Meenoplidae of Taiwan (Homoptera: Fulgoroidea) 81

Meenoplidae of Taiwan (Homoptera : Fulgoroidea) Shun Chern Tsaur*, Chung Tu Yang* and Michael R. Wilson** Received, May 27, 1986 CONTENTS

Abstract ...... · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 83 Introduction ...... · . · · · · · · · · · · · · · · · · · · · · · · · · · · 83 List of meenoplid Species of Taiwan ...... 83 FIRST PART: General ...... · . · · · · · · · · · · · · · · · · · · · · · · 84 I. Adult Morphology ...... · · · · · · · · 84 Head ...... · ... · · · · · · · · · · · · · · · · · · · 84 Thorax ...... · · · · · · 85 Legs ...... · . · · · · . · · · · · · · · · · 86 Pregenital segments of abdomen ...... 86 Male genital segments ...... 86 Female genital segments ...... 86 II. Immature Stages ...... 87 Egg ...... 87 Nymphs ...... 88 Key to nymphal stages ...... 93 SECOND PART: Systematics...... 93 Family Meenoplidae ...... · .. · · . 93 Key to subfamilies ...... 94 Subfamily Kermesiinae ...... 94 Key to genera ...... 94 Genus Nisia Melichar ...... · · · · · · · · · 94 Key to species ...... 95 N. carolinensis Fennah n . status ...... · · . 95 N. australiensis Woodward n. status ...... 96 Genus Eponisiella Emeljanov ...... · · . 98 E. matsumurai n. sp...... 99 Key to matsumurai and guttulinervis ...... 101 Genus Eponisia Matsumura ...... 101 Key to species ...... 101 E. woodwardi n. sp...... 101 E. guttula Matsumura ...... 103 E. macula n. sp ...... · .. 106 Subfamily Meenoplinae ...... 108 Key to genera ...... 108 Genus Metanigrus n. gen ...... 108 M. y ami n. sp...... 109 Genus Anigrus St§l ...... 111 Key to species ...... 111 A. formosanus (Matsumura) ...... 111 A. frequens (Matsumura) ...... 113 Acknowledgements ...... 116 References ...... 116

*Department of Entomology, National Chung Hsing University. ** Commonwealth Institute of Entomology, London, UK. 82 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

M. R. Wilson

-~~~Jll~H-~Jtl;,.{JWfi~ =5lH · 1. Nisia •······-~ae&•·-fi =·~-~· ·*Xtt~=5lHEmA• ae••~-•ff~ 0 z~~~=•zff~·~*~-~•·~ 2. Eponis. iella•······-~· o

0 ~• · -~ae&• · ~liae&f.i 3.Eponisia •······=~•·-Bae& *XEJf1i:z-~Ji8t~ : -~--ff~O I. Kermesiinae 5lH······ =Ji& • -=-~­ II. Meenoplinae5l.fl · · · · ·=• • -~- • ' -~ae&• =Bae&f.i 0 ·=Bae&• 1. Metanigrus •······~~-, -~- o · =·ff~O 2. Anigrus •······=Bae&• 0 Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 83

ABSTRACT

Only 4 meenoplid species belonging to 3 genera, 2 subfamilies recorded in Taiwan pre­ viously. The present paper deals with 9 species of adults and 2 species of nymphs which represent 2 subfamilies, 5 genera. (1) The subfamily Kermesiinae includes 3 genera, 3 new species, 2 known species, 1 newly recorded, and 2 species of nymphal stages. (2) Meenoplinae contains 2 genera, 1 new species, 2 known species. Among the total amount of 9 species, 2 genera and 4 species are new to science, 1 species is newly recorded in the fauna of Taiwan, and 4 species are known. Keys are given to the subfamilies, the genera and the species of adults. New genus : Metanigrus Tsaur and Yang. New species: Eponisia macula Tsaur and Yang, Eponisia woodwardi Tsaur and Yang, Eponisiel/a matsumurai Tsaur and Yang. New synonymy: Anigrus nigricans (Matsumuras) and A . metalces Fennah are reduced to Syns. of A. frequens (Matsumura). Nisia australiensis Woodward and N. carolinensis Fennah are new status.

INTRODUCTION recorded from Taiwan (Matsumura, 1914). This paper deals with 9 species of adults The Meenoplidae is a small family of and 2 species of nymphs which represent 5 fulgoroids , containing IO genera and 105 genera, among them 4 species and 1 genus are species, included in 2 subfamilies : Meenoplinae new to science. Two species (Nisia australiensis Fieb., 1872 and Kermesiinae Kirk., 1906. On & N. carolinensis) have been noted as minor evidences so far obtained, they belong to a rice pests, other are found on herb. Specimens group which include Achilidae, Achilixiidae, are deposited at the Department of Ento­ and Kinnaridae. The fundamental characters mology, National Chung Hsing University, shared by this group are (1) a simple egg, Taichung, Taiwan, and National Taiwan Univer­ (2) a cryptic nymphal life, (3) a reduced or sity, Taipei, Taiwan, Republic of China and obsolete ovipositor, ( 4) a tubular phallobase British Museum (Natural History), London, UK. and a greatly reduced or obsolete phallus, (5) All the meenoplid material available for study a long second post-tarsal joint, (6) a rostrum is collected by the senior author except other­ with a long apical segment, (7) a primitive wise stated, in various areas of Taiwan. tegminal venation (except in a ~ew very special genera) (Fennah, l 950a). They were formerly LIST OF MEENOPLID SPECIES included as a subfamily of Cixiidae but also OF TAIWAN included with Derbidae. They are confined to the Old World and most species are found in I. Subfamily Kermesiinae the tropics and subtropics. 1. Nisia carolinensis Fennah, 1971 n. status This family was established by Fieber in 2. N. australiensis Woodward, 1957 n. 1866 to include a single species from Greece . status Since that time the list of genera and species '. Eponisiella matsumurai Tsaur and Yang has been increased by the work of Kirkaldy n. sp. and Muir principally in the Pacific area; by 4 . Eponisia woodwardi Tsaur and Yang n. Distant, Muir, and Jacobi in the Ethiopian sp. region ; by Distant and· Melichar in India; by 5. E. guttula (Matsumura) Kirkaldy and Woodward in Australia; by 6. E. macula Tsaur and Yang n. sp . Synave in Africa ; by Linnavuori in USSR; by Fennah in various areas; by Stalle in Belgium; II. Subfamily Meenoplinae and by Matsumura in Taiwan and Japan; and 7 . Metanigrus yami Tsaur and Yang n . sp. now being revised by Wilson . 8. Anigrus formosanus (Matsumura) Three genera and 4 species (we agree with 9 . A. frequens (Matsumura) Emeljanov, 1984, excluding Kotonisia Mat­ (=A. nigricans (Matsumura)) sumura, 1938, from Meenoplidae) had been (=A . metalces Fennah) 84 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

FIRST PART: GENERAL taxonomy; some of these are found to be taxonomically useful and dis cussed below. I. Adult Morphology Meenoplidae are small Homoptera, being Head: (Fig. 1, A-D) 2.5 mm to 6.5 mm in size. Their coloration are various from white to brown. It has been The head usually small and connected with fairly easy to investigate a number of external the pro thorax at the posterior border, opis­ characters not previously used in meenoplid thognathous with strongly convex eyes . Vertex

E E "'0

Fig. 1. A. Head of Anigrus frequens. dorsal view . 1 v =length of vertex; wv=width of vetrtex; pa= posterolateral areolets; mc=median carina; pn=pronotum; mn=mesonotum. B. Head, ventral view ; f=frons ; mo= median ocellus; mcc =median carina of clypeus. C. Head, lateral view . D. Rostrum. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 85

small, usually forming two posterolateral lateral carina of frons and clypeus. Postcly­ areolets at base except on part of Kermesia peus with a distinct median carina except in and some Eponisia the latter having their Eponisia macula and Eponisiella . Lateral submedian carinae straight, not touching pos­ carinae of postclypeus are absent in Nisia and terior margin of vertex. In dorsal view, apical Eponisiella. Rostrum slender, reaching over margin of vertex again produced forward hind coxae, 3-segmented, first the longest, medially in species with median carina, strongly second longer than third, third longer than emarginate in species without. In profile vertex wide . Ocelli present. The antenna 3-segment­ roundly running into frons; frons convexed ed, the first segment short and thick, the medially, lateral carina dividing a large area second long and subcylindrical, the third, the around eye. Frans longer in middle line than flagellum , very thin. wide at widest portion, ranging from 1.3 : 1 to 2.3 : 1. Lateral carinae strongly elevated, Thorax: (Fig. 2, A-B) along its inner side has 22-34 sensory pits (including several ones on vertex). Median Pronotum, broader than head, arched, with carina present in Anigrus and Phaconeura, and median carina. Slightly convex on disc of an­ absent in other genera. In some species, an terolateral area in some species. Mesonotum interruption at frontoclypeal suture between with median carina, thumb-like , with large

It _ ___,

2mm

Fig. 2. A. Tegmen of Eponisia guttula (Matsumura) 1 t= length of tegmen ; wt=width of tegmen. B. Wing. 86 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

tegulae connected with pronotum, and as characters at least at species level. But in Nisia broad as or slightly broader than pronotum. the shape and color of pores are uniform Metanotum covered by mesonotum. Tegmina throughout. On the lateral side sternites I and always macropterous and normally helding VIII reduced in both sexes. Sternites II and III vertically in repose. The venation in general fused, only distinguishable in male in cleared cixiid-like. Sc+ R and M with common petiole specimens. Sternites IV-VII are all stout and at base , and forked before middle or after. A easily visible. Spiracles I, II and VIII are located line of sensory pits below Sc+ R and apical dorsally, on the external paratergites. Spiracles portion of common petiole or on above or III-VII can be found in the lateral membrane, not at all. Sc and R with two branches respec­ arranged nearly at same level. According to the tively, M 1+ 2 forked or not, M 3 + 4 fused with position of spiracles it is very easy to point out

Cu 1a two times. Cu 1b fused or slightly con­ which tergite and sternite belong to. Lateral nected with Cu 1a once. Claval veins fused at membranes have depressions anteriorly which middle in Meenoplinae, near apex in Kermesi­ outer part with granulates and tiny process, inae, entering the apex of the clavus. First inner impoverished. The cave, maybe some claval vein with single row of sensory pits in kind of proprioceptor, receive the stout process Meenoplinae, double in Kermesiinae. Many of hind leg (Fig. 4, A-C). sensory pits are scattered on the position of Male genital segments second claval vein in Meenoplinae , remainder without. Wings fan-like, hyaline, Sc+ R forked These are entirely symmetrical. Segments near middle . Sc curved upward, R usually IX and X fused respectively, forming pygofer forked except in Nisia and part of Eponisia. and anal segment, all ring-shaped. Pygofer is r-m present. M 1+ 2 forked. M 3+4 and Cu 1a wider posteriorly than anteriorly, connecting fused throughout. Cu 2 single . 2A forked . with anal segment tightly. Anal segment sclero­ tized, anal style arising from dorsomedial Legs: (Fig. 3, D) portion. The shape of anal segments is diverse Legs simple, the fore and middle legs small. in different genera. In dorsal aspect, slender Hind tibia long and slender, without spines and same width in Nisia, widened at middle in laterally, with basal segment of hind tarsus Eponisia , stout in Metanigrus, deflexed obtuse­ longer than the second and the third together. ly in Anigrus. Spinal formula of hind leg useful for separating Phallus simple and located on the central genera. portion of pygofer, usually with a ventral apodeme connecting with base of genital styles Pregenital segments of the abdomen: (Fig. 3, and a anterior apodeme lying straightly. Phallo­ A-C) base swollen. Processes of phallobase are with Tergite I visible on the dorsal aspect but various types. In lateral view, they are sickle­ very narrow and very weakly sclerotized. shaped in Nisia , irregular in Eponisia and Tergite II found modified conically on the Anigrus, slightly dilating with reverse teeth in dorsal in the male. In both sexes, tergite III Anigrus formosanus, swelling with scale-like widened ventrally. Tergites IV-VIII easily processes in Metanigrus . Genital styles slender, distinguishable. In the female, tergites VI-VIII anterior margin of basal half produced dorsad angulated medially, forming two declivous in a convex lobe which gives off distally a planes, the backward bearing wax-secretion upward spinose process in Nisia and Eponisia, plates (Fig. 4, A-C). The plates usually wide at in other genera curved upward at middle with middle and narrow laterad. Each plate usually a process near middle or apex. separated into two parts by a line of empty area Female genital segments (Fig. 3, C) or the arrangement of marginal pores. Pores of inner plate may be usually darker and larger. Ventral valvifer large and quadrate , in The shape and the size of inner plate are useful Meenoplinae its basocaudal angles turned mesad Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 87

IV v VI - ·-- .. VII A ----- VIII

0 .5mm D

·, .. I I. I

I \ \ \.

Fig. 3. Abdomen and hind leg of Nisia carolinensis Fennah. A. Male abdomen, lateral view; sp=spiracle. B. Pro­ prioceptor. C. Female abdomen, lateral view . D. Hind leg. and produced into a stout process but reduced by a visible membraneous area. in Kermesiinae. Ventral valvula of Meenoplinae except Metanigrus also quadrate and large, IT . Immature stages most portion covered by production of ventral Egg: (Fig. 6 , N-I) Only observed in N. caro­ valvifer with a long process slightly curved linensis, N. australiensis and Eponisia mesad. Shape of ventral valvula in Kermesiinae guttula. and Metanigrus bird-head-like with its process beak-shaped, connected with ventral valvifer Egg ellipsoidal, devoid of ornamentation. 88 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

Fig. 4. Abdominal tergites VI-VIII of Eponisia guttula (Matsumura). A. Tergite VI, dorsal view. B. Tergite VII, dorsal view; mr=median ridge; iwp=inner wax-secretion plate; owp=outer wax-secretion plate. C. Tergite VIII, dorsal view.

Uniformly translucent in Nisia carolinensis, Body length 0.45-0.53 mm, width 0.28- Nisia australiensis and yellowish white in 0.30 mm. Eponisia guttula immediately after laying. Grayish, with abdomen grayish white to Because the female ovipositor is reduced or yellowish white. Body robust in outline, with absent, the eggs are laid on the undersurface thorax and abdomen prominent. Compound of a leaf or in a depression on the stem and eye small and reddish. covered with a thin layer of wax threads, which Head: are excreted from abdominal wax-secretion pores and applied to the egg mass by using the In dorsal view vertex small, with lateral hind tibiae. In Nisia carolinensis about 0.4 mm carinae and elevated median carinae fused in length and 0.3 mm in width. The arrange­ medially at anterior margin , in lateral view ment of eggs in Nisia as Fig. 61, irregular in strongly rounding into frons . The frons with Eponisia. Numbers are from 10 to 25, average two median carinae, convex medially, so that 18 in Nisia, about 27 in Eponisia. Duration of the space between them-the in terfrons- is egg-laying is about half an hour, of the egg stage considerably broader than the lateral areas­ to hatching is about 10 days in summer 20 days laterofrontes medially. The interfrons is in winter. broadest in the middle and become narrower upwards and downwards. Laterofrontes same First instar nymph (Fig. 6, A, E, J) width throughout, each side with 13-14 This and following instar nymphs base on sensory pits arranged in two rows. Ocelli N. carolinensis only. absent. Frontoclypeal suture is slightly convex Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 89

A v

o.2mm 0.3rrlm

v B /[)Vf

0.3mrt1 o.1mm

Fig. 5. Female genitalia. A- B. Meenoplinae; C- D. Kermesiinae. v=Ventral valvula; vf=Ventral valvifer. upward. Clypeus triangular, with steep sides, Thorax: reaching up to convex portion. Postclypeus without carinal margins, with disc flat in ventral Pronotum with prominent anterolateral view but evenly curved in lateral view. Rostrum margin angulating towards side of body. Lateral slender, 3-segmented, reaching slightly over carinae obscure, wide in dorsal view, running hind coxae. Antennae 3-segmented, basal laterad as lateral margins, each side with 11 segment wider than long, the second longer sensory pits. Each inner side with 1 sensory pit than the first , with many sensory organs slightly below anterior margin, 5 medial along company with long setae and tiny processes, posterior margin. Outer 1 at middle, 1 sets at flagellum thin and short, 11-segmented, with anterolateral angle, 3 at posterolateral angle, the first segment dilate ball-like and the last hind margin of pronotum straight. Lateral flame-shaped, wider than long. margins reaching ventrally nearly to base of 90 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

fore coxae. Coxae of fore legs widely sepa­ mately each other, without meracanthae. Inner rated. Fore tarsus 2-segmented, each bearing 2 side of trochanter with about 7 keels. Hind simple claws and a arolium. femur curved dorsally from trochanter, angu­ Mesonotum small , without lateral carina, late. Hind tibia slender, with hind tarsus 2- hind margin straight, without visible wing-pad , segmented, the second longer than the first , each side bears 4 sensory pits, 1 near midway, with two claws and a arolium. Spinal formula 2 lateral. Mid-legs similar to fore legs except of hind leg 4-3-1. larger. Abdomen: Metanotum similar to mesonotum, includ­ ing number and arrangement of sensory pits, Abdomen consists of 9 visible segments, except narrower. Hind legs with coxae approxi- without wax-secretion opening. Segments

0 .3mm 0 .5mm

83 L

00 .3mm N 'l

Fig . 6. Nymphs of Nisia carolinensis fennah. A. First instar, dorsal view; B. Second instar, dorsal view· C. Third instar, dorsal view; D. Fourth instar, dorsal view; E-H. Thorax, flat surface. E. First instar; F. Second instar; G. Third instar; H. Fourth instar; I. Arrangement of eggs. J- M. Antenna; J. First instar; K. Second instar; L. Third instar; M. Fourth instar; N. Egg. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 91

broadest on tergite IV, gradually reducing to Laterofrons with about 19 sensory pits . both ends. Tergites I-III without sensory pits. Flagellum of antenna with its each segment thin Tergites IV- VIII each side with 2 sensory pits. and slender, longer than in third instar nymph. Tergite IX without sensory pit. Each side of tergal pit configuration of thoracic segments 20-17- 5. Two keels present on each Second instar nymph (Fig. 6, B, F, K) side of mesonotum, 1 lies at mi.ddle sinuate, Body length 0.56-0.75 mm, width 0.45- another slightly below ending of lateral carina 0.50 mm. of pronotum. Fore wing-pad developed, Differs from the first instar nymph as covering half length of lateral margin of hind follows: wing-pad. Hind wing-pad reaching to abdo­ Laterofrons with about 16 sensory pits. minal tergite II. Hind coxae each with a Rostrum reaching to sternite VI. Flagellum of meracanthus. Trochanter of hind leg with Antenna 13-segmented, slightly longer than in about 12 keels. Hind tarsus 3-segmented. first instar. Each side of tergal pit configura­ Spinal formula of hind leg 8-7-5. Tergite IV _tion of thoracic segments 14-7-4. Pronotum each side with 4 sensory pits posteriorly. with lateral carinae distinct, fine. Lateral Tergites V- VI each side with 2 anterior, 5 carinae of meso- and metanotum present. Fore posterior. Tergite VII each side with 1 an­ tarsus 2-segmented. Trochanter of hind leg teriorly, 5 posterior. Tergite VIII each side with 9 or 10 keels. Spinal formula of hind leg with 1 near middle, 3 ventral. 5-4-2. Tergites IV-VII each side with 3 Fifth instar nymph (Fig. 7) sensory pits laterally. Body length 1.55-2.03 mm, width 1.03- Third instar nymph (Fig. 6, C, G, L) 1.48 mm. Body length 0.95-1.03 mm , width 0.60- Body white to grayish white , stout, very 0.63 mm. broad across developing wing-pads . Differs from the second instar nymph as Head: follows: Laterofrons with about 18 sensory pits. In dorsal view vertex small, with lateral Rostrum reaching to sternite IV. Flagellum of carinae and elevated median carinae fused antenna 12-segmented, width of each segment medially at anterior margin. In lateral view about 2-3 times as long as wide. Each side of vertex rounding into frons . Frons longer in tergal pit configuration of thoracic segments middle line than wide at widest portion, about 17-13-4. Fore wing-pad slightly developed, 1.3 : 1, widest at level of lower margin of eyes . not reaching half length of metanotum. Hind Interfrons wider than laterofrontes in middle, wing-pad also slightly developed. narrower at base and apex . Laterofrons nearly Tergites IV, VII each side with 1 medial, same width throughout, each with about 20 3 posterolateral sensory pits. Tergite V-VI sensory pits arranged in two rows . Numbers of each side with 1 anterior, 4 posterolateral sensory pits of outer row about two times as sensory pits. Tergite VIII each side with 1 near inner one. Lateral ocelli absent. Median ocellus middle, 2 ventral sensory pits. Trochanter of present. Frontoclypeal suture almost straight. hind leg with 11 keels. Spinal formula of hind Clypeus triangular, with steep sides reaching up leg 6-5- 3. to convex central portion. Postclypeus without carinal margins, with disc flat in ventral view Fourth instar nymph (Fig. 6, D, H, M) but evenly cuvered in lateral view. Rostrum Body length 1.02-1.47 mm, width 0.64- slender, 3-segmented, reaching over hind coxae, 0.97 mm. relative length of each segment about 1.1 : 1.0 : 1.2. Antenna 3-segmented, basal segment wider Differs from the third instar nymph as than long, the second longer than the first follows : about 2.2 : 1, with about 13 sensory organs 92 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

E

E E M 0

Fig. 7. Fifth instar nymph of Nisia carolinensis Fennah. A. Frons and clypeus, ventral view; B. Vertex, dorsal view; C. Thorax, flat surface; D. The same, dorsal view; E. Antennae; F. Hind leg; G-1. Genitalia. G. Dorsal view; H. Lateral view; I. Ventral view.

company with long setae and tiny processes, along lateral carina. Two very small ones an­ flagellum very thin, long, 14-segmented. Com­ terior. Hind margin of pronotum is excavated pound eye large and purplish black. reverse· V-shaped. Lateral margins reaching ventrally to base of fore coxae. Coxae of fore Thorax: legs widely separated. Fore tarsus 2-segmented, Pronotum with prominent anterolateral distal segment longer than proximal and bears margin angulating towards side of body. At two claws apparently attached to long arolium. inner side of lateral carinae each with l 0 sensory Mesonotum very broad, with lateral carinae pits. At outer side, each with 9 sensory pits ar­ elevated. Fore wing-pads strongly produced ranged in semicircle, with another l separate, caudad, cover lateral margins of hind wing-pads, Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 93

but not reaching its posterior margin. Four or 0.90 mm long; wing-pads slightly or well five sensory pits present along each inner side developed ...... 3 of lateral carina, 5 between each lateral carina 2. Spinal formula of hind leg 4-3-1; lat­ and submarginal keel, above median keel. Two erofrons with 13-14 sensory pits; each pits between median keel and submarginal keel, side of tergal pit configuration of thoracic 7 between submarginal keel and lateral margin. segments 11-4-4 . . . . First instar nymph Mid-leg similar to fore leg, except longer. Spinal formula of hind leg 5-4- 2 ; lat­ Metanotum broad, with anterior margin erofrons with 16 sensory pi ts; each side straight hind margin excavate reverse V-shaped, of tergal pit configuration of thoracic lateral margins hidden by fore wing-pads lateral­ segments 14-7-4 .. Second instar nymph ly. Lateral carinae elevated, nearly parallel, 3. Second segment of hind tarsus with 3 teeth inner side with 1 sensory pit medially, outer apically; each side of tergal pit configura­ with 3. Hind legs with coxae approximate each tion of thoracic segments 17- 13-4 .. ... other. Inner side of hind trochanter with about ...... Third instar nymph 14 keels. Hind femur curved dorsally from Second segment of hind leg with 5 teeth trochanter. Hind tibiao slender. Hind tarsus apically ...... 4 3-segmented, the first segment longer than the 4. Spinal formula of hind leg 8-7-5; middle second, the third with claws apparently at­ ocellus absent; each side of tergal pit tached to large arolium. Spinal formula of hind configuration of thoracic segments 20- leg 10-9(8)-5. 17- 5...... Fourth instar nymph Spinal formula of hind leg 10- 9(8)- 5; Abdomen: middle ocellus present ; each side of tergal Abdomen consists of 9 visible segments, pit configuration of thoracic segments 22- without wax-secretion opening. Segments 19(18)- 4 ...... Fifth instar nymph broadest on tergite IV, gradually reducing to both ends. Tergites I-III without sensory pit. SECOND PART: SYSTEMATICS Tergite IV each side with 4-5 sensory pits; tergites V-VI with 4 anterior, 6 posterior; Family Meenoplidae Fieber, 1872 tergite VII with 1 lateral, 6 posterior; tergite VIII with 1 medial, 4 caudad; absent on tergite Type genus: Meenoplus Fieber, 1872 IX . Meenoplidae Fieber, 1872, Ka talog europais­ Nymphal specimens examined: chen Cicadinen : I-IV p. 3. 1st ins tar nymph: 5 Kermesiinae Kirdaldy, 1906, Bull Hawaiian 2nd instar nymph: 5 Sugar Pl. Assoc. Div. Ent. 1(9):425. 3rd instar nymph: 5 Nisiinae Kirkaldy, 1907, Bull. Hawaiian Sugar 4th instar nymph: 23 Pl. Assoc. Div. Ent. 3:163 . 5th instar nymph: 33 Meenoplidae Muir , 1925, Pan-Pacific Ent. 1 :97. NCHU Campus 6-V-1985. All the species, compared with other ful­ goroids, moderately small insects, with body Note: All 71 individuals were carefully ex­ strongly compressed and tegmina tectiform. amined by the senior author. Head usually small with vertex and frons usual­ ly broad and lateral carinae strongly elevated. Key to nymphal stages of Antenna general short and simple with many Nisia carolinensis sensory organs around second segment. Last l . Hind tarsus 2-segmented; body less than segment of the rostrum elongate . Median 0.85 mm long ; wing-pads not developed ocellus usually present. Thorax usually large ...... 2 with carinae generally distinct. Tegulae large . Hind tarsus 3-segmented; body more than Pronotum short and arched, broader than head. 94 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

Mesonotum as broad as or slightly broader than rated medially ; spinal formula of hind leg pronotum. Tegmen usually macropterous, 8-7-6 ...... Eponisia Matsumura normally held vertically in repose. Venation in Tegmen with M 1+ 2 simple; vertex with general cixiid-like, with basal costal area more submedian carinae and basal margin fused or less expanded. Radius usually simple, but medially ...... 2 cubitus complex, general with supernurnerary 2. Spinal formula of hind leg l 0-9(8)5; veins. One or both claval veins typically lateral carinae of postclypeus absent ..... granulate. Abdomen compressed, female with ...... Nisia Melichar six th, seven th, eighth tergi tes bearing three Spinal formula of hind leg 8-7-5 ; lateral areas of wax-secretion pores on each side carina of postclypeus present ...... respectively. Male genitalia superfically re­ ...... Eponisiella n. gen . sembles the genitalia in many derbids, achilids and achilixiids, with genital styles large and Genus Nisia Melichar, 1903 complex. Female genitalia incomplete, ovi­ Nisia Melichar, 1903 , Homopteren-Fauna von positor absent or greatly reduced. Ventral Ceylon 1903 : 53. valvula of female with process at apex, slightly curved mesad. Body small to medium, covered with thick powdery wax. Vertex narrowed medial­ Key to the subfamilies of Meenoplidae ly, with two posterolateral areolets at base. 1. Claval veins fused near apex, first claval Lateral carinae of frons ending at frontocly­ vein with two rows of sensory pits, second peal suture, absent at postclypeus. Clypeus slightly curved, without sensory pits; with median carina. Rostrum reaching over tegmen with Sc+R and M forked near base hind coxae. Pronotum with anterolateral ...... Kermesiinae disc convex, with distinct median carina. Claval veins fused near middle, first claval Mesonotum with distinct median carina. vein with one row of sensory pits, second Tegmen elongate, scarcely dilating towards

strongly curved with many sensory pits apex, with cross veins obscure , M 1+ 2 simple. arranged irregularly; tegmen with Sc+R Below Sc+R with single row of sensory pits. and M forked near apex .... Meenoplinae Wing with R branched. Spinal formula of hind leg 10-9(8)-5 (except N. grandiceps Subfamily Kermesiinae Kirkaldy Kirkaldy with 14-11-6). Male genitalia in (Fig. 5, C-D) dorsal view, anal segment elongate, lateral margins nearly parallel, lateroapical angles Kermesiinae Kirdaldy, 1906 Bull. Hawaiian obtusely rounded at apex. In lateral view , Sugar Pl. Assoc. Div. Ent. 1(9):425. phallus slender, slightly curved downward, Tegmen with Sc+ R and M forked near covered by glove-like process of phallobase. base. Sc with a row of sensory pits. Claval Genital styles elongate, in profile, strongly vein fused near apex, first claval vein with curved at middle, obtusely rounded at apex, two rows of sensory pits, second without. posterior margin convex near base, its anterior

M 1+ 2 single or forked. R of wing simple or margin of basal half produced in a convex lobe forked. Shape of ventral valvula like a bird (except N. grandiceps Kirkaldy and N. langlei head which with its process beak-shaped, Muir). Female genitalia in lateral view, ventral connected with ventral valvifer by a visible valvifer large, caudoventral angle without membraneous area. process. Ventral valvula small, dorsal and caudal margins evenly curved, ventral margin Key to genera of Kermesiinae distinctly incised medially, in caudal view with in Taiwan a small process. Shape of wax-secretion pores

1. Tegmen with M 1+ 2 forked ; vertex with on tergi tes VI- VII all small hexagon um, on submedian carinae and basal margin sepa- tergite VIII large , long hexagonum, all arranged Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 95

tightly. Color uniform, not forming inner and lateroapical angle obtusely rounded at apex. outer plates. Phallus with both sides sclerotized. In lateral view, reaching about as far as anal segment, Type species: Livi/la nervosa Motschulsky apex curved downward, appearing acuminate. ( =Meenoplus atrovenosus Lethierry, Apex of phallus at same level of ventral margin 1888) 1863 (original designation) of phallobase. Anterior apodeme of aedeagus almost in same level as aedeagus in side view. Key to the species of Nisia in Taiwan Process of phallobase bilobed, membraneous, slightly longer than phallus. In lateral view, 1. In lateral view, genital style at basal half convex lobe of genital style (Fig. 8E) gives off with lobe-like process which gives off distally a stout process dorsally, margin of lobe distally a stout process caudally, margin of smooth. In caudal view, divergent, inner lobe sinuate; in caudal view, inner margin margin of styles strongly produced mesad near of style with dorsad trifurcate ...... base, which with basal half convex, with two ...... australiensis Woodward processes apically. Outer one slender, needle­ In lateral view, genital style with lobe-like like, distinctly longer than inner, inner one process at basal half which gives off distally stout, thumb-like, obtuse at apex. a stout process dorsally, margin of lobe Female genitalia: Anal segment quadrate, slightly concave; in caudal view, inner anal style elongate, longer than anal segment, margin of style with dorsad bifurcate ... . the two combined longer than ventral valvifer...... carolinensis Fennah Ventral valvifer in caudal view with wart-like ventral valvula at inner side.

Nisia carolinensis Fennah n. status Length of body (includ. teg.) : Male 2.90- (Fig. 8) 3 .l O mm; Female 3.20-3.70 mm. Length of tegmen : Male 2.40-2.80 mm; Nisia atronervosa carolinensis F ennah, 1971, Female 2.40-3.30 mm . Insects Micronesia 6(8):584-585. Width of head (across eyes): Male 0.53- 0.57 mm; Female 0.53-0.60 mm. General color white. Vertex, frons, an­ Width of mesonotum: Male 0.70-0.73 tennae and pronotum pale yellow. Mesonotum mm; Female 0.77-0.87 mm. dark brown. Ocelli light yellow. Legs yel­ lowish. Tegmen white with veins fuscous Specimens examined: 11 do 2 99 NCHU brown. Abdomen brown. campus, Taichung l 7-IX-1984; 1 d In ventral view, frons longer in middle line 1 9 NCHU campus, Taichung 18-III- than wide at widest portion about 1.3 : 1. 1985; 3 do 3 99 NCHU campus, Tai­ Lateral carinae with about 27 sensory pits chung 7 ·IV-1985; 12 do 8 99 NCHU along each inner side (including several ones campus, Taichung 12-IV-1985; 4 do . on vertex). Postclypeus longer in middle line 4 99 NCHU campus, Taichung 16-IV- than wide at base, median carina obsolete at 1985; 1 d NCHU campus, Taichung basal third. Rostrum with relative length of 3-V-1985; Many specimens Lanshu, each segment about 1 : 1 : 1. Antenna with Taitung Hsien 9-VII-1985; 1 d 1 9 basal segment wider than long, second segment Lanshu, Taitung Hsien l 2-VII-1985 longer than first about 2 .5 : 1. Tegmina about L. C. Tang 1 d 4 99 Lanshu, Taitung 2.5 times as long as wide. Below Sc+R with Hsien 12-VII-1985 C. T. Yang. about 20 sensory pits. First claval vein with Host plant: Cyperus rotundt!S Linn. about 14 sensory pi ts on each side. (Cyperaceae), Oryza sativa L. (rice) Male genitalia: In dorsal view , anal seg­ Distribution: Micronesia, Philippines, ment with lateral margins nearly parallel, each Taiwan. 96 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

0.5mm

"'ci 0.3mm 0 ,3mm G K 0.1mm

Fig. 8. Nisia carolinensis Fennah. A. Frans and vertex, ventral view; B. Vertex, pronotum and mesonotum, dorsal view; C. Tegmen; D. Wing; E. Male genitalia, lateral view; F . Aedeagus, lateral view; G . The same, dorsal view; H. Male genitalia, caudal view; I. Female genitalia, lateral view; J. The same, ventrocaudal view; K. Male anal segment, dorsal view.

Nisia australiensis Woodward n. status Univ. Queensland Pap. (Dept. Ent.) 1(4): (Fig. 9) 67. Nisia atrovenosa australiensis Woodward, 1957 , General color brown. Mesonotum dark Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilso n 97

brown. Ocelli light yellow. Legs pale yellowish Female 3.30-4.10 mm. brown to brown. Tegmina yellowish white Width of head (across eyes) : Male 0.57 - with veins fuscous brown. 0.60 mm ; Female 0.63-0.67 mm. In ventral view , frons longer in middle Width of mesonotum: Male 0.90-1.20 line than wide at widest portion about 1.4: 1. mm; Female 0.80- 1.30 mm. Lateral carinae with 26-29 sensory pits along each inner side (including several ones on Specimens examined: 1 9 Songho, Tai­ vertex). Postclypeus longer in middle line than chung Hsien 2-Vll-1984; 1 d Puli, wid(f at base. Rostrum with relative length of Nantou Hsien 10-VIII- 1984; 25 dd each segment about 1.1 : 1 : 1.1 . Antenna with 12 99 Pingtung, 25-X-1984; 5 d basal segment wider than long, second segment NCHU campus, Taichung l 7- IX- 1984; longer than first about 2.2 : 1. Tegmen about 3 dd 2 99 Yuching, Tainan Hsien, 22- 2.0 times as long as wide. Below Sc+ R with I-1985 C. T. Yang. 1 9 Toubienkeng, about 21 sensory pits. First claval vein with 21-VI-1985; 7 dd 7 99 Peipu , Hualien about 17 , 14 sensory pits at above and below Hsien 12-Vll-1985; 37 dd 45 99 respectively. Fuyuan, Hualien Hsien 13- Vll- 1985; Male genitalia: Pygofer with dorsolateral 5 dd 3 99 Kaochung, Kaohsiung Hsien angles not produced. In dorsal view anal 22-Vll- 1985; Many specimens Fu­ segment with lateral margins parallel , each yuan, Hualien Hsien 10-VIII- 1985 ; lateroapical angle obtusely rounded at apex. 7 dd 8 99 Chushan, Nantou Hsien 26- Phallus with both sides sclerotized. In lateral X-1985 C. T. Yang. view, reaching about as far as anal segment, Host plant: Cyperus rotundus Linn. apex curved downward, appearing acuminate. (Cyperaceae) Commelina diffusa Burm. Apex of phallus at same level of ventral margin f. (Commelinaceae). of phallobase . Anterior apodeme of aedeagus Distribution : Australia, Philippines, Taiwan . almost in same level as aedeagus in side view. Process of phallobase bilobed, membraneous, Notes: This and former species, up to slightly longer than phallus. Genital styles now, were considered as two subspecies. Wil­ nearly same width, nearly straight. In lateral son (1981) had mentioned that "Two forms are view , convex lobe of genital style gives off found together in samples from the Philippines, distally a stout process caudally, margin of lobe with little evidence of intermediates." We have sinuate, anterior nearly at right angle. In caudal carefully examined specimens that were cap­ view, divergent, inner margin of styles strongly tured from NCHU campus and found also that produced mesad near base, which with dorsal · there were two forms without intermediates. margin trifurcate. Outer one slender, acu­ We therefore consider they are sympatric minate at apex, curved at basal half, almost species. Thirty-three individuals collected from reaching at same level of middle one . Middle a square meter were examined. They were all one stout, acute at apex. Inner one straight, N. carolinensis without any another individual directed medially. Inner margin of genital mixed. At Fuyuan, Hualien Hsien, a field left styles with apical two-thirds concave. uncultivated where it was difficult to find any Female genitalia: Anal segment quadrate, sedge, Cyperus rotundus (L.), more than 500 anal styles elongate , longer than anal segment, individuals were found on a square meter the two combined longer than ventral valvifer. (rough estimate). Ten male specimens were Ventral valvifer in caudal view with wart-like examined which were uniformly with N. ventral valvula at inner side. australiensis . We therefore have decided to raise them to full species rank. Nymphs of Length of body (includ. teg.): Male 2.70- those two species are so much alike and cannot 3.60 mm ; Female 3.60-4.70 mm. be separated by us . Length of tegmen: Male 2.30-3.00 mm ; Yang has fo und N. australiensis Woodward 98 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

G

Fig . 9. Nisia australiensis Woodward. A. Frons and clypcus, ventral view; B. Vertex, pronotum and mesonotum; C. Tegmen; D. Male genitalia, lateral view; E. Aedeagus, lateral view; F. The same, dorsal view; G-H. Genitalia. G. Male, ventrocaudal view; H. Female, lateral view; I. The same, caudal view; J. Male anal segment, dorsal view.

of Chiasien, Kaohsiung Hsien was feeding on dery wax. Head concave apically with a black Commelina diffusa Burm. f. (Commelinaceae) stripe medially. Vertex with two posterolateral on 24th, July 1986. They were living beside a areolets at base. Lateral carinae of frons ending stream and about 200 individuals were found in at frontoclypeal suture strongly convergent a square meter (rough estimate). We believe apically. Lateral carinae of postclypeus absent. that this plant must be one of its host plant. Pronotum without, mesonotum with a weak median carina. Tegmina broadened apically Genus Eponisiella Emeljanov, 1984 with M 1+ 2 not branched. Wing with R not Body very small, not covered with pow- forked. Spinal formula of hind leg 8-7-5. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 99

Male genitalia in lateral view pygofer with dor­ Emeljanov, 1984 in frons longer in middle line socaudal angle strongly produced. Female than wide at widest portion about 1.3 : 1 genitalia with ventral valvifer elongate, seemly (about two times in Emeljanov's description forming by produced, directly downward, which was not agree with Vilbaste's material) caudoventral angle nearly without process. also in shape of ventral valvifer. Ventral valvula bird-head shaped, apex with a process directed ven trocaudad. Eponisiella matsumurai* n. sp. (Fig. 10) Type species: E. guttulinervis (Matsumura) 1914, comb. n. General color white. Body not covered Nisia paludicola Vilbaste, 1968, syn. n. with powdery wax. Vertex, frons, antenna, (original designated) mesonotum pale yellow. Pronotum with three This species is not found in Taiwan but longitudinal blackish lines crossing over, one the opportunity is taken here to include the medial, two lateral, down posteriorly to synonomy and designating the lectotype. mesonotum. Legs yellowish. Abdomen black, Figures of guttulinervis may be found under with lateral sides yellow. Tegmina with black paludicola Vilbaste (1968:39). markings as figured. Frans widely expended, slightly longer Distribution: Japan, Korea, Soviet Mari­ in middle line than wide at widest portion. time Territory. Lateral carinae with about 30 sensory pits along Material examined: each inner side (including several ones on vertex). Clypeus without median carina. Loctotype: (here designated), 1 d (dis­ Postclypeus like a homebase plate, longer in sected) mounted on left of two speci­ middle line than wide at base about 1.3 : 1. mens on balsa wood block (1 damaged 0 Anteclypeus elongate. Rostrum reaching to remains undissected), Japan, Tawada, half length of hind femora, relative length of vii. 1905 (sapporo) (examined by Wil­ each segment about 1.1 : 1.4 : 1.0. Antenna son). with basal segment wider than long, second Paralectotypes: 2 do (1 mounted on block segment longer than first about 2.7 : 1. Central with lectotype). Japan, (Sapporo). portion of mesonotum with a week median Other material: series of Paludicola from carina. Spinal formula of hind leg 9(8)-7-5. Soviet Maritime Territory. Tegmen about 2.0 times as long as wide. Below Comments: Matsumura (1914) stated that Sc+ R with about 21 sensory pits. First claval 3 do were examined. These specimens were · vein with 19-20 sensory pits along each side. present in the Matsumura collection in Sapporo Wing with R not branched. Shape of wax­ and have been examined by Wilson. The secretion pores all hexagonum, on tergites VI­ synonomy has been made by direct comparison VII smaller, outer plate the smallest, on tergite with guttulinervis type material and Paludicola VIII outer plate large , all arranged tightly. specimens identified by Vilbaste. Male genitalia: In caudoventral view Notes: This genus from the absence of pygofer strongly concave medially, each side lateral carina of the postclypeus can be appro­ with a thumb-like, long process lateroapically. ximated with Nisia. But differs from other In lateral view anal segment stout, nearly same members of Kermesiinae in combination width, slightly angulate at middle, with stout characters as follows: Ventral valvifer elongate, anal style adherent to anal opening. In dorsal seemly forming by produced directly down­ view subquadrate, trifurcate apically. In lateral ward; spinal formula of hind leg; rostrum view phallus bifurcate, lower one longer than reaching hind femora. Differs from Eponisiella upper, reaching over anal segment, apex curved

* Named in honor of S. Matsumura, a Japanese entomologist. 100 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

, , E A .. .' in \v,ciE

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Fig. 10. Eponisiella matsumurai n. sp. A. Frons and clypeus, ventral view; B. Vertex, pronotum and mesonotum, dorsal view; C. Tegmina; D. Wing; E-F. Female genitalia. E. Caudal view; F. Lateral view. G-I. Male genitalia. G. Lateral view; H. Caudoventral view; I. Anal segment, dorsal view.

downward, appearing acuminate. Genital style ward. In caudoventral view, Ventral valvifer stout, same width throughout, evenly curved strongly curved upward near base, widely upward. In caudoventral view apical third divergent. Ventral valvula ellipse-shaped, at much narrower than the other part, with three middle of inner margin of ventral valvifer, with productions near middle. blade-like process at ellipsoidal top. Distance Female genitalia: In lateral view, anal style between ventral valvulae much wider than stout, anal segment small, both curved down- width of anal segment. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 101

view, covered by membraneous sheath, process Length of body (includ. teg.) : Male 2.93- of phallobase auricle-like. Genital style slightly 3.27 mm; Female 2.93-3.17 mm. bent angularly at middle. Anterior margin Length of tegmen: Male 2.43-2.73 mm; of basal half produced dorsad in a convex Female 2.93-3.17 mm. lobe, another process hidding after it. Female Width of head (across eyes): Male 0.60- genitalia in lateral view, ventral valvula beak­ 0.63 mm; Female 0.63 mm. shaped at apex, slightly longer than process of Width of mesonotum: Male 0.97-1.00 ventral valvifer. Lower margin of ventral mm; 0.93-1 .03 mm. valvifer produced caudoventrad forming a thumb-like process. Holotype: Male, Chintuan, llan Hsien l­ VIII-1985. Type species: Eponisia guttula Matsumura, Paratypes: 1 o Chishingshan, Taipei Hsien 1914. (original designation) 7-IX-1985; 3 od 3 99, Chihtuan, I!an Hsien 1-VIII-1986. (deposited 1d1 9 Key to the species of Eponisia in Taiwan in BMNH). 1. Length of body including tegmen 5.67- Host plant: unknown. 6.50 mm; genital style as long as anal Distribution: Taiwan. segment, equally divided into three rival Key to matsumurai and guttulinervis productions apically ... woodwardi n. sp. Length of body including tegmen less than 1. In caudoventral view, genital style with 5 mm; genital style long, distinctly longer three productions near middle; in lateral than anal segment, unequally divided into view ventral valvifer longest against widest three rival productions apically, ventral one portion about 1.9: 1 .. matsumurai n. sp. extremely long ...... 2 In caudoventral view, genital style only 2. Tegmina with veins white and other por­ with one production near middle; in tion pale black; female genitalia with lateral view ventral valvifer longest against process of ventral valvifer covered by widest portion about 1.2:1 ...... ventral valvula partially, nearly same length ...... guttulinervis (Matsumura) ...... macula n. sp. Tegmina pure white, opaque to somewhat Genus Eponisia Matsumura dark, subhyaline; female genitalia with process of ventral valvifer reaching two­ Eponisia Matsumura, 1914, Ann. Mus . Nat. thirds of ventral valvula, a deep gap be- Hungarici 12:285. tween them ...... guttula (Matsumura) Body median size, covered with or without podwery wax. Vertex without median carina, Eponisia woodwardi* n. sp. with areolets confluent medially, straight (Fig. 11) anteriorly and shallowly concave posteriorly. Median carina of frons absent. An interruption General color grayish yellow. Body more at frontoclypeal suture between lateral carinae or less covered with powdery wax. Vertex, of frons and postclypeus. Clypeus with median frons and antenna yellow. Mesonotum yel­ carina. Pro- _ and mesonotum with distinct lowish brown. Ocelli light yellow. Legs yel­ median carina. Tegmina broad, markedly diver­ lowish. Tegmen dirty yellow. Abdomen dark gent towards apex. Spinal formula of hind leg brown to black, with lateral side orange. 8-7-6. Male genitalia in dorsal view, anal Frans longer in middle than wide at widest segment stout, gradually expending to apex, portion about I .5 : I. Lateral carinae with shallowly emarginate at apex. Phallus in lateral 32-34 sensory pits along each inner side (in-

*Named in honor of Dr. T. E. Woodward, the Australian Hemipterist. 102 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

1mm

0.3mm

Fig. 11. Eponisia woodwardi n. sp. A. Frons and clypeus, ventral view; B. Vertex, pronotum and mesonotum, dorsal view; C. Tegmen; D. Wing; E. Male genitalia, lateral view; F . Aedeagus, lateral view; G. The same, ventral view; H. Male genitalia, caudoventral view; I. Female genitalia, lateral view; J . The same, caudal view; K. Male anal segment, dorsal view. eluding several ones on vertex). Postclypeus first about 2.8 : 1. Tegmen about 2.1 times as longer in middle line than wide at base. Ros­ long as wide. Below Sc+ R with about 20 trum with relative length of each segment about sensory pits. First claval vein with 20, 21 1 .1 : 1 .3 : 1 . Antenna with basal segment sensory pits on upper and lower sides respec­ wider than long, second segment longer than tively. R of wing not branched. Inner wax- Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 103

secretion plate of tergite VII with lateral half Host: plant: unknown. nearly as wide as median. Distribution : Taiwan. Male genitalia: Pygofer with dorsolateral angle not produced. In dorsal view , anal Eponisia guttula Matsumura, 1914 segment elongate, slightly expanding at apex, (Fig. 12) lateroapical angles each produced roundly. Eponisia guttula Matsumura, 1914 Ann. Mus. Phallus with both sides sclerotized, in lateral Nat. Hungarici 12 :285 . view much longer than anal segment, apex membranous, curved upward, at same level of General color white. Body covered with anterior apodeme of aedeagus. In lateral view , powdery wax. Vertex, frons and antenna process of phallobase cylindrically curved yellow. Mesonotum brown with three yel­ downward. Genital styles short and stout, lowish longitudinal stripes from anterior to posterior margin curved near base, anterior posterior. Ocelli light yellow. Legs yellowish. margin slightly produced medially, dorsal Tegmen in color from white, opaque to some­ aspect of convex lobe with many setae along what dark, subhyaline. Abdomen brown with margin, in caudal view divergent, inner margin lateral sides fresh yellow. of styles slightly produced subapically, divided Frans longer in middle line than wide at into three rival productions apically. Inner one widest portion about 1.7 : 1. Lateral carinae higher than the other two. with 25-30 sensory pits along each inner side Female genitalia: Anal segment quadrate. (including several ones on vertex). Postclypeus Anal style elongate. Ventral valvifer distinctly longer in middle line than wide at base. Ros­ produced caudoventrally, just reaching to base trum with relative length of each segment of beak, forming a gap between ventral valvifer about 1.3 : 1.3 : 1. Antenna with basal seg­ and ventral valvula. Ventral valvula like head of ment wider than long, second segment longer bird, as wide as long, beak curved downward than first about 2 : 1. Tegmen about 2.2 at apex. Ventral valvifer and valvula both with times as long as wide. Below Sc+ R with about many setae. 16 sensory pits. First claval vein with 22 sensory pits on each side . R of wing branch­ Length of body (includ. teg.): Male 5.67- ed. Inner wax-secretion plate of tergite VII 5.87 mm; Female 5.70-6.50 mm. with lateral half distinctly narrower than Length of tegmen : Male 5 .03-5 .27 mm ; median. Female 5.00-5.67 mm. Male genitalia: Pygofer with dorsolateral Width of head (across eyes) : Male 0.57- angles not produced. In dorsal view , anal 0.73 mm ; Female 0.77-0.87 mm. · segment strongly expanding to apex, antero­ Width of mesonotum: Male 1.40- 1.43 lateral angle narrowly produced, in lateral mm; Female 1.37 - 1.53 mm. view anal segment long and slender, wide across base of end of anal style, shorter than Holotype: Male, Fuhsing, Taoyuan Hsien greatest length about 1 : 4.1 , apex narrowly 28-IV-1985. (deposited in NCHU) protruding, ventral margin evenly produced Paratypes: 1 9 Wuchieh, Nantou Hsien downward. Phallus with both sides sclerotized , 26-VII-1984; 13 oo 12 99 Fuhsing, in lateral view reaching about as far as anal Taoyuan Hsien 28-IV-1985; 1 o segment, in dorsal view with two auricle­ 4 99 Fuhsing, Taoyuan Hsien 12-V- like processes at basal sixth, apex reflected 1985; 7 oo 8 99 North Tungyoanshan, cephalad ventrally. Apex of phallus at same Nantou Hsien 20-VII-1985 S. J. level of ventral margin of phallobase. In Fang; 6 oo 8 99 North Tungyoan­ dorsal view, process of phallobase acuminate shan, Nantou Hsien 23 - VII-1985 curved downward. Genital styles nearly same C. T. Yang. (deposited 3 oo 3 99 in width, slightly bent angularly at middle , ob­ BMNH) tusely rounded at apex, anterior margin of 104 Meenoplidae of Taiwan (Hornoptera : Fulgoroidea)

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0 .4mm

0.3mm 0.4mm

Fig. 12. Eponisia guttula Matsumura. A. Frans and clypeus, ventral view; B. Vertex, pronotum and mesonotum C. Tegrnen; D. Wing; E. Male genitalia, lateral view; F. Aedeagus, lateral view; G. The same, dorsal view H. Male genitalia, caudoventral view; I. Female genitalia, lateral view; J. Male anal segment, dorsal view K. Female genitalia, ventrocaudal view. basal half produced dorsad in a convex lobe produced mesad. In caudal view, genital styles which gives off distally a stout process, pos­ divergent, converging only apically, inner terior margin at basal third with a scaled area, margin of styles strongly produced mesad anterior margin near base of process, strongly medially. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 105

Female genitalia: Anal segment quadrate. This species is similar to Nisia species but In profile, ventral valvifer distinctly produced differs from latter as follows: caudoventrally, nearly reaching apex of beak, General color yellowish, abdomen pale forming a deep gap between ventral valvifer and yellowish. Laterofrontes each with 13 sensory valvula. Ventral valvula with apex produced pits. Each side of tergal pit configuration of into unequilateral triangle beak dorsocaudal tho_racic segments I 0-4-3. Inner side of side the longest, slightly arched, rounded at trochanter each with 8 keels. Spinal formula apical angle. of hind leg 5-4-1 . Tergites I-III without sensory pit. Tergites IV-VII each with 2 Length of body (includ. teg.): Male 4.54- sensory pits. Tergite VIII with 1. 4.77 mm; Female 4.30-4.83 mm. Length of tegment: Male 3.50-4.07 mm; Material examined: Female 3.83-4.23 mm . Width of head (across eyes): Male 0.57- Lectotype: Male (here designated). 0.6 1 mm; Female 0.57-0.62 mm. Taiwan, labelled; Formosa, Matsumura Width of mesonotum: Male 1.10-1.1 7 (printed), 30.iv.07, Horisha (in Japan­ mm; Female 1.10-1.17 mm. ese, handwritten on reverse) , Eponisia guttula det. Matsumura. (examined by First instar nymph (Fig. 13) Wilson). Body length 0.50-0.60 mm, width 0.32- Paralectotypes: Taiwan, 2 dd 3 99 16 .viii. 0.37 mm. 1906, Taikokan (1 d mouted separated-

0,3mm

Fig. 13. First instar nymph of Eponisia guttula Matsumura. A. Dorsal view; B. Thorax, flat surface. 106 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

ly); 1 d 12.vii.1906, Ako; 1 d 7.viii. Yang. l 9 Takeng, Taichung Hsien 1906, Hoppo. 28- IV - 1986 M. L. Chan. Notes: Four localities (Taikokan, Hoppo, Nymphal specimens examined: Horisha, Ako) were mentioned in the original description but the number of specimens was 1st instar nymph: 7 Takeng, Taichung 5- not stated. Specimens from all these localities VI-1986. were present in the syntypes appear to be Host plant: unknown. specimen figured by Matsumura. The wings Distribution: Taiwan. were possibly removed for drawing (figured by Matsumura) and then reattached. Eponisia macula* n. sp. Additional Specimens examined: 4 do 2 (Fig. 14) 99 Takeng, Taichung 21-Vl-1985; 1 d 1 9 Hsenping, Kaohsiung Hsien General color brownish black. Body not 12-VII-1984; 2 do 4 99 Wuchieh, covered with powdery wax. Vertex, frons Nantou Hsien 26-VII-1984; 1 9 yellowish brown. Mesonotum with three Wuchieh, Nantou Hsien 27-VII-1984; yellowish longitudinal stripes from anterior to 2 do 6 99 Kuanzuling, Tainan Hsien posterior. Ocelli white. Legs yellowish brown. 31-Vll- 1984; 1 d 2 99 Puli, Nantou Tegmina pale black with distinct white veins. Hsien 14-IX- 1984; 1 d Puli, Nantou Abdomen dark brown with lateral sides brown. Hsien 10-Vlll-1984; 1 9 Takeng, Frons longer in middle line than wide at Taichung 19-IV-1985 M. M. Yang; 7 widest portion about 1.6 : 1. Lateral carinae do 7 99 21-IV-1985; Many specimens strongly extended and elevated, with 31-33 Fushing, Taoyuan Hsien 28-IV-1985; sensory pits along each inner side (including Many specimens Shiayunping, 28-IV- several ones on vertex). Postclypeus slightly 1985; 7 do 7 99 Takeng, 2-V-1985; wider at base than long in middle line . Ros­ Many specimens Fushing, ll-V-1985; trum with relative length of each segment about 3 do 2 99 Jihyuehtan, Nantou Hsien 1.2 : 1.3 : 1. Antenna with basal segment 25-V-1985; 2 do Takeng, 5-VI- wider than long, second segment long than first 1985; 1 d Kuantzuling, Tainan Hsien about 2.4 : 1. Tegmina about 2.3 times as long 18-VI-1985; 2 do 2 99 Peipu, Hsin­ as wide. Sc+ R with about 22 sensory pits, 2 chu Hsien 22-Vl-1985; 5 do 18 99, above, another below. First claval vein with Wufeng, Hsinchu Hsien 24-VI-1985; 18-19 sensory pits along each side. R of wing 1 9 Tsaoling, Yunlin Hsien 6-VII-1985 branched. Shape of wax-secretion pores as C. T. Yang; 2 do 1 9 Puli, Nantou Hsien figured, inner plate of tergite VII with lateral 18-VIl-1985; 1 9 Liyan, Taichung half distinctly narrower than median. Hsien 19-VII-1985; 1 d 2 99 Baolai , Male genitalia: Pygofer with dorsolateral Kaohsiung Hsien 21-VII-1985; 1 d angles slightly produced. In dorsal view, anal Kaochung, Kaohsiung Hsien 20 do 22 segment elongate, strongly expending to apex, 99 Chiasien, Kaohsiung Hsien 23-VII- anterolateral angle widely produced, in lateral 1985; 2 do Kuangyinshan, Taipei Hsien view anal segment short and stout, wide across 6-IX-1985; 3 do 6 99, Wulai, Taipei base of end of anal style shorter than greatest Hsien 9-IX-1985; 3 99 Chiasien, length about 1 : 3 .0, apex obtusely protruding Kaohsiung Hsien 11-IX-1985; 9 do ventral margin downward medially. Phallus 6 99 Nanhua, Tainan Hsien 4-IV-1986 straight , membraneous, central portion with C. T. Yang; 7 do 23 99, Chiasien membraneous production anteriorly. In dorsal Kaohsiung Hsien 5-IV-1986 C. T. view with two auricle-like process. Anterior

* L. macula- spot, indicated the black spots of tegmen. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 107

0 .5mm

1mm ____.

. · ~. F ( \. ~~' :.: \,_ ~~.: : :::, 0.2mm

_ _ .2,_4_m_ m-'----

0.2mm Q3mm ~-·--- · · ----' ____0.3"1_'!'

Fig. 14 . Eponisia macu/a n. sp. A. Frans and clypeus, ventral view; B. Vertex, pronotum and mesonotum; C. Tegmen; D. Wing; E. Male genitalia, lateral view; F . Aedeagus, lateral view; G. The same, dorsal view; H. Male genitalia, lateral view; I. The same, caudoventral view; J. Female genitalia, lateral view; K. The same, ventrocaudal view; L. Male anal segment, dorsal view. apodeme of aedeagus rounded, at same plane elongate, curved at apical half, obtusely round­ with phallus and phallobase. Processes of ed at apex, anterior margin of basal half pro­ phallobase decline foliaceous. Genital style duced dorsad in a convex lobe which gives 108 Meenop!idae of Taiwan (Homoptera : Fulgoroidea)

off distally a stout process, posterior margin at Female genitalia usually with ventral valvula basal third with a scale area, anterior margin quadrate, most portion covered by production near base of process, strongly produced mesad. of ventral valvifer with a long process slightly In caudal view, genital styles divergent, inner curved mesad. margin of styles strongly produced mesad medially. Key to genera of Meenoplinae in Taiwan Female genitalia: Anal segment quadrate 1. Frans with a distinct median carina (except with anal style large, both curved downward. A. formosanus); anal segment large, curved Ventral valvifer distinctly produced caudo­ downward at apical half; ventral valvula ventrally, covered by ventral valvula partially. of female quadrate, large, most portion Ventral valvula with apex produced into equi­ covered by production ventrally , with lateral triangle beak, pointed at apical angle. a long process at ventrocaudal angle . . ... In caudoventral view , process of ventral valvula ...... Anigrus Stal directed downward, with a gap between ventral Frans without distinct median carina; anal valvifer and ventral valvula. Ventral valvifer segment stout, with finger-like lateral divergent widely, distance between ventral process basally, directed downward; valvulae wider than width of anal segment. ventral valvula of female reduced into a stout process, apex with a blade-like Length of body (include. teg.): Male 4.33 process, not covered by ventral valvifer mm; Female 3.77-4.27 mm...... Metanigrus n. gen . Length of tegmen: Male 3.70 mm; Female 3.40-3.63 mm. Genus Metanigrus* n. gen. Width of head (across eyes): Male 0.63 mm; Female 0.53-0.65 mm . Body small to medium, covered with a few Width of mesonotum: Male 1.13 mm ; powdery wax. Vertex without median carina, Female 1.10-1.20 mm. with two posterolateral areolets at base. Frans without median carina, an interruption at Holotype: Male, Kuanzuling, Tainan Hsien frontoclypeal suture between lateral carinae of 18-VI-1985. (deposited in NCHU). frons and clypeus. Clypeus with median carina. Paratypes: 5 99 Kuantzuling, Tainan Hsien Lateral carinae of postclypeus present. Pro­ 28-IV-1985 C. T. Yang; 3 99 Kuant­ and mesonotum with median carina. Tegmen zuling, Tainan Hsien 18-VI-1985. slender, near same width. Sc+ R with several (deposited 3 99 in BMNH). sensory pits. Spinal formula of hind leg 8-6- Host plant: unknown. 5. Male genitalia in lateral view, anal segment Distribution: Taiwan. stout with a finger-like lateral process ventro­ basally directed downward. In lateral view, Subfamily Meenoplinae Fieber phallus short, acuminate, a membraneous lobe (Fig. 5, A-B) (phyllobasal process) with many scale-like productions cover it. Genital style inner margin Meenoplinae Fieber, 1872, Katalog europais­ set with long setae, apex of genital style re­ chen Cicadinen: I-IV p. 3. duced into a pointed process, curved cephalad Tegmen with Sc+ R and M forked near apically. Female genitalia in lateral view, apex. Sc+ R with or without sensory pits. ventral valvifer with basocaudal angle turned Claval veins fused near middle, first claval vein mesad and produced into a thumb-like process. with one row of sensory pits, second strongly Ventral valvula not covered by ventral valvifer, curved with many sensory pits arranged ir­ reduced into a stout process, apex with blade­ regularly. M 1+ 2 single. R of wing simple. like process.

* Gr. prefix meta-near+ Anigrus. Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 109

Phallus short, acuminate in lateral view, nearly Type species: Metanigrus yami n. sp. (here same length as anal segment. Process of phal­ designation) lobase swelling on both sides, surface beset with Diagnosis: This genus has the characters, scale-shaped processes obscure the phallus. first claval vein with a row of sensory pits, Genital style curved apically, tapering into a second strongly curved with many sensory pits process, outer side with thumb-like process arranged irregularly; common petiole of claval near apex, directed upward, inner margin with a veins fused near middle; Sc+ R and M forked convex which set with long setae . In caudal near middle are Meenoplinae's. But the ventral view, genital styles divergent, narrowed medial­ valvula of female genitalia connecting with ly. Apical half with outer margin rounded, caudoventral angle of ventral valvifer are similar inner margin ridged, forming a obtusely round­ to those in Kermesiinae. The process of phal­ ed triangle. Apex strongly curved outward, lobase this genus resembles Anigrus but differs forming a hook-like process. in having ventral valvula not covered by ventral Female genitalia: In lateral view, anal valvifer; vertex and frons without median segment stout. Anal style elongate. Ventral carina absolutely; anal segment of male with valvifer with basocaudal angle turned mesad basoventral process. and produced into a thumb-shaped process. Ventral valvula reduced into a stout process, Metanigrus yami* n. sp. apex with blade-like process. In caudodorsal (Fig. 15) view, ventral valvifer approximate each other, separated with ventral valvula, forming a General color dirty yellow. Mesonotum deep gap. dark brown. Ocellus light yellow. Legs yel­ lowish brown. Tegmen hyaline. Abdomen Length of body (includ. teg.): Male 3.00- brown. 3.53 mm; Female 3.77-3.97 mm. Frons longer in middle line than wide at Length of tegmen: Male 2.67-3.07 mm ; widest portion about 1.5 : 1. Lateral carinae Female 3.20-3.40 mm. concave, reaching over median level of antecly­ Width of head (across eyes): Male 1.00- peus, with 22-24 sensory pits along each inner 1.07 mm; Female 1.13-1.20 mm. side (including several ones on vertex). Postcly­ Width of mesonotum: Male 0.53-0.60 peus as long in middle line as wide at base. mm; Female 0.62-0.65 mm. Rostrum with relative length of each segment about 1.1 : 1.4 : 1. Antenna with basal seg­ Holotype: Male, Lanshu, Taitung Hsien ment wider than long, second segment longer 29-IX-1985. (deposited in NCHU). than first about 2.4 : 1. Mesonotum with Paratypes: 1 o Lanshu, Taitung Hsien 4- distinct median carina. Tegmen about 2.5 VII-1985; 2 oo 5 99 Lanshu, Taitung times as long as wide. Sc+ R with 4-8 sensory Hsien 5-VII-1985; 1 o Lanshu, pits. im, r-m and anterior margin at two-thirds Taitung Hsien 8-VII-1985; 3 99 Lan­ with white areas. First claval vein with 9-10 shu, Taitung Hsien 12-Vll-1985 L. C. larger sensory pits at basal half and followed Tang; 1 o 3 99 Lanshu, Taitung Hsien with another about 20 small ones pits. 2A 3 of 12-VII-1985 J. T. Yang; 1 9 Lanshu, wing not reaching margin . Taitung Hsien 12-VII-1985 C. T. Male genitalia: Pygofer with dorsolateral Yang; 1 9 Lanshu, Taitung Hsien 13- angles not produced. In Lateral view, anal VII-1985 L. C. Tang; 3 oo 6 99 segment trapezoid, basolateral side with a Lanshu, Taitung Hsien 29-IX-1985; finger-like process, directed downward. Anal 1 o 4 99 Lanshu, Taitung Hsien 30- style slender, parallel with margin of segment. IX-1985; 1 o Lanshu, Taitung Hsien

* Name of a tribe, indicated collecting from Lanshu. 110 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

Imm

0.Jmm F

H

0.2mm

0.lmm

Fig. 15 . Metanigrus yami n. sp. A. Frons and clypeus, ventral view; B. Vertex, pronotum and mesonotum ; C. Tegmen; D. Wing; E. Male genitalia, lateral view; F. Aedeagus, lateral view; G. The same, ventrocaudal view ; K. Male anal segment, dorsal view .

l-X-1985 . (deposited 2 dd 3 99 in Host plant: unknown. BMNH). Distribution : Taiwan (Lanshu). Shun Chern Tsaur, Chung Tu Yang and Mi chael R. Wilson 111

Genus Anigrus Stal segment concaved medially (Fig. 17 E) ...... frequens (Matsumura) Anigrus Stal, 1866, Hemiptera Africana 4 :172. Body small to median , covered with several Anigrus formosanus (Matsumura, 1914) powdery wax. Vertex with distinct median (Fig. 16) carina down frons, with two posterolateral Paranisia formosana Matsumura, 1914 Ann. areolets at base. Median carina almost reaching Mag. Nat. Hungarici 12 :284. median ocellus. An interruption at frontocly­ Anigrus formosanus : Muir, 1930 Ann Mag . peal suture between lateral carina of frons and Nat. Hist. (10)6 :471. clypeus. Clypeus with median carina. Pro­ and Mesonotum with median carina. Tegmina General color brown to whitish gray. slender, nearly same width. Sc+ R without Bo_dy more or less covered with powdery wax. sensory pit. Claval vein uniting near apex of Vertex, frons , antenna and mesonotum dark clavus , first claval vein with sensory pits on it. brown. A yellowish stripe from vertex to Wing with R not branched, 2A forked near frontoclypeal suture. Eye brownish black. apex. Spinal formula of hind leg 8-6-5. Male Ocellus white. Leg yellowish. Abdomen black, genitalia in lateral view , with anal segment with lateral side orange red. slender, turned downward at apical half, in Head with lateral carinae concave below dorsal view deeply incised at apical margin eye, nearly parallel before areolets, with about forming a sharply pointed angle at each side. 26 sensory pits along each inner side (including In lateral view , phallus straight. Genital style several ones on vertex). Median carina of head nearly same width, strongly angularly bent at replaced by a yellowish vase-shaped marking. middle, at apex obtusely rounded with many Frons longer in middle line than wide at widest setae, inner margin concave beneath rounded portion about 1.4 : 1. Clypeus with median portion at apical fourth, a finger-shaped pro­ carina. Postclypeus as Jong in middle line as duction at apical third, margin with long setae. wide at base. Rostrum with relative length of In caudal view, divergent, inner margin with each segment about 1.1 : 1.2 : 1. Antenna with small production near middle, concave at apical basal segment wider than long, second segment third. Fem ale genitalia in lateral view , anal longer than first about 2 : l. Mesonotum with segment stout, base with a production pos­ two obscure yellowish stripes longitudinally. teriorly. Ventral valvifer with basocaudal angle Spinal formula of hind leg irregular, Post-tibia turned mesad and produced into a finger-like each with 3+5 teeth apically, basal metatarsial process. Ventral valvula quadrate, in caudo­ segment with 5 (male) or 6 (female) teeth, dorsal view , ventral valvula quadrate, with second segment with 6 in right, 5 in left. two productions, the upper one thumb-like, Tegmen about 2.8 times as long as wide. directed cephalad, the lower one acuminate, Sc+ R with 3-5 sensory pits. First claval vein directed upward. Ventral valvifer at base with 9-12 sensory pits, second with 27-32 produced into a elongate process, approximate sensory pits. Shape of wax-secretion pores each other, with la terodorsal angle angula te. uniform hexagonum but different in size , on tergites VI and VII smaller, tergite VIII central Type species: Anigrus sordidus Stal, 1866 portion larger, beehive-like all arranged tightly. (original designation) Male genitalia: Pygofer with dorsolateral angles not produced. In lateral view , anal Key to the species of Anigrus (males only) segment very large , directed vertically down­ ward at apical half, caudal upper margin curved 1. Median carina absent; in caudal view , anal with central portion strongly elevated, forming segment deeply incised medially (Fig. 16 a declivous plane. In caudal view , anal segment H) ...... formosanus (Matsumura) at upper portion with central part elevated, Median carina present; in caudal view , anal both sides deeply concave , lower portion 112 Meenoplidae of Taiwan (Homoptera: Fulgoroidea)

0.5mm

Fig. 16. Anigrus formosanus (Matsumura) . A. frons and clypeus, ventral view; B. Vertex, pronotum and mesono­ tum; C. Tegmen; D. Wing; E. Male genitalia, lateral view; F. Aedeagus, lateral view; G. The same, dorsal view; H. Male genitalia, ventrocaudal view; I. Female genitalia, caudal view; J. The same, caudoventral view. slender, two lobes. Anal style connect with apex, in dorsal view a membraneous area anal segment tightly, directed upward at apex. located between processes of phallobase and Process of phallobase shaped like muffler, with phallus. In lateral view, genital style stout, many reverse teeth. Phallus stout, narrowed to strongly bent angularly at middle, with many Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 113

setae along outer margin, inner margin concave a male from Hokuto is selected as Jectotype. at apical third. A production near middle, with Comparison with nudifrons Fennah, long setae. In caudal view, divergent, inner Anigrus formosanus closely resembles A. margin of styles strongly produced mesad at nudifrons Fennah (1978). The male genitalia basal third, with 4-6 thick setae at middle. serve to separate the two species, particularly Female genitalia: In lateral view, anal the form of the anal tube appendages. In segment stout, anal style elongate. Inner plate formosanus they are considerably longer of tergites VII-VIII with many Jong hairs, (Fig. 16 H) than in nudifrons where they are shape of tergite VIII larger beehive-like. Ven­ short and more or Jess triangular at the tip . tral valvifer at base produced into a elongate process, approximate each other. Ventral Other specimens examined: 2 do 1 9 valvula quadrate, caudal margin sinuate, ventro­ Takeng, Taichung 19-IV-1985 M. M. caudal process slightly produced ventrobased. Yang; 12 do 119921-IV-1985; 2 do In caudodorsal view, ventral valvula curved 2-V-1985; 1 d Songh~, Taichung toward central portion, inner basal angles Hsien 8-V-1985; 2 do takeng, Tai­ roundly produced. Distance between ventral chung 5-IV-1985; 1 9 Kuantzuling, valvulae almost same width of anal style at apex. Tainan Hsien 18-VI-1985; 1 9 C. T. Yang; 1 9 19-VII-1985 M. M. Yang; Length of body (includ. teg.): Male 4.67- 1 9 Baolai, Kaohsiung Hsien 21-VIl- 5.20 mm ; Female 5.20-5.57 mm. 1985; 2 do 4 99 Kaochung, 22-VII- Length of tegmen: Male 4.00-4.53 mm; 1985; 4 do 2 99 Chiasien, 23-VII- Female 4.53-4.90 mm . 1985; 1 d Takeng, Taichung Hsien Width of head (across eyes): Male 0.67- 12-IX-1985; 1o17-X-1985 M. M. 0.70 mm ; Female 0.67-0.70 !Tlm. Yang ; 1 d, Shengsing, Miaoli Hsien Width of mesonotum: Male: 1.13-1.27 13-X-1985 M. L. Chan. 1 o 4 99 mm; Female 1.17-1.30 mm. Nanhua, Taiwan Hsien 4-IV-1986 C. T. Yang. 1 d Takeng, Taichung Iisien Material examined: 28-IV-1986 M. L. Chan. Host plant: unknown. Lectotype: Male (here designated). Distribution: Taiwan. Taiwan, labelled, Formosa, Matsumura (printed), Hokuto, 30.viii.1906 (hand­ Anigrus frequens Matsumura, 1914 written, on reverse). (dissected, ex­ (Fig. 17) amined by Wilson). . Paralectotypes: Taiwan, 4 do 3 99 same Paranisia frequens Matsumura, 1914 Ann. Mus. data as Lectotype but collected 31.viii. Nat. Hungarici 12:285. 1906; 1 d 1 9 Tansui, 29.vii.1906; 1 Paranisia nigricans Matsumura, 1914 Ann. Mus . d Shurui, 3 l.vii.1906, 1 9 Ako, 12.vii. Nat. Hungarici 12:285. syn. nov. 1906; 1 d 2 99 Hoppo, 6.vii.1907. Anigrus frequens: Muir, 1930, Ann. Mag. Nat. Hist. ( 10)6 :4 71. Notes: Matsumura (1914) in his original Anigrus metalces Fennah, 1978 Ann. Zoo!. description did not give any specific localities Warsz. 34(9):237. syn. nov. in Taiwan. A total of 15 specimens were examined from the syntype series. A female General color brown. Vertex and frons from Hoppo is labelled by Matsumura as yellowish brown, mesonotum dark brown. Paranisia formosana and may be have been the Ocellus deep yellow. Leg yellowish brown. specimen figured. However, it agrees well Tegmen semihyaline. Abdomen black, with with the females collected from Hokuto from lateral side orange red. where males were also collected. The single Frons longer in middle line than wide at male from Hoppo is lacking wings. Accordingly widest portion about 2.3 : I. Lateral carinae 114 Meenoplidae of Taiwan (Homoptera : Fulgoroidea)

Imm

___1~

lmm

oJmm

O.Jmm H ~---0 , 4mm

G

0 .3mm

Fig. 17. Anigrus frequens (Matsumura). A. Frans and clypeus, ventral view; B. Vertex, pronotum and mesono­ tum, dorsal view; C. Tegmen; D. Wing; E. Male genitalia, lateral view ; F. Aedeagus, lateral view; G. The same, dorsal view ; H. Male genitalia, ventrocaudal view; I. Female genitalia, lateral view; J. The same, ventrocaudal view; K. Male anal segment, dorsal view. concave , with about 26 sensory pits along each base. Rostrum with relative length of each inner side (including several ones on vertex). segment about 1.2 : 1.2 : 1. Antenna with Postclypeus longer in middle line than wide at basal segment wider than long, second segment Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 115

longer than first about 2.3 : 1. Mesonotum Lectotype: Male (here designated). with median carina at basal two-thirds. Teg­ Taiwan, labelled Formosa, Matsumura men about 2.8 times as long as wide. Sc+ R (printed), Shoka, 21.vii.1906. Hand­ with 8 sensory pits. First claval vein with about written label Paranisia frequens . (dis­ 10 sensory pits, second with about 37 sensory sected, examined by Wilson).

pits. 2A 3 of wing not reaching apex. Shape of Paralectotypes: Taiwan, 1 d 1 9 Shoka; wax-secretion pores uniform hexagonum but 1 9 21.vii.1906, Shako; 1 d 19,20.vii. different in size, on tergite VI the smallest, 1~06, Shoka; 3 do 5 99 Kaoshun; 1 9 tergite VII large, tergite VIII the largest, all Taikokan, 16.viii.1906; 1 d 3 99 arranged tightly. Hokuto, 3 l .vii.1906; 1 d 2 99 Tanshui, Male genitalia: Pygofer with dorsolateral 29 .vii.1906. angles produced. In lateral view, anal segment Lectotype: Male of A. nigricans (here slender, turned downward at apical half, anal designated). Taiwan, labelled, Formosa, style large, distinctly rounded at apex. In Matsumura (printed), Hoppo, 8.viii. dorsal aspect, deeply incised at apical margin, 1906 (handwritten on reverse), labelled forming a sharply pointed angle at each side. Paranisia nigricans. (dissected, ex­ Phallus straight, with central portion sclero­ amined by Wilson). tized. In lateral view, reaching slightly shorter Paralectotypes: 2 do 4 99 same data as than genital styles. Process of phallobase lectotype. bilobed, foliaceous, membranceous, reaching Holotype of Anigrus metalces Fennah about apical third of phallus. Genital styles_ 1978, Type locality; Vietnam. Type nearly same width, strongly angulately bent depository; Insitute Zoo!. Pan, Poland. at middle, obtusely rounded and covered with (examined by Wilson). many setae at apex, inner margin concave beneath rounded portion at apical third, a Notes: Six localities were mentioned by thumb-shaped production at curved inner Matsumura in his original description (Tansui, portion, margin with long setae. In caudal Hokuto, Shoka, Taikokan, Hoppo, Koshun) view, divergent, inner margin with production but the number of specimens examined was near middle, slightly con ~ ave at apical fourth. not stated. The synonomy of nigricans with Female genitalia: In lateral view, anal frequens was made by comparing the male segment stout. Anal style elongate. Ventral genitalia of each species directly. Although valvifer with basocaudal angle turned mesad described by Matsumura in the same paper the and produced into a finger-shaped process. In name frequens has chosen for the two species. caudodorsal view, anal style evenly convex The synonomy of A. metalces with fre­ at apex, slightly produced medially, deeply quens was made on the basis of comparisons concave at basal margin. Distance between two of the male genitalia of the holotype with upper productions about 2 times as wide as frequens. width of anal style at apex. Distance between This species has some kinds of variations processes of ventral valvifer slightly narrower distributed in three main populations which than width of anal style at apex. collected from Songho, Yushih, and other lacalities. The altitudes which they inhabit is Length of body (includ. teg.) : Male 3.46- 500 m, 1,400 m, 300 m respectively. Their 5.10 mm ; Female 3.88-4.12 mm. body coloration also has various forms: white Length of tegmen : Male 3.00-4.67 mm; in Sonr ho population (S. P.); brown in the Female 3.47-4.93 mm. other ' vo. Body size appears the smallest in Width of head (across eyes): Male 0.57- S. P. ,ranged from 3.47-4.07 mm in male; 0.70 mm; Female 0.60-0.73 mm. 3 .88- 4 .12 mm in female), Yushih population Width of mesonotum: Male 0.87-1.27 (Y . P.), the largest, (ranged from 4.80-5.10 mm; Female 1.03-1.30 mm. mm in male ; 5.00-5.53 mm in female), others 116 Meenoplidae of Taiwan (Homoptera: Fulgoroidea)

(0. P.) ranged from 3.93-4.63 mm in male ; type material to M. R. Wilson . Without those 4.57-5.33 mm in female. The ratio of second support this project would have been impos­ against first segment of antannae is 2.8 : 1 in sible. 0. P.; 3.0: 1 in S. P.; 3.3: 1 in Y. P .. Distance between process of female ventral valvifers and References ventral valvulae is widest in P .P . narrowest in S. P. Inspite of having so much minor dif­ Cheng, K. T. 1956. Observatior.s on the ferences in which mentioned above, their male morphology and hibernation of the rice genitalia (so far the most important character fulgorid, Nisia atrovenosa LPth. Ent. and we use to separate species) are the same. So we phytopath. 2:218-219. decide to treat them as the same species. Distant, L. 1906. The fauna of British India, including Ceylon and Burma. Rhychota. Other specimens examined: 7 cid 4 99 Heteroptera-Homoptera 3 :309. Songho, Taichung Hsien 2-VII-1984; Emeljanov, A. F. 1984. A contribution to 13 dd 7 99 Songho, Taichung Hsien knowledge of the families Kinnaridae and 5-VII-1984; 2 dd 2 99 Hsenping, Meenoplidae (Homopcera, Fulgoroidea). Kaohsiung Hsien 10-14-VII-l 984; Ent. Rev. 64(1):61-65. 11 dd 13 99 Yushih, Nan tou Hsi en Fennah, R. G. l 950a. A generic revision of 14-X-1984; 1 d Nanjenshan, Ping­ Achilidae. Bull. · 1us. (Nat. Hist.) Ent. tung Hsien 11-11-1985; 2 9 Nanjen­ 1(1):13. shan, Pingtung Hsien 12-11-1985 L. Y. Fennah, R. G. l 950b. Fulgoroidea of Fiji. Huang; 1 d 2 99 14-Il-1985; 3 dd Bull. B. P. Bishop Mus. 202:46-47. 4 99 Takeng, Taichung Hsien 21-IV- Fennah, R. G. 1956. Fulgoroidea from Sou­ 1985; 1 d 1 9 19-IV-1985 M. M. thern China . Proc. California. Acad. Sci ., Yang; 2 99 Shiayunping, Taoyuan ser. 4, 28(13):474-476. Hsien 28-IV-1985; 9 dd 6 99 Songho, Fennah, R. G. 1971 . Homoptera: Fulgoroidea Taichung Hsien 8-V-1985; 16 dd 6 Suppl. Insects of Micronesia. 6(8):584- 99, Shiayunping, Taoyuan Hsien l 2- 585. V-1985; 1 d Takeng, Taichung Hsien Fennah, R. G. 1978. Fulgoroidea (Homoptera) 5-VI-1985; 4 cid 7 99 Songho , Tai­ from Viet-nam. Ann. Zoo!. Warsz . 34(9): chung Hsien l-Vll-1985; 1 ci Takeng, 237-240. Taichung Hsien 30-IX-1985; 1 d Fieber, F. X. 1872. Katalog der europaischen 7-X-1985; 9 dd 4 99 Yushih, Nantou Cicadinen, nach Originalien mit Benutzung Hsien l -XI-1985. der neueaten Literatur:I-IV p. 3. Host: unknown. Kirdaldy, G. W. 1906. Leafhopper and their Distribution: Taiwan. natural enemies. Bull. Hawaii. Sugar Pl. Assoc. Div. Ent. 1(9):425-429. ACKNOWLEDGEMENTS Kirkaldy, G. W. 1907. Leafhopper supplement. (Homoptera). Bull. Hawaii. Sugar Pl. Fina ial support of the National Science Assoc. Div. Ent. 3:163. Council, Republic of China from December, Lindberg, H. 1958. Hemiptera in sularum 1985 to June 1986 is gratefully acknowledged. Caboverdensium . Comment. biol., Hel­ We wish to express our deepest appreciations to sinki 1( 19): 13 3-13 5 . Dr. Jan Van Stalle, Institut Royal des Sciences, Matsumura, S. 1914. Beitrag zur Kenntnis Naturelles de Belgique, Belgium, the late Dr. der Fulgoroiden Japans. Ann. Mus . Nat. T. E. Woodward, Queensland Museum, Austral­ Hung., 12(1):261-305. ia for their kindly help on sending references. Matsumura, S. 1938. Homopterous insects We also indebted to Dr. S. Takagi, Hokkaido collected by Mr. Tadao Kano at Kotosho, University, Sapporo, for loan of Matsumura Formosa. Ins. Matsumurana. 12(4):147- Shun Chern Tsaur, Chung Tu Yang and Michael R. Wilson 117

153 . Verlaten and Sebesi. Treubia 12(1):29- Melichar, L. 1903. Homopteren-Fauna von 35. Ceylon:52-53. Stal, C. 1866. Hemiptera Homoptera Latr. Metcalf, Z. P. 1945. General catalogue of the Hemiptera Africana 4 : 172. Hemiptera IV(6):219-238. Vilbaste, J. 1968. Contribution to the Au­ Muir , F. 1917. The Derbidae of the Philippine cherorrhyncha fauna of the Maritime. Islands. Philip. Jour. Sci. 12: 52. Territory:38-39. Muir, F. 1923. On the classification of the Wilson, M. R. 1981. Geographical varation Fulgoroidea (Homoptera). Proc. Hawaii. in Nisia nervosa (Motsh.) (Fulgoroidea: Ent. Soc. 5(2):205-247. Meenoplidae ). A preliminary notes. Acta. Muir, F. 1925. On the genera of Cixiidae, Ent. Fenn. 38:53. Meenoplidae and Kinnaridae (Fulgoroidea: Wilson, M. R. 1984. Revision of family Homoptera). Pan-Pacific Ent. 1 :97-100. Meenoplidae (Fulgoroidea): Problems and Muir , F. 1927. New species of African Me­ progress . Mitt. Schweiz. Ent. Ges . 57(4): enoplidae (Fulgoroidea:Homoptera). Ann. 449. Mag. Nat . Hist. 19(9): 197-208. Woodward, T. E. 1957. Studies on Queensland Muir, F . 1930. On a small collection of Hemiptera part II Meenoplidae. Pap. Univ. Fulgoroidea from the Islands of Krakatan, Qd. (Ent.). 1 :57-70.