SSSW*

IP*

BULLETIN OF

THE BRITISH MUSEUM

(NATURAL HISTORY)

ENTOMOLOGY

VOL. XXIII

1969

BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1969

DATES OF PUBLICATION OF THE PARTS

No. I ..... 28 January 1969

No. 2 ..... 28 January 1969

No. 3 ..... 28 January 1969

No. 4 . . . . . .11 April 1969

No. 5 . . . . . .11 April 1969

No. 6 ...... 13 May 1969

No. 7 . . . . . .13 May 1969

No. 8 ...... 13 May 1969

PRINTED IN GREAT BRITAIN
BY STAPLES PRINTERS LIMITED

AT THEIR KETTERING,
NORTHANTS, ESTABLISHMENT

CONTENTS

ENTOMOLOGY VOLUME XXIII

PAGE

No. i. A revision of the genus Telipna Aurivillius (Lepidoptera : Lycaenidae).

By T. H. E. JACKSON i

No. 2. Acridoidea of the Galapagos Islands (Orthoptera). By V. M. DIRSH 25
No. 3. Diptera from Nepal. Sphaeroceridae. By J. C. DEEMING 53

No. 4. Revisional notes on African Charaxes (Lepidoptera : Nymphalidae)

Part V. By V. G. L. VAN SOMEREN 75

No. 5. A revision of the African species of Pseudorhynchus Serville (Orthop-
tera : Tettigoniidae). By D. R. RAGGE 167

No. 6. Studies on Australian Muscidae (Diptera) II. A revision of the Tribe

Dichaetomyiini Emden. By A. C. PONT 191

No. 7. A list of the Type-specimens of Odonata in the British Museum

(Natural History) Part II. By D. E. KIMMINS 287

No. 8. The Family-Group names of the scale insects (Hemiptera : Coccoidea).

By D. J. WILLIAMS 315

Index to Volume XXIII 343

A REVISION OF THE GENUS

TELIPNA AURIVILLIUS
(LEPIDOPTERA :

T. H. E. JACKSON

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. i

LONDON: 1969

A REVISION OF THE GENUS TELIPNA
AURIVILLIUS (LEPIDOPTERA : LYCAENIDAE) ^Njg

BY

T. H. E. JACKSON

Pp. 1-23 : 5 Plates

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. i

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
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hundred pages, and will not necessarily be completed
within one calendar year.

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

This paper is Vol. 23, No. i of the Entomo-
logical 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) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 28 January, 1969 Price Eighteen Shillings.

A REVISION OF THE GENUS TELIPNA
AURIVILLIUS (LEPIDOPTERA : LYCAENIDAE)

By the late T. H. E. JACKSON*

CONTENTS

SYNOPSIS

Page
3

INTRODUCTION ..........

SYSTEMATICS AND TAXONOMY OF THE GENUS

Group i ........... 4

Group ii ........... 16

Species incorrectly placed to Telipna ...... 20

ACKNOWLEDGMENTS ......... 21

REFERENCES ........... 21

INDEX ........... 22

SYNOPSIS

The genus Telipna Aurivillius is revised and four new species and three new subspecies are
described. All species are figured where possible.

INTRODUCTION

A REVISION of the genus Telipna Aurivillius is long overdue and, in addition, many
new species have been discovered since the publication of Seitz, M acrolepidoptera
of the World (1914). An attempt is made here to bring the genus up-to-date.

Insufficient is known of the range and distribution of each species to erect sub-
species and therefore, unless the affinity is obvious, all new forms are treated as
species in this paper. The species divide into two natural groups on the pattern
of the hind wing underside, as noted by Aurivillius, in Seitz, and this arrangement is
followed here.

The genitalia, except in a few instances, e.g. rothi Grose-Smith, are not diagnostic.

The habits of Telipna are interesting. The habitat is floor-level, always in forest,
along forest paths or in places where the undergrowth is not high, and heavy
shade is always preferred. Here they sit, usually at the ends of small dead twigs,
and fly short distances when disturbed, displaying the mimetic Agaristid-like pattern
and settling again and " disappearing ". They are, therefore, easy to capture, but
if missed for any reason, can fly fast right out of the area. They feed on plant sap
from creeper tendrils, leaving as soon as the sun reaches the tendrils. The early
stages are passed on the bark of trees, the larvae feeding on moss and lichens and
are not associated with ants. The larvae are black and densely hairy, similar to
those of Balacra Walker. The pupae are hidden under loose bark on the trunks
or on stumps and dead wood nearby.

* We learned with regret that, whilst this paper was being set up by the printers, the author was
murdered by intruders in his African home. Editor.

ENTOM. 23, i. i

4 T. H. E. JACKSON

All types are in B.M.N.H., unless otherwise stated. In the following text there
are several references to type material lodged in the Hamburg Museum. All this
material is presumed, on good authority, to be destroyed and neotypes are designated
where appropriate.

Note. In the descriptions of the fore wing underside all black marks, including
the two apical, are treated as costal streaks.

Group I

Hind wing beneath at the costal margin with two black transverse streaks
proximal to the median streak.

Telipna acraea (Doubleday, Westwood & Hewitson)

The three species or forms, acraea Doubleday, Westwood & Hewitson, bimacula
Plotz and albofasciata Aurivillius, form a complex which is exceedingly difficult to
unravel. The distributions are peculiar, i.e. acraea, west of the Niger in the true
West African Zone, occurring as echo Grose-Smith & Kirby (subapical spots orange
but not conjoined to the inner-marginal patch) in Ghana and elsewhere; bimacula,
east of the Niger River in Calabar and Cameroons (ex. Brit.) and again in a slightly
different form (ssp. nigrita Talbot) in Equateur and Kasai; albofasciata, type-
series from Fernando Po flies with bimacula in the Cameroons, occurs in a rather
different form in the Moyen Congo, without bimacula and again in a form, nearer the
typical, around Coquilhatville.

On the distribution, therefore, the latter cannot be a subspecies of bimacula
Plotz, as stated by Aurivillius, in Seitz and is here treated as a species.

On the other hand the affinities are obvious and there is little or no difference on
the underside.

Telipna acraea acraea (Doubleday, Westwood & Hewitson)
(PI. i, figs, i, 5)

Pentila acraea Doubleday, Westwood & Hewitson, 1852 : 504, pi. 77, fig. 6.

Type : Ashanti .
Liptena acraea (Doubleday, Westwood & Hewitson) Grose-Smith & Kirby, 1887 : (i), pi. i,

figs. 5, 6.

Telipna acraea (Doubleday, Westwood & Hewitson) Aurivillius (in Seitz), 1914 : 301, pi. 6ic.
Liptena echo Grose-Smith & Kirby, 1890 : 40, pi. X, figs. 4, 5.
Telipna bimacula echo (Grose-Smith & Kirby) Aurivillius (in Seitz), 1914 : 301, pi. 6id.

Subapical spots in the male as in echo Grose- Smith & Kirby and not conjoined to
the discal patch. Spots red. Specimens occur lacking the red subapical spot
altogether. See remarks under semirufa.

In the female the subapical and discal patches are always conjoined by a bar in

3,4-

Distribution. Confined to the true W. African Zone.

REVISION OF GENUS TELIPNA 5

Telipna acraea conjuncta ssp. n.

(PL i, figs. 6, 2)

A well defined subspecies occurs in W. Nigeria, to the Niger River and Port
Harcourt, differing from acraea acraea as follows:

$ Upperside. Fore wing. The subapical spot is very large and is joined distally to the
inner-marginal patch by a thin, sometimes thick, red bar. Rarely this spot is discrete.

9 Upperside. Fore wing. The black costal area invades the orange postdiscal band as far
as the base of space 3, leaving only a broad orange bar, joining the subapical to the inner-
marginal patch. Underside. As in acraea acraea.

Holotype <$. NIGERIA: Benin Prov., Ubiaja, viii.1955 (T. H. E. Jackson).

Allotype - Same data as holotype.

Distribution. NIGERIA: Benin, Ubiaja, Owerri, Port Harcourt.

Telipna acraea bimacula (Plotz) comb. n.
(PI. i, figs. 3, 7)

Pentila acraea Doubleday var. bimacula Plotz, 1880 : 199.

Type: Ashanti.
Telipna bimacula (Plotz) Aurivillius (in Seitz), 1914 : 301, pi. 6ic, d.

Type. Abo : Marz, Mungo : Mai. 2.
Liptena fervida Grose-Smith & Kirby, 1890 : 39, pi. X, figs. 1-3.

Types cJ and $. Cameroons.
Telipna fervida (Grose-Smith & Kirby) Aurivillius, 1898 : 257.

The more eastern subspecies of acraea. Subapical spots small, yellowish and
always divorced from the discal patch in the male. In the female, subapical spots
usually also divorced, though connected by a thin yellow line, but sometimes
indistinguishable from acraea acraea.

Judging from the figures in Grose-Smith & Kirby, 1890, fervida might be a separate
species, but Aurivillius, 1898, who must have seen it, since it was in the Staudinger
collection, sank it as a straight synonym of bimacula. I am informed by Dr.
Hannemann that the types are lost and it seems best, therefore, to follow Aurivillius.

Distribution. NIGERIA: east of the Niger River; Calabar; CAMEROONS (ex Brit.).
Holland, 1920, mentions a female from Niangara, Congo (ex Beige) which I am
inclined to doubt, as also, a male at Tervuren from Congo (ex Beige), Haut Lomami,
Kapanga, Kafakumba.

Telipna acraea nigrita ssp. n.

(PL i, figs. 8, 4)
Telipna acraea bimacula $ f. nigrita Talbot, 1935:70.

In Equateur and in Kasai (Congo, ex Beige) occurs a form of acraea which differs
sufficiently to warrant a subspecific name. By courtesy of the Hope Dept., Oxford,
two pairs of these have been examined and Talbot quotes eight females.

In the males the subapical spots are small and orange, always separated from the inner-
marginal patch and in the females the latter is always confined below the cell.

6 T. H. E. JACKSON

Neallotype $. CONGO STATE, S.W. Upper Kasai R. Distr., 1908 (Paul Laudbeck}.
(Oxford).

Holotype $. CONGO (ex Beige) ;Haut Congo, Basankusu, Bongandanga, 9.iv.i928
(Talbot's type), (Oxford). This locality is in Equateur Province.

Distribution. CONGO (ex Beige) : Equateur and Kasai.

Telipna albofasciata Aurivillius
(PL i, figs. 9, 13)

Telipna albofasciata Aurivillius, 1910 : 520.

Types. Fernando Po. (Genoa, no actual types were designated).
Telipna bimacula albofasciata Aurivillius; Aurivillius (in Seitz), 1914 : 301.
Telipna albofasciata Aurivillius; Stempffer, 1965 : 1450.

<$. Red areas very dark; discal patch in the type series confined below the cell, or just
reaching the median; wide black border, three (or two) white subapical spots in 4, 5 and 6.

?. Red areas more orange, restricted in the discal patch to the lower half of the cell; three
white subapical spots conjoined into a bar and a small white streak in 7; very occasionally
these spots are separate.

The type series consists of 4 ^ and 3 $, all in Mus. Civ. di Stor. Nat., Genoa and
all from Fernando Po.

The series from Moyen Congo differs in that the red discal area in fore wing of the
male covers half the cell as in the female, and the white subapical spots are larger
and broader. The underside is very similar to acraea and bimacula.

Distribution. FERNANDO Po; CAMEROONS; GABON; MOYEN CONGO: Etoumbi,
Kelle; CONGO (ex Beige): Equateur; Coquilhatville, Lake Tumba, Eala.

Telipna semirufa (Grose-Smith & Kirby)
(PI. i, figs, 14, 10)

Liptena semirufa Grose-Smith & Kirby, 1889 : 33, pi. VIII, figs. 5, 6.

Type. Gold Coast (Ghana).
Telipna bimacula semirufa (Grose-Smith & Kirby) Aurivillius (in Seitz), 1914 : 301, pi. 6id.

No subapical spots; discal and hind wing patches red, much as in acraea and confined below
the cell.

It has been found that, at least in the Cote d'lvoire, males of Telipna acraea
(Doubleday, Westwood & Hewitson) can occur without the red subapical spot in
fore wing above, thus closely resembling semirufa. There are, however, two unmis-
takeable characters separating the two species as follows the male of acraea has the
distal edge of red inner-marginal patch of fore wing above nearly straight and the
fore wing submarginal black border below invaded by the yellow ground colour in
spaces 3-5 inclusive. The male of semirufa, on the other hand, has the distal edge
of red inner-marginal patch of fore wing above deeply incised between space i and 2

REVISION OF GENUS TELIPNA 7

and the fore wing submarginal black border below entire. In the females the distal
edge of the fore wing yellow patch is more crenulate in acraea and the submarginal
black border of fore wing below differs as in the males. These characters can be
plainly seen in Smith & Kirby's figure of the type of semirufa.

Distribution. Confined to the W. African Zone, i.e. west of the Niger River.
Clench, 1965, is almost certainly correct in suspecting Aurivillius' locality of the
Congo.

Telipna rufilla (Grose-Smith & Kirby)
(PI. i, figs, u, 15)

Liptena rufilla Grose-Smith & Kirby, 1901 : 133, pi. XXVIII, figs. 4, 5.

Type. (J Nigeria, Warri (unique).
Telipna rufilla (Grose-Smith & Kirby) Aurivillius (in Seitz), 1914 : 301.

Similar to semirufa Grose-Smith & Kirby, but with larger red areas, and differing below.
The red area in fore wing covers most of the cell. $ unknown.

Aurivillius (in Seitz) placed this species to Group II, but it has nothing in common
with sanguined Plotz, nyanza Neave, etc. I think that the median costal streak
on the hind wing below is placed a little more distad than usual and that rufilla
belongs to Group I. It is peculiar that no further specimens of this species have been
taken.

Distribution. NIGERIA: Warri, i.e. west of the Niger River in the true W. African
Zone.

Telipna rothi (Grose-Smith)
(PL i, figs. 12, 16)

Liptena rothi Grose-Smith, 1898 : 353.

Type. Nigeria, near Warri.

Liptena rothi Grose-Smith; Grose-Smith & Kirby, 1901 : 133, pi. XXVII, figs, i, 2, 3.
Telipna rothi (Grose-Smith) Aurivillius (in Seitz), 1914 : 301.

A well known and well defined species in which the genitalia also differ.
The subapical and discal patches in both sexes broadly conjoined.
Distribution. West of the Niger; GUINEA: Mt. Nimba; GHANA; NIGERIA: Warri,
Benin, Lagos.

Telipna citrimacula citrimacula Schultze

(PI. 2, figS. 17, 12)

Telipna citrimacula Schultze, 1916: 141.
Type. Boenga, S.E. Cameroun. (Berlin).

All markings in both sexes orange; underside much more heavily marked than in the next
subspecies.

8 T. H. E. JACKSON

The type of citrimacula Schultze was sent to me by Dr. Hannemann from Berlin.
The type has no subapical spots but is otherwise identical above and below with
angustifascia Joicey & Talbot. The lack of subapical spots is a common aberration
and there is one here exactly similar, belonging to the Uganda subspecies from
Budongo Forest, Uganda, and another from Eala at Tervuren. There is no doubt
that these two subspecies are conspecific.

Distribution. CAMEROONS: Bitje, Djah River; Boenga; CONGO (ex Beige):
Equateur; Eala, Paulis; Coquilhatville.

Note: Monsieur Stempffer states that Schultze' s type of citrimacula was in the
Hamburg Museum and therefore was probably destroyed, but the specimen, received
from Dr. Hannemann in Berlin, was clearly labelled and must be the type.

Telipna citrimacula angustifascia (Joicey & Talbot) comb. n.

(PL 2, figS. 18, 22)

Telipna angustifascia Joicey & Talbot, 1921 : 77, pi. XIV, fig. 43.
Type (J. Bafwaboli, Congo (ex Beige).
Type $. Kasai, Congo (ex Beige).

The name angustifascia may be used for the population in the eastern Congo to
Kasai, and western Uganda. It is intermediate between typical citrimacula and the
Uganda subspecies neavei Baker.

In the male the subapical spots are larger than in the typical race and in the female the hind
wing patch is yellowish, as also the subapical spots of fore wing.

Distribution. E. CONGO (ex Beige) : from Kibale-Ituri to Kasai; Kasai; Equateur;
Kibale-Ituri ; Bafwaboli, Beni, Irumu; W. UGANDA: Budongo, Bwamba, etc.

Telipna citrimacula neavei Baker stat. n.

(PI. 2, figS. 19, 23)
Telipna neavei Baker, 1926 : 386.

Baker's neavei is merely a form in which the subapical patch is usually separated,
but sometimes conjoined to the discal area. It occurs commonly in all populations
in Uganda, and it is impossible to separate the two forms, but as it was described as
a species, the name neavei must be used for the eastern subspecies.

Ssp. neavei differs from typical citrimacula in that the subapical spots in the fore
wing of the male are larger and all spots in the female are pale yellow ; in citrimacula
they are orange. The subapical spots in the fore wing may be missing or small
in all subspecies. The normally red areas in the male are bright orange.

This is the race from eastern Uganda: Kampala, Mabira, Mpigi, Kamengo.

Type <3\ Q. E. UGANDA: Budongo Forest, Unyoro, 3,400 ft.
Distribution. E. UGANDA: In all forested areas.

REVISION OF GENUS TELIPNA g

Telipna sheffieldi Baker

(PI. 2, figS. 2O, 24)

Telipna sheffieldi Baker, 1926 : 387.
Type : Uganda, Toro, Mpanga Forest.

The subapical patch is much longer and broader than in citrimacula Schultze and the underside
differs as can be seen in the plate, particularly in the first costal streak in hind wing, which is
bent distad in all examples.

Distribution. Known only from UGANDA: Toro, Mpanga Forest (= Kibale
Forest).

Telipna aurivillii Rebel

(PL 2, figs. 29, 25)

Telipna aurivillii Rebel, 1914 : 262.

Type <J. Congo (ex Beige): Mawamba (Grauer). (Vienna).
Telipna rothioides Holland, 1920 : 214, pi. XII, fig. 7, syn. n.

Type $. Medje (Amer. Mus. Nat. Hist., New York).

A large species, not very similar to any other. In the male pale orange with
broad subapical band, conjoined to the discal area, in the female pale yellow.

Photos of the types have been received through the kindness of Dr. Kasy in Vienna.
The locality Mawamba is in the north-eastern Congo (ex Beige). The types were
collected by Grauer.

The figure of the female of rothioides, given by Holland, agrees exactly with the
photo of the female of aurivillii Rebel from Vienna, except that the former shows
some white markings along the fore wing costa below. It is significant that Holland,
who was a very careful observer, makes no mention of these white markings in his
description; he merely compares rothioides with rothi Grose-Smith, which has no
white at all. Otherwise his figure agrees exactly with the female of aurivillii, and
in the author's opinion there is no doubt they are synonymous.

Distribution. MOYEN CONGO: Souanke; CONGO (ex Beige): Kasai; Lulua;
Kibale-Ituri ; Mawamba, Beni, Irumu; Equateur; Coquihatville, Eala, Paulis.

Telipna aurivillii jefferyi ssp. n.

(PI. 2, figS. 26, 30)

Differs from the nominate subspecies by the much wider bands in fore wing, particularly
in the female. In jefferyi the band at vein 2 is 8 mm. wide in the male, and 10 mm. wide in the
female. In a. aurivillii these same bands are only 4 mm. wide, in both sexes.

Holotype $. UGANDA: Budongo Forest, ix 1934 (T. H. E. Jackson).
Allotype $. Same data, viii . 1958.

Named after the pioneer G. W. Jeffery, who was one of the first to capture this
species.

Distribution. UGANDA : In all forests.

io T. H. E. JACKSON

Telipna sulpitia Hulstaert

(PI. 2, figS. 31, 27)

Telipna sulpitia Hulstaert, 1924 : 114.

Type ?. Haut Uele, Madyu. (M.R.A.C., Tervuren.)

The type, of this species was kindly sent me by Monsieur L. Berger from Tervuren
and proves to be conspecific with examples from the N.E. Congo to S. Sudan,
which have long remained unnamed in B.M.N.H. The male has not been described.

cj. Upperside. Fore wing. Black, with red inner marginal area covering about half the
cell and extending well over half way in i and 2 ; three smallish, whitish yellow submarginal
spots. Hind wing. Red area covers most of the wing leaving a black marginal border 3 mm.
wide. Underside. As in the female. Length of fore wing 2 mm.

Neallotype $. S. SUDAN : Tembura, xii . 1922 (Jansori) .

In the $ the subapical band may occasionally be divorced from the discal patch
by a broad black line, part of which is always present.

Distribution. N.E. CONGO (ex Beige): Haut Uele, Madyu; S. SUDAN: Yambio,
Tembura, Aza Forest.

Telipna villiersi Stempffer
(PI. 2, figs. 28 and 32)

Telipna villiersi Stempffer, 1965 : 1450-52.

Type c?. Rep. du Congo. Sibiti. (Museum national de Histoire naturelle, Paris.)

Allied to acraeoides Grose-Smith and Kirby and kayonza sp. n., but smaller and
underside heavily marked as in kelle sp. n. ; red areas more orange. The female has
not been described.

o", $. Eyes black ; palps black with traces of white scaling ; frons black with two white lateral
spots; legs black with large white spots at the joints; antennae black above, checkered white
below; abdomen red with brown extremity $, brown $.

$. Upperside. Fore wing. Three white subapical spots in 4-6, broad and conjoined, a
very small white spot in 7. Otherwise as in the male. Underside. Markings, which are the
same as in the male, show through above in both sexes, and the cilia are only minutely white
spotted.

Length of fore wing; $ 21 mm.; $ 23 mm.

Note. The subapical white spots in the male may be conjoined into a bar as in
the type or divided into 3 separate white spots.

Neallotype $. REP. DU CONGO: Moyen Congo, Kelle, x.1963 (T. H. E. Jackson).

Distribution. REP. DU CONGO: Moyen Congo, Sibiti, Kelle, Etoumbi, Mayoumba;
CONGO (ex Beige) : Sankuru.

REVISION OF GENUS TELIPNA n

Telipna cameroonensis sp. n.

(PL 3, figs. 33, 37)

Nearest to atrinervis Hulstaert, but smaller and with white subapical spots in
fore wing.

A small weak species, flying with the common robust Acraea bimacula Plotz in
an obviously protective association.

cJ, ?. Eyes black; palps black; frons black, bordered white; legs black, white on the joints;
antennae black above, checkered white below.

<J. Upperside. Fore wing. Black with dark red basal and discal patch, not quite covering
the cell and leaving a wide black border even in space i ; up to three very small white subapical
spots, sometimes two only, or one, or rarely, absent altogether. Hind wing. The red discal
area is restricted, leaving a wide black border of 3 mm.

Underside. Fore wing. Seven costal streaks, that proximal to the subapical spots being
long and broad; margin narrowly black with white spots in each cellule, slightly wider in i
and 2. Hind wing. Two black proximal costal streaks, all narrow; a wide black border
enclosing two rows of white marginal and submarginal spots.

9. Upperside. Fore wing. Red areas orange, completely covering the cell and leaving
only a very narrow black costal margin; three white subapical spots. Hind wing. As in the
male. Underside. As in the male.

Cilia checkered white in both sexes.

Length of fore wing; <J 20 mm.; 25 mm.; the size is variable and there are much smaller
specimens and an occasional larger.

Holotype <. CAMEROONS (ex Brit.): Mamfe, vii.igGs (T. H. E. Jackson).
Allotype $. Same data, x . 1956.

Distribution. CAMEROONS (ex Brit.): Mamfe, Victoria; NIGERIA: Calabar Prov.,
Ndebiji.

Telipna atrinervis Hulstaert
(PL 3, figs. 38, 34)

Telipna atrinervis Hulstaert, 1924 : 113.

Type?. Kamerun : Bitje. (M.R.A.C., Tervuren).
Telipna ja Baker, 1926 : 388, syn. n.

Type cJ, $. Bitje, Ja River.
Telipna venanigra Baker, 1926 : 387, syn. n.

Type $. Bitje, Ja River.

The types of all the above have been examined and are considered to belong to the
same species. The differences are small and well within the legitimate limits in any
given species; moreover they all come from the same place and the differences are
all present in a series from Moyen Congo before me. One of the main differences
between ja Baker and venanigra Baker is the presence, in both sexes on the underside
in the longest and most distal costal black bar on the fore wing, of a large quadrate
black spot in space 3 of the former and its absence in the latter, but in the series from
Moyen Congo there is every variation in this spot, from large to complete absence.

A larger more robust species, completely lacking the apical spots in the male.

12 T. H. E. JACKSON

In the female three subapical spots, white or pale yellow, sometimes small, sometimes a bar.
The red $ or orange-yellow $ areas are restricted; in the male only covering half the cell of the
fore wing or less and in the female not reaching the upper discocellular and often with one or
two black costal spots invading the cell.

The holotype < of venanigra Baker becomes the neallotype <$ of atrinervis Hulstaert.

Distribution. CAMEROON: Bitje; REP. DU CONGO: Moyen Congo; Etoumbi,
Kelle, Impifondo ; CONGO (ex Beige) : Kapanga.

Telipna transverstigma H. Druce
(PL 3, figs. 39, 35)

$ Telipna transverstigma H. Druce, 1910 : 356, PI. XXXIII, fig. 2.

Type. S.E. Cameroons : Bitje.
Telipna transverstigma Druce; Aurivillius (in Seitz), 1914 : 302, pi. 6$a.

Red areas restricted in fore wing and not covering the cell, but in hind wing covering most of
the wing, leaving only a narrow black border; subapical white spots broad and united into a
bar in both sexes. On the underside 2 black, proximal costal spots in hind wing and in addition
a black streak, nearly horizontal, across the base of i.

Note. Aurivillius placed this species (in the author's opinion incorrectly) to his
Group II.

The male has not been described.

cJ. Upperside. Fore wing. Red area covers two thirds of the cell, then slopes obliquely,
distally ending squarely from midway space 2 to the anal angle; costa narrowly black with a
black spot entering the red area about midway along the cell; a prominent white subapical
bar in 4, 5 and 6. Hind wing. Entirely red, except for a narrow black border about i mm. wide.
Underside. As in the female; the two black bars running at right angles from the inner margin
of the hind wing very prominent.

Length of fore wing; the three males and one female before me, all from Moyen Congo, are
smaller than the female figured by Druce and reproduced in Seitz ; $ 18 mm., $ 21 mm.

Neallotype <$. REP. DU CONGO: Moyen Congo; Kelle. x. 1963 (T. H. E. Jackson}.
Distribution. CAMEROONS: Bitje; REP. DU CONGO: Moyen Congo; Kelle.

Telipna lotti sp. n.

(PI. 3, figs. 36, 40)

Telipna bimaculata [sic]. No author. Wilson, 1953 : 9 6 . pi- I 5- [mis-spelling of bimacula
Plotz.]

Allied to albofasciata Aurivillius, but much larger.

cJ, $. Eyes black; palps black; frons black with white lateral margins; legs black, white at
the joints; antennae black above, faintly checkered white below.

cJ. Upperside. Fore wing. Black; red discal patch restricted, covering only half the cell
and leaving a black distal margin 2 mm. wide in i and 3 mm. in 2; white subapical spots in
4-6, those in 4 and 5 broad, that in 6 small or absent. Hind wing. Red basal and discal

REVISION OF GENUS TELIPNA 13

patch rather restricted, leaving a black marginal border 4 mm. wide ; underside markings show
through. Underside. As in albofasciata; eight costal streaks in fore wing and two in hind
wing, proximal to the median streak.

$. Upperside. Fore wing. Black; orange-red discal patch as in the male, but covering
more of the cell and with two black costal spots invading it, one at cell end and one mid cell;
white subapical spots in 4-6 and a streak in 7, very broad and conjoined. Hind wing. As in
the male. Underside. As in the male.

Cilia in both sexes faintly checkered white.

Length of fore wing; < 25 mm.; $ 29 mm.

Holotype J. S. SUDAN: Lotti Forest (T. H. E. Jackson).

Allotype $. Same data.

Distribution. Known only from the type locality.

Telipna hollandi Joicey & Talbot
(PI. 3, figs. 41, 44)

Telipna hollandi Joicey & Talbot, 1921 : 80, pi. XIV, figs. 46, 47.
Telipna exsuperia Hulstaert, 1924 : 112.

Type. Congo (ex Beige) : $ Beni ; $ Kasai.

A large species with three white subapical spots in male and extensive red areas. In the
female the subapical white spots are very large and joined into a bar and the red area in hind
wing reaches within 4 mm. of the margin. Two proximal costal streaks in hind wing below;
the red areas in fore wing reach almost to the costa, leaving only a narrow black margin.

Distribution. CONGO (ex Beige): Kasai, Kibali-Ituri; Beni.

Telipna kelle sp. n.

(PI. 3, figs. 42, 45)

Nearest to hollandi Joicey & Talbot, differing as follows; red areas very dark in
both sexes.

<J. Upperside. Fore wing. Subapical white spots as in hollandi; red area more extensive,
reaching nearly to the margin in 2, and further beyond the cell; a thick black stigma in the red
patch at the cell end. Hind wing. The red areas are less extensive, reaching only to 3 mm.
from the margin; in hollandi 2 mm.; three white submarginal spots as in hollandi.

Underside. Fore wing. Much as in hollandi, but hind wing red area much narrower, leaving
a black border 4 mm. wide, as against 2-3 mm. in the latter; costal streaks as in hollandi, but
broader, the whole underside being darker.

$. Upperside. Subapical white spots not much larger than in the male in fore wing, other-
wise differing as in the male; red areas darker than in hollandi. Underside. Differs as in the
male, only the reduction of the red patch in hind wing is even more striking.

Length of fore wing; <$ 23 mm.; $ 25 mm.

Holotype ^. REP. DU CONGO: Moyen Congo, Kelle, x.1963 ( T. H. E. Jackson).

Allotype $. Same data.

Distribution. REP. DU CONGO: Moyen Congo: Kelle, Ketta.

14 T. H. E. JACKSON

Telipna acraeoides (Grose-Smith & Kirby)
(PL 3, figs. 43 and 46)

Liptena acraeoides Grose-Smith & Kirby, 1890 : 39.

cJ Liptena sanguinea (Plotz) Grose-Smith & Kirby, 1889; 2, pi. i, figs, i & 2.

Liptena acraea Doubleday, Westwood & Hewitson; Hewitson, 1866, 3 pi. 60, fig. 12.

Type : Angola.
$ Telipna acraeoides (Grose-Smith & Kirby) Aurivillius (in Seitz), 1914 : 301, pi. 6id.

This species has been often confused, but a photo of the type male and two more
males and two females have been received from B.M.N.H. and there is no longer
any doubt as to its identity. It is a good species, apparently confined to N. Angola.

The most important characters are: large species; subapical spots in male 3, of equal size
and well separated in 4, 5 and 6. In female large, broad, finely separated by the veins and an
extra streak in 7; red areas cover most of the cell in both sexes; prominent white submarginal
spots in hind wing above; below; costal streaks finer than usual and an extra series of black
spots in hind wing in the bases of i, 2 and 3, one or other of which may be missing, but always
two.

The series of five in B.M.N.H. are remarkably constant.

The figure of the female in Seitz is a good one and shows the large size and the
hind wing spots showing through from below. N. H. Bennett states there are none
from other localities in B.M.N.H. but that there are a further two males and eight
females from Angola.

The type female is missing, but no neotype is designated, since the type is probably
one of those in the series.

Distribution. ANGOLA : Canhoca, N'dalla Tando.

Note. There is a note in Talbot's handwriting under the male type saying;
" This must be the <$ type vide Rhop. Exot. Lye. Afr. p. 39 (1890) = specimen
figured as sanguinea <$ Lye. Afr. pi. i : figs, i, 2 (1889)".

Telipna kayonza sp. n.

(PL 4, figs. 47, 50)

Allied to acraeoides (Grose-Smith & Kirby), and to villiersi Stempffer but lighter
below.

<$, $. Eyes black; palps black; frons black with white lateral lines; legs black, small white
spots at the joints ; antennae black above and below.

cJ. Upperside. Fore wing. Black, with large red basal and discal area; three white sub-
apical spots in 46, that in 6 sometimes missing and always smaller ; red area to cell end and cover-
ing the base of 3 and most of i and 2, but not beyond the upper discocellular; black costal
border, a little over i mm. wide. Hind wing. The red area reaches to within 2 mm. of the

REVISION OF GENUS TELIPNA 15

margin, hence a narrow black border with no white submarginal spots; the underside costal
streaks show through.

Underside. Fore wing. Reddish yellow; five black costal streaks and a dot in the base, all
short except that proximal to the subapical white spots, which outlines these and is prolonged
by a spot in 3; narrow black margin, interrupted by the veins. Hind wing. Two fine black
proximal costal streaks, black margin narrow with two rows of white spots, but the submarginal
spot in 3 covered by the ground colour.

$. Upperside. Fore wing. As in the male but white subapical spots broad and conjoined
and a white streak in 7; a fine black stigma at cell end. Hind wing. As in the male.

Underside. As in the male.

Length of fore wing; ^ 24 mm.; $ 27 mm.

N. H. Bennett has examined the genitalia of this species and compared it to that
of acraeoides (Grose-Smith & Kirby). He states that although very complicated and
difficult to describe, it is quite different from acraeoides.

Holotype <. UGANDA: Kigezi, Kayonza, v.1954 (T. H. E. Jackson).

Allotype $. Same data, vii . 1952.

Distribution. Only W. UGANDA: Kigezi, Kayonza; Ankole, Kalinzu Forest.

Telipna plagiata Joicey & Talbot
(PI. 4, figs. 48, 51)

Telipna plagiata Joicey & Talbot, 1921 : 79, pi. XIV, figs. 44, 45.
Type $. Lower Butahu River, Semliki Valley, Congo (ex Beige).

Distinguished particularly by the heavily white spotted cilia between the veins.
A large species. Red areas cover all of cell and in female almost to costa.
The male has not been described.

$. Eyes black; palps black; frons black, centrally white; legs black, white at the joints;
antennae black above, checkered white below; abdomen rufous.

Upperside. Fore wing. Black; three small white subapical spots in 4-6; red inner marginal
patch from margin to upper discocellular, extending over most of space i, three-quarters of 2,
half of 3 and the base of 4. A small black triangular spot of ground colour at the cell end.
Hind wing. Black with large red area covering whole wing to within 3 mm. of the margin.
Cilia of both wings heavily spotted white in the interspaces.

Underside. Fore wing. Reddish yellow with six black costal streaks and the white subapical
spots showing through; margin spotted black with two very small white streaks in i and 2.
Hind wing. Of the usual pattern; two black proximal costal streaks; a double row of white
spots in the black margin ; red discal and postdiscal area reaching the margin in 5 and 6, centrally
whitish from costa to beyond cell; a small black streak on the inner margin in the base of i.

Length of fore wing; 25 mm.

Neallotype <. CONGO (ex. Beige) ; Ituri, Beni, (Kivu), iii. 1947 (T. H. E. Jackson).
Distribution. CONGO (ex Beige): Kibale-Ituri ; Semliki Valley, Beni, Itoa River.

16 T. H. E. JACKSON

Group II.

Hind wing below at the costal margin with one black transverse streak,
proximal to the median streak.

Telipna sanguined sanguined (Plotz)
(PL 4, figs. 49, 52)

Pentila acraea var. sanguined Plotz, 1880 : 198.

Type: Victoria (Berlin?).
Liptena anneckei Dewitz, 1886 : 427, pi. II.

Type : Mukenge.

Liptena sanguinea sanguinea (Plotz) Grose-Smith & Kirby, 1887 : 2, pi. I, figs. 3, 4.
Telipna mariae Dufrane, 1945 : 112 [$], syn. n.

Type $ : Congo (ex Beige) : Kivu ; Kamituga.
Telipna sanguinea f. bistrigata Aurivillius, 1925 : 7.

A well known species with a subspecies in Uganda ; subapical white spots variable
but usually large, especially in the female, the spot in 4 occasionally absent or very
small ; one proximal costal streak in hind wing below. The type of mariae Dufrane
has been received through the kindness of Monsieur L. A. Berger and the Director
of I.R.S.N.B., Brussels. It is a typical female of sanguinea Plotz. The type of
f. bistrigata Aurivillius has also been received from the Naturhistoriska Riks-
museum, Stockholm ; the fore wings in this specimen are badly rubbed, but from the
hind wings it appears to be a normal male of sanguinea. The underside also is
typical. It came from north of Lake Edward. The " rubbing " in the fore wings
may be due to lack of pigment on emergence from the pupa.

Distribution. NIGERIA: Calabar, Ikoni; CAMEROONS: Mamfe, Victoria; CONGO
(ex Beige): Kibale-Ituri, Epulu, Beni, Kamituga; Eala, Equateur; Paulis; Uele,
Haut Lornarni; Kapanga; Lake Edward.

Telipna sanguinea depuncta Talbot
(PL 4, figs. 53, 56)

Telipna sanguinea depuncta Talbot, 1937 : 59-

Type <J : Uganda, Tero Forest, Buddu ; $ : Mpanga Forest, Toro.

Distinguished by the much larger white subapical spots in both sexes ; in the female
these are broad and rounded, about 3 mm. wide.
Distribution. UGANDA : all forested areas.

Telipna medjensis Holland
(PL 5, fig- 61)

Telipna medjensis Holland, 1920 : 214, pi. XII, fig. 8 [$].

Type: Congo (ex Beige), Kibale-Ituri; Medje. (Carnegie Museum, Pittsburg).

This species seems to belong to Group II. The $ type is unique, but from the
figure and description it is a good species. On the underside fore wing as stated by

REVISION OF GENUS TELIPNA 17

Holland, " the entire apical region, save immediately on the costa, is immaculately
yellowish red ". This is not the case in any other species known to me. Further
there is only one costal streak in the fore wing below, that proximal to the white
subapical spots.

Distribution. CONGO (ex Beige) : Kibale-Ituri; Medje; only known from the type.

Telipna nyanza Neave
(PL 4, figs. 54, 57)

Telipna nyanza Neave, 1904 : 335, pi. i, fig. 19.
Type : Entebbe. (Hope Department, Oxford).

Red areas very large as in all this group, leaving, on fore wing, only a very narrow
black costal border and on hind wing a black marginal band of only 2 mm. Sub-
apical spots of fore wing small, that in 4 very small or absent, that in 6 shifted
basad as in ruspiniodes Schultze and katangae Stempff er, a small white streak in 7 ;
one proximal costal streak in hind wing below.

Distribution. CONGO (ex Beige): Lubudi and Lufupa Rivers; UGANDA: All
forests.

Telipna erica Suffert
(PI. 4.. figs. 55 and 58)

Telipna erica Suffert, 1904 : 41.

Type $ : Cameroon, Barombi (Berlin.)

Telipna erica Suffert; Aurivillius (in Seitz), 1914 : 302, pi. 6ie.
Telipna acraea $, f. nigra Suffert, 1904 : 42.

The type of this species was examined through the kindness of the authorities in
Berlin. It is a female and differs from all other species by the large white marginal
spots on fore wing above and by the very short costal streaks on hind wing below,
of which there are two including the median. On hind wing above two rows of
white spots, marginal and submarginal.

A male of this apparently very rare species was received from Moyen Congo and is
described as under :

cJ. Upperside. Fore wing. Black, with three white subapical spots in 4-6, that in 6 being
the largest; a white streak in 7; red area large, leaving only a narrow black costal border and
almost reaching the margin in i ; a fine black stigma at cell end. Hind wing. As in the female,
a double row of white spots marginal and submarginal . Underside. Agrees well with the female,
two costal streaks hind wing, all very short.

Length of fore wing; 22 mm.

Neallotype <. REP. DU CONGO: Moyen Congo, Kelle, x. 1962. (T. H. E. Jackson).

A specimen has been received recently by request from Berlin of Telipna acraea
?. f. nigra Suffert. It bears on the bottom of the pin an M.S. label stating, " Type,
Telipna acraea f. nigra Suffert ". Higher up on the pin is another M.S. label,

ENTOM. 23, I 2

i8 T. H. E. JACKSON

" Telipna erica Suffer! . Type ", and a red Type label with no number. The type
of erica, a female, has already been received from Berlin and photographed and this
specimen does not differ in any way from the type, moreover it comes from the same
place, i.e. Cameroon: Barombi. The synonomy above is given on the strength of
this specimen which is the only one which can be found in Berlin, but it is possible
that this is not Suffert's $ f. nigra.

Distribution. CAMEROONS: Barombi (Preuss); REP. DU CONGO: Moyen Congo,
Kelle.

Telipna consanguinea consanguinea Rebel

(PI. 5, figs. 59, 62)

Telipna consanguinea Rebel, 1914 : 262.

Type (J : Congo (ex Beige), Beni. (Vienna).
$ Telipna consanguinea Rebel; Schultze, 1923 : 1150.

Type $: Sud-Kamerum, Molundu. Probably destroyed in Hamburg.
Telipna consanguinea ab. $ exstincta Schultze, 1923 : 1150, fig. 49.

Type $: Moyen Congo, Ouesso probably destroyed in Hamburg.

Red areas very dark above and below; two white subapical spots above in fore
wing, in 5 and 6, occasionally, in the female, a small white streak in 7, all spots well
separated; one costal streak proximal to the median streak in hind wing below.
A photo of the type <$ has been examined, through the courtesy of Dr. Kasy, in
Vienna, and it agrees exactly with the species so named in the author's collection
and in B.M.N.H., except that the white apical spots are inclined to be larger in
Ugandan specimens.

The female was described by Schultze from S. Cameroon. I have not seen this
specimen and it is possible that it does not belong; a description of the true $ is
given below.

$. Upperside. Fore wing. Red areas slightly larger than in the male; two white subapical
spots a little larger than in the male; hind wing as in the male. A replacement Neallotype
is designated.

Length of fore wing; 25 mm.

Neallotype . UGANDA: Mpanga Forest, Mpigi, iv.i959 (T. H. E. Jackson).

Distribution. CONGO (ex Beige): Equateur; Eala, Kibale-Ituri ; Lake Tumba;
Beni; UGANDA: Bwamba.

Note. The specimen, a male, allocated here from Lake Tumba, appears to belong
to this species, but differs as follows ; it is larger and darker above and below ; on the
underside the black patch containing the white subapical spots is larger and the
costal hind wing streaks are larger and thicker.

The specimens from the localities cited by Schultze in S. Cameroon have not been
examined. They were probably in Hamburg and have been destroyed, and the
locality sounds most unlikely. $ ab. exstincta was said to differ by the reduction of
the triangular costal streak in fore wing below.

REVISION OF GENUS TELIPNA 19

Telipna consanguinea ugandae Baker comb. n.
(PI. 5, figs. 60, 63)

Telipna erica ugandae Baker, 1926 : 388.
Type: Uganda, Toro.

This insect is a subspecies of consanguinea Rebel and has nothing to do with erica
Suffert. It is distinguished by the larger subapical spots, especially in the female,
in which they are united into a broad band.

Distribution. UGANDA : all forests except Bwamba where the nominate subspecies
occurs.

Telipna katangae Stempffer
(PI. 5, figs. 64, 66)

Telipna katangae Stempffer, 1961 : 9.

Types: Congo (ex Beige), Kapanga. (coll. Stempffer, Paris).

Very similar to consanguinea Rebel differing as follows; three subapical spots in 5-7, with a
minute dot in 4, that in 6 shifted basad out of line; cilia more conspicuously white, so that in
hind wing there are two rows of white spots, marginal and submarginal; slightly larger than
consanguinea; underside very similar, but distal black margin in fore wing enlarged in i and 2,
enclosing two triangular spots of the ground colour.

A small series from near Coquilhatville appear to belong here.
Distribution. CONGO (ex Beige): Haut Lomami; Kapanga, Sandoa; Equateur;
Kafakumba, Coquilhatville, Lake Tumba, Paulis, Eala.

Telipna ruspinoides Schultze
(PI. 5, figs. 65, 67)

Telipna ruspinoides Schultze, 1923 : 1151, fig. 50.
Type : Sud-Kamerun, Owong.

Similar to katangae Stempffer, but subapical white spots in fore wing large in 4-6, occasionally
a trace of a streak in 7 in the female, the large spot in 6 shifted basad out of line ; cilia very con-
spicuously white; on the underside the first costal streak is divided into two spots.

Schultze based this species on two examples () from Owong and Kom Felle
in S. Cameroon, but did not fix a type. As these specimens were probably destroyed
in Hamburg a NEOTYPE male is proposed here.

The female has not been described.

$. Fore wing costal band narrower than in the Neotype <$ and not invading the cell.
Hind wing marginal band broader than in the g, the orange-red area more angled towards
the outer margin.

NEOTYPE <. CAMEROONS: Bitje, Ja River, dry season (G. L. Bates).
Neallotype $>. FR. EQUAT. AFR. : Moyen Congo, Feb., 1959 (T. H. E. Jackson).

20 T. H. E. JACKSON

Distribution. CAMEROONS (ex Brit.): Mamfe; NIGERIA: Calabar, Ndebije;
S. CAMEROON : Owong, Kom Fe'lle.

Note. In Peters' Check List, 1952, there are two entries which are quite inexpli-
cable; p. 91, Telipna acraea f. rothi Grose-Smith and Telipna erica parva Kirby.
The former is a good species and the latter is the eastern subspecies of Ptelina carnuta
Hewitson, parva is mentioned again in its correct place. I have been unable to
find the following; acraeoides laplumei Devos, 1919.

Species incorrectly placed to Telipna.
Ptelina carnuta Hewitson

Pentila carnuta Hewitson, 1873 : 125

Type : Gabon.

Telipna carnuta (Hewitson) Aurivillius (in Seitz), 1914. PL 6ie.
Ptelina carnuta (Hewitson) Clench, 1965 : 271-273.

Dttrbania carnuta (Hewitson) Grose-Smith & Kirby, 1895 .-91, PL XXI, figs, i, 2.
Telipna kamitugensis Dufrane, 1945.

Type: Congo (ex Beige), Kamitugu.

The type of Telipna kamitugensis Dufrane has been examined through the courtesy
of the Director of I.R.S.N.B., Brussels. It lacks an abdomen, but appears to be a
male of Ptelina carnuta Hewitson. Above it is typical of this species, but has a
small black discocellular spot on hind wing above, present also in a female in this
collection. Below the hind wing marginal band is broader, but I do not think this is
more than an individual variation.

Ptelina carnuta parva (Kirby)

Liptena parva Kirby, 1887 : 362.

Liptena parva Kirby; Grose-Smith & Kirby, 1888 : 15, figs. 1-4.

Ptelina subhyalina (Joicey & Talbot)
Telipna subhyalina Joicey & Talbot, 1921 : 78.

Telipna actinotina Lathy

Telipna actinotina Lathy, 1903 : 194.

Type : Nigeria, Ogruga.

Telipna actinotina Lathy Aurivillius (in Seitz), 1914, PL 62a.
A craea actinotina (Lathy) Peters, 1952 : 88.

T. G. Howarth of the B.M. N.H. states that Lathy misplaced the species and this
was apparently corrected by Neave, according to a label in the collection, who placed
it to the genus Acraea. Peters gives the date as 1915 for this correction, but no
reference to this can be found. Three examples of this rare species were taken by
Mr. R. G. T. St. Leger in December, 1959 at Ahoda, E. Nigeria.

REVISION OF GENUS TELIPNA 21

Pentila rogersi (H. Druce)

Telipna rogersi H. Druce, 1907 : 78.

Type: 14 m. from Mombasa, Rabai. (Oxford.)
Pentila rogersi (H. Druce) Stempffer, 1954 : 7-8.
Telipna rogersi H. Druce; Aurivillius (in Seitz) 1914 : 302.

This species is now in Pentila.

ACKNOWLEDGEMENTS

My thanks are due to Monsieur L. A. Berger of M.R.A.C., Tervuren, the Director,
I.R.S.N.B., Brussels, and to Dr. H. Hannemann of Berlin for the loan of their types;
to Dr. F. Kasy of Vienna for photographs of their types ; to the Director, Natur-
historiska Riksmuseum, Stockholm for the loan of the type of sanguinea bistrigata
Aurivillius; to the Director, Mus. Civico Storia Nat., Genoa, for photographs of the
series of Telipna albofasciata Aurivillius ; to the Curator, University Museum, Oxford.
Without the co-operation of the above this paper could not have been written. To
the Keeper and staff of the B.M.N.H. and particularly to Captain N. D. Riley, C.B.E.,
T. G. Howarth, G. Tite, and N. H. Bennett for much help in arranging the plates,
completing the distributions etc. ; and finally to Monsieur H. Stempffer of Paris for
reading and correcting this paper.

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22 REVISION OF GENUS TELIPNA

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SCHULTZE, A. & AURIVILLIUS, C. 1923. Ergebnisse der Zweiten Deutschen Zentral-Afrika-

Expedition 1910-1911. Lepidoptera III. 1 (17) : 1113-1194, figs.
STEMPFFER, H. 1961. Contribution a l'6tude des L6pidopteres Lycaenidae de 1'Afrique

6quatoriale. Annls Mus. r. Afr. centr. (8) Sci. Zool. 94 : 73 pp., figs.
1965. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descar-

pentries. Bull. Inst. Fr. Afr. noire 27, s6r. A (4) : 1449-1465.
SUFFERT, E. 1904. Neue afrikanische Tagfalter. D. ent. Z. Iris 17 : 12-107, figs.
TALBOT, G. 1935. New species and forms of African Lycaenidae. Entomologist's mon. Mag.

71 : 69-78, 115-127, 147-153, 3 pis, 3 figs.
1937. New African Lycaenidae & Nymphalidae and two new Diestogyna (Lepidoptera).

Trans. R. ent. Soc. Lond. 86 : 59-72, pis. i, 2.
WILSON, C. E. 1953. Butterflies of the Southern Sudan. Sudan Notes Rec. 34 : 73-103, pi. 15.

INDEX

Acraea, 20 fervida, 5
acraea, 4, 5, 14, 17

acraeoides, 10, 14, 15 hollandi, 13
actinotina, 20

albofasciata, 4, 6, 12, 13 ja, n

angustifascia, 8 jefferyi, 9
anneckii, 16

atrimacula, 7, 8 kamitugensis, 20

atrinervis, u katangae, 19

aurivillii, 9 kayonza, 10, 14

kelle, 10, 13
Balacra, 3

bimacula, 4, 5, 6 laplumei, 20

bimaculata, 12 Liptena, 4, 5, 6, 7, 14, 16, 20

bistrigata, 16 lotti, 12

cameroonensis, n mariae, 16

carnuta, 20 medjensis, 16
conjuncta, 5

consanguinea, 18, 19 neavei, 8

nigra, 17, 1 8

depuncta, 16 nigrita, 4, 5

Durbania, 20 nyanza, 7, 17

echo, 4 parva, 20

erica, 17, 18 Pentila, 4, 5, 16, 20, 21

exstincta, 18 plagiata, 15

exsuperia, 13 Ptelina, 20

INDEX 23

rogersi, 21 subhyalina, 20

rothi, 3, 7, g, 20 sulpitia, 10
rothioides, 9

rufilla, 7 transverstigma, 12
ruspinoides, 19

ugandae, 19

sanguinea, 7, 14, 16

semirufa, 4, 6, 7 venanigra, n

Sheffield!, 9 villiersi, 10

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PLATE 5

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. 5.

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp.284; 348 Text-figures. August, 1965. 6.

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ;
18 plates, 270 Text-figures. August, 1965. 4 45.

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October,

1965- 35s.

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 ;
475 Text-figures. November, 1965. 2 155.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae.
Pp. 129 ; 328 Text-figures. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera : Coccoidea). Pp. 168 ; 43 Text-figures. February, 1967.

335.

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the
world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146
Text-figures, 9 maps. February, 1967. 3 los.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 IDS.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa-
locera). Pp. 322; Coloured frontispiece, 348 text-figures. August, 1967. 8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collection's Pp. 184;
82 Text-figures. May, 1968. 4.

12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with an
Account of their World Distribution (Lepidoptera : Drepanidae). Pp. 151:
293 Text-figures, 14 plates. November, 1968. 5.

PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, B A RTH OLOMBW PRESS, DORKING.

,

3^

2 * JAN
ACRIDOIDEA OF THE GALAPAGO:

ISLANDS (ORTHOPTERA)

V. M. DIRSH

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. 2

LONDON : 1969

ACRIDOIDEA OF THE GALAPAGOS ISLANDS

(ORTHOPTERA)

BY

V. M. DIRSH

Anti-Locust Research Centre, London

Pp. 25-51 ; 7 Plates, 79 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 23 No. 2

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

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

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

This paper is Vol. 23, No. 2 of the Entomological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 28 January, 1969 Price i is

ACRIDOIDEA OF THE GALAPAGOS ISLANDS

(ORTHOPTERA)

By V. M. DIRSH

SYNOPSIS

The Acridoidea of Galapagos Islands are revised. The valid species are redescribed and
lectotypes are selected. The systematic position of the species is reviewed.

CONTENTS

Page

INTRODUCTION ........... 27

ACKNOWLEDGEMENTS ......... 27

NAMES OF THE ISLANDS ......... 28

ZOOGEOGRAPHICAL REMARKS ........ 28

TAXONOMY ........... 30

REFERENCES ........... 50

INTRODUCTION

SINCE Darwin's travels the Galapagos Islands have deservedly become famous as
a sanctuary for numerous endemic species of various classes of the animal kingdom.
Of them all, insects have been the least studied. However, lately this gap has
begun to be filled (Linsley and Usinger, 1966).

This paper is an attempt to review the Acridoidea fauna of these islands on the
basis of much material which was not available to earlier authors. The views of
earlier authors are reviewed in the light of the latest taxonomic studies, which have
sometimes demanded redescriptions of the types and revaluation of the taxonomic
status of some of the species and subspecies. All the genera and species known from
these islands are mentioned; they comprise 4 genera and 9 species.

ACKNOWLEDGEMENTS

I wish to express my sincere gratitude to the following colleagues who have helped
me by providing the types and material for this work: Dr. Ch. L. Hogue, Los Angeles
County Museum of Natural History; Dr. P. H. Arnaud, Jr., California Academy of
Sciences; Mr. D. C. Rentz, University of California, Berkeley; Dr. A. B. Gurney,
U. S. National Museum, Washington; Dr. H. E. Evans, Museum of Comparative
Zoology, Harvard University; Dr. P. J. Persson, Naturhistoriska Rijksmuseum,
Stockholm; Dr. H. Radclyffe-Roberts, The Academy of Natural Sciences of Phila-
delphia; Dr. H. Schrader, Naturmuseum und Forschung-Institut, Senckenberg;
Dr. F. Kuhlorn, Zoologische Sammlung des Bayerischen Staates; Mr. J. P. Don-
caster, Keeper of the Department of Entomology of the British Museum (Natural
History) ; and Dr. D. R. Ragge of the same Department. Finally, I would like to
thank Dr. T. H. C. Taylor for reading and editing the manuscript.

ENTOM. 23, 2 4

28

V. M. DIRSH

NAMES OF THE ISLANDS

Because most of the islands of the Galapagos Archipelago have more than one
name (English and Spanish) and sometimes two different islands (Gardner Is.)
have the same name, a list showing the names and synonyms is given below. In
this paper the English names are used.

English names

Abingdon

Albermarle

Barrington

Bartholomew

Bindloe

Champion

Charles

Chatham, Dassigney

Culpepper

Daphne Major

Daphne Minor

Duncan, Dean

Eden

Gardner-near-Charles

Gardner-near-Hood

Hood

Indefatigable, Norfolk, Porter

James, York
Jervis

Narborough
North Seymour

South Seymour
Tower, Ewres
Wenman

Spanish names

Pinta, Geraldino
Isabela, Santa Gertrudis
Santa Fe
Bartholome
Marchena, Torres

Santa Maria, Floreana
San Cristobal, Grande
Darwin, Guerra

Pinzon

Espariola

Santa Cruz, Bolivia, Chavez,

San Clemente, Valdez

San Salvador, Santiago, Gil, Olmedo

Rabida

Fernandina, Plata

Seymour

Plaza

Baltra

Genovesa

Wolf, Gasna, Nunez

ZOOGEOGRAPHICAL REMARKS

The finches observed by Darwin on the Galapagos Islands attracted the attention
of the whole scientific world. Their specialization was attributed to the insular
isolation. Van Denburgh (1914) and many other authors noticed that land tortoises
and iguanas from different islands, although of the same species, could be distin-
guished and their origin in each case from a particular island or even part of an island
could be determined. The differences within species are probably due not only to
physical isolation but also the ' territorial instinct ' of these animals. They are
attached to their breeding places and there is evidence (Eibl-Eibesfeldt, 1960) that
every individual claims its own bit of territory and even fights for it, protecting its

ACRIDOIDEA OF GALAPAGOS ISLANDS 29

rights. This same territorial instinct has been observed also in crabs. In Acridoidea,
species of the genus Schistocerca are good fliers and can easily cover the distances
between the islands, and the territorial instinct has never been observed in Acridoidea.

Snodgrass (1902) erected several subspecies of Schistocerca melanocera (Stal,
1861), and Scudder (1893) several subspecies of Schistocerca literosa (Walker, 1870),
but both authors based their subspecies on insufficient material and considered the
range of variability of the species as racial; when more material from various island
was available it became clear that the ' subspecies ' fit into the range of variability
of the species from a single island. Hebard (1920) and the present author synony-
mized these subspecies and one species.

A different picture arose when the genus Halmenus was studied. The species of
this genus are brachypterous or micropterous and cannot fly. All of them are
morphologically very similar, but the populations from different islands can be
distinguished. It is clear that the various forms of Halmenus could appear only as a
result of physical isolation, and since the males' internal genital organs are very
similar, the isolation has probably not been of long duration.

In the case of the genus Sphingonotus the position is rather similar to Schistocerca.
All the species are macropterous and can fly. However, the species of Sphingonotus
are small and according to observation they are not long-distance fliers but can be
carried by the wind. In the Galapagos Islands much more variability can be
observed in Sphingonotus than in Schistocerca, and possibly there is already a
tendency to form races on the different islands.

Concerning the genera Closteridia (C. bauri Scudder, 1893) and Desmopleura
(D. concinna Scudder, 1893), the former is known by the type only and the latter
by only a few specimens.

How did Acridoidea populate the Galapagos Archipelago? The islands may have
been nearer to the continent, or they may even have been connected with it and
have separated by Continental Drift (Cox, 1966). However, in a later paper it was
suggested that they are of rather recent origin. Cox and Dalrymple (1966), on the
basis of palaeomagnetism and Potassium- Argon measurements, determined that the
rocks exposed above sea level were extruded during the Brunhas Matuyama polarity
epochs and this process took place only about 2-4 million years ago. In any case
the islands could be populated from the continent only, the shortest distance of
which now is about 500 miles. Some species, such as Schistocerca melanocera and
literosa, could easily fly such distances, particularly if winds were favourable. The
small, winged species of Sphingonotus could travel the same way. Wingless and
short-winged species could be carried to the Archipelago by the oceanic currents,
with debris.

Orr (1966) observed that vegetation debris was carried by the rivers of South
America down to the coast and into the sea, where they formed raft-like masses up
to 10 ft. in diameter. There they were caught by the South Equatorial Current and
Humboldt Current and, according to Orr's calculation, could theoretically reach
the Galapagos Islands in less than two weeks.

If we accept the estimate of Cox and Dalrymple (1966) concerning the time of
origin of the Galapagos Islands, then the evolutionary changes in the Galapagos

30 V. M. DIRSH

Acridoidea took a remarkably short time. There is on the continent no species of
Schistocerca very closely related to the Galapagos species. The same applies to
the three genera Desmopleura, Closteridia and Halmenus and partly to Sphingonotus.
The species of the latter have a very near relative on the Puna Islands, which are
only about two miles from the Ecuador shore.

SCHISTOCERCA Stal, 1873

Acridium (Schistocerca) Stl, 1873.
Schistocerca St&l, 1876.

The genus Schistocerca belongs to the subfamily Cyrtacanthacridinae of Acrididae.
It contains numerous species on the American continent and one in the Old World
(Africa and South-west Asia), but occasional specimens sometimes fly, aided by air
currents, far from the usual area of distribution.

In the Galapagos Archipelago, according to the studies presented in this paper,
there are only two species of this genus. How they populated these islands we can
only guess. It could have been by free flight or by being carried by the wind or
oceanic currents. They certainly came from the American continent, and probably
from the northern part of South America. Apparently there are in America no
species closely related to the two Galapagos species. However, it must be noted that
the genus Schistocerca is probably of comparatively recent origin and is expanding,
forming numerous forms and variations. An indication of this possibility is that
the internal genitalia are very similar in all the species of the genus, in America,
the Old World and the Galapagos Islands. The variability within the species,
particularly the American ones, is very great. The external morphological charac-
ters are as similar between species as the phallic complex and in many cases only
morphometric characters, pattern and coloration serve to distinguish the species.

There are not in America or in the Old World any genera which can be considered
to be closely related to Schistocerca; the main characters the flattened plate-like
cerci in combination with the excised distal end of the subgenital plate in males
do not occur in such a combination in other genera of the subfamily.

However, in the genus Halmenus, which occurs only in the Galapagos Islands,
there is a small excision on the male subgenital plate, though smaller than in Schis-
tocerca, and the cerci are flattened as well, but of angular shape, unlike those of
Schistocerca. The internal genitalia are so similar to those of Schistocerca that they
can be easily confused, and the spermatheca is also extremely similar. The meso-
sternal lobes of Halmenus are not fully typical of Cyrtacanthacridinae, but are not
sufficiently atypical to exclude the genus from this subfamily.

There is thus much scope for speculation concerning the origin of Schistocerca
and its relationship with other genera, but the author feels that the time is not at
present ripe for it. The American genera Abracris and Osmilia (both long- winged)
occupy the same indefinite position as Halmenus and they too should perhaps be
placed into Cyrtacanthacridinae. This would complicate the question of the rela-
tionship of Schistocerca even more, particularly because hitherto it has been regarded

ACRIDOIDEA OF GALAPAGOS ISLANDS 31

as the only representative of the subfamily Cyrtacanthacridinae in the whole of
America.

It is obvious that further study of the South and Central American fauna is
urgently needed before definite conclusions can be reached.

The distribution of the two species of the genus on the Galapagos Islands is rather
peculiar. On the information at present available it can be stated (but not yet
explained because of lack of information) that 5. melanocera inhabits almost all the
islands of the Archipelago, while S. literosa occurs only on islands on the south-
eastern fringe of it (see map).

Schistocerca melanocera (Stal, 1860)
(PI. I, Text-figs. r-i2, Map i)

Acridium vnelanocerum Stal, 1860 : 326. Type $. ' Insulae Galapagos '. Naturhistoriska

Rikmuseet, Stockholm.
Acridium tibiale Walker, 1870 : 582 (Kirby, 1910 : 457). Type <J. ' West coast of America '.

British Museum (Natural History), London.

Acridium (Schistocerca) melanocerum Stal, 1860 : 326 (Stal, 1873 : 65).
Schistocerca melanocera (Stal, 1860) (Bruner, 1889 : 193).
Schistocerca melanocera minor Snodgrass, 1902 : 421 (Hebard, 1920 : 323). LECTOTYPE $.

' Tagus Cove, Albemarle '. By present selection from 6 <J and 6 $ syntypes. Los Angeles

County Museum of Natural History, U.S.A.
Schistocerca melanocera pallida Snodgrass, 1902 : 422 (Hebard, 1920 : 323). LECTOTYPE $.

' Barrington Is.' By present selection from 5 $ and 2 syntypes. Los Angeles County

Museum of Natural History, U.S.A.
Schistocerca melanocera lineata Snodgrass, 1902 : 423 (Hebard, 1920 : 323). LECTOTYPE $.

' Albemarle Is. Iguana Cove.' By present selection from 7 $ and 8 $ syntypes. Los Angeles

County Museum of Natural History, U.S.A.
Schistocerca melanocera immaculata Snodgrass, 1902 : 423 (Hebard, 1920 : 323). LECTOTYPE

cj. ' Indefatigable Is.' By present selection from 3 <$ and 3 $ syntypes. Los Angeles

County Museum of Natural History, U.S.A.
Schistocerca intermedia Snodgrass, 1902 : 431. Syn. n. Type $ (lost). ' Duncan Island.'

Neotype $. ' Pinzon Island.' = Duncan Is. California Academy of Sciences, U.S.A.
Schistocerca intermedia borealis Snodgrass, 1902 : 435. Syn. n. LECTOTYPE $ by present

selection from 2 (J and i $ syntypes, Bindloe Is. Los Angeles County Museum of Natural

History, U.S.A.

All subspecies of Schistocerca melanocera were rightly synonymized by Hebard,
1920. They cannot be separated either by morphological, colour or morphometric
characters. Using a few specimens from the different islands, the difference, even
if it is very small, can sometimes be detected, but as soon as a series is available the
material becomes a continuous series, which cannot be separated according to the
islands' sources. For example, the subspecies melanocera melanocera (sensu Snod-
grass) from Charles Island possesses a yellowish spot on the lateral lobe of the prono-
tum. On the basis of this spot, Snodgrass (1902) considered it to be a subspecies.
On Charles Island most specimens possess this spot, but some do not. On the other
hand some specimens from Barrington, Albemarle, North and South Seymour, James,
Narborough, Jervis, Darwin and some other islands also possess it. This invalidates
the character as a subspecific one.

32 V. M. DIRSH

It is well known that coloration and partly pattern depend very much not only
on the genotypical basis of the population but also on the colour of the environ-
ment in which the specimens grow and live.

The same considerations are applicable to the other characters used by Snodgrass
for differentiation of the subspecies. They are even weaker in this respect than the
yellowish spot on the pronotum.

The newly synonymized Sch. intermedia belongs to the same category. The type
of it is lost, but the topotypical material from Duncan Is., which agrees with Snod-
grass' description, differs from the typical melanocera only in the paler greyish general
coloration and in the more pronounced spots on the tegmina. No morphological
or morphometric differences were found. This coloration is probably the result
of the specimens having developed on a lighter-coloured background. To attribute
to them specific or even subspecific value is not possible for the above reasons and
also because some of the specimens from Barrington, Chatham and Jervis Islands
are practically identical with the intermedia from Duncan Island.

The subspecies Sch. intermedia borealis is characterized only by the more developed
pattern of the tegmina ; in all other respects it is the same as melanocera and intermedia.

The opinion was expressed (Hebard, 1920) that Sch. intermedia intermedia is an
intermediate form between Sch. melanocera and liter osa. But a morphometric study
showed that they can be differentiated, the hind femur being relatively longer and
narrower in melanocera than in literosa.

The variability in the size of the body of Sch. melanocera is rather great, but it
cannot be associated with any particular island.

Redescription. <. Large. Integument slightly rugose and pitted. Antenna longer
than head and pronotum together. Fastigium of vertex short, obtuse-angular, with shallow
oval depression in middle; frontal ridge low, with margins parallel and slight depression at
ocellus. Dorsum of pronotum subcylindrical, slightly widening backwards and slightly constric-
ted in anterior half; median carina low, linear, crossed by three deep sulci; metazona slightly
longer than prozona, its posterior margin obtuse-angular to almost rounded; lateral lobes in
prozona concave. Prosternal process from narrow cylindrical to subconical, at apex rounded.
Mesosternal interspace about twice as long as its width in middle, slightly widening in front.
Tegmina and wings far exceeding end of abdomen. Opening of tympanal organ almost semi-
circular, membrane deeply set forming wide angle with body wall. Hind femur comparatively
long and narrow, average ratio of length to maximum width 4-87. Hind tibia as long as femur.
Supra-anal plate elongate-angular, in middle of basal part with longitudinal sulcus merging
with lateral carinulae, sides upcurved, forming ridge-like elevations. Cercus slightly narrowing
towards apical part ; upper half of apex angularly protruding. Subgenital plate in profile short,
narrow, at apex attenuate.

Phallic complex: of usual structure for the genus; apical valves of penis shorter than valves
of cingulum.

Coloration: in general, from shiny dark brown to dull brownish grey. Antenna blackish;
face as general coloration, with longitudinal yellowish or ochraceous stripes on sides of frontal
ridge and on genae ; prozona of pronotum blackish or brown ; metazona from yellowish to dull
ochraceous ; in upper part of lateral lobe of prozona sometimes a small yellowish spot ; head above,
with yellowish narrow stripe, which merges with yellowish median carina; tegmen uniformly
shiny blackish or brown, sometimes with a few dark brown spots in medial area, sometimes dull
brownish or greyish with spots more developed and scattered over whole tegmen; membrane
in dark specimens less transparent than in greyish ones ; wing blackish or inf umate ; outer side
of hind femur blackish with yellowish longitudinal stripe in lower part of medial area, upperside

ACRIDOIDEA OF GALAPAGOS ISLANDS

33

with two indefinite, brown, transverse fasciae, in greyish, specimens the blackish upper part of
medial area greyish and sometimes tending to disappear; hind tibia yellow.

$. Larger than male. Ovipositor, in profile, comparatively short and wide; upper valve
about half as wide as long. In other respects as male.

Measurements

Ratio of length

Length
of body

Length of
pronotum

Length of
tegmen

Length of
hind femur

to width, of
hind femur

Range
Average

30-7-49-1
40-2

6-2-8-2

7'7

32-1-49-0
41-7

17-4-25-5

2I-I

4-4-5-4
4-87

Range
Average

41-4-64-5
54-3

7-7-12-3
10-4

44-9-65-5
55'5

21-5-32-3

27-8

4-5-5-9
5-02

10

12

FIGS. 1-12. Schistocerca melanocera. 1-8, <J: i, prosternal process; 2, end of abdomen,
side view; 3, supra-anal plate; 4, left cercus; 5, phallic complex from above (epiphallus
and most of ectophallic membrane removed) ; 6, the same, lateral view; 7, endophallus,
lateral view; 8, epiphallus (bridge in horizontal position). 9-12, $: 9, end of abdomen
lateral view; 10 sclerotized parts of ovipositor; n, subgenital plate, inner view; 12,
spermatheca.

34

V. M. DIRSH

Geographical distribution: Abingdon, Albemarle, South Seymour, Barrington,
Charles, Chatham, Culpepper, Duncan, Gardner (near Charles), Indefatigable, James,
Jervis, Narborough, Tower.

The main differences between Sch. melanocera and Sch. literosa are in size and
coloration. Their phallic complexes are rather similar. Measurements overlap,
but there is one morphometric character by which it is possible to separate these two
species: the hind femur is relatively more slender and narrower in Sch. melanocera
than in Sch. literosa. This difference can be detected visually, and statistically
it is as follows: Ratio of length to maximum width of hind femur in melanocera is
in males: mean 4-87, standard deviation 0-23, standard error 0-02; and in
females: mean 5-02, standard deviation 0-21, standard error 0-02. The corres-
ponding ratio in literosa is in males mean 4-47, standard deviation i 0-23, standard
error ^ 0-04; and in females: mean 4-55, standard deviation i 0-20, standard
error i 0-04.

GALAPAGOS

o.

^

\ CHARLLS I

IS.

\ .
\ .
\

\ '.

\ \
\:

MAP i . Area of distribution of S. melanocera (dotted line) and S. litorosa (interrupted line) .

ACRIDOIDEA OF GALAPAGOS ISLANDS

35

The difference of means between these two species is: in males 0-41, standard
error ^ 0-04, t 9-37, df 264, P < o-ooi; and in females 0-48, standard error i 0-04,
10-97, df 163, P <o-ooi.

This character places the synonymized Sch. intermedia in Sch. melanocera.

Schistocerca literosa (Walker, 1870)
(PI. 2, Map i, Graph i; Text-figs. 13-24)

Acridium literosum Walker, 1870 : 620. Type <J. ' Galapagos Is.' British Museum (Natural
History), London.

Schistocerca literosa (Walker, 1870) (Scudder, 1893 : 15)

Schistocerca literosa discoidalis Scudder, 1893 : 16 (Hebard, 1920 : 425). LECTOTYPE <$.
' Chatham Is., Wreck Bay.' By present selection. Museum of Comparative Zoology, Har-
vard University, U.S.A. Syntype $. Locality and Museum as for Lectotype.

Schistocerca literosa punctata Scudder, 1893 : 16. Syn. n. Syntype $. ' Hood Is.' Museum of
Comparative Zoology, Harvard University.

Schistocerca literosa hyalina Scudder, 1893 : J 6. Syn. n. Syntype <J. ' Tower Is.' Museum of
Comparative Zoology, Harvard University.

S. literosa can be separated morphometrically (see above) from Sch. melanocera
by the relatively shorter and wider hind femora.

65%T

25% F

155%

35%

GRAPH i . Triangular co-ordinates showing distribution of proportional values of length
of pronotum (P), length of tegmen (T), and length of hind femur (F), expressed as percen-
tages. Chatham Is., A Hood Is., Tower Is. of S. literosa.

36 V. M. DIRSH

The three subspecies of literosa described by Scudder were attributed by him to
three different islands. The differences of characters were tabulated by him, but
when the types and topotypical material were studied, it was clear that not a single
character or any combination could be used to separate the subspecies. These three
forms overlap widely and are present in the population of every island. The popula-
tions from Hood and Tower Islands merge completely in every respect, including
size (Graph I). The Chatham population overlaps widely but not completely with
the populations of Hood and Tower, the tegmina of Chatham specimens being
relatively longer (Graph i), but not so much that it would be advisable to consider
them as a separate race. Probably interchange between these islands occurs
regularly, so that their populations become almost homogeneously mixed.

FIGS. 13-24. Schistocerca literosa. Type. 13-20, <$: 13, prosternal process; 14, end of
abdomen, lateral view; 15, left cercus; 16, supra-anal plate; 17, phallic complex from
above (epiphallus and most of ectophallic membrane removed); 18, the same, lateral
view; 19, endophallus, lateral view; 20, epiphallus (bridge in horizontal position, whole
epiphallus slightly shrunk). 21-24, $: 21, end of abdomen, lateral view; 22, sclerotized
parts of ovipositor; 23, subgenital plate, inner view; 24, spermatheca.

ACRIDOIDEA OF GALAPAGOS ISLANDS

37

Redescription. $. Of medium size. Integument slightly rugose and pitted. Antenna
longer than head and pronotum together. Fastigium of vertex short, obtuse-angular, in middle
with rhomboidal depression; frontal ridge low, with margins parallel and slight depression at
ocellus. Dorsum of pronotum flattened, slightly constricted in prozona and widened in meta-
zona; metazona slightly longer than prozona, its posterior margin rounded; median carina
thin, linear, crossed by three sulci ; lateral lobes of prozona concave in middle. Prosternal process
from narrow, conical, at apex obtuse, to almost cylindrical. Mesosternal interspace twice as
long as its width in middle, slightly constricted and widened in front. Tegmina and wings wide,
exceeding end of abdomen. Opening of tympanal organ almost serai-circular, membrane deeply
set, forming wide angle with body wall. Hind femur comparatively short and wide, average
ratio of length to maximum width 4-47. Hind tibia slightly shorter than femur. Supra-anal
plate elongate-angular, in middle of basal part with longitudinal sulcus, which sometimes
extends to middle of apical half; sides upcurved and elevated. Cercus narrowing towards
apical part, apex oblique, with upper angle protruding. Subgenital plate, in profile, short,
moderately narrow, at apex slightly narrowed.

Phallic comlex : of usual structure for the genus ; apical valves of penis shorter than valves of
cingulum. It differs from melanocera in the shape of the basal valves of the penis and in the
epiphallus, the latter in literosa having more developed and more acute lophi.

Coloration: in general, grey or sometimes brownish. Pronotum uniformly coloured, or mottled
with darkish spots, frequently on upper part of lateral lobe or prozona with a small ochraceous
spot; tegmen covered with dark brownish spots, sometimes poorly defined, sometimes very
sharp; membrane from greyish to transparent colorless; wing hyaline, practically colorless;
hind femur above with three transverse brownish fasciae, sometimes extending to outer and
partly to inner sides; hind tibia yellowish.

. Larger than male. Ovipositor, in profile, comparatively long and narrow; upper valve
about one-third as wide as long.

Measurements

Range
Average
Range
Average

Length of
body

34-6-43-4

39-9

Ratio of length

Length of

Length of

Length of

to width of

pronotum

tegmen

hind femur

hind femur

5-1-6-3

26-4-35-2

13-9-18-9

4-1-4-9

6-1

30-6

16-3

4-47

6-7-7-7

33-7-43-8

17-7-22-6

4-0-5-0

7-5

38-4

20- 1

4-55

Geographical distribution: Chatham, Hood, Gardner (near Hood), Tower, Charles,
Gardner (near Charles).

HALMENUS Scudder, 1893

Four species of this genus have been described. All of them possess strongly
shortened tegmina and wings and cannot fly. They could populate different islands
by means of floating debris brought by currents, but there is also the possibility that
they were able to fly when the islands were populated and afterwards rapidly lost
this ability.

Owing to the loss of the ability to fly, the populations of the islands have been
completely or almost completely isolated. As a result, the genus formed species
which can be differentiated by external morphological characters and partly by the
structure of the phallic complex.

V. M. DIRSH

KEY TO SPECIES

(2) Tegmina overlapping dorsad ; anterior margin of tegmen strongly excurved, apex

narrowed (Text-fig. 25) ....... robustus Scudd.

(i) Tegmina not overlapping dorsad; anterior margin of tegmen straight or only

slightly excurved, apex rounded or slightly attenuate
(6) Male cercus narrow, angular, at apex obtuse or acute. Interocular space

narrower or as wide as antennal scape. Upper outer margin of upper valve

of ovipositor irregularly serrated.
(5) Male cercus narrow, at apex acute or subacute. Tegmina reaching or slightly

exceeding middle of second abdominal tergite . . . cuspidatus Snodgr.

(4) Male cercus narrow, comparatively short, at apex obtuse. Tegmina slightly

exceeding end of first abdominal tergite . . . choristopterus Snodgr.

(3) Male cercus wide, angular, at apex obtuse, almost rounded. Interocular space

wider than antennal scape. Upper margin of upper valve of ovipositor
smooth (Text-fig. 51) eschatus Heb.

26

27

28

FIGS. 25-28. Tegmina of Halmenus. Left, $, right, $. 25, robustus; 26, cuspidatus;
27, choristopterus; 28, eschatus. All drawn to the same scale.

ACRIDOIDEA OF GALAPAGOS ISLANDS 39

In the structure of the phallic complex and of the spermatheca, in the shape of the
lobes of the mesosternum and in the incised male subgenital plate, Halmenus
considerably resembles Schistocerca. It is different from Schistocerca in other respects,
but these similarities suggest strong affinity between the two genera and because
of them Halmenus is here placed in the subfamily Cyrtacanthacridinae.

Type-species: Halmenus robustus Scudder, 1893.

Halmenus robustus Scudder, 1893

(PI. 2, Text-figs. 29-39)

cJ (Type. Redescription). Integument finely rugose and pitted. Antenna filiform, shorter
than head and pronotum together, 24-segmented. Fastigium of vertex longer than wide, slightly
concave in middle, angular, apex obtuse; sides and apex strongly pitted; frontal ridge almost
flat; lateral carinulae obliterated; facial carinulae well developed. Pronotum comparatively
long, slightly constricted at sides in posterior part; dorsum subcylindrical, crossed by 3 deep
sulci ; median carina in prozona almost obliterated, in metazona linear, obtuse ; metazona much
shorter than prozona, its posterior margin obtuse-angular; lateral lobe longer than high, in
prozonal part convex, at posterior sulcus concave, lower margin sinuate. Prosternal process
narrow, conical. Mesosternal interspace about twice as long as wide, slightly widening towards
front; mesosternal lobes long, angular, with angles obtuse. Metasternal interspace strongly
elongate, narrow. Tegmina overlapping dorsad, shortened, reaching fourth abdominal tergite,
venation coarse. Wings shorter than tegmina. Hind femur moderately slender, exceeding
end of abdomen; lower lobes of hind knee rounded. Hind tibia at apical part slightly widened;
external apical spine absent. Arolium large, longer than claw. Last abdominal tergite with a
pair of large, wide projections. Supra-anal plate angular, longer than wide, apex angularly
attenuate; median longitudinal carinula present, bifurcate in basal part; base with a pair of
marginal, lateral projections; transverse fold well pronounced. Cercus acute-angular, com-
pressed from sides, at apex acute. Subgenital plate obtusely conical, at apex excised.

General coloration ochraceous. Lateral lobe of pronotum, in upper part of prozona, with
shiny, dark brown, wide longitudinal stripe, at lower margin of which there is a narrow yellowish
stripe; post-ocular brown stripe similar to pronotal one.

$. Larger than the male. Ovipositor slender; upper outer margin of upper valve of ovi-
positor roughly serrated.

Length: $ 20-0-23-0, $ 26-0-36-2; pronotum $ 4-8-5-7, $ 6-2-7-2; tegmen <$ 5-4-6-I, $ 7-1-
10-9; hind femur $ 12-0-13-4, $ 14-4-17-0 mm. (Measurements based on material studied by the
author.)

Type $. Type locality: ' Conway Bay, Indefatigable Island. Galapagos '.
Type No. 15335. (It must be considered as a Syntype, since Scudder's description
is based on two males and one female.) Museum of Comparative Zoology, Harvard
University, U.S.A.

Geographical distribution : Indefatigable, James.

Halmenus cuspidatus Snodgrass, 1902
(Text-figs. 26, 40-43)

Differs from other species of the genus in the characters indicated in the key.
In female, upper outer margin of upper valve of ovipositor roughly serrated.
Length: $ 19-0-22-8, $ 26-4-29-5; pronotum $ 4-8-5-0, $ 6-4-6-7; tegmen $ 4-8-

4 o

V. M. DIRSH

FIGS. 29-39. Halmenus robustus. 29, phallic complex, from above (epiphallus and most
of ectophallic membrane removed) ; 30, the same, lateral view; 31, endophallus; 32, epi-
phallus. 33, sclerotized parts of ovipositor; 34, spermatheca; 35, end of abdomen;
36, meso- and metasternum; 37, end of abdomen from lateral view; 38, the same, from
above; 39, the same, posterior view, to show excision on end of subgenital plate.

ACRIDOIDEA OF GALAPAGOS ISLANDS 41

5-0, $ 5-2-6-1; hind femur <$ 12-5-13-6, $ 14-8-16-0 mm. (Measurements based on
material studied by the author.)

Type $. Type locality: Albemarle Is., Iguana Cove. Type lost.
Geographical distribution : Albemarle.

Apd ;

--/ Rm

Ects

40

FIGS. 40-43. Halmenus cuspidatus. Phallic complex. 40, from above (epiphallus and
most of ectophallic membrane removed); 41, the same, lateral view; 42, endophallus;
43, epiphallus.

Halmenus choristopterus Snodgrass, 1902
(Text-fig. 27)

Differs from the other species of the genus in the characters indicated in the key.
Tegmina in both sexes shorter than in cuspidatus. In female, upper outer margin
of upper valve of ovipositor roughtly serrated.

Length: <J 22-3, $ 25-1-30-0; pronotum <$ 5-0, $ 6-4-6-5; tegmen 3-6, ? 5-9-6-1;
hind femur ^ 13-4, $ 14-7-17-2 mm. (Measurements based on material studied
by the author) .

Type $. Type locality: Charles Is. Type lost.
Geographical distribution : Known only from Charles Is.

ENTOM. 23, 2 5

42 V. M. DIRSH

Halmenus eschatus Hebard, 1920
(PI. 3, Text-figs. 44-51)

This is the most divergent species of the genus. Characters additional to those
mentioned in the key are: tegmina in both sexes micropterous, being fully lateral
and more strongly reduced than in other species; and interocular space much wider
than in others.

Measurements: length: <$ 20-8-23-5, $ 27-0-32-8; pronotum <$ 5-2-5-9, $ 7-0-7-9;
tegmen $ 3-5-4-0, $ 5-0-6-2; hind femur $ 10-4-11-3, $ 13-2-14-5 mm.

Type $. Type locality : ' Wenman Is.' Museum of California, Academy of Sciences.
(Type No. 717.)

Geographical distribution: Known from type locality only.

FIGS. 44-51. Halmenus eschatus. Paratypes. 44, end of abdomen from above; 45, the
same, lateral view; 46, the same, posterior view, to show excision of apex of subgenital
plate; 47, phallic complex from above (epiphallus and most of ectophallic membrane
removed); 48, the same, lateral view; 49, endophallus; 50, epiphallus; 51, $, end of
abdomen, lateral view.

ACRIDOIDEA OF GALAPAGOS ISLANDS

43

Desmopleura concinna Scudder, 1893
(PL 4, Text-figs. 52-59)

This genus belongs to subfamily Catantopinae of Acrididae, and probably to the
group Podismini judging from the structure of the phallic complex.

No genus examined from the American continent is closely allied to Desmopleura.
It may perhaps be related to the genus Dichroplus, but if so the relationship is
rather remote.

Since only the male type is known, there is no possibility of studying it in detail
and further material is needed.

$ (Type. Redescription). Small. Integument slightly pitted. Antenna filiform, slightly
shorter than head and pronotum together. Head obtuse, conical. Fastigium of vertex short
obtuse-angular, sloping fowards, in middle with deep sulcus bordered by a pair of high, strong
lateral carinulae which extend beyond base of vertex ; linear median carinula running from middle
of fastigium along whole head; frons oblique, ftonral ridge narrow, sulcate, lateral carinulae
obtuse. Pronotum subcylindrical, dorsum flattened, crossed by 4 fine sulci; median carina
linear, crossed by posterior and preposterior sulci; lateral carinae absent; anterior margin
straight, posterior broadly obtuse-angular; lateral lobe longer than high, margin excurved in
middle, transverse sulci deep and comparatively wide. Prosternal process tongue-shaped,
inclined backwards and touching mesosternum. Mesosternal interspace longer than wide;

Ejd

EJS-V--W-4-

// ^x\

/ 59 \

V \

Ejd

Gpr Ejs

FIGS. 5259. Desmopleura concinna. Type. 52, end of abdomen, from above; 53, the
same, lateral view; 54, cercus, from above; 55, the same, lateral view; 56, phallic
complex from above (epiphallus and most of ectophallic membrane removed) ; 57, the
same, lateral view; 58, endophallus; 59, epiphallus.

ENTOM. 23, 2. 5

44 V. M. DIRSH

lateral lobe longer than wide, with rounded angle. Metasternal interspace elongate, closed.
Tegmina and wings fully developed. Tegmen exceeding end of hind femur, rather narrow,
reticulation sparse. Wing comparatively narrow, as long as tegmen. Tympanum well
developed, large. Anterior and middle femora slightly inflated. Hind femur short and moder-
ately slender. Lower basal lobe of hind knee rounded. External apical spine of hind tibia
absent. Arolium large. Last abdominal tergite with a pair of small, narrow projections in
middle. Supra-anal plate short, angular, at apex obtuse. Cercus wide in basal part, narrow-
ing towards apex, which is strongly incurved; apex obtuse. Subgenital plate in profile elongate,
narrow conical.

Phallic complex : Similar to that of the group Podismini : apical valve and valve of cingulum
of penis covered by sheath; apical valves long, slender, at apices acute; valves of cingulum in
profile slender, above fused, shorter than apical valves of penis; basal valves of penis large,
strongly expanded; zygoma very large; apodemes slender, forming U-shaped structure. Epi-
phallus wide-bridged; ancorae small, acute, inclined inwards; lophi long and narrow, forming
narrow lobes.

General coloration buff ; from posterior margin of eye and extending along upper part of lateral
lobe of pronotum, a wide blackish stripe; from apex of fastigium of vertex to posterior margin
of pronotum, a dark brown stripe; carinula on head and median carina of pronotum lighter
coloured than stripe; veins and veinlets of tegmen and wing brownish, membrane colorless.
(Other parts of type specimen discolored due to previous preservation in fluid.)

Measurements: Length: 13-5, pronotum 3-2; tegmen n-o; hind femur 9-0 mm.

Type <. Type locality: 'James Island'. United States National Museum,
Washington.

Closteridia bauri Scudder, 1893
(PL 5, Text-figs. 60-66)

cJ. Of medium size, robust. Integument strongly rugose and granulose. Antenna filiform,
longer than head and pronotum together, ao-segmented. Head subconical; fastigium of vertex
acute-angular, as long as wide, lateral margins raised, median carinula well developed, fastigial
foveolae narrow-rhomboidal, deep, with prominent edges; frons inclined backwards, frontal
ridge narrow, sulcate, lateral carinulae high, almost parallel, slightly diverging basally; facial
carinulae high, slightly excurved; weak occipital carinula present. Pronotum subcylindrical,
widened backwards, dorsum indistinctly separated from lateral lobes; median carina irregularly
linear, in prozona slightly raised; lateral carinulae lyrate, limiting narrow elevation of dorsum;
transverse sulci indistinct; posterior margin of metazona corrugated; posterior angle of lateral
lobe roundly protruding backwards. Prosternal process absent. Mesosternal interspace
twice as wide as long. Metasternal interspace wider than long. Tegmina vestigial; wings and
tympanum absent. Metanotum and first abdominal tergite with rough median carinula.
Hind femur short, stout, reaching end of abdomen. Lobes of hind knee with apices rounded.
Hind tibia slightly shorter than femur; external apical spine absent; spurs short, of equal length;
arolium slightly longer than half length of claws. Last abdominal tergite slightly and regularly
incurved. Supra-anal plate slightly longer than wide, angular, at apex obtuse, slightly incurved,
in basal part with small median concavity, transverse ridge in middle well developed. Cercus
short, conical. Subgenital plate short, subconical.

Phallic complex: of Acridinae type. Apical valve of penis short, much shorter than valves
of cingulum; basal valves large and strongly expanded; apodemes relatively small, forming
wide, arch-like structure. Epiphallus robust, ancorae large, inclined inwards ; lophi very large,
irregularly bilobate.

General coloration rust-brown, mottled with dark brown spots; head above and dorsum of
pronotum lighter, dirty ochraceous; outer and inner sides of hind femur with two transverse,
brown fasciae ; hind tibia dirty ochraceous.

ACRIDOIDEA OF GALAPAGOS ISLANDS

45

4-3, $ 4-5; tegmen 0-3, $ absent (?); hind femur

$ (Type). See original description.
Length: $ 20-0, $ 22-0; pronotum
u-o, ^ 13-5 mm.

Type $ (lost). Type locality ' Wreck Bay, Chatham Is.'

<$. Described by Hebard (1920) and redescribed above. Cowley Mountain,
Albemarle Is. Museum of the California Academy of Sciences.

Only two specimens of this remarkable genus and species are known; and since
the female type is lost, it is not possible to verify that both specimens belong to the
same species. They are practically wingless and were collected on different and
widely separated islands.

This genus is of the subfamily Acridinae of Acrididae (sensu Dirsh, 1961). No
close relatives of it are known from America and its phylogenetic relationship is
obscure.

FIGS. 60-66. Closteridia bauri. 60, phallic complex from above (epiphallus and most of
ectophallic membrane removed) ; 61, the same, lateral view; 62, endophallus; 63, epi-
phallus; 64, end of abdomen, from above; 65, the same, lateral view; 66, meso- and
metasternum.

46 V. M. DIRSH

SPHINGONOTUS Fieber, 1852

It was rather surprising to find the genus Sphingonotus represented on the remote
Galapagos Islands. This presents a puzzling zoogeographical problem, even though
Sphingonotus is the most widely distributed genus of all Acridoidea. It occurs over
the whole of Europe, Africa, most of Asia (including Japan), Australia, Jamaica,
Ecuador and the Galapagos Islands.

The most numerous specific complex of the genus occurs in Central and South-west
Asia and in the palaearctic part of Africa.

It is a peculiar fact that the species from Galapagos (Text-figs. 67-70), Jamaica
(Text-figs. 71-74), and the type-species S. caerulans (Text-figs. 75-79) from Sweden
have remarkably similar phallic complexes. (The one known specimen from Ecua-
dor, actually from Puna Island, is a female, but externally it is very similar to the
species from Galapagos.)

This indicates a relatively recent separation of these species and a relatively recent
penetration by them to America and the Galapagos Islands. We have no indication
of how this occurred. The distances between the Palaearctic Region, Jamaica,
Ecuador and Galapagos are enormous, and Sphingonotus has not so far been found
in intervening countries. At present it is only possible to affirm a close affinity
between the species on the basis of morphological characters, leaving zoogeographical
considerations until more data become available.

Stal in 1860 described Oedipoda fuscoirrorata (at present Sphingonotus fuscoir-
roratus) from two females from ' Galapagos ' and one female from Puna Island,
about two miles from the coast of Ecuador. Snodgrass (1902) , who considered the
specimen from Puna Island different from the Galapagos specimens, restricted the
name fuscoirrorata Stal to the specimen from Puna Island and gave new names,
trinesiotis and tetranesiotis, to two alleged species from Galapagos, which include
Stal's specimens, on the grounds that Stal did not fix the type locality. Here
Snodgrass' action is considered incorrect and not valid, on the grounds that Stal
had two females from Galapagos and one female from Puna Island and in his original
description mentioned their localities as ' Galapagos Is. and Puna Is/, not vice versa
as stated by Snodgrass. For this reason Stal's name Oedipoda fuscoirrorata Stal,
1860 is accepted here as the original name for the Galapagos species, but for the
specimen from Puna Island the specific name punensis nom. n. is proposed.

In the same paper Snodgrass (1902), who assigned trinesiotis and tetranesiotus to
Sphingonotus, described three subspecies of the former and four of the latter. Hebard
(1920) synonymized all of them, including the specimen from Puna Island, under
Stal's original name fuscoirrorata. In the light of the present study Hebard's
synonymy is correct, except for the specimen from Puna Island whose position is
doubtful as only one specimen is known. From the present author's point of view
it seems preferable to retain the specimen from Puna Island as a separate species
of Sphingonotus. By all external characters, this specimen is very similar to the
Galapagos species, except that the metazona of the pronotum (PI. 7) is relatively
shorter and its posterior margin is widely obtuse-angular, almost rounded, while
in the Galapagos species it is acute-angular, and obtuse only at the apex (PI. 6).

ACRIDOIDEA OF GALAPAGOS ISLANDS

47

A study of the series of the species and subspecies of Snodgrass from the topo-
typical localities shows that they cannot be separated by external morphological
characters or by the phallic complex. When studied morphometricaHy they
display some tendency towards forming races on the different islands. When three
morphometric characters (length of pronotum P, length of tegmen T and length
of hind femur F) are studied and plotted as triangular co-ordinates (Graph 2)
these various forms show such a tendency, but they overlap so widely that to consider
them as races would be stretching the point too far. Probably, in spite of being
small and not such good fliers as Schistocerca, they can still fly from island to island,
levelling the morphological and morphometric features of the population.

60% T

30%F

T50/o

P

10/0

F40%

GRAPH 2. Triangular co-ordinates showing distribution of proportional values of length
of pronotum (P), length of tegmen (T) and length of hind femur (F), expressed as per-
centages. Hood Is., A Indefatigable Is., Charles Is., x Narborough Is.,
V Albemarle Is., * Chatham Is., Abingdon Is., ^ James I.

Sphingonotus fuscoirroratus (Stal, 1860)
(PI. 6, Text-figs. 67-70)

Oedipoda fuscoirrorata Stal, 1860 : 345 (partim). LECTOTYPE ? ' Galapagos Is.' by present

selection. Stockholm Museum.

Sphingonotus trinesiotis Snodgrass, 1902 (Hebard, 1920 : 321).
Sphingonotus trinesiotis chathamensis Snodgrass, 1902 : 437 (Hebard, 1920 : 321). Type <$,

lost.
Sphingonotus trinesiotis indefatigabilensis Snodgrass, 1902 : 437 (Hebard, 1920 : 321). Type <$,

lost.

V. M. DIRSH

Sphingonotus trinesiotis albemarlensis Snodgrass, 1902 : 437 (Hebard, 1920 : 321). Type $,

lost.

Sphingonotus tetranesiotis Snodgrass, 1902 : 437 (Hebard, 1920 : 321).
Sphingonotus tetranesiotis charlesensis Snodgrass, 1902 : 437 (Hebard, 1920 : 321). Type <J,

lost.
Sphingonotus tetranesiotis barringtonensis Snodgrass, 1902 : 437 (Hebard, 1920 : 321). Type <,

lost.

Sphingonotus tetranesiotis hoodensis Snodgrass, 1902 : 438 (Hebard, 1920 : 321). Type <J, lost.
Sphingonotus tetranesiotis indefatigabilensis Snodgrass, 1902 : 438 (Hebard, 1920 : 321). Type

<J, lost.

The female specimen of Oedipoda fuscoirrorata Stal, 1860 which was quoted in
Sjostedt's 1932 paper as the type is selected here and described below as lectotype.
All three of StaTs original specimens were studied.

Redescription. 9 Lectotype. Integument rugose and granulose. Antenna shorter then
head and pronotum together. Fastigium of vertex sloping forwards, concave and widening in
front; lateral and median carinulae well developed; frontal ridge slightly widening at ocellus,
constricted below ocellus and undulating below. Pronotum rugose and granulose, dorsum
crossed by 3 sulci ; median carina slightly elevated in front of anterior sulcus, obliterated between

B P

cv

-Apd

-Ap

Ects

68

-Ap

Sps

FIGS. 67-70. Sphingonotus fuscoirroratus, from Charles Is. 67, phallic complex, from
above (epiphallus and most of ectophallic membrane removed) ; 68, the same, lateral
view ; 69, endophallus ; 70, epiphallus.

ACRIDOIDEA OF GALAPAGOS ISLANDS

anterior and posterior sulci and sharply linear in metazona ; in front of posterior sulcus a concavity
with a pair of small elevations posteriorly; metazona about half as long again as prozona, its
posterior end acute-angular, apex obtuse, sides undulated ; anterior angle of lateral lobe of
pronotum obtuse-angular, posterior angle with acute-angular projection. Tegmina exceeding
end of abdomen; intercalary vein of medial area almost parallel to medial vein. Mesosternal
interspace short, twice as wide as long. Subgenital plate at apex truncate. Ovipositor short,
valves curved at apices; lower valve with angular, lateral, external projection.

General coloration brownish; tegmen with 2 dark brown transverse fasciae and with small
spots in apical part; wings hyaline, slightly infumate.

<J. Much smaller than female. Antenna longer than head and pronotum together. Fasti-
gium of vertex more sloping forwards, narrower, more concave. Last abdominal tergite with a
pair of small, rounded projections; supra-anal plate short, angular. Cercus narrow-conical,
at apex obtuse. Subgenital plate short, obtusely subconical.

Phallic complex of usual type for the genus (see Text-figs. 67-70).

General coloration as in female; wing more infumate.

Measurements (mm.

Range
Average

Range
Average

Length of
body

10-9-17-7
13-8

Length of
pronotum

2-3-3-5
2-9

3-2-4-6
4-0

Length of
tegmen

9-3-14-8
12 -6

i 3 8-2 i 2

17-3

Length of
hind femur

6-4-10-2
7-9

8-4-12 -9
10-7

Apd

- Ap

Ects

-Ap

Ejd

73

Ejs

Sps

FIGS. 71-74. Sphingonotus jamaicensis (Saussure, 1884). 71, phallic complex, from
above (epiphallus and most of ectophallic membrane removed) ; 72, the same, lateral
view; 73, endophallus; 74, epiphallus.

V. M. DIRSH

Bp

E JS V- \

Ap

Ects

-Ap

Sps

77

FIGS. 75-79. Sphingonotus caerulans (Linnaeus, 1767). 75, Phallic complex from above
(epiphallus and most of ectophallic membrane removed); 76, the same, lateral view;
77, endophallus; 78, epiphallus; 79, spermatheca.

Lectotype $. Lectotype locality ' Galapagos Is.' Stockholm Museum.

Geographical distribution: Abingdon, Albemarle, Barrington, Charles, Chatham,
Gardner (near Hood), Hood, Indefatigable, James, Jervis, Narborough, Seymour
(both).

REFERENCES

BOWMAN, R. J. edit. 1966. The Galapagos. Proceedings of the Symposia of the Galapagos

International Scientific Project. Univ. of California Press : 1-318.
CAUDELL, A. N. 1932. Insects of the Order Orthoptera of the Pinchot Expedition of 1929.

No. 2921, Proc. U.S. natn. Mus. 80, art. 21 : 1-7.
Cox, A. 1966. Continental Drift in the Southern Hemisphere. The Galapagos. Proc. of the

Sympos. of the Galap. Intern. Scient. Project. Univ. of California Press.
Cox, A. & DALRYMPLE, G. B. 1966. Palaeomagnetism and Potassium- Argon ages of some

volcanic rocks from the Galapagos Islands. Nature, Lond. 209, No. 5025 : 776-7.
EiBL-EiBESFELDT, J. 1960. Galapagos. MacGibbon & Kee, London.
GRESSITT, J. L. 1964. Pacific Basin Biogeography. Bishop Museum Press, Honululu.
HEBARD, M. 1920. Dermaptera & Orthoptera. Expedition of the California Academy of

Sciences to the Galapagos Islands, 1905-1906. 17. Proc. Calif. Acad. Sci. (4), 2 : 311-

346, text-figs. I-II.
HEBARD, M. 1934. Orthoptera of the Galapagos Islands. Meddr. zool. Mus., Oslo Nr. 41.

(Saertryk av Nyt Magazin for Naturvidenskaberne 74).

ACRIDOIDEA OF GALAPAGOS ISLANDS 51

LINSLEY, E. G. & USINGER, R. L. 1966. Insects of the Galapagos Islands. Proc. Calif. Acad.

Sci. 33, No. 7 : 113-196.
ORR, R. T. 1966. Evolutionary Aspects of the Mammalian Fauna of the Galapagos. The

Galapagos. Proc. of the Sympos. of the Galap. Intern. Scien. Project. Univ. of California

Press.
SCUDDER, S. H. 1893. Reports on the dredging operations off the West Coast of Central

America to the Galapagos, to the West Coast of Mexico, and in the Gulf of California, in

charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer 'Albatross ',

during 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. XII. The Orthop-

tera of the Galapagos Islands. Bull. Mus. comp. Zool. Harv. 25, no. i : 1-25, pis. 2.
SJOSTEDT, Y. 1932. Orthopterentypen im Naturhistorischen Reichsmuseum zu Stockholm.

2. Acrididae. Ark. Zool. 24 A No. i : 1-89, 20 pis.
SNODGRASS, R. E. 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898-

1899. VIII. Entomological Results (7). Schistocerca, Sphingonoius and Halmenus.

Proc. Wash. Acad. Sci. 4 : 411-454, 2 pi.
STAL, C. 1861. Orthoptera species novas descripsit C. Stal. In Kongliga Svenska Fregatten

Eugenies Resaomkring jorden under befal af C. A. Virgin, aren 1851-1853. 2, Zoo.ogi.

Insecta, 10 : 299-350. P. A. Norstedt, Stockholm.
VAN DENBURGH, J. 1914. The gigantic tortoises of the Galapagos Archipelago. Proc. Calif.

Acad. Sci. (4), 2 (i) : 203-374.
WILLIAMS, H. 1966. Geology of the Galapagos Islands. The Galapagos. Proc. of the Sympos.

of the Galap. Intern. Scient. Project. 65-70. Univ. of California Press.
WOOSTER, W. S. & HEDGPETH, J. W. 1966. The Oceanographic Setting of the Galapagos.

The Galapagos. Proc. of the Sympos. of the Galap. Intern. Scient. Project. Univ. of

California Press.

PLATE i

Schistocerca melanocera. Type. Dorsal and lateral views, with original Stal labels.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 2

PLATE i

,

ENTOM. 23, 2.

PLATE 2

Above, Schistocerca literosa. Type. Below, Halmenus robustus. Syntype. Lateral view
(black spot in middle of hind femur is hole left by pin) ; and dorsal view (position of tegmina
slightly disarranged, owing to previous preservation in liquid; normally they overlap).

Bull. BY. Mus. nat. Hist. (Ent.) 23, 2

PLATE 2

PLATE 3

Halmenus eschatus. $ and $ paratypes, dorsal and lateral views.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 2

PLATE 3

PLATE 4
Desmopleura concinna. Type and the labels attached to it.

Bull. BY. Mus. nat. Hist. (Ent.) 23, 2

PLATE 4

No,
U.S.K.M.

Acc.21,69?

u.

. c.

PLATE 5

Closteridia bauri. <$ (the specimen described by Hebard, 1920).

Bull. Br. Mus. nat. Hist. (Ent.) 23, 2

PLATE 5

PLATE 6
Sphingonotus fuscoirroratus. Lectotype. Lateral view and from above.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 2

PLATE 6

PLATE 7
Sphingonotus punensis. Type. Lateral view and from above.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 2

PLATE 7

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. 5.

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp. 284 ; 348 Text-figures. August, 1965. 6.

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ;
18 plates, 270 Text-figures. August, 1965. 4 45.

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October,

1965- 35s.

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156

475 Text-figures. November, 1965. 2 155.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae.
Pp. 129 ; 328 Text-figures. May, 1966. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera : Coccoidea). Pp. 168 ; 43 Text-figures. February, 1967.

33s.

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146
Text-figures, 9 maps. February, 1967. 3 T.OS.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera : Rhopalocera). Pp. 509. August 1967. 8 los.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa-
locera). Pp. 322; 233 Text-figures. Coloured frontispiece. September 1967. 8.

11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp.
184; 82 Text-figures. May, 1968. 4.

12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with an
Account of their World Distribution (Lepidoptera : Drepanidae). Pp. 151:
293 Text-figures, 14 plates. November, 1968. 5.

PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, BARTHOLOMEW PRESS, DORKING

DIPTERA FROM NEPAL

SPHAEROCERIDAE

J. C. DEEMING

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. 3

LONDON : 1969

DIPTERA FROM NEPAL

SPHAEROCERIDAE

BY

J. C. DEEMING.,

Institute for Agricultural Research, Samaru, P.M.B. 1044 Zaria, Nigeria

P P- 53-74 ; 32 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. 3

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in Jive series corresponding to the Departments
of the Museum, and an Historical series.

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

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

This paper is Vol. 23, No. 3 of the 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) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 28 January, 1969 Price Ten Shillings

DIPTERA FROM NEPAL
SPHAEROCERIDAE

By J. C. DEEMING

CONTENTS

PAGE

SYNOPSIS ........... 55

INTRODUCTION ........... 55

TAXONOMY ........... 56

REFERENCES ........... 73

INDEX ............ 74

SYNOPSIS

The Sphaeroceridae collected on the 1954 an d 1961-62 British Museum Expeditions to Nepal
are systematically treated along with specimens collected in neighbouring countries. Of the
thirty-eight species collected, nine new species are described.

INTRODUCTION

THIS paper is based mainly on material collected by Mr. R. L. Coe, entomologist on
the 1961-62 British Museum (Natural History) Expedition to Eastern Nepal, from
localities in the Taplejung District and Arun Valley. Also included are a few
specimens collected by Mr. J. Quinlan in Western Nepal during 1954, by Mr. E.
Brunetti at Darjeeling during 1913 and single specimens from Assam collected by
Mr. R. Senior-White and Mr. C. B. Antram. All type material is in the British
Museum (Natural History).

Very little has been written on the Sphaeroceridae of the Himalayas. Brunetti
(1913 : 173-4) described Limosina magna from Kalek on the north-east frontier of
India and Limosina subtinctipennis (now Leptocera (Rachispodd) subtinctipennis
(Brun.)) from Assam. He later (1924 : 103) described Limosina notatipennis and
Limosina tenebrosa and recorded Limosina ornata de Meijere ( = Leptocera (Poecilo-
somella) borboroides (Walker)) from the Siju Cave, Garo Hills, Assam. Limosina
Macquart sensu Brunetti must be regarded as synonymous with Leptocera Olivier
sensu lato. The original descriptions of Limosina magna, notatipennis and tenebrosa
are inadequate for the placing of these species in subgenera, and I have unfortunately
been unable to examine the types in the Indian Museum, Calcutta. Richards
(19626) recorded nineteen species of Sphaeroceridae from Afghanistan, eighteen of
which are found in Europe and one possibly new. Hackman (1965 : 37-38) des-
cribed Copromyza (Crumomyia] deemingi from Sikkim. The mediterranean species
Leptocera (Coproica] digitata (Duda) has been recorded from Turkestan, and I have
seen specimens from Kirgiziya and Kazakhastan, but it is not represented in the
Nepalese material.

My thanks are due to the late Mr. R. L. Coe for making his material available to
me, to Prof. O. W. Richards of Imperial College of Science and Technology, London

ENTOM. 23 3 7

56 J. C. DEEMING

and Messrs. H. Oldroyd, K. G. V. Smith and A. C. Pont of the British Museum
(Natural History) (BMNH) for their kind help and advice, and to the Director,
Institute for Agricultural Research, Samaru, Nigeria for allowing me the facilities
to complete this paper.

TAXONOMY

Sphaerocera (Lotobia) pallidiventris (Meigen)

Borborus pallidiventris Meigen, 1830 : 204.

W. NEPAL: Ulleri, 6,000-7,000 ft., I #, 19^.1954 (J. Quinlari).

Widespread in Europe and Africa. Richards (19626 : 177) recorded a specimen
collected at an altitude of 1,320 m in Afghanistan. There are specimens in BMNH
from Kirgiziya, Kazakhastan and Assam.

Copromyza (Borborillus) marginatis (Adams)

Borborus marginatis Adams, 1905 : 198.

E. NEPAL: Taplejung Distr., Dobhan, c. 3,500 ft., shady places on shrubby slope
above R. Tamur, i <, 8 ?, 21-27.1.1962 (R. L. Coe).

Recorded from the Atlantic Islands, Africa, Madagascar, India, Philippines and
Palau Is.

Leptocera (Leptocera) curvinervis (Stenhammar)
Limosina curvinervis Stenhammar, 1853 : 406.

A very long series of both sexes was collected by Mr. J. Quinlan at Ulleri in Western
Nepal and a similar series by Mr. R. L. Coe from various localities in Eastern Nepal.
It would be pointless to list all the locality-data as the species is so common and
widespread, having been recorded from Europe, the Atlantic Islands, Africa,
Madagascar, Seychelles, Ceylon, India, Formosa, Java, New Guinea, Micronesia
and Samoa. I have seen a single specimen from Australia and a long series from
Sikkim.

Leptocera (L.) nigrolimbata (Duda)

Paracollinella nigrolimbata Duda, 1925 : 60.

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
plants by damp cliff in deep river gorge, i $, 1-^.1962; Taplejung Distr., edge of
mixed forest above Sangu, c. 6,500 ft., i $, I7.x.-i.xi.i96i; Taplejung Distr.,
damp evergreen oak forest above Sangu, c. 9,200 ft., i ?, 2-26 . xi . 1961 ; Taplejung
Distr., Sangu, c. 6,200 ft., mixed vegetation by stream in gully, i ^, 3 ?, xi.i96i-
1.1962; Taplejung Distr., Sangu, c. 6,200 ft., mixed vegetation in deep gully, i ?,
2. i. -13. 11.1962; Taplejung Distr., Dobhan, E. bank of R. Tamur, c. 3,500 ft., mixed
vegetation by stream in deep gully, i $, 1-^.1962 (R. L. Coe}.

INDIA: Darjeeling, 5 $, 4 $, 13-18. ix. 1913 (E. Brunetti}.

Described from Formosa.

DIPTERA FROM NEPAL 57

Leptocera (L.) paranigrolimbata (Duda)

Paracollinella paranigrolimbata Duda, 1925 : 61.

INDIA: Darjeeling, <$, 13-18. ix. 1913 (E. Brunetti).

Described from Formosa. There are two males in the BMNH from Assam.

Leptocera (L.) ? koningsbergeri (Duda)

Paracollinella koningsbergeri Duda, 1925 : 49.

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5, 200 ft., mixed
shrubs in deep gorge, I <J, x-xi.i96i (R. L. Coe).

This specimen agrees well with the original description, but has a faint infuscate
band across the middle of the wing. Described from Sumatra and W. Java. The
" Buitenzon " given by Duda as a type locality is a misspelling of Buitenzorg, now
named Bogor.

Leptocera (Rachispodd) subtinctipennis (Brunetti)

Limosina subtinctipennis Brunetti, 1913 : 174.

E. NEPAL: Taplejung Distr., Sangu, c. 6, 200 ft., mixed vegetation by stream in
gully, I c?, xi . 196 i-i . 1962 ; same locality, evergreen scrub, I $, 5-13 . x . 1961 ; Arun
Valley, below Tumlingtar, River Sabhaya, west shore, c. 1800 ft., dead leaves lying
on sandy shore, i <j>, 22.xii.i96i (R. L. Coe).

A common species distributed across the tropics of the Old World from West
Africa to the Caroline and Solomon Islands.

Leptocera (Rachispodd) sp. near divergens (Duda)

E. NEPAL: Taplejung Distr., damp evergreen oak forest above Sangu, c. 9,200 ft.,
i $, 2-26. xi. 1961 (R. L. Coe).

The single teneral specimen runs to Collinella divergens Duda in Duda's 1925 key,
but differs from his description for that species in having a very weakly developed
interantennal keel, a strongly projecting mouth margin, and the scapular present
only as a very weak hair.

Leptocera (Opacifrons) pseudimpudica sp. n.

(Text-figs, i, 2)

This species agrees almost perfectly with Duda's original description of Opacifrons
impudica. However, as that species was described from the Andes and has not been
recorded from the Pacific, I prefer to regard the two species as separate rather than
risk a misidentification.

<J. A small dull black species, heavily whitish dusted, especially on the face and jowls; legs
dirty yellow, but femora, tibiae and fore tarsus noticeably infuscated; halteres dirty yellow;
wings whitish hyaline with light brown veins. Head deeper than long; jowls one-quarter of

ENTOM. 23, 3 7

58 J. C. DEEMING

height of eye ; face moderately produced between the antennae, which between their bases form
an angle of 100; arista long-haired, nearly four times as long as remainder of antenna, of which
the third segment is noticeably long-haired ; four pairs of fine interfrontal bristles ; postvertical
bristle as long as ocellar ; all head-bristles long ; frontal triangle less heavily dusted than remainder
of frons. Dorsum of thorax less heavily dusted than head, with two pairs of strong dorsocentral
bristles and six rows of interdorsocentral setulae between the more anterior pair; a pair of
prescutellar acrostical hairs enlarged; scutellum long, with the usual two pairs of bristles, the
shorter anterior pair as long as scutellum and apical pair as long as dorsum of thorax; fore
femur with a row of weak posterodorsal and posteroventral hairs ; fore tarsus long, the seg-
ments becoming progressively more flattened towards the apex; mid tibia with a short antero-
dorsal bristle at 0-7 of its length, a medium one at 0-23 and a long one at 0-82, a medium
posterodorsal at 0-7, no apical ventral, and the apical two-thirds of its ventral surface set with
minute stout bristles ; ventral bristle of mid basitarsus longer than the basitarsus is wide ; hind
leg normal. Wing (Text -fig. i) longer than abdomen; second costal sector shorter than third;
costa overpassing the slightly sinuate third vein ; fold of fourth vein extending to wing-margin ;
fifth vein weakly produced beyond discal cell; alula narrow. Abdomen with only weak pale
hairs; pregenital sternite (Text-fig. 2) with two widely-separated thumb-like projections and an
inwardly-directed hook-like projection on the posterior margin; claspers strap-like, weakly
infuscated; posterior gonapophysis long, pointed and downwardly-directed. Length 1-6 mm.

$ resembling <$, but mid tibia lacking the numerous short stout bristles ventrally; last tergite
strongly shining; cercus with pale upwardly-directed sharp apical tooth.

Holotype <$. E. NEPAL: Taplejung Distr., river banks below Tamrang Bridge,
c. 5,500 ft., x-xi.i96i (R. L. Coe).

Paratypes. 2 <$, 4?, same data; Taplejung Distr., above Sangu, c. 6,500 ft.,
evergreen scrub, 2$, 5-13. x. 1961; Taplejung Distr., Sangu, c. 6,200 ft., mixed
vegetation by stream in gully, 2 $, ix-x.i96i; Taplejung Distr., edge of mixed
forest above Sangu, c. 6,500 ft., i $, I7.x.-i.xi.i96i (R. L. Coe).

FIGS. 1-2. Leptocera (Opacifrons) pseudimpudica sp. n. i, right wing; 2,
cJ pregenital sternite.

KEY TO MALES OF SPECIES OF Leptocera (Poecilosomella) RECORDED
FROM THE HIMALAYAS

1 Hairs on arista very long ; costal hairs long and fine ; second costal sector short ; wing

brownish with pale markings on the crossveins

borboroides (Walker) (widespread in Asia)

- Hairs on arista short ; costal hairs minute, but if long on first sector, then robust ; wing

pattern variable ............ 2

2 Second vein bearing a distinct preapical stump-vein ...... 3

- Second vein lacking a stump-vein ......... 4

DIPTERA FROM NEPAL

59

3 Long anterior bristle on mid tibia equidistant between the two long anterodorsals

annulitibia sp. n. (Nepal)
Long anterior bristle on mid tibia situated much nearer to the more basal of the two

long anterodorsals than to the more apical . . nepalensis sp. n. (Nepal, India)

4 Apical ventral bristle on mid tibia very long, half as long as basitarsus ... 5

- Apical ventral bristle on mid tibia shorter, sometimes absent .... 6

5 Genitalia as in Text-fig. 19 ...... brunettii sp. n. (India)

Genitalia as in Text-fig. 20 .... sp. near brunettii Deeming (India)

6 Fore tarsus predominantly vivid white .... aciculata sp. n. (Nepal)

- No segment of fore tarsus white, though some may be dirty yellow ... 7

7 Ventral surface of mid tibia long-haired ........ 8

- Ventral surface of mid tibia short-haired ........ 9

8 Reddish brown species ; apical ventral bristle of mid tibia present though small

punctipennis (Wiedemann) (widespread through Asia to Polynesia)

- Blackish brown species; apical ventral bristle of mid tibia absent

hitnalayensis sp. n. (Nepal)

9 Second and third costal sectors equal, or nearly so various Duda (widespread in Asia)

- Second costal sector much longer than the third

longinervis Duda (India, Burma, Nepal, China and Formosa)

Leptocera (Poecilosomella) punctipennis (Wiedemann)

Borborus punctipennis Wiedemann, 1824 : 559.

E. NEPAL: Arun Valley, below Tumlingtar, River Sabhaya, west shore, c. 1,800 ft.,
human excreta in sandy place, 3 <?, I ?, 9-22 . xii . 1961 (R. L. Coe).

A very common species, recorded from the Belgian Congo, Asia, Micronesia and
Polynesia. There are specimens from Queensland, Australia, in BMNH. Brunetti
collected very long series of this species at Darjeeling, Lucknow and Calcutta.

Leptocera (Poecilosomella) varians Duda

(Text-figs. 8, 14, 21)
Poecilosomella varians Duda, 1925 : 99.

E. NEPAL: Arun Valley, east shore of R. Arun below Tumlingtar, c. 1,800 ft.,
evergreen shrubs bordering dry stream beds, I <$, 14-23 . xii . 1961 ; Arun Valley,
below Tumlingtar, River Sabhaya, west shore, c. 1,800 ft., human excreta in sandy
place, i $, 9-22. xii. 1961 (R. L. Coe).

Described from Singapore and Formosa. There are specimens in BMNH from
India, Ceylon and Celebes.

Leptocera (Poecilosomella) longinervis Duda

Poecilosomella longinervis Duda, 1925 : 103.

E. NEPAL: Arun Valley, below Tumlingtar, River Sabhaya, west shore, c. 1,800 ft.,
human excreta in sandy place, 2 $, 9-22. xii. 1961 (R. L. Coe).

Described from Formosa. There are specimens in BMNH from India, Burma
and China. Duda omitted to give any description of the abdomen in his original
description, but a useful character for recognition is the very small size of the male
genital capsule in comparison with other species of the subgenus.

6o

J. C. DEEMING

Leptocera (Poecilosomelld) aciculata sp. n.

(Text-figs. 3, 9, 15)

A species closely resembling L. pallidimana Duda, but differing from it in the
chaetotaxy of the mid tibia; leg coloration; and the structure of the male genitalia,
in which species the clasper has a semi-triangular ploughshare-shaped anterior lobe
and a narrow posterior lobe bearing a pale apical tooth.

(J. Ground colour black; jowls and frons anteriorly and antenna basally dirty orange;
pleural sutures yellow ; haltere black with a yellow stem ; all tibiae basally and apically narrowly
yellow, the hind tibia distinctly and the mid tibia indistinctly with a yellow band centrally;
basal half of fore tarsus inf uscated, otherwise tarsus vivid white ; mid tarsus entirely yellow ; hind
basitarsus and next segment black, both narrowly white apically, the next two segments
yellowish white, the last black. Wing yellowish, with a white mark halfway along the first
vein extending downwards into the discal cell, another connecting the crossveins and a third
along the centre of the last section of the third vein; dark markings on the extreme wing base,
the humeral vein, from the apex of first vein to junction of second and third veins, on apex of
second vein, and also, though barely visible, on base and apex of ultimate section of third vein.
Lightly dusted with heavier, silvery dusting in patches. Head deeper than long; 4-5 pairs
almost equally-developed interf rental bristles ; the posterior of the two orbital bristles the longer ;
internal and external vertical bristles subequal ; one pair of strong postverticals ; frons strongly
silver-dusted on a median line, around ocelli, at bases of all bristles, on jowls, first antennal

\

FIGS. 3-8. Leptocera (Poecilosomella) spp., $ right tibia and basitarsus. 3, aciculata sp. n.
4, himalayensis sp. n. ; 5, annulitibia sp. n. ; 6, nepalensis sp. n. ; 7, brunettii sp. n.;
varians Duda.

DIPTERA FROM NEPAL 61

segment and at either side of base of antenna; interantennal prominence weakly developed;
vibrissa as long as an antenna; jowls three-sevenths of height of eye; third antennal segment
one and two-thirds times as long as second, rather pointed, with dense pilosity; arista two and
one-third times as long as remainder of antenna and with hairs of medium length. Thorax
with chaetotaxy typical of the subgenus; 8-10 rows of interdorsocentral setulae anteriorly; a
single pair of prescutellar acrostical hairs enlarged; scutellum as broad as long, rounded apically;
fore coxa strongly haired ; fore femur with a few posterodorsal bristles at 0-7 of its length and with
a row of long straight posteroventral hairs along its whole length; fore tibia rather densely
haired; mid femur not as strongly swollen as fore or hind, with one long erect anteroventral
bristle at extreme bare; mid tibia (Text-fig. 3) with a long anterior bristle at 0-6 of its length
and a shorter one at 0-52, a long anterodorsal at 0-28 and another still longer at 0-72, this last
surmounted by a smaller at 0-64, a posterodorsal paired with and shorter than most apical antero-
dorsal, a shorter posterodorsal at 0-32 and another still shorter at 0-56, ventrally on apical
half with long hairs, and amongst these on apical third are shorter, stouter and more erect
hairs ; mid basitarsus half as long as tibia and one and one-third times as long as second seg-
ment; hind femur strongly swollen, with a long erect basal ventral hair. Wing (Text-fig. 9)
with second vein lacking a stump-vein and strongly upcurved to costa, though not quite at
right angles to it; third vein strongly and evenly curved forward, ending a little further in
front of wing-tip than fold of fourth vein ends behind it, and slightly overpassed by costa;
second section of fourth vein slightly longer than posterior crossvein and only two-thirds times
as long as second section of third vein ; second costal sector slightly longer than third ; alula
normal. Abdomen with long lateral marginal bristles on tergites; genital capsule (Text-fig. 15)
with numerous rather sinuate hairs, especially ventrally; clasper posteriorly near base with
numerous downwardly-directed fine hairs, the anterior lobe sickle-shaped, the posterior squat and
bearing a short black curved apical tooth ; the slightly asymmetrical pregenital sternite half as
wide as abdomen, with a small median apical keel bearing minute dense pile. Length 2-5 mm.
$ unknown.

Holotype <$. E. NEPAL: Taplejung Distr., Sangu, c. 6,200 ft., mixed vegetation
by stream in gully, ix-x.i96i (R. L. Coe).

Paratypes. Same locality, rotting fruits of bhor tree on ground, I $, 7-31 . x . 1961 ;
Taplejung Distr., Dobhan, c. 3,500 ft., small pockets of plants on arid slopes above
R. Maewa, i <J, 2.1.1962 (R. L. Coe).

Leptocera (Poecilosomella) himalayensis sp. n.

(Text-figs. 4, ii, 16)

Similar to the preceding species (L. aciculata) but differing from it in the chaetotaxy
of the mid tibia, leg coloration, wing markings and the structure of the male genitalia.

cJ. Brownish black, paler on antenna, face, frons and pleural sutures, and with dusting typical
of the subgenus; all tibiae yellowish on a narrow basal, broad central and broad apical band;
tarsi yellowish, with apical segment infuscated, the bases of all basitarsi and the second segment
of hind tarsus infuscated. Wing (Text-fig. 1 1 ) yellowish with yellow veins ; a white mark covering
and connecting cross veins; dark marks at base, junction of second and third veins, and apices of
first, second and third veins. Haltere yellow. Head deeper than long with semi-oblique eye;
jowls two-sevenths times height of eye; four pairs of weak interfrontal bristles; the posterior of
the two orbital bristles slightly the longer; internal vertical bristle longer than external and as
long as ocellar; postocular hair adjacent to external vertical well developed; vibrissa long, with
only weak hairs behind it; arista two and a half times as long as remainder of antenna, and with
hairs of moderate length. Thorax with chaetotaxy and dusting as in preceding species; mid
femur with a row of five weak anterior bristles on apical half, preapically with a distinct anterior

62

J. C. DEEMING

bristle; mid tibia (Text-fig. 4) with a long anterior bristle at 0-7 of its length surmounted by a
bristle half as long, a long anterodorsal at 0-8 and a shorter one at 0-32, this last surmounted by a
bristle of half its length, a shorter posterodorsal paired with the most apical anterodorsal and
two short posterodorsal bristles at 0-4 and 0-6; apical part ventrally with long fine hairs, lacking a
stronger apical ventral bristle. Wing (Text-fig, n) with second and third costal sectors equal;
costa slightly overpassing apex of third vein; last developed section of fourth vein equal to
length of posterior crossvein; fourth vein reaching wing-margin as a fold. Abdomen with fine
hairs; a long hair on lateral margins of fourth and fifth tergites; genital capsule (Text-fig. 16) with
numerous fine hairs; posterior lobe of clasper bearing a droplet-shaped apical tooth; posterior
gonapophysis broad. Length about 2-7 mm.

? resembling $, but mid tibia lacking the long fine hairing ventrally and armed with an apical
ventral bristle that is longer than width of tibia; cercus rounded apically, densely covered in
short pile and bearing a few long fine rather sinuate hairs.

Holotype <. E. NEPAL: Taplejung Distr., river banks below Tamrang Bridge, c-
5,500 ft., x-xi.i96i (R. L. Coe).

Paratypes. i <$, i ?, same data.

Leptocera (Poecilosomella) annulitibia sp. n.

(Text-figs. 5, 10, 18)

Similar to the preceding species (L. himalayensis), but differing from it in leg
coloration and chaetotaxy, the presence of a stump-vein on the second wing-vein,
the structure of the male genitalia and other minor points.

11

13

14

FIGS. 9-14. Leptocera (Poecilosomella) spp., right wing. 9, aciculata sp. n. ; 10, annulitibia
sp. n. ; ii, himalayensis sp. n. ; 12, nepalensis sp. n. ; 13, brunettii sp. n.; 14, varians Duda.

DIPTERA FROM NEPAL 63

(J. Coloration as in L. himalayensis, but fore tibia lacking a central annulus, and the two
apical segments of fore and hind tarsi infuscated. Head deeper than long; jowls two-fifths times
height of eye; three pairs of weak interf rental bristles; orbitals equal; verticals equal; arista
three times length of remainder of antenna, with hairs of moderate length; fore femur with a row
of weak posterodorsal bristles on apical half and a row of weak posteroventrals on apical quarter,
ventrally with dense fine hairs that are not as long as femur is wide; fore tibia with similar
hairing; mid femur with a strong preapical anterior bristle and a row of four shorter bristles
leading up to it, and an apical posterior bristle ; mid tibia (Text-fig. 5) with a long anterodorsal
bristle at one-quarter and three-quarters of its length, both these bristles surmounted by a
much shorter bristle, much shorter posterodorsals paired with the long anterodorsals and
several short posterodorsals between them, a long anterior bristle surmounted by a much shorter
one intermediate between the long anterodorsals, ventrally on the apical two-thirds with stout
erect hairs and with several short apical bristles; hind leg normal. Wing (Text -fig. 10) with a
distinct stump-vein preapically on second vein; fourth vein distinctly produced as a fold to
wing-margin and ending slightly nearer wing-tip than third vein; fifth vein not reaching wing-
margin as a fold ; alula normal. Abdomen normal ; pregenital sternite (Text -fig. 18) with a narrow
and projecting median apical keel bearing dense pile and with two hairs at its base; posterior
lobe of clasper bearing a black tooth ; posterior gonapophysis broad and pointed. Length about
2-7 mm.

$ resembling <$, but frons darker; the long ventral hairing on fore and mid legs lacking; mid
tibia with one of the apical ventral bristles, though short, distinctly longer than the others;
cercus rounded apically, with a few fine sinuate hairs.

Holotype <J. E. NEPAL: Taplejung Distr., river banks below Tamrang Bridge,
c. 5,500 ft., x-xi.i96i (R. L. Coe).

Paratypes. i <$, i $, same data.

Leptocera (Poecilosomella) nepalensis sp. n.

(Text-figs. 6, 12, 17)

cj. Almost exactly resembling the preceding species (L. annulitibia), but differing from it in
the following respects : Four pairs of weak interfrontal bristles ; the long anterior bristle on mid
tibia (Text-fig. 6) situated much closer to the more basal of the two long anterodorsal bristles;
wing (Text-fig. 12) slightly broader; pregenital sternite (Text-fig. 17) with a wider median apical
keel; apical tooth on posterior lobe of clasper broader, resembling a molar; posterior gonapo-
physis narrower and rounded apically, and several other small differences in male genitalia.
Length about 2-7 mm.

$ unknown.

Holotype <. E. NEPAL: Taplejung Distr., river banks below Tamrang Bridge,
c. 5,500 ft., x-xi.ig6i (R. L. Coe).

Paratypes. i , same data; INDIA: Darjeeling, i <$, 13-18. ix. 1913 (E. Brunetti}.

Leptocera (Poecilosomella) brunettii sp. n.

(Text-figs. 7, 13, 19)

A species more closely resembling L. furcata Duda than any other species, but
easily distinguishable from it on account of the absence of a stump-vein preapically
on the second vein and the presence of a very long apical ventral bristle on the mid
tibia.

J. C. DEEMING

FIGS. 15-21. Leptocera (Poecilosomella) spp., apex of abdomen. 15, aciculata sp. n. ; 16,
himalayensis sp. n. ; 17, nepalensis sp. n. ; 18, annulitibia sp. n. ; ig, brunettii sp. n. ; 20,
sp. near brunettii sp. n.; 21, varians Duda.

DIPTERA FROM NEPAL 65

(J. Ground colour black; jowls, frons anteriorly and antenna dirty yellow; haltere, extreme
bases and apices of all tibiae, apices of all basitarsi, the next three segments on fore and mid
tarsus, and apex of second segment and all of third and fourth segments of hind tarsus yellow.
Wing (Text-fig. 13) yellowish, with rather ill-defined brown markings on humeral crossvein,
from apex of first across junction of second and third veins, at apex of second, across centre of
wing and at apex ; crossveins and section of fourth vein connecting them white. Dusting effect
typical of the subgenus; head deeper than long; eye two and one-third times as high as jowls,
noticeably emarginate at base of antenna; third antennal segment large, wide and heavily
pilose; arista 2-4 times as long as remainder of antenna, with hairs of moderate length; vertical
bristles subequal, as long as vibrissa; postvertical bristle as long as inwardly-directed upper
postocular; no enlarged setae preceding dorsocentral bristles, ten rows interdorsocentral setulae
between the anterior pair; the prescutellar acrosticals enlarged; scutellum broader than long,
rather truncate apically ; all legs short-haired ; mid femur with a row of five anterior bristles on the
apical third; mid-tibia (Text-fig. 7) with a long anterior bristle situated at 0-72 of its length and
surmounted by a much shorter bristle, a long anterodorsal at 0-33 and another at 0-78, both of
these surmounted by a shorter bristle, a rather short posterodorsal at 0-71 and a very short one
at 0-4, the apical ventral bristle half as long as the basitarsus and surrounded at its base by several
much shorter and weaker bristles. Wing (Text-fig. 13) broad; costa slightly overpassing third
vein; second costal sector a little shorter than third; second vein angularly upcurved to costa;
third vein strongly curved forwards, ending further in front of wing-tip than fold of fourth vein
ends behind; posterior crossvein longer than distance between crossveins; alula rather narrow
but rounded apically. Abdomen normal; tergites bearing single apical bristles laterally and a
few weak hairs on posterior margins; pregenital sternite (Text -fig. 19) with a pilose median
apical keel ; posterior lobe of clasper with a black apical tooth and another on external surface ;
posterior gonapophysis short, broad and apically pointed. Length about 2 mm.

$ unknown.

Holotype $. INDIA: Darjeeling, 13-18. ix. 1913 (E. Brunetti}.

Leptocera (Poecilosomella) sp. near brunettii Deeming
(Text-fig. 20)

A single male of an undescribed species very closely resembling the preceding
species, having identical wing-markings and the same distinctive chaetotaxy of the
mid tibia. However, in this species the second vein is less angularly upcurved to
the costa, the second and third costal sectors are equal, and there are differences in the
male genitalia (Text-fig. 20), these being the lack of the external tooth on the posterior
lobe of the clasper, and the long, thin, sinuous and apically rounded shape of the
posterior gonapophysis.

As the specimen is in poor condition it is deemed an insufficient basis for the erection
of a new species. A figure of the male genitalia is given to facilitate the separation
of the two species.

ASSAM: Shillong, i <$, i6.x.ig20 (R. Senior-White).

Leptocera (Limosina) mirabilis Collin

Limosina mirabilis Collin, 1902 : 19.

E. NEPAL: Taplejung Distr., damp evergreen oak forest above Sangu, c. 9,200 ft.,
i $, 2-26 . xi . 1961 ; Taplejung Distr., above Sangu, c. 6,500 ft., swarming on cowdung,
i$, 5.x. 1961 (R. L. Coe).

Recorded from North America, Europe and New Zealand.

66 J. C. DEEMING

Leptocera (Limosina) moesta Villeneuve
Limosina (Leptocera) moesta Villeneuve, 1917 : 337.

W. NEPAL: Bakhri Kharka, 5, 500 ft., i $, 24. iv. 1954 (/. Quinlari); E. NEPAL:
Taplejung Distr., river banks below Tamrang Bridge, c. 5, 500 ft., i $, x-xi.igGi
(R. L. Coe).

A common European species.

Leptocera (Limosina) brevicostata Duda

Limosina brevicostata Duda, 1918 : 183.

E. NEPAL: Taplejung Distr., above Sangu, c. 6,500 ft., swarming on cowdung,
1$, 5.x. 1961; Taplejung Distr., below Sangu, c. 6,000 ft., by stream in shady
ravine, i ?, 30. x. 1961 (R. L. Coe).

Described from Europe, the species is widely distributed in the Old World, its
distribution extending as far east as Polynesia. Duda (1925 : 188) described a
variety from the Orient and New Guinea under the name of Scotophilella brevicostata
var. rufifrons on the basis of a slight variation in coloration, wing venation and
tarsal enlargement. I prefer to regard the species as variable, as specimens that I
have seen from Uganda, which agree with var. rufifrons in having very swollen hind
tarsi, have the venation and coloration of the typical form. Other specimens from
entirely different regions have been difficult to place in either one form or the other.
A specimen in BMNH from N. Rhodesia was reared in company with Sepsis later alis
Wied. from a dead but unparasitized Nomadacris septemfasciata Serv. (Acrididae).

Leptocera (Limosina) lutea Richards

Leptocera (Limosina) lutea Richards, 19636 : 127.

E. NEPAL: Taplejung Distr., Dobhan, c. 3,500 ft., shady places on shrubby slope
above River Tamur, i $, i ?, 21-27.1.1962 (R. L. Coe).

Described from W. Caroline Islands. There are specimens in the Zoological
Museum, Copenhagen from the Philippines.

Leptocera (Limosina) rufilabris Stenhammar
Limosina rufilabris Stenhammar, 1855 : 408.

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
plants by damp cliff in deep river gorge, 2 $, i-ii.1962 (R. L. Coe}.
W. NEPAL: 2 mis. S.W. of Rambrong, i <$, 8,000 ft. (/. Quinlan).
A common European species.

Leptocera (Limosina) sp. near rufilabris Stenhammar

E. NEPAL: Taplejung Distr., Sangu, c. 6,200 ft., mixed vegetation by stream in
gully, i ?, xi< 1961-!. 1962; Taplejung Distr., edge of mixed forest above Sangu, c.
6,500 ft., i ?, I7.x-i.xi.i96i (R. L. Coe).

DIPTERA FROM NEPAL

67

Both specimens are slightly larger and longer-winged than typical rufdabris, and
the legs are more yellowish as in L. femorina Richards, but the second costal sector
is not as thickened as it is in that species.

Leptocera (Limosina) ? fucata Rondani

(Text-figs. 22 to 27)
Limosina fucata Rondani, 1880 : 19.

E. NEPAL: Taplejung Distr., below Sangu, c. 6,000 ft., by stream in shady ravine,
i (, 30.x. 1961 (R. L. Coe).

This species exhibits such a diversity of form that I am inclined to regard it as a
complex of species. Richards (1952 : 89) described a very closely-related species
Leptocera trichopyga from Austria, differing from L. fucata in wing venation and the
chaetotaxy of the mid leg. The Nepalese specimen and specimens from various
localities in Europe that I have seen have just such a wing venation as L. trichopyga,
and between them have a complete range of development of the apical ventral
bristle of the mid tibia from that described by Duda for fucata (normal) to that
described by Richards for trichopyga (absent). However, all these specimens lack
the enlarged mid basitarsal bristle present in trichopyga. Examination of the claspers
and internal genitalia of the Nepalese male and comparison with those of the
European specimens and an uncertainly-identified L. fucata (Coe, 1962 : 118) from
Macedonia leads one to no definite conclusion as to its identity or the relationship
between the European specimens examined. A great deal more material will have to
be examined before it can be decided whether L. fucata is merely a very variable species
or whether there are other species commonly confused with it. L. fucata was
described from Europe, has been redescribed with wing-figures by Duda (1918 : 148,
fig. 42 and 1938 : 123, fig. 39), and a short redescription is given by Collin (1956 :

27

FIGS. 22-28. Leptocera (Limosina) spp. 22-27, ^fucata Rond., $ internal genitalia. 22,
Nepal, Taplejung Distr.; 23, Germany (Gottingen); 24, Germany (Teutoburger Wald);
25, Austria (Otz); 26, Italy (Dolomites); 27, Jugoslavia (Otesevo). 28, coei sp. n., right
wing.

68 J. C. DEEMING

173) in recording the species from Britain under the synonym Limosina verticella
Stenhammar.

Leptocera (Limosina) ? fungicola Haliday
Limosina fungicola Haliday, 1836 : 330.

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
plants by damp cliff in deep river gorge, i $, i-ii. 1962 (R. L. Coe).

This specimen has a median apical keel on the pregenital sternite, but agrees in
all other respects with L. fungicola.

Leptocera (Limosina) sp. (i) near paraminima (Duda)

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
shrubs in deep gorge, i $, x-xi.i96i; Taplejung Distr., river banks below Tamrang
Bridge, c. 5,500 ft., 2 <, x-xi.ig6i; Taplejung Distr., Dobhan, c. 3,500 ft., spray-
splashed rocks in River Maewa, i <$, 25 .i. 1962 (R. L. Coe).

This species runs to Scotophilella paraminima Duda in Duda's 1925 key, but differs
from it in that the second section of the fourth vein is over twice as long as~the
posterior crossvein. As the description, based on a single female, is scanty and too
inadequate for comparison, the species is left undescribed.

Leptocera (Limosina) sp. (2) near paraminima (Duda)

E. NEPAL: Taplejung Distr., Sangu, c. 6,200 ft., mixed vegetation by stream in
gully, i (J, xi . 1961-1 . 1962 (R. L. Coe).

This species differs from the preceding in the presence of a large and rounded median
apical keel on the pregenital sternite in the male. A single female labelled Taplejung
Distr., between Sangu and Tamrang, c. 5,200 ft., mixed plants by damp cliff in deep
"iver gorge, i-ii . 1962 may belong to either this or the preceding species.

Leptocera (Limosina) sp. (i) near schmitzi (Duda)

E. NEPAL: Taplejung Distr., Sangu, c. 6,200 ft., mixed vegetation by stream in
gully, i $, xi . 1961-1 . 1962 (R. L. Coe).

Superficially almost exactly resembling L. schmitzi, but differing from it in having a
well developed median apical keel on the pregenital sternite, a smaller genital
capsule and less quadrate claspers.

Leptocera (Limosina) sp. (2) near schmitzi (Duda)

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
plants by damp cliff in deep river gorge, i <$, i ?, i-ii. 1962 (R. L. Coe).

Similar to the preceding species, but differing from it and L. schmitzi in the dirty
yellow coloration of the fore coxa. In the female the second costal sector is slightly
shorter.

DIPTERA FROM NEPAL 69

Leptocera (Limosina) sp. near boliviensis (Duda)

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,500 ft., mixed
vegetation by stream in deep gully, i $, i-ii.i962 (R. L. Coe).

Differs from L. boliviensis in the wing being narrower, the second costal sector
longer and in other respects. All legs are bright yellow, only the mid and hind femora
being somewhat infuscated; fore coxa with a pair of short stout bristles on inner
surface near base ; pregenital sternite very long and with a distinct median apical keel.

Leptocera (Limosina) sp. near penetralis Collin

E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200 ft., mixed
plants by damp cliff in deep river gorge, I $, 1-^.1962 (R. L. Coe).

Differs from L. penetralis in that the apical section of the third vein, though
straight, ends nearly as far in front of the wing-tip as the fold of the fourth vein ends
behind it.

Leptocera (Limosina) coei sp. nov.

(Text-fig. 28)

A species that runs to L. chilenica Duda (couplet 43) in Duda's 1925 key, but
easily distinguishable from it on account of its single pair of dorsocentral bristles.

cJ. Ground colour dark brown; head, with exception of occiput, upper orbits, clypeus,
palpus and proboscis golden orange; bases of meso- and pteropleuron, whole of metapleuron,
haltere, trochanters, extreme bases and apices of all tibiae, and all tarsi yellow; wing yellowish
hyaline with yellow veins. Head deeper than long, lightly dusted; frons strongly convex;
interantennal keel strongly pronounced; jowls one-quarter height of eye; an upwardly-directed
bristle behind the vibrissa and half its length ; four pairs of strong interfrontal bristles ; ocellar
bristle long, as long as internal vertical ; external vertical shorter ; the orbitals hardly longer than
the postvertical ; third antennal segment large, rather pointed and densely pilose; arista rather
long-haired, two and a half times as long as remainder of antenna. Disc of thorax densely
grey-dusted, uniformly covered with dense short setulae; a single pair of fine dorsocentral
bristles that reach slightly beyond tip of the semicircular scutellum and a slightly enlarged seta
situated just in front of each; fore femur rather swollen, with a row of four fine posterodorsal
bristles on the apical two-thirds and similar posteroventrals on apical third; mid femur with a
strong preapical anterodorsal bristle; mid tibia with a short anteroventral bristle at 0-55 of its
length, progressively longer anterodorsals at 0-27, 0-45 and 0-8, a dorsal as long as this last
bristle at 0-75, longer posterodorsals at 0-35 and 0-8 with shorter bristles between them at 0-5
and 0-7, and an apical ventral that is two-fifths the length of basitarsus; hind leg normal. Wing
(Text-fig. 28) broad; costa rather densely short haired; interspersed amongst the reclinate hairs
on second and third costal sectors are fine short hairs projecting at right angles to costa 1 ;
second and third costal sectors equal ; third vein uniformly curved forwards, not overpassed by
costa, and ending well in front of wing-tip; fold of fourth vein not produced to wing-margin,
strongly curved forwards; outer angles of discal cell rounded, especially the posterior; alula small
and narrow. Abdomen less strongly dusted than disc of thorax, with numerous short hairs

1 This same character, though seemingly insignificant in this species, may be of value in placing the
species in relationship to others as yet undescribed. I have before me an undescribed and obviously
closely related species from the Philippines in which these hairs are outstandingly developed and set at
an angle to the wing-surface in much the same way as they are in species of the subgenus Elachisoma
Rondani.

70 J. C. DEEMING

and weak marginal bristles on the tergites; genital capsule hemispherical and densely short
haired ; clasper with dense sinuate hairs, band -shaped, slightly swollen apically and bearing an
incurved black pointed tooth; posterior gonapophysis curved and pointed; pregenital sternite
simple. Length about 1-7 mm.

$ resembling <$, abdomen broad and flattened ; tergites two and four longer than three and five ;
sternites narrower and broadly separated from the tergites by membrane; cercus weakly
sclerotized, with several fine, sinuate hairs.

Holotype <. E. NEPAL: Arun Valley, below Tumlingtar, River Sabhaya, west
shore, c. 1,800 ft., human excreta in sandy place, 9-22 . xii . 1961 (R. L. Coe).

Paratypes i $, same data ; same locality, dead leaves lying in sun on sandy shore,
2 <$, 22. xii. 1961 (R. L. Coe); INDIA; Silchar, Cachar, i <$, 3 . in . 1911 (C. B. Antram).

Leptocera (Limosind) furculisterna sp. n.

(Text-figs. 29, 30)

A species which, though rather different from L. vitripennis Zetterstedt, is probably
better likened to it than to any other, on account of the structure of the tergites,
which though less strongly differentiated in vitripennis are nevertheless differentiated
to a noticeable degree. In vitripennis the second costal sector is black in contrast
to the brown first and third sectors, but in the species described below the costa is
uniformly yellow.

<J. A jet-black species, lightly dusted on all but the frontal triangle, the antennal foveolae
and a patch anteriorly on the upper sternopleuron ; antenna, stem of haltere, fore coxa, all
trochanters, basal half of fore femur, extreme bases of mid and hind femora, mid tibia and
tarsus, hind tibia with the exception of a broad preapical annulus, last three tarsal segments
of hind leg strikingly yellow; last four tarsal segments of fore leg whitish yellow; wing yellowish
with yellow veins. Head deeper than long ; jowls one-fifth height of eye ; ocellar triangle extending
to interantennal keel, with three pairs fine hairs ; internal vertical bristle longer than the external,
the orbitals shorter still, scarcely longer than the post vertical ; an upcurved mid jowlar bristle that
is nearly half as long as vibrissa; third antennal segment short, long-pilose apically, arista situated
preapically, rather long-haired and slightly over three times as long as remainder of antenna ; two
widely-separated pairs dorsocentral bristles ; only four rows of interdorsocentral setulae between
anterior pair; scutellum semi-triangular; fore femur slightly swollen, with a few fine anterodorsal
and posterodorsal hairs ; fore basitarsus very narrow, the remaining four tarsal segments becoming
progressively more swollen, the most apical broader than the tibia ; mid femur with a few weak ant-
erodorsal hairs ; mid tibia with a long posterodorsal bristle at 0-92 of its length, a slightly shorter
anterodorsal at 0-33 and a short one at 0-8, this last as long as the apical ventral. Wing (Text-fig.
29) with rather convex fore margin ; costa densely short-haired on second and third sectors, over-
passing third vein to a distance equal to length of anterior crossvein, and with second sector
shorter than third; third vein very strongly bowed, firstly towards and then away from costa;
first section of third vein much longer than second, equal to second section of fourth vein;
fourth and fifth veins ending slightly beyond discal cell; alula very narrow and pointed.
Abdomen (Text-fig. 30) fine-haired; tergite two broad, long and heavily sclerotized, the next
three tergites narrower, much shorter, weakly sclerotized and broadly separated by membrane
(in dry specimens these tergites crumpled and scarcely recognizable as such), the remaining
tergites heavily sclerotized ; sternites becoming progressively wider and longer towards the apex,
the pregenital broad, laterally long-haired, bearing a short-bristled, square median apical keel
that is flanked by an inwardly-curved appendage bearing short bristles apically; clasper large,
rather rectangular, the exterior surface and anterior margin clothed with fine hair; posterior

DIPTERA FROM NEPAL 71

gonapophysis short, broad, with a concave posterior margin between two short points. Length
about 1-5 mm.
<j> unknown.

Holotype $. E. NEPAL: Taplejung Distr., between Sangu and Tamrang, c. 5,200
ft., mixed plants by damp cliff in deep river gorge, i-u . 1962 (R. L. Coe).

Paratype. Same locality and altitude, mixed shrubs in deep gorge, I <$, x-xi . 1961
(R. L. Coe).

Leptocera (Limosina) sp. near furculisterna Deeming
(Text-fig. 31)

Due to their rubbed condition, no name is given to a pair of specimens of an
undescribed species closely resembling L. furculisterna, but the characters necessary
for distinguishing it from that species are given.

$. Interantennal keel, extreme anterior margin of frons, and mouth-margin medially yellow;
fore coxa black; fore femur with exception of extreme apex black; hind tibia lacking preapical
black annulus; all tarsi dirty yellow, the apical segments distinctly infuscated; pregenital
sternite (Text-fig. 31) with a double row of bristles bordering the weakly sclerotized median
apical keel, lacking the flanking appendages, and with a pair of long bristles near its lateral
margins ; clasper rectangular, with several rather stout bristles basally on the exterior surface and
a pair of minute teeth apically on the posterior margin.

$ resembling $, tergites of the same differentiated structure as the male; cercus with one long
sinuate hair and several shorter ones.

E. NEPAL: Arun Valley, below Tumlingtar, River Sabhaya, west shore, c. 1,800 ft.,
dead leaves lying in sun on sandy shore, I $,22. xii . 1961 ; Arun Valley, east shore of
River Arun below Tumlingtar, c. 1, 800 ft., evergreen shrubs bordering dry stream
beds, i $, 14-23 . xii . 1961 (R. L. Coe}.

Leptocera (Limosina) monorbiseta sp. n.

(Text-fig. 32)

A species best likened to L. mollis Richards on the basis of the single orbital
bristle, but differing from it in the colour of the head and in other respects. Due to
the appearance of the antenna, it could be mistaken for a species of the subgenus
Acuminiseta Duda, but the position of the arista on the third antennal segment
places it in Limosina.

$. Ground colour black, lightly dusted; head, with exception of clypeus, proboscis, a tri-
angular patch at base of vibrissa, a fine line extending from base of internal vertical bristle
far into the occiput, and apex of third antennal segment, yellow; all trochanters, extreme base
and apex of mid tibia, hind tibia with exception of a broad preapical annulus and all tarsi bright
yellow; haltere, fore femur, and fore tibia with exception of an indistinct central annulus dirty
yellow. Head much deeper than long; frons strongly projecting in front of eye; face retreating
beneath; eye five times as high as jowls; three pairs of short weak interf rental hairs; orbits
bearing a single orbital bristle and a row of minute setae; external vertical bristle longer than
internal, which is equal to the ocellar; postvertical bristle minute, slightly shorter than the
incurved upper postocular; vibrissa long, with a few weak hairs behind; the upwardly-directed

72 J. C. DEEMING

buccal bristle short ; third antennal segment heavily pilose apically, giving it a pointed appearance
although its apex is rounded ; arista inserted almost apically, three times as long as remainder of
antenna and with hairs of moderate length. Setulae on disc of thorax long and fine, the pre-
scutellar pair no longer than the others ; two pairs of long dorsocentral bristles, the more anterior
pair the shorter and with eight rows setulae between; scutellum semicircular, the lateral pair
marginal bristles scarcely longer than the distance separating them from the apical pair; fore
femur with a pair of strong posteroventral bristles preapically ; mid femur with a strong preapical
anterior bristle; mid tibia with short anterodorsal bristles at 0-27 and 0-68 of its length, similar
posterodorsals at 0-37, 0-51 and 0-68, a long dorsal at 0-74 and a long apical ventral; mid basi-
tarsus twice as long as second segment ; second segment of hind tarsus strongly swollen, slightly
wider than basitarsus; wing (Text-fig. 32) much broader than thorax, with densely haired costa
and narrow and pointed alula; second costal sector shorter than third; second vein strongly,
third weakly sinuate; basal section of third vein with two breaks, shorter than second section;
fourth vein produced beyond discal cell as a rather indistinct fold that does not reach wing-
margin; abdomen very flat dorsally, rounded ventrally; first tergite medially unsclerotized ;
tergites 2-5 short, subequal, becoming progressively narrower; sternites weakly sclerotized
and broadly separated from tergites by membrane ; cercus with a long sinuate dorsal and apical
hair and with several much shorter hairs ventrally. Length about 1-7 mm.
$ unknown.

Holotype $. E. NEPAL: Taplejung Distr., old mixed forest above Sangu, c.
6,200 ft., 25-28. x. 1961 (R. L. Coe).

29

FIGS. 29-32. Leptocera (Limosind) spp. 29, furculisterna sp. n., right wing ; 30, furculisterna
sp. n., $ abdomen; 31, sp. near furculisterna sp. n., <$ abdomen; 32, monorbiseta sp. n.,
right wing.

Leptocera (Trachyopella) obliqua Richards

Leptocera (Trachyopella) obliqua Richards, 19636 : 129.

E. NEPAL: Taplejung Distr., above Sangu, c. 6,500 ft., evergreen scrub, i
5-13. x. 1961 (R. L. Coe}.

DIPTERA FROM NEPAL 73

Described from the Mariana and Caroline Islands. There are specimens in
BMNH from Java, Celebes and Malaya.

Leptocera (Coproica) hirtula Rondani
Limosina hirtula Rondani, 1880 : 38.

E. NEPAL: Taplejung Distr., Dobhan, c. 3,500 ft., shady places on shrubby slope
above River Tamur, 2 ?, 21-27^.1962 (R. L. Coe).
A common and cosmopolitan species.

REFERENCES

ADAMS, C. F. 1905. Diptera Africana I. Kans. Univ. Sci. Bull. 3 : 147-208.

BRUNETTI, E. 1913. Zoological Results of the Abor Expedition 1911-12. Diptera. Rec.

Indian Mus. 8 : 149-190.
1924. Diptera of the Siju Cave, Garo Hills, Assam. I. Tipulidae, Tabanidae, Antho-

myidae, Acalyptrate Muscidae and Phoridae. Rec. Indian Mus. 26 : 99-106.
COE, R. L. 1962. A further collection of Diptera from Jugoslavia, with localities and notes.

II. Glas. Muz. srp. Zeml. (B) 18 : 95-136.
COLLIN, J. E. 1902. Four new species of Diptera (Fam. Borboridae) found in Britain.

Entomologist's mon. Mag. 13 : 55-60.

1925. Diptera (Cyclorrhapha) from Spitzbergen. Results of the Oxford Expedition to
Spitzbergen, 1924. Ann. Mag. nat. Hist. (9) 16 : 332-337.

1956. Some new British Borboridae (Diptera) /. Soc. Br. Ent. 5 : 172-178.
DEEMING, J. C. 1964. A remarkable new species of Leptocera from Christmas Island and New

Guinea. Opusc. ent. 29 : 164-167.

DUDA, O. 1918. Revision der europaischen Arten der Gattung Limosina Macquart (Dipteren).
Abh. zool.-bot. Ges. Wien 10 : 1-240.

1920. Revision der altweltlichen Arten der Gattung Sphaerocera Latreille (Dipteren).
Tijdschr. Ent. 62 : 1-38.

1923. Revision der altweltlichen Arten der Gattung Borborus (Cypsela) Meigen (Dipteren).
Arch. Naturgesch. 89 : 35-112.

1925. Die aussereuropaischen Arten der Gattung Leptocera Olivier = Limosina Macquart
(Dipteren) mit Beriicksichtigung der europaischen Arten. Arch. Naturgesch. 90 (A) (1924) :

5-215-

1938. Sphaeroceridae (Cypselidae) . In LINDNER: Fliegen palaearkt. Reg. 57 : 1-182.
HACKMAN, W. 1965. On the genus Copromyza Fall. (Dipt., Sphaeroceridae), with special

reference to the Finnish species. Notul. ent. 45 : 33-46.

1967. Sphaeroceridae (Diptera) from Madagascar. Notul. ent. 47 : 25-36.

HALIDAY, A. H. 1836. British Species of the Dipterous Tribe Sphaeroceridae. Entomological

Magazine^ : 315-336.
MEIGEN, J. W. 1830. Systematische Beschreibung der bekannten Europaischen zweiflugeligen

Insekten. 6 : 1-401. Aachen & Hamm.
RICHARDS, O. W. 1930. The British Species of Sphaeroceridae (Borboridae, Diptera). Proc.

zool. Soc. Lond. 18 : 261-345.

1952. Two new Austrian species of Leptocera Oliv. (Dipt. : Sphaeroceridae). Proc. R.
ent. Soc. Lond. (B) 21 : 89-91.

19620. Species of Copromyza allied to sordida (Diptera : Sphaeroceridae) with notes on
types of African Sphaeroceridae described by C. F. Adams. /. Kans. ent. Soc. 35 : 364-368.

- 19626. Contribution a l'6tude de la faune d' Afghanistan. 42. Diptera, Sphaeroceridae.
Entomologist's mon. Mag. 97 : 177-179.

74

J. C. DEEMING

RICHARDS, O. W. 1963*1. Sphaerocerid flies from South and Central America in the collection

of the California Academy of Sciences (Diptera). Pan-Pacif. Ent. 39 : 231-246.

19636. Diptera : Sphaeroceridae (Borboridae) . Insects Micronesia 14 : 109-134.

RONDANI, C. 1880. Species Italicae ordinis Dipterorum (Muscaria, Rnd.) collectae et

observatae. Stirps XXV. Copromyzinae Zett. Bull. Soc. ent. Ital. 12 : 3-45.
STENHAMMAR, C. 1853. Copromyzidae Scandinaviae granskade och beskrifne. K. svenska

VetenshAkad. Handl. 41 : 257-442.

1855. Copromyzinae Scandinaviae. Holmiae. 184 pp.

VILLENEUVE, J. 1917. Especes nouvelles de Dipteres de la famille des Cypselidae (Borboridae).

Bull. Soc. ent. Fr. 18 : 333-338.
WIEDEMANN, C. R. W. 1824. Analecta Entomologica ex Museo Regio Hafniensi maxime

congesta. (154 Dipter., 3 Hymenopt., 2 Coleop.) Kiliae, (Reg. typogr. Schol.) 4. 60 pp.

aciculata sp. n., 60
annulitibia sp. n., 62

Borborillus Duda, 56
brevicostata Duda, 66
brunettii sp. n., 63

coei sp. n., 69
Coproica Rond., 72
Copromyza Fall., 56
curvinervis Stenh., 56

fucata Rond., 67
fungicola Hal., 68
furculisterna sp. n., 70

himalayensis sp. n., 61
hirtula Rond., 73

koningsbergeri Duda, 57

Leptocera Oliv., 56
Limosina Macq., 65
longinervis Duda, 59
Lotobia Lioy, 56
lutea Rich., 66

INDEX

marginatis Adams, 56
mirabilis Coll., 65
moesta Villen., 66
monorbiseta sp. n., 71

nepalensis sp. n., 63
nigrolimbata Duda, 56

obliqua Rich., 72
Opacifrons Duda, 57

pallidiventris Meig., 56
paranigrolimbata Duda, 57
Poecilosomella Duda, key to species, 58
pseudimpudica sp. n., 57
punctipennis Wied., 59

Rachispoda Lioy, 57
rufilabris Stenh., 66

Sphaerocera Latr., 56
subtinctipennis Brun., 57

Trachyopella Duda, 72
varians Duda, 59

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. 5.

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp. 284 ; 348 Text-figures. August, 1965. 6.

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 ;
18 plates, 270 Text-figures. August, 1965. 4 45.

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 ; 500 Text-figures. October,

1965- 35s.

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 ;

475 Text-figures. November, 1965. 2 i$s.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae.
Pp. 129 ; 328 Text-figures. May, 1966. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera : Coccoidea). Pp. 168 ; 43 Text-figures. February, 1967.

33s.

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera : Geometridae). Pp. 119; 14 plates, 146
Text-figures, 9 maps. February, 1967. 3 los.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 los.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopa-
locera). Pp. 322; 233 Text-figures, coloured frontispiece. August, 1967. 8.

H. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 184;
82 Text-figures. May, 1968. 4.

12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with an
Account of their World Distribution (Lepidoptera : Drepanidae). Pp. 151 :
293 Text-figures, 14 plates. November, 1968. 5.

PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, BARTHOLOMEW PRESS, DORKING

C-

REVISIONAL NOTES ON AFRICAN

CHARAXES

(LEPIDOPTERA : NYMPHALIDAE)

PART V

V. G. L. van SOMEREN

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 23 No. 4

LONDON: 1969

RE VISIONAL NOTES ON AFRICAN

CHARAXES

(LEPIDOPTERA : NYMPHALIDAE)
PART V

BY

V. G. L. van SOMEREN

P.O. Box 24947, Karen, Kenya

Pp. 75-166; 8 Maps, 29 Plates, 31 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. 4

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

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

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

This paper is Vol. 23, No. 4 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) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued ii April, 1969 Price 4 ios.

RE VISIONAL NOTES ON AFRICAN

CHARAXES

(LEPIDOPTERA : NYMPHALIDAE)
PART V

By V. G. L. van SOMEREN

CONTENTS

Page
SYNOPSIS ........... 77

1. SUPPLEMENTARY NOTES AND ADDENDA ON SPECIES PREVIOUSLY DEALT

WITH ........... 77

2. THE Charaxes cedreatis-vetula COMPLEX AND Charaxes chepalungu A

NEW SPECIES ......... 84

Systematic List ......... gi

3. Charaxes virilis ROTHSCHILD, Ch. kheili STAUDINGER, Ch. northcotti

ROTHSCHILD AND Ch. mafuga SP. N. . . . . . . 92

Systematic List ......... 98

4. THE Charaxes etheocles COMPLEX ....... 98

Systematic List ......... 114

5. Charaxes grahamei SP. N. . .... . .115

6. Charaxes contrarius WEYMER AND Charaxes petersi SP. N. . . . 119

7. Charaxes baileyi VAN SOMEREN . . . . . . . 122

Systematic List ......... 125

8. Charaxes viola BUTLER AND ITS SUBSPECIES AND FORMS . . . 125

Systematic List ......... 147

9. Charaxes cynthia BUTLER . . . . . . .149

Systematic List . . . . ..... 159

ACKNOWLEDGEMENTS ......... 160

REFERENCES ........... 165

INDEX ............ 165

SYNOPSIS

In this paper three new subspecies and two new forms are described of species already dealt
with in previous parts of the Bulletin and four new species, six new subspecies and thirteen new
forms are described.

i. SUPPLEMENTARY NOTES AND ADDENDA ON SPECIES

PREVIOUSLY DEALT WITH

Charaxes acuminatus Thurau
(van Someren, 1963 : 211-220)

Charaxes acuminatus obudoensis ssp. n.

(PL 28, figs. 243, 246)

THIS distinct race is represented by several specimens of both sexes, collected by
Mr. R. G. T. St. Leger on the Obudo Plateau at 5,200 ft. in eastern Nigeria, formerly

ENTOM. 23, 4. 8

78 V. G. L. VAN SOMEREN

part of British Cameroons. The discovery of this race extends the range of
Ch. acuminatus very considerably to the west.

MALE. Fore wing length 46 mm. Apex of wing strongly acuminate, tapering and slightly
flattened at end of costa, typical of the species. Upperside. Resembles somewhat Ch. a.
cottrelli of the Mwinilunga area of N.W. Zambia in general appearance, but differs markedly by
the restricted and more defined bluish white basal areas of both wings ; the distal portions of the
wings are a richer orange, with the dark pattern more denned. Underside. Generally darker
in ground colour, but somewhat variable in rufous brownish tones, with a strong satiny sheen
on distal borders; oblique dark line strong.

FEMALE. Fore wing length 53 mm. Apex slightly more attenuated than in the male.
Upperside. Pattern generally similar. Underside. Darker more greyish olive.

Holotype male. E. NIGERIA: Obudo Plateau, 5,200 ft., I2.xii.i963 (R. G. T. St.
Leger). InB.M. (N.H.).

Allotype female. E. NIGERIA: Obudo Plateau, 25^.1965 (R. G. T. St. Leger).
InB.M. (N.H.).

Charaxes druceanus Butler
(van Someren, 1963 : 228-240)

Charaxes druceanus tectonis Jordan
(PL 29, figs. 251, 254)

Charaxes tectonis Jordan, 1937: 324. Type male, Cameroon, Maungli.
Charaxes druceanus tectonis Jordan; van Someren, 1963 : 230-231.

Originally described as a species, possibly allied to Charaxes eudoxus or " fallax "
Richels = richelmanni Rob., tectonis was placed by me (1963) as a subspecies of
druceanus, for reasons stated. Some doubt was expressed as to whether the under-
side pattern of the unique type could be typical of the race as a whole, in view of the
fact that very similar variations had been noted in other subspecies of druceanus.
This doubt has now been dispelled as a result of the capture of additional specimens
on the Obudo Plateau, eastern Nigeria (at one time part of the territory included
within the Kamerun, then under German control, subsequently administered by
Britain under Mandate).

These specimens were taken by Mr. R. G. T. St. Leger in 1964-65, but all were
males, exhibiting a reasonable constancy of underside pattern and agreeing with
that of the type, i.e. suppression of the broad silvery discal bars of fore and hind
wings, but otherwise conforming to the druceanus pattern below and above. Mr.
St. Leger supports my view that tectonis is a subspecies of druceanus.

During a brief visit to Nigeria over the Easter vacation, 1966, a young friend of
mine visited the Obudo Plateau in company of Mr. St. Leger, and by the extensive
use of baited traps, he was fortunate in securing not only males but also two examples
of the hitherto unknown female. These latter exhibit the broad silvery white bars
on the underside, characteristic of druceanus, thus supplying corroborative evidence
that tectonis is a race of that species.

FEMALE. Fore wing length 38mm. Upperside. Base of fore wing dark chestnut; black
spots in the cell and through the discal line and on the distal portion of the wing as in the male ;

REVISIONAL NOTES ON AFRICAN CHARAXES 79

the series of spots forming the discal and postdiscal pattern as in the male, but orange-ochreous,
the marks in ia-3 coalescing, those of the postdiscal series rather more rufous ; margin with distinct
rufous spots, double in ib. Hind wing pattern generally similar to that of the male, but the
basal triangle more blackish brown; the discal band denned on the inner border and almost
straight, extending from the costa, where it is 7 mm. wide, gradually decreasing in width to
above anal angle, colour orange-ochreous, shaded with rufous on outer border. Black band
broad, carrying blue spots in 2-3, double at anal angle; marginal border with well defined
orange-ochre contiguous lunules, accentuated by a marginal black line; margin rather serrate
below 5 ; tails longish, upper 7 mm., lower 6 mm., followed by some olive at anal lobe. Underside.
Fore wing subcostal area and bases of ib-4 chestnut; the subcostal black marks broadly outlined
in silver ; sub-basal black marks in i b-3 strongly black ; the discal-postdiscal band whitish, shaded
with orange on edges, followed by a series of black lunules of decreasing size from ib to subapex,
slightly accentuated by bluish white; marginal border rufous, with greyish olive rays along
veins. Hind wing basal area chestnut, with black lines strongly edged in silvery white, the
black angled line crossing ib-ic to inner fold. Discal band, widest at costa and almost straight
along inner edge, tapers to above the anal angle where it crosses the inner fold at right angles;
postdiscal chestnut bar widest at 3, fading to ochreous olive to above the anal angle and is distally
bordered by a submarginal zone of greyish olive, flushed with rufous scales, the zone outlined
in black, the black lines forming a double ocellus at anal lobe. Marginal border orange-tawny,
paler between tails, more olive at anal lobe; margin black.

Neallotype female. E. NIGERIA: Obudo Plateau, 5,200 ft., i6.iv.66 (Stephen
Collins). Another rather damaged female taken in trap, same data, in Coll. J.
Brock.

Charaxes pythodoris occidens van Someren
Charaxes pythodoris occidens van Someren, 1963 : 223.

When I described this subspecies from several males taken in the Central African
Republic (French Congo) at Ouesso and Mambili, the female was unknown. It
has now been discovered in the Bangui district of the Central African Republic,
and I am indebted to Mr. J. Plantrou for a colour photograph of this female.

It is very similar in general appearance to the male, having the same strong blue areas on
both wings; the upper blue spots of the discal row larger; the submarginal blue spots in the
hind wing more strongly developed. Fore wing length 52 mm. Size larger than the male,
but the fore wing less incised on outer border; the hind wing more rounded.

CENTRAL AFRICAN REPUBLIC: Bangui, ^1964 (P. Quoniari), in Coll. Plantrou.

Charaxes schoutedeni Ghesquiere

(PI. 29, fig. 257)
Charaxes schoutedeni Ghesquiere; van Someren, 1964 : 220.

By an unfortunate oversight I omitted to give a photograph of the unique type
of this insect when I dealt with this species in some detail, pointing out that it
differed in many important respects from the race of Charaxes smaragdalis leopoldi
Ghesquiere of the southern Congo.

Monsieur Jacques Plantrou has recently sent me photos of leopoldi from Brazza-
ville which he suggests are transitional toward schoutedeni and, though exhibiting

8o V. G. L. VAN SOMEREN

an extension of the blue in the fore wing toward the base of the cell, nevertheless have
the large subapical bluish white spot of the fore wing concave and not convex;
moreover, the submarginal blue spots of the hind wing are small, compared with the
large spots of schoutedeni. Furthermore, the undersides do not agree. Until more
specimens of schoutedeni are obtained from the type locality its exact status cannot
be decided.

Charaxes penricei tanganyikae van Someren
Char axes penricei tanganyikae van Someren, 1966 : 67.

$ form caerulescens forma n. (PI. 29, figs. 255, 256)

Fore wing length 40 mm. Upper side. Similar in pattern to the white-barred form of this
subspecies, but the hind portion of the broad discal bar and the broad discal band on the hind
wing strongly flushed with blue. The fore wing subcostal spots white.

Holotype female. TANZANIA: Kigoma district, Mihumo, vi.ig65 (Japanese
Primate Expedition). To be deposited in B.M. (N.H.).

Charaxes berkeleyi van Someren & Jackson
(Text-figs. lo-n [aedeagus])

Charaxes berkeleyi van Someren & Jackson; 1957 : 5 2 -

Charaxes berkeleyi van Someren & Jackson; van Someren, 1966 : 74.

This species was described from the progeny of two females captured in the Karen
area of Ngong, Kenya. Details of the families are given in the original publication.
Since then, several additional families have been bred from known parents. These
families confirm the general characteristics on which the species was based. The
species was redescribed in detail in the first section of my revisions dealing with the
" Black Charaxes " in 1966 (op. cit.).

Two interesting variations were raised in a recent family and deserve mention.

$ form ngonga forma n.
(PI. 29, figs. 249, 250)

Upperside. Similar to the nominate form in pattern, but the fore wing spots in the discal
row larger and creamy white; the postdiscal spots small and creamy; the hind wing band is
white with strong bluish on borders. Underside. Pattern similar to that of the nominate form
but the discal bars are whitish.

A slight variant has the fore wing discal spots white, but the postdiscal spots are orange.

Range: The distribution of Ch. berkeleyi, as known at the time the species was
redescribed in 1966, was given in some detail; it has now been discovered that the
species occurs on the west side of the Rift Valley and extends in a north-west direc-
tion to the eastern side of Mt. Elgon, but in a distinct form worthy of subspecific
rank, which is now described. (Map i).

REVISIONAL NOTES ON AFRICAN CHARAXES

81

*

\jL.Hannington p/'-Meru
ombe . if"'"... 7 /-"""%'"=

'* / *L\\ V

< >. A.L. Nakaru

\ // >! '^*'%. - /
V "V / 4 \ .^Wfc I Aberdare
. NaivashiliJ~ ,=

" Charaxes berkeley

A C. baileyi.

^ C. evansi evansi.

82 V. G. L. VAN SOMEREN

Charaxes berkeleyi masaba ssp. n.

(PL 29, figs. 252, 253)

FEMALE. Fore wing length 38 mm. Upperside. The pattern differs from the nominate race
by having larger and more strongly orange spots in the fore wing and having a wider discal
band in the hind wing especially in the subcostal region. Underside. Pattern generally similar
to that of the nominate race but bolder ; the ground colour exhibiting the same variation, either
cold greyish or warm brownish. (PI. 29, fig. 252.)

A slight variant may have the fore wing spots less strongly orange and thus very similar to
the nominate, but the hind wing bar is always broader.

Variation. Some specimens exhibit a tendency for the discal and postdiscal spots of the
fore wing to be slightly or strongly rayed or conjoined. There is thus a marked general similarity
to the upperside pattern of some examples of Charaxes etheocles evansi van Someren which flies in
the same area, and they could be easily confused. (PL 29, fig. 253).

MALE. This resembles the male of the nominate race, having well developed blue subcostal
marks in the fore wing and a decided greenish postdiscal line in the hind wing.

It is of interest to note that the genitalia of berkeleyi differs markedly from that of Ch. etheocles
evansi van Someren though the general facies of the upper sides are very similar.

Holotype female. N.W. KENYA: East Mt. Elgon, Teldet Estate, 1963. (Coll.
Lloyd.)

Range: Most numerous in the east Elgon area from 7,000 ft. in the riverine forests
and on the lower slopes at Endebess; also noted in the forest around the Keringet
Dam in the Kapenguria area. It thus has a considerable overlap with Ch. e. evansi,
with which it has hitherto been confused.

These two main aggregates, each with its restricted range, east and west of the
Rift Valley, are however partially bridged by a " cline ", the members of which,
though conforming in the main to the nominate race as regards pattern, have the
fore wing spots richly coloured orange-ochre to tawny orange as in the race masaba.
This cline occupies intermediate territory, west of the Rift in the Visoi-Kilombe
Hill area and along the Elgeyu Escarpment to Kaptagat.

Biological Note. Food plants: there is no doubt that the principal food plant
of the nominate race in the Ngong-Kikuyu-Nyeri areas is Albizia gummifera (Gmel.)
(Leguminosae). This is also the food plant of the cline found in the Visoi-Kilombe
Hill area. But, as mentioned in the original description of the species, 1957, females
do occasionally lay on Ochna insculpta Scleumer (Ochnaceae) , and on Scutia myrtina
Kurz (Rhamnaceae). A few specimens were raised on Scutia on the Kinangop area.

The early stages, larvae and pupa have been described (op. cit., 1957).

Charaxes xiphares kilimensis ssp. n.

(PI. 28, figs. 244-245, 247-248)

This new subspecies of Ch. xiphares belongs to the north-eastern group of the
species which, at the present, includes brevicaudatus Sch., maudei J. & T., desmondi
van Som. and kulal van Som., in which the females exhibit a departure from the
usual female pattern of the more southern group, most of which have a large ochre
patch in the disc of the hind wing. In the northern group the hind wing discal

REVISIONAL NOTES ON AFRICAN CHARAXES 83

patch is white with strong blue scaling on the borders or overall. The fore wing
discal bar is white, while the postdiscal spots, which are pronounced, are ochreous
to orange-ochre.

This new subspecies from western slopes of Mt. Kilimanjaro exhibits characters
which place it intermediate between brevicaudatus of the northern end of Lake
Nyasa and the southern highlands of Tanzania, which has very short tails in both
sexes, and maudei of the Usambara Mts., and possibly the Ulugurus, which has very
long tails in both male and female.

MALE. Fore wing length 49 mm. Upper side. Ground colour deep blue-black with strong
blue sheen in side light, base of costa browner. Discal blue spots rather small, two spots just
beyond end of cell, upper one a streak, spot below larger, spot sub-basal in 4 larger and round,
spot below it in 3 more elongate but smaller with a minute dot beyond, no spot in 2 and spot
in ib small and round, but mark in la an elongate streak widest proximally and tapering toward
postdiscal line. Postdiscal series: two subapical in 8-7 white, spots in 6-4 slightly smaller and
blue, spot in 3 slightly larger, that in ib double. Margin without orange spots except for a
slight indication in ib. Hind wing with a large discal bluish white patch not reaching beyond
5, with strong blue suffusion on the borders, represented at the subcosta by a discrete blue spot;
in the postdiscal row there are two discrete blue spots in the upper sector, but there are confluent
blue marks on the outer border of the discal patch, with black scaling in between. Submarginal
blue spots distinct from 2-6; border of wing with rufous orange lunules from anal angle to 6,
edged with black. Tails black, of about equal length, 4 mm. Underside. Fore wing ground
colour earthy grey with a slight brassy tint, the whole with a satiny sheen in side light except
in mid areas of i and 2, which are dull; the base of the wing olive crossed by narrow black lines
outlined in white; the discal spots represented in olive, proximally edged with black with a
suggestion of white in between; the postdiscal spots: the two upper subapical spots ochreous,
the rest golden olive, the tornal mark olive to greyish distally with conspicuous double half
moon, black in centre, spot in 2 above with slight black distally. Margin with very obscure
olive marks, more obvious in ib and 2. Hind wing ground colour as fore, the sub-base crossed
with zigzag olive line narrowly edged in black; the distal zone more bronzy, with a zigzag
narrow whitish line from costa to 2, a narrow black line through end of cell area. Postdiscal
series of spots from costa to anal angle golden olive, narrowly edged with black proximally,
the mark in the anal angle a long crescent double-edged in black; submarginal spots bluish
grey, those toward the anal angle with black dot distally; marginal lunules golden olive, more
greenish at anal angle; margin black with very narrow white fringe. Tails mostly black with
olive mid line at base.

FEMALE. Fore wing length 55 mm. Upperside. Ground colour purplish brown-black in
basal triangle, blacker on distal half of wing. Disc of wing crossed by a broad white curved
band extending from the costa to hind margin, consisting of four elongate white marks, including
white area on costa, beyond end of cell followed by an off-oval spot sub-basal in 3, the mark in
2 more elongate somewhat pear-shaped, the white mark in ib double, small mark above, that
below a blunted triangle, followed by a long streak in la, the marks in la, ib shaded with violet
scales. Postdiscal spots distinct, a large subcostal subapical mark is whitish and rounded,
spot in 6 smaller, those in 5-2 smaller and orange in colour, mark in ib double and contiguous
with the discal marks. Margin with slight internervular orange marks, double in ib. Hind
wing ground colour black in basal triangle, more purplish black in dark border; disc of wing
with large violet-blue patch, more whitish toward bases of 5-6, with a large whitish quadrate
mark at subcosta in 8; the discal patch itself shaded with violet and on its distal border is a
series of dyslegnic rounded orangish marks from 2-5. Submargin with a series of lilac-blue spots,
distinct from anal angle to 5 ; inner margin with strong orange border of confluent lunules which
shade to olive-green at anal angle ; margin black with narrow white fringe. Tails black, 5 mm.
long. Underside. Ground colour as in the male. Basal triangle of fore wing with series of

84 V. G. L. VAN SOMEREN

strong black lines margined with bluish white, three crossing the cell and two at end of cell,
with a black spot sub-basal in ib and at sub-base with a short bar in 2. The discal white bar
conspicuously outlined proximally in black ; postdiscal spot white in subapical area, then increas-
ingly orange to ib, with the double black spot in ib and 2 strongly marked; admargin with
slight orange lunules strongest in ib. Hind wing ground colour as fore; fine black lines outlined
in white at basel triangle; discal bar represented by a pale ochre-greyish band, dyslegnic on
its outer border but edged internally by a narrow black and white zigzag line; postdiscal series
of rather indistinct ochreous lunules, shaded brownish in lower half, slightly more olive and
more defined above anal angle ; submarginal series of greyish lunules, broadly edged with black
distally, touch the marginal orange-ochre lunules which shade to olive at the anal angle ; extreme
edge black.

Holotype male. TANZANIA: Lower slopes of West Kilimanjaro at Maua Estate,
ix.ig66 (Collector Edmund, for Major Grahame).

Allotype female. Taken in the same locality, 11.1964 (A. F. Brown). Both
deposited in the B.M. (N.H.).

Range: This subspecies is at present known only from the West Kilimanjaro area.

2. THE CHARAXES CEDREATIS-VETULA COMPLEX

When Rothschild & Jordan published their monograph of Char axes in 1898-1900,
the position of many of the black males, so confusingly alike, was left in doubt;
equally so, the many female forms, which could not be associated with any given
males, were left in a state of chaos. Typical of this confusion are the many names,
to which they added several, all lumped under the oldest named form, namely
Charaxes etheocles (Cramer, 1777). They predicted however that in the course of
time, with more extensive collecting and breeding from known parents, this aggregate
would be divided into authentic species and subspecies.

A tentative re-classification of the complex was put forward by van Someren
& Jackson (1952). This was based on considerable field work, breeding from known
females and a re-assessment of the vast amount of material in the B.M. (N.H.),
which now combines the bequests by Rothschild, Joicey & Levick, together with
numerous large collections donated in the past fifty years. Very considerable use
was made of collections in Continental and African museums and in private collec-
tions. During the past decade, particular attention has been paid to verifying
published records by examining the actual specimens, to mapping the distribution
of most of the species within this complex and to breeding them where possible.

Charaxes cedreatis was described by Hewitson in 1874, based on two specimens,
one of which was selected as type and recorded as a male; in fact both are females.
The locality was given as " West Africa ", but Hewitson refers to these specimens
again in 1876, and gives the restricted locality of the type as Fernando Po. Roths-
child however placed cedreatis as one of the numerous female forms of etheocles
Cramer, 1777.

In 1932 I suggested that cedreatis Hewitson, protocedreatis Poulton and lutacea
Rothschild were sexes of one species, based upon the evidence of bred families from
cedreatis females. I did not however completely disassociate it from " etheocles "
s. 1., but this was done by van Someren & Jackson (1952).

REVISIONAL NOTES ON AFRICAN CHARAXES 85

When we reviewed this species, it seemed reasonable from the evidence before
us, to recognize two races, a northern and a southern, based on slight differences
in the male and considerable colour differences in the females. All the northern
females were cedreatis, and the very few from the southern zone were vetula Roths-
child, which we considered to be a form of cedreatis. We made a bad mistake, how-
ever, in recording the type locality of vetula as Ogowe River in Angola instead of in
Gabon! Thus the type locality of vetula lies between the two places mentioned by
Hewitson for cedreatis. It is also unfortunate that we selected as allotype male of
vetula, a specimen taken at Geita, south of Lake Victoria, in Tanzania, where a
female vetula had been captured, instead of one of the many male specimens taken
in Gabon, but these we had placed as cedreatis cedreatis. Thus we overlooked that
the two supposed races overlapped to a considerable extent.

It has now been ascertained that vetula occurs in eastern Nigeria, below the Obudo
Plateau (St. Leger in lit.). Moreover, as a result of recent collecting it has been shown
that lutacea males and cedreatis extend to Katanga and to north-western Zambia,
thus cutting right across the supposed range of vetula. Further to the north-east,
in N.W. Kenya, vetula and cedreatis have been taken in the same forests and in
one instance, in the same trap. It is therefore necessary to re-assess the species in
the light of our present knowledge.

Charaxes cedreatis cedreatis Hewitson
(PL i, figs. 1-6)

Charaxes cedreatis Hewitson, 1874 : 247, $, $ (W. Africa).

Charaxes cedreatis Hewitson; Hewitson, 1876 : (Restricted locality, Fernando Po).

Charaxes carteri Butler, 1881 : 108, $ (Accra).

Charaxes etheocles cedreatis Hewitson; Rothschild, 1900 : 484, $ form.

Charaxes etheocles lutacea Rothschild, 1900 : 485, $ form. (Beni, Congo), syn. n.

Charaxes etheocles cedreatis Hewitson; van Someren & Rogers, 1932.

Charaxes etheocles lutacea Rothschild; van Someren & Rogers, 1932.

Charaxes cedreatis Hewitson; van Someren & Jackson, 1952 : 276.

Charaxes cedreatis vetula Rothschild; van Someren & Jackson, 1952 : 277.

It has been proved by breeding that lutacea Rothschild is the male of cedreatis.
The males of this species exhibit little variation on the upperside; the number and
size of the blue spots on the fore wing may vary slightly. The submarginal white
spots on the hind wing, usually distinct, may be small or even obscured. On the
underside, as in many species of this " black complex ", the tone of the ground colour
may vary, and more particularly, the strength of the black and dark brown pattern
may vary from very strongly to lightly marked. These variations are not related to
any geographical area but may occur throughout the distribution of the species;
they may occur in a single bred family.

MALE. Fore wing length 35-36 mm. Shape slightly incurved at 3-4 on outer edge. Upper-
side. Ground colour blue-black but with strong greenish sheen toward and at base; a small
blue spot at upper end of cell, a larger subcostal spot in discal line ; one or two bluish to white

86 V. G. L. VAN SOMEREN

spots in subapex; marginal glaucous spots usually distinct, largest at the tornal angle at ib.
Hind wing ground colour as fore wing, with slight greenish sheen over disc; there may be a
trace of a greenish wavy postdiscal line most evident opposite the tails; submarginal linear
white marks usually distinct, double at anal angle ; border reddish above upper tail, or reddish
mixed with greenish to anal angle; extreme edge black; tails moderately long, of about equal
length, 4 mm. Underside. Ground colour sepia-brown, slightly browner on border; disc of
fore wing with a satiny bar most evident and expanded at 4-5 and toward costa at end of cell;
basal black marks usually strong up to discal line; postdiscal dark brownish spots often strong,
black at tornus and outlined with greyish. Hind wing ground colour as fore wing; basal black
line fine, divided by a satiny bar in sub-base and a more distinct wavy bar through discal line
followed by a strong dark zone bearing the series of olive and maroon lunules which are usually
strong; the border is more satiny and the submarginal line of whitish linear marks edged with
black distally, strong. The margin is usually strongly reddish to upper tail then olive to anal
angle, which carries two black dots.

Variation. Exhibiting little difference on the upper side, the underside ground colour is
less strongly sepia, more greyish brown, thus the satiny bars are not in strong contrast, and the
dark marks at the base and lower part of the fore wing much reduced. A similar suppression
of pattern is seen in the hind wing. This type of variant is well illustrated in the male we selected
and allocated to vetula of the Geita area. (PI. i, fig. 5.) However, typical lutacea also occurs
with vetula females, as in Gabon.

FEMALE. The type specimen is rather worn and somewhat faded. (PL i, figs. 3, 4.) It
exhibits certain features which are of interest from the point of view of evolution of pattern.
Upper side. Ground colour from base of fore wing including most of the cell, the sub-base of
2, the basal area of ab and most of la, dull tawny olive. A curved white bar crosses the wing
from a white spot in upper part of end of cell followed by larger spots at base of 4, a larger mark
basal in 3 in the discal line, a long mark in 2 and ib representing the fusion of white marks in
discal and postdiscal lines; those in the postdiscal line represented in 35 by small white spots
and larger ones in subapex in 6-7. In addition there are two subcostal white dots in the upper
discal line. Apical half of wing and outer border black, and pale whitish lunules are present
on the margin, most obvious at the tornal angle. The hind wing is mostly tawny olive from
the base to postdiscal line, represented at the costa by two discrete spots; the dark blackish
border is narrow, widest at upper angle; the submarginal row of linear white marks distinct;
marginal border above upper tail reddish then olive to anal angle. Underside. Ground colour
brownish grey with slight olive tone; fore wing basal area with strong black marks. Pattern
of above represented as whitish marks in the curved bar and as pale spots in the apical half
of the wing. Black marks at and above the tornus strong. Hind wing pattern rather subdued;
submarginal pale line fairly clear; margin reddish above upper tail, olive to anal angle. (PI. i,
fig- 40

Although form cedreatis has the widest range, and indeed is the only form recorded
from most of W. Africa, with the exception of type vetula from Gabon and another
specimen from eastern Nigeria, and is the dominant one in the Congo and Uganda,
its overall distribution embraces all the localities from which other female forms
have been described. This precludes the recognition of any subspecies. (Vide
Distributional Map 2.)

Variation, (a) Ground colour and pattern very similar to the type, but curved white bar
more restricted in ib in discal line; no white spots in upper discal line; the postdiscal white
spots restricted to two in subapex. Hind wing as in nominate form. Underside dark olive-
grey-brown; pattern strong.

Variation, (b) very similar to (a) but base of fore wing darker, more brownish olive; the
spots of the white bar more restricted especially in ib to postdiscal line. Hind wing discal
area paler tawny olive. Underside more greyish olive and the pattern less strong.

REVISIONAL NOTES ON AFRICAN CHARAXES

4

-""-' ^

'iVORY COAST /GHANA?
'". o ! o '.

NIGERIA f .S \ CENTRAL t.

''AFRICAN REPUBLIC \ SUDAN

r v

KEY

Charaxes cedreatis.

$ f. cedreatis.
cf f. cedreatis.

Type & paratype of cedreatis

g f. protocedreatis.
CH Type of o" lutacea.

2 f. mexpectata (type & paratype).

2 f vetula, & cedreatis taken together.

^ Type of vetula.

o 2 ' pseudosmaragdalis.

j f. dewitzi.

dj chepalungu.

T GABON; m

\MOYEN B

,'CONGC

CONGO

KATANGA

ANGOLA

ETHIOPIA

. D .

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^A ' so

'SOMALIA

KENYA

V,

?M

MALAWI

ZAMBIA

^,, ' ILWI

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MAP -z.

$ form protocedreatis Poulton

Charaxes etheocles protocedreatis Poulton, 1925 : 556.

Charaxes etheocles protocedreatis Poulton; van Someren & Rogers, 1932.

Somewhat like var. (b) above but curved bar of fore wing distinctly creamy ochre in la-ib; the
discal and postdiscal spots rather obscured ; no marginal spots or if present only faintly indicated
at tornus. Hind wing basal area dark brownish olive but the distal half of the patch paler olive
ochre contrasting with the rather wider black border; submarginal spots rather obscured;
marginal border as usual. Underside ground colour olive greyish; pattern rather obscured
except for curved bar of fore wing. (PL 2, fig. 15.)

88 V. G. L. VAN SOMEREN

$form inexpectata forma n.

This remarkable form would seem to be a development from protocedreatis but exhibits a
pattern unlike any other in cedreatis. Upperside. The fore wing pattern is basically similar
to that of nominate cedreatis, but the basal tawny olive area is darker and more restricted in
ia-2 by a prolongation of the discal pale marks of the curved white bar in these areas basad;
this bar is white in its upper segments then becomes creamy in colour. The distal half of the
the wing is black, but the spots in the upper discal line and those of postdiscal row are large
and distinct though that in 4 is obscured. The marginal spots are mostly small, but those
at the tornal angle, large, especially in rb. The base of the hind wing is tawny olive but
encroached upon by the wide creamy band which is complete from the costa to just above the
anal angle. ; there is some tawny olive scaling on the lower outer border. The black border is
widest at 7-6 then narrows slightly and carries the usual series of submarginal linear white
marks; the marginal border reddish above the upper tail becomes olive to the anal angle;
extreme edge black. Tails long, upper rounded at end, 6 mm. long, lower pointed 5 mm.,
black with median orange-red line. Underside. Ground colour greyish olive, slightly darker
brownish at border of fore wing; basal black marks strong to discal band; the pattern of above
showing up strongly and pale cream in colour ; postdiscal row of dark spots distinct from subapex
to tornal angle where they are black. Hind wing ground colour greyish olive at base crossed
by fine black lines; discal band strongly marked but less wide than above tapering from mid
point to above the anal angle, bounded distally by a strong row of olive-maroon lunules ; border
slightly darker olive-grey with submarginal row of linear white marks strong carrying black
dots in lower segment in region of tails; marginal border narrow, reddish above upper tail then
olive to anal angle.

Holotype female. TANZANIA: N.E. of Lake Tanganyika, Kigoma district,
Mukuyu Forest, 5,200 ft., ^.1965. Captured by Major I. Grahame's collector.
To be deposited in the B.M. (N.H.). (PL 2, figs. 13, 14.)

A second specimen has the upper spots of the fore wing white; and those of the
curved bar slightly less ochreous. The hind wing band is wider and slightly more
whitish. Taken at Muhimo, Kigoma district, N.E. of Lake Tanganyika, 1.1965
(Japanese Primate Expedition) (PL i, figs. 7, 8.) The associated males are typical
lutacea.

Two examples of this new form have recently been taken in the Ivory Coast by
A. Gallay, and a third specimen by the late Mr. T. H. E. Jackson's collector in the
same area.

$ form vetula Rothschild

Char axes etheodes f. vetula Rothschild in Rothschild & Jordan, 1900 : 488.

The type came from Ogowe River, Gabon, and is figured in Rothschild's mono-
graph. This type locality is thus within the range of nominate cedreatis Hewitson.

Upperside. Pattern is very similar to cedreatis as regards the curved fore wing bar, but the
basal area is brownish black; the distal half of the wing black, with an indication of a white
subcostal spot in the discal line, two white spots in subapex while those of the postdiscal series
are faintly indicated. The inner ends of the white marks in la-ib are bluish; marginal pale
spots faint, most obvious at tornal angle. Hind wing basal area brownish black; disc of wing
with a large blue patch tapering toward the anal angle, represented at subcosta by two blue
spots one in discal and one in postdiscal line; border black, widest at upper angle; submarginal
row of linear white marks very distinct; marginal border reddish to between tails then olive

REVISIONAL NOTES ON AFRICAN CHARAXES 89

to anal angle. Tails moderately long, 5 and 4 mm. Underside. Ground colour greyish olive
or pale brownish, darker on border of fore wing. Basal black marks strong or weak; white
curved bar distinct; a satiny quadrate mark sometimes present at subapex; submarginal series
of dark spots are black at tornus but fading out toward the apex. Hind wing ground colour
greyish olive or brownish, pattern rather obscured except in lower part of postdiscal line;
submarginal pale line usually distinct ; marginal border mostly brick-red but olive at anal angle.
(PI. 2, figs. 9, 10, Type. Gabon.)

Variation, (a) Upperside. Very similar to nominate specimen but some indication of dark
ground colour invading the white marks in la-ib in the line of junction of the discal and post-
discal spots; upper subcostal spots in discal row free; spots of upper postdiscal row small but
distinct. Underside. With a rather stronger pattern, division of discal and postdiscal white
marks ib-2 clearly indicated; submarginal marks in fore wing not strong. Hind wing with
satiny bar in discal line ; lunules large in postdiscal zone ; submarginal row of white spots distinct.
Geita, Tanzania.

Variation, (b) Upperside. Basal area blacker with distinct blue sheen, particularly toward
the inner edge of the curved white bar, the blue colour suffusing most of the marks in la-ib;
subcostal mark in discal row faint or absent, but postdiscal spots are blue; marginal spots
glaucous but faint except for double mark at tornal angle ib. Hind wing ground colour at
base, black, black also along costal border thus restricting the upper border of the strongly
blue patch in the disc; submarginal row of linear marks bluish white, more strongly blue at
anal angle; marginal border a mixture of reddish and glaucous to upper tail then olive to anal
angle. Tails shorter: 4mm. Underside. Ground colour olive greyish, brownish on border;
pattern rather obscured except for white bar in fore wing and white submarginal spots in hind
wing. (PI. 2, fig. n, Kakamega, Kenya.) (Rydon Coll.).

Variation, (c) Upperside. Very similar to (b) with the same strong blue invading the
proximal two thirds of bar spots in la-ib. The hind wing blue band goes right through to the
costa and crosses the inner fold above the anal angle. Underside. Ground colour and pattern
as in (b). (PI. 2, fig. 17, Visoi Gap, Kenya.) (Bailey Coll.)

$ form pseudosmaragdalis van Someren & Jackson
Charaxes cedreatis vetula $ f. pseudosmaragdalis van Someren & Jackson, 1957 : 50.

At present known only from a single specimen taken in the Djelo-Binza area of
Leopold ville, lower Congo. It bears a superficial resemblance to the male of
Charaxes smaragdalis smaragdalis Butler.

Upperside. Base of fore wing brownish black with a strong greenish blue suffusion up to the
discal line, with a stronger blue sheen in the cell. Discal spots large, two upper elongate, bluish
white, a rounded mark sub-basal in 4, that in 3 more quadrate and set out a little, followed by
and conjoined to larger blue mark in 2, and a larger mark in la-ib conjoined to the blue spot
in postdiscal row, the remaining postdiscal spots in 2-5 in a slight curve terminate in the two
larger subapical spots which are white. Margin with indication of diffuse glaucous marks.
Hind wing basal area as fore wing merging into the blue discal patch which extends from the
costa to above the anal angle. Border black with distinct linear white marks; margin with
reddish scales to upper tail then olive to anal angle. Underside. Light rufescent brownish;
black lines thin to discal zone ; pattern of Upperside represented by slight greyish brown marks,
the lower postdiscal ones margined in black, proximally in ib-2 where the tornal marks are
black. Hind wing ground colour as fore wing, a pale bar present in the discal line finely outlined
in black proximally; the postdiscal row of buff and maroon lunules strong; whitish marks in
submargin strong ; border reddish above upper tail then olive to anal angle carrying black dots
in region of tails. (PI. 3, figs. 18, 19, deposited in Central African Museum, Tervuren, Belgium.)

go V. G. L. VAN SOMEREN

$ form dewitzi Butler

Charaxes alladinis Dewitz, 1887 : fig. 8.
Charaxes dewitzi Butler, 1895 : 255.

The type of this insect is lost. It was described as the male of alladinis Butler
by Dewitz, but was subsequently shown to be a female and was renamed dewitzi
by Butler. It is figured in colour in Dewitz' original paper. It appears to be nearest
to the female form pseudosmaragdalis , differing mainly in the absence of the post-
discal spots in the fore wing other than the two subapical ones; otherwise, the descrip-
tions agree very well. The specimen came from Pungo Andongo in northern Angola.
(PI. 3, fig. 20, [photograph of original figure.])

Charaxes chepalungu sp. n.

(PI. 3, figs. 21-25)

Charaxes cedreatis vetula Rothschild; van Someren & Jackson, 1952 : 278.

This species was briefly referred to, with a query, in van Someren & Jackson
(1952), under the heading Charaxes cedreatis vetula. Two females and several males
were taken in the Chepalungu Forest, lower Sotik in September, 1949 and several
males in the Mara district. Subsequently, an interesting gynandromorph, male
on right side, female on left, was taken by the late R. T. Evans at the edge of the
Chepalungu Forest. The male half agrees exactly with the males taken previously,
and the female side with the previously known females, tentatively placed to vetula.
Since these males do not agree with any form of lutacea, the male of cedreatis vetula,
it becomes desirable and necessary to disassociate the Chepalungu insect from
cedreatis ; moreover, cedreatis <$ lutacea occurs in the Mara area.

MALE. Fore wing length 31-33 mm.; shape of fore wing only slightly incurved on outer
border, thus somewhat resembling Charaxes ethalion. Upperside. Fore wing ground colour
velvety black with slight indication of glaucous spots on margin between veins; blue subcostal
spots very small and limited to one beyond end of cell and one in subapex, or these spots may
be obscured. Hind wing ground colour velvety black; submarginal row of spots whitish, very
small and may be obscured ; marginal border greenish or with very slight maroon scaling above
upper tail, all rather obscured Tails short, robust, upper 3 mm. lower 4 mm. Margin at
veins slightly dentate. Underside. Fore wing ground colour greyish brown basally, more
brownish on distal third; slight satiny bars in discal and postdiscal lines; basal black lines thin
or may be strongly indicated in lower half of wing especially at tornus, with submarginal dark
marks diminishing toward subapex. Hind wing ground colour greyish brown with a satiny
bar in discal line, followed by a darker brownish bar distal to which the postdiscal zigzag band
of lunate olive and maroon may be strong, or may be mainly olive in colour. Submarginal
row of linear marks lilac or greyish with black dots present in region of tails, double at anal
angle; margin narrowly maroon above upper tail then olive to anal angle. There is some
variation in the strength of the underside pattern.

FEMALE. Fore wing length 35-36 mm. Upperside. Ground colour basal area blackish
brown, distal half black, separated by an oblique white bar made up as follows: a small white
spot at upper end of cell, a quadrate mark beyond end of cell followed by an elongate triangular,
at base of 3, a long white mark in 2 extending to postdiscal line, the marks in la-ib blue shading
to white in postdiscal line; the postdiscal spots small and obscured in 3-5, but the subapical

REVISIONAL NOTES ON AFRICAN CHARAXES 91

punctiform and white; one or two obscure whitish marks midway in 6 and subcosta. Margin
with obscure internervular glaucous marks. Hind wing brownish black at base followed by a
large irregular blue discal patch, tapering toward the anal angle, represented in subcosta by
blue marks in postdiscal line; border black, carrying distinct linear white marks on outer edge;
margin reddish from above upper tail to base of lower, anal angle olive. Tails rather thin, upper
5 mm., lower 3 mm., black edged with maroon-olive mid line. Underside. Fore wing ground
colour clayish brown with white pattern of above well represented and with pale satiny spots in
the postdiscal row ; black tornal spots strong but black lines toward base rather fine. Hind wing
ground colour clayish brown with fine black lines at basal area ; postdiscal zigzag line of ochreous
olive-maroon lunules fairly well marked.

Variation. In one specimen, the white marks in the fore wing rather more extended and the
hind wing blue patch more extended basad over cell.

Holotype male. N.W. KENYA: lower Sotik area, Chepalungu, viii.i95o (van
Somereri) .

Allotype female. Same locality, ix.ig46 (van Someren). Deposited in B.M.

(N.H.).

Range: The topotypical series of ten males and three females were taken in the
Chepalungu Forest in lower Sotik and in the riverine forests along the Mara River.
Identical males have been captured in the Serengeti Game Park across the border
in Tanzania. Two males which appear to belong to this species are recorded from
Mukuyu in the Kigoma area, Tanzania. There is thus an overlap with both
cedreatis and ethalion ssp. The species has not been bred, but the capture of the
gynandromorph verifies the association of males and females. (Map 2.)

Males were dissected by Mr. Bennett of the B.M. (N.H.) who reports that they differ
appreciably from ethalion, and are more sinuate than lutacea (Text-fig. 20 [aedeagus])
with the toothed lobe placed less toward the distal end of the aedeagus.

SYSTEMATIC LIST

Charaxes cedreatis Hewitson

Charaxes cedreatis cedreatis Hewitson, 1874. Type locality: Fernando Po. Ranges
of various forms :

Synonym: lutacea Rothschild, 1900.
9 f . cedreatis and variants :

W. Africa: Ivory Coast, Ghana, French Guinea, Fernando Po, French

Cameroons, N. Angola, Congo central, Kasai, Katanga.
Uganda: S.W., W., C. and E. Uganda.

Kenya: N.W. Kenya, Trans Nzoia, Kaimosi, S. Kavirondo, Nandi,
Lumbwa, Ravine, Lembus, Pekera Gorge, Visoi Gap-Kilombe Hill;
all west of the Rift Valley.
Tanzania: Usukuma, Serengeti, Geita, south of Lake Victoria; Bukoba

and south of the Kagera River; N.E. Lake Tanganyika, Kigoma.
Zambia : Mwinilunga area.

$ f. protocedreatis Poulton, 1925. Type locality: Uganda, Jinja.
Uganda: Central and eastern; Jinja.
Tanzania: Serengeti, Orange River forest.

ENTOM. 23, 4. 9

Q 2 V. G. L. VAN SOMEREN

$ f. inexpectata forma n. Type locality: Tanzania, Kigoma, Mukuyu Forest.

Tanzania: Kigoma area, Mukuyu Forest, and Muhimo.
9 f. vetula Rothschild, 1900. Type locality: Gabon, Ogowe River.

W. Africa: Gabon, Ogowe R., Eastern Nigeria below Obudo.

Tanzania: Geita, S.W. Lake Victoria, (i specimen.)

Uganda: Rutenga, Kigezi, S.W. Uganda, (i specimen.)

Kenya: Kaimosi Forest, N. Kavirondo; Kilombe Hill, Visoi Gap,

(4 specimens.)
$ f. pseudosmaragdalis van Someren & Jackson, 1957. Unique.

Type locality: W. Congo, Leopoldville district, Djelo Binza.
$ f. dewitzi Butler, 1895. Var. of above?

3* Type locality: N.W. Angola, Pungo Andongo.

Charaxes chepalungu sp. n.

Charaxes chepalungu sp. n. Type locality: Kenya, Chepalungu Forest, Lower
Sotik. Range: Kenya, Chepalungu, Mara River and Tanzania, Serengeti.

3. CHARAXES VIRILIS ROTHSCHILD, C. KHEILI STAUDINGER,
C. NORTHCOTTI ROTHSCHILD and C. MAFUGA VAN SOMEREN

Charaxes virilis van Someren & Jackson

Charaxes etheocles $ f. virilis Rothschild; van Someren & Jackson, 1900 : 487.

Charaxes etheocles $ f. virilis Rothschild; Aurivillius in Seitz, 1912 : 136.

Charaxes etheocles $ f. virilis Rothschild; Poulton, 1918 : 71-73.

Charaxes etheocles $ f. lenis Jordan, 1929 : 483. Type loc., Uganda, Mulange, Mabira Forest.

Charaxes virilis van Someren & Jackson, 1952 : 283-284.

The type of virilis is a female and was described as a female form of etheocles by
Rothschild in 1900. Poulton (1918) refers to a series of specimens bred from larvae
found feeding on Adenthera pavonina L. (Mimosoidea), (native to Tropical Asia, and
introduced into other parts of the Tropics) at Ibadan, S. Nigeria by Lamborn and
Farquharson. The series contained four males and three females and Poulton
makes special reference to the females which are male-like, though somewhat
variable. No particular mention is made of the males, which is somewhat surprising
since the male of virilis had not been described. Dr. Jordan examined the material
and agreed that the females were nearer to virilis than to alladinis Butler. The
males were apparently considered to be just etheocles (Cramer). In 1929 Dr. Jordan
described etheocles male form lenis (not leonis as given by Wallace Peters, 1952).
There were three specimens taken in the Mabira Forest, Uganda. He did not asso-
ciate these males with female virilis, but he compared them with " etheocles " <$
form cytila which I now know to be the male of manica Trimen, 1894. There is no
doubt that lenis is the male of virilis, which is a species distinct from etheocles (Cramer)
(s. str.). Whether virilis can be divided into western and eastern geographical
races will now be discussed.

Western aggregate, topotypical material: Ghana to Cameroon and Congo.

REVISIONAL NOTES ON AFRICAN CHARAXES 93

MALE. Fore wing length 33-35 mm. Shape somewhat like ethalion, the outer border of
fore wing only slightly incurved at 3-4. Upper side. Ground colour velvety black with a strong
overall blue sheen, more especially over the cell. Blue spots in the subcostal area rather variable,
but there is usually one at the upper end of cell, one beyond in discal line, no spots in subapex ;
marginal glaucous lunules ill-defined but often contiguous at the hind angle. Hind wing black,
sometimes with a very faint indication of a greenish or bluish wavy line, strongest opposite to
tails ; submarginal row of linear marks, bluish white or blue, complete ; border reddish or bluish
green above upper tail then more olive at anal angle; tails short and thin, upper sometimes
stumpy 5-4 mm., lower 3-4 mm. Underside. Somewhat variable, usually clay-coloured with
a purplish brown tone overall; satiny bars more greyish; black dots and lines in basal triangle
of fore wing distinct, but rather obliterated in distal half, but tornal black mark strong. Hind
wing ground colour as fore wing, basal black marks fine and indistinct; sub-basal and discal
satiny bars fairly distinct ; postdiscal wavy line of olive and maroon lunules defined but may be
somewhat obscured in the general brownish tone; submarginal line of lilac marks with black
dots distally in region of tails fairly strong; marginal border narrow, slightly reddish above
upper tail then olive to anal angle.

Variation. Some specimens may have the fore wing devoid of blue spots in the costal area
and the marginal lunules may be very small. The underside ground colour more greyish brown
and on the hind wing the satiny bars may be strong. An aberration is figured in which the
subcostal spot in the discal line is enlarged and elongate, represented by two long, contiguous,
blue streaks; the ground colour more purplish brown. (PI. 4, fig. 32.)

Eastern aggregate. Uganda, from Budongo Forest to Kenya border.

MALE. Eastern males are, on the whole, more compact looking than their western counter-
parts, due mainly to the fact that the outer margin of the fore wing is less incurved, the wing
thus looks less elongate. Upperside. The subcostal blue marks vary in number and size;
the marginal glaucous marks are usually obscured. On the hind wing the submarginal whitish
marks are usually distinct and well developed. Underside. Pattern variable, weak, moderate
or fairly strong but always with a strong vinaceous or rusty purple bloom overall. The
postdiscal wavy line of lunules well marked.

There is thus little to distinguish western from eastern males on the upper side. In shape,
eastern males are more compact and the underside is more rusty, or cinnamon-brownish with
a strong vinaceous tone overall. (PL 4, figs. 34-37.)

Western aggregate.

FEMALES. The type female on the upperside is blue-black with steel-blue reflections especially
on the hind wing. Fore wing marks limited to one blue subcostal spot in discal line; margin
with large glaucous lunules. Hind wing with a indication of obscured greenish blue row of spots
in the postdiscal line ; submarginal white spots large and distinct. Border ochre reddish above
upper tail, more olive to anal angle; edge black; tails thin, upper 6 mm., lower 5 mm. (PI. 4,
figs. 26, 27.)

Variation. Topotypical females exhibit a considerable degree of variation in the upper side
markings. They may be as in the type or (a) there may be obscure marks in 2-3 in the discal
line and obscure marks in the postdiscal row from sub-apex to ib; (b) the spots on the fore wing
may be purplish, though obscured in the postdiscal line and represented in the hind wing in the
same zone by purplish marks; (c) the fore wing spots may be as in (b) but discal and postdiscal
spots slightly whitish.

The undersides in all these forms are similar, clayish brown, slightly darker brownish between
the discal and postdiscal zones and in the curve of the wing. The pattern of black marks usually
thin and faint except for the black tornal mark and one more proximad in the same area, the
tornal mark with a greyish surround. The hind wing ground colour as fore with very faint
marks in the basal half; the postdiscal row of lunules obscured, but the submarginal series of
pale marks to the anal angle fairly strong. The border reddish to the upper tail then olive to the

94 V. G. L. VAN SOMEREN

anal angle; the postdiscal mark above anal angle strong with an irregular whitish mark above it.
Females from the Moyen Congo area exhibit the same variations on the upper side but the
pattern of the underside is stronger. (PI. 4, fig. 31.)

Eastern aggregate. Uganda.

FEMALES. These exhibit the same character as regard shape as noted in the males. The
markings on the upperside of the fore wing as in western examples, but less strong, but the
marginal glaucous lunules are stronger; the submarginal white marks on the hind wing larger
and stronger. The underside pattern is stronger and the ground colour richer vinaceous
cinnamon-brown. (PI. 4, figs. 28, 29.)

Whether or not the combined characters of males and females of the eastern
aggregate warrant subspecific ranking is a matter of opinion; if so, the name lenis
Jordan, applied to the eastern male (Mabira Forest, Uganda) is available, the
western aggregate being virilis.

The species has not been bred in the east.

Range: The Western aggregate: Ivory Coast Ghana to Nigeria, Cameroon,
Central African Republic, Moyen Congo, Central Congo eastward to Semliki River
and south to upper Katanga.

The Eastern aggregate : Uganda, from the Budongo Forest in the west to Busogo
in the east to the Busia area on Kenya border. (Map 3.).

Charaxes kheili Staudinger
(PL 5, figs. 38, 39, 40, 43)

Charaxes kheili Staudinger, 1896, Iris 9 : 216.
Charaxes kheili Staudinger; Rothschild, 1900 : 473.
Charaxes kheili Staudinger ; Jackson, 1957 : 66-67. (partim).

This species is still rare in collections, and little is known about it, the female
being still unknown.* The type is in the Berlin Museum and was kindly forwarded
to the B.M. (N.H.) in 1956 for examination and dissection of genitalia when the late
Mr. T. H. E. Jackson was working on the butterflies of the Kigezi country, Uganda,
more particularly on the species of black males of the " etheodes group " which
exhibited characters very similar to the description of kheili Staudinger. Jackson
came to the conclusion that these black males were not kheili, and referred them to
cedreatis vetula on the evidence of a vetula female taken in the same area. I have
now separated these males under the name Charaxes mafuga sp. n.

The type of kheili was taken in the Njam-Njam savanna country in the north
of what is now the Central African Republic. I have been able to locate a second
specimen taken in the same general area (unfortunately without exact data label)
in the Central African Museum, Tervuren, Belgium and this has been sent to me by
Dr. Berger for critical examination. Dr. Berger informs me that the Museum has
three other male specimens taken at Sassa & Yakoma in the Njam-Njam country,
which agree with the specimen sent.

* The female has now been taken and will be described by MM. J. Plantrou & Drage.

REVISIONAL NOTES ON AFRICAN CHARAXES

95

MALE (type). Fore wing length 36 mm. Upper side. Ground colour black with blue sheen
at base and over cell; pattern: a blue spot at upper cell-end, two spots beyond, the subcostal
one large and whitish the lower small and bluish; three subapical spots in line, upper ones
whitish, in postdiscal row with spots in 4-3 set in a little, that in 2 set out a little followed by an
elongate double mark in ib and a transverse mark in la, on hind margin; marginal glaucous
marks complete and distint. Hind wing ground colour black, with a strong greenish blue
postdiscal irregular band extending from above anal angle and terminating in a small spot in
6; submarginal linear marks bluish and well marked; border bluish to lower tail then olive at
anal angle; extreme edge black with slight white fringe. Tails rather thin, of about equal
length, 5 mm. Underside. Greyish brown with darker brown zone between paler discal line
and postdiscal series of paler spots, both proximally outlined in black ; submarginal dark marks,
black at tornus become less distinct up to subapex. Basal lines moderate. Some satiny

C. kheili.

C. kheili sbsp
C. northcotti.
C. mafuga.

MAP 3.

96 V. G. L. VAN SOMEREN

sheen over discal and postdiscal zones. Hind wing, ground colour as fore wing but more uniform,
as basal lines are fine, but satiny bar in sub-base and in discal line moderately strong ; postdiscal
irregular line of lunate marks, ochre-olive and maroon well marked; submarginal row of linear
marks strongest in region of tails; border reddish to upper tail then olive to anal angle, which
carries double black spots. (PI. 5, figs. 38, 39.)

Variation. Fore wing. Upper side. Subcostal marks not so marked; the postdiscal row
not so strong from 2-4, but there is an indication of a blue mark in discal line sub-basal in 3;
marginal glaucous marks smaller. Hind wing postdiscal blue-green band strong as in type
but extending to 5 only. Underside. Not so strongly marked and more uniform on the hind
wing.

Range: The known range of this species appears to be the savanna country to
the north of the Central African Republic in the region of the Shari-Ubanji River.
How far south it extends is uncertain, but there is a record of the insect from north
of Bangui (J. Plantrou, in lit.). (Map 3).

I have before me an interesting specimen of a male Charaxes taken by one of Mr.
Jackson's collectors at Metu, West Madi, Nile Province, Uganda, which closely
resembles the variety of Charaxes kheili mentioned above as regards the upperside
of both wings with the exception that the marginal glaucous marks of the fore wing
are larger and more diffuse. The underside however, though resembling kheili in
general pattern, is more strongly variegated, the markings being stronger, thus
having a marked resemblance to this respect to the underside of a boldly marked
variation of Charaxes northcotti from the Bouar area of Central African Republic
(Cf. fig.) described later. (PI. 5, figs. 40, 43.)

It is impossible to gauge the degree of variation of the underside of kheili owing
to paucity of material.

Charaxes northcotti Rothschild
(PL 5, figs. 41, 42, 46)

Charaxes northcotti Rothschild, 1899, Nov. Zool. 6 : 171, PI. 8, fig. 5.
Charaxes northcotti Rothschild ; Peters, 1952 : 54.

There appears to be some justification for uniting northcotti with kheili, but the
evidence is not entirely conclusive. The ranges of the two do not seem to overlap,
but adequate material of each is sadly lacking, and only males are known.

MALE. Fore wing length 36-38 mm. Upperside. Ground colour black with a slight greenish
sheen over base of fore wing. The pattern of the fore wing is very similar to that of kheili,
but the blue markings in the postdiscal series are larger, and in the discal row there is a large
spot sub-basal in 3 and a smaller one below in 2. In the hind wing the postdiscal greenish blue
band is wider than in kheili, and the outer margin nearer to the submarginal row of blue linear
marks and on the inner border it extends proximad toward the cell so that the widest portion
is in area 45, but the width is somewhat variable. The border of the wing is blue-green to
lower tail then olive at the anal angle. Tails rather short and sharply tapering, upper 4 mm.,
lower 3-5 mm. Underside. Although the dark pattern is essentially similar to that of kheili,
the black marks are finer and the satiny bars less obvious due to an overall darker more brownish
tone ; this applies to both wings.

This general description is based on the type and a paratype which came from the " Gold
Coast, Gambaga, near the Volta River ". (PI. 5, figs. 41, 42.)

REVISIONAL NOTES ON AFRICAN CHARAXES 97

Variation. An interesting male specimen which would appear to be a variant comes from the
Boar area of the Central African Republic. It differs from the nominate form in having a
much bolder pattern of blue spots on the upperside of the fore wing, with indications of additional
spots in ib; on the hind wing the postdiscal blue patch is wider throughout, especially in 2-4;
the submarginal spots are bolder and the border is wider. On the underside, the pattern is
bolder, the fore wing discal and postdiscal spots more defined and the basal black marks stronger.
The hind wing pattern is also bolder giving the whole underside a " spotted " appearance.
(PI. 5, fig. 46.) More material from this area may show the characters to be constant and the
aggregate may constitute a good subspecies.

Range: The nominate form is known from the Volta River area and Kumasi
of the Ghana Republic (Gold Coast) and extends eastward into Nigeria, N.E. of
Enugu. (Map 3).

Charaxes mafuga sp. n.

(PI. 5, figs. 44, 45, Text-fig. 9 [aedeagus])
Charaxes cedreatis vetula Rothschild; Jackson, 1957 : 66.

This insect is represented by a long series of males taken in the high rain forests
of Mafuga-Rutenga, 7,000-8,500 ft., Kigezi district of Uganda.

Its true position has remained a puzzle; no authentic females can be associated
with it. On the evidence of one very battered female, identifiable as vetula Roths-
child, taken in the Mafuga area, Jackson associated it with these males (tentatively,
it is true) but he placed the males to cedreatis vetula Rothschild. However, these
Mafuga males do not agree with males of cedreatis which Jackson placed to cedreatis
vetula of the Geita area, Tanzania, nor with cedreatis males (lutacea Rothschild)
from northern Angola, or elsewhere. I have already shown that lutacea males are
variable, above and below and have expressed the opinion that cedreatis vetula
cannot be upheld as a subspecies, because the form vetula crops up here and there
within the range of the species, from Nigeria to N.W. Kenya.

There is a superficial resemblance on the upper side of the Mafuga insect and
C. kheili Staudinger of the savanna country to the north of the Central African Republic,
in the Njam-Njam country. (Cf. PI. 5, fig. 38). The habitats of the two are thus
totally different. The shape of the two are unlike, kheili having a more pointed
acuminate fore wing. No significant difference can be noted in the genitalia
according to Bennett.

It seems advisable to consider the Mafuga insect distinct; it is certainly not
conspecific with cedreatis (male lutacea} .

MALE. Fore wing length 44-46 mm. Shape of fore wing broad and slightly incised at 3-4
on outer edge though the tornal angle projects slightly at vein 2. Upperside. Ground colour
blue-black with a slight overall greeny tinge to the hind wing. Fore wing with an obscure
blue spot at end of cell followed by two distinct blue spots sub-basal in 5-6 the upper one large ;
three subapical spots in a row followed by a complete row of less distinct blue postdiscal spots,
largest and double in ib; margin with distinct glaucous marks, double at tornus. Hind wing
with a row of postdiscal greenish contiguous lunules from above anal angle to 5 ; complete row
of submarginal linear marks bluish green, whiter at upper angle, distinct, double at anal angle ;
marginal border dark maroon above upper tail then olive to anal angle; edge black. Tails
rather short, upper 4 mm. lower 5 mm. black-edged, with greenish mid line. Underside.

98 V. G. L. VAN SOMEREN

Ground colour sepia-brown, darker in interspaces, with strong satiny bars, quadrate at end of
cell, a zigzag line in discal zone, with a more continuous zone in postdiscal zone, widest in sub-
apex and most distinct, almost whitish in ib opposite and adjoining the strong black tornal
mark, with more brownish marks to subapex; margin of wing shaded brownish in the curve
but greyish at tornus. Hind wing ground colour slightly stronger brownish but satiny bars
strong, especially in sub-base and through the discal line ; postdiscal row of ochreous olive and
maroon lunules, outlined proximally in black, strongly marked; submarginal row of linear
whitish lilac marks strong; marginal border dark reddish to upper tail then olive to anal angle.
The strength of the variegated underside pattern depends on the distinctness of the satiny bars.

Holotype male. S.W. UGANDA: Kigezi County, at Rutenga-Mafuga Forest,
7,000 ft., vi.1951 (vanSomeren). InB.M. (N.H.)
Paratype male. Same data.

The female is not yet known, with certainty.

Range: The species was particularly plentiful in the high rain forests of Mafuga
and Rutenga, in Kigezi; and the males came readily to ground bait, particularly
excrement of carnivores, less so to banana smeared on tree trunks. (We had not
adopted traps as a method of capture in those early days.) (Map 3) .

SYSTEMATIC LIST

Charaxes virilis van Someren & Jackson

Charaxes etheocles $ f. virilis Rothschild, 1900. Type locality: "West Africa".
Range : Western aggregate ; Ivory Coast, Ghana to Nigeria, Cameroon, Central
African Republic, Moyen Congo, C. Congo eastwards to Semliki River and
south to Upper Katanga. Eastern aggregate; Uganda, from Budongo
Forest in west to Busogo in east to Kenya border (Busia area).

Charaxes kheili Staudinger

Charaxes kheili Staudinger, 1896. Type locality: Central African Republic, Njam-
Njam. Range: Central African Republic in Shari-Ubanji River area and
possibly north of Bangui.

Charaxes northcotti Rothschild

Charaxes northcotti Rothschild, 1899. Type locality: Ghana, Gambaga. Range:
Ghana, Volta River and Kumasi to Nigeria, north-east of Enugu.

Charaxes mafuga sp. n.

Charaxes mafuga sp. n. Type locality: S.W. Uganda, Kigezi County, Rutenga
Forest. Range : S.W. Uganda.

4. THE CHARAXES ETHEOCLES COMPLEX

The bewildering number of African Charaxes with black males exhibiting hardly
any or only slight differences, has given rise to a great deal of confusion. Before
Rothschild and Jordan (1896-1900) produced their monograph of Charaxes, there
was little or no evidence supporting +he allocation of many of the equally numerous

REVISIONAL NOTES ON AFRICAN CHARAXES 99

females with any given males, or vice versa. The confused state confronting them
can be gauged by the long list of described males and females under the heading
" Char axes etheocles " to which were added many more names, both males and females,
as members of the complex.

During the past fifty years some attempt has been made to breed the various
" suspected species " from known females. A study of these families has provided
evidence that certain males are definitely associated with certain females, and the
degree of constancy in this respect, coupled with the constancy of external characters,
supported the view that several distinct species were involved in this " complex ".
Characters of the larvae in various stages and food preference were also noted.

In 1952 an attempt was made to disentangle the muddle, utilizing as evidence
the breeding results obtained up to that date, coupled with close field observations
relative to distribution, general ecology, etc. (van Someren & Jackson, 1952 and

1957).

Additional data has come to hand during the past decade which necessitates a

further modification. Some of the more obvious species in the " complex " have
been dealt with previously in the present series of Revisions; many of the names
listed by Rothschild as " forms of etheocles " have been elevated to specific or
subspecific rank.

The present paper deals with Char axes etheocles (Cramer) (sens, str.) on a Pan-
African basis. The species ranges from Sierra Leone through the Congo to N.W.
Kenya; it also occurs in N. Angola, eastward through Katanga to N.W. Zambia
and western Tanzania. It appears to be divisable into reasonably well defined
subspecies, based mainly on the characters of the female sex in each aggregate;
the males being hardly distinguishable and variable on the underside.

Charaxes etheocles (Cramer)

[Papilio] etheocles Cramer, 1777 : 34, pi. 119.

Nymphalis etheocles Cramer ; Godart, 1823 : 355.

Charaxes etheocles (Cramer) Thon, 1837 : 74, pi. 37, figs. 547, 548 (<j>).

Charaxes etheocles (Cramer); Butler, 1869 : 52.

Charaxes alladinis Butler; Kirby, 1871 : 269, n. 23 ($).

Charaxes etheocles (Cramer) ; Trimen, 1891 : 80.

Charaxes hollandi Butler, 1893 : 266.

Charaxes ephyra ab. catochrous Staudinger, 1896 : 218.

Charaxes etheocles 9 f. regalis Rothschild, 1900 : 486.

Charaxes etheocles $ f. fulgens Rothschild, 1900 : 487.

Only important relevant references are cited; for further references vide
Rothschild & Jordan, 1900 : 481-482, and Bryk, 1939 : 375-542.

Charaxes etheocles etheocles (Cramer)

(Pis. 6, 7; Text-fig, i [aedeagus])

REGION i. W. AFRICA, SIERRA LEONE EAST TO CROSS RIVER, NIGERIA.
The type of the species is the female form etheocles (Cramer), described from
Sierra Leone. No type specimen now exists, but the figure given by Cramer

ioo V. G. L. VAN SOMEREN

though not entirely satisfactory, must be taken as representing the white-barred
female of Sierra Leone. (PI. 7, figs. 55, 56. [Photo of original figure].)

FEMALE. Fore wing length 35-37 mm. margin of wing slightly incurved at 3-4. Upperside.
Fore wing spots well separated and complete; a large spot in the cell; discal spots including a
dot at base of 4 creamy to 2, larger marks in la-ib white, blue-tinged distally; postdiscal spots
larger, ochre in colour, extending from subapex to 2; margin with faint indication of greyish
marks, most obvious at tornus. Ground colour brownish black with blue tinge at base. Hind
wing ground colour brownish black, blacker on border; discal band fairly narrow, almost straight
on outer border, more irregular on inner, generally white with slight blue on inner border;
tapering rapidly to inner border above anal angle where it crosses the inner fold as a partially
separated mark. Submarginal series of linear whitish marks strong; marginal border strongly
reddish to lower tail then olive to anal angle; margin black; tails moderately long, upper 6 mm.
lower 5 mm., black with red midline.

Underside. Ground colour light brown, slightly more rusty on outer border of fore wing;
basal black marks fairly strong ; pattern of above well marked, white, slightly buffish in subapical
area; submarginal ocelli weak except at tornus and area above where the marks are strongly
black outlined distally in whitish. Hind wing ground colour slightly more rusty; basal black
lines thin; discal white band well represented but narrower, broken towards inner border;
postdiscal row of lunules weak in upper sector but stronger from mid point to anal angle;
submarginal row of linear whitish marks strong; marginal border brick-red to lower tail then
olive with black dots to anal angle.

Variations, (a) Generally similar to above, but Upperside spots and marks in fore wing slightly
larger, spots from 2-5 slightly conjoined. Hind wing band slightly wider. Underside. Very
similar to nominate form, basal fore wing spots and pattern slightly stronger; fore wing spots
" rayed ". The figure of the type is intermediate between the two. (PI. 7, fig. 60.)

Variation, (b) Basically similar to (a) but Upperside fore wing spots white and more strongly
conjoined. Hind wing band wider, with slight blue distally on lower half. Underside. Basal
black marks less strong, but white pattern strong and fusion of fore wing spots clearly indicated.
(PI. 7, fig. 63.)

Variation, (c) Upperside. Ground colour blacker; pattern of fore wing very similar to (b).
Underside. Ground colour darker brownish, but pattern more contrasty and stronger; the
submarginal dark spots in fore wing complete to subapex ; the postdiscal row of lunules in hind
wing stronger.

$ form alladinis Butler, 1869 : 5

Fore wing length 40-42 mm. Upperside. Ground colour purplish blue-black, sometimes
rather brownish black. Fore wing with bluish spot toward upper end of cell and a small whitish
one beyond, blue spots sub-basal in 2 and 5 in discal line; a row of postdiscal spots, three in
line in subapex bluish white or bluish, followed by a slightly larger blue spots in 4-ib. Margin
with indistinct glaucous marks, strongest in ib. Hind wing ground colour as fore, blue spots
rather indistinct, one diffuse spot subcostal in discal line; spots in postdiscal line whitish at
subcosta but bluish and almost obscured, slightly more whitish toward inner fold ; submarginal
row of linear whitish marks complete but not strong; marginal row of lunules reddish above
upper tail then olive to hind angle which has two black dots; edge black; tails longish, slightly
spatulate at end, 5-6 mm. lower 4-5 mm., more pointed at end.

Underside. Fore wing ground colour greyish brown, more brownish on the distal border;
basal black lines moderately strong; discal and postdiscal spots of above strongly represented,
greyish white, bolder, edged proximally in black; the double tornal spot black with pale surround,
not very large. Hind wing ground colour more brownish, sub-base with a greyish irregular bar;
discal band irregular and much broken at 5-6, but strong on inner fold ; postdiscal series of olive
and maroon lunules strong and extending from costa to above hind angle; submarginal row of
linear whitish to lilac marks complete and accentuated with black marks opposite tails and
double at anal angle. (PI. 7, figs. 58, 59, 61, 62.)

REVISIONAL NOTES ON AFRICAN CHARAXES 101

Variation. In some specimens the upperside fore wing spots may all be blue and the post-
discal spots in the hind wing limited to the two upper ones only, the rest totally obscured.

$ iormfulgens Rothschild, 1900 : 487

Fore wing length 40 mm. Upperside. Ground colour deeper blue-black, slightly purplish
toward base. Pattern as in alladinis but all spots of fore wing bolder and usually bluer; the
marginal glaucous or greyish lunules more apparent. Hind wing with costal and postdiscal
spots larger and more distinct, blue, often with a bronzy tint. Underside. Pattern as in
alladinis but stronger and bolder, the fore wing postdiscal strongly supported distally by bold
black marks in the submarginal zone, shaded with grey distally; there is also a large black mark
in ib-ia between discal and postdiscal spots. Hind wing pattern generally as in alladinis
but much bolder, the postdiscal zigzag line of lunules heavily brownish distally ; the submarginal
linear lilac to whitish marks strong, followed by the clearer marginal border strongly reddish
above upper tail and olive to anal angle with strong black dots at tails and double at anal angle.
This form is a transition towards the next. (PI. 6, figs. 53, 54 (Type).)

9 form regalis Rothschild, 1900 : 486

Fore wing length 40 mm. Upperside. Ground colour black with blue sheen toward the
base. Pattern very similar to fulgens, but those of the discal series strongly developed and
reaching to the hind margin; the postdiscal spots often large, all spots except those in la-ib
orange-ochre, the latter blue. Hind wing slightly darker in ground colour has a well marked
blue band made up of six marks in the discal row and two in the postdiscal in the subcosta
area. The submarginal row of linear bluish white marks strongly developed and separated
from the marginal border by black; marginal border reddish to lower tail then olive to anal
angle. Tails 6 and 5 mm. long, reddish, narrowly edged black. Underside. The basal area
of fore wing greyish brown, with deeper brownish in the discal area and wing margin ; the discal
and postdiscal light spots strong; the submarginal black spots of tornus and one above strongly
outlined distally in greyish. Hind wing ground colour browner than fore except at base ; discal
band distinct but rather interrupted in mid area ; the postdiscal lunules, olive and maroon bold ;
the submarginal linear marks strong; marginal border reddish to lower tail then olive to anal
angle. In some specimens the fore wing spots are not large and may be entirely blue or with a
suggestion of bronzy red. The type specimen has exceptionally large spots. The pattern suggests
a transition toward the " etheocles " pattern. (PI. 6, figs. 51, 52.)

<$ form ephyra Godart, 1824 : 330 (355)
= <$ form hollandi Butler, 1893 : 266

Fore wing length 35 mm. Upperside. Ground colour black with little or no sheen. Sub-
costal blue spots variable, but usually one at end of cell, one beyond and one or two subapical.
Marginal lunules greyish, small or vestigial. Hind wing ground colour black; submarginal
whitish spots small or ill-defined; margin may be slightly greenish; extreme edge black with
slight white fringe well separated between veins. There may be a suggestion of a wavy greenish
line opposite the tail in the postdiscal zone, usually absent. Underside. Fore wing ground
colour greyish brown with strong satiny bars, one through end of cell, a complete bar through
the disc outlined proximally in black, followed by a less marked bar in the postdiscal line;
border broadly satiny with a row of dark spots from apex to tornus, of increasing size, blackest at
tornus. Basal black marks clear and white-edged. Hind wing ground colour as fore, crossed by a
satiny bar in sub-base; a satiny bar through the disc outlined in black proximally; postdiscal
olive and maroon lunules strong, outlined in black proximally; submarginal line of pale marks
accentuated distally in region of tails with black; marginal border mostly olive with some
reddish scales above upper tail. The underside thus has a variegated appearance. (PI. 6,
figs. 49, 50.)

102 V. G. L. VAN SOMEREN

$ form carteri Butler, 1881 : 108

Very similar above to ephyra Godart but on underside rather variable, satiny bars less
developed and on the whole less variegated and suffused over with a brownish or rusty purplish
tint. There is much intergrading. (PI. 6, figs. 47, 48.)

<$ form catochrous Staudinger, 1896 : 218

This form was described from Cameroon as an aberration of ephyra. When we
wrote our paper on the etheocles complex (van Someren & Jackson, 1952) only one
male specimen was known from the W. African zone, taken at Lagos, Nigeria. It
is now known to occur in Ghana and Ivory Coast and in Liberia. This extension
westward increases the range of overlap of this form with ephyra and carteri. The
question thus arises were we justified in considering catochrous as a distinct species?
Our action received support when in 1955 we bred a large family from a female
which gave thirty-three males of the catochrous form and females similar to the parent.
Two subsequent families also from " catochrous " females produced both carpenteri
and catochrous females and male forms ephyra (hollandi) and catochrous. It is true
that these families were from females captured at Katera, Uganda, within the range
of Ch. etheocles carpenteri and not from Cameroon or W. Africa, nevertheless the
catochrous males are indistinguishable from Cameroon specimens. I am thus of the
opinion that catochrous must be considered merely a male form of etheocles.

Ghana and Ivory Coast specimens :

Fore wing length 35 mm. Upperside. Ground colour black; blue spots small, one at upper
end of cell, one beyond and two in upper subapex slight whitish ; marginal glaucous spots small ;
fringe narrowly white. Hind wing ground colour as fore; submarginal row of small whitish
spots not strong; marginal border greenish. Underside. Basal area to discal bar silvery;
fore wing bar not strong but proximally lined in black ; distal portion of wing browner and post-
discal pale spots not strong; margin of wing brownish. Hind wing silvery at basal half crossed
by paler sub-basal satiny bar; discal white bar strong, bordered brownish, postdiscal line of
olive and maroon lunules strong accentuated proximally in black; marginal border reddish
above upper tail greenish to hind angle ; submarginal lilac spots not very strong but with black
dots at anal angle.

Range : The nominotypical etheocles ranges from Sierra Leone and Liberia to Ivory
Coast, Ghana and Nigeria, where it merges eastward into a slightly different larger
form in Cameroon, Moyen Congo, west of the Congo River. (Map 4).

REGION 2. CAMEROON, CENTRAL AFRICAN REPUBLIC, MOYEN CONGO,
WESTERN CONGO, KATANGA AND ?N. ANGOLA.

Charaxes etheocles biinclinata ssp. n.

(PL 10, figs. 82, 83, 85-87, 89, PL n, figs. 92, 93, 96, 97)

Type locality, Moyen Congo.

This is a mixed aggregate, occupying a very large area, and might be considered
a " cline " between nominate etheocles of the western area and ochracea of the Gabon,

REVISIONAL NOTES ON AFRICAN CHARAXES

103

lower Congo area, and it would appear to merge with carpenteri in the south-eastern
Congo area. In this aggregate, the female forms resemble variations of the nominate
race, alladinis and regalis and the majority of them are larger than those of the
nominate etheocles.

$ f. resembling etheocles

Fore wing length 42-44 mm. Upperside. Ground colour brownish black towards base,
blacker on distal two-thirds of wing. Pattern of white spots bold, the spots being well separated,
a rounded or triangular spot at upper end of cell, a larger ovoid spot at base of 4 set well in
from the two upper spots of the discal line, spots of 3-1 b of increasing size, that of la a streak;
postdiscal series of spots complete from three in line in subapex, spot in 4 set well in, spots in
3-ib following the curve of the wing, spot in ib touching the mark in discal line. All spots

>M~-,.

; \

\ \ r

} \:"'

A

x >' \" CENTRAL ( , __;

'/ ;-AFRICAN REPUBLIC'''.^ C '"\ ETHIOPIA

KEY

KEY JV;- a A.

GABON " oi^
Charaxes etheocles etheocles. V 1 . oo (

\.vAi

O C. etheocles biinclinata. MOYEN CONGO C

Region 2. V/f-T^

C. etheocles ochracea.
Region 3.

D C. etheocles carpenteri.
Region 4.

Region

Charaxes grahamei.
Charaxes contrarius.

ANGOLA l I KATANGA

o.6 o

MAP 4.

104 v - G. L. VAN SOMEREN

creamy in colour. Margin with slight indication of greyish, separated by black at ends on
veins. Hind wing ground colour brownish black at base, border wide and blacker; discal band
whitish to creamy, of about equal width to ic then tapering and crossing the inner fold, inner
border slightly bluish. Submarginal elongate whitish marks distinct and well spaced; marginal
border reddish orange above upper tail, then olive to anal angle. Tails long and slender 6-7
mm. upper, lower 5 mm., black with orange central line. Underside. Ground colour greyish
brown, darker on the border of fore wing. Pattern bold, basal black marks strong. Upper-
side. Discal and postdiscal spots strongly represented; submarginal row of dark marks
complete, strongly black at tornus and space above; a black mark also present in ib. Hind wing
basal area slightly darker brown with a satiny bar in sub-base ; discal whitish bar strong, though
less wide than above ; postdiscal line of olive and maroon lunate marks strong ; submarginal row of
linear whitish marks strong; marginal border reddish to mid tails, then olive to anal angle. (PI.
10, figs. 82, 86.)

Holotype female. MOYEN CONGO: Kelle, x.ig62 (Jackson).

Variation, (a) Very similar to above. Upperside. Ground colour more strongly black;
fore wing spots smaller, and ochre in colour. Hind wing band wide at costa but tapering rapidly
the outer border incurved ; submarginal linear marks more bluish. Underside. Ground colour
more uniform greyish brown, satiny bars more diffuse; pattern of above, well-marked in
discal line but less strong in postdiscal row, with the adjacent dark marks fading out above
ib. Hind wing discal band creamy white and distinct; the zigzag row of olive and maroon
lunules strong; submarginal row of linear pale marks clear; marginal border reddish to upper
tail, then olive to anal angle. (PI. 10, fig. 85.)

Variations, (b) Upperside. General pattern of fore wing very similar to PL 10, fig. 82 but the
spots are conjoined by rays. Underside. Ground colour more greyish; pattern of above well
represented and showing raying for fore wing spots ; marginal pale area divided by black veins.
Hind wing ground colour more brownish and with a sub-basal satiny bar present but not strong
discal creamy white band well represented; the postdiscal lunules not marked in upper half
but strong thereafter to anal angle ; submarginal whitish line broad but rather diffuse ; marginal
border reddish orange, outlined in ochre. (PI. 10, figs. 83, 87.)

Variations, (c) Upperside. Fore wing ground colour brownish black with purplish tinge on
basal area; distal border of wing more brownish. No spot in cell; spot in sub-base 4 very small;
two upper spots in discal row quadrate, those in 3-2 crescentic, marks in ib-ia increasing in
size. Postdiscal spots large, of almost equal size, that in ib in contact with mark in discal row.
All spots orange-ochre except those in la, ib discal row which are creamy. Hind wing ground
colour as fore wing; the discal band rather irregular on borders, showing a tendency to raying
on lower half of outer border, colour creamy with a tinge of purplish colour on inner border in
2-3. Submarginal linear marks creamy with a slight purplish surround; marginal border
orange-red to lower tail, then olive to anal angle. Underside. Ground colour brownish grey
with purplish flush over bases of both wings, more rufous on distal border of fore wing. The
black marks at bases well developed, so also the black line on the proximal side of the discal
ochreous spots; the postdiscal spots strongly ochreous; the postdiscal spots on fore wing only
strong at tornus. Hind wing discal bar ochreous with some indication of breaking up toward
the inner fold. Postdiscal lunate marks not strongly shown and suffused over by a rufescent
bloom; submarginal pale marks strong; border reddish to lower tail then slightly olive to anal
angle. (PL 10, fig. 89.)

This variation shows a strong tendency towards ochracea Rothschild of Gabon.

$ form near regalis Rothschild.

Fore wing length 43 mm. Upperside. Ground colour black with some purplish brown
towards the base; pattern of discal and postdiscal spots much smaller than in the nominate
regalis type (q.v.), particularly in the discal row; all spots ochreous orange except those in la-ib

REVISIONAL NOTES ON AFRICAN CHARAXES 105

which are blue. Hind wing ground colour blue-black, slightly browner at base; discal bar blue,
not so strong as in nominate regalis especially as regards the two postdiscal spots at subcosta
The submarginal row of linear marks bluish, not so pronounced; marginal border reddish to
lower tail, but olive at anal angle. Underside. Ground colour brownish grey, slightly browner
on outer border; black basal marks bold. Discal spots greyish, outlined in black proximally;
postdiscal spots greyish white in ib-3 then ochreous to apex; tornal spot and one above strongly
black, the dark spots in spaces above hardly visible. Hind wing ground colour slightly browner,
sub-base with paler bar ; discal band whitish at costa, irregular and fading out at 2 to inner fold ;
olive and maroon lunules in postdiscal line rather large but not strong except opposite tails;
submarginal row of linear marks rather diffuse, but narrowly outlined distally in black ; marginal
border not very clearcut but reddish to lower tail then olive to anal angle, which has two black
dots.

In some examples, the fore wing spots are blue with little orange tinge. There is thus consider-
able difference between these specimens and nominate regalis, q.v.

$ form near alladinis Butler

Very similar to form alladinis of the nominate race, but slightly larger; fore wing length
43-45 mm. Upperside. Ground colour purplish brown-black, slightly blacker on distal half
of wing. Spot in cell and in discal line rather obscured, purplish blue; postdiscal spots more
distinct and extending from subapex to ib, upper ones whitish, lower purplish blue. Hind
wing with two rather indistinct purplish blue marks subcostal in discal line; postdiscal spots
indistinct and purplish blue ; submarginal row of whitish linear marks distinct ; marginal border
reddish to upper tail then olive to anal angle. Underside. Ground colour rather dull, uniform
greyish brown with rufescent bloom overall. Basal black marks thin ; discal row of paler spots
hardly visible except in subcosta but outlined proximally in black; postdiscal marks more
distinct; tornal black mark distinct. Hind wing almost uniform with only slight indication of
discal and postdiscal spots; submarginal row of linear white marks distinct.

The males associated with this aggregate are of two forms which differ mainly on
underside characters. They are f. ephyra, carteri and f. catochrous.

The male forms are ephyra and carteri.

These three forms are hardly separable and have been referred to when dealing
with the nominate race.

Upperside. Ground colour deep velvety black with an overall blue-green sheen, most pro-
nounced over the disc of the hind wing. Length of fore wing, 33-35 mm. The number of sub-
costal blue spots variable, there is usually one at the upper part of end of cell, a larger one at
the discal line, and sometimes one or two in the subapical area, but these may be absent. Margin
with varying degree of glaucous marks. Hind wing sometimes with a trace of a wavy greenish
line in the lower postdiscal zone opposite the tails, but more often this is absent; submarginal
row of white dots with lilac surround complete; marginal border greenish; edge black, with
very slight white fringe between veins. Tails short, rather thin, of about equal length, 3-4 mm.
Underside. Ground colour rather dark earthy grey-brown with slight purplish flush. A satiny
bar present in the discal line, and a satiny V at apex. Basal black dots and lines usually strong
and outlined in white; submarginal row of dark subdued marks complete, those at tornus and
space above black; margin with dark marks at end of veins. Hind wing ground colour slightly
darker than fore, with a strong satiny bar in sub-base, and a stronger one through the disc,
rather irregular at lower end but represented at the inner fold by a somewhat triangular whitish
mark ; postdiscal row of olive and maroon lunules strong, outlined proximally in black ; submar-
ginal row of linear white marks clear; marginal border dull greenish with some admixture of
red above upper tail then olive to anal angle; usual double black spot at anal angle, and near
bases of tails. (PI. n, figs. 92, 96.)

io6 V. G. L. VAN SOMEREN

$ form catochrous Staudinger

Upperside. Ground colour deep velvety black; fore wing subcostal spots variable in size and
number, sometimes absent; marginal glaucous marks small and often hardly visible. Hind
wing submarginal spots small, bluish or obscured; marginal border narrow, greenish, plainly
visible or obscured, olive at anal angle. Underside. Pattern essentially as in other forms
but bolder, the basal half of the wings silvery to the discal lines, the distal half of the wings
brownish; satiny bars strong. In some specimens the contrast is less strong. (PI. u, figs. 93,
97-)

Range: south-eastern Nigeria and Cameroon, the Central African Republic and
Moyen Congo to the Lower Congo, Kasai and Katanga, where it appears to meet
etheocles carp enter i. (Map 4) .

REGION 3. GABON AND ADJOINING TERRITORY, LEOPOLD AREA, LOWER CONGO
Charaxes etheocles ochracea van Someren & Jackson, 1957

f. violacea Rothschild, 1900 : 485.
f. ochracea Rothschild, 1900 : 486.

This subspecies is concentrated in a comparatively small area of the total distri-
bution of the species. The female forms of this aggregate exhibit a common char-
acter, that of strong purplish admixture with the blackish brown of the ground
colour, and that the pattern of spots on fore and hind wing are predominantly
ochre-yellow to orange. The first name to be applied to any one of this aggregate
was ochracea Rothschild, 1900 : 486. We adopted this name to represent the
subspecies.

The males associated with this aggregate were named violacea Rothschild, 1900,
the type coming from Ogowe River, Gabon. It was described as having the upper
side violet-blue, but the degree of violet sheen is not constant.

Fore wing length 35 mm. Shape slightly variable in regard to the incurvature on the outer
border of the fore wing. Upperside. Ground colour deep velvety black; fore wing blue spots
variable in size and number, from immaculate to one very small subcostal spot or three visible
marks. Marginal marks hardly visible except at hind angle. Hind wing with submarginal
series of white to lilac spots small, clearly marked; edge with slight white fringe. Underside.
Clayish grey-brown at basal half, more brownish with a slight purplish flush beyond; basal
marks strong, often heavy; satiny bars not strong; submarginal dark marks strong at tornus
and space above. Hind wing basal area as fore wing up to discal line; greyish brown, darker
on postdiscal zone where the series of lunate olive and maroon marks are strong; submarginal
row of whitish lilac marks strong; marginal border reddish to upper tail, then olive to anal angle.

Lambarene males are colder in tone on the underside. (PI. 10, figs. 84, 88.)
The females are extremely variable as regards pattern on the upper side.

$ form ochracea Rothschild 1900 : 486

Fore wing length 40-44 mm. (type, 40 mm.) Upperside. Ground colour brownish black
with a strong purplish tone. Pattern bold; in the discal row there is a large subcostal spot with a
slightly smaller one below, the spot at base of 4 is set well in, subsequent marks of increasing
size, somewhat angular, mark on xa, a streak; postdiscal spots large, three subapical in a row,

REVISIONAL NOTES ON AFRICAN CHARAXES 107

elongate; spot in 4 set in a little, those of 3-ib larger, than in ib touching the discal mark in
same area. All spots ochreous or orange-ochre, those in la-ib slighter paler. Margin of wing
with large but diffuse marks, most visible at tornus. Hind wing ground colour as fore, slightly
darker on border. Discal band complete, wide at costa and gradually tapering toward the inner
fold, colour buffish ochreous with slight purplish tinge to borders; marginal border orange to
lower tail then olive to anal angle. Underside. Rufous brown strongly flushed with purplish
especially at bases of wings. Basal black marks in fore wing well marked, but those of hind
wing fine. The discal and postdiscal rows of spots buffish and clearly defined in the discal row,
but less clear and ochreous in postdiscal; tornal black mark small, double; with inner black mark
in same area distinct. Hind wing discal band buffish, rather irregular toward inner fold,
represented at the fold by a triangular pale mark. Postdiscal row of reddish buff lunules pale at
upper end but strong toward lower half; submarginal row of linear marks strong; marginal border
orange red then olive at anal angle. Tails 6 and 5 mm. long. (PI. n, figs. 91, 95.)

Variation, (a) Upperside. Fore wing spots are large and slightly joined by rays, spots
ochreous; marginal marks large and diffuse. Hind wing band ochreous, the outer border
irregular and somewhat dentate. Otherwise very similar to type.

Variation, (b) Upperside. Fore wing spots are smaller, less strongly orange-ochre, the marks
in la-ib whitish tinged with purplish. The hind wing band narrower and more tapering,
whitish, flushed with purplish on outer border. Underside. More rufous with purplish flush;
the pattern of both wings strong in the discal lines.

Variation, (c) Upperside. Fore wing spots are large, whitish and buffish tinged with violet
in ibib; the hind wing band off-white with strong purplish buff suffusion on outer border
which is rather diffuse. Underside. Ground colour is darker in the basal areas of both wings,
but more greyish on outer half beyond the discal line in fore wing with the postdiscal series of
spots not so distinct. The postdiscal row of lunules in the hind wing strong in the lower half.

Variation, (d) Ground colour and pattern as in type ochracea but all marks in fore and hind
wing strongly ochreous. $ f. ochreata van Someren & Jackson, 1957.

$ form seriata Rothschild, 1900 : 487.

Fore wing length 40 mm. Upperside. Ground colour darker than in previously mentioned
forms of ochracea being brownish black with a strong purplish tone. Discal spots represented
by one large at subcosta with a small dot below, marks in 3-2 almost obliterated; postdiscal
spots complete and large though not sharply defined, three subapical in line, spots from 4 to
ib increasing in size; all spots ochreous, marginal lunules large but indistinct. Hind wing
ground colour as fore, slightly darker in disc; no discal spots, but postdiscal series of indistinct
marks are visible ; submarginal linear marks strong and buffish white ; marginal border strongly
developed, orange-red to lower tail, then olive at anal angle ; extreme edge black, tails long and
slender, upper 6 mm., lower 4 mm., black with central orange streak. Underside. Ground
colour grey-brown strongly rufescent with a purplish brown flush, especially on the hind wing,
border of fore wing more rusty; basal black lines in fore wing strong, but spots in discal line
represented by upper ones only; those of the postdiscal series complete but rather suffused over;
the tornal black mark moderately strong, accentuated by whitish surround. Hind wing pattern
rather suffused over, but sub-basal and discal pale bars visible; the postdiscal line of lunules
fairly distinct strongest at lower half; submarginal series of linear marks buffish and well developed,
margin reddish to lower tail, olive to anal angle. (PI. n, figs. 90, 94.)

Variation, (a) Upperside. Resembling somewhat nominate seriata but fore wing post-
discal spots smaller. No suggestion of postdiscal spots in the hind wing. Underside generally
darker than in seriata.

Variation, (b) Upperside. Somewhat like (a) but ground colour stronger, more purplish
brownish black, the margin of the wing more rusty. The discal spots in fore wing limited to one
large subcostal and a dot below, buffish ochre in colour; postdiscal spots large and ochreous
complete to ib, marginal lunules large but diffuse but distinct in ib. Hind whig without, or with

ENTOM. 23, 4. 10

io8 V. G. L. VAN SOMEREN

a very faintly indicated row of postdiscal spots in the hind wing ; submarginal linear marks bold,
huffish in colour; marginal border strong. Underside with black basal marks in fore wing strong
to discal line; rest of pattern partially obliterated by rufous purplish tone to outer border; hind
wing pattern rather subdued, but with a darker zone in the discal line, the postdiscal lunules
though large, rather hazy in upper half but clearer toward lower end; the submarginal row of
linear marks, strong and buffish in colour; marginal border broad, orange-ochre to lower tail,
and olive at anal angle.

Variation, (c) This variation links the seriata group with ochracea. Upperside. Ground
colour purplish brown, rather more rusty on the border of fore wing. The discal spots are
well developed and clear to 2, slightly indicated in ib, whitish or cream in colour; the post-
discal series large and whitish buff extending from subapex to la; margin with large, rather
diffuse buffish marks contiguous with postdiscal spots. Hind wing ground colour darker in the
disc, more blackish, no discal marks, but postdiscal series of spots well denned, buffish white in
upper part but ochreous with purplish brown tinge in lower part to above anal angle. Submar-
ginal linear buffish marks strong; marginal border orange-red to lower tail, then olive at anal
angle. Underside. Ground colour purplish brown, more rusty on outer border with pale margin.
Discal buffy spots distinct, but those of the postdiscal row becoming rather faint toward the
subapex ; tornal black mark double but small. Hind wing pattern rather overshadowed by the
overall purplish brown tone ; the postdiscal lunules also subdued by the rusty ground ; the submar-
ginal linear ochreous marks suffused over in upper half; marginal border orange-red to lower
tail, olive at anal angle.

Range : Restricted mainly to the forest areas of Gabon, with a slight extension
into Leopold area of S. Congo and northward into the southern corner of Cameroon.
There appears to be some intergrading with biinclinata in the southern part of the
Moyen Congo. (Map 4) .

REGION 4. S.W. OF CONGO BASIN, KATANGA, N.W. RHODESIA, UGANDA, S. SUDAN

AND ADJOINING AREA OF ETHIOPIA, N.W. TANZANIA

Charaxes etheocles carpenteri van Someren & Jackson, 1957 : 54.
(Text-figs. 2-4 [Aedeagus])

The eastern aggregate of ethocles was recognized as a subspecies by van Someren
& Jackson (1957 : 54). We selected as type, the eastern representative of form
alladinis, to which Poulton had given the name carpenteri in 1918, this being the
first name to be applied to any member of this aggregate. This form is unfortu-
nately highly variable. The type was taken at Kakindu Hill, south of the Kagera
River, in the Bukoba district of Tanzania.

MALES. Although we designated violacea Rothschild as the male of this race
(van Someren & Jackson, 1957 : 54) this was an error! Form violacea Rothschild
is now restricted as the male of etheocles ochracea of the Gabon area. In our previous
paper (1952), we considered that the male form catochrous Staudinger with a strong
silvery basal area to both wings was the male of a distinct species with only one form
of female. This also has been shown to be incorrect. We are confronted with the
fact that though all these males resemble each other on the upper surface, within
minor limits, the underside may be brownish as in carteri, or silvery in the basal
areas as in catochrous, and there are many intermediates. They can be bred in the

REVISIONAL NOTES ON AFRICAN CHARAXES 109

same family. We however designated a male from Mabira Forest, Uganda, as
male of carpenteri 1957 : 54.

cJ form carpenteri van Someren & Jackson, 1957 : 54.

Fore wing length, average, 35 mm. Outer border of wing slightly concave at 3-4. Upperside.
Ground colour velvety black (may be tinged brownish in very old specimens) . Little or no indi-
cation of glaucous marks on margin; no blue subcostal spots, or there may be very small dots,
one at upper end of cell, a minute dot beyond in discal line, and one or two in subapex. Hind
wing ground colour as fore, submarginal whitish spots may be small and distinct or somewhat
obscured; marginal border narrow and greenish throughout, or with some olive at anal angle.
Underside. Ground colour brownish drab, darker brownish on outer border, satiny bars strong,
a quadrate mark in cell and a broad satiny bar through the discal zone, outlined proximally in
ib~3 in black, a satiny V mark at apex; basal black lines fine to moderate; submarginal dark
marks black at tornus decreasing in size and colour to subapex, slightly ringed in greyish.
Hind wing ground colour as fore slightly darker in basal half crossed by a paler satiny bar, a
stronger satiny bar through the disc outlined proximally in black; postdiscal zone browner,
with postdiscal olive and maroon lunules strongly indicated, finely outlined in black proximally;
submarginal pale line irregular and faintly indicated; marginal border reddish to upper tail
then olive to anal angle with black dots between tails, double at anal angle. Tails short, some-
what stumpy, 4 and 3 mm.

Variation, (a) Upperside. Fore wing margin with distinct glaucous lunules. Submarginal
white spot and border distinct. Underside. The whole more uniform greyish brown, slightly
browner on the border of fore wing and on disc of hind wing; satiny bars more diffuse so that
in side light the whole surface has a glazed appearance, but the black marks may be strong. The
postdiscal line of wavy lunules almost obscured except toward the anal angle, the submarginal
linear pale marks complete but indistinct; but the marginal reddish and olive border is strong.
(PL 8, fig. 69.)

Variation, (b) Upperside. Differs little from the nominate, except that the fore wing
subcostal spot in discal line is distinct, the postdiscal a mere dot. Underside. Pattern is
however, strong and in side light the satiny bars show up strongly as they do in the nominate,
suggestive of an intermediate stage towards f. catochrous.

$ form carpenteri Poulton, 1918 : Ixxxii

Fore wing length 43 mm. Upperside. Ground colour of fore wing brownish black at base,
blacker on the distal half, with a slight purplish sheen. Pattern not strongly marked, in the
discal line are two spots, upper one whitish and large, lower small, no spot in 4, spots in 3-2
whitish, rest greenish blue. Submarginal Imear white marks well developed; marginal border
two whitish, spot in 4 set in, that in 3 slightly in, that in 2 slightly out and crescentic in shape,
spots in la-ib purplish blue. Margin with diffuse glaucous lunules. Hind wing ground colour
black with strong greenish blue sheen in disc. No discal marks ; but postdiscal spots extending
from costa in an irregular line to short of the inner fold where there is a pale mark, upper spots
whitish, rest greenish blue. Submarginal linear white marks well developed; marginal border
a mixture of greenish and orange scales to lower tail then olive to anal angle. Tails longish,
thin, 7 and 5 mm. Underside. Ground colour greyish brown with slight rufous bloom; satiny
bars slight in discal zone, slightly stronger in postdiscal line; basal black marks thin; tornal
mark and one above black, with black marks more proximal in same spaces, with white mark in
between. Hind wing ground colour as fore, basal black marks very thin; satiny bars very
faint, only visible in side light ; postdiscal line of lunate marks rather angled in upper part and
not strong, but more marked toward the inner fold above the anal angle. Submarginal linear
buffish marks present but rather weak ; marginal border orange-red to between tails, then olive
to anal angle.

no V. G. L. VAN SOMEREN

Variation, (a) Upperside. Very similar to the type but fore wing spots smaller and limited
in the discal line; those of the postdiscal series extending to la; hind wing with postdiscal
series of greenish spots clear in upper part but obscured in lower. A faint indication of a spot
in discal line at costa. (PI. 9, fig. 75.)

Variation, (b) Upperside. Discal spots well represented with an extra dot at sub-base of 4,
the upper spots bluish, those faintly indicated in ib and 2 deeper blue to purplish; the post-
discal series complete, double in ib and a trace of spots in la, upper marks bluish white, lower
stronger blue. Hind wing with complete row of postdiscal spots and a faint indication of marks
in upper discal line; submarginal linear marks strong. Underside. As in (a) as regards ground
colour, but basal black marks stronger; pattern of upperside more clearly represented. The
hind wing with rusty flush overall rather obliterating the pattern except at lower end of
postdiscal line. (PI. 9, fig. 73.)

Variation, (c) Upperside. Very like (b) but mid-discal spots larger and white; postdiscal
spots complete to ib, bluish. Hind wing with a trace of a spot in discal line at costa, the
postdiscal spots complete to ib, large spot in upper end, bluish. Underside. Darker greyish
brown especially over hind wing, the fore wing pattern stronger, but hind wing pattern subdued
except for the postdiscal lunules which are strong except at upper end.

Variation, (d) Upperside. An interesting variation in which there is a large blue spot in
upper part at end of cell; the discal spots large with an extra spot at sub-base 4, all these spots
white, an obscured blue mark in ib; postdiscal spots small and bluish, the mark in ib elongate
and set out toward tornus. Marginal glaucous marks more distinct. Hind wing with an indica-
tion of a blue spot at costa in discal line; postdiscal spots very faint except the second upper
one which is white. Underside. Ground colour as in (c) but pattern stronger in fore wing with
postdiscal spots large and more distinct in the browner outer border. The hind wing ground
colour darker, with a satiny sub-basal bar and the discal and postdiscal lines of spots distinct.

The enlarged white discal spots of the fore wing in conjunction with the strong underside
pattern seem to indicate an element of the pattern of female form pallidimacula in the genetical
make-up of this specimen. (PI. 8, fig. 72.)

Variation, (e) Though very battered as a result of being caged for eggs, this specimen
exhibits some interesting and unusual features. Upperside. Ground colour though brownish
black, has a strong greenish tinge over the base of the fore wing and the disc of the hind wing.
The fore wing discal spots though comparatively large are greenish blue and subdued ; the post-
discal marks are also subdued and large extending from subapex in increasing size to the streak
in the hind margin. The hind wing has a much broader postdiscal band than usual made up
of large greenish blue marks indented on the outer ends ; the submarginal line of whitish marks
and the marginal border as usual. Underside. As in usual carpenteri. (PI. 9, fig. 74.)

Variation, (f) Upperside. Fore wing somewhat similar to (e) but postdiscal series of spots
more obscured and beyond these a series of blackish marks. Hind wing as in (e) but without
any definite postdiscal band, this area being broadly greenish blue. The underside is almost
identical to that of (e).

Variations in the females with " etheocles " pattern.

$ form pallidimacula van Someren & Jackson, 1957 : 54

Fore wing length 40-42 mm. Upperside. Ground colour brownish black, blacker on distal
half of wing. Pattern generally similar to that of etheocles etheocles but distal and post discal
spots white as in the type or very slightly creamy-tinged. A large spot at upper end of cell,
two large subcostal spots beyond in discal line, a spot at sub-base 4, set well in, spots from 3 to
hind margin of increasing size, that in la, a streak. Postdiscal series, three in an oblique line
in subapex, spot in 4 set in at an angle, spots from 3 to ib increasing slightly in size, spot in ib
merging with discal mark. Margin without distinct pale marks except at tornus in ib. Hind
wing with a broad white band of about equal width from costa to cell area then tapering to

REVISIONAL NOTES ON AFRICAN CHARAXES in

above anal angle and crossing the inner fold, borders slightly shaded with blue, mostly opposite
tails; submarginal row of linear marks white and strongly developed; marginal border reddish
to upper tail then olive to anal angle; tails moderately long, 7 and 5 mm., upper with rounded
end, lower pointed. Underside. Ground colour greyish brown, slightly darker in interspaces;
black line in basal area fine; pattern of above showing up strongly; tornal black marks and
smaller one in space above, strong. Hind wing basal area with very fine black lines ; discal white
bar strong but narrower than above and restricted on the inner border; postdiscal line of lunules,
olive and brownish, weak at costa but strong to above anal angle; submarginal linear marks
whitish, rather dyslegnic; marginal border reddish to upper tail then olive to anal angle; black
dots present at bases of tails, double at anal angle. (PI. 8, figs. 64, 65.)

Variation, (a) Upperside. Generally similar to nominate but fore wing spots in discal row
slightly larger, the spot in ib not in contact with that of postdiscal row, spotmark in la, a long
streak, extended distally. All fore wing spots white. Hind wing discal bar narrower, straighter
on the outer border and with very little bluish on inner border. Underside. Ground colour
paler, but pattern essentially similar.

Variation, (b) Upperside. Ground colour as in (a); no spot in the fore wing cell; all spots
creamy. Hind wing bar strongly bluish with white line in upper discal area. Underside.
Ground colour rather paler, but pattern clear; hind wing discal band restricted to discal zone.
One of many similar specimens bred in family containing f. carpenteri. (PI. 8, fig. 71.)

Variation, (c) Upperside. Fore wing ground colour brownish black, slightly darker on
border. Fore wing discal spots larger slightly conjoined, that in ib-ia extended distad to post-
discal line, and extended on the proximal side where they are pale bluish; postdiscal spots
elongate ovoid; a minute dot in the cell; all marks creamy ochreous. Hind wing band wide
slightly restricted at costal end, the band is widest at and extends over the cell area where it is
strongly shaded with blue, submarginal linear marks thin; border rather darker maroon above
upper tail but olive at anal angle. Underside. Ground colour as in previous form but rather
rusty on outer border of fore wing, satiny sheen overall; hind wing with darker band proximal
to discal white band and in zone carrying the postdiscal olive-ochre and olive lunules which are
strong, submarginal row of linear marks strong and border well developed but narrow. (PI. 8,
fig. 66.)

Variation, (d) Upperside. Ground colour as above, slightly darker on border which has
indication of large dark spots; edge with pale marks from tornus to 4. Large spot in cell of
fore wing, discal and postdiscal spots large, slightly rayed or conjoined, all marks ochreous.
Hind wing blacker on border ; discal band wide at costa, tapering to above anal angle and crossiug
inner fold, creamy white with bluish on inner border. Underside. Ground colour as previous
form, pattern stronger, fore wing marks creamy; hind wing with brownish zone proximal to
discal white band; rest of pattern strong.

There are many intermediates between these variations.

c form catochrous Staudinger, 1896 : 218

Upperside. Closely resembles the other forms; the subcostal blue spots of the fore wing may
be small yet visible, one in upper part of cell, one in the discal line and two in the subapex,
or they may all be wanting. The margin may be without marks or there may be small white
marks. On the hind wing the submarginal row of white marks may be punctiform or short
linear. Underside. Presents a rather striking appearance, the basal two thirds of both wings
are a silvery greyish white in which the dark marks show up clearly, in addition, the satiny bars
are well developed. The distal third of the fore wing beyond the discal zone is browner and in
this the submarginal spots, black at the tornus extend up to the subapex, are ringed in whitish
and accentuated proximally in black. On the hind wing the satiny bar in sub-base and that of
the discal line show up strongly and in the darker border the postdiscal series of olive and maroon
lunules are strong ; the submarginal linear white line is sharply defined to above the anal angle ;
the marginal border, though narrow is red to upper tail then olive to anal angle with black dots
between tails double at anal angle. (PI. 8, figs. 67, 68.)

ii2 V. G. L. VAN SOMEREN

The underside pattern of all these forms is essentially the same, and though the two extremes
look different, both forms and transitionals often occur in one bred family.

Range: The range of this subspecies is from the border of N.W. Kenya with
Uganda, from Kakamega-Kabras to Busia in eastern Uganda, westward to the eastern
side of the Congo Basin, south to Katanga and the north-eastern side of Tanzania.
There is some apparent contact with biindinata in the Kasai-Katanga area. (Map 4).

REGION 5. N.E. KENYA AND EASTERN AREAS OF Mr. ELGON, TRANSNZOIA TO MAU,

LUMBWA, LONDIANI, SOTIK-MARA, LEMBUS AND RAVINE

Char axes etheocles evansi van Someren & Jackson
(PI. 9, figs. 76-81, Text-figs. 5-6 [aedeagus])

Charaxes etheocles $ form evansi van Someren & Rogers, 1932.
Charaxes etheocles evansi van Someren & Jackson, 1957 : 54-

MALE. Variation, (a) Fore wing length 35-36 mm. sometimes smaller. Shape, apex
rather pointed, in fore wing. Upper side. Ground colour velvety black, slightly tinged brown
at base in some examples. Blue spots along subcosta variable, may be minute or strongly
represented; marginal glaucous marks hardly visible or may be small but strong at tornus.
Hind wing ground colour black; submarginal row of white marks usually distinct; marginal
border may be mostly greenish or reddish to upper tail then olive to anal angle. Tails short
almost stumpy, of about equal length, 4 mm. Underside. The ground colour and strength
varies in the same way as we have noted in the females (a). It may be greyish brown, slightly
browner in the curve of the fore wing and over the discal portion of the hind wing. Satiny bar
of the fore wing moderately strong ; basal black marks fine, black lines in interspaces on hind
half graduated from heavy in ib to fine; submarginal dark marks heavy at tornus and space
above. Hind wing ground colour as fore wing, satiny sub-basal bar strong; that through the
disc strong in upper portion, rather broken in lower ; postdiscal olive and maroon lunules strong ;
submarginal pale line obscured in upper part but clear in lower; marginal border narrowly red
above upper tail, olive to anal angle, between row of black marks strong.

Variation, (b) Upperside. With minute blue marks in subcostal area of fore wing; no
marginal spots. Underside. Ground colour darker grey brown, with richer brown in inter-
spaces and on the outer border of the fore wing; the satiny bars showing up more strongly in
the discal line and inner submargin and at apex. Hind wing with correspondingly strong
satiny bars and a darker zone in the postdiscal line.

These variations can be present in one family.

FEMALE. Fore wing length 40-42 mm. ; shape slightly more pointed, as a rule, than etheocles
carpenteri. Upperside. Ground colour deep brownish black, rather browner at base of wing.
Fore wing pattern strong; a suggestion of a spot toward upper end of cell; subcostal spots in
discal line in 6-5 one above the other, spot in 4 set well in, that in 2 slightly out, spots increasing
in size to hind margin, that in la a streak extending distad. Postdiscal series, three subapical
in an oblique line, spot in 4 set well in, that in 3 slightly in, that in 2 slightly out, with a double
spot in ib set in a little, the two rows completely separate and strongly orange-yellow. No
marginal marks. Hind wing basal area brownish black, but border blacker and in between
a white discal band widest at costa and gradually tapering to above anal angle but not crossing
the inner fold, though there represented by a greyish mark borders suffused with greenish blue
particularly on the outer border; submarginal row of linear white marks complete; marginal
border reddish to upper tail then olive to anal angle. Underside. Ground colour pale brownish

REVISIONAL NOTES ON AFRICAN CHARAXES 113

grey, more brownish between the two rows of spots representing the upperside pattern, and rusty
in the curve of the wing. The basal black marks are fine ; the discal and postdiscal spots buffy
to ochreous; the spot between the two rows in ib, black, so also the tornal marks in ib and space
above. There is an overall sheen to the wings. Hind wing ground colour tinged more brownish
over the disc; satiny bars present at sub-base; the discal band well represented and white in
colour; the postdiscal lunules not strong except in the lower part in the area of the tails where
it is greenish olive; the submarginal linear marks fairly strong; the marginal border reddish
to upper tail then olive to anal angle.

The whole underside has a rather cold greyish tone. (PL 9, figs. 76, 77.)

Variation, (a) Upperside. Colour and pattern very similar to the nominate form but the
fore wing spots slightly larger, those of the discal row large and extended distad so as to almost
make contact with the postdiscal spots in the same spaces, ib la, all the spots deep ochreous.
Hind wing discal band slightly narrower, bluish on borders; submarginal row of linear white
marks tending to fade out opposite tails; marginal border more strongly marked. Underside.
Much as in nominate form but with the exception of the brownish to rusty border to fore wing,
the whole underside of a colder tone with a satiny sheen overall. The pattern of spots whitish
to creamy. The hind wing rather browner at base which is crossed by a narrow satiny bar;
the outer border of the wing widely satiny. The discal band narrower, the postdiscal row of
lunules complete but obscurely represented in the brownish ground; submarginal line of pale
marks rather obscured by satiny sheen; marginal border broad and well marked.

Variation, (b) Upperside. Ground colour of fore wing as in (a) ; series of discal and post-
discal spots small and widely separated most noticable in areas la-ib. Postdiscal spots small
and ill defined. All spots deep ochreous. Hind wing white bar narrow throughout but going
through to inner fold; submarginal linear white marks strong. Underside. Darker in ground
colour especially on the distal half of the wings, the discal band of fore and hind wing whitish
and in strong contrast, but the postdiscal spots of fore wing less strongly marked.

Variation, (c) Upperside. Ground colour of fore wing brownish black at base but strongly
suffused over with rufous red on the distal half especially on the border. Discal and postdiscal
spots as in (a) strongly orange with very narrow rays joining the two series. Hind wing ground
colour brownish black, greyer on the inner fold ; discal band comparatively narrow, white with
greenish blue on both borders ; submarginal linear white marks strong ; marginal border narrow
but strongly orange red above upper tail, olive to anal angle. Underside. The whole surface
flushed over with rusty colour especially on border of fore wing and over the hind wing; fore
wing discal band ochre, the postdiscal spots orange rather suffused over by the rusty bloom.
The tornal black spot and one between spots in ib strongly black. Hind wing slightly more
greyish on distal border, the discal band rusty orange; the postdiscal lunules rusty brown with
slight greenish proximally; the submarginal row of linear marks though strong are partially
obscured ; marginal border narrow but distinctly reddish to upper tail then olive to anal angle.

$ form conjuncta van Someren & Jackson

Upperside. Fore wing ground colour as in form (c) slightly more rusty red on border; discal
and postdiscal spots orange, completely joined by rusty rays. Hind wing discal band mostly
white with faint bluish on borders; submarginal and border as in (c). Underside. Rusty
suffusion overall; fore wing conjoined marks dull orange, white in ib. Tornal black marks
strong. Hind wing discal band tapering rapidly; dull ochreous. Submarginal and marginal
ornamentation bold. (PI. 9, fig. 78.)

Range: The distribution of etheocles evansi is limited to the high country west
of the Rift Valley, centred mostly around Mt. Elgon and TransNzoia, but also
occurs at Nandi, Lumbwa, Londiani, Mau, Cherangani to Ravine and Tembach.
It thus overlaps some of the territory of Charaxes berkeleyi ssp. and also that of
Charaxes baileyi. (Map 4) .

H4 V. G. L. VAN SOMEREN

SYSTEMATIC LIST

Charaxes etheocles etheocles (Cramer)

Charaxes etheocles etheocles (Cramer) 1777. Type locality, Sierra Leone.
Region i. $ f . etheocles & vars.

alladinis Butler, 1869. Type locality, Ghana.
fulgens Rothschild, 1900.

regalis Rothschild, 1900. Type locality, Sierra Leone.
( f. ephyra Godart, 1823. " West Africa ".

hollandi Butler, 1893. Type locality, Sierra Leone.
carteri Butler, 1881. Type locality, Accra.
catochrous Staudinger, 1896. Cameroon.
Range: West Africa, Sierra Leone east to Cross River, Nigeria.

Charaxes etheocles biinclinata ssp. n.

etheocles biinclinata ssp. n. Type locality, Moyen Congo.

Region 2. $ f. etheocles pattern (two vars.).
ochracea pattern.
alladinis pattern.
regalis pattern.
<$ hollandi pattern,
carteri pattern.
catochrous pattern.

Range: Cameroon, Central African Republic, Moyen Congo, western Congo,
? northern Angola, Katanga.

Charaxes etheocles ochracea van Someren & Jackson.

etheocles ochracea van Someren & Jackson, 1957. Type locality, Gabon.
(etheocles f . ochracea Rothschild.)

Region 3. $ f . ochracea Rothschild, 1900. Type locality, Gabon.

ochreata van Someren & Jackson 1957. Type locality, Gabon.

alladinis x regalis.

seriata Rothschild, 1900. Type locality, Gabon.
<$ f. violacea Rothschild, 1900. Type locality, Ogowe Riv., Gabon.
Range: Gabon and adjoining territory Leopold area, lower Congo.

Charaxes etheocles carpenteri van Someren & Jackson

etheocles carpenteri van Someren & Jackson, 1957. Type locality, Kakindu

Hill, Kagera, Tanzania.
(etheocles f . carpenteri Poulton, 1919)

REVISIONAL NOTES ON AFRICAN CHARAXES 115

Region 4. $ f . carpenteri Poulton, 1919 et vars.

pallidimacula van Someren & Jackson 1957, et vars. Type

locality, Jinja, Uganda.
<$ f . carpenteri van Someren & Jackson,
vars. nr. carteri Butler.

nr. catochrous Staudinger and intermediates.

Range: S.W. of Congo Basin, Katanga and N.W. Rhodesia; Uganda, South Sudan,
adjoining area Ethiopia, N.W. Tanzania.

Charaxes etheocles evansi van Someren & Jackson

etheocles evansi van Someren & Jackson, 1957. Type locality, Elgon east,

TransNzoia.
(etheocles f . evansi van Someren & Rogers, 1932.)

Regions. ? f- evansi et vars.

conjuncta van Someren & Jackson, 1957.
$ f. evansi (nr. carteri} et vars.

Range: Kenya, N.E. and eastern areas of Mt. Elgon, TransNzoia to Mau Lunbwa,
Londiani, Sotik-Mara, Lembus and Ravine.

5. CHARAXES GRAHAMEI VAN SOMEREN
Charaxes grahamei sp. n.

(PI. 12, Text-figs. 7-8 [aedeagus])

In dealing with these specimens, one is faced with the problem of reaching a
decision based upon rather subtle evidence. Any student of the " etheocles "
group will admit that even where specific identity has been established beyond any
doubt, many of the males and females are so superficially alike that they can be
misidentified without close examination.

The facts in this particular case are :

1. The males are confusingly like <$ etheocles carpenteri on the upperside; on
the underside they resemble, to a certain degree, that of etheocles $ f . catochrous (a
form which occurs throughout the various subspecies).

2. The nominate female of grahamei superficially resembles members of the group
of ? etheocles, which embraces the forms alladinis, carpenteri, seriata and regalis
and vars., all of which are " western " with the exception of carpenteri, which is the
eastern representative of alladinis. No known form of carpenteri resembles gra-
hamei; all of them have the blue bar in the hind wing in the discal line. The form
regalis of western etheocles does however have the bar in the discal line, but this form
does not occur in eastern Africa.

3. Ch. etheocles carpenteri in its various forms occurs in the forests on the eastern
side of Lake Tanganyika, thus occupying the same habitats as grahamei. These
carpenteri <$ are of normal size.

4. Charaxes grahamei is always considerably larger, in both sexes. I do not
believe that this is a case of " giantism " amongst an otherwise normal population
of etheocles carpenteri.

n6 V. G. L. VAN SOMEREN

5. The genitalia of grahamei and catochrous have been dissected by T. G. Howarth.
They differ considerably (vide Text-figs. 2-4, 7-8).

On this evidence, I am satisfied that grahamei cannot be considered a " form " of
etheocles carp enter i, nor yet a subspecies of etheocles. The material examined consists
of nine males and eight females, all from the Mukuyu-Muhimo forests of the Kigoma
district, north-east of Lake Tanganyika.

MALE. Fore wing length 40 mm. (average 39 mm.) base to apex; 29 mm. along hind portion
of wing from base to end of vein i, (average 28 mm.). Comparable figures for etheocles carpen-
teri, 34-35 mm. and 27 mm. Shape as in etheocles but costa slightly more curved; curve on
outer margin appears stronger because of projecting tornal angle. Margin of hind wing more
serrate than etheocles carpenteri. Upperside. Ground colour deep velvety black, margin of fore
wing with very small internervular glaucous marks, double at tornal angle; edge with very
slight white fringe. Blue marks small, limited to one beyond end of cell in discal line ; (in some
specimens there is a minute dot at upper end of cell, a slightly larger spot beyond and a minute
subcostal spot at subapex) . Hind wing ground colour velvety black, slightly greyish on inner
fold; submarginal row of white spots, slightly blue distally, very distinct but well spaced;
marginal border narrow, greenish lunules indistinct except toward anal angle; margin black.
Tails rather short, 5 and 3 mm., black-edged with white mid line. Underside. Fore wing
ground colour silvery greyish, with rather strong satiny sheen; slightly brownish in interspace
between discal and postdiscal zones and on outer border; basal black spots and lines distinct
but not heavy; wavy black line between discal and submarginal zone distinct up to 2 then
fading out but with distinct black spot in ib; submarginal marks faint but strong and black
at tornus. Hind wing ground colour strong silvery grey up to discal line with a whitish zone
in this area; border slightly browner; postdiscal row of lunules silvery greenish and brownish,
outlined proximally in black, strong from costa to above anal angle; submarginal row of whitish
linear marks complete and accentuated by black marks in region of tails, double at anal
angle; marginal border reddish above upper tail then greenish to anal angle. (PI. 12, figs. 98,
99 type.)

A slight variant may exhibit a stronger brownish tone to the distal halves of both wings and
a slightly stronger pattern, especially on the hind wing on the underside.

The nominate female of grahamei is a blue-patterned form which resembles somewhat the
form regalis of etheocles.

FEMALE. Fore wing length 42 mm. Upperside. Ground colour blue-black with a tinge of
purplish and a blue sheen over basal half; discal row of spots purplish blue and complete from
the subcosta to hind margin, the spot in 4 minute but the rest increasing in size to the streak in
i a on the hind margin; postdiscal spots complete, three subapical in a row, rather whitish, the
remainder bluish to purplish blue, spot in 2-3 following the contour of the wing but that in ib
set in a little. Margin with faint indication of glaucous marks, double and more visible at
tornus; fringe narrowly white between veins. Hind wing ground colour blue-black with strong
blue sheen at basal half, distal border blacker. Discal band blue with slight greenish sheen,
inner border straight, spot at costa more whitish; a discreet spot at subcosta in the postdiscal
line; band widest at 6-7 then tapering toward the hind angle but not crossing the inner fold;
submarginal row of linear white marks strong; marginal border strongly orange-red to lower
tail then olive to anal angle. Tails long, 8 and 6 mm., black with reddish mid line. Margin
of wing black with fine white fringe, and slightly dentate. Underside. As in the male, the
underside is silvery grey with a slight brownish tinge to outer border of fore wing, and to a
lesser degree in the disco-postdiscal interspace ; the whole surface has a satiny sheen so that the
usual satiny bars are obscured. The black lines in the basal area to discal line very fine; the
discal and postdiscal rows of whiter spots not strongly defined; the usual submarginal dark spots
absent except for the double spot in ib which is black and strong as are the two spots in la-ib
more proximad in postdiscal line. Hind wing black lines in basal area very fine; discal bar only
slightly indicated ; the postdiscal wavy line of greyish green and maroon lunules rather lost in

REVISIONAL NOTES ON AFRICAN CHARAXES 117

the greyish ground except those above the anal angle ; marginal border reddish to base of upper
tail, then olive-grey to anal angle, the inner black marks small. (PI. 12, figs. 100, 101, type.)

Holotype male. TANZANIA: Kigoma District in Mukuyu Forest, 1964 (Major
I. Grahame). To be deposited in B.M. (N.H.).

Allotype female. Same locality, 1965 (Major /. Grahame).

A slight variant has the discal and postdiscal spots in the fore wing slightly larger; the marginal
glaucous marks larger. On the hind wing the blue bar is not so broad and the upper spots in
the postdiscal line are free. Otherwise the insect is very like the type. The underside is
silvery greyish with the pattern obscured.

Variation. Upper side. Well-marked, with large fore wing discal and postdiscal blue spots,
the lower marks in the discal line enlarged and extended distad and almost conjoined to the
spots in the postdiscal row by rays and blue scaling; there is an extra blue spot in la in the post-
discal row. The hind wing blue band is strongly blue in the discal line with slight purple tinge
in postdiscal line in 5-6. Underside. Silvery grey with a brownish tinge on the border of the
fore wing which offsets the whitish spots of the postdiscal row. The hind wing has a stronger
pattern than the type but the ground colour is silvery grey. (PI. 12, figs. 104, 105.)

$ form lacteata forma n.

This interesting form exhibits the general pattern of female " etheocles " , i.e. the fore and hind
wing light areas are white instead of blue. Fore wing length 42 mm. Upperside. Ground
colour black, slightly brownish in the basal area. There is a small white dot at upper end of
cell; the spots in the discal row strong and increase in size from the costa to the hind margin,
those in ia-2 extended distad and fused with the postdiscal spots in these areas; the two upper
postdiscal spots in the subapex are large, the third one smaller followed by spots of increasing
size to 2. All these spots are creamy white. Marginal marks are indistinct, whitish but more
distinct at tornal angle in ib. Hind wing brownish black at base, black along the border.
The discal band is very broad, extending from the costa to 3 then tapering to the inner fold
above the anal angle, colour creamy with some bluish scaling on the inner border and more
strongly bluish on the lower half of the outer border. Submarginal row of linear white marks
strong; marginal border ochre-red to base of lower tail then olive to anal angle; edge black with
slight white fringe between tails; tails moderately long upper 7 mm., lower 5 mm. Underside.
Fore wing ground colour light brownish grey, slightly more brownish between discal and post-
discal zones and on the outer border. Basal black marks not heavy but discal and postdiscal
spots clear and whitish; submarginal dark marks strong at tornus and area above. Hind wing
basal area light brownish grey followed by a more brownish bar on proximal side of the well
marked discal creamy band; the postdiscal row of lunules brownish slightly greenish ochre
above anal angle; submarginal linear white marks strong, with usual black dots toward anal
angle; marginal border reddish to base of upper tail then olive to anal angle. (PI. 12, figs. 102,
103, type.)

Holotype female. TANZANIA: Kigoma district, Mukuyu Forest, 1965. Presented
to the B.M. (N.H.) by Major Iain Grahame.

Variation. Upperside. Ground colour of fore wing with rather more purplish blue sheen;
discal and postdiscal spots arranged as in nominate form but less fused in hind area ia-2;
where spots are tinged with violet-blue, others creamy white. Hind wing discal band broad,
rather irregular on outer border, white, with some bluish on lower half of outer border. Marginal
border as in nominate. Underside. Ground colour greyish brown with a slight overall rusty
ochre tinge; the fore wing spots of both series ochre; only one black spot at tornus with pale
surround. Hind wing ground colour as fore wing, the discal band rather suffused over with
greyish but whiter in subcostal area; row of lunules in postdiscal zone only slightly indicated;

n8

V. G. L. VAN SOMEREN

Mocambique

MAP 5.

REVISIONAL NOTES ON AFRICAN CHARAXES 119

submarginal linear marks subdued; marginal border obscured, dull reddish to upper tail then
olive to anal angle.

Range. This species is found in the forests of the Kigoma district, north-east
of Lake Tanganyika, extending southward to the forest of Kungwe Mt. in the
Mpanda district.

It is to be noted that Char axes etheodes carpenteri also occurs within this range,
the male forms carpenteri and catochrous having been taken in considerable numbers.
(Maps 4, 5).

6. CHARAXES CONTRARIUS WEYMER AND C. PETERSI SP. N.
Charaxes contrarius Weymer
(PI. 13, Text-figs. 27, 28 [aedeagus])

Charaxes contrarius Weymer, 1907 : 44.

Charaxes subargentea van Someren & Rogers, 1932 : 167.

Charaxes catochrous contrarius Weymer; van Someren & Jackson, 1952 : 279.

When I described Charaxes subargentea in 1932, I did so on the assurance of the
late Dr. Karl Jordan that the species had not been described previously, but we had
both overlooked Weymer's contrarius, published in 1907. The subsequent association
of contrarius with catochrous by van Someren & Jackson in 1952 was most unfortu-
nate, for catochrous is but a male form of etheodes (vide Ch. etheodes (Cramer) p. 99).
The differences between catochrous and contrarius were clearly set out by us then;
beyond the somewhat silvery underside exhibited by both, there is no real resem-
blance between the two. No subspecies of etheodes is known to occur within a
thousand miles of the range of contrarius, the intervening country being unsuitable
to etheodes.

The recently described Charaxes martini van Someren, 1965, from southern
Malawi, may be related to contrarius, but insufficient material, especially authentic
females of this southern insect, precludes any definite decision on this point.

MALE. Fore wing length 30-40 mm. Shape incurved on outer border at 3-4, apex falcate,
thus very like Charaxes viola kirki. Upperside. Ground colour blue-black with strong blue
sheen at base of costa. Blue spots in subcostal region slightly variable in number and size,
but there is usually a spot at the upper part of end of cell, one large spot with a small one below
in the discal line, two blue spots in the subapex; margin with distinct glaucous lunules, often
strong, and with a white streak in centre of each. Hind wing ground colour blue-black, greyer
on the inner fold ; a slight greenish sheen sometimes present over lower part of disc sometimes
represented by an indistinct wavy greenish line opposite tails; submarginal row of short linear
white marks distinct ; marginal border with maroon lunules outlined in olive, then olive to anal
angle usually strong, but may be obscured, edge black, tails long, of about equal length, 5 mm.,
black with central olive-red streak. Underside. Fore wing ground colour silvery greyish to
discal line, slightly browner beyond to the postdiscal series of greyish spots, outlined proximally
in areas ia-6 in blackish, followed by the submarginal row of darker diffuse marks which are
black in ib at the tornus. Basal black marks well denned but thin. Hind wing basal area
silvery, more whitish toward the inner fold; basal black lines thin; ground colour more tinged
with brownish in the postdiscal line which has conspicuous olive and maroon lunules accentuated
proximally in black ; submarginal row of linear marks rather broad in the type but usually well

120 V. G. L. VAN SOMEREN

defined and outlined distally in black, particularly in region of tails ; border brick-red above upper
tail then olive to anal angle. (PI. 13, figs. 106, 107, type.)

Variations. Upper side. The size and number of blue spots on the fore wing reduced and
the marginal lunules smaller. On the hind wing the submarginal spots are small and partially
obscured while the marginal border is narrow, broken and mainly greenish. Underside. The
fore wing distal pattern is stronger while on the hind wing, the band carrying the postdiscal
lunules is strongly brownish ; the submarginal linear pale marks continuous and not interrupted ;
the marginal border is strongly reddish above upper tail then olive to anal angle. PI. 13, fig. 108.

Female. There are two distinct patterns of female: (a) those with an" etheodes-
like " pattern, and (b) those with a " rosae-like " pattern. The former predominates.

(a) $ form contrarius, nominate form

Size rather variable, fore wing length 35-42 mm., average 38 mm. In shape the fore wing
may be strongly falcate (as in PI. 13, fig. in) or the outer margin may be less incurved at 3-4 as
in the majority. Upper side. Ground colour blue-black or slightly brownish black, with blue
sheen towards the base of fore wing. There may be a white spot at upper angle of end of cell,
more often missing, seldom large; the discal row of spots as follows: two subcostal, the upper
large, quadrate-elongate, usually no spot sub-base in 4, but when present is often large; the
spots from 3 to hind margin increasing in size, the increase usually on the distal end, these marks
are usually white. The spots in the postdiscal row are three subapical in a row of about equal
size, spot in 4, sometimes missing, is set in, those of 3 to 2 may be free or slightly joined to discal
mark in same area, that of ib fused with discal mark. Margin with no marks as a rule but may
be very faintly indicated at tornus. Hind wing basal area blue- or brownish black, slightly
greyer in the inner fold, border black; discal white band broad, commencing at costa and
widening opposite cell, then gradually diminishing and tapering to above the anal and crossing
the inner fold; the band may be slightly blue-tinged on borders from mid point to anal angle;
the submarginal line of linear bluish white marks distinct and well separated; border reddish
to upper tail, then olive to anal angle; margin black. Tails long and thin, upper 8 mm., lower
56 mm., black with reddish or olive mid line. The fore wing white discal bar is the main
feature of the wing and may widen abruptly from 3 to hind margin. (PI. 13, figs. 109-112.)

Underside. The ground colour in all these forms is silvery greyish, slightly tinged with
brownish in the curve of the fore wing. The basal black lines are fine but distinct; the discal
bars are whitish, the upper postdiscal spots buffish; the submarginal dark marks strongly
represented in the tornus at ib and in an area above and are black and touch the postdiscal
spots in these areas. The hind wing ground colour is satiny silvery grey with a slight brownish
tinge in the base; the discal white band shows up strongly but reduced in width and tapering
rapidly toward the inner fold which is crossed above the anal angle. The postdiscal bar of
olive and maroon lunules is very strong except toward the costs. The border is slightly more
brownish and the submarginal row of linear white to lilac marks are continuous and strong,
accentuated distally by black marks; the marginal border strongly reddish to upper tail, then
olive to anal angle.

Variation, (a) Upperside. The fore wing discal and postdiscal spots in the upper section
may be creamy, creamy ochreous or ochreous.

Variation, (b) Upperside. The upper fore wing spots may be slightly conjoined by rays;
the spots are white.

Variation, (c) Upperside. A rather striking variant in which the ground colour is a deep
blue-black; there is a well developed white spot in the fore wing cell; the white marks from 3
to hind margin fully conjoined, the marks in la-ib bluish at the ends; the hind wing band, white
in the mid area, is strongly bluish on both borders, especially on the proximal border. (PI. 13,
fig. in.)

Variation, (d) Upperside. This is an extreme of (b) in which all the wing spots are fused
from 5 to ib, the upper spots larger than usual. The hind wing band is exceptionally broad
and expanded proximally and strongly blue on the inner border.

REVISIONAL NOTES ON AFRICAN CHARAXES 121

(b) form conjugens forma n.

(PL 13, fig. no)
$ form rosae pattern (PL 13, fig. 112)

Size rather variable, fore wing length 35-44 mm. Shape similar to other forms. Upperside.
Ground colour of fore wing brownish black, sometimes with a slight purplish sheen at base.
The conspicuous white curved band commences as a white spot at the upper part of the cell
(it is of interest that a spot in the cell is unusual in the nominate form) which is continuous
with a larger triangular mark beyond at base of 4, and this, with a larger triangular mark at
base of 3, an elongate mark in 2 with obliquely cut proximal end, followed by elongate quadrate
mark in ib and streak in la completes the band. The upper discal and postdiscal spots are
usually large and free, but may be small. Marginal glaucous or greyish marks hardly visible
except at tornus. Hind wing basal area brownish black, greyer on the inner fold. The discal
white band commencing at the costa, expands over the mid-region then tapers to the inner
border and crosses it above the anal angle as a narrow white mark; the band is mainly white
but may have some bluish scaling on the borders especially at the lower half. Border of wing
black and broad, the submarginal linear white marks distinct or slightly obscured opposite
tails, but here bordered by black distally; marginal border reddish to upper tail then olive to
anal angle; margin black; tails long and thin, 8 and 6 mm. long, black edged with olive or whitish
mid-line. Underside. Ground colour slightly variable silvery greyish or with varying amount
of pale brownish suffusion. Fore wing black basal marks thin out distinct; pattern of upperside
strongly represented; tornal black mark and one above very strong. Hind wing silvery grey
at base, browner on border, the discal band strongly marked but narrower than above and tend-
ing to fade out above anal angle, but postdiscal series of olive and maroon lunules strong;
submarginal row of linear marks usually strong; with strong black distally especially in region
of tails; marginal border reddish above upper tail then olive to anal angle. (PI. 13, fig. 112.)

Variation. An occasional specimen may have on the upperside a fore wing bar of more even
width and the white scaling extending onto the costa. The hind wing band however is limited
at the costal end to a large free spot in the discal line, and a reduced mark in 5 ; the lower part
of the band strongly flushed with bluish.

Range: Kenya, occuring in the coastal forests and heavy savanna from just north
of Lamu in the Mundane range to the Sekoke-Arabuko forest, the Rabai Hills
south to the Shimba Hills and the coastal forests at Zombo and Marima Hill in the
Kwale district. Tanzania, the lower forests of the Usambara range, and the Pugu
forest, south to Lindi. (Maps 4, 5).

Charaxes petersi sp. n.

(PL 13, figs. 113, 114; Text-fig. 25 [aedeagus])

This interesting new species was first taken by Dr. W. Peters in Liberia at Kitoma,
i, 600 ft. in August of 1953, and remained unrecognized until 1965 when Dr. Arthur
Rydon made dissections of the genitalia of the type and other specimens. All had
genitalia which differed markedly from those of Charaxes etheocles, which is common
in the area. The species was taken subsequently by Monsieur A. Gallay in the
Ivory Coast at Abidjan; these specimens have similar distinctive genitalia. Unfor-
tunately, all examples are males; the female is not known with certainty.

122 V. G. L. VAN SOMEREN

This species comes within the " etheocles " s.l. aggregate, and indeed is very similar
in general appearance, especially on the underside, to the catachrous form of etheocles
etheocles, and could easily be mistaken for it.

MALE. Fore wing length 30 mm., (type), 29 mm., paratypes. Upperside. General appear-
ance very similar to male etheocles etheocles (Cramer), deep velvety black with slight greenish
sheen at base of the fore wing costa. Fore wing immaculate except for an obscure blue subcostal
spot toward end of the cell, a smaller blue spot at base of 6, two bluish white spots at mid 8-7
and a small white subapical dot in 8. Margin with very obscure glaucous lunules and slight
white fringe between veins. Hind wing ground colour as fore, slightly browner on inner fold;
disc immaculate but submargin with small punctiform white dots, double at anal angle ; margin
with glaucous lunules, edge black with slight white fringe. Tails short, 4 and 3 mm. long, black
with central glaucous streak. Underside. General appearance very similar to the catachrous
form of etheocles, but base of fore wing with more pronounced silvery satiny sheen; discal bar
silvery edged internally with black crescentic marks; postdiscal lunules silvery, inwardly
margined in black and distally with series of rounded dark spots, strongest at tornus, that at
hind angle and in ib, ringed greyish; margin brownish. Hind wing basal area greyish brown
with fine wavy black lines in sub-base; discal bar strongly silvery from costa to above anal
angle where it crosses the inner fold, bordered distally by a brownish zone; the postdiscal
greyish and brownish ocelli well marked; admargin with whitish lunules accentuated distally
with black, most apparent in area between tails and at anal angle ; margin black with very slight
white fringe.

For formation of the aedeagus, vide Text-fig. 25, it will be noted that the main " tooth "or
spine is extremely large and prominent, and is situated on the ventral or ventro-lateral side
of the shaft of the penis.

Holotype male. LIBERIA: Kitoma, 1,600 ft., viii.1953 (W. Peters}. In B.M.
(N.H.).

Paratypes. Same data, in B.M. (N.H.), and one male from IVORY COAST in
National Museum, Nairobi, Kenya. Other males from Ivory Coast in coll. Gallay.

Range: Liberia, in forested areas at Kitoma; also on the Ivory Coast, in forest
at Abidjan. The species has not been bred.

7. CHARAXES BAILEYI VAN SOMEREN

Char axes bailey i van Someren
(PI. 14, Text-figs. 13-14 [aedeagus])
Char axes bailey i van Someren, 1958 : 4.

This insect was referred to very briefly by van Someren & Jackson (1952 : 285).
There were two female specimens of uncertain status, one from Ravine, the other
from Rongai on the western side of the Rift Valley.

The species was subsequently taken at an intermediate locality, Visoi Gap, by
J. H. Bailey. This was a female with a broad blue band on the upperside of the
hind wing, similar to the first specimen taken at Ravine. When on a visit to the
Visoi area, I was successful in obtaining a few females from which I raised families
with sufficient progeny to indicate the constancy of both male and female characters.

REVISIONAL NOTES ON AFRICAN CHARAXES 123

Several additional males and females have been secured by Mr. Bailey during the
past year or so.

MALE. Fore wing length 35-37 mm. with only a very slight inward curve on outer border.
Upper side. Ground colour intense blue black with slight blue sheen at base; glaucous marginal
marks triangular or lunate usually well marked; subcostal blue spots, one minute dot at upper
end of cell, a larger spot in discal line, a small dot in upper subapex. Hind wing ground colour
intense black with some greenish sheen toward postdiscal zone where there may be the slightest
trace of a greenish zigzag bar opposite tails, but usually absent; submarginal row of linear
white marks distinct, well separated; marginal border greenish, more olive at anal angle;
extreme edge black with slight white fringe. Underside. Fore wing ground colour greyish
brown, browner in zone between discal and postdiscal lines, more so in slight curve of the wing
on outer border, but the satiny sheen is strong over the discal zone and toward the subapex;
the pattern is thus rather strong ; the submarginal dark spots, ringed in greyish, black at tornus
then more brownish to subapex. Hind wing ground colour as fore with a slight olive tinge;
sub-basal and discal satiny bars strong; the postdiscal wavy line of lunate marks olive and
maroon, outlined proximad in black; submarginal series of pale linear marks well marked.
(PL 14, figs. 115, 118, holotype.)

There is little variation in the males on the upperside; in the fore wing the subcostal blue
spots may be reduced in size and number; on the hind wing the submarginal linear marks may
be obscured. On the underside, the pattern may be less strong but the satiny zones are still
marked; the postdiscal line may be bronzy. The shape of the hind wing in baileyi is more
rounded and less pyriform than in berkeleyi which flies with it. I selected as the type female
of nominate baileyi, the form predominating with a broad discal band in the hind wing.

FEMALE. Fore wing 38-40 mm. Upperside. Ground colour deep blue-black with strong
blue sheen at base of wing. There is a suggestion of a blue spot at the upper end of the cell;
the discal spots are one large subcostal with a dot below, followed by a small spot sub-basal
in 4 set well in, two larger spots in 2 and 3, and a suggestion of blue scaling in ib; the post-
discal row of spots well marked and blue, three subapical in line, followed by spots in 4-2 slightly
larger and set in, the mark in la and ib larger and bluer, that in la, a streak; margin with
small glaucous marks, largest in ib at tornus. Hind wing ground colour as forewing; a broad
blue band crosses the disc of the wing, represented in subcosta by a free quadrate spot in post-
discal line, the band tapering to the inner fold above the anal angle. Beyond this band a black
border carrying the complete series of crescentic blue and white marks, double and more bluish
at anal angle; marginal border reddish and green slightly outlined in mauve; extreme edge black
with narrow white fringe. Tails rather long, upper 7 mm., lower 5 mm., black edged with
reddish median line. Underside. Basically similar to that of the male as regards pattern,
but the discal and postdiscal spots of upperside are represented by buffish spots and the whole
ground colour is paler and browner but the dark spots toward the hind angle are strong, particu-
larly that at the tornal angle. On the hind wing the pattern is subdued in the basal area; the
postdiscal line is not strong except opposite the lower tail and above this there is a somewhat
quadrate pale mark on the inner fold; the submarginal linear lilac marks are well developed,
the marginal border of reddish to upper tail then olive to anal angle is rather narrow. (PL 14,
figs. 116, 119, allotype.)

There are several variations of this form, (a) Upperside. The fore wing spots are mainly
bluish, the marks in the discal line not extending to the hind margin in la-ib, the postdiscal
series extending to ib as a blue streak. The hind wing blue band as in the nominate form, the
submarginal spots blue and the marginal border strongly reddish to lower tail then olive to
anal angle accentuated proximally in green. Underside. Ground colour more rufescent,
especially on the outer border of fore wing; the hind wing more rusty, and the postdiscal
line of lunules almost entirely reddish.

Variation, (b) Upperside. Fore wing upper discal spots are white, but strongly blue in
la-ib, the marks coalescing with the large blue marks in la-ib of the postdiscal series; the
margin of the wing with diffuse glaucous marks. The hind wing discal blue band broad and

ENTOM. 23, 4. "

124 v - G - L - VAN SOMEREN

complete to the costa; the submarginal bluish white marks large; the border greenish with a
few reddish scales above the upper tail. Underside. As the nominate form but discal spots
in fore wing from 2 costa white.

Variation, (c) Upper side. Fore wing spots in the upper sector in both discal and postdiscal
rows are small and obscured but those in la-ib and slightly in 2 conjoined and strongly blue.
The hind wing band broad and complete to costa; the submarginal linear marks strongly blue
and in marginal border green, slightly more olive at anal angle. Underside. As in nominate
form. These forms occur in bred material from the type locality.

Variation, (d) Merely a colour variant in which the blue spots and hind wing band of upper-
side are strongly greenish blue (Mau-Narok.)

$ form pseudocarpenteri van Someren, 1958 : 5

This form bears a superficial resemblance to female f. carpenteri of etheocles carpenteri, but
the two do not fly together. The length of fore wing is 40-42 mm. Upperside. Ground
colour blue-black with purplish brown shading toward the base, sometimes shot with green.
The spots in the discal zone are limited to those in subcosta to area 2 and these are white or
slightly bluish white; the postdiscal series complete and ending in a streak in la, the subapical
spots white, the remainder blue. The blue band on the hind wing is limited to the postdiscal
zone and extends from the subcosta to just above the anal angle, the mark in 6 is set in so that
there is a double kink in the bar; the submarginal linear marks are bold and bluish white; the
marginal border reddish above upper tail and olive to anal angle. Tails as in nominate form.
Underside. Ground colour purplish grey-brown with a golden olive bloom over the base of the
fore wing and on the border; the pattern very similar to the nominate form but whitish marks
in discal area limited to spots above. Hind wing ground colour as fore but golden olive bloom
covers the whole of the disc, but is traversed by satiny bars in sub-base and discal line. Submar-
ginal line of pale marks strong but rather fused in region of upper tail, spots bordered by olive
marks distally, double in anal angle; border reddish to upper tail then olive to anal angle.
A quadrate whitish grey mark is present at inner fold above the dark postdiscal mark, the rest
of the postdiscal lunate marks not strong, reddish olive.

Variation, (a) Upperside. Similar to typical, but all fore wing spots blue; the postdiscal
blue bar in hind wing bordered with bronzy golden scales. (PL 14, figs. 117, 120.)

Variation, (b) A paratype specimen of this form has the Upperside ground colour of fore
wing blue-black, slightly bluer toward base; postdiscal spots reduced in number and the post-
discal series present but obscured, the mark in la elongate along the hind border. The hind
wing ground colour blue-black shaded with purplish brown towards the base; the blue band is
limited to the postdiscal area but does not reach the costa; the black border carries a row of
linear bluish white marks extending from upper angle to anal angle; the marginal border is a
mixture of greenish and red scales to upper tail then olive to anal angle. Tails as in typical
form.

Bionomics : A full account of the biology of this species can be found in the original
description. (1958 15) The species lays on Scutia myrtinaBurm. s.l. (Rhamnaceae).
Families bred from $ form baileyi produce both baileyi and $ form pseudocarpenteri ;
and pseudocarpenteri also produces both forms; variants appear in some families.
The males are remarkably constant.

Range: So far, this species has only been taken west of the Rift Valley in Kenya
at Visoi Gap between Kilombe Hill and Londiani Hill, at Molo and Ravine, at
Mau-Narok, and recently on the eastern slopes of Mt. Elgon. Thus there is a
considerable overlap with Charaxes berkeleyi ssp. and Charaxes etheocles evansi.
(Map i).

REVISIONAL NOTES ON AFRICAN CHARAXES 125

SYSTEMATIC LIST

Charaxes grahamei sp. n.

Char axes grahamei sp. n. Type locality: Tanzania, Kigoma Dist., Mukuyu Forest.
Range: Tanzania, Kigoma Dist., N.E. of Lake Tanganyika and
southwards to Mpanda Dist., Kungwe Mt.
$ f. lacteata forma n. Type locality: Kigoma Dist., Mukuyu Forest.

Charaxes contrarius Weymer

Charaxes contrarius Weymer, 1907. Type locality: " German E. Africa ". Range:
Kenya, coastal forests and heavy savanna from Mundane Range,
Lamu to Sekoke to Arabuko Forest, Rabai Hills, Shima Hills,
Zombo, Marima Hills, Kwale Dist., Tanzania, Usambara Range
and Rugu Forest, south of Lindi.
$ f. conjugens forma n. Type locality: Shimba Hills, Kwale.

Charaxes petersi sp. n.

Charaxes petersi sp. n. Type locality: Liberia, Kitoma. Range: Liberia and Ivory
Coast.

Charaxes baileyi van Someren

Charaxes baileyi van Someren, 1958. Type locality: Kenya, Visoi Gap. Range:
Kenya, west of Rift Valley at Visoi Gap, Kilombe Hill, Molo,
Ravine, Mau-Narok and Mt. Elgon.

$ f. pseudocarpenteri van Someren, 1958. Type locality: Kenya, Visoi Gap,
Kilombe Hill.

8. CHARAXES VIOLA BUTLER AND ITS SUBSPECIES AND FORMS

Charaxes viola Butler

Butler described this insect as a species in 1865. Subsequently, Rothschild and
Jordan (1900) referred to viola as a subspecies of etheocles (Cramer), 1777. Auri-
villius (1925) and subsequent authors followed the general classification advocated
by Rothschild.

Van Someren and Rogers (1925 : 42-43) drew attention to certain breeding experi-
ments which seemed to indicate that " etheocles ", as denned by Rothschild, was an
aggregate of several well denned species. This view was upheld and amplified by
van Someren and Jackson (1952 : 257-259), and was followed by a tentative division
of the group into species, being further amplified by us in 1957. Charaxes viola
Butler was reinstated to specific rank and several subspecies were recognized. The
present revision carries the investigation still further, and is based on a great deal
of additional material and data.

This species, throughout its range and in all its racial forms, is an insect of the

126 V. G. L. VAN SOMEREN

savanna country, i.e. " thorn-bush " or " orchard country ", light riverine gallery
forest, and dry forest areas, thus seldom extending into areas of heavy forest.
Char axes etheocles on the other hand, is generally an insect of heavy forest.

If one studies a " Vegetational Map " such as that of UNESCO, 1959, it will be
noted that areas of various types of savanna or woodland extend from W. Africa to
Abyssinia and Somalia, eastern Africa to Central Africa and to S. Africa north of
Lat. 25 with varying degrees of tree coverage; Zones 8, 16, 18, 20, 21-22, and 25.
The known ranges of Charaxes viola correspond more or less with these areas which
are described as: Forest-savanna, mosaic woodland, savanna woodlands, wooded
steppe, etc., the main trees being Acacia, Combretum, Commiphora, Terminalia,
Brachystygia, Albizia, Entada and Isoberlina. Many species of some of these genera
are food plants of Charaxes viola,

This type of habitat being more or less continuous, either as large blocks or even
narrow corridors, from west to east and east to south with no natural ecological
barriers completely severing the continuity, we find some evidence of merging of
the subspecies into which viola has been divided. One is thus faced with the problem
of deciding which of these races is taxonomically sound, i.e. do they represent
restricted or isolated breeding communities or are some bio-ecological strains affected
by climatic conditions and restricted food plants, yet exhibiting evidence of consider-
able genetical control of female forms. There is often a size difference.

On the evidence of the considerable material before me, Charaxes viola can be
divided into four main groups, based on female forms (i) a western group (W.
Africa to Uganda) in which the females conform to a fixed pattern and coloration,
with comparative slight variation; (2) a north-eastern group (Ethiopia-Kenya-
Tanzania) in which the female pattern remains more or less constant, but the colour
varies from white to orange; (3) a central southern group (western Tanzania-Malawi-
eastern Zambia- Rhodesia in which the female is of totally different pattern and colour
but in which the white-barred female crops up again, but mostly in the southern
parts of its range ; (3a) S.W. Africa in which a boldly marked, white-banded $ is the
only one recorded; (4) a southern Congo (Katanga), western Zambia- Angolan group
in which the females are highly polymorphic and polychromatic.

One is led to speculate on what was the ancestral female form. By analogy, one
would expect that the original female was somewhat male-like, the first major diver-
gence being the development of discal and postdiscal spots, already present in males,
most apparent on the underside. These two series of spots, originally bluish or
whitish, became creamy, ochreous or orange; at first completely separated, they
became partially united in the lower or hind portion of the fore wing, and in the disc
of the hind wing. The appearance of form phaeus is difficult to explain, but if one
studies the female forms of the polymorphic group 4, the possible development of this
form is sensed; the greenish blue basal area present in the male became gradually
intensified or became paler blue, the postdiscal spots developing concurrently with
some trace of discal spots. If there is any substance in this speculation, one must
assume that the polymorphic aggregates are still in a state of flux of evolutionary
development, whilst the monomorphic have evolved the more rapidly and achieved
stability.

REVISIONAL NOTES ON AFRICAN CHARAXES 127

I deliberately avoid the controversial subject of mimicry as an agent in the
evolution of Char axes viola; it abounds in pitfalls.

Since this is a taxonomic paper one should be able to define, with some degree of
certainty and accuracy, the characters of the subspecies and state their distribution.
In the main, these criteria can be fulfilled, but there is some evidence of apparent
overlap, even merging, of aggregates to which subspecific names have been applied.
There is also a paucity of material and detailed field data from some areas. I
prefer, at this juncture to retain the taxa as given in the systematic list.

Charaxes viola viola Butler

(PI. 15, figs. 121-132)

Charaxes viola Butler, 1865 : 627. (Partim) Type female.

Charaxes ephyra Feistamel, 1850 : 255 (<J).

Charaxes viola Butler, 1876 : 481 ($}.

Charaxes etheocles viola Rothschild, 1900 : 488 ($).

Charaxes etheocles viola Aurivillius in Seitz, 1912 : 137.

Charaxes viola viola Butler ; van Someren & Jackson, 1952 : 263.

GROUP i, AREA i. SENEGAL N. NIGERIA
Females are monomorphic with only slight variation.

MALE. Fore wing length 33 mm., shape falcate. Upperside. Ground colour blue-black
with greenish sheen especially at base of cell. A blue spot in cell, two blue spots beyond, upper
large, lower a mere dot, two subapical bluish white spots, (occasionally only one), or sometimes
three, continued in postdiscal line as faint blue dots from 4 to hind angle; margin with strong
glaucous lunules increasing in size to tornus, slightly separated by dark veins. Hind wing
black, greyer on inner fold, disc immaculate, but in postdiscal zone a wavy bluish green line,
most pronounced opposite tails and fading out toward upper angle; submarginal linear marks
whitish in upper part, then bluish to anal angle; marginal border greenish ochre with reddish
centres, becoming more greenish to lower tail, then olive to anal angle; margin black. Tails
rather short, tapering rapidly to a point, upper 4 mm., lower 3 mm. Underside. Ground
colour greyish drab, slightly more brownish on border, or generally pale earthy brown overall.
Fore wing black marks strongest in lower part of basal triangle in ib; satiny bars faintly indi-
cated in discal line, internally outlined in black especially towards the hind margin ; postdiscal
line proximally faintly outlined in black; submargin with a series of dark marks from apex to
tornus increasing in size and culminating in double black tornal mark. Hind wing with an
indication of a pale sub-basal bar and an irregular pale bar in discal line ; basal black lines very
fine and hardly indicated; postdiscal series of conjoined lunate marks greenish ochre and
reddish, outlined in black proximally, extends from costa to above anal angle; submarginal
whitish lilac marks not strongly developed ; marginal border reddish above upper tail then olive
to anal angle, with black central dots double at anal angle. (PI. 15, figs. 123, 124.)

FEMALE. Fore wing length 40 mm., shape falcate, generally narrow, since the length of
vein i seldom exceeds 24 mm. Upperside. Ground colour brownish black, slightly browner
on the outer border. All fore wing spots are tawny orange except those in la-ib which are
pale. Two spots beyond end of cell, usually no spot basad in 4, but those in 3 to hind margin
gradually increasing in size, their inner ends rounded, the outer ends continuous with the post-
discal but usually with some dark scaling indicating line of junction; postdiscal spots: three
ovoid in subapex, spots from 4 to hind margin increasing in size and somewhat quadrate are
contiguous with the discal row. Margin with conspicuous though ill-defined tawny orange

ia8 V. G. L. VAN SOMEREN

lunules, most clear from 4 to ib. Hind wing basal triangle brownish black, disc of wing with a
broad white patch slightly bluish on borders, the band goes through from the costa where it is
narrow, then widens and fills the greater part of the disc, then tapers to and crosses the inner
fold above the anal angle. The distal dark border is thus narrow, widest in 6-7 but narrowing
to anal angle. The submarginal linear or lunate marks are whitish blue, more blue toward
anal angle, and conspicuous. Marginal border reddish above upper tail then olive to anal
angle, with black marks and double dots at anal angle. Margin black tails long and slender,
upper 6-7 mm., lower 5 mm., black edged with ochre-olive mid line. Underside. Fore wing
ground colour palish grey-brown in basal area, slightly more rusty toward curve of outer border ;
black marks strongest in lower area ib and along the inner border of the tawny ochreous discal
and postdiscal band, the inner border of the latter with subdued dark spots in submarginal
line which become black at tornus at ib and area above, the former accentuated by a pale lilac
surround. Hind wing colour as fore, the basal black lines very thin; the discal band is ochreous,
bordered distally by the postdiscal zigzag ochreous and reddish lunules which extend from
costa to anal angle. The ground colour of the border is slightly greyish, traversed by the com-
plete row of whitish lunate marks, slightly black lined distally, then strongly black opposite the
tails; marginal border rather narrow, reddish above upper tail then olive to anal angle. (PI. 15,
figs. 121, 122.)

Variation, (a) Upperside. In the fore wing the tawny orange discal and postdiscal spots
are conjoined to 5, there being a spot in 4 in discal row; the outer border of wing strongly rufous
from apex to hind angle so that the lunules on the margin are fused. Underside. As in nominate
form.

Variation, (b) Upperside. Fore wing discal and postdiscal spots well separated from 7-4
then almost separated by black scaling in 3-2. Marginal lunules not very distinct. Hind wing
dark border divided into triangles by angular intrusion of the discal white band into it, which
almost meet the submarginal row of linear marks. Underside. Pale greyish putty colour, with
barely indicated black marks, only the tornal black mark distinct. The discal bands only
slightly visible and suffused over.

Range: Nominate viola extends from Senegal to Togoland and Ghana thence to
northern Nigeria and perhaps to Lake Chad, inhabiting the Savanna bush country.
I am unable to check the records of viola from Atbara, as the specimens appear to
hav e been lost ! (Map 6) .

Charaxes viola picta van Someren & Jackson
(PI. 15, figs. 127-130)

Charaxes etheocles <$ f. picta Rothschild, 1900 : 483.

Charaxes etheocles <$ f. picta Rothschild; Aurivillius, 1912 : 553.

Charaxes etheocles $ f. vansomereni Poulton, 1925 : 553.

Charaxes etheocles $ f. illuminata and f. splendens Rousseau-Decelle, 1938 : 161. Locality, Rio

Luvna, E. Congo Beige.
Charaxes viola picta van Someren & Jackson, 1952 : 263.

GROUP i, AREA 2. S. NIGERIA, CAMEROON, THEN SKIRTING NORTH OF EQUATORIAL
FOREST TO E. CONGO, UGANDA, S. SUDAN AND N.W. KENYA

The females are monomorphic with slight variation.

MALE. Fore wing length 33-35 mm. (average 35 mm.), thus larger than the nominate race,
and more robust ; shape falcate. Upperside. Ground colour black with greenish sheen especially
over the cell. This race is usually more strongly marked with blue spots than in nominate: a
large blue spot in the cell, in discal row beyond, a large subcostal whitish spot with a smaller

REVISIONAL NOTES ON AFRICAN CHARAXES 129

below, traces of blue spots in 4-2 ; postdiscal spots, three in a row in subapex, usually white or
with slight ochreous tinge, followed by two to four small blue spots, the lower ones fading out ;
marginal glaucous lunules broad and strong, separated by dark veins. Hind wing ground colour
as fore, greyer on inner fold; disc immaculate, but postdiscal wavy greenish line present in
lower half, less clear in 5-6 ; submarginal bluish white marks complete but often small ; marginal
border reddish above upper tail, then olive to anal angle which has two black spots. Tails
rather short and of equal length 4 mm. Underside. Darker than nominate race with black
marks at base of fore wing more pronounced, so also the satiny bars, which are bordered in lower
half by black; tornal mark strong. Hind wing ground colour slightly more brownish, satiny
discal band more distinct; postdiscal ochreous olive and reddish lunules, outlined proximally
in black stronger; marginal lunules reddish above upper tail, then olive to anal angle; black
dots strong, double at anal angle. (PI. 15, figs. 127, 128, type.)

Variation, (a) Upperside. Very similar to typical but with discal and postdiscal blue marks
stronger and distinct; the subapical spots ochreous. Hind wing with strong postdiscal wavy
green line. Underside. More strongly marked.

Variation, (b) Upperside. A variant in the reverse direction: the subapical spots limited
to two small blue dots; the glaucous marginal lunules strong. Hind wing postdiscal greeny
line hardly visible; marginal border with hardly and red scaling above upper tail. Underside.
As in nominate form. (PI. 15, fig. 129.)

FEMALE. Upperside. Somewhat similar to nominate viola but considerably larger, and,
as in that race, the discal band of the hind wing is restricted at the costa but expands over the
disc in mid area and lower end. (PI. 15, fig. 130.)

$ form vansomereni Poulton, 1925 : 553

Fore wing length usually 40-42 mm., shape falcate. Upperside. Base of wing brownish
shading to black toward the discal band, basal area often with a greenish sheen; distal border
blackish. Usually no spot in the cell, but there may be a slight indication of orange scales,
two spots beyond the cell, part of the discal series, usually free, as are the three subapical in
the postdiscal series; the remaining spots of both series conjoined in varying degrees; all spots
tawny orange to orange but those of la-ib paler to whitish. Margin of wing with tawny brown
lunules separated by dark veins. Hind wing basal area brownish toward costa but distally
merging into the bluish white discal patch which is narrowest at the costa but expanded over
the mid area then narrows to, then crosses, the inner fold, the borders of the patch tinged with
greenish blue scales; the outer border of the wing blackish and wider than in nominate race:
submarginal whitish to bluish marks strong; marginal border reddish above upper tail then
olive to anal angle which has two black dots. Tails moderately long with rounded ends, upper
8 mm., lower 6 mm., black with olive mid line. Margin of wing black with slight white fringe.
Underside. Rusty greyish brown ground colour with varying amount of satiny bars from hardly
visible to strong especially in the subapex; black lines strongest in the basal area of ib and on
either side of the discal band which varies in distinctness, tornal black marks variable, often
strong. Hind wing ground colour as fore wing, black marks fine; discal pale band variable but
narrower than patch above, often limited to the discal zone, sometimes diffused over and hardly
visible; the postdiscal olive ochreous and reddish lunate marks outlined proximally in black,
usually strong ; submarginal pale lunate marks strong to moderate ; border reddish above upper
tail then olive to anal angle, with black dots distinct. (PI. 15, fig. 131.)

Variation. Some indication of variation has been given in the foregoing general description.
Upperside. A marked variant has the ground colour of the fore wing rusty brown to brownish
in the discal line while the outer border is strongly rusty, especially in apical half, so that the
marginal lunules are absorbed into the ground colour except at the tornal angle. The discal
and postdiscal orange spots are conjoined by rays. The hind wing discal band is strongly
suffused over with orange scaling from costa to 5. The whole pattern is suffused over with a
golden rusty bloom. (PI. 15, fig. 132.)

130 V. G. L. VAN SOMEREN

Range: This subspecies occurs in S. Nigeria and Cameroon, then skirts north
of the Congo equatorial forest to E. Congo, Uganda, S. Sudan and N.W. Kenya.
(Map 6).

Charaxes viola kirki Butler
(PI. 16, figs. 133-137, Text-fig. 29 [aedeagus])

Charaxes kirki Butler, 1881 : 105.

Charaxes etheocles $ f. kirki Butler; Rothschild, 1900 : 486.

Charaxes etheocles $ f. kirki Butler; Aurivillius in Seitz, 1912 : 136.

Charaxes etheocles $ f. rogersi, Poulton, 1919 : Ixxxi.

Charaxes etheocles $ f. albifascia Poulton, 1926 : 553, 555.

Charaxes etheocles $ f. handari Poulton, 1926 : 553, 555.

Charaxes viola kirki Butler; van Someren & Jackson, 1952 : 265.

GROUP 2, AREA I. N. AND CENTRAL TANZANIA AND PARTS OF KENYA

Females of viola from most localities in this area are dimorphic on the upperside.
Though somewhat similar to female viola picta they can readily be distinguished by
the form of the hind wing discal band, which is widest at the costa, then tapers
strongly to the inner border, above the anal angle, but usually does not cross it.

MALE. Fore wing length 44 mm., averaging slightly larger than picta; shape falcate. Upper-
side. Ground colour fore wing blue-black with slight greenish sheen at base and along costa.
A large blue spot at upper angle cell end; a large spot and a small dot below beyond cell end; two
subapical spots are white or slightly ochreous; glaucous marginal lunules usually large and
distinct, increasing in size from apex to tornus. Hind wing velvety black with no green in
postdiscal line; submarginal bluish white spots small, often indistinct except in area opposite
tails and at anal angle; marginal border a mixture of reddish and green, then green at upper
tail, then olive to anal angle. Tails tapering abruptly, of about equal length 5-6 mm. Under-
side. Ground colour ashy grey-brown or brownish grey, rather strongly variegated, the satiny
bars and black marks strong. Fore wing with a satiny quadrate mark in cell, a satiny bar
through disc and a patch at upper apex; submarginal dark marks often strong especially toward
tornus. Hind wing with a sub-basal satiny bar, a rather broken bar through the disc; post-
discal zigzag line of olive and maroon lunules strong, outlined proximally in black; marginal
border maroon to upper tail then olive to anal angle, strongly marked. (PI. 16, figs. 136, 137.)

Variation. Upperside. Some specimens exhibit a reduction in the number of fore wing
spots, the number being reduced to two small subapical which may be white or ochreous; the
marginal glaucous lunules are obscured. In the hind wing the submarginal spots are only just
visible in the tail region; the marginal border in this area is broken up into separate spots;
the tails are mostly black. Underside. Black marks are reduced and the whole pattern suffused
over.

FEMALE. The named female forms with orange fore wing pattern are kirki, albifascia and
rogersi. The form with a white fore wing pattern has been named handari. These females can
be distinguished from picta by their darker, blacker ground colour and the form of the hind
wing discal band.

$ form kirki Butler, 1881 : 105

Fore wing length 34-38 mm. ; shape falcate, not so markedly as in nominate viola but more
like picta. Upperside. Ground colour darker, almost black, especially on the hind wing border.
Fore wing pattern: an orange spot usually present in the cell, varying in size and clarity; the
discal and postdiscal orange spots, except for the two subcostal discal ones and the two upper

REVISIONAL NOTES ON AFRICAN CHARAXES 131

subapical, fused together, with occasionally some black scales indicating the line of fusion in
areas 5-3. The colour of this band is stronger orange than in picta, the lower portion of this
band is almost parallel-sided from 3 to hind margin. Marginal lunules rusty to orange, only
slightly divided by dark veins. Hind wing ground colour black, slightly greyer on inner fold;
discal band widest at costa then tapering rapidly to inner fold above anal angle, but usually
not crossing it; the upper or costal portion of the band tinged with orange scaling to about
middle of disc ; borders very slightly tinged blue ; submarginal bluish white marks well separated
but distinct; marginal lunules reddish maroon above upper tail then olive to anal angle; edge
black, tails longish, upper with rounded end, lower pointed, length 7 and 5 mm. Underside.
Ground colour fore wing greyish brown, pattern strong; basal black marks well developed;
satiny bars distinct, a quadrate mark at end of cell and a triangular patch at upper part of
apex; discal and postdiscal light marks here separated, the former pale creamy ochreous, the
latter ochre-orange with a pinkish tinge, both outlined in black proximally; submarginal dark
marks increasing in size and blackness, strong at tornus and area above, the tornal spot with
lilac surround. Hind wing basal area darker chocolate-brown crossed by a sub-basal satiny
bar; black marks fine but distinct; discal band whitish, slightly shaded brownish distally and
outlined in black proximally; zigzag postdiscal olive and maroon line strong, extending from
costa to above anal angle; bordered by a wide greyish brown zone; submarginal row of lilac
whitish marks fairly clear, most distinct in region of tails; marginal border reddish above upper
tail then olive to anal angle and carrying black dots. (PI. 16, fig. 135.)

$ form albifascia Poulton, 1926 : 553, 555

Upperside. Very similar to kirki on fore wing, but discal band of hind wing entirely white.
(PL 16, fig. 133.)

$ form rogersi Poulton, 1919 : Ixxxi

Upperside. Generally similar to kirki, but both fore and hind wing bands orange or orange
tawny though slightly paler on costal portion of hind wing. The ground colour of fore wing
more rusty brown at base and on the distal half of the wing, so that the apical portion is
almost uniform rusty, thus obliterating the marginal lunules. Submarginal spots of hind wing
small, almost obsolete in upper half. Underside. Pattern strong, but slightly suffused with
an ochreous bloom. (PI. 16, fig. 134.)

Variation. Upperside. A boldly marked form in which the ground colour of both wings is
more blackish, thus the orange bands of both wings are in strong contrast.

$ form handari Poulton, 1926 : 553, 555

In this form the upperside patterns of fore and hind wings are creamy white or white.

Variation, (a) Upperside. Ground colour black with slight blue sheen over base of fore
wing; two large white spots beyond the end of cell, an equally large spots at base of 4, set well
in, spots in 3-1 a of gradually increasing size, rounded or straight on inner side, angled on outer,
are very slightly bluish on borders; postdiscal spots are, three subapical in a row, spot in 4
set in followed by free spots in 3-2 ; border of wing with slight indication of glaucous lunules.
Hind wing ground colour black, disc crossed by a white band widest at costa then tapering to
above upper angle, the borders strongly suffused with blue, especially on the lower half; submar-
ginal bluish white spots distinct; marginal border reddish above upper tail then olive-green to
anal angle. Underside. Fore wing ground colour greyish brown with some indication of
satiny bars; pattern of upper side strongly reproduced and accentuated by black on proximal
side of marks ; the border of wing slightly darker brown so that the upper marks in the submargin
are obscured, but tornal black mark and that in space above accentuated by a whitish surround.
Hind wing ground colour more brownish than fore but discal band whitish and strongly indicated

ENT. 23, 4 IJ

I 3 2 V. G. L. VAN SOMEREN

and going through to inner fold above anal angle; postdiscal whitish lilac linear marks strong;
border as usual. (PI. 20, figs. 188, 192.)

Variation, (b) Upper side. Ground colour of fore wing more brownish at base but with a
bluish sheen. A white spot in the cell, discal and postdiscal ones larger than in (a), all spots
creamy white.

Hind wing ground colour as fore but outer border blacker; discal band creamy white, widest
at costa then tapering, lower half tinged with greenish blue scaling. Underside. As in (a) but
ground colour paler, and pattern less strong.

Range: N. and Central Tanzania and along its coast to north of the Ruvuma
River; extending into Kenya along the coast and inland to Ukambani and Masai
country, spreading into the Kikuyu area, Nairobi, Thika, Fort Hall, Sagana and
Nyeri. (Map 6).

Charaxes viola suk Carpenter & Jackson

(PI. 16, figs. 138-144-)

Charaxes etheocles suk Carpenter & Jackson, 1950 : 98.

Charaxes etheocles suk Carpenter & Jackson; van Someren & Jackson 1952 : 266.

GROUP 2, AREA 2. N. AND CENTRAL KENYA

This aggregate comes nearest to viola kirki; it is very variable in size but is usually
smaller.

MALE. Fore wing length 37-40 mm., shape falcate, often strongly. Upperside. Ground
colour of fore wing black with some green sheen at base and along costa. Pattern : a small blue
spot at cell end, two blue spots beyond in discal line, two subapical spots white or slightly
ochreous. Glaucous marginal lunules strong; fringe white separated by black veins. Hind
wing ground colour black; submarginal white dots most evident in region of tails, double at
anal angle; marginal border reddish above upper tail then olive to anal angle. Tails acutely
tapering and of about equal length, 4-5 mm. Underside. Ground colour fore wing ashy
grey-brown with a satiny quadrate mark in the cell, a satiny bar through the disc and a patch
in upper half of the apex; basal black marks strong; subapical white marks show through from
above; submarginal dark marks complete, becoming black in 2 and at tornus, there is also a
black mark in ib in discal line. Hind wing ground colour slightly browner; a satiny sub-basal
bar and a more distinct bar through disc; postdiscal olive and maroon lunules distinct but
submarginal pale spots rather obscured; marginal border as above. (PI. 16, figs. 138, 139.)

In some specimens the underside ground colour is slightly olive-tinged and the pattern is
bolder, especially the satiny greyish bars in the fore wing.

$ form kirkoides Carpenter & Jackson, 1950 : 98

A large to medium form very similar to viola kirki with upperside basal area of fore wing
rather brownish; there may be an orange spot in the cell; the rest of the pattern as in kirki.
The hind wing band is whitish with some orange scaling mostly over the upper-mid area; it is
widest at the costa, tapering to the inner fold. (PI. 16, fig. 141.)

$ form achaemenesopsis Carpenter & Jackson, 1950 : 99

Variable in size, but usually 33-35 mm. in fore wing length; pattern of fore and hind wing
very similar to form handari of viola kirki; the discal and postdiscal series of fore wing spots
white, the band on the hind wing also white with a varying amount of bluish suffusion on the
borders.

133
liscal

SENEGAL

ir^

Vd

IRA LEONE

LIB

CHARAXES

C. viola viola
Dne form of

C. viola picta
One form of

C. viola dona
Three forms

C. viola chan,
One form of

C. viola suk (
Three forms

C. viola kirki
Four forms

C. viola phae

C. viola nr. p
Only one o

C. v/'o/o d/'ver
Numerous 5

C. V/O/O /03H(

Numerous 5

C. v/o/o bro/r
One form o:

C. v/o/o varic
f(f strongly

C. v/'o/fl figini

, but

rymg

ioja,
g to
r isoi.
issia

as a
the

;ts is
viola
ence

;y of
gical

ties,

>and

wing

two

with

tiind

ginal

with

line

inge.

132

and going
border as

Variatic
bluish she
creamy wl

Hind w
at costa ti
ground co

Range
River; e
country,
Nyeri.

Charaxes
Charaxes

This a
smaller.

MALE.
colour of
spot at o
ochreous.
wing groi
anal angL
tapering
grey-brov
in upper ]
above; su
black mai
bar and ;
submargi:

In som
bolder, es

A large
rather br
The hind
widest at

Variab
very simi
white, th
borders.

REVISIONAL NOTES ON AFRICAN CHARAXES 133

An extreme variant has the discal spots in fore wing reduced in size, the lower postdiscal
spots suffused over with greyish scaling; the hind wing band is narrowed considerably.

$ form intermedia Carpenter & Jackson, 1950 : 98

Upperside. In this form the upper spots of the discal and postdiscal series are orange, but
those of 2-ia are whitish; the hind band white. (PI. 16, fig. 140.)

$ form albifascia Poulton, 1926 : 553

The fore wing pattern and colour as in kirkoides; the hind wing band is white with varying
amount of bluish on borders. (PI. 16, fig. 143.)

Range: This aggregate is plentiful in the thorn-bush country of Karamoja,
Turkana, Suk and Kara-Suk south to Kamasia and the Baringo area, extending to
the Kampi-ya-Moto district north-west of Menengai, Lower Molo River and Visoi.
It was taken in considerable numbers on infected, fermenting seed pods of Cassia
at the foot of Kacheliba Rock, Suk. (Map 6).

The late T. H. E. Jackson was of the opinion that viola suk should be sunk as a
synonym of viola kirki. This however, would ignore certain facts such as: (a) the
males of suk and kirki differ; (b) the white-barred female form achaemenesopsis is
extremely plentiful, making up half the female population, whereas handari of viola
kirki is the exception in this population ; (c) there must therefore be some difference
in the genetical make-up of the two races.

It is possible that the two aggregates merge somewhere in the Kedong valley of
the Rift, but this does not discount the fact that in suk we have a bio-ecological
aggregate occupying a large area to the north of the range of viola kirki.

Since taxonomic Rules do not provide for any category lower than a subspecies,
not even localized genetical forms, I prefer to treat viola suk as a subspecies.

Charaxes viola chanleri Holland
(PL 20, figs. 189-191)

Charaxes chanleri Holland, 1895 : 753.

Charaxes etheocles chanleri Holland; Rothschild & Jordan, 1900 : 483.

Charaxes viola chanleri Holland; van Someren & Jackson, 1952 : 268.

GROUP 2, AREA 3. Mr. KENYA AND NJOMBE RANGE TO S. ETHIOPIA
This subspecies belongs to the eastern group of viola, having the hind wing band
widest at the costa and tapering.

MALE. Fore wing length 34 mm., shape falcate. Upperside. Ground colour of fore wing
black with greenish sheen at base of cell. Spots: one large blue spot at upper end of cell; two
beyond, upper large, lower small, subapical spots two or three white or slightly tinged with
ochre; glaucous marginal lunules strong and only narrowly separated by dark veins. Hind
wing black with slight trace of greenish wavy line in postdiscal zone opposite tails ; submarginal
spots suppressed but just visible in lower section; marginal border mostly greenish, with
little red, the lunules somewhat triangular, the apices projecting toward the submarginal line
and touching it. Underside. Ground colour light greyish brown with slight olive tinge.

134 V. G. L. VAN SOMEREN

Fore wing basal black marks strong; discal satiny bars well marked; postdiscal light marks
visible, both outlined proximally in black; submarginal dark marks obscured in upper half
but strong toward tornus where marks are black. Hind wing slightly browner in colour, the
basal black lines thin but discal satiny pale line strong as also the olive-maroon zigzag line;
border narrowly red above upper tail then olive to anal angle.

The underside may sometimes be more uniform, the black marks thin but the tornal black
marks strong. In the hind wing the discal satiny bar is obscured in the general pale ground.
(PI. 20, fig. 191.)

FEMALE. Fore wing length 37-40 mm. ; shape falcate. Upperside. Ground colour fore
wing brownish black at base shading to black at discal band; outer border more brownish,
shading to rusty in curve of wing where there is only a slight indication of ends of dark veins.
In the discal line there is a small tawny spot at end of cell, then spots in sub-base of 65 followed
by a spot in 4 set well in, spots in 3 to hind margin increasing in size, ovoid to quadrate in
shape ; all spots orange-tawny. The postdiscal spots are, three subapical in a row, spot in 4 set
well in, spots in 32 conjoined with discal marks with some dark scaling indicating line of fusion.
Sometimes the orange spots in 4-6 are joined by rays. Hind wing ground colour as fore wing,
but darker on the border which is wide : the discal band which is widest at the costa, tapers to
the inner fold above the anal angle and is white with some blue, especially on the borders of the
lower half ; submarginal line of whitish lilac linear marks distinct and complete ; border reddish
above upper tail then olive to anal angle; margin black with white internervular fringe; tails
long, upper 7 mm., lower slightly shorter. Underside. Ground colour clay-brown; black basal
lines thin; discal and postdiscal bands of above well represented but as separate marks; dark
submarginal marks in curve of wing rather indistinct except in lower half where the spots are
black with a pale surround, mostly on the distal side. Hind wing slightly darker than fore;
satiny sub-basal bar fore wing rather broken; the discal band of above well represented but
narrower; postdiscal olive and maroon lunules not very distinct; submarginal row of linear
marks, lilac and whitish, distinct and complete; submarginal border reddish above upper tail,
then olive to anal angle. (PI. 20, figs. 189, 190, type.)

Variation. Upperside. The conjoined discal and postdiscal orange band of fore wing more
parallel-sided from 2-ia; the hind wing band slightly orange-scaled in outer mid area; the sub-
marginal spots obscured in upper half but distinct in region of tails. Underside. With fore wing
basal marks heavy. The discal band of hind wing silvery and extending through the inner fold.
Postdiscal lunules strong, especially in lower portion.

Range: extending from the lower Meru (Mt. Kenya) area and the Njombe Range
to southern Ethiopia, Mega and Neghelli to Dua River. (Map 6).

Charaxes viola daria Rothschild
(PL 20, figs. 185-187)

Charaxes etheocles daria Rothschild, 1903 : 326.

Charaxes etheocles chanleri Holland; Rothschild, 1900 : 483. Males, Gillett Mts., Abyssinia

(identified as chanleri Holland from Walenso, error!)
Charaxes etheocles daria Rothschild; Carpenter, 1935 : 3^o.
Charaxes viola daria Rothschild; van Someren & Jackson, 1952 : 264.

GROUP 2, AREA 4. EAST SIDE OF ETHIOPIAN RIFT

The only authentic female of this subspecies is black with white or creamy bands'
as described hereafter.

MALE. Fore wing length 41-42 mm., shape falcate. Upperside. Ground colour black with
very slight green sheen. Subcostal blue spots variable but usually with a blue spot at the end
of the cell, one or two blue spots beyond, lower small, two subapical whitish or bluish and small;

REVISIONAL NOTES ON AFRICAN CHARAXES 135

marginal glaucous lunules well separated, often small or only just indicated. (In two specimens
the subcostal spots are missing except for the upper subapical.) Hind wing black, no postdiscal
greenish line as a rule but may be faintly indicated ; submarginal whitish spots usually obscured
in upper part but more distinct in region of tails ; marginal border may be reddish or a mixture
of reddish and greenish then olive to anal angle ; margin black ; tails short of about equal length
5-4 mm., black with olive mid line. Underside. Fore wing ground colour greyish brown with
satiny bar in the cell, a bar through the disc and a patch at upper part of apex; black lines
distinct but not heavy; postdiscal black line thin, then heavy at ib; submarginal dark spots
present but only black at tornus. Hind wing slightly more brownish, but satiny bars not strong ;
postdiscal line of olive and maroon lunules outlined in black, proximally, well marked ; submar-
ginal linear marks faint; marginal border narrow, reddish above upper tail then olive to anal
angle. In some specimens the underside ground colour may be more uniform, the whole surface
shiny, except in the curve of the fore wing which is browner.

Descriptions based on topotypical specimens from Gugura and Gillett Mts. (PI. 20, fig. 187.)

FEMALE. Fore wing length 37 mm., shape slightly falcate. Upperside. Ground colour
black with slight bluish sheen at base and along costa. Pattern : a small white spot in the cell
(absent in the type). In the discal row two elongate spots between end of cell and postdiscal
series, spot in 4 set well in, spots in 3 to hind margin increasing in size, first two rounded on inner
end, that in ib angled. Postdiscal spots, three insubapex in a row, spot in 4 set in, those in 3-2
free, or just touching discal spots. Hind wing ground colour brownish black at base fading
to greyish on inner fold but border of wing black; discal band widest at costa tapers toward
inner fold above anal angle; but there represented by a pale mark, the whole band white,
slightly bluish on borders of lower half. Submarginal line of whitish marks complete and distinct ;
submarginal border reddish above upper tail then olive to anal angle; margin black. Tails,
7 mm. and 5 mm. Underside. Fore wing greyish brown, slightly browner on curve of the wing.
Black marks at base fairly strong on inner border of discal band ; discal band whitish ; postdiscal
spots well represented and outlined in black proximally with a black spot in ib; submarginal dark
spots distinct, increasing in size and blackness to double mark in tornus. Hind wing ground
colour slightly browner, especially on distal border; basal black marks fine; discal band as
above but slightly narrower, whitish and extending to inner fold; postdiscal line of lunules
olive and maroon strongly indicated; submarginal line complete; marginal border as usual with
black dots in area of tails and double at anal angle. (PI. 20, figs. 185, 186, type.)

Variation, (a) Upperside. Very similar to foregoing but upper discal and postdiscal white
marks larger, though lower discal marks are less wide but contiguous with postdiscal marks
in la-ib. Underside. There is a corresponding difference.

Variation, (b) Upperside. Ground colour of both wings above browner; the discal and post-
discal white marks conjoined from 5 to ib, but the lower part of the band hardly wider or less
wide than in nominate form. Underside. Pattern is creamy, the black marks strong.

Variation, (c) A specimen which I place as this race differs in having the upperside ground
colour of both wings almost black, the upper discal and postdiscal spots reduced in size, the
lower marks narrower, all marks creamy. The hind wing band slightly narrowed and cream
in colour; the submarginal linear marks obscured.

Through the kindness of Dr. Viette of the Paris Museum, I have now been able to
examine the material collected by Ungemach.

There are six males which agree very well on the upperside with an equal series
of topotypical males of daria from Walenso and Gillett Mts. On the underside
they are also very similar, the only difference being that three of the Youbdo
males are rather uniform on the underside, lacking any strong pattern, but this
variation is also to be noted in other races of Char axes viola such as picta.

The two females are extremely interesting. That identified as daria Rothschild
differs from nominate daria by the upperside having a much narrower fore wing bar

136 V. G. L. VAN SOMEREN

in ia-ib, the mark in 2 more elongate-quadrate, the one above in 3 same shape but
smaller, marks in 4-5 conjoined, those in 6 free, the subcostal-subapical mark also
free ; all marks from 2 to subcostal slightly creamy tinged. That stated to be near
form vansomereni Poulton of viola picta has a very similar pattern on the upperside
but the bars are broader in both wings, that of the hind wing especially so, being
white with bluish borders; the fore wing spots conjoined to 6, orange-ochre in colour,
thus very similar to form vansomereni of viola picta.

It would thus appear that the Youbdo aggregate may possibly intergrade with the
viola-picta aggregate on the eastern Sudan border.

Ungemach records a female form like $ f. vansomereni Poulton as belonging to
daria, but since his specimen came from Youbdo, west of the Abyssinia Rift in
Walegga country, there is some doubt about its correct position.

Range: The type and topotypes are from the east side of the Abyssinian Rift
from Jabalo, Gugura and Gillett Mts. at Walenso. This race seems to extend to the
upper Schebeli on the east and to Adola on the west. (Map 6).

Records from Walegga (Ungemach & Carpenter) probably represent a mixed
aggregate.

Charaxes viola phaeus Hewitson

(PI. 21, figS. 193-200)

Charaxes phaeus Hewitson, 1877 : 82.

Charaxes phaeus Hewitson ; Trimen & Bowker, 1887 : 344.

Charaxes phaeus Hewitson ; Butler, 1895 : 361.

Charaxes etheocles phaeus Hewitson; Rothschild & Jordan, 1900 : 488.

Charaxes etheocles $ f. coryndoni Rothschild, 1900 : 488.

Charaxes viola phaeus Hewitson; van Someren & Jackson, 1952 : 269.

Charaxes viola phaeus $ f. vansoni van Someren & Jackson, 1957 : 43-

GROUP 3, AREA I, SEE RANGE P. 148

In this group we find the first evidence of complete divergence amongst the female
forms from a " viola-like " pattern to that of phaeus. But amongst the southern
or typical phaeus we find a white-barred female form which resembles somewhat the
female forms daria, handari and achaemenesopsis of northern races of viola. This
form we named vansoni. In this subspecies also, one is faced with the fact that there
are two environmental aggregates: (a) that of the southern areas (Beira-Transvaal-
Rhodesia and part of Bechuanaland) in which phaeus and vansoni occur side by
side, and (b) the aggregate of the northern range of phaeus (Malawi, Tanzania to
the west side of Lake Victoria), in which only the form phaeus has been recorded.

One must assume that there is a genetical foundation to the dual female form in
the south, and that the genes for vansoni are absent in the northern aggregate.
We thus have what I have termed (for want of a better term) bio-ecological strains,
in which ecological environmental conditions have played an important part.

Subarea (a). SOUTHERN AGGREGATE OR NOMINATE

MALE. Fore wing length 27-30 mm., shape falcate. Upperside. Ground colour blue-black
with greenish sheen at base of costa; blue spots limited to one at upper angle of cell, two spots
beyond cell, upper large, lower small, two spots in subapex bluish or whitish; glaucous marginal

REVISIONAL NOTES ON AFRICAN CHARAXES 137

lunules small, well separated. Hind wing ground colour black; some specimens with a varying
amount of greenish lunules in the postdiscal line opposite tails ; submarginal spots blue or bluish
white, complete ; marginal border narrow, greenish red above upper tail then olive to anal angle ;
tails short, 3-4 mm. Underside. Fore wing ground colour earthy grey-brown or even pinkish
grey, very uniform and plain-looking, relieved slightly by usual satiny bars; black lines thin;
submarginal dark marks in curve of wing present but slight, strongest at tornus. Hind wing
ground colour as fore wing, satiny bars weak; postdiscal lunate marks faint narrowly reddish
and olive; margin as usual.

Description based on topotypical specimens.

$ form phaeus Hewitson, 1877 : 82

Fore wing length 33-34 mm., shape slightly falcate. Upperside. Basal triangle of fore wing
pale blue with slight greenish sheen, or slightly violet-tinged in old specimens, the blue not
reaching the end of cell but in its lower angle extending to the postdiscal line on the hind border ;
distal portion of wing black, paler toward the curve on outer border; subapical spots in post-
discal line large and whitish, spot below smaller, other spots faintly indicated and bluish reaching
ib. Hind wing mostly pale blue from base to postdiscal line, often with a dark spot at costa;
border black, widest at upper part in 6-7 then tapering to anal angle; submarginal spots blue
with white central dot; submarginal border reddish above upper tail then olive to anal angle;
margin black, tails 7-5 mm. long. Underside. Fore wing ground colour pale greyish brown
with a pinkish glaze overall, so that the satiny bars are obscured and the pattern subdued, the
black marks, except the tornal black ones, hardly visible. Hind wing as fore, all marks subdued.

$ form coryndoni Rothschild, 1900 : 488

Variation. Upperside. Similar to phaeus but postdiscal spots of fore wing distinct to ib;
the blue of the hind wing often complete in costal area without any interruption.

$ form vansoni van Someren & Jackson, 1957 : 43

Very similar to the white-barred forms in viola kirki or viola suk. Fore wing length 34 mm.,
shape falcate. Upperside. Ground colour of fore wing brownish at base but darker toward
discal line and on outer border. Discal and postdiscal series of spots well apart except that the
spots in ib may touch. Hind wing ground colour brownish black at base, blacker on border;
but paler on inner fold. Discal white band extending from costa, where it is widest, then tapering
toward inner fold and there represented by a pale mark, the lower half of the band shaded with
blue. Submarginal linear marks clear, lilac or bluish with white centres; submarginal border
narrow above upper tail and dull reddish, then olive to anal angle ; margin black ; tails longish,
7-5 mm. Underside. Basal area of fore wing pale greyish shading to brownish toward the
outer border, basal black marks thin but distinct; discal band well represented, creamy to ochre;
postdiscal spots suffused over; submarginal brownish marks obscured though tornal black
marks distinct. Hind wing marks obscured, the discal pale band slightly indicated at costal
end. The whole underside with a satiny glaze. (PI. 21, figs. 199, 200.)

Subarea (b). NORTHERN AGGREGATE

Attention was drawn to this aggregate by van Someren & Jackson, 1957 : 44.
We expressed the view that a satisfactory placing could not be made until material
had actually been bred; unfortunately this has not been possible, but considerable
additional material has been captured, especially in the Kigoma area north-east of
Lake Tanganyika. These specimens bear out the observation that they are larger
than nominate phaeus of southern regions and are rather more brownish below.

V. G. L. VAN SOMEREN

ft

-. Tanganyika

J KATANGA . ^

^t^. .."": \ Ft - Rosebery* W

t Mwinilunga **( ' ', *.

; \Solwezi ... '....*"

T ^ -, --.,

ger Hill

Ndola \

Nkushi

MALAWI

, f . ZAMBIA

lalovale j i / j r.

Kabompo C J / Katet * ei ": v^U*'

T L S 'P on S / \ . A

>oma ^ J^Kafuehoch * ^Luangwa Bdg. '"" \* V

Nankoya T Chisamba ^ :

* Mlange ...

KEY

Charaxes viola phaeus. .^^

+ .^J L. Kariba .

C. viola diversiforma. jp** Salisbury y ,.

or ,-^ -- '' * '

T C. viola variata. !^r f ' '?"'!'.' X/ "*,-

\ .** ". & '' : "' ; "' -i^Vumba

BECHUANALAND\ / SOUTHERN RHODESIA

'.... * Bulawayo

MOZAMBIQUE'
\

MAP 7.

NOTES ON AFRICAN

3. $ form caerulescens van Someren & Jackson, 1957 : 45

(PL 18, figs. 165, 166)

This is a development from the previous two forms toward form phaeus of viola phaeus.
Upperside. The basal areas of both wings are an iridescent greenish blue sharply denned distally.
The fore wing discal spots missing and the postdiscal series represented by two large subapical
spots and obscured spots in 4-2. Underside. Red-brown with strong lilac bloom overall,
thus pattern obscured. This is rather a small form, fore wing length 33 mm.

4. 9 form albocaerulea van Someren & Jackson, 1957 : 45

(PL 18, figs. 159, 160)

A development from form 2. Upperside. The bases of both wings pale blue, that of fore
wing whitish in distal area in za-ib. and part of the enlarged discal blue spots; the postdiscal
spots large and white in subapical area, but blue from 5-1 b. Hind wing blue not reaching the
costa but here represented by a separate blue mark. Underside. Less reddish brown, more
greyish than previous form, with 'a stronger pattern visible especially on the distal borders of
both wings.

5. $ form albimacula van Someren & Jackson, 1957 : 45
(PL 18, figs. 161, 170)

Upperside. Represents a development from form 4 in the direction of a white-barred form,
but retaining the blue scaling at the bases of both wings. The fore wing discal spots are white,
two large spots beyond end of cell, an obscure spot at base of 4, spots from 3 to ib increasing in
size. Postdiscal spots are: two subapical, white, rest from 4 to ib smaller and that in ib bluish.
Hind wing discal band widest at costa is whitish, strongly suffused with blue on lower half,
Underside. Rusty brown but discal band of fore and hind wing visible.

6. $ form ochremaculata van Someren & Jackson, 1957 : 45
(PL 18, figs. 167, 168)

Upperside. Bases of both wings brownish with strong blue to greenish scaling. Fore wing
with a large ochre spot in the cell, discal and postdiscal spots ochreous, well separated up to 2
then fused in za, ib forming a large mark tinged with ochre distally and greenish proximally.
Hind wing band widest at costa and extending toward inner fold, white in middle but borders
suffused with greenish blue. Underside. Greyish brown, rusty on borders but with satiny sheen
overall but not obscuring the bars of above, these are strong and ochreous in colour.

7. $ form diversiforma van Someren & Jackson, 1957 : 44
(PL 18, figs. 162, 169)

In the type, the spots on the upperside of fore wing are separated to area 2 and then conjoined.
In the specimen here figured, the discal and postdiscal tawny orange spots are conjoined from
4 to hind margin. The hind wing band, widest at costa, is creamy, tinged greenish blue on
borders.

I 4 o V. G. L. VAN SOMEREN

strong; discal white bar strong, distally bordered with black especially in subcostal area; post-
discal zigzag reddish and olive lunules present but strongest in lower half toward anal angle;
submarginal lunules well marked and with black distally to double dots in anal angle; border
reddish above upper tail then olive to anal angle.

Holotype female. ZAMBIA : 60 miles on Mwinilunga to Kabompo Rd., 1-5 . vi . 1963
(C.B.Cottrell).

Variation. Fore wing length 34 mm. Upperside. Ground colour and general pattern
similar to nominate form but all white marks larger and the basal areas of both wings more
shaded with pale blue, increasing in intensity at discal row of spots fore wing and on borders
of hind wing discal band. Underside. Very similar to nominate form. The interesting feature
in this variety is the blue suffusion over the bases of the wings.

$ form tricolor forma n.

Upperside. Ground colour rather blacker than in nominate form with a strong greeny
blue sheen at base, more strongly blue on proximal border of fore wing discal spots in la-ib.

General pattern of fore wing as in nominate form but all spots orange-ochre to orange, slightly
paler to whitish on inner half of marks in ib. Hind wing ground colour black with strong
greenish blue on proximal border of discal band, which is white in the upper half. Underside.
Rather darker in ground colour than nominate form, with all marks other than black lines,
white and in strong contrast. (PI. 19, figs. 183, 184.)

Holotype female. Same data as holotype female variata.

$ form rosella forma n.

Upperside. Ground colour similar to nominate form, fore wing with a curved white bar
extending from the costa at and beyond the cell end to hind margin; spot in 2 a long triangle;
spots beyond cell free; postdiscal spots limited to three subapical and one each in 4-3. Under-
side. Fore wing presenting a strong replica of the Upperside pattern with submarginal pale
and dark marks strong. Hind wing discal band strong in upper half but narrower and irregular
in 3-2, widening again toward inner fold. Postdiscal zigzag line of lunules strong, ochreous
at costal end but lilac and maroon to anal angle and outlined in black; submarginal pale line
well marked with large bluish spots, black-centred in region of tails. (PI. 19, figs. 175, 176.)

Holotype female. Same data as previous forms.

$ form cottrelli forma n.

This interesting form exhibits a transition toward the form phaeus of viola phaeus.

Upperside. Basal areas of fore wing including the discal line in la-ib, greenish blue; the
disc of the hind wing also blue, the rest of the wing black. Fore wing with discal spots, two
beyond cell, spot in 4 set well in, all white; spots in 3-2 larger and slightly blue, spot in ib
merged into blue of base ; postdiscal spots, three in subapex white, spot in 4 set in, followed by
ochreous spots in 3-1 b. Hind wing basal area and discal patch greenish blue represented at
costa by a blue quadrate mark; extreme base slightly dark-scaled; submarginal linear marks
bluish white, black distally, very strong; marginal border reddish above upper tail, then olive
to anal angle. (PI. 19, figs. 177, 178, type.)

Variation. Upperside. Very similar to form rosella but basal blue areas inclining to whitish
in lower discal line i-ia-ib. Upper discal spots in cell and in 3 ochreous, that in 2 blue. Post-
discal spots three in subapex, upper two white, lower ochreous, followed by less distinct spots

REVISIONAL NOTES ON AFRICAN CHARAXES 141

in 4-3 also ochreous but spots in 2 and ib bluish, the lower one contiguous with blue in discal
line. Hind wing basal areas and disc blue, but not reaching the costa, here represented by a
free blue mark. Submarginal line of linear marks, bluish white and strong, margined distally
in black; marginal border as usual. Underside. Ground colour earthy brown with rusty
purplish tinge; pattern not very strong but tornal black marks heavy.

Holotype female. ZAMBIA: Mankoya, io.xii.ig55 (C. B. Cottrell}.

Range: This race is known from the N.W. regions of Zambia from Mwinilunga
area and at Kabompo and Mankoya thus mostly between the upper Zambesi and
its tributary, the Kabompo River. (Maps 6, 7).

GROUP 4, AREA 2. S. KATANGA, CONGO
Charaxes viola diversiforma van Someren & Jackson

(PL 18, figs. 157-172)
Charaxes viola diversiforma van Someren & Jackson, 1957 : 44-46.

MALE. Fore wing length 36 mm., shape falcate, but less so than variata or picta, and larger
than either; in size and general appearance more like kirki. Upper side. Ground colour fore
wing blue-black, with slight greeny sheen at base of costa and cell. Pattern : a trace of a blue
spot in the cell, two blue spots beyond, the upper large; two whitish or ochre-tinged spots in
subapex, sometimes a third small dot; traces of blue spots in 4-3. Glaucous marginal lunules
ill-defined, separated by dark veins. Hind wing blue-black, duller on inner fold; a greenish
wavy line in postdiscal zone most distinct opposite tails; submarginal row of bluish white
spots complete; submarginal border slightly reddish green above upper tail then olive to anal
angle; margin black, tails rather short, of about equal length 5-4 mm. Underside. Ground
colour earthy brown; pattern rather obscured, the satiny bars indistinct and the black marks
thin on both wings. (PI. 18, fig. 172.)

Females very variable, the following forms have been named:

i. $ form purpurea van Someren & Jackson, 1957 : 45

Fore wing length 34 mm., shape falcate. Upper side. Ground colour purplish black with
strong bluish green sheen at bases of wings. This form bears a superficial resemblance to
Charaxes virilis Rothschild but differs in having both a discal and postdiscal series of bluish
spots, the two upper subapical whitish. Marginal glaucous lunules slightly developed. Hind
wing basal area suffused with purplish green, distal border black carrying an irregular post-
discal greenish band; submarginal row of whitish lilac linear marks complete and clear; marginal
border as usual. Tails thin, 6-4 mm. long. Underside. Strongly rusty brown, more rusty
on outer border of fore wing; slight indication of satiny bars in sub-base and through disc.
Hind wing ground colour as fore wing, all black marks very fine ; postdiscal line of lunules present
but suffused over with a golden rusty bloom; submarginal spots lilac but obscured; marginal
border very narrow and indistinct. (PI. 18, figs. 157, 158.)

2. $ form viridicaerulea van Someren & Jackson, 1957 : 45

Upper side. Differs from form i in lacking the discal blue spots other than the one beyond
end of cell ; the postdiscal series rather stronger and extending to the hind border in line with
the more pronounced postdiscal bar in the hind wing. The basal areas of both wings iridescent
greenish blue. Underside. Red-brown with rusty golden bloom obliterating all pattern except
the tornal black spot of fore wing. (PL 18, figs. 163, 164.)

ENTOM. 23, 4. I2

V. C. L.

anganyika

TANZANIA

*&

,^, .ZAMBIA

Balovale

Kabompo
/ Namboma

*.

r>.

Ndola \

'-N'kushi f * {MALAWI 1

* / Katefei^V;;.

Ipongo

| e / Nkushi r < MALAWI ^Ci

.Kabompo I / Kate?ei !?": V/*U\

f I S Ipongo / ^ \ t> V \

> oma . ^iJfuehoch ^ /LuangwaBdg. ''" \* Y i

Nankoya T Chisamba & : /\

T U^^^^J^SL^^x / / H

KEY

r Charaxes viola phaeus.
f C. viola diversiforma.
F C. viola variata.

Kariba

V-

MOZAMBIQUE 1
Salisbury ,. Umtali \

'"'"/'SI OVumba

BECHUANALAND\ / SOUTHERN RHODESIA

Bulawayo

MAP 7.

L tfOTES Ott AFRICAN" CHARAXES * 4 $

3. $ form caerulescens van Someren & Jackson, 1957 : 45

(PI. 18, figs. 165, 166)

This is a development from the previous two forms toward form phaeus of viola phaeus.
Upper side. The basal areas of both wings are an iridescent greenish blue sharply denned distally.
The fore wing discal spots missing and the postdiscal series represented by two large subapical
spots and obscured spots in 4-2. Underside. Red-brown with strong lilac bloom overall,
thus pattern obscured. This is rather a small form, fore wing length 33 mm.

4. $ form albocaerulea van Someren & Jackson, 1957 : 45

(PI. 18, figs. 159, 160)

A development from form 2. Upperside. The bases of both wings pale blue, that of fore
wing whitish in distal area in la-ib. and part of the enlarged discal blue spots; the postdiscal
spots large and white in subapical area, but blue from 5-1 b. Hind wing blue not reaching the
costa but here represented by a separate blue mark. Underside. Less reddish brown, more
greyish than previous form, with 'a stronger pattern visible especially on the distal borders of
both wings.

5. $ form albimacula van Someren & Jackson, 1957 : 45
(PL 18, figs. 161, 170)

Upperside. Represents a development from form 4 in the direction of a white-barred form,
but retaining the blue scaling at the bases of both wings. The fore wing discal spots are white,
two large spots beyond end of cell, an obscure spot at base of 4, spots from 3 to ib increasing in
size. Postdiscal spots are: two subapical, white, rest from 4 to ib smaller and that in ib bluish.
Hind wing discal band widest at costa is whitish, strongly suffused with blue on lower half,
Underside. Rusty brown but discal band of fore and hind wing visible.

6. $ form ochremaculata van Someren & Jackson, 1957 : 45
(PI. 18, figs. 167, 168)

Upperside. Bases of both wings brownish with strong blue to greenish scaling. Fore wing
with a large ochre spot in the cell, discal and postdiscal spots ochreous, well separated up to 2
then fused in la, ib forming a large mark tinged with ochre distally and greenish proximally.
Hind wing band widest at costa and extending toward inner fold, white in middle but borders
suffused with greenish blue. Underside. Greyish brown, rusty on borders but with satiny sheen
overall but not obscuring the bars of above, these are strong and ochreous in colour.

7. $ form diversiforma van Someren & Jackson, 1957 : 44
(PL 18, figs. 162, 169)

In the type, the spots on the upperside of fore wing are separated to area 2 and then conjoined.
In the specimen here figured, the discal and postdiscal tawny orange spots are conjoined from
4 to hind margin. The hind wing band, widest at costa, is creamy, tinged greenish blue on
borders.

144 v - G - L - VAN SOMEREN

8. $ form cupreopurpurea van Someren & Jackson, 1957 : 45

(PL 18, fig. 171)

Upper side. Ground colour brownish black with a strong purplish sheen overall, so that the
discal and postdiscal spots which are entirely separate are coppery coloured. Hind wing
ground colour brownish black with strong purplish gloss, the discal band greenish blue with
dyslegnic borders.

The foregoing descriptions are based on paratypes (kindly loaned by the Tervuren
Museum) and are complementary and supplementary to the original descriptions
of the types.

Range: All material of this subspecies has come from the Upembe area of S.
Katanga of the Congo ; Upembe, Lulua, and Kafskumba. (Maps 6, 7) .

Charaxes viola loandae ssp. n.

(PL 17, figs. 145-152)
GROUP 4, AREA 3. N.W. ANGOLA

This subspecies presents features which suggest a parallel evolution to that of
diversiforma, and might even be considered identical to that race but for the fact
that the males differ as do the females.

MALE. Fore wing length 28-31 mm.; shape falcate. Upperside. Ground colour blue-black
with slight greenish sheen at base of fore wing. Blue spots limited to one blue spot in the cell,
one large spot and a dot below beyond cell end, two subapical spots bluish white and blue;
glaucous marginal lunules with central white mark complete from subapex to tornus. Hind
wing ground colour black with a trace of a wavy greenish line in postdiscal zone; submarginal
whitish spots large and distinct in type, (or may be small) ; marginal border largely reddish to
anal angle, or ochreous with some reddish to anal angle; edge black, with white fringe between
veins distinct; tails short, 5-4 mm. long. Underside. Very similar to that of male phaeus, pale
greyish brown with satiny bars very faint, black marks thin, but tornal black mark strong. Hind
wing ground colour as fore wing, the postdiscal line visible but not strong ; pale submarginal line
indistinct; marginal border narrowly red and olive to anal angle. (PI. 17, fig. 150.)

Female forms very variable, the following are named:

i. $ form primitiva forma n.

(PL 17, fig. 152)

Upperside. Male-like, ground colour of fore wing blue-black with very strong greenish
tone over basal area, blacker in disc, but greyer on border in curve of the wing. There is a
blue spot in the cell, an indication of a spot in subcosta beyond the cell; the postdiscal series
of blue spots from subapex to 2 rather indistinct ; marginal lunules glaucous but diffuse. Hind
wing black, strongly suffused over with bluish green, bordered in discal line with stronger bluish
green bar complete from costa to above anal angle; submarginal white spots distinct; marginal
border mostly reddish and mixed with olive toward anal angle. Tails thin, upper 6 mm.,
lower 5 mm. Underside. Ground colour pale rusty brown with a strong purplish bloom overall ;

REVISIONAL NOTES ON AFRICAN CHARAXES 145

satiny bars obscured; black lines faint but those in ib fore wing stronger though tornal mark is
weak.

2. $ form basiviridis forma n.

(PL 17, fig. 151)

Somewhat like form i but differing as follows: Upper side. Ground colour black but bases
of both wings more strongly green and limited to basal triangles. Fore wing with two white
subapical spots, rest in postdiscal line blue and hardly visible. Marginal lunules ill-defined.
Hind wing postdiscal wavy greenish line stronger; submarginal row of spots greenish; marginal
border a mixture of reddish and green scales. Underside. As in form i.

3. $ form violitincta forma n.

(PL 17, fig. 145)

Upperside. Ground colour black, base of fore wing violet-blue to discal line at la-ib, but
fading out at costa at mid cell, discal bar represented by paler blue diffuse spots increasing in
size from 3 to hind margin; margin of wing slightly brownish but lunules not definite. Hind
wing slightly brownish at the base, but disc of wing with a large violet patch not reaching costa
but here represented by a free spot; submarginal linear blue marks distinct; marginal border
a mixture of bluish and ochre then olive to anal angle. Underside. Very similar to previous
forms but black marks stronger and the tornal spot blacker.

4. $ form vansonoides forma n.

(PL 17, fig. 149)

Upperside. A transitional form toward form vansoni of viola phaeus; and somewhat like
form albimacula of viola diversiforma but without any blue at bases of wings, these areas being
black with a strong green sheen. The discal and postdiscal fore wing spots large; the margin
of fore wing with indistinct brownish lunules. Hind wing base black with slight green sheen;
discal band white, widest at costa, strongly suffused with greenish blue on outer border; sub-
marginal linear marks bluish white, well developed; marginal border reddish above upper tail,
ochreous olive to anal angle. Underside. Ground colour as in previous forms but white
areas of Upperside pattern apparent though narrower and ochreous in colour.

5. Nominate $ form loandae forma n.

(PL 17, fig. 146)

Upperside. Basal areas of fore wing brownish, with purplish sheen; pattern as in form 4
but all spots tawny orange, slightly bluish in la. Hind wing basal area brownish with strong
blue sheen in side light; discal band white, extending from costa to above anal angle, strongly
blue-scaled on the borders especially in lower half; submarginal linear marks bluish white and
strong; marginal border reddish above upper tail, then mixed olive and reddish to anal angle.

6. $ form protokirki forma n.

(PL 17, fig. 148)

Upperside. Fore wing basal area brownish black with strong purplish sheen, darker toward
the inner border of the discal band and on border of wing, but margin rusty brown ; discal and
postdiscal spots orange with indications of joining rays between spots in 2-4. Hind wing basal
area brownish black with purplish blue sheen ; discal band, widest at costa, is here orange, then

146 V. G. L. VAN SOMEREN

shades to ochre and whitish toward inner fold above anal angle, the outer border with strong
blue scaling. Submarginal bluish white linear marks strong; marginal border reddish above
upper tail then olive to anal angle. Underside. Purplish brown, black marks in fore wing
distinct, the tornal black spots strong. Pattern of upperside represented below but slightly
reduced and rusty ochre in colour.

7. $ form instabilis forma n.

(PL 17, fig. 147)

Upperside. Somewhat like form of the nominate diver si for ma, but base of fore wing paler
brownish with some green scaling ; discal and postdiscal orange spots fused from 5 to hind margin
but with some indication of dark scales in line of fusion to 2. Margin of wing rusty. Hind
wing basal area light brown with purply greenish sheen ; discal band broad, widest at costa and
slightly ochre-tinged here, but whitish suffused with blue at lower half; border black with
submarginal bluish white linear marks well developed; marginal border reddish above upper
tail then reddish mixed with olive to anal angle. Underside. Rusty greyish; basal black marks
thin ; pattern of upperside only slightly represented.

Range: The series of viola loandae, from which the foregoing forms have been
selected for description, was bred in the Dundo area near Loanda in the north west
district of Angola. The food plant was not specified. (Maps 6, 7).

Biological note
Many of the races have been bred on the following food plants :

Charaxes viola picta : Uganda and N.W. Kenya, Albizia coriaria, Albizia gummifera,

Albizia sp. (sassa auct.), Entada abyssinica, Entada sudanica

(Leguminosae) .
Charaxes viola kirki: Kenya and Tanganyika; Albizia schimperiana, Albizia

adianthifolia (A 1. fastigiata auct.), Entada leptostachya,

Acacia mellifera.
Char axes viola suk: Karamoja, Suk, Kamasia: Acacia mellifera, Albizia coriaria,

Albizia sp. indet.
Charaxes viola phaeus : Transvaal : Acacia caffra.

I have conducted experiments to try and find out the degree of possible inter-
change of food plants, with little success.

a. Newly hatched larvae from eggs laid on Entada sp. (Uganda) refused to feed on
Entada abyssinica (Kenya) ;

b. Larvae first fed on Albizia coriaria (Uganda) refused to feed on Albizia schim-
periana (Nairobi, Kenya).

c. Larvae from eggs laid on Acacia mellifera either refused to feed on Albizia
schimperiana, or those that did so, died in the next two instars ;

d. Captive females, caught in Acacia mellifera country where they were seen laying
on this plant, refused to lay on fresh foliage of Albizia schimperiana, but when
foliage of Acacia mellifera was supplied, they laid readily. But when the newly
hatched larvae were placed on Albizia schimperiana and Al. gummifera, few
survived to pupate.

REVISIONAL NOTES ON AFRICAN CHARAXES 147

I am of the opinion that, in the case of Charaxes viola, selection of food plant by
the female is an important factor in the build-up of localized strains. The degree of
discrimination is often rigid.

The fact that these food plants grow in different ecological environments has
brought about a restrictive choice of habitat, which almost amounts to a form of
ecological barrier between groups of the same species; and this has given rise to
certain morphological characters such as size, coloration and pattern. Cf. viola
picta/viola suk ; viola suk/ viola kirki.

Charaxes viola brainei van Son 1
(PI. 17, figs. 153-156)

Charaxes ephyra Godart ; Trimen, 1891 : 80.
Charaxes viola brainei van Son, 1966 : 3.

I have only been able to examine a single pair of this new subspecies, kindly
presented to the B.M. (N.H.) by the collector. From this meagre evidence, I am
unable to check on the characters claimed for this race, or to evaluate the statement
that this is the only form of female occurring in the northern area of S.W. Africa.

Range: Apparently confined to S.W. Africa, Kombat area. (Map 6).

SYSTEMATIC LIST

Charaxes viola Butler

Char axes viola viola Butler, 1865. Type locality: " W. Africa". Range: Senegal

to N. Nigeria and Lake Chad (?).
viola picta van Someren & Jackson, 1952. Type locality: Uganda, Unyoro,

Kitanwa. Range : S. Nigeria, Cameroon, then skirting north of

equatorial forest to E. Congo, Uganda, S. Sudan, and N.W.

Kenya.

$ f. vansomereni Poulton, 1925. Type locality: Uganda, Kampala, Bugala.
viola kirki Butler, 1881. Type locality: Tanzania, Momboia. Range: N.

and Central Tanzania and along coast to north of Ruvuma

River. Kenya, along coast and inland Ukambani, Masai

Country, Kikuyu area, Nairobi, Thika, Fort Hall, Sagana and

Nyeri.

$ f. albifascia Poulton, 1925. Type locality: Kenya, Teita Hills, Dabida.
$ f. rogersi Poulton, 1919. Type locality: Kenya, Teita Hills, Dabida.
$ f. handari Poulton, 1925. Type locality: Tanzania, nr. Mvumi.
viola suk Carpenter & Jackson, 1950. Type locality; Kenya, Suk, Kach-

eleba. Range: N. and Central Kenya.
$ f. kirkoides Carpenter & Jackson, 1950. Type locality: Kenya, Suk,

Kacheleba.
$ f. achaemenesopsis Carpenter & Jackson, 1950. Type locality: Kenya,

Suk, Kacheleba.

1 On map 6, the author's name is wrongly spelt as "van Som".

148 V. G. L. VAN SOMEREN

$ f. intermedia Carpenter & Jackson, 1950. Type locality: Kenya, Suk,

Kacheleba.

$ f. albifascia Poulton, 1926. Type locality: Kenya, Teita Hills, Dabida.
viola chanleri Holland, 1895. Type locality: Mt. Kenya, Njombeni Range.

Range: Mt. Kenya to S. Ethiopia.
viola daria Rothschild, 1903. Type locality: Ethiopia, Jalalo-Gugura.

Range : east side of Ethiopian Rift.
viola phaeus Hewitson, 1877. Type locality: Mozambique, Delagoa Bay.

Range: Southern aggregate, Mozambique to Transvaal, E.

Bechuanaland, Rhodesia and S.W. Zambia, S. Malawi, Tanzania,

east side of north end of Lake Nyasa.

$ f . coryndoni Rothschild, 1900. Type locality : Upper Zambezi, Gazungula.
f. vansoni van Someren & Jackson, 1957. Type locality: Transvaal,

Malta Forest, Pietersburg Dist. (^ Holotype), Transvaal,

Pretoria ($ Allotype).

Northern aggregate, N. Malawi and Tanzania, S. Highlands,

east side of Lake Tanganyika to S.W. Lake Victoria at Geita

and Kitigati on Kagera R. (Tanzania to Uganda border).
viola variata ssp. n. Type locality: Zambia, 60 miles on Mwinilunga-

Kabompo Rd. Range: N.W. Zambia.

$ f . tricolor forma n. Type locality : Zambia, as nominate form.
$ f . rosella forma n. Type locality : Zambia, as nominate form.
$ f. cottrelli forma n. Type locality: Zambia, Mankoya.
viola diversiforma van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga. Range: Congo, S. Katanga, Upembe area.
$ f. purpurea van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga.
$ f. viridicaerulea van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga.
$ f. caerulescens van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga.
$ f. albocaerulea van Someren & Jackson, 1957. Type locality: Katanga,

Lupweshi R.
$ f. albimacula van Someren & Jackson, 1957. Type locality: Katanga,

Kafakumba.
$ f. ochremaculata van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga.
$ f. diversiforma van Someren & Jackson, 1957. Type locality: Katanga,

Kafakumba.
$ f. cupreopurpurea van Someren & Jackson, 1957. Type locality: Lulua,

Kapanga.
viola loandae ssp. n. Type locality: N.W. Angola, Dundo area, nr. Loanda.

Range : N.W. Angola.
^ f. primitiva forma n. Type locality: N.W. Angola, Dundo area nr.

Loanda.

REVISIONAL NOTES ON AFRICAN CHARAXES 149

$ f. basiviridis forma n. Type locality: As above.

$ f . violitincta forma n. Type locality : As above.

$ f . vansonoides forma n. Type locality : As above.

$ f . loandae forma n. Type locality : As above.

$ f . protokirki forma n. Type locality : As above.

$ f . instabilis forma n. Type locality : As above.

viola brainei van Son, 1966. Type locality: S.W. Africa, Kombat. Range:

north east of S.W. Africa.
2

9. CHARAXES CYNTHIA BUTLER

The species was described by Butler in 1865, based on a male taken in Ashanti,
Ghana. He subsequently placed other material from Cameroon and N. Angola
as this same species without comment. This lead was followed by other writers
and has given rise to some confusion.

When Rothschild discussed the species in 1900, he pointed out that there appeared
to be two distinct aggregates or subspecies, the nominate of the W. African regions,
Sierra Leone to the Niger, the other occupying the area Cameroon, Congo to Unyoro
in Uganda. He set out the differences between the two very clearly, but he refrained
from giving a name to the latter group because of lack of female material.

Subsequent writers appear to have overlooked Rothschild's comments or did
not agree with them. Thus we find that:

In 1912, when Griinberg reported on the Grauer collection from Central Africa, he
described Char axes cynthia ab. mawamba from the country west of Lake Albert, in N.E.
Congo [not in Uganda], thus from the central African region as defined by Rothschild.

In 1920, Holland refers to the material collected by the American Museum Expedi-
tion at Medje, N.E. Congo, as cynthia Butler, without further comment.

In 1923, Le Cerf describes a female from Cameroon as cynthia, form albofascia.
Thus he assumes that Cameroon examples are typical cynthia cynthia Butler. He
however separates insects from Kindu, Upper Lualaba River, in the Kivu Province
of eastern Congo as cynthia kinduana, thus distinguishing them from Cameroon
specimens which he took to be nominate cynthia Butler.

In 1926, Lathy described cynthia, ab. cizeyi, which he compares with " forme
typique " [sic]. His example came from S. Cameroon. In the same paper he
described cynthia parvicaudatus from Mabira Forest, Uganda, which he compares
with examples from " d'afrique occidentale " [sic],

In 1928, Talbot described cynthia sabulosus; the type being a " dry" form and
the paratype a " wet " form, both taken in September; the type from Kinda,
Kafakumbo, Katanga, S.E. Congo (Overlaet) " distinguished by its paler brown
colouration " ; he did not state with what he compared it.

In 1933 Le Moult described cynthia guineensis from Karouane, French Guinea.
Said to have been taken further north than other described races of cynthia. The
description agrees with examples of the nominate race from Liberia, Sierra Leone
and Ghana.

2 viola fagini Storace. Type locality: Eritrea. No material available for examination. Original
description unsatisfactory. On map 6 this name is wrongly spelt "figini".

150 V. G. L. VAN SOMEREN

In 1934 Rousseau-Decelle described a male from Etoumbi, French Congo (Central
African Republic) as form angustidavius. He also described cynthia sabulosus
f. aurantiaca, from Kafakumbo, Katanga. Thus he recognizes the validity of that
subspecies, but his specimen is an extreme " dry " form.

Charaxes cynthia cynthia Bulter
(Pis. 22-27)

Charaxes cynthia Butler, 1865 : 625.
Charaxes lysianassa Westwood, 1874 : 181.
Charaxes guineensis Le Moult, 1933 : 16 syn. n.

MALE. Fore wing length 38-39 mm. Costa curved from mid-third to apex, outer border
incurved at 3-4, hind angle projecting slightly at ib. Upperside. Base of wing and costa to
end of cell, rufous chestnut, discal zone of wing black with extensions toward costa at end of
cell, the black forking at 4 and extending toward costa and enclosing two elongate rufous marks
at mid-point 5-6 in discal line; fore wing bar rufous orange, extending from costa at subapex
as spots of increasing size to hind margin, these spots to 2 usually free, rather ovoid in shape,
those of ra-ib larger and fused; there may sometimes be a suggestion of contiguous rufous
spots proximal in 2-3 in discal line. Apex and border of wing black, with marginal rufous
internervular spots largest at ib. Hind wing basal area brownish black; disc of wing crossed
by an orange ochreous bar, paler, more orange than bar in fore wing, extending from costa,
where it may be even paler, toward the inner margin, usually almost parallel-sided, with slightly
stronger rufous on distal border; postdiscal black band extending from costa to 2 and tapering
to anal angle, 6 mm. wide at 6, even on inner border but slightly dentate at upper end; border
rufous orange 3-4 mm. wide, tapering at anal angle; margin black with slight interrupted white
fringe most evident at anal angle and slightly serrate; tails thin of about equal length 4-5 mm.
long, mostly black, rufous at base. Underside. Fore wing ground colour purplish or vinaceous
brownish grey, paler more shiny bars in subcostal area separated by rufous wavy bars; post-
discal row rufous shaded, paler creamy proximally in la-ib, followed by a row of lilac-grey
lunules in the submarginal zone, rather quadrate in subapex, contiguous in ib and touching
the tornal angle black mark; black marks sub-basal in ib and 2 conspicuous as separate or
fused irregular strong marks. Hind wing ground colour as fore wing, wavy rufous lines in
basal area with paler lines in between; discal silvery creamy bar complete from costa to above
anal angle, may be uniform in width 3 mm. wide, or narrower followed by a series of rufous and
greyish lunules, with larger rufous spots distally. Submarginal row of lilac-grey and rufous
lunules conspicuous, olive at anal angle with two black dots; margin narrowly black. (PI. 22,
figs. 201, 204, 202, 205.)

Variation. There is some slight difference in intensity of the black and rufous coloration
especially in the fore wing, between fresh and partly worn examples as is to be expected, and
the underside also exhibits this. The main characteristics of this race are unaffected.
" Seasonal " forms do not seem to occur.

FEMALE. Considerably larger than the male, and paler; fore wing length 44-47 mm. Upper-
side. Costa, cell area and base rufous orange, with black mid area as in the male with some
extensions of the black into the cell, the black area beyond the cell with two rufous elongate
spots in the discal line ; pale bar orange-ochre with a rufous flush over upper spots, spots more
elongate than in the male; marginal rufous spots more conspicuous, in the black border of wing.
Hind wing pattern as in the male, but discal band wider, pale orange-ochre with rufous on outer
border tapering and crossing the inner fold; black border slightly variable in width, even on
inner border, slightly dentate on outer where it meets the marginal rufous orange border; extreme
margin black; tails long and thin 7-6 mm. Underside. Pattern much as in the male but light
areas larger, the fore wing pattern divided into a basal and sub-basal area with heavy black

REV1SIONAL NOTES ON AFRICAN CHARAXES 151

marks to discal line; discal series of spots fused with those of the postdiscal in ib-2, the remaining
postdiscal spots free up to subapex, creamy in colour; double triangular black mark at tornus,
points toward border, remaining spots to apex less strong bordered distally with greyish rufous
lunules; margin with rufous lunules interrupted by dark veins. Hind wing pattern as in the
male, but pattern bolder, rather variable in width, 5-7 mm. ; the discal bar narrower than above
but edged distally with ochreous lunules, large in upper sector but thin toward inner margin,
distally bordered with stronger rufous marks; submarginal row of lilac-grey and rufous lunules
distinct; extreme margin black; margin slightly serrate. (PL 22, fig. 203, 206.)

Range: French Guinea, Liberia, Sierra Leone, Ghana and Nigeria, west of the
Niger River. To the east it meets the next race. (Map 8).

Charaxes cynthia cameroonensis ssp. n.

(PL 27, figs. 237-242)

Charaxes cynthia f. albofascia Le Cerf, 1923 : 367.

Charaxes cynthia ab. cizeyi Lathy, 1926 : 93-94.

Charaxes cynthia f. angusticlavius Rousseau-Decelle, 1934 : 2 3 r -

I may be criticized for introducing an entirely new name for the mid-African
race of cynthia. I do so because, as indicated in the general introduction, there has
been much confusion between this aggregate and nominate cynthia in the past. The
three names mentioned above have all been given to forms or aberrations, so called,
names which have no standing and are not recognized under the Code.

MALE. Fore wing length 42-43 mm., thus larger than cynthia cynthia, and more heavily
marked. Upperside. Fore wing, the basal rufous chestnut is more intense as a rule, and the
black areas more solidly black especially in the area beyond the cell, thus almost obliterating
the rufous marks. The postdiscal bar, made up of a series of spots of increasing size from subapex
to hind margin, may show a trace of discal marks fused with the postdiscal in 2-ib., all marks
strong rufous orange. The marginal rufous internervular marks strongest at the tornus, each
with a slight white fringe. Hind wing basal area black with chestnut bloom especially toward
costa ; the discal band, slightly variable in width is strong rufous orange, as dark as the fore wing
bar; clear-cut on the inner border more dyslegnic on the outer and tending to become suffused
toward the inner fold; black band slightly varying in width, widest at upper sector, tapering
towards anal angle with outer edge rather serrate where it adjoins the rufous orange border, which
is black on the edge with very slight white fringe. Tails slightly shorter than in nominate
subspecies, sharply pointed, 4 mm. long, rufous in colour slightly margined in black. Underside.
General pattern similar to that of nominate subspecies but not so strong, in the upper half
of the fore wing, the discal-postdiscal lunules more suffused over, but whitish in la-ib but
these in the basal half may be as strongly black and rufous. On the hind wing the silvery white
line, though slightly variable, is on the whole narrower than in the western race. The base of
the costa in all races is white. (PL 27, figs. 239, 242).

<$ form angusticlavius Rousseau-Decelle, 1934 : 231

This is an extreme form, small in size, with the upperside rather dark in ground colour on
both fore and hind wings, with the strong rufous orange bars of the wings reduced in width.
The underside is also unusually dark, but the silvery discal line in the hind wing narrow but well
marked.

FEMALE. On an average larger than cynthia cynthia, fore wing length 45-46 mm. Upperside.
The basal rufous chestnut rather darker; the median black area tending to reduce or almost

152 V. G. L. VAN SOMEREN

obliterate the subcostal marks beyond the cell; the disco-postdiscal bar slightly narrower but
strongly marked, creamy or creamy ochreous; marginal rufous marks strong to moderate.
The hind wing band is generally narrower, creamy ochreous in colour ; the black border beyond
correspondingly broader, the outer border strongly dentate with the black extending along the
veins into the rufous orange marginal border, which is narrowly black on the edge; tails long
and thin. Underside. Pattern essentially similar to that of nominate cynthia but bolder, the
discal and postdiscal pale marks of the fore wing and the discal band on the hind wing paler,
more creamy, and more strongly marked. (PI. 27, figs. 237, 238, 240, 241.)
Variation.

(a) The female figured on PI. 27, figs. 237, 238, is the type of ab. cizeyi Lathy and conforms
with the majority of females of this subspecies.

(b) Another female would appear to agree with albofascia Le Cerf . This has the fore and hind
wing bars pale creamy to whitish.

s

f!_ ' ,- '

,. ,>.^-'..-^. ; '.,"

'-< , cynthia cynthia- ; ;

-' ' *& ' >^ TT

GHANA, '

NIGERIA gja CENTRAL \

corheroonfl

^AFRICAN REPUBLIC-*.

', sobulosu
ANGOLA \ KATANGA
! . \W

MAP 8.

REVISIONAL NOTES ON AFRICAN CHARAXES 153

(c) A few females exhibit a degree of partial " melanism "; the bars of the fore wing and the
band on the hind wing are more strongly orange and dusted over with brownish scales. The
underside is not greatly affected, but in the specimen from Moyen Congo the submarginal and
marginal lunate pale marks are exaggerated.

Although I have linked the specimens of the Central African Republic and Moyen Congo
with the Cameroon subspecies, since the majority in the aggregates agree, one finds some degree
of variation in all three areas.

Range: Cameroon (and adjacent eastern S.E. Nigeria where there is some contact
with nominate cynthia} to Central African Republic (French Congo), the Moyen
Congo to the Congo River basin; making some contact with kinduana in west
central Congo. (Map 8. On this map the specific name is given in error as
" cameroona ".)

Charaxes cynthia kinduana Le Cerf

(PI. 26, figs. 231-236)

Charaxes cynthia kinduana Le Cerf, 1923 : 366.
Charaxes cynthia, ab. mawamba Grunberg, 1912.

Through the kindness of Dr. P. Viette of the Paris Museum, I have obtained a
photograph of the type of kinduana Le Cerf. This depicts a very dark specimen
which appears to be semi-melanistic ; the black area of the fore wing is more extended
than usual, occupying the basal half of the wing with the exception of the cell area ;
the usual subcostal rufous spots are entirely obliterated; the postdiscal series of
spots are reduced in size and thus well separated to 2 and dark rufous ; the marginal
spots absent except at the tornus. The hind wing band is reduced in width, the
upper marks well separated and darker brownish rufous; the marginal border is
narrower and darker. Underside. Pattern of both wings is obscured except for
the black marks at the base of the fore wing and the pale area in la-ib; the hind
wing pattern obscured except for the conspicuous silvery white discal line which is
very narrow.

Since the type specimen appears to be abnormal, it becomes difficult to assess
what are the normal characteristics of this race. The position is further complicated
in that in 1912 Grunberg described what he called cynthia ab. mawamba from what
is generally accepted to be the same ecological area whence came the type of kind-
uana Le Cerf. The description of mawamba agrees more closely with the general
run of males of this area than does that of kinduana. The description which follows
is based on material from the eastern Congo.

MALE. Fore wing length 44-45 mm. Upperside. General pattern as in the more western
ssp. cameroonensis, but basal rufous colour slightly paler, the subcostal rufous spots distinct
though small, (slightly obscured in some specimens) ; the disco-postdiscal rufous orange spots
separated from subapex to 3, the rest to hind margin larger and conjoined, with some indication
of discal marks in 2-3; marginal rufous spots present but most distinct at the tornal angle.
Hind wing basal area brownish black, paler on inner fold; discal band widest at costa 5-6 mm.
then tapering rather strongly toward, but not reaching the inner fold; black band thus wider
at costal end, and tapering toward anal angle; rufous orange marginal border wider, edged
narrowly with black. Tails 4 mm. Underside. Very similar to cameroonensis but rather less

154 V. G. L. VAN SOMEREN

strongly marked with rufous Gti the fore wing, black marks rather overshadowed by darker
ground ; hind wing white bar somewhat variable in width ; postdiscal rufous lunules, with dark
band distally, show up plainly; submarginal row of pale marks indistinct; margin rufous.
(PI. 26, figs. 231, 234, type.)

FEMALE. In general pattern and colour somewhat similar to nominate cynthia but pale bars
on fore and hind wing narrower. Fore wing marginal rufous spots strong; disco-postdiscal
bar rufous orange, rather narrow. Hind wing discal band rufous orange, widest at costa,
and tapering at 2 to above anal angle, slightly dentate in 4 on outer border; the black postdiscal
band widest at 4-6, dentate on outer border and invading the rufous orange border along veins;
margin narrowly black; tails longish, thin, upper 6 mm., lower 4 mm. Underside. Ground
colour vinaceous grey with a rufous flush. Fore wing chestnut bars distinct but narrower; disco-
postdiscal spots creamy ochre, more whitish in lower half flanked by black lines proximally and
by the black tornal spots distally, the other submarginal dark dots to subapex more diffuse.
Hind wing band more creamy ochre, clear-cut on inner border but merging into the
postdiscal greyish and rufous lunules distally, the latter strongly marked; submarginal
lunules large; marginal rufous orange border distinct. (PI. 26, figs. 232, 235, 233, 236.)

Range: eastern Congo from the Kivu Province area along the Lualaba River to
Stanleyville, Medje, Avikuba-Aruwimi, extending westward to the mid Congo
basin, and eastward to the Semliki Valley. (Map 8).

Charaxes cynthia parvicaudatus Lathy

(PI. 24, figs. 215-222)
Charaxes cynthia parvicaudatus Lathy, 1926 : 94.

MALE. Fore wing length 35-40 mm., thus rather smaller than other races so far mentioned.
Upperside. General pattern basically similar, but differing as follows: rufous areas at costa
and base of fore wing paler; the black mid area represented in the cell by an almost free mark;
the two rufous spots in discal line, beyond, usually distinct; the rufous orange bar with the
subapical and other spots to 4 completely free, the spots of increasing size, those of 2-3 showing
some indication of fusing with trace of marks in discal line, the mark in ib more quadrate, that
in la a streak, but the whole bar is relatively narrow. Marginal rufous marks small and indis-
tinct except the double one at the tornus. Hind wing basal area rather more brownish, darkening
to almost blackish at the discal band ; the discal band is narrow, 5 mm. at the costa, irregularly
parallel-sided and stopping short of the inner fold, rufous orange in colour, slightly paler at
costa. The black postdiscal band is thus wide, fairly even on inner border but serrate on outer
margin invading the outer border along the veins, this border rufous orange with slight black
edge. Tails stumpy, 2-3 mm. long, mostly rufous. Underside. Ground colour as in most
other races, but with a rusty bloom overall, the rufous pattern less strong, but black marks in
ib well developed; the disco-postdiscal series of pale spots rather obscured in upper portion
but whitish in ib; the black tornal triangles not very heavy. Hind wing pattern not strong
in the basal area; the white bar narrow, clear-cut on inner border, more diffuse on outer where
it merges into the inner row of postdiscal lunules; a series of white dots in submargin at apices
of marginal lunules which are not usually strong; border rufous. (PI. 24, figs. 215, 216, 219, 220.)

FEMALE. Upperside. In general appearance resembling somewhat the females of cynthia
cameroonensis in respect to the pale bars through the wings. Fore wing length 45-46 mm. ; basal
rufous area at la-ib and over cell rather paler; the ochreous spots at the end of the cell and
in the discal line, distinct and elongate, the upper postdiscal spots, rounded or elongate, the
remaining spots in this line increasing in size and fused with spots in the discal line which may
be rudimentary or of equal size, the mark in ib rather quadrate. Marginal rufous marks large
and conspicuous. Hind wing basal area brownish; the discal band rather parallel-sided up
to cell area then tapering to inner fold, creamy ochreous in colour; the distal black band wide,

155

more or less straight on the inner side but serrate on the distal, with strong projections along
the veins into the rather narrow rufous marginal border; edge narrowly black; tails relatively
short, upper rather spatulate at end 6 mm. long, lower shorter, more stumpy 4 mm. Underside.
Conforming to the general tone of the ground colour of the male; the disco-postdiscal spots
forming the bar in the fore wing clearly indicated, creamy in colour; the black marks in ib
often strong and conjoined. The hind wing band creamy, strongly denned on inner border,
less strong on outer where it merges with the inner lunate pale marks of the postdiscal zone,
the adjacent brownish rufous zone rather well marked ; the submarginal pale marks flanked
proximally with rufous triangles; border rusty brown. (PI. 24, figs. 217, 218.)

Variation. In some specimens the rufous areas of the fore wing may be dusted over with
brownish scales; the marginal spots obscured. On the hind wing, the rufous marginal border
reduced in width.

Specimens from N.W. Kenya in the Kakamega-Kaimosi forests belong to this
aggregate. (PL 24, figs. 221, 222.)

Range: eastern Uganda, especially in the Mabira Forest and adjacent forest
areas extending eastward in the forests of N.W. Kenya, Kaimosi-Kakamega-Nandi.
(Map 8).

Charaxes cynthia propinqua ssp. n.

(PI. 23, figS. 209, 2IO, 213, 214)

This aggregate combines the features of parvicaudatus as regards shape and more
particularly the very short tails, with the eastern Congo subspecies kinduana with
regard to colour and pattern of the fore wing on upper side.

MALE. Fore wing length 36-38 mm. Upperside. Fore wing pattern bolder than in
parvicaudatus, base of wing and costa rufous slightly more extended; the subcostal rufous
spots in discal line and those of the postdiscal row larger throughout but more especially in the
hind half of the bar; the rufous marginal spots larger and more pronounced. The hind wing
discal band is wider, thus reducing the black band beyond; the marginal rufous border is wider.
Underside. Fore wing pattern very similar to parvicaudatus, in general tone; black marks at
base of ib confluent, thus large and conspicuous; the hind wing white bar is narrow and more
defined on both borders, the pattern is otherwise very similar. The tails are extremely short
and stumpy. (PI. 23, figs. 209, 210, type.)

FEMALES. The characteristics of the male reproduced in the female: fore wing length 45-47
mm. Upperside. Basal area bright rufous; rufous spots beyond cell and those in discal
line usually bolder; the upper postdiscal spots large, those of 3 to hind margin large and fused
with the discal spots so that the fore wing bar is wider; the marginal rufous spots bold. Hind
wing band wider from costa to 2, the black band thus reduced in width; marginal rufous border
wider than in parvicaudatus; tails thinner, upper 6 mm. lower 4 mm. with a slight upward kink
as in parvicaudatus. Underside. Very similar to parvicaudatus in the fore wing; on the hind
wing the white bar is more defined. (PI. 23, figs. 213, 214, type.)

Holotype male. UGANDA: Katera Forest, Masaka, xi.i953 (van Someren).

Allotype female. Same data.

Range: From the forests of western Uganda, Bwamba, Toro, Bugoma, Budongo,
to the forests west of Lake Victoria in the Masaka district, Katera, and north of the
Kagera River. (Map 8).

156 V. G. L. VAN SOMEREN

Charaxes cynthia sabulosus Talbot
(PL 25, figs. 223-230)

Charaxes cynthia sabulosus Talbot, 1928 : 229.

Charaxes cynthia sabulosus f. aurantiaca Rousseau-Decelle, 1934 : 231.

Specimens of cynthia from the Katanga area present a rather puzzling picture for it
would appear that the species is here subject to " seasonal variation " [sic] producing
so-called " wet " and " dry " forms. This gives rise to some difficulty in determining
the characteristics of the subspecies as a whole. In the original description Talbot
states " a well defined race from Katanga "; but the description is involved since
it concerns two specimens, the type and a paratype which are slightly different.
I have before me a small number of topotypical specimens which exhibit marked
variation, in both sexes. It seems desirable therefore to quote the original description
of the type.

Fore wing length 40 mm., " distinguished by its paler brown coloration on both sides. Fore
wing above with the postdiscal brown band of spots continuous to cellule 7, the spots above
vein 4 larger than in typical form, and spots in 2 and 3 not invaded by the ground colour . . .
Band of the hind wing broader than in typical form, being extended distad. Brown scaling
in the cell more marked than in typical form . . . Underside with the dark brown markings
obsolete and black spots smaller. . . . The white band on the hind wing is sharply defined
along both its edges. ..."

Type (J, Kinda, Katanga (Overlaet).

The characters of the paratype are: " the marginal spots [f.w.] are merged together to form
a continuous broad marginal line . . . the hind wing cell is strongly brown. . . . The white band
of underside only reaches to just below vein 2."

I have before me two topotypical males taken by Overlaet at Kafakumba, Katanga. One
is described as a " dry form ", the other as a " wet form ".

" Dry form ": Upper side. Fore wing rufous. Coloration extends along the costa, over the
whole cell and slightly beyond and in the bases of xa-ib and slightly in 2-3 so that the black
mark at end of cell is entirely free. The black median band is thus narrow, but the two upper
rufous spots in the discal line are large and free. The rufous orange bar is continuous from 7 to
the hind margin, the spots increasing in size, those in 2-3 with slight indication of remnants of
discal marks. The rufous marginal marks bold. The hind wing discal band is moderately
broad, 6 mm. at costa, almost parallel-sided, tapering out at vein 2 ; black band widest at 5-6, even
on inner edge, slightly serrate on outer; rufous border rather wide; tails thin, upper 7 mm.,
lower 5 mm. Underside. Vinaceous grey-brown, rufous basal bars not strong; black sub-basal
marks in ib relatively small, tornal mark moderate; pattern of disco-postdiscal bar rather
obscured. Hind wing with most of the pattern obscured except that of the white discal line
which is narrow but strong, clearly defined on both edges, and almost straight. (PI. 25, figs. 229,
230.)

" Wet form ": Upperside. Fore wing basal rufous coloration rather darker and slightly less
extensive; black median zone considerably wider, the two subcostal spots in the discal line
almost obscured ; distally, the band encroaches onto the upper postdiscal row of spots which are
considerably smaller, so that the whole bar is narrower and darker rufous than in the " dry "
form; the rufous marginal spots are very small, except those at the tornal angle. The hind
wing basal area darker; the discal band narrow and stops short at 2; the black band wider than
in the " dry " form so that the rufous marginal border is restricted in width. Underside.
Ground colour slightly darker, but the pattern is stronger.

REVISIONAL NOTES ON AFRICAN CHARAXES 157

Both these specimens were taken in September. According to Chapin, the rainy
season in Katanga is October to May, the dry from June to September.

A further dark form, " wet " presumably, was taken at Kindo, Katanga, in April. Upper-
side. In general appearance it resembles the previous specimen, except that the rufous subcostal
spots are obscured to a greater degree; the postdiscal bar narrower in la-ib; the marginal spots
slightly more developed. In the hind wing the discal band is rather narrower and constricted
in 6. Underside. Very similar to the previous specimen. (PI. 25, figs. 223, 224.)

form aurantiaca Rousseau-Decelle, 1934 : 231
(PI. 25, figs. 227, 228)

In sharp contrast to the two foregoing, we have the very " dry " male form
described by Rousseau-Decelle and named aurantiaca which also came from Kafa-
kumba in Katanga. No date of capture is mentioned for any of the three examples
cited.

MALE. Upper side. Pale rufous orange on costa, the cell and just beyond, the bases of ia-ib,
slightly in 2-3, thus enclosing the black mark at end of cell which is entirely free; there are two
spots beyond the cell in the discal line; the postdiscal bar consists of a small subcostal spot,
then a series of larger spots increasing in size to the streak in la, the bar is wider than in other
forms and pale rufous orange; margin with large conspicuous marks from apex to hind angle.
Hind wing basal area rufous slightly darker at base of 6, discal band light rufous orange, pale
at costa but darkening in 2 and along the outer border; black band, comparatively narrow irreg-
ular on both sides; marginal border wide, pale rufous orange. Tails as in other specimens.
Underside. Vinaceous grey with rusty suffusion, the pattern almost obsolete even to the dark
marks in ib. The hind wing basal area almost uniform followed by a distinct narrow white
bar stopping short at 2, the remainder of the pattern obscured, except for a series of pale rounded
marks in submargin. (PL 25, figs. 227, 228.)

FEMALE. The female does not seem to have been described and the only one available to me
appears to belong to the " wet " form. Upperside. Fore wing rufous basal area extending over
the cell and slightly beyond with only slight rufous tinge at base of ib; the black spot at end of
cell almost free. Fore wing bar rather wide since the upper spots of the postdiscal line are large
with gradual increase in size to hind margin, spots in 2-3 with slight conjoined marks of the discal
line, the spots of this line in 5-6 well developed, large. The marginal lunules large and distinct.
Hind wing basal area brownish, more rufous toward inner fold; discal band almost uniform in
width from costa to 2 then tapering to inner fold above the anal angle. Black band moderately
broad, even on inner side very slightly dentate at upper part on distal side; marginal border
fairly wide with very narrow black edge; upper tail robust with rounded end 6 mm., same colour
as border, lower tail of about same length, darker, thinner, bluntly pointed ; anal angle with two
white dots. Underside. Fore wing basal area vinaceous grey with rufous bars and black
marks strong; disco-postdiscal bar fairly clearly indicated, creamy, but whitish in ia-ib;
submarginal pale marks with dark spots proximally fairly strong; marginal lunules well
represented. Hind wing ground colour as fore wing in basal area but dark lines strong, especially
that adjoining the discal white bar which is thus defined on the inner edge but rather merging
with the irregular lunules on the distal side ; dark brownish band strong, less so on outer edge
where it abuts onto the marginal rather ill-defined pale lunules; border narrowly rufous.
(PL 25, figs. 225, 226.)

From the above descriptions it will be noted that the subspecies sabulosus is
represented by a rather mixed aggregate, with combined features which separate
it from other races.

ENT. 23, 4 13

158 V. G. L. VAN SOMEREN

Range: All the specimens I have seen and those on record appear to have been
taken in the Katanga area of S. Congo, at Kinda, Kafakumba and Thisboba so that
its wider range is thus unknown. (Map 8).

Charaxes cynthia rnukuyu ssp. n.

(PI. 23, figs. 207, 208, 211, 212)

The series of specimens on which this race is based represent one of the most
distinctive subspecies of cynthia. In some respects it resembles the race sabulosus
but is easily distinguishable both above and below. It is characterized by the great
reduction of the black areas on the upper side of both fore and hind wing ; the paler
more orange of the rufous areas. On the underside, the pattern is suppressed, includ-
ing the white bar on the hind wing which, in some specimens is almost vestigial;
in such cases there is in fact a slight resemblance to Charaxes buetti macclounii.

MALE. Fore wing length 39-40 mm. Upperside. Costa and basal area bright rufous
orange extending well beyond the end of the cell, the basal areas of la-ib, 2 and 3, the black
mark at end of cell much reduced to an inverted comma ; the black central band thus much re-
duced in width; the two subcostal marks in the discal series large, quadrate and contiguous, with
clear indication of spots in 2-3, the latter often free, and an indication of black in ib showing
line of junction of the marks in this area; the postdiscal spots from subcosta to 4 of about even
size, those from 3 to hind margin increasing is size ; the bar thus appears divided in upper sector ;
marginal spots usually fairly large especially at the tornal angle. Hind wing basal triangle
rufous brown, slightly darker in mid area and where it meets the discal rufous orange band
which is slightly expanded at the costa, more or less parallel-sided to above 2 where it tapers
and fades out before reaching the inner fold. The black postdiscal band is reduced in width,
widest in 6-7 where the outer border is serrate then tapering to above the hind angle, where, the
lower spots may be free or slightly conjoined; the rufous orange marginal border is thus wide
and very narrowly black-edged; tails short and pointed, of about equal length, 3-4 mm.
Underside. Ground colour rather pale rufous ochreous with less vinaceous shade; the rufous
bars finer and less obvious; the black marks less strong, but those in ib often conjoined to form
a rough oval ; the tornal mark rather blurred. The disco-postdiscal pattern of spots not strong,
though whitish in la-ib. On the hind wing the discal white line is thin and though dark edged
on inner side may be much reduced ; the postdiscal lunules, except toward anal angle obscured ;
pale submarginal lunules obscured; the rufous border not strong. (PI. 23, figs. 211, 212.)

FEMALE. Fore wing length 42-44 mm. Basal area including the cell and base of ib and
slightly that of 2-3 bright rufous orange, extending beyond end of cell and here enclosing the
black inverted " comma " mark as in the male. Mid bar narrow black and interrupted in
base of 4 by a quadrate creamy ochre mark, part of the discal series which starts as two elongate
quadrate marks in 5-6, represented in 3 by a crescentic mark, the lower mark fused with the
postdiscal row of large creamy ochre marks extending to the hind margin in la; margin with
conspicuous rufous marks which are almost contiguous. Hind wing basal triangle rufous brown,
rather restricted owing to the width of the discal band which is almost parallel sided, but tapering
to above the anal angle, the outer edge tinged with rufous ; the black band narrow, constricted
at costal end, outer border serrate at upper end, tapering to anal angle where there are two
white dots. Marginal border rufous, wide, narrowly black edged; tails rather thin, upper
6 mm. lower 4 mm. slightly up-curved. Underside. General tone similar to that of the male;
the fore wing rufous bars thin; the black mark sub-base in ib an oval, the tornal mark moderately
strong; the disco-postdiscal bars broad rather diffuse except in la-ib; margin diffusely rufous.
Hind wing, basal area as fore, broad discal band sharp-cut on inner edge but diffuse on outer
where it merges into the pale lunules which are only slightly visible, the dark band is outwardly

REVISIONAL NOTES ON AFRICAN CHARAXES 159

edged with paler lunules; the border rufous, with two black spots in anal angle. (PL 23, figs.
207, 208.)

Variation. FEMALE. Upperside. Pattern for fore wing very similar to nominate mukuyu
but basal rufous coloration darker; the black triangle at bases of ia-3 broader at base.
Disco-postdiscal series of creamy ochre marks as in the other female but broader; marginal
lunules less defined. Hind wing basal are darker rufous brown; discal band wide, broadest at
costa and tapering to above angle ; the black band still narrow consisting of a series of contiguous
oval black marks, small at costa, and extending to anal angle; rufous border broad; tails
shorter. Underside. Ground colour as in other female but pattern stronger and more defined ; this
applies particularly to the creamy bands on both wings and especially to the hind wing where
the postdiscal pale and rufous lunules are strong.

Holotype male. TANZANIA: Kigoma District, at Mukuyu Forest, .1965 (Major
/. Grahame}, deposited in B.M. (N.H.)

Allotype female. Same locality, vii . 1965 .

Material examined, nine males and two females taken by African collector for
Major Grahame and by the Japanese Primate Expedition. (Map 8).

SYSTEMATIC LIST

Charaxes cynthia Butler

Charaxes cynthia cynthia Butler, 1865. Type locality: Ashanti. Type in B.M.

(N.H.)c?.

= lysianassa Westwood, 1874. Type locality: Ashanti. $.

= guineensis Le Moult, 1933. Type locality: French Guinea. $.

Range: W. Africa, Guinea, Liberia, Ivory Coast, Ghana to Nigeria to

Niger River.
cynthia cameroonensis ssp. n.

= ab. cizeyi Lathy, 1926. Type locality: Cameroon. $.

= f. indiv. angusticlavius Rousseau-Decelle 1934. Type locality:

French Congo, Etumbi. <$.

= f. indiv. albofascia Le Cerf, 1923. Type locality: Cameroon, Bit je. $.

Range: Cameroon rain forests, S.E. Nigeria, Central African Republic.

Moyen Congo, west of Congo Basin.
cynthia kinduana Le Cerf, 1923. Type locality: E. Congo, Kinda on upper

Lualaba River. Type in Paris Mus. $.

= ab. mawamba Griinberg, 1912. Type locality: eastern Congo,

Awamba, west of Lake Albert; en route Awamba-Avukuba-Aruwimi.

Range : From the Kivu Prov. of eastern Congo, along the Lualaba River

to Medje N.E. Congo, west to Congo Basin; east to Semliki Valley.
cynthia propinqua ssp. n. Type locality: Uganda, Katera Forest.

Range : W. Uganda, Toro forests, Bugoma, Budongo, Bwamba, Kalinzu,

Katera, Entebbe.
cynthia parvicaudatus Lathy, 1926. Type locality: Uganda, Mulange,

Mabira Forest.

Range: Uganda ; Busoga, Mabira Forest, Jinja, Kenya; N.W. Kapwaren

forests, Kaimosi, Kakamega, Kabras, Nandi, Kipkarren.

ENT. 23, 4. J 3

i6o V. G. L. VAN SOMEREN

cynthia sabulosus Talbot, 1928. Type locality: Kinda, Katanga. <^.

Tervuren Mus.

f. indiv. aurantiaca Rousseau-Decelle, 1934. Type locality: Katanga,

Kafakumba.

Range: Katanga, at Kinda, Kafakumba, Thisbobo. Upembe Park.

lower Kasai.
cynthia mukuyu ssp. n. Type locality. Tanzania, Kigoma, Mukuyu.

Range: Tanzania, N.E. of Lake Tanganyika in the Kigoma district, at

Mukuyu and Muhimo forests.

ACKNOWLEDGEMENTS

A vast amount of topotypical material was brought together for these revisional
notes and I wish to acknowledge my indebtedness to the following for loan of material,
photographs of types, etc. I offer my apologies to some contributors for retaining
their material for such a lengthy period.

The staff of the Entomological Department, B.M. (N.H.), London; Mr. E. Taylor
of the Hope Dept., University Museum, Oxford, England; Major Iain Grahame,
Lamarsh, Suffolk, England; Dr. Arthur Rydon, N. Chailey, Sussex, England; Dr.
Viette and Dr. Bernardi, Natural History Museum, Paris ; Dr. Berger, Central African
Museum, Tervuren, Belgium; Dr. Kasy, Vienna Museum, Austria; Monsieur J.
Plantrou, of Paris; Dr. Fox, Carnegie Museum, Pittsburgh, U.S.A.; Dr. Hannemann,
Natural History Museum, Berlin; Dr. E. Pinhey, National Museum, Bulawayo;
the late Dr. G. van Son, Transvaal Museum, Pretoria, S. Africa; Mr. K. M. Pennington
Balgowan, Natal; Mr. D. C. Plowes, Umtali, Rhodesia; Dr. C. B. Cottrell, Salisbury,
Rhodesia; Mr. H. D. Handman, Zomba, Malawi; Mrs. Joan Wedekind, Mumbwa,
Zambia; Mr. B. Barton-Eckett, Bathurst, Cape Prov., S. Africa; Dr. C. H. McCleery,
Lindi, Tanzania; Mr. J. H. Bailey, Menengai, Kenya; the late Mr. T. H. E. Jackson,
Kitale, Kenya; Mr. R. H. Carcasson, National Museum, Nairobi, Kenya; and Mr.
H. D. van Someren, Endebess, Kenya.

I desire also to record my thanks to the following for financial assistance toward
the cost of photographic work connected with these revisions: Dr. Arthur Rydon,
N. Chailey, Sussex, Major Iain Grahame, Lamarsh, Suffolk; and Mr. C. G. C. Dickson
of Cape Town, S. Africa.

REVISIONAL NOTES ON AFRICAN CHARAXES

161

8

FIGS. 1-9. Aedeagi of Char axes spp. i, etheocles etheocles (Cramer), Nigeria (Prep. 64,
A. H. B. Rydon) ; 2-4, etheocles carpenteri Poulton, $ f. catachrous, Katera, Uganda (Preps.
669, 670, T. G. Howarth; prep. 55, A. H. B. Rydon); 5-6, etheocles evansi van Someren,
Kitale, east Mt. Elgon, Kenya (Preps. 61-62, A. H. B. Rydon) ; 7-8, grahamei van Someren,
Kigoma Dist., N. E. Lake Tanganyika, Tanzania (Preps. 672, 671, T. G. Howarth);
9, mafuga van Someren, Mafuga Forest, Kigezi, Uganda (Prep. 10, A. H. B. Rydon).
Del. A. H. B. Rydon.

1 62

V. G. L. VAN SOMEREN

11

16

18

FIGS. 10-18. Aedeagi of Charaxes spp. 10-11, berkeleyi van Someren & Jackson, Karen,
Ngong, Kenya (Preps. 56-57, A. H. B. Rydon) ; 12, aubyni aubyni Poulton, Teita Hills,
Kenya (Prep. 173, A. H. B. Rydon); 13-14, baileyi van Someren, Visoi Gap, west Rift
Valley, Kenya (Preps. 51-52, A. H. B. Rydon); 15-16, pembanus Jordan, Pemba Island,
N.E. of Zanzibar, East Coast (Prep. 170, A. H. B. Rydon), lateral and dorsal views;
17-18, usambarae van Someren & Jackson, Amani, lower forests, UsambaraMts., Tanzania
(Prep. 169, A. H. B. Rydon), lateral and dorsal views. Del. A. H. B. Rydon.

REVISIONAL NOTES ON AFRICAN CHARAXES

163

FIGS. 19-26. Aedeagi of Charaxes spp. 19, karkloof van Someren & Jackson, Mt. Kark-
loof, Natal, S. Africa (Prep, van Son) ; 20, credreatis Hewitson, $ lutacea Rothschild,
Kisubia Forest, nr. Entebbe, Uganda (Prep. 175, A. H. B. Rydon); 21, alpinus van
Someren & Jackson, Vumba Mts., Rhodesia (Prep, van Son) ; 22, alpinus nyikensis van
Someren, Nyika Plateau, Malawi (Prep, van Son) ; 23, ethalion littoralis van Someren,
Marere Forest, Shimba Hills, Kenya Coast (Prep. 176, A. H. B. Rydon); 24, ethalion
ethalion Boisduval, Natal, South Africa (Prep, van Son) ; 25, petersi van Someren, Liberia
& Ivory Coast (Prep. 50, A. H. B. Rydon); 26, Charaxes sp. nov.? Bunduki Hill,
Uluguru Mts., Tanzania (Prep. 164, A. H. B. Rydon). Del. A. H. B. Rydon.

i6 4

V. G. L. VAN SOMEREN

30

FIGS. 27-30. Aedeagi of Charaxes spp. 27-28, contrarius Weymer, Zombo Forest, Kwale
Dist., Kenya Coast (Preps. 58-59, A. H. B. Rydon); 29, viola kirki Butler, Mackinnon
Road, Coast belt, Kenya (Prep. 177, A. H. B. Rydon); 30, manica Trimen, Mpanda,
Tanzania (Prep. 3, A. H. B. Rydon). Del. A. H. B. Rydon.

FIG. 31. Charaxes kheili Staudinger. Type $ genitalia. B.M. Neg. No. 8749.

V. G. L. VAN SOMEREN 165

REFERENCES
References not given below will be found in Parts I-IV of this revision.

BUTLER, A. G. 1876. On a Collection of Lepidoptera recently received from Abyssinia. Ann.

Mag. nat. Hist. (4) 18 : 480-490.
1881. On a new species of Charaxes from W. Africa. Entomologist's mon. Mag. 18 : 107

108.
1893. Description of a new species of the butterfly genus Charaxes. Ann. Mag. nat. Hist.

(4) 12 : 266.
FEISTHAMEL, M. 1850. Description de quelques Lepidopteres Rhopaloceres nouveaux ou

peu connus provenant de la Cazamance (Afrique). Annls Soc. ent. Fr. 1850 : 247-262.
GRUNDBERG, K. 1911. Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika Expedition

1907-1908. 3(17) : 506-560. 4 pis. Leipzig.
HEWITSON, W. C. 1874. Description of a new species of Charaxes from the West Coast of

Africa. Entomologist's mon. Mag. 10 : 247-248.

1876. Illustrations of new species of Exotic Butterflies. 5 : 1-116, 60 pis. London.

1877. Descriptions of three new species from Delagoa Bay. Entomologist's mon. Mag.
14 : 81-83.

HOLLAND, W. J. 1896. List of the Lepidoptera collected in East Africa 1894, by Mr. William

Astor Chamber and Lieutenant Ludwig von Hohnel. Proc. U.S. natn. Mus. 18 : 741-767.
JACKSON, T. H. E. 1957. Notes on the Rhopalocera of the Kigezi District of Uganda with

descriptions of new species and subspecies. // E. Africa nat. Hist. Soc. 23 : 63-78, 13 pis.
LE MOULT, E. 1933. Formes nouvelles ou peu connus de Charaxinae (Lep. Nymph.). Novit

ent. 2 : 16-18.
ROUSSEAU-DECELLE, G. 1934. ^ e quelque formes nouvelles de Charaxes africaines. Bull.

Soc. ent. Fr. 39 : 228-235, i pi.

STONEHAM, H. F. 1964. Butterflies of Western Kenya. Pts. 8-9 : 109.
TALBOT, G. 1928. New forms of African Nymphalidae (Lep. Rhop.) in the Musee du Congo,

Tervuren. Bull. Hill. Mus. Witley2 : 229-231.

TRIMEN, R. & BOWKER, J. H. 1887. South African Butterflies 1 : 348. London.
VAN SOMEREN, V. G. L. 1958. A new species and a new female form of " Charaxes " from Kenya

Colony. Novos Taxa ent. 11 : 1-12, 2 pis.

1963. Revisional Notes on African Charaxes (Lepidoptera : Nymphalidae). Part I.
Bull. Br. Mus. nat. Hist. (Ent.) 13 (7) : 195-242, 19 pis., 5 text-figs.

1964. Part II. Bull. Br. Mus. nat. Hist. (Ent.) 15 (7) : 181-235, 2 3 P^ 8 -. 4 maps.

1966. Part III. Bull. Br. Mus. nat. Hist. (Ent.) 18 (3) : 45-100, 16 pis., 5 maps.

19660. Part IV. Bull. Br. Mus. nat. Hist. (Ent.) 18 (9) : 277-316, 9 pis., 4 maps.

VAN SON, G. 1966. A new subspecies of Charaxes viola Butler from South- West Africa.

Novos Taxa ent. 49 : 3-7.
WESTWOOD, J. O. 1874. Thesaurus entomologicus Oxoniensis, pp. 1-205, 40 pis. Oxford.

INDEX
New taxonomic names in bold type

achaemenesopsis, $ f., 132 berkeleyi, 80

acuminatus, 77 biinclinata, 102

albifascia $ f., 131, 133 brainei, 147
albimacula $ f., 143

albocaerulea $ f., 143 caerulescens $ f., 80, 143

alladinis $ f., 100, 105 cameroonensis, 151

angusticlavius $ f., 151 carpenteri, 108, 109

aurantiaca $ $ ff., 157 carteri <$ f., 102

catachrous <$ f., 102, 106, in

baileyi, 122 cedreatis, 84, 85

basiviridis $ f., 145 chanleri, 133

1 66

INDEX

chepalungu, 90

conjugens $ f., 121
conjuncta $ f., 113
contrarius, 119, 120
coryndoni $ f., 137
cottrelli $ f., 140
cupreopurpurea $ f., 144
cynthia, 149, 150

daria, 134
dewitzi $ f., 90
diversiforma, 141, ? f., 143
druceanus, 78

ephyra <$ f., 101
etheocles, 99
evansi, 112

fulgens $ f., 101
grahamei, 115

handari $ f., 131
hollandi $ i., 101

inexpectata $ f., 88
instabilis $ f., 146
intermedia $ f., 133

kheili, 94
kilimensis 82
kinduana, 153
kirki, 130
kirkoides $ f., 132

lacteata $ f., 117
loandae, 144, $ f., 145

mafuga, 97
masaba, 82

mukuyu, 158

ngonga $ f., 80
northcotti, 96

obudoensis, 77
occidens, 79
ochracea, 106
ochremaculata $ f., 143

pallidimacula $ f., no
parvicaudatus, 154
penricei, 80
petersi, 121

phaeus, 136, $ f., 137, 138
picta, 128

primitiva ? f., 144
propinqua, 155
protocedreatis $ f., 87
protokirki $ f., 145
pseudocarpenteri $ f., 124
pseudosmaragdalis $ f., 89
purpurea $ f., 141
pythodoris, 79

regalis 9 f., 101, 104
rogersi $ f., 131
rosella $ f., 140

sabulosus, 156
schoutedeni, 79
seriata $ f., 107
suk, 132
tanganyikae, 80
tectonis, 78
tricolor $ f., 140
tincta $ f., 145

vansomereni $ f., 129
vansoni $ f., 137
vansonoides $ f., 145
variata, 138, $ f., 139
vetula $ f., 88
viola, 125, 127
violitincta $ f., 145
viridicaerulea $ f., 141
virilis, 92

xiphares, 82

PLATE i
Charaxes cedreatis Hewitson

FIGS, i & 2. (J = lutacea Rothschild Type. (N.E. Congo: Beni. [B.M. (N.H.)], upper and

underside. Photos B.M. (N.H.) Nos. 39912; 39913.
FIGS. 3 & 4. ? Type. (Fernando Po.) [B.M. (N.H.)], upper and underside. Photos B.M.

(N.H.) Nos. 39918; 39919.
FIG. 5. <J with strongly developed subcostal spots to fore wings and large glaucous lunules

on margins; base of fore wing strongly greenish; hind wing with postdiscal wavy

green line. Underside pattern not strong and suffused brownish. (Tanzania

[Tanganyika], Geita), upper and underside.
FIG. 6. <J boldly marked on upperside; hind wing with strong greenish postdiscal line;

underside pattern strong and colder greyish ground colour with satiny bars

strong. (N. Angola, Amboim), upper and underside.
FIGS. 7 & 8. $ form inexpectata forma n. (Ivory Coast: Abidjan) (Gallay). N.B. The black area

at the base of the wings $ fig. 7 is a stain and should be ignored, upper and

undersides.

Approx. 2/3 nat. size.

Bull. Br. Mus. not. Hist. (Ent.) 23, 4

PLATE i

ENT. 23, 4

4ij hcj h^ hcj

o o Bo

I?

ft

sn

&! s> a

. , p

Bull. Br. Mus. not. Hist. (Ent.) 23, 4

PLATE 2

PLATE 3
Charaxes

FIGS. i8& 19. cedreatis Hewitson, $ form pseudosmaragdalis van Someren & Jackson, Type.
(Central African Museum, Tervuren, Belgium.) Upper and underside. All
light markings blue \vith the exception of large subapical spots fore wing and
submarginal linear marks in hind wing which are white.

FIG. 20. cedreatis Hewitson $ form dewitzi Butler. Photograph of original colour figure,

Type lost. Pattern generally similar to pseudosmaragdalis but lacking the
postdiscal row of blue spots in fore wing, upper and underside.

FIG. 21. chepalungu sp. n. Gynandromorph. (Kenya: Chepalungu Forest.) (National

Museum, Nairobi) (R. T. Evans leg.).

FIGS. 22 & 23. chepalungu, upper and underside. $ Type. (Kenya: Chepalungu Forest, lower
Sotik.)

FIGS. 24 & 25. chepalungu, upper and underside. <$ Type. (Kenya: Chepalungu Forest).

Approx. 2/3 nat. size.

Bull. BY. Mus. nat. Hist. (Ent.) 23, 4

PLATE 3

PLATE 4
Charaxes virilis Van Someren & Jackson

FIGS. 26 & 27. Type of virilis van Someren & Jackson (W. Africa), upper and underside.
Photos B.M. (N.H.) Nos. 34024; 34025.

FIGS. 28 & 29. $ (Uganda: Busia-Tororo Forest) (Bailey}. Ground colour very similar to
figs. 26 & 27 but with stronger blue sheen; fore wing marginal lunules more
denned. Hind wing with postdiscal row of greenish blue spots more distinct;
submarginal white spots larger; margin reddish ochre to lower tail. Borders
of wings on underside satiny with a strong rusty tinge to the postdiscal zone
of both wings ; the submarginal row of whitish marks strong, upper and under-
underside.

FIG. 30. c? (Uganda: Forests of Busiu-Tororo and Mabira Forest, Busoga), upper and

underside.

FIG. 31. (Moyen Congo: Kelle) (Jackson). Ground colour strong blue, marginal

glaucous marks diffuse. Blue spot beyond cell in fore wing distinct, but with
other spots very faintly indicated. Hind wing greenish, postdiscal line stronger.
Underside ground colour dark vinaceous brown, paler on borders and satiny
bars more distinct, upper and underside.

FIG. 32. (J (S. Nigeria: Ibadan), (Oxford). Ground colour blue-black with strong blue

sheen. Blue marks on fore and hind wings smaller, suffused purplish. Under
side rusty greyish brown, border on fore wing lighter. Pattern not strong,
upper and underside.

FIG. 33. $ (Cameroon: Bitje), [B.M. (N.H.)]. An aberrant specimen with a large greenish

blue triangular mark beyond end of cell in fore wing. Underside vinaceous
brown, pattern faint, but satiny bars present particularly in disc of hind wing,
upper and underside.

FIGS. 34 & 35. Type $ form virilis Rothschild. (Uganda: Jinja.) 34, upperside, purplish blue-
black. Fore wing with obscure blue subcostal marks beyond end of cell ; obscure
glaucous marks on margin. Hind wing with faint trace of greenish postdiscal
line; submarginal white spots distinct; some reddish on margin above upper
tail. 35, underside, dark vinaceous brown; faint satiny bars through discs
of wings ; pattern faint.

FIGS. 36 & 37. form lenis Jordan. $ Type. (Uganda: Budongo Forest.) 36, upperside as
in fig. 34. Fore wing with additional blue mark at upper part of end of cell.
No greenish postdiscal line on hind wing; marginal red to lower tail. 37,
underside : ground colour similar to fig. 35 but satiny bars and pattern stronger.

Approx. 3/4 nat. size.

Bull. BY. Mus. nat. Hist. (Ent.) 23, 4

PLATE 4

o o o o o

en en en en

R

111

S a ^

oo * ,

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00 '

a-
I

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Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 5

PLATE 6

Charaxes etheocles etheocles Cramer

FIGS. 47 & 48. cJ form carteri Butler, 1881, Type. (Ghana [Gold Coast], Accra), upper and

underside. Photos B.M. (N.H.) Nos. 41808; 41809.
FIGS. 49 & 50. $ form hollandi Butler, 1893, Type. (Sierra Leone), upper and underside.

Photos B.M. (N.H.) Nos. 41807; 41810.
FIGS. 51 & 52. form regalis Rothschild, Type. (Sierra Leone), upper and underside. Photos

B.M. (N.H.) Nos., 34026; 34027.
FIGS. 53 & 54. $ form fulgens Rothschild, Type. (Sierra Leone), upper and underside. Photos

B.M. (N.H.) Nos. 34032; 34023.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 6

\

brj

*T)

Tj hr]

*~3

*Tj

P

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P

Q

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CD

fore wing spots " rayed ", upper and underside.
$ form alladinis Butler. (Sierra Leone), upper and unders
$ form etheocles (Cramer), topotypical. (" West Coast ").
" rayed " than fig. 60. Hind wing band wider, white,
upper and underside.

white, slightly ochre tinged in postdiscal row. Hind win
inner border. Underside pattern moderately strong, uppe
$ form alladinis Butler var. (Cameroons), upper and under
$ form etheocles (Cramer), topotypical. (Sierra Leone),
hind wing band wider than in figs. 58 and 59. Undersi

-K5

8*

-t

3

k
*.

;s-

TO
O
TO

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I

underside. Photos B.M. (N.H.). Nos. 40800, 40801.

40

I
1
f

O

l-h
f

a

a 3.

1

8 1

T)

E

Charaxes etheocles etheocles (Cramer

PLATE 7

j*

P< en

"-I OQ

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] 1

H ^

5

^ >rj pj

p &

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5

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$ 1

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Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 7

hrj

hcj

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wings, upper and underside.

brownish grey with satiny silvery bars well marked, upper and underside.
$ form catochrous Staud. (Western Uganda: Bwamba Valley). Upperside ground c
similar to fig. 67. Fore wing without blue dots but marginal spots distinct. Underside:
two thirds both wings silvery grey with black markings strong, upper and underside.
<$ (Uganda : Masaka, Katera Forest) . Upperside ground colour blue-black ; fore wing subc
blue dots small ; marginal internervular glaucous dots present. Underside ground colour d;
brownish grey; satiny bars only slightly evident, upper and underside.
$ (Uganda: Jinja). Upperside ground colour blue-black; two minute subapical blue dots
marginal glaucous lunules strong. Underside pattern not strong on a brownish grey gro
satiny bars not strong, upper and underside.
$ (Uganda: Masaka, Katera Forest). Fore wing spots well spaced, creamy, slightly more och]
in postdiscal series. Hind wing band moderately wide but tapering, strongly bluish espe<
on distal border. Bred with $ form carpenteri in same family, upper and underside.
$ (Uganda: Toro area, Kibali Forest). Fore wing with large cell spot; discal spots large
white, that in ib diffuse and blue. Postdiscal spots complete but small, blue. Hind wing,
discal spots incomplete, obscured except for one white spot in 7; a bluish costal spot prese:
discal line. Underside darker greyish brown ground colour with a more decided pattern on

with small blue subcostal spots and indication of marginal dots. Underside ground c

borders. Submarginal linear marks not strong, underside boldly marked, upper and unde]
<$ form catochrous Staud. (Uganda: Budongo Forest). Upperside deep blue-black; fore

$ (Kenya: South Kavirondo, Suna). Ground colour blue-black. Fore wing discal and
discal spots slightly " rayed "; postdiscal row ochreous, discal row slightly paler in upper
but conjoined spots in la-ib bluish on proximal ends. Hind wing band wide, white with t

brownish, upper and underside.

distally opposite tail region. Marginal spots bold. Underside ground colour darker,

$ form pallidimacula van Someren & Jackson. Type. (Uganda: Jinja). Fore wing
white, widely separated, but spots in ib fused. Hind wing band broad, white, with blue b

Charaxes etheocles carpenteri Poulton

PLATE 8

o* p "5 u. a c jo cro^

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Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 8

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H
*

(Uganda: Masaka, Katera Forest, 1952)., Bred in same family as $
ng postdiscal spots complete, bluish white, slightly bluer in la-ib. Discal
e in ib. Hind wing: Postdiscal spots (blue) complete; a trace of bluish
ne. Underside: ground colour greyish brown with rusty flush over hind
ting moderate, upper and underside.
Rogers $ Type. (Kenya: East Elgon, 1932). Upper and underside.
. 42563, 42564.
an Someren & Jackson. (Kenya: Elgon, Kitale.) Fore wing spots
.ed by rufous rays; margin of wing rufous. Hind wing band white with
. Underside rusty brown with subdued pattern but bars well marked and
per and underside.
Rogers $. (Kenya: Elgon, Kitale). Upperside: ground colour deep
ith distinct subcostal blue spots, and obscured marginal glaucous lunules
Underside ground colour greyish drab-brown slightly darker on borders ;
and hind wing fairly strong, upper and underside.
Rogers $. (Kenya: Elgon, Kitale.) Fore wing spots orange-ochreous,
,ving band white with slight blue on borders. Underside ground colour
i borders of fore wing; satiny bars and pattern strong, upper and underside.

>r tornal black marks. Upperside and underside.
'Uganda: Jinja, i.i922). Fore wing blue marks in discal and postdiscal
Hind wing postdiscal zone with large continuous greenish blue marks.
Upper and underside.

(Tanganyika, S. E. Lake Victoria, Orange River, Serengeti Game
specimen is very similar to the type, which came from Kakindu Hill,
er, west side of Lake Victoria, Tanganyika. Fore wing postdiscal spots
[ spots limited to 2-3, and a minute dot at subcosta. Hind wing post-
xcept for bluish white marks in upper part. Underside greyish brown,

Charaxes etheocles (Cramer)

PLATE 9

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 9

ENT., 23 4

PLATE 10

Charaxes etheocles (Cramer)

FIGS. 82 & 86. biinclinata ssp. n. Holotype $. (Moyen Congo: Kelle, x. 1962) (Jackson)-
Fore wing spots bold, all creamy in colour. Hind wing band generally broad'
white with blue or inner border. Underside greyish, darker on borders ; pattern
bold, upper and underside.

FIGS. 83 & 87. biinclinata ssp. n. $. var. (b). (S.E. Nigeria: Ikom, iii. 1958) (Jackson). Very
similar to figs. 82 & 83 but fore spots conjoined by rays; margin with obscure
lunules. Hind wing broader. Underside ground colour as in figs. 82 & 83.
Fore wing spots rayed, upper and underside.

FIGS. 84 & 88. ochracea $ form violacea Rothschild (Gabon: Lambarene). Upperside: deep
blue-black: fore wing immaculate; hind wing with small sub-marginal spots
(white). Underside ground colour grey-brown, darker on postdiscal zone.
Pattern strong, upper and underside.

FIG. 85. biinclinata ssp. n. <j>. var. (a). (French Eq. Africa: Mambili, Ouesso, 1.1962)

(Jackson). Fore wing spots well separated, slightly ochreous, whiter in la-ib.
Hind wing band tapering, white, with blue borders. Underside dark greyish
brown; pattern strong, upper and underside.

FIG. 89. biinclinata ssp. n. $. var. (c). (Mambili, Ouesso, i.ig6o) (Jackson). Fore wing

spots all orange-ochreous except for large marks in la-ib whitish. Hind wing
band narrow, creamy, with purple shade on borders. Ground colour of both
wings above brownish black with purplish tone. Margin with obscure lunules.
Underside darker brownish, pattern bold. Cf. ochracea. Upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 10

PLATE ii

Charaxes etheodes (Cramer)

FIGS. 94 & 90. ochracea Roths. $ form seriata Roths. Type. (Gabon: Ogowe River), upper and

underside. Photos B.M. (N.H.) Nos. 34019; 34020.
FIGS. 91 & 95. ochracea Roths. $ Type (Gabon), upper and underside. Photos B.M. (N.H.)

Nos. 34017; 34018.
FIGS. 92 & 96. biinclinata form ephyra Gdt. #. (Moyen Congo: Etumbe, 1.1963) (Jackson).

Fore wing with subcostal blue spots; glaucous on margin. Hind wing with

distinct blue submarginal spots. Underside ground colour brownish grey,

fore wing pattern faint; satiny bars in hind wing strong. Form ephyra Gdt.,

upper and underside.
FIGS. 93 & 97. biinclinata form catochrous Staud. <?. (Moyen Congo : Kelle, x . 1962) (Jackson).

Fore wing above almost immaculate, hind wing submarginal spots obscured.

Underside: basal half of wings silvery up to discal zone; fore wing pattern

strong, upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE

PLATE 12

Charaxes grahamei sp. nov.
Upper and undersides.

FIGS. 98 & 99. (J Type. (Tanzania [Tanganyika] : Mkuyu Forest, Kigoma district) (Grahame,

1964).
FIGS. 100 & 101. $ Type. (Tanzania [Tanganyika]: Mukuyu Forest, Kigoma district)

(Grahame, 1965).
FIGS. 102 & 103. $ form lacteata f. nov. Type. (Tanzania [Tanganyika] : Mukuyu Forest,

Kigoma Area) (Grahame, 1965).

FIGS. 104 & 105. $ (Tanzania [Tanganyika]: Muhino Forest, Kigoma district) (Jap. Primate
Expdt. 1965).

Approx. 2/3 nat. size.

Bull. Br. Mus. not. Hist. (Ent.) 23, 4

PLATE 12

o o

o*

pu O

1 & ff i f 1 i

^<J^P^<^

Crt <J * K' H **. ^ *.

2. a- g p- L $ S
* <g * * * * f *

cr 40 Q -* ^-to

_, P HK 3 HhCT

s wj

o ET o o d ^
5* 3-' g. Z a I

lot t^ g

1 3 i- %12&

r/1 ^ <K S b fi

itrarius Weymer Type $
itrarius Weymer <J. (K<
upper surface ; marginal
cal band, upper and unde
itrarius $ form contrari
imba Hills, Kwale Distr

1

p

to

uT

a
p
<-*

en

8

en ^

a* <
C ts- B

H -* T

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J? 5

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P. M.?

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^
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P< ?l

g * I P

r+ CD

? 4 8.^.

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gl ? f

B' ^ ^
P- B' ?.

< ^
PS. o

us var. pseudorosae \
ict), upper and undei

2. o5 n

a^ P
f pi

ll 6

|rrw

1 g H

^ 3 *
013 g-S-
o o P

c? 1 ^^

PLATE 13

X^ J

2 p o

C7Q rt- 3

W QJ

a 1

P: B en"

ux !_Li CD
f-* en

p), y

H CD

M ( ^"

CT 1 <*

a c^

S- W

5 c o

a ^

o

PI - CD

Ui g K>

O*i O

g.ff

P

c ^ 5 1

' <D W

CD

a s i- 3

3 3

^ 5:^

CD .
*t) ' ^

^ en C/3

ty^ ti'

<4-' O a'

CD ^"^ P

M. cr

rr- a ^

M

Ft

< i

p. p g
J? 5* d

en ._, en

gal

CD en =

. CD

O

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s c^

d CD | I j

o

Co CO

en' ti t

cT

J4.'-,

' CD
P

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en
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p

^^i

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CD rh
i-l CD

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Q

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P'

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(/i y

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CD

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a

pu P

g cT

1*8' 3

a en

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2 <-*

d CD ff

PI en KJ
p CD en

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9

en

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hi o 4 '

CD &

P f ^

en

S e

Bull. Br. Mus. nat. Hist. )Ent.) 23, 4

PLATE 13

PLATE 14
Charaxes baileyi van Someren

FIGS. 115 & 118. $ Holotype. (Kenya: Visoi Gap, near Kilombe Hill, 3.11.1957) [In B.M.

(N.H.)], upper and underside. Photos B.M. (N.H.) Nos. 42566; 42568.
FIGS. 116 & 119. $ Allotype. (Kenya: Visoi Gap, near Kilombe Hill, 3.^.1967) [In B.M.

(N.H.)], upper and underside. Photos B.M. (N.H.) Nos. 42565; 42567.
FIGS. 117 & 1 20. form, pseudocarpenteri van Someren. (Kenya: Visoi Gap, Kilombe Hill,

western side of central rift), upper and underside.
Approx. 3/4 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 14

r

PLATE 15

Charaxes viola Butler

FIGS. 121 & 122. viola Butler $ Type [In B.M. (N.H.)], upper and underside. Photos B.M.

(N.H.) Nos. 38828; 38829.

FIGS. 123 & 124. viola Butler, typical (J (French W. Africa: Bamako), upper and underside.
FIGS. 125 & 126. viola $ form viola var. (W. Africa), upper and underside.
FIGS. 127 & 128. picta Rothschild <J Type. (Uganda: Unyoro). The underside is usually

plain. Type in B.M. (N.H.), upper and underside. Photos B.M. (N.H.).

Nos. 38825; 38830.
FIG. 129. picta Rothschild $. Very strongly marked. (Uganda: Mpanga, Fort

Portugal), upper and underside.

FIG. 130. picta $ f. picta Rothschild. (Uganda: Jinja Area, Busoga), upper and under-

side.
FIG. 131. picta $ form vansomereni Poulton. (Uganda: Jinja Area, Busoga), upper

and underside.
FIG. 132. picta $ form vansomereni Poulton var. (Uganda: Jinja Area), upper and

underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 15

PLATE 16

Charaxes viola Butler

FIG. 133. kirki Butler, form albifascia Poulton. Narrow hind wing band, upper and

underside.
FIG. 134. kirki Butler, $ form rogersi Poulton. (Kenya: Mile 29, Magadi Rd., Masai),

upper and underside.
FIG. 135. kirki Butler, $ form kirki Butler. (Kenya: Masai Reserve.) Upper and

underside.
FIG. 136. kirki Butler, <J typical specimen (Kenya: Mile 29, Magadi Rd., Masai Reserve.)

Upper and underside.
FIG. 137. kirki Butler, <$ well marked specimen. (Kenya: Mile 29, Magadi Rd., Masai

Reserve.) Upper and underside.
FIGS. 138 & 139. suk Carpenter & Jackson, typical <J. (Kacheliba, Suk). Upperside and

underside.

FIG. 140. suk Carpenter & Jackson, $ form intermedia Carpenter & Jackson. Upperside.

FIG. 141. suk Carpenter & Jackson, $ form kirkoides Carpenter & Jackson. Upperside.

FIG. 142. suk Carpenter & Jackson $ form achaemenesopsis Carpenter & Jackson.

(Kacheliba, Suk.). Upper and underside.
FIG. 143. suk Carpenter & Jackson, $ form albifascia Carpenter & Jackson. Wide

fore wing bar. (Kacheliba Suk). Upper and underside.
FIG. 144. suk Carpenter & Jackson, $ form albifascia Carpenter & Jackson. (Kacheliba,

Suk). Narrow fore wing bar. Upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Entom.) 23, 4

PLATE 16

PLATE 17

Charaxes viola Butler
Upper and undersides.

FIG. 145. loandae ssp. n. $ form violitincta forma n.

FIG. 146. loandae ssp. n. $ form loandae forma n. Allotype.

FIG. 147. loandae ssp. n. $ form instabilis forma n.

FIG. 148. loandae ssp. n. $ form protokirki forma n.

FIG. 149. loandae ssp. n. $ form vansonoides forma n.

FIG. 150. loandae ssp. n. < Holotype. (N. W. Angda: Dundo Area, Loanda).

FIG. 151. loandae ssp. n. $ form basiviridis forma n.

FIG. 152. loandae ssp. n. 9 f orm primitiva forma n.

FIGS. 153 & 154. brainei van Son. Type (S.W. Africa: Kombat, ^.1965). Photo.

Transvaal Museum.
FIGS. 155 & 156. brainei van Son. Type $ (S.W. Africa: Kombat, iv. 1965). Photo. Transvaal

Museum.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 17

ENT. 23, 4

16

ncj

oooooooo

R V> CO 01 (M 0>

pp

&>

O O\ Ln

O O 00

S3 3 3 3 3 3

c^ 5^ O ^*> C2 5i

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^" Cti

s 6 i

cS" '

to 8

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p p

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3 3

^

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s
ts

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a.

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3

Bull. Br. Mus. not. Hist. (Ent.) 23, 4

PLATE 18

9

DC

HI
CC

K)

o"'

N

3

|.

2*

3-

P

M

00

Iv

H

|

9*

'ts

p

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> oo

M
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t

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"N

IGS. 179 & 1 80. $ form rosella var
iinrlersiHp

O

<!

oo

S 3 i

c S. !
"nJ ^
^6 ^
?B g:

Oi -K

O

M
^1

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1?

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IGS. 173 & 174. $ form variata i<
Cottrell)], upper an

!

cr
*!

p"
15

I

3 0^
CL |

p CD

3

P
3

^
3 3

p
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3

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10

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3

upper and undersid*

[. Rhodesia) : Mwinili

J. Rhodesia): Mwini]

cr

5'

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3'
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a

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en

^P_"

o>

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cr

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p

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cr

p"

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Xi

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p

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cr

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Charaxes viola varial

PLATE ig

en

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g

g

J

-^

^

s

i'

CfQ

OQ

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f*

en

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f

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en

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p

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en

TT

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3

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3

cr
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3

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|

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pH

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p

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cr
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p

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1

3

|

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to

Cu

to

Bull. BY. Mus. nat Hist. (Ent.) 23, 4

PLATE 19

PLATE 20
Charaxes viola Butler
FIGS. 185 & 186. daria Rothschild. Type $, upper and underside. Photos B.M. (N.H.) Nos.

33653, 33654-
FIG. 187. daria Rothschild. Typical $ (Abyssinia: Jabalo, Gillet Mts.), upper and

underside.

FIGS. 188 & 192. viola handari Type $ (Tanzania [Tanganyika]), upper and underside.
FIGS. 189 & 190. chanleri Holland. Type $ (S. Abyssinia), upper and underside.
FIG. 191. chanleri Holland. $ (N.E. Mt. Kenya: Meru), upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 20

PLATE 21
Charaxes viola phaeus Hewitson

FIGS. 193 & 194. <J (Zomba Plateau), upper and underside.

FIGS. 195 & 196. $ (Malawi [Nyasaland] : Limbe), upper and underside.

FIG. 197. $ form phaeits Hewitson (W. Lake Victoria: Kitigati, Kagera River), upper

and underside.

FIG. 198. <J (W. Lake Victoria: Kitigati, Kagera River), upper and underside.

FIGS. 199 & 200. $ form vansoni van Someren & Jackson. (Bechuanaland : Mahalapya Drift)

(Penningtori) , upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 21

PLATE 22
Charaxes cynthia Butler

FIGS. 201 & 204. <J Type. Upper and underside. Photos B.M. (N.H.) Nos. 39937, 39938.

FIGS. 202 & 205. = guineensis Le Moult, <$ Type. Upper and underside.

FIGS. 203 & 206. Nominate $ (Nigeria: Ikom, vi.i957) (Jackson coll.). Upper and underside.

Approx 3/4 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 22

204

203

206

PLATE 23
Charaxes cynthia Butler
Upper and undersides.

FIGS. 207 & 208. mukuyu ssp. n. $ Type. (Mukuyu). Cf. with $ from Kafakumba, Katanga

[B.M. (N.H.)].

FIGS. 209 & 210. propinqua ssp. n. <J Type. (Uganda: Katera Forest, Masaka, xi.i953).
FIGS. 211 & 212. mukuyu ssp. n. $ (N.E. Lake Tanganyika: Forests of Mukuyu and Muhimo

in the Kigoma District, ^.1965.) (Grahame).
FIGS. 213 & 214. propinqua ssp. n. $ Type (Uganda: Katera Forest, Masaka, xi. 1953).

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 23

210

ENT. 23, 4

PLATE 24

Charaxes cynthia parvicaudatus Lathy
Upper and undersides.

FIGS. 215 & 216. <3 Type.

FIGS. 217 & 218. 9 Type.

FIGS. 219 & 220. <J (N.W. Kenya: Kakamega Forest, ix.1937) (M. Berkeley coll.).

FIGS. 221 & 222. $ (N.W. Kenya: Kakamega Forest, ix.i93y) (M. Berkeley coll.).

Approx. 2/3 nat. size.

Bull. Br. Mus. not. Hist. (Ent.) 23, 4

PLATE 24

PLATE 25

Charaxes cynthia sabulostts Talbot

FIGS. 223 & 224. Dark form $ (Katanga: Kinda) " wet " form. cf. with dark specimen from

Kafakumba, Katanga. Upper and underside.
FIGS. 225 & 226. Dark form $ aurantaca R.-D (Katanga: Kafakumba.) "wet" form.

[B.M. (N.H.)]. Upper and underside.
FIGS. 227 & 228. c form aurantiaca R.-D (Katanga: Kafakumba.) "dry" form. Upper and

underside.
FIGS. 229 & 230. (J "dry" form agreeing with Type. (Katanga: Kafakumba) (Overlaet).

Upper and underside.

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 25

226

PLATE 26

Charaxes cynthia kinduana Le Cerf
Upper and undersides.

FIGS. 231 & 234. cJType.

FIGS. 232 & 235. ? (Congo: Medje, x. 1910) (Carnegie Mus., Pittsburgh).

FIGS. 233 & 236. $ (Congo: Epulu, x. 1917) (Jackson).

Approx. 3/4 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 26

PLATE 27
Charaxes cynthia cameroonensis van Someren

FIGS. 237 & 238. $ Type, upper and underside.

FIG. 239. <J (Central African Republic: Bangui, 1.1961), upper and underside.

FIGS. 240 & 241. $ (Central African Republic: Bangui, i. 1961), upper and underside.

FIG. 242. <J (Cameroons : Meter, ii . 1921), upper and underside.

Approx. 3/4 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 27

239

PLATE 28
Charaxes

FIGS. 243 & 246. acuminatus obudoensis ssp. n. <J Type (E. Nigeria: Obudo Plateau.). Upper

and underside.

FIGS. 244, 245, xiphares kilimensis ssp. n. 244 and 247 <J Type upper and underside; 245 and
247, 248. 248 $ Type upper and underside, (both Tanzania [Tanganyika]: Lower
slopes of West Kilimanjaro, Maua Estate, ix.ig66 and ^.1964).

Approx. 2/3 nat. size.

Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 28

3

3

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3

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to

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to

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to

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to

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r>

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9,

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Bull. Br. Mus. nat. Hist. (Ent.) 23, 4

PLATE 29

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. MASNER, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. 5.

2. NIXON, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera:
Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6.

3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177;
18 plates, 270 Text-figures. August, 1965. 4 45.

4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172; 500 Text-figures. October,

1965- S5s.

5. AHMAD, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156;
475 Text-figures. November, 1965. 2 155.

6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae & Drosophilidae.
Pp. 129; 328 Text-figures. 3.

7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera : Coccoidea). Pp. 168; 43 Text-figures. February, 1967.

&3S.

8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera: Geometridae). Pp. 119; 14 plates, 146
Text-figures, 9 maps. February, 1967. 3 IDS.

9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera : Rhopalocera). Pp. 509. August, 1967. 8 los.

10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera : Rhopal-
ocera). Pp. 322; 233 Text-figures. Coloured frontispiece. September, 1967. 8.

11. MOUND, L. A. A review of R. S. BagnalTs Thysanoptera Collections. Pp. 184:
82 Text-figures. May, 1968. 4.

12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with an
Account of their World Distribution (Lepidoptera : Drepanidae). Pp. 151:
293 Text-figures, 14 plates. November, 1968. 5.

13. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families
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Text-figures. December, 1968. 5.

PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED, BARTHOLOMEW PRESS, DORKING

A REVISION OF THE AFRICANU' 5 "*"
SPECIES OF PSEUDORHYNCHUS^J^

SERVILLE
(ORTHOPTERA : TETTIGONIIDAE)

D. R. RAGGE

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 23 No. 5

LONDON: 1969

J^H^

A REVISION OF THE AFRICAN SPECIES OF

PSEUDORHYNCHUS SERVILLE \^

(ORTHOPTERA : TETTIGONIIDAE)

BY

DAVID ROBERT RAGGE

/O-'jf-

British Museum (Natural History)

Pp. 167-190 ; 38 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 23 No. 5

LONDON : 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, *s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

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

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

This paper is Vol. 23, No. 5 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) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued ii April 1969 Price Twelve Shillings

A REVISION OF THE AFRICAN SPECIES OF

PSEUDORHYNCHUS SERVILLE
(ORTHOPTERA : TETTIGONIIDAE)

By D. R. RAGGE

CONTENTS

Page
SYNOPSIS ............ 169

INTRODUCTION ........... 169

ACKNOWLEDGMENTS ........ .170

MATERIAL ........... 170

Pseudorhynchus Serville ......... 170

Key to the African species . . . . . . . .172

Descriptions of the African species . . . . . . 175

REFERENCES ........... 190

SYNOPSIS

The African species of Pseudorhynchus Serville are fully revised and an identification key is
given. Two new species and two new subspecies are described and one specific synonym, is
newly established.

INTRODUCTION

IN 1967 Professor D. F. Owen of Fourah Bay College (University of Sierra Leone)
sent me a collection of Orthoptera from Freetown for identification. Two of the
specimens belonged to an undescribed species of Pseudorhynchus Serville and, as
Professor Owen was studying colour polymorphism in this species, he asked me to
publish a name for it. Not wishing to describe a single species in isolation I under-
took the present study, which embraces all the African species of the genus.

The genus Pseudorhynchus was erected by Serville in 1838 for the three species
Ps. sicarius Serville (synonymized in the present paper with Ps. lanceolatus (Fabri-
cius)), Ps. flavescens (Serville) and Ps. lessonii Serville. No type-species was fixed
at that time, but the first of these three species was later designated as such by
Kirby (1906 : 237), who included 15 species (five African) in the genus. The latest
list of the species of Pseudorhynchus is that of Karny (1912 : 25), who included 14
species (five African) without qualification and a further five doubtful ones (one
African). The genus has not been fully revised since the work of Redtenbacher
(1891 : 364), though a key to the species was given by Karny (19070 : 17). The
present revision includes all the African species referred without qualification to
Pseudorhynchus by Karny (1912); five species (of which two are newly described)
and four subspecies (two newly described and one reduced in status from a species)
are recognized and one specific synonym newly established.

All the holotypes and lectotypes of the specific names covered by this revision
still exist and have been examined. The author's usual conventions are observed
(see Ragge, 1957 : 124) and the wing- vein nomenclature is that of Ragge (1955).

ENTOM. 23, 5 l8

170 D. R. RAGGE

The technique used in the examination of the stridulatory organ is described on
p. 172. The length of the fastigium of the vertex was measured from the basal
tooth to the tip.

ACKNOWLEDGMENTS

My sincere thanks are due to the following specialists, who have been kind enough
to send me type-specimens or other material from their respective institutions :

Dr. P. Basilewsky, Dr. M. Beier, Dr. L. Chopard, Dr. M. Descamps, Dr. J. de A.
Fernandes, Dr. Y. Gillon, Dr. K. K. Giinther, Mr. A. U. Oboite, Dr. E. C. G. Pinhey,
Dr. H. R. Roberts, Dr. C. H. Fraser Rowell, Dr. R. Roy, Dr. J. G. Rozen and the late
Mr. V. van Straelen.

I am also most grateful to the following workers, who have very kindly sent me
specimens collected by them personally :

Dr. and Mrs. R. W. Crosskey, Mr. R. A. Farrow, Miss J. M. McKay, Professor
D. F. Owen and Mr. J. Phipps.

My thanks are due to Mr. W. J. Bailey, who kindly drew my attention to the
replica technique for examining surface structure; this technique greatly facilitated
the study of the stridulatory rib of the left male fore wing (see p. 172).

I should also like to thank Miss L. M. MacDonell for her assistance in measuring
specimens and counting the hind tibial spines.

MATERIAL

In addition to the collection of Pseudorhynchus in the British Museum (Natural
History) (often abbreviated in this paper to BMNH), material was lent by the
institutions listed below, through the courtesy of the specialists mentioned above (the
abbreviations used where the material is listed in detail are inserted in parenthesis) .

Musee Royal de 1'Afrique Centrale, Tervuren (MRAC) ; Naturhistorisches Museum,
Vienna (NM) ; Institut de Recherche Scientifique de Madagascar, Tananarive (IRSM) ;
Museum National d'Histoire Naturelle, Paris (MNHN); Museu e Laboratorio
Zoologico e Antropologico, Lisbon (MLZA) ; Office de la Recherche Scientifique et
Technique Outre-Mer, Centre d'Adiopodoume, Abidjan, Ivory Coast (ORSTOM);
Zoologisches Museum of the Humboldt-Universitat, Berlin (ZMHU) ; University of
Ibadan, Nigeria (Univ. Ibadan) ; National Museum, Bulawayo, Rhodesia (NMR) ;
American Museum of Natural History, New York (AMNH); Makerere University
College, Kampala, Uganda (MUC); Institut Fondamental d'Afrique Noire, Dakar,
Senegal (IFAN); Institut des Pares Nationaux, Brussels (IPN).

PSEUDORHYNCHUS Serville

Pseudorhynchus Serville, [1838] : 509. Type-species: Pseudorhynchus sicarius Serville (=Ps.
lanceolatus (Fabricius)), by subsequent designation (Kirby, 1906 : 237).

Diagnosis. ^ $. Fastigium of vertex separated from fastigium of frons, bearing ventral
tooth or tubercle at base. Pronotum without lateral carinae. Prosternum with two well-
developed spines. Meso- and metasternum with paired lobes, not produced into spines. Fore
coxae with well-developed spine. Fore tibiae terete above, with slit-like tympanal opening on
each side. Hind femora with ventral spines.

REVISION OF AFRICAN PSE UDORH YNCH US 17:

Discussion. The known African members of this Copiphorine genus are ah 1 easily
recognized by their long, pointed " snout ", formed by the fastigium of the vertex.
There are shorter snouts in the two African genera Lanista Bolivar and Plastocorypha
Karsch, but in both these genera the fastigium of the vertex is contiguous with the
fastigium of the frons.

The African species of Pseudorhynchus fall into two quite distinct groups: Ps.
lanceolatus (Fabricius) and Ps. pungens (Schaum), in which the fastigium of the
vertex is sharply pointed and the male cerci have an internal spine at the base ; and
Ps. hastifer (Schaum), Ps. robustus sp. n. and Ps. crosskeyi sp. n., in which the
fastigium of the vertex is bluntly pointed and the male cerci have no basal spine.
Each of these two types of male cercus is common in the Copiphorinae, but it is
unusual for both to occur in the same genus. The two types of male cercus also
occur in the Oriental species of Pseudorhynchus, but the correlation between this
feature and the sharpness of the fastigium of the vertex does not hold for these
species : the fastigium is short and blunt in some Oriental species in which the male
cerci have a basal spine. A thorough study of all the species of Pseudorhynchus
might well show that some of them would be better regarded as a distinct genus, but
it is unlikely that such a decision would affect any of the species covered by the
present revision.

One of the most interesting features of this genus in Africa is the colour variation,
both within and between species. Like many Copiphorinae, ah 1 the species occur
in two principal colour forms, one mainly green and the other mainly brown. (The
three known specimens of Ps. crosskeyi sp. n. are all green, but it may be safely
assumed that a brown form exists.) In addition to this general body colour there is
often a pattern of dark pigmentation on the head, particularly on the frons and the
underside of the fastigium of the vertex; leaving aside Ps. crosskeyi sp. n., of which
insufficient material is available, the development of dark pigmentation takes a
different form in each species, as outlined below.

Ps. lanceolatus (Fabricius). The dark pigmentation always forms the same
pattern and is independent of the green/brown polymorphism. There is little
geographical variation.

Ps. pungens (Schaum). The dark pigmentation is independent of the green/
brown polymorphism, but shows marked geographical variation, forming quite
different patterns in different parts of Africa.

Ps. robustus sp. n. The dark pigmentation always forms the same pattern but
is shown only by the brown colour variety. There is little geographical variation.

Ps. hastifer (Schaum). None of the specimens examined of either the green or
the brown colour forms had any dark pigmentation on the underside of the head.

Since the genitalia of Ps. hastifer (Schaum), Ps. robustus sp. n. and Ps. crosskeyi
sp. n. do not show clear-cut differences, it was thought worth making a closer exami-
nation of the stridulatory organ and, in particular, the shape of the stridulatory rib
of the left male fore wing and the number of teeth on it. To avoid the difficulties

ENTOM. 23, 5 l8

172 D. R. RAGGE

of examination with reflected light or the labour of making cleared preparations of
the wing (both of which have severely limited the use of the microscopical structure
of the stridulatory organ in Ensiferan taxonomy), a replica technique was used; this
enabled the structure of the stridulatory rib to be examined without inconvenience
and without removal of the fore wing from the specimen. The technique adopted
is described below.

The left fore wing of each male specimen was relaxed locally by applying a small quantity of
10% ammonia solution to the wing-articulation with a hypodermic syringe or fine brush; any
legs that would have obstructed the extension of the fore wing were relaxed at the base in the
same way. As soon as it was relaxed (after no more than a few minutes) the fore wing was
extended and, if necessary, the underside was allowed to dry. A small quantity of a strong
solution of " Pyroxylin " (low viscosity nitrocellulose) in acetone was then applied to the under-
side of the cubito-anal area with a fine brush. After about two minutes the Pyroxylin replica,
now dry, was peeled off the wing surface and placed between two microscope slides. If the
replica was satisfactory the fore wing was returned to its flexed position on the insect, which thus
resumed its previous appearance. This technique, which is essentially similar to the replica
techniques described by several other authors (e.g. Sampson, 1961), has the advantages of being
quick, leaving the specimen quite unharmed and being repeatable on the same specimen at any
time. The resulting replicas are transparent and show the finest details of the surface structure
of the wing; they can be easily examined using transmitted light and a number can be placed
between each pair of slides for permanent storage. The self-adhesive binding strips used in
making 7x7 cm. permanent mounts of photographic transparencies were found to provide a
convenient way of binding the slides together.

Replicas of the stridulatory rib were taken in this way from all the male specimens
available of all five species. They were used in preparing the diagrams of the
stridulatory rib, in determining its length and in counting the number of stridulatory
teeth ; a number of tests failed to show any significant difference between the measure-
ments taken from the replica and those taken from the actual wing. The small and
sometimes rather irregular teeth that occur towards the end of the stridulatory rib
in some of the specimens were included both in measuring the length of the rib and
in counting the teeth. The results are given separately for each species and will be
seen to show some well marked interspecific differences in spite of considerable
intraspecific variation. The curvature of the stridulatory rib of Ps. pungens (Schaum)
(Text-fig. 16) is characteristic, and the large number of stridulatory teeth in Ps.
crosskeyi sp. n. marks it off well from Ps. robustus sp. n., which it resembles in many
other respects.

Distribution. Pseudorhynchus is widespread in the Old World tropics, including
most of the Oriental Region, Japan and Australia. Its range in Africa extends from
near the northern limit of the grassland (at about latitude 15 N) to the eastern part
of Cape Province in the south.

KEY TO THE AFRICAN SPECIES
MALES

i Cerci as in Text-figs, i or 2, with a basal, upwardly directed spine. Fastigium of the

vertex sharply pointed (Text-figs 24-28) . ...... 2

- Cerci as in Text-figs. 3, 4 or 5, without a basal spine. Fastigium of the vertex bluntly

pointed (Text-figs. 29-32) . * 3

REVISION OF AFRICAN PSEUDORHYNCHUS 173

2 Cerci as in Text -fig. i, with a short basal spine. Fore wings more than 42 mm. long

Ps. lanceolatus (Fabricius) (p. 175)
Cerci as in Text-fig. 2, with a long basal spine. Fore wings less than 41 mm. long

Ps. pungens (Schaum) (p. 179)

3 Fore wings with a pointed tip, as in Text-fig. 8. Stridulatory region of the left fore

wing as in Text-fig, n, lA as prominent as C 2 . Ps. hastifer (Schaum) (p. 186)

I

FIGS. 1-5. Posterodorsal view of the cerci and terminal abdominal tergites of the male of
(i) Pseudorhynchus lanceolatus (Fabricius); (2) Ps. pungens (Schaum); (3) Ps. hastifer
(Schaum) ; (4) Ps. robustus sp. n. ; (5) Ps. crosskeyi sp. n.

FIGS. 6-io. The right male fore wing of (6) Pseudorhynchus lanceolatus (Fabricius);
(7) Ps. pungens (Schaum); (8) Ps. hastifer (Schaum); (9) Ps. robustus sp. n. ; (10) Ps.
crosskeyi sp. n.

174

D. R. RAGGE

Fore wings with a rounded tip, not as in Text-fig. 8. Stridulatory region of the left

fore wing as in Text-figs. 12 or 13, lA much less prominent than Cw 2 . . 4

Stridulatory region of the left fore wing as in Text-fig. 12; lA well separated from
Cw 2 ; cubito-anal area broadened. Stridulatory rib as in Text-fig. 18, with fewer
than 80 Stridulatory teeth. Hind tibiae with fewer than 15 external dorsal spines

Ps. robustus sp. n. (p. 187)

Stridulatory region of the left fore wing as in Text-fig. 13; lA contiguous for part of
its length with Cu z ; cubito-anal area not broadened. Stridulatory rib as in Text-
fig. 19, with more than 80 Stridulatory teeth. Hind tibiae with more than 15
external dorsal spines Ps. crosskeyi sp. n. (p. 188)

14

12

FIGS. 11-14. The Stridulatory organ of Pseudorhynchus Serville. 11-13. Dorsal view of
the Stridulatory organ of (n) Ps. hastifer (Schaum); (12) Ps. robustus sp. n.; (13) Ps.
crosskeyi sp. n. 14. The venation of the Stridulatory area of the left male fore wing of
Ps. robustus sp. n.

FEMALES

Ps. crosskeyi sp. n., in which the female sex is unknown, is omitted from this key.
[ Ovipositor less than 16 mm. long, shaped as in Text-fig. 21 . Ps. pungens (Schaum) (p. 179)

- Ovipositor more than 18 mm. long, shaped as in Text-figs. 20, 22 or 23 . . . 2
z Fore wings with a pointed tip, as in Text-fig. 8 . . Ps. hastifer (Schaum) (p. 186)

- Fore wings with a rounded tip, not as in Text-fig. 8 ...... 3

3 Dark pigmentation of the underside of the head as in Text-fig. 24; fastigium of the

vertex sharply pointed. Ovipositor relatively broad, as in Text-fig. 20. Hind
tibiae usually with fewer than 7 external dorsal spines

Ps. lanceolatus (Fabricius) (p. 175)

- Dark pigmentation of the underside of the head as in Text-fig. 31 or absent; fastigium

of the vertex bluntly pointed. Ovipositor relatively slender, as in Text-fig. 23.
Hind tibiae usually with more than 8 external dorsal spines Ps. robustus sp. n. (p. 187)

REVISION OF AFRICAN PSE UDORH YNCH US

175

DESCRIPTIONS OF THE AFRICAN SPECIES

Pseudorhynchus lanceolatus (Fabricius)
(Text-figs, i, 6, 15, 20, 24, 36)

Locusta lanceolatus Fabricius, 1775 : 284. Holotype $, SIERRA LEONE (BMNH) (see below)

[examined].
Pseudorhynchus sicarius Serville, 1838 : 510. Lectotype <j>, locality unknown (MNHN, Paris)

[examined], syn. n.
Pyrgocorypha hastata Bolivar, 1890 : 222. Lectotype <$, locality unknown (MLZA, Lisbon)

[examined].

Diagnosis. <$ $. Dark pigmentation of underside of head distributed as in Text-fig. 24 ;
fastigium of vertex sharply pointed at tip. Venation and shape of right male fore wing as in
Text-fig. 6; stridulatory rib of left male fore wing shaped as in Text-fig. 15, with about 76-100
teeth (mean of 24 examined : 89) ; female fore wings similar except for cubito-anal area. Male
cerci as in Text-fig, i, with basal, upwardly directed spine. Ovipositor rather broad, shaped as
in Text-fig. 20.

MEASUREMENTS

Males

63 2-81 o, mean 72 oo
6 '9-9-0, mean 8-18
9-2-11-8, mean 10-47
18-0-23-6, mean 20 -64
43 1-55 o, mean 49 26
3-10-3-78, mean 3-45

(42):
(46);

(52):
(50):
(50):

Females

72-5-89-4, mean 79-12
7-5-10-8, mean 8-74
10-0-12-1, mean 11-02
20-1-25-7, mean 22-33
49-3-62-1, mean 54-13

(48): 20-6-27-6, mean 23-64

Total length (36) :

Length of fastigium of vertex (42) :

Median length of pronotum (45) :

Length of hind femur (44) :

Length of fore wing (41) :

Length of stridulatory rib (24) :
Length of ovipositor

Discussion. This species, which is a common insect in West and Central Africa,
may be easily recognized by the arrangement of the dark markings on the underside
of the head together with the large size. The only other African species of Pseudo-
rhynchus in which the head is sometimes similarly marked is Ps. pungens (Schaum),
which is much smaller and has quite different male genitalia and ovipositor.

The brown colour variety has four dark stripes along the top of the head and
pronotum, converging anteriorly and merging into two stripes at the level of the
antennal sockets; there are also one or two dark stripes on each side of the head,
extending posteriorly from the eyes and continuing along the lateral lobes of the
pronotum. The fore wings have less clearly defined dark markings, also tending to
form longitudinal stripes. Longitudinal striping of a similar nature is shown by the
green variety, at least on the head and pronotum, but it is impossible to judge from
dried material the extent to which the stripes are developed in this variety of the
live insect.

The holotype of this species is in the Sir Joseph Banks Collection in the British
Museum (Natural History). As stated by Zimsen (1964 : 618) there are two speci-
mens in this collection above the label " Locusta lanceolata " : one is a female (labelled
on the pin " Sierra Leone ") and is clearly Fabricius's type, and the other, a male
(labelled on the pin " Siam "), belongs to the Oriental species Ps. nobilis (Walker).
It is clear from Fabricius's statements "... fronte porrecta, lanceolata . . . ",
"... frons inter oculos prominet, capite longior ..." and " Habitat in Sierra
Leon Africae ", all ot which are applicable only to the female, that it was on this
specimen alone that the name and description of " Locusta lanceolata " were based

I 7 6

D. R. RAGGE

Ps. sicarius Serville was described on the basis of two female specimens from
unknown localities, one in the collection of " M. le compte Dejean " and the other
in the Museum National d'Histoire Naturelle, Paris. The former specimen is now
lost, but there is no doubt from Serville's description that it belonged to Ps. pungens
(Schaum). The specimen in the Paris museum, for which Serville gave a brief
separate description, still exists and I have been able to examine it : it clearly belongs
to Ps. lanceolatus (Fabricius). In order that the name Ps. sicarius should be based
on an existing and accessible specimen, I here designate the specimen in the Paris
museum as the LECTOTYPE of this species and I have so labelled it.

'"""//////I

|5

|6

|7 HAST

18 ROB

|9 CROS

FIGS. 15-19. Diagrams showing the shape of, and arrangement of teeth in, the stridulatory
rib of (15) Pseudorhynchus lanceolatus (Fabricius); (16) Ps. pungens (Schaum); (17) Ps.
hastifer (Schaum); (18) Ps. robustus sp. n. ; (19) Ps. crosskeyi sp. n.

2O LANC

21

23

(21)

22 HAST

of the ovipositor of (20) Pseudorhynchus lanceolatus (Fabricius) ;
t) ; (22) Ps. hastifer (Schaum) ; (23) Ps. robustus sp. n.

FIGS. 20-23. Lateral view of the ovipositor of (20) Pseudorhy
i) Ps. pungens (Schaum) ; (22) Ps. hastifer (Schaum) ; (23)

REVISION OF AFRICAN PSE UDORH Y NCH US

177

Pyrgocorypha hastata Bolivar was described from a single male and an unknown
number of females in the Museu e Laboratorio Zoologico e Antropologico, Lisbon;
the type-locality is unknown. There are in that museum two males and three
females (all without data) labelled, apparently in Bolivar's handwriting, as belonging
to this species. The label bears an amendment, apparently in the same handwriting:
" i <$ and 2 $<?" has been altered to " 2 <$<$ and 3 $?" (information kindly supplied
by Dr. J. de A. Fernandes). It would thus seem likely at first sight that a second
specimen of each sex came to the attention of Bolivar after he had written his account
of the species. However, Bolivar states in his description that the unique male

FIGS. 24-32. Ventral view of the head of (24) Pseudorhynchus lanceolatus (Fabricius); (25)
Ps. pungens pungens (Schaum); (26) Ps. p. striatus ssp. n.; (27) Ps. p. werneri Karny;
(28) Ps. p. meridionalis ssp. n. ; (29) Ps. hastifer (Schaum); (30) Ps. robustus sp. n. (green
form); (31) Ps. robustus sp. n. (brown form); (32) Ps. crossheyi sp. n.

178 D. R. RAGGE

lacked the terminal segments of its abdomen (" El unico <$ que he visto carece de los
anillos terminales del abdomen . . . "), and in both the existing males the abdomen
is intact. Although it is possible that Bolivar was mistaken I have thought it
unwise to regard either male specimen as a syntype, but it seems highly probable
that at least two of the females are syntypes and I have selected and labeUed one of
them as a LECTOTYPE.

Material examined. Holotype. SIERRA LEONE, 9 (BMNH).

GUINEA: Nimba, 1000 m, i $, ii.vi.ig42 (Lamotte) (IFAN, Dakar); Nimba,
1640 m, i $, xii.i95i (Lamotte & Roy) (IFAN, Dakar); Nimba, Ziela, I <$, 30-31.
111.1957 (Lamotte, Amiet 6- V anderplaetseri) (IFAN, Dakar); Nimba, Yalanzou, i ^,
I5.iii.i942 (Lamotte) (MNHN, Paris); Nimba, Nion, i $, ii-iv.i942 (Lamotte)
(MNHN, Paris); Nimba, Keoulenta, 3 $, i 9, ii-iv.i942 (Lamotte) (MNHN, Paris);
Conakry, i ^ (Bertrand) (MNHN, Paris); Seredou, i <$, xii.igsg (Pujol) (MNHN,
Paris); Toulaya, i $, 23.xii.i95i (MNHN, Paris); SIERRA LEONE: Njala, i $,
xii.i93i (Hargr eaves); Freetown, 4^, 3$, xii.i966 (Owen); IVORY COAST: Mt.
Tonkoui, i <, 15.^.1964 (Gittori) (ORSTOM, Abidjan); TOGO: Bismarckburg,
2$, i-iv.iSgi (Buttner) (ZMHU, Berlin); Misahohe, i 9, 12. v. 1894 (Baumanri)
(ZMHU, Berlin); NIGERIA: Ibadan, 4 <, 5 $, xii-v. 1960-67 (Oboite) (Univ. Ibadan);
near Ibadan, Gombar, at light, i $, 11.1.1965 (Gerard); CAMEROUN: Batouri Distr.,
savanna, 2$, i-ii.1936 (Mer field}; Batouri Distr., 3 45' N, 13 45' E, 750 m, i 9,
v-vi.i935 (Merfield); Abong M'Bang Distr., i 9, ^.1936 (Mer field}; CONGO (BRAZZA-
VILLE): Mayomba, i $, 1899 (Vergues) (MNHN, Paris); Djoue distr., between
Brazzaville and Reneville, i <$, 1912 (Bel) (MNHN, Paris) ; i 9, 1909 (Lebaudy)
(MNHN, Paris) ; M'Bamou, near Brazzaville, 2 $, i 9, 1903 (Montezer} (MNHN,
Paris); CENTRAL AFRICAN REPUBLIC: Boukoko, i $, 1952 (MNHN, Paris); CONGO
(KINSHASA): Kasai, Lomami, i <$, 1928 (Mayne); Uele, Gabu, i <, I4.xii.i932
(Vrydagh); Uele, Ekibondo, between Niangara and Dungu, 2650 ft., 5 ^, 3 $, i.vii.
1935 (Vanderbilt Exp.) (AMNH, New York) ; Luluabourg, i <$, 1936 (Puissant) ;
Mayidi, i <$, 1942 (Van Eyen) ; Bas-Congo, 2 $, 1927 (Sjoren) ; Camp de Lukula, i <$,
1911 (Daniel); Ituri, Bunia, i <$, iii.1939 (Maristes); Ituri, Faradje, i $, 22.1.1930
(Collarf); Elisabetha, i $, 1921 (Tinant}; Ubangi distr., i <$, ix.igio (Schubotz)
(AMNH, New York); Garamba National Park, 7 ^, 9 $, iii-xii. 1950-51 (De Saeger
& Verschuren} (IPN, Brussels, except for 2 <$, 2 $ in BMNH) ; Upemba National
Park, Lusinga, 1760 m, i $, i8.vii-8.viii.i947 (de Witte); Elizabeth ville, in open
high grass country, i$ (AMNH, New York); BURUNDI: Muyeha, R. Kishubi,
1700 m, i$, I3.vii.i952 (Laurent); UGANDA: Entebbe, i ^ (Gowdey); Entebbe,
i $, 27.viii.i9ii (Gowdey); Entebbe, i <$, 1^1.1913 (Gowdey}; Mwera, i <$, 2.viii.
1913 (Gowdey); Mengo, i$ (Millar); Kampala, i $, 13 ix.igiS; Kampala, i $,
11.1934 (Johnston); Kampala, i <$, 1908 (Bayon); Kampala, i $, 7.vi.i942 (Rehn)
(AMNH, New York); Budongo, i$ (Gowdey); Kabule, i $, 19. vi. 1909; Toro,
Nyakasura, i $, 18. in. 1932 (Shillito}; Kakindu, 2^, i 9, 1910 (Bayon}; Bussu,
I $, 1909 (Bayon) ; Busu-Hill, Busoga, i <?, i $ (Carl) (ZMHU, Berlin) ; Jinja, 2 9,
iii-iv.i932 (van Someren}; Tororo distr., Sukulu, i 9, 13.1.1961 (Burtt); ETHIOPIA:
i 9, 1899 (Michel} (MNHN, Paris); TANZANIA: Mahari Peninsula, Kasoge, 2550 ft,

REVISION OF AFRICAN PSE UDORH YNC H US 179

i $, viii-ix.i959 (2nd Oxford Univ. Exp.); ANGOLA: Lunda, Dundo, 2 ^, 2 $, ii-vii.
1949 (Machado).

In the British Museum (Natural History) unless otherwise stated.

Distribution. This species occurs commonly over a large part of West and
Central Africa, extending from Guinea to Ethiopia and southwards into northern
Angola. It is typically a species of high-grass savanna.

I
i

Pseudorhynchus pungens (Schaum)
(Text-figs. 2, 7, 16, 21, 25-28, 34, 35)

Conocephalus pungens Schaum, 1853 : 778. Holotype $, MOZAMBIQUE (ZMHU, Berlin)

[examined].

Diagnosis. <$ ?. Dark pigmentation of underside of head variable, but usually distributed as
in Text-figs. 25-28; fastigium of vertex sharply pointed at tip. Venation and shape of right male
fore wing as in Text-fig. 7; stridulatory rib of left male fore wing shaped as in Text-fig. 16, with
about 50-87 teeth (mean of 16 examined: 66); female fore wings similar except for cubito-anal
area. Male cerci as in Text-fig. 2, with basal, upwardly directed spine. Ovipositor relatively short,
shaped as in Text-fig. 21.

MEASUREMENTS

Males Females

Total length (28): 41 -6-62-8, mean 53-02 (45): 50-4-71 -9, mean 61 -76

Length of fastigium of vertex (29): 5-5-9-4, mean 7-65 (54): 6-0-10-9, mean 8-95

Median length of pronotum (29) : 6 2-8 3, mean 7-31 (56) : 6 8-9 7, mean 8 36

Length of hind femur ( 2 9): 12-0-17-9, mean 15 -14 (51): 13-3-20-7, mean 17-44

Length of fore wing (30) : 25 8-40 2, mean 34 30 (52) : 29 6-46 8, mean 39 82

Length of stridulatory rib (24) : i -58-2-47, mean i -95

Length of ovipositor (49) : 9-1-14-2, mean 1 1 -49

Discussion. The small size of this species, together with the characteristic male
cerci and short ovipositor, enable it to be readily distinguished from the other
African species of Pseudorhynchus.

In addition to showing the usual green and brown colour varieties, which doubtless
both occur throughout its range, Ps. pungens (Schaum) has the dark pigmentation
on the underside of the head and the side of the mesothorax distributed in four
distinct ways. As far as is known at present these colour forms, which appear to
be independent of the green/brown polymorphism, are allopatric (see Text-fig. 33)
and I have considered it convenient to treat them as subspecies. There are some
associated slight differences in size and abdominal terminalia, and when more
material becomes available one or more of these forms may prove to be distinct
species, but the evidence available at present agrees well with the treatment given
to these forms here. When considered as a whole they form a well defined unit
clearly separated from the remaining species of the genus.

Distribution (Text-fig. 33). Ps. pungens (Schaum) probably occurs throughout
the savannas and grasslands of tropical Africa and extends southwards, in the form
of Ps. p. meridionalis ssp. n., into the coastal savanna of eastern South Africa.

i8o

D. R. RAGGE

KEY TO THE SUBSPECIES

1 Dark pigmentation of the underside of the head as in Text-figs 27 or 28, the fastigium

of the vertex darkening towards the tip ........ 2

- Dark pigmentation of the underside of the head as in Text-figs 25 or 26, the fastigium

of the vertex with a dark ventra 1 stripe . . . . ... . 3

2 Mesothorax with a conspicuous dark spot on each epimeron (Text-fig. 38). (South

Africa) ...... Ps. pungens meridionalis ssp. n. (p. 185)

Ps. pungens pungens
A Ps. p. striatus

Ps. p. werneri

Ps. p. meridionalis

FIG. 33. Map showing the known distribution of the subspecies of Pseudorhynchus pungens
(Schaum) .

REVISION OF AFRICAN PSE UDORH YNCH US

181

Mesothorax without a dark spot on each epimeron. (West and Central Africa, north

of the equator) ...... Ps. pungens werneri Karny (p. 184)

Dark pigmentation of the frons as in Text-fig. 26, forming two longitudinal stripes,
sometimes interrupted or restricted to the anterior end of the frons

Ps. pungens striatus ssp. n. (p. 182)

Dark pigmentation of the frons as in Text-fig. 25, forming two conspicuous dark spots

near the mandibular articulations . . Ps. pungens pungens (Schaum) (p. 181)

Pseudorhynchus pungens pungens (Schaum)

(Text-fig. 25)
Conocephalus pungens Schaum, 1853 : 778.

Diagnosis. <$ $. Dark pigmentation of underside of head as in Text-fig. 25 ; fastigium of
vertex with dark ventral stripe ; frons without longitudinal stripes but with dark spot on fastigium
and two dark spots near anterior mandibular articulations. Epimera of mesothorax often with
rather inconspicuous dark spot (position as in Text-fig. 37). Stridulatory rib of left male fore
wing with about 61-76 teeth (mean of 8 examined: 69).

MEASUREMENTS

Total length (10)

Length of fastigium of vertex (10)

Median length of pronotum (10)

Length of hind femur (9) :

Length of fore wing (10)

Length of stridulatory rib (8) :
Length of ovipositor

Males

52-9-62-8, mean 58-16
7-5-9-3, mean 8-57
7-1-8-3, mean 7-79

14-7-17-5, mean 16-28
34-7-40-2, mean 37-86

i 97-2 24, mean 2 08

Females

(18): 59-6-71 -9, mean 66-62
(20): 8-3-10-9, mean 9-48
(21): 8-0-9-7, mean 8-87
(18): 16-2-20-7, mean 18-62
(20) : 39 5~4 6 ' 8 - mean 43 24

(17): ii -3-14-2, mean 12-74

Discussion. This subspecies may be recognized by the colour pattern on the
underside of the head. All the males in the material examined had a dark spot of
varying size and intensity on each mesothoracic epimeron (position shown in Text-
fig. 37), but this feature was completely lacking in many of the females. As no two
specimens of opposite sex had been c