu.
MATERIAL EXAMINED
Holotype rf, New Guinea: Sarmi, W. to Hollandia, 20-23.vii.1959 (Maa) (BPBM, Honolulu).
Paratypes. Borneo: l<$, Sarawak, Bau district, lake area, 30.viii.1958 (Maa) (BPBM, Honolulu); 1^, Sabah, Tawau, Quoin Hill, Cocoa Research Station, primary forest, 3.x. 1962 (Hiroshima); 1 ^, Sabah, Tawau, Quoin Hill, 3-7.vii.1962 (Holtman) (BPBM, Honolulu). New Guinea: 1 J, Nabire, 5-50 m, at light, 25.viii-2.ix.1962 (Sedlacek); 1 <£ mouth of river Tor, 4 km E. of Maffen, 19. vii. 1959 (Maa); 1 & Kokoda, 400 m, at light, 15-20.xi.1965 (Sedlacek); 1 $, Bodem, 100 m, 11 km SE. of Oerberfaren, at light, 7-17.vii.1959 (Maa) (all in BPBM, Honolulu); 15 <£ same data as holotype (BPBM, Honolulu; BMNH, London). Vanuatu: 3$, Espiritu Santo, Apouna river camp, 300 m, 9-12.ix.1971 (Robinson) (SAM, Adelaide); 1 cJ, Espiritu Santo, Narango, 90 m, vi.1960 (Brandt) (BPBM, Honolulu); 1 ?, Santo I., 17.vi.1925 (Buxton) (BMNH, London).
56
M. D. WEBB
RHUTELORBUS gen. n.
Type-species : Rhutelorbus merinoi sp. n.
Yellow or greenish yellow, sometimes with paler patches on head and thorax ; fore wings with a small brown spot at apex of both clavus and claval veins.
Head wider than pronotum; anterior margin acutely rounded in profile, transversely striate, becoming finely carinate and rim-like medially; ocelli on margin, distant from eyes, not visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approxi- mately 2-5 times length next to eyes, sides slightly convex, apex acutely rounded; with a slight longitudinal depression; finely rugose. Face elongate, shagreen; lateral margin adjacent eye visible dorsally; upper margin slightly depressed each side of mid line, forming a faintly striate longitudinal keel medially; face in profile more or less straight or concave anteriorly; clypeus elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; transclypeal suture indistinct; lora large; antennal pit deep with inner margin angularly rounded to clypeus, rim-like and nearly carinate; antennal ledge slight; an- tennae very long, extending to beyond apex of clavus. Pronotum approximately twice as wide as long, side margins very short, without a carina; finely transversely rugose. Scutellum approximately equal in length to
Figs 68-78 Rhutelorbus merinoi. 68, head and thorax, dorsal view; 69, left subgenital plate and apex of left style, dorsal view; 70, <$ genitale capsule, lateral view; 71, left style, ventral view; 72, connective, lateral view; 73, apex of left style, lateroventral view; 74, connective, dorsal view; 75, 9 pregenital sternite, ventral view; 76, second valvulae, lateral view; 77, 78, aedeagus, lateral and posterior views.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 57
pronotum, shagreen, obscurely rugose posteriorly. Fore wing with three subapical cells, first subapical open, second and third subapicals closed; one or two additional veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of two or three fine setae distally on anterior surface; hind femur with apical setal formula 2+1 + 1 with first proximal seta slender.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes with several macrosetae and short fine setae. Xth segment moderately long, compressed dorsoventrally. Subgenital plate elongate, triangular with numerous moderate to long fine setae dorsally and several short to moderately long fine setae ventrally. Valve triangular. Style moderately long, narrow, with lateral lobe and basal apophyses prominent; apical process fairly short, curved ventrally, tapered to apex, crenulate api?ally; few sensory papilla and a single seta adjacent lateral lobe. Connective Y-shaped, stem elongate with lateral margins keel-like dorsally ; arms short. Aedeagus with shaft elongate, curved dorsally with a pair of apical, dorsally directed processes; gonopore small, situated at apex; basal apodeme moderately long, narrow.
Female genitalia with posterior margin of pregenital sternite extended posteriorly; second valvulae united at first dorsal tooth (arrowed in Fig. 76), elongate, very slightly expanded distally; without a basal promi- nence; dorsal teeth robust, unaligned, extending over approximately distal third of valvulae.
REMARKS. This genus is similar externally to Dryadomorpha and some species of Parohinka but can be distinguished from these and other genera of the subfamily by the rugose vertex and pronotum, the lateral margins of the face adjacent to the eyes being visible dorsally, and by the extended posterior corners of the pregenital sternite (also present in some species of Parohinka).
DISTRIBUTION. Philippines, Malaya and Borneo (Sarawak).
Rhutelorbus merinoi sp. n.
(Figs 68-78)
Length: c?, 5-6-6-5 mm, mean, 6-0 mm; 9, 6-2-7-4 mm, mean 6-8 mm.
Colour and external characters as in generic description.
Male genitalia as in generic description with pygophore lobes acutely produced posteriorly and aedeagus with shaft narrow, slightly tapered from base to apex with apical processes fairly short (see 'Remarks' below).
Female genitalia with posterior margin of pregenital sternite extended posteriorly, maximum length as in Fig. 75, sometimes one-third this length.
REMARKS. The male specimen from the Philippines has the apical process of the style curved more ventrally and the apical processes of the aedeagus shorter than in other specimens.
DISTRIBUTION. Philippines, Malaya and Borneo (Sarawak).
MATERIAL EXAMINED
Holotype £, Malaya : Kuala Lumpur, at light, 16.X.1928 (Corbett) (BMNH, London).
Paratypes. Philippines: 1 9, Tawi Tawi, Lapid Lapid at Manalik Channel, at light, 19.xi.1961; 1 ^, Tawi Tawi, Tarawakan, north of Batu Batu, at light, 13.xi.1961 (both in ZM, Copenhagen). Malaya: 1 9, Kuala Lumpur, at light, 1811931 (Pendlebury) (BMNH, London); 2^, 29, Kuala Lumpur, 24-3 l.xii. 1958 (Quate); 4 c?, 1 9, Kuala Lumpur, Klang gates, 3 l.xii. 1958 (Quate); 1 9, Kuala Lumpur, 19 km south of Subang, 23.xii.1958 (Quate); 1 & Kuala Tahan, 15-16.xii.1958 (Quate) (all in BPBM, Honolulu); 1 <J, Selangor, at light, 19.iv.1932 (Pendlebury) (BMNH, London). Borneo: 1 $ no locality, 1898 (Noualhier) (MNHN, Paris); 1 cJ, Sarawak, foot of Mount Dulit, junction of R. Tinjar and Lejok, old secondary forest, at light, 29.viii.1932 (Hobby & Moore) (BMNH, London); 1 9, Sarawak, Bau District, Bidi, 19-240 m, at light, 2.iv.l958 (Maa); 1 9, Sarawak, Kuching, Santubong, 797-1500 m, 18-30.vi.1958 (Maa) (both in BPBM, Honolulu).
PAROHINKA gen. n.
Type-species: Muirella longiseta Melichar, 1914.
Yellow, greenish yellow or brownish yellow; head with yellow or whitish yellow patches, usually mottled with brown. Fore wings with a small brown spot at apex of both clavus and claval veins, often faint. Legs spotted with brown.
58 M. D. WEBB
Head wider than pronotum; anterior margin angularly rounded in profile, transversely striate laterally, becoming finely carinate and usually rim-like medially, ocelli on margin distant from eyes, not visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length 1-5-4-0 times length next to eyes, sides slightly convex to slightly insinuate in longer-headed forms, apex acute to broadly angularly rounded; with a medial longitudinal depression; finely longitudinally striate, transversely striate anteriorly. Face elongate to only slightly longer than wide, shagreen; upper margin usually slightly depressed each side of mid line, forming a faintly striate longitudinal keel medially, indistinct in shorter-headed forms; face in profile convex to concave in longer-headed forms; clypeus elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; transclypeal suture distinct or indistinct; lora large; antennal pit deep with inner margin angularly rounded to clypeus and sometimes faintly rim-like and nearly carinate; antennal ledge absent; antennae very long, when recurved extending beyond apex of clavus. Pronotum approximately twice as wide as long, side margins very short, without a carina; finely and irregularly transversely striate, obscurely rugose anteriorly. Scutel- lum approximately equal in length to pronotum, shagreen with posterior region obscurely rugose. Fore wings with three subapical cells, first subapical cell open, second and third subapical cells closed; with a few additional veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of seven setae distally on anterior surface; hind femora with apical setal formula 2+1 + 1 with first proximal seta slender.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with posterior margin of pygophore straight dorsally, without apodemes; pygophore lobes with a slight oblique internal ledge, usually not attaining posterior margin; posterior margin sometimes serrate and sometimes extended spine-like dorsally; lobes with several macrosetae and numerous short to moderately long fine setae. Xth segment moderate to long, compressed dorsoventrally, narrowing basally in lateral aspect. Valve triangular. Subgenital plate elongate, triangular with basal lobe relatively short and prominent ; outer margin with numerous long to very long fine setae on dorsal and ventral surfaces ; apex with few very short stout setae. Style short with basal apophyses indistinct; apical process short to moder- ately long, curved ventrally, tapered to apex or spatulate, ventral surface crenulate; lateral lobe prominent; few sensory papilla ventrally adjacent lateral lobe. Connective Y-shaped, stem moderate to very long, narrow, with lateral margins keel-like dorsally; arms short. Aedeagus assymetrical ; shaft elongate, curved dorsally, sometimes recurved ventrally at apex, either of similar width throughout length, tapered to apex or expanded distally in lateral aspect, sometimes compressed anteroposteriorly; with from one to three pro- cesses distally; sometimes with an apical membranous region adjacent to gonopore; gonopore small, situated at or near apex on left or right lateral surface within a species or on posterior surface; basal apodeme short to moderately long, compressed laterally or anteroposteriorly.
Female genitalia with posterior margin of pregenital sternite either broadly V-shaped medially or with posterior corners extended posteriorly. Second valvulae united at or before first dorsal tooth (arrowed in Figs 110, 121, 133, 148), expanded distally, fairly robust, without a dorsoanterior prominence; dorsal teeth fine to robust, unaligned, extending 0-25-0-40 times distance from apex to base of valvulae; dorsal scler- otized region moderately long.
REMARKS. This genus is almost identical externally to Dryadomorpha but can usually be dis- tinguished by the pale patches on the anterior margin of the head. Also similar externally to Rhutelorbus (see 'Remarks', p. 57). In the male genitalia the pygophore has an internal ledge as in Dryadomorpha, Karoseefa and Oceanopona, but the asymmetrical aedeagus is not found in any other genus of the subfamily. In the female genitalia the extended posterior corners of the pregenital sternite of some species (dulita-group) are as in Rhutelorbus and the second valvulae are as in Rhutelorbus, Karoseefa and Oceanopona.
On the basis of the male and femal genitalia Dryadomorpha can be divided into two groups. The dulita-group (dulita, brevicephala, philippina, recurva, apicalis, longiseta and malayensis) differs from the morona-group (morona, sinuata, trispicata, lotophagorum and spinosa) in having the apex of the aedeagal shaft continued as a process on one side and the posterior corners of the female pregenital sternite extended posteriorly either process-like or lobe-like in dulita.
DISTRIBUTION. From NE. India to the Philippines and South East Asia and south to New Guinea and the Pacific islands. The dulita-group extends as far south as Indonesia, while the morona- group is found in New Guinea and eastwards to the Cook Is.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 59
Key to species of Parohinka
Males
1 Aedeagus with apex of shaft continued as a process on one side (Figs 105, 125). (Oriental region
and south to Indonesia) 6
Aedeagus with apex of shaft not continued as a process on one side. (New Guinea and eastwards to the Cook Is.) 2
2 Aedeagus with two processes (both lateral or one lateral and one posterior) .... 3 Aedeagus with three processes (two lateral and one posterior) 5
3 Aedeagus with shaft in lateral aspect of similar width throughout length or narrower apically;
with two lateral processes directed ventroanteriorly 4
Aedeagal shaft in lateral aspect expanded apically; with one lateral process directed ventropo- steriorly and one posterior process directed ventroanteriorly (Figs 93, 94)
lotophagorum (Kirkaldy) (p. 63)
4 Aedeagal shaft in lateral aspect evenly curved from base to apex (Fig. 84) . . morona sp. n. (p. 60) Aedeagal shaft in lateral aspect sinuate apically (Fig. 86) sinuata sp. n. (p. 61)
5 Posterior process of aedeagus short, approximately half length of lateral processes (Fig. 91)
trispicata sp. n. (p. 61) Posterior process of aedeagus long, similar in length to lateral processes (Fig. 102)
spinosa sp. n. (p. .64)
6 Apex of aedeagal shaft curved ventrally (Fig. 124). (Borneo (Sarawak)) . . recurva sp. n. (p. 67) Apex of aedeagal shaft curved dorsally 7
7 Aedeagus with one process 8
Aedeagus with two or three processes 9
8 Connective approximately equal in length to style. (Malaya) . . . malayensis sp. n. (p. 70) Connective approximately 1-5 times length of style. (NE. India to Indonesia)
longiseta (Melichar) (p. 68)
9 Aedeagus with two processes. (Borneo (Sarawak, Sabah)) apicalis sp. n. (p. 68)
Aedeagus with three processes ............. 10
10 Lateral aedeagal processes short, arising from near posterior margin of shaft (Fig. 119).
(Philippines) philippina sp. n. (p. 66)
Lateral aedeagal processes moderately long, arising from near anterior margin of shaft (Fig. 107) 11
11 Aedeagal shaft in lateral aspect narrowly club-shaped (Fig. 107). (Borneo (Sarawak))
dulita sp. n. (p. 64) Aedeagal shaft in lateral aspect of similar width throughout length (Fig. 1 15). (Borneo (Sarawak,
Sabah)) brevicephala sp. n. (p. 66)
Females
The following key is incomplete because females of morona, sinuata and trispicata are indistinguishable (grouped together under couplet 2) and spinosa and recurva are known only from the male.
1 Posterior corners of pregenital sternite extended posteriorly, process-like or lobe-like. (Oriental
region as far south as Indonesia) 3
Posterior corners of pregenital sternite not extended posteriorly as above. (New Guinea and eastwards to the Cook Is.) . 2
2 Posterior margin of pregenital sternite oblique and nearly straight each side of mid line (Fig. 83)
morona sp. n. (p. 60), sinuata sp. n. (p. 61), trispicata sp. n. (p. 61) Posterior margin of pregenital sternite angled at mid length each side of mid line (Figs 96, 97)
lotophagorum (Kirkaldy) (p. 63)
3 Posterior corners of pregenital sternite extended process-like 4
Posterior corners of pregenital sternite lobe-like (Fig. 109) dulita sp. n. (p. 64)
4 Vertex with medial length 0-5-0-9 times basal width between eyes ... 5 Vertex with medial length 1-0-1-2 times basal width between eyes 6
5 Vertex with medial length 0-5 times basal width between eyes. Extended posterior corners of
pregenital sternite short (Fig. 118). (Borneo (Sarawak)) . . . . brevicephala sp. n. (p. 66) Vertex with medial length 0-5-0-9 times basal width between eyes. Extended posterior corners of
pregenital sternite long (Fig. 139). (NE. India to Indonesia) . . . longiseta (Melichar) (p. 68)
6 Pale to greenish yellow. Second valvulae fairly broad (Fig. 121). (Philippines)
philippina sp. n. (p. 66) Sordid to brownish yellow. Second valvulae narrow (Fig. 148). (Malaya) . malayensis sp. n. (p. 70)
60
M. D. WEBB
Parohinka morona sp. n.
(Figs 79-85)
Length:^, 5-9-6- 3 mm, mean 6-1 mm;?, 6-6 mm.
Pale to greenish yellow. Vertex, dorsal region of face and sometimes medial region of pronotum mottled with brown. Inner margin of fore wing sometimes tinged with brown (see 'Remarks' below); other markings as in generic description.
External characters as in generic description with vertex long, medial length 2-8-3-5 times length next to eyes and 1-0-1-2 times basal width between eyes. Face longer than wide with upper medial keel present; clypeus elongate.
Male genitalia as in generic description with pygophore moderately long. Xth segment extending to or slightly beyond posterior margin of pygophore. Connective moderately long (as in malayensis, Fig. 145). Style with apical process digitate, apex of process truncate in medial aspect. Aedeagus with shaft in lateral aspect tapered gradually from base to apex and terminating in a pair of ventroanteriorly directed processes; shaft sinuate in posterior aspect; gonopore situated laterally immediately below apical processes.
Female genitalia with posterior margin of pregenital sternite shallowly V-shaped (see 'Remarks' below); second valvulae united at first dorsal tooth; teeth robust, extending to approximately 0-4 times distance from apex to base of valvulae (as in longiseta, Fig. 133).
REMARKS. Some of the specimens examined of this species have a brown medial longitudinal band dorsally, from the head to the apex of the fore wings.
80
81
82
83
85
Figs 79-86 Parohinka species. 79-85. P. morona. (79) head and thorax, dorsal view; (80) apex of left style, ventromedial view; (81) ^ genital capsule, lateral view; (82) left style, ventral view; (83)$ pregenital sternite, ventral view; (84, 85) aedeagus, lateral and posterior views. 86. P. sinuata, aedeagus, lateral view.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 61
The following four species and morona form a distinct group based on distribution (New Guinea and east to the Cook Is.) and also on the shape of the male and female genitalia (see 'Remarks' under generic description, p. 58). On external appearance and in the female genitalia morona is identical to trispicata and sinuata. Females of the three species are therefore included in their respective type-series as they were taken at the same localities as the males therein and there being no sympatry between the males of the three species samples. Only slight differences in the shape of the pregenital sternite distinguishes females of morona, sinuata and trispicata from lotophagorum; in the first three species the posterior margin of the pregenital sternite is oblique and almost straight each side of the mid line while in lotophagorum it is angled at mid length each side of the mid line. In external appearance morona is similar to spinosa and is sympatric with this species over part of its range in Papua New Guinea. Females of spinosa are unknown. In the male genitalia morona differs from the above four species in having the posterior margin of the pygophore non-serrate, the aedeagus with two apical processes and the aedeagal shaft evenly curved from base to apex in lateral aspect.
DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea).
MATERIAL EXAMINED
Holotype ^, New Guinea: Irian Jaya, Cyclops Mts, Sabron, camp 2, 660 m, vii.1936 (Cheesman) (BMNH; London)
Paratypes. New Guinea: 1 <J, 1 ?, Irian Jaya, Torricelli Mts, Mokai Vill., 750 m, 8-15.xii.1958 (Brandt); 2 cJ, I.J., 40 km north of Baliem Val., 1300 m, at light, 5-ll.xi.1961 (Quate); 1 & Papua New Guinea, Swart Val., Karubaka, 1450 m, 17.xi.1958 (Gressitt) (all in BPBM, Honolulu).
Parohinka sinuata sp. n.
(Fig. 86)
Length : <J, 6-2-6-6 mm, mean 6-4 mm; $, 7-5 mm.
Colour and external characters as in morona with medial length of vertex 2-7-3-0 times length next to eyes and 0-8-1-1 times basal width between eyes.
Male genitalia as in morona but aedeagal shaft apically with processes slightly shorter.
Female genitalia as in morona.
REMARKS. This species is closely related to morona differing only in the shape of the aedeagus as noted above (see also 'Remarks' under morona, p. 60).
DISTRIBUTION. New Guinea (Papua New Guinea)
MATERIAL EXAMINED
Holotype cJ, New Guinea: Papua New Guinea, Lae, Singuawa R., 147° 10' E, 6° 45' S, 30 m, at light, Kunai grass, 8.iv.l966 (Wilkes) (BPBM, Honolulu).
Paratypes. New Guinea: 1 <$, 1 9, same data as holotype (BMNH, London; BPBM, Honolulu); 1 £, P.N.G., Wau, Morobe District, 1250m, 2311963 (Sedlacek) (BPBM, Honolulu).
Parohinka trispicata sp. n.
(Figs 87-91)
Length : $, 5-6-6-0 mm, mean 5-8 mm; $, 6-4-7-0 mm, mean 6-7 mm.
Colour and external characters as in morona with medial length of vertex 2-5-3-0 times length next to eyes and approximately equal to basal width between eyes.
Male genitalia similar to morona but apex of aedeagus with a small additional process on one side adjacent to gonopore.
Female genitalia as in morona.
REMARKS. This species is identical externally and in the female genitalia to morona and sinuata (see 'Remarks' under morona, p. 60). In the male genitalia it differs from morona and sinuata in having three processes on the aedeagus.
DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea) and Solomon Is.
M. D. WEBB
98
101
102
Figs 87-102 Parohinka species. 87-91. P. trispicata. (87) $ pygophore and Xth segment, lateral view; (88) apex of style, ventromedial view; (89) connective, dorsal view; (90, 91) aedeagus, lateral and posterior views; 92-97. P. lotophagorum. (92) <$ pygophore and Xth segment, lateral view; (93, 94) aedeagus, lateral and posterior views; (95) apex of left style, ventromedial view; (96,97) $ pregenital sternite, ventral view; 98-102. P. spinosa. (98) <$ pygophore and Xth segment, lateral view; (99) apex of left style, ventromedial view; (100) aedeagus, lateral view; (101) left style, ventral view; (102) aedeagus, posterior view.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 63
MATERIAL EXAMINED
Holotype cJ, Solomon Is.: Kolombangara, Pepele, 30 m, 6.H.1964 (Shanahan) (BPBM, Honolulu) Paratypes. New Guinea: 1 (J, Irian Jaya, 10.xi.1944 (Aarons) (CAS, San Fransisco); 3^, 1 ?, I.J., Hollandia, Binnen, 100 m, at light, 31.X.1958 (Gressitt); 1 9, I.J., Hollandia, at light, 28.ii.1960 (Maa); 1 & Papua New Guinea, Adelbert Mts, Wanuma, 800-1000 m, at light, 27.x. 1958 (Gressitt) (all in BPBM, Honolulu). Solo- mon Is.: 10 cJ, 4 9, Kolombangara, Pepele, 30 m, 6, 10, 15.ii.1964 (Shanahan) (BPBM, Honolulu; BMNH, London)
Parohinka lotophagorum (Kirkaldy) comb. n. (Figs 92-97)
Dryadomorpha lotophagorum Kirkaldy, 1907: 41. 1 <$, 1 9 syntypes, FIJI: Viti Levu, Ruwa, iii-vi (Muir) (lost). NEOTYPE (J, FIJI (BPBM, Honolulu), here designated [examined].
Length: J, 5-4-6-5 mm, mean 6-0 mm; 9, 6-6-7-6 mm, mean 7-0 mm.
Pale to greenish yellow. Vertex and dorsal region of face mottled with brown. Fore wings sometimes tinged with pinkish orange. Other markings as in generic description.
External characters as in morona with medial length of vertex 2-4-3-0 times length next to eyes.
Male genitalia similar to morona but posterior margin of pygophore serrate and apical process of style with a subapical prominence in medial aspect. Aedeagus with shaft tapered from base to near apex and expanded slightly at apex in lateral aspect; a more or less straight process arising at apex on one side, directed ventrolaterally and another process arising on opposite side to first and slightly below apex, strongly curved ventrolaterally; gonopore apical on posterior surface.
Female genitalia with posterior margin of pregenital sternite V-shaped medially; second valvulae as in morona.
REMARKS. The type-series of lotophagorum was listed as missing at the time when the Hawaiian Sugar Planters' Association collection was transferred to the Bishop Museum and is presumed lost. The specimens described above fit the original description almost exactly, only the darker brown spots on the lateral margins of the clypeus, mentioned by Kirkaldy, are not clear.
In external appearance this species is similar to morona, sinuata, trispicata and spinosa and is sympatric with trispicata over part of its range in the Solomon Is. In the female the pregenital sternite is slightly different in shape to that of morona, sinuata and trispicata (see 'Remarks' under morona, p. 60), and in the male genitalia both this species and spinosa can be distinguished from other members of the genus by the serrate posterior margin of the pygophore. From spinosa it differs in the male genitalia having the posterior margin of the pygophore without a spine-like process, the style with the apical process more robust with a preapical prominence, and the aedeagus with two processes.
DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea), Solomon Is., Vanuatu, Fiji and Niue Is.
MATERIAL EXAMINED
Dryadomorpha lotophagorum Kirkaldy, neotype $, Fiji: Lau group, Ono-i-Lau, Nukuni, 0-50 m, 24.ii.1971 (Krauss) (BPBM, Honolulu).
New Guinea: 1 $, 1 9, Irian Jaya, Waigeu, Camp Nok., 750 m, iv.v.1938 (Cheesman) (BMNH, London); 1 (J, I.J., Japen I., SSE. Sumberbaba, Dawai R., at light, 22.x. 1962 (Holtmann); 2 <$, 19, I.J., Waris, S. of Hollandia, 450-500 m, m.v. light-trap, l-15.viii.1959 (Maa); 1 & 2 9, I. J-, Bodem, 100 m, 11 km SE. of Oerberfaren, m.v. light-trap, lO.vii, 16.xii.1959 (Maa); 3 $, Papua New Guinea, Kokoda, 400 m, light-trap, malaise trap, 15-20.xi.1965 (Sedlacek) (all in BPBM, Honolulu); 1 & P.N.G., Madang Dist., Finisterre Mts, Damanti, 105 m, stn no. 46, 2-1 1.x. 1964 (Bacchus) (BMNH). Bismark Archipeligo: 4<J, New Britain, Gazelle Pen., Gaulim, 130 m, 140 m, at light, 23-28.X, 19-20.xi.1962 (Sedlacek); 1 <J, N.B., Gazelle Pen., Mt Sinewit, 900 m, at light, 5-14.xi.1962 (Sedlecek) (all in BPBM, Honolulu). Solomon Is.: 1 9, 2 specimens (abdomens missing), Guadalcanal, Mt Austen, 15.viii.1966 (Greenslade) (BMNH, London); 2^, G., 17-6 km SE. Tetere, Tathimani, light-trap, 12. v. 1960 (O'Brien); 1 J, G., Kukum, 10 m, light-trap, 21.vi.1956 (Gressitt); 4£, 59, Kolombangara, Pepele, 30 m, 6, 10, 15.ii.1964 (Shanahan); 2 cJ, 39, NW. Malaita, Dala, light-trap, 5- 7.vi.l964 (Straatman); 1 <$, San Cristobal, Wairahu R., 100 m, pressure lamp, 9-15.V.1964 (Straatman) (all in BPBM, Honolulu). Vanuatu: 1 £, Lamen I., 0-10 m, i.1976 (Krauss) BPBM, Honolulu); 1 <J, Tanna I.,
64 M. D. WEBB
Isokoai (Enpinana), 28.vii.1971 (Gross) (SAM, Adelaide). Fiji: 2 cJ, 1 $, Lau, Group, Ono-i-Lau, Nukuni, 0-50 m, 24.ii.1971 (Krauss) (BPBM, Honolulu). Cook Is.: 1 ?, Niue I., nr Lefuka, bush area, m.v. light, 18.vi.1975 (Maddison) (BMNH, London); 1 <J, Niue I., Kavara, at light, ll.vi.1975 (Dugdale) (DSIR, Auck- land).
Parohinka spinosa sp. n.
(Figs 98-102)
Length: <$, 5-8 mm.
Pale yellow with vertex and dorsal region of face mottled with brown ; other markings as in generic description.
External characters as in morona with medial length of vertex 3-0 times length next to eyes and 1-2-1-3 times basal width between eyes.
Male genitalia similar to lotophagorum but posterior margin of pygophore extended into a long spine-like process dorsally and apical process of style narrower, without a preapical prominence. Aedeagus with shaft expanded from base to apex in lateral aspect, two lateral and one posterior process arising immediately below apex, directed ventrally, moderately long.
Female genitalia unknown.
REMARKS. In external appearance this species is similar to morona, sinuata, trispicata and lotopha- gorum and sympatric with morona in its single known locality in Papua New Guinea. In the male genitalia spinosa and lotophagorum can be distinguished from other members of the genus by the serrate posterior margin of the pygophore. From lotophagorum it differs in the male genitalia in the shape of the pygophore and style, as noted above, and the aedeagus with three processes.
DISTRIBUTION. Known only from the type-locality in New Guinea (Papua New Guinea).
MATERIAL EXAMINED
Holotype £, New Guinea: Papua New Guinea, Cyclops Mts, Sabron, 279 m, v.1936 (Cheesman) (BMNH, London)
Paratypes. New Guinea: 1 $, same data as holotype except, camp 2, 600 m; 3 <$, same data as holotype except, vi.1936 (BMNH, London)
Parohinka dulita sp. n.
(Figs 103-1 10)
Length: ^, 5-5-5-9 mm, mean 5-7 mm; 9, 5-8-6-2 mm, mean 6-1 mm.
Pale, sordid or greenish yellow. Vertex and dorsal region efface mottled with pale brown; other markings as in generic description.
External characters as in morona but vertex moderately long, medial length 2-1 times length next to eyes and 0-8 times basal width between eyes.
Male genitalia as in generic description with pygophore moderately long and Xth segment extending slightly beyond posterior margin of pygophore. Connective moderately long, upturned apically. Style with apical process digitate. Aedeagus with shaft narrowly club-shaped in lateral aspect with a moderately long process at apex on one side, directed dorsally, and a moderately long subapical process on each side, arising against anterior margin, directed ventrally; apical region of shaft adjacent process and surrounding go- nopore membranous.
Female genitalia with posterior margin of pregenital sternite broadly V-shaped medially, each posterior corner slightly produced lobe-like; second valvulae united at approximately 0-40 distance from apex to base of valvulae ; teeth fine, extending to approximately 0-25 distance from apex to base of valvulae.
REMARKS. The following six species and dulita form a distinct group based on distribution (Oriental region from NE. India to Indonesia) and on the shape of the male and female genitalia (see 'Remarks' under generic description, p. 58). In the length of the vertex dulita is similar to some specimens of longiseta but differs from this and other members of the genus by the lobe-like posterior corners of the pregenital sternite and the valvulae united basad of the teeth. It is very similar to brevicephala in the male genitalia but differs in having the apical process of the style narrower in medial aspect and the shaft of the aedeagus narrowly club-shaped. This species is known to be sympatric with longiseta and brevicephala in Sarawak.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
104 . VA 106
65
108
109
112
116
Figs 103-118 Parohinka species. 103-1 10. P. dulita. (103) head and thorax, dorsal view; (104) left style, ventral view; (105) aedeagus, posterior view; (106) apex of left style, ventromedial view; (107) aedeagus, lateral view; (108) connective, dorsal view; (109)9 pregenital sternite; (110) second valvu- lae, lateral view. 111-118. P. brevicephala. (Ill) head and thorax, dorsal view; (112) face; (113) aedeagus, posterior view; (114) apex of left style, medial view; (115) aedeagus, lateral view; (116) connective, lateral view; (117) left style, ventral view; (1 18) 9 pregenital sternite, ventral view.
DISTRIBUTION. Known only from Borneo (Sarawak, Mt Dulit).
MATERIAL EXAMINED
Holotype <$, Borneo: Sarawak, foot of Mt Dulit, junction of R. Tinjar & Lejok, light-trap, 31.viii.1932 (Hobby & Moore) (BMNH, London)
Paratypes. Borneo: 2 $, same data as holotype, except, old secondary forest, 29.vii.1932; 1 9, same data as holotype except, 29.viii.1932; 1 <$, same data as holotype except, l.ix.1932; 1 9, same data as holotype except, 4.X.1932; 1 $, Sarawak, Mt Dulit, Dulit trail, old secondary forest, 3.ix.l932 (Hobby & Moore) (BMNH, London)
66 M. D. WEBB
Parohinka brevicephala sp. n.
(Figs 11 1-1 18)
Length : <£, 6-2 mm ; 9, 6-9-7- 1 mm, mean 7-0 mm.
Colour as in dulita.
External characters as in generic description with vertex short, medial length 1-5 times length next to eyes and 0.5 times basal width between eyes. Face slightly longer than wide with upper medial keel very slight or absent ; clypeus relatively short and broad.
Male genitalia similar to dulita but style with apical process slightly expanded in medial aspect and aedeagus with shaft of similar width throughout length in lateral aspect.
Female genitalia with posterior margin of pregenital sternite broadly U-shaped with each posterior corner extended posteriorly process-like; second valvulae as in morona.
REMARKS. In external appearance this species can be distinguished by its short head and relatively short and broad clypeus. It is sympatric with dulita and longiseta over part of its range in Sarawak and with apicalis in its single known locality in Sabah.
DISTRIBUTION. Borneo (Sarawak, Sabah).
MATERIAL EXAMINED
Holotype <£, Borneo: Sarawak, Gunong Mulu Nat. Park, Long Pala, base camp, 1978 (Collins) (BMNH, London)
Paratypes. Borneo: 6 <$, 3 $, same data as holotype; 69, Sarawak, foot of Mt Dulit, junction of R. Tinjar & Lejok, light-trap/at light in house, 1, 2, 26.ix., 3, 31.X.1932 (Hobby & Moore) (BMNH, London); 1 & Sabah, Tawau, Quoin Hill, light-trap, 3-7.vii.1962 (Holtmann) (BPBM, Honolulu).
Parohinka phiUppina sp. n.
(Figs 119-123)
Length: $, 6-6-7-3 mm, mean 7-0 mm; 9, 8-4-8-7 mm, mean 8-6 mm.
Pale to greenish yellow. Vertex and dorsal region of face faintly mottled with brown, or markings absent. Fore wings with or without a brown spot at apex of both clavus and claval veins.
External characters as in morona but vertex longer, medial length 2-7-3-5 times length next to eyes and 1-0-1-1 times basal width between eyes.
Male genitalia as in dulita but aedeagus with shaft of similar width throughout length in lateral aspect, processes short with lateral pair directed more anteriorly.
Female genitalia similar to brevicephala but second valvulae broader with distance from apex of first dorsal tooth to dorsal hyaline region relatively long.
REMARKS. The elongate head of this species is similar to that found in apicalis and malayensis and slightly longer than in specimens of longiseta seen from the same localities in the Philippines. The male genitalia are similar to those of dulita and brevicephala but differ in the shape of the aedeagus as noted above and from dulita by the shorter and broader apical process of the style in medial aspect. In the female genitalia philippina can be distinguished by the relatively long distance from the apex of the first dorsal tooth to the dorsal hyaline region of the second valvulae.
DISTRIBUTION. Philippines (Luzon)
MATERIAL EXAMINED
Holotype £, Philippines: Luzon, Ifugao Prov., Liwo, 8 km E. of Mayoyao, 1000-1300 m, light-trap, l-3.vi.1967 (Torrevillas) (BPBM, Honolulu)
Paratypes. Philippines: 9<$, 89, same data as holotype except, 29.v-l.vi.1967, 7,8,ll,12.vi.l967; 2$, Luzon, Ifugao Prov., Jacmal Bunhian, 24 km E. of Mayoyao, 800-1000 m, light-trap, 30.iv., 9- 10. v. 1967 (Torrevillas) (BPBM, Honolulu; BMNH, London).
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
67
120
124
125
Figs 119-128 Parohinka species. 119-125. P. philippina. (119, 120) aedeagus, lateral and posterior views; (121) second valvulae, lateral view; (122) apex of left style, ventromedial view; (123) left style, ventral view. 124, 125. P. recurva, aedeagus, lateral and posterior views. 126-128. P. apicalis. (126) connective, dorsal view; (127, 128) aedeagus, lateral and posterior views.
Parohinka recurva sp. n.
(Figs 124, 125)
Length: c?, 6-6 mm.
Pale yellow. Vertex, dorsal region of face and pronotum mottled with brown ; other markings as in generic description.
External characters as in morona but vertex longer, medial length 4-0 times length next to eyes and 1 -4 times basal width between eyes.
Male genitalia similar to dulita with aedeagal shaft of similar width from base to near apex in lateral aspect, apical region narrower, curved laterally and ventrally and terminating in a moderately long ventrally directed process on one side; a slightly curved, antero ventrally directed process arising laterally against posterior margin slightly basad of narrow apical region of shaft; gonopore apical, adjacent to apical process.
Female genitalia unknown.
REMARKS. This species can be distinguished by its elongate head and ventrally curved apex of the aedeagus with one apical and one lateral process. It is sympatric with apicalis and longiseta (both with shorter heads) in its single known locality in Sarawak.
DISTRIBUTION. Known only from a single specimen from Borneo (Sarawak).
MATERIAL EXAMINED
Holotype <J, Borneo: Sarawak, Kuching, Matang, 450-894 m, m.v. light-trap, 15.ix.1958 (Gressitt & Mad) (BPBM, Honolulu).
68 M. D. WEBB
Parohinka apicalis sp. n.
(Figs 126-128)
Length:^, 5-6-7-1 mm, mean 6-6 mm; $,8-0 mm.
Colour and external characters as in philippina with medial length of vertex 2-7-3-7 times length next to eyes and 1 -2 times basal width between eyes.
Male genitalia with pygophore elongate, as in longiseta (Fig. 131). Xth segment extending to approxi- mately level with posterior margin of pygophore. Connective very long, slightly upturned apically. Style with apical process slightly expanded. Aedeagus with shaft evenly tapered from base to apex in lateral aspect, compressed slightly anteroposteriorly with two short apical processes on one side, directed dorsally; gonopore apical, on one side, adjacent to processes.
Female genitalia unknown.
REMARKS. The elongate head of this species is similar to that of philippina and malayensis and slightly shorter than in recurva. In the male genitalia apicalis differs from philippina and mal- ayensis in having two aedeagal processes and from recurva by having the apex of the aedeagus directed dorsally. Two female specimens, one from Sabah, Quoin Hill (BPBM, Honolulu) and one from Sarawak, Gunong Matang (BMNH, London) may be this species but differ slightly from one another in the shape of the genitalia.
This species is sympatric with brevicephala and longiseta (both with shorter heads) in Sabah and with longiseta and recurva in Sarawak.
DISTRIBUTION. Borneo (Sarawak, Sabah).
MATERIAL EXAMINED
Holotype <$, Borneo: Sarawak, Kuching, Matang, 450-894 m, m.v. light-trap, 15.ix.1958 (Gressitt & Mad) (BPBM, Honolulu).
Paratypes. Borneo: 1 £, Sarawak, Gunong Matang, 120 m, m.v. light-trap, 16.ix.1958 (Gressitt & Mad) (BMNH, London); 1 <$, Sabah, Tawau, Quoin Hill, 1 5-20. vii. 1962 (Hirashimd) (BPBM, Honolulu).
Parohinka longiseta (Melichar) comb. n.
(Figs 129-143) Muirella longiseta Melichar, 1914: 135. LECTOTYPE & JAVA (RNH, Leiden), here designated [examined].
Length: $, 5-1-6-3 mm, mean 5-7 mm; $,5-7-6-3 mm, mean 6-0 mm.
Pale yellow to stramineous, sometimes tinged with green. Head mottled with brown, stramineous or green, or markings absent; clypeus often patched with stramineous to reddish brown laterally. Forewings sometimes with apex and a transverse band at mid length, brown (see 'Remarks' below); other markings as in generic description.
External characters as in morona but vertex shorter, medial length 1-5-2-6 times length next to eyes and 0-5-0-9 times basal width between eyes. Face with upper medial keel faint in shorter-headed forms; clypeus elongate. Male genitalia as in generic description with pygophore and Xth segment elongate, the latter extending considerably beyond posterior margin of pygophore. Connective with stem elongate, slightly upturned apically. Style with apical process expanded. Aedeagus with shaft evenly curved and tapered from base to apex in lateral aspect with a single apical process directed dorsally; often apex of shaft surrounding gonoduct membranous; few faint longitudinal ridges on posterior surface of shaft and variably on apical process ; gonopore apical, adjacent to apical process.
Female genitalia as in brevipennis, with extended posterior corners of pregenital sternite short to long.
REMARKS. This is the most widespread species of the genus and sympatric with all six other species in the dulita-group over parts of its range from the Philippines to Sarawak. Specimens with shorter heads are similar to brevicephala but have the clypeus and extended posterior corners of the pregenital sfernite longer. The male genitalia are similar to malayensis but differing in the longer pygophore, Xth segment and connective.
In the male genitalia the apical process of the aedeagus shows some variation in shape (see Figs 136-138) and a few specimens from Malaya have the fore wings conspicuously marked with brown, as noted above; these specimens also have the markings on the head darker brown and the vertex shorter than in other specimens.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE
129 / \ 131
69
Figs 129-148 Parohinka species. 129-143. P. longiseta. (129, 130) head and pronotum, dorsal view; (131) c£ pygophore and Xth segment, lateral view; (132) apex of left style, medial view; (133) second valvulae, lateral view; (134) $ Xth segment, dorsal view; (135) aedeagus, posterior view; (136) apex of aedeagus, posterior view, Java; (137) same, Sarawak, Paya Paloh; (138) same, Malaya, Selangor; (139) apex of $ abdomen, ventral view; (140) left style, ventral view; (141, 142) connective, lateral and dorsal views; (143) aedeagus, lateral view. 144-148. P. malayensis. (144) head and pronotum, dorsal view; (145) connective, dorsal view; (146) aedeagus, posterior view; (147)$ pregenital sternite, ventral view; (148) second valvulae, lateral view.
70 M. D. WEBB
DISTRIBUTION. Nepal, extreme north-east of India eastwards to the Philippines and south to Java (excluding Borneo).
MATERIAL EXAMINED
Muirella longiseta Melichar, lectotype <$, Java : Wonosobo, v.1909 (Jacobson) (RNH, Leiden).
Nepal: 1 <J, Adhabbar, Godavari, 1500 m, 5.viii.l967 (CNC, Ontario); 1 9, Kathmandu, 1300-1400 m, 7-12.V.1966 (Sedlacek) (BPBM, Honolulu). India: 1 <J, W. Bengal, Darjeeling, 9.xi.l969 (Das) (BMNH, London). Thailand: 1 <J, Chieng Mai Prov., Doisuthep, 100 m, at light, 18.vi.1965 (Miyatake) (ELKU, Fukuoka). Philippines: 17 <J, 4$, Luzon, Ifugao Prov., Liwo, 8 km E. of Mayoyao, 1000-1300 m, light-trap, 30.v-12.vi.1979 (Torrevillas); 1 <J, Ifugao Prov., Jacmal Bunhian, 24 km E. of Mayoyao, 800-1000 m, light-trap, 27-29.iv.1967 (Torrevillas); 1 & Mindanao, Mt Balatukan, 15 km SW. of Gingoog, 1000-2000 m, at light, 27-30.iv.1960 (Torrevillas); 1 9, M., Del Sur, 32 km NW. of Milbuk, 900 m, rain forest, light-trap, 6.viii.l958 (Milliron) (all in BPBM, Honolulu). Malaya: 2 £, 1 ?, Gomak For. Res. near Kuala Lumpur, 28.xii.1958 (Quote); 1 & 2 ?, Pahang, Kuala Tahan, 15-16.xii.1958 (Quate); 3 rf, 2$, Kuala Lumpur, Klang gates, 31.xii.1958 (Quate) (all in BPBM, Honolulu); 1 specimen (abdomen missing), Kuala Lumpur, at light, 6.ii.l931 (Pendlebury); 3 9, West Coast, P. Angsa, Lt. Ho., at light, 26-27.X.1926 (Seimund); 1 <J, Kedah, near Jitra, catchment area, 8.iv.l928 (Pendlebury); 1 cJ, Selangor, Kepong, 7.ii.l958, at light; 2 specimens (abdo- mens missing), Perak, G. Kledang, 795 m, 15.xi.1927 (Seimund) (all in BMNH, London). Singapore: 14 J, 5$, Mandaj, mangrove, 7.ix.l977 (Murphy) (BMNH, London). Borneo: 1 £, Sarawak, Paya Paloh, at light, 4.xi.l964 (Rothschild); 2 9, Sarawak, Mt Dulit, 22 m, moss forest, light- trap, 28.x. 1932 (Hobby & Moore); IJ, Sarawak, Gunong Mulu Nat. Park (Collins); 1 ^, Sarawak, Gunong Mulu Nat. Park, base camp, at light, v.1978; 1 9, Sarawak, Kalabit End (Mjorberg) (all in BMNH, London); 1 <J, Sarawak, Sarikei Dist., Rejang delta, 15-25.vii.1958 (Maa); 1 £, Sarawak, Matang, m.v. light-trap, 13.ix.1958 (Gressitt) (both in BPBM, Honolulu); 1 <$, Sabah, Kalabakan, light-trap, 10-19.xi.1958 (Quate); 1 $, Sabah, Kalabakan, primary forest, ll.xi.1958 (Maa); 1 9, Sabah, Tawau, Quoin Hill Cocoa Res. Sta., 20.viii.1962 (Hiroshima); 1 9, Sabah, Tawau, Quoin Hill, light-trap, 1 5-20. vii. 1962 (Holtman) (all in BPBM, Honolulu); 1 9, Sabah, R. Kara- muak, 7 m SSE. Telupid, 60 m, l-7.ix.1977 (Bacchus) (BMNH, London). Java: 1 & Batavia, xi.1908 (Jacobson) (RNH, Leiden) (paralectotype of Muirella longiseta Melichar).
Parohinka malayensis sp. n.
(Figs 144-148)
Length: <$, 6-2 mm; 9, 6-9 mm.
Sordid to brownish yellow; vertex and dorsal region of face mottled with brown; other markings as in generic description.
External characters as in morona but vertex longer, medial length 3-2 times length next to eyes and 1-2 times basal width between eyes.
Male genitalia similar to longiseta but pygophore and Xth segment shorter, the latter extending only slightly beyond posterior margin of pygophore, connective with stem short rather than long and aedeagus with shaft more robust.
Female genitalia similar to longiseta with extended posterior corners of pregenital sternite short; second valvulae relatively narrow with first dorsal tooth large.
REMARKS. This species is closely related to longiseta but differs in its longer head and in the shape of the male genitalia, as noted above.
DISTRIBUTION. Known only from the type-locality in Malaya.
MATERIAL EXAMINED
Holotyped1, Malaya : West Coast, P. Angsa, Lt. Ho., at light, 1 1.x. 1926 (Seimund) (BMNH, London). Paratypes. Malaya: 1 cJ, 1 9, same data as holotype except, 12/1 6.x. 1926 (BMNH, London).
KAROSEEFA gen. n.
Type-species: Karoseefa brevipenis sp. n.
Yellow tinged with green.
Head wider than pronotum; anterior margin rounded in profile, transversely striate; ocelli on margin distant from eyes, not or slightly visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length slightly less than twice length next to eyes; sides
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 7 1
slightly convex; apex moderately angularly rounded; finely longitudinally striate, transversely striate an- teriorly. Face slightly wider than long, convex in profile, shagreen ; clypeus relatively short and broad, lateral margins slightly constricted near antennae; clypellus moderately long, of similar width throughout length; lora large ; antennal pit deep with inner margin evenly rounded to clypeus ; antennal ledge slight ; antennae very long, extending to near apex of clavus. Pronotum approximately twice as wide as long, sides very short, without a carina; transversely striate, obscurely rugose anteriorly. Scutellum approximately equal in length to pronotum, shagreen, obscurely rugose posteriorly. Fore wings with three subapical cells, first subapical cell open, second and third subapical cells closed ; with a few additional veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of 10 fine setae distally on anterior surface; hind femur with apical setal formula 2+1-1-1 with first proximal seta slender.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore strongly incurved dorsally with a short apodeme on each side; pygophore lobes with an oblique internal ledge extending to posterior margin; lobes with several macrosetae and short fine setae. Xth segment fairly short, compressed dorsoventrally. Valve triangular. Subgenital plate elongate, triangular, evenly tapered to apex or with distal region digitate and lightly sclerotized; apex of basal lobe with a short acute prominence; outer margin of dorsal surface with several moderate to long fine setae; ventral surface with few short fine setae with or without numerous long fine setae along outer margin; apical region of dorsal and ventral surfaces with several short stout setae. Style moderately long with basal apophyses and lateral lobe prominent; apical process moderately long, curved ventrally and tapered to apex, crenulate dorsally or dentate ventrally; few sensory papilla dorsally, adjacent to lateral lobe. Connective Y-shaped, stem moderately long and narrow, lateral margins keel-like dorsally; arms short. Aedeagus with shaft short or long, narrow, tapered from base to apex and terminating in a pair of dorsally directed processes, gonopore small, situated apically; basal apodeme moderately long and narrow.
Female genitalia with posterior margin of pregenital sternite with one or two lobes each side of mid line; second valvulae united at first dorsal tooth (arrowed in Fig. 160), elongate, of similar width throughout length, without a basal prominence; dorsal teeth robust, unaligned, extending approximately one-quarter distance from apex to base of valvulae; dorsal sclerotized region elongate.
REMARKS. This genus can be distinguished from others of the subfamily by the broad clypeus and clypellus with sides parallel rather than concave. In the male genitalia it is similar to Dryado- morpha but the anterior margin of the pygophore is strongly incurved dorsally with a short apodeme on each side. The female genitalia have the second valvulae similar to those of Pa- rohinka and Rhutelorbus.
DISTRIBUTION. Borneo (Sarawak, Sabah). Key to species of Karoseefa
1 Aedeagus short with apical processes more or less parallel ; posterior margin of female genital
sternite with two lobes each side of mid line (Fig. 161) .... brevipenis sp. n. (p. 70)
Aedeagus long with apical processes divergent; posterior margin of female pregenital sternite with a single lobe each side of mid line (Fig. 163) divergent sp. n.(p. 73)
Karoseefa brevipenis sp. n.
(Figs 149-161)
Length: <$, 5-2-5-6 mm, mean 5-4 mm; 9, 5-5-5-8 mm, mean 5-6 mm.
Colour and external characters as in generic description.
Male genitalia as in generic description with apex of subgenital plate digitate and lightly sclerotized, apical process of style dentate dorsally and aedeagus relatively short with processes more or less parallel.
Female genitalia with posterior margin of pregenital sternite with two lobes each side of mid line, the more lateral lobe acute.
REMARKS. I have identified only specimens of uniform colour as this species, a female specimen from Sabah : N. of Kalabakan (BPBM, Honolulu) may also be this species but has the fore wings banded with brown.
In external appearance this species is almost identical to divergens and is sympatric with this species over its known range in Sabah. From divergens it can be distinguished by its slightly
M. D. WEBB
160
161
162
164
166
Figs 149-166 Karoseefa species. 149-161. K. brevipenis. (149) head and thorax, dorsal view; (150) same, lateral view; (151) face; (152) left subgenital plate and apex of left style, dorsal view; (153)^ genitalia capsule; (154) apex of left style, medial view; (155) left style, ventral view; (156) connective, lateral view; (157) same, dorsal view. (158, 159) aedeagus, lateral and posterior views; (160) second valvulae, lateral view; (161) $ pregenital sternite, ventral view. 162-166. K. diver gens. (162) left style, ventral view; (163) 9 pregenital sternite, ventral view; (164) valve and left subgenital plate, lateral view; (165, 166) aedeagus, lateral and posterior views.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 73
smaller size and female pregenital sternite with two lobes each side of the mid line. In the male genitalia brevipenis differs from divergens in having the apical region of the subgenital plate narrower, the ventral surface of the subgenital plate without long fine setae, the apical process of the style dentate and the aedeagus shorter with the apical processes parallel.
DISTRIBUTION. Borneo (Sarawak, Sabah).
MATERIAL EXAMINED
Holotype cJ, Borneo: Sarawak, foot of Mount Dulit, junction of R. Tinjar and Lejok, old secondary forest, at light, 29.viii.1932 (Hobby & Moore) (BMNH, London).
Paratypes. Borneo: 7 $, 10 $, same data as holotype, except 28.viii-4.x.l932; 1 9, Sarawak, Mount Dulit, moss forest, 1200 m, 2 1.x. 1932 (Hobby & Moore) (BMNH, London); 1 & Sabah, Bettotan, near Sandakan, 31.vii.1927 (BMNH, London); 1 ?, Sabah, Tawau, Quoin Hill, at light, 26-29. vii. 1962 (Holtmann) (BPBM, Honolulu).
Karoseefa divergens sp. n.
(Figs 162-166)
Length: $, 6-0 mm; ?, 6-5-6-7 mm, mean 6-6 mm.
Colour and external characters as in generic description.
Male genitalia as in brevipenis but subgenital plate evenly tapered to apex and ventral surface of subgenital plate with numerous long fine setae laterally. Style with apical process less robust and crenulate dorsally and aedeagus with processes divergent.
Female genitalia with posterior margin of pregenital sternite with a single lobe each side of mid line.
REMARKS. This species is almost identical to brevipenis externally and is sympatric with this species over its known range in Sabah. For differences between the two species see 'Remarks' under brevipenis (p. 71).
DISTRIBUTION. Borneo (Sarawak, Sabah).
MATERIAL EXAMINED
Holotype <J, Borneo: Sarawak, Gunong Mulu Nat. Park, Long Pala, 70 m, alluvial secondary forest, on batu canopy, at light, iii.1978 (Holloway) (BMNH, London).
Paratypes. Borneo: 1 $, Sarawak, Gunong Mulu Nat. Park, near Long Pala, 50 m, alluvial forest, understorey, at light, v.1978 (Holloway) (BMNH, London); 1 $, Sabah, Tawau, Quoin Hill, at light, 3-7.vii.1962 (Holtmann) (BPBM, Honolulu).
OCEANOPONA Linnavuori
Oceanopona Linnavuori, 1960: 300. Type-species: Oceanopona croceipennis Linnavuori, by original de- signation.
Yellow to stramineous; male with thorax and fore wing marked with brown.
Head wider than pronotum; anterior margin narrowly rounded in profile, transversely striate; ocelli on margin distant from eyes, visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approximately 1-7 times length next to eyes, sides slightly convex, apex narrowly angularly rounded; shagreen and obscurely rugose, transversely striate anteriorly. Face approximately as long as wide, shagreen; more or less straight in profile; clypeus elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; transclypeal suture indistinct; lora large; antennal pit deep with inner margin angularly rounded to clypeus, rim-like; antennal ledge slight; antennae very long, extending to near apex of clavus. Pronotum approximately twice as wide as long, sides very short, without a carina; finely transversely striate, obscurely rugose anteriorly. Scutellum approxi- mately equal in length to pronotum, shagreen. Fore wings with three subapical cells, first subapical cell open, second and third subapical cells closed. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of six fine setae distally on anterior surface; hind femur with apical setal formula 2+1+0.
Apodemes of male third abdominal segment ventral, reduced.
Male genitalia with anterior margin of pygophore slightly incurved dorsally, without apodemes; pygo- phore lobes with a slight oblique internal ledge, not attaining posterior margin, lobes with several macros- etae and short to moderately long fine setae. Xth segment moderately long, compressed dorsoventrally.
74
M. D. WEBB
178
Figs 167-178 Oceanopona croseipennis. 167, head and thorax, dorsal view; 168, same, lateral view; 169, face; 170, $ genital capsule, lateral view; 171, fore wing; 172, valve and left subgenital plate, dorsal view; 173, connective, lateral view; 174, left style, ventral view; 175, valve and left subgenital plate, ventral view; 176, connective, dorsal view; 177, 178, aedeagus, lateral and posterior views.
Valve triangular. Subgenital plate elongate, triangular, outer margin of dorsal surface with several short fine setae and a group of long fine setae from mid length to near apex ; ventral surface with several short fine setae. Style moderately long with lateral lobe and basal apophyses prominent; apical process moderately long, turned ventrally with sides parallel, crenulate distally; few sensory papilla ventrally adjacent to lateral lobe. Connective Y-shaped, stem moderately long, narrow with lateral margins keel-like dorsally; arms with apophyses relatively long. Aedeagus with shaft elongate, curved dorsally, evenly tapered to apex and terminating in a pair of dorsally directed processes; gonopore apical on posterior surface, small; basal apodeme moderately long, narrow.
THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 75
Female genitalia with second valvulae united at first dorsal tooth, fairly robust, slightly expanded distally, dorsoanterior prominence present; teeth fine, extending along approximately distal third of valvulae; dorsal sclorotized region moderately long.
REMARKS. This genus is similar externally to Dryadomorpha and Parohinka but with the vertex shagreen and obscurely rugose and the setal formula of the hind femur 2+1+0. The male genitalia are similar to those of Dryadomorpha and Karoseefa but with the internal ledge of the pygophore lobes not extending to the posterior margin and the connective with the apophyses of the basal arms relatively long. In the female genitalia the second valvulae are similar to those of Dryadomorpha but with the teeth extending along approximately the distal third of valvulae.
The specimen of Oceanopona recorded by Evans (1966: 247) from Australia is identified as Dryadomorpha metrosideri (Osborn).
DISTRIBUTION. Eastern Caroline Is.
Oceanopona croceipennis Linnavuori
(Figs 167-178) Oceanopona croceipennis Linnavuori, 1960: 301. Holotype cJ, CAROLINE Is. (BPBM, Honolulu) [examined].
Length: <$, 4-4-4-7 mm, mean 4-6 mm; 9, 5-9 mm.
Yellow or stramineous. Male with pronotum brown posteriorly; scutellum turning to brown anteriorly with basal triangles darker brown ; legs heavily marked with brown ; fore wings golden to yellow, sometimes tinged with green, with a broad dark brown band along clavus, on apical region of subapical cells and becoming slightly paler on apical cells and appendix. Female variably tinged with green, without brown markings.
External characters as in generic description.
Male genitalia as in generic description with lateral lobe of style short and angularly rounded in ventral aspect. Aedeagus with apical processes divergent, short.
Female genitalia with posterior margin of pregenital sternite shallowly concave.
DISTRIBUTION. Eastern Caroline Is.
MATERIAL EXAMINED
Oceanopona croceipennis Linnavuori, holotype (J, Caroline Is.: Ponape, Mt Tamatamansakir, 180 m, 17.U953 (Gressitt) (BPBM, Honolulu).
Caroline Is.: 4 & 1 ?, Ponape, Mt Tamatamansakir, 180 m, 15-19.U953 (Gressitt) (BPBM, Honolulu) (paratypes of Oceanopona croceipennis Linnavuori); 1 <$, Kusaie I., Matanluk, Yepan, 23.U953 (Gressitt) (BPBM, Honolulu) (paratype of Oceanopona croceipennis Linnavuori).
References
Bruner, L. 1908. The Acridiidae. Biologia cent.-am. Insecta. Orthoptera 2: 1-342.
Distant, W. L. 1917. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership
of Mr. J. Stanley Gardiner, M.A. Vol. VI, No. vii.-Rhynchota, Part II: Suborder Homoptera. Trans. Linn.
Soc.Lond. 17: 273-322. - 1918. In Blandford, W. T., The fauna of British India including Ceylon and Burma. Rhynchota 7.
Homoptera: appendix. Heteroptera: addenda. vii + 2 10 pp. London. DIabola, J. 1979. Neue Zikaden aus Anatolien, Iran und aus Sudeuropaischen Landern (Homoptera:
Auchenorrhyncha). Acta zool. hung. 25: 235-257. Evans, J. W. 1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera:
Cicadelloidea and Cercopoidea). Mem. Aust. Mus. 12: 1-347. Eyles, A. C. & Linnavuori, R. 1974. Cicadellidae and Issidae (Homoptera) of Niue Island and material from
the Cook Islands. N. Z. Jl Zool. 1 : 29-44. Gardner, J. A. 1916. Systematic paleontology of the Upper Cretaceous deposits of Maryland. Mollusca. Md
geol. Surv. stratigr. Mem. 1916: 371-734. Hamilton, K. G. A. 1975. Review of the tribal classification of the leafhopper subfamily Aphrodinae
(Deltocephalinae of Authors) of the Holarctic region (Rhynchota: Homoptera: Cicadellidae). Can. Ent.
107:477^98.
76
M. D. WEBB
Haupt, H. 1927. Homoptera Palestinae I. Bull, agric. Exp. Stn, Tel- Aviv 8: 5-43.
Ishihara, T. 1954. Homopterous notes. Sclent. Rep. Matsuyama agric. Coll. 14: 1-28.
Kirkaldy, G. W. 1906. Leaf-hoppers and their natural enemies. (Pt. IX. Leaf-hoppers — Hemiptera). Bull.
Hawaiian Sug. Pirs' Ass. Exp. Stn (Ent.) 1 : 271^479.
— 1907. Leaf-hoppers — supplement. (Hemiptera). Bull. Hawaiian Sug. Firs' Ass. Exp. Stn (Ent.) 3: 1-186. Lindberg, H. 1958. Hemiptera Insularum Caboverdensium. Commentat. biol. 19: 1-246. Linnavuori, R. 1960. Insects of Micronesia. Homoptera Cicadellidae. Insects Micronesia 6: 231-344.
- 1978. Revision of the Ethiopian Cicadellidae (Homoptera). Paraboloponinae and Deltocephalinae : Scaphytopiini and Goniagnathini. Revue zool. afr. 92: 457-500.
Matsumura, S. 1912. Die Acocephalinen und Bythoscopien Japans. J. Coll. Agric. Hokkaido imp. Univ. 4:
279-325. Melichar, L. 1914. Homopteren von Java, gesammelt von Herrn Edw. Jacobson. Notes Leyden Mus. 36:
91-147.
Merino, G. 1936. Philippine Cicadellidae (Homoptera). Philipp. J. Sci. 61 : 307^400. Osborn, H. 1934. Cicadellidae of the Marquesas Islands. Bull. Bernice P. Bishop Mus. 114: 239-269. Pruthi, H. S. 1930. Studies on Indian Jassidae (Homoptera). Part I. Introductory and description of some
new genera and species. Mem. Indian Mus. 11 : 1-68.
- 1934. Entomological investigations on the spike disease of sandal. (14) Jassidae (Homopt.). Indian Forest Rec. 19:1-30.
Uhler, P. R. 1896. Summary of the Hemiptera of Japan presented to the United States National Museum by
Professor Mitzukuri. Proc. U.S. natn. Mus. 19: 255-297. Webb, M. D. 1980. The Cicadellidae from Aldabra, Astove and Cosmoledo Atolls collected by the Royal
Society Expedition 1967-68 (Hemiptera, Homoptera). J. nat. Hist. 14: 829-863.
Index
Invalid names are in italics; references to descriptions are in bold.
anacryon 40, 50 antennalis 40, 50 apicalis 58, 59, 66, 67, 68 australis 51
brevicephala 58, 59, 64, 66, 68 brevipenis 70, 71, 73
Calotettix 39, 49 camphorae 39, 40, 48 chatterjeei 40, 50 chinensis 43, 45, 46 croceipennis 39, 73, 75
divergens 71, 73 Dryadomorpha 40, 42, 49, 50,
57 dulita 58, 59, 64, 66, 67, 68
falcata 40
Favintiga 40, 42, 47 fraternus 40, 50
guttata 39, 43, 45, 46
indicus 40, 50 ishihari 43, 45, 46
Khamiria 49
Karoseefa 42, 50, 58, 70, 73
lais 39
longiseta 40, 57, 58, 59, 64, 66,
67, 68, 70 lotophagorum 40, 58, 59, 61,
63,64 lugubris 50 luzonensis 43, 46
malayensis 58, 59, 68, 70 mangrovecola 50 merinoi 56, 57
metrosideri 39, 50, 53, 54, 75 modestus 39
morona 58, 59, 60, 61, 63, 64, 66, 67, 70
Oceanopona 39, 42, 58, 73, 75 Odmiella 40, 42 Osbornitettix 49
pacifica 50, 53 Paganalia 40, 49 pallida 40, 50, 54, 55 Parabolopona 39, 40, 42, 47 Parohinka42, 50, 57, 71, 73 philippina 58, 59, 66, 68 punctatus 40, 50
quadricornis 40, 50
recurva 58, 59, 67, 68 Rhombopsana 40 Rhombopsis 40, 49 Rhutelorbus 42, 50, 56, 58, 71 robustipenis 50, 54, 55 rubrolineata 39
sinuata 58, 59, 61, 63, 64 spinosa 58, 59, 63, 64 Stenomiella 40, 42 Stirellus 50 Stymphylus 39
tincta 39
trispicata 58, 59, 61, 63, 64
virens 40, 50 virescens 40 viridia 54 viridis 40, 50
Yakunopona 40, 49 yakushimensis 40, 51
Zizyphoides 40, 49, 50
British Museum (Natural History) 1881-1981
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A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea) and its allies.
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The Asian, Australasian and Pacific Paraboloponinae (Homoptera : Cicadellidae). A taxonomic revision with a key to all the known genera of the subfamily.
By M. D. Webb.
A revision ofPhyciodes Hiibner and related genera, with a review of the classification of the Melitaeinae (Lepidoptera : Nymphalidae). By L. G. Higgins.
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A revision of Phyciodes HUbner and relate genera, with a review of the classification c the Melitaeinae (Lepidoptera: Nymphalidae
L. G. Higgins
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ISSN 0524-6431 Entomology series
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A revision ofPhyciodes Hiibner and related genera, with a review of the classification of the Melitaeinae (Lepidoptera: Nymphalidae)
L. G. Higgins
Focklesbrook Farm, Chobham, Woking, Surrey
Contents
Synopsis 78
Introduction 78
Historical review 78
Abbreviations of depositories ........... 79
Type-material and the British Museum (Natural History) collections ... 79
Phyciodine genitalia and the identification keys 80
Examination of Phyciodine genitalia 80
Taxonomy 81
Phyciodini trib. n. 81
Tribal characters 81
Distribution 82
Wing patterns 83
Key to genera of Phyciodini 84
Phyciodes Hiibner 85
Phystis gen. n. 93
Anthanassa Scudder 94
Dagon gen. n 108
Telenassa gen. n. . 1 10
Ortilia gen. n .115
Tisona gen. n 120
Tegosa gen. n. .121
Eresia Boisduval .129
Castilia gen. n. .151
Janatella gen. n. .157
Mazia gen. n .159
Species incertae sedis . 160
Supplement on certain genera of Melitaeini ...... . 160
Gnathotriche Felder & Felder • 160
Gnathotrusia gen. n. ........ • 162
Higginsius Hemming • 163
Antillea Higgins .164
Review of the classification of the Melitaeinae ... . 164
Check-list of tribes, generic groups, genera