Bulletin of the British Museum (Natural History) Entomology

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

© British Museum (Natural History), 1989

ISBN 0 565 06037 6 Entomology Series
ISSN 0524-6431 Vol 58, No. 1, pp. 1 — 130

British Museum (Natural History)
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London SW7 5BD Issued 25 May 1989

Typeset by Computer Typesetting Services, Redhill, Surrey
Printed in Great Britain by Henry Ling Ltd, Dorchester, Dorset

Bull. Br. Mus. nat. Hist. (Ent.) 58(1): 1-78

Issued 25 May 1989

The mealybug genus Planococcus (Homoptera:

Pseudococcidae)

JENNIFER M. COX

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

CONTENTS

Introduction

Distinction of Planococcus from similar genera TORE. Pee Ae 2
3

New combinations

Checklist of species here included in Planococcus

Species-groups in Planococcus
Terminology
Depositories

Systematics = . .;
Planococcus Ferris :
Key to species of Planococcus
Descriptions of species

References

Index

SYNOPSIS.

Descriptions and illustrations are provided for the 35 species considered here

to belong in Planococcus, and a key given for their separation. Nine species are newly
described and eight new synonymies are proposed. Dactylopius calceolariae var. minor
Maskell from Mauritius, previously regarded as a synonym of Planococcus citri (Risso), is
resurrected as Planococcus minor, and P. pacificus Cox synonymized with it. Five species,
previously in Planococcus, are transferred to Planococcoides, and one species to Crisicoc-
cus, as new combinations. Three species of Crisicoccus, C. azaleae (Tinsley), C. mat-
sumotoi (Shiraiwa) and C. matesovae Danzig, are illustrated as they might be confused

with similar species of Planococcus.

INTRODUCTION

Scale insects are amongst the most important
pests of agricultural crops, debilitating the plant
by loss of sap, contaminating the plant and its fruit
with honeydew on which sooty mould frequently
grows, transmitting plant viruses, and sometimes
injecting toxins that stunt plant growth. Having
lifecycles in a warm climate of as little as one
month, mealybugs can rapidly attain very high
numbers on their hostplant. Fortunately they are
usually attacked by a wide range of natural
enemies, in particular, encyrtid parasitic wasps

and ladybird beetles. Most serious outbreaks of
mealybugs occur when they are transported to
new countries where their natural enemies do not
occur, but in these cases biological control pro-
grammes have generally been very effective.
Delays in attaining this control, however, have
sometimes been caused by initial uncertainty as to
the identity, and hence origin, of the mealybug
species concerned, as described under the
remarks for Planococcus kenyae (Le Pelley)
below.

The genus Planococcus contains a number of
well-known mealybug pests including P. citri
(Risso) on citrus, cocoa and a wide variety of

z

plants in greenhouses, P. ficus (Signoret) on gra-
pevines, figs and pomegranates, P. kenyae (Le
Pelley) on coffee, and P. lilacinus (Cockerell) and
P. minor (Maskell) on cocoa. P. vovae (Nasonov)
occurs on Cupressus grown as shelter belts in the
Mediterranean Basin and might be manipulated
as a beneficial insect to encourage the presence of
mealybug predators in the vicinity of crops. A
number of other species occur on crop plants but
are not at present regarded as pests, although this
could change should they be introduced to other
parts of the world where their natural enemies do
not occur. Several of the species revised below are
known largely from material intercepted on pro-
duce entering the U.K. or U.S.A. These species
have considerable potential as future pests but, as
their presumed geographical origins are listed
here, they would be suitable candidates for bio-
logical control.

Planococcus citri is a vector of swollen shoot
virus of cacao. However, it is likely that many of
the virus-transmission experiments carried out on
‘citri’ actually involved minor, as these two species
were not distinguished until recently.

Some species of Planococcus show hostplant
specificity at the plant family or genus level. Spe-
cies of the dendrobii-group are apparently specific
to Orchidaceae, the vovae-group to
Cupressaceae, and P. dubius Cox to Drac-
ophyllum (Epacridaceae). Others, despite having
a wide host range, show distinct preferences. For
example, P. citri, generally regarded as being
polyphagous (which it is in greenhouses), is very
rarely found out-of-doors on grapevines, while P.
ficus, common on grapevines, is never found on
citrus.

The identification of Planococcus species is hin-
dered by the morphological variation that occurs
as a result of the conditions under which the indi-
viduals develop. Cox (1983) showed P. citricus
Ezzat & McConnell (1956) to be a high tempera-
ture form of P. citri. Structural characteristics may
vary simply with environmental conditions, or
there may be an ‘optimum’ effect with, for
instance, specimens reared at high and low tem-
peratures having few oral collar tubular ducts and
specimens reared at medium temperatures having
more. The result is that the characteristics dis-
tinguishing species may be different for different-
sized individuals.

In the absence of live material it has been neces-
sary to resort to multivariate analysis in an
attempt to resolve some of the complexes dis-
cussed in the present paper.

Study of the current situation has been further
confused by the shifting of populations by man -
populations that may have been on the verge of

JENNIFER M. COX

speciation have been brought back together
again. Thus, although Planococcus probably has
an Old World origin, six species are now known to
occur in the Neotropical Region: P. citri, P. ficus,
P. halli Ezzat & McConnell, P. lilacinus, P. minor
and P. vovae. All of these species were probably
introduced into the Neotropical Region by man.

So much work has been done on this group that
the time has come to bring together the available
knowledge, and to attempt to resolve some of the
remaining species complexes. This study has
revealed that the the genus is even more complex
than was previously appreciated, and it is appar-
ent that a great deal more experimental work will
be necessary to completely elucidate some of the
complexes.

Distinction of Planococcus from similar
genera

Planococcus is not satisfactorily distinguished
from other genera. The evolution of mealybugs
has apparently involved the loss rather than gain
of characters in the adult females, making phy-
logenetic analysis based on females intractable.
Studies of males would probably lead to a better
understanding of relationships, but associated
males are not available for most species.

One of the consequences of this arbitrary dis-
tinction of mealybug genera is that some species of
Planococcus may be more closely related (by
descent) to species currently placed in other gen-
era than they are to other species of Planococcus.
This is particularly the case with Crisicoccus which
is distinguished from Planococcus only by the loss
of some of the cerarii - an occurrence that is likely
to have happened more than once. In this paper,
such members of Crisicoccus that are very similar
to species of Planococcus have been illustrated.

Cox & Ben-Dov (1986) synonymized Allococ-
cus with Planococcus and transferred a group of
African species previously in Allococcus to a new
genus, Delottococcus. Cox & Freeston (1985)
gave characters for distinguishing between the
closely related Planococcus and Planococcoides
and transferred Pseudococcus celtis Strickland
and Planococcus lamobokensis Matile-Ferrero to
the latter genus. In addition, as mentioned above,
Crisicoccus is very similar to Planococcus. These
four genera may be distinguished by the following
Key.

1 Cerarii numbering less than 18 pairs
CRISICOCCUS Ferris

— Cerarii numbering 18 pairs, although some thoracic
pairs may be indistinct ...................005. 2

MEALYBUG GENUS PLANOCOCCUS

2 Circulus absent, translucent pores absent from hind
COKAe. Yawn: DELOTTOCOCCUS Cox & Ben-Dov

— Circulus usually present, translucent pores usually
Present on hind! coxaewe sah. dawn 5

3 All abdominal cerarii anterior to anal lobe pair with
only 2 conical setae and without slender auxiliary
setae, anal lobe cerarii each with only 2 conical setae

PLANOCOCCUS Ferris

— Some abominal cerarii anterior to anal lobe pair with
either more than 2 conical setae or with slender
auxiliary setae, anal lobe cerarii each with 2 or more
cerarli .. PLANOCOCCOIDES Ezzat & McConnell

New combinations

The following five species hitherto placed in
Planococcus are here transferred to Plano-
coccoides:

Planococcoides crassus (De Lotto, 1961: 219)

(Planococcus) comb. n.

Planococcoides formosus (De Lotto, 1961: 221)

(Planococcus) comb. n.

Planococcides rotundatus (De Lotto, 1954: 110)

(Planococcus) comb. n.

These three African species are very similar to
each other and to P. celtis (Strickland) (trans-
ferred to Planococcides by Cox & Freeston, 1985)
in having a rotund body, multilocular disc pores
confined to mid-regions of the abdomen, flagell-
ate dorsal setae and large simple pores. They dif-
fer from the type species of Planococcoides, P.
njalensis, by lacking dorsal ducts. Another
African species, P. lamabokensis (Balachowsky &
Ferrero), (also transferred to Planococcoides by
Cox & Freeston, 1985), does have dorsal tubular
ducts and may be a synonym of P. njalensis.
Planococcoides anaboranae (Mamet, 1959: 403)

(Planococcus) comb. n.

This species is similar to Planococcoides lin-
dingeri (Bodenheimer) from the Mediterranean
Basin, which was transferred to Planococcoides
by Cox & Ben-Dov (1986).
Planococcoides mumensis (Tang,

(Planococcus) comb. n.

This species was described from the leaves of
Prunus mume in the Zhejiang Province of China.
Although it was not possible to obtain specimens
of P. mumensis for examination, Professor Tang
kindly provided an English translation of his origi-
nal description and a further illustration. From
these it is apparent that this species has auxiliary
setae in all of the cerarii, the anal lobe cerarii each
have 2 or 3 conical setae, multilocular disc pores
are confined to the median areas of the posterior
abdominal segments, and tubular ducts are con-
fined to the posterior abdominal segments of the

1977: 34)

3

venter. Consequently, Planococcus mumensis is
here transferred to Planococcoides.

Also currently included in Planococcoides are
P. ireneus De Lotto from the Afrotropical
Region, Planococcoides robustus Ezzat &
McConnell from India, and P. pauliani Mamet
from Madagascar.
Crisicoccus matesovae

(Planococcus) comb. n.

(Danzig, 1986: 21)

Checklist of species here included in

Planococcus

This list includes new combinations, new synonymies,
distributions and hostplant preferences where known.
The original generic names, if these are not Planococ-
cus, are given in parenthesis.

aemulor De Lotto, 1964.

Afrotropical Region, on Combretum spendens and
Asparagus sp.

aphelus De Lotto, 1967.

Afrotropical Region, on Phylica sp.

boafoensis (Strickland, 1947) (Tylococcus).
Afrotropical Region, on Musanga spp.

citri (Risso, 1813) (Dorthesia).
Phenacoccus spiriferus Hempel, 1900. Syn. n.
Planococcoides cubanensis Ezzat & McConnell, 1956.
Syn. n.
Planococcus cucurbitae Ezzat & McConnell, 1956.
Syn. n.
Cosmopolitan, on many hostplants, especially citrus,
coffee and cocoa.

dendrobii Ezzat & McConnell, 1956.
Oriental Region, on orchids.

dioscoreae Williams, 1960.
Austro-oriental Region, usually on yams.

dorsospinosus Ezzat McConnell, 1956.
Planococcus myrsinephilus Borchsenius,
Syn. n.
Planococcus sinensis Borchsenius, 1962. Syn. n.
Palaearctic, Oriental and Austro-oriental Regions, ona
variety of trees.

dubius Cox, 1987.
New Zealand, on Dracophyllum.

epulus De Lotto, 1964.
Afrotropical Region, on Pterolobium lacerans.

1962.

ficus (Signoret, 1875) (Dactylopius).
Dactylopius subterraneus Hempel, 1901. Syn. n.
Palaearctic, Afrotropical, Oriental and Neotropical
Regions, on a variety of hostplants, especially
grapevines, figs and pomegranates.

flagellatus De Lotto, 1961.
Afrotropical Region, on a variety of hosts.

4

furcisetosus Mamet, 1959.
Malagasian Region, on Diospypros sp.

halli Ezzat & McConnell, 1956.
Neotropical and Afrotropical Regions, on a variety of
hostplants, especially yams.

hospitus De Lotto, 1961.
Afrotropical Region, on orchids.

hosyni Ezzat & McConnell, 1956.
Afrotropical Region on orchids.

japonicus sp. 0.
Japan (frequently intercepted in the U.S.A.), on a
variety of trees, including loquat.

kenyae (Le Pelley, 1935) (Pseudococcus).
Planococcus subukiaensis De Lotto, 1954. Syn. n.
Afrotropical Region, on a variety of hostplants,
especially coffee and cocoa.

kraunhiae (Kuwana, 1902) (Dactylopius).
Planococcus siakwanensis Borchsenius, 1962. Syn. n.
Palaearctic, Oriental and Nearctic Regions, on a variety
of trees, including citrus.

lilacinus (Cockerell, 1905) (Pseudococcus).
Pseudococcus deceptor Betrem, 1937. Syn. n.
Malagasian, Oriental, Austro-oriental and Neotropical
Regions, on a variety of hostplants including citrus,
guava, cocao, coffee, custard apple and mango.

litchi sp. n.

Palaearctic, Oriental and Austro-oriental Regions (all
interceptions in the U.K. or U.S.A.), onlychee and
loquat.

mali Ezzat & McConnell, 1956.

New Zealand and Australia, on a variety of woody
hostplants, especially apple and blackcurrant.

martini sp. A.
Austro-oriental Region, on Araceae.

minor (Maskell, 1897) (Pseudococcus). Nom. rev.,
stat. n., comb. n.
Planococcus pacificus Cox, 1981. Syn. n.
Oriental, Austro-oriental, Australian, Polynesian, Neo-
tropical and Malagasian Regions, on a wide variety
of hostplants including cocoa and coffee.

morrisoni (Ezzat & McConnell, 1956) (Allococcus).

Oriental and Austro-oriental Regions (all intercepted in
the U.S.A.), on mangosteen, lychee, Lansium
domesticum and Melicoccus bijugatus.

nigritulus De Lotto, 1961.

Afrotropical Region, on Phoenix dactylifera and Ficus
sp.

orchidi sp. n.

Afrotropical Region, on orchids.

philippinensis Ezzat & McConnell, 1956.

Austro-oriental Region, on orchids.

principe sp. n.

Afrotropical Region, on cocoa.

JENNIFER M. COX
psidii sp. n.
Austro-oriental Region, on Psidium guajava.

subterraneus De Lotto, 1964.
Afrotropical Region, on Ficus sp.

sulawesi sp. 0.
Austro-oriental Region, on Urticaceae.

taigae Danzig, 1986.
Palaearctic Region, on Cupressaceae.

tanzaniensis sp. n.
Afrotropical Region, on Hevea braziliensis.

vovae (Nasonov, 1908) (Pseudococcus).
Palaearctic and Neotropical Regions, on Cupressaceae.

zairensis sp. n.
Afrotropical Region.

Species-groups in Planococcus

Several, apparently monophyletic, species-groups
can be distinguished amongst this assemblage of
species. The citri-group contains those species
which have marginal multilocular disc pores on
the abdominal venter, tubular ducts on the venter
of all abdominal segments and on the head and
thorax, and flagellate dorsal setae. As it contains
the type-species of the genus, this group con-
stitutes Planococcus sensu stricto, and includes P.
aphelus, P. citri, P. epulus, P. ficus, P. flagellatus,
P. halli, P. minor, P. nigritulus, P. subterraneus,
P. tanzaniensis and probably P. aemulor and P.
kenyae. P. aemulor lacks tubular ducts on the
head and thorax, while P. kenyae lacks marginal
multilocular disc pores and, in small specimens, it
sometimes also lacks tubular ducts on the head
and thorax. All of these species, except P. minor,
occur in the Mediterranean Basin or the
Afrotropical Region, although P. citri, P. ficus
and P. halli have been transported to other parts
of the world. The geographical origin of P. minor
is not clear - its current distribution includes the
South Pacific, the Oriental, Austro-oriental, Aus-
tralian, Malagasian and Neotropical Regions.
Another group, the dendrobii-group, contains
eight species of which five have been found only
on Orchidaceae. All eight species are rotund,
have stout legs and have the multilocular disc
pores and tubular ducts confined to the posterior
abdominal segments. All of the orchid-feeding
species have knobbed dorsal setae. Two of these
species, P. dendrobii and P. philippinensis, are
from the Oriental Region, and 3 species, P. hospi-
tus, P. hosyni and P. orchidi, are from the
Afrotropical Region. P. zairensis from an
unknown hostplant in Zaire and P. martini from
Araceae in Indonesia are similar to the above

MEALYBUG GENUS PLANOCOCCUS

species, but lack knobbed dorsal setae. P. fur-
cisetosus from ebony in Madagascar differs from
the orchid-feeding species only by having furcate
dorsal setae. P. principe may belong in this group,
but this species has a few marginal multilocular
disc pores. The body shape and distributions of
the multilocular disc pores and tubular ducts of
the dendrobii-group are similar to those found in
most species of Planococoides and it may not be a
genuine component of Planococcus.

A third group, the dorsospinosus-group, com-
prises those species which lack marginal multi-
locular disc pores and have conical dorsal setae
with associated aggregations of trilocular pores.
These species are from the Oriental and Austro-
oriental Regions and Japan. The dorsospinosus-
group comprises P. dioscoreae, P. dorsospinosus
and P. litchi. P. sulawesi clearly also belongs to
this group, but in this species the enlarged dorsal
setae are very long and flagellate. P. krauhniae
from Japan and P. psidii from West Malaysia may
also belong to this group, but both have marginal
multilocular disc pores and the trilocular pores are
not as obviously associated with the enlarged dor-
sal setae as in the above-mentioned species.

The two species of the vovae-group, P. taigae
and P. vovae, are both confined to Cupressaceae.
Each has a distinct geographical range, P. vovae
occurring in central and southern Europe, the
Caucasus and montane Central Asia (also Brazil),
and P. taigae in the north-eastern parts of the
Palaearctic Region. These species are
characterized by their flagellate cerarian setae,
numerous dorsal tubular ducts, and by lacking
marginal multilocular disc pores. Otherwise, they
are similar to the P. citri-group. Crisicoccus
matesovae, known only from Cupressaceae in
Altai, U.S.S.R., is very similar to both of these
species, but has fewer than 18 pairs of cerarii and
has consequently been transferred from
Planococcus to Crisicoccus (see above).

The remaining group that can be distinguished
is the mali-group comprising P. japonicus, P.
mali, and perhaps P. morrisoni. These species are
characterized by having short, stout, almost coni-
cal, dorsal setae and a marginal group of tubular
ducts adjacent to the anterior spiracles, while
these ducts are absent, or in very low numbers, on
the margins of the head and mesothorax. In addi-
tion, P. japonicus and P. mali both have more
than 2 conical setae in most of the cephalic and
thoracic cerarii. P. japonicus is known only from
Japan, P. mali only from Australia and New Zea-
land (although its true origin may lie closer to
Japan - see under remarks for this species), and P.
morrisoni is from the Oriental Region. In P. mali,
some of the pairs of cerarii may be indistinct or

=)

apparently absent, making it appear to be a mem-
ber of Crisicoccus, although most specimens have
a full complement of 18 pairs of cerarii. Both P.
japonicus and P. mali are similar to species of
Crisicoccus, such as C. azaleae (Tinsley) and C.
matsumotoi (Shiraiwa), and may, in fact, be more
closely related to these species than to the P. citri-
group. C. azaleae and C. matsumotoi have been
illustrated in this work to allow the easy distinc-
tion of these species.

A few remaining species have no obvious
affinities with any of the above groups. P. boa-
foensis, from the Afrotropical Region, is
remarkable in having the cerarii situated on pro-
tuberances, but otherwise has very reduced num-
bers of multilocular disc pores and tubular ducts
and has short, stout, dorsal setae. It is perhaps
closest to the dendrobii-group. The Oriental P.
lilacinus is an enigma, having a rotund body, stout
legs, reduced numbers of multilocular disc pores,
but tubular ducts extending around the margins of
the entire body. The dorsal setae are very long,
and quite stout. The true origin of this species may
be from outside the genera discussed here.

From the above it can be seen that the species of
Planococcus revised in this work almost certainly
do not comprise a monophyletic group. However,
the prime purpose of this study is not to propose a
generic system based on phylogenetic relation-
ships, but to provide a means for the identification
of a group of similar looking species, many of
which are real or potential pests of economically
important plants. To this end, illustrations have
been provided of some similar (and perhaps
related) species currently placed in other genera
with which some of the species covered here might
be confused.

Terminology

The major characters used are illustrated in Fig. 1.
Cerarii are counted starting at the anterior end.
Abdominal segments are counted in such a way
that the circulus lies between segments III and IV,
and the vulva lies between segments VII and VIII.
Although the translucent pores on the hind legs
usually occur on the hind surface, and hence
would be hidden in the illustrations as presented,
they have been illustrated here as if they are on the
frontal surface of the legs. Numbers of pores and
ducts, when given in the text, are totals of both
sides of the body - mealybugs are frequently not
symmetrical in the distribution of their characters.

PREOCULAR CERARIUS - - —- - >. Ay

6th CERARIUS — — -— —/b

8th CERARIUS —

PROTHORAX

MESOTHORAX

METATHORAX

JENNIFER M. COX

i-
|
I
I
\
|
|
I
I
!

ABDOMEN - — -;
!
|
|
|
I
|
!
|
!
|
j=

ANAL RING SETA ~-~--------—-

Fig. 1 Generalized diagram of Planococcus sp. showing numbering of body

segments and cerarii.

Depositories

BMNH British Museum (Natural History), London,
U.K.

IBSP Instituto Biologico de Sao Paulo, Brazil.

MNHN Muséum National d‘Histoire Naturelle,
Paris, France.

MZSP Museu de Zoologia, Universidade de Sao
Paulo, Brazil.

NZAC_ New Zealand Arthropod Collection, Auck-

land, New Zealand.

SANC

UCD
USNM

VCCB

VCclI

ZIL

South African National Collection of Insects,
Pretoria, South Africa.

University of California, Davis, U.S.A.
United States National Museum, Washington
D.C., U.S.A.

Vernalna Collection, Universidade Federal
do Parana, Curitiba, Brazil.

Coccoidea Collection, Department of Ento-
mology, The Volcani Center, Bet Dagan,
Israel.

Zoological Institute, U.S.S.R. Academy of
Sciences, Leningrad, U.S.S.R.

MEALYBUG GENUS PLANOCOCCUS

Acknowledgements

Ithank Dr E. M. Danzig (ZIL), Mme D. Matile-Ferrero
(MNHN), Dr D. R. Miller, Mr S. Nakahara and Mr V.
L. Blackburn (United States Department of Agricul-
ture), Mrs C. F. Morales (NZAC), Mr R. D. Schuster
(UCD) and Dr Yair Ben-Dov (VCI) for the loan of
specimens; and my colleagues, especially Dr D. J.
Williams, for helpful advice and criticism. Figs 11 and 14
are reproduced with the permission of Bulletin of Ento-
mological Research, and Figs 20, 22 and 27 with the
permission of Journal of Natural History.

SYSTEMATICS

PLANOCOCCUS Ferris

Planococcus Ferris, 1950: 164. Type-species:
Dactylopius citri Risso, by original designation.

Allococcus Ezzat McConnell, 1956: 13. Type-spe-
cies: Pseudococcus inamabilis Hambleton, by
original designation. [Synonymy by Cox & Ben-
Dov, 1986.]

ADULT FEMALE. Body shape oval to rotund.
Cerarii numbering 18 pairs, thoracic cerarii indis-
tinct in some specimens of a few species, but other
specimens of that species having the full number;
abdominal cerarii each with 2 conical setae and
several long flagellate auxiliary setae, thoracic
and remaining abdominal cerarii each with only 2
conical or flagellate setae and without auxiliary
setae, cephalic cerarii each with 1-5 conical or
flagellate setae. Circulus usually present, quadr-
ate in most species, but small and round in some
species. Legs well-developed in most species, but
short and stout in some species; translucent pores
usually present on hind coxae and tibiae only,
sometimes present also on hind femora in P. ficus,
absent from the hind tibiae in P. dendrobii and P.
philippinensis and apparently completely absent
from the hind legs in P. aemulor and P. hospitus;
claw without denticle. Anal lobe bars always
apparent. Anal ring of usual pseudococcid form,
with 6 long setae.

Multilocular disc pores usually confined to ven-
ter, although present on dorsum in P. epulus and
P. dubius; always present in at least single rows
across posterior borders of mid-regions of pos-
terior abdominal segments, variably present on
head, thorax, anterior borders of abdominal seg-
ments and margins of abdominal segments. Tri-
locular pores usually evenly distributed, but
aggregated around bases of enlarged dorsal setae

7

in the dorsospinosus-group. Tubular ducts gener-
ally of the oral collar type, sparse to numerous on
the venter, sometimes present on the dorsum
where they may appear to have rims. Simple pores
generally smaller than the trilocular pores, but
larger in some species; often aggregated in groups
on the dorsal mid-regions of the anterior abdomi-
nal segments. Setae long, fine and flagellate on
median areas of venter; variable on dorsum,
flagellate in the citri- and vovae-groups, stout,
almost lanceolate, in the mali-group, conical in
the dorsospinosus-group and knobbed in the
dendrobii-group; setae on the marginal areas of
the venter often similar to those on the dorsum.

Key to species of Planococcus
1 Multilocular disc pores present on dorsum .... 2
— Miultilocular disc pores absent from dorsum ... 3

2 Most cephalic and thoracic cerarii each with three
conical setae, dorsal setae stout, found in New Zea-
landionly (Bigs 9) Renee Sas dubius

— Most cephalic and thoracic cerarii each with 2 coni-
cal setae, dorsal setae slender, found in Kenya only
(Big FRO PeR wate oateren eine ese atite ee ss epulus

3 Allcerarii situated on very pronounced sclerotized
protuberances, tubular ducts and multilocular disc
pores few in number and restricted to median areas
of the posterior abdominal segments, dorsal setae
short and stout, almost lanceolate, known only
from the Afrotropical Region (Fig. 4) . boafoensis

— Cerarii not situated on protuberances, although
very young adults may sometimes give this
appearance, other characters various ......... 4

4 Anal lobe cerarii each situated on a large, promi-
nent, sclerotized area, body oval to rotund, cisanal
setae longer than anal ring setae, multilocular disc
pores very few or absent on margins and on anterior
edges of median areas of abdominal segments, dor-
sal tubular ducts absent, ventral tubular ducts pre-
sent in marginal groups on all abdominal segments,
often also present on head and thorax, found in the
Afrotropical Region (Fig. 20) ........... kenyae

— Anal lobe cerarii situated on small, not particularly
prominent, sclerotized areas, other characters vari-
SUS eae eth SO RR weed ew ES os 5)

5 Oral collar tubular ducts absent from both median
and marginal areas of head and thorax........ 6

— Oral collar tubular ducts usually present on median
areas of thorax, if absent, then present on margins

Of Neadlan a COLA a ..c5 sreneasi sais evayss biay>euso a0 « 14
6 Many dorsal setae furcate, found in Madagascar
Only (Eiga013)) ote x eas ee -y-teener furcisetosus
= Dorsalisetaenever turcate™ yan vses ace ene. i

10

14

Ventral simple pores as large as the multilocular
disc pores, known only from Uganda (Fig. 15)
hospitus

Ventral simple pores, if apparent, considerably
smaller than the multilocular disc pores ....... 8

No more than 1 tubular duct present on each side of
each abdominal segment, known only from the
Atroiropical Regionian..- 3.264 s. ere eee 9

Tubular ducts present in small groups on either side
of abdominal segments VI and VII, known from the
Afrotropical, Oriental and Austro-oriental
IRESIONS cies csicnegete Mi taegh teas ae bets ier orien ests 10

Dorsal setae short and stout, tubular ducts and
multilocular disc pores usually present singly on
margins of some posterior abdominal segments
(HIG AAO) actrees tva cnpalstele earnest zairensis

Dorsal setae moderately long and fine, margins of
abdominal segments never with tubular ducts or
multilocular disc pores, known only from the
Afrotropical Region (Fig. 16) ........... hosyni

Margins of posterior abdominal segments each with
a few multilocular disc pores, translucent pores
absent from hind legs, dorsal setae not knobbed,
known only from Kenya (Fig. 2) ....... aemulor

Margins of abdominal segments without multilocu-
lar disc pores, translucent pores present on hind
coxae, often also on hind tibiae, dorsal setae some-
times knobbed

Dorsal setae not knobbed, antennae 7-segmented,
known only from Indonesia (Fig. 26) .... martini

Dorsal setae distinctly knobbed, antennae 8-seg-
MEMEO) sais Fee's ery scmer sures evtagne sae Riovevanere se 12

Simple pores on venter very conspicuous and about
twice the size of the trilocular pores, translucent
pores present on hind tibiae, known only from Lib-
eniau(big. 80)» soniaueet aad. beer eee orchidi

Simple pores inconspicuous and about half the size
of the trilocular pores, translucent pores absent
from hind tibiae, found in the Oriental and Austro-
OKEMTAIRE PIONS cr ccc teers recipe eae 13

Circulus present, cisanal setae shorter or about the
same length as the anal ring setae, simple pores on
mid-region of dorsum smaller than the trilocular
POLESH (HIS). cmt cc tale ancen cin dene dendrobii

Circulus absent, cisanal setae longer than the anal
ring setae, simple pores on mid-region of dorsum,
particularly on the posterior segments, larger than
the trilocular pores (Fig. 31) ...... philippinensis

Tubular ducts with pronounced oral rims present on
dorsum adjacent to several cerarii and scattered
over median area of dorsum, dorsal setae short and
stout, almost lanceolate, found in the Oriental and
Austro-oriental Regions (Fig. 28) ..... morrisoni

If more than 2 oral rim tubular ducts present on
dorsum, then dorsal setae long and slender ... 15

15

16

20

21

22

JENNIFER M. COX

Several cephalic and thoracic cerarii usually each
with more than 2 conical setae, some thoracic and
anterior cerarii usually indistinct and with only 1
conical seta jad ce AS. Se eee 16

Cephalic and thoracic cerarii each with only two
conical setae, except for preocular pair which some-
times have three conical setae in each cerarius, all
cerarii distinct and with 2 conical setae ....... 17

Number of ventral tubular ducts on margins of pro-
thorax totalling 0-45, length of hind tibia + tarsus
305-410 um, known only from Australia and New
Lealand (Eig. 24)) 07... ..3656 qeeee epee mali

Number of ventral tubular ducts on margins of pro-

thorax totalling 24-63, length of hind tibia + tarsus

235-315 wm, known only from Japan (Fig. 17)
japonicus

Multilocular disc pores present on margins of
abdominal segments, even if only 1 or 2 per segment

18
Multilocular disc pores completely absent from
margins of abdominal segments ............. 26

Many simple pores on both venter and dorsum
almost as large as the multilocular disc pores,
known only from Principe (Fig. 32) ..... principe

Simple pores on both venter and dorsum never
more than twice the size of the trilocular pores 19

Dorsal setae with distinctly swollen bases, multi-
locular disc pores very sparse on margins of abdom-
inal segments, oral collar tubular ducts on dorsum
in groups of 2-5 ducts adjacent to some abdominal
cerarii, found in Japan and U.S.A. (Fig. 21)
kraunhiae

Dorsal setae without swollen bases, multilocular
disc pores moderately numerous on margins of
abdominal segments, tubular ducts, if present on
dorsum, usually present singly adjacent to cerarii

20
Circulus absent, known only from South Africa
(Hig23) 2. kth ee eee aphelus
Circulus present: o.oo coer ss iene eee 21

Oral collar tubular ducts absent from abdominal
segments IX, and often also from VIII, body
broadly oval to rotund, ventral simple pores at least
same size as the trilocular pores, dorsal setae stout,
found only in the Afrotropical Region ....... 22

Oral collar tubular ducts present on venter of both
abodominal segments VIII IX, body elongate-oval
to broadly oval, simple pores smaller than the tri-
locular pores, dorsal setae slender, widely dis-
11g 10) U10e(0 EMME Oso e oman Aida x54 3 23

Oral collar tubular ducts present on ventral, median
areas of thorax, tubular ducts usually present on
venter of abdominal segment VIII, simple pores
about the same size as the trilocular pores (Fig. 38)

tanzaniensis

MEALYBUG GENUS PLANOCOCCUS

23

24

25

Oral collar tubular ducts absent from median areas
of thorax, although present on margins of head and
thorax, tubular ducts absent from both abdominal
segments VIII IX, ventral simple pores about twice
the size of the trilocular pores (Fig. 29) nigritulus

Dorsal setae long and fleshy, longest setae on
abdominal segment VI or VII up to 100 ym long,
sometimes strongly bifurcate ............... 24

Dorsal setae either short (less than 30 xm long) or if
longer, then slender, never fleshy or bifurcate 25

Several dorsal setae strongly bifurcate, oral collar
tubular ducts absent from head, known only from
South Africa (Fig. 34) subterraneus

No more than 2 dorsal setae bifurcate, at least 4 oral
collar tubular ducts present on head, found in the
Afrotropical Region (Fig. 12) ........ flagellatus

Head with 0-35 oral collar tubular ducts, longest
seta on median area of abdominal segment VI or
VII 17-33 um long, cerarian setae on head and
thorax always conical, dorsum rarely with more
than 6 oral collar tubular ducts, median ventral area
of abdominal segment VII with a single or double
row of multilocular pores, translucent pores never
PLeSeMUOn MING TEMIOLa 2. cc. e cee ewe pes 26

Head with 0-4 oral collar tubular ducts, longest seta
on median area of abdominal segment VI or VII
25-50 yum long, cerarian setae on head and thorax
often long and slender, dorsum with 0-20 oral collar

Table 1 Separation of P. citri and P. minor.

26

27

9

tubular ducts, median ventral area of abdominal
segment VII usually with a single row of multilocu-
lar disc pores, translucent pores sometimes present
on) bind femora) oz. acl easter aaitaccieek 27

Dorsal setae blunt-ended, translucent pores on
hind legs noticeably large and distinct, known only
from West Malaysia (Fig. 33) ............ psidii

Dorsal setae flagellate, translucent pores on hind
legs typical of genus, worldwide
(see Table 1) citri/minor

Translucent pores often apparent on one or both
hind femora; total number of multilocular disc
pores behind the front coxae 0-17; rows of median
multilocular disc pores usually present on abdomi-
nal segments IV-VII, sometimes also on III; 1 or 2
multilocular disc pores usually present on each mar-
gin of abdominal segments II and III; common on
grapevines in the Mediterranean Basin, Pakistan,
Argentina and South Africa (Fig. 11) ...... ficus

Translucent pores absent from hind femora; multi-
locular disc pores behind front coxae totalling 0-5;
rows of multilocular disc pores present on abdomi-
nal segments III-VII, often also on II; multilocular
disc pores seldom present on margins of abdominal
segment II, sometimes present on margins of III;
common on Dioscorea spp. in the West Indies and
the Afrotropical Region (Fig. 14) halli

Character Value Score
(All numbers are the totals from
both sides of the body)
A. Number of ventral tubular ducts on head 0-3 0
4-13 10
14-35 40
B. Number of ventral tubular ducts adjacent to 0-2 0
8th pair (numbering from the anterior) of 3-7 10
ceraril 8-30 40
C. Tubular ducts present between 2nd and 3rd
cerarii on head on at least one side of the yes 10
body no 0
D. Number of multilocular disc pores behind 0-6 5
front coxae 7-12 0
E. Ratio of length of hind tibia + tarsus to 1.00-1.07 0
length of trochanter + femur 1.08-1.17 5
1.18-1.30 10
F. Width of row of multilocular disc pores on single row 15
posterior margin of abdominal segment VI intermediate 5
double row 0

Scores:
35-120 = citri (Fig. 5) (p. 000)

0-35 = minor (Fig. 27) (p. 000)

10

28 Cephalic cerarian setae flagellate, dorsal tubular
ducts in rows across posterior abdominal segments,
OM CuUpTeESSACAE! ja: sree, aster eabis) leaecencue cereus 29

— Cephalic cerarian setae usually conical, dorsal
tubular ducts, if present, occurring singly adjacent
to cerarii, not found on Cupressaceae ....... 30

29 Dorsal tubular ducts in rows of up to 24 ducts across
posterior abdominal segments, total of more than
200 ducts on dorsum, found in the eastern Palaearc-
ticiRe sion (Eisa36) sea e ee eee taigae

— Dorsal tubular ducts in rows of no more than 18
ducts across posterior abdominal segments, total of
13-110 ducts on dorsum, widespread in the western
Palaearctic Region and also in Brazil (Fig. 39)

vovae

30 All dorsal setae noticeably long and flagellate, long-
est seta on abdominal segment VI or VII 50-140 wm
long, with or without trilocular pores associated
with their bases. 22 ee un oe vee eet be: 31

— Most dorsal setae conical, some almost as large as
cerarian setae, and with one or more trilocular
pores associated with their bases ............ yy

31 Legs noticeably short and stout, ratio of hind tibia
+ tarsus to hind trochanter + femur 0.77—0.97;
dorsal setae without trilocular pores associated with
their bases; found in the Oriental, Austro-oriental,
Malagasian and Neotropical Regions (Fig. 22)

lilacinus

— Legs elongate, ratio of hind tibia + tarsus to hind
trochanter + femur 1.06-1.07; larger dorsal setae
with one or more trilocular pores associated with
their bases; known from Indonesia only (Fig. 35)

sulawesi

32 Several dorsal setae associated in pairs (Fig. 23)
litchi

—7) Dorsalisctac notin) parser etree 33

33 Ventral tubular ducts adjacent to eyes totalling
6-22, found in Papua New Guinea and the Solomon
Islands, usually on Dioscorea sp. (Fig. 7)

dioscoreae

— Ventral tubular ducts adjacent to eyes totalling no
more than 7, known only from Japan and the Orien-

tal Region on a variety of hosts (Fig. 8)
dorsospinosus

Descriptions of species

Planococcus aemulor De Lotto
(Fig. 2)

Planococcus aemulor De Lotto, 1964: 372. Holo-
type 9, Kenya: on Combretrum splendens
(BMNH) [examined].

JENNIFER M. COX

ADULT FEMALE. Mounted specimens broadly oval,
length 1.6-1.9 mm, width 1.1-1.4 mm. Antennae
7- or 8-segmented, if 7-segmented, then pseudoar-
ticulation apparent on segment IV. Margin of
body with complete series of 18 pairs of cerarii,
each cerarius with 2 (rarely 3 on head) slender
conical setae. Legs elongate; hind trochanter +
femur 255-280 um long, hind tibia + tarsus 265-
285 um long, ratio of lengths of hind tibia + tarsus
to hind trochanter + femur 1.00—1.07; translucent
pores not apparent on hind legs. Inner edges of
ostioles moderately sclerotized. Circulus large
and probably quadrate, although generally dis-
torted in mounted specimens, width 170-200 um.
Cisanal setae shorter than anal ring setae. Anal
lobe area apparently only slightly sclerotized.

Venter. Multilocular disc pores moderately
numerous around vulva, present in single rows
across posterior edges of median areas of abdomi-
nal segments III-VII or IV-VII, a few pores pre-
sent on anterior edge of median area of segment
VII and on margins of at least some posterior
segments. Trilocular pores numerous and evenly
distributed. Oral collar tubular ducts, apparently
of 1 size and confined to the abdomen, present
sparsely in rows across median areas of abdominal
segments III-VII and in marginal groups on seg-
ments IV-VIII, but usually absent from segments
VIII or IX. Simple pores slightly larger than the
trilocular pores, sparsely scattered over venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores of 2 sizes, larger pores as for venter,
smaller pores about a third of the size of the tri-
locular pores, sparsely scattered over dorsum.
Setae stout, blunt-ended and moderately short,
longest seta on abdominal segments VI or VII
25-35 um long.

MATERIAL EXAMINED

Holotype 2, Kenya: Ruiru, on Combretum splen-
dens, 20.viii.1957 (R. H. Le Pelley) (BMNH). 6
paratype 9, same data as holotype (BMNH).

DistriBuTION. Afrotropical Region: Kenya.

HostTpLants. Combretaceae: Combretum splen-
dens. Also recorded from Asparagus sp.
(Liliaceae) (De Lotto, 1964).

RemarkKS. P. aemulor resembles P. nigritulus in
lacking tubular ducts on the median areas of the
thoracic venter and in usually lacking ducts on
abdominal segments VIII andIX. The two species
may be distinguished by the presence of marginal
ventral tubular ducts on the head and thorax and
translucent pores on the hind coxae and tibiae in
P. nigritulus, both of which characters are lacking

MEALYBUG GENUS PLANOCOCCUS

Fig. 2 Planococcus aemulor De Lotto.

11

12

Fig. 3

Planococcus aphelus De Lotto.

JENNIFER M. COX

MEALYBUG GENUS PLANOCOCCUS

from P. aemulor. The complete lack of tubular
ducts on the thorax in P. aemulor together with
the short, stout, blunt-ended dorsal setae indicate
that it may be related to the orchid-feeding
dendrobii-group rather than to the citri-group.

The only other known species of Planococcus
completely lacking translucent pores on the hind
legs is P. hospitus.

Planococcus aphelus De Lotto
(Fig. 3)

Planococcus aphelus De Lotto, 1967: 16. Holo-
type 9, SoutH Arrica: on roots of Phylica sp.
(SANC) [not examined].

ADULT FEMALE. Mounted specimens oval, length
about 1.4 mm, width about 0.9 mm. Antennae 7-
or 8-segmented, if 7-segmented, then pseudoar-
ticulation apparent on segment IV. Margin of
body with complete series of 18 pairs of cerarii,
each cerarius with 2 slender conical setae. Legs
elongate; hind trochanter + femur 230-265 um
long, hind tibia + tarsus 230-280 um long, ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur about 1.06; translucent pores apparent on
hind coxae and tibiae. Inner edges of ostioles
moderately sclerotized. Circulus absent. Cisanal
setae shorter than anal ring setae. Anal lobe
cerarii each situated on a moderately sized, mod-
erately sclerotized area.

Venter. Multilocular disc pores present spar-
sely around vulva, in single rows across posterior
edges of abdominal segments II-VII, a few pre-
sent on anterior edges of segments V-VII, present
singly or in small groups on margins of at least
some abdominal segments, and sparsely scattered
over median areas of thorax. Trilocular pores
moderately numerous and evenly distributed.
Oral collar tubular ducts apparently of 1 size,
present sparsely in rows across median areas of
abdominal segments II-VII, in small marginal
groups on segments II-VIII, scattered over
median area of thorax, and present singly on the
margins of some thoracic segments. Simple pores
considerably smaller than the trilocular pores,
sparsely but evenly distributed.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores and simple pores as
for venter. Setae flagellate and of moderate
length, longest seta on abdominal segments VI or
VII 25-35 um long.

13
MATERIAL EXAMINED
South Africa: 2 paratype Q, Cape Province,

Somerset West, on roots of Phylica sp., 26.xi.1964
(J. Munting) (BMNH).

DISTRIBUTION.
Africa.

Afrotropical Region: South

HostpLant. Rhamnaceae: Phylica sp.

RemakkKS. The distribution of the multilocular disc
pores shows this species to be a member of the
citri-group. It differs from the other members of
this group by lacking a circulus.

Planococcus boafoensis (Strickland)
(Fig. 4)

Tylococcus boafoensis Strickland, 1947: 151;
Williams, 1958: 30. Syntypes 9, GHANA: on
Musanga_ smithii (BMNH) [11 syntypes
examined].

Planococcus boafoensis (Strickland) De Lotto,
1964: 372.

ADULT FEMALE. Mounted specimens elongate-
oval, length 1.3-2.2 mm, width 0.6-1.3 mm. Mar-
gin of body with complete series of 18 pairs of
cerarii situated on sclerotised protuberances, each
cerarius with 2 (sometimes 3 on head and/or
thorax) slender conical setae. Legs elongate; hind
trochanter + femur 215—265 xm long, hind + tibia
tarsus 225-275 um long, ratio of lengths of hind +
tibia tarsus to hind trochanter + femur 1.04-1.08;
translucent pores apparent on hind coxae and
tibiae. Inner edges of ostioles lightly sclerotized.
Circulus quadrate or oval, width 50-70 um. Cisa-
nal setae shorter than anal ring setae.

Venter. Multilocular disc pores confined to
median areas of abdomen, sparsely present
around vulva and in a single row across posterior
edge of abdominal segment VI. Trilocular pores
very sparse and evenly distributed. Oral collar
tubular ducts of 1 size, confined to abdomen, a
few present on median areas and margins of
abdominal segments VI or VII. Simple pores
apparently absent.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores somewhat sparse
with 1-3 pores associated with the bases of the
larger setae. Simple pores about a third of the size
of the trilocular pores, sparsely but evenly dis-
tributed. Setae short and stout, almost lanceolate,
longest seta on abdominal segments VI or VII
12-16 wm long.

14

JENNIFER M. COX

Fig. 4 Planococcus boafoensis (Strickland).

MATERIAL EXAMINED

12 Q (including type specimens listed below)
(BMNH, MNNH).

Ghana (Gold Coast): 11 syntype 9, Tafo, on Mus-
anga smithii, 8.xi.1945 (E. O. Boafo) (BMNH).

DISTRIBUTION.
Zaire.

Afrotropical Region: Ghana,

Hostp.ants. Urticaceae: Musanga cecropioides,
M. smithii.

REMARKS. This species is readily recognizable
amongst the other known species of Planococcus
by the sclerotised protuberances which bear the
cerarii. Its relationships are difficult to determine,
but it resembles the dendrobii-group in its dis-
tribution of multilocular disc pores and oral collar
tubular ducts. Cyperia angolia De Lotto has very

15

MEALYBUG GENUS PLANOCOCCUS

Ir) «

“BQ.

Fig. 5 Planococcus citri (Risso).

16

similar sclerotized protuberances, but differs in
having flagellate auxiliary setae and more than 2
conical setae in most of the cerarii, and by having
much longer dorsal setae than Planococcus
boafoensis.

Planococcus citri (Risso)
(Fig. 5)

Dorthesia citri Risso, 1813: 416. Syntypes 9,
FRANCE: Menton, on Citrus sp. (probably lost).

Coccus citri (Risso) Boisduval, 1867: 348.

Dactylopius citri (Risso) Signoret, 1875: 312.

Lecanium phyllococcus Ashmead, 1879: 160.
U.S.A.: Florida, on orange trees (status of type
material not known). [Synonymy by Riley,
1888.]

Dactylopius destructor Comstock, 1881: 342.
U.S.A.: Florida, on orange trees (status of type
material not known). [Synonymy by Penzig,
1887.]

Phenacoccus spiriferus Hempel, 1900: 389. Syn-
types 2, BRAZIL: on cultivated tree (MZSP) [5
syntypes examined]. Syn. n.

Phenacoccus spiniferis Hempel; Hempel, 1901a:
110. [Misspelling. ]

Pseudococcus citri (Risso) Fernald, 1903: 99.

Pseudococcus citri var. phenacocciformis Brain,
1915: 116. Syntypes 2, SourH AFRICA: on
Bouvardia sp. (SANC) [not examined]. [Syn-
onymy by Ezzat & McConnell, 1956. ]

Pseudococcus citri var. coleorum Marchal, 1908:
236. Syntypes Q, FRANCE: on Coleus sp.
(BMNH, MNNH) [4 syntypes examined]. [Syn-
onymy by Borchsenius, 1949.]

Planococcus citri (Risso) Ferris, 1950: 165; Ezzat
& McConnell, 1956: 65; Cox & Freeston, 1985:
YER

Planococcus citricus Ezzat & McConnell, 1956:
69. Holotype Q, ITaty (intercepted in U.S.A.):
on Citrus limonia (USNM) [examined]. [Syn-
onymy by Cox, 1981.]

Planococcoides cubanensis Ezzat & McConnell,
1956: 55. Holotype 2, Cusa (intercepted in
U.S.A.): on Ficus pandurate (USNM) [exam-
ined]. Syn. n.

Planococcus cucurbitae Ezzat & McConnell,
1956: 71. Holotype 9, GRENADA: on Cucurbita
sp. (USNM) [examined]. Syn. n.

ADULT FEMALE. Mounted specimens oval, length
1.6-3.2 mm, width 1.0-2.0 mm. Margin of body
with complete series of 18 pairs of cerarii, each
cerarius with 2 conical setae, except for preocular
cerarii which may have 1 or 3 setae each. Legs
elongate; hind trochanter + femur 220-350 um

JENNIFER M. COX

long, hind tibia + tarsus 260-420 um long, ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 1.1—1.3; translucent pores apparent on hind
coxae and tibiae. Inner edges of ostioles moder-
ately sclerotized. Circulus quadrate, width 120—
200 wm. Cisanal setae shorter than anal ring setae.
Anal lobe cerarii each situated on a small, moder-
ately sclerotized area.

Venter. Multilocular disc pores present around
vulva, in single or double rows across posterior
edges of abdominal segments III-VII, in single
rows across anterior edges of segments V-VII, in
marginal groups on abdominal segments IV-VII
and sometimes a few pores scattered over median
area of the thorax and head, but no more than a
total of 6 pores behind the front coxae. Trilocular
pores moderately numerous and evenly dis-
tributed. Oral collar tubular ducts of 2 sizes;
smaller ducts present sparsely in rows across
median areas of abdominal segments I- VII; larger
ducts in marginal groups of variable size around
entire venter including head and thorax, and scat-
tered over median area of thorax. Simple pores
about the same size as the trilocular pores, spar-
sely but evenly distributed.

Dorsum. Multilocular disc pores absent. Tubu-
lar ducts without apparent rims and slightly larger
than the larger ducts on the venter, often present
adjacent to some cerarii, 1 or 2 ducts sometimes
present on median areas. Trilocular pores as for
venter. Simple pores of 2 sizes, smaller pores
smaller than the smaller size on the venter, scat-
tered over entire dorsum, larger pores about twice
the size of the trilocular pores, present in small
groups along mid-line of thoracic and anterior
abdominal segments. Setae flagellate and of mod-
erate length, longest seta on abdominal segments
VI or VII 30-35 um long.

MATERIAL EXAMINED.

Several hundred field-collected adult females
from many countries and host-plants in addition
to the reared material used by Cox (1983)
(BMNH, MNHN, USNM, VCI).

Planococcus spiriferus Hempel. 5 syntype 2, on
one slide labelled ‘Phenacoccus spiniferus n. sp.
Type 2 336’. The original description gives the
habitat as Sao Paulo, in the grooves of leaves of a
cultivated tree.

Planococcus citricus Ezzat & McConnell. Holo-
type Q, Italy (intercepted at New York, Hidalgo):
on Citrus limonia, 20.vii.1948 (USNM).
Planococcoides cubanensis Ezzat & McConnell.
Holotype 9, Cuba (intercepted at Hoboken, New
Jersey): on Ficus pandurate, 11.vi.1948 (Bennatt)
(USNM).

MEALYBUG GENUS PLANOCOCCUS

Planococcus cucurbitae Ezzat & McConnell.
Holotype 2, Grenada: on Cucurbita sp., 2.i.1944
(R. G. Fennah) (USNM).

DisTRIBUTION. Almost worldwide, but apparently
absent from some South Pacific Islands and from
the Malagasian Region.

HostpLants. Although P. citri is generally
regarded as being polyphagous, which is indeed
the case in greenhouses, in the field it is noticeably
absent from certain host-plants. Thus De Lotto
(1975) recorded that P. citri, although common on
citrus in South Africa, was never found on gra-
pevines. The mealybug found on vines throughout
the Mediterranean basin is P. ficus and not P.
cltrt.

Remarks. Although there is no extant type
material of P. citri, the identity of this species is
not in question. Risso’s original illustration of the
live insect clearly shows it to be Planococcus, and
the only species of this genus found on Citrus spp.
in the Mediterranean Basin is P. citri. In fact, this
species is well fitted to both its specific name ‘citri
and its common name, ‘citrus mealybug’, in that it
shows a decided preference for this host
throughout its geographical range.

Other species of Planococcus have been misi-
dentified as P. citri. The very similar P. minor was
only differentiated by rearing experiments study-
ing the variation of both species (see Introduc-
tion) but is now known to be the predominant
species in the South Pacific Islands, the Austro-
oriental Region, the Malagasian Region and the
northern Neotropical Region (Cox, 1981;
Williams, 1982; Cox & Freeston, 1985). At one
time P. kenyae was misidentified as P. citri and
parasites sought in California for its control on
coffee in Kenya (see under Remarks for this spe-
cies). For a long time the distinction between P.
citri and P. ficus was unclear, but the separation of
these two species is now universally accepted
(Ezzat & McConnell, 1956; De Lotto, 1975;
Danzig, 1977; Cox, 1981).

P. citricus was considered by Ezzat & McCon-
nell (1956) to differ from P. citri by having smaller
appendages and fewer pores and ducts. Cox
(1983) showed that these characteristics could be
readily induced in P. citri by rearing specimens at
high temperatures.

The syntypes examined of Phenacoccus spir-
iferus have numerous tubular ducts on the margins
of the head and the thoracic segments, and are
clearly P. citri. In the original description, the
antennae are stated as being 9-segmented, but
those of the syntypes are 8-segmented, with a

17

slight pseudo-articulation indicated on the eighth
segment — the measurements of the antennal seg-
ments given by Hempel show the eighth segment
to comprise his eighth and ninth segments

Planococcoides cubanensis was described from
a single specimen by Ezzat & McConnell (1956).
It differs from normal Planococcus citri only by
having three conical setae in each anal lobe
cerarius. As no other similar specimens have been
observed, and as Planococcus is apparently not
native to the New World, this individual is here
considered to be an aberrant specimen of P. citri.

P. curcurbitae was described from a single spec-
imen by Ezzat & McConnell (1956). This speci-
men is heavily parasitized, and its main distinction
from P. citri is that its anal ring setae are only
about as long as the diameter of the anal ring. This
characteristic has not been observed in any other
specimen of Planococcus and is considered here to
be an aberration. By the criteria given in Table 1,
this specimen has been identified as P. citri, rather
than P. minor, which it also resembles.

Morrison (1925) synonymized Dactylopius cal-
ceolariae var. minor with Planococcus citri. How-
ever, examination of the type material of this
species has revealed that it is conspecific with a
subsequently described species, P. pacificus Cox.
No specimens of P. citri from the Malagasian
Region have been observed during the course of
this study.

EcONoMIC STATUS. P. citri is a serious pest of citrus
in most citrus-growing areas of the world and is a
major greenhouse pest in cooler regions.

Planococcus dendrobii Ezzat & McConnell
(Fig. 6)

Planococcus dendrobii Ezzat & McConnell, 1956:
73. Holotype 9, INp1A (intercepted at Hono-
lulu): on Dendrobium transparens (USNM)
[not examined].

ADULT FEMALE. Mounted specimens rotund,
length 1.7-2.6 mm, width 1.2—2.0 mm. Margin of
body with a complete series of 18 pairs of cerarii,
each with 2 stout conical setae. Legs stout; hind
trochanter + femur 235-280 um long, hind tibia +
tarsus 220-250 um long, ratio of lengths of hind
tibia + tarsus to hind trochanter + femur 0.88—
0.94; translucent pores apparent on hind coxae
only. Inner edges of ostioles moderately strongly
sclerotized. Circulus small and oval, width 75-115
pm. Cisanal setae shorter than anal lobe setae.
Anal lobes not sclerotized.

18

JENNIFER M. COX

Fig. 6 Planococcus dendrobii Ezzat & McConnell.

Venter. Multilocular disc pores confined to
median areas, present around vulva and ina single
row across posterior edge of median area of
abdominal segment VI. Trilocular pores numer-
ous and evenly distributed. Oral collar tubular
ducts of 1 size, confined to abdomen, present
sparsely around vulva, a few ducts sometimes pre-
sent on median area of segment V and in marginal
groups on segments VI & VII. Simple pores
slightly smaller than the trilocular pores, sparsely
scattered over entire venter. Setae flagellate and
moderately short on median areas, stout and
knobbed on margins.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores generally as for venter, but more numer-
ous on median areas of posterior abdominal
segments. Setae short, stout and distinctly knob-
bed, length of longest seta on abdominal segments
VI or VII 13-15 pm.

MATERIAL EXAMINED
10 Q (including type material listed below)
(BMNH, USNM).

19

MEALYBUG GENUS PLANOCOCCUS

Fig. 7 Planococcus dioscoreae Williams.

20

India (intercepted at Honolulu): 2 paratype 2, on
Dendrobium moschatum, 16.iv.1947 (S. Namiki)
(USNM).

DIstTRIBUTION. Oriental Region: Bhutan (inter-
cepted at Edinburgh), India (intercepted at
Honolulu). Also recorded from the Philippines
and Thailand (Ezzat & McConnell, 1956).

HosTPLaNnts. Orchidaceae: Dendrobium
moschatum. Also recorded from D. fimbriatum,
D. transparens and Cypripedium sp. (all
Orchidaceae) (Ezzat & McConnell, 1956).

RemakkKsS. This species is very similar to P. philip-
pinensis, having reduced numbers of multilocular
disc pores and tubular ducts, dorsal setae knobbed
and translucent pores absent from hind tibiae.
The two species may be distinguished by the pres-
ence of a circulus in P. dendrobii and by the
smaller simple pores on the dorsum of the pos-
terior abdominal segments in this species. See also
under P. hospitus for further remarks.

EcoNomIC STATUS. This species is a potential pest
of orchids.

Planococcus dioscoreae Williams
(Fig. 7)

Planococcus dioscoreae Williams, 1960: 39. Holo-
type 2, Papua New GuIneEa: on yam (BMNH)
[examined].

ADULT FEMALE. Mounted specimens oval to
broadly oval, length 2.0-2.5 mm, width 1.2-2.0
mm. Margin of body with a complete series of 18
distinct pairs of cerarii, each cerarius with 2 coni-
cal setae except for the preocular cerarii with 2 or
3 setae, all cerarian setae elongate-conical and
with flagellate tips. Legs elongate, hind trochan-
ter + femur 220-285 um long, hind tibia + tarsus
235-285 ym long; ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 1.00—1.08;
translucent pores visible on hind coxae and tibiae.
Inner edges of ostioles lightly sclerotized. Circulus
quadrate and of moderate size, width 65-170 um.
Cisanal setae shorter than anal ring setae. Anal
lobe cerarii each situated on a small, moderately
sclerotised area.

Venter. Multilocular disc pores confined to
median areas, present around vulva, in single
rows across posterior borders of abdominal seg-
ments II-VII and anterior borders (sometimes
reduced to 1 or 2 pores per segment in small
specimens) of abdominal segments III-VII or
IV-VII, and a few spores scattered over median

JENNIFER M. COX

areas of thorax. Trilocular pores sparsely but
evenly distributed. Oral collar tubular ducts of 2
sizes; smaller ducts present in rows across median
areas of abdominal segments VI or VII; larger
ducts present in rows across abdominal segments
III-V, scattered over median areas of thorax, and
present in groups around entire venter, including
a group adjacent to each postocular cerarius. Sim-
ple pores about half the size of the trilocular
pores, sparsely scattered over entire venter.

Dorsum. Multilocular disc pores absent. Tri-
locular pores moderately numerous and aggreg-
ated around bases of enlarged setae. Tubular
ducts, larger than the larger ducts on the venter
and without apparent rims, sometimes present
singly next to some abdominal cerarii, no more
than 2 present on any specimen and frequently
absent. Simple pores as for venter. Dorsal setae of
various sizes, larger setae elongate-conical with a
flagellate apex, about the same size as the cerarian
setae and each with 24 trilocular pores associated
with their bases giving the appearance of dorsal
cerarii, these enlarged setae present on all seg-
ments except the last and usually solitary with no
more than 4 pairs present.

MATERIAL EXAMINED

53 Q (including type material listed below) (all
BMNH).

Holotype 2, Papua New Guinea: Sepik District,
Bunahoj, on yams, 22.vi.1959 VJ. J. H. Szent-
Ivany). Paratypes, 32 2, same data except various
villages, 22.v.-23.vi.1959.

DIsTRIBUTION. Austro-oriental Region: Papua
New Guinea, Solomon Islands.

HostTpLants. Dioscoreaceae: Dioscorea_ sp.
(yams), D. alata. Araceae: Xanthosoma
sagittifolium.

REMARKS. The enlarged dorsal setae with asso-
ciated aggregations of trilocular pores place this
species in the dorsospinosus-group. It differs from
both of the other two species in this group, P.
dorsospinosus and P. litchi, by having 3-10 ventral
tubular ducts adjacent to each post-ocular
cerarius; from P. dorsospinosus by having larger
and more flagellate dorsal setae; and from P. litchi
by having the dorsal setae associated into no more
than 4 pairs. P. dioscoreae is particularly close to
P. dorsospinosus, and the specimens from India
on yams listed under the latter species are some-
what intermediate in their characteristics. The
whole group needs further study to clarify the
relationship between these two species.

Economic status. This species has been found
heavily infesting yams.

=

MEALYBUG GENUS PLANOCOCCUS

Planococcus Ezzat &

McConnell
(Fig. 8)

dorsospinosus

Planococcus dorsospinosus Ezzat & McConnell,
1956: 75. Holotype 9, Cuina (intercepted in
U.S.A.): on Pueraria hirsuta (USNM)
[examined].

Planococcus myrsinephilus Borchsenius, 1962:
585. Holotype 2, Cuina: on Myrsine africana
(ZIL) [not examined]. Syn. n.

Planococcus sinensis Borchsenius, 1962: 586.
Holotype 9, Cuina: on Buddleia officinalis
(ZIL) [not examined]. Syn. n.

ADULT FEMALE. Mounted specimens oval, length
2.4-3.5 mm, width 1.3-2.1 mm. Margin of body
with a complete series of 18 distinct pairs of
cerarii, each cerarius with 2 conical setae except
for the preocular cerarii each with 3 or 4 setae, all
cerarian setae conical and with slightly flagellate
tips. Legs elongate, hind trochanter + femur 245—
370 um long, hind tibia + tarsus 255-400 wm long;
ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.97—1.12; translucent pores
visible on hind coxae and tibiae. Inner edges of
ostioles lightly sclerotized. Circulus quadrate and
of moderate size, width 85-130 zm. Cisanal setae
shorter than anal ring setae. Anal lobe cerarii each
situated on a moderately sized, well-sclerotized
area.

Venter. Multilocular disc pores usually con-
fined to median areas, rarely | or 2 pores present
on margins of 1 or 2 posterior abdominal seg-
ments; present around vulva, in single or double
rows across posterior edges of abdominal seg-
ments III-VI (sometimes also 1 or 2 pores also on
segment IT) and anterior edge of abdominal seg-
ment VI or VII (sometimes 1 or 2 pores also
present on anterior border of segment V), and a
variable number of pores scattered over thorax.
Trilocular pores sparsely but evenly distributed.
Oral collar tubular ducts of 2 sizes; smaller ducts
in rows across median areas of abdominal seg-
ments VI or VII; larger ducts in rows across
abdominal segments II-VII, scattered over
median areas of thorax and in marginal groups on
all head, thoracic and abdominal segments; up to
4 ducts sometimes present adjacent to each
postocular cerarius. Simple pores about half the
size of the trilocular pores, sparsely scattered over
entire venter.

Dorsum. Multilocular disc pores absent. Tri-
locular pores moderately numerous and aggreg-
ated around bases of enlarged setae. Tubular
ducts, considerably larger than the larger ducts on

21

the venter and without apparent rims, sometimes
present singly next to some abdominal cerarii.
Simple pores as for venter but very sparse. Dorsal
setae of various sizes, conical and without flagell-
ate apices, larger setae slightly smaller than the
cerarian setae and each with 1-3 trilocular pores
associated with their bases giving the appearance
of dorsal cerarii; these enlarged setae present on
all body segments except the last abdominal seg-
ment, generally occurring singly, but sometimes
up to 4 pairs present, length of longest conical seta
on abdominal segments VI or VII 15-20 um.

MATERIAL EXAMINED

21 @ (including type material listed below
(BMNH, USNM, ZIL).

Planococcus dorsospinosus Ezzat & McConnell.
Holotype 9, China (intercepted at San Francisco,
U.S.A.): on Pueraria hirsuta (kudzu vine),
20.i.1947 (F. L. Blane); 1 paratype 9, same
locality, on Colocasia esculentum, 10.1.1947 (F.
M. Thompson) (both USNM).

Planococcus myrsinephilus Borchsenius. | para-
type 2, China: Yunnan Province, near Siakwan,
on leaves and small branches of Myrsine africana,
18.iv.1957 (N. Borchsenius) (ZIL).

Planococcus sinensis Borchsenius. 1 paratype 9,
China: Yunnan Province, near Siakwan,
17.iv.1957 (N. Borchsenius) (ZIL).

DisTRIBUTION. Palaearctic Region: Japan (inter-
cepted in U.S.A.). Oriental Region: China,
Hongkong, India, Taiwan (intercepted in
U.S.A.), Thailand (intercepted in U.S.A.). Aus-
tro-oriental Region: Philippines (intercepted in
U.S.A.).

HostpLants. Aceraceae: Acer sp. (branches).
Araceae: Colocasia esculentum. Dioscoreaceae:
Dioscorea sp. (root). Euphorbiaceae: Euphorbia
longan. Leguminosae: Pueraria _ hirsuta.
Meliaceae: Lansium domesticum (fruit). Myr-
sinaceae: Myrsine africana (leaves and small
branches). Myrtaceae: Eugenia sp. (fruit),
Psidium guajava. Punicaceae: Punica granatum
(fruit). Rubiaceae: Coffea canephora (roots).
Also recorded from Litchi sp. (Sapindaceae) by
Ezzat & McConnell (1956) and as P. sinensis on
the leaves, branches and stems of Ficus gibbosa,
Morus sp. (Moraceae), Buddleia officinalis (Bud-
dleiaceae), Rhus sp. (Anacardaceae) and
Daphniphyllum sp. (Daphniphyllaceae) by
Borchsenius (1962).

REMARKS. This species is very similar to P.
dioscoreae, apparently differing only in having
relatively smaller and stouter enlarged dorsal
setae and by having no more than 4 tubular ducts

JENNIFER M. COX

22

- EE

Fig. 8 Planococcus dorsospinosus Ezzat & McConnell.

MEALYBUG GENUS PLANOCOCCUS

Fig. 9 Planococcus dubius Cox.

23

24

adjacent to each preocular cerarius. The speci-
mens from Dioscorea and Coffea robusta roots
listed above constitute the records from India.
These specimens are somewhat intermediate in
appearance between P. dorsospinosus and P.
dioscoreae — more material from a wide area will
be necessary to completely resolve this complex.
Both of these species may be distinguished from
P. litchi by having the enlarged dorsal setae asso-
ciated into no more than 4 pairs. P. dorsospinosus
appears to be somewhat variable, with larger
specimens having wider bands of multilocular disc
pores across the abdominal segments and more of
these pores on the thorax. The apparent length of
the dorsal setae is somewhat dependent on the
preparation of the specimens onto microscope
slides, as the flagellate tips are easily broken off.
Borchsenius (1962) separated P. sinensis from P.
dorsospinosus and P. dioscoreae by the lack of
sclerotization of the anal lobes in the latter 2 spe-
cies, but this character is also very dependent on
the microscopic preparation

ECONOMIC STATUS. This species is frequently inter-
cepted on produce in the U.S.A.

Planococcus dubius Cox
(Fig. 9)

Planococcus dubius Cox, 1987: 75. Holotype o,
NEW ZEALAND: on Dracophyllum lati-
folium (NZAC) [examined].

ADULT FEMALE. Mounted specimens oval, length
2.4-3.2 mm, width 1.5—1.8 mm. Margin of body
seldom with a complete series of 18 pairs of
cerarii, 12-18 pairs, each cerarius with 1-4 conical
setae on head and thorax, and 1 or 2 conical setae
on the abdomen, occasionally a single flagellate
auxiliary seta associated with one abdominal
cerarius. Legs elongate; hind trochanter + femur
320-390 um long, hind tibia + tarsus 310-360 um
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.93-1.00; translucent pores
apparent on hind coxae and tibiae. Inner edges of
ostioles moderately sclerotized. Circulus, if
apparent, small and quadrate, width 40-50 um.
Cisanal setae shorter than anal ring setae. Anal
lobe cerarii each situated on a moderately sized,
well-sclerotised area.

Venter. Multilocular disc pores present around
vulva, in single rows across posterior edges of
median areas of abdominal segments V or VI, a
few present on anterior edges of median areas of
segments V-VII, and scattered over head, thorax
and margins of entire body. Trilocular pores mod-

JENNIFER M. COX

erately numerous and evenly distributed. Oral
collar tubular ducts of 2 sizes, smaller ducts pre-
sent sparsely in rows across median areas of
abdominal segments IV-VII, larger ducts moder-
ately numerous around margins of entire venter.
Simple pores about half the size of the trilocular
pores, sparsely scattered over entire venter. Setae
somewhat long and stout.

Dorsum. Multilocular disc pores numerous
over entire surface. Tubular ducts, the same size
as the larger ducts on the venter and without
aparent rims, sparse to moderately numerous
over entire dorsum, forming rows across most
body segments. Trilocular pores as for venter.
Simple pores about a quarter of the size of the
trilocular pores, sparsely scattered over entire
dorsum. Setae generally long and with enlarged
bases but almost conical on median areas of
abdominal segments VI or VII, length of longest
of these setae 23-30 um.

MATERIAL EXAMINED

10 2 (including type material listed) (BMNH,
NZAC).

Holotype 2, New Zealand: Auckland, Waitakere
Range, on Dracophyllum latifolium, 16.xi.1973
(J.A. de Boer) (NZAC). Paratypes, 2 2, same
data as holotype (BMNH, NZAC); 1 9, New
Zealand: Lake Waikaremoana, at base of leaves
of Dracophyllum sp., 4.iii.1983 (J. M. Cox)
(BMNH, NZAC).

DIsTRIBUTION. New Zealand only.
HostTPLants. Epacridaceae: Dracophyllum sp..,
D. latifolium.

Remarks. P. dubius was placed in Planococcus by
Cox (1987), despite usually having fewer than 18
pairs of cerarili, because most specimens have
three pairs of cerarii anterior to the eyes and,
taken as a group, the type specimens show cerarii
in all the 18 possible positions on the body.

This species is very variable in its numbers of
cerarii and multilocular disc pores. Moreover, the
non-type material examined, collected in a dif-
ferent part of New Zealand from the type
material, apparently lack circuli— these specimens
may prove to be a different species when more
material has been collected and examined.

The affinities of this species with other mem-
bers of Planococcus are difficult to determine.
The presence of flagellate auxiliary setae in one
abdominal cerarius in some specimens suggests
that this species may actually be closer to
Planococcoides than to Planococcus. The only
other species of Planococcus with dorsal multi-
locular disc pores is the African P. epulus, but the
latter species has more flagellate dorsal setae on

MEALYBUG GENUS PLANOCOCCUS

abdominal segments VI VII and each cerarius is
distinct and with 2 conical setae. P. dubius may be
most closely related to the other species of
Planococcus found in New Zealand, P. mali
(although this species is suspected here of orig-
inating in Japan), as some cerarii are indistinct,
the dorsal setae are stout and some of the cephalic
and thoracic cerarii have more than 2 conical
setae. However, it is also close to the New Zea-
land species placed by Cox (1987) in Paracoccus,
all of which have oral rim tubular ducts and,
except for P. zealandicus (Ezzat & McConnell),
substantially fewer than 18 pairs of cerarii. The
presence of oral rim tubular ducts does not, in
itself, exclude P. zealandicus from Planococcus,
but it was placed in Paracoccus by Cox (1987),
despite sometimes having 18 pairs of cerarii, as it
is clearly congeneric with some of the other New
Zealand species placed in Paracoccus such as P.
drimydis (Brittin) and P. glaucus (Maskell).
Eventually, when the generic concepts of mealy-
bugs are better understood, these New Zealand
species may require the erection of a new genus.

Planococcus epulus De Lotto
(Fig. 10)

Planococcus epulus De Lotto, 1964: 375. Holo-
type 9 Kenya: on Prterolobium lacerans
(BMNH) [examined].

ADULT FEMALE. Mounted specimen elongate-
oval, length 1.7 mm, width 1.1 mm. Margin of
body with complete series of 18 pairs of cerarii,
each cerarius with 2 (rarely 1 on thorax) moder-
ately stout conical setae. Legs elongate; hind
trochanter + femur 315 um long, hind tibia +
tarsus 350 um long, ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 1.11; translu-
cent pores present on hind tibiae but not apparent
on hind coxae. Inner edges of ostioles not noticea-
bly sclerotized. Circulus quadrate, width 145 um.
Cisanal setae shorter than anal ring setae. Anal
lobe cerarii each situated on a small, moderately
sclerotised area.

Venter. Multilocular disc pores moderately
numerous around vulva, present in rows across
anterior and posterior edges of abdominal seg-
ments II-VII, in small marginal groups on seg-
ments III-VII and scattered over median areas of
venter. Trilocular pores moderately numerous
and evenly distributed. Oral collar tubular ducts
of 2 sizes; smaller ducts present sparsely in rows
across median areas of abdominal segments VI or
VII; larger ducts sparsely scattered over median
areas of head and thorax, present in rows across
median areas of abdominal segments, and in mar-

25

ginal groups around entire venter. Simple pores
minute, sparsely but evenly distributed.

Dorsum. Multilocular disc pores sparsely scat-
tered over thorax and abdomen. Tubular ducts of
same form and size as the larger ducts on the
venter, moderately numerous over entire dorsum.
Trilocular pores and simple pores as for venter.
Setae long and flagellate, longest seta on abdomi-
nal segments VI or VII about 45 wm long.

MATERIAL EXAMINED

Known from holotype 2 only, Kenya: Nairobi,
7.1.1957, on Pterolobium lacerans (G. De Lotto)
(BMNH).

DistriBuTION. Afrotropical Region: Kenya.
HostTPLanT. Leguminosae: Pterolobium lacerans.

REMARKS. The only other known species of
Planococcus with dorsal multilocular disc pores is
P. dubius from New Zealand (see under Remarks
for this species). In all other characters, P. epulus
appears to be a member of the P. citri-group.

Planococcus ficus (Signoret)
(Fig. 11)

[Coccus vitis L.; Nedzel’skii, 1869: 19. Mis-
identification. |

Dactylopius ficus Signoret, 1875: 315. Syntypes
©, France: on edible fig (probably lost).

[Dactylopius vitis; Lichtenstein, 1870: L. Misiden-
tification. ]

Dactylopius subterraneus Hempel, 1901b: 388.
Syntypes 2, ARGENTINA: on roots of cultivated
grapes (VCCB) [1 syntype examined by D. J.
Williams]. Syn. n.

Pseudococcus ficus (Signoret) Fernald, 1903: 101.

Pseudococcus subterraneus (Hempel) Fernald,
1903: 110.

[Pseudococcus vitis (Niedielski) Fernald, 1903:
112. Misidentification. |

Pseudococcus citrioides Ferris, 1922: 203. Holo-
type 2, Lisya: on ? (UCD) [examined]. [Syn-
onymized by Cox & Ben-Dov, 1986. ]

Planococcus citrioides (Ferris) Ferris, 1950: 164.

Planococcus ficus (Signoret) Ezzat & McConnell,
1956: 79.

[Planococcus vitis (Niedielski) Ezzat & McCon-
nell, 1956: 103. Misidentification. ]

ADULT FEMALE. Mounted specimens oval, length
1.4-3.2 mm, width 0.8-2.2 mm. Margin of body
with complete series of 18 pairs of cerarii, each
cerarius with 2 conical setae, more slender
towards the anterior of the body. Legs elongate;
hind trochanter + femur 235-355 um long, hind

26

|

Fig. 10 Planococcus epulus De Lotto.

MEALYBUG GENUS PLANOCOCCUS

Fig. 11

Planococcus ficus (Signoret).

2

28

tibia + tarsus 275-395 um long, ratio of lengths of
hind tibia + tarsus to hind trochanter + femur
1.01-1.17; translucent pores apparent on hind
coxae and tibiae, and often also on hind femora.
Inner edges of ostioles moderately sclerotized.
Circulus quadrate, width 115-150 um. Cisanal
setae shorter than anal ring setae. Anal lobe
cerarii each situated on a small, moderately scle-
rotized area.

Venter. Multilocular disc pores present around
vulva; in single rows across posterior edges of
abdominal segments III-VII or II-VII and across
anterior edges of segments V-VII; in marginal
groups on posterior abdominal segments, usually
as far forward as segment II; often present in
groups of up to 12 pores behind each front coxa;
and sparsely scattered over median areas of the
thorax. Trilocular pores moderately numerous
and evenly distributed. Oral collar tubular ducts
of 2 sizes; smaller ducts present sparsely in rows
across median areas of abdominal segments
V-VII; larger ducts present sparsely in rows across
median areas of abdominal segments II-VII, in
marginal groups on segments II-VIII, scattered
over median area of thorax, a single pore some-
times present on head, and up to 3 pores some-
times present on each margin of the thoracic
segments. Simple pores slightly smaller to slightly
larger than the trilocular pores, sparsely but
evenly distributed.

Dorsum. Multilocular disc pores absent. Tubu-
lar ducts, larger than those on the venter and
frequently with distinct rims, usually occurring
singly (rarely in pairs) adjacent to some or most
cerarii. Trilocular pores as for venter. Simple
pores of two sizes; larger pores, if apparent, larger
than the trilocular pores, present in groups of 1-3
pores on mid-line of some thoracic and abdominal
segments; smaller pores minute, scattered spar-
sely over entire dorsum. Setae long and flagellate,
longest seta on abdominal segment VI or VII
23-50 um long.

MATERIAL EXAMINED

About 50 @ (including type material listed)
(BMNH, MNNH, UCD, USNM, VCI).
Pseudococcus citrioides Ferris. Holotype 9,
Libya: Cyrenaica Guarsia, on ?, 4.iv.1922 (F. Sil-
vestri) (UCD); 2 paratype 2, same data as holo-
type (UCD).

DisTRIBUTION. Palaearctic Region: Cyprus,
Egypt, France, Greece, Iran, Iraq, Israel, Italy,
Lebanon, Libya, Saudi Arabia, Spain, Tunisia,
Turkey. Afrotropical Region: South Africa. Ori-
ental Region: Pakistan. Neotropical Region:
Argentina.

JENNIFER M. COX

HostpLants. Anacardaceae: Mangifera indica.
Lauraceae: Persea americana. Leguminosae:
Dichrostachys glomerata, Prosopis fareta,
Tephrosia purpurea. Moraceae: Ficus carica. Pal-
mae: Phoenix dactylifera. Punicaceae: Punica gra-
natum. Rhamnaceae: Zizyphus spina-christi.
Rosaceae: Malus pumila. Salicaceae: Salix sp.
Styraceae: Styrax officinale. Vitidaceae: Vitis vin-
ifera. Most of the specimens examined were from
grapevines or, less commonly, figs.

Remarks. P. ficus was frequently misidentified as
P. citri before the works of Ezzat & McConnell
(1956) and De Lotto (1975), and most records of
P. citri from grapevines should be referred to P.
ficus. As discussed in detail by De Lotto (1975)
and Cox & Ben-Dov (1986), the binomen Plan-
coccus Vitis is invalid, and records under this name
should be referred to P. ficus.

P. ficus is very similar to P. halli from the
Afrotropical Region and the West Indies, where it
is frequently found on yams. The distinction of
these two species is discussed under the Remarks
on P. halli. One of the most distinctive characters
of P. ficus, the presence of translucent pores on
the hind femora, is not apparent in many speci-
mens. Cox & Wetton (in press), who reared this
species, found that its expression was dependent
on temperature, translucent pores being present
on the hind femora of most specimens reared at
high temperatures, but absent from most speci-
mens reared at low temperatures

Dr D. J. Williams, C.A.B. International
Institute of Entomology, examined and sketched
a syntype of Dactylopius subterraneus Hempel
during a recent visit to Brazil, and concluded that
it was P. ficus. According to his notes and sketch,
the specimen has elongate conical setae in the
cephalic cerarii and 5 and 8 multilocular disc pores
behind the respective front coxae. The original
description states that the female is gall-forming,
but it is more likely that the mealybug was merely
occupying a vacated gall formed by some other
organism.

Economic status. P. ficus is the common meal-
ybug on grapevines throughout the Mediterra-
nean Basin and now also occurs on this host in
Argentina, Pakistan and South Africa. It will
probably continue to spread to other areas where
grapevines are grown.

Rosen & DeBach (1977) noted that the parasite
fauna supported by the vine mealybug in Israel is
significantly different from that of the citrus
mealybug.

MEALYBUG GENUS PLANOCOCCUS
Planococcus flagellatus De Lotto
(Fig. 12)

Planococcus flagellatus De Lotto, 1961: 220.
Holotype 2, UGANDa: on roots of Vernonia
auriculifera (BMNH) [examined].

ADULT FEMALE. Mounted specimens oval, length
1.7-3.2 mm, width 1.2-2.2 mm. Margin of body
with complete series of 18 pairs of cerarii, each
cerarius with 2 slender, often very elongate, coni-
cal setae. Legs elongate; hind trochanter + femur
255-335 um long, hind tibia + tarsus 285-370 um
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.93-1.12; translucent pores
apparent on hind coxae and tibiae. Inner edges of
ostioles lightly sclerotized. Circulus variable in
size, small and round to larger and quadrate,
width 15-160 um. Cisanal setae variable, from
shorter than (including type series) to longer than,
anal ring setae. Anal lobe cerarii situated on
small, moderately sclerotized areas.

Venter. Multilocular disc pores present around
vulva, in single or double rows across posterior
edges of median areas of abdominal segments
I-VII or II-VII (occasionally absent from seg-
ments I-III), usually a few present on anterior
edges of segments III-VII, present in small groups
on margins of abdominal segments II-VII or III-
VII (rarely reduced to a single pore on each side of
each segment) and usually scattered over median
areas of the thorax. Trilocular pores moderately
numerous and evenly distributed. Oral collar
tubular ducts of 2 sizes; smaller ducts present spar-
sely in rows across median areas of abdominal
segments II-VII; larger ducts present in marginal
groups on head and all thoracic and abdominal
segments, and scattered over median area of
thorax. Simple pores slightly smaller than the tri-
locular pores, sparsely but evenly distributed.

Dorsum. Multilocular disc pores absent. Tubu-
lar ducts, larger than the larger ducts on the venter
and without apparent rims, usually present singly
adjacent to some abdominal cerarii. Trilocular
pores as for venter. Simple pores about a third of
the size of the trilocular pores, present in small
groups on mid-line of thoracic segments. Setae
long and flagellate, 1 or 2 setae sometimes bifur-
cate, longest seta on abdominal segments VI or
VII 40-100 um long.

MATERIAL EXAMINED

14 Q (including type material listed below) (all
BMNH).

Holotype 2 , Uganda: Sebei, on roots of Vernonia
auriculifera, 18.11.1957 (D. N. McNutt). Para-
types, 5 2, same data as holotype.

29

DistTRIBUTION. Afrotropical Region: Mozambi-
que, Nigeria, Sudan, Uganda.

HosTPLants. Occurring on the roots. Anacar-
daceae: Anacardium occidentale (cashew).
Asteraceae: Vernonia auriculifera. Leguminosae:
Glycine max (soya bean). Sterculiaceae: The-
obroma cacao. Also recorded from the roots of
Cassia petersiana (Leguminosae) by De Lotto
(1964).

RemarKS. This species is clearly a member of the
P. citri-group by virtue of its distribution of multi-
locular disc pores and tubular ducts. It is similar to
P. ficus and P. halli which may also have long
dorsal setae, but differs from both of these species
by usually having numerous marginal tubular
ducts on the head and thorax. It is most similar to
P. subterraneus, known only from the holotype,
which lacks tubular ducts on the head and has
many bifurcate dorsal setae.

Planococcus furcisetosus Mamet
(Fig. 13)

Planococcus furcisetosus Mamet, 1959: 404. Syn-
types 2, MapaGascakr: on ebenier (MNNH) [2
syntypes examined].

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.6-1.8 mm, width 1.2-1.3 mm.
Margin of body with a complete series of 18 pairs
of cerarii, each with 2 conical setae. Legs stout;
hind trochanter + femur 235-260 um long, hind
tibia + tarsus 220-240 um long; ratio of lengths of
hind tibia + tarsus to hind trochanter + femur
about 0.92; translucent pores present on hind
coxae and tibiae. Inner edges of ostioles well scle-
rotized. Circulus small and round, width 55-70
uum. Cisanal setae shorter than anal lobe setae.
Anal lobe cerarii each situated on a small, moder-
ately sclerotized area.

Venter. Multilocular disc pores confined to
median areas of abdomen, present around vulva
and in rows across posterior edges of median areas
of abdominal segments III-VI. Trilocular pores
moderately numerous and evenly distributed.
Oral collar tubular ducts apparently of 1 size,
present in rows across median areas of abdominal
segments III-VII and in marginal groups on seg-
ments VII-IX. Simple pores not apparent on spec-
imens examined.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores moderately numer-
ous and evenly distributed. Simple pores slightly
larger than the trilocular pores, present in small

JENNIFER M. COX

ee)
Ar

«
‘
in

7 oR

Fig. 12 Planococcus flagellatus De Lotto.

Sil

MEALYBUG GENUS PLANOCOCCUS

2005, 0

40%

eo

a

Fig. 13. Planococcus furcisetosus Mamet.

32

groups on the mid-line of the anterior abdominal
segments. Setae short and stout, most furcate,
longest seta of abdominal segments VI or VII
22-25 um long.

MATERIAL EXAMINED

2 syntype 9, Madagascar: Maroantsetra,
Ambodivoany, on ebenier, iii.1952 (R. Paulian)
(MNNH).

DIsTRIBUTION. Malagasian Region: Madagascar.
HostpLant. Ebenaceae: Diospyros sp.

RemMARKS. The furcate dorsal setae distinguish P.
furcisetosus from all other known species of
Planococcus.

Planococcus halli Ezzat & McConnell
(Fig. 14)

Planococcus halli Ezzat & McConnell, 1956: 81.
Holotype 9, Sr Kitts: on Dioscorea sp.
(USMN) [examined].

ADULT FEMALE. Mounted specimens oval, length
1.5-3.4 mm, width 0.9-2.4 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
with 2 conical setae which are more slender and
flagellate towards the anterior end of the body.
Legs elongate; hind trochanter + femur 220-335
um long, hind tibia + tarsus 275-371 um long,
ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.03—1.24; translucent pores
apparent on hind coxae and tibiae. Inner edges of
ostioles lightly sclerotized. Circulus quadrate,
width 55-180 um. Cisanal setae shorter than anal
lobe setae. Anal lobe cerarii each situated on a
moderately sized, lightly sclerotized area.
Venter. Multilocular disc pores situated around
vulva and in rows (usually single, except in very
large specimens where they may be double) across
posterior edges of median areas of abdominal seg-
ments II-VI or III-VII, in single rows across ante-
rior edges of median areas of abdominal segments
IV-VII, in marginal groups on either side of seg-
ments III-VII or IV-VII (rarely 1 or 2 pores pre-
sent on margins of segment II), and a few pores
sometimes scattered over median areas of head
and thorax. Trilocular pores numerous and evenly
distributed. Oral collar tubular ducts of 2 sizes;
smaller ducts present sparsely in rows across
median areas of abdominal segments VI or VII;
larger ducts present in rows across median areas of
segments II-VII, in marginal groups on abdominal
segments I-IX, scattered over median areas of
thorax, 1-3 ducts sometimes present on head and
1-6 ducts sometimes present on each side of each

JENNIFER M. COX

thoracic segment. S'mple pores slightly smaller
than the trilocular pores, scattered over entire
venter.

Dorsum. Multilocular disc pores absent. Tubu-
lar ducts, if present, larger than those on the ven-
ter and without apparent rims, and present singly
(rarely in 2s or 3s) adjacent to some abdominal
cerarii. Trilocular pores as for venter. Simple
pores of 2 sizes; larger pores slightly larger than
the trilocular pores, present in small groups on
mid-line of each abdominal segment; smaller
pores minute, scattered over entire dorsum. Setae
long and flagellate, length of longest seta of
abdominal segments VI or VII 25—45 um.

MATERIAL EXAMINED

About 100 Q (including holotype) (BMNH,
USNM).

Holotype Q, St Kitts (intercepted at Boston, Mas-
sachusetts): on Dioscorea sp., yam, 12.iii.1949
(Hodson Hardy) (USNM). Originally described
from the holotype only.

DisTRIBUTION. Afrotropical Region: Cameroons,
Ethiopia, Gabon, Ghana, Liberia, Nigeria, South
Africa, Sudan. Neotropical Region: Barbados,
Bermuda, Brazil, Guyana, Haiti, St. Kitts,
Trinidad.

HostTpLants. Celestraceae: Gymnosporia spin-
osa. Dioscoreaceae: Dioscorea alata, D. rotun-
data. Euphorbiaceae: Manihot esculentum.
Leguminosae: Arachis hypogaea (groundnut).
Poaceae: Saccharum officinarum. Rubiaceae:
Coffea canephora. Most commonly encountered
on yams (Dioscorea spp.).

Remarks. P. halli is very similar to P. ficus, and
some specimens may be impossible to identify by
the characters given in the key. Cox & Wetton (in
press) carried out multivariate analyses of speci-
mens of this group from yams, pomegranates, figs
and grapevines. The results showed that the speci-
mens from yams were differentiated almost
entirely from those from figs, pomegranates and
grapevines, and that specimens from yams from
the Ethiopian Region and from the West Indies
were intermingled. That this differentiation was
not merely a host effect was demonstrated by
rearing material from each source on potato
tubers. Cox & Wetton (in press) concluded that,
as two names are available, the two groups should
be regarded for the present as separate species.
The two species differ by the frequent presence
of a group of multilocular disc pores behind the
front coxae in P. ficus (never present in P. halli),
the frequent occurrence of translucent pores on
the hind femora in P. ficus (never present in P.
halli), multilocular disc pores usually extending

MEALYBUG GENUS PLANOCOCCUS

Fig. 14

Planococcus halli Ezzat & McConnell.

33

34

further forwards on the marginal than on the
median areas of the body (the reverse situation
usually occurring in P. halli), and large simple
pores seldom apparent on the midline of the
abdominal segments in P. ficus (always apparent
in P. halli).

Specimens from the Sudan and Ethiopia differ
from the usual forms of both P. ficus and P. halli.
More work is needed on this complex, preferably
carried out locally where live material would be
available for experimentation.

Economic status. P. halli is frequently inter-
cepted in the U.S.A. and the U.K. on yam tubers
from Nigeria and the West Indies.

Planococcus hospitus De Lotto
(Fig. 15)

Planococus hospitus De Lotto, 1961 : 221. Holo-
type 92, UGanpa: on Eulophia (BMNH)
[examined].

ADULT FEMALE. Mounted specimen rotund,
length 1.8 mm, width 1.4 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
with 2 conical setae which are stout on abdominal
segments but more slender on head. Legs stout;
hind trochanter + femur 220 um long, hind tibia +
tarsus 200 um long; ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 0.91; translu-
cent pores not apparant on hind legs. Inner edges
of ostioles well sclerotized. Circulus small and
oval, width 65 um. Cisanal setae shorter than anal
lobe setae. Anal lobe cerarii each situated on a
small, lightly sclerotized area.

Venter. Multilocular disc pores confined to
median areas of abdomen, present around vulva
and in single rows across posterior edges of
median areas of abdominal segments V or VI.
Trilocular pores moderately numerous and evenly
distributed. Oral collar tubular ducts of 1 size,
confined to median areas of abdomen, present in
single rows across median areas of abdominal seg-
ments IV-VII. Simple pores about the same size as
the multilocular disc pores, lightly sclerotized,
and scattered over entire venter. Setae on median
areas long, fine and flagellate, but those on mar-
gins moderately long and fine, and with indis-
tinctly knobbed tips.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores numerous and
evenly distributed. Simple pores of 2 sizes, pores
about twice the size of the trilocular pores scat-
tered over entire dorsum, larger pores in groups
on mid-line of thoracic and abdominal segments.

JENNIFER M. COX

Setae moderately long and fine, and with indis-
tinctly knobbed tips; length of longest seta on
abdominal segments VI or VII about 30 um.

MATERIAL EXAMINED

Holotype 9, Uganda: Kampala, on tubers of
Eulophia sp., 18.x.1956 (A. G. P. Michelmore)
(BMNH).

DistriBuTION. Afrotropical Region: Uganda.
HostpLantT. Orchidaceae: Eulophia sp.

REMARKS. This species was originally described
from, and is still only known from, a single
specimen.

P. hospitus is similar to the other 4 species of
Planococcus which occur on orchids, P. hosyni
and P. orchidi from the Afrotropical Region, and
P. dendrobii and P. philippinensis from the Orien-
tal Region. All have low numbers of multilocular
disc pores and tubular ducts, and have knobbed
(although often indistinctly) dorsal setae. P. hos-
pitus and P. hosyni may be distinguished from the
other species by their lack of marginal tubular
ducts and from each other by the presence of
translucent pores on the hind tibiae and tarsi in P.
hosyni and by the very large ventral simple pores
in P. hospitus.

Planococcus hosyni Ezzat & McConnell
(Fig. 16)

Planococcus hosyni Ezzat McConnell, 1956: 83.
Holotype 9, ZimBaBwe: on Aerangis katschii
(USNM) [examined].

ADULT FEMALE. Mounted specimens oval to
rotund, length 1.3-2.1 mm, width 0.8-1.8 mm.
Margin of body with a complete series of 18 pairs
of cerarii, all with moderately stout conical setae.
Legs stout; hind trochanter + femur 180-210 um
long, hind tibia + tarsus 195-210 um long, ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 1.0—1.05; translucent pores present on hind
coxae and tibiae. Inner edges of ostioles moder-
ately sclerotized. Circulus small and round, width
47-57 um. Cisanal setae shorter than anal ring
setae. Anal lobes not noticeably sclerotized.
Venter. Multilocular disc pores confined to
median areas of abdomen, present around vulva,
in a single row across posterior border of median
area of abdominal segment VI, and 1-3 pores
present on segments IV or V. Trilocular pores
moderately numerous and evenly distributed.
Oral collar tubular ducts of 1 size, confined to
median areas of abdomen, occurring sparsely in
rows across median areas of abdominal segments

MEALYBUG GENUS PLANOCOCCUS

Fig. 15 Planococcus hospitus De Lotto.

35

36

JENNIFER M. COX

ONES zi Oe IE ay SS

rae
voi
\
/
ve
—

7

5
.

u
4

See
r mile
s xe Wain
ba Rts A

| co

SY

Fig. 16 Planococcus hosyni Ezzat & McConnell.

MEALYBUG GENUS PLANOCOCCUS

IV-VII. Simple pores about the same size as the
trilocular pores, sparsely but evenly distributed.
Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores of 2 sizes, smaller pores smaller than the
trilocular pores and sparsely scattered over entire
dorsum, larger simple pores slightly larger than
the trilocular pores, present in groups on mid-line
of thoracic and abdominal segments. Setae mod-
erately long and stout, flagellate but with indis-
tinctly knobbed tips, length of longest seta on
abdominal segments VI or VI 23-30 um.

MATERIAL EXAMINED

5 9 (including holotype) (BMNH, USNM).
Holotype 9, Zimbabwe (Rhodesia): on Aerangis
katschii, 2.vii.1953 (D. Kamper) (USNM).

DistRIiBUTION. Afrotropical
Africa, Zimbabwe.

HostpLant. Orchidaceae: Aerangis katschit.

Region: South

REMARKS. This species was originally described
from the holotype only. This specimen is a very
young adult female — although its body shape is
oval, the mature female would probably have
been rotund, as in the additional material exam-
| ined. Other specimens examined from South
_ Africa also on Orchidaceae have the dorsal setae
slightly more distinctly knobbed than those of the
| holotype, but this characteristic can be observed
_ in the latter specimen under high magnification
P. hosyni is most similar to P. hospitus, also
_ from the Afrotropical Region, in completely lack-
ing marginal tubular ducts and in having flagell-
ate, yet knobbed, dorsal setae, but can be
| distinguished by the very large ventral simple
| pores which are present in P. hospitus. The three
_ other known species with knobbed dorsal setae,
| P. orchidi from the Afrotropical Region, and P.
| dendrobii and P. philippinensis from the Oriental
| region, all recorded only from orchids, have mar-
' ginal tubular ducts on abdominal segments VI or
VII and much stouter dorsal setae.

| Planococcus japonicus sp. n.
(Fig. 17)

| [Planococcus azaleae (Tinsley) Ezzat & McCon-
nell, 1956: 63. Misidentification. |

ADULT FEMALE. Mounted specimens oval, length
1.5-3.9 mm, width 0.9-2.3 mm. Margin of body
| with a complete series of 18 pairs of cerarii,

cephalic and thoracic pairs each with 2-5 conical
| setae and abdominal pairs each with 2 conical

aT

setae. Legs elongate, hind femur + trochanter
240-315 um long, hind tibia + tarsus 275-340 um
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.03—1.17; translucent pores
present on hind coxae and tibiae. Inner edges of
ostioles lightly sclerotized. Circulus quadrate,
width 80-145 um. Cisanal setae shorter than anal
ring setae. Anal lobe cerarii each situated on a
small, moderately sclerotized area.

Venter. Multilocular disc pores situated around
vulva, in rows across posterior edges of median
areas of abdominal segments III-VII or IV-VII
and across anterior edges of segments V-VII, a
few pores present on margins of segments VI or
VII, up to7 pores situated on head and 5-16 pores
scattered over median areas of thorax. Oral collar
tubular ducts of 2 sizes; smaller ducts present in
rows across median areas of abdominal segments
II-VII; larger ducts present in rows across median
areas of segments [V-VI, in marginal groups on
segments I-VIII, in a larger marginal group on
each side of prothorax and a few ducts sometimes
present on head and margins of mesothorax. Tri-
locular pores evenly distributed. Simple pores
minute, scattered over venter.

Dorsum. Multilocular disc pores absent. Some
specimens (including the holotype) with 1 or 2
tubular ducts with distinct rims present on the
head. Trilocular pores and simple pores as for
venter. Setae short and stout, sometimes almost
conical or lanceolate, longest seta on abdominal
segments VI or VII 15-20 um long.

MATERIAL EXAMINED

Holotype 9, Japan: Fukuoka, Ryugezi, on Pyrus
sp., 6.vi.1968 (Hiroshi Kajita) (USNM).
Paratypes. Japan: 2 9, same data as holotype; 1
Q, (intercepted at Anchorage), on Eriobotrya
Japonica fruit, 8.vi.1971 (D. Husnik) (USNM); 1
Q, (intercepted at Seattle), on Eriobotrya jap-
onica fruit, 8.vi.1971 (R. F. Goodall) (USNM); 12
Q, (at National Arboretum, Washington, D.C.),
on Rhododendron indicum stem, 3.viii.1978 (S.
Nakahara & R. Brittingham) (BMNH, USNM).

DistTRIBUTION. Palaearctic Region: Japan.

HostTp.ants. Ericaceae: Rhododendron indicum.
Rosaceae: Eriobotrya japonica, Pyrus sp.

RemaRrkKS. This species has frequently been con-
fused with Crisicoccus azaleae Tinsley, and was
redescribed and illustrated as Planococcus azaleae
by Ezzat & McConnell (1956). The latter species
has fewer pairs of cerarii and has much more
flagellate dorsal setae than does P. japonicus (Fig.
18). Some of the material listed above was
labelled as Crisicoccus matsumotoi (Shiraiwa),
but this species has very long flagellate dorsal

38

Fig. 17 Planococcus japonicus sp. n.

JENNIFER M. COX

MEALYBUG GENUS PLANOCOCCUS

3 ES

Fig. 18

Crisicoccus azaleae (Tinsley)

59

40

Fig. 19 Crisicoccus matsumotoi (Shiraiwa).

JENNIFER M. COX

| Pseudococcus kenyaensis Betrem 1936:

MEALYBUG GENUS PLANOCOCCUS

setae (Fig. 19). It is quite probable that all three of
these species are related, as the distinction of
Crisicoccus from Planococcus is only by the loss of
cerarii in the former genus.

P. japonicus is very similar to P. mali which, at
present, is known only from New Zealand and
Australia. It differs from P. mali by having smaller
legs and a larger group of oral collar tubular ducts
on each side of the prothorax. The separation of
these two species is discussed in more detail under
P. mali.

Economic status. This species caused heavy
infestations on Rhododendron at the National
Arboretum, Washington D.C. in 1978.

| Planococcus kenyae (Le Pelley)

(Fig. 20)

Pseudococcus citri var. congoensis Newstead;
Mann, 1922: 629. Nomen nudum.

Pseudococcus kenyae Le Pelley, 1935: 185. Holo-
type 9, Kenya: on Coffea arabica (BMNH,
?missing) [not examined].

129.
Nomen nudum.

Planococcus kenyae (Le Pelley) Ferris, 1950: 164.

Planococcus subukiaensis De Lotto, 1954: 110.
Holotype @, Kenya: on Coffea arabica
(BMNH) [examined]. Syn. n.

_ ADULT FEMALE. Mounted specimens oval to
| rotund, length 1.4-2.7 mm, width 0.8-2.0 mm.
| Margin of body with a complete series of 18 pairs
| of cerarii, all with moderately stout conical setae.

Legs elongate; hind trochanter + femur 225-340

| um long, hind tibia + tarsus 250-355 um long,
| ratio of lengths of hind tibia + tarsus to hind

trochanter + femur 0.97—1.11; translucent pores
present on hind coxae and tibiae. Inner edges of

| ostioles moderately sclerotized. Circulus quadr-

ate and of moderate size, width 50-160 um. Cisa-
nal setae usually noticeably longer than anal ring
setae. Anal lobe cerarii each situated on a large,
pronounced, sclerotized area which is apparent
only in well-stained specimens.

Venter. Multilocular disc pores usually con-
fined to median areas of venter, present around
vulva and in single rows across posterior borders
of median areas of abdominal segments IV-VII, a
few pores sometimes present on anterior borders
of median areas of abdominal segments VI and/or

_ VII and on margins of segments V-VII. Trilocular
| pores moderately numerous and evenly dis-
' tributed. Oral collar tubular ducts of 2 sizes;
| smaller ducts occurring sparsely in rows across

41

median areas of abdominal segments VI or VII;
larger ducts present in rows across median areas of
all abdominal segments, usually sparsely scattered
over median areas of thorax, and often present
singly or in small marginal groups around entire
venter, a single duct often located adjacent to
each postocular cerarius. Simple pores about the
same size as the trilocular pores, sparsely but
evenly distributed.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores of 2 sizes, smaller size smaller than the
trilocular pores and sparsely scattered over entire
dorsum, larger size about twice the size of the
trilocular pores in groups on mid-line of abdomi-
nal segments I-V. Setae flagellate and moderately
short and stout, length of longest seta on abdomi-
nal segments VI or VI usually 15-26 um, rarely up
to 40 um.

MATERIAL EXAMINED

Over 100 @ (including type material listed)
(BMNH, MNNH).

Pseudococcus kenyae. Holotype 2 apparently
missing. Paratypes, 8 0, Kenya: Nairobi, Scott
Agri. Lab., on Coffea arabica, 1.iii.1935 (P. B.
Notley); 1 Q, same data except 23.vi.1933
(BMNH).

Planococcus subukiaensis. Holotype 2, Kenya:
Subukia, ex Coffea arabica, 8.vii.1933 (no collec-
tor) (BMNH). Paratypes, 5 9, same data as holo-
type (BMNH, USNM). The original description
of P. subukiaensis states that the host plant is
unknown, but it is given as Coffea arabica on the
slide labels and by De Lotto (1964).

DistTRIBUTION. Afrotropical Region: Central
African Republic, Ghana, Kenya, Nigeria,
Congo, Sudan, Tanzania, Togo, Uganda, Zaire,
Zimbabwe.

HostpLants. Anacardaceae: Lannea discola.
Costaceae: Costus sp. Euphorbiacae: Croton sp.
Leguminosae: /nga sp. Loranthaceae: Loranthus
sp. Malvaceae: Hibiscus gossypinus. Pas-
sifloraceae: Barteria fistulosa. Rubiaceae: Coffea
sp. C. arabica, Cuviera angloensis, Leptactinia sp.
Sterculiaceae: Theobroma cacao.

Remarks. P. kenyae is readily recognised by its
prominent anal lobes, long cisanal setae, multi-
locular disc pores largely restricted to the pos-
terior borders of the median areas of the
abdominal segments and relatively few marginal
tubular ducts. It is superficially similar to P.
lilacinus in having long cisanal setae and multi-
locular disc pores absent from the margins of the
venter, but it lacks the noticeably stout legs and
very long dorsal setae of P. lilacinus. It is also

42

JENNIFER M. COX

at he
eS
“Ia [9 . ae a 2 ee
eS —
Life ©)
so//a ° f)
“ of, fs Sie
< >
. is e a a ic

Fig. 20 Planococcus kenyae (Le Pelley).

MEALYBUG GENUS PLANOCOCCUS

similar to P. hosnyi which is known only from the
holotype, but differs by having relatively longer
cisanal setae, elongate legs, and tubular ducts pre-
sent on the margins of the body.

P. subukiaensis was described by De Lotto
(1954) and redescribed by Ezzat & McConnell
(1956) on the basis of a single series of specimens.
These were said to differ from P. kenyae by lack-
ing the dorsal sclerotized area around the anal
lobe cerarii, by having more tubular ducts and
multilocular disc pores on the abdomen and by
lacking tubular ducts anterior to the circulus.
Reexamination of the type-specimens during the
course of the present study shows these specimens
to have been over-macerated, with the result that
the dorsal sclerotization of the dorsal lobes is not
apparent. Furthermore there are, in fact, a few
tubular ducts present on the margins of the head
and thorax in all these specimens. The anal lobe
area is somewhat damaged in all the specimens,
but in one instance the cisanal seta is clearly elong-
ate as in normal P. kenyae. Studied in conjunction
with all the material listed below, the type-
material of P. subukiaenis shows no noticeable
deviation from the pattern of variation observed
for P. kenyae, and the species is consequently here
synonomized.

Mann (1922) published the name Pseudococcus
citri var. congoensis Newstead for some ant-
attended mealybugs found on Barteria fistulosa
and Cuviera angolensis in the Republic of the
Congo. He states that the species was apparently
still undescribed, but had been identified by Prof.
R. Newstead. Specimens which are undoubtedly
those seen by Newstead are in the BMNH and
have been identified here as P. kenyae.

| Economic status. P. kenyae is a major pest of
| coffee in Kenya. The history of its identification
| highlights one of the common problems encoun-
| tered in the implementation of biological control
| programmes. In 1923 a mealybug was found caus-
_ ing considerable damage to coffee in Kenya. As

biological control was known to be a sucessful way
of controlling coccoids, parasites were sought for
its control. The mealybug was initially identified
as the citrus mealybug P. citri for which parasites
were available in California. These were intro-
duced into Kenya and tested against the coffee
mealybug, but the results were unsuccessful. The
mealybug was then redetermined as P. lilacinus,
an Oriental species. Parasites were collected from
south-east Asia, taken to Africa, and tested
against the coffee mealybug, again without suc-
cess. Finally, 12 years after this mealybug became
a problem, it was recognized as being an

undescribed species and appropriate parasites

43

found for it in the neighbouring African countries
(Le Pelley, 1935; 1943a). It is interesting that the
specimens labelled as Pseudococcus citri var. con-
goensis by Newstead, discussed above, were col-
lected in 1919. To date, P. kenyae is known only
from the Afrotropical Region.

Planococcus kraunhiae (Kuwana)
(Fig. 21)

Dactylopius kraunhiae Kuwana, 1902: 55. Syn-
types 2, JAPAN: on Wisteria floribunda (as
‘Kraunhia floribunda’) (location of types not
known) [not examined].

Pseudococcus kraunhiae (Kuwana) Fernald,
1903: 104.

Planococcus kraunhiae (Kuwana) Ferris, 1950:
158; Ezzat McConnell, 1956: 87; McKenzie,
1967: 283.

Planococcus siakwanensis Borchsenius, 1962:
586. Holotype 2 , Cuina: on undetermined host
(ZIL) [not examined]. Syn. n.

ADULT FEMALE. Mounted specimens oval, length
2.0-3.3 mm, width 1.1-2.0 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
with 2 slender conical setae, slightly swollen at the
base, more slender towards the anterior end of the
body. Legs elongate; hind trochanter + femur
285-375 um long, hind tibia + tarsus 305-430 wm
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.06—1.20; translucent pores
present on hind coxae and tibiae. Inner edges of
ostioles lightly sclerotized. Circulus small and
quadrate, width 80-190 um. Cisanal setae shorter
than anal ring setae. Anal lobe cerarii each situ-
ated on a small, lightly sclerotized area which is
apparent only in well-stained specimens.

Venter. Multilocular disc pores present around
vulva, in single or double rows across posterior
edges of median areas of abdominal segments
II-VII, in single rows across anterior edges of
segments III-VII or IV-VII, 1-4 pores present on
each margin of segments V-VII, and a few pores
scattered over thorax. Trilocular pores moder-
ately numerous and evenly distributed. Oral col-
lar tubular ducts of 2 sizes; smaller size occurring
sparsely in rows across median areas of abdominal
segments III-VII; larger size in rows across
median areas of segments II-VI, sparsely scat-
tered over median areas of thorax and present in
small marginal groups on head and thorax and in
larger groups on margins of abdominal segments.
Simple pores about half the size of the trilocular
pores, very sparsely scattered over abdominal seg-
ments only.

Dorsum. Multilocular disc pores absent. Tri-
locular pores moderately numerous and generally

44

Fig. 21 Planococcus kraunhiae (Kuwana).

JENNIFER M. COX

MEALYBUG GENUS PLANOCOCCUS

evenly distributed, but 1 or 2 pores sometimes
associated with the bases of some of the larger
setae. Tubular ducts, larger than the larger size on
the venter and without apparent rims, variable in
number, present in groups of 2-5 ducts adjacent to
at least some abdominal cerarii, sometimes also
sparsely in rows across some abdominal segments.
Simple pores apparently absent. Setae with
flagellate tips and distinctly swollen bases, length
of longest seta on abdominal segments VI or VI
18-26 wm.

MATERIAL EXAMINED

19 © (including type material listed below)
(BMNH, ZIL, UCD, USNM).

Planococcus siakwanensis Borchsenius. 1 para-
type 2, China: Yunnan Province, nr Siakwan, on
branches of an undetermined shrub, 17.iv.1957
(Borchsenius) (ZIL).

DisTRIBUTION. Palearctic Region: Korea (inter-
cepted in U.S.A.), Japan. Oriental Region:
» China. Nearctic Region: U.S.A., California. Fre-
quently intercepted in the U.S.A. on fruit
imported from Japan.

HosteLants. Ebenaceae: Diospyros kaki.
Leguminosae: Wisteria sp. Rutaceae: Citrus sp.
Also recorded from Wisteria floribunda (as
‘Kraunhia floribunda’) (Kuwana, 1902), Ficus
carica (Moraceae), Plantanus orientalis (Plan-

tanaceae), Citrus noblis var. unshiu, Citrus para-
_ disi and Ilex sp. (Aquifoliaceae) (McKenzie,
1967). Some of these records may be based on
misidentifications.

The records by Ezzat & McConnell (1956) on
croton (Croton sp., Euphorbiaceae) from Jamaica
| (intercepted in U.S.A.) and Olea chrysophylla
(Oleaceae) from Eritrea, Ethiopia are regarded
_ here as being dubious. Specimens of the latter
| record have been located and are a species of

| Delottococcus.

REMARKS. This is apparently an eastern Palaearc-
tic species that has been introduced into the
U.S.A.

P. kraunhiae is characterised by having only a few
multilocular disc pores on the margins of the
abdomen, short, stout dorsal setae with swollen
bases and flagellate tips, and groups of 2-5 tubular
| ducts on the dorsum adjacent to most of the
abdominal cerarii. P. kraunhiae could be con-
fused with two other species that often have simi-
lar numbers and distribution of dorsal tubular
ducts, P. ficus and P. vovae. It differs from P. ficus
by having considerably fewer multilocular disc
| pores on the margins of the abdominal segments
and by having differently shaped dorsal setae
| (slender and flagellate in P. ficus), and from P.

45

vovae by having at least a few multilocular disc
pores on the margins of the abdominal segments
and by the same difference in the shape of the
dorsal setae as in P. ficus. The swollen-based dor-
sal setae may cause it to be confused with the
dorsospinosus-group, but these species never
have marginal multilocular disc pores, and have
the trilocular pores much more aggregated around
the bases of the dorsal setae.

The paratype of P. siakwanensis examined fits
within the observed variation of P. kraunhiae col-
lected from other parts of the world. The type
specimens of P. siakwanensis represent the only
material of P. kraunhiae known from China.

Planococcus lilacinus (Cockerell)
(Fig. 22)

Pseudococcus lilacinus Cockerell, 1905: 128. Syn-
types 9, PHILIPPINES: on cultivated orange
(USNM) [8 syntypes examined].

Pseudococcus tayabanus Cockerell, 1905: 129.
Syntypes 9, PHILIPPINES: on cultivated cacao
(USNM) [4 syntypes examined]. [Synonymised
by Morrison, 1920: 176.]

Dactylopius crotonis Green, 1906: 44. Nomen
nudum.

Dactylopius crotonis Green, 1911: 35. [As new
species.] Syntypes 2, Sri LANKA: on Castilloa
elastica (BMNH) [4 syntypes examined]. [Syn-
onymised by Morrison, 1920: 176].

Pseudococcus crotonis (Green) Sasscer, 1912: 86.

Pseudococcus deceptor Betrem, 1937: 54. Java:
on coffee roots (status of type material not
known). Syn. n.

Tylococcus mauritiensis Mamet, 1939: 579. Holo-
type 2, Mauritius: on fruits of Eugenia mes-
piloides (BMNH: ?missing) [not examined].
[Synonymized by Ezzat & McConnell, 1956:
89.]

Planococcus crotonis (Green) Ferris, 1950: 164.

Planococcus tayabanus (Cockerell) Ferris, 1950:
164.

Planococcus lilacinus (Cockerell) Ferris, 1950:
164.

[Planococcus citri (Risso) Ferris, 1954: 52.
Misidentification. ]

[Planococcus citri (Risso) McKenzie, 1967: 281;
illustration only. Misidentification. ]

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.2—3.1 mm, width 0.7-3.0 mm.
Margin of body with a complete series of 18 pairs
of cerarii, usually all with stout conical setae,
occasionally some of those on head with elongate,

Cox

JENNIFER M.

46

Fig. 22 Planococcus lilacinus (Cockerell).

MEALYBUG GENUS PLANOCOCCUS

slender setae. Legs stout; hind trochanter +
femur 210-315 wm long, hind tibia + tarsus 210-
275 um long, ratios of lengths of hind tibia +
tarsus to hind trochanter + femur 0.77-0.97;
translucent pores present on hind coxae and
tibiae. Inner edges of ostioles strongly sclerotized.
Circulus large and quadrate, width 105-200 um.
Cisanal setae noticeably longer than anal ring
setae. Anal lobe cerarii each situated on a moder-
ately sized, well-sclerotized area.
Venter. Multilocular disc pores occurring on
median areas only, present around vulva, in single
or double rows across posterior borders of median
areas of abdominal segments [V-VII and usually
in a single row across anterior edge of segment VII
(although the latter is sometimes reduced to a few
pores), and a few pores sometimes present on
anterior edges of median areas of abdominal seg-
ments V and VI. Trilocular pores moderately
numerous and evenly distributed. Oral collar
tubular ducts of 2 sizes; smaller ducts occurring
sparsely in rows across median areas of abdominal
segments III-VII; larger ducts sparsely scattered
over median areas of thorax and present in margi-
nal groups around entire venter, a few ducts also
present adjacent to each postocular cerarius. Sim-
ple pores about the same size as the trilocular
pores, sparsely but evenly distributed.

Dorsum. Multilocular disc pores and tubular
_ ducts absent. Trilocular pores as for venter. Sim-
| ple pores smaller than the trilocular pores, spar-
sely but evenly distributed. Setae very long, stout
/ and flagellate, length of longest seta on abdominal
segments VI or VII 50-140 um.

MATERIAL EXAMINED

| About 200 9 (including type material listed
| below) (BMNH, USNM).

| Pseudococcus lilacinus Cockerell. 8 syntype 9,
' Philippines: Lucan, Tayabas, on orange (cult.),
10.iv.1904 (Townsend) (USNM).

| Pseudococcus tayabanus Cockerell. 4 syntype °,
Philippines: Lucban, Tay[abas], 7 20.iv.1904
(Townsend) (USNM).

Tylococcus mauritiensis Mamet. 8 paratype Q,
' Mauritius: Corpes de Garde Mt, on Eugenia mes-
| piloides, 30.1.1938 (R. Mamet) (BMNH).

| Pseudococcus crotonis Green, 4 syntype Q, Sri
‘Lanka (Ceylon): Gammaduwa, on Castilloa
| [= Castilla] elastica (E. E. Green) (BMNH).

i

DistRIBUTION. Palaearctic Region: Aden. Mal-
Region: Madagascar, Mauritius,

Oriental Region: Bangladesh,
| Burma, China, Formosa, India, Sri Lanka, Viet-
‘nam. Austro-oriental Region: Borneo, Indo-
‘nesia, West Malayasia, Papua New Guinea,

47

Philippines, Java, Cocos Keeling Island. Neo-
tropical Region: Guyana.

HostpLants. Anacardaceae: Mangifera indica.
Annonaneae: Annona sp., Cananga odorata.
Asteraceae: Adenophyllum sp. Bombaceae:
Ochroma sp. Dioscoreaceae: Dioscorea sp. Dip-
terocarpaceae: Dipterocarpus sp. Ehretiaceae:
Cordia myxa. Euphorbiaceae: Codiaeum sp.,
Euphorbia pyrifolia, | Mallotus japonicus.
Tridaceae: Gladiolus carmels. Lecythidaceae:
Couroupita guianensis. Leguminosae: Albizia leb-
beck, Arachis hypogea, Bauhinia monandra,
Cajanus sp., Erythrina lithosperma, E. variegata,
Hymenaea sp., Prosopsis juliflora, Tamarindus
indica. Moraceae: Castilloa elastica, Ficus rubra.
Myrtaceae: Eugenia mespiloides, Psidium guava.
Palmae: Cocos nucifera, Phoenix dactylifera. Pan-
daceae: Pandanus sp. Puniaceae: Punica gra-
natum. Rhamnaceae: Alphitonia — incana,
Zizyphus jujuba. Rubiaceae: Coffea canephora,
C. sepahijala. Rutaceae: Citrus aurantium, C.
grandis. Sapindaceae: Litchi sp. Simaroubaceae:
Ailanthus sp. Solanaceae: Nicotiana tabacum.
Sterculiaceae: Theobroma cacao. Umbelliferae:
Apium graveolens. Verbenaceae: Tectona gran-
dis. Vitidaceae: Vitis vinifera.

ReMARKS. The combination of stout legs, long
dorsal setae and reduced numbers of multilocular
disc pores distinguishes this species from the other
known species of Planococcus. Some variation
occurs in the thickness of the anterior cerarian
setae; usually these are moderately stout and coni-
cal, but some specimens, notably those from
Aden, have some of these setae long and
flagellate.

Avasti & Shafee (1987: 38) described a new
species, Planococcus indicus, from India on a wild
plant. Although no type material has been exam-
ined, from the original description and illustration
it seems likely that P. indicus is conspecific with P.
lilacinus.

In the 1920s and 1930s the African species P.
kenyae was frequently misidentified as P.
lilacinus, but the latter is not, as yet, known from
this continent. The phylogenetic relationships of
P. lilacinus are obscure; it does not seem to fit into
any of the species-groups into which Planococcus
has been divided here.

Economic status. P. lilacinus is a pest of cocoa
throughout the Oriental Region and also occurs
on a wide variety of economically important crops
such as Citrus, guava, coffee, custard apple and
mango. Le Pelley (1943b) discusses the hos-
tplants, biology and natural enemies of this spe-
cies in some detail.

48
Planococcus litchi sp. n.
(Fig. 23)

ADULT FEMALE. Mounted specimens oval, length
1.3-2.7 mm, width 0.7—2.0 mm. Margin of body
with a complete series of 18 distinct pairs of
cerarii, each cerarius with 2 conical setae except
for the preocular and postocular cerarii each with
1-3 setae, all cerarian setae elongate-conical and
with flagellate tips. Legs elongate; hind trochan-
ter + femur 240-325 um long, hind tibia tarsus
270-345 ym long; ratios of lengths of hind tibia +
tarsus to hind trochanter + femur 1.07—1.16;
translucent pores visible on hind coxae and tibiae.
Inner edges of ostioles lightly sclerotized. Circulus
quadrate and of moderate size, width 95-165 um.
Cisanal setae shorter than anal ring setae. Anal
lobe cerarii each situated on a small, moderately
sclerotized area.

Venter. Multilocular disc pores confined to
median areas of body, present around vulva, in
single rows (sometimes double on segment VII in
larger specimens) across posterior edges of
abdominal segments III-VII (sometimes 1 or 2
pores also on segment II) and anterior edge of
abdominal segment VII (sometimes 1—4 pores
also present on anterior border of segment VI and
occasionally a single pore present on anterior
border of segment V); a few pores scattered over
thorax. Trilocular pores somewhat sparse. Oral
collar tubular ducts of 2 sizes; smaller size present
in rows across median areas of abdominal seg-
ments VI or VII; larger size present in rows across
abdominal segments III-V, scattered over median
areas of thorax and occurring in groups around
entire venter, but usually absent, or at most 1 duct
adjacent to each postocular cerarius. Simple pores
about half the size of the trilocular pores, sparsely
scattered over entire venter.

Dorsum. Multilocular disc pores absent. Tri-
locular pores moderately numerous and aggreg-
ated around bases of enlarged setae. Tubular
ducts, larger than the larger ducts on the venter
and without apparent rims, sometimes present
singly next to some abdominal cerarii and sub-
medially on some abdominal and thoracic seg-
ments, up to 11 ducts present, but frequently
absent. Simple pores as for venter but very sparse.
Dorsal setae of various sizes, larger setae elong-
ate-conical and with flagellate apices, about the
same size as the cerarian setae and each with 2-4
trilocular pores associated with their bases giving
the appearance of dorsal cerarii, these enlarged
setae present on all body segments except the last
abdominal segment; several enlarged setae asso-
ciated in pairs, submedially and medially on the

JENNIFER M. COX

thorax, and medially on adbominal segment VII;
length of longest seta on abdominal segments VI
or VII 25-30 um.

MATERIAL EXAMINED

Holotype 2, Hong Kong (intercepted at Heath-
row, U.K.): on lychees, 1980 (no collector)
(BMNH).

Paratypes. Japan (intercepted in Hawaii): 1 9, on
fruit of Litchi sp., 17.v.1959 (R. O. Parsons)
(USNM). Philippines (intercepted at San Fran-
cisco): 6 2, on Litchi chinensis fruit, 26.v.1972 (E.
Roberts) (USNM). Thailand: 1 9 (intercepted in
U.S.A.), on Litchi sp. (fruit), 29.vi.1980 (L.
Roberson) (USNM); 2 @ (intercepted in
England), on lychees, 1984 (BMNH); 1 @ (inter-
cepted in U.S.A.), on Eriobotrya japonica (fruit),
6. viii. 1982 (B. Stephenson) (USNM). Hong Kong:
10 2 (intercepted at Seattle), on Litchi sp. (fruit),
30.v.1976 (C. R. Payne) (USNM); 1 @ (inter-
cepted at Seattle), on Litchi chinensis (fruit),
22.viii.1974 (R. F. Goodall, M. Harris) (USNM);
1 Q (intercepted?), on litchi, 10.viii.1961 (R. T.
Mitsude) (USNM). China (intercepted at Seat-
tle): 3 2, on litchi (twigs and fruit), 22.v.1960 (H.
C. Nelson) (USNM).

DistTRIBUTION. Palaearctic Region: Japan. Orien-
tal Region: China, Hongkong, Thailand. Austro-
oriental Region: Philippines. All of these records
are based on interceptions in either the U.K. or
the U.S.A.

HostpLants. Rosaceae: Eriobotrya japonica.
Sapindaceae: Litchi chinensis. Most commonly
found on lychees.

Remarks. The enlarged dorsal setae with trilocu-
lar pores aggregated around their bases indicate
that this species is a member of the P. dor-
sospinosus-group. It may be distinguished from
the other 2 known species of this group, P.
dioscoreae and P. dorsospinosus, by the larger
size and grouping of the enlarged dorsal setae into
several pairs.

Economic status. P. litchi is frequently inter-
cepted on lychee fruit imported into the U.S.A.
and the U.K.

Planococcus mali Ezzat & McConnell
(Fig. 24)

Planococcus mali Ezzat & McConnell, 1956: 93;
Williams, 1985: 274; Cox, 1987: 75. Holotype
Q, NEw ZEALAND (intercepted in U.S.A.): on
Olearia chathamica (USNM) [examined].

MEALYBUG GENUS PLANOCOCCUS

49

Fig. 23. Planococcus litchi sp. n.

50

JENNIFER M. COX

Fig. 24 Planococcus mali Ezzat & McConnell.

ADULT FEMALE. Mounted specimens oval, length
2.3-3.3 mm, width 1.2—2.0 mm. Margin of body
usually with a complete series of 18 pairs of
cerarii, although some pairs are often indistinct or
even absent, cephalic and thoracic pairs each with
2-5 conical setae and abdominal pairs each with 2
conical setae. Legs elongate, hind femur +
trochanter 275-380 um long, hind tibia + tarsus
305-410 um long, ratio of lengths of hind tibia +

tarsus to hind trochanter + femur 1.03-1.17;
translucent pores present on hind coxae and
tibiae. Inner edges of ostioles lightly sclerotized.
Circulus quadrate, width 80-190 um. Cisanal
setae shorter than anal ring setae. Anal lobe
cerarii each situated on a small, moderately sized,
moderately sclerotized area.

Venter. Multilocular disc pores situated around
vulva, in rows across posterior edges of median

MEALYBUG GENUS PLANOCOCCUS

3:0

2:0

00

PRINCIPAL COMPONENT 2

53:0) =2°0 -10

0:0

51

1:0 2:0 3-0 4:0

PRINCIPAL COMPONENT 1

Fig. 25

Plot of first against second principal components of material of japonicus/mali from different countries.

Squares, New Zealand; diamonds, Australia; circles, Japan.

areas of abdominal segments III-VII or IV-VII
and across anterior edges of segments V-VII or VI
VII, a few pores present on margins of segments
V-VII or VI or VII, a single pore sometimes situ-
ated on head, 0-22 pores scattered over median
areas of thorax, and in larger specimens, up to 7
pores present amongst the tubular ducts grouped
on each side of the prothorax. Trilocular pores
evenly distributed. Oral collar tubular ducts of 2
sizes; smaller ducts in rows across median areas of
abdominal segments II-VII; larger ducts in rows
across median areas of segments IV-VI, in margi-
nal groups on segments I-VIII, sometimes in a
marginal group on each side of prothorax, and
sometimes present in small numbers on head and
margins of mesothorax. Simple pores not
apparent.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores and simple pores as
for venter. Setae short and stout, sometimes
almost conical, longest seta on abdominal seg-
ments VI or VII 15-20 wm long.

MATERIAL EXAMINED

20 © (including type material listed below)
(BMNH, NZAC, USNM).

Holotype 2, New Zealand (intercepted at Hono-
lulu): on Olearia chathamica, 21.ix.1937

(USNM). Paratypes, 1 2, same data as holotype.
Australia: 1 2 , Tasmania (intercepted at Buffalo,
New York), on Pyrus malus [Malus pumila},
26.vi. 1946 (Inman Reeges); 1 2, same locality and
host (intercepted at Boston, Massachusetts),
27.vi.1946 (USNM).

DistTRIBUTION. Australian Region: Tasmania.
New Zealand Region: New Zealand. Also
recorded by Williams (1985) from New South
Wales.

HostpLants. Asteraceae: Olearia chathamica.
Grossulariaceae: Ribes nigrum. Labiatae: Phlo-
mis sp. Leguminosae: Acacia sp. A. verheillata,
Ulex sp. Pittosporaceae: Pittosporum sp. Pri-
mulariaceae: Primula sp. Rosaceae: Cotoneaster
sp., Malus pumila. Also recorded by Williams
(1985) from Acacia longifolia and Psoralea pin-
nata (Leguminosae), and Callitris tasmanica
(Cupressaceae).

Remarks. P. mali is so similar to P. japonicus that
a principal components analysis was carried out to
confirm their distinction (Fig. 25). The two spe-
cies differ primarily by the fewer tubular ducts in
the prothoracic groups and larger legs of P. mali.
This species is very variable, however, and large
specimens may also have large groups of ducts on

52

the prothorax. These specimens usually also have
multilocular disc pores in these groups, a feature
never found in P. japonicus.

It seems unlikely that sister-species should
occur in Japan and New Zealand/Australia re-
spectively. A possible explanation for this is that
P. mali has been introduced into New Zealand
and Australia from somewhere in the vicinity of
Japan. This theory is supported by its host prefer-
ences in New Zealand and Australia; it occurs
most commonly on introduced northern-temper-
ate plants such as Ribes and Malus, and is, in fact,
a pest of blackcurrant in New Zealand. Extensive
collecting in Japan may reveal a complex of spe-
cies. The lack of records of this species from
Japan, if it does indeed occur there, might seem
surprising at first, but it is possible that this species
has been misidentified as Crisicoccus azaleae. This
is discussed further under P. japonicus.

Economic status. P. mali is a pest of blackcur-
rants in New Zealand.

Planococcus martini sp. n.
(Fig. 26)

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.9-2.8 mm, width 1.4-2.1 mm.
Margin of body with a complete series of 18 dis-
tinct pairs of cerarii, each cerarius with 2 conical
setae. Antennae 7-segmented. Legs somewhat
stout; hind trochanter + femur 200—230 um long,
hind tibia + tarsus 185-220 um long, ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 0.91-0.96; translucent pores present on
hind coxae and a few barely apparent on hind
tibiae. Inner edges of ostioles barely sclerotized.
Circulus oval, width 70-105 wm. Cisanal setae
shorter than anal ring setae. Anal lobe cerarii each
situated on a fairly prominent sclerotized area.

Venter. Multilocular disc pores confined to
median areas, present around vulva and in single
rows across posterior edges of abdominal seg-
ments V or VI, sometimes also IV. Trilocular
pores somewhat sparse. Oral collar tubular ducts
of one size, present in rows across median areas of
abdominal segments VI or VII, usually also V.
Simple pores about the same size as the trilocular
pores, sparsely scattered over entire venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores moderately numer-
ous and evenly distributed. Simple pores about
half the size of the trilocular pores, sparsely scat-
tered over dorsum. Setae short and stout, length
of longest seta on abdominal segments VI or VII
16-25 um.

JENNIFER M. COX

MATERIAL EXAMINED

Holotype 9, Indonesia: Sulawesi Utara, nr Base

Camp Toraut, roots of Araceae, 7Epipremnum

sp., 5.ii1.1985 (J. H. Martin) (BMNH).
Paratypes. 19 9, same data as holotype

(BMNH, USNM).

DISTRIBUTION.
Indonesia.

Austro-oriental Region:

HostTpLants. Araceae: ?Epipremnum sp.

REMARKS. This species is most similar to the
dendrobii-group by virtue of its rotund body and
in having the multilocular disc pores and oral col-
lar tubular ducts confined to the abdomen. It dif-
fers from this group by having 7-segmented
antennae and by lacking knobbed dorsal setae,
and from the similar P. zairensis by having groups
of tubular ducts present on, but multilocular disc
pores absent from, the margins of abdominal seg-
ments VI VII.

Planococcus minor (Maskell) nom. rev., stat.
n., comb. n.

(Fig. 27)

Pseudococcus calceolariae var. minor Maskell,
1897: 322. Lectotype , MAuRITIUs: on roots of
‘onion grass’ (NZAC) here designated
[examined].

Planococcus pacificus Cox, 1981: 48. Holotype @,
WESTERN SAMOA (intercepted in quarantine,
Auckland, New Zealand): on croton leaves
(BMNH) [examined]. Syn. n.

ADULT FEMALE. Mounted specimens oval, length
1.3-3.2 mm, width 0.8-1.9 mm. Margin of body
with complete series of 18 pairs of cerarii, each
cerarius with 2 conical setae, except for preocular
cerarii each of which may have 1 or 3 setae. Legs
elongate; hind trochanter + femur 220-360 um
long, hind tibia + tarsus 270-360 ym long, ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 1.05-1.15; translucent pores apparent on
hind coxae and tibiae. Inner edges of ostioles
moderately sclerotized. Circulus quadrate, width
85-160 um. Cisanal setae usually shorter than
anal ring setae, occasionally longer. Anal lobe
cerarii each situated on a small, moderately scle-
rotized area.

Venter. Multilocular disc pores present around
vulva, in double rows across posterior edges of
abdominal segments III-VII (except in very small
specimens, hind tibia tarsus less than 250 wm,
where these rows may be single), in single rows
across anterior edges of segments V-VII or
VI-VII, in small groups on margins of abdominal

MEALYBUG GENUS PLANOCOCCUS

53

Fig. 26 Planococcus martini sp. 0.

segments IV-VII, sometimes a few pores scat-
tered over median area of the thorax and head,
and frequently several pores present behind each
front coxa. Trilocular pores moderately numerous
and evenly distributed. Oral collar tubular ducts
of 2 sizes; smaller ducts present sparsely in rows
across median areas of abdominal segments III-
VII; larger ducts present in marginal groups on
abdominal segments, often present in small num-
bers on margins of head and thoracic segments

(see Table 1), and scattered over median area of
thorax. Simple pores about the same size as the
trilocular pores, sparsely but evenly distributed.
Dorsum. Multilocular disc pores absent. Tubu-
lar ducts, without apparent rims and slightly larger
than the larger size on the venter, sometimes pre-
sent adjacent to some cerarii, 1 or 2 ducts occa-
sionally present on median areas. Trilocular pores
as for venter. Simple pores of 2 sizes, smaller
pores smaller than those on venter, scattered over

‘ JENNIFER M. COX |
,

54

Fig. 27 Planococcus minor (Maskell).

MEALYBUG GENUS PLANOCOCCUS

dorsum; larger pores about twice the size of the
trilocular pores, present in small groups along
midline of thoracic and anterior abdominal seg-
'ments. Setae flagellate and of moderate length,
‘longest seta on abdominal segment VI or VII
25-30 wm long.

MATERIAL EXAMINED

- Over 200 9 (including type material listed below)
(BMNH, NZAC).

Dactylopius calceolariae var. minor Maskell. Lec-
totype 9 here designated, on slide labelled: ‘Dac-
tylopius calceolariae var. minor, adult female,
1896, W. M. M.’ (NZAC).

Planococcus pacificus Cox. Holotype 2, Western
| Samoa (intercepted at Auckland, New Zealand):
/on croton leaves, 10.v.1979 (C. Butcher)
| (BMNH). 385 paratype 9, reared from holotype
| in England, on potato tubers at various tempera-
tures, 18.vi.1979-30.viii.1979 (J. M. Cox)
(BMNH, NZAC, USNM).

DIsTRIBUTION. Oriental Region: Bangladesh,
Chagos Arch., Burma, India, Rodrigues [.,
| Seychelles Is, Thailand. Austro-oriental Region:
' Borneo, West Malaysia, Papua New Guinea, Phi-

lippines, Singapore, Sumatra. Australian Region:
| New South Wales, Northern Territory, Queens-
_ land, South Australia. Polynesian Region: Sol-
' omon Islands, Austral Islands, Cook Islands,
_ American Samoa, Fiji, Kiribati, Niue, Society
Islands, Tokelau, Tonga, Tuvalu, Vanuatu, West-
' ern Samoa. Neotropical Region: Barbados,
Ecuador, Grenada, Jamaica, Trinidad, St Lucia,
| Virgin Islands. Malagasian Region: Madagascar,
_ Mauritius.

HosteLants. Acanthaceae: Pachystachys coc-
cineae. Agavaceae: Furcraea _ gigantea.
Amaranthaceae: Celosia sp. Annonaceae:
Annona muricata. Araceae: Anthurium sp., Colo-
casia antiquorum, Dieffenbachia sp. Pistia strat-
iotes, Xanthosoma sagittifolium. Asclepiadaceae:
Hoya sp. Asteraceae: Emilia sonifolia. Cac-
taceae: Harrisia portaricensis. Cannaceae: Canna
sp. Cucurbitaceae: Luffa actangula. Ebenaceae:
Diospyros discolor. Ehretiaceae: Cordia myxa.
Euphorbiaceae: Alcalypha hispida, Cassava util-
issima, Codiaeum variegatum Croton sp. Gera-
niaceae: Geranium sp. Labiatae: Epimeredi
indicus, Ocimum sanctum. Leguminosae: Glir-
icidia maculata, Glycine max, Psophocarpus
tetragonolobus, Tephrosia sp. Liliaceae:
Asparagus sp. Malvaceae: Gossypium sp. Maran-
taceae: Maranta sp. Moraceae: Artocarpus com-
munis, Ficus elastica. Musaceae: Musa sapientum.
Myrtaceae: Psidium guajava. Nyctaginaceae:
Boerhavia diffusa. Palmae: Chrysalidocarps sp.,

55

Howeia forsteriana. Passifloraceae: Passiflora
edulis. Periplocaceae: Mondia citrifolia.
Piperaceae: Piper nigrum. Poaceae: Bambusa sp.,
Orya sativa. Polygonaceae: Rumex sp. Rosaceae:
Fragaria sp. Rubiaceae: Coffea arabica, C. liber-
ica, C. robusta, Randia heterophylla. Rutaceae:
Choisya sp. Sapindaceae: Nephelium lappaceum.
Sapotaceae: Manilkara zapota. Solanaceae:
Lycopersicon esculentum, Solanum grandiflorum,
S. torrum, S. tuberosum. Sterculiaceae: The-
obroma cacao. Thunbergiaceae: Thunbergia sp.
Verbenaceae: Clerodendrum thompsonae, Ver-
bena sp. Many further host plants are listed by
Williams (1982) under P. pacificus.

Remarks. Dactylopius calceolariae var. minor was
synonymized with Planococcus citri by Morrison
(1925). The type material of the former taxon,
however, although poor, appears to be the same
species as that described as P. pacificus by Cox
(1981). Additional evidence lies in the fact that no
specimens of P. citri from the Malagasian Region
were encountered during this study.

P. minor is very similar to P. citri, and the
existence of the second species was not estab-
lished until the variation of individual populations
was studied using rearing experiments by Cox
(1981, 1983) (see also under Introduction).

Although P. citri has frequently been recorded
from the South Pacific Islands, Williams (1982)
comments that most of these records are misiden-
tifications of P. minor. His records show P. minor
to be much more common than P. citriin this area,
and to have been substantially longer established,
the earliest record given of P. citri from the area
being 1975 and that of P. minor, 1922

Likewise, Cox & Freeston (1985) showed that
the species of Planococcus commonly occurring
on Theobroma cacao in the Oriental and Neo-
tropical Regions was in fact P. minor and not P.
citri, although most published records are under
the latter name.

EconomIc STATuS. P. minor is a common species
on many economically important plants, par-
ticularly cocoa, throughout its geographical
range.

Planococcus morrisoni (Ezzat & McConnell)
(Fig. 28)

Allococcus morrisoni Ezzat & McConnell, 1956:
17. Holotype 2, PHILiPPINEs: on Lansium dom-
esticum (USNM) [examined].

Planococcus morrisoni (Ezzat & McConnell) Cox
& Ben-Dov, 1986: 488.

JENNIFER M. COX |

56

ig. 28 Planococcus morrisoni (Ezzat & McConnell)

F

MEALYBUG GENUS PLANOCOCCUS

ADULT FEMALE. Mounted specimens elongate-
oval, length 1.4-2.4 mm, width 0.6-1.3 mm. Mar-
gin of body with complete series of 18 pairs of
cerarii, each cerarius with 2 slender conical setae,
except for head and thorax where some cerarii

_ may be indistinct and others with up to 4 conical
| setae. Legs elongate; hind trochanter + femur
_ 210-285 um long, hind tibia + tarsus 230-295 um
_ long, ratio of lengths of hind tibia + tarsus to hind
_ trochanter + femur 1.04-1.09; translucent pores
| apparent on hind coxae and tibiae. Inner edges of
| ostioles only slightly sclerotized. Circulus quadr-
| ate, width 40-115 um. Cisanal setae shorter than
| anal ring setae. Anal lobe cerarii situated on

small, moderately sclerotized areas.

Venter. Multilocular disc pores present around
vulva, in double rows across posterior edges of
abdominal segments IV-VII and sometimes in a

| single row on segment III, scattered over anterior
| edges of segments V-VII; also present in small

groups on margins of adominal segments I-VIII

' and scattered over median area of the thorax.
_ Trilocular pores moderately numerous and evenly

distributed. Oral collar tubular ducts of 2 sizes;
smaller ducts present sparsely in rows across
median areas of abdominal segments IV-VII and
scattered over median areas of thorax; larger size
present sparsely in rows across median areas of
abdominal segments III-VI, and in marginal
groups on segments I-VIII and on prothorax adja-
cent to front coxae. Simple pores considerably
smaller than the trilocular pores, sparsely scat-
tered over venter.

Dorsum. Multilocular disc pores ducts absent.
Tubular ducts, larger than the larger size on the
venter and with distinct rims, present singly next

_ tosome cerarii and sparsely scattered over median

areas. Trilocular pores and simple pores as for
venter. Setae short and stout, almost lanceolate,

| longest seta on abdominal segments VI or VII

10-20 pm long.

MATERIAL EXAMINED

9 © (including type material listed below)
(BMNH, USNM).

Holotype 9, Philippines, on Lansium dom-
esticum, 30.viii.1951 (R. O. Parsons) (USNM).

DistRiBUTION. Oriental Region: Thailand. Aus-
tro-oriental Region: Philippines, West Malaysia.
All of these records are based on interceptions in
the U.S.A.

HostpLants. Clusiaceae: Garcinia mangostana.
Meliaceae: Lansium domesticum. Sapindaceae:

Litchi chinensis, Melicoccus bijugatus.

Remarks. The presence of marginal multilocular
disc pores, short, stout, almost lancolate, dorsal

a7

setae, and a group of tubular ducts adjacent to
each anterior coxa, whilst these ducts are absent
from the head, show this species to be allied to P.
mali. The large tubular ducts with pronounced
oral rims, however, clearly distinguish P.
morrison.

ECONOMIC STATUS. This species is frequently inter-
cepted on fruit entering the U.S.A.

Planococcus nigritulus De Lotto
(Fig. 29)

Planococcus nigritulus De Lotto, 1961: 222. Holo-
type 2, Zaire: on Phoenix ‘canariensis’ (given
as dactylifera on the label) (BMNH)
[examined].

ADULT FEMALE. Mounted specimens oval to
rotund, length 1.9-2.4 mm, width 1.3-1.8 mm.
Margin of body with complete series of 18 pairs of
cerarii, each cerarius with 2 moderately stout con-
ical setae. Legs somewhat stout; hind trochanter
+ femur 255-285 um long, hind tibia + tarsus
245-275 um long, ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 0.93-1.10;
translucent pores present on hind coxae and
tibiae. Inner edges of lips of ostioles not noticea-
bly sclerotized. Circulus large and quadrate,
width 130-190 ym. Cisanal setae longer than anal
ring setae. Anal lobe cerarii each situated on a
very small, lightly sclerotized area.

Venter. Multilocular disc pores present around
vulva, in rows across posterior edges of median
areas of abdominal segments [V-VII and across
anterior edges of segments VI or VII, and present
in small marginal groups on segments IV-VII.
Trilocular pores numerous and evenly dis-
tributed. Oral collar tubular ducts of 2 sizes;
smaller ducts present in rows across median areas
of abdominal segments V-VII; larger ducts occur-
ring in rows across median areas of abdominal
segments IV V and in marginal groups around
entire venter except for abdominal segments VIII
or [X from which they are entirely absent. Simple
pores about twice the size of the trilocular pores,
scattered over entire venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores of 2 sizes, larger pores the same size as
those on the venter, in median groups on thoracic
and anterior abdominal segments and scattered
sparsely over thorax, smaller size slightly smaller
than the trilocular pores, scattered over the
abdominal segments. Setae flagellate, stout and
moderately long, longest seta on abdominal seg-
ments VI or VII 25—40 um long.

ee ee ee ae on lel rl... CO aS 2S ES et eM ee SEH Be

cu
oo ",

%
Se
tant

JENNIFER M. COX

Fig. 29 Planococcus nigritulus De Lotto.

58

MEALYBUG GENUS PLANOCOCCUS

MATERIAL EXAMINED
Holotype 9, Zaire (Belgian Congo): Katana, on
Phoenix dactylifera, 15.x.1941 (F. Hendrick)
(BMNH). 11 paratype 2, same data as holotype
(BMNH). The host is given as P. canariensis in the
original description.

DistriBuTION. Afrotropical Region: Zaire. Also
recorded from Tanzania (as Tanganyika) (De
Lotto, 1964).

HosteLants. Palmae: Phoenix dactylifera. Also
recorded from Ficus sp. (Moraceae) (De Lotto,
1964).

Remarks. P. nigritulus is similar to P. aemulor
and P. tanzaniensis in its rotund body shape,
stout, almost fleshy, dorsal setae, large ventral
simple pores, and absence of oral rim tubular
ducts from abdominal segment IX. Unlike P.
nigritulus, P. aemulor \acks both translucent pores
on the hind legs and marginal tubular ducts on the
venter. P. tanzaniensis possesses tubular ducts on
the median areas of the thorax; these are absent in
_ P. nigritulus.

Planococcus orchidi sp. n.
(Fig. 30)

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.1—2.0 mm, width 1.0-1.5 mm.
Margin of body with a complete series of 18 pairs
of cerarii, each with 2 stout conical setae. Legs
stout; hind trochanter + femur 180-210 ym long,
hind tibia + tarsus 175-220 um long; ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 0.96-1.05; translucent pores present on
hind coxae and tibiae. Inner edges of ostioles well-
sclerotized. Circulus small and quadrate, width
40-75 pm. Cisanal setae shorter than anal lobe
setae. Anal lobe cerarii each situated on a small
moderately sclerotized area.

Venter. Multilocular disc pores confined to
median areas of abdomen, present around vulva
and in single rows across posterior edges of
median areas of abdominal segments V VI. Tri-
locular pores moderately numerous and evenly
distributed. Oral collar tubular ducts of 2 sizes,
both confined to abdomen, smaller ducts present
in single rows across median areas of abdominal
segments V-VII, larger ducts present in small
marginal groups on segments V-VII. Simple pores
about twice the size of the trilocular pores, heavily
sclerotized, and scattered over entire venter.
Setae on median areas moderately long and fine,
but those on margins short, moderately fine, and
with distinctly knobbed tips.

59

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores numerous and
evenly distributed. Simple pores of 2 forms; pores
slightly larger than the trilocular pores, and only
lightly sclerotized, scattered sparsely over entire
dorsum; heavily sclerotized pores, noticeably
larger than the trilocular pores, present in small
groups on midline of posterior abdominal seg-
ments. Setae moderately short and stout, and with
distinctly knobbed tips. Longest seta of abdomi-
nal segments VI or VII 15-30 um long.

MATERIAL EXAMINED
Holotype 9, Liberia (intercepted at New York):
on wild orchids, 10.x.1957 (C. E. Andrews, J.
Hidalgo) (USNM).

Paratypes. 4 9, same data as holotype (BMNH,
USNM).

DistriBUTION. Afrotropical
(intercepted in the U.S.A.).

Region: Liberia

HostTp.Lant. Orchidaceae.

Remarks. P. orchidi resembles the Oriental spe-
cies P. dendrobii and P. philippinensis, which also
occur on orchids, in having few multilocular disc
pores and tubular ducts, and in having dorsal setae
which are distinctly knobbed. It may be dis-
tinguished from both of these species by the pres-
ence of translucent pores on the hind tibiae and by
the large, heavily sclerotized simple pores on the
venter. P. orchidi is also similar to two orchid-
infesting Afrotropical species with knobbed dor-
sal setae, P. hospitus and P. hosyni, both of which
lack marginal tubular ducts.

Planococcus philippinensis Ezzat &
McConnell

(Fig. 31)

Planococcus philippinensis Ezzat & McConnell,
1956: 95. Holotype 2, PHILIPPINES (intercepted
in Hawaii): on orchids (USNM) [not
examined].

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.4-2.3 mm, width 1.0-1.9 mm.
Margin of body with a complete series of 18 pairs
of cerarii, each with 2 stout conical setae. Legs
stout; hind trochanter + femur about 240 um
long, hind tibia + tarsus about 220 um long, ratio
of lengths of hind tibia + tarsus to hind trochanter
+ femur about 0.92; translucent pores apparent
on hind coxae only. Inner edges of ostioles appar-
ently not sclerotized. Circulus absent. Cisanal
setae longer that anal ring setae. Anal lobes
apparently not sclerotized.

60

Fig. 30 Planococcus orchidi sp. n.

JENNIFER M. COX

MEALYBUG GENUS PLANOCOCCUS

61

AN
AN
By)

.

Fig. 31 Planococcus philippinensis Ezzat & McConnell.

Venter. Multilocular disc pores sparse and con-
fined to median areas of abdomen, a few pores
present around vulva and on posterior edges of
median areas of abdominal segments V or VI.
Trilocular pores moderately numerous and evenly
distributed. Oral collar tubular ducts of 1 size

| only, confined to abdomen, present in rows across

—_

median areas of abdominal segments VI or VII, in
marginal groups on segments VI or VII, and a few

pores sometimes on margins of segment V. Simple
pores about a third of the size of the trilocular
pores, sparsely but evenly distributed.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores, slightly smaller than the trilocular
pores, sparsely scattered over most of dorsum, but
those on median areas of abdominal segments
numerous and noticeably larger than the trilocular

62

pores. Setae short and stout, slightly curved and
with distinctly knobbed tips.

MATERIAL EXAMINED

1 paratype °, Philippines (intercepted at Hono-
lulu): on Vanda sanderiana, 4.vi.1948 (H. Mak-
ing) (USNM).

DIsTRIBUTION. Austro-oriental Region: Philip-
pines (intercepted in the U.S.A.).

HostTpLants. Orchidaceae: Vanda sanderiana.
Also recorded from Cymbidium finlaysonianum,
Vanda merrillii and Vandopsis lissochiloides (all
Orchidaceae) (Ezzat & McConnell, 1956).

REMARKS. This species is very similar to P.
dendrobii, also from the Oriental Region, in hav-
ing reduced numbers of tubular ducts and multi-
locular disc pores, translucent pores absent from
the hind tibiae and dorsal setae which are short,
stout and knobbed. P. dendrobii, however, has a
circulus, and the simple pores on the dorsum of
abdominal segments VI or VII are smaller than
the trilocular pores. Together, these two species
are probably the sister-group of two African spe-
cies which also occur on orchids, P. hosyni and P.
orchidi. These latter species also have knobbed
dorsal setae (although indistinct in P. hosyni), but
are distinguishable from the Oriental species by
having translucent pores on the hind tibiae.

EcONoMIC STATUS. This species is a potential pest
of orchids.

Planococcus principe sp. n.
(Fig. 32)

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.6—2.8 mm, width 1.2—2.1 mm.
Margin of body with complete series of 18 pairs of
cerarii, each cerarii with 2 (3 or 4 in preocular
cerarii) conical setae. Legs stout; hind trochanter
+ femur 275-295 um long, hind tibia + tarsus
240-270 um long, ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 0.84—0.92;
translucent pores present on hind coxae and
tibiae. Inner edges of ostioles moderately well
sclerotized. Circulus quadrate, width 125-180
pm. Cisanal setae about the same length as the
anal ring setae. Anal lobe area moderately well
sclerotized.

Venter. Multilocular disc pores present around
vulva, in rows across posterior edges of abdominal
segments IV-VII and forming marginal groups on
segments V or VI; a few pores sometimes present
on the head. Trilocular pores numerous and
evenly distributed. Oral collar tubular ducts very
small, in rows across median areas of abdominal

JENNIFER M. COX

segments V-VII and in marginal groups on head,
prothorax and abdominal segments V-VII; a sin-
gle larger duct sometimes present on margin of
segment V. Simple pores about three quarters of
the size of the multilocular disc pores, scattered
over venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores as for venter. Sim-
ple pores of various sizes, larger pores about the
same size as those on the venter, forming a group
on the midline of each thoracic segment and scat-
tered over submarginal areas of head and thorax,
slightly smaller pores scattered over median areas
of anterior abdominal segments. Setae moder-
ately long and stout, those on the thorax some-
times distinctly knobbed, length of longest seta on
abdominal segments VI or VII 30-40 um.

MATERIAL EXAMINED

Holotype Q, Principe: on cocoa, 1956 (F. J. Sim-
monds) (BMNH).

Paratypes. 7 9, same data as holotype (BMNH).

DistriBuTion. Afrotropical Region: Principe.
HostTpLant. Sterculiaceae: Theobroma cacao.

Remarks. The only other known species of
Planococcus which has simple pores of almost the
same size as the multilocular disc pores is P. hospi-
tus. The presence of marginal multilocular disc
pores in P. principe distinguishes the 2 species.

Planococcus psidii sp. n.
(Fig. 33)

ADULT FEMALE. Mounted specimens oval, length
1.42.7 mm, width 0.7-1.8 mm. Margin of body
with complete series of 18 pairs of ceraii, each
cerarius with 2 slender conical setae. Legs elong-
ate; hind trochanter + femur 245-285 um, hind
tibia + tarsus 255-325 um, ratio of lengths of hind
tibia + tarsus to hind trochanter + femur 1.04—
1.18; large transparent pores very conspicuous on
hind coxae and tibiae. Inner edges of ostioles wll
sclerotized. Circulus quadrate, width 105-145
pm. Cisanal setae shorter than anal lobe setae.
Venter. Multilocular disc pores present around
vulva, in rows across posterior edges of median
areas of abdominal segments III-VII and across
anterior edges of segments V-VII, occurring in
marginal groups on segments IV-VIII, a few pores
also present on median areas of the thorax. Tr-
locular pores sparsely but evenly distributed. Oral
collar tubular ducts of 2 sizes; smaller ducts pre-
sent in rows across median areas of posterior
abdominal segments; larger ducts present in rows
across anterior abdominal segments, scattered

MEALYBUG GENUS PLANOCOCCUS

over median areas of thorax, occurring singly on
margins of head and thoracic segments and pre-
sent in marginal groups on abdomen. Simple
pores about the same size as the trilocular pores,
sparsely scattered over entire venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores sparse and some
appear to be associated with the bases of the larger
setae. Simple pores of 2 sizes, smaller pores min-
ute and sparsely scattered over entire dorsum,
larger pores about the same size as those on the
| venter and forming a small group on the midline of

63

Fig. 32 Planococcus principe sp. 0.

each thoracic and anterior abdominal segment.
Setae short and stout, blunt-ended or slightly
knobbed, length of longest seta on abdominal seg-
ments VI or VII 15-22 um.

MATERIAL EXAMINED
Holotype 9, West Malaysia: Flemming Est., on
Psidium guajava, 21.1.1988 (K. C. Khoo)
(BMNH).

Paratypes. 11 9, same data as holotype
(BMNH).

64

Fig. 33

|

Planococcus psidii sp. n.

JENNIFER M. COX

65

MEALYBUG GENUS PLANOCOCCUS

| lI

LA OW, oy

Fig. 34 Planococcus subterraneus De Lotto.

66

DISTRIBUTION.
Malaysia.

Austro-oriental Region: West

Host. Myrtaceae: Psidium guajava.

REMARKS. P. psidii is superficially similar to P.
minor in its distribution of multilocular disc pores
and oral collar tubular ducts. However, the short
blunt-ended dorsal setae and the large translucent
pores on the hind legs distinguish P. psidii. This
species may actually be more closely related to the
dorsospinosus-group, as the trilocular pores are
somewhat associated with some of the larger dor-
sal setae.

Planococcus subterraneus De Lotto
(Fig. 34)

Planococcus subterraneus De Lotto, 1964: 377.
Holotype 2, SourH Arrica: Pretoria, on roots
of Ficus sp. (BMNH) [examined].

ADULT FEMALE. Mounted specimen elongate-
oval, length 2.3 mm, width 1.4 mm. Margin of
body with complete series of 18 pairs of cerarii,
each cerarius with 2 slender conical setae. Legs
elongate; hind trochanter + femur 375 um long,
hind tibia + tarsus 405 um long, ratio of lengths of
hind tibia + tarsus to hind trochanter + femur
1.08; translucent pores present on hind coxae and
tibiae. Inner edges of ostioles lightly sclerotized.
Circulus quadrate, width 170 wm. Cisanal setae
shorter than anal ring setae. Anal lobe cerarii each
situated on a small, moderately sclerotized area.

Venter. Multilocular disc pores numerous
around vulva, present in rows across posterior
edges of median areas of all abdominal segments
and across anterior edge of segments IV-VII, pre-
sent in marginal groups on abdominal segments
II-VIII, and numerous over median area of
thorax. Trilocular pores moderately numerous
and evenly distributed. Oral collar tubular ducts
of 2 sizes; smaller ducts present sparsely in rows
across median areas of all abdominal segments;
slightly larger ducts sparsely scattered over
median areas of head and thorax, in marginal
groups on all abdominal segments, and a few ducts
present on margins adjacent to front coxae. Sim-
ple pores about half the size of the trilocular
pores, sparsely but evenly distributed.

Dorsum. Multilocular disc pores absent. Tubu-
lar ducts, larger than the larger ducts on the venter
and without noticeable rims, present singly adja-
cent to some abdominal cerarii (except for one
side of the preanal segment on which 2 ducts are
present), and a few ducts scattered over median

JENNIFER M. COX

areas of the thorax. Trilocular pores as for venter.
Simple pores not apparent. Setae long, stout and
flagellate, some strongly bifurcate, longest seta on
abdominal segments VI or VII about 75 um long.

MATERIAL EXAMINED

Known from holotype 9 only, South Africa: Pre-
toria, on roots of Ficus sp., 15.viii.1958 (H. K.
Munro) (BMNB).

DISTRIBUTION.
Africa.

HosTpLant. Moraceae: Ficus sp.

Afrotropical Region: South

Remarks. P. subterraneus is very similar to P.
flagellatus in its distribution of multilocular disc
pores and possession of long, fleshy dorsal setae.
It differs primarily by having many bifurcate dor-
sal setae and by lacking tubular ducts on the head.
As P. subterraneus is known from a single speci-
men, and P. flagellatus from only a few specimens
which vary greatly, it is possible that further col-
lecting may show these two species to be the same.

Both P. subterraneus and P. flagellatus belong
to the citri-group by virtue of their arrangement of
multilocular disc pores and tubular ducts. Their
distinctive characteristic is the very long, fleshy
and sometimes bifurcate, dorsal setae.

Planococcus sulawesi sp. 0.
(Fig. 35)

ADULT FEMALE. Mounted specimens oval, length
1.6-1.7 mm, width 0.8-0.9 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
cerarius with 2 stout conical setae. Legs elongate;
hind trochanter + femur 285-305 um long, hind
tibia + tarsus 305-325 um long, ratio of hind tibia
+ tarsus to hind trochanter + femur 1.06—1.07;
translucent pores present on hind coxae and
tibiae. Inner edges of ostioles lightly sclerotized.
Circulus quadrate and of moderate size, width
about 135 um. Cisanal setae shorter than anal ring
setae. Anal lobe cerarii each situated on a small
sclerotized area.

Venter. Multilocular disc pores confined to
median areas of body, present around vulva and in

rows across posterior edges of abdominal seg- |

ments III-VII and across anterior edges of seg-
ments VI or VII; a few pores scattered over
thorax. Trilocular pores moderately numerous
and evenly distributed. Oral collar tubular ducts
of 2 sizes; smaller ducts present in rows across
median areas of abdominal segments IV-VII;
larger ducts present in rows across median areas of
segments II-VII, in marginal groups on all thor-
acic and abdominal segments, and scattered over

67

Ciena ae = 2
. \

7

|e

Fig. 35 Planococcus sulawesi sp. n.

MEALYBUG GENUS PLANOCOCCUS

68

median areas of thorax. Simple pores not appar-
ent in the specimens examined.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores moderately numer-
ous and slightly aggregated around the bases of
the larger setae. Simple pores not apparent in the
specimens examined. Setae stout and very long,
occasionally bifurcate, length of longest seta on
abdominal segment VI or VII about 100 um.

MATERIAL EXAMINED

Holotype @, Indonesia: Sulawesi Utara,
Dumoga-Bone National Park, Gunung
Mogogonipa, on Urticaceae, 6.iv.1985 (J. H.
Martin) (BMNH).

Paratype. 1 2, same data as holotype (BMNH).

DISTRIBUTION. Austro-oriental

Indonesia.

Region:

HostTeLantT. Urticaceae.

REMARKS. P. sulawesi is a member of the dor-
sospinosus-group by virtue of its arrangement of
multilocular disc pores and tubular ducts, and by
having trilocular pores associated with the bases
of the larger dorsal setae. It is readily dis-
tinguished from the other members of this group
by its very long dorsal setae. P. sulawesi super-
ficially resembles P. flagellatus, P. subterraneus
and P. lilacinus in the length of its dorsal setae, but
can be distinguished from the former two species,
both Afrotropical, by lacking marginal multilocu-
lar disc pores, from P. lilacinus by having multi-
locular disc pores present on the thorax and by
having elongate legs, and from all three by having
trilocular pores associated with the bases of the
dorsal setae.

Planococcus taigae Danzig
(Fig. 36)

[ Planococcus vovae (Nasonov) Danzig, 1980: 168,
in part, illustration. Misidentification. }

Planococcus taigae Danzig, 1986: 19. Holotype Q,
U.S.S.R: Southern Sakhalin, on Juniperus sib-
irica (ZIL) [not examined].

ADULT FEMALE. Mounted specimens oval, length
1.4-2.3 mm, width 1.0-1.7 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
with 2 slender, conical setae, noticeably more
slender and flagellate towards the anterior end of
the body. Legs elongate; hind trochanter + femur
210-295 pm long, hind tibia + tarsus 220-315 wm
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.05—1.06; translucent pores

JENNIFER M. COX

present on hind coxae and 2-4 pores just visible on
each hind tibia. Ostioles very weakly defined. Cir-
culus not apparent in specimens examined, but
indicated in illustration by Danzig (1980). Cisanal
setae shorter than anal lobe setae. Anal lobe
cerarii each situated on a small, lightly sclerotized
area.

Venter. Multilocular disc pores confined to
median areas only, situated around vulva, in rows
across posterior edges of abdominal segments
IV-VII, and a few pores present on anterior edges
of segments V-VII. Trilocular pores moderately
numerous and evenly distributed. Oral collar
tubular ducts of 2 sizes, smaller ducts present spar-
sely in rows across median areas of abdominal
segments V-VII, larger ducts present in rows
across median areas of segments II-VII and
numerous around margin of entire venter. Simple
pores minute, sparsely but evenly distributed.

Dorsum. Multilocular disc pores absent. Tri-
locular pores and simple pores as for venter.
Tubular ducts, variable in size but larger than
those on the venter and without apparent rims,
numerous over entire dorsum, occurring in rows
of up to 24 ducts across abdominal segments.
Setae short and stout, longest setae on abdominal
segments VI or VII about 13 um long.

MATERIAL EXAMINED

2 paratype 9, U.S.S.R: Primorye Region,
Sudzukhinskiy Nature Reserve, on Juniperus
rigida, 15.vii.1969 (E. Danzig); 1 paratype 9,
Southern Sakhalin, Staroduskoye, 4.vii.1968 (S.
Barfeneva) (all BMNH).

DIsTRIBUTION. Palaearctic Region: U.S.S.R.,
from Karelia to Kuril Islands (Danzig, 1980).

HostTp.Lants. Cupressaceae: Juniperus communis,
J. rigida, J. siberica (Danzig, 1986).

REMARKS. The above description was based on
only three specimens, so the ranges of characters
given may be less than actually occurs throughout
the species.

P. taigae is very similar to P. vovae which also
occurs only on Cupressaceae, and replaces the
latter species in the eastern Palaearctic Region. P.
taigae can be distinguished from P. vovae by its
greater numbers of dorsal tubular ducts (total of
more than 200), presence of ventral tubular ducts
on the head, and indistinct ostioles. P. vovae has
variable numbers of dorsal tubular ducts, but
never more than a total of 110, lacks ventral tubu-
lar ducts on the head and has well-defined
ostioles. Both of these species are similar to
Crisicoccus matesovae (Fig. 37), also occurring on

MEALYBUG GENUS PLANOCOCCUS

Fig. 36

Planococcus taigae Danzig.

69

JENNIFER M. COX

70

: ae
0» (en I\co eo

Fig. 37 Crisicoccus matesovae (Danzig).

MEALYBUG GENUS PLANOCOCCUS

Fig. 38 Planococcus tanzaniensis sp. n.

ql

W2

Cupressaceae in the U.S.S.R., but this species has
dorsal multilocular disc pores and fewer than 18
pairs of cerarii.

Planococcus tanzaniensis sp. n.
(Fig. 38)

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.5—2.5 mm, width 1.1-2.0 mm.
Margin of body with a complete series of 18 pairs
of cerarii, each cerarius with 2 moderately stout
conical setae. Legs elongate; hind femur +
trochanter 250-305 um long, hind tibia + tarsus
295-345 um long, ratio of lengths of hind tibia +
tarsus to hind trochanter + femur 1.09-1.18;
translucent pores present on hind coxae and
tibiae. Inner edges of lips of ostioles moderately
sclerotized. Circulus quadrate, width 95-125 um.
Cisanal setae shorter than anal ring setae. Anal
lobe cerarii each situated on a small, moderately
sclerotized area.

Venter. Multilocular disc pores present around
vulva, in rows across posterior edges of median
areas of abdominal segments IV-VII and across
anterior edges of segments V-VII, and present in
small numbers on margins of segments V-VII.
Trilocular pores moderately numerous and evenly
distributed. Oral collar tubular ducts of 2 sizes;
smaller ducts present in rows across median areas
of abdominal segments; larger ducts scattered
over median areas of thorax, present in small
numbers on margins of head and thoracic seg-
ments, and in small groups on margins of all
abdominal segments except segment IX and
sometimes VIII. Simple pores about the same size
as the trilocular pores, moderately numerous and
scattered over entire venter.

Dorsum. Multilocular disc pores absent. Tri-
locular pores as for venter. Tubular ducts, larger
than the larger size on the venter and without
apparent rims, sometimes present singly adjacent
to posterior cerarii. Simple pores of 2 sizes, larger
simple pores slightly larger than the trilocular
pores, present in small median groups on anterior
abdominal segments, smaller pores, about half
the size of the trilocular pores, scattered over
entire dorsum. Setae flagellate, long and stout,
longest seta on abdominal segments VI or VII
about 30 um long.

MATERIAL EXAMINED

Holotype 2, Tanzania: Selem, on Hevea bra-
ziliensis, 8.1.1987 (S. Oswald) (BMNH).
Paratypes. 13 2, same data as holotype (BMNH).

DisTRIBUTION. Afrotropical Region: Tanzania.

JENNIFER M. COX
HostTpLant. Euphorbiaceae: Hevea braziliensis.

REMARKS. P. tanzaniensis is most similar to P.
aemulor and P. nigritulus in its body shape, dis-
tribution of multilocular disc pores, large ventral
simple pores, long stout dorsal setae and absence
of tubular ducts from abdominal segment IX. It
can be distinguished from both species by its pos-
session of tubular ducts on the median areas of the
thorax.

Planococcus vovae Nasonov
(Fig. 39)

Pseudococcus (Dactylopius) vovae Nasonov,
1908: 484. Lectotype 9, U.S.S.R.: on
Juniperus communis (ZIL) designated by
Danzig (1980) [examined].

Pseudococcus inamabilis Hambleton, 1935: 112.
Syntype 9, BraziL: on Cupressus glauca
(IBSP, ?lost) [not examined]. [Synonymized by
Cox & Ben-Dov (1986).]

Allococcus inamabilis (Hambleton) Ezzat &
McConnell, 1956: 15.

Allococcus vovae (Nasonov) Zahradnik, 1959:
297.

Planococcus vovae (Nasonov) Danzig, 1980: 168;
Cox & Ben-Dov, 1986: 485; Danzig, 1986: 18.

ADULT FEMALE. Mounted specimens oval, length
1.3-3.4 mm, width 0.7-2.3 mm. Margin of body
with a complete series of 18 pairs of cerarii, each
with 2 slender conical setae, noticeably more
slender and flagellate towards the anterior end of
the body. Legs elongate; hind trochanter + femur
230-355 um long, hind tibia + tarsus 265-400 4m
long, ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.09-1.17; translucent pores
present on hind coxae and a few pores sometimes
visible on each hind tibia. Ostioles well defined,
but with lip edges weakly sclerotized. Circulus
quadrate, width 75-135 um. Cisanal setae shorter
than anal lobe setae. Anal lobe cerarii each situ-
ated on a small, lightly sclerotized area.

Venter. Multilocular disc pores confined to
median areas of abdomen, situated around vulva
and in rows across posterior edges of abdominal
segments IV-VII, and a few pores present on ante-
rior edges of segments V-VII. Trilocular pores
moderately numerous and evenly distributed.
Oral collar tubular ducts of 2 sizes; smaller ducts
present sparsely in rows across median areas of
abdominal segments V-VII; larger ducts present
in rows across median areas of segments II-VII, in
marginal groups on all abdominal segments and
scattered over median areas of thorax, and a few

«

is

}

MEALYBUG GENUS PLANOCOCCUS

Be

: | (e)

‘9

RS

ole)

Fig. 39 Planococcus vovae (Nasonov).

73

74

ducts sometimes present on margins of thoracic
segments. Simple pores minute, evenly but spar-
sely distributed.

Dorsum. Multilocular disc pores absent. Tri-
locular pores and simple pores as for venter.
Tubular ducts, variable in size but larger than
those on the venter and usually with rims, variable
in number, most numerous at posterior end of
body, scattered over dorsum, a total of 13-110
ducts present over entire dorsum. Setae moder-
ately long and fine, longest seta on abdominal
segments VI or VII about 20-35 um long.

MaTERIAL EXAMINED

89 (including type material listed below and
original material of Pseudococcus inamabilis)
(BMNH, MNNH, USNM, VCI, ZIL).
Pseudococcus vovae Nasonov. Lectotype Q,
U.S.S.R.: Varsoviensi Province, on Juniperus
communis, 24.vii.1906 (ZIL).

DIsTRIBUTION. Palaearctic Region: Cyprus,
England, Germany, Greece, Hungary, Iran,
Israel, Italy, Morocco, Poland, Turkey, U.S.S.R.
Neotropical Region: Brazil.

HostTpLants. Cupressaceae: Cupressus glauca, C.
gloveniana, C. macrocarpa, C. sempervirens, C.
virginiana, Juniperus communis, Thuja
occidentalis.

REMARKS. P. vovae is similar to P. taigae, but
differs in having fewer ducts on the dorsum, in
lacking ventral tubular ducts on the head, in hay-
ing more distinct ostioles, and by having more
flagellate dorsal and cerarian setae. Both of these
species occur only on Cupressaceae, as does the
similar Crisicoccus matesovae (Fig. 37).

P. vovae has variable numbers of dorsal tubular
ducts, as discussed by Cox & Ben-Dov (1986). In
specimens from the Mediterranean Basin, these
ducts occur in low numbers and may be confined
to the posterior abdominal segments. In those
from central Europe, however, these ducts are
much more numerous and occur over the entire
dorsum. The latter specimens resemble P. taigae,
from the eastern Palaearctic Region, which has
even more numerous dorsal ducts. Further work is
needed, based on many more specimens from cen-
tral Europe and the eastern Palaearctic Region, to
determine whether there is a single, variable spe-
cies, or a complex of species with different geo-
graphical distributions.

Ezzat & McConnell (1956) erected a new genus
Allococcus, with Pseudococcus inamabilis as the
type-species. Allococcus and Planococcus were
distinguished by these authors by the nature of the
dorsal tubular ducts, those of Allococcus having
rims, and those of Planococcus not having rims.

JENNIFER M. COX

However, dorsal tubular ducts with the
appearance of oral rims occur variably in some
individuals of several species of Planococcus,
including the type-species, P. citri. These ducts
are not as large, or with such pronounced rims, as
those found in other genera such as Pseudococcus.
Following the synonymy of Pseudococcus
inamabilis with Pseudococcus vovae and the
placement of this species in Planococcus, Allococ-
cus became a synonym of Planococcus. Most of
the species contained in Allococcus at that time
were transferred to a new genus, Delottococcus,
by Cox & Ben-Dov (1986).

Economic sTATUus. This species often reaches high
numbers on Cupressus used as shelter trees in
orchards, but is usually heavily parasitized.

Planococcus Zairensis sp. n.
(Fig. 40)

ADULT FEMALE. Mounted specimens broadly oval
to rotund, length 1.2-1.7 mm, width 0.7-0.9 mm.
Margin of body with a complete series of 18 pairs
of cerarii, each with 2, somewhat elongate, coni-
cal setae. Legs elongate; hind trochanter + femur
170-195 um long, hind tibia + tarsus 295-220 wm
long; ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.12—1.14; translucent pores
present on hind coxae and tibiae. Inner edges of
ostioles moderately sclerotized. Circulus small
and quadrate, width 40-55 um. Cisanal setae
shorter than anal lobe setae. Anal lobe cerarii
each situated on a small, moderately sclerotized
area.

Venter. Multilocular disc pores sparsely pre-
sent around vulva, 1—3 pores present on posterior
edges of median areas of abdominal segment VI,
and 1 pore sometimes present on each margin of
segments VI or VII. Trilocular pores sparsely but
evenly distributed. Oral collar tubular ducts of 1
size only, confined to abdomen, present in small
numbers on median areas of abdominal segments
V-VII and single ducts sometimes present on mar-
gins of segments VI or VII. Simple pores about
half the size of the trilocular pores, scattered over
entire venter.

Dorsum. Multilocular disc pores and tubular
ducts absent. Trilocular pores moderately numer-
ous and evenly distributed. Simple pores about
twice the size of the trilocular pores, present in
small groups on midline of head, thorax and ante-
rior abdominal segments. Setae short and stout,
longest seta of abdominal segments VI or VII
12-15 um long.

MEALYBUG GENUS PLANOCOCCUS

Fig. 40 Planococcus zairensis sp. n.

5

sop ACE

75

76

MATERIAL EXAMINED
Holotype 9, Zaire (Belgian Congo): Elis-
abethville [Lubumbashi], 1919 (Ringeot)
(MNHH).

Paratypes. 7 2 (1 on same slide as holotype),
same data as holotype (BMNH, MNNH).

DisTRiBuTION. Afrotropical Region: Zaire.
HostpLants. Unknown.

REMARKS. P. zairensis is similar to the orchid-
feeding dendrobii-group comprising P. dendrobii,
P. hosyni, P. orchidi and P. philippinensis in its
size, body shape, and distribution of multilocular
disc pores and tubular ducts. It differs from these
species in lacking knobbed dorsal setae. It is also
similar to P. boafoensis, but lacks the distinctly
protruding cerarii of this species.

REFERENCES

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Betrem, J. G. 1936. De Oecologie en Epidemiologie van de
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—— 1937. De Morphologie en Systematiek van Enkele van de
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— 1962. Notes on the Coccoidea of China. X. Descriptions of
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—— 1983. An experimental study of morphological variation in
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JENNIFER M. COX

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—— 1986. New data on the systematics of Coccinea
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1961. New Pseudococcidae (Homoptera: Coccoidea) from
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—— 1964. Observations on African mealy bugs (Hemiptera:
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—— 1967. The mealybugs of South Africa (Homoptera:
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MEALYBUG GENUS PLANOCOCCUS

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JENNIFER M. COX

INDEX

Invalid names are in italics; principal references are in bold.

aemulor 3, 4, 7, 8, 10, 59, 72
Allococcus 2, 7, 55, 72, 74
anaboranae 3

angolica 14

aphelus 3, 4, 8, 13

azaleae 5, 37, 52

boafoensis 3, 7, 13, 76

calceolariae 52, 55

celtis 2

citri 3, 4, 7, 9, 16, 17, 43, 45, 55, 74
citricus 2, 16, 17

citrioides 25

Coccus 16, 25

coleorum 16

congoensis 41, 43

crassus 3

Crisicoccus 2, 5, 37, 41, 68, 74
crotonis 45, 47

cubanensis 3, 16, 17
cucurbitae 3, 16, 17

Cyperia 14

Dactylopius 3, 16, 25, 28, 43, 45, 52,
Sp)5 I?

deceptor 4, 45

Delottococcus 2, 3

dendrobii 3, 4, 7, 8, 17, 34, 37, 59, 62,
76

destructor 16

dioscoreae 3, 5, 10, 20, 21, 48

dorsospinosus 3, 5, 10, 20, 21, 48

Dorthesia 16

drimydis 25

dubius 2, 7, 24, 25

epulus 3, 4, 7, 24, 25

ficus 2, 3, 4, 7, 9, 16, 17, 25, 28, 29, 32,
45

flagellatus 3, 4, 9, 29, 66, 68

formosus 3

furcisetosus 4, 5, 7, 29, 32

glaucus 25

halli 2, 4, 9, 28, 29, 32
hospitus 4, 7, 8, 20, 34, 37, 59, 62
hosyni 4, 8, 34, 37, 43, 59, 62, 76

inamabilis 7, 72, 74
indicus 47
ireneus 3

japonicus 4, 5, 8, 37, 41, 51, 52

kenyae 1, 2, 4, 7, 17, 41, 43, 47
kenyaensis 41
kraunhiae 4, 5, 8, 43, 45

lamabokensis 2, 3

Lecanium 16

lilacinus 2, 4, 5, 10, 41, 43, 45, 47, 68
lindingeri 3

litchi 4, 5, 10, 20, 48

mali 4, 5, 8, 41, 48, 51, 52
martini 4, 8, 52

matesovae 5, 68, 74
matsumotoi 5, 37
mauritiensis 45, 47

minor 2, 4, 9, 17, 52, 55, 66
morrisoni 4, 5, 8, 55, 57
mumensis 3

myrsinephilus 3, 21

nigritulus 4, 9, 10, 57, 59, 72
njalensis 3

orchidi 4, 8, 34, 37, 59, 62, 76

pacificus 4, 17, 52, 55

Paracoccus 25

pauliani 3

phenacocciformis 16

Phenacoccus 3

philippinensis 4, 7, 8, 20, 34, 37, 59, 76

phyllococcus 16

Planococcoides 3, 5

Planococcus 7

principe 4, 5, 8, 62

Pseudococcus 2, 4, 16, 25, 41, 43, 45,
47, 52, 72, 74

psidii 4, 5, 9, 62, 66

robustus 3
rotundatus 3

siakwanensis 4, 43, 45

sinensis 3, 16, 21

spiniferus 3, 16

spiriferus 3, 16, 17

subterraneus, Planococcus 4, 9, 29, 66,
68

subterraneus, Dactylopius 3, 25, 28

subukiaensis 4, 41, 43

sulawesi 4, 5, 10, 66, 68

taigae 4, 5, 10, 68, 74
tanzaniensis 4, 8, 59, 72
tayabanus 45, 47
Tylococcus 3, 13, 45, 47

vitis 25, 28
vovae 2, 4, 5, 10, 45, 68, 72, 74

zairensis 4, 8, 52, 74, 76
zealandicus 25

Bull. Br. Mus. nat. Hist. (Ent.) 58(1): 79-130

Issued 25 May 1989

The Simuliidae (Diptera) of the Santiago
onchocerciasis focus of Ecuador

A. J. SHELLEY', M. ARZUBE? & C. A. COUCH!
'Medical Diptera Section*, Department of Entomology, British Museum (Natural History), Cromwell

Road, London SW7 SBD, U.K.

Perez’, Guayaquil, Ecuador

_ *Departamento de Entomologia, Instituto Nacional de Higiene y Medicina Tropical ‘Leopoldo Izquieta

CONTENTS

Introduction ; : ; : 79
Material examined and terminology . 80

Systematics . ‘ ; i : : : ; ; : : i : 82
Keys to the simuliid species of the Santiago onchocerciasis focus ; . 82
Simulium (Notolepria) exiguum Roubaud : ‘ s 84
Simulium (Notolepria) gonzalezi Vargas & Diaz Najera. ; ; , 88
Simulium (Psilopelmia) bipunctatum Malloch. F , . : . 90
Simulium (Psilopelmia) lewisi Ramirez Perez. ‘ ; : : : 93
Simulium (Psilopelmia) escomeli Roubaud 97
Simulium (Hemicnetha) mexicanum Bellardi 100
Simulium (Psilopelmia) quadrivittatum Loew 104

References . 108

Index . 130

Synopsis Keys to the adults, larvae and pupae of the seven species of Simuliidae found in

the Santiago onchocerciasis focus of Ecuador are provided. Full morphological descrip-
tions of each of these stages are given for each species, together with notes on their
taxonomy, distribution and biology. Three new synonyms are established.

INTRODUCTION

The simuliid fauna of Ecuador had been only

_ Superficially studied up until 1980 when the dis-

covery of onchocerciasis in the north of the coun-
try (Arzube, 1981) emphasised the need for a
systematic investigation of the family. Previous
taxonomic work on Simuliidae in Ecuador was
either of a preliminary nature, consisting of notes
on and keys to the then poorly studied fauna in
papers that also covered the medical importance
of the family (Leon & Wygodzinsky, 1953a, b;
Levi-Castillo, 1956), or of a more detailed
revisionary nature on certain high altitude tem-

perate climate species of the Andes (e.g.
Wygodzinsky, 1971). No studies then existed on
the fauna of the lowland tropical forests of north-
ern Ecuador where onchocerciasis is now known
to occur. This paper provides the necessary bio-
systematic data on the Simuliidae of the Santiago
onchocerciasis focus that are prerequisites for
future longitudinal entomological surveys aimed
at providing baseline data for vector control. Only
three of the seven species recorded in the Santiago
onchocerciasis focus are significantly anthro-
pophilic and two of these, Simulium exiguum and
S. quadrivittatum, have been incriminated as vec-
tors of Onchocerca volvulus by Shelley & Arzube
(1985).

*World Health Organization Collaborating Centre for the study of Simuliidae and Phlebotominae in relation to

Onchocerciasis and Leishmaniasis

80

All species found in the Santiago onchocerciasis
focus belong to the genus Simulium and the sub-
generic assignments conform to those published in the

recent checklist of world blackflies (Crosskey, 1987).

Material examined and terminology

All the material studied from the Santiago
onchocerciasis focus was collected in 1981 during
investigations into the vector species of the dis-

Guayaquil

PERU

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

ease. Details of collection sites and the biology
and medical importance of the species in the area
have already been given by Shelley & Arzube
(1985); asummary of this information is given for
each species in the current paper. Data on the
precise location of the focus shown in Figs 1 and 2,
its topography, climate and inhabitants, are given
in Arzube (1982) and Guderian etal. (1983). Spec-
imens from the 1981 collection and subsequent

COLOMBIA

3 = foci of onchocerciasis

seeee = Province of Esmeraldas

boundary

Fig. 1 Distribution of onchocerciasis in Ecuador.

SIMULIIDAE OF ECUADOR

collections made in other regions of Ecuador with
which this material has been compared are deposi-
ted in the British Museum (Natural History),
London, U.K. and the Instituto Nacional de
Hygiene y Medicina Tropical ‘Leopoldo Izquieta
Perez’, Guayaquil, Ecuador. Material has been
conserved in the following ways: larvae, pupae,
individually reared adults with their correspond-
ing pupal pelts and females from biting catches —
in 80% alcohol; individually reared adults in asso-
ciation with their pupal pelts preserved in
glycerine in small polypropylene phials, and
females from biting catches —in pinned collection;
all stages — on microscope slides mounted in
Berlese mountant. Chromosomal preparations
from larval silk glands of some of the species are

Km
WeeeOme20" 630" 40) 50. 60-70" 80) 90 100
Esmeraldas
yes
2,
:
o
He %
ENS
co
RY Jb
~O @es a
o

Sapallo Grange, ey

81

stored on microscope slides with accompanying
photographs of some of the karyotypes in the
Diptera Section of the Entomology Department
of the British Museum (Natural History); larval
cadavers from which silk glands have been extrac-
ted are preserved in 80% alcohol and on micro-
scope slides. With the increasing need for larval
polytene chromosome analysis in taxonomic stud-
ies on Neotropical Simuliidae, preservation of
samples of larvae in Carnoy’s fixative must now
become routine. Carnoy’s fixative accentuates
and in some cases alters the colour pattern of
larvae compared to that seen in material pre-
served in alcohol. Where colour changes occur
these are noted at the head of the larval descrip-
tions. Descriptions of adult coloration for each

Vain te

Borbon “Bo '

ssion

jeg Yh; Ser 5p \
ae Sr a Ye Pe ire,
e Vy Vii th <; |
(

Reed

Gch i rmibetoiati: / /

eee /
Gay llabamb Lees

2
WS

—---- = Boundary of Esmeraldas Province
Fig. 2. The onchocerciasis foci of Esmeraldas Province, Ecuador.

Foci of Onchocerciasis

= Oil Palm Plantations

82

species have been based on pinned specimens
whereas photographs (Figs 138-153) are from
alcohol preserved material prepared in the follow-
ing way. Legs are removed from the body and
directly mounted on a microscope slide in Berlese
mounting medium. The wings are removed and
discarded and the head, thorax and abdomen are
placed in ‘Cellosolve’ for 12 hours, xylene for 6
hours and then air dried and mounted on a micro-
scope slide using a drop of ‘Araldite’ epoxy resin.
The specimens photographed have been pre-
served in alcohol for up to seven years and show
little or no colour changes from pinned material.
However, much of our alcohol material is teneral
and hence full adult coloration, particularly of the
legs, has not been attained. To obviate differences
between descriptions and photographs of each
species where possible, man-biting rather than
reared females have been photographed.
Terminology of structures in adults, larvae and
pupae referred to in the keys and species descrip-
tions follows Crosskey (in press). General figures
for male and female Simuliid genitalia not figured
in that work may be found in Crosskey (1969); the
terms coxite and style used by Crosskey are
replaced with gonocoxite and gonostyle respect-
ively. To avoid confusion with the less usual termi-
nology found in many descriptions of the Latin
American fauna the following equivalents are
given (terms used in the present paper cited first):

nudiocular area (of female head) = fronto-ocular
triangle;

paramere = endoparameral organ;

gonostyle = distimere;

gonocoxite = basimere.

The following acronyms are used for depositories
of specimens referred to in this paper.

BMNH British Museum (Natural History), London,
U.K.

DBAT Dipartimento di Biologia Animale, Univer-
sita di Torino, Torino, Italy

DDSV _ Seccion de Oncocercosis, Division de Der-
matologia Sanitaria, Villa de Cura, Ara-
gua State, Venezuela

DERM Laboratorio de Entomologia de la Division
de Endemias Rurales, Maracay, Aragua
State, Venezuela

INHMT Instituto Nacional de Hygiene y Medicina
Tropical ‘Leopoldo Izquieta Perez’,
Guayaquil, Ecuador

IOC Instituto Oswaldo Cruz, Rio de Janeiro,
Brazil

MCZH Museum of Comparative Zoology, Harvard,
U.S.A.

MLP Museo La Plata, La Plata, Argentina

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

MNHN Muséum National d’Histoire Naturelle,
Paris, France

STMPR_ Department of Microbiology, School of Med-
icine, School of Tropical Medicine, San
Juan, Puerto Rico

USNM _ United States National Museum,
Washington, D.C., U.S.A.

ZM Zoologisches Museum der Humboldt Univer-
sitaet, Berlin, West Germany

ACKNOWLEDGEMENTS. We are indebted to the

UNDP/World Bank/WHO under their Special Pro-
gramme for Research and Training in Tropical Diseases,
and the Pan American Health Organization for financial
assistance, to Dr V. Reyes, Director of the ‘Servicio
Nacional de Erradicacion de Malaria (SNEM), Minis-
terio de la Salud Publica’, Ecuador, for providing river
transport, and to USAID for a visit to the BMNH by the
second author. Drs J. Alarcon & E. Gomez and Messrs
R. Llerena, I. Mera and A. Villavicencio are gratefully
acknowledged for their technical assistance. Drs L.
Matile, B. V. Peterson and E. O. Wilson kindly loaned
specimens from the collections of the MNHN, USNM
and MCZH respectively, and Dr R. W. Crosskey made
valuable suggestions that have been incorporated in this
paper. Miss S. Allington provided the illustrations and
Mr P. York the photographs

SYSTEMATICS

Keys to the simuliid species of the Santiago
onchocerciasis focus

Females
I aScutuml Orange. of tase cus pete seen 2
= 4Scutummblack:... fe ohalsos5:5.0 esc oer eee 4

2 Scutum with white pruinose pattern dorsally. Abdo-
men without yellow terpites, 30.4. eee 3
— Scutum without white pruinose pattern dorsally.
Abdomen with first three (rarely four) tergites
yellow (Fig. 141). [Zoophilic] ............. lewisi

3 Scutum with pair of submedian greyish white bands
running from anterior border for four fifths of scutal
length. Postnotum black with silver pruinosity.
Abdomen with dorsal chequerboard pattern of
prominent black markings on greyish background
(Fig. 142). [Anthropophilic] ............ escomeli

— Scutum with pair of submedian white pruinose
comma-shaped bands beginning at posterior border
of humeri and running half length of scutum.
Postnotum orange. Abdomen dark brown with ante-
rior segments orange (Fig. 140). [Mainly zoophilic]

bipunctatum

4 Scutum velvet-black with silver pruinose pattern for-
med by a pair of submedian longitudinal silver pru-
inose bands reaching silver pruinose hind margin and
a pair of sublateral silver pruinose bands. Abdomen
black dorsally with transverse silver pruinose bands

SIMULIIDAE OF ECUADOR

on segments of anterior three-quarters (Fig. 145).
[Anthropophilic] quadrivittatum
Scutum shiny black without silver pattern. Abdomen
black without transverse silver pruinose bands .. 5

Scutal hairs yellowish white. Legs mainly dark; fore
tarsi dilated (Fig. 144). Basal section of wing vein R
with two or three irregular rows of hairs; Sc with
single row of hairs (Fig. 41). [Zoophilic]
mexicanum
Scutal hairs golden or green (depending on light
incidence). Legs mainly pale; fore tarsi not dilated
(Figs 138, 139). Basal section of wing vein R and Sc
BUMELTON MER MARES Slee deo) cid hays ous) «ii elear eyeraiss« Srermiene: ai aisls 6

Paraproct with long anterior process (Fig. 55).
[Anthropophilic] exiguum
Paraproct with short anterior process (Fig. 56).

PApapniiclitic ke x mneyrgeie rans Says onder wrs gonzalezi
Males
fetiead dichoptic (Fig. 10) ...........5.. gonzalezi
Eaettcad holoptic (Figs 8,11) ...6......ceeecenes 2

2

Scutum orange or brown. Scutellum orange or
Sis COURTS Srey ceaeeYaiafolel alnss Sel «Aste Wier vials} Pins 3
Scutum and scutellum black

Scutum orange or brown with a pair of silver or black
submedian bands in anterior half. Postnotum orange

BEM CHM GMM Ae cy: cr. o's tek sala tele solute cvarate: boats Nowle le. 4
Scutum orange without dorsal ornamentation.
Postnotum black (Fig. 149) ............... lewisi

Scutum orange with silvery white pruinose subme-
dian bands touching anterior border (most obvious
with anterior illumination). Postnotum orange (Fig.
148). Vestiture of scutum of evenly distributed black
hairs. Basal section of wing vein R with single row of
LRCLITES. sc ey ive Ea Cage ROR OM RE bipunctatum
Scutum orange to mid-brown with silver pruinose
submedian bands touching anterior border using
anterior illumination; bands comma-shaped and
black, not touching anterior scutal border with pos-
terior illumination. Postnotum black with grey pru-
inosity (Fig. 150). Vestiture of scutum of evenly
distributed golden hairs. Basal section of wing vein R
RMN PIR Ty evatte = giv ais ad woe» «, Scone escomeli

Scutum with a pair of silver pruinose submedian
cunae in anterior third of scutum touching anterior
border (Fig. 153) quadrivittatum
Scutum without ornamentation

Scutum greyish black with velvet-black median line
extending along its entire length; anterior and pos-
terior scutal borders not pruinose. Scutum with
dense vestiture of long-brass coloured setae. Legs
mainly black; tarsi broad and flattened (Fig. 152).
Large robust fly, 3.0-4.5 mm long (preserved in
RCSLT ie oy ctaycfnns sack arash csyou sain cia ehaadsicsae mexicanum
Scutum velvet-black; anterior and posterior scutal
borders with silver pruinosity. Scutum with dense
vestiture of short golden setae with green reflections.
Legs mainly yellow or light brown with narrow
unflattened tarsi (Figs 146, 147). Small fly 1.3-2.1
mm long (preserved in alcohol)

7

Pupae

83

Femur of mid leg dark brown to black (Fig. 147).
gonzalezi
Femur of mid leg yellow (Fig. 146) ...... exiguum
Gill with six filaments (Fig. 110) ....... gonzalezi
Gill with more than six filaments .............. 2,
Gill with twelve filaments (Fig. 115) .. mexicanum
Gili withieishitilamentsys a-n0seei-1-8 wi sera 3

Cocoon white to light brown, thick walled and witha
longitudinal dorsal ridge. Gill with most distal bifur-
cations of dorsal and median primary branches at
mid point (Fig. 106) lewisi
Cocoon brown to grey, thin-walled and without dor-
sal ridge. Most distal bifurcations of dorsal and
median primary branches within basal third of gill 4

Gills longer than length of ventral surface of cocoon
and with spaced secondary branching (Fig. 116)
quadrivittatum
Gills shorter than length of ventral surface of cocoon
and with secondary branches of dorsal and median
primary branches arising close to one another .. 5

Gill with secondary branching of all primary gill
branches in basal eighth (Fig. 112) ... bipunctatum
Gill with most distal secondary branches of median
and ventral primary branches within basal sixth to
third

Abdominal sternite TV with 2 + 2 fine hairs on
POStEMor mar pil Peres steels ee os arote wie foals exiguum
Abdominal sternite [TV with 1+1 outer simple or

bifid hooks and 1+1 inner fine hairs ..... escomeli
Mature larvae
1 ‘Body length 6.9-9.7 mm... ......0%. mexicanum
=eBodyilensth2:8-s1Siamimi iat a te rele 2

2

Anterior abdominal segments white to grey with two
prontinent darkibands #9700... oe e ssc ve teu see

Anterior abdominal segments white to grey with a
pale or dark band on each segment ............ +

Postgenal cleft small and pointed anteriorly; postge-
nal bridge as long as hypostomium (Fig. 137). Ven-
tral papillae well-developed (Fig. 122)
quadrivittatum
Postgenal cleft large and rounded anteriorly; postge-
nal bridge about one-tenth the length of hypo-
stomium (Fig. 134). Ventral papillae absent or very
small (Fig. 119) lewisi

Abdominal marking dark and very prominent .. 5
Abdominal marking light and indistinct ........ 6

Head capsule without pattern (occasionally a posi-
tive head pattern or an amorphous darker area in
region of head spots occurs) (Fig. 123) .. exiguum
Head capsule with negative pattern (occasionally
without pattern) (Fig. 124) ............ gonzalezi

Postgenal bridge as long as hypostomium (Fig. 135).
Ventral papillae absent (Fig. 120). Anal gills usually
with three simple lobes but outer pair sometimes
with up to two additional secondary lobules
escomeli

84

— Postgenal bridge one-third the length of the hypo-
stomium (Fig. 133). Ventral papillae small (Fig.
118). Anal gills with 6-8 secondary lobules on each
PEUMATV NODES see spree cere aK bipunctatum

Simulium (Notolepria) exiguum Roubaud

(Figs 3, 5, 7, 8, 12-14, 17, 21, 23, 24, 25, 31, 37, 43,
49, 55, 62, 69, 75, 76, 90, 103, 110, 117, 123, 131,
138, 146)

Simulium exiguum Roubaud, 1906: 108. LEC-
TOTYPE Q, VENEZUELA: Haut Sarare, 1899
(F. Geay) (MNHN), here designated.
[examined]

Simulium glaucophthalmum Knab, 1914b: 123.
Holotype 9, Peru: Santa Clara, iv.1914 (C. H.
T. Townsend) (USNM Cat. No. 18494) [exam-
ined]. [Synonymy by Wygodzinsky, 1951: 214.]

Simulium delpontei Paterson & Shannon, 1927:
742. Holotype 9, ARGENTINA: Salta Province,
Embarcacion, 4.v.1926 (Paterson, Shannon &
Shannon) (depository unknown). [Synonymy
by Wygodzinsky, 1951: 214.]

Description. Female. General body colour black.
Body length (alcohol preserved specimens) 1.8—
2.7mm (n = 24), wing length 1.4-2.0mm (n= 21),
wing width 0.7-0.9 mm (n = 19).

Head dichoptic (Fig. 7) with dark red eyes
showing green highlights; nudiocular area absent
(Fig. 25). Frons, clypeus and occiput black with
silver pruinosity. Mouthparts orange-brown.
Antennae dark brown with scape, pedicel and first
flagellomere orange-brown. Cibarium unarmed
(Fig. 31).

Scutum and humeri greyish black with faint sil-
ver pruinosity; one median and a pair of posteri-
orly divergent submedian darker black lines
running along whole length of scutum (best seen
in specimens devoid of setae and with illumination
perpendicular to specimen); scutum also with
small velvet-black spot adjoining paranotal folds
(= paratergites). Paranotal folds black with silvery
grey pruinosity. Scutum with numerous short
adpressed dark setae and discrete groups of short
flattened brass-coloured setae with greenish
reflections (Fig. 138). Pleural region silvery grey
pruinose. Scutellum greyish black and faintly pru-
inose, its vestiture comprised of golden setae
longer than those on scutum and a single row of
black bristles on posterior margin. Postnotum sil-
very grey pruinose.

Subcostal wing vein and basal sector of radius
bare (Fig. 37). Costal base tuft of dark hairs.

Legs yellow to light brown, except fore tarsi,
mid and hind coxae, hind femora and distal three-
fourths of hind tibiae black. All femora and tibiae

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

with scales (Fig. 3). Proportions of legs as in Fig.
138. Claws curved and slender, without basal
tooth on fore and mid legs but with poorly
developed tooth on hind leg (Fig. 43). Halteres
pale yellow with light brown stem.

Abdominal tergites shiny brownish black with
silver pruinosity on second segment (Fig. 138).
Tergal plates well-developed, pattern as in Fig. 4.
Sternites brownish black; genitalia light brown.
Eighth sternite highly sclerotised with a group of
8-12 stout setae in each sclerotised portion (Fig.
49); gonopophyses small, sclerotised on in-
ner margin, glabrous. Cerci hemispherical, light
brown; paraprocts broadly rectangular with
pointed anteriorly directed process (Fig. 55).
Genital fork (Fig. 62) slender, with sclerotised
anteriorly directed processes and stem. Sper-
matheca oval, highly sclerotised, with no external

sculpturing and spicules of inner surface obscured *

by sclerotisation; width of membranous area of
insertion of spermathecal duct large, about half
maximum width of spermatheca (Fig. 5).

Male. General body colour black. Body length
(alcohol preserved specimens) 1.9-2.1mm(n=6),
wing length 1.5—-1.7 mm (n = 5), wing width 0.7—
0.8mm (n=S).

Head holoptic (Fig. 8) with dark red eyes;
lower, smaller facets with greenish reflections.
Clypeus black with silver pruinosity, other head
coloration as in female.

Scutum and humeri velvet-black with anterior
and posterior margins and anterior two-thirds of
lateral margin silver pruinose (Fig. 146). Parano-
tal folds velvet-black with silvery grey pruinosity.
Scutum covered in numerous, short, adpressed,
light brown setae interspersed amongst groups of
brilliant gold, scale-like setae. Coloration and set-
ation of pleural region, scutellum and postnotum
as in female except scale-like setae on scutellum
brilliant gold.

Wing venation, leg coloration (Fig. 146) and
haltere coloration as in female.

Abdominal tergites velvet-black, basal fringe
dark brown with few long hairs. Silver ornamenta-
tion as follows: tergites II and VI all silver except
sometimes in median area on VI; tergite VII all
silver except for median area, some specimens
show a pair of lateral silver pruinose patches on
posterior margin of tergite VIII. Tergite [IX shiny
black (Fig. 146). Sternites grey with well-
developed velvet-black sternal plates on segments
II-VIII (Fig. 6). Genitalia brownish black.
Gonocoxite subrectangular, gonostyle small, sub-
triangular, one-third as long as gonocoxite and
with small distal spine (Fig. 69). Ventral plate with
reduced basal arms, lightly sclerotised, triangular

SIMULIIDAE OF ECUADOR

with small keel, hairs short, diffuse and mainly
occurring around median keel (Figs 75, 76).
Median sclerite subrectangular with deep apical
incision (Fig. 90). Paramere with several apical
spines (as shown in Fig. 97 for S. gonzalezi).

Pupa. Cocoon length dorsally 1.6-2.6 mm, ven-
trally 2.2-3.2 mm; pupa length 1.7—-2.3 mm; gill
length 1.3—-2.1 mm (n = 27).

Cocoon slipper-shaped, mid to dark brown; rim
of aperture dark brown, reinforced and without
central protuberance (Fig. 103). Cocoon com-
posed of elastic, amorphous substance interwoven
with fibres. Gill light brown with eight forwardly
directed slender filaments arranged in the vertical
plane (Fig. 110), main trunk giving rise to three
primary branches, ventral with two filaments and
median and dorsal each with three filaments; ven-
tral branch with bifurcation in basal fourth of gill,
median branch with first bifurcation in basal
fourth and second bifurcation in basal third of gill;
dorsal branch with first bifurcation basally at junc-
tion of median and dorsal primary branches and
second bifurcation within basal fourth of gill; fila-
ments slender with crenate margins and rounded
distally, their surfaces covered in fine spicules
(Fig. 12). Head (frontoclypeus) with 2 + 2 frontal
trichomes of which the more dorsal pair is simple
and poorly developed and the more ventral pair
well-developed and 2-5 branched, and | + 1 well-
developed facial trichomes with 2-5 branches;
surface of head covered with platelets (Figs 13,
14). Thorax with 5 + 5 antero-dorsal, well-
developed trichomes of 2—S branches. Surface of
thorax covered with platelets which are more
densely distributed on anterior half (Fig. 17).
Abdominal tergite II with 4 + 4 simple hairs in a
line on posterior border of segment, III-IV with 4
+ 4 simple hooks, VI-IX with patches of poorly
developed spine combs on antero-lateral margins,
IX with 1 + 1 strong, unbranched spines (Fig. 19);
sternite [V with 1 + 1 simple hairs, V with 2 + 2
bifid or trifid hooks, VI and VII with 2 + 2 hooks,
the inner pairs being bifid or trifid and the outer
pairs simple; 1 + 1 patches of poorly developed
spine combs on postero-lateral borders of ster-
nites [V-VIII (Fig. 20).

Mature larva. Body length 3.64.4 mm (n = 33).
Width of head capsule 0.4-0.5 mm (n = 32). Body
usually white with greyish brown markings (Fig.
117), occasionally almost completely white and
showing only indistinct grey pigmentation. Color-
ation in Carnoy’s fixative is similar except that the
banding patterns are more distinct. Body form as
in Fig. 117.

Head yellow with head spots concolorous; occa-
sional specimens with positive head spot pattern

85

as in Fig. 123 or with an amorphous dark area in
head spot region that obscures the spots. Head
capsule with few, randomly distributed setae on
all surfaces. Postgenal cleft as wide as long,
rounded anteriorly; postgenal bridge about half as
long as hypostomium (Fig. 131). Hypostomium
with strongly pigmented anterior margin and nine
apical teeth: corner teeth large and blunt, median
tooth less developed but larger than subequal
intermediate teeth; 4-5 lateral serrations with
hindmost about level with first hypostomial seta; 1
+ 1 groups of four hypostomial setae lying parallel
to lateral margins of hypostomium; surface of
hypostomium with a few short setae (Fig. 21).
Antennae long, unpigmented with segment ratios
23 : 10 : 12. Mandible (Fig. 23) with first three
comb teeth decreasing in size posteriorly and two
mandibular serrations of which the anterior is
larger. Maxillary palp about twice as long as
breadth at base. Cephalic fan with 30—43 rays (n =
10).

Thorax white with grey ring around anterior
region and grey pattern dorsally of variable form
that almost covers whole area, ventrally with two
or three central patches of grey chromatocytes
posterior to proleg. Cuticle with occasional small
setae dorsally, glabrous ventrally. Proleg plates
lightly sclerotised and with about six processes.
Pupal respiratory histoblast dark brown, clavi-
form.

Abdomen white with a single complete black or
grey ring on each of the four anterior narrow
segments; posterior segments grey or black dor-
sally, white with variable scattered black chro-
matocytes ventrally. Ventral nerve cord black.
Ventral papillae absent. Cuticle with minute hairs
dorsally in dark areas, ventral surface glabrous.
Anterior perianal area with a group of fine spines.
Anal sclerite well sclerotised with posterior arms
extending to twelfth row of posterior circlet
hooks. Posterior circlet with 70-78 rows of 2-14
hooks (n = 8). Anal gill trilobed, each lobe with
6-8 long finger-like lobules (Fig. 24).

MATERIAL EXAMINED

Lectotype 9, paralectotypes 5 Q of S. exiguum,
Venezuela: Sarare, 1899 (F. Geay) (BMNH,
MNHN). Holotype 9 of S. glaucophthalmum,
Peru: Santa Clara, 1914 (C. H. T. Townsend)
(USNM).

Argentina: 3 2, Jujuy, xii.1938 (W. C. Pater-
son) (BMNH).

Bolivia: 2 2, Covende, 1921-22 (W. M. M.
Mann) (BMNH); 1 9, HuachiBeni, viii.1921—22
(W. M. M. Mann) (BMNH).

Brazil: Roraima Territory: 1 9, Posto Meva,
Rio Auaris, 3.iv.1977 (R. R. Pinger) (BMNH); 1

86

2,15 9 6C (ex pupae), Catrimani Mission, Rio
Catrimani, 9.i.1977 and 12, 13.i.1979 (A. J.
Shelley, and A. J. Shelley & A. P. A. Luna Dias)
(BMNH); 1 9 1 oO (ex pupae), Rio Uraricoeira,
20.1.1979 (A. J. Shelley & A. P. A. Luna Dias)
(BMNH); 3 9 2 CO’ (ex pupae), Northern Perime-
ter Road, Rio Agua Preta, 18.xi.1980 (A. J.
Shelley & A. P. A. Luna Dias); 1 9 1 Oo (ex
pupae), Mucajai Mission, Rio Mucajai, 6.i1.1977
(A. J. Shelley) (BMNH); 1 ©, Normandia,
Igarapé Inamaru, 3.xii.1980 (A. J. Shelley & A. P.
A. Luna Dias) (BMNH); 1 2, 1 9 5 0’ (ex pupae),
Vila Pereira, Rio Surumu, 25, 26.xi.1980 (A. J.
Shelley & A. P. A. Luna Dias) (BMNH);2292¢
(ex pupae), nr Bonfim, Rio Arraia, 3.xii.1980 (A.
J. Shelley & A. P. A. Luna Dias); Amazonas
State: 11 2, Mission Post, Rio Toototobi, 16. viii.
and 24, 25.x.1976 (R. R. Pinger) (BMNH); 6 9,
Rio Ituxi, v.1978 (D. Roberts) (BMNH);6 930
(ex pupae), Mato Grosso, Rio Aripuana,
29.vi.1978 (J. D. Charlwood) (BMNH).

Colombia: Sierra Nevada de Santa Marta: 3 9,
Ariguani, 20.xii.1963 (J. P. Lee-Potter) (BMNH);
1 2, Meollaca, 22.xii.1963 (J .P. Lee-Potter),
2 9, Valledupar, 15.i.1963 (J. P. Lee-Potter)
(BMNH); 2 9, Department of Valle, Tunselas,
16.11.1977 (J. Ardila) (BMNH).

Ecuador: numerous man-biting females, reared
adults, pupae and larvae from the following
localities in the Santiago onchocerciasis focus
in Esmeraldas Province: R. Cayapa, San Miguel
de Cayapas, 18-21.vi.1981 (A. J. Shelley &
M. Arzube) (BMNH, INHTM); R. Grande
(Cayapa), Viruela and Calle Mansa, 26-27.v.1981
(A. J. Shelley & M. Arzube) (BMNH, INMHT);
R. Sapallo Grande, Tumbaviro, 18.vi.1981 (A. J.
Shelley & M. Arzube) (BMNH, INHTM); numer-
ous man-biting females, reared adults and larvae
from the Canandé peripheral onchocerciasis focus
in Esmeraldas Province, R. Canandé, Naranjal,
25.ix.1983 and 21—24.vi.1985 (A. J. Shelley & M.
Arzube) (BMNH, INHMT); 1 @, 1 (ex pupa), 1
larva, Esmeraldas Province, nr Concordia, road
to Puerto Quito, R. Caoni, 24.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH); 2 9 1 Oo (ex
pupa), nr Concordia, road to Puerto Quito, Rio
Salazar, 28.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH, INMHT); 5 9 3 CO (ex pupae), nr Con-
cordia, road to Puerto Quito, R. Blanco,
28.ix.1983 (A. J. Shelley & M. Arzube) (BMNH);
1 9 (ex pupa), Santo-Domingo-Esmeraldas road,
Rio Sapotal, 26.ix.1983 (A. J. Shelley & M.
Arzube) (BMNH); 1 9, Imbabura Province, Sali-
nas road, R. Tahuando, 8.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH); 3 9,5 9 3 OC (ex pupae),
Salinas-Lita road, San Juan del Hacha, R. Mira,
11.ix.1983 (A. J. Shelley & M. Arzube) (BMNH);

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

1 (ex pupa), Salinas-Lita road, R. San Pedro,
11.ix.1983 (A. J. Shelley & M. Arzube) (BMNH);
13 2, 16 9 12 CO (ex pupae), Napo Province, nr
Lago Agrio, Rio San Miguel, 9.xii.1982 (A. J.
Shelley & M. Arzube) (BMNH); 1 9, nr Lago
Agrio, Posto 19, R. Tarapa, 9.xii.1982 (A. J.
Shelley & M. Arzube) (BMNH); 1 9, nr Lago
Agrio, Rio Teteye, 9.xii.1982 (A. J. Shelley & M.
Arzube) (BMNH); 15 9°, 8 9, 6 & (ex pupae),
nr Lago Agrio, Dureno, R. Aguarico,
10-13.xii.1982 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); 8 Q, nr Tena, Misahualli, R.
Napo, 7.vi.1985 (A. J. Shelley & M. Arzube)
(BMNH); 6 Q, 1 9 (ex pupa), El Oro Province,
Machala-Naranjal road, canal de riego, Rio
Bucay, 12.vi.1984 (A. J. Shelley & M. Arzube)
(BMNH); 4 2, Machala-Uzcurume road, nr Pas-
aje, Rio Guesha, 20.vi.1984 (A. J. Shelley & M.
Arzube) (BMNH); 1 @ (ex pupa), Machala-
Uzcurume road, near Pasaje, Rio Jubones,
20.vi.1984 (A. J. Shelley & M. Arzube) (BMNH);
22,4920 (ex pupae), Machala-Pinas road, Rio
Carne Amarga, 21.vi.1984 (A. J. Shelley & M.
Arzube) (BMNH); 2 9, Pastaza Province, Tena-
Puyo road, Rio Mira Valle, 8.vi.1985 (A. J.
Shelley & M. Arzube) (BMNH); 1 9, Puyo, Shell-
Mera road, confluence of Rios Alpayacu and Pas-
taza, 10.vi.1985 (A. J. Shelley & M. Arzube)
(BMNH); 6 2,5 9 2 Oo (ex pupae), Cotopaxi
Province, Quevedo-La Mana-Pilalo road, Rio
San Pablo, 8.vi.1984 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); 1 CO (ex pupa), Quevedo-La
Mana-Pilalo road, Recinto Beles, Rio San Pablo,
8.vi.1984 (A. J. Shelley & M. Arzube) (BMNH); 6
Q,1C (ex pupa), Manabi Province, Santo Domi-
ngo-El Carmen road, Rio Suma, 7.vi.1984 (A. J.
Shelley & M. Arzube) (BMNH); 2 @, Los Rios
Province, Babahoyo-Montalvo road, Rio Cristal,
10.vi.1984 (A. J. Shelley & M. Arzube) (BMNH);
3 9, Bolivar Province, Babahoyo-Balzapamba-
Aguaranda road, Rio Chanpiaco, 10.vi.1984 (A.
J. Shelley & M. Arzube) (BMNH); 39 9,2 92c
(ex pupae), Guayas Province, Naranjal-Machala
road, Via Cooperativa 11 de Agosto, Rio Bucay,
19.vi.1984 (A. J. Shelley & M. Arzube) (BMNH);
14 9, 2 Q (ex pupae), Loja Province, Loja-La
Toma road, Hacienda Monterey, Rio Guayabal,
23.vi.1984 (A. J. Shelley & M. Arzube) (BMNB).

Venezuela: numerous man-biting females from
the Caripe onchocerciasis focus, Monagas State,
San Antonio de Maturin (nr Caripe) and Caripe
area, 7.iv.1981 and iv.1981 (D. J. Lewis)
(BMNH); 3 2 5’ (ex pupae), Yaracuy State, Rio
Yaracuy (no collection date) (J. Ramirez Perez)
(BMNH); 1 @ with associated pupal case,
Miranda State, Tacata (no collection date) (J.
Ramirez Perez) (BMNH); 1 2 2 CO (ex pupae),

SIMULIIDAE OF ECUADOR

Trujillo State, Carache (no collection date) (J.
Ramirez Perez) (BMNH).

TAXONOMIC DISCUSSION. Simulium exiguum was
described by Roubaud from a series of females
(one specimen pinned and an undisclosed number
preserved in alcohol) sent from Sarare [presum-
ably the upper reaches of the R. Sarare in Arauca
State in northern Venezuela, where S. exiguum is
commonly found today] by F. Geay. No type des-
ignations were made in the paper so the specimens
have syntype status. Vargas (1945) incorrectly
refers to their status as a type [= holotype] in the
MNHN and a probable paratype in the BMNH.
Six pinned specimens from the original material
have been located in the BMNH and MNHN,
each bearing a printed label of the Paris Museum
as follows: ‘Museum Paris, Venezuela, Sarare,
Geay 1899’. One of the five specimens in the
MNHN bears a ‘type’ label and a handwritten
label with the following details ‘S. exiguum Rou-
baud n.sp. Bull.Mus. 1906’. This specimen is in
relatively good condition, is here designated lec-
totype and has been labelled accordingly. The
other four specimens in the MNHN appear to
have been recovered from alcohol and have been
labelled as paralectotypes. The single specimen in
the BMNH had been previously recovered from
alcohol and although in poor condition is recog-
nisable as S. exiguum. It bears two labels in Aus-
ten’s hand indicating the collection locality and
that it was donated by Roubaud, and it has now
been labelled as a paralectotype.

In Wygodzinsky’s revision (1951) of S. exiguum
two nominal species, S. glaucophthalmum Knab
collected in Peru and S. delpontei Paterson &
Shannon from Argentina, are synonymised with
S. exiguum. His synonymy of S. glaucophthalmum
was based on an examination of types and Knab’s
description (1914b) but no reasons were given for
the synonymy of S. delpontei with S. exiguum. We
here confirm the former synonymy following dis-
section of the S. glaucophthalmum holotype and
comparison of its paraprocts with those of S.
exiguum. Although Paterson & Shannon’s de-
scription (1927) of S. delpontei clearly indicates
it to be within the subgenus Notolepria its syn-
onymy with S. exiguum or the closely related S.
gonzalezi or S. paraguayense may only be clarified
once the form of its paraproct is known. Coscaron
& Wygodzinsky (1975) continue to cite S.
delpontei as a synonym of S. exiguum in a paper
devoted to comparing the latter with S. para-
guayense. We also recommend that the value of
the characters, principally the female paraproct,
used for the separation of S. paraguayense, S.
exiguum and S. gonzalezi are investigated in con-

87

junction with chromosomal analyses in ‘genet-
ically monomorphic’ populations in order to
clarify the taxonomic status of these three species.

Simulium exiguum is one of the more com-
monly encountered anthropophilic South Amer-
ican simuliids and hence has been well de-
scribed and cited in numerous publications. The
most complete taxonomic works on the species
are Wygodzinsky (1951) and Coscaron &
Wygodzinsky (1975). Simulium exiguum is the
type species of the subgenus Nofolepria and is
closely related to S. paraguayense from southern
South America, from which it is distinguished by
adult leg coloration and the morphology of the
male and female genitalia (Coscaron &
Wygodzinsky, 1975). Another close relative, S.
gonzalezi Vargas & Diaz Najera, occurs sym-
patrically in Ecuador with S. exiguum from which
it may be distinguished by adult leg coloration, the
form of the female paraproct and the number of
pupal gill filaments. Takaoka (1983) confirmed
Dalmat’s finding in 1955 that in Guatemala this
latter character is variable within S. gonzalezi and
cannot therefore be used for accurate species
determination there. Similar observations on the
form of the paraproct for distinguishing S.
exiguum from S. gonzalezi were made by Lewis &
Lee-Potter (1964). They noted differences in the
paraproct of S. exiguum from the Sierra Nevada
de Santa Marta in northern Colombia compared
with typical S. exiguum from northern Venezuela.
Lewis (1963) also refers to the presence of eight-
filamented pupae of S. gonzalezi in Antigua but
no material has been found in the BMNH collec-
tions. In Ecuador separation of S. exiguum and S.
gonzalezi has been based on the form of the para-
proct in females in conjunction with the number of
filaments in the gill of the pupa. No variation in
gill filament number was observed, pupae with
eight-filamented gills always producing females
with S. exiguum type paraprocts and those with six
filaments females with S. gonzalezi type para-
procts. Adult females collected biting man and
horses were identified as exiguum or gonzalezi
based on paraproct form, and similar identifica-
tions of males were based on the gill form of pupae
from which they were reared.

Only male S. exiguum with holoptic heads were
recorded in the Santiago onchocerciasis focus but
a dichoptic form occurs in Venezuela (Vargas &
Diaz Najera, 1953b; Lewis 1963) and Darien
Province, Panama (material collected by Dr J.
Petersen and examined by senior author).

The recent discovery (Procunier et al., 1985 and
Procunier, pers. comm.) that S. exiguum is a com-
plex of at least four sibling species in Ecuador now
suggests the need for integrated morphological

88

and cytological studies when investigations on the
biology and medical importance of the sibling spe-
cies in South America are made.

DisTRIBUTION. In Ecuador S. exiguum is found in
most provinces on both the east and west of the
Andean cordillera. It occurs at both the main
onchocerciasis focus in the Santiago river basin
and at the peripheral focus at the R. Canandé.

Simulium exiguum occurs in the following
South American countries: Argentina, Bolivia,
Brazil, Colombia, Peru and Venezuela. It has also
been recently collected in southern Panama (Dr J.
Petersen, pers. comm.). Records of its occurrence
in other Central American countries, especially
Guatemala and Mexico, need to be verified
because in many cases these refer to S. gonzalezi
(Vargas & Diaz Najera, 1953b; Dalmat, 1955;
Takaoka, 1983). Further details of its distribution
and bionomics may be obtained in Shelley (in
press b).

Brotocy. In Ecuador S. exiguum is the most com-
mon man-biting species in the lowland tropical
forests of the eastern and western slopes of the
Andes, where, typically at altitudes of 100-500 m,
high biting rates of up to 2200 flies/man/day occur
in some localities (Procunier et al., 1985; Shelley
& Arzube, 1985, unpublished data); in these areas
horses, when present, are also favoured as a blood
source. Similar host preferences have been noted
for this species in northern Venezuela in an area
where it is a suspected sporadic vector of
onchocerciasis (Lewis & Ibanez de Aldecoa,
1962) and in Colombia where it attacks man,
horses and cattle in large numbers (Guttman,
1972), showing a preference for animal hosts in
some localities (Trapido et al., 1971). Simulium
exiguum bites man in large numbers in Argentina
(Coscaron & Wygodzinsky, 1975) but in contrast
it may be almost entirely zoophilic in some
localities in northern Brazil (Shelley, 1988). In
Bolivia it has been recorded biting tapirs (Vargas
& Diaz Najera, 1953b).

Apart from being a biting nuisance, S. exiguum
is a primary vector of onchocerciasis in Ecuador
(Shelley & Arzube, 1985) and Colombia (Tidwell
et al., 1980) and a suspected sporadic vector in
northern Venezuela (Lewis & Ibanez de Aldecoa,
1962). A review of its biology in relation to
onchocerciasis transmission in Latin America may
be found in Shelley (1988). In Colombia, S.
exiguum is also suspected of being involved in the
transmission of Venezuelan Equine Encephalitis
(Sanmartin et al., 1973), although more recent
work (Homan et al., 1985) showing no viral rep-
lication in S. metallicum Bellardi and S. mex-

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

icanum also suggests that S. exiguum may only be
involved in mechanical transmission.

Simulium exiguum favours open, sunlit rivers
from 5-100 m wide for breeding grounds in
Ecuador; larvae are found on fallen leaves and
submerged tree branches in shallow shingle beds
as well as in deeper parts of the river. Large rivers
are the breeding sites of this species in other coun-
tries of South America (Coscaron & Wygo-
dzinsky, 1975).

Simulium (Notolepria) gonzalezi Vargas &
Diaz Najera

(Figs 9-11, 56, 63, 77-79, 91, 97, 104, 111, 124,
132, 139, 147)

Simulium gonzalezi Vargas & Diaz Najera, 1953b:
235. Syntypes 2, CO and pupae, Mexico and
GUATEMALA (more precise information on
localities, collection date, collectors and depos-
itory not given).

Description. Female. General body colour black.
Body length (alcohol preserved specimens) 1.5—
2.6mm (n = 44), wing length 1.2-1.7 mm (n = 22),
wing width 0.6—-0.8 mm (n = 20).

Coloration and morphology as in S. exiguum
except femora and inner surface of tibiae of mid
leg and coxae of fore leg brownish black (Fig.
139). Tergal plates wider than in S. exiguum and
paraproct with short anteriorly directed process
(Fig. 56).

Male. There are two forms of the male: the pre-
dominant dichoptic form (Fig. 10) and the holop-
tic form (Fig. 11) (in the proportion 44 : 5,n = 49).

Dichoptic form. Body length (alcohol pre-
served specimens) 1.3-1.9 mm (n = 18), wing
length 1.1-1.3 mm (n = 8), wing width 0.5—-0.6mm
(n=6).

Coloration and form of head as in female except
head small relative to that of female and holoptic
male (Figs 9-11). Frons at narrowest point about
one-eighth width of head at that point compared
to about one-sixth in the female. Antennae of
dichoptic males longer in proportion to depth of
head (ommatidial area) (1.25 : 1) than in females
(0.9 : 1), and the antennae of these males are
thinner.

Coloration and setation of thorax and appen-
dages as in female except claws of male type (Fig.
147).

Coloration of abdomen as in S. exiguum male
except central area of tergite V silver. Basal fringe
short as in female S. gonzalezi. Genitalia as in
male §. exiguum except ventral plate more rec-
tangular than triangular and basal arms more

SIMULIIDAE OF ECUADOR

developed (Figs 77, 78) and median sclerite nar-
rower at base (Fig. 91).

Holoptic form. Body length (alcohol preserved
specimen) 2 mm (n = 1). Coloration and morphol-
ogy as in male S. exiguum except in the single
specimen dissected ventral plate shows a small
protuberance adjacent to basal arms (Fig. 79).

Pupa. Female and dichoptic male pupae: cocoon
length dorsally 1.5—2.4 mm, ventrally 1.9-2.7 mm;
pupa length 1.4-2.2 mm; gill length 1.1-1.8 mm (n
= 41). Holoptic male pupae: cocoon length dor-
sally 1.6-2.2 mm, ventrally 2.2-2.6 mm; pupa
length 1.6-2.2 mm; gill length 1.0-1.4 mm (n = 5)

Cocoon as in S. exiguum (Fig. 104). Pupal mor-
phology of female and both male forms as in S.
exiguum except for gill. Gill light brown with six
forwardly directed slender filaments arranged
irregularly in a vertical plane (Fig. 111); main
trunk giving rise to three primary branches each of
which bears a single bifurcation. Dorsal branch
with basal bifurcation at about one-tenth length of
total gill, ventral bifurcation the most distal occur-
ring at about a fourth the length of the gill and
median bifurcation between those of the dorsal
and ventral primary branches, although it may
occasionally arise at the same distance from the
gill base as that on the ventral branch.

Mature larva. Body length 2.8-4.6 mm (n = 43).
Width of head capsule 0.3—0.4 mm (n = 43). Body
colour and form as S. exiguum. S. gonzalezi may
be distinguished from S. exiguum by the negative
head pattern (Fig. 124) although the concolorous
state typical of S. exiguum sometimes occurs.

MATERIAL EXAMINED

Belize: 23 2, nr Caya, Augustine, 27.vii.1981
(D. J. Lewis) (BMNH).

Ecuador: numerous reared adults, pupae and
larvae from the following localities in the Santiago
onchocerciasis focus in Esmeraldas Province: R.
Cayapa, San Miguel de Cayapas, 17-19.vi.1981
(A. J. Shelley & M. Arzube) (BMNH, INHMT);
R. San Miguel, San Miguel de Cayapas,
17.vi.1981 (A. J. Shelley & M. Arzube) (BMNH,
INHMT); R. Sapallo Grande, Tumbaviro, 26.v.
and 18.vi.1981 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); R. Grande (Cayapa), Viruela
and Calle Mansa, 24—27.v.1981 (A. J. Shelley &
M. Arzube) (BMNH, INHMT); numerous reared
adults, pupae and larvae from the Canandé
peripheral onchocerciasis focus in Esmeraldas
Province, R. Canandé, Naranjal, 25.ix.1983 and
21-24.vi.1985 (A. J. Shelley & M. Arzube)
(BMNH, INHMT).

Guatemala: 3 Q, Departamento Chim-
altenango, Finca Sibaja, 6.xi.1974 (R. Garms)
(BMNH).

89

Mexico: 1 9, Tamazunchale, SLP, ix.1944
(M. Macias) (BMNH).

TAXONOMIC DISCUSSION. Simulium gonzalezi is
most closely related to S. exiguum and S. para-
guayense Schrottky as detailed in the taxonomic
discussion on S. exiguum.

In the Santiago onchocerciasis focus, as well as
in the Canandé peripheral onchocerciasis focus,
the dichoptic form of the male head is the more
prevalent. A similar situation has already been
reported for this species from a locality in the
Oaxaca onchocerciasis focus in Mexico, where
Dampf (1944) recorded this species (as exiguum).
In the Yepocapa onchocerciasis focus in
Guatemala, however, the holoptic male condition
is more prevalent (Dalmat, 1955; Takaoka, 1983).

DIstTRIBUTION. In Ecuador Simulium gonzalezi has
only been recorded from the main Santiago and
peripheral Canandé onchocerciasis foci. It is also
known from many localities in southern Mexico
(Vargas & Diaz Najera, 1957), Guatemala
(Dalmat, 1955) and Belize (BMNH collection).
Lewis (1963) records an eight-filamented pupa of
S. gonzalezi from Antigua but there is insufficient
evidence presented for this species determination.

BroLocy. In Ecuador S. gonzalezi appears to be
totally zoophilic although its main host has not yet
been determined; using a horse as bait 401 S.
exiguum and one S. gonzalezi were collected in
two hours at the Canandé peripheral onchocer-
ciasis focus (Shelley & Arzube, unpublished
data). Though biting a variety of animals in Cen-
tral America it will often attack man in large num-
bers there. Dalmat (1955) records S. gonzalezi (as
S. exiguum) as being mainly zoophilic with a pref-
erence for large domestic stock (bovines and equi-
nes) but in the lower altitudes (under 800 m) of the
Yepocapa onchocerciasis focus it is the dominant
man-biter (Gibson & Dalmat, 1952, as S.
exiguum). In Mexico S. gonzalezi probably shows
similar biting habits (Shelley, 1988) and speci-
mens (now in BMNH) have been obtained biting
both man and mules in Belize.

Simulium gonzalezi is found in similar breeding
grounds to those of S. exiguum in Ecuador but, as
in Guatemala (Dalmat, 1955), shows a preference
for larger rivers over 10 m wide.

Although not a vector of onchocerciasis in
Ecuador because of its zoophilic habit, S.
gonzalezi is nevertheless an important species
because of its sympatry with and similarity to the
primary vector S$. exiguum. In Mexico and
Guatemala it is suspected as a secondary vector of
onchocerciasis because of its man-biting habit in
the onchocerciasis foci and the finding of filarial
larvae in wild females (Shelley, 1988).

90
Simulium (Psilopelmia) bipunctatum Malloch

(Figs 26, 32, 38, 44, 50, 57, 64, 70, 80, 81, 92, 98,
105, 112, 118, 125, 133, 140, 148)

Simulium bipunctatum Malloch, 1912: 650. Holo-
type 2, Peru: Rio Charape, 13.ix.1911 (C. H.
T. Townsend) (USNM, Cat. No. 15305) [exam-
ined]. [Synonymised with S. dinellii Joan by
Knab, 1913: 155; revalidated by Coscaron,
1985: 320.]

Simulium antillarum Jennings, 1915: 200. Lec-
totype CO’, VIRGIN ISLANDs: St Croix Island, 1.5
miles west of West End, Frederiksted,
24.xi.1913 (A. H. Jennings) (USNM Cat. No.
19997) by designation of Stone (1969: 313)
[examined]. Syn. n.

Simulium wolcotti Fox, 1953: 138. Holotype CO’,
Puerto Rico: Henry Barracks, near Cayey,
1950 (J. Fox) (STMPR). [Synonymised with S.
antillarum Jennings by Stone, 1969: 313.] Syn.
n.

Simulium pseudoantillarum Ramirez Perez &
Vulcano, 1973: 379. Syntypes 1 9, 1 0’, VENE-
ZUELA: Monagas State, San Antonio de Matu-
rin, (no collection date) (Ramirez Perez &
Vulcano) (DDSV). Syn. n.

Description. Female. General body colour
orange. Body length (alcohol preserved speci-
mens) 2.1-3.5 mm (n = 30), wing length 1.7-2.7
mm (n = 30), wing width 0.9-1.2 mm (n = 30).

Head dichoptic with red eyes; nudiocular area
poorly developed (Fig. 26). Frons, clypeus and
occiput black with silver pruinosity. Mouthparts
black. Antennae orange with distal third to half
dark brown. Cibarium with central trough
unarmed and sclerotised and a group of about 20
small teeth forming a protuberance on each side of
trough that do not reach the sclerotised cornuae
(Fig. 32).

Scutum dirty yellow with three prominent long-
itudinal orange bands that coalesce posteriorly;
median band commences on anterior border of
scutum and occupies three-quarters of its length;
the pair of lateral bands commence in the second
quarter of the scutum and continue to posterior
margin. Scutum with a pair of submedian silver
comma-shaped marks commencing at interface
between yellow anterior margin and orange area
and running half length of scutum. Lateral mar-
gins of scutum yellow and faintly pruinose (not
obvious as in S. Jewisi). Paranotal folds orange-
brown in fresh specimens, often becoming dark
brown in preserved material. Scutum with numer-
ous adpressed black setae lying singly. Pleural
region varying from light orange to mid brown
with faint silver’ pruinosity. Scutellum and

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

postnotum orange; posterior margin of scutellum
with erect black bristles (Fig. 140).

Subcostal wing vein with line of setae almost to
distal extremity, basal section of R with two or
three irregular rows of setae. Costal base tuft dark
brown (Fig. 38).

Fore leg coxae, trochanters and femora of all
legs orange to light brown; coxae of mid and hind
legs light brown on anterior half, dark brown on
posterior half; tibiae and tarsi of all legs dark
brown. Mid and hind leg femora and tibiae with
darker distal articulations. Claws curved with
large basal tooth (Fig. 44). Proportions of legs as
in Fig. 140. Halteres yellow with light brown
stems.

Abdominal tergites from orange to brown
depending on age of specimen and whether it has
blood fed; older blood fed specimens tend to
become dark brown. Tergites I-IV usually mot-
tled light brown and yellow but can be yellowish
orange, particularly in reared material, occasion-
ally mid brown; tergite V usually matt grey but
sometimes matt black; tergites VI-IX dull mot-
tled mid and light brown but sometimes com-
pletely shiny brown or black (Fig. 140). Tergal
plates (Fig. 4) well-developed and generally light
brown, sometimes dark brown. Sternites and gen-
italia orange to light brown becoming dark brown
in preserved specimens. Eighth sternite usually
lightly sclerotised with 20-24 setae on each side;
gonopophyses small, membranous with minute
hairs on inner margin (Fig. 50). Cerci hemispheri-
cal; paraprocts with pronounced ventral extension
(Fig. 57). Genital fork slender with well-
developed triangular anterior processes (Fig. 64).
Spermatheca similar to that of S. exiguum (Fig. 5),
oval, sclerotised, with surface covered in regular
rounded depressions and spicules of inner surface
randomly arranged; area of insertion of sper-
mathecal duct membranous and a third as wide as
maximum width of spermatheca.

Male. General body colour orange. Body length
(alcohol preserved specimens) 2.0-3.4 mm (n =
15); wing length 1.8-2.4 mm (n = 15); wing width
0.9-1.0 mm (n = 15).

Head holoptic with red eyes. Clypeus black
with silver pruinosity. Rest of head coloration as
in female.

Coloration and hairing of scutum, pleural
region, scutellum and postnotum as in female
(Fig. 148).

Subcostal wing vein bare or with variable num-
ber of setae (1-7) in central portion, basal section
of R with a single row of setae.

Leg (Fig. 148) and haltere coloration as in
female.

SIMULIIDAE OF ECUADOR

Abdominal tergites I-IV mottled orange and
light brown; tergites V-IX and genitalia light
brown; basal tuft of light orange hairs. Silver orna-
mentation as follows: tergites VI and VII with a
pair of submedian silver pruinose patches, ante-
rior margin of tergite I and all of tergite IX faintly
pruinose (Fig. 148). Sternites I-IV light orange,
V-IX dark brown; sternal plates well-developed
only on segments V—VIII (cf. Fig. 6). Genitalia
orange to light brown. Gonocoxite longer than
wide; gonostyle small, half the length of the
gonocoxite, curved and conical with apical spine
(Fig. 70). Ventral plate (Figs 80, 81) with reduced
and lightly sclerotised basal arms and a small keel;
hairs long and covering most of ventral plate.
Median sclerite (Fig. 92) slightly longer than wide
with deep apical incision occupying about half the
length of the sclerite. Paramere as in Fig. 98 with
few, well-developed, mainly apical spines and sev-
eral smaller spines.

Pupa. Length of cocoon dorsally 2.1-3.2 mm,
ventrally 2.8-4.2 mm; length of pupa 2.0-3.3 mm;
length of gill 2.5—3.2 mm (n = 38).

Cocoon slipper-shaped, dark brown; rim of
aperture dark brown, reinforced and usually with-
out median protuberance (Fig. 105). Cocoon
composed of thick threads producing an open
weave, particularly laterally at point of adhesion
to substrate. Gill light brown with eight forwardly
directed slender filaments arranged irregularly in
a vertical plane (Fig. 112): main trunk giving rise
to three primary branches, ventral with two fila-
ments and median and dorsal each with three
filaments; filaments arise basally on all primary
branches; filaments slender with crenated margins
and rounded distally, their surfaces covered with
fine spicules as in S. exiguum (Fig. 12). Head
similar to that of S. exiguum (Fig. 13) with 2 + 2
frontal and 1 + 1 facial bifid or trifid well-
developed trichomes; surface of head with spar-
sely distributed platelets. Thorax similar to S.
exiguum (Fig. 17) with S + 5 trichomes on anterior
border, each with two to five trichomes, 1 + 1
postero-dorsal and 1 + 1 ventral unbranched tri-
chomes. Surface of thorax covered with platelets
mainly concentrated around the dorsal region.
Abdominal tergites I-IV with 4 + 4 simple
hooks, more weakly developed on segment II,
VI-IX with spine combs on anterior margins,
tergite IX with 1 +1 strong unbranched spines
(Fig. 19); sternite IV in female with 2 + 2 simple
hooks, in male reduced to fine setae, sternites
V-VII with 2 + 2 simple to bifid hooks; 1 + 1
patches of spine combs on postero-lateral borders
of sternites [V—VIII (Fig. 20).

Mature larva. Body length 3.8-5.8 mm (n = 46).

91

Width of head capsule 0.40.6 mm (n = 46). Body
colour grey in both alcohol and Carnoy’s pre-
served specimens. Body form as in Fig. 118.

Head yellow with brown markings and several
scattered minute setae on all surfaces. Head pat-
tern negative as in Fig. 125, consisting of a central
clear area and 1 + 1 antero-lateral and 1 + 1
postero-lateral clear areas of head spots within a
dark background confined to the posterior half of
the cephalic apotome. More rarely the head pat-
tern may be positive (Fig. 126) and this form has
also been recorded outside the focus. Postgenal
cleft large, longer than wide, with pointed ante-
rior margin; postgenal bridge short, one-third as
long as hypostomium (Fig. 133). Hypostomium of
type seen in S. exiguum (Fig. 21). Antennae long,
brown and with segment ratios 12 : 14: 17. Mandi-
ble as in S. exiguum (Fig. 23) with one to three
mandibular serrations, of which anterior is usually
larger. Maxillary palp short, about twice as long as
breadth at base. Cephalic fan with 3442 rays.

Thorax whitish grey dorsally, darker grey ven-
trally, either diffuse or concentrated into one to
three central patches posterior to the proleg. Cuti-
cle with minute scattered hairs on dorsal and lat-
eral surfaces. Proleg plates lightly sclerotised with
10-12 processes. Pupal respiratory histoblast dark
brown and claviform.

Abdomen whitish with four grey bands encir-
cling the body on the four (narrow) anterior seg-
ments, bands more obvious dorsally; posterior
(wide) segments of abdomen brownish grey dor-
sally and whitish laterally and ventrally. Ventral
papillae small (Fig. 118). Cuticle with minute scat-
tered hairs on dorsal and lateral surfaces as in
thorax but in greater density on postero-dorsal
region. Anal sclerite well sclerotised with pos-
terior arms extending to twelfth row of posterior
circlet hooks. Posterior circlet with about 62-71
rows of 1 to 12 hooks. Anal gill tri-lobed as in S.
exiguum (Fig. 24), each lobe with 4~7 short,
secondary lobules.

MATERIAL EXAMINED

Holotype Q of S. bipunctatum, Peru: Rio
Charape, 13.ix.1911 (C. H. T. Townsend)
(USNM); paratype @ (collection data as for holo-
type) (BMNH). Lectotype ©’, paralectotype 9,
various topotypic pupae of antillarum, St Croix
Island: Frederiksted, 24.xi.1913 (A. H. Jennings)
(USNM).

Brazil: 3 2, Roraima, MEVA Mission post,
Auaris, 7.vii.1976 (A. J. Shelley) (BMNH); 10 ?,
Amazonas, R. Vaupes, Igarapé Tiquié, 15.
xii.1977 (C. Vicente) (BMNH, IOC).

Colombia: 3 Q, Norte de Santander,

92

Arboledas, Siravita, La Esperanza, 25.x1.1984 (B.
Alexander) (BMNH).

Dominica: 6 2, Roseau, 7.vii.1974 (L. J.
Charles) (BMNH).

Ecuador: numerous reared adults, pupae and
larvae from following localities in the Santiago
onchocerciasis focus, Esmeraldas Province, R.
Cayapa, 18-21.vi.1981 (A. J. Shelley & M.
Arzube) (BMNH, INMHT); R. Cayapa, stream 4
km below San Miguel de Cayapas, 17.vi.1981 (A.
J. Shelley & M. Arzube) (BMNH, INHMT); R.
Cayapa, above Sapallo Grande Mission,
28.v.1981 (A. J. Shelley & M. Arzube) (BMNH,
INMHT); R. San Miguel, small stream 100 m
above San Miguel de Cayapas, 17.vi.1981 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); R. San
Miguel de Cayapas, Estero Hacha, 26.v.1981 (A.
J. Shelley & M. Arzube) (BNMH, INMHT).

Jamaica: 8 9, Loaf Water, 6.iii.1958 (D. J.
Lewis) (BMNH); 2 9 2 O (ex pupae), Pindars
River, near Kellits, 17.vii.1970 (Clarendon)
(BMNH).

Montserrat: 4 2, Montserrat Woodlands,
20.ix.1938 (F. A. S.) (BMNH).

Venezuela: 8 2, 10 pupae, Monagas State,
localities near San Antonio de Maturin, v.1961
(D. J. Lewis) (BMNH) [labelled by Lewis as sp. D
and regarded by Ramirez Perez as S.
pseudoantillarum].

The following specimens of two species closely
related to S. bipunctatum were examined and are
referred to in the following taxonomic discussion.

Simulium dinellii Joan

Argentina: 27 2, Jujuy (collection date
unkown) (W. C. Paterson) (BMNH).

Peru: 19 9, Apurimac, Cuzco-Abancay road,
Apurimac crossing at Cuya, 7.vil.1971(C. & M.
Vardy) (BMNH).

Simulium ochraceum Walker

Guatemala: 3 2, Department Chimaltenango,
Finca Sta. Anita, 8.11.1974 (R. Garms) (BMNH);
1 2 ,2 C0, Department Chimaltenango, Acan-

tenango, 2.x.1948, 8.vi.1949 (H. Dalmat)
(BMNH).
Mexico: 3 Q, Chalchihuitan, 4.xii.1940 (J.

Parra) (BMNH); 10 9, Chiapas State, Huixtla,
x.1958 (R. W. Crosskey) (BMNH).

Panama: 1 9, Chiriqui Province, Los Planes de
Hornito, 12.ix.1978 (J. Petersen) (BMNH).

TAXONOMIC DISCUSSION. Simulium bipunctatum
was first described by Malloch (1912) from
females collected in Peru (and presumably biting
man) in a survey for the possible transmitters of
pellagra. This name was then synonymised by
Knab (1913) with S. dinellii Joan from Argentina

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

after he had compared the holotype of S.
bipunctatum with the original description of the
former. He neglected to observe that Malloch
named S. bipunctatum after the two submedian,
white comma-shaped marks on the female scutum
whilst Joan (1912) had clearly noted the absence
of scutal patterns in S. dinellii. Wygodzinsky
(1950) questioned Knab’s synonymy of
bipunctatum with dinellii in a paper redescribing
the latter species. Recently, Coscaron (1985)
recalled S$. bipunctatum from synonymy after
examining the holotype in the USNM and
topotypic material. His study confirmed the dis-
tinctness in scutal patterns of the two species and
also revealed differences in the morphology of the
paraproct. We support this revalidation after
comparing a paratype of S. bipunctatum donated
to the BMNH by the USNM, with specimens of S.
dinelliiin the BMNH collection that correspond to
Joan’s original description and the more detailed
description of Wygodzinsky (1950). We base our
synonymy of S. antillarum Jennings with S.
bipunctatum on a comparison of the male lec-
totype and a female paralectotype (USNM) of the
former with the holotype of the latter. Examin-
ation of large numbers of specimens of ‘anti-
llarum’ from the Caribbean Islands and Ecuador
showed variation in the sclerotisation and denti-
tion of the female cibarium, characters which Cos-
caron (1985) uses to separate it from bipunctatum.
The detailed descriptions and comments of Cos-
caron, (1985), Diaz Najera (1961), Floch & Abon-
nenc (1946), Jennings (1915), Rubtsov & Garcia
Avila (1972), Smart (1940) and Vargas & Diaz
Najera (1951) for S. antillarum and the original
description of S. bipunctatum by Malloch (1912)
were consulted.

We accept Stone’s (1969) synonymy of S.
wolcotti with S. antillarum because of his compari-
son of topotypes of the former with the original
syntype series of S. antillarum, which did not con-
firm Diaz Najera’s observations (1961) supporting
the validity of S. wolcotti based on coloration.
Following our synonymy of S. antillarum with S.
bipunctatum, S. wolcotti becomes a new junior
synonym of S. bipunctatum. Coscaron (1985)
omits S. wolcotti from his revision of the subgenus
Ectemnaspis.

The synonymy of S. pseudoantillarum Ramirez
Perez & Vulcano is based on the following obser-
vations. Simulium pseudoantillarum was des-
cribed by Ramirez Perez & Vulcano (1973) from
collections that they had made in the region of the
onchocerciasis foci of northern Venezuela. The
specimens are listed as paratypes but as no holo-
type was designated they have syntype status. The
authors cite the BMNH as a depository but no

|

SIMULIIDAE OF ECUADOR

specimens have yet been received and all type
material is presumed to be in the ‘Seccion de
Oncocercosis, Villa de Cura, Venezuela’. This
species corresponds with ‘Simulium sp. D’ of
Lewis (1963) who had previously collected in the
same area and made a preliminary description.
Ramirez Perez & Vulcano (1973) state that S.
pseudoantillarum is closely related to S. anti-
llarum but can be distinguished from it by dif-
ferences in size, the form of the pedisulcus and
calcipala in the female and in the more distal
bifurcation of pupal gill filaments. Their con-
clusions are stated to have been made after a
comparison of S. antillarum types in the BMNH
and of descriptions of this species by Jennings
(1915), Diaz Najera (1961) and Floch & Abon-
nenc (1946). We have no record of the S. anti-
llarum types having been on loan to the BMNH
from their depository the USNM. We have exam-
ined topotypic material collected by Lewis in
Venezuela in 1961 and identified as Simulium sp.
D (Lewis, 1963) and found that the figure of a
pupal gill with the relatively distal bifurcations of
filaments arising on the median primary gill
branch is atypical for the sample, the majority of
the specimens having a gill form asin S. antillarum
(Fig. 112). Ramirez Perez & Vulcano (1973: 380,
fig. 2) copied Lewis figure of the gill (Lewis, 1963:
57, fig. 3H) but mistakenly state that it is a more
detailed figure of their figure 1, which is obviously
of another specimen. In later works Ramirez
Perez (1983) and Ramirez Perez et al. (1982) fig-
ure the gill of S. pseudoantillarum with basal
branching typical of S. antillarum. Taking into
account the natural variation in adult size, the lack
of detectable difference in the form of calcipala
and pedisulcus, and the natural variation in gill
configuration, we consider S. pseudoantillarum to
be conspecific with S. antillar'um and hence syn-
onymous with S. bipunctatum. Coscaron (1985)
suggests that S. pseudoantillarum may fall within
the natural variation of S. antillarum.

We retain Simulium bipunctatum in the sub-
genus Psilopelmia and do not follow Coscaron’s
inclusion (1985) of this species in Ectemnaspis. It
is morphologically most closely related to S.
ochraceum which differs in its dark postnotum and
legs. Simuliid species with orange coloration pre-
serve badly, both as pinned and alcohol speci-
mens, and female coloration can be influenced by
whether the specimen is blood fed or not. Vari-
ation in coloration of the key characters of the
thorax and abdomen in both species is apparent in
the specimens in the BMNH collection. The dis-
tinctness of the two species needs to be more
thoroughly investigated to take into account intra-
specific variation by using both morphological and

93

cytological methods on long series of specimens
from different localities.

DIstTRIBUTION. Distribution of S. bipunctatum is
based upon the BMNH collections and Stone
(1969, as antillarum). In Ecuador S. bipunctatum
has only been reported in Esmeraldas Province.
Its distribution in South America includes north-
ern Brazil, Colombia, Peru and northern and
southern Venezuela. In Central America it has
been reported from the following Caribbean
Islands: Cuba, Dominica, Guadeloupe, Jamaica,
Montserrat, Puerto Rico, St Croix, and Trinidad
as well as from mainland Mexico.

BioLocy. Simulium bipunctatum will bite man to
varying degrees in different localities. In the
onchocerciasis focus of Ecuador (Shelley &
Arzube, 1985) and in the majority of the Carib-
bean Islands and Mexico (Stone, 1969; Rubtsov &
Garcia Avila, 1972) it is mainly zoophilic, only
rarely coming to bite man. It is, however, mark-
edly anthropophilic in the Upper Amazon region
of Brazil along the R. Vaupes but only occasion-
ally bites man (possibly due to small fly popula-
tions rather than zoophilic tendencies) farther
north in the Amazonia onchocerciasis focus of
Brazil and Venezuela (Ramirez Perez et al., 1982;
Shelley, 1988; Shelley & Luna Dias, unpublished
data). In parts of Guadeloupe it can also be a
biting nuisance (Floch & Abonnenc, 1946).

Simulium bipunctatum breeds in shaded, slow-
flowing streams in lowland forest of Ecuador
(Shelley & Arzube, 1985) and in fast flowing small
streams in St Croix and Jamaica (Jennings, 1915).
In Cuba (Rubtsov & Garcia Avila, 1972) it occurs
in small streams up to an altitude of 1200 m
throughout the year. In the Amazonia onchocer-
ciasis focus of Brazil and Venezuela it occurs in
slow flowing streams (1 m wide, Ramirez Perez et
al., 1982) and larger rivers (up to 20 m wide,
Shelley & Luna Dias, unpublished data) in high-
land forested areas.

Simulium (Psilopelmia) lewisi Ramirez Perez

(Figs 27, 33, 39, 45, 51, 58, 65, 71, 82, 83, 93, 99,
106, 113, 119, 127, 134, 141, 149)

Simulium lewisi Ramirez Perez, 1971: 349. Holo-
type 2, VENEZUELA: Miranda State, Panaquire
(No collection date; collector presumably J.
Ramirez Perez) (DERM).

Simulium iguazuense Coscaron 1976a: 147. Holo-
type [sex unspecified], ARGENTINA: Parque
Nacional Iguazu, route 101, in unnamed stream
or Arroyo Nandu [not specified], 17.x.1974 (S.
Coscaron) (MLP). [Synonymised with S. lewisi
by Coscaron, 1985: 304.]

94

DEscripTION. Female. General body colour
orange and black. Body length (alcohol preserved
specimens) 1.6—-2.3 mm (n = 11); wing length 1.8-
2.1mm (n = 10), wing width 0.8-1.0 mm (n = 10).

Head dichoptic with dark red eyes; nudiocular
area poorly developed (Fig. 27). Frons, clypeus
and occiput black with silver pruinosity.
Mouthparts mid brown, maxillary palps dark
brown. Antennae dark brown with scape and ped-
icel and first flagellomere orange. Cibarium with
five irregular rows of blunt tubercles in area of
central trough and a group of about 20-30 minute
teeth between this and each cornua; anterior mar-
gin of cibarium sclerotised (Fig. 33).

Scutum orange, humeri yellow to light orange,
lateral scutal margins yellowish orange with bril-
liant white pruinosity. Paranotal folds dark brown
with silver pruinosity. Scutum with numerous
adpressed dark brown hairs lying singly. Pleural
region mainly dark brown with faint silver pru-
inosity although in some specimens orange to light
brown areas may occur in the area adjacent to the
paranotal folds. Scutellum orange with erect
brown hairs on posterior margin. Postnotum dark
brown with faint silver pruinosity (Fig. 141).

Subcostal wing vein usually with one to six setae
in the median third of the vein, in some cases
without setae; basal section of R with single row of
setae to base of vein (Fig. 39). Costal base tuft of
dark brown setae.

Legs black except basal two-thirds of mid and
hind basitarsi which are white (Fig. 141). Claws
curved, each with a small tooth as in Fig. 45.
Proportions of legs as in Fig. 141. Halteres yellow
with light brown stems.

Abdominal tergites I-III bright yellow, some-
times orange, tergite IV velvet-black, tergites
VI-IX shiny black or mottled brown and black
(Fig. 141). Occasionally specimens occur in which
tergites I-IV are yellow, in which case tergite V is
velvet-black. Tergal plates (Fig. 4) well de-
veloped on segments IV-IX. Sternites I-III yel-
lowish brown, rest mid brown, genitalia dark
brown. Eighth sternite well sclerotised with about
15-18 setae on each side; gonopophyses small,
membranous with minute hairs (Fig. 51). Cerci
hemispherical; paraprocts with pronounced ven-
tral extension (Fig. 58). Genital fork slender with
sclerotised, triangular anteriorly-directed pro-
cesses (Fig. 65). Spermatheca similar to that of S.
exiguum (Fig. 5), oval, sclerotised, with no exter-
nal sculpturing and randomly distributed spicules
on internal surface. Area of insertion of sper-
mathecal duct membranous, one-third as wide as
maximum width of spermatheca.

Male. General body colour orange and black.

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Body length (alcohol preserved specimens) 1.9-
2.8mm (n = 15), wing length 1.7-2.1 mm (n= 15),
wing width 0.8-1.0 mm (n = 15).

Head holoptic with dark red eyes. Coloration of
rest of head as in female.

Coloration and hairing of thorax and its appen-
dages as in female (Fig. 149) except subcostal vein
of wing devoid of setae. Morphology of legs as in
Fig. 149.

Abdominal tergites I-III yellow, rest of tergites
and genitalia velvet-black; basal fringe of long
black hairs. Silver ornamentation on tergites as
follows: tergite II faintly silver pruinose and
tergites VI, VII and IX with obvious silver patches
laterally (Fig. 149). Sternites I-III orange, [V-IX
dark brown with well-developed sternal plates
(Fig. 6) on segments IV—VIII. Gonocoxite longer
than wide, gonostyle with distal spine longer than
wide and about half length of gonostyle (Fig. 71).
Ventral plate membranous with lightly scle-
rotised, reduced basal arms, a small keel and hairs
covering most of its surface (Figs 82, 83). Median
sclerite pyriform with small apical incision (Fig.
93). Paramere as in Fig. 99 with few distal spines
of varying sizes.

Pupa. Cocoon length dorsally 1.9-2.7 mm, ven-
trally 2.3-3.0 mm; pupa length 1.6-2.5 mm; gill
length 2.2-2.7 mm (n = 32).

Cocoon slipper-shaped, white under natural
conditions and light brown in alcohol; rim of aper-
ture thickened and without median protuberance,
median thickened dorsal ridge connecting with
rim of aperture (Fig. 106). Cocoon very thick,
composed of amorphous elastic substance con-
taining fibres which are only apparent under
higher magnification. Gill light brown with eight
forwardly directed slender filaments arranged in a
vertical plane (Fig. 113); main trunk giving rise to
three primary branches, ventral with two fila-
ments and median and dorsal each with three
filaments; ventral branch bifurcation at limit of
basal third of total gill length, first division of
median and dorsal branches arise in the same
region but individual specimens vary in the exact
position — some showing divisions at the same
distance from the gill base in all three branches
while in others these divisions occur at different
points on each branch, the most dorsal of the two
filaments arising from this division in the median
and dorsal primary branches again bifurcate, usu-
ally at the same level on each branch, at the mid
point of the gill; filaments slender with crenate
edges, rounded distally, their surfaces covered in
spicules as in S. exiguum (Fig. 12). Head as in Figs
13, 14 of S. exiguum with 2 + 2 frontal trichomes
usually with 4-6 branches but sometimes up to

SIMULIIDAE OF ECUADOR

eight and 1 + 1 facial trichomes with 2-4 branches,
trichomes well-developed; surface of head with
few platelets mainly concentrated around the fac-
ial trichomes. Thorax as in S. exiguum (Fig. 17)
with 5 + 5 well-developed trichomes with 6-8
branches on anterior margin of thorax. Anterior
half of surface of thorax with scattered platelets.
Abdominal tergite II with 4 + 4 simple hairs,
III-IV with 4 + 4 simple hooks, VI-IX with spine
combs on anterior margins, tergite IX with 1 + 1
unbranched spines (Fig. 19); sternite IV in both
sexes with 1 + 1 inner simple or bifid hooks and 1
+ 1 outer fine setae (developed as hooks in
bipunctatum), V—VII with 2 + 2 hooks with 1 to 3
branches; 1 + 1 patches of spine combs on
postero-lateral borders of sternites [V—VIII (Fig.
20).

Mature larva. Body length 3.7-4.9 mm (n = 23).
Width of head capsule 0.4—0.5 mm (n = 23). Body
colour white with either greyish purple or green
markings. These colour variations are not sex
linked (W. S. Procunier, pers. comm.). In Car-
noy’s fixative markings are bright purple or green
and more distinct. Body form as in Fig. 119.

Head yellow with scattered minute setae on all
surfaces. Head spots generally indistinct being the
same colour as the rest of the head capsule, but in
some specimens a negative pattern occurs (Fig.
127). Postgenal cleft large, longer than wide, with
rounded anterior margin; postgenal bridge short,
about one-tenth as long as hypostomium (Fig.
134). Membrane within postgenal cleft containing
green or grey chromatocytes that obscure the out-
line of the cleft. Hypostomium of type figured for
S. exiguum (Fig. 21). Antennae long, light brown,
with segment ratios 14 : 13 : 12. Mandible as in S.
exiguum (Fig. 23) except second comb tooth
shorter than the first or third. Maxillary palp
short, about twice as long as width at base.
Cephalic fan with 30-34 rays.

Thorax whitish grey with few scattered dark
greyish purple spots dorsally and two central
greyish purple patches ventrally posterior to the
proleg. Insome specimens green replaces the grey
coloration. Cuticle with scattered small hairs on
ventral and lateral surfaces but densely dis-
tributed on the dorsum. Proleg plate lightly scle-
rotised with about twelve processes. Pupal
respiratory histoblasts dark brown and ovoid.
Abdomen whitish grey with a prominent dark
greyish purple band at the anterior end and at the
posterior end of the narrow abdominal segments,
posterior (wide segments) of abdomen with four
dorsal dark greyish purple bands that often
coalesce and scattered patches of grey pigment
laterally and ventrally. Pigment in some speci-

95

mens is green. Ventral papillae absent or very
reduced and indistinct. Cuticle with densely dis-
tributed minute hairs on dorsum, more scattered
laterally and absent ventrally. Anal sclerite well
sclerotised with posterior arms extending to the
twelfth row of posterior circlet hooks. Posterior
circlet with 56-60 rows of 4-14 hooks. Anal gill
trilobed as in S. exiguum (Fig. 24), median lobe
with 6-7 secondary lobules, lateral lobes with 8-11
secondary lobules.

MATERIAL EXAMINED

Paratypes of S. lewisi, Venezuela: 1 Q (ex
pupa), Miranda State, Panaquire (no collection
date) (J. Ramirez) (BMNH); 1 Q (ex pupa),
Miranda State, Acevedo District, Panaquire, Rio
Yaguapo, (no collection date) (J. Ramirez)
(BMNH); 1 @ (ex pupa), Carabobo State, Vir-
igima (no collection date) (J. Ramirez) (BMNH);
1 CO (ex pupa), Barinas State, Rio Socopo (no
collection date) (J. Ramirez) (BMNH).

Ecuador: numerous reared adults, pupae and
larvae from the following localities in the Santiago
onchocerciasis focus of Esmeraldas Province: San
Miguel de Cayapas, Rio San Miguel, 25, 26.v. &
17.vi.1981 (A. J. Shelley & M. Arzube) (BMNH,
INMHT); Tumbaviro, R. Sapallo Grande, 26.v.
& 18-25.vi.1981 (A. J. Shelley & M. Arzube)
(BMNH, INMHT); San Miguel de Cayapas and
Calle Mansa, Rio Cayapa, 27.v. & 17.vi.1981 (A.
J. Shelley & M. Arzube) (BMNH); numerous
reared adults, pupae and larvae from the Canandé
peripheral onchocerciasis focus in Esmeraldas
Province, Naranjal, Rio Aguas Negras, Naranjal
and Canandé, 23-25.vi.1983 (A. J. Shelley & M.
Arzube) (BMNH, INMHT); 1 @ (ex pupa),
Esmeraldas Province, Tululbi (Ricaurte), Rio
Bogota, 13.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH); 3 @ (ex pupae), Tululbi (Ricaurte),
Riachuelo Nadadeiro, 14.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH, INMHT); 2 C (ex pupae),
Santo Domingo-Esmeraldas road, R. Caoni,
24.ix.1983 (A. J. Shelley & M. Arzube) (BMNH,
INHMT); 3 9 2 O& (ex pupae), Pichincha
Province, Quito-Santo Domingo road, Rio Tanti,
28.ix.1983 (A. J. Shelley & M. Arzube) (BMNH);
pupae, Napo Province, near Lago Agrio,
14.xii.1982 (A. J. Shelley & M. Arzube) (BMNH,
INMHT).

Venezuela: 1 (ex pupa), Miranda State,
Acevedo District, El Clavo, Rio Sapo (no collec-
tion date) (J. Ramirez) (BMNH); 2 2 1 C’, [Ara-
gua State], Altagracia, Quebrada Caranacare,
28.vi.1961 (D. J. Lewis) (BMNH); 1 9 [Aragua
State], Tucuyito, Rio Aguacatal, 14.vi.1961 (D. J.
Lewis) (BMNH).

96

The following closely related species were also
examined and are referred to in the taxonomic
discussion.

S. alirioi Ramirez-Perez & Vulcano

Venezuela: 2 CO’, Monagas State, San Antonio
de Maturin, Rio Negro, 23.v.1961 (D. J. Lewis)
(BMNH); 4 pupae, Paratebueno, v.1961 (D. J.
Lewis) (BMNH).

S. gabaldoni Ramirez-Perez

Paratypes 2 9 (ex pupae), Venezuela: Monagas
State, San Francisco (no collection date) (J.
Ramirez) (BMNH).

Venezuela: 1 C (ex pupa), Monagas State, San
Antonio (no collection date) (J. Ramirez)
(BMNH).

S. lutzianum Pinto

Ecuador: 1 ? (ex pupa), Imbabura Province,
Salinas-Lita road, 54 km from Ibarra, Rio San
Pedro, 11.1x.1983 (A. J. Shelley & M. Arzube)
(BMNH); 10 2 4 Oo (ex pupae), Pichincha
Province, Quito-Santo Domingo road, Rio Tanti
and Riachuelo Lelia, 28 & 29.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH); 1 CO (ex pupa),
Manabi Province, Chone-Santo Domingo road,
Rio Maceto, 10.xii.1984 (M. Arzube) (BMNH); 2
Q 2 CO (ex pupa), Pastaza Province, Tena-Puyo
road, Rio Puyo, Rio Mira Valle and Riachuelo
Huamayacu, 8 & 9.vi.1985 (A. J. Shelley & M.
Arzube) (BMNH); 3 @ (ex pupae), Cotopaxi
Province, Quevedo-La Mana-La Pilalo road, Rio
San Pablo, 8.vi.1984 (A. J. Shelley & M. Arzube)
(BMNH).

Venezuela: 1 2 1 O (ex pupa), Carabobo State,
Montalban (no collection date) (J. Ramirez)
(BMNH); 2 2 1 Oo (ex pupae), Aragua State,
Rancho Grande (no collection date) (J. Ramirez)
(BMNH); 4 Oo (ex pupae), Merida State, Merida
& Timotes (no collection date) (J. Ramirez)
(BMNH); 5 9 6 O (ex pupae), Miranda State,
Guarenas, Rio Izcaragua, Guatire (no collection
dates) (J. Ramirez) (BMNH).

TAXONOMIC DISCUSSION. Attention was first drawn
to S. lewisi by Lewis (Lewis, 1963; Lewis & Ibanez
de Aldecoa, 1962) during a survey of simuliids as
potential vectors of O. volvulus in the human
onchocerciasis foci in northern Venezuela. Lewis
was unable to identify this species due to the
inchoate state of the taxonomy of Neotropical
Simuliidae at the time and designated it as species
A. During his revisionary work on the Venezue-
lan fauna Ramirez Perez collected Simulium spe-
cies A of Lewis and described it as a new species,
S. lewisi (Ramirez Perez, 1971). The depository of
the holotype is presumed to be the ‘Division de
Endemias Rurales, Maracay, Venezuela’ since

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

the other cited depository, the BMNH, contains
only paratypes. Further descriptions and distribu-
tional records of S. Jewisi were subsequently given
by Ramirez Perez (1983) and Coscaron (1985).
Coscaron’s subsequent descriptions and
nomenclatural changes concerning S. /ewisi and
its near relatives S. iguazuense and S. lutzianum
(Coscaron, 1976a, 1985) have caused uncertainty
over the status of these three species. The con-
fusion concerns the coloration of the abdominal
tergites, which is the only character used to dis-
tinguish them. In 1976 Coscaron described the
new species S. iguazuense, noting the possibility
that it might not be distinct from S. /utzianum; the
females of S. iguazuense were described as having
the first three abdominal tergites yellowish brown
(similar to S. /ewisi) whereas in males no abdomi-
nal segments were yellow (as in S. lutzianum).
Coscaron (1985) later synonymised S. iguazuense
with S. /ewisi, believing that the abdominal tergite
coloration of males of these two species varies
intraspecifically. In this later paper he records a
greater variation in abdominal coloration of male
S. lewisi but it is not clear in which localities this
occurs, and there are inconsistencies in the
descriptions of both sexes of this species in the text
and keys. Female S. /ewisi are said to have the first
three abdominal segments yellow, whereas in the
key segments I-II or III (presumably meaning I-II
or I-III) are yellow or yellowish brown. Similarly,
male S. /ewisi are described as having segments I,
I-II, I-III or none yellow, whereas in the key I,
II, or I-III are recorded as yellow. Similar
inconsistencies are seen with S. /utzianum. The
first two abdominal segments of the female are
described as light brown in the text whereas in the
key the first segment is said to be yellow; in males
tergite I is described as brown in the text but light
brown in the key. Considering the confusion over
this character it is unclear why Coscaron chose S.
lewisi as the senior synonym of S. iguazuense
rather than synonymising both S. iguazuense and
S. lewisi with S. lutzianum. While we accept his
synonymy at this stage the reliability of such
colour differences for species separation needs
confirmation by using morphological and
cytological methods before the status of S.
iguazuense, S. lewisi and S. lutzianum is decided.
Variation in abdominal coloration of both
female and male S. /ewisi was seen in our material,
usually the first three and rarely the fourth
abdominal segments in both sexes being yellow to
orange. The male paratype of S. lewisi in the
BMNH collection shows orange coloration on the
first four abdominal segments. The closest rela-
tive to S. lewisi is S. lutzianum, which differs by
the presence of yellow coloration only on the first

SIMULIIDAE OF ECUADOR

tergite of the female abdomen and none on the
male abdomen. Other closely related species
showing coloration differences in the thorax or in
pupal gill filament number are S. alirioi, S.
gabaldoni, S. romanai Wygodzinsky and S.
adolfolutzi Wygodzinsky.

Coscaron (1985) includes S. /ewisi in the sub-
genus Ectemnaspis but we prefer to place it in the
closely related subgenus Psilopelmia. Whether
these taxa should be maintained as valid sub-
genera awaits further integrated morphological
and cytological studies.

DistriBUTION. In Ecuador S. lewisi is a common
species of the onchocerciasis foci and circumja-
cent lowland areas on the eastern and western
foothills of the Andean cordillera.

Simulium lewisi is widespread in northern
Venezuela (Ramirez Perez, 1983). Coscaron
(1985) reports S. /ewisi from Argentina, Bolivia
and Peru but the records might refer in some cases
to S. lutzianum (see taxonomic discussion).

BroLocy. In Ecuador S. /ewisi occurs in small
shaded streams and in rivers up to 100 m wide in
lowland tropical forest either side of the Andean
cordillera (Shelley & Arzube, 1985, unpublished
data), and in rivers in lowland areas in central and
western Venezuela (Ramirez Perez, 1971). The
species is apparently totally zoophilic in Ecuador
(Shelley & Arzube, 1985) and there are no records
of it attacking man elsewhere.

Simulium (Psilopelmia) escomeli Roubaud

(Figs 28, 34, 40, 46, 52, 59, 66, 72, 84, 85, 94, 100,
107, 114, 120, 128, 135, 142, 143, 150, 151)

Simulium escomeli Roubaud, 1909: 428. Syntypes
Q, Peru: Arequipa (no collection date)
(Escomel) (deposited in MNHN but now lost).

Simulium rufidorsum Enderlein, 1934: 283. Holo-
type 2, Peru: (no locality except ‘highland
area’, collection date and collector unknown)
(ZM). [Synonymised with S. escomeli by
Vargas & Dias Najera 1953a: 146.]

Description. Female. General body colour brown
and grey. Body length (alcohol preserved speci-
mens) 2.1-3.0 mm (n = 24), wing length 2.0-2.5
mm (n = 24), wing width 1.0-1.2 mm (n = 24).

Head dichoptic with red eyes; nudiocular area
poorly developed (Fig. 28). Frons, clypeus and
occiput grey pollinose. Mouthparts dark brown.
Antennae dark brown with basal fourth light
brown. Cibarium with unarmed, lightly scle-
rotised central trough and a group of about 20 fine
teeth in two protuberances lateral to this and
extending in a single line along base of each scle-
rotised cornua (Fig. 34).

97

Scutum chestnut-brown with two submedian
vittae, posterior margin and posterior two-thirds
of lateral margins of scutum grey pruinose; ante-
rior third of lateral margins of scutum, humeri and
anterior scutal border between these and the sub-
median vittae light brown; the 1 + 1 cunae in the
anterior margins of the submedian vittae appear
brown with an anterior light source and white
pruinose with posterior lighting. Paranotal folds
grey pruinose. Scutum with numerous adpressed
golden hairs (Fig. 142). Pleural region with light
and dark brown areas with faint silver pruinosity.
Specimens from high altitudes are darker; the
chestnut-brown of the scutum is darker brown,
often with areas of black at the interfaces with the
grey pruinose areas, and the whole of the pleural
region is dark brown with grey pruinosity (Fig.
143). (Similar findings on high altitude Peruvian
specimens are reported in Knab (1914a, b).)
Scutellum light brown with adpressed golden hairs
and an uneven row of upright dark brown hairs on
posterior margin. Postnotum dark brown with sil-
very grey pruinosity.

Subcostal wing vein usually bare but sometimes
with up to two hairs on median section, basal
section of radius bare (Fig. 40). In specimens from
highland areas Sc with 4-6 hairs on median sec-
tion, basal section of R usually bare but occasion-
ally with a single hair on median section of vein.
Costal base tuft of dark setae.

Legs yellowish with the following areas dark
brown: distal articulation of tibia and whole of
tarsus of fore leg, external face of tibia faintly
pruinose; coxa, distal articulations of femur and
tibia, distal half of basitarsus, second tarsomere
and rest of tarsus of mid leg; coxa, distal fourth of
femur, distal half of tibia, distal half of basitarsus
and second tarsomere and rest of tarsus of hind
leg. Proportions of legs as in Fig. 142. Claws
slender, slightly curved and lacking a basal tooth
(Fig. 46). Halteres white with yellowish stem.

Abdominal tergites yellowish grey. Tergal
plates (Fig. 4) not obviously developed. Tergite II
with median dark brown amorphous spot, tergites
III-VI with well-defined central subrectangular
black spots; tergites III-VII with pairs of subme-
dian black spots and tergites II-VII with pairs of
indistinct sublateral black spots (Fig. 142). Ster-
nites greyish, genitalia brown. Eighth sternite
highly sclerotised with a group of 20-25 hairs on
each side; gonopophyses small, glabrous, with
light sclerotisation on inner margin. Cerci hemi-
spherical, paraprocts with pronounced anterior
extension (Fig. 59). Genital fork with well-
developed sclerotised anteriorly directed pro-
cesses and sclerotised stem (Fig. 66). Sper-
matheca oval as in S. exiguum (Fig. 5), highly

98

sclerotised with no external sculpturing and
spicules of inner surface randomly distributed;
area of insertion of spermathecal duct mem-
branous, about a third as wide as spermatheca at
widest point.

Male. General body colour brown and black,
showing variations in different localities. Popula-
tions from lowland localities have orange thoraces
and in those from highland areas these are gener-
ally black but in some instances orange. Body
length (alcohol preserved specimens) 2.3-2.8 mm
(n=5), winglength2.0-2.6mm(n=S), wing width
1.0-1.2 mm (n =5).

Head holoptic with red eyes. Coloration of rest
of head as in female except specimens from high-
land areas which have black antennae with the
scape and pedicel orange-brown.

Scutum-orange brown to chestnut in lowland
localities. Anterior border of scutum, except for
central area and humeri and lateral margins yel-
lowish, posterior margin black as is interface
between yellow lateral margin with orange-brown
of scutum; a pair of submedian, comma-shaped
marks occupy the anterior half of the scutum (Fig.
150). Direction of light source affects the colora-
tion of some of the scutal patterns; with light
source posterior to specimen the pair of subme-
dian, comma-shaped marks appear black, curved
and almost reach the anterior border but with light
source anterior to specimen they appear silver
pruinose, are more triangular and adjoin the ante-
rior scutal margin; with light source perpendicular
the posterior and lateral margins of the scutum
appear silver pruinose. In some specimens the
black posterior margin of the scutum extends
anteriorly to join the commas. Paranotal folds
black. Scutum with adpressed golden hairs. In
specimens from most highland localities the
orange-brown areas of the scutum are velvet-
black (Fig. 151). Pleural region with superior part
light brown with faint silver pruinosity and
inferior part grey and_ strongly pruinose.
Scutellum light brown with adpressed yellowish
hairs and an uneven row of upright dark brown
hairs on posterior margin. Postnotum black with
silver pruinosity.

Subcostal wing vein and basal section of radius
bare. Costal base tuft of dark setae.

Coxa, trochanter, femur and tibia of fore leg
light brown, tarsus black; external face of tibia
white pruinose. Coxa of mid leg dark brown,
trochanter, femur, basal half of basitarsus and of
second tarsomere light brown, rest dark brown to
black. Coxa of hind leg black, trochanter, basal
two-thirds of femur, proximal fourth of tibia and
basal half of basitarsus light brown, rest black.

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Proportions of legs as in Fig. 150. Haltere colora-
tion as in female.

Abdomen and genitalia velvet-black, except
tergite II and lateral margins of tergite III which
are brown. Basal fringe of first tergite light brown.
Silver ornamentation as follows: tergite II silvery
white pruinose except for central black spot,
tergite V with a pair of submedian silvery white
spots, tergite VI silver pruinose and lateral mar-
gins of tergite VII silvery pruinose (Fig. 150). In
males from highland localities the silvery white
areas on tergite V are usually absent. Sternites
dull brown with poorly developed sternal plates
(Fig. 6) on segments IV—VIII. Gonocoxite one
and a half times as long as wide; gonostyle subrec-
tangular, half as long as gonocoxite and with a
distal spine (Fig. 72). Ventral plate lightly scle-
rotised, subrectangular, with well-developed
basal arms and a small keel; hairs long and densely
distributed over most of ventral plate (Figs 84,
85). Median sclerite elongate, subquadrangular,
with a deep incision in apical third (Fig. 94). Para-
mere with numerous large spines (Fig. 100).

Pupa. Cocoon length dorsally 2.0-3.0 mm, ven-
trally 2.8-3.4 mm; pupa length 2.0-3.0 mm; gill
length 1.6-2.4 mm (n = 40) (all lowland popu-
lations).

Cocoon slipper-shaped, dark brown; rim of
aperture dark brown, reinforced and without
median protuberance (Fig. 107). Cocoon surface
of thin, amorphous, elastic substance containing
loosely interwoven fibres visible at higher magni-
fications. Gill light brown with eight, forwardly-
directed, slender filaments arranged irregularly in
a vertical plane; main trunk giving rise to three
primary branches, ventral with two filaments and
median and dorsal each with three filaments; ven-
tral branch with bifurcation in basal fourth of gill,
median branch with first bifurcation in basal
fourth and second bifurcation in basal third of gill;
dorsal branch with first bifurcation basally at junc-
tion of median and dorsal primary branches and
second bifurcation within basal fourth of gill (Fig.
114); filaments slender with crenate margins and
rounded distally, their surfaces covered in fine
spicules as in S. exiguum (Fig. 12). Head as in S.
exiguum (Fig. 13) with 2 + 2 frontal trichomes, the
most dorsal being simple and the more ventral 14
branched both poorly developed and 1 + 1 simple
or bifid facial trichomes that are small and difficult
to distinguish; surface of head covered with plate-
lets asin S. exiguum. Thorax asin S. exiguum (Fig.
17) with 5 + 5 antero-dorsal poorly developed
trichomes with 2-5 branches. Surface of thorax
covered with platelets. Abdominal tergite II with
4+ 4simple hairs, III-IV with 4 + 4 simple hooks,

SIMULIIDAE OF ECUADOR

VI-IX with spine combs on anterior margins, IX
with 1 + 1 strong unbranched spines (Fig. 19);
sternite IV with 1 + 1 simple or bifid, well-
developed outer hooks and 1 + 1 fine inner hairs,
V with 2 + 2 simple hooks, VI with 2 + 2 simple
hooks, inner pair sometimes bifid and VII with 2
+2 simple hooks, 1 + 1 patches of spine combs on
postero-lateral borders of sternites IV—VIII (Fig.
20).

Mature larva. Body length 4.5-5.3 mm (n = 20).
Width of head capsule 0.4-0.5 mm (n = 20). Body
colour white with indistinct grey markings (Fig.
120). In specimens preserved in Carnoy’s the grey
markings appear greyish green. Body form as in
Fig. 120.

Head yellow with dark brown markings and
occasional setae on all surfaces. Typical head pat-
tern negative as in Fig. 128, with dark areas in
form of ring surrounding a median clear area and 1
+ 1 postero-lateral clear areas of head spots.
Postgenal cleft rounded anteriorly and small rela-
tive to the other Psilopelmia species (/ewisi and
bipunctatum) in the locality; postgenal bridge
almost as long as hypostomium (Fig. 135). Hypo-
stomium of type seen in S. exiguum (Fig. 21).
Antennae long, brown and with segment ratios 12
: 14:20. Mandible as in S. exiguum (Fig. 23), with
two mandibular serrations, the posterior being the
larger. Maxillary palp about twice as long as
breadth at base. Cephalic fan with 30-32 rays.

Thorax white dorsally, sometimes with a grey
band around anterior margin; ventrally with two
median patches posterior to proleg. Cuticle
glabrous. Proleg plate lightly sclerotised with
about ten processes. Pupal respiratory histoblast
mid brown and ovoid.

Abdomen white with 1 + 1 dorso-lateral grey
patches on first six abdominal segments (5 nar-
row, 1 wide), posteriorly patches coalesce and
cover whole of dorsum of expanded region. Ven-
trally patches form indistinct bands on anterior six
abdominal segments. Ventral nerve cord grey.
Occasionally larvae are white and lack grey orna-
mentation. Larvae from highland localities where
dark male forms occur are more variable in color-
ation; they are darker than those from lowland
localities where orange males occur and in some
cases the grey patches on the thorax and abdomen
coalesce to form a totally grey larva. Cuticle with
scattered fine hairs on dorsal surface of posterior
segments. Ventral papillae absent. Anal sclerite
well sclerotised with posterior arms extending to
twelfth row of posterior circlet hooks. Posterior
circlet with about 60 rows of 4-14 hooks. Anal gill
trilobed as in S. exiguum (Fig. 24), usually without
secondary lobules; if present, secondary lobules
short and up to two on each lateral lobe.

99

MATERIAL EXAMINED

Chile: 1 9, Valle de Azapa, vi.1912 (C. E.
Porter) (BMNH); 3 9, Valle de Lluta, vi.1912 (C.
E. Porter) (BMNH).

Ecuador: 3 2, Santiago onchocerciasis focus,
Rio Onzole, v.1982 (J. Mera & Platon) (BMNH);
2 9 4c (ex pupae), numerous pupae, 4 larvae,
Esmeraldas Province, Santo Domingo-
Esmeraldas road, Rio Chigwe, 26.1x.1983 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); 1 Q 1
CO’ (ex pupae), 8 pupae, 5 larvae, Santo Domingo-
Esmeraldas road, Rio Capli, 26.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); 1 9 1
CO (ex pupae), 8 pupae, 10 larvae, Santo Domi-
ngo-Esmeraldas Road, R. Tatica, 26.ix.1983 (A.
J. Shelley & M. Arzube) (BMNH, INHMT);
numerous pupae, Santo Domingo-Esmeraldas
Road, R. Achioti, 26.ix.1983 (A. J. Shelley & M.
Arzube) (BMNH, INHMT); numerous pupae, 2
larvae, Santo Domingo-Esmeraldas road, R. Tab-
uchi, 26.1x.1983 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); 1 Q, near Concordia, road to
Puerto Quito, Rio Blanco, 28.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH); 3 CO (ex pupae),
numerous pupae and larvae, Esmeraldas-
Atacames road, R. Tasechi, 27.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); 13 2 7
CO (ex pupae), Imbabura Province, Salinas road
20 km north of Ibarra, R. Tahuando, 8.ix.1983
(A. J. Shelley & M. Arzube) (BMNH, INHMT);
numerous females, 15 km south of Juncal on
Ibarra road, R. Chota, 7.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH, INHMT); numerous
females, 17 km south of Juncal on Ibarra road, at
Engeno Tababuelo, R. Chota, 7.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); 7 9 5
CO (ex pupae), numerous pupae and larvae,
Otovalo-Ibarra road, stream opposite Cotacachi
turn off, 10.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH); 1 Q, Salinas-Lita road, San Juan del
Hacha, R. Mira, 11.ix.1983 (A. J. Shelley & M.
Arzube) (BMNH); 9 Q, Palacara river, 9.ix.1983
(A. J. Shelley & M. Arzube) (BMNH, INHMT); 1
Q 2 oO’ (ex pupae), numerous larvae and pupae, nr
Ibarra, tributary stream of R. Salado, 11.ix.1983
(A. J. Shelley & M. Arzube) (BMNH);194cC
(ex pupae), numerous larvae and pupae, irriga-
tion canal on Salinas road 4 km from turnoff on
Ibarra-Tulcan road, 9.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH); 1 CO’ (ex pupa), Salinas-Lita
road 54 km from Ibarra, R. San Pedro, 11.1x.1983
(A. J. Shelley & M. Arzube) (BMNH); numerous
pupae, 4 km from Ibarra-Tulcan road, Salinas
road, 9.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); numerous pupae, Salinas-
Lita road, 11.1x.1983 (A. J. Shelley & M. Arzube)
(BMNH, INHMT).

100

Peru: 15 9, Arequipa, [1922] (Escomel)
(MNHN); 2 9, Arequipa (no collection date)
(Escomel) (BMNH) [presented by E. Roubaud
(1923.185)]; 1 2 (no collection date or collector’s
name, but presumed to be Escomel) (BMNH)
[presented by Dr Escomel (1919: 259)].

TAXONOMIC DISCUSSION. Simulium escomeli was
first described by Roubaud (1909) from three
female syntypes collected by Escomel in Are-
quipa, Peru. Roubaud comments on the poor con-
dition of these specimens, which he intended to
describe in detail at a later stage once Escomel had
obtained more material. The fifteen pinned
females in the MNHN are regarded as topotypes
as they bear labels indicating the collection
locality as Arequipa, the collector E. Escomel and
the date 1922; no syntypes were located in this
collection and are presumed lost. The two females
collected by Escomel from Arequipa and deposi-
ted in the BMNH by Roubaud in 1923
(No.1923.185) as well as a third female from the
same locality presented in 1919 (No. 1919: 259)
are also topotypes.

Later, Enderlein (1934a) gave a superficial
description of the nominally new species S.
rufidorsum from an unnamed highland locality in
Peru. Enderlein comments that the holotype was
‘through Staudinger’. Staudinger was a lepidop-
terist and entomological dealer and not the collec-
tor (K. S. Sattler, pers. comm.). Simulium
rufidorsum was synonymised with S. escomeli by
Vargas & Diaz Najera (1953b) after examination
of the holotype female and dissection of its geni-
talia, which conform to those of S. escomeli.

Since Roubaud’s preliminary description S.
escomeli has been redescribed by many authors.
The most detailed is that by Wygodzinsky (1971)
who lists references to all previous descriptions.
(In his work reference to positive head spots on
the larval cephalic apotome should read nega-
tive.) Wygodzinsky’s (1971) work is based on
specimens collected in Chile, Peru and Ecuador at
high altitude localities. His description completely
coincides with the one presented here except that
males always possessed black scuta. The colour
dimorphism described in this paper, particularly
in males, is apparently loosely linked to altitude
and appears to be intraspecific; preliminary analy-
sis of polytene chromosomes of larvae from
localities pure for males of either colour indicate
no obvious differences in their banding patterns
(W. S. Procunier, pers. comm.). Further detailed
chromosome analysis is underway and will be
reported shortly. Although no males, pupae or
larvae were found in the lowland forests of the
onchocerciasis focus we presume that males are

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

identical to those from adjacent lowland areas and
have taken account of this in the identification
Keys.

Simulium escomeli is the type species of the
subgenus Psilopelmia Enderlein.

DISTRIBUTION. Simulium escomeli is a common
species throughout Ecuador, as indicated by the
list of material examined. It is also common in
association with the Andean cordillera of western
South America in Peru, Chile and Colombia
(Knab 1914a; Wygodzinsky, 1971, collections of
BMNH, INHMT).

BrioLocy. In Ecuador Simulium escomeli bites
man voraciously in higher altitude localities but
rarely in the western lowlands — despite its
omnipresence in the slow-flowing lower reaches
of the smaller rivers. In the onchocerciasis focus
its breeding grounds have not been located. In
higher altitudes it occurs attached to trailing vege-
tation in both slow — and fast — running small
streams.

Simulium escomeli has also been recorded bit-
ing man in Chile and Peru where it also breeds in
small streams and rivers (Coscaron, 1976b;
Wygodzinsky, 1971).

Simulium (Hemicnetha) mexicanum Bellardi

(Figs 15, 16, 18, 22, 29, 35, 41, 47, 53, 60, 67, 73,

86, 87, 95, 101, 108, 115, 121, 129, 136, 144, 152)

Simulium mexicanum Bellardi, 1862 (appendix to
part 2): 6. LECTOTYPE Q, Mexico: Veracruz
State, Tuxpango, near Orizaba (collection date
and collector unknown) (DBAT), here desig-
nated [examined].

Simulium aureopunctatum Malloch, 1914: 27.
Holotype 2, GUATEMALA: Livingston (6.v. or
5.vi. year not given) (Barber & Schwarz)
(USNM Cat. No. 15406) [examined]. [Syn-
onymy by Strong et al., 1934: 208.]

Simulium placidum Knab, 1915: 281. Holotype 9°,
TRINIDAD: Arima river, 31.xi1.1913 (F. W.
Urich) (BMNH) [examined]. [Synonymy by
Vargas & Diaz Najera, 1951b: 133.]

Simulium lugubre Lutz & Nunez Tovar in Lutz,
1928: 46. Syntypes 2, CO’, 1, p, VENEZUELA:
Aragua, Rio de Maracay, La Trinidad,
28.viii.1925 (A. Lutz & Nunez Tovar) (IOC)
[examined]. [Synonymy by Fairchild, 1940:
708. |

Simulium turgidum Hoffmann, 1930: 298. Syn-
types 2, Mexico: Chiapas State, Soconusco
District, Finca Santa Anita (vii.1930) (collector
and depository unknown). [Synonymy by
Strong et al., 1934: 208.]

SIMULIIDAE OF ECUADOR

Description. Female. General body colour black.
Body length (alcohol preserved specimens) 3.2-
4.0mm (n= 24). Wing length2.2-3.6mm (n= 24),
wing width 1.0-1.6 mm (n = 24).

Head dichoptic with red eyes; nudiocular area
well-developed (Fig. 29). Frons, clypeus and
occiput black with grey pruinosity, covered in
numerous black bristles that are longer and denser
on upper margin of clypeus and occiput. Proboscis
light brown, maxillary palps black. Antennae
dark brown with scape, pedicel and _ first
flagellomere orange. Cibarium with large central
trough, unarmed; cornuae well-developed and
sclerotised (Fig. 35).

Scutum, humeri and paranotal folds black with
grey pruinosity. Scutum with numerous, ad-
pressed, short, black setae becoming longer and
upright on posterior border, interspersed with
clumps of adpressed, brass-coloured scale-like
setae (Fig. 144). Pleural region dark brown with
grey pruinosity. Scutellum dark brown to black
with grey pruinosity and with scattered upright
black bristles on whole surface except anterior
border and brass-coloured, adpressed, scale-like
setae over whole surface. Postnotum black with
grey pruinosity.

Subcostal wing vein with an irregular row of
setae over entire length; basal section of R with
three irregular rows of setae along entire length
(Fig. 41). Costal base tuft of dark brown setae.

Fore legs with coxae, trochanters, and femora
light brown, tibiae white pruinose with apical
third and inner margin black, tarsi black. Mid leg
coxae dark brown with grey pruinosity, trochan-
ters and femora black, tibiae black with up to
basal third light brown, basal two-thirds of
basitarsus light brown, rest black, other tarsal seg-
ments black. Hind leg coxae dark brown with grey
pruinosity, trochanters light brown, femora black,
basal third of tibiae white merging to black on
apical two-thirds, basal half of basitarsus white,
distal half black, rest of tarsi black. Colour and
proportions of legs as in Fig. 144. The flattened
and widened fore tarsi, mid and hind basitarsi and
hind femora and tibiae are typical of S. mex-
icanum and several other species of Hemicnetha.
Claws curved with subbasal tooth (Fig. 47).
Halteres white with black stems.

Basal scale of abdomen (tergite I) velvet-black
with long brass-coloured basal fringe, tergite II
mottled brown and black with grey pruinosity,
tergites III-V velvet-black, tergites VI-IX shiny
black (Fig. 144). Tergal plates (Fig. 4) highly scle-
rotised especially on tergite II. Sternites I and I
light brown, rest black. Genitalia black. Eighth
Sternite well sclerotised with 4-6 setae on each
side, gonopophyses well-developed, subtriangu-

101

lar, totally membranous and covered with fine
setae (Fig. 53). Cerci hemispherical; paraprocts
large and subquadrangular with long bristles and
short thick setae (Fig. 60). Genital fork short,
strongly sclerotised and with highly developed lat-
eral arms (Fig. 67). Spermatheca oval as in S.
exiguum (Fig. 5), strongly sclerotised with no
external sculpturing and spicules on inner surface
randomly distributed; width of membranous area
of insertion of spermathecal duct large, about half
maximum width of spermatheca.

Male. General body colour black. Body length
(alcohol preserved specimens) 3.14.5 mm (n =
20), wing length 2.6-3.5 mm (n = 20), wing width
1.1-1.6mm (n = 20).

Head holoptic with red eyes. Clypeus black
with grey pruinosity. Rest of head coloration as in
female.

Coloration and setation of scutum, humeri, par-
anotal folds, pleural region, scutellum and
postnotum as in female, except scale-like setae
golden and thin dark median line running whole
length of scutum, free of these scales (Fig. 152).

Subcostal wing vein and basal section of R bare.

Leg coloration and form (Fig. 152) as in female
except white area of hind tibia reduced to point of
articulation with femur. Halteres as in female.

Basal scale of abdomen (tergite I) velvet-black
with dense basal tuft of long black hairs. Tergites
II-IX velvet-black with the following silver pru-
inose ornamentation: tergite II covering whole
segment, tergite IV covering anterior border
except for median portion, tergites V-VII com-
pletely covered except for median triangle on pos-
terior border of each tergite, tergite VIII with a
small lateral area on anterior margin of tergite
(Fig. 152). Sternites mottled brown and black with
poorly developed sternal plates (Fig. 6). Genitalia
velvet-black. Gonocoxite rectangular, wider than
long, gonostyle elongate with a weakly developed
subterminal spine (Fig. 73). Ventral plate rec-
tangular with sclerotised, poorly developed basal
arms and a large keel; ventral plate densely cov-
ered with fine setae and small spines (Figs 86, 87).
Median sclerite elongate with apical depression
(Fig. 95). Paramere (Fig. 101) with enlarged basal
process and few stout spines apically.

Pupa. Cocoon length 2.5-3.6 mm, ventrally 3.3—
4.4 mm; pupa length 3.24.3 mm; gill length 1.6—
2.5 mm (n = 35).

Cocoon shoe-shaped, mid brown; rim of aper-
ture mid brown, reinforced and without festoons
as seen in other Hemicnetha species (Fig. 108).
Cocoon surface of thin, amorphous, translucent,
elastic substance in which thick, interwoven fibres
are sometimes visible. Gill light to dark brown,

102

generally protruding beyond collar of cocoon,
with 12 short, forwardly-directed filaments
arranged in a bunch. Main trunk of gill giving rise
to an inner primary branch bearing five filaments
and an outer branch with seven filaments. The
filaments arise basally on the gill (Fig. 115), are
slender with crenate margins and rounded dis-
tally, their surfaces covered with fine spicules as in
S. exiguum (Fig. 12). Head with 2 + 2 frontal and
1 + 1 facial trichomes, all poorly developed and
unbranched; surface of head with platelets which
in the frontal region are scattered, enlarged and
highly sclerotised and in the facial region are
dense and of normal size (Figs 15, 16). Thorax
with 5 + 5 antero-dorsal, poorly developed
unbranched trichomes. Surface of anterior region
of thorax covered in well-developed highly scle-
rotised platelets which extend to posterior border
of thorax either side of raphe and in a pair of
submedian and lateral bands to posterior thoracic
border which is densely covered by normal size
platelets. Ventral surface of thorax without plate-
lets (Fig. 18). Abdominal tergite II with 3 + 3
well-developed simple hooks and 1 + 1 simple fine
hairs external to these, IIJ-IV with 4 + 4 simple
hooks, IX with no spines, II-IX with 1 + 1 well-
developed areas of spine combs on anterior mar-
gins and I and II with groups of spine combs on
posterior margins (Fig. 19); sternite TV with no
hooks or hairs, V—VII with 2 + 2 simple hooks, 1
+1 patches of spine combs on anterior borders of
sternites [V—VIII (Fig. 20).

Mature larva. Body length 7.0-9.7 mm (n = 14).
Width of head capsule 0.7-0.8 mm (n = 14). Body
colour grey in specimens preserved in both alco-
hol and Carnoy’s fixative. Form as in Fig. 121.
Head yellow with dark brown markings and
occasional setae on all surfaces. Head pattern
positive as shown in Fig. 129, surrounded by dark
pigmentation. Occasionally the 1 + 1 lateral
groups of head spots are negative in relation to the
dark pigmentation of the cephalic apotome in this
region. Postgenal cleft bell-shaped and rounded
anteriorly (Fig. 136); postgenal bridge almost as
long as hypostomium (Fig. 136). Hypostomium
(Fig. 22) rounded anteriorly with strongly pig-
mented anterior margin and nine _ poorly
developed apical teeth; median tooth the most
developed with blunt apex, corner teeth smaller
and blunt and intermediate teeth poorly defined;
4-5 lateral serrations, hindmost level with first
hypostomial seta; 1 + 1 lines of 9-10 hypostomial
setae. Antennae long and faintly pigmented with
segment ratios 8 : 15 : 10. Mandible with subequal
comb teeth and one well-developed backwardly-
slanting mandibular serration; a second short

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

mandibular serration sometimes present posterior
to the first. Maxillary palp heavily pigmented,
brown, about twice as long as width at base.
Cephalic fan with 48-60 rays.

Thorax grey dorsally, white ventrally with a
large greyish brown patch occupying ventral sur-
face of proleg and a median patch posterior to
this. Cuticle with occasional fine hairs on dorsal
and ventral surfaces. Proleg plate well sclerotised
with about 20 processes. Pupal respiratory histo-
blast dark brown, claviform.

Abdomen greyish brown dorsally, usually over-
all, but in some specimens in indistinct bands in
anterior constricted abdominal segments; ven-
trally whitish with irregular grey patches. Ventral
nerve cord greyish. Ventral papillae absent (Fig.
121). Cuticle with occasional fine hairs on dorsal
and ventral surfaces. Anal sclerite well sclerotised
with posterior arms extending to about 65th row
of posterior circlet hooks. Posterior circlet with
214— 228 rows of 31-40 hooks. Anal gill trilobed
with 12-15 lobules of which one on each lobe is
larger than the rest.

MATERIAL EXAMINED

Lectotype 9, paralectotype Q of S. mex-
icanum, Mexico: Veracruz State, Tuxpango, near
Orizaba (collection date and collector unknown)
(DBAT). Holotype @ of S. aureopunctatum,
Guatemala: Livingston, 6.v. or 5.vi. [no year
given] (Barber & Schwarz) (USNM). Holotype
Q, paratype 2 of S. placidum, Trinidad: Arima
River, 31.xii.1913 (E. W. Urich) (BMNH). Syn-
types pupae and larva of S. /ugubre [only remain-
ing material of syntype series], Venezuela: Rio
Limon, Maracay (7 pupae), 28.viii.1925 (A. Lutz
& Nunez Tovar) (IOC, Cat. Nos. 12134, 12173,
12193-12197); Rio Suaire, Caracas (2 pupae),
1925 (A. Lutz & Nunez Tovar) (IOC, Cat. No.
12198); no data on label but part of Lutz collection
from Venezuela (1 larva) (IOC, Cat. No. 12207).

Belize: 2 2 2 0’, Hell’s Teeth, 15.ii.1958 (P. C.
C. Garnham & D. J. Lewis) (BMNH).

Colombia: 3 @, Norte de Santander,
Arboledas, Siravita, La Esperanza, 26.vii.1986
(B. Alexander) (BMNH).

Ecuador: 8 2 8 © (ex pupae), 10 larvae,
Esmeraldas Province, various localities in the
Santiago onchocerciasis focus, San Miguel de
Cayapas, Casacadita, R. Cayapa, 17.vi. &
26.v.1981 (A. J. Shelley & M. Arzube) (BMNH,
INHMT); 1 @ (ex pupa), Agua Blanca, R.
Cayapa, 15.vii.1986 (P. Beech-Garwood)
(BMNH); 8 9 10 Co’ (ex pupae), numerous pupae
and larvae, Esmeraldas Province, Santo Domi-
ngo-Esmeraldas road, Rio Achioti, 26.ix.1983 (A.
J. Shelley & M. Arzube) (BMNH, INHMT);

SIMULIIDAE OF ECUADOR

numerous pupae and larvae, Santo Domingo-
Esmeraldas road, R. Tabuchi, 26.ix.1983 (A. J.
Shelley & M. Arzube) (BMNH, INHMT); 4 9 2
O (ex pupae), numerous pupae and larvae,
Imbabura Province, stream at 40 km from Ibarra
on Salinas-Lita road, 11.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH); 3 2 9 O&O (ex pupae),
numerous pupae and larvae, Salinas-Lita road, R.
San Pedro, 11.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH, INHMT); 1 9 5 CO (ex pupae), numer-
ous pupae and larvae, Pichincha Province, Quito-
Santo Domingo road, Riachuelo Lelia, 29.ix.1983
(A. J. Shelley & M. Arzube) (BMNH, INHMT);
numerous pupae and larvae, Quito-Santo Domi-
ngo road, R. Toachi, 28.ix.1983 (A. J. Shelley &
M. Arzube) (BMNH, INHMT).

Guatemala: 1 2, Yepocapa onchocerciasis
focus, Chimaltenango, Rio Chusita, Finca
Chusita, 26.viii.1948 (H. Dalmat) (BMNH);2 0’,
Acatenango, Chimaltenango, Rio El Arco, Finca
Chantunjay, 23.x. and 20.xi.1948 (H. Dalmat)
(BMNH).

Honduras: 2 larvae, Departamento de Cortes, 5
m stream, El Cacao, treatment point, 23.i11.1984
(L. Lacey) (BMNH).

Mexico: 2 2, Chiapas State, Escuintla, Jalapa
aldeia, xi.1935 (collector unknown) (BMNH).

Panama: 12 larvae and 6 pupae, Chiriqui
Provence, Boquete, 5.i1.1983 (R. A. Cheke)
(BMNH); 1 pupa, Chiriqui Province, Rio Chiri-
qui Viejo, 25.ix.1975 (M. Kerner) (BMNH); 10
(ex pupa), 12 larvae and 5 pupae, Cocle Province,
El Valle, Rio Anton, La Mapolo, 1-6.vii.1985 (A.
J. Shelley) (BMNH).

Trinidad: 3 @, Naranja, Tucucha Trail,

30.ix.1937 (J. Smart) (BMNH); 4. Q,
Blanchisseuse road, 31.x.1937 (J. Smart)
(BMNH).

Tobago: 5 °,, Pigeon Peak, 16.x.1937 (J. Smart)
(BMNH).

Venezuela: 2 2, 3 CO’, [northern Venezuela,] La
Puerta, 21.v.1961 (D. J. Lewis) (BMNH); 1 9 1
CO, [northern Venezuela,] Rio Aguacatal,
14.vi.1961 (D. J. Lewis) (BMNH); 1 Q, [northern
Venezuela,] Monte Oscuro, 6.v.1961 (D. J.
Lewis) (BMNH); 2 @, [northern Venezuela,]
Altamira, x.1969 (W. Biittiker) (BMNH); 1 9 (ex
pupa), Carabobo State, Birigina (no collection
date) (J. Ramirez) (BMNH); 1 ’ (ex pupa), Lara
State, Rio Tocuyo (no collection date) (J.
Ramirez) (BMNH); 2 C (ex pupae), Federal Dis-
trict, Hacienda El Limon, Divisa Colonia Tovar
(no collection date) (J. Ramirez) (BMNH).
TAXONOMIC DISCUSSION. Bellardi’s description
(1862) of Simulium mexicanum gives insufficient
detail to be able to determine the sex of the
material, which he indicates as being male with no

103

reference to number of specimens. The type status
of the specimens (holotype or syntypes) is not,
therefore, known. No subsequent authors have
examined this material because the depository
was not known. However, many descriptions of
all taxonomically important stages have since
been produced, of which that of Dalmat (1955) is
the most complete.

Bellardi’s collection of Neotropical simuliids
has now been located in the University of Turin.
Simulum mexicanum is represented by two female
specimens which are both covered in fungal
hyphae but otherwise well preserved; they are
easily recognisable as what is currently regarded
as this species. The better specimen bears one
label with the word ‘Tuxpango’ and has been des-
ignated as lectotype and labelled accordingly. The
other specimen with a single label bearing the
number ‘193’ has been labelled as paralectotype.
It is apparent that Bellardi had confused the sex of
his type specimens.

Simulium mexicanum shows little morphologi-
cal variation given its wide distribution; only the
branching height of pupal gills, though basal, can
vary within populations.

The synonymies of S. turgidum and S. aureo-
punctatum with S. mexicanum were first estab-
lished by Strong et al. (1934) although no reasons
were given. Hoffmann’s description (1930) of
turgidum and Malloch’s (1914) of aureopunctatum
are sufficiently detailed to show them to be con-
specific with the distinctive S$. mexicanum. An
examination of the well-preserved female holo-
type of S. aureopunctatum showed this specimen
to completely correspond with our modern con-
cept of S$. mexicanum, thus confirming their syn-
onymy. The synonymy of S. lugubre with S.
mexicanum is first listed in Fairchild (1940). We
have examined the only remaining syntypes (lar-
vae & pupae) in the IOC, which together with the
figures and descriptions given by Lutz & Nunez
Tovar (Lutz, 1928) all support this synonymy.
Vargas & Diaz Najera (1951b) synonymised S.
placidum with S. mexicanum following a compari-
son of two specimens in the BMNH identified by
Smart as S. placidum with specimens of S. mex-
icanum from various Latin American localities.
Our examination of the holotype of S. placidum
confirms this synonymy.

In 1934 Enderlein (1934b) described a new spe-
cies from Mexico as Hemicnetha mexicana.
Vargas (1942a) did not accept Enderlein’s generic
classification of the Simuliidae, regarding Hemic-
netha as a subgenus of Simulium, and published
the name paynei as a replacement name for mex-
icanum Enderlein (junior secondary homonym of
mexicanum Bellardi).

104

Simulium mexicanum belongs to the subgenus
Hemicnetha and is closely related to S. smarti
Vargas, although females superficially resemble
S. guianense Wise and S. orbitale Lutz in
coloration.

DISTRIBUTION. In Ecuador S. mexicanum has been
found from the western lowlands of the Andes to
higher altitudes in the north of the country.
Elsewhere it has been recorded from Belize,
Bolivia, Colombia, Guatemala, Mexico, Panama,
Tobago, Trinidad and Venezuela (Barreto et al. ,
1970; Dalmat, 1955; Fairchild, 1940; Knab, 1915;
Lewis, 1963; Ramirez Perez, 1983; Smart, 1940;
Strong et al., 1934; Vargas, 1945; Vargas & Diaz
Najera, 1957; Vargas et al., 1946; Vulcano, 1967).

Bio.ocy. Simulium mexicanum is a zoophilic spe-
cies (Strong et al., 1934; Lewis, 1963; Shelley &
Arzube, 1985) which has been reported to bite
man occasionally in Colombia (Barreto ef al.,
1970) and Venezuela (BMNH collection). In
Trinidad it bites mules and donkeys (Knab, 1915).
Simulium mexicanum and S. metallicum were
originally thought to be associated with the regu-
lar transmission of Venezuelan Equine Encepha-
litis in Colombia (Sanmartin et al. , 1973; Zuluaga
& Yuill, 1978) but recent evidence (Homan et al.,
1985) suggests that at most they only occasionally
transmit the virus mechanically.

Simulium mexicanum breeds, often in enor-
mous numbers, in small, fast-flowing streams at
both low and high altitudes usually in, or in close
proximity to, waterfalls (Barreto et al., 1970;
Shelley & Arzube, 1985).

Simulium (Psilopelmia) quadrivittatum Loew

(Figs 30, 36, 42, 48, 54, 61, 68, 74, 88, 89, 96, 102,
109; 116; 122591305 137, 145, 153)

Simulium quadrivittatum Loew, 1862: 186. LEC-
TOTYPE Q, Cusa: (collection date unknown)
(Gundlach) (MCZH, Cat. No. 12533), here
designated [examined].

Wilhelmia mallochi Enderlein, 1925: 208. LEC-
TOTYPE Q, Costa Rica: La Palma, 6.v. (no
year cited) (Biolley) (USNM, Cat. No. 8998,
Type No. 41595), here designated [examined].
[Synonymy by Vargas, 1945: 189.]

Simulium fairchildi Vargas, 1942b: 458. LEC-
TOTYPE 9, PANama: Juan Mina Station, Rio
Chagres, 8.xi.1939 (G. B. Fairchild) (MCZH),
here designated [examined]. [Replacement
name for S. haematopotum Malloch sensu Fair-
child, 1940.] [Synonymised with S. quadrivit-
tatum by Fairchild, 1943: 574.]

Description. Female. General body colour black.
Body length (alcohol preserved specimens) 2.0—

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

2.7mm (n= 55), wing length 1.6—2.2 mm (n = 55),
wing width 0.7-1.0 mm (n = 55).

Head dichoptic with red eyes; nudiocular area
well-developed (Fig. 30). Frons, clypeus and
occiput black with silver pruinosity. Mouthparts
dark brown. Antennae orange-brown. Cibarium
with sclerotised posterior margin armed with
teeth; median area of margin concave with three
stout teeth, a pair of submedian groups of about
six well-developed teeth with the central being
longer and up to tricusped; posterior margin of
cibarium between these teeth and cornuae
occupied by three to five smaller teeth; cornuae
well-developed and sclerotised (Fig. 36).

Scutum velvet-black with silver pruinose orna-
mentation, which varies in form depending on
angle of illumination. With light source anterior to

specimen silver ornamentation as follows: a pair |

of parallel, submedian bands narrowing posteri-
orly and running from the anterior scutal border
and coalescing with the silver pruinose posterior
border, a pair of dull grey cunae in form of equi-
lateral triangles present in anterior section of
these bands; a pair of sublateral parallel bands
beginning at the golden pruinose humeri and

extending to posterior pruinose margin of scutum |

(Fig. 145). With light source posterior to specimen
the grey cunae become silver and merge with the
submedian bands, the rest of the pattern being
identical to that seen with an anterior light source.
Irrespective of direction of light source S. quad-
rivittatum is unique in that the submedian pru-
inose bands show a constriction at the posterior
border of the cunae, which readily distinguishes it
from S. haematopotum and similarly ornamented
species of the amazonicum group. Paranotal folds
black with silver pruinosity. Scutum with numer-
ous adpressed golden hairs. Pleural region dark
brown with faint silver pruinosity. Scutellum vel-
vet-black with an uneven row of long black bristles
on posterior border and a group of these bristles
on each postero-lateral corner. Postnotum black
with grey pruinosity.

Subcostal vein of wing usually bare but some-
times with up to two hairs on median section and
more rarely a single hair at base of vein amongst
sensilla; basal section of radius bare (Fig. 42).
Costal base tuft of dark setae.

Legs dark brown with coxae lightly grey pru-
inose; distal tip of femur and basal tip of tibia light
brown at articulation on fore leg; distal tip of
femur of mid leg and basal half of basitarsi and
second tarsomeres cream; basal articulation of
tibiae, basal half to two-thirds of basitarsi and
basal half of second tarsomeres of hind leg cream.
Proportions of legs as in Fig. 145. Femora and
tibiae of all legs and trochanters of fore and hind

SIMULIIDAE OF ECUADOR

legs with numerous scales scattered amongst hairs
as in S. exiguum (Fig. 3). Claws curved with sub-
basal tooth (Fig. 48). Halteres bright yellow with
light brown base.

Abdominal tergites black. Tergal plates slightly
developed (Fig. 4). Basal scale (tergite I) velvet-
black, sometimes with faint silver pruinosity in
central part; tergite II black with silver pruinosity;
tergites III-V velvet-black with posterior margins
faintly silver pruinose; tergites VI-IX shiny black.
Sternites and genitalia dull black (Fig. 145).
Eighth sternite sclerotised with a group of 8-10
setae on each side, gonopophyses large, sub-
quadrangular with numerous minute setae (Fig.
54). Cerci hemispherical, paraprocts small and
subrectangular (Fig. 61). Genital fork stout with
well-sclerotised stem, expanded arms and well-
developed lateral wings with sclerotised anterior
processes (Fig. 68). Spermatheca as in S. exiguum
(Fig. 5), oval, sclerotised with no external sculp-
turing and spicules of inner surface arranged in
groups of three. Area of insertion of spermathecal
duct membranous and about one-third maximum
width of spermatheca.

Male. General body colour black. Body length 2.6
mm (alcohol preserved specimen) (n = 1), wing
length 1.8mm (n = 1), wing width 0.9 mm (n= 1).

Head holoptic with dark red eyes. Clypeus
black with silver pruinosity. Rest of head colora-
tion as in female.

Scutum velvet-black with a pair of submedian
silver cunae extending from anterior scutal border
for about two-thirds of its length (most easily seen
with light source anterior to specimen). Posterior
border and anterior three-fourths of lateral
border of scutum silver pruinose. Scutum with
adpressed golden hairs. Paranotal fold velvet-
black (Fig. 153). Coloration and setation of
pleural region, scutellum and postnotum as in
female.

Subcostal vein and basal section of R bare.

Leg coloration as in females, proportions as in
Fig. 153. Scale distribution on legs as in female.
Haltere coloration as in female.

Abdominal tergites velvet-black, basal fringe of
long black hairs. Silver pruinose ornamentation as
follows: tergite IJ completely silver, tergites III, V
and VI completely silver except for median dark
patch (Fig. 153). Sternites and genitalia dark
brown, sternal plates not examined (lack of
material). Gonocoxite slightly longer than wide;
gonostyle subtriangular, about half length of
basimere, with subterminal, rounded spine (Fig.
74). Ventral plate as in Figs 88, 89 with highly
developed lightly sclerotised basal arms and
poorly developed keel; hairs long and restricted to

105

keel; median plate slightly pyriform with small
apical incision (Fig. 96). Paramere as in Fig. 102
with several well-developed distal spines.

Pupa. Cocoon length dorsally 1.8 mm, ventrally
2.6 mm; pupa length 2.5 mm; gill length 3.6 mm
(n=1).

Cocoon slipper-shaped, dark brown opaque;
rim of aperture dark brown, reinforced and some-
times with dorsal protuberance (Fig. 109).
Cocoon surface of thin, densely interwoven
fibres. Gill mid brown with eight, forwardly-
directed, long, slender filaments; primary
branches arise in the vertical plane with filaments
arranged in the horizontal plane. The gill config-
uration description is from a slide-mounted speci-
men (Fig. 116); main trunk giving rise to three
primary branches, inner (= ventral) with two fila-
ments and median and outer (= dorsal) each with
three filaments; inner branch with bifurcation at
basal sixth of gill, median branch with first bifur-
cation at basal sixth and second at basal third,
outer branch with first bifurcation at basal tenth
and second bifurcation at basal sixth of gill; fila-
ments slender with crenate margins and rounded
distally, their surfaces covered with fine spicules
as in S. exiguum (Fig. 12). Head as in S. exiguum
(Figs 13, 14) with 2 + 2 bifid frontal and 1 + 1
simple well-developed facial trichomes; surface of
head glabrous with few platelets. Thorax as in S.
exiguum (Fig. 17) with 5 + 5 well-developed,
bifid, antero-dorsal trichomes. Surface of thorax
with few platelets. Abdominal tergite II witha line
of 4 + 4 fine hairs, I1I-IV with 4 + 4 simple hooks,
IX with 1 + 1 well-developed spines, VI-IX witha
row of well-developed spine combs on anterior
margins of segments (Fig. 19); sternite IV with 2 +
2 simple or bifid well-developed hooks, V—VII
with 2 + 2 bifid or trifid hooks, sternite VIII with 1
+ 1 antero-median patches of spine combs (Fig.
20).

Mature larva. Body length 3.44.5 mm (n = 6).
Width of head capsule 0.4 mm (n = 6). Body
colour white with grey banding (Fig. 122). Body
form as in Fig. 122. This species is easily confused
with S. /ewisi but can be distinguished by the form
of the postgenal cleft and the presence of ventral
papillae. Larval coloration in Carnoy’s fixative
was not observed due to paucity of material.
Head yellow with head spots concolorous or
forming a positive pattern as in Fig. 130 and with
sparsely distributed setae on all surfaces. Postge-
nal cleft small, triangular with thickened lateral
margins; postgenal bridge as long as hypostomium
(Fig. 137). Hypostomium as in S. exiguum (Fig.
21). Antennae long, yellow to pale brown with

106

segment ratios 15 : 20 : 25. Mandible as in S.
exiguum (Fig. 23). Maxillary palp over three times
as long as width at base. Cephalic fan with 36-40
rays. Thorax white with black collar anteriorly,
amorphous, grey patches dorsally and several
small, black patches ventrally posterior to proleg.
Cuticle glabrous ventrally but with occasional
setae dorsally. Proleg plate not observed (insuffi-
cient specimens). Pupal respiratory histoblast
dark brown and claviform.

Abdomen white with a single black ring on first
abdominal segment and on last narrow abdominal
segment, intermediate segments with incomplete
rings usually more obvious dorsally; posterior seg-
ments of abdomen with dark patches distributed
randomly, but dorsally these coalesce. Ventral
nerve cord grey. Cuticle with scattered setae on
dorsal surface of anterior (narrow) constricted
segments, more densely distributed on posterior
(wide) segments, glabrous ventrally. Ventral
papillae well-developed (Fig. 122). Anal sclerite
well sclerotised with posterior arms extending to
twelfth row of posterior circlet hooks. Posterior
circlet with 58 rows of 10-11 hooks. Anal gill
trilobed as in S. exiguum (Fig. 24) with six short
lobules on each lobe.

MATERIAL EXAMINED

Lectotype 2 and paralectotype 2 of S. quad-
rivittatum, Cuba: collection date unknown
(Gundlach) (MCZH). Lectotype 9 and paralec-
totype 2 of Wilhelmia mallochi, Costa Rica: La
Palma, 6.v. (no year given) (Biolley) (USNM) and
2 paralectotype 2 of Wilhelmia mallochi, Cuba:
Cayamas, 6.vi., 10.vi. (no year given) (E. A. Sch-
warz) (USNM). Lectotype @ and 4 paralectotype
Q of S. fairchildi, Panama: Juan Mina, Rio Cha-
gres, Canal Zone, 8.xi.1939 (G. B. Fairchild)
(MCZH), 2 paralectotype 2 and 1 paralectotype
CO of S. fairchildi, Panama: Summit, Canal Zone,
9.1.1940 (G. B. Fairchild) (MCZH) and the fol-
lowing female paralectotypes of S. fairchildi, Pan-
ama: 2 Q, Forest Reserve, Canal Zone,
11.x.1939; 1 9, Las Guacas, Panama Republic,
4.i1.1940; 1 9, Chilibre, Panama Province, Pan-
ama Republic, 23.xi.1939 (all G. B. Fairchild)
(MCZH).

Belize: 46 Q, Privation Creek, 31.ii.1958 (D. J.
Lewis) (BMNH); 3 2, Cool Shade, 11.i.1958 (D.
J. Lewis & P. C. C. Garnham) (BMNH); 6 9,
Middlesex, 8 & 17.11.1958 (D. J. Lewis) (BMNH),
1 9, North Branch, 10.ii.1958 (D. J. Lewis)
(BMNH); 3 9, Rio Frio, 5.ii.1958 (P. C. C.
Garnham & D. J. Lewis) (BMNH); numerous
females, near Cayo, Augustine, 27.vii.1961 (D. J.
Lewis) (BMNH).

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Colombia: 1 9, Choco, El Tigre, x.1959 (P. C.
C. Garnham) (BMNH).

Costa Rica: 5 2, Env. da Cartago & La Palma,
1500 m, 1906 (Biolley) (MNHN) [1 2 in BMNH
from Env. da Cartago presented by Roubaud with
incomplete data but almost certainly from the
series deposited by Biolley in MNHN]; 1 9, Tala-
manca, 22.iv.1917 (C. B. Williams) (BMNH); 4
Q, Orosi, 6.i1.1938 (K. W. Kamm) (BMNH).

Ecuador: 9 9, Esmeraldas Province, various
localities in the Santiago onchocerciasis focus, San
Miguel de Cayapas, R. Cayapa 18—-21.vi.1981 (A.
J. Shelley & M. Arzube) (BMNH, INHMT);7 9,
R. Cayapas, 28.vi.1980 (M. Arzube) (BMNH,
INHMT); 5 9, San Miguel de Cayapas, R.
Cayapa, 28-30.vi.1980 (M. Arzube) (BMNH,
INHMT); 6 larvae, San Miguel de Cayapas,
Estero Hacha, 26.v.1981 (A. J. Shelley & M.
Arzube) (BMNH); 1 @ (ex pupa), San Miguel de
Cayapas, R. Cayapa, Cascadita, 17.vi.1981 (A. J.
Shelley & M. Arzube) (BMNH); 1 CO (ex pupa),
500 m above San Miguel de Cayapas, feeder
stream of R. San Miguel, 17.vi.1981 (A. J. Shelley
& M. Arzube) (BMNH); 1 Q, Sapallo Grande
Mission, R. Cayapa, 28.v.1981 (A. J. Shelley &
M. Arzube) (BMNH); 3 Q, Naranjal, R.
Canandé, 25.ix.1983 (A. J. Shelley & M. Arzube)
(BMNH); 1 9 (ex pupa), 2 larvae, Naranjal (R.
Canandé), R. Aguas Negras, 23.vi.1985 (A. J.
Shelley & M. Arzube) (BMNH); 1 9, 1 @ (ex
pupa), Cotopaxi Province, Quevedo-La Mana-
Pilalo road, La Germania, Riachuelo, 9.vi.1984
(A. J. Shelley & M. Arzube) (BMNH); 1 9, Man-
abi Province, Santa Domingo-El Carmen road,
km 40, 2 km past El Carmen, Rio Suma, 7.vi.1984
(A. J. Shelley & M. Arzube) (BMNH); 2 9, El
Oro Province, Machala-Naranjal road, Rio
Bucay, 12.vi.1984 (M. Arzube) (BMNH).

Jamaica: 1 9, (no locality or collection date)
(H. G. Johnston) (BMNH).

Mexico: 1 2 1 Oo, Las Chuapas, ii.1947 (J.
Parra) (BMNH).

Panama: 8 92, Canal Zone, 1932 (L. H. Dunn)
(BMNH); 3 9, 1 9 (ex pupa), Chiriqui Province,
Fortuna hydroelectric station, Arroyo 47, Q10
E27 and Q47 CUSA, 18.x.1979, 18.11.1980 &
22.i1.1981 (J. L. Petersen) (BMNH).

TAXONOMIC DISCUSSION. Simulium quadrivittatum
was described by Loew (1862) from females col-
lected in Cuba, with no details of the number of
specimens examined or their depository. How-
ever, Smart (1942) noted the presence of two syn-
types in the MCZH (depository incorrectly cited
as in Berlin and not Harvard) where all of Loew’s
New World Diptera types were deposited. This
material has now been examined and the better

SIMULIIDAE OF ECUADOR

specimen designated as lectotype and labelled
accordingly. Many descriptions and misidentifica-
tions of S. quadrivittatum have been made and
these are listed with the most complete descrip-
tions of this species by Wygodzinsky (1953),
Rubtsov & Garcia Avila (1972) and Fox (1953).
Simulium quadrivittatum is most closely related to
S. haematopotum in the subgenus Psilopelmia and
in female coloration resembles S. marathrumi
Fairchild, $. oyapockense Floch & Abonnenc, S.
roraimense Nunes de Mello and S. sanguineum
Knab of the subgenus Psaroniocompsa.

In his revision of the Simuliidae of the Amer-
icas, Malloch (1914) gave a detailed description of
S. quadrivittatum based on material from Cuba,
Costa Rica, Puerto Rico and Panama in the
USNM. Enderlein (1925) later described the spec-
imens from Cuba and Costa Rica as a new species,
Wilhelmia mallochi, although it is unclear whether
he saw Malloch’s material. Wilhelmia mallochi
was later synonymised with S. quadrivittatum by
Vargas (1945) but with no explanation for this
action. Pinned specimens of W. mallochi in the
USNM have now been examined. They consist of
two females labelled as cotypes from La Palma,
Costa Rica, collected by Biolley, a female ‘cotype’
from Cayamas, Cuba, collected by Schwarz in
June (specimen and collection year missing), and
two further females from Cayamas collected in the
same period by Schwarz but with no ‘cotype’
labels. The better specimen from La Palma of
these four syntypes has been designated lec-
totype. The lectotype and paralectotypes of
Wilhelmia mallochi have been compared with the
lectotype of S. quadrivittatum and are conspecific.
The specimens in the BMNH and MNHN col-
lected in La Palma by Biolley are almost certainly
from the same collection as those examined by
Malloch in the USNM. However, they did not
form part of the syntype series of W. mallochi
because they were not cited by Malloch (1914) on
which publication Enderlein (1925) presumably
based his new species. In Fairchild’s (1940)
revision of the Simuliidae of Panama a description
of the female, male and pupa of S. haematopotum
were given although the author discussed the pos-
sibility that this was a misidentification. He noted
differences between his material and the figures of
this species by Dyar & Shannon (1927), the simi-
larity of his material to S$. pseudohaematopotum
Hoffmann, and that a previous worker in the area
had identified this species as S$. quadrivittatum.
Later, Vargas (1942b) proposed the name fair-
childi for Fairchild’s specimens because of dif-
ferences they showed in the genital fork and claw
with S. haematopotum and in the genital fork of S.
haematopotum. Fairchild (1943) later examined

107

true S. haematopotum and concluded that S. fair-
childi should be regarded as asynonym of S. quad-
rivittatum until material from the type locality
(Cuba) of the latter species could be examined.
Vargas (1945) maintained this synonymy. Fair-
child’s description and figures of S. fairchildi (as
S. haematopotum in Fairchild, 1943) indicates S.
quadrivittatum and not S. haematopotum with ref-
erence to the toothed claw and pupal gill. Fair-
child’s material has now been examined and
currently comprises the material listed as S. fair-
childi under material examined. A lectotype from
Juan Mina, Rio Chagres has been selected from
this syntype series and together with the paralec-
totypes is conspecific with the lectotype of S.
quadrivittatum. The male paralectotype of S. fair-
childi from Summit, Panama, has been compared
with a reared S. quadrivittatum from Ecuador and
is conspecific. We therefore agree with the syn-
onymy of S. fairchildi with S. quadrivittatum first
proposed by Fairchild (1943). Various other misi-
dentifications of S. quadrivittatum as S. hae-
matopotum are listed by Vulcano (1967) and
Wygodzinsky (1953).

DistriBuTION. In Ecuador S. quadrivittatum in-
habits lowland forested areas and plantations to
the west of the Andean cordillera and only occurs
in large numbers in the onchocerciasis focus in the
north of the country (Shelley & Arzube, 1985;
Leon & Wygodzinsky 1953a, b; Wygodzinsky
1953). Outside Ecuador S. quadrivittatum has
been recorded from Belize (Lewis & Garnham,
1960), Colombia (BMNH), Costa Rica (Rou-
baud, 1906), Cuba (Rubtsov & Garcia Avila,
1972), Jamaica (BMNH), Mexico (Vargas & Diaz
Najera, 1957), Panama (Petersen et al., 1983) and
Puerto Rico (Fox, 1953), sometimes at high alti-
tudes (Lewis & Garnham, 1960; Petersen ef al.,
1983). The inclusion of Venezuela by Vargas
(1945) in his distribution list for S$. quadrivittatum
is presumably based on Ortiz (1944). However,
Ramirez Perez (1983) believes that a member of
the S. amazonicum group (subgenus Psaro-
niocompsa) and not S. quadrivittatum was involved.

BroLocy. Simulium quadrivittatum is highly
anthropophilic in Ecuador (Shelley & Arzube,
1985) and Central America (Fox, 1953; Lewis &
Garnham, 1960; Rubtsov & Garcia Avila, 1972;
Petersen et al., 1983), but also bites equines
(Vargas, 1945). Simulium quadrivittatum is only
of medical importance in Ecuador where it is a
vector of human onchocerciasis (Shelley, 1988;
Shelley & Arzube, 1985). In Panama it was a
biting nuisance during the construction of a dam in
Chiriqui Province (Petersen et a/., 1983), and has
been shown to be a suitable experimental host to

108

Guatemalan O. volvulus (Schiller et al., 1984),
although human onchocerciasis has not been
recorded from Panama. It breeds in slow-flowing
streams and shows population peaks in the wet
season (Lewis & Garnham, 1960; Petersen et al.,
1983; Rubtsov & Garcia Avila, 1972; Shelley &
Arzube, 1985).

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110

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A. J. SHELLEY, M. ARZUBE & C. A. COUCH

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Figs 3-6 3, scale of leg of female S. exiguum. 4, dorsal view of female abdomen of Simulium showing position of
tergal plates (shaded). 5, spermatheca of female S. exiguum. 6, ventral view of male abdomen of Simulium showing
position of sternal plates (shaded).

SIMULIIDAE OF ECUADOR 111

11

Figs 7-11 Anterior view of head of: (7) dichoptic female S. exiguum; (8) holoptic male S. exiguum; (9) dichoptic
female S. gonzalezi; (10) dichoptic male S. gonzalezi; (11) holoptic male S. gonzalezi.

12) A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Figs 12-16 12, portion of gill filament of S. exiguum. 13, frontoclypeus of female pupa of S. exiguum. 14,
frontoclypeus of male pupa of S. exiguum. 15, frontoclypeus of female pupa of S. mexicanum. 16, frontoclypeus of
male pupa of S. mexicanum.

SIMULIIDAE OF ECUADOR 113

Figs 17-24 17, thorax of S. exiguum pupa. 18, thorax of S. mexicanum pupa. 19, dorsal view of pupal abdomen of

Simulium showing chaetotaxy. 20, ventral view of pupal abdomen of Simulium showing chaetotaxy. 21, hypostomium

of larva of S. exiguum. 22, hypostomium of larva of S. mexicanum. 23, mandible of S. exiguum larva; inset — variations
in mandibular serrations of S. bipunctatum. 24, anal gills of S. exiguum larva.

114 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Figs 25-30 Nudiocular area of: (25) S. exiguum; (26) S. bipunctatum; (27) S. lewisi; (28) S. escomeli; (29) S.
mexicanum; (30) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 115

FEERTETe sn BEF EAE

32

36

Figs 31-36 Cibarium of: (31) S. exiguum; (32) S. bipunctatum; (33) S. lewisi; (34) S. escomeli; (35) S. mexicanum;
(36) S. quadrivittatum.

116 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

_ ALLELE AA yg pep aj yifltl ghee Mp lM della api

—
———

Figs 37-42 Anterior wing veins of: (37) S. exiguum, (38) S. bipunctatum, (39) S. lewisi; (40) S. escomeli; (41) S.
mexicanum; (42) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 1a

Figs 43-54 43-48, claws of: (43) S. exiguum; (44) S. bipunctatum; (45) S. lewisi; (46) S. escomeli; (47) S. mexicanum;
(48) S. quadrivittatum. 49-54, eighth sternite and gonopophyses of: (49) S. exiguum; (50) S. bipunctatum; (51) S.
lewisi; (52) S. escomeli; (53) S. mexicanum; (54) S. quadrivittatum.

118 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Figs 55-61 Cercus and paraproct of: (55) S. exiguum; (56) S. gonzalezi; (57) S. bipunctatum; (58) S. lewisi; (59) S.
escomeli; (60) S. mexicanum; (61) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 119

Figs 62-68 Genital fork of: (62) S. exiguum; (63) S. gonzalezi; (64) S. bipunctatum; (65) S. lewisi; (66) S. escomeli;
(67) S. mexicanum; (68) S. quadrivittatum.

120 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

nal a
i
iil
AY \ |

|

71

Figs 69-74 Gonocoxite and gonostyle of: (69) S. exiguum; (70) S. bipunctatum; (71) S. lewisi; (72) S. escomeli; (73) S.
mexicanum; (74) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 121

Figs 75-89 Ventral view and profile of ventral plates of: (75-76) S. exiguum; (77-78) dichoptic S. gonzalezi; (79)
ventral view of holoptic S. gonzalezi; (80-81) S. bipunctatum; (82-83) S. lewisi; (84-85) S. escomeli; (86-87) S.
mexicanum, (88-89) S. quadrivittatum.

122 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

93 96

Figs 90-102 90-96, median sclerites of: (90) S. exiguum; (91) S. gonzalezi; (92) S. bipunctatum; (93) S. lewisi; (94) S.
escomeli; (95) S. mexicanum; (96) S. quadrivittatum. 97-102, parameres of: (97) S. gonzalezi; (98) S. bipunctatum; (99)
S. lewisi; (100) S. escomeli; (101) S. mexicanum; (102) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 123

ONAN ROM IG
ey) KN F 4

i ant a

109

Figs 103-109 Lateral view of pupa and cocoon of: (103) S. exiguum; (104) S. gonzalezi; (105) S. bipunctatum; (106)
dorso-lateral view of S. lewisi; (107) S. escomeli; (108) S. mexicanum; (109) S. quadrivittatum.

124 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Figs 110-116 Pupal gill of: (110) S. exiguum; (111) S. gonzalezi; (112) S. bipunctatum: (113) S. lewisi; (114) S.
escomeli; (115) S. mexicanum; (116) S. quadrivittatum.

SIMULIIDAE OF ECUADOR 1125

118

120

122

Figs 117-122 Lateral view of larva of: (117) S. exiguum; (118) S. bipunctatum; (119) S. lewisi; (120) S. escomeli; (121)
S. mexicanum; (122) S. quadrivittatum.

126 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

129

Figs 123-130 Larval head patterns of: (123) S. exiguum, atypical form; (124) S. gonzalezi; (125) S. bipunctatum,
typical form; (126) S. antillarum, atypical form; (127) S. lewisi; (128) S. escomeli; (129) S. mexicanum; (130) S.
quadrivittatum.

SIMULIIDAE OF ECUADOR 127

134

135

137

Figs 131-137 Ventral view of head showing postgenal cleft and bridge of: (131) S. exiguum; (132) S. gonzalezi; (133)
S. bipunctatum; (134) S. lewisi; (135) S. escomeli; (136) S. mexicanum; (137) S. quadrivittatum.

128 A. J. SHELLEY, M. ARZUBE & C. A. COUCH

Figs 138-145 Colour patterns of the female thorax, abdomen and legs of: (138) S. exiguum (xX 20); (139)
S. gonzalezi (x 20); (140) S. bipunctatum (xX 10); (141) S. lewisi (x 20); (142) S. escomeli (x 20), lowland
form; (143) S. escomeli (x 20), highland form; (144) S. mexicanum (xX 10); (145) S. quadrivittatum (x 20).

SIMULIIDAE OF ECUADOR 129

“ay : ap-

te a
J\_-
IU
aK
%
A
SS
mh
IN

—
ol
N

Figs 146-153 Colour patterns of the male thorax, abdomen and legs of (146) S. exiguum (x 20); (147) S.

gonzalezi (xX 20); (148) S. bipunctatum (x 20) (composite photograph of thorax and abdomen); (149) S.

lewisi (Xx 20); (150) S. escomeli (x 20), lowland form (anterior light source) (composite photograph of

thorax and abdomen); (151) S. escomeli ( x 20), highland form (anterior light source); (152) S. mexicanum
(x 10); (153) S. quadrivittatum (x 20).

A. J. SHELLEY, M. ARZUBE & C. A. COUCH

INDEX

Synonyms are in italics; principal references are in bold.

adolfolutzi Wygodzinsky 97

alirioi Ramirez-Pérez & Vulcano 96,
97

amazonicum group 104, 107

antillarum Jennings 90, 91, 92, 93, 126

aureopuntatum Malloch 100, 102, 103

bipunctatum Malloch 82, 83, 84,
90-93, 95, 99, 113-129

delpontei Paterson & Shannon 84, 87
dinellii Joan 90, 92

Ectemnaspis 92, 93, 97

escomeli Roubaud 82, 83, 97-100,
114-129

exiguum Roubaud 79, 83, 84-88, 89,
90, 91, 94, 95, 97, 98, 99, 101, 102,
105, 106, 110-129

fairchildi Vargas 104, 106, 107

gabaldoni Ramirez-Pérez & Vulcano
96, 97

glaucophthalmum Knab 84, 85, 87

gonzalezi Vargas & Diaz Najera 83,
85, 87, 88-89, 111, 118, 119, 121-

124, 126-129
guianense Wise 104

haematopotum, Fairchild 104
haematopotum Malloch 104, 107
Hemicnetha 100, 101, 103, 104

iguazuense Coscaron 93, 96

lewisi Ramirez-Pérez 82, 83, 90, 93-97,
99, 105, 114-129

lugubre Lutz & Nunez Tovar 100, 102,
103

lutzianum Pinto 96, 97

mallochi Enderlein 104, 106, 107

marathrumi Fairchild 107

metallicum Bellardi 88, 104

mexicanum Bellardi 83, 88, 100-104,
112-129

mexicanum Enderlein 103

Notolepria 84, 87, 88

ochraceum Walker 92, 93
Onchocerca volvulus 79, 96, 108
orbitale Lutz 104

oyapockense Floch & Abonnenc 107

paraguayense Schrottky 87, 89
paynei Vargas 103
placidum Knab 100, 102, 103
Psaroniocompsa 107
pseudoantillarum Ramirez-Pérez &
Vulcano 90, 92, 93
pseudohaematopotum Hoffmann 107
Psilopelmia 90, 93, 97, 99, 100, 104,
107

quadrivittatum Loew 79, 83, 104-108,
114-129

romanai Wygodzinsky 97
roraimense Nunes de Mello 107
rufidorsum Enderlein 97, 100

sanguineum Knab 107
Simulium 80, 103, 110, 113
Simulium sp.A Lewis 96
Simulium sp.D Lewis 93
smarti Vargas 104

turgidum Hoffmann 100, 103

Wilhelmia 104, 106, 107
wolcotti Fox 90, 92

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1 The mealybug genus Planococcus (Homentors Pseudococe
Jennifer M. Cox

79 The Simuliidae (Diptera) of the Santiago onchocerciasis focus f
A. J. Shelley, M. Arzube & C. A. Couch