/$**

Bulletin of the ( ^

V> tte*A*y

British Museum (Natural Histor^

Entomology series Vol 43 1981

British Museum (Natural History)
London 1982

Dates of publication of the parts

No 1 30 July 1981

No 2 24 September 1981

No 3 29 October 1981

No 4 26 November 1981

ISSN 0524-6431

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset

Contents
Entomology Volume 43

Page

No 1 A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea)
and its allies.
N. D. Jago 1

No 2 The Asian, Australasian and Pacific Paraboloponinae (Homoptera :
Cicadellidae). A taxonomic revision with a key to all the known genera
of the subfamily.
M. D. Webb 39

No 3 A revision of Phyciodes Hiibner and related genera, with a review of
the classification of the Melitaeinae (Lepidoptera : Nymphalidae).
L. G. Higgins 77

No 4 A revision of six minor genera of Myrmicinae (Hymenoptera : Formi-
cidae) in the Ethiopian zoogeographical region.
Barry Bolton 245

GENER

-4AUG

Bulletin of the

British Museum (Natural History

A revision of the genus Usambilla Sjoste(
(Orthoptera: Acridoidea) and its allies

N. D. Jago

Entomology series

Vol 43 No 1 30 July 19:

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued
in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy)
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.)

Trustees of the British Museum (Natural History), 1981

ISSN 0524-643 1 Entomology series

Vol 43 No 1 pp 1-38
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 30 July 1981

A revision of the genus Usambilla Sjostedt
(Orthoptera : Acridoidea) and its allies

N. D. Jago

<f^f

Centre for Overseas Pest Research, College House, Wrights Lane, London W8 5SJ

Contents

Synopsis

Introduction ....

Material

Key to genera in the Lentulidae .
Chromousambilla gen. n.

Key to species ....

Males

Females ....
Microusambilla gen. n. .
Altiusambilla gen. n.
Rhainopomma gen. n. .

Key to species . . . .

Males

Females . . . .
U sambilla Sjostedt

Key to species and subspecies .

Males

Females ....

References

Index .

1

1

2

3

5

6

6

6

10

12

13

14

14

15

20

22

22

24

37

38

GENER

4AUG

LIBRAI

Synopsis

The genus Usambilla Sjostedt is redefined and fully revised with keys to the species and subspecies. Four
new genera, ten new species and three new subspecies are described. Altiusambilla modicicrus (Karsch) is
implicated in the defoliation of exotic Pinus plantations in northern Tanzania.

Introduction

The genus Usambilla has been assumed to consist of a homogeneous group of small lentulid
grasshoppers. The male genitalia of the group reveal, however, that five genera are involved
within the old definition of Usambilla. Chromousambilla has male penis valves of great length
(Fig. 12) which enable the diminutive male to remain firmly attached to the female during
copulation without standing upon her in the usual acridid manner. The penis valves are curved
and fit into the female spermathecal duct, which is of similar shape and length (Fig. 23). Usam-
billa sensu stricto has tapered valvulae (Fig. 150) with barbs on each side short of the apex, while
Rhainopomma (Fig. 72) has these barbs situated apically. Microusambilla clearly has affinity with
Usambilla, while Altiusambilla has affinity with Rhainopomma and Mecostibus.

Reports have recently been published (Whellan, 1975; 1976) of eumasticids and lentulids
causing defoliation of exotic Pinus, particularly P. patula. In Malawi defoliation has been caused
by Plagiotriptus Karsch (Eumastacidae) and members of the lentulid genera Mecostibus, Nyass-
acris and Malawia. In Tanzania attacks so far reported are from W. Kilimanjaro in December,
1975 at Msituni, the species involved being members of the genus Chromothericles Descamps
(Eumastacidae) and the lentulid Altiusambilla modicicrus. This last is morphologically much

Bull. Br. Mus. not. Hist. (Ent.) 43(1): 1-38

Issued 30 July 1981

N. D. JAGO

closer to Mecostibus than to the rest of Usambilla sensu stricto. Indeed the close similarity
between the genera described in this paper may be due to convergent evolution. The external
similarity between Altiusambilla modicicrus and Rhainopomma montanum is particularly striking.

Material

Most of the material used for this study is in the collections of the first two institutions listed
below. Museums loaning specimens have their abbreviations listed in the text as follows.

BMNH

COPR, London
NR, Stockholm
MNHU, Berlin
MR AC, Tervuren

British Museum (Natural History), London

Centre for Overseas Pest Research, London

Naturhistoriska Riksmuseet, Stockholm

Museum fur Naturkunde der Humboldt-Universitat, Berlin

Musee Royal de 1'Afrique Centrale, Tervuren

Kenya series were immensely improved by the addition of COPR material collected recently
by Dr I. A. D. Robertson and Mrs A. Robertson.

Figs 1-9 Chromousambilla species. 1 , lateral aspect of pronotum of male C. veseyi. 2, dorsal aspect of
head of male C. latestriata. 3, apex of male abdomen of C. burtti, lateral aspect. 4, entire male of C.
latestriata. 5-9, dorsal aspect of male abdominal tip in (5) C. burtti; (6) C. mweruensis; (7) C.
robertsoni; (8) C. latestriata; (9) C. veseyi. Scale lines represent 1 mm, that below Fig. 1 applies to Figs
1-4, that above Fig. 9 applies to Figs 5-9.

REVISION OF USAMBILLA 3

Key to genera in the Lentulidae (modified after Dirsh, 1965)

1 External apical spines of hind tibia absent. Head prognathous, or face vertical, or moderately

sloping backwards 2

External apical spine of hind tibia present. Head conical or acutely conical, with face strongly
sloping backwards or vertical 16

2 Coxa of middle leg with large conical process, sometimes with a small tubercle

. MECOSTIBUS Karsch
Coxa of middle leg without process, sometimes with a small tubercle ...... 3

3 Ninth abdominal tergite of male with a median dorsal bifurcate appendage .... 4
Bifurcate appendage on mid-dorsal margin on ninth abdominal tergite absent .... 5

4 Frons vertical or sloping forwards, slightly excurved or straight; fastigium of vertex not protrud-

ing. Body small but comparatively robust SHELFORDITES Karny

Frons sloping backwards, slightly incurved, fastigium of vertex and upper part of frons strongly
protruding forwards. Body elongate, slender KALAHARICUS Brown

5 Dorsum of pronotum crossed by four deep transverse sulci . . . MECOSTIBOIDES Dirsh
Dorsum of pronotum crossed by three or fewer transverse sulci, or sulci not crossing disc at all . 6

6 Frontal ridge deeply sulcate below level of antennal sockets 7

Frontal ridge weakly sulcate to flat below level of an tennal sockets ...... 8

7 Fastigium of vertex horizontal, triangular, seen from above, strongly projecting in front of eyes

. . SYGRUS I. Bolivar

Fastigium of vertex sloping obliquely forwards; as seen from above (Fig. 27) weakly projecting
in front of eyes . MICROUSAMBILLA gen. n. (p. 10)

8 Fastigium of vertex moderately or slightly projecting in front of compound eyes, as seen from

above 9

Fastigium of vertex not projecting in front of eyes NYASSACRIS Ramme

9 Hind femur not inflated bilaterally but strongly bilaterally compressed 10

Hind femur often stocky and inflated, not bilaterally compressed 11

10 Interocular distance in males about as wide, in females twice as wide, as antennal scape.

Fastigium of vertex in males, from above, seen to be excised at apex . PARALENTULA Rehn
Interocular distance in males about twice, in females about four times, as wide as antennal scape.
Fastigium of vertex, as seen from above, widely rounded or truncate . . . LENTULA Stal

1 1 Body moderately to very smooth. Frontal ridge moderately widened in upper part with weak

carinulae and shallow depression. Male supra-anal plate with black tubercles (Figs 35, 50-54,

94-100) 13

Body strongly rugose. Frontal ridge strongly widened in upper half, strongly carinulate. Supra-
anal plate lacking tubercles on disc 12

12 Hind femora stocky, length to depth ratio R about 3.2. Compound eyes strongly inflated, width

across eyes to pronotal width ratio 3 : 2. Length of pronotum at dorsal midline shorter than
its width. Supra-anal plate and ninth abdominal tergite of male lacking any callosities

KARRUACRIS Dirsh

Hind femur slender, ratio R about 4.0. Compound eyes only moderately inflated, width across
eyes rather less than maximum pronotal width as seen from above. Pronotal length at midline
equal to its greatest width. Supra-anal plate and ninth abdominal tergite with small marginal
callosities MALAWIA Dirsh

13 Hind femora of males yellow or greenish yellow with genicular part black. Eyes protruding, red

in males. Penis valves elongate, flagelliform (Fig. 15). Female ventral ovipositor valves shovel-
shaped (Fig. 24) CHROMOUSAMBILLA gen. n. (p. 5)

Hind femora green, brown or yellowish brown with genicular part dark brown, light brown,
blue or some other colour other than black. Eyes in males never red. Penis valves barbed (Figs
38, 72, 133). Female ventral ovipositor valves pointed, tapered (Figs 147, 148) .... 14

14 Margin of abdominal tergite 9 in males bearing one or two pairs of black tubercles (Fig. 35).

Penis valves apically attenuate with barbs apical. Anterior penis valves short, auricular (Fig.

37). Posterior part of median pronotal carina raised into a large tubercle (Fig. 36) in females.

ALTWSAMBILLA gen. n. (p. 12)

Rear margin of tergite 9 of abdomen without conical black tubercles. Penis valves tapering,

conical in dorso-posterior view (Figs 72, 133). Anterior penis valves flat, vertically orientated

and racket-shaped (as in Fig. 12) 15

N. D. JAGO

15

16

17

18

19

20

21

Male supra-anal plate with eight to twelve strong black tubercles on the disc (Figs 50, 54). Male

penis valves with apical barbs (Figs 64, 67, 68, 72) . . . RHAINOPOMMA gen. n. (p. 13)
Male supra-anal plate with eight to ten strong black tubercles on its disc (rarely as in Figs 97, 99,
with pair of weak ones making 12 in all). Male penis valves with pre-apical lateral barbs (Figs
133,135,138,141,142,145,150) . . . USAMBILLA Sjostedt (p. 20)

Fastigium of vertex not projecting in front of eyes, sloping strongly forwards and forming part of
face . . ... . EREMIDIUM Karsch

Fastigium of vertex projecting or strongly projecting in front of eyes, horizontal . . . . 17

Basal angles of fastigium of vertex detached from eyes and projecting laterally; body strongly

rugose SWAZIACRIS Dirsh

Basal angles of fastigium of vertex of normal shape, touching eyes. Body smooth or only

moderately rugose 18

Frontal ridge, in profile, more or less strongly projecting in upper half or upper third . . . 19

Frontal ridge, in profile, straight, not projecting in upper part 22

Fastigium of vertex elongate. Antennae ensiform, serrated . . . . DEVYLDERIA Sjostedt

Fastigium of vertex shorter. Antennae phylliform or filiform 20

Frontal ridge in upper projecting part lamelliformly compressed, below shallowly sulcate . . 21

Frontal ridge sulcate along whole length BASUTACR1S Dirsh

Antennae very short, phylliform, compressed and strongly widened, about four times longer
than wide. Fastigium of vertex with lateral carinulae. Pronotum shorter than its width. Body

rugose KARRUIA Rehn

Antennae narrow, almost filiform. Fastigium of vertex without carinulae. Pronotum longer than
its width. Body smooth GYMNIDWM Karsch

14

16

Figs 10-16 Chromousambilla species, male phallic complex. 10, 14-16, lateral aspect of aedeagal
valves of (10) C. robertsoni; (14) C. mweruensis; (15) C. burtti; (16) C. veseyi. 11, epiphallus of C.
robertsoni. 12, lateral aspect of right side of entire complex in C. latestriata. 13, ventral aspect of
cingular rami and anterior valves of penis of C. latestriata. Scale-lines represent 0.5 mm — that below
Fig. 12 applies to 10 and 12-16; that below 11 to that figure. Small arrows on Figs 10 and 15 show
equivalent position on each set of valvulae.

REVISION OF USAMBILLA 5

22 Fastigium of vertex comparatively short, angular. Body slender, moderately elongate . . 23
Fastigium of vertex elongate, angular or narrow parabolic, with obtusely angular apex. Body

slender and very elongate 24

23 Fastigium of vertex broadly angular, wider than longest diameter of eye. Frontal ridge sulcate

along whole length. QACHAS1A Dirsh

Fastigium of vertex narrowly angular, much narrower than longest diameter of eye. Frontal
ridge in upper fourth compressed, without sulcus, sulcate below . . HEL W1GACRIS Rehn

24 Fastigium of vertex angular. Frontal ridge above base of antennae low, compressed. Antennae

thick, strongly and regularly narrowing towards apex, weakly compressed in basal half. Body
smooth BETISCOWES Sjostedt

Fastigium of vertex narrow parabolic. Frontal ridge above base of antennae produced, strongly
lamelliformly compressed. Antennae very wide, strongly compressed laterally, only slightly
narrowed at apex. Body rugose . . . BACTERACRIS Dirsh

CHROMOUSAMBILLA gen. n.

Type-species: Adolfia latestriata Ramme.

DIFFERENTIAL DIAGNOSIS. Male. Separable from closely related genus Usambilla by great length of aedeagal
valves, which lack barbs (Figs 10, 14, 15, 16). (Proximal section of female spermathecal duct (Fig. 23)
correspondingly stiffened and elastically thickened for a distance corresponding to length of aedeagal valves
distal to their basal attachment below greatly inflated cingular arch.) In life aedeagal valves curled over
greatly inflated membranous arch of cingulum (Fig. 12), their tips touching ectophallic membrane posterior
to epiphallus. Cingular rami also very unusual, small and parallel, being approximated on mid-line so that
they lie between flattened anterior (internal) valvulae of penis (Figs 12, 13). Epiphallus very small and simple,
lacking ancorae, function of which taken over by hook-like development of ventro-lateral corners of
epiphallus (Fig. 1 1). Whole apparatus housed within enlarged pod-like subgenital plate (Figs 4, 8).

Fastigial depression gently concave, its mid-frontal edge being notched as viewed from above (Fig. 2).
Inter-ocular groove well developed, short and wider capitad than caudad. Differs from that of Usambilla
sensu stricto, which is in form of an elongated slot (Fig. 92) or a slightly wider parallel-sided groove. Body
shape like that of Usambilla (Fig. 4).

Supra-anal plate bears callosities of type found in Usambilla, but proportions of supra-anal plate to
sub-genital plate clearly differ (Figs 8, 94-101).

Female. Ventral valves of ovipositor (Fig. 24) straight-edged ; median process of subgenital plate almost
level with their tips.

Coloration. General colouring very uniform in genus. Male eyes vermilion, frons blackish; three lateral
light yellow or whitish stripes on side of head (Fig. 4). Upper pair of light stripes continued along thorax and
abdomen. Subgenital plate yellow. Posterior femora yellow to yellow-green, knees black; posterior tibiae
yellow-green, sooty at extreme tip and near knee. Dark body stripes blue to blue-black or brown. Female
more variable, but in general (Figs 17, 20, 21, 26) lighter brown to olivaceous dorsally, with light banding
like that of male, but with uppermost of light (dorso-lateral) bands weak and lower one on pronotum strong.
Broad dark lateral band variable in intensity. Light bands yellow or creamy colour. Posterior leg colour as
for male but less intense. Body colour of female C. veseyi exceptional in that body and head olive green
dorsally and uppermost dorso-lateral light bands bright yellow. Lateral dark bands very black so that
female of this species resembles male more closely than do those of rest of genus.

There is great instability in the callosities of the male supra-anal plate and too much variation for this
feature to be reliable diagnostically.

DISCUSSION. The genus is known only from NW. Zambia and Tanzania. C. mweruensis in the
Mweru Wa Ntipa and Malagarasi R. drainages, which were once part of the Congo R. drainage
basin before the formation of Lake Tanganyika by rifting, may be indicative of the great antiquity
of the genus. Each species seems confined to the wettest eco-zones of a separate drainage basin.
Some of these, as in the Dodoma area or Mpwapwa basin, are closed internal drainages. Others
like the Ruaha R. valley are open systems, but have no ecological bridges suitable for the genus
by which they can enter other drainages.

6 N. D. JAGO

Key to species

Males

1 Antennae fully twice length of head and pronotum. Large species with blue/black dark lon-
gitudinal stripes on body. Light longitudinal stripes white to cream.
Subgenital plate larger in profile (Fig. 3) than other species in genus (Fig. 4) and narrowed at

level of cercus apices as seen from above (Fig. 5) C burtti sp. n. (p. 9)

Antennae clearly less than twice length of head and pronotum. Dark body stripes grey, blue or

Prussian blue to blue-green. Light longitudinal stripes yellow 2

2 Supra-anal plate laterad, at level of cercus apices (Figs 8, 9), black in addition to black callosities 3
Supra-anal plate without black areas in addition to black callosities (Figs 6, 7)

. . . . . . . . . C. mweruensis sp. n. (p. 9)

3 Pronotum and more particularly subgenital plate clothed with sparse hairs (Fig. 8). Black area of

supra-anal plate broadly pigmented to base 4

Pronotum and subgenital plate almost without hairs (Fig. 9). Black areas of supra-anal plate not
reaching base of plate C. robertsoni sp. n. (p. 10)

4 Lower light yellow band on pronotum laterad, roughly equal in depth to dark band immediately

above it (Fig. 1) C. veseyi sp. n. (p. 8)

Lower light yellow band on pronotum laterad, half depth of dark band immediately above it
(Fig. 3) C. latestriata (Ramme) (p. 6)

Females

1 Median dorsal carinula of pronotum elevated and clear to weak but entire 2

Median dorsal carinula of pronotum weak, flattened (Fig. 22).

Colouring akin to that of males with median dorsal part of body olive brown and dorso-
lateral yellow band visible and entire as far back as 8th abdominal tergite (Fig. 19)

. C. veseyi sp. n. (p. 8)

2 Large species (Fig. 21). Pronotal side stripe white, bordered above and below by black. Antennae

1.50 times longer than head and pronotum together . ..... C. burtti sp. n. (p. 9)

Smaller species. Pronotal side stripe less heavily delineated or even absent. Antennae less than
1.45 times longer than head and pronotum together 3

3 Lower outer lobe of knee of hind femur uniformly creamy or light brown in colour. Ratio of

length of antennae to that of head and pronotum (R) about 1.0 or 1.4. Dorso-lateral yellow

pronotal line very weak or absent (Figs. 20 and 26) 4

Lower outer lobe of knees of hind femur with additional black markings. Ratio (R) about 1.2.
Dorso-lateral yellow pronotal line narrow but usually clear (Fig. 17) ....
C. latestriata (Ramme) (p. 6)

4 Brownish insects (Fig. 26); body length less than 18 mm. Ratio of antennal length to length of

head and pronotum (R) about 1.0 C. robertsoni sp. n. (p. 10)

Olivaceous or brownish insects (Fig. 20); body length more than 18 mm. Ratio (R) 1.4.

C. mweruensis sp. n. (p. 9)

Chromousambilla latestriata (Ramme) comb. n.

(Figs 2, 4, 8, 12, 13, 17, 18,24,25)

Adolfia latestriata Ramme, 1929: 307, fig. 29c. Holotype & TANZANIA: Ukimbu, Nkila, 20-2 l.viii. 1899
(Glauning) (MNHU, Berlin) [examined].

DIFFERENTIAL DIAGNOSIS. Male with supra-anal plate blackened laterally (Fig. 8), thus differing from C.
burtti and C. mweruensis. Aedeagal valves longer than those of C. robertsoni or C. veseyi (see Figs 10, 12, 16);
in former, pronotum and subgenital plate almost lack hairs, and black areas on supra-anal plate do not
extend to its base. In contrast, C. latestriata and C. veseyi are sparsely but clearly hirsute on pronotum and
subgenital plate, and black areas of supra-anal plate are as in Figs 8, 9. C. latestriata differs from C. veseyi in
proportions of lateral pronotal bands (Figs 1, 3).

Female differs markedly in colouring from C. veseyi (see key) and lacks well-developed median pronotal
carina of C. burtti. Lower outer lobe of knee of posterior femora with dark spots, thus differing from
uniformly light brownish lobe of C. robertsoni and C. mweruensis.

REVISION OF USAMB1LLA

26

Figs 17-26 Chromousambilla species, females. 17, lateral aspect of head and thorax of C. latestriata.
18, dorsal aspect of head and pronotum of C. latestriata. 19-21, lateral aspect of head and thorax of
(19) C. veseyi; (20) C. mweruensis; (21) C. burtti. 22, dorsal aspect of head and pronotum of C. veseyi.
23, dissected display of ventral ovipositor valves (vv) and spermatheca showing point distal to which
duct is non-sclerotised (arrow) and apical sac plus preapical diverticulum. 24, ventral valves of
ovipositor, from below, of C. latestriata. 25, dorsal aspect of head and pronotum of C. latestriata. 26,
lateral aspect head and thorax C. robertsoni. Scale line under Fig. 17 represents 1 mm and applies to
all except Figs 23 and 24, to which 1 mm scale line under Fig. 23 applies.

8 N. D. JAGO

MEASUREMENTS

Males Females

Head width (11) 3.4-3.7,3.52 (8) 4.1-4.4,4.26

Antenna length (9) 5.6-6.4,6.72 (7) 6.0-7.2,6.71

Posterior femur lengh (11) 7.1-8.3,7.88 (7) 9.2-10.2,9.73

Posterior femur depth (11) 2.0-2.3,2.11 (7) 2.5-2.9,2.68

Body length (11) 12.0-14.1,13.09 (8) 16.8-18.8,17.81

MATERIAL EXAMINED

Tanzania: 4 & 8 ?, Rukwa, Kapombo Hill, 25.iv.1958 (Vesey-FitzGerald) (BMNH); 1 & 1 ?, Ufipa, Ilemba
gap, 12.iii.1959 (Vesey-FitzGerald) (BMNH); 11 rf, 10 ?, Ufipa plateau, Mkundi, 26 km NNW. of Sumba-
wanga, 16-27.V.1966 (Jago) (5 (J, 2 9; rest COPR, London); 1 cJ, 2 $, Rukwa valley, Red Locust Camp,
Musa, 26-27.ix.1964 (Jago) (BMNH).

DISCUSSION. This species was originally described from 'Ukimbu (S.O.-Tanganyika)'. At that time
Tanganyika' was applied to a region embracing parts of what are now Katanga, northern
Zambia and SW. Tanzania, and 'Ukimbu' was an area to the east of L. Rukwa. The COPR
collections include material from the Rukwa valley and Ufipa plateau west of L. Rukwa. The
females from the valley floor have dark colouring with two light yellowish body stripes laterad,
the rest of the insect being dark greenish. Females from the plateau lack the upper pair of lighter
bands laterad but are lighter insects overall.

A collection was made of the plant association characterising the herb cover of the habitat.
This insect occurred together with Usambilla haematogramma and was particularly abundant at
the base of old termite hills at the edge of a forestry plantation. The plants were kindly identified
in 1967 at the East African Herbarium by S. P. Kibuwa; the species were Bidens steppia (Steetz)
Scherff, Erlangea sp. near E. laxa S. Moore (Compositae); Hypoestes verticillaris R. Brown
(Acanthaceae); Pycnostachys (?) stuhlmanni Guerke (Labiatae); Pseudarthria hookeri Wight &
Arnott (Papilionaceae); Achyranthes aspera L. (Amaranthaceae); Physalis peruviana L. (Sola-
naceae).

Like U. haematogramma, C. latestriata seems to occur in a plant community which is structur-
ally part of a woodland subclimax. It does not live in the forest or grassland and would therefore
seem to be an insect adapted to a transient ecosystem. The Ufipa forests seem to be at a curious
senescent stage in which many of the broad-leaved trees are dying, leaving many areas with tree
euphorbias as dominants. Local ecologists implicate climatic change and the effects of man in
accelerating the decline in this unique forest mosaic.

Chromousambilla veseyi sp. n.

(Figs 1,9, 16, 19,22)
Holotype J, Tanzania: Ruaha. Nat. Park, riverine, 15.iii.1966 (Vesey-FitzGerald) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male with supra-anal plate (Fig. 9) blackened laterally, thus differing from C. burtti
and C. mweruensis. Like latter, however, in having short aedeagal valvulae (Figs 14, 16). Unlike C. robertsoni
in having moderately hirsute pronotum and subgenital plate. Dark body stripes definitely bluish black, not
black or dark brown. Light body stripes golden yellow, differing in proportions from C. latestriata (see Figs
1, 3; also key, p. 6).

Female differs from all other species (Fig. 19): generally darker dorsally and olive green with clear
dorso-lateral yellow side stripes. Dark lateral stripes black.

MEASUREMENTS

Males Females

Holotype

Head width 3.5 3.6 4.1 4.4

Antenna length 6.9 7.6 7.6 7.9

Posterior femur length 7.8 8.1 9.7 10.3

Posterior femur depth 2.1 2.2 2.6 2.8

Body length 14.1 13.2 16.2 15.9

REVISION OF USAMBILLA 9

MATERIAL EXAMINED
Paratypes. Tanzania : 1 <$, 2 $, same data as holotype ( 1 <J, 1 $ in COPR, London ; 1 $ in BMNH).

DISCUSSION. This new species is named after the late L. D. E. F. Vesey-FitzGerald, a close friend
and colleague. The species is unique in having the male and female with similar bright colour
patterns.

Chromousambilla burtti sp. n.

(Figs 3, 5, 15, 21)
Holotype <J, Tanzania: Kikombo, Mpwapwa, 17.iv.1947 (E. Bum) (BMNH).

DIFFERENTIAL DIAGNOSIS. Both sexes distinguished at once by larger size. Antennae in both sexes longer
proportionately than other species, twice length head and pronotum in male, 1.5 times in female. Supra-anal
plate of male lacking areas of black pigmentation (Fig. 5), thus differing from C. latestriata, C. veseyi and C.
robertsoni. Dark body stripes of male dark brown; light body stripes dull cream. Female (Fig. 21) with dark
brown side stripes with other darker regions black. Dark side stripe on metathorax and first abdominal
tergite strongly arched. Back of female pronotum often green. Male and female with pronounced though
slender median dorsal carinula.

Male genitalia (Fig. 15) very long and aedeagus narrowed pre-apically with an oblique tip. Whole
complex enclosed in large pod-shaped sub-genital plate (Fig. 3).

MEASUREMENTS

Male holotype Female allotype

Head width 3.9 5.0

Antenna length 10.5 10.3

Posterior femur length 9.6 11.9

Posterior femur depth 2.6 3.3

Body length 15.7 21.2

MATERIAL EXAMINED

Paratypes. Tanzania: 2 £, 1 £ allotype and 2 9, same data as holotype (1 & 1 $ (allotype), 1 $ BMNH; 1 &
1 ? COPR, London); 1 cJ, 1 $, Kikombo, Mpwapwa, 20.iv.1947 (E. Burn) ($, BMNH; rf COPR,
London); 1 <J, 2 $, Kikombo, Mpwapwa, 18.iv.1947 (E. Bunt) (BMNH).

DISCUSSION. The species is named after the late Eric Burtt who did so much to improve our
knowledge of the acridid fauna of central and eastern Africa.

Chromousambilla mweruensis sp. n.

(Figs 6, 14,20)
Holotype £, Zambia: Mporokosa distr., Mweru wa Ntipa, [8-13.]vii.l952 (Uvarov) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male differs from all other species, except C. burtti, in having supra-anal plate free
of black shading laterally (see Figs 6, 7). Aedeagal valves resemble C. veseyi (see Figs 14, 16), but this differs
on supra-anal plate facies. Male dark body stripes blue-black, those of C. burtti being black. Latter also
bigger species with longer antennae (ratio of antennal length to length of head plus pronotum 2.0 in males,
1.5 in females); antenna in C. mweruensis having equivalent ratios for male 1.8 and female 1.0.

MEASUREMENTS

Males Females
Holotype

Head width 3.5 3.8 4.5 4.3

Antenna length 6.9 7.7 8.1 9.4

Posterior femur length 8.6 8.2 10.2 9.4

Posterior femur depth 2.3 2.2 2.8 3.0

Body length 16.1 14.5 19.5 21.1

10 N. D. JAGO

MATERIAL EXAMINED

Paratypes. Zambia: 1 *, same data as holotype (BMNH). Tanzania: 6 cJ, 2 ?, 35.4 km W. of Kahama,
Mkwemi, 14-29.iii.1947 (£. Burn] (1 J, 1 ? COPR, London; rest BMNH); 4 J, 35.4 km W. of Kahama,
Mkwemi, [xii.1946-ii.1947] (£. Burn) (BMNH); 2 & Old Shinyanga, l.ii.1947 (£. Burn) (BMNH);2 9, 16.1
km N. of Ussure, on Msigiri rd, 12.iv.1936 (£. Bum) (BMNH).

DISCUSSION. The species name originates from the Mweru wa Ntipa area of Zambia.

Chromousambilla robertsoni sp. n.

(Figs 7, 10, 11,26)

Holotype <J, Tanzania: 70.8 km N. of Dodoma, nr Meia Meia, [16-18.]vi.l967 (Jago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Aedeagal valvulae (Fig. 10) of intermediate length, pointed apically like C. la-
testriata and C. veseyi, with black patches laterad on supra-anal plate, but these areas not reaching broadly
to base of plate. Male colour and body markings like those of C. latestriata, but lower outer lobe of hind
knee mostly white with limited dark spots (unlike C. latestriata in which this lobe extensively mottled with
black). Pronotum and subgenital plate almost without hairs. Female almost uniform dull brown (Fig. 26)
with weakly differentiated creamy diagonal side stripe. Hind femora dull olive to yellow with lower outer
knee lobe pale brown, immaculate (unlike C. latestriata which has dark spots on this area). Ratio of antenna
length to length of pronotum plus head about 1.8 in males and 1.4 in females; antennae being shorter in
proportion than any other species.

MEASUREMENTS

Males Females

Head width (12) 3.2-3.5, 3.41 (2) 4.1 4.1

Antenna length (11) 6.2-7.7,6.78 (2) 6.6 6.3

Posterior femur length (12) 7.4-9.1, 8.03 (2) 9.8 9.4

Posterior femur depth (12) 1.9-2.4,2.16 (2) 2.9 2.7

Body length (12)11.0-13.0,12.20 (2)13.1 17.4

MATERIAL EXAMINED
Paratypes. Tanzania: 10 £, 2 9, same data as holotype (9 <J, 1 9, BMNH; 1 £, 1 9, COPR, London).

DISCUSSION. The species is named for Dr I. A. D. Robertson and his wife Ann who are formidable
collectors of acridids. The species occurred in rather dry Acacia woodland in the same biotope as
Physocrobylus burtti Dirsh (described in Jago, 1978).

MICROUSAMBILLA gen. n.

(Figs 27-33)

Type-species : Usambilla cylindricollis Ramme.

DIFFERENTIAL DIAGNOSIS. Male. Vertex with narrow selliform groove opening forward into disc-shaped
fastigium (Fig. 27). Pronotum rounded above, lacking median carina (Fig. 29). Antennae slightly flattened,
not widened and about 1.25 times length of head and pronotum. Ninth abdominal tergite deeply excavate
(Fig. 28), bearing black tubercles on either side of median emargination. Supra-anal plate simple, triangular,
bearing two large submarginal tubercles laterally and up to three pairs of small tubercles on disc. Penis
complex like that in Usambilla, with large racket-shaped anterior valves and aedeagus bearing a single pair
of pre-apical lateral teeth (Figs 32, 33). Epiphallus simple, ancorae (Fig. 31) weak and forwardly directed,
ventro-lateral lophus (Figs 30, 31) at right angles to disc and with single apical hook. Hind femur 3.8 times
longer than deep, rather slender.

Female. Head as seen from above very similar to that of male. Pronotum with traces of median dorsal
carina. Antenna about 1.25 times length of head and pronotum. Ventral ovipositor valves slender, pointed.
Hind femora slender, like those of male.

Coloration. General colour pattern of both sexes similar (Fig. 29). Light brown above, dark brown
laterally. Dorso-laterally, pair of faint whitish stripes, side lobe of pronotum with another arcuate band of
same colour but twice as deep. Hind femora olive, brown above. Hind tibiae brown, sooty below.

REVISION OF USAMBILLA

11

Figs 27-33 Microusambilla cylindricollis. 27, dorsal aspect of male head. 28, dorsal aspect of male
supra-anal plate. 29, dorsal aspect of female pronotum. 30, right side, epiphallus and lateral plate,
male. 31, epiphallus of male, right side. 32, entire male phallic complex (epiphallus removed), lateral
oblique view, left side. 33, cingular rami and apical valvulae, rear aspect. All scale lines represent
0.5 mm.

Figs 34-40 Altiusambilla modicicrus. 34, dorsal aspect of male head. 35, dorsal aspect of male supra-
anal plate. 36, dorsal aspect of female pronotum. 37, entire male phallic complex, apical valvulae, rear
aspect. 39, left side of epiphallus and lateral plate, male. 40, right side of epiphallus. Scale line under
Fig. 37 represents 0.5 mm and applies to Figs 35, 37-40. Other scale lines represent 0.5 mm.

12 N. D. JAGO

DISCUSSION. The genus is monotypic. It is morphologically close to Sygrus from which it differs in
having a flattened sloping fastigium of the vertex and a weaker system of black nodules on the
male ninth tergite. Sygrus also has a more definite dorsal pronotal carinula (see key, p. 3). In
future, when further material is available, it may be best to unite Sygrus and Microusambilla
under one genus.

Microusambilla cylindricollis (Ramme) comb. n.

Usambilla cylindricollis Ramme, 1929: 302, fig. 27a. Holotype <3, ZIMBABWE: Mashonaland, Chirinda forest,
1 150 m, 22.ii.1907 (C. F. M. Swynnerton) (BMNH) [examined].

MEASUREMENTS

Male Female

Head width 3.0 4.9

Posterior femur length 7.7 9.0

Posterior femur depth 2.0 2.0

Body length 12.1 14.2

MATERIAL EXAMINED

Zimbabwe: 2 & 3 ?, Monarch Mine, Umtali, 1213 m, 6.iv.l945 (N. C. E. Miller) (BMNH);6$, same data
but 7.iv. 1945 (BMNH).

ALTIUSAMBILLA gen. n.

(Figs 34-40)

Type-species : Lentula modicicrus Karsch.

DIFFERENTIAL DIAGNOSIS. Male. Head and fastigium rugulose (Fig. 34) with transverse carinulae in depressed
anterior part of fastigium. Upper part of frons, between antennal bases concave and widened. Groove of
vertex ending just behind narrowest approximation of compound eyes dorsally, two oblique ridgelets
separating it from the highly pitted occiput. Pronotum smoothly rounded above, median carinula being
weak but present along whole length of disc. Dorso-lateral light pronotal stripe very narrow while ventral
band on pronotal side lobe very deep, extending from pronotal margin one-third of way up side lobe.
Posterior margin of ninth abdominal tergite gently concave, bearing one and sometimes two black tubercles
on each side (Fig. 35). Supra-anal plate without marginal callosities of the Rhainopomma type (Figs 50-54)
but with at least six pairs of tubercles on the disc and base of distal appendage (Fig. 35). Phallic complex like
that in Lentula, with small auricular anterior penis valves (Fig. 37). Penis valves styliform with apical barbs
(Fig. 38). Epiphallus with no ancorae and with lophal lobes slender, turned upwards at 45° to epiphallic disc
(Fig. 40) and ruggedly hooked at tip (Fig. 39). Length to depth ratio of hind femur 4.

Female. Angular differentiation of frontal groove and fastigium more pronounced than in male, latter
being pitted and sloping forward at 45°C to horizontal body axis. Lateral body stripes relatively undifferen-
tiated. Dorsal median pronotal carina forming a large hump in the metazona (Fig. 36). Tergites of abdomen
sharply carinulate. Tips of ventral ovipositor valves quadrilateral with acutangular apices.

Coloration. Fore and midlegs yellow green. Hind femur yellow green ; knee lenules blackish, rest brown.
Hind tibiae pale bluish yellow-green, sooty apically. Generally olivaceous brown or greenish with creamy
pronotal stripes in males. Dark brown to blackish side stripe in males and females from behind eye to side of
first abdominal segment. On latter segment often black, continuing as broken black zig-zag on each side to
abdominal segment 6.

Altiusambilla modicicrus (Karsch) comb. n.

Lentula modicicrus Karsch, 1896: 280. Holotype J,
TANZANIA: Madjame, Mt Kilimanjaro (MNHU, Berlin) [examined].

MEASUREMENTS

Males Females

Head width (64) 2.5-3.0, 2.69 (42) 3.0-3.5, 3.24

Posterior femur length (60) 6.0-7.0, 6.55 (39) 7.8-9.3, 8.67

Posterior femur depth (62) 1.5-1.8, 1.68 (39) 2.0-2.4, 2.20

Body length (59)10.9-13.4,12.11 (39)13.1-17.1,15.98

REVISION OF USAMBILLA 13

MATERIAL EXAMINED

Tanzania: 1 J, Moshi, 18.xi.1943 (£. Sum) (BMNH); 3 ?, Kirua Vunja, 28.ix.1952 (Guic/jarrf)(BMNH); 2
$, W. Kilimanjaro, Msituni, defoliating Finns patula, 10.xii.1975 (BMNH); 1 9, W. Kilimanjaro, Moshi, on
Pinus patula (BMNH); 23 <$, 5 9, nymphs, Kilimanjaro, S. side nr Mandera hut, 2370-2730 m, 15.xi.1964
(Jago) (BMNH); 30 rf, 29 9, E. of Mt Meru, Ngurdota-Meru N.P., crater lake rim, forest glades, 22.xi.1964
(BMNH); 5 cJ, 1 $, Ngurdota-Meru N.P., Kisari L., 22.xi.1964 (Jago) (BMNH); 3 rf, 3 9, Mt Meru, 1426 m,
25.ii.1967 (£. 5. Brown) (COPR, London); 5 & 7 $, Jekukumia R., 3°14'S, 36°47'E, 2.vi.l972 (Robertson &
Robertson) (COPR, London).

DISCUSSION. The genus is monotypic. The new generic name refers to the preference this species
has for montane forest and lower heath zones on Kilimanjaro and Meru mountains.

The unusual pronotal morphology of this species is reflected in a less extreme manner in other
female Lentulidae, e.g. Usambilla emaliensis, which also have an elevated crest in the pronotal
metazona. Live material of A. modicicrus is interesting since its mating display shows a possible
behavioural function for the dorsal humps. Males approach females, the female responding by
jerking and opening the hind legs sideways, while pedalling the hind tibiae which are folded and
unfolded alternately. The males often orientate incorrectly head to tail, the female responding by
more violent jerking or kicking. If, however, the male approaches correctly from behind he climbs
the female abdomen and nibbles at the female tergites after the fashion of some female cock-
roaches. In A. modicicrus the male climbs forward until he nibbles the pronotal bulge, whereupon
the female presents her abdominal tip for copulation. A similar function may be involved in the
strange bilateral pits on the metazona of many flightless Catantopinae, e.g. Aresceutica and
Serpusia species.

Its attack on Pinus patula was preceded by years in which this grasshopper was very abundant
in the indigenous natural vegetation. At the higher habitats A. modicicrus lives in a zone prone to
unstable weather conditions with irregular rainfall. It is possible that drought conditions, forcing
the insect to sample exotic Pinus, were responsible for the initial phase of the adaptation that
insects have shown to a new food plant. Forestry involving Pinus patula should be free of attack
by this species if trees are planted in areas other than Meru and Kilimanjaro. The very similar
Rhainopomma montanum found in the West Usambara and Pare mountains is unlikely to adapt
to Pinus species. Care should be taken not to introduce A. modicicrus to other highland areas of
East Africa. This could be easily done if young trees were transferred with egg pods hidden in soil
at their roots.

RHAINOPOMMA gen. n.

(Figs 4 1-72)

Type-species : Adolfia usambarica Ramme.

DIFFERENTIAL DIAGNOSIS. Male. Differing from all other Lentulidae and particularly the closely similar genus
Usambilla in the extreme apical position of the aedeagal barbs (Figs 63, 64). Otherwise form of supra-anal
plate (Figs 50-54) and epiphallus as in Usambilla (compare Figs 65, 144). Inter-ocular space very narrow, a
feature correlated with great inflation of the compound eyes (Figs 41-45).

Female. Vertex narrower in proportion between compound eyes than in females of Usambilla (compare
Figs 49, 83), but this apart identical with that genus.

Coloration. Male more brightly coloured than the female. Hind tibiae blue in some species, a character so
far known from only one Usambilla species.

DISCUSSION. Distribution of the genus suggests a long period of evolution in wet lowland to
middle altitude montane forests in eastern Africa, east of the Rift Valley. Habitats generally in
higher rainfall areas than those of Usambilla. Females only surely identified generically by
association with the males.

The genus is distributed in SE. Kenya and the piedmont forest blocks of the Pare, Usambara,
Nguru, Uluguru mountains and Pugu hills. It is thus indigenous to forests on ancient uplifted
mountain blocks which have not experienced the volcanism characteristic of the central rift.
Their speciation suggests isolation in forest islets as a major factor in speciation.

14

N. D. JAGO

Figs 41-49 Rhainopomma species, dorsal aspect of head. 41, R. usambaricum, male. 42, R. montanum,
male. 43, R. nguruense, male. 44, R. wapugu, male. 45, R. magnificum, male. 46, R. usambaricum,
female. 47, R. nguruense, female. 48, R. wapugu, female. 49, R. magnificum, female. All scale lines
represent 0.5 mm, that under Fig. 42 applies only to that figure.

Figs 50-54 Rhainopomma species, male supra-anal plate. 50, R. usambaricum. 51, R. montanum. 52, R.
nguruense. 53, R. wapugu. 54, R. magnificum. Scale line represents 0.5 mm.

Key to species

Males

1 Knees of posterior femora blue R. usambaricum (Ramme) (p. 16)

Knees of posterior femora red-brown 2

2 Posterior tibiae sooty brown in apical four-fifths R. wapugu sp. n. (p. 19)

Posterior tibiae uniformly coloured or only with black or brown pigment at extreme tips . . 3

3 Supra-anal plate bearing fewer than 10 black teeth or denticles on disc (Fig. 42). Small insects

without clear dorso-lateral light body stripes R. montanum (Kevan) (p. 17)

Supra-anal plate with at least 14 black teeth or denticles on disc. Larger insects, usually with clear
dorso-lateral body stripes .... . .... 4

REVISION OF USAMB1LLA

15

Light body stripes white, upper and lower stripe viewed laterally being clearly delineated above.

Smaller insects, body length under 15mm. Hind tibiae bright blue . . R. nguruense sp. n. (p. 17)
Light body stripes creamy to buff in colour with upper lateral stripe more poorly delineated on its

upper margin. Large insects, body length never less than 16 mm. Hind tibiae dull green.

R. magnificum sp. n. (p. 20)

Females

1 Hind tibiae uniform blue, spines black tipped 2

Hind tibiae green with or without blue-black pigment 3

2 Hind knee lunules (Figs 55, 59) dark slatey blue, blacker above. Hind femur otherwise yellowish.

Light dorso-lateral and lateral body stripes golden yellow in colour

R. usambaricum (Ramme) (p. 16)

Hind knee and lunules (Figs 56, 61) light brown. Hind femur dark olivaceous with dorsal part
brown. Light dorso-lateral and lateral side stripes light ochreous brown

R. nguruense sp. n. (p. 19)

3 Hind tibiae distally blue-black; proximal quarter green. Pronotum with weak but distinct convex

carina dorsally R. wapugu sp. n. (p. 19)

Hind tibiae light yellow-green. Pronotum dorsally lacking median carina and decumbent, con-
cave in profile, or lightly convex, especially at rear 4

4 Hind femur 3.25 times longer than deep. Large insects; body length 18 mm. Upper carina of hind

femur convex (Figs 58, 62) R. magnificum sp. n. (p. 20)

Hind femur over 4.00 times longer than deep. Smaller insects with body length 15 mm. Upper
carina of hind femur almost straight R. montanum (Kevan) (p. 17)

Figs 55-62 Rhainopomma species. 55-58, male posterior femora of (55) R. usambaricum; (56) R.
nguruense; (57) R. wapugu; (58) R. magnificum. Scale line under Fig. 58 represents 1 mm and applies
throughout. 59-62, lateral aspect of female head, thorax and hind femur of (59) R. usambaricum; (60)
R. wapugu; (61) R. nguruense; (62) R. magnificum. Scale line under Fig. 61 represents 5 mm and
applies throughout.

16 N. D. JAGO

Rhainopomma usambaricum (Ramme) comb. n.
(Figs 4 1,46, 50, 55, 59, 63-65)

Adolfia usambarica Ramme, 1929: 305, figs 28b, 29b, 30a. Holotype ._?, TANZANIA: Amani, Usambara, iii. 1906
(Vosseler) (MNHU, Berlin) [examined].

DIFFERENTIAL DIAGNOSIS. Male. Antennal segments 1-3 green, 10 and 13 white, rest black. Head (Fig. 41)
with dorsal triangular occipital area black, frons and two stripes behind and from ventro-posterior edge of
compound eye, dark brown. Vertex, a band round back edge of compound eye and lower half of genae,
white. Pronotum dark brown with pair of dorso-lateral and lateral wide white bands, which are yellow in
metazone. White pronotal bands continued along abdomen as yellow bands, abdominal sternites and
sub-genital plate being yellow. Supra-anal plate (Fig. 50) green, with two pairs of basal and three pairs of
submarginal black tubercles. Hind knees (Fig. 55) blue, knee lunules grey, rest of femur yellow-green. Fore
and mid-femora yellow to green. Hind tibiae blue; spines white, black-tipped. Aedeagus (Figs 63-65) and
epiphallus; aedeagal sheath pear-shaped from posterior aspect (Fig. 64) (like that of R. nyuruense, R.
montanum and R. wapugu), lateral epiphallic sclerites large in proportion to rest of epiphallus (unlike those of
R. montanum, Fig. 69), with lower lophal lobe of epiphallus pointed but not hooked (Fig. 65, upper, left).

Female. Dark brown markings on head and pronotum as for male (Fig. 59), but equivalent white areas
yellow, while median dorsal pronotal carina delineated with yellow. Fastigium of vertex (Fig. 46) black with
yellow margins. Dorso-lateral yellow bands on body clearly marked back to at least abdominal tergite 1.
Antennal markings similar to male, but black areas dark brown in this sex. All femora green, hind femora
yellow on inner side. Hind knees and posterior tibiae dull blue.

MEASUREMENTS

Males Females

Head width (13) 3.4^4.0,3.64 (20) 4.0-5.0,4.45

Inter-ocular distance (14) 0.10-0.24, (20) 0.34-0.49,0.42

Posterior femur length (14) 7.6-9.5,8.37 (20) 9.3-11.0,10.18

Posterior femur depth (14) 2.0-2.7,2.32 (20) 2.5-3.1,2.85

Total length (14) 13.4-17.3, 14.90 (20) 17.2-21.0, 18.90

MATERIAL EXAMINED

Tanzania: 1 ?, same data as holotype (BMNH); 1 9, Amani, iv.1905 (BMNH); 3^, Sigi nr Amani, 460 m,
ll.vi.1937 (£. Burn) (BMNH); 7 J, 1 9, same locality but 24.vi.1937 (E. Bunt) (BMNH); 4^, 4?, same
locality but 29.V.1932 (£. Bum) (BMNH); 1 <$, 1 9, same locality but 6.vii.l937 (£. Burtt) (BMNH); 1^, same
locality but 4.ii.l937 (£. Burtt) (BMNH); 3 & 5 9, same locality but v.1937 (£. Bunt) (BMNH); 1 rf, Kwamtili
plant, iii. 1952 (Phipps) (BMNH); 1 J, 2 9, Ngomeni, Mlingano for., i.1953 (Phipps) (BMNH); 1 9, same
locality but vii.1952 (Phipps) (BMNH); 5 rf, 6 9, 1 nymph, E. Usambara Mts, Sigi nr Amani, [2-1 l.]iv. 1964
(N. D. Jago) (COPR, London); 1 <J, 7 ?, 3 nymphs, same locality, [1 8-3 l.]xii. 1965 (N. D. Jago) (BMNH); 1
cJ, 1 9 nymph, W. Usambara Mts, Mkussu for. res., 1 l.xi.1964 (N. D. Jago) (BMNH); 2^, 19, E. Usambara
Mts, Kwamkora for. res., 7.iv.l966 (N. D. Jago) (BMNH); 2^, 1 9, E. Usambara Mts, Amani-Sigi for. res.,
xii.1966 (N. D. Jago) (BMNH); 1 <J, 1 9, E. Usambara Mts, Amani Rest. Ho., 5.iv.l966 (N. D. Jago)
(BMNH); 4 J, 4 9, E. Usambara Mts, nr Amani, 9.xi.l964 (N. D. Jago) (BMNH). Kenya: 1 J, 49, Shimba
hills, iii. 1941 (VanSomeren) (BMNH).

DISCUSSION. R. usambaricum occupies wet lowland forest in the east Usambara mountains and
immediately adjacent hills. Its ecological preference is typical for the other members of the genus,
which also prefer wet, warm and sunny lowland forest, in contrast to upland habitats. Chromous-
ambilla parallels it in having brightly coloured males, but differs in occupying degraded montane
forest at higher altitudes in the hinterland of east Africa. R. montanum (see material studied)
penetrates a little higher into the wetter warmer forest areas of the West Usambara massif and
converges in appearance on A. modicicrus to such an extent that for many years the two were
confused taxonomically.

Rhainopomma montanum (Kevan) comb. n.
(Figs 42, 5 1,68, 69)

Usambilla montana Kevan, 1950: 21 1, fig. 3b. Holotype^, KENYA: Teita hills, 1370-1680 m, shrubby bushes
and forest clearings, 24.xii.1945 (Kevan) (BMNH) [examined].

REVISION OF USAMBILLA 17

DIFFERENTIAL DIAGNOSIS. Male. Much smaller than R. usambaricum. Antennae with segments 1-3 green but
rest dull brown, segments 8 and 12 being light in colour, cream or green. Head (Fig. 42) dark brown with
black markings, at upper end of frons, around antennal sockets, a pair of spots just behind posterior end of
groove of fastigium of vertex, a pair of dorso-lateral narrow post-ocular stripes and a broader one laterally
behind compound eye, all black. Dorso-lateral light brown band on pronotum very narrow (contrasting
with R. usambaricum) but lower lateral cream band wide (half depth of dark brown band above it). Rest of
thorax and abdomen light brown above narrowly edged with lighter brown; black to dark brown laterally.
Abdomen green below; subgenital plate green. Supra-anal plate green with only three pairs of callosities on
disc; marginal callosity on each side continuous, unbroken (Fig. 51). Fore and mid-legs dark green; hind
femora (like Fig. 58) dark green, knees light brown. Posterior tibiae black at the extreme tip, rest green but
light brown adjacent to knee. Spines white with black tips.

Female. Small with proportions of inter-ocular space much like those of/?, magnificum (Fig. 49). Body and
head light brown dorsally with only a faint trace of a fine line of lighter colour along each side. Dark brown
to black lateral pronotal band similar to that in Fig. 60, but lower lateral light brown to cream band deeper
than in R. wapugu (half depth of dark band) and upper edge of dark lateral band on segments 1 and 2 of
abdomen forming a smooth curve, not angulate. Lateral parts of tergites 1-5 with dark brown or black
markings. Hind femora uniformly red-brown, or yellow on inner and lower sides, green on outer area with
knee red brown. Hind tibia green, spines green with black tips; extreme tip of tibiae black, part near to hind
knee light brown. Fore and midlegs green or brown.

MEASUREMENTS

Males Females

Head width (27) 2.7-3.0, 2.87 (31) 3.3-3.8, 3.80

Inter-ocular distance (27) 0.18-0.30,0.22 (31) 0.34-0.63,0.48

Posterior femur length (26) 6.1-7.0, 6.40 (31) 7.1-9.7, 8.07

Posterior femur depth (26) 1.6-2.0,1-79 (31) 2.1-2.4,2.22

Total length (26)11.1-13.2,11.74 (29)13.8-20.3,15.65

MATERIAL EXAMINED

Kenya: 4 <$, 59, 1 9 nymph, Taita hills, vi.1948 (Van Somm>n)(BMNH); 19, Kenya colony, 1921 (/I. F. J.
Gedye) (BMNH); 26 J, 29 9, 9 nymphs, Taita farmers' training centre, 8 km S. of Wundonyi, 6.V.1975 (/. A.
D. & A. Robertson) (COPR, London). Tanzania: 6 J, 7 9, W. Usambara Mts, forest above Mazumbai,
8.vii.l967;3J, 1 9, same locality, nr irrigation canal, 8.vi. 1963; 2 <J, 29, 1 cJ, 29 nymphs, W. Usambara Mts,
Amboni estate, below Mazumbai, 8.vii.l967; 6 J, 2 9, Mombo-Same rd, foot of S. Pare Mts, Gonja, wet
forest, 13.xi.1964; 3 J, 3 9, W. Usambara Mts, Mazumbai F. Res., vi.1967; 1 <£ W. Usambara Mts, nr
Bumbuli, Mazumbai, l.i.1967; 15 J, 1 9, W. Usambara Mts, Sumamagamba F. Res., 12.xi. 1964; 14^,8 9, W.
Usambara Mts, Mkussu F. Res., ll.xi.1964; 1 £, W. Usambara Mts, Lushoto-Shume rd, 9.7 km from
Lushoto, 1 l.xi.1964; 6 £, 4 9, W. Usambara Mts, Lushoto, 1670 m, 8.vii.l967; 1 & 39, W. Usambara Mts,
nr Shume, ll.xi.1964; 1 9, W. Usambara Mts, Lushoto arboretum, ll.xi.1964; 6 J, 89, W. Usambara Mts,
Mazumbai, secondary regrowth in quinine plantation, 8.vii.l963. (All (Jago); all in BMNH except last
COPR, London.)

DISCUSSION. Recorded eating Commellina species (A. and I. A. D. Robertson, pers. comm.) in its
Taita hills habitat in Kenya. Here the partial cultivation of a hill slope at the edge of cultivated
land offered a particularly favourable habitat, partly shaded by an Albizzia species and palms.
Like its sister genus Usambilla this genus is favoured by the opening up of primary forest and
population density may be very high, e.g. 100 + /square metre. This may contribute to the attack
by Usambilla on exotics planted in the course of forestry in such areas.

Rhainopomma nguruense sp. n.

(Figs 43, 52, 56, 6 1,66, 67)

Holotype J, Tanzania: E. foot Nguru Mts, Mtibwa Forest Reserve, nr Turiani, 7.xi.l964 (N. D. Jago)
(BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Segments 1-3 of antennae green; other segments black, except segments 10
and 14 which are white. Colour pattern on head as in R. usambaricum, but fastigium between eyes margin-
ally narrower (Fig. 43). Light pronotal stripes white but otherwise pattern as in R. usambaricum. Meso- and
metathorax dark brown, light body stripes white to pink. Posterior third of abdomen, including sub-genital

18

N. D. JAGO

Figs 63-72 Phallic complex in males of Rhainopomma species. 63, left lateral aspect of R. usambaricum.
64, posterior apical aspect of R. usambaricum. 65, epiphallus of R. usambaricum — upper right half and
lateral plate, lower from right side. 66, left lateral aspect of R. nguruense. 67, posterior apical aspect of
R. nguruense. 68, posterior apical aspect of R. montanum. 69, epiphallus of R. montanum — upper right
half and lateral plate, lower from right side. 70, epiphallus of R. magnificum — upper right half and
lateral plate, lower from right side. 71, left lateral aspect of R. magnificum. 72, posterior apical aspect
of R. magnificum. Scale line under Fig. 71 represents 0.5 mm and applies throughout.

REVISION OF USAMBILLA 19

plate, pale green. Supra-anal plate (Fig. 52) with two pairs of basal tubercles and two or three pairs on disc
(therefore resembling Fig. 53). Posterior femora green with knees red-brown (Fig. 56). Fore and mid-legs
entirely green. Posterior tibiae light red-brown near knee and black at extreme tip; rest blue with black
tipped white spines. Aedeagal complex (Figs 66, 67) showing aedeagal valves less sharply deflexed forwards
than in R. usambaricwn (Fig. 63) but more elongate than those of/?, montanum (Fig. 68). Epiphallus like that
of/?, usambaricum (Fig. 65).

Female. Body colour and pattern (Fig. 61) like that of R. usambaricum but light body stripes white, not
yellow. Median dorsal pale line weak or absent. Hind femora yellow to green; knees brown. Posterior tibiae
pale blue, brown near knee, brown at extreme tip and with red-tipped white spines.

MEASUREMENTS

Males Females

Head width (23) 3.6-4.5, 3.79 (18) 4.2^4.7, 4.54

Interocular distance (23) 0.16-0.30,0.23 (18) 0.46-0.59,0.53

Posterior femur length (23) 7.2-8.5, 7.96 (18) 8.5-10.5, 9.86

Posterior femur depth (23) 2.1-2.6, 2.36 (18) 2.7-3.2, 2.92

Total length (23) 14.0-15.8, 15.03 (18) 17.1-20.0, 18.71

MATERIAL EXAMINED

Paratypes. Tanzania: 16 J, 10 ?, 6 nymphs, same data as holotype but [5-7.]xi.l964 (BMNH); 1 <$, same
data as holotype (BMNH); 1 J, same data but 7.xi.l964; 5<$, 5$, 3 nymphs, same data but 5.xi.l964(COPR,
London).

DISCUSSION. This species is closely similar to R. usambaricum and was probably derived from
common stock. Isolation in the adjacent but ecologically isolated forests of the east Usambara
and Nguru mountains would explain the divergence between the two species.

Rhainopomma wapugu sp. n.

(Figs 44, 53, 57, 60)
Holotype J, Tanzania : Pugu hills, SW. of Dar es Salaam, 1 1 .iii. 1967 (N. D. Jago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Antennal segments 1-5 cream to light brown; rest dark brown excepting
segments 10 and 14 which are white. Interocular space narrowest for genus (Fig. 44). Pattern similar to that
of /?. usambaricum but light areas white to pink, while dark areas are dark red-brown. Fore and mid-legs
entirely green. Hind femora (Fig. 57) light green with light brown genicular part. Hind tibiae light brown
near knee. Distal three-quarters dark brown, remaining proximal section green. Tip of abdomen pale green.
Supra-anal plate (Fig. 53) with two pairs of basal tubercles and three pairs of tubercles on disc; apical
tongue rather large, two-fifths length of whole plate when measured on mid-line.

Female. Antennal segments 1-3, apical parts of 4, 5 and 6, whole of 9, 11 and 15 white, rest black.
Dorso-lateral light body stripes weak, narrow, thus differing from R. usambaricum and /?. nguruense, but
resembling /?. montanum. Dorsal side of head with pair of black triangular marks on either side of dorsal
mid-line. Dark brown lateral stripes (Fig. 60) about 3 times deeper than lower light pink side stripe and
angulate on tergite 1 of abdomen. Fore and mid-legs green, femora brown dorsally. Hind femora light
brown above and internally with large grey spot at base; outer area green; lower outer area yellow; knee
entirely red-brown. Colour of hind tibiae as for male.

MEASUREMENTS

Males Female

Holotype Paratype Paratype

Head width 3.4 3.6 4.37

Inter-ocular distance 0.15 0.17 0.48

Posterior femur length 7.2 7.2 9.54

Posterior femur depth 2.1 2.1 2.77

Total length 13.7 13.4 17.72

MATERIAL EXAMINED

Paratypes. Tanzania: 1 $, 1 9, data as holotype (BMNH).

20 N. D. JAGO

Rhainopomma magnificum sp. n.

(Figs 45, 49, 54, 58, 62, 70-72)

Holotype <$, Tanzania: Mombo-Same rd, foot of S. Pare Mts, Gonja, wet forest, 13.xi.1964 (N. D. Jago)
(BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Large species, total length about 17 mm. Antennae green in basal half,
darkening to brown distally with light cream tip. Vertex and occiput (Fig. 43) with relatively wide ocular
interspace, dark brown dorsally and on frons. Cream area between frons and sub-ocular sulcus; gena dark
brown bordered above by very narrow cream stripe from rear of compound eye to back of occiput.
Pronotum and body back to tergite 3 of abdomen with dark brown dorsal stripe bordered laterally by bold
dull yellow bands equal in width to that on pronotal side lobe (unlike female where lateral pale stripe like
that of male but dorsolateral pale band lacking (Fig. 62)). Fore and mid-legs green. Hind femora green with
knees light brown (Fig. 58). Hind tibiae light green, light brown at each extremity and with black-tipped
green spines. Abdomen, including supra-anal plate, brown above, green below. Sub-genital plate green.
Supra-anal plate (Fig. 54) with up to seven pairs of non-marginal tubercles; much wider than long. Base of
aedeagus (Figs 71, 72) parallel-sided, not flask-shaped. Epiphallus with well-developed hook on apex of
ventral lophi and large anterior projection of upper lophal lobe (Fig. 70).

Female. Head and body light brown above. Antennae like those in male. Dark black band behind
compound eye bordered with narrow cream stripe ventrally (Fig. 62). Upper margin of dark brown side
stripe with distinct up-curved zone at front of pronotal prozona and above side of thoracic tergite III and
abdominal tergite I. Black zone extends from abdominal tergite I along side of tergites to back of segment 4.
Ovipositor valvulae greenish. Fore and mid-legs green. Hind femora and tibiae as in male, but femora light
brown dorsally. Large insects for the group (see measurements).

MEASUREMENTS

Males Females

Head width (5) 4.1^4.4,4.28 (3) 5.1,5.1,4.4*

Inter-ocular distance (5) 0.29-0.32,0.31 (3) 0.79,0.84,0.69*

Posterior femur length (5) 9.0-9.8,9.28 (3) 11.1,11.7,9.4*

Posterior femur depth (5) 2.6-2.9, 2.74 (3) 3.5, 3.4, 2.8*

Total length (5) 16.0-18.7, 17.57 (3) 21.8, 21.5, 18.2*

* Specimen from slightly higher elevation at Soni.

MATERIAL EXAMINED

Paratypes. Tanzania: 3 ^, 2 ?, 1 nymph, same data as holotype (1 <$, 1$, COPR, London; rest BMNH); 1
cJ, data as holotype but 10.vi.1967; 19, Soni, 17.xi.l 950 (J.PWpps) (BMNH).

USAMBILLA Sjostedt

Usambilla Sjostedt, 1909: 191. Type-species: Usambilla olivacea Sjostedt, by original designation.

Adolfia Rehn, 1914: 147. Type-species: Adolfia insolita Rehn, 1914: 148, by original designation. [Homonym

of Adolfia Guerich, 1909 (Brachiopoda).]
Rehnula Uvarov, 1939: 457. [Replacement name for Adolfia Rehn.] [Synonymized by Dirsh, 1956: 152.]

DIFFERENTIAL DIAGNOSIS. Male. Vertex with narrow (Fig. 86) to wide (Fig. 92) inter-ocular sulculus. Frons
depressed (Figs 90, 91) with frontal ridge almost flat, hardly elevated. Dorsal pronotal carinula moderately
developed to weak or absent. Supra-anal plate, but not ninth abdominal tergite, armed with marginal
callosities as well as elevated black nodules on disc (Figs 94-101). Epiphallus simple, lacking true ancorae
but with hook-like ventro-lateral lophus (Figs 144, 145, 150, 131, 134, 137, 140) terminating in one, two or
four hooks. Penis valves with lateral barbs situated short of apex (Fig. 133).

Female. Vertex proportionately wider than male, but with no special generic facies. Ventral ovipositor
valves unspecialised (Figs 148, 149), not shovel-shaped as in Chromousambilla (Fig. 24).

DISCUSSION. The genus is known from predominantly upland areas of eastern Zaire, Ruanda, the
forests of Uganda, south-western, central and northern Tanzania and the southern half of Kenya.
The species occupy a wide range of ecological niches. Thus U. haematogramma lives in the upper
Ufipa plateau forest margins, but U. insolita and U. sagonai in clearings in the wettest tropical
forests of Ruanda, Zaire and Uganda. U. turgidicrus lives in upland scrub and woodland in

REVISION OF USAMBILLA

21

90

87

Figs 73-93 Dorsal or frontal aspect of head of Usambilla species. 73, male V. turgidicrus turgidicrus.
74, female, same. 75, male U. oraria. 76, female, same. 77, male U. ajfinis affinis, Morogoro. 78, male
V. affinis kikomboensis, Ilonga. 79, female, same. 80, male V. emaliensis. 81, female, same. 82, male U.
chlorophrygana. 83, female, same. 84, male U. leptophrygana. 85, female, same. 86, male U. insolita
paratype, Kwidschwi I. 87, same, frontal aspect. 88, male U. sagonai sagonai, Rwanda. 89, male U.
sagonai fractolineata. 90, same, frontal aspect. 91, female, same. 92, male U. haematogramma. 93,
female, same. Scale line under Fig. 92 represents 1.0 mm and applies throughout.

Kenya and northern Tanzania (Combretum, Commiphora and Acacia or Juniperus associations),
while U. leptophrygana and U. chlorophrygana occupy drier warmer woodland at lower altitudes.
The ecological diversity is matched by subspecific variation in several species, suggesting active
speciation in progress.

22

N. D. JAGO

Figs 94-101 Male supra-anal plate of Usambilla species. 94, U. turgidicrus. 95, U. oraria. 96, U. qffinis.
97, U. leptophrygana. 98, U. insolita. 99, U. sagonai sagonai. 100, U. sagonai fractolineata. 101, U.
haematogramma. Scale line under Fig. 101 represents 0.5 mm and applies throughout.

Key to species and subspecies

Males

1 Ocular interspace dorsally more than 2.5 times width of first antennal segment (Fig. 92). Lateral

black side stripe narrowly edged above with red pigment (as in Fig. 129,9). Supra-anal plate
(Fig. 101) with broad black area confluent with both basal tubercles on each side. Hind
femora light olive green, knee light orange-brown . . . .U. haematogramma sp. n. (p. 36)
Ocular interspace narrower, almost twice width of basal antennal segment. Never with red stripe
delineating upper edge of lateral black stripe. Supra-anal plate with marginal rugosity, but
black pigment never confluent with basal tubercles 2

2 Interocular distance dorsally equal to width of basal antennal segment (Figs 75, 86, 87). Supra-

anal plate with no more than a pair of basal tubercles on each side (Figs 95, 98) ... 3
Interocular distance at least 1.5 times width of basal antennal segment. Supra-anal plate with
two pairs of basal tubercles (Figs 96, 99, 100, 152) or up to three basal tubercles on each side
(Figs 94, 97, 99) 4

3 Hind femur (Fig. 116) more than 3 times longer than deep. (Rwanda and Zaire (?))

U. insolita (Rehn) (p. 34)

Hind femur (Fig. 108) very stocky and thickened from side to side, length to depth ratio about
2.6. Subgenital plate very short (Fig. 95) and folding under parameres which are received into
impressions on its upper surface. Subgenital plate protrudes beyond supra-anal plate by
about length of that plate as seen from above (in U. insolita projecting by roughly twice length
of supra-anal plate). (Coastal Kenya and NE. Tanzania) U. oraria sp. n. (p. 32)

4 Dorso-lateral and light lateral body stripes (Figs 102, 103) bright yellow in life. Hind tibiae blue.

Fore and mid-femora conspicuously hairy above and below. Frons declivate, hardly protrud-
ing as seen from above (Figs 88, 89). Median pronotal carinae absent or feeble ... 5
Dorso-lateral light body stripes; if present, dull grey, light brown but never bright yellow. Hind
tibiae yellowish or greenish, with or without black pigment distally and ventrad. Frons
declivate or decidedly protruding (Figs 73, 82). Fore and mid femora almost without hairs
above. Median pronotal carina weakly to strongly developed (Fig. 105) 6

REVISION OF USAMBILLA

23

Yellow genal stripe continuous to back of occiput
Yellow genal stripe interrupted between back

10

11

U. sagonai sagonai (Ramme) (p. 35)
of gena and occiput. (Figs 102, 103)

U. sagonai fractolineata subsp. n. (p. 36)
Hind femora bearing conspicuous black markings (Fig. 115) and of a uniform light brown
colour. Frons protruding (Fig. 84) and incised as seen from above. Fastigium of vertex con-
cave anteriorly. Posterior tibiae sooty below and towards tip . . U. leptophrygana sp. n. (p. 34)
Hind femora immaculate or with light markings dorsally at base (Fig. 107), or with a light
pregenicular spot (Figs 109, 110). Frons declivate, protruding or decumbent. Fastigium of

vertex deeply to shallowly concave anteriorly 7

Lateral dark brown side stripe intense, angularly bent in outline dorso-anteriorly on pronotum
(Fig. 106). Frons in profile and from above (Fig. 73) seen to be very produced at its upper end,
forming two tubercles which form margins of deeply incised fastigium of vertex. Hind femora
light brown with or without darker mottling. Median dorsal pronotal carinula and its con-
tinuation on tergites of abdomen weak to strong. Posterior tibiae sooty below and towards tip

8

Lateral dark brown side stripe with smooth upper outline (Fig. 104). Frons in profile and from
above protruding (Figs 77, 78, 82) to strongly declivate and downwardly sloped (Fig. 90).
Hind femora tending to be green with knee orange-brown. Median dorsal carinula of prono-
tum weak (Figs 104, 105) to absent. Posterior tibiae unicolorous without black colour below 9
Hind femur very short and plump, length to depth about 3.0. (Vicinity of W. Usambara Mts)

V. turgidicrus olivacea Sjostedt (p. 27)

Hind femur less robust, length to depth ratio 3.3-3.4 . . U. turgidicrus turgidicrus (Karsch) (p. 25)
Larger insects, body length just over 15 mm. Length to depth of hind femur (Fig. 113) about

3.40-3.50 U. chlorophrygana sp. n. (p. 32)

Smaller insects, body length just under 13 mm at most. Frons only weakly incised as seen from

above (Figs 77, 78, 80). Femur length to depth ratio 3.32-3.66 10

Pronotum lacking median dorsal carinula except for metazonal tubercle 11

Pronotum with clear median dorsal carinula (Fig. 105) in prozona and metazona. Hind femur
slender (Fig. 112) with outer area green and rest tending to be brownish

U. emaliensis sp. n. (p. 30)

Length to depth ratio of hind femur 3.3-3.4 . . . U. affinis affinis Kevan & Knipper (p. 29)
Length to depth ratio of hind femur about 3.6 . . . V. affinis kikomboensis subsp. n. (p. 30)

102

104

105

Figs 102-106 Lateral or dorsal aspect head and/or pronotum of male Usambilla species. 102, U.
sagonai fractolineata, Uganda, Mpanga (darker form). 103, same, Uganda, Mpanga (lighter form).
104, U. emaliensis. 105, same, from above. 106, U. turgidicrus turgidicrus. Scale line under Fig. 102
represents 1 mm and applies throughout.

24 N I) JA(K)

I .iniil.s

Females of I/, Insoliia were not available for study; females of U, qfflnis offlnin and f. <t//)nfo kikomhoensts
ciinnol he differentiated.

1 I aleial hlack side stripe bordered above with crimson (Fig. 129), Fastigium of vertex very broad

and lightly indented . U. kwmatogramma sp. n. (p. 36)

l.aicnil hlack side stripe, if present, never bordered ahovc with red. Fastigium of vertex much
narrower and more deeply inoiied . . 2

2 Hind lihiac blue. (Colour very variable. No difference between races.) . . £/. saKonai (Kummc) (p. 35)
Hind tibiae light brown, greeniih brown, green or ochrteeoui . . . 3

\ Inieiouilai spaaMloisallv 4 tinuv, width of basalaiitcmi.il segment (Figs 76, 79,81) . 4

Inlerocular space dorsally 5 times width of basal antennal segment (Figs 74, 83, 85) . 6

4 Pi onotal median carina dorsally very pronounced (Fig. 1 25), hind femora very slender, length to

depth ratio about 3, ft . ... U. tmalitnsix sp. n. (p. 30)

Median pronotal carina weak. Mind femur much more robust, length to depth ratio 3.0-3.2 . 5

5 Profile of pronotal carina and thorax curved (Fig. 121). Dark lateral tide stripe with upper edge

markedly bent in pro/ona. Mind lemur length to depth ratio around 3.0, Mind tibiae com-

pletely black vent rally in mature (?) specimens, ycllowish-ochraccous dorsally

. U. oraria sp, n. (p. 32)
Piotile ni pionoial caima Halter (Figs. 122, 123, 124). Lateral side stripe, if developed, with

anterior upper border almost straight. Length to depth ratio of hind femur about 3.2. Hind

tibiae uniformly yellowish . U. qfflitis Kevan & Knippcr (p. 28)

(> Fronx markedly produced in profile (Figs 1 19, 120), Pronotal carina moderately strong, whole

disc tcctiform . ..... ...... 7

Frons less strongly produced in profile (Figs 12ft, 127), Median pronotal carina almost undevel-

oped, disc depressed . 8

Mm, i lemur more robust, length to depth ratio roughly 3.0 or less. Pronotum wrinkled and

punctate above (Fig, 1 20) . (/. turxMicrus olivacta Sjostcdt (p. 27)

Mind femur more slender, length to depth ratio about 3.2. Pronotum not wrinkled, finely punc-

tate (I..- 119). . (/. turxitticrus tur nidi cms Karsch (p. 25)

8 Large insects, body length more than 20 mm, Probably lack lateral body stripe. Posterior femora

dull olivaceous with knee light brown (Figs 83, 12ft) . . . U. cMontphrygaita sp. n. (p. 33)

Siu.illiM rugose msivls. boih length about IS mm. latci.il sulc stupes \\i-.ik cuvpi KM lust

abdominal tcrgitc and latero-antcrior part of pronotum (Fig. 127). Posterior femora brown

with darker mottling , , U. Itptopkrynaita sp. n. (p. 34)

I hambilla turffitlicrus (Karsch)
(Figs 73, 74, 94, 106, 107, 1 19, 120, 131-133)
/.c-Miii/ii lurtiitlicrus Karsch. 1896: 280.

DIIIIKINIIAI DIAONOSIN. Af«i/«', Head uu h deeply grooved fastigium of vertex forming a k notch at its point
of contact with upper eiul of Irons (Fig. 73). Mead transversely ungulate at level of back end of median
fastigial sulculus, occiput being wrinkled and pitted, Sides of vertex in front of eyes and area of ocellus on
each side protruding, Pronotum lightly tcctiform on disc with low median carinula. Upper edge of dark
lateral pronotal side stripe showing sharp angular change of direction between sulculi one and two. Dorso-
lateral light brown to creamy stripes extending to tcrgite 8. Sides of tergites 1-6 of abdomen shiny black.
Supra-anal plate (Fig. 94) with four pairs of strong black nodules on disc, two weaker ones at base.
L'piphallic tophus with lour teeth (Fig, 131). Penis valves (Figs 132, 133) directed forwards at about 45° to
long axis of body,

Fore and midlcgs light olive brown. Mind femora speckled blackish dorso-anteriorly (Fig. 107), with or
without black comma in front of knee lunulc. Hind tibiae light olive brown, spines black-tipped. Hind tibiae
sooty black or darker brown vcntrally in apical two-thirds or half. Lateral dark stripe on pronotal side lobe
extended boldly along side of meso- and metathorax.

. Stocky (Fig. 1 19) with vertex and frontal ridge which meet in a raised protruding shelf in front of

compound eves 1'ionotum lu'htl\ tecliloim above, \\ilh .1 laUM.il d.uk xuK- stupr uiih .in .iiu'.ul.ii up|vi
profile in front of second sulculus .uul i.ulnu-. .il>i upd\ Ivhuul (I iy, I I'M 1 .IUM.I! b.uul on ilioi.u-u- segments
oK, iiu-. il.uk band intensified on alHloinm.il M-I-IIUMIIS I o. hl.uk .nul slum on M-J'.IIUMIIN .' c Ml.uk aiea i>n
side of abdominal segment I variable in si/e.

Kl VISION 01 US

25

01)

ni|l> ill. in, |c liinwil
II I ill VI-

107 118 Posterior femora of male lisunihillti species. 107, (/. nirfiulicrua olhwca, l./D .V(K),
Momhi). IOS. (/. (ir<iri«, L/D 2.97. Momhasa. I(W. (/. «//!MM kikomhtwnsis, l./D 3.6, Kikombo. 110.
same, L/D 3.36, llongti. I II, (/. aljinix u/finis, L/l> 3,28, Uluguru Mis. 112, (/. cmtilu'nsis, l./D «><>.
Mmali range. 1 13, (/. chlorophrygana, L/D 3.46, Kikomho, Mpwapwa. 1 14, (/. chlorophryyana, dark
lot in, l./D 3,26, 10 mis N. of Ussure. 115, (A /«7>f«>/)/irv</<//i<i, l./D 3.32. 1 16, (/. i>i.w//r<i l./D 3.45. 1 17,
U. sagonalfractollntata, l./D 3.47. 1 18, (/. luu'matofirtimmd, l./D 3.55, Dlipa. Scale line under h'ig, 1 12
represents I mm and applies throughout.

Ventral valves of ovipositor simple, pointed. General colour grey to rugose dark brown with black to
blackish pronotal side stripe, bordered below by short curved grey stripe. M-p.u.iti-d from ventral margin of
side lobe by a band of dark brown or grey of roughly same depth.

Usambilla turiiMicrus turgidicrus (Karsch)

(Figs 73, 74,94, 106, 1 14>, 131 133)
U-ntula luryidicrux Karsch. 18%: 280. Holotype •;. KHNYA: Kitui (MNIIll, Merlin) [exannned|

Dn 1 1 RI N IIAI DiA(iNosis. Male hind femora less robust than in subsp. olhwvti, length to depth ratio 3.3 3.4.
Female similarly more elongate with slimmer femora, above ratio 3.2. Pronotum not wrinkled, finely
punctate (Hg. 119).

26

N. D. JAGO

5mm

123

dg dull green

y yellow

ob olive brown

o olive

db dark brown

r red

Figs 119-130 Lateral aspect of head, thorax and posterior femur of female Usambilla species. 119, U.
t. turgidicrus (S2794). 120, U. t. olivacea, W. Usambara. 121, U. or aria, Kenya, Rabai. 122, U. affmis
kikomboensis, Ilonga. 123, same, lighter form. 124, same, intermediate form. 125, U. emaliensis. 126,
U. chlorophrygana. 127, U. leptophrygana. 128, U. sagonaifractolineata. 129, U. haematogramma. 130,
U. sagonai sagonai. Scale line under Fig. 125 represents 5 mm and applies to Figs 1 19-129. Scale line
under Fig. 130 represents 1 mm and applies to that figure.

REVISION OF USAMBILLA 27

MEASUREMENTS

Males Females

Head width (17) 2.9-3.5,3.37 (29) 3.7-4.2,4.01

Posterior femur length (17) 6.3-7.9, 7.23 (28) 7.4-9.4, 8.50

Posterior femur depth (17) 1.8-2.4, 2.20 (28) 2.5-3.0, 2.68

Body length (16) 12.3-15.0, 13.75 (27) 14.8-18.1, 16.69

MATERIAL EXAMINED

Kenya: 1 9, Mtunguni hill nr Tulia, Kitui distr., 150 m, 01°12'S, 38°02'E, hillside with Combretum,
Commiphora and Acacia woodland, 28.iv.1975 (Robertson & Robertson) (COPR, London); 1 <$, 19, nr
Kibwezi, 5 km SE. of Mbuinzau, 02°24'S, 37°55'E, 940 m, lichens and spiney vegetation on recent lava,
5.V.1975 (Robertson & Robertson); 1 £, 1 $, 3 km N. of Mwatate, Wundangi rd, 03°27'S, 38°22'E, 910 m,
fairly dense scrub woodland, 6.V.1975 (Robertson & Robertson) (COPR, London); 1531, 259, Mtunguni hill,
01°12'S, 38°02'E to 01°13'S, 38°02'E, nr Tulia, Kitui distr., 910 m, heavy juniper woodland and plantations,
lO.v.1975 (Robertson & Robertson) (5 & 4 9, BMNH; rest COPR, London); 1 9, Athi R. crossing, 25.6 km
NNE. of Kibwezi, 430 m, 22.vii.1934 (Vanderbilt Exped. Kenya Africa, Rehn) (BMNH); 1 <J, 1 9, S. Pare Mts,
hillside above Gonja, c. 1000 m, [12-16] vi.1974 (Hollis) (BMNH).

DISCUSSION. More material of the nominate subspecies is required before the true junction
between the subspecies can be determined.

Usambilla turgidicrus olivacea (Sjostedt) stat. n.
(Figs 107, 120)

Usambilla olivacea Sjostedt, 1909: 186, 192. Holotype 9, TANZANIA: Usambara, Mombo, vi. (Sjostedt) (NR,
Stockholm) [examined].

DIFFERENTIAL DIAGNOSIS. Male with hind femur length to depth ratio 3. Female with hind femur length to
depth ratio 3 or less. Pronotum wrinkled and punctate above (Fig. 120). Subspecies markedly more stocky
than nominate race. Male genitalia identical.

MEASUREMENTS

Male Females

Holotype

Head width 3.19 4.3 (4.8) 4.20

Posterior femur length 6.66 3.7 (4.0) 8.68

Posterior femur depth 2.25 9.3 (10.0) 2.90

Body length 12.31 14.7 (17.0) 16.15

The female holotype measurements in parentheses are those published by Sjostedt.

MATERIAL EXAMINED
Tanzania: 1 ^, 1 9, Mombo, 9.vii. 1967 (Jago) (BMNH).

DISCUSSION. The dimensions of the holotype published by Sjostedt do not tally with the accurate
modern ones. It is possible that his optical technique gave consistent overestimates. The holotype
is thus a smaller insect than published measurements would indicate. U. turgidicrus olivacea
represents the southernmost outliers of the nominate race.

In life the specimens from Mombo had a light silvery side stripe, the female being light grey in
colour. Both have darkened considerably and are now brown.

Usambilla affinis Kevan & Knipper
(Figs 77-79, 96, 109-111, 122-124, 134-136)
Usambilla affinis Kevan & Knipper, 1961 : 372.

DIFFERENTIAL DIAGNOSIS. Male. Fastigium of vertex (Figs 77, 78) with margins diverging forward, lightly
emarginate in front and with deeply to lightly impressed interocular groove (lightest in subsp. affinis to
deepest in subsp. kikomboensis). Front end of fastigium pitted and sculptured. Frontal ridge and whole of

28

N. D. JAGO

Figs 131-141 Phallic complex of male Usambilla species. 131, epiphallus of U. turgidicrus
turyidicrus — upper right half and lateral plate, lower from right side. 1 32, same, aedeagus, left lateral
aspect. 133, same, posterior apical aspect. 134, epiphallus of U. qffinis ajfinis (Uluguru Mts) — upper
right half and lateral plate, lower from right side. 135, same, aedeagus, posterior apical aspect. 136,
same, left lateral aspect. 137, epiphallus of U. emaliensis (Emali range) — upper right half and lateral
plate, lower from right side. 138, same, aedeagus, posterior apical aspect. 139, same, left lateral aspect
with spermatophore in situ. 140, epiphallus of I/, or aria — upper right half and lateral plate, lower
from right side. 141, same, aedeagus, posterior apical aspect. Scale line under Fig. 138 represents 0.5
mm and applies throughout.

REVISION OF USAMBILLA 29

frons evenly punctate and more protruding than U. oraria, but less so than in U. turgidicrus. Pronotum
hardly to weakly tectiform above with pronotal carina absent to moderate. Abdominal tergites lightly
carinulated dorsally. Supra-anal plate with marginal callosities (Fig. 96) and four to five pairs of tubercles,
two or three pairs being distal. Penis sheath and valves apically vertical (Fig. 136). Epiphallus with lophal
process ending in a simple hook (Fig. 134). Ancorae absent.

Female. Fastigium of vertex sloping forwards in a smooth convexity to meet upper part of frontal ridge
(Figs 79, 122, 123, 124). Pronotum and abdominal tergites quite sharply carinulate dorsally. Tips of ventral
ovipositor valves triangular, pointed.

Coloration of male and female more divergent in the closely allied U. emaliensis than in U. affinis. Male in
general with yellowish to creamy markings at border of compound eye dorsally, on front of fastigium of
vertex, below compound eye straddling fronto-genal suture. Lateral dark brown stripe variable in intensity,
usually extending weakly onto side of abdomen. Light side stripes pale yellowish to creamy white, lower-
most band separated from ventral margin of pronotal side lobe by darker pigment (Fig. 104) and extending
across epimera and episterna of thoracic segments 2 and 3 as a whitish band. Fore and mid legs light
olivaceous green. Hind femora green with ginger-brown knees (Figs 109-111). Very faint brown comma-
shaped marking in front of genicular lunule. Female light brown above with darker median dorsal area, side
stripe dark brown (but variable, see Figs 122, 123, 124). Hind femora light olivaceous green with dorso-basal
brownish spot and weak sigmoid mark in front of outer and inner knee lunule. Knees light ginger-brown.
Hind tibiae, as in male, light yellow-green with spines black-tipped.

Note. In both male and female upper profile of lateral dark pronotal side stripe forms a smooth convex
curve (Fig. 124) unlike that in U. turgidicrus (Fig. 1 19).

DISCUSSION. This species forms a north-south cline in which the southernmost populations have
short stumpy hind femora (subsp. affinis), the northernmost having slender hind femora. Separ-
ating the components of the cline is difficult, though zoogeographically the nominate subspecies
probably occurs in the Uluguru Mts and southwards, while subsp. kikomboensis probably occurs
in scattered pockets from Ilonga through Kikombo to the west Usambaras.

Usambilla affinis affinis Kevan & Knipper
(Figs 77, 96, 111,134-136)

Usambilla affinis Kevan & Knipper, 1961: 372, fig. 3, pi. 3, ff. 3-7. Holotype J, TANZANIA: Morogoro,
Morningside, 31.vii.l954(Pfa>ps)(BMNH) [examined].

DIFFERENTIAL DIAGNOSIS. Male. Main difference in proportions of hind femur, its length to depth ratio at
3.3-3.4 being less than in subsp. kikomboensis. Males tend to have dark dorso-lateral body lines brown.
Penis sheath and aedeagus erect, set at right-angles to long axis of body (Figs 135, 136).
Females cannot be differentiated from other subspecies.

MEASUREMENTS

Males Females

Head width (3) 3.45,3.38,3.74 (3) 4.18,4.08,4.48

Posterior femur length (3) 7.24, 6.99, 7.48 (3) 9.14, 8.30, 9.10

Posterior femur depth (3) 2.21, 2.89, 3.12 (3) 3.00, 2.65, 2.95

Body length (3) 12.3, 12.93, 14.79 (3) 16.5, 17.22, 17.24

MATERIAL EXAMINED

Tanzania: 1 ?, Morogoro, 1954 (Phipps) (paratype of Usambilla affinis Kevan & Knipper) (BMNH); 1 <?,
Morogoro, 1320 m, 24.xi.1939 (£. Burn) (BMNH); 1 rf, 1 $, Morogoro, xi.1939 (E. Burtt) (BMNH); 2cJ,
Morogoro, 8.iii.l955 (Phipps) (BMNH); 1 ?, Morogoro, 4.xii.l939 (£. Burtt); I & Uluguru Mts, W. side, 1.6
km S. of Mgeta, 25.x. 1964 (Jago) (COPR, London); 1 nymph, Uluguru Mts, W. side, Bunduki For. Res.,
fishing camp, [20-23.]x. 1964 (Jago) (COPR, London).

DISCUSSION. The habitats of this subspecies in the Uluguru Mts are characterised by abundant
tree ferns and tree lilies, relatively high rainfall and abundant sunshine.

30 N. D. JAGO

Usambilla affinis kikomboensis subsp. n.

(Figs 78, 79, 109, 110, 122-124)
Holotype ^Tanzania: Ilonga, 16.vi.l 967 (Jago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Hind femora length to depth ratio about 3.6. More brightly marked than
nominate subspecies, with dark brown stripes in that subspecies replaced by black and light body stripes
golden yellow. Penis sheath and aedeagal valves directed vertically (as in Fig. 136).

Female. Pigmentation variable (Figs 122-124). Frons sloping upwards with a gentle angular change into
fastigium of vertex. Face dark brown with black and cream spots or patches. Dark side stripe, when fully
developed (Fig. 122), with a gently sinuous upper margin (may be reduced to a dark oblique bar, Fig. 123).

MEASUREMENTS

Males Females

Head width (14) 3.5-3.7, 3.63 (5) 4.1-4.5, 4.25

Posterior femur length (14) 7.4-8.0, 7.85 (5) 8.9-9.3, 9.13

Posterior femur depth (14) 2.3-2.9, 2.41 (5) 2.6-3.0, 2.79

Body length (14) 13.2-15.0, 14.18 (5) 16.9-17.3, 17.13

MATERIAL EXAMINED

Paratypes. Tanzania: 12cJ,5$, same data as holotype (5(J, 3?,COPR, London, rest BMNH); 1$,
Mpwapwa, Mt Wilkins, 1460 m, 10.iv.1938 (E. Burn) (BMNH); 1 <J, Kikombo, Mpwapwa, 18.ix.1947
(BMNH).

Usambilla emaliensis sp. n.

(Figs 80, 81, 105, 106, 112, 125, 137-139)
Holotype £, Kenya: Emali range, Sultan Hamud, 1210-1790 m, iii.!940(BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Penis sheath bent in a smooth arc capitad as in U. turgidicrus (cf. Figs 1 32,
139). Epiphallic lophus bifurcate at tip (up to four teeth in U. turgidicrus) (Fig. 137). Form of frons and vertex
much as in U. turgidicrus but otherwise differing as follows: upper margins of dark lateral body stripe
smooth in profile (Fig. 104), not angulate as in Fig. 106; pronotal median carina forming a higher crest (Fig.
105) than in U. turgidicrus, in this respect near to V. affinis; hind femur (Fig. 112) distinctly green in outer
and lower outer areas (always light brown with darker mottling in nominate subspecies of U. turgidicrus);
length to depth ratio about 3.66 (only about 3.30 in nominate subspecies of U. affinis); frontal ridge (Figs 80,
81) weakly incised, unlike U. turgidicrus but like U. affinis.

Female. With much more pronounced pronotal crest than U. turgidicrus (Fig. 125). Hind femur much
more long and slender (Fig. 125) (length to depth ratio 3.6; this ratio in subspecies of U. turgidicrus 3.2 or
less).

MEASUREMENTS

Male holotype Female paratype

Head width 3.20 3.75

Posterior femur length 7.36 9.05

Posterior femur depth 2.00 2.56

Body length 13.46 15.78

Figs 142-153 Phallic complex of male Usambilla species. 142, U. chlorophrygana aedeagus, posterior
apical aspect. 143, same, left lateral aspect. 144, same, epiphallus — upper right half and lateral plate,
lower from right side. 145, epiphallus of U. sagonai sagonai (Kibale forest) — left right half and lateral
plate, right — right lateral aspect. 146, same, aedeagus, left lateral aspect. 147, same, posterior apical
aspect. 148, 149, U. sagonai sagonai female, ventral ovipositor valves from below and left side
respectively. 150-152, U. insolita male (150) epiphallus — upper right half and lateral plate, lower right
lateral aspect; (151) aedeagus, posterior apical aspect; (152) aedeagus, left lateral aspect. 153, supra-
anal plate of U. chlorophrygana male. Scale line under Fig. 146 represents 0.5 mm and applies
throughout.

REVISION OF USAMBILLA

31

145

147

152

32 N. D. JAGO

MATERIAL EXAMINED
Paratype. Kenya : 1 9, same data as holotype (in copula) (BMNH).

DISCUSSION. This species may represent an isolated offshoot of U. qffinis with which it shares the
form of the vertex and pronotum. The very slender hind femora and high pronotal crest are,
however, unique.

Usambilla oraria sp. n.

(Figs 75, 76, 95, 108, 121, 140, 141)
Holotype^, Kenya: Mombasa, [3-4.]vii.l939(E. Bum) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Vertex very narrow between compound eyes dorsally (Fig. 75), frontal ridge
depressed, hardly protruding. Dark body stripes dark reddish brown; light longitudinal stripes creamy
white. Lower lateral light band extending across epimera and episterna of thoracic segments 2 and 3. Hind
femora very squatt and inflated, length to depth ratio less than three (Fig. 108). Supra-anal plate (Fig. 95)
very simple with four pairs of black callosities on the disc. Subgenital plate very short, folding under
parameres for which there are two concave depressions in upper surface of subgenital plate itself. Cingular
sheath small, rather abruptly tapered (Fig. 141) compared with U. qffinis (Fig. 135). Epiphallus with bifur-
cate lophi (Fig. 140).

Female. Fastigium of vertex shallowly concave; frons hardly produced. Occiput with two large rugose
ovoid depressions just behind back end of interocular groove. In lateral aspect with smooth convex dorsal
profile giving a hump-backed appearance. Pronotum quite strongly tectiform. Hind femur short and stocky,
length to depth ratio less than 3 (Fig. 121). Ventral ovipositor valves pointed.

General coloration light brown and dark brown pattern in both sexes. Hind femora unicolorous light
brown (Fig. 108), though in darker females with dark comma anterior to knee lunules and marks on
dorso-anterior part. Hind tibiae light ochrous brown, often blackish below, with black-tipped tibial spines.

MEASUREMENTS

Male Females

Head width 3.50 (3) 4.5-5.3,4.79

Posterior femur length 6.95 (3) 8.4^9.0, 8.74

Posterior femur depth 2.35 (3) 2.7-3.1, 2.93

Body length 11.53 (3)15.6-17.1,16.54

MATERIAL EXAMINED

Paratypes. Kenya: 2 ^, Mombasa, [3-4.] vii. 1939 (E. Burn) (BMNH) [compared with holotype of U.
turgidicrus by V. M. Dirsh]; 1 9, Mombasa, on Vernonia hildebrandti, 12.iii.1969 (BMNH); 1 ?, Rabai, viii.
1937 (Van Someren) (BMNH); 3 9, Arabuko for., 17 km W. of Malindi, 03°13'S, 39°56'E, 61 m, ll.vi.1975
(Robertson & Robertson) (COPR, London); 1 9, Arabuko-Sokoke For.Res., Jilore track, c.50 m, [2-3.]vi.l974
(Hollis) (BMNH).Tanzania: 2 9, Mlingano, Ngomeni, v.1952 (Phipps) (BMNH); 1 9, W. Usambara, Suma-
magamba F. Res., 12.xi. 1964 (Jago) (BMNH).

DISCUSSION. Probably one of the most truncated grasshoppers in proportion to its width known
to science. It has a largely coastal forest distribution, with enclaves in piedmont forest in the west
Usambara mountains. It belongs to the affinis-group of species.

Usambilla chlorophrygana sp. n.

(Figs 82, 83, 1 13, 1 14, 126, 142-144)
Holotype <J, Tanzania: Kikombo, Mpwapwa, 16.iv.1947 (E. Burtt) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Large for the genus (see measurements). Fastigium of vertex strongly
declivate, frons only protruding slightly in front of eyes. Anterior part of fastigium heavily pitted. Occiput
with two oblique echelons of finely pitted cuticle terminating just behind interocular groove in two flat
depressions (Fig. 82). Frons and genae wrinkled and punctate. Pronotum finely punctate with weak or
obsolete dorsal carinula. Dorso-lateral and lateral light body stripes dull yellowish brown, upper band weak
dorsad. Lateral creamy stripe extending across epimera and episterna of meso- and metathorax. Supra-anal
plate with at least 3 pairs of distal black tubercles on disc (Fig. 153). Penis sheath strongly decurved capitad
(Fig. 143) so that valves point forward. Epiphallic lophi simple; single hook at apex (Fig. 144).

REVISION OF USAMBILLA

33

Figs 154-161 Phallic complex of male Usambilla species. 154, U. haematogramma aedeagus, posterior
apical aspect. 155, same, left lateral aspect. 156, same, epiphallus — upper right side and lateral plate,
lower right lateral aspect. 157, U. leptophrygana, aedeagus, posterior apical aspect. 158, same, entire
complex less epiphallus from above showing shape of ectophallic sheath and apodemes plus position
of ejaculatory sacs (shaded). 159, same, entire complex less epiphallus, left lateral profile, showing
broad flat spatulate anterior penis valves. 160, same, aedeagus left lateral view. 161, same,
epiphallus — left right half and lateral plate, right lateral aspect. Scale lines represent 0.5 mm. Line
under Fig. 154 applies to all except Figs 158, 159, to which scale line under Fig. 158 applies.

Female. Large (Fig. 126) with cuticle finely pitted. Female paratype badly discoloured but probably with
light band of shiny cuticle across lower edge of pronotal side lobe.

Fastigium very wide, almost flat and punctate anteriorly (Fig. 83). Ovipositor valves apically pointed as in
Fig. 188.

General coloration of male olivaceous brown with creamy orange dorso-lateral and lateral side stripes.
Hind femora light green; knees light red-brown (Fig. 113). Hind tibiae light yellow-green to yellow, with
black spines.

34 N. D. JAGO

MEASUREMENTS

Male holotype Female allotype

Head width 3.71 4.88

Posterior femur length 7.88 10.92

Posterior femur depth 2.51 3.15

Body length 15.36 20.89

MATERIAL EXAMINED

Paratypes. Tanzania: 2 9, same data as holotype (BMNH); 1 $, data as holotype but 17.iv.1947 (£. Bunt)
(BMNH); 1 J, data as holotype but 18.iv.1947 (£. Bunt) (BMNH); 1 & data as holotype but 19.iv.1947 (£.
Burn); 2 <$, Mpwapwa, Mt Wilkins, 1213 m, 4.iv.l938 (£. Bunt) (BMNH); 2 rf, 19, 16.1 km N. of Ussure,
Msigiri road, 12.iv.1936 (£. Bunt) (1 <$, 1 9, COPR, London; 1 <$, BMNH).

DISCUSSION. The name of this new species is derived from the Greek 'chloros' — green and
'phryganos' — twig or stick.

Usambilla leptophrygana sp. n.

(Figs 84, 85, 97, 1 15, 127, 157-161)
Holotype <$, Tanzania: 70.8 km N. of Dodoma, 18.vi.1967 (N. D. Jago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Males. Distinctive (Fig. 127) with well-marked to faint dark brown side stripe and
conspicuous black markings on posterior femora (Fig. 1 15). Frons weakly to strongly incised in front (Fig.
84); fastigium of vertex excavate but much more produced than the similar U. chlorophrygana (Fig. 82).
Overall colour light brown with dark markings. Hind femora not green in our series. Supra-anal plate with
three basal tubercles on each side (Fig. 97) (see U. chlorophrygana, Fig. 153). Epiphallic lophus with single
apical hook (Fig. 161), anterior penis valves (Figs 158, 159) typical for genus and illustrated here as
representative, being flattened and vertically orientated. Aedeagal valves sloping strongly capitad (Figs 157,
159,160).

Female. Dark pronotal side band weak, fading behind pronotal metazone and only intensifying just
behind tympanum. Hind femur with pre-genicular black spot and dorso-basal spot always present, but row
of three dark spots along lower outer carina variably developed (Fig. 127). Overall colour dull brown with
darker brown markings. Median pronotal carina present, if weak, throughout.

MEASUREMENTS

Males Females

Head width (3) 3.5-3.6, 3.58 (5) 4.1-4.5, 4.31

Posterior femur length (3) 7.7-8.4, 8.01 (5) 8.4-9.8, 9.19

Posterior femur depth (3) 2.3-2.5, 2.41 (5) 2.8-3.1, 2.95

Body length (3) 14.2-15.2, 14.81 (5) 16.4-18.5, 17.58

MATERIAL EXAMINED

Paratypes. Tanzania: 3 9, same data as holotype but 16.vi.1967 (BMNH); 1 cJ, 2 ?, Old Shinyanga,
(l-8.)iii.!947 (£. Bunt) (BMNH); 3 & same data, 7.iii.l947 (BMNH); 3 ^,same data, Hi. 1947 (BMNH); 1 &
same data, 12.iv.1947 (BMNH); 1 1 <J, 5 9, same data, early iii.1947 (1 & 1 9, COPR, London; rest BMNH);
7 cJ, Old Shinyanga, 30.iii.1947 (£. Bunt) (BMNH); 1 <J, 1 9, same data 30.iv.1947 (BMNH); 1 <J, Old
Shinyanga, block 9, 23.V.1956 (£. Bunt) (BMNH); 2 & same data, [22-24.]iiil948 (£. Bunt); 1 9, same data,
24.iv.1948 (£. Bunt) (COPR, London); 3 J, Mkwemi, 17.7 km W. of Kahama, 29.iii.1947 (£. Bunt) (BMNH);
1 9, Ruaha N. P., 15.iii.l 966 (Vesey-FitzGerald) (BMNH).

DISCUSSION. The name of this species is derived from the Greek 'leptos' — slender, 'phryganos' — a
twig.

Usambilla insolita (Rehn)

(Figs 86, 98, 150, 151)
Adolfia insolita Rehn, 1914: 148. Holotype^, CONGO: L. Kivu, Kwidschwi I. (MNHU, Berlin).

DIFFERENTIAL DIAGNOSIS. Only male paratype examined (ex alcohol). Very similar to U. sagonai (see Figs 86,
87, 88, 90). Differs in having a much narrower vertex and frontal ridge hardly produced forwards. Zone of
dark brown or black pigment in U. sagonai (Fig. 90) apparently absent in U. insolita, but this may be due to

REVISION OF USAMBILLA 35

loss of colour after attempted preservation in alcohol. Supra-anal plate similar to that of U. oraria (Fig. 98)
but differing in having subgenital plate extending much further beyond tip of supra-anal plate (see key,
couplet 3) and hind femur more slender (see Figs 108, oraria; 116, insolita). Epiphallus (Fig. 150) with stout
hook at tip of ventral lophi and penis apex curved in an arc cephalad (Figs 151, 152).
General colour pattern unknown.

MEASUREMENTS

Male

Head width 3.32

Posterior femur length 7.51
Posterior femur depth 2.07

Body length 13.53

MATERIAL EXAMINED

Paratype. Congo: 1 <$, Kwidschwi I., Lake Kivu, ix.1907 (Adolf Friedrich Duke of Mecklenberg Expdn)
(COPR, London).

DISCUSSION. There is clear confusion in the original descriptions of species allocated to Adolfia by
Ramme (1929) so that his expansion of U. insolita to include material from modern Zaire and the
Ruwenzori region must be open to doubt. The paratype (topotype) examined here is different in
detail from U. sagonai and is probably a species confined to the Kivu area only. It is replaced in
Uganda by U. sagonai.

Usambilla sagonai (Ramme)
(Figs 88, 89, 90, 99, 100, 102, 103, 117, 128, 130, 145-149)

DIFFERENTIAL DIAGNOSIS. Male. Differing from all other Usambilla species by the following combination of
characters: dorso-lateral and light lateral body stripes (Figs 102, 103) bright yellow; hind tibiae blue; frons
only very slightly produced forwards as seen from above (Figs 88, 89); hind femora green, knee lunules light
reddish brown (Fig. 117); supra-anal plate (Figs 99, 100) open to variation but in general with two pairs of
basal black tubercles and two pairs of black tubercles near margin at centre of disc; frontal ridge less
coarsely pitted than that of U. insolita (Figs 87, 90); penis valves sharply curved cephalad (Figs 146, 147);
ephiphallic lophi with single apical hook (Fig. 145).

Generally brightly coloured species found in wet evergreen forest. Oblique yellow stripe across gena from
base of eye complete or broken (subsp.fractolineata, Figs 102, 103).

Female. Ovipositor valvulae unspecialised; bluntly pointed (Figs 148, 149). Brightly coloured in various
complex shades of green and brown (Fig. 130). In the darkest forms lateral pronotal lobe and area across
thoracic segments II and III plus abdominal tergite 1 bear shining black bands. In more lightly pigmented
specimens the black areas may be replaced by brown or dark green (Fig. 128). Side of pronotum, below dark
band, bright yellow.

In both sexes pronotum is smoothly arched dorsally with median dorsal carinula weak or absent and
cuticle finely punctate. In general appearance they converge on Rhainopomma species but have a wider
inter-ocular groove, and, of course, apical barbs of the male aedeagus are preapical.

Usambilla sagonai sagonai Ramme
(Figs 88, 99, 130, 145-149)

Adolfia sagonai Ramme, 1929: 305, fig. 28c. Holotype £, ZAIRE; Lakes Region (MRAC, Tervuren)

[examined].

DIFFERENTIAL DIAGNOSIS. Differs from subsp.fractolineata by possessing a yellow genal stripe which is entire.
MEASUREMENTS

Males Females

Head width (12) 3.1-3.7, 3.46 (9) 3.8-4.1, 3.96

Posterior femur length (12) 7.0-8.6, 7.95 (9) 8.4-9.4, 8.95

Posterior femur depth (12) 2.0-3.5,2.43 (9) 2.5-2.9,2.60

Body length (12)12.4-15.0,14.18 (9)15.7-16.9,16.44

36 N. D. JAGO

MATERIAL EXAMINED

Zaire: 1 & 2 9, La Chute For., Rutchuru, 7.viii.l949 (E. Burtt) (BMNH). Rwanda: 2 <J, Kisenye,
10.viii.1949 (E. Burtt) (BMNH). Uganda: 1 V, Toro, 1.6 km E. of Bundebugyo, 9.viii.l964 (Jago) (BMNH);
1 ?,Kamanve, 17.ix.33 (Johnston) (BMNH); 1 9, Kilembe, 1 370 m,xii. 1934-1.1935 (F. W. Awards) (BMNH);
lc^Gaba,18.x.l931(G.//.£./f0p/uns)(BMNH);l J,Nsagu, 3.ix.l933(Jonnsron)(misdet.as V. insolita Rehn
by B. P. Uvarov) (BMNH); 1 <J, Ruwenzori, 1610 m, 1913 (Scott-Elliott) (misdet. Adolfia insolita Rehn by
Ramme) (BMNH); 8 <J, 6 9, Toro, SE. of Ft Portal, Kibale for. res., 30°25'E, 0°30'N, [13-16.]viii.l964 (Jago)
(2 $, 2 9, COPR, London; rest BMNH); 1 cJ, 1 ?, same data, 30.iv.1967 (Jago) (BMNH); 1 9, Ruwenzori,
between road and Nyabitaba hut below 2610 m, 27.vi.1963 (P. & P. Carter) (BMNH). Kenya: 1^, 19,
Kakamega For. Statn, c. 1520 m, [18-19.] vii. 1974 (Hollis) (BMNH).

DISCUSSION. The recent discovery of this subspecies in Kenya extends its known distribution to
the forests of the east side of Lake Victoria. It is possible that the nominate race occurs south and
west of the lake while subsp. fractolineata represents an isolated series of populations on its
northern side in the Mabira to Mpanga forest blocks of the Nile drainage and northwards to
Lake Albert.

I samhilla sagonai fractolineata subsp. n.

(Figs 89, 90, 100, 102, 103, 117, 128)

Holotype <J, Uganda: Buganda, Mpanga F. Res., km 33.8 Kampala-Masaka road, 32°20'E, 0°15'N,
[2-3.] viii. 1964 (Jago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Differing from nominate subspecies only in respect of the broken lateral yellow
band across the gena in the male (Figs 102, 103).

MEASUREMENTS

Males Females

Head width (24) 3.3-4.4, 3.71 (29) 3.8-4.4, 4.09

Posterior femur length (24) 7.4-8.9,8.09 (29) 8.1-9.8,9.18

Posterior femur depth (24) 2.1-2.4,2.24 (29) 2.3-2.9,2.58

Body length (24) 13.6-15.5, 14.65 (29) 15.7-18.9, 17.15

MATERIAL EXAMINED

Paratypes. Uganda: 4 9, 1 nymph, Bunyoro, W. of Masindi, Budongo F. Res., 31°30'E, 1°48'N, [25-
27.]viii.l964 (Jago (BMNH); 6 & 29, Buganda, Mabira F. Res., nr Jinja, 33°0'E, 0°25'N, 27.iv.1967 (Jago)
(BMNH); 10 £, 169, 7 nymphs, same data as holotype (COPR, London); 1 cJ, 1 ?, Mpanga F. Res., nr
Kampala, 30.viii.1969 (E. S. Brown) (BMNH); 1 9, Mpanga F. Res., 1210 m (BMNH); 1 & Bunyoro, Lake
Albert, Butiaba, [26-28.] viii. 1964 (Jago) (BMNH); 1 & 3 9, Bunyoro, Bugoma F. Res., S. of Hoima, 31°0'E,
1°15'N, [29-31.] viii. 1964 (Jago) (BMNH; 4 <$, 2 9, Mubende, Mubende reservoir, 19.viii.1964 (Jago)
(BMNH).

I samhilla haematogramma sp. n.

(Figs 92, 93, 101, 118, 129, 154-156)

Holotype <$, Tanzania: Ufipa plateau, 25.8 km NNW. of Sumbawanga, Mkundi plantation, [ 1 6-27.] v. 1966
(./ago) (BMNH).

DIFFERENTIAL DIAGNOSIS. Male. Inter-ocular groove and fastigium of vertex wide (Fig. 92), frontal ridge flat.
Antennae longer than head and pronotum. Hind femora (Fig. 118) entirely pale green, except knee which is
light brown; hind tibiae light green, dark brown in apical half on inner and lower side, spines black-tipped.
Supra-anal plate (Fig. 101) with dark lateral infuscate areas and four pairs of simple tubercles. Colour, as for
females (Fig. 129), distinctive — body pale emerald green with median dorsal black line above and broad
black line on each side from behind compound eye to back of tergite 8 of abdomen. This lateral line
demarcated above by a red line extending from front of pronotum to at least rear of tergite 1 of abdomen;
demarcated below by a dull yellow band bordered with black ventrally. Cerci conical with narrow pre-apical
black annulus. Subgenital plate pale green. Aedeagal valves (Figs 154, 155) gently curved capitad; epiphallic
lophi (Fig. 156) apically bifurcate.

Female. Colour as for male (Fig. 129) and in this respect unusual for the genus. Combination of colours
unique. Note that mid-dorsal black line may be very faint or absent.

REVISION OF USAMBILLA

37

Males

(35) 3.3-3.5, 3.37
(35) 7.5-9.2, 8.29
(35) 2.1-2.7, 2.27
(35) 13.9-17.5, 15.18

Females

(25) 4.1-4.8,4.20
(24) 9.0-11.9, 10.42
(24) 2.9-3.2, 2.88
(24) 17.8-21.7, 19.45

MEASUREMENTS

Head width
Posterior femur length
Posterior femur depth
Body length

MATERIAL EXAMINED

Paratypes. Tanzania: 2 cJ, 3 9, 4 nymphs, Ilemba gap, 12.iii.1959 (Vesey-FitzGerald); 4$, 2$, 1 nymph,
Nsangu, 2120 m, 13.iii.1959 (Vesey-FitzGerald); 6^, 3 9, Ufipa escarpment, [6-9.] vii. 1948 (Waloff); 2<$,
Malonje, Ufipa, 8.iv.l951 (Vesey-FitzGerald); 1 9, Mpui, 100 km N. of Abercorn, 17.vi.1947 (B. P. Uvarov); 3
cJ, 5 9, Ufipa plateau, 25.8 km NNW. of Sumbawanga, Mkundi, [1 6-27.] v. 1966 (Jago); 2 & 69, E. of
Sumbawanga, Mbisi F. Res., [23-29.] v. 1966 (Jago), 9 3, 39, 19.3 km E. of Sumbawanga, Malonje Mt,
plateau grassland, disused road to Mpui, [24-28.] v. 1966 (Jago); 2<$, 29, Rukwa valley, 8 km W. of Muse,
bottom descent Red Locust road, 26.V.1966 (Jago); 10 & 5 9, Mbisi For. Res., v.1966 (Jago);9J, 29, Ufipa,
Nsangu F. Res., Sumbawanga-Mpui road, 28. v.1966 (Jago). (Last series COPR, London; rest BMNH.)

DISCUSSION. The species derives its name from the Greek 'haematos' — bloody, 'grammes' — a line,
emphasising the distinctive dorso-lateral orange-red line which delineates the upper margin of
the black lateral stripe. The general coloration is very similar to that of members of the genus
Lentula but the phallic complex shows that U. haematogramma has a strong affinity with the
olivacea-group of lentulids.

References

Dirsh, V. M. 1956. Orthoptera, Acridoidea. In Hanstrom, B., Brinck, P. & Rudebeck, G., S. Afr. anim. Life 3:

121-272, 2 pis, 42 figs.
Dirsh, V. M. 1968. A new genus and species of the family Lentulidae (Orthoptera: Acridoidea). Proc. R. ent.

Soc. Lond. (B) 37: 143-145, 2 figs.
Jago, N. D. 1978. The systematic position of Physocrobylus burtti Dirsh, 1951 with description of the

previously undescribed male sex. Acrida 7: 79-83, 7 figs.

Karsch, F. 1896. Neue Orthopteren aus dem tropischen Afrika. Stettin, ent. Ztg 57: 242-359, 38 figs.
Kevan, D. K. McE. 1950. Orthoptera from the hills of south-east Kenya. Jl E. Africa nat. Hist. Soc. 19:

192-221, pi. 34, 5 figs.
Kevan, D. K. McE. & Knipper, H. 1961. Geradflugler aus Ostafrika (Orthopteroidea, Dermapteroidea,

Blattopteroidea). Beitr. Ent. II: 356-413, 9 pis, 12 figs.
Ramme, W. 1929. Afrikanische Acrididae. Revisionen und Beschreibungen wenig bekannter und neuer

Gattungen und Arten. Mitt. zoo/. Mus. Berl. 15: 247-492, 16 pis, 106 figs.
Rehn, J. A. G. 1914. Orthoptera I. Mantidae, Phasmidae, Acrididae, Tettigoniidae und Gryllidae aus dem

Zentral-Afrikanischengebiet Uganda und dem Ituri-Becken des Kongos. Wiss. Ergebn. dt. Zent Afr.-

Exped. 1907-19085(1): 1-223.
Sjostedt, Y. 1909. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedtion nach dem Kili-

mandjaro, dem Mem und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905-1906. 17. Orthoptera. 7.

Acridoidea: 149-199, 1 pi., 2 figs.

Uvarov, B. P. 1939. Twenty four new generic names in Orthoptera. Ann. Mag. nat. Hist. (11)3: 457-459.
Whellan, J. A. 1975. The Acridoidea of Malawi: an annotated check list. Acrida 4: 105-122.

- 1976. Notes on the genus Mecostibus Karsch, 1896 (Orthoptera: Lentulidae). Arnoldia, (Rhodesia) 8(1):

1-6, 8 figs.

Index

Synonyms are in italics; principal references are in bold.

Acacia 10, 21, 27

Adolfia 20

affinis 21, 22, 23, 24, 25, 27, 28, 29, 30, 32

Albizzia 17

Altiusambilla 1, 3, 12

Aresceutica 13

aspera, Achyranthes 8

Bacteracris 5

Basutacris 4

Betiscoides 5

burtti, Chromousambilla 2, 4, 6, 7, 8, 9

burtti, Physocrobylus 10

Catantopinae 13

chlorophrygana 21, 23, 24, 25, 26, 30, 31, 32, 34

Chromothericles 1

Chromousambilla 1, 3, 5, 16, 20

Combretum 21, 27

Commelina 17

Commiphora 21, 27

cylindricollis 10, 11, 12

Devylderia 4

emaliensis 13, 21, 23, 24, 25, 26, 28, 29, 30
Eremidium 4
Erlangea 8

fractolineata 21, 22, 23, 25, 26, 35, 36
Gymnidium 4

haematogramma 8, 20, 21, 22, 24, 25, 26, 33, 36
Helwigacris 5
hookeri, Pseudarthria 8

insolita 20, 21, 22, 25, 30, 31, 34, 35

Kalaharicus 3

Karruacris 3

Karruia 4

kikomboensis 21, 23, 25, 26, 27, 29, 30

latestriata 2, 4, 5, 6, 7, 8, 9, 10

Lentula 3, 12, 37

leptophrygana 21, 22, 23, 24, 25, 26, 33, 34

magnificum 14, 15, 17, 18, 20
Malawia 1, 3
Mecostiboides 3
Mecostibus 1, 2, 3
Microusambilla 1,' 3, 10, 12
modicicrus 1, 2, 11, 12, 13, 16
montanum 2, 13, 14, 15, 16, 18, 19
mweruensis 2, 4, 5, 6, 7, 8, 9

nguruense 14, 15, 16, 17, 18
Nyassacris 1, 3

olivacea, Usambilla, 20, 23, 24, 25, 26, 27

olivacea, Lentula 37

oraria 21, 22, 24, 25, 26, 28, 32, 35

Paralentula 3
patula, Pinus 1, 13
peruviana, Physalis 8
Plagiotriptus 1

Qachasia 5

Rhainopomma 1, 4, 12, 35

Rehnula 20

robertsoni 2, 4, 6, 7, 8, 9, 10

sagonai 20, 21, 22, 23, 24, 26, 30, 31, 34, 35

Serpusia 13

Shelfordites 3

steppia, Bidens 8

stuhlmanni, Pycnostachys (?) 8

Swaziacris 4

Sygrus 3, 12

turgidicrus 20, 21, 22, 23, 24, 25, 26, 28, 29, 30

usambaricum 13, 14, 15, 16, 17, 18, 19
Usambilla 1, 2, 4, 5, 10, 13, 17, 20

verticillaris, Hypoestes 8
veseyi 2, 4, 5, 6, 7, 8, 9, 10

wapugu 14, 15, 16, 17, 19

38

British Museum (Natural History)
1881-1981

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A revision of the genus Usambilla Sjostedt (Orthoptera : Acridoidea) and its allies.

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The Asian, Australasian and Pacific Paraboloponinae (Homoptera : Cicadellidae).
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Paraboloponinae

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A taxonomic revision with a key to all the known genera
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M. D. Webb

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Vol 43 No 2 pp 39-76
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London SW7 5BD Issued 24 September 1 98 1

The Asian, Australasian and Pacific Paraboloponinae
(Homoptera : Cicadel lidae)

A taxonomic revision with a key to all the known genera
of the subfamily

M. D. Webb

i«*

Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Contents

Synopsis 39

Introduction and historical review 39

Acknowledgements 40

Abbreviations and depositories 40

Paraboloponinae Ishihara 41

Key to the genera of Paraboloponinae 41

Parabolopona Matsumura 42

Key to species of Parabolopona (males only) 43

Favintiga gen. n 47

Dryadomorpha Kirkaldy . 49

Key to Asian, Australasian and Pacific species of Dryadomorpha (males only) . 50

Rhutelorbus gen. n 56

Parohinka gen. n. .

Key to species of Parohinka
Males ....
Females
Karoseefa gen. n. .

Key to species of Karoseefa
Oceanopona Linnavuori

References
Index .

57
59
59
59
70
71
73
75
76

Synopsis

The subfamily Paraboloponinae is redefined and a key is provided to the nine genera recognized, of which
four are new. The Asian, Australasian and Pacific genera are fully revised for these regions, with keys to the
25 species (18 new) recognized. Four new generic synonymies, ten new specific synonymies and six new
combinations are established. Seven lectotypes and one neotype are newly designated.

Introduction and historical review

The family Paraboloponidae was erected by Ishihara in 1953 as a subdivision of the Cicadelloi-
dea (now Paraboloponinae and Cicadellidae respectively) for the Japanese genus Parabolopona
Matsumura, containing two species, P. guttata (Uhler) and P. camphorae Matsumura. In Ishi-
hara's description of the subfamily he distinguished it from others in Japan by the cylindrical
form, conically produced vertex and long antennae. In 1960 Linnavuori listed the group as a tribe
of the Deltocephalinae and described a new genus and species, Oceanopona croceipennis, from the
Caroline Is., and in 1974 Eyles & Linnavuori raised the group to subfamily level and included
Calotettix Osborn, containing metrosideri Osborn and metrosideri var. tincta Osborn, from the
Marquesas Is., and a new species, lais, from the Cook Is. In 1975 Hamilton added the Holarctic
genera Stymphylus Stal, containing rubrolineata Stal and modestus Linnavuori (transferred to the

Bull. Br. Mus. not. Hist. (Ent.) 43 (2): 39-76 Issued 24 September 1981

40 M. D. WEBB

Deltocephalinae by Linnavuori in 1978), Yakunopona Ishihara, with one species, yakushimensis
Ishihara, from Japan, and Zizyphoides Distant, containing indicus Distant, fraternus Distant and
punctatus Rao from India and quadricornis Linnavuori from Africa. In addition Hamilton drew
further attention to the long, dorsally situated antennae as a means of identifying the subfamily.
In 1978 Linnavuori revised the Paraboloponinae from the Ethiopian region and redefined the
group as having falcate anterior tentorial branches, dorsally situated antennae, deep antennal
pits delimited by a relatively distinct dorsal ledge and the ocelli not visible from above. He added
Dryadomorpha Kirkaldy, containing pallida Kirkaldy, lotophagorum Kirkaldy and viridia Osborn
from Australia, Fiji and the Marquesas Is. respectively, and Stenomiella Evans, containing one
species, viridis Evans, from Africa; he also described a new genus Odmiella for Stenomiella falcata
Linnavuori, from Africa. In addition, the genus Paganalia Distant with one species, virescens
Distant, from the Seychelles, was transferred to the subfamily as a senior synonym of Zizyphoides
and Rhombopsana Metcalf, the latter a replacement name for Rhombopsis Haupt and containing
virens (Haupt) from Palestine and chatterjeei (Singh-Pruthi) and viridis (Singh-Pruthi) from India.
The species Z. fraternus was transferred to Stirellus Osborn & Ball (Deltocephalinae), a new
species of Paganalia from Africa (anacryon) was described and Platymetopius antennalis Lind-
berg, from the Canary Is., was synonymized with Paganalia virens. The latter species was synony-
mized with Paganalia virescens by Webb (1980).

Whilst identifying Paraboloponinae from material in the British Museum (Natural History)
and elsewhere I discovered four new genera and 18 new species from Asia, Australasia and the
Pacific and found that many previous descriptions were inadequate. In addition, the species
Parabolopona camphorae, Dryadomorpha lotophagorum and Muirella longiseta Melichar, the
latter previously included in the Coelidiinae, were found to belong to other genera. A number of
generic and specific synonymies were also discovered.

The subfamily characters given by previous workers are for the most part confirmed by the
present study, although of those listed by Linnavuori in 1978 (see above) the antennal ledge is
sometimes absent and the ocelli are often visible from above. The male genitalia of the group are
similar to those of the Deltocephalinae with Y-shaped connective, but those of Parabolopona are
somewhat distinct from the remaining genera in having a long membranous connection between
the connective and the aedeagus, the apex of the connective extended posteriorly and the basal
apodeme of the aedeagus, in some species, horizontal and compressed dorsoventrally. The genus
Favintiga is unusual within the Cicadellids in having a ventroapical process on the connective.

The aim of the present paper is to revise the known genera of Paraboloponinae, other than
Odmiella and Stenomiella from Africa (revised by Linnavuori in 1978), to describe new genera and
species from the Asian, Australasian and Pacific regions, to redescribe the subfamily and to
provide a key to the known genera.

Acknowledgements

For the loan of material in their care I would like to thank the following: Dr P. H. Arnaud, CAS,
San Francisco; Dr M. Boulard, MNHN, Paris; Dr O. B. Chhotoni, ZSI, Calcutta; Dr J. Dlabola,
NM, Prague; Dr P. H. van Doesburg, RNH, Leiden; Dr K. G. A. Hamilton, CNC, Ontario; Dr
Y. Hirashima, ELKU, Fukuoka; Dr J. P. Kramer, USNM, Washington; Dr M. Meinander,
ZMU, Helsinki; Dr G. M. Nishida, BPBM, Honolulu; Dr P. K. Sen-Sarma, FRI, Dehra Dun; Dr
S. Takagi, EIHU, Sapporo; Dr T. Weir, ANIC, Canberra; Dr K. A. J. Wise, IM, Auckland. I
would also like to thank Dr J. Evans for the loan of material in his private collection, Mrs G.
Nakahashi and Professor J. Medler for sorting material from the Bishop Museum collection, Dr
T. Ishihara and Dr C. A. Viraktamath for their helpful communications and Dr W. J. Knight for
the information contained in his unpublished check list on the Pacific Cicadellidae.

Abbreviations of depositories

The specimens studied in the course of this work are deposited in the various institutions and
private collections whose names are abbreviated in the text as follows: The South Australian
Museum, Adelaide, South Australia (SAM, Adelaide); Auckland Institute and Museum, Auck-

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 41

land, New Zealand (IM, Auckland); Department of Scientific and Industrial Research, Auckland,
New Zealand (DSIR, Auckland); Zoological Survey of India, Calcutta, India (ZSI, Calcutta);
[Australian National Insect Collection,] C.S.I. R.O., Canberra, Australia (ANIC, Canberra); The
Ohio State University, Columbus, Ohio, U.S.A. (OSU, Columbus); Zoologisk Museum, Copen-
hagen, Denmark (ZM, Copenhagen); Forest Research Institute, Dehra Dun (U.P.), India (FRI,
Dehra Dun); Entomological Laboratory, Kyushu University, Fukuoka, Japan (ELKU, Fu-
kuoka); Zoological Museum of the University, Helsinki, Finland (ZMU, Helsinki); Bernice P.
Bishop Museum, Honolulu, Hawaii, U.S.A. (BPBM, Honolulu); private collection of Professor
Dr H. J. Muller, Jena, D.D.R. (HJM, Jena); Rijkusmuseum van Natuurlijke Historic, Leiden,
Netherlands (RNH, Leiden); British Museum (Natural History), London, United Kingdom
(BMNH, London); [Canadian National Collection,] Ottawa, Ontario, Canada (CNC, Ontario);
Museum National d'Historie Naturelle, Paris, France (MNHN, Paris); Narodni Muzeum,
Prague, Czechoslovakia (NM, Prague); California Academy of Sciences, San Francisco, Califor-
nia, U.S.A. (CAS, San Francisco); Entomological Institute, Hokkaido University, Sapporo,
Japan (EIHU, Sapporo); private collection of Dr J. W. Evans, Sydney, Australia (JWE, Sydney);
[U.S. National Museum,] National Museum of Natural History, Washington, U.S.A. (USNM,
Washington).

PARABOLOPONINAE Ishihara

Paraboloponidae Ishihara, 1953: 20. Type-genus: Parabolopona Matsumura.
Paraboloponini; Linnavuori, 1960: 299.
Paraboloponinae; Eyles & Linnavuori, 1974: 39.

Yellow, greenish yellow or brownish yellow, often apex of clavus and claval veins with a small brown spot.

Head as wide or wider than pronotum; anterior margin rounded or rim-like, transversely striate, ocelli on
margin distant from eyes, anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly
produced with fine longitudinal striations. Face as wide or wider than long, shagreen ; antennae very long,
arising near dorsal corners of eyes; antennal pits deep; antennal ledges slight or absent; clypeus with lateral
margins constricted near antennae; clypellus elongate, usually with sides concave, rarely with sides parallel;
lora large. Pronotum with sides very short to moderately long, with or without a carina; transversely striate,
with anterior region rugose or shagreen. Scutellum shagreen or shagreen and obscurely rugose posteriorly.
Forewing with three subapical cells, the first subapical cell open and the second and third closed ; subcostal
region usually with a few veinlets near to fifth apical cell. Fore tibia with setal arrangement 1 : 4.

Male genitalia with pygophore lobes long with several long spine-like setae. Xth segment short to long,
without processes. Valve triangulate. Subgenital plate elongate, triangulate with short to long fine marginal
setae dorsally. Connective Y-shaped with stem short to long, arms short. Style with lateral lobe and apical
process short to moderately long, with a few sensory papilla and sometimes setae adjacent preapical lobe;
basal apophyses weak to strongly developed. Aedeagus usually closely attached to connective; shaft cylin-
drical, narrow, tapered to apex, usually symmetrical, processes usually at or near apex sometimes basal;
basal apodeme usually vertical, rarely horizontal.

Female genitalia with second valvulae elongate, usually slightly expanded distally with a short to long
dorsal sclerotized region, with or without a dorsal prominence.

BIOLOGY. The few recorded host plants of Paraboloponinae are shrubs and small trees. Favintiga
camphorae is found on Cinnamomum camphora Nees & Ebermaier, Dryadomorpha pallida on
Zizyphus jujuba Miller and Dryadomorpha metrosideri on Glochidion ramiflorum J. R. & G.
Forster, Rapanea sp., Reynoldsia sp. and Weinmannia parviflora G. Forster.

DISTRIBUTION. The subfamily is confined to the Old World where it is found mainly in Asia and
Australasia but also extends into the Pacific and the Ethiopian region.

Key to the genera of Paraboloponinae

1 Side margins of pronotum carinate, moderately long (Fig. 1). Setal arrangement at apex of hind

femur 2 + 2+1 or 2 + 2 + 0 2

Side margins of pronotum not carinate, short (Fig. 41). Setal arrangement at apex of hind femur

2+1 + 1 or 2+1+0. 4

42 M. D. WEBB

2 Vertex with medial length approximately twice length next to eye. 12-15 setae in fore femur

series (Fig. 3). Setal arrangement at apex of hind femur 2 + 2+ 1. (Asia as far south as Nepal

to the Philippines: Luzon) 3

Vertex with medial length approximately four times length next to eyes. 3-4 setae in fore femur
series. Setal arrangement at apex of hind femur 2 + 2 + 0. (Africa) . . ODMIELLA Linnavuori

3 Dorsum yellow or yellow tinged with green. Fore margin of head rim-like (Fig. 5). Vertex

shagreen PARABOLOPONA Matsumura (p. 42)

Dorsum brownish yellow. Fore margin of head rounded (Fig. 33). Vertex finely longitudinally
striate . .FAVINTIGA gen. n. (p. 47)

4 Vertex shagreen and obscurely rugose. Setal arrangement at apex of hind femur 2+1+0

OCEANOPONA Linnavuori (p. 73)
Vertex longitudinally striate or rugose. Setal arrangement at apex of hind femur 2+1 + 1 . 5

5 Length approximately 10-0 mm. Male pygophore lobes with a long process; subgenital plates

shorter than pygophore. (Africa) STENOMIELLA Evans

Length not exceeding 8-7 mm. Male pygophore lobes without a process; subgenital plates
longer than pygophore (Fig. 53). (Africa and Oriental region) 6

6 Clypellus with sides parallel (Fig. 151). 10 setae in fore femur series . KAROSEEFA gen. n. (p. 70)
Clypellus with sides concave (Fig. 42). 3-7 setae in fore femur series 7

7 Vertex and pronotum rugose. Lateral margins of face adjacent to eyes visible dorsally (Fig. 68)

RHUTELORBUS gen. n. (p. 56)
Vertex and pronotum longitudinally striate. Lateral margins of face not visible dorsally (Fig. 41) 8

8 Vertex without pale patches. Female genitalia with posterior margin of pregenital sternite with a

small protuberance each side of midline (Fig. 51); dorsal margin of second valvulae with an
anterior prominence (Fig. 56). Male genitalia with aedeagus symmetrical

DRYADOMORPHA Kirkaldy (p. 49)

Vertex with or without pale patches. Female genitalia with posterior margin of pregenital
sternite without a protuberance each side of midline; dorsal margin of second valvulae
without an anterior prominence (Fig. 133). Male genitalia with aedeagus asymmetrical

PAROHINKA gen. n. (p. 51)

PARABOLOPONA Matsumura

Parabolopona Matsumura 1912: 288. Type-species : Parabolocratus guttatus Uhler, by original designation.

Yellow to greenish yellow ; forewings with a small brown spot on apex of clavus, on apex of veins of clavus
and apical cells and a variable brown spot on first m-cu cross vein and base of inner vein of second subapical
cell.

Head as wide as pronotum; anterior margin rim-like, carinate; ocelli on margin, distant from eyes, visible
from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced,
medial length approximately twice length next to eyes; sides slightly convex or slightly angularly rounded;
apex narrowly angularly rounded; shagreen, transversely striate anteriorly. Face slightly wider than long,
shagreen; upper margin depressed medially with a few transverse striations; face in profile more or less
straight ; clypeus moderately long and narrow, lateral margins constricted near antennae; clypellus elongate,
expanded apically ; transclypeal suture visible ; lora large ; antennal pit deep with inner margin more or less
angularly rounded to clypeus; antennal ledge slight; antennae very long, when recurved extending to near
apex of clavus. Pronotum approximately twice as wide as long, side margins moderately long, carinate;
irregularly and transversely striate, shagreen anteriorly. Scutellum approximately equal in length to prono-
tum, shagreen, obscurely rugose posteriorly. Fore wing with three subapical cells, first subapical cell open,
second and third subapical cells closed. Fore tibia with dorsal setal arrangement 1:4; fore femur with a
series of 12-14 fine setae distally on anterior surface; hind femur with apical setal formula 2 + 2+1 with the
proximal and more dorsal of the middle setae slightly narrower than others.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore straight in dorsal aspect, with or without an apodeme
on each side; pygophore lobes with several macrosetae and numerous short fine setae. Xth segment moder-
ately long, cylindrical. Valve triangular. Subgenital plate moderately long, triangular, apical region digitate
and lightly sclerotized; ventral surface with several short setae; outer margin of dorsal surface with a few
moderately long fine setae on basal lobe and usually towards apex of plate. Style moderately long with basal
apophysis and lateral lobe prominent; apical process moderately long, curved ventrally and tapered to apex
or with apex foot-like, crenulate dorsally; region adjacent preapical lobe with a few sensory papilla ventrally

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 43

and a few short fine setae dorsally. Connective Y-shaped with stem produced posteriorly, not articulated
with aedeagus at apex but with a long membranous connection from approximately midlength of stem;
arms short. Aedeagus with shaft straight or curved posteroventrally, short with a pair of apical processes;
gonopore situated at apex on posterior surface; basal apodeme large, either vertical and compressed
anteroposteriorly or horizontal around base of shaft and compressed dorsoventrally.

Female genitalia with second valvulae united at midlength (arrowed in Fig. 14) and slightly expanded
distally, without a basal prominence; dorsal teeth very fine, extending approximately one-quarter distance
from apex to base of valvulae; dorsal sclerotized region short to long.

REMARKS. This genus is similar to Favintiga with the lateral margins of the pronotum moderately
long and carinate and the hind femur with a setal formula 2 + 2 + 1 ; it differs, however, in the
more rim-like fore margin of the head and the shagreen rather than longitudinally striate vertex.
In the male genitalia the horizontal and dorsoventrally compressed basal apodeme of the
aedeagus in some species, the produced stem of the connective and the distant relationship of the
aedeagus and connective to each other are unique within the subfamily.

DISTRIBUTION. Asia as far south as Nepal to the Philippines (Luzon).

Key to species of Parabolopona (males only)

1 Aedeagus with a pair of apical processes; stem of connective without setae 2

Aedeagus with a pair of basal processes; stem of connective with setae. (Philippines)

luzonensis sp. n. (p. 46)

2 Aedeagus with apical processes directed away from base of shaft in lateral aspect, gonopore small 3
Aedeagus with apical processes directed towards base of shaft in lateral aspect; gonopore large

(Figs 16, 17) ishihari sp. n. (p. 45)

3 Aedeagal shaft strongly curved (Fig. 8); connective with apex upturned and expanded laterally

(Figs 7, 13). (Japan and Taiwan) guttata (Uhler) (p. 43)

Aedeagal shaft weakly curved (Fig. 20); connective with apex straight and narrow. (China)

chinensis sp. n. (p. 45)

Parabolopona guttata (Uhler)
(Figs 1-14)

Parabolocratus guttatus Uhler, 1896: 291. LECTOTYPE & JAPAN (USNM, Washington), here designated

[examined].
Parabolopona guttata (Uhler) Matsumura, 1912: 288.

Length: cJ, 6-6-7-0 mm, mean 6-8 mm; $, 7-0-8-0 mm, mean 7-5 mm.

Colour and external characters as in generic description.

Male genitalia with pygophore lobes angularly rounded posteriorly; anterior margin of pygophore with a
prominent apodeme on each side. Connective with apex upturned and expanded laterally. Style with apical
process tapered to apex. Aedeagus with shaft elongate, curved posteroventrally and continued ventrally as a
pair of elongate diverging processes; gonopore small; basal apodeme horizontal, compressed dorsoventrally
around base of shaft (Fig. 12).

Female genitalia with posterior margin of pregenital sternite sinuate; second valvulae with dorsal scler-
otized region short.

REMARKS. The male genitalia of guttata are similar to those of chinensis but the pygophore lobes
are broader, the apex of the connective is upturned and expanded laterally and the aedeagal shaft
is strongly curved with the apical processes expanded subapically without a small lateral pro-
tuberance basally.

DISTRIBUTION. Japan and Taiwan.

MATERIAL EXAMINED

Parabolocratus guttatus Uhler, lectotype cJ, Japan: Gifuyama, 7.vii.l888 [in Japanese] (USNM, Wash-
ington).

Japan: numerous specimens from Honshu and Kyushu (BMNH, London; EIHU, Sapporo; ELKU,
Fukuoka); 1 $, Gifuyama, 7.vii.l888 [in Japanese] (USNM, Washington) (paralectotypes of Parabolocratus
guttatus Uhler). Taiwan: 1 <J, 1 ?, Tattaka, 16, 19.viii.1921; 3 9, Hassenzan, Taichu-shu, Reimei, 13-
14.vii.1932 (ELKU, Fukuoka).

44

M. D. WEBB

Figs 1-14 Parabolopona guttata. 1, head and thorax, dorsal view; 2, face; 3, left fore leg, anterior view;
4, left anterior tentorial branch, lateral view; 5, head and thorax, lateral view; 6, $ pregenital
segments, ventral view; 7, connective, lateral view; 8, aedeagus, lateral view; 9, left style, ventral view;
10, cJ genital capsule, lateral view; 11, aedeagus, dorsal view; 12, left subgenital plate and style and
connective, dorsal view; 13, connective, dorsal view; 14, second valvulae, lateral view.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

45

Parabolopona ishihari sp. n.

(Figs 15-19)

Length: ^,6-6 mm; $,7-5 mm.

Colour and external characters as in generic description.

Male genitalia with pygophore lobes acute posteriorly; anterior margin without an apodeme on each side.
Connective with basal stem straight and narrow throughout length. Style with apical process tapered to
apex. Aedeagus with shaft short and robust, more or less straight, terminating in a pair of moderately long
ventrally directed processes; gonopore large; basal apodeme horizontal, compressed dorsoventrally around
base of shaft (Figs 16,17).

Female genitalia as in guttata.

REMARKS. The male genitalia of ishihari are similar to those of guttata but the pygophore lobes
are more acute posteriorly, the apex of the connective is straight and narrow and the aedeagus
has the processes directed towards the base of the shaft in lateral aspect and the gonopore large.

DISTRIBUTION. Japan.

MATERIAL EXAMINED

Holotype J, Japan: Northern Honshu, Towada, vii.1905 (EIHU, Sapporo).
Paratypes. Japan: 1 <$, 1 $, same data as holotype (BMNH, London; EIHU, Sapporo).

Parabolopona chinensis sp. n.

(Figs 20-21)

Length : <J, 6-4 mm.

Colour and external characters as in generic description.

Male genitalia with pygophore and connective similar to ishihari (Figs 15, 18) but pygophore with an
apodeme on each side anteriorly and posterior lobes less acute; remaining structures similar to guttata but

15

20

Figs 15-21 Parabolopona species. 15-19. P. ishihari. (15) ^ pygophore, lateral view; (16, 17) aedeagus,
lateral and posterior views; (18, 19) connective, dorsal and lateral views. 20, 21, P. chinensis. (20)
aedeagus, lateral view; (21) apex of aedeagus, dorsal view.

46

M. D. WEBB

connective straight and narrow apically and aedeagal shaft only slightly curved with apical processes evenly
tapered from base to apex with a small lateral protuberance basally.
Female genitalia unknown.

REMARKS. This species is similar to guttata but differs in the male genitalia as noted above. From
ishihari with a similarly shaped pygophore and connective (see above), chinensis differs in having
the aedeagal processes directed away from the shaft in lateral aspect.

DISTRIBUTION. Central China.

MATERIAL EXAMINED

Holotype cJ, China: Hubei-Sichuan border, trail between Mo-Tai-Chi and Sang-Hou-Ken, 19.vii.1948
(Gressitt & Djou) (CAS, San Francisco).

Parabolopona luzonensis sp. n.

(Figs 22-27)
[Parabolocratus guttatus Uhler; Merino, 1936: 364. Misidentification.]

Length : (J, 6-7-7-3 mm, mean 7-0 mm; 9, 7-3-8-2 mm, mean 8-0 mm.

Colour and external characters as in generic description.

Male genitalia with pygophore lobes narrowly rounded posteriorly; anterior margin of pygophore with-
out apodemes. Connective with stem nearly straight, tapered to acute apex; distal region with numerous
short stout setae dorsally. Style with apical process foot-like apically in lateral aspect. Aedeagus with shaft
short and robust, curved dorsally, apex with a slight lamellate expansion arising from anterior margin on
each side; a pair of moderately long, dorsally directed processes, arising basally from posterior margin;
gonopore moderately large ; basal apodeme vertical, compressed anteroposteriorly.

Female genitalia with posterior margin of pregenital sternite produced medially; second valvulae with
dorsal sclerotized region elongate.

23

27

Figs 22-27 Parabolopona luzonensis. 22, $ genital capsule, lateral view; 23, 9 pregenital sternite; 24,
connective, dorsal view; 25, 26, aedeagus, lateral and posterior views; 27, apex of style, lateral view.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 47

REMARKS. This species can be distinguished from other members of the genus by the foot-like
apex of the style and the aedeagus with the processes arising basally rather than apically and the
basal apodeme being vertical rather than horizontal. The female genitalia can be distinguished by
the medially produced posterior margin of the pregenital sternite and the elongate dorsal scler-
otized region of the second valvulae.

DISTRIBUTION. Philippines (Luzon).

MATERIAL EXAMINED

Holotype J, Philippines: Luzon, Baguio, Benguet (Baker) (USNM, Washington).

Paratypes. Philippines: 10 £, 19 9, same data as holotype (1 (J, 1 $ in BMNH, London; remainder in
USNM, Washington); 1 <J, Luzon, Heights Plane (OSU, Columbus).

FA VINT1GA gen. n.

Type-species: Parabolopona camphor -ae Matsumura, 1912.

Brownish yellow dorsally, pale yellow ventrally; fore wings with a small brown spot near mid length of
subcostal region, on apex of clavus and on apex of veins of clavus, apical cells and additional vein in
subcostal region.

Head as wide as pronotum; anterior margin angularly rounded in profile, transversely striate, becoming
carinate and rim-like medially; ocelli on margin, distant from eyes, visible from above; anterior tentorial
branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approximately twice
length next to eyes, sides slightly convex; apex moderately broadly rounded; finely longitudinally striate,
transversely striate anteriorly. Face slightly wider than long, more or less straight in profile; shagreen;
clypeus moderately long, narrow, lateral margins constricted near antennae; clypellus elongate, expanded
apically; transclypeal suture visible; lora large; antennal pit deep with inner margin obliquely inclined to
clypeus; antennal ledge slight; antennae very long, extending to near apex of clavus. Pronotum approxi-
mately twice as wide as long, sides moderately long, carinate ; finely and transversely striate, shagreen and
more or less smooth anteriorly, Scutellum approximately equal in length to pronotum, shagreen, obscurely
rugose posteriorly. Fore wings with three subapical cells, first subapical cell open, second and third subapi-
cal cells closed; an additional veinlet in subcostal region near to fifth apical cell. Fore tibia with dorsal setal
arrangement 1:4; fore femur with a series of 15 fine setae distally on anterior surface; hind femur with apical
setal formula 2+1 + 1 with the proximal and more dorsal of the middle setae slightly narrower than
others.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes
with several macrosetae and short fine setae. Xth segment moderately long, cylindrical. Valve triangulate.
Subgenital plates moderately long, distal half digitate and lightly sclerotized; dorsal surface with a few
moderatly long fine setae on basal lobe and a few very short setae distally, Style moderately long with basal
apophyses and lateral lobe prominent; apical process moderately long, curved ventrally and tapered api-
cally, crenulate medially; a few sensory papilla medially adjacent lateral lobe. Connective Y-shaped; stem
long, lateral margins keel-like dorsally; a bifurcate process distally on ventral surface; arms short. Aedeagus
large, laterally compressed with an elongate preatrium; shaft curved dorsally and tapered apically with a
pair of basal processes; gonopore situated at apex on posterior surface, elongate; basal apodeme elongate.

Female genitalia with second valvulae united at first dorsal tooth (arrowed in Fig. 38), narrow throughout
length in lateral aspect with a slight dorsoanterior prominence; dorsal teeth very fine, extended over
approximately distal third of valvulae; dorsal sclerified region elongate.

REMARKS. This genus is similar externally to Parabolopona (see remarks under that genus) but the
long preatrium of the aedeagus and the processes on the connective are unique within the
subfamily.

DISTRIBUTION. Japan and Amama-Oshima I. (south of Japan).

48

M. D. WEBB

Figs 28-40 Favintiga camphor ae. 28, head and thorax, dorsal view; 29, face; 30, 9 pregenital segments,
ventral view; 31, fore wing; 32, left style, ventral view; 33, head and thorax, lateral view; 34, left
subgenital plate and apex of left style, dorsal view; 35, 36, aedeagus, lateral and posterior views; 31,3
genital capsule, lateral view; 38, second valvulae, lateral view; 39, 40, connective, lateral and dorsal
views.

Favintiga camphorae (Matsumura) comb. n.
(Figs 28^0)

Parabolopona camphorae Matsumura, 1912: 288. LECTOTYPE 9, JAPAN (EIHU, Sapporo), here designated
[examined].

Length: <$, 6 mm; 9, 6-3-7-0 mm, mean 6-6 mm.

Colour and external characters as in generic description.

Male genitalia as in generic description with apical process of style with a small subapical tooth on medial
surface and connective with each branch of ventral process elongate, tapered to apex and curved ventrome-
dially. Aedeagus with shaft elongate; processes arising against posterior margin on each side, extended
dorsally close to shaft and terminating a little before its apex, evenly tapered from base to apex.

Female genitalia with posterior margin of pregenital sternite convex; second valvulae as in generic
description.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 49

REMARKS. This species is distinguishable by its brownish yellow colour dorsally and its distinctive
male genitalia as noted above.

DISTRIBUTION. Japan and Amami-Oshima I. (south of Japan).

MATERIAL EXAMINED

Parabolopona camphorae Matsumura, lectotype 9, Japan: S. Kyushu, Kagoshima, 10.vii.1903 (EIHU,
Sapporo).

Japan: 1 $, same data as lectotype (paralectotype of Parabolopona camphorae Matsumura); !<£,!$,
Honshu, Gifuyama, 7.vii.l888 (USNM, Washington); 3 9, Amami-Oshima I., Yakkachi, Sumiyo-mura,
17-18.vii.1933 (Esaki & Yasumatsu) (ELKU, Fukuoka, BMNH, London).

DRYADOMORPHA Kirkaldy

Dryadomorpha Kirkaldy, 1906: 335. Type-species: Dryadomorpha pallida Kirkaldy, by monotypy.

Paganalia Distant, 1917: 314. Type-species : Paganalia virescens Distant, by monotypy. Syn. n.

Zizyphoides Distant, 1918: 73. Type-species: Zizyphoides indicus Distant, by original designation. [Synony-
mized by Linnavuori 1978: 459.]

Rhombopsis Haupt, 1927: 22. Type-species: Rhombopsis virens Haupt, by monotypy. [Homonym of Rhom-
bopsis Gardner, 1916: 456.]

Calotettix Osborn, 1934: 247. Type-species: Calotettix metrosideri Osborn, by original designation. [Hom-
onym of Calotettix Bruner, 1908: 309.]

Yakunopona Ishihara, 1954: 12. Type-species: Yakunopona yakushimensis Ishihara, by original designation.
Syn. n.

Rhombopsana Metcalf, 1967: 229. [Replacement name for Rhombopsis Haupt.] [Synonymized by Linna-
vuori, 1978: 459.]

Osbornitettix Metcalf, 1967: 229. [Replacement name for Calotettix Osborn.] Syn. n.

Khamiria Dlabola, 1979: 252. Type-species: Khamiria mangrovecola Dlabola, by original designation.
Syn. n.

Yellow, greenish yellow or stramineous; fore wings with either a small brown spot at apex of both clavus
and claval veins or inner margin of clavus bordered with brown. Legs spotted with brown.

Head wider than pronotum; anterior margin angularly or acutely rounded in profile, transversely striate,
sometimes becoming carinate and rim-like in longer headed forms; ocelli on margin distant from eyes, not
or slightly visible from above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangu-
larly produced, medial length 1-5-3-0 times length next to eyes, sides slightly convex to concave, apex fairly
broadly angularly rounded to acute in longer headed forms; with a longitudinal depression; finely longitudi-
nally striate, transversely striate anteriorly. Face elongate to only slightly longer than wide, shagreen; upper
margin slightly depressed each side of mid line in longer headed forms forming a faintly striate longitudinal
keel medially; face in profile convex to more or less straight, concave anteriorly in longer headed forms;
clypeus elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; trans-
clypeal suture distinct or indistinct; lora large; antennal pit deep with inner margin angularly rounded to
clypeus, sometimes faintly rim-like and nearly carinate; antennal ledge very slight; antennae very long,
extending to beyond apex of clavus when recurved. Pronotum approximately twice as wide as long, sides
very short, without a carina; finely and transversely striate, obscurely rugose anteriorly. Scutellum approxi-
mately equal in length to pronotum, shagreen with posterior region obscurely rugose. Fore wing with three
subapical cells, first subapical cell open, second and third subapical cells closed; one or two additional
veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur
with a series of seven setae distally on anterior surface ; hind femur with apical setal formula 2+1-1-1 with
first proximal seta slender.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes
with an oblique internal ledge terminating at ventroposterior margin with a darkly pigmented area, pygo-
phore lobes with several macrosetae and short to moderately long fine setae. Xth segment moderately long,
compressed dorsoventrally. Valve triangulate. Subgenital plates elongate, triangular; ventral surface of
lateral lobe with a more heavily sclerotized region apically; outer margin with numerous long fine setae on
dorsal and ventral surfaces; apex with a few short stout setae. Style moderately long with basal apophyses
prominent ; apical process moderately long, curved ventrally and tapered to acute or rounded apex, crenu-
late distally; a few sensory papilla ventrally, adjacent lateral lobe. Connective Y-shaped, stem short to
moderately long with lateral margins keel-like dorsally; arms short. Aedeagus with shaft elongate, curved

50 M. D. WEBB

dorsally and tapered to apex with two or four apical, dorsally directed processes ; gonopore small, apical on
posterior surface; basal apodeme moderately long and narrow.

Female genitalia with posterior margin of pregenital sternite with a small protuberance each side of mid
line; second valvulae united at first dorsal tooth (arrowed in Fig. 56), slightly expanded distally, fairly robust
with a dorsoanterior prominence; dorsal teeth robust, unaligned, extending to near mid length of valvulae;
dorsal sclerotized region moderately long.

REMARKS. This genus is represented in the southern Palaearctic region by Dryadomorpha pallida
Kirkaldy (also present in Africa, Asia and Australasia and described below), and D. mangrovecola
(Dlabola) comb. n. Also present in Africa are D. anacryon (Linnavuori) comb. n. and D. quad-
ricornis (Linnavuori) comb. n. (both species adequately described by Linnavuori, 1978: 462). Two
species incorrectly described in Zizyphoides ( = Dryadomorpha) are Z. fraternus Distant belonging
to the genus Stirellus Osborn & Ball (Linnavuori, 1978: 460) and Z. punctatus Rao syn. n. of
Mahalana lugubris Distant.

Externally Dryadomorpha is almost identical to Rhutelorbus and Parohinka but differs in
lacking the pale patches on the head (usually present in Parohinka and sometimes in Rhutel-
orbus), the striate vertex and pronotum (rugose in Rhutelorbus), the lateral margins of the face
adjacent to the eyes not being visible dorsally (just visible in Rhutelorbus) and the posterior
margin of the female pregenital sternite being transverse (posterior corners produced in Rhutel-
orbus and Parohinka or broadly V- or U-shaped in Parohinka). The male genitalia are similar to
Karoseefa but have the anterior margin of the pygophore straight dorsally and without apo-
demes. In the female genitalia the second valvulae are similar to those of Parohinka but with a
dorsoanterior prominence present.

DISTRIBUTION. Ethiopian and southern Palaearctic region, Asia, Australasia and the Pacific.

Key to Asian, Australasian and Pacific species of Dryadomorpha (males only)

1 Aedeagus with apical processes strongly divergent (Fig. 52) 2

Aedeagus with apical processes weakly divergent (Fig. 62) 3

2 Aedeagal processes slightly expanded at mid length in posterior aspect (Fig. 52); fore wings with a

small brown spot at apex of both clavus and claval veins (Fig. 47). (Ethiopian, southern
Palaearctic and Oriental regions as far south as Singapore and Java and northern and

north-eastern Australia) pallida Kirkaldy (p. 50)

Aedeagal processes narrow throughout length in posterior aspect (Fig. 60); fore wings without a
small brown spot at apex of both clavus and claval veins, sometimes inner margin of clavus
brown (Fig. 58). (Marquesas Is., Cook Is. and northern Australia) . . metrosideri (Osborn) (p. 53)

3 Aedeagal shaft robust (Fig. 63); style with apical process tapered to apex (Fig. 61). (Malaya,

Borneo (Sarawak and Sabah)) robustipenis sp. n. (p. 54)

Aedeagal shaft slender (Fig. 66); style with apical process rounded or foot-like apically (Figs 64,

65). (Borneo (Sarawak and Sabah)), New Guinea and Vanuatu . . . pacifica sp. n. (p. 55)

Dryadomorpha pallida Kirkaldy
(Figs 41-56)

Dryadomorpha pallida Kirkaldy, 1904: 336. Holotype?, AUSTRALIA, (BPBM, Honolulu) [examined].

Paganalia virescens Distant, 1917: 314. Lectotype $, SEYCHELLES (BMNH, London), designated by Webb
(1980: 848) [examined]. Syn. n.

Zizyphoides indicus Distant, 1918: 73. LECTOTYPE ?, INDIA (BMNH, London), here designated [exam-
ined]. Syn. n.

Rhombopsis virens Haupt, 1927: 23. Lectotype cJ, ISRAEL (HIM, Jena), designated by Webb (1980: 848)
[examined]. Syn. n.

Rhombopsis viridis Singh-Pruthi, 1930: 34. LECTOTYPE <$, INDIA (ZSI, Calcutta), here designated [exam-
ined]. Syn. n.

Rhombopsis chatterjeei Singh-Pruthi, 1934: 26. LECTOTYPE <3, INDIA (FRI, Dehra Dun), here designated
[examined]. Syn. n.

Platymetopius antennalis Lindberg, 1958: 181. Holotype <$, CAPE VERDE Is. (ZMU, Helsinki) [examined].
Syn. n.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

51

Figs 41-56 Dryadomorpha pallida. 41, head and thorax, dorsal view; 42, face; 43, head and thorax,
lateral view; 44, head and pronotum, dorsal view; 45, head and thorax, lateral view; 46, head and
pronotum, dorsal view; 47, fore wing; 48, face; 49, aedeagus, lateral view; 50, left style, ventral view;
51, apex of $ abdomen, ventral view; 52, aedeagus, posterior view; 53, <$ genital capsule, lateral view;
54, 55, connective, lateral and dorsal views; 56, second valvulae, lateral view.

52 M. D. WEBB

Yakunopona yakushimensis Ishihara, 1954: 13. Holotype 9, JAPAN (ELKU.Fukuoka) [examined]. Syn. n.
Platymetopius australis Evans, 1966: 247. Holotype £, AUSTRALIA (ANIC, Canberra) [examined]. Syn. n.

Length : J, 4-5-5-6 mm, mean 5- 1 mm ; 9, 5-3-6-6 mm, mean, 6-0 mm.

Colour as in generic description with rostrum and sometimes apex of clypellus scarlet; head and thorax
rarely with testaceous marking (see 'Remarks' below), sometimes marked with stramineous or orange at
apex of vertex and on each side at base of vertex, on lateral margins of clypeus and on anterior region of
pronotum. Fore wing variably tinged with brown distally; apex of both clavus and claval veins with a small
brown spot.

External characters as in generic description with vertex short to long, medial length 1-7-3-5 times length
next to eyes; apex acute, narrowly rounded or fairly broadly angularly rounded, sides concave to slightly
convex (see 'Remarks' below).

Male genitalia as in generic description with pygophore variable in shape (see 'Remarks' below). Style
with apical process evenly tapered to acute apex, lateral lobe fairly strongly developed. Connective short.
Aedeagus with shaft slender, apical processes slightly expanded at mid length, strongly divergent.

Female genitalia as in generic description.

REMARKS. Some specimens from India have the markings on the head and pronotum orange (see
description) while one specimen from Australia has the head and thorax with testaceous mar-
kings. Considerable variation occurs in the shape of the head; usually the side margins of the
vertex are nearly straight but a few specimens from India have the side margins considerably
concave (Fig. 46). The face in profile is usually slightly convex to nearly straight but in one
specimen from Australia the face is strongly concave. In the male genitalia slight variation occurs
in the pygophore in the shape of the posterior lobes, the distal darkly pigmented region of the
internal ledge and in the number of macrosetae (compare Fig. 53 from Java with Webb, 1980:
849, fig. 115 from Aldabra).

DISTRIBUTION. Afrotropical, southern Palaearctic and Oriental regions as far south as Singapore
and Java (excluding Malaya and Sumatra) and from northern and north-eastern Australia.

MATERIAL EXAMINED

Dryadomorpha pallida Kirkaldy, holotype 9, Australia: Queensland, Bundaberg, ix-xii.1904 (Koebele)
(BPBM, Honolulu). Paganalia virescens Distant, lectotype 9, Seychelles: Silhouette, 1908 (BMNH, London).
Zizyphoides indicus Distant, lectotype 9, India: Calcutta, on Zizyphus jujuba Miller, 23.vii.1912 (BMNH,
London). Rhombopsis virens Haupt, lectotype <$, Israel: 'Palestine', Ben-Shemen, 17- 18.x. 1925 (HIM, Jena).
Rhombopsis viridis Singh-Pruthi, lectotype £, India: Punjab, Lyallpur, x.1929 (Rahman) (ZSI, Calcutta).
Rhombopsis chatterjeei Singh-Pruthi, lectotype $, India: North Salem, Uduparani, on unspiked sandel,
2911930 (Chatterjee) (FRI, Dehra Dun). Platymetopius antennalis Lindberg, holotype^, Cape Verde Is.:
Fogo I., supra Fte Aleixo, 19.ii.1954 (Lindberg) (ZMU, Helsinki). Yakunopona yakushimensis Ishihara,
holotype 9, Japan: Yakushima, Anbo, 25.viii.1952 (Takeya & Hiroshima} (ELKU, Fukuoka). Platymetopius
australis Evans, holotype <$, Australia: NW., Kimberley Research Station, via Wyndham, 23.viii. 1956
(Langfield) (ANIC, Canberra).

India: l<$, 1 9, Punjab, Lyallpur, x.1929, 7.X.1929 (ZSI, Calcutta) (paralectotypes of Rhombopsis viridis
Singh-Pruthi); 1 £, North Salem, Jawalagiri, 22.xii. 1930 (ZSI, Calcutta) (paralectotypes of Rhombopsis
chatterjeei Singh-Pruthi); 1 9, Coorg, 9.H.1930 (ZSI, Calcutta) (paralectotype of Rhombopsis chaterjeei
Singh-Pruthi); 3 <J, 3 9, Dehli, Indian Agricultural Research Institute, at light; 1 <J, Bihar, Pusa, 14.vi.1931 ; 1
<J, 6 9, Ludhiana, 25.xi.1976 (all in BMNH, London). Nepal: 3 $, 3 9, near Simra, Abhabbar, 180 m,
25-27.viii.1967; 1 9, near Birganj, Lothor, 135 m, 3.ix.l967 (all in CNC, Ontario); 1 9, Kathmandu,
1300-1400 m, 7-12.V.1966 (BPBM, Honolulu); 2 9, Arun Valley, below Tumlingtar, river Sabhaya, west
shore, 460 m, 22.xii.1961 (BMNH, London). Sri Lanka: 1 <J, Amiradhapura, at light, 22.iii.1953 (BMNH,
London). Bangladesh: 1 <£, Lawa Chera forest, Srimangal, 110 m, 27.ix.1961 (CAS, San Francisco). China: 1
9, Macao, i.1907; 1 & 39, Fujian, Yungan, 4.viii-24.ix.l940(all in BPBM, Honolulu). Japan: 1 & 29, Tokara
I., Nakanoshima, 25.v-13.vi.1953; 1 <$, Kyushu, Satamisaki, Osumi, 30.viii.1951 (all in ELKU, Fukuoka).
Hong Kong: 1 ^, Lantau I., Shek Pik Reservoir area, 21.vii.1964; numerous examples, Sai Kung Station,
30.i.l965; 9^, 89, Taipokau, 20.vi.-6.vii.1964 (all in BPBM, Honolulu). Taiwan: 1 & 19, Taipei Hsien
Santiaoling, 100-400 m, 19.xi.1957 (BPBM, Honolulu). Thailand: 1 9, South Banna, Nakhon, 108 m,
5-10.V.1958; 1 <J, Bangkok, at light, 4.xii.l957; 1 9, Chiangmai, Fang, 12-19.iv.1958 (all in BPBM, Hono-
lulu). Laos: 1 9, Vientiane, at light, S.v.1965 (BPBM, Honolulu). Philippines: 2 £, 19, Busuanga I., 4 km
north of San Nicolas, at light, 21-24.V.1962, 1 £, Palawan, 3 km north-east of Tinabog, at light, 14.V.1962;
1 9, Palawan, Tarumpitao Point, at light, 26.V.1958; 1 9, Culion I., 6 km west of Culion, at light, 7.vi.l962 (all

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

53

in BPBM, Honolulu). Singapore: 4 $, 1 ?, Mandai, mangrove, at light (BMNH, London). Java: 1$, Bogor,
at light, v.1954 (BMNH, London). Australia: 1 9, Western Australia, Kimberley Research Station, via
Wyndham, 23.viii.1956 (ANIC, Canberra) (paratype of Platymetopius australis Evans); 1 9, New South
Wales, Sydney (BMNH, London).

Dryadomorpha metrosideri (Osborn) comb. n.
(Figs 57-60)

Calotettix metrosideri Osborn, 1934: 247. Holotype9, MARQUESAS Is. (BPBM, Honolulu) [examined].
Calotettix metrosideri var. tincta Osborn, 1934: 248. LECTOTYPE $, MARQUESAS Is. (BPBM, Honolulu),

here designated [examined]. Syn. n.
Calotettix lais Eyles & Linnavuori, 1974: 40. Holotype^, COOK Is. (IM, Auckland) [examined]. Syn. n.

Length: $, Marquesas Is., 6-3-6-7 mm, mean 6-5 mm; Cook Is., 6-8-7-4 mm, mean 7-2 mm; Australia, 5-5
mm; 9, Marquesas Is., 7-0-7-7 mm, mean 7-2 mm; Cook Is., 7-8-8-5 mm, mean 8-1 mm.

Yellow or yellow tinged with green or orange; darker specimens with additional reddish and dark brown
markings on pronotum and scutellum ; some specimens (see 'Remarks' below) have face reddish, fore wings
either marked with brown on inner margin of clavus and with a brown spot at base of fourth apical cell, or
marked with brown bordering veins of clavus and veins cu and m and with a brown patch at apex of wing,
and female pygophore with a dark brown patch posteriorly on each side.

Figs 57-67 Dryadomorpha species. 57-60. D. metrosideri. (57) head and thorax, dorsal view; (58) fore
wing; (59, 60) aedeagus, lateral and posterior views. 61-63. D. robustipenis. (61) apex of left style,
ventral view; (62, 63) aedeagus, posterior and lateral views. 64-67. D. pacifica. (64) apex of the left
style, ventral view, Sarawak; (65) same, New Guinea; (66) aedeagus, lateral view; (67) connective,
dorsal view.

54 M. D. WEBB

External characters as in generic description with vertex short, approximately 1-5 times as long medially
as next to eyes, sides slightly convex, apex fairly broadly angularly rounded.
Male genitalia as in pallida but aedeagal processes in posterior aspect narrow throughout length.
Female genitalia as in pallida.

REMARKS. In addition to the differences in size noted above specimens from different localities
vary in the following ways: the single specimen from Australia has a medial longitudinal keel
dorsally on the face (absent in other specimens), the clypeus reddish, some veins of the fore wing
bordered with brown, and a brown patch at the apex of the fore wing; specimens from the
Marquesas Is. have the inner margin of the fore wing variably marked with brown (Fig. 58) and
the face sometimes marked with red, the thorax marked with red or brown and the female
pygophore marked with brown posteriorly; the specimens from the Cook Is. are without mar-
kings.

This species can be distinguished from other members of the genus by its short head. The male
genitalia are almost identical to those of pallida, differing only in the shape of the aedeagal
processes as noted above.

DISTRIBUTION. Marquesas Is., Cook Is. and northern Australia.

MATERIAL EXAMINED

Calotettix metrosideri Osborn, holotype 9, Marquesas Is.: Hiva Oa, Kopoa faa, miscel. sweeping, 831 m,
3.viii.l929 (Mwnford & Adamson) (BPBM, Honolulu). Calotettix metrosideri var. tincta Osborn, lectotype$,
Marquesas Is.: Hiva Oa, Feani Ridge, 1 140 m, 22.U932 (Le Bronnec) (OSU, Columbus). Calotettix lais Eyles
& Linnavuori, holotype <$, Cook Is.: Rarotonga, 15.xii.1937 (IM, Auckland).

Marquesas Is.: numerous specimens from Hiva Oa, 630-1140 m, on Weinmannia parviflora G. Forster,
Reynoldsia sp., Glochidion ramiflorwn G. Forster or Rapanea sp., I.viii-5.i.l932 (Le Bronnec or Mumford &
Adamson) (paratypes of Calotettix metrosideri Osborn and paralectotypes of Calotettix metrosideri var.
tincta Osborn) (BPBM, Honolulu; OSU, Columbus). Cook Is. 3 & 19, Rarotonga I., Papua Creek,
Vaimaanga, at light, 1 3.x. 1975; 7 c?, 39, Rarotonga I., Avatu Valley, at light, x. 1975; 2^, 1 9, Rarotonga I.,
Totokoitu Ridge, 210 m, 19.x. 1975; 1 £, Rarotonga I., Totokoitu, at light, 14.x. 1975 (all in DSIR, Auckland
and BMNH, London). Australia : 1 <J, Northern Territory, Darwin (JWE, Sydney).

Dryadomorpha viridia Osborn
Dryadomorpha viridia Osborn, 1934: 244. Holotype?, MARQUESAS Is. (BPBM, Honolulu) [examined].

Length: 9, 6- 7 mm.

Colour as in generic description with inner margin of clavus brown.

External characters as in generic description with vertex moderately long, medial length approximately
twice as long as length next to eyes, apex acutely rounded, sides slightly convex.

Male genitalia unknown.

Female genitalia as in generic description.

REMARKS. The single known specimen of this species is tentatively regarded as distinct from
metrosideri, differing only in its longer head.

DISTRIBUTION. Marquesas Is.

MATERIAL EXAMINED

Dryadomorpha viridia Osborn, holotype 9, Marquesas Is.: Nuku Hiva I., Toovii, 750 m, 4.viii.l931,
beating Metrosideros collina (J. R. & G. Forster) (Le Bronnec & Tauraa) (BPBM, Honolulu).

Dryadomorpha robustipenis sp. n.

(Figs 6 1-63)

Length : $, 5-2-6-0 mm, mean 5-5 mm; 9, 6-0-6-3 mm, mean 6- 1 mm.

Colour as in generic description with a small brown spot at apex of clavus and claval veins of forewing.

External characters as in generic description with vertex moderately long, medial length 2-5-3-0 times as
long as length next to eyes, apex acutely rounded, sides slightly convex to slightly concave.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 55

Male genitalia similar to pallida but connective moderately long and aedeagus more robust with apical
processes shorter and only slightly divergent.
Female genitalia as in generic description.

REMARKS. This species closely resembles pacifica and is known to be sympatric with this species
over part of its range in Sarawak and Sabah. It can be distinguished from pacifica in the male
genitalia by the narrower apical process of the style, the more pronounced lateral lobe of the style
and the more robust shaft of the aedeagus. From the similar but more northerly and southerly
distributed pallida it differs in the male genitalia as noted above.

DISTRIBUTION. Malaya and Borneo (Sarawak, Sabah).

MATERIAL EXAMINED

Holotype <?, Sarawak: foot of Mount Dulit, junction of rivers Tinjar and Lejok, at light, l.x.1932 (Hobby
& Moore) (BMNH, London).

Paratypes. Malaya: 1 <J, Kuala Lumpur, Klang gates, 31.xii.1958 (Quate) (BPBM, Honolulu); 1 9, Kuala
Lumpur, 13.iii.1939 (Pendlebury) (BMNH, London). Borneo: 3 <J, 2 ?, Sarawak, same data as holotype,
26.viii.-25.ix.1932; 1 & Sarawak, Paya, Palah, at light, 4.xi.l964 (Rothschild) (all in BMNH, London); 1 <J,
Sarawak, Bau, lake area, 30.viii.1958 (Mao) (BPBM, Honolulu); 1 £, Sabah, Tawau, Quoin Hill, at light,
3-7. vii. 1962 (Holtman); 1 <J, Sabah, Tawau, Quoin Hill, Cocao Research Station, 22.viii.1962 (Hiroshima)
(all in BPBM, Honolulu).

Dryadomorpha pacifica sp. n.

(Figs 64-67)

Length: <$, 5-2-6-5 mm, mean 5-6 mm; $, 5-5-6-4 mm, mean 5-9 mm.

Colour and external characters as in robustipenis.

Male genitalia similar to robustipenis but apical process of style broader apically and foot-like, or with a
subapical prominence, and lateral lobe ol style less pronounced and aedeagus with shaft narrower and
tapered more at midlength to apex.

Female genitalia as in generic description.

REMARKS. In this species slight variation occurs in the shape of the apical process of the style and
the processes of the aedeagus. In the specimens from Sarawak and Sabah the apical process of the
style is foot-like and the aedeagal processes are narrow and nearly parallel, while specimens from
New Guinea have the apical process of the style with a slight subapical prominence and the
processes of the aedeagus stouter and more divergent. The significance of these differences cannot
be ascertained at the present time.

This species closely resembles robustipenis and is known to be sympatric with this species over
part of its range in Sarawak and Sabah. It can be distinguished from robustipenis in the shape of
the style and aedeagus as noted above. It differs from the similar but more northerly and
southerly distributed pallida in the longer connective and the more robust aedeagus with the
apical processes shorter and only slightly divergent.

DISTRIBUTION. Borneo (Sarawak, Sabah), New Guinea and Vanuatu.

MATERIAL EXAMINED

Holotype rf, New Guinea: Sarmi, W. to Hollandia, 20-23.vii.1959 (Maa) (BPBM, Honolulu).

Paratypes. Borneo: l<$, Sarawak, Bau district, lake area, 30.viii.1958 (Maa) (BPBM, Honolulu); 1^,
Sabah, Tawau, Quoin Hill, Cocoa Research Station, primary forest, 3.x. 1962 (Hiroshima); 1 ^, Sabah,
Tawau, Quoin Hill, 3-7.vii.1962 (Holtman) (BPBM, Honolulu). New Guinea: 1 J, Nabire, 5-50 m, at light,
25.viii-2.ix.1962 (Sedlacek); 1 <£ mouth of river Tor, 4 km E. of Maffen, 19. vii. 1959 (Maa); 1 & Kokoda, 400
m, at light, 15-20.xi.1965 (Sedlacek); 1 $, Bodem, 100 m, 11 km SE. of Oerberfaren, at light, 7-17.vii.1959
(Maa) (all in BPBM, Honolulu); 15 <£ same data as holotype (BPBM, Honolulu; BMNH, London).
Vanuatu: 3$, Espiritu Santo, Apouna river camp, 300 m, 9-12.ix.1971 (Robinson) (SAM, Adelaide); 1 cJ,
Espiritu Santo, Narango, 90 m, vi.1960 (Brandt) (BPBM, Honolulu); 1 ?, Santo I., 17.vi.1925 (Buxton)
(BMNH, London).

56

M. D. WEBB

RHUTELORBUS gen. n.

Type-species : Rhutelorbus merinoi sp. n.

Yellow or greenish yellow, sometimes with paler patches on head and thorax ; fore wings with a small brown
spot at apex of both clavus and claval veins.

Head wider than pronotum; anterior margin acutely rounded in profile, transversely striate, becoming
finely carinate and rim-like medially; ocelli on margin, distant from eyes, not visible from above; anterior
tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial length approxi-
mately 2-5 times length next to eyes, sides slightly convex, apex acutely rounded; with a slight longitudinal
depression; finely rugose. Face elongate, shagreen; lateral margin adjacent eye visible dorsally; upper
margin slightly depressed each side of mid line, forming a faintly striate longitudinal keel medially; face in
profile more or less straight or concave anteriorly; clypeus elongate, lateral margins constricted near
antennae; clypellus elongate, expanded apically; transclypeal suture indistinct; lora large; antennal pit deep
with inner margin angularly rounded to clypeus, rim-like and nearly carinate; antennal ledge slight; an-
tennae very long, extending to beyond apex of clavus. Pronotum approximately twice as wide as long, side
margins very short, without a carina; finely transversely rugose. Scutellum approximately equal in length to

Figs 68-78 Rhutelorbus merinoi. 68, head and thorax, dorsal view; 69, left subgenital plate and apex of
left style, dorsal view; 70, <$ genitale capsule, lateral view; 71, left style, ventral view; 72, connective,
lateral view; 73, apex of left style, lateroventral view; 74, connective, dorsal view; 75, 9 pregenital
sternite, ventral view; 76, second valvulae, lateral view; 77, 78, aedeagus, lateral and posterior views.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 57

pronotum, shagreen, obscurely rugose posteriorly. Fore wing with three subapical cells, first subapical open,
second and third subapicals closed; one or two additional veinlets in subcostal region near to fifth apical
cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of two or three fine setae distally
on anterior surface; hind femur with apical setal formula 2+1 + 1 with first proximal seta slender.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore straight dorsally, without apodemes; pygophore lobes
with several macrosetae and short fine setae. Xth segment moderately long, compressed dorsoventrally.
Subgenital plate elongate, triangular with numerous moderate to long fine setae dorsally and several short
to moderately long fine setae ventrally. Valve triangular. Style moderately long, narrow, with lateral lobe
and basal apophyses prominent; apical process fairly short, curved ventrally, tapered to apex, crenulate
api?ally; few sensory papilla and a single seta adjacent lateral lobe. Connective Y-shaped, stem elongate
with lateral margins keel-like dorsally ; arms short. Aedeagus with shaft elongate, curved dorsally with a pair
of apical, dorsally directed processes; gonopore small, situated at apex; basal apodeme moderately long,
narrow.

Female genitalia with posterior margin of pregenital sternite extended posteriorly; second valvulae united
at first dorsal tooth (arrowed in Fig. 76), elongate, very slightly expanded distally; without a basal promi-
nence; dorsal teeth robust, unaligned, extending over approximately distal third of valvulae.

REMARKS. This genus is similar externally to Dryadomorpha and some species of Parohinka but
can be distinguished from these and other genera of the subfamily by the rugose vertex and
pronotum, the lateral margins of the face adjacent to the eyes being visible dorsally, and by the
extended posterior corners of the pregenital sternite (also present in some species of Parohinka).

DISTRIBUTION. Philippines, Malaya and Borneo (Sarawak).

Rhutelorbus merinoi sp. n.

(Figs 68-78)

Length: c?, 5-6-6-5 mm, mean, 6-0 mm; 9, 6-2-7-4 mm, mean 6-8 mm.

Colour and external characters as in generic description.

Male genitalia as in generic description with pygophore lobes acutely produced posteriorly and aedeagus
with shaft narrow, slightly tapered from base to apex with apical processes fairly short (see 'Remarks' below).

Female genitalia with posterior margin of pregenital sternite extended posteriorly, maximum length as in
Fig. 75, sometimes one-third this length.

REMARKS. The male specimen from the Philippines has the apical process of the style curved more
ventrally and the apical processes of the aedeagus shorter than in other specimens.

DISTRIBUTION. Philippines, Malaya and Borneo (Sarawak).

MATERIAL EXAMINED

Holotype £, Malaya : Kuala Lumpur, at light, 16.X.1928 (Corbett) (BMNH, London).

Paratypes. Philippines: 1 9, Tawi Tawi, Lapid Lapid at Manalik Channel, at light, 19.xi.1961; 1 ^, Tawi
Tawi, Tarawakan, north of Batu Batu, at light, 13.xi.1961 (both in ZM, Copenhagen). Malaya: 1 9, Kuala
Lumpur, at light, 1811931 (Pendlebury) (BMNH, London); 2^, 29, Kuala Lumpur, 24-3 l.xii. 1958 (Quate);
4 c?, 1 9, Kuala Lumpur, Klang gates, 3 l.xii. 1958 (Quate); 1 9, Kuala Lumpur, 19 km south of Subang,
23.xii.1958 (Quate); 1 & Kuala Tahan, 15-16.xii.1958 (Quate) (all in BPBM, Honolulu); 1 <J, Selangor, at
light, 19.iv.1932 (Pendlebury) (BMNH, London). Borneo: 1 $ no locality, 1898 (Noualhier) (MNHN, Paris);
1 cJ, Sarawak, foot of Mount Dulit, junction of R. Tinjar and Lejok, old secondary forest, at light,
29.viii.1932 (Hobby & Moore) (BMNH, London); 1 9, Sarawak, Bau District, Bidi, 19-240 m, at light,
2.iv.l958 (Maa); 1 9, Sarawak, Kuching, Santubong, 797-1500 m, 18-30.vi.1958 (Maa) (both in BPBM,
Honolulu).

PAROHINKA gen. n.

Type-species: Muirella longiseta Melichar, 1914.

Yellow, greenish yellow or brownish yellow; head with yellow or whitish yellow patches, usually mottled
with brown. Fore wings with a small brown spot at apex of both clavus and claval veins, often faint. Legs
spotted with brown.

58 M. D. WEBB

Head wider than pronotum; anterior margin angularly rounded in profile, transversely striate laterally,
becoming finely carinate and usually rim-like medially, ocelli on margin distant from eyes, not visible from
above; anterior tentorial branches curved anteriorly, not bifurcate. Vertex triangularly produced, medial
length 1-5-4-0 times length next to eyes, sides slightly convex to slightly insinuate in longer-headed forms,
apex acute to broadly angularly rounded; with a medial longitudinal depression; finely longitudinally
striate, transversely striate anteriorly. Face elongate to only slightly longer than wide, shagreen; upper
margin usually slightly depressed each side of mid line, forming a faintly striate longitudinal keel medially,
indistinct in shorter-headed forms; face in profile convex to concave in longer-headed forms; clypeus
elongate, lateral margins constricted near antennae; clypellus elongate, expanded apically; transclypeal
suture distinct or indistinct; lora large; antennal pit deep with inner margin angularly rounded to clypeus
and sometimes faintly rim-like and nearly carinate; antennal ledge absent; antennae very long, when
recurved extending beyond apex of clavus. Pronotum approximately twice as wide as long, side margins
very short, without a carina; finely and irregularly transversely striate, obscurely rugose anteriorly. Scutel-
lum approximately equal in length to pronotum, shagreen with posterior region obscurely rugose. Fore
wings with three subapical cells, first subapical cell open, second and third subapical cells closed; with a few
additional veinlets in subcostal region near to fifth apical cell. Fore tibia with dorsal setal arrangement 1:4;
fore femur with a series of seven setae distally on anterior surface; hind femora with apical setal formula
2+1 + 1 with first proximal seta slender.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with posterior margin of pygophore straight dorsally, without apodemes; pygophore lobes
with a slight oblique internal ledge, usually not attaining posterior margin; posterior margin sometimes
serrate and sometimes extended spine-like dorsally; lobes with several macrosetae and numerous short to
moderately long fine setae. Xth segment moderate to long, compressed dorsoventrally, narrowing basally in
lateral aspect. Valve triangular. Subgenital plate elongate, triangular with basal lobe relatively short and
prominent ; outer margin with numerous long to very long fine setae on dorsal and ventral surfaces ; apex
with few very short stout setae. Style short with basal apophyses indistinct; apical process short to moder-
ately long, curved ventrally, tapered to apex or spatulate, ventral surface crenulate; lateral lobe prominent;
few sensory papilla ventrally adjacent lateral lobe. Connective Y-shaped, stem moderate to very long,
narrow, with lateral margins keel-like dorsally; arms short. Aedeagus assymetrical ; shaft elongate, curved
dorsally, sometimes recurved ventrally at apex, either of similar width throughout length, tapered to apex or
expanded distally in lateral aspect, sometimes compressed anteroposteriorly; with from one to three pro-
cesses distally; sometimes with an apical membranous region adjacent to gonopore; gonopore small,
situated at or near apex on left or right lateral surface within a species or on posterior surface; basal
apodeme short to moderately long, compressed laterally or anteroposteriorly.

Female genitalia with posterior margin of pregenital sternite either broadly V-shaped medially or with
posterior corners extended posteriorly. Second valvulae united at or before first dorsal tooth (arrowed in
Figs 110, 121, 133, 148), expanded distally, fairly robust, without a dorsoanterior prominence; dorsal teeth
fine to robust, unaligned, extending 0-25-0-40 times distance from apex to base of valvulae; dorsal scler-
otized region moderately long.

REMARKS. This genus is almost identical externally to Dryadomorpha but can usually be dis-
tinguished by the pale patches on the anterior margin of the head. Also similar externally to
Rhutelorbus (see 'Remarks', p. 57). In the male genitalia the pygophore has an internal ledge as in
Dryadomorpha, Karoseefa and Oceanopona, but the asymmetrical aedeagus is not found in any
other genus of the subfamily. In the female genitalia the extended posterior corners of the
pregenital sternite of some species (dulita-group) are as in Rhutelorbus and the second valvulae
are as in Rhutelorbus, Karoseefa and Oceanopona.

On the basis of the male and femal genitalia Dryadomorpha can be divided into two groups.
The dulita-group (dulita, brevicephala, philippina, recurva, apicalis, longiseta and malayensis)
differs from the morona-group (morona, sinuata, trispicata, lotophagorum and spinosa) in having
the apex of the aedeagal shaft continued as a process on one side and the posterior corners of
the female pregenital sternite extended posteriorly either process-like or lobe-like in dulita.

DISTRIBUTION. From NE. India to the Philippines and South East Asia and south to New Guinea
and the Pacific islands. The dulita-group extends as far south as Indonesia, while the morona-
group is found in New Guinea and eastwards to the Cook Is.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 59

Key to species of Parohinka

Males

1 Aedeagus with apex of shaft continued as a process on one side (Figs 105, 125). (Oriental region

and south to Indonesia) 6

Aedeagus with apex of shaft not continued as a process on one side. (New Guinea and eastwards
to the Cook Is.) 2

2 Aedeagus with two processes (both lateral or one lateral and one posterior) .... 3
Aedeagus with three processes (two lateral and one posterior) 5

3 Aedeagus with shaft in lateral aspect of similar width throughout length or narrower apically;

with two lateral processes directed ventroanteriorly 4

Aedeagal shaft in lateral aspect expanded apically; with one lateral process directed ventropo-
steriorly and one posterior process directed ventroanteriorly (Figs 93, 94)

lotophagorum (Kirkaldy) (p. 63)

4 Aedeagal shaft in lateral aspect evenly curved from base to apex (Fig. 84) . . morona sp. n. (p. 60)
Aedeagal shaft in lateral aspect sinuate apically (Fig. 86) sinuata sp. n. (p. 61)

5 Posterior process of aedeagus short, approximately half length of lateral processes (Fig. 91)

trispicata sp. n. (p. 61)
Posterior process of aedeagus long, similar in length to lateral processes (Fig. 102)

spinosa sp. n. (p. .64)

6 Apex of aedeagal shaft curved ventrally (Fig. 124). (Borneo (Sarawak)) . . recurva sp. n. (p. 67)
Apex of aedeagal shaft curved dorsally 7

7 Aedeagus with one process 8

Aedeagus with two or three processes 9

8 Connective approximately equal in length to style. (Malaya) . . . malayensis sp. n. (p. 70)
Connective approximately 1-5 times length of style. (NE. India to Indonesia)

longiseta (Melichar) (p. 68)

9 Aedeagus with two processes. (Borneo (Sarawak, Sabah)) apicalis sp. n. (p. 68)

Aedeagus with three processes ............. 10

10 Lateral aedeagal processes short, arising from near posterior margin of shaft (Fig. 119).

(Philippines) philippina sp. n. (p. 66)

Lateral aedeagal processes moderately long, arising from near anterior margin of shaft (Fig. 107) 11

11 Aedeagal shaft in lateral aspect narrowly club-shaped (Fig. 107). (Borneo (Sarawak))

dulita sp. n. (p. 64)
Aedeagal shaft in lateral aspect of similar width throughout length (Fig. 1 15). (Borneo (Sarawak,

Sabah)) brevicephala sp. n. (p. 66)

Females

The following key is incomplete because females of morona, sinuata and trispicata are indistinguishable
(grouped together under couplet 2) and spinosa and recurva are known only from the male.

1 Posterior corners of pregenital sternite extended posteriorly, process-like or lobe-like. (Oriental

region as far south as Indonesia) 3

Posterior corners of pregenital sternite not extended posteriorly as above. (New Guinea and
eastwards to the Cook Is.) . 2

2 Posterior margin of pregenital sternite oblique and nearly straight each side of mid line (Fig. 83)

morona sp. n. (p. 60), sinuata sp. n. (p. 61), trispicata sp. n. (p. 61)
Posterior margin of pregenital sternite angled at mid length each side of mid line (Figs 96, 97)

lotophagorum (Kirkaldy) (p. 63)

3 Posterior corners of pregenital sternite extended process-like 4

Posterior corners of pregenital sternite lobe-like (Fig. 109) dulita sp. n. (p. 64)

4 Vertex with medial length 0-5-0-9 times basal width between eyes ... 5
Vertex with medial length 1-0-1-2 times basal width between eyes 6

5 Vertex with medial length 0-5 times basal width between eyes. Extended posterior corners of

pregenital sternite short (Fig. 118). (Borneo (Sarawak)) . . . . brevicephala sp. n. (p. 66)
Vertex with medial length 0-5-0-9 times basal width between eyes. Extended posterior corners of

pregenital sternite long (Fig. 139). (NE. India to Indonesia) . . . longiseta (Melichar) (p. 68)

6 Pale to greenish yellow. Second valvulae fairly broad (Fig. 121). (Philippines)

philippina sp. n. (p. 66)
Sordid to brownish yellow. Second valvulae narrow (Fig. 148). (Malaya) . malayensis sp. n. (p. 70)

60

M. D. WEBB

Parohinka morona sp. n.

(Figs 79-85)

Length:^, 5-9-6- 3 mm, mean 6-1 mm;?, 6-6 mm.

Pale to greenish yellow. Vertex, dorsal region of face and sometimes medial region of pronotum mottled
with brown. Inner margin of fore wing sometimes tinged with brown (see 'Remarks' below); other markings
as in generic description.

External characters as in generic description with vertex long, medial length 2-8-3-5 times length next to
eyes and 1-0-1-2 times basal width between eyes. Face longer than wide with upper medial keel present;
clypeus elongate.

Male genitalia as in generic description with pygophore moderately long. Xth segment extending to or
slightly beyond posterior margin of pygophore. Connective moderately long (as in malayensis, Fig. 145).
Style with apical process digitate, apex of process truncate in medial aspect. Aedeagus with shaft in lateral
aspect tapered gradually from base to apex and terminating in a pair of ventroanteriorly directed processes;
shaft sinuate in posterior aspect; gonopore situated laterally immediately below apical processes.

Female genitalia with posterior margin of pregenital sternite shallowly V-shaped (see 'Remarks' below);
second valvulae united at first dorsal tooth; teeth robust, extending to approximately 0-4 times distance
from apex to base of valvulae (as in longiseta, Fig. 133).

REMARKS. Some of the specimens examined of this species have a brown medial longitudinal band
dorsally, from the head to the apex of the fore wings.

80

81

82

83

85

Figs 79-86 Parohinka species. 79-85. P. morona. (79) head and thorax, dorsal view; (80) apex of left
style, ventromedial view; (81) ^ genital capsule, lateral view; (82) left style, ventral view; (83)$
pregenital sternite, ventral view; (84, 85) aedeagus, lateral and posterior views. 86. P. sinuata,
aedeagus, lateral view.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 61

The following four species and morona form a distinct group based on distribution (New
Guinea and east to the Cook Is.) and also on the shape of the male and female genitalia (see
'Remarks' under generic description, p. 58). On external appearance and in the female genitalia
morona is identical to trispicata and sinuata. Females of the three species are therefore included in
their respective type-series as they were taken at the same localities as the males therein and there
being no sympatry between the males of the three species samples. Only slight differences in the
shape of the pregenital sternite distinguishes females of morona, sinuata and trispicata from
lotophagorum; in the first three species the posterior margin of the pregenital sternite is oblique
and almost straight each side of the mid line while in lotophagorum it is angled at mid length each
side of the mid line. In external appearance morona is similar to spinosa and is sympatric with this
species over part of its range in Papua New Guinea. Females of spinosa are unknown. In the male
genitalia morona differs from the above four species in having the posterior margin of the
pygophore non-serrate, the aedeagus with two apical processes and the aedeagal shaft evenly
curved from base to apex in lateral aspect.

DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea).

MATERIAL EXAMINED

Holotype ^, New Guinea: Irian Jaya, Cyclops Mts, Sabron, camp 2, 660 m, vii.1936 (Cheesman) (BMNH;
London)

Paratypes. New Guinea: 1 <J, 1 ?, Irian Jaya, Torricelli Mts, Mokai Vill., 750 m, 8-15.xii.1958 (Brandt); 2
cJ, I.J., 40 km north of Baliem Val., 1300 m, at light, 5-ll.xi.1961 (Quate); 1 & Papua New Guinea, Swart
Val., Karubaka, 1450 m, 17.xi.1958 (Gressitt) (all in BPBM, Honolulu).

Parohinka sinuata sp. n.

(Fig. 86)

Length : <J, 6-2-6-6 mm, mean 6-4 mm; $, 7-5 mm.

Colour and external characters as in morona with medial length of vertex 2-7-3-0 times length next to eyes
and 0-8-1-1 times basal width between eyes.

Male genitalia as in morona but aedeagal shaft apically with processes slightly shorter.

Female genitalia as in morona.

REMARKS. This species is closely related to morona differing only in the shape of the aedeagus as
noted above (see also 'Remarks' under morona, p. 60).

DISTRIBUTION. New Guinea (Papua New Guinea)

MATERIAL EXAMINED

Holotype cJ, New Guinea: Papua New Guinea, Lae, Singuawa R., 147° 10' E, 6° 45' S, 30 m, at light,
Kunai grass, 8.iv.l966 (Wilkes) (BPBM, Honolulu).

Paratypes. New Guinea: 1 <$, 1 9, same data as holotype (BMNH, London; BPBM, Honolulu); 1 £,
P.N.G., Wau, Morobe District, 1250m, 2311963 (Sedlacek) (BPBM, Honolulu).

Parohinka trispicata sp. n.

(Figs 87-91)

Length : $, 5-6-6-0 mm, mean 5-8 mm; $, 6-4-7-0 mm, mean 6-7 mm.

Colour and external characters as in morona with medial length of vertex 2-5-3-0 times length next to eyes
and approximately equal to basal width between eyes.

Male genitalia similar to morona but apex of aedeagus with a small additional process on one side
adjacent to gonopore.

Female genitalia as in morona.

REMARKS. This species is identical externally and in the female genitalia to morona and sinuata
(see 'Remarks' under morona, p. 60). In the male genitalia it differs from morona and sinuata in
having three processes on the aedeagus.

DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea) and Solomon Is.

M. D. WEBB

98

101

102

Figs 87-102 Parohinka species. 87-91. P. trispicata. (87) $ pygophore and Xth segment, lateral view;
(88) apex of style, ventromedial view; (89) connective, dorsal view; (90, 91) aedeagus, lateral and
posterior views; 92-97. P. lotophagorum. (92) <$ pygophore and Xth segment, lateral view; (93, 94)
aedeagus, lateral and posterior views; (95) apex of left style, ventromedial view; (96,97) $ pregenital
sternite, ventral view; 98-102. P. spinosa. (98) <$ pygophore and Xth segment, lateral view; (99) apex
of left style, ventromedial view; (100) aedeagus, lateral view; (101) left style, ventral view; (102)
aedeagus, posterior view.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 63

MATERIAL EXAMINED

Holotype cJ, Solomon Is.: Kolombangara, Pepele, 30 m, 6.H.1964 (Shanahan) (BPBM, Honolulu)
Paratypes. New Guinea: 1 (J, Irian Jaya, 10.xi.1944 (Aarons) (CAS, San Fransisco); 3^, 1 ?, I.J., Hollandia,
Binnen, 100 m, at light, 31.X.1958 (Gressitt); 1 9, I.J., Hollandia, at light, 28.ii.1960 (Maa); 1 & Papua New
Guinea, Adelbert Mts, Wanuma, 800-1000 m, at light, 27.x. 1958 (Gressitt) (all in BPBM, Honolulu). Solo-
mon Is.: 10 cJ, 4 9, Kolombangara, Pepele, 30 m, 6, 10, 15.ii.1964 (Shanahan) (BPBM, Honolulu; BMNH,
London)

Parohinka lotophagorum (Kirkaldy) comb. n.
(Figs 92-97)

Dryadomorpha lotophagorum Kirkaldy, 1907: 41. 1 <$, 1 9 syntypes, FIJI: Viti Levu, Ruwa, iii-vi (Muir) (lost).
NEOTYPE (J, FIJI (BPBM, Honolulu), here designated [examined].

Length: J, 5-4-6-5 mm, mean 6-0 mm; 9, 6-6-7-6 mm, mean 7-0 mm.

Pale to greenish yellow. Vertex and dorsal region of face mottled with brown. Fore wings sometimes
tinged with pinkish orange. Other markings as in generic description.

External characters as in morona with medial length of vertex 2-4-3-0 times length next to eyes.

Male genitalia similar to morona but posterior margin of pygophore serrate and apical process of style
with a subapical prominence in medial aspect. Aedeagus with shaft tapered from base to near apex and
expanded slightly at apex in lateral aspect; a more or less straight process arising at apex on one side,
directed ventrolaterally and another process arising on opposite side to first and slightly below apex,
strongly curved ventrolaterally; gonopore apical on posterior surface.

Female genitalia with posterior margin of pregenital sternite V-shaped medially; second valvulae as in
morona.

REMARKS. The type-series of lotophagorum was listed as missing at the time when the Hawaiian
Sugar Planters' Association collection was transferred to the Bishop Museum and is presumed
lost. The specimens described above fit the original description almost exactly, only the darker
brown spots on the lateral margins of the clypeus, mentioned by Kirkaldy, are not clear.

In external appearance this species is similar to morona, sinuata, trispicata and spinosa and is
sympatric with trispicata over part of its range in the Solomon Is. In the female the pregenital
sternite is slightly different in shape to that of morona, sinuata and trispicata (see 'Remarks' under
morona, p. 60), and in the male genitalia both this species and spinosa can be distinguished from
other members of the genus by the serrate posterior margin of the pygophore. From spinosa it
differs in the male genitalia having the posterior margin of the pygophore without a spine-like
process, the style with the apical process more robust with a preapical prominence, and the
aedeagus with two processes.

DISTRIBUTION. New Guinea (Irian Jaya, Papua New Guinea), Solomon Is., Vanuatu, Fiji and
Niue Is.

MATERIAL EXAMINED

Dryadomorpha lotophagorum Kirkaldy, neotype $, Fiji: Lau group, Ono-i-Lau, Nukuni, 0-50 m,
24.ii.1971 (Krauss) (BPBM, Honolulu).

New Guinea: 1 $, 1 9, Irian Jaya, Waigeu, Camp Nok., 750 m, iv.v.1938 (Cheesman) (BMNH, London); 1
(J, I.J., Japen I., SSE. Sumberbaba, Dawai R., at light, 22.x. 1962 (Holtmann); 2 <$, 19, I.J., Waris, S. of
Hollandia, 450-500 m, m.v. light-trap, l-15.viii.1959 (Maa); 1 & 2 9, I. J-, Bodem, 100 m, 11 km SE. of
Oerberfaren, m.v. light-trap, lO.vii, 16.xii.1959 (Maa); 3 $, Papua New Guinea, Kokoda, 400 m, light-trap,
malaise trap, 15-20.xi.1965 (Sedlacek) (all in BPBM, Honolulu); 1 & P.N.G., Madang Dist., Finisterre Mts,
Damanti, 105 m, stn no. 46, 2-1 1.x. 1964 (Bacchus) (BMNH). Bismark Archipeligo: 4<J, New Britain, Gazelle
Pen., Gaulim, 130 m, 140 m, at light, 23-28.X, 19-20.xi.1962 (Sedlacek); 1 <J, N.B., Gazelle Pen., Mt Sinewit,
900 m, at light, 5-14.xi.1962 (Sedlecek) (all in BPBM, Honolulu). Solomon Is.: 1 9, 2 specimens (abdomens
missing), Guadalcanal, Mt Austen, 15.viii.1966 (Greenslade) (BMNH, London); 2^, G., 17-6 km SE. Tetere,
Tathimani, light-trap, 12. v. 1960 (O'Brien); 1 J, G., Kukum, 10 m, light-trap, 21.vi.1956 (Gressitt); 4£, 59,
Kolombangara, Pepele, 30 m, 6, 10, 15.ii.1964 (Shanahan); 2 cJ, 39, NW. Malaita, Dala, light-trap, 5-
7.vi.l964 (Straatman); 1 <$, San Cristobal, Wairahu R., 100 m, pressure lamp, 9-15.V.1964 (Straatman) (all in
BPBM, Honolulu). Vanuatu: 1 £, Lamen I., 0-10 m, i.1976 (Krauss) BPBM, Honolulu); 1 <J, Tanna I.,

64 M. D. WEBB

Isokoai (Enpinana), 28.vii.1971 (Gross) (SAM, Adelaide). Fiji: 2 cJ, 1 $, Lau, Group, Ono-i-Lau, Nukuni,
0-50 m, 24.ii.1971 (Krauss) (BPBM, Honolulu). Cook Is.: 1 ?, Niue I., nr Lefuka, bush area, m.v. light,
18.vi.1975 (Maddison) (BMNH, London); 1 <J, Niue I., Kavara, at light, ll.vi.1975 (Dugdale) (DSIR, Auck-
land).

Parohinka spinosa sp. n.

(Figs 98-102)

Length: <$, 5-8 mm.

Pale yellow with vertex and dorsal region of face mottled with brown ; other markings as in generic
description.

External characters as in morona with medial length of vertex 3-0 times length next to eyes and 1-2-1-3
times basal width between eyes.

Male genitalia similar to lotophagorum but posterior margin of pygophore extended into a long spine-like
process dorsally and apical process of style narrower, without a preapical prominence. Aedeagus with shaft
expanded from base to apex in lateral aspect, two lateral and one posterior process arising immediately
below apex, directed ventrally, moderately long.

Female genitalia unknown.

REMARKS. In external appearance this species is similar to morona, sinuata, trispicata and lotopha-
gorum and sympatric with morona in its single known locality in Papua New Guinea. In the male
genitalia spinosa and lotophagorum can be distinguished from other members of the genus by the
serrate posterior margin of the pygophore. From lotophagorum it differs in the male genitalia in
the shape of the pygophore and style, as noted above, and the aedeagus with three processes.

DISTRIBUTION. Known only from the type-locality in New Guinea (Papua New Guinea).

MATERIAL EXAMINED

Holotype £, New Guinea: Papua New Guinea, Cyclops Mts, Sabron, 279 m, v.1936 (Cheesman) (BMNH,
London)

Paratypes. New Guinea: 1 $, same data as holotype except, camp 2, 600 m; 3 <$, same data as holotype
except, vi.1936 (BMNH, London)

Parohinka dulita sp. n.

(Figs 103-1 10)

Length: ^, 5-5-5-9 mm, mean 5-7 mm; 9, 5-8-6-2 mm, mean 6-1 mm.

Pale, sordid or greenish yellow. Vertex and dorsal region efface mottled with pale brown; other markings
as in generic description.

External characters as in morona but vertex moderately long, medial length 2-1 times length next to eyes
and 0-8 times basal width between eyes.

Male genitalia as in generic description with pygophore moderately long and Xth segment extending
slightly beyond posterior margin of pygophore. Connective moderately long, upturned apically. Style with
apical process digitate. Aedeagus with shaft narrowly club-shaped in lateral aspect with a moderately long
process at apex on one side, directed dorsally, and a moderately long subapical process on each side, arising
against anterior margin, directed ventrally; apical region of shaft adjacent process and surrounding go-
nopore membranous.

Female genitalia with posterior margin of pregenital sternite broadly V-shaped medially, each posterior
corner slightly produced lobe-like; second valvulae united at approximately 0-40 distance from apex to base
of valvulae ; teeth fine, extending to approximately 0-25 distance from apex to base of valvulae.

REMARKS. The following six species and dulita form a distinct group based on distribution
(Oriental region from NE. India to Indonesia) and on the shape of the male and female genitalia
(see 'Remarks' under generic description, p. 58). In the length of the vertex dulita is similar to
some specimens of longiseta but differs from this and other members of the genus by the lobe-like
posterior corners of the pregenital sternite and the valvulae united basad of the teeth. It is very
similar to brevicephala in the male genitalia but differs in having the apical process of the style
narrower in medial aspect and the shaft of the aedeagus narrowly club-shaped. This species is
known to be sympatric with longiseta and brevicephala in Sarawak.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

104 . VA 106

65

108

109

112

116

Figs 103-118 Parohinka species. 103-1 10. P. dulita. (103) head and thorax, dorsal view; (104) left style,
ventral view; (105) aedeagus, posterior view; (106) apex of left style, ventromedial view; (107)
aedeagus, lateral view; (108) connective, dorsal view; (109)9 pregenital sternite; (110) second valvu-
lae, lateral view. 111-118. P. brevicephala. (Ill) head and thorax, dorsal view; (112) face; (113)
aedeagus, posterior view; (114) apex of left style, medial view; (115) aedeagus, lateral view; (116)
connective, lateral view; (117) left style, ventral view; (1 18) 9 pregenital sternite, ventral view.

DISTRIBUTION. Known only from Borneo (Sarawak, Mt Dulit).

MATERIAL EXAMINED

Holotype <$, Borneo: Sarawak, foot of Mt Dulit, junction of R. Tinjar & Lejok, light-trap, 31.viii.1932
(Hobby & Moore) (BMNH, London)

Paratypes. Borneo: 2 $, same data as holotype, except, old secondary forest, 29.vii.1932; 1 9, same data as
holotype except, 29.viii.1932; 1 <$, same data as holotype except, l.ix.1932; 1 9, same data as holotype except,
4.X.1932; 1 $, Sarawak, Mt Dulit, Dulit trail, old secondary forest, 3.ix.l932 (Hobby & Moore) (BMNH,
London)

66 M. D. WEBB

Parohinka brevicephala sp. n.

(Figs 11 1-1 18)

Length : <£, 6-2 mm ; 9, 6-9-7- 1 mm, mean 7-0 mm.

Colour as in dulita.

External characters as in generic description with vertex short, medial length 1-5 times length next to eyes
and 0.5 times basal width between eyes. Face slightly longer than wide with upper medial keel very slight or
absent ; clypeus relatively short and broad.

Male genitalia similar to dulita but style with apical process slightly expanded in medial aspect and
aedeagus with shaft of similar width throughout length in lateral aspect.

Female genitalia with posterior margin of pregenital sternite broadly U-shaped with each posterior
corner extended posteriorly process-like; second valvulae as in morona.

REMARKS. In external appearance this species can be distinguished by its short head and relatively
short and broad clypeus. It is sympatric with dulita and longiseta over part of its range in
Sarawak and with apicalis in its single known locality in Sabah.

DISTRIBUTION. Borneo (Sarawak, Sabah).

MATERIAL EXAMINED

Holotype <£, Borneo: Sarawak, Gunong Mulu Nat. Park, Long Pala, base camp, 1978 (Collins) (BMNH,
London)

Paratypes. Borneo: 6 <$, 3 $, same data as holotype; 69, Sarawak, foot of Mt Dulit, junction of R. Tinjar &
Lejok, light-trap/at light in house, 1, 2, 26.ix., 3, 31.X.1932 (Hobby & Moore) (BMNH, London); 1 & Sabah,
Tawau, Quoin Hill, light-trap, 3-7.vii.1962 (Holtmann) (BPBM, Honolulu).

Parohinka phiUppina sp. n.

(Figs 119-123)

Length: $, 6-6-7-3 mm, mean 7-0 mm; 9, 8-4-8-7 mm, mean 8-6 mm.

Pale to greenish yellow. Vertex and dorsal region of face faintly mottled with brown, or markings absent.
Fore wings with or without a brown spot at apex of both clavus and claval veins.

External characters as in morona but vertex longer, medial length 2-7-3-5 times length next to eyes and
1-0-1-1 times basal width between eyes.

Male genitalia as in dulita but aedeagus with shaft of similar width throughout length in lateral aspect,
processes short with lateral pair directed more anteriorly.

Female genitalia similar to brevicephala but second valvulae broader with distance from apex of first
dorsal tooth to dorsal hyaline region relatively long.

REMARKS. The elongate head of this species is similar to that found in apicalis and malayensis and
slightly longer than in specimens of longiseta seen from the same localities in the Philippines. The
male genitalia are similar to those of dulita and brevicephala but differ in the shape of the
aedeagus as noted above and from dulita by the shorter and broader apical process of the style in
medial aspect. In the female genitalia philippina can be distinguished by the relatively long
distance from the apex of the first dorsal tooth to the dorsal hyaline region of the second valvulae.

DISTRIBUTION. Philippines (Luzon)

MATERIAL EXAMINED

Holotype £, Philippines: Luzon, Ifugao Prov., Liwo, 8 km E. of Mayoyao, 1000-1300 m, light-trap,
l-3.vi.1967 (Torrevillas) (BPBM, Honolulu)

Paratypes. Philippines: 9<$, 89, same data as holotype except, 29.v-l.vi.1967, 7,8,ll,12.vi.l967; 2$,
Luzon, Ifugao Prov., Jacmal Bunhian, 24 km E. of Mayoyao, 800-1000 m, light-trap, 30.iv., 9- 10. v. 1967
(Torrevillas) (BPBM, Honolulu; BMNH, London).

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

67

120

124

125

Figs 119-128 Parohinka species. 119-125. P. philippina. (119, 120) aedeagus, lateral and posterior
views; (121) second valvulae, lateral view; (122) apex of left style, ventromedial view; (123) left style,
ventral view. 124, 125. P. recurva, aedeagus, lateral and posterior views. 126-128. P. apicalis. (126)
connective, dorsal view; (127, 128) aedeagus, lateral and posterior views.

Parohinka recurva sp. n.

(Figs 124, 125)

Length: c?, 6-6 mm.

Pale yellow. Vertex, dorsal region of face and pronotum mottled with brown ; other markings as in generic
description.

External characters as in morona but vertex longer, medial length 4-0 times length next to eyes and 1 -4
times basal width between eyes.

Male genitalia similar to dulita with aedeagal shaft of similar width from base to near apex in lateral
aspect, apical region narrower, curved laterally and ventrally and terminating in a moderately long ventrally
directed process on one side; a slightly curved, antero ventrally directed process arising laterally against
posterior margin slightly basad of narrow apical region of shaft; gonopore apical, adjacent to apical process.

Female genitalia unknown.

REMARKS. This species can be distinguished by its elongate head and ventrally curved apex of the
aedeagus with one apical and one lateral process. It is sympatric with apicalis and longiseta (both
with shorter heads) in its single known locality in Sarawak.

DISTRIBUTION. Known only from a single specimen from Borneo (Sarawak).

MATERIAL EXAMINED

Holotype <J, Borneo: Sarawak, Kuching, Matang, 450-894 m, m.v. light-trap, 15.ix.1958 (Gressitt & Mad)
(BPBM, Honolulu).

68 M. D. WEBB

Parohinka apicalis sp. n.

(Figs 126-128)

Length:^, 5-6-7-1 mm, mean 6-6 mm; $,8-0 mm.

Colour and external characters as in philippina with medial length of vertex 2-7-3-7 times length next to
eyes and 1 -2 times basal width between eyes.

Male genitalia with pygophore elongate, as in longiseta (Fig. 131). Xth segment extending to approxi-
mately level with posterior margin of pygophore. Connective very long, slightly upturned apically. Style
with apical process slightly expanded. Aedeagus with shaft evenly tapered from base to apex in lateral
aspect, compressed slightly anteroposteriorly with two short apical processes on one side, directed dorsally;
gonopore apical, on one side, adjacent to processes.

Female genitalia unknown.

REMARKS. The elongate head of this species is similar to that of philippina and malayensis and
slightly shorter than in recurva. In the male genitalia apicalis differs from philippina and mal-
ayensis in having two aedeagal processes and from recurva by having the apex of the aedeagus
directed dorsally. Two female specimens, one from Sabah, Quoin Hill (BPBM, Honolulu) and
one from Sarawak, Gunong Matang (BMNH, London) may be this species but differ slightly
from one another in the shape of the genitalia.

This species is sympatric with brevicephala and longiseta (both with shorter heads) in Sabah
and with longiseta and recurva in Sarawak.

DISTRIBUTION. Borneo (Sarawak, Sabah).

MATERIAL EXAMINED

Holotype <$, Borneo: Sarawak, Kuching, Matang, 450-894 m, m.v. light-trap, 15.ix.1958 (Gressitt & Mad)
(BPBM, Honolulu).

Paratypes. Borneo: 1 £, Sarawak, Gunong Matang, 120 m, m.v. light-trap, 16.ix.1958 (Gressitt & Mad)
(BMNH, London); 1 <$, Sabah, Tawau, Quoin Hill, 1 5-20. vii. 1962 (Hirashimd) (BPBM, Honolulu).

Parohinka longiseta (Melichar) comb. n.

(Figs 129-143)
Muirella longiseta Melichar, 1914: 135. LECTOTYPE & JAVA (RNH, Leiden), here designated [examined].

Length: $, 5-1-6-3 mm, mean 5-7 mm; $,5-7-6-3 mm, mean 6-0 mm.

Pale yellow to stramineous, sometimes tinged with green. Head mottled with brown, stramineous or
green, or markings absent; clypeus often patched with stramineous to reddish brown laterally. Forewings
sometimes with apex and a transverse band at mid length, brown (see 'Remarks' below); other markings as
in generic description.

External characters as in morona but vertex shorter, medial length 1-5-2-6 times length next to eyes and
0-5-0-9 times basal width between eyes. Face with upper medial keel faint in shorter-headed forms; clypeus
elongate. Male genitalia as in generic description with pygophore and Xth segment elongate, the latter
extending considerably beyond posterior margin of pygophore. Connective with stem elongate, slightly
upturned apically. Style with apical process expanded. Aedeagus with shaft evenly curved and tapered from
base to apex in lateral aspect with a single apical process directed dorsally; often apex of shaft surrounding
gonoduct membranous; few faint longitudinal ridges on posterior surface of shaft and variably on apical
process ; gonopore apical, adjacent to apical process.

Female genitalia as in brevipennis, with extended posterior corners of pregenital sternite short to long.

REMARKS. This is the most widespread species of the genus and sympatric with all six other
species in the dulita-group over parts of its range from the Philippines to Sarawak. Specimens
with shorter heads are similar to brevicephala but have the clypeus and extended posterior corners
of the pregenital sfernite longer. The male genitalia are similar to malayensis but differing in the
longer pygophore, Xth segment and connective.

In the male genitalia the apical process of the aedeagus shows some variation in shape (see Figs
136-138) and a few specimens from Malaya have the fore wings conspicuously marked with
brown, as noted above; these specimens also have the markings on the head darker brown and
the vertex shorter than in other specimens.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE

129 / \ 131

69

Figs 129-148 Parohinka species. 129-143. P. longiseta. (129, 130) head and pronotum, dorsal view;
(131) c£ pygophore and Xth segment, lateral view; (132) apex of left style, medial view; (133) second
valvulae, lateral view; (134) $ Xth segment, dorsal view; (135) aedeagus, posterior view; (136) apex of
aedeagus, posterior view, Java; (137) same, Sarawak, Paya Paloh; (138) same, Malaya, Selangor;
(139) apex of $ abdomen, ventral view; (140) left style, ventral view; (141, 142) connective, lateral and
dorsal views; (143) aedeagus, lateral view. 144-148. P. malayensis. (144) head and pronotum, dorsal
view; (145) connective, dorsal view; (146) aedeagus, posterior view; (147)$ pregenital sternite, ventral
view; (148) second valvulae, lateral view.

70 M. D. WEBB

DISTRIBUTION. Nepal, extreme north-east of India eastwards to the Philippines and south to Java
(excluding Borneo).

MATERIAL EXAMINED

Muirella longiseta Melichar, lectotype <$, Java : Wonosobo, v.1909 (Jacobson) (RNH, Leiden).

Nepal: 1 <J, Adhabbar, Godavari, 1500 m, 5.viii.l967 (CNC, Ontario); 1 9, Kathmandu, 1300-1400 m,
7-12.V.1966 (Sedlacek) (BPBM, Honolulu). India: 1 <J, W. Bengal, Darjeeling, 9.xi.l969 (Das) (BMNH,
London). Thailand: 1 <J, Chieng Mai Prov., Doisuthep, 100 m, at light, 18.vi.1965 (Miyatake) (ELKU,
Fukuoka). Philippines: 17 <J, 4$, Luzon, Ifugao Prov., Liwo, 8 km E. of Mayoyao, 1000-1300 m, light-trap,
30.v-12.vi.1979 (Torrevillas); 1 <J, Ifugao Prov., Jacmal Bunhian, 24 km E. of Mayoyao, 800-1000 m,
light-trap, 27-29.iv.1967 (Torrevillas); 1 & Mindanao, Mt Balatukan, 15 km SW. of Gingoog, 1000-2000 m,
at light, 27-30.iv.1960 (Torrevillas); 1 9, M., Del Sur, 32 km NW. of Milbuk, 900 m, rain forest, light-trap,
6.viii.l958 (Milliron) (all in BPBM, Honolulu). Malaya: 2 £, 1 ?, Gomak For. Res. near Kuala Lumpur,
28.xii.1958 (Quote); 1 & 2 ?, Pahang, Kuala Tahan, 15-16.xii.1958 (Quate); 3 rf, 2$, Kuala Lumpur, Klang
gates, 31.xii.1958 (Quate) (all in BPBM, Honolulu); 1 specimen (abdomen missing), Kuala Lumpur, at light,
6.ii.l931 (Pendlebury); 3 9, West Coast, P. Angsa, Lt. Ho., at light, 26-27.X.1926 (Seimund); 1 <J, Kedah, near
Jitra, catchment area, 8.iv.l928 (Pendlebury); 1 cJ, Selangor, Kepong, 7.ii.l958, at light; 2 specimens (abdo-
mens missing), Perak, G. Kledang, 795 m, 15.xi.1927 (Seimund) (all in BMNH, London). Singapore: 14 J, 5$,
Mandaj, mangrove, 7.ix.l977 (Murphy) (BMNH, London). Borneo: 1 £, Sarawak, Paya Paloh, at light,
4.xi.l964 (Rothschild); 2 9, Sarawak, Mt Dulit, 22 m, moss forest, light- trap, 28.x. 1932 (Hobby & Moore); IJ,
Sarawak, Gunong Mulu Nat. Park (Collins); 1 ^, Sarawak, Gunong Mulu Nat. Park, base camp, at light,
v.1978; 1 9, Sarawak, Kalabit End (Mjorberg) (all in BMNH, London); 1 <J, Sarawak, Sarikei Dist., Rejang
delta, 15-25.vii.1958 (Maa); 1 £, Sarawak, Matang, m.v. light-trap, 13.ix.1958 (Gressitt) (both in BPBM,
Honolulu); 1 <$, Sabah, Kalabakan, light-trap, 10-19.xi.1958 (Quate); 1 $, Sabah, Kalabakan, primary forest,
ll.xi.1958 (Maa); 1 9, Sabah, Tawau, Quoin Hill Cocoa Res. Sta., 20.viii.1962 (Hiroshima); 1 9, Sabah,
Tawau, Quoin Hill, light-trap, 1 5-20. vii. 1962 (Holtman) (all in BPBM, Honolulu); 1 9, Sabah, R. Kara-
muak, 7 m SSE. Telupid, 60 m, l-7.ix.1977 (Bacchus) (BMNH, London). Java: 1 & Batavia, xi.1908
(Jacobson) (RNH, Leiden) (paralectotype of Muirella longiseta Melichar).

Parohinka malayensis sp. n.

(Figs 144-148)

Length: <$, 6-2 mm; 9, 6-9 mm.

Sordid to brownish yellow; vertex and dorsal region of face mottled with brown; other markings as in
generic description.

External characters as in morona but vertex longer, medial length 3-2 times length next to eyes and 1-2
times basal width between eyes.

Male genitalia similar to longiseta but pygophore and Xth segment shorter, the latter extending only
slightly beyond posterior margin of pygophore, connective with stem short rather than long and aedeagus
with shaft more robust.

Female genitalia similar to longiseta with extended posterior corners of pregenital sternite short; second
valvulae relatively narrow with first dorsal tooth large.

REMARKS. This species is closely related to longiseta but differs in its longer head and in the shape
of the male genitalia, as noted above.

DISTRIBUTION. Known only from the type-locality in Malaya.

MATERIAL EXAMINED

Holotyped1, Malaya : West Coast, P. Angsa, Lt. Ho., at light, 1 1.x. 1926 (Seimund) (BMNH, London).
Paratypes. Malaya: 1 cJ, 1 9, same data as holotype except, 12/1 6.x. 1926 (BMNH, London).

KAROSEEFA gen. n.

Type-species: Karoseefa brevipenis sp. n.

Yellow tinged with green.

Head wider than pronotum; anterior margin rounded in profile, transversely striate; ocelli on margin
distant from eyes, not or slightly visible from above; anterior tentorial branches curved anteriorly, not
bifurcate. Vertex triangularly produced, medial length slightly less than twice length next to eyes; sides

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 7 1

slightly convex; apex moderately angularly rounded; finely longitudinally striate, transversely striate an-
teriorly. Face slightly wider than long, convex in profile, shagreen ; clypeus relatively short and broad, lateral
margins slightly constricted near antennae; clypellus moderately long, of similar width throughout length;
lora large ; antennal pit deep with inner margin evenly rounded to clypeus ; antennal ledge slight ; antennae
very long, extending to near apex of clavus. Pronotum approximately twice as wide as long, sides very short,
without a carina; transversely striate, obscurely rugose anteriorly. Scutellum approximately equal in length
to pronotum, shagreen, obscurely rugose posteriorly. Fore wings with three subapical cells, first subapical
cell open, second and third subapical cells closed ; with a few additional veinlets in subcostal region near to
fifth apical cell. Fore tibia with dorsal setal arrangement 1:4; fore femur with a series of 10 fine setae distally
on anterior surface; hind femur with apical setal formula 2+1-1-1 with first proximal seta slender.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore strongly incurved dorsally with a short apodeme on
each side; pygophore lobes with an oblique internal ledge extending to posterior margin; lobes with several
macrosetae and short fine setae. Xth segment fairly short, compressed dorsoventrally. Valve triangular.
Subgenital plate elongate, triangular, evenly tapered to apex or with distal region digitate and lightly
sclerotized; apex of basal lobe with a short acute prominence; outer margin of dorsal surface with several
moderate to long fine setae; ventral surface with few short fine setae with or without numerous long fine
setae along outer margin; apical region of dorsal and ventral surfaces with several short stout setae. Style
moderately long with basal apophyses and lateral lobe prominent; apical process moderately long, curved
ventrally and tapered to apex, crenulate dorsally or dentate ventrally; few sensory papilla dorsally, adjacent
to lateral lobe. Connective Y-shaped, stem moderately long and narrow, lateral margins keel-like dorsally;
arms short. Aedeagus with shaft short or long, narrow, tapered from base to apex and terminating in a pair
of dorsally directed processes, gonopore small, situated apically; basal apodeme moderately long and
narrow.

Female genitalia with posterior margin of pregenital sternite with one or two lobes each side of mid line;
second valvulae united at first dorsal tooth (arrowed in Fig. 160), elongate, of similar width throughout
length, without a basal prominence; dorsal teeth robust, unaligned, extending approximately one-quarter
distance from apex to base of valvulae; dorsal sclerotized region elongate.

REMARKS. This genus can be distinguished from others of the subfamily by the broad clypeus and
clypellus with sides parallel rather than concave. In the male genitalia it is similar to Dryado-
morpha but the anterior margin of the pygophore is strongly incurved dorsally with a short
apodeme on each side. The female genitalia have the second valvulae similar to those of Pa-
rohinka and Rhutelorbus.

DISTRIBUTION. Borneo (Sarawak, Sabah).
Key to species of Karoseefa

1 Aedeagus short with apical processes more or less parallel ; posterior margin of female genital

sternite with two lobes each side of mid line (Fig. 161) .... brevipenis sp. n. (p. 70)

Aedeagus long with apical processes divergent; posterior margin of female pregenital sternite
with a single lobe each side of mid line (Fig. 163) divergent sp. n.(p. 73)

Karoseefa brevipenis sp. n.

(Figs 149-161)

Length: <$, 5-2-5-6 mm, mean 5-4 mm; 9, 5-5-5-8 mm, mean 5-6 mm.

Colour and external characters as in generic description.

Male genitalia as in generic description with apex of subgenital plate digitate and lightly sclerotized,
apical process of style dentate dorsally and aedeagus relatively short with processes more or less parallel.

Female genitalia with posterior margin of pregenital sternite with two lobes each side of mid line, the
more lateral lobe acute.

REMARKS. I have identified only specimens of uniform colour as this species, a female specimen
from Sabah : N. of Kalabakan (BPBM, Honolulu) may also be this species but has the fore wings
banded with brown.

In external appearance this species is almost identical to divergens and is sympatric with this
species over its known range in Sabah. From divergens it can be distinguished by its slightly

M. D. WEBB

160

161

162

164

166

Figs 149-166 Karoseefa species. 149-161. K. brevipenis. (149) head and thorax, dorsal view; (150)
same, lateral view; (151) face; (152) left subgenital plate and apex of left style, dorsal view; (153)^
genitalia capsule; (154) apex of left style, medial view; (155) left style, ventral view; (156) connective,
lateral view; (157) same, dorsal view. (158, 159) aedeagus, lateral and posterior views; (160) second
valvulae, lateral view; (161) $ pregenital sternite, ventral view. 162-166. K. diver gens. (162) left style,
ventral view; (163) 9 pregenital sternite, ventral view; (164) valve and left subgenital plate, lateral
view; (165, 166) aedeagus, lateral and posterior views.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 73

smaller size and female pregenital sternite with two lobes each side of the mid line. In the male
genitalia brevipenis differs from divergens in having the apical region of the subgenital plate
narrower, the ventral surface of the subgenital plate without long fine setae, the apical process of
the style dentate and the aedeagus shorter with the apical processes parallel.

DISTRIBUTION. Borneo (Sarawak, Sabah).

MATERIAL EXAMINED

Holotype cJ, Borneo: Sarawak, foot of Mount Dulit, junction of R. Tinjar and Lejok, old secondary forest,
at light, 29.viii.1932 (Hobby & Moore) (BMNH, London).

Paratypes. Borneo: 7 $, 10 $, same data as holotype, except 28.viii-4.x.l932; 1 9, Sarawak, Mount Dulit,
moss forest, 1200 m, 2 1.x. 1932 (Hobby & Moore) (BMNH, London); 1 & Sabah, Bettotan, near Sandakan,
31.vii.1927 (BMNH, London); 1 ?, Sabah, Tawau, Quoin Hill, at light, 26-29. vii. 1962 (Holtmann) (BPBM,
Honolulu).

Karoseefa divergens sp. n.

(Figs 162-166)

Length: $, 6-0 mm; ?, 6-5-6-7 mm, mean 6-6 mm.

Colour and external characters as in generic description.

Male genitalia as in brevipenis but subgenital plate evenly tapered to apex and ventral surface of
subgenital plate with numerous long fine setae laterally. Style with apical process less robust and crenulate
dorsally and aedeagus with processes divergent.

Female genitalia with posterior margin of pregenital sternite with a single lobe each side of mid line.

REMARKS. This species is almost identical to brevipenis externally and is sympatric with this
species over its known range in Sabah. For differences between the two species see 'Remarks'
under brevipenis (p. 71).

DISTRIBUTION. Borneo (Sarawak, Sabah).

MATERIAL EXAMINED

Holotype <J, Borneo: Sarawak, Gunong Mulu Nat. Park, Long Pala, 70 m, alluvial secondary forest, on
batu canopy, at light, iii.1978 (Holloway) (BMNH, London).

Paratypes. Borneo: 1 $, Sarawak, Gunong Mulu Nat. Park, near Long Pala, 50 m, alluvial forest,
understorey, at light, v.1978 (Holloway) (BMNH, London); 1 $, Sabah, Tawau, Quoin Hill, at light,
3-7.vii.1962 (Holtmann) (BPBM, Honolulu).

OCEANOPONA Linnavuori

Oceanopona Linnavuori, 1960: 300. Type-species: Oceanopona croceipennis Linnavuori, by original de-
signation.

Yellow to stramineous; male with thorax and fore wing marked with brown.

Head wider than pronotum; anterior margin narrowly rounded in profile, transversely striate; ocelli on
margin distant from eyes, visible from above; anterior tentorial branches curved anteriorly, not bifurcate.
Vertex triangularly produced, medial length approximately 1-7 times length next to eyes, sides slightly
convex, apex narrowly angularly rounded; shagreen and obscurely rugose, transversely striate anteriorly.
Face approximately as long as wide, shagreen; more or less straight in profile; clypeus elongate, lateral
margins constricted near antennae; clypellus elongate, expanded apically; transclypeal suture indistinct;
lora large; antennal pit deep with inner margin angularly rounded to clypeus, rim-like; antennal ledge
slight; antennae very long, extending to near apex of clavus. Pronotum approximately twice as wide as long,
sides very short, without a carina; finely transversely striate, obscurely rugose anteriorly. Scutellum approxi-
mately equal in length to pronotum, shagreen. Fore wings with three subapical cells, first subapical cell
open, second and third subapical cells closed. Fore tibia with dorsal setal arrangement 1:4; fore femur with
a series of six fine setae distally on anterior surface; hind femur with apical setal formula 2+1+0.

Apodemes of male third abdominal segment ventral, reduced.

Male genitalia with anterior margin of pygophore slightly incurved dorsally, without apodemes; pygo-
phore lobes with a slight oblique internal ledge, not attaining posterior margin, lobes with several macros-
etae and short to moderately long fine setae. Xth segment moderately long, compressed dorsoventrally.

74

M. D. WEBB

178

Figs 167-178 Oceanopona croseipennis. 167, head and thorax, dorsal view; 168, same, lateral view;
169, face; 170, $ genital capsule, lateral view; 171, fore wing; 172, valve and left subgenital plate,
dorsal view; 173, connective, lateral view; 174, left style, ventral view; 175, valve and left subgenital
plate, ventral view; 176, connective, dorsal view; 177, 178, aedeagus, lateral and posterior views.

Valve triangular. Subgenital plate elongate, triangular, outer margin of dorsal surface with several short fine
setae and a group of long fine setae from mid length to near apex ; ventral surface with several short fine
setae. Style moderately long with lateral lobe and basal apophyses prominent; apical process moderately
long, turned ventrally with sides parallel, crenulate distally; few sensory papilla ventrally adjacent to lateral
lobe. Connective Y-shaped, stem moderately long, narrow with lateral margins keel-like dorsally; arms with
apophyses relatively long. Aedeagus with shaft elongate, curved dorsally, evenly tapered to apex and
terminating in a pair of dorsally directed processes; gonopore apical on posterior surface, small; basal
apodeme moderately long, narrow.

THE ASIAN, AUSTRALASIAN AND PACIFIC PARABOLOPONINAE 75

Female genitalia with second valvulae united at first dorsal tooth, fairly robust, slightly expanded distally,
dorsoanterior prominence present; teeth fine, extending along approximately distal third of valvulae; dorsal
sclorotized region moderately long.

REMARKS. This genus is similar externally to Dryadomorpha and Parohinka but with the vertex
shagreen and obscurely rugose and the setal formula of the hind femur 2+1+0. The male
genitalia are similar to those of Dryadomorpha and Karoseefa but with the internal ledge of the
pygophore lobes not extending to the posterior margin and the connective with the apophyses of
the basal arms relatively long. In the female genitalia the second valvulae are similar to those of
Dryadomorpha but with the teeth extending along approximately the distal third of valvulae.

The specimen of Oceanopona recorded by Evans (1966: 247) from Australia is identified as
Dryadomorpha metrosideri (Osborn).

DISTRIBUTION. Eastern Caroline Is.

Oceanopona croceipennis Linnavuori

(Figs 167-178)
Oceanopona croceipennis Linnavuori, 1960: 301. Holotype cJ, CAROLINE Is. (BPBM, Honolulu) [examined].

Length: <$, 4-4-4-7 mm, mean 4-6 mm; 9, 5-9 mm.

Yellow or stramineous. Male with pronotum brown posteriorly; scutellum turning to brown anteriorly
with basal triangles darker brown ; legs heavily marked with brown ; fore wings golden to yellow, sometimes
tinged with green, with a broad dark brown band along clavus, on apical region of subapical cells and
becoming slightly paler on apical cells and appendix. Female variably tinged with green, without brown
markings.

External characters as in generic description.

Male genitalia as in generic description with lateral lobe of style short and angularly rounded in ventral
aspect. Aedeagus with apical processes divergent, short.

Female genitalia with posterior margin of pregenital sternite shallowly concave.

DISTRIBUTION. Eastern Caroline Is.

MATERIAL EXAMINED

Oceanopona croceipennis Linnavuori, holotype (J, Caroline Is.: Ponape, Mt Tamatamansakir, 180 m,
17.U953 (Gressitt) (BPBM, Honolulu).

Caroline Is.: 4 & 1 ?, Ponape, Mt Tamatamansakir, 180 m, 15-19.U953 (Gressitt) (BPBM, Honolulu)
(paratypes of Oceanopona croceipennis Linnavuori); 1 <$, Kusaie I., Matanluk, Yepan, 23.U953 (Gressitt)
(BPBM, Honolulu) (paratype of Oceanopona croceipennis Linnavuori).

References

Bruner, L. 1908. The Acridiidae. Biologia cent.-am. Insecta. Orthoptera 2: 1-342.

Distant, W. L. 1917. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership

of Mr. J. Stanley Gardiner, M.A. Vol. VI, No. vii.-Rhynchota, Part II: Suborder Homoptera. Trans. Linn.

Soc.Lond. 17: 273-322.
- 1918. In Blandford, W. T., The fauna of British India including Ceylon and Burma. Rhynchota 7.

Homoptera: appendix. Heteroptera: addenda. vii + 2 10 pp. London.
DIabola, J. 1979. Neue Zikaden aus Anatolien, Iran und aus Sudeuropaischen Landern (Homoptera:

Auchenorrhyncha). Acta zool. hung. 25: 235-257.
Evans, J. W. 1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera:

Cicadelloidea and Cercopoidea). Mem. Aust. Mus. 12: 1-347.
Eyles, A. C. & Linnavuori, R. 1974. Cicadellidae and Issidae (Homoptera) of Niue Island and material from

the Cook Islands. N. Z. Jl Zool. 1 : 29-44.
Gardner, J. A. 1916. Systematic paleontology of the Upper Cretaceous deposits of Maryland. Mollusca. Md

geol. Surv. stratigr. Mem. 1916: 371-734.
Hamilton, K. G. A. 1975. Review of the tribal classification of the leafhopper subfamily Aphrodinae

(Deltocephalinae of Authors) of the Holarctic region (Rhynchota: Homoptera: Cicadellidae). Can. Ent.

107:477^98.

76

M. D. WEBB

Haupt, H. 1927. Homoptera Palestinae I. Bull, agric. Exp. Stn, Tel- Aviv 8: 5-43.

Ishihara, T. 1954. Homopterous notes. Sclent. Rep. Matsuyama agric. Coll. 14: 1-28.

Kirkaldy, G. W. 1906. Leaf-hoppers and their natural enemies. (Pt. IX. Leaf-hoppers — Hemiptera). Bull.

Hawaiian Sug. Pirs' Ass. Exp. Stn (Ent.) 1 : 271^479.

— 1907. Leaf-hoppers — supplement. (Hemiptera). Bull. Hawaiian Sug. Firs' Ass. Exp. Stn (Ent.) 3: 1-186.
Lindberg, H. 1958. Hemiptera Insularum Caboverdensium. Commentat. biol. 19: 1-246.
Linnavuori, R. 1960. Insects of Micronesia. Homoptera Cicadellidae. Insects Micronesia 6: 231-344.

- 1978. Revision of the Ethiopian Cicadellidae (Homoptera). Paraboloponinae and Deltocephalinae :
Scaphytopiini and Goniagnathini. Revue zool. afr. 92: 457-500.

Matsumura, S. 1912. Die Acocephalinen und Bythoscopien Japans. J. Coll. Agric. Hokkaido imp. Univ. 4:

279-325.
Melichar, L. 1914. Homopteren von Java, gesammelt von Herrn Edw. Jacobson. Notes Leyden Mus. 36:

91-147.

Merino, G. 1936. Philippine Cicadellidae (Homoptera). Philipp. J. Sci. 61 : 307^400.
Osborn, H. 1934. Cicadellidae of the Marquesas Islands. Bull. Bernice P. Bishop Mus. 114: 239-269.
Pruthi, H. S. 1930. Studies on Indian Jassidae (Homoptera). Part I. Introductory and description of some

new genera and species. Mem. Indian Mus. 11 : 1-68.

- 1934. Entomological investigations on the spike disease of sandal. (14) Jassidae (Homopt.). Indian
Forest Rec. 19:1-30.

Uhler, P. R. 1896. Summary of the Hemiptera of Japan presented to the United States National Museum by

Professor Mitzukuri. Proc. U.S. natn. Mus. 19: 255-297.
Webb, M. D. 1980. The Cicadellidae from Aldabra, Astove and Cosmoledo Atolls collected by the Royal

Society Expedition 1967-68 (Hemiptera, Homoptera). J. nat. Hist. 14: 829-863.

Index

Invalid names are in italics; references to descriptions are in bold.

anacryon 40, 50
antennalis 40, 50
apicalis 58, 59, 66, 67, 68
australis 51

brevicephala 58, 59, 64, 66, 68
brevipenis 70, 71, 73

Calotettix 39, 49
camphorae 39, 40, 48
chatterjeei 40, 50
chinensis 43, 45, 46
croceipennis 39, 73, 75

divergens 71, 73
Dryadomorpha 40, 42, 49, 50,

57
dulita 58, 59, 64, 66, 67, 68

falcata 40

Favintiga 40, 42, 47
fraternus 40, 50

guttata 39, 43, 45, 46

indicus 40, 50
ishihari 43, 45, 46

Khamiria 49

Karoseefa 42, 50, 58, 70, 73

lais 39

longiseta 40, 57, 58, 59, 64, 66,

67, 68, 70
lotophagorum 40, 58, 59, 61,

63,64
lugubris 50
luzonensis 43, 46

malayensis 58, 59, 68, 70
mangrovecola 50
merinoi 56, 57

metrosideri 39, 50, 53, 54, 75
modestus 39

morona 58, 59, 60, 61, 63, 64,
66, 67, 70

Oceanopona 39, 42, 58, 73, 75
Odmiella 40, 42
Osbornitettix 49

pacifica 50, 53
Paganalia 40, 49
pallida 40, 50, 54, 55
Parabolopona 39, 40, 42, 47
Parohinka42, 50, 57, 71, 73
philippina 58, 59, 66, 68
punctatus 40, 50

quadricornis 40, 50

recurva 58, 59, 67, 68
Rhombopsana 40
Rhombopsis 40, 49
Rhutelorbus 42, 50, 56, 58, 71
robustipenis 50, 54, 55
rubrolineata 39

sinuata 58, 59, 61, 63, 64
spinosa 58, 59, 63, 64
Stenomiella 40, 42
Stirellus 50
Stymphylus 39

tincta 39

trispicata 58, 59, 61, 63, 64

virens 40, 50
virescens 40
viridia 54
viridis 40, 50

Yakunopona 40, 49
yakushimensis 40, 51

Zizyphoides 40, 49, 50

British Museum (Natural History)
1881-1981

Centenary Publications
Chance, change & challenge

Two multi-author volumes from one of the foremost scientific institutions in the world.
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A revision of Phyciodes HUbner and relate
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L. G. Higgins

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ISSN 0524-6431 Entomology series

Vol 43 No 3 pp 77-243
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 29 October 1981

/*

A revision ofPhyciodes Hiibner and related genera, with
a review of the classification of the Melitaeinae
(Lepidoptera: Nymphalidae)

L. G. Higgins

Focklesbrook Farm, Chobham, Woking, Surrey

Contents

Synopsis 78

Introduction 78

Historical review 78

Abbreviations of depositories ........... 79

Type-material and the British Museum (Natural History) collections ... 79

Phyciodine genitalia and the identification keys 80

Examination of Phyciodine genitalia 80

Taxonomy 81

Phyciodini trib. n. 81

Tribal characters 81

Distribution 82

Wing patterns 83

Key to genera of Phyciodini 84

Phyciodes Hiibner 85

Phystis gen. n. 93

Anthanassa Scudder 94

Dagon gen. n 108

Telenassa gen. n. . 1 10

Ortilia gen. n .115

Tisona gen. n 120

Tegosa gen. n. .121

Eresia Boisduval .129

Castilia gen. n. .151

Janatella gen. n. .157

Mazia gen. n .159

Species incertae sedis . 160

Supplement on certain genera of Melitaeini ...... . 160

Gnathotriche Felder & Felder • 160

Gnathotrusia gen. n. ........ • 162

Higginsius Hemming • 163

Antillea Higgins .164

Review of the classification of the Melitaeinae ... . 164

Check-list of tribes, generic groups, genera and species of the Melitaeinae . . 165
Distribution .......... • 171

Palaearctic region .......

Nearctic and Neotropical regions ....

Acknowledgements .174

References • 175

Index 238

Bull. Br. Mus. Hist. (Ent.) 43 (3): 77-243 Issued 29 October, 1981

78 L. G. HIGGINS

Synopsis

The new tribe Phyciodini is defined and its constituent taxa revised at the generic and specific levels, thus
completing an account of the nymphalid subfamily Melitaeinae commenced by the author over 40 years ago.
An historical account of the taxonomy of the group is included, together with information on type-material,
and the taxonomic and practical methods employed. The geographical distribution and wing patterns
of the 137 included species are discussed. A key to the twelve genera is provided and a key to species is
given for each genus in turn. The account of each species includes a full synonymy of nominal species
and forms, and a brief morphological description and a summary of distribution. A supplementary section
deals with four divergent genera of American Melitaeini previously often confused with the Phyciodini.
Finally, an overview of the author's conclusions concerning the classification of the Melitaeinae is given,
including a complete check-list of the tribes, generic groups and species of the subfamily. The taxonomic
work results in the description often new genera, five new species, and three new subspecies; 135 changes
in combination, one new generic synonym, 35 new specific synonyms, and 38 changes of specific status
are established; three species are raised from synonymy; one new specific name is proposed; and
106 lectotypes are designated.

Introduction

This account of the Phyciodini has two main objectives. The first one is to examine the
characters of this large tribal group of over 130 species, and to analyse the relationships of the
various subgroups of generic status which are defined and named. The second objective,
dependent upon the first, needs explanation. I have had a special interest in the Melitaeinae
for a great many years, and reviews of the Palaearctic species, and of the American Chlosyne
and their allies, have already appeared (Higgins, 1941; 1950; 1955; 1960). The present review of
the Phyciodini, and the accompanying account of the small Melitaeine genera related to
Gnathotriche Felder & Felder, will complete the analysis, by a single worker, of the entire
subfamily, including about 240 species. Apart from a consistent taxonomic treatment, the
comprehensive account could reveal interesting features which may be overlooked when the
series is considered, as is usual, as consisting of independent Eurasiatic and American groups.

Historical review

The history of the Phyciodini goes back to 1819, when Hubner introduced the generic name
Phyciodes for two American species, cocyta Cramer and liriope Cramer, united generically by
the rather unimpressive character of 'reticulate markings' on the undersides of the hindwings.
The new genus was placed by the author in his group of Dryads, a creditable decision at that
early date. Seventeen years later Boisduval (1836) published the name Eresia eunica (recte eunice)
by means of a figure only, erecting a new genus without any description or definition. Soon
afterwards, as interest increased in the butterflies of Central and South America, many new
species were described by various authors including the Felders, Bates, Hewitson, Druce and,
later, Godman & Salvin, using the generic names Melitaea, Eresia, Phyciodes etc. according to
personal preference — since the characters of these genera were not defined.

In fact, definition was not possible, at this date, since characters used in the nineteenth, and
in the early years of the present, century were not adequate for the taxonomic analysis of these
butterflies. Nevertheless, in 1875 Scudder proposed the genus Anthanassa, with somewhat
equivocal characters and a type-species of uncertain identity, evidence that the complex nature
of the tribe was becoming recognised. Seven years later, in 1882, Godman & Salvin made a
distinction between Eresia and Phyciodes, and separated these from the American Chlosyne
(Melitaeini) and their allies, partly on the characters of their genitalia. This useful hint was
not followed up by later authors. Schatz (1892), in his article on the Melitaeinae, separated
Melitaea from Phyciodes, with a note that it was customary to do so, but that there was not any
good distinctive character.

Schatz's careful review provided the basis for Rober's long article on Phyciodes in Seitz
(1913-1914). A similar system was adopted by the late Arthur Hall in his 'Revision of the genus
Phyciodes' (1928-1930).

REVISION OF PHYCIODES 79

In his account Hall recognised only the genus Phyciodes, to include not only Eresia and
Anthanassa, but also numerous American species of Melitaeini such as harrisii Scudder and
its allies, already distinguished and very properly removed from Phyciodes by Godman & Salvin,
Staudinger, Schatz and others. In doing this, Hall made a serious taxonomic mistake, but in
other respects his work was a most important contribution. He had the advantage of knowing
many species in the field, and had easy access to the extensive collections of Rothschild and of
Joicey, and to the collection in the British Museum (Natural History), all containing much
original type-material. With excellent library facilities he succeeded in establishing the com-
plicated synonymies of most species and also assembled the confusing polymorphic and mimetic
phenotypes in an outstanding study which has made his work essential to all later students.
There are occasional references to relationships disclosed by examination of the male genitalia,
but by modern standards this part of his work is not helpful. Revision of Hall's work was
clearly necessary. This was undertaken by W. T. M. Forbes (1945), who made free use of the
genitalia in an improved taxonomic arrangement on modern lines. In this review the North
American species allied to palla are placed in Melitaea, and the Felders' genus Gnathotriche
is retained, while the remaining species are all included in Phyciodes, used in a tribal sense,
and divided into 13 groups with three named subgenera. This paper marks an important advance
in the study of the group, but still fails to provide a generic definition for Phyciodes. Forbes
contented himself with the remark '. . . the boundary against Melitaea is universally agreed to
be indefinite, one might almost say non-existent, if we consider only superficial characters'.
Certainly this extensive group does present difficulties. Natural affinities are so often obscured
by mimetic resemblances to unrelated, naturally protected species. In the present paper the species
retained by Forbes in Phyciodes have been allocated to 12 genera, based principally upon the
characters of the male genitalia.

Abbreviations of depositories

AMNH, New York American Museum of Natural History, New York

BM, Brighton Booth Museum, Brighton

BMNH British Museum (Natural History), London

CAS, San Francisco California Academy of Sciences, San Francisco

CM, Pittsburgh Carnegie Museum, Pittsburgh

CNC, Ottawa Canadian National Collection, Ottawa

FMNH, Chicago Field Museum of Natural History, Chicago

IML, Tucuman Institute Miguel Lillo, Tucuman

MACN, Beunos Aires Museo Argentine Ciencias Naturales, Buenos Aires

MN, Rio de Janeiro Museu Nacional, Rio de Janeiro

MNHN, Paris Museum National d'Histoire Naturelle, Paris

MNHU, Berlin Museum fur Naturkunde der Humboldt-Universitat, Berlin

NM, Vienna Naturhistorisches Museum, Vienna

NR, Stockholm Naturhistoriska Riksmuseet, Stockholm

RSM, Edinburgh Royal Scottish Museum, Edinburgh

UM, Oxford University Museum, Oxford

USNM, Washington National Museum of Natural History, Washington, D.C.

ZI, Leningrad Zoological Institute, Leningrad

Type-material and the British Museum (Natural History) collections

I have never been in Central or South America, and this review is based on the material in the
BMNH, most of which was assembled between the middle and the end of the last century. This
huge collection includes not only the entirety of the Rothschild Collection, but much other
material from Bates, Hewitson, Godman & Salvin, Oberthiir and Joicey. The Rothschild
Collection itself included much material from the Felders, and from Rober. Finally, the BMNH
collection includes many specimens collected by Arthur Hall, who was the last person to
re-arrange the BMNH Phyciodini, although Hall's private collection (together with his
voluminous manuscript notes) passed to the BM, Brighton, Sussex (where I have examined it).

80 L. G. HIGGINS

Thus the BMNH collection includes type-material covering over 75 per cent, of the
phyciodine species recognised at present (including approximately 50 per cent of nominal species).
With this, and much of the other material, the provenance from individual collections is almost
always recorded, sometimes the names of the actual collectors, and often the year of collection,
although very few specimens are accurately dated. As always with material of this vintage, even
the actual locality data are often vague, totally wanting, or even downright misleading; however,
most of the false data labels are now tolerably well known, and their 'information' can be
discarded or re-interpreted.

In their descriptions the early authors sometimes referred to 'types', but the actual specimens
were rarely so-labelled. Throughout this treatment, original specimens have been regarded as
holotypes only where there is clear evidence or a statement in the original description as to
their unique status. In all other cases, where original material has been positively identified and
examined, lectotypes have been designated, in accordance with the suggestions put forward by
Vane- Wright (1975: 26-28). Where original material has not been traced or examined,
information has been included, as far as is available from the original description, as to the
number, sex and status of primary type-specimens, and their provenance and present depository,
where known. With respect to the BMNH types in particular, a great many were previously
listed in A. G. Gabriel's excellent catalogue, published in 1927; wherever possible the present
type designations have been cross-referenced to Gabriel's catalogue and type-specimen numbers.

Phyciodine genitalia and the identification keys

In making this study it has been necessary to examine the genitalia of every available species,
and these are now illustrated, with other anatomical characters, in a series of some 313 figures,
taken from camera lucida drawings made by myself. The original preparations naturally vary
greatly in size, and it has not been possible to make the drawings really uniform in this
respect. In all cases size is indicated by a 1 mm scale line. Keys to genera and species have
been included. Unfortunately external characters are often misleading at the generic level, so the
key to genera is based largely upon genitalic characters. I have not been able to devise for the
species of the whole tribe a single workable key based upon wing markings, size and similar
external features. Keys to the species of each genus are provided, however, based upon external
characters as far as possible, but such keys are only usable after a specimen has been correctly
placed generically. This is easy if the male genitalia are examined. Distinction between closely
similar species may be difficult and the keys will be useful in making the final identification,
with the help of the photographic plates.

Examination of Phyciodine genitalia

Preparation of the male genitalia of the Phyciodini needs special care. The organs are usually
very small, fragile, and unsuitable for dissection. Every care must be taken to avoid distortion,
and the genitalia should be mounted in a shallow cell, free from pressure, in a position which will
give a view of the parts at exactly the same angle in all cases, in order to allow comparison
between different preparations. Owing to the oval shape of the valves, the only position in which
the organs are stable is erect, the dorsal structures upwards, resting on the saccus and the lower
borders of the valves. This position gives a good view of all taxonomically important parts.
The penis should be extracted and mounted to give a lateral (side) view. There is a difficulty
here when a single specimen only is available, as it is easy to damage the dorsal structures
when the penis is removed. Any distortion will destroy the symmetry of the organs and mounting
in the correct position will be difficult. The penis shape in most species is flattened antero-
posteriorly, and manipulation will be required to secure a lateral view in a permanent
preparation. It is often best to leave the penis in position until immediately before mounting, as
it is easily lost if it is removed at an early stage of preparation.

The elaborate sterigma of the female genitalia is displayed if the abdomen is opened along
the dorsum and the lateral walls are spread flat on each side. When the parts are cleaned, care
must be taken to leave the bursa in its correct position, lying along the ventral abdominal plates.

REVISION OF PHYCIODES 81

Prepared in this way and allowed to dry upon the slide for a day or so, the specimen can be
taken off by undercutting with a thin knife when it will remain flat while it is dehydrated and
cleared for mounting in the usual way. A few drops of alcohol will help to loosen the dried
specimen before it is removed from the slide.

Taxonomy

The collections with which I have worked have covered the whole, vast area of tribal
distribution, but representation is uneven. Often the material has been collected from a single
locality, perhaps during a single visit, and so for many species it is not possible to define
distributional frontiers. This leads to taxonomic difficulties as often there is little information
about seasonal variation or possible distributional overlap or clinal series. In the absence of
this information it is only by guessing that the status, local or individual form, seasonal
modification or geographical subspecies, can be decided, when specimens with slightly differing
phenotypical characters are reported flying in widely separated localities. In such cases, when
the differing characters are constant and easily recognisable, the butterflies are treated here as
distinct species, in order to emphasise their individuality. I have found it necessary to
introduce 10 new genera and to describe eight new species or subspecies. A single familiar
name has been found to be invalid. This is Eresia 'c/to' of Aurivillius (1882) and of many
later authors, which certainly is not the species named Papilio clio by Linnaeus, which is probably
an Ithomiine, perhaps Ithomia aegle (F.) as figured in 'Seitz' (pi. 38, row f [Fig. 5]). I understand
from Dr Holm, the Curator of the Linnaean (i.e. Queen Ulrica's) collection at Uppsala, that
the single original specimen of 'clio auctt' is there labelled nauplius, as it is in the figures in
Clerck's Icones.

In the synonymy given for each species, I have indicated when I have examined the type-
material of each nominal taxon by the annotation '[examined]'. Where type-material has not
been examined, the assignment of any name must be regarded in some sense as tentative or
provisional, as the examination of the genitalia is of critical importance in most species.

PHYCIODINI trib. n.

(Figs 178-185)
Type-genus: Phyciodes Hiibner.

In the Melitaeinae, of which the Phyciodini form the largest section, the structure of the
antennae, palpi, legs and the wing venation is almost uniform throughout the subfamily. It has
not been generally recognised that almost all such characters are of subfamily value, but their
uniformity has proved to be a serious difficulty in recognition of the tribal status of the
Phyciodini and for their classification at generic levels. It is in the structure of the genitalia
that characters below the subfamily level are well defined and the individuality of the Phyciodini
is seen to isolate them from other melitaeine tribes. Careful examination of every available
species (about 135) has failed to show any with intermediate or ambiguous tribal features.

Tribal characters

Small or medium-sized butterflies, forewing 11-30 mm (Fig. 178), cell closed by vestigial veins,
vll arises near cell-end except in a few genera (Mazia, p. 159; Castilia, p. 151) in which it arises
at or beyond cell-end. Hindwing (Fig. 179) cell open, precostal vein well developed.

Palpi (Fig. 180) porrect or semi-ascending, terminal segment narrow, middle segment usually
slightly dilated, clothed beneath with long or, in Castilia, very long hair.

Legs of usual nymphaline type, J foreleg greatly atrophied (Fig. 181), tarsus often reduced
to a single segment, tibiae of mid- and hindlegs with slender spines and a single pair of spurs.

Early stages. Characters of ovum, larva and pupa, when known, similar to those of Melitaea;
larval spines in Phyciodes represented by little more than hairy tubercles (P. tharos).

Male genitalia (Figs 182-184). In all melitaeine butterflies the genitalic structure is unusual,
especially in respect of the absence of an uncus and in the fusion of the saccus and juxta to

82 L. G. HIGGINS

form a strongly chitinised basal plate (Higgins, 1941). In the Phyciodini the genitalia show little
variation from this basic type. As discussed above, the organs are .best examined entire, without
any dissection of the valves, which may be confusing, since their lateral walls are rounded and
the absence of a flat lateral surface to support the isolated organ at a constant angle prevents
accurate comparison between different preparations. In most cases examination of a dorsal view
of the intact organs is most valuable, showing the special features of tribal and generic value,
as follows.

Tegumen (A) and scaphial extension (E) (Figs 182, 183). These dorsal structures, taken
together, are almost always well developed, and form an important feature of the organs in
Eresia, Phyciodes, Tegosa and Janatella. This is in contrast with the usual structure in the
Melitaeini, in which the tegumen is commonly small or even vestigial. In the Phyciodini minor
variations in shape and in the development of spinous areas provide generic characters of value.
In many species the lateral walls of the scaphial extension (E) are more or less chitinised
(Figs 182, 183), with the space between covered by a membranous curtain, which may extend as
an inferior layer, sometimes partly chitinised, forming the floor of the anal compartment.

Saccus (S). This structure, fused with the juxta (J), is always well developed. It is often deeply
cleft, as in the Melitaeini, but it is entire (Ungulate) in the genera Eresia, Janatella, Ortilia and
Tegosa (part), often with a small apical notch and associated with unusual development of the
dorsal structures. Among the Melitaeini, a saccus of this type is found only in Didymaeformia
trivia and in Melitaea lukto in its various forms, the former usually considered a divergent relict
species.

Valves (V). Throughout the entire tribe these are oval, elongate posteriorly, with relatively
little variation. Near the apex the costal border often has small teeth which are enlarged in
Anthanassa and in Ortilia, to give the appearance of a bifid apex (Fig. 225). In the genera
Telenassa and Castilia the posterior sections of the valves have a characteristic form, giving a
generic character. The harpe (H) is always well developed, the structure usually simple,
occasionally with a few basal teeth on the underside (Fig. 268).

Penis (P) or aedeagus (Fig. 184). In all species this is straight or nearly so, with a well-defined
morula (M). An ostium-keel (OK) is present in the genera Eresia, Castilia, Janatella and
Phyciodes (part), not so prominent as it is in the melitaeine genus Mellicta, and in Phyciodes
quite small, but large enough in most species to show distant relationship. The penis may be
slender or stout, e.g. in Telenassa (Fig. 297) in which the organ is massive.

Female genitalia (Fig. 185). The structure of the sterigma is distinctive and often elaborate.
The ostium bursae (O) opens between segments 7 and 8, usually at the bottom of a deep pit;
the bursal duct (BD) is partly chitinized and surmounted by a structure that I have called the
bursal support (BS), often egg-shaped, sometimes with three lateral projections or ribs (e.g.
Eresia coela, Fig. 389), to which the bursa copulatrix (B) is attached. This structure, with the
formation of the bursal support, is peculiar to the Phyciodini and forms an excellent tribal
character. The genital plate extends across segment 8 as a modified area named the scutum (S).

Distribution

The Phyciodini are restricted to America, and widely distributed throughout the continent from
about 55°N to 35°S. The species usually occur in localised colonies, and do not appear to have
any tendency to migrate or to wander. Owing to lack of information and of material from
large areas, especially in South America, it is not possible to be precise about distribution
patterns of genera or species, but in general terms I think the following picture is reasonably
accurate.

The genus Phyciodes alone is represented in Canada and in most of the United States,
excepting the fringe areas in the south where some tropical species have penetrated. Anthanassa,
which begins to appear in Mexico and in Arizona, is a large genus developed extensively in
Central America, Colombia and Venezuela, with single divergent species in the east in Argentina
and Brazil (hermas) and in Jamaica and Cuba (frisia). Further south in S. Peru, Bolivia and
in S.W. Argentina, there is Dagon, with three (or four?) rather isolated and small, rare species,
all closely related. The genus Telenassa, with about 16 species, occurs in the mountainous

REVISION OF PHYCIODES 83

western regions of Peru, Colombia and Ecuador. The saccus is deeply cleft in all species. In
eastern South America its place is taken by Ortilia, with 10 species, widely distributed in
C. and S. Brazil, Paraguay, and Argentina, but overlapping the Telenassa area in Peru and
Bolivia. All species have a single (entire) saccus, and this genus is accompanied over much of
the region by Phystis, a monotypic genus, the single small and variable species recalling
Phyciodes in certain respects. The largest genus, Eresia, with about 35 species, includes many
mimetic and polymorphic forms. Although especially characteristic of Central America, it
extends throughout the area south of the United States, with endemic species in the Guyanas
and in NE. Brazil (Amazon). A large genus, Tegosa, including many small species with orange-
yellow wings with dark borders, is widely distributed in Central and South America, and
includes several small and localised species endemic in Ecuador and Peru. Probably closely
related is Tisona, endemic in the Chaco district of Argentina and in Bolivia. The male genitalia
of the single species have several unusual features (Figs 321-323). The genus Castilia includes
about 12 species, of which four are mimetic of Actinote, occurring over a wide area in tropical
Central America, Colombia and Guatemala. In three of these species the venation of the
forewing is atypical. The external features of the remaining species are variable, but good generic
characters are provided by the genitalia. Another small group is Janatella with three species
occurring from Ecuador and Central America to the Guyanas, probably related most closely
to Eresia. Finally, in Mazia, the third monotypic genus, the single species amazonica is divergent
in wing markings, wing venation and in the characters of the genitalia, and although undoubtedly
a member of the Phyciodini, it is difficult to suggest any near relative. The species appears to be
widely distributed over the vast drainage system of the Amazon. In hindsight, the astonishing
concentration of species in Central America, Venezuela and Colombia, is most striking, yet only
one species, Anthandssa frisia, has become established in Cuba and in a few other islands,
to represent the Phyciodini in the Greater Antilles. Phyciodes phaon, which occurs in the
Cayman Islands and probably in Cuba, is an outlier from North America where all its
relationships lie. It appears as an alien among the true fauna of Mexico and Central America.

Wing patterns

On the upperside of the Melitaeinae there is a remarkably constant basic or standard pattern,
with black striae and/or spots in series upon a buff coloured or orange-red ground-colour,
present and immediately recognisable in all Palaearctic and Nearctic species, with the exception
of two euphydryad butterflies, phaeton (both sexes) and cynthia (d only). On the underside of the
hindwing the arrangement of reddish spots in basal and submarginal series is even more constant
and characteristic. This scheme of markings, which appears in all species distributed over the
areas of Quarternary glaciation, must be recognised as an important character of the subfamily
Melitaeinae. In the tribe Phyciodini, the standard markings are present in the species of the
genus Phyciodes, which inhabit principally the temperate northern regions of America, but in
Mexico and further south to Argentina, Bolivia and Peru, the standard markings do not appear,
but are altered in all species by mimicry (Chlosyne) or they become variable (Phyciodini),
sometimes with mimicry, but often with different arrangements characteristic of different
genera (e.g. Tegosa, Ortilia, Anthanassa, etc.).

Mimicry. In Eresia and Castilia many species are mimetic, and wing shapes and markings may be
altered from the usual generic patterns. Heliconiine, ithomiine and acraeine species are the most
frequent models, including such genera as Ceratinia Hiibner, Napeogenes Bates, Ithomia Hiibner,
Eueides Hiibner and Actinote Hiibner, all recognised distasteful groups. In Eresia eutropia,
E. pelonia and E. eunice, polymorphism, especially among females, is so varied that it is impossible
to find a satisfactory name for every form. In all such cases the mimicry appears to be of the
classical Batesian type (pseudaposematic), and much information on this subject, based upon
personal observation, is recorded by Hall (1928-1930). In addition there is a strong tendency
towards mimicry, or to independent development of similar wing markings, between species
in different genera among the Phyciodini, or even between phyciodine species and those of other
unprotected groups. It is difficult to explain the value of repetitive wing markings (or mimicry?)

84 L. G. HIGGINS

of this type, but it may be that the protection afforded by certain simple wing patterns is not
understood. The following examples are especially striking.

Ortilia liriope and Tegosa claudina, confused by everyone until separated by Forbes (1945)

after genitalia examination.
Janatella leucodesma and Dynamine Hiibner spp.
Telenassa abas and Janatella fellula. Wing markings almost identical but genitalia do not

indicate close relationship.
Janatella hera and Castilia ofella. The similarity is remarkable. It seems unlikely that the two

species ever fly together or even in the same regions.

Adams & Bernard (1979) have described some rather similar examples of puzzling convergence
between unrelated but, in this case, definitely sympatric members of the Andean pronophiline
satyrid fauna. One possible explanation involves the concept of arithmetic mimicry, as
originally put forward by Van Someren & Jackson (1959), and discussed by Vane-Wright
(1976: 37-38). Robson & Richards (1936: 260) refer to the earlier observations by Chapman
(1913) and myself (Higgins, 1930) on conceivably similar phenomena among alpine species of
Erebia Dalman in the Palaearctic region.

Key to genera of Phyciodini

1 Antennal club slender, elongate (Fig. 491) MAZIA (p. 159)

Antennal club larger, pyriform (Fig. 227) 2

2 Penis apex with crossing ostium-folds (Fig. 323) TISONA (p. 120)

Penis apex not so formed 3

3 Scaphial extension of tegumen elongate, terminal angles armed with hooks or spines (Fig. 383) 4
Scaphial extension of tegumen small, terminal angles lacking hooks or spines ... 7

4 Upperside of forewing usually yellow, marginal borders and oblique subapical bar black

* TEGOSA (p. 121)
Upperside of forewing not so marked 5

5 Saccus deeply cleft PHYCIODES (p. 85)

Saccus single ................ 6

6 In dorsal view lateral angles of tegumen armed with 3 or 4 strong teeth (Fig. 471) .

JANATELLA (p. 157)
In dorsal view lateral angles of tegumen appearing as rounded, spiculate bosses (Fig. 364)

ERESIA (p. 129)

7 Hindwing upperside submarginal band formed of 5 ocellar spots . . . PHYSTIS (p. 93)
Hindwing upperside submarginal band not so formed, or absent 8

8 Saccus single, more or less expanded to broadly notched apex (Fig. 270) . . DAGON (p. 108)
Saccus not so formed .............. 9

9 Saccus single, narrow or tapering to apex (Fig. 302) ORTILIA (p. 115)

Saccus cleft 10

10 Posterior section of valve short, penis slender ANTHANASSA (p. 94)

Posterior process of valve longer, penis massive 11

11 Posterior section of penis short, about one-third total length, ostium-keel lacking (Fig. 279)

**TELENASSA (p. 110)
Posterior section of penis about half total length, ostium-keel prominent (Fig. 465)

CASTILIA (p. 151)

* In Tegosa the black marginal upperside borders may be extended into wide, dusky suffusions, e.g. T. etia (p. 127);
T. nigrella (p. 128).

** The male genitalia show other distinctive characters but the features of the penis are sufficiently striking for the
purposes of this key.

REVISION OF PHYCIODES 85

PHYCIODES Hubner

Phyciodes Hubner, [1819]: 29. Type-species: Papilio cocyta Cramer [= Phyciodes tharos (Drury)], by
subsequent designation (Scudder, 1872: 46). Gender: masculine.

Small butterflies, forewing outer margin straight or slightly excavate; upperside orange-fulvous marked
with black spots and stripes, recalling Palaearctic species of Melitaeini; hindwing underside a pale yellow
or silvered marginal crescent usually present in s3. Sexes similar, mimetic forms absent.

Genitalia. 3, in dorsal view, tegumen elongate, scaphial extension tapering slightly to a wide apex,
terminating in one or more small hooks at each lateral angle, valve elongate, tapering gradually to an
incurved, pointed apex, preceded by one or more small costal teeth; penis slender, almost straight, with
small ostium-keel and morula. 4 ductus chitinised, rather long, well defined, bursal support elongate
(Fig. 192).

DISTRIBUTION. The species are widely distributed and often common in North America,
especially in the western states of the U.S.A. Four small species, almost restricted to the
southern states, range further south into Mexico and Guatemala. The little P. phaon has spread
from Georgia to the Cayman Islands.

DISCUSSION. The species tharos, batesii, campestris and montanus all have similar markings, are
sometimes difficult to identify, and specific characters in the male genitalia are poorly developed.
In some specimens the scaphial hooks are bent over in a curious way, but without obvious
fracture. It seems possible tkat the distortion happens during life when the tissues are elastic.

Key to species of Phyciodes

Note. It has not been practicable to include P. herlani (p. 90), P. pallidus (p. 89) and P. orseis (p. 90) in this key.
Most species are variable in size. Identification may be difficult, especially in P. mylitta, in which the
black upperside markings may be expanded (rare). It has proved difficult to key out P. montanus, lightly
marked but otherwise resembling P. campestris, and considered a high-level form of the latter by many
authorities. Except for P. vesta, and perhaps for P. mylitta (including herlani, pallidus and orseis), the
male genitalia are not really helpful in making identifications.

1 Two or more hooks at each angle of scaphial extension (Fig. 217) . . . vesta (p. 92)
Single hook at each angle (Fig. 186) 2

2 Small species, o forewing 11-14 mm, markings pale yellow, colour contrast brilliant* . . 3
Larger species, o forewing 15-18 mm, colour contrast slight or absent ..... 4

3 Forewing underside pale apical area unmarked pictus (p. 90)

Forewing underside apical area with linear markings phaon (p. 91)

4 Hindwing underside outer margin with dark shade in s2-s5 tharos (p. 85)

Hindwing underside marginal shade slight or absent ........ 5

5 Forewing upperside base dusky, yellow mark near apex of cell prominent .... 6
Forewing upperside base rarely dusky, yellow mark near apex of cell not prominent

mylittus (p. 88)

6 Forewing underside black costal and discal markings heavy batesii (p. 86)

Forewing underside black costal and discal markings usually vestigial or absent .

montanus (p. 88), campestris (p. 87)

Phyciodes tharos (Drury)
(Figs 1, 178-182, 186-192)

Papilio tharos Drury, [1773]: index to vol. 1; [1770]: 43, pi. 21, figs 5, 6. Syntype(s), U.S.A.: New York

(depository unknown) [not examined].
Papilio morpheus Fabricius, 1775: 530. Syntype(s), NORTH AMERICA ('in America boreali') (presumed lost;

not listed by Zimsen, 1964).
Papilio cocyta Cramer, [1777]: 7, pi. 101, figs A, B. j syntype(s), [NORTH AMERICA] ('Surinam') (presumed

lost).
Papilio euclea Bergstrasser, 1780: 23, [pi. 79, figs 1, 2]. Holotype (sex?) [NORTH AMERICA] ('England')

(d'Orcy) (depository unknown, presumed lost) [not examined].

* In specimens of P. phaon, spring brood, colour contrast may be greatly reduced.

86 L. G. HIGGINS

Argynnis tharossa Godart, [1819]: 289. cJ, 9 syntypes, U.S.A.: New York, etc. (one syntype in RSM,

Edinburgh; Grimshaw, 1898: 4). [There would be good grounds for considering this, and many other

Godart names, as unjustified emendations; in this case, of tharos Drury, to which Godart includes a

reference in his synonymy.]

Melitaea tharos (Drury); Boisduval & LeConte, 1833: 170, pi. 47, figs 3, 4.
Melitaea selenis Kirby, 1837: 289. Syntype(s), CANADA ('North America') (depository unknown; probably

lost).
Melitaea pulchella Boisduval, 1852: 306. Syntypes, U.S.A.: [? New York] ('California') (not in BMNH). [See

Tilden, 1970, concerning the type-material of this taxon.]
Melitaea pharos [sic!] (Drury); Harris, 1862: 289, figs 116, 117.
Melitaea marcia Edwards, [1869]: 207. Lectotype <£, U.S.A.: New York, Hunter, Greene Co. (CM,

Pittsburgh), designated by Brown (1966: 432, fig. 25).

Melitaea packardii Saunders in Packard, 1869: 256. Syntype(s), CANADA (depository unknown).
Phyciodes pascoensis Wright, 1906: 165, pi. 21, figs 198, a. Lectotype <$, U.S.A.: E. Washington, Pasco

(W. G. Wright] (CAS, San Francisco, Type no. 4308), designated by Tilden (1975: 23).
[Phyciodes nycteis (Doubleday); Wright, 1906: 164, pi. 21, fig. 197. Misidentification.]
Phyciodes tharos f. reaghi Reiff, 1913: 305, pi. 10, figs 1, 2. Holotype 9, U.S.A.: Massachusetts, Franklin

(A. L. Reagh) (depository unknown).
Phyciodes tharos (Drury); Rober, 1913: 436, pi. 89, row d [figs 1, 2]; Forbes, 1945: 154, 189; Holland,

1947: 135, pi. 18, figs 1-4, pi. 5, figs 20-22 (pupa).
Phyciodes tharos pascoensis f. vern. herse G. C. Hall, 1924: 110. Holotype 9, CANADA: British Columbia,

Taft (AMNH, New York).
Phyciodes tharos pascoensis f. ab. nigrescens G. C. Hall, 1924: 110. Holotype 9, CANADA: British Columbia,

Taft (AMNH, New York).
Phyciodes tharos tr. f. dyari Gunder, 1928: 167, fig. 18. Holotype cJ, U.S.A.: New Hampshire, Webster

(USNM).
Phyciodes tharos tharos (Drury); Hall, 1928ft: 35; Klots, 1951: 101, pi. 6, fig. 11 (pupa), pi. 13, fig. 14;

Bauer, 1975: 144, pi. 44, figs 6, 8.

Phyciodes tharos f. t. marcia (Edwards); Hall, 19286: 37.
Phyciodes tharos arctica dos Passes, 1935: 87; Forbes, 1945: 155; Klots, 1951: 101; Bauer, 1975: 144.

Holotype <J, CANADA: Newfoundland, Port au Port, Table Mountain (G. C. Hall) (AMNH, New York).
Phyciodes tharos pascoensis Wright; Bauer, 1975: 144, pi. 77, figs 9, 10.
Phyciodes tharos distincta Bauer, 1975: 144. Syntype(s), U.S.A.: California, Calexico, Imperial Co.

(depository not indicated).

c? forewing 15-17 mm, upperside fulvous, markings black, without colour contrast, discal marks in s2 and
s3 small or vestigial; hindwing fulvous discal band wide, the veins which cross the band not pigmented;
underside with a prominent dark mark on outer margin between s2 and s5, often enclosing a pale lunule.
9 similar, often larger.

Genitalia. In dorsal view,^ tegumen elongate, tapering slightly to rather wide apex, posterior border of
juxta with central prominence. 9, in dorsal view, sterigma deeply excavated, bursal duct firmly chitinised,
bursal support rather long.

DISTRIBUTION. Widely distributed in North America, especially east of the Rocky Mountains;
in Canada to 52°N., and southwards to Mexico.

DISCUSSION. There are two or more annual generations in suitable localities, with marked
seasonal variation. In spite of wide distribution, regional (geographical) variation is slight, and
Vawter & Brussard (1975) have recently commented on the apparent lack of geographical
variability of various enzyme systems in this species. Oliver (19780; 19786; 1979) comments
further on the genetics of this species, and reports on results from experimental hybridisations
with P. batesii and P. montanus.

Phyciodes batesii (Reakirt)
(Figs 2, 193)

[Melitaea tharos (Drury); Boisduval & Leconte, 1833: pi. 47, fig. 5. Misidentification.]
Eresia batesii Reakirt, 1865: 226. J1, 9 syntypes, U.S.A.: Virginia, Winchester; New Jersey, Gloucester,
Reakirt Coll. (lost?).

REVISION OF PHYCIODES 87

Phyciodes batesii (Reakirt); Rober, 1913: 436, pi. 89, row d [fig. 5]; Hall, 19286: 38; Forbes, 1945: 154, 189;

Holland, 1947: 136, pi. 17, figs 35, 36; Klots, 1951: 100, pi. 13, fig. 12; Bauer, 1975: 145, pi. 44, fig. 9.
Phyciodes batesii IT. f. harperi Gunder, 1932: 283. Holotype ?, CANADA: Manitoba, McCreary, Gunder

Coll. (present depository unknown).

3 like P. tharos on both surfaces, but forewing underside black discal markings more extensive; hindwing
underside yellow, faintly marked, dark shade along outer margin vestigial (if present). 4 similar.

Genitalia. j like P. tharos, in five specimens examined tegumen slightly more tapered, hooks closer
together and more slender, probably within the range of normal variation.

DISTRIBUTION. E. Canada and NE. states of U.S.A., from Ontario and Quebec to Virginia and
Nebraska, including New Jersey.

Phyciodes campestris (Behr)

(Figs 4, 5, 194-202)
Melitaea campestris Behr, 1863: 86.

Like Phyciodes tharos, j forewing 17 mm, variable, upperside black pattern more heavily marked, discal
band yellow, paler than postdiscal bands, yellow mark present at cell-end; hindwing underside marginal
lunule in s3 large, yellow or white.

Genitalia. Like tharos, J in dorsal view, scaphial hooks usually slightly smaller, v as in tharos.

DISTRIBUTION. Western regions of North America from Alaska southwards to Nevada and
Mexico. Two subspecies with similar genitalia.

Phyciodes campestris campestris (Behr)
(Figs 4, 194-199)

Melitaea campestris Behr, 1863: 86. + syntypes, U.S.A.: California (various unstated localities) (destroyed

by fire).

Melitaea pratensis Behr, 1863: 86. J syntypes, U.S.A.: California, near San Francisco (destroyed by fire).
Eresia campestris (Behr); Reakirt, 1866a: 142.

[Phyciodes orseis (Edwards); Godman & Salvin, 1882: 193. Misidentification.]
Phyciodes pratensis (Behr); Rober, 1913: 436, pi. 89, row d [figs 3, 4].

Phyciodes campestris (Behr); Comstock, 1927: 116, pi. 39, figs 4-6; Holland, 1947: 137, pi. 17, figs 37, 38.
Phyciodes campestris campestris (Behr); Hall, 1929: 46; Forbes, 1945: 154-155, 189 (part); Bauer, 1975: 145,

pi. 44, figs 10, 12.

As described above, colour contrast usually subdued.

DISTRIBUTION. Occurs especially in northern localities, Alaska, British Columbia etc., at
moderate altitudes. The figure in 'Seitz' is too dark.

Phyciodes campestris camillus Edwards
(Figs 5, 200-202)

Phyciodes camillus Edwards, 18716: 268. Lectotype j, U.S.A.: Colorado, Fairplay, Park Co. (Mead)

(CM, Pittsburgh), designated by Brown (1966: 451, fig. 30).
Phyciodes emissa Edwards, 18716: 269. Lectotype ?, U.S.A.: Colorado, Denver, Denver Co. (Mead)

(CM, Pittsburgh), designated by Brown (1966: 453, fig. 31).
Phyciodes camillus ab. rohweri Cockerell, 1913: 308, fig. 1. Holotype, U.S.A.: Colorado, North Boulder

Creek, Boulder County, Canadian Zone, viii. 1907 (S. A. Rohwer) (depository unknown).
Phyciodes camillus ab. tristis Cockerell, 1913: 308, fig. 2. Holotype, U.S.A.: Colorado, Jim Creek, Boulder

County, 7. ix. 1907 (S. A. Rohwer) (depository unknown).
Phyciodes camillus Edwards; Rober, 1913: 437, pi. 89, row d [figs 7, 8]; Holland, 1947: 138, pi. 17,

figs 32-34; Klots, 1951: 99, pi. 12, fig. 5.
Phyciodes campestris tr. f. mcdunnoughi Gunder, 1928: 167, figs 19, 19a. Holotype j, ? CANADA: 'T.N.W.,

Olds?' (CNC, Ottawa).

88 L. G. HIGGINS

Phyciodes campestris camillus Edwards; Hall, 1929: 47; Forbes, 1945: 155; Bauer, 1975: 146, pi. 78,
fig. 5.

^ like P. campestris campestris, but upperside markings brighter, black pattern reduced and colour contrast
often quite lively, with bands of red and yellow; hindwing underside usually greyish rather than yellow.

DISTRIBUTION. South-western states of U.S.A., especially Colorado and California, flying there
at 2000-3000 m.

Phyciodes montanus (Behr) stat. rev.
(Figs 3, 203-208)

Melitaea montana Behr, 1863: 85. Syntypes, U.S.A.: California, Los Angeles; headwaters of Tuolumne

River; Yosemite Valley (destroyed by fire).
Melitaea orsa Boisduval, 1869: 55. LECTOTYPE J, U.S.A.: California, 'Interior' (BMNH), here designated

[examined]. [Lectotype figured by Oberthiir, 1914: 81, fig. 2178.]
Phyciodes montana (Behr); Rober, 1913: 437, pi. 89, row e [fig. 3] (^); Comstock, 1927: 116, pi. 39,

figs 7, 8; Hall, 1929: 45; Holland, 1947: 138, pi. 17, figs 26, 27.
Phyciodes campestris montanus (Behr); Forbes, 1945: 155, 189; dos Passes, 1964.
Phyciodes campestris montana (Behr); Bauer, 1975: 145.

Like P. campestris camillus but slightly larger, e? forewing 16-18 mm, upperside black markings reduced,
colours brighter. ? similar.

Genitalia. 3 like campestris, in three specimens apical section of valve slightly shorter and more massive.
Hall (1929: 46) also considered 'valve shorter than in P. campestris, the apical process less developed'.

DISTRIBUTION. Seen only from California, especially the Sierra Nevada; recorded flying at
2000-3000 m.

DISCUSSION. Dos Passos (1964), probably following Forbes (1945), included montanus with
P. campestris as a subspecies. Hall (1929: 45) considered that montanus should have specific
rank, and I agree that there are small distinctive characters in the male genitalia; further, the
external characters are constant and recognisable. It is uncertain whether P. campestris camillus
also flies with montanus at high levels in the Sierra Nevada, California.

Phyciodes mylittus (Edwards)

(Figs 6, 7, 209-211)
Melitaea mylitta Edwards, 1861: 160.

J forewing 16-20 mm, variable, outer margin usually slightly excavate. Upperside fulvous yellow, black
pattern well defined, basal areas not suffused black ; hindwing upperside fulvous, discal area unmarked,
black postdiscal dots usually forming a complete series (6 dots). ? similar.

Genitalia. j tegumen like P. campestris, but terminal lateral hooks smaller, more widely separated.

DISTRIBUTION. From S. Canada and British Colombia through the western states of U.S.A. to
California (eastwards to foothills of the Rocky Mts.) to Mexico and Guatemala. Three
subspecies with similar genitalia.

Phyciodes mylittus mylittus (Edwards)
(Figs 6, 209-211)

Melitaea mylitta Edwards, 1861: 160. Neotype j, U.S.A.: California, Stanyan Hill, San Francisco

(CAS, San Francisco), designated by Brown (1966: 438, fig. 27).
Melitaea collina Behr, 1863: 86. Syntypes, U.S.A.: California, near San Francisco; hills of Contra Costa

(? destroyed by fire).
Melitaea epula Boisduval, 1869: 54. LECTOTYPE J, U.S.A.: California, 'Interior' (BMNH), here

designated [examined]. [Lectotype figured by Oberthiir, 1914: 80, pi. 259, fig. 2176.]

REVISION OF PHYCIODES 89

Phyciodes mylitta (Edwards); Rober, 1913: 437, pi. 89, row e [fig. 1] (^); Comstock, 1927: 117, pi. 39,

figs 13-15; Forbes, 1945: 153, 189; Holland, 1947: 138, pi. 17, figs 40, 41; Tilden, 1970: 97.
Phyciodes mylitta mylitta (Edwards); Hall, 1928ft: 42; Bauer, 1975: 149, pi. 16, figs 9, 10.
Phyciodes mylitta tr. f. collinsi Gunder, 1930: 62. Holotype J, U.S.A.: California, Collin's Ranch, Voltair,

Gunder Coll. (present depository unknown).
Phyciodes mylitta ab. macyi Fender, 1930: 182. Holotype, U.S.A.: Oregon, McMinnville (Fender)

(depository unknown).
? Phyciodes mylitta arizonensis Bauer, 1975: 149, pi. 43, fig. 8. Syntypes, U.S.A.: central & south-eastern

Arizona; New Mexico, Sonora; south-western Colorado (depository not indicated).

o forewing usually small, 16-18 mm, described above.

FLIGHT. In lowland areas, occurs in a succession of broods from April to October, but
double-brooded at altitudes of 2300 m or more.

DISTRIBUTION. Western states, from British Colombia to California, and east to foothills of the
Rocky Mts. in Montana and Colorado. Guppy (1974) and Shepard (1977) discuss the apparently
recent establishment of this species on Vancouver Island.

Phyciodes mylittus mexicanus Hall

Phyciodes mylitta mexicana Hall, 1928ft: 44. Holotype ^, MEXICO: Jalapa (Hoege) (BMNH) [examined].
Phyciodes mylitta mexicana Hall; Bauer, 1975: 150.

Both sexes like P. m. mylittus, but upperside fulvous ground colour slightly darker and all black markings
slightly heavier.

FLIGHT. November, December, April, probably in two broods. Occurs at 1300 m or more.
DISTRIBUTION. Eastern Mexico.

Phyciodes mylittus thebais Godman & Salvin
(Fig- 7)

Phyciodes thebais Godman & Salvin, 1878a: 267. Holotype ^, MEXICO: Mountains of Oaxaca (Fenocchio)

(BMNH, Type no. Rh. 8434; Gabriel, 1927: 118) [examined].

Phyciodes thebais Godman & Salvin; Godman & Salvin, 1882: 194, pi. 21, figs 13-15; Rober, 1913: 436.
Phyciodes mylitta thebais Godman & Salvin; Hall, 1928ft: 44; Bauer, 1975: 150.

Like P. mylittus mylittus, ground-colour paler fulvous, almost obscured by greatly extended black markings.
9 similar.

FLIGHT. Specimens dated April and July.
DISTRIBUTION. Western Mexico, Guatemala.

Phyciodes pallidus (Edwards)
(Fig. 8)

Melitaea pallida Edwards, 1864: 505. Neotype $, U.S.A.: Colorado, Flagstaff Mt, Boulder Co. (CM,

Pittsburgh), designated by Brown (1966: 443, fig. 28).
Eresiamata Reakirt, 1866a: 142. Syntype(s), U.S.A.: Rocky Mountains, Colorado Territory (Reakirt Coll.;

lost?). Syn. n.
Phyciodes barnesi Skinner, 1897: 154. 'Many' j syntypes, U.S.A.: Colorado, Glenwood Springs (W. Barnes)

(CM, Pittsburgh).
Phyciodes barnesi Skinner; Rober, 1913: 436, pi. 89, row e [fig.2] (J); Holland, 1947: 138, pi. 18,

fig. 5 (type).

Phyciodes mylitta barnesi Skinner; Hall, 1928ft: 43.
Phyciodes mylitta pallida (Edwards); Brown, 1966: 443.

Phyciodes pallida pallida (Edwards); Tilden, 1970: 96; Bauer, 1975: 148, pi. 44, fig. 5.
Phyciodes pallida barnesi Skinner; Tilden, 1970: 96; Bauer, 1975: 149, pi. 43, fig. 7, pi. 78, fig. 6.

90 L. G. HIGGINS

3 (brewing 18-21 mm, upperside like P. m. mylittus, no constant distinctive external character except larger
size. 9 black markings sometimes extended.
Genitalia. J like P. m. mylittus.

FLIGHT. Occurs as a single annual brood.

DISTRIBUTION. SW. Canada and Washington State southwards to Colorado and Arizona.

DISCUSSION. Larval and pupal characters are said to differ from those of P. m. mylittus.

Phyciodes orseis Edwards
(Fig. 212)

Phyciodes orseis Edwards, 1871a: 206. Lectotype ^, U.S.A.: California, Napa County, Mt. St. Helena

(Henry Edwards) (AMNH, New York), designated by Brown (1966: 450, fig. 29).
Phyciodes orseis Edwards; Rober, 1913: 436, pi. 89, row d [fig. 5]; Comstock, 1927: 117, pi. 39, figs 9-12;

Hall, 1929: 48; Forbes, 1945: 153, 155, 189; Holland, 1947: 137, pi. 17, fig. 31 ('type'; designation

rejected by Brown, 1966: 450); Scott, 1974 (early stages).
Phyciodes orseis ir. f. edwardsiGunder, 1927: 135, pi. 2, fig. 5. Holotype +, U.S.A.: California, W. Barnes Coll.

(present depository uncertain).
Phyciodes orseis orseis Edwards; Bauer, 1975: 147, pi. 43, fig. 6.

3 forewing 20 mm, like P. campestris, upperside black pattern extensive, colour contrast usually present in
pale areas. -4 similar.

Genitalia. J like P. m. mylittus.

DISTRIBUTION. Mountainous areas of western states of U.S.A. (coastal ranges), from Washington
State to California.

DISCUSSION. The species is said to fly in association with P. campestris and P. m. mylittus
(Scott, 1974), but separation from campestris may be extremely difficult.

Phyciodes herlani Bauer stat. n.

Phyciodes (Phyciodes) orseis herlani Bauer, 1975: 148, pi. 45, figs 9, 10. J, 9 syntypes, U.S.A.: Nevada,
Glenbrook Creek, Douglas Co. (depository not indicated).

Like P. orseis and P. campestris, upperside black markings greatly reduced, fulvous areas paler, colour
contrast present but less striking. + similar.
Genitalia. $ like P. m. mylittus.

DISTRIBUTION. California and Nevada (Douglas Co.), not rare at 2000 m or more.

Phyciodes pictus (Edwards)

(Figs 9, 213, 214)
Melitaea picta Edwards, 1865: 201.

cJ small, forewing 11-12 mm, like P. campestris but with colour contrast on upperside, cell-spot and discal
spots pale yellow; underside apex bright yellow, usually unmarked; hindwing underside yellow, darker
markings vestigial or absent. ? similar, often slightly larger.
Genitalia. Described below.

DISTRIBUTION. South-western states of U.S.A., and Mexico.
Two subspecies with slightly different genitalia.

Phyciodes pictus pictus (Edwards)
(Figs 9, 213)

Melitaea picta Edwards, 1865: 201. Lectotype _>, U.S.A.: Nebraska, North Platte, Lincoln Co., (Ridings)
(FMNH, Chicago), designated by Brown (1966: 457, fig. 32).

REVISION OF PHYCIODES 91

Phvciodes canace Edwards, 1871a: 206. Neotype J, U.S.A.: Arizona, nr Tucson, Pima Co. (Morrison)

(CM, Pittsburgh), designated by Brown (1966: 461, fig. 331).
Phyciodes picta (Edwards); Rober, 1913: 437, pi. 89, row e [figs 4, 5]; Comstock, 1927: 1 18; Holland, 1947:

139; pi. 17, figs 20, 21.
Phyciodes picta ab. jemezensis Brehme, 1913: 194, pi. 7, figs 7, 8. Holotype J, U.S.A.: New Mexico, Jemez

Springs, bred by J. Woodgate, Brehme Coll. (present depository unknown).
Phyciodes picta picta (Edwards); Hall, 1929: 49; Forbes, 1945: 154-155, 189; Brown, 1966: 455; Bauer,

1975: 146, pi. 44, fig. 11.
Phyciodes picta canace (Edwards); Brown, 1966: 457; Bauer, 1975: 147, pi. 16, fig. 11.

Upperside colour contrast very bright.
Genitalia. 3 tegumen short, wide, terminal lateral hooks relatively large and strong. ? not examined.

DISTRIBUTION. U.S.A., southern states, from Arizona and Kansas southwards to N. Mexico
(N. Sonora). Two broods, upperside black markings slightly heavier in first brood.

Phyciodes pictus pallescens (Felder)
(Fig. 214)

Eresia pallescens Felder, 1869: 469. cJ, V syntypes, MEXICO: Puebla, region of Cuernavaca (Hedemann)
(NM, Vienna; BMNH) [3 J, 3 9 ? syntypes from Puebla in BMNH examined].

Phyciodes pallescens (Felder); Godman & Salvin, 1882: 195, pi. 21, figs 18, 19; 1901: 678; Rober,
1913: 437, pi. 89, row f [fig. 10] (J).

Phyciodes picta pallescens (Felder); Hall, 1929: 50; Forbes, 1945: 154.

$ like P. pictus pictus, upperside markings less brilliant, forewing underside apical area with vestigial
markings; hindwing underside dark markings slightly indicated. 9 similar.

Genitalia. $ like P. pictus pictus but tegumen perhaps slightly narrower in two preparations. The
genitalia are minute and it is difficult to make a comparison.

DISTRIBUTION. According to Hall (1929: 50) this subspecies is restricted to central and southern
Mexico.

DISCUSSION. P. pallescens is placed as a distinct species by some authors.

Phyciodes phaon (Edwards)
(Figs 10, 215, 216)

Melitaea phaon Edwards, 1864: 505. Neotype^, U.S.A.: Georgia, St. Simon's Island, Glynn Co. (CM,

Pittsburgh), designated by Brown (1966: 437, fig. 26).

Phyciodes phaon (Edwards), winter form ('aestiva'); Edwards, 1878 (descriptive term).
Phyciodes phaon (Edwards), summer form ('hiemalis'); Edwards, 1878 (descriptive term).
Phyciodes phaon (Edwards); Godman & Salvin, 1901: 677; Rober, 1913: 436, pi. 89, row c [figs 7, 8];

Comstock, 1927: 116, pi. 39, figs 1-3; Holland, 1947: 137, pi. 17, figs 22, 23; Bauer, 1975: 143, pi. 44,

fig. 7; Riley, 1975: 79, pi. 12, fig. 18.
Phyciodes phaon maya Hall, 19286: 41. Holotype 0*, GUATEMALA: Lake Amatitlan, 3800 ft [1260 m],

October (BM, Brighton) [examined]. Syn. n.
Phyciodes phaon phaon (Edwards); Hall, 19286: 40.
Phyciodes phaon phaon ft. 'aestiva'; Hall, 19286: 40.
Phyciodes phaon f. phaon; Forbes, 1945: 154.

Phyciodes phaon f. 'hiemalis'; Forbes, 1945: 154; Brown, 1966: 464.
Phyciodes phaon f. 'aestiva'; Brown, 1966: 464.

cJ forewing 11-13 mm, upperside like P. pictus, but colour contrast less brilliant, post-discal band from
slb-s4 often fulvous in spring brood but in later broods bright yellow and very prominent, forewing
underside apex with usual marginal and submarginal markings; hindwing underside all markings well
defined, ground colour greyish in early brood, pale cream in later broods, v similar.

Genitalia. J like P. pictus, in dorsal view not quite so wide, terminal hooks very robust; penis slender.
2 not examined.

92 L. G. HIGGINS

DISTRIBUTION. U.S.A. (all southern states), Cayman Islands, Mexico, Guatemala, British
Honduras. Recorded from Cuba (Riley, 1975). Flies in two or three annual broods, with marked
seasonal variation.

Phyciodes vesta (Edwards)

(Figs 11,217,218)
Melitaea vesta Edwards, 1869: 371.

d small, forewing 13-14 mm, upperside like P. phaon, but without colour contrast and yellow markings;
hindwing underside markings well defined, especially postdiscal macular band. ? similar.

Genitalia. J distinctive; tegumen short, massive, two hooks at each angle of scaphial extension,
additional short hook sometimes present, y bursal duct short, chitinised, scutum large.

DISTRIBUTION. Local but widely distributed in south-western states of U.S.A., Guerrero in
Mexico, Guatemala.
Two subspecies, with similar genitalia.

Phyciodes vesta vesta (Edwards)
(Figs 11,217,218)

Melitaea vesta Edwards, 1869: 371. Neotype <J, U.S.A.: Texas, Neu Braunfels, Comal Co. (CM,

Pittsburgh), designated by Brown (1966: 463, fig. 34).
Phyciodes boucardi Godman & Salvin, 1878a: 268. Holotype J1, MEXICO: Putla (Rebouch) (BMNH

Type no. Rh. 8438; Gabriel, 1927: 23) [examined].
Phyciodes vesta (Edwards); Godman & Salvin, 1882: 195; 1901: 678; Rober, 1913: 436, pi. 89, row c

[fig. 5]; Hall, 1929: 50; Forbes, 1944; Holland, 1947: 136, pi. 17, figs 17-19; Bauer, 1975: 142, pi. 45,

fig. 14.
Phyciodes boucardi Godman & Salvin; Godman & Salvin, 1882: 194, pi. 21, figs 16, 17; 1901: 678

(gen. 2); Rober, 1913: 437, pi. 89, row e [figs 8, 9].
Melitaea arida Skinner, 1917: 328. Holotype, U.S.A.: Arizona, Cochise Co. (F. Haimbach) (CM

Pittsburgh).

Phyciodes vesta vesta (Edwards); Hall, 1929: 51.
[? Phyciodes thebais Godman & Salvin; Holland, 1947: 137, pi. 59, fig. 22. Misidentification.]

The markings of the upperside are regular and complete.

DISTRIBUTION. South-western states of U.S.A., Northern Mexico. Flies in two annual broods,
the summer brood (gen. 2, f. boucardi) has the underside hindwings heavily marked with
purplish brown.

Phyciodes vesta graphica (Felder)

Eresia graphica R. Felder, 1869: 470. J, V syntypes, MEXICO: Huahuapan, Hedemann (NM, Vienna ?).
Phyciodes vesta vestalis Hall, 1929: 52. Holotype^, GUATEMALA (BM, Brighton) [examined]. Syn. n.
[Phyciodes vesta vesta (Edwards); Hall, 1929: 51, in part. Misidentification.]

c? slightly larger than nominate P. vesta, upperside tone of fulvous ground colour more intense and
black markings slightly extended.

DISTRIBUTION. S. Mexico and Guatemala, doubtfully recognisable as a valid subspecies.

DISCUSSION. Eresia graphica was described by R. Felder from specimens collected at Huahuapan
by Hedemann; these are presumed to be in the NM, Vienna. A male from 'Cuernavaca',
ex Felder Coll., is now in the BMNH and bears Felder's determination label 'Eresia graphica
Feld.', and an ex Rothschild 'type' label. However, it seems doubtful that this specimen is part
of the original type-series, although it was possibly in the Felders' collection at the time of
description.

REVISION OF PHYCIODES 93

PHYST/S gen. n.

Type-species: Eresia simois Hewitson. Gender: feminine.

Small butterflies, wings broad, forewing outer margin convex. Upperside brown, markings small yellow
spots arranged across the wings in regular transverse series; on hindwing upperside each spot of the
postdiscal series encloses a small blind pupil.

Genitalia. j tegumen in dorsal view short, wide, slightly chitinized, scaphial extension vestigial, posterior
border ofjuxta gently convex or almost straight, saccus deeply cleft, posterior section of valve terminating
in a simple, incurved pointed apex, harpe slightly variable; penis slender, probably lacking ostium-keel.
? genitalia unusually simple, in dorsal view ostium bursae exposed, the duct not chitinized, sterigma
shallow, the cup-shaped bursal support arising very close to ostium, post-vaginal scutum large.

DISTRIBUTION. Brazil, Argentina, Uruguay, Bolivia, Paraguay, NE. Peru.

DISCUSSION. The genus includes a single divergent and very variable species, with wing markings
which suggest relationship with the Phyciodes of North America. This is not confirmed by the
genitalia, which are distinctive, especially those of the female, and unlike those of any other
species.

Phystis simois (Hewitson) comb. n.

(Figs 12, 13, 219-223)
Eresia simois Hewitson, [1864].

Size very variable, o forewing 10-15 mm, upperside markings yellow to cream, prominent or greatly
reduced in size by extension of the brown ground colour. On underside of forewing a black postdiscal
mark is characteristic and usually prominent. On hindwing upperside a postdiscal series of ocellar spots
recalls Phyciodes tharos and its allies.

Genitalia. See generic description. The organs are very small. In a series of seven preparations I have not
found any consistent variation to suggest separation into two species. The variation observed has been due,
probably, to small differences in rotation of the parts within the mountant.

DISTRIBUTION. Widely distributed in Brazil, extending southwards to N. Argentina and westwards
to Peru, Uruguay and Bolivia.
Two subspecies.

Phystis simois simois (Hewitson)
(Figs 12,219,220)

Eresia simois Hewitson, [1864]: [21], pi. [11], figs 30, 31. LECTOTYPE ?, BRAZIL (BMNH, Type no.

Rh. 8435), here designated [examined].
Phyciodes pedrona Moulton, 1909: 103. HolotypeJ, BRAZIL: 'Minas Geraes, Discoberto, near Joao Pedro's

house' (W. J. Burchell) (UM, Oxford, Type no. 1149) [examined]. Syn. n.
Phyciodes pedrona Moulton; Rober, 1913: 436.
Phyciodes simois simois (Hewitson); Hall, 1929: 53.
Phyciodes simois pedrona Moulton; Hall, 1929: 53.
Phyciodes (Tritanassa) simois (Hewitson); Forbes, 1945: 189.
Phyciodes (Tritanassa) pedrona Moulton; Forbes, 1945: 189.

j forewing 10-11 mm, usually very small, upperside yellow markings at base and in cell obscured by
extension of brown ground colour, but with an oblique discal series of small yellow spots rather better
defined; hindwing upperside markings small and regular; underside bright yellowish brown, on forewing
a wide, black, postdiscal area extends from costa to inner margin, with white spots, as on upperside, very
prominent. ? similar, but larger.

DISTRIBUTION. N. Brazil: Pernambuco, Bahia, etc.

94 L. G. HIGGINS

Phystis simois variegata (Rober)
(Figs 13, 221-223)

Phyciodes variegata Rober, 1913: 437, pi. 89, row f [fig. 3] (J). Holotype ^, ARGENTINA: La Soledad,

Prov. Entre Rios (E. A. Britton) (BMNH) [examined].
Phyciodes simois ab. nigrina Hayward, 1931: 51, pi. 11, fig. 10. Holotype J, ARGENTINA: Prov. Cordoba,

Yacanto (Breyer Coll.; present depository unknown).
Phyciodes simois variegata Rober; Hall, 1929: 53.
Phyciodes simois pratti Hall, 1935: 221. Holotype J, PERU: W. slopes of Andes, 4000 ft [1330 m] (BM,

Brighton) [examined]. Syn. n.
Phyciodes pedrona variegata Rober; Hayward, 1952: 290; 19646: 342, pi. 18, fig. 6.

cJ forewing 13-15 mm, variable, upperside with yellow spots in pattern similar to simois simois but larger,
postdiscal costal bar of 4 macules prominent and spots in sib, s2 and s3 enlarged, forming with the
costal bar, a broken band across the wing; forewing underside black postdiscal area reduced, spots usually
yellow (not white), submarginal area with single spot in s3; hindwing underside pale buff, markings
vestigial, if present. 9 usually slightly larger, markings also larger and paler.

DISTRIBUTION. Brazil, especially in southern regions, N. Argentina, Uruguay, Bolivia, Peru.

DISCUSSION. The two subspecies, as described, represent the extremes of a well-defined cline. The
geographical limits of simois simois are found in northern Minas Gerais, and from there
southwards through C. and S. Brazil there are larger forms with intermediate characters, often
difficult to place with one or other of the recognised subspecies. In Argentina and in Uruguay
simois variegata is fully developed. The species is represented in the BMNH by occasional
specimens from Bolivia and Peru, but the material is very scanty and insufficient to show local
character.

The c? holotype of Phyciodes pedrona Moulton is very small and in my view is referable to
nominate simois. On the forewing upperside the yellow discal spots are slightly more prominent
than usual. A single paratype in the BMNH has similar characters.

A NTH AN ASS A Scudder

Anthanassa Scudder, 1875: 239, 268. Type-species: Melitaea texana Edwards, by subsequent designation
[= Eresia cincta Edwards, by original designation. Misidentification] (see ICZN, 1967, Bull. zool. Nom.
24: Opinion 839: 337). Gender: feminine.

Tritanassa Forbes, 1945: 171 (as subgenus of Phyciodes). Type-species: Eresia drusilla Felder & Felder,
by original designation. Syn. n.

Butterflies of less than median size, forewing apex often truncate, outer margin excavate. On uppersides
all species are black or dark brown with white or yellow markings arranged in a characteristic pattern,
on forewing rounded white or yellow submarginal spots in sib and s4 are constant, making a useful
generic character; hindwing upperside with a transverse or slender discal band, and a series of submarginal
and marginal lunules present in most species. Sexes similar, or nearly so.

Genitalia. In most species £ tegumen is little chitinised, scaphial extension often extremely fragile,
membranous; valve apex slender, preapical tooth well developed, so that the apex appears to be bifid in
dorsal view (Fig. 229); saccus deeply cleft; penis slender with a small morula, ostium keel absent.
? bursal duct short, bursal support usually small but well defined.

In A. drusilla and the 1 1 species which follow it in this work, the genitalia are uniform, but good
specific characters are present in the genitalia of the seven remaining species.

DISTRIBUTION. The genus is well developed in Mexico, Central America and western South
America, from Venezuela and Colombia to Bolivia, in mountainous regions. It is represented in
eastern regions, including Brazil, Paraguay and northern Argentina, by one species, A. hermas.
A single species, A.frisia, occurs in Jamaica, Cuba and in other islands of the Greater Antilles.
The species are not mimetic.

REVISION OF PHYCIODES 95

Key to species of Anthanassa

1 In dorsal view, lateral walls of scaphial extension dentate (Fig. 255) 2

In dorsal view scaphial walls not dentate 6

2 Spiny area of scaphium in a wide band (Fig. 255) hermas (p. 104)

Spines confined to lateral walls of scaphium (Fig. 258) 3

3 Upperside markings very pale on dark grey-brown tulcis (p. 105)

Upperside markings fulvous or yellow 4

4 Upperside markings yellow on medium-brown dubia (p. 106)

Upperside markings fulvous 5

5 Hindwing upperside prediscal band short frisia (p. 105)

Hindwing upperside prediscal band well developed taeniata (p. 106)

6 Valve apex with 2 or 3 prominent teeth 7

Valve apex appears bifid 10

7 Valve apical teeth short sosis (p. 106)

Valve medial apical tooth long 8

8 Underside forewing base bright orange sitalces (p. 107)

Underside forewing base grey-brown ........... 9

9 Valve apex medial tooth very long (Fig. 267) drymaea (p. 107)

Valve apex medial tooth shorter (Fig. 269) cartes (p. 107)

10 Hindwing upperside transverse band well-defined 11

Hindwing upperside transverse band obscure or absent 17

11 Hindwing upperside transverse band yellow 12

Hindwing upperside transverse band white 15

12 Band wide at outer margin, tapering rapidly 13

Hindwing not so marked 14

13 c? forewing 19 mm, single spot in sib, hind wing underside pallid buff, markings obscure

dracaena (p. 99)
<3 forewing 17 mm, two spots in sib, hindwing underside markings brighter, grey and brown

phlegias (p. 99)

14 Hindwing upperside marginal lunules of even size drusilla (p. 96)

Hindwing upperside marginal lunules enlarged in s6 and s7 .... ptolyca (p. 97)

15 Forewing base orange-red t ex ana (p. 99)

Forewing not so marked 16

16 Forewing upperside markings yellowish alexon (p. 100)

Forewing markings white on black ardys (p. 98)

17 Forewing upperside with large orange costal mark, other markings vestigial or absent

fulviplaga (p. 104)
Forewing not so marked 18

18 Forewing upperside with orange postdiscal oblique band .... crithona (p. 104)
Forewing not so marked 19

19 Hindwing upperside gleaming blue-black, marginal border black, otherwise unmarked . . 20
Hindwing upperside not so marked 21

20 Forewing upperside with yellow markings in pattern like drusilla . . . otanes sopolis (p. 103)
Upperside markings greatly reduced or absent ...... otanes otanes (p. 103)

21 Upperside hindwing dark brown, markings vestigial, if present 22

Upperside hindwing postdiscal markings combine to form a series of ocelliform circles . 23

22 Forewing upperside discal area chestnut-brown ....... argentea (p. 102)

Forewing upperside very dark, markings obscure if present ..... atronia (p. 102)

23 Forewing upperside dark brown, markings vestigial if present .... acesas (p. 101)
Forewing upperside markings well developed 24

24 Hindwing upperside yellow discal band obscure, divided into macules by dark veins

nebulosa (p. 101)
Hindwing upperside discal band absent annulata (p. 103)

96 L. G. HIGGINS

Anthanassa drusilla (Felder & Felder) comb. n.

(Figs 14-16, 224-228)
Eresia drusilla Felder & Felder, 1861: 103.

cJ forewing size variable, 14-17 mm, upperside dark brown, markings orange-fulvous to yellow-buff,
individually and locally variable; hindwing upperside pale discal band present, usually with postdiscal and
submarginal spots and lunules. 9 slightly larger, with similar markings.

Genitalia. <3 in dorsal view, tegumen short, membranous, extremely fragile; posterior border of juxta
wide, concave or slightly wavy, lateral angles prominent. 9 ductus chitinised, short, bursal support shallow,
post-vaginal scutum extensive.

DISTRIBUTION. From Mexico across central and western South America to Peru, Ecuador and
Bolivia.
Four subspecies, all with similar genitalia.

•

Anthanassa drusilla drusilla (Felder & Felder)
(Figs 14, 224-228)

Eresia drusilla Felder & Felder, 1861 : 103. LECTOTYPE rf, VENEZUELA (Moritz) (BMNH, specimen bears

original label of Felders' inscribed 'drusilla H.-S.', locality data, and 'type') here designated [examined].

(A specimen labelled BM Type no. Rh. 8447 and listed by Gabriel, 1927: 42, is a 9 paralectotype.)
Phyciodes flavimacula Rober, 1913: 442, pi. 90, row c [fig. 2] (9). LECTOTYPE <J, COLOMBIA: Coreato,

Cauca Valley (Paine & Brinkley) (BMNH, specimen bears Rober's determination label, and is also

labelled 'type') here designated [examined].
Phyciodes drusilla drusilla (Felder & Felder); Hall, 1929: 86.

c? forewing 15-16 mm, variable, upperside markings bright orange-fulvous, hindwing underside pale discal
band usually prominent. 9 usually larger, upperside markings slightly paler, vague basal markings often
present on forewing upperside.

DISTRIBUTION. Venezuela, Colombia, Ecuador, Panama (rare).

Anthanassa drusilla lelex (Bates)

Melitaea lelex Bates, 1864a: 82. LECTOTYPE rf, PANAMA: Lion Hill (McLeannan) (BMNH, Type no.

Rh. 8445; Gabriel, 1927: 72), here designated [examined].
Melitaea alethes Bates, 1864a: 82. LECTOTYPE J, GUATEMALA: Central Valleys (Godman & Salvin)

(BMNH Type no. Rh. 8443; Gabriel, 1927: 9), here designated [examined].
Melitaea stesilea Bates, 1864a: 82. LECTOTYPE 9, GUATEMALA: interior (BMNH Type no. Rh 8444;

Gabriel, 1927: 113), here designated [examined]. [9 off. alethes.]
[Phyciodes ptolyca (Bates); Godman & Salvin, 1882: 201, pi. 21, fig. 34 (9), fig. 35 (J underside),

fig. 36 (^ underside). Misidentification.]

[Phyciodes ptolyca (Bates); Rober, 1913: pi. 90, row b [fig. 3] (£), [fig. 4] (<J). Misidentification.]
Phyciodes drusilla lelex (Bates); Hall, 1929: 88.
Phyciodes drusilla alethes (Bates); Hall, 1929: 88.

<3 upperside like A. drusilla drusilla but markings paler, ochre-yellow or buff. In some districts postdiscal
and submarginal markings become less distinct (f. alethes). 9 similar, markings often pale.

DISTRIBUTION. Mexico and Central America, widely distributed, with transition zone to nominate
drusilla in Panama, with minor local variation.

Anthanassa drusilla alceta (Hewitson)
(Fig. 15)

Eresia alceta Hewitson, 1869a: 28, [index]. LECTOTYPE^, ECUADOR: Rio Verde (Buckley) (BMNH,
Type no. Rh. 8454; Gabriel, 1927: 9), here designated [examined]. [Incorrectly identified and labelled
9 by Hewitson.]

REVISION OF PHYCIODES 97

Phyciodes flavimacula conflua Rober, 1913: 442, pi. 90, row c. Syntype(s), PERU: Chanchamayo (depository

unknown).
Phyciodes drusilla alceta (Hewitson); Hall, 1929: 89.

cJ like A. drusilla drusilla, forewing 16-18 mm, upperside markings orange-fulvous, postdiscal costal bar
and discal spots fused to form a band 2-3 mm wide; hindwing upperside fulvous discal band 3 mm wide,
veins crossing band little, if at all, darkened.

DISTRIBUTION. Ecuador, N. Peru. The distribution of this rather striking form appears to be well
defined, but Hall records intermediate specimens grading into nominate drusilla.

Anthanassa drusilla verena (Hewitson)
(Fig. 16)

Eresia verena Hewitson, [1864]: [20], pi. [10], figs 27, 28. LECTOTYPE rf, BOLIVIA (BMNH, Type

no. Rh. 8455; Gabriel, 1927: 122), here designated [examined].
Phyciodes verena (Hewitson); Rober, 1913: 440, pi. 90, row a [fig. 1] (^).
Phyciodes drusilla verena (Hewitson); Hall, 1929: 89.

J forewing 16-17 mm, upperside like drusilla drusilla, orange-fulvous discal band complete from costa to
inner margin, 2-3 mm wide; hindwing upperside postdiscal area clear orange-fulvous, 3-4 mm wide.
$ larger, forewing 19 mm, upperside markings slightly paler and more extensive.

DISTRIBUTION. Bolivia. This striking form appears to be uncommon but constant where it occurs.

Anthanassa ptolyca (Bates) comb. n.

(Figs 17, 229, 230)
Melitaea ptolyca Bates, 1864a: 81.

Like Anthanassa drusilla, o forewing 15-18 mm, upperside markings yellow to white, hindwing postdiscal
markings vestigial or absent, submarginal lunules present, enlarged in s7 and s8. * larger, markings usually
extended.

Genitalia. 3 in dorsal view like A. drusilla, slightly less massive, tegumen often better defined, scaphial
extension tapering, y bursal support small, scutum very large.

DISTRIBUTION. From Mexico southwards through Central America to Venezuela.
Two subspecies or clinal forms.

Anthanassa ptolyca ptolyca (Bates)
(Figs 17, 229, 230)

Melitaea ptolyca Bates, 1864a: 81. LECTOTYPE J, GUATEMALA: Chisoy Valley (Godman & Salvin)
(BMNH, Type no. Rh. 8448; Gabriel, 1927: 101), here designated [examined].

Phyciodes ptolyca (Bates); Godman & Salvin, 1882: pi. 21, fig. 32 (J), fig. 33 (<J underside), fig. 34
($ underside).

Phyciodes carrera Hall, 1917: 162. LECTOTYPE J, GUATEMALA: L. Amatitlan, 4000 ft [1330 m]
(BM, Brighton), here designated [examined]. [Specimen bears Hall's ms type label.]

Phyciodes ptolyca ptolyca (Bates); Hall, 1929: 92.

Upperside markings bright orange-yellow in both sexes, J forewing underside base yellow; hindwing discal
band well-defined.

DISTRIBUTION. Mexico, Guatemala, Nicaragua, Venezuela.

Anthanassa ptolyca amator (Hall)

Phyciodes ptolyca amator Hall, 1929: 92. Holotype j, MEXICO: Venta de Zopilote, Guerrero, 2800 ft
[930 m], (H.H. Smith) (BMNH) [examined].

Upperside markings pale yellow to cream-white, hindwing discal band narrow ; underside forewing base pale
yellow-brown.

98 L. G. HIGGINS

DISTRIBUTION. Western Mexico, flying at altitudes of 600-1800 m.

DISCUSSION. The status of this form is uncertain since all specimens in the BMNH, and in
the BM, Brighton (Hall Coll.), came from a single locality.

Anthanassa ardys (Hewitson) comb. n.

(Figs 18, 19, 231-233)
Eresia ardys Hewitson, [1864]: [22].

3 forewing 16-19 mm, upperside black with small white markings, which include a cell-spot slightly before
cell-end, and a small submarginal spot in s3, vestiges of yellowish basal striae sometimes present;
hindwing upperside discal band white; hindwing underside variable, marbled in grey, pale and dark brown.
9 larger, upperside markings cream-white, usually expanded, wing-bases with obscure yellowish striations.
Genitalia. 3 like Anthanassa drusilla, tegumen better defined, posterior border of juxta gently concave,
penis slender. 9 not examined.

DISTRIBUTION. From S. Mexico through Central America to Colombia.
Two subspecies with similar genitalia.

Anthanassa ardys ardys (Hewitson)
(Figs 18,231,232)

Eresia ardys Hewitson, [1864]: [22], pi. [11], figs 35, 36. LECTOTYPE 3, MEXICO: Orizaba (BMNH,

Type no. Rh. 8442; Gabriel, 1927: 15), here designated [examined].
? Eresia genigueh Reakirt, 1865: 225. Syntype(s)^, U.S.A.: Los Angeles, California (Reakirt Coll.; present

depository unknown). [Treated by Hall, 1929: 90, as a synonym of ardys ardys, without comment; if this

is correct, the type-locality given by Reakirt must be in error — if not, then Hall's synonymy must be

in error!]
Phyciodes ardys ardys (Hewitson); Hall, 1929: 90.

cJ forewing upperside cell-spot vestigial; hindwing upperside white discal band narrow, 1-2 mm wide,
slightly sinuous, broken by black veins, series of slender submarginal lunules complete. 9 underside
hindwing white discal stripe narrow and irregular if present.

DISTRIBUTION. S. Mexico, Nicaragua, Costa Rica, Colombia.

Anthanassa ardys subota (Godman & Salvin)
(Figs 19, 233)

Phyciodes subot a Godman & Salvin, 1878a: 268. LECTOTYPE^, GUATEMALA: Polochic Valley (Godman &

Salvin) (BMNH, Type no. Rh. 8505; Gabriel, 1927: 114), here designated [examined].
Phyciodes subota Godman & Salvin; Godman & Salvin, 1882: 204, pi. 22, figs 7, 8.
Phyciodes ardys subota Godman & Salvin; Hall, 1929: 91.

Usually slightly larger in overall size than ardys ardys; upperside markings milk-white, slightly larger,
cell-spot larger, wing bases black; hindwing upperside white discal band wider, 2 mm, veins less heavily
marked, submarginal lunules absent; hindwing underside with white discal band prominent.
Genitalia. 3 in dorsal view, posterior section of valve more elongate (single preparation).

DISTRIBUTION. Guatemala (Polochic Valley), Costa Rica.

DISCUSSION. A. ardys subota appears to be a constant form, and widely distributed in Guatemala.
Its taxonomic position is a little uncertain as A. ardys ardys occurs in neighbouring countries,
but an overlap of distribution has not been recorded. Hall (1929: 91) states that A. ardys ardys
does not occur in Guatemala. The figure in Godman & Salvin (1882: pi. 22, fig. 7) is
considerably enlarged (forewing 20 mm) compared with the actual specimen, now preserved in
the BMNH.

REVISION OF PHYCIODES 99

Anthanassa dmcaena (Felder & Felder) comb. n.
(Figs 20, 234-236)

Eresia dracaena Felder & Felder, 1867: 393. LECTOTYPE £, COLOMBIA: 'Bogota' (Lindig) (BMNH),

here designated [examined]. [Specimen bears Felders' original ms label.]
Phyciodes dracaena (Felder & Felder); Rober, 1913: 442, pi. 90, row c [fig. 3] (underside).
Phyciodes dracaena dracaena (Felder & Felder); Hall, 1929: 100, pi. 1, fig. 11 (^).

o (brewing 19 mm, elongate, upperside dark brown, markings orange, like Anthanassa drusilla but with
oblique mark before cell-end and lacking proximal orange spot in sib; hindwing underside very pale buff,
markings not well defined, basal area with irregular markings limited by broken brown discal line
(most prominent on the costa), outer margin orange-brown from s5 to anal angle. ? similar, forewing
20 mm, markings white, hindwing upperside discal band faintly yellow.

Genitalia. $ like A. drusilla, tegumen and scaphial extension better defined, posterior border of juxta
almost straight, valve apex and harpe more slender. ? sterigma not large, ductus lightly chitinized, bursal
support large, cylindrical, a most distinctive feature, post-vaginal scutum extensive.

DISTRIBUTION. Colombia, not recorded elsewhere.

Anthanassa phlegias (Godman & Salvin) comb, n., stat. rev.

(Fig. 21)

Phyciodes phlegias Godman & Salvin, 1901: 680, pi. 108, figs 21, 22. LECTOTYPE J, HONDURAS
(BMNH, ex Staudinger, ex Godman & Salvin Coll.; Type no. Rh. 8502; Gabriel, 1927: 97), here
designated [examined].

Phyciodes carigia Schaus, 1902: 395. Holotype (sex ?), COLOMBIA (USNM, Type no. 5889).

Phyciodes carigia Schaus; Schaus, 1913: 346, pi. 50, fig. 9 (^).

Phyciodes platytaenia Rober, 1913: [no text] pi. 90, row b [fig. 5] (j). Syntype(s) o, [HONDURAS]
(depository unknown).

Phyciodes dracaena phlegias Godman & Salvin; Hall, 1929: 100.

J1 forewing 17 mm, like Anthanassa dracaena but smaller, outer margin deeply excavate, upperside
markings orange-fulvous as in Anthanassa drusilla, but wing-base dark, spot at base of space Ib present;
hindwing uppermost orange discal band wide, tapering to inner margin, submarginal lunules well defined
but narrow postdiscal band absent; hindwing underside discal band grey, brightly marked, expanded at
outer margin, brown costal mark conspicuous.
Genitalia. Not examined.

DISTRIBUTION. Honduras (1 o ), Costa Rica (1 o )• A rare species.

Anthanassa texana (Edwards)

(Figs 22, 237-239)
Melitaea texana Edwards, 1863: 81.

j forewing 18-19 mm, outer margin deeply scalloped, upperside dark brown, markings small white spots,
small spot in cell near apex, another in s2 near base; hindwing upperside white discal band narrow,
crossed by dark veins, small orange spot in cell; forewing underside base orange-red; hindwing white
discal band not prominent, marginal area strongly marbled in grey, light and dark brown. ? larger,
markings similar, upperside with confused orange basal markings more prominent.

Genitalia. J like Anthanassa drusilla, in dorsal view narrower, valve apex more slender, harpe less
massive. 9 like A. drusilla.

DISTRIBUTION. Southern States of U.S.A., from Arizona and Texas to N. Mexico and eastwards
to Georgia and Florida.
Two subspecies with similar genitalia.

100 L. G. HIGGINS

Anthanassa texana texana (Edwards)
(Figs 22, 237-239)

Melitaea texana Edwards, 1863: 81. ?Holotype .3, U.S.A: Texas [New Braunfels, Comal Co.] (CM,

Pittsburgh) (see Brown, 1966: 421, fig. 22).
Eresia smerdis Hewitson, [1864]: [21], pi. [11], figs 33, 34. LECTOTYPE J, MEXICO (BMNH, type no.

Rh. 8497; Gabriel, 1927: 112), here designated [examined].
[Eresia cincta Edwards; Scudder, 1875: 239, 268. Misidentification.]
Phyciodes texana (Edwards); Rober, 1913: 442, pi. 90, row c [fig. 5] (3), [fig. 6] (?).
Anthanassa texana (Edwards); Barnes & McDunnough, 1917: 10; Holland, 1947: 141, pi. 18, figs 8, 9.
Phyciodes texana texana (Edwards); Hall, 1929: 96.
Phyciodes texana (Edwards); Forbes, 1945: 178.

3 forewing upperside wing bases brown with obscure orange markings.
DISTRIBUTION. U.S.A: Arizona, Texas, New Mexico, Nebraska; Mexico.

Anthanassa texana seminole (Skinner)

Eresia texana seminole Skinner, 1911: 412. cJ, 9 syntypes, U.S.A.: Bainbridge, Georgia (CM, Pittsburgh).
Anthanassa texana var. seminole Skinner; Holland, 1947: 141, pi. 59, fig. 14 (3).

Differs from A. texana texana on upperside, bases of both wings being orange-red.
DISTRIBUTION. Only south-east U.S.A.: Florida, Georgia.

DISCUSSION. There is little information about this butterfly; its status as a subspecies needs
confirmation.

Anthanassa alexon (Godman & Salvin) comb, n., stat. rev.

(Figs 23, 240)
Phyciodes alexon Godman & Salvin, 1889: 353.

Like Anthanassa texana, 3 forewing 17-19 mm, upperside dark brown, markings white or yellowish;
forewing underside base yellow-grey; hindwing underside with triangular, brown costal mark, other
markings indistinct. 9 larger, markings similar.

Genitalia. Like A. ardys, J differs slightly in shape of tegumen and in bilobed scaphial extension,
posterior border of juxta with central prominence. 9 not examined.

DISTRIBUTION. Arizona, Mexico (Guerrero).
Two subspecies with similar genitalia.

Anthanassa alexon alexon (Godman & Salvin)
(Fig. 23)

Phyciodes alexon Godman & Salvin, 1889: 353. LECTOTYPE .J, MEXICO: Cuernavaca, Morelos (H. H.

Smith) (BMNH, type no. Rh. 8499: Gabriel, 1927: 9), here designated [examined].
Phyciodes alexon Godman & Salvin; Godman & Salvin, 1901: 681, pi. 108, figs 23-26.
? Phyciodes natalces Dyar, 1913a: 279. Holotype ?, MEXICO: Rascon, San Luis Patosi (R. Mutter] (USNM,

Type no. 14487).
Phyciodes nebulosa alexon Godman & Salvin; Hall, 1929: 102.

3 forewing 17-18 mm, upperside markings chiefly white or cream, with small, confused brown marbling
at base; hindwing orange-brown markings more prominent, * similar, slightly larger.

DISTRIBUTION. Mexico.

REVISION OF PHYCIODES 101

Anthanassa alexon subconcolor (Rober)
(Fig. 240)

Phyciodes subconcolor Rober, 1913: 441, pi. 90, row b [fig. 1] (<$). LECTOTYPE J, U.S.A.: Benson,
Arizona (0. T. Baron) (BMNH), here designated [examined]. [Specimen bears Rober's original
determination label, and a separate, pink 'type' label, ex Rothschild Coll.]

Phyciodes nebulosa subconcolor Rober; Hall, 1929: 102.

3 forewing 19 mm, larger than alexon alexon, upperside yellow-brown markings more prominent and
with general brown flush overall. 9 larger, markings similar.

DISTRIBUTION. Arizona.

Anthanassa acesas (Hewitson) comb. n.
(Figs 24, 241)

Eresia acesas Hewitson, [1864]: [24], pi. [12], figs 48, 49. LECTOTYPE 9, VENEZUELA (BMNH, Type no.

Rh. 8466; Gabriel, 1927: 5), here designated [examined].
? Phyciodes annita Staudinger, 1885: 92, pi. 36. ^ syntype(s), VENEZUELA: Merida (Hahnel) (? MNHU,

Berlin).
Phyciodes acesas acesas (Hewitson); Hall, 1929: 107, pi. 1, fig. 2 (3).

This and the following species (A. nebulosa) must be distinguished from species of Telenassa
with rather similar ocellar ring markings on the upperside of the hindwings.

3 forewing 15 mm, upperside very dark with a few indistinct or obsolescent yellowish markings in the
submarginal area, otherwise unmarked; underside discal and postdiscal markings well developed as in
A. nebulosa', hindwing upperside no basal markings, postdiscal series of yellowish lunules forming 5 rings
in slb-s5, rings completed by short striae proximal to submarginal lunules. 5 slightly larger, upperside
markings better defined.

Genitalia. J like A. ardys, apical teeth of valve rather long, harpe slender, posterior border of juxta
almost straight.

DISTRIBUTION. Venezuela.

Anthanassa nebulosa (Godman & Salvin) comb. n.

(Figs 25, 242, 243)

Phyciodes nebulosa Godman & Salvin, 1878a: 269. LECTOTYPE o, GUATEMALA: Las Nubes (O. Salvin)

(BMNH, Type no. Rh. 8501; Gabriel, 1927: 85), here designated [examined].
Phyciodes castianira Godman & Salvin, 1880: 123, 131, pi. 4, fig. 10. LECTOTYPE J, COLOMBIA: Manaure,

Sierra Nevada de Santa Marta (F. Simons) (BMNH, Type no. RH. 8504; Gabriel, 1927: 27), here

designated [examined]. Syn. n.
Phyciodes nebulosa Godman & Salvin; Godman & Salvin, 1882: 205, pi. 22, figs 13, 14; Rober, 1913:

443, pi. 88, row i [fig. 7].
Phyciodes nebulosa nebulosa Godman & Salvin; Hall, 1929: 101.

c£ forewing 16-17 mm, small, upperside dark brown, markings orange-yellow, in general pattern like
A. drusilla, postdiscal spots in sib and s4 prominent, spot at middle of sib present; hindwing upperside
with basal striae, a narrow, inconspicuous discal band divided by dark veins, and postdiscal lunules
forming a series of 5 ocelliform rings, each completed by short striae placed before the submarginal
lunules. 9 larger, 18 mm, all markings brighter, well defined, postdiscal spots joined to make a wide
yellowish band.
Genitalia. J like A. acesas.

DISTRIBUTION. N. Colombia, Guatemala.

DISCUSSION. The holotype of A. nebulosa is a rather dark specimen of the species later described
by Godman & Salvin as castianira, and this name must fall as a synonym, but remains
available if required.

102 L. G. HIGGINS

The two forms are not truly identical. The lectotype of nebulosa is slightly darker on the
upperside, with markings somewhat obscured by brown dusky suffusion, when compared with
the original type-material of castianira, five males all from the Cinchicua Valley in N. Colombia.

Anthanassa argentea (Godman & Salvin) comb, n., stat. rev.

(Figs 26, 244)

Phyciodes argentea Godman & Salvin, 1882: 207. LECTOTYPE rf, GUATEMALA: Choctum (Hague)

(BMNH, Type no. Rh. 8468; Gabriel, 1927: 16), here designated [examinedj.
Phyciodes atronia atronia f. argentea Godman & Salvin; Hall, 1929: 111.

<$ forewing 16 mm, like A. atronia, upperside very dark, but with wide field of chestnut-brown over
inner margin and discal area, pale postdiscal spots in sib and s4 distinct, other markings more or less
vestigial; hindwing upperside brown flush generally extends along inner margin; forewing underside costa
and much of hindwing pale grey with fine dark striae. V upperside less dark, markings yellow, spots
arranged as in A. drusilla; hindwing underside yellow-grey with few markings.

Genitalia. 3 like A. ardys, tegumen membranous and very fragile, valve apex slender in dorsal view,
posterior margin of juxta straight. 9 not examined.

DISTRIBUTION. Mexico, Guatemala, Nicaragua.

DISCUSSION. This taxon is treated by Hall as a 'form' of A. atronia, with the observation
that both forms fly together at Orizaba in Mexico. In the short series present in the BMNH
and in the Hall coll. (BM, Brighton) the distinctive markings are constant and specific rank
seems to be more appropriate.

Anthanassa atronia (Bates) comb. n.

(Figs 27, 28, 245)

Melitaea atronia Bates, 1866: 113. LECTOTYPE <J, GUATEMALA: Duenas (BMNH, Type no. Rh. 8470;

Gabriel, 1927: 18), here designated [examined].
? Eeresia sydra Reakirt, 18666: 335. Syntype(s), MEXICO: near Vera Cruz (W. H. Edwards) (depository

uncertain, possibly destroyed).
Eresia obscurata Felder, 1869: 471. LECTOTYPE J, MEXICO: Cuernavaca (BMNH) here designated

[examined].

? Phyciodes diallus Godman & Salvin, 18786: 260. Syntype(s) V, PANAMA: Chiriqui (Ribbe) (MNHU).
? Phyciodes chromis Godman & Salvin, 18786: 260. Syntype(s) $, PANAMA: Chiriqui (Ribbe) (MNHU).
Phyciodes cassiopea Godman & Salvin, 18786: 262. LECTOTYPE 9, COSTA RICA: Cache (H. Rogers)

(BMNH, Type no. Rh. 8471; Gabriel, 1927: 27), here designated [examined].
Phyciodes diallus Godman & Salvin; Godman & Salvin, 1882: 207, pi. 22, figs 24, 25.
Phyciodes cassiopea Godman & Salvin; Godman & Salvin, 1882: 207, pi. 22, figs 22, 23; Rober, 1913:

441, pi. 90, row a [fig. 4] (J).

Phyciodes atronia (Bates); Godman & Salvin, 1882: 206, pi. 22, figs 19-21, Forbes, 1945: 172-178.
? Phyciodes albofascia Rober, 1913: 441, pi. 90, row b [fig. 6] (V, as albifascia). Syntype(s) ?, GUATEMALA

(depository unknown).

Phyciodes atronia atronia (Bates); Hall, 1929: 109.
Phyciodes atronia diallus Godman & Salvin; Hall, 1929: 111.

cJ forewing 17 mm, upperside very dark brown, markings paler brown, usually obscure, small white or
yellowish postdiscal spots in sib and s4 always present on both surfaces; hindwing upperside costal area
dark brown, unmarked; forewing underside medium brown, discal and postdiscal markings very obscure
or absent. 9 forewing 19-21 mm, larger, markings variable, in a pattern like A. drusilla, yellowish
(nominotypical form) or white (V -forms cassiopea, albofascia). In +-form diallus markings white, costal bar
and postdiscal spots united to form a prominent fascia, recorded by Hall as a constant form at Chiriqui.
Genitalia. _> like A. ardys, posterior margin of juxta straight in dorsal view, penis slender. f not examined.

DISTRIBUTION. Mexico, Guatemala, Costa Rica, Nicaragua, Panama.

DISCUSSION. The ^-form chromis is included here with reserve. It is not figured and the type-
material has not been examined. Described from one or more specimens in the Staudinger
Coll., perhaps similar to ^-form diallus.

REVISION OF PHYCIODES 103

Anthanassa otanes (Hewitson) comb. n.

(Figs 29, 30, 246)
Eresia otanes Hewitson, [1864]: [24], pi. [12], fig. 47.

c? forewing 18 mm, upperside dark brown with or without small yellow markings in the general pattern
of A. drusilla; hindwing leaden blue, unmarked except for a narrow black marginal border; hindwing
underside marbled brown and yellow, brown mark at apex of costa prominent. $ larger, upperside
markings more complete.

Genitalia. J tegumen and scaphial extension well defined, lateral walls partly chitinized, posterior
border of juxta sinuous with central boss, posterior section of valve rather long, harpe stout, penis
slender. 9 not examined.

DISTRIBUTION. Mexico, Guatemala, Costa Rica.
Two subspecies with similar genitalia.

Anthanassa otanes otanes (Hewitson)
(Figs 29, 30)

Eresia otanes Hewitson, [1864]: [24], pi. [12], fig. 47. LECTOTYPE ^, GUATEMALA (0. Salvin) (BMNH,

Type no. Rh. 8473; Gabriel, 1927: 91), here designated [examined].
Phyciodes otanes (Hewitson) Godman & Salvin, 1882: 208, pi. 22, figs 28, 29.
Phyciodes otanes otanes (Hewitson); Hall, 1929: 112.

c? upperside very dark, markings vestigial if present. 9 larger, upperside markings complete, white or
shaded brown, in general pattern as A. drusilla.

DISTRIBUTION. Guatemala, only in western areas (Hall).

Anthanassa otanes sop oils (Godman & Salvin)

Phyciodes sopolis Godman & Salvin, 18786: 262; Godman & Salvin, 1882: 209, pi. 22, figs 30-32.

LECTOTYPE £, GUATEMALA: Vera Paz, Choctum (Hague) (BMNH, Type no. Rh. 8474; Gabriel,

1927: 112), here designated [examined].
Phyciodes cyno Godman & Salvin, 1889: 354; Godman & Salvin, 1901: 680, pi. 108, figs 27-30. Holotype

cJ, MEXICO: Orizaba (H. J. Elwes) (BMNH, Type no. Rh. 8472; Gabriel, 1927: 37), [examined]. Syn. n.
Phyciodes dora Schaus, 1913: 345, pi. 51, fig. 9; Hall, 1929: 122; Forbes, 1945: 174. Syntype(s) 9,

COSTA RICA: Poas (? USNM). Syn. n.

Phyciodes otanes sopolis Godman & Salvin; Hall, 1929: 113.
Phyciodes otanes cyno Godman & Salvin; Hall, 1929: 114.

c? forewing like otanes otanes, but upperside yellow markings complete, discal macules sometimes enlarged
(f. cyno). 9 upperside markings well developed as in A. drusilla, forewing markings yellow (sopolis) or
white (cyno), clearly quite variable. The form dora is probably a florid modification of sopolis.
Genitalia. <$ like otanes otanes. 9 not examined.

DISTRIBUTION. Guatemala (in eastern regions), Mexico, Costa Rica.

DISCUSSION. All forms of this species are rare and material available is scanty. It is probably
that otanes and sopolis represent parts of a cline.

Anthanassa annulata sp. n.
(Figs 31, 247, 248)

cJ forewing 15 mm, apex truncate, upperside dark brown, markings like A. otanes, small yellow spots,
cell spot small, narrow oblique postdiscal mark in s4-s6, spot at base of s2 prominent, extending into
sib and s3, small round spots in sib and s4 proximal to scanty vestigial submarginal markings, fringes
chequered, brown and white; hindwing upperside dark brown, base unmarked, vestigial stria on
discoidal veins, discal and postdiscal lunules joined to form circles in slb-s3, narrow submarginal lunules
in complete series from slb-s7, fringes chequered. Underside forewing markings as above but paler buff

104 L. G. HIGGINS

and orange-brown mark present on outer margin; hindwing grey with confused brown striae, conspicuous
brown mark on costa and along outer margin to s4. V similar, larger.

Genitalia. _> tegumen very fragile, membranes, bilobed, unfortunately distorted in preparation, posterior
border of juxta concave, saccus small, posterior section of valve short, stout, harpes massive. There is
marked similarity to the organs of A. crithona.

Holotype J, Ecuador: Paramba, 3500 ft [1 160 m], iv. 1897 (Rosenberg) (genitalia slide no. 1351) (BMNH).
Paratypes. Colombia: 2 o, Rio Dagan, (Rosenberg). Ecuador: 2 3, 1 9, Cachabe, low country, 1897
(Rosenberg). (All in BMNH.)

DISTRIBUTION. Ecuador and Colombia.

^ Anthanassa crithona (Salvin) comb. n.

(Figs 32, 249-251)

Melitaea crithona Salvin, 1871: 415. LECTOTYPE 3, PANAMA: Chiriqui (Arce) (BMNH, Type no. Rh.

8477; Gabriel, 1927: 36), here designated [examined].
Phyciodes crithona (Salvin) Godman & Salvin, 1882: 206, pi. 22, figs 17, 18; Rober, 1913: pi. 89, row k

[figs 2-4]; Hall, 1929: 127.

3 forewing 17 mm, upperside dark, prominent postdiscal band yellow; hindwing dark with yellow striae
in discal, postdiscal and submarginal series. V forewing 24 mm, upperside markings paler and extended.
Genitalia. 3 in dorsal view short, wide, valve apex conical, harpe massive, posterior border of juxta
slightly concave. 9 ductus short, bursal support well defined.

DISTRIBUTION. Nicaragua, Costa Rica, Panama.

Anthanassa fulviplaga (Butler) comb, n., stat. rev.

(Figs 33, 252-254)

Phyciodes fulviplaga Butler, 1872: 77; Butler, 1874: 183, pi. 63, fig. 2; Godman & Salvin, 1882: 208, pi. 22,
figs 26, 27; Rober, 1913: 441, pi. 90, row a [figs 2, 3]; Hall, 1929: 114. LECTOTYPE 3, COSTA RICA:
Cartago (van Patten) (BMNH, Type no. Rh. 8476; Gabriel, 1927: 53), here designated [examined].

Phyciodes crithona stenotaenia Rober, 1913: 440, pi. 89, row k [fig. 8]. Holotype V, COSTA RICA: Juan
Vinas (BMNH) [examined].

Phyciodes otanes fulviplaga Butler; Forbes, 1945: 175, 190.

3 forewing 16 mm, upperside brown with large costal mark orange-yellow and small yellow spots in
postdiscal area; hindwing leaden-grey with yellow submarginal lunules in series. V forewing 21 mm,
upperside markings more extensive.

Genitalia. 3 tegumen membranes in dorsal view, posterior border or juxta with central prominence,
apical section of valve short, harpe very massive; penis apex slender. $ ductus partly chitinized, bursal
support not well defined.

DISTRIBUTION. Costa Rica, Panama.

Anthanassa hernias (Hewitson) comb, n., stat. rev.

(Figs 34, 255-257)

Eresia hernias Hewitson, [1864]: [21], pi. [11], fig. 32. LECTOTYPE 3, BRAZIL: Recife, Pernambuco, (Mrs

G. Smith) (BMNH), Type no. Rh. 8515; Gabriel, 1927: 59), here designated [examined].
Eresia conferta Felder & Felder, 1867: 394. LECTOTYPE ?, BRAZIL: Bahia, Luschnath (BMNH), here

designated [examined]. [Bears Felders' original M/S label E. conferta Feld.)

? Phyciodes brancodia Schaus, 1902: 394. Holotype J, BRAZIL: Sao Paulo (USNM, type no. 5887).
Phyciodes aequatorialis Rober, 1913: 442, pi. 90, row c [fig. 4] (? *). Syntype(s), ECUADOR (depository

unknown).
? Phyciodes aequatorialis gisela Rober, 1913: 442. Syntype(s), BRAZIL: Mato Grosso (depository

unknown).

Phyciodes conferta (Felder & Felder) Rober, 1913: 443, pi. 90, row d [fig. 8] (J).
Phyciodes hernias (Hewitson) Hall, 1929: 98.

REVISION OF PHYCIODES 105

Phyciodes (Eresia) frisia hermas (Hewitson); Forbes, 1945: 156, 172, 189.
Phyciodes frisia hermas (Hewitson); Hayward, 19646: 330, pi. 18, fig. 14 (3).

3 forewing 16 mm, upperside markings small white spots on dark brown ground-colour; V similar, usually
larger. In both sexes upperside of forewings 2 pairs of prominent white spots form a distinctive specific
character (Hall).

Genitalia. 3 in dorsal view tegumen wide, scaphial extension very short, lateral walls minutely toothed,
posterior border of juxta convex, valve apical teeth blunt, saccus massive; penis slender. y ductus partly
chitinized, bursal support not well defined.

DISTRIBUTION. Brazil, Paraguay, N. Argentina, ? Ecuador.

Anthanassa frisia (Poey) comb. rev.
(Figs 35, 258-260)

Melitaea frisia Poey, 1832: [9], pi. [2], figs (J, y). Syntype(s) c?, y', CUBA (depository unknown).

Eresia gyges Hewitson, [1864]: [24], pi. [12], figs 45, 46. LECTOTYPE <$, JAMAICA (BMNH, Type no.

Rh. 8498; Gabriel, 1927: 56), here designated [examined].
Eresia frisia (Poey); Hewitson, [1866]: iv (corrections).
Phyciodes frisia frisia (Poey); Hall, 1929: 83.

Phyciodes (Eresia) frisia frisia (Poey); Forbes, 1945: 156; Bauer, 1975: 142, pi. 44, fig. 16.
Phyciodes (eresia) frisia gyges (Hewitson); Forbes, 1945: 157.
Anthanassa frisia (Poey); Holland, 1947: 140, pi. 17, fig. 42.
Phyciodes frisia (Poey); Riley, 1975: 79, pi. 6, fig. 12 (3).

J forewing 15-16 mm, variable, upperside brown, markings orange-yellow, spot near anal in sib prominent,
usually with minute spot above it in s2; hindwing upperside prediscal band wide at costa, tapering
gradually to vie. y similar, often larger.

Genitalia. y tegumen large, scaphial extension short, lateral walls toothed, posterior border of juxta
curved; penis in side view slender. V: short ductus partly chitinised, bursal support defined.

DISTRIBUTION. Cuba, Trinidad, Jamaica, Hispaniola, Puerto Rico, Bahamas, and probably on
other islands, and U.S.A. (Texas, Florida: Key West).

Anthanassa tulcis (Bates)
(Figs 36, 261-263)

Melitaea tulcis Bates, 1864a: 82. LECTOTYPE j, GUATEMALA: Central Valleys (BMNH, Type no. Rh.

8513; Gabriel, 1927: 120), here designated [examined].

? Eresia archesilea Felder, 1869: 471. Syntypes J, y MEXICO: Cordoba (Hedemann) (? NM, Vienna).
Eresia punctata Edwards, 1871c: 191. Holotype j, {MEXICO]: 'S. Arizona' (Palmer) (CM, Pittsburgh). [See

Brown, 1966: 424, fig. 23.]

Phyciodes tulcis (Bates); Rober, 1913: 441, pi. 90, row b [fig. 8] (3).
Phyciodes frisia tulcis (Bates); Hall, 1929: 84.

Phyciodes (Eresia) frisia tulcis (Bates); Forbes, 1945: 156; Bauer, 1975: 142, pi. 44, fig. 19.
Anthanassa tulcis (Bates); Holland, 1947: 141, pi. 17, fig. 39.

c? forewing 14 mm, small but variable, like A. frisia, upperside dark grey-brown, markings very pale,
yellowish; hindwing upperside prediscal band very small or absent, postdiscal line absent, pale discal band
relatively wide, prominent. ? similar, larger.

Genitalia. 3 like frisia, in dorsal view scaphial extension wider, lateral teeth small, more numerous,
posterior border of juxta more convex, slightly irregular with low central bulge. 9 bursal duct short,
partly chitinized, support oval.

DISTRIBUTION. From U.S.A. (Texas), Mexico and Guatemala through Central America to
Panama.

106 L. G. HIGGINS

Anthanassa dubia (Hall) comb, n., stat. n.

(Figs 37, 264)

[Phyciodes frisia (Poey); Godman & Salvin, 1882: pi. 22, fig. 6; Rober, 1913: pi. 89, row e [fig. 6] (<}).

Misidentifications.]
Phyciodes frisia dubia Hall, 1929: 85. Holotype c?, VENEZUELA: San Esteban, Puerto Cabello, ex Hahnel

(BMNH; note: there may be a 'false type' of this taxon in BM, Brighton) [examined].
Phyciodes (Eresia) frisia dubia Hall; Forbes, 1945: 156.

cJ forewing 13-15 mm, like A. tulcis but upperside ground-colour brown, markings yellow, hindwing
upperside prediscal band present but short, extending to v5, discal band relatively wide, postdiscal
markings present. 9 similar, larger.
Genitalia. Like A. frisia.

DISTRIBUTION. Venezuela and coastal districts of Colombia.

Anthanassa taeniata (Rober) comb, n., stat. rev.

(Figs 38, 265)

Phyciodes taeniata Rober, 1913: 438, pi. 89, row h [figs 3, 4] (^, 9). £, 9 Syntypes ^, 9, PERU: Tarapoto

(depository unknown).

Phyciodes frisia taeniata Rober, Hall, 1929: 85.
Phyciodes (Eresia) frisia taeniata Rober; Forbes, 1945: 156.

cJ forewing 16-17 mm, upperside brown, markings bright fulvous, like frisia but with additional spot in s2
near anal angle; hindwing upperside prediscal band well developed, extending to v2, discal band, postdiscal
line and submarginal lunules all present in complete series. 9 similar, slightly larger.

Genitalia. Like A. frisia but wider, lateral shoulders better marked, scaphial extension slightly narrower,
dentate borders almost horizontal with larger teeth, saccus enlarged, posterior section of valve shorter,
posterior border of juxta straight. 9 not examined.

DISTRIBUTION. Ecuador, W. Peru. Distribution probably restricted to this area.

DISCUSSION. The four preceding species are clearly closely related and might be grouped as
subspecies (following Hall). I find this treatment not really satisfactory since the different
phenotypes are so well marked and easily distinguished, and intermediates have not been found
in the extensive series in the BMNH. Each form appears to be vicarious and allopatric. The
presence of two forms flying in the same district would confirm specific separation, but so far
this has not been recorded.

Anthanassa sosis (Godman & Salvin) comb. n.
(Figs 39, 266)

Phyciodes sosis Godman & Salvin, 18786: 262; Godman & Salvin, 1882: 204, pi. 22, figs 11, 12; Rober,
1913: 442, pi. 90, row d [fig. 1] (J); Hall, 1929: 95; Forbes, 1945: 178. LECTOTYPE rf, COSTA RICA:
Irazu, 6000-7000 ft [2000-2300 m] (H. Rogers) (BMNH, Type no. Rh. 8482; Gabriel, 1927: 112), here
designated [examined].

cJ forewing 16-17 mm, upperside black, markings small, yellow, spots arranged as in A. otanes sopolis;
hindwing upperside with 3 series of white or yellowish lunules parallel with the outer margin, hindwing
underside marbled in shades of brown and grey, t larger, markings extended.

Genitalia. J tegumen short, firmly chitinized, scaphial extension well defined, saccus narrow but deep,
valve in dorsal view shows 3 strong apical teeth, harpe slender, slightly sinuous. $ not examined.

DISTRIBUTION. Costa Rica, Panama, Colombia.

REVISION OF PHYCIODES 107

Anthanassa drymaea (Godman & Salvin) comb. n.
(Figs 40-267)

Phyciodes drymaea Godman & Salvin, 1878a: 268; Godman & Salvin, 1882: 205, pi. 22, figs 9, 10;
Rober, 1913: 442, pi. 90, row c [fig. 7] (J); Hall, 1929: 93; Forbes, 1945: 177-178. LECTOTYPE ^,
GUATEMALA: hacienda Las Nubes (0. Salvin) (BMNH, Type no. Rh. 8481; Gabriel, 1927: 42), here
designated [examined].

? forewing 16-17 mm, like A. ardys, upperside dark brown (? black when fresh) with small white
markings; forewing underside base grey-brown; hindwing upperside with curved discal series of 5 very
small white spots; underside brown marbled with grey, with darker costal mark. V slightly larger,
hindwing underside markings paler.

Genitalia. <$ organs narrow, elongate, tegumen chitinized, lateral walls gently concave, scaphial extension
very short, saccus conspicuously narrow, cleft short, valve apex with 3 teeth, one very long directed
medially, harpe short, slightly curved with a small, ventral tooth; penis slender.

DISTRIBUTION. Guatemala. Recorded at altitudes of 4000-7000 ft [1460-2560 m] at Calderas,
Purula, Las Nubes, etc.

NOTE. Can be distinguished from A. ardys (p. 98) by the very small, widely separated white discal
spots on hindwing upperside.

Anthanassa sitalces (Godman & Salvin) comb. n.
(Figs 41, 268)

Phyciodes sitalces Godman & Salvin, 1882: 201, pi. 21, figs 30, 31; Rober, 1913: 442, pi. 88, row i [fig. 5];

Hall, 1929: 95. LECTOTYPE J, GUATEMALA: Chilasco (G. C. Champion) (BMNH, Type no. Rh. 8489;

Gabriel, 1927: 111), here designated [examined].
Phyciodes sitalces saltator Hall, 1929: 96. Holotype J, MEXICO: Guerrero, Omilteme, July, 8000 ft [2600 m]

(H. Smith) (BMNH) [examined]. Syn. n.

cJ forewing 19-20 mm, like A. drymaea but larger; upperside dark brown (? black when fresh), markings
small white spots; hindwing upperside postdiscal white spots very small, the series strongly curved,
forewing underside base bright orange-brown. ? similar, slightly larger.

Genitalia. 3 like A. drymaea but less elongate, saccus wider, apical horizontal tooth of valve shorter,
curved downwards, harpe with 2 or 3 basal teeth below. 9 not examined.

DISTRIBUTION. Guatemala, Mexico. Flies at 6000-8000 ft [2000-2600 m] in mountains.

Anthanassa cortes (Hall) comb. n.
(Figs 42, 269)

Phyciodes cortes Hall, 1917: 161. Lectotype rf, MEXICO: Cuautla, 4000 ft [1300 m] June-July (A. Hall)
(BM, Brighton), designated by Hall, 1929: 93 [examined].

c? forewing 18-19 mm, upperside like A. ardys etc., but white spots are slightly larger and well defined;
hindwing upperside white postdiscal spots in series all well separated, unlike A. ardys in which the white
band is only divided by the intersection of black veins. V upperside ground-colour paler, grey-brown,
markings larger, underside forewing bases orange-brown.
Genitalia. 3 like A. sitalces, in dorsal view the teeth below the base of harpe less prominent.

DISTRIBUTION. SW. Mexico. The type— series of 11 rj, 1 ? were all taken by Hall near Cuautla.
I have not seen or heard of other material.

DISCUSSION. The species A. sosis, A. drymaea, A. sitalces and A. cortes form an interesting and
clearly related group, indicated by the presence of elaborate teeth at the apices of the valves.
All appear to be rare, three of them being represented in the BMNH by short series, while
A. sitalces is known only from the original series. The taxonomic position of A. cortes is
uncertain; possibly it is better graded as a subspecies with A. sitalces, but the difference in
external characters is most definite.

108 L. G. HIGGINS

DAGON gen. n.

Type-species: Eresia catula Hopffer. Gender: masculine.

Small butterflies, forewing 13-15 mm, outer margin straight; hindwing outer margin slightly wavy; in all
species upperside dark brown or black with small white or yellow markings; underside markings more
variable, hindwing with small ocellar postdiscal spots in slc-s6. Sexes similar.

Genitalia. J organs narrow, elongate, tegumen well defined, posterior border of juxta convex, saccus
prominent, elongate, broad, widest at its apex which is broadly notched, subapical tooth present before
valve apex; penis slender, ostium-kneel absent (D. catula and D. pusillus).

DISTRIBUTION. NW. Argentina (Salta), S. Peru, Bolivia.

DISCUSSION. The four small butterflies included in this genus form an interesting group, the males
being distinguished at once by their narrow, elongate genitalia which are similar in all species. The
sharply pointed twin teeth at the apices of the valves, and the slender penes, suggest relationship
with Anthanassa, but this is not confirmed by the shapes of the wings (the forewings with
straight outer margins), the pattern of wing markings, and especially by the long, wide saccus,
with no more than a shallow terminal notch. Only a single specimen is known of the little
species fontus, and the genitalia have not been examined, but judging by the external characters
it probably belongs to this genus.

Forbes (1945) keys out three of the species as his group 13; the fourth species (morenus)
he includes in his key to combined groups 11 and 15.

NOTE. The mythological Dagon was like a mermaid — 'upwards man and downwards fish'. The
name is proposed with allusion to the extraordinary development of the saccus, shaped like the
tail of a fish, in this small and obscure genus.

Key to species of Dagon (based on Forbes, 1945)

1 Upperside forewing dark brown, almost unmarked except for traces of submarginal lunules

morenus (p. 109)
Dark upperside forewing with small yellowish white discal spots ...... 2

2 Spot in hindwing cell sib isolated, in line with postmedian band, the spot in s2 offset outward

catula (p. 109)
Hindwing pattern not as above 3

3 Postmedian spots in hindwing cells s2 and sib in line, that in cell s3 offset outward

pusillus (p. 108)

Postmedian spots in cells slb-s3 all in line, that in s4 offset outward; underside much paler,
yellowish fontus (p. 109)

Dagon pusillus (Salvin) comb. n.
(Figs 43, 270, 271)

Eresia pusilla Salvin, 1869: 172. LECTOTYPE J, PERU: Cosnipata Valley (H. Whitely) (BMNH, Type no.

Rh 8533; Gabriel, 1927: 101), here designated [examined].

Eresia calena Hopffer, 1874: 350, Syntypes 3, PERU: Chanchamayo (Thamm) (? MNHU, Berlin).
Eresia adoxa Erschoff, 1874: 142, pi. 3, fig. 5. Syntypes J, PERU: near Monterico (? ZI, Leningrad).
Phyciodes calena (Hopffer) Rober, 1913: 445, pi. 90, row g [fig. 2] (^).
Phyciodes pusilla (Salvin) Hall, 1929: 118.
Phyciodes (Tritanassa) pusilla (Salvin); Forbes, 1945: 186, 190.

3 forewing rather elongate, hindwing upperside discal white stripe narrow, postdiscal and submarginal
markings vestigial, if present.

Genitalia. S in dorsal view with tegumen short, fragile, posterior margin of juxta strongly convex,
saccus elongate, valve apex short, stout, harpe relatively massive; penis slender, f not available.

DISTRIBUTION. Peru, Bolivia.

REVISION OF PHYCIODES 109

Dagon catula (Hopffer) comb. n.

(Figs 44, 272, 273)

Eresia catula Hopffer, 1874: 350. Holotype J, 'BOLIVIA' ( Warscew icz) (? MNHU, Berlin) [not examined;

see note below].

Phyciodes catula (Hopffer) Rober, 1913: 445, pi. 90, row g [fig. 4].
Phyciodes minima Rober, 1913: 445, pi. 90, row g [fig. 3]. Syntype(s), ARGENTINA: Salta (depository

unknown). Syn. n.
Phyciodes catula extincta Rober, 1913: 445, pi. 90, row g [fig. 5]. Syntype(s), PERU (? MNHU, Berlin).

Syn. n.

Phyciodes abasina Staudinger (in litt.); Rober, 1913: 445 (invalid: proposed in synonymy with catula).
Phyciodes birivula Dyar, 19136: 632. Holotype rf, PERU: San Miguel, 6000 ft [2000 m] (USNM,

Washington). Syn. n.
Phyciodes catula catula ab. fulvocincta Hall, 1929: 117. Holotype v>, PERU/BOLIVIA boundary (BMNH)

[examined].

Phyciodes catula catula (Hopffer); Hall, 1929: 117; Forbes, 1945: 187.
Phyciodes catula birivula Dyar; Hall, 1929: 118; Forbes, 1945: 186.
Phyciodes catula minima Rober; Hall, 1929: 118; Forbes, 1945: 186.
Phyciodes catagraphus Hay ward, 1933: 214, 2 figs. Holotype 9, ARGENTINA: Rosario de la Frontera

(A. Breyer) (? IML, Tucuman). [Placed by Hayward, 1949: 12, as synonym of P. teletusa signata.]

Syn. n.

cJ forewing 15 mm, like D. pusillus but broader (less elongate), upperside black, markings small, white or
yellow; hindwing upperside postdiscal and submarginal markings present. ? larger, similar to 3 but
ground-colour variable, white to orange-brown (rare).

Genitalia. $ in dorsal view like D. pusillus but valve apex more slender, lower (apical) tooth long,
sharply curved inwards, harpe slender, undulant, tegumen defined but fragile, short, posterior margin of
juxta strongly convex, saccus long, wide; penis slender. $ not examined.

DISTRIBUTION. Peru, Bolivia, W. Argentina.

NOTE. According to Prof. K. S. Brown (in litt.) the Warscewicz material probably came from
W. Colombia, not Bolivia; clearly it is important to examine the holotype of catula at the first
opportunity.

Dagon morenus (Rober) comb. n.

(Figs 45, 274)

Phyciodes morena Rober, 1913: 443, pi. 90, row d [fig. 7]; Hall, 1929: 126, pi. 2, fig. 3; Forbes, 1945:
180. syntype(s) .3, PERU: Cuzco (MNHU, Berlin).

d forewing 15 mm, upperside dark brown, almost unmarked, both wings with traces of submarginal
lunules, hindwing with obscure, yellowish postdiscal ocelli. ? unknown.
Genitalia. 3 organs like D. catula, apparently without individual specific characters.

DISTRIBUTION. Peru, Cuzco Province.

NOTE. There are 2 J in the BMNH and 3 3 in coll. Hall (BM, Brighton), one of which has
been dissected to make the preparation shown in the genitalia figure.

Dagon fontus (Hall) comb. n.
(Fig. 46)

Phyciodes fontus Hall, 1928a: 11; Hall, 1929: 75, pi. 2, fig. 2. Holotype 3, GUYANA (BMNH) [examined].
Phyciodes (Tritanassa) fontus Hall; Forbes, 1945: 186, 190.

3 forewing 14 mm, narrow, upperside black, markings yellowish white, several small discal spots as in
D. pusillus, with suggestion also of a pale costal bar and an isolated small postdiscal spot in s5; hindwing
upperside black, pale discal band narrow ; underside general colour scheme pale brown, markings yellowish.
Genitalia. Not examined, abdomen missing.

DISTRIBUTION. The single known specimen is labelled 'British Guyana'.

1 10 L. G. HIGGINS

TELENASSA gen. n.

Type-species: Argynnis teletusa Godart. Gender: feminine.

Rather small butterflies, outer margin of forewing excavate or entire, upperside usually with well-developed
subapical bar; hindwing upperside usually with 3 rows of lunules parallel to outer margin, underside
markings variable. Sexes similar or nearly so. Specific characters well marked.

Genitalia. d in dorsal view with tegumen short, fragile; posterior border of juxta often with central
prominence, valve long, narrow, apex curved inwards to terminate in a single point; saccus deeply cleft;
penis straight, robust, distal section short, about one-third total length or little more, morula prominent
but ostium-keel not developed. 9 genitalia with ductus very short, bursal support cup-shaped.

DISTRIBUTION. The species occur over an extensive area in tropical South America, northwards
to Colombia, but apparently absent from Central America.

NOTE. Of the species of Telenassa represented in the BMNH, eight lack females, although
in some cases the males appear to be common.

Key to species of Telenassa

Note. It has not been practicable to include T. burchelli (p. Ill) in this key.

1 Forewing upperside with orange markings

Forewing upperside without orange markings 5

2 Forewing orange subapical band wide, entire

Forewing not so marked • 4

3 Hindwing orange submarginal lunules vestigial jana (p. 112)

Hindwing orange submarginal lunules developed elaphina (p. 113)

4 Forewing subapical band broken trimaculata (p. 1 14)

Forewing discal band broken delphia (p. 115)

5 Hindwing with orange discal band 6

Hindwing without discal band 8

6 Forewing with cell-spot well developed signata (p. 112)

Forewing with cell-spot vestigial or absent

7 Forewing upperside postdiscal spot in si, s2, s3 small, not extending to inner margin

berenice (p. Ill)
Forewing upperside postdiscal spot in si, s2, s3 large, extending to inner margin . . 9

8 Forewing upperside with proximal border of postdiscal spot vertical to inner margin

teletusa (p. Ill)

Forewing upperside markings white 10

Forewing upperside lacking white markings 12

10 Forewing with white subapical band nana (p. 113)

Forewing not so marked . . 11

11 Forewing with white submarginal spot in s3, but lacking subapical band; hindwing dark

nussia (p. 113)
Forewing not so marked, hindwing with white discal band abas (p. 112)

12 Forewing apex truncate • sepulta (p. 115)

Forewing apex not truncate 13

13 Forewing underside crossed by 4 or 5 dark lines catenaria (p. 114)

Forewing not so marked 14

14 Hindwing upperside with wide yellowish discal band flavocincta (p. 1 14)

Hindwing not so marked 15

15 Forewing underside with 2 dark postdiscal bands ....... gaujoni (p. 114)

Forewing underside postdiscal area dark, bands not defined not us (p. 113)

REVISION OF PH YCIODES 1 1 1

Telenassa teletusa (Godart) comb. n.

(Figs 47, 275-277)

Argynnis teletusa Godart, [1824]: 817. Syntype(s), BRAZIL (depository unknown).

Phyciodes teletusa teletusa (Godart); Hall, 1929: 76.

Phyciodes (Tritanassa) teletusa (Godart); Forbes, 1945: 183-184, 190.

? Phyciodes teletusa douglasi Bryk, 1953: 87. Holotype <3, PERU: Roque (NR, Stockholm).

Phyciodes teletusa (Godart); Hayward, 19646: 350, pi. 18, fig. 15.

Upperside dark brown (? black when fresh), markings deep yellow; $ forewing 16-18 mm, an erect
postdiscal mark in si, s2, s3, extends from middle of inner margin to v3, the proximal border vertical to
inner margin; transverse band of hindwing 3-4 mm wide, often tapering slightly and shading to white
at inner margin. 9 similar, often larger and markings slightly paler, forewing usually with obscure pale
mark at cell-end.

Genitalia. J in dorsal view: posterior border of juxta straight with a small central prominence, all
generic characters well developed. 9 chitinized bursal duct short, bursal support well defined.

DISTRIBUTION. Central and S. Brazil, Peru.

Telenassa burchelli (Moulton) comb, n., stat. rev.

(Figs 49, 278, 279)

Phyciodes burchelli Moulton, 1909: 100. Holotype 3, BRAZIL: Meiaponte to S. Joaquim (Joaq. Alves/W. J.

Burchell) (UM, Oxford, Type no. 1 108) [examined].

[Phyciodes teletusa (Godart); Rober, 1913: pi. 89, row h [fig. 5] (d). Misidentification.]
Phyciodes teletusa peruana Rober, 1913 : 438, pi. 89, row h [fig. 7]. Syntype(s) 3, PERU (depository unknown).
Phyciodes teletusa burchelli Moulton; Hall, 1929: 78; Forbes, 1945: 182-183.

cJ forewing 15-16 mm, narrow, outer margin straight, upperside markings yellow, like T. teletusa, usually
smaller, postdiscal mark in si, s2 and s3, proximal border oblique, obscure spot at cell-end often present;
hindwing upperside transverse band wide, usually extending to postdiscal area. 9 similar, larger.
Genitalia. Like T. teletusa.

DISTRIBUTION. Brazil, in Dept. Goias and westwards through Mato Grosso to Peru. Recorded
from Bolivia and Ecuador.

DISCUSSION. Hall records this species flying at low altitudes in the Amazon valleys, while
T. berenice was present at high altitudes, not below 1,100 m (but see also Forbes, 1945: 183).
T. burchelli shows little variation in the series of 49 ^, 2 $ in the BMNH in spite of its wide
distribution.

Telenassa berenice (Felder & Felder) comb, n., stat. rev.

(Figs 48, 280, 281)

Eresia berenice Felder & Felder, 1862a: 110. LECTOTYPE (9?), PERU: 'Rio Negro' (BMNH), here

designated [examined]. [Specimen bears Felders' M/S determination label.]
Phyciodes berenice (Felder & Felder) Rober, 1913: 438, pi. 89, row i [fig. 1] (j).
? Phyciodes drusinilla Rober, 1913: 441, pi. 90, row a [fig. 8]; Hayward, 19646: 337. Syntype(s) <J,

ARGENTINA (depository unknown).
Phyciodes teletusa berenice (Felder & Felder); Hall, 1929: 78; Forbes, 1945: 182-184.

c? forewing 16 mm, upperside dark brown, basal area unmarked, postdiscal area markings yellow like
T. teletusa but small, costal bar short, postdiscal mark in s2 not large, usually extends into sib and s3, but
not reaching inner margin, small submarginal marks in s3 and in s4 present; hindwing upperside transverse
band narrow, 2 mm or less, thin postdiscal line and submarginal lunules present. 9 similar.

Genitalia. Like T. teletusa, perhaps a little wider; tegumen and scaphial extension differ, the latter less
well defined, bilobed. 9 not examined.

DISTRIBUTION. Peru (Dept. Amazonas), Ecuador, Bolivia?, Argentina?.

112 L. G. HIGGINS

Telenassa signata (Hall) comb, n., stat. n.

(Figs 50, 282)

Phyciodes teletusa signata Hall, 1928a: 11; Hall, 1929: 77; Forbes, 1945: 182; Hayward, 1949: 12; 1964ft:
352, pi. 18, fig. 16 (<^). Holotype rf, ARGENTINA: Salta, Sierra de Aconquija, 3300 ft [1200 m] (BM,
Brighton) [examined].

<3 forewing 15-16 mm, like T. teletusa but upperside markings slightly reduced, paler yellow; forewing
with oblique prominent yellow mark at cell-end ; hindwing yellow discal band narrow, not tapered at inner
margin; hindwing underside pale yellow-buff, markings faint, lacking dark clouding along outer margin
in s2-s5. ? not seen.
Genitalia. <3 like T. teletusa but wider in dorsal view, tegumen shorter, posterior section of valve slender.

DISTRIBUTION. Argentina (Jujuy, Salta, Catmarca), Bolivia (Cochabamba).
NOTE. For distinction from Ortilia sejona, see p. 118.

Telenassa abas (Hewitson) comb. n.
(Figs 51, 283, 284)

Eresia abas Hewitson, [1864]: [22], pi. [11], figs 37, 38. LECTOTYPE £, COLOMBIA: New Grenada,

Bogota (BMNH, Type no. Rh. 8512; Gabriel, 1927: 5), here designated [examined].
Phyciodes abas (Hewitson) Rober, 1913: 443, pi. 90, row e [figs 6, 7]; Hall, 1929: 115.
Phyciodes (Eresia) abas (Hewitson); Forbes, 1945: 160-161, 189.

cJ forewing 18 mm, outer margin very slightly excavate, upperside markings white, cell-spot often slightly
elongate, placed a little before the disco-cellular vein; hindwing upperside discal band 2-3 mm wide,
extending to sic but not reaching inner margin, postdiscal yellowish line and white submarginal lunules
usually vestigial. $ similar, markings often better developed.

Genitalia. 3 in dorsal view like T. teletusa, tegumen and scaphial extension fragile, posterior border of
juxta gently convex, terminal section of valve short, harpe rather slender. $ not examined.

DISTRIBUTION. Colombia, not seen from elsewhere.

NOTE. This species is often confused (e.g. Rober, 1913; Hall, 1929) with Janatella fellula. The
latter can be identified by the white discal band on the upperside of the hindwing, which is
continued to reach the inner margin. Specimens of T. abas with the white markings slightly
reduced may be mistaken for A. ardys subota.

Telenassa jana (Felder & Felder) comb. n.
(Figs 52, 285, 286)

Eresia jana Felder & Felder, 1867: 394. LECTOTYPE rf, COLOMBIA: Bogota (Undig (BMNH) here
designated [examined]. [Specimen bears original M/S Felder determination label.]

Eresia elaphiaea Hewitson, [1868]: [33], pi. [19], figs 50, 51. LECTOTYPE rf, ECUADOR (BMNH, Type no.
Rh. 8464; Gabriel, 1927: 43), here designated [examined]. Syn. n.

Phyciodes jana (Felder & Felder) Rober, 1913: 439.

Phyciodes elaphiaea (Hewitson) Rober, 1913: 439, pi. 89, row i [fig. 7] (^).

Phyciodes jana jana (Felder & Felder); Hall, 1929: 129; Forbes, 1945: 190.

Phyciodes jana elaphiaea (Hewitson); Hall, 1929: 129; Forbes, 1945: 190.

c? forewing 17-19 mm, upperside dark brown with wide orange subapical band, hindwing dark, triple series
of submarginal lunules rather faintly marked. ? similar, forewing 20 mm, markings paler.
Genitalia. Like T. teletusa in both sexes.

DISTRIBUTION. Colombia, Peru, Bolivia, Ecuador.

DISCUSSION. It is difficult to maintain a subspecific distinction between nominate T. jana from
Colombia and specimens from Bolivia and Peru, referred by Hall to elaphiaea.

REVISION OF PHYC10DES 113

Telenassa elaphina (Rober) comb. n.

(Figs 53, 287)

Phyciodes elaphiaea elaphina Rober, 1913: 439, pi. 89, row i [fig. 9]. Syntype(s) 3, BOLIVIA (depository

unknown).
Phyciodes elaphina Rober; Hall, 1929: 130; Forbes, 1945: 180, 190.

3 (brewing 19-20 mm, outer margin deeply excavate, like T. jana, but upperside subapical band paler
yellow, slightly irregular; hindwing upperside marginal lunules yellowish, usually well developed. 9
unknown.
Genitalia. £ in dorsal view like T. jana Felder.

DISTRIBUTION. Bolivia.

Telenassa nana (Druce) comb. n.

(Figs 54, 288, 289)

Phyciodes nana Druce, 1874a: 156. LECTOTYPE 3, PERU: Cosnipata Valley (Whitely) (BMNH, Type

no. Rh. 8480; Gabriel, 1927: 84), here designated [examined].
? Eresia geminia Hopffer, 1874: 351. Syntypes 3, PERU: Chanchamayo (Thamm), BOLIVIA: Moxos (Pavon)

(? MNHU, Berlin).

? Phyciodes omosis Dyar, 1913b: 632. Syntypes, PERU: Pampaconas River (USNM, Washington).
Phyciodes nana Druce; Rober, 1913: 439, pi. 89, row i [fig. 6].
Phyciodes geminia (Hopffer); Rober, 1913: 439, pi. 89, row i [figs 4, 5].
Phyciodes nana nana Druce; Hall, 1929: 131; Forbes, 1945: 180.
Phyciodes nana omosis Dyar; Hall, 1929: 131; Forbes, 1945: 180.

3 forewing 17-18 mm, upperside dark brown, subapical band white or yellowish (omosis), variable;
hindwing upperside with triple series of lunules, postdiscal and submarginal lunules rather obscure, greyish.
? not seen.
Genitalia. 3 like T. teletusa but larger, more robust; penis massive.

DISTRIBUTION. Peru, Bolivia.

Telenassa nussia (Druce) comb. n.
(Figs 55, 290)

Phyciodes nussia Druce, 1876: 222; Rober, 1913: 439; Hall, 1929: 132; Forbes, 1945: 180. LECTOTYPE
3, PERU (BMNH, Type no. Rh. 8478; Gabriel, 1927: 88), here designated [examined].

d forewing 17 mm, like T. nana but on upperside the white subapical band is replaced by a single white
submarginal spot in s3. Perhaps better graded as a local form of T. nana.
Genitalia. £ like T. nana. $ not available.

DISTRIBUTION. Peru: Chachapoyas (BMNH series 11, 1 J, all from the same locality). Hall has
recorded a specimen (very doubtfully) from Paraguay.

Telenassa notus (Hall) comb. n.
(Figs 56, 291-293)

Phyciodes notus Hall, 1917: 163. Holotype 3, PERU: Pozzuzo (BM, Brighton) [examined].

Phyciodes notus notus Hall, 1929: 123, pi. 2, fig. 1 (<^).

Phyciodes notus f. pullopicta Hall, 1929: 124. Holotype 3, PERU: Cushi, Huanuco Province (BMNH)

[examined].
Phyciodes (Tritanassa) notus Hall; Forbes, 1945: 179, 190.

3 forewing 16-17 mm, upperside dark brown, 2 small postdiscal spots in s4 and s6. ^ forewing with 2
white subapical spots and fulvous submarginal spot in s3; hindwing with 4 fulvous discal spots.
Genitalia. 3 in dorsal view like Telenassa teletusa, tegumen fragile.

DISTRIBUTION. Peru, Bolivia.

114 L. G. HIGGINS

Telenassa gaujoni (Dognin) comb. n.
(Figs 57, 294)

Phyciodes gaujoni Dognin, 1887: 175, fig.; Hall, 1929: 125; Forbes, 1945: 179, 183, 190. Lectotype <J,
ECUADOR: Zamora (BMNH), designated by Hall, 1929: 125 [examined].

c? [brewing 14-17 mm, outer margin slightly excavate, upperside dark grey, submarginal lunules yellowish,
preceded by a double series of slender lunules which tend to fuse, forming ocellar spots; underside marbled
brown on pale grey, on forewing postdiscal area dark with confused marbling, on hindwing dark
submarginal spots in s2-3-4 prominent. 9 unknown.
Genitalia. <$ in dorsal view like T. teletusa.

DISTRIBUTION. Ecuador, Colombia.

Telanassa trimaculata (Hewitson) comb. n.
(Figs 58, 295)

Eresia trimaculata Hewitson, 1869a: 28 [index]. LECTOTYPE^, ECUADOR: Rio Verde (Buckley) (BMNH,

Type no. Rh. 8458; Gabriel, 1927: 120), here designated [examined].
Phvciodes taphius Godman & Salvin, 1878/r. 263. LECTOTYPE cJ, ECUADOR: Canelos (Buckley) (BMNH,

Type no. Rh. 8459; Gabriel, 1927: 116), here designated [examined].
Phyciodes elaphiaea abrupta Rober, 1913: 439, pi. 89, row i [fig. 8]. Syntype(s) rf, ECUADOR/BOLIVIA:

'Andes' (depository unknown).
Phyciodes trimaculata (Hewitson) Hall, 1929: 127; Forbes, 1945: 180, 183, 190.

c? forewing 15-17 mm, outer margin slightly excavate; upperside dark brown, markings yellow, an
oblique subapical band present, formed of small spots ; hindwing with ocellate postdiscal markings and
lunulate submarginal line. 9 not available.
Genitalia. <$ like T. teletusa.

DISTRIBUTION. Ecuador, Bolivia.

Telenassa flavocincta (Dognin) comb. n.
(Figs 59, 296, 297)

Phyciodes flavocincta Dognin, 1887: 174, fig.; Hall, 1929: 119; Forbes, 1945: 179, 183, 190. Lectotype 9,
ECUADOR: Zamora (BMNH), designated by Hall, 1929: 120 [examined].

$ forewing 16-17 mm, outer margin entire, both wings upperside brown, with postdiscal areas and
submarginal lunules orange-yellow. 9 not seen.

Genitalia. $ in dorsal view rather massive, like T. nana, tegumen short, bilobed scaphial extension
short and very fragile.

DISTRIBUTION. Ecuador, N. Peru (Rio Tabacones, 2000 m).

Telenassa catenaria (Godman & Salvin) comb. n.
(Figs 60, 298)

Phyciodes catenarius Godman & Salvin, 1880: 123, 131, pi. 4, fig. 11; Hall, 1929: 124; Forbes, 1945: 179,
190. Holotype ^, COLOMBIA: Sierra Nevada de Santa Marta, Chinchicua Valley (Simons) (BMNH,
Type no. Rh. 8465; Gabriel, 1927: 27), [examined].

cJ forewing 17 mm, upperside dark grey-brown, markings yellow-grey, delicate, 2 rings in sib near anal
angle, continued with vestigial lunules to costa, discal markings vestigial if present, series of submarginal
lunules complete; hindwing upperside postdiscal and premarginal lunules combine to form a complete
series of postdiscal rings, submarginal lunules well defined. Underside pale buff, forewing crossed by brown
lines, with small, blind ocelli in s5 and in s6. 9 similar, slightly larger, forewing upperside with red-brown
postdiscal shade.

REVISION OF PHYCIODES 115

Genitalia. £ like T. teletusa. 9 not examined.

DISTRIBUTION. Colombia: Sierra Nevada de Santa Marta; recently also found in the Cordillera
Occidental (G. Bernard & M. Adams).

Telenassa delphia (Felder & Felder) comb. n.
(Figs 61, 299)

Eresia delphia Felder & Felder, 1861: 103. LECTOTYPE 9, COLOMBIA: near Muzo, Bogota Province
(Uricoechea) (BMNH), here designated [examined]. [Specimen bears original label of Felders'.]

Eresia aceta Hewitson, [1864]: [23], pi. [12], figs 39, 40. LECTOTYPE $, COLOMBIA: 'New Grenada'
(BMNH, Type no. Rh. 8467; Gabriel, 1927: 6), here designated [examined].

Phyciodes delphia delphia (Felder & Felder); Rober, 1913: 440, pi. 89, row k [fig. 1].

Phyciodes delphia aceta (Hewitson); Rober, 1913: 440.

Phyciodes delphia (Felder & Felder); Hall, 1929: 120; Forbes, 1945: 183, 190.

c? forewing 15-17 mm, outer margin slightly excavate, upperside dark brown, irregular broken discal band
fulvous, and dark-pupilled postdiscal ocelli of similar colour in series across both wings, followed by
submarginal lunules. 9 slightly larger with markings better developed.
Genitalia. $ like T. teletusa. 9 not examined.

DISTRIBUTION. Colombia: Muzo, El Baldio (5000 ft [1600 m]), Ecuador?

NOTE. In the BMNH there is a single specimen with upperside dark grey-brown and prominent
unpupilled ocellar postdiscal spots, otherwise unmarked. It is clearly closely related to T. delphia,
and possibly represents an unnamed species. Since only a single specimen is known, I have not
named it at present.

Telenassa sepulta (Hall) comb. n.
(Figs 62, 300, 301)

Phyciodes sepultus Hall, 1928a: 12; Hall, 1929: 125, pi. 2, fig. 12; Forbes, 1945: 179, 190. Holotype <J,
PERU: Chachapoyas (de Mathan) (BMNH) [examined].

c? forewing 19 mm, outer margin excavate, apex truncate and apical angle acute; upperside pale postdiscal
and submarginal lunules obscure; underside yellow-brown, markings better defined, forewing with 4 pale
postdiscal spots. 9 not known.

Genitalia. Like T. teletusa, 3 base wide, tegumen and scaphial extension well defined, penis with usual
generic characters.

DISTRIBUTION. Peru.

ORT1L1A gen. n.

Type-species : Papilio liriope Cramer. Gender : feminine.

Rather small butterflies, uppersides usually dark brown or black with yellow or white markings. Specific
characters are often poorly defined in the male genitalia. On the forewing upperside a large white or yellow
postdiscal spot is present in sib, s2, and s3, in several species. Markings and genitalia are somewhat
atypical in the divergent species ithra.

Genitalia. 3 tegumen usually short, fragile, sometimes vestigial, terminal section of valve curved inwards
and a large pre-apical tooth is present in all species, forming a bifid or 'double' apex, harpe variable,
exceptionally long in liriope and in gentina, saccus always single (entire), penis slender, ostium-keel
absent. 9 examined in all species, bursal support shaped like an egg-cup, and often arising directly from
a deeply excavated pocket in the genital plate, post-vaginal scutum large.

DISTRIBUTION. French Guiana, Guyana, Surinam, Brazil, Argentina, Bolivia, W. Peru (0. gentina
only).

DISCUSSION. The genus is especially characteristic of the eastern regions of South America.
In genitalia preparations the twin apical teeth at the valve apices may prove difficult to display.

116 L. G. HIGGINS

Key to species of Ortilia

2 Upperside markings fulvous 2

Upperside markings chiefly white , 9

2 Fulvous areas very extensive 3

Fulvous areas macular, not extensive 4

3 In dorsal view valve apex almost straight liriope (p. 116)

In dorsal view valve apex strongly curved gentina (p. 116)

4 Hindwing upperside discal band tapered to inner margin 5

Hindwing discal band wide, not tapered 6

5 Small, forewing 16 mm, hindwing discal band not well defined . . . . polinella (p. 119)
Large, forewing 19 mm, hindwing discal band well defined zamora (p. 119)

6 Forewing elongate dicoma (p. 119)

Forewing not elongate 7

7 Upperside suffused fuscous, markings obscure velica durnfordi (p. 118)

Upperside markings well defined ' . . 8

8 Upperside markings yellow, cell-spot prominent, well formed sejona (p. 118)

Upperside markings orange-brown, cell-spot, if present, not prominent, small

velica velica (p. 118)

9 Upperside hindwing submarginal spot in s6-7 fulvous orthia (p. 117)

Upperside hindwing with this spot white 10

10 Upperside forewing postdiscal spot in s4 prominent ithra (p. 120)

Upperside forewing postdiscal spot in s4 lacking orticas (p. 117)

Ortilia liriope (Cramer) comb. n.
(Figs 63, 302-304)

Papilio liriope Cramer, 1775: 2, pi. 1, figs C, D. Syntype(s), SURINAM (depository unknown).

? Argynnis liriope (Cramer); Fabricius, 1807: 285.

? Dryas reticulata liriope (Cramer); Hiibner, 1808: pi. 40.

Phydodes liriope (Cramer); Rober, 1913: 435, pi. 89, row a [figs 1, 2]; ? Hayward, 1964b: 359, pi. 18,

fig. 10.

? Phydodes liriope liriope (Cramer); Hall, 1929: 64.
Phydodes (Tritanassa) liriope (Cramer); Forbes, 1945: 157-158, 188, 190.

c£ forewing 16-17 mm, upperside rather dark fulvous, markings dark brown (? black when fresh); hindwing
underside with an irregular reddish discal line. $ similar, slightly larger.

Genitalia. <$ tegumen short, fragile, difficult to define; valve with distal section short, apex slightly
inturned, harpe rather long; saccus narrow; penis straight, suspensory ligament near mid-point. 9 genital
plate small, ductus not clearly chitinised, bursal support relatively large.

DISTRIBUTION. French Guiana, Guyana, Surinam and N. Brazil.

DISCUSSION. This species is easily confused with Tegosa orobia, but liriope can be identified by
the slightly wavy red line across the discal area on the underside of the hindwing. Forbes (1945:
188-189) discusses the confusion between liriope and claudina (in which he includes orobia).

Ortilia gentina sp. n.

(Figs 64, 305, 306)

c£ forewing 14-17 mm, upperside ground-colour fulvous-yellow, markings black, slightly variable, wing
borders black, 2-5 mm or even more, prominent, subapical band rather irregular, often imperfectly defined
medially, extending into submarginal area in s3, some dark striae across cell; hindwing upperside basal
area slightly dusky, dark marginal border prominent, including poorly defined submarginal lunules in
some specimens, underside postdiscal spots usually well marked.

Genitalia. J in dorsal view with organs wider, distal section of valve longer, apex slender, strongly
curved inwards, harpe placed further forwards at centre of valve. $ not examined.

REVISION OF PHYCIODES 117

Holotype £, Brazil: Sao Paulo, Anhangahy, xi. 1926 (R. Spitz genitalia slide no. 2698) (BMNH).

Paratypes. Brazil: 1 J, Mato Grosso (genitalia slide no. 826). Bolivia: 1 3, no further data (genitalia
slide no. 1082); 1 ,3, Santa Cruz de la Sierra (genitalia slide no. 825); 1 3, Bueyes (genitalia slide no.
1081). Argentina: 1 3, Tucuman (genitalia slide no. 1083).

DISTRIBUTION. Brazil, Argentina (Tucuman), Peru, Bolivia, Paraguay. Collected in November,
February, April, July. Probably emerges in two or perhaps three annual broods.

DISCUSSION. This species resembles 0. liriope closely. Distinctive features include the paler
upperside ground colour, and on the hindwing the better-defined basal dark area. On the
hindwing underside the reddish oblique line is prominent in liriope, and the series of postdiscal
spots is obvious in gentina, while the oblique line is vestigial and is not prominent. The male
genitalia are easily distinguished.

Ortilia orthia (Hewitson) comb. n.
(Figs 65, 307)

Eresia orthia Hewitson, [1864]: [19], pi. [10], figs 21, 22. LECTOTYPE J, BRAZIL: Minas Geraes (BMNH,

Type no. Rh. 8449; Gabriel, 1927: 91), here designated [examined].
Phyciodes poltis Godman & Salvin, 18786: 261; Godman & Salvin, 1882: 200, pi. 21, figs 28, 29; Rober,

1913:438, pi. 88, row i [fig. 3] (very poor fig.); Hall, 1929: 81). LECTOTYPE J [BRAZIL: Minas Geraes]

(BMNH, Type no. Rh. 8450; Gabriel, 1927: 98), here designated [examined]. [For locality see Hall,

1929: 81.]
Phyciodes orthia (Hewitson) Rober, 1913: 438, pi. 89, row h [figs 8-10]; Hall, 1929: 80; Hayward, 19646:

346, pi. 18, fig. 17.

Phyciodes orthia ab. evanescens Rober, 1913: 438. Syntype(s), [? BRAZIL] (depository unknown).
Phyciodes orthia f. roeberi Hall, 1935: 221. Holotype *, BRAZIL: Santa Maria, Rio Grande do Sul, 1200 ft

[420 m], i. 1920 (A. Hall) (BM, Brighton).
Phyciodes (Tritanassa) orthia (Hewitson); Forbes, 1945: 177, 190.

•$ forewing 15 mm, upperside dark brown, markings pale yellow, sometimes shading to white, cell-spot well
developed, postdiscal spots in sib, s2 and s3 rather small with smaller spot placed more distally and well
separated in sib; hindwing discal band usually darker orange anteriorly in s7. 1 larger, submarginal
markings often better defined.

Genitalia. J like 0. liriope, in dorsal view slightly narrower, harpe shorter, straight, posterior border of
juxta gently convex.

DISTRIBUTION. S. Brazil, Paraguay, Argentina.

NOTE. The form evanescens (black and white) is mostly found in Minas Gerais (Prof. Keith
Brown, pers. comm.).

Ortilia orticas (Schaus) comb. n.
(Figs 71, 308-309)

Phyciodes orticas Schaus, 1902: 394; Rober, 1913: 445. Holotype J1, BRAZIL: Castro, Parana (USNM,

Type no. 5888) [examined].
Phyciodes orticas orticas Schaus; Hall, 1929: 105.
Phyciodes (Tritanassa) orticas orticas Schaus; Forbes, 1945: 182, 190.

o forewing 18 mm, upperside black, markings white, cell-spot minute, vestigial, oblique bar in s5-s7 short,
postdiscal spot in slb-s3 large; hindwing upperside white discal band crossed by black veins, submarginal
lunules vestigial or absent; forewing underside base orange-fulvous. V similar, slightly larger.

Genitalia. J in dorsal view like O. orthia but base wider, discal section of valve longer. 9 bursal support
well chitinized.

DISTRIBUTION. Brazil (Parana, Rio de Janeiro, etc.), Argentina (Entre Rios).

118 L. G. HIGGINS

Ortilia sejona (Schaus) comb, n., sp. rev.

(Figs 66, 310)

Phyciodes sejona Schaus, 1902: 393; Rober, 1913: 438, pi. 89, row i [figs 2, 3] (£, 9). Holotype 9, BRAZIL:

Sao Paulo (USNM, Type no. 5886).
[Phyciodes velica velica (Hewitson); Hall, 1929: 81. Misidentification.]

Upperside brown, markings yellow, like Telenassa teletusa. <$ forewing 16-17 mm, cell spot prominent,
submarginal markings present, postdiscal oval mark in si, s2 and s3 entire (distinction from O. orthia);
hindwing upperside transverse band usually wide, blending anteriorly with submarginal lunules in s7;
underside pale transverse band well defined. 9 similar, larger.

Genitalia. In dorsal view £ organs narrow, like O. orticas, tegumen fragile, vincula wide, posterior
section of valve relatively stout. 9 not examined.

DISTRIBUTION. Brazil, restricted to the states of Goias and Minas Gerais, to the border of
Parana and Santa Catarina (Ebert).

DISCUSSION. This species was misidentified by Hall as a synonym of velica, and consequently
overlooked by Forbes (1945).

Ortilia velica (Hewitson) comb. n.

(Figs 67, 311, 312)
Eresia velica Hewitson, [1864]: [20], pi. 10, figs 25, 26.

c? forewing 15 mm, upperside dark brown, with or without fulvous markings arranged as in O. orthia.
9 similar, usually larger.
Genitalia. d in dorsal view, like O. orthia, harpes slightly curved, vincula appear wider.

DISTRIBUTION. S. Brazil, Uruguay, Paraguay, Argentina.
Two subspecies with similar genitalia.

Ortilia velica velica (Hewitson)

Eresia velica Hewitson, [1864]: [20], pi. [10], figs 25, 26. LECTOTYPE <J, [BRAZIL: Minas Gerais] (BMNH,

Type no. Rh. 8451; Gabriel, 1927: 121], here designated [examined].
Phyciodes velica velica (Hewitson) Rober, 1913: 438; Hall, 1929: 81; Forbes, 1945: 172, 175, 179, 190.

cJ forewing 16 mm, markings of upperside dull fulvous, arranged as in O. orthia, hindwing upperside
fulvous discal area wide.

DISTRIBUTION. S. Brazil (only in southern Minas Gerais and Sao Paulo), and Uruguay (here
often with dark upperside suffusion forming transition to following subspecies).

Ortilia velica durnfordi (Godman & Salvin)

Phyciodes durnfordi Godman & Salvin, 1878ft: 263. LECTOTYPE (J, ARGENTINA: Buenos Aires (H.

Durnford) (BMNH, Type no. Rh. 8452; Gabriel, 1927: 43), here designated [examined].
Phyciodes velica dictynna Rober, 1913: 438, pi. 89, row k [figs 5-7]. Syntypes J, 9, BRAZIL; ARGENTINA:

Palermo, Gardens of Agriculture (Seitz) (? MNHU, Berlin).
Phyciodes velica durnfordi Godman & Salvin; Hall, 1929: 82; Forbes, 1945: 179; Hayward, 1964ft: 348,

pi. 18, fig. 19.

J upperside fulvous markings reduced by dark suffusion, markings almost absent in extreme examples.

DISTRIBUTION. Brazil (Sao Paulo and southwards), Argentina, Uruguay and Paraguay.
Specimens with intermediate characters common in northern localities.

REVISION OF PHYCIODES 119

Ortilia zamora (Hall) comb, n., stat. rev.

(Figs 68, 313)

Phyciodes zamora Hall, 1917: 161. Lectotype d, 'Venezuela' [S. Brazil] (BM, Brighton), designated by Hall,

1929: 105 [examined].
Phyciodes orticas zamora Hall; Hall, 1929: 105, pi. 1, fig. 13 (rf); Forbes, 1945: 182.

cJ forewing 19 mm, outer margin slightly excavated, upperside dark brown, markings orange-yellow,
like O. orticas but with vestiges of basal markings, small spot present in cell before disco-cellular vein, a
smaller spot on the vein, postdiscal markings in darker tone of orange-brown in s3-s5; hindwing yellow
discal band as in 0. orticas, extending to inner margin, not crossed by dark veins. 9 not seen.

Genitalia. o like O. orticas, tegumen very fragile, short, posterior border of juxta gently curved, saccus
tapering.

DISTRIBUTION. Brazil: restricted to the mountains from central Espirito Santo southwards to
central Sao Paulo (Prof. Keith Brown, Pers. comm.).

Ortilia dicoma (Hewitson) comb. n.
(Figs 69, 314-316)

Eresia dicoma Hewitson, [1864]- [23], pi. [12], figs 41, 42. LECTOTYPE J, [BRAZIL] (BMNH, Type no.

Rh. 8546; Gabriel, 1927: 40), nere designated [examined].
Phyciodes dicoma (Hewitson) Rober, 1913: 445, pi. 90, row g [fig. 6] (J); Hall, 1929: 138; Forbes, 1945:

187, 190; Hay ward, 19646: 360, pi. 18, fig. 18.

cJ forewing 19 mm, elongate, upperside orange-yellow, costa and outer margin bordered black, a black
oblique postdiscal bar; hindwing with outer margin broadly black and base narrowly black. 9 similar,
upperside with fulvous tone paler.

Genitalia. <$, in dorsal view, base wide, then tapering, valve apex strongly inturned, posterior border
of juxta convex with deep central notch, harpe slender, inclined at an angle of 45°, penis with apical
section slender, lacking ostium-keel. 9 like 0. velica.

DISTRIBUTION. S. Brazil (from Dist. Federal southwards to Rio Grande do Sul, always in
forest), Paraguay, Argentina (Hayward, 1964b: 361).

Ortilia polinella (Hall) comb. n.
(Figs 70, 317)

Phyciodes polinella Hall, 1928a: 12, Hall, 1929: 139, pi. 2, figs 10 (^), 11 ($); Forbes, 1945: 188, 190.
Holotype 9, BRAZIL: San Jacintho Valley, Minas Geraes (BMNH) [examined].

cJ forewing c. 15 mm, upperside dark brown, obscurely marbled with orange; hindwing with rather broad
discal orange band (Fig. 70), and faint, very fine submarginal lunules; brownish orange below, with the
fine pattern rather indistinct. 9 forewing 16-17 mm, upperside dark brown, markings orange-yellow,
postdiscal spot in slb-s2 present, paler spot in s4; hindwing discal band orange-yellow, becoming pale
yellow and very narrow near inner margin; underside pattern rather better defined than in male.

Genitalia. <3 like 0. dicoma, in dorsal view tegumen fragile, narrow, with central sulcus, valve apex less
curved, harpe more massive. (N. B. tegumen slightly distorted in figure.)

DISTRIBUTION. Brazil: Minas Geraes, Rio de Janeiro and Espiritu Santo. A rare and local
species; there are three males and six females in the BMNH.

DISCUSSION. Hall's original description refers only to the female sex, the 'type ?' of the original
description being stated to be in the Rothschild (Tring) collection. Later, Hall (1929: 139)
claims that the holotype is a male in the National (BMNH) collection. Finally, I have seen a
male in the Booth Museum, Brighton (Hall coll.) which also appears to be a 'holotype'! The true
holotype 9, labelled 'type' by Hall and now in the BMNH, has now been clearly and
unequivocally so labelled.

120 L. G. HIGGINS

Ortilia ithra (Kirby) comb. n.
(Figs 72, 318-320, 490)

[Argynnis ianthe (Fabricius); Godart, [1824]: 818. Misidentification.]

[Acca hera (Cramer); Hiibner, [1826]: pi. 44, figs 1-4. Misidentification.]

Neptis ithra Kirby, 1871: 252. Syntypes d, 9, BRAZIL (lost). [N. ithra was named by reference to Hiibner's

misidentified figures of Papilio (as Acca) hera Cramer; these specimens are lost.]
Phyciodes ithra (Kirby) Kirby, 1900: 18; Hayward, 1964: 357, pi. 18, figs 20 (rf), 21.
[Phyciodes ianthe ianthe (Fabricius); Rober, 1913: 443, pi. 90, row d [fig. 9], row e [fig. 1]. Misidentification.]
Phyciodes ianthe atra Rober, 1913: 443, pi. 90, row b [fig. 7], row c [fig. 8]. [Hall, 1929: 104 considers

atra to be a misprint for ithra.]
Phyciodes janthe ab. nigra Giacomelli, 1928: 679. Holotype J, ARGENTINA: Rioja, Cosquin (depository

unknown).
Phyciodes ithra rufocincta Hall, 1928a: 11; Hall, 1929: 104, pi. 1, fig. 3; Forbes, 1945: 182. Holotype <J,

ARGENTINA: Cordoba Hills (BM, Brighton) [examined]. Syn. n.
Phyciodes ithra ithra (Kirby); Hall, 1929: 103; Forbes, 1945: 182, 187, 190.
Phyciodes janthe ab. schaeferi Kohler, 1945: 256, pi. 20, fig. 4. Holotype, ARGENTINA: Catamarca, Sierra

de Ambato, 2000 m ('Schaeffer Coll.').

cJ forewing 17-20 mm, elongate, upperside markings white, cell-spot small, postdiscal spot in sib, s2 s3
large; hindwing upperside discal band narrow, crossed by dark veins, small submarginal spot in s7 clearly
marked, other submarginal spots variable, well developed in Mato Grosso specimens; forewing underside
base bright orange-yellow, enclosing white, dark-ringed cell-spot. 9 similar.

Genitalia. 3 organs large, in dorsal view tegumen short, but well defined, scaphial extension very short,
saccus long, massive, apical section of valve short; penis in side view slender. 9 bursal duct partly chitinized.

DISTRIBUTION. Brazil, Argentina, Paraguay, Uruguay, Bolivia. Absent from most western
regions, but one of the commonest and most widely distributed species in the East.

NOTE. Two figures giving the dorsal view of male genitalia are included. In Fig. 318 from
Paraguay, the apical section of the valve is slightly longer and the harpe less massive than
that shown in Fig. 319 from the Mato Grosso.

TISONA gen. n.

Type-species: Phyciodes saladillensis Giacomelli. Gender: feminine.

The single known species is a small butterfly with superficial resemblance to species of Tegosa, but
differing from all other known Phyciodini in wing-markings and in genital structure.

cJ upperside of both wings orange-fulvous, markings black in a rather complicated pattern defining a
series of submarginal spots on both wings. Underside paler, yellow, markings orange, hindwing with
a prominent brown postdiscal stripe. Sexes similar.

Genitalia. j, in dorsal view, tegumen bilobed, short and very fragile, valve wide, tapering abruptly to a
short posterior process, posterior border of juxta convex, saccus entire (Ungulate) with a shallow terminal
notch; penis about 3 times the length of valve, slender, apex with 2 twisted tapering processes. 9 bursal
duct partly chitinized, bursal support well formed.

Tisona saladillensis (Giacomelli) comb. n.

(Figs 73, 74, 321-324)
Phyciodes saladillensis Giacomelli, 1911: 22.
For description of the species, the main features are covered in the generic diagnosis.

DISTRIBTION. N. Argentina, Bolivia.
Two subspecies with similar genitalia.

REVISION OF PHYCIODES 121

Tisona saladillensis saladillensis (Giacomelli)
(Figs 73, 321-324)

Phyciodes saladillensis Giacomelli, 1911: 22; Rober, 1913: 437, pi. 89, row f [figs 1, 2]; Hall, 1929: 55;
Forbes, 1945: 187, 190; Hayward, 19646: 355, pi. 18, fig. 8 (^). Holotype (sex ?), ARGENTINA: Saladillo-
Santa Cruz (MACN, Buenos Aires).

<$ forewing 17 mm, upperside bright fulvous yellow, black markings heavy and complex on both wings.
DISTRIBUTION. N. Argentina: Tucuman, Catamarca, La Rioja, Salta.

Tisona saladillensis clarior suhsp. n.

(Fig. 74)

Differs from nominate race in the following aspects: upperside paler yellow, black markings more delicate;
c? hindwing upperside with wide discal area unmarked, black postdiscal lunules vestigial or absent, but
series of submarginal lunules complete; underside paler, discal band grey-brown. 9 not available.
Genitalia examined (genitalia prep. no. 878).

Holotype <3, Bolivia: Cochabamba, Yunga del Espiritu Santo (P. Germain) (BMNH).
Paratypes. Bolivia: 3 3, same data as holotype; 2 J, 'Bolivia', ex Germain; (BMNH).

DISTRIBUTION. Bolivia: Cochabamba.

TEGOSA gen. n.

Type-species: Acraea claudina Eschscholtz. Gender: feminine.

Small butterflies, forewing length 13-18 mm, uppersides characteristically yellow-buff, wing margins black,
forewing with an oblique, black postdiscal bar, often incomplete, enclosing a sub-apical band or bar of
ground-colour. Venation and other anatomical characters show the usual tribal features, forewing outer
margin gently convex, never excavate. Sexes similar, or nearly so.

Genitalia. <3 tegumen well formed, chitinized, followed by a relatively long scaphial extension, well
defined, ending abruptly with minute spines at the posterior lateral angles, which appear in dorsal view
as spiny bosses; valves oval, tapering to a slender terminal process with a small tooth immediately above
it; harpe slender, in dorsal view usually almost straight; saccus deeply cleft or entire, or with a terminal
notch; penis straight, with a small ostium-keel. The shape of the posterior border of the juxta may be
useful. $ (examined in T.fragilis and T. anieta) with bursal duct chitinized, arising from a deep antevaginal
pit, bursal support globular.

DISTRIBUTION. The species are very widely distributed through Central and South America,
from southern Mexico to about 43°S in Argentina and Peru.

DISCUSSION. The male genitalia show a striking difference between the species of the eastern and
western regions of South America. In the eastern region, from Trinidad across Brazil and Argen-
tina, southwards to Buenos Aires, and including part of Venezuela, Paraguay and Bolivia, the
saccus is deeply cleft and the tegumen elongate, wide, tapering gradually to an abrupt, truncate
apex, with minor variation in all eastern species, and including two rare species endemic in Ecua-
dor and N. Peru. In the western region, from Mexico through Central America and along the
Andes, the saccus is entire, the tegumen and scaphial extension raquet-shaped, i.e. the extension
abruptly tapered and then expanded again to its apex, in dorsal view appearing forked with
spiny boss at the end of each lateral rib. These differences might suggest a generic distinction,
but there are some species with intermediate characters and the wing-patterns are so similar
that a close relationship is clearly present between all species within the group.

While there is not an obvious mimetic association with protected species, there is a
remarkable similarity in wing-markings between Tegosa claudina and two species of a distinct
genus, Ortilia liriope and 0. gentina. The general structure of the genitalia in Tegosa suggests

122 L. G. HIGGINS

relationship with Eresia, and it is interesting that the colour scheme and upperside wing-
markings of Eresia selene are similar to those of Tegosa, e.g. T. etia, probably indicating a
mimetic relationship.

The male genitalia of Tegosa species are among the smallest known among the Rhopalocera.
Examination is difficult as the globular shape does not present a flat surface to support the
parts in a constant position for close comparison. The organs are too small for successful
dissection, and they should be mounted entire. A dorsal view showing the shape of the tegumen
and the scaphial extension is the most helpful.

Key to species of Tegosa

Note. It has not been practicable to include T. tissoides (p. 120) and T. pastazena (p. 125) in this key.

1 Male tegumen racquet-shaped (Fig. 346)

Male tegumen not racquet-shaped (Fig. 334) 12

2 Upperside unmarked except forewing apex black, hindwing outer margin with or without

black marginal border guatemalena (p. 125)

Upperside not so marked ............. 3

3 Forewing upperside black costal bar incomplete or vestigial .... anieta luka (p. 126)
Forewing not so marked 4

4 Wing-base, costa and marginal borders broadly black, postdiscal bar isolating narrow

subapical orange band anieta anieta (p. 126)

Upperside not so marked • 5

5 Like T. anieta anieta, but black upperside markings extended, forewing pointed

anieta cluvia (p. 126)
Like T. anieta cluvia, but forewing upperside with small orange cell-spot . . . nazaria (p. 127)

6 Upperside ground-colour pale yellow flavida (p. 124)

Upperside not so marked

7 Upperside forewing with additional markings at base and in cell .... ursula (p. 124)
Forewing not so marked

8 Upperside hindwing with coloured area reduced to discal band 9

Upperside hindwing black 11

9 Hindwing upperside discal band orange nigrella f. niveonotis (p. 128)

Hindwing not so marked • 10

10 Hindwing discal band white nigrella f. niveonotis (p. 128)

Hindwing discal band obscured by general fuscous suffusion . . . nigrella f. nigrella (p. 128)

11 Forewing upperside with orange markings etia f. etia (p. 128)

Forewing upperside entirely black etia f. tissa (p. 128)

12 <3 tegumen base wide, tapering evenly to truncate apex (Fig. 334) 13

• c? tegumen base wide, tapering abruptly with angle at point of contraction (Fig. 325) . 14

13 Forewing upperside markings well developed .... orobia (p. 123), infrequens (p. 124)
Forewing upperside postdiscal markings vestigial ... . fragilis (p. 124)

14 Forewing upperside markings well developed claudina (p. 122)

Forewing upperside markings incomplete ........ ami/is (p. 123)

Tegosa claudina (Eschscholtz) comb. n.
(Figs 75, 325, 326)

Acraea claudina Eschscholtz, 1821 : 212, pi. [8], figs 18a, b. Syntype(s), [ARGENTINA] (depository unknown).

? Argynnisflavia Godart, [1824]: 818. Syntype(s), BRAZIL (? MNHN, Paris).

? [Phyciodes liriope (Cramer); Hall, 1929: 63, in part. Misidentification.]

? Phyciodes liriope claudina (Eschscholtz); Hall, 1929: 65.

? Phyciodes (Eresia) claudina (Eschscholtz); Forbes, 1945: 157-160, 188-9, in part.

? Phyciodes claudina (Eschscholtz); Hayward, 19646: 334, pi. 18, fig. 9.

Upperside fulvous yellow; 3 forewing 15-16 mm, cell crossed by black striae, post-discal black bar
complete, subapical yellow bar divided into two almost equal areas, wing apex and outer margin black;
hindwing black marginal border about 2 mm wide with vestigial submarginal lunules near anal angle,

REVISION OF PHYCIODES 123

postdiscal dots in each space almost always present; hindwing underside with reddish striae in a confused
pattern, almost always with a small, darker spot at cell-end, postdiscal spots in complete series, spot in s3
emphasised, often with a small dark suffusion. 9 similar, forewing 18 mm, upperside slightly paler.

Genitalia. <$ in dorsal view, tegumen short, scaphial extension tapered with a blunt lateral angle, spiny
bosses well defined as rather small spiny knobs, posterior border of juxta slightly prominent, saccus deeply
cleft; penis in side view slender, ostium keel present, morulae small.

DISTRIBUTION. Brazil (Parana), Argentina (especially in north-west; Chaco, Resistencia, Villa
Ana etc.), Paraguay (common), ? Peru.

NOTE. Eight males have been checked by dissection (3 <$, Paraguay, slide nos 821, 2711, 2763;
1 cJ, Peru, no. 1074; 1 cJ, Brazil, Parana, no. 863; 3 ^, Argentina, slide nos 2764, 2766(1), 2766(2)).

Tegosa similis nom. n.

(Figs 76, 327-333)

Papilio thymetus Fabricius, 1787: 30. LECTOTYPE <J, [BRAZIL: Minas Geraes] 'Indiis') (UM, Glasgow),

here designated [examined]. [Junior primary homonym of Papilio thymetus Cramer, 1777.]
[Phyciodes fragilis (Bates); Rober, 1913: 445, pi. 89, row a [fig. 9] (^). Misidentification.]
? Phyciodes liriope thymetus (Fabricius); Hall, 1929: 66.
[Phyciodes (Eresia) claudina fragilis (Bates); Forbes, 1945: 157-160. 189. ? Misidentification.]

Upperside usually pale orange-yellow; <$ forewing 16-18 mm, costa dark with striae running into cell,
postdiscal dark bar incomplete, often vestigial, then defined only by a mark on costa, apex and outer
margin dark, subapical yellow band not divided; hindwing dark marginal border prominent, including
marginal lunules, dark postdiscal dots usually present in each space; underside paler yellow, with faint
brownish markings. 9 similar, slightly larger.

Genitalia. $ tegumen short, in dorsal view scaphial extension tapered with blunt lateral angle, apex
truncate, spiny bosses conspicuous, close together, lying transversely, posterior of juxta with low central
prominence, saccus cleft often incomplete. 9 ductus chitinized, arising from a deep pit, surmounted by
globular bursal support.

DISTRIBUTION. Trinidad, Brazil, N. Argentina, Venezuela, Colombia, Guatemala, Peru, Ecuador,
Cayenne, Paraguay, Mexico.

NOTE. Identification has been confirmed by genital examination of the following specimens.
Mexico, Santos (slide no. 894); Colombia, Cauca V. (no. 2716); Venezuela (no. 856); Peru,
Huancamayo (no. 2759); Ecuador (no. 2715); Guatemala (no. 858); Brazil, Para (no. 874);
Trinidad, (no. 893); Brazil, Bahia (?, slide no. 1080); Rio de Janeiro (no. 2701).

Tegosa orobia (Hewitson) comb, n., sp. rev.

(Figs 77, 334-336)

Eresia orobia Hewitson, [1864]: [19], pi. [10], figs 23, 24. LECTOTYPE 9, [BRAZIL] (BMNH, Type no.

Rh. 8532; Gabriel, 1927: 91), here designated [examined].

[Phyciodes liriope claudina (Eschscholtz); Rober, 1913: 435, pi. 89, row a [fig. 4] (9). Misidentification.]
Phyciodes liriope orobia (Hewitson); Rober, 1913: 435, pi. 89, row b [fig. 7].
Phyciodes liriope claudina f. orobia (Hewitson); Hall, 1929: 66.
? Phyciodes liriope claudina ab. immaculata Hayward, 1935: 223. Holotype, ARGENTINA: Santiago del

Estero ('Breyer Coll.').
Phyciodes (Eresia) claudina anieta ab. orobia (Hewitson); Forbes, 1945: 160, 189.

Upperside fulvous-yellow to darker orange-fulvous; <$ forewing 15-19 mm, black markings rather extensive
with black striae at wing-base and in cell, subapical orange bar divided into two or three areas; hindwing
upperside marked with basal striae, postdiscal dots and submarginal lunules; hindwing underside with
brown markings, a dark marginal patch in s4 often extended basally.

Genitalia. $ in dorsal view, tegumen wide, scaphial extension evenly tapered to truncate apex, terminal
bosses appearing as small, round spiny knobs, posterior border of juxta gently convex, harpe slender,
saccus deeply cleft; penis slender, ostium keel small. 9 not figured.

DISTRIBUTION. Brazil (Parana etc.), Paraguay (Sapucay), Argentina (Buenos Aires).

124 L. G. HIGGINS

Tegosafragilis (Bates) comb, n., stat. rev.

(Figs 78, 337, 338)

Melitaea fragilis Bates, 1864ft: 189. LECTOTYPE <J, BRAZIL: banks of Cupari, branch of the Tapajos

(H. W. Bates) (BMNH, Type no. Rh. 8531; Gabriel, 1927: 52), here designated [examined].
[Phyciodes guatemalena (Bates); Rober, 1913: 435, pi. 89, row b [figs 1-3]. Misidentification.]
[Phyciodes liriope thymetus (Fabricius); Hall, 1929: 66. Misidentification.]
Phyciodes (Eresia) claudina fragilis (Bates); Forbes, 1945: 160, 189.

Upperside fulvous-yellow, markings black ; J forewing 15-17 mm, elongate, costa narrowly dark with black
striae running into cell, postdiscal bar incomplete, dark suffusions at wing-bases minimal; hindwing
underside markings yellow-brown, a strongly convex postdiscal line usually well developed. 9 similar,
upperside markings better defined.

Genitalia. $ in dorsal view like T. orobia, tegumen evenly tapering to truncate apex, saccus deeply
cleft; penis in side view slender.

DISTRIBUTION. Venezuela (Siapure), Brazil.

NOTE. Identification has been confirmed by genitalic examination of the following specimens :
2 (J, Venezuela, Siapure (slide nos 848, 866); 1 £, Brazil, Alta Mira (no. 2595).

Tegosa infrequens sp. n.

(Figs 79, 339, 340)

c? upperside fulvous, like T. anieta, forewing 16 mm, oblique postdiscal bar complete, enclosed fulvous
bar not divided; hindwing underside markings very faint, pale orange-brown. 9 slightly larger, markings
as in cJ.

Genitalia. J1 like T. claudina, apex of scaphial extension of tegumen slightly narrower, posterior border
of juxta with low central prominence; penis slender. 9 not dissected.

Holotype <J, Brazil: Sao Paulo, Bahuru (Hempel) (genitalia slide no. 1075) (BMNH).

Paratypes. Brazil: 2 <$, Petropolis (genit. si. nos 2764); 2 <$, same data as holotype (genit. si. no. 857);
1 c£, Minas Gerais, San Jacintho Valley (genit. si. no. 2765); 1 9, Quipapa, Pernambuco. Bolivia: 2 J,
R. Tanampaya (genit. si. no. 2714, 2759). Peru: 1 cJ, Chanchamayo (genit. si. no. 1074). (All in BMNH.)

DISTRIBUTION. Brazil, Bolivia (R. Tanampaya), Peru (Chanchamayo).

Tegosa ursula (Staudinger) comb. n.
(Figs 80, 341)

Phyciodes ursula Staudinger, 1894: 70, pi. 2, fig. 3; Rober, 1913: 435, pi. 89, row c [fig. 3] (<J); Hall, 1929:
54; Forbes, 1945: 158, 189; Hayward, 1964ft: 333, pi. 18, fig. 11 (rf). 1 £, 2 9 syntypes, BOLIVIA:
Cocapata; Bueyes (Garlepp) (MNHU, Berlin).

cJ forewing 15-16 mm, upperside fulvous with irregular reticulate markings black; hindwing black marginal
lunules and marginal border conspicuous. 9 similar.

Genitalia. $ tegumen wide in dorsal view, slightly tapered posteriorly, apex truncate, terminal bosses
represented by slightly oblique dense bars each with 5 or 6 short teeth on each side, dorsal and ventral
plates lightly chitinized, harpes sharply curved, saccus deeply cleft. 9 not examined.

DISTRIBUTION. Argentina (Tucuman, Salta, Jujuy— specimens in BMNH), Bolivia (no specimens
seen).

Tegosa flavida (Hewitson) comb. n.
(Figs 81, 342)

Eresia flavida Hewitson, [1868]: [34], pi. [19], fig. 61. LECTOTYPE J, ECUADOR (BMNH, Type no. Rh.

8527; Gabriel, 1927: 51), here designated [examined].
Phyciodes albescens Rober, 1913: 444, pi. 90, row f [fig. 4]. Holotype 9, ECUADOR: Quevedo (Buchwald)

(BMNH) [examined].

REVISION OF PHYCIODES 125

Phyciodes flavida (Hewitson) Rober, 1913: 444, Hall, 1929: 75; Forbes, 1945: 158, 189.

$ forewing 14-15 mm, upperside pale yellow, apex and outer margin dark brown (? black when
fresh); hindwing outer margin dark; no other markings. 9 similar, slightly larger.

Genitalia. Like T. claudina, $ tegumen moderately tapered in dorsal view, spinous areas of scaphial
extension small, approximately rounded, harpes slender, slightly curved, saccus pedunculate, partially cleft
(single preparation). 9 not examined.

DISTRIBUTION. Ecuador, N. Peru.

Tegosa tissoides (Hall) comb. n.
(Figs 90, 343)

Phyciodes tissoides Hall, 1928a: 11; Hall, 1929: 72, pi. 1, fig. 1 (J); Forbes, 1945: 158, 189. Holotype
cJ, ECUADOR: Angamarca, Salidero, 350 ft [110 m] (BMNH) [examined].

3 forewing 14 mm, uppersides of both wings black with brown reflections; forewing underside brown,
marginal markings vestigial if present; hindwing underside anal area and disc white shading into brown
near outer margin.

Genitalia. In dorsal view $ tegumen slightly elongate, tapered, scaphial extension with terminal bosses
not prominently spined but appearing on each side as oblique oval structures; harpe relatively stout;
saccus elongate, entire, with shallow terminal notch, preparation no. 1117 (single preparation).

DISTRIBUTION. Ecuador. BMNH collection includes 4 males with holotype, all from Angamarca,
110 m, February (Hall, 1929: 73 records one of these specimens as a female).

Tegosa pastazena (Bates) comb, n., stat. n.

(Figs 82, 344, 345)

Melitaea liriope var. pastazena Bates, 18646: 189 (footnote). LECTOTYPE d, ECUADOR: Canelos, banks of

the Pastaza (Spruce) (BMNH, Type no. Rh. 8530; Gabriel, 1927: 94), here designated [examined].
Phyciodes liriope pastazena (Bates); Hall, 1929: 67.
Phyciodes (Eresia) claudina anieta f. pastazena (Bates); Forbes, 1945: 160.

$ forewing 16-17 mm, like T. anieta but slightly larger; upperside tawny orange, costa black to origin
of oblique costal bar, enclosed orange subapical band not divided, outer margin broadly black ; hindwing
dark border along outer margin narrow but clearly defined; hindwing underside postdiscal dark spots
well developed in slb-s6. 9 similar, slightly larger.

Genitalia. J tegumen rather narrow, scaphial extension narrow, elongate, lateral walls almost parallel,
each posterior spiny angle scarcely defined as a boss, saccus single with terminal notch.

DISTRIBUTION. Bolivia, Peru, Ecuador.

DISCUSSION. This species appears to be very local but widely distributed in the south-western
area of generic distribution. In four specimens examined the characters of the genitalia are
constant and the external markings also recognisable, but identification must be confirmed by
examination of the genitalia.

Tegosa guatemalena (Bates) comb, n., stat. n.

(Figs 83, 346-348)

Melitaea fragilis var. guatemalena Bates, 1864/x 192-193. LECTOTYPE <J, GUATEMALA: central valleys

(BMNH, Type no. Rh. 8529; Gabriel, 1927: 56), here designated [examined].
[Phyciodes fragilis (Bates); Godman & Salvin, 1882: 198, pi. 21, fig. 23 (^). Misidentification.]
[? Phyciodes liriope pastazena (Bates); Rober, 1913: 435, pi. 89, row b, [fig. 4] (^). Misidentification.]
Phyciodes liriope guatemalena (Bates); Hall, 1929: 67.
Phyciodes (Eresia) claudina guatemalena (Bates); Forbes, 1945: 160, 189.

c? forewing 16-17 mm, upperside orange-yellow, almost unmarked, apex slightly shaded black; hindwing
with narrow dark marginal border in some specimens; underside with confused pale reddish striae. 9
slightly larger, forewing upperside dark markings more complete.

126 L. G. HIGGINS

Genitalia. £ tegumen wide, tapered rather abruptly to narrow scaphial extension, apex forked and
expanded with spiny terminal bosses on each side, posterior border of juxta gently curved, saccus conical,
entire or rarely apex slightly notched. 9 not examined.

DISTRIBUTION. Guatemala, Mexico, Ecuador, Belize.

Tegosa anieta (Hewitson) comb, n., stat. rev.

(Figs 84-88, 349-356)
Eresia anieta Hewitson, [1864]: [23].

cJ forewing 14-15 mm, upperside orange-fulvous, costa, base and wing-borders black, oblique postdiscal
bar variable, typically well developed but incomplete or even vestigial in many areas, subapical orange
band, when defined, rather small, narrow, rarely partly divided; hindwing margin black, otherwise
unmarked. 9 upperside usually slightly paler, often with small submarginal yellowish spots in s5 and s6.
Genitalia. <$ tegumen wide, abruptly tapered to a narrow neck and expanded again at apex, saccus single,
posterior border of juxta almost straight, penis straight, slender, morula small. 9 not examined.

Four subspecies.

Tegosa anieta anieta (Hewitson)

(Figs 84, 85, 349, 350, 351, 355, 356)

Eresia anieta Hewitson, [1864]: [23], pi. [12], figs 43, 44. LECTOTYPE rf, VENEZUELA: Caraccas (BMNH,

Type no. Rh. 8528; Gabriel, 1927: 14), here designated [examined].
Phyciodes anieta (Hewitson) Godman & Salvin, 1882: pi. 21, fig. 20 (^); Rober, 1913: 435, pi. 89, row a

[fig. 6], row b [fig. 5, 6].

Phyciodes liriope anieta (Hewitson); Hall, 1929: 68.
Phyciodes (Eresia) claudina anieta (Hewitson); Forbes, 1945: 159, 189.

c? upperside with all black markings well defined; forewing subapical orange band not divided; hindwing
postdiscal black dots and vestiges of submarginal lunules often present.
Genitalia. As described above.

DISTRIBUTION. Panama, Costa Rica, Venezuela.

Tegosa anieta cluvia (Godman & Salvin)
(Figs 86, 352, 353)

Phyciodes cluvia Godman & Salvin, 1882: 198, pi. 21, figs 21, 22; Rober, 1913: 435; Hall, 1929: 70.

LECTOTYPE <J, GUATEMALA: Cerro Zunil (Champion) (BMNH, Type no. Rh. 8524; Gabriel, 1927: 32),

here designated [examined].

Phyciodes liriope f. lirina Rober, 1913: 435. Syntype(s), BOLIVIA (? MNHU, Berlin). Syn. n.
Phyciodes liriope lirina Rober; Hall, 1929: 68; Forbes, 1945: 188, 190.
Phyciodes (Eresia) claudina cluvia Godman & Salvin; Forbes, 1945: 159, 189.

Like T. anieta anieta but upperside with all black markings extended; forewing subapical orange band
reduced, often partly divided, discal fulvous area reduced; hindwing marginal black border wide.
Genitalia. As described above.

DISTRIBUTION. Guatemala, ? Bolivia.

Tegosa anieta luka suhsp. n.

(Figs 87, 354)
[Phyciodes liriope flavia (Godart); Rober, 1913: 435, pi. 89, row a [fig. 6] (<$). Misidentification.]

cJ upperside fulvous-yellow; forewing costa usually dark to postdiscal bar, the latter, although often
extending only halfway across the wing and then tapering to a point, is variable and sometimes vestigial;
hindwing underside often with darker marginal shade in s3 and s4. 9 slightly larger, upperside dark
markings often more extensive.

REVISION OF PHYCIODES 127

Genitalia. Compared with nominate anieta, the scaphial expansion may be slightly wider.

Holotype J, Mexico: Tamazunchale, San Luis Potose, i.1963 (£. C. Welling) (genitalia slide no. 2596)
(BMNH).

Paratypes. Colombia: 5 <$ (genitalia slide nos. 1223, 2708, 839, 1224, 2709. Mexico: 1 ?, same data as
holotype. Ecuador: 2 <$ (genitalia slide nos. 1068, 886). (All in BMNH.)

DISTRIBUTION. Mexico, Panama, Ecuador, Honduras, Belize, Costa Rica, Colombia, Peru. The
most widely distributed form of Tegosa and often common.

Tegosa anieta serpia subsp. n.

(Fig. 88)

c? forewing 15-16 mm, apex rounded, hindwing relatively small; upperside pale orange-fulvous, black
markings not heavy; forewing oblique black postdiscal bar short; hindwing black margin well defined,
underside postdiscal ocellar spots well marked in sib, sic, and in s2, bordered proximally by darker
suffusion. $ not identified.
Genitalia. $ like T. anieta anieta.

Holotype J, Peru: Rio Chuchurras, R. Palcazu, 320 m ( W. Hoffmanns) (genitalia slide no. 1070) (BMNH).
Paratype. Bolivia: 1 cJ, Cochabamba (Germain) (genitalia slide no. 2761) (BMNH).

DISTRIBUTION. Peru (Tarapoto), Bolivia (Cochabamba), W. Columbia?

DISCUSSION. T. anieta, in its various forms, is very widely distributed in the western mountains,
especially in the form or subspecies anieta luka. The relationship of this phenotype remains
rather uncertain, with the small difference in genitalia present in most specimens, but wing
markings are variable and the transformation series to typical anieta anieta appears to be
complete. T. anieta serpia, which shows quite striking features in wing-shape and markings,
is perhaps specifically distinct, but with only 7 male specimens I think it is better included with
anieta in the absence of more information about distribution.

Tegosa nazaria (Felder & Felder) comb, n., stat. rev.

(Figs 89, 357, 358)

Eresia nazaria Felder & Felder, 1867: 394. LECTOTYPE <J, COLOMBIA: Bogota (Lindig) (BMNH),

here designated [examined].

Phyciodes mazaria; Kirby, 1871: 172. [Misspelling.]

Phyciodes nazaria (Felder & Felder); Rober, 1913: 439; Hall, 1929: 69, pi. 1, fig. 12 (<?).
Phyciodes aquila Hall, 1917: 162. 7 <J syntypes, COLOMBIA: El Baldio, 5400 ft [1750 m] (BM, Brighton)

[examined].
Phyciodes (Eresia) claudina nazaria (Felder & Felder); Forbes, 1945: 159, 189.

c? forewing 16 mm, upperside base broadly black, extending across cell, a small orange cell-spot present,
black marginal borders wide, orange discal area extending from inner margin to v3, subapical orange band
composed of 3 small spots; hindwing base and outer margin broadly black, vestigial submarginal lunules
present. 9 similar.

Genitalia. d in dorsal view like T. anieta, scaphial extension wider, terminal spiny bosses small, saccus
single with small apical notch.

DISTRIBUTION. Colombia.

Tegosa etia (Hewitson) comb, n., stat rev.

(Figs 91, 92, 359, 360)

Eresia etia Hewitson, [1868]: [33], pi. [19], figs 56, 57. LECTOTYPE j, [BOLIVIA] ('Ecuador') (BMNH,

Type no. Rh. 8526; Gabriel, 1927: 47), here designated [examined].
Eresia tissa Hewitson, 1869a: 27 [index]. LECTOTYPE J, ECUADOR: Mapoto (Buckley) (BMNH, Type

no. Rh. 8517; Gabriel, 1927: 119), here designated [examined]. Syn. n.

128 L. G. HIGGINS

Phyciodes etia (Hewitson) Rober, 1913: 445, pi. 90, row g [fig. 1].

Phyciodes etia selenoides Hall, 1928a: 11; Hall, 1929: 71, pi. 1, fig. 4 (^). Holotype rf, PERU: Cachapoyas

(de Mathan) (BMNH) [examined]. Syn. n.
Phyciodes etia etia (Hewitson); Hall, 1929: 70.
Phyciodes etia tissa (Hewitson); Hall, 1929: 72.
Phyciodes (Eresia) claudina etia (Hewitson); Forbes, 1945: 159, 189.
Phyciodes (Eresia) claudina tissa (Hewitson); Forbes, 1945: 189.
Phyciodes (Eresia) claudina selenoides Hall; Forbes, 1945: 159, 189.

<3 forewing 14-15 mm, upperside with or without fulvous discal and subapical markings; underside fulvous
markings replaced by paler yellow, forewing margins and all hindwing brown. 9 similar.
The wing markings of this small species are very variable, with 3 named forms.

1. Fulvous markings present on upperside of forewing only, f. etia.

2. Markings present on hindwing, reduced on forewing, f. selenoides.

3. Markings present on uppersides of both wings, but partly obscured by dusky suffusion, f. tissa.

All forms occur, with intermediates, throughout the range.

Genitalia. <$ like T. nazaria, scaphial extension of tegumen shorter and wider, saccus short with apical
notch.

DISTRIBUTION. In the mountains of Ecuador, Bolivia and Peru.

Tegosa nigrella (Bates) comb, n., stat. rev.
(Figs 93, 94, 361-363)

Melitaea nigrella Bates, 1866: 133. LECTOTYPE <J, GUATEMALA: central valleys (BMNH, Type no. Rh.

8521; Gabriel, 1927: 86), here designated [examined].
Phyciodes niveonotis Butler & Druce, 1872: 100; Godman & Salvin, 1882: 200; Rober, 1913: 441, pi. 90,

row a [fig. 7] (J). Holotype '?' [J], COSTA RICA: Cartago (van Patten) (BMNH, Type no. Rh. 8518;

Gabriel, 1927: 87) [examined]. Syn. n.
Phyciodes lutescens Godman & Salvin, 1882: 199, pi. 21, fig. 26; Rober, 1913: 444. LECTOTYPE <J,

GUATEMALA: Purula (Champion) (BMNH, Type no. Rh. 8519; Gabriel, 1927: 76), here designated

[examined]. Syn. n.
Phyciodes nigrella (Bates) Godman & Salvin, 1882: 199, pi. 21, fig. 24 (^); Rober, 1913: 444, pi. 88, row i

[fig. 8].

? Phyciodes drusinilla Rober, 1913: 441, pi. 90, row a [fig. 8]. Syntype(s), ARGENTINA (depository unknown).
Phyciodes nigrella nigrella (Bates); Hall, 1929: 73.
Phyciodes nigrella f. lutescens Godman & Salvin; Hall, 1929: 74.
Phyciodes nigrella niveonotis Butler & Druce; Hall, 1929: 74.
Phyciodes (Eresia) claudina nigrella (Bates); Forbes, 1945: 158-159, 189.
Phyciodes (Eresia) claudina niveonotis Butler & Druce; Forbes, 1945: 159, 189.

c? forewing 14 mm, upperside black, markings white or yellow, large discal spot and small subapical
spots; hindwing with coloured discal band on black ground. $ similar.
The wing markings vary in colour, with three named forms.

1. Markings more or less obscured by dusky suffusion, f. nigrella.

2. Upperside markings white, f. niveonotis.

3. Upperside markings yellow, f. lutescens.

Genitalia. Like T. anieta, small, with similar tegumen and scaphial extension raquet-shaped, posterior
border of juxta gently convex, harpe almost straight. $ not examined.

DISTRIBUTION. Guatemala, Costa Rica (f. niveonotis only).

DISCUSSION. All forms of T. nigrella are rare; their distribution and relationships are not well
understood. I have not seen any specimen which agrees with the figure of Phyciodes drusinilla
Rober (1913: 441, pi. 90, row a [fig. 8]), described from Argentina, and included by Hall
(1929: 73) as a possible synonym of nigrella.

REVISION OF PHYCIODES 129

ERESIA Boisduval

Eresia Boisduval, [1836]: pi. 11 [= pi. 7B], fig. 8. Type-species: Nereis eunice Hiibner, by monotypy.
Gender, feminine.

Butterflies of moderate size, forewings elongate, upperside markings very varied. In females the forewings
are often more ample, apices more rounded, sometimes with striking sexual dimorphism. Astonishing
mimicry is a prominent feature, with the genera Eueides Hiibner, Ceratinia Hiibner, and Mechanitis
Fabricius the most frequent models, but a precise specific model is often difficult to establish, especially
in western Brazil, N. Peru, Guatemala and Colombia, where many species of various families appear to
be unstable. The genus is extensive and includes the largest species of the tribe.

Genitalia. In males the outstanding feature is the long scaphial extension which overhangs the valves
like a roof (Fig. 383). Slight dorsal constrictions in the lateral margins represent the posterior limit of the
9th tergum (tegumen) in many species, beyond which the anal compartment is roofed by a curtain of
delicate fibrous tissue, while the strongly chitinised lateral margins are continued to terminate on each side
in a spiny area, forming a prominent boss when viewed from above. The oval valve tapers to a single
terminal tooth, usually preceded by one or more smaller teeth upon the costal border. The medial border
proximal to the apex usually bears bristles or fine, short spines, not visible in the small figures reproduced
on the plates. In all species the saccus is single, without any cleft or terminal notch; penis straight,
sometimes massive, with a small ostium-keel. In dorsal view the female genitalia are sculptured to form a
deep pit, from which arises a lightly chitinised cone representing the ductus bursae, surmounted by a
cup-shaped bursal support. The post-vaginal scutum is not well developed.

LARVAL FOOD PLANT. Fittonia (Acanthaceae) is recorded for E. eunice by Hayward (1966).

DISTRIBUTION. The genus is widely distributed through Central and South America, from
Mexico in the north, across the tropical regions to northern Argentina (E. lansdorfi), Bolivia and
Peru. Eresia eunice, with its different modifications, is the most widely distributed species.
In the eastern districts of Brazil, and in the Guyanas, it flies with six other species, E. perna,
E. lansdorfi, E. erysice (excessively rare), E. nauplius, E. plagiata and E. clara. The remaining
species, numbering about 30, are all restricted to Central America and to the countries of
western South America, with a marked concentration of species in Colombia, Costa Rica,
Venezuela and Peru.

DISCUSSION. Throughout the genus polymorphism is often confusing, and associating the sexes
correctly may be difficult. In about half the known species the male genitalia do not show specific
characters in a convincing manner and therefore are useless for specific definition. In several of
the most variable species the status of certain phenotypes is uncertain; these are recorded here as
subspecies in all doubtful cases. Specific characters of the male genitalia are probably constant,
except in the case of E. nauplius, in which confusing variation has been observed.

Key to species of Eresia (males).

Note. It has not been practicable to include E. coela (p. 135), E. mimas (p. 138), E. phaedima (p. 143),
E. melaina (p. 139), E. anomala (p. 141) and E. etesiae (p. 146) in this key.
The species E. datis keys out twice.

1 Hindwing upperside black with transverse band of white, yellow or fulvous

Hindwing not so marked ......... . . 12

2 Band white or pale cream

Band yellow or fulvous

3 Veins crossing white band firmly lined with black ... . letitia (p. 133)
Veins crossing white band not prominent ... 4

4 Hindwing underside lacks brown basal stripe in s7 . . clara (p. 131)
Hindwing underside brown basal stripe in s7 present

5 Forewing underside cell-base yellow-brown nauplius nauplius (p. 132)

Forewing underside cell-base white or pale yellow . 6

6 Hindwing upperside transverse white band 3 mm broad, posterior border concave

plagiata (p. 133)
Hindwing upperside transverse white band 4 mm broad, posterior border straight

nauplius extensa (p. 132)

130 L. G. HIGGINS

7 Hindwing upperside transverse band yellow 8

Hindwing upperside transverse band fulvous 10

8 Forewing extremely narrow perna (p. 150)

Forewing of usual shape ' 9

9 Forewing upperside postdiscal area shaded brown lansdorfi (p. 134)

Forewing upperside black, 3 yellow discal spots in oblique row . . . polina (p. 136)

10 Forewing upperside with wide longitudinal fulvous band from base to postdiscal area

sestia (p. 134)
Forewing not so marked 11

11 Hindwing upperside fulvous band tapered from costa to inner margin . carme (p. 135)
Hindwing upperside slightly or not at all tapered oblita (p. 135)

12 Hindwing upperside disc yellow, crossed by black veins cissia (p. 137)

Hindwing not so marked 13

13 Forewing upperside black, with or without small fulvous basal suffusion .... 14
Forewing not so marked 19

14 Forewing with broad yellow oblique discal band 15

Forewing not so marked 16

15 Forewing underside with pale submarginal spots margaretha (p. 144)

Forewing underside without submarginal spots datis (p. 143)

16 Forewing upperside with many white or yellowish spots, apex truncate .... 17
Forewing upperside markings similar, apex rounded 18

17 Forewing underside spots well defined sticta (p. 140)

Forewing underside spots replaced by stripes nigripennis (p. 141)

18 Hindwing underside space 8 completely white poecilina (p. 139)

Hindwing underside pale basal mark very small (Note : markings variable, fulvous basal area

of forewing extensive in some specimens.) ithomioides (p. 140)

19 Forewing upperside fulvous, marked with 2 or 3 black oblique stripes .... 20
Forewing not so marked 21

20 Forewing proximal black stripe arises very near wing-base alsina (p. 137)

Forewing proximal black stripe, sometimes incomplete, arises more distally . .eutropia (p. 137)

21 Forewing fulvous, apex broadly black with prominent white mark in s4, s5 and s6

emerantia (p. 142)
Forewing not so marked 22

22 Wings partly translucent 23

Wings not so marked 24

23 Forewing basal and postdiscal black bands present datis (p. 143)

Forewing lacking basal and postdiscal markings moesta (p. 142)

24 Hindwing underside grey-brown, veins radiating, heavily scaled black . . actinote (p. 150)
Hindwing not so marked 25

25 Hindwing upperside fulvous, with black transverse band 26

Hindwing not so marked 36

26 Forewing exceedingly narrow aveyrona (p. 149)

Forewing of usual shape 27

27 Forewing apex truncate phillyra (p. 148)

Forewing of usual shape, apex rounded 28

28 Forewing upperside fulvous basal area restricted to s2 casiphia (p. 147)

Forewing upperside fulvous basal area larger, including cell 29

29 Forewing upperside apex black 30

Forewing upperside apex not so marked 34

30 Forewing apical black area unmarked 31

Forewing apical area pale markings more or less developed 32

31 Forewing upperside black discal band reduced to 2 black spots . . pelonia f. pelonia (p. 148)
Forewing upperside black discal band wide, prominent .... eunice esora (p. 145)

32 Forewing upperside black discal band short, restricted to cell .... erysice (p. 146)
Forewing not so marked 33

33 Forewing upperside with short black mark in cell, another mark at cell-end, yellow post-discal

area extensive quintilla (p. 138)

Forewing upperside postdiscal black band complete, reaching outer margin .... 34

34 Forewing upperside apical markings present, forming short, yellow band . mechanitis (p. 147)

REVISION OF PHYCIODES 131

Forewing apical markings absent, rarely vestigial ......... 35

35 Forewing upperside black markings well defined, oblique costal bar complete to outer margin

eunice eunice (p. 145)
Forewing upperside black markings reduced, costal black bar incomplete eunice olivencia (p. 145)

36 Upperside black, with a blue discal band on forewing ...... levina (p. 150)

Upperside black, markings fulvous 37

37 $ forewing 27 mm; hindwing with wide fulvous area along distal marginpelonia f. ithomiola (p. 148)
- c? forewing 16 mm; fulvous discal mark on hindwing not extending to outer margin (upperside

marking resembles Telenassa teletusa) selene (p. 151)

Eresia clara Bates sp. rev.
(Figs 177, 183, 184, 364-367)

[Papilio nauplius Linnaeus; Clerck, [1764]: pi. 46 [fig. 2]; Cramer, [1780]: 55, pi. 316, figs D-G.

Misidentifications.]
[Eresia clio (Linnaeus); Doubleday, 1850: 184; Aurivillius, 1882: 172; Rober, 1913: pi. 92, row h [figs 1, 2].

Misidentifications.]
Eresia clara Bates, 18646: 192. LECTOTYPE rf, BRAZIL: Belem ('Para') (H. W. Bates) (BMNH,

Type no. Rh. 8536; Gabriel, 1927: 32), here designated [examined].

[Phyciodes clio (Linnaeus); Kirby, 1871: 177; Rober, 1913: 446; Hall, 1929: 144. Misidentifications.]
Phyciodes clio estebana Hall, 1929: 146. Holotypec?, VENEZUELA: San Esteban Valley, nr Puerto Cabello

(ex Hahnel) (BMNH) [examined]. Syn. n.
Phyciodes clio reducta Hall, 1929: 146. Holotype J, ECUADOR: La Chima, Rio de las Juntas, pr.

Bahahoyo, vi-vii. 1893 (de Mathan) (BMNH) [examined]. Syn. n.
[Phyciodes (Eresia) clio (Linnaeus); Forbes, 1945: 165-166, 189. Misidentification.]

cJ forewing 17-19 mm, upperside black, forewing with 4 well-defined white or yellowish spots; on
hindwing upperside a white discal band of variable width; hindwing underside base narrowly white with
broader white stripe below, above the white discal band. 9 similar, usually slightly larger.

Genitalia.J, in dorsal view, tegumen wide, lateral shoulders well developed, posterior border of juxta
gently convex, terminal bosses of scaphial extension small, spines not prominent; penis with large
ostium-keel.

DISTRIBUTION. From S. Mexico through central and western South America to Peru and
Bolivia, and in all the Amazon region, including the Guyanas, and southwards to western
Mato Grosso.

DISCUSSION. External markings may be white or slightly yellowish; the extent of markings,
especially that of the white transverse bar on the hindwing, may be slightly enlarged (estebana),
or reduced in small specimens (reducta); the hindwing outer margin may be straight or
gently convex. These variants appear to represent clinal phases with only minor taxonomic
significance.

NOMENCLATURAL NOTE. The name Papilio clio Linnaeus, used for this species by many authors,
is based upon the figure by Merian (1705: pi. 35), and placed by Linnaeus in his group
'Heliconii', with Papilio apollo, etc. Merian's figure shows an Ithomiine, and was so recorded
by Butler (1870: 126), possibly Leucothyris aegle (Fabricius) as later figured in 'Seitz' (pi. 38f).
Confusion was introduced by Doubleday (1850: 184) when 'Papilio clio L.(?)' was included
in the list of species of Eresia, and again by Kirby (1871: 177) when he transferred P. clio L.
to Phyciodes with the usual references, but now without the interrogation mark. Perhaps this
incorrect identification led Aurivillius (1882) to introduce the name clio for the common
Phyciodes species, as later accepted by Rober in 'Seitz', and by Hall. I imagine that Aurivillius
never saw Merian's original figure, nor the paper by Bates (1864b), in which action was taken
to give valid names to the two species figured as nauplius by Clerck and by Cramer.

Papilio clio was recorded by Linnaeus as a species represented in the collection of Queen
Ludovica Ulrica, now preserved at Uppsala. I understand that there is no species in the collection
today under this name, but there is a specimen of Eresia clara Bates under the name
P. nauplia which corresponds in every way with the lower figure in Clerck ([1764]: pi. 46).

132 L. G. HIGGINS

Eresia nauplius (Linnaeus) comb. rev.
(Figs 95, 96, 368-374)

Papilio nauplius Linnaeus, 1758: 488.
Eresia nauplia (Linnaeus) Bates, 1864b: 192.

c? (brewing 20-22 mm, narrow, outer margin slightly concave; upperside grey-black with chalk-white
markings like E. clio and additional small white submarginal mark in s3, s4; hindwing rather elongate,
outer margin scalloped and fringes slightly chequered pale and dark ; upperside with a broad white discal
band extending from inner margin into s7 ; forewing underside grey with orange-brown and large white
markings as on upperside; hindwing underside costa broadly white, divided by black-lined v8, followed by
narrow brown and black stripes before the white discal band, then a white submarginal band divided
into discrete lunules.

Genitalia. cJ, in dorsal view, variable, tegumen generally narrow, shoulders little developed (Fig. 369),
lateral borders of scaphial extension straight or slightly divergent, terminal bosses well developed, rarely
very large (Fig. 368) or otherwise modified (Fig. 370), posterior border of juxta prominent, penis slender,
ostium-keel well developed. 9 bursal duct chitinised, rather long, conical, bursal support cup-shaped,
scutum well defined.

DISTRIBUTION. Brazil, the Guyanas.
Two subspecies, with similar genitalia.

Eresia nauplius nauplius (Linnaeus)
(Figs 95, 368-374)

Papilio nauplius Linnaeus, 1758: 488; Clerck, [1764]: pi. 46, fig. [1]. Syntype(s), [?SURINAM] ('In Indiis')

(University of Uppsala).

Papilio nauplia Linnaeus; Linnaeus, 1767: 783; Aurivillius, 1882: 105.
[Phyciodes clara (Bates); Rober, 1913: 446, pi. 92, row h [fig. 5]. Misidentification.]
Phyciodes nauplia nauplia (Linnaeus); Hall, 1929: 147 [note: Hall's synonymy is in part erroneous];

Forbes, 1945: 165-6, 189.

Forewing underside cell-base orange-brown to round white cell-spot, with orange mark beyond ; width of
hindwing upperside white discal band about 3 mm.

DISTRIBUTION. Probably restricted to the Guyana shield, and Amazon region of Brazil.

Eresia nauplius extensa (Hall)
(Fig. 96)

Phyciodes nauplia extensa Hall, 1929: 148. Holotype ^, BRAZIL: Chapada, Mato Grosso (H. H. Smith)

(BMNH) [examined].
Phyciodes (Eresia) nauplia extensa Hall; Forbes, 1945: 166.

Like E. nauplius nauplius, but hindwing upperside white discal band about 4 mm wide; forewing
underside cell-base white, sometimes fusing with round cell-spot; underside of both wings with white
markings wider, more extensive.

DISTRIBUTION. Brazil: Mato Grosso (5 specimens), lower R. Madeira (1 specimen).

DISCUSSION. The six specimens in the BMNH show constant characters, and others in the
Hall Coll. (BM, Brighton) are similar. Accepted by Prof. Keith Brown (pers. comm.) as a
geographical subspecies, distributed from central Mato Grosso westwards.

E. nauplius has proved to be, taxonomically, the most difficult species of the genus. The
external features appear to be well defined, but the male genitalia show marked variation in the
shape of the dorsal structures and of the posterior border of the juxta, to an extent not seen in
any other species. It may be that two or more species are present in the complex. Four
figures of male genitalia are included to show the extent of variation found in a series of
14 preparations.

REVISION OF PHYCIODES 133

Eresia plagiata (Rober) comb, n., stat. n.

(Figs 97, 375-379)

Phyciodes nauplia plagiata Rober, 1913: 446; Hall, 1929: 148; Forbes, 1945: 166. 9 syntype(s), PERU:

upper Madre de Dios, 1500-3000 ft [500-1000 m] (A. H. Fassl) (depository unknown).
[Eresia nauplia (Linnaeus); Rober, 1913: pi. 92, row h [fig. 5]. Misidentification.]

cJ forewing 23-24 mm, like E. nauplius; upperside white markings slightly larger; hindwing outer margin
not scalloped, fringes brown, white discal band about 3.5 mm wide, rarely invades s7; forewing underside
with cell white, fusing with white cell-spot, brown mark beyond vestigial, if present; hindwing underside
markings as in E. nauplius, brown postdiscal line and white submarginal line both continuous, not
lunulate as in E. nauplius. 9 slightly larger, underside marginal markings better defined.

Genitalia. $ in dorsal view, tegumen wide, lateral walls of scaphial extension divergent, strongly
chitinized, terminal bosses large, posterior border of juxta curved; penis slender; only minor variation has
been seen. 9 like E. nauplius, bursal duct perhaps shorter and wider (single preparation).

DISTRIBUTION. Brazil (including Amazon regions, Iquitos, Teffe etc.), Colombia (Florida),
Peru (R. Chuchurras, La Merced), Ecuador. Not seen from the Guyanas, or lower Amazon.

Eresia letitia Hewitson comb. rev.
(Figs 98, 99, 380-384)

Eresia letitia Hewitson, 1869a: 24 [index].
Phyciodes letitia (Hewitson); Rober, 1913: 448.

c? forewing 24 mm, outer margin gently convex, upperside black, markings white or pale grey; hindwing
transverse discal band white or yellowish, submarginal striae flat. 9 ground-colour yellow-brown or
white (form leucophaea}, forewing 26 mm or more, markings as in d, often enlarged.
Genitalia. Variable, see below.

DISTRIBUTION. Colombia, Ecuador, Peru.
Two subspecies with similar genitalia.

Eresia letitia letitia Hewitson
(Figs 98, 382, 384)

Eresia letitia Hewitson, 1869a: 24 [index]; Hewitson, [1870]: [38], pi. [21], figs 70, 75, 76; Rober, 1913:
pi. 91, row a [fig. 3]. LECTOTYPE 9, ECUADOR: St. Inez, Buckley (BMNH, Type no. Rh. 8538;
Gabriel, 1927: 73), here designated [examined].

? Eresia leucophaea Weymer, 1890: 54. 9 syntypes, ECUADOR (? MNHU, Berlin).

Phyciodes letitia letitia (Hewitson); Hall, 1929: 149; Forbes, 1945: 165.

cJ upperside markings well defined, hindwing transverse white band 4 mm wide in specimens from
Colombia, slightly narrower in those from Ecuador.

Genitalia. d tegumen wide, lateral shoulders prominent in three examples from Colombia, scaphial
extension wide in two specimens but narrower in a third specimen.

DISTRIBUTION. Colombia, Ecuador (hindwing upperside white discal band often narrower).

Eresia letitia ocellata (Rober) stat. rev.
(Figs 99, 380, 381, 383)

Phyciodes letitia ocellata Rober, 1913: 448. Syntype(s), PERU: Chanchamayo (depository unknown).
Eresia neptoides Rosenberg & Talbot, 1914: 675. Holotype <$, PERU: El Porvenir, 900 m, April 1908

(BMNH) [examined].
Eresia letitia nigra Rosenberg & Talbot, 1914: 676. Holotype c?, PERU: Huancabamba, Cerro del Pasco

(BMNH) [examined]. Syn. n.

[Phyciodes leucophaea (Weymer); Rober, 1913: 448. Misidentification.]
[Eresia leucophaea Weymer; Rober, 1913: pi. 91, row a [fig. 4]. Misidentification.]

134 L. G. HIGGINS

Phyciodes letitia nigra (Rosenberg & Talbot); Hall, 1929: 150; Forbes, 1945: 165.

Phyciodes letitia nigra f. leucophaeoides Hall, 1929: 151. Holotype 9, PERU : Chanchamayo (BM, Brighton).

Phyciodes ocellata Rober; Hall, 1929: 151; Forbes, 1945: 165; 189.

<$ upperside darker, pale markings suffused with dark scales; hindwing upperside pale transverse band
narrow, 2.5 mm, widest at centre, white or yellowish.

Genitalia. Of five preparations, the lateral shoulders of the tegumen are prominent in three, but
in two examples they do not project laterally; scaphial extensions wide, or narrower and longer
(Fig. 381).

DISTRIBUTION. Peru, in which it appears to be rather widely distributed. The nominate form
has not been seen from this region.

DISCUSSION. Specimens with discal band yellowish on upperside of the hindwing are typical
of ocellata, later named neptoides by Rosenberg & Talbot. In the neptoides type-series, seven
specimens all from El Porvenir in Peru, the outer margin of the hindwing is straight.

Eresia lansdorfi (Godart) comb. rev.
(Figs 100, 385)

Heliconia lansdorfi Godart, 1819: 209. Syntype(s), BRAZIL (depository unknown).

Melinaea langsdorfii (Godart) Hiibner, [1821-1822]: pi. [68], figs 389, 390; 1823: 31.

Heliconia langsdorfii Godart; Godart, [1824]: 806.

Phyciodes lansdorfi (Godart) Staudinger, 1885: 92, pi. 36 (^); Rober, 1913: 448; Hall, 1929: 154; Forbes,

1945: 162, 164, 189; Hayward, 1964ft: 338, pi. 18, fig. 12.
Eresia lansdorfi (Godart) Rober, 1913: pi. 92, row a [figs 1, 2].
Phyciodes lansdorfi t.jacinthica Rober, 1913: 448, pi. 90, row k [fig. 6]. LECTOTYPE 9, BRAZIL: San Jacintho,

Theophilo Ottoni, Minas Geraes (F. Birch) (BMNH), here designated [examined].
Phyciodes lansdorfi f. veternosa Ferreira d'Almeida, 1922: 181. Holotype 9, BRAZIL: Jacarepagua (Tanque)

(depository uncertain).
Phyciodes lansdorfi 9 f. sulphurata Zikan, 1937: 386. Holotype 9, BRAZIL: 'Siidabhang des Itatiaya,

700 m., 27th December' (depository uncertain).
Phyciodes lansdorfi (Latreille); Hayward, 1952: 290.

J forewing upperside black, obscure basal stripe yellowish, post-discal area light red-brown, including
costal bar; hindwing upperside black, transverse band pale yellow. 9 variable, usually like 3, but in
some areas (form jacinthica) forewing upperside basal stripe well defined, light brown, and hindwing
upperside submarginal lunules present in slb-s4. More rarely upperside forewing red-brown markings
more or less replaced by yellow (? f. sulphurata).

Genitalia. ^ distinctive, tegumen wide, shoulders not defined, lateral walls of scaphial membrane
chitinised, diverging, terminal bosses small, posterior border of wide juxta almost straight, harpes short,
tapering. 9 not examined.

DISTRIBUTION. Brazil (from Espirito Santo and Bahia southwards through Parana to Rio
Grande do Sul), N. and C. Argentina, Paraguay, Uruguay (and Peru?).

Eresia sestia Hewitson comb. rev.
(Figs 101, 102, 386, 387)

Eresia sestia Hewitson, 1869a: 26 [index]; Hewitson, [1870]: [37], pi. [21], figs 68, 69, 74; Fassl, 1912: 122.

LECTOTYPE ^, ECUADOR: Jorge (Buckley) (BMNH, Type no. Rh. 8548; Gabriel, 1927: 109], here

designated [examined].

Phyciodes sestia (Hewitson) Rober, 1913: 446, pi. 90, row g [fig. 9], row h [figs 1, 2].
Phyciodes saturata Rober, 1913: 446, pi. 90, row h [fig. 3] [valid name? — apparently proposed in

synonymy of P. sestia (Hewitson)]. LECTOTYPE 9, ECUADOR: 'Cachabe, low c., xi. 1896, ex Rosenberg'

(BMNH), here designated [examined].

Phyciodes sestia sestia (Hewitson); Hall, 1930: 171; Forbes, 1945: 170.
Phyciodes sestia sestia 9 f. satura Rober; Hall, 1930: 172; Forbes, 1945: 170.

REVISION OF PHYCIODES 135

$ forewing 23-24 mm, upperside black with fulvous markings in a pattern very like E. letitia 9, a curved
longitudinal streak from wing-base to s2 prominent. 9 wings broader, postdiscal markings white, discal
markings as in J, cream-yellow or fulvous (f. saturata).

Genitalia. o, in dorsal view, tegumen shoulders not prominent, scaphial extension rather narrow but
spiny terminal bosses well defined, posterior border of juxta prominent, saccus slender, harpe slender,
almost straight. 9 not examined.

DISTRIBUTION. Ecuador (also recorded from Colombia by Hall, 1929: 172).

Eresia coela Druce comb, rev., stat. rev.

(Figs 103, 104, 388, 389)

Eresia coela Druce, 18746: 37; Godman & Salvin, 1882: 187, pi. 21, figs 2, 3; Schaus, 1913: 344, pi. 50,
fig. 8; Rober, 1913: p. 91, row c [fig. 4]. LECTOTYPE 9 ('cT), COSTA RICA: Limon (BMNH, Type no.
Rh. 8550, Gabriel; 1927: 33), here designated [examined].

Phyciodes coela (Druce) Rober, 1913: 444.

Phyciodes sestia coela (Druce); Hall, 1930: 172; Forbes, 1945: 170.

c? forewing 22 mm, like E. sestia, upperside black, markings fulvous; hindwing upperside fulvous discal
band wider, submarginal spots not prominent; hindwing underside discal band terminating before black
marginal border, submarginal spots small, discrete and well defined. 9 forewing 23-25 mm, broad,
apex rounded, upperside black, markings white as in E. sestia but lacking basal coloured area; hindwing
fulvous discal field larger, otherwise as in E. sestia.

Genitalia. $ tegumen wide, shoulders defined, scaphial extension narrow, spiny bosses not well formed,
posterior border of juxta slightly prominent, saccus narrow, posterior section of valve slender, harpes slender,
almost straight. 9 with rami of the bursal support unusually large.

DISTRIBUTION. Costa Rica, coastal regions (Schaus).

NOTE. I have examined 5 ? in the BMNH, and 2 <$ and 1 ? in the Hall Coll. (BM, Brighton).

Eresia oblita (Staudinger) comb, rev., stat. rev.

(Figs 105, 390)

Phyciodes oblita Staudinger, 1885: 93; Rober, 1913: 446. 1 c?, 6 9 syntypes, VENEZUELA: Puerto Cabello

(MNHU, Berlin).

Eresia oblita (Staudinger); Rober, 1913: pi. 92, row d [fig. 5].
Phyciodes carme oblita Staudinger; Hall, 1929: 141; Forbes, 1945: 164.

c? forewing 21 mm, upperside dark brown, markings fulvous yellow, large spots in oblique row as in
E. polina, additional spots in cell and a costal bar less prominent; hindwing upperside discal bar 4-5 mm
wide at cell, tapering slightly to inner margin, narrow post-discal and submarginal lunules well defined;
forewing underside markings similar, paler; hindwing underside markings like E. polina but submarginal
band orange with black proximal border. 9 similar, slightly larger.

Genitalia. <3 tegumen in dorsal view rather narrow, shoulders not prominent, scaphial bosses large,
spines numerous, posterior border of juxta gently convex. 9 not examined.

DISTRIBUTION. Venezuela (San Esteban Valley) (Hall, 1929: 141 erroneously indicates
Staudinger's material as coming from Colombia).

Eresia carme Doubleday comb. rev.
(Figs 106-108, 391-393)

Eresia carme Doubleday, [1847]: pi. 20.

Phyciodes carme (Doubleday) Staudinger, 1885: 93.

cJ forewing 20-23 mm, upperside dark brown, markings fulvous, large spots in slb-s2, and in s4 in an
oblique series; hindwing upperside discal band usually tapers sharply to inner margin, white or pale in

136 L. G. HIGGINS

sla-slc; underside hindwing and forewing markings yellow-buff, usually gleaming. + larger, forewing
upperside with additional postdiscal costal mark, spots yellow, hindwing fulvous discal band wider.
Genitalia. See below.

DISTRIBUTION. Venezuela and Colombia.
Two subspecies, genitalia probably differ slightly.

Eresia carme carme Doubleday
(Figs 106, 107, 391, 392)

Eresia carme Doubleday, [1847]: pi. 20, fig. 5; [1848]: 183. LECTOTYPE ?, VENEZUELA (BMNH,

Type no. Rh. 8541; Gabriel, 1927: 26), here designated [examined].
Phyciodes carme (Doubleday); Staudinger, 1885: 93; Rober, 1913: 446.
Phyciodes carme carme (Doubleday); Hall, 1929: 140; Forbes, 1945: 164.

cJ forewing upperside fulvous markings scanty, post-discal spot in sib quadrilateral, spot in s2 similar but
displaced slightly distad, spot in s4 similar, spot in s5 and costal markings vestigial.

Genitalia. $ , in dorsal view, tegumen wide, shoulders variable, scaphial extension short, terminal spines
large but bosses not well defined, posterior border of juxta gently curved, saccus tapering; penis robust,
ostium-keel small.

DISTRIBUTION. Venezuela, Colombia (?).

Eresia carme laias Godman & Salvin, comb. rev.
(Figs 108, 393)

Eresia laias Godman & Salvin, 1879: 151, pi. 14, fig. 1. LECTOTYPE^, COLOMBIA: Frontino, Antioquia
(T. K. Salmon) (BMNH, Type no. Rh. 8542; Gabriel, 1927: 70), here designated [examined].

Phyciodes laias (Godman & Salvin) Rober, 1913: 446, pi. 90, row g [figs 7, 8].

Phyciodes laias lycus Hall, 1928a: 12; Hall, 1929: 144. Holotype <3, COLOMBIA: Rio Aguaca Valley
(BM, Brighton) [examined]. Syn. n.

Phyciodes laias laias (Godman & Salvin); Hall, 1929: 143.

Phyciodes (Eresia) carme laias (Godman & Salvin); Forbes, 1945: 164.

Phyciodes (Eresia) carme lycus Hall; Forbes, 1945: 164.

S like carme carme but forewing upperside markings differ slightly; fulvous postdiscal spots in sib + s2 + s3
united to form an elongate macule, spot in s5 and small submarginal spots in s3 and s4 all well
defined (3 specimens); markings may be extended by a cell-spot and a postdiscal costal spot, both well
defined (form lycus). 9 unknown.

Genitalia. <3 , in dorsal view, like carme carme but posterior section of valve very slender (£. carme laias
(lycus), single preparation).

DISTRIBUTION. W. Colombia.

DISCUSSION. E. carme is a rare species, not well represented in the BMNH. E. laias was
described as an independent species, with lycus as a subspecies. I have followed Forbes and
retained laias as a subspecies of carme, but it seems more likely that it simply represents a local
form of minor taxonomic importance. The BMNH includes three specimens of laias and two
specimens sufficiently well marked to be placed as f. lycus. The additional markings can often
be seen, vaguely outlined, on normal $ specimens of carme. The underside wing-markings
show close relationship to E. polina (included by Forbes as a subspecies of carme).

Eresia polina Hewitson comb, rev., stat. rev.

(Figs 109, 394, 395)

Eresia polina Hewitson, [1852]: [60], pi. [30], fig. 6. LECTOTYPE J, ECUADOR: 'Quito' (BMNH,

Type no. Rh. 8540; Gabriel, 1927: 98), here designated [examined].
Eresia encina Felder & Felder, 1861 : 103. LECTOTYPE rf, ECUADOR (BMNH), here designated [examined].

[Specimen bears Felders' original determination label.]

REVISION OF PHYCIODES 137

Phyciodes polina f. intermedia Rober, 1913: 446, pi. 92, row g [fig. 5]. c? syntype(s), BOLIVIA (depository

unknown).

Phyciodes polina (Hewitson) Staudinger, 1885: 93; Rober, 1913: 445; Hall, 1929: 142.
Phyciodes (Eresia) carme polina (Hewitson); Forbes, 1945: 164.

3 forewing 27-28 mm, upperside black with 3 large yellow spots placed obliquely in sib, s2 and s4,
discoidal and costal markings vestigial; on hindwing a wide yellow transverse band tapering slightly to
inner margin; underside forewing base fulvous, additional yellow spots at cell-end, on costa beyond cell
and small apical markings; hindwing underside pale yellow, gleaming, veins brown, and basal, sub-basal,
post-discal and submarginal stripes brown, the last proximally bordered black and wide. cJ similar, yellow
markings often enlarged.

Genitalia. <$ organs large, in dorsal view tegumen strongly chitinised, scaphial extension short, terminal
bosses large with prominent spines, juxta with posterior border gently curved, harpes slender, almost
straight. 9 not examined.

DISTRIBUTION. Colombia, Ecuador, Peru, Bolivia.

Eresia alsina Hewitson comb. rev.
(Figs 110, 111,396-398)

Eresia alsina Hewitson, 1869b: 33; Hewitson, [1870]: [35], pi. [20], figs 62, 63; Godman & Salvin, 1882: 186;

1901: 671. LECTOTYPE 9, NICARAGUA: Chontales (T. Belt) (BMNH, Type no. Rh. 8552; Gabriel,

1927: 10), here designated [examined].

Phyciodes alsina (Hewitson); Rober, 1913: 448, pi. 90, row k [fig. 5]; Hall, 1930: 174; Forbes, 1945: 169, 189.
[Eresia heliconina Rober; Rober, 1913: pi. 91, row a [fig. 2] (d). Misidentification.]
[Phyciodes (Eresia) eutropia Hewitson; Young, 1973: 87. Misidentification.]

c? forewing 25-26 mm, upperside orange-fulvous, wing margins black, 3 very oblique black bars, proximal
bar narrow, tapering to a single line as it meets the outer margin, apical markings and marginal spots
yellowish. 9 larger, forewing 28-29 mm, wider, apex more rounded, markings similar.

Genitalia. £ like E. eunice, tegumen short, in dorsal view shoulders well defined, lateral walls of
scaphial extension slightly divergent, terminal spiny bosses rounded, posterior border of juxta almost
straight, saccus wide, tapering. 9 like E. eunice (not figured).

DISTRIBUTION. Nicaragua, Costa Rica.

BIOLOGY. Young (1973) describes the early stages, host plant, mimicry and other aspects of the
biology of what appears to be this butterfly, under the name eutropia Hewitson.

Eresia cissia (Hall) comb, n., stat. n.
(Figs 112,399-401)

Phyciodes ithomioides cissia Hall, 1928a: 12; Hall, 1930: 181, pi. 2, fig. 5 (J); Forbes, 1945: 169.
Holotypec?, COLOMBIA: Juntas, Cauca Valley (de Mathan) (BMNH) [examined].

c? forewing 25 mm, apex rounded, compared with E. ithomioides by Hall, but upperside ground-colour
yellow, forewing markings more regular, basal area yellow, with black postdiscal costal bar followed by
small yellow postdiscal and submarginal markings; hindwing base and discal area yellow, series of yellow
submarginal spots regular, enclosed in wide black marginal band. 9 not examined.

Genitalia. d in dorsal view, tegumen short, shoulders prominent, terminal bosses with a few large spines
but not really well defined, posterior border of juxta straight, saccus wide, tapering rapidly to pointed apex;
penis as in Fig. 401.

DISTRIBUTION. Colombia (Cauca Valley, Juntas (2 ^)).

Eresia eutropia Hewitson comb. rev.
(Figs 113, 114,402)

Eresia eutropia Hewitson, 18746: 56. LECTOTYPE^, PANAMA: Santa Fe (BMNH, Type no. Rh. 8553;
Gabriel, 1927: 48), here designated [examined].

138 L. G. HIGGINS

[Phyciodes eutropia (Hewitson); Godman & Salvin, 1882: 187, pi. 21, fig. 1 ; Rober, 1913 : 447; Hall, 1930: 175;
Forbes, 1945: 169-70, 189. Misidentifications: all referred to E. melaina.]

<$ forewing 22-24 mm, outer margin slightly excavate, upperside black, markings pale yellow or white,
much of sib and lower half of cell fulvous, but inner margin black, oblique discal band yellow or white,
irregular, broken into spots by black veins, spot at base of s3 out of line, postdiscal band usually somewhat
irregular, spot in s4 more or less divided, submarginal spots small; hindwing upperside fulvous, costa and
outer margin black, series of submarginal spots complete from slc-s7; underside markings enlarged, often
confluent; hindwing costa black. 9 forewing 27 mm, wide, apex rounded, outer margin convex, markings
as in c£, submarginal spots well developed, hindwing underside submarginal spots white.

Genitalia. $ tegumen rather wide in dorsal view, shoulders developed, lateral walls of scaphial extension
slightly divergent, terminal bosses not well defined, spines large, posterior border of juxta prominent,
saccus tapering. 9 not examined.

DISTRIBUTION. Panama, W. Colombia.

NOTE. This species was incorrectly identified by Godman & Salvin, Rober, Hall and Forbes;
their descriptions and figures refer to the new species described below under the name melaina.

Eresia mimas (Staudinger) comb, n., stat rev.

(Figs 116, 174)

Phyciodes mimas Staudinger, 1885: 93; Rober, 1913: 448. <£, 9 syntypes, COLOMBIA: Rio San Juan
(E. Troetsch) (? MNHU, Berlin; BM, Brighton) [d syntype in BM, Brighton, examined].

Phyciodes alsina subfasciata Rober, 1913: 448, pi. 92, row c [fig. 3] (as mimas). Syntype(s), COLOMBIA
(depository unknown). Syn. n.

Phyciodes eutropia subfasciata Rober; Hall, 1930: 177; Forbes, 1945: 170.

Phyciodes eutropia mimas Staudinger; Hall, 1930: 178; Forbes, 1945: 170.

<J forewing 26 mm, like E. eutropia, but differs on forewing upperside, postdiscal (subapical) band
regular, crossed by v5 and v6, slightly curved and continued by enlarged submarginal spot in s3 (perhaps
variable), black mark in mid-cell terminates abruptly at median vein, inner margin not black, pale
submarginal spots vestigial or absent. 9 not identified (Hall, 1930: 178, records 1 9 in BMNH).
Genitalia. Not examined.

DISTRIBUTION. W. Colombia (R. San Juan).

DISCUSSION. Rober appears to have been confused about the identity of mimas, of which the
type-locality is not Rio Dagua as he states. His name subfasciata is accepted here for the
figure on his pi. 92, row c [fig. 3], which shows nominate mimas. There is a single specimen in
BMNH and two specimens in Hall Coll., all ex Staudinger, one of the latter recorded as a
'co-type' (Hall, 1930: 178), and all labelled Rio San Juan. The phenotype is easily recognisable,
the markings constant, clearly related to E. eutropia, which also occurs in Colombia. No
specimen is available for dissection.

Eresia quintilla Hewitson comb, rev., stat rev.

(Figs 122, 403, 404)

Eresia quintilla Hewitson, [1872]: [30], pi. [15], fig. 83. LECTOTYPE 9, ECUADOR (Buckley] (BMNH,

Type no. Rh. 8554; Gabriel, 1927: 102), here designated [examined].
Phyciodes quintilla (Hewitson) Rober, 1913: 447, pi. 90, row i [fig. 2].
Phyciodes eutropia quintilla (Hewitson); Hall, 1930: 178; Forbes, 1945: 170.

cJ like E. mimas but forewing upperside discal and post-discal markings bright yellow, fused and expanded
to enclose the short postdiscal costal mark at cell-end, subapical orange band absent and submarginal
pale spots present, especially well defined at wing apex; hindwing underside with or without black basal
shade in s7. 9 similar, usually slightly larger.
Genitalia (single preparation). J tegumen wide, tapering, shoulders well defined, scaphial extension lateral

REVISION OF PHYCIODES 139

walls not diverging, terminal bosses with large spines, posterior border of juxta gently convex, saccus
tapering, harpes slightly sinuous (slide no. 2576). 9 with usual generic characters, bursal support well
defined.

DISTRIBUTION. Ecuador.

Eresia poedlina Bates comb, rev., stat. rev.

(Figs 115, 121)

Eresia poedlina Bates, 1866: 133; Godman & Salvin, 1882: 188, pi. 20, figs 19, 20 [lectotype].

LECTOTYPE 9, PANAMA: Veragua, Santa Fe (Arce} (BMNH, Type no. Rh. 8566; Gabriel, 1927: 98),

here designated [examined].
Phyciodes poedlina (Bates) Rober, 1913: 448.

Phydodes ithomioides poedlina (Bates); Hall, 1930: 181; Forbes, 1945: 169.
Phydodes eutropia confirmans Hall, 1930: additional page. Holotype cJ, COSTA RICA (A. G. M. Gillott)

(BMNH) [examined]. Syn. n.

cJ upperside black, markings white or yellowish, forewing 26 mm, a short red basal flush present, 3 pale
spots in s2, basal and submarginal spots in s3, oval spot in cell, 2 spots in s4 (an additional basal spot
may be present), 3 spots in s5, series of small submarginal spots complete from slb-s8; hindwing
upperside orange-red, costa and outer margin black, tapering to anal angle, with prominent white
submarginal spots in s6 and s7; underside forewing spots similar, slightly larger. 9 forewing wider,
markings slightly larger but arrangement similar, hindwing submarginal white spots in complete series;
hindwing underside often with disk flushed grey-blue (absent in lectotype).
Genitalia. Not examined.

DISTRIBUTION. Panama (Santa Fe, Chiriqui, Laguna River), Costa Rica.

DISCUSSION. The postdiscal spots grouped together in sib + s2 and in s4 + s5 form a useful
character. The area between the postdiscal costal bar and the apical submarginal spots is
unmarked. I have examined 1 d, 3 9 in the BMNH, and 1 $, 1 $ in the Hall coll. (BM,
Brighton). Both males and 3 9 show the blue-grey, slightly vitreous ('violaceous-grey') suffusion
on the hindwing underside referred to by Hall in his description of confirmans; this does not
appear in the 9 lectotype of poedlina, although this looks identical in all other respects.

Eresia melaina sp. n.

(Figs 117, 118)

[Eresia eutropia Hewitson; Godman & Salvin, 1882: 187, pi. 21, fig. 1 (J); Rober, 1913: pi. 92, row b

[fig. 5] (c?). Misidentifications.]

Eresia dismorphina Butler; Godman & Salvin, 1882: 187, pi. 20, figs 17, 18 (9). Misidentification.]
Phydodes eutropia (Hewitson); Rober, 1913: 447. Misidentification.]

Phydodes eutropia eutropia (Hewitson); Hall, 1930: 176; Forbes, 1945: 170. Misidentifications.]
Phydodes eutropia eutropia ab. nivifera Hall, 1930: 177. Syntypes 4 $, PANAMA, COLOMBIA (BMNH)
[examined].

cJ forewing 25-26 mm, upperside black, with scanty yellowish markings, a fulvous flush below cell in sib,
pale mark near cell-end usually elongate, followed by a small spot at base of s3, post-discal spots in s2
and s4 prominent, isolated, forming a useful specific character, postdiscal bar from costa to s4, submarginal
spots small, often indistinct but series usually complete from slb-s8; hindwing upperside fulvous, costa
black, continued narrowly along outer margin, enclosing white spots in s6 and s7; underside markings
similar. 9 similar, upperside markings yellowish or white, slightly enlarged; forewing base of cell and much
of s2 fulvous but base of s3 black ; hindwing black marginal border wider, series of submarginal spots
complete, a faintly yellow flush often present internal to marginal border in s5.
Genitalia. Not examined.

Holotype o, Panama: Veraguas (Arce), ex Godman & Salvin Coll. (BMNH).

Paratypes. Panama: 1 $, same data as holotype; 2 9, Santa Fe (Arce); 2 9, Lion Hill (McLeannan)
(BMNH). Colombia: 1 9 (BMNH).

140 L. G. HIGGINS

DISTRIBUTION. Panama (Veragua, El Valle, Santa Fe), Colombia.

NOTE. No male specimen is available for dissection. The <$ figured by Godman & Salvin and
erroneously named eutropia was accepted as 'type' by Rober and later by Hall, which has led,
for many years, to confusion in identification of these species. This specimen is now fixed as the
holotype of melaina.

Eresia sticta Schaus comb, rev., stat. rev.

(Figs 119,405)

Eresia sticta Schaus, 1913: 344, pi. 50, fig. 7. rf, ? syntypes, COSTA RICA (? BMNH) [identity of these

types not confirmed].

[Eresia coela Druce; Rober, 1913: pi. 91, row c [fig. 5] ($). Misidentification.]
Phyciodes sticta (Schaus) Rober, 1924: 1030, pi. 192, row h [fig. 4] (^).
Phyciodes ithomioides sticta (Schaus); Hall, 1930: 181; Forbes, 1945: 169.

cJ forewing 26-27 mm, narrow, apex elongate, slightly truncate, upperside black with white markings
consisting of small spots; a prominent spot over discoidal vein and many spots in postdiscal and
submarginal areas; hindwing upperside orange-red, costa and inner margin broadly black, margin
tapering in anal angle; underside similar, but hindwing black borders extended and series of white
submarginal spots complete. $(?) larger, forewing 30 mm, wide, apex rounded.

Genitalia.d tegumen short, wide, scaphial bosses not well defined, posterior border of juxta gently curved
(slide no. 1023).

DISTRIBUTION. Costa Rica, ? Colombia (Hall, 1930: 182).

DISCUSSION. In my opinion, the ?-form attributed to this species by Hall may be incorrectly
placed. The species seems to be rare; 2 £ in BMNH, 1 J in Hall Coll. (BM, Brighton). The
fate and identity of Schaus' original syntypes are uncertain; a pair from Carillo in the BMNH,
collected in February (the female certainly by Schaus) may represent the original material,
described from specimens in the BMNH.

Eresia ithomioides Hewitson comb. rev.
(Figs 120, 124, 124a, 406, 407)

Eresia ithomioides Hewitson, [1864]: [18].

Phyciodes ithomoides [sic] (Hewitson) Rober, 1913: 448.

c? forewing 28-29 mm, narrow, apex rounded, outer margin straight, markings consisting of white or yellow
spots in discal and postdiscal areas; hindwing oval, disc fulvous, black outer margin enclosing pale
submarginal spots; hindwing underside with black basal mark in s7, black marginal border extended along
v7. ? differs, larger, forewing 36 mm, broad, apex rounded; upperside markings as in<3 but spots larger;
hindwing upperside orange-fulvous, submarginal spots obscure in wider dark border; underside black
borders wide, series of submarginal spots prominent and complete (described from single specimen in
BMNH).

Genitalia.J organs wide, tegumen short, in dorsal view scaphial bosses not well defined, posterior border
of juxta wide, gently convex, base of saccus wide; penis in side view slender, ostium-keel prominent.

DISTRIBUTION. Colombia, ? Panama.
Two subspecies.

Eresia ithomioides ithomioides Hewitson
(Figs 120, 124a)

Eresia ithomioides Hewitson, [1864]: [18], pi. [9], fig. 20. LECTOTYPE rf, COLOMBIA: 'New Granada',

Bogota (E. Birchell) (BMNH, Type no. Rh. 8565; Gabriel, 1927: 66), here designated [examined].
Phyciodes ithomoides [sic!] (Hewitson); Rober, 1913: 448, pi. 90, row k [fig. 2].
Phyciodes ithomioides ithomioides (Hewitson); Hall, 1930: 179; Forbes, 1945: 169.

REVISION OF PHYCIODES 141

cJ upperside forewing base black ; hindwing fulvous discal area reduced by extension of black marginal
border, rarely entirely black.

DISTRIBUTION. Colombia.

Eresia ithomioides pseudocelemina (Strand) stat n.
(Fig. 124)

Phyciodes pseudocelemina Strand, 1916: 6, pi. 16, fig. 13. Holotype^, COLOMBIA (BMNH) [examined].
Phyciodes ithomioides ithomioides <$ f. pseudocelemina Strand; Hall, 1930: 180.

cJ differs from nominate form: forewing upperside base and discoidal cell bright fulvous; hindwing
fulvous area extending to black marginal border, black extension along v7 variable, sometimes vestigial;
hindwing underside black mark in s7 reduced.

DISTRIBUTION. Colombia (Magdalena Valley, May (C. Allen); 5^ in BMNH).

DISCUSSION. The BMNH has two males of the nominate form, and five males of pseudocelemina,
all from Colombia, and it may be that they represent a simple cline, but with so little material it
is not possible to be sure of the best taxonomic rank. There is a single <$ in the Hall Coll.
(BM, Brighton).

The Eresia species poecilina, sticta, ithomioides and melania form a difficult group. It is
especially difficult to assemble the females with the appropriate male. The arrangement adopted
here may need revision when more material is available. The species are all rare, poorly
represented in collections, most often by single specimens taken at long intervals by different
collectors, in various localities. I have accepted the precise arrangement of spots on the upperside
of the forewing as the most reliable character for definition of species and for association of
sexes. It seems possible that one or more of the females may be polymorphic, but I have
had to describe as a distinct species (anomala) one well-known but previously unnamed $ form,
for which I have not been able to identify a male.

Eresia anomala sp. n.

(Fig. 123)

[Phyciodes ithomioides (Hewitson); Rober, 1913: 447, pi. 90, row k [figs 3, 4] (9, as Eresia on pi.).

Misidentification.]
[Phyciodes ithomioides ithomioides (Hewitson); Hall, 1930: 180 (9 only). Misidentification.]

$ forewing 30 mm, upperside dark grey-brown, markings yellowish spots arranged as in E. poecilina but
spot at base of s2 constantly absent, postdiscal spot in s2 well separated from submarginal spot; hindwing
upperside dark grey-brown, series of submarginal spots yellowish, large, complete, a slightly variable fulvous
flush extending along inner margin and across s2, a postdiscal series of yellow, rectangular marks present in
s2-s5. All markings similar on underside.

Holotype 9, Colombia: Muzo, 1924 (Apollinar Maria) (ex Joicey Coll.) (BMNH).
Paratypes. Colombia: 4 9, (no further data); 3 ?, Bogata; 2 9, Valdevia, 1897 (Pratt) (BMNH);
3 9, 'Colombia' (BM, Brighton).

There is little variation in the series of ten specimens in the BMNH and three specimens in
the Hall Coll. (BM, Brighton). Compared with other species in this section, E. anomala appears
to be relatively common. It was attributed to ithomioides by Rober, but the absence on the
forewing of the spot at base of s2 makes this attribution doubtful.

Eresia nigripennis Salvin comb, rev., stat. rev.

(Figs 125, 125a, 408)

Eresia nigripennis Salvin, 1869: 170; Godman & Salvin, 1882: 186, pi. 20, figs 15, 16. LECTOTYPE <J,
COSTA RICA: Cache (Carmiol) (BMNH, BM Type no. Rh. 8563; Gabriel, 1927: 86), here designated
[examined].

142 L. G. HIGGINS

Eresia dismorphina Butler, 1872: 78; Butler, 1874: 182, pi. 63, fig. 1 (?). LECTOTYPE 9, COSTA RICA:

(van Patten) (BMNH; BM Type no. Rh. 8564), here designated [examined].
Phyciodes nigripennis (Salvin) Rober, 1913: 447, pi. 90, row i [fig. 4] #); Hall, 1929: 169.
Phyciodes (Eresia) phillyra nigripennis (Salvin); Forbes, 1945: 168.

cJ forewing 26-27 mm, elongate, apex truncate, outer margin excavate; upperside black with rather small
postdiscal and submarginal white or yellowish spots; hindwing orange-red, costal and marginal borders
black, enclosing pale submarginal spots in s7 and s8; 9 larger, forewing 30-31 mm, wider, less abruptly
truncate, pale markings usually yellowish, fulvous basal stripe indistinct; hindwing as in<^.

Genitalia.d like E. phillyra, shoulders of tegumen not greatly developed, posterior border of juxta curved,
saccus rather slender (slide no. 896). 9 not examined.

DISTRIBUTION. Restricted to Costa Rica.

Eresia emerantia Hewitson comb. rev.
(Figs 126, 127, 409, 410)

Eresia emerantia Hewitson, [1857]: [43], pi. [22], figs 7, 11; Rober, 1913: pi. 92, row d [fig. 6] (5).

LECTOTYPE J, COLOMBIA: 'New Grenada' (BMNH, Type no. Rh. 8551; Gabriel, 1927: 45), here

designated [examined].

Eresia emerentia [sic!] Hewitson; Hewitson, [1864]: pi., fig. 14 (d).
Phyciodes emerantia (Hewitson); Rober, 1913: 446.
Phyciodes celemina Rober, 1913: 448, pi. 90, row k [fig. 1] (9). LECTOTYPE 9, COLOMBIA: Cundinamarca,

vii. 1903 (de Mathau) (BMNH), here designated [examined].
Phyciodes emerentia [sic!] (Hewitson); Hall, 1930: 173; Forbes, 1945: 170, 189.

d forewing 25 mm, upperside fulvous, black margins wide, apex black, enclosing 3 white subapical spots;
hindwing underside yellowish, marginal spots pale yellow, large. 9 differs : forewing wider, postdiscal area
black with irregular white spots, submarginal spots white, series complete on both wings, resembling
E. ithomioides pseudocelemina <$ .

Genitalia. 3 in dorsal view elongate, shoulders of tegumen not conspicuous, inferior lamina of scaphial
membrane well developed, spiny bosses defined, posterior border of juxta prominent, falces relatively very
slender (slide no. 2740). 9 with usual generic characters, rami of bursal support long.

DISTRIBUTION. Colombia.

Eresia moesta Salvin & Godman comb. rev.
(Figs 173,411,412)

Eresia moesta Salvin & Godman, 1868: 145. LECTOTYPE 9, ECUADOR: Canelos (Pearce) (BMNH,

Type no. Rh. 8577; Gabriel, 1927: 82), here designated [examined].
Eresia ildica Hewitson, 1869a: 24 [index]; Hewitson, [1872]: [30], pi. [15], fig. 84 (9). LECTOTYPE 9,

ECUADOR: Aguano (Buckley) (BMNH, Type no. Rh. 8578; Gabriel, 1927: 62), here designated

[examined].
Eresia cerquita Dognin, 1894: 680. LECTOTYPE^, ECUADOR: environs de Loja (BMNH), here designated

[examined]. [Specimen bears Dognin's m/s type label.]
Eresia (Phyciodes) ildica var. heliconiformis Strand, 19 12a : 143. Holotype, ECUADOR : Macas (' Niepelt Coll. ' ;

MNHU, Berlin ?).
Phyciodes ildica fassli Rober, 1913: 447, pi. 90, row i [figs 5, 6]. cJ, 9 syntypes, COLOMBIA: Upper Rio

Negro, 2500 ft [830 m] (A. H. Fassl) (Rothschild Coll., BMNH; Hall, 1930: 189; not found, but one

c? from Bogota, ex Rothschild, is labelled 'Phyciodes fassli Rob./Type'. in Rober's handwriting). Syn. n.
Phyciodes moesta moesta (Salvin & Godman); Hall, 1930: 187; Forbes, 1945: 170.
Phyciodes moesta moesta f. cerquita (Dognin); Hall, 1930: 188; Forbes, 1945: 171.
Phyciodes moesta moesta f. ildica (Hewitson); Hall, 1930, 1930: 188, Forbes, 1945: 171.
Phyciodes moesta fassli Rober; Hall, 1930: 188; Forbes, 1945: 170.

cJ forewing 27-28 mm, upperside grey or rarely with slight reddish basal flush, more or less translucent,
costa and outer margin rather broadly black, otherwise unmarked; hindwing usually similar, occasionally
with red discal flush, costa and outer margin black, white submarginal spots usually present. + similar,
slightly larger.

REVISION OF PHYCIODES 143

The following colour forms have been named: 1, inner margin of hind wing red, f. cerquita; 2, forewing
base and discal field of hindwing brick -red, f. ildica; 3, hindwing inner margin red, shading to yellow
over disc, Lfassli.

Genitalia. J organs large, tegumen short, shoulders not well developed, scaphial extension short,
terminal bosses well defined with large spines, saccus base wide, tapering rapidly; penis with usual generic
characters (slide no. 2793).

DISTRIBUTION. S. Colombia, Ecuador, Peru.

Eresia phaedima Salvin & Godman comb. rev.
(Figs 128, 413)

Eresia phaedima Salvin & Godman, 1868: 146. LECTOTYPE J, PERU: Pozzuzo (Pearce) (BMNH,
Type no. Rh. 8579; Gabriel, 1927: 97; genitalia slide no. 639), here designated [examined].

Eresia prisca Hopffer, 1874: 349; Rober, 1913: pi. 92, row e [fig. 1] ($). 3 syntypes, 'Mexico'; PERU:
Chanchamayo (Thamm) (MNHU, Berlin).

Phyciodes prisca (Hopffer); Rober, 1913: 447.

Melitaea (Phyciodes) magniplaga Rober, 1927: 98, fig. LECTOTYPE <J, COLOMBIA: Canungucho,
iii.-vi. 1926 (BMNH), here designated, [examined]. Syn. n.

Phyciodes phaedima (Salvin & Godman) Hall, 1930: 185; Forbes, 1945: 169, 189.

Phyciodes magniplaga (Rober) Hall, 1930: 186; Forbes, 1945: 170, 189.

c? forewing 29-30 mm, like moesta, costa and outer margin black, base with fulvous extending along sib
to outer margin, short, oblique black band across cell, cell apex and postdiscal area translucent except
postdiscal black band from costa to v4; hindwing like moesta but disc fulvous-brown, costa and outer
margin broadly black, white submarginal spots prominent, apex of s6 usually yellow. 9 similar, larger.

A single colour form has been named : upperside pale areas yellow, not fulvous-brown, f. magniplaga.

Genitalia. d organs wide, tegumen narrow, scaphial extension longer than that of moesta, saccus rather
narrow, elongate. 9 not examined.

DISTRIBUTION. Peru, Colombia. According to Hall (1930: 186), Hopffer's record for Mexico is
erroneous.

Eresia datis Hewitson comb. rev.
(Figs 129, 414)

Eresia datis Hewitson, [1864]: [17].
Phyciodes datis (Hewitson) Hall, 1930: 182.

c? forewing 29-30 mm, upperside black, a broad discal band cream-white, a short basal streak below cell
and field of hindwing brown or orange-brown; hindwing black marginal border rather wide, enclosing
white submarginal dots, more distinct on underside. 9 unknown.

DISTRIBUTION. Peru, Bolivia.
Two subspecies.

Eresia datis datis Hewitson
(Fig. 129)

Eresia datis Hewitson, [1864]: [17], pi. [9], fig. 14. LECTOTYPE^ [UPPER AMAZON— see Hall, 1930: 183]
(BMNH, Type no. Rh. 8575, Gabriel, 1927: 38), here designated [examined].

? Phyciodes fenestrata Rober, 1914; 450, pi. 91, row c, fig. 3. Lectotype ^, PERU: Huayabamba, SE. of
Chachapoyas, 3500 ft [1300 m] (O. T. Baron) (BMNH) [examined]. [Specimen bears Robers M/S
determination labels.]

Phyciodes datis datis (Hewitson); Hall, 1930: 182; Forbes, 1945: 169, 189.

? Phyciodes datis fenestrata Rober; Hall, 1930: 183; Forbes, 1945: 169.

? Phyciodes fenestrata derivata Bryk, 1953: 92. Holotype 9, PERU: Roque (NR, Stockholm).

S upperside basal streak of forewing and field of hindwing orange-brown.

144 L. G. HIGGINS

Genitalia. <$, in dorsal view, like clara, tegumen with a curious small central hiatus in two specimens
examined, shoulders defined, scaphial extension with moderately divergent lateral walls, posterior border
of juxta with small central prominence, saccus Ungulate (slide no. 623).

DISTRIBUTION. 'Upper Amazon' (lectotype), Peru.

Eresia datis corybassa Hewitson
(Fig. 414)

Eresia corybassa Hewitson, 1874a: 6. LECTOTYPE^, BOLIVIA (Buckley) (BMNH, Type no. Rh. 8574;

Gabriel, 1927: 35), here designated [examined].
Phyciodes cornelia Staudinger in litt.; Rober, 1914: 450, pi. 92, row g [fig. 1] (as Eresia). [Invalid:

proposed in synonymy with corybassa.]

Phyciodes datis corybassa (Hewitson); Hall, 1929: 184; Forbes, 1945: 169.
? Phyciodes corybassa mimicry Bryk, 1953: 91. Holotypec?, PERU: Roque (NR, Stockholm).

Like E. datis datis, but <3 upperside basal streak of forewing and field of hindwing dark mahogany brown.
$ similar, but larger.
Genitalia. Like E. datis datis, lateral walls of scaphial extension more divergent (single preparation).

DISTRIBUTION. S. Peru, Bolivia.

Eresia margaretha Hewitson comb. rev.
(Figs 130, 415)

Eresia margaretha Hewitson, [1872]: [29], pi. [15], fig. 77. LECTOTYPE rf, COLOMBIA: Bogota ('New
Granada') (Chesterton) (BMNH, Type no. Rh. 8576; Gabriel, 1927: 78), here designated [examined].

Phyciodes margaretha (Hewitson) Rober, 1914: 450, pi. 91, row b [fig. 1] (as Eresia).

Phyciodes margaretha manto Hall, 1930: 185. Holotype^, COLOMBIA: Rio Dagua (BMNH) [examined].
Syn. n.

Phyciodes margaretha margaretha (Hewitson); Hall, 1930: 184; Forbes, 1945: 169.

Phyciodes (Eresia) margaretha manto Hall; Forbes, 1945: 169.

3 forewing 25 mm, upperside black, discal band and subapical spots yellow; basal streak and discal field
of hindwing brick-red. 9 unknown.

Genitalia. <$ , in dorsal view, organs wide, tegumen wide, scaphial extension and terminal spiny bosses
well defined, posterior border of juxta almost straight, saccus wide, tapering rapidly to a pointed apex
(slide no. 2718).

DISTRIBUTION. Colombia.

Eresia eunice (Hiibner) comb. rev.
(Figs 131, 132, 135, 416-420)

Nereis fulva eunice Hiibner, [1807]: pi. 9.
Eresia eunice (Hiibner) Hewitson, [1857]: [44].

cJ forewing 22-25 mm, upperside base fulvous, distally paler, yellowish, markings black, a short stripe on
median vein, a wide oblique band across end of cell which terminates abruptly in s3, but continues narrowly
along v3 to blend with black outer margin, apex black, sometimes with traces of yellowish markings;
hindwings similar, yellowish anterior to the wide black subcentral transverse band, submarginal area usually
darker fulvous, subcostal stripe and outer margin black. ? similar, often slightly larger.

Genitalia. 3 tegumen wide, tapering to well-formed shoulders, lateral walls of scaphial extension divergent,
terminal bosses well defined with small spines, posterior border of juxta convex, valve apex slender, penis
apex slender, ostium-keel prominent. ? bursal duct chitinised, short, bursal support globular, supporting
rami short.

DISTRIBUTION. Brazil, the Guyanas, Panama, Venezuela, Colombia, Bolivia, Peru, Ecuador.
DISCUSSION. Three subspecies (or major colour phenotypes) are described below. It is difficult

REVISION OF PHYCIODES 145

to define these forms on a geographical basis although one or other of them may preponderate
in a given area. Hall (1929: 157) states that nominate eunice is almost constant in Guyana and
W. Brazil if the form pella is excluded. Intermediates between the several forms are very common
and in some districts, according to Hall, two or more may fly together.

Eresia eunice eunice (Hiibner)
(Figs 131, 418-420)

Nereis fulva eunice Hiibner, [1807]: pi. 9, 4 figs. Syntype(s)^, 9, [SOUTH AMERICA] (depository unknown).
Eresia pella Hewitson, [1852]: [59], pi. [30], fig. 2. Syntypes, [BRAZIL] ('Amazon'): W. W. Saunders & W. C.

Hewitson Colls (not in BMNH, probably in UM, Oxford).
Eresia eunice (Hiibner) Hewitson, [1857]: [44]; Bates, 18646: 191.
[Eresia olivencia Bates; Rober, 1913: pi. 92, row b [fig. 1] (?). Misidentification.]
Phyciodes eunice eunice (Hiibner); Hall, 1929: 156; Forbes, 1945: 167, 171.

c£ upperside fulvous, distal areas slightly paler in tone; hindwing yellow transverse discal band with
black band posteriorly, followed by fulvous area. 9 similar, often showing greater colour contrast, on
forewing distal pale areas may be yellow.
Genitalia. As described above.

DISTRIBUTION. Brazil (including Lower Amazon), the Guyanas, Colombia, Peru, Ecuador.

Eresia eunice olivencia Bates
(Fig. 132)

Eresia eunice var. olivencia Bates, 18646: 191. LECTOTYPE rf, BRAZIL: Sao Paulo de Olivencia

(H. W. Bates) (BMNH, Type no. Rh. 8555; Gabriel, 1927: 89), here designated [examined].
Eresia drypetis Godman & Salvin, 1878: 269; Godman & Salvin, 1882: 184, pi. 20, figs 11, 12. Holotype^,

PANAMA: Lion Hill (McLeannan) (BMNH, Type no. Rh. 8556; Gabriel, 1927: 43) [examined]. Syn. n.
Phyciodes olivencia (Bates) Staudinger, 1885: 193.

Ecesia [sic!] klagesii Weeks, 1906: 195. Syntypes, VENEZUELA: Suapure (? MCZ, Cambridge).
Eresia klagesii Weeks; Weeks, 1911: 1, pi. 1 (?^).
Phyciodes olivencia f. gudruna Rober, 1913:447, pi. 92, row b [fig. 2] (as Eresia). Syntype(s), ' Upper Amazon'

(depository unknown).
Phyciodes olivencia f. brunhilda Rober, 1913: 447, pi. 92, row b [fig. 3] (as Eresia). Syntype(s), ' Upper Amazon '

(depository unknown).
Phyciodes olivencia polymnia Rober, 1913: 447. <$, 9 syntypes, COLOMBIA: Medina, 1600 ft [530 m] (Fassl)

(depository unknown).

Phyciodes eunice eunice f. olivencia (Bates); Hall, 1929: 157.
Phyciodes eunice eunice f. klagesii (Weeks); Hall, 1929: 158.
Phyciodes eunice eunice f. brunhilda Rober; Hall, 1929: 158.

Phyciodes eunice drypetis (Godman & Salvin); Hall, 1929: 158; Forbes, 1945: 167, 171.
Phyciodes (Eresia) eunice brunhilda Rober; Forbes, 1945: 166.
Phyciodes (Eresia) eunice olivencia (Bates); Forbes, 1945: 167.
Phyciodes (Eresia) eunice gudruna Rober; Forbes, 1945: 167.
? Phyciodes eunice homogena Bryk, 1953: 89-90. cJ, 9 syntypes, BRAZIL: Amazonas, Taracua (NR,

Stockholm).

<3 Upperside orange-brown, variable, black markings often reduced, pale areas prominent but without
colour-contrast. Several named phenotypes, as noted in synonymy above. 9 similar.

DISTRIBUTION. Brazil (locally), Venezuela, Guatemala, Colombia, Peru, Panama.

Eresia eunice esora Hewitson
(Figs 135, 417)

Eresia esora Hewitson, [1857]: [44], pi. [22], fig. 12. LECTOTYPE^, BRAZIL: [Espirito Santo] (BMNH,

Type no. Rh. 8558; Gabriel, 1927: 47), here designated [examined].
Phyciodes esora (Hewitson) Rober, 1913: 446.

146 L. G. HIGGINS

[Eresia eunice (Hiibner); Rober, 1913: pi. 92, row a [fig. 3] (y). Misidentification.]

Phyciodes eunice esora (Hewitson); Hall, 1929: 158; Forbes, 1945: 167, 171; Hayward, 19646: 340,

pi. 18, fig. 13.
? Phyciodes eunice 9 f. albella Zikan, 1937: 385. Holotype 9, BRAZIL: [Fazenda] 'Jerusalem', Alegre,

Espirito Santo, May (? Rio de Janeiro).

cJ forewing 22-23 mm, upperside apex solid black, oblique black costal bar wide, tapering, pale discal and
postdiscal markings yellow, with marked colour-contrast; hindwing upperside yellow anterior to the black
transverse band, £ genitalia examined, slide no. 2717. 9 similar, usually larger.

DISTRIBUTION. Brazil (constant from Pernambuco, Bahia and Mato Grosso southwards;
northern and north-western populations are more variable but remain distinct from eunice
eunice), Ecuador, Paraguay, NE. Argentina.

DISCUSSION. Of all the modifications of this variable species, esora most nearly shows the
characters and distribution of a true geographical race.

Eresia etesiae (Hall) comb, n., stat. n.
(Figs 133, 134, 421)

Phyciodes erysice etesiae Hall, 1929: 160, pi. 1, figs 8, 9. Holotype cJ, FRENCH GUIANA: 'Cayenne'

(BMNH) [examined].
Phyciodes (Eresia) erysice etesiae Hall; Forbes, 1945: 168.

c? like E. eunice, forewing 25-26 mm, base fulvous, including most of cell and base of s2, oblique mark
in mid-cell and irregular discal bar black, the last followed by yellow postdiscal band heavily suffused
black, round yellow spots in s2 and in s4 rather prominent; hindwing upperside fulvous, costa and
postdiscal band black, narrow submarginal lunules present from slc-s8; forewing underside markings
expanded, pale yellow.

Genitalia. 3 tegumen like eunice, slightly wider, posterior border of juxta gently convex, valve apex more
robust, penis more massive, slightly sinuous (slide no. 1026). 9 not examined.

DISTRIBUTION. French Guiana, N. Brazil.

Eresia erysice (Geyer) comb. n.
(Fig. 136)

Melinaea erysice Geyer, 1832: 28, figs 717, 718. Syntype(s), 'SOUTH AMERICA' (depository unknown) [For

date and authorship see Hemming, 1937(1): 478.]
Phyciodes erysice Hiibner; Kirby, 1871: 176.
Phyciodes erysice erysice Geyer; Hall, 1929: 160
Phyciodes (Eresia) erysice erysice (Hiibner); Forbes, 1945: 168, 189.

c? forewing 27 mm, upperside like E. etesiae, forewing black discal band irregular but complete from
costa to outer margin at v2, followed by well-developed, regular yellow oblique band, pale postdiscal spots
in s2 and s4 absent (prominent in etesiae). Hindwing upperside transverse fulvous band broadly suffused
with yellow; underside markings similar, hindwing underside submarginal spots white, series complete.
9 larger, wings more rounded, pattern paler and more diffuse.
Genitalia. Not available for dissection.

DISTRIBUTION. Brazil, restricted to dense forest in Bahia, very rare and local (K. Brown, pers.
comm.).

NOTE. The original figures 717, 718 were published by Hiibner in [1826], but the name was
published by Geyer in 1832, who also indicates that the original material was in the Franck
Collection (Strasbourg). Hiibner's anonymous catalogue of the Franck Collection ([Hiibner],
[1825]) does not list the species by name, but it may have been included by him as one of the
un-named entries; it is possible that Hiibner made his figures of the insect during his visit to
Strasbourg.

REVISION OF PHYCIODES 147

Eresia casiphia Hewitson comb. rev.

(Figs 137, 138, 422)

Eresia casiphia Hewitson, 1869a: 25, [index]; Hewitson, [1870]: [38], pi. [21], figs 72, 73. LECTOTYPE^,
ECUADOR: Jorge (Buckley] (BMNH, Type no. Rh. 8561 ; Gabriel, 1927: 27), here designated [examined].
Phyciodes casiphia (Hewitson) Rober, 1913: 446; Hall, 1929: 166, pi. 1, fig. 7 (?).
Phyciodes (Eresia) casiphia (Hewitson); Forbes, 1945: 167, 189.

c? upperside fulvous, markings black ; forewing 23 mm, apex and wing-margins black, proximal oblique
bar wide at costa, tapering to anal angle, distal oblique bar shorter, wider, fusing with outer margin,
yellowish preapical bar present but rather obscure; hindwing post-discal transverse bar complete, fusing
with marginal black border, a small white spot at s7 enclosed within the border. 9 similar, forewing 27 mm,
wider, upperside postdiscal band pale yellow, series of white submarginal spots complete around both wings.
Genitalia. Like E. eunice, slightly wider, in dorsal view posterior border of juxta almost flat (slide
no. 1027). 9 genitalia not examined.

DISTRIBUTION. Ecuador.

Eresia mechanitis Godman & Salvin comb, rev., stat. rev.

(Figs 139, 423)

Eresia mechanitis Godman & Salvin, 1878a: 269; Godman & Salvin, 1882: 183, pi. 20, figs 13, 14.

Lectotype 9, NICARAGUA: Chontales (Janson) (BMNH, Type no. Rh. 8557; Gabriel, 1927: 79),

designated by Godman & Salvin, 1882: 184 [examined].

Phyciodes mechanitis (Godman & Salvin) Rober, 1913: 447, pi. 90, row h [fig. 5] (^).
? Phyciodes mechanitis f. daguana Bargmann, 1928: 236. Holotypec?, COLOMBIA: Rio Dagua (via Joicey to

BMNH?— not found).
Phyciodes eunice mechanitis (Godman & Salvin); Hall, 1929: 159; Forbes, 1945: 167.

Like E. eunice, <$ forewing 27 mm, upperside first oblique band continues to outer margin (in eunice ceases
abruptly in s3, or dwindles to a narrow line along v3), apical yellow band defined, forming a wide black
preapical band (in eunice yellow apical markings are vestigial or absent); hindwing outer margin angled at
v7 (less acutely angled in eunice}. 9 similar.

Genitalia. Like E. eunice, scaphial extension short, spiny bosses relatively large, posterior border of juxta
convex (slide no. 1243). 9 not examined.

DISTRIBUTION. Nicaragua, Costa Rica, ? Colombia.

DISCUSSION. Hall considered E. mechanitis to be a subspecies of E. eunice. Represented in the
BMNH by 4^, 3 ?, in which the markings are constant; the series does not associate well
with E. eunice. The genitalic characters are equivocal in this group.

Eresia pelonia Hewitson comb. rev.
(Figs 140-142, 424, 425)

Eresia pelonia Hewitson, [1852]: [59], pi. [30], fig. 3. LECTOTYPE c?, ECUADOR: 'Quito' (BMNH, Type

no. Rh. 8559; Gabriel, 1927: 95), here designated [examined].
Eresia ithomiola Salvin, 1869: 171. LECTOTYPE^, PERU: Cosnipata Valley (H. Whitely) (BMNH, Type

no. Rh. 8560; Gabriel, 1927: 66), here designated [examined].

Eresia pelonia Hewitson; Hewitson, [1870]: [35], pi. [20], figs 64-67 [fig. 66, f. ithomiola].
Phyciodes callonia Staudinger, 1885: 92, pi. 36. 2 <$, 1 9 syntypes, PERU: Pebas; Jurimaguas (Hahnel)

(MNHU, Berlin).

Phyciodes callonia var. murena Staudinger, 1885: 92. 2 syntypes, PERU: Chanchamayo (? MNHU, Berlin).
Eresia callonioides Strand, I9l2b: 181. Holotype 9, PERU (BMNH) [examined].
Phyciodes pardalina Rober, 1913: 447, pi. 90, row h [fig. 6]. LECTOTYPE rf, PERU: Pebas, xii. 1906

(de Mathan) (BMNH) here designated [examined]. [Specimen bears Rober's M/S determination label.]
Phyciodes pardalina apicalis Rober, 1913: 447, pi. 90, row i [fig. 1]. LECTOTYPE^, PERU: Rio Chuchurras,

R. Palcazu, 320 m (W. Hoffmanns] (BMNH) here designated [examined]. [Specimen bears Rober's M/S

determination label.]

148 L. G. HIGGINS

Phyciodes murena heliconina Rober, 1913: 447. LECTOTYPE 9, PERU: Pebas, xii. 1906 (de Mathan)
(BMNH), here designated [examined]. [Specimen bears Rober's M/S' determination label.]

Phyciodes murena Staudinger; Rober, 1913: 447, pi. 92, row c [fig. 1] (9 — as Eresia).

Phyciodes pellonia [sic!] (Hewitson) Rober, 1913: 446, pi. 92, row b [fig. 4] (as Eresia).

Eresia callonioides var. pastazana Strand, 1920: 147. Holotype 9, ECUADOR: Upper Pastaza, c. 1000 m,
x-xii. 1906 (BMNH) [examined].

Phyciodes pelonia f. hewitsonii Hall, 1929: 164. Holotype ?, ECUADOR (BMNH) [examined].

Phyciodes pelonia pelonia (Hewitson); Hall, 1929: 161; Forbes, 1945: 167.

Phyciodes pelonia f. callonia Staudinger; Hall, 1929: 162.

Phyciodes pelonia f. pardalina Rober; Hall, 1929: 163.

Phyciodes pelonia f. apicalis Rober, Hall, 1929: 163.

Phyciodes pelonia f. ithomiola (Salvin); Hall, 1929: 163.

Phyciodes pelonia f. heliconina Rober; Hall, 1929: 163.

Phyciodes pelonia f. callonioides (Strand); Hall, 1929: 164.

Phyciodes (Eresia) pelonia apicalis Rober; Forbes, 1945: 167.

Phyciodes (Eresia) pelonia callonia Staudinger; Forbes, 1945: 168.

Phyciodes (Eresia) pelonia callonioides (Strand); Forbes, 1945: 168.

Phyciodes (Eresia) pelonia heliconina Rober, Forbes, 1945: 168.

Phyciodes (Eresia) pelonia hewitsonii Hall; Forbes, 1945: 168.

Phyciodes (Eresia) pelonia ithomiola (Salvin); Forbes, 1945: 168.

cJ forewing 26-28 mm, upperside fulvous, with or without yellow postdiscal band, and with black markings,
forewing apex usually broadly black; hindwing upperside fulvous, subcostal and discal stripes black,
submarginal spots white (if present), best developed on underside. ? similar, larger. (For principal colour
forms see 'Discussion'.)

Genitalia. 3, in dorsal view, tegumen rather narrow, shoulders not prominent, scaphial bosses appear
incompletely developed with large spines on the external aspects on each side, posterior border of juxta
almost straight; penis specifically distinctive, posterior section massive, slightly sinuous (slide no. 2558).
9 not examined.

DISTRIBUTION. Peru, Ecuador, Bolivia, Brazil ('Amazonas').

DISCUSSION. This species is extremely variable. Three principal phenotypes are recognised which,
according to Hall (1929), may all fly together; their genitalia are similar. These are described
below as 'forms'.

1. c? upperside fulvous, forewing postdiscal band yellow, wing apex black; hindwing fulvous, costal and
transverse bars black.

Recorded from Upper Amazon, Peru and Ecuador, f. pelonia.

2. $ upperside as in f. pelonia, but forewing upperside without yellow post-discal band ; hindwing broadly
black, by extension of costal and transverse black bars, submarginal black border absent. This form occurs
in both sexes.

Recorded from Peru, Ecuador and Bolivia, f. ithomiola (= murena; apicalis is similar).

3. Like f. pelonia, but forewing apical areas enclose yellow subapical spots which may be large ; hindwing
upperside often more or less suffused with black. A form close to f. pelonia, but with forewing apical areas
slightly spotted, was named pardalina by Rober.

Recorded from Ecuador (most common in $), f. callonia (= hewitsonii).

Eresia phillyra Hewitson comb. rev.
(Figs 143, 144, 185, 426, 427)

Eresia phillyra Hewitson, [1852]: [59], pi. [30], fig. 1. 9 syntype(s), MEXICO (W. W. Saunders Coll.;

probably in UM, Oxford), several 9 specimens of 'phillyra' ex Hewitson in BMNH].
Eresia ezorias Hewitson, [1857]: [44], pi. [22], fig. 13. Lectotype^, MEXICO (BMNH, Type no. Rh. 8562;

Gabriel, 1927: 49), here designated [examined].

Phyciodes phillyra (Hewitson) Rober, 1913: 448; pi. 92, row c [figs 3, 4] (as Eresia); Hall, 1929: 167.
Phyciodes (Eresia) phillyra phillyra (Hewitson); Forbes, 1945: 168, 189.

REVISION OF PHYCIODES 149

cJ upperside fulvous, markings black; <$ forewing 26-27 mm, narrow, apex truncate, outer margin
excavate, wing margins black, oblique post-discal band complete; hindwing with black transverse bar
below centre, prominent. 9 larger, forewing 32 mm, broader, apex more rounded, outer margin straight,
upperside markings as in 3, but apical and marginal spots pale yellow.

Genitalia. <3 like E. eunice, terminal bosses of scaphial extension well defined, posterior border of juxta
almost straight; penis massive (slide no. 2569).

DISTRIBUTION. Mexico, Guatemala, ? Honduras, ? Brazil.

Eresia aveyrona Bates comb, rev., stat. rev.

(Figs 145, 428)
Eresia aveyrona Bates, 1864: 192.

$ forewing very narrow, upperside fulvous, oblique costal bar and wing margins black. 9 similar, forewing
slightly wider.

Genitalia. J like E. perna, scaphial extension narrower, spiny bosses less prominent (single preparation).
9 not examined.

DISTRIBUTION. The Guyanas, Brazil (on Lower Amazon), Venezuela, Costa Rica, Ecuador.
Two subspecies, with similar genitalia.

Eresia aveyrona aveyrona Bates
(Figs 145, 428)

Eresia aveyrona Bates, 1864b: 192, pi. 10, fig. 4. Holotype 9, BRAZIL Aveyros, on the Tapajos

(H. W. Bates) (BMNH, Type no. Rh. 8543; Gabriel, 1927: 20), [examined].

Phyciodes aveyrona (Bates) Staudinger, 1885: 93; Rober, 1913: 446, pi. 92, row d [fig. 3] (as Eresia).
Phyciodes perna aveyrona (Bates); Hall, 1929: 153; Forbes, 1945: 165.

c£ forewing 24 mm, outer margin slightly excavate, upperside fulvous yellow, veins lined black, fulvous
apical macule divided. 9 forewing wider, upperside hindwing orange submarginal line more prominent.

DISTRIBUTION. The Guyanas, Brazil, Venezuela, Costa Rica.

Eresia aveyrona mylitta Hewitson nom. rev., stat. n.

Eresia mylitta Hewitson, 1869a: 26 [index]. LECTOTYPE rf, ECUADOR: St. Ines (Buckley) (BMNH, Type

no. Rh. 8544; Gabriel, 1927: 21, as bella), here designated [examined].
Eresia bella Hewitson, 1869a: ii. [Unjustified replacement name for Eresia mylitta Hewitson; originally

proposed by Kirby who intended to place mylitta Hewitson in the same genus as the older proposed

nominal species Melitaea mylitta Edwards.]

[Eresia bella Hewitson; Hewitson, [1870]: [37], Eresia pi. 9, fig. 71.]
Phyciodes bella Kirby, 1871 : 177. [Replacement name proposed by Kirby on transferring mylitta (Hewitson)

and the older established mylitta (Edwards) both. to Phyciodes; Kirby was apparently unaware that

Hewitson had already published the name in the form of an unjustified replacement.] Syn. n.
Phyciodes bella Kirby; Rober, 1913: 446, pi. 88, row i [fig. 2] (tf).
Phyciodes perna bella (Hewitson); Hall, 1929: 153; Forbes, 1945: 162, 166.

c? forewing 22.5 mm, smaller, upperside bright fulvous, veins not lined black, apical fulvous macule not
divided. 9 not seen.

DISTRIBUTION. Ecuador.

DISCUSSION. Eresia mylitta is placed here as a subspecies with E. aveyrona. It appears to be
present only in Ecuador, and rather isolated from nominate aveyrona, which is not recorded
from this region. The butterflies are rare and more information is required before their true
relationship can be decided.

150 L. G. HIGGINS

Eresia perna Hewitson comb. rev.
(Figs 146, 429)

Eresia perna Hewitson, [1852]: [60], pi. [30], fig. 5. LECTOTYPE J, BRAZIL: Rio de Janeiro (BMNH,

Type no. Rh. 8545; Gabriel, 1927: 95), here designated [examined].
Phyciodes alma Staudinger, 1885: 93, pi. 36 (3).c?, 9 syntypes, BRAZIL: Sta Catharina, Blumenau (MNHU,

Berlin).

Phyciodes perna (Hewitson) Rober, 1913: 446, pi. 92, row g [figs 4, 5] (as Eresia).
Phyciodes perna perna (Hewitson); Hall, 1929: 152; Forbes, 1945: 165.

J forewing 21 mm, very elongate, upperside black, markings yellow to cream, an obscure basal stripe across
cell terminates in a yellow spot in s2, a second more distal spot is placed in s4; hindwing upperside black,
with yellow transverse band. 9 larger, forewing 25 mm, broader, apex more rounded, post-discal spots
tend to cream-white.

Genitalia. J tegumen short in dorsal view, shoulders prominent, scaphial terminal bosses appear oval,
well defined, spines small and irregular, posterior border of juxta with central prominence, saccus wide,
tapering (slide no. 2716). 9 not examined.

DISTRIBUTION. Brazil (coastal districts from Espirito Santo southwards to Santa Catarina, rare
and local, flying in dense vegetation; Keith Brown, pers. comm.).

Eresia levina Hewitson comb. rev.
(Figs 151, 430, 431)

Eresia levina Hewitson, [1872]: [29], pi. [15], fig. 78; Fassl, 1912: 122 (9). Syntype(s) rf, COLOMBIA:

'New Granada' (W. W. Saunders Coll.; probably in UM, Oxford).
M orpheis felderi Hopffer, 1874: 351. Holotype -J, [COLOMBIA] (not 'Bolivia' according to K. S. Brown

in litt.) (Warscewicz) (MNHU, Berlin).
Phyciodes levina (Hewitson) Staudinger, 1885: 94, pi. 36 (9); Rober, 1914: 450, pi. 92, row e [fig. 3]

fa— as Eresia); Hall, 1930: 189; Forbes, 1945: 163, 189.

Phyciodes levina f. decorata Rober, 1914: 450. Syntypes, COLOMBIA (depository unknown).
? Phyciodes levina werneri Hering, in Hering & Hopp, 1925: 193. Holotype 9, COLOMBIA: Rio Micay

(MNHU, Berlin).
Phyciodes levina ab. decorata Rober; Hall, 1930: 180; Forbes, 1945: 163.

c? forewing 24 mm, upperside black, discal band gleaming blue, otherwise unmarked. 9 slightly larger,
blue discal band edged white.

Genitalia. Distinctive, large, £ tegumen firmly chitinized, shoulders prominent, dorsal and ventral aspects
of scaphial extension extensively chitinized but spiny bosses weakly developed, posterior border of juxta
sinuous, terminal section of valve armed along inner aspect with fine teeth; penis apex rounded,
ostium-keel absent (slide no. 98). 9 not examined.

DISTRIBUTION. Colombia.

Eresia actinote Salvin comb. rev.
(Figs 152, 153, 432)

Eresia actinote Salvin, 1869: 171. LECTOTYPE^, PERU: Cosnipata Valley (H. Whitely) (BMNH, Type

no. Rh. 8573, Gabriel, 1927: 6), here designated [examined].
Eresia acraea Hopffer, 1874: 349.^ syntype(s), PERU (MNHU, Berlin).
Phyciodes actinote (Salvin) Rober, 1913: 448; pi. 92, row f [fig. 2] (^ — as Eresia).
Phyciodes actinote limbata Rober, 1913: 448; Hall, 1930: 191. c? syntype(s), BOLIVIA (depository unknown)

Syn. n.

Phyciodes actinote actinote (Salvin); Hall, 1930: 190.
Phyciodes (Eresia) actinote (Salvin); Forbes, 1945: 164, 189.

J forewing 24-25 mm, upperside both wings orange-brown, veins and margins black, forewing with a wide
black band which runs obliquely from costa to outer margin at s2. j unknown.

Genitalia. J1 tegumen narrow in dorsal view, scaphial extension short, shoulders well defined, terminal

REVISION OF PHYCIODES 151

bosses large, heavily spined, posterior border of juxta gently convex, saccus short, harpes robust, slightly
curved (slide no. 618).

DISTRIBUTION. Peru, Ecuador, Bolivia.

Eresia selene (Rober) comb, n., stat. rev.

(Figs 154, 433, 434)

Phyciodes selene Rober, 1913: 445, pi. 90, row f [figs 7, 8]. LECTOTYPE J, COLOMBIA: Canon de Tolima,
1700 m, i. 1910 (A. H. Fassl) (BMNH), here designated [examined]. [Specimen bears Robers M/S
determination label.]

Phyciodes etia selene Rober; Hall, 1929: 72.

Phyciodes (Eresia) claudina selene Rober; Forbes, 1945: 159, 189.

cJ forewing 16-17 mm, upperside fulvous, margins and costal band black to outer margin; hind wing discal
area fulvous, underside paler with confused grey markings. 9 similar.

Genitalia. 3 tegumen wide, strongly chitinized, scaphial extension well defined, lateral walls and terminal
spiny bosses chitinized, posterior border of juxta gently convex, saccus wide, short, posterior process
of valve massive but short (slide no. 832). 9 not examined.

DISTRIBUTION. Colombia, Ecuador.

DISCUSSION. The $ genitalia of this species are typical of Eresia, but in general habitus and
wing-markings there is an approach to species of the genus Tegosa, which has led to previous
uncertainty as to the position of this species.

CASTILIA gen. n.

Type-species: Eresia castilla Felder & Felder. Gender: feminine.

In external characters the species are most variable, but they divide naturally into a non-mimetic group,
and a mimetic group composed of the first five species described below, all mimics of Actinote species. All
these occur in the mountainous regions of north-western South America; the females are rare, perhaps
unknown in C. neria and C. northbrundii. In four species (C. perilla, C. castilla, C. occidentalis and
C. neria) the venation of the forewings is unusual, vein 1 1 arising at or a little beyond the cell-end. In
the hindwings the lower discocellular vein arises at or close to the junction of v2 and v3. These four species
also show a distinctive character in the palpi, which are clothed with such long and abundant hair-scales
that the terminal segments are almost hidden. The sexes differ.

The non-mimetic species have the forewings elongate, with pale markings on a dark ground, and are
variable in size, with one species (C. angusta) quite small. Wing venation is normal and the palpal
hairs are not conspicuously long, but relationship with the mimetic group is evident in the structure of the
male and female genitalia. With the exception of C. fulgora, all these species were associated with
C. perilla by Forbes (1945) in his group 12. In the non-mimetic forms the sexes are usually similar. In
the mimetic group the special characters of the palpi and the unusual wing-venation could be taken as
calling for generic separation. I have not suggested this as the group is small and generic separation would
mask the relationship to the remaining species.

Genitalia. $ in dorsal view, tegumen short, un-armed, often feebly chitinised, valve terminating in a single
point, curved strongly upwards in characteristic fashion, preceded by a slight dilatation and lacking
pre-apical teeth, saccus deeply cleft in all species; penis straight, often massive, ostium-keel well developed.
9 sterigma like Eresia, bursal duct usually chitinised and well formed, bursal support large, the usual
lateral ribs exceptionally well developed in some species (e.g. C. eranites).

DISTRIBUTION. The generic range extends through central and western South America, including
Colombia, Venezuela, Peru, Ecuador and Bolivia. The species C. angusta reaches NW. Argentina
and Brazil in the Mato Grosso. With the exceptions of C. fulgora and C.fausta, there are adequate
series of all species in the BMNH collections, but often from a single locality only. Especially
in the non-mimetic group, the external characters of all species seem to be stable.

152 L. G. HIGGINS

Key to species of Castilia

Males.

Note. It has not been practicable to include C. griseobasalis (p. 156) and C. chinantlensis (p. 157)
in this key.

In C. eranites, females are like males with additional white postdiscal spots. Females of C. neria and
of C. nortbrundii are not known; those of C. occidentalis are very rare. Females resemble males in
C. angusta, C. fulgora, C. ofella and C. myia (probably also in C. fausta, but not yet described). The
species C. griseobasalis and C. chinantlensis have not been included in the key; the former is closely
related to C. myia, the latter to C. eranites.

1 Mimetic of Actinote, hindwing underside lined dark along and between veins ... 2
Markings variable, not mimetic of Actinote 5

2 Hindwing upperside costal margin white nortbrundii (p. 154)

Hindwing upperside costal margin dark .......... 3

3 Forewing upperside basal area fulvous, postdiscal area black, unmarked . . neria (p. 154)
Forewing upperside not so marked 4

4 Forewing upperside postdiscal band red, yellow or white, width 8 mm or more . perilla (p. 153)
Forewing upperside postdiscal band red, width 4 mm or less castilla (p. 152), occidentalis (p. 153)

5 Forewing 14 mm, very elongate, width 6 mm, or less .... angusta (p. 157)
Forewing wider, species larger 6

6 Upperside markings fulvous on black ........... 7

Upperside markings white or pale yellow 8

7 Forewing upperside wide fulvous band extending from base to post-discal area eranites (p. 155)
Forewing upperside base black fulgora (p. 155), fausta (p. 155)

8 Forewing upperside with white spot at cell-end ofella (p. 156)

Forewing upperside lacking white spot at cell-end myia (p. 156)

Females, mimetic species.

1 Forewing upperside base yellow-buff perilla (p. 153)

Forewing upperside base black 2

2 Hindwing upperside discal area red-brown, black area matt .... castilla (p. 152)
Hindwing upperside inner margin shaded red, black area with blue reflections occidentalis (p. 153)

Castilia castilla (Felder & Felder) comb. n.
(Figs 155, 156, 435-437)

Eresia castilla Felder & Felder, 1862b: 419; Felder & Felder, 1867: 395, pi. 50, figs 7-10; Rober, 1913:
pi. 91, row b [fig. 2]; pi. 92, row e [fig. 2]. LECTOTYPE <$, COLOMBIA: Bogota Cordillera, near Pandi,
5600 ft. [1700 m] (Lindig) (BMNH), here designated [examined].

Phyciodes castilla (Felder & Felder) Rober, 1914: 450.

Phyciodes castilla ;) f. virilis Rober, 1914: 450. 9 syntype(s), COLOMBIA (depository unknown).

Phyciodes castilla castilla (Felder & Felder); Hall, 1930: 191.

Phyciodes (Tritanassa) castilla castilla (Felder & Felder); Forbes, 1945: 185, 190.

c2 forewing 25-26 mm, upperside black, discal band red, oblique, 3-5 mm wide; hindwing upperside black,
unmarked; hindwing underside yellow-grey, 2 small basal red marks and radiating black stripes along and
between veins. $ larger, forewing 27-28 mm, upperside dark grey, discal band irregular, yellow,
4-6 mm wide; hindwing upperside dark grey, base and discal area brown with black radiating stripes as on
underside; a male-like v form (virilis), with upperside dark grey-brown and discal band red, has been
recorded by Rober.

Venation (Fig. 437). Forewing stalk of veins 7-10 arises with vll and v6 at cell-end.

Genitalia. 3 tegumen short, scaphial extension weakly chitinised, conical, posterior border of juxta slightly
concave, harpe massive at base but tapering rapidly (slide no. 615). ? ductus and bursal support firmly
chitinised, the large scutum especially noticeable.

DISTRIBUTION. Colombia (not recorded elsewhere).

REVISION OF PHYCIODES 153

Castilia occidentalis (Fassl) comb, n., stat. n.

(Figs 147, 148, 438-440)

Eresia castilla f. occidentalis Fassi, 1912: 122. J, 9 syntypes, COLOMBIA: West Cordillera, 1500-2000 m

(depository uncertain — MNHU, Berlin?).

Eresia castilla occidentalis Fassl; Rober, 1913: pi. 91, row c [fig. 1]; pi. 92, row e [fig. 2].
Phyciodes castilla occidentalis (Fassl) Rober, 1914: 450; Hall, 1930: 192; Forbes, 1945: 185.
? Phyciodes aurora Rober, 1914: 449; pi. 91, row b [fig. 3] (as Eresia); Hall, 1930: 192; Forbes, 1945: 190

(as ? 9 of castilla). LECTOTYPE 9, ECUADOR: 'Zamora, 3000-4000 ft.' (0. T. Baron) (BMNH) here

designated [examined]. [Specimen bears Rober's M/S determination label.]

cJ forewing 21 mm, like C. castilla but smaller, upperside black, red postdiscal band more oblique and
narrower. 9 larger, upperside gleaming black, forewing discal band slightly wider, red or yellowish ; hindwing
with red anal mark which fades away near discoidal cell. The nominal species aurora Rober may represent
a 9-form of occidentalis; specimens of this sex are extremely rare.

Venation. Stalk of v7-vll divides a little beyond cell, first discocellular vein present but short, v6 free.

Genitalia. 3 similar to C. castilla but smaller, in dorsal view structure of tegumen less well defined,
posterior border of juxta gently convex, valve apex bowed inwards, pre-apical bulge not greatly developed
(slide no. 626).

DISTRIBUTION. Colombia, Ecuador.

Castilia perilla (Hewitson) comb. n.
(Figs 149, 150, 175, 176, 441-443)

Eresia perilla Hewitson, [1852]: [60], pi. [30], fig. 4. LECTOTYPE rf, ECUADOR: 'Quito' (BMNH, Type

no. Rh. 8567; Gabriel, 1927: 95), here designated [examined].
Eresia acraeina Hewitson, [1864]: [17], pi. [9], fig. 15. LECTOTYPE^, [PERU] 'Upper Amazon' (BMNH,

Type no. Rh. 8568; Gabriel, 1927: 6)., here designated [examined].

Eresia aricilla Hopffer, 1874: 350. d syntypes, PERU: Chanchamayo (Thamm) (MNHU, Berlin).
Eresia heliconoides Butler, 1877: 120. LECTOTYPE £, PERU: Ucayali (W. Davis) (BMNH, Type no. Rh.

8569; Gabriel, 1927: 58), here designated [examined].
Phyciodes perilla (Hewitson) Staudinger, 1885: 94.
Phyciodes acraeina f. hilarina Rober, 1913: 448, pi. 92, row e [fig. 5] (as Eresia acraeina). Syntypes,

'Andes' (depository unknown).
? Phyciodes neria f. microdryope Rober, 1913: 448, pi. 92, row e [fig. 4] (as Eresia). Syntype(s),

COLOMBIA (depository unknown).

Eresia amoenides Rober, 1913: pi. 92, row f [fig. 1]. [Published without description.]
Phyciodes acraeina 9 f. aberrans Rober, 1913: 448, pi. 91, row a [fig. 5] (as Eresia acraeina).

9 syntype(s), 'Andes' (depository unknown).

Phyciodes perilla lugubris Rober, 1914: 450. Syntypes, PERU (depository unknown).
Phyciodes pellenea Rober, 1914: 450, pi. 91, row c [fig. 2] (as Eresia). LECTOTYPE 9, BOLIVIA: Reyes

(Stuart) (BMNH), here designated [examined]. [Specimen bears Rober's M/S determination label.]
Phyciodes perilla perilla (Hewitson); Hall, 1930: 194.

Phyciodes perilla f. acraeina (Hewitson); Hall, 1930: 194; Forbes, 1945: 186.
Phyciodes perilla f. aricilla (Hopffer); Hall, 1930: 195; Forbes, 1945: 186.
Phyciodes perilla f. lugubris Rober; Hall, 1930: 195.
Phyciodes perilla 9 f. aberrans Rober; Hall, 1930: 196; Forbes, 1945: 185.
Phyciodes perilla 9 f. pellenea Rober; Hall, 1930: 196; Forbes, 1945: 185.
Phyciodes (Tritanassa) perilla f. perilla (Hewitson); Forbes, 1945: 185, 190.
? Phyciodes pellenea crucifera Bryk, 1953: 92. Holotype 9, PERU: Roque (NR, Stockholm).

A polymorphic and variable species;^ forewing 23 mm, upperside black, wing bases usually more or less
flushed red, discal band on forewing in various shades of red to yellow and cream-white; hindwing
underside yellow-brown, base often flushed reddish, and rayed black along and between veins. 9 larger,
forewing 27-33 mm, upperside dark grey, base yellow-brown, discal band curved, yellow, slightly irregular;
hindwing broadly yellow-brown, margin grey, lightly rayed dark along and between veins. For description
of colour forms, see 'Discussion'.

Venation. Forewing vll and v7-vlO arise together.

154 L. G. HIGGINS

Genitalia.J in dorsal view like C. castilla, slightly smaller, tegumen weakly chitinised, exposing on each
side a broad plate of the inferior lamina of the scaphial extension; posterior border of juxta slightly
irregular, convex; valve terminal process short, turned medially (slide no. 627). 9 ductus short, bursal
support with strong ribs.

DISTRIBUTION. Ecuador, Peru, Bolivia, ? Colombia.

DISCUSSION. The polymorphism has not been investigated as far as I know. Hall (1930: 194)
states 'forms are not geographically separated, although some are more prevalent in certain
localities than others'. The principal colour forms are named as follows.

1, cJ band of forewing pale yellow, bases of both wings broadly flushed red, f. perilla;

2, cJ band of forewing and wing-bases red, f. acraeina ;

3,<J band of forewing red, base of hindwing black, f. aricilla;

4, like aricilla, but band of forewing yellow (mimic of Eueides aides Stichel and Archonias bella
(Cramer), f. lugubris;

5, ? forewing basal suffusion and band yellow, and vestigial yellow apical markings, f. pellenea;

6, forewing band and both wing bases yellow, colour extensive on hindwing, f. aberrans.

Castilia neria (Hewitson) comb. n.
(Figs 157, 444-446)

Eresia neria Hewitson, 1869c: 98; Hewitson, 1869a: 27; [1872]: [29], pi. [15], fig. 80. LECTOTYPE J,
ECUADOR: Sarayacu (Buckley) (BMNH, Type no. Rh. 8572; Gabriel, 1927: 85), here designated
[examined].

Phyciodes neria (Hewitson) Rober, 1914: 449; Hall, 1930: 197.

Phyciodes (Tritanassa) neria (Hewitson); Forbes, 1945: 185, 190.

c? forewing 21 mm, upperside black, wide basal area orange-fulvous, hindwing black; underside forewing
apex and all hindwing grey-brown, rayed black along and between veins. ? unknown.

Venation. Forewing vll and stalk of v7-vlO arise together, beyond cell-end.

Genitalia.J like C. perilla, tegumen short, scaphial extension fragile, membranous; posterior border of
juxta rather prominent; terminal process of valve curved sharply medially; penis robust (slide no. 619).

DISTRIBUTION. Ecuador.

Castilia nortbrundii (Weeks) comb. n.
(Figs 158, 447-449)

Phyciodes nortbrundii Weeks, 1901: 359; Weeks, 1905: 96, pi. 42, fig. 1 ($). 4 J syntypes, BOLIVIA:

Cochabamba district (MCZ, Cambridge).

Eresia nebrites Weymer, 1907: 19. Holotype^, BOLIVIA: Cochabamba (Fruhstorfer) (? MNHU, Berlin).
Phyciodes actinotina Rober, 1914: 449; pi. 91, row b [fig. 7] (as Eresia). J syntype(s), BOLIVIA (depository

unknown).

Phyciodes northbrundii Weeks; Hall, 1930: 197.
Phyciodes (Tritanassa) northbrundii Weeks; Forbes, 1945: 185, 190.

cJ forewing 19 mm, upperside black, large basal area and oblique subapical band orange-fulvous; hindwing
discal area orange-fulvous; underside hindwing pale grey, rather faintly rayed dark brown along and
between veins, sometimes a few small orange markings near base. V unknown.

Venation. In forewing vll arises from subcostal vein before end of cell. This is the normal arrangement
found in other Phyciodini.

Genitalia. _J tegumen short, lateral walls of scaphial extension chitinised and projecting posteriorly;
posterior border of juxta gently curved, saccus short, posterior process of valve long, slender, curved
medially, harpe slender; in side view penis like C. perilla (slide no. 611).

DISTRIBUTION. Bolivia.

REVISION OF PHYCIODES 155

Castilia eranites (Hewitson) comb. n.
(Figs 159, 160, 450-452)

Eresia eranites Hewitson, [1857]: [43], pi. [22], figs 8-10; Godman & Salvin, 1882: 185; 1901: 673.

LECTOTYPE J, COLOMBIA: Bogota ('New Granada') (BMNH, Type no. Rh. 8547; Gabriel, 1927: 46),

here designated [examined].
Phyciodes eranites (Hewitson) Staudinger, 1885: 93; Rober, 1913: 446; Hall, 1929: 164; Forbes, 1945: 184,

190.

Eresia evanides [sic!] Hewitson; Rober, 1913: pi. 92, row c [fig. 5]; row d [figs 1, 2].
[Eresia carme Doubleday; Rober, 1913: pi. 92, row d [fig. 3]. Misidentification; aberrant specimen with

unusual hindwing marking.]

Phyciodes eranites mejicana Rober, 1913: 446. Syntype(s), MEXICO: Presidio (depository unknown).
Phyciodes (Eresia) eranites mejicana Rober; de la Maza, 1978: 39-44, figs 2a, b, 3b.

cJ forewing 22 mm, upperside fulvous with black markings along costa and margins, defining a large basal
and discal orange area, and smaller orange costal and submarginal spots; hindwing orange, borders black
with prominent black macular band almost parallel with the outer margin, equally conspicuous on
underside. 9 larger, markings differ slightly, post-discal area of forewing spots more numerous, white.

Genitalia. <3 apical section of valve slender, apex curved strongly inwards, harpe firmly chitinized,
tegumen weakly chitinized, slightly conical, fragile, posterior border of juxta protruding posteriorly, forming
an acute angle, saccus large, deeply cleft; penis straight, massive, ostium-keel small (slide no. 2587).
9 genitalia, ductus short, bursal support large, posterior rib prominent.

DISTRIBUTION. Mexico, Costa Rica, Nicaragua, Salvador, Panama, Guatemala, Venezuela,
Colombia.

Castilia fulgora (Godman & Salvin) comb. n.

(Figs 161, 453, 454)

Phyciodes fulgora Godman & Salvin, 18786: 261; Godman & Salvin, 1882: 206, pi. 22, figs 17, 18; Rober,
1913:445, pi. 90, row f [fig. 6]; Hall, 1929: 106; Forbes, 1945: 182, 190. LECTOTYPE rf, COSTA RICA: Rio
Sucio (H. Rogers) (BMNH, Type no. Rh. 8456; Gabriel, 1927: 52), here designated [examined].

Phyciodes levana Rober, 1913: 440, pi. 88, row h [fig. 7]. 9 syntype(s), COSTA RICA: Orosi, 1280 m
(A. H. Fassl) (depository uncertain).

3 forewing 15 mm, like C. fulgora but smaller, outer margin slightly excavate, upperside dark brown
bar on discoidal vein, a large oval postdiscal spot in sib, s2 and s3, and a subapical costal bar before a very
small, white apical spot; hindwing with a narrow orange transverse band tapering to vlb; underside
hindwing pale brown with confused cryptic markings. 9 similar, forewing upperside orange postdiscal spot
expanded to form a wide postdiscal band; hindwing upperside white submarginal spot in s7 prominent.
Genitalia. d in dorsal view rather narrow, tegumen and scaphial extension lightly chitinised ; posterior
border of juxta gently undulant, terminal process of valve curved medially, harpe slender; penis like
C. perilla (slide no. 1021). 9 not examined.

DISTRIBUTION. Seen only from Costa Rica.

Castilia fausta (Godman & Salvin) comb. n.
(Figs 162, 455)

Phyciodes faustus Godman & Salvin, 1897: 243; 1901: 680, pi. 108, figs 19, 20 (^); Rober, 1913: 441;
Hall, 1929: 79; Forbes, 1945: 181, 190.^ syntype(s), PANAMA: Chiriqui, ex Staudinger (MNHU, Berlin).

3 forewing 15 mm, like C. fulgora but smaller, outer margin slightly excavate, upperside dark brown
(black when fresh?), markings fulvous, spot at cell-end small; hindwing fulvous transverse band tapering
to inner margin, submarginal lunules obscure excepting well-marked spot in s7; underside marbled light
and dark brown. 9 not seen.

Genitalia. 3 in dorsal view like C. fulgora, tegumen shorter, scaphial extension not well defined,
posterior border of juxta wide with small undulations; falces parallel, not oblique (slide no. 1345).

DISTRIBUTION. Panama (Chiriqui).
A rare species, two males only in BMNH.

156 L. G. HIGGINS

Castilia ofella (Hewitson) comb. n.
(Figs 163, 456-458)

Eresia ofella Hewitson, [1864]: [18], pi. [9], figs 18, 19; Godman & Salvin, 1882: 189; 1901 : 674. Syntype(s),
COLOMBIA: 'New Granada' (E. Birchall Coll.; present depository uncertain, possibly in UM, Oxford;
not recorded by Gabriel, 1927, but 'type' noted by Hall, 1929: 135, as being in BMNH).

Phyciodes ofella (Hewitson) Rober, 1913: 444, pi. 90, row f [fig. 3] (3).

Phyciodes ofella ofella (Hewitson); Hall, 1929: 134.

? Phyciodes ofella guaya Hall, 1929: 135. Holotypec?, ECUADOR (BM, Brighton) [paratype 3 in BMNH
examined].

Phyciodes (Tritanassa) ofella ofella (Hewitson); Forbes, 1945: 183, 185, 190.

Phyciodes (Tritanassa) ofella guaya Hall; Forbes, 1945: 183.

c? forewing 19 mm, outer margin excavate, upperside black, markings white; small spot over cell-end, large
postdiscal mark in slb-s3 which is placed to continue the white transverse band of the hindwing, a few
small spots near wing apex; hindwing upperside veins crossing white transverse band not pigmented;
hindwing underside distal area marbled brown, a row of small, dark spots present in sic, s2 and s3.
$ similar.

Genitalia. d in dorsal view with tegumen very short, scaphial extension greatly reduced ; posterior border
of juxta with low central prominence, apical process of valve very small; harpe slender; penis as in
C. perilla (slide no. 1063). 9 distinctive, ductus long, chitinised, bursal support of unusual shape, partly
membranous.

DISTRIBUTION. Guatemala, Costa Rica, Panama, Colombia, Venezuela, Trinidad, ? Ecuador.

Castilia myia (Hewitson) comb. n.
(Figs 164, 459-461)

Eresia Myia Hewitson, [1874]: [17], pi. [9], figs 16, 17; Godman & Salvin, 1882: 188; 1901: 674.
LECTOTYPE rf, MEXICO (BMNH, Type no. Rh. 8534; Gabriel, 1927: 84), here designated [examined].
Phyciodes myia (Hewitson) Rober, 1913: 444, pi. 90, row e [figs 4, 5].
Phyciodes myia myia (Hewitson); Hall, 1929: 136; Forbes, 1945: 183.

cJ forewing 18 mm, upperside black, markings white, like C. ofella but lacking white cell-spot, round
postdiscal spot in sbl and sb2 conspicuous; hindwing transverse band narrow, crossed by black veins,
small submarginal spot conspicuous in s7; forewing underside base orange-yellow; hindwing underside
marbled brown, with a row of dark submarginal lunules in slc-s5. $ similar.

Genitalia. d like C. eranites, tegumen fragile, posterior border of juxta wide with small undulations, valve
tapering posteriorly to slender, upturned apex, rather sharply curved, harpe gently curved (slide no. 2586).
? genitalia: bursal duct chitinised, support well defined with prominent ribs.

DISTRIBUTION. Mexico, Honduras, Nicaragua, Guatemala.

Castilia griseobasalis (Rober) comb, n., stat. n.

(Figs 165, 462, 463)

Phyciodes myia f. griseobasalis Rober, 1913: 444. Syntype(s), [CENTRAL AMERICA] (depository unknown).

Phyciodes myia griseobasalis Rober; Hall, 1929: 137.

Phyciodes (Anthanassa) myia griseobasalis Rober; Forbes, 1945: 183, 190.

c? forewing 15 mm, variable, upperside like C. myia, but smaller, pale markings reduced, especially the white
postdiscal spot in slb-s2; hindwing white transverse band narrow; forewing underside base yellow-grey.
Genitalia. 3 like C. myia, in dorsal view posterior section of valve conspicuously long, harpe straight,
long and slender (slide no. 1038).

DISTRIBUTION. Guatemala, El Salvador, Costa Rica, ? Mexico.

DISCUSSION. Hall states that C. griseobasalis replaces C. myia in western Guatemala, El Salvador
and Costa Rica. The distinction between the two in size and markings is far from striking, but
it appears to be constant; I have not seen specimens with intermediate characters.

REVISION OF PHYCIODES 157

Castilia angusta (Hewitson) comb. n.
(Figs 166, 464-466)

Eresia angusta Hewitson, [1868]: [34], pi. [19], figs 58, 59. LECTOTYPE^, locality unknown (BMNH,

Type no. Rh. 8535; Gabriel, 1927: 13), here designated [examined].

Phyciodes angusta (Hewitson); Rober, 1913: 445, pi. 92, row h [fig. 6] (^); Hall, 1929: 137.
Phyciodes (Tritanassa) angusta (Hewitson); Forbes, 1945: 185, 190.

c£ forewing 14 mm, narrow, elongate, outer margin weakly excavate, upperside black, markings pale yellow,
small narrow bar at cell-end, round postdiscal spots in s2 and s3 prominent; hindwing upperside with
transverse band and isolated submarginal spot in s7, submarginal lunules vestigial if present. 9 similar.

Genitalia. In dorsal view J organs wide, tegumen fragile, posterior border of juxta very prominent,
valve tapering, rather massive, apex inclined medially, harpe slender; penis ostium-keel present, morula
small (slide no. 2762). 9 bursal duct chitinised, support well formed, scutum large.

DISTRIBUTION. Colombia, Ecuador, Peru, Bolivia, Brazil (Mato Grosso), Argentina (Tueuman).

Castilia chinantlensis (de la Maza) comb. n.
(Figs 171, 172)

Phyciodes (Eresia) chinantlensis de la Maza, 1978: 39, figs la-c, 3a [3, 4-cover]. Holotype^, MEXICO: La
Esperanza, Sierra de Juarez, Oaxaca, 1750 m, 27.xi.1976 (R. de la Maza) (Museo de Historia Natural
de la Ciudad de Mexico) [1^,19 paratypes examined in BMNH].

De la Maza, in his original description, provides good photographs of the adults (including colour figures
of both sexes on the rear cover of the journal issue), and a dorsal view of the male genitalia.

Genitalia (based on examination of paratype in BMNH).^ very close to C. eranites; in dorsal view the
tegumen is well defined, nearly quadrilateral; valve less curved posteriorly; posterior margin of juxta
appears evenly and gently convex.

DISTRIBUTION. Mexico (known only from the Sierra de Juarez, Oaxaca).

DISCUSSION. C. chinantlensis is compared by de la Maza with Eresia phillyra, and particularly
with C. eranites (for the Mexican population of which de la Maza retains the subspecies
mejicana). On account of the close similarity to C. eranites, chinantlensis is transferred here to
Castilia.

JAN A TELLA gen. n.

Type-species: Eresia leucodesma Felder & Felder. Gender: feminine.

Rather small butterflies, forewing outer margin straight or slightly convex, never excavate, uppersides black
with white markings; forewing with costal mark beyond cell and large white mark on centre of inner
margin; hindwing upperside discal band prominent; underside dark ocellar postdiscal spots well defined
in s3, s4 and s5. Sexes similar.

Genitalia. J in dorsal view with tegumen elongate, not tapered, scaphial extension short, posterior angles
well defined, each bearing 3-5 teeth or small hooks (Fig. 467); saccus single (Ungulate) with shallow apical
notch in J. hera, posterior border of juxta gently convex, penis slender, morula not prominent, ostium-keel
very small (possibly absent in the type-species). 9 ductus not chitinised, bursal support attached to a low
elevation, ribs little developed.

DISTRIBUTION. Nicaragua, Panama, northern South America, St. Vincent and Trinidad.

DISCUSSION. The three species C. leucodesma, C. hera and C. fellula were united by Forbes
(1945: 189) as his group 6 (Eresia). The facies of all are very different from other species in the
Eresia series, and by the shape of the dorsal structures of the male genitalia, appear allied more
closely to Phyciodes. All are clearly related and they form a small group most suitably placed
in a separate genus.

158 L. G. HIGGINS

Key to species of Janatella (based on Forbes, 1945: 161)

1 Base of forewing beneath light buff, ground colour of hindwing white, except towards the margin 2
Base of forewing underside ash grey, ground colour of hindwing dull grey; forewing rarely,

if ever, with white in cell fellula (p. 159)

2 White the dominant colour, extending over more than half of hindwing, and nearly half of

forewing (Colombia to Trinidad) leucodesma (p. 158)

- White markings limited to spots on forewings and a median band on the hindwings (Guianas)

hem (p. 158)

Janatella leucodesma (Felder & Felder) comb, n., stat. rev.

(Figs 167, 467-470)

Eresia leucodesma Felder & Felder, 1861: 103; 1867: 394, pi. 50, figs 11, 12. LECTOTYPE 9,

VENEZUELA: Caracas Province, Moritz (BMNH, specimen ex Felders' Coll., ex Rothschild Coll.), here

designated [examined].
Eresia cincta Edwards, 1864: 502. ^ syntypes, U.S.A.: Texas, Florida' [C. America?] (lost) (see Brown,

1966: 428).
Phyciodes leucodesma (Felder & Felder) Godman & Salvin, 1882: 197; Staudinger, 1885: 92, pi. 36;

Rober, 1913: 444, pi. 90, row f [figs 1, 2]; Hall, 1929: 132.
Phyciodes (Eresia) ianthe leucodesma (Felder & Felder); Forbes, 1945: 161, 189.
Anthanassa cincta Edwards; Holland, 1947: 141. [Holland indicates that this taxon is probably identical

with leucodesma.]
Anthanassa (?) leucodesma (Felder & Felder); Holland, 1947: 141, pi. 73, fig. 7.

c? forewing 16 mm, upperside wing-base and costa black to postdiscal costal bar, a large white area
extends from costal margin to meet costal mark ; hindwing upperside white except extreme base and black
marginal border. 9 similar, often slightly larger.

Genitalia. <$ in dorsal view, posterior angles of long, scaphial extension each bears 5 small teeth curved
medially, saccus single, valve apex with 2 terminal teeth (slide no. 1349). 9 genitalia, bursal duct
incompletely chitinised.

DISTRIBUTION. Trinidad, Island of St. Vincent, Panama, Nicaragua, Colombia, Venezuela.

DISCUSSION. In spite of the wide distribution, the species is remarkably stable in external
characters and genital structure.

Janatella hera (Cramer) comb. n.
(Figs 168, 471, 472)

Papilio hera Cramer, [1779]: 108, pi. 253, figs F, G. Syntype(s), [SOUTH AMERICA] ('Sierra Leona')

(depository unknown).
Papilio ianthe Fabricius, 1781: 80. Syntype(s), FRENCH GUIANA: Cayenne ('Mus Dom Yates', presumably

lost, illustrated in 'Jones Icones', Oxford, not listed by Zimsen). [Junior homonym of Papilio ianthe

Pallas, 1771.]

Phyciodes hera (Cramer) Kirby, 1871: 175; Rober, 1913: 443, pi. 90, row e [figs 2, 3].
Phyciodes ianthe (Fabricius) Hall, 1929: 133.
Phyciodes (Eresia) ianthe ianthe (Fabricius); Forbes, 1945: 161, 189.

c? forewing 20 mm, upperside markings like Castilia ofella, but outer margin of wing not excavate, hindwing
upperside white submarginal spot absent (constantly present in C. ofella). § similar.

Genitalia. 3 like J. fellula, dorsal structures elongate, in dorsal view scaphial angles rather densely
chitinised with 3-4 large teeth, saccus apex weakly notched (slide no. 830). 9 not examined.

DISTRIBUTION. French Guiana, Surinam.

NOTE. The excellent figure in Jones's unpublished Icones (Hope Department of Zoology, Oxford)
shows ianthe Fabricius to be identical with hera Cramer.

REVISION OF PHYCIODES 159

Janatella fellula (Schaus) comb. n.
(Figs 169, 473, 474)

Phyciodes fellula Schaus, 1902: 393. Holotype <J, COLOMBIA (USNM; Type no. 5885) [examined].
[Phyciodes abas (Hewitson); Rober, 1913: 443, pi. 90, row e [fig. 6] (J); Hall, 1929: 115; Forbes, 1945: 161,
189. Misidentifications.]

e? forewing 17-19 mm, variable, upperside black, markings white, cell-spot very small, larger spots in s2 s3
and s4 in oblique series; hindwing upperside white transverse band 2.0-2.5 mm, extends from s7 to inner
margin, 9 similar.

Genitalia. 3, in dorsal view, tegumen wide, elongate, well chitinised, scaphial terminal angles armed with
3 or 4 teeth, posterior border of juxta gently convex, saccus variable, wide, tapering rapidly, or longer and
more uniform in width in some specimens (slide no. 101). 9 genitalia like J. leucodesma, ductus partly
chitinised with goblet-shaped bursal support.

DISTRIBUTION. Ecuador, Colombia.

NOTE. The wing markings of this species are very like those of Telenassa abas, in which the
white transverse band on the hindwing upperside extends only from s6 to sib, i.e. it does not
quite reach to the inner margin.

MAZIA gen. n.

Type-species: Melitaea amazonica Bates. Gender: feminine.

In both sexes wings broad, appearing large in comparison with the thorax and abdomen; upperside
ground-colour fulvous with delicate black markings, arranged in series parallel with the wing-margins.
Antennae less than half the length of forewing, club slender, cylindrical (Fig. 491). Palpi elongate,
slightly ascending, terminal segment acicular.

Venation. Forewing cell closed, vll arising beyond cell-end from stalk of v7-10; hindwing cell open.

Genitalia. J in dorsal view, tegumen very short, a bunch of strong spines arising at each posterior angle
of the scaphial extension, valve in side view wide, posterior process short with 2 terminal teeth, posterior
margin of juxta gently convex; saccus elongate, apex notched; penis straight, morula inconspicuous.
9 genitalia, ductus bursae appears to be firmly chitinised as a saccular structure with bursa attached by
a membranous collar, unlike the arrangement present in all other genera.

DISCUSSION. The single species, M. amazonica, has strongly divergent characters, not only in the
structure of antennae, palpi and in venation, but also the wing-markings, which show a pattern
dissimilar to that of any other known species. Forbes (1945) placed amazonica as the sole species
in his Phyciodes (Eresia) group 7. The distribution is confined to the Amazon Region, but
quite extensive, reaching the eastern areas of Ecuador and Peru.

Mazia amazonica (Bates) comb. n.
(Figs 170, 475-479)

Melitaea amazonica Bates, 18646: 190. LECTOTYPE rf, BRAZIL: Tapajos (H. W. Bates) (BMNH, Type

no. Rh. 8522; Gabriel, 1927: 11), here designated [examined].

Phyciodes amazonica (Bates) Staudinger, 1885: 91, pi. 36 (9); Rober, 1913: 435, pi. 89, row b [figs 8, 9]
(5); Hall, 1929: 62, pi. 3, fig. 3 (genit.).
Phyciodes (Eresia) amazonica (Bates); Forbes, 1945: 161, 189.

3 upperside bright fulvous, markings black, striae arranged partly in series parallel with the black outer
margins of the wings, hindwing with a row of postdiscal dots; underside yellowish, markings as on
upperside but grey, series more complete. 9 similar.
Genitalia. As described above.

DISTRIBUTION. Brazil (including Mato Grosso), E. Peru, Ecuador.

160 L. G. HIGGINS

Species incertae sedis

Phyciodes melini Bryk

Phyciodes melini Bryk, 1953: 88. Holotype 9, PERU: Roque, 6.iv.l925 (NR, Stockholm)

Phyciodes metharmeoides Fassl
Phyciodes metharmeoides Fassl, 1922: 38. Holotype ?, BRAZIL: 'Innern von Teffe' (depository unknown).

Phyciodes rima Hall

Phyciodes rima Hall, 1929: 79. Holotype^, SURINAM ('interior') (C. W. Ellacombe) (BMNH) [examined].
Phyciodes (Tritanassa) rima Hall; Forbes, 1945: 190.

This taxon is based on a single specimen, now lacking the abdomen. It probably belongs to
the genus Telenassa.

Phyciodes mirabilis Hayward

Phyciodes mirabilis Hayward, 1967: 13-15, fig. Holotype ?, ECUADOR: Puyo, ix.1960 (J. Foerster) (IML,
Tucuman).

Phyciodes eucrasia Zikan

Phyciodes eucrasia Zikan, 1937: 385. !<?, 2 9 syntypes, BRAZIL: Sao Gabriel, Rio Negro, viii, is, x (MNRJ,
Rio de Janeiro).

Phyciodes chinchipensis Hayward

Phyciodes chinchipensis Hayward, 1964a: 63, fig. Holotype^, PERU: Rio Chinchipe, afluente norteno del
Rio Maranon, 500-700 m (W. Weyrauch) (IML, Tucuman).

This species is not represented in the BMNH collection. It is a small butterfly, J wing
expanse 20-22 mm, $ 23-24 mm. On the upperside the ground colour is yellow, markings black,
with a wide postdiscal area of the forewing with small yellow spots and markings; on the
hindwing upperside the postdiscal series of spots is rather prominent, as seen from the original
figure. Hayward considered P. chinchipensis to be closely related to Phystis simois. The genitalia
have not been figured.

Supplement on certain genera of Melitaeini

The genus Gnathotriche, and the closely related genus Gnathotrusia, were not included in my
descriptive account of the American Chlosyne and related genera (Higgins, 1960). This omission
is corrected here, and the butterflies described in similar style and detail. Most of the species
were included by Hall (19286-1930) in Phyciodes, but Forbes (1945) corrected this error,
grouping them in Gnathotriche. The four species involved, together with the small genus
Higginsius, form a group of special interest, divergent from all other American Melitaeini,
although the usual tribal characters are present. Outstanding group characters include the
unusual venation of the forewing and the structure of the male genitalia, which are generally
elongate, the posterior section of the valve very slender, curved and tapering to a pointed apex.
The species all occur in Colombia, Venezuela and N. Peru. In Gnathotrusia, wing-markings in
the male are mimetic and without any value to show relationships, but the genitalia structure
shows, as does that of Gnathotriche, an astonishing similarity in general structure, and in several
details, to the genitalia of Didymaeformia didyma and its allies, with extensive Palaearctic
distributions. The similarity includes the shape of the tegumen, the presence of brachia
(sub-unci), and the shape of the valve and harpe. Within the subfamily these features are so
unusual that a real relationship must be accepted to exist between these species and the
didyma-group, which suggests very wide dispersal of the didyma precursors at some distant epoch.

GNATHOTRICHE Felder & Felder
Gnathotriche Felder & Felder, 1862ft: 420, nota. Type-species: Euterpe exclamationis Kollar, by monotypy.

Rather small or medium-sized butterflies, forewing outer margin excavate below v5; palpi ascending,
middle segment slightly inflated and with dense long black hairs, terminal segment acicular. Upperside <$
markings dark brown with white or pale yellow spots. The females differ.

REVISION OF PHYCIODES 161

Venation. Forewing vlO arises near vll and before cell-end, branching to v9 running to wing-apex, v8
running to outer margin with v7 branching at about one-third of its length; beyond vlO the subcostal
continues as first discoidal vein with branch as v6, then continues as middle discoidal vein with branch as
v5, and then continues as lower discoidal vein to meet the median vein at v4. Hindwing venation
normal, with cell open.

Genitalia.d, in dorsal view, tegumen weakly chitinised, narrow, tapering, terminal section membranous,
posterior border of juxta conical (exclamationis) or rounded (sodalis), lingulae densely chitinised, saccus
cleft, valve elongate, the posterior section slender, strongly curved inwards and tapering gradually to a
single point, the saccular element well defined with a small harpe directed forwards; penis slender,
slightly longer than saccus plus valve, dilated beyond manica, then tapering to a gaping ostium, lacking
ostium crest and morula.

DISTRIBUTION. Colombia, Venezuela.

The species are said to inhabit forests in hilly and mountainous country; they do not occur
at high altitudes.

Gnathotriche exclamationis (Kollar)
(Figs 480-483)

Euterpe exclamationis, [1849]: 359, pi. 45, figs 5, 6.<3 syntype(s), VENEZUELA (NM, Vienna).
Gnathotriche exclamationis (Kollar) Felder & Felder, 18626: 420, nota; Staudinger, [1885]: 95, pi. 36;

Rober, 1914: 454, pi. 88, row g [figs 2-5] (rf, 9); Forbes, 1945: 190; 1946: 196, fig. 1 (genit.).
Eresia elaea Hewitson, 1869a: 25 [index]; [1872]: [29], pi. [15], figs 81, 82. Lectotype 9, ECUADOR: Rio

Verde (Buckley] (BMNH, Type no. Rh. 8581; Gabriel, 1927: 43), here designated [examined].
Eresia eleates Weymer, 1907: 18, pi. 2, fig. 5. Holotype 9, COLOMBIA, 'coll. Fruhstorfer' (depository

uncertain).
Phyciodes vanessoides Rober, 1914: 449, pi. 91, row b [fig. 5]. 9 syntype(s), COLOMBIA: Bogota (NMHU,

Berlin).

Phyciodes eleates (Weymer) Rober, 1914: 449, pi. 91, row a [figs 6, 7].
Gnathotriche exclamationis f. eresia Rober, 1914: 454, pi. 88, row g [fig. 6]. 9 syntypes, COLOMBIA and

VENEZUELA (MNHU, Berlin).

Gnathotriche exclamationis 9 f. eresia Rober, 1946: 196.
Gnathotriche exclamationis 9 f- eleates (Weymer); Forbes, 1946: 196.
Gnathotriche exclamationis 9 f. elaea (Hewitson); Forbes, 1946: 196.
Gnathotriche exclamationis 9 f. vanessoides (Rober); Forbes, 1946: 196.

c? forewing 19 mm, apex truncate, outer margin excavate, upperside black, markings cream-white, a short
basal streak in cell and a spot at cell-end, postdiscal costal bar of 3 elongate spots, and other postdiscal and
submarginal spots ; hindwing upperside a wide transverse band from costa to inner margin, crossed by dark
veins. 9 slightly larger, variable and differs greatly. In one form similar to^ but all markings orange-fulvous;
more often wing-apex rounded, outer margin of forewing less excavate, upperside black with an irregular
discal cream-white bar on forewing upperside, designated 'typical' by Rober. In a third female form
(f. eresia Rober), the upperside is black, except a wide fulvous area on the forewing extending from the
base along inner margin and costa about half-way to apex.

Genitalia.3, in dorsal view, posterior border of juxta conical; harpe of valve well developed. 9 ostium
bursae open, leading directly to bursal duct, bacilli weakly chitinised, bifid, but accompanied by numerous
membranous fibrils to compose a single united rod, scutum not large.

DISTRIBUTION. Colombia, Venezuela.

DISCUSSION. In females the variation is very great and specific attribution is very largely a
matter of guesswork.

Gnathotriche sodalis Staudinger
(Figs 484-486)

Gnathotriche sodalis Staudinger, [1885]: 95; Rober, 1914: 454, pi. 88, row g [figs 7, 8] (^); Forbes, 1945:
190; 1946: 196. 3 syntypes, COLOMBIA: Cauca Valley (MNHU, Berlin).

162 L. G. HIGGINS

J forewing 20 mm, outer margin deeply excavate; ground-colour brown, markings yellow, rather obscured
by brown scales, spot in mid-cell and at cell-end, discal spots in sib, and at bases of s2 and s3, costal
bar of 4 or 5 short stripes, submarginal series of spots often incomplete, .but a white spot in s6 constant;
hindwing upperside with 4 or 5 small discal spots forming a broken transverse band not suffused brown,
postdiscal and submarginal markings vestigial. 9 not known.

Genitalia.J densely chitinised, in dorsal view tegumen fragile, easily distorted, posterior border of juxta
semicircular, prominent, harpe very small; penis slender, as long as valve, ostium crest prominent
(?everted), morula absent.

DISTRIBUTION. Colombia (Cauca Valley).

NOTE. G. sodalis appears to be a very local and rare species. Seven males in the BMNH are
all from Staudinger, ex Godman & Salvin, clearly part of the original type-material. The species
is not represented in Hall's collection.

GNATHOTRUSIA gen. n.

Type-species: Eresia mundina Druce. Gender: feminine.

Butterflies of medium size, palpi like Gnathotriche, hairy, terminal segment acicular; forewing apex
rounded, outer margin straight. Upperside markings mimetic of Actinote species, polymorphic, underside
like Gnathotriche, apex of forewing and all hind wing greyish, lined black along and between veins.

Venation. Like Gnathotriche but in forewing v5 arises close to v6 so that middle discocellular vein is
shortened, but remains easily identifiable.

Genitalia. Male like Gnathotriche but differ in the densely chitinised conical tegumen, produced posteriorly
to terminate abruptly ; differs also in presence of prominent, cylindrical brachia, strongly chitinised, their
medial borders armed with fine spines; saccus cleft.

DISTRIBUTION. Restricted to Colombia and N. Peru.

Gnathotrusia mundina (Druce) comb. n.
(Fig. 487)

Eresia mundina Druce, 1876: 221, pi. 18, fig. 4. Holotype^, PERU: Huiro, Santana Valley (H. Whitely)

(BMNH, Type no. Rh. 8570; Gabriel, 1927: 83), [examined].

Phyciodes fallax Staudinger, 1885: 94. 1 3, 3 ? syntypes, PERU: Chanchamayo (MNHU, Berlin).
Phyciodes rosina Dognin, 1888: 48, fig.; Rober, 1914: 449. Lectotype <^, ECUADOR: 'Numbala River'

('environs de Loja' on specimen) (BMNH) here designated [examined]. [Specimen bears Dognin's M/S

'type' label.]

Eresia crina Schaus, 1902: 392. Holotype, ECUADOR (USNM, Type no. 5884) [examined].
[Eresia acraea Hopffer; Rober, 1913: pi. 92, row f [fig. 3]. Misidentification.]
Phyciodes erebia Rober, 1914: 449, pi. 91, row b [fig. 6]. LECTOTYPE, PERU: Cushi, Huanuco province,

1900 m, ( W. Hoffmans) (BMNH) here designated [examined]. [Specimen bears Rober's M/S determination

label.]
Phyciodes hopfferi Rober, 1914: 449; pi. 92, row f [fig. 3] (as Eresia acraea). Syntype(s), PERU (depository

unknown).

Phyciodes crina (Schaus) Rober, 1913: 448.
Phyciodes mundina (Druce) Rober, 1914: 450.

Phyciodes mundina peraea Hall, 1928a: 13. Holotype c?, ECUADOR (BMNH) [examined].
Phyciodes mundina f. callianira Hall, 1930: 200. Holotype c?, PERU (BM, Brighton) [examined].
Phyciodes mundina mundina (Druce); Hall, 1930: 198.
Phyciodes mundina f. hopfferi Rober, Hall, 1930: 199.
Phyciodes mundina L fallax Staudinger; Hall, 1930: 199.
Phyciodes mundina f. rosina Dognin; Hall, 1930: 200.
Phyciodes mundina f. crina (Schaus); Hall, 1930: 201.
Phyciodes mundina f. erebia Rober; Hall, 1930: 201.
Phyciodes mundina f. peraea Hall; Hall, 1930: 201, pi. 2, fig. 7 (^).
Phyciodes mundina f. testacea Hall, 1935: 221. LECTOTYPE J, BOLIVIA: Coroico (BMNH), here

designated [examined: Hall stated 'type' to be from Coroico, in BM; I have labelled middle specimen

of 3 from Coroico as lectotype, no specimens having been labelled by Hall].

REVISION OF PHYCIODES 163

Gnathotriche mundina (Druce); Forbes, 1945: 141-5, 190.

? Phyciodes erebia mima Bryk, 1953: 90. HolotypeJ, PERU: Roque (NR, Stockholm).

<$ forewing 22-23 mm, upperside black, polymorphic, markings most variable, orange-fulvous, rosy red,
pink etc.; in nominate form forewing a wide orange-red postdiscal bar extends from costa to s3; hindwing
upperside gleaming blue-black, underside, all veins lined sooty-black ; in f. hopfferi, upperside forewing basal
area orange-fulvous, hindwing with basal area of same colour; in f. peraea, forewing upperside postdiscal
band narrow, buff, running to sib, basal colour absent from both wings; f. rosina is like hopfferi,
but with upperside markings rose-red. These and other variants occur at random in individual specimens,
and usually an obvious model in Actinote can be recognised.

Genitalia. 3 in dorsal view like Gnathotriche exclamations, but in dorsal view tegumen large, an
elongate conical shape; brachia prominent.

DISTRIBUTION. Restricted to Colombia and N. Peru.

Gnathotrusia epione (Godman & Salvin) comb. n.
(Figs 488, 489)

Eresia epione Godman & Salvin, 18786: 263. Lectotype^, COLOMBIA: Antioquia (Salmon) (BMNH, Type

no. Rh. 8571; Gabriel, 1927: 46), here designated [examined].
Phyciodes epione (Godman & Salvin) Staudinger, 1885: 94; Rober, 1914: 449, pi. 91, row b [fig. 4]

(as Eresia).

Phyciodes styx Staudinger, 1885: 94. 10 syntypes, COLOMBIA: Antioquia (MNHU, Berlin).
Eresia callianthina Hall, 1921: 279. 2 [9] syntypes, COLOMBIA: Santa Elena, 2900 m (BM, Brighton).
Phyciodes epione elenae Hall, 1928: 13. Holotype J, COLOMBIA: Santa Elena, 2900 m (BM, Brighton)

[examined].

Phyciodes epione epione (Godman & Salvin); Hall, 1930: 202; Forbes, 1945: 190.
Phyciodes epione f. styx Staudinger; Hall, 1930: 203.
Phyciodes epione elenae Hall; Hall, 1930: 203, pi. 2, fig. 8 (^).
Phyciodes callianthina (Hall) Hall, 1930: 203, pi. 2, fig. 9 (9).
Phyciodes epione callianthina Hall; Forbes, 1945: 190.

cJ forewing 23 mm, upperside both wings gleaming metallic deep blue, shading to black along forewing
costa and outer margins; hindwing inner margin bright red; underside black, apex of forewing and all
hindwing shading to greyish, with black lines along and between veins. Wing-markings variable, in f. styx
hindwing upperside inner margin black; in f. elenae similar with forewing underside with red basal area;
f. callianthina has forewing upperside base red, underside red area extensive. 9 illustrated by Hall (1930:
pl. 2, fig. 9).

Genitalia. J like G. mundina, in dorsal view elongate, tegumen firmly chitinised, domed with a narrow
central extension ending abruptly, brachia stout, juxta wide, tapering to a rounded apex, lingulae very dense.

DISTRIBUTION. Restricted to Colombia.

HIGGINSIUS Hemming

Fulvia Higgins, [1959]: 162. Type-species: Melitaeafasciata Hopffer, by original designation. [Homonym

of Fulvia Gray, 1853, and Fulvia Adams, 1957.]
Higginsius Hemming, 1964: 139. [Replacement name for Fulvia Higgins.]

The genus is defined, the species described, and anatomical characters figured in Higgins (1960: 460-462).
Important characters include forewing venation as in Gnathotriche and in hindwing v7 branches from
subcostal vein some distance before the fork with v5 and v6; palpi conspicuous, long and very hairy.
In the minute genitalia the tegumen is narrow, the apical section of the valve curved inwards, very
slender and sharply pointed, the saccus small with shallow terminal notch.

DISTRIBUTION AND RELATIONSHIPS. The genus Higginsius includes two species (H. fasciatus,
H. miriam) with restricted distributions in Ecuador, W. Colombia and probably in Venezuela.
The curious wing-venation, the long, densely hairy palpi and the genitalic characters, with the

164 L. G. HIGGINS

slender, pointed valve apex, all point to close relationship with Gnathotriche, in spite of the very
different fades. The species are not mimetic or cryptic, and in fasciata the upperside wing-
markings show the common ancestral pattern of black striae on a fulvous ground. Both
species appear to be very local and rare. Females are unknown.

ANTILLEA Higgins

Antillea Higgins, [1959]: 164; Riley, 1975: 78-79, pi. 6. Type-species: Papilio pelops Drury, by original
designation.

The genus is defined, the species are described and anatomical characters are figured in Higgins (1960:

462-465).

Important generic characters include forewing venation as in Gnathotriche. In the hindwing veins 5, 6 and
7 all arise from a common point on the subcostal vein; veins la and Ib are slightly displaced to make
room for a hair-pencil. The minute genitalia are unusual, the tegumen very shallow, the valve in side
view blunt, the harpe relatively large, and with a prominent saccular tooth near the blunt apex. The penis
massive, with ostium-crest and vestigial morula. 9 (a single specimen examined) appears to have a short,
double bacillus.

In spite of the apparently primitive venation of the forewing, this genus does not agree well
with the genus Didymaeformia. With several unusual and unique characters, the two species
(pelops, proclea) are probably relics of great antiquity, and, without obvious relationship to any
other extant group, difficult to place today. The $ examined cannot belong to the Phyciodes
section, and I have included Antillea within the Melitaeini as an independent genus.

Review of the classification of the Melitaeinae

The publication of this account of the tribe Phyciodini, and of the genera Gnathotriche and
Gnathotrusia, completes the study of the Melitaeinae, which was begun in the Illustrated
Catalogue of the Palaearctic Melitaea, published by the Royal Entomological Society of London
in 1941. This is, therefore, a suitable opportunity to review the conclusions reached then and in
subsequent papers, and to examine the classification of the whole subfamily. In considering
the relationships of the three principal tribes, I have commented upon species living in restricted
habitats, divergent species, and especially upon such interesting features of distribution as specific
and generic disjunctions. It is necessary to include a formal description of the subfamily, and a
catalogue at tribal and generic levels. Roger Verity, who was the first entomologist to introduce
subfamily status for the Melitaeinae, insisted that the large genus Melitaea required further
division, and I agree with his suggestions. To make a more meaningful arrangement it is
necessary to divide Melitaea into four generic groups. For one of these, the name Mellicta is now
generally recognised; for the two remaining groups, suitable generic names are available, and
these are now resurrected. A new genus, for which the name Gnathotrusia has been proposed
above, is required for two species associated with the divergent Gnathotriche group of South
America. No other taxonomic changes are suggested.

On the basis of the characters of the male and female genitalia, it is easy to recognise three
major groups of tribal rank, and possibly a fourth tribe is required for the small genus
Atlantea, the three species of which appear divergent in relation to all other genera. All species
show the key family and subfamily characters, but at tribal level each group is entirely
isolated. Species with intermediate or equivocal characters have not been seen. The tribes can
be defined precisely as follows.

1 Male genitalia, saccus absent . . . EUPHYDRYINI
Male genitalia, saccus present ... 2

2 Female genitalia with bacillus . . . MELITAEINI
Female genitalia without bacillus . . 3

3 Female genitalia with bursal support . PHYCIODINI
Female genitalia without bursal support . ? tribe

REVISION OF PHYCIODES 165

Check-list of tribes, generic groups, genera and species of the Melitaeinae

The type-species of genera are marked with an asterisk; junior synonyms are not included unless they are
type-species.

EUPHYDRYINI Higgins (Holarctic)
EUPHYDRYAS Scudder

*phaeton (Drury)
HYPODRYAS Higgins

*maturna (L.)

intermedia (Menetries)

cynthia (Denis & Schiffermuller)

iiluna (Dalman)

gillettii (Barnes)
OCC/DRYAS Higgins

*anicia (Doubleday)

chalcedona (Doubleday)

editha (Boisduval)

colon (Edwards)
EURODRYAS Higgins

*aurinia (Rottemburg)

alexandrina (Staudinger)

desfontainii (Boisduval)

orientalis (Herrich-Schaffer)

MELITAEINI Tutt (Holarctic and Neotropical)
MELLICTA-group (Palaearctic)
MELLICTA Billberg

*athalia athalia (Rottemburg)

athalia celadussa (Fruhstorfer)

athalia ambigua (Menetries)

deione (Geyer)

parthenoides (Keferstein)

aurelia (Nickerl)

britomartis (Assmann)

varia (Meyer-Dur)

asteria (Freyer)

centralasiae (Wnukowsky)

rebeli (Wnukowsky)

menetriesi (Caradja)

plotina (Bremer)

alatauica (Staudinger)

DIDYMAEFORMIA-group1 (Palaearctic except Poladryas)
MEL1TAEA Fabricius

avinovi Sheljuzhko

arduinna (Esper)

*cinxia (L.)

halha Evans

minerva Staudinger

pallas Staudinger

turanica ErschofT

agar Oberthiir

asteroidea Staudinger

di ami nu diamina (Lang)
diamina regama Fruhstorfer

1 Verity's genus Didymaeformia has not been accepted by most authors, partly because the designation of a type-species
is somewhat ambiguous in the author's original introduction of the name (1950). It is, however, inconvenient to be
without a generic name for Papilio didyma Esper and its many relatives in Central Asia. Verity's intention was perfectly
clear and Didymaeformia Verity, 1950, is accepted here as a valid generic name, with type-species Papilio didvma Esper, as
recorded by Hemming (1967).
(Note. The name is spelt Didymaeformis by Hemming, clearly a lapsus calami\)

166 L. G. HIGGINS

lukto Evans

mimetica Higgins

tangigharuensis de Freina

romanovi Grum-Grshimailo

arcesia Bremer

balbita Moore

sindura Moore

amoenula Felder & Felder
jezabel Oberthiir

bellona Leech
POLADRYAS Bauer

miniita (Edwards)

*pola (Boisduval)

arachne (Edwards)
D/DYMAEFORMIA Verity
didyma-group

*didyma (Esper)

deserticola (Oberthiir) comb. n.

transcaucasica (Turati) comb. n.

persea (Kollar) comb. n.

afghana (Heydemann) comb. n.

abyssinica (Oberthiir) comb. n.

gina (Higgins) comb. n.

mixta (Evans) comb. n.

casta (Kollar) comb. n.

yuenty (Oberthiir) comb. n.

saxatilis (Christoph) comb. n.

didymina (Staudinger) comb. n.

didymoides (Eversmann) comb. n.

ala (Staudinger) comb. n.

pseudoala (Sheljuzhko) comb. n.

chitralensis (Moore) comb. n.

sutschana (Staudinger) comb. n.

acraeina (Staudinger) comb. n.
fergana-group

fergana (Staudinger) comb. n.

ambrisia (Higgins) comb. n.

lunulata (Staudinger) comb. n.

shandura (Evans) comb. n.

macarandica (Staudinger) comb. n.

athene (Staudinger) comb. n.

infernalis (Grum-Grshimailo) comb. n.
collina-group

collina (Lederer) comb. n.

turkmanica (Higgins) comb. n.

consulis (Wiltshire) comb. n.

kuchi (Wyatt) comb. n.

trivia (Denis & Schiffermiiller) comb. n.
C1NCL1DIA Hiibner

*phoebe (Denis & Schiffermiiller)

aetherie (Hiibner)

scotosia (Butler)

sibina (Alpheraky)

sarvistana (Wiltshire)

CHLOSYNE-group (Nearctic and Neotropical)
CHLOSYNE Butler

nycteis (Doubleday)

gorgone (Hiibner)

REVISION OF PHYCIODES 167

harrisii (Scudder)

hoffmanni (Behr)

malcolmi (Comstock)

gabbii (Behr)

damaetas (Skinner)

neumeogeni (Skinner)

acastus (Edwards)

palla (Boisduval)

definita (Aaron)

marina (Geyer)

melitaeoides (Felder & Felder)

gloriosa Bauer

*janais (Drury)

poecile (Felder & Felder)

erodyle (Bates)

melanarge (Bates)

lacinia lacinia (Geyer)

lacinia saundersi (Doubleday)

calif arnica (Wright)

riobalensis Bauer

ehrenbergi (Geyer)

narva (Fabricius)

gaudealis (Bates)

hippodrome (Geyer)

rosita Hall

marianna Rober
T HESS ALIA Scudder

*leanira (Felder & Felder)

theona (Menetries)

cynisca (Salvin)
TEXOLA Higgins

*elada (Hewitson)

anomalus (Godman & Salvin)

coracara (Dyar)
DYMASIA Higgins

*dymas (Edwards)
MIC ROT I A Bates

*elva elva Bates

elva draudti Rober

GNATHOTRlCHE-group (Neotropical — South American and Antillean)
GNATHOTRICHE Felder & Felder

*exclamationis (Kollar)

sodalis Staudinger
GNATHOTRUSIA gen. n.

*mundina (Druce)

epione (Godman & Salvin)
HIGGINSWS Hemming

*fasciatus (Hopffer)

miriam (Dognin)
ANTILLEA Higgins

*pelops (Drury)

proclea (Doubleday)

PHYCIODINI trib. n. (Nearctic and Neotropical)
PHYCIODES Hiibner
tharos (Drury)

*cocyta (Cramer)
batesii (Reakirt)
campestris campestris (Behr)

168 L. G. HIGGINS

campestris camillus Edwards

montanus (Behr)

mylittus mylittus (Edwards)

mylittus mexicanus Hall

mylittus thebais Godman & Salvin

pallidus (Edwards)

orseis Edwards

herlani Bauer

pictus pictus (Edwards)

pictus pallescens (Felder)

phaon (Edwards)

vesta vesta (Edwards)

vesta graphica (Felder)
PHYSTIS gen. n.

*simois simois (Hewitson)

simois variegata (Rober)

? chlnchipensis (Hayward)
ANTHAHASSA Scudder

drusilla drusilla (Felder & Felder)

drusilla lelex (Bates)

drusilla alceta (Hewitson)

drusilla verena (Hewitson)

ptolyca ptolyca (Bates)

ptolyca amator (Hall)

ardys ardys (Hewitson)

ardys subota (Godman & Salvin)

dracaena (Felder & Felder)

phlegias (Godman & Salvin)

texana texana (Edwards)
*cincta Edwards; Scudder

texana seminole (Skinner)

alexon alexon (Godman & Salvin)

alexon subconcolor (Rober)

acesas (Hewitson)

nebulosa (Godman & Salvin)

argentea (Godman & Salvin)

atronia (Bates)

otanes otanes (Hewitson)

otanes sopolis (Godman & Salvin)

annulata sp. n.

crithona (Salvin)

fulviplaga (Butler)

hermas (Hewitson)

frisia (Poey)

tulcis (Bates)

ilubia (Hall)

taeniata (Rober)

sosis (Godman & Salvin)

drymaea (Godman & Salvin)

shakes (Godman & Salvin)

cortes (Hall)
DAGON gen. n.

pusillus (Salvin)

*catula (Hopffer)

morenus (Rober)

fontus (Hall)
TELENASSA gen. n.

*teletusa (Godart)

burchelli (Moulton)

REVISION OF PHYCIODES 169

berenice (Felder & Felder)

signal a (Hall)

abas (Hewitson)

jana (Felder & Felder)

elaphina (Rober)

nana (Druce)

nussia (Druce)

notus (Hall)

gaujoni (Dognin)

trimaculata (Hewitson)

flavocincta (Dognin)

catenaria (Godman & Salvin)

delphia (Felder & Felder)

sepulta (Hall)
ORTILIA gen. n.

*liriope (Cramer)

gentina sp. n.

orthia (Hewitson)

orticas (Schaus)

sejona (Schaus)

velica velica (Hewitson)

velica durnfordi (Godman & Salvin)

zamora (Hall)

dicoma (Hewitson)

polinella (Hall)

ithra (Kirby)
T1SONA gen. n.

* saladillensis saladillensis (Giacomelli)

saladillensis clarior subsp. n.
TEGOSA gen. n.

*claudina (Eschscholtz)

similis nom. n.

orobia (Hewitson)

fragilis (Bates)

infrequens sp. n.

ursula (Staudinger)

flavida (Hewitson)

tissoides (Hall)

pastazena (Bates)

guatemalena (Bates)

anieta anieta (Hewitson)

anieta cluvia (Godman & Salvin)

anieta luka subsp. n.

anieta serpia subsp. n.

nazaria (Felder & Felder)

etia (Hewitson)

nigrella (Bates)
ERESIA Boisduval

clara Bates

nauplius nauplius (Linnaeus)

nauplius extensa (Hall)

plagiata (Rober)

letitia letitia Hewitson

letitia ocellata (Rober)

lansdorfi (Godart)

sestia Hewitson

coela Druce

oblita (Staudinger)

carme carme Doubleday

170 L. G. HIGGINS

carme /a/as Godman & Salvin

polina Hewitson

alsina Hewitson

cissia (Hall)

eutropia Hewitson

mintas (Staudinger)

quintilla Hewitson

poecilina Bates

mclaina sp. n.

sticta Schaus

ithomioides ithomioides Hewitson

ithomioides pseudocelemina (Strand)

anomala sp. n.

nigripennis Salvin

emerantia Hewitson

moesta Salvin & Godman

phacdima Salvin & Godman

datis datis Hewitson

ilatis corybassa Hewitson

mat-yard ha Hewitson

*eunice eunice (Hiibner)

eunice olivencia Bates

eunice esora Hewitson

etesiae (Hall)

erysice (Geyer)

casaphia Hewitson

mechanitis Godman & Salvin

pelonia Hewitson

phillyra Hewitson

aveyrona aveyrona Bates

aveyrona mylitta Hewitson

perna Hewitson

levina Hewitson

actinote Salvin

selene (Rober)
CASTILIA gen. n.

*castilla (Felder & Felder)

accident alls (Fassl)

perilla (Hewitson)

neria (Hewitson)

nortbrundii (Weeks)

eranites (Hewitson)

fulgora (Godman & Salvin)

fausta (Godman & Salvin)

ofella (Hewitson)

myia (Hewitson)

griseobasalis (Rober)

angusta (Hewitson)

chinantlensis (de la Maza)
JAN AT ELL A gen. n.

*leucodesma (Felder & Felder)

hera (Cramer)

fellula (Schaus)
MAZIA gen. n.

*amazonica (Bates)

Tribe?

ATLANTEA Higgins

*perezi (Herrich-Schaffer)

REVISION OF PHYCIODES 171

pantoni (Kaye)
tulita (Dewitz)
cryptadia Sommer & Schwartz

Distribution

The following account is not intended to be exhaustive, but the principal features of distribution
of the Melitaeinae are described and some anomalous features are pointed out. Especially in the
Palaearctic Region, the situation is complicated, and a serious difficulty arises in the absence of
information and material from many parts of the U.S.S.R.

PALAEARCTIC REGION

The subfamily is represented by the two tribes Melitaeini and Euphydryini, with about 75 species
widely distributed, forming an important faunal element. Four regions in this vast area are
sufficiently distinct to need separate consideration.

European Region, including Europe with Russia to the Urals, NW. Africa and W. Asia (Pontic
distribution area)

There is an important concentration of 22 species in W. Europe and NW. Africa, involving two
faunal regions. Euro-Siberian elements are widespread in northern and central parts of the area,
including the arctic, and on all mountains. Six genera are represented : Melitaea (M. cinxia and
M. diamina); Cinclidia (C. phoebe) (see also Siberian Region below); Didymaeformia (D. didyma};
Mellicta, the dominant genus (M. athalia with two major subspecies; M. britomartis occurs from
Piedmont to Transbaical; two species, M. asteria and M. varia are endemic in the high Alps
and central Apennines, two species, M. parthenoides and M. aurelia, fly at moderate altitudes in
W. Europe, the latter rather widely distributed to Baltic countries and to the Caucasus);
Hypodryas (H. intermedia wolfensbergeri, H. maturna, H. iduna not south of 60°N lat., H. cynthia
only on the Alps and Balkan Mts. (also S. Caucasus?)); Eurodryas (E. aurinia).

Most of these species occur in suitable localities in S. Europe and in NW. Africa, with
two additional species, Eurodryas desfontainii and Cinclidia aetherie, their ranges restricted to
SW. Europe and Morocco, the latter also in Sicily. Although situated within the Mediterranean
area, the taxonomic characters of these two species are closely related to the Euro-Siberian fauna.
They are best placed as pre-glacial relicts (Atlanto-Mediterranean). E. desfontainii is closely
related to E. orientalis, locally endemic in W. Asia. The distribution of Cinclidia is extensive (see
below, E. Siberia).

There is an interesting situation with the two subspecies of M. athalia, identifiable by the
distinctive characters of the male genitalia. In W. Europe the distribution of the eastern taxon
M. a. athalia meets the western M. a. celadussa in central France. Here the two are known to
breed together along a frontier that extends from the Atlantic to the Adriatic, producing
individuals having male genitalia of intermediate character, forming a hybrid zone up to 40 miles
wide. I have no doubt that the western M. a. celadussa, with typical Atlanto-Mediterranean
distribution, is the original pre-glacial species, its range now reduced by the post-glacial invasion
of M. a. athalia, a Siberian (Angaraland) element.

Ponto-Mediterranean species in S. Europe are few in number, as follows : Melitaea arduinna
is local in the Balkans and in N. Greece; Didymaeformia trivia is widespread in SE. Europe, with
scattered colonies in N. and C. Italy, N. Spain and in Portugal. A single species occurs in
NW. Africa, D. deserticola in S. Algeria and Tunisia.

In Asia Minor and to the east and south, there is a vast area, partly semi-desert, where
the genus Didymaeformia is dominant at low and medium altitudes, often numerous in
individuals, but the number of species is not large and includes D. didyma and D. trivia, often
common in northern areas (see p. 166). D. deserticola macromaculata is local in Syria, Lebanon
and Israel. D. persea is widely distributed, with several local races and marked seasonal variation.
The range extends south to Oman in Arabia, D. abyssinical (D. a. scotti Higgins) occurring in
the mountains of Yemen and D. a. abyssinica, described from Ethiopia, but never recorded again
from that country. The mountain fauna is remarkable for presenting several small species of

172 L. G. HIGGINS

Melitaea, flying in rare widely distributed colonies, usually at rather high altitudes, as follows:
M. collina, Syria, Lebanon, Iraq (Kurdistan); M. turkmanica, Turkmen S.S.R. ('Askabad'),
Armenia ( = vedica}', M. consults, Iran, S. Zagros, perhaps a subspecies of M. turkmanica. These
three species are closely related. D. saxatilis, N. Iran (Mt. Demavend, Mt. Shahkuh, flying at
4,000 m). Cinclidia sarvistana, Iran, Zagros Mts at high altitudes (male genitalia very divergent).

Central Asia

In this region the Tian Shan, Pamirs and other great mountain ranges form a specialised
biotope with the greatest concentration of Melitaeini in the Palaearctic region. Didymaeformia
didyma and three related species fly in the western districts, but in general the western species
disappear and their places are taken by other taxa endemic in central Asia. Among the most
interesting is the D. fergana-group of seven species, most of which fly at altitudes of 2,500 m or
more, in oreo-tundral biotopes, including D. lunulata, D. shandura, D. ambrisia etc. All are rare,
their habitats strictly localised, but in all species the male genitalia are distinctive, with good
specific characters. These stand apart from the western D. didyma, as a distinct endemic group
within the genus. They are accompanied by four or five species of Melitaea, closely related to
M. cinxia, the uppersides bright fulvous with delicate black markings. All are distributed at high
altitudes on the Ala Tau, Changai, Pamirs etc., and extend into Afghanistan, each major mountain
group having its special phenotype, but with the male genitalia similar in all cases. Of these,
M. minerva is probably the most important, with a wide distribution in the Ala Tau, Pamirs, and
Hindu Kush, the different races flying at various altitudes up to 3,000 m or more, with a
confusing range of different phenotypes, variously graded by different authors as species or
subspecies. Cinclidia sibina is widely distributed in E. Turkestan, Alai Mts., etc. In many ways
Afghanistan is closely related to the Hindu Kush and Pamirs, with similar high mountains, but
the butterfly fauna differs considerably. Five species are known which appear to be endemic to
the area, all with restricted distributions as follows: Melitaea avinovi has been known for
many years; Didymaeformia kuchi is a small species that flies at high altitudes; D. afghana
data are scanty at present. In the southern districts, which lead into Baluchistan, Melitaea lukto
and M. mimetica occur, flying at moderate levels, perhaps subspecies of a single taxon.

The Himalayas and Mongolia

In the west these mountains merge into the central Asian Karakorums and Hindu Kush in
Ladak and Chitral. Two or three species of Didymaeformia occur here including D. chitralensis,
which flies at moderate levels in the valleys. The principal interest lies in the small Melitaea
amoenula (male forewing length 12 mm), which flies in Khema and Ladak at 3,500 m or more,
and the larger M. balbita which occurs at lower levels. These are the most westerly
representatives of a remarkable series of distinct but closely related races (? species) which
occurs through the length of the Himalayas to the exclusion of other Melitaeinae. The series
includes M. sindura in the north-west and M. sikkimensis in Sikkim and Nepal. Further east,
in south-western Tibet, the series merges into M. jezebel, slightly larger, with the very distinct
M. bellona at lower altitudes. In the Amdo region, the Himalayan races are represented by
M. arcesia which flies in suitable localities near the Kuku Nor and northwards to the Changai
Mts and Transbaicalia (Sajan and Kentei Mts). The entire series forms a 'Rassenkreis' which is
remarkable among Palaearctic butterflies. The male genitalia of the different races vary in size
but are similar in all other respects. Beyond India, the Chinese province of Yunnan in the
extreme east of the Palaearctic region supports two divergent species, M. yuenty and M. agar,
the latter extending into Tibet at high altitudes, with considerable racial variation. One other
species inhabits the north of this inhospitable region, M. romanovi, about which there is little
information.

Eastern Siberia, including N. China, Mongolia and N. Korea

Near Lake Baikal (100°E) the picture changes. While some of the western dominants are still
to be found, more often their eastern ranges are continued by subspecies or closely related
species across temperate Siberia, China and Mongolia to Korea and often to Japan. Six genera
are present: Melitaea, with one species, M. diamina protomedia, Amur etc.; Cinclidia, with two
species, C. scotosia, also in Mongolia and Japan, and C. phoebe whose range extends to N. China,

REVISION OF PHYCIODES 173

but probably it is not truly Siberian; Didymaeformia with two species, D. didymoides, Changai
Mts, Kansu, Korea etc., and D. sutschana, Amur, Korea; Mellicta with five species, M. athalia
ambigua, male genitalia with small subspecific characters, M. britomartis amurensis, Amur,
Sutschan etc., M. centralasiae, Shigansk, Kentei Mts, Chingan Mts, etc.; M. menetriesi, Amur
and Kamschatka, M. plotina, Ussuri, Amur (Bureja Mts), Transbaicalia (?). The known data
of these species are not satisfactory, and it is not possible at present to record their individual
ranges. Two other Mellicta species, M . alatauica and M. rebeli, from the Alatau and Altai Mts
respectively, really belong to this group; as they are represented by only a few specimens, their
true distribution is quite unknown. Hypodryas, with three species, H. intermedia konumensis
from Sutschan and Korea, larger than western specimens, H. iduna, recorded from the far north,
H . maturna recorded from S. and E. Siberia. Eurodryas, with one species, E. aurinia siberica from
Transbaical, Mongolia, Kansu, Korea etc., very variable.

The generic list in E. Siberia closely resembles that of W. Europe. The genus Cinclidia is
probably best regarded as Euro-Siberian. Five species are widely distributed in warm and
temperate areas of the Palaearctic, as follows. C. phoebe: S. and C. Europe and eastwards to
S. Siberia and N. China; C. aetherie: Morocco, Spain, Sicily; C. sibina: Turkestan, Thian Shan;
C. scotosia: E. Siberia, N. China, Korea, Japan; C. sarvistana: S. Iran (Zagros Mts), very
divergent. While the genus does not reach N. America, the Chlosyne series in the western
mountains, by habitus and wing markings, appears more closely related to Cinclidia than to
any other melitaeid genus.

NEARCTIC AND NEOTROPICAL REGIONS

There is no reason to suppose that the Eurasiatic continent has been other than a single united
land mass throughout the development and dispersal of the present Melitaeine fauna. It is
otherwise with the Americas. Across the Behring Straits North America has been connected
many times with Asia. On the other hand, it has been separated by marine barriers from
South America until junction occurred relatively recently in geological terms. The situation in
Central America in the distant past appears to be uncertain, but the present Melitaeid fauna is
distinctive. The Antillean archipelago, perhaps the remaining traces of a former continent, is
still unconnected with the mainland. These archaic isolations of two, or perhaps three large
continents, are reflected in the distribution and the character of the butterflies. With few
exceptions the Palaearctic Melitaeine genera are absent from the American continents. An
Exception is Hypodryas gillettii, which occurs in the mountains of Wyoming and Montana,
and is a close relative of the European H . maturna.

North America

The Euphydryini are well represented by four species of Occidryas in the western Mountains.
These show great differentiation compared with the Eurasiatic series. Hypodryas gillettii, referred
to above, is of great interest. It is the single representative within the subfamily of a truly
Palaearctic Melitaeine genus present in America. Euphydryas phaeton, with eastern distribution,
is not closely related to Occidryas. It is most divergent in wing markings, and the structure of
the male genitalia suggests relationship to the European Eurodryas aurinia. The tribes Melitaeini
and Phyciodini are both well represented, and again principally in the western areas and
mountains, each with a single genus, Chlosyne and Phyciodes respectively. Although belonging
to different tribes, the two genera have certain features in common in their American ranges. First,
the genitalic characters and wing markings in each genus are almost identical in all species, so that
generic identification is very easy but specific identification usually difficult. Secondly, all species
of both genera have the common basic Melitaeine pattern of upperside wing markings, with black
spots and striae on a fulvous ground. This pattern is rare further south, and almost non-existent
south of Mexico. The situation is difficult to explain, since in both cases these northern popula-
tions probably represent dispersals from the more primitive populations in Central America
(Chlosyne) and South America (Phyciodes). In both genera the almost complete lack of diversity in
genitalia and in wing markings does not suggest an ancient origin. In North America there
do not appear to be examples of the impressive specific disjunctions that are not uncommon
in Eurasia, nor are there any isolated relict forms among the Melitaeinae. On the other hand

174 L. G. HIGGINS

there are two unexpected continental disjunctions in Hypodryas gillettii, referred to above, and
the Sonoran Poladryas, in my opinion a displaced Eurasiatic genus. Nor is there any basically
allopatric Melitaeine group in North America; all genera are traceable to dispersal from
neighbouring populations. The tropical region is usually considered to have its frontier in
southern Mexico. Between this and Nearctic North America there is an important transitional
area, continuous with the southern states of Arizona, Texas, New Mexico and S. California,
forming the so-called Sonoran region. The situation here is exceptional, with vast areas of dry,
semi-desert country and an interesting and individual butterfly fauna. Melitaeine butterflies in
this region include the very small species of Texola and Dymasia, and Thessalia leanira, with its
remarkable series of local forms or subspecies, each with rather grotesque facies and not obviously
related to Melitaea. The structure of the genitalia in these interesting little species (valves with
three terminal processes) shows them all to be members of the Chlosyne series, of which such
tropical forms as C. lacinia and C.janais begin to appear. The Phyciodini are also well represented,
especially species of the genus Anthanassa, very different taxonomically and in appearance from
the typical Phyciodes that fly in North America.

Central and South America

These regions are taken together as their Melitaeine faunas are so similar. In Central America
the genus Chlosyne attains its maximum tropical development with 16 species or more, in a
compact group centred in Mexico and extending across Central America to Colombia and
Venezuela. A single species, C. lacinia, with many confusing polymorphic forms, has extended its
distribution northwards as an occasional migrant in the Sonoran region, and southwards to
include Brazil and northern Argentina. This species is exceptional in showing normal family
coloration of dark markings upon a fulvous ground in the local forms californica and
saundersi, the latter widespread in many parts of South America, and treated with specific rank
by many local entomologists. In Central and South America the Phyciodini, with 13 genera, is
the dominant tribe, extending southwards to Bolivia and northern Argentina, forming the
principal subject of this paper. Two small divergent genera, not members of the Phyciodini,
occur in N. Peru, Bolivia and S. Colombia. Two of these, Gnathotriche and Gnathotrusia, are
dealt with on pp. 160-163. Another genus, Higginsius, has a single species which flies in the same
area, and appears equally out of place among the Phyciodes series. The genitalic characters and
primitive type of wing venation suggest relationship with Gnathotriche, although the wing
markings are quite different. In fact, in Central and South America, the very frequent modification
of the wing markings to form cryptic or mimetic patterns has greatly reduced the taxonomic
importance of wing markings in these Melitaeinae.

The Antilles

The subfamily is poorly represented with seven species, of which six are truly endemic,
representing two, or perhaps three tribes. Of the two Phyciodes species, P. phaon is probably a
recent introduction from the U.S.A., but the second, P. frisia, is endemic on each of the larger
islands. The genus Antillea, with two very small species endemic on Jamaica, Cuba and Haiti,
is probably best placed with Melitaea, but its characters are very divergent, and it is certainly
unrelated to the American fauna. A third genus, Atlantea, is represented by a single species on each
of the larger islands. The species are closely related to each other, but their characters, especially
those of the females, are so unusual that I cannot place the genus in any tribal group.
Satisfactory material, especially of the rare females, is not available for examination.

Considering the situation of these islands so close to mainland America, it is remarkable that
there is so little evidence of relationship with the American Melitaeine fauna. The analysis
suggests a highly individual fauna, distantly related to Europe.

Acknowledgements

I have to record my grateful thanks to the Trustees of the British Museum (Natural History)
for their permission to study the collections and for encouragement in the preparation of this
review. I wish to thank Mr C. A. B. Steele, the Curator of the Booth Museum, Brighton,

REVISION OF PHYCIODES 175

Sussex, for permission to study the Hall collection of Phyciodini, and for allowing me to
dissect several interesting specimens from this remarkable collection. I must record my special
gratitude to my late friend Mr Norman Riley for many useful suggestions and for his help in
settling many difficulties of nomenclature. Dr D. R. Ragge, Mr R. I. Vane-Wright, Mr W. G.
Tremewan and Mrs R. Arora have given indispensable help, especially in editorial matters and
in settling the arrangement of figures and text in the final stages. I owe several useful specimens
to the kindness of Prof. Keith Brown, now working at the University of Campinas, Sao Paulo,
Brazil, who has helped greatly with information about the habitats and distribution of
Brazilian species.

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REVISION OF PHYCIODES 179

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REVISION OF PHYCIODES

181

21

•*w - i^- 19 ^W 1 ? *^P^ 20

Figs 1-21 1-11, Phyciodes species. (1) P. tharos (Drury), Scranton, Pennsylvania; (2) P. batesii
(Reakirt), Scranton, Pennsylvania; (3) P. montanus (Behr), Reno, Nevada; (4) P. campestris campestris
(Behr), Sherman Camp, Oregon; (5) P. campestris camillus Edwards, Divide, Colorado; (6) P. mylitta
mylitta (Edwards), Crater Lake, Oregon; (7) P. mylitta thebais (Godman & Salvin), Calderas,
Guatemala; (8) P. pallidus (Edwards), Warm Springs, Jefferson County; (9) P. pictus pictus
(Edwards), Arizona; (10) P. phaon (Edwards), Louisiana; (11) P. vesta vesta (Edwards), Bexar
County, Texas. 12, 13, Phystis species. (12) P. simois simois (Hewitson), San Antonio da Barra,
Bahia, Brazil; (13) P. simois variegata (Rober), Santiago de Chaquitos, Bolivia. 14-21, Anthanassa
species. (14) A. drusilla drusilla (Felder & Felder), Merida, Venezuela; (15) A. drusilla alceta
(Hewitson), Chanchamayo, Peru; (16) A. drusilla verena (Hewitson), Cochabamba, Bolivia; (17)
A. ptolyca ptolyca (Bates), San Geronimo, Guatemala; (18) A. ardys ardys (Hewitson), Cordoba,
Mexico; (19) A. ardys subota (Godman & Salvin), Vera Paz, Guatemala; (2) A. dracaena (Felder
& Felder), 'New Granada' [Colombia!: (2H A. ohleaias (Godman & Salvini Honduras.

42

Figs 22-42 Anthanassa species. (22) A. texana texana (Edwards), Sonora, Mexico; (23) A. alexon
alexon (Godman & Salvin), Guerrero, Mexico; (24) A. acesas (Hewitson), Venezuela; (25) A. nebulosa
(Godman & Salvin), Manaure, N. Colombia; (26) A. argentea (Godman & Salvin), Atoyac, Vera
Cruz, Mexico; (27) A. atronia (Bates) J, Zapoto, Guatemala; (28) A. atronia (Bates) £, Atoyac, Vera
Cruz, Mexico; (29) A. otanes otanes (Hewitson) ^, Duenas, Guatemala; (30) A. otanes otanes
(Hewitson) ?, Christobal, Vera Paz, Guatemala; (31) A. annulata sp. n., Rio Dagua, Colombia; (32)
A. crithona (Salvin), Chiriqui, Panama; (33) A.fulviplaga (Butler), Cache, Costa Rica; (34) A. hermas
(Hewitson), Sapucay, Paraguay; (35) A. frisia (Poey), Holgunin, Cuba; (36) A. tulcis (Bates),
Yucatan, Mexico; (37) A. dubia (Hall), San Esteban, Venezuela; (38) A. taeniata (Rober), Vina,
N. W. Peru; (39) A. sosis (Godman & Salvin), Irazu, Costa Rica; (40) A. drymaea (Godman &
Salvin), Calderas, Guatemala; (41) A. sitalces (Godman & Salvin), Chilasco, Guatemala; (42)
A. cortes (Hall), Omilteme, Guerrero, Mexico.

REVISION OF PHYCIODES

183

64

Figs 43-64 43-46, Dagon species. (43) D. pusillus (Salvin), Rio Colorado, Peru; (44) D. catula
(Hopffer), Charaplaya, Bolivia; (45) D. morenus (Rober), Cuzco, Peru; (46) D. fontus (Hall),
Guyana. 47-62, Telenassa species. (47) T. teletusa (Godart), Novo Friburgo, Brazil; (48)
T. berenice (Felder & Felder), Chanchamayo, Peru; (49) T. burchelli (Moulton), Chanchamayo,
Peru; (50) T. signata (Hall), La Rioja, Argentina; (51) T. abas (Hewitson), Siato, W. Colombia;
(52) T. jana (Felder & Felder), Moyobamba, Peru; (53) T. elaphina (Rober), Cochabamba,
Bolivia; (54) T. nana (Druce), Rio Colorado, Peru; (55) T. nussia (Druce), Chachapoyas, Peru;
(56) T. notus (Hall), Pozzuzo, Peru; (57) T. gaujoni (Dognin), Ecuador; (58) T. trimaculata
(Hewitson), Ecuador; (59) T.flavocincta (Dognin), Chachapoyas, Peru; (60) T. catenaria (Godman
& Salvin), W. Colombia; (61) T. delphia (Felder & Felder), Magdalena Valley, Colombia; (62)
T. sepulta (Hall), Chachapoyas, Peru. 63, 64, Ortilia species. (63) O. liriope (Cramer), Obydos,
Amazon; (64) 0. gentina sp. n., Tucuman, Argentina.

184

L. G. HIGGINS

80

Figs 65-85 65-72, Ortilia species. (65) 0. orthia (Hewitson), Sapucay, Paraguay; (66) 0. sejona
(Schaus), Goias, Brazil; (67) O. velica (Hewitson), Sao Paulo, Brazil; (68) O. zamora (Hall),
Espiritu Santo, Brazil; (69) O. dicoma (Hewitson), Sapucay, Paraguay; (70) 0. polinella (Hall),
San Fidelis, Rio de Janeiro; (71) O. orticas (Schaus), Rio de Janeiro, Brazil; (72) 0. ithra (Kirby),
Rio Iguassu, Brazil. 73, 74, Tisona species. (73) T. saladillensis saladillensis (Giacomelli), Salta,
Argentina; (74) T. saladillensis clarior subsp. n., Bolivia. 75-85, Tegosa species. (75) T. claudina
(Escholtz), Iguassu, Parana, Brazil; (76) T. similis nom. n., Siapure, Venezuela; (77) T. orobia
(Hewitson), Sapucay, Paraguay; (78) T. fragilis (Bates), Altamira, Brazil; (79) T. infrequent sp.
n., Petropolis, Brazil; (80) T. ursula (Staudinger), Sierra de Aconquija, N. Argentina; (81) T.flavida
(Hewitson), N. Peru; (82) T. pastazena (Bates), Pozzuzo, E. Peru; (83) T. guatemalena (Bates),
Oaxaca, Mexico; (84) T. anieta anieta (Hewitson) j, El Valle, Panama; (85) same V, San Luis
Potozi, Mexico.

REVISION OF PHYCIODES

185

102

Figs 86-102 86-94, Tegosa species. (86) T. anieta cluvia (Godman & Salvin), La Paz, Bolivia; (87)
T. anieta luka subsp. n., Pacho, Colombia; (88) T. anieta serpia subsp. n., Tarapoto, Peru; (89)
T. nazaria (Felder & Felder), Bogota, Colombia; (90) T. tissoides (Hall), Salidero, N. W. Ecuador;
(91) T. etia (Hewitson), Cochabamba, Bolivia; (92) same, Chachapoyas, Peru; (93) T. nigrella
(Bates), San Geronimo, Guatemala; (94) same, Cache, Costa Rica. 95-102, Eresia species. (95)
E. nauplius nauplius (L.), Cayenne, Guyana; (96) E. nauplius extensa (Hall), Matto Grosso, Brazil;
(97) E. plagiata (Rober), Santo Paulo d'Olivencia, Brazil; (98) E. letitia letitia Hewitson, Tolima,
Colombia; (99) E. letitia ocellata (Rober), Chanchamayo, Peru; (1 10) E. lansdorfi (Godart), Iguassu,
Parana, Brazil; (101) E. sestia Hewitson ^, Paramba, Ecuador; (102) same V, Zaruma, Ecuador.

186

L. G. HIGGINS

108

114

Figs 103-114 Eresia species. (103) E. coela Druce d, Limon, Costa Rica; (104) same y, Guapiles,
Costa Rica; (105) E. oblita (Staudinger), San Esteban, Venezuela; (106) E. carme carme Doubleday
J, Venezuela; (107) same ? holotype, no locality [Venezuela]; (108) E. carme laias Godman &
Salvin, Rio Aquatil, W. Colombia; (109) E. polina Hewitson, Cochabamba, Bolivia; (110) E. alsina
Hewitson j1, Cache, Costa Rica; (111) same 'y, Jinotega, Nicaragua; (112) E. cissia (Hall), Bogota,
Colombia; (113) E. eutropia Hewitson J, El Valle, Panama; (114) same ?, El Valle, Panama.

REVISION OF PHYCIODES

187

120

Figs 115-124 Eresia species. (115) E. poecilina Bates 3, Chiriqui, Panama; (116) E. mimas
(Staudinger) <£, Rio San Juan, W. Colombia; (117) E. melaina sp. n. 3, Veraguas, Panama; (118)
same v, El Valle, Panama; (119) E. stricta Schaus, Guapiles, Costa Rica; (120) E. ithomiodes
ithomiodes Hewitson J, Rio Chili, Colombia; (121) E. poecilina Bates V holotype, Santa Fe;
(122) E. quintilla Hewitson, Cauca, Colombia; (123) £. anomala sp. n. ? holotype, Muzo, Colombia;
(124) E. ithomioides pseudocelemina (Strand), Bogota, Colombia; (124a) E. ithomioides ithomioides
Hewitson ?, Juntas, Choco, Colombia.

188

L. G. HIGGINS

134

Figs 125-134 Eresia species. (125) E. nigripennis Salvin J, Costa Rica; (125a) same *, Cache, Costa
Rica; (126) E. emerantia Hewitson J1, Colombia; (127) same v, Cananche, Colombia; (128)
E. phaedima Salvin & Godman, Chanchamayo, Peru; (129) E. datis datis Hewitson, Cochabamba,
Bolivia; (130) E. margaretha Hewitson, Colombia; (131) E. eunice eunice (Hiibner), Para, Brazil;
(132) E. eunice olivencia Bates, Sao Paulo da Olivencia, Brazil; (133) E. etesiae (Hall) j, French
Guiana; (134) same *, Cayenne, French Guiana.

REVISION OF PHYCIODES

189

146

Figs 135-146 Eresia species. (135) E. eunice esora Hewitson, Sao Paulo, Brazil; (136) E. erysice
(Geyer) *, Bahia, Brazil; (137) E. casiphia Hewitson J, Balsapamba, Bolivar, Ecuador; (138) same
i, Balsapamba, Bolivar, Ecuador; (139) E. mechanics Godman & Salvin, Chontales, Costa Rica;
(140) E. pelonia Hewitson J, Sarayacu, Ecuador; (141) same, Peru; (142) same i, Pebas, Amazon;
(143) E. phillyra Hewitson j , Cordoba, Mexico; (144) same i, Honduras; (145) E. aveyrona aveyrona
Bates, Cayenne, French Guiana; (146) E. perna Hewitson, Espirito Santo, Brazil.

190

L. G. HIGGINS

150

151

Figs 147-157 147-150, Castilia species. (147) C. occidental (Fassl) <$, Cauca, Colombia; (148)
same 9, lectotype of aurora Rober, Zamora, Ecuador; (149) C. perilla (Hewitson) 3, Peru; (150)
same ?, Moyabamba, Peru. 151-154, Eresia species. (151) E. levina Hewitson, Manizales, Colombia;
( 1 52) E.act inote Salving, Peru; (153) same V, Carillo, Costa Rica; (154) E. selene (Rober), Manizales,
Colombia. 155-157, Castilia species. (155) C. castilla (Felder & Felder) ^, Bogota, Colombia; (156)
same ?, Bogota, Colombia; (157) C. neria (Hewitson), Ecuador.

REVISION OF PHYC10DES

191

Figs 158-174 158-166, Castilia species. (158) C. nortbrundii (Weeks), Cochabamba, Bolivia; (159)
C. eranites (Hewitson) J, Balzapamba, Ecuador; (160) same ?, Costa Rica; (161) C.fulgora (Godman
& Salvin), Castajal, Costa Rica; (162) C.fausta (Godman & Salvin), Chiriqui, Panama; (163)
C. ofella (Hewitson) Muzo, Colombia; (164) C. myia (Hewitson), Cordoba, Mexico; (165) C.
griseobasalis (Rober), Santa Tecla, Guatemala; (166) C. angusta (Hewitson), Peru. 167- 169, Janatella
species. (167) J. leucodesma (Felder & Felder), Merida, Venezuela; (168) J. Hera (Cramer), French
Guyana; (169) J.fellula (Schaus), Cauca, Colombia. 170, Mazia amazonica (Bates), Sao Paulo
d'Olivencia, Amazon. 171, 172, Castilia species. (171) C. chinantlensis (de la Maza) j, Oaxaca,
Mexico; (172) same ?, Oaxaca, Mexico. 173, 174, Eresia species. (173) E. moesta Salvin & Godman,
Env. Bogota, Colombia; (174) E. mimas (Staudinger), Cauca, Juntas, Colombia.

192

L. G. HIGGINS

176

177

Figs 175-177 175, 176 Castilia species. (175) C. perilla f. aricilla (Hopffer), Pozuzo, Peru; (176)
C. perilla f. acraeina (Hewitson), Moyobamba, Peru. 177, Eresia clara (Bates), Rio Cachiyaca,
Iquitos, Peru.

REVISION OF PHYCIODES

193

178

H

Figs 178-183 178-182, Phyciodes species. (178) P. tharos, forewing venation; (179) same, hindwing
venation; (180) same, palpus, lateral view; (181) same, J1 foreleg; (182) same, J genitalia, dorsal
view, anatomical structures lettered, Missouri, g/s 134. The posterior border of the juxta is seen
rather obscurely through the chitinized tegumen. 183, Eresia clara, 3 genitalia, lateral view, penis
removed, showing dorsal structures. A = tegumen; E = scaphial extension; S = saccus; J = juxta;
H = harpe; V = valve. Scale = 1 mm.

194

L. G. HIGGINS

184

185

188

190

187

Figs 184-191 184, 185, Eresia species. (184) E. clara, penis, lateral view of distal section, showing
apical structures lettered. OK = ostium keel of penis; M = morula of penis. (185) E. phillyra,
t sterigma, showing structure of the genital plate and bursal support. B = position of bursa;
BS = bursal support; BD — bursal duct; O = ostium bursae; S = scutum. 186-191 Phyciodes
species. (186) P. tharos, $ genitalia, dorsal view, Chicago, g/s 1230; (187) same, 3 genitalia, lateral
view, Connecticut, g/s 1233; (188) same, tegumen + scaphial extensions; (189) same, posterior
border of juxta; (190) same, saccus; (191) same, penis, lateral view, Chicago, g/s 1230. Scale = 1 mm.

REVISION OF PHYCIODES

Figs 192-198 Phyciodes species. (192) P. tharos, ? genitalia, dorsal view, Chicago, g/s 128; (193)
P. batesii, 3 genitalia, dorsal view, Scranton, Perm., g/s 130; (194) P. campestris campestris, <$
genitalia, dorsal view, Oregon, g/s 422; (195) same, lateral view, Yosemite, g/s 2597; (196) same,
tegumen, California, g/s 421; (197) same, posterior border of juxta, California, g/s 421; (198) same,
penis, lateral view, Wyoming, g/s 2592. Scale — 1 mm.

196

L. G. HIGGINS

Figs 199-208 Phyciodes species. (199) P. campestris campestris, 9 genitalia, Alberta, g/s 1057; (200)
P. c. camillus, 3 tegumen, Wyoming, g/s 1592; (201) same, posterior border of juxta, Wyoming,
g/s 1592; (202) same, saccus, Wyoming, g/s 1592; (203) P. montanus, j genitalia, California, g/s
423; (204) same, j tegumen, California, g/s 423; (205) same, posterior border of juxta, California,
g/s 423; (206) same, saccus, California, g/s 423; (207) same, penis, lateral view, California, g/s 778;
(208) same, * genitalia, California, g/s 2617. Scale = 1 mm.

REVISION OF PHYCIODES

197

Figs 209-214 Phyciodes species. (209) P. mylitta, j genitalia, dorsal view, California, g/s 119; (210)
P. m. mylitta, penis, lateral view, Golden, Colorado, g/s 794; (211) same, * genitalia, Oregon,
g/s 1056; (212) P. orseis, tegumen and scaphial extension, Oregon, Gold Hill, g/s 780; (213)
P. pictus pictus, j genitalia, dorsal view, Tucson, Arizona, g/s 792; (214) P. pictus pallescens,
•3 genitalia, dorsal view, Guadalajara, g/s 793. Scale = 1 mm.

198

L. G. HIGGINS

Figs 215-223 215-218, Phyciodes species. (215) P. phaon, 3 genitalia, dorsal view, no locality, g/s
396; (216) same, penis, lateral view, no locality, g/s 396; (217) P. vesta vesta, $ genitalia, dorsal
view, Texas, g/s 773; (218) same, V sterigma, Colorado Springs, g/s 1079. 219-223, Phystis species.
(219) P. simois simois, 3 genitalia, dorsal view, Pernambuco, g/s 790; (220) same, penis, lateral
view, Pernambuco, g/s 790; (221) P. simois variegata intermediate form, j genitalia, dorsal view,
Bahia, g/s 2755; (222) same, penis, lateral view, Bahia, g/s 2755; (223) same, 9 genitalia, Cordoba,
g/s 1078. Scale = 1 mm.

REVISION OF PHYCIODES

224

226

229

227

228

230

Figs 224-230 Anthanassa species. (224) A. drusilla drusilla, _> genitalia, dorsal view, Colombia,
g/s 1036; (225) same, valve, lateral view, Venezuela, g/s 2776; (226) same, penis, lateral view,
Venezuela, g/s 1032; (227) same, pyriform antennal club; (228) same, ? genitalia, Colombia,
g/s 1033; (229) A. ptolyca ptolyca, J genitalia, dorsal view, Guatemala, g/s 106; (230) same, penis,
lateral view, Guatemala, g/s 106. Scale = 1 mm.

200

L. G. HIGGINS

232

234

235

236

233

Figs 231-236 Anthanassa species. (231) A. ardys ardys, d genitalia, dorsal view, Costa Rica, g/s 814;
(232) same, penis, lateral view, Colombia, g/s 1350; (233) A. ardys subota, 5 genitalia, dorsal
view, Guatemala, g/s 2565; (234) A. dracaena, j genitalia, dorsal view, Colombia, g/s 817; (235)
same, penis, lateral view, Colombia, g/s 8 17 ; (236) same, ? genitalia, Colombia, g/s 8 1 7. Scale = 1 mm.

REVISION OF PHYCIODES

237

238

239

Figs 237-241 Anthanassa species. (237) A. texana texana, J genitalia, dorsal view, Texas, g/s 1042;
(238) same, penis, lateral view, Texas, g/s 1042; (239) same, ? genitalia, Texas, g/s 1050; (240)
A. alexon subconcolor, _> genitalia, dorsal view, Arizona, g/s 801; (241) A. acesas, j genitalia, dorsal
view, Venezuela, g/s 1024. Scale = 1 mm.

242

L. G. HIGGINS

244

243

245

246

Figs 242-246 Anthanassa species. (242) A. nebulosa, j genitalia, dorsal view, N. Colombia, Sierra
Nevada de Sante Marta, Chichihua Valley (f. castianira), g/s 110; (243) same, tegumen and
scaphial extension, g/s 1252; (244) A. argentea, j genitalia, dorsal view, Guatemala, g/s 1018; (245)
A. atronia, j genitalia, dorsal view, Guatemala, g/s 829; (246) A. otanes, j genitalia, dorsal view,
Guatemala, g/s 1247. Scale = 1 mm.

REVISION OF PHYCIODES

250

249

252

251

Figs 247-252 Anthanassa species. (247) A. annulata, J genitalia, dorsal view, Ecuador, g/s 1351;
(248) same, penis, lateral view of distal section, Ecuador, g/s 1351; (249) A. crithona, j genitalia,
dorsal view, Costa Rica, g/s 2578; (250) same, penis, lateral view, Costa Rica, g/s 807; (251) same,
$ genitalia, dorsal view, Panama, g/s 1049; (252) A. fulviplaga, J genitalia, dorsal view, Costa Rica,
g/s 833. Scale = 1 mm.

L. G. HIGGINS

254

Figs 253-259 Anthanassa species. (253) A. fulviplaga, penis, lateral view, Costa Rica, g/s 833; (254)
same, ? genitalia, Costa Rica, g/s 1066; (255) A. hernias, ,3 genitalia, dorsal view, Paraguay, g/s 111 ;
(256) same, penis, lateral view, Paraguay, g/s 111; (257) same, 9 genitalia, dorsal view, Brazil,
g/s 1067; (258) A. frisia, .3 genitalia, Cuba, g/s 2780; (259) same, 3 genitalia, Jamaica, g/s 879.
Scale = 1 mm.

REVISION OF PHYCIODES

260

262

265

263

Figs 260-265 Anthanassa species. (260) A. frisia, 3 genitalia, lateral view, Jamaica, g/s 879; (261)
A. tulcis, ^ genitalia, Panama, g/s 882; (262) same, penis, Panama, g/s 882; (263) same, * genitalia,
Panama, g/s 117; (264) A. dubia, 3 genitalia, Venezuela, San Esteban, g/s 2783; (265) A. taeniata,
o genitalia, Peru, g/s 880. Scale = 1 mm.

206

L. G. HIGGINS

266

269

268

270

267

Figs 266-271 266-269, Anthanassa species. (266) A. sosis, d genitalia, dorsal view, Panama, Chiriqui,
g/s 1017; (267) A. drymaea, <$ genitalia, dorsal view, Guatemala, g/s 836; (268) A. shakes, £ genitalia,
dorsal view, Guatemala, g/s 1346; (269) A. cartes, <$ genitalia, Mexico, g/s 1029. 270, 271, Dagon
species. (270) D. pusillus, J genitalia, dorsal view, no locality, g/s 802; (271) same, penis, lateral
view, no locality, g/s 802. Scale = 1 mm.

REVISION OF PHYCIODES

Figs 272-279 272-274, Dagon species. (272) D. catula, 3 genitalia, dorsal view, Bolivia, g/s 812;
(273) same, penis, lateral view, Bolivia, g/s 812; (274) D. morenus, j genitalia, dorsal view, Peru,
g/s 2768. 275-279, Telenassa species. (275) T. teletusa, j genitalia, dorsal view, Brazil, Novo
Friburgo, g/s 2563; (276) same, penis, lateral view, Brazil, Novo Friburgo, g/s 2563; (277) same,
9 genitalia, Brazil, Tucuman, g/s 1049; (278) T. burchelli, o genitalia, dorsal view, Peru,
Chanchamayo, g/s 799; (279) same, penis, lateral view, Peru, Chanchamayo, g/s 799. Scale = 1 mm.

L. G. HIGGINS

Figs 280-285 Telenassa species. (280) T. berenice, <J genitalia, Peru, Amazonas, g/s 2801; (281)
same, penis, Peru, g/s 936; (282) T. signata, <$ genitalia, dorsal view, Argentina, Salta, g/s 2749;
(283) T. abas, <$ genitalia, dorsal view, Colombia, Frontino, g/s 2749; (284) same, penis, lateral view,
Colombia, 'Bogota', g/s 2752; (285) T. jana, <$ genitalia, dorsal view, Peru, Amazonas, g/s 109.
Scale = 1 mm.

REVISION OF PHYCIODES

287

290

286

Figs 286-290 Telenassa species. (286) T. jana, 9 genitalia, Peru, Chachapoyas, g/s 1062; (287)
T. elaphina, 3 genitalia, dorsal view, Bolivia, g/s 809; (288) T. nana, <3 genitalia, dorsal view,
S. Peru, Cosnipata Valley, g/s 2575; (289) same, penis, lateral view, g/s 810; (290) T. nussia, <$
genitalia, Peru, g/s 1028. Scale = 1 mm.

210

293

L. G. HIGGINS

291

295

Figs 291-297 Telenassa species. (291) T. notus, 3 genitalia, dorsal view, Peru, g/s 808; (292) same,
penis, lateral view, Peru, g/s 808; (293) same, ? genitalia, dorsal view, Peru, g/s 1085; (294) T.
gaujoni, -3 genitalia, dorsal view, Ecuador, g/s 1016; (295) T. trimaculata, 3 genitalia, dorsal view,
Bolivia, g/s 899; (296) T. flavocincta, 3 genitalia, dorsal view, Ecuador, g/s 1025; (297) same,
penis, Ecuador, g/s 1025. Scale = 1 mm.

REVISION OF PHYCIODES

298

299

303

Figs 298-303 298 301, Telenassa species. (298) T. catenarie, $ genitalia (excluding tegumen),
W. Colombia, g/s 2757; (299) T. delphia, 3 genitalia, dorsal view, Colombia, g/s 812; (300) T. sepulta,
d genitalia, dorsal view, Peru, g/s 898; (301) same, penis, lateral view, Peru, g/s 898. 302, 303,
Ortilia species. (302) O. liriope, 3 genitalia, dorsal view, Guyana, g/s 1559; (303) same, penis, lateral
view, Brazil, Para, g/s 816. Scale = 1 mm.

212

L. G. HIGGINS

Figs 304-310 Ortilia species. (304) O. liriope, ? genitalia, Lower Amazon, g/s 817; (305) O. gentina,
-j genitalia, dorsal view, Bolivia, g/s 825; (306) same, J genitalia, lateral view, Bolivia, g/s 1082;
(307) 0. orthia, J genitalia, dorsal view, Castro, Parana, Brazil, g/s 797; (308) O. orticas, 5 genitalia,
dorsal view, Brazil, Parana, g/s 883; (309) same, * genitalia. Brazil, g/s 1086; (310) O. sejona, j
genitalia, dorsal view, Brazil, Goias, Campinas, g/s 2800. Scale = 1 mm.

REVISION OF PHYCIODES

Figs 311-317 Ortilia species. (311) O. velica, £ genitalia, dorsal view, Brazil, Sao Paulo, g/s 800;
(312) 0. velica durnfordi, V genitalia, dorsal view, g/s 1090; (313) O. zamora, <$ genitalia, dorsal
view, Rio de Janeiro, g/s 2753; (314) 0. dicoma, J genitalia, dorsal view, Brazil, g/s 834; (315)
same, penis, lateral view, Brazil, g/s 834; (316) same, ? genitalia, dorsal view, Brazil, g/s 1089; (317)
0. polinella, -S genitalia, dorsal view, Brazil, Minas Geraes, g/s 1084. Scale = 1 mm.

214

L. G. HIGGINS

320

326

Figs 318-326 318 320, Ortilia species. (318) O. ithra, 3 genitalia, dorsal view, Paraguay, g/s 853a;
(319) same, ,J genitalia, dorsal view, Brazil, Mato Grosso, g/s 853b; (320) same, penis, distal section,
lateral view, Brazil Mato Grosso, g/s 852. 321 324, Tisona species. (321) T. saladillensis, 3 genitalia,
dorsal view, Salta, g/s 878; (322) same, J genitalia, lateral view, Salta, g/s 881; (323) same, penis,
lateral view, Salta, g/s 878; (324) same, + genitalia, Salta, g/s 1077. 325, 326, Tegosa species.
(325) T. claudina, _> genitalia, dorsal view, Paraguay, g/s 135; (326) same, penis, lateral view,
Brazil, Parana, g/s 136. Scale = 1 mm.

REVISION OF PHYCIODES

328

Figs 327-333 Tegosa species. (327) T. similis, tegumen + scaphial extension, 'Indiis', lectotype, g/s
1557; (328) same, o genitalia, 'Indiis', lectotype, g/s 1557; (329) same, o genitalia, Trinidad, g/s
893; (330) same, penis, lateral view, Brazil, g/s 2700; (331) same, saccus and tegumen + scaphial
extension, Brazil, g/s 2700; (332) same, o genitalia, lateral view, Brazil, Minas Geraes, g/s 2700;
(333) same, $ genitalia, Brazil, g/s 1080. Scale = 1 mm.

216

L. G. HIGGINS

Figs 334-341 Tegosa species. (334) T. orobia, 3 genitalia, Brazil, Castro, Parana, g/s 892; (335) same,
penis, lateral view, Brazil, Castro, Parana, g/s 892; (336) same, J genitalia, lateral view, g/s 1073;
(337) T.fragilis, J genitalia, dorsal view, 'Brazil', g/s 2595; (338) same, penis, lateral view, 'Brazil',
g/s 2595; (339) T. infrequens, J genitalia, dorsal view, Bolivia, g/s 2714; (340) same, penis, lateral
view, Bolivia, g/s 2714; (341) T. ursula, 3 genitalia, Argentina, Tucuman, g/s 888. Scale = 1 mm.

REVISION OF PHYCIODES

Figs 342-348 Tegosa species. (342) T. flavida, 3 genitalia, dorsal view, Ecuador, g/s 2583; (343)
T. tissoides, 3 genitalia, Ecuador, g/s 1117; (344) T. pastazena, 3 genitalia, Peru, Pozzuzo,
Huanuco, g/s 827; (345) same, penis, lateral view, Peru, Pozzuzo, Huanuco, g/s 827; (346) T.
guatemalena, 3 genitalia, dorsal view, Guatemala, g/s 2590; (347) same, 3 genitalia, lateral view,
Mexico, Oaxaca, g/s 1193; (348) same, j genitalia with tegumen displaced and right valve omitted,
Mexico, g/s 125. Scale = 1 mm.

L. G. HIGGINS

Figs 349-356 Tegosa species. (349) T. anieta anieta, •$ genitalia, Panama, g/s 1020; (350) same,
•3 genitalia, Peru, g/s 1072; (351) same, J genitalia, lateral view, Venezuela, Caracas, g/s 1202;
(352) T. anieta cluvia, J1 genitalia, Guatemala, g/s 1116; (353) T. anieta luka, j genitalia, Mexico,
g/s 2596; (354) same, J genitalia, Ecuador, g/s 855; (355) T. anieta anieta, j genitalia, lateral view,
Venezuela, g/s 876; (356) same, penis, Panama, Onaca, g/s 839. Scale = 1 mm.

REVISION OF PHYCIODES

219

358

364

Figs 357-367 357-363, Tegosa species. (357) T. nazaria, 3 genitalia, Colombia, g/s 877; (358) same,
penis, lateral view, Colombia, g/s 877; (359) T. etia, <$ genitalia, Guatemala, g/s 1125; (360) same,
penis, lateral view, Guatemala, g/s 1125; (361) T. nigrella, 3 genitalia, dorsal view, Costa Rica,
g/s 2571 ; (362) same, o genitalia, lateral view, Costa Rica, g/s 831 ; (363) same, side view of palpus.
364 367 Eresia species. (364) E. clara, S genitalia, Venezuela, g/s 2559; (365) same, penis, Ecuador,
g/s 2559 (2); (366) same, ? genitalia, Mexico, g/s 1061; (367) E. clara f. reducta, tegumen, g/s
2559 (1). Scale = 1 mm.

220

L. G. HIGGINS

372

Figs 368-377 Eresia species. (368) E. nauplius nauplius, J genitalia, dorsal view, Brazil, Obidos,
g/s 2726; (369) same, tegumen, Brazil, Obidos, g/s 2742 (1); (370) same, J genitalia, dorsal view,
Guyana, g/s 636; (371) same, penis, distal section, lateral view, French Guiana, g/s 2584; (372) same,
j genitalia, lateral view, Guyana, g/s 2733; (373) same, V genitalia, dorsal view, Brazil, Obidos, g/s
2734; (374) same, j1 genitalia, dorsal view, 'Brazil', g/s 2736; (375) E. plagiata, dorsal structures,
Brazil, Cucaru, g/s 2742 (2); (376) same, juxta, Brazil, Cucaru, g/s 2742(2); (377) same, j genitalia,
dorsal view, Peru, g/s 2560. Scale = 1 mm.

REVISION OF PHYCIODES

221

378

383

Figs 378-384 Eresia species. (378) E. plagiata. j genitalia, lateral view, Brazil, Teffe, g/s 2737; (379)
same, ? genitalia, Brazil, g/s 637; (380) E. letitia ocellata (neptoides), j genitalia, dorsal view,
Peru, g/s 1244; (381) E. letitia ocellata (nigra), j genitalia, dorsal view, Peru, g/s 91; (382) E. /efm'a
letitia, j genitalia, dorsal view, Ecuador, g/s 2782; (383) E. letitia ocellata, j genitalia, lateral view,
Peru, g/s 186; (384) E. terma, penis, lateral view, g/s 1242. Scale = 1 mm.

L. G. HIGGINS

Figs 385-390 Eresia species. (385) E. lansdorfi, _> genitalia, dorsal view, Brazil, g/s 92; (386) E. sestia,
3 genitalia, dorsal view, Ecuador, g/s 633; (387) same, penis lateral view, Ecuador, g/s 2713; (388)
E. coela, _> genitalia, dorsal view, Costa Rica, g/s 2790; (389) same, + genitalia, g/s 2740; (390)
E. oblita, genitalia, dorsal view, Venezuela, g/s 2769. Scale = 1 mm.

REVISION OF PHYCIODES

223

393

Figs 391-397 Eresia species. (391) E. carme carme, j genitalia, dorsal view, Colombia, g/s 1241;
(392) same, penis, lateral view, Colombia, g/s 1241 ; (393) same, j genitalia, dorsal view, W. Colombia,
g/s 2723 (2); (394) E. polina, _> genitalia, dorsal view, Colombia, g/s 89; (395) same, penis, lateral
view, Colombia, g/s 89; (396) E. alsina, j genitalia, dorsal view, Costa Rica, g/s 628; (397) same,
penis, lateral view, Costa Rica, g/s 628. Scale = 1 mm.

224

L. G. HIGGINS

398

Figs 398-403 Eresia species. (398) E. alsina, Y genitalia, Nicaragua, g/s 629 ; (399) E. cissia, 3 genitalia,
dorsal view, W. Colombia, g/s 2771; (400) same, penis, lateral view, W. Colombia, g/s 2771; (401)
same, penis, dorsal view, W. Colombia, g/s 2771; (402) E. eutropia, j genitalia, dorsal view, W.
Colombia, g/s 630; (403) E. quintilla, J genitalia, dorsal view, Ecuador, g/s 631. Scale = 1 mm.

REVISION OF PHYCIODES

406

408

407

Figs 404-408 Eresia species. (404) E. quintilla, * genitalia, Ecuador, g/s 632; (405) E. sticta, J
genitalia, dorsal view, Costa Rica, g/s 1023; (406) E. ithomioides, J genitalia, dorsal view, Colombia,
g/s 1240; (407) same, penis, lateral view, Colombia, g/s 1240; (408) E. nigripennis, j genitalia, dorsal
view, Costa Rica, g/s 896. Scale = 1 mm.

L. G. HIGGINS

410

Figs 409-413 Eresia species. (409) E. emerantia, j genitalia, dorsal view, Colombia, nr Larogosa,
g/s 620; (410) same, ? genitalia, Colombia, g/s 2740; (411) E. moesta, j genitalia, no locality
[Ecuador?], g/s 2793; (412) same, penis, no locality [Ecuador?], g/s 2793; (413) E. phaedima,
3 genitalia, Peru, g/s 639. Scale = 1 mm.

REVISION OF PHYCIODES

419

Figs 414-419 Eresia species. (414) E. datis corybassa, j genitalia, dorsal view, Bolivia, g/s 622; (415)
E. margaretha, j genitalia, dorsal view, Colombia, g/s 2718; (416) E. eunice, 3 genitalia, dorsal
view, Peru, g/s 897; (417) E. eunice esora, S genitalia, Brazil, Sao Paulo, g/s 2717; (418) £.
eunice eunice, penis, lateral view, Brazil, g/s 1265; (419) same, j genitalia, lateral view, Peru, g/s
2561. Scale = 1 mm.

L. G. HIGGINS

422

424

Figs 420-425 Eresia species. (420) E. eunice eunice, ? genitalia, Peru, g/s 1048; (421) E. etesia, •$
genitalia, French Guiana, g/s 1026; (422) E. casiphia, j genitalia, dorsal view, Ecuador, g/s 1027;
(423) E. mechanitis, 3 genitalia, dorsal view, Nicaragua, g/s 1243; (424) E. pelonia, 3 genitalia,
dorsal view, Brazil, g/s 92; (425) same, penis, lateral view, Ecuador, g/s 2558. Scale = 1 mm.

REVISION OF PHYCIODES

429

Figs 426-431 Eresia species. (426) E. phillyra, 3 genitalia, dorsal view, Mexico, g/s 2569; (427) same,
penis, lateral view, Mexico, g/s 1239; (428) E. aveyrona aveyrona, 3 genitalia, French Guiana, g/s
635; (429) E. perna, 3 genitalia, Brazil, g/s 2716; (430) E. levina, 3 genitalia, Colombia, g/s 98;
(431) same, penis lateral view, Colombia, g/s 98. Scale = 1 mm.

L. G. HIGGINS

436

435

Figs 432-437 432-434, Eresia species. (432) E. actinote, 3 genitalia, Peru, g/s 618; (433) E. selene,
<$ genitalia, Colombia, Manisales, g/s 832; (434) same, penis, Colombia, g/s 837. 435-437, Castilia
species. (435) C. castilla, 3 genitalia, dorsal view, Colombia, g/s 615; (436) same, 9 genitalia, g/s
1164; (437) same, wing venation diagram. Scale = 1 mm.

REVISION OF PHYCIODES

440

443

Figs 438-443 Castilia species. (438) C. occidental, j genitalia, dorsal view, Ecuador, g/s 626; (439)
same, penis, lateral view, Colombia, g/s 2715; (440) same, wing venation diagram, slide 1167;
(441) C. perilla, J genitalia, dorsal view, penis removed, Ecuador, g/s 627; (442) C. perilla f. acraeina,
j1 genitalia, dorsal view, penis removed, Brazil, upper Amazons, g/s 1058; (443) C. perilla, wing
venation diagram, discoidal area. Scale = 1 mm.

L. G. HIGGINS

446

448

Figs 444-451 Castilia species. (444) C. neria, <$ genitalia, dorsal view, Ecuador, g/s 619; (445) same,
penis apex, lateral view, Ecuador, g/s 619; (446) same, wing venation diagram, discoidal area of
forewing, slide 1169; (447) C. nortbrundii, <$ genitalia, dorsal view, Bolivia, g/s 611; (448) same,
penis apex, lateral view, Bolivia, g/s 611; (449) same, palpus; (450) C. eranites, £ genitalia, dorsal
view, Colombia, g/s 2587; (451) same, penis, lateral view, Colombia, g/s 2587. Scale = 1 mm.

REVISION OF PHYCIODES

233

452

453

Figs 452-458 Castilia species. (452) C. eranites, 9 genitalia, Colombia, g/s 1065; (453) C.fulgora,
3 genitalia, dorsal view, Costa Rica, g/s 1021; (454) same, penis apex, lateral view, Costa Rica,
g/s 1021; (455) C.fausta, £ genitalia, dorsal view, Panama, g/s 1345; (456) C. ofella, <$ genitalia,
Colombia, g/s 2585; (457) same, penis, lateral view, Colombia, g/s 2585; (458) same, 9 genitalia,
g/s 1063. Scale = 1 mm.

L. G. HIGGINS

461

464

Figs 459-465 Castilia species. (459) C. myia, J genitalia, dorsal view, Mexico, Teapa, g/s 2586;
(460) same, palpus; (461) same, 4 genitalia, Guatemala, g/s 1064; (462) C. griseobasalis, j genitalia,
dorsal view, Guatemala, Volcan Santa Maria, g/s 1038; (463) same, penis, lateral view, Guatemala,
Volcan Santa Maria, g/s 1038; (464) C. angusta, .$ genitalia, dorsal view, Peru, Chanchamayo,
g/s 835; (465) same, penis, lateral view, Bolivia, g/s 2762. Scale = 1 mm.

REVISION OF PHYCIODES

Figs 466-474 466, Castilia angusta, j genitalia, oblique view, Bolivia, g/s 2762. 467 474, Janatella
species. (467) J. leucodesma, 3 genitalia, dorsal view, Trinidad, g/s 1349; (468) same, penis, lateral
view, Trinidad, g/s 1349; (469) same, <$ genitalia, lateral view, Venezuela, Caucu River, g/s 1348; (470)
same, 9 genitalia, Panama, g/s 1090; (471) J. hera, $ genitalia, dorsal view, Surinam, g/s 830; (472)
same, distal section of penis, lateral view, Surinam, g/s 830; (473) J.fellula, o genitalia, dorsal view,
Colombia, g/s 101; (474) same, 9 genitalia, Colombia, g/s 1088. Scale = 1 mm.

479

481

Figs 475481 475-479, Mazia species. (475) M. amazonica, 3 genitalia, dorsal view, 'Amazons', g/s
1365; (476) same, <$ genitalia, lateral view, no locality, g/s 459; (477) same, y genitalia, 'Amazon', g/s
1076; (478) same, palpus; (479) same, ^ forewing venation diagram, discoidal area. 480, 481,
Gnathotriche species. (480) G. exclamationis, J genitalia, dorsal view, Colombia, g/s 82; (481) same,
y genitalia, dorsal view, 'Bogota', g/s 2611. Scale = 1 mm.

REVISION OF PHYCIODES

482

487

483

486

484

Figs 482-487 482-486, Gnathotriche species. (482) G. exclamationis, forewing venation; (483) same,
hindwing venation; (484) G. sodalis, j genitalia, dorsal view, Colombia, g/s 2614; (485) same,
J genitalia, lateral view, Colombia, g/s 2814; (486) same, penis apex, dorsal view, Colombia, g/s
2814. 487, Gnathotrusia mundina, 3 genitalia, dorsal view, Colombia, g/s 547. Scale = 1 mm.

L. G. HIGGINS

\

489

Figs 488-490 488, 489, Gnathotrusia species. (488) G. epione, J genitalia, dorsal view, Colombia,
g/s 548; (489) same, saccus and juxta, dorsal view, showing conical posterior border and densely
chitinized lingulae, Colombia, g/s 548. 490, Ortilia ithra, ? genitalia, g/s 1060.

abas 84, 110, 112, 169

abasina 109

aberrans 153

abrupta 114

abyssinica 166

acastus 167

acesas 95, 101, 169

aceta 115

acraea 150

acraeina Hewitson 153

acraeina Staudinger 166

actinote83, 130,150, 170

actinotina 154

adoxa 108

Index

Principal references are in bold; invalid names are in italics.

aequatorialis 104
aetherie 166
afghana 166
agar 165
ala 166
alatauica 165
albescens 124
alceta96, 168
alethes 96
alexandrina 165
alexon 95, 100, 168
alma 150

alsina!30, 137, 138, 170
amator97, 168

INDEX

239

amazonica 159, 170
ambigua 165
ambrisia 166
amoenides 153
amoenula 166
angusta 152, 157, 170
anicia 165
anieta!22, 126, 169
annita 101
annulata!03, 168
anomala 141, 170
anomalus 167
Anthanassa 84, 94, 168
Antillea 167
apicalis 147
aquila 127
arachne 166
arcesia 166
archisilea 105
arctica 86
arduinna 165
ardys95,98, 168
argentea95, 102, 169
aricilla 153
arida 92
arizonensis 89
asteria 165
aster oidea 165
athene 166
athalia 165
Atlantea 170
atra 120

atronia 58, 102, 168
aurelia 165
aurinia 165
aurora 153

aveyrona 130,149, 170
avinovi 165

balba 165
balbita 166
barnesi 89
batesii 65, 86, 167
bella 149
bellona 166
berenicellO, 111, 169
birivula 109
boucardi 92
brancodia 104
britomartis 165
brunhilda 145
burchelli 11 1,168

calena 108
californica 167
callianira 162
callianthina 163
callonia 147
callonioides 147
camillus 87, 168
campestris 85, 87, 167

canace 91
carigia 99

carmelSO, 135, 136, 169
carrera 97

casiphia 130, 147, 170
cassiopea 102
casta 166
castianira 101
Castilia 170
castilla!52, 170
catagraphus 109
catenariallO, 114, 169
catula 108, 109, 168
celadussa 165
celemina 142
centralasiae 165
cerquita 142
chalcedona 165
chinantlensis 157, 170
chinchipensis, 160, 168
chitralensis 166
Chlosyne 166
chromis 102
Cinclidia 166
cincta 94, 100
cinxia 165
cissia!30, 137, 170
clara 129, 131, 169
clarior 121, 169
claudina 84, 121, 122, 169
c/io81, 131
cluvia!22, 126, 169
cocyta 85, 167
coela 135, 140, 169
collina 166
collina-group 166
collinsi 89
colon 165
confer ta 104
confirmans 139
conflua 97
consulis 166
coracora 167
cornelia 144
cortes95, 107, 168
corybassa 144, 170
crina 162

crithona95, 104, 168
cynisca 167
cyno 103
cynthia 165
crucifera 153

Dagon84, 108, 168
daguana 147
damaetas 167
datisBO, 143, 170
decor ata 150
definita 167
deione 165

240

delphiallO, 115, 169
derivata 143
deserticola 166
desfontainii 165
diallus 102
diamina 165
dicomall6, 119, 169
dictynna 118
dicomal!6, 119, 169
didyma 166
didyma-group 166
Didymaeformia 165
didymina 166
didymoides 166
dismorphina 142
dor a 103
douglasi 111
dracaena 95, 99, 168
draudti 167
drymaea95, 107, 168
drypetis 145
drusilla 94, 95, 96, 168
drusinilla 11 1,128
dubia95,106, 168
durnfordil!6, 118, 169
dymas 167
Dymasia 167

editha 165

edwardsi 90

ehrenbergi 167

elada 167

elaea 161

elaphiaea 112

elaphinallO, 113, 169

eleates 161

elenae 163

elva 167

emerantia 142, 170

emissa 87

encina 136

epione 163, 167

epula 88

eranites!52, 155, 170

erebia 162

Eresia 83, 84, 129, 161,169

erodyle 167

erysice 130, 146, 170

esora!30, 145, 170

eslebana 131

etesiae 146, 170

etia 122, 127, 151,169

euclea 85

Eueides 83

eucrasia 160

eunice 130, 144, 145, 170

Euphydryas 165

Euphydryini 165

Eurodryas 165

eutropia 130, 137, 138, 170

INDEX

evanescens 117
evanides 155
exclamationis 161, 167
extensa 129, 132, 169
extincta 109
ezorias 148

fallax 162
fasciatus 167
fassli 142, 143
fausta!52, 155, 170
felderi\5Q
fellula 84, 159, 170
ferghana 166
ferghana-group 166
flavia 122

flavida 122, 124, 169
flavocincta 1 10, 114, 169
fenestrata 143
fontus 108, 109, 168
fragilis 122, 124, 169
frisia83,95, 105, 168
fulgora!52, 155, 170
fulviplaga 95, 104, 168
fulvocincta 109

gabbii 167
gaudealis 167
gaujonillO, 114, 169
geminia 113
gentinal!6, 169
gillettii 165
gina 166
gloriosa 167
Gnathotriche 160, 167
Gnathotriche-group 167
Gnathotrusia 162, 167
gorgone 166
graphica92, 168
griseobasalis 156, 170
guatemalena 122, 124, 125, 170
guaya 156
gudruna 145

harperi 87
harrisii 167
heliconiformis 142
heliconina 148
heliconoides 153
hera84, 158, 170
herlani90, 168
hermas95, 104, 168
hewitsonii 148
Higginsius 163, 167
hilarina 153
hippodrome 167
hoffmanni 167
homogena 145
hopfferi 162
Hypodryas 165

INDEX

241

ianthe 158

iduna 165

ildica 142, 143

immaculata 123

infernalis 166

infrequens 122, 124, 169

intermedia Menetries 165

intermedia Rober 137

ithomioides 130, 137, 139, 140, 141, 170

ithomiola 131

ithral!6, 120, 169

jacinthica 134
janallO, 111, 169
janais 167
Janatella 157, 170
jemezensis 91
jezabel 166

klagesii 145
kuchi 166

lacinia 167
Iaiasl36, 170
lansdorfi 130, 134, 169
leanira 167
Ielex96, 168
letitia 129, 133, 169
leucodesma 158, 170
leucophaea 133
levana 155
levina 131, 150, 170
limbata 150
Iiriopell6, 169
lugubris 153
luka 122, 126, 169
lukto 166
lunulata 166
lutescens 122, 128
lycus 136

macarandica 166

macdunnoughi 87

macyi 89

magniplaga 143

malcolmi 167

manto 144

marcia 86

margaretha 130, 144, 170

marianna 167

marina 167

mata 89

maturna 165

may a 91

Mazia84, 159, 170

mechanitis 130,147, 170

mejicana 155

melaina!39, 170

melini 160

Melitaea 165

Melitaeini 165
melitaeoides 167
Mellicta 165
Mellicta-group 165
menetriesi 165
metharmeoides 160
mexicanus89, 168
microdryope 153
Microtia 167
mima 163
mimas 138, 170
mimetica 166
mimicry 144
minerva 165
minima 109
minuta 166
mirabilis 160
miriam 167
mixta 166

moesta 130,143, 170
montanus 85, 88, 168
morenus 108, 109, 168
morpheus 85
mundina 162, 167
murena 147, 148
myia!52, 156, 170
mylitta 149, 170
mylittusSS, 168

nanallO, 113, 169

narva 167

natalces 100

nauplius 129, 131, 132, 169

nazaria 122, 127, 169

nebrites 154

nebulosa 95, 101, 168

neptoides 133

neria 152, 154, 170

neumeogeni 167

nigra Giacomelli 120

nigra Rosenberg & Talbot 133

nigrella!22, 128, 169

nigrina 94

nigripennis 130, 141, 170

niveonotis 128

nivifera 139

nortbrundii 152, 154, 170

notusllO, 113, 169

nussiallO, 113, 169

nycteis 166

oblita!30, 135, 169
obscurata 102
occidentalis 152, 153, 170
Occidryas 165
ocellata!33, 169
ofella 152, 156, 170
olivencia 13 1,145, 170
omosis 113
orientalis 165

242

INDEX

orobia 122, 123, 169
orthial!6, 117, 169
orticasl!6, 117, 169
OrtiliallS, 169
orsa 88
orseis90, 168
otanes95, 103, 168

packardii 86
palla 167
pallas 165
pallescens91, 168
pallidus 89, 168
pardalina 147
parthenoides 165
pascoensis 86
pastazana 148
pastazena 125, 169
pedrona 93
pella 145
pellenea 153

pelonia 130, 147, 148, 170
pelops 167
peraea 162
perezi 170
perilla 152, 153, 170
perna!30, 149,150,170
persea 166
peruana 111
phaedima 143, 170
phaeton 83, 165
phaon83,85,91, 168
phillyra!30, 142, 148, 170
phlegias 95, 99, 168
phoebe 166

Phyciodes 79, 84, 85, 167
Phyciodini 81
Phystis 84, 93, 168
pictus85,90, 168
plagiata!29, 133, 169
platytenia 99
plotina 165
poecile 167

poecilina 130, 139, 170
pola 166
Poladryas 166
polina!30, 136, 170
polinellall6, 119, 169
poltis 117
polymnia 145
pratensis 87
pratti 94
prisca 143
proclea 167

pseudocelemina 141, 170
ptolyca 95, 96, 97, 168
pulchella 86
pullopicta 113
punctata 105

pusillus 108, 168
quintilla 130, 138, 170

reaghi 86
rebel! 165
reducta 131
regama 165
rima 160
riobalensis 167
roeberi 117
rohweri 87
romanovi 166
rosina 162
rosita 167
rufocincta 120

saladillensis 120, 121, 169
saltator 107
sarvistana 166
satura 134
saturata 134
saundersi 167
saxatilis 166
schaeferi 120
scotosia 166
sejonal!6, 118, 169
selene!31,151, 170
selenis 86
selenoides 128
seminole 100, 168
sepultallO, 115, 169
serpia 127, 169
sestia!30, 134, 169
shandura 166
sibina 166

signatallO, 112, 169
similis 122, 123, 169
simois93, 168
sindura 166
sitalces95, 107, 168
sodalis 161, 167
sopolis 93, 103, 168
sosis95, 106, 168
stenotaenia 104
stesilea 96
sticta!30, 140, 170
styx Staudinger 163
subfasciata 138
subconcolor 101, 168
subota98, 168
sulphurata 134
sutschana 166
sydra 102

taeniata95, 106, 168
tangigharuensis 166
taphius 1 14

Tegosa 83, 84, 121, 169
Telenassa 82, 84, 110, 168

INDEX 243

teletusa 110, 111, 168 ursula 122, 124, 169

texana 94, 95, 100, 168

Texolal67 vanessoides 161

tharos85,167 varia 165

tharossaS6 variegata 93, 94, 168

thebais89,168 velica 116, 118, 169

theona!67 verena97, 168

Thessahal67 vesta 85,92, 168

thymetusm Vestalis92

Tisona84,120,169 veternosalU

tissfl!27,128 virilis 152

tissoides 125, 169

transcaucasica 166

trimaculata 1 10, 114, 169 wernen 15°

tristis 87

trivia 166 yuenty 166

tulcis 95, 105, 168

turkmanica 1 66 zamora 1 1 6, 1 19, 1 69

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Barry Bolton

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

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This number completes Volume 43

ISSN 0524-643 1 Entomology series

Vol 43 No 4 pp 245-307
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 26 November 1981

-5

A revision of six minor genera of Myrmicinae
(Hymenoptera : Formicidae) in the Ethiopian
zoogeographical region

Barry Bolton

Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

1982

Contents

Synopsis .
Introduction

Measurements and indices
Abbreviations of museums
Ankylomyrma Bolton .
Atopomyrmex Andre .

Synonymic list of species

Key to species (workers)

Treatment by species
Baracidris gen. n.

Key to genera of Ethiopian region Myrmicinae with 2-segmented
antennal club (workers) .......

List of species .........

Key to species (workers)

Treatment by species
Cyphoidris Weber

List of species

Ket to species (workers)

Treatment by species
Ocymyrmex Emery

Synonymic list of species

Key to species (workers)

Treatment by species
Pristomyrmex Mayr .

Synonymic list of species

Key to species (workers)

Treatment by species
Terataner Emery

List of species

Key to species (workers)

Treatment by species

Synopsis of Malagasy region species

Key to species (workers)
Acknowledgements •

References

Index

246
246
246
247
247
249
250
250
250
252

253
255
255
255
256
257
257
257
260
262
262
264
282
283
283
284
288
290
290
291
296
297
298
298
307

Bull. Br. Mus. nat. Hist. (Ent.) 43(4): 245-307

Issued 26 November 1981

246 B. BOLTON

Synopsis

The ant genera Ankylomyrma Bolton, Atopomyrmex Andre, Cyphoidris Weber, Ocymyrmex Emery, Pri-
stomyrmex Mayr ( = Odontomyrmex Andre, = Hylidris Weber, = Dodous Donisthorpe) and Terataner
Emery ( = Tranetera Arnold) are revised for the Ethiopian zoogeographical region. Keys and descriptions of
species are presented for each genus and the genera are defined on a world-wide basis. In Atopomyrmex two
species are recognized and four new infraspecific synonyms are established. Three new species are described
in the previously monotypic genus Cyphoidris. Twenty-three species of Ocymyrmex are recognized of which
seven are described as new; seven new synonyms are established and new status as valid species is granted to
seven previously infraspecific forms. Five Pristomyrmex species are recognized of which one is new; five new
synonyms are proposed in this genus. In Terataner the former subgenus Tranetera is newly synonymized
and six species recognized, of which one is new. The six Terataner species of Madagascar are summarized,
one new species is described and a key presented. The genus Baracidris is described as new, containing two
new species from west and central Africa. A key to Ethiopian region myrmicine genera in which the antennal
club has two segments is given under Baracidris.

Introduction

This paper is presented as a contribution towards a revision of the subfamily Myrmicinae in the
Ethiopian zoogeographical region. The subfamily Myrmicinae is by far the largest and most
diverse in the region (and the world), containing some 43 genera in sub-Saharan Africa at present.
These genera can be categorized roughly as small (1-10 species), moderate (10-40 species) and
large (more than 40 species). The vast majority of Ethiopian region myrmicine genera fall into the
small category (29 genera), about nine genera come in the moderate category and only five reach
the final category, of which three (Tetramorium Mayr, Pheidole West wood and Crematogaster
Lund) are huge, with well over 100 species each in the region. A good indication of distribution of
genera on a world basis is given by Brown (1973), and a discussion of the most prevalent ant
genera is presented by Wilson (1976).

Of the seven genera discussed in this paper only Ocymyrmex, with 23 species, reaches the
moderate category; the rest, Ankylomyrma (1 species), Atopomyrmex (2 species), Baracidris (2
species), Cyphoidris (4 species), Pristomyrmex (5 species) and Terataner (6 species), are all small in
terms of number of species. Only two of the genera under consideration here have species which
occur outside of the Ethiopian region, Terataner and Pristomyrmex; the rest are peculiar to the
region. Terataner, apart from its six African species, has another six in the Malagasy region
whilst Pristomyrmex species are widely distributed in the Old World tropics, reaching a peak in
the Indo-Australian region.

Measurements and indices

Total Length (TL). The total outstretched length of the individual, from mandibular apex to
gastral apex.

Head Length (HL). The length of the head proper, excluding the mandibles, measured in a
straight line from the anteriormost point of the median clypeal margin to the mid-point of the
occipital margin, in full-face view. (In species with strongly concave occipital margin the head
length is measured to the mid-point of a line connecting the posterolateral projections.)

Head Width (H W). The maximum width of the head in full-face view, measured behind the eyes
except in Ankylomyrma (eyes in the posterior corners) and in Ocymyrmex (head often narrows
rapidly behind eyes) where HW is measured immediately in front of the eyes.

~ HW x 100

Cephalic Index (CI).

HL

SIX MINOR GENERA OF MYRMICINAE

247

Scape Length (SL). The maximum straight-line length of the antennal scape excluding the basal
constriction or neck close to the condylar bulb.

Scape Index (SI).

SL x 100
HW

Pronotal Width (PW). The maximum width of the pronotum in dorsal view. In Terataner PW is
measured behind the prominent angular or dentiform corners.

Alitrunk Length (AL). The diagonal length of the alitrunk in lateral view from the point at
which the pronotum meets the cervical shield to the posterior base of the metapleural lobes or
teeth.

All measurements are expressed in millimetres.

Abbreviations of museums

AMNH, New York

BMNH

CAS, San Francisco

IE, Bologna

IRSNB, Brussels

MCSN, Genoa

MCZ, Cambridge
MHN, Geneva

MNHN, Paris
MNHU, Berlin

MR AC, Tervuren

NM, Basle
NM, Bulawayo
USNM, Washington
ZM, Kiev

American Museum of Natural History, New York, U.S.A.

British Museum (Natural History), London, U.K.

California Academy of Sciences, San Francisco, California, U.S.A.

Istituto di Entomologia del'Universita, Bologna, Italy.

Institut Royal des Sciences Naturelles de Belgique,

Brussels, Belgium.

Museo Civico di Storia Naturale 'Giacomo Doria', Genoa,

Italy.

Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.

Museum d'Histoire Naturelle, Geneva, Switzerland.

Museum National d'Histoire Naturelle, Paris, France.

Museum fur Naturkunde der Humboldt-Universitat,

Berlin, Germany (D.D.R.).

Musee Royal de 1'Afrique Centrale, Tervuren, Belgium.

Naturhistorisches Museum, Basle, Switzerland.

National Museum, Bulawayo, Zimbabwe.

United States National Museum, Washington, D.C., U.S.A.

Zoological Museum, Institute of Zoology, Academy of

Sciences of Ukrainian, S.S.R., Kiev, U.S.S.R.

ANKYLOMYRMA Bolton

(Figs 1,2)

Ankylomyrma Bolton, 19736: 235. Type-species: Ankylomyrma coronacantha Bolton, loc.cit.; by original
designation.

DIAGNOSIS OF WORKER. Monomorphic arboreal myrmicine ants. Mandibles with 5 sharply defined large
triangular teeth, the mandibles almost entirely concealed by the clypeus when closed. Palp formula 5, 3, the
palpomeres very long (apical maxillary palpomere equal in length to apical antennal segment). Clypeus very
large, projecting forwards over the mandibles as a broad shelf. Median portion of clypeus raised, broad
posteriorly and broadly inserted between the widely separated frontal lobes. Frontal lobes concealing
antennal insertions, prolonged back by a pair of irregular frontal carinae which run past the inner margins
of the eyes to the occipital margin. Frontal carinae forming dorsal margins of a weak scrobe which is
bounded below by a ridge running from below the eye to the mandibular insertions. Eyes large, situated at

248 B. BOLTON

extreme posterolateral corners of head, within the scrobal area as defined by the frontal carinae and ridge.
Antennae 1 2-segmented, without a strongly defined club, the flagellomeres increasing in size apically.
Occipital margin bounded by a broad transverse lamella which projects into a series of dentiform processes ;
occiput behind the lamella broad and flat. Promesonotum swollen, large and convex, without sutures but
with 4 pairs of roughly triangular teeth or prominences. Propodeum short and strongly bispinose. Meta-
pleural lobes absent. Mesopleuron divided by a broad transverse suture and a broad suture separating
meso- and metapleuron present. Petiole strongly bispinose, with a broad anterior peduncle which grades
into the node. Gaster consisting almost entirely of the massively enlarged and strongly vaulted first tergite.
The first sternite is visible as a narrow collar fringing the ventral portion of the forward-pointing orifice of
the tergite; remaining gastral segments very small and telescoped inside. Sting strongly developed and
projecting anteriorly below the pedicel segments.

In the original description of this remarkable ant I placed the genus tentatively in the tribe
Meranoplini. I am aware now that this move was incorrect and that Ankylomyrma is not close to
Meranoplus or any other member of that now-disbanded tribe (for discussion see Bolton, 1981).
The real relationships of Ankylomyrma are an enigma for, although there are a number of charac-
ters implying alliance with genera such as Atopomyrmex, Terataner and their allies, such as low
dental count, high palp formula, broad clypeus and structure of petiole, there are also objections
to such a placement. Chief among these must be the position of the eyes, situated as they are at
the extreme posterior corners of the sides of the head and within what is strictly the scrobal area.
In Terataner and allies the eyes are always positioned well forward of the occipital corners and
below the scrobes when such are present. The incredible occipital fringe and unique gastral
development of Ankylomyrma are of course very derived characters which, though they serve to
isolate the genus, do nothing to indicate its relationships. The only known species is as follows.

Ankylomyrma coronacantha Bolton
(Figs 1,2)

Ankylomyrma coronacantha Bolton, 19736: 235, figs 1-3. Holotype worker, GHANA: Eastern Region, Mt
Atewa; by pyrethrum knockdown, sample A4/3, 12.vii. 1969 (D. Leston) (BMNH) [examined].

WORKER. TL 6-3-6-8, HL 1-48-1-50, HW (at maximum in front of eyes) 1-38-1-42, CI 92-96, SL 1-06-1-12,
SI 76-80, PW 1-20-1-22, AL 1-92- 1-95 (4 measured).

Mandibles delicately and superficially longitudinally striate, with scattered fairly conspicuous pits. Palpi
very long, the maxillary palps projecting beyond the posteroventral margin of the head when stretched out,
their total length c. 1-60. Anterior clypeal margin with a median impression. Main features of head as given
in generic diagnosis and Fig. 2. Eyes large, maximum diameter 0-36-0-38, about 0-27-0-28 x HW, situated
at posterolateral corners of head. Frontal carinae irregular, running inside eyes to occipital margin. Scrobal
area bounded below by a ridge which runs from the mandibular insertions to the extensive prominence at
the occipital corner, below the eye. Occipital margin with a transverse raised broad lamella which projects
into a number of teeth or spines, projecting especially strongly at the occipital corners. Promesonotum fused
and swollen, the dorsum much higher than the propodeum in profile. Metanotal groove absent. Pronotum
with three pairs of teeth or tubercles; dorsally with a pair of broad tubercles which are large and are seen to
be blunt or truncated in anterior view; dorsolaterally, just anterior to pro-mesonotal junction, with a pair of
broad acute, sometimes conical projections; humeral angles bluntly dentate, the teeth prominent in dorsal
view. Mesonotum at point where sclerite begins to slope down to the propodeum with a pair of broad, acute
teeth. Propodeal dorsum short, the entire dorsal face forming the base for a pair of long, stout spines. Apices
of tibiae each terminating in a pair of cuticular tooth-like outgrowths. Petiole strongly bispinose, in profile
the peduncle short and continuous with the dorsal surface of the node. Subpetiolar process a small tooth,
anteriorly situated. First gastral tergite massive, developed into a subspherical ball which has an opening
anteriorly, below the postpetiole. First sternite forms a narrow collar around the ventral portion of the
orifice so that almost all of the visible gaster consists only of the first tergite. Sting strong and projecting
anteriorly. Dorsum of head coarsely reticulate-rugose, the rugae strongly raised and enclosing broad foveo-
late spaces. Ground-sculpture of fine shagreening or superficial punctulation. This sculpture also present on

SIX MINOR GENERA OF MYRMICINAE 249

sides of head below the scrobes but the scrobal area itself merely densely reticulate-punctate. All pronotum
and raised dorsum of mesonotum sculptured as dorsum of head but the pleurae, the sloping portion of the
mesonotum and the propodeum with fine dense but superficial reticulation only. Petiole, postpetiole and
first gastral tergite coarsely foveolate, the interspaces finely reticulate or reticulate-punctulate. Dorsum of
head with numerous stout erect hairs which are also present, but sparser, on dorsum of promesonotum and
dorsal portion of first gastral tergite; the hairs are denser and finer ventrally on the first tergite but do not
occur on the sloping posterior half of the mesonotum, the propodeum or the petiole. Black with appendages
lighter, orange-brown to red-brown.

This large and very spectacular ant is arboreal, but beyond that nothing is known of its biology.

MATERIAL EXAMINED

Ghana: Mt Atewa (D. Leston). Cameroun: Korup (D. Jackson).

A TOPOMYRMEX Andre
(Figs 3-11)

Atopomyrmex Andre, 1889: 226. Type-species: Atopomyrmex mocquerysi Andre, op. cit.: 227; by monotypy.

DIAGNOSIS OF WORKER. Polymorphic arboreal myrmicine ants. Mandibles short and stout, the apical (mas-
ticatory) margin armed in smallest workers with 2 teeth followed by 2 denticles and an unarmed straight
edge; in slightly larger workers the edge crenulate or feebly denticulate. Most medium-sized and large
workers with 2 teeth + 4-5 denticles but in large workers all the teeth may be worn down and rounded. Palp
formula 4, 3 in all sizes; in smallest workers the two basalmost maxillary palp segments may be partially
fused. Median portion of clypeus shield-like, broad, posteriorly broadly inserted between the frontal lobes.
Anterior clypeal margin indented to concave medially, the median portion separated from the lateral parts
by a longitudinal carina on each side. Anterior tentorial pit represented by a deep and sharply incised hole
which is roughly circular, situated immediately behind the clypeus close to the antennal insertions and more
obvious in larger workers. Development of frontal carinae varying with worker size. In smallest workers
short and only feebly divergent, ending in front of the level of the anterior margins of the eyes. In largest
workers extending back beyond the level of the posterior margins of the eyes and strongly divergent from
source to level of eyes; behind this roughly parallel. Workers between largest and smallest showing inter-
mediate development of frontal carinae. Antennal scrobes absent in smallest workers, becoming longer and
deeper with increasing size; conspicuous and capable of accommodating the scape in largest workers.
Antennae 12-segmented with a 3-segmented club. With head in full-face view the eyes situated behind the
midlength of the sides, and the occipital corners broadly and evenly rounded. Pronotum more or less flat to
shallowly concave transversely, bluntly marginate laterally, the margination more acute in smaller workers.
Promesonotal suture vestigial to absent from dorsum but at sides forming an impression separating prono-
tum and mesonotum. Mesonotum in profile usually broadly and bluntly bituberculate behind, then sloping
almost vertically to the broad metanotal groove. In medium to large workers the mesonotum with a shallow
but quite broad transverse impression at about the midlength. Propodeum in profile raised immediately
behind the metanotal groove then sloping downwards to a pair of strong spines. Metapleural lobes incon-
spicuous, very narrow and low. Propodeal spiracle circular. Lower margin of metapleuron without a broad
groove running forward from the orifice of the metapleural glands; instead the margin rounded and folded
under, giving the appearance of being smoothly eroded away, the hind coxa appearing to rest upon the bulla
of the metapleural gland. Ventral surface of alitrunk with a very conspicuous roughly circular deep pit
between the hind coxae, a sieve-plate apparently present at the bottom of the pit. Petiole dorsally with a pair
of short stout spines of variable size. Dorsal surfaces of head and body without standing hairs, such hairs
present only on mouthparts and gastral sternites.

Atopomyrmex is a small genus of strongly polymorphic arboreal ants. The two species included,
mocquerysi and cryptoceroides, nest in the wood of standing trees and forage arboreally, fre-
quently coming down the trunk but only rarely venturing onto the ground. Arnold (1916) points
out that mocquerysi is usually carnivorous and, when disturbed, exudes a whitish secretion from
the anal glands. Nests are made in hollow stems or rotten parts of standing timber but it is not

250 B. BOLTON

known if the species tunnel their own galleries or take over the galleries of termites and boring
beetles. Of the two species mocquerysi is very widely distributed, being found in wooded and
forested areas almost throughout sub-Saharan Africa; Wheeler (1922: 181) gives a distribution
map. A. cryptoceroides has a more limited range, being confined to the rain-forest zones of west of
central Africa, where it is sympatric with mocquerysi.

The genus most closely related to Atopomyrmex is the Ethiopian and Malagasy genus Tera-
taner. Differences separating them, and other related genera, are noted in the discussion of
Terataner (p. 290). At its inception Atopomyrmex contained only mocquerysi. Soon afterwards
cryptoceroides was added and this was followed by a number of others, added later by several
authors. Later still these extra species were progressively removed from Atopomyrmex until the
present time, when mocquerysi and cryptoceroides are again its only members. These species,
originally described in Atopomyrmex but now placed elsewhere, are as follows. The species
alluaudi Emery, bottegoi Emery, foreli Emery, luteus Emery, scotti Forel, and steinheili Forel were
transferred to Terataner by Emery (1912); nodifier Emery was originally made type-species of
genus Atopula Emery but is now included in Tetramorium Mayr, see Bolton (1976; 1980);
selebensis Emery was made type-species of Dilobocondyla Santschi, by Santschi (1910); escherichi
Forel was transferred to Dilobocondyla by Forel (1913c); ceylonicus Emery was made type-species
ofParatopula Wheeler, by Wheeler (1919).

Synonymic list of species

cryptoceroides Emery

deplanatus Mayr

mocquerysi var. curvispina Forel

mocquerysi subsp. cryptoceroides var. melanoticus Wheeler (unavailable)
mocquerysi Andre

mocquerysi var. australis Santschi

mocquerysi var. obscura Santschi syn. n.

mocquerysi var. arnoldi Santschi syn. n.

mocquerysi si. opaca Santschi syn. n.

mocquerysi var. erigens Santschi syn. n.

mocquerysi st. opacus var. nigellus Santschi (unavailable).

Key to species (workers)

1 Sides of head behind eyes blanketed by a fine and very dense reticulate-puncturation. (Woodland

and forest zones throughout Africa) mocquerysi (p. 251)

Sides of head behind eyes smooth and glossy between widely separated small pits. (Rainforests of
West and Central Africa) cryptoceroides (below)

Treatment by species

Atopomyrmex cryptoceroides Emery
(Figs 9, 11)

Atopomyrmex cryptoceroides Emery, 1891 : 561, pi. 15, figs 5, 6. Holotype female, IVORY COAST: Assinie (C.
Alluaud) (MCSN, Genoa) [examined].

Atopomyrmex deplanatus Mayr, 1895: 113. Holotype worker, SIERRA LEONE: 'Riv. N' Gamie, Chutes de
Samlia(A MocguerysMIRSNB, Brussels). [Synonymy by Emery, 1899: 477.]

Atopomyrmex mocquerysi var. curvispina Forel, 1911: 311. Syntype workers, ZAIRE: Kondue (£. Luja)
(MHN, Geneva) [examined]. [Synonymy by Emery, 1922: 240.]

Atopomyrmex mocquerysi subsp. cryptoceroides var. melanoticus Wheeler, 1922: 182, fig. 44. Syntype wor-
kers, ZAIRE : between Lukolela and Basoko (H. 0. Lang) (BMNH) [examined]. [Name not available.]

WORKER. Answering to the description of mocquerysi and falling into the size range noted there; differing
from mocquerysi as follows.

SIX MINOR GENERA OF MYRMICINAE

251

cryptoceroides

Sides of head behind eyes smooth and shining be-
tween widely scattered pits, not blanketed by
reticulate-punctate sculpture and never with ru-
gulae in this area.

Propodeal dorsum strongly rugose, without or
only with vestiges of punctate sculpture.

Pronotal dorsum closely and coarsely rugose,
without dense punctate ground-sculpture.

Propodeal spines in dorsal view with their basal
portions projecting outwards before angling
backwards (Fig. 1 1), the projecting portion con-
cealing the spiracle which is not at all visible
from above.

Propodeal spiracle large and relatively close to the
margin of the declivity below the spine; diam-
eter of spiracle equal to or greater than the dis-
tance separating the spiracular hind margin
from the edge of the declivity at its closest point
(Fig. 9).

mocquerysi

Sides of head behind eyes blanketed by dense re-
ticulate punctate sculpture; sometimes rugulae
may also occur in this area.

Propodeal dorsum predominantly or entirely

reticulate-punctate, if rugulae occur they are

secondary to the punctation.
Pronotal dorsum usually with rugae present but

with punctate ground-sculpture which is usually

conspicuous and dense.
Propodeal spines in dorsal view directed more or

less evenly backwards, not projecting outwards

basally; the spiracle (or at least its annulus)

clearly visible from above (Fig. 10).

Propodeal spiracle smaller and some distance
away from the margin of the declivity below the
spine; diameter of spiracle less than the distance
separating the spiracular hind margin from the
edge of the declivity at its closest point (Fig. 8).

MATERIAL EXAMINED

Ghana: Tafo (G. S. Cotterelf); Tafo (Strickland); Mampong (D. Lesion); Kibi (D. Leston); Bunso
(D. Leston); Osenasi (D. Leston); Asamankese (D. Leston); Etukrom (D. Leston); Mpraeso (D. Leston).
Zaire: Ituri For., Beni (T. H. E. Jackson).

Atopomyrmex mocquerysi Andre
(Figs 3-8, 10)

Atopomyrmex mocquersyi Andre, 1889: 227. Syntype workers, SENEGAL: Dakar (A. Mocquerys) (MNHN,

Paris) [examined].

Atopomyrmex mocquerysi var. australis Santschi, 1914a: 16. Syntype workers, SOUTH AFRICA: Natal, Zulu-
land (/. Tragardh) (NM, Basle) [examined]. [Synonymy by Wheeler, 1922: 885.]
Atopomyrmex mocquerysi var. obscura Santschi, 1923: 283. Syntype workers, IVORY COAST: Jacqueville

(Lohier); and BENIN: Cotonou (Silvestri) (MRAC, Tervuren; NM, Basle) [examined]. Syn. n.
Atopomyrmex mocquerysi var. arnoldi Santschi, 1923: 283. Syntype workers, ZAIRE: Eala (R. Mayne) (NM,

Basle; MRAC, Tervuren) [examined]. Syn. n.
Atopomyrmex mocquerysi st. opaca Santschi, 1923: 283. Syntype workers, ANGOLA : 'Riviere Cubia, entre

Combo et Cubra' (Rohan-Chabot) (MRAC, Tervuren; NM, Basle) [examined]. Syn. n.
Atopomyrmex mocquerysi var. erigens Santschi, 1924: 205. Syntype workers, ZAIRE: Yambata (Di Giorgi)

(NM, Basle; MRAC, Tervuren) [examined]. Syn. n.
Atopomyrmex mocquerysi st. opacus var. nigellus Santschi, 1930: 72. Syntype workers, ANGOLA: Rio Mbale

and Chimporo (A. Monard) (NM, Basle) [examined]. [Name not available.]

WORKER. Standard measurements are obviously not of great value where continuously polymorphic species
are involved, as one size grades into another without any break. However, when graphs of the relationships
of various dimensions are plotted a number of allometric relationships become apparent. Most easily
noticeable of these are the following. The CI increases with increase in HW; the relative lengths of the scapes
(SI) decrease as HW increases; the frontal carinae increase in length and strength as HW increases. The size
of the eye has little or no dependence on the size of the head, the eyes of the largest workers being relatively
only slightly increased in size over those of the smallest (as expressed by the ratio of ocular diameter to HW).
Overall size range in the species is TL 4-0-8-7, HL 0-96-2-24, HW 0-88-2-30, CI 92-104, SL 0-70-1-22,
SI 49-80, PW 0-68-1-40, AL 1-34-2-50 (85 measured).

Basic characters as given under generic diagnosis, differentiation from cryptoceroides as tabulated above.
Mandibles pitted, the surface between the pits finely and densely shagreened to finely striate. Frontal carinae
increasing in length and strength with increasing worker size (Figs 5-7). In smallest workers short, ending in

252 B. BOLTON

front of the level of the anterior margins of the eyes, forming a short and narrow laterally projecting flange
on each side and only very slightly divergent. In larger workers the frontal carinae lengthening and
becoming more obviously divergent, the laterally projecting flange broadening. In largest workers the
carinae reaching back beyond the level of the eyes, divergent to eye level then becoming almost parallel, the
laterally projecting flange very prominent throughout most or all of the length of the carinae. Antennal
scrobes absent in small workers, becoming better defined with increased size; in large workers the scrobe
conspicuous, narrow but quite deep and capable of accommodating the scape. Change in head shape with
increased size as in Figs 5-7. Maximum diameter of eye 0-18-0-36, about 0-14-0-20 x HW, the relative size of
the eyes not radically increased in larger workers. Outline shape of alitrunk as in Figs 3, 4. Propodeal spines
very variable in length, thickness and degree of curvature. In dorsal view the spines not projecting outwards
in their basal portions, the propodeal spiracle or at least its annulus visible from above. In profile the
propodeal spiracle some distance away from the margin of the declivity below the spine, the diameter of the
spiracle less than the distance separating it from the margin of the declivity. Petiolar teeth conspicuous,
varying in length and thickness. Sculpture in general increasing in intensity and frequently also in density
from smaller to larger workers. Dorsum of head with scattered shallow pits, the surface between them
varying from smooth or almost smooth to densely reticulate-punctate. This ground-sculpture is overlaid
between the frontal carinae by fine dense longitudinal striation. As the frontal carinae increase in length so
the area of striate sculpture becomes stronger and extends further back on the head. In larger specimens the
space between the frontal carinae becomes strongly rugose or costate and this sculpture may reach back
almost to the occipital margin. Sides of head densely reticulate-punctate everywhere. With increasing size
there is a tendency for the progressive encroachment of rugulose sculpture across the reticulate-punctate
surface from the front to the back of the sides. Small workers have only the punctate sculpture but as size
increases rugulae appear anteriorly which gradually strengthen and spread further back on the head.
Pronotal dorsum longitudinally rugose at least centrally, the rugae varying in intensity and sometimes
divergent posteriorly. Ground-sculpture reticulate-punctate and usually distinct, sometimes faint and fre-
quently with larger superimposed punctures present. Pronotal sculpture continuing onto mesonotum in
smaller workers, but in larger individuals (and also in some smaller ones) the sculpture becomes reduced on
the mesonotum so that only the punctate ground-sculpture is present or rugae occur but are restricted to the
anterior part of the sclerite. In large workers there is usually a striking reduction in mesonotal sculpture so
that most or all of the dorsum is feebly punctulate or even smooth and shining. Propodeal dorsum densely
reticulate-punctate, usually without trace of rugulose sculpture but sometimes with one or two weak rugulae
present. Petiole, postpetiole and first gastral tergite finely and very densely reticulate-punctate to densely
shagreened. Dorsal surfaces of body without standing hairs of any description. Colour very variable, ranging
from dull yellowish brown to blackish brown but most commonly bicoloured, with head and alitrunk
reddish, gaster dark brown to black. In some the head alone reddish and the rest of the body darker, in
others the head and gaster dark and the alitrunk lighter.

MATERIAL EXAMINED

Ghana: Tafo (B. Bolton); Mampong (P. Room); Bunso (D. Lesion); Wiawso (D. Lesion); Legon
(D. Lesion); Mepom (D. Lesion); Okumaning (D. Lesion). Nigeria: Gambari (B. Bolton); Mokwa (C. Long-
hursl). Cameroun: Nkoemvon (D. Jackson). Zaire: Yangambi (N. L. H. Krauss); Lukolela to Basoko
(H. O. Lang). Sudan: Keilak (R. C. H. Sweeny). Kenya: Kibwesi (S. A. Neave). Uganda: Masindi (R. Lucius).
Tanzania: Kilossa (S. A. Neave); Morogoro (A. Loveridge); Zanzibar (W. M. Aders); Zanzibar
(L. F. Brown); Duthumi (A. Loveridge); Kigoma Reg., Mahale Mts. (S. Uehara). Zambia: N'Changa
(C. T. Macnamara); Mwengwa (Dollman). Malawi: Port Herald (J. E. S. Old). Zimbabwe: Umgusa Riv.,
Sipopoma (G. Arnold); Victoria Falls (G. Arnold).

BARACIDRIS&n.n.

(Figs 12-14)
Type-species : Baracidris meketra sp. n.

DIAGNOSIS OF WORKER. Minute monomorphic myrmicine ants. Outline shape of head as in Figs 13, 14.
Mandibles narrow, the apical (masticatory) margin armed with 5 teeth which decrease in size from apical to
basal, the two basalmost teeth small. Basal borders of mandibles unarmed; the mandibles enclosing a space
between their basal borders and the anterior clypeal margin when their apical margins are overlapping. Palp
formula 2, 2, the first maxillary palpomere small (worker and queen of meketra dissected). Median portion of
clypeus sharply raised centrally and in the form of a narrow longitudinal ridge which runs from the anterior
margin to the frontal lobes. Anterior clypeal margin projecting medially (meketra) or in the form of a

SIX MINOR GENERA OF MYRMICINAE

253

truncated lobe (sitra). Lateral portions of clypeus unmodified, not prominent not raised into a wall in front
of the antennal insertions. Frontal lobes small, very closely approximated, almost touching anteriorly and
separated only by the extremely narrow strip of the median clypeus which is inserted between them. The
frontal lobes end immediately behind the antennal fossae; frontal carinae and antennal scrobes are absent.
Antennae with 12 segments, the flagellum ending in a strong 2-segmented club of which the apical segment is
much the larger. Scapes short, when laid back on the head conspicuously failing to reach the occipital
margin. Eyes minute and inconspicuous, situated approximately at the midlengths of the sides of the head.
Alitrunk and pedicel segments as in Fig. 12. Promesonotum fused and forming a single long shallow
convexity in profile. Metanotal groove impressed. Propodeum short, much shorter than the promesonotum,
armed with a pair of blunted, broad but short, triangular teeth. Metapleural lobes very broad and rounded,
strongly prominent, linked to the propodeal teeth above by a lamella. Petiole nodiform with a short, thick
anterior peduncle; in dorsal view the peduncle about as broad as the node. Postpetiole with a strongly
projecting blunt or truncated ventral process in profile; cylindrical in dorsal view. Standing hairs absent
from dorsal surfaces of head and body, present only on the clypeus and gastral apex.

Known from two species, the minute ants of the genus Baracidris seem to be endemic in the wet
forest zones of West and Central Africa where they inhabit the leaf-litter and topsol layers. The
genus most closely related to Baracidris is Adelomyrmex Emery whose known distribution in-
cludes the Neotropics, New Guinea, Fiji and Samoa. One of the African species was wrongly
referred to Adelomyrmex in an earlier publication (Bolton, 1973a). Adelomyrmex shares a number
of diagnostic characters with Baracidris, particularly in having similarly constructed antennae of
12 segments ending in a large 2-segmented club. Differences separating the genera may be
tabulated as follows.

Baracidris
Basal border of mandibles unarmed.

Maxillary palp 2-segmented.

Median strip of clypeus raised into a narrow ridge;

anterior clypeal margin unmodified or with a

simple lobe.

Hairs absent from dorsal surfaces of head and

body.
Postpetiole short-cylindrical in dorsal view, with a

large truncated ventral process.
Petiole low in profile.

Metapleural lobes very large and continuous with

the propodeal teeth above.
Range: West & Central Africa.

Adelomyrmex

Basal border of mandibles with a tooth at or prox-
imal to the midlength of the border.

Maxillary palp 1 -segmented (Gotwald, 1969).

Median portion of clypeus swept upwards into a
strongly raised sharp-edged longitudinal plat-
form which projects sharply forwards into a
lobe; anterior clypeal margin sweeping down-
wards and outwards away from and behind the
apex of this lobe.

Hairs present on dorsal surfaces of head and body.

Postpetiole not short-cylindrical in dorsal view, us-
ually without a truncated ventral process.

Petiole usually high and narrow in profile, only
rarely low.

Metapleural lobes small, separated from the pro-
podeal spines above.

Range: Neotropics, New Guinea, Fiji, Samoa.

Among the Myrmicinae of the Ethiopian zoogeographical region Baracidris is unique in pos-
sessing 1 2-segmented antennae with a 2-segmented club. This character, coupled with the very
closely approximated frontal lobes with the median clypeus narrowly inserted between them, the
short 5-dentate mandibles, reduced palp formula of 2, 2 and the shape of the pedicel segments,
renders Baracidris quickly recognizable. The key presented below will separate the myrmicine
genera of the region which have a conspicuously 2-segmented antennal club. Crematogaster is
included as a few species have such a club although the vast majority of species in this genus have
the club 3-segmented.

Key to genera of Ethiopian region Myrmicinae with 2-segmented antennal club (workers)

1 Mandibles elongate and linear, produced into long narrow projecting blades; never triangu-
lar/subtriangular, never serially dentate .......

Mandibles triangular or subtriangular, not produced into long narrow projecting blades ; apical
(masticatory) margins usually serially dentate but teeth sometimes reduced .... 6

254 B. BOLTON

2 Apex of each mandibular blade armed with a fork of 2-3 spiniform teeth set in a more or less

vertical series, with or without denticles between these teeth * 3

Apex of each mandibular blade either with a single long tooth at the dorsal apex subtended by a
series of denticles, or with a series of denticles only 5

3 Apical fork of mandibles with 3 spiniform teeth. Blades of mandibles without preapical teeth.

Petiole node with a pair of teeth or short spines. Antennal scrobes absent

MICRODACETON Santschi

Apical fork of mandibles with 2 spiniform teeth. Blades of mandibles with preapical teeth.
Petiole node unarmed. Antennal scrobes present 4

4 Antennae with 4 segments. Head with large orbicular hairs present. Mandibles very strongly

bowed, with a single preapical spiniform tooth .... QUADRISTRUMA Brown
Antennae with 6 segments. Head without orbicular hairs. Mandibles not strongly bowed, with
preapical armament of 1-2 teeth or denticles STRUMIGENYS F. Smith

5 Head with large orbicular hairs present; clypeal margin with spatulate or strap-like projecting

hairs. Antennal scape with a broad anteriorly projecting lobe. Head shield-like and broad

EPITRITUS Emery

Head with simple hairs; clypeal margin without spatulate or strap-like projecting hairs. Anten-
. nal scape linear, without broad anteriorly projecting lobe. Head not shield-like, longer than

broad CLADAROGENYS Brown

6 Antennae with 4-6 segments

Antennae with 8- 12 segments .12

7 Antennal scrobes absent. Propodeum unarmed, smoothly rounded. Basal tarsal segment on

each leg (but especially the first) very strongly swollen . . . MELISSOTARSUS Emery
Antennal scrobes present above the eye. Propodeum bidentate or bispinose. Basal tarsal seg-
ment on each leg slender 8

8 Clypeus very large, covering most of mandibles so that only their apices are visible in full-face

view, the mandibles thus appearing very short. Dorsal surfaces of head capsule and body
hairless but specialized hairs present on clypeus and scapes . . . M1CCOSTRUMA Brown
Clypeus smaller, not covering most of mandibles; in most cases merely covering the basal
borders so that the whole length of the mandible is visible in full-face view. Dorsal surfaces of
head and body usually conspicuously hairy 9

9 Apical margin of mandible with > 20 denticles, the basal 4-8 of which may be enlarged

SERRASTRUMA Brown

Apical margin of mandible with < 10 teeth of variable size .... 10

10 With mandibles fully closed their basal borders separated from the anterior clypeal margin by a

conspicuous impression or gap. Cephalic hairs reduced to 2-3 on posterior dorsum

TR1CHOSCAPA Emery

With mandibles fully closed their basal borders contiguous with or overlapped by the anterior
clypeal margin. Cephalic hairs numerous ..........

1 1 Mandibles very strongly convex and downcurved, armed with relatively few, strongly devel-

oped, stout teeth COD1OMYRMEX Wheeler

Mandibles not strongly convex nor downcurved, armed with numerous small spiniform teeth or
denticles . SMITHISTRUMA Brown

12 Antennae with 8-9 segments

Antennae with 10- 12 segments 15

13 Propodeum bidentate, bispinose or sharply angled. Dimorphic species

OL1GOMYRMEX Mayr (part)

Propodeum unarmed. Monomorphic species .......... 14

14 Eyes absent. Mandibles with 5-6 teeth. Promesonotum not marginate laterally

CAREBARA Westwood

Eyes present. Mandibles with 4 teeth. Promesonotum marginate laterally . . PAEDALGUS Forel

15 Postpetiole articulated on dorsal surface of first gastral segment. Petiole dorsoventrally flattened

and without a node CREMA TOG ASTER Lund

Postpetiole articulated on anterior surface of first segment. Petiole with a node of some form . 1 6

16 Propodeum unarmed and rounded. Antennae with 10 segments . . SOL ENOPSIS Westwood
Propodeum bidentate or bispinose. Antennae with 10-12 segments 17

17 Frontal lobes very close together so that the portion of the clypeus running between them is

extremely narrow, reduced to a line. Antennae with 12 segments . . . BARACIDRIS gen. n.
Frontal lobes not very close together, the portion of the clypeus running between them not

reduced to a line. Antennae with 10-1 1 segments . . . . . . . . • 18

SIX MINOR GENERA OF MYRMICINAE 255

18 Clypeus bicarinate. Dimorphic species without intermediates . . OLIGOMYRMEX Mayr (part)
Clypeus not bicarinate. Polymorphic species PHEIDOLOGETON Mayr

List of species
meketra sp. n.
sitra sp. n.

Key to species (workers)

1 Central portion of anterior clypeal margin not produced into a narrow truncated lobe (Fig. 13).
Occipital margin in full-face view indented or concave medially. Head slightly narrower and
antennal scapes relatively longer, CI 80-82, SI 75-79. (Ivory Coast, Ghana, Nigeria)

meketra (p. 255)

Central portion of anterior clypeal margin produced into a narrow truncated lobe (Fig. 14).
Occipital margin in full-face view not indented or concave medially. Head slightly broader
and antennal scapes relatively shorter, CI 86, SI 70. (Gabon) sitra (p. 256)

Treatment by species

Baracidris meketra sp. n.

(Figs 12, 13)

HOLOTYPE WORKER. TL 1-8, HL 0~44, HW 0-36, CI 82, SL 0-28, SI 78, PW 0-26, AL 0-50.

Mandibles sculptured with fairly large scattered pits, shining, without striate sculpture. Apical tooth of
mandible large, the second smaller, the remaining three teeth very small. Anterior clypeal margin projecting
forwards medially but not forming a narrow truncated lobe. Structure of clypeus and frontal lobes as given
in generic diagnosis. Antennal scapes short and stout (SI 75-79 in entire type-series), when laid back on the
head distinctly failing to reach the occipital margin. First funicular segment large, longer than broad;
segments 2-9 of funiculus annular and distinctly broader than long. The two apical funicular segments
which form the strong club dissimilar in size, the preapical much shorter than the apical (0-05 as opposed to
0-22). Eye minute, with a single ocellus of diameter c. 0-015 — 0-020, approximately 0-05 x HW. With head
in full-face view the occipital margin indented or slightly concave medially. Promesonotum forming a single
long shallow convexity in profile, the metanotal groove impressed. Propodeal dorsum in profile feebly
convex and sloping posteriorly to the short but quite broad triangular teeth. The propodeal teeth are joined
by a lamella below to the enlarged, strongly prominent and very broadly rounded plate-like metapleural
lobes, the two together forming an efficient shield all down the propodeal declivity. Propodeal spiracle
round. With alitrunk in dorsal view the promesonotum three times longer than the propodeum (0-32 and
0-10 respectively), the pronotal shoulders broadly rounded. Propodeal dorsum terminating posteriorly in a
sharply defined arch between the bases of the laterally flattened teeth, the declivity almost vertical and
bounded on each side by the very prominent teeth and metapleural lobes, and the lamella which links them.
A single small carina traverses the declivity at the level of the propodeal teeth. Petiole in profile with a short,
thick anterior peduncle which is distinctly shorter than the length of the node; the peduncle with a small
anteroventral process. Postpetiole in profile with the sternite produced into a large truncated ventral
process. In dorsal view the petiole node as long as broad (c. 0-12), the anterior peduncle broader than long
and much shorter than the dorsal length of the node; the node itself scarcely broader than the peduncle.
Postpetiole dorsally slightly broader than long, about equal in length to the petiole node and parallel-sided,
appearing short-cylindrical. Base of first gastral tergite concave at the postpetiolar articulation. Dorsum and
sides of head with small, close-packed foveolate punctures, the spaces between which are smooth and
shining. Spaces between punctures usually smaller than the diameter of the punctures. On the sides of the
head above and behind the eyes there is a tendency for the punctures to be aligned. Dorsal alitrunk similarly
sculptured but the punctures less conspicuous and more widely scattered, with extensive smooth shining
spaces between them. Sides of alitrunk and all surfaces of pedicel segments punctate, denser on the meta-
pleuron than elsewhere on the alitrunk and denser on the sides of the petiole and postpetiole than dorsally.
First gastral tergite unsculptured except for scattered minute punctulae. Hairs absent except on clypeus,
mouthparts and gastral apex. All dorsal surfaces of head and body thinly and sparsely clothed with
extremely fine short appressed pubescence. Colour uniform light brown, the appendages lighter in shade
than the body.

PARATYPE WORKERS. TL 1-7-1-8, HL 0-44-0-47, HW 0-36-0-38, CI 80-82, SL 0-27-0-30, SI 75-79,
PW 0-26-0-28, AL 0-49-0-50 (4 measured). As holotype but the Ghana specimens slightly darker in colour

256 B. BOLTON

than those from Nigeria and with the cephalic sculpture somewhat more sharply defined. Propodeal teeth
vary slightly in length and width but are always shorter than the metapleural lobes. One or two fine
transverse carinae may be present on the propodeal declivity, between the teeth.

PARATYPE FEMALE (dealate). TL 2-0, HL 0-46, HW 0-39, CI 85, SL 0-28, SI 72, PW 0-30, AL 0-56. As
worker but eyes larger (maximum diameter 0-10, c. 0-26 x HW) and with ocelli present. Alitrunk with full
complement of flight sclerites, the pronotum forming a clearly visible collar anteriorly in dorsal view.
Mesoscutellum much higher than and somewhat overhanging the propodeal dorsum. Mesoscutum and
scutellum with foveolate punctures and a smooth unsculptured median longitudinal strip; otherwise as
holotype.

Holotype worker, Nigeria : Ile-Ife, 2 l.vii. 1971(7. T. Medler)(EMNH).

Paratypes. Nigeria: 1 worker with same data as holotype. Ghana: 1 worker and 1 dealate female, Tafo,
4.ix.l970, cocoa litter sample (B. Bolton). Ivory Coast: 1 worker, Abidjan, Banco Forest, i.1963 (W. L.
Brown) (BMNH ; MCZ, Cambridge).

All specimens originated in forest; those from Ghana were extracted from a sample of cocoa leaf
litter which had built up between the roots of a large forest tree, left in the plantation to provide
shade for the cocoa.

Baracidris sitra sp. n.

(Fig. 14)

HOLOTYPE WORKER. TL 2-0, HL 0-51, HW 0-44, CI 86, SL 0-31, SI 70, PW 0-31, AL 0-56.

Answering to the description of meketra above as regards general shape, sculpture etc., but differing as
follows.

Anterior clypeal margin produced into a short truncated simple lobe. Occipital margin of head in full-face
view transverse or exceedingly feebly convex, not indented or concave medially. Larger, more stockily built
species with relatively broader head and shorter arrtennal scapes — compare standard measurements. Apart
from these the lengths of apical and preapical funicular segments in sitra are 0-26, 0-07 respectively, and the
dorsal lengths of promesonotum and propodeum are 0-35, 0-12 respectively. Petiole node in dorsal view
slightly broader than long (0-13 by 0-10). Propodeal dorsum more strongly convex and the lamella linking
the teeth to the metapleural lobes broader, minimum width of the lamella 0-06 (as opposed to 0-03 in
meketra holotype).

Holotype worker, Gabon: Plateau d'Ipassa, VM9, IPA7 (J. A. Barra) (MCZ, Cambridge).

The two minute species of this genus are closely related, but the first and second characters noted
above, plus the differences in dimensions, serve to separate them. Like most of the meketra
material the holotype and only known specimens of sitra seems to come from a leaf-litter sample.

CYPHOIDRIS Weber

(Figs 15-17)
Cyphoidris Weber, 1952: 26. Type-species: Cyphoidris spinosa Weber, loc. cit.; by original designation.

DIAGNOSIS OF WORKER. Monomorphic myrmicine ants. Mandibles triangular with an elongate apical (mas-
ticatory) margin bearing 10-14 small teeth or denticles which decrease in size from apex to base. Palp
formula 4, 3. Median portion of clypeus narrow and raised, bicarinate above and narrowly inserted between
the frontal lobes. Lateral portions of clypeus unmodified, not forming a shield-wall or raised ridge in front of
the antennal insertions. Frontal lobes not strongly expanded but covering the antennal insertions, pro-
longed posteriorly as a pair of strongly developed frontal carinae which form the dorsal margins of a pair of
strong and conspicuous broad scrobes; the scrobes run back almost to the occiput. Ventral margin of scrobe
a longitudinal ridge or ruga running above the eye, the latter of moderate size and situated in front of the
midlength of the sides. Antennae 11 -segmented, with a conspicuous 3-segmented club apically. Alitrunk in
profile with promesonotum fused and swollen, the dorsum dome-like and strongly convex in outline, much
elevated above the level of the propodeum. Propodeum bispinose; the spiracle close to the margin of the
declivity, the orifice circular and directed posteriorly. Metapleural lobes low and triangular. Petiole with an
elongate anterior peduncle and well developed node. Sting terminating in a narrow spatulate appendage
apically.

SIX MINOR GENERA OF MYRMICINAE 257

An easily defined genus, Cyphoidris is the only African representative of a group of genera
centring on Lordomyrma Emery, most of which have an Indo-Australian or Neotropic distri-
bution. Cyphoidris is close to Lordomyrma itself but differs as the latter has 12-segmented an-
tennae, a reduced palpomere count, and has the propodeal spiracle set well forward from the
margin of the declivity. Of the known species of Cyphoidris, exalta and spinosa are of Central
African origin and inhabit the leaf litter layer; parissa originates in West Africa, and werneri is the
only known East African representative of the genus.

List of species
exalta sp. n.
parissa sp. n.
spinosa Weber
werneri sp. n.

Key to species (workers)

1 Basal half of first gastral tergite densely and strongly shagreened. (Liberia) . . . parissa (p. 258)
Basal half of first gastral tergite unsculptured except for hair-pits 2

2 Propodeal dorsum flat in profile, not continuing curve of mesonotum. Dorsal alitrunk not

strongly densely reticulate-rugose everywhere 3

Propodeal dorsum in profile continuing curve of mesonotum. Dorsal alitrunk strongly densely
reticulate-rugose everywhere. (Rwanda) werneri (p. 259)

3 Dorsal alitrunk with abundant conspicuous standing hairs (Fig. 15). (Ivory Coast, Zaire, Angola)

. spinosa (p. 257)
Dorsal alitrunk with inconspicuous short decumbent hairs (Fig. 17). (Cameroun) exalta (p. 258)

Treatment by species

Cyphoidris spinosa Weber
(Figs 15, 16)

Cyphoidris spinosus Weber, 1952: 26, figs 7, 8. Holotype worker, ZAIRE: Ituri Forest, 15 miles [24 km] N. of
Beni, 25.ii.1948, no. 2129-2 (N. A. Weber) (AMNH, New York) [examined].

WORKER. TL 3-8-4-3, HL 0-90-0-98, HW 0-81-0-91, CI 88-93, SL 0-68-0-76, SI 83-86, PW 0-60-0-72,
AL 1-08-1-22 (10 measured).

Mandibles smooth with scattered pits or at most with faint traces of fine longitudinal striation; number of
small teeth on mandible varying from 10-12. Anterior clypeal margin arcuate or with a very shallow
impression medially, the median portion of the clypeus raised and bicarinate longitudinally. Main features
of head as in generic diagnosis and Fig. 16. Eyes of moderate size, maximum diameter 0-16-0-18, about 0-20-
0-22 x HW, situated below the ventral margin of the scrobe and in front of the midlength of the sides of the
head. Frontal carinae sharply defined, relatively close together and diverging slightly posteriorly but in
general not becoming radically broader than the distance across the frontal lobes. At eye level the separation
of the frontal carinae is 0-36-0-42, about 0-44-0-46 x HW.

Outline of alitrunk as in Fig. 15, the promesonotum conspicuously swollen and on a much higher level
than the propodeal dorsum. Propodeum armed with a pair of strong spines which are straight to feebly
upcurved. Metanotal groove absent; metapleural lobes triangular and conspicuous. Petiole in profile with
an elongate peduncle and a well developed node, the dorsum of the node sloping downwards posteriorly so
that the posterior face is short. Anterior and dorsal surfaces of postpetiole forming a single convexity, the
posterior face truncated. In dorsal view the nodes of both the petiole and postpetiole broader than long.
Dorsum of head strongly longitudinally rugose, with 5-7 rugae between the frontal carinae at eye-level. The
rugae are irregular and tend to meander slightly, a few anastomoses usually being present. Scrobal area for
the most part smooth but usually with 1-2 fine rugulae traversing the width of the scrobe behind the antennal
fossa. Sides of head behind eyes finely reticulate-rugose, this sculpture extending round the posterior
margins of the scrobes to the occiput. Spaces between rugae everywhere on head unsculptured or at most
with the faintest superficial traces. Dorsal alitrunk everywhere finely but strongly reticulate-rugose with
broad shining interspaces. Petiole dorsum rugulose, the postpetiole varying from rugulose to only faintly
sculptured. Caster unsculptured. All dorsal surfaces of head and body densely clothed with fine acute hairs.
Colour dark reddish brown to blackish brown.

258

B. BOLTON

C. spinosa, known only from Zaire, Ivory Coast and Angola, is a leaf litter inhabiting species of
the forest zone. The closest related species of the genus, exalta, is compared with spinosa below.

MATERIAL EXAMINED
Angola: Duque de Braganca Falls (P. M. Hammond). Ivory Coast: Agboville(7. Lobl).

Cyphoidris exalta sp. n.

(Fig. 17)

HOLOTYPE WORKER. TL 4-3, HL 0-96, HW 0-88, CI 92, SL 0-74, SI 84, PW 0-64, AL 1-16.

Answering to the description of spinosa in general characters but differing markedly in sculpture and
pilosity, as follows.

exalta

Sides of pronotum smooth.
Promesonotal dorsum weakly and predominantly

transversely rugulose, with few meshes.
Postpetiole in dorsal view unsculptured.
Occipital corners without long fine hairs.

Occipital margin and sides of head behind eyes

with short, curved, decumbent to appressed

hairs.
Dorsal margins of frontal carinae without a spaced

row of long curved hairs; such hairs also absent

elsewhere on head where only very short, curved

pilosity is present.
Dorsal surfaces of alitrunk, petiole and postpetiole

with inconspicuous short, curved decumbent

hairs (Fig. 17).
First gastral tergite with short curved hairs.

spinosa

Sides of pronotum reticulate-rugose.
Promesonotal dorsum strongly and conspicuously

reticulate-rugose.

Postpetiole in dorsal view sculptured.
Occipital corners each with a single long fine hair

which is prominent and conspicuous.
Occipital margin and sides of head behind eyes

with projecting curved hairs.

Dorsal margins of frontal carinae with a spaced
row of long curved hairs, such hairs also present
elsewhere on head and projecting freely above
the level of the shorter ground-pilosity.

Dorsal surfaces of alitrunk, petiole and postpetiole
with conspicuous long standing hairs (Fig. 15).

First gastral tergite with elongate projecting hairs.

PARATYPE WORKER. TL 4-2, HL 0-95, HW 0-85, CI 89, SL 0-74, SI 87, PW 0-62, AL 1-12. As holotype.
Holotype worker, Cameroun. Korup Reserve, 14.ii.1980, in rotten log (D. Jackson) (BMNH).
Paratype. 1 worker with same data as holotype (MCZ, Cambridge).

Cyphoidris parissa sp. n.

HOLOTYPE WORKER. TL 3-7, HL 0-84, HW 0-76, CI 90, SL 0-62, SI 82, PW 0-58, AL 1-00.

Mandibles smooth with scattered small pits, the apical margin armed with 1 1-12 low denticles. Anterior
clypeal margin convex but medially slightly flattened and apparently with a minute median indentation.
Median portion of clypeus raised and the raised section bicarinate above. Frontal carinae strongly devel-
oped, running back beyond the level of the eyes but fading out well in front of the occipital margin, forming
the dorsal borders of the broad but shallow conspicuous scrobes. Frontal carinae slightly diverging po-
steriorly, their separation at the level of the midlengths of the eyes only c. 0-30, about 0-40 x HW. Eyes
distinctly longer than wide, their maximum diameter 0-18, about 0-24 x HW. With the head in full-face view
the sides rounding broadly and evenly into the occipital margin, without trace of an occipital corner; the
occipital margin itself evenly shallowly convex, not impressed or concave medially. Alitrunk in profile with
the promesonotum strongly swollen and dome-like, much higher than the surface of the propodeum; the
latter sloping posteriorly to a pair of narrow spines which are very feebly sinuate along their length.
Metapleural lobes short-triangular and acute. Petiole node in profile with the anterior face higher than the
posterior so that the dorsal surface slopes downwards posteriorly; posterodorsal angle of node more obtuse
than anterodorsal angle. Postpetiole with a sharp anteroventral dentiform process (which is seen in ventral
view as a short transverse flange). In dorsal view both petiole node and postpetiole broader than long.
Dorsum of head unsculptured except for a median carina between the frontal carinae and some extremely
faint vestiges of feeble sculpture between the frontal carinae. Alitrunk unsculptured and shining. Petiole and
postpetiole unsculptured, the latter with traces of punctulate sculpture posteriorly. First gastral tergite
densely, strongly and conspicuously shagreened on the basal half, this sculpture fading out apically on the

SIX MINOR GENERA OF MYRMICINAE 259

sclerite. Dorsum of head with numerous short fine curved hairs. Dorsal alitrunk and pedicel segments
without hairs but with scattered sparse short pubescence which is appressed and very inconspicuous. First
gasttal tergite without hairs but with a fairly dense coat of short appressed pubescence superimposed upon
the shagreened surface of the sclerite and quite conspicuous. Scapes and tibiae with pubescence but without
hairs. Colour dark reddish brown, the gaster darker in shade than the head.

Holotype worker, Liberia : Gibi, Smithsonian Firestone Exp. 1940 (W. M. Mann) (USNM, Washington).

The only known representative of Cyphoidris from West Africa, parissa is easily separated from
both its Central African congeners by its lack of strong cephalic sculpture, unsculptured alitrunk,
strongly shagreened first gastral tergite, lack of hairs on the dorsal body behind the head, and
presence of a sharp subpostpetiolar process.

Cyphoidris werneri sp. n.

HOLOTYPE WORKER. TL 3-7, HL 0-91, HW 0-83, CI 91, SL 0-62, SI 75, PW 0-63, AL 1-08.

Mandibles smooth and shining with scattered small pits, the apical margin armed with 10-11 small
denticles. Anterior margin of clypeus conspicuously impressed medially. Narrow median portion of clypeus
bicarinate above. Frontal carinae strongly developed and forming the dorsal margins of the broad but
shallow antennal scrobes, the carinae diverging posteriorly and fading out well before reaching the occipital
margin. Separation of the frontal carinae at the level of the midlengths of the eyes c. 0-37, about 0-45 x HW.
Eyes longer than wide, their maximum diameter 0-17, about 0-20 x HW. Alitrunk with promesonotum
swollen but not more or less evenly convex in profile as is usual in the genus. Instead there is a long
plateau-like dorsum which is much less strongly convex than the ascending face of the pronotum in front or
the descending face of the mesonotum behind. Propodeal dorsum in profile continuing the downward slope
of the mesonotum, the surfaces not separable. Propodeal spines with their extreme apical portions upcurved.
Metapleural lobes low and broadly triangular. Alitrunk in dorsal view broadest across the pronotal shoul-
ders, evenly narrowing posteriorly. Petiole node low and quite small in profile, its dorsal surface sloping
downwards posteriorly and the anterodorsal angle better defined than the posterodorsal which tends to
round into the posterior face. In dorsal view nodes of both petiole and postpetiole conspicuously broader
than long and the latter much broader than the former. Dorsum of head predominantly longitudinally
rugose, the rugae irregular and with cross-meshes developing behind the level of the eyes. Cross-meshes
increase in density posteriorly and a rugoreticulum is present occipitally. Sides of head below the scrobes
finely reticulate-rugose, the scrobes themselves much less strongly sculptured than the rest of the head, being
mostly smooth with a few feeble transverse rugulae. Entire dorsum of alitrunk very densely strongly and
closely reticulate-rugulose, the reticular meshes small and the rugulae raised so that in places the surface
appears reticulate-foveolate. Dorsal surfaces of rugulae with a beaded appearance due to presence of aligned
minute punctures; the entire surface blanketed with sculpture, without smooth areas. Sides of alitrunk
similarly but more loosely sculptured, the pleurae and sides of propodeum with smooth areas between the
more widely separated rugulae. Propodeal declivity smooth, with vestiges of transverse rugulae between the
spines. Nodes of petiole and postpetiole both dorsally and laterally sculptured as dorsal alitrunk. First
gastral tergite unsculptured except for fairly conspicuous pits from which hairs arise. All dorsal surfaces of
head and body densely clothed with short fine curved hairs which are subdecumbent to decumbent. Long
hairs absent except on clypeus and a row on the upper surface of each frontal carina. Colour blackish brown
with a dull reddish tint, the latter most apparent on the sides of the alitrunk. Antennae and legs dull
orange-yellow.

PARATYPE WORKERS. TL 3-5-3-8, HL 0-84-0-92, HW 0-76-0-84, CI 90-93, SL 0-60-0-66, SI 74-79,
PW 0-58-0-64, AL 0-98-1-08 (14 measured). As holotype but maximum diameter of eye 0-15-0-17, about
0-18-0-20 x HW.

Holotype worker, Rwanda : Rangiro, ix.1976, litter (P. Werner) (MHN, Geneva).

Paratypes. 14 workers with same data as holotype (MHN, Geneva; BMNH ; MCZ, Cambridge).

C. werneri is the only known species of this genus from East Africa. It is easily distinguished from
its congeners in West and Central Africa by its strong blanketing alitrunkal sculpture and by the
fact that the propodeal dorsum continues the steep slope of the posterior part of the mesonotum.
Apart from these features werneri lacks the gastral shagreening typical of parissa, is much more
densely hairy than exalta, and has shorter scapes and generally much coarser sculpture than
spinosa.

260 B. BOLTON

OC YM YRMEX Emery

(Figs 18-32)
Ocymyrmex Emery, 1886: 364. Type-species: Ocymyrmex barbiger Emery, loc. cit.; by monotypy.

DIAGNOSIS OF WORKER. Monomorphic myrmicine ants. Mandibles short and powerful, armed with five
sharp teeth which decrease in size from apex to base. The third and fourth teeth, counting from the apical,
are paired, having flanking teeth internally on the masticatory margin which are only visible when the
mandibles are open. Palp formula 4, 3 in barbiger, but 3, 3 is the predominant count (thus in ankhu, celer,
foreli, fortior, micans, monardi, nitidulus, phraxus, picardi, shushan, sobek, sphinx, velox). Ventral surface of
head with a strongly developed psammophore, the ammochaete hairs arising on the gular surface, base of
the ventral borders of the mandibles and bases of the mouthparts. Clypeus large, projecting over the basal
borders of the mandibles; posteriorly the clypeus broadly inserted between the frontal lobes. Frontal lobes
well developed but short, mostly or wholly covering the antennal insertions, ending at the same level as do
the antennal fossae; frontal carinae and antennal scrobes absent. Antennae with 12 segments, filiform,
without an apical club. Eyes well developed, situated slightly behind the midlength of the sides of the head
and usually failing to break the outline of the sides in full-face view. Mesothoracic spiracles opening high on
the sides, clearly visible in dorsal view, with a slit-like or crescent-shaped orifice. Propodeal spiracle ex-
tremely elongate, slit-shaped and very conspicuous. Propodeum unarmed, rounded in all known species.
Legs extremely long and slender, their coxae large and powerful. Petiole with a long narrow anterior
peduncle and with a rounded node which is usually low and small. Behind the node a short posterior
peduncle is present which runs to the articulation with the postpetiole. Postpetiole low and generally
shallowly curved, often voluminous but not usually forming a prominent node. Sting small, perhaps not
functional. First segment of gaster often with a narrow neck-like constriction basally (not in barbiger and
allies), the sides of the tergite usually with a series of roughly transverse, parallel indentations or grooves.

DIAGNOSIS OF FEMALES (queens). Extremely ergatoid, answering to all the characters stated above and dif-
fering from the workers only slightly, having thicker scapes, broader and more parallel frontal lobes, and
usually possessing conspicuous transverse sculpture on the head. Characters normally associated with
female ants, such as larger eyes, presence of ocelli, swollen alitrunk with flight sclerites and wings etc., are
never developed. Females are discussed in more detail below.

This easily defined and spectacular genus, which has not been revised previously, is confined to
the Ethiopian zoogeographical region where its 23 species inhabit dry to semi-desert conditions
in the eastern and southern parts of the continent. All the species nest directly into the ground,
either in the open or at the bases of plants. In the former case the nest is usually in sandy soil and
a crater is formed around the entrance hole. Arnold (1916), who was acquainted with and
reviewed the South African species, pointed out the remarkable swiftness of these ants, saying
that for speed they far outstrip 'all other ants with which I am acquainted, so much so that they
appear almost to fly over the surface of the ground'. Prins (1965) has recorded that Ocymyrmex
species are granivorous but will also attack and destroy other insects.

In the classification of both Emery (1922) and Wheeler (1922) the genus Ocymyrmex is the sole
constituent of its own tribe, the Ocymyrmecini, and Kugler's (1978) study of the sting structure
does nothing to undermine this view. In general the construction of the head in Ocymyrmex
suggests affinities with the pheidoline genera, but so many specialized characters are present that
this cannot be certain. What does seem certain is that Ocymyrmecini is best retained as a
separate tribe. It is easily isolated by the form of the alitrunk spiracles, which are unique amongst
the Myrmicinae.

Arnold (1916) and Emery (1922) both recorded that no females of Ocymyrmex had ever been
found, but they were both aware of the presence of strange variants in a number of nest-series
which had transverse sculpture on the head instead of the usual longitudinal form. Several of
these variants were described from isolated examples as separate species or subspecies, despite
the fact that long ago Arnold (1916) had recorded that they occurred in the same nests as the
more normally sculptured form.

It is now apparent that these forms are in fact the ergatoid females of the species; their
resemblance to the workers is truly remarkable. Most characters regarded as normal for female
ants are absent or have been suppressed, the body is extremely worker-like, without trace of flight
sclerites, and the head lacks ocelli or enlarged eyes. The head, however, has three specializations
which serve to distinguish the females from their workers; compare Figs 22 and 23.

SIX MINOR GENERA OF MYRMICINAE

261

Firstly, the outer margins of the frontal lobes are more widely separated in their posterior
halves in females, and the margins of the frontal lobes behind the level of the antennal insertions
are parallel or nearly so, whereas in workers they are obviously convergent behind.

Secondly, the antennal scapes are broader and frequently slightly shorter in females than in
workers.

To illustrate these two points all available females were measured for width across the margins
of the frontal lobes at their posteriormost point (FW), and the maximum width of the shaft of the
scape (SW) discounting the apical swelling when present. The same measurements were taken for
an equal number of workers chosen at random from the series in which the respective females
originated. These measurements were compared with the standard measures of HW and SL, as
follows (where n = number of females measured).

species

HW

FW

FW/HW

SL

SW

SW/SL

fortior

female

1-68-1-

80

0-48-0-54

0-28-0-31

1

48-1-66

0-18-0-19

0-11-0-12

(n

-5)

„

worker

1-72-1-

96

0-40-0-46

0-23-0-26

1

46-1-70

0-12-0-16

0-08-0-Q9

nitidulus

female

1-67-1-

76

0-48-0-52

0-28-0-30

1

•40-1-52

0-15-0-16

0-10-0-11

(n

= 8)

worker

1-72-1-

78

0-40-0-44

0-22-0-25

1

•48-1-58

0-12-0-14

0-08-0-10

picardi

female

2-48-2-

50

0-70

0-28

2

•28-2-36

0-22-0-24

0-09-0-10

(n

= 2)

„

worker

2-44-2-

60

0-58-0-62

0-24

2

40-2-54

0-18-0-20

0-08

velox

female

2-08-2-

18

0-50-0-56

0-24-0-26

2'

08-2-16

0-20-0-22

0-10

(n

= 3)

M

worker

2-14-2-

18

0-44-0-46

0-20-0-21

0-20-0-26

0-18-0-20

0-08-0-09

barbiger

female

1-48-1-

60

0-44-0-46

0-29-0-30

1

34-1-48

0-16

0-11-0-12

(n

= 2)

n

worker

1-52-1-

60

0-40-0-44

0-26-0-27

1

40-1-52

0-12-0-14

0-09

flaviventris

female

1-50-1-

64

0-44-0-46

0-28-0-31

1

40-1-48

0-15-0-17

0-11

(n

= 3)

55

worker

1-54-1-

76

0-36-0-40

0-23-0-24

1

46-1-66

0-12-0-14

0-08-0-09

weitzeckeri

female

1-64

0-49

0-30

1-42

0-17

0-12

(n

= 1)

,,

worker

1-60

0-38

0-24

1-42

0-14

0-10

foreli

female

1-64-1-

76

0-50-0-53

0-30

1

52-1-65

0-16-0-18

0-11

(n

= 2)

n

worker

1-68-1-

76

0-40

0-23-0-24

1

56-1-76

0-14

0-08-0-09

sobek

female

1-80-1-

84

0-50-0-54

0-28-0-29

1

62-1-68

0-17-0-18

0-10-0-11

(n

= 4)

«

worker

1-76-1-

84

0-42-0-44

0-23-0-24

1

66-1-70

0-14-0-15

0-08-0-09

Finally, the dorsum of the head behind the level of the eyes usually has strong regular trans-
verse sculpture in females, whereas such sculpture is generally longitudinal in workers. Excep-
tions to this occur in robecchii where both known specimens have transverse sculpture, implying
that they are female, but lack the specialized characters of scapes and frontal lobes noted above.
They are treated as workers in this paper although it is realized that they may turn out to be
females. In hirsutus^most workers have arched-transverse sculpture on the head, but in this case it
is coarse, sharply developed, irregular and vermiculate. In some species (velox) the cephalic
sculpture is reduced in both castes and may not be apparent.

A few individual workers in any species may show some transverse costulae or rugulae close to
the occipital margin, especially in species where the main longitudinal cephalic sculpture arches
outwards towards the occipital corners, but none have the extensive transverse sculpture devel-
oped by their respective females and their frontal lobes and antennal scapes are of the worker
form.

As the females are apterous the founding of new nests must be by colony fission or by single
newly mated females setting out on their own. Quite probably a relatively large number of
females are retained in the nest at all times as captures are frequent. Whether all females in a
colony lay eggs or whether this function is dominated by a single laying female who suppresses
the rest is not known, but I suspect the latter as stray females appear to be fairly common in
worker samples collected outside the nest and which are, apparently, behaving like workers.

Males, not dealt with here, are very poorly represented in collections, being known only for the
species/orf/or, barbiger and weitzeckeri.

262 B. BOLTON

Synonymic list of species

ankhu sp. n.
barbiger Emery

barbatus Emery

barbiger var. robustior Stitz syn. n.

barbiger var.flavescens Stitz syn. n.
cavatodorsatus Prins
celer Weber
cursor sp. n.

flaviventris Santschi stat. n.
foreli Arnold stat. n.
fortlor Santschi stat. n.

weitzeckeri st. transversus Santschi syn. n.

arnoldi Forel syn. n.
weitzekeri [sic] st. abdominalis Santschi

weitzaeckeri [sic] var. usakosensis Stitz syn. n.
hirsutus Forel
laticeps Forel
micans Forel stat. n.
monurdi Santschi stat. n.
nitidulus Emery stat. n.
phraxus sp. n.
picardi Forel

carpenteri Donisthorpe syn. n.
robecchii Emery
shushan sp. n.
sobek sp. n.
sphinx sp. n.
turneri Donisthorpe
velox Santschi
weitzeckeri Emery

weitzeckeri subsp. wroughtoni Forel syn. n.
zekhem sp. n.

Key to species (workers)

1 Anterior clypeal margin without a conspicuous semicircular median impression, the margin

entire or at most flattened or feebly eroded in the middle 2

Anterior clypeal margin with a conspicuous semicircular median impression, the impression
usually flanked by a pair of denticles or teeth 8

2 With head in full-face view the large eyes (0-28 x HW) very obviously breaking the outline of

the sides (Fig. 25). Middle of anterior clypeal margin with a low, broad bluntly triangular

prominence. (South West Africa) turneri (p. 279)

With the head in full-face view the smaller eyes( < 0-25 x HW) failing to break the outline of the
sides. Middle of anterior clypeal margin without a triangular prominence .... 3

3 First gastral tergite in dorsal view strongly constricted basally and forming a narrow neck, the

sclerite in this region roughly parallel-sided and no broader than the postpetiole. Petiole node
low and broadly rounded in profile, with a blunt angular ventral process about half-way

along the peduncle (Fig. 29) 4

First gastral tergite in dorsal view not constricted basally, without a narrow neck, the sclerite
broadening evenly from its articulation with the postpetiole. Petiole node high and domed in
profile, without trace of a ventral process at the midlength of the peduncle (Fig. 32) . . 5

4 Larger species, HW 2-04, PW 1-30; head narrower, CI 98. Metapleural lobes large and very

strongly prominent, plainly visible with alitrunk in absolute profile, not concealed by the

bulge of the metapleural glands. (Angola) cur sor (p. 267)

Smaller species, HW 1-70-1-74, PW 1-08-1-14; head broader, CI 102-103. Metapleural lobes
very small, not prominent, scarcely or not visible with alitrunk in absolute profile, mostly or
entirely concealed by the bulge of the metapleural glands (Fig. 29). (Angola) . . laticeps (p. 27 1 )

5 Larger species, HL > 2-0. Promesonotal outline in profile low and evenly rounded, propodeal

SIX MINOR GENERA OF MYRMICINAE 263

dorsum more or less flat; dorsal outline of entire alitrunk not saddle-shaped (Fig. 32) . . 6
Smaller species, HL < 1-50. Promesonotal outline in profile high and dome-like, propodeal
dorsum sloping sharply upwards posteriorly; dorsal outline of entire alitrunk strongly saddle-
shaped. (South Africa) cavatodorsatus (p. 266)

6 Basal half of first gastral tergite with conspicuous hairs which are as long as those on the

mesonotal and propodeal dorsa. Smaller species with longer scapes, HW 1-90, SI 116. Entire
ant black to the naked eye, the gaster the same colour as the alitrunk. (South West Africa)

zekhem (p. 281)

Basal half of first gastral tergite without hairs or at most with 1-2 minute inconspicuous hairs
which are much shorter than those on the mesonotal and propodeal dorsa. Larger species
with shorter scapes, HW > 2-0, SI range 94-105. Entire ant not black to the naked eye . 7

7 Tricoloured species with alitrunk dull red to blackish red, gaster orange to yellow and head an

intermediate shade. Hairs on dorsal alitrunk usually white, only rarely coloured. SI 101-105,

CI 95-97 (Angola, South West Africa) . . . velox (p. 280)

Colour uniform rich orange-brown, or sometimes with the gaster slightly lighter. Hairs on
dorsal alitrunk reddish brown. SI 94-102, CI 97- 101. (Angola) . . " . . ankhu (p. 265)

8 Large or very large species, HW > 2-00, SL> 1-90 9

Smaller species, HW < 2-00, SL < 1-90 11

9 Hairs on dorsal alitrunk dark reddish brown to blackish. Extremely large species, HW 2-30 or

more. (Zimbabwe, Botswana, South West Africa, Angola) picardi (p. 275)

Hairs on dorsal alitrunk white to silvery. Large species but not approaching the above in size,
H Win range 2-02-2- 15 10

10 Antennal scapes relatively longer, SI > 100 (range 103-106). Dorsum of head behind level of

eyes with dense transverse costulate sculpture, ground-sculpture between the costulae vesti-
gial. (Somali Republic) robecchii (p. 276)

Antennal scapes relatively shorter, SI < 100 (range 93-98). Dorsum of head behind level of eyes
with dense longitudinal rugular sculpture, ground-sculpture between the rugulae a coarse
conspicuous punctulation or granulation. (Botswana) sphinx (p. 278)

1 1 First gastral tergite not constricted basally, not forming a roughly parallel-sided neck behind the

postpetiole, the gaster broadening more or less evenly from immediately behind the postpetio-
le. Palp formula 4, 3. (South Africa, South West Africa) barbiger (p. 265)

First gastral tergite constricted basally, forming a roughly parallel-sided neck behind the postpe-
tiole which is no wider than the postpetiole. Palp formula 3, 3 12

12 Spaces between costulae on dorsum of head smooth and shining, either without or with only

vestigial ground-sculpture; the surface smooth and polished, often with a slick and very shiny
appearance. Commonly the costulae themselves are low, rounded and glossy, not sharply
defined ; sometimes the costulae effaced in places leaving unsculptured areas . . . . 13
Spaces between costulae or rugulae on dorsum of head with punctulate or granular ground-
sculpture which is conspicuous, the surface not appearing smooth and polished, without a
slick and very shiny appearance. Costulae or rugulae on dorsum of head always strongly and
often sharply developed, very distinct ........... 14

1 3 Pronotum in profile with dorsal outline more or less flat (Fig. 19) or even slightly concave. Head

longer and narrower, CI 92-93. (Sudan) celer (p. 267)

Pronotum in profile with dorsal outline evenly convex (Fig. 18). Head shorter and broader,
CI 96-100. (Ethiopia, Somali Republic, Kenya, Uganda, Tanzania) . nitidulus (p. 274)

14 Petiole node much enlarged, swollen in profile and very conspicuous (Fig. 20). Sculpture of

petiole node coarse, strongly developed everywhere, with strong coarse rugae running right
round the node ........... 15

Petiole node not greatly enlarged (Figs 26-28, 30, 31). Sculpture of petiole node not strongly
developed everywhere, usually feebly and unevenly sculptured to unsculptured; commonly
with fine or weak rugulae but these only rarely encircling the whole node. A few sharp
transverse rugulae may be present dorsally but in this case the node is not much enlarged in
profile ............... 16

1 5 Alitrunk very dark dull reddish brown or reddish black, usually appearing as black to the naked

eye. Gaster yellow or yellowish red, much lighter than and contrasting strongly with the dark
alitrunk. Head a shade of dull red intermediate between that of alitrunk and gaster. (Zim-
babwe, Botswana) sobek (p. 277)

Colour uniform orange-red to red throughout, the gaster usually the same shade as the head and

alitrunk but sometimes slightly lighter or darker. (Zimbabwe) .... foreli (p. 269)

264 B. BOLTON

16 Dorsum of head behind level of posterior margins of eyes mostly or wholly sculptured with

extremely tightly packed fine rugulae which are sharply and narrowly vermiculate and which
are mostly or entirely transverse in direction. (South West Africa) . . . . hirsutus (p. 271)
Dorsum of head behind level of posterior margins of eyes not sculptured as above, usually with
longitudinal costulae or rugulae which may be obscured in some places by dense ground-
sculpture. Transverse sculpture usually absent but if a few transverse components are present
then they are not sharply and narrowly vermiculate . . . . . . . . 17

17 Alitrunk dull brick-red to black 18

Alitrunk yellowish orange to bright red ........... 21

18 Dorsum of head from inner margin of eye to antennal fossa and the area extending back from

this to the occiput blanketed by strong irregular granular sculpture which masks or replaces
the costulae or rugulae usually seen in this area. (South West Africa, Angola) . . monardi (p. 273)
Dorsum of head from inner margin of eye to antennal fossa and the area extending back from

this to the occiput costulate or rugulose, without blanketing granular sculpture . . . 19

19 Petiole node in dorsal view very broad, distinctly much broader than long (Fig. 30), the maxi-

mum width of the node greater than the length of the petiole from its spiracle to the apex of
the collar where it articulates with the postpetiole. (Lesotho, South Africa) weitzeckeri (p. 280)

Petiole node in dorsal view more slender, at most as broad as long (Figs 26, 27), the maximum
width of the node usually distinctly less than the length of the petiole from its spiracle to the
apex of the collar where it articulates with the postpetiole ........ 20

20 Petiole in profile shaped as in Fig. 27, the dorsal surface of the peduncle and the anterior face of

the node confluent or nearly so, without a marked change of slope where the two surfaces

meet. (Tanzania) phraxus (p. 274)

Petiole in profile shaped as in Fig. 26, the dorsal surface of the peduncle and the anterior face of
the node not at all confluent, with a marked change of slope where the two surfaces meet.
(Zambia, Zimbabwe, Botswana, South Africa, South West Africa, Angola) • fortior (part; p. 269)

21 With the alitrunk in profile the promesonotum forming a high dome-like convexity (Fig. 31).

(South West Africa) . .... shushan (p. 277)

With the alitrunk in profile the promesonotum evenly shallowly convex 22

22 Sculpture of dorsum of head between eyes and from this level to occiput everywhere longitudi-

nally costulate, not vermiculate nor dominated by coarse punctulate ground-sculpture, the
costulae quite regular and evenly spaced. (Zambia, Zimbabwe, Botswana, South Africa, South

West Africa, Angola) fortior (part; p. 269)

Sculpture of dorsum of head between eyes and from this level to occiput densely finely longitudi-
nally rugulose, the rugulae conspicuously irregular, being wavy to vermiculate and with a
strong granular or punctulate ground-sculpture which may become the dominant component
in places 23

23 Peduncle of petiole ventrally with an elongate convex keel-like process (Fig. 28). Slightly larger

species, HW 1-54-1-70, SL 1-44-1-54. (Botswana, South West Africa) . . flaviventris (p. 268)
Peduncle of petiole ventrally without a keel-like process. Slightly smaller species, HW 1-40-1-56,
SL 1-38-1 -44. (South West Africa) micans (p. 272)

Treatment by species

The members of the genus Ocymyrmex are very uniform in structure and are not easily divisible
into meaningful species-groups. For this reason they are set out below in alphabetical order.

To some extent the genus can be split into three unequal complexes of species based on the
degree of constriction of the first gastral tergite. 0. barbiger, cavatodorsatus and turner! have the
base of the gaster unconstricted, quite broad basally and continuing to broaden behind. In velox,
picardi, ankhu and zekhem the gaster is narrow basally and gradually broadens behind without
forming a narrow neck. In all other species a conspicuous narrow neck, often parallel-sided, is
developed. 0. sphinx forms an intermediate stage between these last two complexes.

The presence or absence of a median clypeal impression seems at first sight a means of dividing
the genus into groups but, although a useful key character, it does not serve to aggregate related
species. Examination of species without the clypeal impression shows that in most cases they are
more closely related to forms possessing it than to the others which lack it.

SIX MINOR GENERA OF MYRMICINAE 265

Ocymyrmex ankhu sp. n.

HOLOTYPE WORKER. TL 1O2, HL 2 34, HW 2-35, CI 100, SL 2-30, SI 98, PW 1-50.AL 2-96.

Anterior clypeal margin entire, without a strong median notch flanked by a pair of teeth but only with a
tiny erosion of the apron where the weak median clypeal carina runs into it. Maximum diameter of eye 0-42,
about 0-18 x HW, the eyes conspicuously failing to break the outline of the sides of the head in full-face
view. Sides of head extremely weakly divergent anteriorly, rounding broadly and evenly into the occipital
margin which is shallowly convex on each side of a median indentation. Alitrunk in profile with the
promesonotum low and very shallowly convex, almost flat dorsally but with the posterior half of the
mesonotum sloping more steeply downwards. Propodeal dorsum almost flat, with an exceedingly shallow
depression in the surface just in front of the level of the spiracle. Posteriorly the propodeal dorsum rounding
broadly and evenly into the shallowly convex declivity. Metapleural lobes low and truncated posteriorly.
Petiole in profile large, broadly dome-like and rounded, the anterior peduncle without trace of a ventral
process. In dorsal view the petiole node very slightly longer than broad, rounded and with evenly convex
sides. Postpetiole in dorsal view broader than long. Base of first gastral tergite no broader than the
postpetiole but not constricted to a narrow neck; instead the sides diverge quickly and evenly from the base.
Dorsum of head between eyes with faint superficial vestiges of fine and quite dense rugular or costulate
sculpture which in places is almost effaced. Ground sculpture absent except for the faintest remnants of a
minute superficial reticulation, the surface mostly smooth and glossy and the scattered hair pits quite clearly
visible. Occipital region of head mostly smooth but a narrow strip in front of this with feeble transverse
sculpture. Pronotal dorsum mostly smooth, with marginal remnants of fine arched rugulae and vestiges of
the longitudinally sculptured area between the mesothoracic spiracles just visible. Remainder of dorsal
alitrunk very finely and feebly transversely rugulose, with a tendency for the rugulae to fade out centrally.
Sides of alitrunk more strongly and more sharply rugulose or costulate, weaker on the pronotal sides than
elsewhere. Petiole, postpetiole and gaster unsculptured except for a fine superficial reticulation. Dorsal
surfaces of head, alitrunk, petiole and postpetiole with numerous strong reddish brown hairs. Basal half of
first gastral tergite hairless but more apically the segment with 1-2 very short, inconspicuous hairs. Colour a
uniform rich orange-brown, the gaster very slightly lighter in shade than the head and alitrunk.

PARATYPE WORKERS. TL 9-7-10-2, HL 2-26-2-34, HW 2-22-2-34, CI 97-101, SL 2-22-2-34, SI 95-102,
PW 1-40-1-48, AL 2-84-2-92 (7 measured). Maximum diameter of eye 0-42-0-44, about 0-18-0-19 x HW.
As holotype.

Holotype worker, Angola: 5 miles [8 km] E. of Vila Arriaga, 1000 m, 21.V.1958 (E. S. Ross & R. E. Leech)
(CAS, San Francisco).

Paratypes. 7 workers and 1 female with same data as holotype (CAS, San Francisco; BMNH; MCZ,
Cambridge).

0. ankhu is closest related to velox and shares most of its diagnostic characters. However, ankhu is
more or less uniformly coloured and has the body pilosity reddish brown. In velox the body is
conspicuously tricoloured and the hairs are usually white.

Ocymyrmex barbiger Emery

Ocymyrmex barbiger Emery, 1886: 364, pi. 17 figs 9-11. Syntype workers and male, SOUTH AFRICA: Cape of

Good Hope (L. Peringuey) (MHN, Geneva; MCSN, Genoa) [examined].
Ocymyrmex barbatus Emery, 1892: 1 14, 1 17. [Lapsus for barbiger Emery.]
Ocymyrmex barbiger var. robustior Stitz, 1923: 146. Syntype worker, SOUTH WEST AFRICA: Luderitzbucht,

5-1 3.vii. 1911; and Swakopmund, 12-19.iv.1911 (W. Michaelsen)(MNWJ, Berlin) [examined] Syn.n.
Ocymyrmex barbiger \ar.flavescens Stitz, 1923: 147. Syntype worker, SOUTH WEST AFRICA: Okaputa,

5.v.l9ll(W. Miduw?/sen)(MNHU, Berlin) [examined]. Syn.n.

WORKER. TL 6-7-7-2, HL 1-54-1-84, HW 1-42-1-76, CI 90-97, SL 1-38-1-58, SI 87-98, PW 0-94-1-10,
AL 2-00-2-20 (20 measured).

Anterior clypeal margin with a semicircular median impression which is flanked by a pair of teeth. The
impression is usually deep and conspicuous but in some individuals may be broad and quite shallow. The
flanking teeth are generally well developed but commonly are broadly triangular and blunted, sometimes
little more than broadly rounded prominences. Maximum diameter of eye 0-32-0-35, about OT 9-
0-22 x HW, in full face view not breaking the outline of the sides of the head. Promesonotum forming an
evenly rounded low convexity which slopes downwards behind to the propodeum, the dorsum of which
varies from more or less flat to slightly inclined. Propodeal dorsum rounding broadly and evenly into the

266 B. BOLTON

declivity. Metapleural lobes low but prominent, their free edges rounded to bluntly truncated posteriorly.
Petiole node in profile usually quite high, bluntly rounded-subconical in shape, but sometimes the dorsum
more flattened and the node appearing less regular in shape. Petiole node in dorsal view usually broader
than long, less commonly only about as broad as long. Postpetiole always broader than long in dorsal view,
discounting the anterior articulating portion. Base of first gastral tergite without a neck-like constriction.
Head finely and densely rugulose, with fine punctulate ground-sculpture; the pattern formed by the rugulae
very variable on the dorsum but apparently following a step by step change. In many the cephalic rugulae
are regular and longitudinal, parallel on the central strip and running straight back to the occipital margin,
but the more lateral rugulae tending to diverge and arch outwards behind the eyes. This seems the basic
pattern from which the following derive sequentially. Firstly, the median-line rugulae begin to diverge
posteriorly, forcing the more lateral rugulae to arch outwards even more. Next, the point at which the
median-line rugulae begin to diverge shifts gradually forward, arching the lateral components more strongly
outwards all the time. When the point of divergence of the mid-line rugulae has shifted a certain distance
forwards a number of V-shaped rugulae appear in front of the impression in the centre of the occipital
margin, which occupy the space vacated by the now divergent mid-line rugulae. Then, as the point of
divergence of the mid-line rugulae shifts still further forward the V-shaped rugulae also shift forward on the
head and their angle becomes more obtuse the further forward they shift. This process continues until
ultimately the head is mostly transversely sculptured behind the eyes, although the rugulae tend to retain
their broadly V-shaped nature more posteriorly on the head. Pronotal dorsum usually with extensive
longitudinal sculpture, the more lateral components arching across in front of the central longitudinals.
Space between mesothoracic spiracles usually longitudinally rugose, sometimes obliquely so and very rarely
the sculpture here more or less transverse. Remainder of dorsal alitrunk transversely rugose. Petiole with
transverse rugulae ventrally which usually extend for some distance up the sides of the node, commonly to
the dorsum. In more strongly sculptured samples the rugulae continue across the top of the node but often
the dorsum is more weakly sculptured. Postpetiole finely shagreened or with fine superficial patterning only.
All dorsal surfaces of head and body with hairs, those on the first gastral tergite more numerous and longer
in barbiger than is usual in the genus, the gastral hairs frequently approaching the length and density seen on
the alitrunk. Elsewhere in the genus gastral hairs on the first tergite are much shorter and much sparser than
on the alitrunk. Colour of head and alitrunk orange to dull brick red, the gaster darker, frequently dull
brown with a reddish tint or even blackish brown.

One of three known species in which the base of the first gastral tergite is not constricted to a
narrow neck, barbiger is separable from both others showing this character (cavatodorsatus,
turneri) by its possession of an impressed anterior clypeal margin and its strong sculpture.

MATERIAL EXAMINED

South West Africa: Maltahoe dist., Sesriem Farm (M. C. Day). South Africa: Cape Prov., Orange Riv.,
Kakamas (G. Arnold); C.P., Orange Riv., no loc. (G. Arnold); C.P., Betty's Bay (G. Arnold); C.P., Willow-
more (G. Arnold); Willowmore (ex coll. Mayr); Willowmore (H. Brauns); C.P., Table Mt (G. Arnold); C.P.,
Victoria West (G. Arnold); C.P., Mossel Bay (R. E. Turner); C.P., Camps Bay (R. E. Turner); C.P., Die
Panne (M. C. Day); C.P., Pt Elizabeth (W. L. Brown); C.P., Grahamstown (L. Weatherill & W. L. Brown).

Ocymyrmex cavatodorsatus Prins

Ocymyrmex cavatodorsatus Prins, 1965: 1021 figs 1, 2. Syntype workers, SOUTH AFRICA: Cape Prov., Dist.
Upington, Louisvale, 6.viii.l964, AcAx 3412 (A. J. Prins) (Res. Inst. for Plant Protection, Pretoria, South
Africa).

I have not seen the types of this species but it is obvious from Prins' original description that
cavatodorsatus is a very distinctive species. Its main diagnostic characters are as follows.

WORKER. TL 4-5, HL 1-02, AL 1-41. Anterior clypeal margin without a semicircular median impression.
Eyes not breaking outline of sides of head in full-face view. Alitrunk in profile characteristically shaped, with
the promesonotum forming a single high, strongly arched convexity, the posterior mesonotum and anterior
propodeum forming a uniform deep concavity and the remainder of the propodeum arching upwards again
posteriorly before rounding narrowly into the deep and almost vertical declivity; the entire alitrunk having
a strong saddle-shaped appearance in profile. Petiole node high-subconical in profile, longer than broad in
dorsal view. Postpetiole broader than long in dorsal view. Base of first gastral tergite in dorsal view without
a neck-like constriction, the sides of the tergite evenly convex behind the point of articulation with the
postpetiole. Dorsum of head from level of eyes to occiput smooth and shining. Promesonotal arch dorsally

SIX MINOR GENERA OF MYRMICINAE 267

smooth and shining, including the area between the mesothoracic spiracles. Remainder of dorsal alitrunk
with faint transverse rugae which are almost effaced; the propodeal declivity smooth. Head bright red to
yellowish red, the alitrunk yellowish red to brick red, the gaster distinctly darker, piceous.

As Prins points out, this species is related to barbiger in its lack of a basal gastral constriction. It
is separated from barbiger by its lack of a clypeal impression, much reduced sculpture and
strongly saddle-shaped alitrunk. In these aspects it seems closely related to turneri, another
species without a gastral constriction, which also lacks a clypeal impression and has reduced
sculpture. However, turneri has the alitrunk jet black, the clypeal margin prominent medially, the
eyes large and breaking the outline of the sides of the head, the scapes with SI > 100 (apparently
SI < 100 in cavatodorsatus to judge by Prins' fig. 1.) and has a transverse arched crest on the
alitrunk at the promesonotal junction.

Ocymyrmex celer Weber stat. n.
(Fig. 19)

Ocymyrmex weitzeckeri subsp. celer Weber, 1943: 368. Syntype workers, SUDAN: Torit, N. of Imatong Mts,
6.viii.l939, no. 1462 (N. A. Weber) (MCZ, Cambridge) [examined].

WORKER: TL 8-1-8-5, HL 1-98-2-04, HW 1-82-1-88, CI 91-94, SL 1-68-1-73, SI 89-95, PW 1-12-1-16,
AL 2-30-2-46 (4 measured).

Anterior clypeal margin with a conspicuous median impression which is flanked on each side by a low
broad tooth. Sides of head in front of eyes more or less parallel in full-face view, not obviously diverging
anteriorly as is frequent in the genus. Maximum diameter of eye 0-38-0-39, about 0-21 x HW. Dorsum of
pronotum in profile flat to indented, sloping upwards behind to its junction with the mesonotum; the
promesonotum not forming a single even convexity. Dorsum of propodeum rounding broadly and evenly
into the sloping declivity. Metapleural lobes deep but narrow, rounded and little projecting. Petiole node in
profile small, low and rounded, in dorsal view longer than broad. Postpetiole in dorsal view slightly longer
than broad. First gastral tergite with a neck-like constriction basally. Dorsum of head finely densely and
regularly longitudinally costulate, the costulae low and superficial, weakly developed, parallel and arching
outwards behind the eyes. Spaces between the costulae highly polished, without strong granular or punctu-
late ground-sculpture; the whole head with a slick and glossy appearance. Pronotum with weak to feeble
arched-transverse costulate sculpture dorsally, which may be almost effaced centrally, the space between the
mesothoracic spiracles with longitudinal or transverse costulate sculpture. Remainder of dorsal alitrunk
transversely and more strongly costulate. Sides of alitrunk regularly densely sharply costulate everywhere.
Petiole node with transverse rugulae ventrally which may extend for some distance up the sides, and with
faint rugulae on the dorsal peduncle, but the dorsum of the node unsculptured or only with the faintest
vestiges present. Postpetiole only with superficial patterning. All dorsal surfaces of head and alitrunk with
numerous hairs of varying length; first gastral tergite with hairs shorter and much sparser than on alitrunk.
Colour glossy dull red, the gaster dark brown.

Along with nitidulus this species can be separated from all others in which the clypeus is im-
pressed and the first gastral segment constricted by the slick and glossy appearance of the head.
In all others the ground-sculpture on the head consists of quite conspicuous granulation or
punctulation, but in celer and nitidulus this is very reduced or absent, leaving the spaces between
costulae smooth or very nearly so, and shining. The two may be distinguished by the shape of the
pronotum, which in nitidulus is convex and, together with the mesonotum, forms an evenly
convex surface. In celer the pronotum is flat and an even convexity does not result, compare Figs
18 and 19.

MATERIAL EXAMINED
Sudan: Torit (N. A. Weber).

Ocymyrmex cursor sp. n.

HOLOTYPE WORKER. TL 9'7, HL 2-08, HW 2-04, CI 98, SL 1 94, SI 95, PW 1-30, AL 2-74.

Anterior clypeal margin with a minute and very shallow inconspicuous indentation in the apron medially,
without a conspicuous semicircular impression flanked by a pair of teeth. With the head in full-face view the

268 B. BOLTON

occipital margin very shallowly but quite broadly concave. Maximum diameter of eye 0-44, about
0-22 x HW. Promesonotum in profile evenly rounded, sloping posteriorly. Anterior half of propodeal
dorsum very feebly concave but above the spiracle very shallowly convex before rounding broadly and
evenly into the declivity. Metapleural lobes prominent and narrowly rounded, easily visible in profile, not
concealed by the bulge of the metapleural glands. Peduncle of petiole with a broadly triangular low blunt
process on its ventral surface, about half-way between the insertion and the level of the spiracle. Node of
petiole in profile low and broadly rounded, with a differentiated dorsal surface which is almost flat. In dorsal
view the petiole node as long as broad, the postpetiole broader than long. Base of first gastral tergite
constricted, forming a neck in dorsal view. Dorsum of head costulate, predominantly longitudinally so but
with a few arching in over the eye. Ground-sculpture a fine superficial granulation or punctulation which is
more conspicuous away from the midline of the head. Centre of pronotal dorsum almost smooth, with only
vestigial sculpture; in front of this arched-transverse costulae are present and behind it longitudinal costulae
run back between the mesothoracic spiracles. Remainder of dorsal alitrunk and propodeal declivity trans-
versely rugose. Sides of alitrunk densely and evenly costulate-rugose, the sculpture regular except around
the propodeal spiracle. Petiole ventrally transversely rugose from the level of the process and the peduncle
dorsally with a few weak transverse rugae. The node itself more weakly sculptured, mostly with superficial
patterning only but the posterior face with a few weak transverse rugulae. Postpetiole only with superficial
patterning. Dorsal surfaces of head and alitrunk with numerous hairs of varying length but first gastral
tergite only with scattered short hairs. Alitrunk glossy dull red, the head lighter and with an orange tint, the
gaster darker, reddish brown.

Holotype worker, Angola: Kopeio, vii.1931 (T. D. A. Cocfcm?//)(BMNH).

The closest relative of cursor is laticeps, also from Angola. Differentiation of the two species is
discussed under laticeps.

Ocymyrmexflaviventris Santschi stat. n.
(Fig. 28)

Ocymyrmex hirsutus \ar.flaviventris Santschi, 1913: 431. Holotype worker, SOUTH WEST AFRICA: Windhoek
(Viehmeyer)(NM, Basle) [examined].

WORKER. TL 7-1-7-4, HL 1-64-1-80, HW 1-54-1-70, CI 93-97, SL 1-44-1-58, SI 90-96, PW 0-98-1-04,
AL 2-04-2- 28 (17 measured).

Anterior clypeal margin with a narrow but deep semicircular impression medially, the impression flanked
by a pair of teeth. Occipital corners broadly rounded, the margin medially with a small indentation. Eyes
with maximum diameter 0-38, about 0-22 x HW. Promesonotum in profile evenly shallowly convex, the
propodeal dorsum posteriorly rounding narrowly into the declivity which is almost vertical. Metapleural
glands swollen and projecting strongly to the rear, in profile concealing all but the extreme tips of the
metapleural lobes; the projection of the metapleural glands enhanced by the near-vertical propodeal decliv-
ity. Peduncle of petiole ventrally with an elongate keel-like process which is semitranslucent and unsculp-
tured, evenly shallow convex throughout its length. Petiole node small in profile, evenly rounded. In dorsal
view the petiole node broader than long, the maximum width of the node about equal to the distance from
the spiracle to the apex of the collar where the petiole articulates with the postpetiole. Postpetiole in dorsal
view slightly longer than broad. Base of first gastral tergite constricted and forming a neck. Dorsum of head
longitudinally very densely finely rugulose, the rugulae close-packed and irregular, being narrowly wavy or
even minutely vermiculate in places. Ground-sculpture a conspicuous granulation or punctulation. Rugulae
between and on median strip just behind the frontal lobes more regular than elsewhere. Dorsal alitrunk
transversely densely rugose, the sculpture longitudinal only between the mesothoracic spiracles and on the
arched portion of the pronotum. Sides of alitrunk rugose everywhere. Petiole with a few transverse rugae
beneath the node and on the dorsum of the peduncle. Elsewhere on the petiole sculpture is vestigial to
absent. Postpetiole unsculptured except for faint superficial patterning. All dorsal surfaces of head and
alitrunk with numerous hairs of varying length. Propodeal dorsum with long hairs arising from a fairly
dense mat of much shorter hairs. First gastral tergite with sparse scattered hairs which are much shorter
than those on the alitrunk. Colour bright orange-yellow, the gaster lighter and more yellow than the head
and alitrunk.

0. flaviventris is characterized by its light orange-yellow colour, keel-like process below the
petiole peduncle, broad node, prominent metapleural glands and uneven cephalic sculpture. It is

SIX MINOR GENERA OF MYRMICINAE 269

closest related to shushan and hirsutus, but in the former the promesonotum forms a conspicuous
high dome and the latter lacks a keel-like subpeduncular process as well as having the cephalic
sculpture transverse behind the level of the eyes.

MATERIAL EXAMINED
Botswana: Damara Pan (G. V. Son); nr Nkata (G. U. Son).

Ocymyrmex foreli Arnold stat. n.

Ocymyrmex weitzaeckeri [sic] var. foreli Arnold, 1916: 197. Syntype workers, ZIMBABWE: Redbank, 7.iv.l912
(G. 4rnoW)(BMNH;NM, Bulaway) [examined].

WORKER. TL 7-3-8-0, HL 1-76-1-86, HW 1-64-1-76, CI 92-95, SL 1-56-1-72, SI 92-98, PW 1-08-1-14,
AL 2-28-2-44 (14 measured).

Anterior clypeal margin with a semicircular impression but this impression frequently shallower and
broader than is usual in the genus; flanked by a pair of low broad tubercles or blunt small teeth formed by a
thickening of the clypeal apron. Maximum diameter of eye 0-34-0-36, about 0-20-0-22 x HW. Promesono-
tum in profile evenly shallowly rounded and convex. Propodeal dorsum more or less flat to slightly convex,
rounding broadly and evenly into the declivity. Metapleural lobes small and rounded. Petiole node in profile
large, almost or quite as massively developed as in sobek, Fig. 20. In dorsal view the petiole node appearing
swollen, as broad as or broader than long; postpetiole dorsally as broad as long, discounting the anterior
articulatory section. Base of first gastral tergite constricted and forming a neck behind the postpetiole.
Dorsum of head finely and densely longitudinally regularly costulate-rugulose, the components sharply
defined and parallel. On the central part of the dorsum the sculpture is longitudinal, running straight back
to the occiput or at most diverging slightly on each side of the occipital impression. More laterally on the
dorsum the rugulae are divergent and arch outwards behind the eye. Ground-sculpture of head finely
punctulate or granular. Pronotal dorsum usually with arched-transverse costulate sculpture followed by a
patch of longitudinal sculpture which runs back between the mesothoracic spiracles. However, in some
workers the sculpture here is oblique and in a few is more or less transverse. Remainder of dorsal alitrunk
and also propodeal declivity transversely costulate or rugose. Petiole node coarsely sculptured everywhere,
with strong, sharply defined rugae which encircle the node, running continuously across the dorsal and
ventral surfaces and down the sides. Peduncle of petiole also with transverse rugulae both dorsally and
ventrally, but these are weaker or effaced on the sides. All dorsal surfaces of head and alitrunk with
numerous hairs; first gastral tergite also with hairs but these are shorter and much sparser than those on the
alitrunk. Colour usually uniform orange-red to red throughout, but sometimes the gaster slightly lighter or
darker than the alitrunk.

This species, known at present only from Zimbabwe, is closest related to sobek with which it
shares the character of possessing a massively developed petiole node which is coarsely sculp-
tured. The two are separable on colour pattern as in sobek the alitrunk is dark reddish brown to
almost black, the gaster yellow and contrasting strongly with the alitrunk. The head is dull red,
intermediate in colour between alitrunk and gaster. Beside this, the sculpture on the petiole is
more sharply defined and regular in foreli than in sobek.

MATERIAL EXAMINED

Zimbabwe: Bembesi Riv. (G. Arnold).

Ocymyrmex fortior Santschi stat. n.
(Fig. 26)

Ocymyrmex weitzeckeri st. fortior Santschi, 1911: 209. Syntype workers, ANGOLA: Benguela, Cucala (J. Cru-

chet) (NM, Basle) [examined].
Ocymyrmex weitzeckeri st. transversus Santschi, 1911: 209. Holotype female [not worker], ANGOLA:

Benguela, Cucala (J. Cruchet) (NM, Basle) [examined]. Syn. n. [Types of fortior and transversus originate

in a single series.]
Ocymyrmex arnoldi Forel, 1913fo: 138. Syntype workers, males, ZIMBABWE: Bulawayo (G. Arnold) (MHN,

Geneva) [examined]. Syn. n.
Ocymyrmex weitzekeri [sic] st. abdominalis Santschi, 1914a: 16. Syntype workers, SOUTH AFRICA: Natal,

Zululand, Entendweni, 20.viii.1905 (/. Trdgardh) (NM, Basle) [examined]. [Synonymized with arnoldi by

Arnold, 1916: 197.]

270 B. BOLTON

Ocymyrmex weitzaeckeri [sic] var. usakosensis Stitz, 1923: 146. Syntype workers, SOUTH WEST AFRICA:
Usakos, iv-vi.191 1 (W. Michaelsen) (syntypes presumed lost, not in MNHU, Berlin).Syn. n.

WORKER. TL 6-7-8-2, HL 1-68-2-00, HW 1-58-1-98, CI 94-99, SL 1-40-1-70, SI 85-91, PW 1-04-1-22,
AL 2-04-2-44 (20 measured).

Anterior clypeal margin with a semicircular median impression which is flanked on each side by a small
tooth or denticle. Maximum diameter of eye 0-36-0-40, about 0-20-0-23 x HW. Promesonotal dorsum
evenly shallowly convex in profile, the convex portion not strongly raised above the level of the propodeum
so that the slope of the posterior half of the mesonotum is very shallow indeed. Propodeal dorsum flat or
slightly sloping, rounding evenly into the declivity, the slope of which is quite steep but by no means vertical.
Metapleural lobes low and bluntly rounded, sometimes mostly concealed by the bulge of the metapleural
glands but usually easily visible. Peduncle of petiole commonly without a ventral process but quite fre-
quently a low rounded bulge is present, which in a few may be shorter and more prominent, forming a
broad, low and rounded angle. Petiole node small and low in profile, evenly rounded, the transition from
dorsal surface of peduncle to anterior face of node involving a marked change of slope. Petiole node in
dorsal view slender, small, varying from longer than broad to slightly broader than long, but the maximum
width of the node usually less than the length from the petiolar spiracle to the apex of the collar where
petiole and postpetiole articulate. Postpetiole in dorsal view longer than broad, sometimes only slightly so,
but usually the difference easily visible. Base of first gastral tergite strongly constricted and forming a narrow
neck behind the postpetiole Dorsum of head finely, densely and usually very regularly sharply longitudinally
costulate, the costulae usually parallel or nearly so over most or all of the area. In many samples all costulae
run straight back on the head, but commonly the outermost components tend to curve outwards behind the
eyes. Very rarely there is a tendency for the costulae to converge on the midline posteriorly, in which case a
few transverse members may be developed on the occipital surface. Ground-sculpture of fine punctulation is
present everywhere. Dorsal alitrunk densely costulate or rugose, the usual pattern being with arched
transverse sculpture on the anterior part of the pronotum followed by an area of longitudinal sculpture
which runs back just beyond the mesothoracic spiracles, followed by coarser transverse sculpture on the
remainder of the alitrunk dorsum. Exceptions to this are usually due to the extension of the longitudinal
component on the pronotum at the expense of the transverse. At its most extreme the longitudinal compo-
nent reaches forward almost to the cervical shield, and the other costulae are arched so steeply around it
that they appear longitudinal everywhere except on the extreme anterior part. Very rarely the longitudinal
costulae may extend back to the mesonotal-propodeal junction. In a few cases the costulae between the
mesothoracic spiracles are oblique, and now and then an individual is found in which the entire dorsal
alitrunk is transversely sculptured. Ventral surface of petiole with transverse rugulae of variable intensity,
usually fairly distinct but grading through to very faint. These rugulae may extend for some distance up the
sides of the node before fading out, but rarely reach the dorsum. Dorsum of peduncle and anterior and
posterior faces of node usually with weak transverse rugulae, very faint and scratch-like in places; the
dorsum of the node itself only rarely with vestiges of rugular sculpture, generally unsculptured or with a
superficial patterning. Postpetiole only with a superficial patterning or more or less smooth. All dorsal
surfaces of head and alitrunk with hairs of varying length, the hairs of the first gastral tergite much shorter
and sparser than on the alitrunk. Head and alitrunk varying from dull brick-red to lighter red, the two
always the same colour; gaster darker, blackish brown to black.

O. fortior is one of the more widely distributed and commoner species of the genus, ranging
widely from Angola to Zimbabwe and South Africa. Among the species with a strongly constric-
ted base to the gaster and a developed clypeal impression fortior is defined more by its lack of
specialized characters than the possession of them, as can be seen in the key. The closest related
species appear to be phraxus and micans. The former has a differently shaped petiole than fortior,
which is evenly rugulose dorsally on the node as opposed to the feebly or unsculptured surface
seen in fortior; phraxus is also darker in colour, appearing black with a red head to the naked eye.
O. micans has different cephalic structure from fortior and is also orange to orange-red in colour,
with a lighter yellowish gaster.

MATERIAL EXAMINED

Zambia: Mwengwa (H. Dollman). Zimbabwe: Bulawayo (G. Arnold); Khami Riv. (G. Arnold); Lonely
Mines (H. Swale); R. Zambesi (H. Swale); Victoria Falls (M. Grabham); Victoria Falls (W, L. Brown);
Bindurg (G. H. Bunzli). Botswana: R. Semowane (M. C. Day); between Kastwe and Damara Pan (H. Lang).
South Africa: Transvaal, Barberton (F. S. Parsons); Transvaal, Saltpan (H. Lang); Transvaal, Lydenburg
(H. Lang); Natal (G. Arnold).

SIX MINOR GENERA OF MYRMICINAE 271

, Ocymyrmex hirsutus Forel

Ocymyrmex weitzeckeri subsp. hirsutus Forel, 19106: 13. Syntype workers, SOUTH WEST AFRICA: Severelela

and Kooa (L. Schultze) (MHN, Geneva) [examined].
Ocymyrmex hirsutus Forel; Santschi, 1913: 431. [Raised to species.]

WORKER. TL 6-9-7-8, HL 1-54-1-82, HW 1-48-1-74, CI 93-97, SL 1-40-1-62, SI 90-95, PW 0-96-1-14,
AL 2-04-2-36 (7 measured).

Anterior clypeal margin with a conspicuous semicircular median impression which is flanked by a pair of
teeth or denticles. Occipital margin in full-face view slightly indented or flattened to feebly concave medially,
not evenly transversely convex. Maximum diameter of eye 0-32-0-38, about 0-21-0-23 x HW. Alitrunk in
profile with promesonotum evenly convex, sloping behind to the propodeum. Dorsum of propodeum
sloping very weakly, rounding broadly and evenly into the declivity. Metapleural lobes short and bluntly
triangular. Petiole in profile with a small, low, evenly rounded node, the peduncle without a ventral process
but broadly and very shallowly sinuate in some workers. Petiole node in dorsal view varying from as broad
as long to distinctly broader than long. Postpetiole dorsally longer than broad. Base of first gastral tergite
constricted and forming a narrow neck behind the postpetiole. Rugulose sculpture on dorsum of head fine,
irregular and very densely packed, with fine punctulate to granular ground-sculpture between the narrow
rugulae. To the level of the posterior margins of the eyes the rugulae are mostly or entirely longitudinal to
arched-longitudinal, the pattern varying from specimen to specimen. Behind this level the rugulae are
transverse or arched-transverse, tightly packed and narrowly vermiculate. In some the rugulae are so fine
and close together, and so narrowly vermiculate, that the occipital sculpture appears as a disorganised mass
of narrow irregular wiggly transverse lines. Dorsal alitrunk and declivity of propodeum transversely rugose
except between the mesothoracic spiracles where the sculpture is longitudinal. Extent of this longitudinally
sculptured area variable, the further forward the rugae extend the more strongly arched is the transverse
sculpture of the pronotum. In some the pronotal dorsum is mostly arched-longitudinally rugose. Sides of
alitrunk rugose, the sides of the pronotum finer and less densely so than the pleurae. Petiole with a few
transverse rugae ventrally and the peduncle also with a few dorsally, but otherwise the segment only
superficially sculptured. Postpetiole unsculptured. All dorsal surfaces of head and body except first gastral
tergite densely clothed in acute hairs of varying length, very numerous on the dorsal alitrunk. On the sides of
the pronotum the hairs are directed forwards; on the pleurae they point backwards and downwards. First
gastral tergite with sparse short hairs on the surface and with a denser transverse apical row. Colour reddish,
the gaster the same colour as the alitrunk or lighter.

Apart from its dense pilosity hirsutus can quickly be recognized by the distinctive sculpture of the
head, described above. Transverse sculpture on the area of the head behind the eyes is also found
in robecchii and in females of the various species of Ocymyrmex, but in all of these the sculpture
consists of regular transverse costulae, not narrowly vermiculate rugulae.

Ocymyrmex laticeps Forel
(Fig. 29)

Ocymyrmex laticeps Forel, 1901: 306. Syntype workers, ANGOLA: Mossamedes, Cubango-Cuito (H. H.
Braun & Van der Kellen) (MHN, Geneva) [examined].

WORKER. TL 7-2-7-6, HL 1-66-1-70, HW 1-70-1-74, CI 102-103, SL 1-56, SI 90-92, PW 1-08-1-14,
AL 2-20-2-24 (2 measured).

Middle of anterior clypeal margin flat to very feebly concave, without a semicircular notch or impression.
With head in full-face view the occipital margin with only the shallowest of faint indentations medially.
Maximum diameter of eye 0-36-0-38, about 0-21-0-22 x HW. Alitrunk in profile with promesonotum
evenly shallowly convex, the posterior part of the mesonotum very shallowly concave and sloping down to
the propodeum, the anterior half of which is itself slightly sloping; behind this the propodeum levels out
before rounding broadly and evenly into the declivity. Metapleural lobes vestigial, merely a thin laminar
strip on each side of the petiolar articulation, their width distinctly much less than (about half of) the width
of the propodeal spiracle. Petiole in profile as in Fig. 29, the anterior peduncle with a broad low triangular
process about half way along its ventral surface. Petiole node in dorsal view narrow, its maximum width
c. 0-35, less than the distance from the spiracle to the posterior collar of the petiole. Postpetiole in profile
swollen, low in front then forming an evenly convex low dome; the sternite strongly developed and bulging.
In dorsal view the postpetiole much longer than broad, gradually increasing in width from front to back.

272

B. BOLTON

First gastral tergite strongly constricted basally, forming a narrow neck behind the postpetiole. Dorsum of
head with longitudinal fine dense rugulae which become less regular and more disorganized away from the
midline and which tend to arch outwards posteriorly, towards the occipital corners, where they become
fainter. Spaces between the rugulae with punctulate ground-sculpture except in the median strip behind the
frontal lobes. Genae and sides of head below eyes regularly costulate, the costulae fine and dense, sharply
defined and fading out on the occipital corners. Sides of alitrunk costulate to rugose, the sculpture finer on
the sides of the pronotum, coarser elsewhere, regular except for an area in front of the propodeal spiracle
where some wavy rugae are present. Petiole and postpetiole unsculptured except for a vestigial superficial
patterning; the petiole ventrally with vestiges of a few transverse rugulae which are faint or incomplete.
Dorsal alitrunk transversely or arched-transversely rugulose except between the mesothoracic spiracles
where longitudinal sculpture is present. Convex portion of promesonotum less strongly sculptured than
remainder of dorsal alitrunk. All dorsal surfaces of head and body with sparse hairs which are apparently
absent from the first gastral tergite, but the specimens available have been much abraded so this is not
certain. Colour red, the alitrunk darker than the head and pedicel segments where the cuticle has an orange
tint. Base of gaster yellowish orange, much darker posteriorly.

This species, known only from the type-series, is closest related to cursor, which is also from
Angola. They differ as follows

laticeps
HW 1-70-1-74,

SL 1-56,

Smaller species,
PW 1-08-1-14.

Head slightly broader than long, CI 102-103.

Metapleural lobes vestigial, scarcely or not visible
in absolute profile.

Anterior half of propodeal dorsum sloping down-
wards, posterior half more or less level.

Petiole node in profile with a short, narrowly
rounded dorsum.

Postpetiole longer than broad in dorsal view.

cursor
Larger species, HW 2-04, SL 1 94, PW 1-30.

Head slightly longer than broad, CI 98.

Metapleural lobes large, conspicuous in absolute
profile.

Anterior half of propodeal dorsum shallowly con-
cave, posterior half rising and shallowly convex.

Petiole node in profile with an elongate, almost
flat dorsum.

Postpetiole broader than long in dorsal view.

Sculpture in the two species is very similar and they also share the characters of lacking a
semicircular clypeal impression and possessing a broad ventral process on the peduncle of the
petiole. Five other known species do not have a semicircular impression in the middle of the
anterior clypeal margin, cavatodorsatus, velox, ankhu, zekhem and turneri, but none of these has
the strongly developed constriction of the first gastral tergite seen in laticeps and cursor, and all
lack a subpetiolar process. Apart from this turneri is small (HW < 1-30) and has large eyes which
break the outline of the sides in full-face view; velox has a high, strongly convex petiole node
(Fig. 32) and very feeble cephalic sculpture; and cavatodorsatus has the alitrunk strongly saddle-
shaped in profile and the posterior half of the cephalic dorsum unsculptured.

Ocymyrmex micans Forel stat. n.

Ocymyrmex weitzeckeri var. micans Forel, 1910b: 12. Holotype worker, SOUTH WEST AFRICA: Okahandja
(Peters) (MHN, Geneva) [examined].

WORKER. TL 6-2-6-9, HL 1-54-1-70, HW 1-40-1-56, CI 91-92, SL 1-38-1-44, SI 92-98, PW 0-92-1-00,
AL 2-00-2- 16 (2 measured).

Anterior clypeal margin with a semicircular median impression which is flanked by a pair of small teeth.
Occipital margin very slightly concave or indented medially. Maximum diameter of eye 0-32-0-36, about
0-23 x HW. With alitrunk in profile the promesonotum evenly shallowly convex. Propodeal dorsum round-
ing broadly and evenly into the declivity. Metapleural lobes small and only feebly projecting, in profile
mostly or wholly concealed by the bulge of the metapleural glands. Petiole node well defined, the dorsum
narrowly and evenly rounded in profile, dome-like. Postpetiole swollen, the tergite low in front then rising
behind into a distinctly convex, smoothly rounded node. In dorsal view the petiole node broad and
conspicuous, its maximum width slightly greater than the distance from the spiracle to the apex of the
petiolar collar where it articulates with the postpetiole. Postpetiole in dorsal view narrow in front, becoming

SIX MINOR GENERA OF MYRMICINAE 273

much broader behind, the width greater than the length. Base of first gastral tergite strongly constricted and
forming a narrow neck behind the postpetiole. Sculpture of dorsum of head of dense, closely packed fine
longitudinal irregular rugulae which, away from the midline in the area behind the level of the eyes, curve
out towards the occipital corners. Behind the level of the eyes a conspicuous punctulate ground-sculpture is
present which in places may be the dominant component of the sculpture. Where this is the case the rugulae
are distinctly uneven, becoming wavy or even vermiculate. Dorsal alitrunk and propodeal declivity trans-
versely rugose except between the mesothoracic spiracles where the sculpture is longitudinal. On the
pronotum the rugae are arched-transverse around the longitudinal component. Sides of alitrunk regularly
rugose, the rugae weakest and most widely spaced on the sides of the pronotum. Petiole and postpetiole
unsculptured or the former with a few weak transverse rugulae ventrally and scattered vestigial marks
elsewhere. All dorsal surfaces of head and body with numerous hairs of varying length except for the first
gastral tergite where the hairs are short and sparse. Colour orange to orange-red, the gaster lighter in shade
than the alitrunk.

O. micans, known at present only from the Okahandja region of South West Africa, is closest
related to the widely distributed fortior. The two are separated on cephalic sculpture which is
stronger, more sharply defined and more regular in fortior, and the punctulate ground-sculpture
never dominates the rugulose/costulate component in this species. Besides this, the postpetiole of
fortior is relatively long and narrow, always longer than broad, whereas in micans the postpetiole
is distinctly swollen posteriorly and its width exceeds its length. Similarly, the petiole node in
dorsal view is always markedly expanded in micans, only moderately so in fortior.

MATERIAL EXAMINED.
South West Africa: Okahandja (M. C. Day).

Ocymyrmex monardi Santschi stat. n.

Ocymyrmex weitzeckeri st. monardi Santschi, 1930: 68. Syntype workers, ANGOLA: Cakindo (A. Monard)
(NM, Basle) [examined].

WORKER. TL 7-4-8-1, HL 1-78-1-88, HW 1-68-1-86, CI 95-99, SL 1-60-1-72, SI 92-95, PW 1-06-1-24,
AL 2-22-2-40 (3 measured).

Anterior clypeal margin medially with a distinct semicircular impression which is flanked on each side by
a small tooth. Eyes with maximum diameter 0-37-0-40, about 0-21-0-22 x HW. Promesonotum in profile
evenly rounded, sloping posteriorly to the propodeum which itself slopes very shallowly to the evenly
rounded posterior angle where the dorsum meets the declivity. Metapleural lobes small, broadly rounded to
truncate posteriorly, not triangular. Metapleural glands not strongly swollen nor strongly projecting
posteriorly, in profile not concealing even the bases of the metapleural lobes. Petiole in profile with the node
evenly rounded, dome-like; in dorsal view the node as broad as long or slightly longer than broad.
Postpetiole in dorsal view very slightly longer than broad. Base of first gastral tergite constricted. Sculpture
on dorsum of head characteristic; space between frontal lobes and median strip of dorsum behind the
frontal lobes finely and densely longitudinally costulate, with feeble punctulate ground-sculpture between
the costulae. Behind the level of the eyes the costulae of this median area become much weaker and show
signs of diverging, or fade out altogether, being replaced partially or entirely by coarse punctulate or
granular sculpture which is very dense and conspicuous. The space between the inner margin of the eye and
the antennal fossa, and the area extending back from this level to the occiput covered with dense irregular
granular sculpture. Dorsal alitrunk and propodeal declivity transversely rugose except for the space between
the mesothoracic spiracles and the median strip of the pronotum in front of this level, where the rugae are
longitudinal. Petiole and postpetiole unsculptured except for faint superficial patterning or the former at
most with a few vestigial transverse rugulae ventrally. All dorsal surfaces of head and body with hairs, but
those on the first gastral tergite shorter and much sparser than on the alitrunk, where strong hairs are
conspicuous. Alitrunk dull red to reddish tinted black, the head somewhat lighter in shade.

This moderately sized quite darkly coloured species is closely related to fortior, but is separated
from it and from other close forms by the distinctive pattern of sculpture on the head.

MATERIAL EXAMINED
South West Africa: Swakopmund (M. C. Day).

274 B. BOLTON

Ocymyrmex nitidulus Emery stat. n.

(Figs 18, 22, 23)

Ocymyrmex robecchii st. nitidulus Emery, 1892: 116. Holotype female [not worker], SOMALI REPUBLIC:
Obbia (Robecchi) (MCSN, Genoa) [examined].

WORKER. TL 7-4-7-8, HL 1-70-1-90, HW 1-64-1-82, CI 96-100, SL 1-48-1-60, SI 86-96, PW 1-08-1-20,
AL 2-20-2-30 (20 measured).

Anterior clypeal margin with a conspicuous median semicircular impression which is flanked on each side
by a low tooth or denticle, this tooth usually quite low and rounded, uncommonly strongly prominent.
Maximum diameter of eye 0-34-0-40, about 0-20-0-22 x HW. Shape of alitrunk as in Fig. 18, the promes-
onotum evenly and broadly convex, sloping posteriorly to the propodeal dorsum; the latter rounding evenly
into the declivity. Metapleural lobes small and low, bluntly rounded or broadly and bluntly truncated,
sometimes reduced to a narrow little-projecting strip. Petiole node in profile small, low and smoothly
rounded. In dorsal view the petiole node varying from slightly longer than broad to slightly broader than
long. Postpetiole as broad as long to slightly broader than long in dorsal view. First gastral tergite
constricted and forming a neck basally. Dorsum of head sculptured with low, rounded, feeble longitudinal
costulae which may be very reduced or even effaced in places. Spaces between the costulae without ground-
sculpture, the surface smooth and polished, with a slick and glossy appearance. Sides of alitrunk closely and
finely sharply constulate, the dorsum sharply transversely costulate except between the mesothoracic spira-
cles where the sculpture is usually longitudinal. Pronotal dorsum in front of the longitudinal sculpture finely
arched-costulate to smooth, the sculpture always weaker than elsewhere on the alitrunk, sometime partially
or wholly effaced or the costulae reduced to vestiges. This weakening of the sculptural intensity may also
affect the longitudinal costulae between the mesothoracic spiracles. Ventral surface of petiole node with a
few transverse rugulae which may be very feeble but which normally extend for some distance up the sides of
the node before fading out. Dorsum of petiole node unsculptured or at most with vestiges remaining.
Dorsum of peduncle usually with transverse fine rugulae but these may be very reduced and faint. Postpet-
iole unsculptured or at most with some fine superficial patterning. All dorsal surfaces of head and alitrunk
with numerous hairs of varying length, the first gastral tergite with hairs shorter and sparser than elsewhere.
Colour glossy dull red, the gaster darker, blackish brown to black; entire body with a polished and shining
appearance.

Only nitidulus and celer, among the species with the clypeus impressed and gaster constricted
basally, lack strong ground-sculpture between the costulae or rugulae of the dorsal head. This
gives them a slick and very polished appearance which immediately separates them from their
allies in which granular or punctulate ground-sculpture is present, and which in consequence are
dull and less polished. 0. nitidulus and celer are separated by the shape of the pronotum, which is
flat in the latter, rounded in the former (compare Figs 18 and 19), and by the shape of the head
which is longer and narrower in celer than in nitidulus.

MATERIAL EXAMINED

Ethiopia: Meisso (K. Guichard). Kenya: Kajiado (G. Nyamasyo), Kajiado (J. Darlington). Uganda: N.
Turkana Prov. (Lake Rudolf expd.). Tanzania: Longido West (A. Loveridge).

Ocymyrmex phraxus sp. n.

(Fig. 27)

HOLOTYPE WORKER. TL 7-9, HL 1-92, HW 1-82, CI 95, SL 1-58, SI 87, PW 1-15, AL 2-32.

Anterior clypeal margin with a deep median impression which is flanked on each side by a tooth. Sides of
head in front of eyes more or less parallel, the head not broadening anteriorly. Maximum diameter of eye
0-38, about 0-21 x HW. Promesonotum forming an even, low convexity in profile which slopes gently
downwards posteriorly. Propodeal dorsum rounding evenly into the declivity. Metapleural lobes low and
rounded. Petiole in profile with a small node which is not sharply differentiated from the peduncle; the
dorsum of the peduncle runs into the anterior face of the node without a marked change in slope. This
surface is confluent behind with the dorsum, which is low and broadly evenly rounded. In dorsal view the
node is narrow and almost parallel-sided, its maximum width not much greater than that of the posterior
peduncle. Postpetiole longer than broad in dorsal view, discounting the anterior articulatory section. First
gastral tergite constricted basally, forming a narrow neck behind the postpetiole. Dorsum of head densely
sculptured with fine, sharply defined parallel longitudinal rugulae which are slightly divergent posteriorly

SIX MINOR GENERA OF MYRMICINAE 275

and more divergent laterally where they tend to arch outwards behind the eyes. Spaces between the rugulae
filled with a fine dense and conspicuous punctulate ground-sculpture. Pronotum with arched-transverse
costulae, with a patch of longitudinal sculpture between the mesothoracic spiracles. Remainder of dorsal
alitrunk transversely rugose except for a small disorganized patch between the propodeal spiracles. Alitrunk
dorsally lacking the punctulate ground-sculpture seen on the head. Petiole with a number of very fine
rugulae which encircle the node, the postpetiole only with fine superficial patterning, or light shagreening.
Dorsal surfaces of head and alitrunk all with numerous hairs, those on the first gastral tergite much shorter
and sparser than those on the alitrunk. Alitrunk a dark, deep red, dully shining. Head dark red but lighter in
shade than the alitrunk, the contrast easily visible to the naked eye. Caster blackish brown to black, darker
than the rest of the body.

PARATYPE WORKERS. TL 7-7-8-0, HL 1-90-1-96, HW 1-80-1-86, CI 94-96, SL 1-56-1-60, SI 85-89, PW
1-14-1-16, AL 2-26-2-34 (5 measured). Maximum diameter of eye 0-36-0-38, about 0-20-0-21 x HW. As
holotype but some with the petiole node slightly broader in dorsal view and with the disorganized rugular
patch on the propodeal dorsum less obviously developed.

Holotype worker, Tanzania : Shinyanga, viii.1949 (B. K. Coll.) (BMNH).
Paratypes. 4 workers with the same data as holotype (BMNH ; MHN, Geneva).
Non-paratypic material examined. Tanzania: no further data.

The non-paratypic series matches the type-series well but has sculpture slightly more intense in
development and has some workers which are slightly smaller, HL 1-78-1-90, HW 1-70-1-82,
CI 95-96, SL 1-54-1-64, SI 90-93.

The sculpture of the petiole in phraxus is reminiscent of a reduced version of that found in
sobek, but in the latter the node is greatly swollen and very distinctive (Fig. 20), and the gaster is
yellowish, much lighter than the alitrunk.

Ocymyrmex picardi Forel

Ocymyrmex picardi Forel, 1901: 306. LECTOTYPE worker, ANGOLA: Mossamedes, Cubango-Cuito

(MHN, Geneva), here designated [examined].
Ocymyrmex carpenter! Donisthorpe, 1933: 195. Holotype female [not worker], BOTSWANA: Ngamiland,

ix.1930-i.1931 (G. D. H. Carpenter) (BMNH) [examined]. Syn. n.

Note. The number of specimens in the original syntypic series of picardi was not stated by Forel.
At the time of this study only two specimens, mounted on a single pin, were found bearing the
label 'typus'. Of these the top specimen is a worker and fits the original description perfectly; it is
here designated as lectotype of picardi. The lower specimen, now remounted on a separate pin, is
a female and is not mentioned in the original description.

WORKER. TL 11-6-12-6, HL 2-52-2-80, HW 2-44-2-68, CI 93-97, SL 2-30-2-52, SI 90-98, PW 1-60-1-80,
AL 3-40-3-68 (20 measured).

Very large species. Anterior clypeal margin with a conspicuous median semicircular impression which is
flanked by a small tooth on each side. Maximum diameter of eye 0-46-0-51, about 0-19-0-20 x HW. With
the head in full-face view the occipital margin varying from approximately transverse to feebly indented
medially. Promesonotum in profile evenly and broadly convex, sloping posteriorly to the propodeal dorsum
which, in this species, is not as strongly depressed below the level of the promesonotum as is usual elsewhere
in the genus. Propodeal dorsum rounding broadly and evenly into the declivity. Metapleural lobes rounded,
visible in profile, not concealed by the bulge of the metapleural glands. Petiole node in dorsal view as broad
as or broader than long. Postpetiole slightly longer than broad in dorsal view, discounting the anterior
articulatory portion. Base of first gastral tergite narrow, in dorsal view no broader than the postpetiole, but
the sides evenly diverging from their junction with the postpetiole, without a roughly parallel-sided neck.
Dorsum of head finely, densely and more or less evenly longitudinally costulate, the costulae rarely approxi-
mately straight, much more commonly diverging behind towards the occipital corners. Infrequently a few
transverse costulae may be present occipitally. Individual costulae commonly irregular, tending to be
narrowly sinuate or wavy, especially away from the midline of the dorsum. Ground-sculpture a fine dense
punctulation. Dorsal alitrunk and propodeal declivity transversely rugose except for the space between the
mesothoracic spiracles and part of the pronotal dorsum, where sculpture is longitudinal to oblique. Sides of
alitrunk rugose, the sculpture less regular on the pleurae than on the sides of the pronotum. Petiole with
transverse rugae ventrally, below the node, and also with a few dorsally on the peduncle in front of the node.

276 B. BOLTON

On the node itself sculpture is usually restricted to a superficial patterning with vestigial rugulae, but
occasionally one or two stronger transverse rugulae may be present dorsally, or vertically on the sides, or
both. Postpetiole unsculptured except for superficial patterning. All dorsal surfaces, of body with scattered
strong dark hairs which are reddish brown to blackish; those on the first gastral tergite very sparse and
much shorter than those on the alitrunk. Colour very dark red to black, the head usually slightly lighter in
shade than the alitrunk and the gaster generally darker.

A very conspicuous species, the largest known in the genus, picardi appears to be quite widely
distributed in southern Africa. By its size alone it is unlikely to be confused with any other
species.

The female (queen) of picardi was first described by Donisthorpe as a worker, under the name
of carpenteri. The holotype matches the female in the same series as the picardi lectotype and the
synonymy is thus absolute. The female of picardi fits the description given above and its dimen-
sions fall within the ranges given. The only differences from the worker lie in those characters
discussed under the generic diagnosis, namely the broader straighter margins to the frontal lobes,
broader antennal scapes and sharp transverse sculpture on the posterior portion of the dorsum of
the head.

MATERIAL EXAMINED

Zimbabwe: Umgusa Riv., Sawmills (G. Arnold): Insiza Riv. (no name). Botswana: Sevrelela (L. Schultze);
Okavango Delta, Smiti (A. Russell-Smith); Kalabura (ex coll. Donisthorpe); Tsabong (G. Arnold); Nkate
(Vernay-Lang expd.); Matopo Pan (G. V. Son); Shaleshonto (G. U. Son). South West Africa: Kalahari
Desert (no name).

Ocymyrmex robecchii Emery
(Fig. 24)

Ocymyrmex robecchii Emery, 1892: 1 14, fig. Syntype workers, SOMALI REPUBLIC: Uebi (Robecchi), and Erdal
(Pavesi) (MHN, Geneva; MCSN, Genoa) [examined].

WORKER. TL 9-2-9-3, HL 2-20-2-22, HW 2-02-2-04, CI 92, SL 2-08-2-16, SI 103-106, PW 1-26-1-28, AL
2-52-2-60 (2 measured).

Anterior clypeal margin with a strong semicircular median impression flanked by a pair of short rounded
denticles. Outline shape of head as in Fig. 24, the head longer than broad (CI < 100), with sides which
converge posteriorly, and with the occipital margin strongly impressed medially. Maximum diameter of eye
0-40, about 0-20 x HW. Scapes long, SI > 100. With alitrunk in profile the mesonotum anteriorly forming a
distinct hump behind the pronotum, the two not forming a single continuous convexity. Orifices of meso-
thoracic spiracles protected by a pair of low tumuli or welts which project from the surface. Mesonotal
dorsum sloping shallowly downwards posteriorly and confluent with the propodeal dorsum which is flat to
very feebly concave to the level of the spiracle where it becomes shallowly convex and rounds broadly and
evenly into the declivity. Metapleural lobes large and strongly developed, prominent, broadly subtriangular
in shape and slightly upcurved. Peduncle of petiole without a ventral process. Petiole node in dorsal view
very narrow, slightly expanded. Postpetiole in dorsal view much longer than broad. Basal portion of first
gastral tergite constricted, forming a narrow neck behind the postpetiole. Dorsum of head with arched-
longitudinal costulae on the genae and frontal lobes, the costulae curving in towards the midline posteriorly.
Remainder of dorsum to occipital margin densely transversely costulate. Ground-sculpture vestigial, merely
a superficial patterning between the costulae. Sides of alitrunk strongly, regularly and sharply costulate
everywhere, the dorsum similarly sculptured, as is the propodeal declivity. All dorsal sculpture transverse
except between the mesothoracic spiracles. Petiole encircled by fine rugulae or costulae which are most
strongly developed ventrally. Postpetiole and gaster unsculptured. All dorsal surfaces of head and body with
scattered acute hairs of varying length; those on the first gastral tergite shorter and sparser than elsewhere.
Colour dark dull red, the postpetiole and gaster lighter, yellowish.

A large and conspicuous species, robecchii is characterized by its strongly impressed median
clypeal notch, transverse cephalic sculpture, strongly impressed occipital margin, long scapes and
large size. Although the two specimens seen possess the transverse cephalic sculpture character-
istic of females in this genus, they lack the associated characters of shorter thicker scapes and
broader, more parallel-sided frontal lobes which are usually associated with this caste (see
discussion under the generic diagnosis). For this reason I have chosen to treat these specimens as

SIX MINOR GENERA OF MYRMICINAE 277

bona fide workers for the time being, and have used the direction of cephalic sculpture as a key
character in case they do not turn out to be females. This of course cannot be ascertained until
more material becomes available.

Ocymyrmex shushan sp. n.

(Fig. 31)

HOLOTYPE WORKER. TL 6-7, HL 1-66, HW 1-58, CI 95, SL (antennae missing), PW 1-02, AL 2-14.

Anterior clypeal margin with a small but distinct median semicircular impression, which is flanked by a
pair of short, acute teeth. Maximum diameter of eye 0-34, about 0-22 x HW. Sides of head behind eyes
evenly convex and narrowing, rounding into the occipital margin with which, if it were not for the median
impression in the latter, they would form a very regular arc. In profile the posterior pronotum and anterior
part of mesonotum high and strongly convex, forming a dome-like outline above the level of the mesothora-
cic spiracles. Pronotum in front of this and mesonotum behind it sloping away evenly, increasing the
dome-like appearance. Posterior portion of mesonotum and dorsum of propodeum more or less flat in
profile, the latter rounding broadly into the convex declivity. Metapleural lobes small and low, rounded.
Peduncle of petiole flat dorsally but sinuate ventrally. Petiole node in profile low dome-like, smoothly and
evenly rounded. Postpetiole in profile subglobular, Fig. 31. In dorsal view the petiole node much broader
than long, all surfaces smoothly rounded; postpetiole slightly longer than broad. Base of first gastral tergite
constricted and forming a neck behind the postpetiole. Dorsum of head finely and densely longitudinally
rugulose, the rugulae diverging away from the midline posteriorly. Ground-sculpture between the rugulae a
strong and conspicuous punctulation which in places seems as strongly developed as the rugular sculpture.
Pronotum with a few strong transverse costulae on the cervical shield but behind this the costulae which
arch up from the sides become very faint or vestigial when traversing the dorsum. Between the mesothoracic
spiracles and extending for a short distance forwards and backwards is a patch of low longitudinal rugosity.
Remainder of alitrunk and propodeal declivity transversely rugose or costulate. Petiole node smooth and
shining, the anterior peduncle with some very faint transverse striae. Postpetiole unsculptured. Body every-
where lacking the conspicuous punctulate ground-sculpture which is so well developed on the head. All
dorsal surfaces of head and body with numerous hairs, those on the first gastral tergite shorter and sparser
than those on the alitrunk. Colour uniform bright orange.

Holotype worker, South West Africa : Gobasis, 17.xii.1933 (J. Ogilvie) (BMNH).

This bright orange species is characterized amongst forms with both a gastral constriction and an
impressed clypeal margin by the strongly convex anterior portion of the dorsal alitrunk and the
shape of the petiole, as described above.

Ocymyrmex sobek sp. n.

(Fig. 20)

HOLOTYPE WORKER. TL 7-8, HL 1-90, HW 1-82, CI 96, SL 1-70, SI 93, PW 1-18, AL 2-44.

Anterior clypeal margin with a conspicuous median semicircular impression which is flanked by a pair of
small teeth. Occipital margin in full-face view feebly indented medially, the margin continuous with the sides
through a broad, shallow curve. Maximum diameter of eye 0-40, about 0-22 x HW. Promesonotum in
profile evenly convex, the dorsum of the propodeum rounding broadly and evenly into the declivity.
Metapleural lobes small, subtriangular in shape, but plainly visible in profile and not concealed by the bulge
of the metapleural glands. Ventral surface of peduncle of petiole without a projecting process, the node
strongly swollen and conspicuous (Fig. 20). In profile the large node with a developed posterodorsal angle,
not rounded as is usual in the genus. In dorsal view the petiole node massive, almost as large as the
postpetiole; the latter also somewhat swollen, about as high as long in profile and broader than long in
dorsal view. Base of first gastral tergite constricted, in dorsal view forming a neck behind the postpetiole
which is narrower than the maximum width of the postpetiole itself. Dorsum of head finely and densely
costulate to rugulose, the sculpture most regular on the central strip of the dorsum. On each side of this
central strip the rugulae more wavy and irregular, and tending to diverge posteriorly towards the occipital
corners, arching round above the eyes. Spaces between the rugulae with fine dense punctulate ground-
sculpture. Anterior portion of pronotum with arched-transverse rugae, behind this with an area where the
rugae are almost longitudinal, very slightly oblique, running back between the mesothoracic spiracles;
remainder of dorsum and also propodeal declivity coarsely transversely rugose. Sides of alitrunk coarsely

278 B. BOLTON

rugose. Node of petiole coarsely rugose, the rugae continuous, running transversely on the ventral surface,
up the sides of the node and across the dorsum; on the dorsum less regular than elsewhere. Anterior face of
node and dorsum of peduncle in front of node also transversely rugose. Postpetiole unsculptured except for
fine superficial shagreening. Dorsal surfaces of head, alitrunk and pedicel segments with numerous strong
hairs. First gastral tergite with hairs much shorter and sparser than elsewhere. Alitrunk dull reddish black
(appearing black to the naked eye); gaster dull yellowish red, much lighter than the alitrunk and contrasting
strongly with it; head a dull red intermediate in shade between gaster and alitrunk so that the ant appears
tricoloured to the naked eye.

PARATYPE WORKERS. TL 7-6-8-0, HL 1-70-1-90, HW 1-64-1-88, CI 93-100, SL 1-56-1-74, SI 92-99,
PW 1-08-1-20, AL 2-20-2-44 (11 measured). Maximum diameter of eye 0-36-0-40, about 0-22-0-24 x HW.
Answering description of holotype but some with the posterodorsal angle of the petiole more rounded and
the pronotal sculpture showing the usual variation, with differences in extent and direction of the longitudi-
nal component. In most the longitudinal rugae are antero-posteriorly straight, but in some (as in the
holotype) they are oblique and in a few decidedly transverse. Sculpture pattern on the dorsum of the head is
usually as described above but in a few the rugulae run straight back everywhere, not diverging towards the
occipital corners, and in one the rugulae between the eye and the central strip of the head are more or less all
transverse.

Holotype worker, Botswana : Smiti, no. 16, mopane woodland, 1 l.ix.1975 (A. Russell-Smith) (BMNH).

Paratypes. 11 workers with same data as holotype (BMNH; MCZ, Cambridge; NM, Basle; MHN,
Geneva; NM, Bulawayo).

Non-paratypic material examined. Zimbabwe: Victoria Falls (G. Arnold); Victoria Falls (H. Swale); Vic-
toria Falls (M. Grabham). Botswana: Kabulabula (G. U. Son).

Size range in the non-paratypic material is TL 7-6-8-8, HL 1-80-2-00, HW 1-70-1-96, CI 93-98,
SL 1-64-1-80, SI 91-98, PW 1-12-1-28, AL 2-24-2-52 (15 measured). Measurements of eyes all fall
within the range of the paratype-series.

This strongly sculptured and conspicuously coloured species is closest related toforeli. Among
the species with a clypeal impression and a distinctly constricted first gastral segment only two,
sobek andforeli, have the petiole much enlarged and very strongly sculptured. The two are best
separated on their colour, as indicated in the key, but it is also notable that the rugae on the
petiole node are narrower and more sharply defined inforeli than in sobek, where they tend to be
blunted and much less regular on the dorsum.

Ocymyrmex sphinx sp. n.

(Fig. 21)

HOLOTYPE WORKER. TL 9-3, HL 2-18, HW 2-04, CI 94, SL 1-96, SI 96, PW 1-32, AL 2-86.

Large species. Anterior clypeal margin with a conspicuous semicircular impression medially which is
flanked on each side by a low but broad triangular tooth. Occipital margin feebly indented medially in
full-face view. Maximum diameter of eye 0-43, about 0-21 x HW, the eyes distinctly failing to break the
outline of the sides of the head in full-face view. Promesonotum rounded in profile, sloping posteriorly to the
propodeum which is almost flat and which rounds evenly into the steep declivity. Metapleural lobes small,
narrowly rounded and prominent, in absolute profile just visible behind the bulge of the projecting meta-
pleural glands. Petiole node small and rounded in profile, the peduncle with a feebly convex area midway
along its ventral surface but without a developed process. Petiole node in dorsal view distinctly broader
than long, strongly developed, constricted behind at the posterior peduncle. Postpetiole about as broad as
long in dorsal view, excluding the anterior articulatory portion. Base of first gastral tergite narrow, no wider
than the postpetiole, but not forming as conspicuously narrowed a neck as is usual in this genus. Instead the
gaster begins to widen gradually almost immediately behind its articulation with the postpetiole, the sides at
first gradually and then more strongly divergent in dorsal view. Dorsum of head finely, densely and
irregularly longitudinally rugulose everywhere, with dense conspicuous coarse punctulate to granular
ground-sculpture between the rugulae. On the central strip of the dorsum the rugulae are most regular
behind the frontal lobes, posteriorly they become narrowly vermiculate. Occipitally and laterally on the
dorsum the rugulae are narrowly vermiculate everywhere and tend to arch outwards behind the eyes. Rugae
on pronotal dorsum arched-transverse anteriorly. An area of longitudinal sculpture occurs between the
mesothoracic spiracles, visible in paratypes, but in the holotype a pin is inserted at this point and the

SIX MINOR GENERA OF MYRMICINAE 279

sculpture destroyed. Remainder of dorsal alitrunk and declivity coarsely transversely rugose. Petiole and its
peduncle finely transversely rugose ventrally; behind the level of the spiracle the rugae continuing up the
sides of the node and across the dorsum. Transverse rugae also present on dorsum of peduncle. Dorsum of
postpetiole finely granular and matt. All dorsal surfaces of head and alitrunk densely clothed with hairs,
many of them quite short. Propodeum dorsally also with dense pubescence. Hairs on first gastral tergite very
short and sparse. Colour a very dull dark red, the gaster black.

PARATYPE WORKERS. TL 9-5-10-1, HL 2-16-2-28, HW 2-08-2-14, CI 93-96, SL 1-95-2-06, SI 94-98,
PW 1-38-1-44, AL 2-90-3-02 (12 measured). Maximum diameter of eye 0-44-0-46, about 0-20-0-22 x HW.
As holotype but some showing shorter pilosity than others, apparently due to the hairs having been broken
off.

Holotype worker, Botswana: 18 miles [29 km] NE. of Kalkfontein, 12-13.iv.1972, no. B3 (M. C. Day)
(BMNH).

Paratypes. Botswana: 19 workers, Kuke Pan, 26.iii.1930, Vernay-Lang Kalahari Expd. (G. U. Son) (MCZ,
Cambridge; BMNH; NM, Bulawayo).

At first glance this species appears to be a smaller version of picardi but, apart from being
consistently smaller it has silvery body hairs where those of picardi are dark reddish brown to
blackish, has much denser pilosity and pubescence on the propodeal dorsum and has much
stronger, denser and more sharply defined ground-sculpture on the dorsum of the head than is
seen in picardi. The cephalic sculpture of sphinx is much the same as that seen in monardi, but this
latter species is much smaller and has the petiole node unsculptured.

Ocymyrmex turneri Donisthorpe
(Fig 25)

Ocymyrmex turneri Donisthorpe, 1931 : 499. Holotype worker, SOUTH WEST AFRICA: Walvis Bay, l.xii.1927
(R. E. Turner) (BMNH) [examined].

WORKER. TL 5-8, HL 1-40, HW 1-28, CI 91, SL 1-40, SI 109, PW 0-80, AL 1-76.

Small species. Anterior clypeal margin without a median impression, on the contrary the middle of the
anterior clypeal margin projecting forward as a low, broad triangular prominence. Eyes relatively large,
maximum diameter 0-36, about 0-28 x HW. In full-face view the eyes very conspicuously breaking
the outline of the sides of the head. Antennal scapes relatively long, SI > 100. Promesonotum in profile
rounded, with a low transversely arched crest running across the dorsum at the junction of pronotum and
mesonotum, arching forward from the mesothoracic spiracle on each side. Posterior part of mesonotum and
anterior part of propodeum concave in profile, the posterior portion of the propodeum convex and curving
evenly into the declivity behind. Petiole node in profile high, subconical, with a narrowly rounded dorsum,
the posterior face convex, the anterior face longer, almost flat and less steeply sloped than the posterior face.
Petiole node in dorsal view longer than broad, the postpetiole broader than long. First gastral tergite
without a constricted basal neck. Dorsum of head glossy, sculptured with very feeble superficial rugulae
which are transversely arched and are almost effaced occipitally. Ground-sculpture between the faint
rugulae absent. Dorsal alitrunk unsculptured, smooth and highly polished except for vestiges of faint
transverse rugulae on the extreme anterior portion of the pronotum and the propodeal declivity. On the
sides of the alitrunk the pleurae with strong, widely spaced and roughly parallel rugae, otherwise unsculp-
tured except for vestiges on the sides of the pronotum near the base. Petiole and postpetiole only with faint
superficial patterning which is almost effaced in places. All dorsal surfaces of head and body with scattered
hairs of varying length but the holotype (and only known specimen) appears to be considerably abraded and
in life this species may well be as hairy as other species in the genus. Alitrunk jet black and shiny, head and
gaster blackish brown.

0. turneri is one of the smallest species known in the genus and is easily distinguished from all
others by having an unconstricted base to the gaster, large eyes which break the outline of the
sides of the head, reduced sculpture, relatively long antennal scapes and a clypeal margin which
projects medially into a low triangular prominence. To date it remains known only from the
holotype.

280 B. BOLTON

Ocymyrmex velox Santschi
(Fig. 32)

Ocymyrmex velox Santschi, 1932: 387. Syntype workers, SOUTH WEST AFRICA: Otjimbimbi, Kunene R.,
Hi. 1923 (G. Arnold) (NM, Basle; NM, Bulawayo) [examined].

WORKER. TL 10-0-10-7, HL 2-20-2-40, HW 2-08-2-30, CI 95-97, SL 2-10-2-29, SI 101-105, PW 1-30-
1-42, AL 2-48-2-96 (8 measured).

Large species; anterior clypeal margin entire or very feebly eroded medially giving a weakly crenellated
appearance, without a median semicircular impression. Maximum diameter of eyes 0-40-0-43, about 0-19-
0-20 x HW. Frontal lobes distinctly convergent posteriorly, the occipital margin indented medially in
full-face view. Antennal scapes relatively long, SI above. Promesonotum forming an even low convexity in
profile which may be somewhat flattened above in some individuals. Propodeal dorsum more or less flat,
usually sloping shallowly downwards posteriorly to the evenly rounded junction with the declivity; the
alitrunk not saddle-shaped. Metapleural lobes low and rounded. Petiole node relatively large in profile,
high, dome-like and evenly rounded, the anterior peduncle without a ventral process. In dorsal view the
petiole node longer than broad. Postpetiole in dorsal view slightly broader than long. Base of first gastral
tergite in dorsal view not forming a narrow bottle-neck-like constriction. The tergite basally is no wider than
the postpetiole but the sides diverge evenly from immediately behind the articulation. Sculpture of head very
feeble, faint and even effaced in places, at most consisting of a few weak and superficial irregular rugulae.
Ground-sculpture where present only of a weak superficial patterning or shagreening. Dorsal alitrunk
transversely rugulose, with or without a patch of longitudinal sculpture on the pronotum and between the
mesothoracic spiracles. Rugulae on the propodeal dorsum may be irregular or broken. Petiole and postpe-
tiole dorsally only with superficial patterning, or the former with faint rugular vestiges. Ventral surface of
petiole with weak transverse rugulae which may extend up the sides of the node and onto the dorsum, but
these are vestigial in some individuals. Dorsal surfaces of head and alitrunk with hairs but these are very rare
or even absent on the first gastral tergite. Alitrunk dull red to blackish red, the head a lighter red and the
gaster lighter still, orange or even yellow in some individuals.

0. velox is one of the seven known species in this genus which lack a conspicuous semicircular
impression in the middle of the anterior clypeal margin. The others are turneri, laticeps, cursor,
ankhu, zekhem and cavatodorsatus. Of these turneri is a small shiny species with large eyes which
break the outline of the sides of the head, and cursor and laticeps have the dorsum of the head
strongly sculptured and have the first gastral tergite constricted into a narrow neck basally.
0. cavatodorsatus shares most characters with velox, but the shape of the alitrunk is very dif-
ferent; besides this cavatodorsatus is much smaller and has the sculpture of the alitrunk much
reduced. Separation of velox from ankhu and zekhem is discussed under those names.

MATERIAL EXAMINED
South West Africa: Otjiwarongo (G. Hobohm). Angola Cahama (E. S. Ross & R. E. Leech).

Ocymyrmex weitzeckeri Emery
(Fig. 30)

Ocymyrmex weitzeckeri Emery, 1892: 116. Syntype workers, LESOTHO: Leribe (Weitzecker) (MCSN, Genoa)

[examined].
Ocymyrmex weitzeckeri subsp. wroughtoni Forel, 1910b: 13. Syntype workers, male, SOUTH AFRICA: Natal

( Wrought on) (MHN, Geneva) [examined]. Syn. n.

WORKER. TL 6-8-7-8, HL 1-70-1-80, HW 1-62-1-72, CI 94-99, SL 1-44-1-58, SI 88-94, PW 1-02-1-10, AL
2- 16-2- 30 (8 measured).

Anterior clypeal margin with a conspicuous semicircular median impression flanked by a pair of short
teeth. Occipital margin in full-face view with a median indentation. Maximum diameter of eyes 0-36-0-40,
about 0-22-0-23 x HW. Promesonotum evenly convex in profile, the propodeal dorsum flat to very shal-
lowly convex, rounding evenly into the declivity which is almost vertical. Metapleural lobes small but
prominent, their apices narrowly rounded. Petiole in profile as in Fig. 30. Petiole node in dorsal view very
broad, distinctly broader than long and its width greater than the distance from the spiracle to the apex of
the collar where the petiole articulates with the postpetiole. Postpetiole node in dorsal view broader than
long (excluding the anterior articulating portion). Base of first gastral tergite strongly constricted behind the

SIX MINOR GENERA OF MYRMICINAE 281

postpetiole and forming a distinct neck. Head finely and densely rugulose everywhere, with punctulate or
granular ground-sculpture. Pattern of the rugulae varying between individuals. On the dorsum from the
level of the eyes to the occiput the rugulae never all running straight back. Either the rugulae away from the
median strip diverge towards the occipital corners, or the rugulae close to the inner or posterior margins of
the eyes are irregular to vermiculate; in some the rugulae are extensively vermiculate on the head. Dorsal
alitrunk and propodeal declivity transversely rugose except for the area between the mesothoracic spiracles
where the sculpture is longitudinal. Other components of the pronotal sculpture arch around the anterior
end of these longitudinal rugae. In some workers the pronotum appears to be entirely longitudinally rugose
in dorsal view as the median rugae extend so far forward that the more laterally situated rugae must parallel
them almost to the cervical shield before arching round. Petiole with a few transverse rugae ventrally which
may extend for some distance up the sides ; the node with a few weak to vestigial transverse rugulae, those
traversing the peduncle in front of the node stronger than those on the node itself. Postpetiole unsculptured
except for the usual faint superficial patterning. All dorsal surfaces of head and body with fairly dense
pilosity, the hairs of varying length; those on the first gastral tergite shorter and sparser than elsewhere on
the body. Colour everywhere dark red to blackish red, in some specimens very dark indeed, almost entirely
black.

Known only from the two type-series above and the two short series recorded below, weitzeckeri
has served as the base from which many infraspecific forms have been described in the past. Of all
the names formerly attached to it only wroughtoni has proved to be synonymous; the treatment
of the remainder is summarized under the synonymic list of species.

MATERIAL EXAMINED
South Africa : Orange Free State, Donga (R. W. Slobey); Transvaal, no loc. (G. Arnold).

Ocymyrmex zekhem sp. n.

HOLOTYPE WORKER. TL 84, HL 2-06, HW 1-90, CI 92, SL 2-20, SI 116, PW 1-40, AL 2-50.

Anterior clypeal margin entire, without trace of a median notch or impression. Maximum diameter of eye
0-40, about 0-21 x HW, the eyes only just failing to break the outline of the sides in full-face view. Sides of
head in front of eyes straight, diverging anteriorly; behind the eyes the sides rounding broadly and evenly
into the occipital margin, the latter very shallowly impressed medially in full-face view. Antennal scapes the
longest yet known in the genus (see SI, above). With the alitrunk in profile the promesonotum low, evenly
shallowly convex. Posterior part of mesonotum and anterior part of propodeum sloping gently downwards
posteriorly, the posterior half of the propodeal dorsum levelling off for a short distance before rounding
smoothly into the gently convex declivity. Metapleural lobes low and narrow, rounded and only slightly
projecting, but not at all concealed by the metapleural gland bulla in absolute profile. Petiole in profile with
ventral surface of peduncle shallowly sinuous but without a developed process. Dorsal surface of peduncle
also irregular and passing through a blunt angle about one-third the way from the base. Petiole node evenly
rounded and dome-like in profile; long and narrow in dorsal view, longer than broad and no broader than
the posterior peduncle, the sides of the node scarcely convex. Postpetiole in dorsal view broader than long.
Base of first gastral tergite no wider than the postpetiole in dorsal view but not forming a narrow neck;
instead the sides of the tergite diverge quickly and evenly from the base. Dorsum of head with sculpture
almost effaced, the surface between the eyes with faint narrow longitudinal costulae which are quite close-
packed and almost effaced in places. Occipitally the costulae present are even weaker than between the eyes
and are transverse. Ground-sculpture between the narrow costulae everywhere vestigial or absent, the
surface shining and mostly smooth. Dorsum of pronotum transversely arched-rugose, centrally with an area
of longitudinal rugosity; everywhere else the alitrunk transversely rugose. Sides of alitrunk more strongly
sculptured than dorsum, the rugae no denser but more sharply defined and more strongly developed.
Petiole, postpetiole and gaster unsculptured except for a faint superficial reticular pattern. All dorsal
surfaces of head and body with numerous standing hairs. Basal half of first gastral tergite with a number of
conspicuous long hairs which are as long as those on the dorsal alitrunk. Head very dark dull red, alitrunk
glossy jet black, remainder of body blackish brown but the pedicel segments with a reddish tint. Legs and
antennae dark dull red to reddish dark brown, approximately the same colour as the sides of the head.

Holotype worker, South West Africa: Tsisab Cyn., Brandberg Mts, 550 m, ll.v.1958 (E. S. Ross & R. E.
Leech) (CAS, San Francisco).

0. zekhem is one of the seven species known in this genus which lack a notch or impression in the
anterior clypeal margin. Of these seven species two, laticeps and cursor, have the base of the first

282 B. BOLTON

gastral tergite constricted and forming a narrow neck, which quickly separates them from
zekhem. Two other species, turneri and cavatodorsatus, are differentiated from zekhem by being
much smaller and having shorter scapes, as well as by their possession of specializations not seen
in zekhem. In turneri the eyes are large (0-28 x HW) and very conspicuously break the outline of
the sides of the head in full-face view; in cavatodorsatus the alitrunk outline is strongly saddle-
shaped. The only other known species which lack a clypeal notch are ankhu and velox, a close
species-pair. In both these species, however, the scapes are shorter than in zekhem and the first
gastral tergite lacks conspicuous long hairs on the basal half. Such pilosity is distinct in zekhem
where the hairs are as long as those on the dorsal alitrunk, whereas in both ankhu and velox hairs
are frequently absent from the first tergite, and when present they are very sparse, short and
inconspicuous.

PRISTOMYRMEX Mayr
(Figs 33-37)

Pristomyrmex Mayr, 1866: 903. Type-species: Pristomyrmex pungens Mayr, op. cit.: 904; by monotypy.

Odontomyrmex Andre, 1905: 207. Type-species: Odontomyrmex quadridentatus Andre, op. cit.: 208; by
monotypy. [Synonymy by Mann, 1919: 341.]

Hylidris Weber, 1941: 190. Type-species: Hylidris myersi Weber, loc. cit. (= Pristomyrmex africanus Kara-
vaiev); by original designation. [Synonymy by Brown, 1953: 9.]

Dodous Donisthorpe, 1946: 145. Type-species: Dodous trispinosus Donisthorpe, loc. cit.; by original de-
signation. [Synonymy by Brown, 1971 : 3.]

DIAGNOSIS OF WORKER. Distal portion of each mandible in anterior view suddenly broadened, much broa-
der than proximal portion. Apical (masticatory) margin usually with 4 teeth arranged as apical + preapical-
+ short diastema + 2 basals, of which the first basal is usually the smallest tooth. An offset basal denticle
may also be present as may a tooth about midway on the basal margin. In some species there is a tendency
for the two basal teeth to fuse, resulting in a broad basal tooth with two points. With wear these appear as a
single broad blunt tooth. Less commonly the mandible with 3 teeth only (apical + preapical + long dia-
stema + single basal), or with 5 teeth. Palp formula usually 1, 3 or 2, 3(1, 3 in africanus and orbiceps; 2, 3 in
trogor; 5 extralimital species also dissected) but higher in pungens-group with 4, 3 in cribrarius and 5, 3 in
pungens itself. Lateral portions of clypeus in front of antennal insertions reduced to a thin plate or ridge,
often translucent and usually projecting. Median portion of clypeus shield-like, broad between the antennal
insertions and frequently with a median carina, its anterior margin prominent and with a projecting apron
which is usually equipped with teeth or denticles but sometimes is merely crenulate. Median portion of
clypeus projecting over the mandibles when the latter are closed. Frontal lobes reduced or absent so that the
articulations of the antennae are mostly or wholly exposed and the roughly circular depressed areas
containing the antennal sockets are clearly visible. Frontal carinae variously developed, ranging from absent
to conspicuous, sometimes bounded below by a genal carina running longitudinally outside the depressed
area of the antennal insertions. Antennae with 1 1 segments, with a strong 3-jointed apical club. Eyes present
in all known species, situated approximately at the midlength of the sides of the head, very variable in size.
Alitrunk fusiform, without sutures dorsally. Pronotum often armed with spines, teeth or tubercles of varying
size but frequently only with minute prominences or unarmed. Propodeum with a pair of spines or teeth.
Propodeal spiracle circular. Metapleural lobes present and usually conspicuous. Orifice of metapleural
glands situated some distance up the pleuron, not in the posterior lower corner. Below the gland orifice is a
smooth and usually concave area running down towards the coxa, the concave area is bounded by a ridge
on each side and appears to be a specialized surface from which the products of the glands may evaporate
easily. Petiole nodiform, with a long anterior peduncle. Gaster commonly without hairs or only very
sparsely hairy. Sting long and very slender, the apical portion often very thin and hair-like.

Discounting synonyms and infraspecific forms the genus Pristomyrmex contains about 38 named
species to date. Of these names five valid species belong to the Ethiopian region (see below), six to
Australia (Taylor, 1965; 1968), and two to the Malagasy region (Brown, 1971). The remaining 26
names apply to the Oriental/Indo-Australian forms for which no synthesizing taxonomic study
has ever been undertaken. Without doubt a proportion of these names represent synonyms and
equally without doubt a fair number of new species from these regions await description, so 26
may in fact be a good indication of the actual number of species in the Oriental and Indo-
Australian regions, with a preponderance of forms in the latter.

SIX MINOR GENERA OF MYRMICINAE 283

The genus most closely related to Pristomyrmex is the monotypic Perissomyrmex Smith (1947:
281), which differs mainly in having the antennae with only 9 segments, as opposed to 11 in
Pristomyrmex. Perissomyrmex was based on a couple of specimens intercepted in quarantine in
the U.S.A., ostensibly on a ship from Guatemala. But, although Kempf (1972: 182) includes
Perissomyrmex in his Neotropic catalogue and that area has been tacitly accepted as the place of
origin, there is no hard evidence that Guatemala is in fact the original home of the genus. It so
resembles Pristomyrmex that an Old World origin must be suspected, and the recent discovery by
Cesare Baroni Urbani of a second Perissomyrmex species in Bhutan (as yet undescribed) adds
weight to the argument that South America is not the place of origin but that the specimens of the
type-species were brought to the U.S.A., via Guatemala, from somewhere in the Oriental region,
or possibly the Indo-Australian region, by human commerce.

Most Pristomyrmex species nest in rotten wood, either in fallen twigs in the litter layer or in
larger pieces of timber. Some nest in rotten parts of standing trees but most appear to prefer the
ground, foraging in the leaf litter and top soil. Of the five African species three, fossulatus,
cribrarius and trogor, seem fairly limited in distribution, with the first known from South Africa,
the second from South Africa and Mozambique, and the latter only from Zaire. P. orbiceps is
widely distributed throughout the wet forest zones of west and central Africa whilst the last
species, africanus, has an extremely wide range and seems able to inhabit woodlands and forests
virtually throughout sub-Saharan Africa.

Synonymic list of species

africanus Karavaiev

myersi Weber syn. n.

myersi subsp. mbomu Weber syn. n.

myersi subsp. primus Weber syn. n.

myersi subsp. beni Weber syn. n.
cribrarius Arnold
fossulatus (Forel)
orbiceps (Santschi)

laevigatus Weber syn. n.
trogor sp. n.

Key to species (workers)

1 Sides of petiole and postpetiole with coarse rugose sculpture. Sides of head behind eyes with

abundant short stout projecting straight hairs. (South Africa, Mozambique) . cribrarius (p. 285)
Sides of petiole and postpetiole smooth and shining. Sides of head behind eyes without projecting
straight hairs

2 Frontal carinae absent ; the backward projections of the frontal lobes terminating at the ends of

the antennal fossae, not reaching back even as far as the level of the anterior margins of the
eyes (Fig. 36). Antennal scapes relatively long, SI 100 or more. (Zaire) . . . trogor (p. 287)
Frontal carinae present ; the backward projections of the frontal lobes running back to or beyond
the level of the posterior margins of the eyes (Fig. 37). Antennal scapes relatively short, SI in
range 80-94

3 Eyes relatively small, maximum diameter only 0-12-0-15 x HW, with only 4-5 ommatidia in the

longest row. Pronotum armed with a pair of acute teeth or short broad spines. (Extremely

widespread in Africa) africanus (p. 284)

Eyes relatively large, maximum diameter 0-18-0-29 x HW. with 7 or more ommatidia in the
longest row. Pronotum only with a pair of low blunt tubercles, without acute teeth or short
spines

4 Head between frontal carinae to occiput smooth and shining, without foveolate punctures.

Maximum diameter of eye 0-18-0-21 x HW. Petiole node not wedge-shaped in profile. (Ivory
Coast, Ghana, Nigeria, Cameroun, Gabon, Zaire, Angola) . . orbiceps (p. 286)

Head between frontal carinae to occiput with distinct foveolate punctures. Maximum diameter of
eye 0-26-0-29 x HW. Petiole node in profile wedge-shaped, roughly triangular and tapering
dorsally. (South Africa) fossulatus (p. 285)

284 B. BOLTON

Treatment by species

Pristomyrmex africanus Karavaiev
(Fig. 34)

Pristomyrmex africanus Karavaiev, 1931: 47, fig. 5. Holotype worker, KENYA: Mabira, no. 5322 (Dogiel &

Sokolov) (ZM, Kiev) [examined].
Hylidris myersi Weber, 1941 : 190, figs 8, 9. Syntype workers, SUDAN: Equatoria, Aloma Plateau, Khor Aba,

3700 ft [1290 m], 10.viii.1939, no. 1470, 1474 (N. A. Weber) (BMNH; MCZ, Cambridge) [examined].

Syn. n.
Hylidris myersi subsp. mbomu Weber, 1952: 19. Holotype worker, CENTRAL AFRICAN REPUBLIC: Ubangi-

Shari, Bas Mbomu, 5 miles [8 km] W. of Bangassau, 12.iii.1948, no. 2210 (N. A. Weber) (AMNH, New

York) [examined]. Syn. n.
Hylidris myersi subsp. primus Weber, 1952: 19. Holotype worker, ZAIRE: Stanleyville [Kisangani],

19.iii.1948, no. 2235 (N. A. Weber) (AMNH, New York) [examined]. Syn. n.
Hylidris myersi subsp. beni Weber, 1952: 20. Syntype workers, ZAIRE: 15 miles [24 km] N. of Beni,

25.ii.1948, no. 2129 (N. A. Weber) (AMNH, New York; MCZ, Cambridge) [examined]. Syn. n.

WORKER. TL 2-7-3-4, HL 0-70-0-90, HW 0-68-0-92, CI 97-104, SL 0-62-0-86, SI 85-94, PW 0-46-0-60,
AL 0-68-0-92 (20 measured).

Base of mandible with 1-2 fairly strong rugulae present ; frequently these fade out distally but in a few they
may extend to the apical margin. Apical (masticatory) margin with a strong apical and preapical tooth
followed by a diastema and a broad basal tooth formed by the fusion of two basal denticles. Frequently this
broad basal tooth shows two points but these are often worn down and only a single blunt prominence
remains. Clypeus usually with a median longitudinal carina but this is variously reduced or lost in different
samples. Anterior clypeal margin with a median denticle and 1-2 pairs on each side of it, sometimes the
lateral pairs not strongly developed. Frontal carinae present, running back to or beyond the level of the
posterior margins of the eyes and strongly divergent in their anterior halves. Strongly developed scrobes
absent but sides of head between frontal carinae and eyes slightly concave and forming an unsculptured
scrobal area. Maximum diameter of eye 0-09-0-12, about 0-12-0-15 x HW and with 4-5 ommatidia in the
longest row. With the head in full-face view the occipital margin broadly and shallowly concave to conspicu-
ously indented medially, the sides convex. With the alitrunk in profile the pronotum armed with a pair of
acute triangular teeth or short spines; propodeum armed with a pair of spines which are somewhat variable
in length and thickness. Metapleural lobes prominent and rounded. Petiole and postpetiole in profile
rounded, without acute angles. Shape of petiole node variable but generally as in Fig. 34. Dorsum of head
between frontal carinae to occipital margin with foveolate punctures present at least from level of eyes
backwards. Both intensity of development and number of punctures very variable; at one extreme the
punctures are dense, sharply incised and conspicuous whilst at the other extreme the punctures are sparse,
shallow and feebly incised. Sides of head in front of, below, and behind eyes also with foveolate punctures,
and punctures usually also present at the posterior end of the scrobal area. Alitrunk unsculptured or at most
the dorsum with a few feeble rugular traces on the pronotum. Petiole, postpetiole and gaster unsculptured.
Mouthparts, ventral surface and dorsum of head with dense fine pilosity, on the dorsum the hairs arising
along the line of the frontal carinae longer than those arising between the carinae. Pronotal dorsum with a
transverse row of 3-4 pairs of hairs anteriorly; mesonotum with 3-4 pairs of hairs arising on the lateral
margins; propodeum hairless. Petiole, postpetiole and first gastral tergite without hairs but hairs present on
apex of gaster. Scapes and tibiae with pubescence, more conspicuous on the former than on the latter.
Colour varying from orange-brown to blackish brown, frequently with the gaster darker in shade than the
alitrunk and head.

Differentiation of this, the most widely distributed species of this genus, from orbiceps is tabulated
under the latter name. P. africanus separates easily from cribrarius as the latter is densely hairy,
strongly sculptured, has a palp formula of 4, 3 (as opposed to 1, 3 in africanus), and has much
larger eyes. P. trogor differs from africanus as the former lacks frontal carinae and has longer
scapes, a less densely hairy alitrunk and lacks foveolate punctures on the dorsum of the head. The
foveolate cephalic sculpture seen in africanus is, however, also present in fossulatus, but in this
species the eyes are much larger (0-26-0-29 x HW) and the pronotum has only a pair of blunt
tubercles, not sharp teeth such as are seen in africanus.

MATERIAL EXAMINED

Ghana: Kibi (D. Lesion); Mt Atewa (B. Bolton). Cameroun: Nkoemvon (D. Jackson). Gabon: Plateau

SIX MINOR GENERA OF MYRMICINAE 285

d'Ipassa (J. A. Barra). Kenya: 1°25'S, 35°10W [sic] to 1°38'S, 35°17'E (N. A. Weber). Zaire: Yangambi
(M. Maldague). Angola: R. Chicapa, Saurimo (Luna de Carvalho); Dundo (no name); Dundo (Luna de
Carvalho), R. Kahingo (Mwaoka); Salazar (P. M. Hammond).

Pristomyrmex cribrarius Arnold
(Fig. 35)

Pristomyrmex cribrarius Arnold, 1926: 281, fig. 81. Holotype female, MOZAMBIQUE: Amatongas Forest
(G. Arnold) (NM, Bulawayo) [examined].

WORKER. TL 3-2-3-4, HL 0-84-0-90, HW 0-86-0-92, CI 101-102, SL 0-66-0-72, SI 77-79, PW 0-56-0-60,
AL 0-82-0-86 (3 measured).

Basal portions of mandibles rugulose but this fading out distally so that the area near the apical margin is
mostly or entirely smooth. Apical (masticatory) margin of mandible with three teeth; apical and preapical
tooth acute and roughly the same size, behind them is a long diastema followed by the broad and truncated
basal tooth. The broad basal tooth may be the result of fusion of two denticles and specimens with a
broad-based tooth but a bidenticulate crown must be expected. Clypeus with a strong sharp median
longitudinal carina; anterior clypeal margin with a small truncated lobe medially where the carina meets the
margin, this lobe flanked on each side by a few smaller denticles which are variable in number and shape.
Palp formula 4, 3. Frontal carinae present and strongly divergent, reaching back to, or just beyond, the level
of the posterior margins of the eyes. Below the frontal carinae to eye level is a short, weakly impressed
scrobal area, bounded below by a weak genal carina which runs above the eye. Eyes large, maximum
diameter 0-18-0-20, about 0-21-0-22 x HW and with 9-10 ommatidia in the longest row. With head in
full-face view the occipital margin transverse to evenly very feebly concave, not indented medially. Prono-
tum armed with a pair of short but broad-based acute triangular teeth. Propodeum with a pair of long
spines which are slightly sinuate along their length. Metapleural lobes small, bluntly triangular in shape.
General shape of alitrunk and pedicel segments as in Fig. 35. Dorsum of head between frontal carinae and
back to occipital margin densely covered with broad, shallow foveolate punctures which are much broader
than the distances separating them. In places the foveolae are roughly aligned so that the cuticle separating
them appears as rugular sculpture. Similar sculpture is present below, behind and in front of the eyes, but the
scrobal area lacks such sculpture and the clypeus is unsculptured except for the strong median carina.
Entirety of dorsal alitrunk with strongly developed but blunt rugae which are predominantly longitudinal.
Sides of alitrunk rugulose. Sides and dorsum of petiole node with a few coarse longitudinal rugae. Sides of
postpetiole with a few strong rugae, the tergite in profile bounded by a raised ridge or rim. In dorsal view the
rim appearing as in Fig. 35, the space between the converging lines being smooth and shiny. Gaster
unsculptured. Dorsal surfaces of head, alitrunk, petiole and postpetiole with abundant short erect to
suberect hairs. Scapes and tibiae with similar freely projecting hairs and numerous hairs projecting from the
sides of the head in full-face view. First gastral tergite without hairs. Colour brown, the gaster darker than
the head and alitrunk but the appendages lighter.

This very distinctive species is apparently rare, being known only from the type-collection made
in Mozambique and a short series from South Africa noted below. It is easily separable from all
the other African Pristomyrmex by its dense pilosity, coarse sculpture, high palp formula and
oddly shaped postpetiole.

MATERIAL EXAMINED
South Africa: Natal, Dukuduku For. Res. (W. L. & D. E. Brown).

Pristomyrmex fossulatus (Forel)

Tetramorium (Xiphomyrmex) fossulatum Forel, 1910a: 428. Syntype workers, SOUTH AFRICA: Natal, Will

Broak [Willbrook] (Wroughton)(MHN, Geneva) [examined].
Pristomyrmex fossulatus (Forel) Santschi, 1916:51.

WORKERS. TL 2-7-3-2, HL 0-65-0-72, HW 0-62-0-70, CI 95-97, SL 0-54-0-60, SI 82-90, PW 0-46-0-52,
AL 0-68-0-70(3 measured).

Mandibles with vestiges of striate sculpture basally but mostly smooth with scattered small pits. Apical
(masticatory) margin of mandible with strongly developed apical and preapical teeth followed by a diastema
and a basal tooth which may be truncated. Clypeus with a sharp median longitudinal carina. Median

286 B. BOLTON

portion of clypeus with the anterior margin shallowly concave and armed with denticles; a median denticle
and 2-3 others on each side. Frontal carinae present and distinct, running back to the level of the posterior
margins of the eyes and forming the dorsal margins of the narrow, short antennal scrobes. Lower margin of
scrobe delimited by a longitudinal ruga above the eye, which runs back from the antennal fossa approxi-
mately to the midlength of the eye. Eyes large, maximum diameter c. 0-18, about 0-26-0-29 x HW, with
8-10 ommatidia in the longest row. With the head in full-face view the side convergent behind the eyes and
rounding into the occipital margin which is straight to very feebly and shallowly concave. With the alitrunk
in profile the pronotum only with a low, broad, blunt tubercle, without the teeth or spines frequently
encountered in this genus; in dorsal view the tubercles appear as low, bluntly rounded angles. Propodeum
with a pair of strong spines which are distinctly longer than their basal width. Metapleural lobes narrow and
strongly prominent. Petiole in profile wedge-shaped, strongly tapering dorsally and with the apex of the
triangular shape blunted. In dorsal view the petiole node broader than long. Subpetiolar process a long
narrow low flange. Dorsum of head between frontal carinae with scattered large shallow foveolate punc-
tures. Similar punctures also present on the sides of the head but generally less conspicuous. Dorsal alitrunk
also with foveolate punctures but here they are very sparse, widely separated, very shallow and inconspicu-
ous. Apart from these punctate areas the entirety of the head and body smooth and shining, with pedicel
segments and gaster completely unsculptured. Hairs present on mouthparts and gastral apex, otherwise the
dorsum only with 4-5 pairs on the head behind the level of the antennal insertions, following the line of the
frontal carinae; alitrunk with a single pair, on the mesonotum; petiole with 0-1 and postpetiole with 1-2
pairs dorsally; first gastral tergite hairless. Colour glossy light brown.

Known only from the type-series collected in South Africa, fossulatus is related to africanus
and orbiceps. It separates easily from the former as its eyes are much larger (0-26-0-29 x HW
as opposed to 0-12-0-15 x HW) and its alitrunk lacks the sharp pronotal teeth or broad spines
seen in africanus. The eye size of orbiceps is closer to that of fossulatus (but still smaller); orbiceps
lacks the foveolate cephalic sculpture of fossulatus and also lacks the very distinctively shaped
petiole node seen in the latter.

Pristomyrmex orbiceps (Santschi)
(Fig. 37)

Xiphomyrmex orbiceps Santschi, 19146: 367, fig. 30. Syntype workers, CAMEROUN: Victoria (Silvestri); and

GHANA: Aburi (Silvestri) (NM, Basle) [examined].
Pristomyrmex orbiceps (Santschi) Santschi, 1916:51.
Hylidris laevigatus Weber, 1952: 20, fig. 12. Holotype worker, ZAIRE: 13 miles [21 km] S. of Asa, lat. 4°40'N,

long. 25°40'E., 3.iii.l948, no. 2170.1 (N. A. Weber) (AMNH, New York) [examined]. Syn. n.

WORKER. TL 2-9-3-4, HL 0-72-0-90, HW 0-75-0-94, CI 100-106, SL 0-62-0-74, SI 80-87, PW 0-48-0-62,
AL 0-72-0-88 (25 measured).

Base of mandible usually with 1-2 rugulae present but these fading out distally so that the blade near the
apical margin is almost or entirely smooth. Apical (masticatory) margin with strong apical and preapical
tooth followed by a diastema and a broad basal tooth formed by the fusion of two basal denticles. In many
specimens this broad basal tooth shows two points but with wear only a single blunt prominence remains.
Clypeus frequently with a weak median longitudinal carina but this tends to be reduced, present only
posteriorly, or entirely absent. Anterior clypeal margin equipped with small denticles, usually 5 in number,
consisting of a median and 2 on each side. Degree of development of the denticles is variable, ranging from a
bluntly crenulate appearance to very distinct. Frontal carinae present, running back at least to the level of
the posterior margins of the eyes and strongly divergent in their anterior halves. Strongly developed scrobes
absent but the sides between the frontal carinae and the eyes sloping outwards and flat to very shallowly
concave. Maximum diameter of eye 0-14-0-19, about O-lS-Q-21 x HW and usually with 7-9 ommatidia in
the longest row, though in smaller individuals only 6 may be present. With the head in full-face view the
occipital margin shallow concave to quite conspicuously indented medially, the sides convex. With the
alitrunk in profile the pronotum armed with a low broad blunt tubercle, without the conspicuous acute teeth
or spines seen in other species; in dorsal view these tubercles appearing as low bluntly rounded angles.
Propodeum armed with a pair of spines which are usually longer than their basal width but which are
variable in length and, at minimum, may be only as long as their basal width. Metapleural lobes prominent
and rounded. Petiole and postpetiole in profile rounded, without acute angles. Dorsum of head between
frontal carinae to occipital margin smooth and highly polished. Sides of head mostly smooth but with some

SIX MINOR GENERA OF MYRMICINAE

287

scattered foveolate punctures in front of, below, and behind the eyes. These punctures are generally fairly
conspicuous but in some individuals they may be faint or even vestigial, especially behind the eyes. In most
specimens, but especially in larger ones, a few punctures occur on the side of the head on a line linking the
posterior margins of the eyes with the posterior extension of the frontal carinae, but these are frequently
vestigial or absent. Alitrunk, petiole, postpetiole and gaster unsculptured. Hairs very sparse, present on
mouthparts, ventral surface of head and gastral apex, but otherwise as follows. Dorsum of head behind level
of antennal insertions with 2-3 pairs of hairs along the line of the frontal carinae. Occipital corners with one
pair of hairs, Dorsal alitrunk without hairs except for a single pair on the mesonotum, and this pair is
frequently lost by abrasion. Petiole, postpetiole and first gastral tergite without hairs. Scapes and tibiae
lacking hairs but with fairly conspicuous pubescence which is usually more obvious on the former than on
the latter. Colour uniform blackish brown to black, the appendages lighter.

P. orbiceps is a widely distributed species, occurring throughout the wet forest zones of West and
Central Africa. Nests are constructed in fallen twigs or larger pieces of wood in the leaf litter layer
and workers forage singly in the litter. The separation of orbiceps from other African species is
straightforward. It differs from cribrarius as that species is heavily sculptured and densely hairy.
P. trogor lacks frontal carinae, has long scapes and relatively small eyes. The widely distributed
africanus differs from orbiceps as follows :

orbiceps
Pronotum with a pair of blunt tubercles.

Maximum eye diameter 0-18-0-21 x HW.

Eye with usually 7-9 ommatidia in the longest row

(sometimes with 6).
Dorsum of head behind antennal insertions with

2-3 pairs of hairs.

Dorsal alitrunk with 1 pair of hairs.
Head between frontal carinae smooth.

Scapes relatively somewhat shorter, SI range
80-87.

africanus
Pronotum with a pair of acute teeth or short

spines.

Maximum eye diameter 0-1 2-0- 15 x HW.
Eye with only 4-5 ommatidia in longest row.

Dorsum of head behind antennal insertions with
more than 5 pairs of hairs.

Dorsal alitrunk with 4 or more pairs of hairs.

Head between frontal carinae with foveolate punc-
tures, at least posteriorly.

Scapes relatively somewhat longer, SI range 85-94.

MATERIAL EXAMINED

Ivory Coast: Divo (L. Brader); Banco For. nr Abidjan (W. L. Brown); Orstom Exp. Sta. (W. L. Brown).
Ghana: Kibi(D. Lesion); Bunso(D. Lesion); Mampong(D. Leston); Mampong(P. Room) ; Tafo (B. Bolton);
Mt Atewa (B. Bolton). Nigeria : Gambari (B. Bolton); Gambari (B. Taylor). Cameroun : Nkoemvon (D. Jack-
son). Gabon: Plateau d'Ipassa (J. A. Barra); Makokou (W. H. Gotwald). Angola: Dundo (no name).

Pristomyrmex trogor sp. n.

(Figs 33, 36)

HOLOTYPE WORKER. TL 4-0, HL 0-96, HW 0-99, CI 103, SL 1-00, SI 101, PW 0-61, AL 0-96.

Mandibles basally with some weak rugular sculpture but this fading out distally so that near the apical
margin the blade is smooth. Apical (masticatory) margin with a large apical tooth followed by a slightly
smaller preapical, a diastema and two basal denticles which arise at each end of a raised welt representing
the fused bases of the two denticles (in worn specimens this would appear as a single broad truncated basal
tooth). Median portion of clypeus without a longitudinal median carina except posteriorly where a vestige
remains. Anterior clypeal margin with a small median tooth and a couple of smaller denticles on each side.
Frontal carinae absent, the posterior extensions of the frontal lobes strongly divergent but short, fading out
in front of the level of the anterior margins of the eyes. The genal carina which bounds the outer margin of
each antennal fossa strong, curving in towards the extensions of the frontal lobes but not meeting them.
Antennal scrobes absent, the scapes relatively long (SI, above). Eyes small, maximum diameter 0-15, about
0-15 x HW. With the head in full-face view the occipital margin indented medially, the sides shallowly but
evenly convex. Pronotum armed with a pair of short triangular spines, propodeum with a pair of slightly
larger spines; outline shape of alitrunk as in Fig. 33. Metapleural lobes fairly large, rounded. Dorsum of
alitrunk flat to shallowly concave between the pronotal spines and between the lateral hair-bearing welts of
the mesonotum. Petiole node high in profile, the dorsum sloping downwards posteriorly and rounding into
the posterior face. Anterior and dorsal faces of postpetiole in profile forming a single evenly curved surface.

288 B. BOLTON

In dorsal view the petiole node about as long as broad, the postpetiole very slightly longer than broad and
broadening from front to back. Entirety of head and body smooth and glossy, unsculptured except for a few
ridges on the metapleuron leading up to the orifice of the metapleural glands. Dorsum of head with
numerous fine curved hairs, some of which are very long. Mandibles, clypeal margin and ventral surface of
head with equally dense but generally shorter fine hairs; similar hairs also present on anterior coxa. Alitrunk
without hairs except for 2 pairs arising from the mesonotal welt. Petiole, postpetiole and first gastral tergite
without hairs; apex of gaster and sternites behind the first with a few hairs present. Scapes and tibiae with
short, fine, apically directed hairs. Colour uniform glossy chestnut-brown.

PARATYPE WORKERS. TL 3-4-4-0, HL 0-88-0-96, HW 0-89-0-98, CI 101-103, SL 0-90-0-98, SI 100-102,
PW 0-56-0-61, AL 0-86-0-98 (4 measured). Maximum diameter of eye 0-14-0-16, about 0-14-0-16 x HW.
As holotype but some darker brown in colour.

Holotype worker, Zaire (B. Congo on data label): S. Slope of Mt Kahuzi, 1900 m, 5.ix.l957 (£. S. Ross &
R. E. Leech) (CAS, San Francisco).

Paratypes, 19 workers and 1 male with same data as holotype (BMNH; MCZ, Cambridge; CAS, San
Francisco).

P. trogor is related to africanus, fossulatus, and orbiceps but is easily recognizable as in all of these
species frontal carinae are strongly developed whereas in trogor they are absent, compare Figs 36
and 37. Besides this the antennal scapes in trogor are relatively long, with SI 100 or more,
whereas the scapes are shorter in africanus, fossulatus and orbiceps with SI range 82-90. P.
africanus and fossulatus also differ from trogor by having broad foveolate punctures on the
dorsum of the head. P. orbiceps lacks the strong pronotal spines seen in trogor, having instead a
pair of low broad rounded tubercles.

TERATANER Emery
(Figs 38-55)

Terataner Emery, 1912: 103. Type-species: Atopomyrmexforeli Emery, 1900: 274; by original designation.
Tranetera Arnold, 1952: 130 [as subgenus of Terataner]. Type-species: Atopomyrmex bottegoi Emery, 1896:
155; by original designation. Syn. n.

DIAGNOSIS OF WORKER. Monomorphic arboreal myrmicine ants. Mandibles armed with 5 or 6 teeth. Palp
formula 5, 3 (alluaudi) or 4, 3 (bottegoi, elegans, luteus, piceus, scotti). Anterior clypeal margin with a median
notch or impression. Median portion of clypeus broad and broadly inserted between the frontal lobes,
bounded laterally by a pair of widely separated carinae which run to the anterior margin ; lateral portions of
clypeus unmodified. Frontal lobes narrow, continuing back into a pair of more or less straight frontal
carinae which are usually roughly parallel and relatively close together on the dorsum of the head. Towards
the occiput the frontal carinae either fade out or are sharply angled outwards as a ridge or row of tubercles
which runs to the sides of the head. Antennal scrobes absent or at most the sides of the head below the
frontal carinae with a broad and very shallow concavity. Antennae 12-segmented with a 3-segmented club,
the scapes when laid back failing to reach the occipital margin. Eyes large and conspicuous, situated at or in
front of the midlength of the head. Occipital corners tuberculate or denticulate in full-face view. Pronotum
marginate laterally and usually also anteriorly, the lateral marginations generally simple but sometimes
expanded into ornate lobes or flanges. Pronotal shoulders angulate, denticulate or tuberculate in dorsal
view. Promesonotal suture absent on the dorsum or represented by a line or slight indentation, only rarely
easily visible. Mesonotum usually marginate laterally and forming a low projecting angle or tubercle in
dorsal view; rarely immarginate and armed with a sharp denticle laterally. Metanotal groove impressed,
most frequently only shallowly so but deep in some species; very shallow indeed in some samples of elegans.
Propodeum bluntly marginate to rounded laterally, unarmed or with a pair of denticles or teeth. Meta-
pleural lobes large and strongly developed; ventral margin of metapleuron with a strong broad groove
running forward from the orifice of the metapleural glands. Ventral surface of alitrunk between hind coxae
entire, simple, without a broad deep circular pit. Middle and hind tibiae frequently with a distinct simple
spur, the spur reduced in some and indistinguishable from the hairs of the tibial apex in others. Petiole with
a short, stout anterior peduncle, the node narrow and tapering dorsally so that it appears triangular or
conical in profile. In anterior or posterior view the narrow dorsum of the node either forms a transverse crest
or is indented medially so that a pair of blunt prominences are formed laterally. In some these prominences
are acute and dentiform, in others developed into quite long teeth; rarely the petiole is strongly bispinose.

SIX MINOR GENERA OF MYRMICINAE 289

In one species (scott i) the petiole is developed into a very high plate dorsally which has a central emargina-
tion. Postpetiole simple or armed dorsally with a transverse crest or a single spine. Pilosity very variable,
some species densely hairy, others almost hairless. Sculpture generally of coarse rugae or sulci, but reduced
in the African species piceus, elegans, luteus and velatus.

Terataner is a small genus of arboreal ants containing 12 species, six of which occur in the
Ethiopian region and six in the Malagasy region. Nests are constructed in rotten parts of
standing timber, often some considerable distance above the ground. Of the African species four
occur quite widely in West and Central African forests (luteus, elegans, piceus, velatus), one is East
African (bottegoi) and the last (transvaalensis) is known only from South Africa. The first four
named form a complex of closely related species; the last two form a close species-pair which
shows marked similarity to the Malagasy species foreli, rufipes, steinheili and xaltus. Apart from
these Madagascar has another species, alluaudi, which is certainly the most bizarre representative
of the genus as it is presently understood. The final species of the Malagasy region, scotti, is
known only from a single worker from the Seychelles. A synopsis of the Malagasy species is given
at the end of this section.

Females (queens) are known for a few species and in general show the same characters as the
workers, except for the usual modifications associated with this caste. Males are very poorly
known, having been recorded only for elegans, scotti and foreli, the total number of specimens
amounting to six or seven.

Arnold (1952) proposed a subgenus of Terataner which he called Tranetera, erected to include
only the species bottegoi and transvaalensis, with the former nominated as type-species. In the
same paper he chose to treat Atopula Emery as a subgenus of Terataner. It has since been shown
(Bolton, 1976; 1980) that the type-species of Atopula, A. nodifera (Emery), is in fact a tetramoriine,
and the name Atopula has fallen as a straight synonym of Tetramorium Mayr.

Turning now to Tranetera, it seems probable that Arnold erected this name on the strength of
original descriptions alone, and did not see any material other than that of transvaalensis which,
however, he did recognize as being close to bottegoi. From his description of the subgenus only
three characters emerge to differentiate Tranetera from Terataner, namely that in Tranetera the
promesonotal suture was clearly defined, the metanotal groove (= meso-epinotal suture) was not
depressed, and that the petiole was quadrate and without spines. Opposed to this the six species
which he left in Terataner (alluaudi, foreli, luteus, rufipes, scotti and steinheili; piceus is not
mentioned and the description of elegans had not then appeared) were supposed to have the
promesonotal suture obsolete or slightly indented at the sides, the metanotal groove deeply
depressed and the petiole not quadrate, armed with two long spines.

Taking the last character, it is obvious when specimens are compared that an almost complete
morphocline is present. Only one species has the petiole transverse above, transvaalensis, and
even here a feeble indentation can be seen; only one species, alluaudi, has the petiole strongly
bispinose. Between these two extremes the petiole dorsally is indented to emarginate in bottegoi,
bilobate in scotti, shortly and bluntly bidentate in luteus, long bidentate in steinheili. The pres-
umed separation petiole not quadrate and with two long spines versus petiole quadrate and
without spines, does not exist in fact; all major steps between them being bridged in already
described species.

Similarly with the degree of definition of the promesonotal suture. It is quite clearly marked in
transvaalensis (though fused), weakly defined in bottegoi, vestigial in scotti where it is indicated
more by a change in sculpture pattern, almost invisible in steinheili where at certain angles the
strong sulcate sculpture is very feebly indented along the former track of the suture; obliterated
elsewhere. It should be noted that all species have an impression, notch or groove at each side of
the dorsum where pronotum and mesonotum meet, which indicates the ends of the former track
of the promesonotal suture, and which separates the marginations of the two segments.

Finally, the degree of impression of the metanotal groove varies considerably from species to
species. The variation does not allow a split such as that proposed by Arnold and a number of
species were incorrectly placed by him, as in bottegoi the metanotal groove is impressed where it
is as shallow in luteus as it is in transvaalensis.

290

B. BOLTON

Thus the concept of a subgenus Tranetera collapses and the name is relegated to the syn-
onymy. In point of fact Terataner, as presently constituted, forms a fairly compact genus, within
which the following species-complexes can be discerned.

luteus-complex (elegans, luteus, piceus, velatus). Frontal carinae more or less straight, fading out posterior-
ly, not angled outwards towards the sides of the head. Sculpture fine. Hairs very sparse and scattered or
absent on first gastral tergite. Dorsal (outer) surfaces of middle and hind tibiae without projecting hairs.
West and Central Africa.

/ore/i-complex (bottegoi, foreli, rufipes, scotti, steinheili, transvaalensis, xaltus). Frontal carinae angled
outwards posteriorly, running across the head either to the sides or towards the occipital corners as a
crest, ridge or row of tubercles. Sculpture coarse. Hairs dense and very conspicuous on first gastral
tergite. Dorsal (outer) surfaces of middle and hind tibiae with projecting hairs. East and South Africa,
Madagascar, Seychelles.

alluaudi-complex (alluaudi). As/ore/i-complex but frontal carinae feeble, sometimes almost indistinguish-
able from the cephalic sculpture, not running transversely on the head posteriorly. Postpetiole with a
single long median dorsal spine. Margins of pronotum expanded into a pair of broad laminae. Madaga-
scar.

Terataner belongs to a group of genera which also includes the African genus Atopomyrmex
Andre, the Oriental/Indo-Australian genus Dilobocondyla Santschi and the predominantly Aus-
tralian genera Dacryon Forel, Peronomyrmex Viehmeyer, Podomyrma F. Smith and Pseudo-
podomyrma Crawley (Taylor, 1970). Possibly also the strange monotypic genus Ireneopone Don-
isthorpe of Mauritius belongs to this assemblage. As can be seen, the only other African genus
noted is Atopomyrmex; the two are separated as follows in the worker.

Terataner
Monomorphic

Occipital corners tuberculate or denticulate.
Ventral surface of alitrunk simple and entire, with-
out a broad deep pit between the hind coxae.
Lower margin of metapleuron with a broad
groove running forward from the orifice of the
metapleural glands. (Figs 38, 39, 50-55).

Frontal carinae close together, long, usually ap-
proximately parallel, running back well beyond
the level of the eyes (Figs 42-45).

Propodeum unarmed to bidentate.

Atopomyrmex
Polymorphic

Occipital corners evenly rounded.

Ventral surface of alitrunk with a very conspicu-
ous broad deep pit between the hind coxae.

Lower margin of metapleuron without such a
groove, the margin rounded and folded in so
that the hind coxa appears to rest on the bulla of
the metapleural glands (Figs 3, 4).

Frontal carinae absent in smallest workers, length-
ening as size increases, running back beyond the
eyes only in large workers; when present always
very broadly separated and distinctly divergent
anteriorly (Figs 5-7).

Propodeum with a pair of long spines.

List of species

bottegoi (Emery)
elegans Bernard
luteus (Emery)
piceus Menozzi
transvaalensis Arnold
velatus sp. n.

Key to species (workers)

1 Dorsal (outer) surfaces of hind tibiae with projecting hairs. With the head in full-face view the
frontal carinae posteriorly sharply angled outwards, forming a ridge or row of tubercles which
runs towards the sides (Figs 42, 43) ..........

Dorsal (outer) surfaces of hind tibiae without projecting hairs. With the head in full-face view the
frontal carinae fading out posteriorly in a straight line, not sharply angled outwards
(Figs 44, 45)

SIX MINOR GENERA OF MYRMICINAE 291

2 Eyes more or less flat, not or only weakly breaking the outline of the sides of the head in full-face

view (Fig. 43). Sides of pronotum in dorsal view strongly convergent posteriorly (Fig. 47).
Black species; dorsal surfaces of alitrunk with abundant stout hairs. (Somali Republic, Kenya,

Tanzania) bottegoi (p. 291)

Eyes convex, conspicuously breaking the outline of the sides of the head in full-face view (Fig. 42).
Sides of pronotum in dorsal view roughly parallel, not strongly convergent posteriorly
(Fig. 46). Head and gaster black, alitrunk red; dorsal surfaces of alitrunk with sparse short
hairs. (South Africa) transvaalensis (p. 294)

3 With the postpetiole in dorsal view the posterior arch of the dorsum very broad and broadly

rounded (Fig. 41) 4

With the postpetiole in dorsal view the posterior arch of the dorsum narrow and narrowly
rounded (Fig. 40) • 5

4 Yellow. Pronotal dorsum distinctly broader than long (Fig. 49). Pronotal width almost equalling

to distinctly exceeding the combined lengths of pronotum and mesonotum in dorsal view.

(Ghana, Cameroun) luteus (p. 293)

Dark brown to black, the alitrunk usually reddish, lighter than the head and gaster. Pronotal
dorsum about as long as broad (Fig. 48). Pronotal width less than the combined lengths of
pronotum and mesonotum in dorsal view. (Guinea, Ghana, Nigeria) . . . elegans (p. 292)

5 First gastral tergite without hairs. Pronotal dorsum reticulate-punctate, without strong longi-

tudinal rugulae (Ghana, Nigeria, Cameroun, Equatorial Guinea) .... piceus (p. 293)
First gastral tergite with strong hairs on basal half. Pronotal dorsum reticulate-punctuate and
with strong overlying longitudinal rugulae. (Ivory Coast, Ghana) velatus (p. 295)

Treatment by species

The six species known from the Ethiopian zoogeographical region are discussed below in alpha-
betical order.

Terataner bottegoi (Emery)
(Figs 39, 43, 47)

Atopomyrmex bottegoi Emery, 1896: 155, fig. Holotype worker, SOMALI REPUBLIC: Lugh (V. Bottego)

(MCSN, Genoa) [examined].
Terataner bottegoi (Emery) Emery, 1912: 103.

WORKER. TL 4-8-6-0, HL 1-24-1-42, HW 1-12-1-34, CI 90-94, SL 0-76-0-90, SI 67-69, PW 0-90-1-12,
AL 1-46- 1-64 (5 measured).

Mandibles delicately longitudinally striate to almost smooth. Frontal carinae sharply developed as a pair
of raised ridges which are set quite close together on the dorsum and which are almost parallel, only very
slightly divergent and feebly sinuate along their lengths. Posteriorly each carina angles sharply outwards,
running towards the sides of the head as a weak ridge or row of linked tubercles. The frontal carinae and
their transverse posterior extensions form the borders of a weakly defined and shallowly impressed scrobal
area on each side. Eyes flat to extremely feebly convex, not prominent, maximum diameter 0-26-0-30, about
0-22-0-24 x HW. Pronotum in dorsal view marginate anteriorly and laterally, the lateral marginations
projecting and overhanging the sides. In dorsal view the pronotal corners sharply angulate to denticulate
and the sides distinctly convergent posteriorly. Sides of pronotum without a sharp strong margin running
between the anterodorsal and the anteroventral corners. Mesonotum marginate laterally; propodeum with
sides separated from dorsum by a blunt angle. Promesonotal suture visible at the sides of the dorsum but
usually absent centrally; however, in a few larger workers a faint track is visible arching across the dorsum.
Metanotal groove impressed, the propodeal dorsum sloping downwards posteriorly. Propodeum armed
with a pair of denticles, a pair of minute blunt tubercles, or unarmed. No two workers of the five seen have
the same propodeal armament. Metapleural lobes large and rounded. Petiole node roughly triangular in
profile, rising to a point dorsally. In anterior view the dorsum very shallowly impressed and the corners
rounded, not projecting as teeth. Dorsum of head strongly longitudinally rugose between the frontal carinae,
the rugae quite widely separated and the spaces between them filled with dense fine reticulate-punctulate
ground-sculpture. Sides of head between frontal carina and eye with similar dense ground-sculpture. In the
upper part of this scrobal area the punctulation is the only sculpture present, but in the lower half it is
overlaid by a series of rugae which became stronger closer to the eye; none of these rugae are as strong as
those on the dorsum. Sides of head below eye with regular strong, more or less parallel, longitudinal rugae.

292 B. BOLTON

Pronotal dorsum longitudinally rugose, the rugae becoming finer and less regular away from the midline but
present right up to the lateral marginations. Mesonotum similarly sculptured. Propodeal dorsum with a few
short rugae anteriorly but these fade out posteriorly leaving only the reticulate-punctate ground-sculpture.
Sides of alitrunk with regular more or less parallel longitudinal rugae. Petiole and postpetiole with
reticulate-punctulate ground-sculpture, the anterior face of the petiole node only with such sculpture but the
posterior face of the node and the dorsum of the postpetiole also with coarse rugose to almost sulcate
sculpture. First gastral tergite finely and superficially densely reticulate-punctulate, the base of the tergite
also finely longitudinally costulate. All dorsal surfaces of head and body with numerous conspicuous stout
hairs which are white to silvery in colour. Dorsal surfaces of middle and hind tibiae with standing hairs, but
such hairs absent from leading edges of antennal scapes. Colour black, the appendages brown.

Together with transvaalensis, bottegoi forms the African mainland representation of the predomi-
nantly Malagasy /ore//-complex of this genus. Characters useful in separating the two species are
discussed under transvaalensis.

MATERIAL EXAMINED

Kenya: Diani Beach (N. L. H. Krauss); Simu Beach, Kwale (£. S. Ross & R. E. Leech). Tanzania: Mafia I.
( Vesey-Fitzgerald).

Terataner elegans Bernard

(Figs 41, 45, 48)

Terataner elegans Bernard, 1952: 243. fig. 13 A. Holotype female, GUINEA: Keoulenta (Lamotte) (MNHN,
Paris) [examined].

WORKER. TL 4-9-5-6, HL 1-18-1-40, HW 1-12-1-30, CI 92-97, SL 0-72-0-82, SI 58-65, PW 0-78-0-88,
AL 1-50- 1-76 (20 measured).

Mandibles weakly longitudinally rugulose to almost smooth, frequently with distinctive large pits close to
the masticatory (apical) margin. Frontal carinae a pair of close-set straight, almost parallel ridges which fade
out posteriorly between the level of the posterior margins of the eyes and the occiput. Antennal scrobes
absent. Maximum diameter of eye 0-24-0-28, about 0-22-0-24 x HW. Pronotum in dorsal view marginate
anteriorly and laterally, with strongly angulate or short-denticulate anterior corners. On sides of the
pronotum margination runs from the anterodorsal to anteroventral angles. Mesonotum marginate laterally
and propodeum also with lateral margination though here it is much less sharply defined than on the
pronotum and in a few specimens is better described as bluntly angular. Lateral parts of promesonotal
suture visible on dorsum but centrally it is usually completely effaced. Metanotal groove feebly indicated in
profile by a shallow concavity or feeble indentation. Dorsum of pronotum about as long as broad, its width
obviously less than the combined lengths of the pronotum and mesonotum. Propodeum unarmed. Petiole
node low-triangular in profile, rising to a dorsal peak. In anterior view the dorsal margin of the node
strongly emarginate medially, the portions on each side of the emargination appearing as a pair of stout
tubercles or blunt teeth. Postpetiole in dorsal view flattened, in some the dorsum appearing shallowly
longitudinally concave; the posterior margin of the postpetiolar dorsum broad and broadly rounded.
Dorsum of head between frontal carinae with superficial but dense punctulate to granular ground-sculpture,
and with a few very weak longitudinal rugulae which are commonly broken or interrupted. Sides of head
above eyes with the same ground-sculpture and also with longitudinal rugulae which are usually somewhat
stronger than those on the dorsum. Dorsal alitrunk finely and densely reticulate-punctulate and with feeble
longitudinal rugulae, at least on the promesonotum. Postpetiole dorsally with a few stout conspicuous
rugae. First gastral tergite shagreened to finely superficially punctulate, with fine costulae on the basal
portion. Hairs very sparse on dorsal surfaces of head and body, present on mouthparts and gastral apex but
otherwise the maximum complement seeming to be 3-4 pairs on the head along the lines of the frontal
carinae, one pair on the pronotal corners, one pair on the propodeum, one pair on the posterior face of the
petiole node and 1-2 pairs each on the postpetiole and first gastral tergite. These hairs appear to be lost
easily by abrasion and completely hairless individuals are frequent. Scapes and tibiae without standing hairs
of any description. Colour usually with head and gaster blackish brown to black, the alitrunk lighter brown
or more usually reddish. Mandibles and clypeus usually lighter in colour than rest of head.

The four known species of the luteus-comp\e\ are confined to the forests of West and Central
Africa. The complex is diagnosed by the short straight frontal carinae which fade out on the head
behind the level of the eyes, fine sculpture, and sparse pilosity on the body; the tibiae lack

SIX MINOR GENERA OF MYRMICINAE

293

standing hairs. Of the species thus defined two, elegans and luteus, have a broad flattened
postpetiole, the posterior dorsal margin of which is broad and very broadly, evenly rounded. The
other two species, piceus and velatus, have the posterior arch of the postpetiole narrow and
narrowly rounded in dorsal view, compare Figs 40 and 41.

T. elegans and luteus are very closely related, being separated mainly on colour (luteus is
uniformly yellow), and on the dimensions of the pronotum as noted in the key. Apart from this
the pronotum of luteus in dorsal view usually has the lateral margins more strongly convex than
in elegans (Figs 48 and 49), though in smaller individuals this difference is often unapparent.
MATERIAL EXAMINED

Ghana: Tafo (C. Campbell); Tafo (B. Bollon); Kunso (D. Cross); Adeiso (D. Lesion); Bunso (D. Lesion);
Sajimasi (D. Lesion); Aburi (P. Room). Nigeria: Ife (B. Taylor).

Terataner luteus (Emery)
(Figs 38, 44, 49)

Alopomyrmex luteus Emery, 1899: 477. Syntype workers, CAMEROUN (Conradt) (MCSN, Genoa; MCZ,

Cambridge) [examined].
Terataner luteus (Emery) Emery, 1912: 103.

WORKER. TL 5-0-5-8, HL 1-22-1-40, HW 1-16-1-32, CI 94-98, SL 0-74-0-80, SI 59-66, PW 0-86-1-04
AL 1-52- 1-78 (15 measured).

Answering to the description of eleyans and very closely related to it. T. luteus differs from elegans as
follows.

luteus
Colour uniform dull yellow.

Pronotum broader than long, the pronotal width
about equalling or distinctly exceeding the com-
bined lengths of the pro- and mesonotum in
dorsal view.

At least in larger specimens the sides of the prono-
tum in dorsal view distinctly convex (Fig. 49).

Ground-sculpture on dorsum of head very feeble,
sometimes almost effaced.

MATERIAL EXAMINED

Ghana: Kade (D. Lesion); Bunso (D. Lesion).

elegans
Colour blackish brown to black on head and

gaster, alitrunk reddish or lighter brown.
Pronotum about as long as broad, the pronotal

width less than the combined lengths of the pro-

and mesonotum in dorsal view.

In all specimens the sides of the pronotum in
dorsal view only very feebly convex (Fig. 48).

Ground-sculpture on dorsum of head superficial
but dense and distinct.

Terataner piceus Menozzi
(Fig. 40)

Terataner piceus Menozzi, 1942: 173. Holotype worker, EQUATORIAL GUINEA: Rio Benito, 1939-40 (//.
Eidmann) (holotype not found in IE, Bologna; presumed lost).

WORKER. TL 4-6-5-5, HL 1-12-1-28, HW 1-06-1-20, CI 94-98, SL 0-64-0-70, SI 59-62, PW 0-75-0-84,
AL 1-38- 1-54 (9 measured).

Mandibles smooth with scattered large pits, those nearest the apical (masticatory) margin frequently
elongate. Frontal carinae straight, close together and approximately parallel, fading out between the level of
the posterior margins of the eyes and the occiput. Maximum diameter of eyes 0-26-0-30, about 0-25-
0-27 x HW. Pronotum marginate laterally and more weakly so anteriorly, the pronotal angles bluntly
denticulate and prominent. Sides of pronotum somewhat convergent posteriorly. Mesonotum marginate
laterally, the propodeal dorsum separated from the sides by a blunt angle, not nearly so sharply marginate
as the pronotum. Promesonotal suture absent from dorsum except laterally where it forms a break between
the pronotal and mesonotal marginations. With the alitrunk in profile the metanotal area shallowly im-
pressed. Propodeum unarmed or at most with a pair of minute low blunt tubercles where the sloping
dorsum meets the declivity proper. Metapleural lobes large and rounded. Petiole node in profile appearing
as a broad-based short triangular tooth, tapering to a point apically and slightly curved backwards. In
anterior view the dorsum of the petiole quite broadly and deeply impressed, the portion on each side of the

294 B. BOLTON

impression projecting as a blunted tooth which is directed slightly outwards. Postpetiole in dorsal view with
the arch of the posterodorsal surface narrow and narrowly rounded. Dorsum and sides of head above eyes
covered with fine reticulate-punctulate ground-sculpture which is usually weaker between the frontal ca-
rinae than elsewhere. Superimposed on this are a few weak fine longitudinal rugulae between the frontal
carinae and some stronger rugulae on the sides of the head above the eye. In this latter position cross-meshes
are frequently developed between the longitudinal components, more strongly so in front of the level of the
eye than behind it. Dorsal alitrunk finely and densely reticulate-punctulate, without trace of rugular sculp-
ture on the promesonotum. Pedicel segments and gaster similarly but more lightly sculptured. Basigastral
costulae very reduced, short and inconspicuous, restricted to the small area immediately behind the
postpetiolar-gastral junction. Hairs on dorsal surfaces of body very sparse, present on mouthparts and
gastral apex but otherwise the fullest complement seeming to be 3-4 pairs on head along the lines of the
frontal carinae, 1 pair each on mesonotum and propodeum (but none on pronotum in any specimens seen),
1-2 pairs on petiole behind node, 2-3 pairs on postpetiole, none on first gastral tergite. Outer surfaces of
middle and hind tibiae and antennal scapes without standing hairs. Colour uniform mid-brown to blackish
brown.

Of the luteus-complex of species two, piceus and velatus, are diagnosed by the shape of the
postpetiole which in dorsal view is narrow and narrowly rounded posteriorly, rather than being
broad and broadly evenly rounded posteriorly as is the case in elegans and luteus, compare Figs
40 and 41. T. piceus is separated from velatus by its lack of rugular sculpture on the promesono-
tum and lack of hairs on the first gastral tergite.

MATERIAL EXAMINED

Ghana: Sajimasi (D. Lesion); Aburi (D. Leston); Mampong (P. Room). Nigeria: Gambari (B. Bolton).
Cameroun : Nkoemvon (D. Jackson).

Terataner transvaalensis Arnold
(Figs 42, 46)

Terataner (Tranetera) transvaalensis Arnold, 1952: 130, fig. Holotype worker, SOUTH AFRICA: e. Transvaal,
Marieps (or Mariepskop), v.1951 (J. C. Faure) (NM, Bulawayo) [examined].

WORKER. TL 6-8, HL 1-60,HW 1-58.CI 98, SL 1-08, SI 68, PW 0-94, AL 2-02.

Mandibles longitudinally rugulose. Frontal carinae strongly developed as a pair of sharp raised ridges
which are almost parallel throughout their length, only very feebly sinuate. Posteriorly each frontal carina
angled sharply outwards and running towards the side of the head as a weak ridge. Antennal scrobes absent
but the head with a narrow indentation below the frontal carinae. Eyes convex and conspicuous, maximum
diameter 0-32, about 0-20 x HW. Pronotum in dorsal view marginate anteriorly and laterally, the anterior
corners prominently angulate; these angles also forming the origin of a sharp margin running down the sides
of the pronotum to the anteroventral angles. Sides of pronotum only very feebly sinuate in dorsal view,
roughly parallel, not strongly convergent posteriorly. Mesonotum marginate laterally, propodeum not
marginate. Promesonotal suture feeble medially on dorsum but its track visible; strongly defined laterally
on the dorsum where it separates the pronotal and mesonotal marginations. With alitrunk in profile the
metanotal region very shallowly concave only, not distinctly impressed. Propodeum rounded and unarmed,
without trace of teeth. Metapleural lobes conspicuous, large and rounded. Petiole node broadly triangular
in profile, rising to a peak dorsally. In anterior view the dorsum of the petiole indented medially, the dorsum
on each side of the indentation rounded, not developed into teeth. Dorsum of head with 11 sharp longitudi-
nal rugae between the frontal carinae at the level of the posterior margins of the eye. Sides of head between
frontal carinae and eyes more strongly rugose than dorsum and sides below eyes more strongly rugose still,
almost sulcate. Ground-sculpture on all surfaces of head a fine and very dense reticulate-punctulation.
Dorsal alitrunk sculptured with very broad low strong longitudinal rugae, almost sulcate. On the pronotum
these strong rugae are roughly parallel and are restricted to the central, transversely convex, portion and do
not occur on the flatter more lateral areas close to the lateral marginations. On the mesonotum the rugae
become weaker posteriorly and are distinctly divergent. On the propodeal dorsum the rugae are broader
and even less well defined. Sides of alitrunk bluntly rugose, on the pronotum the rugae diagonal and running
from the anteroventral to the posterodorsal angle. Elsewhere on the sides the rugae longitudinal, coarsest on
the propodeum above the spiracle. Ground-sculpture of entire alitrunk a fine and very dense reticulate-
punctulation. Anterior face of petiole node with dense punctulate ground-sculpture and a few rugular
vestiges but posterior face deeply sulcate, the sulci regular and parallel. Postpetiole sulcate in dorsal view,

SIX MINOR GENERA OF MYRMICINAE 295

the sculpture divergent posteriorly and overlaid by punctulate ground-sculpture. First gastral tergite dense-
ly reticulate-punctulate everywhere and with strong longitudinal costulae or rugae on the basal quarter. All
dorsal surfaces of head and body with short, quite stout hairs. These are quite numerous on head, pedicel
segments and gaster but are decidedly sparser on the alitrunk, being obvious only on the pronotum where
several pairs are present, but apparently represented by only a single pair each on the mesonotum and
propodeum. Dorsal surfaces of femora and tibiae and also leading edges of antennal scapes with scattered
stout standing hairs. Head and gaster blackish brown to black, alitrunk and appendages red.

T. transvaalensis, known only from the holotype worker, is one of the two African species
referable to the predominantly Malagasy /ore/i-complex. The other species in Africa is bottegoi.
The two are separated by the characters given in the key and by the fact that bottegoi tends to be
somewhat smaller than transvaalensis. Also, in transvaalensis the rugose sculpture of the pronotal
dorsum does not run to the lateral marginations, the lateral marginations of the pronotum do not
overhang the sides of the pronotum, the metanotal area is not sharply impressed and the side of
the head between the eye and frontal carina is more strongly rugose than the dorsum. In contrast
the rugose sculpture of the pronotal dorsum runs to the lateral marginations in bottegoi, the
lateral pronotal marginations overhang the sides, the metanotum is sharply impressed and the
side of the head between eye and frontal carina is less strongly rugose than the dorsum of the
head.

At present the ranges of the two are not known to overlap; transvaalensis is known only from
Transvaal, South Africa, whilst bottegoi has been recorded from Somali Republic, Kenya and
Tanzania.

Terataner velatus sp. n.

HOLOTYPE WORKER. TL 4-5, HL 1-14, HW 1-07, CI 94, SL 0-64, SI 60, PW 0-76, AL 1-40.

Mandibles smooth basally but in the apical halves with a number of large elongate pits which, where
aligned, give the appearance of short impressions. Frontal carinae fine, close together, parallel, more or less
straight, fading out on the dorsum between the level of the posterior margins of the eyes and the occiput.
Maximum diameter of eyes 0-27, about 0-25 x HW. Pronotum marginate laterally and more weakly so
anteriorly, the pronotal corners bluntly dentate and prominent. Sides of pronotum shallowly convex and
somewhat convergent posteriorly. Mesonotum marginate laterally, the propodeal dorsum separated from
the sides by a blunt angle, not at all as sharply marginate as the pronotum. Promesonotal suture absent on
dorsum except laterally where it forms a break in the margination. Metanotal groove only very shallowly
impressed in profile. Propodeum unarmed. Metapleural lobes large and rounded. Petiole node in profile
appearing as a triangular tooth, broad-based but short, tapering to an acute point apically and very slightly
curved backwards. In anterior view the dorsal surface of the node strongly emarginate in the middle, the
portions on each side of the emargination projecting slightly outwards as a pair of blunt teeth. Postpetiole in
dorsal view narrow and narrowly rounded posteriorly. Dorsum of head and sides of head above the eyes
with a fine dense reticulate-punctulate ground-sculpture which is fainter between the frontal carinae than
outside them. Overlying the ground-sculpture are a few fine longitudinal rugulae between the frontal carinae
and a series of somewhat stronger longitudinal rugulae running above the eye. Dorsal alitrunk finely and
densely reticulate-punctate everywhere. On the propodeal dorsum this is the only sculpture but the prono-
tum, and to a lesser extent the mesonotum, has fine but conspicuous fairly dense longitudinal rugulation.
Petiole, postpetiole and first gastral tergite finely and densely reticulate-punctate everywhere. Basigastral
costulae very reduced, the individual components short, fine and restricted to the area immediately behind
the postpetiolar-gastral junction. Standing hairs sparse on dorsal surfaces of the body, consisting only of 4
pairs on the head following the lines of the frontal carinae, one pair each on the pronotum (at the corners),
mesonotum and propodeum, 3 pairs each on the petiole and postpetiole, and 3 pairs on the first gastral
tergite situated in the basal half towards the sides of the sclerite. Other than this hairs are present only on the
mouthparts and gastral apex; standing hairs are absent on the scapes and tibiae. Colour uniform dark
brown.

PARATYPE WORKERS. TL 4-1-4-4, HL 1-04-MO, HW 0-98-1-06, CI 94-96, SL 0-60-0-64, SI 60-64,
PW 0-70-0-74, AL 1-24-1-36 (5 measured). Maximum diameter of eye 0-24-0-26, about 0-24-0-25 x HW.
As holotype but several paratypes with hairs variously lost by abrasion from the dorsal body surfaces.
Colour varying from mid-brown to blackish brown.

296 B. BOLTON

Holotype worker, Ghana: Okumaning, 12.viii.1969, ant ecology sample K12 (D. Lesion) (BMNH).

Paratypes. Ghana : 1 worker with same data as holotype but sample K 1 1 ; 1 worker, Kade, ant ecology
sample K96 (D. Lestori); 1 workers, Enchi, 17.V.1969, ant ecology sample E 5-8 (D. Leston). Ivory Coast: 1
worker, Banco Forest near Abidjan, 10.U963, no. A30(W. L. Brown). (BMNH; MCZ, Cambridge.)

T. velatus is closest related to piceus in the luteus-comp\e\, the two species sharing the character-
istic form of the postpetiole which in dorsal view is narrow and narrowly rounded posteriorly.
The two are easily separated as velatus has longitudinal rugular sculpture on the pronotum which
overlies the dense reticulate-punctulate ground-sculpture, whilst in piceus such rugulae are
absent. Besides this velatus has hairs present on the basal half of the first gastral tergite where in
piceus such hairs are lacking.

Synopsis of Malagasy region species

Terataner alluaudi (Emery)
(Fig. 50)

Atopomyrmex alluaudi Emery, 1895: 341, fig. 2. Syntype workers, MADAGASCAR: Diego-Suarez, iv-viii.1893

(C. Alluaud) (MCSN, Genoa; MCZ, Cambridge) [examined].
Terataner alluaudi (Emery) Emery, 1912: 103.

Terataner foreli (Emery)
(Fig. 55)

Atopomyrmex foreli Emery, 1900: 274, figs. Syntype workers, MADAGASCAR: Baia di Antongil, 1897-98

(A. Mocquerys) (MCSN, Genoa) [examined].
Terantaner foreli (Emery) Emery, 1912: 103.

Terantaner mfipes Emery
(Fig. 53)

Terataner rufipes Emery, 1912: 104. Holotype worker, MADAGASCAR: Fort Dauphin (M. Sikora) (MCSN,
Genoa) [examined].

Terataner scotti (Forel)
(Fig. 51)

Atopomyrmex scotti Forel, 1912: 160. Syntype worker and males, SEYCHELLES: Praslin I. (worker), and

Silhouette I. (males) (H. Scott) (BMNH) [examined].
Terataner scotti (Forel) Emery, 1912: 103.

Terataner steinheili (Forel)
(Fig. 54)

Atopomyrmex steinheili Forel, 1895: 485. Holotype worker, MADAGASCAR: 'Madagascar central' (M. Sikora)

(MHN, Geneva) [examined].
Terataner steinheili (Forel) Emery, 1912: 103.

Terataner xaltus sp. n.

(Fig. 52)

HOLOTYPE WORKER. TL 8-0, HL 1-82.HW 1-78.CI 98, SL 1-16, SI 65, PW 1-50,AL 2-22.

Mandibles with 6 teeth; with strong rounded longitudinal rugular sculpture. Median portion of clypeus
bounded by a carina on each side which runs to the anterior margin. Median clypeal carina present, running

SIX MINOR GENERA OF MYRMICINAE 297

to the clypeal notch. On each side of the median carina the clypeus with 2-3 weaker carinae which may be
broken or discontinuous. Eyes large, maximum diameter 0-40, about 0-22 x HW, situated approximately at
the midlengths of the sides and breaking the outline of the sides in full-face view. Frontal carinae sharply
developed, roughly parallel and about 0-76 apart (c, 0-43 x HW) at the level of the midlength of the eyes.
Posteriorly the frontal carinae sharply divergent and running towards the occipital corners as a raised crest
on each side. Lateral marginations of pronotum extended outwards as a semitranslucent lamella on each
side, the maximum width of which is c. 0- 14. On the left side of the holotype the lamella is continuous from
the pronotal corner but on the right the corner forms a separate angle, separated from the lamella proper by
an indentation. Pronotum (including the lamellae) much broader than long. Promesonotal suture absent on
dorsum centrally, but its former track indicated by a feeble indentation running across the unbroken strong
longitudinal sculpture. Lateral parts of promesonotal suture on the dorsal alitrunk, and its track down the
sides of the alitrunk, represented by a conspicuous cleft filled with very dense off-white pubescence. Mesono-
tum not marginate but armed with a low broad tubercle on each side. Promesonotum convex both longi-
tudinally and transversely. Metanotal groove impressed, narrow. Propodeum weakly and irregularly mar-
ginate laterally, armed with a pair of small denticles. Dorsum of propodeum rounding evenly into the short
declivity; the dorsum between the denticles transversely shallowly concave. Metapleural lobes conspicuous,
deep and broad. Peduncle of petiole thick, with a low and rounded anterior process ventrally. Petiole node
in profile low and roughly triangular, tapering dorsally to a narrow peak which is inclined backwards from
the vertical. In anterior view the dorsum of the petiole node emarginate and armed with a pair of low, broad
triangular teeth. Postpetiole in profile rising to a sharp peak dorsally. In dorsal view this peak is seen to be
the topmost point of a knife-edged transverse crest which falls away on each side, separating the anterior
face of the postpetiole from the rest. Dorsum of head between the frontal carinae with eight roughly parallel,
quite widely separated, strong longitudinal costae; the spaces between them with superficial granular
ground-sculpture. Sides of head between frontal carinae and eye rugose, with granular ground-sculpture
between the rugae and with a few cross-meshes above and in front of the eys. Sides of head below and
behind eye strongly sulcate. Sides of pronotum strongly sulcate, the sulci inclined slightly backwards from
the vertical. Remainder of sides of alitrunk longitudinally rugose except in the area below the propodeal
margin where the rugae are irregular. Promesonotal dorsum strongly and regularly longitudinally sulcate.
Propodeal dorsum with some longitudinal rugulae on the anterior half but behind this the surface merely
superficially reticulate. Anterior faces of both petiole and postpetiole with fine superficial reticulation, the
latter also with some low rugae which radiate from the petiolar junction. Posterior faces of both segments
with coarse rugae. Basal third of first gastral tergite densely longitudinally costulate, the spaces between the
costulae smooth or nearly so. Posteriorly, as the costulae fade out, a fine superficial reticulation or shagreen-
ing fades in, and this continues to the hind margin of the sclerite. All dorsal surfaces of head and body with
numerous short white hairs. Femora and tibiae with numerous standing hairs. Black, the mandibles,
antennae and legs dark brown.

Holotype worker, Madagascar: Foret de Zombitsy, near Sakaraha, 650 m, 16.xii.1959 (no collector's
name) (MCZ, Cambridge).

The shape of the postpetiole allies xaltus to rufipes, but this latter is a much smaller species
with simple pronotal margination and less regular pronotal sculpture. The following key will
separate the known species of the Malagasy region.

Key to species (workers)

1 Postpetiole armed dorsally with a single median curved spine (Fig. 50). Petiole with a pair of
long, usually back-curved, spines which are distinctly longer than the maximum diameter of the

eye. (Madagascar) • alluaudi

Postpetiole unarmed dorsally, without a median spine (Figs 51-55). Petiole with a pair of lobes or
short teeth which are distinctly shorter than the maximum diameter of the eye ....

2 Petiole node in profile produced into a very high thin plate-like prominence above (Fig. 51). In
anterior view the petiole broadly bilobate dorsally. (Seychelles) • scotti

Petiole node in profile roughly triangular in shape, tapering dorsally. In anterior view the petiole
emarginate to bidentate dorsally ............. 3

3 Postpetiole in profile rising to a sharp point or peak dorsally; in dorsal view this peak seen to be
the dorsalmost point of a sharp, knife-edged, transverse crest which slopes away on each side (Figs
52,53) . . 4

Postpetiole in profile bluntly rounded dorsally; in dorsal view without a knife-edged transverse
crest (Figs 54, 55) ............... 5

298 B. BOLTON

4 Lateral marginations of pronotum projecting into a broad semitranslucent lamina on each side

(Fig. 52). Larger species, HW > 1-50, PW > 1-30. (Madagascar) ....... xaltus

Lateral marginations of pronotum merely a sharp angle, not projecting into broad laminae
(Fig. 53). Smaller species, HW < 1-50, PW < 1-30. (Madagascar) rufipes

5 Posterior half of first gastral tergite densely reticulate. Lateral marginations of pronotum re-
duced, blunt and rounded, no sharper than the coarse sulci which make up the sculpture. Metano-
tal groove extremely deep, the anterior face of the propodeum concave so that the anterodorsal
angle of the propodeum overhangs the base of the metanotal groove (Fig. 55). (Madagascar) foreli

Posterior half of first gastral tergite smooth. Lateral marginations of pronotum sharp, very
distinctive and much sharper than the pronotal sculpture. Metanotal groove present but shallow,
unspecialized (Fig. 54). (Madagascar) steinheili

Acknowledgements

My profound thanks and gratitude for the loan of types and other important material necessary to complete
this study go to Dr Paul H. Arnaud (CAS, San Francisco); Dr Claude Besuchet (MHN, Geneva); Prof.
William L. Brown (Cornell University, Ithaca, New York); Mrs Cathy Car (NM, Bulawayo); Dr Jean
Decelle (MRAC, Tervuren); Mrs Marjorie Favreau (AMNH, New York); Dr Eberhard Konigsmann
(MNHU, Berlin); Dr Roberto Poggi (MCSN, Genoa); Dr A. G. Radchenko (ZM, Kiev); Dr David R. Smith
(USNM, Washington); Mrs Margaret K. Thayer (MCZ, Cambridge); Dr Cesare Baroni Urbani (NM,
Basle); Mme J. Casevitz Weulersse (MNHN, Paris).

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237, 3 figs.

- 1976. The ant tribe Tetramoriini. Constituent genera, review of smaller genera and revision of Tri-
glyphothrix Ford. Bull. Br. Mus. nat. Hist. (Ent.) 34: 281-379, 73 figs.

- 1980. The ant tribe Tetramoriini. The genus Tetramorium Mayr in the Ethiopian zoogeographical
region. Bull. Br. Mus. nat. Hist. (Ent.) 40: 193-384, 145 figs.

1981. Revision of the ant genera Meranoplus F. Smith, Dicroaspis Emery, and Calyptomyrmex Emery

in the Ethiopian zoogeographical region. Bull. Br. Mus. nat. Hist. (Ent.) 42: 43-81, 44 figs.
Brown, W. L. 1953. Characters and synonymies among the genera of ants, part 1. Breviora no. 11: 1-13.
- 1971. Characters and synonymies among the genera of ants, part 4. Some genera of subfamily Myrmi-
cinae. Breviora no. 365: 1-5.

1973. A comparison of the Hylean and Congo-West African rain forest ant faunas, pp. 161-185. In

Meggers, B. J., Ayensu, E. S., Duckworth, W. D., Tropical forest ecosystems in Africa and South America,
a review. Washington, D.C.

Donisthorpe, H. StJ. K. 1931. Descriptions of some new species of ants. Ann. Mag. nat. Hist. (10) 8: 494-501.
- 1933. Descriptions of three new species of Formicidae and a synonymical note. Ann. Mag. nat. Hist.
(10)11:194-198.

1946. A new genus and species of Formicidae from Mauritius. Proc. R. ent. Soc. Lond. (B) 15: 145-147,

4 figs.
Emery, C. 1886. Alcune formiche africane. Boll. Soc. ent. ital. 18: 355-366, pi. 17.

- 1891. Voyage de M. Ch. Alluaud dans le territoire d'Assinie (Afrique occidentale) en juillet et aout
1886. Annls Soc. ent. Fr. 60: 553-574, pi. 15.

SIX MINOR GENERA OF MYRMICINAE 299

- 1892. Sopra alcune formiche raccolte dall'Ingegnere L. Bricchetti Robecchi nel paese del Somali.
Annali Mus. civ. Stor. nat. Genova (2) 12: 1 10-122, figs.

- 1895. Mission scientifique de M. Ch. Alluaud dans le territoire de Diego-Suarez (Madagascar-Nord).
Annls Soc. ent. Belg. 39: 336-345, 3 figs.

- 1896. Formiche raccolte dal Cap. V. Bottego nella regione dei Somali. Annali Mus. civ. Stor. nat.
Genova (2) 17: 153-160, figs.

- 1899. Fourmis d'Afrique. Annls Soc. ent. Belg. 43: 459-504, figs.

- 1900. Formiche di Madagascar raccolte dal Sig. A. Mocquerys nei pressi della Baia di Antongil
(1897-1898). Boll. Soc. ent. ital. 31 : (1899): 263-290, figs.

- 1912. Etudes sur les Myrmicinae. Annls Soc. ent. Belg. 56: 94-105, 5 figs.

1922. In Wytsman, P. A. G., Genera Insect. Hym. fam. Formicidae subfam. Myrmicinae. fasc. 174c:

207-397.
Forel, A. 1895. Quelques fourmis du centre de Madagascar. Annls Soc. ent. Belg. 39: 485-488.

- 1901. Einige neue Ameisen aus Siidbrasilien, Java, Natal und Mossamedes. Mitt, schweiz. ent. Ges. 10:
297-310.

- 1910u. Note sur quelques fourmis d'Afrique. Annls Soc. ent. Belg. 54: 421-458.

- 19106. In Schultze, L. S., Zoologische und anthropologische Ergebnisse einer Forschungsreise im westli-
chen und zentralen Sudafrika. Formicidae: 1-30.

- 1911. Ameisen des Herrn Prof. v. Ihering aus Brasilien (Sao Paulo usw.) nebst einigen anderen aus
Sudamerika und Afrika. Dt. ent. Z 1911 : 285-312.

- 1912. The Percy Sladen expedition to the Indian Ocean in 1905, no. 11. Fourmis des Seychelles et des
Aldabras, recues de M Hugh Scott. Trans. Linn. Soc. Lond. 15: 159-167.

- 1913a. Formicides du Congo Beige recoltes par MM. Bequaert, Luja, etc. Revue Zoo/. Bot. afr. 2:
306-351.

19136. Fourmis de Rhodesia etc. recoltees par M. G. Arnold, le Dr H. Brauns et K. Fikendey. Annls

Soc.ent.Belg.51:m-\41,

- 1913c. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, ausgefuhrt im Auftrage
der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen, 2. Ameisen aus Su-
matra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr v. Buttel-Reepen in den Jahren
191 1-1912. Zoo/. Jb. Syst. 36: 1-148, figs.

Karavaiev, V. 1931. Ameisen aus Englisch-Ostafrika. Zoo/. Anz. 95: 42-51, 7 figs.
Kempf, W. W. 1972. Catalago abreviado das formigas da Regiao Neotropical. Studia ent. 15: 3-344.
Kugler, C. 1978. A comparative study of the myrmicine sting apparatus. Studia ent. 20: 413-548, 244 figs.
Mann, W. M. 1919. The ants of the British Solomon Islands. Bull. Mus. comp. Zoo/. Harv. 63 : 271-391, 59 figs.
Mayr, G. 1866. Diagnosen neuer und wenig gekannter Formiciden. Verh. zool.-bot. Ges. Wien 16: 885-908,
pi. 20.

- 1895. Afrikanische Formiciden. Annln naturh. Mus. Wien 10: 124-154, 3 figs.

Menozzi, C. 1942. Formiche dell'isola Fernando Po e del territorio del Rio Muni (Guinea Spagnola). Zoo/.

Anz. 140: 164- 182, 4 figs.

Prins, A. J. 1965. African Formicidae. Description of a new species. S. Afr. J. agric. Sci. 8: 1021-1024, 2 figs.
Santschi, F. 1910. Deux nouvelles fourmis du Tonkin. Naturaliste 32: 283-284.

- 1911. Nouvelles fourmis du Congo et du Benguela. Revue Zoo/. Bot. afr. 1 : 204-217.

- 1913. Melanges myrmecologiques. Annls Soc. ent. Belg. 57: 429-437.

- 1914a. Meddelanden fran Goteborgs Musei Zoologiska Afdeling no. 3. Fourmis du Natal et du
Zululand recoltees par le Dr I. Tragardh ; avec un appendice: Notes biologiques par I. Tragardh.
Goteborgs K. Vetensk.-o. VitterhSamh. Handl. 15: 1^7, 10 figs.

- 19146. Formicides de 1'Afrique occidentale et australe du voyage de M le Professeur F. Silvestri. Bull.
Lib. Zoo/, gen. agr. Portici 8: 307-385, 34 figs.

- 1916. Description d'un nouveau formicide de 1'Afrique occidentale. Bull. Soc. ent. Fr. 1916: 50-51, 1 fig.

- 1923. Descriptions de nouveaux formicides ethiopiens et notes diverses 1. Revue Zoo/. Bot. afr. 11:
259-295, 5 figs.

- 1924. Descriptions de nouveaux formicides africains et notes diverses 2. Revue Zoo/. Bot. afr. 12:

195-224, 10 figs.
- 1930. Formicides de 1'Angola. Revue suisse Zoo/. 37: 53-82, 10 figs.

1932. Formicides sud-africains. Soc. ent. Fr. Livre du centenaire, 1932: 381-392, 12 figs.

Smith, M. R. 1947. A new genus and species of ant from Guatemala. Jl N.Y. ent. Soc. 55: 281-284, 2 figs.
Stitz, H. 1923. Beitrdge zur Kenntnis der Land- und Siisswasser fauna Deutsch-Sudwestafrikas. Ergebnisse der

Hamburger deutsch-siidwestafrikanischen Studienreise 1911. Hymenoptera 7 Formicidae: 143-167, 2

figs. Hamburg.

300 B. BOLTON

Taylor, R. W. 1965. The Australian ants of the genus Pristomyrmex, with a case of apparent character
displacement. Psyche, Camb. 72: 35-54, 15 figs, 1 pi.

- 1968. A supplement to the revision of Australian Pristomyrmex species. J. Aust. ent. Soc. 7: 63-66,
2 figs.

1970. Characterization of the Australian endemic ant genus Peronomyrmex Viehmeyer. J. Aust. ent.

Soc. 9: 209-2 11, 3 figs.

Weber, N. A. 1941. Four new genera of Ethiopian and Neotropical Formicidae. Ann. ent. Soc. Am. 34:
183-194, 12 figs.

- 1943. Ants of the Imatong Mountains, Anglo-Egyptian Sudan. Bull. Mus. comp. Zool. Harv. 93:
261-389, 16 pis.

1952. Studies on African Myrmicinae, 1. Am. Mus. novit. no. 1548: 1-32, 36 figs.

Wheeler, W. M. 1919. The ants of Borneo. Bull. Mus. comp. Zool. Harv. 63: 43-147.

- 1922. Ants of the Belgian Congo, parts 1-8. Bull. Am. Mus. nat. Hist. 45: 1-1 139, 23 pis, 76 figs.
Wilson, E. O. 1976. Which are the most prevalent ant genera? Studia ent. 19: 187-200.

SIX MINOR GENERA OF MYRMICINAE

10

Figs 1-11 1, 2. Ankylomyrma worker. 1, profile of coronacantha, 2, head of coronacantha. 3-11. Ato-
pomyrmex workers. 3, profile of large worker of mocquerysi, 4, profile of small worker of mocquerysi.
5-7, heads of mocquerysi workers: (5) large (HW 2-20), (6) medium (HW 1-90), (7) small (HW 1-30). 8,
9, propodeal profile of (8) mocquerysi, (9) cryptoceroides. 10, 1 1, propodeal dorsum of (10) mocquerysi,
(11) cryptoceroides. Sculpture omitted.

302

B. BOLTON

14

15

Figs 12-17 12-14. Baracidris workers. 12, profile of meketra. 13, 14 heads of (13) meketra, (14) sitra.
15-17 Cyphoidris workers. 15, profile of spinosa. 16, head of spinosa. 17, alitrunk and pedicel of
exalta. Sculpture omitted.

SIX MINOR GENERA OF MYRMICINAE

Figs 18-25 Ocymyrmex workers. 18, profile of nitidulus. 19-21, alitrunk and pedicel segments of (19)
celer, (20) sobek, (21) sphinx. 22-25, head of (22) nitidulus worker, (23) nitidulus female, (24) robecchii,
(25) turneri. Sculpture omitted in 24, 25.

304

B. BOLTON

Figs 26-32 Ocymyrmex workers. 26-30, pedicel segments of (26) fortior, (27) phraxus, (28) flaviventris,
(29) laticeps, (30) weitzeckeri. 31, 32, alitrunk and pedicel segments of (31) shushan, (32) velox. Sculp-
ture omitted.

SIX MINOR GENERA OF MYRMICINAE

305

36

35

40

Figs 33-41 33-37. Pristomyrmex workers. 33, profile oftrogor. 34, 35, alitrunk and pedicel segments of
(34) africanus, (35) cribrarius. 36, 37, heads of (36) trogor, (37) orbiceps. 38-41. Terataner workers. 38,
39, profiles of (38) luteus, (39) bottegoi. 40, 41, pedicel in dorsal view of (40) piceus, (41) elegans.
Sculpture omitted.

B. BOLTON

Figs 42-55 Terataner workers. 42-45, heads of (42) transvaalensis, (43) bottegoi, (44) luteus, (45) elegans.
46-49, dorsal alitrunk of (46) transvaalensis, (47) bottegoi, (48) elegans, (49) luteus. 50-55, alitrunk and
pedicel of (50) alluaudi, (51) scotti, (52) xaltus, (53) ruftpes, (54) steinheili, (55)foreli. Sculpture omitted.

SIX MINOR GENERA OF MYRMICINAE

307

Index

Synonyms are in italics.

abdominalis 269
africanus 284
alluaudi 296
ankhu 265
Ankylomyrma 247
arnoldi (Atopomyrmex) 251
arnoldi (Ocymyrmex) 269
Atopomyrmex 249
australis 251

Baracidris 252
barbatus 265
barbiger 265
beni 284
bottegoi 291

carpenteri 275
cavatodorsatus 266
celer 267

coronacantha 248
cribrarius 285
cryptoceroides 250
cursor 267
curvispina 250
Cyphoidris 256

deplanatus 250
Dodous 282

elegans 292
erigens 251
exalta 258

flavescens 265
flaviventris 268
foreli (Ocymyrmex) 269
foreli (Terataner) 296
fortior 269
fossulatus 285

hirsutus 271
Hylidris 282

laevigatus 286
laticeps 271
luteus 293

mbomu 284
meketra 255
melanoticus 250
micans 272

mocquerysi 251
monardi 273
myersi 284

nigellus 251
nitidulus 274

obscura 251
Ocymyrmex 260
Odontomyrmex 282
opaca 251
orbiceps 286

parissa 258
phraxus 274
picardi 275
piceus 293
primus 284
Pristomyrmex 282

robecchii 276
robustior 265
rufipes 296

scotti 296
shushan 277
sitra 256
sobek 277
sphinx 278
spinosa 257
steinheili 296

Terataner 288
Tranetera 288
transvaalensis 294
transversus 269
trogor 287
turned 279

usakosensis 269

velatus 295
velox 280

weitzeckeri 280
werneri 259
wroughtoni 280

xaltus 296
zekhem 281

British Museum (Natural History)
1881-1981

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