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BULLETIN OF fr) THE BRITISH MUSEUM Sern wi (NATURAL HISTORY)

ENTOMOLOGY VOL. XA

1968

THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1969

DATES OF PUBLICATION OF THE PARTS

CON ADU W DN

23 February 1968 . 28 May 1968

- 28 May 1968

5 May 1968

a 3 September 1968 . 20 September 1968

4 October 1968

. 21 December 1968

PRINTED IN GREAT BRITAIN BY ADLARD AND SON LIMITED

DORKING, SURREY

CONTENTS

ENTOMOLOGY VOLUME XXII

. The evolution and dispersal of the grassland leafhopper genus Exitianus,

with keys to the old world species (Cicadellidae : Hemiptera). By H. H. Ross

. A revision of the genus Mucrogaster Latreille (Hymenoptera :

Braconidae). By G. E. J. NIxon

. An index-catalogue of African Phaneropterinae (Orthoptera : Tetti-

goniidae). By D. R. RAGGE

Hymenoptera from Turkey (Symphyta). By R. B. BENSoNn

The subgeneric divisions of the genus Bombus Latreille (Hymenoptera :

Apidae). By O. W. RicHARDS

. Alist of the type-specimens of Libellulidae and Corduliidae (Odonata)

in the British Museum (Natural History). By D. E. KiMmins

A revision of the genus Atolopus Fieber (Orthoptera : Acridoidea). By D. Ho tis

Revision of the genus Catasarcus Schénherr (Coleoptera : Curculion-

idae). By R. T. THompson

Index to Volume XXII

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109

209

277

307

397

455

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THE EVOLUTION AND DISPERSAI% OF THE GRASSLAND LEAFHOPPER GENUS EXITIANUS, WITH KEYS TO THE OLD WORLD SPECIES (CICADELLIDAE: HEMIPTERA)

H. H. ROSS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 22 No. 1

LONDON: 1968

THE EVOLUTION AND DISPERSAL OF THE GRASSLAND LEAFHOPPER GENUS EXITIANUS, WITH REYS [OTHE OLD WORLD SPECIES (CICADELLIDAE: HEMIPTERA)

BY

H. H. ROSS |}

a

Illinois Natural History Survey, Goeana

Pp. 1-30; 82 Text-figures

BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 22 No. 1 LONDON : 1968

THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.

In 1965 a separate supplementary serves of longer papers was instituted, numbered serially for each Department.

This paper is Vol. 22, No. 1 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.

World List abbreviation : Bull, Br. Mus. nat. Hist. (Ent.).

TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 23 February, 1968 Price Thirteen Shillings

Dae EVOLUTION AND DISPERSAL OF THE GRASSLAND LEAFHOPPER GENUS EXITIANUS, WITH KEYS) 1O THE-OLD WORLD SPECIES (CICADELLIDAE: HEMIPTERA)*

By H. H. ROSS CONTENTS

Page

INTRODUCTION . 3 EXITIANUS BALL ' : ; ; ‘ 4 KEy TO GROUPS AND OLD WORLD SPECIES—MALES 4 Nanus-Group . 7 Obscurinervis-Group 8 Distanti-Group 8 Taeniaticeps-Group . 9 Okahandia-Group 16 UNPLACED SPECIES : : ; 21 SPECIES TRANSFERRED TO OTHER GENERA 21 PROBLEMS OF SPECIES RECOGNITION 22 PHYLOGENY 24 PALAEODISPERSAL 28 ACKNOWLEDGMENTS 28 REFERENCES 29

SYNOPSIS

Keys and illustrations are presented for the identification of the five species-groups of Exitianus and for the identification of the species occurring in the Old World, the latter including seventeen species, nine new to science. Synonymy is included for the described Old World species plus a brief synopsis of their distribution. The phylogenetic relationships are adduced for the world fauna. When geographic distribution is compared with the postulated family tree there are strong indications that the genus evolved primarily in Africa with an ancient dispersal to the Americas and more recent dispersals to the Oriental and Australian regions.

INTRODUCTION

AMONG the most widespread and frequently the most abundant species of grassland leafhoppers are the moderately large tawny forms comprising the genus Exztianus, a member of the subfamily Deltocephalinae. This genus differs from related forms in having only a small number of apical, stout setae on the male pygofer; the female has a relatively long ovipositor extending conspicuously beyond the last dorsal segment. These characters are shared only by the presumably sister genus Nepho- tettix, readily differentiated from Exitianus by having the crown sharply ridged where it meets the face, and in being opaque green with various black markings.

* This work was supported in part by a research grant from the National Science Foundation, U.S.A. ENTOM. 22, I. I

4 H. H. ROSS

Based on similarities of male and female genitalia, Exitianus and Nephotettix are probably most closely related to Doratura and its allies; the entire complex can be segregated as a grass-feeding tribe, the Doraturini.

On every continent at least one common form of Exitianus has a conspicuous black band across the top of the head. So similarly coloured are these forms that considerable difference of opinion has existed concerning the number of species involved. The same difference has been expressed concerning the number of species occurring on the same continent. DeLong & Hershberger (1947) segre- gated eleven American species on the basis of colour and the number and position of the large pygofer spines. Linnavuori (1959) added a few more segregates from South America. No one appears to have undertaken a study of the Old World fauna of this genus. An opportunity in 1964 to examine types of the genus at the British Museum (Natural History) and Stockholm and to see large series of specimens from Africa, India, and Madagascar led me to attempt such a study. As new characters and species were found, the possibility became evident of reconstructing the phylogeny of the entire genus with a high degree of probability.

This paper treats the genus Exitianus in the following topical order: (1) a diagnosis and description of the world groups and the Eastern Hemisphere species, (2) a discussion of the peculiar problems encountered concerning species distinctions in the Eastern Hemisphere populations, (3) a derivation of the phylogeny of the genus, and (4) an application of these hypotheses to the question of its palaeodispersal.

This study is based entirely on the male sex. In several species, associated females are known but they have proven difficult to diagnose. Sexual dimorphism in colour is common, females of certain indubitably different species appear identical, and certain females that appear distinctive have as yet not been associated with males. In the face of these circumstances it appears that our present knowledge of Exitianus females does not lend itself to either keys or diagnoses of worthwhile predictive value.

EXITIANUS Ball

Exitianus Ball, 1929: 5. Type-species: Cicadula obscurinervis Stal. Mimodrylix Zachvatkin, 1935 : 108. Type-species: Athysanus capicola Stal.

Body usually about 4-5 mm. long. Colour chiefly tawny with various patterns of brown or black. Head only moderately produced (Text-figs. 1-14), the crown rounding into the face. Wings elongate and having typical leafhopper venation. Male pygofer having from two to six large dark or black spines along apical margin and only one or two other conspicuous spines. Aedeagus having an articulation between shaft and base, the shaft relatively simple in structure. Female ovipositor extending a considerable distance beyond abdominal tergites.

Distribution. Recorded from all continents except Antarctica.

KEY To GROUPS AND OLD WoRLD SPECIES—MALES

I Plate short and ovoid, without lateral setae (Text-figs. 54, 55) okahandia-group 18 = Plate elongate-triangular, with a row of long, stout lateral setae extending the length of the plate (Text-fig. 78) . ; : : . ‘ : ; 2

THE OED WORLD EXTCIANUS 5

2 (1) Pygofer having four to six apical brown or black spines (Text-fig. 15)

nanus-group 5 Pygofer having two or three apical brown or black spines (Text-figs. 19, 21) . 3

3 (2) Dorsal margin of aedeagus with only a slight, irregular concavity at apex (Text-fig. 20) 5 : . obscurinervis-group 6 -- Dorsal margin of aedeagus with 2 a large arcuate concavity at apex (Text-fig. 22) 4

4 (3) Spine 2 of pygofer as long and slender as spine 1 (Text-fig. 21)

distanti-group 4

- Spine 2 of pygofer much shorter than and thicker than spine 1 (Text-fig. 24) taeniaticeps-group 8

NANUS-GRovuP

5 (2) Known only from one widespread Old World species. ; : nanus (p. 7)

OBSCURINERVIS-GRoup

6 (3) Known from several New World species not treated further here. For diag- nostic treatments see DeLong & Hershberger (1947) and Linnavuori (1959).

DISTANTI-Grovup 7 (4) Known only from one African species : : : d distanti (p. 8)

Fics. 1-6. Head, pronotum and mesonotum of Evitianus spp. 1, E. nanus (India); 2, same, dark form (Okahandia, Rhodesia); 3, same, light form (Okahandia, Rhodesia) ; 4, E. okahandia (holotype); 5. E. twrneri (holotype); 6, E, natalensis (holotype).

6 H.H. ROSS

TAENIATICEPS-Grovup 8 (4) Socket of spine 2 of pygofer almost contiguous with that of spine 1 (Text-fig.

23); head with extensive dark markings (Text-fig. 13) : kilimanus (p. 16) _ Socket of spine 2 a considerable distance from that of spine 1 (Text-fig. 24); head variously marked . : : ‘ é 9

9 (8) Spine 2 of pygofer only slightly asymmetrical (Text- figs. 64, 6 5); aedeagal shaft tapering almost uniformly from base to gonopore (Text-fig. 33) attenuatus (p. 10) _ Spine 2 of pygofer markedly twisted or bent at tip (Text-figs. 24, 26); aedeagal shaft either parallel-sided, or enlarged near gonopore, or markedly con- stricted at base of dorsal points ‘ : ; ; ; ; ; d 10

Fics. 7-14. Head, pronotum and mesonotum of Evxitianus spp. 7, E. picatus (Mexico; 8, E. exitiosus (Illinois); 9, E. indicus (China); 10, E. indicus (holotype); 11, E. coronatus (holotype); 12, E. ootacamundus (holotype); 13, E. kilimanus (holotype); 14, E. greens- ladei (paratype).

to (9) Shaft of aedeagus as thick as in Text-figs. 25, 28, 30, 31, its paired dorsal points usually well separated at base as in Text-fig. 27, its apex long,

slender and markedly curved . ; ; . : : II — Shaft of aedeagus thinner (Text-figs. aata6, 39-51), or its dorsal points bias contiguous at base : ‘ 13

1r (10) Dorsum of head yellow with rigidly defined black spots as in ‘Text-fig. II coronatus (p. 14) — Dorsum of head otherwise (Text-figs. 10, 12) : : : 12 12 (11) Dark markings on top of head as extensive as in Text- fig. 12 ootacamundus (p. 14) — Dark markings on top of head chiefly a single, arcuate, dark line (Text-fig. 10), sometimes with additional light brown suffusions . ; indicus (p. 12) 13 (10) Dorsal profile of shaft with a slight notch on which dorsal points are situated

(Text-figs. 34-36, 39-42, 47) - + ee ee eK

16 (13)

17 (16)

tS (1)

19 (18)

THE OLD WORLD EXITIANUS

Dorsal profile of shaft with no indication of a notch where dorsal points are situated (Text-figs. 43-46, 48-51) F Profile of aedeagal shaft markedly angulate below gonopore, Poaopore opening

very long (Text-fig. 34) . pondus (p.

Either profile of aedeagal shaft rounded below gonopore (Text- -figs. 39-42) or gonopore opening much shorter (Text-figs. 35, 47) Either aedeagal shaft slightly to moderately flared before gonopore (Text- figs.

39-41) or dorsal points close to base of shaft (Text-fig. 42) taeniaticeps (p.

Aedeagal shaft not flared and dorsal points as far from base of shaft as shown

in Text-figs. 35, 36, 47 . ; ; frontalis (p. Head with extensive dark markings (Text- fig. 14) : ; as tisdaieas (p.

Head with only a dark arcuate band, as in Text-fig. ro Profile of aedeagal shaft with lateral ridge close to margin and apical excava- tion forming an obtuse angle with body of shaft (Text-figs. 48-51)

plebeius (p.

Profile of aedeagal shaft with lateral ridge closer to central line of shaft and apical excavation forming nearly a right angle with body of shaft (Text-

figs. 43-45) . : : ; : ; : 3 . mucronatus (p.

OKAHANDIA-GrRoupP

Apex of pygofer forming an upturned, fingerlike tip bearing one extremely stout, dorsally projecting spine and sometimes another much more slender spine (Text-figs. 60, 63); apex of style scarcely widened (Text-fig. 58) .

Apex of pygofer wider, not upturned, and bearing two fairly stout spines (Text-figs. 52, 56); apex of style greatly widened (Text-figs. 54, 55) ‘

Apex of pygofer bearing only one spine, this spine wider than pygofer tip

(Text-fig. 63); profile of aedeagus wider (Text-fig. 62) . zuluensis (p.

Apex of pygofer bearing a slender seta mesad of large one, the latter narrower than pygofer tip (Text-fig. 60); profile of aedeagus narrower (Text-fig. 59)

natalensis (p.

Style markedly shorter than plate (Text-fig. 55); costal area of front wing not

forming a conspicuous light stripe . : okahandia (p.

Style projecting beyond plate (Text-fig. 54): costal area a of front wing forming

a conspicuous light stripe ‘ ; é ; : é . turneri (p.

NANUS-GRovupP Exitianus nanus (Distant) comb. n.

(Text-figs. 1, 2, 3, 15, 16, 17, 18, 76)

Athysanus nanus Distant, 1908 : 345. India.

Athysanus insularis Distant, 1909 : 47. Amirante Is. syn. n. Athysanus simillimus Matsumura, 1914 : 185. Formosa. syn. n. Euscelis vulnerans Bergevin, 1925 : 42. Southern Algeria. syn. n.

This is the most widespread Old World species of the genus, occurring throughout Africa, Asia Minor, southern Asia, Formosa, islands in the Indian Ocean, and

Australia. Well-marked specimens show considerable geographic variation.

bars.

14)

19 20

18)

20) 17)

17)

The dorsal head markings may be faint or form dark spots or crescentic In speci-

mens from Asia, the dark markings usually form a solid curved arc interrupted on the meson (Text-fig. I, as in the type of manus); specimens from the southern half

8 H. H. ROSS

of Africa invariably have each half of the mark either narrowed in the middle (Text- fig. 2) or completely pinched off into a pair of black spots (Text-fig. 3). In specimens from Asia Minor and North Africa, the dark marks intergrade between the two darker types and occasionally these intergradations are encountered in Asian material also.

The type male of insularis (Text-fig. 15) has pygofer spines 2-4 relatively slender; males from Tanzania have them slightly shorter and more robust (Text-fig. 17) ; specimens from Johannesburg, Republic of South Africa, have a mixture of slender and stouter spines on each specimen, and sometimes five or six spines on a side (if so, usually not the same number on both sides). The spination of the pygofer therefore mirrors the variation found in the colour pattern.

I have studied the types of manus Distant and imsularis Distant but not those of simillimus Matsumura and vulnerans Bergevin. The latter two species I am placing on the basis of illustrations by Ishihara (1954, fig. 5) and illustrations in the original description, respectively.

The female seventh sternite is characterized by the bi-emarginate apical margin and the slightly bowed mesal area (Text-fig. 76).

OBSCURINERVIS-GRovupP (Text-figs. 7, 8, 19, 20)

The members of this American group of species are remarkably homogeneous in shape of aedeagus and pygofer, and in spines 1 and 2 of the pygofer. In all members spine 3 is absent, in most members spine 4 is present, and in occasional individuals of the latter species an additional spine 5 may be present on one or both sides of the capsule. The females exhibit a considerable number of inter-specific differences in the shape of the apical margin of the seventh sternite.

DeLong & Hershberger (1947) and Linnavuori (1959) deal with eleven and seven American species, respectively. I am making no attempt to correlate the two accounts pending the availability of longer tropical series needed to establish the associations of the two sexes and possible synonymy.

DISTANTI-Grovup Exitianus distanti sp. n. (Text-figs. 21, 22)

3g. Length 4mm. Colour of dorsum pale greenish yellow, face the same but with arcuate dark lines, venter of thorax and abdomen almost black, legs with coxae black, the remaining segments tawny with dark streaks. Dorsum of head proportioned as in Text-fig. 9, without any trace of arcuate banding. General structure typical for genus. Pygofer having only setae 1 and 2, both of them straight and slender (Text-fig. 21). Aedeagus with shaft elongate, slender, its apex arcuately incised, as seen from lateral view, and the shaft bearing a pair of dorsal projections below middle (Text-fig. 22).

Holotype ¢ (in glycerine). [TANZANIA] Tanganyika: Ilonga, 28.vi.1962 (L. Robertson). In B.M.(N.H.).

THE OLD WORLD EXITIANUS 9

Paratype. [TANZANIA] Tanganyika: Ukirigiru, 1 3 (pinned, abdomen in gly- cerine), 27.iv.1g61, light trap. In B.M.(N.H.).

The two specimens have been preserved in alcohol, and the greenish cast may be due to the action of that preservative. This species forms a unique group all of its own, characterized by the possession of only spines I and 2 on the pygofer, and the long and slender spine 2 that is much like spine 1. In all other known species having only spines 1 and 2, spine 2 is much thicker and more densely pigmented than spine I.

Fics. 15-22. Male genital parts of Exitianus spp. 15, 16, E. nanus (holotype of insularis) ; 17, 18, E. nanus (Tanzania); 19, 20, E. exitiosus (Illinois); 21, 22, E. distanti (holotype) ; 15, 17, 19, 21, pygofer; 16, 18, 20, 22, aedeagus, lateral aspect.

TAENIATICEPS-Grovup

In this group the aedeagus has a well-defined preapical notch, a sharp, curved apex, and a pair of small, dorsal, sclerotized points; the pygofer usually possesses only spines 1 and 2, of which spine 1 is slender and elongate, and spine 2 is much shorter, black, and more or less twisted at the tip (Text-figs. 23, 24, 64, 65). In most species the vertex of the head has only a single transverse-arcuate black band (Text-fig. 9), often with suffusions of paler brown (Text- fig. 10); in a few the vertex is darker (Text-fig. 12) or has a different pattern of banding (Text- fig. 11).

In those species for which the two sexes are associated (indicus, taentaticeps, frontalis, and plebeius), the female eighth sternite has a narrow, V-shaped apical incision bordered with darker colouring (Text-fig. 69). In the case of mucronatus,

ENTOM. 22, I. 1§

10 Hots KOSS

pondus, and several other species, females of the same type were present from the same or nearby localities. It seems likely that females of all members of this group share this distinctive configuration of the eighth sternite. The unassociated female type of capicola (Stal) (Text-fig. 70) can therefore at present be assigned to this group but not definitely to any species. As explained below under indicus Distant, geographic location gives a clue as to the possible identity of fusconervosus (Mots.).

30 »

28

ieee

rt

Fics. 23-32. Male genital parts of Exitianus spp. 23, E. kilimanus (holotype); 24, 25, E. ootacamundus (holotype); 26, 27, 28, E. indicus (type series); 29, 30, E. indicus (Ceylon); 31, 32, E. coronatus (holotype). 23, 24, 26, 29, 32, pygofer; 25, 28, 30, 31, aedeagus, lateral aspect; 27, aedeagus, posteroventral aspect.

Exitianus attenuatus sp. n. (Text-figs. 33, 64, 65, 66)

6. Length 445 mm. Colour above and below tawny, with a thin arcuate black line across dorsum of head (as in Text-fig. 9) and frontal area of head and legs with small dark markings. General structure typical for genus. Pygofer with spines 1 and 2, or 1, 2 and 3; spine 1 is long and slender, spine 2 or spines 2 and 3 are shorter, thicker, darker and very slightly asymmetrical at apex (Text-figs. 64, 65). Aedeagus with shaft tapering almost uniformly from base to

THE OLD WORLD EXITIANUS II

gonopore, the portion beyond that slender and arcuately incised (Text-fig. 33); near the base of the shaft are a pair of dorsal points or projections that are finger-like, flared laterally, the two well separated at the base (Text-fig. 66).

Holotype ¢ (pinned, genitalia in glycerine). SourH AFRICA: Cape Province, Somerset East, xi.1930 (R. E. Turner). In B.M.(N.H.).

Paratype. Same data, I § (pinned, genitalia in glycerine). In B.M.(N.H.).

Both type specimens have the pygofer spines somewhat damaged, but the remain- ing evidence indicates several interesting items: (I) spine I is long and slender; (2) spine 2 is only slightly asymmetrical (Text-figs. 64, 65), not approaching the bent condition (Text-figs. 23, 24, 26) of related species; and (3) the genetic control for spine 3 has not been completely lost. The latter two points warrant a serious consideration of this species as the most primitive of its group.

Exitianus pondus sp. n. (Text-figs. 34, 67)

3. Length 5mm. Colour tawny except for a fine dark arcuate line across anterior portion of vertex and a few dark marks on the face and legs. General structure typical for genus. Pygofer with spines 1 and 2 only, spine 1 long and slender, spine 2 deeper, shorter, almost jet black, and markedly asymmetrical (Text-fig. 67), the extreme tip slightly but definitely hooked ventrally. Aedeagus (Text-fig. 34) with profile of shaft wide at base and tapering slightly to gonopore, the apex with an arcuate incision occupying the apical third of the shaft and forming a long narrow apex sharply hooked at tip; near the base of the shaft is a prominent pair of dorsal points.

Holotype 3 (pinned, abdomen in glycerine). S. ArricA: Pondoland, Port St. John, 1-9. vii.1923 (R. E. Turner). In B.M.(N.H.).

Paratypes. Same data, 2 J. In B.M.(N.H.).

Three specimens that resemble the above types very closely but lack dorsal points at the base of the shaft are tentatively considered as this species but not included in the types, as follows: S. Arrica: Zululand, Gingindhlovu, 2 ¢, 15.vi.1926 (R. EF. Turner); MADAGASCAR: Morondova, forest south of Befasy, 1 3, 1.1956 (R.P.).

Exitianus taeniaticeps (Kirchbaum) (Text-figs. 37, 39-42, 71)

Athysanus taeniaticeps Kirchbaum, 1868 : 87. Messina, Syracuse.

I have not seen the type of this species, but the description fits perfectly the species here considered as taeniaticeps, which is the only species of the group so far identified from the circum-Mediterranean countries, Specimens from Iraq, Lebanon, Cyprus, France, and Libya are remarkably uniform, in contrast with other popula- tions in the species, in having the dorsal points some distance from the base of the shaft (Text-figs. 39-41). Males from Rhodesia, considered as this species, usually

12 We. ROSS

have these points almost at the base of the shaft (Text-fig. 42). Large series of males from central Sudan are much more variable in this character, completely bridging the morphological gap between the Mediterranean and Rhodesian popu- lations.

This situation suggests that the Mediterranean and South African populations represent previously isolated segments of an ancestral population that became morphologically different, then were brought together by changing ecological con- ditions and have since formed an intermingled, freely-hybridizing population in at least the Sudan.

In a few Rhodesian males the apico-central corner of the pygofer is almost square (Text-fig. 37), in contrast to the sinuate condition normally found throughout other populations of the species (as indicated by the broken line in Text-fig. 37).

The illustrations of Ribaut (1952 : 138, figs. 245-249) undoubtedly refer to this species, although pygofer spine 2 is shown as straight rather than contorted at the tip. It should be noted that in Fig. 248 the shaft is folded into the socket so that the latter hides the real base of the shaft and the dorsal points of the shaft therefore erroneously appear to be basal.

Exitianus frontalis (Distant) comb. n.

(Text-figs. 35, 36, 47) Athysanus frontalis Distant, 1917 : 317. Seychelles.

An extremely close relative of taeniaticeps, this species as here recognized is diagnosed by the uniformly thick shaft (Text-figs. 35, 36) in contrast with the slightly to markedly clavate shaft of taeniaticeps (Text-figs. 39-42). In frontalis the dorsal points of the shaft are some distance from the shaft’s base, as sometimes occurs in Mediterranean specimens but not in South African specimens of taensaticeps, sug- gesting that in the latter species there has been a displacement of this character in the southern part of Africa where both taeniaticeps and frontalis occur in the same area.

To date this species has been found in the Seychelles, Madagascar (Diego-Suarez, Montagne des Francais), and South Africa (Pondoland). It is interesting that the three specimens from Pondoland exhibit almost exactly the same range of variation in the aedeagal shaft as found in the two specimens illustrated from Madagascar. The male from the type series (Text-fig. 47) is a slender extreme, matched almost perfectly by a specimen from Pondoland in the Turner collection, B.M.(N.H.). I have studied the female type, in which the eighth sternite is notched as in other species of the group.

Exitianus indicus (Distant) comb. n.

(Text-figs. 9, 10, 26-30, 69)

?Athysanus fusconervosus Motschulsky, 1863 : 97. Ceylon. Athysanus indicus Distant, 1908 : 344. India. Athysanus atkinsoni Distant, 1908 : 345. India. syn. n.

THE OLD WORLD £EXITIANUS

50-51

Fics. 33-51. Male genital parts of Exitianus spp. 33, E. attenuatus (holotype); 34, E.

pondus (holotype); 35, 36, E. frontalis (Madagascar); 37, E. taeniaticeps (Rhodesia) ; 38, E. frontalis (type series) ; 39, 40, 41, E. taeniaticeps (Tripoli) ; 42, the same (Rhodesia) ; 43, 44, 45, E. mucronatus (holotype and two paratypes, respectively); 46, E. greensladet (holotype); 47, E. frontalis (type lot); 48, E. plebeius (Queensland); 49, the same (para- type of plebeius); 50, the same (Samoan Is.); 51, the same (homeotype of norrist). 37, 38, pygofer; all others, aedeagus or shaft only, lateral aspect,

14 H. H. ROSS

This is a pale species having a faint or thin, dark, transverse, arcuate band on the vertex. It has been taken at several localities in India (in great abundance at Kharagpur), Ceylon, Formosa, and Japan.

I have studied the female types of both Distant names. These are in B.M.(N.H.), bearing identical data ‘Calcutta, Atkinson Col. 92.6.’ The common type series of the two species contains several males that demonstrate the characters of this sex (Text-figs. 26-28).

There is a possibility that fusconervosus Motschulsky, described from the summit of Mt. Patannos, Ceylon, may apply to this species, which is the only one of the genus thus far recorded for the island, but the unavailability of the type mitigates against the use of its carrier name at the present time. Ishihara’s illustrations of E. capicola (Stal), listed under Mimodrylix Zakhvatkin, undoubtedly apply to this species (Ishihara, 1954 : 6).

Exitianus coronatus (Distant) comb. n. (Text-figs. II, 3I, 32)

Phrynomorphus coronatus Distant, 1918 : 52. India.

This species is remarkably like indicus in morphological details of the male genitalia (Text-figs. 31, 32) but differs markedly in the dorsal spotted pattern of the head (Text-fig. rr).

The species is known only from the type, which I have studied. It is in B.M. (N.H.) and was collected at Kumaon, N.W. India.

Exitianus ootacamundus (Distant) comb. n. (Text-figs. 12, 24, 25) Phrynomorphus ootacamundus Distant, 1918 : 51. India.

As is true for the above-mentioned species, this one also is practically identical in details of male genitalia (Text-figs. 24, 25) with imdicus, differing primarily in having a much darker head (Text-fig. 12).

To date the only known record is the type, which I have studied. It is in B.M. (N.H.) and was collected at Ootacamund, South India.

Exitianus mucronatus sp. n.

(Text-figs. 43, 44, 45)

3. Length 4-5 mm. Colour pale tawny except for a black arcuate band across the vertex, much as in Text-fig. 9, and a few pale marks on face and legs. General structure typical for genus. Pygofer with spines 1 and 2 only, spine 1 very long slender, spine 2 much shorter, asymmetrical, and slightly hooked at apex as in Text-fig. 37. Aedeagus with shaft moderately long and thick (Text-figs. 43-45), with an apical arcuate incision resulting in a sharp hooked tip; the dorsal points are minute and their bases do not interrupt the even contour of the dorsal surface,

THE OLD WORLD EXITIANUS 15

Holotype 3 (pinned, abdomen in glycerine). MApAGAscaAR: Ft. Dauphin (R.P.). In collection of Madagascar Scientific Institute.

Paratypes. MADAGASCAR: Ft. Dauphin, Tananarive, Bas Mangoky, Nosivola, Perinet, Monronova forest south of Befasy, Montagne des Francais and Andova, 28 3. In collections of Madagascar Scientific Institute, Illinois Natural History Survey and B.M.(N.H.).

This series of specimens appears to represent a distinctive and perhaps an endemic Madagascar species differing from plebeius in having a stouter aedeagal shaft whose ventral lateral ridges curve dorsally and disappear toward the apex. In Madagascar, mucronatus appears to displace taeniaticeps almost completely, although the latter species appears to be abundant in Madagascar at Mohali, lac Iconi, and one specimen has been taken from Rte de Fianarantsoa, Km. 334. These specimens of taeniaticeps are like the more massive Rhodesian type rather than the narrower type found around the Mediterranean.

Exitianus plebeius (Kirkaldy) sp. rev., comb. n. (Text-figs. 48-51, 68)

Nephotettix plebeius Kirkaldy, 1906 : 331. Queensland. Eutettix norrist Evans, 1938 : 14. West Australia. syn. n. Euscelis transversus Metcalf, 1946: 122. Guam. syn. n.

This species has the same tawny colour and black arcuate stripe on the vertex as is found in zmdicus and several other species of the genus (as in Text-fig. 9). It is most closely related to frontalis, from which it differs in lacking a notch on the dorsal profile of the shaft where the dorsal spines arise (Text-figs. 48-51), and to E. mucronatus, from which it differs in the longer and more slender aedeagal shaft.

The holotype 2 and paratype ¢, which I have examined, are from Cairns, Queens- land, Australia, deposited in the Bernice P. Bishop Museum, Honolulu. Both appear to be the same species and I am herewith designating the ¢ paratype as the allotype. I have also examined the ¢ holotype of tvansversus, which agrees exactly with males from Australia.

Concerning norrist, I am following the synonymy of Evans (1966). To date the only records of this species I have personally examined are from Australia, Guam, and Samoa. The records of capicola cited by Linnavuori (1960) from Fiji are based on females, but probably refer to this species. Previously this species has been confused with other species in the Old World having the same colour, and the Australian records have been labelled as capicola. The distinct, long shaft and the inconspicuous dorsal aedeagal spines indicate, however, that this species is different from those occurring on other continents,

16 H.H. ROSS

Exitianus greensladei sp. n. (Text-figs. 14, 46, 78)

$. Length 4 mm. Background colour tawny with many dark areas; vertex with a wide black arcuate band covering most of the area, face almost entirely black, the facial marks running up slightly over the edge of the vertex; exposed thorax with heavy black spots (Text- fig. 14); legs and venter of body with irregular dark marks; membrane of wings almost hyaline, the veins bordered by dark brown striping. General structure typical for genus. Pygofer with only spines 1 and 2, spine 1 long and slender, spine 2 shorter, thicker, darker, and slightly hooked at apex, as in Text-tig. 24. Aedeagus with shaft long and slender (Text-fig. 46), the dorsal points minute and their bases below the dorsal contours.

Holotype 3 (pinned, abdomen in glycerine). SoLomMon ISLANDS: Guadalcanal, Kau Kau, 17.xi.1962, no. 2924 (P. Greenslade). In B.M.(N.H.).

Paratype. Same data, 1 ¢ (pinned, abdomen not cleared). In B.M.(N.H.).

This heavily marked species is a close relative of plebeius, which it resembles in structure of genitalia and from which it differs primarily in the dark colour pattern. The two specimens of the type series are both markedly smaller (4 versus 5 mm.) than any specimens of plebevus I have yet seen.

Exitianus kilimanus( Jacobi) comb. n. (Text-figs. 13, 23) Phrynomorphus kilimanus Jacobi, 1910 : 131. Kilimandjaro, Africa.

This species is readily distinguished by the twisted spine 2 of the pygofer and the very close proximity of the bases of spines 1 and 2 (Text-fig. 23). The head and pronotum are heavily marked with black (Text-fig. 13), much as in greensladet. The type ¢ (which I have seen) bears the collection data: Meru, 3,000—3,500 m Jan. (Sjostedt). Another female from the same locality on Mount Kilimandjaro bears a type label but a second female bearing a red ‘Allotypus’ label is evidently another species. I was not in the Stockholm Museum long enough to clear the genitalia of the type but the diagnostic position of the pygofer spines was readily visible on the dried specimen.

OKAHANDIA-GRovup

Among the extensive grass-inhabiting species collected in Africa by R. E. Turner, I found four species that form a distinctive group of Exitianus in which the plates are short and lack lateral spines, and the styles have the apex truncate and projecting nearly posteriorly, rather than being sharp and curved sharply laterally (Text-fig. 82) as is true of all other members of the genus,

LAE OLD WORLD EXATTLANGUS 17

Exitianus okahandia sp. n.

(Text-figs. 4, 55, 56, 57, 72)

3. Length 4 mm. Colour tawny with the following black marks: a fairly narrow arcuate band on vertex, small black spots on pronotum, several black spots on scutellum (Text-fig. 4), plus markings on the face, legs, and venter. General structure typical for genus. Plates ovate, without lateral spines, and about as long as wide (Text-fig. 55), extending considerably beyond apex of styles. Styles with apex broadened and truncate (Text-fig. 55). Pygofer of usual shape for genus, with only spines 1 and 2, both symmetrical, spine 2 shorter, thicker, and darker than spine 1 (Text-fig. 56). Aedeagus moderately long, its apex with only a slight and inconspicuous incision (Text-fig. 57).

Q. Length 5-5 mm. Colour and general structure similar to male. Apex of eighth sternite slightly produced in middle, the segment often with a small dark spot on each side near the middle of the lateral half of the segment (Text-fig. 72).

Holotype 3 (pinned, abdomen in glycerine). SoutTH AFRIca: Natal, Van Reenen, Drakensberg, I-22.1.1927 (R. E. Turner). In B.M.(N.H.).

Allotype 2. SouTHWEsT AFRICA: Okahandja, 10-16.i1.1928 (R. E. Turner). In B.M.(N.H.).

Paratypes. Same data as allotype 9, 3 g, 89, with inclusive dates 27.i—18. iii. I g, 1 in Illinois Natural History Survey collection, remainder in B.M.(N.H.).

Additional females that appear to belong to this species are from SOUTH AFRICA: Cape Province, Queenstown; Transkei, Untata. [TANZANIA] Tanganyika: Ukiriguru.

Exitianus turneri sp. n.

(Text-figs. 5, 52, 53, 54, 73)

6. Length 4-5 mm. Colour tawny, head with a black arcuate band slightly interrupted at middle (Text-fig. 5), face with a narrow mesal stripe in addition to more lateral markings; legs with dark marks; tegmina with all veins pale brown except costal margin which is con- trastingly cream coloured. General structure typical for genus. Pygofer moderately narrow at apex, which bears spines 1 and 2, both long and symmetrical, spine 2 thicker and shorter than spine 1 and with a marked crease at its base (Text-fig. 52). Plates short and obliquely truncate, without large lateral spines (Text-fig. 54). Style with flared apex, extending slightly beyond plate (Text-fig. 54). Aedeagus with shaft short and slightly sinuate (Text-fig. 53).

9. Length 5-5 mm. Colour and general structure similar to male except for the median stripe on the face which is usually paler. Eighth sternite (Text-fig. 73) with a fairly deep broad cleft apically, and with a black oblique bar on each side almost at the apical margin.

Holotype ¢ (pinned, abdomen in glycerine). SoutH AFrrica: Pondoland, Port St. John, 1-11. vi.1923 (R. E. Turner). In B.M.(N.H.).

Allotype 2. SoutH Arrica: Zululand, Gingindhlovu, 15.vi.1926 (R. E. Turner). In B.M.(N.H.).

18 H. H. ROSS

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Fics. 52-54. Male genitalia of Exitianus turneri. 52, pygofer; 53, aedeagus, lateral aspect; 54, genital assemblage, ventral aspect (one style omitted). a, aedeagus; c, connective; pb, pygofer; pi, plate; s, style.

Paratypes. Same data as allotype, 3 ¢ (one without abdomen), 4 9, with inclu- sive dates 9.vi-5.vii.1926; Natal, Kloof, 1500 ft., 1 9, vill.1926 (R. E. Turner). In B.M.(N.H.), one pair in Illinois Natural History Survey collection.

This species differs from the preceding in the shorter plates and other details of the genitalia, and in the contrasting pale costal margin of the tegmina. In this latter character it resembles the following species, from which it differs in details of the male genitalia and the female eighth sternite.

Exitianus zuluensis sp. n. (Text-figs. 61, 62, 63, 74)

3. Length 5 mm. Colour tawny, the head with an arcuate black band on vertex, and almost equally dark median stripe on the face. Lateral areas of face and legs with dark mark- ings. Tegmina with all veins brown except the costal margin which is cream colour. General

THE-OLD WORLD E XTTIAN GS 19

structure typical for genus. Pygofer (Text-fig. 63) with apical half strongly narrowed and upturned, ending in a single, dorsally-directed heavy black spine (presumably number 2). Plates short, broadly rounded at apex, and without large lateral setae (Text-fig. 61). Style with apex obliquely truncate, only moderately wide, extending almost to the apex of the plate (Text-fig. 61). Aedeagus with shaft relatively small, slightly angulate near middle, and narrowed at extreme tip (Text-fig. 62).

2. Length 5:5 mm. Colour and general structure same as male. Eighth segment with transverse posterior margin and with a pair of dark areas, one on each side, just in front of posterior margin (Text-fig. 74).

Fics. 55-63. Male genital parts of Exitianus spp. 55, 56, 57, E. okahandia (holotype) ; 58, 59, 60, E. natalensis (holotype); 61, 62, 63, E. zuluensis (holotype). 55, 58, 61, style, ventral aspect, with outline of plate shown beneath; 56, 60, 63, pygofer; 57, 59, 62, aedeagus, lateral aspect,

20 H:. H. ROSS

Holotype $ (pinned, abdomen in glycerine). SouTH AFRICA: Zululand, Eshowe, vi.1926 (R. E. Turner). In B.M.(N.H.).

Allotype 2. Same data as holotype. In B.M.(N.H.).

Paratypes. Same data as holotype, 3 9, 3 without abdomens, with inclusive dates, vi-vii. In B.M.(N.H.), 1 Qin Illinois State Natural History Survey collection.

This species is readily distinguished from the preceding by the peculiar upturned pygofer, and from the species that follows by its pale colour pattern.

66

Fics. 64-67. Male genital parts of Exitianus spp. 64, E. attenuatus (holotype), apex of right side of pygofer; 65, same (paratype), apex of right and left sides of pygofer, respectively; 66, same (paratype), quartering posterior view of aedeagal shaft; 67, E. pondus (holotype), apex of pygofer.

Exitianus natalensis sp. n.

(Text-figs. 6, 58, 59, 60, 75)

3. Length 4:5 mm. Base colour of dorsum chiefly tawny, vertex of head (Text-fig. 6) with a very wide black arcuate band, pronotum with lateral dark marks, mesonotum with lateral black triangles, face mostly black with tawny enclosed areas, the facial marks running up onto vertex; venter and legs mostly black; tegmina with veins brown and costa contrast- ingly cream coloured. General structure typical for genus. Pygofer (Text-fig. 60) with apical half strongly narrowed and upturned, the tip bearing spines 1 and 2, their bases contiguous, spine 1 very slender, shorter than 2, spine 2 long and massive. Plates fairly short, and apex broadly rounded, the lateral margin without large setae (Text-fig. 58). Style with relatively long apex, which is truncate and which almost reaches the margin of the plates (Text-fig. 58). Aedeagus with shaft relatively small, curved near middle, and narrowed at extreme tip (Text-

fig. 59).

Holotype ¢ (pinned, abdomen in glycerine). SoutTH AFrRIca: Natal, Van Reenen, Drakensberg, 1-22.1.1927 (R. E. Turner). In B.M.(N.H.).

Three females from Pondoland, Port St. John (B.M.(N.H.)), may belong to this species. In general colour pattern they resemble the male closely, but the prono- tum and mesonotum are both much darker and therefore there is a possibility that they may represent another undescribed species. The eighth sternite has the apical margin transverse and uniformly dark coloured (Text-fig. 75).

THE OLD WORLD EXITIANUS zai

mrad

[ y ne ay

De ee ae 74 [ = i te nee 75

Fics. 68-77. Apex of female eighth sternite. 68, E. plebeius (holotype); 69, E. indicus (holotype); 70, E. capicola (holotype); 71, E. taeniaticeps (Libya); 72, E. okahandia (allotype); 73, E. turneri (allotype); 74, E. zuluensis (allotype); 75, E. natalensis (Pondoland); 76, E. nanus (holotype); 77, E. africanus (holotype).

UNPLACED SPECIES

Bythoscopus africanus Walker, 1851 : 873. Type @ seen in B.M.(N.H.), from Sierra Leone (Morgan), 42, 31. The banded head (much as in Text-fig. 10) and the elongate ovipositor indicate that this species is a member of Exitianus, but the eighth sternite has a straight posterior margin and a pair of large posterior dark spots (Text-fig. 77). No males have been associated with this distinctive specimen, apparently representing a valid species.

Athysanus capicola Stal, 1855 : 99. Type 9 seen in the Stockholm Museum, from Meru, Neider. This is a typical member of the ¢aeniaticeps group, with head banded much as in Text-figure 9 and the eighth sternite with a median notch (Text-fig. 70). At present it is impossible to associate this specimen with a recognized species.

Eutettix pallida Evans and selbyi Evans, 1938 : 14. Both of these species were described from @ types collected in Queensland, and might prove to be additional junior synonyms of plebeius. The leafhopper fauna of Australia is as yet not well enough collected to make undue assumptions concerning the identity of females in this group of species.

SPECIES TRANSFERRED TO OTHER GENERA

Two species placed in Exitianus by Evans (1966) do not belong in