ali Beal stot a aL
rly easel ay
eat ae
Ny
eels
Bait : i hae eit eye fing tne tie
ARUSHA TAs Cala Win rC IO ae Ma e nyt
NURI SRA aa GaN a oD
‘ RSE NRARN Apter aie se PEI Hi tip
Ati pave 4 ee ;
Tees ‘4 tie
nv
ast
.
eS
5
. ae
Sra
ab eats
Be
5
yiis
ol
La ; if at bas
at ICRC AT RI re :
Higa eaneeenctt nas Pane an ‘
piel : 2 ;
ite boa Nar eee s Sey atye ser
PAC TaeE Une aie ec Heca tran del var eae Girt nt toet
Trsestys rr ; Mate Sin es
SS nA ep a OSs MER a
Man Tt wy) ? ane + bose
Reta i iy Hie
mits
eel
on
See
aig
Pye
Hay hee 2
iNet s
#
K
rs
'S,
ae
we eae
re
3a:
oy
Y Phe
4 vsbaiey’
ceo i Re
pa)
tis
3 ?
sh iy
Ueaat?
“sik Ps
4 $
ye eH
Wishes?
Tea
ats
ca
re)
iseat
t? fy ye
Lk . t tee 3 y. f
PONSA fest stele ty panes
De ey ce
Breen ee piesa Wale
a > Batay aa é ;
Fit) Fst tp ( i on ie
+
ths
sect
Fee ety
ie
te REE t
Ave
ie
i.
*y
Raa
ya yt
es
Sis
eee
ee
rt oe id i oe
Pere ars
i
rerieltatl
dy relyt gen ap lis
Wis tik
* ‘S
‘4 5 i
cay ; 1, Hosta:
ae ; ery per
Hep tiertutete 4 in a
Msuoieae raat bss Wisp seeain gaye tia ey
Biot,
esos
ae ee ae em
Carty
crs
3
=
Z
2 ~
ote
eae ae
fe
; i Hi
Ak:
we as),
re
ay
is
ee
f Sie
fe
=
ay
Moe
aoe
nary
Tepe atee
10
ty ts
8
Breath
ie bee as
SRE
: 2,
eaten ts
A
eee
4
Me,
Hi Sistine
ue Nis a Bena He Bastia
radial Pat aay Latest
Resi ere Nonsetatnt}
eae
ope
ae
Se
eee
Votes
ee
cacé
es
=
‘ae
Heinys
* +3
Hiatal
EsCh pat take
i cst ieee ns) at)
isteale; Ht vircded say
Ahh ? Y
BP Ri
ee
isan
ie
tis
He, Nivel
Lea
eee
=
ap a
i aiid
Ao ‘
ine is it
Beasties
¥, i,
Ts nat
y x
2. BSP HPer stacy a ga es? Sake
iin ioe Saami
7 ‘ . . 0 .
ee ae) \ ee) i
REE tN 5 soi ih iM
4 7 4, ~ ; < .
SNORT AB . (eaten NS
al i tet *
bi
5
Nh
ter
oe ey
+7
RMON!
Rasterirsett
Palmns
a
b On
SPAR ciara sisstecraas
ote tet ek F iy E nes
Seri
ir,
tee
tess]
we
a, Rem :
ead be
i -.
‘
4
fe
we
ti
* ' a
“~ 7 ~
‘ bas
hy of
- -
tse
Osa. >
on te
a
or
J
, +
’ of
: int,
zs .
‘
a .
\
- an ‘ r
oT Oe &
mh ioe! eee
.
: ha ; 3
"ste a .
‘
i
‘
,
i
:
‘
i
¥
= ~
-
~
}
‘
1
D f
(es
=
ees
ro i
Pe
- 7 o
,
7
'
4
=
-
1
I
a a
0. aL. .
= a
1
cP ee
We ee
‘
i“ ear!
e * pa
fat
a
3
a.
-
a eee
ae hoy
as" =
‘ ’
£
:
30 Feo 2
, q “w St
i”
‘
a
¥
'
‘
a T
‘
By
‘a
‘a —
iW
i
i
—. ae ee eo ei ese Uf
a eee ~ T ral
vv? ww
Bulletin of the
British Museum (Natural History)
A revision of the African pod bugs of the
tribe Clavigrallini (Hemiptera: Coreidae)
with a checklist of the world species
W. R. Dolling
Entomology series Vol39 Nol 26 July 1979
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and
an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum’s resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and are not
necessarily completed within one calendar year. Subscriptions may be placed for one or more
series. Subscriptions vary according to the contents of the Volume and are based on a forecast
list of titles. As each Volume nears completion, subscribers are informed of the cost of the
next Volume and invited to renew their subscriptions. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW7 5BD,
England.
World List abbreviation: Bull. Br. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1979
ISSN 0524-6431 | Entomology series
Vol 39 No | pp 1-84
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 26 July 1979
A revision of the African pod bugs of the tribe
Clavigrallini (Hemiptera: Coreidae) with a checklist of
the world species
W. R. Dolling ©
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis . 1
Introduction 1
Abbreviations of depositories 2
Terminology and measurements . 2
Systematic section . 3
Key to the genera of Clavigrallini 3
Oncaspidia Stal , , 3
Description of species . 3
Clavigralla Spinola . 6
Key to groups and subgroups 8
Key to African and Malagasy species . : y , . 2 ‘ - ws
Descriptions of species : . ; : : , : : ; mae
The tuberculicollis-group . : ; : . ; ’ , : ty Wate
The elongata-group . ° . 7 : ; ; : y : : i: to
The tomentosicollis-group . : A ; ; ' ; : ; . 48
The Jeontjevi-subgroup . : ” . ; , : : , . 48
The ruandana-subgroup . : : z , 3 : : ; Oe
The spiniscutis-subgroup : : ; F ; : : 1. 354
The tomentosicollis-subgroup . , ; : ; : : = 3163
The wittei-subgroup__.. j , : : ; . 74
Nomenclature of economically important species : : : ; ; . 80
Check-list of the species of the tribe Clavigrallini . ; : : 4 : ae)
Acknowledgements. , : : : : : : ; ; ; Pane} |
References : : : : ; , ; ; : ; : : te
Index : ; ; ‘ : : , : : : : ; £1483
Synopsis
All known African and Malagasy species of the tribe Clavigrallini are described and a key is provided for
their separation. Two genera and 44 species are recognized in the area covered. Two generic and two
specific synonymies are newly established, one species is raised from synonymy and three specific synonym-
ies are confirmed. Two subordinate taxa are raised to the rank of species. Twenty-three species are
described as new. Twelve new combinations are established, five combinations are revived and one com-
bination is used in its correct sense for the first time. A type-species is designated for one generic name and
lectotypes are designated for 18 names in the species-group. Biological information derived from published
works and from data labels is summarized for each species. The implications of this revision for the
nomenclature of the more important pest species are summarized separately.
Introduction
The general morphology and systematic position of the tribe Clavigrallini, which belongs to the
subfamily Pseudophloeinae of the family Coreidae, are discussed by Dolling (1978). That paper
and the present work together constitute a complete revision of the tribe.
Bull. Br. Mus. nat. Hist. (Ent.) 39 (1) : 1-84 Issued 26 July 1979
2 W. R. DOLLING
The tribe is represented throughout the Ethiopian and Oriental regions, extending into tropical
Australia and the southern fringes of the Palaearctic region. Of the four genera, two are exclusively
Oriental, one (Oncaspidia Stal, with a single species) is African and one (Clavigralla Spinola)
occurs throughout Africa, Madagascar and associated islands and has a few species belonging to
one of its more highly evolved sections in the Oriental region.
The major recorded host plants are herbaceous Leguminosae (Fabaceae), including many
cultivated pulses. A rapidly growing body of literature exists concerning the bionomics, pest
status and control of several species regarded as pests. In the English language these pests are
called spiny brown bugs or pod bugs (Afrikaans: Ysterbek).
Clavigrallini may be recognized by their small size (length usually less than 11 mm), generally
spiny outline and the characteristically spined, clavate posterior femora (Fig. 185). Where more
spines are present on the posterior femur (extreme case in Fig. 124), Clavigrallini may be dis-
tinguished from other Pseudophloeinae of similar build by the absence of a tubercle from the base
of the posterior femur on its posterior face close to the apex of the trochanter, a feature which is
readily visible if present. Risbecocoris Izzard lacks this tubercle but is unlikely to be confused
with the Clavigrallini. It has several long, laterally directed spines arising from the lateral margins
of the pronotum.
Abbreviations of depositories
The specimens studied in the course of this revision are deposited in the various institutions whose
names are abbreviated in the text as follows: Museum fiir Naturkunde der Humboldt-Universitat,
Berlin, D.D.R. (MNHU, Berlin); Institut Royal des Sciences Naturelles de Belgique, Brussels,
Belgium (IRSNB, Brussels); National Museum, Bulawayo, Rhodesia (NM, Bulawayo); Zoo-
logical Museum of the University, Helsinki, Finland (ZMU, Helsinki); University of Ife, Ie-Ife,
Nigeria (University of Ife); University of Ghana, Legon, Ghana (UG, Legon); the collection of
Dr P. Duarte Rodrigues, at present at Museu e Laboratorio Zooldgico e Antropolégico (Museu
Bocage), Lisbon, Portugal (Duarte coll.); British Museum (Natural History), London, United
Kingdom (BMNH, London); American Museum of Natural History, New York, U.S.A.
(AMNH, New York); University Museum, Oxford, United Kingdom (UM, Oxford), Muséum
National d’Histoire Naturelle, Paris, France (MNHN, Paris); Plant Protection Research Insti-
tute, Pretoria, Republic of South Africa (PPRI, Pretoria); Transvaal Museum, Pretoria, Republic
of South Africa (TM, Pretoria); Institute of Agricultural Research, Samaru, Nigeria (IAR,
Samaru); California Academy of Sciences, San Francisco, California, U.S.A. (CAS, San
Francisco); Naturhistoriska Riksmuseum, Stockholm, Sweden (NR, Stockholm); the private
collection of Professor J. A. Slater, at present at University of Connecticut, Storrs, Conn., U.S.A.
(Slater coll.); Musée Royal de l’Afrique Centrale, Tervuren, Belgium (MRAC, Tervuren); Uni-
versity of Uppsala, Uppsala, Sweden (University of Uppsala); Naturhistorisches Museum,
Vienna, Austria (NM, Vienna).
Terminology and measurements
The segments of the antennae and rostrum are numbered I-IV starting with the segment nearest
the body; lengths exclude basal parts of segments capable of retraction into the preceding arti-
culation and also exclude the antennal ring segment between III and IV. The origin of the first
rostral segment is taken as the base of the labrum rather than the base of the first labial segment,
which is usually concealed, whence the use of the term ‘rostrum’ (‘rostral’) in preference to
‘labium’ (‘labial’). The angles at the point of greatest width of the pronotum are termed the
posterolateral angles and the spines they bear the posterolateral spines. In the presentation of
linear data, ratios are preferred to direct scalar measurements. Where practicable, means, ranges
and the number of specimens measured are given, the number of specimens, n, being indicated
by the formula ‘(n = x)’ following the mean. For a discussion of the rationale of these methods
of presenting data see Dolling (1978 : 283).
AFRICAN POD BUGS 3
Systematic section
Key to the genera of Clavigrallini
1 Side of head above base of antennifer bearing an erect spine (Fig. 1). Pronotum without large
spines or tubercles on disc or near lateral margins. (Throughout Ethiopian region except
Mascarenes) . : : : F . ONCASPIDIA Stal (p. os
— Side of head without spine (Figs 9, 51, 137)
2 Pronotum without large tubercles sublaterally or on disc; female abdomen with apex strongly
deflexed; pubescence of pronotum uniform. (Oriental region eastwards to Solomon Islands
and northern tropical Australia) . ; ; . GRALLICLAVA Dolling (1978 : 304)
— Pronotum with large tubercles on disc or sublaterally or, if without tubercles, pubescence of
anterior two-thirds of pronotum strikingly different in colour and texture from that of
posterior one-third . 3
3 Pronotum with a group of four large tubercles on disc. (Oriental region eastwards to New
Guinea and northern tropical Australia) ; . CLAVIGRALLOIDES Dolling (1978 : 293)
— Pronotum with a pair of large tubercles sublaterally behind level of calli (Figs 138, 157, 188,
etc.) or a semicircle of tubercles on disc (Figs 10, 50, 98, 126, etc.), often with additional
tubercles but never with a group of four, rarely without tubercles (Fig. 131). (Africa and
Mascarenes with a few species in Oriental region eastwards to China and Java)
CLAVIGRALLA Spinola (p. 6)
ONCASPIDIA Stal
Oncaspidia Stal, 1873 : 81, 83. Type-species: Clavigralla pilosicollis Stal, by monotypy.
Length of head about two-thirds length of pronotum. Side of head in front of each eye just above base of
antennifer with an erect spine. Antennifer with outer apical process porrect. Antenna with segment I
longest, III shortest, IV shorter than II and with specialized sensory area occupying about nine-tenths
of its length. Rostrum at rest with apex reaching to disc of metasternum, segment IV slightly longer and
II slightly shorter than I, III about two-thirds as long as I. Bucculae small, occupying about one-third of
length of ventral midline of head.
Pronotum strongly declivent, posterolateral angles produced, disc without large tubercles or spines,
prescutellar spines present, small. Scutellum convex with a pair of small knobs at base. Mesosternum and
metasternum broadly and shallowly longitudinally sulcate along midline. Dorsal ridge of metathoracic
scent-gland peritreme entire, not bilobed. Anterior and intermediate femora each with 0-1 subapical
spine beneath, posterior femur with two major spines and a terminal series, without basal tubercle.
Abdominal sterna III-VII with posterolateral angles prominent. Male genital capsule closed posteri-
orly, parameres obliquely capitate. Female with valvulae of ovipositor very long in comparison with other
Coreidae.
REMARKS. A single species, similar in build to C. tomentosicollis, readily distinguished by the spines
in front of the eyes. The ovipositor is remarkable for its length, even in comparison with other
Pseudophloeinae, in which the structure of the ovipositor tends more towards the ‘laciniate’ type
than that of other Coreidae.
DISTRIBUTION. Tropical and southern Africa.
INCLUDED SPECIES. A single known species, O. pilosicollis (Stal).
Description of species
Oncaspidia pilosicollis (Stal)
(Figs 1-8)
Clavigralla pilosicollis Stal, 1855 : 31. LECTOTYPE 2, Soutu Arrica (NR, Stockholm), here designated
[examined].
Clavigralla similis Signoret, 1860 :944. LECTOTYPE 3, ZANZIBAR (NM, Vienna), here designated
[examined]. [Synonymized by Stal, 1866 : 107.]
Oncaspidia pilosicollis (Stal) Stal, 1873 : 83.
Length: 3, 7:3-9:1 mm, mean 8-2 mm (n = 11); 9, 8:2-9:-4 mm, mean 9-0 mm (n = 11).
Form robust, deep-bodied.
4 W. R. DOLLING
C Te
Figs 1-8 Oncaspidia pilosicollis. (1) head, left lateral view; (2) pronotum, dorsal view; (3) genital
capsule with parameres, dorsal view; (4) conjunctiva, left lateral view (vesica strongly flexed to
right, strongly foreshortened in this view); (5) genital capsule, right lateral view; (6) left paramere,
medial view; (7) VII and succeeding segments of 2 abdomen with ovipositor exserted; (8)
spermatheca.
Spine above antennifer slender, curved (Fig. 1); external apical process of antennifer porrect, apically
rounded, its dorsal margin flexed inwards almost horizontally. Ratio of lengths of antennal segments
1:11: Ill :IV not differing significantly between the sexes, about 1-00 : 0-80 : 0-65 : 0-86; length of
segment I divided by width of head including eyes in both sexes 1:06-1:32, mean 1-17 (n = 21). Ratio of
lengths of rostral segments I : II : III : IV about 1-00 : 0-90 : 0-70 : 1-08 in both sexes.
Pronotum (Fig. 2) steeply declivent anteriorly, its posterolateral angles rather narrowly produced
anterolaterally and tapering into slender spines; width of pronotum across tips of spines divided by width
of head including eyes in male 2-62-3-09, mean 2-80 (n = 11), in female 2-76-3-07, mean 2:89 (n = 10).
Corium with apex slightly produced, at rest reaching posteriorly to a point level with the end of the basal
third of laterotergite VI. Anterior and intermediate femora each with 0-1 very small subapical spine
beneath; posterior femur with 2 major spines, the first about half as long as the second, with about 3 short
spines or granules between them, and a terminal series of 4 spines. Posterior tibia not arcuate basally, its
length divided by length of posterior femur 0-75—0-84, mean 0-80 (n = 21).
Abdominal sternites III-VII each with posterolateral angles drawn out into narrowly triangular spines
about one-quarter as long as lateral margin of segment. Male genital capsule (Figs 3, 5) closed posteriorly,
lip distinctly produced and pouched, tongue triangular. Paramere with apex capitate (Fig. 6), the capitate
ends of the two parameres almost horizontal at rest and largely occluding, in dorsal view, space between
tongue and lip of capsule (Fig. 3). Conjunctiva (Fig. 4) with dorsomedian lobe broad, low, transverse;
intermediate dorsal lobe broad, high; distal dorsomedian lobe small, rounded; distal dorsolateral lobes
AFRICAN POD BUGS 5
absent; apical ventral lobes small, transverse; distal ventrolateral lobes each with a rounded, membranous,
posterior subdivision and a reflexed, dorsally sclerotized, narrow, anterior subdivision; no other lobes
sclerotized. Ejaculatory reservoir complex with wings and straps long, symmetrical; vesica long, sclerites
protecting its base large, almost symmetrical. Ovipositor with valvulae very long (Fig. 7). Sclerites of
dorsal wall of gynatrium narrowly V-shaped, longitudinal arms slightly more than twice as long as trans-
verse arms, both arms ribbon-like in vertical plane. Spermatheca (Fig. 8) with bulb fusiform, duct tightly
convoluted in part adjacent to bulb. Ovarian egg (only 1 examined) 1-38 x 0-52 mm, with 6 micropyles.
Head weakly granulate, antennal segments I-III very weakly and minutely granulate. Pronotum granu-
late, disc also punctate, propleura granulate-punctate, area of pronotum and propleura immediately
anterior to each posterolateral angle with about 8 small, blunt, piceous tubercles. Scutellum with entire
surface coarsely and irregularly granulate. Meso- and metapleura granulate-punctate. Clavus seriately
punctate in 3 rows. Corium more weakly punctate than clavus, especially at apex, punctures in basal half
seriate. Veins in basal half of corium very weakly granulate. Femora and tibiae minutely and weakly
granulate. Abdominal sterna transversely rugose, laterotergites weakly granulate, tergites I and II trans-
versely rugose, III and IV with large, sometimes confluent punctures largely restricted to median half of
each, V-VII with smaller punctures occupying whole width of V and VI but restricted to centre third of
VII.
Long, erect, colourless pubescence present on head, antennal segments I-III, pronotum, scutellum,
thoracic pleura, abdominal sterna and throughout all legs. Shorter, brown, erect pubescence arising from
tubercles in anterior midline of pronotum. Shorter, suberect pubescence present on clavus, corium,
abdominal laterotergites and rostrum and also mixed with the longer type of hairs on antennae, legs and
abdominal sterna, usually colourless but pale or dark brown where it arises from similarly coloured areas
of corium. Short to moderately long, generally decumbent and more or less tomentose pubescence present
on head, antennal segments I-III (here very sparse), pronotum, scutellum, thoracic pleura, abdominal
sterna and femora, usually appearing white or silvery but brown in 4 slightly oblique longitudinal bands on
pronotum (Fig. 2).
Colour of integument generally pale pinkish brown. Ocellar tubercles, extreme base of scutellum,
anterior midline of pronotum, posterolateral spines of pronotum, ventral surface of head, thoracic sterna,
spots on coxae, centre of abdominal sterna II and III and parts of abdominal terga, especially punctures,
piceous to black. Antennal segments I-III except for apices, bases of femora, tibiae except for basal,
apical and sometimes median annuli, parts of tarsi and almost the whole of laterotergites VI and tergum
VII stramineous. Hemelytra with clavus and basal half of corium stramineous, with a few spots on anterior
veins dark brown, apical half of corium variegated pale and dark brown, membrane colourless, hyaline.
REMARKS. A distinctive species, as indicated in the generic description above. I follow Stal’s
(1865 : 107) interpretation of Clavigralla similis Signoret, which is based on a specimen in Vienna
apparently determined by Stal as being Signoret’s type-material. Signoret’s (1860 : 944) descrip-
tion is inadequate and confusing.
DISTRIBUTION. Tropical and southern Africa but not Madagascar. Apparently less common in
West Africa. Perhaps avoids densely forested areas.
MATERIAL EXAMINED
Clavigralla pilosicollis Stal, lectotype 2, South Africa: ‘Caffraria’ (J. Wahlb.) (NR, Stockholm). Clavi-
gralla similis Signoret, lectotype 3, Zanzibar [bears labels: (1) Zanzibar/ coll. Signoret, (2) similis/ det.
Signoret, (3) pilosicollis/ det. Stal] (NM, Vienna).
Liberia: 1 9, Kpaine, 30.vii.1953 (W. Peters) (BMNH, London). Ghana: 1 9°, Wegbe, Volta River,
1.xi.1967 (D. Leston) (UG, Legon). Nigeria: 1 9°, Ibadan, 15-19.xii.1975 (J. C. Deeming) (IAR, Samaru).
West Africa: 3 2, ‘Cameroons’ (ex coll. Distant); 1 2, ‘W. Afr.’ (BMNH, London). Zaire: 1 9, Kasongo,
viii.1959 (P. L. G. Benoit); 1 9, Sankuru, M’Pemba Zeo (Gandajika), 9.i.1960 (R. Maréchal) (MRAC,
Tervuren); 1 3, Ngowa Kwango, 3.ii.1938 (J. Mertens); 1 3, Ngowa, 9.v.1939 (R. P. J. Mertens); 1 3,
Nguela, Usambara (no other data); 1 3, Marais Kululu, 8.vi.1939 (H. J. Brédo); 1 3, Lubumbashi,
9.iii.1939 (Brédo); 1 3, Mpese, v-vi.1937 (R. P. J. Cooreman) (IRSNB, Brussels); 2 3, E. of R. Semliki,
1931-1932 (E. B. Worthington); 1 2°, Katanga, Kambove, 4000-5000 ft (1200-1500 m), 25.iii.1907 (Neave)
(BMNH, London). Uganda: 1 3, Mabira, on dung, 16.viii.1925 (G. L. R. Hancock); 1 3, Mbarara,
29.v.1911 (C. C. Gowdey); 1 3, top of escarpment N. of Fort Portal, 4700 ft (1430 m), 24.xi.1911 (S. A.
Neave); 1 2, Mpumu (Miss M. Robertson) (BMNH, London). Kenya: 1 3, 12 miles (19 km) NW. of
Mazeras, c. 500 ft (150 m), 23.ix.1905 (E. B. Poulton) (UM, Oxford); 1 3, Chyulu Hills, 5600 ft (1680 m),
vii.1938 (Coryndon Mus. Exped.); 1 2, Mombasa (van Someren) (BMNH, London). Tanzania (Mainland):
6 W. R. DOLLING
1 3, Mbeya Mountain, 33° 25’ E, 8° 48’ S, 7000 ft (2100 m), 5.viii.1959 (Cambridge Univ. Exped.); 1 3, E.
rim of Ngoro-ngoro Crater, 7000 ft (2100 m), 20.vii.1959 (Cambridge Univ. Exped.) (BMNH), London).
Zanzibar: 2 3, 1 9, i-ii.1925 (H. J. Snell) (BMNH, London). Malawi: 2 3, Mlanje, 1.vii.1913 and 15.viii.
1913 (S. A. Neave); 1 2, Mlanji Boma, 2400 ft (720 m), 26.iv—5S.v.1910 (Neave) (BMNH, London).
Rhodesia: 1 3, 1 2, Mashonaland, Salisbury, 1898 (BMNH, London). South Africa: 2 3, 3 2 (no further
locality data) (A. L. Capener); 2 3, 2 9, N. Transvaal, Louis Trichard, 20—30.xii.1956 (Capener) (Slater
coll.); 8 3, 2 2°, Natal, Durban (AMNH, New York); 1 3, 1 9, Wonderboom, 12.iii.1905 (Swierstra) (TM,
Pretoria); 1 9, Natal, Lake St Lucia, Charters Creek, 12.xi.1967 (J. A. & S. Slater, T. Schuh) (PPRI,
Pretoria); 1 3, Pondoland, Port St John, 5—30.vi.1928 (R. E. Turner); 1 3, 1 9, Natal, Umbilo, 14.vi.1914
(L. Bevis); 3 3, 1 9, Natal, Malvern, 13.vi.1897, 16.vi.1897, vii.1897, 10.vi.1897; 1 3, Zululand, Mtunzini,
7.vi.1926 (Turner); 1 3, Pretoria, 12.iii.1905 (C. J. Swierstra) (BMNH, London).
CLAVIGRALLA Spinola
Clavigralla Spinola, 1837 : 200. Type-species: Clavigralla gibbosa Spinola, by monotypy.
Acanthomia Stal, 1873 : 81, 82. Type-species: Clavigralla natalensis Stal, by PRESENT DESIGNATION.
Syn. n.
Lancha Shadab, 1972 : 3-7. Type-species: Myla schnelli Villiers, by monotypy. Syn. n.
Length of head half to three-quarters that of pronotum. Antennifers weakly to strongly divergent, process
at outer apical angle of antennifer short and porrect or long and deflexed with its apex in contact with
maxillary plate. Side of head in front of eye without spine.
Pronotum rather strongly to very strongly declivent anteriorly, posterolateral angles weakly to very
strongly produced, each bearing apically a usually slender spine. Disc of pronotum rarely without tubercles
or spines, never with a group of four stout, blunt tubercles in middle, usually with a stout, blunt tubercle
or spine close to lateral margin about half way between posterolateral and anterolateral angles, often with
a more or less regular, anteriorly concave semicircle of spines or tubercles posterior to this pair. Scutellum
equilateral or slightly longer than its basal width, flat to strongly convex, midline impressed in convex
forms. Metapleural scent-gland peritreme with dorsal ridge entire and shortly reniform or bilobed. Corium
with apex scarcely to distinctly produced.
Abdomen with posterolateral angles of sternites IIJ-VII usually prominent, often strongly and spin-
ously produced. Male genital capsule with lip narrow or broad, often filling posterior emargination of
capsule, tongue entire or apically trifid or bifid. Tergum IX of female usually in horizontal plane, rarely
apically deflexed. Valves of ovipositor not especially long. Spermatheca with bulb narrowly lunate, duct
looped to strongly convoluted. Sclerites of wall of gynatrium basically V-shaped, not deep in vertical
plane.
REMARKS. When Stal (1873) differentiated his new genus Acanthomia from Clavigralla he listed
five species under the latter but separated the first of these, C. horrens Dohrn, from the remaining
four species (including gibbosa, the type-species) by a horizontal rule across the page. This con-
vention presumably indicates that he had not seen specimens of the species below the rule. The
character he uses to separate Clavigralla sensu Stal from Acanthomia in his key is the relative
lengths of rostral segments II and IV, with IV equal to or longer than II in Acanthomia and shorter
than II in Clavigralla. While IV is slightly shorter than II in horrens, it is definitely longer in
gibbosa. This confirms that gibbosa, which is closely related to tomentosicollis, was unknown to
Stal. I have selected C. natalensis, a species described by Stal himself, as type-species of Acan-
thomia. If it were necessary to treat the species related to e/ongata in a different generic or sub-
generic taxon from those related to tomentosicollis and gibbosa, the name Acanthomia would be
available for the former. However, this course of action seems inadvisable in view of the size of
the genus and the fact that a further generic name would be required for tuberculicollis and its
allies. Shadab (1972) recognized that Myla schnelli Villiers did not belong to the genus Myla but
overlooked its close affinity with C. elongata and erected the unnecessary new genus Lancha for it.
SPECIES GROUPS AND SUBGROUPS. There are three major external features which may be used as a
basis for defining groups of species within the genus. First, the external apical process of the
antennifer may be short and porrect (Fig. 9) or long and deflexed (Fig. 51). Secondly, the dorsal
AFRICAN POD BUGS a
ridge of the metathoracic peritreme may be bilobed (Fig. 12) or entire (Fig. 54). Thirdly, the
pilosity of the pronotum may be uniform (Fig. 50) or divided into an anterior area of pale,
tomentose pubescence and a posterior area of darker, erect to decumbent pubescence (Fig. 188).
On the basis of comparisons with related genera and of correlation with other presumed apo-
morphies, the second alternative state of each character mentioned seems to be apomorphic with
respect to the first. Three groups of species may be recognized: (1) a group of species related to
C. tuberculicollis in which the apical process of the antennifer is short and porrect, the pubescence
of the pronotum is uniform and, except in the case of C. uelensis, the dorsal ridge of the meta-
thoracic peritreme is bilobed; (2) a more advanced group related to C. elongata in which the apical
process of the antennifer is long and deflexed, the dorsal ridge of the metathoracic peritreme is
entire and the pubescence of the pronotum uniform; (3) a group of species related to C. tomento-
sicollis which differs from the e/ongata-group in having the pubescence of the anterior and poster-
ior parts of the pronotum dissimilar. C. uelensis has probably acquired the entire state of the dorsal
ridge of the peritreme independently from the e/ongata- and tomentosicollis-groups, as it is very
similar in many details to C. /eroyi. In all members of the tuberculicollis-group where the male is
known (i.e. all except C. uelensis) the posterior lip of the genital capsule does not completely fill
the posterior emargination and the apices of the parameres are visible in posterior view (Fig. 11),
as in most other Pseudophloeinae, whereas in the elongata- and tomentosicollis-groups the
posterior lip is much better developed and conceals the apices of the parameres in posterior view.
This confirms the conclusion that these two groups are more highly evolved than the tuberculi-
collis-group. In the elongata-group the pronotum bears numerous to few spines on the disc, usually
with some of them organized into a semicircle terminating anteriorly in a pair of sublateral spines
(e.g. Fig. 50) as in the tuberculicollis-group. In the tomentosicollis-group most of the spines are
very small and concealed by the pubescence, only the large, sublateral pair being retained (e.g.
Fig. 188). Even these are lacking in C. /eontjevi (Fig. 131). The posterolateral pronotal angles in
the e/ongata-group are in general more strongly produced than those of the tomentosicollis-group
and the posterolateral spines arise less abruptly from them. These two characters tend to confirm
the validity of the groups and the relatively greater apomorphy of the tomentosicollis-group, as
the pattern of pronotal spines and the shape of the posterolateral pronotal angles in the e/ongata-
group are shared with the less highly evolved tuberculicollis-group.
The tuberculicollis-group is too small to warrant subdivision and the e/ongata-group does not
usefully lend itself to such treatment as the patterns of relationship within it are obscured by the
striking apomorphies of several of its included species. In this group, close relationships obviously
exist between hystrix and hystricodes and between mira and annectans; natalensis is rather isolated
and the remainder of the group fall into two series, those in which the veins of the hemelytral
membrane are finely pigmented (e/ongata to schnelli in the check-list) and those in which they are
heavily and broadly pigmented (insignis to aculeata). The tomentosicollis-group can usefully be
divided into five subgroups. The first contains only /eontjevi, which is unique in lacking sublateral
pronotal tubercles. It shares the character of an entire tongue in the genital capsule with the next
two subgroups. Primitively, the tongue is triangular when seen in dorsal view (Figs 140, 181) with
a prominent, ventral, longitudinal keel. Fhere isa tendency for the tongue to become apically
rounded and in C. strabo (Fig. 170) it appears in dorsal view to be mucronate as the keel can be
seen projecting beyond the rounded apex of the dorsal surface of the tongue. In the ruandana-
subgroup the pigmentation of the hemelytral membrane is almost uniformly diffused and the head
is elongate while in the spiniscutis-subgroup the membrane is spotted and the head is of normal
length for the genus with the exception of bovilla in which the head is elongate. Possibly this
species would be better placed in the ruandana subgroup, although its spotted membrane would
be anomalous. The trend in the ruandana- and spiniscutis-subgroups for the tongue of the genital
capsule to depart from the ancestral triangular shape is continued in the tomentosicollis-subgroup.
Here the tongue is apically emarginate and the apex of the ventral keel projects into this emargina-
tion, giving the tongue a trifid appearance (Fig. 194). In all members of this subgroup the pig-
mentation of the membrane of the hemelytron is evenly diffused. The most highly evolved form
of the tongue is encountered in the wittei-subgroup, where it is deeply bifid and the keel is greatly
reduced (Fig. 222). The hemelytral membrane in the species of this subgroup is spotted.
8 W. R. DOLLING
Key to groups and subgroups
1 Antennifer with outer apical process short, porrect (Fig. 9) : . tuberculicollis-group (p.
— Antennifer with outer apical process long, deflexed (Fig. 51) :
2 Pubescence of pronotum almost uniform. . ; elongata-group (p. ey
— Pubescence of anterior part of pronotum differing markedly i in colour and texture from that of
posterior part (tomentosicollis-group) . , Z : 3
3 Pronotum without sublateral tubercles (Fig. 131) : ; : leontjevi-subgroup (p. 48)
— Pronotum with a pair of large, sublateral tubercles (Fig. 138) . : , ; 4
4 Tongue of male genital capsule entire. : F ; : ‘ ; g ; 5
— Tongue of male genital capsule apically emarginate F ; : ‘ ; : : : 6
5 Membrane of hemelytron with diffuse pigmentation . ‘ : . ruandana-subgroup (p. 50)
— Membrane of hemelytron spotted . : : . spiniscutis-subgroup (p. 54)
6 Tongue of male genital capsule apically trifid (Fig. 194) 4 : tomentosicollis-subgroup (p. 63)
— Tongue of male genital capsule apically bifid (Fig. 221) . ; ; . wittei-subgroup (p. 74)
DISTRIBUTION. Tropical and southern Africa, Madagascar and islands adjacent to these, southern
Arabia, Oriental region eastwards to Java. The genus is essentially African. Two invasions of the
Oriental region have occurred, once by the common ancestor of gibbosa and orientalis and once
by scutellaris, which is also present in Arabia and in the more arid areas of Kenya and Sudan.
C. scutellaris is the sister-species of C. tomentosicollis; orientalis and gibbosa are more distantly
related but belong to the tomentosicollis-subgroup. All three species-groups are represented in
Madagascar and the two known Malagasy species of the tomentosicollis-group belong to different
subgroups. Only C. elongata seems to be present on the more remote and smaller islands in the
Ethiopian region, occurring in the Canary and Cape Verde Islands, Principe, Sido Tomé, Sey-
chelles, Réunion, Mauritius and Rodriguez as well as Arabia (where it is the only species to be
found other than scutellaris), Madagascar and the African mainland. C. tomentosicollis is repre-
sented on the Comoro Islands. The area of greatest species diversity is the highlands of central
Africa, especially northern Zambia and the vicinity of the Rift Valley. Several species have very
wide distributions within Africa. The most important are tomentosicollis, elongata, shadabi, wittei,
spiniscutis, curvipes, simillima, leontjevi, hystrix and hystricodes. Of these, curvipes and simillima
are sister species with vicarious distributions and shadabi appears to be replacing elongata in the
mainland of West Africa, where e/ongata is restricted to the offshore island groups only, and in
lowland central Africa, where the two species coexist over a wide area.
Key to African and Malagasy species
Note. Beware the superficially similar Oncaspidia Stal and Myla Stal. The former is distinguished by the
spine above the base of the antenna (Fig. 1). Myla belongs to a large group of genera of Pseudophloeini
in which there is a small tubercle close to the base of the posterior face of the posterior femur (Fig. 247).
This tubercle is lacking in Clavigrallini. The pubescence of Myla is never tomentose.
1 Antennifer with apical process short, porrect (Figs 9, 34). : : : A 4 ; 2
— Antennifer with apical process long, deflexed (e.g. Figs 51, 137) : >
2 Robust species with posterolateral angles of abdominal sternites (Fig. 13) and tubercles and
posterolateral angles of pronotum (Fig. 10) very prominent (Madagascar)
C. tuberculicollis (Reuter) (p. 12)
— Less robust species with posterolateral angles of abdominal sternites (Figs 22, 29, 35) and
tubercles and posterolateral angles of pronotum (Figs 20, 21, 27, 31) less prominent.
(African mainland, not Madagascar) . ‘ 3
3 Metathoracic scent-gland peritreme with dorsal ridge entire (Fig. 33). (Central Africa)
C. uelensis (Schouteden) (p. oy
— Metathoracic scent-gland peritreme with dorsal ridge bilobed (as Fig. 12) . d
4 Pronotum (Fig. 27) with posterolateral angles scarcely produced, width across apices of
posterolateral spines divided by width of head sic eyes 2:10 or less. (Southern central
Africa) . : C. zambiae sp. n. (p. 17)
— Pronotum (Figs 20, 21) with posterolateral angles more strongly produced, width across apices
of posterolateral spines divided by width of head including eyes 2:14 or more. (Widespread
in tropical Africa from Guinea and Sudan to Angola and Malawi)
C. leroyi (Schouteden) (p. 14)
13
14
\.15
16
AFRICAN POD BUGS 9
Pubescence of anterior, declivent part of pronotum of almost uniform colour and texture with
that of posterior, horizontal part. Posterolateral angles of pronotum strongly produced,
passing gradually into posterolateral spines (e.g. Figs 36, 43, 50, 78, 83, 87-89, 126) . : 6
Pubescence of anterior, declivent part of pronotum pale, silvery or golden, tomentose, that of
posterior, horizontal part darker, brown, erect to decumbent, the two areas abruptly
demarcated each from the other along an almost straight to sharply undulating or zigzag
line joining posterolateral angles. Posterolateral angles usually only slightly prominent with
spines arising abruptly from them (e.g. Figs 131, 138, 144, 148) . , 25
General coloration blackish brown, forewing membrane heavily blotched with dark chocolate
brown . |
General coloration greyish, yellowish or reddish brown, forewing membrane with brown lines
generally following course of veins. 8
Antennal segment I longer than 1-7 times width of head including eyes. Antennae with rather
obscure pale and dark annulations. (Widespread in equatorial Africa) C. hystrix (Dallas) (p. 19)
Antennal segment I shorter than 1-5 times width of head including eyes. Antennae with con-
spicuous pale and dark annulations. (Widespread in tropical Africa) C. hystricodes Stal (p. 21)
Scutellum with two longitudinal rows of outstanding, blunt spines on disc (Fig. 123). (South
Africa and Rhodesia) ‘ : : : : . C. horrida (Germar) (p. 44)
Scutellum never with outstanding spines : . 3 : : : : : ‘ 9
Head with frontal area inflated (Figs 97, 243). ‘ : : ; 10
Head with frontal area declivent from behind level of antennae (Fig. 51). : : ie |
Distal dorsomedian lobe of conjunctiva bearing sclerotized cues (Figs 100, 101).
(Western, central and eastern Africa) . . CC. mira sp. n. (p. 36)
Conjunctiva without sclerotized appendages (Figs 245, 246). (Zaire) . C. annectans sp. n. (p. 38)
Pronotum (Fig. 126) with posterolateral lobes and spines very short. Scutellum weakly convex.
Spines at posterolateral angles of abdominal sterna very small (Fig. 128). (Southern Africa)
C. natalensis Stal (p. 46)
Pronotum (e.g. Figs 50, 104, 117) with posterolateral angles and spines more strongly
produced. Scutellum usually more strongly convex and abdominal spines longer _.. 12
Pubescence of head, pronotum and scutellum not or poorly organized into narrow, longitudinal
lines of white hairs; if such pale lines are partially discernible they are never present on
declivent part of pronotal disc posterior to calli. Forewing membrane colourless, its veins
finely picked out in brown. (Several common species, throughout Africa and Madagascar). 13
White pubescence organized into conspicuous, narrow longitudinal lines on head, pronotum
and scutellum, including three lines traversing entire length of pronotum. Forewing
membrane with broad bands of brown coloration following course of veins and usually
spreading onto membrane between veins especially on disc. (Species of restricted distribu-
tion, mostly confined to high ground in eastern, central and southern Africa; not known
from West Africa or Madagascar) . 19
Scutellum flat, not elevated above level of posterior margin of pronotum, Pronotum with
posterolateral angles strongly produced anterolaterally (Figs 87, 88, 89, 91). (Western and
central Africa). : . _C. schnelli (Villiers) (p. 34)
Scutellum convex (Figs 53, aN. elevated above level of posterior margin of pronotum when
viewed in profile. i.
Spines at posterolateral angles of abdominal ‘sternites long, almost perpendicular to lateral
margins of abdomen (Figs 79, 80). Scutellum without long, erect pubescence (Madagascar is
only). ;
Spines at posterolateral angles of abdominal sternites shorter, usually distinctly deflected
posteriad (Fig. 52). Scutellum with at least some long, erect pubescence (Africa and
Madagascar) . 16
Antennal segment I longer than 1 4 times width of head including eyes. Produced, postero-
lateral angles of pronotum and of abdominal sternites (Figs 78, 79) more slender. (Mada-
gascar) . E . _C. madagascariensis sp. n. (p. 31)
Antennal segment I shorter than i: ‘4 times width of head including eyes. Produced, postero-
lateral angles of pronotum and of abdominal sternites (Figs 83, 80) stouter. (Madagascar)
C. asterix sp. n. (p. 33)
Scutellum moderately convex and strongly tuberculate (Fig. 77). (Madagascar)
C. ankatsoensis sp. n. (p. 31)
Scutellum strongly convex with less prominent tuberculation (Fig. 53) : : : : WF
10
17
21
22
25
26
29
W. R. DOLLING
Head short (Fig. 72). Tibiae with dark, basal, apical and median annuli. (West and central
Africa) . : ; 7 C. breviceps sp. n. (p. 30)
Head longer (Figs 50, 66). Tibiae without care peedien aati ; ’ , 18
Semicircle of spines on disc of pronotum with anterior pair much more prominent than the
pair behind them (Fig. 66). (West and central Africa) . : : C. shadabi sp. n. (p. 28)
Semicircle of spines on disc of pronotum with the first two pairs of spines of comparable size
(Fig. 50). (Central, eastern and southern Africa and Canary Is., Sao Tomé, Principe, Sey-
chelles, Madagascar, Réunion, Mauritius, Rodriguez, S. Arabia) . C. elongata Signoret (p. 24)
Width across apices of posterolateral spines of pronotum greater than 3-0 times width of head
including eyes . é 20
Width across apices of posterolateral spines of pronotum less than a ‘0 times width of head
including eyes . ' 21
Length of antennal segment I greater than 1: 6 times width of head including eyes. Spinously
produced posterolateral angles of abdominal sternites perpendicular to lateral margins of
abdomen. Width of pronotum across apices of posterolateral spines more than 3-5 times
width of head including eyes. (Malawi) . ; . C. egregia sp. n. (p. 41)
Length of antennal segment I less than 1-6 times ondih of heaa lachiding eyes. Spinously pro-
duced posterolateral angles of abdominal sternites deflected posteriad. Width of pronotum
across apices of posterolateral spines less than 3-5 times width of head including eyes.
(Zambia, Zaire) : : : ; C. longispina sp. n. (p. 43)
Posterolateral spines of abdconnal fernitee more neatly perpenalicules to lateral margins of
abdomen and broader at the base (Figs 113, 115) ; F 22
Posterolateral spines of abdominal sternites strongly deflected bpueaca and narrower acre
106) : : 23
Width of pronotum across apices of posterolateral spines greater than 2:9 times width of head
including eyes. Male paramere with apical tooth less prominent (Fig. 114). (Angola)
C. angolensis sp. n. (p. 41)
Width of pronotum across apices of posterolateral spines less than 2-9 times width of head
including eyes. Male paramere with apical tooth more prominent (Fig. 119). (South Africa)
C. aculeata sp. n. (p. 43)
Scutellum weakly convex (Fig. 108). Male paramere with apical margin strongly oblique (Fig.
111). (East Africa) . ; C. andersoni sp. n. (p. 41)
Scutellum strongly convex (Fie. 107). Male paramere with apical margin less strongly oblique
(Figs 109, 110) : : 24
Male paramere with apical margin straight, ‘blade longer in ‘relation to shaft (Fie, 109).
(Highlands of central Africa) . : C. insignis (Distant) (p. 39)
Male paramere with apical margin convex, Biude Reentel in reigion to length of shaft (Fig.
110). (Burundi) : ‘ C. minor (Schouteden) (p. 39)
Pronotum without large, blunt tubercles at sides of disc (Fig. 131). (Western, central and
eastern Africa) : : i C. leontjevi (Bergroth) (p. 48)
Pronotum with a pair of fares. usually, blunt fibeicles at sides of disc behind level of calli (e.g.
Figs 188, 209, 220) . : 26
Membrane of hemelytron with disc evenly suffused with pale to medium brown coloration,
rarely with a few spots of slightly more intense pigmentation between ee mae veins near
basal cells, in which case male genital capsule with tongue trifid . , 27
Membrane ot hemelytron largely colourless or very faintly milky white, with 1 < neon or
piceous spots between bases of longitudinal veins where these emerge from basal cells (Fig.
219); spots sometimes very faint; membrane occasionally with more extensive piceous
blotching (Fig. 178); tongue of male genital capsule never trifid . ; 34
Length up to 8-5 mm (largest females); posterior tibia short, strongly arcuate at base (Fig.
212) ; A 28
Length greater than 8: 5 mm except for smallest males, in which case posterior tibia long and
only weakly arcuate at base (Fig. 185) : ‘ ; 29
Antennal segment I slender (Fig. 211). (Tropical Africa north of latitude 3° S)
C. curvipes (Stal) (p. 70)
Antennal segment I stouter (Fig. 210). (Africa south of latitude 6° S) . C. simillima sp. n. (p. 73)
Length of antennal segment I divided by width of head including eyes 1-11 or less and head
elongate (Fig. 137). Male genital capsule with tongue entire. : : : ' ; 30
33
34
35
36
40
AFRICAN POD BUGS
Length of antennal segment I divided by width of head including eyes greater than 1-11 or, if
less, head not elongate (Figs 187, 189) and tongue of male genital capsule trifid (Fig. 194)
Pronotum with posterolateral spines very short (Fig. 148). Abdominal sternites with postero-
lateral angles not at all prominent (Fig. 150). (Northern Zambia) . . C. oxonis sp. n. (p.
Pronotum with posterolateral spines longer (Fig. 138). Abdominal sternites with posterolateral
angles distinctly prominent (Fig. 139). (Highlands of central Africa)
C. ruandana (Schouteden) (p.
Pronotum with posterolateral spines directed strongly posteriad (Fig. 144). Male genital cap-
sule with tongue entire. (Highlands of central Africa) . : ; . C. biston sp. n. (p.
Pronotum with posterolateral spines directed laterad (Figs 188, 190, 203). Male genital capsule
with tongue trifid :
Band of dark pubescence on pronotum posterior to posterolateral angles almost completely
divided into two by a narrow, V-shaped, median, longitudinal band of pale pubescence
(Fig. 203). Posterior tibia distinctly arcuate at base. (Madagascar) C. annulipes Signoret (p.
Band of dark pubescence on pronotum posterior to posterolateral angles not divided (Figs
188, 190). Posterior tibia only weakly arcuate at base (Fig. 192). (Not occurring in Mada-
gascar) :
Pronotum with area of pale mabewcsnce Zonpletely divided ae dark. lonigituditial bbaud:
junction with area of dark pubescence with two small, posteriorly projecting extensions of
pale pubescence close to midline (Fig. 188). (Africa south of the Sahara except for the
north-eastern part; Comoro Is.) : C. tomentosicollis Stal (p.
Pronotum with area of pale pubescence incompletely divided , junction of pale and dark areas
following a gently undulating line without distinct, narrow projections of pale pubescence
into dark area (Fig. 190). (Northern Kenya, Sudan, Arabia, SW. Asia)
C. scutellaris (Westwood) (p.
Form of body depressed. Scutellum only moderately convex (Figs 231, 232). Abdominal
sternites with posterolateral angles only slightly prominent (Figs 233, 234). Pronotum with
posterolateral angles triangularly prominent, their spines very short (Figs 229, 230). Length
of antennal segment I divided by width of head including eyes 0-80 or less ;
Deeper bodied. Scutellum usually more strongly convex. Abdominal sternites with postero-
lateral angles usually more strongly produced (Figs 154, 158, 169, 179, 224). Posterolateral
angles of pronotum, if produced, not triangularly so, their spines longer (Figs 157, 166, 167,
177, 220). Length of antennal segment I divided by width of head including eyes greater
than 0-80 : :
Abdominal spines slightly shorter (Fig. 233). Body and appendages with all erect hairs shorter
than transverse dorsal diameter of eye. (Highlands of north-east and central Africa)
C. alpica (Bergroth) (p.
Abdominal spines slightly longer (Fig. 234). Many erect hairs on head, thorax and appendages
as long as transverse dorsal diameter of eye (e.g. pubescence of scutellum, Fig. 232).
(Cameroun) . C. montana sp. n. (p.
Antennal segment I very long, its length greater than 13 times width of head including eyes.
Posterior tibia very weakly arcuate at base (Fig. 185), more than 0-75 times length of
posterior femur. (Southern Africa) . . C. marmorata sp. n. (Pp.
Antennal segment I shorter than 1-25 times width of head including eyes or, if longer, then
posterior tibia more strongly arcuate at base and not exceeding 0-70 times length of
posterior femur
Posterolateral angles of abdominal sternites very weakly produced (Fig. 154). Head elongate
(as in ruandana, Fig. 137). (Zambia) . : . C. bovilla sp. n. (p.
Posterolateral angles of abdominal sternites more strongly produced (Figs 158, 169, 179, aia
Head less elongate
Male genital capsule with tongue narrowly triangular. (Madagascar). ay a pusilla sp. n. (p.
Male genital capsule with tongue of various shapes but never narrowly triangular. (Not in
Madagascar) .
Pronotum with posterolateral angles rather strongly produced (Fi g. 220). Male genital capsule
with tongue bifid.
Pronotum with posterolateral angles weakly produced (Figs 166, 167 175, 177). Male genital
capsule with tongue entire
Length greater than 7-5 mm. Lobes of tongue of male genital capsule laminar, externally and
apically deflexed (Fig. 221). (Widespread in Africa) . 7 . C. wittei (Schouteden) (p.
33
63)
69)
35
36
tA)
80)
63)
37
54)
38
56)
39
40
41
74)
12 W. R. DOLLING
— Length less than 7-5 mm. Lobes of tongue of male genital capsule cylindrical (Fig. 222).
(Uganda, Zaire) ; : C. neavei sp. n. (p. 77)
41 Posterior tibia not arcuate at base. Antennal segment I with many outstanding hairs longer
than diameter of segment. Tongue of male genital capsule apically rounded (Fig. 170).
(Southern Africa) . ; . C. strabo sp. n. (p. 57)
— Posterior tibia gently or strongly arcuate at base. Antennal segment I with no hairs as long as
its diameter. 42
42 Posterior tibia gently arcuate at base. Tongue of male genital capsule narrow, truncate (Fig.
174). (Southern Africa) . . _C. pabo sp. n. (p. 59)
— Posterior tibia strongly arcuate at base (as i in C. Saline Fig. 212). Tongue of male genital
capsule broad, mucronate (Fig. 181). (Widespread in Africa) C. spiniscutis (Bergroth) (p. 60)
Descriptions of species
The tuberculicollis-group
Antennifer with apical process short, porrect; metathoracic peritreme with dorsal ridge bilobed or entire;
male genital capsule with tongue entire; lip of genital capsule not completely filling posterior emargination
in posterior view; pubescence of anterior, declivent area of pronotum uniform with that of posterior,
horizontal area.
Clavigralla tuberculicollis (Reuter) comb. n.
(Figs 9-18)
Acanthomia tuberculicollis Reuter, 1887 : 90. Holotype 2, MADAGASCAR [destroyed].
Length: 3, 9:7-10:8 mm, mean 10:2mm (n = 7); 9, 11-0-11-5mm, mean 11-3 mm (n = 5). Body
robust, connexivum broad.
Antennifers strongly divergent, outer apical angle of each with a well-developed, porrect, triangular
process (Fig.9). Ratio of lengths of antennal segments I: II : III : IV in male about 1-00 : 0-91 : 0-83:
0-83, in female about 1-00 : 0-92 : 0-84 : 0-71. Length of antennal segment I divided by width of head
including eyes in male 1-23-1-28, mean 1-26 (n = 7), in female 1-29-1-44, mean 1-35 (n = 4). Ratio of
lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0-89 : 0:57 : 0-88.
Pronotum (Fig. 10) strongly declivent, disc tuberculate, sublateral pair of spines very prominent and
robust, posteriorly with a raised ridge parallel to posterior margin, the ridge expanded medially to form a
low boss. Width of pronotum across apices of posterolateral spines divided by width of head including
eyes in male 2:67-2:91, mean 2:84 (n = 7), in female 2:67-3:21, mean 2:94 (n = 4). Scutellum strongly
convex, disc with a pair of prominent tubercles, apex elevated slightly and knobbed. Metathoracic scent-
gland peritreme (Fig. 12) with dorsal ridge bilobed, anterior lobe slightly larger than posterior. Corium
with apex slightly produced, at rest reaching level of suture between laterotergites V and VI. Anterior and
intermediate femora each with a single, strong, subapical spine beneath. Posterior femur subapically with
a rather small major spine followed after a short space by a larger major spine and a terminal series of 4
spines decreasing in length towards apex of femur. Posterior tibia almost straight, about 0-88 times length
of posterior femur.
Abdominal sterna III-VII with posterolateral angles produced into broad, triangular teeth (Fig.13).
Male genital capsule (Fig. 11) with lip small, not filling posterior emargination, tongue long and narrowly
triangular. Paramere (Fig. 15) broad and flattened. Aedeagus (Figs 16-18) with conjunctival lobes all
membranous; dorsomedian and intermediate dorsal lobes broad, low, M-shaped, distal dorsolateral lobes
long, distal dorsomedian lobe narrow, finger-like, small, proximal ventrolateral lobes present, distal
ventrolateral lobes long, apical ventral lobes large. Female sternum VII narrow, especially in midline,
cleft for about one-third of its length. Second valvula truncate apically. Sclerites of gynatrial wall simply
V-shaped. Spermatheca (Fig. 14) with bulb long and narrow, duct rather short and scarcely convoluted.
Antennal segments I-III minutely granulate; head granulate, more coarsely in dorsal midline. Pronotum
sparesly punctate posteriorly, granulate anteriorly, tuberculate along lateral and posterolateral margins.
Scutellum, apart from the two prominent tubercles, and thoracic pleura sparsely punctate-granulate.
Thoracic sterna minutely rugose. Legs almost smooth, posterior femora with a few shallow punctures in
apical half. Clavus and corium except for apex evenly punctate, corium with veins anteriorly in basal
half bearing a few large granules. Abdominal sterna and laterotergites neither granulate nor punctate.
Antennae with short to moderately long, suberect, brown pubescence, segment I also with short, ad-
pressed, silvery hairs, head with both of these types of pubescence throughout. Pronotum, scutellum and
AFRICAN POD BUGS 13
Ree
11
12
Figs 9-15 Clavigralla tuberculicollis. (9) head, right lateral view; (10) pronotum, dorsal view; (11)
genital capsule with parameres, posterior view; (12) left metathoracic peritreme, lateral view; (13)
lateral margins of ¢ abdominal sternites III-VII, ventral view; (14) spermatheca; (15) paramere,
dorsal view.
thoracic pleura with moderately long to long, erect, brown hairs and a thin covering of adpressed, silvery
hairs, the latter type only present on thoracic sterna. Legs with short to long, semidecumbent to erect,
pale or dark brown hairs, femora and tibiae, especially near base, with sparse, adpressed, silvery hairs
also. Clavus and corium with short, suberect, pale brown hairs. Abdominal sternites with sparse, moder-
ately long, erect, pale hairs and a thin covering of short, adpressed, silvery hairs; laterotergites with short,
slightly crisped, semidecumbent, pale brown hairs.
Colour generally dark brown to piceous. Antennae pale brown with obscure stramineous rings. Tarsi,
two broad rings on each tibia, some large spots in apical half of each femur and most of basal half of all
femora pale brown to stramineous, legs otherwise piceous. Clavus and corium mid-brown, apex of corium
piceous with small cream spots, apical margin of corium cream with a few piceous spots. Punctures of
clavus and corium piceous. Membrane fuscous with indistinct, paler blotches. Abdominal sterna with
creamy yellow markings which are most extensive on disc of sternites 1V and V and sides of VII. Latero-
tergites IV, V and VI each with a transverse cream-coloured band, VII almost entirely cream.
14 W. R. DOLLING
REMARKS. This species is distinguished from all others with a short, non-deflexed process at the
apex of the antennifer by its large size and dark coloration. It is the only species of its group so
far recorded from Madagascar. The unique holotype, collected in Nossibé, was deposited by
Reuter in the Natiirhistorisches Museum, Liibeck. The entomological collections of this museum
were totally destroyed by bombing and subsequent fire on 28-29 March 1942 (Anonymous, 1947).
DISTRIBUTION. Madagascar.
Figs 16-18 Clavigralla tuberculicollis. (16) aedeagus and basal apparatus, dorsal view; (17) aedeagus,
ventral view; (18) aedeagus and basal apparatus, left lateral view.
MATERIAL EXAMINED
Madagascar: 1 3, 1 92, 1900 (Fairmaire) (MNHN, Paris); 1 3, 3 2°, Diego Suarez, 1893 (Ch. Alluaud)
(MNHN, Paris and BMNH, London); 3 3, Région du Sud, Andrahomana, 1901 (Al/uaud) (MNHN, Paris
and BMNH, London); 1 3, Bezanozano, 1898 (Noualhier) (MNHN, Paris); 1 3, Ambovombe, 1925 (R.
Decary) (BMNH, London); 1 2, Région du Sud de I’Isle, Bekily, viii.1936 (A. Seyrig) (MNHN, Paris).
Clavigralla leroyi (Schouteden) comb. n.
(Figs 19-26)
Acanthomia leroyi Schouteden, 1938 : 290. LECTOTYPE d, Zaire (MRAC, Tervuren), here designated
[examined].
Acanthomia mixta Schouteden, 1938 : 290-291. LECTOTYPE 6, ZatrE (MRAC, Tervuren), here
designated [examined]. Syn. n.
Length: 3, 8-3-9-8 mm, mean 9:0 mm (n = 28); 9, 8:8-10°8 mm, mean 9:5 mm (n = 25).
Antennifers strongly divergent, process at external apical angle well developed, triangular, porrect.
Ratio of lengths of antennal segments I : II : III] : IV in male about 1-00 : 0:92 : 0-76 : 0-91, in female
about 1-00 : 0-92 : 0-78 : 0-85. Length of segment I divided by width of head including eyes in male
1:03-1:25, mean 1-15 (n = 27), in female 1-07—1-44, mean 1:19 (n = 23). Ratio of lengths of rostral seg-
ments I : II : III : IV in both sexes about 1-00 : 0-85 : 0:52 : 0-79.
Pronotum moderately strongly and convexly declivent, disc with a stout, conical tubercle near mid-
point of each anterolateral margin and several other spines and tubercles of similar size, the more
AFRICAN POD BUGS 15
Figs 19-26 Clavigralla leroyi. (19) conjunctiva, left lateral view; (20) pronotum of ¢ lectotype of
mixta, dorsal view; (21) pronotum of ¢ lectotype of /eroyi, dorsal view; (22) lateral margins of 2
abdominal sterna III-VII, ventral view; (23) conjunctiva, dorsal view; (24) spermatheca; (25)
conjunctiva, ventral view; (26) left paramere, dorsal view.
posterior ones being blunt, apically rounded and approximately hemispherical in form. Prescutellar
spines small or absent. Posterolateral angles moderately produced, posterolateral spines variable in length,
thickness and direction, generally slightly anterolaterally directed if longer and thinner (Fig. 20) and
slightly posterolaterally directed if shorter and thicker (Fig. 21), but various intermediates and combina-
tions of features occur. Width of pronotum across apices of posterolateral spines divided by width of head
including eyes in male 2:14-2:65, mean 2:35 (n = 27), in female 2:24-2:95, mean 2:43 (n = 25). Postero-
lateral spines in male more usually directed forwards and in female more usually directed laterally, but
not consistently so in either sex. Scutellum distinctly but rather weakly convex, granulate. Dorsal ridge of
metathoracic scent-gland peritreme bilobed, the lobes subequal in size or the anterior one slightly larger.
Apical margin of corium only very shallowly concave, apex scarcely produced, when at rest attaining only
level of suture between laterotergites V and VI or slightly exceeding this. Anterior and intermediate
femora each with 1 small subapical spine beneath, posterior femur with 2 major subapical spines, the more
basad less than half the length of the other, with or without a small spine or tubercle between them and
16 W. R. DOLLING
with an apical series of 4 spines, very rarely a third, minute, major spine is discernible basad of the sub-
apical group. Posterior tibia straight, about 0-93 times length of posterior femur.
Abdominal sternites III-VII with posterolateral angles prominent as small, triangular spines (Fig. 22).
Male genital capsule with lip shallow, obtusely angled or truncate, not filling posterior emargination,
tongue triangular, entire. Paramere (Fig. 26) with the two teeth on inner side about equally prominent and
forming a slightly acute angle between themselves. Conjunctiva (Figs 19, 23, 25) with dorsomedian lobe
low, simple; intermediate dorsal lobe large, its lateral angles slightly prominent and its apex triangularly
produced posteriad; distal dorsomedian lobe small, bifid; distal dorsolateral lobes deflexed ; posterior face
of conjunctiva with a single, subspherical lobe above the large, paired sclerites protecting the coiled base
of the vesica; apical ventral lobes large, with dorsal surface lightly sclerotized and ornamented with
numerous small warts; distal ventrolateral lobes large; paired ventral lobes present at level of dorsomedian
lobe. Vesica with non-coiled portion about as long as conjunctiva, sclerites protecting its base about equal
in length to straps of ejaculatory reservoir complex. Female sternum VII cleft along distal third of ventral
midline. Spermatheca (Fig. 24) with duct nearest to bulb curving in three-quarters of a circle round to-
wards bulb, then turning back on itself and forming another three-quarters circle closely appressed to
inside of first loop, followed by a small kink and a U-shaped loop; thin-walled portion of duct not quite
half length of bulb.
Head and antennal segment I weakly granulate, antennal segments II and III minutely granulate.
Pronotum bearing granules of a wide range of sizes, disc punctuate. Scutellum with large granules mostly
arranged in two parallel, longitudinal bands. Thoracic pleura granulate-punctuate. Thoracic sterna and
abdominal sternites and laterotergites almost lacking in macrosculpture apart from weak transverse
striations on abdominal sternites. Abdominal tergites I and II transversely striate, III with a group of
about 12 large, often confluent punctures in middle, IV-VII with smaller punctures throughout, decreas-
ing in size and density laterally and posteriorly. Clavus seriately punctate in about four rows, corium
punctate throughout, seriately so basally, veins in basal half granulate. Femora very weakly and tibiae
minutely granulate.
Pubescence all white to silvery, fine. Short, decumbent pubescence present on head, pronotum,
scutellum, thoracic pleura and sterna, abdominal sterna and, sparsely, on femora and antennal segment I,
often organized into a conspicuous white line along midline of posterior lobe of pronotum and scutellum.
Short, semidecumbent pubescence present on abdominal laterotergites, clavus and corium. Rather short,
suberect pubescence present on head, pronotum, scutellum and all antennal segments, rostrum and legs.
Longer, suberect to erect pubescence also present on antennal segment I, dorsum of head, femora, tibiae,
tarsi, pronotum and abdominal sterna.
Colour generally grey-brown, mottled. Dorsum of head, pronotum and scutellum reddish brown; head
with a pair of cream-coloured stripes running along inner border of compound eyes and thence towards
posterior margin of head, slightly convergent posteriorly and usually edged narrowly piceous; ocellar
tubercles often piceous. Larger spines and tubercles of pronotum and often lateral margins and raised
apex of scutellum also piceous. Head laterally and ventrally red-brown but usually largely or almost
entirely covered with piceous markings. Thoracic pleura red-brown with piceous markings; sterna entirely
piceous. Abdominal sterna red-brown with numerous, variable, cream and piceous markings, sterna
III-VII almost always with diagonal piceous and cream markings about half-way between midline and
lateral margins. Laterotergites red-brown, each with a transverse pale cream band occupying the middle
third. Clavus and corium pale grey-brown, corium with a few spots and granules brown, veins at and near
its apical margin partly or largely cream; membrane slightly infuscate, veins brown, interrupted by
unpigmented areas. Antennae pale flavous, often tinged rufous, segments I-III typically each with a line
along dorsolateral surface and an apical annulus not continuous with the line red-brown to piceous or
black, these darker antennal markings sometimes more extensive and sometimes absent. Legs stramineous;
femora heavily marked with irregular, red-brown and frequently also piceous spots, especially in apical
two-thirds; tibiae each with a basal and apical annulus and a median dorsal spot or complete annulus
red-brown or piceous; base and apex of first and second tarsal segments and whole of third tarsal segment
of all legs red-brown.
Egg with 9 aeromicropyles (3 ovarian eggs taken from a single female).
REMARKS. There is considerable but continuous variation in the shape of the pronotum and the
extent of piceous markings, which may be completely absent. These characters vary independ-
ently. The complex form of the conjunctiva and that of the spermatheca, however, show little
variation, suggesting that the material examined comprises a single, variable species rather than
a complex of closely similar species. The type-series of Acanthomia leroyi comprises specimens
with short, laterally directed pronotal posterolateral spines (Fig. 21) and about average intensity
AFRICAN POD BUGS Ey
and extent of the piceous markings while the type-specimens of A. mixta have longer, anterolater-
ally directed pronotal posterolateral spines (Fig. 20) and more extensive piceous markings.
DISTRIBUTION. Central and West Africa, usually at high altitudes.
MATERIAL EXAMINED
Acanthomia leroyi Schouteden, lectotype 3, Zaire: Ituri, Nioka, vii.1934 (J. V. Leroy) (MRAC, Tervuren).
Acanthomia mixta Schouteden, lectotype 3, Zaire: Ituri, Nioka, vii.1934 (J. V. Leroy) (MRAC, Tervuren).
Guinea: 2 °, Mt To, 1660 m, 28.vi.1954 (BMNH, London). Sudan: 1 3, W. Darfur, S. Jebel Murra,
Dimbiti, 7600 ft (2350 m), 28.v.1932 (M. Steele) (BMNH, London). Ethiopia: 1 3, Lake Zwai, v.1914 (O.
Kovacs); 1 3, Maraco, 8.iv.1915 (Kovacs); 1 °, Urgessa, R. Simu, 2300 ft (700 m), 3.v.1905 (Ph. C.
Zaphiro) (BMNH, London). Uganda: 1 2°, Kampala, 15-30.x.1917 (C. C. Gowdey) (BMNH, London).
Burundi: 1 3, Mugera, end of 1965 (J. J. Rwabuneza) (MRAC, Tervuren). Zaire: 1 3, 2 9, Ituri, Nioka,
vii.1934 (J. V. Leroy) (paralectotypes of Acanthomia leroyi Schouteden) (MRAC, Tervuren); 1 3, 29
Ituri, Nioka, vii.1934 (J. V. Leroy) (paralectotypes of Acanthomia mixta Schouteden) (MRAC, Tervuren) ;
1 3, Kinda, N’Zazi, 15.i.1949 (R. P. Lefebvre); 1 3, Lualaba, Jadotville, Kakanda, vii—viii.1953 (R. P. Th.
de Caters); 1 9, Bas-Congo, Lemfu, x—xii.1944 (Rév. P. De Beir); 1 3, Lulua, Kapanga, ii.1933 (F. G.
Overlaet); 3 9, Lulua, Kapanga, 14.xii.1932, i.1933 and iii.1933 (Overlaet); 1 3, Kivu, Mulungu, 1939
(Hendrickx); 1 9, Kivu, Mulungu, Tshibinda, xi.1951 (P. C. Lefévre) (MRAC, Tervuren); 1 9, Lubum-
bashi, 10.iv.1939, at light (H. J. Brédo); 1 2, Kaniama, 8.iv.1939 (Brédo) (IRSNB, Brussels); 35 3, 44 9,
Faradje, 29° 40’ E, 3° 40’ N, i.1913 (Lang & Chapin) (AMNH, New York); 1 3, Katanga, Lufira River,
31.viii.1907, 3500 ft (1050 m) (Neave); 1 3, 1 9, Katanga, Kambove, 4000-5000 ft (1200-1500 m),
23.iv.1907 and 21.vi.1907 (Neave); 1 3, Lualaba River, 24.v.1907, 2500-4000 ft (750-1200 m) (Neave); 1 9,
SE. Katanga, 4000 ft (1200 m), i.xii.1907 (Neave); 1 9, Kando, 9.ix.1944 (BMNH, London), Angola: 1 3,
Duque de Braganca Falls, 11-12.iii.1972, by sweeping (BMNH Southern Africa Expedition) (BMNH,
London). Zambia: 4 3,'1 9, Upper Kalungwisi Valley, 4200 ft (1280 m), 11.ix.1908 (S. A. Neave); 1 d,
Lower Kalungwisi Valley, 3500 ft (1050 m), in dense forest, 15.ix.1908 (Neave); 1 3, Chisinga Plateau,
Kalungwisi District, 4500 ft (1350 m), 23.ix.1908 (Neave); 3 3, 2 2, same locality, 25.ix.1908 (Neave) (UM,
Oxford). Malawi: 1 3, Mlanje Plateau, 6500 ft (2000 m), 18.xii.1913 (S. A. Neave); 1 9, Ft Jameson to
Dowa, 4000-4500 ft (1200-1350 m), 4-9.x.1910 (Neave); 1 9, Kotakota, xi.1910 (¥. E. S. Old) (BMNH,
London).
Clavigralla zambiae sp. n.
(Figs 27-30)
Length: 3, 8-2 mm (n = 1); 9, 8-4mm (n = 1). Similar to C. /eroyi but smaller and body slightly more
depressed. Known from only one specimen of each sex.
Antennifers strongly divergent, external apical processes porrect, triangular. Ratio of lengths of
antennal segments 1 : II : III : 1V in male 1-00 : 0-95 : 0-79 : 0-84, in female 1-00 : 0-97 : 0:74:-(segment
IV missing). Length of segment I divided by width of head including eyes in male 1-08, in female, 1-04. Ratio
of lengths of rostral segments.I : II : I] : IV in male 1-00 :.0-87 : 0-49 : 0-78.
Pronotum (Fig. 27) moderately strongly declivent, posterolateral angles scarcely produced, postero-
lateral spines short, width across tips of spines divided by width of head including eyes in male 1-94, in
female 2-10. Prescutellar spines absent. Dorsal ridge of metathoracic peritreme bilobed.
Abdominal sternites III-VII with posterolateral angles (Fig. 29) much less strongly produced than in
leroyi. Paramere (Fig. 30) with teeth on inner side very unequal in length, apical tooth much the more
prominent of the two. Conjunctiva (which could not be fully inflated) apparently similar to that of leroyi,
having apical ventral lobes large and ornamented with small warts. Vesica with non-coiled portion as
long as expanded conjunctiva and phallotheca together. Spermatheca (Fig. 28) with duct near bulb
following several short convolutions, then with short U-shaped portion.
Sculpture, pubescence and coloration similar to /eroyi.
REMARKS. Distinguished from C. lJeroyi by the generally shorter antennae, narrower pronotum,
longer vesica and different configuration of spermathecal duct.
DIsTRIBUTION. So far known only from Zambia, a country with a large number of endemic species
of very restricted distribution.
MATERIAL EXAMINED
Holotype 2, Zambia: Mpika, 4700 ft (1450 m), 1904 (F. H. Mellard) (BMNH, London).
Paratype. Zambia: 1 3, Abercorn, x.1943 (H. J. Brédo) (IRSNB, Brussels).
18 W. R. DOLLING
31
Figs 27-35 Clavigralla species. (27) zambiae, pronotum of holotype 3, dorsal view; (28) zambiae,
spermatheca; (29) zambiae, lateral margins of abdominal sternites III—-VII, ventral view; (30)
zambiae, paramere of holotype, dorsal view; (31) uelensis, pronotum of 2, dorsal view; (32) uelensis,
spermatheca; (33) uelensis, left metathoracic peritreme, lateral view; (34) uelensis, head, right
lateral view; 35 uelensis, lateral margins of abdominal sternites III-VII, ventral view.
Clavigralla uelensis (Schouteden) comb. n., stat. n.
(Figs 31-35)
Acanthomia mixta var. uelensis Schouteden, 1938 : 291. LECTOTYPE 9, ZAIRE (MRAC, Tervuren), here
designated [examined].
Length: 9, 9-6-9-7 mm, mean 9-6 mm (n = 5). Male unknown. Very similar in size, build and appearance
to C. leroyi.
Antennifers strongly divergent, external apical processes large, triangular, porrect (Fig. 34). Ratio of
lengths of antennal segments I : II : III : IV about 1-00 : 0-88 : 0-81 : 0-87; length of segment I divided
by width of head including eyes 1:10-1:23, mean 1:15 (n = 5). Ratio of lengths of rostral segments I : II :
III : IV about 1-00 : 0-84 : 0:54 : 0-77.
Pronotum (Fig. 31) with posterolateral angles moderately produced, posterolateral spines slender,
directed obliquely forwards; width across apices of spines divided by width of head including eyes 2:27—
2-57, mean 2:40 (n = 5). Metathoracic scent-gland peritreme with dorsal ridge entire, not bilobed (Fig
33). Posterior femur with first subapical spine very short, much smaller than the second, or only one, long,
subapical spine present, not preceded by smaller spines or tubercles and followed by a terminal series of
four spines. Length of posterior tibia divided by that of posterior femur 0:86-0:92, mean 0-88 (n = 3).
AFRICAN POD BUGS 19
Posterolateral angles of abdominal sternites III-VII (Fig. 35) less prominent than those of /eroyi.
Spermatheca (Fig. 32) with duct long and loosely convoluted, thin-walled portion very short.
REMARKS. Differs from all other species by the combination of the porrect external apical process
of the antennifer and the entire, not bilobed, dorsal ridge of the metathoracic scent-gland peri-
treme. Otherwise, in size, shape, sculpture, pubescence and coloration this species is very similar
to Jeroyi, and it was originally described as a variety of mixta, synonymized above with /eroyi.
DISTRIBUTION. Central Africa.
MATERIAL EXAMINED
Acanthomia mixta var. uelensis Schouteden, lectotype °, Zaire: Dungu, xi.1919 (P. van den Plas) (MRAC
Tervuren).
Zaire: 3 2, Dungu, xi.1919 (P. van den Plas) (paratypes of Acanthomia mixta var. uelensis Schouteden)
(MRAC, Tervuren). Central African Republic: 1 2, Bambari, i.1964. (G. Pierrard) (MRAC, Tervuren).
The elongata-group
Antennifer with outer apical process long, deflexed; metathoracic peritreme with dorsal ridge entire; male
genital capsule with tongue entire; lip of genital capsule filling posterior emargination in posterior veiw;
pubescence of anterior, declivent area of pronotum almost uniform with that of posterior, horizontal part.
Second valvula of ovipositor bearing at apex a usually bifid process. Pronotum with posterolateral angles
usually strongly produced, tapering gradually into posterolateral spines.
Clavigralla hystrix Dallas comb. rev.
(Figs 36-42)
Clavigralla hystrix Dallas, 1852 : 512-513. Holotype 3, SIERRA LEONE (BMNH, London) [examined].
Acanthomia hystrix (Dallas) Stal, 1873 : 83.
Length: 3, 7°6-8:7 mm, mean 8:1 mm (n = 28); 9, 8:3-9:8 mm, mean 9:0 mm (n = 43). Body robust,
aspect very spinose, connexivum broad in middle, coloration very dark.
Antennifers rather strongly divergent. Ratio of lengths of antennal segments I : I : III : IV in male
about 1-00 : 0-67 : 0-69 : 0-62, infemale about 1-00 : 0-68 : 0-7 0: 0-61. Segments I and III gently curved.
Length of segment I divided by width of head including eyes in male 1:81-2:26, mean 2-01 (n = 28), in
female 1-72-2-08, mean 1-92 (n = 41). Ratio of lengths of rostral segments I : II : III : IV in bothsexes
about 1-00 : 0-83 : 0:51 : 0-79.
Pronotum (Fig. 36) moderately strongly declivent, posterolateral spines drawn out into very long,
slender spines, with several prominent spines in the vicinity of the posterolateral angles, disc with a semi-
circle of about six spines, the most anterior spine on each side (close to middle of lateral margins) very
much more prominent than the rest, spines at prescutellar angles small. Width of pronotum across apices
of posterolateral spines divided by width of head including eyes in male 2:78-3:68, mean 3-32 (n = 23),
in female 3-10—3-95, mean 3-51 (n = 35). Scutellum strongly convex. Corium with apex slightly produced,
at rest reaching to level of middle of laterotergite V. Anterior and intermediate femora each with one small
subapical spine beneath, or that of intermediate femora absent, or, rarely, anterior femur with a second
subapical spine proximal to and much smaller than the other. Posterior femur with two major spines, the
more proximal about half as long as the other and separated from it by a row of about four small spines,
apical series of four spines present. Posterior tibia gently bowed near base, its length divided by that of
posterior femur in both sexes 0:85-0:97, mean 0:90 (n = 68).
Abdominal sternites III-VII with posterolateral angles produced into long, stout spines (Fig. 37). Male
genital capsule with lip broad, tongue rather short. Paramere (Figs 39, 40) with apical margin weakly
convex to straight and slightly emarginate in middle, varying slightly in the angle it forms with axis of
shaft. Conjunctiva (Fig. 38) with distal ventrolateral lobes large, apical ventral lobes of moderate size,
distal dorsolateral lobes obsolete, dorsomedian and distal dorsomedian lobes low. Female sternite VII
deeply cleft. Second valvula (Fig. 41) with a single, narrow process at apex. Spermatheca (Fig. 42) with
bulb narrow and duct short. Sclerites of dorsal wall of gynatrium (Fig. 42) with median, longitudinal arms
broad, strongly divergent posteriorly.
Head granulate throughout, antennal segments I-III sparsely and minutely granulate. Pronotum in
posterior half and all pleura coarsely punctate, pleura with a few granules or short, blunt spines. Scutellum
very coarsely granulate. Clavus and corium coarsely and rather sparsely punctate, veins in basal two-thirds
20 W. R. DOLLING
Figs 36-42 Clavigralla hystrix. (36) pronotum, dorsal view; (37) lateral margins of abdominal
sternites III-VII, ventral view; (38) conjunctiva and vesica, ventral view; (39) paramere, dorsal
view, Uganda; (40) same, Zaire; (41) left second valvula, medial view; (42) sclerites of dorsal wall
of gynatrium and spermatheca, dorsal view.
of corium granulate. Femora and tibiae almost obsoletely granulate. Thoracic and abdominal sterna
impunctate, abdominal sternite VII shallowly and sparsely rastrate.
Head, antennae, rostrum and legs with short, decumbent, golden hairs and short, semidecumbent to
suberect white hairs, the decumbent pubescence dense on head, sparse elsewhere, semierect pubescence
longer on tibiae and tarsi. Pronotum and scutellum with short, dense, decumbent, golden pubescence and
fairly short, erect, sparse, brown pubescence. Clavus and corium with short, decumbent, golden pubesc-
ence and short, crisped, semidecumbent, white pubescence intermingled. Thoracic pleura, abdominal
sternites and laterotergites with short, dense, decumbent, golden pubescence and short, dense, semi-
decumbent, white pubescence intermingled. Thoracic sterna with decumbent, golden pubescence only.
Colour dark brown to piceous. Antennal segments I-III each with a more or less obscure pale annulus
distad of middle, that on segment II the most distinct of the three. Antennal segment IV ivory-white, not
at all to heavily infuscate. Tibiae each with two pale brown to stramineous annuli, the distal annulus
frequently continuous to apex of tibia. First segment of all tarsi stramineous above except at extreme base
and apex. Posterior femur with a pale annulus basad of middle. Midline of anterior half of pronotum
broadly darker than ground colour. Abdominal sternites and laterotergites with obscurely paler areas.
Corium stramineous, heavily spotted with brown and with a more extensive patch of brown on disc.
Membrane dark brown with scattered white spots and a white crescent in proximal angle. Distal margin
of corium stramineous, forming a large, pale capital A together with the white crescent on membrane
when elytra are at rest.
AFRICAN POD BUGS 21
REMARKS. This species varies in the length of its spines and appendages and to a lesser degree in
colour in different parts of its range. The greatest development of the pronotal spines is found in
the centre of its range, in Zaire. Means of the ratio pronotal width/head width in various parts of
the range of the species are as follows: West Africa (Sierra Leone to Nigeria): 3, 3-25; 9, 3-31;
Cameroun: 4, 3-38; 9, 3-61; Zaire: 3, 3:44; 9, 3-67; East Africa (Uganda and Kenya): 4, not
available; 9, 3-19. The ratio length of antennal segment I/head width decreases from west to east
as follows: West Africa: g, 2:11; 9, 1:99; Cameroun: 4g, 1:97; 9, 1-95; Zaire: J, 1-89; 9, 1-84;
East Africa: 3, 1-87; 92, 1:74. Antennal segments II and III maintain their length relationship with
segment I throughout the range of the species but segment IV in both sexes averages 0-66 times
the length of segment I in Zaire and 0-60 elsewhere. Antennal segment IV is more heavily
infuscate in examples from Zaire and East Africa than in those from Cameroun and West Africa
and the more distal pale annulus of the tibiae is more often continuous to the apex of each tibia
in Zaire and East Africa than in Cameroun and West Africa. Paramere shape does not appear to
vary systematically from place to place. The egg is figured by Cobben (1968 : 97, fig. 85).
DISTRIBUTION. Equatorial Africa, from Sierra Leone to Uganda, in forest areas.
MATERIAL EXAMINED
Clavigralla hystrix Dallas, holotype 3, Sierra Leone (D. J. Morgan) (BMNH, London).
Sierra Leone: 1 3, Njala, viii.1928 (E. Hargreaves) (BMNH, London). Liberia: 1 2, 32 km S. of Voin-
jama, 13.viii.1966 (EZ. S. Ross, K. Lorenzeu) (CAS, San Francisco). Ivory Coast: 1 3, Adiopodoumé,
iv—v.1964 (R. H. Cobben) (BMNH, London); 1 3, Akoupe, 25 km N. of Abidjan, x.1961 (J. Decelle); 1 2,
Akoupe, v.1961 (Decelle); 2 3, 5 2, Mouyassoué, Aboisso, xii.1962 (Decelle) (MRAC, Tervuren). Ghana:
1 2, ‘Owealé Akem, Aschanti-Guinée’ (IRSNB, Brussels); 1 3, 1 9, Aburi (A. R. Gould); 1 9, ENE. of
Tafo, 26.iii.1948, on herbage (BMNH, London); 1 9, Tafo, 14.x.1966, on cocoa, by pyrethrum knock-
down (R. Kumar); 1 2, Adukrom, 14.x.1967 (D. Leston); 1 3,1 2, Kukurantumi, 13.i.1968, on Legumin-
osae (Leston); 3 3, Tafo, 13.v.1966, 3.i.1967, 11.1.1968 (Kumar); 1 3, Mt Atewa, 22.x.1967, in secondary
forest (Leston); 1 3, Adonkwanta, 1.viii.1967 (Leston); 1 3, Volta River Forest Reserve, 5.ix.1965 (Leston)
(UG, Legon). Togo: 1 3, Missahoué, 650 m, vi.1963 (Mme Y. Schach) (MRAC, Tervuren). Nigeria: 1 2,
Old Calab[ar] (J. Gray) (BMNH, London); 1 2, Lagos Colony, Isheri, 9-10.iv.1949 (Borys Malkin); 1 &,
Oyo Prov., Ogbomosho, 12.xii.1948 (Malkin) (CAS, San Francisco). Cameroun: 1 3, 4 2, Lolodorf (G.
Schwab); 4 2, Metet (Schwab) (AMNH, New York); 1 2, Mukonje Farm (F. Rohde) (IRSNB, Brussels);
1 3, 3 2, Nkolbisson, Nyong-Sanaga, x.1963 (L. G. Segers) (MRAC, Tervuren); 1 3, 1 9, Escalera; 2 9°,
Batouri District, 3° 45’ N, 13° 45’ E, 750 m, 1.v—6.vi.1935 (F. G. Merfield) (BMNH, London); 9 3,82,1V
instar nymph, Victoria Div., Mabete, 24.v—7.vi.1949 (B. Malkin); 3 3, Victoria, Mabete, 15.vi.1949
(Malkin); 11 3,10 9, 1 V instar nymph, Victoria Div., Mabeta, vii—viii.1949 (S. Tita); 1 3, Victoria Div.,
Muyuka, 24-29.vi.1949 (B. Malkin); 4 3, 3 2, Sasse, Buea, iv-v.1951 [no collector stated]; 1 9, Sasse,
Sappo, 22-26.ii.1952 (S. Tita); 1 3, 2 2, Sappo, near Buea, iii.1951 (S. Tita); 1 3,2 2, Mamfe, 22—24.1.1949
(B. Malkin) (CAS, San Francisco). Zaire: 1 3, Luhoho River, Bunyakiri, 1100 m, 6.ix.1957 (E. S. Ross,
R. E. Leech) (CAS, San Francisco); 1 3, 1 9, Ituri Forest, 40 miles (64 km) NNE. of Beni, 3000 ft (900 m),
12-10.ix.1959 [sic], in indigenous forest (Cambridge E. African Exped.), 1 2°, Mt Hoyo, 1° 10’ N, 30° 0’ E,
3000 ft (900 m), 14-16.ix.1959, in forest clearings (Cambridge Exped.) (BMNH, London); 1 3, 1 9, Medje,
2° 25’ N, 27° 15’ E, vi.1914 (Lang & Chapin); 1 3, Medje, vii.1910 (Lang & Chapin); 1 9, Kisangani (as
Stanleyville), iii.1915 (Lang & Chapin) (AMNH, New York), 2 3, Bambesa, 17.vi.1937, 26.i.1939 (J.
Vrydagh); 2 2, Bambesa, 29.vi.1937, 20.xii.1939 (Vrydagh); 1 3, Bukolela, 1938 (R. Massart); 3 2, Ngowa,
16.xi.1938 (R. P. J. Mertens); 1 2, Wenga Ifomi (E. Quineaux); 1 2, Kisangani (J. Muller) (IRSNB,
Brussels); 1 3, Kasongo, R. Lumami, ii.1960 (P. L. G. Benoit); 1 3, Kasongo, Mufala, xi.1959-i.1960
(Benoit); 1 3, Equateur, Bokuma, vii.1952 (R. P. Lootens); 1 9, Kivu, Kavumu to Kabunga, km 82,
viii.1951 (H. Bomans); 1 2, Kivu, Lukando (Bunyakiri) 1959-1960 (J. Hecg) (MRAC, Tervuren). Uganda:
1 3, Entebbe, 16.xii.1912 (C. C. Gowdey); 1 2, Entebbe, 12.ix.1912 (Gowdey) (BMNH, London); 1 3, 1 8,
Mulange, xi.1922 (R. Summer) (PPRI, Pretoria). Kenya: 1 2°, Mabira F. (A. F. J. Gedye) (BMNH, London).
Angola: 1 9, Carmona, Faz. Bambi, ii.1972 (J. A. Quarteau) (Duarte coll.).
Clavigralla hystricodes Stal comb. rev.
(Figs 43-49)
Clavigralla hystricodes Stal, 1866 : 109. LECTOTYPE 9, SteRRA LEONE (NR, Stockholm), here designated
[examined].
Acanthomia hystricodes (Stal) Stal, 1873 : 83.
22 W. R. DOLLING
44
NG
Figs 43-49 Clavigralla hystricodes. (43) pronotum, dorsal view; (44) conjunctiva and vesica, left
lateral view; (45) lateral margins of ¢ abdominal sternites III-VII, ventral view; (46) paramere,
dorsal view, Sierra Leone; (47) same, Nigeria; (48) same, Tanzania; (49) sclerites of dorsal wall of
gynatrium, dorsal view.
Length: 3, 8-6-9-8 mm, mean 9:2 mm (n = 21); 9, 9:2-10-8 mm, mean 9:8 mm (n = 37). Body robust,
coloration generally piceous.
Antennifers strongly divergent. Ratio of lengths of antennal segments I : II : III : IV in male about
1-00 : 0-76 : 0-80 : 0-75, in 2 about 1-00 : 0:76 : 0-79 : 0:73, segments II and III straight. Length of segment
1 divided by width of head including eyes in male 1:25-1-49, mean 1:36 (n = 23), in female 1-18-1-46,
mean 1-33 (n = 37). Ratio of lengths of rostral segments I : II : III : 1V in both sexes about 1:00: 0-83:
0-62 : 0-90.
Pronotum (Fig. 43) strongly declivent, spines of disc short, broad, irregularly distributed. Width of
pronotum across apices of posterolateral spines divided by width of head including eyes in male 2:63-3:07,
mean 2:88 (n = 15), in female 2:63-3:25, mean 2:97 (n = 31). Scutellum strongly convex. Spines on
femora as in C. hystrix except that anterior femur usually bears two subapical spines (usually one in
hystrix). Posterior tibia weakly angled, rather than curved, near base, its length divided by that of posterior
femur in both sexes 0:74-0:84, mean 0-80 (n = 57). Corium with apex produced, at rest reaching to level
of suture between laterotergites V and VI.
AFRICAN POD BUGS 23
Abdominal sternites III-VII with posterolateral angles strongly produced (Fig. 45). Male genital
capsule with lip moderately broad, tongue rather narrow. Paramere (Figs 46-48) with apical margin
weakly to rather strongly convex, making a variable angle with axis of shaft. Conjunctiva (Fig. 44)
similar to that of C. hystrix but with distal dorsomedian lobe larger. Female sternite VII deeply cleft.
Spermatheca with bulb narrow, duct short. Sclerites of wall of gynatrium (Fig. 49) slender, apically
slightly convergent. Second valvula with apex broad, bearing apically a bifid process similar to that of C.
elongata (cf. Fig. 57).
Sculpture and pubescence as in C. hystrix except that thoracic pleura lack spines and large granules.
Colour piceous, with pale brown or stramineous markings. Head with a diagonal pale line running from
inner dorsal angle of each eye towards posterior margin of head behind ocellus. Antennal segments I,
II and III each with a conspicuous pale stramineous annulus at base and another about two-thirds of the
way towards apex, IV brown with basal annulus pale stramineous. Pronotum with granules along midline
of declivent area black. Thoracic pleura reddish piceous, with black and pale rufous patches. Clavus and
corium pale brown, corium with some spots near anterior margin and a large area near apical margin
piceous to black, a line along posterior two-thirds of apical margin and some large spots on and near
anterior margin stramineous. Membrane dark brown with extensive piceous area on disc and numerous
obscure paler spots. Connexivum piceous, laterotergites III-VII each with a narrow, transverse band in
anterior half stramineous. Most lateral one-quarter of abdominal sternites on each side dark, rufous
brown with a stramineous spot on each margin adjacent to the pale band on the corresponding latero-
tergite, remaining area of each sternite stramineous with confused, rufous brown and piceous markings.
Femora dark rufous brown with extensive area near base and some mottling near apex stramineous.
Tibiae piceous, each with a distinct, stramineous annulus in basal and apical half.
Remarks. West African (Sierra Leone to Cameroun) individuals were found on average to have
longer pronotal spines and a longer antennal segment I than the Central and East African in-
dividuals. The ratio width of pronotum/width of head in West African individuals was: male,
2-92-3-07, mean 3-00 (n = 5); female, 2:93-3:31, mean 3-02 (n = 7) and the ratio length of antennal
segment I/head width was: male, 1:37-1:49, mean 1-44 (n = 5); female, 1:34-1-46, mean 1-40
(n = 8). The single female seen from South Africa was the largest at 10-75 mm in length (next
longest specimen 10-4 mm) and had the lowest value, 1-18 for the ratio length of antennal segment
I/head width (next lowest value 1-20 in individuals from Malawi and Zaire). In males from West
and Central Africa the angle between the apical and lateral margins of the paramere is more
narrowly rounded (Figs 46, 47) than in individuals from East Africa (Fig. 48). This does not of
itself seem to justify taxonomic separation of the populations and no males were available from
the most southerly part of the range or from Uganda, so that at present it is not possible to
interpret the pattern of variation in full. Feeds on pulse crops in East Africa (Bohlen, 1973 : 95).
A coloured figure of the adult is given by Bohlen (1973 : 33, fig. 159).
DISTRIBUTION. Tropical Africa, extending westward to Sierra Leone, eastward to Uganda and
Tanzania and southward to the Transvaal.
MATERIAL EXAMINED
Clavigralla hystricodes Stal, lectotype 2, Sierra Leone (no other data) (NR, Stockholm).
Sierra Leone: 1 3, Njala, 15.x.1960, on Homalium angustifolium Smith (Flacourtiaceae) (M. P. Rushton)
(BMNH, London). Ivory Coast: 1 9, Daloa, Zepreghé, i.1963 (J. Decelle) (MRAC, Tervuren). Ghana:
3 3, Tafo, 14.iv.1966, 23.iv.1967, 13.ii.1967, the first two by pyrethrum knockdown from cocoa (R.
Kumar); 1 °, Kade, cocoa farm, 22.viii.1969 (G. Benson) (UG, Legon). Nigeria: 2 3, 2 9, Calabar,
3-6.xi.1955 (Bechyne, Exped. Mus. G. Frey, Nigeria-Kamerun) (BMNH, London); 1 9, Ibadan, 22.iii.1949
(B. Malkin) (CAS, San Francisco). Cameroun: 2 2, Nyong-Sanaga, Nkolbisson, x.1963 (L. G. Segers)
(MRAC, Tervuren). Zaire: 1 2, Lubumbashi, 11.ix.1931 (Miss A. Mackie) (BMNH, London); 46 3, 39 9,
i ex without abdomen, Faradje, 3° 40’ N, 29° 40’ E, i.1913 (Lang & Chapin) (AMNH, New York); 2 d,
Kasongo Terr., Lumani River, ii.1960 (P. L. G. Benoit); 1 3, Eala, 29.x.1938 (G. Couteaux); 1 2, Equateur,
Bokuma, vii.1952 (R. P. Lootens); 1 °, Sankuru, M’Pemba Zeo (Gandajika), 1960 (R. Maréchal) (MRAC,
Tervuren); 2 ¢, Ngowa, 9.v.1939 (R. P. J. Mertens); 6 2, Ngowa, 7-27.iv.1939, 5—16.vi.1939 (Mertens);
2 3, 2 2, Mpese, ii-iv.1937, v-vi.1937 (R. P. J. Cooreman); 2 3,1 2, Kibangula, ii.1955, 1956 (L. Henry);
1 3, Malongi, vi.1943 (H.J. Brédo); 1 2, Wenga Ifomi (E. Quineaux); 1 2, Kongolo (Mad. Passau); 1 °,
Botuna, Bokungu, 1950 (M. Boel); 1 2, Libenge, 12.x.1947 (R. Cremer, M. Neuman) (IRSNB, Brussels).
Uganda: 1 9, Entebbe, x.1912 (C. C. Gowdey) (BMNH, London). Tanzania: 1 2, Morogoro, 20.vi.1922,
on legumes (A. H. Ritchie); 3 3, 1 2, Morogoro, 7-8.v.1973, on cow pea (Vigna catjang Endlicher); 1 3,
24 W. R. DOLLING
Figs 50-57 Clavigralla elongata. (50) head and pronotum, dorsal view; (51) head, right lateral view;
(52) lateral margins of 2 abdominal sternites III-VII, ventral view; (53) scutellum, left lateral view;
(54) left metathoracic peritreme, lateral view; (55) abdominal tergites III and IV, 2, South Africa;
(56) same, 9, Uganda; (57) left second valvula of ovipositor, lateral view.
Ilonga, 12.vi.1968, on grams (Phaseolus) (C.J. Thomas); 1 2, longa, 26.ii.1968, on cow pea (Thomas); 1 2,
Tlonga, 24.iv.1968, on cow pea (Thomas) (BMNH, London). Malawi: 2 9, vii-viii.1895 (A. Whyte)
(BMNH, London). Rhodesia: 1 2, Cashel, xii.1947 (NM, Bulawayo). South Africa: 1 2, Transvaal,
Marieps Mountain, iv.1932 (G. van Son) (TM, Pretoria).
Clavigralla elongata Signoret sp. rev.
(Figs 50-65)
[Zicca horrida (Germar) Herrich-Schaeffer, 1851 : 271, pl. 320, fig. 991. Misidentification.]
Clavigralla elongata Signoret, 1860 : 944. LECTOTYPE 3, TANZANIA (NM, Vienna), here designated
[examined]. [Synonymized with Clavigralla horrida (Germar) sensu Herrich-Schaeffer by Stal,
1866 : 108.]
Clavigralla flavipennis Signoret, 1860 : 945. LECTOTYPE 2, MADAGASCAR (NM, Vienna), here design-
ated [examined]. [Synonymized with Clavigralla horrida (Germar) sensu Herrich-Schaeffer by Stal,
1866 : 108.]
AFRICAN POD BUGS 25
[Clavigralla horrida (Germar) Stal, 1866 : 108. Misidentification.]
[Acanthomia horrida (Germar) Stal, 1873 : 83. Misidentification.]
Length: 3, 8:3-9-6 mm, mean 9:1 mm (n = 23); 2, 8:3-10-8 mm, mean 9-6 mm (n = 26). Form elongate-
oblong, aspect spinose, coloration greyish yellow.
Head (Figs 50, 51) about as long as wide, antennifers moderately divergent. Ratio of lengths of antennal
segments I : II : III : 1Vin male about 1-00 : 0-85 : 0-82 : 0-81, in female about 1-00 : 0-85 : 0-0-82 : 0-80.
Length of antennal segment I divided by width of head including eyes in male 1:29-1:72, mean 1-49
(n = 23), in female 1:18-1:59, mean 1-42 (n = 24). Ratio of lengths of rostral segments I : II : III : IV in
male about 1-00 : 0-90 : 0-58 : 0-89, in female about 1-00 : 0-87 : 0°58 : 0:88.
58 a9
Figs 58-62 Clavigralla elongata. (58) left paramere, dorsal view, Tanzania; (59) same, Madagascar;
(60) phallotheca, expanded conjunctiva and vesica, left lateral view; (61) same, dorsal view; (62)
lip of 3 genital capsule, posteroventral view.
Pronotum (Fig. 50) strongly declivent, disc typically with an often irregular semicircle of about six
large, spine-like tubercles, the anterior pair, situated close to mid-point of lateral margins, being only
slightly longer and stouter than the pair behind them. Posterolateral angles of pronotum strongly pro-
duced, slightly elevated and directed slightly anteriorly, each terminating in a slender, very gently curved
spine. Width of pronotum across apices of posterolateral spines divided by width of head including eyes
rather variable between populations, in male 2:56-3:27, mean 2°84 (n = 23), in female 2:57-3:27, mean
2-91 (n = 25). Scutellum (Fig. 53) strongly convex, without outstanding spines or tubercles. Dorsal ridge
of metathoracic peritreme (Fig. 54) entire, reniform. Corium with apex narrowly produced, at rest reach-
ing posteriorly to about level of apex of laterotergite VI. Anterior and intermediate femora each with a
single, moderate sized, subapical spine ventrally, posterior femur typically with two major spines with four
minor spines between them and an apical series of four spines, rarely with additional major spines proxi-
mal to the main pair, number of small spines between main pair rather variable. Posterior tibia almost
straight, its length divided by that of posterior femur in both sexes 0-76—-0-88, mean 0-83 (n = 49).
Abdominal sternites III-VII (Fig. 52) each with a slender spine at posterolateral angles directed
obliquely posteriad. Male genital capsule with lip (Fig. 62) narrowly rounded, tongue acutely pointed,
entire. Paramere (Figs 58, 59) with blade rather broad, apex with a small tooth, basal tooth pointing back
along shaft. Conjunctiva (Figs 60, 61) without ventral or distal dorsolateral lobes, dorsomedian lobe large,
distal dorsomedian lobe small with narrow, distally projecting, finger-like, apical process, ventral apical
and distal ventrolateral lobes well developed. Vesica long, left sclerite protecting base of vesica slightly
longer than right, apices of both sclerites simple. Wings and straps of ejaculatory reservoir complex long.
26 W. R. DOLLING
Female sternum VII cleft for about one-third of its median length. Spermatheca (Fig. 63) typically with
bulb narrow, duct convoluted for a distance equal to about twice the length of bulb, then simply curved
for a distance about equal to length of bulb, then wider and thinner walled for about the same distance
before entering gynatrial sac. Sclerites of wall of gynatrium with apices of median arms abruptly diverging,
usually slender (Fig. 63). Apex of second valvula (Fig. 57) bearing a bifid process.
Figs 63-65 Clavigralla elongata. (63) spermatheca and sclerites of dorsal wall of gynatrium, dorsal
view, South Africa; (64) same, Canary Is.; (65) same, Cape Verde Is.
Head weakly granulate throughout, more strongly on frons. Antennae with segment I minutely and
sparsely granulate, other segments almost smooth. Pronotum with posterior half of disc punctate, lateral
margins and area around posterolateral angles granulate to tuberculate. Scutellum granulate. Thoracic
pleura punctate and very sparsely granulate, thoracic sterna minutely roughened. Femora weakly and
sparsely granulate, very obscurely punctate in apical half, tibiae minutely granulate. Clavus and much of
corium strongly punctate, disc of corium more weakly punctate, apex impunctate. Abdominal sternites
and laterotergites minutely rough, lacking macrosculpture. Abdominal tergites I and II transversely
coarsely striate; III and IV with a few large pores, very few pores of intermediate size and some small or
minute ones, the large pores restricted to the midline of tergite III and its anterior and posterior borders
and sometimes also to midline of segment IV (Fig. 55), from which they are often entirely lacking (Fig.
56), if present on segment IV then distributed along its whole length and concentrated near anterior
margin; puncturation of remaining abdominal tergites small or minute.
Head with short, decumbent, silvery pubescence and a few, suberect colourless or pale brown hairs, the
latter restricted to dorsal surface, especially frons. All antennal segments with very short, decumbent,
silvery much sparser pubescence, on segments II and III than on I and lacking on specialized sensory area
of IV, and short, suberect, pale hairs. Pronotum with silvery, decumbent pubescence anteriorly and very
sparse, decumbent, golden pubescence posteriorly, with moderately long, pale brown to colourless, erect
pubescence throughout. Scutellum with short, decumbent, silvery, tomentose pubescence mixed with long,
erect, pale brown pubescence which becomes paler towards lateral margins. Thoracic pleura with silvery,
decumbent pubescence and longer, sparse, semierect colourless hairs; thoracic sterna with short, decumb-
ent, silvery white hairs. Legs with short, pale brown, suberect hairs and short, silvery, decumbent hairs
throughout, the latter type denser on femora which also bear a few long, erect hairs ventrally. Clavus and
corium with short, semidecumbent, pale brown to colourless pubescence and sparse, scarcely visible,
decumbent, silvery pubescence. Abdominal sterna with short, suberect to semidecumbent, colourless hairs
and decumbent, silvery pubescence. Decumbent, silvery pubescence of thoracic pleura and outer thirds of
abdominal sterna organized in some places into longitudinal or oblique streaks of denser, tomentose or
slightly tufted hairs.
Head, thorax and femora slightly ferrugineous brown, with ventral surface of head, thoracic sterna and
apices of pronotal spines piceous. Antennae stramineous, segments I and IV darker. Tibiae stramineous,
each with a narrow, basal annulus and sometimes a similar, apical annulus brown. Clavus, corium, tarsi
AFRICAN POD BUGS ZI
and a few spots on apical halves of femora stramineous with a slightly greyish tinge. Membrane of
hemelytra colourless, narrowly pigmented brown along course of veins, pigment interrupted in places.
Abdominal sternites ferrugineous brown with middle, especially of more anterior ones, more or less
broadly piceous and a slightly oblique line on each side about one-third of the way from lateral margin to
midline stramineous; a stramineous spot also present on lateral margin of each. Laterotergites ferrugine-
ous brown, III-VII each with a large, stramineous spot adjacent to the similarly coloured spot on lateral
margin of corresponding sternite. Abdominal tergites I and II piceous, III-VII orange, III and lateral
areas of IV-VI more or less heavily infuscate to piceous, centre of VI and VII occasionally with piceous
markings.
REMARKS. There is considerable variation in the degree of puncturation of the abdominal terga
and in the number of spines on the pronotal disc but the general pattern of both punctures and
spines is characteristic. There are isolated populations of this species in the Canary Islands and
the Cape Verde Islands. Morphometric data for specimens from these areas are not included in
the description above. On average, the antennae are shorter than in the mainland population. In
two females seen from the Canary Islands the length of antennal segment I divided by the width
of the head including eyes was 1-18 and 1-30. In two males from the Cape Verde Islands this ratio
was 1:37 and 1-43 and in four females from the same area it ranged from 1-31 to 1:39, mean
1-36. In females from both island groups the convoluted part of the duct of the spermatheca is
shorter than in the mainland population (Figs 64, 65; compare Fig. 63). In both females from the
Canary Islands the sclerites of the dorsal wall of the gynatrium (Fig. 64) were unusually short and
much thicker apically than in the mainland form (Fig. 63), but this was not true of specimens from
the Cape Verde Islands (Fig. 65). Many specimens from the Cape Verde Islands bear three major
subapical spines on the posterior femur. Such a specimen is figured by Lindberg (1958 : 33) who
records the species from the following islands in the group: Santo Antaéo, Sao Vecente, Sado
Nicolau, SAo Tiago, Fogo, Brava and Ilheus de Rombo. Because these populations differ some-
what from the mainland form recent introduction seems unlikely. The nearest localities to these
islands where C. elongata is now found are Sio Tomé, Principe and Zaire. Perhaps the species was
once present in the intervening area of West Africa and has now been displaced by C. shadabi
which appears from the recorded host-plant data to occupy a similar niche. The form from the
Canary Islands may have been isolated long enough to have acquired specific status, to judge
from the differences in the sclerites of the wall of the gynatrium when compared with material
from other parts of the range of C. elongata. Unfortunately, no male was available from the
Canary Islands and the question of the status of this population must remain unresolved for the
present.
This is the species known to East African agricultural entomologists as Acanthomia horrida. It
is a pest of many cultivated Leguminosae, in particular Dolichos lablab Linnaeus, Cajanus cajan
Linnaeus (Millspaugh), Phaseolus spp. and Vigna spp. Natural enemies include Mormonomyia
argentifrons Walker, a tachinid fly parasitic on the adult (Materu, 1971 : 376) and guineafowl. A
scelionid (Hymenoptera), Hadronotus gnidus Nixon, parasitized a low percentage of eggs in the
laboratory (Materu, 1971 : 380). None of these appears likely to cause a significant degree of
mortality in the field. Materu (1970; 1971) describes the damage caused by this species and gives
an account of its biology in Tanzania. The same author (Materu, 1972) describes and figures egg,
nymph and adult. Accounts of control measures are given by Burnett, Lee & Park (1966), Materu
& Makusi (1972; 1973) and Swaine (1969). Bohlen (1973 : 33, fig. 157) gives a coloured figure of
the adult.
DIsTRIBUTION. Much of central, eastern and southern Africa (but not the extreme south), Mada-
gascar, Seychelles, Réunion, Mauritius, Rodriguez, Sio Tomé, Principe, Canary Islands, Cape
Verde Islands, Yemen; recorded by Chiaromonte (1934) from Somalia; absent from the mainland
of West Africa.
MATERIAL EXAMINED
Clavigralla elongata Signoret, lectotype 3, Tanzania: Zanzibar (NM, Vienna). Clavigralla flavipennis
Signoret, lectotype 2°, Madagascar (NM, Vienna).
28 W. R. DOLLING
Numerous specimens from the following localities. Canary Islands. Cape Verde Islands: Sao Nicolau,
Ribiera do Recanto. Yemen: Hodeida, Wadi Bohl. Ethiopia: Mt Chilalo. Sao Tomé: Vallée Ribiera
Palma. Principe. Zaire: Kongo Central, Banana; Kongo Central, Mpese; Bandundu, Popokabaka;
Bandundu, Ngowa (near Popokabaka); Kasai Orientale, Sankuru, Gandajika; Orientale, Faradje; Kivu,
Lwiro; Kivu, Uvira; Kivu, Butembo; Kivu, Terr. Beni, M. Bau; Kivu, Plaine de la Ruzizi; Kivu, Kas-
ongo; Lulaba River; Katanga, Kambove; Katanga, Lubumbashi; Katanga, Moliro. Rwanda: Rubona.
Burundi: Bururi. Uganda: Mabira Forest; Jinja; Entebbe; Kampala; Koki Country, SW. Buddu; Mt
Kokanjero, SW. of Elgon; Valley of Kafu River, Unyoro; Mpanga Forest, Toro; Maramas District,
Ilala, 22 km E. of Mumias; near Masindi; S. Lake Albert; Yingo; Namanve; Bugoma; Masindi. Kenya:
Mombasa, 19 km NW. of Mazera; Mombasa, 16 km inland, Changamwe; Mombasa; Kwale; Langata;
Teita Hills; Teita Hills, Wundanyi; Rabai; Kaimosi; Lake Jipe; Nyeri; Tumutumu; Bwamba Valley;
Naivasha; Diani Beach; Nairobi. Tanzania: Zanzibar; Mt Chala; Biharamulo; Ukiriguru; E. Rukwa;
Rukwa Rift; Kafukola; Dar-es-Salaam; Bukoba. Angola: 19km SW. of Luimbale; 6km SW. of
Quirimbo. Zambia: Lake Bangweolo, Chishi Island; Upper Luangwa River; between Ft Jamieson and
Lundazi; Lake Bangweulu, Mbawala. Malawi: Nkata Bay; Zomba; between Ft Maguire and Ft Johnston,
SE. shore of Lake Nyasa; Mlanje Plateau; Mlanje. Mozambique: Ruo Valley; valley of Kola River, near
E. Mt Chiperone; Serra da Gorongosa; Lower Zambesi River, Luambo. Rhodesia: Umtali; Mashonaland.
South Africa: Transvaal, Elandshoek; Tvl, Louis Trichard; Tvl, Karino; Natal, Malvern; Natal, Pine-
town; Natal, Durban; Natal, Verulam; Natal, Tugela River, near Weenen; Natal, Umbilo; Natal,
Isipingo; Natal, Kloof; Natal, Port Shepstone; Natal, Umkomaas River; Natal, Port Natal; Natal,
Pondoland, Port St Johns; Natal, 32 km S. of Durban, Illova River Mouth; Natal, Umtentweni; Natal,
Eshowe, Natal, Umkomaas; Natal, Umhlali; locality uncertain, ‘C. Bon. Spei’. Seychelles: Praslin; Mahé,
Silhouette. Madagascar: Lac Alaotra; Centre Ferme de Nanisana; Baie d’Antongil; Fénérive, Soaniérana;
Diego-Suarez; Rég. Sud, Bekily; Amparafaravola (E. of Lake Alaotra); Rég. Sud, Andrahomana;
Antanambé; South-east, Plaines d’Ambolisatra. Réunion: Bretagne. Mauritius: Vacoas. Rodriguez.
Altitudes, recorded on labels: 150-2000 m, mostly in the range 1000-1700 m, one locality at
2600 m in Ethiopia.
Host-plants recorded on labels: ‘Canadian Wonder’ beans (Vicia faba Linnaeus), cowpea
(Vigna sp.) and ‘C. procera’ in Tanzania, Soja (Glycine) in Madagascar and Cajanus indicus
Sprenger in Réunion.
Other habitat and biological information on labels: in forest (twice); beside lake; among lake
shore vegetation; “‘cocotiers morts, rid. forét’; ex crop of guinea fowl (twice).
Depositories of material: IRSNB, Brussels; NM, Bulawayo; Duarte coll.; BMNH, London;
AMNH, New York; UM, Oxford; MNHN, Paris; PPRI, Pretoria; TM, Pretoria; CAS, San
Francisco; MRAC, Tervuren; Slater coll.
Clavigralla shadabi sp. n.
(Figs 66-71)
Length: 3, 7-9-9-8 mm, mean 8-9 mm (n = 18); 9, 8-5-10°5 mm, mean 9:7 mm (n = 27). Very similar in
overall appearance to C. elongata.
Ratio of lengths of antennal segments I : II : III : IV in male about 1-00 : 0-81 : 0-81 : 0-77, in female
about 1-00 : 0-81 : 0-80 : 0-74. Length of antennal segment I divided by width of head including eyes in
male 1-49-1-67, mean 1:57 (n = 18), in female 1-39-1-62, mean 1:50 (n = 26). Ratio of lengths of rostral
segments I : II : III : 1V in male about 1-00 : 0-92 : 0:62 : 0-96, in female about 1:00 : 0-90 : 0-60 : 0-94.
Pronotum (Fig. 66) with posterolateral angles in general more strongly produced, slightly more anteri-
orly directed and narrower than in C. elongata; disc with a semicircle of spines distributed as in elongata
but the anterior pair much more prominent than the rest; width across apices of posterolateral spines
divided by width of head including eyes in male 2:73-3:52, mean 3-03 (n = 18), in female 2:67-3:59, mean
3-03 (n = 25). Posterior tibia almost straight, its length divided by length of posterior femur 0-71-0-92,
mean 0°81 (n = 43).
Male genital capsule with lip (Fig. 67) broad, paramere (Fig. 68) with prominent ridge on dorsomedial
face. Conjunctiva (Fig. 70) differs from that of elongata and its close allies in that well-developed,
membraneous distal dorsolateral lobes are present and apex of distal dorsomedian lobe is narrow and
directed anteriorly. Vesica rather short, sclerites protecting its base both with outer apical angle at least
right-angled and usually, in either or both, acutely projecting laterad. Sclerites of dorsal wall of gynatrium
AFRICAN POD BUGS 29
(Fig. 69) slender, with a subterminal expansion on median arm of each. Duct of spermatheca (Fig. 69)
with convoluted portion short, broad, thin-walled section communicating with gynatrium very short.
Abdominal terga III and IV with a patch of punctures common to both, size of punctures greatest in
middle of segment III, decreasing fairly uniformly in all directions away from this area (Fig. 71). Scutellum
frequently with pale, silvery white hairs organized into a distinct, narrow, longitudinal median line;
pronotum with obscure traces of three longitudinal lines of similar pubescence. Sculpture, pubescence and
coloration otherwise very similar to those of C. elongata.
68
70 7
Figs 66-71 Clavigralla shadabi. (66) head and pronotum, dorsal view; (67) lip of 3 genital capsule,
posteroventral view; (68) paramere, dorsal view; (69) spermatheca and sclerites of dorsal wall of
gynatrium, dorsal view; (70) aedeagus, dorsal view; (71) abdominal tergites III and IV.
REMARKS. This is the insect known to West African entomologists as Acanthomia horrida. Under
this name it was recorded as a pest of cowpea (Vigna) by Booker (1965), Taylor & Omoniyi (1972)
and Aina (1972). The last-named author recorded an unnamed ectoparasitic mite and Rhinocoris
bicolor (Fabricius), a predaceous reduviid bug, as natural enemies. The range of this species
overlaps with that of C. elongata in a large area of Zaire. The two species may be readily dis-
tinguished by comparison of the sizes of the spines on the disc of the pronotum. The length and
thickness of the prominent posterolateral angles of the pronotum vary markedly between series
from different localities. For example, these spine-like processes are much longer and thinner in
specimens from Nero Mer, Ivory Coast and Bambesa, Zaire than they are in specimens from
Lamto, Ivory Coast and Faradje, Zaire. There is marked variation between individuals in the
shape of the apices of the sclerites protecting the base of the vesica and also in the length of the
posterior tibia. None of these variations appears to be linked with any other, nor do they vary
systematically with large-scale geographical range.
30 W. R. DOLLING
DISTRIBUTION. West and Central Africa, extending north-eastwards to S. Sudan. Not recorded to
the east of the Great Rift Valley or south of latitude 12° S.
MATERIAL EXAMINED
Holotype 3, Nigeria: Samaru, ix.1961, on cowpea (Vigna catjang Endlicher) (Dept of Agric, N. Nigeria)
(BMNH, London).
Paratypes. Sudan: 1 2, Blue Nile, Abu Hashim to Galegu, 23-24.xi.1962 (R. Linnavuori) (BMNH,
London). Sierra Leone [?]: 1 2, ‘Sierra. Haw. W.’ (UM, Oxford). Ivory Coast: 2 3, 8 2, Toumodi, Lamto,
30.iii, 3.vii, 4.xii.1962, 2.1, 4.vi, 11.vi.1963, 21.vii.1964 (D. Gillon); 4 3, 4 9, Nero-mer, 30.x.1963 (Gillon)
(BMNH, London). Ghana: 1 3, Tafo, 27.xi.1962 on Asclepias (B. Gerard) (BMNH, London). Togo: 1 3,
2 2, Yoh, 12.vi.1963 (Mme Y.Schach) (MRAC, Tervuren). Nigeria: 1 3, 4 2, Bida, weeds on fallow land,
22-23.ix.1955 (M. G. Emsley); 1 3, no exact locality, on leaf of Phaseolus aureus, 22.viii.1953 (Emsley);
1 2, Ibadan, on cowpea (Vigna), 17.vi.1962 (T. A. Taylor); 1 3, Shendam, at light, 26.ix.1955 (Emsley);
1 3, 1 2, Akpashe District, Udi, 28.x.1955 (Bechyne) (BMNH, London). Cameroun: 1 2, Batouri District,
savannah country, 1.i-24.ii.1936 (F. G. Merfield) (BMNH, London). Zaire: 72 3, 90 2, Orientale, Faradje,
29° 40’ E, 3° 40’ N, i.1913 (Lang & Chapin) (AMNH, New York); 5 3, 6 2, Orientale, Bambesa, 1937-1938
(J. Vrydagh); 1 3, Kivu, Kasongo, ix.1959 (P. L. G. Benoit); 1 9, Katanga, Kiambi, Manono, Riv. Luvua,
x.1956 (N. Leleup) (MRAC, Tervuren); 1 3, Katanga, Nyunzu, 1955 (L. Henry); 1 3,2 2, Kivu, Kibangula,
11.1955, 1956 (Henry); 2 3, 2 2, Katanga, Lubumbashi, i.1939, 9.iii.1939 (H. Brédo); 1 2, Kongo Central,
Mpese, v—vi.1937 (R. P.J. Cooreman); 16 3, 10 2, Orientale, Bambesa, 11-12.vi, 13.xi.1937, 18-21.xii.1939,
6-8.1.1940 (J. Vrydagh); 7 3, 17 9, Bandundu, Ngowa (near Popokabaka), 16-17.ii.1938, 16.xi.1938,
17.vi.1939, 2.vii.1939 (R. P. J. Mertens) (IRSNB, Brussels).
Material excluded from type-series. Ghana: 1 2, Mt Atewa, 15.xii.1967 (D. Leston); 1 2, Oyoko,
24.xi.1966 (Leston); 2 2, Legon, 2.x.1968 (R. Kumar); 1 °, Tafo, 20.iv.1966, by pyrethrum knockdown
from cocoa; 1 3, 3 9, Tafo, 13.v.1966, 14.v.1966, 3.i.1967 (Kumar) (UG, Legon). Nigeria: 3 3, 6 9, Zaria,
Samaru, all on cowpea (Vigna), ix.1961, 31.viii.1966, 14.ix.1966 (J. C. Deeming) (IAR, Samaru). (Received
after description was prepared.)
Clavigralla breviceps sp. n.
(Figs 72-76)
Length: 3, 7-5-8-3 mm, mean 7:8 mm (n = 4); 2, 7:8-8:3 mm, mean 8:1 mm (n = 5). Closely resembles
C. elongata in body form but smaller.
Head short, broad, eyes prominent (Fig. 72). Ratio of lengths of antennal segments I : II : III : IV in
male about 1-00 : 0-83 : 0-82 : 0-77, in female about 1-00 : 0-81 : 0-81 : 0-77. Length of antennal segment I
divided by width of head including eyes in male 1:16-1:24, mean 1-21 (n = 4), in female 1-11-1-20, mean
1-15 (n = 5). Ratio of lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0-88 : 0-49 :
0-82.
Pronotum (Fig. 72) with a pair of stout, blunt spines near anterolateral margins of disc with a few,
blunt spines behind them. Width of pronotum across apices of posterolateral spines divided by width of
head including eyes in male 2:67-2:91, mean 2°80 (n = 4), in female 2:74-2:95, mean 2-82 (n = 4).
Length of posterior tibia divided by length of posterior femur 0-71-0-80, mean 0-75 (n = 9).
Male genital capsule with lip (Fig. 75) broad, paramere (Fig. 73) broader at apex in proportion to length
compared to that of C. elongata. Conjunctiva as in elongata. Sclerites of dorsal wall of gynatrium (Fig. 74)
slender, duct of spermatheca (Fig. 74) very short.
Abdominal tergites III and IV (Fig. 76) with numerous, large and medium sized punctures over most of
their width. Tibiae with brown annuli at base, apex and middle, tarsi brown except for most of dorsal
surface of first segment. Sculpture, pubescence and coloration otherwise similar to that of elongata.
REMARKS. The smallest of the species closely allied to C. elongata and readily distinguished from
its relatives by the short head and the colour of the tibiae and tarsi.
DISTRIBUTION. West Africa and Zaire, not common.
MATERIAL EXAMINED
Holotype 3, Nigeria: near Zaria, Samaru, at grass roots, 12.ix.1954 (M. G. Emsley) (BMNH, London).
Paratypes. Nigeria: 1 ¢, Samaru, in mercury vapour light-trap, 1—-8.viii.1970 (P. H. Ward); 1 9, swept
from grasses and flowers, 30—31.vii.1970 (Ward) (BMNH, London). Zaire: 2 3, 4 9, 1 ex. without
abdomen, Orientale, Faradje, 3° 40’ N, 29° 40’ E, i.1913 (Lang & Chapin) (AMNH, New York and
BMNH, London).
AFRICAN POD BUGS 31
Clavigralla ankatsoensis sp. n.
(Fig. 77)
Length: 9, 9:9 mm (n = 2). ¢ unknown. Very similar to C. elongata in almost all respects.
Ratio of lengths of antennal segments I : II : III : IV in holotype 1-00 : 0-85 : 0-81 : 0-91; length of
segment I divided by width of head including eyes in holotype 1:31. Antennae completely lacking in
paratype. Ratio of lengths of rostral segments I : II : III : IV about 1-00 : 0:90 : 0:55 : 0-91. Width of
pronotum across apices of posterolateral spines divided by width of head including eyes in paratype 2°81.
Apices of both posterolateral spines lacking in holotype. Scutellum (Fig. 77) much less strongly convex
than in C. elongata and with more prominent tubercles. Posterolateral angles of abdominal sternites
II-VI produced into spines which stand almost perpendicular to lateral margin of abdomen (as in C.
madagascariensis and C. asterix, Figs 79, 80). Length of posterior tibia divided by that of posterior femur
about 0-78. Pubescence of scutellum distinctly less dense than in C. elongata.
74
rhe
Figs 72-77 Clavigralla species. (72) breviceps, 3 head and pronotum, dorsal view; (73) same, para-
mere, dorsal view; (74) same, spermatheca and sclerites of dorsal wall of gynatrium, dorsal view;
(75) same, lip of 3 genital capsule, posteroventral view; (76) same, 9 abdominal tergites III and
IV; (77) ankatsoensis, scutellum, left lateral view.
REMARKS. Distinguished from C. madagascariensis and C. asterix by the presence on the scutellum
of sparse, erect pubescence and larger tubercles, but resembles them in the form of the postero-
lateral spines of the abdominal sternites.
DISTRIBUTION. Known only from the type-locality in Madagascar.
MATERIAL EXAMINED
Holotype °, Madagascar: Ankatso, ii.1921 (R. Decary) (MNHN, Paris).
Paratype. Madagascar: 1 2, same data as holotype (MNHN, Paris).
Clavigralla madagascariensis sp. 0.
(Figs 78, 79, 81, 84)
Length: 3, 8-0-9-0 mm, mean 8-5 mm (n = 5); 9, 8:7-9-6mm, mean 9:2 mm (n = 4). Similar to C.
elongata in overall appearance but differs in the longer spines at sides of thorax and abdomen, which are
more nearly perpendicular to body margin.
Ratio of lengths of antennal segments I : II : III : IV in male about 1-00 : 0-81 : 0-74 : 0-77, in female
about 1-00 : 0-83 : 0-75 : 0-78. Length of antennal segment I divided by width of head including eyes in
W. R. DOLLING
Figs 78-86 Clavigralla species. (78) madagascariensis, 2 pronotum, dorsal view; (79) same, lateral
margins of 2 abdominal sternites III-VII, ventral view; (80) asterix, holotype 3, lateral margins
of abdominal sternites III-VII; (81) madagascariensis, paramere, dorsal view; (82) asterix, para-
mere, dorsal view; (83) same, holotype 3, pronotum, dorsal view; (84) madagascariensis, sperm-
atheca and sclerites of dorsal wall of gynatrium, dorsal view; (85) asterix, aedeagus, left lateral
view; (86) same, dorsal view, basal apparatus omitted.
AFRICAN POD BUGS 33
male 1:56-1:66, mean 1-61 (n = 5), in female 1-42-1-64, mean 1-54 (n = 4). Ratio of lengths of rostral
segments I : II : Ill : IV about 1-00 : 0-88 : 0-46 : 0-77.
Pronotum (Fig. 78) with posterolateral angles drawn out into long, slender, anterolaterally directed
spines; width across apices of spines divided by width of head including eyes in male 3:36-3:90, mean
3-58 (n = 4), in female 3-00-3.67, mean 3-40 (n = 4). Scutellum convex, but less so that in C. elongata,
depressed in midline. Corium with apex at rest reaching level of middle of laterotergite VI. Posterior tibia
almost straight, its length divided by length of posterior femur about 0-83.
Abdominal sterna III—VII with posterolateral angles drawn out into long spines (Fig. 79), the spines
almost perpendicular to abdominal margin and rather slender. Paramere (Fig. 81) and conjunctiva very
similar to those of C. elongata. Ovipositor as in elongata; sclerites of wall of gynatrium (Fig. 84) slender,
sharply angled outwards apically ; spermatheca (Fig. 84) with bulb narrow. duct short and not convoluted.
Sculpture, pubescence and colour generally as in C. elongata except that pubescence of pronotum and
scutellum is much shorter, scutellum completely lacking erect or suberect pubescence and only a few hairs
of this type present on pronotum.
RemaRKS. Differs from e/ongata in the average longer pronotal posterolateral spines, in the
arrangement of spines on pronotal disc, which resembles that of shadabi, in the form of the
sclerites of the gynatrial wall and the form of the spermatheca and in the spines at the postero-
lateral angles of the abdominal sternites, which are almost perpendicular in this species and
deflected posteriorly in elongata. Clearly forms a group with ankatsoensis and asterix but differs
from the former in the absence of erect pubescence on the scutellum and from the latter in the
form of the pronotum. In a long series of elongata from Baie d’Antongil in the MNHN, Paris are
three females which appear to be intermediate between madagascariensis and elongata. In all three
the pronotal posterolateral angles are very strongly produced and in two of them the spines at
the posterolateral angles of the abdominal sternites stand out almost perpendicular to the sides
of the body. In all three the scutellum is as strongly convex as in elongata but has only a few out-
standing hairs in addition to the dense, decumbent pubescence. One female was dissected and
proved to have the spermathecal duct and the sclerites of the wall of the gynatrium intermediate
in form between the two species. This mixture of characters suggests that there may be some
genetic interchange between the two species.
DISTRIBUTION. Madagascar.
MATERIAL EXAMINED
Holotype 3, Madagascar: Antalaha, xii.1935 (Vadon) (MNHN, Paris).
Paratypes. Madagascar: 1 3, 1 9, Tananarive, 1898 (Noualhier) (MNHN, Paris); 4 3, 2 9°, Fampa-
nambo, iii.1959, x.1959, 1962 (J. Vadon) (MRAC, Tervuren and BMNH, London).
Clavigralla asterix sp. n.
(Figs 80, 82, 83, 85, 86)
Length: 3, 8-0-8:2 mm (n = 2); 2, 8:7 mm (n = 1). Closely related to C. elongata, ankatsoensis and mada-
gascariensis, which it resembles in most features.
Ratio of lengths of antennal segments I : II : III : IV in males 1-00 : 0-80 : — and 1-00 : 0:86 :-:-,
in female 1-00 : 0-82 : 0-78 : — (—- denotes missing segment). Length of segment I divided by width of head
including eyes in males 1:26 and 1-35, in female 1:36. Ratio of lengths of rostral segments I : II : II : IV
in both sexes about 1-00 : 1:00 : 0:56 : 0-93.
Pronotum (Fig. 83) with posterolateral angles produced into long, very broadly based, laterally directed
spines; width across apices of spines divided by width of head including eyes 3:79 in holotype male, apices
of spines missing in paratypes. Arrangement of semicircle of spines on disc of pronotum as in shadabi and
madagascariensis. Scutellum less strongly convex than that of C. elongata. Corium with apex at rest reach-
ing to level of basal half of laterotergite VI. Posterior tibia very weakly curved at base, its length divided
by that of posterior femur about 0-79.
Abdominal sternites III-VII with posterolateral angles produced into long, broadly based spines per-
pendicular to abdominal margin (Fig. 80). Paramere (Fig. 82) and conjunctiva (Figs 85, 86) differing little
from those of C. elongata. Sclerites of wall of gynatrium slender, long, resembling those of madagascar-
iensis. Spermatheca with bulb longer and more slender than that of madagascariensis, with a slender, distal
tail, duct not convoluted and about one and one-third times as long as bulb. Sculpture, colour and
34 W. R. DOLLING
pubescence as in elongata, except that pubescence of scutellum and pronotum resembles that of mada-
gascariensis in that scutellum lacks outstanding hairs and pronotum has only a few, short, erect hairs
arising from granules at margins.
REMARKS. Resembles madagascariensis in most of the features in which it differs from elongata;
readily distinguished from all its relatives by the form of the posterolateral pronotal angles.
DISTRIBUTION. Known only from the type-locality in Madagascar.
MATERIAL EXAMINED
Holotype 3, Madagascar: Ambohimitombo, 1894 (Forsyth-Major) (BMNH, London).
Paratypes. Madagascar: 1 3d, 1 2, Ambohimitombo Forest, 24.i.1895 (Forsyth-Major) (BMNH,
London). .
Clavigralla schnelli (Villiers) comb. n.
(Figs 87-96)
Myla schnelli Villiers, 1950 : 656-658, fig. 16. Holotype 3, GUINEA (MNHN, Paris) [not examined].
[Myla gracilis Schouteden; Linnavuori, 1971 : 175. Misidentification.]
Lancha schnelli (Villiers); Shadab, 1972 : 1-11.
Length: 3, 8-8-11:2 mm, mean 10-1 mm (n = 24); 9, 9-3-12:0 mm, mean 11-1 mm (n = 21). Resembles
C. elongata in general appearance.
Antennifers slightly divergent. Ratio of lengths of antennal segments I : II : II] : IV in male about
1-00 : 0:75 : 0-77 : 0-76, in female about 1-00 : 0-74 : 0-75 : 0-71. Length of segment I divided by width of
head including eyes in male 1:68-2:14, mean 1-96 (n = 22), in female 1:79-2:12, mean 1-98 (n = 20).
Ratio of lengths of rostral segments I : II : III : 1V in both sexes about 1-00 : 0-87 : 0-67 : 0-99.
Pronotum (Figs 87-89, 91) moderately declivent anteriorly, prescutellar spines small, disc with a few,
small, spine-like tubercles and a pair of larger spines close to mid-point of anterolateral margins, postero-
lateral angles drawn out into long, tapering spines which are very variable in shape and are directed more
or less obliquely forward. Length of posterolateral spine, measured from extreme posterolateral angle of
pronotum to apex of spine, divided by width of head including eyes in male 1:29-2:09, mean 1-82 (n = 21),
in female 1-78—2-23, mean 1:97 (n = 19); distance between apices of spines divided by width of head in-
cluding eyes in male 2:71-3-94, mean 3-23 (n = 21) in female 2:77-3:83, mean 3:20 (n = 19); length of
spines not closely correlated with distance between their apices due to extreme variability of angle at which
they project from pronotum. In most examples seen apices of posterolateral spines fall anterior to level of
anterior margin of pronotum, but in some examples with short, widely spreading spines the apices do not
project anteriorly beyond anterior margin of calli. Scutellum slightly longer than wide, flat or with disc
very slightly convex, coarsely granulate-punctate; its most prominent feature the two basal, raised knobs.
Corium with apex narrowly produced, its apex at rest reaching posteriorly to level of basal one-quarter
of laterotergite VI. Metathoracic wing with antevannal vein well defined (not shown by Shadab, 1972 :
fig. 8). Anterior and intermediate femora without subapical spines, posterior femur with two long, sub-
apical spines beneath, about 6 very small spines between them and an apical series of 3-4 spines; posterior
tibia straight, its length divided by length of posterior femur 0:76-0:88, mean 0-82 (n = 39).
Abdominal sternites III-VII with posterolateral angles produced into very slender, posteriorly directed
spines (Shadab, 1972 : fig. 1; Villiers, 1950 : fig. 16). Male genital capsule with posterior lip entire, moder-
ately broad, tongue narrowly pointed. Paramere (Figs 92-96) with apical margin strongly oblique, apex
rarely more prolonged (Fig. 94), external angle of blade rarely more broadly rounded (Fig. 96) than usual.
Conjunctiva entirely membraneous, ventrally with distal ventrolateral and apical ventral lobes normally
developed, dorsally with distal dorsolateral lobes absent, distal dorsomedian lobe obsolete, intermediate
dorsal lobe very large, high, conical, slightly bifid laterally at apex, dorsomedian lobe transverse, each of
its dorsal angles with a vertical, finger-like lobe about as high as main part of dorsomedian lobe. Female
with abdominal sternite VII cleft for one-third of its length. Second valvula with a single, entire process
at apex (Fig. 90). Spermatheca (Shadab, 1972 : fig. 18) with bulb very narrow, duct slightly undulating but
not convoluted. Sclerites of wall of gynatrium V-shaped; median, longitudinal arms gently convergent
apically, about one and one-third times as long as anterior, transverse arms.
Head obsoletely granulate-punctate. Pronotum fairly strongly, thoracic pleura and scutellum less
strongly granulate-punctate. Basal halves of anterior veins of corium granulate, clavus and corium
punctate. Antennal segments I-III and femora minutely and sparsely granulate. Thoracic and abdominal
sterna rastrate.
AFRICAN POD BUGS 35
Figs 87-96 Clavigralla schnelli. (87) pronotum of 9, Zaire, Faradje; (88) pronotum of 3, Ivory
Coast, Lamto; (89) pronotum of 3, Zaire, Mpese; (90) apex of second valvula of ovipositor, left
lateral view; (91) pronotum of 3, Zaire, Macaco, Luebo, dorsal view; (92) left paramere, dorsal
view, Ivory Coast; (93) same, Zaire, Macaco, Luebo; (94) same, Luebo; (95) same, Zaire, Mpese;
(96) same, Zaire, Ngowa.
Head, pronotum, scutellum, clavus, corium, thoracic pleura, abdominal sterna, legs and antennae with
short, semierect pubescence. Decumbent, white pubescence present on head, all exposed parts of thorax
and abdomen and on clavus, corium and femora, this pubescence forming dense, white, narrow bands
along midline of pronotum and scutellum and along anterolateral margins of pronotum, four narrow,
longitudinal bands on vertex and a broad, transverse band on disc of pronotum.
General coloration of exposed parts of head, thorax and abdomen and of clavus and corium pale
ferruginous-brown, becoming darker and redder towards apices of posterolateral pronotal angles; spines
36 W. R. DOLLING
at apices of pronotal posterolateral angles and spines at abdominal margin piceous. Abdomen sometimes
with 2-5 paler, continuous, longitudinal bands beneath. Rostrum concolorous with body, antennae and
legs stramineous, sometimes with apical part of posterior femur ferruginous-brown.
REMARKS. A distinctive species, figured in dorsal view by Villiers (1950 : fig. 16) and Shadab
(1972 : fig. 1). The form of the pronotum and to a lesser extent of the parameres are variable and
are often characteristic for a particular locality though they do not vary systematically on a large
geographical scale. The extreme forms of the paramere shown in Figs 94 and 96 are from in-
dividuals which appear to have suffered some damage during growth. There is no evidence from
the genitalia to justify the division of this taxon into more than one species despite the great
variability of the pronotum. Specimens with less well-developed pronotal posterolateral angles
might be confused with C. shadabi but may be distinguished from that species by the narrower
body and from it and all species of the elongata-group by the flat or just perceptibly convex
scutellum.
There is a specimen of this species in MRAC, Tervuren labelled as ‘Holotype’ of Myla gracilis
Schouteden. It was not designated by Schouteden (1938 : 293) in his original description of M.
gracilis and, although it is evidently part of his (mixed) type-series, it disagrees with the detailed
description of gracilis in respect of the antennifers, rostrum, antennae, femora and genital
capsule.
DISTRIBUTION. West and Central Africa.
MATERIAL EXAMINED
Guinea: 1 3, Mt Nimba, Kéoulenta, 11.vi.1942 (M. Lamotte) (MNHN, Paris) (paratype of Myla schnelli
Villiers). Ivory Coast: 10 3, 10 2, Toumodi, Lamto, 11.ix, 6.xi, 4.xii.1962, 2.i, 26.ii, 13.iii, 23.iv, 7.v,
14.v.1963, 13.vii, 24.vii.1964 (BMNH, London). Zaire: 6 3, 10 2, Faradje, 3° 40’ N, 29° 40’ E, i.1913
(Lang & Chapin); 1 3, Faradje, i.1912 (Lang & Chapin); 1 3, 1 9, Medje, 2° 25’ N, 27° 15’ E, 10.iv.1910
(Lang & Chapin) (AMNH, New York); 2 3, Mpese, v—vi.1937 (R. P. J. Cooreman); 1 3, Ngowa, 16.xi.1938
(R. P. J. Mertens); 1 9, Ngowa, 1938 (J. Mertens) (IRSNB, Brussels); 1 3, Luebo, on Urena lobata
Linnaeus (Malvaceae) viii.1921 (Lt Ghesquiére) (labelled ‘holotype’); 1 3, Macaco (Luebo), 30.ix.1921 (Dr
H. Schouteden) (labelled ‘paratype’); 1 9, Kikwit, xi.1920 (P. Vanderijst) (MRAC, Tervuren) (the last three
specimens are syntypes of Myla gracilis Schouteden); 1 3, 110 km S. of Sampwe, 980 m, 21.i.1958 (E. S.
Ross, R. E. Leech) (CAS, San Francisco).
Clavigralla mira sp. n.
(Figs 97-103)
Length: 3, 7:-5-8:7 mm, mean 7:1 mm (n = 12); 2, 8-0-8:8 mm, mean 8-3 mm (n = 11). Rather similar
in build and appearance to C. elongata.
Antennifers slightly to moderately divergent. Ratio of lengths of antennal segments I : II : III : IV in
male about 1:00 : 0:89 : 0-89 : 0-81, in female about 1-00 : 0-87 : 0-87 : 0-74. Length of antennal segment I
divided by width of head including eyes in male 1:26-1:56, mean 1-38 (n = 13), in female 1-23-1-38, mean
1-32 (n = 11). Rostrum short, at rest with apex of segment I reaching posteriorly to level of centre of eye
and apex of segment IV reaching posterior margin of mesosternum. Ratio of lengths of rostral segments
I : II : III : IV in both sexes about 1-00 : 0-89 : 0-68 : 0-89. Frons strongly elevated between antennifers
(Fig. 97), not evenly declivent.
Pronotum (Fig. 98) more shallowly declivent than in C. elongata, collar broad, posterolateral angles”
more nearly perpendicular to sides of body and more slenderly produced; disc of pronotum with a pair
of slender spines near middle of anterolateral margins and several much shorter spines behind them;
width across apices of posterolateral spines divided by width of head including eyes in male 2-29-2-73,
mean 2°54 (n = 10), in female 2:27-2:69, mean 2°48 (n = 11). Scutellum convex, but less so than in C.
elongata. Corium with apical margin almost straight, apex scarcely produced and reaching only to base
or middle of laterotergite VI. Anterior and intermediate femora each with or without a small, subapical
spine beneath, posterior femur with two major subapical spines, 0-2 very small spines between them and an
apical series of 3-4 spines. Posterior tibia straight, its length divided by that of posterior femur in both
sexes 0:78-0:86, mean 0-82 (n = 19).
Abdominal sternites III-VII with posterolateral angles more strongly produced than in C. elongata,
directed posterodorsally. Male genital capsule with lip broad, tongue entire. Paramere (Fig. 99) with apical
margin oblique. Conjunctiva (Figs 100, 101) with dorsomedian lobe low, confluent with distal dorso-
AFRICAN POD BUGS EY;
Figs 97-103 Clavigralla mira. (97) head, 3, right lateral view; (98) pronotum, 3, dorsal view; (99)
paramere, dorsal view; (100) phallotheca, conjunctiva and vesica, left lateral view; (101) con-
junctiva and vesica, ventral view; (102) spermatheca, Zaire, Kapanga; (103) spermatheca and
sclerites of dorsal wall of gynatrium, dorsal view, Ivory Coast.
median lobe and only visible as a small lobe at each side of it; distal dorsomedian lobe voluminous, with
a pair of slender, gently curved, sclerotized processes apically; distal dorsolateral lobes absent; apical
ventral lobes present, membranous; distal ventrolateral lobes divided, one part of each forming a long,
narrow lobe, the other part fused with its fellow on the opposite side to form a low ridge across ventral
| surface of conjunctiva; left end of this ridge produced into a short, membranous extension. Female with
apex of second valvula bearing a more or less acutely pointed, bifurcate process similar to that of C.
elongata. Spermatheca (Fig. 103) with duct short, gently curved or longer and slightly convoluted. Sclerites
of wall of gynatrium as in Fig. 103; gynatrial sac large.
Colour generally dark brown; tibiae, except basal and apical annuli, and first segment of tarsi stramine-
Ous; spines of pronotum and abdominal margin black. Pubescence distributed as in C. elongata, with the
shorter hairs of body, femora and hemelytra white, making whole insect appear grey; midline of pro-
notum and scutellum marked by a conspicuous line of adpressed, white hairs; two additional, similar lines
of pubescence sometimes present, at least anteriorly, on pronotum parallel to midline; dorsum of head,
thoracic pleura and lateral areas of abdominal sterna with faint, oblique lines of adpressed, white hairs.
Veins of hemelytral membrane dark brown, uninterrupted.
RemaRKS. Although closely related to C. elongata and its allies by reason of the bifurcate apical
appendage of the second valvula, this species is unique in the genus in having sclerotized append-
38 W. R. DOLLING
ages to the distal dorsomedian lobe of the conjunctiva. The asymmetry of the ventral, membranous
lobes of the conjunctiva is also remarkable. The spermathecae of two females from Thysville and
Kapanga (Zaire) had longer bulbs and ducts than those of examples from Lamto (Ivory Coast)
and Ngowa (Zaire) (compare Figs 102, 103). These specimens are excluded from the type-series
and the above description is based only on the typical females. No difference was found between
the genitalia of males from Lamto and from Nasisi Hills (Tanzania). Males from Zaire and females
from Tanzania were unavailable, but see C. annectans, below.
DISTRIBUTION. Tropical Africa from Ivory Coast to Tanzania.
MATERIAL EXAMINED
Holotype 3, Ivory Coast: Lamto (Toumodi), 28.v.1964 (D. Gillon) (BMNH, London).
Paratypes. Ivory Coast: 12 3, 10 2, Lamto, 3.vii, 4.xii, 18.xii.1962, 15.i, 26.11.1963, 21.i, 28.i, 1.iv, 28.v,
13.vii, 16.vii, 21.vii, 28.viii.1964 (D. Gillon) (BMNH, London). Zaire: 1 2, Bandundu, Ngowa, xii.1937
(R. P. J. Mertens) (IRSNB, Brussels). Tanzania: 1 3, Nasisi Hills, 20 miles (32 km) N. of Mumias, 4800 ft
(1460 m), 14-15.vi.1911 (S. A. Neave) (BMNH, London).
Material excluded from type-series. Zaire: 1 9, Kongo Central, Thysville, 1959-1963 (R. Michaux);
1 2, Kasai Orientale, Kapanga, xi.1933 (G. F. Overlaet) (MRAC, Tervuren).
Clavigralla annectans sp. n.
(Figs 243-246)
Length: 3, 8-2 mm(n = 1). Known only from the holotype. Indistinguishable in external appearance from
C. mira, which it resembles in form, sculpture, pubescence and coloration, and especially in the bulging
frons (Fig. 243).
Length of antennal segment I divided by width of head including eyes 1-50; ratio of lengths of antennal
segments I : II : III : IV as 1-00 : 0-83 : 0-79 : 0-69. Length of rostral segment I divided by width of head
including eyes 0-59, ratio of lengths of rostral segments I : II : III : IV as 1-00 : 0-90 : 0-66 : 0-90. Width
of pronotum across apices of posterolateral spines divided by width of head including eyes 2-35. Length
of posterior femur divided by length of posterior tibia 0-81. Paramere (Fig. 244) with first tooth of blade
well developed (obsolete in mira). Conjunctiva (Figs 245, 246) with dorsomedian lobe well developed,
extending medially in an erect, narrowly conical lobe and laterally in similar sized, posteriorly directed
lobes; distal dorsomedian lobe very small and narrow, distal dorsolateral lobes obsolete, intermediate
dorsal lobe absent; no sclerotized appendages present on any dorsal lobe; ventral apical lobes of normal
size and shape for the genus, their surfaces minutely warty; distal ventrolateral lobes normally developed,
lightly sclerotized; no other lobes present; all of conjunctiva symmetrical. Ejaculatory reservoir complex
with wings and straps long.
REMARKS. Differs from C. mira, to which it is obviously closely related, in the more strongly
toothed paramere and particularly in the symmetrical conjunctiva lacking dorsal sclerotized
appendages. The form of the conjunctiva is typical of the genus but the inflated frons is a character
shared only with C. mira. Thus C. annectans forms a link between that bizarre species and the
normal members of the e/ongata-group. A single specimen of this species was received in a con-
signment of material sent after the descriptions of the e/ongata-group were prepared. It is possible
that the two females excluded from the type-series of C. mira, above, on the basis of their longer
spermathecal ducts, may prove to belong to the present species if more material with associated
males and females should become available. The localities of these two females, Lulua-Kapanga
and Thysville, are at 8° 22’S, 22°37’ E and 5° 16’S, 14° 15’ E respectively, while the type-
locality of annectans is at 7° 11’ S, 29° 09’ E. A female of typical mira form is recorded above from
Ngowa at 5° 41’ S, 16° 41’ E. These locality data suggest that annectans may be sympatric with
mira in the middle part of the latter’s range, in which case the profound differences in the genitalia
between these two superficially indistinguishable species may be the result of strong selection for
an isolating mechanism in the recent history of the species pair.
DISTRIBUTION. Zaire.
MATERIAL EXAMINED
Holotype 3, Zaire: 10 miles (16 km) S. of Kapona, 13.i.1958 (E. S. Ross, R. E. Leech) (CAS,
San Francisco).
AFRICAN POD BUGS 39
Clavigralla insignis (Distant) comb. n.
(Figs 104-107, 109)
Acanthomia insignis Distant, 1908 : 442. Holotype 2, UGANDA (BMNH, London) [examined].
Length: 3, 8:7-9:2 mm(n = 2); 9, 9:1-9:2 mm(n = 2). Similar to C. elongata but more brightly coloured
and more strikingly patterned and with apical margin of corium almost straight.
Ratio of lengths of antennal segments I : II : III : [Vin both sexes about 1-00 : 0-83 : 0-79 : 0-81. Length
of segment I divided by width of head including eyes in male 1-39-1-58 (n = 2), in female 1:36-1:47
(n = 2). Ratio of lengths of rostral segments I : II : III : [V about 1-00 : 0-88 : 0-70 : 0-94 in both sexes.
Pronotum (Fig. 104) with posterolateral spines directed almost perpendicularly to sides of body, width
across apices of spines divided by width of head including eyes in male 2:72-2-81 (n = 2), in female
2:64-2:67 (n = 2). Disc of pronotum with a short, stout spine near middle of each anterolateral margin
and a number of smaller, short, blunt spines behind them. Scutellum (Fig. 107) rather strongly convex
and a little longer than broad. Corium with apical margin almost straight, apex reaching extreme base of
laterotergite VI at rest. Posterior tibia straight, its length divided by that of posterior femur about 0°83.
Abdominal sternites III-VII with posterolateral angles drawn out into spines (Fig. 106) which are
directed posteriorly as in C. elongata. Male genital capsule with lip broad, tongue acute; paramere (Fig.
109) with apical margin slightly concave. Aedeagus with sclerites protecting base of vesica longer than
those of C. elongata; conjunctiva with all lobes membranous; paired apical ventral and distal ventro-
lateral lobes present, distal dorsolateral and ventral lobes absent, dorsomedian lobe low, flat-topped,
distal dorsomedian lobe obsolete, intermediate dorsomedian lobe present, slightly larger than dorsomedian
lobe. Female genitalia generally similar to those of C. elongata but spermatheca (Fig. 105) with bulb
longer, narrow part of duct less convoluted and broad part of duct longer.
Sculpture and pilosity generally as in C. elongata but short, decumbent, white pubescence forming four
more or less distinct longitudinal lines on dorsal surface of head, three very distinct longitudinal lines on
pronotum and three to five distinct lines along scutellum; sides of head, thoracic pleura and sides of
abdominal sterna each with two oblique lines of similar white pubescence. Scutellum much less densely
pubescent than in C. elongata.
Head, thorax, abdomen, femora and antennal segment I dark red-brown; spines of pronotum and at
sides of abdomen black; antennal segments II-VI yellow-brown with IV infuscate; tibiae and tarsi
stramineous, all tibiae with distinct basal annulus piceous to black, tarsi and apices of tibiae infuscate.
Clavus and corium bright orange-brown, clavus with median piceous stripe, corium with a narrow stripe
along part of R and a broad, irregular stripe along parts of Cu, M and apical margin piceous; membrane
milky white, veins boldly marked with chocolate brown.
REMARKS. The male specimen from Ethiopia has the posterolateral angles stouter and the pronotal
spines and granules less pronounced than in the other specimens seen. The holotype is figured by
Distant (1909 : pl. 2, fig. 7).
DISTRIBUTION. Highlands of East Africa.
MATERIAL EXAMINED
Acanthomia insignis Distant, holotype 9, Uganda: Ruwenzori, 5600 ft (1700 m) (G. F. Scott-Elliot)
(BMNH, London).
Ethiopia: 1 3, Borodda, 2500 m, 18—19.xi.1948 (H. Scott) (BMNH, London). Zaire: 1 3, 1 9, Kivu,
Butembo, ix—x.1965 (P. Célis) (MRAC, Tervuren).
Clavigralla minor (Schouteden) comb. n., stat. n.
(Fig. 110)
Acanthomia insignis forma minor Schouteden, 1938 : 290. LECTOTYPE 3, BuRUNDI (MRAC, Tervuren),
here designated [examined].
Length: 3, 8-1 mm (n = 1); 9, 8-6 mm (n = 1). Very similar to C. insignis but smaller.
Ratio of lengths of antennal segments I : II : III : [Vin male as 1:00 : 0-85 : 0-85 : 0-89, in female as
1-00 : 0-83 : 0-87 : 0-87; length of segment I divided by width of head including eyes in male 1-39 (n = 1),
in female 1-32 (n = 1). Ratio of lengths of rostral segments I : II : III : IV in male as 1-00 : 0-87 : 0-63 :
0-99, in female as 1-00 : 0:87 : 0-69 : 0-99.
40 W. R. DOLLING
105
;
109 110 (
Figs 104-111 C/avigralla species. (104) insignis, 3, pronotum, dorsal view; (105) same, holotype 9,
spermatheca, dorsal view; (106) same, 3, lateral margin of abdominal sternites III-VII, ventral
view; (107) same, holotype 9, scutellum, left lateral view; (108) andersoni, holotype 3, scutellum,
left lateral view; (109) insignis, left paramere, dorsal view; (110) minor, lectotype, left paramere,
dorsal view; (111) andersoni, holotype, left paramere, dorsal view.
Width of pronotum across apices of posterolateral spines divided by width of head including eyes in
female 2:59 (n = 1), male pronotum damaged.
Paramere of male (Fig. 110) with apical margin convex and shaft proportionately longer than in
C. insignis.
Sculpture, pubescence and coloration as in C. insignis.
REMARKS. Because of the difference in the shape of the paramere this ‘form’ is treated as a full
species, distinct from C. insignis.
DISTRIBUTION. Known only from the type-locality in Burundi.
MATERIAL EXAMINED
Acanthomia insignis minor Schouteden, lectotype 3, Burundi: Kitega, xi.1935 (P. Lefevre) (MRAC,
Tervuren).
Burundi: 1 2, same data as lectotype (paralectotype of Acanthomia insignis minor) (MRAC, Tervuren).
AFRICAN POD BUGS 41
Clavigralla andersoni sp. n.
(Figs 108, 111)
Length: 3, 9-4 mm (n = 1). Very similar to C. insignis. Known only from the male holotype.
Ratio of lengths of antennal segments I : II : III : IV as 1-00 : 0-83 : — :— (segments III and IV missing);
length of segment I divided by width of head including eyes 1-42. Ratio of lengths of rostral segments
I: 1: III :1V as 1-00 : 0-88 : 0-62 : 0-96.
Posterolateral angles of pronotum directed more anteriad than in C. insignis; width across apices of
posterolateral spines divided by width of head including eyes 2:58. Scutellum (Fig. 108) weakly convex.
Length of posterior tibia divided by length of posterior femur 0-87.
Paramere of male (Fig. 111) with apical margin strongly oblique.
Dark markings of clavus, corium and membrane veins much less evident than in C. insignis.
REMARKS. Differs from C. insignis chiefly in the form of the scutellum and the paramere.
DISTRIBUTION. East Africa.
MATERIAL EXAMINED
Holotype 3, ‘British East Africa’ (7. J. Anderson) (BMNH, London).
Clavigralla angolensis sp. n.
(Figs 112-114)
Length: 3, 8-1 mm (n = 1). Similar to C. elongata and C. insignis but smaller and with posterolateral
angles of abdominal sternites III-VII produced almost perpendicularly to sides of body. Known only from
the male holotype.
Ratio of lengths of antennal segments I : II : III : IV as 1-00 : 0-80 : 0-78 : 0:89; length of segment I
divided by width of head including eyes 1-42. Ratio of lengths of rostral segments I : II : III : [Vas 1:00:
0-87 : 0-69 : 1:04.
Pronotum (Fig. 112) strongly declivent, posterolateral angles strongly produced and directed slightly
forwards; width across apices of posterolateral spines divided by width of head including eyes 2-95; disc
with a slender spine near middle of each anterolateral margin and a few very short spines behind these.
Scutellum strongly convex, biseriately granulate longitudinally. Corium with apical margin almost
straight, at rest with apex reaching to level of base of laterotergite VI. Length of posterior tibia divided by
length of posterior femur 0:81.
Abdominal sternites III-VII with posterolateral angles produced laterad into rather slender spines
standing almost at right angles to the line of the abdominal margin (Fig. 113). Male genital capsule with
lip broad, tongue acute; paramere (Fig. 114) rather thick, apical margin short and oblique.
Pubescence and colour pattern as in C. insignis except that ground colour of head, thorax, abdomen,
antennal segment I and femora is orange-brown, the same colour as clavus and corium; femora with a
few, obscure, paler markings in apical third; tibiae with basal annuli orange-brown, not black.
REMARKS. Easily distinguished from the other species of the C. elongata and C. insignis groups by
the combination of almost perpendicular abdominal marginal spines with the small size.
DISTRIBUTION. Known only from the type-locality in Angola.
MATERIAL EXAMINED
Holotype 3, Angola: Capeio, 22.vii.1931 (Mrs W. P. Cockerell) (BMNH, London).
Clavigralla egregia sp. n.
(Fig. 116)
Length: 2, 9:5 mm (n = 1). Similar to C. insignis but differs in the longer, laterally directed posterolateral
spines of the pronotum and abdominal sterna. Known only from the female holotype.
Ratio of lengths of antennal segments I : II : III : IV as 1-00 : 0:75 - 0-72 : 0-76; length of segment I
divided by width of head including eyes 1-74. Ratio of lengths of rostral segments I : II : III : [Vas 1-00 :
0-93 : 0-61 : 1-00.
Pronotum (Fig. 116) with posterolateral angles very strongly produced laterally, width across apices of
posterolateral spines divided by width of head including eyes 3-97; disc with several small, blunt spines
42 W. R. DOLLING
119
Figs 112-119 Clavigralla species. (112) angolensis, holotype 3, head, pronotum and scutellum,
dorsal view; (113) same, lateral margins of abdominal sternites III-VII, ventral view; (114) same,
left paramere, dorsal view; (115) aculeata, holotype 3, lateral margins of abdominal sternites II-
VU, ventral view; (116) egregia, holotype °, pronotum, dorsal view; (117) aculeata, holotype 3d,
pronotum, dorsal view; (118) same, paratype 3, conjunctiva and vesica, left lateral view; (119)
same, paratype 3, left paramere, dorsal view.
and one larger, slender spine near the middle of each anterolateral margin. Corium with apical margin
straight, at rest with apex reaching to level of base of laterotergite VI. Posterior tibia straight, 0-81 times
as long as posterior femur.
Abdominal sternites III-VII with posterolateral angles drawn out laterally into spines almost per-
pendicular to lateral margin of abdomen. Spermatheca and female genital sclerites as in C. shadabi (cf.
Fig. 69).
Coloration, pilosity and sculpturing as in C. insignis.
AFRICAN POD BUGS 43
REMARKS. Distinguished from similarly coloured species of this group by the very elongate
pronotal posterolateral angles, and from all except C. angolensis by the radiating abdominal
spines.
DISTRIBUTION. Known only from the type-locality in Malawi.
MATERIAL EXAMINED
Holotype °, Malawi: Mlanje, 17.iii.1913 (S.A. Neave) (BMNH, London).
Clavigralla longispina sp. n.
Length: 3, 8:7 mm (n = 1); 9, 9:2-9:5 mm, mean 9-3 mm (n = 3). Closely allied to C. insignis which it
very closely resembles in appearance and coloration.
Ratio of lengths of antennal segments I : II : III : IV in both sexes about 1-00 : 0-76 : 0-75 : —- (IV miss-
ing in all examples seen); length of segment I divided by width of head including eyes in male 1:53 (n = 1),
in female 1:32-1:48 (n = 2). Ratio of lengths of rostral segments I : II : III : IV in both sexes about
1-00 : 0-87 : 0-59 : 0-95.
Pronotum with posterolateral angles strongly produced, width across apices of posterolateral spines
divided by width of head including eyes in male 3:25 (n = 1), in female 3-02-3-16 (n = 2). Length of
posterior tibia divided by length of posterior femur in both sexes 0:80-0:83, mean 0-82 (n = 4).
Abdominal sternites III-VII with posterolateral angles produced into spines which are longer and less
posteriorly directed than in C. insignis. Male paramere indistinguishable from that of C. insignis; inter-
mediate dorsal lobe of conjunctiva rather long.
Coloration, sculpture and distribution of pubescence as described for C. insignis except that basal
annuli of tibiae are red-brown and femora bear pale marmoration in their apical thirds.
REMARKS. Distinguished from C. insignis chiefly in the form of the posterolateral spines of the
pronotum and abdominal sternites.
DISTRIBUTION. Southern Zaire and northern Zambia.
MATERIAL EXAMINED
Holotype 3, Zambia: Lower Kalungwisi Valley, 1100 m, dense forest, 12—13.ix.1908 (S. A. Neave) (UM,
Oxford).
Paratypes. Zaire: 1 9, Bandundu, Malongi, vi.1943 (H. J. Bredo) (IRSNB, Brussels). Zambia: 1 9,
Chisanga Plateau, Kalungwisi District, 1400 m, 25.ix.1908 (S. A. Neave); 1 9°, N. Lake Bangweolo,
Luwingu, 1300 m, 3.vi.1908 (S.A. Neave) (UM, Oxford).
Material excluded from type-series. Zaire: 1 2, 62 km NE. of Lubumbashi (Elizabethville), 1225 m,
22.1.1958 (E. S. Ross, R. E. Leech) (CAS, San Francisco). (Received after description was prepared.)
Clavigralla aculeata sp. n.
(Figs 115, 117-119)
Length: 3, 8-75-9.1 mm (n = 2); 2, 9:-6-10-:0 mm, mean 9:8 mm (n = 3). Rather similar to C. insignis in
general appearance and coloration.
Antennifers moderately divergent. Ratio of lengths of antennal segments I : II : II : IV in male (1
specimen only) as 1:00 : 0:82 : 0-85 : 0-95; in female (1 specimen only) as 1-00 : 0-86 : 0-90 : 0-82; length
of segment I divided by width of head including eyes in male 1-48-1-49 (n = 2), in female 1:39-1:45, mean
1-42 (n = 3). Ratio of lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0-88 : 0-70 :
1-01.
Pronotum (Fig. 117) about as steeply declivent as in C. elongata, with posterolateral angles directed
more strongly forwards; disc with stouter, stronger spines than in C. insignis or C. elongata; width across
apices of posterolateral spines divided by width of head in male 2:68-2:84 (n = 2), in female 2:58-2:86,
mean 2-69 (n = 3). Scutellum strongly convex, with two longitudinal rows of 3-4 short, blunt tubercles.
Corium with apex distinctly but shortly produced, its apex at rest reaching posteriorly to level of base of
laterotergite VI. Anterior and intermediate femora each with a small subapical spine beneath, posterior
femur with two major subapical spines, some very small spines between them and an apical series of four
spines. Posterior tibia straight, its length divided by that of posterior femur in both sexes 0-82-0-84, mean
0-83 (n = 5).
Abdominal sternites III-VII with posterolateral angles (Fig. 115) produced into strong spines which are
slightly but distinctly curved posteriad and more nearly perpendicular to lateral margins of abdomen than
44 W. R. DOLLING
are those of C. elongata and C. insignis. Male genital capsule with lip broad, paramere (Fig. 119) of similar
shape to that of C. insignis. Conjunctiva (Fig. 118) with ventral apical lobes and distal ventrolateral lobes
developed as in C. elongata, dorsomedian lobe low, flat-topped, distal dorsomedian lobe very weakly
developed, narrow intermediate dorsal lobe present between dorsomedian and distal dorsomedian lobes,
distal dorsolateral lobes absent. Spermatheca as in C. insignis (cf. Fig. 105) except that the part of the duct
between the ‘valve’ and the junction with the gynatrium is shorter. Sclerites of wall of gynatrium as in
C. shadabi (cf. Fig. 69).
Resembles C. insignis in distribution of pubescence and in colour pattern but head, thorax, abdomen,
femora and antennal segment I paler, gingery brown, and brown markings of clavus and corium less
intense.
REMARKS. Distinguished from C. insignis by stouter spines of pronotal disc, paler colour and
differently shaped abdominal spines and from C. angolensis by the shorter pronotal posterolateral
spines. Rather similar in appearance to C. horrida but lacks the long spines on the scutellum and
the additional femoral spines.
DISTRIBUTION. South Africa.
MATERIAL EXAMINED
Holotype 3, South Africa: Natal, Estcourt, 1896 (BMNH, London).
Paratypes. South Africa: 1 3, 2 2, Natal, Estcourt, 1896 (BMNH, London); 1 9, Transvaal, Zoutpans-
berg, 800 m, -.x.— (Schouteden) (MRAC, Tervuren).
Clavigralla horrida (Germar) nec auctt. post.
(Figs 120-125)
[Alydus acantharis (Fabricius); Thunberg, 1822 : 1. Misidentification based on specimen now at University
of Uppsala, examined.]
[Cimex acantharis Linnaeus; incorrectly cited in synonymy with Alydus acantharis (Fabricius) by Thun-
berg: 1822 > 1.]
Syromaster horridus Germar, 1840 : 145. LECTOTYPE 2, SoutH ArricaA (MNHU, Berlin), here design-
ated [examined].
Clavigralla muricata Stal, 1855 : 31. LECTOTYPE ¢, SoutH Arrica (NR, Stockholm), here designated
[examined]. [Synonymized with Clavigralla acantharis (Fabricius) sensu Thunberg by Stal, 1866 : 108.]
Syn. n. of horrida.
[Clavigralla acantharis (Fabricius) sensu Thunberg; Stal, 1866 : 108.]
Acanthomia muricata (Stal); Stal, 1873 : 83.
(The binomen Clavigralla horrida (Germar) was first used by Stal, 1866 : 108 for the species treated above
under the name of C. elongata Signoret, q.v.)
Length: 3, 8-4-9-8 mm, mean 9:2 mm (n = 12); 9, 9-7-11- ‘Omm, mean 10:5 mm (n = 13). Related to
C. elongata but body form more robust.
Antennifers moderately divergent. Ratio of lengths of antennal segments I : II : III : IV in male about
1-00 : 0-91 : 0-76 : 0-70, in female about 1-00 : 0-94 : 0-78 : 0-70; length of segment I divided by width of
head including eyes in male 1-51—1-81, mean 1-65 (n = 13), in female 1-44-1-67, mean 1-54 (n = 13).
Ratio of lengths of rostral segments I : II : III : [Vin male about 1-00 : 0-94 : 0-56 : 0-80, in female about
1:00 : 0-90 : 0-54 : 0-76.
Pronotum (Fig. 120) with two prominent, stout spines near middle of anterolateral margins and some
smaller ones on disc, some of these arranged in a semicircle terminating anteriorly at the two larger spines;
posterolateral angles strongly produced anterolaterally and moderately elevated, each terminating in a
short spine; width across apices of posterolateral spines divided by width of head including eyes in male
2:41-2:86, mean 2:64 (n = 11), in female 2:65-3:07, mean 2:86 (n = 13). Scutellum (Fig. 123) strongly
elevated with two longitudinal rows of up to 5 stout, erect spines. Corium with apex very slightly produced,
at rest reaching posteriorly to level of suture between laterotergites V and VI. Anterior and intermediate
femora each with a single subapical spine beneath; posterior femur (Fig. 124) with 3-4 major subapical
spines, sometimes 1 or 2 minor spines between prepenultimate and penultimate major spines and between
penultimate and last major spines, an apical series of 4 spines and also with a subapical row of up to 3
small spines on posterior edge of lower face of femur. Posterior tibia straight, its length divided by that of
posterior femur in both sexes 0:87-0:96, mean 0-91 (n = 25).
AFRICAN POD BUGS 45
121
122
Figs 120-125 Clavigralla horrida. (120) 2 pronotum, dorsal view; (121) left paramere, dorsal view;
(122) lateral margins of 2 abdominal sternites III-VII, ventral view; (123) lectotype, scutellum,
left lateral view; (124) apex of 2 right posterior femur, posterior view; (125) spermatheca and
sclerites of dorsal wall of gynatrium, dorsal view.
Abdominal sternites III-VII with posterior angles (Fig. 122) produced into broad spines which are much
less strongly deflected posteriorly than those of C. elongata. Male genital capsule with lip broad, tongue
entire; paramere (Fig. 121) with apical margin slightly sinuate. Aedeagus with long, membranous distal
dorsolateral lobes, resembling aedeagus of C. shadabi (cf. Fig. 70). Spermatheca (Fig. 125) with duct only
about as long as bulb, portion of duct between ‘valve’ and gynatrium very short. Process at apex of second
valvula bifid with the lower part of the fork shorter than the upper.
Sculpture of integument as in C. elongata but more pronounced on pronotum, where granules on area
around posterolateral angles are developed into small, black-tipped spines or tubercles. Pilosity generally
as in C. elongata but colour of hairs dead white and hairs organized into two more or less distinct
longitudinal broad bands on pronotum with a fine band between them along midline. Pubescence of
46 W. R. DOLLING
pronotum otherwise sparse. Colour of head, thorax and abdominal sternites red-brown with piceous
markings as in C. elongata; spines of pronotum, scutellum and abdominal margins black, at least at apex;
antennal segments I and IV, femora and basal annuli of tibiae uniform red-brown; remainder of tibiae,
clavus and corium pinkish yellow; antennal segments II and III red. Membrane of hemelytra milky white,
its veins in part brown but contrast in colour between membrane and veins not nearly as pronounced as
in C. insignis and its close allies.
REMARKS. The long, outstanding spines of the scutellum at once distinguish this species from all
others in the genus.
DISTRIBUTION. Rhodesia and South Africa.
MATERIAL EXAMINED
Syromaster horridus Germar, lectotype 2, South Africa: ‘Promontorium Bonae Spei’ (C. F. Drége)
(MNHU, Berlin). Clavigralla muricata Stal, lectotype 3, South Africa ‘Cap. B. Spei’ (Victorin) (NR,
Stockholm) [Stal, 1855 : 31 cites locality as ‘in terra Natalensi’].
Rhodesia: 1 3, Vumba Mts, xii.1933 (NM, Bulawayo). South Africa: 1 9, ‘Caffraria’ (NR, Stockholm);
1 ex., ‘Caput Bonae Spei’ [det. by C. P. Thunberg as Alydus acantharis] (University of Uppsala); 1 3, 1 9,
‘C.G.H.’;1 3,1 2, ‘Cape’; 1 3, ‘S. Africa’ (BMNH, London); 1 9, ‘British Caffrara Castilneau 1862’ (UM,
Oxford); 1 3, Natal, Cathedral Peak, 18.xii.1966 (J. G. H. Londt) (TM, Pretoria); 1 2, Cape Prov.,
Grootvatersbosch, For. Res. 22km N. of Heidelburg, 5.ii.1968 (Schuh, Slater, Sweet & Slater) (Slater
coll.); 1 3, Cape Prov., Simonstown, ii.1893 (P. de la Garde) (UM, Oxford); 1 9, Cape Prov., 21 km E. of
Gansbaai, 2.ii.1968 (J. A. & S. Slater & M. Sweet) (PPRI, Pretoria); 1 3, 2 2, Cape Prov., Stellenbosch,
5.xi.1925 (H. Brauns) ; 1 3, 1 9, Cape Prov., Swellendam, 17.xii.1931-18.i.1932 (R. E. Turner); 1 9, no
data; 8 3, 9 9, Cape Prov., Ceres, 450 m, xii.1920, 1.1921, ii.1921, 2-21.ii1.1921, 1-12.xi.1924, xii.1924,
ii.1925 (R. E. Turner) (BMNH, London); 1 2°, Cape Prov., Pearl, 16.xi.1949 (Malkin); 1 3, Cape Prov., Du
Toits Kloof, 22.xi.1949 (B. Malkin) (CAS, San Francisco).
Clavigralla natalensis Stal
(Figs 126-130)
Clavigralla natalensis Stal, 1855 : 31. LECTOTYPE 9°, SoutH Africa (NR, Stockholm), here designated
[examined].
Acanthomia natalensis (Stal) Stal, 1873 : 83.
Length: 3, 9:2-10:4 mm, mean 9:9 mm (n = 4); 2, 10-6-12:0 mm, mean 11-8 mm (n = 14). The most
robust of the species allied to C. elongata.
Antennifers moderately divergent. Ratio of lengths of antennal segments I : II : III : IV in male about
1-00 : 0-86 : 0-80 : 0-92, in female about 1-00 : 0-85 : 0-78 : 0-85 (figure for male segment IV is based on
only two observations and is the mean of 0-82 and 1-02); length of segment I divided by width of head
including eyes in both sexes 1:16-1:38, mean 1-27 (n = 18). Ratio of lengths of rostral segments I : II:
III : IV in male about 1-00 : 0-97 : 0-71 : 1:06, in female about 1-00 : 0-90 : 0-67 : 1-00.
Pronotum (Fig. 126) strongly declivent, with a semicircle of small spines on disc, posterolateral angles
much less strongly produced than in related species, tipped with short spines; width across apices of
posterolateral spines divided by width of head including eyes in male 2:39-2:69, mean 2°53 (n = 4), in
female 2:43-2:77, mean 2-62 (n = 14). Scutellum slightly longer than broad, weakly convex, with two
longitudinal rows of large granules. Corium with apex produced, at rest reaching posteriorly to about one-
third of the way along laterotergite VI. Anterior and intermediate femora each with a strong subapical
spine beneath; posterior femur (Fig. 127) with 3-5 major spines beneath, increasing in length towards
apex of femur, without minor spines between them, apical series of 4 spines present but no additional
posterior row of spines (cf. C. horrida). Posterior tibia straight, its length divided by that of posterior
femur in both sexes 0:83-0:92, mean 0-88 (n = 18).
Abdominal sternites III-VII with posterior angles (Fig. 128) produced into short, posteriorly directed
spines. Male genital capsule with lip broad, tongue entire; paramere (Fig. 129) stout. Conjunctiva gener-
ally as in C. elongata except for distal dorsomedian lobe, which is lower. Female with bifid apical process
of second valvula well developed. Spermatheca (Fig. 130) with bulb narrow and duct short, sclerites of
dorsal wall of gynatrium (Fig. 130) slender.
Sculpture of integument generally as in C. elongata but granulation everywhere weaker. Pubescence of
head, pronotum, scutellum, thoracic pleura, thoracic and abdominal sterna, femora and antennal segment
I almost uniform, consisting of short, colourless, suberect to erect hairs interspersed with short, decum-
AFRICAN POD BUGS 47
127
. VY
130
128
Figs 126-130 Clavigralla natalensis. (126) pronotum of 9, dorsal view; (127) apex of left posterior
femur, anterior view; (128) lateral margin of $ abdominal sternites IIi—VII, ventral view; (129)
left paramere, dorsal view; (130) spermatheca and sclerites of dorsal wall of gynatrium, dorsal
view.
bent, white, curled but not tomentose hairs, organized into a thin, sometimes indistinct line along midline
of pronotum and a line on each side of vertex contiguous with each eye. Clavus, corium and abdominal
laterotergites with similar but shorter and sparser pubescence of both types; antennal segments II-IV,
tibiae and tarsi with short, suberect pubescence interspersed with very short, indistinct, decumbent
pubescence. Colour largely red-brown; antennae with segments I-III paler and segment IV darker than
general body coloration; abdominal sternites slightly paler; clavus and corium testaceous, veins coloured
brown or red-brown, especially in anterior and distal areas of corium, membrane very pale fuscous, its
veins brown, uninterrupted; all tibiae stramineous with basal annulus piceous and apical annulus
red-brown.
REMARKS. A distinctive species by reason of the heavy build and the shape of the pronotum.
DIsTRIBUTION. Africa south of the Zambesi river.
MATERIAL EXAMINED
Clavigralla natalensis Stal, lectotype 2, South Africa: ‘Caffraria’ (J. Wahlb.) (NR, Stockholm).
Rhodesia: 1 9, Mashonaland, Salisbury, 1898 (BMNH, London). South Africa: 1 3, Transvaal,
Johannesburg E., Bedford Ridge, 25.x.1953 (A. L. Capener) (Slater coll.); 1 9, Johannesburg, 5.ii.1958
48 W. R. DOLLING
(Capener) (Slater coll.); 1 3, Johannesburg, 1800 m, x.1898 (J. P. Cregoe); 1 2, Johannesburg, 1800 m,
vi.1899 (Cregoe); 1 2, Johannesburg, 1.v.1905 (Cregoe); 1 3, Transvaal, Klerksdorp, Snowball, ix.1899;
1 3, Transvaal, Lydenberg District (BMNH, London); 2 9, Orange Free State, Bothaville (Dr Brauns)
(TM, Pretoria); 2 2, Natal, Pietermaritzburg, 1902 (Cregoe); 1 9, Natal, Drakensberg, Van Reenen,
1-22.1.1927 (R. E. Turner); 1 9, Natal, Estcourt, viii.1895; 1 2, Natal, Howick, 1806; 1 9, Howick
(BMNH, London); 1 3, Transvaal, Veekraal, 14.xii.1963 (A. L. Capener) (PPRI, Pretoria); 1 2, Natal,
Bray Hill, viii, Gram (BMNH, London).
The tomentosicollis-group
Antennifer with apical process long, deflexed; metathoracic peritreme with dorsal ridge entire; tongue of
male genital capsule trifid, bifid or entire; lip of genital capsule filling posterior emargination in posterior
view; pubescence of anterior, declivent*area of pronotum contrasting strongly in form and colour with
that of posterior, horizontal part. Second valvula of ovipositor variously shaped but never with apical
appendage. Pronotum with posterolateral angles not or relatively weakly prominent, posterolateral spines
arising abruptly from them.
The leontjevi-subgroup
Tongue of male genital capsule entire; pronotal disc without sublateral tubercles; membrane of hemely-
tron unpigmented.
Clavigralla leontjevi (Bergroth) comb. n.
(Figs 131-136)
Acanthomia leontjevi Bergroth, 1908 : 107. LECTOTYPE 6, Erniopia (ZMU, Helsinki), here designated
[examined].
Length: 3, 6:0-7-1 mm, mean 6-6 mm (n = 22); 9, 5-9-7:5 mm, mean 6:9 mm (n = 21). A small insect,
similar in appearance to a small C. tomentosicollis.
Head subquadrate; antennifers very slightly divergent. Length of antennal segment I divided by width
of head including eyes in male 1:07-1:29, mean 1:21 (n = 22), in female 1-06—1-30, mean 1-17 (n = 21).
Ratio of lengths of antennal segments I : II : III : IV in male about 1-00 : 0-78 : 0-66 : 1-00, in female
about 1:00 : 0-78 : 0-67 : 0-96. Length of basal segment of rostrum divided by width of head including
eyes in male 0:60-0:72, mean 0:65 (n = 20), in female 0:61-0:69, mean 0-66 (n = 16). Ratio of lengths of
rostral segments I : I] : III : IV in male about 1-00 : 0-91 : 0-63 : 0-95, in female about 1-00 : 0-88 :
0-60 : 0-91.
Pronotum (Fig. 131) with posterolateral angles bearing slender spines directed laterally or slightly
antero- or posterolaterally, spines gently upcurved; width of pronotum across apices of spines divided by
width of head including eyes in male 2:36-2:95, mean 2:54 (n = 18), in female 2-34-3-00, mean 2°55
(n = 18). Disc of pronotum with a cluster of granules in anterior part of midline but completely without
projecting tubercles or spines, even near lateral margins. Scutellum strongly convex. Anterior and inter-
mediate femora each with 0-1 subapical spine beneath; posterior femur with 2 major subapical spines
beneath, the second about one and one-half times as long as the first, about 5 small spines present between
the major spines and a terminal series of four spines present. Posterior tibia distinctly curved at base,
length of posterior tibia divided by length of posterior femur in male 0:66-0:74, mean 0-70 (n = 21), in
female 0:64-0:74, mean 0-69 (n = 19). Apex of corium at rest reaching to level of suture between
laterotergites V and VI.
Abdominal sternites III-VII with posterolateral angles produced into well-developed, acute spines, each
spine about one-third as long as lateral margin of corresponding sternite. Male genital capsule with tongue
broadly rounded distally; paramere (Fig. 132) with apex strongly inflexed and with two well-developed
teeth on blade, the distal one more strongly sclerotized than the other and sited on the dorsal face.
Aedeagus (Figs 133, 134) with vesica long, sclerites at its base large, together almost as wide as con-
junctiva. Conjunctiva with dorsomedian lobe very low, its lateral angles strongly produced, intermediate
dorsal lobe separated from distal dorsomedian lobe, which is weakly developed as are distal dorsolateral
lobes; distal ventrolateral lobes the largest, apical ventral and ventral lobes present; none of the lobes
sclerotized. Female tergite IX apically deflexed, anus directed posteroventrally. Spermatheca (Fig. 136)
with duct short, slightly convoluted and rather wide especially near junction with gynatrium. Sclerites of
dorsal wall of gynatrium (Fig. 135) each with an anterolaterally directed spur arising from the longitudinal
arm. Laterotergite VII clearly separated from mediotergite VII in both sexes.
AFRICAN POD BUGS 49
Sculpture of integument as in C. tomentosicollis. Pubescence and coloration generally similar to those of
C. tomentosicollis. Pale, declivent area of pronotum interrupted by a small patch of piceous granules in
anterior midline; area of pale pubescence projecting posteriorly into the patch of darker pubescence be-
hind it in four places, the two pale prominences nearest midline tending to project as backward sloping
tufts and without a tuft of darker hairs between them; midline of posterior, dark brown area frequently
with an incipient line of pale, silvery hairs. Hemelytral membrane slightly milky, hyaline, its only dark
markings being two very small piceous or brown blotches on two veins at the point where these enter the
membrane from the apical margin of the corium.
133
136
135
Figs 131-136 Clavigralla leontjevi. (131) pronotum of 3, dorsal view; (132) left paramere, dorsal
view; (133) conjunctiva and vesica, dorsal view; (134) same, left lateral view; (135) sclerites of
dorsal wall of gynatrium, dorsal view; (136) spermatheca.
REMARKS. This species differs from all others in the genus in the complete absence of large
tubercles or spines from the declivent area of the pronotum. Another unique feature is the partially
deflexed tergum IX and anus of the female, approaching the condition which obtains in the genus
Gralliclava in the Oriental region. The complete separation of laterotergites VII from medio-
tergite VII in the male is also unique. However, in most features this species accords well with the
characters of the tomentosicollis-group of species. There is some variation in the pronotal postero-
lateral spines, which may be directed slightly anterolaterally, laterally or slightly posterolaterally,
and in the amplitude of the undulations in the line marking the boundary between the pale and
dark areas of pronotal pubescence. The male paramere is frequently slightly narrower than the
example illustrated (Fig. 132), in which an incipient third tooth is visible on the ventromedian
area of the blade at its junction with the shaft. This third tooth is usually not as strongly developed
as that illustrated and may often be absent. From the material available it is not possible to make
a definite statement as to the status of these variants but they do not appear to be correlated one
with another, nor do they seem to vary consistently on a geographical basis. Risbec (1950; 1951;
1955) records the following Hymenoptera as parasites of the egg: Anastatus aliberti Risbec and
A. nezarae Risbec (Eupelmidae), Paravignalia hemipterae Risbec (Pteromalidae) and Gryon gnidus
Nixon (Scelionidae).
DISTRIBUTION. Africa between the Sahara desert and the Zambesi river.
MATERIAL EXAMINED
Acanthomia leontjevi Bergréth, lectotype 3, Ethiopia: Adue, 1896 (other data illegible) (ZMU, Helsinki).
50 W. R. DOLLING
Senegal: 3 3, Bambey, 29.i.1943, on potato (J. Risbec); 2 2, Bambey, iv.1940 (Risbec) (BMNH, Lon-
don). Ivory Coast: 2 3, Lamto (Toumodi), 24.vii.1964; 1 9, Lamto, 4.xii.1962 (BMNH, London). Ghana:
2 3, 3 2, Kintampo, 7.xii. 1965 (D. Leston); 4 3, 1 9, Tafo, 3.i1.1967, 21.1.1967, 13.11.1967, 30.11.1967, by
pyrethrum knockdown from cocoa (Theobroma) (R. Kumar, D. Leston); 1 3, locality illegible, 7.x.1967
(Leston); 1 3, Twenadurasi, 28.i1.1967 (Leston); 1 3, Tapa Amonya, Volta Region, 3.xi.1967; 1 9, Pram-
pram, 27.viii.1966 (UG, Legon). Togo: 1 9°, Missahoué, 650 m, vi.1963 (Mme Y. Schach) (MRAC,
Tervuren). Nigeria: N. Nigeria, Chafe, 7.vii.1977 (J. C. Deeming, A. O. Medaiyedu); 1 °, Zaria, Samaru,
12.i1.1970 (J. C. Deeming) TAR, Samaru); 1 9, Ilorin, 13.xii.1929, on cotton (Gossypium) (F. D. Golding);
1 3, 2 2, Jos, 9.1.1956 (Bechyne) (BMNH, London). Zaire: 1 9, Faradje, 29° 40’ E, 3° 40’ N, xii.1912
(Lang, Chapin) (AMNH, New York); 1 3, Ngowa, 5.vi.1939 (R. P. J. Mertens); 2 3, Musoa, vi-vii.1939,
xi.1939 (H.-J. Brédo) (IRSNB, Brussels). Angola: 2 9, no exact data (Welwitsch) (BMNH, London and
Duarte coll.). Sudan: 1 3, 2 °, Blue Nile, Singa-Damazin, 15—17.xi.1962 (R. Linnavuori) (BMNH, Lon-
don). Kenya: 1 2, Kisumu, 1143 m, xi.1920 (Gedye); 1 3, Thika, ix.1936 (F. J. Gedye) (BMNH, London).
Tanzania: 1 3, E. Rukwa, 1050 m, iv.1938 (D. G. McInnes); 1 3, Urikiriguru, 25.viii.1958 (J. A. D.
Robertson); 1 2, Urikiriguru, 19.ix.1958, on cotton (Gossypium) (Robertson) (BMNH, London). Malawi:
1 9, Valley of N. Rukuru, Karonga District, 600-1200 m, 15-18.vii.1910 (S. A. Neave) (BMNH, London).
Zambia: 1 2, Abercorn, 15.xii.1943 (H.-J. Brédo) (IRSNB, Brussels). Rhodesia: 1 2, Lomagundi, vi.1929
A. Cuthbertson) (NM, Bulawayo).
Clavigralla griseola (Linnavuori) comb. n.
Acanthomia griseola Linnavuori, 1978: 35. Holotype 9, SOUTHERN YEMEN (Linnavuori coll.) [not
examined].
This species was described too late to be included in the key. It is very close to /eontjevi. 1 have seen a single
2 paratype, from Eritrea. This specimen differs from /eontjevi in the more intense dark markings, parti-
cularly those of the femora; the width of the pronotum divided by that of the head is only 2:29 and the
antennae are longer, the length of segment I divided by width of head being 1:31. Southern Yemen and
Ethiopia (Eritrea).
The ruandana-subgroup
Tongue of male genital capsule entire; pronotal disc with a pair of large, blunt tubercles sublaterally;
membrane of hemelytron suffused with brown pigment.
Clavigralla ruandana (Schouteden) comb. n.
(Figs 137-143)
Acanthomia ruandana Schouteden, 1957 : 309-310. LECTOTYPE 3, RwanpDA (MRAC, Tervuren), here
designated [examined].
Length: 3, 9:0-10.0 mm, mean 9.5 mm (n = 8); 9, 9:3-10°6 mm, mean 9:9 mm (n = 9). Rather more
slender than C. tomentosicollis, which it somewhat resembles.
Head (Fig. 137) elongate. Length of antennal segment I divided by width of head including eyes in
male 0:87-1:11, mean 1-00 (n = 8), in female 0-80-1-10, mean 0-97 (n = 9). Ratio of lengths of antennal
segments I : II : II] : IV in male about 1-00 : 1-05 : 0-73 : 1-12, in female about 1-00 : 1-07 : 0-75 : 1-06.
Length of basal segment of rostrum divided by width of head including eyes in both sexes 0:81-1:01, mean
0-91 (n = 17). Ratio of lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0-94 : 0:59 :
0-90.
Pronotum (Fig. 138) with posterolateral spines short, directed slightly posteriorly. Width of pronotum
across apices of posterolateral spines divided by width of head including eyes in male 2:48-3:06, mean
2:66 (n = 8), in female 2:31-2:79, mean 2:56 (n = 9). Scutellum strongly convex. Corium with apex pro-
duced, at rest reaching level of middle of laterotergite VI. Posterior tibia almost straight; length of poster-
ior tibia divided by that of posterior femur 0:73-0:82, mean 0-77 (n = 17).
Abdominal sternites III-VI with posterolateral angles (Fig. 139) very slightly prominent, not acute,
posterolateral angle of sternite VII acutely projecting in male, narrowly rounded in female. Male genital
capsule (Fig. 140) with lip shallowly emarginate, tongue narrowly rounded. Paramere (Fig. 141) falcate,
apex strongly curved medially, tooth at mid-point of blade small. Conjunctiva with dorsomedian lobe
bearing large, finger-like, erect lobes at its posterolateral angles; intermediate dorsal, distal dorsomedian
and distal dorsolateral lobes poorly developed; apical ventral and distal ventrolateral lobes well developed;
ventral lobes very small; none of the lobes sclerotized. Second valvula of ovipositor (Fig. 142) slender,
140
143
146
147
Figs 137-147 Clavigralla species. (137) ruandana °, head, right lateral view; (1 38) same, lectotype 3,
pronotum, dorsal view; (139) same, left lateral margin of ¢ abdominal sternites III—-VII, ventral
view ; (140) same, apical half of 3 genital capsule, dorsal view; (141) same, left paramere, dorsal
view; (142) same, second valvula, lateral view; (143) same, sclerites of dorsal wall of gynatrium,
dorsal view; (144) biston, 2 pronotum, dorsal view; (145) same, left lateral margin of 2 abdominal
sternites III—V11; (146) same, left paramere, dorsal view; (147) same, sclerites of dorsal wall of gyn-
atrium, dorsal view.
52 W. R. DOLLING
apical third abruptly narrower than basal two-thirds; sclerites of dorsal wall of gynatrium (Fig. 143) with
longitudinal arms abruptly divergent at apices and barbed; spermatheca similar to that of C. tomentosi-
collis except that bulb is distinctly narrower, duct slightly longer and sometimes with incipient
convolutions.
Sculpture as in C. tomentosicollis. General coloration a slightly rufous brown, clavus and corium paler.
Piceous markings variable, generally comprising a streak occupying ventral surface of antennal segment I,
the whole ventral surface and often parts of sides of head, thoracic sterna, a horizontal bar on each of the
thoracic pleura, a patch in anterior midline of pronotum, posterolateral spines and sublateral tubercles
of disc of pronotum, base of scutellum, spots on femora, base of abdominal venter except midline and a
broad, lateral, marginal band on abdominal sterna III-VI, sometimes III—VII, rarely only II-IV.
Stramineous areas include tibiae except for basal, apical and usually median annuli, bases of femora and
often antennal segments II and III. Membrane of hemelytron infuscate with veins and basal cells colour-
less. Pubescence generally as in C. tomentosicollis but often with an incipient longitudinal band of pale
hairs along midline of posterior lobe of pronotum and usually with a narrow band of dense, silvery or
golden hairs along apical margin of corium and clavus contrasting strongly with the almost uniform
colouring and pubescence of the remainder of these parts. Colour of abdominal dorsum varying from
almost uniformly orange to largely piceous.
REMARKS. C. ruandana belongs to a group of species in which the head is elongate and the rostrum
at rest reaches beyond the posterior margin of the mesosternum. Within this group it is distingu-
ished by the basal antennal segment being subequal in length to the width of the head, the non-
acute posterolateral angles of abdominal sterna III-VI, the absence of discrete spots on the
forewing membrane and the moderately long posterolateral pronotal spines. A single female from
Rhodesia is of a pale, golden-brown colour with few piceous markings. It is not possible to say if
this difference is due to a difference in the genetic constitution of the populations in Rhodesia
and elsewhere or if the specimen is merely teneral.
DISTRIBUTION. Highlands of central Africa, mostly around the Rift Valley.
MATERIAL EXAMINED
Acanthomia ruandana Schouteden, lectotype 3, Rwanda: Contrefort Est Muhavura, 2100 m, 28.1.1953 (P.
Basilewsky) (MRAC, Tervuren).
Rwanda: | 3, same data as lectotype; 2 2, Contrefort Sud Nyamateke, 2200 m, 15.xi.1953 (Basilewsky),
all paralectotypes of Acanthomia ruandana (MRAC, Tervuren). Zaire: 3 3, 2 2, Tshibinda [‘Tanganyika
T., Tshibinda’ on labels], 21-27. viii.1931 (Prof. T. D. A. Cockerell, Miss A. Mackie) (BMNH, London);
1 3,1 9, Kivu, Mulugu, Tshibinda, xi.1951 (P. C. Lefevre) (MRAC, Tervuren); 3 9, Lualaba river, 750—
1200 m, 31.v.1907 (Neave) (BMNH, London); 1 3, Malongi, vi.1943 (H. J. Brédo) (IRSNB, Brussels).
Zambia: 1 3, 1 2, Chisinga Plateau, Kalungwisi District, 1400 m, 17.ix.1908 (S.A. Neave) (UM, Oxford).
Rhodesia: 1 2°, Mtarazi Falls, iii.1957 (NV. L. H. Krauss) (BMNH, London).
Clavigralla biston sp. n.
(Figs 144-147)
Length: 3, 9:3 mm (n = 2); 9, 8-7-10:7 mm, mean 9-6 mm (n = 7). Resembles C. ruandana in most
features and probably closely related to that species, differing chiefly in the more pronounced pronotal
spines.
Head elongate. Length of antennal segment I divided by width of head including eyes in male 1-32-1-38
(n = 2), in female 1:35-1:43, mean 1-38 (n = 7). Ratio of lengths of antennal segments I : II : HI : TV in
male about 1-00 : 0-85 : 0-68 : — (IV missing in both specimens seen), in female about 1-00 : 0-87 : 0-68 :
0-92. Length of basal segment of rostrum divided by width of head including eyes in both sexes 0:70-0:79,
mean 0-74 (n = 9). Ratio of lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0-86 :
0:54 : 0-87.
Pronotum (Fig. 144) with poterolateral spines rather long, directed posterolaterally; width across
apices of spines divided by width of head including eyes in male 2:58-2:72 (n = 2), in female 2:69-2:93,
mean 2-79 (n = 5). Scutellum strongly convex. Corium with apex narrowly produced, at rest reaching to
level of base of laterotergite VI. Posterior tibia gently curved at base, its length divided by that of posterior
femur 0:73-0:79, mean 0:75 (n = 8).
Abdominal sternites with posterolateral angles triangularly produced (Fig. 145). Male genital capsule
with lip broadly rounded, tongue rather narrow and parallel-sided for most of its length, apex rounded or
AFRICAN POD BUGS 53
bluntly angulate. Paramere (Fig. 146) with apex strongly angled inwards, tooth at middle of blade moder-
ately prominent. Aedeagus as described for C. oxonis (Fig. 152) except that all parts of ejaculatory
reservoir complex are smaller and vesica is shorter, its non-coiled portion subequal in length to rest of
aedeagus. Second valvula of ovipositor of similar general shape to that of C. oxonis (Fig. 153) but with
apical lobe rather broader. Spermatheca as in C. oxonis (cf. Fig. 149). Sclerites of dorsal wall of gynatrium
(Fig. 147) with recurved processes on median arms large. Coloration generally bright orange-brown (not
red-brown as in its relatives), with darkest markings, on sides and ventral surface of head, thorax and
abdomen, usually red-brown or brown (not piceous). Pronotum (Fig. 144) with midline of declivent area
occupied by a band of brown pubescence terminating posteriorly in a tuft of erect, brown hairs flanked by
two narrow projections of the area of silvery pubescence, these projections themselves each flanked on the
outer side by a triangular area of erect, brown pubescence. Posterior lobe of pronotum behind this area
with decumbent, golden-brown pubescence except for posterior margin and a more or less distinct,
longitudinal, rectangular or triangular area in midline, where the decumbent pubescence is silvery.
Scutellum with rather long, slightly tomentose pubescence and a pair of tufts of erect, brown hairs on
disc. Clavus and most of corium pale orange-brown with short, crisped, semidecumbent, silvery to golden-
brown pubescence, corium with a very distinct band close to apical margin brown, with short, suberect,
brown pubescence, apical margin with a narrow band of silvery pubescence contrasting strongly with the
broader, brown band. Produced apical part of corium pale rufous with short, silvery to golden, semi-
decumbent pubescence. Membrane infumate between the colourless longitudinal veins, the brown
coloration slightly more intense in middle.
REMARKS. A distinctive species by reason of the shape of the pronotum and bright coloration.
DISTRIBUTION. Highlands in the vicinity of the Rift Valley.
MATERIAL EXAMINED
Holotype 3, Malawi: Cholo, 1919-1920 (R. C. Wood) (BMNH, London).
Paratypes. Malawi: | 3, 1 2°, Cholo, 1919-1920 (R. C. Wood) (BMNH, London). Zambia: 1 3, 4 2°, Mid
Luangwa Valley, 600 m (2000 ft), 14-16.viii.1900 (S. A. Neave) (BMNH, London). Tanzania: 1 9, Lulan-
guru, 27 km (17 miles) W. of Tabora, xi.1917, on bushes (G. D. H. Carpenter) (BMNH, London).
Rhodesia: 1 9, Manica, Mpudzi River, c. 900 m (3000 ft), 26.x.1905 (Guy Marshall) (UM, Oxford).
Material excluded from type-series. Angola: 1 3, Caconda, Huile, 30.ix.1949 (Borys Malkin) (CAS, San
Francisco). (Received after description was prepared.)
Clavigralla oxonis sp. n.
(Figs 148-153)
Length: 3, 8-9 mm (n = 1); 2, 9:0-9:1 mm (n = 2). Resembles C. ruandana in general build, but differs
in many details including shape of pronotum.
Head elongate. Antennal segment I very short, its length divided by width of head including eyes in
male 0-75 (n = 1), in female 0-79-0-84 (n = 2). Ratio of lengths of antennal segments I : II : II : IV in
male unknown (specimen mutilated), in female about 1-00 : 1-13 : 0-84 : 1-33. Length of segment I of
rostrum divided by width of head including eyes in both sexes 0:94-1:02, mean 0-99 (n = 3). Ratio of
lengths of rostral segments I : II : III : IV in both sexes about 1-00 : 0:93 : 0-61 : 0-87.
Pronotum (Fig. 148) with posterolateral angles not prominent, posterolateral spines reduced to short
tubercles or (male holotype) evanescent; width across apices of these posterolateral tubercles divided by
width of head including eyes in male 2-15 (n = 1), in female 2:24-2:41 (n = 2). Scutellum strongly
convex. Corium with apex at rest reaching to about level of suture between laterotergites V and VI.
“yin tibia gently arcuate in basal third, its length divided by that of posterior femur 0:68-0:73, mean
“70 (n = 3).
Abdominal sternites III-VII with posterolateral angles rounded, not prominent except in segment VII
of male (Fig. 150). Male genital capsule with lip evenly rounded, tongue narrowly rounded as in C.
ruandana (cf. Fig. 140). Paramere (Fig. 151) falcate, tooth at middle of blade obsolete. Aedeagus (Fig. 152)
with vesica very long, sclerites at its base and wings and straps of ejaculatory reservoir complex large,
symmetrical. Conjunctiva with dorsomedian lobe large, bearing well-developed, sac-like outgrowths at its
posterolateral angles, intermediate dorsal, distal dorsomedian and distal dorsolateral lobes weakly
developed, apical, ventral and distal ventrolateral lobes well developed, ventral lobes absent. Second
valvula of ovipositor (Fig. 153) slender. Sclerites of dorsal wall of gynatrium not differing from those of
C. ruandana. Spermatheca (Fig. 149) with wide portion of duct rather long, narrow portion not convoluted,
bulb narrow.
54 W. R. DOLLING
Pronotum, scutellum, thoracic pleura, apical half of each femur and apical one-third to half of corium
reddish brown; ventral surface of head, base of scutellum, thoracic sterna, midline of declivent part of
pronotum, base of abdominal venter medially and sometimes lateral stripes on head and thoracic pleura
piceous; head, clavus, base of corium, antennal segments I and IV and abdominal sternites pale reddish
brown; laterotergites obscurely patterned with pale and dark reddish brown; antennal segments II and
lll, tibiae, tarsi and bases of femora stramineous; forewing membrane brown, veins and basal cells
colourless. Distribution of hair types as in C. ruandana and C. tomentosicollis; apical margin of corium
with a narrow band of silvery hairs; junction between areas of pale and dark pubescence of pronotum
(Fig. 148) strongly M-shaped, with a tuft of erect, brown hairs entirely surrounded by pale, tomentose
hairs between the anterior corners of the ‘M’.
149
153
Figs 148-153 Clavigralla oxonis. (148) pronotum, 2 paratype, dorsal view; (149) spermatheca;
(150) left lateral margins of abdominal sternites III-VII, ¢ holotype, ventral view; (151) left para-
mere of holotype, dorsal view; (152) conjunctiva and vesica of holotype, left lateral view; (153)
second valvula of paratype, lateral view.
REMARKS. Distinguished from all other members of the genus by the very short pronotal postero-
lateral spines.
DISTRIBUTION. Known only from a small, hilly area of northern Zambia.
MATERIAL EXAMINED
Holotype 3, Zambia: N. Lake Bangweolo, Luwingu, 4200 ft (1300 m), 9.vi.1908 (S. A. Neave) (UM,
Oxford).
Paratypes. Zambia: 2 2, Abercorn, 15.xii.1943 and 4.iv.1944 (H. J. Brédo) (IRSNB, Brussels).
The spiniscutis-subgroup
Tongue of male genital capsule entire; pronotal disc with a pair of large, blunt, sublateral tubercles;
membrane of hemelytron with a row of discrete, brown or piceous spots between bases of longitudinal
veins, sometimes additionally blotched brown or piceous.
Clavigralla bovilla sp. n.
(Figs 154-156)
Length: 3, 8:3 mm (n = 1); 9, 8-4-8-8 mm, mean 8-6 mm (n = 3). Similar in build to C. tomentosicollis;
resembles C. ruandana in the elongate head and posteriorly directed pronotal posterolateral spines.
AFRICAN POD BUGS es Py
135
163
y
Figs 154-165 Clavigralla species. (154) bovilla, left lateral margin of ¢ abdominal sternites III—-VII,
ventral view; (155) same, left paramere, dorsal view; (156) same, second valvula, lateral view;
(157) pusilla, 3 pronotum, dorsal view; (158) same, left lateral margin of $ abdominal sternites
IfI-VII, ventral view; (159) same, left paramere, dorsal view; (160) same, aedeagus, left lateral
view; (161) same, aedeagus, ventral view; (162) same, aedeagus, dorsal view; (163) same, second
valvula, lateral view; (164) same, sclerites of dorsal wall of gynatrium, dorsal view; (165) same,
spermatheca.
Length of antennal segment I divided by width of head including eyes in male 0-96 (n = 1), in female
0-90-0-94, mean 0-92 (n = 3). Ratio of lengths of antennal segments I : II : IIIf : [V in male as 1-00:
0-91 : 0-69 : 1-13, in female about 1-00 : 0-85 : 0-70 : 1-06. Length of basal segment of rostrum divided
by width of head including eyes in both sexes 0:86-0:90, mean 0-88 (n = 4). Ratio of lengths of rostral
segments [ : II : III : IV in both sexes about 1-00 : 0-94 : 0-60 : 0-87.
Pronotum similar in shape to that of C. ruandana (cf. Fig. 138), with the posterolateral spines directed
slightly posteriorly; width across apices of spines divided by width of head including eyes in both sexes
56 W. R. DOLLING
2:57-2:60, mean 2:59 (n = 4). Corium with apex slightly produced, at rest reaching to level of base of
abdominal laterotergite VI. Posterior tibia with gentle curvature near base, its length divided by length of
posterior femur 0:71-0:75, mean 0-73 (n = 5).
Abdominal sternites III-VII (Fig. 154) with posterolateral angles acute, slightly prominent. Male genital
capsule with lip evenly rounded, not emarginate, tongue rounded as in C. ruandana. Paramere (Fig. 155)
with blade broad, thick, apex bluntly hooked, tooth at middle of blade well developed. Conjunctiva could
not be fully expanded in only specimen available. Female with sclerites of wall of gynatrium similar to
those of C. ruandana; spermatheca like that of C. tomentosicollis but bulb narrower; second valvula of
ovipositor (Fig. 156) unevenly narrowed towards apex.
Colour and pubescence generally as in C. ruandana but piceous markings absent from antennal segment
1 and absent or almost absent from sides of abdominal sternites; laterotergites marked with stramineous
and rufous brown, not piceous; junction between pale and dark areas of pubescence on pronotum slightly
undulating, with four evenly spaced tufts of pale hairs at junction, without any pale hairs in midline of
posterior lobe, a tuft of slightly darker hairs present behind each large sublateral tubercle of disc, about
equidistant from tubercle and from junction of pale and dark areas; no line of silvery or golden hairs
present along apical margin of corium or clavus; membrane of hemelytron colourless with a row of
piceous spots between longitudinal veins just distal to basal cells and parallel with apical margin of
corium.
REMARKS. Differs from the other species with long heads (ruandana-subgroup) in the possession of
discrete piceous spots in the forewing membrane, rather than an even suffusion of pigment. The
shape of the second valvula and paramere are characteristic. During dissection of the abdomen
of a female paratype a single, very small, hymenopterous larva was found in the haemocoel. It
was presumably a first instar, being slender, tailed and provided with a broad head bearing large
mandibles. It is possibly a euphorine braconid.
DISTRIBUTION. Known only from the type-locality in the hills of N. Zambia.
MATERIAL EXAMINED
Holotype 3, Zambia: Abercorn, 15.xii.1943 (H.J. Brédo) (IRSNB, Brussels).
Paratypes. Zambia: 4 2, Abercorn, x.1943, xi.1943, 15.xii.1943, 4.iv.1944 (Brédo) (IRSNB, Brussels).
Clavigralla pusilla sp. n.
(Figs 157-165)
Length: 3, 6:5-7:0 mm, mean 6:7 mm (n = 7); 9, 6:7—-7:3 mm, mean 7:0 mm (n = 3). A small, rather
slender species.
Antennifers almost parallel, juga slightly tumid. Ratio of lengths of antennal segments I : II : HI : IV
in male about 1-00 : 0-82 : 0-72 : 0:98, in female about 1-00 : 0-88 : 0-75 : 0-95; length of segment I
divided by width of head including eyes in male 1-28—1-41, mean 1-32 (n = 7), in female 1-22—1-26, mean
1-24 (n = 3). Ratio of lengths of rostral segments I : II : II] : 1V in both sexes about 1-00 : 0-94 : 0-52 :
0-84; length of segment I divided by width of head including eyes 0:57-0:64, mean 0-62 (n = 5).
Pronotum (Fig. 157) strongly declivent, centre of posterior lobe with a short, longitudinal ridge inter-
secting a similar transverse one; spines of posterolateral angles long, laterally directed, straight or gently
upcurved; posterolateral angles just anterior to base of each spine with a pair of short, stout spines; paired,
sublateral spines of disc very prominent; width across apices of posterolateral spines divided by width of
head including eyes in male 2:50-3:07, mean 2:77 (n = 4), in female 2:40-2:73, mean 2-57 (n = 3).
Scutellum convex, midline slightly depressed, apex slightly produced. Corium narrow, its apex slightly
produced, at rest reaching suture between abdominal laterotergites V and VI. Posterior tibia gently curved
near base, its length divided by length of posterior femur 0:65-0:70, mean 0-68 (n = 5).
Abdominal sternites III-VII with posterolateral angles produced into strong spines (Fig. 158). Male
genital capsule with lip rounded, tongue entire and narrowly triangular. Paramere (Fig. 159) stout, with
shallow depression on dorsolateral face, tooth at middle of blade projecting dorsally. Conjunctiva (Figs
160-162) with all lobes membranous, dorsomedian lobe low, its lateral angles projecting almost independ-
ently as small globular lobes slightly distal to main, transverse part of lobe; intermediate dorsal lobe large,
conical, overhanging dorsomedian lobe; distal dorsomedian lobe poorly developed; distal dorsolateral
lobes obsolete; apical ventral, distal ventrolateral and ventral lobes well developed; vesica large in com-
parison with size of conjunctiva. Female with second valvula (Fig. 163) bearing a short, apical process;
sclerites of dorsal wall of gynatrium (Fig. 164) with short ‘barb’ on longitudinal arms; spermatheca (Fig.
165) with bulb short, duct short and simply looped.
AFRICAN POD BUGS 57]
Head, legs and antennae obsoletely granulate, granulation more distinct on apices of femora and tibiae,
pronotum anteriorly granulate, posteriorly punctate, pleura punctate-granulate, thoracic sterna and
abdominal sterna and laterotergites minutely rough, clavus with three rows of punctures, corium in basal
half with a few granules on veins and a few punctures between veins, apically unsculptured. Head and
antennae with short, suberect, colourless pubescence, head and antennal segment I also with short,
decumbent, silvery pubescence; pronotum anteriorly, on disc and along lateral and posterolateral margins
with sparse, moderately long, erect, white pubescence, collar and declivent area also with rather dense,
white, tomentose pubescence, posterior lobe with moderately long, decumbent, silvery hairs mixed with
dense, suberect to decumbent, brown hairs, a small tuft of brown hairs present at middle of junction
between areas of dark and pale pubescence; scutellum with long, dense, tomentose to suberect, silvery
hairs, tinged more or less heavily with brown on disc; thoracic pleura and sterna with rather dense, white,
decumbent, somewhat tomentose pubescence, pleura also with long, suberect, white pubescence; legs
with rather short, suberect, white pubescence, femora also with short, semidecumbent to decumbent,
white pubescence; clavus and corium with short, crisped, semidecumbent to decumbent, white pubescence;
abdominal sterna with dense, decumbent to slightly tomentose, white pubescence, interspersed with long,
erect, colourless hairs; laterotergites with white, decumbent pubescence. Colour generally as in C.
tomentosicollis but antennae pale to mid-brown throughout and membrane colourless except for a row of
piceous spots between bases of longitudinal veins parallel to apical margin of corium and piceous marks
on two veins where they enter membrane from corium.
Ovarian egg (4 ex. seen) ovate, flattened dorsally, with chorion minutely reticulate; aeromicropyles
separated into two groups as follows: 2 contiguous ones ventral to eclosion suture and 3 or 4 contiguous
in a transverse row dorsal to the suture.
RemaRKS. Differs from the only other Malagasy species of the tomentosicollis-group in the
characters of the pigmentation of the membrane and the shape of the tongue of the male genital
capsule, as well as the smaller size, more slender build and less strongly curved posterior tibia.
Superficially similar to the African species C. strabo and C. pabo but body more slender and para-
mere stouter.
DISTRIBUTION. Southern Madagascar.
MATERIAL EXAMINED
Holotype 3, Madagascar: Tulear Province, Betioky, 275 m, 15.iii.1968 (K. M. G. & P. D.) (BMNH,
London).
Paratypes. Madagascar: 2 3, 2 9, same data as holotype (BMNH, London); 1 3, Région Sud de I’Ile,
Bekily, vi.1936 (A. Seyrig); 1 3, Région du Sud, Andrahomana, vii.1901 (Ch Alluaud); 1 3, Région du
Sud, Androy, between Antanimora and Ifotaka, xi.1901 (Al/uaud); 1 3, Région du Sud, Androy, Ambo-
vombé, xi.1901 (Al/uaud); 1 2, Belumbe, 1900 (Fairmaire) (MNHN, Paris).
Clavigralla strabo sp. n.
(Figs 166-173)
Length: 3, 7-0-7:3 mm, mean 7-1 mm (n = 3).; 9, 6-6-8:3 mm, mean 7:7 mm (n = 9). Similar in size
and build to C. spiniscutis.
Head short, antennifers slightly divergent. Ratio of lengths of antennal segments I : II : Il : 1V in male
about 1-00 : 0-77 : 0-65 : 0-89, in female about 1-00 : 0-81 : 0-69 : 0-87. Antennal segment I moderately
robust and bearing long, erect hairs, length of segment divided by width of head including eyes in male
1-06—1-23, mean 1-14 (n = 3), in female 1-05-1-15, mean 1-09 (n = 9). Ratio of lengths of rostral segments
I: 11: Ill : 1V in both sexes about 1-00 : 0-89 : 0-62 : 0-92; length of basal segment divided by width of
head including eyes 0-56-0-66, mean 0-59 (n = 10).
Pronotum (Figs 166, 167) with posterolateral angles moderately prominent, usually with a distinct
spine or tubercle laterally on anterior part of each prominence; posterolateral spines slender, gently up-
curved and also curved slightly anteriorly, variable in length (extreme variants figured in Figs 166, 167);
width across apices of posterolateral spines divided by width of head including eyes in male 2:19-2:27,
mean 2:22 (n = 3), in female 2:13-2:50, mean 2:27 (n = 7). Corium with apex not produced, at rest with
apex reaching posteriorly to level of apical one-third of laterotergite V, rarely as far as suture between
laterotergites V and VI. Posterior tibia straight, its length divided by length of posterior femur 0-65—-0-70,
mean 0-68 (n = 11).
Abdomen with posterolateral angles of sternites III-VI1 produced into rather stout but very acute spines
(Fig. 169); tergum 1X of female slightly deflexed apically. Male genital capsule (Fig. 170) with lip broad,
58 W. R. DOLLING
168
: i
Figs 166-176 Clavigralla species. (166) strabo, 3 pronotum, dorsal view, from Rustenburg; (167)
same, 3 pronotum, dorsal view, Louis Trichardt; (168) same, left paramere, dorsal view; (169)
same, left lateral margin of abdominal sternites III-VII, ventral view; (170) same, apical half of 3
genital capsule, dorsal view; (171) same, second valvula, lateral view; (172) same, spermatheca;
(173) same, sclerites of dorsal wall of gynatrium, dorsal view; (174) pabo, apical half of 3 genital
capsule, dorsal view; (175) same, 3 pronotum, dorsal view; (176) same, left paramere, dorsal view.
170
prominent, slightly truncate, tongue rounded. Paramere (Fig. 168) with apex strongly incurved, blunt,
tooth at middle of blade prominent, well developed. Ejaculatory reservoir complex with wings, straps and
vesica moderately long. Conjunctiva entirely membranous; dorsomedian lobe low, with a pair of short,
finger-like lobes arising from its base posterolaterally; intermediate dorsal lobe large, higher than finger-
like lobes; distal dorsomedian and distal dorsolateral lobes small; apical ventral and distal ventrolateral
lobes normally developed; small, paired, ventral lobes present. Female with apex of second valvula of
ovipositor (Fig. 171) broadly prominent; spermatheca (Fig. 172) with bulb narrow, duct about three times
as long as bulb, not convoluted, with broad and narrow regions about equal in length; sclerites of dorsal
wall of gynatrium (Fig. 173) with broad expansions on median arms.
Head, thoracic pleura and sterna, declivent part of pronotum, scutellum and abdominal sterna with
rather dense, decumbent, silvery white, tomentose pubescence; head dorsally also with very long, sparse,
AFRICAN POD BUGS 59
erect pubescence, laterally, and ventrally with shorter, suberect pubescence. Antennal segment I bearing
short, rather sparse, decumbent, silvery pubescence and long, erect, colourless hairs much longer than
diameter of segment; segments II and III with rather long, suberect hairs the longest of which are longer
than width of segments; pubescence of rostrum mainly consisting of rather short, suberect hairs and, on
segments I and II, some decumbent, silvery hairs; propleura and declivent part of pronotum also with
long, erect hairs in addition to the tomentose indumentum; horizontal part of pronotum with golden-
brown to dark brown, decumbent to erect pubescence, mostly of short to moderately long hairs, in the
posterior part mostly decumbent with a few pale brown, erect hairs intermixed; centre of junction between
anterior, pale and posterior, dark areas of pronotal pubescence with a small brush of erect, brown hairs;
disc and sides of scutellum with long pubescence, white except for anterior part of disc where hairs are
brown; thoracic and abdominal sterna also with moderately long, suberect, silvery hairs; clavus and
corium with short, crisped, silvery to golden pubescence with an admixture of short, slightly tomentose
hairs of the same colour along costal margin of corium and apical margins of clavus and corium; ab-
dominal laterotergites with a mixture of these two types of pubescence; legs with long, erect, colourless to
pale brown hairs much longer than width of tibiae and shorter, suberect hairs, femora also with de-
cumbent, silvery hairs. Head, thoracic pleura and abdominal sterna pale brown with obscure, darker
brown to piceous markings; thoracic sterna piceous; pronotum pale brown anteriorly, chestnut-brown in
posterior, horizontal part, tubercles in anterior midline piceous, this group of tubercles extending laterally
to embrace calli for one-third to one-half of their width but not extending posteriorly beyond level of the
large, sublateral tubercles, sublateral tubercles and posterolateral spines of pronotum piceous; scutellum
pale brown, broadly piceous at base; clavus and corium stramineous, veins anteriorly with some granules
piceous, apical one-third of corium heavily marked with dark or reddish brown mottling; membrane
milky-hyaline, bases of two veins at junction with corium and 1-5 discrete spots between longitudinal
veins close to basal cells dark brown or piceous; femora pale yellowish brown in basal one-quarter to
one-third, otherwise dark reddish brown with a few paler spots, pale area at base of femur sometimes with
extensive darker markings; tibiae and tarsi yellowish brown to stramineous, tibiae with basal and apical
annuli reddish brown; antennae pale yellowish brown, apices of segments I-III and whole of IV sometimes
infuscate; spines at posterolateral angles of abdominal sternites piceous; abdominal laterotergites dark
brown, III—V each with a narrow, transverse band near anterior margin stramineous, VI with a broader
stramineous band in the same position, lateral exposed areas of tergite VII coloured as laterotergites VI.
Ovarian egg with granular, polygonal sculpture on chorion and with 3-4 aeromicropyles close together
in a straight line on rim where flat, dorsal and convex, ventral sides of egg meet.
REMARKS. This species closely resembles C. spiniscutis in general appearance but differs from it in
the pubescence of the scutellum, which is not organized into spine-like tufts, the coloration of the
hemelytral membrane which never has irregular piceous patches but only a few discrete spots,
the straight posterior femur, the much stronger dorsal tooth of the paramere and other details of
the male genitalia.
DISTRIBUTION. All specimens seen were taken at medium altitudes in southern Africa between
latitudes 17° S and 30° S.
MATERIAL EXAMINED
Holotype 3, Botswana: 1 3, Bakgatla, Sebele, in grass, 8.viii.1968 (R. E. Roome) (BMNH, London).
Paratypes. Botswana: 1 9, Ghanzi, 13.ix.1926 (J. Maurice) (BMNH, London); 1 2, Samengeigei,
Kaukau Kungv., vi.1951 (C. Koch) (TM, Pretoria). Rhodesia: 1 2, Sawmills, 1.iv.1923 (BMNH, London).
South West Africa: 1 2, Abachaus, Otjiwarongo District, iii.1950 (G. Hobohm); 1 9°, Ovamboland,
Oshikango, vii.1948 (TM, Pretoria). South Africa: 1 3, 1 2, Rustenburg, xii.1961 (A. L. Capener); 1 3,1 9,
Louis Trichardt, 6.ii.1941 (Capener) (TM, Pretoria); 1 2, Natal, Durban (J. P. Cregoe); 1 °, Transvaal,
Zoutpansberg (H. Junod) (BMNH, London).
Clavigralla pabo sp. n.
(Figs 174-176)
Length: 3, 6-4-7:1 mm, mean 6:7 mm (n = 3). Female unknown. Very similar to C. strabo and C.
spiniscutis in general appearance.
Head short, antennifers distinctly divergent, especially at the base. Ratio of lengths of antennal seg-
ments I : II : III : [V about 1-00 : 0-82 : 0-70 : 0-97; segment I slender, bearing short, suberect hairs much
shorter than diameter of segment; length of segment I divided by width of head including eyes 1-17—1-22,
mean 1-20 (n = 3). Ratio of lengths of rostral segments I : II : III : IV as 1-00 : 0-93 : 0-53 : 0-87 and
60 W. R. DOLLING
length of basal segment divided by width of head including eyes 0-60 in only specimen (from Natal) where
rostrum is visible.
Pronotum (Fig. 175) with posterolateral spines long and slender, width across apices of spines divided
by width of head including eyes 2:19-2:37 (n = 2, both from Mozambique). Posterior tibia distinctly
arcuate basally, its length divided by length of posterior femur 0:70-0:72, mean 0-71 (n = 3).
Genital capsule (Fig. 174) with lip prominent, broadly rounded, tongue very narrow. Paramere (Fig.
176) with tooth at middle of blade very large, apex of blade strongly incurved, acutely pointed. Aedeagus
as in C. strabo.
Coloration and distribution of pubescence as in C. strabo except that the length of the hairs, particularly
the outstanding hairs of the appendages, is much less; longest hairs on antennal segments I-III suberect,
shorter than diameter of segment II; longest hairs on legs suberect to erect, shorter than or just equal to
diameter of tibiae. Hemelytral membrane with rather extensive pale brown markings in addition to a row
of piceous spots between veins close to basal cells. Group of piceous tubercles in anterior midline of
pronotum extending posteriorly well behind level of sublateral tubercles, not deeply embracing calli.
REMARKS. The unique shape of the tongue of the genital capsule distinguishes this species from all
its relatives. Superficially it might be confused with C. strabo, from which it differs in the much
shorter pubescence of the appendages and the shape of the patch of piceous granules in the
anterior midline of the pronotum, or with C. spiniscutis from which it differs in the much paler
pigmented patches of the hemelytral membrane and in lacking the specialized arrangement of
pubescence on the scutellum of that species.
DiIsTRIBUTION. Known only from the Delagoa Bay — Lake St Lucia area on the eastern coast of
southern Africa.
MATERIAL EXAMINED
Holotype 3, Mozambique: I. de Inhaca, 20—22.ii.1971 (F. Simoes) (Duarte coll.).
Paratypes. Mozambique: 1 3, I. de Inhaca, 27—30.iii.1971 (F. Madureira) (Duarte coll.). South Africa:
1 3, Natal, Lake St Lucia, Charters Creek, 12.xi.1967 (J. A. & S. Slater, T. Schuh) (Slater coll.).
Clavigralla spiniscutis (Bergroth) comb. n.
(Figs 177-184)
Acanthomia spiniscutis Bergréth, 1913 : 328-329. Type 3, CENTRAL AFRICAN REPUBLIC [not seen; perhaps
present but unrecognized at ZMU, Helsinki].
Length: 3, 6:9-7:8 mm, mean 7:3 mm(n = 10); 2, 6:8-8-4 mm, mean 7:6 mm(n = 7). A small, generally
dark coloured species of rather stout build.
Head short, antennifers very slightly divergent. Ratio of lengths of antennal segments I : II : II] : [Vin
male about 1-00 : 0-81 : 0-76 : 0-99, in female about 1-00 : 0:79 : 0-69 : 0-92. Antennal segment I rather
stout, gradually and uniformly expanding towards apex, its length divided by width of head including
eyes in male 0:96-1:15, mean 1-04 (n = 9), in female 0:96-1:16, mean 1-05 (n = 7). Ratio of lengths of
rostral segments I : II : III : IV in both sexes about 1-00 : 0-86 : 0-58 : 0-90; length of segment I divided
by width of head including eyes 0:56-0:71, mean 0:61 (n = 17).
Pronotum (Fig. 177) strongly declivent anteriorly, posterolateral angles strongly inflated but not pro-
duced laterally, posterolateral spines short, triangular; width of pronotum across apices of spines divided
by width of head including eyes in male 2:11-2:41, mean 2:28 (n = 9), in female 2:13-2:40, mean 2-22
(n = 7). Scutellum convex, with a pair of callosities on disc, its shape usually obscured by dense tufts of
pubescence (Fig. 177). Corium with apex weakly produced, at rest reaching posteriorly to level of suture
between laterotergites V and VI. Posterior femur with the 2 major subapical spines almost equal in length,
with 4-5 minor spines between them and a terminal series of four spines; posterior tibia short, strongly
arcuate at base, its length divided by that of posterior femur 0:63-0:70, mean 0-66 (n = 16).
Abdominal sterna III-VII with posterolateral angles (Fig. 179) abruptly prominent as short, triangular
teeth. Male genital capsule (Fig. 181) short, lip strongly and broadly projecting, tongue mucronate.
Paramere (Fig. 180) strongly expanded towards apex, especially in vertical plane, tooth at middle of
dorsal surface of blade very broad, apex of blade narrowly hooked but not acute. Conjunctiva (Fig. 182)
rather similar to that of C. curvipes, with dorsomedian lobe broad and bearing small subsidiary lobes at
its posterolateral basal angles; intermediate dorsal lobe large, conical; distal dorsomedian and distal
dorsolateral lobes poorly developed but distinct, a finger-like median lobe arising from posterior face of
conjunctiva above base of vesica; apical ventral lobes large, less rough on surface than those of C.
AFRICAN POD BUGS 61
180
QS 186
184
Figs 177-186 Clavigralla species. (177) spiniscutis, pronotum and scutellum, 9, dorsal view; (178)
same, ¢ left hemelytral membrane and apex of corium, dorsal view; (179) same, left lateral margin
of 2 abdominal sternites III-VII, ventral view; (180) same, left paramere, dorsal view; (181) same,
apical half of 3 genital capsule, dorsal view; (182) same, conjunctiva and vesica, dorsal view; (183)
same, second valvula, lateral view; (184) same, sclerites of dorsal wall of gynatrium, dorsal view;
(185) marmorata holotype 3, right posterior femur and tibia, anterior view; (186) same, left para-
mere of holotype, dorsal view.
curvipes; distal ventrolateral lobes large; ventral lobes absent. Female with second valvula of ovipositor
(Fig. 183) narrowly produced at apex. Spermatheca as in C. curvipes. Sclerites of dorsal wall of gynatrium
(Fig. 184) with anterior arms very broad in the oblique transverse/frontal plane, median longitudinal arms
each with a small external tooth.
Pubescence of antennae of short, suberect, pale hairs shorter than width of segment II, segment I also
with short, decumbent, silvery hairs. Head with white, decumbent pubescence, dorsally very dense and
obscuring ground colour, also with colourless, suberect, short hairs throughout and with long, erect,
colourless hairs along dorsal midline; rostrum with short, suberect, colourless hairs and segments I and I
also with short, decumbent, silvery hairs. Thoracic sterna and pleura and abdominal sterna with moder-
ately long, tomentose, white pubescence interspersed with long or medium length, suberect, colourless
62 W. R. DOLLING
hairs; declivent area of pronotum (Fig. 177) with dense, tomentose, decumbent, white pubescence inter-
spersed with long, sparse, erect, usually brown hairs; a short, transverse ridge of pale, tomentose
pubescence present posterior to each sublateral tubercle of disc; posterior horizontal lobe with decumbent,
golden-brown pubescence; junction between the two areas of pronotal pubescence marked by a median
tuft of moderately long, dark brown hairs flanked by two triangular patches of short, erect, dark brown
hairs, median tuft largely or entirely surrounded by white, tomentose pubescence which continues in a
band along midline of pronotum posterior to tuft. Scutellum (Fig. 177) in fresh, unabraded specimens
with two tufts of dark brown hairs on disc, flanked by long, white hairs which are organized into two
thorn-shaped lateral brushes and an irregularly rounded or bifid posterior fringe. Clavus, corium and
abdominal laterotergites with short, crisped to semidecumbent, silvery and golden-brown pubescence.
Legs with suberect to erect, colourless pubescence the longest hairs of which are about as long as diameter
of tibiae; femora also with decumbent, silvery hairs.
Head pale brown, ventral surface and some lateral markings and ocellar tubercles piceous. Antennae
stramineous, segment I with external basal streak and broad, apical annulus brown, segments II and III
each usually with an apical or subapical annulus brown, segment IV pale brown. Rostrum brown, seg-
ment IV largely piceous. Thorax and abdomen mainly brown, thoracic sterna, larger granules of thoracic
pleura, median and lateral markings on abdominal sterna, pronotal midline and sublateral tubercles,
pronotal posterolateral spines, integument beneath the 3 patches of erect, brown hairs on pronotum and
base of scutellum piceous. Abdominal laterotergites III-V red-brown, VI and corresponding area of
tergum VII stramineous with narrow basal and apical bands red-brown. Femora basally stramineous,
apically confusedly mottled piceous and various shades of brown; tibiae stramineous with basal, apical
and medial annuli brown; tarsi pale brown. Hemelytra with clavus and corium stramineous to pale brown,
punctures pale brown, corium anteriorly with some large granules on basal parts of major veins piceous,
apical one-third of corium confusedly mottled piceous and various shades of brown; membrane (Fig.
178) largely colourless with more or less discrete piceous markings basally which vary in extent, but
always with some distinct spots between longitudinal veins near basal cells.
REMARKS. The small size and arcuate posterior tibiae could lead to confusion of this species with
C. curvipes, from which it may always be distinguished by the presence of distinct, piceous mark-
ings on the hemelytral membrane which in C. curvipes is evenly suffused pale fuscous. The shape
of the pronotum and the pattern of pubescence on the scutellum also serve to distinguish the two
species. Examination of the form of the tongue of the genital capsule will confirm the identity of
male specimens. Specimens from eastern and southern Africa differ from those from West Africa
in having a slightly narrower head and slightly longer tibiae. In West African specimens the means
of pronotum width, first antennal segment length and first rostral segment length divided by head
width were found to be 2:24, 1-01 and 0-58 respectively, compared with values for eastern and
southern specimens of 2-26, 1-09 and 0-64. The mean value of posterior tibia length divided by
posterior femur length was 0-66 for West African specimens and 0-68 for eastern and southern
specimens.
In his original description of this species, Bergrdéth (1913 : 328-329) apparently mistook the
tufts or brushes of hairs on the scutellum for solid spines. This cannot be verified as the type-
specimen cannot be located at present. According to Grant & Stys (1970 : 113) many of Bergroth’s
types have been recognized among the general collection of the ZMU, Helsinki, so it is possible
that type-material of spiniscutis may one day come to light. For this reason and because the
species is readily recognizable from the original description it seems inappropriate to designate a
neotype.
DISTRIBUTION. A crescent-shaped area connecting Senegal, Kenya, Malawi, Botswana and
Angola. No specimens attributable to this species were found in the extensive collections of
Hemiptera from Zaire held by the IRSNB, Brussels and the MRAC, Tervuren.
MATERIAL EXAMINED
Senegal: 1 3, Nioro, 12.x.1943, on rice (Oryza sativa) (J. Risbec) (BMNH, London). Ivory Coast: | 3,
Lamto, 11.xi.1965 (Mme D. Gillon) (BMNH, London). Ghana: | 3, Tafo, 8.i.1967 (R. Kumar); 1 3,2 9,
Kpandu, 2.xi.1967 (Kumar) (UG, Legon). Nigeria: 2 3, 1 9, Samaru, 8.ix.1954, 12.1.1955, 12.ii.1956, at
grass roots and on cotton (Gossypium) (M. G. Emsley) (BMNH, London). Kenya: | 3, Migori Valley, S.
Kavirondo, 1300 m, 2-4.v.1911 (S. A. Neave); 1 3, 1 9, Kisumu, 1145 m, xi.1920 (A. F. J. Gedye); 1 9,
Nairobi, iv.1936 (Gedye); 1 2, N.F.D., Wajir, 25.i.1955 (/. Lansbury) (BMNH, London). Tanzania: 1 4,
AFRICAN POD BUGS 63
Meru, vii.1943 (Van Someren); 2 3, no detailed data (W. H. Potts) (BMNH, London). Malawi: 1 9,
Mlanje, 5.ix.1913, 700m (S. A. Neave) (BMNH, London). Angola: 1 9, Luimbale, Mt Moco, 1800-
1900 m, iii.1934 (K. Jordan) (BMNH, London). Botswana: 1 3, Moremi Reserve, 19° 23’S, 23° 33’ E,
19.iv.1972 (BMNH Southern Africa Expedition) (BMNH, London).
Clavigralla marmorata sp. n.
(Figs 185, 186)
Length: 3, 7:0-7:7 mm (n = 2). Female unknown. Resembles C. spiniscutis in colour and pubescence but
of a much more slender build, with longer appendages and non-arcuate posterior tibiae. The paratype
lacks antennal segments III and IV.
Head rather short, antennifers slightly divergent ; antennae with segment I very slightly thickened in its
apical one-third, its length divided by width of head including eyes 1-34 in both specimens. Ratio of
lengths of antennal segments I : II : II] : [V as 1-00 : 0-87 or 0-83 : 0-78 : 0-98; segment III slightly longer
than width of head, other segments distinctly longer. Rostrum at rest reaching to disc of metasternum,
ratio of lengths of segments I : II : III : 1V about 1-00 : 0-93 : 0-53 : 0-83; length of basal segment divided
by width of head including eyes 0-64—0-69.
Pronotum rather similar to that of C. spiniscutis but posterior, horizontal lobe behind posterolateral
angles much shorter; width across apices of posterolateral spines divided by width of head including eyes
2:13-2:23. Apex of corium at rest reaching to level of base of abdominal laterotergite VI. Posterior tibia
(Fig. 185) almost straight, its length divided by that of posterior femur 0-76-0-77.
Abdominal sternites III-VII with posterolateral angles projecting as broad, rather short spines. Male
genital capsule with lip rounded, prominent, tongue triangular. Paramere (Fig. 186) moderately broad,
apex curved towards midline of body, very blunt. Conjunctiva as in C. spiniscutis except for the presence
of a pair of small ventrolateral lobes at about the level of the dorsomedian lobe; finger-like lobe on
posterior face of conjunctiva above base of vesica much stouter at its base than that of C. spiniscutis.
Colour and distribution of pubescence as in C. spiniscutis but pubescence everywhere sparser; pubes-
cence of antennae very short; pattern of hairs on pronotum exactly as in C. spiniscutis and that of scutellum
very similar but less compactly concentrated into tufts; colour pattern of hemelytral membrane as in C.
spiniscutis; abdominal laterotergites III-V dark brown with narrow, pale brown, transverse band close to
anterior border of each, VI with a much broader band occupying basal half except for very narrow dark
brown anterior margin, corresponding area of tergite VII pale brown except for narrow, dark brown
bands at anterior and posterior margins.
REMARKS. The small size, dark coloration, non-arcuate posterior tibiae and particularly the very
long antennae distinguish this species from its relatives. The shape of the paramere indicates a
close relationship with C. spiniscutis.
DISTRIBUTION. Known only from Rhodesia and Transvaal.
MATERIAL EXAMINED
Holotype 3, South Africa: Transvaal, Wylies Poort, 3—8.ii.1941 (A. L. Capener) (TM, Pretoria).
Paratype. Rhodesia: 1 3, Salisbury, 1898 (BMNH, London).
The tomentosicollis-subgroup
Tongue of male genital capsule trifid; pronotal disc with a pair of large, blunt, sublateral tubercles;
membrane of hemelytron suffused fairly evenly with brown pigment, only rarely with some indication of
spotting between the veins close to the basal cells.
This is the only subgroup of the genus which contains species found in the Oriental region. The
three Oriental species are C. orientalis, C. gibbosa (the type-species of the genus) and C. scutellaris,
which is also found in Africa.
Clavigralla tomentosicollis Stal
(Figs 187, 188, 191-202)
Clavigralla tomenticollis Stal, 1855 : 31. LECTOTYPE 2, SoutH Arrica (NR, Stockholm), here design-
ated (examined).
Clavigralla tomentosicollis Stal, 1865 : 107. [Emendation.]
Acanthomia tomentosicollis (Stal) Stal, 1873 : 82.
64 W. R. DOLLING
Length: 3, 8-3-9-7 mm, mean 9-1 mm (n = 32); 2, 9:-3-11:5 mm, mean 10-6 mm (n = 30). Form robust,
oblong, deep-bodied.
Head (Fig. 187) anteriorly declivent at an angle of about 45° to the vertical. Antennifers scarcely
divergent, process at outer apical angle broad, deflexed. Bucculae occupying about one-third of length of
ventral midline of head. Antennae with segment I slightly curved laterad and slightly thickened in its
apical third, its length divided by width of head including eyes in male 1-17—1-39, mean 1-26 (n = 30), in
female 1:10-1:32, mean 1-25 (n = 30). Ratio of lengths of antennal segments I : II : II : 1V in male
about 1-00 : 0-85 : 0-72 : 1-03, in female about 1-00 : 0-88 : 0-75 : 1:01. Basal segment of rostrum at rest
(Fig. 187) with apex distinctly posterior to level of posterior margin of eye; length of basal segment divided
by width of head including eyes in male 0:68-0:81, mean 0-73 (n = 30), in female 0-63—-0-83, mean 0-76
(n = 30). Ratio of lengths of rostral segments I : II : III : IV in male about 1-00 : 0-92 : 0-71 : 1-15, in
female about 1-00 : 0-89 : 0-70 : 1-10.
Pronotum (Fig. 188) strongly declivent anteriorly, with a pair of large, blunt, sublateral tubercles;
posterolateral angles bluntly projecting, each bearing a short, slender, laterally directed spine with slight
anteriad and upward curvature; width across apices of posterolateral spines divided by width of head
including eyes in male 2:35-3:10, mean 2-70 (n = 30), in female 2:54-3:18, mean 2-87 (n = 30); pre-
scutellar spines small, strongly divergent. Scutellum with disc very strongly convex. Metathoracic scent
gland peritreme of the usual form for the species-group (Fig. 191). Apex of corium produced, at rest
reaching posteriorly to level of about basal one-third of abdominal laterotergite VI. Anterior and inter-
mediate femora each with a single small, subapical spine beneath, rarely with two; posterior femur (Fig.
192) with two major subapical spines beneath, the more distal about one and one-half times as long as the
other, with about 5 very small spines or granules between them and a terminal series of 4 or 5 spines.
Posterior tibia straight except for slight basal curvature (Fig. 192), its length divided by length of posterior
femur in both sexes 0:68-0:88, mean 0:75 (n = 60).
Abdominal sternites III-VII with posterior angles acute, slightly prominent (Fig. 193). Male genital
capsule (Fig. 194) with lip filling posterior emargination, tongue trifid. Paramere (Fig. 198) with apex
strongly incurved, acute, tooth at middle of blade strongly developed, blade scarcely broader than shaft.
Conjunctiva (Figs 195-197) with dorsomedian lobe low, transverse, simple; distal dorsomedian lobe
larger, transverse, with conical intermediate dorsal lobe arising from its anterior face and short, distal
dorsolateral lobes arising at its lateral angles; posterior face of conjunctiva above base of vesica with a
pair of T-shaped or capitate lobes slightly larger than distal dorsolateral lobes; apical ventral lobes about
equal in size to these capitate lobes; distal ventrolateral lobes large, spreading, lightly sclerotized on their
posterior faces; ventral lobes small, short and finger-like; no lobes, apart from distal ventrolaterals,
sclerotized. Ejaculatory reservoir (Fig. 199) small, wings and straps long, sclerites protecting base of vesica
small and closely associated with ejaculatory reservoir; vesica tightly coiled at base, moderately long.
Female abdominal sternite VII cleft along rather less than half the length of its midline. Second valvula of
ovipositor (Fig. 201) with small, rounded, apical prominence bearing about 10 stout bristles; ventral half
of lateral surface with numerous, finer bristles. Spermatheca (Fig. 202) with duct short, not convoluted.
Sclerites of dorsal wall of gynatrium (Fig. 200) V-shaped, anterior, spreading arms broad in slightly
oblique vertical plane, median, longitudinal arms with half-hastate expansion in horizontal plane.
Head dorsally and laterally granulate, antennal segments I-III minutely granulate. Pronotum with
declivent area, collar and lateral margins granulate, disc and posterior lobe granulate-punctate, with a
few, very small tubercles. Scutellum granulate. Thoracic pleura granulate-punctate. Femora and abdom-
inal laterotergites weakly granulate. Clavus seriately punctate in three to four rows, corium weakly
punctate, in basal half only; anterior veins in basal half with a few, coarse granules.
Head with short, decumbent, tomentose, silvery to golden pubescence throughout, interspersed with
moderately long, suberect, colourless hairs and with a few, long, erect, colourless hairs along dorsal mid-
line. Antennal segments I-IV with short, semidecumbent, colourless pubescence, I also with distinct, short,
decumbent, silvery pubescence, traces of which are also present on II and III. Rostral segments I-IV with
short, suberect, colourless pubescence, I and II also with decumbent, crisped, silvery pubescence. Pro-
notum (Fig. 188) in declivent part with dense pubescence composed of long, tomentose, silvery-white or
slightly yellow hairs and long, erect, pale yellow to pale brown hairs, this area of pale pubescence com-
pletely divided into two halves by a median longitudinal band devoid of tomentose pubescence; posterior,
horizontal lobe of pronotum with darker pubescence, the two areas of different coloured pubescence
sharply demarcated, the junction between them following an undulating, transverse line between postero-
lateral angles of pronotum, two posterior projections of pale pubescence cause this line to undulate
sharply in middle. Hairs of posterior lobe in band immediately adjacent to pale area short, erect, dark
brown; remainder of this area with short, decumbent, crisped, golden-brown pubescence; posterior
margin of pronotum fringed with long, erect, colourless hairs. Scutellum laterally and in posterior two-
AFRICAN POD BUGS 65
Figs 187-193 Clavigralla species. (187) tomentosicollis, head of °, right lateral view; (188) same, 2
pronotum, dorsal view; (189) scutellaris, head of 2, right lateral view; (190) same, 2 pronotum,
dorsal view; (191) tomentosicollis, left metathoracic peritreme, lateral view; (192) same, right
posterior femur and tibia of 3, anterior view; (193) same, left lateral margin of 2 abdominal
sternites III—VII, ventral view.
66 W. R. DOLLING
194
Figs 194-202 Clavigralla tomentosicollis. (194) apical two-thirds of 3 genital capsule, right paramere
removed, dorsal view; (195) conjunctiva and vesica, posterior view; (196) same, dorsal view; (197)
same, left lateral view (broken lines show position of wings and straps of ejaculatory reservoir
complex); (198) left paramere, dorsal view; (199) ejaculatory reservoir complex, left lateral view
with left wing and strap removed, right wing stippled, right strap obscured by body of reservoir;
(200) sclerites of dorsal wall of gynatrium, dorsal view; (201) left second valvula, lateral view; (202)
spermatheca.
thirds with long, erect, colourless and long, tomentose, silvery types of hair intermingled, anteriorly with
two tufts of long, erect, dark brown hairs. Thoracic pleura, thoracic and abdominal sterna and abdominal
laterotergites with short, decumbent, slightly tomentose, silvery pubescence; propleura and thoracic and
abdominal sternites with sparse, short, erect, colourless hairs intermixed with the silvery ones; mesopleura
and metapleura with similar but denser pubescence intermixed; laterotergites with very short, semi-
decumbent, pale to dark brown pubescence. Clavus and corium with very short, sparse, crisped, decum-
AFRICAN POD BUGS 67
bent, silvery to golden pubescence intermixed with very short, semidecumbent to suberect, pale to dark
brown, bristle-like pubescence. Femora throughout and tibiae basally with short, rather sparse, decumb-
ent, silvery to golden-brown pubescence; legs throughout, including these areas, with short, suberect to
rather long, erect, colourless pubescence.
Head pinkish brown, dorsally with numerous granules piceous, laterally and ventrally almost entirely
piceous except for stramineous bucculae. Antennae stramineous, often suffused pink, segment I usually
with a few granules in basal half and a streak or blotch ventrolaterally in apical third piceous, apical
one-quarter usually pinkish brown. Thorax and abdomen pinkish brown; main colour pattern of pro-
notum due to the pubescence, declivent area appearing off-white, completely divided by a dark, longi-
tudinal band along midline where piceous granules are visible; large tubercles on lateral margins and a
few scattered tubercles on disc piceous, projecting through indumentum; posterior lobe of pronotum rich
brown, due to coloration of both surface and pubescence; posterolateral spines piceous or dark brown;
scutellum pinkish brown, disc anteriorly piceous, colour mostly obscured by pubescence; thoracic pleura
each usually with a large, brown or piceous, median macula; acetabula often brown; meso- and meta-
sternum piceous; abdominal sterna with ill-defined stramineous or pale pinkish brown spots contrasting
weakly with pinkish brown ground colour, sterna III—VII each with a small, stramineous spot in the middle
of each lateral margin, sterna III and often IV medially and II-IV, V or VI laterally with poorly defined
piceous areas; laterotergites each with a small, stramineous spot on lateral margin adjacent to stramineous
spot on margin of corresponding sternum. Clavus and corium stramineous, suffused with pink or pinkish
brown especially on veins and on apical one-third of corium, anterior veins of corium in basal half with a
few large granules piceous. Membrane of hemelytron evenly infuscate except for basal cells and veins of
disc which are milky-hyaline. Legs stramineous; coxae and trochanters with large, piceous patches; each
femur with apical half rufous brown with obscurely paler markings, apical area of darker colour where it
borders on paler, basal area with a more or less extensive piceous band, so that junction between the pale
and dark areas is abrupt and very distinct; all femora with a few, small spots in rows in basal halves
piceous (Fig. 192); tibiae usually with basal and apical annuli red-brown and median penannular mark-
ing, interrupted on dorsal surface of tibia, piceous; all tarsal segments infuscate apically, at least beneath.
Egg usually with 5—6 aeromicropyles in a ring.
REMARKS. Within its subgroup, this species is distinguished by its large size and almost straight
posterior tibia from all species except C. scutellaris, from which it may be distinguished by a care-
ful examination of the head shape, length of first rostral segment and details of the pronotal
pubescence. A summary of morphometric variation in C. tomentosicollis and C. scutellaris is
presented in Table 1. C. tomentosicollis is a common pest species throughout most of subsaharan
Africa, though absent from the north-east (where it is replaced by C. scutellaris) and rare in forest
areas including the Congo basin. Most records of ‘Acanthomia brevirostris’ refer to this species.
In economic literature the species is usually recorded as Acanthomia tomentosicollis. English
names are ‘bean bug’ and ‘spiny brown bug’. According to Smit (1964) it is called ‘ysterbek’ in
Afrikaans. A colour figure is given by Bohlen (1973 : 33, fig. 158). Materu (1972) should be con-
sulted for descriptions of the egg and nymphal instars. Recorded host-plants are ‘beans’ (Phaseolus
spp.), cowpea (Vigna), hyacinth bean (Dolichos), chick pea (Cicer), pigeon pea (Cajanus) and
Tephrosia. Publications on the pest status, biology and control include those of Fuller (1922),
Anonymous (1926; 1927), Johnston (1930-1931), Smit (1964 : 134), Swaine (1969), Materu (1970;
1971), Materu & Makusi (1972; 1973), Bohlen (1973) and Taylor & Omoniyi (1972) discuss
host-plant preference, pest status and variation in the morphology of the egg and the same paper
and those of Taylor (1975) and Egwuatu & Taylor (1977a; 1977b) deal with various aspects of
the parasitization of the eggs by Gryon gnidus (Nixon) (Hymenoptera : Scelionidae) and the same
authors (1977c) give a detailed account of the biology of C. tomentosicollis under field and labora-
tory conditions.
DISTRIBUTION. Throughout the African mainland from Senegal and Sudan in the north to Cape
Province in the south; absent from extreme north-eastern part of Ethiopian region (drier parts of
Sudan and Kenya); present in Zanzibar and Comoro Is. but not recorded from other islands.
MATERIAL EXAMINED
Clavigralla tomenticollis Stal, lectotype 2, South Africa: ‘Caffraria’ (J. Wahlberg) (NR, Stockholm).
Numerous specimens from the following localities. Senegal: Bambey. Gambia: Jambour. Ivory Coast:
Bouake. Ghana: Yapi; Tamale; Zuo; Jimle; Zau; Krachi; Abutia; Buriso. Togo: Sambu; Yendi; Sag-
68 W. R. DOLLING
Table 1 Morphometric variation in Clavigralla tomentosicollis Stal and C. scutellaris (Westwood)
Length Width of Length of Length of Length of Size of
(mm) pronotum/ IAS/width of IRS/width of posterior tibia/ sample
width of head head head length of
posterior femur
Males
tomentosicollis 8:8 2°65 1-27 0:73 0:77
South Africa (8:3-9-4) (2:35-2:77) (1:19-1:34) (0°68-0°81) (0-75-0-83) 10
tomentosicollis 9-1 2:66 1-29 0-75 0:75
Central Africa (8-6-9:7) (2-51-2°84) (1:18-1:39) (0:69-0:79) (0:73-0:77) 10
tomentosicollis 9-4 2-79 1:23 0:72 0-72
West Africa (8:8-9:7) (2:56-3:10) (1:17-1:28) (0:68-0:74) (0:70-0:75) 10
scutellaris 9-6 2:69 1-26 0:62 0:72
Africa and Arabia (8-8-10°3) (2:58-2:81) (1:18-1:39) (0°-55-0-66) (0:69-0:77) 9
scutellaris 9-5 2-80 1-31 0-66 0-74
Asia (9:3-9:9) (2:68-2:97) (1:21-1:39) (0°63-0-69) (0:73-0:77) 5
Females
tomentosicollis 10-2 277 1:26 0:77 0:76
South Africa (9-3-11-0) (2:54-2:93) (1:21-1:31) (0:69-0:83) — (0-73-0-80) 10
tomentosicollis 10-4 2:84 1-28 0:77 0:76
Central Africa (9-8-11-1) (2:70-3:05) (1:24-1°32) (0-73-0-81) (0-73-0-88) 10
tomentosicollis 10-6 3°1 1-20 0:74 0-73
West Africa (9:8-11-5) (2-67-3:18) (1:10-1:27) (0:63-0:78) (0-68-0-80) 10
scutellaris i koe 2:96 1-26 0-63 0:72
Africa and Arabia (10-3-12:2) (2:73-3:12) (1:15-1:38) (0:56-0:67) (0-68-0-76) 9
scutellaris 10-5 2:97 1-26 0-66 0-73
Arabia (9-8-11-3) (2-81-3-12) (1:16-1:32) (0:58-0:76) (0-70-0-76) 9
IAS: basal segment of antenna. IRS: basal segment of rostrum. Figures quoted are means, with ranges beneath in
brackets. South Africa: Republic of South Africa only. Central Africa: Zaire, Zambia, Tanzania. West Africa:
Ghana, Togo, Nigeria.
ranga. Nigeria: Ile-Ife; Zaria, Samaru; Samaru; Samaru, Dam; Samaru-Da; Kano, Gumel; Kano;
Katsina Prov.; Malumfashi; Bida; Ibadan. Zaire: Faradje; Gandajika; Lubumbashi; Ngowa; Kwango;
Albertville; Moliro. Sudan: Torit; W. Darfur; Gebel Murra, Killing, W. side of mountain. Uganda:
Kampala; Entebbe; Eastern Prov., Kadunguru. Kenya: Ol Donto Sabuk; Rabai; Mombasa. Tanzania:
Zanzibar ; Ilonga; Arusha; Morogoro; Rukwa Drift, west escarpment; Kilimanjaro, west; Lake Manyara.
Malawi: Zomba; Ruo Valley; Mlanje. Zambia: Niamadzi River, near Nawalia; Mid Luangwa Valley;
Serenje District. Mozambique: Valley of Kola River, near E. Mt Chiperone; Vila Paiva d’Andrada.
Rhodesia: Salisbury; Sawmills; Bulawayo, Siloswe, Matopos; Hope Fountain. Angola: Vila Flor,
Huambo. Botswana: Bakgatla, Sebele. South West Africa: Ovamboland, Oshikango. South Africa:
Transvaal, Rustenburg; Tvl, Warmbaths; Tvl, Warmbad; Tvl, Towoomba; Tvl, 22 km NE. of Pot-
gieterarus; Tvl, base Magoebaskloof; Tvl, Premiermyn; Tvl, Waterval Boven; Tvl, Tierpoort; Tvl,
Pretoria; Tvl, Pretoria, Roodeplaat; Tvl, Silikat nek; Tvl Johannesburg;,Tvl, Barberton; Natal, Lake St
Lucia, Charters Creek; Natal, Port Shepstone; Natal, Scotsbrough; Natal, Zululand, Hedley; Natal,
Zululand, Mtunzini; Natal, Howick; Natal, Tugela River near Weenen; Natal, Estcourt; Natal, Hilton;
Natal, Greytown; Natal, Durban; Natal, Port St John; Natal, Selati, Zoutpansberg; Cape Province,
Ceres; C. P., Krakeel; C. P., Krakeel River; C. P., Barrydale; C. P., Riversdal; C. P., Ladismith; C. P.,
Oudshorn; C. P., Outshorn; C. P., Port Elizabeth; C. P., Knysna District, Harkerville Forest Reserve;
C. P., Alice; C. P., Stellenbosch; C. P., Ongeluks nek; C. P., Queenstown; C. P., Swellendam; C. P.,
Cape Town; C. P., Cape Peninsula, Camps Bay; C. P., Keurboom River; C. P., Du Toits Kloof.
Altitudes recorded on labels: mostly 300-1500 m; one locality in Sudan at 2100 m.
Host-plants recorded on labels: cowpea (Vigna) in Senegal, Nigeria, Tanzania and Botswana;
pigeon pea (Cajanus) in Nigeria, Zaire and Rhodesia; Dolichos lablab Linnaeus in Ghana and
AFRICAN POD BUGS 69
Tanzania; beans (Phaseolus) in Tanzania and South Africa; soya bean (Glycine) in Nigeria;
Tephrosia in Gambia; ‘peas’ in Ivory Coast; unidentified legume in Nigeria; cotton (Gossypium)
in Ghana; spinach (Spinacea) in South Africa and ‘weeds’ in Nigeria.
Habitats recorded on labels: mango orchard, vegetable garden, farm nutwoods and on plant
bases.
Depositories of material: IRSNB, Brussels; NM, Bulawayo; University of Ife, Ile-Ife; UG,
Legon; Duarte coll.; BMNH, London; AMNH, New York; PPRI, Pretoria; TM, Pretoria;
CAS, San Francisco; IAR, Samaru; MRAC, Tervuren; Slater coll.
Clavigralla scutellaris (Westwood)
(Figs 189, 190)
Coreus scutellaris Westwood, 1842 : 24. Lectotype 3, INDIA (UM, Oxford), designated by Dolling (1978 :
291) [examined].
Clavigralla scutellaris (Westwood) Dallas, 1852 : 514.
Acanthomia brevirostris Stal, 1873 : 82. Lectotype 9, SUDAN (NR, Stockholm), designated by Dolling
(1978 : 291). [Synonymized by Dolling, 1978 : 291.]
Length: 3, 8-8-10:3 mm, mean 9:6 mm (n = 14); 2, 9:-8-12:2 mm, mean 10-8 mm (n = 18). Very similar
indeed to C. tomentosicollis. A full account of this species is given by Dolling (1978 : 291-293). Morpho-
metric data for this species are given in Table 1 (above), where C. tomentosicollis is compared. Additional
data for African and Arabian specimens are as follows. Ratio of lengths of antennal segments I : II : III :
IV in males about 1-00 : 0-84 : 0-71 : 1:05, in female about 1-00 : 0-85 : 0-73 : 1-03. Ratio of lengths of
rostral segments I : II : III : Vin male about 1-00 : 0-91 : 0-76 : 1-19, in female about 1-00 : 0:92 : 0-76 :
1-16.
Head (Fig. 189) more strongly declivent than that of C. tomentosicollis, frons making an angle of about
30° with the vertical. Rostrum shorter, first segment at rest with apex reaching posteriorly to a point
distinctly anterior to posterior margin of eye (see Fig. 189 and Table 1). Stal (1873 : 82) states that the
antennae are shorter and the posterolateral angles of the abdominal sternites less prominent in this species
than in C. tomentosicollis. These differences are not substantiated by the larger number of specimens now
available.
Sculpture, pubescence and coloration as in C. tomentosicollis except that antennal segment I is almost
uniform pinkish brown, median annuli of tibiae are evanescent and area of pale pubescence on pronotum
is divided by a dark band for only its anterior half to two-thirds (Fig. 190) and extends posteriorly in a
single median undulation.
REMARKS. This species is extremely close to C. tomentosicollis, from which it may be distinguished
by the steeper declivity of the frons, the shorter rostrum and the slight difference in pronotal
pubescence mentioned above. No differences between the two species could be discerned in either
the male or female genitalia. Most records of ‘Acanthomia brevirostris’ in the literature, including
that of Johnston (1930), refer to C. tomentosicollis. | have seen no specimens attributable to this
species from localities west of Sudan. Nixon’s (1943) record of the egg parasite Microphanurus
striaticeps Dodd (Hymenoptera: Scelionidae), from ‘Acanthomia brevirostris’ in Sudan could
refer to either species; its hosts also include Pentatomidae. Authentic host-plant records are
Dolichos lablab Linnaeus in South Yemen and Cajanus cajan (Linnaeus) Millspaugh in India.
DISTRIBUTION. A crescent-shaped area from Kenya through Arabia and Pakistan to western
India.
MATERIAL EXAMINED
Coreus scutellaris Westwood, lectotype ¢, India: Gogo (UM, Oxford). Acanthomia brevirostris Stal, lecto-
type 9, Sudan: Chartum ( = Khartoum) (Schauf) (NR, Stockholm).
Kenya: | 2, Marsabit Forest, 19.viii.1977, by sweeping (J. H. Martin); 1 3, Northern Frontier District,
Wajir, 25.i1.1955 (/. Lansbury); 1 3, S. Nyeri, vi.1949 (van Someren); 1 2, Lukenia, iii.1937 (van Someren)
(BMNH, London). Sudan: 1 3, no precise locality, 16.iii.1926 (G. R. F. Medani, F. G. S. Whitfield); 2 9,
Berber Prov., Lulua, 1908 (H. H. King); 3 3, 1 9, Ed Damer, 5—10.vii.1961 (R. Linnavuori) (BMNH,
London). Southern Yemen: 3 3, 3 9, El Kod, xii.1957, on Dolichos lablab Linnaeus (R. C. M. Darling);
1 9, Wadi Dareija, SW. of Dhala, c. 1400 m, 6-9.xi.1937 (H. Scott, E. B. Britton) (BMNH, London).
Oman: 1 2, Oman, Al Wafi, 18.iii.1976 (K. M. Guichard) (BMNH, London).
70 W. R. DOLLING
Extralimital specimens. Afghanistan: Nengrahar Province. Pakistan: Hangu, Hyderabad, Karachi.
India: Hyderabad (A. P.), Bombay, Coimbatore, Thekkadi, Trivandrum, Chickballapur. (Moravské
Museum, Brno; Karachi University; BMNH, London. For details see Dolling, 1978 : 293.)
Clavigralla annulipes Signoret comb. rev.
(Figs 203-208)
Clavigralla annulipes Signoret, 1860 : 943. Holotype, MADAGASCAR (‘Musée de Naples’, fide Bergréth,
1903 : 297) [not examined].
Acanthomia annulipes (Signoret) Bergréth, 1903 : 297.
Length: 3, 9:-4-10-:1 mm, mean 9-7 mm (n = 12); 9, 10-0-11:3 mm, mean 10:4 mm (n = 11). Similar in
size and build to C. tomentosicollis, but differing in many details.
Antennifers almost parallel; head shape similar to that of C. tomentosicollis. Length of antennal seg-
ment I divided by width of head including eyes in male 1 -08—1-:28, mean-1-18 (n = 12), in female 1-14~1-22,
mean 1-17 (n = 11). Ratio of lengths of antennal segments I : II : HI : IV in male about 1-00 : 0-82 :
0-74 : 1-11, in female about 1-00 : 0-81 : 0-72 : 1-03. Length of basal segment of rostrum divided by width
of head including eyes in both sexes 0:66-0:75, mean 0:69 (n = 6). Ratio of lengths of rostral segments
1:11: 111: IV in male about 1-00 : 1-06 : 0-84 : 1-32, in female about 1-00 : 0-94 : 0-69 : 1-21.
Pronotum (Fig. 203) strongly declivent, posterolateral angles not prominent, each bearing a short,
slightly posteriorly directed spine, disc with a few, short, blunt, projecting tubercles; posterior lobe with a
low, transverse keel in middle, bisected by a more prominent longitudinal keel; width across apices of
posterolateral spines divided by width of head including eyes in male 2:50-2:75, mean 2°63 (n = 12), in
female 2:55-2:76, mean 2:65 (n = 8). Scutellum convex, midline depressed, apex slightly produced.
Corium with apex slightly produced, at rest reaching posteriorly to level of suture between laterotergites
V and VI. Posterior tibia distinctly arcuate in basal third, its length divided by that of posterior femur
0:69-0:74, mean 0-71 (n = 6).
Abdominal sternites III-VII with posterolateral angles acute, prominent (Fig. 204). Male genital
capsule similar to that of C. tomentosicollis. Paramere (Fig. 205) rather similar to that of C. tomentosicollis
but apex distinctly angled inwards, not curved. Conjunctiva (Figs 206-208) with dorsomedian lobe low,
M-shaped, intermediate dorsal lobe conical, distal dorsomedian lobe obsolete, distal dorsolateral lobes
rather large, almost globular, a pair of large, T-shaped lobes arising from posterior face of conjunctiva
above origin of vesica, apical ventral, distal ventrolateral and ventral lobes similar in form to those of C.
tomentosicollis; all lobes membranous. Vesica rather short. Female genitalia as for C. tomentosicollis.
Sculpture, pilosity and coloration generally as in C. tomentosicollis. Antennae with segment I entirely
brown. Membrane of hemelytron evenly suffused with brown pigment except for colourless basal cells.
Area of brown pubescence on posterior lobe of pronotum almost completely divided into two triangular
areas by a band of pale, silvery hairs extending posteriorly from declivent area along raised midline of
posterior lobe and enclosing a tuft of erect, brown hairs between the apices of the two lateral triangles.
Scutellum with erect, slightly tomentose, pale golden to silvery pubescence, in an almost globular arrange-
ment, very slightly embrowned anteriorly. Tibiae without median pigmented annuli, or annuli present
but faint.
REMARKS. Within the tomentosicollis-subgroup this species seems to be more closely allied to
C. curvipes and C. simillima than to C. tomentosicollis and C. scutellaris by reason of the distinctly
curved posterior tibiae and the pattern of pronotal pubescence. It differs from C. curvipes and C.
simillima and from the only other Malagasy species of the tomentosicollis-group, C. pusilla, by its
larger size, and from the last named species by the even pigmentation of the hemelytral membrane.
DISTRIBUTION. Southern Madagascar.
MATERIAL EXAMINED
Madagascar: 8 3, 10 9, Région Sud de I’Ile, Bekily, vi.1936 (A. Seyrig) (MNHN, Paris and BMNH,
London); 3 3, 2 2, Ampandrandava (Seyrig) (MNHN, Paris); 1 3, Région du Sud, Andrahomana, 1901
(Ch, Alluaud) (MNHN, Paris).
Clavigralla curvipes (Stal) comb. n.
(Figs 211, 213-217)
Acanthomia curvipes Stal, 1873 : 82. LECTOTYPE 3, GuINgeA-BissAu (NM, Vienna), here designated
[examined].
AFRICAN POD BUGS
71
203
204 205
208 \
207
Figs 203-208 Clavigralla annulipes. (203) pronotum of 2, dorsal view; (204) left lateral margins of 3
abdominal sternites III—-VII, ventral view; (205) left paramere, dorsal view; (206) aedeagus, dorsal
view; (207) same, left lateral view; (208) same, ventral view.
Length: 3, 7-3-8-:3 mm, mean 7:8 mm (n = 9); 9, 7-6-8:5 mm, mean 8:0 mm (n = 8). A small species,
similar in build to the much larger C. tomentosicollis but with posterior tibia strongly arcuate at base,
abdomen proportionally narrower and pubescence of pronotum differently organized.
Head short, antennifers parallel to slightly divergent. Antennal segment I (Fig. 211) slender, narrowest
at a point about one-third of its length from base, then gradually increasing in thickness towards apex;
length of this segment divided by width of head including eyes in male 1:03-1:13, mean 1-09 (n = 9), in
female 1-00-1-06, mean 1-02 (n = 8). Ratio of lengths of antennal segments I : II : II] : 1V in male about
1-00 : 0-78 : 0-70 : 0-94, in female about 1-00 : 0-80 : 0:72 : 0-96. Length of basal segment of rostrum
divided by width of head including eyes in both sexes 0:63-0:73, mean 0:67 (n = 16). Ratio of lengths of
rostral segments I : II : III : IV in both sexes about 1-00 : 0:90 : 0-49 : 0:86; apex of rostrum at rest
reaching disc of mesosternum.
72 W. R. DOLLING
t
ee
ANG NW 209
Ie
U
Figs 209-218 Clavigralla species. (209) simillima, holotype 3, pronotum and scutellum, dorsal view;
(210) same, first antennal segment of 2, dorsal view; (211) curvipes, first antennal segment of 9,
dorsal view; (212) simillima 9, right posterior femur and tibia, posterior view; (213) curvipes, left
paramere, dorsal view; (214) same, conjunctiva and vesica, left lateral view, outline of left wing of
ejaculatory reservoir complex dotted; (215) same, left lateral margin of 2 abdominal sternites III-
VII, ventral view; (216) same, second valvula, lateral view; (217) same, sclerites of dorsal wall of
gynatrium, dorsal view; (218) simillima, sclerites of dorsal wall of gynatrium, dorsal view.
213
Pronotum with posterolateral spines slender, directed laterally, slightly upcurved, width across apices
of spines divided by width of head including eyes in male 2:24-2:72, mean 2°52 (n = 5), in female 2-39-
2:64, mean 2:46 (n = 7). Scutellum rather strongly convex. Corium with apex produced, at rest reaching
posteriorly to level of basal one-third of laterotergite VI. Posterior tibia strongly curved at base, its length
divided by that of posterior femur 0:63-0:68, mean 0-66 (n = 17).
Abdomen with posterolateral angles of sternites III-VII (Fig. 215) acute, slightly projecting. Male
genital capsule with lip rounded, tongue apically trifid. Paramere (Fig. 213) with strong tooth near middle
of blade on dorsal surface and with apex strongly curved inward. Ejaculatory reservoir with wings and
straps long; vesica of moderate length. Conjunctiva (Fig. 214) with dorsomedian lobe low, rounded, a
pair of finger-like lobes arising from its base posterolaterally; intermediate dorsal lobe high, rather
narrowly conical; distal dorsomedian and distal dorsolateral lobes obsolete; a large, downcurved, median
lobe arising near top of posterior face of conjunctiva; apical ventral lobes large, with distinct surface
roughness; distal dorsoventral lobes of the form usual in the genus; ventral lobes absent. Female with
tergum IX distinctly but not strongly deflexed at apex. Second valvula of ovipositor (Fig. 216) bearing a
prominent dorsal tooth. Sclerites of dorsal wall of gynatrium (Fig. 217) U-shaped, median arms with
broad distal expansion. Spermatheca with bulb narrowly lunate, less than three times as wide as main part
of duct, which is attached subterminally to it and is not convoluted but simply U-shaped; total length of
duct about twice that of bulb, length of broad portion about two-thirds as long as bulb.
AFRICAN POD BUGS a3
Sculpture and distribution of pubescence generally as for C. tomentosicollis. Junction between anterior,
pale and posterior, dark areas of pronotal pubescence strongly M-shaped (as in C. simillima, Fig. 209),
with an isolated tuft of erect, brown hairs in the triangle of pale pubescence between the anterior angles of
the ‘M’; pale pubescence extending posteriorly in a band along midline of posterior lobe to its posterior
margin. Scutellum with two tufts of dark brown pubescence anteriorly on disc, otherwise with long,
slightly tomentose, white pubescence shaped roughly into a hemisphere, not organized into spine-like
tufts. Coloration in general as in C. tomentosicollis but usually with brown or piceous markings at apical
border of corium more pronounced. Membrane of hemelytron suffused with brown, especially on disc,
occasionally pigment also concentrated into 1 or 2 darker brown spots between bases of longitudinal
veins where these arise from basal cells; veins themselves colourless except for bases of 2 longitudinal
veins entering membrane from corium, which are dark brown.
Ovarian egg with 3 (in two cases) or 4 (in six cases) aeromicropyles.
REMARKS. The small size, pronotum shape, basally arcuate posterior tibiae and coloration dis-
tinguish this species from all others except C. simillima, which differs chiefly in the shape of the
first antennal segment and the sclerites of the dorsal wall of the gynatrium.
DISTRIBUTION. This species occupies a band of territory between latitudes 15° N and 3°S from
Guinea-Bissau to Uganda.
MATERIAL EXAMINED
Acanthomia curvipes Stal, lectotype 3, Guinea-Bissau: ‘Bissao’ (in Signoret coll.) (NM, Vienna).
Ghana: 1 3, Legon, 14.viii.1964 (S.A. Whyte); 1 2, Legon, 29.xi.1968; 1 3, Legon, 1.ix.1968 (R. Kumar);
3 3, 1 2, Legon, 6.xii.1968 (Kumar); 1 3, 1 2, Tafo, 1.iv.1966, 3.i.1967, by pyrethrum kndéckdown from
cocoa; 1 9, Tafo, 22.iv.1967; 1 9, Tafo, on ground, 15.ii.1956 (D. Leston); 1 3, Tafo, 19.i.1966, in u.v.
light-trap (Leston); 1 ex. without abdomen, Keta, 8.xi.1967 (Leston); 1 3, Wegbe, V. R., 1.xi.1967 (Leston)
(UG, Legon). Togo: 1 3, Missahoué, 650 m, vi.1963 (Mme Y. Schach) (MRAC, Tervuren). Nigeria: 2 3,
N. Nigeria, Gusau, 1.ix.1975, at u.v. light (L. E. Wallace, J. C. Deeming) IAR, Samaru); 1 3, 1 9, Bida,
on weeds, 22.ix.1955; 1 9, N. W. State, 18 km NW. of Mokwa, Zugurma, in riparian rainforest by River
Eku, 11-19.viii.1970 (P. H. Ward) (BMNH, London). Zaire: 2 3, 2 2, Faradje, 29° 40’ E, 3° 40’ N, i.1913
(Lang, Chapin) (AMNH, New York). Uganda: 1 3, Northern Buddu, 1160 m, 16—18.ix.1911 (S. A. Neave);
1 $, Kampala, 2-20.iv.1918 (C. C. Gowdey); 2 9, Kawanda, 11.iv.1940, on Desmodium adscendens De
Candolle and 14.iv.1940 on ‘lulimilumw’ (H. H.) (BMNH, London). Rwanda: 1 3, Rubona, 1.v.1963 (G.
Pierrard) (MRAC, Tervuren).
Clavigralla simillima sp. n.
(Figs 209, 210, 212, 218)
Length: 3, 7-2-8-1 mm, mean 7:8 mm (n = 5); 2, 7-8-8:5 mm, mean 8-1 mm (n = 9). Very similar
indeed to C. curvipes.
Antennal segment I (Fig. 210) rather stout, of almost uniform thickness in the basal half, then expanding
slightly and gradually towards apex. Length of antennal segment I divided by width of head including eyes
in male 1-00-1-21, mean 1:09 (n = 5), in female 1:01-1:14, mean 1:09 (n = 9); ratio of lengths of segments
I: II : I : [Vin male about 1-00 : 0-78 : 0-70 : 1-00, in female about 1-00 : 0-77 : 0-68 : 0-91. Length of
rostral segment I divided by width of head including eyes in both sexes 0:55-0:71, mean 0°65 (n = 14);
ratio of lengths of segments I : II : If] : IV about 1-00 : 0-91 : 0-48 : 0-87.
Width of pronotum across apices of posterolateral spines divided by width of head including eyes in
male 2:57-2:65, mean 2:61 (n = 4), in female 2:44-2:75, mean 2:64 (n = 7). Length of posterior tibia
(Fig. 212) divided by length of posterior femur 0:63-0:69, mean 0:65 (n = 14).
Male genitalia as in C. curvipes but vesica on average shorter. Second valvula of ovipositor with dorsal
tooth less prominent than in C. curvipes and sclerites of dorsal wall of gynatrium (Fig. 218) with distal
expansions of median arms less pronounced.
Pattern of pubescence on pronotum and scutellum (Fig. 209) and elsewhere and coloration and sculpture
of integument as in C. curvipes.
REMARKS. This is clearly a sister-species to C. curvipes, differing little from it in appearance and
morphology. There is a slight tendency to sexual dimorphism in the length of antennal segment
IV, a phenomenon not encountered in C. curvipes, and the pronotal posterolateral spines are on
average slightly longer. The differences in the form of the sclerites of the gynatrium are not as
clear-cut as Figs 217 and 218 would suggest, since the median arms of these sclerites are broad in
74 W. R. DOLLING
the sagittal plane and a slight alteration in the angle from which they are viewed can result in a
pronounced change of outline. Other characters of the genitalia do not provide reliable characters
for separation of the species unless several specimens are available for comparison. However, the
thickness of the first antennal segment is a reliable diagnostic character. It may be that the two
populations treated here as species are not separated by a fertility barrier but only by the physical
barriers of the Congo rainforest and the highlands of eastern central Africa.
DISTRIBUTION. The eastern half of southern Africa from Tanzania to Natal.
MATERIAL EXAMINED
Holotype 3, Tanzania: E. Rukwa, 3500 ft (1100 m), iv.1938 (D. G. MacInnes) (BMNH, London).
Paratypes. Tanzania: | 2, data as holotype (BMNH, London). Zaire: 2 9, Lubumbashi, 9.iii.1939,
30.iv.1939 (H. J. Brédo) (IRSNB, Brussels). Malawi: 1 3, SW. Lake Malawi, Monkey Bay, 2.v.1966 (NM,
Bulawayo). Mozambique: 1 3, Chiluvo Hills, Vila Machado, 31.x.1967 (E. Pinhey) (NM, Bulawayo); 1
ex. without abdomen, Luabo, lower Zambesi River, viii.1957 (P. J. Usher) (TM, Pretoria). South Africa:
1 3, ‘Stella Bush’, xi.1905 (Mailer); 1 2, no data (probably leg. Mailer); 1 3, Natal, Durban (/. P. Cregoe);
1 9, Natal (Bell Morley); 1 2, Natal, Umbilo, 16.x.1914 (L. Bevis); 1 2, Natal, Port Shepstone, v.1897;
2 9, Natal, Malvern, vi.1897, 10.vi.1897; 1 2, Natal, Umkomaas River, 1897 (BMNH, London).
The wittei-subgroup
Tongue of male genital capsule bifid; pronotal disc with a pair of large, blunt, sublateral tubercles;
membrane of hemelytron with a row of discrete, piceous spots, sometimes very pale, between bases of
longitudinal veins. Pronotum with posterolateral angles more prominent that in other members of the
tomentosicollis-group.
Clavigralla wittei (Schouteden) comb. n.
(Figs 219-221, 223-228)
Acanthomia wittei Schouteden, 1938 : 291-292. LECTOTYPE 64, ZatrE (MRAC, Tervuren), here
designated [examined].
Length: 3, 7:-6-9:5 mm, mean 8:5 mm (n = 48); 2, 8:3-10:2 mm, mean 9:2 mm (n = 56). Of a rather
more slender form than C. tomentosicollis, less deep-bodied, with more prominent pronotal posterolateral
angles.
Antennifers slightly divergent. Length of antennal segment I divided by width of head including eyes in
male 0:86-1:12, mean 0:99 (n = 46), in female 0:81-1:09, mean 0-95 (n = 56); ratio of lengths of segments
I : II : HI : [Vin male about 1-00 : 0-75 : 0-72 : 1-32, in female about 1-00 : 0-76 : 0-73 : 1:28. Length of
basal segment of rostrum divided by width of head including eyes in both sexes 0:62-0:81, mean 0-70
(n = 103); ratio of lengths of segments I : II : III : IV about 1-00 : 0-93 : 0-55 : 0-89.
Pronotum (Fig. 220) strongly declivent, posterolateral angles prominent, posterolateral spines slender,
curved slightly upwards and slightly anteriad, usually long; margins of pronotum behind spines with a
few prominent tubercles; width across apices of posterolateral spines divided by width of head including
eyes in male 2:31-3:14, mean 2:80 (n = 42), in female 2-47—3-32, mean 2:87 (n = 48). Scutellum strongly
convex. Corium with apex distinctly produced (Fig. 219), at rest reaching posteriorly to about level of
middle of laterotergite VI. Anterior femur without subapical spines beneath or rarely with a single, very
small one; intermediate femur usually with a single, small, subapical spine beneath, which is sometimes
absent; posterior femur usually with 2-4 small spines between the two major spines. Posterior tibia
almost straight, its length divided by that of posterior femur 0-68-0-79, mean 0-74 (n = 97).
Abdominal sternites III-VII with posterolateral angles (Fig. 224) produced into short, triangular spines.
Male genital capsule (Fig. 221) with lip short, tongue deeply bifid, each half of tongue laminar, distinctly
deflexed apically and laterally. Paramere (Fig. 225) strongly curved inward at apex, dorsomedial tooth of
blade well developed, blade rather broad. Conjunctiva (Fig. 223) with dorsomedian lobe bearing a pair of
small, finger-like lobes arising from its posterior face; intermediate dorsal and distal dorsomedian lobes
low, conical; distal dorsolateral lobes absent; apical ventral lobes large, distal ventrolateral lobes of the
form usual in the genus; ventral lobes poorly developed; posterior face of conjunctiva without prominent
lobes. Ejaculatory reservoir complex with wings and straps long. Vesica very long, sclerites at its base
large. Female sternum VII cleft for about half its median length. Second valvula (Fig. 226) with dorsal
margin strongly undulating and apex produced into a lobe which is more weakly sclerotized than main
body of valvula. Spermatheca (Fig. 228) with duct long, narrow portion convoluted. Sclerites of dorsal
wall of gynatrium (Fig. 227) with large, anterodorsally directed tooth on each median arm.
AFRICAN POD BUGS 15
ee ee 221
: : | 222 |
\ 225
a, 224
227
226
228
Figs 219-228 Clavigralla species. (219) wittei 3, apical half of left hemelytron, dorsal view; (220)
same, pronotum of 3, dorsal view; (221) same, apical half of 3 genital capsule, dorsal view, with
parameres removed; (222) neavei, holotype 3, apical half of genital capsule with parameres re-
moved, dorsal view; (223) wittei, conjunctiva and vesica, left lateral view, outline of left wing of
ejaculatory reservoir complex dotted; (224) same, left lateral margin of 2 abdominal sternites
III-VII, ventral view; (225) same, left paramere, dorsal view; (226) same, second valvula, lateral
view; (227) same, sclerites of dorsal wall of gynatrium, dorsal view; (228) same, spermatheca.
Pronotum and thoracic pleura granulate-punctate; head, antennae, legs, sterna and abdominal latero-
tergites weakly to very weakly granulate; major veins of clavus and corium with a few, small granules;
clavus with three rows of rather small but deep punctures, corium near base weakly punctate. Abdominal
terga I-VI and base of VII with fine, reticulate sculpture, I coarsely, and II finely, transversely furrowed,
central one-third of III and IV with a few moderate sized to large punctures, III laterally and [V-VII with
numerous, small punctures.
Head dorsally with sparse, long, erect, colourless pubescence, everywhere with fairly dense, short,
decumbent and semidecumbent, silvery pubescence; antennal segment I with similar, short pubescence
and some longer, suberect to erect hairs one-half to one times the width of the segment, II and III with
semidecumbent pubescence about one-third as long as width of respective segments. Pronotum (Fig.
76 W. R. DOLLING
220) in declivent, anterior part with long, sparse, erect, colourless pubescence, shorter, brown, erect
pubescence arising from granules in midline, a pair of pale to dark brown tufts on disc behind sublateral
tubercles and of comparable size to these, and everywhere with dense, short to moderately long, slightly
tomentose, silvery pubescence which extends onto anterior faces of the prominent posterolateral angles
and is produced posteriorly into two tufts flanking midline; posterior, horizontal lobe of pronotum with
dense, moderately long, brown pubescence which is erect anteriorly and decumbent posteriorly and later-
ally and with a poorly developed stripe of short, decumbent, silvery pubescence in part of midline.
Scutellum with pubescence long, dense, slightly tomentose, silvery laterally and posteriorly, forming two
brown tufts on disc anteriorly. Thoracic pleura with sparse, long, erect, colourless pubescence and with
dense, shorter, slightly tomentose, silvery pubescence. Thoracic sterna with fairly dense, short, decumbent,
crisped, silvery pubescence. Femora and coxae with fairly dense, short, decumbent, silvery pubescence,
legs throughout with long, colourless, suberect to erect pubescence. Clavus, corium and abdominal
laterotergites with dense, short, crisped, semidecumbent, brown and silvery hairs intermingled. Abdom-
inal sternites with sparse, moderately long, suberect, colourless pubescence mixed with much denser and
shorter, decumbent, slightly tomentose, silvery pubescence.
Coloration generally stramineous, variously suffused with pale pinkish brown, darker markings very
variable in intensity and extent. Head typically with ocellar tubercles, ventral surface and most of lateral
surfaces, apex of rostrum and a basal streak on lateral surface of antennal segment I brown to piceous.
Anterior midline, tubercles and posterolateral angles and spines of pronotum, thoracic sterna, anterior
half of disc of scutellum and two oblique streaks on each of the thoracic pleura piceous, posterior lobe of
pronotum brown. Coxae, trochanters and femora except basal halves of intermediate and posterior pairs
mottled with brown and piceous markings; tibiae usually with brown to piceous annuli at base, apex and
middle. Corium with brown spotting or mottling on anterior veins and in apical one-third; hemelytral
membrane (Fig. 219) milky hyaline with 2-4 discrete, brown spots between bases of longitudinal veins,
bases of two veins where they enter membrane from corium brown; sometimes other, vaguer brown
markings present at base of membrane. Abdominal sternites in middle with longitudinal markings and
laterally with oblique stripes piceous or brown and variable in extent; laterotergites and corresponding
area of tergite VII anteriorly each with a narrow, brown to piceous band, followed by a stramineous band
which is narrow on segments III-V and broad on VI and VII, behind which is a broad, brown or piceous
band continuous to posterior margin of each segment. Abdominal tergites varying from largely piceous
with only a large spot above each abdominal scent-gland rudiment, a pair of lateral markings on tergites
VI and VII and a narrow posterior border of VI and posterior half of VII stramineous, to a condition
where tergites III-VII are largely stramineous or pale brown with piceous markings restricted to lateral
areas of each tergite. Sometimes almost the entire ventral and lateral surfaces of the insect are piceous or
black.
Ovarian egg with chorion strongly reticulate, each polygon of reticulation separately convex; about
eight aeromicropyles present.
REMARKS. The posterolateral angles of the pronotum are more prominent in this species than in
any other of the tomentosicollis-group. It is the most common species with distinct spots on the
membrane of the hemelytron. Recorded host-plants which may be food-plants are Schotia
(Leguminosae, Caesalpiniaceae) and Vigna.
DISTRIBUTION. Widespread in western, eastern and southern Africa, less common in central
Africa and apparently absent from South Africa except the Transvaal.
MATERIAL EXAMINED
Acanthomia wittei Schouteden, lectotype 3, Zaire: Stanleyville ( = Kisangani), v.1926, on Schotia (Lt. J.
Ghesquiére) (MRAC, Tervuren).
Ivory Coast: 1 3, Orumbe-boka, 11.x.1964 (Gillon); 2 3, 2 9, Man, 19.x.1968 (J. Breniére) (BMNH,
London). Ghana: 2 3, Tafo, 13.xi.1965 (Leston) (BMNH, London); 1 3, 1 9, Tafo, 8.1.1967 (R. Kumar);
2 3, 1 2, Legon, Botanic Garden, 14.v.1967 (UG, Legon). Nigeria: 1 9°, Ibadan, 21.i11.1955, on Zea mays
Linnaeus leaf (G. H. Caswell) (BMNH, London); 2 3, 3 2, Zaria Province, Dumbi Wood, 26.x.1969,
15.xi.1969, 17.xi.1969 (J. C. Deeming); 1 °, Zaria, Samaru, 3.x.1971 (Deeming) (IAR, Samaru). Zaire:
1 3, Lake Tanganyika, Baie de Burton, 17.iv.1947 (Miss. Tanganyika); 1 2, near Mwashia, grassy plain,
1 km from saltpan, 3.v.1939 (H. J. Brédo) (IRSNB, Brussels); 1 3, Yangambi, ix—xii.1958 (J. Decelle); 1 9,
Lulua, Muteba, i.1932 (G. F. Overlaet); 1 3, 2 2, Lulua, Kapanga, xii.1932 (F. G. Overlaet) (MRAC,
Tervuren). Uganda: 12 3, 14 9, Kafu River, near Hoima, Kampala Road, 1050 m, 29-31.xii.1911 (S. A.
Neave); 1 2, Entebbe, 5-9.iv.1914 (C. C. Gowdey) (BMNH, London). Kenya: 1 3, Mombasa Island,
AFRICAN POD BUGS DE
Kilindini, 19-23.viii.1955 (L. F. Brown); 1 3, Mombasa, iv.1930 (R. E. Dent); 1 3, Maramas District,
lala, 22 km E. of Mumias, 1350 m, 18-21.vi.1911 (S. A. Neave); 1 3, 2 2, Naivasha, vii.1937 (H. J. A.
Turner) (BMNH, London). Tanzania: 2 3, 1 2, Mbeya Mt, 33° 25’ E, 8° 48’ S, 2100 m, trees and herbage
on grass slopes, 5.viii.1959 (Cambridge E. African Exped.); 1 2, Elton Plateau, 2100 m, 34° 10’ E, 9° 20’ S,
7.viii.1959 (Cambridge E.. African Exped.) (BMNH, London). Malawi: 1 2°, Zomba (H. S. Stannus); 2 3,
4 2, Mlanje, 23.iv.1913, 2.vi.1913, 13.vi.1913 (S.A. Neave) (BMNH, London). Botswana: | °, Batawana,
Totena, 9.v.1968, on cow pea (Vigna sp.) (R. E. Roome) (BMNH, London). Zambia: 1 3g, Mweru-
Wantipa, 1944 (H.J/. Brédo) (IRSNB, Brussels). Rhodesia: 3 2, Bindura, ii.1959 (NM, Bulawayo). South
Africa: 15 3, 15 2, Transvaal, Waterburg District, Plat River, 6-18.iv.1905 (C. Swierstra) (TM, Pretoria).
Clavigralla neavei sp. n.
(Fig. 222)
Length: 3, 6-75-7-0 mm (n = 2). Female unknown. Very similar to C. wittei but smaller and with pro-
notal posterolateral angles less prominent.
Length of antennal segment I divided by width of head including eyes 0-89-0:94 (n = 2); length of
segments II and III divided by length of segment I in holotype 0-74 and 0-64 respectively, these segments
missing in paratype, IV missing in both specimens. Length of basal segment of rostrum divided by width
of head including eyes 0-68-0-73 (n = 2).
Pronotum with posterolateral angles less prominent than in C. wittei and spines shorter, width across
apices of spines divided by width of head incuding eyes 2:45-2:52 (n = 2). Length of posterior tibia
divided by that of posterior femur 0:73-0:78 (n = 2).
Posterolateral angles of abdominal sternites less strongly produced than the average for C. wittei. Lip
of male genital capsule (Fig. 222) more prominent than in C. wittei and tongue narrower, the two lobes
cylindrical, apically rounded, the sinus between them U-shaped, not V-shaped as in C. wittei.
Other features, including sculpture, pubescence and coloration, as for C. wittei.
REMARKS. Could be confused with small specimens of C. wittei but distinguished by the form of
the tongue of the male genital capsule.
DISTRIBUTION. Central Africa.
MATERIAL EXAMINED
Holotype 3, Uganda: Entebbe, 12-20.1.1912 (S. A. Neave) (BMNH, London).
Paratype. Zaire: 1 3, Lubero, 14.viii.1932 (L. Burgeon) (MRAC, Tervuren).
Clavigralla alpica (Bergroth) comb. n.
(Figs 229, 231, 233, 235, 236, 238-242)
Acanthomia alpica Bergréth, 1927 :10. LECTOTYPE 92, Zaire (NR, Stockholm), here designated
[examined].
Length: 3, 7:1-7-7 mm, mean 7:3 mm (n = 9); 2, 7:3-8:3 mm, mean 7:7 mm (n = 12). Form depressed
in comparison with other species of the genus.
Antennifers slightly divergent. Antennae short, length of segment I divided by width of head including
eyes in male 0:64-0:69, mean 0-66 (n = 9), in female 0:59-0:72, mean 0-64 (n = 12). Ratio of lengths of
segments I : II : III : [V in male about 1-00 : 1-05 : 0-95 : 1-54, in female about 1-00 : 1-08 : 0-98 : 1-50;
segment I shortly fusiform. Length of basal segment of rostrum divided by width of head including eyes
in both sexes 0:68-0:76, mean 0-73 (n = 18), ratio of lengths of segments I : II : III : 1V about 1-00:
0-93 : 0°51 : 0-77.
Pronotum (Figs 229, 231) strongly declivent despite dorsoventral compression of body, its postero-
lateral angles strongly produced as in C. wittei but bearing much shorter spines; width across apices of
posterolateral spines divided by width of head including eyes in male 2:33-2:55, mean 2-47 (n = 8), in
female 2:28-2:75, mean 2-45 (n = 12). Scutellum (Fig. 231) weakly convex. Corium with apex produced,
at rest reaching posteriorly to level of middle of laterotergite VI. Anterior and intermediate femora with-
out or each with a very small subapical spine beneath, posterior femur usually with two major spines, the
more proximal about half the length of the other, and an apical series of four spines; rarely 1 or 2 small
spines are present between the major spines of the posterior femur and in one abnormal specimen
examined there was an additional rather large spine just proximal to the first major spine. Posterior tibia
not curved at base, its length divided by length of posterior femur 0:78-0:86, mean 0-82 (n = 15).
78 W. R. DOLLING
242
237, 241
Figs 229-242 Clavigralla species. (229) alpica, 3 pronotum, dorsal view; (230) montana, holotype
3, pronotum, dorsal view; (231) alpica, left lateral profile of pronotum, scutellum and base of
corium of 3; (232) montana, holotype 3, scutellum, left lateral view; (233) alpica, left lateral
margins of ¢ abdominal sternites III-VII, ventral view; (234) montana, holotype 3, left lateral
margins of abdominal sternites III-VII, ventral view; (235) alpica, 3 genital capsule, dorsal view,
parameres removed; (236) same, left paramere, dorsal view; (237) montana, holotype, left para-
mere, dorsal view; (238) alpica, conjunctiva and vesica, left lateral view; (239) same, conjunctiva
and vesica, dorsal view; (240) same, second valvula, lateral view; (241) same, sclerites of dorsal
wall of gynatrium, dorsal view; (242) same, spermatheca.
Abdominal sterna III-VII with posterolateral angles (Fig. 233) very slightly prominent, acute. Male
genital capsule (Fig. 235) with posterior emargination completely obliterated by lip, tongue deeply bifid.
Paramere (Fig. 236) with apex hooked. Conjunctiva (Figs 238, 239) with dorsomedian lobe large, flat-
topped, its posterolateral angles produced posteriorly into small, round, accessory lobes; intermediate
dorsal, distal dorsomedian and distal dorsolateral lobes present but small; apical ventral and distal
ventrolateral lobes moderately well developed; small ventral lobes present; vesica long. Female with
second valvula of ovipositor (Fig. 240) strongly sclerotized, apex very prominent, dorsal surface strongly
corrugated. Spermatheca (Fig. 242) with duct moderately and variably convoluted. Sclerites of dorsal wall
AFRICAN POD BUGS
243
244
247
Figs 243-247 Clavigralla annectans and Myla sp. (243) C. annectans, holotype 3, head, left lateral
view; (244) same, left paramere, dorsal view; (245) same, inflated conjunctiva and vesica, dorsal
view, Outline of apices of ‘wings’ of ejaculatory reservoir complex dotted; (246) same, inflated con-
junctiva and vesica, left lateral view, ejaculatory reservoir and associated structures indicated by
dotted outlines; (247) Myla sp., left posterior trochanter and base of femur, posterior face.
of gynatrium (Fig. 241) with a large, barb-like process situated at about middle of median, longitudinal
arms.
Sculpture and distribution of pubescence generally as in C. tomentosicollis but decumbent pubescence
much less dense and all erect and suberect pubescence short, no hairs as long as transverse diameter of an
eye. Coloration of head, exposed areas of thorax and abdomen, clavus, corium and apical two-thirds of
femora pinkish brown with varying amounts of piceous coloration. Ventral surface of head, mesosternum,
metasternum, disc of abdominal venter piceous. Head laterally streaked piceous, dorsally with only
granules piceous or more or less heavily streaked piceous throughout. Antennal segment I always with a
larger or smaller ventral piceous streak, at least towards base. Rostrum of variable coloration. Pronotum
with rather sparse pubescence compared with that of other members of the tomentosicollis-group, dividing
line between anterior, decumbent, silvery pubescence and posterior, erect to decumbent, brown pubescence
almost straight (Fig. 229), posterolateral margins, patch in anterior midline of disc and at least apices of
sublateral tubercles piceous, punctures of posterior region of disc often piceous. Scutellum black, with
apex and distal half or more of midline cream, with pubescence along midline silver, elsewhere black or
brown. Thoracic pleura and abdominal sterna more or less heavily streaked piceous; dorsal midline of
abdomen always pale, cream or buff, remainder of tergites largely piceous; laterotergites and correspond-
ing areas of tergum VII dark brown or piceous, IV and V each with a narrow band and VI and VII each
with a broad transverse band buff. Corium with punctures and veins apically more or less heavily marked
with brown or piceous, punctures of clavus sometimes piceous. Hemelytral membrane hyaline, with faint
to well-marked brown streaks between longitudinal veins and a row of piceous spots between bases of
longitudinal veins where these arise from basal cells. Femora moderately to heavily marked with piceous
streaks and spots, especially on posterior surfaces, sometimes almost wholly piceous. Tibiae stramineous,
basally and sometimes apically, but never medially, with darker, brown or partly piceous annuli.
REMARKS. This species varies considerably in colour, specimens from Ethiopia being in general
the palest, with less extensive piceous markings than those from central Africa. Apparently re-
stricted to areas above the 2000 m contour. The compressed form and very short pubescence
distinguish this species from all of its congeners.
DIsTRIBUTION. Highlands of central Africa and Ethiopia. Schouteden (1957 : 308) records this
species from Rwanda and Burundi.
80 W. R. DOLLING
MATERIAL EXAMINED
Acanthomia alpica Bergroth, lectotype °, Zaire: Mt Muhavura, Birunga, 2500 m (Pr. W. Exp., Gyld) (NR,
Stockholm).
Ethiopia: | 2, Gamo Province, Mt Tola (Gughe highlands), c. 3200 m, 10-14.xii.1948 (H. Scott); 1 9,
edge of Djem-Djem Forest, c. 2700 m, 4.x.1926 (Scott); 1 9, Simien, below Atgheba Ghiyorghis, c
3200 m, from wet rocks below a spring, 7.xii.1952 (Scott); 1 9, Simien, near Enchetcab, above 2700 m,
beneath large stones round drinking pool for cattle, 25.xii.1952 (Scott); 1 2, Simien, Arghine, 3500 m or
higher, near torrent in peat soil, under boulders or at roots of plants, 24.xi.1952 (Scott) (BMNH, London).
Zaire: 1 °, Birunga, Kariss, 3000 m (Pr. W. Exp., Gyld) (paralectotype of Acanthomia alpica Bergréth)
(NR, Stockholm). Uganda: | 9, Butandiga, 12.i1.1930 (H. Hargreaves) (BMNH, London). Kenya: 1 2,
Muguga, 28.1.1969 (EZ. S. Brown) (UM, Oxford); 1 9, Naivasha, ix.1939 (H. J. A. Turner); 7 3, 1 9, West
Aberdares, 3700 m, xii.1935 (Turner); 3 2, West Aberdares, 3000-3500 m, xi.1934 (A. F. J. Gedye); 1 3,
Londiani, v.1936 (Turner); 1 ex. without abdomen, Kinangop, iii.1930 (Turner) (BMNH, London).
Clavigralla montana sp. n.
(Figs 230, 232, 234, 237)
Length: 3, 7-3 mm (n = 1). Known only from the unique male holotype. Very similar to C. alpica in
general form and in many details.
Length of antennal segment I divided by width of head including eyes 0-80. Ratio of lengths of antennal
segments I : II : III : 1V as 1-00 : 0-86 : 0-75 : 1-35. Length of basal segment of rostrum divided by width
of head including eyes 0-66, ratio of lengths of rostral segments I : II : III : [V as 1-00 : 0-93 : 0-50 : 0-81.
Pronotum (Fig. 230) with posterolateral angles less strongly produced than those of C. alpica but bear-
ing slightly longer spines; width across apices of spines divided by width of head including eyes 2-41.
Scutellum (Fig. 232) distinctly more convex than that of C. alpica. Length of posterior tibia divided by
length of posterior femur 0°81.
Abdominal sternites III-VII with posterolateral angles (Fig. 234) acutely produced, more prominent
than in C. alpica. Male paramere (Fig. 237) with apical margin distinctly notched. Coloration as in darker
examples of C. alpica, except that tibiae, tarsi, ground coloration of femora and antennal segments I-III
red. Pubescence distinctly longer than that of C. alpica, many of the outstanding hairs on head, thorax
and appendages as long as transverse diameter of an eye (e.g. pubescence of scutellum, Fig. 232).
REMARKS. Obviously very closely related to C. alpica, from which it may be distinguished by the
longer pubescence and other characters presented above. This species probably occupies the same
montane niche in West Africa as that favoured by C. alpica in Ethiopia and central Africa.
DISTRIBUTION. Cameroun mountains.
MATERIAL EXAMINED
Holotype 3, Cameroun: Bamenda, 17.xiii.1955 (Bechyne) (BMNH, London).
Nomenclature of economically important species
As a result of this revision it has proved necessary to make changes in the names of the African
species formerly referred to the genus Acanthomia Stal (variously mis-spelt in collections and
published work as Acanthomya and Acanthomyia). This generic name falls as a synonym of Clavi-
gralla Spinola, thus placing the African ‘pod bugs’ or ‘spiny brown bugs’ in the same genus as the
Indian minor pest known as the ‘tur pod bug’, Clavigralla gibbosa, which has been the subject of
several biological and anatomical studies (for details of which see Dolling, 1978 : 288-289).
The well-known and widespread Acanthomia tomentosicollis (Stal) reverts to Clavigralla
tomentosicollis Stal and the less important ‘A. brevirostris’ is shown to be restricted to a small area
in the north-eastern part of the Ethiopian region, southern Arabia and parts of the Oriental
region, its correct name being Clavigralla scutellaris (Westwood).
The minor pest species known as A. hystricodes (Stal) reverts to Clavigralla hystricodes Stal.
The name Acanthomia horrida (Germar), correctly applied, should refer (as Clavigralla horrida
(Germar)) to an economically unimportant species with a restricted distribution in South Africa
and Rhodesia. In West Africa the name horrida has been misapplied to the pest species which
should now be called Clavigralla shadabi Dolling. The same name has been misapplied in East and
AFRICAN POD BUGS 8]
Southern Africa to a different species for which the name Clavigralla elongata Signoret is avail-
able. Both shadabi and elongata are widespread in Zaire.
Check-list of the species of the tribe Clavigrallini
CLAVIGRALLOIDES Dolling
acantharis (Fabricius)
tuberculata Dallas
spinigera Walker
spinosus Dolling
quadrituberculatus (Breddin)
enkidu Dolling
tuberosus tuberosus (Hsiao)
tuberosus indicus Dolling
GRALLICLAVA Dolling
indica Dolling
soror Dolling
rubra Dolling
solitaria Dolling
insularia Dolling
montana montana Dolling
montana sinensis Dolling
dissimilis Dolling
horrens horrens (Dohrn)
spinofemoralis Shiraki
horrens palawanensis Dolling
indecora (Walker)
irianensis Dolling
australiensis Dolling
ONCASPIDA Stal
pilosicollis (Stal)
similis Signoret
CLAVIGRALLA Spinola
Acanthomia Stal syn. n.
Lancha Shadab syn. n.
tuberculicollis (Reuter) comb. n.
leroyi (Schouteden) comb. n.
mixta (Schouteden) syn. n.
zambiae sp. n.
uelensis (Schouteden) comb. n.
hystrix Dallas
hystricodes Stal
elongata Signoret
flavipennis Signoret
shadabi sp. n.
breviceps sp. n.
ankatsoensis sp. n.
madagascariensis sp. n.
asterix sp. n.
schnelli (Villiers) comb. n.
mira sp. n.
annectans sp. n.
insignis (Distant) comb. n.
minor (Schouteden) comb. n.
andersoni sp. n.
angolensis sp. n.
egregia sp. n.
longispina sp. n.
aculeata sp. n.
horrida (Germar)
muricata Stal syn. n.
natalensis Stal
leontjevi (Bergroth) comb. n.
griseola (Linnavuori) comb. n.
ruandana (Schouteden) comb. n.
biston sp. n.
oxonis sp. n.
bovilla sp. n.
pusilla sp. n.
strabo sp. n.
pabo sp. n.
spiniscutis (Bergroth) comb. n.
marmorata sp. nN.
tomentosicollis Stal
tomenticollis Stal
scutellaris (Westwood)
brevirostris Stal
annulipes Signoret
gibbosa Spinola
orientalis orientalis Dolling
orientalis serica Dolling
curvipes (Stal) comb. n.
simillima sp. n.
wittei (Schouteden) comb. n.
neavei Sp. Nn.
alpica (Bergroth) comb. n.
montana sp. n.
Acknowledgements
My sincere thanks are due to the following people for the loan of material in their care: Dr
P. H. Arnaud, CAS, San Francisco; Mr J. C. Deeming, IAR, Samaru; Dr P. Duarte Rodrigues,
Faculdade de Ciéncias, Lisbon; Dr U. Gdllner-Scheiding, MNHU, Berlin; Dr L. Hedstrém,
University of Uppsala; Dr A. Kaltenbach, NM, Vienna; Dr R. Kumar, UG, Legon; Mr I.
Lansbury and Dr M. W. R. de V. Graham, UM, Oxford; Dr M. Meinander, ZMU, Helsinki;
Dr T. Nyholm, NR, Stockholm; Dr E. C. G. Pinhey, NM, Bulawayo; Dr G. L. Prinsloo, PPRI,
82 W. R. DOLLING
Pretoria; Mr J. A. van Reenen, TM, Pretoria; Dr G. Schmitz, MRAC, Tervuren; Mr M. U.
Shadab and Dr P. Wygodzinsky, AMNH, New York; Prof. J. A. Slater, University of Con-
necticut, Storrs; Dr H. Synave, IRSNB, Brussels; Dr A. Villiers, MNHN, Paris.
References
Aina, J. O. 1972. The biology of Acanthomia horrida Germ. (Heteroptera, Coreidae), a pest of fresh
cowpea pods. Bull. ent. Soc. Nigeria 3 : 85-90, 4 figs.
Anonymous. 1926. Miscellaneous entomological notes. Fmg S. Afr. 1 : 23.
1927. Overcoming the bean bug. Fmg S. Afr. 2 : 87.
1947. Museen und Sammlungen. Jn Warnecke, G. (ed.), Interne Mitglieder-Nachrichten. Faunistiche
Arbeitsgemeinschaft fiir Schleswig-Holstein, Hamburg und Liibeck. Rundschreiben 1947 (1) : 3-5.
Bergroth, E. E. 1903. Rhynchota aethiopica. III. Annls Soc. ent. Belg. 47 : 288-297.
1908. Neue Hemiptera aus Siid-Abyssinien. Russk. ent. Obozr. 7 : 106-110.
—— 1913. On a small collection of Hemiptera from north eastern French Congo. Annls Soc. ent. Belg.
57 : 237-239.
—— 1927. Zoological results of the Swedish expedition to central Africa 1921. Insecta. 21. Hemiptera
Heteroptera. Ark. Zool. 18 (A) 26 : 1-12.
Bohlen, E. 1973. Crop pests in Tanzania and their control. 142 pp., 270 figs. Berlin & Hamburg.
Booker, R. H. 1965. Pests of cowpea and their control in N. Nigeria. Bull. ent. Res. 55 : 663-671.
Burnett, G. F., Lee, C. W. & Park, P. O. 1966. Aircraft application of insecticides in East Africa. XV. Very
low-volume treatment of a seed-bean crop with DDT in oil solution. Bull. ent. Res. 56 : 701-714, 5
tables.
Chiaromonte, A. 1934. Considerazioni entomologiche sulla coltura delle piante ortensi nella Somalia
Italiana. Agricoltura colon. 27 : 523-529.
Cobben, R. H. 1968. Evolutionary trends in Heteroptera. Part I. Eggs, architecture of the shell, gross
embryology and eclosion. Meded. LandbHoogesch. Wageningen 151 : 475 pp., 316 figs.
Dallas, W. S. 1852. List of the specimens of hemipterous insects in the collection of the British Museum. Part
II. Pp. 369-590, pls 12-15. London.
Distant, W. L. 1908. Descriptions of some Rhynchota from Ruwenzori. Ann. Mag. nat. Hist. (8) 2:
436-444.
—— 1909. Ruwenzori expedition reports. 9. Rhynchota. Trans. zool. Soc. Lond. 19 : 67-84, pl. 2.
Dolling, W. R. 1978. A revision of the Oriental pod bugs of the tribe Clavigrallini (Hemiptera : Coreidae).
Bull. Br. Mus. nat. Hist. (Ent.) 36 : 281-321, 108 figs.
Egwuatu, R. I. & Taylor, T. A. 1977a. Development of Gryon gnidus (Nixon) (Hymenoptera : Scelionidae)
in eggs of Acanthomia tomentosicollis (Stal) (Hemiptera : Coreidae) killed either by gamma irradiation
or freezing. Bull. ent. Res. 67 : 31-33.
1977b. Bionomics of Gryon gnidus Nixon (Hymenoptera : Scelionidae), an egg parasite of
Acanthomia tomentosicollis Stal (Hemiptera : Coreidae) in Nigeria. Appl. Ent. Zool., Tokyo 12 (1):
76-78.
—— —— 1977c. Studies on the biology of Acanthomia tomentosicollis (Stal) (Hemiptera : Coreidae) in
the field and insectary. Bull. ent. Res. 67 : 249-257.
Fuller, C. 1922. Report of the Division of Entomology. J. Dep. Agric. Un. S. Afr. 5 : 542-545.
Germar, E. F. 1840. Hemiptera Heteroptera Promontorii Bonae Spei, nundum descripta, quae collegit
C. F. Drége. Revue Ent. (Silbermann) 5 : 121-192.
Grant, J. A. & Stys, P. 1970. Hyocephalus aprugnus Bergroth, 1906 (Insecta, Hemiptera—Heteroptera,
Hyocephalidae): request for the retention of the neotype in preference to the rediscovered holotype.
Z.NAS.) 1916. Bull. zool. Nom. 27 : 113-114.
Herrich-Schaeffer, G. A. W. 1851. Die Wanzenartigen Insekten. 9 (6) : 257-348, pls 319-324. Niirnberg.
Johnston, H. B. 1930-1931. Summary report on the entomological work of the Gezira Laboratory for the
year 1929. Bull. Wellcome trop. Res. Labs. (Ent. Sect.) 31 : 42-47.
Lindberg, H. 1958. Hemiptera Insularum Caboverdensium. Systematik, Okologie und voc der
Heteropteren. Commentat. biol. 19 (1). 246 pp., 114 figs.
Linnavuori, R. 1971. On the family Coreidae (Het.). Suom. hydént. Aikak. 37 : 163-178, 24 figs.
—— 1978. Hemiptera of the Sudan, with remarks on some species of the adjacent countries 6. Aradidae,
Meziridae, Aneuridae, Pyrrhocoridae, Stenocephalidae, Coreidae, Alydidae, Rhopalidae, Lygaeidae.
Acta zool. fenn. 153. 108 pp., 74 figs.
Materu, M. E. A. 1970. Damage caused by Acanthomia tomentosicollis Stal and A. horrida Germ. (Hemip-
tera, Coreidae). E. Afr. agric. For. J. 35 : 429-435, 5 tables.
AFRICAN POD BUGS 83
—— 1971. Population dynamics of Acanthomia spp. (Hemiptera, Coreidae) on beans and pigeon peas in
the Arusha area of Tanzania. E. Afr. agric. For. J. 36 : 361-383, 4 figs, 13 tables.
—— 1972. Morphology of adults and description of the young stages of Acanthomia tomentosicollis Stal
and A. horrida Germ. (Hemiptera, Coreidae). J. nat. Hist. 6 : 427-450, 15 figs, 4 tables.
—— & Makusi, R. A. 1972. Toxicity of technical endosulfan applied topically to Acanthomia spp (Coreidae,
Hemiptera). E. Afr. agric. For. J. 37 : 313-317.
1973. Toxicity of synergised pyrethrum to Acanthomia spp. 1. Acanthomia tomentosicollis Stal.
E. Afr. agric. For. J. 39 : 141-153.
Nixon, G. E. J. 1943. A synopsis of the Ethiopian and Indo-Malayan species of Microphanurus (Ser-
phoidea, Scelionidae). Bull. ent. Res. 34 : 135-144, 9 figs.
Reuter, O. M. 1887. Ad cognitionem Heteropterorum madagascariensium. Ent. Tidskr. 8 : 77-109.
Risbec, J. 1950. Sur la ponte et le développement larvaire des insectes dans les cultures au Sénégal. Agron.
trop., Nogent 5 : 227-267, 10 figs.
1951. Les Chalcidoides d’A.O.F. Mém. Inst. fr. Afr. noire 13. 409 pp., 179 figs.
—— 1955. Hyménoptéres parasites du Cameroun. Bull. Inst. fr. Afr. noire Sér. A. Sci. nat. 17 : 191-266,
11 figs.
Schouteden, H. 1938. Catalogues raisonnés de la faune entomologique du Congo Belge. Hémiptéres,
Coreidae. Annis Mus. r. Congo Belge C. Zool. Sér. 3 (2) 1 : 221-308.
—— 1957. Contributions a l’étude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky
1953). CXXVI. Heteroptera Plataspidae, Acanthosomidae, Pentatomidae et Coreidae. Annls Mus. r.
Congo Belge (Sér. 8vo Sci. zool.) 58 : 269-310.
Shadab, M. U. 1972. A new genus of pseudophloeine bugs from the Democratic Republic of the Congo.
Am. Mus. Novit. no. 2493. 11 pp., 23 figs.
Signoret, V. 1860. Faune des Hémiptéres de Madagascar. 2. Hétéroptéres. Annis Soc. ent. Fr. (3) 8 :
917-972, pls 13, 14.
Smit, B. 1964. Insects in southern Africa: how to control them. xiv + 399 pp., 8 pls. Cape Town.
Spinola, M. 1837. Essai sur les genres d’insectes appartenants a l’ordre des Hémiptéres, Lin. ou Rhyngotes,
Fab. et ala section des Hétéroptéres, Dufour. 383 pp., 15 tables. Genoa.
Stal, C. 1855. Hemiptera fran Cafferlandet. Ofvers. K. Vetensk Akad. Férh. 12 : 27-46.
1866. Hemiptera Africana. 2. 181 pp. Stockholm.
—— 1873. Enumeratio Hemipterorum 3. K. svenska Vetensk Akad. Handl. 11 (2). 163 pp.
Swaine, G. 1969. Studies on the biology and control of pests of seed beans (Phaseolus vulgaris) in northern
Tanzania. Bull. ent. Res. 59 : 323-338.
Taylor, T. A. 1975. Gryon gnidus, a Scelionid egg-parasite of Acanthomia tomentosicollis (Hem : Coreidae)
in Nigeria. Entomophaga 20 : 129-134.
— & Omoniyi, O. 1972. Variation in number of micropyles in eggs of Acanthomia tomentosicollis (Stal)
(Hemiptera : Coreidae). Niger. ent. Mag. 2 : 70-73, 4 figs.
Thunberg, C. P. 1822. Dissertatio entomologica de Hemipteris Rostratis capensibus. Ill. 8 pp. Upsala.
Villiers, A. 1950. Note d’entomologie Ouest-africaine. 1. Descriptions de nouveaux Hémiptéres Hétér-
optéres. Bull. Inst. fr. Afr. noire 12 : 634-659, 17 figs.
Westwood, J. O. 1842. A catalogue of Hemiptera, in the collection of the Rev. F. W. Hope, M.A. F.R.S.
F.L.S. F.Z.S. M.E.S. etc. etc. etc. etc. with short Latin descriptions of the new species. Part II. 26 pp.
London.
Index
Synonyms are in italics; references to descriptions are in bold. Entries in checklist (p. 81) are not indexed.
acantharis 44, 46 biston 11, 52
Acanthomia 6, 80 bovilla 7, 11, 54
aculeata 7, 10, 43 breviceps 10, 30
alpica 11, 77, 80 brevirostris 69, 80
andersoni 10, 41
angolensis 10, 41, 43, 44 Glavi .
: Z gralloides 3
ankatsoensis 9, 31, 33 :
annectans 7, 9, 38 curvipes 8, 10, 60-62, 70, 73
annulipes 11, 70 egregia 10, 41
asterix 9, 31, 33 elongata 7, 8, 10, 19, 23, 24, 28-46 passim, 81
Clavigralla 2, 3, 6, 80
84
flavipennis 24
gibbosa 6, 8, 63, 80
gracilis 34, 36
Gralliclava 3
griseola 50
horrens 6
horrida 24, 25, 27, 29, 80
horrida 9, 44, 46, 80
horridus 46
hystricodes 7, 8, 9, 21, 80
hystrix 7, 8, 9, 19, 21
insignis 7, 10, 39, 40-46 passim
Lancha 6
leontjevi 7, 8, 10, 48, 50
leroyi 7, 8, 14, 17-19
longispina 10, 43
madagascariensis 9, 31, 33, 34
marmorata 11, 63
minor 10, 39
mira 7, 9, 36, 38
mixta 14, 17, 19
montana 11, 80
muricata 44, 46
Myla 6, 8
W. R. DOLLING
natalensis 6, 7, 9, 46
neavei 12, 77
Oncaspidia 2, 3, 8
orientalis 8, 63
oxonis 11, 53
pabo 12, 57, 59
pilosicollis 3
pusilla 11, 56, 70
Risbecocoris 2
ruandana 7, 8, 11, 50, 52-56 passim
schnelli 6, 7, 9, 34
scutellaris 8, 11, 63, 67, 68, 69, 70, 80
shadabi 8, 10, 28, 33, 36, 42, 44, 45, 80, 81
similis 3
simillima 8, 10, 70, 73
spiniscutis 7, 8, 12, 54, 57, 59, 60, 63
strabo 7, 12, 57, 59, 60
tomenticollis 63
tomentosicollis 6, 7, 8, 11, 48-79 passim, 63, 80
tuberculicollis 7, 8, 12
uelensis 7, 8, 18
wittei 7, 8, 11, 74, 77
zambiae 8, 17
eee
British Museum (Natural History) Monographs
A Study of the Aphididae (Homoptera) of West Africa.
V. F. Eastop 1961
The Morphology and Taxonomy of Male Scale Insects (Homoptera: Coccoidea)
M. S. K. Ghauri 1962
Whitefly of the World: A systematic catalogue of the Aleyrodidae
(Homoptera) with host plant and natural enemy data.
L. A. Mound & S. H. Halsey. Co-published with J. Wiley & Son, 1978
Classification of the Aradidae (Hemiptera—Heteroptera)
R. L. Usinger & R. Matsuda 1959
Lists are available free on request to:
Publications Sales
British Museum (Natural History)
Cromwell Road
London SW7 SBD
w
Titles to be published in Volume 39
A revision of the African pod bugs of the tribe Clavigrallini
(Hemiptera: Coreidae) with a checklist of the world species.
By W. R. Dolling.
A revisionary classification of the Cynipoidea of the Ethiopian
Zoogeographical Region. By J. Quinlan.
A review of the genera of the Hydroptilidae (Trichoptera).
By J. E. Marshall.
A revision of Vitessidia Rothschild & Jordan and Vitessa Moore
(Lepidoptera: Pyralidae). By E. G. Munroe & M. Shaffer.
Printed by Henry Ling Ltd, Dorchester
—
Bulletin of the
British Museum (Natural History)
A revisionary classification of the
Cynipoidea (Hymenoptera) of the
Ethiopian Zoogeographical Region
Aspicerinae (Figitidae) and Oberthuerellinae
(Liopteridae)
J. Quinlan
Entomology series Vol 39 No2 25 October 1979
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and
an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum’s resources. Many of the papers
are works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and are not
necessarily completed within one calendar year. Subscriptions may be placed for one or more
series. Subscriptions vary according to the contents of the Volume. and are based on a forecast
list of titles. As each Volume nears completion, subscribers are informed of the cost of the
next Volume and invited to renew their subscriptions. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW7 5BD,
England
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1979
ISBN 0524-6431 Entomology series
Vol 39 No 2 pp 85-133
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 October 1979
A revisionary classification of the Cynipoidea
(Hymenoptera) of the Ethiopian Zoogeographical
Region
Aspicerinae (Figitidae) and Oberthuerellinae (Liopteridae)
J. Quinlan
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 SBD
Contents
Synopsis. : : 7 : ; E : ‘ : ‘ : ; 85
Introduction , 2 ; : : ; 85
Material examined, terminology and acknowledgements P ; : ; : 86
Family classification of Cynipoidea , : ; : : : ‘ j 86
Taxonomic characters of Cynipoidea : ; : : : ; 88
Key to families and subfamilies of world Cynipoidea ; : : : : 90
Checklist of the families and subfamilies of Cynipoidea_. ; F 91
Checklist of the Ethiopian species of Aspicerinae and Oberthuerellinae : ¢ 92
Aspicerinae Kieffer : . : A : : ; , 93
Key to genera of Ethiopian Aspicerinae , : ‘ ; : ; : 93
Anacharoides Cameron. : : : A , : 94
Key to species of Uneeharoides Cameron : ; ; : : ; 94
Females . ‘ ; ; ; , : : y j : : 94
Males. : F ' : ; : 3 p : ’ : 95
Proaspicera Kieffer ; ‘ , f : : 104
Key to Ethiopian species of Proaspicera. Kieffer : ; : : ; 105
Females . : ? 3 : ; ' ‘ ‘ ; : , 105
Males. : : , ‘ ; 3 : : : ; ; 106
Oberthuerellinae Kieffer ; ‘ ; : : : : : . 109
Key to genera of Oberthuerellinae : ‘ , P : . : ‘ 110
Oberthuerella Saussure. ; ; : . F ; 110
Key to species of Oberthuerella Saussure : : é : : : 110
Tessmannella Hedicke . ; : : : : : 116
Key to species of Tessmannella Hedicke : : : , , : 116
Xenocynips Kieffer : ‘ : 5 ‘ : : ; : : 118
References . : : ; ; A ; : A ; P : ; 119
Index : : , ; ; : ; ; : : ; 5 : 133
Synopsis
This is the first of a series of papers which will cover the taxonomy of the cynipoid fauna of the Ethiopian
Region. Keys to the world families and subfamilies are provided. In addition, keys to the genera and
species of the Ethiopian Aspicerinae and Oberthuerellinae are given and are based on a study of type-
specimens and all other available material. Twelve new species are described, and five new combinations
and two new specific synonymies are established.
Introduction
This is the first attempt to review the Ethiopian Cynipoidea as a whole. Except where specific
exceptions are mentioned the term Ethiopian Region refers to the original concepts of Sclater
Bull. Br. Mus. nat. Hist. (Ent.) 39 (2) : 85-133 Issued 25 October 1979
85
86 J. QUINLAN
(1858) and Wallace (1876). The most recent work covering the Cynipoidea at the family, sub-
family and generic level is that of Weld (1952); this work dealt with the fauna of all geographical
regions and superseded that of Dalla Torre & Kieffer (1910), which was the first comprehensive
treatise on the Cynipoidea. More recently Benoit (1952-56) described various genera and species
from Africa but did not include generic keys. All known species of which type-material is available
have been fully redescribed. Where the type-material is lost its presumed taxonomic position is
indicated in the text. In very few instances are host data given, and these are based solely on
labels attached to the specimens.
Material examined, terminology and acknowledgements
I have been fortunate in being able to borrow large amounts of material from Dr J. Decelle,
Musée Royal de |’Afrique Centrale, Tervuren; Madame S. Kelner-Pillault, Muséum National
d’Histoire Naturelle, Paris; Dr E. K6nigsmann, Museum fiir Naturkunde der Humboldt-
Universitat, Berlin. All have been kind enough also to loan type-material from their respective
collections. This material, together with R. E. Turner’s cynipoid collections in the British Museum
(Natural History) and more recent material obtained from Malaise traps, has formed the basis
for this revision. There are, however, large areas in the Ethiopian Region from which no material
has been available for examination. The terminology used is as in Richards (1977), but some older
names are used particularly in reference to wing venation. Some of the more general taxonomic
terms used in this paper are illustrated by labelled figures (Figs 1-21). It is hoped that this will
provide a correlation between the terms used by Richards (1977) and by Ross (1936). Type
depositories are given in abbreviated form in the text and are listed below in alphabetical order
of the towns in which they are situated.
MNHU, Berlin Museum fiir Naturkunde der Humboldt-Universitat, Berlin
IP, Eberswalde Institut fiir Pflanzenschutzforschung, Eberswalde, Germany
AM, Grahamstown Albany Museum, Grahamstown, South Africa
KSM, Kansas Snow Entomological Museum, University of Kansas, Lawrence, Kansas
BMNH British Museum (Natural History), London
MNHN, Paris Muséum National d’Histoire Naturelle, Paris
MRAC, Tervuren Musée Royale de |’Afrique Centrale, Tervuren, Belgium
I am indebted to Mr A. Sutton for mounting and labelling the drawings and Miss V. I. Dick for
typing the manuscript. My sincere thanks are extended to Mr T. Huddleston for his constructive
and helpful advice during the preparation of this paper
Family classification of Cynipoidea
In preparing the key to families account has been taken of subfamilies proposed since Weld (1952).
Five families and thirteen subfamilies are recognised in this paper. Yoshimoto (1970) proposed the
name Himalocynipinae to include the genus Himalocynips. According to Yoshimoto this new
subfamily is distinguished from Figitinae on the number of antennal segments, a partial nervulus
and post nervulus in the forewing, and a projecting clypeus. I have not seen the unique specimen
on which the subfamily is based but have included it in the key to subfamilies on the basis of the
description. Similarly, to accommodate the genus Austrocynips Riek (1971) erected the sub-
family Austrocynipinae which, because it has a strongly produced hypopygium, he associated
with the subfamily Cynipinae. The wing venation, however, appears to be closer to that of the
Pycnostigmatinae, which have a radial cell resembling at first sight a pterostigma.
In the family and subfamily keys the Eucoilidae have been treated as a family and not as a
subfamily of the Figitidae as in Weld (1952) and earlier authors. The reason for this is stated by
Eady & Quinlan (1963), namely that the differences between Eucoilidae and subfamilies Anach-
aritinae, Figitinae and Aspicerinae are much greater than the differences between these three
subfamilies of Figitidae.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 87
The Ibaliidae are distinguished from all other families by the elongate radial cell (Fig. 9),
distinct areolet, smooth and inconspicuous petiole and elongate metatarsus (Fig. 10). As far as
is known all species are parasites of Siricidae (Symphyta). They are probably the largest cynipoids,
being between 10 and 25 mm in length. Their distribution is Holarctic and Australasian.
The Liopteridae are distinguished from the Ibaliidae by the short radial cell and from other
families by the relative lengths of the gastral segments. The Liopteridae are large, 5-14 mm in
length. The family consists of three subfamilies, mainly tropical in distribution, of which two
(Oberthuerellinae and Mesocynipinae) are known to occur in the Ethiopian Region. The life
history is not known.
The Eucoilidae are parasites of Diptera, and are clearly distinguished from other Cynipoidea
by their distinctive scutellum, which comprises an outer disc surmounted by a cup-shaped
elevation in the centre (Fig. 14). The pronotum is very often distinctly produced medially into an
anterior plate with a strong posterior margin (pronotal plate). In the forewing the cubitalis
(Rs+M) is usually visible and emerges from a point very close to the lower end of the basalis
(close to the junction of M+Cu) or combined with the median (Cu,) for a short distance (Fig.
15). The life histories of some of the more common species are known; they are of economic
importance in that they parasitise Diptera attacking cereal crops. Eucoilidae have a wide range
of dipterous hosts including the stem-boring and leaf-mining larvae of the family Agromyzidae,
the plant-feeding families Chloropidae and Anthomyiidae and those larvae of the family Sphaero-
ceridae and other flies associated with animal excreta. Adult Eucoilidae vary in size from 1-5 to
5-0 mm and are world-wide in distribution.
The Figitidae include four subfamilies, each resembling the others in the majority of characters.
The Aspicerinae are generally the largest, 4-5-5-0 mm in length, and are parasites of the larger
Diptera such as Syrphidae. The gaster has a very distinctive second tergite which is shaped rather
like a saddle and termed liguliform (Fig. 5). The Anacharitinae have wedge-shaped heads and
generally the second tergite of the gaster is longer than the third (Fig. 8). The forewing has at the
most a vestigial cubitus (vein M) and a distinctive radial cell (Fig. 21). The adults vary in size
from 3 to 4 mm depending on the length of the petiole. Some genera have a short petiole and can
be confused wth the subfamily Figitinae. The Anacharitinae are parasites of lace-wings (Neur-
optera). The Figitinae are very similar to the Anacharitinae and Aspicerinae, but in the majority
of genera the second tergite is shorter than the third or equal to it and generally the cubitus (vein
M) is distinct, thus separating it from the Anacharitinae. They are parasites of smaller Diptera and
range in size from 2 to 4 mm. They are world-wide in distribution. However, all three subfamilies
have genera that can be confused with those of other subfamilies, e.g. in Figitinae the genera
Neralsia Cameron and Xylophora Kieffer have a very reduced second gastral tergite which can be
mistaken for that of an Aspicerinae, and in some cases the cubitus is vestigial as in the Ana-
charitinae.
The subfamily Himalocynipinae is similar to the Figitinae (Yoshimoto, 1970) but differs in
having 20-segmented antennae, the head unusually small in relation to the thorax, the second
tergite of the gaster considerably longer than the third, a partial nervulus and postnervulus present
in the forewing, and a small subcordate clypeus strongly projecting forward and upward. Size
5:5 mm.
The family Cynipidae is represented by three subfamilies, each having distinctive characters,
but some genera are nevertheless difficult to place in a subfamily. Cynipidae are separated from
related families on the forewing venation and the relative lengths of the gastral segments. In the
forewing the cubitalis (Rs+M), when visible, arises from a point midway along the basalis
(Rs+M) (Fig. 17) and in the female the largest free tergite is either the second or the second and
third fused (Figs 18-19). The Pycnostigmatinae contains two genera; as far as is known both are
restricted to Africa. The forewing is very distinctive in having a reduced radial cell which, unless
closely examined, can be mistaken for a pterostigma, the wing veins are very thick (Fig. 23), and
the number of antennal segments is 12, 15 or 19. When viewed laterally, the gaster has what
appears to be one large segment occupying almost the whole of the visible surface area; this
consists of segments two and three fused together, the remaining segments generally being
obscured. The first segment of the gaster in one genus is longitudinally ridged as in the cynipine
88 J. QUINLAN
genus Synergus Hartig. This appearance of the gaster has been taken by some authors as suggest-
ing the habitus of an inquiline. At present its life history is unknown. Size 2-5—4-0 mm.
The Cynipinae are usually sculptured and the middle and hind tibiae each have two distal spurs.
They are either gall-causers or inquilines in galls of other Cynipinae. The gall-causers attack a
range of plants: Compositae, Labiatae, Papaveraceae, etc. (herbacous plants); Rosaceae;
Fagaceae (Quercus). The only gall-causer at present known from the Ethiopian Region is
Rhoophilus Mayr on Rhus (Anacardiaceae). In the extreme northern parts of Africa where there
are oaks various European genera (Andricus, Neuroterus, etc.) are found. Some species of
Cynipinae on oakexhibit the phenomenon of alternation of generations. One generation consisting
of both males and females and the other entirely of agamic females reproducing partheno-
genetically. One genus, Ceroptres, has been bred commonly from galls on oak; Blair (1949),
however, stated that C. arator Hartig was parasitic on Andricus kollari (Hartig). Riek (1971) refers
to Ceroptres as being an inquiline in dipterous galls. The size varies from 2 to 7 mm.
The Alloxystinae are very small in relation to other Cynipoidea, the larger specimens being
only about 2mm long, and are separated from other Cynipidae by the characteristic shiny
appearance although sculpture is revealed by the scanning electron microscope, see Andrews
(1978). The whole insect lacks the rugose type of sculpture found on the mesonotum of Cynipinae,
and the scutellum only rarely has distinct foveae; occasionally notaulices are present and in others
a mesopleural suture is present. They are hyper-parasites of aphids through the braconid sub-
family Aphidiinae. Distribution is world-wide.
Taxonomic characters of Cynipoidea
Antennae
The antennae of the 9 are generally 13-segmented and vary in length and shape but are constant
within a species. Some species have filiform antenna and others are club-shaped. The 3 antennae
are usually 14-segmented and filiform. However, in certain genera the number of segments can
vary by the occasional extra segments and in one or two subfamilies by up to a maximum of
7 extra segments inthe 9. The antenna consists of a scape, pedicel and flagellum (Fig. 28). In
describing the females of some species the term club is used and in this paper, in order that a club
segment is not confused with other segments, I have counted as club segments those with rhinaria.
The relative lengths of antennal segments are given as ratios, e.g. Segment Numbers (SN) 1, 2,
3, 4=Antennal Ratio (AR) 5 :3 : 18 : 12, to further explain the overall shape of the antennae.
In some males some of the basal segments of the flagellum are modified, i.e. swollen or excised
(Figs 38-40).
Head
Characters most useful are the distance between the compound eyes (inner orbits) compared with
the height of an eye measured medially. The variation in the shape of the vertex viewed posteriorly
and in some subfamilies the shape of the occipital carina and sculpture in the occipital area are
important. Some species have heads broader than the thorax. In general, however, head characters
have not been used at the family and subfamily level but within genera and species a wide variety of
different parts have been used. In some groups the form of mandible varies and probably relates
to the biology of particular groups and their various methods of emerging when adult, be their
habitat as larvae a gall or an insect. The sculpture of the face, the presence or absence of striae
and of pubescence are usually important at the specific level.
Thorax
The thorax has many important characters (Fig. 29). As in Apocrita generally the first abdominal
segment has become fused with the thorax and is connected to the abdomen by a narrow petiole.
This post-propodeal part of the abdomen is termed the gaster (Figs 1-8). In some families the
pronotum has a distinctive medial protrusion (pronotal plate) which varies in shape and this is a
useful character at the generic and species level (Figs 13, 22, 30). This is best seen in anterodorsal
view. In certain species the pronotal plate is either absent or represented by small pits or foveae.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 89
The mesoscutum can have notaulices (parapsidal grooves) which vary in length, width and
surface sculpture. Anterior parallel lines, lateral lines and a median scutal line are present in some
species, absent in others. The mesopleuron sometimes has a mesopleural suture (precoxal suture
or carina of Richards, 1977; median episternal groove of Snodgrass, 1910). This suture, when
present, can be very distinct or lost in a mass of striations or reticulate-rugose sculpture. Its form
within a species is fairlyconstant and therefore a useful character either at the species level or at
the generic level in some families. An area present in some genera in the upper anterior part of
the mesopleurae and more recently termed mesopleural triangle (Andrews, 1978) is useful at the
generic level. In the majority of genera the scutellum has either one or two pits or foveae at the
base bordering the suture separating the mesoscutum from thescutellum, althoughinsome genera,
e.g. Neuroterus, the mesoscutum is fused to the scutellum without a suture. In Eucoilidae some
genera have a distinctive lateral bar on either side of the foveae. The scutellum is usually separated
from the mesoscutum by a suture and varies enormously between families, being smooth and
shiny in the Alloxystinae, coarsely sculptured in the Figitinae and Aspicerinae or with a distinc-
tive disc or cup in the Eucoilidae. The term cup used by Weld (1952) is employed here to differ-
entiate that centre part of the scutellum from the outer part termed the disc. The disc itself has
dorsal and lateral faces in some families (Fig. 29). In some genera of Aspicerinae, Figitinae and
Eucoilidae the apex of the scutellum is produced to form a very distinct spine. The propodeum
generally has parallel or bowed carinae which are sometimes obscured by pubescence.
Gaster
The gaster is extremely variable in the relative lengths of the segments and in their shape viewed
laterally and dorsally. In a number of instances there has been fusing together of segments. The
first segment of the gaster is petiolate, either long or short, smooth or sulcate and sometimes
hidden by a ring of pubescence at the base of tergite 2 in the Eucoilidae. The surface of tergite 2
at the base is sometimes striated and other segments can be punctate. The hypopygium is very
clearly produced in the Cynipinae, the ventral spine varying in length.
Legs
The legs vary in size and the shape of the various parts, and can be densely pubescent on the
coxae in some species. In some genera the hind tibiae are longitudinally rigid or furrowed, in
others the hind femur has a tooth. The mid and hind tibial spurs can vary in length. In the
Cynipidae the hind tarsal claws provide useful generic and specific characters. In the Ibaliidae
the metatarsus is exceptionally long, being twice as long as tarsal segments 2—5 combined.
Wings
Cynipoidea generally are fully winged although some families have brachypterous or apterous
forms. The venation of the forewing, though somewhat reduced, generally retains two branches
of vein Rs (Fig. 15). The presence of vein Rs+M and the point of juncture with Rs+M and its
closeness to Cu, are important at the subfamily level in those groups where it is present. The
radial cell varies significantly in shape within families and from species to species, being either open
or closed on the wing margin. The presence or absence of the areolet and cubitus is important in
all groups. Generally the wing surface has fine scattered pubescence, although in some species this
character is completely absent or reduced to a dotted surface and is invariably ciliate on the wing
margins. In some species the apex of the forewing can be arcuate, incised or truncate though
generally it is rounded.
Sculpture
Cynipoidea exhibit extremes of sculpture, both in form and intensity, on various parts of the
insect. Only the Alloxystinae appear completely devoid of sculpture (visible under scanning
electron microscope). In other subfamilies, e.g. Liopteridae, the mesonotum has a coarse trans-
versely sculptured surface. Inthe Figitidae the sculpture of the mesonotum varies in form from dull
and sculptured to smooth and shining. The Cynipinae gall-causing genera with alternate genera-
tions exhibit extreme diversity. In the agamic forms the face, mesoscutum, scutellum and
90 J. QUINLAN
mesopleurae have different types of sculpture and some are pubescent. In the sexual forms the
various parts can be alutaceous or finely coriaceous. The. terminology for sculpture follows
Eady (1968).
Colour
Most Cynipoidea are sombre in colour. They are more generally a mixture of black, red or
yellow but never metallic as in some other Hymenoptera. In the larger species the thorax tends
to be blackish with a distinctive reddish brown gaster, e.g., Liopteridae, Ibaliidae. In the smaller
species a mixture of red, black and yellow is normal, and exceptionally in the agamic forms of some
Cynipidae, a dull darkish yellow. Some species are predominantly blackish, in other species
reddish blackish yellowish colour patterns are exhibited. The use of colour in identification has
to be treated cautiously because, as with other Hymenoptera, factors such as humidity, tempera-
ture, host or plant habitat of the larval stages can have a strong influence on the colour of the
adult insect.
Key to families and subfamilies of world Cynipoidea
1 Largest segment of the gaster the fourth, fifth or sixth viewed laterally (Figs 1-4) with two to
four small segments preceding largest segment é 2
— Largest segment of the gaster the second or third (Figs 5- 8), or formed by these two segments
fused together (Fig. 7), with at most only one small segment preceding the largest : 5
2 Radial cell of forewing at least nine times as long as broad, closed on front margin (Fig. 9).
Segment 2 of gaster longer than 3 viewed laterally along the dorsal curvature, segment 1 smooth
and indistinct, segment 6 the largest. Segment 1 of tarsus (metatarsus) of hind leg as long as
segments 2-5 combined (Fig. 10) . , ‘ ‘ IBALIIDAE
— Radial cell of forewing less than nine times as long as s broad. Segment 1 of hind tarsus not as
long as 2-5 combined. Petiole distinct and sulcate (except the genera Kiefferiella and
Heteribalia) (LIOPTERIDAE) _. 3
3 Hind femur with a distinct tooth on underside (Fig. 11) between middle and apex. Mid and
hind coxae almost round and strongly swollen. Scutellum with a distinct spine at apex (Fig.12)
(exceptionally without). Segment 1 of gaster generally at least as long as wide, sulcate, segment
5 the largest with three short segments preceding it (except in Xenocynips which has 2-4 fused)
CE a2 )ees : OBERTHUERELLINAE (p. 109)
— Hind femur without a tooth on ‘the underside, mid and hind coxae elongated. Scutellum without
a spine. Segment 1 of gaster sometimes shorter than wide : 4
4 Segment 1 of gaster at least twice as long as wide and attached tangently (Fig. a segment 4 of
gaster the largest : : LIOPTERINAE
— Segment | of gaster in the form of a ring or ‘collar, never r longer than wide and attached normally
(Fig. 4); segment 5 or 6 of gaster in 2 (except in Paraegilips, a genus based on a ¢ from the
Philippines) the largest ; : MESOCYNIPINAE
5 Scutellum with a disc or cup on dorsal surface (Fig. 14). Pronotum often very pronounced,
produced frontodorsally into an anterior plate with a strong posterior margin (pronotal plate)
(Fig. 13). Forewing with cubitalis (Rs+ M) usually visible and arising from a point very close
to lower end of the basalis (near junction with M+ Cu) (Fig. 15), areolet not indicated, or
combined with median (Cu,) for a short distance. Segments 2 and 3 of gaster fused, without
visible suture. (Brachypterous forms sometimes occur.) . : EUCOILIDAE
— Scutellum without a disc or cup dorsally, sometimes with a posterior fovea or apical spine
(Fig. 33) (Aspicerinae). Pronotum if raised dorsally into an anterior aoe then without
a posterior margin, very often indistinct or absent . ; 6
6 Alate; forewing with cubitalis (Rs+M), when visible, arising from a point at or close to junction
of basalis (Rs or Rs and M) with median (Cu,) (Fig. 16), areolet indicated. Female often with
either second or third tergite of gaster the largest (Fig. 6). Male generally with cylindrical
gaster (except Aspicerinae). Mesopleuron sculptured medially at least, or scutellum with
apical spine (FIGITIDAE) . ; 7
— Winged, brachypterous or apterous. Forewing with cubitalis (Rs+M), when visible, arising
from a point nearer the middle of basalis (Rs and M) than to the junction of basalis with
median (Cu,) (Fig. 17). Female with the largest free tergite either the second (Fig. 19) or the
second and third fused together, with or without an obsolete suture visible (Fig. 18). Both sexes
12
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 91
with gaster laterally compressed ; sometimes whole insect smooth and shining. Scutellum
never with an apical spine (CYNIPIDAE) : 10
Tergite 2 of gaster liguliform (Fig. 5). Hind tibiae j in most genera longitudinally ridged or
furrowed on outer margins or posteriorly (Fig. 34). Scutellum with one or more longitudinal
carinae and subapically with either a spine or foveae. Thorax sculptured ASPICERINAE (p. 93)
Tergite 2 of gaster not liguliform (Fig. 6). Hind tibia not longitudinally ridged or furrowed
externally or posteriorly, at most with longitudinal carinae or groove internally. Scutellum
usually without three longitudinal carinae or subapical fovea, though more frequently
produced apically to form aspine. Thorax generally smooth and shining, occasionally dull
and sculptured . : 8
Tergite 2 of gaster shorter than third (Fig. 6): segment 1 never as Jong as wide. Forewing with
cubitalis (Rs+M) dividing externally before point of emission of 2rm (i.e. areolet present,
though often obsolete) (Fig. 16). : FIGITINAE
Tergite 2 of gaster longer than third (Fig. 8), segment | at least as Jong. as wide, generally more
than twice as long as wide (Fig. 8). Forewing with cubitalis (Rs+ M) dividing ena at
point of emission of vein 2rm (i.e. areolet vestigial) (Figs 20, 21) ; : 9
Head distinctly broader than thorax viewed dorsally. Gaster wedge-shaped (except i in canine:
gilips). Antenna of 2 13-segmented, 3 14-segmented. Radial cell of forewing open or closed,
cubitus vestigial, nervellus and post nervellus absent. Segment 1 of gaster at least as long
as broad, anges much longer and gaan: sade 2 of gaster longer than third (Fig.
SP 1s : ANACHARITINAE
Head small, narrower than thorax, oblong in “dorsal view. Antenna of 2 20-segmented (3
unknown). Radial cell of forewing open on front margin (R, and Rs, not aching margin of
wing), cubitus (M) almost reaching apex of forewing, areolet absent, nervellus (Cu and Cu-a)
and post nervellus (M-cu) indicated. Clypeus projecting forward and upward away from
the unsclerotized area and truncate labium. Segment | of gaster shorter than wide, tergite 2
at least half as long as remaining visible segments viewed laterally . HIMALOCYNIPINAE
Radial cell of forewing with a distinct pterostigma (Fig. 20).
Antennae of 2 15-segmented, g unknown . : : ; AUSTROCYNIPINAE
Forewing without a true pterostigma ; : : : . ek
Radial cell much reduced, closed, its veins thick and heavy (Fig. pen
Gaster with segments 2 and 3 completely fused (as in Synergus, cf. Fig. 18). Antennae of
2 12-, 15- or 19-segmented. . ' PYCNOSTIGMATINAE
Radial cell normal (at most very little reduced), open or closed os 17, 35) (apterous forms
occur) , 7 ee
Mid-tibia often with only one spur: hind tibia with one spur o or more often with two very unequal
spurs. Vertex, mesoscutum, scutellum, mesopleuron and gaster smooth and shiny. Pronotum
generally sharply angled anteriorly (Fig. 24), this forming a lateral vertical carina (Fig. 25).
Gaster with a pubescent ring at base of tergite 2. Male antenna with either the fourth, rarely
the third, sometimes the third to fifth segment modified (Figs 39,40). . ALLOXYSTINAE
Mid and hind tibiae each with two distinct spurs. Sculpture present at least on the vertex,
mesoscutum, scutellum, mesopleuron or gaster. Pronotum generally not angled sharply
anteriorly (Figs 26, 27), but if some carinae indicated (Figs 36, 37) then head and thorax
strongly sculptured. Gaster very rarely with pubescence at base of tergite 2 of gaster. Antenna
of male with modified segment, when present, always the third (Fig. 38). . CYNIPINAE
Checklist of the families and subfamilies of Cynipoidea
IBALIIDAE Thomson, 1862
LIOPTERIDAE Ashmead, 1895
OBERTHUERELLINAE Kieffer, 1903
LIOPTERINAE Ashmead, 1895
MESOCYNIPINAE Hedicke & Kerrich, 1940
EUCOILIDAE Thomson, 1862
FIGITIDAE Thomson, 1862
ASPICERINAE Kieffer, 1910
FIGITINAE Thomson, 1862
92 J. QUINLAN
ANACHARITINAE Thomson, 1862
HIMALOCYNIPINAE Yoshimoto, 1970
CYNIPIDAE Hartig, 1840
AUSTROCYNIPINAE Riek, 1971
PYCNOSTIGMATINAE Cameron, 1905
ALLOXYSTINAE Hellén, 1931
CYNIPINAE Hartig, 1840
Checklist of the Ethiopian species of Aspicerinae and Oberthuerellinae
FIGITIDAE Thomson, 1862
ASPICERINAE Kieffer, 1910
ANACHAROIDES Cameron, 1904
arcus sp. Nn.
astrida sp. n.
decellius sp. n.
elongaticornis Benoit, 1956
eurytergis Benoit, 1956
nigra sp. n.
pallida sp. n.
paragi Benoit, 1956
quadrus sp. n.
rufa (Kieffer, 1910)
sanitus sp. n.
striaticeps Cameron, 1904
spinosipes (Kieffer, 1910)
gibbosus Benoit, 1956 syn. n.
stygius Benoit, 1956
suspensus sp. Nn.
PROSASPICERA Kieffer, 1907
africana (Kinsey, 1919) comb. n.
antennata (Benoit, 1956) comb. n.
kisantua (Benoit, 1956) comb. n.
optivus sp. n.
paragicida (Benoit, 1956) comb. n.
tropica (Kieffer, 1910) comb. n.
LIOPTERIDAE Ashmead, 1895
OBERTHUERELLINAE Kieffer, 1903
OBERTHUERELLA Saussure, 1890
abscinda sp. n.
aureopilosa Benoit, 1955
breviscutellaris Benoit, 1955
crassicornis Benoit, 1955
compressa Benoit, 1955 syn. n.
lenticularis Saussure, 1890
longicaudata Benoit, 1955
longispinosa Benoit, 1955
nigra Kieffer, 1910
nigrescens Benoit, 1955
tibialis Kieffer, 1904
triformis sp. Nn.
TESSMANNELLA Hedicke, 1912
expansa sp. n.
nigra Hedicke, 1912
spinosa Hedicke, 1912
transiens Benoit, 1955
XENOCYNIPS Kieffer, 1910
subsquamata Kieffer, 1910
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 93
ASPICERINAE Kieffer
Aspicerinae Kieffer, 1910 : 50. Type-genus: Aspicera Dahlbom.
Tergite 2 of gaster less than half the total length of gaster and distinctly liguliform (Fig. 5). Hind tibia in
most genera with a longitudinal ridge. Scutellum with one or more longitudinal carinae and subapically
with a tooth or spine. Forewing with radial cell completely open on front margin (Fig. 45), invariably
open at base; wing surface generally bare, margins either bare or with short hair fringe.
Discussion. Eight genera are at present recognised as being in the subfamily Aspicerinae. Only
two of these, Prosaspicera Kieffer and Anacharoides Cameron, are known to occur in the Ethiopian
Region. Previously four species of Aspicera Dahlbom were recognised; these are now transferred
to Prosaspicera. Two other species of Aspicera have been described from North Africa and do
not occur in the region covered by this paper. They are A. aegyptica Hedicke from Egypt and
A. lobata Hedicke from Tunisia. The holotypes of these two species were lost due to bombing in
the 1939-45 war. Aspicera has been included in the key to genera in order that it may be recog-
nised if it is found to extend into the Ethiopian Region. Of the genera now recognised from the
region, Prosaspicera contains, together with the one new species described here, six species. Of
these, five are based on females, and one on a male; in only two species are males and females
correlated. The genus Anacharoides now contains 13 recognised species of which seven are
known from both males and females; six of these species are new and described here for the first
time. Two Kieffer species, spinosipes and rufa, both originally described in the genus Coelonychia,
are lost and could not be found by Dr KGnigsmann. In correspondence he indicated that they
might be in another collection of which loan records have been lost. C. spinosipes was made a
junior synonym of Anacharoides striaticeps Cameron by Weld (1930).
The Aspicerinae have many close affinities with the Figitinae but are generally distinguished
from them on the shape of tergite 2 of the gaster, which is liguliform or saddle-shaped (Fig. 5),
and the longitudinally ridged or furrowed hind tibiae. The thorax is invariably sculptured and the
scutellum has one or more longitudinal carinae and subapically a spine or foveae.
Key to genera of Ethiopian Aspicerinae
1 Scutellum with a sharply pointed apex in the form of a spine, not rounded, truncate or emar-
ginate; scutellar foveae not extending to middle of scutellum (Fig. 33). Scutellar spine with
longitudinal striations; mesonotum sculptured, rarely shining. Segment | of gaster wider than
long and crenulate. Radial cell of forewing open at base, generally with R, indicated (Figs 41,
42). Hind tibiae with hairs but without erect spines (Fig. 43). 2
— Scutellum not ending in a spine, truncated at apex (Fig. 44); scutellar foveae extending at least
half the length of scutellum, separated by a septum, the apex of each fovea enclosing a
triangular area; apex of scutellum with a margined semicircular depression (Fig. 44). Segment
1 of gaster at least twice as long as broad. Radial cell of forewing completely open at base
(Fig. 45). Hind tibiae with perpendicular spines on outer margin (Fig. 46)
ANACHAROIDES Cameron (p. 94)
2 Méesoscutum dull and conspicuously reticulate-coriaceous with medial scutal line in form of a
raised ridge starting at base, dividing at a point two-thirds down from the apical margin of the
pronotum and forming a depressed area at its apex (Fig. 47); anterior parallel lines ridged,
visible in basal third of mesoscutum; notaulices wide at apex, tapering at base, distinct and
crenulate; lateral lines indicated at apex of mesoscutum, scutellum with two foveae at base
separated by aseptum, lateral margins of scutellum reticulate-coriaceous; scutellar spine broad-
based, arising from a point near middle of truncate scutellum, measured from apex of scutellum
subequal to length of mesoscutum and tapering to a sharp point (Fig. 47); spine viewed
laterally curved upward. Vertex generally incised (Fig. 48) . PROSASPICERA Kieffer (p. 104)
— Mesoscutum shiny with fine reticulate sculpture between strong widely spaced transverse
strigose sculpture; median scutal line and parallel lines continued from base of scutellum to
centre of mesoscutum, median scutal line forming a triangular area at apex; notaulices wide,
distinct and crenulate, lateral lines indicated as ridges. Scutellar foveae separated by a
septum; lateral margins of scutellum strigose, apex of scutellum truncated, spine arising at a
94 J. QUINLAN
point close to apex of scutellum, not involving apex of scutellar foveae, distinctly shorter than
mesoscutum (Fig. 33). Vertex not incised, at most arched, occipital carinae not pronounced
ASPICERA Dahlbom
ANACHAROIDES Cameron
Anacharoides Cameron, 1904a:160, Type-species: Anacharoides striaticeps Cameron, by original
designation and monotypy.
Coelonychia Kieffer, 1910c: 19-20. Type-species: Coelonychia spinosipes Kieffer, by monotypy.
[Synonymised by Weld, 19306: 139.]
DIAGNOSIS. 2 antenna 13-segmented, weakly subclavate, sometimes moniliform; ¢ antenna filiform,
14-segmented with modified third segment. Head viewed frontally with eyes generally as far apart as the
height of an eye (cf. Fig. 74), frontal carinae extending from outer margin of antenna to lateral ocellus,
occiput strongly striated, occipital carinae distinct. Pronotum angular on anterior margin (cf. Fig. 44).
Mesoscutum with notaulices distinct, median scutal line, anterior parallel and lateral lines present or
indicated. Scutellum viewed dorsally with apical depression, base of scutellum with foveae. Mesopleurae
generally sculptured on lower half, with variable sculpture above to a completely polished surface.
Gaster with segment 1 (petiole) longer than broad, smooth, segment 2 liguliform, segment 3, viewed
laterally, the largest, generally punctate on lateral and dorsal surfaces. Hind tibia longitudinally ridged
or furrowed with distinctive suberect spines on outer margins (Fig. 46). Radial cell of forewing open on
base and front margin.
DISTRIBUTION. South Africa; Malawi.
Key to species of Anacharoides Cameron
Females (unknown in astrida)
1 Apical segment of antenna longer than segment 3 and longer than 11 plus 12, segments 3 and 4
not swollen (Fig. 59).
Scutellum with apical depression rounded on basal margin; scutellar foveae angled with a
small triangular depression at base (Fig. 60), apex of scutellum arched
elongaticornis Benoit (p. 98)
— Apical segment of antenna equal to or shorter than segment 3, only rarely as long as segments 11
plus 12 (Fig. 87) ; 2
2 Inner basal margin of apical scutellar depression viewed dorsally almost straight medially, side
margins of depression straight, rarely converging basally, apex of depression sometimes
arcuate, the whole more generally appearing square (Figs 61,62) : b
— Inner basal margin of apical scutellar depression strongly arched medially, sometimes arcuate on
apical margin (Fig. 63), side margins of apical scutellar depression converging towards base,
viewed dorsally, apical scutellar depression usually wider than long measured at centre
(Fig. 63)" —. F
3 Mesoscutum medially with strong widely spaced broken transverse rugae ona ‘reticulate surface,
anterior parallel lines distinct anteriorly in first quarter, not converging (Fig. 64).
Hind coxae swollen, strongly coriaceous and pubescent with a smooth shiny area on inner
margin. Forewings with veins Sc+ R angled at a point where R, would normally emanate
(Fig. 65). Rs, weakly projecting along wing margin. Antennal segment 13 shorter than 3
(Fig. 66) . : r stygius Benoit (p. 103)
— Méesoscutum either with closely spaced regular vugee or finely reticulate-coriaceous j : ie
Forewing with Rs, projecting along wing margin (Fig. 67). Pronotum without a median tooth . 5
— Forewing with Rs, not projecting along wing margin (Fig. 68). Pronotum with a small tooth
medially (Fig. 82) : : ; suspensus Sp. n. (p. 104)
5 Anterior parallel lines distinct, converging to join scutal furrow, “notaulices crenulate, wide at
apex, narrow basally. Rs, of forewing not reaching wing margin but weakly extended, Rs, of
radial cell not sharply angled (Fig. 67) . 5 : paragi Benoit (p. _
— Anterior parallel lines obsolete. Rs, of forewing sharply angled or not angled
6 Mesoscutum reticulate medially with rugae at apex and base, notaulices distinct, not crenulate,
pubescence weak on outer margins of notaulices, median scutal line complete, raised above
the surface sculpture, triangular area extending halfway along mesoscutum; scutellar foveae
BSS
10
11
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 95
elongate, shiny reticulate. Apical segment of antennae dark, remainder orange-yellow. R, of
forewing weakly indicated (Fig. 69) ; ; : sanitas sp. n. (p. 102)
Mesoscutum finely reticulate, notaulices crenulate, complete, whole surface of mesoscutum
covered with hairs obscuring notaulices, median scutal line complete, obscured by pubescence;
apical depression of scutellum angular at apex; scutellar foveae round, smooth and polished.
Apical segment of antennae variable in colour, remaining segments orange-yellow. R, of
forewing not indicated : b quadrus sp. n. (p. 101)
Mesoscutum with widely spaced strong irregular transverse rugae medially on finely reticulate
surface; notaulices with crenulations, median scutal line extending at least to the middle of
the mesoscutum (Fig. 63).
Antennae yellow or yellow with the apical segments dark yellow . F 8
Mesoscutum finely reticulate-coriaceous, at most with weak transverse rugae on outer margins of
notaulices; median area finely reticulate, notaulices either distinct or in the form of widely
spaced crenulations, median scutal line either complete or not reaching past middle of
mesoscutum : 9
Mesoscutum at most with fine scattered pubescence, surface finely reticulate- -coriaceous between
widely spaced transverse rugae, notaulices with widely spaced crenulations, pubescence
sparse, not obscuring sculpture, anterior parallel lines converging to join at median scutal line
(Fig. 63), apex of median scutal linein form of a small triangular polished area apically, fading
out before reaching middle of mesoscutum. Sc+ R of forewing weakly angled at point where
R, would emanate if present, Rs, strongly curved, weakly extended along the wing margin,
2rm short and broad (Fig. 86) : : ; striaticeps Cameron (p. 103)
Mesoscutum with long scattered pubescence, its surface reticulate- -coriaceous between strong
transverse rugae; notaulices wide with close crenulations but obscured by scattered pubes-
cence, anterior parallel lines obscured by whitish pubescence, median scutal line obscured by
long hairs, not distinctly widened at apex, merging into the transverse rugae of mesoscutum.
Radial cell of forewing with Sc+ R angled at point close to where R, would emanate, Rs,
weakly curved, strongly extended along wing margin, 2rm long and narrow (Fig. 70)
arcus sp. n. (p. 96)
Forewing with Rs, not reaching wing margin, extended weakly below margin (Fig. 71). Median
scutal line distinct. Head, mesoscutum and mesopleurae either completely black or reddish
orange : 10
Forewing with Rs, reaching wing margin, not extended along margin ( Fig. 72). Median scutal
line obsolete, at most with weak triangular area at apex. Head and mesoscutum never com-
pletely black, variably red- to black-marked . , 11
Median scutal line complete with a triangular depression at apex, deeply impressed, extending
to middle of mesoscutum (Fig. 75). Head, mesonotum and mesopleurae completely black,
gaster bright yellow. é ' . nigra sp. n. (p. 117)
Median scutal line incomplete with a ‘small triangular shallow depression at apex not extending
to middle of mesoscutum (Fig. 76). Head, mesonotum and mesopleurae reddish orange to
brownish, gaster chestnut-brown . : ; : decellius sp. n. (p. 97)
Mesoscutum finely reticulate-coriaceous with weak transverse rugae On inner margins basally,
median area without pubescence, median scutal line obsolete in basal half, triangular area at
base distinct, extending to a point midway on mesoscutum (Fig. 77), notaulices complete,
distinct with close shallow Seated scutellar foveae deep basally, polished. Colour
orange-yellow. ; pallida sp. n. (p. 100)
Mesoscutum densely pubescent ona . fine reticulate- -coriaceous surface, notaulices with widely
spaced crenulations obscured by pubescence, median scutal line obsolete basally, visible in
apical quarter, triangular area at apex weak and obscure (Fig. 73); scutellar foveae shallow.
Viewed dorsally, head orange-yellow, mesoscutum dark brown except apex orange-yellow,
scutellum orange-yellow, gaster chestnut-brown : ‘ 4 eurytergis Benoit (p. 100)
Males (unknown in elongaticornis, nigra, pallida, paragi and stygius)
1
Segment 3 of antenna weakly expanded apically, slightly curved, never with flattened surface on
basal half (Fig. 78). ; 2
— Segment 3 of antenna generally strongly expanded apically and medially, always with basal half
distinctly narrower than apex and flattened (Fig. 79) ‘ ; 3
2 Segments 4-14 of antenna moniliform, not swollen distally; notaulices distinct and crenuilate,
anterior parallel lines converging to join median scutal line; apical depression of scutellum
96 J. QUINLAN
without triangular-shaped area at base; median scutal line with obsolete horseshoe-shaped
area at apex, mesoscutum reticulate-coriaceous with scattered pubescence quadrus sp. n. (p. 101)
— Segments 4-6 of antenna weakly swollen distally (Fig. 80); notaulices weakly crenulate, apical
depression of scutellum viewed dorsally with pronounced triangular-shaped area at base,
median scutal line strongly raised, the apical area in form of acute triangular shape extending
to middle of mesoscutum, mesoscutum reticulate with scattered pubescence on outer margins
of notaulices : arcus sp. n. (p. 96)
3 Segment 3 of antenna strongly expanded on outer apical margins, ‘segments 4-14 moniliform or
only very weakly expanded medially (Figs 79, 84) . , 4
— Segments 3 and 4 of antenna swollen or expanded apically on outer margins, base of segments
clearly narrower than apex (Fig. 81) ; ; 5
4 Mesoscutum with closely spaced strong transverse rugae between notaulices, median scutal line
triangular at apex and transversely striated (Fig. 82), notaulices distinct, not obscured by
the fine pubescence. Head and thorax predominantly red-reddish brown, hind coxae strongly
swollen (Fig. 83) : : ; suspensus sp. n. (p. 104)
— Méesoscutum without transverse rugae, surface fine reticulate- coriaceous, notaulices distinct but
obscured by fine scattered pubescence. Head and thorax completely black, hind coxae long
and narrow (Fig. 85) . , : decellius sp. n. (p. 97)
5 Segments 3 and 4 of antenna swollen at apex, 5- 14 moniliform (Fi ig. 87) striaticeps Cameron (p. 103)
— Segments 3-5 of antenna swollen on outer margins (Fig. 88) F , 6
6 Segments 3-5 of antenna distinctly swollen on outer margins apically, a 14 moniliform,
notaulices distinct, without crenulations, mesonotum finely reticulate-coriaceous. Head black
suffused with red, thorax reddish black . ; ; . Sanitas sp. n. (p. 102)
— Segments 3-6 of antenna swollen apically. Head and thorax completely black : ~
7 Segments 3-6 of antenna distinctly swollen apically, 7-14 moniliform (Fig. 91). Mesoscutum fine
reticulate-coriaceous with closely spaced broken rugae medially, not pronounced
eurytergis Benoit (p. 99)
— Segments 3-7 of antenna distinctly swollen apically (Fig. 90), segment 8 very weakly swollen,
9-14 moniliform. Mesoscutum fine reticulate-coriaceous with widely spaced transverse rugae,
notaulices deeply crenulate . d j ; ; : ‘ ; . astrida sp. n. (p. 97)
Anacharoides arcus sp. 0.
(Figs 44, 70, 80)
DESCRIPTION. 2. Antenna 13-segmented, segment 3 longer than apical segment, shorter than 4 and S.
SN 3-13=AR S50: 28: 28: 25: 25: 20: 20: 20: 20: 20: 45. ¢ antenna 14-segmented, segment 3 subequal
in length to 4 plus 5, 3 weakly curved on outer margin, 4, 5 and 6 expanded apically (Fig. 80). Head
viewed frontally with eyes closer together than height of eye measured midway between eyes, vertex
incised, frontal carinae distinct, face reticulate-rugose, pubescent on the inner orbits, occiput strongly
striated, strigose between lateral ocellus, vertex incised, pronotum viewed dorsally angled on the basal
side margins, mesoscutum with reticulate surface sculpture and transverse rugae, anterior parallel lines
aberrant, obscured by pubescence, notaulices in form of weak crenulate suture, complete, median scutal
line indistinct basally, ending in weak triangular area apically, obscured by the surface sculpture of
mesoscutum, scutellar fovea extending half the length of scutellum, finely sculptured, the apex of each
fovea triangular, separated by a septum, apical depression of scutellum, arcuate with strong raised
margins, lateral bars of scutellum reticulate. Mesopleural carinae distinct, the surface below carinae
sculptured and pubescent, carinae of propodeum bowed. Segment 1 of gaster petiolate, sulcate, three
times as long as broad, segment 2 of gaster liguliform, segment 3 the largest, closely punctate on apical
half in dorsal and lateral views, propodeum and coxae strongly pubescent. Hind tibia with longitudinal
furrow and suberect spines on outer margin viewed dorsally. Colour: 2 antenna dark yellow, apical
segments dark, legs brownish yellow, head brownish orange, pronotum reddish orange on front margin;
mesopleura dark brownish red; mesoscutum blackish on front and side margins; apex reddish orange;
scutellum reddish orange; gaster chestnut-brown; 3, antenna and legs yellow, head and thorax black,
scutellum and gaster chestnut-brown. Radial cell of forewing completely open at base and on wing margin,
veins Rs+M not indicated, Rs, weakly extended along margin of wing, R, not indicated, wing surface
with long scattered cilia on apical third and scattered shorter ciliae on remainder, wing margin with a
distinct hair fringe apically.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 97
MATERIAL EXAMINED
Holotype 9, Rwanda: env. Astrida, 1954/1955 (G. Foucart) (MRAC, Tervuren).
Paratypes. Rwanda: 1 3, same data as holotype (badly damaged) (MRAC, Tervuren). Uganda: 1 9,
Kampala, 24.x.1921 (H. Hargreaves) (BMNH).
REMARKS. This species is closely related to striaticeps but is distinguished by the long scattered
pubescence on the mesoscutum.
Anacharoides astrida sp. n.
(Fig. 90)
DESCRIPTION. ¢. Antenna 14-segmented. Segment 3 of antenna as long as 3 plus 4, 3-8 each strongly
swollen, remaining segments moniliform (Fig. 90). Head viewed frontally strongly reticulate and pubescent,
eyes as far apart as the height of an eye, vertex arcuate, frontal carinae on inner orbits of eye crenulate,
head viewed dorsally with frontal carinae extending from lateral ocellus, enclosing a finely reticulate area
with transverse rugae, occiput strongly striated. Pronotal plate produced, pubescent either side with
pronounced foveae medially; pronotum viewed dorsally angular, pubescent, mesoscutum finely reticulate
with widely spaced irregular rugae striate on side margins, anterior parallel lines distinct in basal third,
median scutal line complete, widened in apical quarter, notaulices complete, crenulate, obscured by the
pubescence on the surface of mesoscutum; scutellum with apical depression almost quadrate, weakly
arched on basal margin, scutellar foveae rounded basally, wider than apical half with fine reticulate area
between apex and apical depression, apex of depression almost straight, sides of scutellum with large
reticulate-rugose surface sculpture. Mesopleurae below mesopleural suture densely pubescent; carinae of
propodeum almost parallel. Gaster with segment 1 smooth, petiolate, twice as long as broad, segment 2
liguliform, segment 3 viewed laterally the largest, densely punctate, remaining segments not visible.
Hind tibia longitudinally ridged or furrowed with pronounced suberect spines on outer margin. Colour:
antenna and legs orange-yellow, head and thorax black, gaster dark chestnut-brown. Forewing with short
apical hair fringe, surface with widely scattered short pubescence, Rs, of forewing weakly produced on
wing margin, R, absent, i.e. radial cell completely open at base, Sc+R not COUSPICUO MEY angled at
point where R, would normally emanate.
2 not known.
MATERIAL EXAMINED
Holotype 3, Rwanda: env. Astrida, 1954/1955 (G. Foucart) (MRAC, Tervuren).
Paratypes. Tanzania: 1 3, Ngorongoro, Pest Camp, 2400-2500 m, 6-19.vi.1957 (P. Basilewsky &
N. Leleup) (MRAC, Tervuren). Zaire: 1 ¢, Kivu Sac (Lac Kivu), 1460 m, 19-22.ii1.1934 (G. F. de Witte)
(MRAC, Tervuren); 1 ¢ Secteur Tshiaberimu, Clieu-dit, 2450 m, 24.iii1.1954 (P. Vanschuytbroeck &
H.. Synave) (MRAC, Tervuren); 1 3, P.N.A. Secteur Tshiaberimu, Riv. Kalvina, affl. dr. Talia N. 2720 m,
27.111.1953 (P. Vanschuytbroeck & V. Hendricx); 1 3, Secteur Tshiaberimu, riv. Talia Nord, 2340 m,
28.i11.1954 (P. Vanschuytbroeck & H. Synave) (MRAC, Tervuren).
REMARKS. Distinguished from eurytergis Benoit by the seventh antennal segment being swollen
apically.
Anacharoides decellius sp. n.
(Figs 76, 84, 85)
DESCRIPTION. 2. Antenna 13-segmented, weakly subclavate, segment 3 longer than apical segment,
shorter than 4 and 5. SN 1-13=AR 5:3:18:12:12:11:9:9:10:10:9:9: 10. The apical segment
darker than remainder. ¢ antenna 14-segmented, segment 3 expanded apically, 3 and 4 very weakly
swollen medially on outer margin, remaining segments moniliform. Head viewed frontally with eyes as
far apart as height of eye, vertex incised, frontal carina extending from outer margin of antenna to lateral
ocellus, enclosing a reticulate area with weak transverse rugae, face below antenna with dense pubescence;
occipital carinae pronounced, occiput strongly striate. Pronotum viewed dorsally, angular on side
margins which are striated, with a small pronotal plate medially. Mesopleurae smooth and polished,
mesosternum pubescent, mesonotum finely reticulate, notaulices distinct, not crenulate, median scutal
line visible in apical quarter only as a triangular area, anterior parallel lines absent, surface of meso-
scutum with fine scattered pubescence; scutellar foveae extending past middle of scutellum deep and
polished, angular at apex, apical depression of scutellum rounded on basal margin, scutellum with
98 J. QUINLAN
triangular area medially between foveae and apical depression, sides of scutellum strongly rugose, propo-
deum smooth and polished, carinae almost parallel, bowed at either end. Gaster with segment 1 smooth
and polished, twice as long as wide, segment 2 liguliform, segment 3 the largest, closely punctate on
lateral and dorsal margins, sides of propodeum and coxae pubescent, hind tibia with longitudinal furrow
and suberect spines on outer margin viewed dorsally. Colour: 2 antenna orange-yellow except apical
segment dark brownish, legs brownish with yellowish tarsal segments, head reddish yellow, pronotum
reddish dorsally, brownish laterally, mesoscutum black dorsally, scutellum reddish yellow, gaster chestnut-
brown; 3 head and thorax black. Radial cell of forewing completely open at base and on wing margin,
veins Rs+M not indicated, Rs, weakly extended along margin of wing. Wing surface ciliate, apical
margin of wing with hair fringe, surface of wing with short scattered hairs apically.
MATERIAL EXAMINED
Holotype °, South Africa: Natal, Kloof, 1500 ft [450 m], August (R. E. Turner) (BMNH),.
Paratypes. Ethiopia: 8 2, Mt Chillala Forest, circa 9000 ft [2700 m], 12.xi.1926 (H. Scott); 1 3, same
data except 27.xi.1926 (BMNH); 1 3, Harrar, iii.1939 (H. C. Taylor) (BMNH). Kenya: 1 2, Chiromo,
7.viii.69, ex Allograph calopus (BMNH). Nigeria: 1 2 Njawai, NE. State, 24.viii.1973 (J. 7. Medler)
(BMNH). Rhodesia: 1 2, Chishawasha, nr Salisbury, ii.1978 (A. Watsham) (BMNH). South Africa:
6 2, Port St Johns, Pondoland, 1-14.v.1923 (R. E. Turner) (BMNH); 2 2, same data except 15-31.v.1923
(BMNH); | 2, same data except 5—30.iv.1923 (BMNH); 2 9, same data except 12-30.vi.1923 (BMNH);
1 2, same data except 10-31.vii.1923 (BMNH); 4 3, same data except 12-30.vi. (R. E. Turner) (BMNH);
1 d, same data except 1-11.vi.1923 (BMNH); 1 2, Natal Kloof, 1500 ft [450 m], viii.1926 (R. E. Turner);
1 3, Natal, Weenen, vi.1925 (H. P. Thomasset) (BMNH); 1 °, Cape Province, Mossel Bay, v.1930 (R. E.
Turner) (BMNH); 1 2, same data except x.1931 (BMNH) | 3, same data except v.1930 (R. E. Turner)
(BMNH); 1 9°, Assegaaibosch, E. Stellanbosch, 4-5.i.1972 (BMNH). Uganda: 1 3, Kazhara, iii.1939
(H. C. Taylor) (BMNH). Zaire: 2 2, Secteur Nord, riv. Butahu, affl. Semliki, 1420 m, 26.vi.1957 (P.
Vanschuytbroeck) (MRAC, Tervuren); 1 2 Mont Hoyo, 1250 m, ‘sur plantes bases’, 1—-15.vii.1955 (P.
Vanschuytbroeck) (MRAC, Tervuren); 1 2, Secteur Tshiaberimu, Mon. Kitwa, 2840 m, 29. viii.—7.ix.1953
(P. Vanschuytbroeck & V. Hendricx) (MRAC, Terveren); 1 2, Massif Ruwenzori, ri. Kaklari, affl. Bombi
(P. Vanschuytbroeck & H. Synave) (MRAC, Tervuren).
REMARKS. This species has close affinities with nigra but lacks the complete median scutal line
and bright yellow gaster. The male resembles suspensus but is distinguished by the absence of
transverse rugae on the mesoscutum.
Anacharoides elongaticornis Benoit
(Figs 45, 59, 60)
Anacharoides elongaticornis Benoit, 1956 : 201. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antenna 13-segmented, moniliform, 3 shorter than 4 plus 5, 4 and 5 equal, 6-12 subequal
in length, 13 twice length of 12 (Fig. 59), 5-13 with rhinaria indicated. SN 1-13=AR 4:2:8:5:5:5:5:
5:5:4:5:4:5:4:5:4:4:9. Head viewed from the front with eyes as far apart as height of an eye,
frontal carina extending from outer margin of antenna to lateral ocellus, enclosing a rugose area, vertex
incised, occiput strongly striated with strong curved striae. Pronotum viewed dorsally strongly coriaceous,
angled on margins, mesoscutum with strong coriaceous-reticulate surface, anterior parallel lines indicated,
notaulices complete though obscured in sculpture and by pubescence, median scutal line weakly indicated
posteriorly; scutellum viewed laterally with pointed apex, fine coriaceous, viewed dorsally with apical
depression, scutellar foveae not extending past middle of scutellum, separated by a septum, the surface
coriaceous, the apex of each fovea angled, with a small triangular depression in front of apical depression,
lateral bars with fine coriaceous sculpture, apex of scutellum arched (Fig. 60), propodeum with bowed
carinae, sides densely pubescent. Mesopleura smooth and polished, episternum with longitudinal striae.
Segment | of gaster petiolate, twice as long as wide, segment 2 liguliform, segment 3 the largest, closely
punctate on lateral and dorsal margins of the upper apical half. Hind tibia with longitudinal furrows and
suberect spines on outer margins, basitarsi as long as remaining combined segments.
Colour: antenna yellowish, head dark blackish red, thorax black, gaster black, front and middle legs
yellowish, hind legs blackish brown, tarsi yellow. Forewing surface with widely scattered short pub-
escence, apical hair fringe very short, radial open on front and basal margins, Rs, almost closing wing
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 99
margin (vein at juncture with wing margin barely pigmented) (Fig. 45), veins Rs+M not indicated, M
(cubitus) weakly indicated.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Uele Paulis, xii.1947 (P. L. G. Benoit) (MRAC, Tervuren).
Anacharoides eurytergis Benoit
(Figs 2..9,5,.7 45.91)
Anacharoides eurytergis Benoit, 1956: 195-196. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DEscRIPTION. @. Antenna 13-segmented, moniliform, 3 shorter than 4 plus 5, 4 and 5 subequal, 5 and 6
subequal, 6 longer than 7, 7 and 8 subequal, 8 longer than 9, 9 shorter than 10, 10 equal to 7, 10 and
11 equal, 11 shorter than 12, 13 twice as long as 12. SN 1-13=AR 8:3:11:65:6:6:5:5:45:5:5:
5-5: 11, segments 6-13 of equal width, Ist segment the widest. ¢. Antenna 14-segmented, segment 3 the
longest, strongly excavated on outer margin, 4-6 distinctly swollen. SN 1-14=AR 63:3:14:10: 10:
10:10:8:8:8:8:8:8:11-5. Head viewed frontally with eyes as far apart as the height of an eye,
frontal carina extending from outer margin of antenna to lateral ocellus, enclosing a finely reticulate,
coriaceous area with transverse rugae, lower face pubescent, vertex incised (Fig. 74), occiput strongly
striate with a strong occipital carina. Thorax: pronotum viewed dorsally strongly sculptured, obliquely
angled apically with a small plate produced forward on front margin, surface of mesoscutum reticulate-
coriaceous, obscured by dense pubescence, notaulices with widely spaced crenulations, median scutal
furrow obsolete basally, visible apically, anterior parallel lines indicated, scutellum viewed laterally with
pointed apex, reticulate rugose, viewed dorsally with apical depression; scutellar fovea extending past
middle of scutellum, foveae separated by a septum and highly polished on basal half, apical half of each
fovea elongated and weakly strigose, posterior part margined and excavate, without median carina;
propodeum with bowed carinae. Mesopleurae smooth and polished, episternum sculptured. Segment | of
gaster petiolate, longer than broad, smooth segment 2 of gaster liguliform, segment 3 the largest, closely
punctate in the lateral and dorsal areas of the upper apical half. Hind tibia with a longitudinal furrow and
suberect spines on outer margins. Colour: ¢, head, apex of mesoscutum, thorax and mesopleurae red,
gaster black, front and middle legs yellow to brown, hind legs brownish black; 3, head, thorax and hind
legs black, antenna reddish brown basally, black at apex, front and middle legs brownish yellow. Surface
of forewings with widely scattered short pubescence, apical hair fringe short, radial cell completely open
at base and on wing margin, R, not indicated, Sc+ R at point where R, would emanate, angled (Fig. 72),
veins Rs+M not indicated, Rs, not reaching margin of wing.
MATERIAL EXAMINED
Holotype 9, Zaire: Kivu, Ibanda, 1952 (M. Vandelannoite) (MRAC, Tervuren).
Nigeria: 2 2, W. State, Owena, ii.1970 (J. T. Medler) (BMNH); 1 2, Umuahia, Crin, EC. State, 10.iv.
1975 (J. T. Medler) (BMNH). Rhodesia: 3 3, Chishawasha, nr Salisbury, 11.1978 (A. Watsham) (BMNH.)
South Africa: 1 2, 5 ¢ Grahamstown, Southwell Rd, em. xi.1975, ex pupa of /schiodon aegyptus reared on
Aphis nerii on Gomophocarpus physocarpus (J.B.) (BMNH); 4 2, Zululand, Eshowe, vii.1926 (R. E. Turner)
(BMNH); 3 9, Port St. Johns, Pondoland, 1—14.v.1923 (R. E. Turner) (BMNH); 2 2, same data except date,
1-11.vi.1923; 1 9, same data except 5—30.iv.1923; 1 9, same data except 12-30.vi.1923; 1 3d, same data
except 15—31.v.1923 (R. E. Turner) (BMNH); | 2, Asseganibosch, E. Stellenbosch, 4—5.i.1972 (BMNH);
2 2, Natal, Kloof, 1500 ft, 450 m, viii.1926 (R. E. Turner) (BMNH); | 2, Cape Province, Somerset East,
x.1930 (R. E. Turner) (BMNH). Zaire: 2 3, Secteur Tshiaberimu, riv. Talia, Nord, 2340 m, 28.iii1.1954
(P. Vanschuytbroeck & H. Synave) (MRAC, Tervuren); | 3, same data except 23.iii1.1954 (MRAC,
Tervuren); | 3, Secteur Tshiaberimu Husiangwa aff. Musabaki, 19.iii.1954 (P. Vanschuytbroeck & H.
Synave) (MRAC, Tervuren); | 3 (allotype), same data as holotype; | 3 (paratype), Rwanki, 15.11.1952
(J. Leroy) (MRAC, Tervuren).
Anacharoides nigra sp. n.
(Figs 71, 75, 105)
DEscrIPTION. ©. Antenna 13-segmented, subclavate, segment 3 subequal to apical segment, shorter than
4+-5, segment 3 weakly curved apically, 4-12 moniliform, each progressively shorter than the preceding
100 J. QUINLAN
one, SN 3-13=AR 50: 30: 30: 25:25: 25:24:20: 20:20:47. Head viewed frontally with eyes as
far apart as height of an eye measured midway between the eyes, frontal carina pronounced, extending
from outer margin of antenna to lateral ocellus, face coarsely reticulate rugose with fine pubescence,
vertex incised, occiput strongly striated. Lateral margins of pronotum viewed dorsally angular. Meso-
scutum dull, finely coriaceous, with very weak scattered transverse aberrant rugae and hairs, median
scutal line visible in apical two-thirds, forming a long triangular impressed area, notaulices distinct,
crenulate, wider at apex than at base, anterior parallel lines obsolete, lateral lines weakly indicated;
scutellum viewed laterally with pointed apex dorsally with apical depression, strongly arched on basal
margin. Scutellar foveae extending to middle of scutellum with apical area merging into a triangular shape
in front of apical depression, sides of scutellum variably reticulate, carinae of propodeum parallel at
midway point, converging at top; mesopleurae smooth and polished. Gaster with segment 1 petiolate,
weakly furrowed, less than three times as long as broad, segment 2 liguliform, segment 3 the largest, closely
punctate on apical two-thirds. Hind tibia longitudinally ridged or furrowed with suberect spines on the
outer margins, hind coxae viewed dorsally longer than petiole, strongly swollen with long pubescence on
outer margins, inner margins flattened. Colour: antenna, legs and gaster orange-yellow, head and thorax
black. Forewings with apical hair fringe, surface with a few dotted hair bases at basal half, Rs, of forewing
weakly produced, not reaching margin of wing, vein R, absent, i.e. radial cell open at base and on wing
margin. Sc+ R angled at point where vein R, would normally be.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Uganda: Kawanda, viii.1943 (7. H. C. Taylor) (BMNH).
Paratype. Zaire: 1 9°, P. N. A. Secteur Tshiaberimu Riv. Mbulikere, aff. dr. Talia N. 2700 m, 26-28. viii.
1953 (P. Vanschuytbroeck & V. Hendricx) (MRAC, Tervuren).
REMARKS. Closely related to dece//ius but distinguishable on characters given in the key.
Anacharoides pallida sp. n.
(Figs 77, 93)
DEscCRIPTION. 2. Antenna 13-segmented, weakly subclavate, segment 3 shorter than 4+-5, 4 and 5 subequal,
6 and 7 subequal, 8 shorter than 7, 9-12 shorter than 8, subequal. SN 1-13 =AR 12:5: 4:5: 22:5: 12:5:
12°5:11:11:10:7:5:7:5:7:5: 7:5: 15. Head viewed frontally with eyes as far apart as the height of
an eye, frontal carinae crenulate, extending from outer margin of antenna to lateral ocellus, enclosing a
fine rugose area with strong transverse rugae in upper area, vertex incised, subantennal sutures pro-
nounced, face pubescent, occiput strongly striated. Pronotum viewed dorsally obliquely angled with a
small protrusion medially; surface of mesoscutum finely reticulate-coriaceous with traces of weak
transverse rugae on inner margins basally, anterior and lateral lines obsolete, median scutal line very
weakly present in basal half, diverging to form a small obscure triangular area at apex, weakly strigose;
notaulices complete and distinct with weak shallow crenulations. Scutellum viewed laterally, pointed,
dorsally with apical depression straight on apical margin, base of depression arched, scutellar fovea
elongate, polished, not extending past middle of scutellum, scutellar foveae separated by a broad septum,
lateral bars broad, not sculptured, apical depression with a triangular area between its base and apex of
scutellar foveae, scutellar disc with strong reticulate sculpture, mesopleurae polished with light pubescence
below the mesopleural suture, carinae of propodeum weakly bowed. Segment | of gaster slightly longer
than broad, petiolate, smooth and polished, segment 2 liguliform, segment 3 the largest, strongly punctate,
remaining segments enclosed by the third. Hind coxae swollen, weakly pubescent, hind tibiae longi-
tudinally ridged or furrowed with suberect spines on the outer margins. Colour: antenna orange-yellow,
head, thorax and legs reddish orange, gaster reddish orange dorsally, yellow basally. Forewings with
apical hair fringe, apical surface with scattered hairs, remainder dotted with hair bases, Rs, of forewing
extending to margin of wing, not extended along margin, radial cell completely open at base, Sc+R
angled at point where vein R, would normally be.
3 unknown.
MATERIAL EXAMINED
Holotype 2, South Africa: E. Cape Province, Kathberg, 11—18.ii.1938 (R. E. Turner) (BMNH).
REMARKS. This species has close affinities with eurytergis but lacks the dense pubescence.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 101
Anacharoides paragi Benoit
(Fig. 67)
Anacharoides paragi Benoit, 1956 : 200. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antenna 13-segmented, moniliform, weakly clavate, rhinaria present on segments 3-13,
3 shorter than 4+5, segments 6-12 gradually decreasing in length, segment 3 twice the length of 8, 9, 10,
11 or 12, 13 equal in length to segment 3, as long as 11+12. SN 1-13=AR 6:2:8:7:5:5:5:4:4:
4:4:4:8. Head viewed from front with eyes as far apart as height of an eye, frontal carina extending
from outer margin of antenna to lateral ocellus and enclosing a reticulate area, vertex incised, occiput
strongly striate with strong occipital carinae. Pronotum viewed dorsally obliquely angled apically,
mesoscutum reticulate rugose; notaulices complete and distinct with crenulations on apical section, the
side margins of mesoscutum with short pubescence, median scutal furrow indicated posteriorly, anterior
margins of mesoscutum with strigose sculpture; scutellum viewed laterally with pointed apex, reticulate
rugose, viewed dorsally with apical depression, scutellar foveae round, extending past middle of scutellum,
separated by a septum, apex of scutellum with large depression, straight on apical margin, sides of
scutellum strongly reticulate rugose; carinae of propodeum weakly bowed at anterior. Mesopleura
smooth and polished, suture distinct. Segment 1 of gaster petiolate, longer than broad, smooth,
segment 2 of gaster liguliform, segment 3 the largest, closely punctate dorsally and laterally. Hind tibia
with longitudinal furrows with dense comb of hairs on inner margin and a row of widely spaced spines on
outer margin. Colour: antenna, head and thorax reddish brown, legs yellow to reddish brown, gaster
chestnut-red-black. Surface of forewing with widely scattered hair bases and a few short hairs near apex,
hair fringe on wing margin very short, vein Rs, thin, not reaching margin, R, absent, Rs+M and M not
indicated (Fig. 67), Sc+R angled at point where R, would emanate.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Gandajika, 22.111.1947 (P. Henrard) (MRAC, Tervuren).
Ghana: | 2, Kumasi, Kwadaso, 28.iii.1977 (Scheibelreiter) (BMNH). Nigeria: 1 2, Umuahia Crinec
State, 10.iv.1975 (J. T. Medler); 1 2, Zungeru, xi.1910 (J. W. Scott-Macfie) (BMNH); 4 8, Ile-Ife, W.
State, v.1973 (J. T. Medler) (BMNH). Rhodesia: Chishawasha, nr Salisbury, ii.1978 (A. Watsham)
(BMNH). Sierre Leone: 2 2, Njala, 2—15.i.1932 and 1931, ex syrphid puparium (£. Hargreaves) (BMNH).
Uganda: 2 2, Kampala, 6.x.1928 and 8.x.1928, ex Paragus burbonicus (G. L. R. Hancock) (BMNH);
1 9, Kawanda, vi.1941 (7. H. C. Taylor). Zaire: 1 ° (paratype), same data as holotype except 5.iii.1948
(MRAC, Tervuren); 1 2, Haut-Uele, Moto, iv—v.1923 (L. Burgeon) (MRAC, Tervuren); 1 2, Leverille,
ix.1920 (P. Vanderjist) (MRAC, Tervuren).
Anacharoides quadrus sp. n.
(Figs 78, 89)
DEscRIPTION. ?. Antenna 13-segmented, very weakly subclavate, segment 3 longer than 4, shorter than
4+-5, 4-6 subequal in length, 6-12 gradually decreasing in length, 13 as long as 3. SN 1-13=AR 10:3:
15:9:9:9:9:7:7:7:7:6:15; apical segment darker than remainder. 3 antenna moniliform, 14-
segmented, segment 3 weakly expanded apically (Fig. 78), SN 1-5 =AR 15: 8:45: 30: 30. Head viewed
frontally with eyes as far apart as height of eye, vertex incised, frontal carinae extending from outer
margin of antennae to lateral ocellus, enclosing a reticulate area covered with scattered long hairs,
occipital carina pronounced, occiput strongly striate. Lateral margins of pronotum with striations,
pronotum viewed dorsally, angular, with a small pronotal plate on the front margin, no tooth present.
Mesoscutum finely reticulate, covered with long white widely spaced hairs, median scutal line obscured
by pubescence, notaulices complete with wide-spaced crenulations, obscured by long hairs, anterior
parallel lines obsolete. Scutellum viewed laterally with pointed apex, dorsally with apical depression, not
strongly arched on basal margin (almost straight), scutellar foveae elongate, extending past the middle of
scutellum, basal half of foveae with triangular carinae (Fig. 89); sides of scutellum strongly rugose,
carinae of propodeum almost parallel. Mesopleurae smooth and polished above the suture, densely
pubescent below. Gaster with segment | petiolate, smooth, segment 2 liguliform, segment 3 viewed
laterally the largest, closely punctate in dorsal and lateral margins, remaining visible segments punctate.
Hind tibia longitudinally ridged or furrowed with pronounced suberect spines on outer margins. Colour:
2 antenna, head, thorax and legs orange-red, gaster chestnut-brown, 3 antenna, fore and mid legs yellowish,
102 J. QUINLAN
head, mesonotum predominantly black, scutellar foveae reddish, hind legs brownish, gaster chestnut-
brown. Forewings with apical hair fringe, surface with scattered hairs, Rs, of forewing produced along
wing margin, vein R, absent i.e. radial cell completely open at base, Sc-+ R angled at point where vein
R, would normally be.
MATERIAL EXAMINED
Holotype 2, Rhodesia: Salisbury, iv.1974 (A. Watsham) (BMNH).
Paratypes. Rhodesia: 2 3, Salisbury, same data as holotype (BMNH). Uganda: 1 3g, 2 2, Kawanda,
vii—vili.1943, 3, viii.1942 (7. H. C. Taylor) (BMNH); 1 9, Bushenyi, iii.1939 (7. H. C. Taylor) (BMNH).
REMARKS. Related to the group of species with the apical depression of the scutellum appearing
almost quadrate. The male is distinguished from all others by the shape of the antennal segments.
Anacharoides rufa (Kieffer)
Coelonychia rufa Kieffer, 1912 : 451. Holotype, sex unknown, SouTH Arrica: Port Elizabeth (Brauns)
(originally in MNHU, Berlin, now lost).
The holotype of this species cannot be traced. According to information received from Dr
K6nigsmann in 1976 it could be in a collection on loan of which records have been lost.
Anacharoides sanitas sp. n.
(Figs 46, 62, 69, 88, 92)
DEscRIPTION. 2. Antenna 13-segmented, filiform, 3 shorter than 4+5, SN 3-13=AR 12:7:7:7:7:7:
6:6:6:6: 12, 4-8 equal in length, 9-10 equal, 11-12 equal, 13 twice length of 12. 3. Antenna with
segment 3 swollen on apical half, longer than segment 4, less than 4+-5 (Fig. 88), 4 and 5 swollen apically,
remaining segments moniliform, subequal in length. Head viewed frontally with eyes as far apart as the
height of an eye, frontal carina extending from outer margin of antenna to lateral ocellus, enclosing a
rugose area (pentagonal area), vertex incised, occiput striated, striae curved (Fig. 92). Pronotum viewed
dorsally angled on side front margins, mesoscutum with reticulate-rugose sculpture, anterior parallel
lines indicated, notaulices complete with weak crenulate appearance, median scutal line complete, apical
triangular area extending to middle of mesoscutum, lateral lines indicated. Scutellum viewed laterally
with pointed apex, reticulate-rugose, viewed dorsally with apical depression, scutellar foveae extending
past middle of scutellum, separated by aseptum, the apex of each fovea triangular, enclosing between them
a triangular area, apex of scutellum not arched, with a semicircular depression, lateral bars of scutellum
striate (Fig. 62). Mesopleurae smooth and polished, episternum with longitudinal striae, propodeum
with weakly bowed carinae, weakly pubescent either side of carinae. Segment 1 of gaster petiolate, longer
than broad, smooth, segment 2 of gaster liguliform, segment 3 the largest, closely punctate on apical half
dorsally and laterally. Hind tibia with longitudinal furrows and suberect spines on outer margins, with a
dense pubescent comb of hairs on inner margin and a row of widely placed spines on outer margin.
Colour: head red, pronotum red, mesoscutum reddish brown, gaster reddish on lateral margins, reddish
brown at apex, antennae and legs reddish. Forewings with short apical hair fringe, surface with dotted hair
bases, radial cell of forewing open on the margin and at base, Rs, not reaching margin of wing, cubitus
(M) not indicated.
MATERIAL EXAMINED
Holotype 2, South Africa: Cape Province, Mossel Bay, x.1921 (R. E. Turner) (BMNH).
Paratypes. Ethiopia: 1 2, Mt Zuguala, c. 9000 ft [2700 m], 22.x.1926 (H. Scott) (BMNH). Kenya:
1 2, Kiambu, 12.viii.1929 ex syrphid pupa (R. H. Le Pelley); 1 2, Mt Kenya, Kathita river, 9000 ft
[2700 m], 9.viii.1949 (J. A. Riley) (BMNH). Nigeria: 1 9, Zaria, Samaru, 22.ii1.1974 (J. L. Musa) (BMNH).
South Africa: 1 2, Cape Province, Somerset East, 10-22.xii.1930 (R. E. Turner) (BMNH); 9 3, same data
as holotype, varying dates (BMNH); | 3, Cape Province, Mossel Bay, 28.vi.1938 (R. E. Turner) (BMNH);
1 3, Capetown, x.1937 (R. E. Turner) (BMNH); | 2, Port St. Johns, Pondoland, 10—31.vii.1923 (R. E.
Turner) (BMNH). Uganda: 1 2, Mt Elgon, 6.viii.1934 (J. Ford) (BMNH). Yemen: 2 9, Usaifira, 1 mile N.
of Ta’izz, c. 4500 ft [1350 m], 12.xii.1937 (H. Scott & E. B. Britton) (BMNH); 1 3, Sana, c. 7900 ft
[2370 m], 19.1.1938 (H. Scott & E. B. Britton) (BMNH). Zaire: 1 2 Katanga, Dilolo, 24—27.vii.1931
(J. Ogilvie) (BMNH).
REMARKS. Resembles qguadrus but is clearly separated from it on a number of characters as
indicated in the key.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 103
Anacharoides striaticeps Cameron
(Figs 63, 81, 86, 87)
Anacharoides striaticeps Cameron, 1904a : 160. Holotype 2, SoUTH AFRICA (BMNH) [examined].
Coelonychia spinosipes Kieffer, 1910c : 19-20. Holotype 2, MALAwI: Nord-Nyassa, Langenburg (Fulle-
born) (originally in MNHU, Berlin, now lost ?). (Synonymy by Weld, 1930: 139.)
Anacharoides gibbosus Benoit, 1956 : 198. Holotype 2, ZAIRE (MRAC, Tervuren) [examined]. Syn. n.
DESCRIPTION. 2. Antenna 13-segmented, weakly clavate, 3 longer than 4, shorter than 4+5, 4 and 5
subequal, 7-12 decreasing gradually in length (Fig. 87). SN 1-13=AR 8:3:9:5:5:5:45:45:4:
4:4:4:7, the apical two segments darker than remainder, segments 4-13 with rhinaria, 13 as long as
11+12. 3. Antenna 14-segmented, segment 3 the longest, strongly excavated on outer margin, 4 strongly
swollen, narrow at base, 5-13 not swollen, SN 1-5=AR 5:3 :9-5 : 5-5 : 5-5. Head viewed frontally
with eyes as far apart as the height of an eye, frontal carina extending from outer margin of antenna to
lateral ocellus, enclosing a strongly reticulated area between median ocellusand antennal sockets. Occiput
strongly striate with a strong occipital carina. Pronotum with weak strigose sculpture, pubescent on sides,
angled apically with a small indistinct plate on front margin; no tooth medially. Mesoscutum reticulate-
coriaceous with fine evenly spaced white pubescence and transverse rugae, notaulices complete, but
broken by crenulations, anterior parallel lines indicated, median scutal furrow obsolete. Scutellum viewed
laterally with pointed apex, viewed dorsally with apical depression, scutellar fovea elongate, shallow,
extending past middle of scutellum, foveae separated by a septum, each fovea polished, carinae of
propodeum almost parallel; mesopleurae smooth and polished, suture indicated. Segment 1 of gaster
petiolate, three times as long as wide, smooth; segment 2 of gaster liguliform, segment 3 the largest
viewed laterally, closely punctate in lateral and dorsal views. Hind tibia with longitudinal furrows and
distinctive suberect spines on outer margins. Colour: 2, head, pronotum, scutellum, antenna and legs
reddish, mesoscutum blackish brown, gaster chestnut-brown-red; ¢, antenna, legs orange-yellow, head
and thorax black, gaster chestnut-brown-red. Forewings with only sparse hair fringe on lower apical
margin, surface of wing with scattered pubescence on apical half, middle with dotted hair bases, Rs,
extends along margin of wing for a short way without touching margin, vein R, not present, Sc+ R not
distinctly angled at the point where R, would normally emanate (Fig. 86).
MATERIAL EXAMINED
Anacharoides striaticeps Cameron, holotype °, South Africa: Pearston (Robert Brown) (BMNH) [right
antenna, left midleg, one fore and hind leg on right side, wings on left side missing]. Anacharoides
gibbosus Benoit, holotype °, Zaire: Rutshuruji, 1.1937 (J. Ghesquiere) (MRAC, Tervuren).
Angola: 1 2, 7 miles [11 km] W. Gabela, 16-18.iii.1972 (BMNH). Nigeria: 1 9, Ile-Ife, W. State, x.1973
(J. T. Medler). South Africa: 1 2, Cape Province, Somerset East, x.1930 (R. E. Turner) (BMNH); 2 8,
Port St Johns, Pondoland, 1-4.v.1923 (R. E. Turner) (BMNH). Uganda: 1 2, Budongo Forest, 7.ii.1935
(F. W. Edwards) (BMNH); 3 9, Kazhara, 111.1939 (7. H. Taylor) (BMNH); 1 °, ? Ruzumbra, c. 1939
(R. E. Turner) (BMNH). Zaire: 1 3 (allotype of gibbosus), same data as holotype except 15.vi.1937
(MRAC, Tervuren); 1 2 (paratype of gibbosus), same data as holotype (MRAC, Tervuren); 1 °, Massif
Ruwenzori, Kiurama, 2100 m, 26.x.1953 (P. Vanschuytbroeck & V. Hendricx) (MRAC, Tervuren);
1 9, Secteur Nord, Mulingo, 1300 m, Secteur Kikura, 9.vii.1954(P. Vanschuytbroeck & H. Synave)(MRAC
Tervuren).
Anacharoides stygius Benoit
(Figs 64, 65, 66)
Anacharoides stygius Benoit, 1956 : 197-198. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antenna 13-segmented, moniliform, weakly clavate, segment 3 1-5 times length of 4,
4 and 5 equal, 13 larger than 114+-12. SN 1-13=AR 5:3:11:5:65:6:6:5:5:4:4:4:4:9. Head
viewed frontally with eyes slightly further apart than height of an eye, frontal carina extending from outer
margin of antenna to lateral ocellus, enclosing a strongly reticulated area between median ocelli and
antennal sockets; occiput strongly striate with pronounced genal carinae. Pronotum with strong striate
strigose sculpture, angled apically with a small indistinct plate on front margin; mesoscutum with fine
reticulate sculpture and widely spaced transverse rugose sculpture, notaulices complete, distinct anterior
parallel lines indicated, median scutal furrow extending to middle of mesoscutum. Scutellum in lateral
view pointed at apex, viewed dorsally with apical depression, scutellar fovea extending past middle of
104 J. QUINLAN
scutellum, separated by a narrow septum with small margined triangular area in front of apical depression
(Fig. 64), sides of scutellum strongly reticulate. Mesopleurae smooth and polished, propodeum with
parallel carinae converging and joining at either end. Segment 1 of gaster petiolate, longer than wide,
smooth, segment 2 of gaster liguliform, segment 3 the largest, apical two-thirds dorsally and laterally
closely and finely punctate, punctures extending to remaining visible tergites. Hind tibia with longitudinal
furrows and widely spaced spines on outer margins (cf. Fig. 46). Colour: antenna, head and mesopleurae
reddish yellow, thorax blackish, gaster and legs chestnut-red-brown, hind coxa swollen, strongly cori-
aceous with a polished shiny area on inner margins. Forewing with ciliate surface on apical half and
scattered hair bases; hair fringe on apical margin short, vein Rs, thick, reaching wing margin, R, absent
Rs+M and M not indicated (Fig. 65), Sc+ R angled at point where R, would emanate.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Kiniata-Zobe, end xii.1915 (R. Mayne) (MRAC, Tervuren).
Madagascar: 1 9, Bekily, iv.1942 (A. Seyrig) (MRAC, Tervuren); 1 9, Bekily, vii.1940 (A. Seyrig)
(MNHN, Paris). Nigeria: 2 9, Ile-Ife, W. State, v. and x.1973 (J. T. Medler) (BMNH). Tanzania: 2 9,
Kilimanjaro Marangu, 13-20.vii.1957 (P. Basilewsky & N. Leleup) (MRAC, Tervuren). Zaire: P. N. A.
Secteur Tshiaberimu, Kirungu (lieu-dit) 2720 m, 28.ii1.1954 (P. Vanschuytbroeck & H. Synave) (MRAC,
Tervuren).
Anacharoides suspensus sp. n.
(Figs 61, 68, 79, 82, 83)
DESCRIPTION. ¢. Antenna 13-segmented, elongate moniliform, segment 3 longer than 4, shorter than
415, 4-12 gradually decreasing in length, apical segment equal in length to 3. SN 3-13=AR 12:9:8:
8:7:7:6:6:6:6:12. Head viewed frontally with frontal carina extending from outer margin of
antenna to lateral margin of eye, face densely pubescent, obscuring the sculptured surface, mandibles with
three teeth; vertex incised with frontal carina continued between lateral ocelli, eyes as far apart as height
of an eye. Pronotum with a small median dorsal tooth obliquely angled on side margins with a small
pronotal-type plate centrally. Mesoscutum with close transverse rugae and scattered pubescence, notau-
lices complete and crenulate, median scutal line distinct from posterior to middle, obscured anteriorly,
triangular at posterior (Fig. 61), anterior parallel lines obsolete. Scutellum viewed laterally with apex
pointed, viewed dorsally with apical depression, the basal margin straight, scutellar foveae not extending
past middle of scutellum, angled at apex to form a triangular area, lateral bars with fine broken striate
sculpture, apex of apical depression viewed dorsally, straight, posteriorly excavate. Mesopleura smooth
and polished, episternum pubescent, carinae of propodeum weakly bowed on inner margins. Segment | of
gaster petiolate, smooth dorsally, weakly furrowed laterally, segment 2 liguliform, segment 3 the largest,
closely punctate on lateral and dorsal surfaces, the visible part of segment 4 punctate. Hind tibia with
longitudinal furrows, closely pubescent on inner margin with long scattered spines on outer margin,
basitarsi as long as remaining tarsal segments. Hind coxae strongly swollen viewed laterally, with polished
excavation on inner margins, remainder of surface densely pubescent, inner margins with comb-like hair
fringe (Fig. 83). Colour: 2, antenna entirely orange-yellow, head orange-yellow, pronotum reddish orange,
mesonotum reddish suffused with black on outer margins, gaster chestnut-red, legs reddish orange;
3, same colour as 2, antenna 14-segmented, segment 3 distally expanded (Fig. 79), 4-14 moniliform.
Forewings dotted with short hair bases, apical hair fringe short, radial cell open on front and basally,
vein Rs not reaching margin, not extending along wing margin, veins Rs+M not indicated, vein R,;
absent, 2rm distinct.
MATERIAL EXAMINED
Holotype °, Senegal: Bambey, ex Ischiodon aegypticum (J. Risbec) (BMNH).
Paratypes. Nigeria: 1 2, Zaria, Samaru, 19.ix.1973 (B. Yashim) (BMNH). Senegal: 1 3, same data as
holotype (BMNH).
REMARKS. The male has similar antenna to the female of decellius but differs by the sculpture of
the mesoscutum. The female has affinities with those species bearing a quadrate scutellar depres-
sion.
PROSASPICERA Kieffer
Prosaspicera Kieffer, 1907 : 157. Type-species: Prosaspicera ensifera Kieffer, by subsequent designation
(Rohwer & Fagan, 1907).
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 105
DIAGNOsIS. 2 antenna 13-segmented, filiform, ¢ 14-segmented with a modified segment 3. Head viewed
frontally, coriaceous, inner margins of eye with a carina extending from lateral ocellus to the occiput,
vertex incised. Pronotum viewed dorsally raised in the form of a lip, mesoscutum reticulate to coriaceous,
median scutal line, anterior parallel lines and notaulices present; scutellar foveae distinct, apical spine
of scutellum long and sharply pointed at apex, in dorsal view extending back to base of radial cell.
Segment 1 of gaster short, wider than long, weakly furrowed, segment 2 of gaster liguliform, spherical
viewed laterally, segment 3 the largest, punctate, almost reaching apex of gaster. Hind tibia longitudin-
ally furrowed on inner margins. Forewings hyaline, usually bare, sometimes with apical hair fringe, radial
cell open on front margin, generally open at base.
DISTRIBUTION. Belize; Brazil; Chile; Formosa; Malawi; U.S.A.
DIscussION. The type-species of this genus is extra-limital and the generic description is based on
the original description by Kieffer, examination of a syntype (labelled paratype), a 2 No. 26017
in the United States National Museum, Washington, and on the Ethiopian species of Prosaspicera
discussed in this paper. I have been unable to see the syntypes housed at Pomona referred to by
Weld (1952). One other specimen labelled ‘Prosaspicera ensifera K. det. Weld 1934’ with the
locality ‘Mex 215H collection C. F. Baker’ has been examined.
Key to Ethiopian species of Prosaspicera Kieffer
Females (unknown in fropica)
1 Antenna weakly subclavate, segment 3 longer than 4 en 49, 100). Wing margin with complete
hair fringe . . 9)
— Antenna filiform (Figs 50, 94, 99), segment 3 at most as long as 4. Wing margin with hair fringe
either present or absent : ; 3
2 Notaulices very wide at apex with wide- spaced transverse carina on a reticulate base, median
scutal line forked near apex to forma polished horseshoe-shaped area, lateral bars of scutellum
reticulate, merging into lateral lines of scutellum. Scutellar foveae deep, very finely reticulate,
separated by a shallow septum. Base of scutellar spine with three carinae merging into a
striate medial area (Fig. 51). Radial cell of forewing with R, distinctly longer than wide, R,
curved, extended along wing margin (Fig. 41). Mesopleurae polished on upper half, mesopleural
suture obscured by strigose sculpture on lower half (Fig. 52).
Apical segment of antenna longer than segment11+12 . ’ . antennata (Benoit) (p. 107)
- Notaulices wide with closely spaced transverse carina, median scutal line forked near apex to
form a polished triangular area, lateral bars of scutellum reticulate at base, merging into
lateral margins of scutellum, scutellar foveae deep, reticulate-coriaceous, separated by a broad
septum. Base of scutellar spine broad with longitudinal carinae extending two-thirds of its
total length. Radial cell of forewing with R, short, not longer than wide, R, straight, not
projecting along wide margin os 53). ga rs sculptured on the lower half, coriaceous
on upper half. : : paragicida (Benoit) (p. 108)
3 Gena viewed laterally reticulate- -coriaceous, expanded (Fig. 54); upper facial area between the
frontal carinae reticulate-coriaceous (Fig. 54). Median scutal line dividing medially to form a
horseshoe-shaped area apically, notaulices not crenulate (Fig. 55). Scutellar foveae polished,
scutellar spine narrow basally (Fig. 55).
Forewing without apical hair fringe. Apical segment of antenna as long as segment 11+12
optivus sp. n. (p. 108)
— Gena viewed laterally not expanded. Median scutal line dividing medially to form an angular
shaped area apically (Fig. 47); notaulices complete, crenulate, foveae sculptured : age |
4 Frontal area between median ocellus and antennal sockets weakly strigose, becoming finely
reticulate-rugose, eyes viewed frontally strongly swollen closer together than height of an eye
measured at the median (Fig. 57), face with strong triangular depression covered in dense
white pubescence extending up between antennal sockets, vertex sharply incised
kisantua (Benoit) (p. 107)
- Frontal area between median ocellus and antennal sockets strongly strigose, eyes viewed
frontally not strongly swollen, as far apart as height of an eye (Fig. 56), face with weak
depression below median ocellus, weakly pubescent, vertex not sharply incised .
africana Kinsey (p. 106)
106 J. QUINLAN
Males (unknown in antennata and optivus)
This key should be treated with caution as only a single male is known of each species.
1 Median scutal line with triangular depression at apex dull and coriaceous. Scutellar fovea each
with a central longitudinal carina continued together with median carinae separating the
foveae down the scutellar spine. Upper face with strong radiating striae extending to antennal
scrobes (Fig. 58) 4 , . tropica (Kieffer) (p. 109)
— Median scutal line with depression at apex either polished and triangular or arcuate basally and
coriaceous. Scutellar foveae smooth and polished or coriaceous-reticulate, with a central
carina in each, not continued along scutellar spine. Upper face coriaceous, rugose or at most
with broken striae not extending to antennal scrobes : Z
2 Apical depression of median scutal line polished with aberrant sculpture. Outer orbits of face
with radiating striae, median area reticulate . ; 3
— Apical depression of median scutal line dull with fine sculpture. Outer orbits of face with faint
broken striae merging into coriaceous sculpture.
Scutellar foveae deep, surface reticulate-coriaceous, scutellar spine with three distinct carinae
extending two-thirds of way along the spine from apex of scutellar foveae. Spine completely
black ‘ . paragicida (Benoit) (p. 108)
3 Upper face strongly rugose. Median scutal line with apical depression arcuate basally, weakly
coriaceous on a shiny surface (Fig. 47). Scutellar foveae weakly striated on a polished reticulate
surface. Notaulices weakly crenulate. Scutellar spine completely black africana (Kinsey) (p. 106)
— Upper face with broken striae. Median scutal line with apical depression triangular, basally deep
and shiny, notaulices with strong crenulations. Scutellar spine chestnut-brown
kisantua (Benoit) (p. 107)
Prosaspicera africana (Kinsey) comb. n.
(Figs 47, 56, 94, 95, 101)
Aspicera africana Kinsey, 1919 : 162-163. LECTOTYPE 2, SouTH AFRICA (KSM, Kansas), here desig-
nated [examined].
DESCRIPTION. 2. Antenna filiform, segments 3-6 subequal in length, 7-12 gradually decreasing in length,
segment 13 longer than 12, rhinaria present on segments 3-13. SN 3-13=AR 32: 32: 31:31:25:25:
25: 25:25:25:52. $. Antenna 14-segmented, moniliform (9 segments only left on the paralectotype),
segment 3 excavate at base (Fig. 101). Head with face viewed frontally, coriaceous, eyes as far apart as
height of an eye, frons raised, inner margins of eye with a carina extending forward, head viewed dorsally
with distinct carina extending from lateral ocellus to antenna and beyond, upper face irregularly rugose,
pubescent, cheeks pubescent, mandibles dark orange-brown, occipital carina distinct, occiput striated;
vertex with angular depression (Fig. 95). Sides of pronotum pubescent on a coriaceous surface, front of
pronotum viewed dorsally raised in form of a lip, mesoscutum reticulate-coriaceous, median scutal line
visible anteriorly in the form of an arcuate depression at posterior, anterior parallel lines raised, extending
almost half way to scutellar foveae, notaulices complete, weakly crenulate, wide posteriorly, narrowing
anteriorly. Lateral bars of scutellum reticulate-coriaceous, extending past the scutellar fovea, scutellar
disc truncate either side of scutellar spine and with hair tufts. Viewed dorsally, scutellar foveae polished,
deep with striations, foveae separated by a septum which continues along the scutellar spine, spine of
scutellum wide-based with striated dorsal surface, narrowing to a sharp spine (Fig. 47). Mesopleurae
polished with a suture and weak strigose sculpture, propodeum and metapleurae pubescent. Segment | of
gaster short, wider than long viewed laterally, weakly furrowed, segment 2 of gaster liguliform, spherical
viewed laterally, segment 3 the longest, almost reaching apex of gaster, punctate on apical two-thirds,
remaining segments short, punctate. Hind tibia longitudinally furrowed. Colour: antenna, legs reddish
brown, head, thorax and gaster brownish black. Forewing with short apical hair fringe in 9, present on
upper margin only in 3, apex of wing with sparse scattered hairs, remainder with widely spaced spicules,
radial cell distinctly open on front and apex (Rs, and R, not reaching wing margin), areolet absent.
MATERIAL EXAMINED
Lectotype 2, Rhodesia: Salisbury, 5050 ft [1515 m], vi.1900 (F. L. Snow) (KSM, Kansas).
Rhodesia: 4 2, Chishawasha, nr Salisbury, ii.1978 (A. Watsham) (BMNH): 1 9, Salisbury, 1969 (A.
Watsham) (BMNH); 1 3 (paralectotype), Salisbury, 5050 ft [1515 m], xii.1900 (F. L. Snow) (KSN, Kansas).
REMARKS. A second male paralectotype in the Kinsey collection (same data as lectotype except
xii. 1900) was not examined.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 107
Prosaspicera antennata (Benoit) comb. n.
(Figs 41, 43, 49, 51, 52, 96)
Aspicera antennata Benoit, 1956 : 202. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antenna 13-segmented, subclavate, segment 3 longer than 4, 4 and 5 equal, 6 as long as 3,
7 as long as 4, 8 as long as 4, 9, 10, 11 and 12shorter than 8, 13 longer than 11+ 12 (Fig. 49), SN 3-13 =
AR 14:13: 13:14:13:13:11:11:10: 10:22, 3-13 with rhinaria. Head viewed dorsally with carina
extending from outer margin of antenna to lateral ocellus enclosing a fine reticulate area, occipital carinae
pronounced, vertex incised, back of head below ocelli with canaliculate sculpture (Fig. 96); mesoscutum
with distinct notaulices, wider at apex and crenulate, anterior parallel lines indicated, median scutal line
extending to middle of mesoscutum where it widens and joins the scutellum at the scutellar suture, surface
finely reticulate. Scutellar spine viewed laterally in same plane as scutellar foveae, extending almost to
basalis (Fig. 51). Mesopleura polished medially with lateral striations on lower half; lateral bars of
scutellum reticulate at base, becoming striated on side margins, scutellar foveae very finely coriaceous,
separated by a narrow septum, surface of disc finely reticulate, base of spine broad, gradually tapering
towards apex (Fig. 51), base either side of spine with tufts of pubescence. Mesopleura polished on upper
half, strongly strigose on lower half (Fig. 52). Segment 1 of gaster viewed laterally, wider than long,
crenulate, segment 2 of gaster liguliform, segment 3 the largest, the apical half dorsally and laterally with
dense punctures. Hind tibia with longitudinal furrows. Colour: 2 antenna with segments | and 2 chestnut-
brown (ferruginous), 3-13 orange-yellow, head and thorax black, gaster dorsally and upper lateral
margins chestnut-brown, apical margins yellowish, legs orange-yellow. Forewings with a colourless apical
fringe, surface of wing with spicules except for a few sparse hairs apically, radial cell open at base andapex
(Rs, and R, not reaching wing margin) (Fig. 41), R, short, longer than wide, weakly projecting (Fig. 41),
areolet absent, 2rm in form of a spur, cubitus (/) absent, hair fringe on apical margin short.
3 unknown.
MATERIAL EXAMINED
Holotype 2°, Zaire: Basoko, xii. 1948 (P. L. Benoit) (MRAC, Tervuren).
Prosaspicera kisantua (Benoit) comb. n.
(Figs 42, 48, 57, 98, 99)
Aspicera kisantua Benoit, 1956 : 203-204. Holotype 2, ZAIRE (MRAC Tervuren) [examined].
DESCRIPTION. °. Antenna 13-segmented, filiform, segments 3—6 equal in length, 7-9 equal in length, each
shorter than 3, 10 and 11 each shorter than 9, 12 shorter than 11, 13 longer than 11+12. Sn 3-13=
AR 17:17:17:17:15:15:15:14: 14:13:28, rhinaria present on segments 3-13. 3 antenna 14-
segmented, segment 3 expanded apically, 4-13 moniliform, subequal in length, apical segment longer than
penultimate. Head viewed dorsally with distinct frontal carina extending from outer margin of lateral
ocellus to antenna, enclosing a finely reticulate area, eyes further apart than height of an eye, occipital
carinae distinct, vertex incised, back of head with wide longitudinal furrows (Fig.98).Pronotum viewed
dorsally densely pubescent, angled on side margins with a small protruding pronotal plate; surface of
mesoscutum with reticulate-coriaceous sculpture, notaulices complete with strong crenulate sculpture,
median scutal line extending from base, diverging at a point midway to enclose a polished area, anterior
parallel lines not extending to halfway down mesonotum, lateral bars of scutellum reticulate at base,
weakly striate apically. Scutellar foveae shallow with wide-spaced crenulations separated by a septum,
scutellar disc with longitudinally strigose sculpture interspaced with reticulations at apex; scutellar spine
wide at base, rugose, extending as far as apical margin of tergite two viewed dorsally. Mesopleura smooth
and polished anteriorly, with weak striations and coriaceous sculpture basally; sides of propodeum densely
pubescent. Segment | of gaster wider than long, obscured by tergite 2, tergite 2 liguliform, tergite 3 the
largest, strongly punctate on lateral and dorsal surfaces of apical two-thirds. Hind tibia with longitudinal
furrows. Colour: antenna and legs reddish yellow, head and thorax black, gaster black with dark chestnut
tinge at base and on ventral margins. Forewings with hair fringe present on upper apical margin only,
surface of wing dotted with widely spaced spicules, radial cell open on front margin (Rs, and R, not
reaching wing margin), areolet absent, R, short and weakly projecting, 2rm in form of a spur, cubitus (M)
absent (Fig. 42).
108 J. QUINLAN
MATERIAL EXAMINED
Holotype °, Zaire: Bas-Congo, Kissantu, 1931 (R. P. Vanderyst) (MRAC, Tervuren).
Ghana: | 2, ex syrphid on Gusicidia (BMNH). Nigeria: 1 °, Ile-Ife, 2.viii.1969 (J. T. Medler) (BMNH).
Sierra Leone: | 3, 1 2, Njala, 1931 ex syrphid on groundnuts (E. Hargreaves) (BMNH).
Prosaspicera optivus sp. n.
(Figs 50, 54, 55, 97)
DEscRIPTION. 2. Antenna 13-segmented, filiform, 3 shorter than 4, 4 and 5 subequal, 6-12 each decreasing
very slightly in length, 13 longer than 12. SN 3-13=AR 17: 19: 19: 16: 14: 13-5: 12-5: 11:10:11: 22,
rhinaria present on segments 3-13. Head viewed dorsally with distinct frontal carina extending from outer
margin of lateral ocellus to antenna, viewed laterally with reticulate-coriaceous sculpture between
occipital carina and margin of eye, genal area wide (Fig. 97), vertex incised, occipital carinae viewed dorso-
posteriorly, expanded, vertex with canaliculate sculpture (Fig. 97). Pronotum weakly pubescent on side
margins with a small protruding pronotal plate. Surface of mesoscutum with reticulate sculpture, median
scutal line dividing medially to form a horseshoe-shaped area, anterior parallel lines short, not extending
to middle of mesoscutum, notaulices complete, smooth and polished, lateral bars of scutellum reticulate,
merging into margin of the scutellar disc. Scutellar foveae smooth and shiny, with a septum continued
along the dorsal surface of the acutely pointed spine, apex of disc truncate, spine at base arising from
above the apex of scutellar disc (Fig. 55). Mesopleurae smooth and polished anteriorly, strigose in part
basally; sides of propodeum pubescent. Segment 1 of gaster wider than long, obscured by tergite 2,
tergite 2 liguliform, tergite 3 the largest viewed laterally, the apical third strongly punctate, remaining
segments hardly visible. Hind tibia with longitudinal furrows on inner margins. Colour: antenna and
legs reddish yellow, head, mesoscutum, scutellum and apical segments of gaster black, anterior half of
mesopleurae and basal segments of gaster reddish. Forewings without apical hair fringe, surface of wing
dotted with widely spaced spicules, radial cell open on front margin (Rs, reaching wing margin, R, not
reaching wing margin), areolet absent, 2rm not in form of a spur.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Congo: Cayo Dist., Central Farm, x.1.1975, on Zea mays L. (R. Akers) (BMNH).
REMARKS. This species is distinguished from ethets with a filiform antenna by the very distinctive
. expanded gena. — riche
Prosaspicera paragicida (Benoit) comb. n.
(Figs 53, 100, 102)
Aspicera paragicida Benoit, 1956 : 202-203. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antenna 13-segmented, weaklyclavate, segment 3 longer than4, 4and Sequal, SN 1-13=
AR 20:6: 16:13:15: 15:13:13: 13:13:13:13: 26, segments 9-12 almost as broad as long (Fig.
100), rhinaria present on 3-13. 3. Antenna 14-segmented. SN 3-5=AR 20: 16: 17. Segment 3 strongly
expanded apically (Fig. 102), rhinaria on segments 3-14. Head viewed dorsally with frontal carina extend-
ing from outer margin of antenna to lateral ocellus, not distinct, enclosing a fine reticulate area with
radiating sculpture in part, occipital carina distinct, occiput with widely spaced canaliculate sculpture
(cf. Fig. 97). Pronotum viewed dorsally, angled on the margins. Mesoscutum with fine reticulate sculp-
tured surface, notaulices distinct, complete, wide at apex and crenulate, lateral lines complete, anterior
parallel lines not reaching halfway to apex, median scutal line reaching halfway to apex and branching
to enclose a polished wide triangular area. Mesopleurae polished on upper half with fine striate sculpture
and a carina (a mesopleural suture) in middle section. Scutellar spine viewed laterally in the same plane
as scutellar foveae, strongly ridged; scutellum finely reticulate with longitudinal strigose sculpture
continued along spine, scutellar fovea deep, surface reticulate-coriaceous, separated by a weak septum,
the apex of scutellum either side of spine pubescent. Segment | of gaster viewed laterally wider than long,
crenulate, segment 2 of gaster liguliform, segment 3 the largest, the apical two-thirds with dense punctures
dorsally and laterally. Hind tibia with longitudinal furrows. Colour: antenna, legs reddish yellow, head,
thorax and gaster black, apex of gaster with dark chestnut-brown tinge. Forewing with radial cell open
at base and apex (Rs, and R, not reaching margin of wing), R, short, conical, as wide as long measured at
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 109
base, weakly projecting, cubitus (M) absent, Rs, almost straight, venation very pale, surface of wing
dotted with spicules on apical half, hair fringe on apical margin short.
MATERIAL EXAMINED
Holotype 2°, Zaire: Samkuru, Gandajika, 5.iii.1948 (P. Henrard) (MRAC, Tervuren).
Zaire: 2 2, 2 3, same data as holotype (all parasites of larvae of Paragus borbonicus Macquart (Syr-
phidae) (MRAC, Tervuren).
Prosaspicera tropica (Kieffer)
(Fig. 58)
Aspicera tropica Kieffer, 1910c : 20. Holotype 3, MALAWI (MNHU, Berlin) [examined].
Prosaspicera tropica (Kieffer) Weld, 1952 : 167.
DESCRIPTION. 3. Antenna 14-segmented. SN 3-5=AR 14:13:12. Segment 3 swollen medially, 4-13
moniliform, subequal in length, 14 longer than 13. Head viewed frontally arcuate on vertex, inner orbits
of face with radiating striae (Fig. 58), face above clypeus densely pubescent, occipital carinae distinct,
vertex crenulate (Fig. 58). Pronotum angled on dorsal margin with a coriaceous pronotal plate, sides of
pronotum coriaceous and slightly pubescent. Surface of mesocutum coriaceous, median scutal line distinct,
in form of a wide deep coriaceous triangular area apically, its side margins bordered by the inner margins
of the notaulices, notaulices wide at apex, crenulate, anterior parallel lines pronounced, extending to a
point midway on the mesoscutum, lateral lines almost complete; scutellar foveae separated by carinae,
each fovea with a longitudinal carina extending to and along the scutellar spine, scutellar spine viewed
dorsally reaching as far as margin of segment 3 of gaster viewed dorsally; propodeum carina not obvious.
Mesopleural suture distinct, area above and below suture reticulate-coriaceous. Segment | of gaster not
visible. Segment 2 liguliform, segment 3 the largest, viewed laterally the apical two-thirds strongly
punctured, remaining segments not visible. Hind tibia with longitudinal ridges. Radial cell of forewing
open at base and on wing margin, vein m indicated as a short spur. Colour: head and thorax black, gaster
chestnut-brown to black, antenna and legs orange-yellow.
2 unknown.
MATERIAL EXAMINED
Holotype 3, Malawi: Nyassa-See, Langenburg, vi—vil. 1898 (S. Fiil/lehorn) (MNHU, Berlin).
OBERTHUERELLINAE Kieffer
Oberthuerellinae Kieffer, 1903 : 88. Type-genus: Oberthuerella Saussure.
Segment 5 of gaster the largest, preceded by three short segments (Fig. 2). Scutellum more usually with
a spine on the apical margin. Hind femur with a distinct tooth on the underside (Fig. 11).
Discussion. The Oberthuerellinae was treated as one of three subfamilies in the Liopteridae by
Hedicke & Kerrich (1940) and again by Weld (1952). It is closely related to the Mesocynipinae
but distinguished from it by a conspicuous tooth or spine on the underside of the hind femur
and the scutellum invariably has a distinct medial spine at the apex with a tooth or spine on either
side. It is separated from the Liopterinae by the gaster being attached normally at an angle
(Fig. 2). The Oberthuerellinae are not known to occur outside the Ethiopian Region and comprise
three genera, Oberthuerella Saussure, Tessmannella Hedicke and the monotypic Xenocynips Kieffer.
Twelve species of Oberthuerella, including two newly described here, are at present recognised. Of
the twelve, two are omitted from the key to species either because the syntype-material is lost or
the type-depository is not known. Only four species of Tessmannella are known, including one
described in this paper. Hedicke (1912a) separated Tessmannella and Xenocynips from Ober-
thuerella on the median dorsal tooth of the pronotum being present in both but completely absent
in Oberthuerella. Xenocynips was erected by Kieffer (1910) and is separated from Oberthuerella
and Tessmannella by segments 2-4 of the gaster being fused.
110 J. QUINLAN
Key to genera of Oberthuerellinae
1 Pronotum without a median dorsal tooth on hind margin; propodeum with pronounced side
margins in the form of flaps, strongly sculptured (Fig. 117); tooth on hind femur almost erect
(Fig. 11); segment 1 of gaster sulcate, at most twice as long as broad, almost rectangular
(Fig. 114) . : : OBERTHUERELLA Saussure (p. 110)
— Pronotum viewed laterally with a median dorsal tooth on hind margin anterior of mesoscutum
(Fig. 116), propodeum without pronounced side margins (Fig. 115); angle of tooth on hind
femur variable; segment | of gaster variable in length 2
2 Tergites 2, 3 and 4 short, 5 the largest; hind femur with a rounded lobe between medial area
and apex, tooth angled, not erect (Fig. 109); segment | of gaster at least three times as long
as broad, petiolate (Fig. 120); hind tibia with distinct lobe opposite the two tibial spines
apically; scutellar foveae three in number : : . TESSMANNELLA Hedicke (p. 116)
~ Tergites 2-4 fused without trace of suture (Fig. 32); hind femur without a rounded lobe at apex,
tooth between medial area and apex erect (Fig. 111); segment 1 of gaster sulcate, less than
three times as long as broad, almost rectangular; hind tibia without a distinct lobe apically;
scutellar foveae obscure (Fig. 110) ; ; ; , . XENOCYNIPS Kieffer (p. 118)
OBERTHUERELLA Saussure
Oberthuerella Saussure, 1890 : 20, pl. 20. Type-species: Oberthuerella lenticularis Saussure, by monotypy.
DIAGNOSIS. 2 antenna 13-segmented, clavate; ¢ antenna 14-segmented. Face viewed frontally reticulate
to rugose with a frontal line or keel extending from anterior ocellus to clypeus in most species, eyes
further apart than the height of an eye viewed frontally. Pronotum sculptured without a median tooth
on hind margin. Mesoscutum variously sculptured, scutellum apically tridentate the median tooth
generally in the form of a long spine except in friformis, base of scutellum with a variable number of
between 2 and 4 foveae each separated by a longitudinal carina. Segment |! of gaster (petiole) sulcate,
variable in length, segments 2—4 each short, 5 the largest in lateral view. Hind femur with erect tooth.
DISTRIBUTION. Cameroun; Madagascar; Nigeria; Zaire.
DIscussiON. The original publication contained only the name and a figure. Kieffer (1903 : 88),
however, gave an extended description of /enticularis. Weld (1952) stated ‘that a female in Paris
collected by Alluaud in 1901 in Madagascar may be the type of /enticularis Sauss’. To this
specimen Weld added his determination label ‘Oberthurella lenticularis Sauss. det. Weld 1931”.
This specimen cannot be a syntype because Saussure’s original figure of /enticularis was published
in 1890. The specimen determined by Weld agrees in detail with Kieffer’s description and with
the figure of /enticu/aris illustrated by Saussure (1890). I have examined a specimen standing
under /enticu/aris in Geneva labelled ‘Typus’ and ‘Madasgasc’; it is a male and therefore cannot
be the type figured by Saussure (1890) or the specimen described as /enticu/aris by Kieffer (1903)
Key to species of Oberthuerella Saussure
1 Frontal line in form of a keel or crest running from median ocellus to clypeus through rugose
sculpture; gaster viewed laterally generally with conspicuous hairs on segments 2—5 viewed
dorsally with or without long golden pubescence; 2 with weak apical hair fringe on segments
5 and 6 viewed laterally : 2
— Frontal line not present, face entirely reticulate- -rugose; gaster viewed dorsally without con-
spicuous hairs on the segments; 2 with sparse hairs and apical fringe sometimes present on
segments 5—7 - ; 8
2 Tergites 2-5 of gaster viewed dorsally with dense long golden pubescence. Segment I of gaster at
least as broad as long (Fig. 119); areolet absent.(¢ unknown) aureopilosa Benoit (p. 112)
— Tergites 2-3 of gaster viewed dorsally without pubescence, 4-5 with fine pubescence or bare
(Fig. 120); areolet present or absent : 3
3 Apical margin of tergite 4 of gaster almost straight medially (Fig. 120): segment 1 of gaster
longer than broad.
Areolet absent ? 5
— Apical margin of tergite 4 of gaster strongly arcuate medially (Figs £21, 127), segment 1 of gaster
either longer than broad or as broad as long . ; : ; ‘ : : : . ae
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 111
4 Segment 5 of gaster viewed dorsally with long scattered pubescence; first cubital cell of forewing
closed, areolet distinct though sometimes weakly indicated (Fig. 122) ‘ : 6
— Segment 5 of gaster viewed dorsally without long scattered pubescence; first cubital of forewing
open, areolet absent (Fig. 123) . : 7
5 Pronotum distinctly striated; scutellar spine short, viewed dorsally not extending o1 one- -fourth of
the length of segment | of gaster, segment 5 of gaster viewed laterally shiny with fine scattered
punctures, segments 5 and 6 with apical hair fringe; fore and mid legs black, hind coxae dark
chestnut, femur and tibia yellow, tarsi reddish black. (¢ unknown) breviscutellaris Benoit (p. 112)
— Pronotum reticulate-rugose; scutellar spine long, viewed dorsally extending complete length of
segment | of gaster in male, short in female; segment 5 of gaster viewed laterally closely and
densely punctate, dull with long scattered hairs on dorsal surface (Fig. 126); fore, mid and
hind legs reddish yellow “ 5 longispinosa Benoit (p. 114)
6 Pronotum reticulate-rugose; gaster and legs orange- -yellow, segments S5—7 of gaster with fine
scattered pubescence on upper lateral surface, lower lateral surface very finely punctate;
forewing with areolet very weakly indicated, first cubital cell appearing closed basally (Fig. 122).
($ unknown) é ; crassicornis Benoit (p. 112)
— Pronotum punctate- -reticulate (the punctures widely spaced); gaster and legs chestnut-brown,
segments 5-7 of gaster with long golden scattered pubescence on upper lateral surface,
lower lateral surface strongly punctate; forewing with areolet distinct, first cubital cell
closed (Fig. 124).(¢ unknown) ; . nigrescens Benoit (p. 115)
7 Frontal line in form of a sharp keel, narrow; segment 3 of antenna shorter than 4 in both sexes;
segment | of gaster longer than broad, black . ‘ : abscinda sp. n. (p. 111)
— Frontal line not sharp, raised and broadened apically, obscured by sculpture; segments 3 and 4
of antenna subequal; segment | of gaster as broad as long, chestnut-brown. (¢ unknown)
longicaudata Benoit (p. 114)
8 Forewing with a distinct areolet, first cubital cell closed (Fig. 128); scutellar spine long, viewed
dorsally extending well past middle of segment 1 of gaster; pronotum and mesoscutum
coarsely reticulate rugose; legs entirely yellow . ; lenticularis Saussure (p. 113)
— Forewing without an areolet, first cubital cell closed; scutellum tridentate but without a spine
medially (Fig. 125); pronotum and mesoscutum aciculate; fore and mid legs reddish black, hind
femur and tibia reddish yellow. (¢ unknown) . ; ; ; : triformis sp. n. (p. 115)
Oberthuerella abscinda sp. n.
DESCRIPTION. 2. Antenna 13-segmented, clavate, segment 3 shorter than 4. SN 3-13=AR 17:23:18:
20:15:17:14:14:12: 12:20, segments 5-13 each progressively thicker than preceding segment.
g. Antenna 14-segmented, segment 4 twice the length of 3, 5 longer than 3, shorter than 4, remainder
subequal to 5. Face viewed frontally reticulate-rugose with a distinct frontal line or keel extending from
anterior ocellus to clypeus, eyes further apart than height of an eye, cheeks converging sharply. Pronotum
and mesoscutum reticulate-rugose viewed dorsally. Notaulices submerged in coarse reticulate sculpture.
Scutellum with 3-4 irregular foveae at base, apical margin tridentate with the median tooth in form of a
distinct spine. Propodeum viewed posteriorly, angular, carina of propodeum weakly bowed with trans-
verse canaliculate sculpture either side. Mesopleura with strigose sculpture on a polished surface. Segment
1 of gaster sulcate, longer than broad, gaster viewed dorsally without pubescence, segment 4 of gaster
arcuate medially; hypopygium protruding, ventral spine short. Legs black, hind femur with erect spine
between middle and apex, punctate; hind tibia foveolate-reticulate on outer margins, inner margins with
longitudinal carinae. Wings infuscate, densely ciliate on the surface, margins bare, radial cell of forewing
closed, first cubital cell open, second cubital cell absent (areolet), cubitus almost reaching apex of wing.
Colour: antenna, head, segment | of gaster (petiole) and legs black; gaster yellow.
MATERIAL EXAMINED
Holotype 2, Zambia: Mbala (‘Abercorn’), 31.xii.1943 (BMNH).
Paratypes. Zambia: | 3, same data as holotype (BMNH). Zaire: Eala, xii.1932 (A. Corbiser) (MRAC,
Tervuren).
REMARKS. This species is in the group with a frontal line present on the face but lacks long
scattered pubescence on the gaster. Segment 3 of the antenna is shorter than 4 in both sexes, thus
separating it from /ongicaudata.
112 J. QUINLAN
Oberthuerella aureopilosa Benoit
(Fig. 119)
Oberthuerella aureopilosa Benoit, 1955 : 290. Holotype 9, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. °. Antennae |3-segmented (only basal three segments of type present, five median segments
mounted on card point). Face viewed frontally with a pronounced median carina, extending to median
ocellus, surface reticulate-rugose with fine pubescence, cheeks not sharply converging. Pronotum and
mesoscutum with broken transverse reticulate-rugose sculpture, median scutal line complete and distinct,
notaulices distinct. Scutellum coarsely reticulate-rugose with long apical spine medially, lateral bars
extended at apex to form short spines, scutellar foveae canaliculate. Propodeum angular viewed from
behind, sculpture obscured by dense pubescence; mesopleura with strigose sculpture on upper half,
polished on lower half. Segment 1 of gaster viewed dorsally slightly broader than long, crenulate, gastral
segments 2-5 on dorsal surface with long golden pubescence on a foveolate-reticulate surface, viewed
laterally apical half of segment 5 pubescent, remaining visible segment pubescent, hypopygium not
protruding, apical sternite produced, apical margin with long setae, ventral spine short. Legs dark
chestnut-red, tarsi black, hind femur with perpendicular spine between middle and apex, hind tibia
polished, with widely spaced foveolae, inner side with a longitudinal carinae. Wings infuscate, densely
ciliate on surface, margins bare, radial cell of forewing closed, first cubital cell closed, 2nd cubital (areolet)
not indicated, cubitus almost reaching apex of wing. Colour: antenna, head, thorax, segment | of gaster
with black and brown crenulations, gaster orange-red.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Maniema, Kindu, xi.1973 (L. Burgeon) (MRAC, Tervuren).
Oberthuerella breviscutellaris Benoit
(Fig. 120)
Oberthuerella breviscutellaris Benoit, 1955 : 286. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antennae 13-segmented, subclavate, segment 3 shorter than 4, 4 shorter than 5, 6 shorter
than 5, subequal to 7, 8-13 subequal in length, 8-12 as wide as long, 13 longer than wide. SN 3-13=
AR 10:12:15:12:11:10:8:8:8:8:14. Face viewed frontally with coarse reticulate sculpture,
striate in region of clypeus and cheeks, median area in form of a crest viewed laterally, eyes further apart
than height of an eye, cheeks converging sharply. Pronotum transversely striate dorsally. Mesoscutum
with coarse transverse ridge-like sculpture, notaulices crenulated, impressed, median scutal line absent.
Scutellum with foveae at base (obscured in the holotype by the pin on which it is mounted being through
the centre), sculpture large reticulate-rugose, apex of scutellum with medial spine spoon-shaped and
lateral bars tooth-shaped apically. Propodeum viewed dorsally, foveolate. Mesopleura with diagonal
striate sculpture, mesopleural triangle finely coriaceous. Segment 1 of gaster sulcate, longer than broad
viewed dorsally (length 3-5, breadth 2-5) but broader apically than at base (Fig. 120). Tergites 2-4 short,
tergite 4 of gaster as long as 3+ 4 (Fig. 120), 5 the longest, all segments with sparse punctures visible on
the dorso-lateral margins, apex of 5, 6 and 7 with long setae, hypopygium distinct. Coxae and tibiae
smooth and shining, tarsi dull, sculptured and pubescent, hind metatarsus longer than remaining seg-
ments combined, hind femur with distinct perpendicular tooth or spine. Wings entirely fuscous, forewing
with radial cell completely closed, radial and first cubital cell darker, first cubital open, second cubital
(areolet) not indicated, wing surface lightly ciliate, margins bare, cubitus almost reaching apex of wing.
Colour: antenna, head, thorax, fore and mid legs black, hind coxae black, femur, tibiae and tarsi red,
segment | of gaster black, remaining segments yellowish orange.
MATERIAL EXAMINED
Holotype 2, Zaire: Lalua, Luashi, x.1938 (F. Freyne) (MRAC, Tervuren).
Rhodesia: | 2, Mt Chirinda, Gasaland (Marshall) (BMNH). Zaire: 1 2, Lubumbashi, 1.iii.1975 (W.
Beun) (MRAC, Tervuren).
Oberthuerella crassicornis Benoit
(Figs 121, 122, 127)
Oberthuerella crassicornis Benoit, 1955 : 289. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
Oberthuerella compressa Benoit, 1955 : 292. Holotype 2, ZAIRE (MRAC, Tervuren) [examined]. Syn. n.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 113
DESCRIPTION. ¢. Antenna 13-segmented, clavate. SN 3-13=AR 12:15:12:12:12:10:10:10:10:
10 : 23, 5-12 each progressively thicker. Face viewed frontally reticulate-rugose, with a distinct frontal
line or keel extending from median ocellus to clypeus, eyes further apart than height of an eye, strongly
swollen in frontal view, cheeks converging, pronotum and mesoscutum foveolate to reticulate-rugose
viewed dorsally, notaulices present, impressed in the sculpture, base of scutellum with three large foveae,
median area reticulate-rugose, apex of scutellum tridentate, the medial spine viewed dorsally as long as
segment | of gaster (petiole). Mesopleurae transversely strigose to rugose-foveolate anteriorly. Propodeum
with parallel carinae and a transverse medial ridge. Segment 1 of gaster sulcate, viewed dorsally longer
than wide, segment 2 of gaster viewed dorsally shorter than 1, 3 subequal to 2 (Fig. 127); apical margins
straight medially, segment 4 longer measured medially than 2+ 3, apical margin arcuate, segment 5 the
largest with scattered pubescence and strongly punctate in lateral view, apical margin of tergite 6 with a
fringe of hairs, hypopygium short, not protruding. Legs orange-yellow, smooth and shining. Hind femur
with suberect median spine; hind tibia weakly sculptured. Wing surface ciliate, margins bare, radial cell
of forewing completely closed, cubitus extending almost to edge of wing, first cubital cell closed, areolet
weakly indicated. Colour: antenna, head and thorax black, petiole black and red, gaster orange-red.
3 unknown.
MATERIAL EXAMINED
Oberthuerella crassicornis Benoit, holotype 2, Zaire: Tshuapa, Eala, vii. 1936 (J. Ghesquiére) (MRAC,
Tervuren). Oberthuerella compressa Benoit, holotype 2, Zaire: dist. Lac Léopold II, Bena Bendi, v.1915
(R. Mayne) (MRAC, Tervuren).
Malawi: | °, Malanje, 24.iv.13 (S. A. Neave) (BMNH).
Oberthuerella lenticularis Saussure
(Figs 114, 117)
Oberthuerella lenticularis Saussure, 1890: pl. 20. Holotype 2, MADAGASCAR: (? MHN, Geneva). (Holo-
type not found; description based on material determined by Weld, 1931.).
DEscRIPTION. ?. Antenna 13-segmented, subclavate, segment 3 shorter than 4, 5-12 cylindrical, longer
than: broad, 13. subequal: .to- 11-12; -SN' 3=!13=AR117-2.21 $222.20 +182 1512442 122 122 20 s.
Antenna 14-segmented. SN 3-14=AR 10:17:15:15:15:13:13:13:13:13:13:20. Head viewed
frontally with a median ridge extending from median ocellus to a point midway between antennal sockets,
face and dorsal surface of head coarsely reticulate-rugose with scattered pubescence, face without a
medial carina. Pronotum and mesoscutum shining, coarsely reticulate, median scutal line and notaulices
obscure, impressed in the sculpture, centre of scutellum dished with longitudinal carina within each of
two foveae, apex of scutellum reticulate-rugose, ending in a smooth spine almost as long as breadth of
scutellum, lateral bars extending at apex to form short spines. Propodeum angled either side of centre
which is broad and crenulate (Fig. 117). Mesopleurae with strigose-reticulate sculpture on upper half,
polished on lower half. Segment 1 of gaster viewed laterally as broad as long, crenulate (Fig. 114), segment
2 viewed dorsally short, 3 twice as long as 2, 4 as long as wide, 5 the longest, viewed laterally apex of
tergite 4 with scattered setae, apex of 5 and 6 finely reticulate, hypopygium not protruding, ventral spine
short. Petiole in ¢ longer than broad, segments 2 and 3 of gaster viewed dorsally of equal length, 4 twice
length of 3. Legs orange-yellow, coxae and femur shiny, hind femur with distinct perpendicular tooth.
Wings infuscate, darker basally, surface finely ciliate, margins bare, radial cell of forewing completely
closed, first cubital cell closed, second cubital (areolet) weakly indicated, cubitus almost reaching apical
margin of wing (Fig. 128). Colour: antenna, head and thorax black, petiole and gaster orange-red.
MATERIAL EXAMINED
Madagascar: | 2, Region du Sudest, Vallée du Fanjahira Isakam, xii.1901 (Ch. Alluaud) (MNHN, Paris);
1 2,13, 1919 (J. de Gaulle) (MNHN, Paris); 1 2, ? Somlirano (A. Seyrig)(MNHN, Paris); 2 2, Fanovano,
ili.34 (A. Seyrig) (MNHN, Paris); 1 2, Manara, x.1963 (J. Vadon) (MRAC, Tervuren); 1 9°, Manguzi
River, nr Maputa Z., xii.1945 (H. Bell Marley) (BMNH); 1 2, Ambodivoangy, xii.1961 (J. Vadon) (MRAC
Tervuren); 1 9, Fanpanambo, ii.1960 (J. Vadon) (MRAC, Tervuren); 1 9, same data except ii.1961
(MRAC, Tervuren); 1 2, Manara, 1963 (J. Vadon) (MRAC, Tervuren); °, Bekily, Reg. Sud De Lile,
iii. 1933 (A. Seyrig) (MNHN, Paris). Malawi (Nyasaland): Chiromo (R. C. Wood)(BMNH). South Africa:
1 9, Natal, Lake Sibayi, 13-24.iii.1968 (D. J. Brothers) (AM, Grahamstown).
114 J. QUINLAN
Oberthuerella longicaudata Benoit
(Fig. 123)
Oberthuerella longicaudata Benoit, 1955 : 291. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DEscrRIPTION. 2. Antenna 13-segmented (apical three segments missing), subclavate, segments 3 and 4
subequal, 5 longer than 4, remaining segments gradually decreasing in length. SN 3-1O=AR 20: 20:
22:19:18: 18:18:18. Face with reticulate-rugose sculpture and fine scattered pubescence, medial
carina extending to median ocellus. Pronotum viewed dorsally foveolate, mesoscutum with transversely
wrinkled reticulate-rugose surface, notaulices distinct but submerged in the surface sculpture. Scutellum
coarsely foveolate with five canaliculate foveae at base, median apical spine short, equal to apical width
of scutellum, lateral bars produced to form weak spines on either side of medial spine. Propodeum
angular on side margins, weakly pubescent, carinae bowed with foveolae within medial area. Mesopleurae
with transversely striate sculpture on upper half, lower half polished. Segment 1 of gaster viewed dorsally
broader than long, crenulate, segment 2 slightly shorter than 3, 3 longer than 2, 4 longer than 3, viewed
laterally finely punctate, apical margins of 4, 5 and 6 with fringe of hairs, hypopygium short, not pro-
truding, ventral spine short. Legs orange, hind femur with distinct perpendicular tooth, hind tibiae
polished with widely spaced foveolae medially, hair fringe on margins weak. Wings infuscate, surface
ciliate, margins bare, radial cell of forewing closed, first cubital cell open, second cubital (areolet) not
indicated, cubitus almost reaching apex of wing. Colour: antenna, head and thorax black, segment 1 of
gaster mainly black, apex reddish, gaster orange-red.
3 unknown.
MATERIAL EXAMINED
Holotype °, Zaire: Tshuapa, Eala, v.1935 (J. Ghesquiére) (MRAC, Tervuren).
Oberthuerella longispinosa Benoit
(Fig. 126)
Oberthuerella longispinosa Benoit, 1955 : 290. Holotype 3, ZAIRE: (MRAC, Tervuren) [examined].
DESCRIPTION... Antenna 13-seemented: SN 3-13=AR: 107117: 1413::'122 10: 10: 102 828% 13:
$. Antenna 14-segmented, subclavate, segment 3 shorter than 4, 4 longer than 5, 5 and 6 subequal, 7-13
subequal, each shorter than 6, 14 shorter than 12+13. SN 3-14=AR 13:20:18:18:16:16:16:15:
15: 15:15:26. Face with median carina extending to median ocellus, surface rugose-reticulate with fine
sparse pubescence and strigose sculpture medially. Pronotum and mesoscutum coarsely reticulate-rugose,
median scutal line obsolete, visible apically, notaulices submerged in surface sculpture but distinct.
Scutellum coarsely reticulate-rugose with median apical spine as long as apical width of scutellum, lateral
bars in form of spines at apex, scutellar foveae canaliculate, five in number. Propodeum angular viewed
from behind, carinae parallel on sculptured surface, pubescent on side margins only. Mesopleura with
canaliculate sculpture on upper surface, polished below. Segment | of gaster longer than broad, sulcate,
segment 2 viewed dorsally shorter than 3, 3 shorter than 4, 5 largest with long scattered hairs, viewed
laterally segment 5 with fine dense punctures. Legs completely orange-red, hind femur with distinct
perpendicular tooth, hind tibia coarsely reticulate and pubescent on the outer margins, inner side with a
longitudinal carina. Wings infuscate, surface ciliate, margins bare, radial cell of forewing completely
closed, first cubital cell appearing open (venation not indicated), second cubital (areolet) absent, cubitus
extending past middle of wing. Colour: antenna, head and thorax black, segment | of gaster chestnut-
red, remaining segments orange-red.
MATERIAL EXAMINED
Holotype 3, Zaire: Iles des Elephants (en aval de Roma), 30.xi.1939 (H. J. Bredo) (MRAC, Tervuren).
Gabon: | ¥, ‘Congo Franc’, Ogooue N’kogo, 1901 (J. Bouysson) (MNHN, Paris); 1 9°, Ogooue,
Lambarene, 1912 (R. Ellenberger) (MRAC, Tervuren). Ivory Coast: 1 9, Andé Bongouuno, iii.1962
(J. Decelle) (MRAC, Tervuren). Malawi: | 2, Ruo Valley, 2000 ft [600 m], 14.xii.1913 (S. A. Neave)
(BMNH).
Oberthuerella nigra Kieffer
Oberthuerella nigra Kieffer, 1910 : 110. Holotype 3, EQUATORIAL GUINEA (‘Spanish Guinea’) (originally in
MNHU, Berlin, now missing).
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 115
I have not included this species in the key as the taxonomic characters used by Weld (1952) and
Benoit (1955) are based on colour only. Neither author appears to have seen the type-material,
which is now missing.
Oberthuerella nigrescens Benoit
(Fig. 124)
Oberthuerella nigrescens Benoit, 1955 : 288. Holotype 2, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 2. Antennae 13-segmented, subclavate, SN 1-13 =AR 22:7:25:26:24:20:17:17:16:
16: 14:13:20. Face viewed frontally coarsely reticulate-rugose with fine scattered pubescence, median
carina extending to median ocellus, eyes viewed frontally further apart than the height of an eye, cheeks
weakly converging, head viewed dorsally reticulate-rugose either side of lateral ocelli, weak striae
medially. Pronotum dorsally scaly-reticulate with interspaces as big as the reticulate areas. Mesoscutum
coarsely reticulate-rugose, median scutal line obsolete, notaulices distinct though submerged in surface
sculpture of mesoscutum. Scutellum with canaliculate foveae at base, apex of scutellum with a distinct
medial spine as long as apical width of scutellum, smooth and polished, medial area of scutellum coarsely
reticulate-rugose, lateral bars of scutellum produced apically to form weak spines either side of median
spine. Propodeum with distinct carinae medially and wide flanged-shaped areas on either side. Meso-
pleurae with reticulate-rugose sculpture in upper half, lower half mainly polished with broken reticulate
outer margins. Segment | of gaster viewed dorsally slightly longer than broad, sulcate, segment 2 viewed
dorsally one-third as long as segment 3, 3 half as long as 4, apical margin of segment 2 reflexed inward,
side margins of tergites 2 and 3 with fine short scattered pubescence, tergite 5 the largest viewed laterally
with fine scattered punctures in median basal area, long scattered pubescence anteriorly on apical region
and dense small punctures in basal apical area, tergites 5 and 6 densely punctate with scattered long hairs,
hypopygium short, not protruding, ventral spine short. Hind femur with a suberect median tooth or
spine, hind femur coarsely foveolate-reticulate with comb of hairs on margin and a distinct longitudinal
carina medially on inner side. Wings infuscate, surface ciliate, margin of wing bare, radial cell of forewing
completely closed, cubitus almost reaching apex of wing, first cubital cell closed, second cubital (areolet)
complete, closed (Fig. 124). Colour: antenna, head, thorax and segment 1 of gaster (petiole) black,
remaining segments dark chestnut-brown, legs dark brown.
3 unknown.
MATERIAL EXAMINED
Holotype 2°, Zaire: Tshuapa, Eala, ix.1935 (J. Ghesquiére) (MRAC, Tervuren).
Oberthuerella tibialis Kieffer
Oberthuerella tibialis Kieffer, 1904 : 107. Holotype 2, CAMEROUN (? depository) [not examined].
The whereabouts of the holotype of this species is not known. Benoit (1955) includes it in his
key but gives no indication as to whether he examined the type-material.
Oberthuerella triformis sp. n.
(Fig. 125)
DESCRIPTION. 2. Antenna 13-segmented, subclavate, SN 3-13=AR 15:15:20:15:12:12:10:8:8
8 : 10, segments 5-13 each very slightly broader than preceding one, segment 5 the largest. Face viewed
frontally transversely strigose medially, reticulate-rugose laterally, frontal line or keel not apparent,
eyes further apart than the height of an eye, cheeks weakly converging. Pronotum and mesoscutum
transversely aciculate, notaulices submerged in the transverse sculpture. Scutellum with five foveae at
base, medially reticulate-rugose, tridentate apically, the medial tooth not in the form of a spine. Pro-
podeum viewed posteriorly, angular, carinae of propodeum bowed, surface coarsely reticulate-rugose.
Mesopleura with strigose sculpture in part, smooth on lower half. Segment | of gaster longer than broad,
sulcate, segment 4 of gaster arcuate viewed dorsally, segment 5 the largest viewed laterally, segments 5-7
of gaster viewed laterally with a distinct apical fringe, hypopygium not protruding, ventral spine short.
Fore and mid legs reddish black, hind coxa and femur reddish yellow, tarsi black, hind femur with erect
spine between median area and apex, impunctate, hind tibia smooth and shining on outer margins, inner
margins with weak longitudinal carinae. Wings infuscate, densely ciliate on surface, margins bare,
116 J. QUINLAN
radial cell closed, first cubital cell closed, second cubital (areolet) absent, cubitus almost reaching apex of
wing. Colour: antenna, head, thorax and segment | of gaster (petiole) black, gaster reddish yellow, fore
and mid legs reddish brown, hind coxae black, femur and tibia reddish yellow, tarsi blackish.
3 unknown.
MATERIAL EXAMINED
Holotype °, Tanzania (‘Tanganyika’): Mshughaa, 30 miles [48 km] east of Singida, xii.1935-i.1936
(E. Burtt) (BMNH).
REMARKS. Distinguished from all other species in the genus by the absence of a long distinctive
scutellar spine (see Fig. 125).
TESSMANNELLA Hedicke
Tessmannella Hedicke, 1912a : 303. Type-species: Tessmannella spinosa Hedicke, by original designation
DIAGNOSIS. 2 antenna 13-segmented, subclavate; ¢ antenna 14-segmented. Face with reticulate to rugose
sculpture and scattered pubescence. Pronotum coarsely rugose with median tooth or spine viewed laterally.
Mesonotum with coarse variable sculpture, propodeum without pronounced side margins. Segment 1 of
gaster (petiole) three times as long as broad, segments 2-4 short viewed laterally and dorsally, segment 5
the largest. Hind femur with a rounded lobe between medial area and apex, tooth on hind femur angled,
hind tibia with a distinct lobe apically, opposite the tibial spines, scutellar foveae three in number.
DISTRIBUTION. Gabon; Zaire.
Discussion Only four species are at present known in this genus, including one described for
the first time in this paper. Hedicke (1912a) separated Tessmannella from Oberthuerella by the
presence of a median tooth on the pronotum and by the spine on the hind coxae being obliquely
angled.
Biology unknown.
Key to species of Tessmannella Hedicke
1 Forewing infuscate medially, with distinctive light-coloured area basally and medially in the
cubital cell (Fig. 108). (Q unknown) : A : : : . transiens Benoit (p. 118)
— Forewings completely infuscate : Z
2 Scutellar foveae either angular or rounded apically, not extending past the middle of lateral bars
(Fig. 106), legs red except for black hind tibia ; 3
— Scutellar foveae extending past middle of lateral bars, triangular apically (Fig. 105); fore and
mid legs dark chestnut-brown, hind legs entirely black. (¢ unknown) nigra Hedicke (p. 117)
3 Notaulices obscured by strong crenulations along whole length; scutellar foveae angled at apex;
facial cheeks converging weakly towards clypeus (Fig. 104), face rounded medially, weakly
pubescent, clypeus with transverse striations, mandibles with lobed teeth; hind tarsi red;
lateral bars (outer apical spines of scutellum) parallel. (¢ unknown) . spinosa Hedicke (p. 117)
— Notaulices broad basally, not obscured by crenulations, weakly crenulate at base, mainly shiny,
scutellar spine long and narrow at apex, foveae rounded at apex (Fig. 106), facial cheeks
strongly converging towards clypeus, face flattened medially, pubescent, mandibles with
incised teeth, hind tarsi black, lateral bars of scutellum acutely angled. (¢ unknown) expansa sp. n.
(p. 116)
Tessmannella expansa sp. n.
(Figs 106, 107, 118)
DESCRIPTION. °. Antenna 13-segmented, subclavate. SN 3-13=AR 22: 38 : 36: 35: 28:26:24: 22:22:
20: 35. Face with irregular reticulate sculpture with long hairs issuing at base of each reticulation,
mandibles with basal tooth blunt, incision deep, clypeus with weak coriaceous sculpture, eyes less than
twice as far apart as height of eye viewed frontally, equal to length of malar space, cheeks weakly con-
verging towards clypeus. Pronotum viewed laterally with median tooth or spine, frontal surface coriaceous,
sides reticulate-rugose. Mesoscutum with broken widely spaced transverse sculpture, notaulices impressed
on the surface sculpture and wide, median scutal furrow indicated by a depression, obscured by the
transverse sculpture. Scutellum with three foveae at base, apex of scutellum with median spine and lateral
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION bags
bars extended to form striate spines either side of median spine, surface of scutellum with irregular rugose
sculpture. Mesopleurae with canaliculate sculpture on upper half and long hairs on a weakly reticulate
polished surface on lower half. Propodeum sulcate at apex. Segment 1 of gaster sulcate, 3-5 times as long as
broad, tergites 2-4 short, tergite 5 the largest, apex of tergite 5 and remaining visible tergites with long
scattered hairs and dense punctures interspaced between the hairs, ovipositor sheath with dense hairs on
the lower margins. Hind femur with a distinct tooth on inner margin which is directed backwards almost
in same plane as femur, apex of femur emarginated (Fig. 129), hind tibia with reticulate-strigose sculpture
and long hairs or spines, apex of hind tibia with a distinct lobe, tarsus 5-segmented with long hairs or
spines on a reticulate surface. Radial cell of forewing completely closed, long and narrow, cubitus extend-
ing almost to edge of wing, wing surface ciliate, margin bare, whole wing infuscate. Colour: antenna
blackish brown, head reddish on face, blackish dorsally, thorax black, gaster chestnut-brown, all legs
reddish brown except hind tibia and tarsi black.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Gabon: Ogooue, Lambarene, 1910 (R. Ellenberger) (MNHN, Paris).
REMARKS. This species has close affinities with spinosa but is separated from it by a number of
characters (see key).
Tessmannella nigra Hedicke
(Fig. 105)
Tessmannella nigra Hedicke, 1912a : 304. Holotype 2, ZAIRE (MNHU, Berlin) [examined].
DESCRIPTION. 2. Antenna 13-segmented, subclavate. Face with reticulate surface sculpture and fine
white hairs issuing at base of reticulations, clypeus with very weak transverse striations, eyes further apart
than height of eye, cheeks converging sharply towards clypeus viewed frontally. Pronotum viewed
laterally with median tooth or spine, frontal surface polished, sides coarsely reticulate-rugose, meso-
scutum appearing coarsely rugose with rugosity consisting of short transverse scales, notaulices and
median scutal line impressed in the sculptured surface. Scutellum with three foveae at base, the outer
two extending to apex of lateral bars which are finely striated and ending in a spine or tooth (Fig. 105),
apex of scutellum with long median spine, surface of scutellum deeply rugose. Mesopleurae with canal-
iculate sculpture on upper half, lower half with long pubescence. Propodeum sulcate at apex. Segment 1
of gaster sulcate, three times as long as broad, tergites 2-4 short, tergite 5 the largest, apex of tergite 5 and
remaining visible tergites with long scattered hairs and dense punctures interspaced between the hairs,
Ovipositor sheath with dense hairs on the lower margins. Hind femur with a distinct tooth on inner
margin which is directed backwards almost in the same plane as femur, apex of femur rounded-emarginate
(cf. Fig. 109), hind tibia with strong reticulate-canaliculate sculpture and long hairs, apex of hind tibia
with a distinct lobe and two spines, tarsus 5-segmented with fine coriaceous sculpture. Radial cell of
forewing completely closed, long and narrow, cubitus extending past middle of wing, surface of wing
ciliate, margin bare, whole wing infuscate. Colour: antenna black, head, thorax and gaster black, front
legs dark brownish black on coxae and femur, mid and hind legs completely black.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Uelleburg, vi—viii.1908 (von Tessmann) (MNHU, Berlin).
Zaire: 1 2 Kibombo, 2.xi.1910 (Bequaert) (MRAC, Tervuren).
Tessmannella spinosa Hedicke
(Figs 103, 104)
Tessmannella spinosa Hedicke, 1912a : 303. Holotype 2°, ZAIRE (MNHU, Berlin) [examined].
DESCRIPTION. 2. Antenna 13-segmented, subclavate. Face with reticulate surface sculpture and fine white
hairs issuing at base of reticulations, clypeus with transverse striations, eyes further apart than height of
eye viewed frontally, less than length of malar space, cheeks strongly rounded or curved, converging
strongly towards clypeus (Fig. 104). Pronotum viewed laterally with a median tooth or spine (Fig. 116),
frontal surface polished, sides coarsely sculptured. Mesoscutum coarsely reticulate-rugose with notaulices
118 J. QUINLAN
and median scutal line impressed (Fig. 103), scutellum with spine at apex and shorter ones either side
(Fig. 103), scutellum with three foveae at base, not extending to apex at lateral bars, surface of scutellum
strongly reticulate-rugose. Mesopleurae with canaliculate sculpture on upper half, lower half shiny with
scattered pits. Propodeum sulcate at apex. Segment 1 of gaster sulcate, almost four times as long as wide,
tergites 2-4 short, tergite 5 the largest, apex of tergite 5 and remaining visible segments with fine light
scattered hairs and dense punctures, ovipositor sheath with dense hairs on lower margin. Legs red except
for hind tibia blackish, hind femur with a distinct tooth on inner margin which is directed backward
almost in same plane, apex of femur with rounded lobe (cf. Fig. 109), hind tibia with reticulate-rugose
sculpture with long hairs, tarsus 5-segmented with long hairs, apex of hind tibia with a distinct lobe and
two spines. Radial cell of forewing completely closed, long and narrow, cubitus extending to past middle
of wing, surface of wing ciliate, margin bare, whole wing infuscate. Colour: antenna brownish black,
head and thorax black, gaster chestnut-brown, front and mid legs reddish brown, hind femur and tarsi
reddish brown, tibia blackish brown.
3 unknown.
MATERIAL EXAMINED
Holotype 2, Zaire: Uelleberg, vi-viii.1908 (von Tessman) (MNHU, Berlin).
Tessmannella transiens Benoit
(Figs 108, 116, 118)
Tessmannella transiens Benoit, 1955 : 283. Holotype 3d, ZAIRE (MRAC, Tervuren) [examined].
DESCRIPTION. 3. Antenna 14-segmented, subclavate. Face with coarse reticulate sculpture and scattered
pubescence, frontal line distinct, branching before reaching frontal area, occipital carinae pronounced,
eyes further apart than height of eye, cheeks converging weakly. Pronotum viewed laterally with a
median tooth or’spine (Fig. 116), dorsal surface with coarse reticulate-rugose sculpture and foveolae.
Mesonotum scaly-reticulate, notaulices impressed in surface sculpture, indistinct. Scutellum with three
apical spines, scutellar foveae extending partly along lateral bars. Mesopleurae with canaliculate sculpture
on upper half, lower half smooth with weak pits apically, propodeum sulcate. Segment | of gaster sulcate,
four times as long as broad, tergites 2-4 short, viewed dorsally evenly curved (Fig. 118), tergite 5 the
largest, viewed laterally densely punctate, apex of tergites 5 and remaining visible segments with long
distinct scattered hairs. Legs black, hind femur with a distinct tooth on inner margin directed weakly
towards apex, apex of femur with a distal lobe, hind tibia scaly-reticulate with long scattered hairs, apex
of hind tibia with two spines and a distinct lobe, tarsus 5-segmented, surface sculpture fine reticulate.
Radial cell of forewing completely closed, five times as long as broad, cubitus faint, extending almost to
apex of wing, wing surface ciliate, margin bare, wing infuscate medially to apex except for the medial
area of the cubital cell.
Colour: completely black except for lower margins of basal segments of gaster which are chestnut-
brown.
2 unknown.
MATERIAL EXAMINED
Holotype 3, Zaire: Ditri Tanganikam Albertville, 1-20.i1.1919 (R. Mayne) (MRAC, Tervuren).
XENOCYNIPS Kieffer
Xenocynips Kieffer, 1910: 340. Type-species: Xenocynips subsquamata Kieffer, by original designation
and monotypy.
DIAGNosis. 2. Antenna 13-segmented, claviform. Head and thorax strongly rugose. Scutellar foveae
obscured by longitudinal carinae, scutellum reticulate-rugose, apex triangular. Pronotum viewed laterally
with erect tooth or spine on hind margin anterior of mesoscutum (Fig. 32). Hind femur without a lobe
distally, with an erect spine medially. Gaster with tergites 2-4 fused without trace of a suture (Fig. 32).
DISTRIBUTION. Zaire.
Discussion. Only one species, based on a single specimen, has so far been placed in this genus.
It differs from Tessmannella primarily by segments 2-4 of the gaster being fused and the spine
on the hind coxae being erect. Weld (1952) suggested that at first appearance the size and shape
of the gastral tergites could confuse it with the Figitinae.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 119
Xenocynips subsquamata Kieffer
(Figs 32) 1105 119, 112; 113)
Xenocynips subsquamata Kieffer, 19106 : 340. Holotype 2, CAMEROUN (IP, Eberswalde) [examined].
DESCRIPTION. °. Antenna 13-segmented, claviform, rhinaria on segments 4-13, segment 4 three times
length of 3 (Fig. 112), basal segment brown, 2-8 yellowish, remainder brownish yellow. Head, pronotum
and mesoscutum dorsally strongly rugose. Scutellar foveal depression obscured by longitudinal carinae,
surface of scutellum reticulate-rugose, apex triangular, ending in conical protrusion; propodeum with
parallel carinae. Pronotum viewed laterally with erect tooth on hind margin at anterior end of meso-
scutum. Mesopleura polished with broken striate sculpture, fore and mid coxae yellowish brown, hind
coxae blackish, elongate and swollen with scattered pubescence, hind femur with erect spine (Fig. 111),
without a distal lobe. Segment | of gaster measured at extreme points less than three times as long as
broad, crenulate, segment 2 of gaster small, fused with segments 3-4, segment 5 the largest with short
regular hairs on apical half. Colour: antenna yellow basally, becoming darker towards apex, head and
thorax black, gaster chestnut-red basally, brownish black on apical half, fore and mid legs yellowish,
hind legs reddish brown on coxae and femur, tibia and tarsi dark reddish brown. Forewings with scattered
short pubescence, wing margin without an apical hair fringe, radial cell completely closed, long and
narrow, 4:5 times as long as its widest point, with a brownish patch extending throughout cell and well
below, vein 2rm short, M (cubitus) weak, extending almost to wing margin (Fig. 113).
3 unknown.
MATERIAL EXAMINED
Holotype °, Cameroun (Conradt) (IP, Eberswalde).
References
Andrews, F. G. 1978. Taxonomy and host specificity of Nearctic Alloxystinae with a catalog of the world
species (Hymenoptera : Cynipidae). Occ. Pap. Bur. Ent. Calif. 25: 1-128.
Benoit, P. L. G. 1955. Les Liopteridae Oberthurellinae du Congo Belge. Revue Zool. Bot. afr. 52 : 283-293.
—— 1956. Figitidae — Aspicerinae nouveaux du Congo Belge. Revue Zool. Bot. afr. 53 : 195-204.
Cameron, P. 1904a. Descriptions of new genera and species of Hymenoptera from Dunbrody, Cape
Colony. Rec. Albany Mus. 1: 125-160.
— 1904b. On the Hymenoptera of the Albany Museum, Grahamstown, S. Africa. Rec. Albany Mus.
$161:
Crosskey, R. W. & White, G. B. 1977. The Afrotropical Region, a recommended term in Zoogeography.
J. nat. Hist. 11: 541-544.
Dalla Torre, K. W. & Kieffer, J. J. 1910. Cynipidae. Tierreich 24: 1-891.
Eady, R. D. 1968. Some illustrations of microsculpture in the Hymenoptera. Proc. R. ent. Soc. Lond. (A)
42 : 66-72.
—— 1974. The present state of nomenclature of wing venation in the Braconidae (Hymenoptera); its
origins and comparison with related groups. J. Ent. (B) 43 : 63-72, 10 figs.
Eady, R. D. & Quinlan, J. 1963. Hymenoptera Cynipoidea. Key to families and subfamilies and Cynipinae
(including galls). Handbk Ident. Br. Insects 8 (la) : 1-81.
Hedicke, H. 1912a. Beitrige zur Kenntnis der Cynipiden (Hym.) I. Eine neue Gattung aus der Unter-
familie der Oberthiirellinae. Dt. ent. Z. 38 : 303-304.
—— 19126. Beitrage zur Kenntnis der Cynipiden. II. Zur Systematik der Oberthiirellinae K. Ent. Rdsch.
29 : 81-82.
Hedicke, H. & Kerrich, G. J. 1940. A revision of the family Liopteridae (Hymenopt., Cynipoidea).
Trans. R. ent. Soc. Lond. 90 : 177-225.
Kerrich, G. J. 1973. On the taxonomy of some forms of /balia Latreille (Hymenoptera : Cynipoidea)
associated with conifers. Zool. J. Linn. Soc. 53 : 65-79.
Kieffer, J. J. 1903. Description de trois Cynipides exotiques. Marcellia 2 : 84-90.
—— 1904. Beitrage zur Kenntnis der Insektenfauna von Kamerun. N : 0 26. Beschreibung einer neuer
Cynipide aus Kamerun. Ent. Tidskr. 25 : 107-110.
1907. Beschreibung neuer parasitischer Cynipiden aus Zentral- und Nord-Amerika. Ent. Z.,
Frankf. a.M. 21 : 151-152, 157.
1910a. Description de nouveaux Hyménoptéres. Boll. Lab. Zool. gen. agr. Portici 4: 105-117.
—— 1910b. Nouveaux Cynipides exotiques. Boll. Lab. Zool. gen. agr. Portici 4: 340-341.
120 J. QUINLAN
1910c. Serphidae, Cynipidae, Chalcidae, Evaniidae und Stephanidae aus Aquatorialafrika. Wiss.
Ergebn. dt. Zent Afr. Exped. 1907-1908 3 (Zool.) 2: 1-20.
1912. Cynipides et Bethylides de l’Afrique du Sud. Annls Soc. ent. Fr. 80: 451-462.
Kinsey, A. 1919. An African Figitidae. Psyche, Camb. 26 : 162-163.
Riek, E. F. 1971. A new subfamily of cynipoid wasps (Hymenoptera : Cynipoidea) from Australia.
Entomological Essays to Commemorate the Retirement of Professor K. Yasumatsu. Hokuryukem Ltd.,
Tokyo.
Richards, O. W. 1978. Hymenoptera. Introduction and key to families. Handbk Ident.Br. Insects 6 (1):1-100.
Rohwer, S. A. & Fagan, M. M. 1917. The type-species of the genera of the Cynipoidea, or the galls and
parasitic Cynipoids. Proc. U.S. natn. Mus. 53 : 357-380.
Ross, H. H. 1936. The ancestry and wing venation of the Hymenoptera. Ann. ent. Soc. Am. 29 : 99-111.
Saussure, H. de. 1890. Histoire physique, naturelle et politique de Madagascar. xxi+ 590 pp., 27 pls. Paris.
Sclater, P. L. 1858. On the general geographical distribution of members of the class Aves. J. Proc. Linn.
Soc. (Zool.) 2 : 130-145.
Snodgrass, R. E. 1910. The thorax of the Hymenoptera. Proc. U.S. natn. Mus. 39 : 37-91.
Wallace, A. R. 1876. The geographical distribution of animals, with a study of the relations of living and
extinct faunas as elucidating the past changes of the earth’s surface. 1, 503 pp. London.
Weld, Lewis H. 1930. Notes on types (Hymenoptera, Cynipidae). Proc. ent. Soc. Lond. 32 : 137-144.
— 1952. Cynipoidea (Hym.) 1905-1950. 351 pp. Ann Arbor, Michigan (privately published).
Yoshimoto, Carl M. 1970. A new subfamily of Cynipoidea (Hymenoptera) from Nepal. Can. Ent. 102:
1583-1585.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 121
metatarsus
10
es
11
Figs 1-11 1-8, gaster, lateral view of 2 of (1) Ibaliidae; (2) Oberthuerellinae; (3) Liopterinae;
(4) Mesocynipinae; (5) Aspicerinae; (6) Figitinae; (7) Eucoilidae; (8) Anacharitinae. 9, forewing,
Ibaliidae. 10, hind tarsus, Ibaliidae. 11, hind femur, Oberthuerellinae.
122 J. QUINLAN
Figs 12-21 12, scutellar spine, Oberthuerellinae. 13, pronotal plate, Eucoilidae. 14, scutellum,
Eucoilidae. 15-17, forewing of (15) Eucoilidae; (16) Figitidae; (17) Cynipidae. 18, 19, gaster of
(18) Rhoopilus; (19) Andricus. 20,21, forewing of (20) Himalocynipinae; (21) Anacharitinae.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 123
23
ee
lateral disc
lines =
pronotal Nis bars
ae Gan eee
“rhinaria
ao =
1
1
1
1
1
1
scutellar
notaulices or foveae=
hair-lines — nterior Cup = inner disc !
anterior parallel foveae YY '
'
1
1
1
I
'
!
ln eee eee ew ewes flagellins sia. slot e = wrote ele wee oe
29
ventral border metapleura
of pronotum
mesosternum
Figs 22-30 22, head, dorsal view, Anacharitinae, 23, forewing, Pycnostigmatinae. 24, pronotum,
dorsal view, Alloxystinae, 25, pronotum, lateral view, Alloxystinae. 26, pronotum, dorsal view,
Cynipinae. 27, pronotum, lateral view, Cynipinae. 28, 2 antenna, Eucoilidae. 29, thorax, Eucoilidae.
30, pronotal plate, Eucoilidae.
124 J. QUINLAN
Din Ges ess
| oe pee
Figs 31-42 31, 32, gaster of (31) Synergus; (32) Xenocynips. 33, mesonotum, Aspicera. 34, hind tibia,
Aspicera. 35, radial cell, Synergus. 36, pronotum, dorsal view, Synergus. 37, pronotum, lateral view,
Synergus. 38-40, basal antennal segments of ¢ of (38) Cynipinae; (39, 40) Alloxystinae. 41, 42,
forewing of (41) Prosaspicera antennata; (42) P. kisantua.
ETHIOPIAN ZOOGEOGRAPHICAL REGION 125
CYNIPOIDEA OF THE
'
we
Hy '
(AR
re beanie
Pye!
Fx
ms
een IN
Figs 43-53 43, hind tibia, Prosaspicera. 44, mesonotum, Anacharoides arcus. 45, forewing, A
elongaticornis. 46, hind tibia, A. sanitas. 47, mesonotum, Prosaspicera africana. 48, head, P.
kisantua. 49, 50, $ antenna of (49) P. antennata; (50) P. optivus. 51, mesonotum, P. antennata.
52, mesopleura, P. antennata, 53, forewing, P. paragicida.
126 J. QUINLAN
Figs 54-64 54, head, Prosaspicera optivus. 55, mesonotum, P. optivus. 56-58, head of (56) P. africana;
(57) P. kisantua; (58) P. tropica 3. 59, 2 antenna, Anacharoides elongaticornis. 60, scutellar foveae,
A. elongaticornis. 61, mesonotum, A. suspensus. 62, scutellar foveae, A. sanitas. 63, 64. mesoscutum
of (63) A. striaticeps; (64) A. stygius.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 127
spar Boe,
wr? Pag peor,
<A! ae
ay it A)
\\)
et 3
Ae Nt
70
Figs 65-74 65, forewing, Anacharoides stygius. 66, 2 antenna, A. stygius. 67-69, forewing of (67) A.
paragi; (68) A. suspensus; (69) A. sanitas. 70, 71, radial cell of (70) A. arcus; (71) A. nigra. 72, fore-
wing, A. eurytergis. 73, head, A. eurytergis. 74, mesonotum, A. eurytergis.
128 J. QUINLAN
SU)
80
Figs 75-86 75-77, mesonotum of (75) Anacharoides nigra; (76) A. decellius; (77) A. pallida. 78-81,
3 basal antennal segments of (78) A. quadrus; (79) A. suspensus; (80) A. arcus; (81) A. striaticeps.
82, mesonotum, A. suspensus. 83, hind coxa, A. suspensus. 84, $ antenna, A. decellius, 85, hind
coxa, A. decellius. 86, forewing, A. striaticeps.
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION 129
Figs 87-99 87, 2 antenna, Anacharoides striaticeps. 88, 3 basal antennal segments, A. sanitas.
89, scutellar foveae, A. quadrus. 90, 91, 3, antenna of (90) A. astrida; (91) A. eurytergis. 92, head,
vertex, A. sanitas. 93, radial cell, A. pallida. 94, 2 antenna, Prosaspicera africana. 95-98, occiput of
(95) P. africana; (96) P. antennata; (97) P. optivus; (98) P. kisantua. 99, 2 antenna, P. kisantua.
130 J. QUINLAN
100
lateral ocellus
median ocellus
compound vertex antenna
eye
‘frontal
line
mandible
1 6) 4 clypeus
Figs 100-108 100, 2 antenna, Prosaspicera paragicida. 101, 102, 3 antennal segments of (101) P.
africana; (102) P. paragicida. 103, mesonotum, Tessmannella spinosa. 104, head, T. spinosa. 105,
106, mesonotum of (105) 7. nigra; (106) T. expansa. 107, head, T. expansa. 108, forewing, T.
transiens.
131
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION
My, aN!
a) ii
fl i
i nr inf i OY Mt iN"
rn a AY
Ne Cy \ Hy)
Figs 109-119 109, hind femur, Tessmannella sp. 110, scutellum, Xenocynips subsquamata. 111, hind
femur, X. subsquamata. 112, 2, basal antennal segments, X. subsquamata. 113, forewing, Xenocynips
subsquamata. 114, first segment of gaster, Oberthuerella lenticularis. 115, propodeum, Tessmannella
sp. 116, pronotum, lateral view, 7. transiens. 117, propodeum, Oberthuerella sp. 118, first segment
of gaster, Tessmannella expansa.
132 J. QUINLAN
(=
127
Figs 120-129 120, 121, gaster of (120) Oberthuerella aureopilosa; (121) O. compressa. 122-124, radial
cell of (122) O. compressa; (123) O. longicaudata; (124) O. nigrescens. 125, mesonotum, O. triformis.
126-127, gaster of (126) O. longispinosa; (127) O. crassicornis. 128, forewing, O. /lenticularis.
129, apex of hind femur, Tessmannella expansa.
128
CYNIPOIDEA OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION
Index
Principal references are in bold, synonyms in italics.
abscinda 92, 111
aegyptica 93
africana 92, 105, 106
Alloxystinae 88, 89, 91, 92
Anacharitinae 86, 87, 91, 92
Anacharoides 88, 92, 93, 94
Andricus 88
antennata 92, 105, 107
arcus 96
Aspicera 93, 94
Aspicerinae 86, 87, 89, 91, 92, 93, 96
astrida 92, 94, 95, 96, 97
aureopilosa 92, 110, 112
Austrocynipinae 91, 92
Austrocynips 86
breviscutellaris 92, 111, 112
Ceroptres 88
Coelonychia 93, 94
compressa 92, 112
crassicornis 92, 111, 112
Cynipidae 87, 88, 89, 90, 91, 92
Cynipinae 88, 91, 92
decellius 92, 95, 96, 97, 100
elongaticornis 92, 94, 98
ensifera 105
Eucoilidae 86, 87, 89, 90
eurytergis 92, 95, 97, 99, 100
expansa 92, 116
Figitidae 87, 89, 90, 91, 92
Figitinae 86, 87, 89, 91, 93
gibbosus 92, 103
Himalocynipinae 86, 87, 91, 92
Himalocynips 86
Ibaliidae 87, 90, 91
kisantua 92, 105, 106, 107
lenticularis 92, 110, 111, 113
Liopteridae 89, 90, 91, 92
Liopterinae 90, 91
lobata 93
longicaudata 92, 111, 114
longispinosa 92, 111, 114
Mesocynipinae 87, 90, 91
Neralsia 87
Neuroterus 88, 89
nigra 92, 95, 98, 99, 114, 117
nigrescens 92, 111, 115
Oberthuerella 92, 110, 116
Oberthuerellinae 87, 90, 91, 92, 109
optivus 92, 105, 108
pallida 95, 100
paragi 92, 94, 95, 101
paragicida 92, 105, 106, 108
Prosaspicera 92, 93, 104, 105
Pycnostigmatinae 86, 87, 91, 92
quadrus 92, 95, 96, 101, 102
Rhoopilus 88
rufa 92, 93, 102
Synergus 88
sanitas 92, 95, 102
spinosa 92, 117
spinosipes 92, 93, 103
striaticeps 92, 93, 94, 95, 103
subsquamata 92, 93, 118, 119
suspensus 92, 94, 95, 98, 104
stygius 92, 94, 95, 103
Tessmannella 92, 109, 110, 116
tibialis 92, 115
transiens 92, 116, 118
triformis 92, 111, 115
tropica 92, 105, 106, 109
Xenocynips 87, 92, 109, 110, 118
133
ae a ae
ee
British Museum (Natural History)
Monograph
The social wasps of the Americas
O. W. Richards 1978
Social wasps are particularly numerous in South America, both in genera and
species. Their nest-building habits are of great interest because of the great variety
of architecture, sometimes even in closely allied species. This volume deals with
the American social wasps except the Vespinae (those resembling the British
wasps) which are northern in distribution and only just extend to Mexico. The
nests, habits and larvae of the wasps are described as far as they are known. The
main purpose of the work, however, is to make it possible to identify the 500
species of wasps. There is no recent work for this purpose and there has never
been a really comprehensive one. In preparing this volume a great many of the
types of earlier authors have been examined, including those of Zikan which have
hitherto been difficult to trace.
240x160mm Hard-bound PP 571 Index 159 text figures 4 colour plates
ISBN 0 565 00785 8
£32.50
Titles to be published in Volume 39
A revision of the African pod bugs of the tribe Clavigrallini
(Hemiptera: Coreidae) with a checklist of the world species.
By W. R. Dolling.
A revisionary classification of the Cynipoidea (Hymenoptera) of the
Ethiopian Zoogeographical Region. By J. Quinlan.
A review of the genera of the Hydroptilidae (Trichoptera).
By J. E. Marshall.
A revision of Vitessidia Rothschild & Jordan and Vitessa Moore
(Lepidoptera: Pyralidae). By E. G. Munroe & M. Shaffer.
Printed by Henry Ling Ltd, Dorchester
w GJ! f° ww
Bulletin of the
British Museum (Natural History)
A review of the genera of the Hydroptilida
(Trichoptera)
J. E. Marshall
Entomology series Vol 39 No3 20 December 1979
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and
an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum’s resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and are not
necessarily completed within one calendar year. Subscriptions may be placed for one or more
series. Subscriptions vary according to the contents of the Volume and are based on a forecast
list of titles. As each Volume nears completion, subscribers are informed of the cost of the
next Volume and invited to renew their subscriptions. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW7 SBD,
England.
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1979
ISSN 0524-6431 Entomology series
Vol 39 No 3 pp 135-239
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 20 December 1979
A review of the genera of the Hydroptilidae
(Trichoptera)
J. E. Marshall
Department of Butemoloey, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis. , 2 : ‘ ‘ ; > j : ‘ ; , 135
Introduction : F ? ; ‘ : : : ; ; ‘ ‘ 136
Material. F ; : 5 ; ; ’ 3 j : : , 136
Historical review : : : , : : : : . ; 136
Subfamily classification A : : : : : : ; 137
Systematic position within the Trichoptera ; : : ; ; : 138
General morphology. ; , , ; d ; ; ; : ; 139 -
Adult : ‘ : ; : : ; i : : : , : 139
Larva , : : ; : ; : : ‘ : : ; : 147
Pupa é : : : : : ; : : 2 : : , 152
Systematics . : " . ; : : : : ; 153
Family Hydroptilidae Stephens . : 3 : : : ; ‘ 153
Keys to genera. : ; : : i . : : : 153
Adult : ; : ' ‘ : ; ? 2 : : ; 153
Larva : : : ‘ ’ : : ; : 156
Subfamily Ptilocolepinae Martynov : 157
Keys to differentiate the Ptilocolepinae, Glossosomatidae and Rhyacophilidae 15?
Adult ; : ‘ : . ; ; , : j : : 159
Larva : : : : : : ak ok : ; 159
Pupa ‘ i : : : ‘ : , 159
Subfamily Hydroptilinae Stephens : : : : : i : : 161
Key to tribes . : : : ; : : F Z : 162
Tribe Stactobiini Botosaneanu , : : F ; ‘ ‘ : 163
Tribe Leucotrichiini Flint . ; F ; , , : ‘ : 17S
Tribe Ochrotrichiini trib. n. . : : . F , ; : : 184
Tribe Neotrichiini Ross : ; : ; ‘ ; : ; : 188
Tribe Hydroptilini Stephens : : : 5 ‘ , : ; 191
Tribe Orthotrichiini Nielsen : : as : : ; ; F 212
Incertae sedis. : ; : : ; : , , : 217
Phylogeny of the Hydroptilidae : : : ; . , : : 22
Genera here transferred from the Hydroptilidae : : : ‘ : : 223
Checklist of species of Hydroptilidae ; ; : F : ‘ : . Zar
Acknowledgements : ; : ; : ; ? : : , : 234
References . : ; : ’ ; ; : : : : F : 234
Index : : ; : ; : ; ‘ , : ‘ : ; 239
Synopsis
The Hydroptilidae, a world-wide group of small caddis flies, are reviewed down to the generic level for
the first time. As previously, two subfamilies are recognized, the primitive Ptilocolepinae and the more
typical Hydroptilinae, the latter being subdivided provisionally into six tribes, Stactobiini, Leucotrichiini,
Ochrotrichiini (new), Neotrichiini, Hydroptilini and Orthotrichiini. Thirteen genera and two species are
newly synonymized, three new combinations are established, one previously synonymized genus is re-
instated and three genera (Padunia Martynov, Tsukushitrichia Kobayashi and Petrotrichia Ulmer) are
transferred to other families of Trichoptera; thus 46 genera with 616 species are considered here as valid
Bull. Br. Mus, nat. Hist. (Ent.) 39 (3) : 135-239 Issued 20 December 1979
135
136 J. E. MARSHALL
members of the Hydroptilidae. For each genus accounts are given of the nomenclature, distribution,
morphology of both adults and immature stages, biology and, where appropriate, species groupings. Keys
are given to tribes for adults and to the world genera for both adults and larvae; keys are also provided
to differentiate the Ptilocolepinae from the closely related families Rhyacophilidae and Glossosomatidae.
The phylogeny of the family is briefly discussed and a synonymic checklist is given of all the species.
Introduction
As a result of a recent investigation into the morphology and biology of the Hydroptilidae, it
became apparent that very little was known of the basic systematics of the family. Previous
studies on the group have been very fragmentary and have consisted mainly of faunal descriptions
and keys, there being no comprehensive appreciation of the family as a whole. The aim of this
review has therefore been to investigate the higher classification of the Hydroptilidae with respect
to the present knowledge of adult and larval morphology, biology and distribution. Provisional
generic keys are given to the adults and to the known larvae, the latter incorporating morpho-
logical and case-form data. A full species checklist has also been included which, where possible,
indicates tentative species groupings. However, a complete species review was beyond the scope
of this work since the family contains over 600 species in 46 genera.
A number of interesting discoveries has been made regarding the systematics and taxonomy of
the Hydroptilidae. Fourteen genera and one species have been synonymized for the first time, three
species have been transferred to different genera and one genus has been reinstated; three genera
have been transferred to other families of Trichoptera. Two subfamilies have been recognized,
the primitive Ptilocolepinae and the more typical Hydroptilinae; within the latter six tribes have
been provisionally proposed. The Stactobiini is a predominantly Palaearctic group with a small
Nearctic component, the larval habits of which are paralleled by the exclusively New World
Leucotrichiini. Two other small but distinct Nearctic tribes are the Ochrotrichiini and the Neo-
trichiini while the remaining assemblage of the generally more successful and more widely dis-
tributed genera has been tentatively split into the Hydroptilini and Orthotrichiini.
This revision has been based on the existing literature and on the examination of available adult
and immature material. Each genus is separately discussed and data such as distribution, adult
and larval diagnostic features, biological details and species groupings are given where possible.
Before the detailed systematics are discussed, however, a brief historical review and introduction
to the general morphology of the adult and immature stages are given.
Material
Most of the material examined is in the collections of the British Museum (Natural History)
(BMNH); additional material was loaned or donated from the collections of the following
individuals and institutions.
Higler, L. W. G. Rijksinstitut voor Natuurbeheer, Leersum, Netherlands
Nybom, O. Imatra, Finland
Ross, H. H. Athens, Georgia, U.S.A.
Statzner, B. Kiel, West Germany
MCM Merseyside County Museums, Liverpool, England (Dr I. D. Wallace)
UHZIM Universitat Hamburg Zoologisches Institut und Zoologisches Museum, Hamburg,
West Germany (Dr H. Weidner, G. Ulmer Collection)
USNM Smithsonian Institution, National Museum of Natural History, Washington, D.C.,
U.S.A. (Dr O. S. Flint, Jr)
ZI Zoological Institute, Leningrad, U.S.S.R. (Dr L. Zhiltzova, S. I. Lepneva Collection)
Historical review
‘Trichoptera in general, but more especially the Hydroptilidae, are an annoyance to the Lepidop-
terist. If he lives near a river they are a constant source of disappointment to him whenever, on a
summer evening, he inspects the entomological miscellany which accumulates upon his tablecloth
beneath the gaslight. Down falls a crippled “‘Micro-’’; he has the trouble of looking at it; but his
THE GENERA OF THE HYDROPTILIDAE 137
first glance detects the scabrous clothing of the anterior wings of a Micro-Trichopteron, whose
hair looks as if it has been brushed the wrong way, in lieu of the scales of a Micro-Lepidopteron;
and his only consolation is that he may leave the pinning of that kind of things to the
Trichopterists.’
According to the attitude of most present day entomologists these words could well have been
written just yesterday but they were, in fact, part of the introduction to the Reverend A. E. Eaton’s
paper ‘On the Hydroptilidae, a family of the Trichoptera’ written over 100 years ago in 1873.
Since then the number of described genera and species has gradually increased from 6 and 21 to
approximately 50 and 600 respectively but, as stated in the Introduction, little work has been done
on the basic systematics of the group and much more material still needs to be collected. It is
probably the small size of these insects which is the main deterrent to their collection and identifi-
cation but, as in all groups, once they are purposefully sought they can be found in quite signifi-
cant numbers.
The first review of the family was, as already mentioned, that of Eaton (1873) in which he erec-
ted four new genera and described six new species. Since then the number of described taxa has
steadily increased due to the work of such notable trichopterists as McLachlan, Morton, Mosely
and Kimmins in Britain; Ris, Klapalek, Martynov and Schmid on the Continent, and Ross and
Flint in the U.S.A. Amongst these, Mosely is perhaps best known for his singular contribution to
the description of many new hydroptilid taxa not only from Great Britain and Europe but also
many then less well-known regions such as South America, Australia and New Zealand; indeed,
according to Kimmins (1951), the family Hydroptilidae was Mosely’s favourite group. Kimmins,
continuing the work of his predecessor, Mosely, also described a number of new taxa but his most
notable contribution was probably his revision of the British species of Oxyethira (Kimmins,
1958), a genus with rather complex male and female genitalia, despite which Kimmins succeeded
in providing descriptions, figures and keys to both sexes of almost every British representative.
Of the Continental contributors, Schmid is perhaps best known for his descriptions of many new
exotic genera and species from Sri Lanka, Iran and Pakistan (Schmid, 1958a; 19596; 1960) and
for his review of the genus Stactobia (Schmid, 1959a).
Ross, as well as describing many new Nearctic taxa, was the first to break away from the almost
classical use of wing venation in the taxonomy of the Hydroptilidae which, due to the small size
of these insects, does not provide such reliable characters as it does in the groups of larger caddis
flies. Instead Ross employed basic structures of the head, thorax and abdomen which provide
more stable features for the differentiation of genera and are also indicative of possible phylo-
genetic relationships within the Hydroptilidae and between trichopterous families in general. The
features thus used by Ross have provided the foundation for the studies outlined in this review.
Finally, Flint has made impressive contributions to our knowledge of the Central and South
American hydroptilid fauna, especially in his reviews of the Leucotrichiini and the genus Ochro-
trichia (Flint, 1970; 1972a) and his studies on certain faunal regions such as Jamaica and Surinam
(Flint, 19685; 1974).
The larvae, onthe other hand, are less welldocumented as regards specific descriptions although
a significant amount is known at the generic level due to the work of a number of authors of
whom special mention should be made of Nielsen. In his monumental work on ‘The biology of the
Hydroptilidae’, Nielsen (1948) describes the morphology, life histories, feeding and case building
behaviour of five hydroptilid genera with such detail and accuracy that no subsequent work on
this subject has surpassed or even equalled it. Unfortunately it does tend to give rather a restricted
impression of hydroptilid larvae in general because it concerns only five of the relatively more
advanced and specialized genera and does not stress the diversity of the group as a whole. Never-
theless, it is hoped that this present review will give a much broader appreciation of the family, not
only in its immature but also its adult stages, because this cosmopolitan group probably has the
greatest ecological, morphological and behavioural diversity within the whole of the order
Trichoptera.
Subfamily classification
Stephens (1836) erected the family Hydroptilidae for the genera Hydroptila Dalman, Agraylea
138 J. E. MARSHALL
Curtis and Narycia Stephens of which, however, the only species figured, Narycia elegans Stephens,
subsequently proved to be a tineid moth of the family Psychidae. The Hydroptilidae were basically
distinguished from other families of Trichoptera by the unfolded posterior wings and the filiform
antennae of the adults and the ‘cleft-like’ openings of the larval cases. The cases and larvae of
Hydroptila pulchricornis Pictet and Oxyethira flavicornis (Pictet) (as Hydroptila) had first been
described and illustrated by Pictet (1834) and, in fact, McLachlan (1880) considered Pictet to be
the true founder of the family since he had been the first to consider the group as a distinct taxon
under the general name of ‘les Hydroptiles’.
Nielsen (1948) was the first to attempt to divide the then large and heterogeneous group of
hydroptilid genera into subfamilies. Thus Nielsen proposed the Orthotrichiinae for Orthotrichia
and Ithytrichia and the Hydroptilinae for Agraylea, Oxyethira and Hydroptila on the basis of the
morphological affinities of the larvae. However, although Nielsen was aware of the other genera
known at that time, he only remarked on the relative positions of Ptilocolepus and Stactobia in
his scheme. The subfamily Stactobiinae was erected by Botosaneanu (1956) for Stactobia and ‘its
immediate relatives’ which, although not expressly named at the time, probably included Stacto-
biella, Plethus, Plethotrichia, Lamonganotrichia and, possibly, Catoxyethira. Ross (1956) proposed
a new Classification of the Hydroptilidae to include the Ptilocolepinae, a small subfamily con-
sisting of just two small genera with glossosomatid-like adults but typically hydroptilid-like larvae.
Ross thus divided the Hydroptilidae into the Ptilocolepinae and the Hydroptilinae, the latter
being further subdivided into the Hydroptilini and the Neotrichiini. Flint (1970) subsequently did
not adopt Ross’s (1956) classification when he proposed the subfamily Leucotrichinae (later
emended to Leucotrichiinae by Flint, in litt.) for the Neotropical genus Leucotrichia and its related
genera, since he retained the Ptilocolepinae, Stactobiinae and Orthotrichiinae as distinct sub-
families. Once again, however, the Hydroptilinae was left as a large and very heterogeneous group;
Table 1 shows the two principal approaches to the subfamily classification of the family and com-
pares them withthe system to be adopted here. The present views are based on those of Ross (1956)
in that only two subfamilies are recognized, with the Hydroptilinae being split into six tribes
which correspond to the subfamilies proposed by Flint (1970). These tribes, however, appear to
exhibit a number of common features which unite them and distinguish them as a whole from the
Ptilocolepinae.
Table 1 Summary of the three views of the subfamily classification of the
Hydroptilidae
Nielsen (1948)
Botosaneanu (1956) Proposed
Flint (1970) Ross (1956) classification
PTILOCOLEPINAE PTILOCOLEPINAE PTILOCOLEPINAE
HYDROPTILINAE HY DROPTILINAE
STACTOBIINAE HYDROPTILINI STACTOBIINI
LEUCOTRICHIINAE LEUCOTRICHIINI
ORTHOTRICHIINAE ORTHOTRICHIINI
HY DROPTILINAE OCHROTRICHIINI
HYDROPTILINI
NEOTRICHIINI NEOTRICHIINI
Systematic position within the Trichoptera
Eaton (1873) first recognized the relationships between adult hydroptilids and rhyacophilids
after agreeing with Pictet’s (1834) observations that the maxillary palpi of both sexes of the former
are five-segmented, thus placing the family in the old subdivision Aequipalpidae (later known as
the Aequipalpia). McLachlan retained the Hydroptilidae in the Aequipalpia but, on the basis of
the case building habits of the larvae, placed the family between the ‘tube-case’ building families
Sericostomatidae and Leptoceridae. McLachlan also regarded Ptilocolepus as a member of the
Glossosomatinae (‘section II of the Rhyacophilidae’) but noted that ‘the affinities are certainly
THE GENERA OF THE HYDROPTILIDAE 139
with Agapetus’ (Glossosomatinae) ‘but the thickened erect hairs of the anterior wings show an
analogy in the direction of the Hydroptilidae’.
Mosely (1939), on the basis of adult features, considered the family to be closely related to the
Rhyacophilidae and had particular difficulty in assigning the Nearctic Protoptila group to either
one of these families, at first including it in the Hydroptilidae but later transferring it to the Glos-
sosomatinae—Rhyacophilidae (Mosely, 1954) (see discussion on Padunia).
Nielsen (1948), using larval and pupal characters, placed the Hydroptilidae between the ‘saddle-
case’ building Glossosomatinae and the ‘tube-case’ making Integripalpia, the affinities with the
latter being particularly evident in the form of the larval antennae and prolegs. Nielsen concluded
that the subfamily Hydroptilinae sensu Nielsen (see Table 1) was closest to the Integripalpia,
while the Orthotrichiinae sensu Nielsen retained the primitive features of the Rhyacophilidae.
Nielsen’s views were, however, based entirely on the external morphological features of the larvae
which, it should be noted, may be subject to functional modifications correlated with the case
building habit. His conclusions regarding the affinities of the Hydroptilinae and the Integripalpia
may thus have been influenced by the occurrence of superficially similar characters which have
resulted from parallel evolution within the two groups.
Milne & Milne (1939), on the basis of larval anatomy, behaviour and case-form, concluded this
of the family: ‘the Hydroptilidae is probably more closely related to the Glossosomatinae than
the Glossosomatinae to the Rhyacophilidae . . . Hydroptilid larvae being more specialized in the
direction of the differentiation of the ends of the case (feeding from one particular end, etc.) than
Glossosomatinids.’
The first serious attempt to construct a phylogenetic classification of the order Trichoptera was
made by Ross (1956) who based his conclusions on a comparative study of adult and immature
morphology and larval case making behaviour patterns. From this study Ross deduced a set of
primitive adult and larval characters and followed the development of these within each group, his
findings regarding the evolution of the order being summarized in Chart 1; in this chart the rela-
tionships of the Hydroptilidae have been emphasized.
According to Ross (1956), the ancestral caddis fly gave rise to two major evolutionary lines:
the ‘fixed retreat division’, where the larvae simply spin silken nets in which they live and trap
their food (the Philopotamidae and the Hydropsychidae—Psychomyiidae complex), and the ‘case-
maker division’, containing all of the remaining families. Within the latter group the most primi-
tive representatives are considered to be the free-living Rhyacophilidae which build stone-covered
silken pupal shelters. The early ancestors of the Rhyacophilidae gave rise to the primitive glos-
sosomatid-line, in which the larvae build transportable cases and which in turn gave rise to the
more advanced ‘saddle-case’ building Glossosomatidae and the ‘purse-case’ building Hydrop-
tilidae. The final step in the evolution of the case building habit was the adoption of the ‘tube-case’
form as seen in the higher limnephiloid families. Ross considers this major branch to have arisen
from the same ancestor as that which gave rise to the hydroptilid line and not from the ‘tube-case’
building hydroptiline tribe, the Neotrichiini, which would appear to have evolved directly from
an early ancestor of the Hydroptilinae. The Ptilocolepinae are considered to be an early offshoot
of the hydroptilid line in which many adult features of the more primitive Glossosomatidae have
been retained.
In conclusion it would thus seem that the Hydroptilidae may be regarded as a specialized but
early offshoot of the case-making branch of the Trichoptera. The family is included in the Rhya-
cophiloidea, along with the Rhyacophilidae and Glossosomatidae, the other main subdivision
being the Limnephiloidea which contains the Limnephilidae, Sericostomatidae and the Lepto-
ceridae. The Hydroptilidae share a common ancestry with the Glossosomatidae with which they
have both larval and adult affinities, the latter being most marked in the Ptilocolepinae which also
exhibit pupal features intermediate between the Glossosomatidae and the Hydroptilinae. The
Hydroptilidae is also regarded as being an early offshoot of the line leading to the Limnephiloidea.
General morphology
Adult
There is a marked difference between the general appearance of the two hydroptilid subfamilies.
140
J. E. MARSHALL
Psychomyiidae
net-spinners _— Hydropsychidae
me Philopotamidae
A
Annulipalpia
larvae. free-living
Integripalpia
Rhyacophilidae
saddle-cases
ae J Glossosomatidae
purse-cases
Ptilocolepinae |
Hydroptilidae
Hydroptilinae J
tube-cases
Limnephilidae
ae Sericostomatidae
poe deans Leptoceridae
Chart I The position of the Hydroptilidae within the order Trichoptera (after Ross, 1967).
ha
Daplous4sd oupAH
Dapio}iydooDAyy
baplo}iydeuwly
THE GENERA OF THE HYDROPTILIDAE 141
1mm
a
' ‘
Fig. 1 Hydroptila tineoides Dalman, ¢.
The Hydroptilinae have what is normally regarded as the typical hydroptilid form, being small
with narrow, pointed wings, long costal fringes, reduced venation and an overall pubescent ap-
pearance due to the wings and parts of the body being densely covered in setae (Fig. 1). The
Ptilocolepinae, however, look more like small rhyacophilids and glossosomatids, the wings being
relatively broad with rounded apices, short costal fringes, analmost full complement of veins (Fig.
5) and a granulose, rather than pubescent appearance due to the presence of sparsely scattered
short, unmodified macrotrichia more typical of the order Trichoptera.
The Ptilocolepinae tend to have a rather uniform dark brown to black appearance, reflecting
the basic colour of the body, whereas the coloration and markings of the Hydroptilinae are de-
pendent on the arrangement of the setae. These have the same effect, to a certain extent, as the
overlapping scales of Lepidoptera and hydroptilids are often mistaken for small moths at first
sight, as first noted by Eaton (1873). Hydroptilid setae are usually black, or brown, and white and
give the insects a mottled or ‘salt and pepper’ appearance although, in Agraylea for example, this
may be partly due to spotting of the wing membrane itself. Distinct metallic hues are seen in some
tropical genera and are often caused by the presence of localized groups of modified, scale-like
setae.
142 J. E. MARSHALL
The Hydroptilidae are described as being small to minute insects and are often termed the
‘micro-caddis’. The average forewing length (middle of prothorax to apex of wing) of the Ptilo-
colepinae is 5mm (4-6 mm), while that of the Hydroptilinae is 3-5 mm (1:2-6:0 mm). A full
account of adult hydroptilids will not be given here; however, those features of taxonomic
importance referred to in the systematic section will briefly be described below.
Head capsule
The number of dorsal ocelli varies from the basic complement of 3, to 2 and 0. Posteriorly there
is usually a pair of dorsal postoccipital lobes or warts which are somewhat variable in shape, and
in the males of the genus Hydroptila these are modified as hinged caps which conceal eversible
scent dispersing organs lodged in the rear of the head cavity (Fig. 114).
Thorax
The small, ring-like prothorax bears a pair of setigerous protuberances or ‘warts’ which, in the
Hydroptilidae, are characteristically set close together (Figs 53, 62, 83, 108, 114, etc.). The shape
of the meso- and metathoracic nota and the presence or absence of transverse sutures provide
excellent diagnostic characters at both the subfamily and generic levels; Fig. 2 shows the general
terminology employed here. The posterior mesothoracic katepisternal suture (Fig. 3, suture ‘k’)
is characteristically present in the Ptilocolepinae. Ross (1956) first noted the ‘general absence’ of
this suture in the Hydroptilinae (Fig. 3, inset), but I have found this to be a consistent feature of
the subfamily. An important feature of hydroptilid taxonomy is the spur formula which refers
to the number of tibial spines commonly present on each leg. The maximum number on any one
leg is four, consisting of two preapical and two apical spurs; the spur formula gives the number
of spines on the fore-, mid- and hindlegs respectively and, for example, in Stactobia (Fig. 4) is
given-as 1.2.4.
The general form of the wings has already been described, the more primitive condition being
seen in the Ptilocolepinae (Fig. 5). Here the venation is well developed and resembles that of
primitive rhyacophilids (vide Ross, 1956 : figs 154, 155) from which it differs in the subcosta (Sc)
of the forewing and, from rhyacophilids in general, in the fusion of various veins in the hindwing.
The Ptilocolepinae differ from the Hydroptilinae in possessing a distinct discofdal cell (dc),
separate M, and M, and a forked Cu, in the forewing (Fig. 5). The Hydroptilinae are noted for
their narrow pubescent wings, reduced venation and long costal fringes (Figs 1, 6-10). In the
larger genera, such as Ugandatrichia and Agraylea (Figs 6, 7), the wings are slightly broader and
less pointed apically, the costal fringes are short, the raised setae less dense and the neuration more
regular. However, the wings of most hydroptilines are long, slender and highly acuminate. As the
overall size of the insect decreases so the wings become reduced and the venation less distinct
while the costal fringes become longer to compensate for the loss in wing membrane area. The
most extreme examples of wing reduction are seen in Chrysotrichia and Neotrichia (Fig. 8)
(average forewing length 1-5 mm) where the wings are reduced to thin, ribbon-like strips and the
venation reduced essentially to the main longitudinal branches R, M and Cu. The compensatory
development of the costal fringes parallels the condition seen in other insect groups such as the
Thysanoptera, the Trichogrammatidae and Myrmaridae (Hymenoptera) and the Ptiliidae,
Clambidae, Corylophidae and some Staphylinidae (Coleoptera). Venational features of the Hy-
droptilinae do not provide reliable taxonomic characters because, at suchreduced dimensions, small
variations in crossveins and forks, for example (all of taxonomic importance in the families of
larger Trichoptera), are not constant in genera and species of the Hydroptilinae.
Abdomen
This consists of the usual eleven basic segments (X and XI being regarded as one) and the sclerites
of the posterior segments are modified to form the genitalia. In all hydroptilid genera examined
there is a pair of small sternal glands of unknown function which open anterolaterally on the
fifth abdominal sternite in both sexes. The external structures associated with these glands differ
markedly between the two subfamilies. The Ptilocolepinae again resemble the Rhyacophilidae
and Glossosomatidae in having a well-developed sternal ridge (Fig. 11) which, in Ptilocolepus,
THE GENERA OF THE HYDROPTILIDAE 143
tegula coxa
pronotum setal fringe
femur el i
\ = anterior notal process I
TB ~ jst axillary sclerite ( ax1)
SS pleural process
ax2
mesoscutum
tibia
mesoscutellum _1 ,
posterior notal
& 2 process II
ens > 3 ad. Fane A ax 3
See — ‘ ax4
postnotum~ y, oa
~~ lateral
trochanter
postnotum I ‘
metascutum \ ; p-imm_,
BOS, jugum
ce a ad ie 4
ee ee posterior notal
metascutellum 1. / f aroceas. Ur
preapical
/ lateral postnotum IL spurs
abdominal 2 foreleg
tergitel 0-Imm
midtibia
apical
hindtibia spurs
postnotum _Spiracle (sp) .
scutum (s) I toate pronotum
subalare (sa) I | sy 9 anepisternum (an) I
Ke : occipital condyle
abdominal
tergite (tA)2
abdominal : ‘
sternite: 2)——$——-. | 9 lateral cervical
a sclerite
katepisternum (k) I
epimeron (em) I es trochantin
iN “ Pleural process
meron (m) II ae
: 3
coxa (cx) E pre-episternum
furca
0.5mm SUCK mas eee
: =F compare
Figs 2-4 Adult thorax and legs. 2, Agraylea sp., thorax and wing base, dorsal view (ax, axillary
sclerite); 3, Ptilocolepinae, thorax, lateral view (inset, Hydroptilinae, mesopleuro-coxal sclerites);
4, Stactobia mclachlani Kimmins, legs and spur formula (1.2.4).
has a pair of long, slender, membranous filaments arising from its anterolateral corners. In the
Hydroptilinae there is simply a pair of shallow, subcircular pits in the anterolateral corner of the
fifth sternite, from which there also arises a pair of short, posteriorly directed, setate, finger-like
processes (Figs 12, 14). Tricholeiochiton fagesii (Guinard) is so far unique within the subfamily
since there is no process and the surface of the relatively broader pit is irregularly sculptured
(Fig. 13). Ithytrichia lamellaris (Eaton) (Fig. 14) appears to exhibit an intermediate condition with
its reduced process (but relatively elongate seta) and faintly sculptured pit.
The external genitalia of the Hydroptilidae provide very useful diagnostic characters at both
the generic and specific levels. Most species are described from the males only but it has been
144 J. E. MARSHALL
Figs 5-10 Wing venation. 5, Ptilocolepinae, Prilocolepus granulatus (Pictet). 6-10, Hydroptilinae,
(6) Ugandatrichia cyanotrichia (Kimmins); (7) Agraylea sp.; (8) Neotrichia anahua (Mosely); (9)
Abtrichia antennata Mosely, 3; (10) Costatrichia lodora Mosely, 3.
found that, if carefully prepared and studied, the female genitalia can also provide useful taxono-
mic and phylogenetic features. Very little is known of the comparative morphology of hydroptilid
genitalia and, as may be expected, authors vary considerably in their terminology of the various
structures involved. As a detailed account of hydroptilid genitalia morphology and nomenclature
cannot be given here, reference should be made to the generalized diagrams of male and female
hydroptilid genitalia (Figs 15-17) which show the terminology to be adopted in this account.
However, a short description of the basic features will be given below and a more detailed study of
the male genitalia of Hydroptila occulta (Eaton), Agraylea multipunctata Curtis and Orthotrichia
costalis (Curtis) is given by Nielsen (1970).
Male. Segment IX forms a distinct genital capsule with a membranous posterior depression
(genital chamber) from which arise segment X (the dorsal plate) and the ventral inferior appen-
dages, the aedeagus lying between these two structures. Internal apodemes may arise from the
lateral anterior margins (as in the Stactobiini, Figs 65-68) and various external processes may
project posteriorly from the anal margin (e.g. Hydroptila, Figs 115-117 and Orthotrichia, Fig.
135); the latter are termed the side pieces or the lateral or intermediate processes. Segment X is
represented by the tergite only and projects from the dorsal posterior margin of segment IX; it is
usually entirely membranous, or sometimes weakly sclerotized, and its size and shape vary con-
siderably between taxa. The ventral face of tergite X may fuse with structures ventral to the aedea-
gus to form a distinct phallic tube, or phallocrypt.
THE GENERA OF THE HYDROPTILIDAE 145
spiracle lateral filament ; lateral
* lateral pit process
11
Figs 11-12 Abdomen and sternal gland, lateral view. 11, Ptilocolepinae (Prilocolepus
granulatus (Pictet)); 12, Hydroptilinae, generalized.
ee
sterniteY ———— __~_ lateral process
t= —
sternite V
Figs 13-14 Sternal gland, abdominal sternite V, antero-dorsal corner, lateral view.
13, Tricholeiochiton fagesii (Guinard); 14, Ithytrichia lamellaris Eaton.
There has been much confusion over the use of the terms ‘superior’, ‘intermediate’ and ‘inferior’
appendages. True superior appendages, as found in the majority of Trichoptera, are considered by
Nielsen (1970) to be absent in the Hydroptilidae; when used in context with this family the term
usually refers to a secondarily developed structure. The term ‘intermediate’ has been variously
applied to what has also been referred to as ‘superior’ and also to what are here called the ‘sub-
genital appendages’. The latter is a non-committal term used to refer to any structure ventral to
the aedeagus; when paired they are called appendages (=‘lateral penis-sheaths’, ‘parameres’,
‘intermediate appendages’) and when fused the structure is termed the ‘subgenital plate’ (= ‘lower
penis cover’, ‘ventral plate of X’).
The inferior appendages (gonopods), or ‘claspers’ as they are usually termed, are one-segmented
in the Hydroptilidae as opposed to the two-segmented condition normally found in Trichoptera.
They vary considerably in shape, size, additional processes and setae, and provide very useful
diagnostic characters at both generic and specific levels (see Figs 55-57, 65-68, 84, 94, 101, 109,
115-117, 122, 129,.131, 135, 140). Ross (1948) introduced the term ‘bracteole’ for the structure
arising from the base of the clasper (Fig. 101), while ‘bilobed process’ is introduced here to refer
to the delicate membranous processes which are seen in a number of genera of the Neotrichiini,
Hydroptilini and Orthotrichiini (e.g. Figs 16, 122, 129, 135, Bp). The bilobed process usually lies
just ventral to the subgenital plate and, although its form is somewhat variable, it consists basically
of a pair of short, posteriorly directed processes, each with an apical seta. In Orthotrichia (Fig.
135, Bp) they may be represented by the single median process formed by the fusion of the paired
organs, while in Oxyethira (Fig. 122, Bp) two pairs of processes would appear to be present which
apparently arise from the base of the subgenital plate. It is obvious that much more research into
the homologies of hydroptilid genitalia is required.
146 J. E. MARSHALL
internal
apparatus
(trident)
16
Figs 15-17 Hydroptilid genitalia, generalized. 15, 3, lateral view; 16, 3, ventral view; 17, 9,
ventral view. (Ae, aedeagus; Bp, bilobed process; Ja, inferior appendage; Sg, subgenital process.)
The copulatory organ (‘penis’, ‘phallus’) will simply be referred to here as the aedeagus (Figs
15, 16, Ae) as I have made no attempt to homologize the various structures that occur in the
Hydroptilidae. The aedeagus is thus basically a long, slender sclerotized tube which, in the Neo-
trichiini, Hydroptilini and Orthotrichiini (Figs 101, 109, 122, 135, 140, Ae) is distinctly divided
into two regions, the proximal, muscular-walled ejaculatory duct and the slender, distal intro-
mittent organ, and also bears a single, spiral ‘titillator’. In the other hydroptilid groups where the
aedeagus is apparently undivided, the homologies are unknown; each group would appear to
have its own specialized structure. In the Stactobiini there is basically a common median duct
with a pair of lateral processes which may be variously fused (Figs 69-73). The Leucotrichiini
have a very complex structure with ‘basal loops’ and ‘windows’ and a membranous apex (Fig. 84)
while the Ochrotrichiini have an aedeagus which may be very slender (Fig. 94) or heavily spined
(Fig. 96). Again, research into the homologies of the hydroptilid aedeagus could prove very re-
warding both taxonomically and phylogenetically.
Female. In the Hydroptilidae the female genitalia (Fig. 17) are of the generalized trichopteran
condition which is of the simple ‘telescopic ovipositor’ or ‘oviscapt’ form (Scudder, 1971). This
essentially involves modifications of segments VIII-X and the gonopore is situated ventrally
between segments IX and X or, when the sternite IX is reduced as in Oxyethira (Fig. 121),
apparently just behind segment VIII.
The posterior margin of the ring-like segment VIII bears a number of useful characters such as
rows of setae and dorsal and ventral processes and excisions (see Marshall, 1978 for figures of the
female genitalia of the British Hydroptilidae). The anterior margin may bear a pair of slender
lateral apodemes (Figs 17, 82, 121, 134) and the sternite may have patches of modified setae or, as
in Hydroptila occulta (Eaton) (Marshall, 1977; 1978), a characteristically shaped sclerite termed
the ‘ventral plate’ by Nielsen (1970) (Fig. 17). The long, slender segment IX forms a retractile
‘oviscapt’ (Scudder, 1971) or ‘false ovipositor’ (Nielsen, 1970) which is essentially membranous and
has a pair of long, rod-like apodemal sclerites to which the retractile muscles are probably
THE GENERA OF THE HYDROPTILIDAE 147
attached (Fig. 17). Segment IX often bears an anterior ventral sclerite, termed by Nielsen (1970)
the ‘dorsal plate’ (Fig. 17), and the distal end of the segment forms the lower lip of the genital
opening. Segment X is greatly reduced and forms the fleshy upper lip of the gonopore; it also
bears a pair of short, one-segmented cerci.
In preparations of female abdomens the form of the posterior, partly sclerotized genital ducts
(‘internal apparatus’, Nielsen, 1970) may be more or less discernible, the most noticeable struc-
tures being the sclerites of the vaginal wall (Fig. 17). These are what Mosely (1939a) termed the
‘trident-like’ structures and they are often erroneously referred to as the ‘bursa’ or ‘bursa copula-
trix’ which are terms indiscriminately used to refer to almost any part of the internal female
reproductive system visible in preparations. Such vaginal sclerites, however, along with the dorsal
and ventral plates and the posterior margin of segment VIII provide useful specific characters
which, I believe, could be put to more use in the future.
The generalized oviscapt form appears to have been modified in certain genera such as Oxyethira
(Fig. 121) and Orthotrichia (Fig. 134) where the genitalia are reduced and specialized in that they
are non-retractile and therefore do not have an oviscapt function. Both forms are described in the
systematic account and it is possible that their modifications are associated with the oviposition
habits of the females which have not, however, been investigated here.
Larva
The family Hydroptilidae is perhaps the most diverse of the Trichoptera with regard to the form
of the larvae and their cases since they are structurally adapted to suit a wide range of aquatic
environments. The members of the family do, however, share a number of basic morphological
and behavioural features which distinguish them from all other Trichoptera and have led to the
inclusion of the Ptilocolepinae within the group. Nielsen’s (1948) classic work on hydroptilid
larvae unfortunately tends to give a restricted view of the family since it deals with only five of the
more advanced genera, namely Agraylea, Hydroptila, Oxyethira, Orthotrichia and Ithytrichia
which are all specialized members of the Hydroptilinae. However, Nielsen (1948) does provide
an excellent foundation for all studies, whether morphological, biological or behavioural, on
the Hydroptilidae and without it the following more general account would have been almost
impossible to conceive.
The following account has been based on the literature and on the examination of material in
the BMNH or loaned from various institutions. The genera which are known in their larval stages
are given below (brackets signify a proposed synonym of the preceding genus; a single asterisk, *,
indicates material examined; two asterisks, **, denote a previously undescribed genus).
Ptilocolepinae: Ptilocolepus, Palaeagapetus*. Hydroptilinae: Stactobia*, (Lamonganotrichia),
Plethus, Stactobiella*, Catoxyethira, Leucotrichia*, Zumatrichia, Peltopsyche** (previously
known from case only), Anchitrichia, Alisotrichia, Neotrichia*, Mayatrichia*, Agraylea*, Allo-
trichia, Ugandatrichia, Hydroptila*, (Oeceotrichia, Pasirotrichia, Sumatranotrichia), Oxyethira*,
Paroxyethira, Xuthotrichia, Tricholeiochiton*, Ithytrichia*, (Saranganotrichia), Orthotrichia*,
(Javanotrichia, Baliotrichia, Orthotrichiella), Dibusa*, Caledonotrichia**, Dicaminus (case only).
A characteristic feature of the family is that the larvae undergo a simple form of hypermeta-
morphosis in which instars I-IV (‘young larvae’) are minute, free-living, caseless and of very short
duration, while the fifth instar (‘old larva’) builds a portable or secondarily fixed case and is the
principal feeding and growing stage of the life-cycle. The following account of the general exter-
nal appearance of hydroptilid larvae refers only to the fifth instar; the earlier instars will be dis-
cussed later.
General appearance
Despite their case building habits, hydroptilid larvae are prognathous and campodeiform as are
the more primitive, free-living and ‘saddle-case’ bearing groups in comparison with the hypo-
gnathous, eruciform, ‘tube-case’ building limnephiloid groups.
The Hydroptilidae may be distinguished from all other caddis larvae by a combination of the
following characters: small size (2-0—-7-0 mm body length), enlarged abdomen, three pairs of well-
developed thoracic tergites, the absence of segmentally arranged tracheal gills and the fusion of the
148 J. E. MARSHALL
labial pal
labrum mandible Be Gi maxillary
prementum : we palp
anteclypeus
postmental
sclerites
anterior
ventral apotome
4 Lanterior
4 tentorial
dorsal pit
apotome
lateral
stemmata
ventral 4
S F ecdysial 4 parietal
dorsal ue 4 ea eae
ecdysial line 3
it _—+# Parietal
4 sclerite
posterior
ventral
18 ce opto me 9
a,
Figs 18-19 Larva, generalized head. 18, dorsal view; 19, ventral view.
(Terminology after Hinton, 1963.)
abdominal prolegs to the sides of segment X. The number of abdominal tergites is variable but
there is always one on segment IX (the ‘anal plate’) and in the Hydroptilinae there is usually a
pair of small sclerotized (sometimes fused) ‘rings’ on the terga of segments II/III to VII/VIII.
A detailed account of the external morphology of hydroptilid larvae will not be given here; for
this, reference should be made to Nielsen (1948). However, features of taxonomic importance or
over which there has been some controversy regarding nomenclature will be briefly mentioned
below.
Head capsule
There has been some confusion over the homologies of the various regions of the head capsule of
trichopteran and endopterygote larvae in general, especially with respect to the ventral sclerites
Figs 20-28 Ventral apotomes and postmental sclerites, larva. 20, Palaeagapetus celsus Ross; 21,
Agraylea sexmaculata Curtis; 22, A. multipunctata Curtis; 23, Tricholeiochiton fagesii (Guinard);
24, Ithytrichia lamellaris Eaton; 25, Stactobia caspersi Ulmer; 26, Hydroptila occulta (Eaton);
27, Oxyethira flavicornis (Pictet); 28, Orthotrichia costalis (Curtis). (ava, pva, anterior and posterior
ventral apotomes; par, parietal sclerites; pms, postmental sclerites.)
THE GENERA OF THE HYDROPTILIDAE 149
penicillus
cutting teeth
\ penicillus
membranous
ventral bladd
condyle 31 e
scraping and scooping
Figs 29-32 Mandibles, larva. 29, Palaeagapetus celsus Ross (after Flint, 1962); 30, Stactobia
caspersi Ulmer (after Botosaneanu, 1956); 31, Hydroptila occulta (Eaton); 32, Orthotrichia
costalis (Curtis) (after Nielsen, 1948).
and the maxillolabial and hypopharyngeal complex. For simplification the terminology proposed
by Hinton (1963) has been adopted here and is illustrated in Figs 18, 19. The variety of ventral
apotomes and postmental sclerites found within the family is shown in Figs 20-28. The mouth-
parts are of the mandibulate type and were probably basically suited for biting through vegetable
material as in the Ptilocolepinae (Fig. 29) but, within the Hydroptilinae, two main lines of feeding
adaptations may be traced. In the Stactobiini and Leucotrichiini the larvae are primarily detritus-
feeding and the mandibles tend to have a form suited to scraping the substrate and scooping up
diatoms and other small organic particles (e.g. Fig. 30). In the Hydroptilini and Orthotrichia,
however, the larvae feed on the fluids of green filamentous algae and the mandibles are adapted
for biting or piercing a hole in the cell wall (Figs 31, 32). The mandibles are basically short and
robust and one or both may bear a tuft of hair-like spinules (the ‘penicillus’) in the median con-
cavity and which, as in Hydroptila occulta (Eaton), for example (Fig. 31), may be replaced in one
mandible by a small, membranous ‘bladder’. The form of the mandibles may provide useful
generic or specific characters but it should always be borne in mind that such structures may be
worn down with age and hence lose some of their external sculpturing.
Thorax
Well-developed tergites are present on all three thoracic segments and each has a distinct mesal
suture which, in the prothorax, probably serves as an ecdysial line (Fig. 33). Pleurites are present
on all three segments and essentially consist of the fused episternite (es) and epimeron (aem, oem)
(collectively termed the ‘pleurite’) separated by the raised pleural ridge (p/r) which continues
ventrally as a short pleural process and articulates with the antero-lateral coxal process (vide Fig.
34). A small anterior sclerite derived from the episternite (the preepisternite) may also be present.
The various forms of the pleurites found in hydroptilid larvae are shown in Figs 34-37. The
thoracic sternites are shown in Figs 38-46.
The thoracic legs of fifth instar hydroptilid larvae are basically of the ambulatorial type which
may be modified as short, robust clinging organs in torrenticolous forms (e.g. Fig. 85) or as long,
slender appendages in vegetation-dwellers (Figs 123, 132). The general form is shown in Fig. 33,
150 J. E. MARSHALL
tergite Anterior
tibial
process
12a
Figs 33-46 Thorax, larva. 33, thorax and legs, Hydroptila sp., lateral view. 34-37, thoracic pleurites,
lateral view, (34) Palaeagapetus celsus Ross; (35) Agraylea multipunctata Curtis; (36) Oxyethira
flavicornis (Pictet); (37) Dibusa angata Ross (aem, analepimeron; cx, coxa; es, episternite; oem, oral
epimeron; p/, pleurite; p/p, pleural process; p/r, pleural ridge; pes, preepisternite.) 38-46, thoracic
sternites; (38) Stactobia caspersi Ulmer; (39) Hydroptila occulta (Eaton); (40) Agraylea multi-
punctata Curtis; (41) Oxyethira flavicornis (Pictet); (42) Ithytrichia lamellaris Eaton; (43) Plethus
sp.; (44) Ochrotrichia sp.; (45) Caledonotrichia sp.; (46) Tricholeiochiton fagesii (Guinard).
the foreleg always being the shortest and sometimes adapted for special feeding or case-construc-
tion purposes. In Agraylea, Oxyethira, Paroxyethira and Hydroptila the foreleg (Fig. 47) possesses
a distinct modified seta-bearing process against which, according to Nielsen (1948), the tarsal
claw may be apposed thus forming a specialized chelate organ used in the manipulation of algal
filaments. In general, features of the legs of hydroptilid larvae often provide useful taxonomic
characters, especially at the generic level.
Abdomen
In fifth instar, case-bearing hydroptilid larvae, this is uniquely distended in shape and its overall
form is usually characteristic for each genus. There are ten distinct segments (XI regarded as
being fused with X) which are usually membranous except for the tergite of segment IX; segment
X always bears a pair of well-developed anal prolegs which are usually fused to the sides of the
segment in case-bearing larvae but project from the sides of the body in free-living forms andin the
‘tube-case’ building Neotrichiini (Figs 48, 49, cf. 50, 51, 92, 106). In Hydroptila and Ithytrichia
(Figs 48, 49, 118, 141) long filiform caudal filaments are present arising on segments IX and X.
Hydroptilinae larvae have dorsal sclerotized rings on abdominal segments II/III to VII/VIII
which seem to be a constant feature of the subfamily (Figs 74, 97, 110, 118, 123, 132, 136, 141,
THE GENERA OF THE HYDROPTILIDAE 151
articular rod
tibi isternit sclerite ‘b’
ia pre episternite pleural anal claw
ridge
episternite tergite IX
tarsus sclerite‘c’
) oral
: | RG epimeron % se
47
Figs 47-49 Larval thoracic leg and anal prolegs (fifth instar). 47, foreleg and pleurites, Hydroptila
sp. 48-49, anal prolegs, generalized hydroptilid, (48) dorsal view; (49) lateral view.
143, 147). Nielsen (1948) referred to them as ‘dorsal abdominal glands’ but they do not appear to
have a glandular structure internally and simply appear to be modified epithelial cells. According
to Wiggins (1977) they may be regions of specialized chloride epithelial cells adapted for ionic
absorption and osmoregulation.
Case
Final instar hydroptilid larvae construct distinctive cases of silk secretion which may be externally
sculptured or into which may be incorporated various organic or inorganic particles such as
bryophyte fragments, algal filaments and diatoms or small sand grains (which may be added to
provide ballast). Ross (1967) coined the term ‘purse-case’ to distinguish hydroptilid cases from
the ‘saddle-cases’ of the Glossosomatidae and the ‘tube-cases’ of the higher limnephiloid groups.
The “purse-case’ essentially consists of two silken ‘valves’ closely apposed and joined along the
lateral margins (seams) leaving slit-like anterior and posterior openings. It may be laterally (Figs
79, 98, 111, 119, 123, 128, 132, 145) or dorso-ventrally (Fig. 61) compressed and in the latter in-
stance may be temporarily or permanently fixed to the substrate (Figs 75, 76, 78, 81, 86, 148) or it
may have a more fusiform shape (Figs 99, 138). As the abdomen of the fifth instar larva increases
in size the larva enlarges the case by splitting the dorsal and ventral seams and adding new layers
of silk before closing them again (Nielsen, 1948). The Neotrichiini are distinct in having slightly
tapering ‘tube’-type cases (Figs 103, 107) with circular oral and anal apertures, thus superficially
resembling the higher Limnephiloidea. Alisotrichia (Leucotrichiini) and Ugandatrichia (Hydro-
ptilini) are distinct in that the final instar appears to have lost its case building habit and
consequently retains the general appearance of the free-living early instars. The cases of the
Hydroptilidae are very distinct for each genus and provide very useful key characters.
The ‘young larvae’: instars I-IV
Larval hypermetamorphosis is characteristic of the Hydroptilinae, although it has not yet been
observed in the Ptilocolepinae, and has so far been recorded in Agraylea, Hydroptila, Oxyethira
Ithytrichia, Orthotrichia (vide Nielsen, 1948); Stactobia (Botosaneanu, 1956; Danecker, 1961;
Lepneva, 1964); Ochrotrichia, Mayatrichia (Ross, 1944), Paroxyethira (Leader, 1970) and Ugan-
datrichia (Scott, 1976).
The young larvae are characterized by their relatively smaller size (0-5—2-7 mm, I-early V) and
features associated with the absence of a case. They thus have narrow posteriorly tapering abdo-
mens and long, freely projecting anal prolegs while the whole body is covered in long, fine setae
152 J. E. MARSHALL
anal claw
50
mandible
32
scierite ‘c’
sclerite ‘b’ articular rod
=)
y l
[ ey i reat i ( Ad thoracic
ie \ ‘i wingsheaths y: et
ae Uy
ee. y
. vp
So : ery
Sh” <GD <a
anterior
hook plate
; ay a posterior
A =< hook plate
ZF” a « NN : at
, A gh Ne : i
-: ~ y
Lee se
eee p—~% pharate papel
small abdomen aS elongate setae cuticle
JJ ate pc BS a Q oviscapt
0.1mm
ee
free anal
Prolegs
Figs 50-52 Early instars and pupa. 50, anal prolegs, early instar, dorsal view; 51, generalized early
instar, dorsal view; 52, pharate adult (‘pupa’), Hydroptila vectis Curtis, 2, dorsal view.
which give them the appearance of small, planktonic Crustacea (Fig. 51). The setae offer resistance
to sinking and aid the larvae in remaining suspended in the water and in swimming, these instars
being the dispersive stages of the larval life-history. In those genera which live in rapids and tor-
rents the larvae are more robust and dorso-ventrally flattened with short legs and few setae, thus
being adapted for moving over the surface of the substrate rather than swimming freely. A more
detailed account of the early instars is given by Nielsen (1948) who describes both their morphology
and their behaviour.
Pupa
The general appearance of the exarate, decticous pupae of most Trichoptera is very uniform
throughout the order and the cast pupal skins are difficult to identify beyond the family level.
Hydroptilid pupae may be recognized by their lack of any positive distinguishing structural fea-
tures which variously characterize the other families of Trichoptera. In addition to their relatively
small size (1-5-6-0 mm body length), the following features are useful in recognizing hydroptilid
pupae (Fig. 52): (i) absence of abdominal gills or lateral lines; (ii) presegmental dorsal abdominal
plates on segments III-VII, postsegmental plates on segments III-V; (iii) mandibles symmetrical,
short with one or two small median teeth. or long or short without teeth or with a fine, serrate
cutting edge; (iv) anal seta-bearing processes absent (these only occur in forms where the posterior
end of the pupal case is open to admit a water current, the setate processes serving to keep the
apertures clear of debris).
Finally, the pupal case generally resembles that of the larva except that the oral and anal aper-
tures are sealed and it is fixed to the substrate. Having attached and sealed the case, the larva then
spins the final internal lining and adopts the characteristic prepupal resting attitude in which,
according to Barnard (1971), the thorax becomes distended, the abdomen straightens and the
THE GENERA OF THE HYDROPTILIDAE 153
intersegmental grooves become less distinct. According to Nielsen (1948) there is a distinct anterior
and posterior differentiation of the larval case and this becomes reversed in the pupal stage. When
the pharate pupa is fully developed, ecdysis takes place and the larval exuviae are pushed to the
posterior end of the pupal case. The fully developed pharate adult cuts an exit hole in the pupal
case and wriggles out with the aid of the dorsal abdominal hook plates. It then makes its way to
the surface where it swims around erratically, back uppermost, until it encounters a support on
which it climbs out of the water and rests before the final ecdysis.
Systematics
Family HYDROPTILIDAE Stephens
Hydroptilidae Stephens, 1836 : 151. Type-genus: Hydroptila Dalman, 1819.
DISTRIBUTION. Cosmopolitan (excluding polar regions).
D1aGnosis. Adult (Fig. 1) Forewing length 1-2-6-0 mm; forewings moderately to densely pubescent; wings
broad with relatively complete venation in the more primitive Ptilocolepinae (Fig. 5) to narrow with
highly reduced venation and tapering apices in the more advanced Hydroptilinae (Figs 6-10); ocelli 3, 2
or 0, lateral pair set close to compound eyes some distance posterior to antennal bases; post-occipital
warts prominent, meeting postero-medially or represented by loosely hinged sclerities or ‘lobes’ (Fig. 114);
2 genitalia usually an oviscapt with an elongate telescopic segment IX and a small, cerci-bearing segment
X (Fig. 17) (IX rarely reduced and fixed (e.g. Fig. 121)); ¢ inferior appendages one-segmented, genitalia
otherwise very variable (generalized plan, Fig. 15).
Larva (e.g. Fig. 118). Campodeiform, prognathous; usually exhibiting hypermetamorphosis; dorsa of
all three thoracic segments with paired nota; abdominal gills and lateral line absent; anal prolegs usually
fused to sides of segment X in the fifth instar (free in instars I-IV); abdominal tergites present or absent on
segments I-VIII, always present on segment IX; dorsa of abdominal segments I—-VIII variously with small
cuticular rings which may be regions of chloride epithelia; abdomen usually greatly distended in fifth
instar; case basically of ‘purse-type’ (e.g. Fig. 119).
Pupa (Fig. 52). Mandibles usually short with one or two small median teeth (Ptilocolepinae) or with
fine serrations along inner edge only (Hydroptilinae): abdominal gills and lateral line absent; apex of
abdomen without lobes, processes or specialized cleaning appendages; presegmental hook plates present
dorsally on abdominal segments III-VII, postsegmental plates on III-V.
Keys to genera
Adult
1 Forewings broad with rounded apices, sparsely pubescent, fringes relatively short; venation
well developed, discoidal cell (dc) present, Cu, forked in forewing (Fig. 5); mesonotum con-
vex, prescutum distinct, scutellum subtriangular with a straight posterior margin and a large
oval wart (Fig. 53); mesokatepisternal suture (Fig. 3, suture ‘k’) present; latero-ventral ridge
of abdominal sternite V present (with or without a pair of antero-dorsal filaments) (Fig. 11)
(Ptilocolepinae) , 2
— Forewings narrow, usually acuminate, ‘densely pubescent, fringes long (Fig. 1): venation
variable, usually reduced (Figs 6-10), forewing with Cu, unbranched; mesonotum flat,
prescutum absent, scutellum with posterior section forming a flat triangular area with steep
sides and a warty texture along edges only (e.g. Figs 62, 114); mesokatepisternal suture
absent (Fig. 3, inset); sternite of abdominal segment V with a pair of shallow antero-dorsal
pits, each with a short, setae-bearing digitate process (Figs 12-14) (Hydroptilinae) . : 3
2 Veins R., R;, M,, M, free in the hindwing; spur formula 2.4.4 (North America; Baltic Amber)
PALAEAGAPETUS (p. 160)
— Veins R,+ R;, M,+M, fused in the hindwing (Fig. 5); spur formula 1.3.4 (Palaearctic)
PTILOCOLEPUS (p. 160)
4
3 Transverse suture present on mesoscutellum (Figs 62, 83, 93) ; : : ;
— Transverse suture absent on mesoscutellum (e.g. Figs 108, 114, 120) ‘ ; 20
4 Metascutellum subpentagonal to triangular (Figs 83, 93) (Nearctic and Neotropical) . 3 10
Metascutellum subrectangular to short pentagonal (Palaearctic) or as wide as scutum, very
short and rectanguloid (Figs 62-64) (Holarctic) : g : i , i ; 5
154
Nil wn
J. E. MARSHALL
Metascutellum as wide as scutum, short and subrectangular (Fig. 62) : : ; : 6
Metascutellum not as wide as scutum (Figs 63, 64) ‘ ; r ; ; : ; 8
Spur formula 0.2.4
(Oriental) CHRYSOTRICHIA (p. 170) (Neotropical: Antilles) BREDINIA (p. ni
Spur formula 1.3.4
3 genitalia simple, segment VIII unmodified (Fig. 65) (Holarctic) , STACT OBIELLA (p. a
3 genitalia very complex, sternite VIII elongate with posterior dorso-lateral spines, segment
IX much reduced and withdrawn into VIII (Fig. 68) (Africa) . CATOXYETHIRA (p. 171)
Spur formula 0.2.3 (SE. Asia) : : - ‘ j : ; ; PLETHUS (p. na
Spur formula 1.2.4 ‘
Fore-tibial spur reduced to a small process; postoccipital lobes narrow, elongate; metascu-
tellum wide, short, rectangular to pentagonal (Fig. 64); ¢ genitalia very complex; 9 internal
apparatus with distinct loop in anterior duct (Fig. 82) (Africa; SE. Asia ?)
MADIOXYETHIRA (p, 173) (possibly including PSEUDOXYETHIRA (p. 174) and
SCELOTRICHIA (p. 174))
Fore-tibial spur unmodified; postoccipital lobes broad, hemispherical; metascutellum narrow,
long, rectangular (Fig. 63); ¢ genitalia relatively simple, sternite VIII produced postero-
ventrally, IX reduced, VII with elongate process (Fig. 66); 2 internal apparatus without
anterior loop (Palaearctic) ; ; STACTOBIA (p. 165)
3 inferior appendages large and distinct, projecting well beyond ‘segment IX, often armed with
various spines and processes (Figs 94-96); head and antennae always simple (Fig. 93) _. 11
3 inferior appendages usually short, hardly projecting beyond segment IX (Fig. 84) but if
narrow and elongate with elaborate baso-dorsal processes then head and antennae (3)
modified (Figs 90, 91) : : : ’ ; 13
Fore-tibia without apical spur ; ; F OCHROT. RICHIA (OCHROT. RICHIA) = 186)
Fore-tibia with apical spur . , 12
3g segment IX produced posteriorly as broad dorso- lateral lobes; inferior appendages narrow,
elongate, projecting ventrally just beyond lobes of IX . , . RHYACOPSYCHE (p. 187)
3 segment IX without lateral lobes, inferior appendages broad and elongate, projecting well
beyond IX (Fig. 94) é : F OCHROTRICHIA (METRICHIA) (p. 186)
Spur formula 0.3.4; 0.2.4; 0.2.3; 3 seca with median constriction only; @ tergite VII
modified, internal apparatus simple, ring-like . . ALISOTRICHIA (p. 183)
Spur formula 1.3.4; ¢ aedeagus with midlength complex (Fig. 84); Q pees VII unmodified,
internal apparatus well developed (Leucotrichia-group) : s ; ; 14
Basal costal pouch present on ¢ forewing (Figs 9,10) . ; : F d ; : 15
Basal costal pouch absent on ¢ forewing : 17
Basal costal pouch large, almost half length of forewing (Fig. 9): head and basal antennal
segment (3) complex (Fig. 91); ocelli 2 (3) ABTRICHIA (p. 183), Betrichia bispinosa (p. 182)
Basal costal ‘bulla’ small (Fig. 10); head and basal antennal segment (3) unmodified; ocelli 3 16
Middle antennal segments (3) wide and flat (Fig. 90). COSTATRICHIA (lodora-group) (p. 181)
Middle antennal segments (¢) unmodified . . ACOSTATRICHIA (p. 182)
Basal segment of g antennaenlarged . PELT OPS YCHE AB: 179), ZUMATRICHIA (p. 179)
Basal segment of ¢ antenna unmodified : : : : ‘ 18
Ocelli 3 (3, 2)
COSTATRICHIA (simplex-group) (p. 181), Betrichia argentinica (p. 182),
LEUCOTRICHIA (melleopicta-group) me se and CELAENOTRICHIA (p. 183)
Ocelli 2 (3) : : : 19
3 abdominal sternite VIII with postero- -lateral processes ; : ANCHITRICHIA (p. 181)
3g abdominal sternite VIII unmodified
LEUCOTRICHIA : tage aed a 2 and BETRICHIA easiits (p. 182)
Ocelli absent ‘ ‘ : 21
Ocelli present. ; ‘ : 23
Fore-tibia with apical spur; poe élliptical. vovoid: pasha — lorewink length 5-5 mm)
(Nearctic) , , : ; : ; DIBUSA (p. 218)
Fore-tibia without apical spurt wing Bounties , j ; Ze
Metascutellum subrectangular (Fig. 133); spur formula 0. Ef 4 3 eehitalia markedly asym-
metrical (Fig. 135); 2 genitalia with segment IX relatively short (Fig. 134)
ORTHOTRICHIA (p. 213)
THE GENERA OF THE HYDROPTILIDAE
Metascutellum pentagonal to triangular; spur formula 0.2.4; 3g genitalia symmetrical;
segment IX of 2 long, forming an oviscapt; postoccipital lobes of ¢ modified as hinged caps
concealing eversible scent organs : , : ; : HYDROPTILA (p.
Mid-tibia with no preapical spur (0.2.3, 0.2. 4)
Mid-tibia with preapical spur (1.3.4, 0.3.4) é : ; ; 2 A :
Spur formula 0.2.3 (Nearctic) é d ; : 7 : P NEOTRICHIA (p.
Spur formula 0.2.4
3 segment VIII fused, annular; segment IX almost ‘entirely concealed within VIII; forewing
length c. 1-6 mm (SE. Asia) F , : : : : STENOX YETHIRA (p.
$ genitalia not as above
3 segment IX with prominent postero- -lateral processes and broad apices: inferior appendages
with broad setae-bearing posterior margin and small digitate dorso-lateral basal projection
(Nearctic, Neotropical) . : MAYATRICHIA (p.
3 segment IX without processes, inferior appendages apparently absent (Australia)
ORPHNINOTRICHIA (p.
Fore-tibia with apical spur ned Chile). : . . NOTHOTRICHIA (p.
Fore-tibia without apical spur ;
Mesoscutellum diamond shaped, narrow (Figs 108, 112, 113)
Mesoscutellum with anterior margin evenly convex, wide (Figs 120, 126, 130, 139)
$ ventral process of sternite VII short, laterally compressed, triangular; inferior appendages
large, elongate, often broad
3 ventral process of sternite VII usually long and spatulate; inferior appendages short and
broad, sometimes with short postero-median processes
Lateral filaments of abdominal segment II present in ¢ (India, SE. Asia)
UGANDATRICHIA (MOSELYELLA) (p.
Lateral filaments absent (Africa) . F . UGANDATRICHIA (UGANDATRICHIA) (p.
$ inferior appendages with concave posterior margins; subgenital plate with a pair of asym-
metrical processes bearing asymmetrical spines or filamentous appendages (Fig. 109)
ALLOTRICHIA (p.
3 inferior appendages with convex posterior margins (sexmaculata-group) or very short with
postero-median processes (multipunctata-group); processes of subgenital plate absent
AGRAYLEA (p.
$ abdominal segment VIII fused, annular; segment IX mostly concealed within VIII (Fig. 122);
2 genitalia not oviscapt, segment IX reduced, tergite usually fused with that of VIII (Fig.
121) : : d OXYETHIRA (p.
3 segment VIII with tergite and sternite distinct; 2 genitalia usually oviscapt
$ genitalia very distinctive, sclerities VII and VIII reduced to thin strips, sternite IX enlarged,
tergite X forming an elongate narrow plate, inferior appendages short, slender with in-
curved apices (Sri Lanka) ; ’ ‘ , : : MACROSTACTOBIA (p.
3 genitalia not as above
3 segment IX with deep dorsal U-shaped i incision revealing aedeagus ii in distinct sheath usually
with subapical processes; inferior appendages elongate, concave, broad; ventral processes
on segments VII and VIII, latter long and spatulate (Fig. 127) (New Zealand)
PAROXYETHIRA (p.
3 genitalia not as above :
3 segment IX produced postero-ventrally (obliquely truncate in 1 lateral view); genital appen-
dages concealed within IX, difficult to homologize (Fig. 131) (NW. Palaearctic, SE. Asia)
TRICHOLEIOCHITON (p.
$ genitalia not as above
3 sternite VI with ventral process: sides of segment IX tapering postero- -medially; inferior
appendages elongate, parallel, arising near antero-ventral margin of IX, scarcely projecting
posteriorly beyond IX (Fig. 140) ; ; ; : : ; ITHYTRICHIA (p.
3 genitalia not as above
$ inferior appendages apparently arising dorso- posteriorly from segment Ix, ‘bean-shaped’
with distinct ventrally directed processes; aedeagus slender, tapering (New Caledonia)
CALEDONOTRICHIA (p.
3 genitalia not as above ,
$ sternite VII with long, slender process: segment IX with broad postero-lateral truncate
side-pieces; aedeagus with distinct apical or sub-apical hook (Fig. 129) (Australia)
XUTHOTRICHIA (p.
155
200)
24
Zi
189)
25
207)
26
191)
220)
219)
28
29
a2
30
31
198)
198)
196)
193)
203)
33
217)
34
208)
35
210)
36
216)
a7
221)
38
209)
156 J. E. MARSHALL
— genitalia not as above : 39
39 J segment IX with postero-lateral side-pieces produced as ; broad, truncate lobes, segment 1X
reduced dorsally to thin median transverse strip; aedeagus truncate with well-developed
spiral ‘titillator’ (Yemen, Africa, Zaire) : DHATRICHIA (p. 199)
— segment IX relatively complete dorsally; aedeagus slender with pointed apex or short, spiral
‘titillator’ (Europe, Pakistan, Aden) . : ; : : ‘ MICROPTILA (p. 197)
Larva
1 Small tergites present on abdominal segments I—VIII; larval case dorso-ventrally flattened,
fixed to substrate or absent :
Segments I-VIII without distinct tergites; larval case ‘variable but usually not fixed to substrate
WNAON
2 Median sutures of meso- and metathoracic nota indistinct (Nearctic, Neotropical)
— Median sutures of all three thoracic nota distinct ¢ ;
3. Fifth instar larva free-living, without case; pupal case sac-like (Africa)
UGANDATRICHIA (p. 198)
— Fifth instar larva with distinct case 4
Posterior margin of fourth tergite appearing ‘crenellated’; case dorso-ventrally flattened,
barrel-shaped with mid-dorsal and ventral longitudinal seams (Figs 74, ie STACTOBIA (p. 165)
— Tergite IX of larva and case not as above... 5
5 Case rectangular in dorsal view (Fig. 78) with mid- dorsal and ventral seams; larva (Fig. 71)
(SE. Asia) : : PLETAUS (p. 168)
— Case tapering anteriorly and posteriorly (Fig. 81) without dorsal and ventral seams (Africa)
CATOXYETHIRA (p. 171)
6 Fifth instar larva free-living, without case; abdomen slender, tapering posteriorly; anal prolegs
projecting from sides of abdominal segment X (Fig. 92) : ; ALISOTRICHIA (p. 183)
— Fifth instar in a fixed, oval-circular case, abdomen enlarged, abruptly widened from segments
V-VII; anal prolegs fused to sides of segment X 4 : 7
fi Anterolateral angle of pronotum produced anteriorly; femora with arborescent setae
ANCHITRICHIA (p. 181)
— Anterolateral angle of pronotum not produced; arborescent setae absent : : : 8
8 Tergite IX with short, stout scattered setae (vide Figs 87-89)
ZUMATRICHIA (p. 179), ABTRICHIA (p. 183), PELTOPSYCHE (p. 179)
— Tergite IX usually without short, stout setae but if present then in a transverse band (vide
Figs 85, 86) . ; ; : . LEUCOTRICHIA Ne 178)
9 Mid- and hindlegs slender, much longer than forelegs ; d P ; 10
— Thoracic legs short, subequal in length : ; : ; : E : ; : 12
10 Case ‘bottle’- or ‘jug’-shaped (Figs 123, 124) : é : ; OXYETHIRA (p. 203)
— Long edges of case almost parallel . ; : F : F P ; 11
11 Mid- and hindlegs greater than 4 = forelegs (Fig. 132) , . TRICHOLEIOCHITON (p. 210)
— Mid- and hindlegs approximately 2:5 x forelegs (Fig. 128) (New Zealand)
PAROXYETHIRA (p. 208)
12 Abdominal segments with dorsal and ventral ‘bud-like’ processes; a single filamentous caudal
gill present (Fig. 141) : ’ : ITHYTRICHIA (p. 216)
— Abdominal segments not as above, caudal filaments usually absent : , 13
13. Abdomen more or less cylindrical; head tapering anteriorly; anal prolegs not fused to sides of
segment X; case cylindrical, tapering posteriorly (Nearctic) . : 14
— Abdomen flattened dorso-ventrally or -laterally; head not tapering anteriorly; anal prolegs
fused to sides of segment X; case flattened with slit-like anterior and posterior openings . 15
14 Abdomen slightly depressed, intersegmental grooves sede aa case of secretion and fine sand
grains (Figs 102-104). ; ; ; .NEOTRICHIA (p. 189)
— Abdomen not depressed, intersegmental grooves indistinct; case of secretion only, surface
sculptured usually with transverse or longitudinal ridges (Figs 105-107) MA YATRICHIA (p. 191)
15 Labrum with asymmetrical ‘beak’ (Fig. 137); abdominal segment II with lateral humps (Fig.
136); case fusiform, of secretion only with longitudinal ridges (Fig. 138)
ORTHOTRICHIA (p. 213)
— Labrum, abdomen and case not as above : : : , ‘ : ‘ ‘ 16
16 Tibia of foreleg with pronounced baso-ventral process ' : , : ; : , 17
— Tibia of foreleg without baso-ventral process ; 20
17 Abdomen with three filamentous caudal gills; case laterally compressed, of two silken valves
covered in fine sand grains and diatoms (Figs 118, 119) : : HYDROPTILA (p. 200)
THE GENERA OF THE HYDROPTILIDAE 157
— Caudal gills absent. 18
18 Ventral intersegmental grooves of abdomen very pronounced; case of two silken valves
covered in radially arranged algal filaments (Figs 110, 111)
AGRAYLEA (p. 193), ALLOTRICHIA ? (p. 196)
— Abdomen and case not as above, exclusively ‘New World’ : 19
19 Tarsus with large, plate-like seta and hook claw; antero-lateral expansion of metanotum
absent; case fusiform, of secretion only (Figs 99, 100) (Central America)
RHYACOPSYCHE (p. 187)
— Tarsal seta not plate-like; metanotum with antero-lateral expansion; case usually laterally
compressed, of two silken valves covered in fine sand grains, sometimes carried horizontally
(dorsal valve vaulted, ventral face flat, of secretion only) (Figs 97, 98)
OCHROTRICHIA (p. 185)
20 Abdomen and case compressed dorso-ventrally ; 21
— Abdomen and case not compressed dorso-ventrally; tarsal claws stout, abruptly ‘curved with
thick blunt spur at base ' 23
21 Case of secretion only with a central tubular section ‘surrounded by lateral fringe, ‘former with
a pair of small dorsal pores (Fig. 148; larva, Fig. 147) (New Caledonia)
CALEDONOTRICHIA (p. 221)
— Case of two silken valves covered in Eee oe of moss and liverwort (Holarctic, montane
regions (Ptilocolepinae)) . 22
22 Abdomen with a pair of lateral tubercules on 1 each segment; abdominal sesmient I without
small tergite (Figs 60, 61) (Holarctic) , ; PALAEAGAPETUS (p. 160)
— Abdomen without lateral tubercules; abdominal segment I with small dorsal tergite
(Palaearctic) . 5 — PTILOCOLEPUS (p. 160)
23 Dorsal abdominal setae stout, dorsal rings of abdominal segments distinct; larva feeding on red
algae which are incorporated into silken valves of purse-like case; tarsal claw spurs short,
thick and blunt (Figs 143-145) (eastern U.S.A.) . ‘ : DIBUSA (p. 218)
— Abdominal setae slender, dorsal ridges indistinct; case of two symmetrical silken valves;
tarsal claw spurs elongate, thick, blunt (Figs 79, 80) (Holarctic) . STACTOBIELLA (p. 169)
Subfamily PTILOCOLEPINAE Martynov
Ptilocolepinae Martynov, 1913a : 22. Type-genus: Prilocolepus Kolenati.
DISTRIBUTION. Holarctic.
The Ptilocolepinae contains two small montane genera which are confined to the Holarctic
region. Ptilocolepus, containing four extant Palaearctic species, and Palaeagapetus which is known
from the type-species in Baltic Amber and three extant Nearctic species. The adults bear more
resemblance to small glossosomatids with which they were once originally classified until dis-
covery of their larval stages indicated that their true affinities were with the Hydroptilidae (Ross,
1956). Ptilocolepus granulatus (Pictet) was originally described in Rhyacophila but, as Pictet (1834)
himself noted, could be distinguished from all other members of the latter genus by its ‘rough
wings and very ciliated legs’. Rhyacophila granulata was later shown to be a senior synonym of
Ptilocolepus turbidus Kolenati by Hagen (1855), the type-species thus becoming Ptilocolepus
granulatus. Thienemann (1904) described the larva of Pt. granulatus and noted its hydroptilid
affinities which prompted Ulmer (1907) to place the genus in the Hydroptilidae. Subsequently,
however, Martynov (1913a) erected the Ptilocolepinae for the nominate genus but retained it in
the Rhyacophilidae. It was Ross (1956) who finally transferred the subfamily to the Hydroptilidae,
redefining it to include the genus Palaeagapetus (earlier in that paper, chart 2, p. 18, Ross refers
erroneously to the subfamily as the Palaeagapetinae), the inclusion of Palaeagapetus later being
justified by the discovery of the immature stages of Palaeagapetus celsus Ross, described by Flint
(1962) and shown to have many affinities with those of Prilocolepus granulatus.
The following subfamily diagnosis is based on the literature and on examination of adult
material of Prilocolepus granulatus and Palaeagapetus celsus in the BMNH collections and larvae
of P. celsus loaned from the USNM collection.
158 J. E. MARSHALL
Figs 53-54 Adult head and thorax, dorsal view. 53, Prilocolepus granulatus (Pictet);
54, Agapetus sp., Glossosomatidae.
DiaGnosis. Adult. Forewing length 4-6 mm; general body colour dark brown to black, wings somewhat
iridescent; wings (Fig. 5) broad with rounded apices, very sparsely pubescent; costal fringes short; vena-
tion almost complete, resembling glossosomatid and primitive rhyacophilid conditions in that forewing
with discoidal cell (dc) closed, Cu, forked and Cu, curved: head (Fig. 53, cf. Agapetus, Fig. 54) strongly
sclerotized, z-shaped tentorium well developed; ocelli 3, lateral pair set close to compound eyes some
distance posterior to bases of antennae; antennae short, of approximately 24-30 segments; postoccipital
warts large, ovoid, meeting medially: pronotum (Fig. 53, cf. Fig. 54) with median warts close-set; mesono-
tum strongly convex, prescutum delineated by a row of punctate setal sockets; scutellum subtriangular,
convex with a straight posterior margin and a large oval wart; katepisternal suture (‘k’) present (Fig. 3):
metascutellum subtriangular: preapical spur absent from foretibia (spur formula 2.4.4 in Palaeagapetus
and 1.3.4 in Ptilocolepus): abdominal sternite V (Fig. 11) with a well-developed sternal ridge running from
anterior dorsal fold, posteriorly and ventrally to meet in midline: ventral processes variously present on
sternites VI and VII: 2 genitalia forming an oviscapt: ¢ genitalia simplified but very specialized within
each genus (Figs 55-59) with segment VIII unmodified ; segment LX fused, produced antero-ventrally with
strongly developed oblique lateral ridges, strongly excised dorsally and ventrally (latter excision con-
taining a triangular ventral plate in Prilocolepus above which lie the fused, generally bifid, ‘inferior
appendages’ (homology uncertain)); ‘inferior appendages’ apparently fused with segment IX in
Palaeagapetus; tergite X forming a simple dorsal plate (which in Ptilocolepus bears a pair of setose lobes);
aedeagus greatly simplified, short, broad and membranous with a dilated apex and a median ventral
sclerite.
Larva (Fig. 60). Slightly flattened dorso-ventrally: head with distinct dorsal ecdysial lines; anterior
ventral apotome large and triangular, postmental sclerites paired, posterior ventral apotome small and
triangular (Fig. 20); labrum symmetrical, anterior margin strongly emarginate with a row of many short
setae; mandibles (Fig. 29) not markedly symmetrical, short and stumpy, medial brush present in both
right and left, cutting edges strongly ridged: thoracic nota present on all three segments, longitudinal
median ecdysial lines distinct on each; thoracic sternites absent; legs short, subequal; preepisternite free
on all thoracic segments (Fig. 34); abdomen entirely membranous except for the well-developed tergites
of segments I and IX in Prilocolepus; anal prolegs fused to segment X; abdomen with prominent sucker-
like tubercules on segments II—VII in Palaeagapetus.
THE GENERA OF THE HYDROPTILIDAE 159
Case (Fig. 61). Purse-type, dorso-ventrally compressed (carried horizontally), constructed of two silken
valves covered with small fragments of liverwort and moss.
Pupa. Mandibles long with two medial teeth in Ptilocolepus, one in Palaeagapetus.
BIoLoGy. The larvae occur in small montane springs and seeps, amongst luxuriant growths of moss
and liverwort, on stones and other submerged objects; they are often found in the cold damp
vegetation above the water level. Habitats in cool, shaded, montane coniferous forests, which
favour the growth of bryophytes, are likely to support small colonies of these very local and primi-
tive caddis flies; indeed, McLachlan (1880) noted that the adults of Ptilocolepus granulatus were
found in European mountainous regions ‘on driblets on faces of rocks’ where water was soaked
up by the moss. The habitat preferences of the Ptilocolepinae are thus very similar to those of the
hypothetical ancestral caddis larva as postulated by Ross (1956).
The early stages (instars I-IV) of the Ptilocolepinae have not yet been discovered; the pheno-
menon of hypermetamorphosis has therefore not been verified in this subfamily.
DISCUSSION. The Ptilocolepinae appear to represent a natural extant link between the primitive
Glossosomatidae, with which they share many adult features, and the very specialized Hydrop-
tilinae with which they are classified on the basis of the affinities of the distinctive larvae and their
cases. The subfamily is confined to cool Holarctic regions, being found locally in clear, shaded
montane springs in association with bryophytes as is typical of the more primitive Trichoptera.
The Ptilocolepinae probably thus arose from the main hydroptiline stock very early in the evolu-
tion of the family before the typical adult hydroptilid form had been derived. It has since remained
very little changed except perhaps with respect to the development of the highly specialized and
characteristic male genitalia.
Keys to differentiate the Ptilocolepinae, Glossosomatidae and Rhyacophilidae
Adult
1 Discoidal cell of forewing open; spur formula 3.4.4 . RHYACOPHILIDAE
— Discoidal cell of forewing closed = >; oe spur formula 2.4. 4 Or 13.4) bs 2
2 Spur formula 1.3.4 : i . PTILOCOLEPUS (p. Ty
— Spur formula 2.4.4
3 Posterior dorsal head warts small, “oval - or round, widely separated: pronotal warts widely
separated (Fig. 54) . F GLOSSOSOMATIDAE
— Posterior dorsal head warts large, ‘almost meeting medianly; pronotal warts closely set (cf.
Fig. 53) . ; : : ; : 7 , : 2 PALAEAGAPETUS (p. 160)
Larva (partly after Ross, 1944)
1 A pair of well-developed nota present on all three thoracic segments; larvae usually inhabiting
purse-type cases of silk with particles of moss and liverwort attached PTILOCOLEPINAE
— Pronotum only sclerotized; larva without case or with small saddle-like case (flat ventrally,
vaulted dorsally) covered i in small stones or large sand grains . F Z
2 Anal prolegs with large elongate hooks; abdominal tracheal gills present or absent; free- living,
without case. RHYACOPHILIDAE
— Anal prolegs with very small retractile hooks; tracheal gills absent: with small saddle-like cases
covered in small stones ‘ ; : ; 5 : ; GLOSSOSOMATIDAE
Pupa
1 Presegmental and postsegmental dorsal hook plates on abdominal segments III-VII and
III-V respectively (Fig. 52) ; : ‘ ; 2
— Presegmental and postsegmental hook plates not with full complement as above . a
2 Pupal mandibles without teeth : ; : P . : - HYDROPTILINAE
— Pupal mandibles with teeth (Ptilocolepinae) : ‘ ‘ ; : : i : : 3
3 Pupal mandibles with 2 teeth , ; F ; : ‘ PALAEAGAPETUS (p. 160)
— Pupal mandibles with 1 tooth : ; . PTILOCOLEPUS (p. 160)
4 Presegmental hook plates always absent from abdominal segment III (i.e. present on IV—VIII;
post-: III-IV); spur formula 3.4.4 : ‘ . ‘ ‘ ‘ RHYACOPHILIDAE
160 J. E. MARSHALL
ventral
plate(vpl)
A
Figs 55-59 ¢ genitalia, Ptilocolepinae. 55-57, Ptilocolepus, lateral view, (55) Pt. granulatus (Pictet);
(56) Pt. colchicus Martynov; (57) Pt. extensus McLachlan. 58-59, Palaeagapetus celsus Ross,
(58) lateral view; (59) ventral view. (vide Figs 15, 16).
— Presegmental hook plates sometimes absent from segment III; postsegmental hook plates
always absent from segment III, sometimes also from segment V; spur formula 2.4.4
GLOSSOSOMATIDAE
Genus PTILOCOLEPUS Kolenati
Prtilocolepus Kolenati, 1848 : 102. Type-species: Ptilocolepus turbidus Kolenati, by monotypy.
DISTRIBUTION. Palaearctic.
Ptilocolepus contains four species which occur in the western Palaearctic region eastwards to the
Urals, the Caucasus and Iran, the commonest and most widespread species being granulatus. The
identity of vi/losus from Spain is questionable due to the inadequacy of the original description
(Navas, 1916); the species may prove to be synonymous with either granulatus or extensus (N.B.
P. villosus was not referred to by Schmid, 1949, in his paper on the Navas types).
The following key to the males of colchicus, granulatus and extensus is based on a study of
material in the BMNH collections. Females of granulatus are in the BMNH collections but have
not been described; the larva of this species has been described by Thienemann (1904) and Jacque-
mart & Coineau (1962).
DiaGnosis. Adult. Branches R,+ R; and M,+ M, fused in hindwing (Fig. 5), spur formula 1.3.4.
Larva. Abdomen without lateral tubercules, segment I with small dorsal tergite.
Pupa. Mandibles with two medial teeth.
Key to Ptilocolepus males (excluding villosus)
1 Hindwing with patch of modified setae; ‘inferior appendages’ (Fig. 56, /a) bifurcate with dorsal
process not longer than ventral, ventral process broad; ‘ventral plate’ with process (Fig. 56).
Caucasus and Iran . ; : é : : F : ‘ ; : . colchicus
— Hindwing without patch of modified setae; ‘inferior appendages’ not as above; ‘ventral plate’
without process. : ; ; : : . ; : : : : ; : 2
2 ‘Inferior appendages’ (Fig. 55, Ja) bifurcate, dorsal process longer than narrow ventral process;
‘ventral plate’ without process (Fig. 55). Spain, Italy, Central Europe ’ . granulatus
— ‘Inferior appendages’ (Fig. 57, Ja) non-bifurcate; ‘ventral plate’ without process (Fig. 57).
Portugal, Spanish Pyrenees. ; : , : ; ; : : : : extensus
Genus PALAEAGAPETUS Ulmer
Palaeagapetus Ulmer, 1912a : 35. Type-species: Palaeagapetus rotundatus Ulmer, by monotypy.
DISTRIBUTION. Nearctic (and Baltic Amber).
THE GENERA OF THE HYDROPTILIDAE 161
Palaeagapetus is a small genus containing one fossil species (the type) and three extant Nearctic
species. P. rotundatus is known only from Baltic Amber and is therefore probably of western
European origin (and thus possibly Eocene according to Ross, 1956). The three North American
species agree with the type form in features of venation, spur formula and male genitalia. P.
nearcticus Banks and guppyi Schmid are western species, from California to Vancouver Island,
while ce/sus Ross occurs in the east from the western Appalachians of North Carolina and Ten-
nessee, New Hampshire to the Laurentians of Quebec (Roy & Harper, 1975). According to
Wiggins (1977) it is likely that ce/sus will be found in montane areas between these two extremes.
There is no key to the species of Pa/aeagapetus and only the larva of ce/sus has been described
(Flint, 1962).
dorsal ‘maculations’ lateral tubercule
]
mm
Figs 60-61 Palaeagapetus celsus Ross. 60, larva, lateral view; 61, case, lateral view.
DraGcnosis. Adu/t. Branches R2, R; and M,, M, free in the hindwing (cf. Fig. 5); spur formula 2.4.4.
Larva. Abdomen with lateral pairs of segmental tubercules, segment I without tergite (Figs. 60, 61).
Pupa. Mandibles with one medial tooth.
Subfamily HYDROPTILINAE Stephens
Hydroptilidae Stephens, 1836 : 151. Type-genus: Hydroptila Dalman, 1819.
DIsTRIBUTION. Cosmopolitan (excluding polar regions).
The Hydroptilinae is a very heterogeneous subfamily considered here to contain six distinct
tribes characterized by fundamental adult, larval and pupal morphological features which unite
them and distinguish them from the Ptilocolepinae.
DiaGnosis. Adult (Fig. 1). Forewing length 1:2-6:0 mm: wings narrow with rounded apices and almost
complete venation in the larger, more generalized forms (Figs 6, 7) to highly attenuate with consequently
reduced venation in the smaller, more specialized groups (Figs 8, 9, 10); Cu, unbranched in the forewing
(Fig. 6, arrowed); wings densely pubescent, costal fringes usually long and well developed: sclerotization
of head capsule reduced in the smaller forms; z-shaped tentorium basically well developed, anterior arms
reduced medially to weakly sclerotized filaments (scarcely detectable in KOH-treated specimens) in
smaller, specialized genera; ocelli 3, 2 or 0, lateral pair close to eyes and some distance posterior to anten-
nae; antennae usually short, of approximately 30 segments (rarely 40, sometimes 18); postoccipital warts
well developed, ovoid to subspherical, often represented by distinct sclerites hinged to the head capsule
which, in Hydroptila males, conceal eversible scent-organs; compound eyes of variable size but usually
occupying whole width of head capsule: thorax (e.g. Fig. 2) with close-set median pronotal warts; mesono-
tum flat, prescutum absent, scutellum with posterior half forming a triangular flat area with steep sides
and a warty texture along edges only, anterior half triangular to arc-like, transverse suture present or
absent; mesokatepisternal suture absent (Fig. 3, inset) (I have found this to be a constant feature of the
Hydroptilinae, confirming Ross’s (1956) view that it is ‘usually’ absent): fore-tibia with never more than
one apical spur; meso-tiia with never more than one preapical spur (Fig. 4): abdomen with ventral
processes variable (segments VI, VII or VIII); sternite V (Fig. 12) without postero-lateral ridge but with
162 J. E. MARSHALL
a pair of small dorso-anterior pits usually containing a short, posteriorly directed setose membranous
process: 2 genitalia an oviscapt; segment IX usually long and telescopic (Fig. 17), rarely short and fixed
with segment VIII modified (Figs 121, 134): ¢ genitalia variable but usually with a characteristic generic
or tribe form (generalized form, Figs 15, 16); segment VIII usually with tergite and sternite distinct, rarely
fused; segment IX forming an annular ‘genital capsule’, sometimes with various postero-lateral processes,
antero-lateral apodemes and dorsal and ventral excisions, sometimes concealed by segment VIII; inferior
appendages (Ja) usually present, size and shape variable, sometimes with baso-dorsal processes (‘brac-
teoles’), sometimes fused and at times withdrawn into segment IX; tergite X reduced to dorsal plate (may
be absent), size and degree of sclerotization variable; subgenital plate (process) or appendages (Sp)
present; bilobed process (Bp) present ventral to subgenital plate, consisting of a membranous process with
two short apical setae (sometimes appears to be fused to subgenital plate); aedeagus (Ae) variable but
basically a long narrow sclerotized tube enclosing the phalloduct which opens apically, a pair of lateral
processes may occur along apical half (sometimes fused together or with central tube) which may be
represented by a single spiral ‘titillator’ (homologies uncertain).
Larva. Early instars (I-IV) free-living (Fig. 51), fifth instar case-bearing (e.g. Figs 74, 75). All three
thoracic segments with paired nota completely covering dorsa; abdomen distended, membranous, com-
pressed laterally or dorso-ventrally, rarely cylindrical, abdominal gills absent. Ecdysial lines of head dis-
tinct (Figs 18-19), sometimes fused; ventral apotome and postmental sclerites variable (Figs 21-28),
small posterior ventral apotome sometimes present (Fig. 27); labrum basically symmetrical; mandibles
(Figs 30-32) primitively symmetrical and each with mesal brush (in left mandible only in specialized
groups), cutting edges ridged or modified as sharp blades or ‘scoops’ (Figs 31-32): thoracic sternites vari-
able (Figs 38-46) with a maximum of three per segment (one median ‘oral’ and two lateral ‘anal’);
pleurites primitively with preepisternite free on all segments, usually free on segment I only; anal epimeral
pleurite sometimes present (Figs 35-37): legs variable, basically short and subequal, modified variously
as robust, strongly clawed clinging organs (e.g. Fig. 89) or with mid- and hindlegs long and slender as in
vegetation dwellers (e.g. Figs 123, 128, 132): abdominal tergites present or absent, tergite IX always
present; dorsal ‘rings’ usually present on segments I/II—VIII; anal prolegs usually fused to sides of segment
X (except in early instars and fifth instar of Alisotrichia and the Neotrichiini).
Case. Variable, basically purse-like of two silken valves with anterior and posterior slit-like openings
(e.g. Fig. 119), sometimes with incorporated inorganic or organic particles; usually held horizontally.
Sometimes, however, the case may be cylindrical or fusiform with slit-like (Fig. 138) or circular (Figs 103,
107) openings; in specialized torrenticolous or madicolous forms the case is greatly flattened dorso-ven-
trally and firmly fixed (temporarily or permanently) to the substrate (Figs 75, 78, 86) and may sometimes
have lateral ‘wings’ (Figs 76, 81, 148).
Pupa. Of the typical hydroptilid form (Fig. 52); presegmental and postsegmental plates on abdominal
dorsa III-VII and III-V respectively; mandibles without teeth, usually long. Case formed by sealing the
larval case which has been previously attached to the substrate by adhesive discs at the corners or by one
seam.
BioLoGy. The subfamily contains both cool- and warm-adapted genera which occur in a wide
variety of habitats; it is almost cosmopolitan in distribution, being absent only from the extreme
polar regions. Habitat preferences range from seeps, springs and splash zones of waterfalls to
clear, fast-flowing montane streams, and from all sizes of rivers to still (but rarely stagnant) ponds
in lowland regions.
The larvae are basically detritus-feeders but some groups have adopted special methods for
feeding on the fluid contents of the cells of filamentous green algae, the most specialized genus in
this respect being Orthotrichia. The distribution and life-histories of the algal-feeding groups are
dependent on those of the algae, and genera vary as to whether they occur with the algae on the
substrate or amongst vegetation. The primitive Nearctic genus Dibusa is unique in that it is
associated with the red fresh-water alga, Lemanea (Wiggins, 1977).
Key to tribes (for keys to genera see p. 153)
Adult
1 Transverse suture present on mesoscutellum (Figs 62, 83,93). ’ ? ‘ ; 2
— Transverse suture absent from mesoscutellum (e.g. Figs 108, 114, 120) 3
2 Metascutellum subpentagonal to triangular (Figs 83, 93) (Nearctic, Neotropical)
LEUCOTRICHIINI (p. 175)
— Metascutellum subrectangular to short pentagonal (Palaearctic) or as wide as scutum, very
short and rectanguloid (Figs 62-64) (Holarctic, Palaeotropical) : STACTOBIINI (p. 163)
THE GENERA OF THE HYDROPTILIDAE 163
3 dg aedeagus thread-like or armed with thickened spines, spiral ‘titillator’ absent (Figs 94, 96)
(Nearctic) ; : OCHROTRICHIINI (p. 184)
— 6 aedeagus usually divided into proximal and distal ‘sections, spiral ‘titillator’ usually present
(Figs 15, 101, 109, 122, 135, 140) : 4
4 ¢ genitalia asymmetrical (Fig. 135) or with segments Ix and x membranous dorsally, IX
incomplete ventrally with lateral sides tapering posteriorly, inferior appendages slender
(Fig. 140) (cosmopolitan) . : : : : : P ORTHOTRICHIINI (p. 212)
— (genitalia symmetrical, not as above. - 5
5 Adults small, forewing length 1-5-2:0 mm; segment 1X or - inferior appendages of 3 genitalia
with a pair of broad, flat, spatulate processes or lobes (bracteoles) (Fig. 101) (Nearctic,
Neotropical) . i : NEOTRICHIINI (p. 188)
— Adults small to large, forewing length 16 oe 6: 01 mm; 3 genitalia very variable, usually without
broad bracteoles (Figs 109, 115-117, 122, 127, 129, 131) (cosmopolitan)
HYDROPTILINI (p. 191)
Tribe STACTOBIINI Botosaneanu stat. n.
Stactobiinae Botosaneanu, 1956 : 382. Type-genus: Stactobia McLachlan, 1880.
DisTRIBUTION. Holarctic.
Botosaneanu (1956) erected the subfamily Stactobiinae for Stactobia and ‘les genres étroitement
apparentes’ (Stactobiella, etc.). Although Botosaneanu did not name these genera, he probably
intended the group to include Stactobiella, Plethus, Plethotrichia and Lamonganotrichia as sug-
gested by the knowledge of the group at that time. Ulmer (1957) considered Plethus and Lamong-
anotrichia to be closely related according to their larval affinities; on adult features alone, however,
Plethus bears more resemblance to Plethotrichia while Lamonganotrichia is more like Orthotrichia.
Ulmer also considered the larva of Plethotrichia to have more affinity with that of Hydroptila and
consequently placed it in the Hydroptilinae sensu Nielsen (1948) (vide Table 1).
Schmid (1959a), despite recognizing Botosaneanu’s subfamily grouping, remarked on the
relationships of Stactobia with Stactobiella, Chrysotrichia, Madioxyethira, Pseudoxyethira,
Parastactobia, Macrostactobia and Plethus (the last of which Schmid considered Plethotrichia to
be a possible synonym). Flint (1970), on the basis of larval morphology and case form, considered
Plethus and Lamonganotrichia to belong in the Stactobiinae but placed Stactobiella in the
Hydroptilinae sensu Nielsen (1948) along with Hydroptila, Agraylea and Oxyethira.
Morse (1974) has independently remarked on the relationship of Catoxyethira veruta Morse,
from S. Rhodesia, with the Holarctic genus Stactobiella according to features of the adults;
Ulmer (1957) had previously noticed the relationship of the larvae of Catoxyethira and Stactobia.
Examination of adult type-material has revealed errors in the original diagnoses of the genera
which are henceforth considered to comprise the tribe Stactobiini; such anomalies have occurred
in the interpretation of spur formulae, presence of ocelli and wing venation features, all of which
were once considered to be very important in deducing generic relationships. The present grouping
is thus based on more reliable features such as the male and female genitalia, head and thoracic
structures and amended ocellar counts and spur formulae; a number of previously unsuspected
relationships and possible synonymies have also been discovered.
Since the constituent genera exhibit a number of larval and adult features which are unique to
the stactobiines the group is treated as a distinct tribe although three main genus-groups may be
recognized. These are the Stactobia-group (Stactobia and Plethus), the Stactobiella-group (Stac-
tobiella, Bredinia, Parastactobia and Chrysotrichia) and the Madioxyethira-group (Madioxyethira,
Pseudoxyethira and Scelotrichia). Within the tribe the genera show various lines of development
and specialization in either the larval or the adult stages or both, but they all share the basic
common features (set out below) from which the principal characters of a hypothetical stactobiine
ancestor may be deduced. As more becomes known of the group, especially of the larval stages,
the following tentative conclusions may accordingly require modification.
Diacnosis. Adult (Figs 62-64). Head with tentorium complete; antennae short, usually 18-segmented;
ocelli 3: mesoscutellum with*transverse suture; metascutellum with anterior edge approximately straight,
parallel with posterior edge: abdominal segment V with typical short, lateral setose processes: 2 genitalia
164 J. E. MARSHALL
an oviscapt: ¢ genitalia (Figs 65-68) with tergite and sternite VIII distinct; IX reduced ventrally, sternite
vestigial (absent or fused with tergite ?); inferior appendages small, distinct; paired subgenital structures
(‘subgenital appendages’, Schmid, 1959a) heavily sclerotized, arched downwards, sometimes represented
by a fused sclerite (?); bilobed process absent; tergite X absent or reduced; aedeagus (Figs 69-73) long,
straight, without median constriction or spiral titillator, basically tripartite with a single apical process
arising midlength beneath (or above) which are two flattened processes fused along most of their length
(often only the apex reflects this form as a dilated, trilobed structure), sometimes rows of short apical
setae or heavily sclerotized subapical spines may be present.
Larva. The larval stages are only known for Stactobia, Plethus, Stactobiella and Catoxyethira (including
also Lamonganotrichia and Plethotrichia). The genera have different structural adaptations associated with
their various habits and ecological preferences and do not appear superficially to be as homogeneous as
the adults. The generalized larval form, as seen in Stactobiella, is that of the typical hydroptilid with the
fifth instars inhabiting transportable purse-type cases. However, the main evolutionary trend within the
group has been the adaptation to life in fast-running water and the thin surface film of water on rocks
(the madicolous, hygropetric or hydropetric habitat). Various degrees of specialization occur but the basic
trends are towards the dorso-ventral flattening of the body and case, the attachment of the case to the
substrate, the protection of the body by fusion and thickening of dorsal sclerites and the development of
the thoracic legs as strong clinging appendages. The most advanced genus in this respect is Stactobia,
which is truly madicolous, while Plethus, according to Schmid (1958a), although morphologically similar
to Stactobia does not share its ‘hydropetric’ habits. Schmid (1960) also states that Madioxyethira milinda
is exclusively madicolous, but this is purely speculative since the larva of this species is unknown.
Pupa. Typically hydroptilid with no distinguishing characters; mandibles long, without teeth.
DiscussION. On the basis of adult and, to a lesser extent, larval affinities the genera of the Stac-
tobiini as defined here appear to fall into three natural groups. Each group contains both
generalized and highly specialized representatives of which the former exhibit the basic features
by which the relationships of the three groups may be recognized.
The Stactobia-group is distinguished by the shape of the metascutellum and the general form
of the larvae. Plethus would appear to be the most typical and generalized representative on ac-
count of the unmodified male genitalia, which are highly specialized in Stactobia, and the larvae
which, although obviously adapted for life in running water, are not, according to Schmid (1958a),
as hygropetric in their habits as are those of Stactobia. Plethus is restricted to the SE. Asian region
while Stactobia appears to be a more successful group with a continuous recent temperate distribu-
tion in the Old World and a few subtropical representatives. The Stactobia-group appears to fill
the Old World niche occupied by the Leucotrichiini in the New World. Lamonganotrichia is
grouped with Stactobia and Plethus on the basis of larval affinities whereas Plethotrichia, although
almost identical with Plethus in male genitalic features, most closely resembles Stactobiella in
larval features; the latter may simply be due, however, to the retention of the generalized
hydroptilid form in both these genera.
The Stactobiella-group is characterized by the distinctive narrow, strip-like form of the metascu-
tellum of the adults. Stactobiella, Bredinia and Chrysotrichia have generalized male genitalia very
similar to those of Plethus; Catoxyethira and Parastactobia, however, both have very unusual,
specialized genitalia. The group as a whole has a wide distribution with Palaearctic, Nearctic,
Oriental and African representatives. Only the larval stages of two genera are known, Stactobiella
from a single Nearctic species (which has the unmodified hydroptilid form) and Catoxyethira from
essentially an unidentified African species which, according to Ulmer (19126; 1957), resembles
that of Stactobia). It may thus be that the larvae of Stactobiella (and Plethotrichia) have retained
the basic hydroptilid form while those of Stactobia (including Lamonganotrichia), Plethus and
Catoxyethira have become adapted and structurally modified for life in fast-running water
conditions.
Madioxyethira, Pseudoxyethira and Scelotrichia are African and SE. Asian genera grouped
here according to the shape of the mesoscutellum, the reduced apical process of the fore-tibia,
the shape of the post-occipital lobes and the unique forms of the male genitalia (and possibly also
the genital-duct loop of the female). The larvae of these genera are completely unknown but they
are included in the Stactobiini on the basis of the form of the thorax and the short antennae, and
because the male genitalia may be derived from the basic stactobiine form.
THE GENERA OF THE HYDROPTILIDAE 165
It would thus appear that the Stactobiini is essentially an early offshoot of the main Hydropti-
linae branch with representatives which have become isolated in the now equatorial regions of
Africa and SE. Asia. Stactobiella is distinct in that it has a more northerly Holarctic distribution
with local species in montane habitats extending from Central Europe to the eastern United
States. The most successful and best represented genus is Stactobia. This may be due to its special-
ized larval habits which allow it to fill a unique niche with no competition from any other Palae-
arctic hydroptilid. Stactobia also has a continuous Palaearctic distribution.
The Stactobiini may have arisen from originally ‘cool-adapted’ montane stream-dwelling
ancestors which gave rise to two main lines, the first (1) becoming ‘cool-temperate-adapted’ while
the second (2) became more ‘warm-adapted’. Line | is the Stactobiella-group containing the ‘cool-
adapted’ genus Stactobiella which has retained a northerly distribution. It appears to have crossed
via the Bering route down into the North American mountain chains of the western ‘Rockies’
and eastern Laurentians and Appalachians, where isolated populations are still maintained. A
more ‘temperate-adapted’ line leading to the African and SE. Asian genera Chrysotrichia,
Catoxyethira and Parastactobia may also have arisen fromline 1. The North American Stactobiella
line may have given rise to (if Stactobiella is not synonymous with) the Central American genus
Bredinia.
Line 2, consisting of the Stactobia- and Madioxyethira-groups, became more ‘warm-adapted’
and probably had a past continuous distribution extending from Africa through North Africa,
SW. Asia to SE. Asia at a time when these areas were linked by one great forest. When the climate
became drier and severed the links between these two continents, representatives of these once
widespread genera became isolated in the Oriental and African regions where the original con-
ditions were, and still are, maintained. In the latter continent these areas occur in the forests
surviving in western Africa, the mountain slopes of eastern Africa and along the rivers through to
southern Africa which are precisely the regions from which these hydroptilids have been recorded,
namely Nigeria and Zaire (compare also the distribution of Ugandatrichia). The most recent suc-
cessful representative of line 2, Stactobia, now has a continuous subtropical Palaearctic
distribution in the mountain ranges extending from eastern Asia (Japan) through Asia Minor, the
Mediterranean region and North Africa out to the Atlantic Islands. Some species of the furcata-
group have spread northwards into the more temperate Central European regions while
representatives of the vaillanti-group have been discovered in Africa (Guinea and Uganda).
In conclusion, the Stactobiini appears to have arisen from an ancestor which evolved early in
the history of the Hydroptilinae. The tribe is quite distinct from any other belonging to this sub-
family and its possible relationships with the New World tribe Leucotrichiini will be discussed
later.
Genus STACTOBIA McLachlan
Stactobia McLachlan, 1880 : 505. Type-species: Hydroptila fuscicornis Schneider, by subsequent designa-
tion by Mosely (1933).
Afritrichia Mosely, 1939b : 35. Type-species: Afritrichia aurea Mosely, by original designation and
monotypy. [Synonymized by Schmid, 1959a.]
Aratrichia Mosely, 1948 : 76. Type-species: Aratrichia fahjia Mosely, by original designation and mono-
typy. [Synonymized by Schmid, 1959a.]
Lamonganotrichia Ulmer, 1951 : 68. Type-species: Lamonganotrichia crassa Ulmer, by original designa-
tion and monotypy. Syn. n.
DISTRIBUTION. Palaearctic, SE. Asia, Africa.
McLachlan erected this genus for fuscicornis and a new species, eatoniella, the true identities
of which were subject to a careful investigation by Kimmins (1949). Larvae of the type-species were
collected by Zeller from Messina (Sicily) along with the adults on which Schneider based his
original description. Eaton (1873) referred to the biology of the larvae remarking that they ‘. . . at
first roam at large, caseless; when they become corpulent they construct oval cylindrical cases of
fine mud. They abound on rocks suffused with an extremely thin film of water resulting from the
spray and dribbling of trickling streamlets, especially in places exposed to the sun!’; he was thus
166 J. E. MARSHALL
Figs 62-64 Adult head and thorax, Stactobiini. 62, Catoxyethira sp.;
63, Stactobia sp.; 64, Madioxyethira sp.
the first to record both larval hypermetamorphosis in the Hydroptilidae and the madicolous
habits of Stactobia.
Prior to Schmid’s (1959a) paper on Stactobia the major contributions, albeit fragmentary, to
the knowledge of the genus were by Mosely (1933), Vaillant (195la; 19516; 1952; 1956) and
Botosaneanu (1956). Schmid subdivided the genus, which contains approximately 45 species, into
four main species-groups, of which a summary of the main features is given below (partly adapted
from Jacquemart, 1973). In the checklist the species are arranged according to these groups with
new species (i.e. post Schmid, 1959a) being assigned correspondingly. Examination of the type
of Lamonganotrichia crassa reveals that the genus Lamonganotrichia is synonymous with Stactobia
according to structural affinities of the adult head, thorax and genitalia and similarities in the
larva as described by Ulmer (1957).
D1acnosis. Adult. Forewing length 1-5—-4-(0 mm: compound eyes relatively small; postoccipital lobes wide,
suboval (Fig. 63): mesoscutellum narrow, deep (Fig. 63): spur formula 1.2.4 (Fig. 4) (0.2.4 in radavanovici
Schmid); fore-coxae notched and fringed (Fig. 4): ¢ genitalia (Fig. 66) specialized; sternite VIII displaced
posteriorly; segment IX reduced ventrally, often produced anteriorly as a pair of lateral apodemes; in-
ferior appendages small, rarely elongate; subgenital appendages strongly sclerotized, arched; tergite X
semimembranous; aedeagus long, straight, often considerably developed but sometimes thin and simple,
usually with a dilated, heavily spined, membranous apex; ventral process of VII long, sinuous and thick-
ened apically.
Larva. Early instars free-living, caseless, dorso-ventrally flattened and with tergites present on all
abdominal segments. Fifth instars (Fig. 74) case-bearing, dorso-ventrally flattened; sclerotization well
developed; setae modified as short, stout, protective spines. Sclerites of head fused; labrum symmetrical ;
mandibles (Fig. 30) not markedly asymmetrical, massive, heavily ridged, ‘shovel-like’, brushes present in
both left and right (Botosaneanu, 1956); anterior ventral sclerites paired (Fig. 25): thoracic nota paired,
medial lines distinct; preepisternite free on all three segments; sternites as in Fig. 38; legs short, subequal,
robust, adapted for clinging: tergites present on all abdominal dorsa; posterior margin of tergite IX with
‘crenellations’ (modified flattened setae 7).
Case (Fig. 75). Dorso-ventrally compressed; transportable but usually temporarily fixed to substrate;
barrel-shaped with dorsal and ventral longitudinal seams and slit-like anterior and posterior openings;
THE GENERA OF THE HYDROPTILIDAE 167
constructed of silk and fine grains of mud or sand. The case of Lamonganotrichia (Fig. 76, after Ulmer,
1957) is suboval or shield-shaped, with a central rectangular section occupied by the larva, and is bordered
by a wide lateral folded fringe; a ventral longitudinal seam appears to be present.
BIoLoGy. The greatest contribution to the knowledge of the biology of Stactobia was made by
Vaillant (1956) while detailed life-history data are given by Danecker (1961). The larvae are unique
within the Hydroptilidae because they are madicolous (hygropetric, hydropetric), that is they are
adapted to live in the tenuous habitat formed, according to Hynes (1970), by ‘thin sheets of water
flowing over rock faces’ which ‘although it is hardly a running-water habitat in the ordinary sense,
it is often situated very close to one’. Thus the habitat often occurs at the edge of streams by the
side of waterfalls and on rocky chutes, the faces of which may be almost vertical. Such conditions
require very special structural adaptations of the inhabitants which can be clearly seen in the
larvae and cases of Stactobia (Figs 74, 75). Both are dorso-ventrally flattened and capable of
maintaining a thin film of water over their dorsal surfaces by surface tension; the cases are tem-
porarily fixed but may be moved if conditions become unfavourable; the larvae are protected
dorsally by heavily sclerotized fused tergites and spine-like setae and their short, robust legs are
used for clinging to the substrate. The larval mouthparts are adapted for feeding on organic
particles (‘phytosaprophagous’ according to Vaillant, 1956), the mandibles (Fig. 30) being robust,
strongly ridged and having a scraping function. According to Vaillant (1956) the larvae are ex-
clusively petrimadicolous, that is they occur only on rocky substrates almost devoid of sediment
and filamentous algae. The pupae are often gregarious forming a ‘Puppenneste’ (Danecker, 1961)
in which the cases are closely packed in a single ‘colony’.
SPECIES-GROUPS (after Schmid, 1959a and Jacquemart, 1973). The furcata-group (20 species):
this is the largest and most homogeneous group, its main feature being the constancy of the form
of the male subgenital appendages; its distribution extends throughout central and southern
Europe, the Atlantic Islands, North Africa, Israel and Turkey.
The martynovi-group (13 species); this is less homogeneous than the previous group and is
characterized by the dorsal thickenings of tergite X and the anterior displacement of the inferior
appendages of the males (most marked in ulmeriana Schmid to which, in my opinion, schmidi
Kimmins from Nepal is most closely related); the group is restricted to southern Asia — Iran,
Pakistan, Sri Lanka, Burma, Nepal and Turkestan (U.S.S.R.).
The nielseni-group (seven species including Lamonganotrichia crassa): this group is distin-
guished by the modified sternite VIII and the absence of any subgenital appendages in the males;
representatives occur in Southern Yemen (Aden), Iran, Pakistan and Indonesia (Java).
The vaillanti-group (two species): exclusively an African group with vaillanti Schmid from Guinea
and aurea (Mosely) from Uganda.
S. bolzei Jacquemart (Turkey) is considered by Jacquemart (1965) to be distinct although
I believe that it may belong to the nie/seni-group (original description inadequate for direct
comparison).
S. japonica \wata (Japan) is known only from the larva of which the case bears more resemblance
to that of Plethus as figured by Ulmer (1957); the japonica of Tsuda & Nakagawa (1959) may be a
distinct species since again only the larvae were examined.
S. megalatlantica Vaillant and mallorcensis Vaillant are nomina nuda.
‘
DiscussION. Stactobia is a very specialized member of the Stactobiini. Although it is one of the
more successful hydroptilid genera in terms of both species and individual numbers it does not
have the widespread geographical ranges of, for example, Hydroptila, Oxyethira or Orthotrichia.
The madicolous habits of the larvae perhaps enable Stactobia to occupy a niche where it is free
from competition from these other genera, at least in the Palaearctic and parts of the Palaeotropi-
cal regions. The absence of Stactobia from the New World may be due to two factors: (i) the
slow rate of dispersal of montane stream-dwelling forms in comparison with lowland vegetation
dwellers where geographic barriers are less pronounced (see later discussions) and (ii) competition
with the highly successful New World Leucotrichiini which exhibit parallel modifications suiting
them to a similar mode of life.
168 J. E. MARSHALL
Figs 65-73 Generalized ¢ genitalia, Stactobiini. 65-68, lateral view, (65) Stactobiella, Plethus,
Chrysotrichia; (66) Stactobia (after Schmid, 1959a); (67) Madioxyethira; (68) Catoxyethira. 69-73,
aedeagus, dorso-ventral views, (69) Stactobiella; (70) Chrysotrichia; (71) Madioxyethira; (72)
Pseudoxyethira; (73) Catoxyethira. (vide Figs 15, 16.)
Stactobia could have arisen from temperate or subtropical Palaearctic stock which penetrated
southwards into Africa and SE. Asia and northwards into central Europe. It is totally absent
from the northern Palaearctic region, there being no Scandinavian or British representatives of
the genus.
Genus PLETHUS Hagen
Plethus Hagen, 1887 : 643. Type-species: Hydroptila cursitans Hagen, by monotypy.
Plethotrichia Ulmer, 1951 : 65. Type species: Plethotrichia baliana Ulmer, by original designation and
monotypy. Syn. n.
DISTRIBUTION. S. and SE. Asia.
Plethus was erected for the type-species from Sri Lanka and now contains another five species
from this island (Schmid, 1958a), one from Pakistan (Schmid, 1960) and three from Indonesia
(Ulmer, 1951); the larvae of acutus and cruciatus were described by Ulmer (1957). Plethotrichia
baliana was originally considered to be closely allied to Plethus but was separated on account of
the unmodified larval case (Ulmer, 1957); a close examination of the adult features, however,
indicates that the two genera are synonymous. Stactobia japonica |wata from Japan also resembles
Plethus in the form of the larval case.
D1acnosis. Adu/t. Forewing length 1-2 mm: head and thorax similar to those of Stactobia (cf. Fig. 63):
spur formula 0.2.3: wing venation greatly reduced; veins of forewing confluent before margin; veins of
THE GENERA OF THE HYDROPTILIDAE 169
hindwing reduced to the two main longitudinal subparallel stems of R and M; costal fringes well devel-
oped: ¢ genitalia (Fig. 65) simple, generalized without ventral processes; VIII unmodified; LX reduced
ventrally although the sternite may be vestigial in some species, anterior apodemes short; inferior ap-
pendages unmodified; subgenital appendages heavily sclerotized, arched; tergite X variable; aedeagus
long and straight without apical spines, sometimes with a pair of short basal processes: the anterior
abdominal segments of the male may bear sensorial organs (Schmid, 1958a) which, in baliana, occur on
tergites V and VI according to Ulmer (1951).
Larva (Fig. 77). Dorso-ventrally compressed; small abdominal tergites present, posterior margin of
tergite IX simple; legs short, robust, subequal, unmodified; thoracic sternites as in Fig. 43.
Case (Fig. 78). Dorso-ventrally compressed, barrel-shaped with dorsal and ventral seams and a lateral
fringe (‘wings’).
According to Ulmer (1957) the larvae of Plethotrichia are more like those of Oeceotrichia (here
synonymized with Hydroptila) in that they are not dorso-ventrally compressed, there are no
abdominal tergites and the fore-tibia bears ashort process. The case is of the purse-type, flattened
laterally and constructed of secretion and diatoms. However, the larvae were not reared and no
pupae were collected to confirm the association of larvae and adults.
BIoLoGy. The larvae inhabit brooks amongst ‘spongy algal masses’ of Cladophora and ‘silicaceous’
algae (Ulmer, 1957). According to Schmid (1958q) the adults are abundant by streams, fast-flowing
rivers and rocky streams and are not attracted to light. The larvae are not hygropetric although
Schmid (1958a) suspects that they live amongst the rocks in shallow water. According to Ulmer
(1957) the larvae of Plethotrichia live in flowing water, springs (including warm springs), waterfalls
and brooks, often in moss and algae.
DiIscussION. The adults of Plethus retain many primitive features of the generalized stactobiine
hydroptilid, their main specializations being their reduced size and wing venation and the presence
of specialized male abdominal sensorial organs. The larvae, although adapted to live in fast-
flowing shallow water, are not as specialized as the petrimadicolous larvae of Stactobia. Plethus
is essentially a warm-adapted genus confined to the Oriental region.
Plethus is most closely related to Stactobia and was probably an early offshoot of the Stactobia-
group branch of the Stactobiini and which has retained certain primitive attributes. Its main
diagnostic features are the adult spur formula, overall reduced size and less specialized genitalia,
larva and case (i.e. with respect to Stactobia).
Genus STACTOBIELLA Martynov
Stactobiella Martynov, 1924 : 57. Type-species: Stactobia ulmeri Siltala, by monotypy.
Tascobia Ross, 1944 : 124. Type-species: Stactobia palmata Ross, by original designation. [Synonymized
by Ross, 1948.]
DISTRIBUTION. Holarctic.
Stactobiella is a small Holarctic genus containing six species, three of which are Palaearctic
(northern and eastern Europe, Siberia) and three Nearctic which, according to Wiggins (1977),
have been recorded over much of the North American continent west to the Rocky Mountain
states and north to Minnesota, Ontario and Maine. S. risi from Switzerland was first described
in Microptila but was transferred to Stactobiella by Ulmer (1929); Dohler (1963) synonymized
risi with ulmeri but the species were later considered to be distinct by Botosaneanu (1967) and
are regarded as such here. Ross (1944) gives diagnostic characters for the males and females of
delira and palmata and (Ross, 1948) a key to the males of all six species.
The species are grouped below according to features of the male genitalia (after Ross, 1948).
The biramosa-group: biramosa (Siberia, ‘Russia’); palmata (Oklahoma, Wisconsin, Illinois,
Kentucky, Maine, Tennessee, Oregon).
The u/meri-group: ulmeri (Finland, Germany, Poland, Rumania, U.S.S.R.); risi (Switzerland) ;
delira (Wyoming, Colorado, Minnesota, Wisconsin, Maine, Kentucky, New Hampshire).
The brustia-group: brustia (Wyoming).
170 J. E. MARSHALL
DiaGnosis. Adult. Forewing length 1-5-3-0 mm: metascutellum narrow, as wide as metascutum (cf.
Fig. 62): spur formula 1.3.4: 3 genitalia (Fig. 65) with segment VIII unmodified; LX well developed, es-
pecially dorsally, with long anterior apodemes; inferior appendages fused in brustia, sometimes with baso-
dorsal bracteoles; subgenital plate strongly sclerotized, arched; tergite X absent or membranous; aedeagus
(Fig. 69) simple, tubular, with fused lateral processes and a free median process in delira.
Larva (after Ross, 1944 and Wiggins, 1977). Body slightly compressed laterally: abdomen without
tergites (except on segment IX); dorsal abdominal ‘rings’ present (II-VIII): legs short, subequal, stout;
tarsal claws distinct, sharply curved and with thickened basal spur nearly half as long as claw (Fig. 80):
setae unmodified: preepisternite free on all thoracic segments, episternite and epimeron fused in meso-
and metathorax.
Case (Fig. 79). Purse-type, laterally compressed, constructed of siik only.
BIOLOGY. The larvae of pa/mata are found in small, fairly swift streams on stones in riffles and they
mature in the early spring in Illinois (Ross, 1948). According to Wiggins (1977) it is therefore
likely that this species overwinters as final instar larvae.
Discussion. Adult features indicate that Stactobiella is related to Stactobia and Plethus although
the metascutellum shows a close resemblance to those of Chrysotrichia, Parastactobia and
Catoxyethira. The larvae are unspecialized and exhibit the basic hydroptilid form, the case being
of the purse-type and laterally compressed. The genus thus appears to be an early offshoot of the
Stactobiini which has retained many primitive larval and adult features. It is a temperate group
which may have originated in the Palaearctic region and has had at least two subsequent migra-
tions into the Nearctic region (via the Bering route) as indicated by the relationships of delira
with u/meri and risi, and of palmata with biramosa. The affinities with the other genera of the
Stactobiella-group will be discussed later.
Genus BREDINIA Flint
Bredinia Flint, 1968c : 50. Type-species: Bredinia dominicensis Flint, by original designation and
monotypy.
DISTRIBUTION. Lesser Antilles (Dominica).
The following diagnosis is adapted from the original description, the genus being known only
from the type-series collected on Dominica in 1965. The larval stages are unknown.
DiAGnosis. Adult. Forewing length 1-5 mm: ocelli 3: mesoscutellum with transverse suture; metascutellum
as wide as scutum, short and rectangular: spur formula 0.2.4: 3 genitalia with sternite VIII divided mid-
ventrally; IX narrow, oblique, lateral halves divided ventrally by the inferior appendages and with long
anterior apodemes; tergite X a large membranous lobe; inferior appendages small, quadrate; subgenital
plate elongate, rectangular; aedeagus tubular, apex flat and tridentate, central tubule in apical quarter.
BIioLoGy. The adults were taken only near the larger lowland rivers of the island.
DIscussIoN. Flint was uncertain of the relationships of this genus and simply listed its affinities
with Neotrichia (presence of ocelli, minute size and spur formula), with Mayatrichia (spur for-
mula) and with Alisotrichia (transverse suture of the mesoscutellum and lateral apodemes of segment
IX in the male genitalia). Although Flint noted the similarity of the thoracic nota of Bredinia to
those of Stactobiella, he did not consider these genera to be closely related. According to Flint’s
(1968c; 1970) descriptions, Bredinia bears more resemblance to Stactobiella than to Alisotrichia,
especially with respect to the thoracic nota (cf. Figs 62, 83), the lateral apodemes of the male
segment IX and the tridentate aedeagus (Figs 65, 69-73, cf. Fig. 84). The main distinction between
Bredinia and Stactobiella is their spur formulae (0.2.4 and 1.2.4 respectively). As Flint notes,
however, the affinities of Bredinia may be clarified when the larval stages have been discovered.
Genus CHRYSOTRICHIA Schmid
Chrysotrichia Schmid, 1958a : 54. Type-species: Chrysotrichia hatnagola Schmid, by original designation.
DISTRIBUTION. S. and SE. Asia (Sri Lanka, India and Sarawak).
Five species described from Sri Lanka (Schmid, 1958a) and India (Schmid, 1960) comprise this
small SE. Asian genus. The immature stages are unknown but the adults appear to be closely
THE GENERA OF THE HYDROPTILIDAE 171
allied to those of Stactobiella according to features of the head, thorax and male genitalia.
Chrysotrichia may be distinguished by its distinctive spur formula and reduced size and wing
venation. Examination of more material, especially of the larval stages, will be needed before any
definite conclusions regarding relationships can be reached. The following generic diagnosis is
based on original descriptions and examination of paratype material in the BMNH collections.
Series of what may prove to be one or more new species of Chrysotrichia have recently been
collected from Sarawak, 1977 (B. Bolton); 1978 (J. E. Marshall). The specimens are in the BMNH.
DiaGnosis. Adult. Forewing length 1:25-1:50 mm: head and thorax as in Stactobiella (Fig. 62): ocelli 3
(N.B. absent according to Schmid, 1958a): wings essentially reduced to thin ‘ribbon-like’ strips; venation
represented by only two main longitudinal veins in the hindwings, veins of forewings reduced but not
concurrent at the margin as in Plethus; costal fringes very long: spur formula 0.2.4: ¢ genitalia general-
ized as in Stactobiella and Plethus (Fig. 65); segment VIII simple; [X reduced ventrally, anterior apodemes
present; inferior appendages small; subgenital plate strongly sclerotized, arched; tergite X membranous;
aedeagus (Fig. 70) long, simple, with a trilobate apex and sometimes with minute apical spines (as in
hatnagola).
BrioLocy. According to Schmid (1958a) the adults occur along calm rivers at moderate altitudes.
C. badhami from the Punjab was taken by a fairly large river rich in aquatic vegetation (Schmid,
1960).
Discussion. Schmid (1958a) states that the male genitalia greatly resemble those of Plethus
from which they may be distinguished by the small size and weak sclerotization of the two
terminal segments in comparison with the remaining abdominal segments. The male genitalia
are also similar to those of the u/meri-group of Stactobiella; the diagnostic features at present used
to distinguish this genus from Chrysotrichia (spur formula, wing size and venation) may not be
as critical as originally thought and the genera may prove to be synonymous.
Genus CATOXYETHIRA Ulmer
Catoxyethira Ulmer, 19126 : 82. Type-species: Catoxyethira fasciata Ulmer, by monotypy.
Sperotrichia Marlier, 1978 : 294. Type-species: Sperotrichia mali Marlier, by original designation and
monotypy. Syn. n.
DISTRIBUTION. Africa.
Catoxyethira was erected for fasciata from Zaire by Ulmer (19126) who described and figured
the male genitalia. Ulmer (19125) also outlined the main features of the larva and case of an un-
identified species which he placed in Catoxyethira of which the generic identity is questionable
since no positively determined adults were associated with it. C. pinheyi and veruta have since been
described from the Victoria Falls and Rhodesia respectively and two new species, ocellata and
improcera have been described by Statzner (1977) from Zaire; undetermined material collected by
Prof. J. Medler from Nigeria is in the BMNH collection. Examination of the holotype and para-
type of Sperotrichia mali from Mali reveals that this genus is synonymous with Catoxyethira.
As independently noted by Morse (1974), Catoxyethira has a number of structural affinities
with the Holarctic genus Stactobiella especially with regard to the adult spur formula, thoracic
nota and male genitalia. Although the latter are very specialized in Catoxyethira they may be
derived from the basic stactobiine form. As Morse points out, however, further studies are needed
to clarify the taxonomic limits of the genera.
Hydroptila formosae Iwata from Taiwan (Formosa) was described from the larva and case only,
according to which Nielsen (1948) and Ulmer (1957 : 186) suggested that the species might belong
in Catoxyethira. However, neither Nielsen nor Ulmer formally transferred the species which was
done without reasoning by Fischer (1971) in his catalogue. Thus the species remains in Catoxye-
thira until examination of additional material can establish its true identity.
The generic diagnosis is based on examination of the following material (all in BMNH collec-
tions except where otherwise stated): fasciata (UHZIM); pinheyi (¢ type); improcera and ocellata
(paratypes); undetermined species (Medler).
172 J. E. MARSHALL
dorsal
seam
foreleg
lateral
flange
tergite
75
\ ventral
\ seam
dorsal
ring
margin of
81
80
79 panel claw 2 J
— ie
Imm
end-on view
Figs 74-82 Stactobiini. 74-81, larvae and cases. 74-75, Stactobia (adapted from Vaillant, 19515),
(74) larva, dorsal view; (75) case, ventral view. 76, Lamonganotrichia, case, dorsal view (after Ulmer,
1957). 77-78, Plethus cruciatus Ulmer (after Ulmer, 1957), (77) larva, dorsal view, and foreleg,
lateral view; (78) case, dorsal view. 79-80, Stactobiella, (79) case, lateral view; (80) larval tarsus,
lateral view. 81, Catoxyethira, case, dorsal and end-on views (after Ulmer, 19125). 82, Madi-
oxyethira, 2 genitalia, ventral view.
DiaGnosis. Adult. Forewing length 1-5 mm: head and thorax as in Stactobiella (Fig. 62): ocelli 3 (cf.
absent according to Ulmer, 19125): spur formula 1.3.4 (cf. 0.3.4 according to Ulmer, 19125): 3 genitalia
(Fig. 68) very specialized (main diagnostic feature of genus); sternite VIII elongate with a pair of posterior
dorso-lateral spines; IX reduced, withdrawn into VIII, sternite may be present although small and fused
with tergite, anterior apodemes narrow and produced beyond VIII; inferior appendages may be present
(fused with tergite X in veruta); tergite X semimembranous in veruta, membranous and covered in micro-
scopic setae in pinheyi; aedeagus long, straight, tapering to acute apex (Fig. 73) (the characteristic features
of the male genitalia of fasciata, pinheyi and veruta are summarized by Morse, 1974).
Larva. Young larva 1:2 mm long, fifth instar 2-5 mm. According to Ulmer (19125) the larva is somewhat
similar in appearance to that of Stactobia from which it may be distinguished by the absence of the
posterior ‘crenellations’ of tergite IX (cf. Fig. 74), the short spines of the posterior margin of tergite VIII
and stout spines on tergites VIII and IX. According to Iwata (1928) formosae has sternites on abdominal
segments IIIJ-V which do not occur in the African species of Catoxyethira and appear to be unique to this
species within the Hydroptilidae.
Case (Fig. 81). According to Ulmer (19125) this resembles the ‘shield-case’ of Molanna Curtis (Molan-
nidae : Trichoptera) as it is dorso-ventrally flattened and has a central tubular region from which arise
THE GENERA OF THE HYDROPTILIDAE 173
the lateral ‘wings’. The case, constructed of silk and fine sand-grains, tapers anteriorly and posteriorly
and is attached to the substrate by the edges of the ‘wings’.
BrioLoGy. Nothing is known of the biology of Catoxyethira but, from the dorso-ventrally flattened
larva and fixed case, it would seem likely that the larva lives in fast-flowing water habitats such as
swift streams (somewhat stony ?). Ulmer’s (19125) specimens were taken from a tributary of the
Butagu, Ruwenzori West (Zaire) at 1800 m, February 1908; the adults of improcera and ocellata
(Statzner, 1977) were from a ‘spring-brook’ community (Kalengo stream, Zaire) and the larvae
of formosae were taken in ‘rapid mountain streams’ (Iwata, 1928).
Discussion. The form of the adult head and thorax indicate that Catoxyethira belongs to the
Stactobiella-group of the Stactobiini. Discounting the dubious Taiwan species formosae, Catoxye-
thira appears to contain five exclusively African species which are characterized by the highly
specialized and distinctive male genitalia. The uniqueness of the latter within the Stactobiella-
group suggests that the ancestors of Catoxyethira reached Africa quite early in the history of the
group and there evolved as a more tropical, warm-adapted line in isolation from the basic tem-
perate stock. The recent discovery of a number of new species, all collected in considerable num-
bers, indicates that the genus is more widespread and successful than may have previously been
thought.
Genus PARASTACTOBIA Schmid
Parastactobia Schmid, 1958a : 48. Type-species: Parastactobia talakalahena Schmid, by original designa-
tion and monotypy.
DISTRIBUTION. S. Asia (Sri Lanka).
Parastactobia is known only from the type-species in which the male genitalia are unique and
difficult to homologize with those of any other hydroptilid. Examination of the female paratype
(loaned by the USNM) has revealed that, contrary to Schmid’s original description, three ocelli
are present and that the thorax most closely resembles that of the Stactobiella-group (assuming
that the female had correctly been associated with the male type). The immature stages are
unknown.
DiaGcnosis. Adult. Forewing length 1:75-2:25 mm: head and thorax as in Stactobiella (cf. Fig. 62): ocelli
3: spur formula 0.3.4: ¢ genitalia distinct (after Schmid, 1958a); segment IX with a large ventral notch
(in which sternite VIII is completely encased) and a long asymmetrical dorsal spine and two tapering
ventral appendages; tergite X reduced; aedeagus large with paired internal supports and a very complex
apex.
BioLoGy. Schmid (1958a) records the adults from beside a small, stony, fairly calm river in a dense
forest in a small montane locality.
Discussion. Little can be said regarding the relationships of Parastactobia until more adult (and
larval) material is available for study. If the female paratype examined had been correctly associated
with the male type then the genus would appear to be a highly specialized member (with respect
to the male genitalia) of the Stactobiella-group as suggested by the structure of the head and tho-
rax. I do not agree with Schmid (1958a) that the male genitalia resemble those of Macrostactobia
(q.v.).
Genus MADIOXYETHIRA Schmid
Madioxyethira Schmid, 1960 : 89. Type-species: Madioxyethira milinda Schmid, by original designation
and monotypy.
DISTRIBUTION. S. and SE. Asia (Pakistan, Nepal, Sarawak); Africa (Zaire, Congo).
Schmid erected this genus for the type-species from Pakistan. Kimmins (1964) described nepalen-
sis from Nepal and Statzner (1977) has described a new species from Zaire (marshalli) which, on
the basis of affinities of the metascutellum and the male and female genitalia, he has placed in
Madioxyethira. The occurrence of the genus in the African continent is supported by the opinion
that Hydroptila trifurcata Jacquemart from the Congo should rightly belong in Madioxyethira
174 J. E. MARSHALL
according to the form of the male genitalia (however, the figures given by Jacquemart (1962a) are
not easy to compare with those of established species). The immature stages are unknown.
A 3 of what is possibly a new species of Madioxyethira was recently collected in Sarawak, 1978
(J. E. Marshall), in BMNH.
The following diagnosis is based on examination of the male type of nepalensis and a female
paratype of marshalli (BMNH coll.).
DraGnosis. Adult. Forewing length 2:25-3:25 mm: head and prothorax typical of the Stactobiini although
the metascutellum (Fig. 64) is intermediate between those of Stactobia and Stactobiella (Figs 62, 63) and
the postoccipital lobes are characteristically narrow: ocelli 3: spur formula 1.2.4, fore-tibial spur reduced
to a small subspherical process (diagnostic): 2 genitalia a typical oviscapt but with a characteristic loop
in the anterior duct of the internal system (Fig. 82): 3 genitalia very complex and difficult to homologize
(Fig. 67); segment IX relatively small but produced anteriorly as a long point into the preceding segments;
inferior appendages large, concave and ventrally prolonged as a unique plate which terminates on sternite
VII; segment X forming two lateral obtuse plates (‘bourrelet bombé’ of Schmid, 1960) which Kimmins
(1964) considers to be prolongations of tergite IX (Kimmins also considers that the ‘inferior appendages’
may have arisen from the fusion of sternites VIII and IX and the true inferior appendages); the aedeagus is
narrow and simple in milinda while in nepalensis it has a complex apex divided into two narrow foliate
lobes with two narrow divergent spines (Fig. 71).
BioLoGcy. According to Schmid (1960) milinda is common and abundant in parts of Pakistan
between 1450 and 3000 m and its habits are exclusively madicolous (although Schmid did not
collect or, therefore, examine larvae).
Discussion. As will be shown later, Madioxyethira may prove to be synonymous with Pseudoxye-
thira Schmid and Scelotrichia Ulmer (in which case the last name takes priority). Schmid (1960)
remarked on the possible synonymy of Madioxyethira and Pseudoxyethira since he could only
distinguish these genera (apart from using genitalic features) on the venation of the hindwings,
especially with respect to vein ScR.
Genus PSEUDOXYETHIRA Schmid
Pseudoxyethira Schmid, 1958a : 44. Type-species: Pseudoxyethira asgiriskanda Schmid, by original
designation and monotypy.
DISTRIBUTION. S. Asia (Sri Lanka).
Only the adult stages of this highly specialized genus, represented by a single species from Sri
Lanka, are known. Examination of a paratype female (USNM) reveals that the mesoscutellum
and postoccipital lobes are identical to those of Madioxyethira (q.v.) (the specimen was returned
before the significance of the fore-tibial spur and genital-duct loop were discovered and these were
therefore not investigated).
DiaGnosis. Adult. Forewing length 2:75-3:00 mm: head and thorax as in Madioxyethira (Fig. 64): ocelli
3: spur formula 0.2.4 (according to Schmid, 1958a): 3 genitalia very specialized with the ventral half of
segment IX extending anteriorly as a plate to the middle of sternite VII, but not invaginated into the
preceding segments; dorsal half of segment IX massive; appendages absent; aedeagus ‘long and spinifer-
ous’ (Fig. 72).
BioLoGy. According to Schmid (1958a) the adults frequent rivers at moderate altitudes and also
elevated regions; they run about on rocks on the river banks.
Discussion. If, as Schmid (1960) suggests, this genus is synonymous with Madioxyethira (q.v.)
the name Pseudoxyethira takes priority. However, this is complicated by the possibility that
Scelotrichia (see below) may also be a synonym since this would then take precedence.
Genus SCELOTRICHIA Ulmer
Scelotrichia Ulmer, 1951 : 73. Type-species: Scelotrichia saranganica Ulmer, by original designation and
monotypy.
DISTRIBUTION. Indonesia (Java).
Only two male specimens of the type-species of this genus are known. The type-specimen of
saranganica (UHZIM) was examined but was not in a condition favourable for the recognition of
THE GENERA OF THE HYDROPTILIDAE 175
critical features; however, the postoccipital lobes resemble those of Madioxyethira and the meso-
scutellum has a transverse suture. The following diagnosis is partly adapted from Ulmer (1951).
DiaGcnosis. Adult. Forewing length 2-8 mm: head as in Madioxyethira (Fig. 64): thorax with transverse
mesoscutellar suture (cf. Fig. 62): ocelli 3: spur formula 1.2.4, fore-tibial spur minute (Ulmer, 1951 : pl. 2,
figs 31B, C): 3 genitalia difficult to interpret from Ulmer’s figures.
Discussion. According to features of the head and mesothorax, Sce/otrichia is a member of the
Stactobiini. The shape of the postoccipital lobes and the presence of a small fore-tibial spur (and
also the general appearance of the male genitalia) indicate that the genus may be very close to, if
not synonymous with, Madioxyethira and Pseudoxyethira (see above); in that case the senior
name would be Scelotrichia. Further examinations of material and discovery of the larval stages
are required to establish the true identities of these genera.
Tribe LEUCOTRICHIINI Flint stat. n.
Leucotrichinae Flint, 1970 : 2. Type-genus: Leucotrichia Mosely, 1934.
DISTRIBUTION. North, Central and South America.
Flint (1970) erected the subfamily Leucotrichiinae (as Leucotrichinae) for the distinct group of
Nearctic and Neotropical hydroptilids consisting of Leucotrichia and its closely related genera.
Since the group has many of the features characteristic of the Hydroptilinae in general it is con-
sidered here at the status of tribe.
The tribe is predominantly of Central American distribution (including the West Indies)
although a few species occur in North America while some are found as far south as Chile in
South America; the latter continent is very under-collected and will undoubtedly yield many new
species.
Flint (1970) stated that he could give no single character by which the group could be distin-
guished from other hydroptilids in the adult stages but he did list the following as being diagnostic
when present: ocelli reduced to 2; head modified; antennae modified; reflexed costal cell (basal
costal ‘pouch’ or ‘bulla’) present on male forewing. Flint did not define the basic structure of the
male genitalia although, in his words, ‘there is . .. something characteristic’ about their form. As
will be shown in the diagnosis given below, the adults of the Leucotrichiini do possess a number
of distinguishing features which clearly set the group apart from the other hydroptiline tribes;
the larvae (at least in those genera which are known) also appear to be very distinct due to their
structural modifications associated with their torrenticolous habits. The following list of diagnos-
tic adult and larval characters has been based on an examination of Mosely’s type-material
(BMNH) and a study of Flint’s (1970; 19726; 1974) papers on Leucotrichia and related genera.
The tribe Leucotrichiini is considered tocontain those genera originally included by Flint (1970),
with the addition of Ce/aenotrichia, of which Flint considered Alisotrichia to be distinct according
to features of the male aedeagus, the female genitalia and the larval stages. The tribe thus contains
the Leucotrichia-group: Leucotrichia, Zumatrichia, Peltopsyche, Anchitrichia, Costatrichia,
Acostatrichia, Betrichia, Abtrichia and Celaenotrichia, and the Alisotrichia-group: Alisotrichia.
DiaGnosis. Adu/t. Forewing length 1-2-4-0 mm; wings brilliantly coloured, sometimes spangled with green
and silver; the male forewing sometimes has a basal costal pouch (bulla) which may contain modified
setae; head and tentorium well developed, former often with modifications in the male; antennae generally
short, of approximately 13-20 segments, male basal or median segments sometimes modified; ocelli 3,
often reduced to 2 in the male (rarely also in the female): mesoscutellum with transverse suture; metascu-
tellum pentagonal or subtriangular: spur formula 1.3.4 (except Alisotrichia, 0.2/3.3/4): fifth abdominal
segment with typical lateral setose processes: 2 genitalia a simple oviscapt, internal apparatus well de-
veloped (in Alisotrichia segment VII is modified and the internal apparatus is simple and ‘ring-like’):
3 genitalia (Fig. 84) often with ventral processes; sternite VIII distinct, produced posteriorly beneath
segment IX, sometimes with postero-lateral processes; segment IX fused, incomplete ventrally, sometimes
with postero-lateral processes; inferior appendages narrow, short, rod-like, basically unmodified but
sometimes fused medially and bearing baso-dorsal bracteoles; tergite X membranous or well sclerotized,
fused latero-ventrally with the subgenital appendages which may be indistinct or developed as heavily
176 J. E. MARSHALL
sclerotized plates; latter fused ventrally with subgenital plate which may be produced ventrally and may
articulate with an unidentified structure lying between the inferior appendages; aedeagus (Fig. 84) an
elongate tube with a median constriction and (except in Alisotrichia) a complex medial structure consisting
of a basal loop and a pair of basally directed processes, sometimes with a pair of circular ‘windows’ in
lateral view, apex membranous with various spinose processes and lateral plates.
Larva (Figs 85-89, 92). Larvae have been associated with Leucotrichia, Zumatrichia, Peltopsyche,
Anchitrichia, Abtrichia and Alisotrichia; they typically exhibit hypermetamorphosis (Flint, 1970), the
early instars being entirely free-living and caseless. From a specimen of what would appear to be a young
larva froma small collection of Peltopsyche sieboldii Miller (BMNH collection), the early stages are dorso-
ventrally flattened with slender, tapering abdomens and short, robust clinging legs; there is a single
tergite on each abdominal segment and the long, slender anal prolegs project laterally from the sides of
segment X. In all genera (except Alisotrichia) the final (fifth) instar builds a permanently fixed case in
which it remains throughout the stage. The fifth instar exhibits structural modifications suiting it for such
a sedentary existence in comparison with the earlier free-living stages: the body again is dorso-ventrally
flattened; the ecdysial lines of the rugose or papillate head are variously fused, as are those of the meso-
and metathoracic nota; the legs are short and robust; the abdomen is distended, especially segments V and
VI which are abruptly enlarged, and small tergites are present on segments I-VIII while tergite IX is large,
shield-shaped and sometimes bears enlarged spine-like setae; the anal prolegs are fused to the sides of
segment X, the claws alone being free. The genus Alisotrichia is unique in that the fifth instar larva does
not construct a case until just prior to pupation. The larva accordingly retains the structural adaptations
of the earlier stages (cf. Figs 92, 51) with a slender, tapering abdomen, large abdominal tergites (i.e.
relative to the size of the segments) and long, slender, freely projecting anal prolegs which are not fused
to the sides of segment X.
Case (Fig. 86). The cases of the Leucotrichiini are said to resemble leech egg-cases. They are strongly
depressed dorso-ventrally and composed of tough secretion, rarely embedded with small inorganic par-
ticles, and sometimes with transverse strengthening ridges; the dorsal surface is slightly vaulted while the
flat ventral surface is firmly attached to the substrate; there is a small circular opening at either end.
Pupa. Typically hydroptilid: mandibles long, without teeth. The pupal case varies between genera and
species but basically resembles the larval case with sealed anterior and posterior ends. In Alisotrichia
hirudopsis, where the case is not built until just before pupation, the central suboval region is surrounded
by an irregular flange by which the case is attached to the substrate and which has two to six small round
openings (possibly for water circulation according to Flint, 1964; 1970). In A. spangleri the pupal case is
torpedo-shaped and is attached to the substrate from one end by a silk thread 2-8 mm long (Flint, 1970);
in this way the case floats freely just below the surface of the water and this may serve to prevent desiccation
due to fluctuating water levels (Flint, 1970).
BIOLOGY. The larvae (early and fifth instars) are adapted to live in torrenticolous conditions, usually
being found in great numbers on boulders in the rapid sections of fast-flowing rivers. Some are also
found in the thin surface film of water on partly exposed rock surfaces moistened by spray from
nearby cascades (similar to the madicolous habitat of Stactobia). Structural modifications include
dorso-ventral flattening, fixed cases, well-developed and fused sclerites, protective spinose setae
and short robust clinging legs.
The larvae are reported to feed on ‘periphyton’ and small particles of detritus on the surround-
ing rock surfaces. In the case-dwelling forms the larva extends its slender anterior end through
either of the narrow case openings and grazes on the immediate surroundings. The larva never
leaves the case and this, in the later stages of the instar, would be impossible anyway due to the
distension of the middle abdominal larval segments. The retention of the free-living habit by the
final instar of Alisotrichia may be an adaptation serving to increase the feeding range of the larva.
In all genera the mandibles are adapted for scraping and grazing as they are strong with blade-like,
sometimes dentate, edges.
The adaptation of the pupal case of Alisotrichia spangleri for the prevention of desiccation due
to fluctuating water levels shows a remarkable parallelism with that of the larva of Rhyacopsyche
hagenii which also inhabits a torpedo-shaped case anchored by a thread to the substrate; however,
in this species, the pupal case is fixed.
Adult Leucotrichiini are reported to be most active on the exposed parts of the rocks on which
the larvae dwell. They prefer bright sunlight and congregate on the rock surfaces and run around
often describing semicircular paths. Such habits are typical of hydroptilids in general.
THE GENERA OF THE HYDROPTILIDAE Lies?
abdominal
tergite
Figs 83-86 Leucotrichia. 83, adult head and thorax, dorsal view; 84, generalized 3 genitalia, lateral
view (vide Figs 15, 16); 85, larva, lateral view; 86, larva in case, dorsal view.
Discussion. As can be seen from the generic key, couplets 13 to 19, it has not been possible to
define the genera of the Leucotrichiini satisfactorily since species from two or more genera often
key out with one another (e.g. Abtrichia and Betrichia; Costatrichia partim, Betrichia partim,
Leucotrichia partim and Celaenotrichia). Flint (e.g. 19726; 1974) appears to be very inconsistent
in his choice of criteria when assigning new Neotropical species to genera, using features of the
head, antennae and wings in some cases and genitalic features in others. Genera were originally
erected on the basis of certain (then apparently distinctive) characters common to a group of
species from a particular geographical area but, as new species have been discovered from other
areas, the generic limits do not appear to be so clear-cut. The larval generic key is also very un-
satisfactory at present since it is based on only a few species of each genus (only one in some cases
and, in Anchitrichia, association with adults is purely assumptive). The tribe is thus in need of a
thorough review which, however, would probably best be done by American workers due to the
location of most of the material.
In general, however, the Leucotrichiini appears to form a distinct group within the Hydropti-
linae and is characterized by a number of both adult and larval features. The origins of the tribe
178 J. E. MARSHALL
are speculative but the ancestral form appears to have given rise to a very specialized group with
an essentially Central American distribution. In this region the ecological conditions favour short
generation cycles resulting in high speciation rates and species diversity.
The tribe has greatest superficial affinities with the essentially Old World Stactobiini, particu-
larly the Stactobia-group. Common adult features are the short antennae, the transverse meso-
scutellar suture (cf. Figs 62, 83) and the simple basic form of the male genitalia (cf. Figs 65, 84)
while the larvae are similarly dorso-ventrally flattened, have clinging legs, protective abdominal
tergites, modified setae and fixed cases (cf. Figs 74, 75, 77, 78, 85-87, 92). However, the larval
similarities between the Leucotrichiini and the Stactobia-group are probably due to their com-
parable madicolous habits since they are all functional and protective adaptations to living in this
type of aquatic habitat and are also seen in a number of unrelated insect groups (see Vaillant, 1956;
Hynes, 1970). The larvae of the two tribes may be distinguished by certain fundamental differences
in, for example, the form of the head, the complete fusion of all thoracic nota (i.e. medially) and
the characteristic enlargement of the abdomen in the Leucotrichiini; the cases of the Leucotrichiini
do not have dorsal and ventral seams and are of a different shape. The adults differ in the shape of
the metascutellum, the reduced spur formula of the Stactobiini and the different modifications of
the male genitalia.
Nevertheless, the Leucotrichiini and the Stactobiini appear to have more in common with
one another than with any other group of hydroptilids and may have arisen from a common
ancestor very early in the evolutionary history of the Hydroptilinae. This common ancestor may
have arisen in the Old World temperate region and given rise to two main lines, one of which
colonized the cool, montane streams of the Palaearctic and Palaeotropical regions (the present-day
Stactobiini) while the other penetrated the American continent (probably via the Bering connec-
tion) and passed down to the Central American region where the descendants of these early
arrivals evolved into the highly successful Neotropical Leucotrichiini.
However, this hypothesis is tentative and further evidence from new species and distribution
records is required in order to present a more accurate picture of the evolutionary history of the
Leucotrichiini. The tribe may prove to be more distantly related to the Stactobiini than indicated
above, the morphological similarities being the result of convergent rather than parallel evolution.
In conclusion the Leucotrichiini is a very specialized and successful member of the Hydroptilinae
which has a predominantly Central American distribution and may share a common ancestry
with the Stactobiini.
Genus LEUCOTRICHIA Mosely
Leucotrichia Mosely, 1934 : 157. Type-species: Leucotrichia melleopicta Mosely, by original designation.
DisTRIBUTION. North and Central America and the Antilles.
Leucotrichia contains ten species of predominantly Central American distribution although
sarita and limpia have been recorded from the southern United States, while pictipes is widespread
throughout the U.S.A. (but not so far recorded from Canada). The larva of pictipes was first
described by Lloyd (1915) (as Ithytrichia confusa) and larvae have now been associated with all
species except melleopicta, virida and fairchildi.
DiaGcnosis. Adult. Forewing length 2-5 mm: dorsal region of head sometimes modified; ocelli 3 or 2
(3 only): metascutellum pentagonal (Fig. 83): wings unmodified: spur formula 1.3.4: 3 genitalia simple
(Fig. 84) with a single posterior spine or paired setal brushes on sternite VII; sternite VIII posteriorly
produced beneath segment IX; 1X completely open ventrally, postero-lateral margin with a row of stout
setae; tergite X heavily sclerotized; subgenital plate connected dorsally to tergite X and produced ven-
trally as a narrow, elongate mesal sclerite which projects down to the base of the inferior appendages; sub-
genital appendages small, semimembranous; inferior appendages unmodified, elongate and usually fused
meso-ventrally; aedeagus with median complex and spinous, membranous sac-like apex (Fig. 84).
Larva (Fig. 85). Typical of the Leucotrichiini but with the following characteristics: head rugose or
papillate; pronota with antero-lateral angles not produced; femora with spiniform dorsal setae, tarsal
claw single; abdominal tergites II-VII smaller than in Zumatrichia, without central pores; tergite 1X
rarely with enlarged basal setae (after Flint, 1970).
Case (Fig. 86). Typically flat, silken, ovoid and slightly domed.
THE GENERA OF THE HYDROPTILIDAE 179
BioLoGcy. According to Wiggins (1977) the larvae occur on rocks in strong currents of running
water where they graze on the surrounding periphyton and detritus. Collection data indicate that
pictipes probably overwinters as a final instar larva and pupates from May to August.
SPECIES-GROUPS. Flint (1970) splits the genus into two main species-groups on the basis of adult
features.
The melleopicta-group is characterized by the three ocelliin both sexes, the unmodified male head
(except in chiriquiensis) and the single process of sternite VII in the male.
The pictipes-group is distinguished by the males having only two ocelli, a modified head (except
in imitator) and a brush of setae or a point on sternite VII.
Flint (1970) gives keys to the males of all species and to all described larvae.
Genus ZUMATRICHIA Mosely
Zumatrichia Mosely, 1937 : 187. Type-species: Zumatrichia filosa Mosely, by original designation.
DISTRIBUTION. North and Central America and the Antilles.
Zumatrichia contains 18 species of essentially Central American distribution although notosa
is known from Montana only and was originally described in Leucotrichia. The immature stages
are known only for antilliensis, anomaloptera and multisetosa.
Diacnosis. Adult. Forewing length 2:-5—4-0 mm: head unmodified; ocelli 2 (3), 3 (2); male basal antennal
segment elongate, enlarged, with a ‘button-like’ appendage which covers half of the slightly concave face
(cf. Fig. 91): mesoscutellum pentagonal: wings unmodified: spur formula 1.3.4: 3 genitalia (cf. Fig. 84)
with a short ventral process on sternite VII; sternite VII] produced beneath segment IX, sometimes with
a pair of lateral processes; segment IX open ventrally with a postero-lateral lobe or process; tergite X is
heavily sclerotized and sometimes produced posteriorly ; subgenital appendages large and plate-like, fused
with subgenital plate ventro-medially; inferior appendages usually fused baso-medially and often with a
baso-dorsal bracteole; aedeagus with median complex and membranous apex bearing spines and plates.
Larva (Fig. 87). Typical of the Leucotrichiini but distinguished by the rugose, but non-papillate, head;
simple pronota; femora with spiniform baso-dorsal setae (according to Wiggins, 1977, paired tarsal claws
are present in antilliensis); abdominal tergites larger than those of Leucotrichia and with paired contiguous
central pores; tergite IX covered in short, stout, spinous setae.
Case. Typically flat, silken (sometimes with embedded sand-grains), ovoid and dome-like (cf. Fig. 86).
BioLoGy. Wiggins (1977) states that the larvae inhabit fast-flowing sections of running-water
(preferring larger rivers according to Flint, 1968a). The larvae of antilliensis have successfully
adapted to living on boulders in the fast-flowing sections of the larger lowland rivers of Dominica
(Flint, 1968q).
SPECIES-GROUPS. Flint (1970) splits the genus into five species-groups (here reduced to four) on
the basis of features of the male genitalia.
The multisetosa-group (lobe of segment IX multisetate).
The galtena-group (sternite VIII without lateral processes, bracteole present).
The filosa-group (sternite VIII with lateral processes, no bracteole).
The palmara-group (sternite VIII with lateral and ventral processes, bracteole present; Flint,
1970, distinguishes anomaloptera and palmara on the colour and structure of the forewing only).
Genus PELTOPSYCHE Miller
Peltopsyche Miiller, 1879c : 144. Type-species: Pe/ltopsyche sieboldii Miiller, by subsequent designation by
Fischer, 1961.
DISTRIBUTION. Brazil.
This genus is known only from descriptions and crude figures of two species collected by
Miiller from the Santa Catarina region of southern Brazil. Although Miller (18795; 1879c; 1880)
gave a general account of the habitat and adults he only figured the general larval form and the
basal antennal segments of the males of both species. Thus, since Miiller’s original adult specimens
have not been traced, the genitalia and, therefore, the true identities of the two species remain un-
known. However, I have been able to examine larvae and cases of sieboldii which were sent to
McLachlan by Miiller and are now in the BMNH collection.
180 J. E. MARSHALL
91 epicranial ‘beak *
lateral ocellus
flagellum _
Pedicel
scape
Figs 87-92 Leucotrichiini. 87, Zumatrichia antilliensis Flint, larva, dorsal view (after Flint, 1970).
88-89, Peltopsyche sieboldii Miller, larva; (88) head and thorax, dorsal view; (89) thoracic legs,
lateral views. 90, Costatrichia lodora Mosely, 3, basal antennal segments; 91, Abtrichia squamosa
Mosely, 3, head, lateral view (after Mosely, 1939c); 92, Alisotrichia hirudopsis Flint, larva (fifth
instar), dorsal view (after Flint, 1970).
The actual spelling of the name of the type-species is in need of clarification. Miiller continually
used sieboldii except in a communication to McLachlan which was published in the Proceedings
of the Royal Entomological Society (Miller, 18795) where he used sieboldi. P. sieboldi was used by
Ulmer (1907) and Fischer (1961) lists the species under this name with all of Miiller’s references
to sieboldii being treated as synonyms even though Fischer was aware that sieboldii was the first
published name. I therefore propose to accept Miiller’s original orthography.
The following diagnosis is based on Miiller’s original descriptions with additional observations
on the larvae in the BMNH collection. The genus is clearly a member of the Leucotrichiini accord-
ing to the general appearance of the larvae and the modified male antennae.
DiaGnosis. Adult. According to Miiller the adults are distinguished by the modified basal segments of the
male antennae (13-segmented, longer in the female). In maclachlani the second segment is merely some-
what longer and wider than the others, but in sieboldii this segment is enlarged, ovoid and bears a small,
rounded process as in Zumatrichia. Miller suggests that these structures may have an ‘odoriferous’
function. The spur formula is quoted as 2.4.4 but this may be an error, the usual leucotrichiine count
being 1.3.4.
Larva (Figs 88, 89). According to Miiller’s descriptions and figures the abdomen of the larva is greatly
distended in segments V-—VII and fills nearly the entire case (cf. Figs 85, 87) and is thus typical of the
Leucotrichiini. Examination of the BMNH specimens of sieboldii (dried and originally glued to card)
THE GENERA OF THE HYDROPTILIDAE 181
reveals that the general morphology is very similar to that of Zumatrichia with respect to the head, legs,
thoracic nota, wide abdominal tergites (with their paired contiguous central pores) and the short, stout
spines of segment IX; unlike Z. antilliensis, however, the tarsal claws are unpaired, according to Wiggins
(1977).
Case. This is identical to that of Leucotrichia, being ovoid and flattened (‘leech egg-case’ case); the
dorsal surface is transversely ridged in siebo/dii and smooth in maclachlani.
Pupa. According to Miiller this is remarkable for the unusually great difference in the ‘complicated cor-
neus patches dorsally on the abdomen’ between the two species. He is probably referring to the pre- and
postsegmental dorsal abdominal plates but, since Miiller does not describe the specific differences, I
cannot comment on their significance.
BIoLoGy. The cases were found in very large numbers fixed to the upper side of stones in rapids
in the larger tributaries of the Itahajy (Garcia, Encana, Warnow, etc.), Santa Catarina. P.
maclachlani was only found in a single rapid near the mouth of the Warnow along with sieboldii.
Discussion. Peltopsyche may prove to be a senior synonym of one or more genera of the Leuco-
trichiini described subsequently by Mosely and Flint (see later discussion).
Since Miiller obviously encountered what he identified as siebo/dii in a number of localities
more than one species may be involved; this problem will only be resolved when Miiller’s original
material has been located and examined.
Genus ANCHITRICHIA Flint
Anchitrichia Flint, 1970 : 14. Type-species: Anchitrichia spangleri Flint, by original designation and
monotypy.
DISTRIBUTION. Central America (Mexico, Guatemala, Honduras, Costa Rica, Panama).
This genus contains only the type-species and is very closely related to Zumatrichia from which
it is distinguished by a number of adult features including its relatively larger size (4-5 mm fore-
wing length), its habit of holding its wings roof-like over the body, the unmodified antennae, the
prescence of 2 ocelli in both sexes, and the general form of the male genitalia (cf. Fig. 84). The
larva, described by Flint (1970), was not reared but associated ‘by supposition’ and is distinguished
by the bilobed anterior margin and postero-medial process of the frontoclypeus, the enlarged
antero-lateral angles of the pronota, the arborescent baso-dorsal seta of the femur and the form
of the abdominal tergites. The larval case is typical of the Leucotrichiini but has an enlarged,
transversely oval, protective hood at one end and a flared collar at the other. The pupal case is
unique in that it is quite unlike that of the larva, being torpedo-shaped and attached at one end
to the substrate by a silk strand 2-8 mm long. In this way the case floats just below the surface
of the water and may be an adaptation to prevent desiccation due to fluctuating water levels and
appears to parallel the condition seen in the larval case of Rhyacopsyche (Fig. 99).
The true status of Anchitrichia has yet to be established; the examination of more species is
required to define the limits of the genus. It may prove to be synonymous with one or more of the
other leucotrichiine genera.
Genus COSTATRICHIA Mosely
Costatrichia Mosely, 1937 : 166. Type-species: Costatrichia lodora Mosely, by original designation and
monotypy.
DISTRIBUTION. Central America.
Costatrichia was erected for a single Mexican species and now contains three additional species
from Panama (Flint, 1967a; 1970), one from Nicaragua and one from El Salvador (Flint, 1970).
The immature stages are unknown.
DraGnosis. Adult. Forewing length 2-0-2-5 mm; head unmodified; ocelli 3; basal antennal segments simple,
middle segments sometimes broad and flat (Fig. 90): metascutellum subtriangular: forewing often with
basal costal ‘bulla’ (Fig. 10): spur formula 1.3.4: ¢ genitalia (cf. Fig. 84) with lateral processes often on
sternite VIII; segment IX with setose postero-lateral processes; tergite X strongly sclerotized and fused
with segment IX; subgenital appendages elongate, separate, often with baso-dorsal bracteoles; aedeagus
with median complex and spinose membranous apex.
182 J. E. MARSHALL
SPECIES-GROUPS. Flint (1970) splits the genus into two distinct species-groups on the basis of
features of the male genitalia, head and wings.
The simplex-group (simplex, spinifera): unmodified antennae, no basal costal bulla.
The /odora-group (/odora, panamensis, tripartita, bipartita): modified antennae, basal costal
bulla present.
Flint (1970) gives a key to separate the males of all six species. He distinguishes the genus from
Zumatrichia by the presence of three ocelli and the unmodified basal segments, but sometimes
modified middle segments, of the male antennae.
Genus ACOSTATRICHIA Mosely
Acostatrichia Mosely, 1939c : 228. Type-species: Acostatrichia plaumanni Mosely, by original designation.
DISTRIBUTION. South America (Brazil, Surinam).
Mosely erected this genus for two south Brazilian species, plaumanni and simulans, and Flint
(1974) later described three new species from Surinam. The immature stages are unknown.
DiaGnosis. Adult. Forewing length 2:0-2:5 mm: head unmodified; ocelli 3; antennae unmodified; fore-
wing with basal costal pouch (cf. Fig. 9) filled with broadened hairs: spur formula 1.3.4: ¢ genitalia (cf.
Fig. 84) usually with two short ventral processes on segment VII; sternite VIII with postero-lateral pro-
cesses; inferior appendages sometimes with bracteoles; aedeagus with median complex and spinose mem-
branous apex.
Discussion. The three Surinamese species are said by Flint (1974) to resemble the two original
Mosely species although the basic form of the male genitalia is slightly different (but not enough
to warrant the erection of a new genus). According to Flint, fimbriata isclosest to brevipenis, while
spinifera resembles simulans and plaumanni. As Mosely (1939c) points out, the genus is closely
related to Costatrichia Mosely, differing only in features of wing venation and in having unmodified
antennae but, again as noted by Mosely, these may be specific rather than generic characteristics
(see later discussion). The larvae will probably prove to be of the general Leucotrichia type, as will
those of Costatrichia.
Genus BETRICHIA Mosely
Betrichia Mosely, 1939c : 230. Type-species: Betrichia zilbra Mosely, by original designation and
monotypy.
DISTRIBUTION. South America (Brazil, Argentina).
Erected for a single Brazilian species, this genus now also contains argentinica from north
Argentina and surinamensis, bispinosa and occidentalis from Surinam (Flint, 19726; 1974).
Mosely gives no precise reasons for erecting this genus but it may be assumed that it was mainly
on account of venational and antennal features as well as the general form of the male genitalia;
in the male type-specimen (BMNH) there are 2 ocelli, the antennae are 19-jointed with an elongate
basal segment. Flint (19725) places argentinica in this genus on genitalic features alone although
there are 3 ocelli and the antennae are simple. Of Flint’s (1974) Surinamese species only occiden-
talis resembles the type-species, while in surinamensis the head is greatly modified and the genitalia
are quite different; in bispinosa, although the genitalia resemble those of zi/bra, the forewing has a
basal costal pouch. Flint (1974) states that ‘he would prefer to wait until the South American
fauna is better known... before erecting more genera’ but I am of the opinion that all the small
leucotrichiine genera, such as Betrichia, Costatrichia and Acostatrichia for example, should be
grouped together as the characters originally given as diagnostic generic features are now proving
to be specific instead ; this, however, will be discussed more fully later. As such no precise diagnosis
of the genus Betrichia can be given; only the main features are listed below. The immature stages
are unknown.
DiaGnosis. Adult. Forewing length 2-4 mm: head and antennae slightly modified or simple; ocelli 2 or 3:
forewing with or without basal costal pouch: spur formula 1.3.4: 3 genitalia (cf. Fig. 84) with a process on
sternite VII; sternite VIII usually without postero-lateral processes: segment IX with or without postero-
lateral processes; inferior appendages fused medially; tergite X and subgenital plate variable; aedeagus
with median complex and spinose membranous apex.
THE GENERA OF THE HYDROPTILIDAE 183
Discussion. As is evident from the above account this genus is very difficult to key out satisfac-
torily. As with a number of other genera in the Leucotrichiini all species descriptions must be
referred to before a specimen suspected of belonging to Betrichia can be positively identified.
Genus ABTRICHIA Mosely
Abtrichia Mosely, 1939c : 224. Type-species: Abtrichia antennata Mosely, by original designation.
DISTRIBUTION. South America (Brazil, Argentina).
Abtrichia was erected for two very distinct species from southern Brazil, antennata and squa-
mosa, of which the former has since been recorded, along with its immature stages, from northern
Argentina by Flint (19720). Abtrichia is characterized by the highly modified head of the male with
the enlarged, process-bearing, basal antennal segment (Fig. 91) and the large basal costal pouches
of the male forewings which contain small scales or coarse hairs (Fig. 9). According to Flint
(19725), the larvae bear most resemblance to those of Zumatrichia antilliensis, differing only in
having a papillate dorsal head surface.
DiaGnosis. Adult. Forewing length 4 mm: head of male modified with dorsal and ventral anterior beak-
like processes (Fig. 91); ocelli 2 (both sexes); antennae 12-jointed in the male with an enlarged, process-
bearing, basal segment (Fig. 91): forewing with costal pouch along basal half (Fig. 9): spur formula 1.3.4:
3 genitalia (cf. Fig. 84) with a short process on sternite VII; sternite VIII and segment IX without postero-
lateral processes; subgenital appendages well developed; inferior appendages distinct, with or without
bracteoles; aedeagus with median complex and spinose membranous apex.
Larva and case. Typical of the Leucotrichia-group.
BioLocy. According to Flint (19725) the cases are tightly attached to rocks in riffles.
Genus CELAENOTRICHIA Mosely
Celaenotrichia Mosely, 1934 : 158. Type-epecies: Celaenotrichia edwardsi Mosely, by original designation
and monotypy.
DISTRIBUTION. South America (Chile).
C. edwardsi was originally described from Chiloe Island and Flint and Barria have since collec-
ted additional specimens (USNM) of which I have examined a female example. From the adult
features the genus appears to belong in the Leucotrichiini; it is characterized by the unmodified
antennae and forewings and the distinctive genitalia of the male. The immature stages are un-
known.
DiaGcnosis. Adult. Forewing length 3 mm: head and antennae simple, latter 26-segmentced; ocelli 3:
forewings unmodified: mesoscutellum with transverse suture; metascutellum pentagonal: spur formula
1.3.4: ¢ genitalia relatively simple (cf. Fig. 84) with a process on sternite VII; sternite VIII produced
postero-ventrally, without postero-lateral processes; segment IX open ventrally with deep postero-dorsal
V-shaped cleft enclosing the membranous tergite X which has two narrow longitudinal supporting scler-
ites; subgenital appendages heavily sclerotized and spinose; subgenital plate well developed; inferior
appendages curved inwards, without bracteoles; aedeagus appears to have a median complex (microscope
slide preparation of the type-specimen (BMNH) indistinct here), apex membranous and spinose.
Genus ALISOTRICHIA Flint
Alisotrichia Flint, 1964 : 46. Type-species: Alisotrichia hirudopsis Flint, by original designation and
monotypy.
DISTRIBUTION. Central America (including the Antilles).
Alisotrichia contains 12 species, three of which are also known in their larval stages, and is a
member of the Leucotrichiini but forms a distinct subgroup according to both adult and larval
features. The insects are very small and, although adult characters are somewhat variable, cons-
tant diagnostic features include the absence of a fore-tibial spur, the simplified aedeagus and
modified tergite VII of the male and the simple internal apparatus of the female genitalia. The
larvae are unique within the tribe in that the fifth instars remain free-living until pupation and
thus retain the general appearance of the earlier instars of this and other leucotrichiine genera
(cf. Ugandatrichia). The following diagnosis is adapted from Flint (1970).
184 J. E. MARSHALL
DiaGnosis. Adult. Forewing length 1-2—2-5 mm: basal antennal segment of male often enlarged; ocelli
2 or 3: metascutellum pentagonal to subtriangular: spur formula 0.2.4, 0.2.3 or 0.3.4: 2 genitalia simple,
tergite VII modified (no details or figures given by Flint, 1970), internal apparatus with a simple ring or
sphere: ¢ genitalia (cf. Fig. 84) with sternite VIII and segment IX often with postero-lateral processes;
inferior appendages, tergite X, subgenital appendages and plate very variable and difficult to interpret;
aedeagus simple with a median constriction and a simple to complex internal structure but never with a
midlength complex (basal loop, ‘windows’, etc.).
Larva (Fig. 92). Abdomen not distended, tapering posteriorly; tergites I-VIII covering most of dorsum
of each segment and each with three contiguous central pores, tergite IX shield-shaped without pores;
anal prolegs long, projecting freely from segment X.
Pupal case (after Flint, 1964, for hirudopsis). This is built just prior to pupation and is similar to the
typical leucotrichiine larval and pupal case. It is silken, dorso-ventrally flattened, with an ovoid central
region (containing the pupa) which is surrounded by an irregular flange which attaches the case to the sub-
strate and has two to six small round openings (for water circulation ? (Flint, 1964)).
BioLoGy. The larvae of hirudopsis are found on rocks in fast-flowing water at various altitudes
and occur abundantly on exposed boulders kept wet by the spray from nearby cascades. The
adults are also abundant and run about in the sunlight on the dry exposed parts of the rocks on
which the larvae live.
SPECIES-GROUPS. Flint (1970) splits the genus into several groups of species on the basis of adult
characters such as the number of ocelli, spur formula and antennal modifications. However, as
Flint does not define the groups clearly, I have not discussed them here.
Discussion. The fifth instar larva of Alisotrichia conforms to the basic leucotrichiine plan but
retains the free-living adaptations of the earlier instars; on the basis of this and the characteristic
adult features this genus would appear to be a specialized and distinct member of the Leucotrichiini
and is placed here in its own subgroup.
Tribe OCHROTRICHIINI trib. n.
Type-genus: Ochrotrichia Mosely, 1934.
DISTRIBUTION. North and Central America.
The tribe Ochrotrichiini is proposed here for the small group of Nearctic and Neotropical
hydroptilids composed of Ochrotrichia (O. Ochrotrichia and O. Metrichia) and Rhyacopsyche. As
will be discussed in more detail below, Metrichia was reduced to a subgenus of Ochrotrichia by
Flint (1972a) on the larval affinities, Flint (1971) having previously indicated the relationships of
the adults of Metrichia and Rhyacopsyche. The relationships of these genera have not been
thoroughly investigated due to the lack of comparative material. Flint’s views have therefore been
accepted and, as the group exhibits a number of characteristic adult features which distinguish
it from the other hydroptiline groups, it has been accorded the status of tribe. However, this
arrangement is not entirely satisfactory and the group may subsequently prove to be a subgroup
of the tribe Hydroptilini. This will be discussed in more detail in the discussion below.
DiaGnosis. Adu/t. Forewing length 1-5-3-5 mm: head and antennae unmodified (Fig. 93); tentorium
complete; ocelli 3; postoccipital lobes small, subspherical: mesoscutellum with transverse suture (Fig.
93); metascutellum pentagonal or convexly subtriangular: spur formula 0.3.4 or 1.3.4: abdomen with
typical hydroptiline setate processes of segment V: 2 genitalia a simple oviscapt: ¢ genitalia variable but
basically with segment VIII unmodified and inferior appendages well developed and elongate.
Larva (Figs 97-99). General appearance typical of basic, unspecialized hydroptilid: head unmodified,
labrum symmetrical: three pairs of thoracic nota, each divided by a distinct median line: abdomen dis-
tended, slightly compressed; abdominal tergites absent except on segment IX; anal prolegs fused to sides
of segment X.
Case. Variable between genera: basic ‘purse-type’ hydroptilid case in Ochrotrichia, usually covered in
fine sand grains; modified in Rhyacopsyche as an elongate, tubular case tapering at both ends and attached
at one end to the substrate by a short silken thread.
Pupa. No description available, but probably of the typical hydroptiline form with long, untoothed
mandibles.
THE GENERA OF THE HYDROPTILIDAE 185
Discussion. The Ochrotrichiini is a distinct, warm-adapted, New World group of the subfamily
Hydroptilinae. It may be distinguished from the Leucotrichiini, with wlsich it shares the feature
of a transverse mesoscutellar suture, by the characteristic and highly specialized male genitalia
and the basic hydroptilid appearance of the larva. The tribe may later prove to be a subgroup of
the Hydroptilini but is provisionally treated here as distinct on the basis of certain fundamental
morphological and behavioural features. These include the presence of a transverse mesoscutellar
suture and the characteristic male and female genitalia (including the absence of a spiral ‘titillator’
on the male aedeagus) and the associations of the larvae with running water habitats and their
possible detritus-feeding habits (cf. the predominantly algal-feeding habits of the Hydroptilini).
However, such features may be secondarily derived from the basic Hydroptilini condition, a
thorough examination of representative adult and larval material being essential for the establish-
ment of the true relationships of the group within the Hydroptilinae and of the genera with one
another.
Genus OCHROTRICHIA Mosely
Ochrotrichia Mosely, 1934 : 162. Type-species: Ochrotrichia insularis Mosely, by original designation and
monotypy.
DISTRIBUTION. North, Central and South America.
The genus Ochrotrichia has recently been reviewed by Denning & Blickle (1972) who list the 40
known species and give descriptions of 15 additional new species from the western United States
and Mexico. In the same year Flint (1972a) published a paper on ‘The genus Ochrotrichia from
Mexico and Central America (Trichoptera : Hydroptilidae)’ in which he described a number of
new species and proposed Metrichia as a subgenus of Ochrotrichia, and Rhyacopsyche as a close,
but distinct, relative. Flint (19685) had already stated his views regarding the congenericity of
Metrichia and Ochrotrichia but they were not accepted by Denning & Blickle (1972). As it has
not been possible to investigate this question in detail due to the paucity of material in the BMNH
collections, Flint’s views have been accepted here. There is obviously a need for a thorough and
critical review of the group to establish the relationships of the genera concerned and to investigate
the position of the group within the Hydroptilinae.
DiaGnosis (Ochrotrichia sensu Flint, 1972a). Adult. Forewing length 1-5-3-0 mm: head unmodified;
tentorium complete (Fig. 93); antennae simple (c. 33-segmented in insularis); mesoscutellum (Fig. 93)
with transverse suture; metascutellum convexly subtriangular: spur formula 0.3.4 or 1.3.4: 2 genitalia a
simple oviscapt; internal surface of sternite VIII often sclerotized and reticulate in Ochrotrichia (Metrichia
not described): 3 genitalia distinct (Figs 94-96) with segment VIII unmodified; segment IX fused, annular,
deeply incised dorsally; tergite X often highly developed with numerous spines and processes; inferior
appendages usually strongly developed, basically broad and elongate, usually with various processes and
rows of spines; aedeagus simple and thread-like (Ochrotrichia), sometimes armed with strong spines
(Metrichia).
Larva (Fig. 97) after Flint (1972a). General appearance typically hydroptilid: head unmodified; mandi-
bles robust, broadly subtriangular, medial brush in left only: meso- and metanota with pronounced
antero-dorsal processes; prothorax with a pair of anal sternites (Fig. 44, after Wiggins, 1977); pleurites
well developed (at least on prothorax): abdomen similar to that of Hydroptila (cf. Fig. 118) being slightly
distended and laterally compressed; dorsal ‘rings’ present.
Case. Similar to that of Hydroptila; constructed of two laterally compressed silken valves, ‘purse-type’,
covered in fine sand-grains and, occasionally, filamentous algae (Wiggins, 1977); sometimes the case con-
sists of a single dorsal convex valve carried ‘tortoise-shell’ like, the ventral valve being flat and of secretion
only.
Pupa. No description available.
BIoLoGy. The larvae occur in a wide variety of running water habitats (rivers to warm streams
and cold springs) and temporary streams according to Ross (1944); the specimens with cases
carried ‘tortoise-shell’ like were from a spring stream on rocks in a thin film of water; the feeding
habits are unknown but it is possible that the larvae are detritus feeders.
186 J. E. MARSHALL
Key to subgenera (adults) (adapted from Flint, 1972a)
1 Spur formula 0.3.4; 3 inferior appendages and tergite X complex, aedeagus simple and thread-
like : ; ; : : : : ; : : . OCHROTRICHIA (p. 186)
— Spur formula 1.3.4; ¢ inferior appendages and tergite X simple, aedeagus with well-developed
spines : , ; ; : ‘ : : ; : : . METRICHIA (p. 186)
Subgenus OCHROTRICHIA Mosely
Polytrichia Sibley, 1926 : 102. Type-species: Ithytrichia confusa Morton, by monotypy. [Preoccupied by
Polytrichia Borg de St Vincent, 1831 in Protozoa.]
Ochrotrichia Mosely, 1934 : 162. Type-species: Ochrotrichia insularis Mosely, by original designation and
monotypy. [Synonymized with Polytrichia by Mosely, 1937 and reinstated by Ross, 1944.]
This subgenus is distributed throughout the U.S.A., with a few species recorded from southern
Canada, Panama and the Antilles, although the group appears to be predominant in the midwest
and montane regions of the United States.
DiaGcnosis. Adult. Spur formula 0.3.4; 3 genitalia with inferior appendages and tergite X often very com-
plex, aedeagus simple and thread-like.
The subgenus contains about 70 species which Flint (1972a) splits into two distinct groups (for
the Central American species only). The xena-group is characterized by the male genitalia with
its relatively simple segment IX and tergite X and the aedeagus, which varies from a simple tube
to one which bears various processes (not as well developed as in Metrichia). It appears to be the
simpler of the two groups with respect to the structure of the male genitalia and also seems to be
intermediate between the next group and Metrichia. The group contains xena, unio, flagellata,
pectinata, brayi, caligula, marica, spinosissima and verda.
The second group consists of all other Central American and Antillean species which Flint
(1972a) further splits into six subgroups; the group is characterized by the fusion of tergites IX
and X in the male genitalia to form a complex structure bearing many spines and plates (Fig. 95,
terminology after Ross, 19415), and also the very simple, thread-like aedeagus. A key to the males
of the Central American species is given by Flint (1972a).
Larvae have been associated with the following species: anisca, riesi, spinosa, tarsalis, unio and
xena, a key to which is given by Ross (1944). According to Flint (1972a), the larvae of this sub-
genus are found in running water, usually in small to moderately sized streams, sometimes in very
shallow water and often in streams that dry up during the dry season. Flint also remarks that the
adults come readily to light unlike those of Metrichia.
Subgenus METRICHIA Ross
Metrichia Ross, 1938 : 9. Type-species: Orthotrichia nigritta Banks, by monotypy.
Argentitrichia Jacquemart, 1963 : 339. Type-species: Argentitrichia bulbosa Jacquemart, by monotypy.
Syn. n.
This subgenus has a more southerly distribution than Ochrotrichia, occurring in the south-west
United States, throughout Central America and the Antilles, Peru, north-west Argentina, central
Chile and Surinam. According to Flint (1972a) the area of greatest diversity occurs throughout
Central America and the Antilles.
DiaGnosis. Adult. Spur formula 1.3.4; ¢ inferior appendages and tergite X simple, aedeagus with well-
developed spines; 3 with internal abdominal sacs (between IV-V, V-VI or VI-VIII) and dorso-lateral
hair brushes on abdominal segments V and VI.
There are 19 distinct species of which Flint (1972a) splits the Central American representatives
into five groups on the basis of the presence and positioning of the internal abdominal sacs of the
males, the dorso-lateral hair brushes and the structure of the aedeagus (Fig. 96). Flint also gives
a key to the males of the Central American species.
Larvae have been associated with nigritta by Edwards & Arnold (1961) and juana by Flint
(1968a). According to Flint (1972a) the larvae build ‘purse-type’ cases of silk and organic particles.
On the basis of features of the two known species, Flint concludes that the subgenus appears to
THE GENERA OF THE HYDROPTILIDAE 187
97
anterolateral process
Figs 93-100 Ochrotrichiini. 93, adult head and thorax, dorsal view, Ochrotrichia. 94-96, generalized
3 genitalia. 94-95, Ochrotrichia (Ochrotrichia); (94) lateral view; (95) tenth tergite, dorsal view
(terminology after Ross, 19415). 96, O. (Metrichia), aedeagus, lateral view (vide Figs 15, 16).
97-98, Ochrotrichia (O.) jauna Flint; (97) larva, lateral view; (98) case, lateral view. 99-100,
Rhyacopsyche mexicana (Flint); (99) larva in case, lateral view; (100) larval foretibio-tarsus, lateral
view.
be closely associated with springs, seeps, waterfalls and the like (rather than the larger, relatively
more slowly flowing streams preferred by Ochrotrichia). Flint also states that the adults are more
frequently taken by sweeping vegetation than at light.
Flint (1972a), following Schmid (1958a), considers Metrichia to be most closely related to the
Palaearctic genus Microptila (sensu Schmid, vide the Hydroptilini, Agraylea-group) on the basis
of the form of the male genitalia and wing venation. Healsorecognizes the possibility that Microp-
tila may eventually come to be considered as a third subgenus of Ochrotrichia. As will be seen
in the account of Microptila, Metrichia is really very dissimilar to and quite distantly related to
Microptila. These conclusions were based on a comparative study of features of the head, thorax
and male genitalia of the two groups.
Finally, Argentitrichia bulbosa, described by Jacquemart (1963) from the Argentine, appears to
be a species of Metrichia according to the characteristic form of the male genitalia (material not
examined).
Genus RHYACOPSYCHE Miller
Rhyacopsyche Miiller, 1879a : 40. Type-species: Rhyacopsyche hagenii Miller, by subsequent monotypy.
Rhyacopsyche Miiller, 1879c : 143.
DISTRIBUTION. Central America.
Miiller (1879a) erected Rhyacopsyche for a single Brazilian species, which he subsequently
188 J. E. MARSHALL
(1879c) named hagenii, on the basis of the larval cases only. Figures and full descriptions of the
larval and pupal cases were not published until 1880 by Miiller, the first descriptions of the adults
and larvae being given by Thienemann (1905). Mller (1921) subsequently reproduced a previously
unpublished figure by Miiller of the general appearance of the larva.
Following the discovery of immature and adult stages of Metrichia mexicana in Guatemala
(first described from Mexico by Flint, 1967a), Flint (1971) considers this species to be congeneric
with Rhyacopsyche hagenii. Consequently Flint (1971) described the genus Rhyacopsyche and
gave a key to the males of mexicana and three new species.
D1aGnosis (after Flint, 1971). Adult. Forewing length 2-5-3-5 mm: head and antennae simple; ocelli 3:
mesoscutellum with transverse suture (cf. Fig. 93); metascutellum pentagonal: spur formula 1.3.4: 9
genitalia a simple oviscapt, tergite VIII with posterior lobes; 3 genitalia with segment VIII unmodified;
IX produced into dorso-lateral lobes; tergite X contracted inside lobes of IX; inferior appendages large
and distinct, elongate and curved upwards; single, reduced, median subgenital sclerite; aedeagus with
tubular basal half, apical half with central tube and thin spiral filament, apex with thickened spine.
Larva (Fig. 99). Typically hydroptilid with thoracic nota present on all three segments, divided medially;
sternites apparently absent; legs short, subequal; apex of fore-tibia with a distinct process bearing several
enlarged setae; tarsus short with one or two enlarged, plate-like setae (Fig. 100): abdomen distended,
slightly compressed; tergites absent; anal prolegs fused to segment X.
Case (Fig. 99). Larval case elongate, tubular, tapering at both ends; silken, covered in sand grains
(mexicana) or of secretion only (Aagenii); attached to substrate by a long silken thread from the anterior
end. Pupal case of mexicana apparently split along posterior end according to the figure in Flint (1971),
but this may have been drawn after emergence of the pharate adult; pupal cases of both mexicana and
hagenii anchored by a short silken stalk.
BioLoGy. The larvae and pupae of mexicana were found on an exposed boulder subjected to con-
stant wetting from spray and a thin film of water running down from a nearby cascade. The pupal
cases were attached at right angles to the boulder on the moist, but not submerged, surfaces.
According to Flint (1971) the larval cases of mexicana, unlike those of hagenii, are not attached
by a thread to the substrate until just prior to pupation.
Discussion. According to Flint (1971) the adults of Rhyacopsyche may be distinguished from those
of Metrichia only by the form of the male and female genitalia (the lobes of segment IX and the
relatively simple aedeagus of the male and the lobes of tergite VIII of the female in Rhyacopsyche).
The larvae of Rhyacopsyche are, according to Flint (1971), very similar to those of Ochrotrichia
s.1. and Hydroptila, being distinguished by the plate-like setae of the fore-tibia and the hook-like
tarsal claws. The larval and pupal cases are, however, so far unique to Rhyacopsyche and are
probably adapted to exposed conditions in fast-flowing water, being fusiform and anchored by a
silk thread to the substrate.
The morphology of the larva has not been described in sufficient detail to allow adeeper investi-
gation into the generic relationships of Rhyacopsyche but adult features suggest that the genus is
closely allied to Ochrotrichia s.1. I do not agree with Schmid (1958a) and Flint (1971) that these
genera are closely related to Microptila (q.v.).
Tribe NEOTRICHIINI Ross
Neotrichiini Ross, 1956 : 18. Type-genus: Neotrichia Morton, 1905.
DISTRIBUTION. Nearctic, Neotropical.
Ross first used the name Neotrichiini in a phylogenetic diagram (Ross, 1956 : 18, chart II)
showing the coincidence of the primitive lines of Trichoptera with the ‘cool-adapted habitat’, the
Neotrichiini representing a specialized ‘warm-adapted’ offshoot of the main Hydroptilini branch.
Ross, however, did not then or subsequently define the tribe although he is still of the opinion
that the group should be maintained as distinct from the rest of the Hydroptilidae, principally
on account of the characteristic form of the larvae and their ‘Limnephilid-like’ cases (Ross, in
litt.). Therefore, although Ross’s (1956) classificatory scheme has been modified here in that the
Hydroptilini sensu Ross are treated as a subfamily, the status of the Neotrichiini is retained as
originally proposed.
THE GENERA OF THE HYDROPTILIDAE 189
The Neotrichiini consists of two exclusively warm-adapted New World genera, Neotrichia and
Mayatrichia, the combined distribution of which extends from Canada, through the United States
and Central America (the regions of greatest species and individual numbers) down into central
Argentina. The genera share a number of unique adult and larval characters which distinguish
them from all other Hydroptilinae, although they still retain the basic diagnostic features of this
subfamily. As indicated above, the immature stages characterize the Neotrichiini since the larvae
construct cylindrical cases, with circular anterior and posterior openings, reminiscent of the higher
limnephiloid groups and quite unlike the basic ‘purse-type’ case of the Hydroptilidae. The larvae
accordingly exhibit parallel limnephiloid-like morphological features associated with the form of
the case. These include the cylindrical abdomen and the free anal prolegs which may be extended
through the wide posterior opening of the case, and in one species of Neotrichia a lateral line
appears to be present.
The following diagnosis is based on Ross (1944) and examination of adult material in the BMNH
collection and larvae of both genera (USNM loan).
DiaGnosis. Adult. Forewing length 1-5-2-0 mm: head and antennae simple; tentorium complete; ocelli 3:
mesoscutellum without transverse suture, anterior margin convex; metascutellum convexly subtriangular:
fore-tibia without apical spur: abdomen with typical lateral setate processes on sternite V: 9 genitalia a
simple oviscapt: ¢ genitalia (Fig. 101)-—segment VIII unmodified; IX fused, annular, open postero-
ventrally, well-developed postero-lateral processes present; inferior appendages and subgenital plate well
developed; bilobed process sometimes present and often may be fused to the underside of the subgenital
plate in some species of Neotrichia; tergite X usually membranous with various lobes; subgenital appen-
dages present (homologies uncertain).
Larva (Figs 102, 104-106). Head conical, tapering anteriorly, antennae relatively long (cf. other Hydrop-
tilidae): thoracic nota paired: abdomen only slightly distended, cylindrical, abruptly enlarged from
segment II, from there gradually tapering posteriorly; tergites VIII and IX present only, latter covering
whole of dorsum; anal prolegs slender, long, projecting freely from segment X; lateral fringe of short
setae (‘lateral line’) sometimes present along sides of abdomen: thoracic legs slender, tarsal claws long.
Case. Cylindrical, slightly tapering posteriorly, constructed of secretion only (surface strongly ridged)
(Fig. 107) or with a covering of small sand-grains (Fig. 103); openings suboval to circular, wider
anteriorly.
BioLoGy. Larvae occur on rocks in rapid sections of rivers and streams; details of feeding habits
unknown, but probably detritus-feeders; according to Wiggins (1977) the guts of three specimens
of Mayatrichia contained almost exclusively fine organic particles although, from the shape of the
head, he suspects them to have specialized feeding habits.
Discussion. The Neotrichiini is exclusively a New World group containing two very closely re-
lated genera which are distinguished from each other by features of the male genitalia, adult spur
formula and the general form of the larva. The structure of the adult thorax and presence of the
spiral aedeagal process in the males of Neotrichia indicate that the group may have arisen relatively
recently from the main Hydroptilini branch and evolved its characteristic features in isolation on
the American continent. The most notable feature, apart from their minute size, is the distinctive
appearance of the larvae which seem to have developed characteristics similar to those of the
limnephiloid groups in association with the possession of cylindrical cases. As Ross (1956) points
out, however, it isunlikely that the Neotrichiini gave rise to the limnephiloid branch of the Trichop-
tera since this tribe is a very specialized member of the Hydroptilinae, the superficial similarities
having arisen by parallel evolution. In addition, the larvae do not seem to have the habit of feeding
on filamentous green algae, as is characteristic of the Hydroptilini, and are supposed to be detritus-
feeders. From the form of the larval mandibles, which are robust and not markedly dentate or
asymmetrical, the detritus-feeding habit would appear to be a primary characteristic and not
secondarily derived from the algal-feeding condition (cf. /thytrichia). The position of the Neo-
trichiini within the Hydroptilidae will be discussed later.
Genus NEOTRICHIA Morton
Cyllene Chambers, 1873 : 124. Type-species: Cyllene minutisimella Chambers, by monotypy. [Preoccupied
several times, vide Fischer, 1961.]
190 J. E. MARSHALL
101
‘rostrum’
anal proleg
=—*t— attachment
a
enenenccvsancapc wr CCTAHiCcttt WHOMMLLGLE IMC CCY
Po cate A
ew ite elarea”,
tubular case :
Figs 101-107 Neotrichiini. 101, 3 genitalia, lateral view, Neotrichia (vide Figs 15, 16). 102-104,
Neotrichia sp.; (102) larva, lateral view; (103) case, lateral view; (104) head, larva, dorsal view.
105-107, Mayatrichia ayama Mosely; (105) head, larva, dorsal view; (106) anal proleg, larva,
lateral view; (107) larval and pupal cases, lateral views.
Neotrichia Morton, 1905 : 72. Type-species: Neotrichia collata Morton, by monotypy.
Exitrichia Mosely, 1937 : 170. Type-species: Exitrichia anahua Mosely, by original designation. [Synony-
mized by Ross, 1944.]
Dolotrichia Mosely, 1937: 177. Type-species: Dolotrichia canixa Mosely, by original designation. [Synony-
mized by Ross, 1944.]
Guerrotrichia Mosely, 1937 : 179. Type-species: Guerrotrichia caxima Mosely, by original designation.
[Synonymized by Ross, 1944.]
Lorotrichia Mosely, 1937 : 181. Type-species: Lorotrichia hiaspa Mosely, by original designation and
monotypy. [Synonymized by Ross, 1944.]
DISTRIBUTION. Nearctic, Neotropical.
Neotrichia is exclusively a New World genus with representatives occurring most abundantly
in the U.S.A. and Mexico, although a few species have been recorded from Canada, the Antilles
Surinam and Brazil. There are 46 species which fall into a number of distinct groups which are
characterized by features of the male genitalia and which correspond with the genera described
by Mosely (1937) and subsequently synonymized by Ross (1944). The Surinamese species (Flint,
1974) appear to fall into two additional groups.
According to Ross (1944) the larvae of minutisimella, okopa, collata and riegeli are known but
only those of the first named have been described. However, Flint (1964; 19685; 1968c) has since
described the larva of iridescens.
DiaGnosis. Adult. Spur formula 0.2.3: ¢ genitalia (Fig. 101) sometimes with ventral process on sternite
VIII; segment IX annular, generally complete dorsally and ventrally; tergite X variable but usually form-
ing a well-developed dorsal plate; inferior appendages distinct, somewhat elongate, usually with promi-
nent, spatulate bracteole (Br); subgenital plate well developed with a pair of apico-ventral setate lobes
(representing fused bilobed process ?); a pair of narrow subgenital appendages sometimes present (‘slender
structure’ (Flint, 1964); ‘filiform appendage’ (Ross, 19415)); aedeagus elongate, generally with a wide
tubular base narrowing to a median constriction or neck from which arises a stout spiral process, apical
half more slender, divided at apex or bearing apical spines.
Larva (Figs 102, 104). Head cone-shaped, tapering anteriorly; labrum symmetrical; mandibles sub-
symmetrical, short, robust, not markedly dentate: thoracic nota covered in long sparse setae; legs long
THE GENERA OF THE HYDROPTILIDAE 191
and slender (cf. Mayatrichia): abdomen cylindrical but not as rotund as in Mayatrichia, intersegmental
grooves distinct.
Case (Fig. 103). Cylindrical, tapering posteriorly, basically of silk and often covered in small sand-
grains.
BioLoGcy. Larvae found in rapid sections of rivers and streams.
Genus MAYATRICHIA Mosely
Mayatrichia Mosely, 1937 : 182. Type-species: Mayatrichia ayama Mosely, by original designation.
DISTRIBUTION. Nearctic.
There are only four species in this genus which are all of local occurrence but have an overall
distribution ranging from Saskatchewan to Mexico and from Ontario to Maine. Ross (1944 : 278)
gives a key to the males of the three North American species (ayama, ponta and acuna) with
rualda from Mexico being described by Mosely (1937 : 183). The larvae and cases of ayama (in-
cluding an early instar) and ponta have been described by Ross (1944) and Wiggins (1977)
respectively.
Diacnosis. Adult. Spur formula 0.2.4: 3 genitalia with sternite VI with a long, slender ventral process;
segment IX annular, deeply incised along postero-ventral margin, accentuated by postero-lateral pro-
cesses; tergite X membranous, hood-like; inferior appendages with broad, setae-bearing posterior margin
and a small, digitate, dorso-lateral projection; subgenital plate well developed with postero-medial,
ventrally directed processes and produced dorso-laterally to meet tergite X; bilobed process absent;
aedeagus simple, long, apex filiform or with three ‘staggered tubercules’ (Ross, 1944).
Larva (Figs 105, 106). Head very attenuate anteriorly: thoracic nota covered in short, stout setae; legs
relatively shorter and more robust than in Neotrichia: abdomen rotund, cylindrical, with faint inter-
segmental grooves; lateral setal fringe present only in ayama.
Case (Fig. 107). Cylindrical, tapering posteriorly, constructed of secretion only but with strengthening
transverse or longitudinal ridges.
BioLoGcy. According to Wiggins (1977) the larvae occur on rocks in rapid sections of rivers and
streams, generally in rather large bodies of running water.
Tribe HYDROPTILINI Stephens
Hydroptilidae Stephens, 1836 : 151. Type genus: Hydroptila Dalman, 1819.
DISTRIBUTION. Cosmopolitan (excluding polar regions).
The Hydroptilini, as interpreted here, contains the most widely distributed and successful
contemporary genera of the subfamily Hydroptilinae. Although the tribe may appear to be very
heterogeneous when considering the variety of both adult and larval morphological features,
habits and habitat preferences, all members do have a number of basic affinities. These include
features of the adult thorax and the male genitalia, and also the fundamental association of the
larvae with green filamentous algae in more slowly moving bodies of water. Owing to the diversity
of the group, however, it is difficult to give a precise diagnosis of the tribe. The following is only
a preliminary attempt at a formal diagnosis of the tribe, since this must await further more detailed
research into the possible relationships of the genera and the discovery of all the larval stages.
Diacnosis. Adult. Forewing length 1-5—6-0 mm. Head basically unmodified (e.g. Fig. 108); antennae simple;
ocelli 3 or 0; postoccipital lobes variable, modified in Hydroptila (3) as ‘scent caps’; tentorium variable,
distinct or reduced: shape of thoracic nota variable (Figs 108, 139); mesoscutellum without transverse
suture, diamond-shaped or with convex anterior edge; metascutellum pentagonal or with convex anterior
edge; spur formula 0.2.3, 0.2.4, 0.3.4 or 1.3.4: lateral setate process of abdominal segment V present, pit
sometimes sculptured: ? genitalia (Figs 17, 121) basically a simple telescopic oviscapt, sometimes fused and
non-retractile: ¢ genitalia very variable with form characteristic for each genus (Figs 109, 115, 122),
aedeagus usually with spiral ‘titillator’.
Larva. The larvae of Microptila, Dhatrichia, Stenoxyethira and Xuthotrichia have not yet been described.
Basically of the typical hydroptilid ‘purse-case’ building form. Head unmodified, mouthparts usually
showing various degrees of specialization associated with feeding on the cell contents of green filamentous
algae; lateral line and gills absent, body often slightly compressed laterally, anal prolegs fused to sides of
segment X, abdominal tergites usually absent, ‘dorsal rings’ present. The forms of the larva and its case
192 J. E. MARSHALL
vary according to habits and habitat preferences. Substrate dwellers such as Hydroptila retain the gen-
eralized hydroptilid form and their ‘purse-type’ cases often incorporate sand-grains and diatoms, possibly
to aid ballast. Larvae found amongst aquatic vegetation are usually characterized by their lighter cases,
often of secretion only, and the elongation of the mid- and hindlegs. The recently described larva of
Ugandatrichia is very specialized, being adapted for life in torrenticolous conditions. Many of its features
parallel those of Stactobia and leucotrichiine larvae, for example, in that it is strongly dorso-ventrally
flattened and heavily sclerotized, with well-developed abdominal tergites. Ugandatrichia is also unique in
that the fifth instar larva does not build a case until just prior to pupation.
Pupa. Typically hydroptilid, mandibles long and without teeth.
DiscussiON. The tribe Hydroptilini is considered here to consist of three subgroups which are
distinguished by male and female genitalic characters and the general appearance and habits of
the larvae. Within each subgroup the genera variously exhibit both generalized and specialized
hydroptilid characteristics. The subgroups are defined below.
The Agraylea-group. This contains some of the larger and more generalized hydroptilids with
respect to the overall appearance of the adults. The male genitalia, however, are rather distinctive
and conform to a common basic pattern within the subgroup. The larvae of Agraylea are essen-
tially adapted for life in vegetation thickets in slow-moving to static bodies of water, whereas those
of Ugandatrichia are adapted for life in torrenticolous habitats and are free-living in the fifth
instar.
The Hydroptila-group. This essentially consists of the nominate genus Hydroptila with which a
number of genera have here been synonymized. The adults exhibit generalized but distinctive
patterns of genitalia which vary slightly between species-groups; the larvae are basically typical
of the hypothetical generalized hydroptilid but are adapted for feeding on filamentous algae on
more or less exposed substrates in moderate to fast-flowing water. The genus is distinguished by
the specialized cephalic (postoccipital) scent-caps of the males and the absence of ocelli in both
sexes. Hydroptila contains the greatest number of species within the Hydroptilidae and has the
widest geographical distribution; it therefore may be regarded as perhaps the most successful
genus of the family.
The Oxyethira-group. This is a more heterogeneous group with regard to adult features, each
genus possessing a very distinct set of genitalia, but it is characterized by the relatively uniform
appearance of the larvae. The latter are specialized filamentous algal feeders and are generally
associated with vegetation thickets in slowly moving to static bodies of water. Within the group
there is a definite trend towards the elongation of the mid- and hindlegs in the larvae which
reaches its extreme condition in Tricholeiochiton fagesii. Oxyethira, like Hydroptila, has a continu-
ous world-wide distribution (although represented by fewer species) while Stenoxyethira,
Paroxyethira and Xuthotrichia are restricted to SE. Asia, New Zealand and Australia respectively ;
Tricholeiochiton has one west Palaearctic and two SE. Asian representatives only.
Thus the Hydroptilini as interpreted here consists of three distinct and more or less specialized
groups which have been united on the basis of the form of the male aedeagus and the basic asso-
ciations of the larval stages with green filamentous algae. The three groups probably diverged
relatively early in the evolution of the Hydroptilini branch although somewhat later than the
Ochrotrichiini, Neotrichiini and Orthotrichiini split off from this main stem (vide Chart II). The
hypothetical ancestral Hydroptilini larva probably differed little in appearance from that of the
present-day generalized condition as seen, for example, in Hydroptila, with the slightly laterally
compressed abdomen, subequal legs and simple ‘purse-type’ case. Such a condition is also seen
in the unspecialized members of other groups such as the Ptilocolepinae, Stactobiella, Ochrotrichia
and Dibusa. Within the Hydroptilini, however, the form of the larval case varies little from the
generalized condition in Agraylea, Allotrichia, Hydroptila, Tricholeiochiton, Paroxyethira and
perhaps Xuthotrichia, the only specialized genus in this respect being Oxyethira with its distinct
‘jug-shaped’ cases. The main differences in case-form appear to be associated with corresponding.
differences in larval morphology which reflect trends towards adaptation to life in vegetation
thickets in slow-moving bodies of water. Thus the cases lose any extraneous particles which might
weigh them down and the larvae tend to have longer mid- and hindlegs.
Within the Agraylea-group, Agraylea and Allotrichia have a temperate (to cool temperate)
THE GENERA OF THE HYDROPTILIDAE 193
distribution, Dhatrichia and Microptila are more subtropical (to temperate) while Ugandatrichia
is tropical. Hydroptila and Oxyethira are cosmopolitan genera and occur throughout the Ameri-
cas and through SE. Asia and Indonesia to the Australian region. The endemic Australian and
New Zealand genera, Xuthotrichia and Paroxyethira respectively, probably represent descendants
of an early branch of the Oxyethira-line which occurred in the Australian region but which was
cut off from the mainland leaving the two genera to evolve in isolation ever since. Tricholeiochiton
is distinguished by its rather unusual distribution pattern with one western Palaearctic species and
two SE. Asian species. The origins of this genus are obscure but, from the general form of the
larva, it may again have been derived from an early offshoot of the Oxyethira-line.
The success of the Hydroptilini (in terms of its present-day distribution and numbers) may be
due partly to its larval associations with green filamentous algae and partly to its tolerance of a
wider range of aquatic habitats in comparison with the more specialized Stactobiini and Leuco-
trichiini and the more restricted Ochrotrichiini and Neotrichiini. The utilization of algal filaments
(i.e. their cell fluid contents) may have opened the pathway from a life restricted to rocky and stony
substrates, which is maintained to a certain extent in Hydroptila, to one in vegetation thickets. The
latter may have at first been restricted to the banks of streams and larger, moderately flowing
rivers but thence radiated into the more slowly moving waters of backwaters, lakes and ponds
culminating in artificial, almost static, bodies of water such as reservoirs, etc. The distribution of
vegetation-dwellers, or rather their dispersal, may also be enhanced by the accidental transport
of the immature stages with the aquatic plants on which they dwell; here again the influence of
man may play an important part in the success of these insects. Hydroptila shows great species
variation as to habitat preference (as does Oxyethira to a certain extent); some, such as forcipata
and angulata in Britain, favour the more stony fast-flowing streams of the north and west uplands
while others, such as sparsa and simulans, occur more frequently in the slower and relatively more
vegetated streams and canals of the lower regions.
Recent discovery of the larva of Ugandatrichia (Scott, 1976) indicates that this genus may have
branched off from the Hydroptilini line at a very early stage in the evolution of the group, the
larvae becoming specialized tropical members of the hydropetric fauna while the adults retained
the generalized primitive form as seen also in Agraylea, for example.
In conclusion the Hydroptilini, although superficially seeming to be a somewhat heterogeneous
tribe, can be seen to have a relatively recent point of origin along the main Hydroptilinae branch
(although this was still geologically quite distant as indicated by the very modern looking fossil
amber species of Agraylea and Allotrichia). The group has radiated out to filla number of aquatic
niches in association with green filamentous algae and larger and more slowly moving bodies of
water. Early offshoots of the Hydroptilini-line may have given rise to the Ochrotrichiini and Neo-
trichiini in the Americas (and the unique, red-alga associated, Nearctic genus Dibusa) and the
highly specialized Orthotrichiini (q.v.).
Genus AGRAYLEA Curtis
Agraylea Curtis, 1834 : 217. Type-species: Agraylea sexmaculata Curtis, by subsequent designation by
Westwood, 1840.
Agraules Agassiz, 1846 : 32. [Unjustified emendation of Agraylea according to Fischer, 1961.]
Hydrorchestria Kolenati, 1848 : 103. Type-species: Agraylea sexmaculata Curtis, by subsequent designa-
tion by Kimmins, 1950. [Synonymized by Kimmins, 1950.]
DIsTRIBUTION. Holarctic.
There are nine species in this small Holarctic genus although some of these are of doubtful
status and are very restricted in occurrence. The most successful species are sexmaculata, which
has a wide western Palaearctic distribution, and mu/tipunctata, which is reputedly Holarctic but
which probably involves a species complex. In my opinion the Nearctic form of multipunctata, as
figured in Ross (1944), is distinct from the Palaearctic form (the type) in features of both the male
and female genitalia (notably the relative size and shape of the inferior appendages and ventral
process of sternite VII of the male) and should be referred to by its original designation (signata
Banks). However, Ross (in litt.) has Alaskan material of what may prove to be a third species and
he agrees that a thorough study should be made of all records previously referable to multipunctata
194 J. E. MARSHALL
from throughout the Holarctic region, including the records of the closely related European species,
cognatella.
The following list summarizes the status of the species at present comprising the genus Agraylea.
A. spathifera Ulmer: Baltic Amber; closely related to multipunctata.
A. insularis Hagen: known only from a single female specimen (now lost) from Madeira which,
according to Nybom (1948), was probably a female of Stactobia atra (Hagen).
A. drosima Navas: Navas (1917) gives only a crude description and figure of this subsequently
unrecorded species from Spain; its identity remains doubtful until the type-specimen can be
located.
A. argyricola Kolenati: originally described from Sweden and subsequently from Finland, this
species has been regarded of doubtful identity ever since Eaton (1873) considered it to be a possible
synonym of multipunctata; Nybom (1960) regards it as an ‘unintelligible species’ and has with-
drawn it from the Finnish list.
A. multipunctata Curtis: possibly represented by a species complex (see above) consisting of
multipunctata Curtis (Palaearctic) and multipunctata Curtis sensu Morton (1905) (Nearctic)
(=signata Banks 2).
A. sexmaculata Curtis: a distinct Palaearctic species.
A. cognatella McLachlan: in this species the abdomen of the type-specimen, which has been
crudely figured by McLachlan (1880), has been lost; however, it appears to be most closely
related to multipunctata; the larva has been described by Solem (1972), although he had no spare
adult specimens available for examination.
A. saltesea Ross: Nearctic (Montana, California); according to Ross (1944) this bears most
resemblance to multipunctata.
A. costello Ross: Nearctic (Ontario, Maine); Ross (1941a) considers this to be most closely
related to Allotrichia pallicornis (Eaton) (q.v.).
The larvae of the following species have been described: multipunctata (Nielsen, 1948; Lepneva,
1964; Hickin, 1967); sexmaculata (Lepneva, 1964; Barnard, 1971) and cognatella (Solem, 1972).
Agraylea is most closely allied morphologically to Allotrichia from which it was originally dis-
tinguished by the presence of fork 1 (R, and Rs) in the hindwing of Agraylea. This has not
proved to be a consistent generic feature and the genera may later be found to be synonymous.
They have been treated separately in this account, however, because the male genitalia of the
Allotrichia-group of species conform to a distinct pattern (which may, nevertheless, be derived
from the basic Agraylea plan). Also, the form of the male genitalia appears to link Agraylea with
Ugandatrichia (and Moselyella) and Dhatrichia, the relationships of which will be discussed later.
The following diagnosis is based on specific descriptions and examination of adult and larval
material in the BMNH collections. The general morphological features, apart from those of the
genitalia, also apply to Allotrichia.
D1acnosis. Adult. Forewing length 4-5 mm; wings (Fig. 7) relatively broad, apices slightly tapered, vena-
tion fairly complete (cf. Figs 5 and 8): head (Fig. 108) and antennae simple; ocelli 3; postoccipital lobes
unmodified; tentorium distinct: mesoscutellum (Fig. 108) narrow, diamond-shaped; metascutellum
narrow, pentagonal, parallel-sided: spur formula 0.3.4: lateral setate processes of abdominal segment V
present, typical: 2 genitalia a simple oviscapt: ¢ genitalia (Fig. 109) generally with a long ventral process
on segment VIII; segment IX forming an annulus, usually with a deep dorso-anal indentation; tergite X
reduced to a short membranous lobe; inferior appendages generally well developed; subgenital plate
triangular, tapering posteriorly (ventral view) and with a median, ventrally projecting process; aedeagus
relatively short and stout with distinct proximal and distal halves, proximal broad ejaculatory duct
giving rise to the slender distal intromittent organ from which arises the spiral ‘titillator’ (midway between
the end of the ejaculatory duct and the constriction at the base of the intromittent organ).
Larva (Fig. 110). Body slightly compressed laterally: dorsal ecdysial lines of head indistinct; labrum
symmetrical, short and broad with convex sides, concave anterior margin and small, antero-lateral pro-
jections; mandibles asymmetrical, robust (left with apical tooth and projecting dorsal blade without teeth
but with small apical denticles, median brush present; right mandible without teeth which are replaced
distally by two tubercules, inner median margin with central membranous bladder-like structure) ;
‘postmental sclerites’ entire in multipunctata, paired in sexmaculata (Figs 22, 21); posterior ventral apo-
tome absent: legs short, subequal; fore-femur short and broad, baso-ventrally produced into a short
THE GENERA OF THE HYDROPTILIDAE 195
Figs 108-113 Hydroptilini, Agraylea-group. 108, 112, 113, head and thorax, dorsal view; (108)
Agraylea; (112) Ugandatrichia (Moselyella); (113) Ugandatrichia s.str. 109, generalized 3 genitalia,
lateral view, Allotrichia (vide Figs 15, 16). 110-111, Agraylea sp., lateral view; (110) larva; (111)
case.
process with apico-ventral projection of trochanter; distal process of tibia with typical spinose setae:
small oral sternite present on prothorax only (Fig. 40): abdominal segment I larger than metathorax,
abdomen widest at segment IV, intersegmental constrictions deep; dorsal ‘rings’ present on segments
II-VIII; tergite present on segment IX; anal prolegs fused to X, anal claws relatively longer than in other
hydroptilids. The early instars are described by Nielsen (1948).
The larvae of multipunctata, sexmaculata and cognatella can be distinguished by the markings of the
dorsal head capsule (Solem, 1972).
Case (Fig. 111). ‘Purse-type’, constructed of silk and filaments of algae. The case building behaviour of
multipunctata is described by Nielsen (1948).
Pupa. Typically hydroptilid, mandibles long, without median teeth; case a sealed larval case attached
at each corner by a short silken pad to the surrounding vegetation.
BioLoGy. The larvae of Agraylea occur in plant thickets in lakes and slowly flowing rivers in
association with green filamentous algae. The larvae of mu/tipunctata and sexmaculata have been
observed to feed (Nielsen, 1948; Barnard, 1971) by grasping each algal filament with their modified
chelate forelegs and passing it upwards between the mandibles with which they bite into each
cell; the cell contents are then sucked out through this excision. According to Wiggins (1977) gut
contents of multipunctata (Nearctic) included diatoms while Siltala (1907) records Fucus, algae
and diatoms from the guts of Palaearctic specimens of this species.
196 J. E. MARSHALL
Genus ALLOTRICHIA McLachlan
Allotrichia McLachlan, 1880 : 508. Type-species: Agraylea pallicornis Eaton, by monotypy.
DISTRIBUTION. Palaearctic (including N. Africa).
This genus was erected for a single male specimen of what McLachlan regarded as a new species
from Worcester, England but, just prior to publication, he discovered that it was identical to
Agraylea pallicornis Eaton which thus became the type-species. McLachlan regarded the genus
as distinct from Agraylea on the basis of the absence of fork 1 (R, and R;) (vide Fig. 7, arrowed)
in the hindwing. As pointed out previously, this has not proved to be a constant diagnostic
feature and the genera may later come to be regarded as synonyms. Allotrichia is retained here,
however, as the male genitalia of the constituent species conform to a characteristic pattern and
the female genitalia and larval habits of pallicornis are distinct from those of Agraylea (although
it is not certain whether pallicornis is typical of the Allotrichia group in general).
Apart from pallicornis, which has a widespread western Palaearctic distribution, the other
members of this genus are all very local in occurrence; the following list summarizes the distribu-
tion and status of the species of A//otrichia.
A. ampullata Ulmer: Baltic Amber.
A. succinica Hagen: Baltic Amber.
A. heterocera Navas: based on a single female specimen from Spain with a vague description
accompanied by a very nondescript figure and said by Navas to ‘resemble pallicornis’ ; this species
is therefore unrecognizable and must await the discovery and examination of the type.
A. pallicornis (Eaton): central and southern Europe, Algeria, Iran.
A. vilnensis Raciecka: refigured by Schmid (1959a) and known from the type-specimen from
Poland and further material from Iran (Schmid); this species is distinct but very closely related to
pallicornis.
A. tauri Jacquemart: described from a single male specimen from Turkey; it may prove to be
synonymous with pallicornis.
A. teldanica Botosaneanu: a distinct species from Israel, closely related to vilnensis.
A. laerma Malicky: a distinct species from Greece.
[A. africana Marlier & Vaillant: I believe this species from the Congo to belong to the genus
Ugandatrichia and to be closely related to, if not synonymous with, U. nigra Mosely (q.v.).]
The members of the genus Allotrichia are morphologically very similar to those of Agraylea
and the adult generic diagnosis given for the latter, excluding the male and female genitalia, also
applies to Al/otrichia and therefore has not been repeated here. New species are allocated to either
genus not according to the fusion of R, and R, in the hindwings, as was previously the practice,
but by the general form of the male genitalia. However, Ross (1941a) states that the genitalia of
Agraylea costello Ross are more like those of Allotrichia pallicornis, thus implying that the two
genera are synonymous. Before any definite conclusions can be drawn, however, a thorough study
of the Agraylea—Allotrichia complex needs to be carried out, incorporating as many male, female
and larval features as possible; only the larva of pallicornis has so far been described (Giudicelli &
Vaillant, 1967).
The following accounts of the male and female genitalia and the immature stages are based on
original descriptions and examination of adult material of pallicornis in the BMNH collections.
DiAGnosis (see also Agraylea). Adult. 2 genitalia an oviscapt; segment VIII without a ventral sclerite but
with a characteristic asymmetrical groove. ¢ genitalia (Fig. 109) with concave inferior appendages, broad
with concave posterior margins; the subgenital plate bears a pair of characteristic asymmetrical dorsal
processes.
Larva (cf. Fig. 110) (after Giudicelli & Vaillant, 1967). Characterized by the lightness of the pigmentation
of the sclerities and absence of dorsal markings (cf. Agraylea): setae of head and labrum much shorter
than in Agraylea; dorsal ecdysial lines of head appear to be more distinct than in Agraylea; mandibles
similar to those of Agraylea although the apices are more dentate; foreleg with distinctive chelate form but
devoid of the specialized distal process of the tibia: abdominal tergites absent, including tergite [X (un-
usual); anal prolegs simple, claws apparently short.
Case. Identical with that of Agraylea (cf. Fig. 111).
THE GENERA OF THE HYDROPTILIDAE 197
Pupa. Typical; mandibles long with fine serrations along inner edge; case similar to that of larva but
sealed, with two attachment pedicels (cf. four in Agraylea) at either end of one long side by which the
case is held vertical to the substrate.
BioLoGy. The immature stages were found in a rapid stream at 750 m with little shelter or marginal
vegetation. Larval gut contents revealed, according to Giudicelli & Vaillant (1967), that the larvae
are secondary substrate feeders which have been derived from the basic filamentous algal-feeding
Agraylea stock; they have retained certain features associated with algal-feeding such as the asym-
metrical mandibles, chelate forelegs, lateral flattening of the body and case and the incorporation
of algal filaments into the latter.
The larvae of Allotrichia pallicornis thus differ from those of the known species of Agraylea
in the lack of pigmentation, absence of abdominal tergite IX, simplicity of the anal prolegs,
shorter head and labral setae, dentate mandibles and the attachment of the pupal case by two
(cf. four) adhesive discs. These differences are apparently due to structural adaptations to living
in fast-flowing water conditions and detriphagous feeding habits. As only the larva of pallicornis
has so far been described, it is not known whether these features are diagnostic of Allotrichia
species in general.
Genus MICROPTILA Ris
Microptila Ris, 1897 : 416. Type-species: Microptila minutissima Ris, by monotypy.
DISTRIBUTION. W. Palaearctic.
Microptila was erected for a new Swiss species, minutissima and bejela Mosely was subse-
quently described from Southern Yemen (Aden). Schmid (1960), as a result of studies on the
trichopterous fauna of Sri Lanka and Pakistan, synonymized the African genus Ugandatrichia
Mosely and the SE. Asian genus Moselyella Kimmins with Microptila on the basis of features of
the wing venation and the general form of the male genitalia. Schmid described a number of new
species, mainly from Pakistan and adjacent areas, which he considered to exhibit male genitalic
features intermediate between those of the Ugandatrichia—Moselyella complex, which contains
some of the largest known hydroptilids, and Microptila, which contains some of the smallest.
I have examined type-material of all three genera (BMNH collections) (including paratypes of
three species described by Schmid (1960) and placed in Microptila sensu Schmid) but only one
specimen of the type-species, minutissima, of Microptila was available for study. This is mounted
in Canada Balsam and is now in a very poor condition, the features of the genitalia (male) being
indistinct and the thorax completely destroyed thus rendering a comparative study impossible.
Nevertheless, Ugandatrichia and Moselyella are considered to be distinct from Microptila s.str.,
the two former genera comprising a distinct group which has more affinities, at least in the adult
stages, with Agraylea and Allotrichia, whereas Microptila s.str. bears more resemblance to Dhatri-
chia Mosely. This raises the problem of the correct generic identity of the species described by
Schmid as Microptila s.|., which will be dealt with below.
On the basis of adult features alone, I propose to reinstate the genus Ugandatrichia, with
Moselyella as a junior synonym, and to interpret Microptila in its original sense. These views may
need to be modified with the discovery of new species and the larval stages, which have so far been
described only for Ugandatrichia.
DiaGnosis. Adult. Forewing length 1-5—3-0 mm; antennae 20- to 25-segmented; ocelli 3: mesoscutellum
short, anterior margin convex; metascutellum trapezoidal: spur formula 0.3.4: 2 genitalia an oviscapt:
3 genitalia simple; segment IX large, fused, with a shallow excision along dorso-posterior margin;
tergite X forming a short membranous dorsal lobe; subgenital appendages elongate, slender, apparently
arising lateral to tergite X; inferior appendages elongate, apices slightly incurved; subgenital plate elon-
gate, semi-tubular; aedeagus simple, long and slender, tapering to a pointed apex in bejela (vide Mosely,
1948) and with a short spiral ‘titillator’ in indra and apsara.
M. indra is retained in this genus on the basis of the form of the thoracic nota; apsara is only
doubtfully retained.
198 J. E. MARSHALL
Genus UGANDATRICHIA Mosely gen. rev.
Ugandatrichia Mosely, 19396 : 36. Type-species: Ugandatrichia minor Mosely, by original designation.
Moselyella Kimmins, 1951 : 195. Type-species: [thytrichia violacea Morton, by original designation and
monotypy. [Synonymized by Schmid, 1960.]
DISTRIBUTION. Africa (Congo, Uganda, Kenya, Zimbabwe); S. and SE. Asia (Burma, Assam,
Pakistan, Sri Lanka, Sarawak).
Ugandatrichia is a small genus (10 species) of large, primitive-looking tropical hydroptilids.
Schmid (1960) synonymized the then distinct genera Ugandatrichia and Moselyella with the Palae-
arctic genus Microptila but, for the reasons set out in the discussion below, Ugandatrichia (in-
cluding Moselyella) has here been reinstated.
The following diagnoses are based on examination of adult material in the BMNH collections
and on Scott’s (1976) descriptions of the adults, pupae and larvae of rhodesiensis.
D1aAGnosis. Adult. Forewing length variable, 2-6 mm; wings relatively broad but still long and acuminate
as typical of the Hydroptilinae; forewings with dense setal covering (dark) with an oval patch of creamy
scale-like hairs arising centrally from the membrane in cyanotrichia (Kimmins) (Fig. 6): tentorium com-
plete (Figs 112, 113); antennae usually long (32-37 segments); ocelli 3: mesoscutellum diamond-shaped
(Figs 112, 113) as in Agraylea (Fig. 108); metascutellum ‘convexly subtriangular’, narrow: spur formula
0.3.4: violacea, cyanotrichia and nikitaruwa (Schmid) each has a pair of long, membranous extrusible
processes arising laterally between the tergite and sternite of segment II in the male: 9 genitalia an ovis-
capt; sternite VII with diagnostic patch or row of setae in certain species; sternite VIII with process or
some other distinctive marking: ¢ genitalia with segment IX well developed, variously with dorsal and
ventral posterior and anterior excisions; tergite X a median lobe with well-developed elongate lateral
processes; inferior appendages large, broad and elongate; subgenital plate trilobed (or emarginate);
aedeagus with distinct proximal and distal regions, divided by a constriction adjacent to which arises
a short, spiral ‘titillator’.
The thoraces of nikitaruwa (Schmid) and sourya (Schmid) are almost identical to that of Ugandatrichia
(as seen in species of Moselyella), while roudra (Schmid) is included here on the basis of size and male
genitalic affinities with Ugandatrichia species as noted by Schmid (1960). M. apsara Schmid may also be-
long here (see Microptila s.str. section).
Larva. The larva of U. rhodesiensis Scott has recently been described by Scott (1976) from which the
following description of the immature stages and biology has been taken (for figures see Scott, 1976).
Fifth instar free-living, robust and heavily sclerotized, flattened dorso-ventrally: head relatively small,
oval, tapering anteriorly; eyes large, distinct; ecdysial lines distinct; antennae well developed; anterior
and posterior ventral apotomes small, former very narrow with long, slender anterior arms; labrum well
developed, rectanguloid with convex sides, convex anterior margin and dense setal brushes arising
postero-ventrally and projecting beyond anterior margin; mandibles almost symmetrical, apices blunt
with no lateral teeth, left with dense inner brush of predominantly feathered spines (‘penicillus’), right
ridged externally and with long inner feathered spine and tuft of shorter bifid spines: all three thoracic
nota paired; pleurites with preepisternite, episternite and epimeron; sterna entirely membranous and
strongly muscular; legs short, stout and serving as powerful clinging appendages: abdomen dorso-ven-
trally flattened, gills absent; well-sclerotized tergites present dorsally on all segments, each with clear
central spot on segments I-VIII; tergite IX with emarginate posterior margin; all segments distinctly
demarcated and strongly muscular, ventral surfaces of segments II-VIII each with a pair of suckers close
to lateral margins (adaptation for clinging to rocks); anal prolegs short, stout, fused to X and with simple
large, hooked anal claws.
The larval stages are free-living throughout, a case being built only at the end of the final instar in which
the latter pupates.
Pupa. Typically hydroptilid. Pupal case sac-like, tubular but not flattened and anchored by a holdfast
at one end; case of tough, semi-transparent, parchment-like secretion.
BioLoGy. The larvae and pupae of rhodesiensis were collected mainly from the most torrential
parts of the waterfall on the Bundi River in Rhodesia. The larvae are clearly adapted for living in
such harsh hydropetric conditions as indicated by their strongly sclerotized, robust, dorso-
ventrally flattened form, clinging appendages and loss of the case building habit in the final instar.
The form of the labrum and mandibles suggests that the larvae are substrate feeders.
Discussion. According to the basic form of the male genitalia and the structure of the thoracic
nota, especially with regard to the characteristic diamond-shaped mesoscutellum, I consider
THE GENERA OF THE HYDROPTILIDAE 199
Ugandatrichia and Moselyella to be synonymous and closely related to Agraylea as apposed to
Microptila in which the mesoscutellum is more ‘kite-shaped’ with a convex anterior margin. Only
specimens of nikitaruwa (Schmid), sourya (Schmid) and indra Schmid were available for study
(BMNH collection, paratypes) and, from their thoracic features, the last mentioned would appear
to belong to Microptila s.str. while the other two strongly resemble Ugandatrichia species. The
affinities of apsara (Schmid) and roudra (Schmid) are still doubtful.
Thoracic structure alone, however, cannot be used as an indication of generic affinity. As
Schmid (1960) rightly states, the genitalia of the Microptila-~Ugandatrichia—Moselyella complex
do have a common basic form but this is very generalized and has certain features in common with
the genitalia of Agraylea, Allotrichia and Dhatrichia. Indeed, Allotrichia africana Marlier &
Jacquemart appears to belong to Ugandatrichia and is very close to, if not synonymous with,
U. nigra Mosely. Contrary to Schmid (1960), it is unlikely that there is any relationship between
Microptila s.1. and the Nearctic-Neotropical subgenus Ochrotrichia (Metrichia) since the genitalia
of the two groups are quite distinct and the characteristic mesoscutellar suture of Ochrotrichia is
absent in Microptila.
Due to the recent description of the immature stages of rhodesiensis by Scott (1976), I have been
able to include the following remarks concerning the affinities of the larvae with those of other
hydroptilid genera. As Scott notes, the only other hydroptilid genus to retain the free-living habit
throughout its larval existence is the Central American genus Alisotrichia. The larvae appear to
have developed parallel structural adaptations suiting them for life in torrenticolous conditions
but, as can be seen from a comparison of basic larval and, especially, adult features, the genera
are totally unrelated, any affinities being purely superficial. Alisotrichia is a distinct, though rather
isolated, member of the exclusively New World Leucotrichiini whereas Ugandatrichia would ap-
pear to be a distinct Palaeotropical component of the Hydroptilini.
The structural modifications of Ugandatrichia larvae are also paralleled to some extent by those
of Stactobia, which are also heavily sclerotized, dorso-ventrally flattened with robust clinging
appendages and similar madicolous habits. Fundamental differences can be seen, for example, in
the fusion of the ecdysial lines of the head, the crenellated posterior margin of tergite IX and the
retention of a case in the final instar of Stactobia. Comparison of the adult stages will immediately
discount any close affinity between the two genera. It is interesting to note here that Ugandatrichia
is essentially Palaeotropical whereas Stactobia is principally Palaearctic.
The discovery of the larva of Ugandatrichia does not, unfortunately, shed any light on its true
generic affinities as may have been hoped since, as has been emphasized throughout this account,
it is highly specialized and possesses structural modifications which tend to mask fundamental
characters. It is, however, still possible that Ugandatrichia may be a descendant of an early ancestor
of the Hydroptilini line which spread throughout the Palaeotropical regions, the larvae becoming
specialized while the adults retained the primitive generalized form similar to that seen in Agraylea
for example.
It would now seem essential to discover the immature stages of the SE. Asian Moselyella group
of Ugandatrichia, as well as those of the Palaearctic Microptila group to establish the true affinities
of this generic complex, since basic larval forms seem to serve as useful generic indicators in the
Hydroptilidae.
A single specimen (3) of Ugandatrichia (Moselyella group) has recently been collected from
Sarawak, 1978 (J. E. Marshall), BMNH coll.
Genus DHATRICHIA Mosely
Dhatrichia Mosely, 1948 : 78. Type-species: Dhatrichia inasa Mosely, by original designation and
monotypy.
DISTRIBUTION. Yemen, Zaire.
This small genus was first described for a single male specimen from the Yemen which is now
mounted as a microscope slide preparation in the BMNH collection. A new species from Zaire,
bipunctata, has been described by Statzner (1977), paratypes of which have been donated to the
BMNH. The immature stages are unknown.
200 J. E. MARSHALL
The following generic diagnosis is based on examination of the above material.
DiaGnosis. Adult. Forewing length 2mm; wings narrow, tapering: tentorium complete but very fine
medially; ocelli 3; antennae 19-segmented (male inasa); postoccipital lobes ovoid: mesoscutellum with
convex anterior margin; metascutellum pentagonal, sides convergent anteriorly; spur formula 0.3.4:
typical lateral setate processes of abdominal segment V present: 2 genitalia an oviscapt: ¢ genitalia with
segment IX annular, dorsally reduced to a narrow chitinized band and with well-developed postero-lateral
sides (almost concealing genitalia in lateral view); tergite X (‘upper penis cover’ (Mosely, 1948)) broad,
bilobed in inasa; inferior appendages short, apically broad, trilobed in inasa; subgenital plate an elongate
flat process with a pair of apical setae and not extending beyond the sides of IX; aedeagus with a broad
proximal ejaculatory duct and slender distal intromittent organ and with a spiral ‘titillator’ arising mid-
length adjacent to the median constriction.
DISCUSSION. From the structure of the male aedeagus and the thoracic nota, Dhatrichia would
appear to be a member of the Hydroptilini. The form of the male genitalia indicates an affinity
with Agraylea, especially in the shape of segment IX and the form of the aedeagus; however, the
thorax bears more resemblance to that of Microptila s.str. and these two genera (i.e. Microptila
and Dhatrichia) can be separated in the generic key only by the form of the male genitalia.
Genus HYDROPTILA Dalman
Hydroptila Dalman, 1819 : 125. Type-species: Hydroptila tineoides Dalman, by monotypy.
Phrixocoma Eaton, 1873 : 132. Type-species: Hydroptila sparsa Curtis, by original designation. [Synony-
mized by McLachlan, 1880.]
Hydropneuma Enderlein, 1929 : 232. Type-species: Hydropneuma juba Enderlein, by original designation
and monotypy. [Synonymized by Nybom, 1963.]
Hydroptilina Martynov, 1934 : 144. Type-species: Hydroptilina angustipennis Martynov, by monotypy.
[Synonymized by Fischer, 1971.]
Oxydroptila Martynov, 1935 : 114. Type-species: Oxydroptila furcata Martynov, by original designation
and monotypy. Syn. n.
Oeceotrichia Ulmer, 1951 : 85. Type-species: Oeceotrichia elongata Ulmer, by original designation and
monotypy. Syn. n.
Pasirotrichia Ulmer, 1951 : 90. Type-species: Pasirotrichia crenata Ulmer, by original designation and
monotypy. Syn. n.
Sumatranotrichia Ulmer, 1951 : 87. Type-species: Sumatranotrichia trullata Ulmer, by original designation
and monotypy. Syn. n.
DISTRIBUTION. Cosmopolitan (excluding polar regions).
Hydroptila is the most successful genus of the Hydroptilidae with over 150 species and with an
almost world-wide distribution; species have been recorded from Lapland and the northern
U.S.S.R. to South Africa and Australia (including Tasmania) in the ‘Old World’ and throughout
the Americas as far south as Nova Teutonia in Brazil. A number of geographically more or less
distinct species-groups may be recognized by the forms of the male and female genitalia, and may
eventually come to be considered as distinct subgenera. However, the species all have the follow-
ing features in common: basic genitalia patterns; thoracic nota; absence of ocelli and presence
of male dorsal postoccipital scent-organs in the adults, and the general appearance of the larvae
and their cases. The immature stages are difficult to identify at the species level; Ross (1944) gives
head and thoracic pigmentation patterns as rough guides to the identity of some Nearctic species
although these tend to be very variable intraspecifically. My own observations indicate the pos-
sible use of features of the sculpturing and shape of the mandibles as specific guides but a full
investigation was not carried out. The only satisfactory means of identification is by association
with correctly identified pupae and adults.
There is no single comprehensive work on the genus as a whole, any one species-group or any
particular faunal group, nor is there a key to the species of Hydroptila, identification at present
relying on faunal keys (of which there are few) and reference to original descriptions, some of
which are unsatisfactory and in need of redefinition. A species key was beyond the scope of this
project but I have attempted to indicate the main species-groups, these groupings being based on
affinities suggested by the original authors and, in many instances, by my own observations.
THE GENERA OF THE HYDROPTILIDAE 201
Diacnosis. Adult (Fig. 1). Forewing length 3-4 mm: head (Fig. 114) with tentorium reduced medially;
ocelli 0; antennae c. 30-segmented: mesoscutellum (Fig. 114) subtriangular with convex anterior margin;
metascutellum pentagonal to triangular, anterior margin strongly convex: spur formula 0.2.4: setate
processes present on abdominal segment V: 2 genitalia a simple oviscapt (Fig. 17); sternite VIII (‘ventral
plate’, vp) and small sternite/s on IX (‘dorsal plate/s’, dp) sometimes present: 3 genitalia (Figs 115-117)
distinctive for each species-group but basically as follows — sternite VII often with ventral process; segment
VIII rarely with ventral process; segment IX fused, annular, sometimes with antero-lateral apodemes
(Fig. 117) and often with postero-lateral processes; inferior appendages distinct, usually simple, straight
and elongate, sometimes modified (Fig. 117); tergite X forming a variable ‘dorsal plate’; subgenital ap-
pendages variable (=‘parameres’, ‘lateral penis sheaths’, ‘intermediate appendages’); subgenital plate
simple, broad, slightly emarginate in the sparsa- and consimilis-groups but absent or inconspicuous
in others; bilobed process apparently absent (but may be incorporated into subgenital plate) ; aedeagus with
distinct proximal and distal ejaculatory duct and intromittent organ respectively and a well developed
spiral ‘titillator’.
Larva (Fig. 118). The following species have been described (* indicates material examined in BMNH
collections): H. acuta (Jacquemart & Coineau, 1962); ajax, albicornis, angusta, armata (Ross, 1944);
capensis (Barnard, 1934); consimilis (Ross, 1944); delineata (Sibley, 1926); emarginata (Lepneva, 1932),
forcipata* (Fahy, 1972); grandiosa, hamata (Ross, 1944); martorelli (Flint, 1964); pulchricornis (Lepneva,
1964); sparsa* (Hanna, 1961; Lepneva, 1964; Hickin, 1967); spatulata (Ross, 1944); taurica (Botosaneanu
& Sykora, 1963); tineoides* (Nielsen, 1948; Lepneva, 1964; Hickin, 1967); trilobata (Jacquemart, 1965);
vectis* (Jacquemart & Coineau, 1962; Lepneva, 1964); waubesiana (Ross, 1944).
Head elongate with almost parallel sides, almost circular in cross-section; ecdysial lines indistinct;
‘postmental’ sclerites (Fig. 26) fused, although a faint median line may sometimes be detected, cresentic,
without posterior process (cf. Agraylea); labrum long, anterior margin deeply indented, lateral lobes slightly
asymmetrical, row of setae anteriorly; mandibles asymmetrical (Fig. 31), form variable between species,
median brush in left only: thoracic pleurites as in Agraylea (cf. Fig. 35) but with preepisternite free on
prothorax only; single oral and paired anal sternites on prothorax only (Fig. 39); legs short, subequal
(Fig. 33) with distal ventral process of fore-tibia well developed and armed with specialized setae (Fig. 47)
as in Agraylea: abdomen typically distended; dorsal ‘rings’ present on segments I-VIII; tergites absent
except on IX; anal prolegs fused to X; three filamentous caudal gills present (Figs 48, 49).
Case (Fig. 119). ‘Purse-type’, laterally compressed, constructed of silk and covered in fine sand-grains
and other inorganic particles and occasionally with diatoms and algal filaments.
Pupa (Fig. 52). Typically hydroptilid, mandibles long, without teeth; pupal case simply a sealed larval
case, attached to substrate by anterior and posterior adhesive discs on ventral edge, held vertically.
BIoLoGy. The larvae prefer running water in lakes, streams and rivers, usually on the bottom
substrate in association with the green filamentous algae on which they feed. According to
Nielsen (1948) the typical habitat of tineoides is that of ‘stony brooks in shallow water of lakes’
while others tend to prefer swifter flowing water and I agree with his observation that this is
probably due to true species preferences, as distinct differences have been observed in the habitats
of the British species.
Life-cycles are very variable according to species and locality; both univoltine and bivoltine
species have been observed. The larvae feed on filamentous green algae by grasping the filament
with the modified chelate forelegs, biting into each cell and sucking out the fluid contents.
SPECIES-GROUPS (for species see checklist). The sparsa-group: a very large Palaearctic group
recognized by the form of the male genitalia (Fig. 115).
The wncinata-group: a small Palaearctic group with characteristic male genitalia which have an
affinity with those of the sparsa-group.
The capensis-group: a small African group possibly derived from the sparsa-group.
The consimilis-group: a large Nearctic group with a distinctive basic male genitalia form which
is close to that of the Palaearctic sparsa-group.
The occulta-group: a distinct group of approximately 25 species with a distribution extending
throughout Europe into Asia Minor, Africa and SE. Asia and with at least 4 Nearctic representa-
tives; male genitalia (Fig. 116).
The tineoides-group: tineoides is the only Palaearctic member of this predominantly Nearctic
group (moselyi Ulmer from Peking may also belong here); male genitalia (Fig. 117).
202 J. E. MARSHALL
Figs 114-119 Hydroptila. 114, adult head and thorax, dorsal view (3). 115-117, generalized 3 geni-
talia; (115) sparsa-group; (116) occulta-group; (117) tineoides-group (vide Figs 15, 16). 118, larva,
lateral view; 119, case, lateral view.
The dikirilagoda-group: a distinct group of three species from Sri Lanka; possibly related to
the tineoides-group.
The /osida-group: a small group of Australasian species (Australia, Tasmania, Solomon Islands
(Guadalcanal), New Guinea) which may have some affinity with the SE. Asian members of the
occulta-group such as panchaoi.
The pulchricornis-group: a small group of Palaearctic species with possibly one N. African
representative.
The forcipata-group: of three Palaearctic species; Schmid (19595) considers that armathai
belongs here but this is dubious.
The vectis-group: two Palaearctic species.
The tigurina-group: three distinctive European species with very characteristic sidepieces to
segment IX of the male genitalia.
The waubesiana-group: a very distinct Nearctic group of seven species.
Incertae sedis: a number of species have not been assigned to any group owing to the un-
availability of material for study and the inadequacy of descriptions in the literature.
GENERIC SYNONYMS. Oxydroptila is a small genus of only two species, furcata Martynov from
India and kirilawela Schmid from Sri Lanka, which is distinguished from Hydroptila on minor
venational differences and the form of the male genitalia which is said to resemble that of Oxye-
thira, especially with respect to segment VIII. As in Hydroptila, ocelli are absent and the spur
formula is 0.2.4; examination of a male paratype of kirilawela (BMNH) has revealed the presence
of postoccipital scent organs and a thoracic structure identical to that of Hydroptila. Contrary to
Martynov (1935) and Schmid (1958a), the male genitalia do not resemble those of Oxyethira and
THE GENERA OF THE HYDROPTILIDAE 203
the female genitalia of kirilawela, as figured by Schmid (1958a), form a simple oviscapt and not the
modified, specialized condition characteristic of Oxyethira (q.v.). The larvae are unknown. It is
therefore proposed to sink Oxydroptila as a junior synonym of Hydroptila and, although both
furcata and kirilawela are included in the incertae sedis group, they probably form a distinct
species unit.
Oeceotrichia, Pasirotrichia and Sumatranotrichia are three monotypic genera described by
Ulmer (1951) from Indonesia (the Sunda Islands) and distinguished from Hydroptila solely on
minor venational differences. Each has a spur formula of 0.2.4, ocelli absent, male genitalia of the
Hydroptila pattern and, from examination of type-material (UHZIM loan), postoccipital scent-
organs (males) and a thorax identical with those of Hydroptila. According to the descriptions and
figures of the larvae and cases given by Ulmer (1957), these are also almost identical with those of
Hydroptila (notably with respect to the fore-tibial process and the three caudal filaments). It
would thus appear that Oeceotrichia, Pasirotrichia and Sumatranotrichia are all junior synonyms
of Hydroptila, the constituent species showing certain affinities, according to the forms of the male
genitalia, to the SE. Asian members of the occulta-group (e.g. panchaoi Schmid) and the Austra-
lasian /osida-group.
As an additional point it was noted, during the examination of Ulmer’s Sunda Island material
that two females in spirit labelled Pasirotrichia bore no resemblance to Hydroptila: sp. A had no
ocelli but had a thorax resembling that of Orthotrichia, and sp. B had 3 ocelli and a thorax similar
to that of Oxyethira or Tricholeiochiton. However, I was not able to identify the specimens posi-
tively owing to their fragile conditions.
The following records of material found in the BMNH collections may also be of interest (all
det. J. E. Marshall): palestinae Botosaneanu & Gasith (1 9, Southern Yemen ‘W. Aden Protec-
torate’, 1937, Scott & Britton); triloba Kimmins (1 3, Solomon Islands, Guadalcanal, J. D.
Bradley); vectis Curtis (1 3, Spain, Prov. Granada, Valley of Guadelfeo, 3.vii.1969, Sattler &
Carter).
Genus OXYETHIRA Eaton
Oxyethira Eaton, 1873 : 143. Type-species: Hydroptila costalis Curtis sensu Eaton, by original designation.
Lagenopsyche Miiller, 1879a : 39. Type-species: Lagenopsyche spirogyrae Miiller, by subsequent designa-
tion by Fischer, 1961. [Synonymized by Fischer, 1961.]
Argyrobothrus Barnard, 1934 : 392. Type-species: Argyrobothrus velocipes Barnard, by monotypy.
[Synonymized by Ross, 1948.]
Loxotrichia Mosely, 1937 : 165. Type-species: Loxotrichia azteca Mosely, by original designation.
[Synonymized by Ross, 1944.]
Dampfitrichia Mosely, 1937 : 169. Type-species: Dampfitrichia ulmeri Mosely, by original designation and
monotypy. [Synonymized by Ross, 1944.]
Oxytrichia Mosely, 1939a : 289. Type-species: Oxytrichia mirabilis Mosely, by original designation and
monotypy. [Synonymized by Kimmins, 1966.]
DISTRIBUTION. Cosmopolitan (excluding polar regions).
Oxyethira is a very successful genus of approximately 75 species with a world-wide distribution
and with very distinctive larvae which feed on green filamentous algae. The true identity of the
type-species is still in dispute. Eaton originally proposed the genus for two species, Hydroptila
costalis Curtis and H. albiceps McLachlan, the latter being an endemic New Zealand species.
Eaton (1873) considered costalis (Curtis) to be a senior synonym of H. tineoides Dalman sensu
Stephens and, doubtfully, of H. flavicornis Pictet. However, subsequent dissection of Curtis’s
male type of costalis by Neboiss (1963) showed this to be conspecific with Orthotrichia tetensii
Kolbe, the latter being placed as a junior synonym of costalis (Curtis) and congeneric with angus-
tella (McLachlan), the type-species of Orthotrichia. The next available name for costalis (Curtis)
sensu Eaton is Oxyethira flavicornis (Pictet).
This is still not an unequivocal solution to the problem, however, since Pictet’s type-specimen
of flavicornis has not yet been located (one female specimen in Geneva is labelled ‘Zurich 9/81’
and a second specimen has no abdomen according to Neboiss, 1963). Neboiss therefore merely
proposes the synonymy as ‘one of the possible answers’ but does not ignore the fact that flavicornis
204 J. E. MARSHALL
Pictet could be a synonym of H. tineoides Dalman sensu Stephens, thus leaving costalis (Curtis)
sensu Eaton without a name. However, until the problem can be resolved (and this is unlikely
since Pictet’s type appears to be missing), the proposal of Neboiss (1963) must be accepted. Never-
theless, there is no doubt as to the actual species referred to by Eaton since his figures of the dis-
tinctive male genitalia are clearly recognizable (he merely considered that costalis Curtis was the
same species).
The larval stages of Oxyethira (discounting Pictet’s (1834) figures of H. flavicornis) were first
described by Miiller (1879a) as Lagenopsyche spirogyrae Miller from Brazilian material. The
first description of British material was by Morton (1887) from specimens of O. costalis (Curtis)
sensu Eaton sent to him by McLachlan and Hudson (1886) described larvae of an unidentified
New Zealand hydroptilid which subsequently proved to be O. albiceps McLachlan. The most
detailed larval description is that of O. costalis (Curtis) sensu Eaton as given by Nielsen (1948)
upon which the generic diagnosis given below is partly based. The following list summarizes the
published descriptions of species of Oxyethira larvae to date (the type-species will now be called
flavicornis).
O. albiceps (Mosely & Kimmins, 1953); delcourti (Jacquemart, 1973); distinctella (Lepneva,
1964); dualis (Sibley, 1926); ecornuta (Lepneva, 1964); flavicornis (Nielsen, 1948; Lepneva, 1964;
Hickin, 1967); frici (Lepneva, 1964); hyalina (Miller, 1879a); incana (Ulmer, 1957); meridionalis
(=unidentata) (Jacquemart & Coineau, 1962); puertoricensis (Flint, 1964); sagittifera (Lepneva,
1964); serrata (Ross, 1944); simplex (Macdonald, 1950; Hickin, 1967); tristella (Lepneva, 1964);
velocipes (Barnard, 1934).
Apart from Kimmins’s (1958) paper on the British species of Oxyethira there has been no
previous review of the genus as a whole. The following diagnosis and species-groupings have been
based on published species descriptions and examination of adult and larval material in the BMNH
collections. To date there are approximately 75 species which fall into more or less distinct species-
groups each distinguished by the form of the male genitalia. The status of Stenoxyethira Kimmins
will be discussed later but, according to Schmid (1958a), it should be considered a synonym of
Oxyethira.
DiaGnosis. Adult. Forewing length 2:5—3-5 mm: head and antennae unmodified (Fig. 120); anterior ten-
torial arms reduced medially; ocelli 3; thorax (Fig. 120) — mesoscutellum with anterior margin convex and
posterior margins slightly concave, metascutellum convexly subtriangular: spur formula 0.3.4: 9 genitalia
a modified oviscapt (Fig. 121) (after Kimmins, 1958) with sternite VI with a short, acute ventral process;
segment VII with tergite and sternite more or less fused, ventral apical margin sometimes excised; tergite
VIII reduced to a narrow arched band, the centre of its apical margin produced as a spatulate lobe; ster-
nite VIII forming a short ‘subgenital plate’; tergite IX either free or fused to VIII, short and hood-like;
tergite X forming a short, broad plate carrying two short slender cerci; details of internal vaginal struc-
tures obscure but apparently with an upper and lower lobe, the latter with a conspicuous baso-ventral
process directed caudad: ¢ genitalia (Fig. 122) (after Kimmins, 1958) with sternite VII with short, acute
ventral process; segment VIII with tergite and sternite more or less fused, apical margins dorsally and
ventrally excised, that of tergite with dorsal or lateral processes, the latter sometimes armed with stout
spines; segment IX mostly withdrawn into VIII, its upper half generally much shorter than lower, often
reduced to a short transverse band, lateral apical margins of IX sometimes produced or with variously
formed spiniform processes, ventral margin generally excised and from which arises a lobe; tergite X
obscure, membranous; subgenital appendages represented by two sclerotized plates attached to segment
1X, fused basally with their apices variously hooked downwards (‘fused claspers’ (Kimmins, 1958);
‘lower penis cover’ (Mosely, 1939a); ‘subgenital plate’ (Ross, 1948)); above and arising from them are two
pairs of short, membranous ‘bilobed’ processes each terminating in a short apical seta; aedeagus long and
slender with a spiniform ‘titillator’ arising at midlength.
Larva (Fig. 123). Head elongate, of uniform diameter, slightly laterally compressed; dorsal and ventral
ecdysial lines distinct; postmental sclerites paired, anterior ventral apotome well developed, U-shaped
with a distinct transverse ridge, posterior ventral apotome present (Fig. 27); labrum with strongly convex
lateral margins and deep median anterior indentation; mandibles asymmetrical — left with projecting
blades bearing small teeth and with a median brush, right with blades poorly defined and distal only, teeth
only near apices: thoracic pleurites of each segment fused except for the free anal epimeron of the pro-
thorax; three sternites present on each thoracic segment (Fig. 41); forelegs short with small baso-ventral
setate lobe on femur and long distal process bearing a ventral row of short, spine-like setae on tibia
THE GENERA OF THE HYDROPTILIDAE 205
122
Figs 120-125 Oxyethira. 120, adult head and thorax, dorsal view; 121, generalized ° genitalia, ventral
view; 122, generalized 3 genitalia, lateral view (vide Figs 15, 16); 123, early fifth instar larva in
case, lateral view; 124, late fifth instar larval case, lateral views; 125, pupa in case, ventral view.
(chelate); mid- and hindlegs twice as long as forelegs, tarsal claws long and slender: abdomen typically
distended, intersegmental constrictions distinct; laterally compressed and usually held in characteristic
comma-shaped attitude; dorsal ‘rings’ present on segments II—VII; tergites absent except on segment IX;
anal prolegs fused to side of segment X; caudal gills absent.
The early instars of flavicornis (Pictet), as costalis (Curtis), are described and figured in Nielsen (1948).
Case (Figs 123, 124). Laterally compressed, semi-transparent, constructed of secretion only; ‘jug-’ or
‘bottle’-shaped; posterior end broad with slit-like opening, anterior end narrow, constricted to form a
neck with a well-defined rim around the opening. The pupal case (Fig. 125) is basically a sealed larval
case attached horizontally to the substrate by four short ligaments, each with an adhesive disc and arising
from each corner (in some species there are two ligaments from each of the anterior corners); the pupa
206 J. E. MARSHALL
lies with its head towards the broad (hence anterior) end (cf. larva), with its dorsum towards the substrate;
the case is closed anteriorly by a narrow curved membrane, the posterior neck being filled with a plug of
silk; the pupal cases usually occur on aquatic plants, rarely on submerged rocks or wood.
Pupa. Typically hydroptilid; mandibles long with serrate inner blades.
BioLoGy. The larvae are generally considered to occur in plant thickets in slow-moving and static
bodies of water in association with the green filamentous algae upon which they feed. This is true
for distinctella, ecornuta and sagittifera, for example (Lepneva, 1964), and also velocipes (Barnard,
1934) and spirogyrae (Miiller, 1879c) but a number of species are reputed to frequent faster-flowing
streams where the larvae are found on the rocky substrate in areas covered with algae; these are
frici and tristella (Lepneva, 1964), azteca (or janella) (Flint, 1968a) and hyalina (Miller, 1879c).
O. flavicornis (Pictet) (costalis Curtis sensu Eaton) is a pond dweller although, in Pictet’s (1834)
original description, the larva of flavicornis is reputed to occur in fast-flowing streams with
Hydroptila pulchricornis Pictet, thus casting further doubts on the identity of flavicornis (see
above). In Illinois, Ross (1944) records pupae of serrata from the underside of rocks in about 1 m
of water along the open beaches of glacial lakes. Pupal cases of albiceps in New Zealand, according
to Hudson (1886), were found attached to stones in streams, especially in the late summer when
‘the rivers are generally very low ... and a luxurious growth of green slime weed is present’.
Finally, puertoricensis is found in all types of streams and at all elevations in Jamaica (Flint,
19685) and, on Puerto Rico (Flint, 1964), the cases are found ‘in tangles of filamentous green algae’
and are fixed to irregularities in the rocks at pupation.
The larvae are highly specialized for feeding on filamentous algae and for living in masses of
vegetation as indicated by their chelate forelegs, laterally compressed bodies and very long, slender
mid- and hindlegs. According to Siltala (1907) gut contents of an Oxyethira species included
diatoms and entire algal filaments and Jacquemart & Coineau (1962) similarly record diatoms and
desmids (the normal food source is reputed to be simply the fluid contents of algal cells). Nielsen
(1948) describes the case building behaviour of flavicornis while Jacquemart & Coineau (1962)
describe the similar habits of an unidentified Oxyethira species.
SPECIES-GROUPS. At least 10 species-groups may be recognized by the form of the male genitalia.
The following groupings have been based on relationships suggested in original species descrip-
tions and have been verified by me as far as the literature and available material would allow. The
lists of species are given in the checklist.
The falcata-group: essentially a western Palaearctic group of seven species, three occurring
only in the Atlantic Islands, and with possibly one Nearctic representative. Two species (falcata
and frici) are British. The group is distinguished by the spinose postero-lateral processes of the
male segment VIII. There is still some doubt as to the status of fischeri and dentata which both
have very similar genitalia to those of falcata; as Higler (1974) points out in his paper on fischeri,
falcata may exhibit ‘eco-geographic’ forms (after Jacquemart & Coineau, 1962) and he considers
that dentata may belong to one of these forms. However, Higler believes that fischeri is distinct
from falcata, mainly on account of features of the male aedeagus (based on the figures of falcata
in Jacquemart & Coineau, 1962) although Mosely’s (1939a) figures of the aedeagus of falcata are
very similar to Higler’s figures of fischeri! A thorough study of falcata and its reputed close
relatives is obviously required in order to sort out this species-complex. O. boreella also is close
to falcata but would seem to be distinct according to both male and female genitalic characters
(Svensson & Tjeder, 1975).
The flavicornis-group: a Palaearctic group extending from Europe (and Egypt) to Korea with
perhaps one species from Sri Lanka. It may be related to the Nearctic rivicola-group and is dis-
tinguished by the absence of a median ventral lobe on the fused claspers and the broad, widely
separated subgenital plates (in ventral view).
The rivicola-group: possibly related to the flavicornis-group and, although essentially Nearctic,
may contain a number of Neotropical members (from Surinam, described by Flint, 1974).
The simplex-group: again possibly related to the flavicornis-group; this group may require
further investigation as, at present, it contains two geographically distinct elements: simplex and
THE GENERA OF THE HYDROPTILIDAE 207
tristella from northern Europe and NW. U.S.S.R., and paramartha, ramosa, harpagella and
galekoluma (and, possibly, Stenoxyethira excisa) from SE. Asia.
The distinctella-group: a small Holarctic group distinguished by the prominent dorso-lateral
posterior serrate processes of the male segment VIII.
The azteca-group (=Loxotrichia Mosely): a small group from Central America and the Antilles,
characterized by the form of the male segment IX. O. archaica Malicky from Portugal would also
appear to belong here, its occurrence in this region possibly being a result of a chance introduction
from S. America (the group occurs, for example in Brazil); it is unlikely that it is an early Tertiary
relict as stated by Malicky (1975).
The ulmeri-group (= Dampfitrichia Mosely): another small Nearctic group distinguished by
the dorso-lateral tapering processes of segment IX.
The zeronia-group: essentially Central American and characterized by the anterior prolongation
of the ventral margin of segment IX.
The pallida-group: a small but distinct Nearctic group of uncertain affinities.
The bidentata-group: a Nearctic—-Neotropical group of three species.
The mirabilis-group: containing mirabilis (Palaearctic) and flagellata (Reunion Island); the
latter has some affinity with bidentata which itself was once included with mirabilis in the genus
Oxytrichia.
Incertae sedis: of these, albiceps is endemic to New Zealand and velocipes to South Africa;
spirogyrae and hyalina (Lagenopsyche) from Brazil are known only in their larval stages; incana
(Indonesia) was described from the female and larva only (Ulmer, 1951; 1957), while angustella
is known from the wings alone. O. anabola and berneri are distinct Nearctic species while mithi,
from Greece, has not yet been placed.
Genus STENOXYETHIRA Kimmins
Stenoxyethira Kimmins, 1951 : 207. Type-species: Stenoxyethira minima Kimmins, by original designa-
tion.
Gnathotrichia Ulmer, 1951 : 59. Type-species: Gnathotrichia isabellina Ulmer, by original designation and
monotypy. Syn. n.
DISTRIBUTION. SE. Asia (Burma, Indonesia).
Kimmins (1951) erected Stenoxyethira principally for the type-species, minima, but included,
somewhat doubtfully, excisa Kimmins on the basis of the general similarity of the wing venation.
The genus is here considered to contain just these two species with Gnathotrichia isabellina treated
as a synonym of Stenoxyethira excisa. The following generic diagnosis is based on Kimmins
(1951). The immature stages are unknown.
DiaGnosis. Adult. Forewing length 1-6 mm: head and antennae unmodified; ocelli 3; spur formula 0.2.4
(0.3.4 in excisa and isabellina): 3 genitalia (cf. Fig. 122) with segment VIII forming a ring concealing
segment IX, latter narrow and lightly sclerotized dorsally, ventral margin produced as a rounded, bilobed
plate (fused inferior appendages ?) above which is a triangular, trough-shaped subgenital plate; tergite X
difficult to interpret; aedeagus rod-like, with a single spine or sheath.
S. excisa was included in this genus on the basis of venational features, being distinguished from
minima by the presence of a small spine or microscopic spur on the mid-tibia, the form of the
aedeagus and the produced spines of segment VIII of the male genitalia. Kimmins (1951) was
in no way satisfied with this arrangement and noted on the card for Stenoxyethira in the BMNH
index (Hydroptilidae) ‘?=Gnathotrichia Ulmer, 1951’. I noted independently of Kimmins the
similarity between these two genera when examining type-material of G. isabellina Ulmer (UHZIM
loan) and concluded that this species, from Java and Sumatra, was probably synonymous with
S. excisa Kimmins. According to the dates of publication (Kimmins, March, 1951; Ulmer, April,
1951) S. excisa takes priority. However, as Schmid (1958a) points out, S. excisa may truly belong
in Oxyethira, being closely related by the form of the male genitalia and wing venation to O.
galekoluma Schmid from Sri Lanka (the venation of the latter approaching that of G. isabellina
and O. angustella Martynov). I agree that there is a similarity in form which is also supported by
the spur formula (0.3.4) and the structure of the thoracic nota, but I do not agree with Schmid
208 J. E. MARSHALL
that Kimmins’s figures of the male genitalia of S. excisa are inverted in comparison with those
of O. galekoluma (Schmid, 1958); on the contrary, it is Schmid who has misinterpreted their
orientation.
It would thus seem that Stenoxyethira and Gnathotrichia should be regarded as synonyms of
Oxyethira, at least with regard to S. excisa. The status of S. minima is not so easily resolved — as
regards venational and male genitalic characters it could well be a specialized member of Oxye-
thira, although it is the only species so far recorded without a preapical spur on the mid-tibia.
This may simply prove to be a specific or even individual variation; as in most of these problems
the solution must await the collection and examination of further material. In conclusion, there-
fore, it is proposed to retain the genus Stenoxyethira for the time being with Gnathotrichia as a
junior synonym although the latter should probably more correctly be placed as a synonym of
Oxyethira along with Stenoxyethira excisa. In this case the genus Stenoxyethira may need to be
retained solely for the type-species, minima, although this may also prove to belong in Oxyethira.
Genus PAROXYETHIRA Mosely
Paroxyethira Mosely, 1924 : 670. Type-species: Paroxyethira hendersoni Mosely, by subsequent designa-
tion by Mosely & Kimmins, 1953.
DISTRIBUTION. New Zealand.
This endemic New Zealand genus contains five species although I believe that further species
are at present being described (F. B. Michaelis, pers. comm.). Mosely (1924) originally described
eatoni, tillyardi and hendersoni; hintoni and kimminsi being subsequently described by Leader
(1972) in which paper a key to all males and females was provided. Leader (1970) also discussed
the function of the setal equipment of the early and fifth instar larvae of Paroxyethira, giving very
generalized figures of each but no detailed morphological descriptions. Paroxyethira is charac-
terized by the unique forms of both the male and female genitalia, especially the elongate inferior
appendages and ventral processes of the former, and the general appearance of the larva and its
case. The following diagnosis is based on Leader (1972) and Mosely’s original material (BMNH
collection).
DiaGcnosis. Adult. Forewing length 2:5—3-5 mm: anterior tentorial arms reduced to fine strands medially
(Fig. 126); antennae of approximately 25-40 segments; ocelli 3: thorax (Fig. 126) as in Xuthotrichia: spur
formula 0.3.4: 2 genitalia a modified oviscapt with sternite VI with a small ventral ‘tooth’; posterior mar-
gin of sternite VII with a triangular ventral process; tergite VIII deeply excised dorsally and elongate
postero-ventrally, almost meeting mid-ventrally; sternite VIII small, appearing as a small lobe just above
the projection from VII; segment IX small, mostly concealed by VIII; segment X small, bilobed, each
lobe with a short apical cercus: ¢ genitalia (Fig. 127) with sternite VII with an elongate flattened ventral
process extending beyond the posterior margin of segment IX; segment IX fused, with a deep postero-
dorsal excision such that the segment is incomplete dorsally; tergite X absent; inferior appendages basically
elongate, concave with apices directed inwards; aedeagus with a very characteristic basal, sleeve-like
sheath with the posterior margin produced as a long spine extending almost to the apex of the aedeagus,
latter long and slender with a short, broad proximal region which tapers anteriorly to a distinct constric-
tion just before which arises a spiral process or ‘titillator’.
Larva (Fig. 128). According to Leader (1972) the larva of Paroxyethira differs from that of Oxyethira
only in the possession of a complex spine on the ventral process of the fore-tarsus which resembles the
fan-like seta present on the fore-tarsus of Hydroptila (cf. Fig. 47). As in Oxyethira, the legs of Paroxyethira
are longer and more slender than those of Hydroptila but apart from these general points no further com-
parisons can be drawn. Leader (1972) states that the five species of Paroxyethira can only be distinguished
by the fine sculpturing of the thoracic sternites which, however, tends to be rather an inconsistent feature
and thus not a reliable taxonomic character.
Case (Fig. 128). ‘Purse-type’, constructed of secretion only, oblong and sometimes with outward pointing
anterior and posterior ventral spines as extensions of the ventral seam by which the case is attached to the
substrate (pupal).
BioLoGy. According to Pendergrast & Cowley (1966), the larvae of hendersoni are abundant in
vegetation in lakes, ponds and streams where filamentous green algae occur. Leader (1972)
briefly gives collecting data for each species.
THE GENERA OF THE HYDROPTILIDAE 209
‘penis sheath’
Figs 126-129 Hydroptilini, Oxyethira-group. 126-128, Paroxyethira; (126) adult head and thorax,
dorsal view; (127) generalized ¢ genitalia, lateral view; (128) larva in case, lateral view. 129,
Xuthotrichia, generalized 3 genitalia, lateral view (vide Figs 15, 16).
DISCUSSION. From features of the adult head and thorax and the basic form of the female genitalia,
I consider Paroxyethira to be closely allied to the endemic Australian genus Xuthotrichia Mosely.
The general morphology and habits of the larvae suggest that it may, along with Xuthotrichia,
be an offshoot of the Oxyethira branch of the Hydroptilini.
Genus XUTHOTRICHIA Mosely
Xuthotrichia Mosely, 1934 : 139. Type-species: Xuthotrichia ochracea Mosely, by original designation.
DISTRIBUTION. Australia.
This endemic Australian genus is known from the four original species described by Mosely
(1934) (ochracea, simplex, fimbriata and eskensis), and I believe that more species are being des-
cribed (A. Wells, in /itt.) and there are specimens of an unidentified (and possibly new) species from
Western Australia in the BMNH collections.
The genus is characterized by the unique form of the male genitalia which show considerable
specific variation; they are asymmetrical and often very complex, and identification of species is
very difficult from Mosely’s original descriptions. The females have not been described but from
examples in the unidentified material in the BMNH collections mentioned above the genitalia
appear to bear some resemblance to those of Paroxyethira. The immature stages are unknown
although a figure of an unidentified Australian hydroptilid larva in Riek (1970) greatly resembles
that of Paroxyethira in general appearance (cf. Fig. 128) and, if so, may be the larva of Xutho-
trichia (see Paroxyethira account). The case of the larva figured in Riek (1970) is constructed of
silk and sand-grains although, according to A. Wells (in /itt.), the cases of larvae which she believes
210 J. E. MARSHALL
to be of Xuthotrichia (larvae not described, however) are of secretion only but of a slightly differ-
ent shape to those of Paroxyethira (also of secretion only). It would be necessary to compare
specimens of the two genera, however, before any conclusions regarding affinities could be drawn.
The following diagnosis is based on a study of material in the BMNH collections (Mosely’s
types) and the original descriptions.
Diacnosis. Adult. Forewing length 2-5—3-0 mm: anterior tentorial arms reduced to fine strands medially;
antennae c. 40-segmented; ocelli 3: mesoscutellum similar to that of Hydroptila and Oxyethira (cf. Figs
114, 120); metascutellum pentagonal with slightly anteriorly convergent lateral sides: spur formula
0.3.4: ¢ genitalia (Fig. 129) — sternite VII with a moderately long, slender ventral process; segment VIII
unmodified; segment IX fused, usually obliquely truncate in lateral view, dorsal and ventral margins
more or less excised; tergite X, inferior appendages, subgenital appendages, subgenital plate and bilobed
process very variable; aedeagus long and slender, without a median constriction or spiral process but
sometimes with additional short lateral processes arising just before apex,
BioLoGy. According to A. Wells (in /itt.), the larvae of Xuthotrichia occur in all types of still
water in Australia such as lakes, billabongs, etc.
Discussion. Xuthotrichia may be closely related to Paroxyethira as indicated by adult head and
thoracic features (and also, possibly, by the general form of the larva); these genera may in turn
be allied to the cosmopolitan genus Oxyethira.
Genus TRICHOLEIOCHITON Kloet & Hincks
Leiochiton Guinard, 1879 : 139. Type-species: Leiochiton fagesii Guinard, by monotypy. [Preoccupied by
Leiochiton Curtis, 1831 in Coleoptera.]
Tricholeiochiton Kloet & Hincks, 1944 : 97. [Replacement name for Leiochiton Guinard.]
Synagotrichia Ulmer, 1951 : 81. Type-species: Synagotrichia fortensis Ulmer, by original designation and
monotypy. Syn. n.
DIsTRIBUTION. W. Palaearctic; SE. Asia (Burma, Indonesia).
The genus Tricholeiochiton contained originally only two species, fagesii (Guinard), which is
known locally from Europe and the U.S.S.R., and /acustris Kimmins from Burma (S. Shan
States). Examination of type-material of Synagotrichia fortensis Ulmer (UHZIM loan) reveals
that this genus should be regarded as a junior synonym of Tricholeichiton.
The unique larva of fagesii was first noticed by Bremi who gave an extended description of it
under the name of Hydroptila flabellifera in a letter to Hagen which was published by the latter
in 1864. Hagen considered the species to belong to the genus Agraylea due, possibly, to the shape
of the case which was covered with filaments of ‘Confervae’ (?). However, it is now known that the
cases of Tricholeiochiton are constructed of secretion only and that Bremi’s H. flabellifera con-
sisted of a mixed series of T. fagesii and Agraylea sexmaculata Curtis.
The next reference to the larva was in 1867 when Dr Barker presented a series to the Dublin
Microscopical Society as specimens of Diptera. Then, in 1878, Guinard published figures of the
larva and, albeit rather crudely, the male genitalia, and named the species Leiochiton fagesii. The
species has since been variously referred to as Oxyethira fagesii and O. felina Ris, and was first
recorded on the British list by Mosely (1932) who subsequently recognized the synonymy with
L. fagesii (Mosely, 1939a). The generic name was changed to Tricholeiochiton by Kloet & Hincks
(1944) when it was discovered that Leiochiton was preoccupied in Coleoptera.
The larva of fagesii has been described a number of times but not in any great detail. The
account given below is based on Lepneva (1964) and a study of material donated to the BMNH
by Dr B. Higler (Netherlands) and Dr I. D. Wallace (MCM). A formal description has not been
given since it is understood that Wallace (pers. comm.) proposes to publish a full account based
on recently discovered British material.
Kimmins (1951) described a second species, /acustris, from Burma in which the male genitalia
are almost identical with those of fagesii except for the presence of a long ventral process on the
seventh abdominal sternite. The immature stages of /acustris are unknown.
THE GENERA OF THE HYDROPTILIDAE eal
Figs 130-132 Tricholeiochiton. 130, adult head and thorax, dorsal view; 131, generalized
3 genitalia, lateral view; 132, larva in case, lateral view.
From the form of the male genitalia, wing venation and shape of the metascutellum, Synago-
trichia fortensis Ulmer, from Indonesia (Sumatra), probably represents a third species of Tricho-
leiochiton. Discovery of the larval stages, however, would confirm this view.
Diacnosis. Adult. Forewing length 2:25—3-00 mm: anterior tentorial arms reduced to fine strands medially
(Fig. 130); antennae of c. 37 segments in /acustris; ocelli 3: metascutellum pentagonal, lateral sides slightly
divergent anteriorly: spur formula 0.3.4: lateral processes of abdominal sternite V absent, basal pit with
characteristic sculpturing in fagesii (Fig. 13): 2 genitalia an oviscapt: ¢ genitalia very distinct, but very
similar in the three known species (Fig. 131) — sternite VII sometimes with a long, slender ventral process;
segment VIII unmodified; segment IX fused, produced ventro-posteriorly (obliquely truncate in lateral
view), dorsal and ventral posterior margins deeply incised; segment X forming a dorsal membranous lobe
with a pair of postero-ventrally produced lateral processes which partly enclose the aedeagus; inferior
appendages small, fused baso-medially, contained within ventral incision of IX and concealed by latero-
ventral processes of IX; subgenital plate transverse with down-curved posterior ‘lip’ and produced antero-
ventrally to join the inferior appendages, base of plate with a pair of small seta-bearing processes, junction
of plate and inferior appendage produced anteriorly as a slender truncate process; aedeagus short and
robust with a distinct median constriction, just before which arises a slender spiral ‘titillator’ which has a
slightly dilated apex (homologies of genitalia uncertain).
Larva (Fig. 132). Head long and slender with parallel sides; dorsal ecdysial lines indistinct anteriorly;
tentorial pits distinct; ‘postmental sclerites’ crescent-shaped, undivided, with short blunt postero-median
process (Fig. 23); mandibles typically subtriangular with slender pointed apices, left with bluntly toothed
dorsal blade: thoracic nota paired; three well-developed sternites on prothorax only (Fig. 46); pleurites
with preepisternite fused in all three segments (according to Lepneva, 1964); forelegs short, tibiae with
distinct disto-ventral processes bearing apical spines and a row of short ventral setae; mid- and hindlegs
characteristically very long and slender, achieved by elongation of trochanter, femur, tibia and tarsal
claw: abdomen long, more slender than in other Hydroptilidae, characteristically curved; tergite on
segment IX only; anal prolegs fused to X, anal claw short; dorsal ‘rings’ present on segments III-VIII.
Case (Fig. 132). ‘Purse-type’, oblong, ‘sac-shaped’, sides slightly convex, constructed of secretion only.
Pupa. Undescribed but pupal case with two stalked attachment discs at each anterior corner and one at
each posterior corner.
#4 We J. E. MARSHALL
BioLoGy. According to Lepneva (1964) the larvae of fagesii inhabit stagnant and slow-moving
bodies of water in plant thickets, the pupal cases being attached to the undersides of aquatic
plants.
Discussion. The form of the male aedeagus and the algae-associated habits of the larval stages
indicate that Tricholeiochiton is a member of the Hydroptilini. Although obviously very specialized
with respect to both the male genitalia and the morphology and habits of the larvae, the general
appearance of the latter and the form of the adult head and thorax are suggestive of the genus
Oxyethira, of which branch of the Hydroptilini Tricholeiochiton may be an early offshoot. The
disjunctive distribution of this genus is rather unusual with only two Oriental and one Palaearctic
representatives. The latter may have been due to an early introduction of ancestral stock from the
Oriental region.
Tribe ORTHOTRICHIINI Nielsen stat. n.
Orthotrichiinae Nielsen, 1948 : 186. Type-genus: Orthotrichia Eaton, 1873.
Nielsen erected the subfamily Orthotrichiinae for two genera, Orthotrichia and Ithytrichia, on the
basis of larval morphological and behavioural affinities. Although the adults and larvae are very
distinctive for each genus, the larvae exhibit a number of fundamental similarities which dis-
tinguish them from other members of the Hydroptilinae. However, according to certain adult
features, such as the structure of the head and thorax and the male aedeagus, the Orthotrichiini
appears to have arisen from an early Hydroptilini-type ancestor. I therefore propose to accept
Nielsen’s original grouping but as a tribe of the subfamily Hydroptilinae and with the inclusion
of Ithytrichia remaining open to further speculation; the latter may later be found to form a distinct
unit on its own.
Diacnosis. Adult. Forewing length 2-4 mm; tentorium complete (Figs 133, 139); ocelli 3 or 0; post-
occipital lobes unmodified: thoracic nota variable (Figs 133, 139); mesonotum without transverse suture:
spur formula 0.3.4: abdominal sternite V with typical setate process: 2 genitalia basically a simple oviscapt,
modified in Orthotrichia (Fig. 134): 3 genitalia very characteristic for each genus, highly asymmetrical in
Orthotrichia (Figs 135, 140); aedeagus with spiral ‘titillator’.
Larva. Both Orthotrichia and Ithytrichia have very distinct and specialized larval stages. Their charac-
teristic features, on the basis of which Nielsen (1948) grouped the genera together, are as follows. Labium
broad and flat; mandibles flattened, tapering; general form of the ventral apotome and ‘postmental
sclerites’ (Figs 24, 28); the thoracic sternites (Fig. 42); the comb-like setae of the fore-coxae; the
circle of ‘rings’ on abdominal segment I and the single median caudal filament from segment X. The cases
of the larvae of each genus are also very characteristic, neither being of the typical ‘purse-type’.
Pupa. Typically hydroptilid, mandibles long and untoothed.
Discussion. According to Nielsen (1948) /thytrichia and Orthotrichia may be derived from a com-
mon ancestor which was probably associated with filamentous green algae; this, in my opinion,
probably occurred early in the evolution of the Hydroptilini. Orthotrichia became a highly effi-
cient algal feeder living in plant thickets in sluggish to still water while /thytrichia, possibly in
association with the colonization of rheophilous and semi-exposed habitats, secondarily adopted
the detritus-feeding habit but retained a number of morphological features derivable from algal-
feeding ancestors. Such features which are shared by /thytrichia and Orthotrichia larvae are the
broad, flat labium; the flattened mandibles; the form of the ventral apotomes and postmental
sclerites (cf. Figs 24, 28) and the thoracic sternites (Fig. 42); the comb-like setae of the fore-
coxae; the complete circle of abdominal ‘rings’ on segment I and the possession of a single median
caudal filament which, according to Nielsen (1948), is present in the early instars of Orthotrichia.
On the basis of these larval affinities, Nielsen proposed the subfamily Orthotrichiinae for
Orthotrichia and Ithytrichia and, although the larvae and adults of these two genera have evolved
along different lines, I am retaining Nielsen’s grouping but as a tribe of the Hydroptilinae. The
Orthotrichiini is thus, on the basis of the form of the adult thorax, male aedeagus and fundamental
larval associations with filamentous algae, a highly specialized and probably early offshoot of the
Hydroptilini line. Orthotrichia is a very successful genus which has a wide present-day distribution
THE GENERA OF THE HYDROPTILIDAE PANS
although no species have yet been recorded from the Neotropical and Australian regions. /thy-
trichia is less well represented with only three recognized Holarctic species and possibly one In-
donesian. Until the genus is better known, however, its origins and evolutionary history must
remain speculative.
Genus ORTHOTRICHIA Eaton
Orthotrichia Eaton, 1873 : 141. Type-species: Hydroptila angustella McLachlan, by original designation.
Clymene Chambers, 1873 : 114. Type-species: Clymene aegerfasciella Chambers, by monotypy. [Synony-
mized by Flint, 1966.]
Javanotrichia Ulmer, 1951 : 75. Type-species: Javanotrichia maeandrica Ulmer, by original designation.
Syn. n.
Orthotrichiella Ulmer, 1951 : 79. Type-species: Orthotrichiella ranauana Ulmer, by original designation
and monotypy. Syn. n.
Baliotrichia Ulmer, 1951 : 88. Type-species: Baliotrichia litoralis Ulmer, by original designation and
monotypy. Syn. n.
DISTRIBUTION. Holarctic; Africa; SE. Asia (to New Guinea).
Orthotrichia is a very successful genus with approximately 40 species and with an almost con-
tinuous world-wide distribution, although it has not so far been recorded from Central or South
America or the Australian region. The genus is well represented in the Palaeotropical regions of
Africa and SE. Asia, through Indonesia (Sunda Islands) to New Guinea (Papua) and, possibly,
Fiji and Borneo (unidentified species, BMNH collections, det. J. E. Marshall). The adults are
characterized by the uniquely developed asymmetrical male genitalia and the absence of ocelli
(cf. Hydroptila which may be distinguished by the presence of postoccipital scent organs in the
males). The larvae are associated with aquatic vegetation in standing waters and are highly
specialized for feeding on the cell contents of green filamentous algae.
The genus was erected by Eaton (1873) for angustella (McLachlan) and atra (Hagen), the latter
being transferred to Stactobia by McLachlan (1884). O. tetensii Kolbe was subsequently found by
Neboiss (1963) to be a junior synonym of Hydroptila costalis Curtis (vide Oxyethira) and Flint
(1966) showed Clymene aegerfasciella to be a senior synonym of Orthotrichia americana Banks.
The larva of costalis was described in detail by Nielsen and that of angustella by Jacquemart
(1962b), although no distinguishing features were indicated in the latter work for separating the
two species. It should be noted that Jacquemart’s figure of the dorsal view of the head of angus-
tella does not make it clear that the anterior beak is an extension of the labrum; it has also been
drawn pointing in the opposite direction to that as figured by Nielsen.
Regarding the genera described by Ulmer (1951) from the Sunda Islands, larvae are described
by Ulmer (1957) for Orthotrichiella ranauana and Baliotrichia litoralis. The relationships of these
two genera, as well as that of Javanotrichia, with Orthotrichia are discussed below.
The following generic diagnosis is based on the literature and examination of adult and larval
material in the BMNH collections.
Diacnosis. Adult. Forewing length 2-4 mm; wings markedly attenuate, forewings with a row of black,
flattened basal subcostal setae in males of the angustella-group: tentorium (Fig. 133) complete; antennae
of 30-40 segments; postoccipital lobes prominent, subspherical, but not modified as scent-organs; ocelli
0: thorax (Fig. 133)-—metascutellum characteristically rectangular: spur formula 0.3.4: 2 genitalia a
modified oviscapt (Fig. 134), segment VIII fused and sclerotized, segments IX and X membranous, IX
short: ¢ genitalia (Fig. 135) characteristically asymmetrical; sternites VI and VII variously with ventral
processes or tufts of setae; segment VIII unmodified ; segment IX fused, often with asymmetrical, postero-
lateral processes; inferior appendages usually small and fused medially, but well developed, porrect and
strongly curved in costalis; subgenital appendages indistinct or absent; bilobed process present (Bp);
tergite X forming a large, asymmetrical dorsal plate, ventrally concave, often with laterally directed dorsal
spines; beneath X is a variously shaped, slender, asymmetrical sclerite usually with two asymmetrical
processes, one of which is long and slender and lies diagonally beneath X and alongside the aedeagus
(according to Nielsen, 1970, a similar structure was described by Ulmer, 1951, for Javanotrichia, Ortho-
trichiella and Sumatranotrichia); aedeagus very long and slender with distinct proximal and distal halves
near the junction of which arises a slender, spiral ‘titillator’.
214 J. E. MARSHALL
Larva (Fig. 136). Slightly depressed dorso-ventrally: dorsal ecdysial lines of head distinct; postmental
sclerites and anterior ventral apotomes as in Fig. 28; labrum asymmetrical with a beak-like median
process (Fig. 137); mandibles slender, flat, asymmetrical (Fig. 32): thoracic nota paired; mid- and hind-
legs just slightly longer than forelegs; fore-tibia without a ventral process or modified setae; each tarsus
with a distal, flattened, plate-like spur; fore-coxa with comb-like rows of setae; preepisternite of pro-
thorax free; paired anal sternites present on all three thoracic venters, fused in prothorax (Fig. 42):
abdomen cylindrical, slightly broader at midlength; segment II with a pair of lateral ‘humps’; dorsal
‘rings’ present on segments III—-VIII, segment I with a complete circle of ‘rings’ near anterior margin;
tergite present on segment IX only; median caudal filament in early instars only.
Case (Fig. 138). Of the ‘wheat-seed’ type, almost circular in cross-section, tapering gradually towards
slit-like anterior and posterior openings; convex dorsal surface characteristically ribbed, ventral surface
flat.
Pupa. Typically hydroptilid but mandibles short; pupal case attached by a single attachment disc at
each end.
BioLocy. The larvae prefer slowly running and sometimes almost stagnant water in plant thickets.
According to Nielsen (1948) the larva bites into each cell of a filament of green algae and enlarges
the hole by inserting the labral beak; the cell contents are then sucked out. This is a more efficient
mechanism than that found in Agraylea, Hydroptila and Oxyethira. In addition there have been
two reports (Burton & McRae, 1972; Disney, 1973) of Orthotrichia larvae feeding on Simulium
(Diptera) larvae, but it was not evident as to whether these were cases of active predation or of
the hydroptilids mistaking the Simulium for algal filaments.
SPECIES-GROUPS (see checklist). The 36 species may be split into four main groups on the basis of
the form of the male genitalia.
The angustella-group: exclusively ‘Old World’ with the majority of representatives being re-
corded from the regions of Africa and SE. Asia (the latter region may include the species originally
described by Ulmer, 1951, in Javanotrichia, Baliotrichia and Orthotrichiella from Indonesia, see
group 2); angustella is the only truly Palaearctic representative and has a very wide distribution
extending from northern Europe (including the British Isles) to northern Africa and from Iberia
to the Urals; the group is characterized by the development of the lateral processes of segment
IX in the males and the row of subcostal scales on the forewings of the males of most species.
The /itoralis-group: the species mentioned above from Indonesia originally described by Ulmer
(1951; 1957) as Javanotrichia, Baliotrichia and Orthotrichiella may form a distinct group; it also
includes species described by Schmid (1958a) from Sri Lanka.
The costalis-group: a small ‘Old World’ group; costalis is the most common species and has a
wide European distribution (including the British Isles) and has also been recorded from the
Sudan, U.S.S.R. and Japan; the group is distinguished by the absence of lateral processes on
segment IX and the small, more or less symmetrical inferior appendages in the males.
The aegerfasciella-group: principally Nearctic in distribution with one well-established Palaearc-
tic representative (tragetti); the group is characterized by the lateral development of the asym
metrical inferior appendages of the males.
A fifth group may also be recognized.
The kokodana-group: this consists of two distinct species from Papua New Guinea charac-
terized by the simplicity of the male genitalia. (N.B. there are unassociated 2 specimens of Ortho-
trichia spp. from Fiji and Borneo in the BMNH collections.)
GENERIC SYNONYMS. The genera discussed below are here considered to be junior synonyms of
Orthotrichia.
Javanotrichia Ulmer contains three species, maeandrica Ulmer and curvata Ulmer from In-
donesia and dampfi Ulmer from Egypt. Ulmer (1951) distinguished the genus from Orthotrichia
solely on venational characters but, as can be seen from other adult features such as the absence
of ocelli, spur formula 0.3.4 and the distinctive asymmetrical form of the male genitalia in
Javanotrichia, these two genera are probably synonymous. Examination of type-material (UHZIM
loan) reveals that the structure of the thorax of J. curvata is typical of Orthotrichia; in addition the
male genitalia are very similar to those of O. avicularis Kimmins from India. J. dampfi greatly
THE GENERA OF THE HYDROPTILIDAE ZS
Figs 133-138 Orthotrichia. 133, adult head and thorax, dorsal view; 134, generalized 2 genitalia,
ventral view; 135, generalized ¢ genitalia, ventral view and aedeagus (vide Figs 15, 16); 136, larva,
dorsal view; 137, larval labrum, dorsal view; 138, pupal case, lateral and dorsal views.
resembles other African Orthotrichia species, such as O. benguelensis, especially with respect to
the spinose lateral processes of segment IX in the male. The immature stages are unknown.
Baliotrichia Ulmer contains /itoralis Ulmer from Indonesia (Bali) and four species from Sri
Lanka (Schmid, 1958a). As in Javanotrichia the genus is only distinguished on minor venational
features. According to Schmid (1958a) the male genitalia of B. udawarama bear a ‘curious simi-
larity’ to those of O. extensa Martynov from India while, in my opinion, the genitalia of B. /itoralis
are most like those of O. avicularis Kimmins. The larva and case of B. /itoralis, as described by
Ulmer (1957), are unmistakably those of Orthotrichia and thus, taking both adult and larval
features into account, I propose that Baliotrichia be placed as a synonym of Orthotrichia.
Orthotrichiella Ulmer is a monotypic genus from Indonesia again only distinguished from
Orthotrichia on minor venational features. Examination of type-material (UHZIM loan) indicates
216 J. E. MARSHALL
that the two genera are synonymous and this is supported by the appearance of the immature
stages described by Ulmer (1957).
In conclusion these three small tropical genera provide a natural geographical link with the
Orthotrichia species from the mainland and New Guinea, this genus being previously unrecorded
from the Indonesian region.
Discussion. The genus Orthotrichia is a specialized and very successful recent member of the
Hydroptilinae with a wide continuous present-day distribution. It is the only hydroptilid genus
which has a truly tropical component (cf. the Stactobiini which, although tropical, are confined
to the relict cool montane areas of SE. Asia and Africa) and probably owes its success to its
tolerance of still water conditions and its specialized, efficient larval feeding habits.
Genus ITHYTRICHIA Eaton
Ithytrichia Eaton, 1873 : 139. Type-species: Ithytrichia lamellaris Eaton, by original designation and
monotypy.
Saranganotrichia Ulmer, 1951 : 83. Type-species: Saranganotrichia decussata Ulmer, by original designa-
tion and monotypy. Syn. n.
DISTRIBUTION. Holarctic.
Ithytrichia is a small genus at present containing only four species. J. /amellaris is the most com-
mon Palaearctic species while c/lavata Morton, originally described from North America, was
found by Tjeder (1930) to occur in Sweden and has since been recorded from France, Great
Britain and Finland (Karelia) and may prove to be the only hydroptilid with a truly Holarctic
distribution (cf. Agraylea multipunctata which may involve a species complex). /. mazon Ross from
Illinois is known only from the holotype male while bosniaca Botosaneanu, a Jugoslavian species
described from the immature stages only, is regarded as a doubtful species by Botosaneanu (1967).
The larva of /amellaris has been described several times, the most detailed account being given
by Nielsen (1948). That of clavata has not positively been identified although the larva referred
to by Needham (1902) and figured by Ross (1944), as well as that described by Wiggins (1977), is
probably this species.
Saranganotrichia decussata Ulmer from Indonesia (Java) is known from both the adult and
larval stages (Ulmer, 1951 and 1957 respectively) of which the latter show distinct affinities with
Ithytrichia in the form of the case, larval head shape and possession of abdominal ‘buds’. How-
ever, the male genitalia of Saranganotrichia are atypical of Ithytrichia and thus cast doubts on the
associations of the adults and larvae. Examination of Ulmer’s specimen of decussata (adult)
failed to reveal any diagnostic characters as the microscopical preparation was in a very poor
condition. Nevertheless, on the basis of the affinities of the larval stages, it is proposed here that
Saranganotrichia be placed as a synonym of /thytrichia until the true identity of the adults is
discovered.
The following generic diagnosis is based on the literature and on examination of material in
the BMNH collections.
Diacnosis. Adult. Forewing length 3 mm: tentorium complete (Fig. 139); antennae with 20-25 segments;
ocelli 3: thorax (Fig. 139) with mesoscutellum with anterior margin convex, posterior edge separated from
posterior margin of mesonotum by a narrow strip; metascutellum with a strongly convex anterior margin:
spur formula 0.3.4: abdominal sternite V with typical setate processes, although shorter than usual and
in a faintly sculptured pit (Fig. 14): 2 genitalia an oviscapt, segment VIII with median ventral sclerite:
3 genitalia distinct (Fig. 140); sternite VI sometimes with a short ventral spine; segments VI and VII un-
modified; IX and X membranous dorsally, former incomplete ventrally but with lateral sides tapering
posteriorly (‘side-pieces’ (Tjeder, 1930)) and with a hook-like structure arising ventrally near the apex; in
lamellaris tergite X bears two characteristic parallel rod-like sclerites; inferior appendages slender, arising
from antero-ventral margin of segment IX and extending to its lateral apex; subgenital plate with short
postero-medial processes bearing short setae and attached basally to the apical hooks of IX; aedeagus long,
well developed, with distinct proximal and distal halves divided by a median constriction by which arises
the spiral ‘titillator’.
THE GENERA OF THE HYDROPTILIDAE ay
Larva (Fig. 141). Head broad with slight median lateral bulges in region of ‘eyes’, flattened, tapering
anteriorly both dorso-ventrally and laterally: dorsal ecdysial lines distinct; postmental sclerite hemi-
spherical, ventral apotome V-shaped, prominent (Fig. 24); labrum long and narrow, curved sides tapering
anteriorly, anterior margin of sclerite with three shallow indentations, narrow anterior membranous por-
tion with straight anterior margin; mandibles very flat (even more so than in Orthotrichia, cf. Fig. 32),
left with short lower blade, serrated upper blade and median brush, apices of both mandibles pointed with
a small basal tooth on upper blade: pleurites of prothorax only with three distinct sclerites; prothoracic
venter with unpaired anal sclerite (as in Orthotrichia) with lateral ends fused with posterior corners of
nota (Fig. 42); median oral sternites present on meso- and metathoracic venters only: legs short, subequal,
forelegs without trochanteral, femoral or tibial processes; fore-coxae with comb-like rows of setae:
abdomen greatly compressed laterally, segment I very small, II with large lateral processes, III-VI with
characteristic dorsal and ventral ‘bud-like’ processes, VII with dorsal process only, VIII with postero-
ventral projection; segment I with complete circle of dorsal ‘rings’; [LX with tergite which projects ‘roof -
like’ above anal prolegs (fused to X) and covering the base of the single median caudal filament.
Case (Fig. 142). This consists of secretion only and resembles a pumpkin seed; narrow anteriorly with
an oval ridged opening flanked by the lateral valves, ‘slit? only open medially. The thorax of the larva may
be extended through the narrow anterior opening and, due to the ability of the thorax and first abdominal
segment to rotate on the rest of the abdomen, the case is usually carried horizontally (cf. the lateral com-
pression of the abdomen). Consequently, the ventral surface of the case is slightly more convex while the
anterior ventral margin is slightly indented so that the hindlegs may be held free of the case.
Pupa. Typical; mandibles long, without teeth. The pupal case is sealed and held, with the ventral side
towards the substrate, by a single median posterior filament with a broad attachment disc, and an anterior
cylindrical plug which also broadens into a single attachment disc.
BioLocy. According to Lepneva (1964) the larvae of /amellaris prefer running water, brooks and
rivulets in plant thickets while Nielsen (1948) records them as being distinctly rheophilous,
occurring in the strong current on exposed parts of submerged vegetation and smooth, current-
swept stones, the cases being held flat against the substrate. I. D. Wallace (pers. comm.) has
collected /thytrichia larvae (probably /amellaris) from amongst willow roots in fine silt along with
larvae of an Orthotrichia species. According to Ulmer (1957) the larvae and pupae of S. decussata
occur in flowing water amongst moss and leaves, in waterfalls, spring-channels and mountain
brooks.
Although Nielsen (1948) did not observe the feeding behaviour of /amellaris larvae, he considers
them to be detritus-feeders unlike the specialized algal-feeding larvae of Orthotrichia to which
Ithytrichia is supposed to be allied. The flattened labial lobe may be adapted to serve as a shovel
for scooping up diatoms and other detritus from the substrate, the food then being conveyed to a
groove on the ventral surface of the labrum along which it passes to the pharynx. Nielsen considers
the anterior tapering of the head to be associated with such feeding habits while the flattened
pointed mandibles, although apparently not adapted for scraping or scooping, may be derived
from the Orthotrichia-like condition (cf. Fig. 32), their function being taken over by the labium.
INCERTAE SEDIS
The following group of essentially monotypic and geographically distinct genera have not been
assigned to any particular tribe due to the inadequacy of the existing data, the insufficiency or
unavailability of well-preserved material or, as in the case of Caledonotrichia and Dibusa, the
unique morphological features which cannot be likened to those of any other hydroptilid genus.
All would, however, appear to be members of the subfamily Hydroptilinae; Caledonotrichia and
Dibusa have typically hydroptilid larvae while all have hydroptilid adult features except Dicaminus
which is known from the larval cases only. Electrotrichia is only known from Baltic Amber and
appears to be a distinct but now extinct genus.
Genus MACROSTACTOBIA Schmid
Macrostactobia Schmid, 1958a : 46. Type-species: Macrostactobia elawalikanda Schmid, by original
designation and monotypy.
DISTRIBUTION. Sri Lanka.
This monotypic genus is known only from the original series from Sri Lanka, of which there
218 J. E. MARSHALL
Figs 139-142 Ithytrichia. 139, adult head and thorax, dorsal view; 140, generalized ¢ genitalia,
lateral view (vide Figs 15, 16); 141, larva, lateral view; 142, larval case, lateral view.
is a single paratype male in the BMNH collections. The adult diagnosis given below is based on
Schmid (1958a), the main characteristic being the unique form of the male genitalia. The larvae
are unknown.
DiaGnosis. Adult. Forewing length 2-5-3-0 mm; head without large postoccipital lobes; antennae 18-
segmented; ocelli 3: mesoscutellum subtriangular (BMNH specimen): ¢ genitalia with sternites VII and
VIII very narrow; sternite IX enlarged ventrally but not invaginated into the preceding segments; segment
X of two distinct halves; inferior appendages complex and fused along their midlength; aedeagus a long
narrow tube.
BroLoGy. The species is reported to inhabit ‘rivers at moderate altitudes’.
Discussion. Schmid places Macrostactobia at the base of the Macrostactobia—Parastactobia-
Plethus—Chrysotrichia branch (of the Stactobiini), its primitiveness being indicated by its more
complete wing venation and its relatively larger size. The genitalia are stated to exhibit the typical
(stactobiine) characters of the group, the line of evolution resembling that of Parastactobia as
suggested by the great development of segment IX at the expense of those preceding, and the long
and rigid appendages.
The male genitalia are unique but while the antennae are of 18 segments only, as is typical of
the Stactobiini, the metathorax does not have the rectangular form characteristic of this tribe.
Consequently I can give no positive indications as to the true affinities of this very specialized
monotypic genus but can merely stress the need for the examination of additional material.
Genus DIBUSA Ross
Dibusa Ross, 1939 : 66. Type-species: Dibusa angata Ross, by original designation and monotypy.
DisTRIBUTION. U.S.A. (N. Carolina, Tennessee, Arkansas, Kentucky, Oklahoma).
This North American genus is known only from the type-species. Ross (1939) first described
the male and the larva has since been described by Wiggins (1977). The form of the male genitalia
is unique within the Hydroptilidae but the associated larvae show unequivocally that Dibusa is a
true member of the family. The following generic diagnosis is based on Ross (1939; 1944) and
Wiggins (1977) and examination of larvae (USNM loan).
DiaGnosis. Adult. Forewing length 5-5 mm: head and antennae unmodified; ocelli 0; wing venation
relatively complete, wings elliptical, ovate; spur formula 1.3.4: ¢ genitalia with segment IX retracted into
THE GENERA OF THE HYDROPTILIDAE 219
eighth, with sclerotized sides and a membranous dorsum; tergite X fused with IX, divided into a pair of
lateral sclerotized plates separated on the meson by membranous folds and curved downward at the apex
to form a short hook; the ‘cerci’ arise as a small lobe in the middle of the lateral margin at the base of
tergite X; inferior appendages biramous, ventral lobe slightly enlarged and upturned at apex, dorsal lobe
more slender, out-curved and widest at middle; aedeagus with a filiform basal portion, markedly enlarged
near base, and a semimembranous irregularly tapering pointed apex.
Larva (Figs 143, 144), after Wiggins (1977). Head suboval in lateral view (i.e. not compressed): distinctly
paired thoracic nota onall segments; pleurites fused on all thoracic segments; legs short, subequal, robust;
tarsal claws unique being stout and strongly curved with a thick, blunt spur basally on each (but smaller
than that of Stactobiella, cf. Figs 144 and 80): abdomen moderately distended, not markedly depressed
or compressed, intersegmental grooves distinct; abdominal tergites present only on segment IX; dorsal
‘rings’ present on segments I-VIII; anal prolegs fused to segment X.
Case (Fig. 145). Typically ‘purse-like’, of two adpressed silken valves covered with pieces of the red
freshwater alga Lemanea.
BioLoGy. The larvae collected by Neff and Resh (Wiggins, 1977) were associated with the red
freshwater alga Lemanea, the pupal cases being attached to the base of the algal thallus.
DISCUSSION. The relationships of Dibusa are not clear because no material was available for an
examination of the diagnostic features of the adult head and thorax. However, from the general
features of the adults, larvae and cases, Dibusa would appear to be a very specialized early off-
shoot of the tribe Hydroptilini. The relatively complete adult wing venation and the generalized
form of the larva with its ‘purse-type’ case are very primitive features whereas the absence of
ocelli, the unique male genitalia and the larval associations with Lemanea (and not the usual, more
common green filamentous algae as in the Hydroptilini) indicate specialization. The more
primitive features are suggestive of affinities with Agraylea (Hydroptilini) and Nothotrichia (in-
certae sedis) (vide Flint, 19675).
Genus NOTHOTRICHIA Flint
Nothotrichia Flint, 1967b : 56. Type-species: Nothotrichia illiesi Flint, by original designation and
monotypy.
DISTRIBUTION. South America (Chile).
This monotypic genus has so far been recorded only from Chile, the type being collected in the
Cautin Province in 1957 by Professor J. Illies with further specimens being taken by Flint and
Cekalovic in 1966.
Flint (1967b) provides the basis of the generic diagnosis given below which is supplemented by
observations on the head and thoracic structure as seen in a female specimen of i/liesi (USNM
loan). The larval stages are unknown.
DiaGnosis. Adult. Forewing length 3-5 mm: head and antennae unmodified; ocelli 3; wings not markedly
narrowed but with pointed apices; venation relatively complete; spur formula 1.3.4: thorax (Fig. 146);
mesoscutellum without transverse suture, anterior margin strongly convex; metascutellum deep, sub-
quadrate, but with strongly convex anterior angles: ¢ genitalia (after Flint, 19675) with segment IX
incomplete dorsally, with a broad flap from dorsal angles; tergite X membranous, posterior margin
trilobate; inferior appendages broad, large, ‘mitten-like’, with a small baso-dorsal ‘thumb’; aedeagus with
slightly expanded apex and apico-lateral process arising at midlength and contiguous with the central
aedeagal duct, non-spiral ‘titillator’ present.
DISCUSSION. This unique endemic Chilean genus may represent a descendant of an early offshoot
of the Hydroptilini branch which evolved in isolation in this part of the Neotropical region. This
is indicated by the relatively complete venation, presence of 3 ocelli, presence of a fore-tibial spur,
an undivided mesoscutellum and the non-spiral ‘titillator’ of the aedeagus. The male genitalia
appear to be derived from the basic hydroptilid form but are specialized in that they are greatly
simplified. Discovery of further species and the immature stages may give some insight into the
true relationships of this genus.
220 J. E. MARSHALL
144
spur
Figs 143-145 Dibusa angata Ross. 143, larva, lateral view; 144, larval tarsus, lateral
view; 145, larval case, lateral view.
Genus DICAMINUS Miller
Dicaminus Miller, 1879a : 39. Type-species: Diaulus ladislavii Miller, by subsequent monotypy.
Diaulus Miller, 1879c : 142. Type-species: Diaulus ladislavii Miller, by monotypy. [Synonymized by
Fischer, 1961.]
DISTRIBUTION. South America (Brazil).
Miiller (1879a) described a number of caddis cases from Brazil amongst which were some very
distinct forms which had two dorsal ‘chimneys’ and were covered in diatoms. Miller considered
these to belong to hydroptilid larvae and described them under the generic name of Dicaminus
(Latin: of two chimneys), although neither a description of the larva nor a species name were
given. In subsequent papers Miller (1879c; 1880) refers to these cases as being of the hydroptilid
Diaulus ladislavii, thus naming the species and giving Diaulus (the Greek equivalent of Dicaminus)
as the generic name. As stated by Fischer (1961), Dicaminus takes priority over Diaulus and con-
sequently receives its type-species.
The following description of the cases is from Miller (1879c).
DESCRIPTION. The strongly laterally compressed oval cases have slit-like anterior and posterior
openings and are covered in diatoms. On the dorsal margin they bear two (rarely three) cylindrical
chimneys which facilitate the passage of water through the case without, according to Miiller,
the incessant exertions of the larva as seen in other hydroptilids where the case simply has slits
at either end. The pupal cases are fixed vertically along their whole ventral margin to the upper
side of stones ‘and often these little houses form large villages of a rather picturesque aspect’. The
larval case is reported to be 2:5 mm long by 0-75 mm broad.
Discussion. The relationships and validity of this genus are not known since only the cases of the
immature stages have been described. Miiller’s original material, which has not yet been traced,
may contain larvae and pharate adults which will permit a more positive identification.
Genus ORPHNINOTRICHIA Mosely
Orphninotrichia Mosely, 1934 : 138. Type-species: Orphninotrichia maculata Mosely, by original designa-
tion and monotypy.
DISTRIBUTION. Australia (New South Wales).
The holotype of this endemic Australian genus is the only known specimen and it is in such a
THE GENERA OF THE HYDROPTILIDAE POA
poor condition (BMNH collection, slide mount) that details of the genitalia and thorax cannot be
interpreted clearly.
Mosely erected the genus on the basis of the unique wing venation while the male genitalia also
appear to be unlike those of any other hydroptilid genus. Since the thoracic structure, female geni-
talia and larval stages are unknown, the genus cannot be placed with any certainty at present.
However, from the simplicity of the male genitalia and the form of the aedeagus, with its distinc-
tive spiral ‘titillator’, the genus may have affinities with the Hydroptilini, differing from Hydrop-
tila in the possession of ocelli, the absence of cephalic scent-organs and venation, and from
Oxyethira in venation and spur formula.
DriaGnosis. Adult. Forewing length 2:75 mm: tentorium incomplete; antennae 32-segmented; ocelli 3:
spur formula 0.2.4: ¢ genitalia with sternite VII bearing a short spinous process; segment VIII unmodified ;
segment IX fused, deeply excised dorsally, appearing bifurcate from above and containing the mem-
branous tergite X which tapers posteriorly to a pointed apex; subgenital plate not distinguishable unless
it is fused with or replaces the ‘inferior appendages’; ‘inferior appendages short but slender, fused along
almost entire mid-line except for a short distance apically (this structure may represent the subgenital
plate, the inferior appendages being absent or vestigial as the two small medio-ventral processes of segment
1X); aedeagus with a broad proximal region separated from the longer more slender distal region by a
median constriction just anterior to which arises the spiral process or ‘titillator’.
Genus CALEDONOTRICHIA Sykora
Caledonotrichia Sykora, 1967 : 585. Type-species: Caledonotrichia illiesi Sykora, by original designation.
DISTRIBUTION. New Caledonia.
This small endemic New Caledonian genus contains two species, , illiesi and minor, both des-
cribed by Sykora (1967) from material collected in 1966 by Professor J. Illies. I have been able to
examine adult, larval and pupal material of unidentified and undescribed species of Caledono-
trichia kindly loaned to me by Professor H. H. Ross of Athens, Georgia.
The genus is unique within the Hydroptilidae with respect to the very distinctive male genitalia,
the corresponding parts of which, as noted by Sykora, are difficult to homologize with those of
any other genus. The following diagnosis is based on Sykora (1967) and examination of Ross’s
material (the larvae have not previously been described).
Diacnosis. Adult. Forewing length 2:5—3-0 mm: head unmodified; tentorium complete; antennae c. 30-
segmented: wings elongate, tapering to narrow apex; base of forewing with costal area thickened with a
dense fringe of black androconia, oval patch of androconia between Cu, and Rs in centre of wing (males):
mesoscutellum without transverse suture: spur formula 0.3.4: 2 genitalia an oviscapt: 3 genitalia charac-
terized by the inferior appendages (‘claspers’ (Skyora, 1967)) which are ‘bean-shaped’ with dorsal and
ventral lobes; ‘subgenital plate’ of Sykora with two long rods; tergite X short, membranous; aedeagus
simple, long, rod-like.
Larva (Fig. 147). Head relatively short, wide posteriorly, narrowing anteriorly; ‘postmental sclerites’
difficult to interpret, but apparently U-shaped: body somewhat compressed dorso-ventrally : thorax widest
at segment III; mid- and hind-legs held out more to side than in, for example, Hydroptila; legs short,
subequal; tibia of each leg with a short apico-ventral process bearing two stout, blunt spines; tarsal claw
very small, hook-like; preepisternite free on all segments, epimeron of each apparently fused; single pair
of anal sternites present on prothoracic venter only: abdomen not greatly distended, tapering slightly
back from metathorax; intersegmental grooves distinct; dorsal ‘rings’ present on segments II-VIII; in
dorsal view the Malpighian tubules can be seen just under the abdominal cuticle, one pair passes along
the abdomen laterally and has a distinct loop in each of segments II—VII (possibly having some special
osmoregulatory or excretory function ?); tergite IX present; anal prolegs fused to sides of segment X.
Case (Fig. 148). This consists of secretion only and is dorso-ventrally compressed with a central tubular
section (in which the larva lives) surrounded by a folded lateral fringe by which the case is probably fixed
to the substrate. The central region has two small dorsal holes (possibly to aid water circulation) and the
whole case characteristically ‘curls up’, at least in fluid-preserved specimens.
BIoLoGy. There are no data available on this subject but, from the general appearance of the larva
and its case, the genus would seem to be associated with fast-running water habitats (case fixed
and flattened).
222 J. E. MARSHALL
Malpighian
tubule
Figs 146-148 Incertae sedis. 146, Nothotrichia, adult head and thorax, dorsal view.
147-148, Caledonotrichia; (147) larva, lateral view; (148) case, dorsal view.
Discussion. Caledonotrichia is a very distinct hydroptilid genus both with respect to the adult and
immature stages, neither of which provide any clues as to the relationships of the group. It would
appear to have evolved in complete isolation on the island of New Caledonia to which it is
endemic.
Genus ELECTROTRICHIA Ulmer
Electrotrichia Ulmer, 1912a : 42. Type-species: Electrotrichia subtilis Ulmer, by monotypy.
DISTRIBUTION. Baltic Amber (therefore possibly Eocene, of western Palaearctic origin).
This genus is known from specimens of the type-species in Baltic Amber only; no material was
available for examination. From Ulmer’s original description (ocelli present; spur formula 0.3.4;
forewing length 2:0-2-6 mm; male genitalia figured) it is impossible to place the genus with any
certainty within the Hydroptilidae, although the narrow tapering wings and spur formula suggest
that it may have affinities with the Hydroptilinae.
Phylogeny of the Hydroptilidae
The family Hydroptilidae may be derived from a primitive glossosomatid-like ancestor early in
the evolutionary history of the Trichoptera. Unlike the typical glossosomatids which construct a
primitive ‘saddle-type’ transportable case, the larvae of these early hydroptilid ancestors probably
adopted the more advanced type of ‘purse-like’ case. This basic ancestral stock then appears to
have given rise to two distinct lines which have resulted in the present-day subfamilies, the
Ptilocolepinae and the Hydroptilinae. The former retained primitive adult features similar to
those of the other Rhyacophiloidea, especially the Glossosomatidae, while the latter evolved the
more characteristic form associated with present day adult Hydroptilidae.
THE GENERA OF THE HYDROPTILIDAE 223
The larvae of the Ptilocolepinae have retained the basic form of the hypothetical generalized
hydroptilid and in addition they exhibit the primitive, cool montane habitat preferences sup-
posedly shown by the early, ancestral Trichoptera. Thus the Ptilocolepinae are now restricted to
cool springs in shaded forest regions in the mountainous areas of western Europe and North
America. In the Hydroptilinae, on the other hand, the present-day success of the subfamily may
be due to the adaptive radiation of the larval stages into a wide variety of freshwater aquatic
habitats. These range from springs and seeps to the larger, moderately flowing rivers and streams.
Certain groups have become structurally, behaviourly and physiologically adapted for living in
conditions requiring specialized modifications. These include the splash zones of waterfalls and
rapids, exposed substrates in the more rapid sections of rivers and streams and, at the other extreme,
vegetation thickets in slowly moving to almost static, larger bodies of water.
An early member of the ancestral Hydroptilinae stock possibly gave rise to two, or perhaps
three, subsidiary evolutionary lines. The first led to the Stactobiini and possibly also the Leuco-
trichiini, although it is more likely that the latter arose from a distinct, but closely related, ancestor.
The second line eventually radiated into the Ochrotrichiini, Neotrichiini, Hydroptilini and
Orthotrichiini which appear to have diverged relatively more recently but which have each under-
gone such remarkable larval morphological modifications in relation to habitat specialization
that the larvae at least are quite distinct. A study of both fundamental adult and larval characters
has revealed a number of basic relationships between the four last mentioned tribes which, al-
though not yet fully satisfactorily resolved, are suggestive of the following evolutionary pathways.
The early ancestors of the second (third ?) branch of the Hydroptilinae may have given rise to
two distinct and independent Nearctic—Neotropical tribes, the Ochrotrichiini and the Neotrichiini.
This may have occurred before the larvae adopted associations with green filamentous algae as is
typical of the remaining two tribes, the Hydroptilini and the Orthotrichiini. The latter tribe
would appear to be the most advanced tribe since the larvae are highly specialized algal feeders,
at least in Orthotrichia, and are often capable of tolerating stagnant and polluted conditions.
The relationships of the genera and generic groups within each tribe have been discussed in the
relevant sections of the systematic account. The evolutionary steps have been summarized in
Chart II.
Genera here transferred from the Hydroptilidae
As a result of a study of all genera previously attributed to the Hydroptilidae the following three
small, geographically restricted groups are considered to belong to other Trichopteran families
as indicated.
Padunia Martynov, 1910 (= Uenotrichia Tsuda, 1942); to GLOSSOSOMATIDAE: PROTO-
PTILINAE.
Tsukushitrichia Kobayashi, 1964; possibly a synonym of Kibuneopsychomyia Tsuda, 1942; to
PSYCHOMYIIDAE.
Petrotrichia Ulmer, 1910; possibly related to the primitive, unplaced African genera Petro-
thrincus Barnard and Hydrosalpinx Barnard.
Material examined: Padunia spp. (loaned from the Lepneva collection, ZI); Petrotrichia
palpalis Ulmer, paratypes, BMNH;; the transfer of Tswkushitrichia is based on a study of the orig-
inal description and figures, material being unavailable for study.
Genus PADUNIA Martynov (Glossosomatidae: Protoptilinae)
Padunia Martynov, 1910 : 425. Type-species: Padunia adelungi Martynov, by monotypy.
Uenotrichia Tsuda, 1942 : 228. Type-species: Uenotrichia fasciata Tsuda, by monotypy. [Synonymized by
Fischer, 1971.]
DISTRIBUTION. Asia (central and eastern U.S.S.R.; Korea).
This small Asian genus contains four species described by Martynov from the U.S.S.R.
(adelungi, forcipata, lepnevae and bikinensis) and one from Korea originally described as Ueno-
trichia fasciata Tsuda. Fischer (1971) synonymized these two genera but wrongly gave preference
to Uenotrichia since Padunia clearly snould have date priority.
224
Ptilocolepinae
—_
\
Hydroptilinae ik
J. E. MARSHALL
Palaeagapetus
Ptilocolepus
Stactobia
Plethus
Madioxyethira
Scelotrichia
Ce.
[ \ Pseudox yethira
Paras tactobia
Chrysotrichia
i
Bredinia
é Cn Stactobiella
Catoxyethira
Leucotrichia-gp.
ee Alisotrichia
/ Rhyacopsyche
Metrichia
Ochrotrichia
— ee
Neotrichia
CO Mayatrichia
Caledonotrichia
Dibusa
Allotrichia
{ Agraylea
i Ugandatrichia
Microptila
| - Dhatrichia
Hydroptila
Stenoxyethira
L Oxyethira
Xuthotrichia
Paroxyethira
Tricholeiochiton
Orthotrichia
e Ithytrichia
DISTRIBUTION FOOD HABITAT
= is)
Se) ae
avv (= ere
SCS8 gues
825084
rile dane neat, = 4
X Bryophyte montane-
> 4 ragments ;
springs
Xx ) madicolous
X X fast streams
X X
Xx
X
4
Xx
Xx
XX Detritus | fast
x streams
. x { rapids
X springs &
re waterfalls ak.
X Xx streams
xX X rapids
X X \
4 ? fast streams
4 Red algae’ streams
x Detritus fast streams
X X * rivers, lakes
X X madicolous
X
X X
A Me Green rivers etc.
filamentous
X algae
SOEX KE GCG rivers,lakes
Xx
x lakes etc.
> Game, G4 lakes etc.
XXXXXX ¥ —— lakes ete
OG x Detritus streamsetc
Chart II The phylogeny of the Hydroptilidae: generic relationships (I, Stactobiini; II, Leucotrichiini;
IW, Ochrotrichiini; IV, Neotrichiini; V, Hydroptilini; VI, Orthotrichiini).
THE GENERA OF THE HYDROPTILIDAE pips
Examination of adult features indicates that Padunia, especially with respect to the structure of
the thorax, the male genitalia and the shape and venation of the wings, is closely related to
Matrioptila Ross and Nepaloptila Kimmins, primitive members of the Protoptilinae (Glosso-
somatidae) from the SE. United States and SE. Asia respectively. Figs 149, 150 show the resem-
blance of the head and thorax of Padunia to those of Nepaloptila.
In 1954, Mosely transferred Mortoniella Ulmer, Protoptila Banks, Mexitrichia Mosely,
Canoptila Mosely and Antoptila Mosely, which are essentially Neotropical genera, from the Hy-
droptilidae to the Glossosomatinae, then a subfamily of the Rhyacophilidae. Subsequently, Ross
(1956) elevated the group, with the inclusion of Culoptila Mosely, to the subfamily Protoptilinae
within the Glossosomatidae, at the same time erecting the genus Matrioptila for a single primitive
species originally described in Protoptila.
Representatives of the Protoptilinae occur from southern Canada to central Chile and the
subfamily now contains the following additional genera: Campsiophora Flint, Cariboptila Flint,
Mastigoptila Schmid, Merionoptila Schmid and Tolhuaca Schmid. The description of Nepaloptila
Kimmins (1964) from a single Nepalese species furnished the first Palaearctic record of this sub-
family and this may now be supported by this discovery of the true identity of Padunia Martynov.
Ulmer (in litt. to Mosely, vide Mosely, 1954) suggested that Padunia might also belong with the
Protoptila-group but, possibly due to not having any material to study, Mosely did not transfer
it with the others. He did, however, remark upon its more primitive venation (and its resemblance
to Antoptila) which is now substantiated if this character is compared with those of Matrioptila
and Nepaloptila which are both considered to be the more primitive members of the Protoptilinae.
Ross (1956) considered the subfamily to have originated in the New World due to the occurrence
there of Matrioptila, but this theory may now need to be modified on account of the discovery of
the equally primitive genera Nepaloptila and Padunia in the Old World. The possibility that
Matrioptila and Padunia were derived from New World ancestors which passed into Asia, how-
ever, cannot be ruled out. The immature stages of Padunia are unknown.
Genus TSUKUSHITRICHIA Kobayashi (Psychomyiidae)
Tsukushitrichia Kobayashi, 1964 : 211. Type-species: Tsukushitrichia forficula Kobayashi, by monotypy.
DISTRIBUTION. Japan.
Kobayashi originally described this monotypic genus in the Hydroptilidae but stated that it was
unique within the family on account of its atypical spur formula (2.4.3), the structure of the fifth
abdominal segment and the wing venation. In addition ocelli are absent, the fifth segment of the
maxillary palp is long and flexible and the male genitalia are quite unlike those of any other
hydroptilid.
If the spur formula is considered to be derived from the combination 2.4.4, the species keys out
in Ross (1944) to the family Psychomyiidae, very close to the genus Psychomyia especially with
respect to the wing venation and the distinctive elongate fifth segment of the maxillary palp,
the latter always being proportionately shorter in the Hydroptilidae. Of the known species of
Japanese psychomyiids, Kibuneopsychomyia kibuneana Tsuda bears most resemblance to T.
forficula, both in the venation, which is almost identical, and the general aspect of the male geni-
talia. It is thus proposed that the genus Tsukushitrichia be transferred from the Hydroptilidae to
the Psychomyiidae where it may be synonymous with the genus Kibuneopsychomyia Tsuda.
Examination of material would be necessary to confirm these propositions. Neither the immature
stages of Tsukushitrichia nor Kibuneopsychomyia are known.
Genus PETROTRICHIA Ulmer
Petrotrichia Ulmer, 1910 : 43. Type-species: Petrotrichia palpalis Ulmer, by monotypy.
DISTRIBUTIONS. Seychelles.
This isolated monotypic genus is known only from the original series of males and females from
226 J. E. MARSHALL
150
prescutum
Figs 149-150 Glossosomatidae, Protoptilinae, adult head and thorax. 149, Nepaloptila;
150, Padunia.
the Seychelles, the immature stages being unknown. Examination of paratype material in the
BMNH indicates that this genus is definitely not a member of the Hydroptilidae, in which it was
originally placed, although difficulty has been encountered in assigning it to its proper family.
The adult features are as follows: head without ocelli or postoccipital warts; basal antennal
segment elongate; maxillary palpi (male and female) 5-segmented (cf. Ulmer, 1910); spur formula
originally stated to be 0.2.4 but a minute spur has been detected on the fore-tibia, hence 1.2.4;
mesothorax elongate (cf. Hydroptilidae), mesoscutellum ovoid, with a short distance between
posterior margins of mesoscutellum and notum; metascutellum elongate, subtriangular; wings
with rounded apices (slightly tapered); abdominal sternite V without lateral setate processes or
sternal ridge; 3 tenth segment fused, with a pair of parallel, tapering, ventro-lateral processes
(=fused inferior appendages), dorsal cerci each with a short median process and longer lateral
baso-ventral processes, aedeagus ‘fan-like’.
Following Scott’s (1967) key to South African caddis-flies, Petrotrichia keys out with Hydro-
salpinx Barnard and Petrothrincus Barnard which have been placed in the Beraeidae and Molan-
nidae respectively. The head and thorax of Petrotrichia accord well with those of Molanna
flavicornis as figured by Wiggins (1968 : 11, Fig. 12). Professor H. H. Ross (pers. comm.) is at
present studying the relationships of Hydrosalpinx and Petrothrincus which may prove, at least in
the case of the latter, to be primitive relict Trichoptera so far found only in the southern African
continent. Ross agrees (pers. comm.) that it is possible that Petrotrichia might be a beraeid or a
small molannid, especially with respect to the ‘weird dorsal processes (cerci)’ and the venation.
Discovery of the immature stages would greatly aid in establishing the relationships of these
genera.
Although such a conclusion is very unsatisfactory, it is proposed that Petrotrichia should be
removed from the Hydroptilidae, on the basis of the great differences in the structures of the head,
antennae, thorax, abdominal segment V and the male genitalia, and that it should be grouped,
for the time being, with Hydrosalpinx and Petrothrincus.
THE GENERA OF THE HYDROPTILIDAE 2a)
Checklist of species of Hydroptilidae
(* indicates material examined; + indicates fossil species)
Family HYDROPTILIDAE Stephens
Subfamily PTILOCOLEPINAE Martynov
Genus PALAEAGAPETUS Ulmer
Genus PTILOCOLEPUS Kolenati
*colchicus Martynov
*extensus McLachlan
*oranulatus (Pictet)
dilatatus Martynov
turbidus Kolenati
villosus Navas
*celsus Ross
guppyi Schmid
nearcticus Banks
trotundatus Ulmer
Subfamily HYDROPTILINAE Stephens
Tribe STACTOBIINI Botosaneanu
STACTOBIA-group
Genus STACTOBIA McLachlan
Afritrichia Mosely
Aratrichia Mosely
Lamonganotrichia Ulmer syn. n.
furcata-group
*algira Vaillant
*atra (Hagen)
*beatensis Mosely
*caspersi Ulmer
decosterai Jacquemart
*eatoniella McLachlan
oredonensis Mosely
eretziana Botosaneanu & Gasith
freyi Nybom
*furcata Mosely
fuscicornis Schneider sensu
McLachlan, 1884 (partim)
*fuscicornis (Schneider)
obscura (Kolenati)
jacquemarti Malicky
*kimminsi Schmid
*maculata Vaillant
*malacantosa Schmid
margalitana Botosaneanu
*mclachlani Kimmins
botosaneanui Schmid
delamerei Coineau & Jacquemart
fuscicornis Schneider sensu
McLachlan, 1884 (partim)
monnioti Jacquemart
*moselyi Kimmins
eatoniella McLachlan sensu
Mosely
fuscicornis Schneider sensu
McLachlan
nybomi Schmid
atra (Ragen) sensu Morton
atra (Hagen) sensu Nybom
atra (Hagen) sensu Schmid
storai Nybom
martynovi-group
*dohleri Schmid
fischeri Schmid
*forsslundi Schmid
klapaleki Schmid
marlieri Schmid
*martynovi Schmid
*mayeri Schmid
olgae Martynov
*quadrispina Kimmins
*schmidi Kimmins
*tjederi Schmid
ulmeriana Schmid
urania Malicky
nielseni-group
bolzei Jacquemart
*crassa (Ulmer) comb. n.
*fahjia (Mosely)
moretti Schmid
*nielseni Schmid
radavanovici Schmid
risiana Schmid
vaillanti-group
*aurea (Mosely)
* vaillanti Schmid
Incertae sedis
japonica Iwata
mallorcensis Vaillant (nomen nudum)
megalatlantica Vaillant (nomen nudum)
Genus PLETHUS Hagen
Plethotrichia Ulmer syn. n.
*acutus Ulmer
amogawarsa Schmid
*baliana (Ulmer) comb. n.
*bodikatuwa Schmid
cilamegha Schmid
cruciatus Ulmer
*cursitans (Hagen)
kala Schmid
228
usawasadenna Schmid
*vajhrobodhi Schmid
STACTOBIELLA-group
Genus STACTOBIELLA Martynov
Tascobia Ross
biramosa-group
biramosa Martynov
*palmata (Ross)
ulmeri-group
delira (Ross)
*ulmeri (Siltala)
risi (Felber)
brustia-group
brustia (Ross)
Genus BREDINIA Flint
dominicensis Flint
Genus CHRYSOTRICHIA Schmid
*aranuwa Schmid
badhami Schmid
*dotalugola Schmid
hapitigola Schmid
*hatnagola Schmid
J. E. MARSHALL
Genus PARASTACTOBIA Schmid
talakalahena Schmid
Genus CATOXYETHIRA Ulmer
Sperotrichia Marlier syn. n.
*fasciata Ulmer
formosae (Iwata)
*improcera Statzner
mali (Marlier) comb. n.
*ocellata Statzner
*pinheyi Kimmins
veruta Morse
MADIOXYETHIR4-group
Genus MADIOXYETHIRA Schmid
*marshalli Statzner
*milinda Schmid
*nepalensis Kimmins
trifurcata (Jacquemart) comb. n.
Genus PSEUDOXYETHIRA Schmid
asgiriskanda Schmid
Genus SCELOTRICHIA Ulmer
*saranganica Ulmer
Tribe LEUCOTRICHIINI Flint
LEUCOTRICHIA4-group
Genus LEUCOTRICHIA Mosely
melleopicta-group
chiriquiensis Flint
gomezi Flint
*limpia Ross
*melleopicta Mosely
tubifex Flint
viridis Flint
pictipes-group
fairchildi Fiint
imitator Flint
pictipes (Banks)
sarita Ross
Genus ZUMATRICHIA Mosely
multisetosa-group
multisetosa Flint
galtena-group
antilliensis Flint
attenuata Flint
bifida Flint
chiriquiensis Flint
diamphidia Flint
echinata Flint
*galtena Mosely
notosa (Ross)
saluda Flint
strobilina Flint
vieja Flint
filosa-group
caudifera Flint
*filosa Mosely
rhampoides Flint
teapa Flint
palmara-group
anomaloptera Flint
palmara Flint
Genus PELTOPSYCHE Miller
maclachlani Miller
*sieboldii Miiller
Genus ANCHITRICHIA Flint
spangleri Flint
Genus COSTATRICHIA Mosely
simplex-group
simplex Flint
spinifera Flint
lodora-group
bipartita Flint
*lodora Mosely
panamensis Flint
tripartita Flint
Genus ACOSTATRICHIA Mosely
brevipenis Flint
fimbriata Flint
*plaumanni Mosely
*simulans Mosely
spinifera Flint
Genus BETRICHIA Mosely
argentinica Flint
bispinosa Flint
occidentalis Flint
surinamensis Flint
*zilbra Mosely
Genus ABTRICHIA Mosely
*antennata Mosely
*squamosa Mosely
Genus CELAENOTRICHIA Mosely
*edwardsi Mosely
THE GENERA OF THE HYDROPTILIDAE
ALISOTRICHIA-group
Genus ALISOTRICHIA Flint
argentilinea Flint
blantoni Flint
chorra Flint
dominicensis Flint
hirudopsis Flint
lobata Flint
orophila Flint
quemada Flint
septempunctata Flint
tamaza Flint
trifida Flint
wirthi Flint
Tribe OCHROTRICHIINI trib. n.
Genus OCHROTRICHIA Mosely
Polytrichia Sibley
Subgenus OCHROTRICHIA
*aldama (Mosely)
alexanderi Denning & Blickle
alsea Denning & Blickle
*anisca (Ross)
argentea Flint & Blickle
arizonica Denning & Blickle
*arranca (Mosely)
*arva (Ross)
attenuata Flint
brayi Flint
buccata Denning & Blickle
caimita Flint
caligula Flint
capitana Ross
chiapa Denning & Blickle
confusa (Morton)
*contorta (Ross)
cruces Flint
dactylophora Flint
denningi Blickle & Morse
eliaga (Ross)
escoba Flint
felipe Ross
filiforma Flint
flagellata Flint
gurneyi Flint
hadria Denning & Blickle
ildria Denning & Blickle
*insularis Mosely
intermedia Flint
lobifera Flint
*logana (Ross)
*lometa (Ross)
lucia Denning & Blickle
marica Flint
*mono (Ross)
moselyi Flint
nacora Denning & Blickle
okanogensis Flint
oregona (Ross)
pacifica Flint
palitla Flint
panamensis Flint
pectinata Flint
pectinifera Flint
phenosa Ross
ponta Flint
potomus Denning
provosti Blickle
quadrispina Denning & Blickle
riesi Ross
rothi Denning & Blickle
shawnee (Ross)
spinosa (Ross)
spinosissima Flint
spinulata Denning & Blickle
*stylata (Ross)
susanae Flint & Herrmann
tagala Flint
tarsalis (Hagen)
*tenanga (Mosely)
*trapoiza Ross
*unio (Ross)
verda Flint
vertreesi Denning & Blickle
weddleae Ross
wojcickyi Blickle
xena (Ross)
zioni Denning & Blickle
Subgenus METRICHIA Ross
Argentitrichia Jacquemart syn. n.
aberrans Flint
argentinica Schmid
arizonensis Flint
biungulata Flint
bulbosa (Jacquemart) comb. n.
campana Flint
continentalis Flint
dietzi Flint
exclamationis Flint
juana Flint
lemniscata Flint
neotropicalis Schmid
nigritta Banks
penicillata Flint
quadrata Flint
similis Flint
trigonella Flint
warema Flint
yalla Flint
Genus RHYACOPSYCHE Miller
hagenii Miiller
mexicana (Flint)
obliqua Flint
torulosa Flint
turrialbe Flint
229
230
J. E. MARSHALL
Tribe NEOTRICHIINI Ross
Genus NEOTRICHIA Morton
Cyllene Chambers
Exitrichia Mosely
Dolotrichia Mosely
Guerrotrichia Mosely
Lorotrichia Mosely
collata-group
alata Flint
*anahua (Mosely)
collata Morton
*digitata (Mosely)
elerobi Blickle
*eroga (Mosely)
ersitis Denning
*esmalda (Mosely)
*exicoma (Mosely)
falca Ross
heleios Flint
interrupta Flint
iridescens Flint
minutisimella (Chambers)
*notuena (Mosely)
*novara (Mosely)
numii Ross
okopa Ross
*olorina (Mosely)
osmena Ross
*ovona (Mosely)
*oxima (Mosely)
panneus Denning
riegeli Ross
sonora Ross
*tertia (Mosely)
vibrans Ross
ranea Denning
canixa-group
*canixa (Mosely)
corniculans Flint
*dubitans (Mosely)
*xicana (Mosely)
caxima-group
*cameria (Mosely)
*caxima (Mosely)
costaricensis Flint
edalis Ross
rotundata Flint
hiaspa-group
halia Denning
*hiaspa (Mosely)
kitae Ross
biuncifera-group
biuncifera Flint
bullata Flint
falcifera Flint
proboscidea Flint
unispina Flint
bifida-group
bifida Flint
lobata Flint
Genus MAYATRICHIA Mosely
acuna Ross
*ayama Mosely
ponta Ross
*rualda Mosely
Tribe HYDROPTILINI Stephens
AGRA YLEA-group
Genus AGRAYLEA Curtis
argyricola Kolenati
*cognatella McLachlan
costello Ross
drosima Navas
insularis (Hagen)
*multipunctata Curtis
multiguttata Uljanin
signata Banks
flavida (Banks)
fraterna Banks
saltesea Ross
*sexmaculata Curtis
pallidula McLachlan
flabellifera (Bremi) (partim)
tspathifera Ulmer
Genus ALLOTRICHIA McLachlan
tampullata Ulmer
heterocera Navas
laerma Malicky
marinkoviciae Malicky
*pallicornis (Eaton)
succinica Hagen
tauri Jacquemart
teldanica Botosaneanu
vilnensis Raciecka
Genus MICROPTILA Ris
apsara Schmid
*bejela Mosely
*indra Schmid
*minutissima Ris
Genus UGANDATRICHIA Mosely gen. rev.
Moselyella Kimmins
nigra-group
*acuta Mosely
africana (Marlier & Vaillant) comb. n.
*minor Mosely
*nigra Mosely
rhodesiensis Scott
violacea-group
*cyanotrichia (Kimmins)
THE GENERA OF THE HYDROPTILIDAE 231
*nikataruwa (Schmid)
roudra (Schmid) comb. n.
*sourya (Schmid) comb. n.
*violacea (Morton)
Genus DHATRICHIA Mosely
*bipunctata Statzner
*inasa Mosely
HYDROPTIL4A-group
Genus HYDROPTILA Dalman
Phrixocoma Eaton
Hydropneuma Enderlein
Hydroptilina Martynov
Oxydroptila Martynov syn. n.
Oeceotrichia Ulmer syn. n.
Pasirotrichia Ulmer syn. n.
Sumatranotrichia Ulmer syn. n.
sparsa-group
*acuta Mosely
*africana Kimmins
*angulata Mosely
*campanulata Morton
cintrana Morton
*cornuta Mosely
emarginata Martynov
friedeli Malicky
*fuentaldeala Schmid
*hochyangha Schmid
*lotensis Mosely
*simulans Mosely
*sparsa Curtis
uncinata-group
angulifera Kumanski
*fortunata Morton
juba (Enderlein)
insubrica Ris
kalonichtis Malicky
bureschi Kumanski
*stellifera Morton
*sylvestris Morton
*uncinata Morton
vichtaspa Schmid
capensis-group
calundoensis Marlier
*capensis Barnard
consimilis-group
*ajax Ross
albicornis Hagen
maculata (Banks)
transversa Banks
ancistrion Flint
*angusta Ross
antilliarum Flint
arctia Ross
berneri Ross
broweri Blickle
*consimilis Morton
denza Ross
ditalea Flint
grenadensis Flint
*icona Mosely
latosa Ross
lloganae Blickle
martorelli Flint
medinai Flint
*melia Ross
*meralda Mosely
*mexicana Mosely
novicola Blickle & Morse
*paschia Mosely
*pecos Ross
*perdita Morton
*producta Mosely
pullatus Denning
quinola Ross
scolops Ross
strepha Ross
surinamensis Flint
tusculum Ross
valhalla Denning
veracruzensis Flint
occulta-group
*adana Mosely
*armata Ross
*cognata Mosely
*cruciata Ulmer
*gandhara Schmid
*hirra Mosely
*martini Marshall
occulta (Eaton) sensu auctt.
*occulta (Eaton)
insignis Martynov
*kimminsi Mosely
*parthava Schmid
palestinae Botosaneanu & Gasith
panchaoi Schmid
phaon Malicky
rhodica Jacquemart
kumanskii Malicky
*sanghala Schmid
*spatulata Morton
taurica Martynov
*vala Ross
*valesiaca Schmid
waskesia Ross
tineoides-group
amoena Ross
*ampoda Ross
callia Denning
fiskei Blickle
*hamata Morton
*perplexa Mosely
lennoxi Blickle
metoeca Blickle & Morse
*modica Mosely
moselyi Ulmer
remita Blickle & Morse
*rono Ross
spinata Blickle & Morse
232
*tineoides Dalman
femoralis (Eaton)
longispina McLachlan
tortosa Ross
wyomyia Denning
dikirilagoda-group
dikirilagoda Schmid
*kurukepitiya Schmid
mitirigalla Schmid
losida-group
*bispina Kimmins
*incertula Mosely
*losida Mosely
*tasmanica Mosely
*triloba Kimmins
pulchricornis-group
aegyptia Ulmer
*dampfi Ulmer
kurnas Malicky
phenianica Botosaneanu
*pulchricornis Pictet
forcipata-group
*bifurcata Mosely
*forcipata (Eaton)
ivisa Malicky
vectis-group
*vectis Curtis
corsicanus Mosely
maclachlani Klapalek
viganoi Botosaneanu
tigurina-group
*cortensis Mosely
tigurina Ris sensu Mosely
rheni Ris
*tigurina Ris
waubesiana-group
*acadia Ross
*delineata Morton
nicoli Ross
salmo Ross
wakulla Denning
*waubesiana Betten
xoncla Ross
Incertae sedis
acoma Denning
angustipennis (Martynov)
annulicornis Matsumura
*argosa Ross
*armathai Schmid
brincki Jacquemart
crenata (Ulmer) comb. n.
decia Etnier & Way
dentata Ross
dodgei Denning
elongata (Ulmer) comb. n.
eramosa Harper
Suentelarbola Schmid
furcata (Martynov) comb. n.
grandiosa Ross
J. E. MARSHALL
gunda Milne
jackmanni Blickle
*kirilawela (Schmid) comb. n.
lonchera Blickle & Morse
molsonae Blickle
pectinifera Schmid
tphileos Cockerell
*protera Ross
sengavi Schmid
*serrata Morton
simplex Nielsen (doubtful sp. acc.
Nielsen, in /itt.)
*touroumaya Schmid
trilobata Jacquemart
trullata (Ulmer) comb. n.
usurigonis Matsumura
*virgata Ross
xella Ross
xera Ross
OXYETHIRA-group
Genus OXYETHIRA Eaton
Lagenopsyche Miiller
Argyrobothrus Barnard
Loxotrichia Mosely
Dampfitrichia Mosely
Oxytrichia Mosely
falcata-group
boreella Svensson & Tjeder
delcourti Jacquemart
dentata Nybom
bidentata Nybom
*falcata Morton
rhodani Schmid
fischeri Higler
*frici Klapalek
*spinosella McLachlan
flavicornis-group
*bogambara Schmid
campanula Botosaneanu
ecornuta Morton
*flavicornis (Pictet)
costalis (Curtis) sensu Eaton
sida Blickle & Morse
*tenuella Martynov
*yunidentata McLachlan
fuentejalona Schmid
meridionalis Jacquemart & Coineau
rivicola-group
allagashensis Blickle
dualis Morton
*forcipata Mosely
grisea Betten
lumosa Ross
*michiganensis Mosely
novasota Ross
obscura Flint
obtatus Denning
rivicola Blickle & Morse
rossi Blickle & Morse
THE GENERA OF THE HYDROPTILIDAE 233
setosa Denning
sodalis Ross & Spencer
unispina Flint
simplex-group
galekoluma Schmid
*harpagella Kimmins
paramartha Schmid
ramosa Martynov
angustella Martynov
*simplex Ris
*tristella Klapalek
distinctella-group
araya Ross
*distinctella McLachlan
*sagittifera Ris
*serrata Ross
azteca-group
archaica Malicky
*azteca (Mosely)
*dalmeria (Mosely)
glasa (Ross)
janella Denning
neglecta Flint
puertoricensis Flint
*zilaba (Mosely)
ulmeri-group
aculea Ross
arizona Ross
cirrifera Flint
tega Flint
*ulmeri (Mosely)
zeronia-group
jamaicensis Flint
longissima Flint
macrosterna Flint
simulatrix Flint
verna Ross
walteri Denning
zeronia Ross
pallida-group
florida Denning
*maya Denning
pallida (Banks)
cibola Denning
viminalis Morton
bidentata-group
abacatia Denning
aeola Ross
*bidentata Mosely
mirabilis-group
flagellata Jacquemart
*mirabilis Morton
Incertae sedis
*albiceps McLachlan
anabola Blickle
angustella Martynov
berneri Etnier & Way
coercens Morton
hyalina (Miller)
incana Ulmer
spirogyrae (Miiller)
*velocipes (Barnard)
Genus STENOXYETHIRA Kimmins
Gnathotrichia Ulmer syn. n.
*excisa Kimmins
*isabellina (Ulmer) syn. n.
*minima Kimmins
Genus PAROXYETHIRA Mosely
*eatoni Mosely
*hendersoni Mosely
hintoni Leader
kimminsi Leader
*tillyardi Mosely
Genus XUTHOTRICHIA Mosely
*eskensis Mosely
*fimbriata Mosely
*ochracea Mosely
*simplex Mosely
Genus TRICHOLEIOCHITON Kloet &
Hincks
Leiochiton Guinard
Synagotrichia Ulmer syn. n.
*fagesii (Guinard)
felina (Ris)
flabellifera (Bremi) partim.
*fortensis (Ulmez) comb. n.
*Jacustris Kimmins
Tribe ORTHOTRICHIINI Nielsen
Genus ORTHOTRICHIA Eaton
Clymene Chambers
Javanotrichia Ulmer syn. n.
Orthotrichiella Ulmer syn. n.
Baliotrichia Ulmer syn. n.
angustella-group
*angustella (McLachlan)
brunneicornis (Pictet) (partim)
*avicularis Kimmins
barnardi Scott
benguelensis Marlier
damasi Marlier
dampfi (Ulmer) comb. n.
flagellum Marlier
indica Martynov
kivuensis Jacquemart
melitta Malicky
*moselyi Tjeder
*sanya Mosely
*spinicauda Kimmins
straelini Jacquemart
verbeki Jacquemart
234
litoralis-group
*curvata (Ulmer) comb. n.
guruluhela (Schmid) comb. n.
hinipitigola (Schmid) comb. n.
*litoralis (Ulmer) comb. n.
* maeandrica (Ulmer) comb. n.
medipitigola (Schmid) comb. n.
*ranauana (Ulmer) comb. n.
udawarama (Schmid) comb. n.
costalis-group
*aequatoriana Kimmins
alboguttata Jacquemart
*costalis (Curtis)
tetensii Kolbe
extensa Martynov
aegerfasciella-group
*aegerfasciella (Chambers)
*americana Banks
dorsalis Banks
brachiata Morton
Genus MACROSTACTOBIA Schmid
*elawalikanda Schmid
Genus DIBUSA Ross
angata Ross
Genus NOTHOTRICHIA Flint
illiesi Flint
Genus DICAMINUS Miiller
J. E. MARSHALL
baldufi Kingsolver & Ross
*cristata Morton
americana Betten
curta Kingsolver & Ross
instabilis Denning
*tragetti Mosely
kokodana-group
*kokodana Kimmins
*obscura Kimmins
Incertae sedis
dentata Kingsolver & Ross
petiti Jacquemart
trilineata Jacquemart
Genus ITHYTRICHIA Eaton
Saranganotrichia Ulmer syn. n.
bosniaca Botosaneanu
*clavata Morton
*decussata (Ulmer) comb. n.
*Jamellaris Eaton
brunneicornis (Pictet) (partim)
INCERTAE SEDIS
Genus ORPHNINOTRICHIA Mosely
*maculata Mosely
Genus CALEDONOTRICHIA Sykora
*illiesi Sykora
minor Sykora
Genus ELECTROTRICHIA Ulmer
tsubtilis Ulmer
Diaulus Miller
ladislavii (Miller)
Acknowledgements
Most of the research for this paper was carried out during the tenure of a Postgraduate Student-
ship grant awarded by the Trustees of the British Museum (Natural History) to whom thanks are
due for the use of their facilities and access to their collections. ,
I am grateful to my two supervisors, Mr R. G. Davies (Department of Zoology and Applied
Entomology, Imperial College, London) and Dr P. E. S. Whalley (Department of Entomology,
BMNH), for their continued guidance and assistance during the course of my studies.
Thanks are due to everyone who provided me with additional material, records, information,
advice or any other form of assistance. They are too numerous to list in full here but the more
notable are: Dr L. Botosaneanu, Bucharest, Rumania; Dr O. S. Flint Jr, Washington, D.C.,
U.S.A.; Dr L. W. G. Higler, Leersum, The Netherlands; Dr H. Malicky, Lunz-am-See, Austria;
Dr O. Nybom, Imatra, Finland; Professor H. H. Ross, Athens, Georgia, U.S.A.; Dr B. Statzner,
Kiel, West Germany; Dr H. Weidner, Hamburg, West Germany; Dr G. B. Wiggins, Ontario,
Canada and Dr L. Zhiltzova, Leningrad, U.S.S.R.
Finally, I am especially grateful to my colleagues at the BMNH for a variety of reasons, again
too numerous to list in full, but of whom I particularly thank Mr W. G. Tremewan and Dr P. C.
Barnard.
References
Agassiz, J. L. R. 1846. Nomenclator zoologicus. Index Universalis: 32.
Barker J. 1867. Larval state of a dipterous insect. Q. J/ microsc. Sci. (N.S.) 7 : 175-176.
THE GENERA OF THE HYDROPTILIDAE 235
Barnard, K. H. 1934. South African caddis-flies (Trichoptera). Trans. R. soc. S. Afr. 21 : 291-394.
Barnard, P. C. 1971. The larva of Agraylea sexmaculata Curtis (Trichoptera, Hydroptilidae). Entomolo-
gist’s Gaz. 22 : 253-257.
Botosaneanu, L. 1956. Recherches sur les Trichoptéres de Bulgarie recuellis par MM. le Prof. A. Balkanov
et B. Rusev (Trichoptera). Beitr. Ent. 6: 354-402.
— 1967. Trichoptera. Jn Illies, J. (Ed.), Limnofauna Europaea. Pp. 285-309. Stuttgart.
& Sykora, J. 1963. Nouvelle contribution ala connaissance des Trichoptéres de Bulgarie. Sb. faun.
Praci ent. Odd. nar. Mus. Praze 9 : 121-142.
Burton, G. J. & McRae, T. M. 1972. Observations on trichopteran predators on aquatic stages of Simulium
damnosum and other Simulium spp. in Ghana. J. med. Ent. Honolulu 9 : 289-294.
Chambers, V. T. 1873. Micro-Lepidoptera. Can. Ent. 5 : 110-115, 124-128, 147-152.
Curtis, J. 1834. Descriptions of some hitherto nondescript British species of mayflies of anglers. Phil.
Mag. (3) 4 : 212-218.
Dalman, J. W. 1819. Nagra nya insekt-genera, beskrifna. K. svenska Vetensk Akad. Handl. 40 : 117-127.
Danecker, E. 1961. Studien zur hygropetrischen Fauna. Biologie und Okologie von Stactobia und Tinodes
(Insecta, Trichoptera). Int. Revue ges. Hydrobiol. 46 : 214-254.
Denning, D. G. & Blickle, R. L. 1972. Review of Ochrotrichia (Trichoptera : Hydroptilidae). Ann. ent. Soc.
Am. 65 : 141-151.
Disney, R. H. L. 1973. Larval Hydroptilidae (Trichoptera) that prey upon Simuliidae in Cameroon.
Entomologist’s mon. Mag. 108 (1972) : 84-85.
Dohler, W. 1963. Liste der deutschen Trichopteren. NachrBl. bayer. Ent. 12 : 17-22.
Eaton, A. E. 1873. On the Hydroptilidae, a family of the Trichoptera. Trans. ent. Soc. Lond. 1873 : 125-
150.
Edwards, S. W. Arnold, & C. R. 1961. Thecaddis flies of the San Marcos River. Texas J. Sci. 13 : 398-415.
Enderlein, G. von 1929. Entomologia Canaria. 2. Zool. Anz. 84 : 221-234.
Fahy, E. 1972. The larva of Hydroptila forcipata (Eaton) (Trichoptera, Hydroptilidae). Entomologist’s
mon. Mag. 107 (1971) : 145-148.
Fischer, F. C. J. 1961. Trichopterorum Catalogus 2. Philopotamidae, Hydroptilidae, Stenopsychidae.
iv+ 190 pp. Amsterdam.
1971. Trichopterorum Catalogus 12. Supplement to 1 & 2. (Hydroptilidae: 73-177). Amsterdam.
Flint, O. S. 1962. The immature stages of Palaeagapetus celsus Ross (Trichoptera, Hydroptilidae). Bull.
Brooklyn ent. Soc. 57 : 40-44.
1964. The caddisflies (Trichoptera) of Puerto Rico. Tech. Pap. agric. Exp. Stn P. Rico 40 : 1-80.
—— 1966. On the identity of Clymene aegerfasciella Chambers (Trichoptera : Hydroptilidae). Proc. ent.
Soc. Wash. 68 : 135.
—— 1967a. Studies of Neotropical caddisflies, IV: new species from Mexico and Central America
(Trichoptera). Proc. U.S. natn. Mus. 123 : 1-24.
—— 1967b. Studies of Neotropical caddis flies, II: Trichoptera collected by Prof. Dr. J. Illies in the Chilean
subregion. Beitr. neotrop. Fauna 5 : 45-68.
—— 1968a. New species of Trichoptera from the Antilles. Fla Ent. 51 : 151-153.
—— 1968b. The caddisflies of Jamaica (Trichoptera). Bull. Inst. Jamaica Sci. Ser. 19 : 1-68.
—— 1968c. Bredin—Archbold-Smithsonian Biological Survey of Dominica. 9. The Trichoptera (caddis
flies) of the Lesser Antilles. Proc. U.S. natn. Mus. 125 : 1-86.
—— 1970. Studies of Neotropical caddisflies, X: Leucotrichia and related genera from North and Central
America (Trichoptera, Hydroptilidae). Smithson. Contr. Zool. 60 : 1-64.
— 1971. Studies of Neotropical caddisflies, XI: the genus Rhyacopsyche in Central America (Hydrop-
tilidae). Proc. biol. Soc. Wash. 83 : 515-526.
— 1972a. Studies of Neotropical caddisflies, XIII: the genus Ochrotrichia from Mexico and Central
America (Trichoptera : Hydroptilidae). Smithson. Contr. Zool. 118 : 1-28.
—1972b. Studies of Neotropical caddisflies, XIV: on a collection from northern Argentina. Proc. biol.
Soc. Wash. 85 : 223-248.
— 1974. The Trichoptera of Surinam. Studies of Neotropical caddisflies, XV. Uitg. natuurw. StudKring
Suriname 14 (55) : 1-151.
Fredeen, F. J. H. 1972. The temporary abatement of some nuisance species of Trichoptera with D.D.D.
(T.D.E.) larvicide. Can. Ent. 104 : 145-163.
Giudicelli, J. & Vaillant, F. 1967. Lalarve et la nymphe d’Allotrichia pallicornis (Eaton) (Trichoptera).
Trav. Lab. Hydrobiol. Piscic. Univ. Grenoble 57-58 (1965-66) : 29-36.
Guinard, E. 1879. Métamorphoses d’un genre nouveau de Phryganide (Leiochiton fagesii). Mém Acad.
Sci. Lett. Montpellier. Section des Sciences 9 : 139-144.
236 J. E. MARSHALL
Hagen, H. A. 1855. Versuch die Phryganiden Pictet’s zu bestimmen. Stettin. ent. Ztg 16 : 204-210.
— 1864. Ueber Phryganiden-Gehause. Stettin. ent. Ztg 25 : 221-266.
— 1887. Ueber Plethus cursitans. Verh. zool.-bot. Ges. Wien 37 : 643-645.
Hanna, H. M. 1961. The larva of Hydroptila sparsa Curtis (Trichoptera : Hydroptilidae). Entomologist’s
Gaz. 12 : 69-75.
Hickin, N. E. 1967. Caddis larvae. 476 pp., 980 figs. London.
Higler, L. W. G. 1974. Oxyethira fischeri n. sp. A new Oxyethira species from Madeira. Ent. Ber., Amst.
34 : 62-63.
Hinton, H. E. 1963. The ventral ecdysial lines of the head of endopterygote larvae. Trans. R. ent. Soc. Lond.
115 : 39-61.
Hudson, G. V. 1886. On the metamorphosis of the caddis fly. Trans. N.Z. Inst. 18 : 213-214.
Hynes, H. B. N. 1970. The ecology of running waters. xxiv+555 pp. Liverpool.
Iwata, M. 1928. Five new species of trichopterous larvae from Formosa. Annotnes zool. jap. 11 : 341-343.
Jacquemart, S. 1962a. Trois Trichoptéres nouveaux d’Afrique Centrale. Bull. Inst. r. Sci. nat. Belg. 38
(34) : 1-11.
— 19625. La larve d’Orthotrichia angustella Mcl. (Trichoptére. Hydroptilidae). Bull. Inst. r. Sci. nat. Belg.
38 (15) : 1-8.
— 1963. Deux Trichoptéres nouveaux d’Argentine. In Biologie de Il’ Amerique Australe. II, Etudes sur la
faune du sol : 339-342.
— 1965. Résultats de l’expedition belge au Moyen-Orient. (Premiére note). Sept Trichopteres Hydrop-
tilides nouveaux de Turquie et d’Iran. Bull. Inst. r. Sci. nat. Belg. 41 (33) : 1-19.
— 1973. Descriptions des deux Trichoptéres Hydroptilides nouveaux et de l’imago de Stactobia monnioti
Jacquemart (Ile de Rhodes). Bull. Inst. r. Sci. nat. Belg. (Ent.) 49 (4) : 1-16.
— & Coineau, Y. 1962. Missions S. Jacquemart dans les Pyrénées Orientales (2e note). Les Trichoptéres
Hydroptilides des Albéres. Bull. Inst. r. Sci. nat. Belg. 38 (24) : 1-181.
Kimmins, D. E. 1951. Indian caddis flies — IV. New genera and species of the family Hydroptilidae. Ann.
Mag. nat. Hist. (12) 4 : 193-213.
— 1958. The British species of the genus Oxyethira. Entomologist’s Gaz. 9 : 7-17.
— 1964. On the Trichoptera of Nepal. Bull. Br. Mus. nat. Hist. (Ent.) 15 : 35-55.
— 1966. A revised checklist of the British Trichoptera. Entomologist’s Gaz. 17 : 111-120.
Kloet, G. S. & Hincks, W. D. 1944. Nomenclatural notes on two generic names in the Trichoptera. Ento-
mologist 77 : 97.
Kobayashi, T. 1964. A new genus and species of Hydroptilidae from Japan (Trichoptera). Kontyé 32 : 211-
213:
Kolenati, F. 1848. Genera et species Trichopterorum. Pars prior. Heteropalpoidea. vi+ 108 pp., 3 pls.
Prague. [First published as a separate, subsequently reprinted in 1851 in Abh. K. bohm. Ges. Wiss. (5)
6 with same pagination.]
Leader, J. P. 1970. Hairs of the Hydroptilidae (Trichoptera). Tane 16 : 121-129.
— 1972. New Zealand Hydroptilidae. J. Ent. (B) 41 : 191-200.
Lepneva, S. G. 1932. Zum Studium der Trichopterenlarven in den Wasserbecken der Systeme des Dnipro
und des Sud-Bugs. Zh. bio-zool. Tsyklu Kyyir 3 : 71-115.
— 1964. Larvae and pupae of the Suborder Annulipalpia Trichoptera 2 (1). [In Russian.] Fauna SSSR
(N.S.) 88 : 1-566, 773 figs.
Lloyd, J. T. 1915. Notes on Jthytrichia confusa Morton. Can. Ent. 47 : 117-121.
Macdonald, W. W. 1950. The larvae of Mystacides azurea L., Cyrnus flavidus McLachlan and Oxyethira
simplex Ris (Trichoptera). Proc. R. ent. Soc. Lond. (A) 25 : 19-28.
McLachlan, R. 1880. 4 monographic revision and synopsis of the Trichoptera of the European fauna. (Part
IX with supplement of part II.) Pp. 501-523, xiii-Ixxix. London.
— 1884. Ibid. (1st additional supplement). 76 pp. London.
Malicky, H. 1975. Fuenfzehn neue mediterrane K6cherfliegen. Mitt. ent. Ges. Basel (N.F.) 25 : 81-96.
Marlier, G. 1978. Sur une collection de Trichoptéres de |’Afrique occidentale. Revue Zool. afr. 92 : 283-
301.
Marshall, J. E. 1977. Hydroptila martini sp. n. and Hydroptila valesiaca Schmid (Trichoptera : Hydrop-
tilidae) new to the British Isles. Entomologist’s Gaz. 28 : 115-122.
— 1978. Trichoptera : Hydroptilidae. Handbk Ident. Br. Insects. 1 (14a) : 31 pp.
Martynov, A. V. 1910. Les Trichoptéres de la Sibérie et des régions adjacentes, ii-partie. [In Russian.]
Ezheg. zool. Muz. 15 : 351-429.
— 1913a. Zur Kenntnis der Trichopteren-fauna des Kaukasus. [In Russian.] Rab. Lab. zool. Kab. imp.
varsh. Univ. 1913 : 1-111.
THE GENERA OF THE HYDROPTILIDAE 237
— 19135. Contributions a la faune des Trichoptéres du Caucase. Trichoptéres de la province de Batoum
et environs du Novyj Afon. Trudy russk.. ent. Obosch. 40 : 1-30.
— 1924. Trichoptera. Jn Boghdanov-Katkov, N. N., Practical Entomology, 5. \xvii+ 388 pp. Leningrad.
— 1934. Tableaux analytiques de la faune de l’U.R.S.S. Trichoptera Annulipalpia. I. Opred. Faune SSSR
13 : 343 pp.
— 1935. Ona collection of Trichoptera from the Indian Museum. Rec. Indian Mus. 37 : 93-209.
Milne, M. J. & Milne, L. S. 1939. Evolutionary trends in caddis worm case construction. Ann. ent. Soc.
Am. 32 : 533-542.
Miller, A. 1921. Fritz Miiller. Werke, Briefe und Leben. 2. Briefe. 667 pp., 239 figs. Jena.
Morse, J. 1974. New caddis flies (Trichoptera) from Southern Africa. J. Kans. ent. Soc. 47 : 328-344.
Morton, K. J. 1887. On the cases, &c., of Oxyethira costalis, Curt., and another of the Hydroptilidae.
Entomologist’s mon. Mag. 23 : 201-203.
— 1905. North American Hydroptilidae. Bull. N.Y. St. Mus. 86 : 63-75.
Mosely, M. E. 1924. New Zealand Hydroptilidae. (Order Trichoptera.) Trans. N.Z. Inst. 55 : 670-673.
— 1932. Notes. Entomologist 65 : 176.
— 1933. The genus Stactobia McL. (Trichoptera). Sty/ops 2 : 162-165.
— 1934. New exotic Hydroptilidae. Trans. R. ent. Soc. Lond. 82 : 137-163.
— 1937. Mexican Hydroptilidae (Trichoptera). Trans. R. ent. Soc. Lond. 86 : 151-190.
— 1939a. The British caddisflies (Trichoptera): a collector’s handbook. 320 pp. London.
— 1939b. Trichoptera. Ruwenzori exped. 1934-35. 3 : 1-39.
— 1939c. The Brazilian Hydroptilidae (Trichoptera). Novit. zool. 41 : 217-239.
— 1954. The Protoptila group of the Glossosomatinae. (Trichoptera : Rhyacophilidae). Bull. Br. Mus.
nat. Hist. (Ent.) 3 : 315-346.
— & Kimmins, D. E. 1953. The Trichoptera of Australia and New Zealand. 550 pp. London.
Miiller, F. 1879a. Ueber Phryganiden. Zool. Anz. 2 : 38-40.
— 1879. Extracts from letters of F. Miiller. Proc. ent. Soc. Lond. 1879 : viii.
— 1879c. Notes on the cases of some South Brazilian Trichoptera. Trans. ent. Soc. Lond. 1879 : 131-144.
— 1880. Sobre las cases construidas palas larvas de insectos Trichopteros da Provincia de Santa Catharina.
Archos. Mus. nac. Rio de J. 3 : 99-134, 209-214.
Navas, L. 1916. Trichoptera de Aragan. Revta Acad. Cienc. exact. fis. quim. nat. Zaragoza 1 : 73-85.
— 1917. Trichopteros nuevos de Espana III. Broteria (Ser. Zool.) 15 : 5-17.
Neboiss, A. 1963. The Trichoptera types of species described by J. Curtis. Beitr. Ent. 13 : 582-635.
Needham, J. G. 1902. A probable new type of hypermetamorphosis. Psyche, Camb. 9 : 375-378.
Nielsen, A. 1948. Postembryonic development and biology of the Hydroptilidae. Biol. Skr. 5 (1) : 1-200,
3 pls, 64 figs.
— 1970. Trichoptera. Jn Tuxen, S. L. (Ed.), Taxonomist’s glossary of genitalia in insects. 2nd edn. Pp.
104-115. Munksgaard, Copenhagen.
Nybom, O. 1948. The Trichoptera of the Atlantic Islands. Commentat. biol. 8 (14) : 1-19.
— 1960. List of Finnish Trichoptera. Fauna fenn. 6 : 1-56.
Pendergrast, J. G. & Cowley, D. R. 1966. An introduction to New Zealand freshwater insects. ix—100 pp.,
31 figs. Auckland.
Pictet, F. J. 1834. Recherches pour servir a histoire et l’'anatomie des Phryganides. 233 pp. Geneva.
Riek, E. F. 1970. Trichoptera (caddis-flies, caddises). In Mackeras, E. M., (Ed.), The insects of Australia.
A textbook for students and research workers. Pp. 741-764. CSIRO, Melbourne University Press,
Victoria.
Ris, F. 1897. Neue schweizerischen Hydroptiliden. Mitt. schweiz. ent. Ges. 9 : 415-423.
Ross, H. H. 1938. Lectotypes of North American caddis flies in the Museum of Comparative Zoology.
Psyche, Camb. 45 : 1-61.
— 1939. New species of Trichoptera from the Appalachian Region. Proc. ent. Soc. Wash. 41 : 65-72.
— 1941a. New species of Trichoptera from Canada and northern United States. Can. Ent. 73 : 15-19.
— 1941b. Descriptions and records of North American Trichoptera. Trans. Am. ent. Soc. 67 : 35-126.
— 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Ill. St. nat. Hist. Surv. 23 : 1-326.
— 1948. Notes and descriptions of Nearctic Hydroptilidae (Trichoptera). J. Wash. Acad. Sci. 38 : 201-
206.
— 1956. The evolution of the mountain caddisflies. 213 pp. Urbana, Illinois.
— 1967. The evolution and past dispersal of the Trichoptera. A. Rev. Ent. 12 : 169-206.
Roy, D. & Harper, P. P. 1975. New records of Trichoptera from Quebec and description of Limnephilus
nimmoi sp. nov. (Limnephilidae). Can. J. Zool. 53 : 1080-1088.
Schmid, F. 1949. Les Trichoptéres de la collection Navas. Eos, Madr. 25 : 305-426.
238 J. E. MARSHALL
— 1958a. Trichoptéres de Ceylon. Arch. Hydrobiol. 54 : 1-173.
— 1958. Contributions a |’étude des Trichoptéres Neotropicaux. III. Mitt. zool. Mus. Berl. 34 : 183-217.
— 1959a. Le genre Stactobia McL. Misceldnea zool. 1 (2) : 1-56.
— 1959b. Trichoptéres d’Iran. Beitr. Ent. 9 : 200-219; 376-412; 683-698.
— 1960. Trichoptéres du Pakistan III. Tijdschr. Ent. 103 : 83-109.
Scott, K. M. F. 1967. Trichoptera: provisional key to the families (imagos) from Africa south of the
Sahara. News]. limnol. Soc. sth. Afr. 8 : 21-34.
— 1976. The larval and pupal stages of Ugandatrichia Mosely (Trichoptera : Hydroptilidae) from
Rhodesia, with the description of a new species. Ann. Cape Prov. Mus. (nat. Hist.) 11 : 117-127.
Scudder, G. G. E. 1971. Comparative morphology of insect genitalia. A. Rev. Ent. 16 : 379-406.
Sibley, C. K. 1926. A preliminary biological survey of the Lloyd—Cornell Reservation: studies on Trichop-
tera. Bull. Lloyd Libr. 27 (Ent. Ser. 5) : 102-108; 185-247.
Siltala, A. J. 1907. Trichopterologische Untersuchungen. 2: Ueber die postembryonal Entwicklung der
Trichopteren-Larven. Zool. Jb. Suppl. 9 : 309-626.
Solem, J. O. 1972. The larva of Agraylea cognatella McLachlan (Trichoptera, Hydroptilidae). Norsk. ent.
Tidsskr. 19 : 77-79.
Statzner, B. 1977. Taxonomische Studien an den Hydroptilidae-Imagines aus dem zentralafrikenischen
Bergbach Kalengo (Trichoptera). Beitr. Ent. 24 : 393-405.
Stephens, J. F. 1836. //ustrations of British entomology (Mandibulata) 6 : 151-154.
Svensson, B. 1974. Population movements of adult Trichoptera at a south Swedish stream. Oikos 25 :
157-175.
— & Tjeder, B. 1975. Oxyethira boreella n. sp. from northern Sweden (Trichoptera, Hydroptilidae).
Entomologica Scand. 6 : 131-133.
Sykora, J. 1967. Trichoptera collected by Prof. J. Illies in New Guinea and New Caledonia. Pacif.
Insects 9 : 585-595.
Thienemann, A. 1904. Prilocolepus granulatus Pictet, eine Ubergargsform von den Rhyacophiliden zu den
Hydroptiliden. Allg. Z. Ent. 9 : 418-424; 437-441.
—1905. Trichopteren-Studien. Z. wiss. InsektBiol. 1 : 285-291.
Tjeder, B. 1930. Ithytrichia lamellaris Eaton and I. clavata Morton in Dalecarlia. Ent. Tidskr. 51 : 134-138.
Tsuda, M. 1942. Zur Kenntnis den Koreanischen Trichopteren. Mem. Coll. Sci. Kyoto Univ. (B) 17 : 227-
237.
—— & Nakagawa, A. 1959. Fauna hygropetrica of a mountain stream. Jap. J. Ecol. 9 : 134-136.
Ulmer, G. 1907. Trichoptera. Jn Wytsman, P. A. G., Genera Insectorum. Fasc. 60. 259 pp. Hamburg.
— 1910. Trichoptera. Percy Sladen Trust Expedition to the Indian Ocean III. Trans. Linn. Soc. Lond.
14 : 41-54.
— 1912a. Die Trichopteren des baltischen Bernsteins. Schr. phys.-dkon. Ges. Kénigsb. 10 : 380 pp.
—— 19126. Trichoptera aus Aquatorial-Afrika. Wiss. Ergebn. dt. ZentAfr. Exped. 4 : 81-125.
—— 1929. Uber einige deutsche Hydroptiliden. Ecol. Anz. 80 : 253-266.
— 1951. K6cherfliegen (Trichoptera) von den Sunda-Inseln (Teil I). Arch. Hydrobiol. Suppl.19 :528 pp.
— 1957. Kocherfliegen (Trichoptera) von den Sunda-Inseln (Teil III). Arch. Hydrobiol. Suppl. 23 :
109-470.
Vaillant, F. 1951a. Contributions a l’étude des Trichoptéres du genre Stactobia Mcl. Bull. Soc. zool. Fr.
76 : 13-17.
— 1951b. Les larves de Stactobia McL. Bull. Soc. zool. Fr. 76 : 205-208.
—— 1952. Au sujet des Stactobia (Trichoptera). Bull. Soc. zool. Fr. 77 : 285-286.
—— 1956. Recherches sur la faune madicole de France, de Corse et d’Afrique du Nord. Mém. Mus. natn.
Hist. nat. Paris (A) 11 : 1-258.
Westwood, J. O. 1840. An introduction to the modern classification of insects. 2. xi+587 pp., 133 figs.
London.
Wiggins, G. B. 1968. Contributions to the systematics of the caddisfly family Molannidae in Asia (Trichop-
tera). Contr. Life Sci. Div. R. Ont. Mus. 72 : 1-26.
—— 1977. Larvae of the North American caddisfly genera (Trichoptera). xi—401 pp. Toronto.
Abtrichia 183, 228
Acostatrichia 182, 228
Afritrichia 165, 227
Agraules 193
Agraylea 193, 230
Alisotrichia 183, 229
Allotrichia 196, 230
Anchitrichia 181, 228
Aratrichia 165, 227
Argentitrichia 186, 229
Argyrobothrus 203, 232
Baliotrichia 213, 233
Betrichia 182, 228
Bredinia 170, 228
Caledonotrichia 221, 234
Catoxyethira 171, 228
Celaenotrichia 183, 228
Chrysotrichia 170, 228
Clymene 213, 233
Costatrichia 181, 228
Cyllene 189, 230
Dampfitrichia 203, 232
Dhatrichia 199, 230
Diaulus 220, 234
Dibusa 218, 234
Dicaminus 220, 234
Dolotrichia 190, 230
Electrotrichia 222, 234
Exitrichia 190, 230
Gnathotrichia 207, 233
Guerrotrichia 190, 230
Hydropneuma 200, 231
Hydroptila 200, 231
Hydroptilidae 153, 227
Hydroptilina 200, 231
Hydroptilinae 161, 227
Hydroptilini 191, 230
Hydrorchestria 193
Ithytrichia 216, 234
Javanotrichia 213, 233
Lagenopsyche 203, 232
Lamonganotrichia 165, 227
Leiochiton 210, 233
Leucotrichia 178, 228
Leucotrichiini 175, 228
Lorotrichia 190, 230
Loxotrichia 203, 232
Macrostactobia 217, 234
Madioxyethira 173, 228
THE GENERA OF THE HYDROPTILIDAE
Index
Synonyms are in italics; references are to main sections and checklist only.
Mayatrichia 191, 230
Metrichia 186, 229
Microptila 197, 230
Moselyella 198, 230
Neotrichia 189, 230
Neotrichiini 188, 230
Nothotrichia 219, 234
Ochrotrichia 185, 186, 229
Ochrotrichiini 184, 229
Oeceotrichia 200, 231
Orphninotrichia 220, 234
Orthotrichia 213, 233
Orthotrichiini 212, 233
Orthotrichiella 213, 233
Oxydroptila 200, 231
Oxyethira 203, 232
Oxytrichia 203, 232
Padunia 223
Palaeagapetus 160, 227
Parastactobia 173, 228
Paroxyethira 208, 233
Pasirotrichia 200, 231
Peltopsyche 179, 228
Petrotrichia 225
Phrixocoma 200, 231
Plethotrichia 168, 227
Plethus 168, 227
Polytrichia 186, 229
Pseudoxyethira 174, 228
Ptilocolepinae 157, 227
Ptilocolepus 160, 227
Rhyacopsyche 187, 229
Saranganotrichia 216, 234
Scelotrichia 174, 228
Sperotrichia 171, 228
Stactobia 165, 227
Stactobiella 169, 228
Stactobiini 163, 227
Stenoxyethira 207, 233
Sumatranotrichia 200, 231
Synagotrichia 210, 233
Tascobia 169, 228
Tricholeiochiton 210, 233
Tsukushitrichia 225
Ugandatrichia 198, 230
Uenotrichia 223
Xuthotrichia 209, 233
Zumatrichia 179, 228
239
British Museum (Natural History)
Monographs on Trichoptera
A revision of the genus Leptonema (Trichoptera). M. E. Mosely
1933, 69 pp, 200 text figures, frontispiece, 8vo, £2.20
The Francis Walker types of Trichoptera in the British Museum
(Natural History). (Re-described and figured). C. Betten and M. E.
Mosely
1940, ix+248 pp, frontispiece, 122 text figures, 4to £7.75
The Trichoptera (Caddis-Flies) of Australia and New Zealand. M. E.
Mosely and D. E. Kimmins
1953, 550 pp, 364 text figures, £10.50
Titles to be published in Volume 39
A revision of the African pod bugs of the tribe Clavigrallini
(Hemiptera: Coreidae) with a checklist of the world species.
By W. R. Dolling.
A revisionary classification of the Cynipoidea (Hymenoptera) of the
Ethiopian Zoogeographical Region. By J. Quinlan.
A review of the genera of the Hydroptilidae (Trichoptera).
By J. E. Marshall.
A revision of Vitessidia Rothschild & Jordan and Vitessa Moore
(Lepidoptera: Pyralidae). By E. G. Munroe & M. Shaffer.
Printed by Henry Ling Ltd, Dorchester
Bulletin of the .
British Museum (Natural History)
A revision of Vitessidia Rothschild &
Jordan and Vitessa Moore (Lepidoptera:
Pyralidae)
E. G. Munroe & M. Shaffer
Entomology series Vol 39 No4 31 January 1980
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum’s resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and are not
necessarily completed within one calendar year. Subscriptions may be placed for one or more
series. Subscriptions vary according to the contents of the Volume and are based on a forecast
list of titles. As each Volume nears completion, subscribers are informed of the cost of the
next Volume and invited to renew their subscriptions. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW7 S5BD,
England.
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1980
ISBN 0524-6431 Entomology series
Vol 39 No 4 pp 241-260
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 31 January 1980
A revision of Vitessidia Rothschild & Jordan and \
Vitessa Moore (Lepidoptera : Pyralidae)
E. G. Munroe
Biosystematics Research Institute, Research Branch, Agriculture Canada, Ottawa, Ontario
K1A 0C6
M. Shaffer
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis : é : ; , ' ‘ ‘ : : ; : . 241
Introduction . ; ‘ ; ; 3 ~ 241
Abbreviations of museums, institutions and collections : : , : . 243
Acknowledgements . ; ; : : ; : ; ; ; . 244
Species transferred from Vitessa : 4 , ; : . 244
Check-list of genera, species-groups, species and subspecies , ; : ‘ . 244
Key to genera : ' : : ; ; : . 245
Vitessidia Rothschild & Jordan, 1905 F : ; ; j : : 240
Key to the species of Vitessidia : F F : ‘ g , : eos
Cosmethella gen. n. : ; : : : ‘ : ; . 254
Key to the species of Cosmethella ‘ a : : : : : ‘ i 256
Vitessa Moore, [1860] : : ; : : : ; F OO
Key to the species and subspecies of Visa ; : : : : . . 264
The griseata-group . j ; ‘ ; : ‘ : , : e2 2267
The zemire-group : . ; : ; : ; : ; ; ree -7 fi!
The hieratica-group . : , F : ; : § ; : . 284
The stettina-group. ; ‘ ; ; ‘ : . , : , 200
The hemiallactis-group : ; : : : : ; ; § =, 291
The pyraliata-group . = : ‘ : : ‘ : * , - = 303
The plumosa-group . , ‘ ‘ : . , ; ' 5 * 314
The suradeva-group . F ; ‘ : : ; : , : Fae
References. . : ; ; : : , : . ; ; . 324
Index. . : : ; ; : : é ; : ‘ F 2 359
Synopsis
The genera Vitessidia Rothschild & Jordan and Vitessa Moore are revised and a new genus is described.
Two species of Vitessidia, three species of Cosmethella gen.n., and twenty-eight species and sixteen
subspecies of Vitessa are described, of which fifteen species and fourteen subspecies are new. One new
synonym, two new status and two new combinations are established, eleven lectotypes are designated.
Keys to the genera, species and subspecies are provided, the distributions of the species and subspecies
are discussed and are illustrated with maps, and notes on the biology and descriptions of the immature
stages are given for Vitessa suradeva rama Moore. Wing patterns and genitalia are illustrated for all
species and subspecies, and for both sexes where available.
Introduction
This paper revises the classification of three genera of large, conspicuous and at least partly
diurnal moths of the family Pyralidae (subfamily Pyralinae). Except for revisional notes on
Endotrichini (Whalley, 1961; 1963), there has been no study of the inter-relationships of genera
of the Pyralinae since Hampson (18695), who used characters now known to be inadequate. The
Bull. Br. Mus. nat. Hist. (Ent.) 39 (4): 241-360 Issued 31 January 1980
941
3RAR‘
242 E. G. MUNROE & M. SHAFFER
three genera can now be defined as belonging to the Pyralinae from the following characters:
chaetosema present; maxillary palpus present; proboscis well developed; fore wing with R;
stalked with R, and R,; hind wing with Rs not anastomosed with Sc+R, (Pyralini), or Rs
anastomosed with Sc+ R, (Endotrichini); median vein non-pectinated. According to the current
definition of the Endotrichini, Vitessidia keys to this tribe, whereas Cosmethella and Vitessa key
to the Pyralini. Certain features of the ¢ genitalia in Vitessidia appear to be endotrichine in
character: the uncus is wide and flattened dorsoventrally, and distally bilobed in this genus,
whereas in typical Pyralini, including Cosmethella and Vitessa, the uncus is single; the gnathos in
Vitessidia has anteroventrally directed lateral arms and a strongly bifid medial element, as
opposed to the transverse lateral arms and single, laterally compressed, blade-like medial element
in Cosmethella and Vitessa. Vitessidia, however, differs from known Endotrichini, as well as
from Cosmethella and Vitessa, in the following features: the eyes bear interfacetal hairs, a very
unusual character in the Pyralidae; and R; separates from R, distal to the separation of R;, not
basal to it as in Endotrichini and many Pyralini. In external features and in ¢ and 9 genitalia,
Vitessidia is not, however, closely related to any Endotrichini known to us at present, and we
cannot place it in this tribe with any certainty. The genera Cosmethella and Vitessa, though
typical of the Pyralini in most characteristics, have wing markings that are quite distinct from
those of any other genera, and at present their generic relationship is uncertain. Because Vitessidia
has been associated with Vitessa (including Cosmethella) in the generally accepted classification,
and because their relationship has not previously been questioned, we think it useful to consider
the genus in the present paper.
The life-history of this group is known through the work of T. R. Bell, who made manuscript
notes on one species of Vitessa, and preserved larval and pupal exuviae. These are discussed
under the genus Vitessa (p. 262) and under Vitessa suradeva rama (p. 322).
Geographically the genera extend to North India and Sri Lanka in the west, to Queensland,
Australia, in the south, and to Fiji and Samoa in the east. All three genera occur in New Guinea;
Vitessidia appears to be confined to that island. Cosmethella is represented also in the Solomon
Islands; Vitessa occupies the whole range of the group but the largest concentration of species
and species-groups is in the Papuan region. This pattern of distribution is consistent either with
evolution in and radiation from a Papuan centre, or with radiation from an Asiatic centre, with
outward displacement and secondary local radiation of successive ancestral stocks. The latter
alternative would agree with the interpretation of Belkin (1962) for mosquitoes and the model of
Wilson (1959) based on ants. The distribution of Vitessa is discussed in greater detail under the
generic heading.
The colour patterns in these genera are striking and depart widely from those normal in the
Pyralinae. The nature of the patterns and their boldness suggest that they may be mimetic or
aposematic. There is indirect evidence that aposematism, Miilerian mimicry and Batesian
mimicry may all occur among the genera considered. The hypothesis of mimicry is supported by
the convergent resemblance of the moths to members of several different groups of Lepidoptera
and other insects, some of them potential or possible models.
Considering first the genus Vitessidia, the resemblance is to ithomiine butterflies of the genus
Tellervo Kirby (see Vane-Wright, 1971), with which other Lepidoptera of several families con-
stitute a mimetic association, probably partly Miillerian, partly Batesian. The relative rarity of
the two Vitessidia species as compared with the more abundant Tel/ervo butterflies suggests that
the former may be Batesian mimics of the latter.
We have not investigated in detail the possible mimetic associates or models of Cosmethella
species. The two rare New Guinea species are Vitessa-like in fore wing pattern and probably
belong to the same mimetic groupings as members of that genus. The more common Cosmethella
unipectinalis, from the Solomon Islands, is reminiscent of certain Homoptera, but the similarity
has not been assessed under field conditions. j
Vitessa species very closely resemble such Agaristidae as Argyrolepidia megisto (Boisduval) and
its relatives. Certain Agaristidae are thought to be aposematic models in Africa (Guillaumin,
1976), but so far as we know this has not been demonstrated for Argyrolepidia. Munroe and
Holland collected Argyrolepidia species in New Guinea at mercury-vapour light, but in a ratio
REVISION OF VITESSIDIA & VITESSA 243
of abundance of only | : 100 to the similarly coloured Vitessa species. However, many Agaristidae
are mainly diurnal, and this may well be true of Argyrolepidia. At least some Vitessa species are
also diurnal in part. In India, T. R. Bell (manuscript notes in BMNH) recorded V. suradeva rama
as flying in small numbers in the daytime at flowers of Acanthocephalus indicus Achille Richard
(Myrsinaceae) in company with larger numbers of the hypsid moth Macrobrochis gigas Walker,
which has only a very general resemblance to this Vitessa. Any mimetic association between these
two species is presumably only of partial efficiency. Any advantages that arise from these
resemblances between Vitessa species and Agaristidae most likely depend on selection by diurnal,
visually searching predators. Such selection might be exercised as easily on resting moths as on
active ones.
A number of other moths resemble Vitessa species and probably enter the same mimetic
associations. Tyspanodes hillalis (Schaus), celebensis Munroe and radiata Kenrick, and Nevrina
procopia (Stoll) (Pyralidae : Pyraustinae) bear some resemblance to Vitessa species. Heortia
polyplagalis Hampson and dominalis Lederer (Pyralidae : Odontiinae; see Munroe, 1977) closely
resemble species of the Vitessa zemire- and griseata-groups, whilst Heortia vitessoides (Moore)
and H. iridia Munroe resemble species of Vitessa that have black and white hind wings. Jmma
saturata (Walker) (Immidae) resembles Vitessa cyanea.
As well as resembling other moths, Vitessa species have a general appearance much like that of
some lycid beetles. This group is generally considered to be aposematic, and its species are thought
to be mimicked by a number of other insects. However, Drs Henry F. Howden, P. J. Darlington
Jr and J. Linsley Gressitt, all of whom are familiar with the Papuan beetle fauna, were unable to
suggest likely beetle models in that geographical region.
We think it possible in view of their relative abundance, rather slow flight and somewhat
leathery texture that Vitessa species are aposematic and that they may act as models or Miillerian
co-mimics rather than as Batesian mimics. Snellen (1890 : 560) mentions a 3 of Vitessa suradeva
from the Naga Hills with a label stating that the anal tuft had a scent of bitter almonds. We
have not located this specimen. Much stronger corroborative evidence will, of course, be required
before we can firmly conclude that Vitessa species are aposematic.
Throughout the text wing measurements given are double the distance from wing apex to
centre of thorax. Original altitude records from data labels have been converted from feet to
metres and rounded to the nearest 50 metres except in the cases of very low altitudes.
Abbreviations of museums, institutions and collections
ANIC Australian National Insect Collection, C.S.I.R.O., Canberra, Australia.
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.
BMNH British Museum (Natural History), London, U.K.
CAS California Academy of Sciences, San Francisco, California, U.S.A.
CNC Canadian National Collection, Biosystematics Research Institute, Agriculture Canada, Ottawa,
Canada.
CM Carnegie Museum, Pittsburgh, Pennsylvania, U.S.A.
CMAG City Museum & Art Gallery, Birmingham, U.K.
CU Cornell University, Ithaca, New York, U.S.A.
HI Private collection of H. Inoue, Iruma City, Saitama Prefecture, Japan.
KRS Koronivia Research Station, Nausori, Fiji.
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A.
MM Manchester Museum, Manchester, U.K.
MNHN Muséum National d’Histoire Naturelle, Paris, France.
NMNH National Museum of Natural History, Washington, D.C., U.S.A.
NMNZ National Museum of New Zealand, Wellington, N.Z.
RNH Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands.
RSM Royal Scottish Museum, Edinburgh, U.K.
™ Transvaal Museum, Pretoria, South Africa.
UM Ubersee-Museum, Bremen, West Germany.
UM University Museum, Oxford, U.K.
ZM Zoologisk Museum, Copenhagen, Denmark.
244 E. G. MUNROE & M. SHAFFER
Acknowledgements
We thank the following colleagues for the loan of specimens for study: Mr A. Brindle, MM,
Manchester; Dr J. F. Gates Clarke, NMNH, Washington; Mr H. K. Clench, CM, Pittsburgh;
Dr I. F. B. Common, ANIC, Canberra; Dr P. J. Darlington, Jr, MCZ, Cambridge, Mass.; Dr
J. G. Franclemont, CU, New York; Dr M. W. R. de V. Graham, Mr J. W. Ismay and Mr E.
Taylor, UM, Oxford; Dr J. Linsley Gressitt, Dr F. G. Howarth and Rev. Father J. C. E. Riotte,
BPBM, Honolulu; Mr P. Hanney, CMAG, Birmingham; Dr H. Hohmann, UM, Bremen;
Dr H. Inoue, Iruma City, Saitama Prefecture, Japan; Dr R. de Jong, RNH, Leiden; Mr H. B.
Leech, CAS, San Francisco; Dr P. A. Maddison, UNDP/FAO Survey of Agricultural Pests and
Diseases, Fiji; Mr E. S. Nielsen, ZM, Copenhagen; Mr R. D. Ordish, NMNZ, Wellington; Mr
E. C. Pelham-Clinton, RSM, Edinburgh; Dr L. Vari, TM, Pretoria; and Dr P. Viette, MNHN,
Paris.
Mr Douglas Kritsch of the Entomology Research Institute, Canada Department of Agriculture,
Ottawa, gave extensive and invaluable technical help. Most of the photographs of moths were
made by Mr Orin Hanright of the Graphics Unit, Scientific Information Service, Agriculture
Canada, with the assistance of Mr Kritsch; photographs of some moths and all genitalia except
for the photograph of the ¢ genitalia of Vitessa brandti were made by Mr Peter York of the
Photographic Unit, British Museum (Natural History).
Dr H. F. Howden of Carleton University, Ottawa, Dr P. J. Darlington Jr and Dr J. Linsley
Gressitt advised on possible coleopterous models for Vitessa species. Dr P. E. S. Whalley, British
Museum (Natural History), initiated the studies for this present paper and continued giving
valuable advice and criticism during its preparation. Dr K. Sattler, British Museum (Natural
History), also made many useful suggestions.
We are indebted to our colleagues in Ottawa and London for their comments and advice.
Species transferred from Vitessa
Vitessa pyraliformis Walker, 1864 : 220 (Oxford Type no. 546; UM, Oxford) [examined], not Vitessa
pyraliata Walker, 1864 : 220, though so misinterpreted by Ragonot (1891 : 646). Transferred to the
family Agaristidae by Swinhoe (1892 : 159); now regarded as a synonym of Argyrolepidia megisto
Boisduval, 1832.
Vitessa triplaga Walker, 1869 : 8 (holotype ¢ in BMNH) [examined]. Transferred to the subfamily
Pyraustinae as a synonym of Heortia dominalis Lederer, 1863, by Hampson (1898 : 751). Later
transferred to the subfamily Odontiinae by Munroe (1977 : 436).
Check-list of genera, species-groups, species and subspecies
VITESSIDIA Rothschild & Jordan, 1905
diaphana Rothschild & Jordan, 1905
geometrina sp. n.
COSMETHELLA gen. n.
unipectinalis (Hampson, 1906) comb. n.
minor sp. n.
major sp. n.
VITESSA Moore, [1860]
Cosmethis sensu auct. nec Hiibner, [1820]
griseata-group
griseata Kenrick, 1907
brandti sp. n.
talboti (Janse, 1924) comb. n.
nobilis sp. n.
zemire-group
cyanea ceramensis subsp. n.
cyanea cyanea Hampson, 1906
REVISION OF VITESSIDIA & VITESSA
zemire expansa subsp. n.
zemire zemire (Stoll, 1782)
zemire temerata Swinhoe, 1906 stat. n.
zemire nephritica subsp. n.
zemire coronis subsp. n.
zemire novaebritanniae subsp. n.
zemire solomonis subsp. n.
zemire cheesmanae subsp. n.
vitialis Hampson, 1906
tamsi sp. n.
hieratica-group
hieratica Swinhoe, 1900
stettina-group
Philippina sp. n.
stettina Swinhoe, 1906
hollandi wallacealis subsp. n.
hollandi hollandi subsp. n.
hemiallactis-group
ternatica Lederer, 1863
sarumensis Holland, 1900 syn. n.
hemiallactis moluccana subsp. n.
hemiallactis hemiallactis Meyrick, 1887
hemiallactis admiralitatis subsp. n.
hemiallactis lustrans subsp. n.
intermedia sp. n.
barretti sp. n.
glaucoptera Hampson, 1906
cristobalensis sp. n.
pyraliata-group
nicobarica Hampson, 1896
pyraliata latialbata subsp. n.
pyraliata triangulifera subsp. n.
pyraliata pyraliata Walker, 1864
kolakalis sp. n.
sulaensis sp. n.
muluana sp. n.
gemina sp. n.
splendida Schultze, 1908
plumosa-group
plumosa plumosa Hampson, 1896
plumosa salayerensis subsp. n.
suradeva-group
teleroma Swinhoe, 1906
suradeva suradeva Moore, [1860]
formosa Felder & Rogenhofer, 1874
suradeva rama Moore, 1885 stat. n.
Key to genera
245
1 Antenna simple, either ciliated or pilose, weakly compressed, its length more than three-
quarters that of fore wing; fore wing with discocellular vein at centre strongly angled toward
base
VITESSIDIA (p. 245)
— Antenna laminate, unipectinate or bipectinate, its length less than three-quarters that of fore
wing; fore wing with discocellular vein at centre not strongly angled toward base . ‘ 2
2 Antenna unipectinate in both sexes; fore wing with R, stalked with Rs, ;
COSMETHELLA (p. 254)
— Antenna bipectinate in 3, laminate in 2; fore wing with R, arising from cell VITESSA (p. 258)
VITESSIDIA Rothschild & Jordan, 1905
Vitessidia Rothschild & Jordan, 1905, Novit. zool. 12 : 475. Type-species: Vitessidia diaphana Rothschild
& Jordan, 1905, by original designation and monotypy.
E. G. MUNROE & M. SHAFFER
246
‘sdeuwl uonnqiisiq p-[ ssi
y st[tqou a eulwas
a (oq[ed
@ Qpueag gs s0lew
@ vieastis @® 10uULW
r wy 008 ly wy 008
S01 $014
0o ©0
wR v4 ae a aR
- YOOV 14d IHOUY
: MOUVWSIC
ete a
VSSHLIA
eauing
MAN Endeg
eauing
MIN ended
uoquy
v-
22 -
Ps a,
e —
wRias
VSSSLIA
W\v-riantawsoo hai ts
oes 1O0jJWNN ¢
is "| LeQoisisy ues
VSSHLIA
V'TIAHLANSOO
f W100 7
4,09
SGNV1SI NOWO10S
@ stsua[eqois1io
e@ stpeunsadiun
$014
t 4yYOO8
“TP RLLaABT RIIIA
Bouingy
MIN endeg
C VICISSA.LIA
B eUlI9UI0a3
e@ eueydeip
$014
247
REVISION OF VITESSIDIA & VITESSA
‘sdveul uonnqijsiq g
¢ SSI
T T
@ sluolwo[os ar11waz : a oavluuryliqaeaou aitwaz
@ stuotod a1lwaz
E “| peueopepens S,0I4
d ie tA O02. at
“s] TE 8 Q 2 - eer
S! BAOpUsY
& @ omy : e\
£ | toqtun ©
$8
r Wx 007 =) é
b Sr
puelri}
4 9 Si m2N ODVIAdIHOUV i.
: . WoOUVWSIa
SANV1S1 NOWO10S © ;
a
.
"| Aulesnog JIAOUURH MAN 7
EOL 38st oe 4,0SL EM} al
VSSALIA cai oF VSS:LLIA ¥ | Xv 2 |
@ eoltaydou artuaz . i uoquy B® d911WI9Z JIIW9Z
enuedeg @
@ ev eIDWI9A] J11WUI9Z ~ a I e@ esuedxa d1lulaz
~ | °
ac
Av)
O
oa
SV¥)INI10OW \
41008 1
“] equies
e $014
J 0
~N g eo oe
eg, Q: Omar
1 pe ar ve ° _
eauinn Ae mo
MON BNdeg nay hl
uene(ndsy s.s4
‘ . ° t wx O07 1
— =
h A PB
= ° <
; A zel ¢
VSSALIA VSSHLIA {
=
E. G. MUNROE & M. SHAFFER
248
*sdeul uonnqidjsiq
71-6 S8IY
a SISUJLUBI9D BQURAD
e sents
@ voueaod vourad oe
% [>
naoq vee ue ¢
ecu Qn
neqelerynweg s
7 vy.
e c
b-- dnoig e § $815
neq :
‘ wy 008 ° : at oa
S017 ; © 4 . i
ee * 4 :
t- i SoZl4
a t
*S] Xneaiseo2i1ug Gg
r Ww
— “| vossn8i34@—7~ 0 ‘oe
d Bauingn ila
MON BNdeg
SOS ~
Fe 4y6Zl 4,82
VSSALIA of of OL
t J @ aeUueWSaayo arIWiaz
& i oS
nr
EMT At 4,0rt
VSSALIA a oF C L :
' Lf
@ tsweq
21ejq
ey
e .
ry t wyxOOL ’
ne
he 7 a lI4
a eypnyaew
O¢ P
ase =—1 VOWVS NUYalsam e
r S3GIYaH MAN \ : Sia
€
“| ojuReg Mytuidsy
S.€l4
©
Mott MotZt LL An 4 0Zt 45691 Go 6
VSSALIA Sn a VSSALIA am i 1
249
REVISION OF VITESSIDIA & VITESSA
‘sdew uonNqiysiq 9T-€T 88
' 41 008 1
eauing
MIN ENdeg
X.
VSSALIA
VSSALIA
@ (pueljoy ipueypoy
r wx O08
uainoyos,
—
> se
5, Ce
soo funn 9 bvssauia Aig
oo
eauingy
mIN endeg
@® euljjo1s
ue gy
B esajynsueis eierpesdd RSS
v epipuatds
4 e@ eutddiyiyd ngs
N,0l7
N§t5
SA VA Lyssa
VISANOQNI
4, Ol
@ eoneioly
“1 008 1
E. G. MUNROE & M. SHAFFER
250
‘sdew uonngiysiq = O7-LI S81
VSSAHLIA
vy esraidoone[3s
m@ (o11eq
@ eipowtaiul
Puejsusand
$014
fo
eauing
MAN endeg
YAR ALIA
ox T
B® SUPIISN] SIOLI[ eWay
@ sleilpeatwupe siovjperuoay
f WIGS 07. 4
bee [euedjepeny :
: rae 10219
|
ed | epee
Sal
SGNVTSI
NOWO10S
on “TP atjiaute3nog
ia
' 4
, OOV14dIHIUV
: WOUVASIB
i . -
my 37
| eeu Bs i :
yaaouueH MON gf eseey Sonics
by | snuew *.
44091 15081 6 |
VSSALIA
r wy 008 1
@ Sijoeetuay SsHoelpermoy
S,Ol>
gauing
MIN ended
oan ey
Oo
$o$4
piey Se
tre
4,071
leas
VSSHLIA
@ eoneuial
L, iS a y vy eueoON[OW soe] [eWay
> 4 .
Payee ees
~
; @ sljeaoR][eM Ipuel[loy
VISANOGNI ' t
cee 6
eae wO8 ae al ; Sf
SoSs
XN 5
] {)
fe 2
<4
-
ning
wesas
SV. INIOW i
Gio ; aan.
< GOS
S,
BL
Le “ Y Som
.
ee—ue
aieusay
1,0C1 ny
Pas)
REVISION OF VITESSIDIA & VITESSA
‘sdew uonnginsiq = p7-17 ssl
f 4*7OOSL 1
~OTRjadiyIsy 4
VSSALIA ove
J @ eUIBI BAQpLANS
%, e@evAapeins eAapeins
eyucy] tus
ewin
a Ysape|3ueg
eipuy
(183u2g “M)
eipuy
VSSAHLIA
T T @ eBWO1a] 31
eilewng
| eunieN
f “YOO9
1,011
a sisuaidAejes esound
vy esouinjd esoun|d
= euen[nw
@ SIsuoeins
b ST[exel[OY
@ ejelesdd eqerpeshd
sow 4
oo ~
e¢
Ss
VSSALIA oa
ia Ip, ‘ ;
Se ‘@
SVDDNI1OW
Pijesisny
VISINOGNI
No ‘ YWAOO SL 1
°
(ise3) VISAV IVAW
Zé VSSALIA
T
@ eleq(ene, eyerperAd
@ eolreqoolu
eyuey las
JEQOIIN 18H
moouen%*— @ @)
B —o- 31 18Q05IN
.
ewing }
252 E. G. MUNROE & M. SHAFFER
Head small. Frons rounded, shaggily or smoothly scaled, anteriorly directed plume of scales between
antennae. Vertex short, scaling erect, woolly. Eye well developed, with long interfacetal hairs over
central area. Ocellus large, conspicuous. Chaetosema elliptical, directed laterally, with long radiating
setae. Antenna nearly as long as fore wing; thick, compressed, heavily ciliated in 3; evenly short-pilose
in 2. Labial palpus in $ prominent, curved, upturned; first segment short, weakly curved; second segment
thick, long, strongly curved, extending well above vertex, distally expanded, obliquely truncate; third
segment acuminate, obliquely ascending from a subterminal origin on anterior surface of second segment.
Labial palpus in 2 shorter, obliquely ascending; first segment as in $; second segment slender, nearly
straight, reaching about level of vertex; third segment straight, acuminate, continuing line of second.
Maxillary palpus minute, directed medially toward base of proboscis. Proboscis long, strong, scaled at
base. Neck region with slender erect scales. Body slender; abdomen long, exceeding hind wing. Legs
long, slender; outer spurs shorter than inner; tarsi with long ventral spines. Fore wing in ¢ broad; costa
nearly straight; apex rounded; termen curved, oblique; tornus angled; posterior margin nearly straight,
narrowly curved at base. Wing in 9 broader, costa convex, apex and termen more rounded. Costal area
wide. Cell about half length of wing. Discocellular vein strongly angled at middle. Sc approaching or
anastomosing with R, postmedially. R, from before apex of cell, approaching or anastomosing with Rp.
R; stalked with R, ,;. Ra, Rs long-stalked. M, stalked for a short distance with R; .;. M2, M; stalked from
posterior angle of cell. Cu, from just behind posterior angle. Cu, from cell at middle. CuP preserved near
wing-base and termen. A, straight. A, forming a small loop and spur with A,. Hind wing broad; apex,
termen, anal angle rounded. Costal area wide. Cell about half length of wing. Discocellular vein anteriorly
straight, erect; at middle obtusely angled, with a distinct medial spur extending toward base; posterior
part straight, oblique distad. Sc+ R, anastomosed for most of postcellular length with Rs. M, basally
apposed to or stalked with Rs. M, and M; stalked from posterior angle of cell. Cu, from behind posterior
angle. Cu, from cell at middle. CuP and two anals present. Frenulum of 2 with two long setae.
GENITALIA ¢ (Fig. 135). Uncus wide, deeply bifid, each process trumpet-shaped, with shaggy dorsal scales.
Gnathos wide; lateral arms broad, dorsoventrally flattened, anteroventrally directed; medial element
formed of paired claw-like, admedial, posterior processes, arising from a transverse band. Transtilla with
broad triangular lateral elements, joined by a weak semimembranous medial section. Tegumen wide,
arched; sclerotization narrowly stronger at origin of gnathos. Vinculum broad and rounded. Valve
with broad rounded distal expansion; costa sigmoidal; a strengthening ridge, obsolescent distally;
narrowly inflated and sclerotized, with small triangular expansion dorsally at tip. Juxta ovate, lightly
sclerotized, with large V-shaped posteromedial emargination; on dorsal surface of anellus a posteriorly
pointed V-shaped catena, with strong medial ridge. Aedeagus long, narrow and sinuate, basally rounded;
junction of ductus ejaculatorius subbasal; manica without spines; vesica minutely spinose. Cornuti small,
few and isolated, possibly deciduous.
GENITALIA ° (Figs 219-222). Ovipositor lobes low, cushion-like, dorsally rounded and separated. Setae
short, longer on outer margin, all simple. Posterior apophysis short, with a symmetrical vertical bar.
Anterior apophysis slightly longer, stronger, distally tapering, weakly bent near base. Eighth sternite
triangular, deeply indented anteriorly, sides elongated and acuminate. Eighth tergite rectangular, ventro-
laterally rounded, covered with setae. Ostial chamber small, unarmed. Ductus bursae long, slender,
membranous; base with a pair of lateral sclerites extending to junction of ductus seminalis. Corpus
bursae oval or comma-shaped, finely denticulate posteriorly, with weakly sclerotized signum.
REMARKS. This genus is known from two species, both from the central mountain ranges of New
Guinea. We have no record of the habits, but the pattern suggests that the moths are diurnal.
The genus differs from Cosmethella and Vitessa in a number of striking external and genital
characters (see notes in Introduction), though there is a similarity in the structure of the labial
palpi. In Vitessidia, however, they are longer in the gf than the 9, whereas in Cosmethella they are
longer in the 9, and in Vitessa of the same length in both sexes. In the $ genitalia the anellus
lacks lateral and medial spines, and is not reflexed after eversion; the ventral junction of the
vesica to the aedeagus is plain, not rugose, hence the vesica is not reflexed after eversion. In this
respect Vitessidia is similar to Cosmethella, but differs from Vitessa. In the 2 genitalia the
ovipositor lobes have simple setae, as in Cosmethella, unlike Vitessa where the ovipositor bears
flattened spiny setae in addition to simple setae.
BIOLoGy. Early stages and host-plants unknown. Moths taken by various collectors from January
to April and September to December.
REVISION OF VITESSIDIA & VITESSA 250
DISTRIBUTION (Fig. 1). New Guinea: Irian Jaya; Papua New Guinea.
Key to the species of Vitessidia
1 Fore wing upperside with broad proximal and narrow curved preapical, translucent white areas;
veins on white areas of upperside of fore and hind wings narrowly black . diaphana (p. 253)
— Fore wing upperside with irregularly ovoidal distal translucent white area; veins on white areas
of fore and hind wings whitish buff, with little trace of infuscation . ‘ geometrina (p. 253)
Vitessidia diaphana Rothschild & Jordan, 1905
(Figs: 1,:45;,46, 135, 219, 221)
Vitessidia diaphana Rothschild & Jordan, 1905, Novit. zool. 12 : 475. Holotype 3, NEw GuINEA: Papua,
Angabunga R., affl[uent] of St Joseph R., 1850 m (‘6000 ft’) upwards, xi.1904-ii.1905 (Meek) (genitalia
slide no. 1503; BMNH) [examined].
Vitessidia diaphana Rothschild & Jordan; Janse, 1928 : 88.
3, 2. 47-56 mm. Frons shaggily scaled; in ¢ white, laterally bordered with black; in 2 black, admixed
and laterally bordered with white. Plume of scales between antennae white; in 2 smaller, with less white.
Vertex black. Eye black with pale interfacetal hairs. Ocellus buff, shiny. Chaetosema with pale yellowish
setae. Antenna black. Labial palpus black; weak pale grey scaling on dorsal surfaces of second and
- third segments in 3g; pale grey scaling absent in 2; ventral surfaces of first and second segments lightly
admixed with white. Maxillary palpus buff-tinted. Basal scaling of proboscis black, minutely admixed
with white at base in 3, with more white in 9. Neck region with black and orange scaling. Thorax above:
blackish fuscous; patagium and tegula orange, scales black-tipped. Abdomen above: black, terminal
segments orange. Body below and legs black; pectus, fore coxa, parts of mesal surfaces of all femora, and
terminal abdominal segments orange. Fore wing above: ground colour, including fringe, black, with weak
violaceous reflections. An irregularly quadrate, nacreous, translucent white area occupying much of
proximal half of wing. A similarly coloured curved preapical area from cell at R; to cell at M3. Both areas
intersected by narrow black vein-lines. Hind wing above: ground colour as on fore wing. A large ovate,
nacreous, translucent white subbasal and discal area, intersected by black vein-lines, narrower than those
of fore wing. Underside of fore and hind wings like upperside but less violaceous; retinaculum of fore
wing glossy dark grey.
GENITALIA ¢ (Fig. 135). As described for genus.
GENITALIA @ (Figs 219, 221). In general as described for genus. Posterior apophysis slightly curved and
tapering. Anterior apophysis evenly curved and weakly sinuate, tapering and finely acuminate. Ostial
chamber weakly sclerotized, posterior opening wide, narrowing sharply anteriorly; proximal part of
ductus bursae about six times as long as wide, paired sclerites long, narrow, strong. Corpus bursae oval;
signum on posterior lobe near entrance to ductus bursae, a weakly sclerotized incurved plate, finely
denticulated.
REMARKS. The wing markings, though similar to those of V. geometrina, are quite distinct. In the
2 genitalia the apophyses are shorter and thicker; both the whole ductus bursae and its proximal
parts and sclerites are longer. The corpus bursae is oval, not comma-shaped, and the signum is
smaller and simpler.
BioLocy. Early stages and host-plant unknown. Moths collected in January to November.
DISTRIBUTION (Fig. 1). New Guinea: Irian Jaya; Papua New Guinea.
MATERIAL EXAMINED
New Guinea: 6 3, 8 9, Irian Jaya, Weyland Mts, Menoo Valley, Mt Kunupi, 1850 m, xi.1920-i. 1921
(C., F. & J. Pratt) (BMNH; TM, Pretoria). In addition, Dr Vari examined 19 3, 11 2 not seen by us (TM,
Pretoria).
Vitessidia geometrina sp. n.
(Figs 1, 47, 220, 222)
3. Unknown.
2. 44-48 mm. Frons smoothly scaled, black, laterally bordered with white. Plume of scales between
antennae black with violaceous reflections; vertex of same colour, smoothly scaled; a small yellow tuft
254 E. G. MUNROE & M. SHAFFER
behind antenna. Eye, ocellus, chaetosema as in V. diaphana. Antenna black, scape with slight admixture
of white on inner surface. Labial palpus black, ventral surface of first and base of second segment admixed
with white. Maxillary palpus buff-tinted. Basal scaling of proboscis black, admixed with white at extreme
base. Neck region with black-based yellow scales. Body above, including patagium and tegula: black with
violaceous reflections; border of anterior portion of terminal abdominal segment yellow. Thorax below
and legs black; pectus, parts of fore and mid coxae yellow admixed with white; ventral surfaces of mid
and hind femora white. Abdomen below: black, posterior margins of segments broadly white admixed
with yellow, continuing narrowly on lateral surfaces to height of spiracles; terminal segment completely
yellow. Fore wing: ground colour above, including fringe, black, with weak violaceous reflections. An
irregularly ovoidal, nacreous, translucent white area on distal part of wing. Hind wing above: ground
colour as on fore wing. A large, ovate, nacreous, translucent, white subbasal and discal area extending
broadly to anal margin, showing whitish buff veins with slight infuscation; vein-lines of same thickness
as in fore wing. Underside of fore and hind wings like upperside but less violaceous; on fore wing white
admixture from base, spreading along anal margin and lightly up the termen; retinaculum glossy light
grey; on hind wing white at base spreading along base of costa.
GENITALIA ° (Figs 220, 222). In general as described for genus. Posterior apophysis slightly curved and
tapering. Anterior apophysis evenly curved and weakly sinuate, noticeably widened for first third of
length, distally tapering, small central swelling at beginning of last third, tip blunt. Ostial chamber weakly
sclerotized, posterior opening narrow, leading abruptly to ductus bursae. The latter shorter than in V.
diaphana, its proximal part two to three times as long as wide, the paired sclerites relatively short. Corpus
bursae distinctly comma-shaped; signum weak, elongate, nearly half as long as corpus, extending basad
from junction of ductus bursae, forming deep, curved indentation along side of corpus, with evenly spaced,
rib-like ripples at right-angles to margin along entire length. Opposite margin of corpus bursae finely
denticulated.
REMARKS. The moths are smaller than in V. diaphana and have several distinctive external and
genital characters. The ground colour of the fore and hind wings is lighter than V. diaphana. The
white part of the fore wing is reduced to one patch. The white areas in this species are less
nacreous.
BioLoGy. Early stages and host-plant unknown. Moths collected in January to April, September
to December.
DISTRIBUTION (Fig. 1). New Guinea: Irian Jaya; Papua New Guinea.
MATERIAL EXAMINED
Holotype 2, New Guinea: Papua, Upp[er] Aroa R., i-iv. 1903 (Meek) (BMNH).
Paratypes. New Guinea: 2 9, Irian Jaya, Snow Mts, nr Oetakwa R., up to 1050 m (*3500 ft’), x—xii.1910
(Meek) (genitalia slide no.’s 1504, 15282; BMNH); 1 9, I. J., Snow Mts, Upper Setekwa R., 600-900 m
(‘2000-3000 ft’), ix.1910 (Meek) (genitalia slide no. 14675; BMNH); 1 9, I. J., Araucaria Camp, 800 m,
ili. 1939 (Toxopeus) (RNH, Leiden).
COSMETHELLA gen. n.
Gender: feminine
Type-species: Vitessa unipectinalis Hampson.
Head small. Frons rounded, narrow, smoothly scaled. Vertex flat, somewhat elevated, with rough,
forward-directed scales. Eye large. Ocellus absent. Chaetosema elliptical, close to eye, directed postero-
laterally, bearing long radiating setae. Antenna about two-thirds length of wing, compressed, distinctly
unipectinate in both sexes. Labial palpus in 3 shorter than in 9; slender, obliquely upturned; first segment
short; second segment long, in ¢ reaching level of vertex, in 2 well above vertex, third segment equal in
length to second in J, shorter than second in 2, acuminate and usually continuing line of second segment.
Maxillary palpus short, directed medially, with distal, weakly dilated, triangular tuft of scales. Proboscis
weak but coiled, scaled at base. Body slender; abdomen long, exceeding hind wings. Legs short, robust,
with long strong tibial spurs, outer spurs shorter than inner; tarsi ventrally spined. Fore wing narrow;
costa nearly straight, strongly curved near base; apex narrowly rounded; termen slightly curved and
oblique; tornus obtuse; posterior margin slightly curved, more strongly at base. Costal area wide. Cell
about half length of wing. Discocellular vein concave distad. Sc well separated from R and R,. R, from
REVISION OF VITESSIDIA & VITESSA 255
cell at two-thirds from base. R,.,; from apex of cell; R, stalked with R,;,,;; R; stalked for a variable
distance with R;,,. M, from anterior angle of cell, or stalked for a very short distance with R, ,,; except
for this stalking its base not apposed to that of R,,;. M, and M, short-stalked from posterior angle of
cell. Cu, from just behind posterior angle. Cu, from cell at four-fifths from base. CuP preserved near wing
base and termen. A, straight. A, forming a small loop and spur. Hind wing narrow; apex, termen and
anal angle rounded. Costal area wide. Cell about half length of wing. Discocellular vein with strong
basally directed angulation before middle, posterior part distally oblique. Sc+ R, not anastomosed with
Rs. M, stalked with Rs for short distance beyond cell. M, and M, stalked from posterior angle of cell.
Cu, from behind posterior angle. Cu, from cell at three-fourths from base. CuP, A, and A, present.
Frenulum of 2 with two long setae.
GENITALIA ¢ (Fig. 136). Uncus narrow, parallel-sided, constricted at base, rounded at tip; tip and postero-
lateral margin with fine setae dorsally. Gnathos with curved lateral arms, meeting at mid-line in an acute
angle, forming strong medial spur. Transtilla strong, complete, with dorsal arch in mid-line. Tegumen
short, wide, angular, with strong shoulders. Vinculum broad, thickened ventrally, dorsally with a curved,
medially directed process on each side, supporting a membranous setose area. Valve short, wide, fan-
shaped; costa nearly straight, with narrow tubular thickening; terminal margin long, curved; sacculus
weakly thickened, not strongly sclerotized, generally recurved, ending in a produced posteroventral angle;
mesal surface densely covered with setae arranged on radiating striations. Juxta large, deeply bifurcate
dorsally; each side with a long slender process, in slide preparations reaching half length of uncus; also
a short, sharp, laterally directed process on each side near the curved ventral edge; anellus simple.
Aedeagus short, narrow, slightly sinuate, basally rounded; ductus ejaculatorius arising mid-way; posterior
end minutely spinose; vesica large, membranous, sac-like, lacking spines but with small, thin sclerotized
bands at either end; cornuti absent.
GENITALIA 2 (Figs 223-225, 269-271, 315). Ovipositor lobes small, narrow, dorsally rounded and
connected. Setae short, longer on outer margin, all simple (Fig. 315). Posterior apophysis long and
slender, vertical bar incomplete. Anterior apophysis as long as but thicker than posterior, slightly sinuate.
Eighth sternite triangular. Eighth tergite rectangular, several transverse rows of setae on posterior portion.
Ostial chamber wide, with long, wide, simple, scobinated antrum in dorsal wall, extending anteriorly
into a short caecum of ostial chamber. Ductus bursae short and wide. Corpus bursae large, cylindrical,
more or less J-shaped; signum absent.
REMARKS. This genus is known from three species, two represented only by holotypes. Though
the holotype of C. minor was taken at light, the apparently mimetic pattern suggests that the
moths are probably mainly diurnal. The genus differs from Vitessidia in both external and genital
characters; as explained in the Introduction, the two genera are probably not closely related.
The eye, unlike that of Vitessidia, lacks interfacetal hairs. The antenna of Cosmethella is uni-
pectinate, not simple as in Vitessidia. The labial palpus is longer in the 9 than in the ¢ in Cos-
methella, the opposite in Vitessidia. The two genera share the relatively plain anellus, lacking
spines and not reflexed after eversion. In both genera the vesica lacks the reflexing mechanism
found in Vitessa.
Cosmethella is closely related to Vitessa but differs in a number of characters. The antenna is
unipectinate in both sexes in Cosmethella, not bipectinate in the g and laminate in the 9. The
labial palpus is longer in the 3 of Cosmethella and much longer in the 9, not shorter and of equal
length in the sexes as in Vitessa; in Cosmethella it has the second segment straight and oblique,
not curved basally. Cosmethella shares with Vitessa the prominent maxillary palpus. The cell is
shorter in both fore and hind wings. R, of the fore wing is stalked with R;,;, whereas in Vitessa it
arises from the cell. Like Vitessa, this genus has Sc+ R, anastomosed with Rs. The genitalia in
both sexes differ very markedly from those of Vitessa, particularly in shape of valve in the 3.
In the 2 the ovipositor lobes have simple setae; in all examined species of Vitessa some setae
are covered with spines. There are also distinct differences in the shape of the antrum, ductus
bursae and corpus bursae.
BioLoGy. Early stages and host-plants unknown. Moths so far taken by only a few collectors
from January to March and from October to December.
DISTRIBUTION (Figs 2, 3). New Guinea: Papua New Guinea. Solomon Islands.
256 E. G. MUNROE & M. SHAFFER
Key to the species of Cosmethella
1 Thorax purplish fuscous; fore wiug with pale antemedial fascia about one-sixth as wide as long.
Solomon Islands : . _ unipectinalis (p. 256)
— Thorax orange-yellow with black spots: fore wing with pale antemedial fascia more than one-
third as wide as long. New Guinea : : 2
2 Longest antennal pectinations in 2 about three times as long as ‘width of antennal shaft; fore
wing with antemedial fascia narrowed behind, forming a semicircular patch; expanse under
30mm _. 4 minor (p. 256)
— Longest antennal pectinations in 9 about s Six times as longa as width of antennal shaft; fore wing
with antemedial fascia scarcely narrowed behind, meeting posterior margin broadly; expanse
over 40 mm : ; : j : / f : ; ; : ; major (p. 257)
Cosmethella unipectinalis (Hampson, 1906) comb. n.
(Figs 2, 48, 49, 136, 223, 269)
Vitessa unipectinalis Hampson, 1906, Ann. Mag. nat. Hist. (7) 17: 217. LECTOTYPE ¢, SoLoMoNn
IsLANDs: Gizo I. (Meek) (genitalia slide no. 15283; BMNH), here designated [examined].
3, 2. 32-38 mm. Frons, vertex orange, with darker shading laterally. Eye black. Chaetosema with setae
yellow. Antenna fuscous. Labial palpus fuscous, ventral surface of second segment in ¢ light brown,
tip of palpus light brown in both sexes. Maxillary palpus fuscous. Basal scaling of proboscis blue-black.
Neck region orange. Thorax above: patagium, tegula and abdomen above metallic blue-black; in 3 a
slight admixture of orange in most abdominal segments; terminal segment orange. Underside of thorax
and abdomen like upperside, terminal segment admixed with black. Legs metallic blue-black. Fore wing:
ground colour above metallic blue-black. A narrow, oblique, white antemedial fascia, slightly curved at
costal end. A white longitudinal stripe in cell R;, a smaller white stripe above. A narrow triangular white
apical patch. Fringe fuscous. Hind wing above metallic purplish fuscous, unmarked. Underside of wings
purplish fuscous. On fore wing faint traces of proximal line and stripe in cell R;; apical patch distinct.
GENITALIA ¢ (Fig. 136). As described for genus.
GENITALIA ° (Figs 223, 269). In general as described for genus. Apophyses nearly straight. Eighth tergite
with irregularly distributed setae. Eighth sternite indented at posterior angle. Caecum slightly wider than
antrum. Sides of antrum slightly curved, broadly rounded posteriorly.
REMARKS. This geographically isolated species can be distinguished by its narrow proximal fascia
and smaller, distinct apical marking, also by its blue-black thorax.
BioLoGy. Early stages and host-plant unknown. Moths collected from January to March and in
November and December.
DISTRIBUTION (Fig. 2). Solomon Islands.
MATERIAL EXAMINED
Paralectotype. 1 3, data as lectotype (abdomen missing; BMNH).
Solomon Islands: 4 3, north side of Choiseul I., xii.1903 (Meek) (BMNH); 3 2, south side of Choiseul
I., 1.1904 (Meek) (BMNH); 1 3, 1 9, Vella Lavella I., ii. and iii. 1908 (Meek) (BMNH); 9 2, 4 9, Gizo L.,
1903 and xi.1903 (Meek) (BMNH; NMNH, Washington).
Cosmethella minor sp. n.
(Figs 3,:50,-224, 270,315)
3. Unknown.
2.28 mm. Frons, vertex orange. Eye fuscous. Chaetosema setae yellow. Antenna black. Labial palpus
black, some orange scaling basally and distally. Maxillary palpus basally black, distally orange. Basal
scaling of proboscis black. Neck region orange. Thorax above: tegula orange, with an anterior black spot.
Abdomen above: metallic blue-black, terminal segment orange. Thorax and abdomen below: blue-black,
terminal segment orange. Legs blue-black, grey-banded. Fore wing above: ground colour black with
purplish reflections. A small orange basal spot on Cu. A narrow, erect, diffuse, white subbasal fascia. A
large, white, nearly semicircular antemedial fascia, not reaching costa or posterior margin. Discocellular
vein and longitudinal veins in terminal half of wing outlined in whitish grey. A creamy apical patch
REVISION OF VITESSIDIA & VITESSA 250
diffusing into the vein-lines. Fringe fuscous. Hind wing and fringe with purple reflections, unmarked. Fore
wing below: fuscous, shading to pale grey on posterior margin. A large, posteriorly narrowing, white
subterminal fascia, narrowly outlined with black anteriorly and distally. Underside of hind wing similar
to upperside.
GENITALIA @ (Figs 224, 270, 315). In general as described for genus. Posterior apophysis slender and nearly
straight. Anterior apophysis curved. Eighth sternite a nearly equilateral triangle. Caecum short and weak.
REMARKS. This species is distinct from C. unipectinalis in fore wing markings and colour of thorax
(see key). The @ genitalia differ from those of C. unipectinalis in the long ovipositor lobes, curved
posterior apophysis, more anteriorly concentrated setae of eighth tergite, shorter caecum, and
broader antrum, with posterior end angled medially rather than round. C. minor resembles C.
major in general appearance, but differs in the shorter antennal pectinations and in the markings
of the fore wing, as follows: the proximal fascia of the upperside is posteriorly constricted in the
present species and the pale apical patch of the upperside is narrow; on the underside the sub-
terminal pale fascia is posteriorly elongate. The eighth sternite forms a nearly equilateral triangle
rather than a wider, truncated one asin C. major. The fore wing markings bear aslight resemblance
to those of Vitessa cyanea, enhanced by the uniformly dark hind wing; in all other aspects the
two species are quite distinct.
BIoLoGy. Early stages and host-plant unknown. Holotype collected in October at mercury-vapour
light.
DISTRIBUTION (Fig. 3). New Guinea: Papua New Guinea.
MATERIAL EXAMINED
Holotype 2, New Guinea: North-east New Guinea, Wewak, Dagua Road, near sea level, 21.x.1957, at
mercury-vapour light (Munroe & Holland) (genitalia slide no. DK 107; CNC, Ottawa; type no. 11,012).
Cosmethella major sp. n.
(Pigs 3.015 2295 201)
3. Unknown.
2. 42 mm. Frons orange. Vertex orange, posteriorly yellow. Eye fuscous. Chaetosema setae yellow.
Antenna fuscous. Labial palpus fuscous; orange scaling on ventral surface of second segment; tip of
palpus light brown. Maxillary palpus fuscous, orange-tipped. Basal scaling of proboscis black. Neck
region orange. Thorax above: patagium and tegula orange, each with a black anterior spot. Abdomen
black above, terminal segment orange. Thorax, abdomen below: black; a thin band of orange on pectus;
terminal segment orange. Legs black, grey-banded. Fore wing above: ground colour black with faint
purplish lustre; a small orange basal spot on Cu; a narrow, arcuate, creamy white subbasal fascia. Ante-
medial fascia broad, weakly oblique medially, creamy white; commencing narrowly behind costa, rapidly
widening and extending broadly to posterior margin. A creamy spot on discocellular vein near posterior
angle of cell. Longitudinal veins in terminal half of wing outlined in creamy yellow. A fairly wide,
triangular, creamy yellow subapical patch, bordered by a narrow black marginal line. Fringe fuscous.
Hind wing and fringe black with purple reflections, unmarked. Fore wing below: fuscous with purple
reflections; subapical patch as on upperside; a diffuse, pale proximal area and pale posterior margin.
Hind wing below as on upperside.
GENITALIA @ (Figs 225, 271). In general as described for genus. Posterior apophysis weakly sinuate.
Anterior apophysis angled near base. Eighth tergite with one irregular, major, transverse, posterior row
of setae, numerous setae scattered over remaining area. Eighth sternite subtriangular, wider than long,
posteriorly truncated. Sclerotized antrum with narrow diverticulum and long subacute process.
REMARKS. This species resembles C. minor; differential characters are noted under that species.
The development of the antrum is the most distinctive genital character.
BioLoGcy. Early stages and host-plant unknown. Holotype collected in March.
DISTRIBUTION (Fig. 3). New Guinea: Papua New Guinea.
MATERIAL EXAMINED
Holotype 2, New Guinea: Mambare R., Biagi, 1500 m (‘5000 ft’), iii.1906 (Meek) (genitalia slide no.
14698, BMNH).
258 E. G. MUNROE & M. SHAFFER
VITESSA Moore, [1860]
Vitessa Moore, [1860,] in Horsfield & Moore, Cat. lepid. Insects Mus. nat. Hist. East-India House 2 : 299.
Type-species: Vitessa suradeva Moore, [1860], by original designation and monotypy.
[Cosmethis Hiibner; sensu de Joannis, 1930 : 637. Erroneous type-species designation.]
Head large. Frons rounded, narrow, its scaling rough or smooth. Vertex short, with erect, compact,
woolly scaling. Eye large. Ocellus usually present, inconspicuous, sometimes weak or absent. Chaetosema
elliptical, adjacent to eye, directed laterally, with long radiating setae. Antenna variable in length, more
than half, less than three-quarters length of fore wing; bipectinate in 3, compressed, laminate in 9,
postaxial pectinations in ¢ usually considerably longer than preaxial ones; pectinations flat, spatulate,
capable of folding like slats of a Venetian blind. Labial palpus long, more or less equal in the sexes,
obliquely or fully upturned, exceeding level of vertex; first segment very short; second segment long,
cylindrical, slightly curved, not reaching level of vertex; third segment more slender, cylindrical, acum-
inate, varying from two-thirds to fully as long as second segment..Maxillary palpus short, distally
somewhat dilated with scales, mesally directed. Proboscis long, strong, basally widened and scaled. Body
robust, abdomen exceeding hind wing. Legs fairly long, robust; spurs long, outer somewhat shorter than
inner; tarsi ventrally spined. Fore wing long, narrow, usually two and a half to three times as long as
wide; in ° a little wider and with apex more rounded than in ¢; costa straight, gently arched or weakly
sinuate; apex rounded; termen generally weakly rounded and a little oblique; tornus oblique or rounded;
posterior margin nearly straight, but convex near base. Costal area wide. Cell more than half length of
wing. Anterior discocellular vein short, at a distinct angle to middle discocellular. Sc separated from radial
stem and R,. R, arising from cell. R, from before end of cell. R; ,, stalked with R; for a short distance;
R; and R, with a short stalk beyond separation of R;. M, arising at junction of anterior and middle
discocellular veins, obliquely divergent from R; at base. M, and M; short-stalked from posterior angle of
cell. Cu, from a little behind posterior angle. Cu, variable in position, always from well out on cell. Anal
fold strong. Terminal part of CuP developed or absent. A, long and strong, connected with A, by an
outwardly oblique cross-vein, or A, looped to A,. Hind wing long and narrow; apex narrowly rounded,
termen broadly rounded, anal angle rounded. Costal area wide. Cell about half length of wing. Dis-
cocellular vein strongly angled or incurved at middle; anterior and posterior angles of cell acute, posterior
angle projecting further distad than anterior angle. Sc+ R,; long and straight, reaching apex of wing. Rs
weaker than Sc+ R,, paralleling it closely to beyond middle of wing, but not anastomosing with it,
diverging distally. M, stalked with Rs. M, and M; from posterior angle of cell, basally approximated or
short-stalked. Cu, from cell somewhat behind posterior angle; Cu, from cell at two-thirds from base.
CuP, A, and A, present. Frenulum of 2 with two long setae.
GENITALIA ¢ (Figs 25-41, 43, 44, 137-218). Uncus slender: in griseata- and hieratica-groups shorter and
pyriform, bearing anteriorly directed setae on each side; in other groups with distal part slender, rod-
shaped, with sparse, laterally directed setae on each side. Gnathos wide, with broad, flat, posteriorly
curved arms, acutely angled mesally, joined to form a long, strong, tapering and acuminate process,
broadly curved dorsad at some point in its length. Transtilla strong, of even width, V-shaped or arcuate,
often with a more heavily sclerotized dorsal bar. Tegumen wide, with pronounced shoulders; its anterior
margin with a narrow, V-shaped, medially deeply emarginated sclerite and its posterior margin with a
broad semicircular or subtriangular sclerite. Vinculum narrow, ventrally flattened, with a pair of small
accessory sclerites. Valve variable in form; costa strengthened; mesal surface strongly setose; usually
with a low ridge-like clasper of varying form, absent in most of zemire-group. Juxta well developed, divided
into a pair of lateral sclerites, each with dorsal and ventral processes directed medially, dorsal process
articulating with lateral sclerite of anellus; this sclerite of varying form, generally linguiform and spinose,
directed anteromedially when retracted (Figs 25-32). Dorsal medial catena spines on anellus large, small
or absent (Figs 33-40). Manica long, tapering; medially directed sclerotized bands supporting part of
length. Lateral sclerite of anellus hinging on articulation with juxta, capable of reflexion or extension
as aedeagus is retracted or protruded (Fig. 33). Extension or reflexion of lateral sclerite independent of
eversion or inversion of anellus in dissected genitalia with acdeagus removed. Aedeagus long, narrow,
tubular, expanded distally, narrowly rounded basally; ductus ejaculatorius arising subbasally; aedeagus
rugose at ventral connection to vesica, usually heavily spinose; rugose area longitudinal or transverse,
apposed on dorsal connection by longitudinal rugose tissue or longitudinal bands of sclerotization.
Junction of aedeagus with vesica at rugose area capable of reflexion, both before and after eversion
(Fig. 41). Vesica long and tubular, with variable area and number of minute spines dorsally and ventrally
at base. Cornuti present, deciduous to some extent, grouped in clusters or long bands of varying size, in
various combinations: two larger dorsolateral clusters and one smaller ventromedial cluster, or, two
large dorsolateral clusters, sometimes with one either smaller or absent.
Figs 25-32 ¢ genitalia. Details of left juxta sclerite and its connection with lateral sclerite of anellus
in representative species of Vitessa to show variation within the genus. (25) V. griseata Kenrick.
(26) V. zemire nephritica subsp. n. (27) V. hieratica Swinhoe. (28) V. stettina Swinhoe. (29) V.
hemiallactis hemiallactis Meyrick. (30) V. pyraliata latialbata subsp. n. (31) V. plumosa plumosa
Hampson. (32) V. suradeva rama Moore.
E. G. MUNROE & M. SHAFFER
TaN
> \
peed S92
lateral
sclerite
anellus
Figs 33-40 ¢ genitalia. Details of dorsal catena of anellus in representative species of Vitessa to
show variation within the genus. Fig. 33 also shows the juxta, lateral sclerites and everted, reflexed
anellus to illustrate the position of the catena in relation to these parts. (33) V. griseata Kenrick.
(34) V. cyanea cyanea Hampson. (35) V. hieratica Swinhoe. (36) V. stettina Swinhoe. (37) V.
hemiallactis admiralitatis subsp. n. (38) V. pyraliata latialbata subsp. n. (39) V. plumosa plumosa
Hampson. (40) V. suradeva suradeva Moore.
REVISION OF VITESSIDIA & VITESSA 261
Fig. 41 3 genitalia. Lateral view of aedeagus illustrating everted vesica in reflexed position.
Vitessa pyraliata pyraliata Walker.
GENITALIA ° (Figs 226-268, 272-314, 316-318). Ovipositor lobes long, fairly wide, dorsally fused and
produced posteriad; posterior surface setose, margins with a moderately thick fringe of longer setae;
longer setae and some of the smaller, round, tubular and simple, a varying number of smaller setae
flattened and bearing spines for most of their length. Posterior apophysis curved, of moderate thickness,
mostly blunt-tipped. Anterior apophysis longer, thicker, weakly sinuate, mostly blunt-tipped. Eighth
tergite broad, indented posteromedially, bluntly protruded anteromedially; densely setose posteriorly
and laterally. Ostium wide. Posterior ostial sclerite with posterior margin curved; anterior margin with
lateral prolongations of varying length and shape, sometimes expanded toward mid-line to form large
bullae. Anterior ostial sclerite with posterior margin recurved, wrinkled. Antrum wide, variably extended
anteriorly, posterior part sometimes weak. Ostial end of ductus bursae mostly funnel-shaped, wrinkled,
with weak lateral sclerotizations; continuing as a slender membranous tute to corpus bursae. Corpus
bursae globular or oval, membranous; with two depressed, finely denticulate signa, with paired denti-
culate areas, or without signa.
REMARKS. We recognize eight species-groups. In the griseata-group the valve has a small, central,
almost vestigial clasper; the uncus is pyriform-subtriangular, as in Cosmethella; the hind wing, as
in Cosmethella, is black. In the hieratica-group the uncus is similar in shape but the valve is armed
on the posterior margin with a small, pointed clasper; the discal area of the hind wing is white.
In the remaining six groups the uncus has a wide basal portion and a rod-like distal portion, the
latter decurved, fitting closely into the curve of the gnathos and bearing sparse lateral setae. In
the suradeva-group the costa of the valve is curved at the base, enabling the valve to open
horizontally rather than obliquely; in the remaining groups the costa of the valve is straight
basally, as in the griseata-group. The zemire-group has black hind wings and a fluted valve,
mostly with no clasper; the condition of the uncus is transitional from that in the griseata- and
hieratica-groups to that of the remaining five groups (including the suradeva-group, above),
the basal portion being less sharply differentiated from the rod-like distal portion than in the
latter series of groups. The stettina-group has a weak clasper on posterior margin of the narrow
valve and in the 2 the antrum is weakly sclerotized. In the hemiallactis-group the valve is wide
262 E. G. MUNROE & M. SHAFFER
and basally constricted, the clasper is long, serrate and well developed and in the 9 the ostial
sclerotization is strong. In the p/umosa-group the clasper of the valve has a strong, truncate and
bifid-tipped ventral process, the ostial chamber has a complex folded sclerite on each side. In the
pyraliata-group the clasper of the valve is coarsely serrate and the margins of the ostial chamber
are wide and heavily sclerotized. Within the species-groups, spined setae on the ovipositor lobes
are most easily seen in the pyraliata-group and least easily seen in the griseata- and zemire-groups.
To be distinguished easily the spined setae require study under a compound microscope; the
examples shown in Figs 316-318 were photographed at about x 250 magnification by the use
of differentiated interference-contrast microscopy.
BrioLoGy. Available information on the life-history is given in detail below, under Vitessa suradeva
rama. The larvae of the species are gregarious web-makers on the undersides of leaves of Chailletia
sumatrana Miquel (= Dichapetalum gelanioides subsp. gelanioides (Beddome) Engler) (Dicha-
petalaceae). They are gregarious when young, solitary later, commonly sitting in a ‘semilooper’
position. They are long, oily-black, marked with white, with orange head, anal segment and
prolegs. Their bright colours may be aposematic. The pupa is formed in a cocoon just under the
ground against sides of stones or in a similar protected place. V. suradeva ordinarily flies in the
daytime and visits flowers in company with hypsids. Specimens of V. hemiallactis hemiallactis
from Waigeo, Camp Nok, collected by Cheesman, are labelled as being diurnal. However, the
adults of at least some of the species, including V. suradeva, griseata, zemire, hollandi and
hemiallactis, are active nocturnally, though the bright colours might seem better adapted to
diurnal life. The general aspect is that of aposematic or mimetic insects (see Introduction, p. 242).
This is supported by the presence of noctuoids of closely similar appearance in the same areas,
and by the similar patterns and parallel geographical variation of the forewings in species of
different groups, e.g. the greenish V. griseata and V. barretti in the highlands of New Guinea, the
black and white V. zemire, V. stettina and V. hemiallactis at lower altitudes in New Guinea, the
greenish V. glaucoptera and V. zemire nephritica in Queensland, and the dark, white-spotted V.
pyraliata triangulifera and V. philippina in Luzon.
DISTRIBUTION. Northern India and Sri Lanka to Queensland and Samoa.
BIOGEOGRAPHY (Figs 2-24). The griseata-, zemire-, stettina- and hemiallactis-groups are mainly
Papuan; all extend west into the Moluccas and the stettina-group reaches the Philippines. The
hemiallactis- and zemire-groups extend south to Queensland and south-east into the Solomon
Islands. The zemire-group ranges farther, to the New Hebrides, Fiji and Samoa. The hieratica-,
plumosa-, and pyraliata-groups are centred on the Malayan subregion; the hieratica-group
ranges from Sumatra to Borneo, the p/umosa-group through the Lesser Sunda Islands and north
to Salajar, and the pyraliata-group from the Nicobar Islands and possibly Sri Lanka to the
Philippines, Flores and possibly northern Australia. The suradeva-group is Indo-Malayan,
ranging from Sri Lanka and the Himalayas to Borneo.
Turning to distributions within the groups, the griseata-group has four known species. V.
griseata is widely distributed in New Guinea at altitudes from about 450 to 1850 m. This species
is replaced by the closely related V. brandti in the mountains of New Britain. The remaining
two species of the group, V. talboti and V. nobilis, occur in Seram and Ambon respectively;
the former at least appears to be a mountain species.
The zemire-group consists of four species. The polytypic V. zemire extends from the Moluccas
to the New Hebrides. V. vitialis in Fiji and V. tamsi in Samoa form a closely related pair. V.
cyanea, the fourth species of the group, overlaps V. zemire in eastern New Guinea and adjacent
coastal islands and has a widely disjunct subspecies in the Moluccas. The subspecies of V. zemire
form regional groupings. In the New Guinea subspecies the postmedial fascia of the fore wing is
very wide; in the Queensland subspecies the fascia is strongly green-tinted; it is equally wide, but
tends to diffuse across the medial region to meet similar diffusion from the antemedial fascia.
The two Moluccan and one New Hebrides subspecies have somewhat narrower postmedial
fasciae; the remaining two subspecies from the Bismarck and Solomon Islands have much
narrower postmedial fasciae.
REVISION OF VITESSIDIA & VITESSA 263
The single known species of the hieratica-group is rare in collections. So far it is known from
Borneo and Sumatra.
The three species of the stettina-group are largely allopatric. V. stettina, a species with wide
pale fasciae on the fore wing, ranges from Waigeo and the Arfak Mountains to eastern Papua.
It is the dominant species in southern New Guinea, but its range is widely overlapped by that of
V. hollandi, a species with restricted pale fasciae. V. hollandi occurs in the Schouten Islands
sympatrically with the externally almost indistinguishable V. gemina of the pyraliata-group. A
distinct subspecies of V. hollandi occurs in Sulawesi; two specimens are known from the islands
in the Moluccan group and a single specimen from New Britain; though these probably represent
distinct subspecies we do not name them here. In the Philippines the stettina-group is represented
by a distinct species, V. philippina, which is dark, with reduced light spots; externally this species
closely resembles the sympatric V. pyraliata triangulifera, of the pyraliata-group.
The hemiallactis-group has six species. Of these the polytypic V. hemiallactis is the most widely
distributed, with one subspecies in Halmahera and Obi in the Moluccas, a second in New Guinea,
a third in the Admiralty Islands, a fourth in the Bismarcks and a fifth in the Solomon Islands.
The fore wing patterns parallel those of sympatric members of the stettina-group. The New Guinea
subspecies has moderately narrow, at most weakly interrupted, white fasciae, almost identical
with those of V. hollandi hollandi, whereas in the remaining subspecies the fasciae are very much
reduced, being both narrower and more widely interrupted. Overlapping V. hemiallactis
hemiallactis in geographical range, though mostly occurring at somewhat higher altitudes, is the
slightly larger second species, V. intermedia. The wide greenish white fasciae of this species parallel
the markings of V. stettina, which occurs both with V. intermedia and at lower altitudes. In
Queensland the geographically representative species V. glaucoptera has the fore wing green-
suffused, like that of V. zemire nephritica from the same area. At altitudes of 600 to about
1850 m and possibly higher in New Guinea there occurs another representative species with
strongly greenish fore wings, namely V. barretti; this species superficially resembles the more
abundant V. griseata, which occurs together with it. A fifth species, V. ternatica, is known from
the Moluccas from Ternate to Buru and Seram. A sixth, V. cristobalensis, is known only from
San Cristobal in the Solomon Islands. A seventh species is known to us from a single 3 specimen
which most closely resembles V. ternatica; however, its locality, given on the data label as ‘Japan’,
is regarded as erroneous and the species is described without being named in this paper.
The pyraliata-group includes seven species. The most widely distributed is V. pyraliata, which
ranges from Sri Lanka to the Philippines, Sulawesi and Flores, in three subspecies. V. pyraliata
latialbata has the widest range. It occurs in the Malay Peninsula, Sumatra, Java, Borneo and part
of Sulawesi; there are dubious records from Sri Lanka and Australia. The rather similar V.
pyraliata triangulifera replaces it in the Philippines. The more divergent nominate subspecies is
found in part of Sulawesi and in Flores. The remaining species of the group have restricted ranges.
V. nicobarica is restricted to the Nicobar Islands. Four vicarious species, V. muluana, kolakalis,
sulaensis and gemina, occur in Sarawak, Sulawesi, Sula Besi and the Schouten Islands, Geelvink
Bay, respectively. V. splendida lives sympatrically with V. pyraliata triangulifera in the Philippines.
The single species of the plumosa-group is geographically variable. It extends through the Lesser
Sunda Islands from just west of Wallace’s Line in Bali to Dammer Island; a distinct subspecies is
found in Salajar, off the south coast of Sulawesi. Although we include all the populations from the
Lesser Sunda Islands provisionally in the nominate subspecies, the limited material we have seen
suggests that there may be geographical variation; specimens from the middle of the chain in
Lombok and Sumbawa are large and dull-coloured, whereas those from Bali in the west and
from the eastern islands of the group are smaller and brighter. More material is needed before
these differences can be evaluated.
The suradeva-group has two species. V. teleroma ranges from Perak to Borneo; there is some
variation in this species which more material may show to be geographically based. V. suradeva
has two subspecies. One ranges from W. Bengal and Assam to Burma, Thailand and Vietnam, the
other is found in southern India and in Sri Lanka.
There appear to be two possible interpretations of the geographical distribution of the Vitessa
complex. On the one hand, the concentration of species and species-groups in the Papuan region
264 E. G. MUNROE & M. SHAFFER
may be taken as evidence that the complex has originated and undergone most of its evolution
there, and that it has spread secondarily into the Indo-Malayan region. Alternatively, it may be
supposed that the groups now confined to the Papuan region were previously widespread, but
that they have been displaced in the west by the species that now exist there. On either hypothesis
it must be assumed that there has been active local differentiation in most of the surviving species-
groups.
NOMENCLATURE. The genus has mostly been cited as Vitessa Moore. However, the name was
changed to Cosmethis Hiibner, [1820] in the BMNH collection many years ago, presumably by
Hampson when he recognized the validity of Hiibner’s generic names, though Hampson (1896a :
145; 18966 : 502) had earlier used Vitessa in his published work on the genus. Cosmethis was used
by Janse (1924; 1928) and by de Joannis (1930). The latter author designated Phalaena zemire
Stoll as type-species of Cosmethis. However, this designation is antedated by that by Kirby
(1892 : 498) of Phalaena barbara Stoll, 1781, now referred to the Geometridae. The name
Cosmethis therefore cannot be applied to the present genus and the more familiar name Vitessa
must be used.
Key to the species and subspecies of Vitessa
1 Hind wing upperside uniformly black, fringe white . ; : : 2
Hind wing whitish buff, with or without broad black or fuscous ‘terminal band , 17
2(1) Fore wing upperside greenish grey, with orange basal area outwardly bordered by fine
blue-black subbasal fascia. New Guinea : : : : : griseata (p. 267)
Fore wing with fascia in medial area , ; ; F 3
3(2) Fore wing upperside with broad violet-grey medial fascia. New Britain . brandti (p. 269)
- Fore wing with medial fascia other than violet-grey . ; ‘ : 4
4(3) Fore wing upperside with large, square white medial fascia. Seram : é talbott C 269)
Fore wing with more than one white marking in medial area. 5
5 (4) Fore wing upperside with narrow white medial fascia divided into two spots. Ambon
nobilis (p. 270)
Fore wing with more than one fascia : 6
6(5) Fore wing upperside with two white transverse fasciae, ‘sometimes joined by medial
diffusion; basal area orange ; 7)
Fore wing with two pale yellow transverse fasciae, always separated: subbasal area orange 16
7(6) Fore wing upperside with postmedial fascia joining white terminal lines (cyanea sub-
species) ; A 8
Fore wing with postmedial fascia not soinine white terminal lines emis subspecies) ; 9
8(7) Fore wing upperside with orange basal area not passing Cu; ¢ genitalia with termen of
valve narrowly rounded. Seram . , . cyanea ceramensis (p. 273)
- Fore wing with orange basal area indented at Ci. but reaching posterior margin; ¢
genitalia with termen of valve truncated. Papua New Guinea and Fergusson Island
cyanea cyanea (p. 273)
9(7) Fore wing upperside with wide creamy white postmedial fascia, width nearly equal to half
width of wing at centre; antemedial fascia narrower than subbasal fascia; basal area
complete. Northern Moluccas . ; ; ? : : zemire expansa (p. 275)
Fore wing with markings not as above . 10
10(9) Fore wing upperside with wide chalky white postmedial fascia, width equal to half width
of wing at centre; antemedial fascia narrower than subbasal fascia; basal area indented;
hind wing underside with white apical area. Southern Moluccas . Zzemire zemire (p. 276)
Fore wing with markings not as above . 1]
11 (10) Fore wing upperside with antemedial fascia diffused medially but not joining postmedial
fascia, as wide as subbasal fascia; width of postmedial fascia more than half width of
wing at centre. New Guinea and coastal islands. : zemire temerata (p. 276)
Fore wing with medial diffusion of fasciae joining, or fasciae well separated : : 12
12 (11) Fore wing upperside with antemedial fascia irregularly diffused medially, meeting diffusion
from postmedial fascia, especially along posterior margin. Queensland
zemire nephritica (p. 278)
- Fore wing with fasciae well separated, without such diffusion . : : : ‘ 13
13 (12)
14 (13)
15 (14)
16 (6)
17 (1)
18 (17)
19 (18)
20 (19)
21 (20)
22 (18)
23:(22)
24 (23)
25 (24)
26 (25)
REVISION OF VITESSIDIA & VITESSA 265
Fore wing upperside with orange basal area bearing subcostal black dot half width of eye;
postmedial fascia twice as wide as antemedial fascia in 3, antemedial fascia in 2 reduced
to hair-line. Umboi Island . : : : , ; ; . zemire coronis (p. 278)
Fore wing with markings not as above . : 14
Fore wing upperside with subcostal black dot half width of eye or much less; both white
fasciae reduced in width, postmedial fascia one-third width of wing at centre. New
Britain and New Hannover : : zemire novaebritanniae (p. 279)
Fore wing similar to above; hind wing underside lacking white apical area but having
white spot at apex of cell. ‘ 15
Fore wing upperside with orange basal area indented at ve 1 with blue-black; postmedial
fascia narrow, less than half width of wing at centre. Solomon Islands
zemire solomonis (p. 280)
Fore wing with postmedial fascia broad, equal to half width of wing at centre. New
Hebrides . , . zemire cheesmanae (p. 280)
Fore wing upperside witli postmedial eee directed foward posterior margin well basad
of tornus, narrow, width less than half width of wing at centre. Fiji F vitialis (p. 281)
Fore wing with postmedial fascia directed towards tornus, broad, width equal to half
width of wing at centre. Samoa . : j ‘ tamsi (p. 283)
Hind wing upperside with infuscation from costa reaching CuP or farther, eliminating
white from cell; fore wing upperside with orange subbasal area reaching posterior
margin. Borneo ud Sumatra , : : : : . heiratica (p. 285)
Wings not with this combination of characters ; : ” 18
3 genitalia with anellus lacking catena spines; 9 genitalia with corpus bursae lacking
signum or small denticulate area ‘ 19
3 genitalia with anellus bearing catena spines; 2 genitalia with corpus bursae bearing signa
or small denticulate areas . : 22
Fore wing above with about six white terminal lines; Q genitalia with funnel- shaped
antrum (¢ unknown). Philippines ; : Philippina (p. 286)
Fore wing with more than six white terminal lines: 9 genitalia with cup-shaped
antrum. 20
Fore wing upperside with postmedial and antemedial fasciae broad and complete. New
Guinea. Stettina (p. 287)
Fore wing upperside with both fasciae narrow, ‘incomplete or indented : : . zl
Fore wing underside with postmedial fascia divided into two spots. Sulawesi
hollandi wallacealis (p. 289)
Fore wing with postmedial fascia a single large oval spot. New Guinea
hollandi hollandi (p. 290)
3 genitalia with clasper on valve long, strap-shaped and serrulate along part or whole of
free edge; 2 genitalia with ductus seminalis arising below or just above base of antrum 23
3 genitalia with clasper short, mostly parallel to posterior margin, free edge with small
ridge and rounded scoop-like process; 2 genitalia with ductus seminalis ane suis
up swollen extended base of antrum (suradeva-group) . : : 42
3 genitalia with clasper with large ridges or tooth-like points at posterior margin with
pronounced acuminate ridges mid-way; 2 genitalia with posterior portion of antrum
trapezoidal, or broadly extended and cup-shaped anteriorly . : : : : 24
3 genitalia with clasper with pronounced truncate process mid-way; 2 genitalia with
antrum broadly rounded posteriorly, acutely Sai eis with V-shaped
dorsal crease (plumosa-group) . : F 41
Fore wing upperside with antemedial fascia eudinsede to minute ae on costa aa iatsed
narrow triangular spot on Cu pointing medially; hind wing upperside with white area
reduced to a comma-shape directed basally. Moluccas . ; é ternatica (p. 292)
Combined fore wing and hind wing markings not as above : : - 25
Fore wing upperside with postmedial fascia divided into two spots Sail jenmiinal margins
at right-angles to posterior margin of wing. Moluccas . hemiallactis moluccana (p. 295)
Fore wing with postmedial fascia divided into two spots or complete but terminal margins
oblique to posterior margin : 26
Fore wing upperside with postmedial and setemedal Pacis indented or e CORIBIEtE. fascia
approaching but not joining at any point. New Guinea . _—_hemiallactis hemiallactis (p. 296)
266
27 (26)
28 (27)
29:28)
30 (29)
31 (30)
32 (31)
33 (32)
34 (33)
35 (34)
36 (35)
37 (36)
38 (37)
39 (38)
40 (39)
41 (23)
E. G. MUNROE & M. SHAFFER
Fore wing with postmedial and sometimes antemedial fascia distinctly divided into two or
more spots, or fasciae complete and partially fusing
Fore wing upperside with antemedial fascia narrowing almost toa - hair-line but not
divided; postmedial fascia with two spots; hind wing upperside with broad black
terminal band. Admiralty Islands : A : . hemiallactis admiralitatis (p.
Fore wing with antemedial fascia narrow, or broad, or divided into two spots; postmedial
fascia divided into two or more spots, or complete; hind wing upperside wholly creamy
white or with black terminal band :
Hind wing upperside with large oval white area not reaching anal margin; fore wing
upperside with terminal white lines well apart from postmedial fascia. Bismarck
Archipelago , F . hemiallactis lustrans (p.
Hind wing with white area reaching anal margin Teeny where white almost obliterated by
infuscation encroaching from costa; fore wing with terminal white lines separate from
or joining postmedial fascia
Fore wing upperside with fasciae partially fusing leaving variable amounts of blue- black
medial fascia exposed; orange subbasal area reduced leaving blue-black subbasal fascia
broad at costa. Papua New Guinea : intermedia (p.
Fore wing with fasciae more completely fda or Bifte coed and divided into spots
Fore wing upperside with fasciae almost completely fused leaving medial fascia as a
longitudinal short streak; expanse of fore wings 60 mm or over. Papua New Guinea
barretti (p.
Fore wing with fasciae similar, but fore wing expanse less, or fasciae separated
Fore wing upperside with fasciae almost completely fused leaving medial fascia as a short
triangular spot; expanse of fore wings 50 mm or less. Queensland . glaucoptera (p.
Fore wing with fasciae separated and divided into spots :
Fore wing underside with white terminal lines present from R, to Pies ean Cowal
Island : ; . cristobalensis (p.
Fore wing with HS feamial fines absent or from R, to Cu
Hind wing upperside of 3 unicolorous whitish buff, of 2 mainly eiatele buff, par narrow
and ill-defined black costal and terminal borders. Nicobar Islands . nicobarica (p.
Hind wing of ¢ and 2 with broad well-defined black costal and terminal borders
Hind wing upperside with white extending along over half of anal margin; fore wing
upperside with lower spot of antemedial fascia distinctly rectangular. Malaysia
pyraliata latialbata Me
Hind and fore wing markings other than in above combination .
Fore wing upperside with orange subbasal area restricted between Sc and Wie : hind wing
upperside with white extending along half of anal margin. Philippines
pyraliata ricci iss (p.
Fore and hind wing markings other than in above combination
Hind wing upperside with white variably extended and restricted along anal margin, ea
a third to complete restriction; fore wing upperside with orange subbasal area from just
above Sc to A, .2.; patagium mostly black. Sulawesi ; . pyraliata pyraliata (p.
Hind and fore wing markings and colour of patagium other than in above combination
Hind wing upperside with white variably extended along anal margin from half to about a
third; fore wing upperside with orange subbasal area to within hair-line of costa;
patagium mostly orange. Sulawesi : : . kolakalis (p.
Hind wing with white extending along half or more of anal area
Fore wing upperside with orange subbasal area not passing A, ,.3; white terminal vein- lines
very short. Moluccas . ; ; . sulaensis (p.
Fore wing with subbasal area nearly reaching posterior margin , ,
2 genitalia with corpus bursae with two round, depressed denticulate signa (3 unknown).
Sarawak, Sabah : : : ; . muluana (p.
2 genitalia with corpus bursae with minute denticulate patches d :
9 genitalia with antrum funnel-shaped (¢ unknown). Schouten Islands, Geelvink Bay
gemina (p.
2 genitalia with antrum deep and cup-shaped (¢ unknown). Philippines . Splendida (p.
Hind wing upperside with black border along termen narrowing abruptly before reaching
tornus. Sumbawa ; . P 4 : 3 : : plumosa plumosa (p.
27
297)
28
298)
29
299)
30
300)
31
301)
32
302)
eK)
303)
34
306)
5D
308)
36
308)
OM
309)
38
310)
39
311)
40
312)
313)
316)
REVISION OF VITESSIDIA & VITESSA 267
~ Hind wing with black border narrowing evenly from costa to tornus. Salajar
plumosa salayerensis (p. 317)
42 (22) Fore wing upperside with antemedial fascia finely parted at cell. Malaysia . teleroma(p. 317)
- Fore wing with antemedial fascia indented but complete . ; : 43
43 (42) 3 genitalia with aedeagus with right cluster of cornuti about half as long as left: 2 genitalia
with antrum nearly as wide as long. Northern India to Vietnam _suradeva suradeva (p. 320)
~ 3 genitalia with right cluster of cornuti noticeably smaller, about one-third as long as left;
2 genitalia with antrum much narrower than long. Southern India and Sri Lanka
suradeva rama (p. 321)
The griseata-group
DiAGnosis. Fore wing above: mostly greenish, with or without a broad, diffuse, violet-grey transverse
medial fascia but without a distinct black wedge-shaped discocellular dot or dash or narrow transverse
medial fascia as in greenish-winged species of the zemire- and hemiallactis-groups; or fore wing above:
largely black, with complete or interrupted whitish transverse medial fascia, without postmedial and
antemedial fasciae as in other species-groups. Hind wing black, as in zemire-group. Genitalia in ¢ with
uncus narrowly pyriform, a short zone of oblique, close-set setae dorsolaterally, as in the hieratica-group,
but with setae less differentiated; valve falcate, with small, central, almost vestigial clasper, not ridge-like
or projecting marginal clasper as in other species-groups; anellus with large lateral clusters of spines as
in zemire-group. Genitalia in 2 with weakly developed spined setae on ovipositor lobes; posterior ostial
sclerite bearing two lateral bullae; anterior margin of anterior sclerite strongly wrinkled; antrum
sclerotization short; anterior end of ductus bursae membranous, broadly funnel-shaped; corpus bursae
with two round, depressed, denticulate signa.
Vitessa griseata Kenrick, 1907
(Figs 4,25, 33;552553, 138,173, 204, 226; 316)
Vitessa griseata Kenrick, 1907, Proc. zool. Soc. Lond. 1907 : 75, pl. 3, fig. 45. LECTOTYPE 3, New
GUINEA: Papua, Dinawa, 4000 ft [1200 m], ix.1902 (A. E. Pratt) (BMNH), here designated [examined].
Cosmethis griseata (Kenrick); Janse, 1928 : 88.
3, 2. 44-68 mm. Frons pale orange. Vertex bright orange. Eye dark brown. Ocellus weak, not conspicuous,
dark brown. Chaetosema with pale yellowish setae. Antenna dark fuscous, anterior surface of scape
and basal segment orange; posterior surface of apical third of shaft white. Labial palpus blackish fuscous;
orange on ventral surfaces of first and second segments, minute amount at base of third. Maxillary palpus
orange. Basal scaling of proboscis black, admixed with orange at base. Neck region orange. Thorax
above: orange; patagium and tegula orange, each with blue-black spot at centre. Abdomen above: black;
narrow whitish grey posterior line on each segment, anal tuft orange. Body below: black with bluish
reflections; pectus with considerable orange scaling; whitish grey abdominal lines wider than above; anal
tuft orange. Legs blackish fuscous, with extensive pale grey scaling on coxae, femora and tibiae. Fore wing
above: basal area orange bearing two black spots on R, some black posteriorly; oblique subbasal fascia
a blue-black line. Remainder of wing silky greenish grey, with fine black terminal lines, mostly between
veins, from R to Cu, variable in number. Fringe anteriorly black, posteriorly pale grey. Hind wing above:
black with some metallic lustre. Fringe: base grey, terminally whitish grey, anal margin black. Fore wing
below: black, slightly metallic; an oblique, oblong, cream-coloured postmedial fascia, almost reaching
posterior margin; apex with diffuse longitudinal greenish grey lines; fringe dark grey anteriorly, whitish
grey posteriorly; retinaculum dark grey. Hind wing below: similar to fore wing but with fringe uniformly
whitish grey.
GENITALIA ¢ (Figs 25, 33, 138, 173, 204). Uncus relatively short and wide, base constricted, basal part
laterally convex with ventral foramen; distal part flattened, curved, acuminate, with a dorsolateral zone
of oblique setae. Gnathos with flattened, medially decurved, wide lateral arms and with compressed,
basally straight, distally upturned and acuminate, medial process. Transtilla shallowly V-shaped, of
even width. Tegumen with medially broadened posterior sclerite and with anterior sclerite widely
emarginated medially. Vinculum ventrally narrow and flattened. Valve long and narrow, distally falcate,
extreme tip very narrowly rounded and terminal margin excised before it; costa strongly inflated; mesal
surface of valve heavily setose in distal two-thirds of costal region only; clasper a small erect three-sided
ridge at axis of valve, less than half valve length from base. Juxta with nearly rectangular lateral sclerites,
268 E. G. MUNROE & M. SHAFFER
each bearing a dorsal, somewhat decurved, thorn-shaped, thinly setose process and ventral, slender,
ribbon-shaped process, ventral process directed posteriorly and nearly reaching mid-line, supporting base
of anellus; anellus (Figs 33, 173) arising from juxta processes with large lateral, medially directed sclerite
bearing large anteriorly directed spines, sclerite connected directly to dorsal process (Fig. 25); ventral
surface of anellus between the sclerites longitudinally rugose, lightly sclerotized and centrally spinose;
dorsal surface with medial catena (Fig. 33) consisting of an elongated, ovoid area, minutely spinose; most
of manica supported by dorsal sclerotized caliper-shaped bands, narrowing and less sclerotized towards
tip. Aedeagus of moderate length, expanded distally, slightly curved, base narrowly rounded; ductus
ejaculatorius arising basad of middle; scale-like spines extending medially for a short distance from vesica
along dorsal and ventral surfaces of aedeagus; base of vesica bulbous dorsally at posterior end of aedeagus;
dorsal and ventral surfaces covered with scale-like spines from base of vesica to about half-way along
larger cornuti clusters, the spinose area longitudinally rugose between cornuti clusters; vesica bulbous
immediately distad of this region. Cornuti (Fig. 204) grouped in three clusters of spines: two large, more
basal, dorsolateral clusters of unequal size, the left one longer and extending farther distad than the right,
and a much smaller, distal, mid-ventral cluster, bearing thinner spines and arising from distal end of an
oblique sclerotized band.
GENITALIA 2 (Figs 226, 272, 316). High, wide ovipositor lobes, dorsally fused and produced posteriorly;
surface densely setose, margin with row of well-separated longer setae; spiny setae not easily seen (Fig.
316). Posterior apophysis fairly long, slender, weakly curved, with spatulate tip. Anterior apophysis
similar but longer. Posterior ostial sclerite (Fig. 272) broadly sclerotized, posterior margin angularly
excised, anteromedial part broadly divided on mid-line by a narrow sulcus, laterally a pair of wrinkled
bullae with sinuate, posteriorly converging inner margins, posteriorly denticulate. Anterior sclerite
narrow, wrinkled, reflected, heavily denticulate. Antrum short, collar-shaped. Ductus bursae membranous,
posteriorly widened and flattened, lightly sclerotized. Corpus bursae globular, membranous, with a pair
of large, heavily sclerotized, depressed, denticulate signa.
REMARKS. This species is fairly common at most altitudes in many parts of New Guinea. The
only sympatric species with which it might be confused is V. barretti, which can be separated by
the black discocellular spot of its fore wing and the large white area on the hind wing. The closest
relative of V. griseata is V. brandti from New Britain, which has a contrasting violet-grey trans-
verse medial fascia on the upperside of the fore wing.
BrioLoGy. Early stages and host-plant unknown. Moths collected by many collectors in most
months of the year.
DISTRIBUTION (Fig. 4). New Guinea: Irian Jaya; Papua New Guinea, 500-2150 m.
MATERIAL EXAMINED
Paralectotypes. 6 3, 2 2, data as lectotype (genitalia slide no. 15286; BMNH; CMAG, Birmingham).
New Guinea: 1 3, 3 9, Irian Jaya, C. Arfak Mts, Ninay Valley, 1050 m, xi.1908-i.1909 (A. E. Pratt)
1 9,1. J., Arfak Mts, 1850 m, iii.1910 (C. B. & F. B. Pratt) (BMNH); 1 9, I. J., Snow Mts, Setekwa R.,
ix.1910 (Meek) (BMNH); 8 2, I. J., Snow Mts, Upper Setekwa R., 600-1050 m, ix.1910 (Meek) (BMNH);
2 2, 1. J., Snow Mts, nr Oetakwa R., up to 1050 m, x—x1i.1910 (Meek) (BMNH); 24, 11 9,1. J., Mt Goliath,
1500-2150 m, about 139° long. [4°40’S 139°52’ E], i-ii.1911 (Meek) (BMNH; NMNH, Washington);
2 3, 2 2, Papua New Guinea, Sepik District, Telefomin (Eliptamin), 1350-1700 m, vi-ix.1959 (Brandt)
(ANIC, Canberra); 1 3, 3 2, P. N. G., Sepik District, Bainyik, 300 m, x.1957 (Munroe & Holland) (CNC,
Ottawa); 1 9, P. N. G., Western Highlands, Jimi R., 1450 m, vii—ix.1961 (Brandt) (ANIC, Canberra);
5 3, 1 9, P. N. G., Finisterre Range, Kiambari, 1350 m, vii—viii.1958 (Brandt) (ANIC, Canberra); 1 9,
P. N. G., Madang District, Finisterre Mts, Naho R. Valley, Budemu, 1250 m, x.1964 (Bacchus) (BMNH);
2 3, 8 2, P. N. G., Morobe District, Finisterre Mts, Wantoat, 1200 m, ix.1957 (Munroe & Holland)
(CNC, Ottawa); 3 9, P. N. G., Gurukor, 600 m, ii.1978 (Hébert) (CNC, Ottawa); 8 2, P. N. G., Morobe
District, near Bulolo, Araucaria forest, 1200 m, ix.1957 (Munroe & Holland) (CNC, Ottawa); 1 9, P. N.
G., Morobe District, Edie Creek, 2050 m, ix.1957 (Munroe & Holland) (CNC, Ottawa); 8 9°, P. N. G.,
Bulldog Rd, 3 km S. Edie Creek, 2200 m, viii.1975 (Hébert & Ward) (CNC, Ottawa); 1 2, P. N. G., same
locality, xii.1977 (Hébert & Harmsen) (CNC, Ottawa); 5 2, P. N. G., Eastern Highlands, Okapa, 1850 m,
x.1957 (Munroe & Holland) (CNC, Ottawa); 44 3, 9 9, P. N. G., Dinawa, 1200 m, ix.1902 (A. E. Pratt)
(BMNH; NMNH, Washington); 11 ¢, 22 9, P. N. G., Mt Kebea, 1100 m, and 1850 m, iii—viii.1903 (A. E.
Pratt) (BMNH; CMAG, Birmingham; NMNH, Washington); 4 ¢, 4 9, P. N. G., Upper Aroa R., iii-
iv.1903 (Meek) (BMNH); 5 9, P. N. G., Mafalu [sic], 1850 m, viii.1903 (A. E. Pratt) (BMNH); 1 6, 2 2,
REVISION OF VITESSIDIA & VITESSA 269
P. N. G., Mafulu, 1200 m, i.1934 (Cheesman) (BMNH); 3 9, P. N. G., Avola, 1850 m, viii.1903 (A. E.
Pratt)(BMNH); 2 2, P. N. G., Babooni, 1100 m, ix.1903 (A. E. Pratt) (BMNH); 1 92, P. N. G., Angabunga
R., affluent] of St Joseph R., 1850 m upwards, xi.1904-ii1.1905 (Meek) (BMNH); 2 3, 3 9, P. N. G.,
Tapini, Loloipa R., Bome, 1900 m, ii—v.1958 (Brandt) (ANIC, Canberra); 13 3, 19 2, P. N. G., Sogeri
Plateau, Bisianumu, 500 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 26 3, 60 9, P. N. G., Sogeri
Plateau, Mageri, vili.1957 (Munroe & Holland) (CNC, Ottawa); 10 3, 11 9, P. N. G., Mambare R., Biagi,
1500 m, i-iv.1906 (Meek) (BMNH; MNHN, Paris; NMNH, Washington); 8 3, 1 9, P. N. G., Hydro-
grapher Mts, 750 m, i-iii.1918 (Eichhorn Bros.) (BMNH); 1 3, 1 9, P. N. G., Amazon Bay area, Doveta,
750 m, vii-ix.1962 (Brandt) (ANIC, Canberra); 1 3, 1 2, no data (CMAG, Birmingham).
Vitessa brandti sp. n.
(Figs 4, 54, 137)
3. 54mm. Frons yellow. Vertex orange. Eye and ocellus brown. Chaetosema with pale yellowish setae.
Antenna dark fuscous; posterior surface of apical third of shaft white. Labial palpus black, orange on
ventral surfaces of first and second segments, third segment entirely black. Maxillary palpus orange.
Basal scaling of proboscis black. Neck region orange. Thorax above: orange; patagium orange with
posteromesal blue-black spot; tegula orange with similar anterolateral spot. Abdomen above: blue-black,
narrow white terminal line on each segment, anal tuft orange. Body below: blue-black; pectus orange;
whitish grey terminal abdominal lines, anal tuft orange. Legs blue-black, mid coxa and mid and hind
femora with whitish grey scales; hind tibia with shaggy black scales dorsally on basal half. Fore wing
above with basal area orange, bearing single small basal blue-black spot and two larger subbasal spots,
posterior spot diffuse; subbasal fascia erect, fairly wide, glossy blue-black, sharply defined on both sides.
Antemedial fascia a narrow area of jade green, diffuse medially and followed by a broad violet-grey medial
fascia, slightly expanded posteriorly. Terminal part of wing jade green with wide black stripes and narrow
wedges between veins. Costa narrowly black from base to apex. Fringe black, grey towards tornus. Hind
wing above: blue-black with metallic lustre. Fringe fuscous, distal half whitish grey, anal margin black.
Fore wing below: blue-black; oval postmedial patch white; faint light grey streaks on a few veins sub-
apically. Hind wing below: like upperside, but with veins of subapical region broadly jade green.
2. Unknown. ;
GENITALIA ¢ (Fig. 137). In general as for V. griseata (p. 267). Uncus broad, gradually tapering from
convex base, slightly waisted. Transtilla very shallowly V-shaped. Valve relatively short and broad, distally
falcate, extreme tip narrowly rounded; terminal margin at right angles to costa, becoming broadly rounded
before forming posterior margin; clasper more than half valve length from base. Aedeagus base broadly
rounded.
REMARKS. This species is similar in general appearance to V. griseata, but is separated externally
by the broad violet-grey medial fascia on the fore wing, and by the larger spots on the patagium
and tegula. In the ¢ genitalia the broader uncus and the relatively short, broad and truncated
appearance of the valve separate this species from V. griseata and V. talboti. We take pleasure in
dedicating this species to Mr W. Brandt, collector of the holotype and of many other interesting
Papuan Lepidoptera.
BioLoGy. Early stages and host-plant unknown. Holotype collected some time between June and
September.
DISTRIBUTION (Fig. 4). Bismarck Archipelago: New Britain.
MATERIAL EXAMINED
Holotype 3, New Britain: Mt Sinewit, 1050 m (‘3500 ft’), 27.vi.-17.ix.1963 (Brandt) (genitalia slide no.
DK 738; ANIC, Canberra).
Vitessa talboti (Janse, 1924) comb. n.
(Figs.4, 55,.56, 139, 174, 227, 273)
Cosmethis talboti Janse, 1924, Bull. Hill Mus. Witley 1: 503. LECTOTYPE 3, Motuccas: Central Ceram
[Seram], Manusela [3°15’S, 129°38’ E], 1850 m (‘6000 ft’), x—xii.1919 (C., F. & J. Pratt) (genitalia
slide no. 14676/a, Shaffer; TM, Pretoria), here designated [examined].
3, 2. 50-56 mm. Frons orange. Vertex orange with a few medial black scales. Eye and ocellus brown.
Chaetosema with pale yellowish setae. Antenna black; posterior surface of apical third of shaft white.
270 E. G. MUNROE & M. SHAFFER
Labial palpus black, orange on ventral surfaces of first and second segments, third segment entirely black.
Maxillary palpus orange, inner surface black. Basal scaling of proboscis brownish fuscous, a slight
sprinkling of orange at extreme base and on sides. Neck region orange. Thorax above: orange; patagium
orange with a posteromesal blue-black spot; tegula orange with diffuse anterolateral blue-black spot.
Abdomen above: blue-black, a narrow whitish grey terminal line on each segment, anal tuft orange. Body
below: black; pectus orange; abdomen black, a whitish grey terminal line on each segment, wider than
on dorsal surface; anal tuft orange. Legs black; whitish grey scaling on coxae, ventral surfaces of femora,
tips of mid and hind tibiae and bases of tibial spurs; hind tibia with shaggy black scales dorsally on
basal half. Fore wing above: basal area orange, reaching Cu, bearing single large subbasal spot; broad
subbasal fascia black with bluish green reflections; white square medial fascia, reaching posterior margin,
postmedial border weakly oblique; costa narrowly black from apex to base, in ¢ costa anterior to medial
fascia admixed with white; apical part of wing blue-black with whitish grey lining the veins, separate
from medial fascia in 3, almost touching in 9; fringe dark grey, white at tornus. Hind wing above: blue-
black with metallic reflections; fringe white admixed with grey, black at tornus and anal margin. Fore
wing below: dull blackish fuscous; white medial fascia narrower, closer to apex than on upperside,
antemedial margin irregular; apical region with traces of whitish grey; fringe as on upperside. Hind wing
below: dull blackish fuscous; apical region with white vein-lines; fringe as on upperside.
GENITALIA ¢ (Figs 139, 174). In general as for V. griseata. Uncus broad, gradually tapering from convex
base, slightly waisted. Anterior sclerite of tegumen triangular with keyhole-shaped medial anterior
emargination. Vinculum broadly rounded. Valve long and extremely narrow, distally falcate, extreme tip
very narrowly rounded and posterior margin deeply excised before it; clasper less than half valve length
from base. Juxta with rectangular lateral sclerites, each bearing broadly triangular, blunted dorsal process,
and long, slender, upturned ventral process; anellus (Fig. 174) with large, slightly sinuate, lateral sclerites
bearing large anteriorly directed spines, anterolateral edge medially protruded, bearing long, more erect
spines. Aedeagus base broadly rounded; ductus ejaculatorius subbasal. Cornuti: basal dorsolateral
clusters of slightly unequal length, left one shorter than, but extending distad of, right.
GENITALIA ° (Figs 227, 273). In general as for V. griseata, see Remarks. Posterior sclerite of ostial chamber
denticulate on lateral portions.
REMARKS. The almost square white medial fascia on the upperside of the fore wing makes this
species quite distinct. Of the known males of this group, V. ta/boti has the most narrowly falcate
valve. The 9 genitalia resemble those of V. griseata and V. nobilis, differing only slightly; posterior
apophysis slightly curved, thicker than in V. griseata and terminally truncated as in V. nobilis;
anterior apophysis similar, differing from other species by being medially angled near base;
lateral bullae on posterior sclerite smaller than in V. griseata but larger than in V. nobilis, differing
also from both in that the inner margins are nearly parallel and touching; posteriorly widened
ductus bursae smaller than in V. griseata and more nearly square, larger than in V. nobilis; ductus
bursae and corpus bursae smaller than in V. griseata and V. nobilis; signa smaller than in V.
griseata but larger than in V. nobilis.
BioLoGy. Early stages and host-plant unknown. Moths collected between October and December.
DISTRIBUTION (Fig. 4). Moluccas: Seram.
MATERIAL EXAMINED
Paralectotypes. 1 3, 2 9, data as lectotype (1 9, abdomen missing; genitalia slide no.’s 14676/b, Shaffer,
14676; TM, Pretoria; BMNH) [1 2 syntype not seen].
Vitessa nobilis sp. n.
(Figs 4, 57, 228, 274)
3g. Unknown.
2. 58 mm. Frons and vertex orange. Eye and ocellus brown. Chaetosema with pale yellowish setae.
Antenna black, posterior surface of apical third of shaft white. Labial palpus [missing since time of
description] black, first and second segments with considerable orange scaling basally and ventrally; third
segment black, nearly as long as second. Maxillary palpus orange. Proboscis with black basal scaling.
Neck region orange. Thorax above: orange; patagium black, laterally orange; tegula: black extending
whole length, laterally orange. Abdomen above: black with grey terminal line on each segment, anal tuft
orange. Body below: black; pectus black admixed with orange; abdomen black with wider grey terminal
REVISION OF VITESSIDIA & VITESSA 271
lines on segments than above; anal tuft orange; legs black, whitish grey scaling on coxae, ventral surfaces
of femora, tip of hind tibia. Fore wing above: black with metallic blue reflections; orange basal area
completely divided, leaving minute basal area with larger subbasal area reaching Cu; costal area black
from apex to base; wing veins bordered with greenish grey; medial fascia represented by a slightly post-
medial, oblong, greenish grey patch on discocellular vein, and a diffuse bilobed greenish grey patch at
middle of posterior margin; fringe black, with pale terminal scaling at tornus. Hind wing above: dull
black with faint metallic reflections; fringe whitish grey, black at anal margin. Fore wing below: dull
black with large, trapezoidal, whitish medial fascia; short whitish streak extending basally along R from
anterolateral corner of fascia; faint traces of white vein-lines at apex; fringe as on upperside. Hind wing
below: dull black; whitish grey vein-lines at apex; small greenish grey dash at angle of discocellular vein;
fringe as on upperside.
GENITALIA 2 (Figs 228, 274). In general as for V. griseata, see Remarks. Posterior sclerite of ostial
chamber (Fig. 274) coarsely denticulate on lateral portions.
REMARKS. This species occurs sympatrically with V. zemire in Ambon. The fore wing is broader
than in other species of the species-group; the upperside markings are completely distinct from
those of all other species of Vitessa. The 2 genitalia resemble those of V. griseata and V. talboti,
but differ in the following characters: the posterior apophysis is thick and truncated as in V.
talboti but straight; the anterior apophysis is long, slightly curved, thicker than in V. griseata,
without basal angle as in V. ta/boti; the inner margins of the lateral bullae on the posterior wall
of the ostial chamber diverge posteriorly, and do not converge medially as in V. griseata, or run
parallel as in V. talboti; the anterior margin of the anterior sternite is recurved and deeply
wrinkled as in V. griseata, more strongly wrinkled than in V. talboti; the ductus bursae is
posteriorly widened and funnel-shaped as in V. griseata but narrower, not square as in V. talboti;
the signa in the corpus bursae are smaller than in V. griseata or V. talboti.
BrioLoGy. Early stages and host-plant unknown. Holotype collected in November.
DISTRIBUTION (Fig. 4). Moluccas: Ambon.
MATERIAL EXAMINED
Holotype 9, Moluccas: Ambon, Mt Salohotus, no. 17844, xi.1923 (C. J. Brooks) (genitalia slide no.
14697; BMNH).
The zemire-group
D1aGnosis. Hind wing above: black, as in griseata-group, but fore wing above with two transverse whitish
fasciae on blackish ground colour, in addition to orange basal or subbasal patch and pale terminal vein-
lines. 3 genitalia with uncus slender or broadly flattened, decurved, with weakly differentiated distal
portion bearing laterally directed setae; anellus with large lateral clusters of spines as in griseata-group;
valve distally rounded or truncate, not falcate, fluted with oblique striations and mostly without clasper.
2 genitalia more variable, with well-developed lateral, sclerotized, pouch-like invaginations beside ostial
chamber, or twin bullae on posterior sclerite as in griseata-group; anterior margin of anterior
sclerite strongly wrinkled; anteriorly extended antrum with conical or broadly flattened sclerotization,
or antrum greatly reduced; corpus bursae with two round, depressed denticulate signa.
REMARKS. Within the zemire-group there are three subgroups: (1) V. cyanea, with 3 genitalia like
those of V. zemire and with @ genitalia similar to those of the griseata-group; (2) V. zemire, with
valve in 3 not bearing a clasper and with @ genitalia having the distinctive long flattened antrum;
(3) V. vitialis and V. tamsi, where the 3 has a long club-shaped clasper on a valve otherwise similar
to that of V. zemire, and where the sclerotized antrum of the Q genitalia is greatly reduced or
lacking.
Vitessa cyanea Hampson, 1906
(Figs 12, 34, 58-61, 140, 141, 175, 176, 205, 206, 229, 230, 275, 276)
Vitessa cyanea Hampson, 1906 : 217.
3, 2. 38-46 mm. Frons and vertex orange. Eye fuscous. Ocellus well developed, brown. Chaetosema with
pale yellowish setae. Antenna black, some orange scaling at base of scape: posterior surface of apical
IA 92 E. G. MUNROE & M. SHAFFER
third in ¢ with some dark grey scaling. Labial palpus with first and second segments orange, second
segment black dorsally, lightly admixed with orange; third segment black with some orange at base.
Maxillary palpus orange with ventral black spot, some black scales admixed terminally. Basal scaling
of proboscis black, lightly admixed with orange at extreme base. Neck region orange. Thorax above:
orange; patagium orange with posteromedial blue-black spot; tegula orange with an anterolateral blue-
black spot. Abdomen above: blue-black; terminal margins of most segments finely edged with white,
broader laterally, last segment with terminal margin orange, acuminately blue-black at centre. Body below
and laterally: blue-black; pectus orange; terminal margins of abdominal segments broadly white; last
segment with terminal margin completely orange. Legs blue-black; base of fore coxa black, broadly
bordered with orange; light grey scaling at base of fore femur and ventral surface of fore tibia with reddish
white scaling on ventral surface of epiphysis. Fore wing rather narrow; above with basal area orange,
strongly indented at regions of A, and A,, outer margin broadly curved, not reaching costa; single
subbasal spot blue-black ; subbasal, medial and terminal areas blue-black ; creamy white antemedial fascia
very narrow, sometimes incomplete, erect, not reaching costa or posterior margin, reduced to shorter
hair-line in 9; creamy white postmedial fascia large, oval, obliquely placed at end of cell, terminally
extending into weak white terminal lines; fringe dark grey. Hind wing above: black with weak violaceous
reflections; fringe dark grey apically, white towards anal margin, anal margin black. Fore wing below:
black with violaceous reflections; postmedial fascia of similar proportions to that of upperside but
without apical diffusion, diffusion stronger medially; posterior margin basally orange brown, medially
white; fringe dark grey. Hind wing below: black with violaceous reflections; large white apical patch
diffusing slightly along costa; fringe terminally white, anal margin black.
GENITALIA ¢ (Figs 34, 140, 141, 175, 176). Uncus slender, abruptly tapering from convex base, distal
portion down-curved, then up-curved, with sparse scattering of long, laterally directed setae. Gnathos
with flattened, medially angled, wide lateral arms and with compressed, basally straight, gently upturned
and finely acuminate process. Transtilla V-shaped, medially broadened or narrowed. Tegumen with
posterior sclerite short, wide; anterior sclerite with keyhole-shaped medial anterior emargination.
Vinculum ventrally narrow, broadly rounded posteriorly. Valve long, widest at centre, distally finely or
broadly truncated; costa basally inflated; many fine oblique striations on mesal surface, slightly stronger
striations towards posterior margin. Juxta with two lateral sclerites, each bearing a dorsal bluntly
triangular process, deeply recurved and sparsely setose, supporting base of anellus, and a ventral,
posteriorly directed, slender, sinuate and acuminate process, tip nearly reaching mid line; anellus (Figs
175, 176) arising from dorsal processes of juxta, large lateral sclerites, anteromedially directed, bearing
large anteriorly directed spines; on dorsal surface of anellus between anterior ends of lateral sclerites,
an elongated, oval, medial catena (Fig. 34), consisting of a cluster of minute posteriorly directed spines;
remainder of anellus just reaching medial catena, minutely spinose; manica supported by lightly
sclerotized elongated bands continuous with lateral sclerites of anellus, narrowing towards tip. Aedeagus
of moderate length, expanded distally, slightly curved, base narrowly rounded; ductus ejaculatorius
subbasal; scale-like spines extending dorsally and ventrally for a short distance onto aedeagus from
vesica; base of vesica bulbous ventrally at distal end of aedeagus; dorsal and ventral surfaces covered with
scale-like spines from base of vesica to halfway along dorsolateral clusters of cornuti, the spinose areas
longitudinally rugose. Two large basal lateral cornuti clusters of equal length, the left placed a little distad
of the right; oblique posterior ventromedial cluster reduced to sclerotized bases (Figs 205, 206).
GENITALIA @ (Figs 229, 230, 275, 276). High, wide ovipositor lobes, dorsally fused and prolonged
posteriorly, densely setose, setae longer on margin; spiny setae not easily seen. Posterior apophysis
thick, curved, distally blunt and enlarged. Anterior apophysis thick, slightly curved, widened near base,
distally blunt and enlarged. Posterior sclerite of ostial chamber (Figs 275, 276) broadly sclerotized, antero-
medially broadly divided by a narrow sulcus, laterally a pair of wrinkled denticulate bullae with rounded,
posteriorly converging inner margins. Anterior sclerite narrow, wrinkled, denticulate and reflected.
Antrum short, wide. Ductus bursae membranous, posteriorly widened and flattened. Corpus bursae oval,
membranous, with pair of large, heavily sclerotized, depressed, denticulate signa.
REMARKS. This species resembles V. zemire in general appearance, but has narrower wings and
slighter build. The postmedial fascia on the upperside of the fore wing is clearly connected to the
white terminal lines. The $ genitalia are like those of V. zemire but differ in having the valve
distally truncate, not narrowly rounded, and with a more pronounced ridge in normal position of
clasper. The catena is reminiscent of that of the griseata-group. The 2 genitalia most clearly
resemble the griseata-group in the structure of the sclerites around the ostial chamber, in the
development of the lateral bullae divided by a sulcus, and in the shape and size of the antrum.
REVISION OF VITESSIDIA & VITESSA ab
The @ genitalia differ in important characters from those of V. zemire, the ostial sclerites and
antrum being very different in shape.
We recognize two subspecies from the following geographical areas: (1) southern Moluccas;
(2) eastern New Guinea.
BioLoGy. Early stages and host-plants unknown. Moths collected from January to March,
September to December.
DISTRIBUTION (Fig. 12). Southern Moluccas: Seram. New Guinea: Papua New Guinea. D’Entre-
casteaux Islands: Fergusson Island.
Vitessa cyanea ceramensis subsp. n.
(Figs 12;-58; 59; 140}175,-205,; 229,275)
3, 9. 38-46 mm. Generally as described for species. Fore wing above with white postmedial fascia
broadly reaching main stem of R in 3; in 2 anteromedial corner of postmedial fascia protruded basally
into cell. White antemedial fascia in 2 narrow and complete but not reaching costa or posterior margin,
diffuse medially.
GENITALIA 3 (Figs 140, 175, 205). Generally as described for species. Uncus slender, gently tapering.
Transtilla V-shaped and medially narrow. Valve widest a little beyond centre, distally finely truncated.
Anellus (Fig. 175) with dorsal medial catena oval.
GENITALIA 2 (Figs 229, 275). Generally as described for species, but see Remarks.
REMARKS. The specimens are externally in poor condition, especially the 3. The fore wing above
has the postmedial fascia broader than in the nominate subspecies and, in the Q at least, the
antemedial fascia is broader. In the ¢ there is virtually no sign of the antemedial fascia or orange
basal area, probably due to wear. The 3 genitalia are distinguished from those of the nominate
subspecies by the narrower uncus, somewhat narrower transtilla, broader valve with narrower
apical truncation, shorter medial catena and shorter cornuti clusters. The 2 genitalia are dis-
tinguished by the smaller, not distally enlarged apophyses, smaller bullae on the posterior ostial
sclerite (Fig. 275), smaller corpus bursae and larger signa. The genital differences are considerable,
and may prove to be of specific value.
BIoLoGy. Early stages and host-plant unknown. Allotype collected in December.
DISTRIBUTION (Fig. 12). Southern Moluccas: Seram.
MATERIAL EXAMINED
Holotype 9, Moluccas: Seram (‘Ceram’), ‘Jllo’, 1901 (Coll. E. L. Ragonot) (genitalia slide no. 15302/a,
Shaffer; MNHN, Paris).
Paratype. Moluccas: 1 3 (allotype), Central Ceram, 29.20, 3000 ft [900 m], xii.1919 S F. & J. Pratt)
(genitalia slide no. 15302/b, Shaffer; TM, Pretoria).
Vitessa cyanea cyanea Hampson, 1906
(Figs 12, 34, 60, 61, 141, 176, 206, 230, 276)
Vitessa cyanea Hampson, 1906, Ann. Mag. nat. Hist. (7) 17: 217. Holotype 3, D’ENTRECASTEAUX
ISLANDS: Fergusson I., x.1894 (Meek) (BMNH) [examined].
3, 2. 40-46 mm. Generally as described for species. Fore wing above with white postmedial fascia
narrowly reaching main stem of R veins in both sexes, medial margin broadly rounded. White antemedial
fascia in 2 and sometimes in ¢ reduced to a hair-line.
GENITALIA ¢ (Figs 34, 141, 176, 206). Generally as described for species. Transtilla medially broadened.
Valve apex broadly truncated. Medial catena (Figs 34, 176) with additional smaller lateral patches.
GENITALIA & (Figs 230, 276). As described for species.
REMARKS. The distinguishing features from V. cyanea ceramensis are given under that subspecies.
BioLoGy. Early stages and host-plant unknown. Moths collected from January to March,
September to December.
274 E. G. MUNROE & M. SHAFFER
DISTRIBUTION (Fig. 12). New Guinea: Papua New Guinea. D’Entrecasteaux Islands: Fergusson
Island.
MATERIAL EXAMINED
New Guinea: 8 3, Papua New Guinea, Hydrographer Mts, 750 m, i-iii.1918 (Eichhorn Bros.) (BMNH).
D’Entrecasteaux Islands: 2 3, 1 9, Fergusson I., ix—xii.1894, xii.1914 (Meek) (BMNH).
Vitessa zemire (Stoll, [1781])
(Figs 5-9, 26, 62-77, 142-148, 177-182, 207, 208, 231-238, 277-284, 317)
Phalaena zemire Stoll, [1781] : 149, pl. 367, fig. 1.
3, 2. 37-62 mm. Frons and vertex orange. Eye and ocellus fuscous. Chaetosema with pale yellowish setae.
Antenna black; anterior surface of scape orange; posterior surface of apical third of shaft white to variable
extent; terminal segments sometimes black. First and second segments of labial palpus either with
ventral and lateral surfaces orange, dorsal surfaces black, or completely orange; third segment black.
Maxillary palpus basally black, distally orange. Basal scaling of proboscis black, admixed with orange at
extreme base. Neck region orange. Thorax above: orange; patagium with large, posteromedial, metallic
blue-black spot; tegula orange, with anterolateral metallic blue-black spot varying in length. Abdomen
above: matt black, a narrow whitish grey terminal line on most segments, posterior half of terminal
segment and anal tuft orange. Body below and laterally: blue-black; pectus orange; whitish grey
abdominal lines wider than above; posterior half of terminal segment and anal tuft orange. Legs black;
tibiae with metallic green reflections; variable amounts of whitish grey on some parts; anterior end of
upper surface of hind tibia with long black scales. Fore wing rather wide; above: basal area orange, outer
margin straight, slightly sinuate to greatly indented at A, oblique, bearing small black subbasal spot;
ground colour black with metallic blue or green reflections; subbasal fascia variable in width, glossy blue-
black; white or greenish white, parallel, somewhat oblique, antemedial and postmedial transverse fasciae
of variable width, sometimes irregularly extended towards each other, rarely joining, the postmedial fascia
always the broader; whitish vein-lines in terminal and apical regions; fringe fuscous with bronze gloss at
intervals. Hind wing above: black with violaceous reflections; fringe glossy fuscous, whitish grey towards
anal margin. Fore wing below: blue-black; single white postmedial fascia present, of variable width;
posterior margin basally orange brown, medially white; retinaculum dark grey. Hind wing below: blue
black; variable amount of white in apical region, either solid or consisting of wide bars on veins; some-
times a white discal spot present; fringe wholly grey or white.
GENITALIA ¢ (Figs 26, 142-148, 177-182, 207, 208). Uncus slender, gradually tapering, distal portion
down-turned then up-turned, with sparse scattering of long, laterally directed setae. Gnathos with
flattened, medially angled, wide lateral arms and with compressed, basally straight, but dorsoventrally
widened, up-turned and sickle-shaped, acuminate medial process. Transtilla shallowly V-shaped and
medially narrowed or of even width. Tegumen with posterior sclerite short and wide, anterior sclerite
widely emarginated medially. Vinculum ventrally broad and basally convex. Valve long, widest at centre,
distally tapering to a narrowly rounded apex; costa inflated; some oblique striations on ventral surface,
clasper absent. Juxta with lateral sclerites chelate, each bearing a dorsal, bluntly triangular and thickly
setose process and a ventral, slender, straight and acuminate process nearly reaching mid-line, supporting
base of anellus. Anellus (Figs 177-182) with large lateral sclerites, strongly angled medially, bearing large
anteriorly directed spines, sclerite connected directly to dorsal process (Fig. 26); ventral surface of anellus
above lateral sclerite longitudinally rugose and lightly sclerotized, not spinose medially; medial catena
absent; manica supported by dorsal, sclerotized, caliper-shaped bands arising from lateral sclerites on
anellus, narrowing and Jess sclerotized towards tip. Aedeagus of moderate length, slightly expanded
distally, straight, base narrowly rounded; ductus ejaculatorius subbasal; vesica bulbous ventrally at distal
end of aedeagus and transversely rugose; ventrally, longitudinally rugose between cornuti clusters;
dorsally and ventrally covered with scale-like spines; slightly bulbous at distal end of cornuti. Cornuti
(Figs 207, 208) grouped in two clusters of spines, the left cluster ventrolateral and much smaller than the
dorsolateral right cluster, or sometimes completely absent. No ventral cluster of cornuti.
GENITALIA & (Figs 231-238, 277-284, 317). Ovipositor lobes long, slender, dorsally fused; surface densely
setose, margin with longer setae; spiny setae not easily seen (Fig. 317). Posterior apophysis with weak
vertical bar and slender shaft, distally spatulate. Anterior apophysis slightly longer, thicker and curved,
distally spatulate. Posterior ostial sclerite (Figs 277-284) sclerotized as a broad band, transversely wrinkled
and finely denticulate, forming laterally a pair of shallow invaginations slightly intruded into the pouches
of anterior sclerite, sclerite bearing faintly produced bullae meeting in mid-line. Anterior sclerite forming
REVISION OF VITESSIDIA & VITESSA 21D
laterally a pair of shallow, pouch-like invaginations, connected medially, posterior margin wrinkled and
reflected. Antrum wide and long, slightly waisted, sclerotized, broadly rounded posteriorly, fitting between
invaginations, heavily sclerotized laterally, forming rounded bars, anteriorly ending irregularly in widened
ductus bursae. Ductus bursae funnel-shaped at base of antrum, remainder narrow, membranous and
minutely spinose. Corpus bursae globular, membranous, with a pair of large, heavily sclerotized,
depressed, denticulate signa.
REMARKS. This species is most closely related to V. vitialis and V. tamsi, but does not occur
sympatrically with these species, which are limited to Fiji and Samoa respectively. Differential
characters are discussed under V. vitialis and V. tamsi (pp. 111, 112).
Individual populations of V. zemire vary in external colour and pattern, but the structural
differences are slight. We recognize eight subspecies, from the following geographical areas:
(1) northern Moluccas; (2) southern Moluccas; (3) Misodl, Salawati, Waigeo, New Guinea,
Kepulauan Aru and D’Entrecasteaux Islands; (4) Queensland; (5) Umboi Island; (6) New Britain
New Hannover and presumably New Ireland; (7) Solomon Islands; and (8) New Hebrides. More
material may show that additional subspecies should be distinguished in the Moluccas and eastern
Melanesia.
BioLoGy. Early stages and host-plants unknown. Moths collected by many collectors in all
months of the year.
DISTRIBUTION (Figs 5-9). Moluccas; New Guinea and its coastal islands; Kepulauan Aru;
D’Entrecasteaux Islands; Queensland; Bismarck Archipelago; Solomon Islands and New
Hebrides.
Vitessa zemire expansa subsp. n.
(Figs 5, 62, 63, 231, 277)
3, 2. 48-58 mm. Generally very similar to the nominate subspecies. Fore wing above: orange basal area
large, distal margin slightly protruded, straight or slightly sinuate; subbasal spot distinct. Antemedial
and postmedial fasciae creamy white; antemedial fascia very narrow, diffuse medially; postmedial fascia
broad, diffuse medially at base, anteriorly rounded, not reaching costa, hardly or not reaching posterior
margin, terminal margin of fascia curved. Vein-lines in terminal area not reaching postmedial fascia.
Hind wing below with pale apical patch tending to break into bars along veins.
GENITALIA 3. Generally as described for species. Right lateral cornuti cluster of vesica long and narrow
with well-developed spines; left lateral cornuti cluster much narrower, spines greatly reduced in size,
largest posteriorly.
GENITALIA 2 (Figs 231, 277). As described for species.
REMARKS. This subspecies is distinguished from V. zemire zemire by creamy white as opposed to
chalky white fasciae; the antemedial fascia is slightly wider than in V. zemire zemire. The ¢ is
known only from a single non-topotypic specimen and may therefore not be typical of the
subspecies. The 9 is like that of V. zemire zemire but larger and with larger orange basal area. In
the ostial chamber of the genitalia (Fig. 277) the anterior margin of the posterior sclerite is
broader than in V. zemire zemire, also the antrum is longer and broader, with a slight waist near
anterior end. The subspecies is distinguished from others by the geographical range, wing
markings and genital characters.
BioLoGy. Early stages and host-plant unknown. Moths collected in May and August.
DISTRIBUTION (Fig. 5). Northern Moluccas: Halmahera and Batjan.
MATERIAL EXAMINED
Holotype 2, Moluccas: Halmahera (‘Halmaheira’), Toeelo [Tobelo?] (Bought from E. Le Moult)(BMNH).
Paratypes. Moluccas: 19 9, data as holotype (genitalia slide no.’s 14707, 14708, 15288; BMNH); 1 9,
Halmahera (Waterstradt) (BMNH); 1 2, Halmahera, Dodinga, 1892 (Doherty) (BMNH).
Material excluded from paratype series. Moluccas: | 3, 2 2, Batjan, v.1892, viii.1897 (Doherty) (BMNH);
1 9, Ins. Obi (Bernst.) (RNH, Leiden).
276 E. G. MUNROE & M. SHAFFER
Vitessa zemire zemire (Stoll, [1781])
(Figs 5, 64, 65, 142, 232, 278, 317)
Phalaena zemire Stoll, [1781], in Cramer, Papillons Exotiques, 4: 149, pl. 367, fig. I. Holotype 32,
Mo uccas: Ambon (‘Amboina’) (type lost; at the time of description in the collection of the Prince of
Orange). [Subspecies identified from coloured figure and tocality.]
Cosmethis zemire (‘Cramer’); Hiibner, [1820] : 179; Janse, 1924 : 503.
Vitessa zemira (‘Cramer’); Lederer, 1863 : 335, pl. 6, fig. 8; Pagenstecher, 1884 : 266, 314; 1888 : 184;
Ragonot, 1891 : 110. [Incorrect subsequent spellings.]
Vitessa zemire (‘Cramer’); Walker, 1864 : 220; Swinhoe, 1900 : 429; Rothschild, 1915 : 223.
Vitessa zalmira (‘Cramer’); Hampson, 18965 : 503. [Incorrect subsequent spelling. ]
Cosmethis zemire (Stoll); de Joannis, 1930 : 637.
3, 2. 37-56 mm. Generally as described for species. Fore wing above: orange basal area variable, outer
margin straight, curved or indented at cell and A; subbasal spot distinct, or merging with ground colour
when area indented. Antemedial and postmedial fasciae chalky white; antemedial fascia very narrow,
sometimes incomplete; postmedial fascia broad, slightly diffuse medially at base, anteriorly rounded, not
reaching costa, hardly or not reaching posterior margin, terminal margin curved. Vein-lines in terminal
area reduced or obsolete, not reaching postmedial fascia. Hind wing below with pale apical patch tending
to break into bars along veins.
GENITALIA 3 (Fig. 142). Generally as described for species, but see Remarks.
GENITALIA ° (Figs 232, 278, 317). Generally as described for species, but see Remarks.
REMARKS. Distinguished from V. zemire expansa by chalky white fasciae as opposed to creamy
white fasciae. Antemedial fascia slightly narrower than in V. zemire expansa. The examined
specimens are very variable and may include as yet unrecognized subspecies; they are generally
smaller than V. zemire expansa. The only known @ is in poor external condition and has genitalia
identical with those of V. zemire expansa, also known from only one 3; in both specimens the
genitalia are somewhat damaged. In the 9 genitalia the anterior margin of the posterior ostial
sclerite (Fig. 278) is narrower than in V. zemire expansa, also the antrum is shorter and narrower,
with a slight waist in the mid region. The subspecies is distinguished from others by the geo-
graphical range, wing markings and genital characters.
BioLoGy. Early stages and host-plant unknown. Moths collected in January and February.
DISTRIBUTION (Fig. 5). Southern Moluccas: Seram, Ambon and Saparua.
MATERIAL EXAMINED
Moluccas: 1 2, Seram (ex Felder coll.) (BMNH); 1 9, Central Seram, Mansela [sic] [3°15’ S, 129°38’ E],
650 m, 1912 (Stresemann) (BMNH); 1 9, Seram, Wahaai (Moens) (RNH, Leiden); 1 2, Ambon (ex
Musaeo Dr. Boisduval) (BMNH); 1 2, Ambon (Doleschal) (BMNH); 4 2, Ambon, ii.1892 (Doherty)
(BMNH); 1 2, Ambon (Wallace) (UM, Oxford); 1 2, Saparua, Oeliasers, 1.1892 (exp. Martin) (BMNH);
1 3, Saparua (Fruhstorfer) (BMNH); 1 2, no data (ex Felder coll.) (BMNH).
Vitessa zemire temerata Swinhoe, 1906 stat. n.
(Figs 6,:66, 67, 143, 177,233,279)
Vitessa zemira (‘Cramer’); Pagenstecher, 1886 : 167, 188. [Partim; incorrect subsequent spelling. ]
Vitessa temerata Swinhoe, 1906, Ann. Mag. nat. Hist. (7) 18 :413. LECTOTYPE 3, NEw Guinea: Papua,
Granville [‘Port Moresby’] 1907-128 (genitalia slide no. 15289; BMNH), here designated [examined].
Vitessa zemira (‘Cramer’); Kenrick, 1907 : 75. [Incorrect subsequent spelling. ]
Cosmethis temerata (Swinhoe); Janse, 1928 : 88.
3, 2. 38-62 mm. Generally as described for species. Fore wing above with orange basal area slightly
variable, large, distal margin straight or curved; subbasal spot distinct. Antemedial and postmedial
fasciae creamy white; antemedial fascia as wide as or wider than glossy blue-black subbasal fascia,
medially diffuse; postmedial fascia very wide, width more than half length of fascia, slightly diffuse
medially, distal margin sinuate, fascia truncate, short of costa but reaching posterior margin. Vein-lines
in terminal area not reaching postmedial fascia. Hind wing below with pale apical patch tending to break
into bars along veins but solid till past M,.
REVISION OF VITESSIDIA & VITESSA ZN.
GENITALIA ¢ (Figs 143, 177). Generally as described for species, but see Remarks.
GENITALIA @ (Figs 233, 279). Generally as described for species, but see Remarks.
REMARKS. This is distinguished from all other subspecies of V. zemire by the very broad, sharply
defined postmedial fascia and fairly wide antemedial fascia, being closest in appearance to V.
zemire expansa and V. zemire zemire; the subspecies nephritica from Queensland has equally
broad fasciae but is distinguished by the diffusion of both into the medial region. In the 3 genitalia
the cornuti clusters are larger than in previous subspecies, the left cluster has much reduced, almost
obsolete spines for most of posterior length. The 2 genitalia are almost identical to those of
previous subspecies, but most closely resemble those of V. zemire expansa in the shape of the
antrum (Fig. 279), though in the present subspecies the antrum is shorter.
BioLoGy. Early stages and host-plant unknown. Moths collected by many collectors in all months
of the year.
DISTRIBUTION (Fig. 6). Miso6l; Salawati; Waigeo; New Guinea; Kepulauan Aru; D’Entre-
casteaux Islands.
MATERIAL EXAMINED
Paralectotypes. New Guinea: 1 3, 2 2, labelled 6846, otherwise data as lectotype (genitalia slide no.
15290; BMNH); 11 3, 14 2, Mount Kebea, 1100 m (‘3600 ft’), vi—viii.1903 (A. E. Pratt) (BMNH).
Misodl: 5 3, Mysol, foot-hills, 30-150 m, wet season, x—xi.1916 (Frost) (BMNH); 1 9, central Mysol,
bush country, about 60 m, wet season, x.1916 (Frost) (BMNH); 1 2, Misol, i.1899 (Kiihn) (BMNH).
Salawati: 1 3, (Bernst) (RNH, Leiden); 1 3, Salvati [Salawati ?], 1875 (Bruijn) (BMNH). Waigeo: 1 2,
2 2, Waigeu (Waterstradt) (BMNH); 1 3, Waigeoe, 1875 (Bruijn) (BMNH); 2 3, 5 2, Waigeu, Camp Nok,
750 m, v.1938 (Cheesman) (BMNH). New Guinea: 2 92, Irian Jaya, Mamberamo, Pionierbivak, vii.1920
(Heurn) (RNH, Leiden); 4 3, I. J., Sorong, 1875 (Bruijn) (BMNH); 2 2, I. J., Dorey Bay, Andai (Doherty)
Arfak Mts, Ninay Valley, 1050 m, xi.1908-i.1909 (Pratt) (BMNH); 1 9, I. J., Arfak Peninsula, Momi
Coast (= Wariab), 11.1928 (Mayr) (BMNH); 2 4, 6 9, I. J., Kapaur, low country, xii.1896—i.1897 ( Doherty)
(BMNH); 1 9, I. J., Kapaur (Doherty) (MM, Manchester); 1 9, I. J., River Tor (mouth), 4 km E. of
Hol[andia], Maffen, vii.1959 (Maa) (BPBM, Honolulu); 1 3, I. J., Wandammen Mts, 900-1200 m,
xi.1914 (A. C. & F. Pratt) (TM, Pretoria); 1 92, I. J., Geelvink B[a]y, Nomnagihé, 600 m, i-ii.1921 (Pratt)
(TM, Pretoria); 2 3g, 2 2, I. J., Nomnagihé, 15-33 km south of Wangaar, 600 m, i-ii.1921 (C., F. & J.
Pratt) (TM, Pretoria); 1 9, I. J., Etna Bay, viii.1896 [3°58’ S, 134°47’ E] (Kiihn) (BMNH); 4 2, 4 9, I. J.,
Snow Mts, nr Oetakwa R., 1050 m, x—xii.1910 (Meek) (BMNH); 1 9, I. J., Utakwa [‘Oetakwa’] R.,
750-900 m, ii.1913 (Wollaston) (BMNH); 1 3, 3 9, I. J., Snow Mts, Upper Setakwa R., 600-900 m,
viii-ix.1910 (Meek) (BMNH); 1 9, I. J.. Mt Goliath, 1500 m, about 139° Long. [4°40’S, 139°52’ E],
ili.1911 (Meek) (BMNH); 1 3, 1 92, I. J., Cyclops Mts, Sabron, Camp 2, 600 m, vii.1936 (Cheesman)
(BMNH); 1 9, I. J., Humboldt Bay, ix—x.1892 (Doherty) (BMNH); 1 3, I. J.. Humboldt Bay Dist., Uskwar,
v.1937 (Stiiber) (BMNH); 1 3, Papua New Guinea, Humboldt Bay Dist., Wembi, vii.1937 (Stiiber)
(BMNH); 1 2, P. N. G., Njau-limon, vi. 1937 (Stiiber) (BMNH); 1 9, P. N. G., Torricelli Mts, Sea Falls
between Afua & Chinapelli villages, iv.1939 (Moore) (BMNH); 2 2, P. N. G., Torricelli Mts, Mokai,
800 m, xii.1958 (Brandt) (ANIC, Canberra); 1 3, 1 2, P. N. G., Sepik Dist., Bainyik, 300 m, x.1957
(Munroe & Holland) (CNC, Ottawa); 1 2, P. N. G., Sepik Dist., Angoram, 6 m, 1959 (Brandt & Hallstrom)
(ANIC, Canberra); 1 9, P. N. G., Telefomin, 1350-1700 m, vi—ix.1959 (Brandt) (ANIC, Canberra); 8 9,
P. N. G., Astrolabe Bay, iii-x (Wahnes) (BMNH); 1 9, P. N. G., Finisterre Range, Kiambavi, 1350 m,
vi-vii.1958 (Brandt) (ANIC, Canberra); 1 9, P. N. G., Finisterre Mts, Wantoat, 1200 m, ix.1957 (Munroe
& Holland) (CNC, Ottawa); 3 3, 4 9, P. N. G., Mount Kebea, 1100 m, vi—-viii.1903 (A. E. Pratt) (BMNH;
CMAG, Birmingham); 2 2, P. N. G., Aroa R., Booboomie, 600 m, v.1905 (Meek) (BMNH); 6 2, P. N. G.,
Aroa R. (Meek) (BMNH); 3 3, 3 9, P. N. G., Lower Aroa R., xi.1904-iii1.1905 (Meek) (BMNH); 2 9°,
P. N. G., Upper Aroa R.., iii.1903 (Meek) (BMNH); 7 3, 7 2, P. N. G., Babooni, 900-1100 m, vii-ix.1903
(A. E. Pratt)(BMNH; CMAG, Birmingham); 1 9, P. N. G., Ekeikei, 450 m, iii-iv.1903 (Pratt) (BMNH);
2 3, 1 2, P. N. G., Foula, 1350 m, viii.1903 (Pratt) (BMNH); 1 9, P. N. G., St Joseph R. [8°56’S,
147°20’ E] (Weiske) (BMNH); 1 2, P. N. G., Brown R. (Weiske) (BMNH); 4 2, 6 2, P. N. G., Sogeri
Plateau, Bisianumu, 500 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 5 3, 11 2, P. N. G., Sogeri
Plateau, Mageri, 500 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 1 2, P. N. G., Central Dist.,
Subitana, 550 m, i.1949 (Brandt & Hallstrom) (ANIC, Canberra); 2 3, 2 2°, P. N. G., Mambare R., Biagi,
1500 m, i-iii.1906 (Meek) (BMNH); 3 3, 6 2, P. N. G., Hydrographer Mts, 750 m, i-iii.1918 (Eichhorn
278 E. G. MUNROE & M. SHAFFER
Bros.) (BMNH); | 2, P. N. G., Collingwood Bay, Haidana (Meek) (BMNH); 2 4, 2 2, P. N. G., Amazon
Bay area, Doveta, 750 m, vii—ix.1962 (Brandt) (ANIC, Canberra); 1 3, 13 2, P. N. G., Milne Bay, xii.1898—
ii.1899 (Meek) (BMNH; NMNH, Washington); 2 9, P. N. G., Sariba I. (Meek) (BMNH); 1 2, no data
(Barnett) (BMNH); 1 2, no data (Joicey coll.) (BMNH); 2 2, no data (Rosenberg) (RSM, Edinburgh);
2 2, no data (Walsh) (RNH, Leiden); 2 9, no data (BMNH); | 9, no data (UM, Oxford). Kepulauan Aru:
1 3, Aroe [Aru] (Kahn) (BMNH); 1 3, Ureiuning (BMNH); 1 9, Dobbo (Dobo), 1897 (Doherty) (BMNH).
D’Entrecasteaux Islands: 3 2, Fergusson I., xii.1894—xii.1895 (Meek) (BMNH).
Vitessa zemire nephritica subsp. n.
(Figs 6, 26, 68, 69, 144, 178, 234, 280)
3, 2. 43-56 mm. Generally as described for species. Fore wing above with orange basal area large, outer
margin finely protruded at costa, mid-line and posterior margin; subbasal spot distinct. Antemedial
and postmedial fasciae pale jade green; antemedial fascia broad, diffuse medially and at points meeting
diffusion from very broad postmedial fascia; postmedial fascia finely separated from costa but reaching
posterior margin, terminal margin sinuate. Vein-lines in terminal area strongly developed and at some
points faintly joining postmedial fascia. Hind wing below with pale apical patch tending to break into
bars along veins, but solid till past M,, faintly extended down to Cuz.
GENITALIA ¢ (Figs 26, 144, 178). Generally as described for species, but see Remarks.
GENITALIA & (Figs 234, 280). Generally as described for species, but see Remarks.
REMARKS. This is distinguished from all other subspecies of V. zemire by the very broad fasciae,
which diffuse medially and join in places. The jade green coloration is striking in fresh material,
but is faded in old specimens. In the 3 genitalia the right cornuti cluster is slightly shorter than in
V. zemire temerata, the left cornuti cluster is greatly reduced in length, its anterior spines being
nearly obsolete, mostly represented only by sclerotized bases. The 9 genitalia have the antrum
(Fig. 280) slightly narrower than in V. zemire temerata, its sides more nearly parallel but the
lateral areas much less heavily sclerotized, and the anterior waist hardly accentuated. The signa
are smaller than in all previous subspecies.
BrioLoGy. Early stages and host-plant unknown. Moths collected from March to May.
DISTRIBUTION (Fig. 6). Australia: Queensland.
MATERIAL EXAMINED
Holotype 3, Australia: Queensland, Mt Edith, 18 mls [29 km] north-east of Atherton, 3400 ft [1050 m],
18.i11.1964 (Common & Upton) (ANIC, Canberra).
Paratypes. Australia: 1 2 (allotype), Queensland, 27 km (‘17 mls’) south of Atherton, 900 m (‘3000 ft’),
19.i11.1964 (Common & Upton) (ANIC, Canberra); 5 3, 2 9, Q., 18 km (‘11 mls’) south of Ravenshoe,
800 m (‘2700 ft’), iii.1964 (Common & Upton) (ANIC, Canberra; CNC, Ottawa); 1 3, Q., Mt Bartle Frere,
East Base [17°27’ S, 145°53’ E], 24 m (‘80 ft’), iii.1955 (Common) (ANIC, Canberra); 1 2, Q., L. Barrine
[17°16’ S, 145°38’ E], v.1939 (ANIC, Canberra); 5 3, 8 2, Q., Atherton Tableland, Millaa Millaa, 750 m
(‘2500 ft’), iv.1932 (Darlington) (Genitalia slide no.’s DK 69, DK 70; MCZ, Cambridge, Mass.; CNC,
Ottawa); 1 3, 2 2, Q., Kuranda, n. Cairns, 1910 (Dodd) (Genitalia slide no.’s 15291, 15292; BMNH);
1 2, same locality, 26.v.[19]02 (F.7.) (ANIC, Canberra); 1 2, Q., Atherton (ANIC, Canberra); 1 3, Q.,
5 km (‘3 mls’) west of Mossman, 4.iii.[19]64 (Common & Upton) (ANIC, Canberra); | 3, Q., Palmerston
National Park, on Tully-Cairns Power Line, at light, in rain forest, 6.xi.1966 (Britton) (ANIC, Canberra);
1 9, Q., 14km (‘9 mls’) west of Kennedy, 610 m (‘2000 ft’), 17.iv.1969 (Common & Upton) (ANIC,
Canberra); 1 3, no data (MNHN, Paris).
Vitessa zemire coronis subsp. n.
(Figs 7; 70, 71;-145;-1795-207.235, 28)
3, 2. 44-58 mm. Generally as described for species. Fore wing above with orange basal area large, outer
margin straight, slightly produced distad in 3 at posterior margin, in 2 broadly produced at base of cell;
subbasal spot distinct. Antemedial and postmedial fasciae narrow, variable; antemedial fascia narrower
than postmedial, greenish white, slightly diffuse medially; postmedial fascia creamy white, generally widest
in mid region, anteriorly rounded, not reaching costa, posteriorly with slight diffusion medially, not
REVISION OF VITESSIDIA & VITESSA 279
reaching posterior margin, outer margin in ¢ rounded, in 2 broadly indented between M, and M,. Vein-
lines in terminal area of ¢ extending back to Cu, well separated from postmedial fascia; in 2 far less
strongly developed. Hind wing below with pale apical patch tending to break into bars along veins.
GENITALIA ¢ (Figs 145, 179, 207). Generally as described for species, but see Remarks.
GENITALIA @ (Figs 235, 281). Generally as described for species, but see Remarks.
REMARKS. This is distinguished from previous subspecies by the much narrower white fasciae;
from V. zemire novaebritanniae by the generally broader white fasciae and a more distinct subbasal
spot in the orange basal area; from remaining subspecies by not having the orange basal area
indented, by having white apical markings on the underside of the hind wing and lacking the
white spot over the cell. In the 3 genitalia the right cornuti cluster (Fig. 207) is relatively longer
than in previous subspecies, the left cornuti cluster is just as long, but has the spines developed
only on the posterior half; on the anterior half they are reduced to sclerotized bases. In the 2
genitalia the antrum (Fig. 281) is longer than in V. zemire nephritica and laterally more heavily
sclerotized; it is broadest at posterior end, with a slight waist anteriorly. The corpus bursae has
the signa larger than in V. zemire nephritica.
BioLoGcy. Early stages and host-plant unknown. Moths collected in July and August.
DISTRIBUTION (Fig. 7). Bismarck Archipelago: Umboi Island.
MATERIAL EXAMINED
Holotype 3, Bismarck Archipelago: Rook[e] Island, viii.1913 (Meek) (BMNH).
Paratypes. Bismarck Archipelago: 3 3, 2 2 (including allotype), vii—vili, data as holotype (genitalia
slide no.’s 14716, 15137, 15293, 15294; BMNH).
Vitessa zemire novaebritanniae subsp. n.
(Figs 7, 72, 73, 146, 180, 236, 282)
3, 2. 40-54 mm. Generally as described for species. Fore wing above with orange basal area large, outer
margin straight or variably sinuate; subbasal spot much reduced. Antemedial and postmedial fasciae
very narrow, greenish white; antemedial fascia reduced to sometimes incomplete hair-line; postmedial
fascia not reaching costa or posterior margin, posteriorly with slight diffusion medially, outer margin in
both sexes broadly indented between M, and M,. Vein-lines in terminal area of ¢ reaching down to M;,
well separated from postmedial fascia; far less developed in 2. Hind wing below with pale apical markings
breaking into bars along veins at least to R;, sometimes as far as M3.
GENITALIA ¢ (Figs 146, 180). Generally as described for species, but see Remarks.
GENITALIA 2 (Figs 236, 282). Generally as described for species, but see Remarks.
REMARKS. This is distinguished from all other subspecies by the much narrower white fasciae.
Among the subspecies having large, non-indented, orange basal areas, this subspecies shows
greatest reduction in the size of the subbasal spot. Among the subspecies with white apical
markings on the underside of the hind wing, these are the most reduced in this subspecies.
Specimens from New Hannover tend to have the antemedial fascia and the subapical vein-streaks
better developed, but are not always distinguishable. In the ¢ genitalia the right cornuti cluster is
slightly smaller than in V. zemire coronis, the left cornuti cluster is shorter than the right, with
short spines present on posterior quarter of cluster, the remainder reduced to sclerotized bases.
In the 2 genitalia the antrum (Fig. 282) is similar to that of V. zemire coronis but narrower
posteriorly; the corpus bursae and signa are of the same size as in that subspecies.
BioLoGy. Early stages and host-plant unknown. Moths collected from January to March.
DISTRIBUTION (Fig. 7). Bismarck Archipelago: New Britain; New Hannover; New Ireland.
MATERIAL EXAMINED
Holotype 3, Bismarck Archipelago: New Britain, Talesea, i.1925 (A. F. Eichhorn) (genitalia slide no.
14632; BMNH).
Paratypes. Bismarck Archipelago: 3 3, 2 2 (including allotype), data as holotype (genitalia slide no.’s
14634, 15134, 15295, 15296; BMNH).
280 E. G. MUNROE & M. SHAFFER
(Meek) (BMNH; TM, Pretoria); 1 3, New Ireland, Schleinitz Mountains, 915 m (‘3000 ft’), 2.x.-15.xii.1959
(Brandt) (ANIC, Canberra).
Vitessa zemire solomonis subsp. n.
(Figs 8, 74, 75, 147, 181, 237, 283)
3, 2. 46-58 mm. Generally as described for species. Fore wing above with orange basal area reduced,
split at A, by V-shaped indentation from outer margin; subbasal spot large, sometimes joining costa.
Antemedial and postmedial fasciae very narrow, creamy white; antemedial fascia narrow in 2, slightly
short of costa and posterior margin, in 2 reduced to hair-line; postmedial fascia well short of costa and
posterior margin, slightly diffuse medially, outer margin in both sexes slightly rounded. Vein-lines in
terminal area in 3 from between R, to Cu, in 2 variable, from just a trace between R, and R; to lines
reaching between Cu, and CuP; mostly separated from postmedial fascia. Hind wing below with pale
apical marking reduced to faintest trace, but with white streak along costa from apex to about level with
medial side of postmedial fascia of fore wing; small, triangular, oblique, white spot at apex of cell,
commencing at and following lines of discocellular vein.
GENITALIA ¢ (Figs 147, 181). Generally as described for species. Left cornuti cluster absent; right cornuti
cluster short, crescent-shaped, spines longest at centre. See Remarks.
GENITALIA 9 (Figs 237, 283). Generally as described for species. Antrum laterally asymmetrical, right
lateral margin concave, left lateral margin convex. See Remarks.
REMARKS. This is distinguished from previous subspecies by having a white spot at the apex of cell
on the underside of the hind wing; by lacking a definite apical marking but having a short white
streak on the costa. On the upperside of the fore wing the deeply indented orange basal area is
shared only by V. zemire zemire, from which the present subspecies is distinguished by the
narrower postmedial fascia. V. zemire solomonis is distinguished from V. zemire cheesmanae by
the slightly narrower postmedial fascia and by the predominant tendency for the terminal vein-
lines on the upperside of the fore wing not to reach the postmedial fascia, the only exception seen
being a 2 from Guadalcanal Island. The ¢ genitalia are easily distinguished from those of the
previous subspecies by the absence of the left cornuti cluster; the 2 genitalia by the asymmetry of
the antrum (Fig. 283), though this asymmetry is less accentuated than in V. zemire cheesmanae.
BioLoGcy. Early stages and host-plant unknown. Moths collected in January, February, April,
June, November and December.
DISTRIBUTION (Fig. 8). Solomon Islands: Bougainville Island to Guadalcanal Island.
MATERIAL EXAMINED
Holotype 3, Solomon Islands: Florida I., 1.1901 (Meek) (genitalia slide no. 15297; BMNH).
Paratypes. Solomon Islands: 2 2, Bougainville, iv.1904 (Meek) (BMNH); 1 9, north side of Choiseul I.,
xil.1903 (Meek) (BMNH); 3 2, Guizo I., xi.1903 (Meek) (genitalia slide no. 15135; BMNH); 1 9, Rendova,
11.1904 (Meek) (BMNH); 6 @ (including allotype), data as holotype (BMNH); 1 2, Tugela (Woodford)
(genitalia slide no. 15298; BMNH); 1 2, Guadalcanal, Honiara, vi.1971 (H. S. Robinson) (BMNH). .
Vitessa zemire cheesmanae subsp. n.
(Figs 9, 76, 77, 148, 182, 208, 238, 284)
3, 2. 48-52 mm. Generally as described for species and for V. zemire solomonis. Fore wing above with
considerable green in the blue reflections of the dark ground colour. Orange basal area deeply indented,
subbasal spot always separated from costa. White antemedial fascia narrow but complete, slightly diffuse
medially. Postmedial fascia broad, not reaching costa or posterior margin, medially extended and slightly
diffuse, terminal margin broadly rounded and sinuate. Vein-lines in terminal area extending back to
Cuz, separated from but close to postmedial fascia. Hind wing below with white costal streak and white
marking on discocellular vein.
GENITALIA ¢ (Figs 148, 182, 208). Generally as described for species. Left cornuti cluster greatly reduced
to minute group of short spines and sclerotized spine-bases. Right cornuti cluster short, crescent-shaped,
spines longest at centre. See Remarks.
REVISION OF VITESSIDIA & VITESSA 281
GENITALIA 2 (Figs 238, 284). Generally as described for species. Antrum (Fig. 284) laterally asymmetrical.
See Remarks.
REMARKS. This is distinguished from V. zemire solomonis by the broader postmedial fascia and
by the terminal white lines being closer to the postmedial fascia. It differs from all other subspecies
of V. zemire by the presence of a white discocellular marking on the underside of the hind wing.
In the ¢ genitalia this subspecies differs from V. zemire solomonis in the presence of a small left
cornuti cluster (Fig. 208) consisting of sclerotized bases. The 9 genitalia are distinguished by the
more strongly asymmetrical, narrower and longer antrum (Fig. 284) and by the larger signa. This
subspecies and V. zemire solomonis are somewhat atypical of the species, but the general similarity
to other subspecies is so great that separate specific status is unwarranted. We take pleasure in
dedicating this subspecies to Miss L. E. Cheesman, collector of the holotype, in recognition of
her extensive contribution to the knowledge of Pacific entomology.
BioLoGcy. Early stages and host-plant unknown. Moths collected from March to May and in
August. A specimen from Efate recorded as flying at 1600 h.
DISTRIBUTION (Fig. 9). New Hebrides.
MATERIAL EXAMINED
Holotype 3, New Hebrides: Malekula, Ounua [Bay, 16°14’ S, 167°36’ E, nr Bansarik], iv-v.1929 (Cheesman)
(genitalia slide no. 14630; BMNH).
Paratypes. Hew Hebrides: 1 9° (allotype), data as holotype (genitalia slide no. 14631; BMNH); 1 9,
Efate, near Vila, Montmartre, flying 1600 h, 9.iii.1974 (Maddison) (genitalia slide no. 15299; BMNH);
6 3, 14 2, Espiritu Santo, south coast, Narango, v—vii.1960 (Brandt) (ANIC, Canberra; CNC, Ottawa);
2 3, Espiritu Santo I. (SW), above Namatasopa [15°31’ S, 166°49’ E] 400 m, viii.1957 (Gressitt) (BPBM,
Honolulu); 1 3, [no exact locality] (HMS Dart) (BMNH).
Vitessa vitialis Hampson, 1906
(Figs 10, 78-80, 149, 183, 209, 239, 285)
Vitessa vitialis Hampson, 1906, Ann. Mag. nat. Hist. (7) 17: 217. Holotype 2, Fist (Crowley Bequest)
(genitalia slide no. 15300; BMNH) [examined].
3, 2. 54-60 mm. Frons smoothly scaled, glossy blue-black. Vertex with erect bushy orange scaling,
extending forward between antennae. Eye and ocellus black. Chaetosema with orange setae. Antenna with
pectinations in ¢ very short, black; dorsal scaling of antennal shaft glossy black, except basal half of
apical third white. Labial palpus with first and second segments orange, third segment black. Maxillary
palpus basally black, distally orange. Basal scaling of proboscis blue-black with violaceous reflections.
Neck region orange. Thorax above: orange, with narrow, medial, longitudinal blue-black stripe; patagium
glossy blue-black ; tegula orange with anterolateral blue-black spot. Abdomen above: matt black, a narrow
white terminal line on each segment, except last segment with posterior margin orange, anal tuft orange.
Body below: glossy blue-black; pectus orange; white terminal lines on abdominal segments broader than
above, last segment with posterior margin orange, anal tuft orange. Legs in 3 with most parts of coxae,
femora and tibiae covered with bushy orange scales, remaining parts dark fuscous; legs in 2, glossy green-
black, ventral surfaces of mid and hind femora white, tibiae white-tipped. Fore wing above: base blue-
black with strong gloss; subbasal area orange, reaching Cu, short of costa, finely edged with weakly
glossy blue-black; subbasal fascia blue-black with strong gloss; yellowish white antemedial fascia erect
narrow, short of costa and posterior margin, finely diffuse medially; medial fascia blue-black; yellowish
white postmedial fascia slightly oblique, pointing towards posterior margin well basad of tornus, narrow,
widest at centre, medial and terminal margins irregular; terminal area blue-black, yellowish white vein-
lines short of postmedial fascia; dark areas of wing with strong greenish hue; fringe basally dark grey,
distally white, flecked with dark grey. Hind wing above blue-black with weak violaceous reflections,
cubital and anal area covered with long black hairs; fringe basally dark grey, distally white flecked with
dark grey. Fore wing below: blue-black, in ¢ with strong diffusion of orange at base, finely diffuse on
costa and posterior margin; dark areas with violaceous reflections, weak greenish hue especially at costa;
retinaculum dark grey; yellowish white postmedial fascia as above but more diffuse; vein-lines short,
apical; fringe as above. Hind wing below as above, apical vein-lines fine; in ¢ basal and discal areas with
orange suffusion, strongest on veins, especially discocellular vein; 2 without orange suffusion but with
minute patch of white scales on angle of discocellular.
282 E. G. MUNROE & M. SHAFFER
GENITALIA ¢ (Figs 149, 183, 209). Uncus wide, depressed, concave ventrally like an inverted trough,
decurved then upcurved to tip, bearing lateral setae for most of its length. Gnathos arms broadening
medially, flattened medial process deep at base, acutely sickle-shaped, tip finely acuminate. Transtilla
shallowly V-shaped, medially narrow. Tegumen with medially broadened narrow posterior sclerite;
anterior sclerite medially upturned, with broad keyhole-shaped emargination. Vinculum wide, laterally
broadened, base strongly concave. Valve long, very narrow, costa inflated at base, apex narrowly rounded;
with narrow, central, longitudinal sclerotized band; mesal surface with long oblique striations; posterior
margin heavily sclerotized, especially at broadened base; long curved club-shaped clasper at mid length of
posterior margin, on widened oblique base, extending posteriorly at right angles then curving basad. Juxta
wide; lateral sclerites broad, heavily sclerotized, each bearing short wide dorsal and ventral processes,
posteromedially directed; anellus (Fig. 183) supported by dorsal juxta process, with large, lateral, antero-
medially directed sclerites, bearing large anteriorly directed spines; manica supported by sclerotized
bands continuous with lateral sclerites. Aedeagus long, distally widened, base narrowly rounded, ductus
ejaculatorius subbasal; spines of vesica mainly basal, vesica ventrally bulbous at distal end of aedeagus
and transversely rugose, vesica with ventrolateral left and dorsolateral right cornuti clusters (Fig. 209),
longitudinally rugose dorsally between cornuti clusters; right cluster longer than left, with longer spines.
GENITALIA @ (Figs 239, 285). Ovipositor lobes dorsally fused, dorsally narrowly produced, densely setose,
setae longer on margins; spiny setae not easily seen. Posterior apophysis long, straight, distally widening,
spatulate at tip. Anterior apophysis longer, thicker, sinuate, with inwardly curved spatulate tip. Posterior
ostial sclerite (Fig. 285) broad, heavily sclerotized, posterior margin with small V-shaped medial incision;
small anteromedial bullae; sulcus wide, margins separated. Anterior margin of ostium mostly
membranous, inner part weakly sclerotized, wrinkled, laterally minutely denticulate. Antrum short,
weakly sclerotized, except for a small anteromedial sclerotized area. Ductus bursae irregularly swollen
posteriorly; narrow anterior part minutely spined. Corpus bursae large, oval, membranous, with small,
paired, heavily sclerotized, depressed, denticulate signa.
REMARKS. This species bears considerable resemblance to V. zemire, but otherwise it can be
confused only with V. tamsi from Samoa. It differs from V. zemire in the markings of the fore
wing; the orange area is subbasal, the white fasciae are distinctly yellowish, the very wide terminal
area has almost equally long pale lines on most veins. The 3 is unlike other species except V.
tamsi, in the presence of massive orange scaling (androconia ?) on the femora and tibiae and
unlike other species, including V. tamsi, with sex-limited orange scaling on the underside of the
hind wing. The 3 genitalia are unlike those of any other species except V. tamsi, which shares the
wide, flattened uncus, the long narrow valve and the club-shaped clasper. In V. vitialis the transtilla
is V-shaped, the clasper lacks a ridge from base, the juxta is narrower and the left cornuti cluster
is larger than in V. tamsi. The @ genitalia are distinctive in the shape of the swollen posterior end
of ductus bursae and in the much reduced antrum, but those of V. tamsi are almost identical
(see Remarks under that species). There is some individual variation in external and genital
characters. The two specimens from the Lau group, though worn, appear to have the postmedial
fascia of the fore wing wider, but whether this is a significant difference remains to be seen. This
is the only known species in the Fiji Islands.
BioLoGy. Early stages and host-plant unknown. Moths collected in January, from June to
October and in December.
DISTRIBUTION (Fig. 10). Fiji: Vanua Levu, Viti Levu and Lau Group.
MATERIAL EXAMINED
Fiji: 6 2, Vanua Levu, Navonu, Forest Area Niuvundi, ix.1975 (Maddison) (BMNH; KRS, Fiji); 1 2,
Viti Levu, Vuni[n]dawa, vii.1932, caught in butter dish at tea (Phillips) (BMNH); 1 2, V. L., Nandarivatu,
x.1937 (Valentine) (BPBM, Honolulu); 1 9, V. L., Nandarivatu, 800 m, ix.1955 (Simmonds) (BMNH);
1 9, V. L., Nandarivatu, vi.1968 (H. S. Robinson) (BMNH); 1 3,1 9, V. L., Nandarivatu, ix.1974 (G. S.
Robinson) (BMNH); 2 9, V. L., Savura Creek, 1966-67, 1.1967 (H. S. Robinson) (BMNH); 1 2, V. L.,
Savura Creek, vii.1969 (H. S. & G. S. Robinson) (BMNH); 14, 1 9, V. L., Tholoisuva, xii.1958 (Simmonds)
(BMNH); 1 2, V. L., Colo-i-suva, Malaise trap, iii.1963 ( Yoshimoto) (BPBM, Honolulu); 1 9, V. L.,
Coloi Suva [Tholoisuva], viii.1968 (H. S.&G.S. Robinson) (BMNH); 1 9, V. L., Suva, vii.1954 (Simmonds)
(BMNH); 1 3, Lau Group, Namuka [-i-Lau], xii.1924 (Bryan) (BPBM, Honolulu); 1 9, Ongea, viii.1924
(Bryan) (BPBM, Honolulu).
REVISION OF VITESSIDIA & VITESSA 283
Vitessa tamsi sp. n.
(Figs 11, 42, 43, 81, 240, 286)
3, 2. 52-58 mm. In general as described for V. vitialis. Tegula orange with anterior blue-black spot.
Thorax above: orange with wide, medial, longitudinal blue-black stripe. Fore wing above with yellowish
white postmedial fascia, in 2 half as wide as long, wider in 3, medially rounded, not reaching costa or
posterior margin, very oblique, pointing towards tornus; orange subbasal area narrow. Fore wing below:
postmedial fascia as wide as above; antemedial fascia faintly indicated. Hind wing below: white spot on
discocellular vein varying from a few scales to a spot over half width of cell. Wings of 3 below like those
of 2, without orange scaling, and with white discocellular marking.
Fig. 42 Vitessa tamsi sp. n., 3 allotype, Western Samoa.
GENITALIA ¢ (Fig. 43). In general as described for V. vitia/is. Transtilla nearly straight. Genitalia damaged,
valve tips missing. Club-shaped clasper with short, straight, raised ridge extending from base towards
base of valve. Anterior portion of juxta very broad. Aedeagus with slightly reduced left cornuti cluster.
GENITALIA 2 (Figs 240, 286). In general as described for V. vitialis. Posterior apophysis short, slightly
sinuate, distally widening and spatulate at tip. Anterior apophysis short, thicker, straight, distally tapering,
tip spatulate. Posterior ostial sclerite (Fig. 286) narrow, strongly sclerotized, posterior margin with small
V-shaped medial incision; anterior margin broadly upturned medially, small anteromedial bullae, wide
sulcus between, margins separated; sclerite slightly bulging laterally. Antrum sclerotization very weak,
hardly visible.
REMARKS. This species is very closely related to V. vitialis, of which it is the Samoan vicariant.
It is distinguished externally by the broader and more oblique postmedial fascia on the upperside
of the fore wing pointing to the tornus, not to the posterior margin basad of the tornus as in V.
vitialis, and by the slightly narrower orange subbasal area. The postmedial fascia is noticeably
wider in the 3 than in the 2. The tegula bears an anterior blue-black spot rather than an antero-
lateral spot as in V. vitialis. V. tamsi shares with V. vitialis the strong greenish hue over all dark
areas on wings and legs. In the ¢ the underside of the wings lacks the extensive diffusion of orange
scaling seen in the § of V. vitialis. The 3 genitalia differ from those of V. vitialis by the straighter
transtilla, the short raised ridge extending from the base of the clasper, by the broader anterior
portion of the juxta, and by the smaller left cornuti cluster of the vesica.
In the 2 genitalia the posterior ostial sclerite is much narrower medially in V. tamsi, almost
making two triangular areas. The antrum is hardly sclerotized, whereas in V. vitialis a small
triangular sclerotized plate is present. The ductus bursae is longer than the bursa in V. tamsi,
shorter in V. vitialis. These differences seem of specific rather than subspecific value. We take
pleasure in dedicating this species to our esteemed colleague Mr W. H. T. Tams, in recognition
of his contributions to the knowledge of the moths of Samoa.
BioLoGy. Early stages and host-plant unknown. Moths collected in January, March and
September.
DISTRIBUTION (Fig. 11). Western Samoa; so far known only from Upolu.
284 E. G. MUNROE & M. SHAFFER
Fig. 43 ¢ genitalia. Vitessa tamsi sp. n.
MATERIAL EXAMINED
Holotype 2, Samoa: Upolu, Moamoa, 2.1x.1951 (Armstrong) (genitalia slide no. 14629; BMNH).
Paratypes. Samoa: | ¢ (allotype), 1 2, Samoa, 1914-18 (Tate) (genitalia slide no. NMNZ 141 =15301/a,
3, Shaffer; NMNZ, Wellington); 1 2, Upolu, Hellesoi’s Paddock, 500 m(‘1600 ft’), 15.iii.1953 (Armstrong)
(genitalia slide no. 15301; BMNH); 1 9, Upolu, Tialevi, 10.1.1967 (Catley) (BMNH).
The hieratica-group
DIAGNOsIS. Fore wing above with orange subbasal area and two whitish transverse fasciae, the latter
crowded closer together near medial third of wing than in other groups with black and white hind wing;
antemedial fascia complete, postmedial fascia undivided, narrowly oval, not reaching costa or posterior
margin as in the few species of succeeding groups with undivided postmedial fasciae. Hind wing with pale
medial area, tending to be diffuse on a fuscous background. ¢ genitalia with uncus subpyriform, bearing
relatively short, poorly differentiated, laterally directed setae, contrasting with the basally broad, distally
rod-like uncus with strong laterally directed setae found in other groups with white areas on hind wing.
Valve with oblique serrate clasper and intersecting ridge. Anellus with heavily sclerotized lateral arms in
REVISION OF VITESSIDIA & VITESSA 285
the form of thick straight bars not bearing spines; dorsal medial catena very large, oval and heavily spined;
three cornuti clusters in aedeagus. 2 genitalia with posterior sclerite of ostial chamber W-shaped, anterior
sclerite semicircular and strongly wrinkled; corpus bursae without signa.
Vitessa hieratica Swinhoe, 1900
(Figs 13, 27, 35, 82, 83, 150, 184, 210, 241, 287)
Vitessa hieratica Swinhoe, 1900, Cat. Eastern and Australian Lepid. Het. Coll. Oxford Univ. Mus. 2 : 430,
pl. 6, fig. 22. Holotype 3, BORNEO: Sarawak (Wallace) (Type no. 873; genitalia slide no. Oxford 717-
1977, Shaffer; UM, Oxford) [examined].
3, °. 32-40 mm. Frons and vertex orange. Eye dark brown. Ocellus minute, oval, yellow. Chaetosema with
pale yellowish setae. Antenna black, anterior surface of scape orange; dorsal scaling of nearly half apical
half of shaft white. Labial palpus: first and second segments orange, third segment black. Maxillary
palpus basally black, terminally orange. Basal scaling of proboscis black, ventrally flecked with orange.
Neck region orange. Thorax above: orange with wide, medial, longitudinal, blue-black stripe; patagium
orange with posteromedial blue-black spot; tegula orange with long anterior and terminal blue-black
spots. Abdomen above: blue-black with broad whitish grey terminal lines on segments, anal tuft orange.
Body below: fuscous; pectus orange; abdomen as on upperside. Legs fuscous; fore coxa with large orange
patch bearing central blue-black spot; mid and hind coxae, femora and tibiae banded with whitish grey.
Fore wing short and narrow; above: basal area blue-black with small patch of white at base of posterior
margin; subbasal area of irregular width, basally orange and medially white, slightly short of costa but
reaching posterior margin, both margins acutely angled at base of cell, sharply widened again to region of
CuP, then truncated and widened on reaching posterior margin. Antemedial fascia white, reaching margins
of wing, at posterior margin narrowly joined to subbasal area, the fascia wide, medially radiating in
narrow lines towards postmedial fascia at R stem, Cu,, A, and posterior margin, line at Cu touching
postmedial fascia. Postmedial fascia white, large, oval, not reaching margins of wing. Dark areas on wings
blue-black ; terminal area wide with white vein-lines. Fringe greyish fuscous. Hind wing above with basal
area whitish, variably suffused with fuscous from margins, costal and distal margins diffusely blue-black.
Fringe basally fuscous, distally white. Fore wing below blue-black; retinaculum light grey; postmedial
fascia smaller than on upperside; apex and anterior part of fringe narrowly whitish grey, posterior margin
white. Hind wing below: as on upperside but with blue-black margin demarcated from whitish basal area.
Hind wing of 2 with well-demarcated margins above and below.
GENITALIA ¢ (Figs 27, 35, 150, 184, 210). Uncus short, laterally swollen at base, distally tapering and with
laterally directed setae, tip broadly acuminate. Gnathos long, basal arms nearly straight, tapering distally
and converging at an acute angle, medial process strong, tapering, tip hooked. Transtilla strong, shallowly
V-shaped, of even width. Tegumen anteriorly directed, posterior and anterior sclerites constricted, anterior
margin of anterior sclerite with large posteriorly directed keyhole-shaped emargination; posterior sclerite
weak. Vinculum broad, with base slightly concave. Valve long and wide, costa heavily sclerotized and
sinuate, valve broadest postmedially, apex narrowly rounded; clasper small, angulate with a curved carina.
Juxta with narrow lateral sclerites bearing long dorsal and ventral processes; ventral process supporting
base of anellus (Fig. 184); dorsal process (Fig. 27) posteriorly setose and connected to long, straight, heavily
sclerotized, unspined, lateral sclerite on anellus, terminally connected to flat, wide, sclerotized bands on
manica; long dorsal catena (Fig. 35), narrow, oval, heavily spined. Aedeagus of moderate length, distally
expanded, base narrowly rounded; ductus ejaculatorius subbasal; spines on vesica dorsally and ventrally,
vesica on ventral surface at distal end of aedeagus bulbous and transversely rugose; dorsal side of vesica
between cornuti clusters longitudinally rugose. Three cornuti clusters (Fig. 210); two long curved lateral
proximal clusters, right slightly longer than left and with shorter spines; single distal cluster minute with
about four spines.
GENITALIA 2 (Figs 241, 287). Ovipositor lobes high, narrow, dorsally fused and prolonged posteriad;
posterior surface densely setose, longer setae on margin, spiny setae visible at 100 magnification.
Posterior apophysis short, narrow, angled slightly near tip. Anterior apophysis short, narrow, angled
slightly at base and near tip. Posterior ostial sclerite (Fig. 287) long and wide, with three well-defined and
highly sclerotized border regions laterally and anteriorly, a central roughly circular area well defined but
not sharply demarcated in sclerotization. Anterior sclerite membranous with sclerotized incurved wrinkled
base, laterally denticulate. Antrum broadly oval. Ductus bursae with funnel-shaped swelling posteriorly,
lightly spinose. Corpus bursae large, membranous, oval; signa absent.
286 E. G. MUNROE & M. SHAFFER
REMARKS. This rare species is easily distinguished externally from V. suradeva and other species
with generally similar maculation by the undivided oval postmedial white fascia of the fore wing,
connected to the antemedial white fascia by a white streak along Cu,. In other species of similar
appearance the postmedial fascia is bi- or tripartite and is usually not connected to the antemedial
white fascia along Cu,. The species is distinct from all other species in both 3 and 9 genitalia, as
indicated in the diagnosis of the species-group.
BioLoGy. Early stages and host-plant unknown. Moths collected in January, February, July to
September and November.
DISTRIBUTION (Fig. 13). Sumatra; Borneo: Sarawak, Sabah.
MATERIAL EXAMINED
Sumatra: 1 9, Benkulen [Bengkulu] [3°47’S, 102°15’ E], xi.1921 (Brooks) (TM, Pretoria); 1 °, Lebong
Tandai [3°02’S, 101°56’ E], i.1922 (Brooks) (BMNH). Borneo: 1 3, Sarawak, Mt Matang [1°36’ N,
110°11’ E] (Mjoberg) (BMNH); | 9, S., Bidi, 1907-1908 (Brooks) (TM, Pretoria); 1 2, S., Mt Penrissen
[1°07’ N, 110°13’ E] (Janse) (TM, Pretoria); 2 9, S., Kuching, Semongok, secondary forest, ii.1976
(Classey) (BMNH); 2 2, S., Semongok, 19.3 km (‘12 mls’) south of Kuching, iii, v.1974 (Earnshaw)
(CNC, Ottawa); 2 g, S., Gunong Mulu Nat. Park, Mulu, 1790 m, 452463 lower montane (moss) forest,
MV - canopy, 1.1978 (Holloway et al.) (BMNH); 1 3, Sabah, Mt Kinabalu, ‘Power Station’, 1950 m,
vii-1x.1965 (Holloway) (BMNH).
The stettina-group
DIAGNOsIs. Fore wing above with two transverse whitish fasciae and an orange subbasal area; the post-
medial white fascia usually and the antemedial fascia sometimes divided into two or more spots. Hind wing
with white or whitish discal area and black terminal margin, the latter narrowing posteriorly. In the 3
genitalia uncus differentiated into a short, relatively wide basal portion and a long, slender, rod-like,
down-turned, then up-turned, distal portion, with conspicuous, laterally directed setae. Valve long, with
regular fluting and weak angular clasper. Anellus with undifferentiated spineless sclerotization at junction
with dorsal process of juxta. Vesica with three cornuti clusters, but, unlike the condition in previous
groups, one cluster basal and two distal. In the 9 genitalia the posterior ostial sclerite short, wide; anterior
sclerite wrinkled and laterally differentiated into three portions; antrum short and wide, merging in
outline with funnel-shaped posterior end of ductus bursae. Corpus bursae long and narrow, signa absent.
Distinguished from the hieratica-group by the shape of the whitish fasciae on the fore wing, by the differen-
tiated uncus of the 3, and by the structure of the ostial chamber in the 2. Differing from the hemiallactis-,
plumosa- and pyraliata-groups by the weak clasper of the valve and from the suradeva-group by the oblique
position and more rounded outline of the valve. Differing from all other groups by the anellus of the 3
lacking a catena, and the corpus bursae of the 9 lacking signa. The @ is distinguished from the hemiallactis-
group by having the posterior margin of the seventh abdominal sternite straight or concave, not convex.
Vitessa philippina sp. n.
(Figs 14, 84, 242, 288)
3g. Unknown.
2. 48 mm. Frons and vertex orange. Eye and ocellus dark brown. Chaetosema with yellowish setae.
Antenna black, scape basally orange, probably with some white on posterior surface of apical third of
shaft (antennae of holotype badly damaged). Labial palpus pale orange from base to near end of second
segment, tip of second, all of third segment black. Maxillary palpus basally black, distally pale orange.
Basal scaling of proboscis black. Neck region pale orange. Thorax above: black; patagium blue-black,
bordered with pale orange; tegula blue-black, anteromedially with faint border of pale orange. Abdomen
above blue-black, with narrow whitish grey terminal lines on segments, anal tuft orange. Body below black,
but with white terminal lines on segments wider; pectus pale orange; legs black, banded with white. Fore
wing long and narrow; above: ground colour blue-black; a subbasal orange spot, roughly triangular,
anteriorly not reaching costa, posteriorly extending to A,; white antemedial fascia divided, small spot
behind costa, larger triangular spot between Cu and A,; white postmedial fascia oblique, divided into two
spots, a circular spot on discocellular vein and a more distal, smaller, triangular spot on CuP, some white
scaling anterior to terminal end of this spot; in terminal area white vein-lines from R, to M; or Cu, well
separated from postmedial fascia; fringe dark grey. Hind wing above with base, costa and termen broadly
black, anal margin with thin black fringe, narrow black scaling along some veins from base, discal area
and middle of anal margin white, faintly nacreous. Fore wing below: postmedial fascia almost as above,
REVISION OF VITESSIDIA & VITESSA 287
faint traces of antemedial fascia, posterior margin white, remainder black; retinaculum dark grey;
fringe apically white, dark grey below. Hind wing below as above.
GENITALIA ° (Figs 242, 288). High, narrow ovipositor lobes, dorsally fused and produced posteriorly,
densely setose, with longer setae at margin; spiny setae visible at x 100 magnification. Posterior apophysis
long, slender and slightly curved. Anterior apophysis similar but thicker and longer. Posterior ostial
sclerite (Fig. 288) with irregular transverse wrinkles, posterior margin medially concave, anterior margin
deeply incurved but with medial area strongly convex. Anterior sclerite with posterior margin of even
width, medially with weak folds, whole margin denticulated, somewhat differentiated into three sections,
protruded posterolaterally and rounded, anterior margin membranous, laterally denticulate. Antrum
broad and funnel-shaped. Posterior end of ductus bursae swollen, funnel-shaped; short narrow portion
of ductus spinose, merging gradually into long, narrow corpus bursae; signa absent.
REMARKS. This has fewer white vein-lines in the terminal area of the fore wing than any other
species of the group. In general reduction of markings it closely resembles V. ternatica of the
hemiallactis-group, but it is distinguishable by genitalia. In the Philippines, V. pyraliata
triangulifera is very similar in appearance, but V. philippina is distinguishable by the larger orange
subbasal area of the upperside of the fore wing and by lacking white vein-lines on the apical
region of the underside of the hind wing. The 9 genitalia of the species in this group are very
similar, but in no other species are the ovipositor lobes so acutely narrowed posteriorly.
BioLoGy. Early stages and host-plant unknown. Moths collected in April, September and
October.
DISTRIBUTION (Fig. 14). Philippines: Luzon, Mindanao.
MATERIAL EXAMINED
Holotype 2, Philippines: Luzon, Manila, 12.ix.1912 (Wileman) (genitalia slide no. 14701; BMNH).
Paratypes. Philippines: 1 2°, Mindanao, Davao Province, 30.iv.1930 (Clagg) (CM, Pittsburgh); 1 9,
Mindanao, 24-13 km (‘15 mls’) SW. Davao, Mt Apo School, 22-31.x.1965 (Davis) (NMNH, Washington).
Vitessa stettina Swinhoe, 1906
(Figs 15, 28, 36, 85, 86, 151, 185, 211, 243, 289)
Vitessa stettina Swinhoe, 1906, Ann. Mag. nat. Hist. (7) 18 : 412. LECTOTYPE 2, NEw Guinea: Papua,
Granville [Port Moresby] 845 (genitalia slide no. 15305; BMNH), here designated [examined].
3, 2. 46-56 mm. Frons and vertex orange. Eye and ocellus brown. Chaetosema with pale yellowish setae.
Antenna black; ventral surface of scape mainly or wholly orange; dorsal scaling of basal half of apical
third of shaft white. Labial palpus with first two segments orange, third segment black. Maxillary palpus
basally black, distally orange. Basal scaling of proboscis black. Thorax above: orange with central round
blue-black spot; patagium orange with posteromedial glossy blue-black spot; tegula orange with
anterolateral glossy blue-black spot. Abdomen above: blue-black with narrow whitish grey terminal lines
on segments, anal tuft orange. Body below: blue-black; pectus orange; abdomen as above with wider
terminal lines. Legs blue-black, with faint greenish hue in places; posterior surfaces of fore coxa, mid and
hind femora whitish grey; whitish grey bands on mid and hind tibiae. Fore wing long and of moderate
width; above: ground colour glossy blue-black, some orange scales at costa; orange subbasal area tapering
to near posterior margin in 9, not passing A, in 3; whitish grey antemedial fascia wide, complete, almost or
quite reaching posterior margin, separated from costa by a narrow line of ground colour, basal margin
erect, straight, medial margin irregular, diffuse, sometimes joining postmedial fascia, especially along
posterior margin; whitish grey postmedial fascia wide, complete, rounded anteriorly, sometimes reaching
posterior margin, sometimes indented at Cu,, terminal margin sinuate; terminal area with all veins whitish
grey; fringe glossy dark grey. Hind wing above: base, costa and termen blue-black with violaceous
reflections, white medial area extending broadly to anal margin; fringe glossy dark grey. Fore wing below:
bluish fuscous with violaceous reflections; reduced whitish grey postmedial fascia; few whitish grey apical
vein-lines; posterior margin white; retinaculum dark grey; fringe dark grey. Hind wing below as above,
apical part of costa, few apical vein-lines and anterior portion of fringe whitish grey.
GENITALIA ¢ (Figs 28, 36, 151, 185, 211). Uncus long, narrow, basally widened, distally tapering with long
laterally directed setae, down-curved then up-curved, apex sharp. Gnathos with flat lateral arms con-
verging gradually to the rounded, hook-like, acuminate medial process. Transtilla gently curved to centre,
288 E. G. MUNROE & M. SHAFFER
medially widened. Tegumen: posterior sclerite with wide rounded shoulders, anterior sclerite posteriorly
directed medially with large keyhole-shaped medial emargination. Vinculum broad, basally truncated.
Valve long, narrow, costa inflated, apex moderately rounded, termen broadly rounded, posterior margin
almost straight for most of length; narrow longitudinal striations on mesal surface; clasper short, oblique,
slightly separated from curved carina, a small serrate ridge ending ventrally before margin in a basally
directed tooth-like process. Juxta with large narrow lateral sclerites, ventral process strongly upturned,
flat, narrow and twisted, supporting base of anellus (Fig. 185); dorsal process shorter, wider, bearing
setae, connected to sinuate, distally reticulated sclerite supporting side of anellus (Fig. 28); medial to tips
of sclerites are two small irregular sclerotized and ridged areas (Fig. 36); in continuation with sclerites,
flat sclerotized bands supporting base of manica. Aedeagus of moderate length, distally expanded, base
narrowly rounded; ductus ejaculatorius subbasal. Vesica with spines spreading ventrally to distal end of
aedeagus, here the vesica bulbous and transversely rugose; vesica wide to just beyond area of cornuti,
membrane between cornuti clusters longitudinally rugose; cornuti (Fig. 211) in three clusters, a single small
basal cluster with tight pack of small spines, two larger distal clusters with longer spines, left cluster shorter
than right.
GENITALIA 2 (Figs 243, 289). Ovipositor lobes high, narrow, dorsally fusing, posteriorly produced;
posterior surface densely setose, longer setae on margin, spiny setae visible at x 100 magnification.
Posterior apophysis long, slender, weakly curved, blunt; anterior apophysis longer, thicker, blunt;
Posterior ostial sclerite (Fig. 289) broad, longitudinally wrinkled, posterior margin with medial thickened
bar, anterior margin medially upturned, with prominent curved flap, margin finely denticulated; anterior
sclerite broad, posterior margin concave, slightly wrinkled, denticulate, differentiated into three slightly
overlapping denticulate plates, each rounded posteriorly, lateral plates posterolaterally protruded; antrum
wide and fairly long, conforming with swollen end of ductus bursae, cup-shaped; ductus bursae minutely
spinose, short, merging imperceptibly with elongated narrow corpus bursae; signum absent.
REMARKS. This species most closely resembles externally the superficially almost identical V.
intermedia of the hemiallactis-group, with which it flies, at least at higher altitudes, in New Guinea.
It can be distinguished from V. intermedia by having less diffusion of white into the medial area
of the fore wing; in V. intermedia the fusion between the white fasciae is more extensive, showing
little of the blue-black medial area. The species-group characters of the genitalia are of course
diagnostic. In the 2 a quick diagnosis for all the stettina-group can be made by dry examination
of the posterior margin of the ventral posterior sternite; this is lightly sclerotized and fairly
straight in the stettina-group, heavily sclerotized and convex in the hemiallactis-group, showing as
pale yellowish brown in the former and dark reddish brown in the latter, the colours being
enhanced by using toluene. V. stettina is distinct from others in the group in the greater develop-
ment of the white fore wing fasciae. It is distinct from all other species in the $ genitalia, as far as
known, by the shape of the valve and clasper and especially by the absence of dorsal catena spines
on the anellus; and in the 2 genitalia by the shape of the anterior ostial sclerite and by the absence
of signa.
BioLoGy. Early stages and host-plant unknown. Moths collected in most months of the year.
DISTRIBUTION (Fig. 15). Waigeo; New Guinea: Irian Jaya, Papua New Guinea.
MATERIAL EXAMINED
Paralectotypes. New Guinea: 1 2, Papua New Guinea, data as lectotype (genitalia slide no. 15321; BMNH);
5 3, 6 2, P. N. G., Mount Kebea, 1100-1850 m, iii—viii.1903 (A. E. Pratt) (BMNH). [1 3, 1 9, see under
paratype series of Vitessa intermedia (p. 299).]
Waigeo: 1 3, 1 2, 1904 (Waterstradt) (BMNH). New Guinea: 1 9, Irian Jaya, Ninay Valley, Central
Arfak Mts, 1050 m, ii-iii.1909 (Pratt) (BMNH); 1 4, 2 9, I. J., Nomnagihe, 40-23 km south of Wangaar,
600 m, i-ii.1921 (C. F. & J. Pratt) (TM, Pretoria); 1 9, I. J., Hollandia [Sukarnapura], iii.1960 (Maa)
(BPBM, Honolulu); 1 9, I. J., Hollandia area, Sentani, 100m, vi.1959 (Gressitt & Maa) (BPBM,
Honolulu); 1 9, I. J., Waris, S. of Hollandia, 450-500 m, viii.1959 (Maa) (BPBM, Honolulu); 1 3, no
data (Janse) (TM, Pretoria); 4 3, 3 2, Papua New Guinea, Sepik District, Bainyik, 300 m, x—xi.1957
(Munroe & Holland) (CNC, Ottawa); 4 9, P. N. G., Ekeikei, 450 m, i-iv.1903 (Pratt) (BMNH); 1 2,
P. N. G., Mount Kebea, 1100-1850 m, iii—viii.1903 (A. E. Pratt) (CMAG, Birmingham); 1 9, P. N. G.,
Mafalu, 1850 m, viii.1903 (Pratt) (BMNH); 1 92, P. N. G., Babooni, 1100 m, ix.1903 (Pratt) (BMNH);
37 3, 16 2, P. N. G., Sogeri Plateau, Mageri, viii.1957 (Munroe & Holland) (CNC, Ottawa); 2 3, 2 9,
P. N. G., Sogeri Plateau, Bisianumu, viii.1957 (Munroe & Holland) (CNC, Ottawa); 1 3, P. N. G., Port
REVISION OF VITESSIDIA & VITESSA 289
Moresby, Mt Lawes, 400 m, iii-v.1967 (Brandt) (ANIC, Canberra); 1 3, 2 2, P. N. G., Hydrographer
Vitessa hollandi sp. n.
(Figs 16, 17, 87-90, 152, 153, 186, 187, 244, 245, 290, 291)
3, 2. 36-50 mm. As in V. stettina, except as follows. Only base of scape ventrally orange. Thorax above:
variable, orange, yellowish white or greyish white with blue-black medial spot variable in size; patagium
orange with large posteromedial glossy blue-black spot; tegula glossy blue-black, laterally orange.
Abdomen above with narrow whitish grey terminal lines on first three segments only; below with broad
whitish grey terminal lines on most segments. Fore wing appearing narrower; upperside with large orange
subbasal area not reaching costa or passing A,, basally acutely angled, medially obliquely curved. Ante-
medial fascia white, basal edge nearly straight and erect, slightly short of both costa and posterior margin,
broadly indented or divided at cell, medially broadened and diffuse; postmedial fascia broad, divided into
two spots at about CuP, upper spot large and round, lower spot triangular, sometimes with small white
spot terminally above, regularly followed by a streak on posterior margin, often fusing with medial
extension of antemedial fascia. Hind wing above with white medial area variably extended to anal margin,
anal margin with thin black scaling. Fore wing below: postmedial fascia complete and obliquely oval, or
divided at CuP, spots as on upperside.
GENITALIA 3 (Figs 152, 153, 186, 187). In general as described for V. stettina. Valve apex narrowly rounded,
termen short, posterior margin broadly rounded.
GENITALIA ° (Figs 244, 245, 290, 291). In general as described for V. stettina, but see Remarks.
REMARKS. This species is nearly indistinguishable in genitalia from the largely allopatric V.
stettina; in the 3 genitalia the valve apex is more narrowly rounded and the termen more acute;
in the Q genitalia the lateral margins of the antrum (Figs 290, 291) and the swollen end of the
ductus bursae are more oblique than in V. stettina. The antenna appears to be more slender in
both sexes in the present species. Externally, the reduced pale markings of the upperside of the
fore wing are distinctive from V. stettina, but are deceptively similar to those of V. hemiallactis
hemiallactis of the hemiallactis-group, with which V. hollandi flies. The posteriorly tapering
terminal band of the hind wing usually distinguishes the present species from V. hemiallactis
and the genitalia are very distinct in both sexes (see species-group diagnoses, also Remarks under
V. stettina). V. hollandi is superficially indistinguishable from V. gemina of the pyraliata-group
but again differs in genitalia (see species-group diagnosis). We recognize two subspecies, one
from New Guinea and the Kepulauan Schouten, the other from Sulawesi. Limited material
from the Moluccas and New Britain may represent further subspecies. We take pleasure in
dedicating this species to Dr G. P. Holland, co-collector of the holotype, and companion of the
senior author in a highly productive collecting expedition to Papua New Guinea.
BioLoGy. Early stages and host-plant unknown. Moths collected in almost every month of the
year.
DISTRIBUTION (Figs 16, 17). Sulawesi; Moluccas; New Guinea; Kepulauan Schouten; New
Britain.
Vitessa hollandi wallacealis subsp. n.
(Figs 17, 87, 88, 152, 186, 244, 290)
3, 2. 36-46 mm. Generally as described for species. Thorax above: anteriorly orange, posteriorly whitish
grey, large arrow-shaped medial blue-black spot; patagium orange with very large posteromedial glossy
blue-black spot; tegula glossy blue-black, laterally yellowish white. Fore wing above: subbasal area
orange tinged with yellow; antemedial fascia white, divided at cell into two spots in ¢ and mostly in 9;
postmedial fascia divided into two spots at CuP; terminal white vein-lines very weak posterior to Cm,
touching postmedial fascia at Cu,. Hind wing above: blue-black marginal band wide, its basal margin less
strongly concave than in nominate subspecies, basal white area restricted. Fore wing below: postmedial
fascia with strong round spot but weak triangular spot; white apical lines hardly visible. Hind wing below:
white apical lines hardly visible.
290 E. G. MUNROE & M. SHAFFER
GENITALIA ¢ (Figs 152, 186). Generally as described for species. Valve narrow; uncus gradually tapering
from basal section.
GENITALIA &° (Figs 244, 290). As described for species.
REMARKS. This is distinguished from the nominate subspecies by the narrower and generally
restricted white markings; the antemedial fascia is mostly divided into two spots. On the hind
wing the black border is broader in the middle of the wing, its basal margin is less strongly
concave, and the white area is more restricted, especially in the $. The underside markings are
greatly reduced; the postmedial fascia on the fore wing is divided into two spots, whereas it is
generally a single large spot in V. hollandi hollandi; on the hind wings the apical white lines are
reduced. In the 3 genitalia the uncus tapers more evenly and the valve is narrower (see also
Remarks under V. hollandi hollandi).
BIoLoGy. Early stages and host-plant unknown. Moths collected in May and June.
DIsTRIBUTION (Fig. 17). Sulawesi: central, north-eastern and south-eastern regions.
MATERIAL EXAMINED
Holotype 3, Sulawesi: Menado (Van Braeckel) (genitalia slide no. 14635; BMNH).
Paratypes. Sulawesi: | 9 (allotype), data as holotype (BMNH); 2 9, Paloe, Sidaonta, 1350 m (‘4500 ft’),
vi.1937 (Kalis) (genitalia slide no. 15308; BMNH); 1 9, Paloe, Koelawi 1°25’ S, 119°58’ E, 950 m (‘3100
ft’), 111.1937 (Kalis) (genitalia slide no. 14686; BMNH); 1 9, Paloe, Loda, 1200 m (‘4000 ft’), v.1937 (Kalis)
(BMNH); 2 2, Minahassa, Tomohon, 1-9.vi & 26-29. vii.1954 (Alston) (genitalia slide no. 15307; BMNH);
2 2, Ulu Kolaka, 500 m, v—vi.1939 (Kalis) (BMNH).
Vitessa hollandi subsp.
The following two specimens probably represent a distinct subspecies, but without more material
we prefer not to name it.
3, 2. 46-50 mm. Fore wing above: orange subbasal area of similar size to that of V. hollandi wallacealis;
antemedial fascia indented but not interrupted, wider than in V. hollandi hollandi though not as wide as in
V. stettina; postmedial fascia oblique, divided into two spots, as in V. hollandi hollandi. Hind wing above
with tapering black terminal band. White markings on underside of both wings as in hollandi hollandi.
DISTRIBUTION. Moluccas.
MATERIAL EXAMINED
Moluccas: 1 3, Batjan, viii.1897 [abdomen missing] (Doherty) (BMNH); 1 9, Obi, Laiwui, ix.1897
(Doher ty) (BMNH).
Vitessa hollandi hollandi subsp. n.
(Figs 16, 89, 90, 153, 187, 245, 291)
[Cosmethis suradeva (Moore); Janse, 1928 : 88. Misidentification.]
3, 2. 40-50 mm. Generally as described for species. Thorax above: orange with large arrow-shaped medial
blue-black spot; patagium orange with large posteromedial glossy blue-black spot; tegula glossy blue-
black, laterally pale orange. Fore wing above: subbasal area orange; antemedial fascia white, broadly
indented at cell; postmedial fascia white, divided into two spots, upper large round spot over discocellular
vein, not reaching costa, lower triangular spot over CuP, smaller dot above, between Cu, and Cu,; white
streak along posterior margin joining streak from antemedial fascia; terminal area with white vein-lines,
sometimes absent on CuP, touching postmedial fascia at A,. Hind wing above: blue-black terminal band
wide at apex, gradually tapering, narrow near anal margin. Fore wing below: postmedial fascia with large
round spot and adjoining lower marking; white apical lines weak. Hind wing below: white apical lines
weak.
GENITALIA ¢ (Figs 153, 187). Generally as described for species. Valve broad; uncus with basal and distal
sections well demarcated.
GENITALIA 2 (Figs 245, 291). As described for species.
REMARKS. The subspecies is restricted to New Guinea and adjacent islands; it is rarer in Papua,
where V. stettina occurs sympatrically, than in north-eastern New Guinea. In New Guinea it
REVISION OF VITESSIDIA & VITESSA 291
occurs with V. hemiallactis hemiallactis, which it closely resembles externally (see Remarks under
V. hollandi and V. stettina, pp. 288-289; for distinguishing features from V. hollandi wallacealis see
Remarks under the subspecies, above). In Kepulauan Schouten V. hollandi hollandi bears a
similar relation to V. gemina (p. 312). We have not examined the specimen recorded by Janse as
V. suradeva from Mefor, Kepulauan Schouten. It may be referable to this subspecies or to V.
gemina. It is almost certainly not V. suradeva.
BioLoGy. Early stages and host-plant unknown. Moths collected in most months of the year.
DISTRIBUTION (Fig. 16). Kepulauan Schouten; New Guinea: Papua New Guinea.
MATERIAL EXAMINED
Holotype 3, New Guinea: Papua New Guinea, Finisterre Mts, Wantoat, 1200 m (‘4000 ft’), 13.ix.1957
(Munroe & Holland) (genitalia slide no. DK 113; CNC, Ottawa; type no. 11,021).
Paratypes. New Guinea: 3 9, Papua New Guinea, vic. Wewak, Koigin, 350 m (‘1200 ft’), 22—24.x.1957
(Munroe & Holland) (CNC, Ottawa); 3 9, P. N. G., Sepik District, Padwe R., 26.x.1957 (Munroe &
Holland) (CNC, Ottawa); 30 9, P. N. G., Sepik District, Bainyik, 350 m (‘1200 ft’), 29.x.-28.xi.1957
(Munroe & Holland) (genitalia slide no.’s DK 133, DK 134, DK 136; CNC, Ottawa); 1 9, P. N. G.,
Astrolabe Bay (Wahnes) (genitalia slide no. 15336; BMNH); 2 3, 32 2 (including allotype), P. N. G., data
as holotype, 5.ix.1957 (genitalia slide no.’s DK 114, 15336/a, Shaffer; CNC, Ottawa); 6 2, P. N. G.,
Morobe District, Araucaria Forest, Bulolo, 1200 m (*4000 ft’), 23.ix.1957 (Munroe & Holland) (genitalia
slide no.’s DK 112, DK 120; CNC, Ottawa); 3 9, P. N. G., Hunstein-Gebirge, Wagu/Black River,
i-ii.1973 (Hohmann) (UM, Bremen); 2 2, P. N. G., Mount Kebea, 1100 m (‘3600 ft’), iii.viii.1903 (A. E.
Pratt) (genitalia slide no.’s 15329, 15335; BMNH); 4 9, P. N. G., Sogeri Plateau, Mageri, 500 m (‘1600 ft’),
15.viii.1957 (Munroe & Holland) (genitalia slide no.’s DK 517, DK 520, DK 521; CNC, Ottawa); 1 3,
P. N. G., Sogeri Plateau, Bisianumu, 500 m (‘1600 ft’), 11.viii.1957 (Holland) (genitalia slide no. DK 175;
CNC, Ottawa); 1 9, P. N. G., Bisianumu, NE. of Port Moresby, 150m, vi.1957 (Hardy) (BPBM,
Honolulu).
Material excluded from paratype series. Kepulauan Schouten: 1 2°, Mefor (“Numfoor’), Suer, v—vi.1897
(Doherty) (BMNH). New Guinea: 1 9, P. N. G., Mount Kebea, 1100 m., vii.1903 (A. E. Pratt) (fore wing,
thorax and head of V. hemiallactis hemiallactis; hind wing and abdomen, glued on, of V. hollandi hollandi
(BMNHB)).
Vitessa hollandi subsp.
The following specimen probably represents a distinct subspecies, but without more material
we prefer not to name it.
2. 50 mm. Fore wing above: orange subbasal area of similar size to that of V. hollandi hollandi; narrow
antemedial and postmedial fasciae divided into two spots as in V. hollandi wallacealis, but lower spot of
postmedial fascia more oblique; terminal area with white vein-lines much shorter than in any other
subspecies, from M; to Cu,. Hind wing above and white markings on underside of both wings as in V.
hollandi hollandi.
DISTRIBUTION. Bismarck Archipelago: New Britain.
MATERIAL EXAMINED
Bismarck Archipelago: 1 9°, New Britain, Yalom, 1000 m, v.1962, at mercury vapour light (ZM,
Copenhagen).
The hemiallactis-group
DIAGNosIs. Wing markings either like those of stettina-group, or with the pale fasciae on the upperside
of fore wings greenish, expanded and partly or wholly merged, leaving a wedge-shaped black spot in cell.
Hind wing with white medial area and contrasting black terminal band. $ genitalia with uncus basally
wide, distally rod-like, with a row of laterally directed setae on each side. Valve less regular in outline than
in stettina-group, posterior margin concave at centre; clasper larger and more prominent than in stettina-
group, forming a finely serrulate ridge, differing from the coarsely serrate ridge of the pyraliata-group and
the bifid process of the p/umosa-group; costal margin of valve weakly curved subbasally and valve oblique
in dorsoventral preparation, as compared with the strongly curved costa and horizontal valve of the
suradeva-group. Large dorsal catena present on anellus. Cornuti consisting of three clusters, two basal
dorsolateral and one distal ventrolateral. 2 genitalia with much heavier ostial sclerotization than in
292 E. G. MUNROE & M. SHAFFER
stettina-group, similar to that of pyraliata-group. Posterior sclerite with lateral prolongations anteriorly
directed; anterior sclerite recurved and wrinkled, differentiated at centre, forming an anteriorly directed
pouch; antrum greatly extended posteriorly to form a trapezoidal plate fitting between the prolonged
sides of the posterior sclerite. Corpus bursae with signa consisting of paired denticulate areas. 2 seventh
abdominal segment with posterior margin convex, not concave or straight as in stettina-group.
Vitessa ternatica Lederer, 1863
(Figs 17, 91, 92, 154, 188, 212, 246, 292)
Vitessa ternatica Lederer, 1863, Wien. ent. Monatschr. 7 : 335, 456, pl. 6, fig. 7. Holotype 9, MOLUCCAs:
[Ternate] ex Felder coll. (genitalia slide no. 15306; BMNH) [examined].
Vitessa ternatica Lederer; Hampson, 1896a : 503.
Vitessa sarumensis Holland, 1900, Novit. zool. 7 : 582. LECTOTYPE 92, Mo.uccas: Buru (Doherty)
(genitalia slide no. 155 DK; CM, Pittsburgh), here designated [examined]. Syn. n.
3, 2. 42-44 mm. Frons orange. Vertex orange admixed with black. Eye, ocellus fuscous. Chaetosema with
pale yellowish setae. Antenna black, ventral surface of scape orange at base; dorsal scaling of basal half
to three-quarters of apical third of shaft white; terminal segments pale fuscous. Labial palpus: first
segment orange, second segment orange with black admixed distally, third segment black. Maxillary
palpus basally black, distally orange. Proboscis basally black. Thorax above: blue-black, with an orange
line anteriorly, terminally and laterally marked with greyish white; patagium glossy blue-black, anteriorly
and laterally admixed with orange; tegula glossy blue-black, anteromedially and laterally orange.
Abdomen above black, some pale grey laterally, terminal margins of segments narrowly pale grey, anal
tuft orange. Body below fuscous; pectus orange; abdomen black, terminal margins of segments pale grey,
anal tuft orange. Legs black, ventral surfaces of mid and hind femora white, mid tibia with white band at
centre, tips of mid and hind tibiae and spurs white. Fore wing long and narrow; above: glossy blue-black;
orange subbasal area restricted to small spot on CuP with isolated orange scales above and at base;
antemedial fascia white, single triangular spot on CuP, sometimes minute dot near costa; postmedial fascia
white, divided into two spots, large round spot over discocellular vein, more distal smaller spot on CuP;
terminal area with white vein-lines on most veins, as far as Cu, in 3; fringe basally dark grey, distally light
grey. Hind wing above mostly blue-black with violaceous reflections, white medial area small, comma-
shaped, directed towards base; fringe basally dark grey, distally white at apex, continuing as light grey to
anal margin. Fore wing below: blue-black with violaceous reflections; retinaculum rosy pale grey; post-
medial fascia forming one large oblique oval spot in ¢; similar to that of upperside in 2; apex with white
vein-lines from R, to Cu,; posterior margin white; fringe as above. Hind wing below similar to upperside,
but apex with white lines from costal margin to Cu.
GENITALIA ¢ (Figs 154, 188, 212). Uncus fairly long, slender, basal third wider and flattened, distal two-
thirds rod-like, laterally setose, recurved, then up-turned, terminally blunt. Gnathos with lateral arms
wide, posteriorly curved, central portion compressed, sickle-shaped and acuminate. Transtilla broadly
V-shaped, medially widened. Tegumen with posterior sclerite semicircularly expanded, anterior sclerite
with anterior margin strongly angled at centre with keyhole-shaped emargination. Vinculum broad and
basally convex. Valve long and broad; costa inflated, termen shallowly rounded, meeting costa at right
angles, posterior margin broadly concave at centre; striations on mesal surface fine; clasper ridge pro-
minent, oblique, joining carina ridge; at mid-length rising to isolated acuminate peak, falling gradually
with two or more minor points, parallel to mesal surface for short distance, rising and rounded with serrate
edge to posterior margin. Juxta with large narrow lateral sclerites; ventral process wide at base, strongly
up-turned, abruptly narrowing; dorsal process broadly rectangular, bearing setae at tip, joined to heavily
sclerotized and almost tubular lateral sclerite on anellus (Fig 188); sclerite connected to dorsal catena
via heavily sclerotized ridged bars; catena with compressed spines posteriorly, longitudinal ridges
anteriorly; manica with light sclerotization for most of length. Aedeagus long, distally tapering, base
narrowly rounded; ductus ejaculatorius nearly subbasal. Vesica with spines spreading ventrally to distal
end of aedeagus, membrane in this region bulbous and transversely rugose; vesica wide to just beyond area
of cornuti, membrane anterior to cornuti longitudinally rugose and spinose; cornuti (Fig. 212) with two
basal dorsolateral clusters, left cluster broadly connected with gradually shortening spines to heavily
sclerotized cap at angle of reflexion, right with almost isolated cluster of spines connected to cap by very
thin bar; posterior ventral cluster isolated and opposite to cap.
GENITALIA & (Figs 246, 292). High, narrow ovipositor lobes, dorsally fused, posteriorly produced; surface
densely setose, longer setae on margin, spiny setae visible at x 100 magnification. Posterior apophysis
short, slender, curved, tip truncated. Anterior apophysis much larger, thicker, curved, tip truncated.
REVISION OF VITESSIDIA & VITFESSA 293
Posterior sclerite of ostial chamber (Fig. 292) with lateral prolongations anteriorly directed; membrane
above finely denticulate; posterior margin broadly concave, medial anterior margin slightly projected to
meet antrum; anterior sclerite well developed, ventrally with recurved broad band, slightly wrinkled,
membrane beyond finely denticulate, sclerite laterally widening and greatly extended anteriorly, combining
with antrum to form a high wide dorsal surface; antrum extended posteriorly into a trapezoidal plate
fitting between the lateral prolongations of posterior margin. Basal end of ductus bursae wide and stretched
at junction with broad antrum, remainder of ductus narrow, spinose. Corpus bursae oval, bearing small
paired denticulate areas.
REMARKS. The small size and greatly reduced antemedial fascia and orange subbasal area give
this species a distinctive appearance; the broadly dark anal margin of the hind wing and the
enclosed comma-shaped white area distinguish the species from V. hemiallactis moluccana. We
find no obvious differences in specimens from the Moluccan localities. The New Guinea 3 has
white markings on the fore and hind wings noticeably reduced; this locality is questionable for
the specimen bears no other data though it is labelled as coming from the de Joannis collection.
The genitalia of both sexes are distinct from those of other species in this group; the clasper on
the valve of the ¢ and the great length of the antrum in the 2 are the most striking features.
BioLoGy. Early stages and host-plant unknown.
DISTRIBUTION (Fig. 17). Moluccas: Ternate, Buru, Seram; New Guinea.
MATERIAL EXAMINED
Moluccas: 2 2, [Ternate] ex Felder coll. (BMNH); 1 3, 3 9, Buru (Doherty) (CM, Pittsburgh; CNC
Ottawa) (paralectotypes of Vitessa sarumensis Holland); 1 9, Seram, Wahaai (Moens) (RNH, Leiden);
1 9, Seram (R.) (RNH, Leiden); 1 2, Seram, JIlo (BMNH); 1 2, Seram (BMNH); 2 9, no data, ex Moore
coll. (BMNH); 1 2, no data (Piepers) (RNH, Leiden). New Guinea: | 3, no data, 1920-1932 coll. L. & J.
de Joannis (MNHN, Paris).
Vitessa sp.
(Figs 93, 155, 189)
The following specimen represents a distinct species, but because it bears very dubious data we
prefer not to name it.
3. 44 mm. Generally as described for ternatica. Frons and vertex bright dark orange. Antennae missing.
First two segments of labial palpus orange, third segment missing. Thorax blue-black, posteriorly
bordered with white; patagium anteriorly orange, posteriorly glossy blue-black; tegula glossy blue-black,
anteriorly flecked with orange. Fore wing above, subbasal area bright dark orange, extending from costa
to A,; white antemedial fascia extending from costa to A,, medially diffuse; white postmedial fascia
divided into two spots, anterior round spot over discocellular vein, posterior triangular spot over CuP
directly behind anterior spot; terminal area with white vein-lines; fringe missing. Hind wing with very
broad blue-black margin, white area much restricted by infuscation from base; fringe basally dark grey,
distally white.
GENITALIA ¢ (Figs 155, 189). Generally as described for V. ternatica. Clasper on valve a prominent oblique
ridge, at mid length rising to isolated acuminate peak, curving convexly to posterior margin with irregular
serrations. Cornuti consisting of three clusters, left dorsolateral cluster with long sclerotized base and small
cluster of spines distally; right dorsolateral cluster consisting of a few sclerotized spine bases; single
ventral cluster a tight pack of shorter spines.
REMARKS. This distinctive species seems mid way in some of its features between V. ternatica and
V. hemiallactis. It bears a strong resemblance to V. hemiallactis moluccana but is distinguished by
the orange subbasal area reaching the costa, and by characters of the genitalia. In accordance
with resemblances already noted between other species within Vitessa which occur in the same
island or island-group it is reasonable to assume that this species might originate from the
Moluccas or somewhere nearby. The data label gives Japan, but it has been confirmed by Dr H.
Inoue (in /itt.) that no Pyralinae of this genus or any moths resembling this species are known
from the Japanese Islands. The genus occurs as far north as north India, Burma and north
Vietnam, but in the Pacific is not known to extend north of Luzon in the Philippines. It is possible
that the label is a misspelling of Japen (Kepulauan Schouten), but the Rev. J. C. E. Riotte
294 E. G. MUNROE & M. SHAFFER
informs us there is no direct evidence at the BPBM of the true provenance of this specimen or
others similarly labelled.
MATERIAL EXAMINED
Locality unknown: | ¢, Japan (BPBM, Honolulu).
Vitessa hemiallactis Meyrick, 1887
(Figs 17-19, 29, 37, 94-101, 156-159, 190-193, 213, 247-250, 293-296)
Vitessa hemiallactis Meyrick, 1887 : 193.
3, 2. 40-56 mm. Frons and vertex orange, sometimes admixed with black scales. Eye and minute ocellus
brown. Chaetosema with pale yellowish setae. Antenna black, anterior surface of scape basally orange;
dorsal scaling of apical third of shaft white to a variable extent, last several segments black. Labial palpus:
first segment orange; second segment orange, distally black; third segment black. Maxillary palpus
basally black, distally orange. Basal scaling of proboscis black, sometimes laterally admixed with orange.
Neck region orange. Thorax above with variable amounts of blue-black, anteriorly with some orange,
posteriorly yellowish white; patagium either orange with large posteromedial glossy blue-black spot, or
mostly glossy blue-black with anterior margin orange; tegula orange with anterolateral glossy blue-black
spot and distally admixed with blue-black, or mostly blue-black with some anteromedial and outer orange
scaling, or entirely blue-black. Abdomen above: matt black, posterior margins of segments with variable
amounts of whitish grey, terminal segment posteriorly bordered with orange, anal tuft orange. Body
ventrally and laterally blue-black; pectus orange; whitish grey abdominal lines broader than above;
posterior half of terminal segment and anal tuft orange. Legs blue-black with variable amounts of white
banding, a few orange scales at base of fore coxa. Fore wing rather narrow in 3, broader in 9; above:
ground colour blue-black; orange subbasal area variable in size, not reaching costa or posterior margin;
two whitish grey transverse fasciae of variable width, sometimes irregularly extending medially but never
joining; antemedial fascia sometimes as wide as postmedial fascia, indented at cell or divided, variable
amounts of medial diffusion; postmedial fascia broad or narrow, medially and terminally indented behind
cell or divided; terminal area with variably developed white vein-lines, not reaching postmedial fascia;
fringe dark grey. Hind wing above: blue-black marginal band with violaceous reflections, of variable
width; black anal margin variable in width; large white medial area broadly oval or rounded; fringe dark
grey basally, distally with variable amounts of whitish grey towards apex. Fore wing below: black with
violaceous reflections; posterior margin variably suffused with light brown; postmedial fascia larger than
above, complete or divided at Cu,, sometimes medial streaking between Cu, and CuP; terminal area with
apical white vein-lines; retinaculum dark grey. Hind wing below as above, with or without white apical
vein-lines, these reaching termen when present.
GENITALIA ¢ (Figs 29, 37, 156-159, 190-193, 213). Uncus slender, basally flattened and laterally con-
stricted, abruptly narrowing, terminal portion narrow, rod-shaped, decurved then up-turned with sparse
laterally directed setae, tip blunt. Gnathos with widely arched lateral arms and compressed sickle-shaped
medial process, tip finely acuminate. Transtilla shallowly V-shaped with strong S-shaped lateral flexure.
Tegumen: posterior sclerite broad, rounded; anterior sclerite with wide medial emargination. Vinculum
broad, basally concave. Valve narrow at base, widely inflated distally; costa sinuate, basally inflated;
termen broadly rounded; posterior margin deeply excised medially; mesal surface with fine longitudinal
converging striations; clasper a variably serrulate, continuous, diagonal ridge, with small curved carina
at base. Juxta with large narrow lateral sclerites, ventral process flat, narrow and up-turned; dorsal
process (Fig. 29) broader, bearing setae, directly connected to sclerotized flat sclerite supporting side of
anellus (Figs 190-193), sclerotization finely reticulated dorsally, longitudinally ridged; between sclerites a
large, narrow, heavily sclerotized catena (Fig. 37), constricted anteriorly, deeply ridged, posteriorly with
multiple spines; manica supported by lightly sclerotized bands. Aedeagus long, distally and basally
expanded, base narrowly rounded; ductus ejaculatorius subbasal; vesica with spines spreading ventrally
to distal end of aedeagus, here membrane bulbous and transversely rugose; vesica evenly tapering distally,
membrane between cornuti clusters longitudinally rugose; cornuti (Fig. 213) in three clusters, two basal
dorsolateral clusters bearing large spines, left cluster longer than right, and a single distal ventrolateral
cluster bearing smaller spines.
GENITALIA 2 (Figs 247-250, 293-296). High, narrow ovipositor lobes, dorsally fusing, posteriorly pro-
duced; surface densely setose, larger setae on margin, spiny setae easily visible at « 100 magnification.
Posterior apophysis short, slender, curved or straight; tip blunt, not expanded; anterior apophysis
longer, thicker, tip slightly spatulate. Posterior ostial sclerite (Figs 293-296) with lateral prolongations
REVISION OF VITESSIDIA & VITESSA 295
anteriorly directed, posterior margin straight or broadly convex, anterior margin medially concave;
anterior sclerite ventrally with recurved broad band, slightly wrinkled, finely denticulate, bearing broad,
curved medial pouch and much smaller lateral pouches; broad trapezoidal wall of antrum extending
posteriorly between lateral prolongations of posterior sclerite, antrum extending anteriorly, ending in a
broad curved margin; sclerotization at base of ductus seminalis; ductus bursae slightly expanded
posteriorly, remainder narrow, minutely spinose; corpus bursae globular, bearing small paired denticulate
areas.
REMARKS. This species is related to V. ternatica, but the 3 genitalia differ considerably in the shape
of the valve and clasper; in the 2 genitalia the antrum, though sharing the posteriorly developed
trapezoidal wall, is not nearly as long anteriorly. Externally V. hemiallactis is more lustrous,
with the white area on the hind wing relatively larger and with at most a narrow black band along
the anal margin. The various subspecies are easily confused externally with those of the stettina-
group (see Remarks under V. hollandi), and the subspecies admiralitatis and lustrans strongly
resemble the various species of the pyraliata-group, though these are not known to occur in the
same geographical areas; nevertheless, they are easily distinguishable in the genitalia, as pointed
out in the diagnosis of the species-group. In genitalia hemiallactis is closely similar to other
species in its own group, but it is distinct externally. We recognize four subspecies, from the
following geographical areas: (1) Moluccas; (2) Kepulauan Kai, Waigeo, New Guinea; (3)
Admiralty Islands; (4) Bismarck Archipelago, Solomon Islands.
BIioLoGy. Early stages and host-plants unknown. Moths collected in all months of the year.
DISTRIBUTION (Figs 17-19). Moluccas; Kepulauan Kai; Waigeo; New Guinea; Bismarck
Archipelago and Solomon Islands.
Vitessa hemiallactis moluccana subsp. n.
(Figs 17, 94, 95, 156, 190, 247, 293)
[Vitessa suradeva Moore; Pagenstecher, 1886 : 167, 188, partim. Misidentification. ]
3, 2. 42-50 mm. Generally as described for species. Frons in ¢ orange at base with extensive blue-black
above, completely orange in 9. Vertex orange with extensive black scaling admixed. Basal scaling of
proboscis blue-black, orange admixed laterally. Thorax posteriorly tipped with white; patagium glossy
blue-black, medially edged with orange: tegula glossy blue-black, laterally edged with orange. Fore wing
above, orange subbasal area from near costa to A,; narrow antemedial fascia divided at cell, anterior
spot minute, triangular, sub-costal, posterior spot large, oblong, medially diffuse; postmedial fascia
divided at Cu., anterior spot large, round, over discocellular vein from stem of R to Cuz, posterior spot
small, triangular and over CuP, terminal margin level with anterior spot; terminal vein-lines extensive, not
reaching postmedial fascia. Hind wing above: black terminal band broad, slightly diffuse medially; white
medial area narrow, reaching anal margin. Fore wing below: postmedial fascia large, oval or narrowly
divided at Cu2, apical area with variable number of veins-lines. Hind wing below: apical vein-lines variable.
GENITALIA ¢ (Figs 156, 190). Generally as described for species. Base of valve wide. See Remarks.
GENITALIA @ (Figs 247, 293). Generally as described for species. Posterior edge of trapezoidal wall of
antrum very broad, anterior development very narrow (Fig. 293).
REMARKS. Although only eight specimens are available they fall outside the range of variation of
an extensive series of the nominate subspecies. We conjecture that Pagenstecher’s record from
Ambon is referable to this species and subspecies. The subspecies is distinguished from the
nominate subspecies by the reduced pale markings on the wings. In the § genitalia the valve base
is slightly wider than in the nominate subspecies and the valve is slightly longer. In the 2 genitalia
the antrum has the trapezoidal area broader and the anterior development narrower than in any
other subspecies.
BioLoGy. Early stages and host-plant unknown. Holotype collected in September.
DISTRIBUTION (Fig. 17). Moluccas.
296 E. G. MUNROE & M. SHAFFER
MATERIAL EXAMINED
Holotype 3, Moluccas: Obi, Laiwui, ix.1897 (Doherty) (genitalia slide no. 14627; BMNH).
Paratypes. Moluccas: 1 2 (allotype), Halmaheira (Waterstradt) (genitalia slide no. 15818; BMNH);
1 3, Halmaheira, Toeelo (Le Moult) ({abdomen missing] BMNH); 1 3, Seram, 1895 (RNH, Leiden);
1 3, 1 2, Seram (Piepers) (RNH, Leiden); 1 3, 1 2, no data (Piepers) (genitalia slide no. 15818/a, Shaffer;
RNH, Leiden).
Vitessa hemiallactis subsp.
The following specimen probably represents a distinct subspecies, but without adequate material
or data we prefer not to name it.
2. 56 mm. Specimen in poor condition; in most characters similar to V. hemiallactis moluccana. Fore wing
above: lower spot of antemedial fascia broadly triangular, larger than in subspecies moluccana; postmedial
fascia also similar but lower spot more terminally oblique. Hind wing with narrow black margin and
extremely wide white medial area, exceeding any other species and subspecies in this group. Genitalia
much as in V. hemiallactis moluccana; posterior apophyses proportionally longer.
DISTRIBUTION. Unknown.
MATERIAL EXAMINED
1 2, no data, Moore coll. (BMNH).
Vitessa hemiallactis hemiallactis Meyrick, 1887
(Figs 18, 29, 96, 97, 157, 191, 213, 248, 294)
[Vitessa suradeva Moore; Pagenstecher, 1886 : 167. Misidentification.]
[Vitessa pyraliata Walker; Meyrick, 1886 : 213. Misidentification.]
Vitessa hemiallactis Meyrick, 1887, Trans. ent. Soc. Lond. 1887 : 193; Ragonot, 1891 : 110; Hampson,
1896b : 503; Swinhoe, 1900 : 429. LECTOTYPE 3, New GuINEA: Papua, 87:50 Port Moresby
(Mathews) (genitalia slide no. 8257; BMNH), here designated [examined].
[Vitessa suradeva Moore ; Kenrick, 1907 : 75. Misidentification.]
3, 2. 42-56 mm. Generally as described for species. Frons orange at base, with extensive blue-black above
in 3, completely orange in 9. Vertex orange with extensive blue-black admixed. Basal scaling of proboscis
blue-black. Thorax above posteriorly tipped with pale yellow; patagium orange with glossy blue-black
posteromedially, blue-black more extensive in ¢ than in 9; tegula mostly glossy blue-black, laterally
edged with orange, with more orange medially in 9 than in 3. Fore wing above: orange subbasal area
from near costa to A,; broad antemedial fascia widely indented at cell, sometimes finely divided in 2,
medial streaking approaching postmedial fascia; broad postmedial fascia strongly indented at Cus,
sometimes divided, anterior part broad and rounded, posterior part triangular, more oblique to termen,
streaks extending medially but not reaching antemedial fascia; terminal area with vein-lines more
extensive in ° than in g, not reaching postmedial fascia. Hind wing above: black terminal band narrow,
sharply demarcated from white medial area; medial area broad, fine black line at anal margin. Fore wing
below; postmedial fascia a large oval; apical area with slightly variable number of vein-lines. Hind wing
below: apical vein-lines slightly variable.
GENITALIA ¢ (Figs 29, 157, 191, 213). Generally as described for species. Clasper variable in outline.
GENITALIA 2 (Figs 248, 294). Generally as described for species. Trapezoidal wall of antrum noticeably
broadening anteriorly; anterior development of antrum exceeding medial pouch (Fig. 294).
REMARKS. The white markings on the upperside of the wings are more extensive than in other
subspecies. The subspecies differs from more easterly subspecies in the less intense metallic gloss
of the upperside markings. In markings it is almost indistinguishable from V. hollandi hollandi
in the stettina-group, with which it flies (see also Remarks under V. stettina and V. hollandi), but
the genitalia differ as indicated in the species-group diagnosis, and in the 2 the posterior margin
of the seventh abdominal sternite is convex, not straight or concave as in V. hollandi.
BIoLoGy. Early stages and host-plant unknown. Moths collected in most months of the year.
Specimens from Waigeo, Camp Nok, are labelled as being diurnal; in the same series one is
labelled as being caught at light.
REVISION OF VITESSIDIA & VITESSA 297
DISTRIBUTION (Fig. 18). Kepulauan Kai; Waigeo; New Guinea.
MATERIAL EXAMINED
Paralectotype. 1 2, data as lectotype (genitalia slide no. 15326; BMNH).
Kepulauan Kai: | 3, 1 2, Little Kei (“Kai Ketjil’), iv. (Kiihn) (BMNH). Waigeo: 1 3, 1 2 (Waterstradt)
(BMNH); | 3 (Wallace) (UM, Oxford); 3 3, 3 2 [2 d, labelled diurnal, 1 °, labelled at light], Camp Nok,
750 m, v.1938 (Cheesman) (BMNH). New Guinea: 7 9°, Irian Jaya, Central Arfak Mts, Ninay Valley,
1050-1100 m, xi.1908-i-iii.1909 (A. E. Pratt) (BMNH; CU, Ithaca); 1 2, I. J., Arfak Mts, 1200 m,
ii-iii.1909 (C. B. Pratt) (CMAG, Birmingham); | 9, I. J., Arfak Mts, 1850 m, iii.1910(C. B. & F. B. Pratt)
(BMNH); | 92, I. J., Dorey, vi.1897 (Doherty) (BMNH); 1 9, I. J., Kapaur, xii.1896-i.1897 (Doherty)
(BMNH); 3 8, I. J., Geelwink [sic] Bay, near Wandesi, 1892 (Doherty) (BMNH); 2 2, I. J., Nomnagihe,
40:23 km south of Wangaar, 600 m, i-ii.1921 (C. F. & J. Pratt) (TM, Pretoria); 1 9, I. J., Mt Goliath,
about 139° Long., 1500 m, 1ii.1911 (Meek) (BMNH); | 9,1. J., River Tor (mouth), 4kmE. of Hol.[landia],
Maffen, vii.1959 (Maa) (BPBM, Honolulu); 2 3, I. J., Maffin Bay, x.1944 (Ross) (CAS, San Francisco;
CNC, Ottawa); 2 9, I. J., Waris, S. of Hollandia, 450-500 m, viii.1959 (Maa) (BPBM, Honolulu); 6 2,
I. J., Humboldt Bay, ix—x.1892 (Doherty) (BMNH); 1 9, I. J., 1939, ex Guedet collection (CAS, San
Francisco); 1 9, Papua New Guinea, Telefomin (Eliptamin), 1350-1700 m, vi-ix.1959 (Brandt) (ANIC,
Canberra); 1 3, 6 9, P. N. G., Hunstein-Gebirge, Wagu/Black River, ix.1972-iii.1973 (Hohmann) (UM,
Bremen); 11 9, P. N. G., Sepik R., Padwe, x.1957 (Munroe & Holland)(CNC, Ottawa); 7 3, 65 9, P. N. G.,
Sepik District, Bainyik, 300 m, x—xi.1957 (Munroe & Holland) (CNC, Ottawa); 1 3, 1 2, P. N. G., Koigin,
nr Wewak, 350 m, x.1957 (Munroe & Holland) (CNC, Ottawa); 3 9, P. N. G., Astrolabe Range, winter,
1917 (F. P. & W. D. Dodd) (BMNH); 3 3, 6 2, P. N. G., Astrolabe Bay (Wahnes) (BMNH); 13 6, 33 9,
Finisterre Mts, Wantoat, 1200 m, ix.1957 (Munroe & Holland) (CNC, Ottawa); 2 2, P. N. G., Western
Highlands, Mt Hagen Valley, Keltiga, 1700 m, ix—x.1961 (Brandt) (ANIC, Canberra); 1 9°, P. N. G.,
Kainantu, 1650 m, x.1959 (Maa) (BPBM, Honolulu); 1 9, P. N. G., west of Lae, Bubia, sea level, viii. 1957
(Munroe & Holland) (CNC, Ottawa); 1 9, P. N. G., Lae, Botanic Garden, 30 m, viii.1957 (Munroe &
Holland) (CNC, Ottawa); 1 2, P. N. G., Lae, vi.1951 (Brandt & Hallstrom) (ANIC, Canberra); 2 9,
P. N. G., Busu River, E. of Lae, 0-60 m, vili.1957 (Munroe & Holland) (CNC, Ottawa); 1 9, P. N. G.,
8-04 km north-east of Lae, 30 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 1 2, P. N. G., south side
of Markham Valley, Oomsis, 0-30 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 12 3, 7 9, P. N. G.,
Morobe District, Bulolo, Araucaria forest, 1200 m, ix.1957 (Munroe & Holland) (CNC, Ottawa); 1 9,
P. N. G., Fly River, Kiunga, vii-x.1957 (Brandt) (ANIC, Canberra); 1 9°, P. N. G., Omati, iii.1952
(Barnett) (BMNH); 10 3, 9 9, P. N. G., Mount Kebea, 1100 m, vii—viii.1903 (4A. E. Pratt) (BMNH;
CMAG, Birmingham); 1 9, P. N. G., Ekeikei, 450 m, iii-iv.1903 (A. E. Pratt) (BMNH); 1 9, P. N. G.,
Aroa River (Meek) (BMNH); 1 3, 1 2, P. N. G., Lower Aroa R., xi.1904-11i.1905 (Meek) (BMNH);
43,39, P. N. G., Port Moresby, Mt Lawes, 400 m, ili-v.1963 (Brandt) (ANIC, Canberra); 1 2, P. N. G.,
Central District, Brown R., Karema, sea level, viii.1957 (Munroe & Holland) (CNC, Ottawa); 1 2, P. N. G.,
Bisianumu, NE. of Port Moresby, 150 m, vi.1957 (Hardy) (BPBM, Honolulu); 10 3, 18 2, P. N. G., Sogeri
Plateau, Bisianumu, 500 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 59 3, 96 2, P. N. G., Sogeri
Plateau, Mageri, 500 m, viii.1957 (Munroe & Holland) (CNC, Ottawa); 6 9, P. N. G., Lejo, 15 km
WNW - Popondetta, v.1973 (Hohmann) (UM, Bremen); 1 3, 3 2, P. N. G., Hydrographer Mts, 750 m,
i-ili.1918 (Eichhorn Bros.) (BMNH); 1 2, P. N. G., Biagi, Mambare R., 1500 m, iii.1906 (Meek) (BMNH);
1 9, P. N. G., Amazon Bay area, Dogon, 700 m, ix—xii.1962 (Brandt) (ANIC, Canberra); 4 2, P. N. G.,
Milne Bay, xi.1898-ii.1899 (Meek) (BMNH); 1 9, no data, 600 m (Barnett) (BMNH); | 2, no data (Walsh)
(RNH, Leiden).
The following data are dubious. Bismarck Archipelago: 1 3, New Britain, Rabaul District (Verteuil)
(BMNH).
Vitessa hemiallactis admiralitatis subsp. n.
(Figs 19, 37, 98, 99, 158, 192, 249, 295)
3, 2. 40-50 mm. Generally as described for species. Frons and vertex orange. Basal scaling of proboscis
glossy blue-black. Thorax above: glossy blue-black, posteriorly white-tipped; patagium glossy blue-black,
lateral extremities orange; tegula glossy blue-black. Abdomen above: glossy black; grey terminal lines on
segments much reduced or absent; terminal segment with orange posterior margin. Fore wing above:
ground colour glossy blue-black, basal areas and posterior margin with greenish hue; orange subbasal
area small, in 2 short of costa, reaching to A,, in ¢ restricted to smaller dot posterior to base of cell;
white antemedial fascia very narrow, basally straight, medially diffuse and indented at cell, short of costa
and posterior margin; white postmedial fascia oblique, narrowly divided at Cu., round anterior spot
298 E. G. MUNROE & M. SHAFFER
slightly larger than triangular posterior spot; terminal area with vein-lines short and restricted, not
reaching termen or postmedial fascia, more extensive in 2. Hind wing above: small black basal area, well
demarcated terminal black band, anal margin narrowly black, angled from basal area. Wings below with
faint, much reduced apical vein-lines.
GENITALIA ¢ (Figs 37, 158, 192). As described for species.
GENITALIA 2 (Figs 249, 295). Generally as described for species. Posterior trapezoidal wall of antrum with
lateral margins posteromedially oblique, posterior margin concave; long anterior development of antrum
(Fig. 295).
REMARKS. This subspecies more closely resembles V. hemiallactis lustrans than the nominate
subspecies, but differs from V. h. /ustrans in the less extensive orange markings of the thorax,
wing base and subbasal area, the very narrow but generally undivided antemedial fascia of the
fore wing and the more extensive development of the terminal white vein-lines. It differs from the
nominate subspecies in having the frons and vertex wholly orange, in the less extensive pale
markings on the fore wing, and the thicker black terminal band on the hind wing. In the 3
genitalia the clasper is more irregular in outline than in previous subspecies; in the 9 genitalia
the shape of the antrum is different.
BIioLoGy. Early stages and host-plant unknown. Moths collected in April, June, September and
October.
DISTRIBUTION (Fig. 19). Admiralty Islands.
MATERIAL EXAMINED
Holotype ¢, Admiralty Islands: Manus, ix—x.1913 (Meek) (BMNH).
Paratypes. Admiralty Islands: 2 3, 4 9 (including allotype), data as holotype (genitalia slide no.’s 15140,
15819; BMNH); 1 3, 6 2 [no exact data] (Meek) (genitalia slide no.’s 14658, 14659, 15820; BMNH);
1 9, Manus I., Rossum, 35-125 m, vi.1959 (Gressitt) (BPBM, Honolulu); 1 3, Los Negros I., iv.1952,
(Brandt & Hallstrom) (ANIC, Canberra).
Vitessa hemiallactis lustrans subsp. n.
(Figs 19, 100, 101, 159, 193, 250, 296)
[Vitessa ternatica Lederer; Pagenstecher, 1900 : 169. Misidentification. ]
3, 2. 42-54 mm. Generally as described for species and previous subspecies. Frons and vertex dark orange
sometimes lightly admixed with black in g. Patagium medially glossy blue-black, laterally dark orange,
with medial streak of orange in 9; tegula glossy blue-black, small orange spot anteromedially, larger
orange spot on outer margin. Abdomen above glossy black, terminal segment with orange posterior
margin. Fore wing above: orange subbasal area large, slightly short of costa, anteriorly extending beyond
A,, of similar size in both sexes; white antemedial fascia narrow, broadly divided, anterior spot minute,
subcostal; posterior spot broadly triangular, extending from cell to beyond A,; white postmedial fascia
oblique, broadly divided at Cu,, round anterior spot larger than triangular posterior spot; terminal area
with vein-lines larger and more extensive in 9 than in 3g, longest on R; and M,, much restricted in 3, not
reaching termen or postmedial fascia. Hind wing above, black band on terminal margin very wide. Wings
below with faint, much reduced apical vein-lines, sometimes absent on hind wing in both sexes.
GENITALIA ¢ (Figs 159, 193). Generally as described for species. Clasper with free margin deeply concave
at centre, basally with high acuminate ridge.
GENITALIA 2 (Figs 250, 296). Generally as described for species. Posterior trapezoidal wall of antrum with
posterior margin slightly convex (Fig. 296).
REMARKS. This subspecies most closely resembles V. hemiallactis admiralitatis; it differs in the 3
genitalia by the shape of the clasper and in the 2 by the shape of the posterior wall of the antrum.
There is little variation within the Bismarck Archipelago. A specimen from the Rabaul District
of New Britain is unlike all others from this area and exactly matches the New Guinea subspecies
to which we have referred it; the specimen may be mislabelled or it may represent an accidentally
introduced New Guinea individual. We have not seen Pagenstecher’s material, collected by
Ribbe in New Britain (Neu Pommern), Kinigunang, but it almost certainly belongs to this species
REVISION OF VITESSIDIA & VITESSA 299
and subspecies. It is possible that the bulk of the Solomon Islands specimens represent a distinct
subspecies. The fore wing markings are slightly different from the more northerly material but are
also variable over the Solomon Islands range. In the 3 genitalia the valve is narrower and longer;
in an isolated specimen from Florida Island the right cornuti cluster is much reduced, though this
may be due to broken-off spines. A description and naming will await the discovery of more 3
specimens from a range of the islands.
BioLoGy. Early stages and host-plant unknown. Moths collected in most months of the year.
DISTRIBUTION (Fig. 19). Bismarck Archipelago; Solomon Islands.
MATERIAL EXAMINED
Holotype 3, Bismarck Archipelago: New Hannover, iii.1923 (Meek) (BMNH).
Paratypes. Bismarck Archipelago: | 9° (allotype), New Hannover, iv.1923 (Meek) (BMNH); 5 2, 2 9,
New Hannover, ii-iii.1923 (Meek) (genitalia slide no.’s 14660, 14661, 15822, 15825; BMNH).
Material excluded from paratype series. Bismarck Archipelago: 1 3, 4 2, Rooke I. (‘Umboi Island’),
vii—viii.1913 (Meek) (BMNH); 2 3, 12 2, New Ireland, xi.1923-i1.1924 (A. F. Eichhorn) (BMNH; CMAG,
Birmingham; TM, Pretoria); 4 2, New Britain (Wahnes) (BMNH); 3 2, New Britain, Talesea, i-iv.1925
(A. F. Eichhorn) (BMNH); 1 9, New Britain, Keravat, 135 m, xi.1959, at mercury vapour light (Maa)
(BPBM, Honolulu); 1 2, New Britain, Yalom, 1000 m, v.1962 (ZM, Copenhagen); 2 2, New Britain, Mt
Sinewit, 1050 m, vi-ix.1963 (Brandt) (ANIC, Canberra; CNC, Ottawa). [1 3, New Britain; see Remarks.]
Solomon Islands: 2 2, Bougainville, Buin, Kukugai Village, x.1960-i1.1961 (Brandt) (ANIC, Canberra;
CNC, Ottawa); 8 ¢, 9 2, Guizo (‘Gizo Island’), xi.1903 (Meek) (BMNH; CNC, Ottawa; NMNH,
Washington); | 3, Florida, 1.1901 (Meek) (BMNH); 1 2, Guadalcanal, Betikama R., viii-x.1960 (Brandt)
(ANIC, Canberra).
Vitessa intermedia sp. n.
(Figs 20, 102, 103, 160, 194, 251, 297)
3, 2. 50-60 mm. Frons and vertex orange, admixed with black in 3; scaling of vertex forming strong crest.
Eye brownish fuscous. Ocellus minute, brown. Chaetosema with pale yellowish setae. Antenna black,
anterior surface of scape basally orange; posterior surface with basal half of apical third of shaft white.
Labial palpus: first segment orange; second segment orange, distally black; third segment black. Maxillary
palpus basally black, distally orange. Basal scaling of proboscis black, sometimes laterally admixed with
orange. Neck region orange. Thorax above: anteriorly orange, medially blue-black, posteriorly white
admixed with yellow; patagium orange with posteromedial glossy blue-black spot; tegula orange with
anterolateral glossy blue-black spot and posteriorly admixed with black. Abdomen above matt black,
posterior margins of segments whitish grey, posterior half of terminal segment and anal tuft orange. Body
below blue-black with oblique white bands; pectus orange; whitish grey abdominal lines broader than
above, posterior half of terminal segment and anal tuft orange. Legs blue-black; greenish hue on fore
coxa; white on ventral surfaces of femora; medial white band on mid tibia; white at tips of tibiae and
ventral surfaces of tibial spurs. Fore wing above: orange subbasal area large with narrow subcostal streak
to base, basal blue-black spot partially demarcated, orange short of costa and posterior margin; glossy
blue-black subbasal fascia with greenish hue; whitish grey transverse fasciae with pale greenish hue,
antemedial and postmedial fasciae wide, medially diffuse, irregularly joining, usually both fasciae slightly
short of costa and posterior margin; medial blue-black fascia reduced to two irregularly shaped patches,
or, more rarely, a single longitudinally triangular spot on cell; terminal area wide, greenish white vein
lines thick on all veins, most lines joining postmedial fascia and termen, completely merging in costal
region; fringe dark grey. Hind wing above: broad black terminal band violaceous, gradually tapering
from costa, acuminate at anal margin, close to infuscation from wing base, anal margin edged with
black; white medial area large, oval; fringe dark grey, distally white near apex. Fore wing below: black,
weakly violaceous; retinaculum light grey; a single large, oval postmedial fascia, in ¢ extending to near
posterior margin, in 2 hardly passing Cu,; terminal area with apical white vein-lines, posteriorly shorter
and weaker; posterior margin white; fringe as above. Hind wing below: much as above, long white
vein-lines near costa and apex, small vein-lines on posterior part of termen; fringe as above.
GENITALIA ¢ (Figs 160, 194). As described for V. hemiallactis, but see Remarks.
GENITALIA 2 (Figs 251, 297). As described for V. hemiallactis, but see Remarks.
REMARKS. This species is externally almost identical with the sympatric V. stettina, which also
has very broad white fasciae on the fore wing, but differs by having the fasciae converging on the
300 E. G. MUNROE & M. SHAFFER
medial area and fusing; its genitalia are very distinct. V. intermedia flies over the same range of
altitudes as V. stettina, though it appears to be more common in the upper part of its range than
in the lower, where it is largely replaced by V. hemiallactis. The paratype series includes 3 and
2 paralectotypes of V. stettina, from which Swinhoe failed to distinguish the present species. V.
intermedia closely resembles the Australian V. g/aucoptera in appearance and general structure;
one 3 from Avola, Papua, has wing markings almost identical with those of V. glaucoptera, but
represents an extreme of variation. V. intermedia differs from V. barretti, which overlaps the
upper part of its altitude range, by the smaller size, smaller orange subbasal area, less completely
fused fasciae, thinner vein-lines, merging only slightly in the costal region; in the ¢ genitalia the
posteroventral angle of the valve is less extended; in the 2 genitalia the posterior development of
the trapezoidal wall of the antrum is longer but narrower (Fig. 297). V. intermedia closely
resembles V. hemiallactis hemiallactis but has larger wing markings; the 3 genitalia have a larger
valve and a more extended posteroventral angle; the 2 genitalia most closely resemble those of
V. hemiallactis lustrans. In external and genital characters this species is intermediate between V.
hemiallactis and V. barretti and, apart from the latter species, is the largest species in the
hemiallactis-group.
BioLoGy. Early stages and host-plant unknown. Moths collected in February, March, June and
August.
DISTRIBUTION (Fig. 20). New Guinea: Papua New Guinea.
MATERIAL EXAMINED
Holotype 3, New Guinea: Papua, Biagi, Mambare R[iver], 1500 m (‘5000 ft’), ii.1906 (Meek) (genitalia
slide no. 15311; BMNH).
Paratypes. New Guinea: | 9 (allotype), Papua New Guinea, data as holotype, iii.1906 (genitalia slide
no. 15347; BMNH); 1 3, P. N. G., Granville (“Port Moresby’) [paralectotype of Vitessa stettina Swinhoe]
(genitalia slide no. 8259; BMNH); 1 3, P. N. G., Granville, 6845 (genitalia slide no. 15309; BMNH);
1 3, P. N. G., Milne Bay, 20.viii.1903 (Meek) (genitalia slide no. 15340; BMNH);1 9, P. N. G., Mount
Kebea, 1100 m (‘3600 ft’), vi.1903 (A. E. Pratt) [paralectotype of Vitessa stettina Swinhoe] (genitalia
slide no. 15346; BMNH); 1 9, P. N. G., Foula, 1350 m (*4500 ft’), viii.1903 (A. E. Pratt) (genitalia slide
no. 15348; BMNH); 8 4, 1 2, P. N. G., Avola, 1850 m (‘6000 ft’), viii.1903 (A. E. Pratt) (genitalia slide
no.’s 14718, 15310, 15313, 15316, 15319, 15339, 15341 [1 3, 1 2, abdomens missing]; BMNH).
Vitessa barretti sp. n.
(Figs 20, 104, 105, 161, 252, 298)
3, 2. 58-63 mm. Frons and vertex unusually narrow, orange. Eye and ocellus fuscous. Chaetosema with
pale yellowish setae. Antenna black; dorsal scaling on basal half of apical third of shaft white. Labial
palpus mainly black with considerable orange admixture dorsolaterally and ventrally. Maxillary palpus
basally black, distally orange. Basal scaling of proboscis black. Neck region orange. Thorax above black,
anteriorly orange; patagium orange with large medial glossy blue-black spot; tegula orange with antero-
lateral glossy blue-black spot. Abdomen above black, posterior margins of segments narrowly white; anal
tuft orange. Body below black, abdomen as above; thorax and legs black with extremely restricted areas
of white. Fore wing rather wide; above: orange subbasal area large, reaching posterior margin; narrow
subcostal streak to base, partially demarcating basal blue-black spot; very narrow glossy blue-black
subbasal fascia; antemedial and postmedial greyish green fasciae completely fused, reaching costa and
posterior margin, medial fascia a small, longitudinal, triangular, blue-black streak in cell; thick greyish
green vein-lines in terminal area, all merging with postmedial fascia, especially at costa, blue-black streaks
between lines; fringe basally black, distally pale grey. Hind wing above: broad black terminal band
violaceous, inner margin angled from M, to Cuz, abruptly narrowing then tapering to anal margin; deep
infuscation from base; white medial area oval, reaching anal margin; fringe basally fuscous, distally white.
Fore wing below: black; single, small, irregularly ovate, white postmedial fascia in $ with diffuse white
distally in cubital area; subterminal whitish grey vein-lines; fringe basally blackish fuscous, distally paler.
Hind wing below: like upperside; white vein-lines apically and terminally.
GENITALIA ¢ (Fig. 161). Generally as described for V. hemiallactis. Large valve distally subtruncate;
termen narrowly rounded; base of uncus fairly wide.
GENITALIA 2 (Figs 252, 298). As described for V. hemiallactis.
REVISION OF VITESSIDIA & VITESSA 301
REMARKS. This species resembles V. griseata in the almost unicolorous medial area of the fore
wing, but is distinguished by the black medial streak on the fore wing and the white medial area
on the hind wing, as well as by genitalia (see diagnoses of the griseata- and hemiallactis-groups).
This species is distinguished from V. stettina by the loss of much of the black pattern of the fore
wing, and by the group characters of the genitalia; it shows similar pattern differences from V.
intermedia and also has the orange subbasal area larger; the 3 genitalia, though almost identical
with those of V. intermedia, have the posteroventral angle of the valve more extended; in the 2
genitalia the posterior development of the trapezoidal wall of the antrum is shorter but broader
(Fig. 298). This species, however, is most easily confused with V. glaucoptera from Australia,
which has almost identical wing markings and genitalia; it is distinguished by the much larger
size and by having the black streak between M, and M, of the fore wing more constricted at
junction with fasciae and the black streak in the cell longer; the genitalia are correspondingly
larger. Though the members of the hemiallactis-complex in New Guinea and Australia are closely
related to one another, it seems best at present to consider V. hemiallactis, intermedia, barretti
and glaucoptera different species. We take pleasure in dedicating the present species to Mr J.
Barrett, collector of the holotype.
BioLoGy. Early stages and host-plant unknown. Moths collected in February and September, at
altitudes from 600 to over 1800 m.
DISTRIBUTION (Fig. 20). New Guinea: Papua New Guinea, in the central mountains.
MATERIAL EXAMINED
Holotype 3, New Guinea: North-east New Guinea, Western Highlands, Mt Hagen, 22.ix.1951 (Barrett)
(genitalia slide no. DK 78; CNC, Ottawa).
Paratypes. New Guinea: | 9 (allotype), Papua New Guinea, Eastern Highlands, Aiyura, 27.ix.1957
(Munroe & Holland) (genitalia slide no. DK 79; CNC, Ottawa); P. N. G., 1 9, Gurukor, 600 m, 10-
12.11.1978 (Hébert) (CNC, Ottawa).
Vitessa glaucoptera Hampson, 1906
(Figs 20, 106, 107, 162, 195, 253, 299)
Vitessa glaucoptera Hampson, 1906, Ann. Mag. nat. Hist. (7) 17 : 216. Holotype 3, AUSTRALIA: Queens-
land, Cedar Bay, s[outh] of Cooktown (Meek) (genitalia slide no. 8258; BMNH) [examined].
3, 2. 34-52 mm. [Holotype 46 mm, incorrectly given by Hampson as 30 mm.] Frons orange; vertex orange
admixed with dark grey. Eye and ocellus brown. Chaetosema with pale yellowish setae. Antenna black;
base of scape orange; dorsal scaling of basal half of apical third of shaft white. Labial palpus: first two
segments orange, third segment black. Maxillary palpus basally black, distally orange. Basal scaling of
proboscis black, laterally orange, orange admixed dorsally. Neck region orange. Thorax above orange,
with large central blue-black spot; patagium orange with small posteromedial glossy blue-black spot;
tegula orange, distally admixed with black, small anterolateral glossy blue-black spot. Abdomen above:
matt black, posterior margins of segments narrowly white, terminal segment with posterior margin and
anal tuft orange. Body below blue-black; pectus orange; legs blue-black with dark green hue; white bands
on mid and hind coxae; white on ventral surfaces of mid and hind femora; medial white band on mid
tibia; tibiae white-tipped; tibial spurs dorsally white; abdomen with white margins broader than above.
Fore wing above: orange subbasal area large, with narrow subcostal streak to base; basal blue-black spot
partially demarcated; orange sometimes reaching posterior margin in 2; narrow glossy blue-black subbasal
fascia; greyish green antemedial and postmedial fasciae almost completely fused, reaching costa and
posterior margin, small broadly triangular blue-black spot in cell, sometimes with small black subcostal
streak above and diffuse black patch below in cubital area; thin greyish green vein-lines in terminal area,
on M, and M, slightly short of fasciae, sometimes also on CuP; fringe basally dark grey, distally light
grey. Hind wing above: broad black terminal band violaceous, broad at apex, basal margin angled from
M, to Cuz, abruptly narrowing and tapering to anal margin; fringe basally dark grey, distally white. Fore
wing below: black; single, small, irregularly ovate, white postmedial fascia, diffuse in cubital region;
scattered white scales along costa; posterior margin white; fine, small, white apical vein-lines; fringe as
above. Hind wing below as above; small, faint, white apical vein-lines.
GENITALIA ¢ (Figs 162, 195). Generally as described for V. hemiallactis. Posteroventral angle of valve
broadly rounded, valve narrow.
302 E. G. MUNROE & M. SHAFFER
GENITALIA 2 (Figs 253, 299). As described for V. hemiallactis.
REMARKS. In general as in Remarks for V. barretti. The greenish colour of the fore wing led
Swinhoe to associate V. stettina with the present species, but the two are not closely related.
The independent acquisition of the same colour by the Australian V. zemire nephritica suggests
adaptation to a common environmental factor, and can be compared to the parallel acquisition
of a similar greenish colour by V. barretti and the unrelated V. griseata in the highlands of New
Guinea. Though externally V. glaucoptera most closely resembles V. barretti, in size and fore wing
markings it is also comparable to V. intermedia (see Remarks under V. intermedia for Avola
specimen). The ¢ genitalia lack the extended posteroventral angle of the valve seen in V. intermedia
and barretti; the valve is most like that of V. hemiallactis hemiallactis but narrower; the 9
genitalia are also most like those of that subspecies.
BioLoGy. Early stages and host-plant unknown. Moths collected in March, September, November
and December.
DISTRIBUTION (Fig. 20). Australia: Queensland.
MATERIAL EXAMINED
Australia: 1 2, Queensland, Cedar Bay, south of Cooktown (Meek) (BMNH); 1 9, Q., Evelyn Scrub
[145°27’ E, 17°30’ S], xi.1910 (Dodd) (ANIC, Canberra); 3 3, 1 9, Q., Kuranda, iii, ix, xii.1904, iv.1905
(Dodd) (ANIC, Canberra); 1 3, 7 2, Q., Kuranda, n. Cairns 1910 (Dodd) (BMNH); 2 2, 13 2, Q., 17-6 km
south of Ravenshoe, 800 m, i1i.1964 (Common & Upton) (ANIC, Canberra; CNC, Ottawa); 2 3, 7 2, Q.,
Mt Lewis, 12:8 km north-west of Mt Molloy, 800 m, iii.1964 (Common & Upton) (ANIC, Canberra);
3 g, 2 9, Q., 4:8 km west of Mossman, iii.1964 (Common & Upton) (ANIC, Canberra); 1 2, Q., 27 km
(‘17 mls’) south of Atherton, 915 m, iii.1964 (Common & Upton) (ANIC, Canberra).
Vitessa cristobalensis sp. n.
(Figs 2, 108, 254, 300)
3. Unknown.
9. 38mm. Frons orange. Vertex orange with central admixture of black. Eye and ocellus black.
Chaetosema with pale yellowish setae. Antennae black, base of scape orange; dorsal scaling of basal half
of apical third of shaft white. Labial palpus: first two segments orange, third segment black. Maxillary
palpus basally black, distally orange. Basal scaling of proboscis glossy blue-black. Neck region orange.
Thorax above: glossy blue-black, laterally edged and posteriorly tipped with orange; patagium glossy
blue-black, laterally tipped with orange; tegula glossy blue-black with anteromedial and outer lateral
edges finely orange. Abdomen above black (markings not recorded). Body below blue-black; pectus
orange; legs black, mid and hind coxae banded with white, posterior surfaces of mid and hind femora
white, mid tibia with medial white band, mid and hind tibiae tipped with white. Fore wing narrow; above:
ground colour glossy blue-black with dark green hue; orange subbasal area small, restricted between stem
of Rand CuP; faintly greenish white antemedial fascia narrow, deeply indented medially at cell by wedge
of ground colour, anterior portion triangular, posterior part oblong and larger, narrowly separated from
costa and posterior margin; greenish white postmedial fascia divided into two spots, large oval anterior
spot over discocellular vein, extending from stem of R to Cuz, posterior spot far distad of anterior,
narrowly triangular, spanning CuP; terminal area with narrow greenish white vein-lines extending from
termen, well short of oval spot of postmedial fascia, but nearly touching posterior triangular spot; fringe
glossy, basally dark grey, distally pale grey. Hind wing above: broad blue-black terminal band violaceous,
very broad from infuscated costa, commencing from upper end of cell, sharply angled to just beyond Cuz,
continuing narrowly and parallel to margin, finely upturned and ending at anal margin; anal margin finely
edged with black; white medial area roughly rectangular; fringe basally dark grey, distally white, anal
angle black. Fore wing below: blue-black, weakly violaceous, shading to brownish grey and admixed with
white at posterior margin; retinaculum light grey; postmedial fascia diffuse, oblique, narrowly oval,
white; white vein-lines in terminal area weak and shorter than above; fringe dark grey. Hind wing below
as above, fringe mostly white; weak, short apical and terminal vein-lines extending to Cuz, joining fringe.
GENITALIA 2 (Figs 254, 300). High, dorsally fused, posteriorly narrow and produced ovipositor lobes;
surface densely setose, larger setae on margin, spiny setae easily visible at < 100 magnification. Posterior
apophysis short, slightly curved. Anterior apophysis longer, thicker, inwardly curved at tip. Posterior
ostial sclerite (Fig. 300) very wide with long lateral prolongations anteriorly directed, medial portion
REVISION OF VITESSIDIA & VITESSA 303
narrow with anterior margin recurved, thickened, forming broad short flap; posterior margin broadly
concave; anterior sclerite very wide, short, with recurved and slightly wrinkled ventral margin, minutely
denticulate, bearing small weakly developed medial pouch; very broad trapezoidal wall of antrum
extending posteriorly between lateral prolongations of posterior sclerite, laterally acutely angled, posterior
edge half width of anterior sclerite, antrum anteriorly produced for shorter distance and narrowing; base
of ductus seminalis mostly free of sclerotization; ductus bursae slightly expanded posteriorly, remainder of
ductus narrow, minutely spinose; corpus bursae comma-shaped, bearing small, paired, denticulate areas.
REMARKS. This is distinguished from all other species and subspecies with narrow, wholly or
partly divided white fascia by having the antemedial fascia almost and the postmedial fascia
clearly divided into two spots, in combination with the following characters: terminal area with
all veins bearing long white vein-lines, nearly touching posterior spot of postmedial fascia; much
reduced orange subbasal area. This species most closely resembles V. hemiallactis lustrans, but is
distinguished by longer vein-lines of the fore wing, the greater width of the antrum and the shape
of the corpus bursae.
BioLoGy. Early stages and host-plant unknown. Holotype collected at black light.
DISTRIBUTION (Fig. 2). Solomon Islands: San Cristobal Island.
MATERIAL EXAMINED
Holotype 2, Solomon Islands: San Cristobal, Warahito—Pogato confluence, 1965, at black light (genitalia
slide no. 14263; BMNH).
The pyraliata-group
DIAGNOsIS. Pectinations of ¢ antennae larger than in other species-groups. Wing markings similar to those
of stettina- and hemiallactis-groups, but with pale markings of fore wing relatively restricted in most
species. Hind wing markings very varied, in V. nicobarica 3 all dark markings absent, in 2 white area
heavily infuscated; in V. pyraliata similar infuscation, but in some subspecies with a very wide terminal
band, well demarcated from the white medial area, as also in V. kolakalis and V. sulaensis; in V. muluana,
V. gemina and V. splendida the terminal band narrow, as in plumosa- and suradeva-groups. 3 genitalia
with basal portion of uncus wide, distal portion rod-like, down-turned then up-curved, with laterally
directed setae. Valve of moderate width, shape somewhat similar to that of hemiallactis-group, but
narrower, clasper coarsely serrated basally and with tooth-like ridges on posterior margin, not finely
serrulate as in hemiallactis-group. 2 genitalia with very large ostial chamber; posterior sclerite wide, medial
section narrowed but with produced medial or lateral portions; lateral section of sclerite anteriorly
elongated, in V. muluana with an oblique raised ridge. Anterior sclerite broad, ventrally narrow, antrum
very well-developed anteriorly, posterior development similar to that of hemiallactis-group. Ostial
sclerotization relatively larger than in stettina- or hemiallactis-groups, smaller and less regular than in
suradeva-group. Corpus bursae with either fine paired denticulate areas or heavily sclerotized, depressed,
paired denticulate signa.
REMARKS. Four of the species assigned to this group are known only from the 2. When the 3
characters are known some of these species may require reassignment or the definition of the
species-group may have to be revised.
Vitessa nicobarica Hampson, 1896
(Figs 21, 44, 109, 110, 163, 214, 255, 301)
Vitessa nicobarica Hampson, 1896a, The Fauna of British India, including Ceylon and Burma. Moths
4: 146. LECTOTYPE 3, Nicopar IsLANDs: Nancowry, Moore coll. 94-106 (genitalia slide no. 8268;
BMNH), here designated [examined].
Vitessa nicobarica Hampson; Hampson, 18966 : 503.
3, 2. 44 mm (all specimens damaged). Frons, vertex (from remaining scales) orange. Eye, ocellus black.
Chaetosema with pale yellowish setae. Antenna black, base of scape orange (at least in 9; terminal
segments lost). Labial palpus blue-black; base of first segment orange; tip of third segment fuscous.
Maxillary palpus basally black, distally orange. Basal scaling of proboscis black. Neck region orange.
Thorax above: orange with large central blue-black spot; patagium blue-black laterally tipped with orange;
tegula orange, with broad, oblique, blue-black band. Abdomen above: ochreous buff, anal tuft orange.
304 E. G. MUNROE & M. SHAFFER
Body below: pectus orange; thorax blue-black; legs black, possibly with some white on coxae (legs mostly
damaged or missing); abdomen blackish fuscous, posterior margins of segments whitish buff, pale bands
progressively wider on the posterior segments, anal tuft orange. Fore wing narrow, costa convex; above
(very worn): ground colour blue-black ; orange subbasal area more restricted in 3, short of costa, reaching
to A,, in 9 extending past A,, subcostal streak to base partially demarcating blue-black basal spot; narrow
white antemedial fascia divided into two spots at cell, smaller triangular anterior spot, larger, rectangular,
medially diffuse posterior spot, short of posterior margin, fascia more restricted in ¢; broader white
postmedial fascia, divided into two primary spots, anterior spot over discocellular vein, entire, large, oval,
short of costa, posterior spot more distal, triangular, divided into four smaller short streaks by black lines
on veins Cu,, Cu, and A,, reaching posterior margin in 2, in some specimens a trace of an additional black
line on CuP; posterior white marking absent in 3; terminal area with short, thin apical and terminal white
vein-lines, short of termen and postmedial fascia, extending to Cu, in 9, very much weaker in 3; fringe
dark grey. Hind wing above: in 3 uniformly pale yellow with isolated black scales at apex, in 2 costa black
narrowly edged with white, termen black, narrow, diffuse, more strongly black for short distances on M3,
Cu, and Cuz, medial area broadly white to anal margin; fringe white, basally admixed with black towards
apex. Fore wing below: in 3, costa narrowly black, apex and termen triangularly black to tornus, slight
black admixture from base weakly spreading over cell; remainder of wing and retinaculum pale yellow;
fringe black slightly admixed with yellow at tornus; in 2 white postmedial fascia visible as a broad L-shape
broken by black vein-lines as above, fascia very diffuse, posterior margin and retinaculum light grey,
remainder of wing black, weakly violaceous. Hind wing below: in 3 mainly pale yellow with black at base
along veins, large black apical patch weakly extended down part of termen, costa and fringe pale yellow;
in 2 exactly like upperside but weakly violaceous.
Fig. 44 ¢$ genitalia. Detail of left clasper. Vitessa nicobarica Hampson.
GENITALIA ¢ (Figs 44, 163, 214). Uncus slender, basal portion flattened, laterally constricted, foramen
posteriorly acuminate, terminal portion narrow, decurved then up-turned with sparse, long, laterally
directed setae, tip broadly pointed. Gnathos with widely arched lateral arms and compressed, relatively
short, talon-shaped medial process, tip finely acuminate. Transtilla broadly arcuate, medially widened.
Tegumen with posterior sclerite broad and rounded, anterior sclerite widely emarginated medially.
Vinculum broad, anteriorly slightly concave. Valve narrow, costa narrowly inflated, apex acutely rounded,
distal margin weakly excised below middle, mesal surface with fine oblique converging striations; clasper
(Fig. 44) a long diagonal ridge with basal curved carina, base of ridge edged with a few points, continuing
to form three tooth-like, medially ridged processes, diminishing in size towards distal margin of valve,
smaller ridges merging with mesal striations; largest process bearing raised serrulate ridge. Juxta with
large narrow lateral sclerites, ventral process long, flat, narrow, up-turned, supporting base of anellus,
dorsal process shorter, broader, bearing setae, connected directly to sclerotized longitudinally ridged
narrow plates supporting sides of anellus; between sclerites and connected medially, a catena divided into
two large, narrow, heavily sclerotized ridges, distally spined and medially separated, basally rounded,
connected by a lightly sclerotized wrinkled element (area damaged); manica supported by lightly
sclerotized bands (anellus not everted in slide preparations). Aedeagus long, distally expanded, base
narrowly rounded; ductus ejaculatorius subbasal; vesica with spines spreading ventrally to distal end of
REVISION OF VITESSIDIA & VITESSA 305
aedeagus, here membrane bulbous and transversely rugose; between cornuti clusters vesica longitudinally
rugose and sclerotized at point of reflexion, probably making an elbow-shaped angle (vesica not properly
everted in slide preparations); cornuti (Fig. 214) in three clusters, two proximal dorsolateral clusters of
same size bearing large spines, one distal ventrolateral cluster bearing smaller spines.
GENITALIA @ (Figs 255, 301). High, posteriorly narrow and produced ovipositor lobes, dorsally fused;
surface densely setose, larger setae on margin, spiny setae easily visible at x 100 magnification. Apophyses
fairly short, slender, weakly curved; anterior apophysis longer and thicker than posterior apophysis.
Posterior ostial sclerite (Fig. 301) wide with short lateral prolongations anteriorly directed, medial portion
prolonged with short constricted base, posterior margin rounded and medially emarginate; anterior
sclerite wide with ventral margin short, recurved, slightly wrinkled and laterally denticulate; very long
dorsal wall of antrum extending with curved margin to base of medial prolongation of posterior sclerite,
anteriorly protruded and funnel-shaped; ductus seminalis arising at junction of antrum with swollen base
of ductus bursae, remainder of ductus bursae narrow, anteriorly spinose; corpus bursae oval, bearing
minute paired denticulate areas.
REMARKS. Despite the unusual lithosiid-like external appearance of this species, it is close in
genital structure to V. pyraliata. It is distinguished by the rounded costa of the fore wing and in
the 3 by the yellow hind wing devoid of a dark terminal band; also on the underside of the hind
wing by the black triangular apical marking; the 9 hind wing is more typical, but has a large
white medial area and diffuse black markings. The 3 genitalia are distinguished by the smaller
talon-shaped gnathos, the narrower valve and the arrangement of ridges and points on clasper,
with very distinctive serrulate ridge on basal process; the divided catena is unique; in the 2
genitalia the shape of the sclerites in the ostial chamber is distinctive.
BIoLoGy. Early stages, host-plant, and season of flight unknown.
DISTRIBUTION (Fig. 21). Nicobar Islands.
MATERIAL EXAMINED
Paralectotypes. Nicobar Islands: 1 9°, data as lectotype (genitalia slide no. 13168; BMNH); 1 3, Great
Nicobars, 873 (genitalia slide no. 14682; BMNH).
Vitessa pyraliata Walker, 1864
(Figs 14, 21, 22, 30, 38, 41, 111-116, 164-166, 196, 197, 256-258, 302-304)
Vitessa pyraliata Walker, 1864 : 220.
3, 2. 46-58 mm. Frons: ¢ black lightly admixed with orange, 2 orange, laterally black. Vertex: ¢ matt
black lightly admixed with orange, 2 orange lightly admixed with black. Eye, ocellus black. Chaetosema
with pale yellow setae. Antenna black, base of scape in 2 orange, dorsal scaling of apical third of shaft
white, last several segments black. Labial palpus: 3, first segment orange, second segment black or
admixed with orange, third segment black; 9, first segment dorsally black, ventrally orange, second seg-
ment black laterally streaked with orange to near tip, third segment black. Maxillary palpus basally black,
distally orange. Basal scaling of proboscis glossy blue-black. Neck region orange. Thorax above: glossy
blue-black, anteriorly bordered with variable amounts of orange, posteriorly tipped with white; patagium
glossy blue-black, laterally and sometimes medially edged with variable amounts of orange; tegula glossy
blue-black, laterally edged with variable amounts of orange. Abdomen above: black, posterior margins of
segments with variable amounts of white, terminal segment posteriorly finely bordered with orange, anal
tuft orange, medially brown. Body below: black; pectus orange; white abdominal lines broad, posterior
margin of terminal segment and anal tuft orange. Legs black, white markings mostly on mid and hind
legs; fore coxa black or black streaked with orange. Fore wing above: ground colour blue-black, fresh
specimens with dark green hue; orange subbasal area variable, in ¢ restricted to small spot at base of cell,
sometimes with orange scaling forming a weak subcostal streak, in 2 sometimes large with broad subcostal
streak continuing to base and joining white spot sometimes present at base of posterior margin, partially
demarcating basal black spot; white antemedial fascia narrow, divided at cell into two spots, in 3 anterior
spot sometimes reduced to minute speck, posterior spot rectangular or triangular, not reaching posterior
margin; white postmedial fascia oblique, divided at Cu, into two spots, large anterior spot circular, over
discocellular vein, not reaching costa, posterior spot triangular in °, minute in 3, sometimes with smaller
spot anterior to it in 2; terminal area with weak white vein-lines, sometimes absent in 3; fringe basally
dark grey, distally light grey. Hind wing above varying with subspecies; black terminal band broad, weakly
306 E. G. MUNROE & M. SHAFFER
violaceous; infuscation from base sometimes extending across wing, almost obliterating white medial area,
otherwise white medial area broadly reaching anal margin; fringe dark grey, distally white at apex and
part of termen. Fore wing below: in ¢ dark brown with shaggy scaling, in 9 blue-black, violaceous, partly
with dark green hue, smoothly scaled, brownish fuscous in cubital region; in both sexes postmedial fascia
as above, or with spots fused, white streak along anal fold; apical white vein-lines weakly developed in
2, sometimes absent as in 3; posterior margin white; retinaculum light grey. Hind wing below: ground
colour as on fore wing, markings as above, apical vein-lines in 2 weak or absent, in 3 absent.
GENITALIA 6 (Figs 30, 38, 41, 164-166, 196, 197). Generally as described for V. nicobarica. Uncus:
marginal setae very long. Gnathos long, strongly curved, sickle-shaped. Transtilla V-shaped. Anterior
sclerite of tegumen with long narrow medial emargination on anterior margin. Vinculum broad, anteriorly
concave. Valve narrow, costa upturned to apex, apex narrowly rounded, termen broadly rounded, slightly
oblique, ventral margin weakly excised below middle; clasper variable, a long diagonal ridge, basal part
with cluster of short spines, remainder produced into three or four tooth-like, medially ridged processes of
irregular size, those on ventral margin smallest and merging with mesal striations. Catena (Fig. 38) single,
large, long, basally constricted, medially sulcate, tips diverging, varying in width posteriorly with multiple
short spines, catena slightly separated from lateral ridged sclerites attached to dorsal processes of juxta
(Figs 30, 196, 197). Aedeagus with strong sclerotization at junction with vesica, forming acute elbow-joint
when vesica everted and reflexed (Fig. 41); vesica bearing three cornuti clusters, two large proximal
dorsolateral clusters, one small distal ventrolateral cluster.
GENITALIA ° (Figs 256-258, 302-304). Generally as described for V. nicobarica. Posterior ostial sclerite
(Figs 302-304) wide with short triangular lateral prolongations anteriorly directed, medial portion of
anterior margin prolonged into a V-shaped flap with rounded upturned tip. Anterior sclerite with ventral
margin short, recurved, slightly wrinkled and laterally denticulate; antrum posteriorly extending to
anterior margin of posterior sclerite, medially narrowed and joined to base of upturned flap; anteriorly
extending into large cup-shaped chamber. Ductus seminalis arising from base of antrum; ductus bursae
slightly widened posteriorly, remainder of ductus narrow, anteriorly spinose; corpus bursae oval bearing
minute denticulate areas, these sometimes absent.
REMARKS. This species is related to V. nicobarica, but is distinguished by possessing typical black
and white hind wing markings in both sexes. It is variable over its range but the two subspecies
V. pyraliata latialbata and V. pyraliata pyraliata are distinguished in the genus by a predominantly
black and white appearance due to great reduction in all pale markings, especially on the fore wing
and the undersides of both wings; the nominate subspecies from Sulawesi is best distinguished
from V. kolakalis, from the same island, by the reduced orange subbasal area, by the rough
underside scaling of the 3 and by the genital characters of both sexes. The subspecies V. pyraliata
triangulifera can easily be confused externally with V. philippina of the stettina-group, but is
distinct on genital characters (see Diagnoses of the'species-group, pp. 286, 303). Other species in the
pyraliata-group are distinguished by their more extensive pale tore wing markings and by genital
characters. We recognize three subspecies, from the following geographical areas: (1) Burma,
Malaya, Sumatra, Java, Natuna Island, Borneo and western central Sulawesi; (2) Philippines;
(3) south and south-western Sulawesi and Flores.
BioLoGy. Early stages and host-plant unknown. Moths collected in all months of the year.
DisTRIBUTION (Figs 14, 21, 22). Burma; Malaya; Sumatra; Java; Natuna Island; Borneo;
Philippines; Sulawesi; Flores.
Vitessa pyraliata latialbata subsp. n.
(Figs: 205 3053810) 12, 164, 19622565302)
3, 2. 46-58 mm. Generally as described for species. Labial palpus: in ¢ second segment laterally admixed
with orange; in 2 mostly orange. Patagium: in ¢ mostly blue-black, laterally and medially edged with
orange; in 2 orange with posteromedial blue-black spot; tegula: in ¢ blue-black with inner and outer
edges finely orange; in 2 more distinctly orange with anterolateral and posterior blue-black spot. Legs in
3 black with mid and hind tibiae bearing small dorsal white spot and distally tipped with white; in 9
most of fore coxa orange, ventral margins of mid and hind femora white, mid and hind tibiae medially
banded with white, tibiae and tibial spurs distally tipped with white. Fore wing above with variable
orange subbasal area, in ¢ restricted between posterior margin of cell and A,, with faint orange scaling or
REVISION OF VITESSIDIA & VITESSA 307
defined subcostal streak continuing to base; in 2 orange area larger, subcostal streak stronger. White
antemedial fascia with fairly well-defined anterior spot and large rectangular posterior spot. White post-
medial fascia sometimes with faint white scales behind posterior spot in 9. Hind wing above with broad,
well demarcated, blue-black terminal band, acutely angled and narrowing at Cu,; infuscation from base
weak, medial white area broadly extended to anal margin. Fore wing below with postmedial fascia diffusely
fused into large oval in 3, spots separate with slight diffusion in 9.
GENITALIA ¢ (Figs 30, 38, 164, 196). Generally as described for species. Clasper with three or four tooth-
like processes, largest being second from base.
GENITALIA ° (Figs 256, 302). As described for species.
REMARKS. This is distinguished from other subspecies by the very large white area of the hind
wing, and the correspondingly minute amount of infuscation from base and costa. it is easily
distinguished from V. teleroma of the suradeva-group, also from Malaysia, by the larger pectina-
tions of the g antenna; the smaller orange subbasal area and white antemedial fascia on the fore
wing; the broader black terminal band on the hind wing; the rough scaling on the underside of
the wings in the 3; and by genital characters (see species-group diagnoses, pp. 303, 317). The
subspecies is slightly variable over its considerable range. The arrangement of spines and
processes on the clasper of the ¢ genitalia is somewhat variable. The data labels of Ceylon [Sri
Lanka] and Australia on two specimens are dubious.
BIoLoGy. Early stages and host-plant unknown. Moths collected from January to July, September,
November and December.
DISTRIBUTION (Fig. 21). Burma; Malaya; Sumatra; Java; Natuna Island; Borneo; western central
Sulawesi.
MATERIAL EXAMINED
Holotype 3, Malaya: Maxwell Hill [4°52’ N, 100°48’ E], 13.ix.1929 (Corbet) (BMNH).
Paratypes. Malaya: 1 3, Penang, 1896 (Curtis) (genitalia stide no. 15832; BMNH); 1 3, Pulo Penang
(‘Penang Island’), ex Boisduval coll. (BMNH); 1 3, Pulo Penang (Teylingen) (UZM, Copenhagen); 1 2,
Pulo Penang (Lévendal) (UZM, Copenhagen); 1 3, 2 3, Taiping, 1972, ili, vili.1973 (HI, Iruma City);
1 3, Pahang, Fraser’s Hill, 1.1955 (Tayleur) (BMNH); 1 9, Pahang, Fraser’s Hill, 27.iv.1958 (Tayleur)
(BMNH); 1 3,3 2, Pahang, Genting Tea Estate, 600 m (‘2000 ft’), 1.iv, 22.vi, 16.x.1976 (Barlow) (BMNH);
7 3, Perak, 550-1200 m (‘1800-4000 ft’), xii.1896—vi.1897 (Curtis) (BMNH); 1 2, Selangore, R[B]ukit
Putus, 900 m (‘3000 ft’), v.1896 (Curtis) (BMNH); 3 2, Selangor, 100 m (‘300 ft’), x.1954 (Banks) (BMNH);
1 3, Perak (Banks) (BMNH); 2 6, 12 2, Bukit Kutu [3°32’ N, 101°44’ E], 1000 m (‘3300 ft’), ix.1927-
vii.1928, and no date (Sanderson) (genitalia slide no.’s 14651, 15833; BMNH); 18 3, 15 2 (including
allotype), Maxwell Hill, 9-21.ix.1929 (Corber) (genitalia slide no. 8261; BMNH); 1 3, Perak, Maxwell
Hill, 1350 m (‘4500 ft’), 1.1955 (Morrell) (BMNH); 2 °, Perak, Maxwell Hill, 1350 m, iii.1958 (Maa)
(BPBM, Honolulu); 6 ¢, 2 2, [Mt] Gunong Ijan (BMNH).
Sumatra: 1 9, south-western Sumatra [?], W. slopes Barisan Range, 750 m, ix—xii.1921 (Pratt) (TM,
Pretoria); 4 3, 2 2, south-western Sumatra, North Korintji Valley, 1500 m, ix-x.1921 (C. F. & J. Pratt)
(BMNH; TM, Pretoria); 1 2, Lebong Tandai, Mt Lalangir, 1050 m, vii.1923 (Brooks); 1 3, 1 2, Fort de
Kock, 900 m, iv.1921, 1924 (Jacobson) (RNH, Leiden); 1 2, east coast, Darat Bangkat, 1909 (De Bussy)
(BMNH). Java: 3 3, Res. [Dutch health resort] Soekaboemi [Sukabumi, 6°58’S, 106°47’ E], 1895
(Ledru) (BMNH); 1 3, Malang (Walsh) (RNH, Leiden); 1 2, no data (BMNH). Natuna Island: 2 3,
Bunguran, vii-x.1894 (Hose) (BMNH). Borneo: 6 3, 8 2, Kalimantan, Dent Province, Mt Marapok
(BMNH); 1 3, 4 9, K., Samarinda, viii-x.1938 (Walsh) (BMNH); 1 3, no data (BMNH); 5 3, Sarawak,
Kuching (BMNH; UM, Oxford); 4 3, S., Kuching (Hewitt) (RSM, Edinburgh); 1 2, S., Matang, 1100 m,
vi.1898 (BMNH); | 9°, S., Gunong Matang, 100 m, ix.1958 (Gressitt & Maa) (BPBM, Honolulu); 1 2,
S., Bidi, 1907-1908 (Brooks) (BMNH); 1 9°, S., Bau District, Bidi, 90-250 m, ix.1958 (Maa) (BPBM,
Honolulu); 2 3, S., Gunong Mulu Nat. Park, Mulu, 1790 m, lower montane (moss) forest, MV — canopy,
1.1978 (Holloway et al.) (BMNH); 1 2, S., Gunong Mulu Nat. Park, Mulu, 1780 m, lower montane forest,
MV - canopy/understorey, i.1978 (Holloway et al.) (BMNH); 3 2, S., Gunong Mulu Nat. Park, Mulu,
150 m, mixed dipt. for. MV — mainly canopy, ii.1978 (Holloway et al.) (BMNH); 2 3, 1 9, S., Gunong
Mulu Nat. Park, Mulu, 1000 m, lower montane for., MV — canopy/understorey, ii.1978 (Holloway et al.)
(BMNH); 1 2, S., Gunong Mulu Nat. Park, Long Pala (Base), 70 m, Alluv./second. for., MV — on batu—
canopy, iii.1978 (Holloway et al.) (BMNH); | 2, S., Gunong Mulu Nat. Park, W. Melinau Gorge, 130 m,
308 E. G. MUNROE & M. SHAFFER
alluvial/Kerangas bank, Acl.—u’ storey, ili.1978 (Holloway et al.) (BMNH); 1 3, Gunong Mulu Nat. Park,
G. Api, 900 m, lower montane forest, MV and Act., iv.1978 (Holloway et al.) (BMNH). Sulawesi: 1 9,
Koelawi Paloe, 950 m, iii.1937 (Kalis) (BMNH). No data: 2 9, Dutch East Indies, no exact data (BMNH);
1 2, 999 (TM, Pretoria).
The following data are dubious: Sri Lanka: | 3, no exact data (BMNH). Australia: 1 3, ‘Australie’, ex
Guenée coll. (BMNH).
Vitessa pyraliata triangulifera subsp. n.
(Figs 14, 113, 114, 165, 257, 303)
3, 2. 50-58 mm. Generally as described for species. Labial palpus in ¢ black; in 9 first segment dorsally
black, ventrally orange, second segment black, with strong lateral orange streak, third segment black.
Patagium and tegula glossy blue-black, latter with anteromedial orange spot. Legs mostly black in 3, in 2°
hind femur ventrally white, mid tibia bearing small dorsal white spot, mid and hind tibiae tipped with
white. Fore wing above: orange subbasal area restricted from below costa to A,, smaller in $. White ante-
medial fascia with anterior spot minute, mostly anterior to cell, posterior spot large, triangular, from
behind cell to just past A,. White postmedial fascia sometimes with faint white scales anterior and posterior
to posterior spot in 2. Hind wing above with broad, well demarcated, blue-black terminal band, acutely
angled and narrowing at Cu,; infuscation from base fairly heavy, restricting white medial area to about
half of anal margin. Fore wing below: postmedial fascia, spots separate with slight diffusion.
GENITALIA ¢ (Fig. 165). Generally as described for species. Clasper with two tooth-like processes, the
more basal one the larger.
GENITALIA @ (Figs 257, 303). As described for species.
REMARKS. The external characters are much as in V. pyraliata latialbata, but the posterior white
spot of the antemedial fascia on the upperside of the fore wing is smaller and triangular, not
rectangular; the orange subbasal area is small in both sexes and lacks the subcostal streak to the
base. In the hind wing this subspecies is distinct from V. pyraliata latialbata in having the white
area more constricted by infuscation from base, especially in the J, but not so constricted as in
the nominate subspecies. The 9 is hardly distinguishable in external appearance from V. philippina
of the stettina-group, which occurs sympatrically (the 3 of V. philippina is unknown); V. philippina
is smaller, with orange patagium and larger orange subbasal area, nearly reaching costa; its
genitalia differ by the species-group characters (see Diagnosis, p. 286).
BioLoGy. Early stages and host-plant unknown. Moths collected in February, May and August
to November.
DISTRIBUTION (Fig. 14). Philippines.
MATERIAL EXAMINED
Holotype 3, Philippines: Luzon, Benguet, Baguio (Baker) (NMNH, Washington).
Paratypes. Philippines: 1 9 (allotype), data as holotype (NMNH, Washington); | 2, Luzon, Tabato,
ii.1916 (NMNH, Washington); 14 9, Luzon, Mt Makiling (Baker) (genitalia slide no. 342 DK) (NMNH,
Washington; CNC, Ottawa); 1 3g, Luzon [with conflicting labels], Mt Makiling, 350m (‘1070 ft’),
12.ix.1936 (Salmon) [and] Los Banos, 22.viii.1921 (Pearson) (NMNH, Washington); 7 2, Luzon, Manila,
19. viii-16.x.1912 (Wileman) (genitalia slide no.’s 14704, 14705, 15144, 15145, 15835; BMNH); 1 °, Luzon
prov. Rizal, Manila, plains, v.1912 (Wileman) (genitalia slide no. 15834; BMNH); 4 2, Mindanao, Surigao,
xi. (NMNH, Washington; CNC, Ottawa); 1 2, Mindanao, 15 km south-west of Davao, Mt Apo School,
500 m, x.1955 (Davis) (NMNH, Washington); 1 2, Mindanao, Lanao, Butig Mts, 24km NE. of Butig,
1080 m, in rain forest, 21.vi.1958 (Milliron) (BPBM, Honolulu); 1 3, no data (Type no. 2636; genitalia
slide no. Oxford 723-1977, Shaffer; UM, Oxford).
Vitessa pyraliata pyraliata Walker, 1864
(Figs 22, 41, 115, 116, 166, 197, 258, 304)
Vitessa pyraliata Walker, 1864, List Specimens lepid. Insects Colln Br. Mus. 31 Supplement: 220.
LECTOTYPE 2 [not ¢ as stated by Walker], SULAWEsI (‘Celebes’) [(Wallace)] (genitalia slide no.
15828; BMNH), here designated [examined].
Vitessa pyraliata Walker; Ragonot, 1891 : 110.
Vitessa pyraliata Walker; Hampson, 18966 : 503.
Vitessa pyraliata Walker; Swinhoe, 1900 : 429.
REVISION OF VITESSIDIA & VITESSA 309
3, 2. 46-58 mm. Generally as described for species. Labial palpus in 3: second segment lightly admixed
with orange; in 2 second segment laterally orange to near tip. Patagium in ¢ glossy blue-black; in @
glossy blue-black, finely edged medially and laterally with orange. Tegula glossy blue-black with orange
anteromedial spot. Legs in 3: mid tibia with medial dorsal white spot; mid and hind tibiae tipped with
white; in 2, ventral surfaces of mid and hind femora white; medial white band and white tips on mid and
hind tibiae. Fore wing above: orange subbasal area variable in size, from behind costa to CuP, smaller in
3g. White antemedial fascia with anterior spot minute, anterior to cell; posterior spot large, roughly tri-
angular, from cell to just past A,. White postmedial fascia with very large round spot over end of cell and
discocellular vein, posterior spot on CuP minute. Hind wing above with broad, well-demarcated, blue-
black terminal band in both sexes, with heavy infuscation from base; white medial area narrow in 2,
very narrow in 34, almost to obliteration. Fore wing below: postmedial fascia forming large oval in 3,
two spots in 2. Hind wing below: similar to upperside of 2, sometimes restricted white area to centre of
wing cut off from anal margin by infuscation.
GENITALIA 6 (Figs 41, 166, 197). Generally as described for species. Clasper with two or three tooth-like
processes, largest being second from base.
GENITALIA 2 (Figs 258, 304). As described for species.
REMARKS. This subspecies is similar in size and external appearance to V. pyraliata latialbata,
but is distinguished by the much-restricted white area of the hind wing in both sexes; it is dis-
tinguished from V. pyraliata triangulifera by the much-reduced posterior white spot of the
postmedial fascia. It strongly resembles V. kolakalis, with which it flies over part of its range, but
is distinguished by the smaller orange subbasal area and by the genitalia of both sexes, with clasper
of valve in 3 and without sclerotized depressed signa on corpus bursae in 9. We have not traced
the paralectotype from Seram collected by Wallace, mentioned by Walker in his description of
pyraliata. It is probably not conspecific with the lectotype. All available Wallace specimens from
the BMNH and UM, Oxford, collections have been examined. Swinhoe (1900 : 429) recorded
two specimens from Makian, an island in the Moluccas south of Ternate, but these have not
been traced.
BioLoGy. Early stages and host-plant unknown. Moths collected in February, March, May, June
and October.
DISTRIBUTION (Fig. 22). South-west and south-east Sulawesi; Flores.
MATERIAL EXAMINED
Sulawesi: 1 9°, Makasar (Wallace) (UM, Oxford); 2 3, South Celebes (Everett) (BMNH); 1 3, 1 9,
Bonthain, Indrulaman, 700 m, x.1895 (Everett) (BMNH); 4 9, near Maros, Tjamba, 450 m, ii.1938
(Kalis) (BMNH); 3 2, near Maros, Pangean, 600 m, 111.1938 (Kalis) (BMNH); 6 2, Goa, Malino, 1100 m,
vi.1938 (Kalis) (BMNH); 1 2, Tombugu [Boengkoe 2°32’ S, 121°58’ E](MNHN, Paris); 1 2, Ulu Kolaka,
500 m, v—vi.1939 (Kalis) (BMNH). Flores: 1 3, no exact data (Everett) (BMNH); 1 9°, no exact data
(Wallace) (UM, Oxford).
Vitessa kolakalis sp. n.
(Figs 22, 117, 118,167, 198, 259; 305)
3, 9. 46-50 mm. Generally as described for V. pyraliata. Frons and vertex orange. Antenna black, in 3
white on terminal region more extensive. Labial palpus: first two segments orange, third segment black,
buff-tipped. Patagium orange with medial blue-black spot in 3, anteromedial in °. Tegula blue-black,
anteromedially orange at extreme base, inconspicuous in 3, more obvious in 9. Abdomen above: black, a
few light scales on posterior margins of some segments; anal tuft orange. Abdomen below: black, segments
with broad white posterior bands, narrowed or interrupted medially. Legs black, coxae with scattered
white spots; hind femur ventrally white; mid tibia with dorsal white spot; tips of mid and hind tibiae
and of spurs white. Fore wing above: orange subbasal area broad, narrowly short of costa, extending to
A,; white streak at base of posterior margin; white antemedial fascia broadly divided at cell, anterior spot
short of costa, posterior spot larger, triangular, short of posterior margin; white postmedial fascia divided
at Cu., large anterior spot round, posterior spot more distal, triangular, medially directed; apical white
vein-lines weak in 3, strong in 9, short of termen and postmedial fascia; fringe dark grey. Hind wing above:
medial white area narrowly restricted; broad blue-black terminal band, medial edge angled at base of cell;
310 E. G. MUNROE & M. SHAFFER
infuscation from base stronger in 3; fringe dark grey, distally white at apex. Fore wing below: postmedial
fascia narrowly divided into two spots; apical vein-lines weak in 9, absent in 3; pale antemedial streak on
anal fold; without shaggy scaling in g. Hind wing below as above.
GENITALIA ¢ (Figs 167, 198). Generally as described for V. pyraliata. Uncus slender, evenly tapering, basal
portion wider, basally waisted, foramen wide, posteriorly rounded; lateral setae of moderate length.
Gnathos with medial process wide, short, strongly angled, hook-shaped. Transtilla arcuate. Valve narrow,
costa very broadly inflated, apex and termen broadly angled, posterior margin strongly excised ante- and
postmedially; strong subcostal ridge to apex; mesal surface with fine oblique striations, more lateral near
clasper; clasper a long diagonal ridge, continuous with straight carina, free edge rising gradually to near
posterior margin, rising abruptly, forming single, slightly recurved, acuminate process, the falling distal
edge joining posterior margin of valve at upper end of postmedial excision. Sclerotized dorsal surface of
sclerite on anellus reticulated (Fig. 198); broadly rounded emargination at base of sclerotized manica,
filled with very wide dorsal catena; catena slightly narrowing basally and longitudinally ridged, distally
spined.
GENITALIA 2 (Figs 259, 305). Generally as described for V. pyraliata. Posterior ostial sclerite (Fig. 305)
with V-shaped medial flap on anterior margin, recurved, extending above posterior margin. Antrum
very wide posteriorly, slightly asymmetrical, box-shaped. Corpus bursae globular, with single round,
depressed, denticulate signum and single sclerotized, diffuse, denticulate patch.
REMARKS. In Sulawesi this species is easily confused with V. pyraliata pyraliata, but is dis-
tinguished by the largely orange patagium, the wider white area on the hind wing of the 3; on
the fore wing by the more distinct spots of both fasciae in the 3 and the larger orange subbasal
area in both sexes; in the ¢ by the absence of shaggy scaling on the undersides of the wings; the
genitalia have larger spines on the cornuti clusters in the 3 and different ostial sclerotization in the
Q. V. kolakalis is distinguished from V. hollandi wallacealis of the stettina-group, which occurs in
Sulawesi, by the smaller white markings on the fore and hind wings; similarly it differs from all
other members of the pyraliata-group by the reduced markings. The ¢ genitalia differ from
those of other members of the pyraliata-group (so far as known) by the shape of the valve and
clasper and the size of the dorsal catena; the 2 genitalia differ in the asymmetrical development
of the signa. The species is most closely related to V. pyraliata and V. sulaensis.
BioLoGy. Early stages and host-plant unknown. Moths collected from May to June.
DISTRIBUTION (Fig. 22). South-eastern Sulawesi.
MATERIAL EXAMINED
Holotype 3, Sulawesi: Ulu Kolaka, 500 m, v—vi.1939 (Kalis) (genitalia slide no. 8265; BMNH).
Paratypes. 10 3, 6 % (including allotype), data as holotype (genitalia slide no.’s 14699, 14700, 15836,
15837, 15838, 15839; BMNH).
Vitessa sulaensis sp. n.
(Figs 22, 119, 260, 306)
3. Unknown.
2. 46 mm. Generally as described for V. pyraliata and V. kolakalis. Frons and vertex orange. Antenna
black, base of scape orange, terminal third of dorsal scaling of shaft white, last few segments fuscous.
Labial palpus: first two segments orange, third segment fuscous. Patagium orange with anteromedial
blue-black spot. Tegula blue-black, anteromedially orange at extreme base. Abdomen above: black,
posterior margins of segments narrowly white, anal tuft orange. Abdomen below: like upperside, wider
segmental white margins. Legs black, hind femur ventraliy white; mid tibia with dorsal white spot; tips
of mid and hind tibiae white. Fore wing rather wide; above: ground-colour blue-black with dark-green
hue; orange subbasal area almost circular, narrowly short of costa, extending to A,; white scales at base
of posterior margin; white antemedial fascia divided at cell into two spots, anterior spot small, short of
costa, posterior spot larger, broad, triangular, short of posterior margin; postmedial fascia divided at Cu,
into two spots, anterior spot round, very large, posterior spot more distal, triangular, medially directed,
with minute spot anterior to distal end; five short but wide vein-lines in apical area, widely short of
termen and postmedial fascia; fringe dark grey. Hind wing above: base and costa narrowly infuscated;
terminal band blue-black, violaceous, broad, tapering to anal margin, inner border irregular; anal margin
edged with black; white medial area broadly extended to anal margin; fringe [worn] dark grey. Fore wing
REVISION OF VITESSIDIA & VITESSA 311
below: postmedial fascia as above, more diffuse, posterior spot less distal; isolated white scales at apex;
retinaculum light grey. Hind wing below as above.
GENITALIA ° (Figs 260, 306). Generally as described for V. pyraliata. Posterior ostial sclerite (Fig. 306)
with low, broad, quadrangular-based, V-shaped medial flap on anterior margin, narrowly recurved.
Dorsal wall of antrum broadly extended posteriorly, distinctly trapezoidal and broadly connected to
V-shaped flap on posterior sclerite; sclerotization narrowly extended anteriorly, with concave end formed
within funnel-shaped expansion of ductus bursae. Corpus bursae globular, with an unequal pair of
elongate, depressed denticulate signa.
REMARKS. This species is close to V. pyraliata, but differs externally in the more distal posterior
white spot of the postmedial fascia of the fore wing; it differs from V. kolakalis by the more
extensive white wing markings, and from remaining species in the group by the broader black
terminal band of the hind wing. It is distinct from all other species of the group in genital
characters.
BioLoGy. Early stages and host-plant unknown. Holotype collected in October.
DISTRIBUTION (Fig. 22). Moluccas: Sanana (‘Sula Besi’).
MATERIAL EXAMINED
Holotype °, Moluccas: Sula Besi, x.1897 (Doherty) (genitalia slide no. 14628; BMNH).
Vitessa muluana sp. n.
(Figs 22, 120, 261, 307, 318)
3. Unknown.
2. 52 mm. Frons, vertex orange. Eye, ocellus black. Chaetosema with pale yellowish setae. Antenna
black, ventral and inner surfaces of scape orange; dorsal scaling of apical third of shaft white. Labial
palpus, first two segments orange, dorsal surfaces admixed with black, third segment black. Maxillary
palpus basally black, distally orange. Basal scaling of proboscis basally black. Neck region orange. Thorax
above: orange with medial 8-shaped blue-black spot; patagium orange with small anteromedial blue-black
spot; tegula orange, anterolaterally and distally blue-black. Abdomen dark brown, anteriorly matt,
posteriorly glossy. Body below: pectus orange; thorax black, flecked with orange; legs blue-black, fore
and hind coxae mostly orange, mid and hind femora ventrally creamy white, mid and hind tibiae with
medial and distal creamy white spots. Fore wing broad; ground-colour blue-black, violaceous; orange
subbasal area large, narrowly short of costa, obliquely extended to near posterior margin, narrow sub-
costal streak extending to base, partially demarcating blue-black basal spot, medial margin jagged;
yellowish white streak at base of posterior margin; creamy white antemedial fascia divided at cell, small
triangular anterior spot subcostal, posterior spot broadly triangular, short of posterior margin, medially
diffuse; creamy white postmedial fascia divided at Cu, anterior spot large, round, over discocellular vein,
posterior spot a broad dash, more distal and medially diffuse; smaller streaks anteriorly and posteriorly
at distal end; an additional minute spot between M, and Cu,; terminal area with subapical white vein-lines
extending to Cu,; fringe dark grey. Hind wing above: infuscation from base and costa very narrow;
terminal band blue-black, violaceous, broad at apex, acutely angled to past Cu,, narrowing and finely
terminating at anal margin; anal margin finely edged with black; white medial area very broad, extending
over most of anal margin; fringe basally dark grey, distally light grey at apex, white to anal margin. Fore
wing below: black, slightly violaceous; posterior margin basally white; white postmedial fascia with spots
less clearly defined than above, finely separated at Cu., basally diffuse; retinaculum light grey. Hind wing
below: as above with weak white vein-lines to Cuz.
GENITALIA © (Figs 261, 307, 318). High, wide, posteriorly prolonged ovipositor lobes, dorsally fused,
surface densely setose, longer setae on margin, spiny setae conspicuous at * 25 magnification (Fig. 318).
Apophyses fairly short, slightly curved, tips truncated, anterior pair thicker. Posterior ostial sclerite
(Fig. 307) wide, posterior margin with medial sclerotization weak and partially emarginated; sclerite
anteriorly tripartite, short lateral prolongations ending anteriorly in tooth-like points; medial section
also tripartite, with short triangular, anteriorly directed lateral prolongations, medial area concave;
ventrally, a shallow, slightly oblique, lateral pouch extending from mid way along prolongation of medial
area to lateral margin of sclerite, free edge finely denticulate. Anterior sclerite with ventral margin very
short, sclerotization strongest laterally, broadly wrinkled, inner surface finely denticulate, medially
membranous; antrum mostly anterior to ventral margin, asymmetrical, very narrow, about half width of
312 E. G. MUNROE & M. SHAFFER
posterior sclerite; posteriorly pyramidal, but last third quadrangular and fused to concave medial area of
posterior sclerite; anteriorly with a weak transverse fracture-line in sclerotization, separating pyramidal
area from asymmetrical, funnel-shaped and collar-like area, ventral surface of latter present on one side
only, triangular. Ductus bursae narrowly expanded posteriorly, twisted to one side at junction with
antrum, short duct anteriorly spinose; corpus bursae large and bearing a pair of sclerotized, depressed,
irregularly shaped, denticulate signa.
REMARKS. We do not know the ¢ of this striking and distinctive species and its systematic position
is as yet not fully ascertained; it may well need its own species-group. The white markings of the
fore wing suggest relationship with the pyraliata-group, particularly with V. nicobarica, though
they are closest in appearance to those of V. teleroma of the V. suradeva-group. The narrowness
of the terminal band of the hind wing closely resembles those of the V. plumosa-group and V.
teleroma. In the genitalia the antrum bears some resemblance to that of V. gemina, of the present
group; the signa show affinity to this group as well; the spiny setae, which tend to be well
developed in the pyraliata-group, are more conspicuous than in any other species of the genus.
BioLoGy. Early stages and host-plant unknown. Moths collected in March and August to
December.
DISTRIBUTION (Fig. 22). Borneo: Sarawak; Sabah.
MATERIAL EXAMINED
Holotype °, Borneo: Sarawak, Mt Mulu, 300-1200 m (‘1000-4000 ft’), viii—xii.1894 (Hose) (genitalia
slide no. 15846; BMNH).
Paratypes. Borneo: | 2, Sabah, Park HQ, Mt Kinabalu State Park, 1550 m, 20.iii.1975, m/v light
(Munroe) (genitalia slide no. 5196 DK; CNC, Ottawa).
Vitessa gemina sp. n.
(ligs:3.. 1201222624308)
3. Unknown.
2. 42-48 mm. Frons, vertex orange, latter admixed with black. Eye, ocellus brown. Chaetosema with
pale yellowish setae. Antenna black, ventral and inner surfaces of scape orange, dorsal scaling of apical
third of shaft heavily admixed with white. Labial palpus: first two segments orange, third segment black.
Maxillary palpus basally black, distally orange. Basal scaling of proboscis black, heavily admixed with
orange. Neck region orange. Thorax above: blue-black, anteriorly and posteriorly orange; patagium
orange, with large, posteromedial, glossy blue-black spot; tegula orange, with very large anterolateral and
terminal glossy blue-black spots. Abdomen above: blackish fuscous, anal tuft orange. Body below: pectus
orange; thorax blue-black, large yeliow spot below tegula; abdomen with posterior margins of segments
narrowly creamy white, anal tuft orange. Legs black, partly with dark green hue; fore coxa dorsally flecked
with orange, ventrally white; mid and hind coxae with whitish grey bands; mid and hind femora ventrally
white; mid tibia with dorsal medial white spot; mid and hind tibiae tipped with white. Fore wing of
moderate width; above: ground colour glossy blue-black; orange subbasal area large, narrowly short of
costa, obliquely extended to near posterior margin, narrow subcostal streak, sometimes incomplete,
extending to base, partially demarcating large blue-black basal spot, medial margin broadly indented at
cell and A,; white antemedial fascia divided at cell, small anterior triangular subcostal spot, large posterior
triangular spot short of posterior margin, both medially directed; white postmedial fascia divided at
Cuz, anterior, large, round spot over discocellular vein, posterior, small, diffusely triangular, more distal
spot; terminal area with white vein-lines on most veins, indistinct after Cu,, reaching termen, but narrowly
short of postmedial fascia; fringe dark grey. Hind wing above, small amount of infuscation from base and
narrowly from costa, terminal band blue-black, violaceous, broad at apex, acutely angled to past Cu,
very narrow at termen, abruptly terminating at anal margin; anal margin finely edged with black; white
medial area wide, broadly extended to anal margin; fringe basally dark grey, distally light grey. Fore wing
below: black, slightly violaceous; posterior margin basally white; white postmedial fascia with spots less
clearly defined than above, finely separated at Cu., basally diffuse; weak white apical vein-lines to M,;
retinaculum dark grey. Hind wing below as above, weak vein-lines to Cu.
GENITALIA ° (Figs 262, 308). High, broad, posteriorly prolonged ovipositor lobes, dorsally fused, surface
densely setose, longer on margin; spiny setae fairly conspicuous at x 25 magnification. Posterior apophysis
short, slender, slightly curved, tip spatulate. Anterior apophysis long, thick, curved, tip spatulate. Posterior
REVISION OF VITESSIDIA & VITESSA a13
ostial sclerite (Fig. 308) broad, long, with wide lateral prolongations, each with two broad transverse
constrictions; posterior margin with shallow medial emargination; anterior margin with medial area
sharply differentiated into a laterally constricted quadrangle, demarcated posteriorly by a convex fold.
Anterior sclerite broad, with ventral margin recurved, heavily wrinkled; posterior dorsal wall of antrum
medially prolonged, narrowly dome-shaped, connected to medial quadrangular area of posterior sclerite;
dorsal and ventral wall of antrum narrowly extended anteriorly, slightly asymmetrical, broadly concave
and funnel-shaped at junction with expanded end of ductus bursae; ductus bursae slightly convoluted
posteriorly, spinose anteriorly; corpus bursae oval, bearing minute paired denticulate areas.
REMARKS. This species closely resembles V. hollandi hollandi of the stettina-group in external
appearance, and can be distinguished reliably only by examination of the genitalia. The 9 genitalia
are distinct from those of the rest of the pyraliata-group.
BioLoGy. Early stages and host-plant unknown. Moths collected between May and June and
between August and September.
DISTRIBUTION (Fig. 3). New Guinea: Irian Jaya (Kepulauan Schouten).
MATERIAL EXAMINED
Holotype 2, New Guinea: Irian Jaya (Kepulauan Schouten), Mefor (“Numfoor’), Suer, v—vi.1897 (Doherty)
(genitalia slide no. 14638; BMNH).
Paratypes. New Guinea: 3 9, Irian Jaya (Kepulauan Schouten), data as holotype (genitalia slide no.’s
14684, 15139, 15840; BMNH); 1 9, I. J., Mefor Island, Geelvink Bay, 15.viii-10.ix.1920 (C., F., & J
Pratt) (TM, Pretoria); 3 9, 1. J., Biak, 1896 (Doherty) (genitalia slide no. 14683; BMNH).
Vitessa splendida Schultze, 1908
(Figs 14, 123, 263, 309)
Vitessa splendida Schultze, 1908, Philipp. J. Sci. 3 (1) A : 35, pl. 1, fig. 11. Holotype 9°, PHILIPPINES:
Negros Cccidental, Maao, xi.1902 (Banks) (originally in Entomological Collection, Bureau of Science,
Manilla; now destroyed).
3. Unknown.
2. 44-46 mm. Frons, vertex orange. Eye, ocellus black. Chaetosema with pale yellowish setae. Antenna
black, ventral surface of scape orange, dorsal scaling of apical third of shaft [presumably] white [antennae
damaged]. Labial palpus: first two segments orange, third segment black. Maxillary palpus basally black,
distally orange. Basal scaling of proboscis blue-black. Neck region orange. Thorax above: orange, large
blue-black medial spot, posteriorly tipped with yellowish white; patagium glossy blue-black, laterally
tipped and medially edged with orange; tegula orange with longitudinal, broad, glossy blue-black stripe
along entire length. Abdomen above, matt fuscous; posterior margins of segments narrowly white, anal
tuft orange. Body below: pectus orange; thorax black, banded with white; abdomen with posterior
margins white, broader than above, anal tuft orange. Legs blue-black, partly with dark green hue; fore
coxa dorsally flecked with orange, ventrally white; mid and hind coxae spotted with white; mid and hind
femora ventrally white; mid and hind tibiae with medial bands and tips white, outer surfaces of spurs
white; hind tarsus with dorsal white spot at end of first segment. Fore wing of moderate width; above:
ground-colour glossy, green-black ; orange subbasal area large, narrowly short of costa, short of posterior
margin but passing A,, medial margin sinuate, narrow streaks at subcostal and A, regions extended to base,
the subcostal streak broader, the anal streak weak and sometimes parted at centre, the streaks demarcating
a blue-black oval basal spot; creamy white antemedial fascia divided by a wedge at cell, anterior spot
subcostal, narrowly triangular, medially directed, posterior spot larger, basal margin level with anterior
spot, slightly short of posterior margin, broadly triangular, medially diffuse; creamy white postmedial
fascia divided at Cu,., large round anterior spot over discocellular vein, posterior spot smaller, narrowly
triangular, more distal, medially directed, minute white scaling anterior and posterior to terminal edge;
terminal area with strong creamy white vein-lines, extending from termen but short of postmedial fascia;
fringe dark grey. Hind wing above: slight basal and narrow costal infuscation; violaceous blue-black
terminal band, broad at apex, evenly tapering to anal margin, white medial area large, broadly extended
to white anal margin; fringe basally dark grey, distally white at apex. Fore wing below: black, slightly
violaceous; weak subcostal white streak from base; posterior margin shiny yellowish fuscous, broadly
white at base; well-developed white postmedial spots; weak white apical vein-lines; fringe dark grey;
retinaculum light grey. Hind wing below as above, weak white apical vein-lines; fascia white, dark grey
at base of termen.
314 E. G. MUNROE & M. SHAFFER
GENITALIA & (Figs 263, 309). High, wide, posteriorly prolonged ovipositor lobes, dorsally fused, surface
densely setose, larger setae on margin, spiny setae visible but not conspicuous at 25 magnification.
Apophyses short, slightly curved, tips bluntly spatulate, anterior pair thicker. Posterior ostial sclerite
(Fig. 309) very broad, densely sclerotized, with long, wide, anteriorly directed, lateral prolongations;
posterior margin heavily sclerotized, medially swollen and slightly recurved, curved transverse pleats
converging on swollen margin; anterior margin medially with short lateral prolongations joining antrum.
Anterior sclerite very broad, ventral margin long, recurved, medially wrinkled, laterally minutely spinose
on membranous area; posterior dorsal wall of antrum with posterior edge recurved, converging broadly
and joining medial margin of posterior sclerite; very long anterior extension of antrum symmetrical,
heavily sclerotized, dorsal wall broadly bowl-shaped, ventral wall with deep triangular folds on either
side of opening of ductus bursae; posterior end of ductus bursae slightly swollen and pleated, remainder
minutely spinose, merging gradually with oval corpus bursae; the latter bearing a single denticulate area.
REMARKS. The wing markings resemble those of V. plumosa and V. suradeva but differ in having
less extensive white fasciae, and in the fasciae not being joined by medial streaking; the white
terminal vein-lines are close to, but do not join the postmedial fascia; the shorter orange subbasal
area does not reach the posterior margin. Superficially the species resembles other Philippine
species: V. philippina of the stettina-group and V. pyraliata triangulifera of the pyraliata-group,
but it differs by the larger orange subbasal area and larger white medial area of the hind wing.
This species is very distinct on genital characters. The holotype of V. splendida was destroyed
during the Second World War, but it was well figured by Schultze and the NMNH specimens have
at one time been compared with the holotype. As material from Negros Island is not available,
we do not designate a neotype.
BIoLoGy. Early stages and host-plant unknown. Moths collected in May, August, September and
November.
DISTRIBUTION (Fig. 14). Philippines: Manila to Mindanao and Palawan.
MATERIAL EXAMINED
Philippines: 1 2, Luzon, Manila, ix.1912 (Wileman) (BMNH); 3 2, Luzon, Mt Makiling (Baker) (NMNH,
Washington; CNC, Ottawa); 1 2°, Mindanao, Surigao, xi (BMNH); | 2, Mindanao, Subprov. Lanao,
Kolambugan, sea level, v.1914 (Wileman) (BMNH); | °, Palawan, Brookes Point, Uring Uring, vili.1961
(ZM, Copenhagen).
The plumosa-group
DIAGNosIs. Wing markings basically as in the stettina- and suradeva-groups, but white antemedial fascia
with two long acuminate streaks on R and Cu, bordering a V-shaped indentation, extending to postmedial
fascia; white postmedial fascia divided into an oval anterior spot and two distinct, narrowly triangular,
basal spots, anterior and posterior to A, ... 3 genitalia with distal portion of uncus down-curved then
up-turned, narrowed, with laterally directed setae, as in the pyraliata-group and several others; valve with
slight distal expansion; clasper with prominent, bifid, anteromedially directed process, not paralleled in
other groups; base of long oblique clasper showing affinities with the hemiallactis- and pyraliata-groups.
Ostial chamber of 2 genitalia with long ventral margin of anterior sclerite more strongly developed than
in any other species-group.
Vitessa plumosa Hampson, 1896
(Figs 22, 31, 39, 124-127, 168, 169, 199, 200, 216, 264, 265, 310, 311)
Vitessa plumosa Hampson, 18966 : 503.
3, 2. 38-52 mm. Frons, vertex orange. Eye, ocellus brown. Chaetosema with pale yellowish setae. Antenna
black, anterior surface of scape basally orange; dorsal scaling of apical third of shaft black, sometimes
with isolated white scales, exceptionally with an extensive white zone; terminal segments fuscous. Labial
palpus: first two segments orange, sometimes admixed with black, third segment black. Maxillary palpus
basally black, distally orange. Basal scaling of proboscis blue-black. Neck region orange. Thorax above:
orange, 8-shaped blue-black medial area, posteriorly tipped with pale yellow; patagium orange with
variable-sized, posteromedial, glossy blue-black spot; tegula orange with variable-sized, longitudinal,
glossy blue-black medial band. Abdomen above: black, posterior margins of segments broadly white,
REVISION OF VITESSIDIA & VITESSA 315
posterior segment blue-black, laterally orange, anal tuft orange. Body below: black with greenish hue;
pectus orange; white abdominal margins broader than above; posterior half of terminal segment and anal
tuft orange. Legs black with greenish hue; fore coxa dorsally orange, ventrally white; mid and hind coxae
banded with white; fore femur ventrally edged with orange; mid and hind femora ventrally white; mid
and hind tibiae medially banded and tipped with white; tibial spurs white; hind tarsus with dorsal posterior
tip bearing white dot. Fore wing rather narrow; above: ground-colour glossy blue-black with dark green
hue; orange subbasal area large, narrowly short of costa and posterior margin, medial margin sinuate,
sometimes with subcostal medial extension, subcostal streak extending to base, basal margin protruding
at CuP, basal blue-black spot partly demarcated; white antemedial fascia with basal margin incurved,
especially towards costa, narrow black medial wedge at cell sometimes dividing fascia, anterior spot a
narrow triangular streak, medially directed and acuminate, sometimes touching postmedial fascia,
posterior area with medially directed and acuminate triangular extension touching postmedial fascia, base
narrowing to near posterior margin; white postmedial fascia distinctly divided into four markings,
divisions at Sc, Cu, and A,, a long thin subcostal line, a large oval discocellular spot with slight terminal
indentation, posterior to this a narrowly triangular, medially directed spot, nearly touching antemedial
fascia, shorter in 3, the most posterior streak slightly thinner, short of posterior margin, terminal margin
of fascia slightly oblique or at right angles to posterior margin; terminal area with thick white vein-lines
extending from termen to postmedial fascia; fringe dark grey. Hind wing above: narrow infuscation from
costa, violaceous blue-black terminal band, broad at apex, gradually tapering or strongly angled at Cu,
and narrow to anal margin; large white medial area, broadly extending to white anal margin; fringe basally
grey, distally white. Fore wing below: ground-colour blue-black, violaceous; orange-brown area around
white retinaculum; posterior margin broadly white from base; short white subcostal streak from base,
lightly admixed with yellow; white postmedial fascia with large discocellular spot indented terminally, a
smaller, less distinct, more distal spot posterior to Cu,; white vein-lines apically, minutely at termen to
Cuz; fringe grey. Hind wing below as above, white along costa apically, apical vein-lines continued
minutely on termen to Cu,; fringe white.
GENITALIA ¢ (Figs 31, 39, 168, 169, 199, 200, 216). Uncus long, slender, narrowly constricted at tegumen,
with pronounced lateral flanges; base slightly constricted, gradually tapering, bearing long narrow
foramen; distal portion rod-like with sparse, short lateral setae, de-curved then up-turned to acuminate
tip. Gnathos with wide lateral arms, converging to a long, compressed, acuminate, scythe-shaped medial
process, not as long as uncus. Transtilla arcuate, medially broadened, posterior margin sometimes medially
excised, lateral flexure pronounced. Tegumen with posterior sclerite broad, rounded; anterior sclerite
with very broad medial emargination. Vinculum broad, basally concave. Valve narrow, base narrowly
inflated, costa sinuate, apex pronounced, narrowly rounded, termen broadly rounded, posterior margin
sinuate towards base; mesal surface with finely converging oblique striations; clasper with oblique base
separated from crease-like carina, base abruptly upturned, forming prominent, bifid, anteromedially
directed process, distal side curving gradually, forming raised serrate ridge terminating close to posterior
margin shortly before terminal angle. Juxta with large, narrow lateral sclerites (Fig. 31), very long, flat,
narrow, upturned, ventral process; dorsal process broader, shorter, bearing setae, directly connected to
ridged sclerite on anellus (Figs 199, 200); large narrow catena (Fig. 39) between sclerites, heavily sclero-
tized, constricted anteriorly, deeply ridged, tips laterally diverging, multiple spines posteriorly; manica
supported by sclerotized bands. Aedeagus long, distally expanded, base narrowly rounded; ductus
ejaculatorius subbasal; vesica with spines spreading ventrally to distal end of aedeagus, here membrane
bulbous and transversely rugose; vesica sharply tapering distally; dorsally spinose, longitudinally rugose,
lightly sclerotized; sclerotized junction with aedeagus forming a sharp elbow-joint when vesica everted
and reflexed; three cornuti clusters (Fig. 216); two dorsolateral; the left with long, short-spined sclerotized
bar, large spines posteriorly, the right lacking bar, smaller patch of spines posteriorly; one ventrolatera!
cluster with large spines.
GENITALIA ° (Figs 264, 265, 310, 311). High, wide, posteriorly prolonged ovipositor lobes, dorsally fused ;
surface densely setose, longer setae on margin, spiny setae visible but not conspicuous at x 25 magnifica-
tion. Apophyses slightly curved, tips truncate; anterior pair longer, thicker. Posterior ostial sclerite (Figs
310, 311) broad with long lateral prolongations; posterior margin medially emarginate, thickened, slightly
recurved; sclerite with lateral transverse ridge and oblique, shallow, pouch-like invagination; lateral
prolongation constricted, with ventromedial edge deeply recurved; anterior margin medially elongated,
bearing small recurved flap. Anterior sclerite with very long ventral margin, half as wide as sclerite,
recurved, wrinkled, minutely spinose laterally; antrum posteriorly broad and rounded, margin joined
to recurved edge of lateral prolongation and medial flap of posterior sclerite; anterior development long,
forming large funnel-shaped structure, ventral wall producing sclerotized base of ductus seminalis, dorsal
316 E. G. MUNROE & M. SHAFFER
wall with deep V-shaped in-fold; antrum merging with swollen end of ductus bursae; ductus bursae long,
slender, medialiy spinose; corpus bursae globular, bearing a pair of minute denticulate areas.
REMARKS. This species most closely resembles V. suradeva, but the long triangular medial streaks
of the antemedial fascia on the fore wing are very distinctive. An unusual feature of V. plumosa
is the frequent great restriction of white dorsal scaling on the antenna. Very few specimens
examined had the normal antennal pattern. The genitalia are distinctive in both sexes. There is
some variation in this species, at least partially geographically correlated. Specimens from
Sumbawa and Lombok are larger than others and have the subbasal area of the fore wing
relatively restricted; those from Bali in the west and from the eastern part of the Lesser Sunda
chain are smaller and have the orange area larger. More material will be needed to determine
whether these apparent geographical correlations are consistent. A long series from Salajar
appears consistently different from specimens from the Lesser Sunda Islands. For the present
we recognize two subspecies, from the following geographical areas: (1) Bali; Lombok; Sumbawa;
Kalao; Timor and Damar; (2) Salajar.
BioLoGy. Early stages and host-plants unknown. Moths collected in January, April and from
September to December.
DISTRIBUTION (Fig. 22). Lesser Sunda Islands and Salajar.
Vitessa plumosa plumosa Hampson, 1896
(Figs 22, 31, 39, 124, 125, 168, 199, 216, 264, 310)
Vitessa plumosa Hampson, 1896), Trans. ent. Soc. Lond. 1896 : 503. Holotype 3, SUMBAWA: ix.1891
(Doherty) (genitalia slide no. 8256; BMNH) [examined].
Vitessa plumosa Hampson; Pagenstecher, 1898 : 200.
3, 2. 41-52 mm. Generally as described for species. Antenna completely black. Fore wing above: orange
subbasal area variable in width; white antemedial fascia sometimes divided at cell. Hind wing above:
black terminal band broad at apex, abruptly narrowing at Cu,, narrow to anal margin.
GENITALIA 6 (Figs 31, 39, 168, 199, 216). As described for species.
GENITALIA ° (Figs 264, 310). As described for species.
REMARKS. Topotypical specimens from Sumbawa and specimens from Lombok are larger than
the material from other islands, including the subspecies sa/ayerensis, with proportionally larger
genitalia and broader fore wings. Material from Bali and the islands east of Sumbawa do not
conform exactly with the Lombok-Sumbawa series; these specimens have slightly wider orange
subbasal areas, similar to those in the Salajar series, and the hind wing markings are somewhat
intermediate between the two subspecies. However, the material grouped under the nominate
subspecies differs consistently from the Salajar subspecies by the shape of the terminal band on
the hind wing, the narrowness of which, after passing Cu, is similar to that in V. teleroma of the
suradeva-group. It is likely that more material will show that additional subspecies should be
recognized in the lesser Sunda Islands.
BioLocy. Early stages and host-plant unknown. Moths collected in January, April, and from
September to December.
DISTRIBUTION (Fig. 22). Lesser Sunda Islands from Bali to Damar.
MATERIAL EXAMINED
Bali: 1 3, West Bali, Mondoktoempang, 750 m, x.1934 (Kalis) (BMNH). Lombok: 4 3, 1 2, Sapit, 600 m,
iv—vi.1896 (Fruhstorfer) (BMNH; RNH, Leiden). Sumbawa: | 3, ix.1891 (Doherty) (BMNH). Kalao:
1 2 [paralectotype of Vitessa teleroma Swinhoe] (BMNH). Timor: 1 3, Oinainisa (“Oinanissa’) [64-3 km
from Kupang, 600 m], xi-xii.1891 (Doherty) (BMNH); 1 9, Suai, i.1918 (Wahr) (BMNH). Damar: | 2°,
Dammer, xii.1898 (Kiiin) (BMNH).
REVISION OF VITESSIDIA & VITESSA Aly
Vitessa plumosa salayerensis subsp. n.
(Figs 22, 126, 127,169, 200, 265, 311)
3, 2. 38-44 mm. Generally as described for species. Antenna black, posterior surface of apical third of
shaft either with isolated white scales or with extensive white scaling. Fore wing above: orange subbasal
area slightly variable but moderately wide; white antemedial fascia with large medial indentation, but
never divided completely. Hind wing above: black terminal band broad at apex, gradually tapering to
anal margin.
GENITALIA ¢ (Figs 169, 200). As described for species.
GENITALIA 2 (Figs 265, 311). As described for species.
REMARKS. Specimens of this subspecies are consistently smaller than topotypical specimens of
the nominate subspecies, but about the same size as specimens from Bali, Kalao, Timor and
Damar. The fore wing is narrower, the orange subbasal area is generally wider. The hind wing
has the black terminal band more evenly curved, wider in the cubital area than in the nominate
subspecies, and with the medial margin either fairly straight or evenly curved. In the ¢ genitalia
the valve is slightly narrower, especially at the terminal margin.
BIoLoGy. Early stages and host-plant unknown. Moths collected in December.
DISTRIBUTION (Fig. 22). Salajar.
MATERIAL EXAMINED
Holotype 3, Salajar (‘Seleyer’): Somarisi, 500 m (‘1660 ft’), xii.1938 (Kalis) (BMNH).
Paratypes. 54 3, 73 2 (including allotype), data as holotype (genitalia slide no.’s 14652, 14653, 15843,
15844; BMNH).
The suradeva-group
D1aGnosis. Wing pattern variable, but basically as in the stettina-group; antemedial fascia not with long,
acuminate, distal streaks on R and Cu as in the plumosa-group. Hind wing with white basal and distal
area and contrasting blackish fuscous terminal band. ¢ genitalia with basal portion of uncus very wide,
distal portion rod-shaped, down-curved then up-turned, with long lateral setae; valve with distinctive
subquadrate outline, costa strongly bent subbasally, valve consequently opening in dorsal-ventral pre-
paration at right angles, not at an acute angle, to tegumen and uncus; termen of valve nearly truncate;
clasper a low flange, with small angular, ventrally directed, scoop-like process. The ? genitalia with
posterior sclerite of ostial chamber reduced and lightly sclerotized; anterior sclerite with antrum heavily
sclerotized, very large and pouch- or sack-shaped, more extensive than in other species-groups.
Vitessa teleroma Swinhoe, 1906
(Figs 23, 128-130, 170, 201, 217, 266, 312)
Vitessa teleroma Swinhoe, 1906, Ann. Mag. nat. Hist. (7) 18 : 412. LECTOTYPE 3, MAtaya: Perak,
1907-128 (genitalia slide no. 8255; BMNH), here designated [examined].
3, 9. 42-50 mm. Frons, vertex orange. Eye brown. Ocellus minute or absent. Chaetosema with pale
yellowish setae. Antenna black, anterior surface of scape basally orange; dorsal scaling of apical third of
shaft white, last several segments fuscous. Labial palpus: first two segments orange, third segment black,
inner surface and tip with light admixture of orange in °. Maxillary palpus basally black, distally orange.
Basal scaling of proboscis glossy blue-black. Neck region orange. Thorax above: orange, 8-shaped medial
blue-black area, posterior margin white; patagium orange with posteromedial glossy blue-black spot;
tegula with anterolateral glossy and terminal blue-black spot. Abdomen above: first segment white,
remaining segments black, posterior margins white, progressively narrowing to anal end, terminal segment
with posterior margin and anal tuft orange. Body below: black; pectus orange; abdomen as above from
second segment on. Legs black; fore coxa dorsally black admixed with orange, ventrally edged with white;
fore femur posteroventrally tipped with white; mid and hind femora ventrally white; mid and hind tibiae
with broad medial white band; tibiae posteriorly tipped with white; tibial spurs white, admixed with black
in 9. Fore wing above: ground-colour blue-black with dark green hue. Orange area basal, subcostal,
sometimes extending to posterior margin, base of posterior margin white; orange subcostal streak to base,
narrower streak to base on CuP, sometimes broken, demarcating large round blue-black spot from Sc to
CuP, narrower oblique blue-black spot in cubital area; fairly wide blue-black subcostal fascia; white
318 E. G. MUNROE & M. SHAFFER
antemedial fascia just short of costa and posterior margin, divided at cell, small anterior spot narrow and
triangular, posterior spot larger, rectangular; white postmedial fascia short of costa and posterior margin,
divided at Cu, in 3, anterior spot large, oval, over discocellular vein, traces of subcostal streak towards
apex, posterior spot smaller, more distal, triangular, directed along CuP; in 9 subcostal streak stronger,
smaller white streaks anterior and posterior to posterior spot; terminal area with all veins white, thicker
in °, extending from termen, short of posterior spots of postmedial fascia; fringe dark grey. Hind wing
above: narrow infuscation from costa, blue-black, violaceous terminal band, broad at apex, gradually
tapering to anal margin in 3, narrower in 2 and abruptly angled at Cu., finely tapering to anal margin;
iarge white medial area broadly extending to and including most of anal margin. Fore wing below: black,
violaceous; white subcostal streak at base; broad white streak at posterior margin; white postmedial fascia
as above, weaker, more diffuse; short, weak, white apical vein-lines, more extensive in °; retinaculum
white; fringe as above. Hind wing below: as above; costa edged with white; short, weak apical vein-lines,
more extensive in 9; fringe white.
GENITALIA ¢ (Figs 170, 201, 217). Uncus long, de-curved then up-turned; basal portion very wide,
flattened, laterally constricted, with long narrow foramen; distal portion rod-like with short sparse lateral
setae. Gnathos with wide lateral arms. converging to a long, compressed, acuminate, claw-shaped medial
process, not as long as uncus. Transtilla V-shaped, medially broadened, high, pronounced lateral flexure.
Tegumen: posterior sclerite broad with posterior margin straight; anterior sclerite with narrow medial
emargination. Vinculum broad, basally concave. Valve long, costa basally strongly arched, then straight
and parallel to posterior margin, narrowly inflated for most of length, up-turned to apex; apex sub-
truncate; termen broadly rounded; posterior margin sinuate; mesal surface with fine longitudinal and
oblique striations; clasper with small, oblique, raised basal ridge, separated from crease-like carina, not
meeting free edge of distal part of clasper, a free edge rising abruptly, recurving to form round raised
scoop-like process, becoming tubular and distally tapering, running parallel to posterior margin, ter-
minating at two-thirds length. Juxta with large, narrow lateral sclerites; ventral process very long, flat,
narrow, up-turned; dorsal process broader, shorter, up-turned near centre and bearing setae, directly
connected to long curved sclerite on anellus, longitudinally ridged and bearing spines; between spiny
sclerites a large, narrow, oval, sclerotized catena, longitudinally creased, medially and distally spined
(Fig. 201); manica supported by sclerotized bands. Aedeagus long, distally expanded, base narrowly
rounded; ductus ejaculatorius subbasal; vesica with spines spreading ventrally to distal end of aedeagus,
here membrane bulbous and transversely rugose; vesica of even width, abruptly narrowing after cornuti
clusters; vesica dorsally spinose and longitudinally rugose; junction with sclerotized aedeagus forming
elbow-joint when vesica everted and reflexed; two ventrolateral cornuti clusters (Fig. 217), variable in
length, each with long spines, left cluster two to four times as long as right, with about 40-80 spines, right
cluster with about 20-30 spines.
GENITALIA © (Figs 266, 312). High, narrow, posteriorly prolonged ovipositor lobes, dorsally fusing, surface
densely setose, longer setae on margin, spiny setae visible but not conspicuous at 25 magnification.
Eighth tergite with anteromedial margin long, broadly rounded. Posterior apophysis short, thin, nearly
straight; anterior apophysis longer, broader, curved, tip spatulate. Posterior ostial sclerite (Fig. 312)
weakly sclerotized, weakly constricted laterally and ventrally; posterior margin shallowly V-shaped;
short lateral prolongations; short medial development on anterior margin. Anterior sclerite with weakly
sclerotized ventral margin, narrow, finely wrinkled, laterally setose. Antrum very large, heavily sclerotized,
pouch-shaped, slightly waisted posteriorly; dorsal part posteriorly broadly rounded, with margin ventrally
recurved at junction with anterior margin of posterior sclerite; anteriorly with strong dorsal reflexion;
ventral part with irregular transverse and oblique creases, especially on raised margins of aperture of
ductus seminalis about mid-way, these creases and those at junction with sclerotized raised base of ductus
bursae making antrum asymmetrical. Ductus bursae minutely spinose along entire length. Corpus bursae
globular, with very minute paired denticulate areas.
REMARKS. This species is closely related to V. suradeva. It differs externally by the restricted pale
markings and the complete black subbasal fascia of the fore wing, and by the appreciable posterior
tapering of the terminal band of the hind wing. The $ genitalia are very similar to those of V.
suradeva, but differ in the larger cornuti clusters and the narrower longitudinal element of the
clasper. The 2 genitalia differ in the generally larger, broader antrum and in the reflexion of its
anterior margin. This species bears a superficial resemblance to some members of the stfettina-
and pyraliata-groups; the 2, with extra white markings of the fore wing postmedial fascia, closely
resembles V. muluana. However, V. teleroma is distinguishable from all these by the genital
REVISION OF VITESSIDIA & VITESSA 319
characters of the species groups (see diagnoses of stettina-group, p. 286, pyraliata-group, p. 303,
suradeva-group, p. 317).
BIoLoGy. Early stages and host-plant unknown. Moths collected in March, May, June, September
and October.
DISTRIBUTION (Fig. 23). Malaya; Sumatra; Natuna Island; Borneo. Dubious records from Sri
Lanka, India: Darjiling.
MATERIAL EXAMINED
Paralectotypes. Malaya: | 3, Perak, 6844 (genitalia slide no. 15845; BMNH). Borneo: 1 3, 2 °, no data,
Moore coll. (genitalia slide no.’s 8263, 15850; BMNH). [1 9°, Kalao Is., see under Vitessa plumosa plumosa
Hampson.]
Malaya: | 3, Pulo Pinang (‘Penang Island’), ex Boisduval coll. (BMNH); 1 3, 1 2°, Penang, 1896 (Curtis)
(BMNH); 5 3, Penang (BMNH; CMAG, Birmingham, UM, Oxford; MM, Manchester); 3 3, Penang,
Government Hill, 300 m, v.1898 (Curtis) (BMNH); | 3, Penang Hill, ix.1932 (Corbet) (BMNH); 3 3,
Fraser Hill, x.1930 (Corbet) (BMNH); 5 3, Bukit Kutu, 1000 m (Sanderson) (BMNH); | ¢, Kutu Hill,
1050 m, iii.1931 (Corbet) (BMNH); 4 3, Wellesley Province (Distant) (TM, Pretoria). Sumatra: 2 3, SW.
Sumatra, N. Korintji Valley, 1500 m, ix—x.1921 (Pratt) (TM, Pretoria); 1 3d, no exact data, Adams
Bequest (BMNH). Natuna Island: 1 3, Bunguran, vii-x.1894 (Hose) (BMNH). Borneo: | 2, Kalimantan,
Island of Pontianak, Sintang, xii.1909 (Martin) (BMNH); 1 2, K., Samarinda, ix.1938 (Walsh) (BMNH);
2 2, no exact data (BMNH; CAS, San Francisco); | 3, Sarawak, Matang, 1100 m, vi.1900 (Shelford)
(BMNH); 44, S., Gunong Mulu Nat. Park, Mulu, 1790 m, lower montane (moss) forest, MV — canopy.
1.1978 (Holloway et al.) (BMNH); 1 3, S., Gunong Mulu Nat. Park, G. Api, 1500 m, scrub, pandanus,
Act. iv.1978 (Holloway et al.) (BMNH). No data: 1 3, 1 2, ex Felder coll. (BMNH); 1 3, ex Joicey coll.
(BMNH); 1 ¢ (UZM, Copenhagen). Dubious data: | 2, Ceylon (Niet.) ex Felder coll. (BMNH); 1 &,
Darjeeling (BMNH).
Vitessa suradeva Moore, [1860]
(Figs 24, 32, 40, 131-134, 171, 172, 202, 203, 218, 267, 268, 313, 314)
Vitessa suradeva Moore, [1860] : 299.
3, 2. 30-54 mm. Frons, vertex orange. Eye, ocellus brown. Chaetosema with pale yellowish setae. Antenna
black, anterior surface of scape basally orange; dorsal scaling of apical third of shaft white, extending to
nearly half length of shaft in 9, last several segments fuscous. Labial palpus: first two segments orange,
third segment black, sometimes minutely tipped with white. Maxillary palpus basally black, distally
orange. Basal scaling of proboscis blue-black, ventrally admixed with orange. Neck region orange. Thorax
above: large medial black oval spot, anteriorly and laterally orange, posteriorly white; patagium orange
with posteromedial glossy blue-black spot; tegula orange, anterolateral glossy blue-black spot, tip black.
Abdomen above: black, posterior margins of segments white, widest on second segment, progressively
narrowing, terminal segment with posterior margin and anal tuft orange. Body below blue-black; pectus
orange; abdomen as above from second segment on, white margins broader. Legs black ; fore coxa dorsally
orange, tipped with black, ventrally white; mid and hind coxae banded with white; fore femur ventrally
orange, mid and hind femora ventrally white; mid and hind tibiae with broad medial white band; tibiae
posteriorly tipped with white; tibial spurs white, admixed with black in 9; all first tarsal segments bearing
posterior dorsal white spot. Fore wing above: ground-colour blue-black, dark green hue along costa.
Basal orange area narrowly subcostal, extending to posterior margin, white at base; orange subcostal
and CuP streaks to base, demarcating anterior round basal black dot, posterior oblique basal black streak,
the latter reaching posterior margin; orange medial margin with CuP streak joining white antemedial
fascia, anterior and posterior margins basally arcuate, narrowly short of white fascia at costa, becoming
white and extending along posterior margin to antemedial and postmedial fasciae; glossy blue-black
subbasal fascia demarcated and divided into two quadrate spots by orange streak, anterior spot to costa,
posterior spot short of posterior margin. Greyish white antemedial fascia narrowly short of costa, basal
margin slightly arcuate anterior and posterior to CuP; medial margin broadly indented at cell, a triangular
streak on either side, short of or touching white postmedial fascia; greyish white postmedial fascia divided
into two major spots at Cu, or diffusely joined at this point in 9, anterior spot large, oval, over dis-
cocellular vein, short of costa, posterior spot irregular, more distal in 2, sometimes divided at Cu, and A,
in 3, large central portion medially extended on CuP and sometimes touching antemedial fascia. Terminal
area with greyish white on all veins, always thin or greatly reduced on CuP, short of or touching termen,
320 E. G. MUNROE & M. SHAFFER
sometimes joining posterior spot of postmedial fascia on Cu, and Cu,; fringe light grey, sometimes
admixed with white at apex. Hind wing above: costal infuscation narrow, black terminal band violaceous,
broad at apex, tapering very weakly and gradually towards anal margin; large medial white area broadly
extending to anal margin; fringe basally light grey, distally white. Fore wing below: black, slightly
violaceous, particularly anterior to cell, brownish fuscous in cubital region, broadly white along posterior
margin; short subcostal white streak at base; retinaculum whitish grey; white postmedial fascia a long
narrow oval in 4, divided at Cu, in 2; short, greyish white apical and terminal vein-lines to Cu, in 2, only
short faint apical lines in 3; fringe light grey in 3, basally white in 9. Hind wing below as above; costa
broadly edged with white; short faint apical and terminal white vein-lines to Cu.; fringe white.
GENITALIA ¢ (Figs 32, 40, 171, 172, 202, 203, 218). Generally as described for V. teleroma. Base of uncus
very wide, flattened and laterally constricted. Valve with costa incurved for most of length, apex sharply
subtruncate, posteroventral angle broadly rounded; basal portion of clasper joining raised rim of scoop-
shaped process, distal portion tubular and tapering, running parallel to posterior margin; catena (Fig. 40)
on anellus (Figs 202, 203) constricted at base and distally; cornuti clusters (Fig. 218) variable in length.
GENITALIA ° (Figs 267, 268, 313, 314). Generally as described for V. teleroma. Antrum (Figs 313, 314) very
large, heavily sclerotized, pouch- or sack-shaped, slightly waisted medially or tapering anteriorly, roughly
square or noticeably longer than wide; anteriorly with slight dorsal reflexion; ventral wall with strong
oblique creases giving rise to sclerotized base of ductus seminalis at mid-length, these creases producing
slight asymmetry of antrum.
REMARKS. This species is distinguished by the strong division of the blue-black subbasal fascia
of the fore wing by an orange streak on CuP joining the orange basal area to the white antemedial
fascia; and by the presence of two distinctly demarcated blue-black basal spots. The white
fasciae of the fore wing are basically similar to those of other groups where these markings are
extensive and the fasciae more or less compiete. The basal and subbasal dark markings most
closely resemble those in V. hieratica, but here the division and demarcation of the spots is less
complete; the white fasciae most closely resemble those in V. plumosa, but in that species the
medial streaking of the antemedial fascia is more accentuated than in V. suradeva. The genitalia
closely resemble those of V. teleroma; the 3 genitalia are distinguished by the generally wider
base of the uncus, the incurved costa, the general shape of valve, the slightly wider ridge of the
distal portion of the clasper and its complete joining to the base of the ridge; in the 2 genitalia
by the distinctly longer antrum and weaker dorsal reflexion, and by the origin of the ductus
seminalis mid-way on the antrum. We recognize two subspecies, from the following geographical
areas: (1) north India, W. Bengal and Assam; Bangladesh; Burma; Thailand and Vietnam;
(2) southern India and Sri Lanka. These have been considered distinct species by some authors,
synonyms by others.
BioLocy. Early stages and host-plant information given under V. suradeva rama, below. Moths
collected by various collectors throughout the year.
DISTRIBUTION (Fig. 24). Sri Lanka; India; Bangladesh; Burma; Thailand; Vietnam.
Vitessa suradeva suradeva Moore, [1860]
(Figs 24, 40; 131,132, 171,/202, 267, 313)
Vitessa suradeva Moore, [1860], in Horsfield & Moore, Cat. lepid. Insects Mus. nat. Hist. East-India House
2 : 299, pl. 7a, fig. 7; Lederer, 1863 : 335, pl. 6, fig. 6; Walker, 1864 : 219; Moore, 1886 : 58; Swinhoe
& Coates, 1889 : 664; Swinhoe, 1890 : 292; Snellen, 1890 : 560; Ragonot, 1891 : 110; Swinhoe,
1900 : 429. LECTOTYPE 32, INpDIA: Assam, Chera Poonjee (‘Cherrapunji’) (Buckley) (Type no. 2637;
genitalia slide no. Oxford 718-1977, Shaffer; UM, Oxford), here designated [examined].
Vitessa formosa Felder & Rogenhofer, 1874, Reise dst. Fregatte Novara, Zool. Theil, 2: pl. 137, fig. 1.
Holotype 9, INp1A (‘Ind. or.’) (genitalia slide no. 16021; BMNH) [examined]. [Synonymized by
Hampson, 1896a : 146.]
Vitessa suradeva Moore; Hampson, 1896a : 146; 18966 : 503, partim.
[Vitessa suradeva Moore; Kenrick, 1907 : 75. Misidentification: see Vitessa hemiallactis hemiallactis
Meyrick, above.]
REVISION OF VITESSIDIA & VITESSA 321
[Cosmethis suradeva (Moore); Janse, 1928 : 88. Misidentification: see Vitessa hollandi hollandi subsp. n.,
above. ]
Cosmethis suradeva (Moore); de Joannis, 1930 : 637.
3, 9. 40-54 mm. As described for species.
GENITALIA 6 (Figs 40, 171, 202). Generally as described for V. teleroma and V. suradeva. Base of uncus
very wide above lateral constriction. Scoop-shaped process on clasper very large, usually exceeding
posterior margin. Left cornuti cluster about two and a half times length of right.
GENITALIA ° (Figs 267, 313). Generally as described for V. teleroma and V. suradeva. Antrum (Fig. 313)
very large, roughly square, pouch-shaped, slightly waisted medially; posterior margin of dorsal wall
medially convex, recurved, covering opening to antrum; deep, oblique, longitudinal and transverse folds
on ventral wall giving rise to raised sclerotized base of ductus seminalis at about mid-length; lower third
of antrum with slight dorsal reflexion; posteriorly produced, narrowed and convex beyond broad-
shouldered entry of ductus bursae; reflexion and creases causing asymmetry of antrum.
REMARKS (see also Remarks under species). Apart from being generally larger than V. suradeva
rama and having a somewhat more finely tapering terminal band on the hind wing, this subspecies
is indistinguishable externally. The genitalia, however, are distinct in both sexes. The ¢ genitalia
have the base of the uncus slightly broader, the costal margin of the valve more strongly curved,
the scoop-shaped process of the clasper larger, and the cornuti clusters proportionally larger.
In the @ genitalia the antrum is more distinctly pouch-shaped, wider but shorter, with a more
noticeable dorsal reflexion, and with generally even width.
BioLocy. Early stages and host-plant unknown. Moths collected from February to May, August
to October and December.
DISTRIBUTION (Fig. 24). Northern India; Bangladesh; Burma; Thailand; Vietnam.
MATERIAL EXAMINED
Paralectotypes. India: 1 3 (Argent) (Type no. 2637; UM, Oxford). [There were originally two more syn-
types, one collected by Buckley, one by Argent. These have not been traced; they may be among the
material listed below, but we have not been able to establish this.]
India: 1 3, Malnold [?] [with label citing original reference for V. suradeva], ex Felder coll. (BMNH);
1 2, no exact data, ex Felder coll. (BMNH); 2 3, 1 2, no exact data, ex Swinhoe coll. (TM, Pretoria);
1 2, no exact data (TM, Pretoria); 1 2, no exact data (E/wes) (RNH, Leiden); 1 2, Darjeeling, Gopaldhara,
1400 m (Stevens) (BMNH); 3 9, Darjiling, vili.1912 (Carmichael) (UM, Oxford); 3 9, Darjiling
(Carmichael) (RSM, Edinburgh); 1 3, 27 9, Darjiling (Méller) (BMNH); 1 3, Darjeeling, v.1935
(Sevastopulo) (BMNH); 3 3, 5 2, Darjiling (BMNH); 1 3, 1 2, Khasi Hills, Moore coll. (BMNH); 14 3,
4 9, Khasis, ix-x.1894 (CMAG, Birmingham; MNHN, Paris; UM, Oxford); 3 2, Khasi Hilll[s], ii, v,
vi.1973 (HI, Iruma City); 21 3, 19 9, Assam, Khasia Hills [1 3, 1x.1894] (BMNH; NMNH, Washington;
TM, Pretoria; UM, Oxford); 1 3, Khasis [sic] Hills (Hamilton) (BMNH); 2 2, Assam, Cherrapunji,
v.1893 (C. F. H.) (BMNH); 3 3, Assam (BMNH; NMNH, Washington); 1 3, Assam, Jaintia Hills (MM,
Manchester); 1 ¢, Cachar (UM, Oxford); 1 ¢, Naga H[ills] (BMNH). Bangladesh: 2 3, 2 2, Sylhet (UM,
Oxford); 3 3, 1 2, Silhet (BMNH). Burma: 1 2, no exact data (TM, Pretoria); 4 2, East Pegu, 1200-1500 m,
iii-iv.1890 (Doherty) (BMNH); 1 2, Mergui, [xii, (Anderson),] ex Moore coll. (BMNH); 5 3, Mergui
(Doherty) (TM, Pretoria). Thailand: 1 2, Khao Sabap, Chentaban, 1935 (Macbeth) (BMNH); | 3, Pyayen
Range, Pakchong, xii.1957 (Munroe) (CNC, Ottawa). Vietnam: 2 3, Tonkin, Cha pa, 11.1930 (Lemai)
(MNHN, Paris); 1 3, Tonkin, Son La (Lemai) (MNHN, Paris); 1 2, Tonkin (MNHN, Paris); 1 3, 2 2,
Hanoi [(Lerminat)] (MNHN, Paris); 1 2, Cochin China [?], Laokai, vi.1936 (Masseyeff) (BMNH);
1 3, Haut Donai, E. Poilane (NMNH, Washington). No data: 6 3, 2 2, (BMNH; NMNH, Washington;
UM, Oxford).
Vitessa suradeva rama Moore, 1885, stat. n.
(Figs 24, 32, 133, 134, 172, 203, 218, 268, 314)
Vitessa rama Moore, 1885, Lepid. Ceylon 3 : 256, pl. 178, fig. 4. LECTOTYPE 3, Sri LANKA (‘Ceylon’):
Moore coll. 94-106 (BMNH), here designated [examined].
3, 2. 30-50 mm. As described for species.
522 E. G. MUNROE & M. SHAFFER
GENITALIA 6 (Figs 32, 172, 203, 218). Generally as described for V. teleroma and V. suradeva. Left cornuti
cluster (Fig. 218) over three times length of right. See also Remarks, below.
GENITALIA & (Figs 268, 314). Generally as described for V. teleroma and V. suradeva, Antrum (Fig. 314)
very large, sack-shaped, posteriorly wide, gradually tapering anteriorly, distinctly longer than wide.
Posterior margin of dorsal wall medially with broadly conical recurved flap, covering opening to antrum;
about lower third of antrum with very slight dorsal curve, not causing reflexion; anteriorly narrowly
rounded, tip level with opening of ductus bursae; deep, oblique longitudinal and transverse folds on
ventral wall giving rise to raised sclerotized base of ductus seminalis, opening posterior to mid-length of
antrum; creases causing only slight asymmetry of antrum.
REMARKS. This has often been treated as a distinct species, but the differences from V. suradeva
suradeva are small, and we consider them as of subspecific value only. This subspecies is on the
average smaller than the nominate subspecies, but the measurements given include a series
bred in southern India by T. R. Bell; this series contains most of the smallest specimens. The pale
markings of the fore wing are very slightly reduced and the black terminal band of the hind wing
is generally broader in V. suradeva rama than in V. suradeva suradeva, otherwise specimens of
comparable size are externally indistinguishable. The 3 genitalia of the present subspecies have
the basal part of the uncus slightly narrower, the valve slightly narrower, with a smaller scoop-
shaped process on the clasper and, as the most striking feature, the right cornuti cluster much
reduced. The 9 genitalia are distinct by the shape and size of the antrum, which in the present
subspecies is larger, narrower and generally shows little sign of dorsal reflexion.
BIoLoGy. The description of immature stages is adapted from the manuscript notes of T. R. Bell
preserved in the British Museum (Natural History). Cast larval skins and pupal cases, included
with material from south India in the Bell collection, have been examined for comparison with
Bell’s descriptions. The terminology used here for larval characters is mainly based on Peterson
(1962), that for pupal characters on Kuznetsov (1967); the setal nomenclature of the larva and
any ambiguous terms follow Bell’s own system.
LarvA. Length 35-40 mm; width, middle of body 4:5 mm, head 3-5 mm. Oily-looking; general coloration
glossy black with white and orange markings. Head-capsule light orange, broadly semi-oval, surface
glossy and smooth. Tubercles wider than setae they bear, circular, not prominent; setae fine; subdorsal
setae near apex of front nearly as long as height of head, those of adfrontals shortest, not as long as width
of front. Front equilaterally triangular, apex widely arched, each side slightly incurved, reaching more than
one-third height of head. Adfrontals half height of head, apex more broadly arched, the whole not clearly
defined. Clypeus transverse, transparent or nearly so, posterior margin strongly curved, anterior margin
quite straight. Labrum glossy, blackish and kidney-shaped, as long as but narrower than clypeus. Basal
segment of antenna orange, second and third segments glossy black. Mandible black, cutting edge entire
or nearly so. Six ocelli arranged in a semicircle, of equal size, three, four and six black, remainder yellow
with black centres. Legs long, glossy, black, claw rose-tinged. Prolegs thin, held well under abdomen, with
infuscated glassy-colourless cylindrical shanks and light coloured planta bearing crochets in complete
circle. Strongly interrupted subspiracular white line running whole length of thoracic and abdominal
segments; below, an interrupted line of similar spots along the line of legs. White spots on mesothorax
and first abdominal segment larger than rest and nearly coalescent to form a short subdorsal line. A
supraspiracular, interrupted, white, longitudinal line or narrow band from first to eighth abdominal
segments, formed of three spots and lines on each segment. Seventh abdominal segment longer than
eighth; eighth segment contracted posteriorly by deeply impressed line, widely convex anteriorly from
spiracle to spiracle, dividing off nearly one-third of segment which is twice as long as ninth, appearing
somewhat tumid medially; ninth segment with posterior margin shelving shortly to tenth. Ninth segment
black; tenth segment, including anal claspers, orange; claspers short, erect, bases thickly conical, ends
broadly rounded, planta light orange, two-lobed, with crochets arranged in straight row. Ventral surface
of body black. Spiracles broadly oval, reddish brown, with coarse, raised, glossy, black rims; those of
prothorax and eighth abdominal segment fully twice as wide and long as any other; those from first to
third segments slightly larger than spiracles from fourth to seventh, which are about one-sixteenth as long
as segment. Setae white, arising from black tubercles, erect, fine, as long as width of body. Setae of all
subspiracular tubercles, especially on ninth and tenth abdominal segments, as long as a segment, sub-
dorsals and dorsolaterals a little over half as long. Setal arrangement of fifth abdominal segment: dorsal
REVISION OF VITESSIDIA & VITESSA 323
trapezoidal arrangement of setae one-third length of segment; subdorsals the same distance apart; dorso-
laterals very little more; supraspiracular a spiracle length in front of and above spiracle; two sub-
spiraculars close together and with no postspiracular setae, a little less than three spiracle lengths below
spiracle, one being directly below spiracle, the other a spiracle length in front of first and slightly higher up;
of five others, one behind and below the subspiraculars and four on the leg-base.
Pupa. Length 16mm, width 6mm. Glossy; dorsally bright orange-chestnut, ventrally lighter. Stout,
widest at centre, anteriorly conical, head anteriorly square; abdomen weakly tapering posteriorly, sharply
tapering from seventh segment, anal end narrowly rounded. Surfaces of head and thoracic areas minutely
corrugated; surfaces of first to eighth abdominal segments minutely pitted, posterior margins of segments,
especially of seven and eight, broadly smooth and without pits; segments nine and ten without pits. Dorsal
surface: areas well demarcated except between mesonotum and fore wings, metanotum and hind wings;
hind wings narrowly visible to past centre of third tergite; fore wings visible to near centre of fourth. Line
of fore wing straight on narrow convergence to anterior end of metanotum; anterior margin of metanotum
broadly concave. Abdominal tergites two and three noticeably wider than remainder. Spiracles from
tergites two to eight with raised dark rims, those of tergites two, three and eight noticeably larger than
remainder. Ventral surface; tips of fore wings medially reaching fifth sternite; antennae converging with
mid and hind legs, with tips raised from surface, reaching sternite six. Cremaster composed of eight
hooks, two central hooks stouter and larger.
Hapsits. Larvae communal when young, up to six together in a clean, clear, laxly woven web
spun on undersides of young leaves; separating when fully grown, each making a similar web for
itself amongst older leaves towards the tops of the plants; preferring to feed on young leaves
throughout their existence. Often found sitting highly looped like a semilooper of the noctuid
type. Larvae active and moving about the web with alacrity but rarely leaving it. When ready to
pupate, larvae descending to ground and making a cocoon of paper-like consistency with light
fawn-coloured silk, shining smooth inside and more fluffy outside. Cocoon made against the
side of a stone just under the surface of the earth or in a similar place; nearly circular or broadly
oval and slightly pointed at the ends; attached by threads of silk all round. Outer threads made
first, the larva then retiring into the middle and making the cocoon proper. The latter very roomy,
measuring 23mm x 10mm x 5mm. Pupa attached inside by cremaster; imago emerging
through a slit at other end. Moth flying by day, seen at Castle Rock at 1500 ft [450 m] in
September, feeding from flowers of a large tree called Kadam tree (Rubiaceae), relatively few
V. suradeva in the company of large numbers of Macrobrochis gigas (Walker) (Hypsidae). Pupa
described about 20.viii and the first moth emergence on the night of 13.ix.
HOST PLANT OF LARVA. This is a shrub that in Kanara only grows in the evergreen jungles on the
Western Ghats but is otherwise widely distributed throughout India and Sri Lanka to Sumatra.
It is a small tree, practically a shrub, that grows in the underwood, often forming nearly the whole
underwood in parts. The plant is Chailletia sumatrana Miquel [= Dichapetalum gelonioides
gelonioides (Beddome) Engler (Dichapetalaceae)].
GENERAL. Moths collected by various collectors in most months of the year.
DISTRIBUTION (Fig. 24). Sri Lanka; southern India.
MATERIAL EXAMINED
Paralectotypes. Sri Lanka: 4 3, 1 9°, data as lectotype (genitalia slide no.’s 1501 [abdomen only, genitalia
missing], 1502, 15855; BMNH).
Sri Lanka: 5 3, 12 2, no exact data (BMNH; CAS, San Francisco; MM, Manchester); 2 2, Kandy,
1886 (BMNH); 2 3, 2 2, Kandy, ix.1902-xii.1909 (Mackwood) (BMNH); 1 2, Kandy, ix.1914 (V. H.)
(BMNH); 1 3, Dambulla, vii.1903 (Newman) (BMNH); 1 2, Maskeliya, vii.1885 (BMNH); 2 3, 7 2,
Maskeliya, i-ix (Mackwood) (BMNH); 2 2, Pundaluoya, xi.1903 (BMNH); 1 3, 1 9, Ellahara, vii.1903
(Newman) (BMNH); 3 3, Nalanda, vii.1903 (Newman) (BMNH); | 9, no data (Thwaites) (UM, Oxford);
1 2, no data (UM, Oxford). India: 1 2, Mui, v.1912 (Maxwell) (BMNH); 1 3g, Tranvancore (Place)
(BMNH); 1 2, Annamali (‘Anaimalai’) Hills (Godman) (BMNH); | 3, Karwar, rains (Davidson) (BMNH);
8 3, 15 2°, Karwar, ix.1923-iv.1924 (Bell) (BMNH); 1 3, Kanara (Bell) (BMNH); 10 3, 6 2, Castle Rock,
v-vi.1920 (Bell) (BMNH); 2 3, 1 2, no data (Mackwood) (BMNH); 2 3, no data(BMNH; MNHN, Paris).
Various cocoons, pupal cases and last instar larval skins, from localities in southern India (Bell)
(BMNH).
324 E. G. MUNROE & M. SHAFFER
References
Belkin, J. N. 1962. The mosquitoes of the South Pacific. 2 vols, xii+608 pp., 412 pls. Berkeley and Los
Angeles.
Cowan, C. F. 1975. Horsfield, Moore, and the Catalogues of the East India Company Museum. J. Soc.
Biblphy nat. Hist. 7 : 273-284, figs 1, 2.
Felder, C., Felder, R. & Rogenhofer, A. F. 1864-1875. Reise der Gsterreichischen Fregatte Novara um die
Erde in den Jahren 1857, 1858, 1859. Zoologischer Theil. 2. 2. Abtheilung: Lepidoptera 2. 9+ 10+ 20
pp., 140 pls. Wien.
Guillaumin, M. 1976. Participation des Euphaedra et Bebearia (Lep. Nymphalidae) aux associations
mimétiques de la forét équatoriale africaine. I. - Associations ayant des Hétérocéres 4 moeurs diurnes
comme modeéles. Bull. Soc. zool. Fr. 101 : 603-611, pls 1-4.
Hampson, G. F. 1896a. The fauna of British India, including Ceylon and Burma. Moths 4. xxvi+ 594 pp.,
287 figs. London.
1896. On the classification of three subfamilies of moths of the family Pyralidae: the Epipaschiinae,
Endotrichinae, and Pyralinae. Trans. ent. Soc. Lond. 1896 : 451-550, text-figs.
1898. A revision of the moths of the subfamily Pyraustinae and family Pyralidae. Proc. zool. Soc.
Lond. 1898 : 590-761, pls 49, 50.
1906. On new Thyrididae and Pyralidae. Ann. Mag. nat. Hist. (7) 17 : 189-222.
Holland, W. J. 1900. The Lepidoptera of Buru. Part 2. Heterocera. Novit. zool. 7 : 555-591.
Hiibner, J. [1820]. Verzeichniss bekannter Schmettlinge [sic]. Signature 12. 431 pp. Augsburg.
Janse, A. J. T. 1924. List of species, including descriptions of new species, belonging to the family
Pyralidae, collected by Messrs. C., F. and J. Pratt, in the mountains of central Ceram during October,
1919, to February, 1920. Bull. Hill Mus. Witley 1 : 489-507.
—— 1928. List of Pyralidae collected by Messrs. C., F. and J. Pratt, in Dutch New Guinea, during 1920
and 1921. Bull. Hill Mus. Witley 2 : 75-88, pl. 1, figs 1-7.
Joannis, J. de 1930. Lépidoptéres Hétérocéres du Tonkin (3° partie). Annis Soc. ent. Fr. 98 : 559-834.
Kenrick, G. H. 1907. A list of moths of the family Pyralidae collected by A. E. Pratt in British New Guinea
in 1902-3, with descriptions of new species. Proc. zool. Soc. Lond. 1907 : 68-87, pl. 3, figs 6-213,
pl. 4, figs 5-209.
Kirby, W. F. 1892. A synonymic catalogue of Lepidoptera Heterocera (Moths). 1. Sphinges and Bombyces.
xii+ 951 pp. London.
Kuznetsov, N. Y. 1967. Fauna of Russia and adjacent countries. Lepidoptera. 1. iv+ 305 pp., 212 figs, 7
maps. (English translation, Jerusalem.)
Lederer, J. 1863. Beitrag zur Kenntniss der Pyralidinen. Wien. ent. Monatschr. 7 : 243-280, pls 2-5,
331-502, pls 6-18.
Meyrick, E. 1886. Descriptions of Lepidoptera from the South Pacific. Trans. ent. Soc. Lond. 1886 : 189-
296.
—— 1887. On Pyralidina from Australia and the South Pacific. Trans. ent. Soc. Lond. 1887 : 185-268.
Moore, F. 1858 [1860]. /n Horsfield, T. & Moore, F., A catalogue of the lepidopterous insects in the Museum
of Natural History at the East-India House 2 : 279-440+ 1-6, pls 13-23, 7A-13A. London. (For dates
see Cowan, 1975).
—— 1885. The Lepidoptera of Ceylon. 3 : xv+ 578 pp., pls 144-215. London.
—— 1886. List of the Lepidoptera of Mergui and its Archipelago collected for the Trustees of the Indian
Museum, Calcutta, by Dr. John Anderson, Superintendent of the Museum. J. Linn. Soc. (Zool.)
21 : 29-60, pls 3, 4.
Munroe, E. 1963. Collecting insects in the Territory of Papua and New Guinea, 1957. Proc. Pacif. Sci.
Congr. 9 : 32-41, figs 1-12.
1977. Synopsis of Heortia Lederer, with descriptions of a new species and a new subspecies (Lepi-
doptera : Pyralidae : Odontiinae) Can. Ent. 109 : 429-441, figs 1-20.
Pagenstecher, A. 1884. Beitrige zur Lepidopteren-Fauna von Amboina. Jb. nassau. Ver. Naturk, 37 : 150-
326, pls 6, 7.
—— 1886. Beitrage zur Lepidopteren-Fauna des Malayischen Archipels. (3) Heteroceren der Aru-Inseln,
Kei-Inseln, und von Siidwest-Neu-Guinea. Jb. nassau. Ver. Naturk, 39 : 104-194.
— 1888. Beitrage zur Lepidopteren-Fauna des Malayischen Archipels. (5) Verzeichniss der Schmetter-
linge von Amboina. Jb. nassau. Ver. Naturk, 41 : 85-217.
— 1898. Beitrage zur Lepidopteren-Fauna des Malayischen Archipels. (12) Jb. nassau. Ver. Naturk.
51 : 179-200.
—— 1900. Die Lepidopterenfauna des Bismarck-Archipels. Zweiter Theil: Die Nachtfalter. 268 pp.,
2 pls. Stuttgart.
REVISION OF VITESSIDIA & VITESSA 325
Peterson, A. 1962. Larvae of insects. An introduction to Nearctic species. Part 1. Lepidoptera and plant
infesting Hymenoptera. 315 pp., figs O1-14, L1—-58, H1—12. Ohio.
Ragonot, E. L. 1891. Essai sur la classification des Pyralites. Annis Soc. ent. Fr. 60 : 15-114.
1891. Essai sur la classification des Pyralites. Note supplémentaire et rectificative (1). Annls Soc.
ent. F. 60 : 599-646.
Rothschild, Lord 1915. On Lepidoptera from the islands of Ceram (Seram), Buru, Bali, and Misol. Novit.
zool. 22 : 209-227.
Rothschild, W. & Jordan, K. 1905. On some new Lepidoptera discovered by A. S. Meek in British New
Guinea. Novit. zool. 12 : 448-478.
Schultze, W. 1908. New and little-known Lepidoptera of the Philippine Islands. Philipp. J. Sci. 3 (1)
A : 27-37, pl. 1, figs 1-13.
Snellen, P. C. T., with notes by Elwes, H. J. 1890. A catalogue of the Pyralidina of Sikkim collected by
Henry J. Elwes and the late Otto MGller. Trans. ent. Soc. Lond. 1890 : 557-647, pls 19, 20.
Stoll, C. [1781]. Zn Cramer, P., Papillons exotiques des trois parties du monde, Il’ Asie, Il’ Afrique
et l Amerique. 4, 29+ 252 pp., pls 289-400. Amsterdam & Utrecht.
Swinhoe, C. 1890. The moths of Burma. Part 2. Trans. ent. Soc. Lond. 1890 : 201-296, pls 7, 8.
—— 1892. Catalogue of eastern and Australian Lepidoptera Heterocera in the collection of the Oxford
University Museum. Part 1. Sphinges and Bombyces. viii+ 324 pp., 8 pls. Oxford.
—— 1900. Catalogue of eastern and Australian Lepidoptera Heterocera in the collection of the Oxford
University Museum. Part 2. Noctuina, Geometrina and Pyralidina. vii+ 540 pp., 8 pls. Oxford.
—— 1906. New and little-known species of Eastern and Australian Heterocera. Ann. Mag. nat. Hist.
(7) 18 : 403-416.
Swinhoe, C. & Cotes, E. C. 1889. A catalogue of the moths of India. Part 5. Pyrales. 591-670 pp. Calcutta.
Vane-Wright, R. I. 1971. The systematics of Drusillopsis Oberthiir (Satyrinae) and the supposed
Amathusiid Bigaena van Eecke (Lepidoptera : Nymphalidae), with some observations on Batesian
mimicry. Trans. R. ent. Soc. Lond. 123 : 97-123, figs 1-47.
Walker, F. 1864. List of the specimens of lepidopterous insects in the collection of the British Museum.
31. Supplement. 321 pp. London.
—— 1869. Characters of undescribed Lepidoptera Heterocera. 112 pp. London.
Whalley, P. E. S. 1961. A change of status and a redefinition of the subfamily Endotrichinae (Lep.
Pyralidae) with the description of a new genus. Ann. Mag. nat. Hist. (13) 3 : 733-736, fig.
—— 1963. A revision of the world species of the genus Endotricha Zeller (Lepidoptera : Pyralidae).
Bull. Br. Mus. nat. Hist. (Ent.) 13 : 397-453, pls 1-37, figs 1-314.
Wilson, E. O. 1959. Adaptive shift and dispersal in a tropical ant fauna. Evolution, Lancaster, Pa.
13 : 122-144.
Figs 45-51 (45) Vitessidia diaphana Rothschild & Jordan, g, New Guinea. (46) V. diaphana
Rothschild & Jordan, °, New Guinea. (47) V. geometrina sp.n., 2 holotype, New Guinea. (48)
Cosmethella unipectinalis (Hampson), 3, Solomon Islands. (49) C. unipectinalis (Hampson), °,
Solomon Islands. (50) C. minor sp. n., 2 holotype, New Guinea. (51) C. major sp. n., 2 holotype,
New Guinea.
5/7
Figs 52-57 (52) Vitessa griseata Kenrick, 3, New Guinea. (53) V. griseata Kenrick, 2, New Guinea.
(54) V. brandti sp.n., ¢ holotype, New Britain. (55) V. ralboti (Janse), ¢ lectotype, Moluccas.
(56) V. talboti (Janse), 2 paralectotype, Moluccas. (57) V. nobilis sp. n., 2 holotype, Moluccas.
64
Figs 58-65 (58) Vitessa cyanea ceramensis subsp. n., 3 allotype, Moluccas. (59) V. cyanea ceramensis
subsp. n., 2 holotype, Moluccas. (60) V. cyanea cyanea Hampson, 3, D’Entrecasteaux Islands.
(61) V. cyanea cyanea Hampson, °, D’Entrecasteaux Islands. (62) V. zemire expansa subsp. n., 3,
Moluccas. (63) V. zemire expansa subsp. n., 2 paratype, Moluccas. (64) V. zemire zemire (Stoll),
3, Moluccas. (65) V. zemire zemire (Stoll), 2, Moluccas.
Figs 66-73 (66) Vitessa zemire temerata Swinhoe, 3, New Guinea. (67) V. zemire temerata Swinhoe,
2, New Guinea. (68) V. zemire nephritica subsp. n., 3 paratype, Australia. (69) V. zemire nephritica
subsp. n., 2 paratype, Australia. (70) V. zemire coronis subsp. n., ¢ holotype, Umboi Island. (71) V.
zemire coronis subsp. n., 2 allotype, Umboi Island. (72) V. zemire novaebritanniae subsp. n., 3
holotype, New Britain. (73) V. zemire novaebritanniae subsp. n., 2 allotype, New Britain.
80
Figs 74-81 (74) Vitessa zemire solomonis subsp. n., 3 holotype, Solomon Islands. (75) V. zemire
solomonis subsp. n., 2° paratype, Solomon Islands. (76) V. zemire cheesmanae subsp. n., ¢ paratype,
New Hebrides. (77) V. zemire cheesmanae subsp. n., 2° paratype, New Hebrides. (78) V. vitialis
Hampson, 3, Fiji. (79) V. vitialis Hampson, 3, Fiji (Lau group). (80) VY. vitialis Hampson, 2, Fiji.
(81) V. tamsisp.n., 2 paratype, Western Samoa.
Figs 82-90 (82) Vitessa hieratica Swinhoe, 3, Borneo: Sarawak. (83) V. hieratica Swinhoe, °,
Sumatra. (84) V. philippina sp. n., 2 holotype, Philippines. (85) V. stetrina Swinhoe, 3, New Guinea.
(86) V. stettina Swinhoe, 2°, New Guinea. (87) V. hollandi wallacealis subsp. n., 3 holotype, Sulawesi.
(88) V. hollandi wallacealis subsp. n., 2 allotype, Sulawesi. (89) V. hollandi hollandi subsp. n., 3
holotype, New Guinea. (90) V. hollandi hollandi subsp. n., 2 paratype, New Guinea.
Figs 91-99
(91) Vitessa ternatica Lederer, 3, New Guinea. (92) V. ternatica Lederer, 2 (paralectotype
of sarumensis Holland), Moluccas. (93) V. sp., d, Japan [?]. (94) V. hemiallactis moluccana subsp.
n., ¢ holotype, Moluccas. (95) V. hemiallactis moluccana subsp. n., 2 allotype, Moluccas. (96)
V. hemiallactis hemiallactis Meyrick, 3, New Guinea. (97) V. hemiallactis hemiallactis Meyrick,
2, New Guinea. (98) V. hemiallactis admiralitatis subsp. n., ¢ paratype, Admiralty Islands. (99)
V. hemiallactis admiralitatis subsp. n., 2 paratype, Admiralty Islands.
103 "|
Figs 100-107 (100) Vitessa hemiallactis lustrans subsp. n., 3, New Ireland. (101) V. hemiallactis
lustrans subsp. n., 2 allotype, New Hannover. (102) V. intermedia sp. n., 3 holotype, New Guinea.
(103) V. intermedia sp. n., 2 allotype, New Guinea. (104) V. barrettisp. n., ¢ holotype, New Guinea.
(105) V. barretti sp. n., 2 allotype, New Guinea. (106) V. glaucoptera Hampson, 3, Australia. (107)
V. glaucoptera Hampson, °, Australia.
115
Figs 108-116 (108) Vitessa cristobalensis sp. n., 2 holotype, Solomon Islands. (109) V. nicobarica
Hampson, ¢ paralectotype, Nicobar Islands. (110) V. nicobarica Hampson, 2 paralectotype,
Nicobar Islands. (111) V. pyraliata latialbata subsp. n., 3 holotype, Malaya. (112) V. pyraliata
latialbata subsp. n., 2 paratype, Malaya. (113) V. pyraliata triangulifera subsp. n., 3 holotype,
Philippines. (114) V. pyraliata triangulifera subsp. n., 2 paratype, Philippines. (115) V. pyraliata
pyraliata Walker, 3, Sulawesi. (116) V. pyraliata pyraliata Walker, °, Sulawesi.
: 124 125
Figs 117-125 (117) Vitessa kolakalis sp.n., 3 holotype, Sulawesi. (118) V. kolakalis sp.n., 2
allotype, Sulawesi. (119) V. sulaensis sp. n., 2 holotype, Moluccas. (120) V. muluana sp. n., ° holo-
type, Borneo: Sarawak. (121) V. gemina sp.n., 2 holotype, Schouten Islands. (122) V. gemina
sp. n., 2 paratype, Schouten Islands. (123) V. splendida Schultze, 2, Philippines. (124) V. plumosa
plumosa Hampson, 3, Lesser Sunda Islands. (125) V. plumosa plumosa Hampson, 2, Lesser Sunda
Islands.
ae
or
: a
| 133 } 134
Figs 126-134 (126) Vitessa plumosa salayerensis subsp.n., ¢ holotype, Salajar Island. (127) V.
plumosa salayerensis subsp.n., Q allotype, Salajar Island. (128) V. teleroma Swinhoe, 3 para-
lectotype, Borneo: Kalimantan. (129) V. teleroma Swinhoe, 3, Malaya. (130) V. teleroma Swinhoe,
2, Borneo: Kalimantan. (131) V. suradeva suradeva Moore, 3, North India. (132) V. suradeva
suradeva Moore, 9, North India. (133) V. suradeva rama Moore. &. Sri Lanka. (134) V. suradeva
rama Moore, 2, South India.
Figs 135-141 ¢ genitalia. (135) Vitessidia diaphana Rothschild & Jordan. (136) Cosmethella
unipectinalis (Hampson). (137) Vitessa brandti sp.n. (not to scale). (138) V. griseata Kenrick.
(139) V. talboti (Janse). (140) V. cyanea ceramensis subsp. n. (141) V. cyanea cyanea Hampson.
Figs 142-149 ¢ genitalia. (142) Viressa zemire zemire (Stoll). (143) V. zemire temerata Swinhoe.
(144) V. zemire nephritica subsp. n. (145) V. zemire coronis subsp. n. (146) V. zemire novaebritanniae
subsp. n. (147) V. zemire solomonis subsp. n. (148) V. zemire cheesmanae subsp. n. (149) V. vitialis
Hampson.
aneltus
G- quel \ 2
Figs 150-157 ¢ genitalia. (150) Vitessa hieratica Swinhoe. (151) V. stettina Swinhoe. (152) V.
hollandi wallacealis subsp. n. (153) V. hollandi hollandi subsp. n. (154) V. ternatica Lederer. (155) V.
sp. Japan [ ?] (156) V. hemiallactis moluccana subsp. n. (157) V. hemiallactis hemiallactis Meyrick.
——anellus
158
Figs 158-165 <¢ genitalia. (158) Vitessa hemiallactis admiralitatis subsp. n. (159) V. hemiallactis
lustrans subsp. n. (160) V. intermedia sp. n. (161) V. barretti sp. n. (162) V. glaucoptera Hampson.
(163) V. nicobarica Hampson. (164) V. pyraliata latialbata subsp. n. (165) V. pyraliata triangulifera
subsp. n.
Figs 166-173 ¢ genitalia. (166) V. pyraliata pyraliata Walker. (167) V. kolakalis sp. n. (168) V.
plumosa plumosa Hampson. (169) V. plumosa salayerensis subsp. n. (170) V. teleroma Swinhoe.
(171) V. suradeva suradeva Moore. (172) V. suradeva rama Moore. (173) Detail of anellus and valve
region of V. griseata Kenrick.
Figs 174-182 ¢ genitalia. Details of anellus and valve regions. (174) Vitessa talboti (Janse). (175) V.
cyanea ceramensis subsp. n. (176) V. cyanea cyanea Hampson. (177) V. zemire temerata Swinhoe.
(178) V. zemire nephritica subsp. n. (179) V. zemire coronis subsp. n. (180) V. zemire novaebritanniae
subsp. n. (181) V. zemire solomonis subsp. n. (182) V. zemire cheesmanae subsp. n.
Figs 183-191 ¢ genitalia. Details of anellus and valve regions. (183) Vitessa vitialis Hampson.
(184) V. hieratica Swinhoe. (185) V. stettina Swinhoe. (186) V. hollandi wallacealis subsp. n. (187) V.
hollandi hollandi subsp. n. (188) V. ternatica Lederer. (189) V. sp. Japan [?] (190) V. hemiallactis
moluccana subsp. n. (191) V. hemiallactis hemiallactis Meyrick.
Figs 192-200 ¢ genitalia. Details of anellus and valve regions. (192) Vitessa hemiallactis admiralitatis
subsp. n. (193) V. hemiallactis lustrans subsp. n. (194) V. intermedia sp. n. (195) V. glaucoptera
Hampson. (196) V. pyraliata latialbata subsp. n. (197) V. pyraliata pyraliata Walker. (198) V.
kolakalis sp. n. (199) V. plumosa plumosa Hampson. (200) V. plumosa salayerensis subsp. n.
Figs 201-209 ¢ genitalia. Figs 201-203. Details of anellus and valve regions. (201) Vitessa teleroma
Swinhoe. (202) V. suradeva suradeva Moore. (203) V. suradeva rama Moore. Figs 204-209. Details
of cornuti. (204) V. griseata Kenrick. (205) V. cyanea ceramensis subsp. n. (206) V. cyanea
cyanea Hampson. (207) V. zemire coronis subsp. n. (208) V. zemire cheesmanae subsp. n. (209) V.
vitialis Hampson.
é, ‘ Pw ike : \
Be vasa Giriate et Ait Srey >
oe : or ¥
og | $ :
Pies Sosa : : ,
f
Hampson. (215) V. pyraliata pyraliata Walker. (216) V. plumosa plumosa Hampson. (217) V.
teleroma Swinhoe. (218) V. suradeva rama Moore.
Figs 219-228 9° genitalia. Figs 219, 220. Details of corpus bursae. (219) Vitessidia diaphana
Rothschild & Jordan. (220) V. geometrina sp. n. (221) V. diaphana Rothschild & Jordan. (222) V.
geometrina sp. n. (223) Cosmethella unipectinalis (Hampson). (224) C. minor sp. n. (225) C. major
sp. n. (226) Vitessa griseata Kenrick. (227) V. talboti (Janse). (228) V. nobilis sp. n.
Figs 229-238 ° genitalia. (229) Vitessa cyanea ceramensis subsp. n. (230) V. cyanea cyanea Hampson.
(231) V. zemire expansa subsp. n. (232) V. zemire zemire (Stoll). (233) V. zemire temerata Swinhoe.
(234) V. zemire nephritica subsp. n. (235) V. zemire coronis subsp. n. (236) V. zemire novaebritanniae
subsp. n. (237) V. zemire solomonis subsp. n. (238) V. zemire cheesmanae subsp. n.
Figs 239-248 2 genitalia. (239) Vitessa vitialis Hampson. (240) V. tamsi sp. n. (241) V. hieratica
Swinhoe. (242) V. philippina sp. n. (243) V. stettina Swinhoe. (244) V. hollandi wallacealis subsp. n.
(245) V. hollandi hollandi subsp. n. (246) V. ternatica Lederer. (247) V. hemiallactis moluccana
subsp. n. (248) V. hemiallactis hemiallactis Meyrick.
Figs 249-258 ©° genitalia. (249) Vitessa hemiallactis admiralitatis subsp. n. (250) V. hemiallactis
lustrans subsp. n. (251) V. intermedia sp. n. (252) V. barretti sp. n. (253) V. glaucoptera Hampson.
(254) V. cristobalensis sp. n. (255) V. nicobarica Hampson. (256) V. pyraliata latialbata subsp. n.
(257) V. pyraliata triangulifera subsp. n. (258) V. pyraliata pyraliata Walker.
Figs 259-268 2 genitalia. (259) Vitessa kolakalis sp. n. (260) V. sulaensis sp. n. (261) V. muluana sp.
n. (262) V. gemina sp. n. (263) V. splendida Schultze. (264) V. plumosa plumosa Hampson. (265) V.
plumosa salayerensis subsp. n. (266) V. teleroma Swinhoe. (267) V. suradeva suradeva Moore. (268)
V.suradeva rama Moore.
» “a ey 975 3 i | 276
Figs 269-276 & genitalia. Details of ostial chamber. (269) Cosmethella unipectinalis (Hampson).
(270) C. minor sp. n. (271) C. major sp. n. (272) Vitessa griseata Kenrick. (273) V. talboti (Janse).
(274) V. nobilis sp. n. (275) V. cyanea ceramensis subsp. n. (276) V. cyanea cyanea Hampson.
| om
4 . 7
'
if
Figs 277-284 2 genitalia. Details of ostial chamber. (277) Vitessa zemire expansa subsp. n. (278) V.
zemire zemire (Stoll). (279) V. zemire temerata Swinhoe. (280) V. zemire nephritica subsp. n.
(281) V. zemire coronis subsp. n. (282) V. zemire novaebritanniae subsp. n. (283) V. zemire solomonis
subsp. n. (284) V. zemire cheesmanae subsp. n.
Figs 285-292 © genitalia. Details of ostial chamber. (285) Vitessa vitialis Hampson. (286) V. tamsi
sp. n. (287) V. hieratica Swinhoe. (288) V. philippina sp. n. (289) V. stettina Swinhoe. (290) V.
hollandi wallacealis subsp. n. (291) V. hollandi hollandi subsp. n. (292) V. ternatica Lederer.
Figs 293-300 2 genitalia. Details of ostial chamber. (293) Vitessa hemiallactis moluccana subsp. n.
(294) V. hemiallactis hemiallactis Meyrick. (295) V. hemiallactis admiralitatis subsp. n. (296) V.
hemiallactis lustrans subsp. n. (297) V. intermedia sp. n. (298) V. barretti sp. n. (299) V. glaucoptera
Hampson. (300) V. cristobalensis sp. n.
Figs 301-306 & genitalia. Details of ostial chamber. (301) Vitessa nicobarica Hampson. (302) V.
pyraliata latialbata subsp. n. (303) V. pyraliata triangulifera subsp. n. (304) V. pyraliata pyraliata
Walker. (305) V. kolakalis sp. n. (306) V. sulaensis sp. n.
Figs 307-314 9 genitalia. Details of ostial chamber. (307) Vitessa muluana sp. n. (308) V. gemina
sp. n. (309) V. splendida Schultze. (310) V. plumosa plumosa Hampson. (311) V. plumosa salayerensis
subsp. n. (312) V. teleroma Swinhoe. (313) V. suradeva suradeva Moore. (314) V. suradeva rama
Moore.
Figs 315-318 2 genitalia. Details of setae on ovipositor lobes. (315) Cosmethella minor sp. n.
(316) Vitessa griseata Kenrick. (317) V. zemire zemire (Stoll). (318) V. muluana sp. n.
REVISION OF VITESSIDIA & VITESSA 359
Index
Synonyms and unavailable names are in /talics; principal references are in bold.
admiralitatis subsp. n. 245, 266, 295, 297, 298
Agaristidae 242, 243
Argyrolepidia sp. 242, 244
barbara Stoll 264
barretti sp. n. 245, 262, 263, 266, 268, 300, 301,
302
brandti sp. n. 244, 262, 264, 268, 269
celebensis Munroe 243
ceramensis subsp. n. 244, 264, 273
cheesmanae subsp. n. 245, 265, 280
coronis subsp. n. 245, 265, 278, 279
Cosmethella gen. n. 242, 244, 245, 252, 254, 255,
256, 261
Cosmethis auctorum 244, 258, 264, 267, 269, 276,
290, 321
cristobalensis sp. n. 245, 263, 266, 302
cyanea Hampson 243, 244, 257, 262, 264, 271, 273
diaphana Rothschild & Jordan 244, 245, 253, 254
dominalis Lederer 243, 244
Endotrichini 241, 242
expansa subsp. n. 245, 264, 275, 276, 277
formosa Felder & Rogenhofer 245, 320
gemina sp. n. 245, 263, 266, 289, 291, 303, 312
Geometridae 264
geometrina sp. n. 244, 253
gigas Walker 243, 323
glaucoptera Hampson 245, 262, 263, 266, 300,
301, 302
griseata-group 243, 244, 258, 261, 262, 267, 271,
272, 301
griseata Kenrick 244, 262, 263, 264, 267, 268, 269,
210; 271, 301, 302
hemiallactis-group 245, 261, 262, 263, 267, 286,
287, 288, 289, 291, 300, 301, 303, 314
hemiallactis Meyrick 262, 263, 265, 266, 289, 291,
293, 294, 295, 296, 299, 300, 301, 302, 303, 320
Heortia sp. 243, 244
hieratica-group 245, 258, 261, 262, 263, 267, 284,
286
hieratica Swinhoe 245, 265, 285, 320
hillalis (Schaus) 243
hollandi sp. n. 245, 262, 263, 265, 289, 290, 291,
295, 296, 310, 313, 321
Homoptera 242
Hypsidae 243, 323
Imma sp. 243
Immidae 243
intermedia sp. n. 245, 263, 266, 288, 299, 300, 301,
302
iridia Munroe 243
kolakalis sp. n. 245, 263, 266, 303, 306, 309, 310
latialbata subsp. n. 245, 263, 266, 306, 308, 309
lustrans subsp. n. 245, 266, 295, 298, 300, 303
Macrobrochis sp. 243, 323
major sp. n. 244, 255, 256, 257
megisto (Boisduval) 242, 244
minor sp. n. 244, 255, 256, 257
moluccana subsp. n. 245, 265, 293, 295, 296
muluana sp. n. 245, 263, 266, 303, 311, 318
nephritica subsp. n. 245, 262, 263, 264, 277, 278,
279, 302
Nevrina sp. 243
nicobarica Hampson 245, 263, 266, 303, 306, 312
nobilis sp. n. 244, 262, 264, 270
novaebritanniae subsp. n. 245, 265, 279
Odontiinae 243, 244
Phalaena sp. 264, 274, 276
philippina sp. n. 245, 262, 263, 265, 286, 287, 306,
308, 314
plumosa-group 245, 262, 263, 265, 286, 291, 303,
S12. 314,317
plumosa Hampson 245, 266, 267, 314, 316, 319,
320
polyplagalis Hampson 243
procopia (Stoll) 243
pyraliata-group 245, 262, 263, 286, 289, 291, 292,
295, 303, 306, 310, 312, 313, 314, 318, 319
pyraliata Walker 244, 245, 262, 263, 266, 287,
296, 303, 305, 306, 308, 309, 310, 311, 314
Pyralidae 241, 242, 243
pyraliformis Walker 244
Pyralinae 241, 242, 293
Pyralini 242
Pyraustinae 243, 244
radiata Kenrick 243
rama Moore 242, 243, 245, 262, 267, 320, 321, 322
salayerensis subsp. n. 245, 267, 316, 317
sarumensis Holland 245, 292, 293
saturata (Walker) 243
solomonis subsp. n. 245, 265, 280, 281
splendida Schultze 245, 263, 266, 303, 313, 314
stettina-group 245, 261, 262, 263, 286, 288, 291,
295, 296, 303, 306, 308, 310, 313, 314, 317, 319
360 E. G. MUNROE & M. SHAFFER
stettina Swinhoe 245, 262, 263, 265, 287, 288, 289,
290, 292, 296, 299, 300, 301, 302, 318
sulaensis sp. n. 245, 263, 266, 303, 310
suradeva-group 245, 261, 263, 265, 286, 291, 303,
3073122914, 510, 387,019
suradeva Moore 242, 243, 245, 258, 262, 263, 267,
286, 290, 291, 295, 296, 314, 316, 318, 319, 320,
EV aK 7 an Pe
talboti (Janse) 244, 262, 264, 269, 270, 271
tamsi-sp: n;245, 262, 265, 271, 275,282, 283
teleroma Swinhoe 245, 263, 267, 307, 312, 316,
317-3 18320, 321322
Tellervo sp. 242
temerata Swinhoe 245, 264, 276, 278
ternatica Lederer 245, 263, 265, 287, 292, 293, 294,
295, 298
triangulifera subsp. n. 245, 262, 263, 266, 287,
306, 308, 309, 314
triplaga Hampson 244
Tyspanodes sp. 243
unipectinalis (Hampson) 242, 244, 254, 256, 257
Vitessa Moore 242, 243, 244, 245, 252, 254, 255,
256, 257, 258, 264
Vitessidia Rothschild & Jordan 242, 244, 245,
25282595255
vitessoides (Moore) 243
vitialis Hampson 245, 262, 265, 271, 275, 281, 282,
283
wallacealis subsp. n. 245, 289, 290, 291, 310
zalmira misspelling 276
zemira misspelling 276
zemire-group 243, 244, 258, 261, 262, 267, 271
zemire (Stoll) 245, 262, 263, 264, 265, 271, 272,
214,215, 210, 211, 2105 219, 200; 201, 202
British Museum (Natural History)
The Generic Names of Moths of the World
The aim of this series is to provide a complete list of all the generic
names, their type-species, designations, and appropriate references,
for each of the families of moths treated. By ensuring that each
generic name has its type-species fixed in accordance with the
International Code of Zoological Nomenclature, this series should
rectify present confusion and form a basis for a more stable
nomenclature. All generic names have been checked for
homonymy, and if necessary replaced.
The series is a companion to The Generic Names of the Butterflies
and their Type-species by F. Hemming, published in 1967
(reprinted 1972). It is expected to run into some seven volumes
which will not necessarily appear in numerical order.
The authors are all specialists on the staff of the Department of
Entomology, British Museum (Natural History) and have the
unrivalled collections and resources of the Museum on which to
base their work.
Volume 1; Noctuoidea (part): Noctuidae, Agaristidae and Nolidae.
by I. W. B. Nye
1975 pp. viii+568 frontispiece Crown Quarto Cloth.
ISBN 0 565 00770 X £26.00
Volume 2; Noctuoidea (part): Arctiidae, Ctenuchidae, Dioptidae, Lymantriidae,
Notodontidae, Thaumetopoeidae and Thyretidae.
by A. Watson, D. S. Fletcher and I. W. B. Nye.
Publishing 1980
Volume 3; Geometroidea: Apoprogonidae, Axiidae, Callidulidae, Cyclidiidae,
Drepanidae, Epicopeiidae, Epiplemidae, Geometridae, Pterothysanidae,
Sematuridae, Thyatiridae and Uraniidae.
by D. S. Fletcher. ;
1979 pp xx+244 frontispiece A4, Cloth Bound £22.50
ISBN 0 565 00812 9
A STANDING ORDER placed with us now will ensure that you
are kept informed of forthcoming volumes of this important work.
As each volume is published you will receive a pro forma invoice
informing you of the cost. Your copy will be forwarded on receipt
of your payment.
The companion volume
The Generic Names of the Butterflies and their Type-species, by
F. Hemming
Bulletin of the B.M.(N.H.) Entomology Supplement No. 22.
1967 (reprinted 1972) SO9pp 4to paper covers £22.00
Titles to be published in Volume 39
A revision of the African pod bugs of the tribe Clavigrallini
(Hemiptera: Coreidae) with a checklist of the world species.
By W. R. Dolling.
A revisionary classification of the Cynipoidea (Hymenoptera) of the
Ethiopian Zoogeographical Region. By J. Quinlan.
A review of the genera of the Hydroptilidae (Trichoptera).
By J. E. Marshall.
A revision of Vitessidia Rothschild & Jordan and Vitessa Moore
(Lepidoptera: Pyralidae). By E. G. Munroe & M. Shaffer.
Printed by Henry Ling Ltd, Dorchester
Se ote
Spa’
:
fr
3
Sreita
be
~ =
at wat ~ 3
aS
ee
>
3
peter?
Se
"he
an
=
mesa
Seas
oye
ee
aes
a
viene
AHS
tes
‘<
ans
ee
Eye
eee
(y i
y ‘
(a
es
a
thy
SMEG Tt
roe iio
ine ti { vie
i
Fast
Ast
aia ‘
—
<i
jo
8
fh 3
rite:
He i hah
asia
a
2
Settee
Se
ye
ite,
pate
ithe *
st asatity | 4
atte
~%
alt yes
Hi ¢ at
A
Sea ers :
a 3. oes
rae
anh
—
Ay
Sar
mae
peace ate ate Dy
eer
t
5, ty
x!
ff
sbntanrath
open
¢ MR
9
eral
a
PP Rita:
(peciets
iter Rides
Retz insttate Aide
ote a ‘ au 9
ey gh anette!
= 4 : hi
Vs c Se
i
oo
.
nine
ST eb Utne a ate et ah! ath
b Fate Gy tests espteti at APP ry
Mtn nen Dts An ne, t
Perk
dhe i
wt
~~
“S
or,
eae
erase
=
SS
aye
aie
et re on
Pa
neeeey
i See
— Sas
reste
aren
oe
pnuaratioes
2) ete Re Ee
is ite ke
cite f
* i
<
rats Ki
5, steed,
=
note vie =
Peg ee ats
EY oa? or
Sr sig ein bees ten
ee be eee
rete —
xc
se
a
ai
o
Fre
S “3
ae.
rs
: see ‘4
! i
: aut
‘Sh
poe
ee
BN,
aby
cde Bea sat
Crt rth o Tua,
cena
pee sees Mente eh pe
nissan
ie’ ty
Sad ok
Ay
aria
%
Seg: ie
, ne
t,
ae
4
Samer ena oa eoahy
+" oe
Se
a,
pea
ee as
paraka =
Letts
~
Sete
:
a.
Lye
ean
%
‘ ~
paoeeret
is
rs eat
eee
er ator eee
dah eet
ea Aet ey ave sect
HTH >
stot
ih
oe
ie
aut
pea wef
Pro trod owed
Paetetatas
he
SEN
4h
<pteea!
-
Nae
fetgty
THe
a)
ea Ule tele teat :
rei hebetse eAlentay arta tstnns persed a}
: Ayal, yi
stile tala
:
- j ‘
Pat terete eet
SWE AAO ae ae
NOUN cies tote td stad
: i
Tae
Fate Pt)
Aiea ASE dTAL eA ie
psiud
HAGE
ae
ve >
bs
$,
. 3 i
seesheete eset e taht isthe ses i
TR Aeron
Uileenmcaisrieiet
Aust Fetes
Hane nee
iy
Sue
pose cecee
=
<3
Let
.
J Reg
zs
shakes
mre
Rotsnt
Sie
v
wax
=
pcs
’
: Stok r
site vit eet ata
ee
Raed pd dtd ips
ite yiaty the
i 2 Watt
ty
Beige
|
yeiete
ete masee
miatarete
ts
— aart
a Sipser:
eas
. Bet yehra fgets
+ =o ae, > =
Stat hs ye ema
2 . atta
a is tee
GS ares
eo teehe
Fey
a
ie
ae
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11