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© The Natural History Museum, 1999

Entomology Series
ISSN 0968-0454 Vol. 68, No. 2, pp. 93-193

The Natural History Museum
Cromwell Road
London SW7 5BD Issued 25 November 1999

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset

Bull. nat. Hist. Mus. Lond. (Ent.) 68(2): 93-128

A revision of the African and
Malagasy species of the genus
Leptomastix (Hymenoptera, { 7
Encyrtidae), parasitoids of |
mealybugs (Homoptera:

Pseudococcidae) 3 MERIAL

JEAN-MARC ANGA
International Cocoa Organisation, 22 Berners Street, London WIP 3DB, England.
JOHN S. NOYES
The Natural History Museum, London SW7 5BD, England.
CONTENTS
IS TOP SISB a got haus teers pcacieas tata ns Sisnoale kde ual duciannsns deter sts vscqaddaaendedeadeetsscedtanassceshs catedssievacssusosacts 93
Jingite oa (UTehn Co) aTNRe eer RaSecR Ac Pace ccre cc corte CORAM tC RREMEREP ce ae er ere era Per ner PP eeere Perc Pepe neee rer ee AR 93
MEW OSTLOUIE Shires crea eras Saaece Pee AER Ae coc cvs os Maeucc du enon a Benacuite Gout Radeon dais catvcun Cov sneiceas tenses 94
PS GRNOWICG CCM OTIS Sn. c.0. estat anes cunetnas ne dew ea Desc sxe neanesavesanaduneudadnesWdsanesudvasevncdenadsenaausaac <Gsesmrdacas 94
SST SrA NMEA GUILE IMC MES eas vaaee sas cavaeeexcnawapcbsassosessec auvutesetacstansavgserecsavausvcscareettaversanssaraseeeseasssees 94
OG STUN PIECE fa ass Bre aca aa carn ees emgn aes es az ads sonatas case suneananeedutedaceeae ttaercasadisccevonecscssuescosvonze 95
DSc OHA OSES oh ges cat cuts 1d Sis Sega eh oes Seth ode eects one vncda taeeeee amteerees esac meecvaeweateten mace metueeevevossacevavenchee 95
Ifo lS {ahi (os fo) ol appear Meroe ee meer ncr Ar orrCe ct er eee MER or er crop er ree recon ot rr ore E A Ar ener eer eee eee 96
Piste bution Of SPECIES ri saatac sui hte uecnoupticdes nau snsonaee ote ee reams name So Mente vacate desta easreceee stot evans 96
15310) [ys Ape eae ERI er ore ne EPI RE cer cer co rr dase rere eee CoE 96
Wises biOC ONO 5 sscc5 cas onceecssasdaeensvesaiacssassades.scsseneverbedseetensoer dias Veaadebetaens cv Toestees cottcteseoudeTicats 96
Kev tovAti Can SHeCles) Ol: CpTOMASti Nes acss...<c---cemmar ee eee eee aoe 96
FREVASLOM OLS POGUES cnn cast ot ope te rare caven ac cv'vn svn atenen rea conan eae ama awenea wee cct ea noah oa cu ne ANAS 98
LOSLSIOL, Au GAME CP LOMAS TI mace x- x oaccase sce ncas uc <adtceevme rte retet aera ste eau seate aida nesses valeeseolees 111
PRE LETEMGC ES se sirc cv ccansdeececades ders costa. sade sc shlcXuvis slssas sis ageahemsnaastrasveaansae ni diitasy sweetsnanct ites Men wiae. tv tsse 112
TMA RALONSCLEM HURTS MANES eee theese. csanavsusecvensssnceneses@uakedersedeeusvavauen chur aencuaivenvsaaedvaedvanssedtares esas 116
MULTE Creel EL OMS Ae aedee tects eet sree eet ce esc Sadan heanonice cownannswasees uty dic tenr seve sastenceudsuituedesnspeeennasdnancanusies 117

SyNopsis. The 11 species of Leptomastix known from Africa and Madagascar are
revisedIllustrations 117. Three are described as new (herreni, africana, jonesi), three synonymies
are proposed (/yciae with nigrocincta, phenacocci with nigrocoxalis, superba with dactylopii),
and a lectotype is designated for algirica Trjapitzin. A dichotomous key to all species is provided
and each species is further characterised by a taxonomic diagnosis or description, and notes are
provided on distribution, hosts and use in biocontrol.

iy gf

X

Issued 25 November 1999

INTRODUCTION

In recent years, increasing consideration has been

given to alternative methods to the use of chemicals
for pest control resulting from growing public concern

© The Natural History Museum, 1999

about the effect of the indiscriminate use of pesticides
on the environment. This is especially so in Africa
where the continued use of pesticides may be unrealis-
tic for reasons other than environmental risk alone, eg.
poor financial resources or the remoteness of some of

94

the areas involved. In recent years, the use of natural
enemies to control pests has proven to be a valuable
alternative to pesticides. This form of control is rela-
tively cheap, self sustaining and is not harmful to the
environment. Furthermore, many biocontrol agents
can disperse very rapidly over large areas thus greatly
facilitating control in remote places. The recent spec-
tacular successes of two encyrtid parasitoids introduced
against two mealybug (Homoptera: Pseudococcidae)
pests in Africa, Phenacoccus manihoti and
Rastrococcus invadens, has done much to promote the
use of biological control on the continent (see
Neuenschwander, 1989, 1990; Herren &
Neuenschwander, 1991; Agricola et al, 1989). How-
ever, these successes have also highlighted the need
for reliable taxonomic work to facilitate the identifica-
tion of pests and their parasitoids in order to provide
early control. For instance, Rosen and DeBach (1979)
pointed out that their taxonomic revision of Aphytis
(Hymenoptera: Aphelinidae) resulted in some 35 new
species being made available for biological control
projects against armored scale pests (Homoptera:
Diaspididae). On the other hand, misidentifications of
pest or natural enemies can have potentially disastrous
consequences. The control of Phenacoccus manihoti
in Africa was delayed for several years because the
pest mealybug was initially wrongly identified as
having originated in Central America and northern
South America whereas the species was actually of
central South American origin (Herren &
Neuenschwander, 1991; Noyes & Hayat, 1994).

Worldwide, several mealybug pests have been the
subject of biocontrol projects utilising species of the
encyrtid genus Leptomastix. This genus includes about
35 described species, all of which are almost certainly
parasitoids of mealybugs and therefore have the po-
tential to be used as biocontrol agents against these
pests. To date, three species of Leptomastix have been
associated with biocontrol projects (see below), espe-
cially in Africa and in greenhouses in Europe and
North America. However, in Africa the species are
poorly known, most being difficult to identify with
any degree of certainty. The only available key to
species is that of Compere (1938) which relies heavily
on colour to discriminate between them. The present
study is therefore an attempt to provide a reliable
means of identifying the African species of Leptomastix
by means of a dichotomous key to new and known
species, together with diagnostic notes, illustrations
and information on their distribution and hosts. The
work is based on material collected from all over the
African continent including north Africa and also
Madagascar.

J-M. ANGA AND J.S. NOYES

DEPOSITORIES

BMNH The Natural History Museum, London, United
Kingdom

IEEM Instituto di Entomologia Espanol, Madrid, Spain

IITA International Institute of Tropical Agriculture,
Cotonou, Benin

INMP Universita Degli Studi Di Napoli Federico II,
Italy

MNHN_ Muséum National d’ Histoire Naturelle, Paris,
France

NMP Natural History Museum, Prague, Czech
Republic

PPRI Plant Protection Research Institute, Pretoria,
South Africa

UCR University of California, Riverside, California,
USA

USNM _ United States National Museum, Washington
D.C., USA

ZAMU_ Zoological Museum of the Aligarh Muslim

University, India.

ACKNOWLEDGEMENTS We are grateful to the following
for the loan of material: E. Guerrieri (INMP), J. Macek
(NMP), J-L. Nieves Aldrey (IEEM), G.L. Prinsloo (PPR),
M.E. Schauff (USNM), S. Triapitsyn (UCR) and C. Villemant
(MNHN). Thanks are expressed also to the Biological Con-
trol Center for Africa of the International Institute of Tropical
Agriculture, Cotonu, Peoples’ Republic of Benin, for finan-
cial support and to the Government of Cote d’ Ivoire for study
leave for the first author from the Ministry of Agriculture and
Animal Resources. We also thank the Trustees of The Natural
History Museum for providing facilities.

TERMS AND MEASUREMENTS

Clava Terminal segments of antenna, composed of one
to three segments. Segments are separated by
partial or complete sutures and are not as clearly
separated as funicle segments.

EL Maximum length of eye (see eye).

EW Maximum width of eye (see eye).

Eye The measurements of length (EL) and breadth
(EW) are the maximum and minimum diameters
respectively. The points from which the
measurements are taken should be equidistant
from the objective of the microscope (i.e. both in
focus simultaneously). Occasionally, when
stated, the length of an eye is measured in facial
view from a slide mounted specimen.

Fl, F2, First funicle segment, second funicle segment,

etc. etc.

Flagellum All segments constituting the funicle and clava
(sometimes termed flagellomeres).

Funicle This is part of the antenna between the pedicel

REVISION OF LEPTOMASTIX
and clava. There are no anelli present in
Leptomastix.

FV Relative width of the frontovertex. The

measurement is taken either in frontal view or
from above and is the measurement across the
narrowest part of the frontovertex which is
normally near the anterior ocellus.

FWL Length of forewing excluding the marginal
fringe and measured from the extreme base of
the costal cell to the apex of the wing.

FWW Maximum width of the forewing excluding the
marginal fringe and normally measured along an
imaginary line at right angles to the anterior
margin of the wing to the anal angle opposite.

GL Relative length of gonostylus.

Gono- Third valvula, or ovipositor sheath, as seen in

stylus slide-mounted material (Fig. 7).

HW Head width measured in facial view.

Hypo- __ Apical, externally visible sternite of abdomen.

pygium

Linea An oblique hairless line extending from the

calva stigmal vein to posterior margin of the forewing.

Marginal Measured from the distal margin of the

vein subhyaline break at the apex of the submarginal
vein to the distal margin of the stigmal vein
where it joins the marginal and postmarginal
veins. Occasionally the subhyaline break is
obliterated but its position can be determined by
the position of a single placoid sensillum on the
dorsal surface of the submarginal vein a little
before it joins the marginal vein. The position of
the hyaline break is immediately distad of this
sensillum.

MS Malar space or the minimum distance between
eye and mouth margin. The measurement is
taken as for the eye (above). Occasionally, when
stated, the malar space is measured in facial view
from a slide mounted specimen.

MT Length of middle tibia for comparison with
length of the ovipositor or last tergite.

MV Relative length of marginal vein.

OCL The minimum distance between the posterior
ocelli and occipital margin.

OL Relative length of ovipositor.

OOL Minimum distance between the eye margin and
the nearest posterior ocellus.

Post- Measured from the distal side of the stigmal vein

marginal where it joins the marginal vein to the apex of the

vein postmarginal vein (Fig. 6). Its apex is occasion-
ally indicated by a single erect seta, although in
anagyrines this is frequently difficult to see or
absent.

PMV Relative length of postmarginal vein.

POL Minimum distance between the posterior ocelli.

Scale-like Structures on apical segments of antennae of

structures males which are usually scale-like in appearance
(see Figs 60-68). These are almost certainly not
sensilla but ‘release and spread structures’
associated with mating (see Isodoro et al., 1996).

SL Relative length of the scape. This excludes the

95

radicle and is most accurately measured along
the internal surface of the scape.

Stigmal Measured from its proximal margin, where it

vein joins the marginal vein, to its apex (Fig. 6).

SV Relative length of stigmal vein.

SW Maximum width of scape. This is best measured
on a card-mounted specimen.

LEPTOMASTIX FORSTER

Leptomastix Forster, 1856: 34, 37. Type-species
Leptomastix histrio Mayr by subsequent monotypy.

Sterrhocoma Forster, 1856: 33, 36. Type-species
Sterrhocoma histrio Forster by monotypy. Syn-
onymy with Leptomastix by Graham, 1969a:
216-217.

Stenoterys Thomson, 1876: 115, 128. Type-species
Stenoterys orbitalis Thomson, by monotypy. Syn-
onymy with Leptomastix by Mercet, 1921: 119.

Leptomastix belongs to the encyrtid subfamily
Tetracneminae, tribe Anagyrini (see Noyes & Hayat,
1994 for diagnostic characters of these taxa) and can
be separated from all other genera of Encyrtidae using
the following combination of character states (charac-
ters separating Leptomastix from other anagyrine
genera in italics).

DIAGNOSIS.Body moderately robust and stout, yellow,
orange, dark brown or completely black; mandibles
bidentate; maxillary palpi 4-segmented; labial palpi 3-
segmented; frontovertex between top of antennal
scrobes and anterior ocellus with fairly regular,
hexagonally reticulate sculpture of mesh size, about
equal to that of an eye facet; eyes not reaching occipi-
tal margin, normally separated by at least the diameter
of an ocellus; eyes inconspicuously hairy, the hairs
sparse and shorter than diameter of a facet; funicle 6-
segmented; mesoscutum without notauli; sculpture on
mesoscutum and scutellum finely reticulate but not
hexagonally reticulate; linea calva interrupted, but
not closed; filum spinosum absent; forewing normally
with postmarginal vein at least as long as stigmal,
rarely shorter; hindwings about 4—6x as long as broad;
Female: scape subcylindrical, at least 4x as long as
broad; all funicle segments clearly longer than broad,
F6 shortest; clava 3-segmented; costal cell narrow
with only a single complete line of setae ventrally;
forewings varying, about 2.5-4x as long as broad;
gaster with hypopygium reaching apex and paratergites
present; ovipositor not longer than mid tibia and never
exserted. Male: all funicle segments much longer than
broad and clothed in long setae, normally at least about
3x as long as diameter of segments; F'6 without scale-
like structures ventrally (but see digitariae, Fig. 65);

96

clava solid and with a line scale-like structures ven-
trally in basal half (see Figs 61-68); forewings normally
about 2.5—3x as long as broad; phallobase with a pair
of apical digiti, each with two or three hooks (Figs 69,
70).

IDENTIFICATION OF SPECIES. Identification keys to
species of Leptomastix include those of Compere,
1938 (Afrotropical); Trjapitzin, 1978 (western
Palaearctic); Trjapitzin, 1989 (Palaearctic) and Noyes
& Hayat, 1994 (Oriental).

DISTRIBUTION. Leptomastix includes about 35 de-
scribed species and is almost exclusively Old World in
distribution. The only possible exception is
Leptomastix dactylopii, first described from Brazil
and introduced throughout the New World and else-
where. However it is very likely that this species is of
Afrotropical origin (see Noyes & Hayat, 1994). The
known species are fairly evenly distributed between
the Palaearctic (12), Africa and Madagascar (11) and
the Oriental region (16) with only three species known
from Australia and New Zealand (one introduced),
with some overlap between these regions.

BIOLOGY. Several workers have investigated the bi-
ology of Leptomastix species. Their records suggest
that they are primary, solitary endoparasitoids of
mealybugs (Moursi, 1948; Zinna, 1959). Bess (1939)
and Lloyd (1958) recorded Leptomastix dactylopii
Ovipositing in several mealybug species while Bess
(1939) and Su & Li (1993) found that, of the hosts
tested, it would only develop successfully in
Planococcus citri. However, numerous other mealybug
species have been recorded as hosts of dactylopii (see
Noyes & Hayat, 1994). Lloyd (1964) showed that host
preference cannot be changed by rearing the parasitoid
for several generations on an ‘unnatural’ host species.
Van Baaren and Nenon (1994) reported some param-
eters relating to host discrimination and potential
superparasitism in Leptomastix species, arguing that
females can distinguish between unparasitised and
parasitised hosts and generally avoid the latter more
often, whatever the time elapsed since the first ovipo-
sition. Discrimination seems to be based on several
markers which interact either simultaneously or suc-
cessively. Kirkpatrick (1953) and Jong & Alphen
(1989) reported that a female of Leptomastix dactylopii
can lay up to ten eggs per day and that oviposition can
take place into third instar or adult mealybugs. Fecun-
dity is greatest at about 30°C (Tingle & Copland,
1989) and the egg is coated with minute spherulae
which contain a protein which probably prevents en-
capsulation of the egg by the host (Barbier & Rambault,
1985; Barbier et al., 1988). Zinna (1959) and Moursi
(1948) respectively reported four larval instars for
Leptomastix dactylopii and six for nigrocoxalis
(=phenacocci). According to them, the number of

J-M. ANGA AND J.S. NOYES

segments varies from 11 to 13, the last four not clearly
segmented in the earlier instars. Lloyd (in Zinna,
1959) reported that at 27°C, development in
Leptomastix dactylopii is complete in a little over two
weeks, development of the male generally being com-
pleted a day or two before the females. In Leptomastix
nigrocoxalis (=phenacocci) the rate of development is
linked to the age of the host into which oviposition has
occurred. In 10-day-old hosts at 27°C, development
takes 30 days, whilst in 20-day-old hosts, develop-
ment is complete in 20 days (Moursi, 1948). Su & Li
(1993) reported that the sex ratio of progeny of
Leptomastix dactylopii was female biased, with males
generally developing in smaller hosts. Adults of
Leptomastix nigrocoxalis (=phenacocci) can live up to
70 to 80 days, females generally living longer than
males (Moursi, 1948). In adults of Leptomastix
dactylopii, longevity is slightly greater at 26°C.
Battaglia and Tranfaglia (1994) reported that isolated
adults of Leptomastix dactylopii lived longer than
those in other conditions. Survival was strongly af-
fected by diet; longevity without food was nearly three
days while the highest values of longevity were re-
corded for adults fed with honey solution and pure
honey.

USE IN BIOCONTROL. The use of the three species of
Leptomastix which have been associated with
biocontrol projects is summarised in Table 1.

Key to African species of Leptomastix
(females and males)

1 Females: clava 3-segmented; flagellum with short
AP Pressed Wasws) 0.1. .ccsceseecscevese-s-22cescseeseeetee epee meavereee 2

— Males: clava entire; flagellum with whorls of long setae,
each longer than diameter of segments. .............-.:...+++ 12

FEMALES

2 Head completely yellow, without even dark areas on
temples behind eyes; forewings at least about 3.5x as
long as broad .............. tsukumiensis Tachikawa (p. 98)

— Head with at least temples behind eyes brown, some-
times completely blackish; forewings at most about 3x as
| Ca) aff e219] 0) (Ox: (0 eee eee Re is a Pree eaters ca 3

3 Forewing with postmarginal vein not more than one-
third longer than stigmal vein (Figs 11, 18) ................ 4

— Forewing with postmarginal vein at least about 1.5x as
long as stigmal vein (Figs 25, 29, 35, 43, 53, 59) ....... 6

4 Linea calva interrupted by at least 5 lines of setae (Fig.
10); tegulae with base white and apex brown ...............
cd SSR Ss 3a, oclna eae or RE digitariae Risbec (p. 100)

— Linea calva interrupted by at most 3 lines of setae (Fig.
17); tegulae with base orange and apex greyish brown or
Completely bro wmlees -cecse eateries cece eee eee 5

REVISION OF LEPTOMASTIX

Table 1.
Parasitoid

dactylopii

abyssinica

nigrocoxalis

* Parasitoid misidentified as Leptomastix flava (see below under L. dactylopii)

Target pest

Planococcus citri

Planococcoides
njalensis
Planococcus kenyae
Pseudococcus
comstocki
Pseudococcus
calceolariae

Planococcus citri

Country/Region

Australia (Qd)
Belgium
Bermuda

USA (California)
Canada

Chile

Costa Rica
Cyprus

UK

Georgia
Greece
India

Israel
Israel
Italy

Morocco
Netherlands

Pakistan
Portugal
South Africa
Spain

Sweden
Turkmenia

USA (California)
USA (Florida)
USA (Texas)

Uzbekistan
Yugoslavia (former)

Ghana

Kenya
USA (California)

Chile

USA (California)

Maconellicococcus hirsutus

Nipaecoccus viridis

Jordan

Year

1980
1989
1952
1934
1939-1945

1930-1958

1968
1966-1977

1982-1987

196]
?

1983-1985

1984, 1987
1941
1956-1980

2
1984

1985-1985
?

1935
1948,1977

1989-1990
1968
1934
1934
1970-1983

1959-1960

q
1949-1955

1937,1938
1973-1975

1931

1931

Egypt
1986

Result

SCI
SCG

SCG

SCG

SCI

A summary of the worldwide use of Leptomastix species in biocontrol projects.

Reference

Smith et al. (1988)

Ronse (1990)

Cock (1985)

Compere (1939b), Bartlett (1978)
Baird (1938,1939,1940a,1941,
1942,1943,1944,1946,1947),
McLeod (1962),

Turnbull & Chant (1961)

Duran (1944), Gonzalez & Rojas
(1966), Graf Marin & Pena (1940)
Molina (1977), Noyes & Hayat (1994)
Cock (1985), Greathead (1976),
Krambias & Kotzionis (1980)
Copland (1983), Copland & Varley
(1987), Tingle & Copland
(1988a,b, 1989)

Kobakhidze (1965)

Jourdheuil (1986)

Manjunath (1985b), Krishnamoorthy &
Singh (1987), Ramesh (1987),
Prakasan (1987)

Argov & Rdéssler (1988), Rubin (1985)
Rivnay (1960)

Greathead (1976), Luppino (1979),
Viggiani (1975a,c), Barbagallo et
al.

(1983), Longo & Benfatto (1982)
Jourdheuil (1986)

Hennekam er al. (1987):

Kole & Hennekam (1990)

CIBC (1985,1986,1987)
Jourdheuil (1986)

Greathead (1971)

Carrero (1980a,b), Greathead (1976),
Gomez Clemente (1951)

Noyes & Hayat (1994)

Niyazov (1969b)

Bartlett (1978)

Watson & Thomson (1940a,b)
Meyerdirk et al. (1978),

Summy et al. (1986)

Rozanova & Loseva (1963)
Jourdheuil (1986)

Greathead (1971)

Greathead (1971)
*Meyerdirk & Newell (1979)

Marco (1959), Rojas (1967)

Compere (1931)

P Kamal (1951)
Meyerdirk et al. (1988)

? —no subsequent information; E — established but no further information available; NC — established but no significant control
achieved; NE — not established; NR — not released; P — partial control achieved; PG — partial control in greenhouses only; R —

released but no further information available; SC — successful control achieved; SCG — successful control in greenhouses only; SCI
— successful control by inundative releases only.

98

10

11

12

14

Scape less than 7x as long as broad (Fig. 15); mesoscutum
clothed in fairly conspicuous translucent, silvery setae;
forewing with marginal vein about as long as postmarginal
(Eig) eens herreni Anga & Noyes sp. n. (p. 101)

Scape at least 9x as long as broad; mesoscutum clothed in
brown setae; forewing with marginal vein about 2x as
long as postmarginal vein ... algirica Trjapitzin (p. 102)

Antenna with F1 at least 4x as long as pedicel (Fig. 22)
wae yndseateasetticeents’ africana Anga & Noyes sp. n. (p. 103)

Antenna with F1 not more than 3x as long as pedicel
CEWES 28) BF B35 SE ADs S10) S19) ececocecocecectcontseccosersaspee i]

Bases of mandibles blackish, occasionally mouth margin
also blackish; forewings completely hyaline, without any
longitudinal fuscous streaks (Fig. 27, 34) «0... 8

Bases of mandibles yellow or orange, mouth margin
never blackish; forewings frequently with two longitudi-
nal fuscous streaks (Figs 41, 51) ......eeeeceeeeseeseeeeeeeees 9

Distance of posterior ocelli from eye margin (OOL) less
than 1.4x distance from occipital margin (OCL); ocellar
area of frontovertex orange, axillae orange ...................
see ebeeseces es permanente nigocoxalis Compere (p. 00)

Distance of posterior ocelli from eye margin more than
1.4x distance from occipital margin; ocellar area of
frontovertex and axillae dark brown ..........:.:cescceceeeees
nad due wasuctusnabevdase sensgesesteeceuweden ove nigra Compere (p. 103)

Mid coxae completely yellow ...........:.::ccceseeeseeseeeeeeeees
biases Sd esta ease daca as dertereaes dactylopii Howard (p.106)

Mid coxae at least partly brown ..............:.:ssseseeeee 10

Linea calva interrupted by at least 4 lines of setae; mid
coxae generally orange and only brown laterally ..........
ave eneeet neuen sae enee Jonesi Noyes sp. n. (p. 108)

Linea calva interrupted by not more than 4 lines of setae
(Figs 52, 58); mid coxae completely brown or almost so
stg wecnaus tes Naeater et tts saNee cteaten ae cec vet gt ine st Mea deta ea ee 11

Hind and fore coxae concolorous, yellow ................0
ie ee eee ee nigrocincta Risbec (p. 109)

Hind coxae brown contrasting with yellow fore coxae
cect setnbcocedttelerstrs aera’ abyssinica Compete (p. 110)

Forewings with postmarginal vein not longer than stig-
mal vein; head mostly dark brown (Figs 14, 20, 21) .....

Either forewings with postmarginal vein clearly longer
than stigmal vein (Figs 31, 38, 46, 55, 60) or head orange
without any brown afeas ............scceceeceeeeeseeneeeeneeneeeee 15

Tegulae white with brown apex; F6 with scale-like struc-
(HETCEAS (LEN (OB) coseetbecoccsoonccoacccaceenoreete digitariae (p.1 00)

Tegulae with base orange or completely dark brown;
only clava with scale-like structures (Figs 61, 62) (some-
TOMES AP PabeNLly TaD SCL) leaeeeeee tenes eee enn 14

Clava with 5-7 longish, slender, recurved scale-like

I

20

21

J-M. ANGA AND J.S. NOYES

structures (Fig. 61); head and thorax usually largely
C0) 2101 2 «ase ee Per SSO ea APF herreni (p. 101)

Clava with 5—7 short, straight, scale-like structures (Fig.
62); head and thorax dark brown......... algirica (p. 102)

Fain di COX ae ONO WN Feeeeess oes es ceo enon eee eee 16
Hindicoxae yellow ceeacat asec co oes cece eee ee 17
OLE COKAe DLOW Dee ceceree eee eee africana (p. 103)
ore:coxae yellow jcc abyssinica (p. 110)
Temples and occiput yellow .......... tsukumiensis (p. 98)
Temples and occiput largely blackish ...............0.00+ 18

Scale-like structures arranged in line at base of clava
very slender and elongate and difficult to distinguish
from setae) (igh O4) x seneerce eee Jonesi (p. 108)

Scale-like structures arranged in line at base of clava
flattened and short and relatively easy to distinguish
from setae | (Eigs163; (66,67) s..1ncee eee eee eens 19

Longest setae on F1 at least 4x as long as diameter of
segment measured in middle (Fig. 63); mesosternum
dark brown, almost black, mid coxae yellow; forewing
with linea calva interrupted by not more than two setae
(Pig 245) ee seaeacs. coe te tcensonedeesssee dactylopii (p. 106)

Longest setae on Fl less than 4x as long as diameter of
segments measured in middle (Figs 66, 67); mesoster-
num orange, or pale brown, mid coxae sometimes brown;
forewing with linea calva normally interrupted by at least
three setae, frequently by three or more lines of setae
(Figs 30,38; S5)) <scss:sccstencesstupeettec eee ee eee 20

Forewing with linea calva interrupted by at most only
three or four setae (Fig. 55); mandibles and mouth mar-
PIN MEVer Gankened esccceeeoceeseeeees nigrocincta (p. 109)

Forewing with linea calva interrupted by at least five
setae (Figs 31, 38); bases of mandibles and mouth margin
SOmetimMeS| DlACkas haan ccczeceecenccescee eee ae acne ee eee necro 21

Frontovertex and axillae almost completely orange ......
ee ec AP an SR PP PN nigrocoxalis (p. 103)

Frontovertex and axillae dark brown....... nigra (p. 105)

REVISION OF SPECIES

Leptomastix tsukumiensis Tachikawa
(Figs 1-7, 68)

Leptomastix tsukumiensis Tachikawa, 1963a:66-68.

Holotype, 9 Japan, ELKU, not examined.

Leptomastix singularis Shafee,
Holotype 9, India, ZAMU, not examined).
Synonymized with tsukumiensis by Noyes & Hayat
(1994).

1971:50-51.

[Leptomastix longipennis Mercet; Otanes, 1935:503—

DIAGNOSIS.

504. Misidentification. ]

Female (length 1.42-2.41): body

REVISION OF LEPTOMASTIX

slender; including gaster generally dusky yellow to
orange; head dusky yellow; mesopleuron orange,
concolorous with mid coxa and mesosternum; forewing
hyaline (Fig. 3) or with variable fuscous markings
adjacent to anterior and posterior margins distad of
venation (Figs 4, 5), sometimes also adjacent to linea
calva (Fig. 5); posterior ocelli equidistant from eye
and occipital margins or hardly closer to eyes; antenna
(Figs 1, 2) with F1 generally at least about 2.5x as long
as pedicel; forewing comparatively long and narrow,
from slightly more than 3x as long as broad to about
4x (Figs 3-5); postmarginal vein varying from about
as long as stigmal vein to about 1.3x as long and
varying from slightly longer than marginal to a little
shorter (Figs 6, 7); linea calva interrupted by 4 or 5
lines of setae (Figs 3—5); gaster as long as thorax. Male
(length 0.90-1.80 mm): generally similar to female;
funicular segments clothed in relatively short setae
(Fig. 68), each generally not longer than twice as long
as diameter of segments, although this can be variable
(see below); a line of about 10 scale-like structures
present on the basal half of clava; forewing completely
hyaline, about 3.0x as long as broad; marginal vein
slightly longer than postmarginal which is subequal to
stigmal vein in length; aedeagus about one-third as
long as midtibia; phallobase with elongate digiti, each
with two elongate apical spines.

VARIATION. Slightly more extensive than in
extralimital material (see Noyes & Hayat, 1994) in
that in the female the forewing may vary from com-
pletely hyaline to almost uniformly dusky or with a
distinct fuscous pattern (see Figs 3—5), the posterior
ocelli may be conspicuously closer to the eye than
occipital margin, the funicle segments vary from rela-
tively slender (Fig. 1) to distinctly flattened (Fig. 2)
with Fl varying from only 4x to nearly 7x as long as
broad and the forewing may be only slightly more than
3x as long as broad. The males show some consider-
able variation in the density and length of setae on the
funicle, some specimens having the setae relatively
dense and short (about 2x as long as diameter of
segments, see Fig. 68) whilst in others they may be
relatively sparse and long (about 4x as long as diam-
eter of segments, similar to Fig. 63).

Hosts. Leptomastix tsukumiensis has been recorded
from an unknown mealybug on Citrus sp. by Shafee
(1971) and from Ferrisia virgata by Otanes (1935).
This species is also recorded for the first time in
Africa, from undetermined hosts.

DISTRIBUTION. Gambia, Ivory-Coast, Benin, Ni-
geria, Cameroon, Somalia, Kenya, Zambia, Zimbabwe,
South Africa, Madagascar, India, Laos, Philippines,
China and Japan.

99

MATERIAL EXAMINED.

Non-type material. Gambia, o&2, Bakau, 1.1978
(Huggert); 10, Fajara, 1.1978 (Huggert); Ivory Coast,
1c, Lamto, 6°13'N 5°02'W, 16.v.1985 (Rasplus); 8 & ,
130°, Lamto, 6°13'N 5°02'W, xi.1988 (Noyes); 1,
Bouake, xi.1981 (Cochereau); R.P. Benin, 1c, 25Km
N Ohicon, 4.xii.1988 (Noyes); Nigeria, 119, 200’,
Kaduna St., 20 Km N Kaduna, 8.xi.1987 (Noyes,
Neuenschwander); 10°, 81Km NW Jos, 1.xi.1987
(Neuenschwander); 62 , 60", Plateau St., 20 Km NW
Jos, 12.xi.1987 (Noyes, Neuenschwander); 12, 60,
Bauchi St., 74 Km W. Bauchi, 14.xi.1987 (Noyes,
Neuenschwander); | 9 , Sokoto St., 54 Km E. Sokoto,
8.x1.1987 (Noyes, Neuenschwander); Cameroon, | 9 ,
Sanyere, 18.x11.81 (Compton); Somalia, 19, Lr.
Shabelli Valley, Mogadiscio, v.1977 (Bin); Uganda,
10, Serere, 22. vii. 1934, BM 1935-459 (Ford); Kenya,
19, Nairobi, Karen, vi.1982 (Dewhurst); Zambia,
12, 15Km E Lusaka, 11—19.ii.1980 (Beaver); Zim-
babwe, 149, 20’, Harare, Chishawasha, i.8 1—xii.84
(Watsham); South Africa, 32, Cape Province,
Swellendam, 17.xii.31—18.i.32 (Turner), 169, 9,
Cape Province, Ceres, various dates iii.1921—iv.1925
(Turner); 19, Cape Province, Matjesfontein, 7—
13.xi.1928 (Turner); 19, Cape Province, summit of
Du Toits Kloof, 4.v.1972 (BMNH Southern African
Expedition); 19, Natal, Kloof, 1500ft (=572m),
ix.1926 (Turner), 19, Natal, Cape Vidal, 28°10'S
32°32'E, 13.1.1981 (Prinsloo); 3 9 , Tvl, D’Nyala Nat.
Res. Ellisras District, 23°45'S 27°49'E, 23—26.ii. 1987
(Prinsloo); 19, Natal, Vernon Crookes Nat Res.,
Umzinto, 30°17'S 30°37'E, 443m, 25-26.111.1985
(Prinsloo); 19, C.P., Andries Vosloo Kudu Res. nr.
Grahamstown, 33°07'S 26°38'E, 30.xi.1983 (Prinsloo,
Grobbelaar); 29, Tvl, Blyderivierspoortdaam Nat.
Res., 24°32'S 30°47'E, 25-26.x.1984 (Prinsloo); 1,
C.P., Grahamstown, 33°19'S 26°32'E, 13.1.1983
(Prinsloo, Grobbelaar); | 9 , Tvl, Hans Merensky nat.
Res., 23°40'S 30°39'E, 27-30.i.1981 (Prinsloo); 1°,
Tvl, Mogol Nat. Res., Ellisras District, 23°58'S
27°A45'E., 21.1.1983 (Prinsloo); 10°, Tvl, Pretoria,
T3754, iii.1971 (Annecke); 1 9 , Tvl, Letsitele, xi.1978
(Prinsloo); also material from India, Laos, China and
Philippines as detailed in Noyes & Hayat (1994).
Material in BMNH, PPRI.

COMMENTS. In Africa, Leptomastix tsukumiensis can
be recognized by its relatively narrow forewing which
is usually infuscate along its anterior margin distad of
the venation. Itis most similar to Leptomastix dactylopii
and can be separated on the coloration of the temple
and mesosternum and relative lengths of the forewings.
In tsukumiensis the temples and mesosternum are
yellow and the forewings are 3.3-4.1x as long as
broad whilst in dactylopii the temple and mesoster-
num are dark brown and the forewings are 2.6—3.0x as
long as broad.

100

The considerable variation in both sexes as outlined
above initally led us to believe that at least two species
were present. After careful examination of all the
material at hand we have been unable to recognise any
discrete groups. Intermediate forms for all combina-
tions of the variable character states are present in both
African and Oriental material and therefore we are
assigning all material listed above to tsukumiensis.

Leptomastix digitariae Risbec
(Figs 8-14, 65, 69)

Leptomastix digitariae Risbec, 1959:23—6. Lectotype
9 , designated by Noyes & Prinsloo (1998), Mada-
gascar, examined, MNHN.

DIAGNOSIS. Female (length 1.70-1.85 mm): body,
including gaster, dark-brown to black; head and dor-
sum of thorax clothed in white setae; tegulae yellow
basally; legs brown, except mid tibia which is yellow;
posterior ocelli closer to eye margin than occipital
margin; antenna (Fig. 8) with Fl about 2x as long as
pedicel, segments subequal in length; forewing (Fig.
9) hyaline and about 2.6x as long as broad; linea calva
interrupted by five lines of setae; postmarginal vein
slightly shorter than either marginal or stigmal veins
(Fig. 11); gaster as long as thorax; ovipositor slightly
exserted. Male (length 1.34—1.42 mm): generally simi-
lar to female but antenna (Fig. 65) with setae on
funicle longer, up to 3.5x as long as diameter of F1;
scale-like structures on F6 and clava.

FEMALE (PARALECTOTYPE). Body, including gaster,
dark-brown to black; head and thoracic dorsum clothed
in white setae; mandibles brown; antennae dark-brown;
coxae dark brown; legs generally dark brown except
for distal apex of mid femur and entire mid tibia which
are yellow; mid tarsi testaceous; subalar sclerites pale-
yellow; tegulae whitish basally, dark brown apically.

Head with inner eye margins diverging ventrad;
frontovertex with regular, polygonally reticulate sculp-
ture of fine mesh size which is on average a little
smaller than an eye facet; piliferous punctures small,
but distinct; posterior ocelli closer to eye margin than
occipital margin; antennae (Fig. 8) as long as body;
scape 5x as long as broad; pedicel 2x as long as broad;
funicle segments subequal in length, at least 2x as long
as broad; F1 1.6x as long as pedicel; clava 1.7x as long
as FI and twice as long as F6. Relative measurements:
HW 85; HL 75, FV 35,,POL 155, OCLs, OOLG, EBL
52, EW 37, MS 23, SL 43, SW 12.

Thorax with mesoscutum regularly clothed in white
setae; mesoscutum and scutellum with similar sculp-
ture to frontovertex; wings hyaline; forewing (Fig. 9)
about 2.6x as long as broad; linea calva interrupted by
5 lines of setae (Fig. 10); postmarginal vein (Fig. 11)
about 0.75x as long as either marginal or stigmal
veins; hindwing hyaline, 4.6x as long as broad.

J-M. ANGA AND J.S. NOYES

Gaster as long as thorax, with ovipositor (Fig. 12)
slightly exserted, the exserted part slightly shorter
than the mid tibial spur.

MALE (PARALECTOTYPE). Similar to female, differ-
ing slightly as follows. Posterior ocelli equidistant
from eye and occipital margins; linea calva interrupted
by 3 lines of setae; antennae (Fig. 65) clothed with
long setae, each about 4x as long as diameter of
segments; F6 ventrally with 5 scale-like structures and
clava with two. Aedeagus and phallobase as in Fig. 69.

VARIATION. Little in material available.

Hosts. Leptomastix digitariae was reared originally
from mealybugs on leaves of Digitaria humberti.
(Poaceae) (Risbec, 1959).

DISTRIBUTION. Madagascar.

MATERIAL EXAMINED.
Type material. Lectotype 9 originally mounted in slide
labelled “Leptomastix digitariae Risbec 1079’
‘Leptomastix nigrocoxalis Compere 1079’ ‘1079 1 ex,
Leptomastix nigrocoxalis Compere 13 Leptomastix’,
remounted on card 21.vii.1988 by JSN and labelled
‘LT Q’ on ventral side of card. Paralectotypes: 89,
60", same data as lectotype and remounted on cards.
According to Risbec (1958:26) the type material was
reared in Madagascar from mealybugs on the leaves of
Digitaria humberti by Renaud Paulian. Lectotype and
paralectotypes in MNHN, 1 9, 2° paralectotypes in
BMNH.

Non type material. Madagascar, | 9 , Tananarive,
Tsimbazaza Gdns, 27.v.1983 (Noyes, Day). Material
in BMNH.

COMMENTS. Leptomastix digitariae, although super-
ficially similar to several species of Leptomastix may
be incorrectly placed in the genus and is retained in
Leptomastix pending further work on the phylogenetic
relationships of the taxa currently included in the tribe
Anagyrini. A preliminary phylogenetic analysis of the
taxa of this tribe undertaken by the first author was
inconclusive with regards to the recognition of two
separate lineages identified previously (see Noyes &
Hayat, 1994). These lineages were defined on the
sculpture of the frontovertex and the distribution of the
scale-like structures on the antenna of the male. The
lineage including Leptomastix includes species which
have regular polygonal sculpture on the frontovertex
and males with scale-like structures on the clava only.
The second lineage, which includes Anagyrus, is de-
fined by irregular, fine sculpture on the frontovertex
and presence of scale-like structures on F6 of the male
antenna. In L. digitariae the sculpture is somewhat
intermediate, and scale-like structures are present on
F6.

Leptomastix digitariae is superficially similar to
Leptomastix herreni, both sharing the same general

REVISION OF LEPTOMASTIX

coloration of the body (dark-brown to black), white
setae on the head and relatively short postmarginal
vein of the forewing. Apart from the sculpture of
frontovertex and distribution of scale-like structures
on the antenna of the male as stated above, they can be
separated by the relative position of the posterior
ocelli, the colour of the tegulae and in the females by
the number of lines of setae interrupting the linea
calva. In digitariae the posterior ocelli are closer to the
eye than the occipital margin and the tegulae are
basally yellow and the linea calva of the female is
interrupted by five lines of setae (Fig. 10) whilst in
herreni the posterior ocelli are closer to the occipital
margin than the eye, the tegulae are completely brown
and the linea calva of the female is interrupted by only
three lines of setae (Fig. 17). There is also some
similarity between digitariae and algirica (see com-
ments under algirica).

Leptomastix herreni Anga & Noyes sp. n.
(Figs 15-20, 61)

DIAGNOSIS. Female (length 0.50—1.70 mm): colour
varying from almost completely orange to almost
completely dark brown (see below); at least mid and
hind coxae orange-brown to dark brown; wings hya-
line; inner eye margin diverging ventrad; minimum
width of frontovertex 0.8—1.2x eye length; posterior
ocelli closer to occipital margin than eye margin; F1
about as long as pedicel to twice as long; forewing
about 2.5x as long as broad (Fig. 16); postmarginal
vein only about as long as either marginal or stigmal or
even slightly shorter (Fig. 18); linea calva interrupted
by 2 or 3 lines of setae (Fig. 17); gaster as long as
thorax. Male (length 0.50—1.35 mm): generally simi-
lar to female in coloration, but each torulus always
connected to mouth margin by a yellowish streak even
in the darkest specimens; antennae (Fig. 61) clothed
with long setae, the longest about 3x as long as diam-
eter of Fl, Fl twice as long as pedicel, basal half of
clava with about 8-10 very fine, recurved scale-like
structures ventrally; forewings about 2.3x as long as
broad; postmarginal vein very nearly as long as stig-
mal (Fig. 21).

FEMALE (HOLOTYPE). Length 1.48 mm. Head dark-
brown, with inner eye margins yellow; temples
dark-brown, occiput dark orange-brown, dark brown
adjacent to temples; clypeal area pale orange, with
white setae; mandibles dark-brown; radicle dark-
brown; scape brown dorsally and dusky-orange
ventrally; flagellum unicolorous brown; dorsum of
thorax dusky-orange, dark brown on dorsal part of
pronotum, anterior margin of mesoscutum, axillae,
anterior median part of scutellum and metanotum;
mesopleuron orange; prosternum anteriorly and mes-
osternum brown; mid coxa orange-brown; fore coxa

101

brown anteriorly; subalar sclerites below base of
forewing dusky-yellow; propodeum dark brown with
spiracular area orange; gaster dark-brown.

Head with inner eye margin diverging ventrad;
width of frontovertex 0.85x eye length; posterior ocelli
closer to occipital margin than eye margin; antennae
(Fig. 15) 0.9x as long as body; scape 5x as long as
broad; F1 about as 1.5x as long as pedicel; clava 1.5x
as long as Fl and 2.1x as long as F6; pedicel 1.9x as
long as broad. Relative measurements (holotype): HW
101, HH 92, FV 50, POL 19.5, OCL 5, OOL 11, EL 58,
EW 37, MS 28, SL 52, SW 10.

Thorax and gaster: mesoscutum and scutellum with
finely reticulate sculpture; wings hyaline; forewing
(Fig. 16) about 2.6x as long as broad; linea calva
interrupted by two or three lines of setae (Fig. 17);
postmarginal vein about 0.7x as long as either mar-
ginal or stigmal veins (Fig. 18); hindwing 4.2x as long
as broad; gaster as long as thorax, with ovipositor
hidden. Relative measurements: FWL 230, FWW 90;
HWL 153, HWW 36.

Gaster as long as thorax, with ovipositor (Fig. 19)
hidden. Relative measurements (paratype): OL 35,
MT 51.

MALE. See diagnosis. Relative measurements: HW
AS, FV 27, POL 10, OOL 7, OCL 25, EL 23, EW 16,
MS 14, SL 20, SW 5; FWL 103, FWW, 45.

VARIATION. The most noticeable variation is that of
colour, in both sexes from almost completely orange
with only the mandibles, interantennal prominence,
occiput, pronotum, tegulae and gaster brown to com-
pletely brown, including all coxae and femora, but not
the tibiae and tarsi which are always more or less
orange. Morphological variation is mostly related to
size. The smallest females have a slightly wider
frontovertex, antennae about as long as the body, and
the postmarginal vein slightly longer than either the
marginal or stigmal veins. Some larger females may
have the scape only about 4x as long as broad and F1
about 2x as long as the pedicel. Smaller males tend to
have relatively shorter setae on the funicle and fewer
scale-like structures on the clava.

Hosts. Leptomastix herreni recorded below from
Octococcus sp. on Compositae and ‘mesem’, from
Octococcus pentziae on Pentzia globosa, Octococcus
sp. nr pentziae on Athenasia trifurcata, Octococcus sp.
on Oedera imbricata, and unidentified mealybugs on
Rehlania ericoides, Chrysocoma ?coma-aurea, and
?Athenasia sp.

DISTRIBUTION. South Africa.

MATERIAL EXAMINED.

Type-material: Holotype? , South Africa, C.P.,
Klipheuwel, ex Octococcus sp. on Compositae, T2783,
x.1968 (Insley). Paratypes: South Africa, 99, 5c’,

102

C.P., x.1968, ex mealybug on Rehlania ericoides,
T2779, x. 1968 (Insley); 22, 2¢', C.P., Klipheuwel,
ex Octococcus sp. on Compositae, T2783, x.1968
(Insley) 39, 20°, same data as holotype; 32, 2c,
C.P., Tulbagh district, ex mealybug on Rehlania
ericoides, T2833, x.1968 (Gilliomee); 692,50, C.P.,
Caledon, x.1969, ex Octococcus sp.n. on Oedera
imbricata, T3219, x.1969 (Insley); 32, 10%, C.P.,
Villiersdorp District, ex mealybug on Chrysocoma
?coma-aurea, T3217, x.1969 (Insley);12, 10%, C.P.,
Paarl, x.1971, H.P.Insley, ex mealybug on Cape
Macchia, T3971, x.1971 (Insley); 19, 4c, C.P.,
Worcester, ix.1972, ex mealybug on unknown plant,
T4525, ix.1972 (Neser); 3 2, C.P., Windmeul nr Paarl,
ex Octococcus sp. nt. pentziae on Athenasia trifurcata,
T4922, 3.x.1974 (Prinsloo) 29, 10°, C.P., Ceres,
9.x.1974, on Rehlania genistaefolia, T4927, 9.x.1974
(Prinsloo); 92, 60°, C.P., Tulbagh, ex ?Octococcus
sp. on ?Athenasia sp., T5469, vi, 1978 (Neser); 59,
90, OFS, Edenville, OFS, ex Octococcus pentziae on
Pentzia globosa, T3503, i.1970 (Insley); 29, 3c,
C.P., Oudtshoorn, ex Octococcus sp. on “Mesem’,
T6915, i1.1979 (Laubscher). Holotype in PPRI,
paratypes in PPRI and BMNH.

COMMENTS. Leptomastix herreni is most similar to
nigra, both species having the same variable colour
from largely orange to more or less completely dark
brown or blackish and the ocelli closer to the occipital
margin than to the eyes. They differ in the number of
lines of setae interrupting the linea calva and the
relative length of the postmarginal vein of the forewing.
In herreni there are at most three lines of setae inter-
rupting the linea calva (Fig. 17) and the postmarginal
vein is about as long as the stigmal (Fig. 18) whereas
in nigra there are generally at least four lines of setae
interrupting the linea calva (Fig. 34) and the
postmarginal vein is at least about 1.5x as long as the
stigmal (Fig. 35). The males can be separated as in the
females by the relative length of the postmarginal vein
(compare Figs 20, 38) and number of lines of setae
interrupting the linea calva and also by the number and
structure of the scale-like structures on the clava. In
herreni there are about 8-10 very fine, almost seta-
like, recurved structures (Fig. 61) whilst in nigra there
are only about 6 relatively broad, distinctly flattened
structures (Figs 66, 67). The species is also very
similar to Leptomastix nigrocoxalis and Leptomastix
digitariae (see under nigrocoxalis and digitariae for
comments). Paler forms of herreni might be mistaken
for flava but the females differ in having the forewings
completely hyaline, the mandibular bases almost black,
and only two or three lines of setae interrupting the
linea calva whereas in flava the forewings are fre-
quently distinctly infuscate, the mandibles are orange
and there are four or five lines of setae interrupting the
linea calva. The males on the other hand can be

J-M. ANGA AND J.S. NOYES

separated by the elongate, recurved scale-like struc-
tures on the clava whilst those in flava are very short
and broad.

The species is named in honour of Dr Hans Herren
for his contribution to biological control in Africa.

Leptomastix algirica Trjapitzin
(Fig. 21, 62)

Leptomastix algirica Trjapitzin 1989:126.
LECTOTYPE © (here designated), Algeria, NMP,
examined.

DIAGNOSIS. Female (length 1.40-1.50 mm): body
moderately robust; generally dark brown, including
coxae, femora and gaster; mouth margin yellowish;
tibiae and tarsi pale orange; wings hyaline; Fl about
2x as long as pedicel; posterior ocelli slightly closer to
occipital margin than eyes; forewing 2.5x as long as
broad; postmarginal vein variable, from slightly shorter
than stigmal to about one-third longer; stigmal vein
about as long as marginal vein; linea calva interrupted
by 2 to 4 lines of setae; gaster as long as thorax. Male
(length 0.90 mm): similar to female; antennae clothed
with long setae, longest about 3.3x as long as diameter
of F1; base of clava with 5 or so scale-like structures
ventrally; forewing venation as in Fig. 21.

VARIATION. Little in material examined, but see com-
ments.
Hosts. Unknown, but see comments.

DISTRIBUTION. Greece, Cyprus, Algeria

MATERIAL EXAMINED.

Type material. LECTOTYPE 9, here designated: Al-
geria, | 2, Aures: Ain Zaatout, |.—4.vi.1971 (Hoffer,
Horak [Expedition “SCARABAEUS’]) ‘Holotypus
Leptomastix algirica Hoffer 9’. Paralectotypes: Al-
geria, 19, Biskra, 7.-8.vi.1971 (Hoffer, Horak
[Expedition ‘SCARABAEUS’]); 1c, Auresi,
Ainzaatout, 1.—4.vi.1971 (Hoffer, Horak [Expedition
‘SCARABAEUS J. Lectotype in NMP, paralectotypes
in BMNH.

Non-type material. Greece, 1 9 , Kos, 4Km W Kos
Town, 20.viii.1994 (Noyes); Cyprus, 12, 13Km SW
Kyrenia, Karaman, 22—23.viii.1993 (Noyes). Material
in BMNH, both specimens part of longer series of
probably the same species.

COMMENTS. The name algirica was made available
in a review of the Palaearctic Encyrtidae by Trjapitzin
(1989) and accredited to Hoffer. However, in that
publication there is no indication that Hoffer was
responsible for meeting the criteria for making the
name available and therefore in compliance with Arti-
cle 50 of the International Code of Zoological
Nomenclature (1985) the name is here attributed to

REVISION OF LEPTOMASTIX

Trjapitzin. No reference was made to the primary type
by Trjapitzin and therefore a lectotype is selected
above.

We are of the opinion that algirica may be a dark
form of epona Walker since we are unable to find any
consistent morphological characters to separate the
two. We have examined a long series (BMNH) of
specimens reared in Israel from Pseudococcus cryptus
which have been identified as algirica. Females of this
series vary from entirely dark brown as in the type
material of algirica to the scutellum being almost
entirely orange. The males are inseparable from darker
forms of epona from western Europe. For the present
we are not including this material in the present study
because this ‘Israeli’ form is currently the subject of
experimental studies to ascertain its status.

Leptomastix africana Anga & Noyes sp. n.
(Figs 22-25)

DIAGNOSIS. Female (length 1.30—1.60 mm): body
including gaster dark-brown to black; all coxae dark
brown; Fl nearly 4x as long as pedicel (Fig. 22);
posterior ocelli closer to occipital margin than eye
margin; wings hyaline; forewing (Fig. 23) about 2.7x
as long as broad; postmarginal vein about twice as
long as stigmal vein which is subequal to marginal
(Fig. 25); linea calva interrupted by two lines of setae
(Fig. 24); gaster as long as thorax. Male (1.10—1.40
mm): generally similar to female but antennae clothed
with long setae, the longest about 5x as long as diam-
eter of Fl; basal half of clava with 9 or so scale-like
structures ventrally.

FEMALE (HOLOTYPE). Length 1.48 mm. Body in-
cluding gaster dark-brown to black; lower face with
conspicuous silvery setae; antennae dark-brown; meso-
scutum with constrasting silvery setae; scutellum
similar to mesoscutum but setae not as conspicuous;
wings hyaline; all coxae brown; femora brown but for
apex of mid femora which is yellow; fore and hind
tibiae and tarsi brown; mid tibia and tarsi pale orange;
propodeum laterally with silvery setae.

Head with inner eye margins diverging ventrad;
minimum frontovertex width slightly less than eye
length; posterior ocelli closer to occipital margin than
eye; antennae (Fig. 22) 1.3x as long as body, scape 5x
as long as broad, pedicel 1.25x as long as broad, Fl
nearly 4x as long as pedicel, clava 1.2x as long as F1
and 2x as long as F6. Relative measurements
(holotype): HW 43, HH 40, FV 22, POL 15, OCL 5,
OOL 8, EL 22, EW 7, MS 12, SL 25, SW S.

Thorax with finely reticulate sculpture on meso-
scutum and scutellum; forewing (Fig. 23) about 2.7x
as long as broad, with postmarginal vein twice as long
as either marginal or stigmal veins (Fig. 25); linea
calva interrupted by two lines of setae (Fig. 24);
hindwing hyaline, about 4.4x as long as broad.

103

Gaster as long as broad, with ovipositor hidden.
Relative lengths (paratype): OL 29, MT 65.

MALE. Generally similar to female but inner eye
margin subparallel, minimum frontovertex width about
equal to eye length, forewing about 2.6x as long as
broad, linea calva interrupted by a single line of setae,
antennae clothed with long setae, the longest about 5x
as long as diameter of F1; basal half of clava with 9 or
so scale-like structures ventrally.

VARIATION. Very little except in body colour and
size: smaller specimens tend to be brownish and larger
ones dark-brown to black.

Hosts. Leptomastix africana has been reared froma
mealybug, probably Octococcus sp., on Elytropappus
rhinocerotis (see below).

DISTRIBUTION. South Africa.

MATERIAL EXAMINED.

Type material. Holotype 9: South Africa, C.P.,
Stellenbosch, Idas Valley, ex ?Octococcus sp. on
Elytropappus rhinocerotis, T5976, 11.1979 (Neser).
Paratypes: South Africa, 32, lo’, same data as
holotype; 20°, C.P., Paarl, ex ?Octococcus on
Elytropappus rhinocerotis, T2737, ex 14.x.1968
(Insley); 72, 3o&°, Cape Province, Ceres, 111.1925
(Turner). Holotype in PPRI, paratypes in PPRI, BMNH
and IITA.

COMMENTS. Leptomastix africana is most similar to
abyssinica and nigrocincta (see comments under those
species).

Leptomastix nigrocoxalis Compere
(Figs 26-31)

Leptomastix nigrocoxalis Compere, 1928:219—220.
Holotype 2, South Africa, USNM, not examined.

Leptomastix phenacocci Compere, 1938:325. Holotype
Q , Egypt (ex. Java), BMNH, examined. syn. n.

[Leptomastix flavus Mercet; Risbec 1951:103.
Misidentification (part).]

Leptomastix brevis Hayat, Alam & Agarwal, 1975:14—
16. Holotype 9, India, BMNH, examined.
Synonymized with Leptomastix nigrocoxalis by
Noyes & Hayat (1994).

Leptomastix aligarhensis Khan & Shafee, 1975:194.
Holotype 9, India, ZAMU, not examined.
Synonymized with Leptomastix nigrocoxalis by
Noyes & Hayat (1994).

Leptomastix longiscapus Khan & Agarwal, 1976:378—
380. Lectotype 9 (designated Noyes & Hayat,
1994), India, ZAMU. Synonymized with
Leptomastix nigrocoxalis by Noyes & Hayat
(1994)).

DIAGNOSIS. Female (length 0.90—2.10 mm): body

104

including gaster orange to dusky orange; head with a
dark brown area adjacent to each eye which extends
across occiput towards foramen; mandibles always
with bases dark brown, almost black; mesoscutum
completely orange or with a brown wedge-shaped
mark medially; mesosternum always brown and mid
coxa always dark-brown; wings hyaline; position of
ocelli relative to eye and occipital margin variable,
generally about equidisant from eye and occipital
margin, but sometimes clearly a little closer to occipi-
tal margin; antenna (Fig. 26) with Fl about 1.3—2.0x
length of pedicel; forewing (Fig. 27) about 2.7x as
long as broad; postmarginal vein about 1.5x as long as
stigmal or slightly longer (Fig. 29); linea calva inter-
rupted by 4 or 5 lines of setae (Fig. 28); gaster as long
as thorax. Male (length 1.06—1.50 mm): generally
similar to female, but head and thorax predominantly
yellow; head with a dark-brown area on occiput adja-
cent to each eye, extending towards foramen;
mandibles varying from orange to dark brown; dor-
sum of thorax orange or with anterior part of
mesoscutum marked with dark brown; metanotum and
sides of propodeum yellow; side and sternum of tho-
rax mostly dusky-yellow; mid coxae occasionally
hardly darkened; legs pale orange except mid and hind
tibia which is brown dorsally; tarsi of fore and hind
legs brownish, those of midlegs dusky-yellow except
towards apex; funicle clothed in setae about 3x as long
as diameter of segments; antenna with 5 or 6 scale-like
structures on base of clava; forewing about 2.5—2.8x
as long as broad; linea calva interrupted by about four
lines of setae; aedeagus a little more than one-third as
long as mid tibia, each digitus with a pair of moderate
apical teeth.

VARIATION. The colour of the dorsum of the thorax
of both sexes varies as outlined above. In general the
frontovertex is not more than half head width with
posterior ocelli about equidistant from eye and occipi-
tal margin, OOL:OCL never greater than 1.4.

Hosts. Leptomastix nigrocoxalis has been recorded
as a parasitoid of Nipaecoccus graminis on
Hyparrhenia hirta (Prinsloo, 1981); ?Nipaecoccus
graminis on wild grass (Compere, 1928);
Pseudococcus sp. on Melia azedarach, ‘coccid’ on
hibiscus (Risbec, 1959); Planococcus citri on Citrus
medica (Agarwal, 1965) and on coffee (Prakasan and
Kumar, 1985); and Rastrococcus cappariae (Avasthi
and Shafee, 1983); Maconellicoccus (=Phenacoccus)
hirsutus (Compere, 1939c) and Nipaecoccus viridis
(Moursi, 1948; as Pseudococcus filamentosus
Cockerell, misidentification). It has also been recorded
by Hayat et al. (1975) from Coccidohystrix
(=Centrococcus) sp. on Achyranthes aspera and
Pupalia lappacea; Coccidohystrix (=Centrococcus)
insolita on Achyranthes aspera and Wittania somnifera;
Nipaecoccus sp. on citrus, Acacia, Peritropha

J-M. ANGA AND J.S. NOYES

bicalyculata and Morus alba; Nipaecoccus viridis on
Ziziphus, Acacia and Tephrosia purpurea, a ‘coccid’
on Tamarindus indica, Leucas cephalotus,
Hemigraphes and Acacia, and from the margarodid
Icerya aegyptica. Additionally recorded from
Rastrococcus iceryoides on Ziziphus and from
Pseudococcus sp. on tamarind (Noyes & Hayat, 1994).
The record of this species as a parasitoid of Nimboa
basipunctata (Neuroptera, Coniopterygidae) by
Sharma etal. (1985) is unlikely and requires confirma-
tion as does the record from Saissetia nigra (CIBC,
1970).

DISTRIBUTION. Egypt, Sudan, Zimbabwe, Namibia,
South Africa, India and Indonesia.

MATERIAL EXAMINED.

Type material. Holotype 9 of Leptomastix phenacocci:
‘Egypt sent by Priesner’ ‘ex Pseudococcus hirsutus’ ;
according to Priesner in Compere (1939c) this was
from material originally obtained in Indonesia (Java)
(BMNH); paratypes of Leptomastix phenacocci: 6 & ,
50°, same data as holotype (BMNH, UCR). Holotype
9 of Leptomastix brevis, India, Kerala, Kalamasseri,
ex Icerya aegyptica (Douglas), 12.11.1967 (Hayat)
(BMNH). Lectotype Qof Leptomastix longiscapus:
India, Uttar Pradesh, Garhwal, ex coccid on citrus,
5.ix.1975 (Khan) (ZAMU); Paralectotypes of
Leptomastix longiscapus: India, 99 , 40°, same data
as holotype (ZAMU).

Non-type material. Sudan, 1 9 , Wad Medani, from
mealybugs on Tamarindus indica, 17.v.1939
(Cowland); 1 9, Wad Medani, ex mealybug on Citrus
medica, ii.1938 (Ruttledge); Egypt, 5Q Giza,
7.1x.1929, ? hibiscus mealybug breeding room,
(Kamal); Senegal, 12 9 , 60°, Louga, ex cochenille sur
Euphorbia, 1945 (Risbec) (misidentified as
Leptomastix flava by Risbec); 12, Bambey, sur
Sourour (misidentified as Leptomastix flava by Risbec);
South Africa, 3 9 , 50’, Tvl, Dendron, ex, mealybug
on grass, T6067, i.1980 (Prinsloo); 1 9 , Tvl, Soutpan,
Pretoria Dist., 25°24'S 28°06'E, 11-12.1.1984
(Grobbelaar); 11 9,10, Tvl, Levubu Tvl, ex mealybug
on grass, T6924, 1.1987 (Prinsloo); 1 9 , Tvl, Entabeni
For. Res., 33°00'S 30°16'E, 7—1.1.1987 (Prinsloo);
192, Tvl, Weltevreden Farm nr Karino, ii.1990
(Prinsloo); India, material as listed in Noyes & Hayat
(1994). Material in BMNH, PPRI.

COMMENTS. Although there are slight differences in
the coloration of the head and mesoscutum of type and
other material identified as phenacocci and
nigrocoxalis (see Noyes & Hayat, 1994) we are unable
to find any significant morphological characters by
which to separate the two and therefore we treating the
two names as synonyomous.

Leptomastix nigrocoxalis is very close to nigra, the
species possibly being synonymous (see comments

REVISION OF LEPTOMASTIX

under nigra). It is also very similar to nigrocincta (see
comments under nigrocincta). At a glance, females of
nigrocoxalis may be mistaken for those of herreni but
can be separated readily on the relative length of the
postmarginal vein. In nigrocoxalis the postmarginal
vein is at least nearly twice as long as the stigmal (Fig.
29) whereas in herreni it is only about as long as the
stigmal vein (Fig. 18). Leptomastix nigrocoxalis might
be confused with epona and histrio, both of which are
Palaearctic species and yet to be recorded in Africa.
Females of nigrocoxalis can be distinguished from
both of these species by the blackish mandibular bases,
postmarginal vein at least about twice as long as the
stigmal, brown mid coxae and hyaline forewings. In
both epona and histrio the mandibular bases are al-
ways orange or orange-brown, the postmarginal vein
is at most only about 1.5x as long as the stigmal, the
mid coxae are orange, except in uniformly dark brown
specimens, and the forewings almost always have two
faint longitudinal fuscous streaks (similar to
dactylopii). The males are very similar to those of
dactylopii, nigrocincta, and nigra, but can be can be
separated with difficulty using the characters given in
the key. It is possible that these characters are unreli-
able.

Leptomastix nigra Compere
(Figs 32-38, 66-67)

Leptomastix nigrum Compere 1938:324—5.
Holotype 2 , South Africa, examined, BMNH.

DIAGNOSIS. Female (length 0.50—2.10 mm): head
and thorax varying from orange with frontovertex and
axillae dark brown, to almost completely dark brown,
blackish; paler specimens always with a dark-brown
area adjacent to each eye, extending across occiput,
towards foramen, mandibles always with bases dark
brown, almost black; mesosternum always brown and
mid coxa always dark-brown; wings hyaline; gaster
mostly dark brown, occasionally mixed with orange
basally; posterior ocelli clearly closer to occipital
margin than to eye; antenna (Figs 32, 33) with Fl
about as long as pedicel to about twice as long; forewing
(Fig. 34) about 2.7x as long as broad; postmarginal
vein about 1.5x as long as stigmal or slightly longer
(Fig. 35); linea calva interrupted by about 4 or 5 lines
of setae, 3 lines in the smallest specimens; gaster as
long as thorax; ovipositor as in Fig. 36. Male (length
0.50-1.50 mm): generally similar to female, but head
and thorax always largely yellow; head with a dark-
brown area on occiput adjacent to each eye which
extends towards the foramen; median area of
frontovertex brown; mandibles varying from orange
to dark brown; mesoscutum orange laterally and brown
to dark-brown medially; axillae dark-brown; scutellum
orange to dark brown; metanotum and sides of

105

propodeum yellow; side and sternum of thorax mostly
dusky-yellow; mid coxae occasionally hardly dark-
ened; legs pale orange except mid and hind tibia which
are brown dorsally; tarsi of fore and hind legs brown-
ish, those of midlegs dusky-yellow except towards
apex; funicle (Fig. 66, 67) clothed in setae about 3x as
long as diameter of segments; five or six scale-like
structures on base of clava; forewing (Fig. 37) about
2.5—2.8x as long as broad; linea calva interrupted by
about four lines of setae (Fig. 38); aedeagus a little
more than one-third as long as mid tibia, each digitus
with a pair of moderate apical teeth.

VARIATION. As mentioned in the diagnosis for the
females there is a notable amount of variation in the
colaration of the body, some specimens being largely
yellow or pale orange, but almost without exception
with the axillae dark brown whilst in others the head
and thorax is more or less completely dark brown. We
have also noted variation in the relative lengths of the
antennal segments with the smallest specimens having
F1 only about as long as the pedicel (Fig. 33) whilst in
larger specimens it may be about twice as long (Fig.
32). On the other hand, apart from a slight variation in
the relative lengths of the funicle segments (Figs 66,
67), males appear to vary little. Some males have the
brown areas of the dorsum of the thorax more exten-
sive than others with sometimes the whole of the
scutellum almost completely dark brown.

Hosts. Leptomastix nigra was recorded as a
parasitoid of Pseudococcus sp. on oleander by Com-
pere (1938). The host is very probably Paracoccus
burnerae, as this mealybug on Nerium oleander has
been recorded as the host for Leptomastix nigra by
Prinsloo (1981). In addition, the species is recorded
below from Octococcus africanus on Stoebe cinerea;
Octococcus ?minor on Athenasia trifurcata;
Octococcus pentziae on Metalasia muricata and
Pentzia globosa; Octococcus on Elytropappus;
2Octococcus on Stoebe and Elytropappus
gnaphaloides; Paracoccus burnerae on ?Diospyros
lycioides; Spilococcus on Metalasia muricata and an
unidentified mealybug on Gazania. Also recorded
below in association with Parasaissetia nigra
(Homoptera: Coccidae) on Loranthus elegans,
Saissetia somereni (Coccidae) on grapefruit, and
Ceroplastes sp. (Coccidae) on Metalasia muricata,
but these are unlikely to be the true hosts.

DISTRIBUTION. Zimbabwe, Namibia, South Africa.

MATERIAL EXAMINED.

Type material. Holotype 2 of Leptomastix nigra, South
Africa, Tvl, Johannesburg, May 1937 (Compere)
(BMNH). Paratypes: South Africa, 19, 20°, Tvl,
Johannesburg, ex mealybug, iv.1937 (Compere).
Other material. Zimbabwe, 59, Salisbury, near Harare
and Chishawasha, various dates 111.1979—vi1.1982

106

(Watsham); Namibia, 1, Helmeringhausen, with
Parasaissetia nigra on Loranthus elegans, T6925,
y.1973 (Prinsloo); 29, 20°, Otavi, ex mealybug on
unidentified plant, T6928, 17.v.1973 (Prinsloo); 4°,
Karibib, ex mealybug, T6931, 16.11.1978 (Kok); 62,
1c, Otavi, ex mealybug, T6930, 24.11.1978 (Kok);
South Africa, 6 9 and 9c’, OFS, Edenville, OFS, ex
Octococcus pentziae on Pentzia globosa, T2503,
i.1970 (Insley); 179, C.P., Jeffreys Bay, ex mealybug
in axils of Gazania rigens, T6916, 3.1.1980 (Neser);
190, C.P., Jeffreys Bay, ex mealybug in axils of
Gazania, T6917, 3.i1.1980 (Neser); 22, 70, Tvl, S.
of Pongola, 27°23'S 30°38'E, ex Saissetia somereni
on grapefruit, T6932, vii.1979 (Kambourov); 1 9 ,C.P.,
Witzenberg Vall., 3, OOOft, 19.1.1921 (Turner); 12,
1o, C.P, Paarl District, ex ?Octococcus on Stoebe,
T2736, x.1968 (Insley); 29,20, C.P., Du Toits Kloof
Pass, ex ?Octococcus sp. on Elytropappus
gnaphaloides, T2795, x.1968 (Insley); 42, 1c, Du
Toits Kloof Pass, ex mealybug on Metalasia muricata,
T2832, x.1968 (Insley); 22,20, C.P., Villiersdoorp,
ex. mealybug on Stoebe sp., T3205, x.1969 (Insley);
82, 90°, C.P., Swellendam, T3226, ex Spilococcus
sp. on Metalasia muricata, x.1969 (Insley); 3 2, C.P.,
Stellenbosch, ex Octococcus ?minor on Athenasia
trifurcata, T3231, x.1969 (Insley); 89, 150°, C.P.,
Jeffreys Bay C.P., ex Octococcus sp. on Metalasia
muricata, T3378, iii.1970 (Insley); 32, C.P., Garcia
Pass, ex Octococcus sp., T3414, iii.1970 UInsley); 19,
4c, C.P., Stellenbosch, on Stoebe plumosa, T5383,
v.1978 (Neser); 42, 3c, C.P., Kleimond, ex
Octococcus africanus on Stoebe cinerea, T5475,
viii.1978, (Neser); 22,40, C.P., Floraberg, ex prob.
Octococcus sp. on Metalasia sp., T5793, vi.1978
(Neser); 72, 10, C.P., Franschhoek Pass, ex
Octococcus pentziae on Metalasia muricata, T5883,
xii.1978 (Neser); 22, 150°, C.P., D.F Malan Airport
C.P., ex Octococcus prb. pentziae on Metalasia
muricata, T6114, iv.1979 (Neser); 3c’, C.P., D.F.
Malan Airport, with Ceroplastes sp. on Metalasia
muricata, 10.iv.1979 (Neser); 42, 1c", C.P. Goudini,
ex. prob Octococcus sp., on Stoebe sp., T6914 (Neser);
52,1c, C.P., Goudini, ex prob. Octococcus sp. on
Elytropappus rhinocerotis, T6920, 25.1x.1979 (Neser);
50’, CP, Algeria, Cadarberg, ex prob. Octococcus sp.
on Elytropappus sp. cf. gnaphaloides, T6921
16.vii.1979 (Neser); 42, 10°, C.P., Sir Lowry’s Pass
rd. nr. Somerset, ex Octococcus sp. on Athenasia sp.,
T6922, 29.viii.1979 (Neser); 60°, Tvl, Irene, ex
Paracoccus burnerae on ?Diospyros lycioides, T6927,
16.iii1.1969 (Insley); 1492, 150°, C.P., Capetown, D.F
Malan Airport, ex Octococcus prb. pentziae on
Metalasia muricata, T6230, vii.1980 (Neser); 29,
1o', C.P., Gamtoos Sta., with Ceroplastes sp. on
Metalasia muricata, T6933, 22.xi.1983 (Grobbelaar).
Material in BMNH, PPRI.

J-M. ANGA AND J.S. NOYES

COMMENTS. Leptomastix nigra is extremely close to
nigrocoxalis and may represent different forms of that
species. In general, the two can be separated by the
relative width of the frontovertex and position of the
ocelli in the females, with some degree of overlap
between unusually small specimens of nigrocoxalis
from India and specimens identified here as nigra. In
nigra the frontovertex is almost always half as wide as
the head or wider and OOL:OCL is at least 1.4, whilst
in nigrocoxalis the frontovertex is less than half the
head width except in the smallest specimens and
OOL:OCL is less than 1.4. Other differences, such as
coloration, may be unreliable, but in nigra the axillae
and frontovertex are nearly always dark brown and in
nigrocoxalis at most only the occipital margin is marked
with brown and the axillae are always orange. The
males can be separated only by the coloration of the
frontovertex and axillae (see key), but this is probably
unreliable.

Darker females of Leptomastix nigra are similar to
Leptomastix abyssinica, both species having a general
dark brown coloration and forewing about 2.7x as
long as broad. They can be separated by the colour of
the clypeal area and prosternum, and the number of
lines interrupting the linea calva. In nigra the clypeal
area and prosternum are dark brown and there are four
or five lines of setae interrupting the linea calva (Fig.
34), whereas in abyssinica the clypeal area and
prosternum are yellow and there are only two lines of
setae interrupting the linea calva (Fig. 58). Paler fe-
males of nigra, those which are generally orange but
have dark axillae, can be mistaken for flava, a species
known from Mediterranean Europe and the Middle
East. However, in flava the forewing has the
postmarginal vein about as long as the stigmal vein
and the mandibles are orange, whereas in nigra the
postmarginal vein is at least 1.5x as long as the stigmal
and the mandibles are dark brown. Leptomastix nigra
can also be very similar to herreni, both species exhib-
iting the same range of colour variation (see comments
under herrent)

Leptomastix dactylopii Howard
(Figs 39-46, 63, 70)

Leptomastix dactylopii Howard, 1885:23-24.
Lectotype 9 (designated Noyes & Hayat, 1994),
USA, USNM examined.

Leptomastix superba Silvestri, 1915:348. Holotype @ ,
Eritrea, INMP examined. Syn.n.

Leptomastix longipennis Mercet, 1927:497—498.
Lectotype 9 (designated Noyes, 1981), Eritrea,
IEEM, examined. Synonymized with Leptomastix
dactylopii by Noyes & Hayat (1994).

Leptomastix bifasciatus Compere, 1938:323-325.
Holotype 2 , Tanzania, examined. Synonymized by
Noyes & Hayat (1994).

REVISION OF LEPTOMASTIX

Paraleptomastix dactylopii (Howard); Mani, 1939:70-
Til

Leptomastix tambourissae Risbec, 1952:16-19.
Lectotype 9 (designated Noyes & Hayat, 1994),
Madagascar, MNHN, examined. Synonymized by
Noyes & Hayat (1994).

[Leptomastix flavus Mercet; Risbec, 1951:103;
Meyerdirk & Newell, 1979:72. Misidentification. ]

DIAGNOSIS. Female (length 1.10—2.30 mm): body
including gaster yellow to orange, moderately robust
and stout; head with dark-brown area adjacent to each
eye, extending across occiput towards foramen;
mesopleuron yellow, concolorous with mid coxa; mes-
osternum dark-brown; wings more or less hyaline
(Fig. 40), sometimes with two distinct fuscous longi-
tudinal streaks (Fig. 41); position of posterior ocelli
relative to eyes and occiput variable; Fl about 2x
length of pedicel; forewing fairly wide, about 3x as
long as broad (Figs 40, 41); postmarginal vein at least
1.5x as long as stigmal (Fig. 43); linea calva inter-
rupted by at most two lines of setae (Fig. 42); gaster as
long as thorax; ovipositor about 12x as long as
gonostyli. Male (length 0.90-1.54 mm): generally
similar to female, but posterior margin and dorsal part
of pronotum and anterior margin of mesoscutum
brownish; antenna (Fig. 63) clothed with long setae,
each at least 4x as long as diameter of F1; basal two-
thirds of clava with 9 or so scale-like structures present
ventrally; aedeagus about half as long as midtibia;
phallobase (Fig. 70) with elongate digiti, each with a
pair of apical hooks.

Hosts. Leptomastix dactylopii has been reared from
many species of mealybugs. Ashmead (1900) and
later Viereck (1916) first recorded Planococcus citri
(as Dactylopius destructor) as its host whilst Mercet
(1927) noted that the species (as L. longipennis) was a
parasitoid of Ferrisia virgata and Compere (1938)
recorded it from Pseudococcus longispinus as
Leptomastix bifasciatus. Leptomastix dactylopii has
also been recorded from Birendracoccus saccharifolii
on sugarcane by Mani (1939), but this is a possible
misidentification of Leptomastix tsukumiensis. Kerrich
(1953) noted it as a parasitoid of Planococcus vovae
(as Pseudococcus inamabilis) while Annecke & Insley
(1971) recorded it from Planococcus aemulor on
Combretum splendens. Leptomastix dactylopii has also
been recorded from stem galls on Tambourissa sp. by
Risbec (1952), from Pseudococcus sp. on guava by
Compere (1938), Pseudococcus bukbensis,
Planococcus citri, Ferrisia virgata, Phenacoccus
madeirensis, Pseudococcus longispinus (= adonidum)
and Pseudococcus concavocerarii by Donald (1956),
Planococcus kenyae on coffee (Noyes & Hayat, 1994),
Pseudococcus occiduus on Psidium guajava by
Annecke & Insley (1971), Planococcoides njalensis
by Strickland (1951), Planococcoides lamabokensis

107

by Prinsloo (1983a), Planococcus kraunhiae in Japan
by Tachikawa (1963a) and Dysmicoccus brevipes in
Hawaii by Bartlett (1978). The parasitoid has been
laboratory reared from the last host in Ghana by
Anonymous (1953). Tachikawa (1963b) recorded
Leptomastix dactylopii from Phenacoccus gossypii
but this was based on a record of specimens reared
from a mixed population of Phenacoccus gossypii and
“Pseudococcus krauhniae’, the latter being a
misidentification of Planococcus citri and therefore
the most likely host. Leptomastix dactylopii has been
reared from several mealybug species in the labora-
tory: Phenacoccus gossypii, Phenacoccus solani,
Pseudococcus calceolariae, Pseudococcus longispinus
and Pseudococcus maritimus by Bess (1939),
Pseudococcus comstocki by Clancy (1944) and
Planococcus lilacinus by Krishnamoorthy (1988).
Manichote & Middlekauff (1967) noted that
Leptomastix dactylopii would not attack Spilococcus
leucopogi (= cactearum ) in the laboratory. Recorded
below from Delotococcus (=Allococcus) quaesitus on
Acacia and Planococcus ficus on Ficus carica and an
unidentified species of Planococcus on Erythrina
lysistemon.

DISTRIBUTION. Senegal, Gambia, Sierra Leone,
Ivory-Coast, Togo, Benin, Nigeria, Cameroon, Ga-
bon, Uganda, Zimbabwe, Zambia, Kenya, Seychelles,
South Africa, Mauritius, Canada (introduced), USA
(introduced), Hawaii (introduced), Caribbean, Costa
Rica, Chile, Brazil, Mediterranean Europe (intro-
duced), Pakistan (introduced), India (introduced),
Taiwan (introduced) and Australia (introduced).

MATERIAL EXAMINED.

Type material. Leptomastix dactylopii: lectotype 9 on
card point labelled ‘11700 Pars on Dactylopius de-
structor Oct 4.[18]84’ “Type 2649 U.S.N.M. (USNM)’.
Paralectotypes 10’, on single point, labelled ‘11700,
Oct.4.1884 “Type No 2649 U.S.N.M.’; 19, same
data but 14.10[18]84 (USNM). Leptomastix superba:
holotype 9 fragmented on a slide, labelled
‘Leptomastix superbus Silv. F Nefasit’ (INMP).
Leptomastix longipennis: lectotype 9 on card labelled
‘ex Pseudococcus virgata’ ‘Somalia Italiana’
“Leptomastix longipennis Mercet’ ‘Leptomastix
longipennis Mercet Lectotype det. J.S. Noyes, 1979’
(IIEM). Leptomastix bifasciata: holotype 2, South
Africa, Uitenhage, ex Pseudococcus longispinus on
oleander, 12.1.1937 (Compere) (BMNH). Paratypes:
Tanzania (as Tanaganyika), 39, Bukoba, ex
Pseudococcus on guava, 20.vii.1935 (Ritchie)
(BMNH). Leptomastix tambourissae: lectotype @ , la-
belled “Elevage du 1.4.50 Eclos 20.6.50 No G20P
galles de tege des Tambourissa sp. d’ Ambatoloana
(R.P.)’ ‘Encyrtidae Leptomastix tambourissae Risbec
T’ (MNHN). Paralectotypes: 32, 1c (not four as
stated by Risbec, 1952) remounted from same slide

108

onto card rectangles as lectotype (MNHN, BMNH).

Non-type material. No data, 1 9 (misidentified as L.
flava by Risbec); Senegal, 19, 39, Bambey, ex
coccid, (Risbec); 29, Bambey, ex Pseudococcus on
Prosopis, 12.x.1945 (Risbec) (misidentified as L. flava
by Risbec); Gambia, 1’, Bakan, i.78 (Huggert); 20’,
Fajara, i.78 (Huggert); Sierra Leone, 10’, Njala, ex
mealybug on Cassiea javanica; Ivory Coast, 7°,
1c, Lamto, 6°13'N 5°02'W, MT/YPT, x1.1988 (Noyes);
20, Korhogo, 18—21.iii.1984 (M. Matthews); 19,
30°, Gagnoa, 2—5.i11.1984 (Matthews); 192, 30,
Bouaffle; R.P. Benin, 52, 7c’, 15Km N. Cotonou,
Abomey-Calavi, MT, xii.1988 (Noyes); Togo: 29,
Kpalime, xii.1988 (Noyes); Nigeria, 3 9 , 1c’, Kaduna
St. 20km N., 8.xi.1987, (Noyes, Neuenschwander),
52,50, Ibadan, ITA compound, MT/YPT, x1.1987
(J.S.Noyes); 60’, Zaria, 8.x1.1987 (J.S. Noyes); Cam-
eroun, 22, 20°, S.W. Cameroun, Musone, Lake
Mboandong, 11.xii.1981 (Compton); 2 2, 3.0, Victo-
ria Bot. Gdns, 6.xii.1981, (Compton); 1 9 , Nyassosso,
Mt Koupe, 8.i.1982 (Compton); Gabon, 29, 20’,
Foret de la Mondah, 15—25km N.Libreville, 25.xi—
3.xii.87 (Noyes); Uganda, 1 9, Bujumbura, iv.1939
(Taylor); 32, 4c, ex Pseudococcus kenyae on C.
arabica, 12.iv.1938 (Melville); Zimbabwe, 62 , 3c,
Salisbury, i.1979 (Watsham); 62, 20, Harare,
Chishawasha, xii. 1982 (Watsham); Zambia, 9 9 , 130",
15km E.Lusaka, 27.xi-29.xii.1979 and 11—19.11.1980
(Beaver); Kenya, 19, Nairobi, 11.1982, Brit. Mus.
1982-347; 30%, Gazi, vili.1982 (Barnett); South Af-
rica, 18 9,290", Cape Province, 10—22.xii.1930, Brit.
Mus. (Zurner); 19, Mossel Bay, April.1921, 1921—
210, (Turner); 39, Blyderivierspoortdam Nat.
Reserve, 29°32'5, 30°47'E, 25—26.x.1984, (Prinsloo);
1 9, Cape Province, Grahamstown, 1.1979 (Prinsloo);
19, 1o& Letaba, ex suction trap on citrus, T2504,
xi. 1966 (Catling); 4 9 , 1c" Cape Province, Ashton, ex
mealybug on Ficus carica, T5306, 111.1978 (Neser,
Kok & Urban); 19, Okahandja, 2—18.i11.1928; 19,
Transvaal, Roodeplaat, nr Pretoria, ex Allococcus
quaesitus on Acacia sp., T4292, iii. 1992, (Insley); 1%,
Mogol Nature Reserve, Ellisras Dist. 23°58'S 27°45'E,
27-29.ii1.1984 (Prinsloo); 1c, Entabeni Forest Res.
Soutlanberg, 25°00'S 30°16'E, 3—7.xi.1980 (Prinsloo);
1o,C.P., Jeffreys Bay, with unidentified coccoidea on
Euclea sp., T6918, 13.1.1980 (Neser); 1co, Natal,
Umzinto, ex Planococcus sp. on Erythrina lysistemon,
T4135 1.1972 (Insley); 29, C.P., De Dooms, ex
Planococcus ficus on Ficus carica, T6919; Mauri-
tius, 20’, Rose Hill, 29.ix.32 (Mamet). Material in
BMNH, PPRI, MNHN.

Voucher material. Israel, 19, California, River-
side, lab culture, ex Planococcus citri, 30.viii.1974,
R74-81-1, shipped, reared and released in California,
Porterville by Meyerdirk as Leptomastix flavus; 19,
lo, Negev, lab reared, California, Porterville, ex
comstock mealybug, Warkentin, R76—42, 17.v.1976,

J-M. ANGA AND J.S. NOYES

shipped and released in California, Porterville, by
Meyerdirk as Leptomastix flavus; 1c, Sde Boker, lab
culture, California, Porterville, ex Planococcus vitis
on grape, R-75—34—2B, 30.iv.1975, shipped, reared
and released in California, Porterville by Meyerdirk as
Leptomastix flavus. Material in UCR and part of a
longer series of voucher material of the species re-
leased in California from Israel and referred to by
Meyerdirk & Newell (1979) as Leptomastix flava.

COMMENTS. The holotype of Leptomastix superba
is mounted on a slide and in very poor condition, but it
is clearly the same as Leptomastix dactylopii and
therefore we have no hesitation in treating the two
names as synonymous.

Females of Leptomastix dactylopii are similar in
general appearance to Leptomastix tsukumiensis, both
species having similar habitus and generally yellow
coloration of body and mid coxae (see comments
under tsukumiensis). The males are very similar to
those of nigrocoxalis, nigrocincta, and nigra and can
be difficult to separate. Males of these four species can
be separated using the characters given in the key,
although it is likely that these characters are not totally
reliable.

Leptomastix jonesi Noyes sp. n.
(Figs 47-48, 64)

DIAGNOSIS. Female (length 1.58—2.24 mm): body
mostly yellow, but with temples part of occiput, mes-
osternum, mid coxae externally and occasionally
mesopleuron brown; gaster orange brown; posterior
ocelli about equidistant from eye and occipital mar-
gins; antenna with Fl about 3x as long as pedicel;
forewing about 3x as long as broad, with a longitudi-
nal fuscous streak along anterior wing margin from
apex of venation and another parallel streak in disc
(similar to Fig. 47); postmarginal vein about 2x as
long as stigmal and slightly longer than marginal;
linea calva interrupted by 4 or 5 lines of setae. Male
(length 1.77—1.83 mm): generally similar to female,
antenna (Fig. 64) clothed with long setae, each at least
3x as long as diameter of F1; basal two-thirds of clava
with 9 or 10 slender, elongate and straight scale-like
structures ventrally; forewings (Fig. 47) with two
fuscous longitudinal streaks as in female; postmarginal
vein about 1.5x as long as stigmal (Fig. 48); aedeagus
less than one-third as long as midtibia; phallobase with
elongate digiti, each with three apical hooks.

FEMALE (HOLOTYPE). Length 2.21 mm. Head or-
ange with a small brown spot on interantennal
prominence dorsally, temples and occiput laterally
dark brown; radicle dark brown; scape brown with
ventral margin yellow; pedicel brown; flagellum dark
brown; dorsum of thorax mostly orange with neck of
pronotum and extreme anterior part of mesoscutum

_

REVISION OF LEPTOMASTIX

brown; mesosternum and metanotum dark brown, re-
mainder of thorax orange; legs orange except for mid
coxae which are dark brown laterally; gaster orange-
brown dorsally, orange laterally and ventrally.

Head with inner eye margins subparallel, diverging
slightly ventrally; posterior ocelli about equidistant
from eye and occipital margins; scape about 5.5x as
long as broad; Fl 3x as long as pedicel; clava very
slightly shorter than F1 and 1.6x as long as F6; pedicel
about 2x as long as broad. Relative measurements:
HW 50.5; HH 44; FV 21; SL 33.5; SW 6.

Mesoscutum and scutellum with coriaceous sculp-
ture; forewings 3x as long as broad, with postmarginal
vein very nearly 2x as long as stigmal and a little
longer than marginal; linea calva interrupted by 4 or 5
lines of setae; hindwing 4.5x as long as broad.

Gaster slightly longer than thorax, with ovipositor
hidden. Relative measurements: OL 38.5, GL5.5 [MT
84].

MALE. Seediagnosis. Relative measurements (speci-
men 1): HW 93, FV 48, POL 17, OOL 10.5, OCL 7,
EL 53, EW 35, MS 23, SL 52, SW 11.5; (specimen 2):
FWL 148, FWW, 54.5, AL 22, MT 80.

VARIATION. Very little, but the female excluded from
the type series has the mesopleuron brown instead of
orange.

Hosts. Unknown.

DISTRIBUTION. South Africa.

MATERIAL EXAMINED.
Type material. Holotype 9 (slide-mounted), South Af-
rica, Cape Province, Mossel Bay, xii.1921 (Turner).
Paratypes: 10°, same data as holotype; 19, Cape
Province, Mossel Bay, 18—30.xi.1921 (Turner); 10,
Cape Province, Mossel Bay, vi—vii.1930 (Turner);
392, Cape Province, Somerset East, 1—26.i1.1931
(Turner). Holotype in BMNH, paratypes in BMNH
and PPRI.

Non type material. South Africa, 1 9 , Pondoland,
Port St John, ix.1923 (Turner)

COMMENTS. Leptomastix jonesi is superficially very
similar to dactylopii and females of the two species
can be difficult to separate with certainty. In jonesi
the mid coxae are partially brown, the linea calva is
interrupted by at least 4 lines of setae, the ovipositor
is less than half as long as the mid tibia and only about
7x as long as the gonostyli whereas in dactylopii the
mid coxae are completely orange or yellow, the linea
calva is generally interrupted by only 2 lines of setae,
the ovipositor is at least about 10x as long as the
gonostyli and more than half as long as the mid tibia.
The males of jonesi are fairly easy to separate from
those of dactylopii and other species of the genus by
the scale-like structures at the base of the clava being
very elongate, slender and straight (Fig. 64). In the

109

only other species where these structures are slender
(herreni) they are relatively short and apically strongly
curved (Fig. 61).

The species is named in honour of Mr Peter Jones
for his contribution to children’s education.

Leptomastix nigrocincta Risbec
(Figs 49-55)

Leptomastix nigrocincta Risbec, 1959:27. Lectotype ° ,
designated by Noyes & Prinsloo (1988), Madagas-
car, MNHN, examined.

Leptomastix lyciae Noyes & Hayat, 1994:275-278.
Holotype 9 , India, BMNH, examined. Syn.n.

DIAGNOSIS. Female (length 0.91—2.20 mm): body
including base and venter of gaster yellow to orange,
dorsum of gaster at least partially dark brown; head
always with a dark-brown area on temple adjacent to
each eye extending a short way onto occiput towards
foramen; base of mandibles yellow; mesopleuron gen-
erally orange, rarely dark brown (see below);
prosternum and mesosternum adjacent to coxae usu-
ally with a distinct brown tinge; mid coxae always
dark-brown or black; propodeum dark-brown above
coxae; wings hyaline or with a pair of longitudinal
fuscous streaks(Fig. 51); position of posterior ocelli
variable (see below), but usually ocelli about equidis-
tant from eye and occipital margin; antenna (Figs 49,
50) with Fl about 1.7 to 2.3x as long as pedicel;
forewing (Fig. 51) about 3x as long as broad;
postmarginal vein slightly longer than marginal and at
least nearly twice as long as stigmal (Fig. 53); linea
calva interrupted by two or three lines of setae, rarely
by four (Fig. 52); gaster as long as thorax; ovipositor
about 6.5x as long as gonostyli. Male (length 0.80—
1.59 mm): similar to female but darker, although mid
coxae not or hardly dusky; funicle clothed in long
setae, longest about 4x as long as diameter of F1; a line
of six or so scale-like structures ventrally on base of
clava; forewing (Fig. 54) about 2.5x as long as broad;
postmarginal vein distinctly longer than marginal which
is subequal in length to stigmal (Fig. 55); aedeagus
about one-third length of mid tibia.

VARIATION. Very little variation has been noted in
material from India, China and the African mainland,
but specimens from Madagascar exhibit a striking
degree of variation in coloration and some minor
morphological characters. In Malagasy females the
area between the toruli and eyes is sometimes marked
with black as is the median area of the pronotum; the
mesoscutum may have a wedge-shaped dark brown
area; the tegulae vary from almost completely orange
to completely dark brown or black; the axillae may be
dark brown laterally; the mesopleuron is sometimes
almost entirely black. In the Malagasy material the

110

frontovertex varies from slightly less than two-fifths
head width to nearly half with a corresponding varia-
tion in the relative position of the posterior ocelli. In
some specimens the posterior ocelli are distinctly
nearer the occipital margin whilst in others they are
distinctly nearer the eyes with the angle formed vary-
ing from about 50° to only slightly less than 90°. In the
Malagasy material the relative width of the funicle
segments varies with some specimens having slender
antennae with F6 about 2.5x as long as broad and
others having slightly broader segments with F6
slightly less than 1.5x as long as broad (compare Figs
49, 50); the postmarginal vein varies from about 2.0—
2.5x as long as the postmarginal vein. Males from
Madagascar exhibit a similar, but less marked range of
variation.

Hosts. Leptomastix nigrocincta was recorded as a
parasitoid of mealybugs on Hibiscus sp. by Risbec
(1959). Recorded in India from Coccidohystrix insolita
and Phenacoccus sp. (probably misidentification of C.
insolita) on aubergine, Solanum melongena L. (Noyes
& Hayat, 1994).

DISTRIBUTION.
China.

Togo, Zambia, Madagascar, India,

MATERIAL EXAMINED.

Type material. Lectotype 9 of nigrocincta: ‘Elevage
du 5.12.50 No 810 Eclose du 20.12.50 Parasites des
cochenilles sur le feuille d’ Hibiscus sp. de Tsimbazaza
(R.P.) Inst. Scient. Madagascar’ “Leptomastix
nigrocincta’, remounted on a card rectangle and la-
belled ‘LTQ’ on ventral side of card (MNHN).
Paralectotypes: 69, 32, remounted on card rectan-
gles, same data as lectotype. Lectotype of nigrocincta
in MNHN, paralectotypes in MNHN and BMNH.
Holotype 9 of lyciae: India, Delhi, [ARI area, x.1979
(Boucek). Paratypes: 53 9 , 13% from India and China
as detailed in Noyes & Hayat (1994). Holotype of
lyciae in BMNH, paratypes in BMNH, USNM, CNC,
AMU, PPRI, QM.

Non-type material. Togo, 10 2 , 1Okm NW Kpalime,
xii.1988 (Noyes); Zambia, | 9 , Lusaka, 6—12.xi.1979
(Beaver); Madagascar, | 0’, Tananarive, Angavokely
for sta., 28km E. Angavokely, 24.iv.1983 (Noyes) 2 9 ,
20, Perinet, 27.iv.—3.v. 1983 (Noyes, Day); 519,
700, Tulear, Berenty 12km, NW Amboasary, 5—
15.v.1983 (Noyes, Day); 32, 110°, Tulear, Bereboka
60km, N.E. Morondova, 18—23.v.1983 (Noyes, Day);
19, 20km S. Mandrivazo, 24.v.1983 (Noyes, Day).
Material in BMNH, PPRI.

COMMENTS. As we are unable to find any consistent
differences to discriminate between the type series of
lyciae and the long series of nigrocincta from Mada-
gascar, we are here treating the two species as
synonymous.

Females of Leptomastix nigrocincta can be very

J-M. ANGA AND J.S. NOYES

similar to nigrocoxalis and might be mistaken for that
species. In general, both species have similar colora-
tion with the body generally yellowish with the mid
coxae partially dark brown. The two species can be
separated by the coloration of the base of the mandi-
bles and number of lines of setae interrupting the linea
calva. In nigrocincta the mandibles are entirely yel-
lowish and the linea calva is interrupted by at most
three lines of setae (very rarely four), whilst in
nigrocoxalis the bases of the mandibles are dark brown
and the linea calva is interrupted by four or more lines
of setae. In general, males are very similar to those of
nigrocoxalis, nigrocincta, and nigra and can be diffi-
cult to separate. They can be separated using the
characters given in the key, although it is likely that
these characters are not totally reliable. Females of
Leptomastix nigrocincta can also be similar to those of
Leptomastix africana, both species having the poste-
rior ocelli closer to eyes than occiput, F1 at least 2x as
long as pedicel; mesopleuron brown, postmarginal
vein distinctly longer than stigmal, and only two lines
of setae interrupting the linea calva. They differ in
general coloration, and relative length of postmarginal
vein. In nigrocincta the body is largely pale orange
and the postmarginal vein is about 2.6x as long as the
marginal vein and about 3x as long as the stigmal vein,
whereas in africana the body is entirely dark brown
and the postmarginal vein is twice as long as either the
marginal or stigmal veins. At a glance, females of
some colour forms of nigrocincta might be confused
with those of herreni, but the two species can be
separated easily on the relative length of the
postmarginal vein. In nigrocincta the postmarginal
vein is at least nearly twice as long as the stigmal (Fig.
53) whilst in herreni it is about as long as the stigmal
(Fig. 18). Leptomastix nigrocincta might also be mis-
taken for histrio or epona, both of which are Palaearctic
and currently unknown in the Afrotropical region.
Females of nigrocincta differ from those of the two
other named species in always having the postmarginal
vein at least nearly twice as long as the stigmal, the
mid coxae dark brown and the linea calva generally
interrupted by not more than three lines of setae,
whereas in histrio and epona the postmarginal vein is
only about 1.5x as long as the stigmal and the mid
coxae are yellow, except in the darkest specimens
where the linea clava is interrupted by at least five
lines of setae.

Leptomastix abyssinica Compere

(Figs 56-60)

Leptomastix abyssinica Compere, 1931:267-9.
Holotype @ , Eritrea, not examined, USNM.

DIAGNOSIS. Female (length 1.60—2.40 mm): body
including gaster generally brown to dark-brown; head

REVISION OF LEPTOMASTIX

with inner eye margins, genal area and area below
toruli yellow; antennae dark brown with ventral mar-
gin of scape yellow or translucent; prosternum and
fore coxae yellow; mid and hind coxae orange-brown;
wings hyaline; posterior ocelli about equidistant from
eye and occipital margins; antenna (Fig. 56) with F1
about 2.5x length of pedicel; forewing (Fig. 57) about
2.9x as long as broad, with postmarginal vein about
1.75x as long as stigmal (Fig. 59); linea calva inter-
rupted by 2 lines of setae (Fig. 58); gaster as long as
thorax; ovipisitor about 10x as long as gonostyli. Male
(length 1.30—1.70 mm): generally similar to female
but for antennae and genitalia; yellow coloration of
head more extensive then in female; antennae with
about 10 scale-like structures ventrally on base of
clava; forewing (Fig. 60) about 2.5x as long as broad;
linea calva interrupted by only a single setae; aedeagus
about one-third as long as mid tibia, each digitus with
two long apical teeth.

Hosts. Leptomastix abyssinica has been recorded
from Planococcus citri on Olea chrysophylla (Com-
pere, 1931, 1938).

DISTRIBUTION. Eritrea.

MATERIAL EXAMINED.

Type material. Paratypes: Eritrea, 11 9 , 30’, Nefasit,
ex Pseudococcus citri on Olea chrysophylla, i11.1930
and iv.1930(Compere); | 9 , Nefasit, ex Pseudococcus
citri on Olea chrysophylla, iv.1930 (Compere); 3 & ,
20, Nefasit, ex Pseudococcus No 4630 on Olea
chrysophylla, 15.iv.1930 (Compere); 10, Nefasit, ex
Pseudococcus No 4630 on Olea chrysophylla,
16.iv.1930 (Compere); 1 2 , Nefasit, on window, out of
mealybug, 31.iii.1930 (Compere). Material in BMNH
and UCR.

Non type material. Eritrea, 50’, Nefasit, ex
Pseudococcus No 4630 on Olea chrysophylla,
11.iv.1930 (Compere); 40°, Nefasit, ex Pseudococcus
No 4630 on Olea chrysophylla, 10.iv.1930 (Com-
pere); 19, Ezdaclesan, ex Croton macrostachys,
1.v.1930 (Compere); 592, 1o&, Nefasit, ex
Pseudococcus citri (Compere). Material in UCR.

COMMENTS. Leptomastix abyssinica is close to
Leptomastix africana, females of both species having
similar dark-brown general coloration, the forewing
about 2.7—2.9x as long as broad and two lines of setae
interrupting the linea calva. They differ by the colour
of the scape, lower face, prosternum and forelegs,
position of posterior ocelli in relation to occipital
margin and relative length of the clava. In abyssinica
the scape is yellow ventrally, the lower face, prosternum
and fore coxae are yellow, the posterior ocelli are
equidistant from the eye and occipital margins and the
clava is longer than F1 (Fig. 56), whereas in africana
the scape is entirely brown, the prosternum and coxae
are dark brown, the posterior ocelli are nearer the

111

occipital margin and the clava is shorter than F1 (Fig.
22). Males of the two species can be separated by
colour as in the female. Leptomastix abyssinia might
also be mistaken for Leptomastix nigra (see comments
under nigra).

HOSTS OF AFRICAN SPECIES OF
LEPTOMASTIX

* extralimital record
? doubtful record
! laboratory rearing

HEMIPTERA
Coccidae

Parasaissetia nigra (Nietner) (=Saissetia nigra) —
?*nigrocoxalis
Saissetia nigra, see Parasaissetia nigra

Dactylopiidae

Dactylopius destructor, see Planococcus citri
(Pseudococcidae)

Margarodidae
Icerya aegyptiaca (Douglas) — *nigrocoxalis
Pseudococcidae

Allococcus quaesitus, see Delottococcus quaesitus

Birendracoccus saccharifolii (Green) (=Phenacoccus
saccharifolii) — ?*dactylopii

Coccidohystrix — *nigrocoxalis

Coccidohystrix insolita (Green) — *nigrocincta,
*nigrocoxalis

Delottococcus quaesitus (Brain) (= Allococcus
quaesitus) — dactylopii

Dysmicoccus brevipes (Cockerell) — *dactylopii

Ferrisia virgata (Cockerell) —dactylopii, *tsukumiensis

Maconellicoccus hirsutus (Green) — nigrocoxalis

Nipaecoccus — *nigrocoxalis

Nipaecoccus graminis (Maskell) — nigrocoxalis

Nipaecoccus viridis (Newstead) — nigrocoxalis

Octococcus — herreni, africana, nigra

Octococcus africanus (Brain) — nigra

Octococcus minor De Lotto — nigra

Octococcus pentziae Hall — herreni, nigra

Paracoccus burnerae (Brain) — nigra

Phenacoccus — *nigrocincta

Phenacoccus gossypii Townsend & Cockerell —
!2*dactylopii

Phenacoccus madeirensis Green — dactylopii

Phenacoccus saccharifolii, see Birendracoccus
saccharifolii

Phenacoccus solani Ferris — !dactylopii

Planococcoides njalensis (Laing) — dactylopii

112

Planococcoides lamabokensis Balachowsky & Ferrero
— dactylopii
Planococcus — dactylopii
Planococcus aemulor De Lotto — dactylopii
Planococcus citri (Risso) (= Dactylopius destructor) —
abyssinica, dactylopii, *nigrocoxalis
Planococcus ficus (Signoret) — dactylopii
Planococcus kenyae (Le Pelley) — dactylopii
Planococcus kraunhiae (Kuwana) — *dactylopii
Planococcus lilacinus (Cockerell) — dactylopii
Planococcus vovae (Nasonov) — *dactylopii
Pseudococcus — dactyolopii, nigra, nigrocoxalis
Pseudococcus adonidum, see Pseudococcus
longispinus
Pseudococcus bukbensis Laing — dactylopii
Pseudococcus calceolariae (Maskell) — !dactylopii
Pseudococcus comstocki (Kuwana) — !dactylopii
Pseudococcus concavocerarii James — dactylopii
Pseudococcus inamabilis, see Planococcus vovae
Pseudococcus longispinus (Targioni Tozzetti) (=
Pseudococcus adonidum) — dactylopii
Pseudococcus occiduus De Lotto — dactylopii
Rastrococcus cappariae Avasthi & Shafee —
*nigrococalis
Rastrococcus iceryoides (Green) — *nigrocoxalis
Spilococcus — nigra

NEUROPTERA

Nimboa basipunctata Withycombe — ?*nigrocoxalis

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Tingle, C.C.D. & Copland, M.J.W. 1988b. Effects of temperature
and host-plant regulation of glasshouse mealybug (Hemiptera,
Pseudococcidae) populations by introduced parasitoids (Hymenop-
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135-142.

Tingle, C.C.D. & Copland, M.J.W. 1989 Progeny production and
adult longevity of the mealybug parasitoids Anagyrus pseudococci,
Leptomastix dactylopii and Leptomastidea abnormis (Hymenop-
tera: Encyrtidae) in relation to temperature. Entomophaga 34:
111-120.

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(Edited: Medvedev, G.S.)

Trjapitzin, V.A. 1989. Parasitic Hymenoptera of the Fam. Encyrtidae
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REVISION OF LEPTOMASTIX

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115

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Entomologia Agraria ‘Filippo Silvestri’, Portici 18: 1-148.

INDEX

Synonyms and misidentifications are in italics; main citations in bold.

abyssinica (Leptomastix) 97, 98, 103,
106, 110-111, 112, 126

Acacia 104, 107

Achyranthes 104

adonidum (Pseudococcus) 107, 112

aegyptiaca (Icerya) 104, 111

aemulor (Planococcus) 107, 112

africana (Leptomastix) 98, 103, 110, 111,
120-121

africanus (Octococcus) 105, 106, 111

alba (Morus) 104

algirica (Leptomastix) 98, 101, 102-103,
120, 127

aligarhensis (Leptomastix) 103

Allococcus 107, 108, 111

Anagyrus 100

Aphytis 94

arabica (Coffea) 108

aspera (Achyranthes) 104

Athenasia 101, 102, 105, 106

azedarach (Melia) 104

basipunctata (Nimboa) 104, 112
bicalyculata (Peritropha) 104
bifasciatus (Leptomastix) 106, 107
Birendracoccus 107, 111

brevipes (Dysmicoccus) 107

brevis (Leptomastix) 103, 104, 111
bukbensis (Pseudococcus) 107, 112
burnerae (Paracoccus) 105, 106, 111

cactearum (Spilococcus) 107

calceolariae (Pseudococcus) 97, 107, 112

cappariae (Rastrococcus) 104, 112

carica (Ficus) 107, 108

Cassiea 108

Centrococcus 104

cephalotus (Leucas) 104

Ceroplastes 105, 106

Chrysocoma 101, 102

chrysophylla (Olea) 111

cinerea (Stoebe) 105, 106

citri (Planococcus, Pseudococcus) 96, 97,
104, 107, 108, 111, 112

Citrus 99, 104

Coccidohystrix 104, 110, 111

Coffea 108

coma-aurea (Chrysosoma) 101, 102

Combretum 107

comstocki (Pseudococcus) 97, 107, 108,
112

concavocerarii (Pseudococcus) 107, 112

Croton 111

cryptus (Pseudococcus) 103

dactylopii (Leptomastix) 96, 97, 98, 99,
105, 106-108, 109, 111, 112, 123-124,
127, 128

Dactylopius 107, 111, 112

Delotococcus 107, 111

destructor (Dacylopius) 107, 111, 112

Digitaria 100

digitariae (Leptomastix) 96, 98, 100-101,
102, 118-119, 127, 128

Diospyros 105, 106

Dysmicoccus 107, 111

elegans (Loranthus) 105, 106
Elytropappus 103, 105, 106

epona (Leptomastix) 103, 105, 110
ericoides (Rehlania) 101, 102
Erythrina 107, 108

Euclea 108

Euphorbia 104

Ferrisia 99, 107, 111

ficus (Planococcus) 107, 108, 112

Ficus 107, 108

filamentosus (Pseudococcus) 104

flava (Leptomastix) 97, 102, 103, 104,
106, 107, 108

Gazania 105, 106

genistaefolia (Rehlania) 102

globosa (Pentzia) 101, 102, 105, 106
gnaphaloides (Elytropappus) 105, 106
gossypii (Phenacoccus) 107, 111
graminis (Nipaecoccus) 104, 111

guajava (Psidium) 107

Hemigraphes 104

herreni (Leptomastix) 98, 100, 100-102,
105, 106, 110, 111, 119-120, 127

Hibiscus 104, 110

hirsutus (Maconellicoccus) 97, 104, 111

hirta (Hyparrhenia) 104

histrio (Leptomastix) 95, 105, 110

histrio (Leptomastix, Sterhocoma) 95

humberti (Digitaria) 100

Hyparrhenia 104

Icerya 104, 111

iceryoides (Rastrococcus) 104, 1112

imbricata (Oedera) 101, 102

inamabilis(Pseudococcus) 107, 112

indica (Tamarindus) 104

insolita (Centrococcus,
Coccidohystrix) 104, 110, 111

invadens (Rastrococcus) 94

javanica 108
jonesi (Leptomastix) 98, 108-109, 124,
127

kenyae (Planococcus) 97, 107, 108, 112
kraunhiae (Planococcus) 107, 112

lamabokensis (Planococcoides) 107, 112
lappacea (Pupalia) 104

Leucas 104

leucopogi (Spilococcus) 107

lilacinus (Planococcus) 107, 112
longipennis (Leptomastix) 98, 106, 107
longiscapus (Leptomastx) 103, 104
longispinus (Pseudococcus) 107, 112
Loranthus 105, 106

lyciae (Leptomastix) 109

lycioides (Diospyros) 105, 106
lysistemon (Erythrina) 107, 108

Maconellicoccus 97, 104, 111
macrostachys (Croton) 111
madeirensis (Phenacoccus) 107, 111

116

manihoti (Phenacoccus) 94
maritimus (Pseudococcus) 107
medica (Citrus) 104

Melia 104

melongena (Solanum) 110
Metalasia 105, 106

minor (Octococcus) 105, 106, 111
Morus 104

muricata (Metalasia) 105, 106

Nerium 105

nigra (Leptomasti 98, 102, 104, 105-106,

110, 111, 122-123

nigra (Parasaissetia, Saissetia) 104, 105,
106, 108, 111, 112

nigrocincta (Lepomastix) 98, 103, 105,
108, 109-10, 111, 112, 125-126, 127

nigrocoxalis (Leptomastix) 96, 97, 98,
102, 103-105, 106, 108, 110, 111, 112,
121-122

Nimboa 104, 112

Nipaecoccus 97, 104, 111

njalensis (Planococcoides) 97, 107, 111

occiduus (Pseudococcus) 107, 112
Octococcus 101, 102, 103, 105, 106, 111
Oedera 101, 102

Olea 111

oleander (Nerium) 105

orbitalis (Stenoterys) 95

Paracoccus 105, 106, 111

Parasaissetia 105, 106, 111

Pentzia 101, 102, 105, 106

pentziae (Octococcus) 101, 102, 105, 106,
111

Peritropha 104

phenacocci (Leptomastix) 96, 103, 104

Phenacoccus 94, 104, 107, 110, 111, 112

Planococcoides 97, 107, 111, 112

Planococcus 96, 97, 104, 107, 108, 111,
112

plumosa (Stoebe) 106

Prosopis 108

Pseudococcus 97, 103, 104, 105, 107,
108, 112

Psidium 107

Pupalia 104

purpurea (Tephrosia) 104

quaesitus (Allococcus,
Delotococcus) 107, 108, 111

Rastrococcus 94, 104, 112

Rehlania 101, 102

rhinocerotis (Elytropappus) 103, 106
rigens (Gazania) 106

saccharifolii (Birendracoccus, Phena-

J-M. ANGA AND J.S. NOYES

coccus) 107, 111
Saissetia 104, 105, 106, 111
singularis (Leptomastix) 98
solani (Phenacoccus) 107, 111
Solanum 110
somereni (Saissetia) 105, 106
somnifera (Wittania) 104
Spilococcus 105, 106, 107, 112
splendens (Combretum) 107
Stenoterys 95
Sterrhocoma 95
Stoebe 105, 106
superba (Leptomastix) 106, 107, 108

Tamarindus 104

Tambourissa 107

tambourissae (Leptomastix) 107

Tephrosia 104

trifurcata (Athenasia) 101, 102, 105, 106

tsukumiensis (Leptomastix) 96, 98-100,
107, 108, 111, 117-118, 128

virgata (Ferrisia) 99, 107, 111
viridis (Nipaecoccus) 97, 104, 111
vitis (Planococcus) 108

vovae (Planococcus) 107, 112

Wittania 104

Ziziphus 104

Sp 25

REVISION OF LEPTOMASTIX 117

Figs 1-6. Leptomastix tsukumiensis — 1, antenna, variant, 9 ; 2, antenna, variant, Q : 3, forewing, variant, 9 ; 4, forewing,
variant, ; 5, forewing, variant, 2 ; 6, forewing, apex of venation, variant, 9 .

J-M. ANGA AND J.S. NOYES

118

fe an tek

ee

Figs 7-12. Leptomastix tsukumiensis — 7, forewing, apex of venation, variant, 2 . Leptomastix digitariae — 8, antenna, Os

9, forewing, @ ; 10, forewing base, 2 ; 11, forewing, apex of venation, 2 ; 12, ovipositor.

REVISION OF LEPTOMASTIX MG)

Figs 13-18. Leptomastix digitariae — 13, forewing,C’; 14, forewing, apex of venation,©’. 15-18, Leptomastix herreni — 15,
antenna, © ; 16, forewing, ; 17, forewing base, 2 ; 18, forewing, apex of venation, 2.

120 J-M. ANGA AND J.S. NOYES

Figs 19-24. Leptomastix herreni — 19, ovipositor; 20, forewing, apex of venation, O’. Leptomatix algirica — 21, forewing,
apex of venation. Leptomastix africana — 22, antenna, 9 ; 23, forewing, 9 ; 24, forewing base, 2 .

REVISION OF LEPTOMASTIX 121

Figs 25-30. Leptomastix africana — 25, forewing, apex of venation, 9 . Leptomastix nigrocoxalis — 26, antenna, @ ; 27,
forewing, 9 ; 28, forewing base, 9 ; 29, forewing, apex of venation, 9 ; 30, forewing,c’.

122 J-M. ANGA AND J.S. NOYES

Figs 31-36. Leptomastix nigrocoxalis — 31, forewing base,O’. Leptomastix nigra — 32, antenna, large specimen, es 313)
antenna, small specimen, @ ; 34, forewing, 9 ; 35, forewing, apex of venation, Q ; 36, ovipositor.

REVISION OF LEPTOMASTIX 123

Figs 37-42. Leptomastix nigra — 37, forewing,’ ; 38, forewing base, 0’. Leptomastix dactylopii — 39, antenna, @ ; 40,
forewing, variant, ? ; 41, forewing, variant, 2 ; 42, forewing base, @ .

124 J-M. ANGA AND J.S. NOYES

Figs 43-48. Leptomastix dactylopii — 43, forewing, apex of venation, ? ; 44, forewing,’ ; 45, forewing base, 0’; 46,
forewing, apex of venation, O’. Leptomastix jonesi — 47, forewing,’ ; 48, forewing, apex of venation,@’.

REVISION OF LEPTOMASTIX W3)

Figs 49-54. Leptomastix nigrocincta — 49, antenna, variant, 9 ; 50, antenna, variant, 9 ; 51, forewing, 2 ; 52, forewing
base, 9 ; 53, forewing, apex of venation, 9 ; 54, forewing, C’.

126 J-M. ANGA AND J.S. NOYES

Figs 55-60. Leptomastix nigrocincta — 55, forewing base,O". Leptomastix abyssinica — 56, antenna, ? ; 57, forewing, Q;
58, forewing base, 9 ; 59, forewing, apex of venation, 9 ; forewing, ©’.

REVISION OF LEPTOMASTIX 127

Figs 61-65. Leptomastix herreni— 61, antenna, scale-like structure enlarged," . Leptomastix algirica — 62, antenna, scale-
like structure enlarged,’ . Leptomastix dactylopii — 63, antenna, scale-like structure enlarged, C’. Leptomastix jonesi — 64,
antenna, scale-like structure enlarged, ’. Leptomastix digitariae — 65, antenna, scale-like sstructure enlarged, O’.

128 J-M. ANGA AND J.S. NOYES

69 70

Figs 66-70. Leptomastix nigra — 66, antenna, larger specimen, scale-like structure enlarged, ©’; 67, antenna, smaller
specimen, . Leptomastix tsukumiensis — 68, antenna, scale-like structure enlarged,’ . Leptomastix digitariae — 69,
genitalia, (a) complete, (b) apex of phallobase enlarged, &’. Leptomastix dactylopii — 70, genitalia, (a) complete, (b) apex
of phallobase enlarged, oO’.

Bull. nat. Hist. Mus. Lond. (Ent.) 68(2): 129-144

A revision of the north-west
European species of the formosus
species group of Spilomicrus
(Hymenoptera, Diapriidae)

DAVID G. NOTTON

Reading Museum Service, The Museum of Reading, Blagrave Street, Reading, RG1
1QH, United Kingdom.

CONTENTS
SWIMO SIS cesnceaeceesstessecesas<cdetassset unease secsscoaccaanieist ssuesutensasodeseeee, Couameaters camaro tnczescas cate onde ec etes 129
ISL IN CUS EL OD waecyas ace oats cs trpiserscdls ay sspscayncatcsivineuadustendtacee trance sstaauasataaperetsdvareues etucasatswceeeeettecraas 129
IDEMOSILQUIES, cos cers cvscas acts hzgscuinassasncneidetscasssestnendsaesdaaceatcuumerceesesteone tvee ta aacaten oe acai textoba te ceucesenates 130
PERO WIEUP CMCALS iesactssesucacascenaceoeeeeoccacthccavee-vans sncasToetis esac eeteee co eaee see este nue ta sa oe eae cnet 130
SS PIULONTICI TASS WCSUMOOW es ccaceceocsacseverecscevarecsavovztsaranndesseetuacodetnusvorentetararsces to Nore n enema cena eras 130
MEY OPIMOSTES! SPECICS I PLOUP eeazasectctesce=suc---va<ssentexasasanews cuscetasavdvesaseuecdat sete -cercet lavastean-caceers 130
Key to north-west European species of the formosus SpeCie€S QrOUP ...........:ceccceceeseeteeeeeeee 132
SU MOTCESSTCLO VES INTELECD coe one seme arestinesterese vt vos vas sunsek xacscessincl deapesuacepseisaaeusesasadadesesvadedansasaasess 132
DVM OF FLO SEIS ALLSS OL cece deere cect tess se eee sca cnn cu scesatenaabtoestadcbcciee sccbesuheteedesuxeeueadustousecccsesensees 134
SSSRNCAILOOVIECIANTASIIEIN: «. pteentenscecasteeetrereccsidesscaacouersccavetescepceacssdsaasacsssecadacansateeesaccecotenentettte 136
IRELEMEM CES Mramnec setts tres setcedecnduse heatoetots covtscuececcecccesscdeccusestendtedavtsaceavscescegcesceccesesddeeteoneadesseatades 137
Ertl eae wemne sae renee. mee reir neat at ce cote SMMMNR ALES 1s wa cocepetacehasiecctensunuanadentctudisaduastvestecenssscesseteaes 138

Synopsis. A revisionof north-west European species of the formosus species group of Spilomicrus
is given including an outline of the characters of the group, a key to all species and descriptions
of all species. S. crassiclavis is removed from synonymy with S. integer. S. pelion is placed in
synonymy with S. crassiclavis. Lectotype designations are made for S. crassiclavis and S.
formosus. S. sanbornei is recorded as new to the Palaearctic and S. crassiclavis and S. formosus

Issued 25 November 1999

as new to the east Palaearctic.

INTRODUCTION

The formosus group of Spilomicrus is a small species
group of inconspicuous diapriid parasitoid wasps with
seven described species from the Palaearctic and
Nearctic regions. The group may distinguished from
other Spilomicrus by the unusual form of the clypeus,
which protrudes in front of the mandibles as a triangu-
lar or rounded projection and by associated
modifications of the face (Figs 2-3, 5-6). Some mem-
bers of this group have an unusual way of life as
pseudohyperparasitoids of cicadellid bugs via the
puparia of pipunculid flies. The formosus group is also
significant because, although only a small group, it
includes two species, S. formosus and S. sanbornei,
with Holarctic distributions. It is becoming clear that
such distributions are not uncommon among
Diapriinae; examples are known within other species

© The Natural History Museum, 1999

groups of Spilomicrus, e.g. S. antennatus (Jurine) and
S. stigmaticalis Westwood (Masner, 1991) as well as
species belonging to a number of other diapriine genera.

The aim of this paper is to revise the formosus
species group of Spilomicrus occurring in north-west
Europe. A revision was needed because of the confu-
sion over species resulting from the mixed and partly
lost type series of S. crassiclavis and from later
misidentifications. Type material of all species was
examined and type restrictions made where necessary.
Nomenclatural changes were made as follows:
lectotype designations were made for S. formosus and
S. crassiclavis; S. crassiclavis was removed from
synonymy with S. integer; S. pelion was placed in
synonymy with S. crassiclavis. Currently three spe-
cies are recognised in the region and, since previous
keys are now inadequate, a new partial key to
Spilomicrus is presented here. S. crassiclavis is re-
corded as new to Czechia, Denmark, Finland, Japan,

130

Norway and Sweden and S. formosus as new to Bel-
gium, Ireland, Japan and Norway. S. sanbornei is
recorded as new to Britain and the Palaearctic.

Morphological terminology largely follows Masner
(1991). Wing vein terminology follows the conven-
tion of Mason (1986).

DEPOSITORIES

BMNH The Natural History Museum, London, United
Kingdom (formerly the British Museum, Natural

History)

CNC Canadian National Collection of Insects,
Arachnids and Nematodes, Ottawa, Canada

DN D. G. Notton collection, Reading, United
Kingdom

NMS National Museums of Scotland, Edinburgh,

United Kingdom
RAS Russian Academy of Sciences, Palaeontological
Institute, Arthropod Laboratory, Moscow, Russia
SEL Department of Applied Zoology, University of
Helsinki, Helsinki, Finland

ZI Universitets Zoologiska Institutionen, Lund,
Sweden
ZM Zoological Museum, University of Copenhagen,

Copenhagen, Denmark

ACKNOWLEDGEMENTS. Thanks are due to the above insti-
tutions and the following people: Ferdinando Bin; Peter Buhl;
Roy Danielsson; the staff of the Entomology Library of the
Natural History Museum, London; Jeremy Field; Mike Fitton;
Gary Gibson; the Hamiltons of Abbots Moss; Peter Holmes;
Karen Hull; Viktor Kolyada; Martti Koponen; Nina Laurenne;
Suzanne Lewis; Andrew Polaszek; Mark Shaw; Lars
Vilhelmsen; Claire Villemant and especially to Lubomir
Masner.

SPILOMICRUS WESTWOOD

Spilomicrus Westwood, 1832: 129. Type species:
Spilomicrus stigmaticalis Westwood by monotypy.
Extensive generic synonymy may be found in
Masner (1991) and Johnson (1992).

DIAGNOSIS (north-west European _ species
only). Antenna 13-segmented in both sexes; for the
female, apical antennal segment subequal to the
preapical; for the male the fourth antennal segment
usually modified; lateral surface of pronotum without
a row of foveae; scutellum with two anterior pits,
sometimes faint in some females of S. antennatus;
macropterous to apterous; winged forms with costal
cell wide, 2—3 times width of submarginal vein and

D.G. NOTTON

marginal vein short, not more than twice length of
stigmal vein; anterior margin of metasomal syntergite
(2+3) entire, and without keels, ridges or grooves,
fitting loosely over apex of petiole and more or less
hiding it; female with apex of gaster blunt to moder-
ately conical.

DISTRIBUTION. Spilomicrus is a worldwide genus
with many species: Johnson includes 144 described
species in his World catalogue and there are many
more undescribed. The number of species in north-
west Europe is unclear: Nixon (1980) recorded 14 in
Britain and Kozlov (1978) 22 in Europe, however
there is no recent authoritative treatment covering
north-west Europe. Of the four species groups recog-
nised by Masner (1991) only the stigmaticalis group
and the formosus group have been found in north-west
Europe, where they are both widely distributed.

BIOLOGY. Spilomicrus species have been recorded
mostly as solitary endoparasitoids of the puparia of
Diptera, including the host families Tephritidae,
Tachinidae and possibly Bibionidae (Hoffmeister,
1989); Syrphidae, Pipunculidae, Muscidae,
Sciomyzidae (Masner, 1991); Phoridae (Disney, 1994).
A small number of species have been reared from the
pupae (not larvae) of Staphylinidae (Coleoptera)
(Nixon, 1980; Masner, 1991; Notton, 1996).
Hoffmeister (1989) gives an account of the laboratory
culture of Spilomicrus hemipterus Marshall on a spe-
cies of Drosophila (Diptera, Drosophilidae) including
descriptions of larval morphology. Spilomicrus spe-
cies occur mainly in forest habitats although some
may be common in grassland or wetland.

COMMENTS. North-west European Spilomicrus may
be recognised using the generic key of Nixon (1980).
The taxonomy of Spilomicrus on a world scale is
poorly understood as are the interrelationships of its
major species groups. Within north-west Europe many
types need to be located and reinterpreted and a number
of disparate regional studies need to be reconciled (see
refs. in Masner, 1991 and Johnson, 1992). The species
groups recognised by Masner (1991) are admittedly
only provisional but are practical working groups: in
particular the formosus group is easily recognisable in
north-west Europe from the form of the clypeus (see
key below).

formosus species group: Masner

(Figs 2-20)

formosus species group of Spilomicrus: Masner, 1991:
111.

DESCRIPTION. Habitus robust. Head in dorsal view
transverse, 1.1—1.4x as wide as long; surface of head
with scattered, long, semidecumbent hairs; occipital

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 131

flange narrow but distinct, not foveolate medially;
head in frontal view subcircular, slightly transverse;
space between anterior margins of toruli without a
carina; face just below toruli smooth, without punc-
tures; clypeal margin medially with a ventrally directed,
triangular or rounded projection, either side of this
projection the clypeal margin is strongly impressed
forming two deep concavities; tentorial pits absent;
malar sulcus complete, strongly curved. Antenna: scape
smooth and shining at least in part, if sculptured then
no more than weakly coriaceous; female antenna with
5- to 8-segmented club; male antenna with segment 3
equal to or a little shorter than segment 4, segment 4
cylindrical to weakly expanded apically with a weak
to moderately deep emargination and carina or flange
in basal 0.3-0.7. Mandibles stout, with upper tooth
short, about 0.5—0.6 of length of the lower. Mesosoma:
pronotum without epomia, weakly concave between
the projecting pronotal shoulder and spiracle, anteriorly
with long, scattered grey hairs which do not form a
distinct cushion, posteriorly with a narrow strip of
short, fine hairs extending downwards from the
spiracle; acetabular carina complete, extending from
the lower corner of the pronotum to the mid-line of the
mesosternum; transscutal sulcus weakly curved; ante-
rior scutellar pits large, deep, rounded, 0.35—0.4 of
length of scutellum, separated by a narrow medial
carina, which broadens out posteriorly into the scutellar
disc; lateral scutellar pits broad, deep, broadened
posteriorly; 7—12 distinct, posterior, scutellar pits;
medial propodeal keel low posteriorly, its highest
point anteriorly well below level of mesoscutum. Fore
wing with subcostal, marginal and stigmal veins tubu-
lar; costal, basal, cubital and radial sector veins
nebulous; costal cell broad, at widest point at least
2.5x as wide as subcostal vein; marginal vein short,
thick, about 1.7—3.0x as long as wide when its length
is measured from the start of its basal expansion to the
fork of the postmarginal and stigmal veins, and its
width measured at the mid-point of this length; stigmal
vein short to very short; radial area indicated by distal
abscissa of radial sector and anterior wing margin
long, extending 0.45—0.60 of distance between mar-
ginal vein and wing tip. Legs unremarkable; tarsal
segments slender; basitarsi longer than telotarsi.
Metasoma with dorsally visible part of petiole rela-
tively short, 1.2—2.2x as long as wide, a little flared
anteriorly, sparsely hairy at base; tergites 4-7 with
scattered micropunctures and scattered hairs; sternites
3-6 (2) or 3-7 (c) with deep, fine micropunctures.
Ovipositor sheaths and apical sternite without pegs or
dense pilosity. Colour: body mainly black; fore wing
weakly tinged brown, without maculae.

DISTRIBUTION. The formosus group is widely dis-
tributed throughout the Nearctic, Palaearctic, Oriental
and Australian regions (Masner, 1991) although only

seven species are described so far from the Palaearctic
and Nearctic. Currently three species are recognised
from north-west Europe. A number of species are
widespread: S. formosus is known to be Holarctic
(Masner, 1991) and S. sanbornei was found to be so in
the current study.

BIOLOGY. The only published host record for the
formosus group (Masner, 1991) is for S. formosus
itself: much reared material of this species was also
seen during this study and supported Masner’s obser-
vation that it attacks the puparia of Pipunculus species
(Diptera, Pipunculidae) in Europe. It is therefore a
pseudohyperparasitoid of cicadellid bugs (the primary
hosts of Pipunculus species — Coe, 1966). Curiously
Spilomicrus formosus and S. crassiclavis show a
number of morphological similarities with some
diapriines of the genus Basalys including B. fumipennis,
B. tuberculatus and B. erythropus which also attack
pipunculid puparia (Lundbeck, 1922; this study). These
similarities include: a compact, robust habitus, trans-
verse head and a short mesosoma with domed
mesoscutum. It is speculated that this apparent conver-
gence has resulted from independent adaptation to
development inside the short ovoid puparia of
pipunculids.

COMMENTS. The formosus group was recognised by
Masner (1991) who defined the group by the posses-
sion of the following putative apomorphies: presence
of malar sulci; a pointed clypeus; mandibles with the
lower tooth stronger than the upper. On the basis of
species included by Masner, the group is difficult to
define because of a number of exceptions to the above
characters: malar sulci do not occur throughout the
formosus group and occur in some other Spilomicrus
and other Spilomicrini such as /diotypa; when viewed
from in front, the clypeus is not pointed in all formosus
group species, although clearly pointed in S. formosus,
it is rounded in a number of species including S.
sanbornei; lastly, while the mandibles do have the
lower tooth stronger in the formosus group, this can
occur in other Spilomicrus. Thus none of the three
characters given by Masner uniquely defines the
formosus group, however, from material seen during
this study and from Masner’s descriptions, this group
may still be defined by a character of the clypeus, that
is, in all species the clypeus protrudes well over the
mandibles as a pointed or rounded projection. In addi-
tion to the malar sulcus and mandible characters noted
by Masner, other supporting characters which occur
frequently though not universally in this species group
include: absence or reduction of tentorial pits; absence
of the epistomal suture; moderately to strongly im-
pressed clypeal margin either side of the medial
projection; weakly marked dorsal margin of the clypeus
so that in lateral view the surface of the face forms an
even curve.

132

Key to north-west European species of
the formosus species group of
Spilomicrus

1 Clypeus not protruding over mandibles (Fig. 1); clypeal
margin not strongly impressed submedially; malar sulcus
usually absent; tentorial pits usually present .................

a eee Aer een Jae Spilomicrus stigmaticalis group

— Clypeus protruding over mandibles as a pointed or
rounded projection (Figs 2, 3, 5, 6); clypeal margin
strongly impressed submedially; malar sulcus present
and strong; tentorial pits absent ..............:cccceeeeeeeeeenees
cen oso ati (Spilomicrus formosus group) 2

2 Antennal flagellum apically clubbed (Figs 7, 10, 13);
post-petiole with 6 stermites ..............eeeeee (females) 3

— Antennal flagellum not clubbed (Figs 9, 12, 15); post-
petiole with) 7 stemites -....c2.-0...ee.cc-bececeenceseee (males) 5

3. Antennal club 5- to sub-6-segmented (Fig. 13); in frontal
view, apex of medial projection of clypeus rounded,
blunt (Fig. 5); mesosoma depressed, no more than 0.8x
as high as wide, mesoscutum weakly convex (Fig. 16);
medial propodeal keel low, hardly raised in front (Fig.
16); body length 1.9-2.5 mm ........... eee sanbornei

— Antennal club 7- to 8-segmented (Figs 7, 10); in frontal
view, apex of medial projection of clypeus triangular,
pointed (Figs 2-3); mesosoma less depressed, at least
0.9x as high as wide, mesoscutum strongly convex (Fig.
18); medial propodeal keel raised in front to form a right-
angled to acute projection (Fig. 18); body length 2.23.1

4 Antennal club 7-segmented, with segments 8—12 strongly
transverse (Fig. 7); notauli convergent anteriorly, ex-
tending over no more than basal 0.25 of mesoscutum
(Fig. 19); eye height 1.4-1.5x as long as malar space
(BIG D2) veces concn cen conn sctese cose rere sctesteertecateccnes crassiclavis

— Antennal club 8-segmented, with segments 8—12 quad-
rate to slightly elongate (Fig. 10); notauli divergent
anteriorly, extending over at least basal 0.35 of meso-
scutum (Fig. 20); eye height 1.7—2.2x as long as malar
SPace*(HIGNS) ee. cccestceecess tees e eieeesee toot erieetess formosus

5 In frontal view, apex of medial projection of clypeus
rounded, blunt (cf. 9— Fig. 5); antennomere 4 with a
moderately deep, curved emargination over basal 0.45—
0.5 (Fig. 14); mesosoma depressed, no more than 0.8x as
high as wide (cf. 9 — Fig. 16); medial propodeal keel low,
hardly raised in front (cf. 2 — Fig. 16); body length 2.0—
QBN SE o RSee ses decease eieate eecGces ceteae aeteesee ee sanbornei

— In frontal view, apex of medial projection of clypeus
triangular, pointed (cf. 9 Q— Figs 2-3); antennomere 4
with at most a shallow emargination, over basal 0.3—0.7
(Figs 8, 11); mesosoma less depressed, at least 0.9x as
high as wide (Fig. 17); medial propodeal keel raised in
front to form a right-angled to acute projection (Fig. 17);
bodyslenm thn 2 UE 3 S/n sexes eee cceeeeseeteenete see erate 6

6 Méesopleuron with a sternaulus, delimited dorsally by a
sharp carina (Fig. 17); apical antennomere no longer than

D.G. NOTTON

preapical (Fig. 9); notauli absent or vestigial, extending
over no more than posterior 0.2 of mesoscutum (cf. 2 —
Fig. 19); eye height 1.0x as long as malar space; surface
of eye with sparse hairs ...........:.ceeeeereeeees crassiclavis

— Mesopleuron without such a sternaulus or carina (cf. 9 —
Fig. 18); apical antennomere at least 1.5x as long as
preapical (Fig. 10); notauli extending over at least poste-
rior 0.35 of mesoscutum (cf. 9 — Fig. 20); eye height 2.3x
as long as malar space; surface of eye without hairs .....

eevee daha eth baa Geesuei ston a cote eae Dees, ee eee formosus

Spilomicrus crassiclavis Kieffer
(Figs 2, 7-9, 17, 22)

Spilomicrus crassiclavis Kieffer, 1911: 788, 797.
LECTOTYPE, Great Britain: Mugdock
(BMNH), here designated [examined]. Stat. rev.
Lectotype labels: Mugdock, 11—7 [under mount];
Cameron coll., 1910-302; Mugdock; Determined
by Dr. Kieffer.; Spilomicrus crassiclavis K.;
Spilomicrus formosus Janss., G. Nixon det., 1972;
B.M. TYPEHYM. 9.645a. Lectotype glued onto a
card, with the right fore wing disarticulated, apical
segment of left antenna lost and right hind telotarsus
disarticulated.

[Spilomicrus formosus Jansson: Nixon, 1980: 16, Figs
30, 31. Misidentification in part (2 ).]

Spilomicrus pelion Nixon, 1980: 17. Holotypeo’, Great
Britain: Surrey (BMNH) [examined]. Syn. n.
Holotype labels: Holotype [red BMNH type label];
Surrey, Horsley, 14.vi.1930, G. Nixon; Spilomicrus
pelion Nixon, Typec’, 197; B. M. TYPE HYM.
9.811. Holotype entire, mounted on a card point.

DESCRIPTION. 9. Head (Fig. 2) in dorsal view 1.25—
1.32x as wide as long; temples weakly to moderately
contracted just behind eyes; dorsal part of occipital
flange barely widened medially; post-genal pilosity
sparse, not forming a defined cushion; head, in lateral
view suboval, 1.13—1.17x as high as long; frons be-
hind toruli weakly convex, almost flat; eye sparsely
hairy; eye height 1.4—1.5x as long as malar space; in
frontal view, apex of medial projection of clypeus
triangular, pointed when seen from in front; lateral
margins of clypeus undefined; malar sulcus a finely
impressed line becoming a little deeper towards man-
dible. Antenna (Fig. 7): scape entirely smooth or at
most weakly coriaceous at base; segment 3 about 2.5x
as long as wide; segments 7-13 expanded to form a
long-fusiform 7-segmented club; segments 8-12
strongly transverse; apical segment 1.49-1.74x as
long as the preapical and slightly narrower than it;
apical segment without a ventral pit. Mesosoma 1.29—
1.37x as long as wide; anteriorly mesoscutum usually
without a pair of short, submedial, parallel lines, rarely
these are weakly indicated; notauli short but sharply
incised, deep, weakly sinuate, narrowed anteriorly,

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 133

extending over posterior 0.18-0.24 of mesoscutum
with no trace in front of this, convergent anteriorly
(Fig. 19); humeral sulcus running 0.5—0.7 of the way
from the transscutal suture to the posterior corner of
pronotum; mesoscutum without other impressions;
transscutal suture deeply incised; anterior scutellar
pits subround, medial carina separating pits narrow;
floor of anterior pits usually with one or two fine
carinae running from front to back, rarely entirely
smooth; lateral scutellar pits broad, deep, broadened
posteriorly; posterior margin of scutellum with a row
of 7—9, small, deep pits; dorsellum differentiated into
dorsal and posterior faces, with three keels, which
form three blunt projections dorsally; mesosoma
slightly compressed laterally, 1.11—-1.16x as high as
wide; pronotum evenly concave between spiracle and
pronotal shoulder; mesoscutum strongly convex;
acetabular carina evenly curved across mesosternum
and close behind fore coxae; sulcus just behind acetabu-
lar carina foveolate, with scattered pilosity; sternaulus
absent; two to four deep foveolae just above the mid
coxa and a weak flange immediately above the mid
coxa; medial propodeal keel anteriorly forming a right-
angled to acute, dorsally directed triangular projection.
Fore wing with basal vein strongly pigmented and
well defined; marginal vein 1.8—2.6x as long as wide;
stigmal vein short, about as long as width of marginal
vein; distal abscissa of radial sector moderately
pigmented. Metasoma: dorsally visible part of petiole
short, 1.3-1.5x as long as wide, longitudinal rugose,
superimposed with weak coriaceous sculpture; in dor-
sal view, gaster short oval with apex obtusely pointed;
metasomal syntergite (2+3) strongly convex, without
micropunctures posteriorly; in lateral view metasomal
tergites 7-8 almost perpendicular to long axis of gaster;
metasomal sternite 2 without micropunctures, with
two submedial tracts of hairs arising from fine punc-
tures. Colour: mandibles dark brown to black; palps
orange-yellow; radicle and basal 0.6—0.7 of scape
black; apex of scape and antennal segments 2-6 or-
ange-yellow, the more apical of these brownish;
segments 7—13 black; tegulae orange-yellow; basal
vein brown and marginal vein brown to dark brown;
legs including coxae orange-yellow except for telotarsi
and basal 0.6—0.7 of femora which are more or less
darkened. Body length: 2.2—2.9 mm, fore wing length
2.00—2.46 mm.

©. Differing from female as follows: Head in dorsal
view 1.13—1.17x as wide as long; temples weakly
contracted just behind eyes; head, in lateral view 1.08—
1.22x as high as long; eye height 1.0x as long as malar
space. Antenna (Fig. 9): scape mostly smooth,
coriaceous as base; segments 3-13 cylindrical; third
segment 2.1—2.5x as long as wide; segment 4 (Fig. 8)
cylindrical, very slightly expanded apically, 2.6—2.8x
as long as wide, about 1.2x as long as segment 3,
weakly emarginate and with a low carina extending

over basal 0.7; preapical segment 1.6—1.7x as long as
wide; apical segment no longer than and a little nar-
rower than preapical. Mesosoma (Fig. 17) in dorsal
view 1.46—1.52x as long as wide; notauli absent or
represented by weak traces or at most represented by
two small pits just in front of transscutal suture, ex-
tending over no more than posterior 0.2 of mesoscutum;
humeral sulcus broad, running all the way from the
transscutal suture to the posterior corner of the
pronotum; mesoscutum sometimes with very shallow
parapsidal impressions; posterior margin of scutellum
with a row of 8-10 indistinct pits, which tend to
coalesce; dorsellum less clearly differentiated into
dorsal and posterior faces and with lateral keels indis-
tinct; mesosoma depressed, 0.91—0.92x as high as
wide; acetabular carina bowed posteriorly, almost
reaching mid coxae; sternaulus present, defined
dorsally by a strong carina which extends right across
mesopleuron; mesopleuron with 3-4 rugae just above
mid coxa. Fore wing with basal vein moderately
pigmented; marginal vein 1.8—3.0x as long as wide;
distal abscissa of radial sector weakly pigmented,
almost spectral. Metasoma: dorsally visible part of
petiole 1.3—2.2x as long as wide; in dorsal view, gaster
oval, with apex rounded; in lateral view metasomal
tergites 7-8 at about 45° to long axis of gaster;
metasomal sternite 2 with medial area of
micropunctures in apical half; metasomal syntergite
(2+3) moderately convex with a small area of
micropunctures posteriorly. Colour: mandibles dark
brown to black; palps brown; antenna brown except
for scape which has the basal 0.5—0.7 dark brown and
apex orange to brown; tegulae brown; legs variable,
orange to dark brown with coxae, base of tibiae and
telotarsi darkened. Body length 2.1—3.7 mm; fore
wing length 2.03—3.11 mm.

MATERIAL EXAMINED.Czechia: ? , Bohemia, Sumava,
Hurka v PoSum, 20.vii.1954 (Hoffer);o%, Moravia,
Javorina, 800-900 m, 10.viii.1991, climax eufagion
for. (Masner) (CNC). Denmark: 9, north-east Zea-
land, Fergelunden, 55°51'N 12°02'E, 21.vi.1988
(Buhl) (DN);c’, north-east Zealand, Boserup Skov,
55°40'N 12°01'E, 12.vi.1988 (Buhl) (DN);c,
28.vi.1878 (Schlick) (ZM). Finland: 2, Pohjois-
Karjala, Tohmajarvi, 6908:660, 17.vii.1982 (Koponen)
(SEL). Great Britain: 9, Stirling, Mugdock,
11.vii.[pre-1910] (Cameron) (BMNH) (lectotype); 2,
W. Ross, Rassal National Nature Reserve, NG845432,
1-15.vi.1992, Malaise trap (Brown) (NMS); 9, West
Glamorgan, Crymlyn Bog, SS6995, 14.vi.1989
(Holmes) (DN); 39, Cheshire, Abbots Moss,
$J596680, 1-19.v1.1990, Malaise trap/swept (Notton)
(DN); 9%, Cambridgeshire, Chippenham Fen,
TL650693, 6.v11.1983, Malaise trap, carr/reed bed
(Field) (DN); 2 9 , Norfolk, Catfield, TG374204, 21.vi—
5.vil.1988, water trap (DN);o’, Surrey, Horsley,

134

14.vi.1930 (Nixon) (BMNH) (holotype);c’, Kent,
Eynsford, 7.vi.1930 (Nixon) (BMNH) (paratype).
Japan: 2 , Iwate Prefecture, Mount Hayachie, 400 m,
19-26.vii.1989 (Makihara & Sharkey) (CNC).
Norway:c', Rgyken, Kinnertangen, EIS 28 BQ,
vii.1993, Malaise trap (Hansen) (DN). Sweden: 9 ,
Nike, Orebro, 12.vii.1957 (Jansson) (Z1); 9 , Rérvik,
Lammhult, 15.vii.1986 (Buhl) (ZM);2, Uppsala,
Ericksburg, 14—-17.vii.1986, pan trap (Ronquist)
(CNC).

DISTRIBUTION. Widespread in north-west Europe
including: Czechia, Denmark, Finland, Great Britain,
Japan, Norway and Sweden (Fig. 22). Previously re-
corded from Britain (Kieffer, 1911; Nixon, 1980 as
Spilomicrus pelion; Notton, 1996 as S. near formosus)
and Algeria (Kieffer, 1911; cited in Jansson 1942). No
evidence was found to support the occurrence of S.
crassiclavis in Africa, the identity of the syntype
material Kieffer recorded from Algeria is uncertain
now that the type series has been found to be mixed.

BIOLOGY. Host unknown. Flight period June—July.
Recorded mainly from woodland, fen habitats and
other wetland.

COMMENTS. As well as the male syntype from
Bishopton (Masner, 1965) a female syntype of
Spilomicrus crassiclavis was discovered (BMNH) but
the Algerian syntypic material collected by du Buysson
is lost. Horn & Kahle (1937) note that the du Buysson
collection is at the Muséum National d’ Histoire
Naturelle, Paris but this material could not be found
there (Claire Villemant, pers. com.). Since the two
syntypes located are considered to belong to different
species the female was chosen as a lectotype. The
previous synonymy established by Nixon (1980) of S.
crassiclavis is not supported by this designation and so
S. crassiclavis is removed from synonymy with S.
integer. Also S. pelion is here synonymised with S.
crassiclavis. Character states associating the sexes
include the following: shorter antennal segments,
shorter notauli, less bulging temples and generally
smaller size than for S. formosus. A re-examination of
the material seen by Nixon (1980) showed that the
female he referred to as S. formosus was in fact S.
crassiclavis. Teodorescu (1970; 1986) figured the male
genitalia, however, owing to the taxonomic problems
outlined above, the identity of this material is doubt-
ful.

Spilomicrus formosus Jansson

(Figs 3-4, 10-12, 18, 20-21)

Spilomicrus formosus Jansson, 1942: 215.
LECTOTYPE @,, Sweden: Orebro, Oset (ZI), here

designated [examined]. Lectotype labels: Or. Oset,
10-8 39 A. J.; Typus [red label]; Spilomicrus

D.G. NOTTON

formosus Janss.,2; Zool. Mus. Lund Sweden,
Diapriidae Type No. 1411:1-7; ZML 1997.026.
Lectotype entire, clean and glued onto a card with
its wings raised.

DESCRIPTION. @ . Head (Fig. 3) in dorsal view 1.36—
1.42x as wide as long; temples weakly expanded to
parallel just behind eyes; dorsal part of occipital flange
widened medially; post-genal pilosity sparse, not form-
ing a defined cushion; head, in lateral view suboval,
1.04—1.20x as high as long; frons behind toruli weakly
convex, almost flat; eye sparsely hairy; eye height
1.7-2.2x as long as malar space; apex of medial
projection of clypeus triangular, pointed when seen
from in front; lateral margins of clypeus undefined;
malar sulcus a finely impressed line, becoming a little
deeper towards mandible. Antenna (Fig. 10) slender;
scape entirely smooth or at most weakly coriaceous at
extreme base of dorsum; segment 3 about 3.5x as long
as wide; segments 7-13 expanded to form a long,
cylindrical, 8-segmented club; segments 8—12 quad-
rate to slightly elongate; apical segment 1.7—1.8x as
long as the preapical and as wide or very slightly
narrower than it; apical segment without a ventral pit.
Mesosoma (Fig. 18) short, 1.25—1.26x as long as
wide; anteriorly mesoscutum usually without a pair of
short, submedial, parallel lines, rarely these are weakly
indicated; notauli sharply incised, deep, weakly sinuate,
narrowed anteriorly, extending over posterior 0.35—
0.45 of mesoscutum with no trace in front of this,
divergent anteriorly (Fig. 20); humeral sulcus running
all the way from the transscutal suture to posterior
corner of pronotum; mesoscutum without other im-
pressions; transscutal suture deeply incised; anterior
scutellar pits subround, medial carina separating pits
narrow; floor of anterior pits smooth; lateral scutellar
pits broad, deep, broadened posteriorly; posterior mar-
gin of scutellum with a row of 7—9 small, deep pits;
dorsellum differentiated into dorsal and posterior faces,
with three keels which project dorsally as three, sharp
points; mesosoma slightly depressed, 0.90—0.93x as
high as wide; pronotum evenly and shallowly concave
between spiracle and pronotal shoulder; mesoscutum
strongly convex; acetabular carina evenly curved
across mesosternum and close behind fore coxae;
sulcus just behind acetabular carina foveolate with
scattered pilosity; sternaulus absent; four to five deep
foveolae just above mid coxa and a broad flange
immediately above mid coxa; medial propodeal keel
anteriorly forming a right-angled to acute, dorsally
directed, triangular projection. Fore wing with basal
vein strongly pigmented and well defined; marginal
vein 2.0-2.3x as long as wide; stigmal vein short,
about as long as width of marginal vein; distal abscissa
of radial sector moderately pigmented. Metasoma:
dorsally visible part of petiole short, 1.2—1.4x as long
as wide, longitudinally rugose superimposed with weak

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 135

coriaceous sculpture; in dorsal view, gaster short oval,
with apex obtusely pointed; metasomal syntergite (2+3)
strongly convex, without micropunctures posteriorly;
in lateral view metasomal tergites 7—8 almost perpen-
dicular to long axis of gaster; metasomal sternite 2
without micropunctures, with two submedial tracts of
hairs arising from large punctures. Colour: mandibles
dark brown to black; palps orange-yellow; antennal
segments 1—5 orange-yellow except for the radicle
which is black and basal 0.2 of scape which is occa-
sionally black as well; segment 6 brown and segments
7-13 usually black; rarely the sixth orange-yellow and
seventh dark brown; tegulae transparent, orange-yel-
low; fore wing with basal vein brown and marginal
vein brown to dark brown; legs including coxae or-
ange-yellow. Body length 2.2-3.1 mm; fore wing
length 2.70-2.86 mm.

©. Differing from female as follows: head (Fig. 4)
in dorsal view 1.35—1.39x as wide as long; head, in
lateral view, 1.11—1.20x as high as long; eye height
2.3x malar space; eye without hairs. Antenna (Fig.
12): segments 3-13 cylindrical; third segment 3.5—
3.7x as long as wide; fourth segment 3.4—3.6x as long
as wide and as long as segment 3; segment 4 cylindri-
cal, with at most a weak basal emargination and a low,
weak carina in basal 0.3-0.4 (Fig. 11); preapical seg-
ment 2.0—2.1x as long as wide; apical segment 1.5—1.6x
as long as wide as the preapical and a little narrower
than it. Mesosoma in dorsal view 1.31—1.35x as long
as wide; notauli extending over posterior 0.3-0.4 of
mesoscutum; posterior margin of scutellum with a row
of 9-12 small deep pits; dorsellum with keels usually
projecting dorsally as sharp points but sometimes
rounded; mesosoma slightly depressed to slightly com-
pressed, 0.98-1.15x as high as wide. Fore wing:
marginal vein 1.7—2.2x as long as wide. Metasoma:
dorsally visible part of petiole 1.3-1.5x as long as
wide; gaster short oval, its apex rounded; in lateral
view metasomal tergites 7—8 at about 45° to long axis
of gaster. Colour: mandibles black to dark brown;
palps orange-yellow to brown; antennal colour vari-
able, in lightest individuals, scape orange yellow with
basal third black, pedicel orange-yellow and flagellum
brown, in darkest individuals, scape all black, pedicel
dark brown and flagellum black; tegulae transparent
orange-yellow; legs with ground colour varying from
orange-yellow to brown; telotarsi and basal 0.5—0.65
of femora blackened and coxae and trochanters brown.
Body length 2.8—3.4 mm; fore wing length 2.71—3.20
mm.

MATERIAL EXAMINED. Belgium:co’, Mont Rigi, stat.
se. Hautes Fagnes, 18—24.viti.1983, Université Liege
(CNC). Czechia: 29, Moravia, Lednice, env., 7—
9.vili.1991, yellow pan trap, riparian forest
(Masner);C’, Bohemia C., Revnice (Masner) (CNC).
Denmark: 29,0’, north Zealand, iv.1891, each

mounted with a pipunculid puparium (Schlick)
(ZM);o, v.1889; 49, 20%, iv.1890, each mounted
with a pipunculid puparium (Schlick) (ZM); Q , Zea-
land, Ruderhegen, v.1910, ex. Pipunculus sp.,
mounted with a pipunculid puparium (Kryger) (CNC).
Finland: 9 , Uusimaa, Espoo, 6669:373, 27.viii.1982
(Koponen) (SEL); 2 , Kemin Lappi, Kittila, 7469:409,
17.viii.1983 (Koponen) (DN);o, Etela-Savo,
Mantyharju, 6795:508, 1.viii.1981 (Koponen)
(SEL);o, Etela-Pohjanmaa, Toysa, 6948:336,
23.vii.1987 (Koponen) (SEL);c’, Helsinki, 28.vi—
11.vii.1989, Malaise trap (Goulet) (CNC). Germany:
30, Bayern, Oberstdorf, 12—29.viii.1936 (Nixon)
(BMNH). Great Britain: 20°, Elgin, Aviemore, 26.vii
& 5.viii.1946 (Harwood) (BMNH); 9, 20", W. Ross,
Rassal National Nature Reserve, NG845432, x.1991,
Malaise trap (Brown) (DN); 29, Cambridgeshire,
Chippenham Fen, TL6469, 17—24.vi.1985, Malaise
trap (Field) (DN),c’, Chippenham Fen, TL650693,
6-20.vii.1985, Malaise trap (Field) (DN);o,
Pembrokeshire, Brynberian Moor, SN106346,
20.vii.1987, pitfall trap, Juncus sp., soligenous flush
(Holmes) (DN);c&, Wiltshire, Savernake Forest,
$U21366708, 26.vii—16.viii.1990, Malaise trap
(DN);o, Oxfordshire, Weston Fen, SP526194, 22. vii—
20.viii. 1987, Malaise trap (DN);o", Surrey, Oxshott,
9.vili. 1930 (Nixon) (BMNH). Ireland:o’, Co. Wex-
ford, Benroe, Killybegs district, 18.viii.1933 (Stelfox)
(BMNH). Japan: 20°, Hokkaido, Tomuraushi area,
500 m, 13.vili.1996 (Masner) (CNC). Norway:
29,0, Berum, Kjaglidalen, EIS 28 AK, 27.vii-—
11.viii.1990, Malaise trap (Falck) (DN);c’, Rollag,
Varviken, EIS 35 BV, viii.1992, Malaise trap
(Sagvolden) (DN). Russia: 2, Moscow, Bitsa Park,
18.viil.1993 (Kolyada) (RAS);c', Moscow,
13.vii.1993 (Kolyada) (RAS). Slovakia:o’, {illeg-
ible}, Nizké Tatry, Kysla, 10.viii.1989 (Macek)
(CNC). Sweden:c’, Blekinge, R6deby Gagnekulla,
56°16'N 15°34 'E, 8.ix.1956 (Sundholm) (ZI); 2, 20°,
Dalarna, Al, 10 km south east of Leksand, 60°42'N,
15°02'E, 4.vii.1968 (Sundholm) (ZI); 3c, Dalarna,
Jarna, 60°33'N, 14°15'E, 3.viii.1968 (Sundholm) (ZI);
29, Hialsingland, Loos, pre-1942 (Sjéberg) (ZI)
(paralectotype); 5c, Jamtland, Fors, 63°01'N,
16°37'E, 7.viii.1964 (Sundholm) (ZI);:o,
Lulelappmark, Jokkmokk Messaure, 66°43'N,
20°20'E, 7.vili.1966 (Sundholm) (ZI);9, Narke,
Orebro, Oset, 59°17'N, 15°10'E, 10.viii.1939
(Jansson) (ZI) (lectotype);o", Narke, Orebro, Oset,
59°17'N, 15°10'E, 17.viii.1944 (Jansson) (ZI); @,
Narke, Orebro, O. Mark, 59°15'N, 15°12'E,
20.viii.1944 (Jansson) (Z1):o', Narke, Orebro,
22.viii.1937 (Jansson) (ZI) (paralectotype); 9 , Narke,
Orebro, 20.viii.1939 (Jansson) (ZI) (paralectotype); ? ,
Narke, Orebro, 16.viii.1941 (Jansson) (ZI)
(paralectotype); 29, 11co, Niarke, Orebro,
28.vii.1941—20.vii.1960 (Jansson) (ZI);c', Narke,

136

15.vii.1955 (Sundholm) (Z1);c’, Norrbotten, Edefors,
66°OS'N, 20°54'E, 31.vii.1958 (Sundholm) (ZI):o,
Norrbotten, Nedertornea, 65°50'N, 24°07'E,
24.v1i.1966 (Sundholm) (Z1);9, Nu., 15.viti.1955
(Jansson) (ZI);o", Oland, Gardby, 56°37'N, 16°05'E,
T.viii.1967 (Sundholm) (ZI);o, Ostergéthland,
Skedevi, 58°08'N, 15°42'E, 2.viii.1970 (Sundholm)
(ZI);0", Skane, Skaralid, 16.vii.1937 (Jansson) (ZI)
(paralectotype); 30°, Skane, Yngsj6, 55°57'N,
14°11'E, 23.vii.1964 (Sundholm) (ZI);c', Smaland,
Halltorp Varnands, 56°30'N, 16°07'E, 8.viii.1967
(Sundholm) (Z1);c', Smaland, 20.ii.1943 (Jansson)
(ZI); 5o&’, S6dermanland, Dalar6, Malmen, viii.1976
(BMNH);c", Sodermanland, Lanna, 10 km south of
Strangnas, 59°18'N, 16°54'E, 9.vii.1958 (Sundholm)
(ZI);0, 24.vii.1940 (ZI);2, 25.vii.1943 (ZI);c,
1.viii.1944 (Jansson) (ZI). Canada: °,c’, British Co-
lumbia, Hollyburn, 24.vii.1930 (Whittaker) (BMNH).
U.S.A.:0", Washington, Okanagan C., N. Waconde,
25.vii.1985 (Tinnamore) (BMNH);@, Virginia,
Shenandoah N. P., Big Meadow, 1,300 m, vii-—
vili.1987, Malaise trap, natural meadow (BRC Hym.
Team) (BMNH).

DISTRIBUTION. Widespread in north-west Europe,
extending well inside the Arctic circle (Fig. 21) in-
cluding: Belgium, Czechia, Denmark, Finland,
Germany, Great Britain, Ireland, Norway, Russia,
Slovakia and Sweden, also occurring in Japan, Canada
and the U.S.A. Previously recorded from Czechia
(Masner, 1991), Denmark (Masner, 1991), Finland
(Hellén, 1963; Masner, 1991), Germany (Nixon, 1980),
Great Britain (Nixon, 1980 — males only), Russia
(Hellén, 1963), Slovakia (Masner, 1991) and Sweden
(Masner, 1991).

BIOLOGY. Eleven Danish specimens of S. formosus
were seen each mounted with the puparium of a
pipunculid (Diptera). Of these Kryger’s specimen was
identified as Pipunculus sp., supporting Masner’s
(1991) observation that this genus is attacked in Eu-
rope. The developmental stage of host attacked is
unknown, although by analogy with other Spilomicrus,
it is likely that oviposition is into the young host
puparium. This would necessarily be after the
pipunculid has emerged from its own cicadellid
(Homoptera) host, and is therefore likely to be a case
of pseudohyperparasitism sensu Shaw & Askew
(1976). There are no host records yet from the Nearctic.
In Europe both sexes are found most often from July to
August, with a few females also in June, the dates of
reared Danish material, April to May, may not be
representative as they are much earlier and these wasps
may have been reared indoors in warmer conditions
than would be natural. In Europe S. formosus is re-
corded from a range of habitats including fen and other
wetland, forest and grassland. This is very similar to
its behaviour in north America where the flight period

D.G. NOTTON

is from July to October and it frequents mainly forest
habitats (Masner, 1991).

COMMENTS. Nixon(1980) misinterpreted the female
of S. formosus although he correctly identified the
male. The single female he mentioned belongs to S.
crassiclavis q. v. One male S. formosus from Orebro,
dated 28.vii.1941 (ZI), is erroneously labelled ‘typus’ ;
its date is not among those given in the original
description and so it was not considered a syntype.
Hellén’s (1963) material of this species is thought to
be at the Museum of Helsinki University but is appar-
ently lost.

Spilomicrus sanbornei Masner 1991
(Figs 5—6, 13-16, 23)

Spilomicrus sanbornei Masner 1991: 145. Holotype @ ,
Canada: Ontario, Thunder Bay (CNC) [not examined].

DESCRIPTION. 9. Head (Figs 5-6) in dorsal view
1.12—1.17x as long as wide; dorsal part of occipital
flange of even width; post-genal pilosity, forming only
a small cushion; temples parallel just behind eyes;
head in lateral view very rounded, subcircular 1.00—
1.05x as high as long; frons convex; vertex evenly
rounded; eye sparsely hairy, small, its height 1.6—-1.8x
as long as malar space; apex of medial projection of
clypeus rounded when seen from in front; malar sul-
cus, narrow near eye and distinctly widened towards
mandible, where it has the form of a step rather than a
groove, so that the anterior gena is depressed relative
to the posterior gena; lateral margin of clypeus defined
by a deep groove. Antenna (Fig. 13) robust: scape
smooth dorsally, weakly coriaceous ventrally; seg-
ment 3 about 1.7x as long as wide; segment 8 slightly
expanded and segments 9-13 strongly expanded to
form a short-fusiform 5- to sub-6-segmented club;
segments 8-12 weakly to strongly transverse; apical
segment 1.0—1.1x as long as the preapical and only
0.85x as wide as it; apical segment with a large,
shallow, ventral pit. Mesosoma (Fig. 16) in dorsal
view 1.45—1.54x as long as wide; notauli deeply im-
pressed over posterior 0.30-0.45 of mesoscutum,
indicated as a shallow trace for a short distance in front
of this; anteriorly mesoscutum with a pair of short,
submedial, parallel lines usually distinct but some-
times weak or absent; mesoscutum with a slight
impression in front of each hind corner of the meso-
scutum; humeral sulcus fine, running all the way from
the transscutal suture to the posterior corner of the
pronotum; transscutal suture moderately incised be-
coming weaker laterally; anterior scutellar pits deep,
oblique oval, the medial carina separating the two pits
usually narrow but sometimes a little wider; floor of
anterior pits with fine carinae running from front to
back; lateral scutellar pits usually deep and slightly

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS S77

broadened posteriorly but shallower in specimens with
a more depressed mesosoma; posterior margin of
scutellum with a row of 9-12 small pits; dorsellum
rounded, not differentiated into dorsal and posterior
faces, its keels weakly developed, the lateral keels
vestigial, the medial keel rounded, only weakly pro-
jecting dorsally; mesosoma strongly depressed,
0.73-0.77x as high as wide; pronotum with a shallow
but distinct fovea between spiracle and pronotal shoul-
der; mesoscutum only weakly convex, flattened
dorsally; sulcus just behind acetabular carina not fo-
veolate, almost glabrous; sternaulus absent; area above
middle coxa with only a few rugae; medial propodeal
keel low, hardly raised in front. Fore wing with basal
vein moderately pigmented; marginal vein 2.1x as
long as wide; stigmal vein very short, about half as
long as width of marginal vein; distal abscissa of radial
sector barely pigmented, almost spectral. Metasoma
with dorsally visible part of petiole short, 1.4x as long
as wide with 3-6 strong, almost parallel, longitudinal
keels; in dorsal view gaster elongate oval, its apex
pointed, approximately right-angled; metasomal
syntergite (2+3) weakly convex, with area of
micropunctures postero-medially; in lateral view,
metasomal tergites 7 to 8 at about 45° to long axis of
gaster; metasomal sternite 2 without micropunctures,
with sparse, scattered hairs arising from fine punc-
tures. Colour: antennal segments 1-8, palps and legs
including coxae orange yellow, telotarsi more or less
darkened; antennal club shading from brown to dark
brown or black; mandibles and tegulae brown. Body
length: 1.9-2.5 mm; fore wing length 1.51—1.80 mm.

©. Differing from female as follows: head in dorsal
view 1.18—1.25x as long as wide; temples parallel to
very slightly receding just behind eyes; head in lateral
view 1.03—1.07 as high as long; eye height 2.1 as long
as malar space. Antenna (Fig. 15) with flagellar seg-
ments cylindrical; flagellum slightly narrowed apically
and basally; third segment 2.2—2.5x as long as wide;
fourth segment 2.2—2.4x as long as wide and about
1.1x as long as segment 3; emargination of segment 4
moderately deep, curved, with a low carina over basal
0.45—0.5, ending in a small tooth (Fig. 14); preapical
segment 1.6—1.7x as long as wide; apical segment
1.3—-1.4x as long as preapical and barely narrower than
it; apical segment without a ventral pit. Mesosoma in
dorsal view 1.34—1.43x as long as wide; notauli deeply
impressed over posterior 0.25—0.45 of mesoscutum;
mesosoma strongly depressed, 0.76—0.79x as long as
high; marginal vein 1.8—2.2x as long as wide.
Metasoma with dorsally visible part of petiole short,
1.2-1.4x as long as wide; in dorsal view gaster with
apex rounded. Colour: antennal scape varying from
orange-yellow to brown, pedicel orange-yellow;
flagellum orange-yellow to yellow brown, usually
darkened apically; palps yellow; legs, including coxae,
orange-yellow with coxae, femoral clubs, tibial clubs

and telotarsi more or less darkened. Body length: 2.0—
2.3 mm; fore wing length 1.84-1.96 mm.

MATERIAL EXAMINED. Great Britain:o’, Cheshire,
Abbots Moss, SJ596680, 1—19.6.1990, Malaise trap
(Notton) (NMS);2,0, Cheshire, Abbots Moss,
S$J596680, 19.6-10.7.1990, Malaise trap (Notton)
(DN). Canada: 9 , Ontario, Guelph, 8—24.vi.1981, pan
trap (Barber) (CNC) (paratype);o", Ontario, Rondeau
Prov. Pk, 9—26.vi.1980, interception trap (Goulet)
(CNC) (paratype);o’, Quebec, Gatineau Pk, 28.vi—
5.vii.1983, beaver pond, interception trap (Denis)
(CNC) (paratype);2, Quebec, Old Chelsea, 5—
15.vi.1987, NCC woodpile, pan trap (Masner) (CNC)
(paratype);o", Québec, Old Chelsea, 5—15.vi.1987,
pan trap in dead wood pile (Masner) (BMNH)
(paratype).

DISTRIBUTION. Newly recorded from Britain and
the Palaearctic (Fig. 23). Previously recorded from the
Nearctic including Canada and the United States
(Masner, 1991).

BIOLOGY. Host unknown. In both North America
and Britain the flight period is June to July and forest
habitats are frequented.

COMMENTS. Spilomicrus sanbornei is superficially
similar in appearance to S. abnormis but easily distin-
guished from it by the presence of a malar sulcus, lack
of pubescence at the base of metasomal syntergite
(243). British material of S. sanbornei does not en-
compass the full range of variation shown by Nearctic
material in the length of the notauli, sculpture of the
floor of the basal scutellar pits and body length. This is
probably because few British specimens were exam-
ined. British material also has the anterior parallel
lines of the mesoscutum either absent or less well
developed than in Nearctic specimens and the
mesosoma very slightly more depressed. This is not
considered significant because such variation in these
features of the mesosoma is common in diapriines,
particularly in groups where wing reduction is frequent.

REFERENCES

Buhl, P. N. 1994. Fortegnelse over Danmarks proctotruper (Hy-
menoptera, Proctotrupoidea s. 1.). Entomologiske Meddelelser,
62(1): 13-24.

Coe, R. L. 1966. Diptera, Pipunculidae. Handbooks for the Identifi-
cation of the British Insects, X(2c): 1-83.

Disney, R.H.L. 1994. Scuttle flies: The Phoridae. Chapman & Hall,
London. Pp xii + 467.

Hoffmeister, T. 1989. Biologie und Wirtskreis parasitischer
Hautfliigler der Familie Diapriidae. Natur und Museum, 119(10):
327-334.

Horn, W. & Kahle, I. 1937. Uber entomologische Sammlungen
Entomologen und Entomo-Museologie. Entomologische Beihefte,
2-5: vi + 536 pp.

138

Jansson, A. 1942. Neue Proctotrupiden aus Schweden. I.
Entomologisk Tidskrift, 1942: 210-216.

Johnson, N. 1992. Catalog of World Proctotrupoidea excluding
Platygastridae. Memoirs of the American Entomological Institute,
51: 1-825.

Kieffer, J. J. 1911. Proctotrypidae, Diapriinae. Jn: André, E. (Ed.)
Species des Hyménopteéres d’Europe et d’Algerie, 10: 753-912.
Kieffer, J. J. 1916. Diapriidae. Das Tierreich, 44. Walter de Gruyter

& Co., Berlin. 617 pp.

Kozlov, M. A. 1978. [Proctotrupoidea except Serphitidae]. Pp 538—
664 in: Medvedev, G. S. (Ed.) Opredeliteli Nasekomykh Evropeiskoi
Chasti S. S. S. R. [Determination of the Insects of the European
portion of the U. S. S. R.], 3(2): 1-758. Leningrad.

Lundbeck, W. 1922. Diptera Danica, 6 (Pipunculidae, Phoridae): 1—
447. Copenhagen.

Masner, L. 1991. Revision of Spilomicrus Westwood in America
North of Mexico (Hymenoptera: Proctotrupoidea, Diapriidae).
Canadian Entomologist, 123: 107-177.

Mason, W. R. M. 1986. Standard drawing conventions and defini-
tions for venational and other features of wings of Hymenoptera.

D.G. NOTTON

Proceedings of the Entomological Society of Washington, 88: 1-7.

Nixon, G. E. J. 1980. Diapriidae (Diapriinae). Hymenoptera,
Proctotrupoidea. Handbooks for the Identification of the British
Insects, VY(3di): 1-55.

Notton, D. G. 1996. Diapriid Wasps (Hym., Proctotrupoidea) from
Abbots Moss, Cheshire. Lancashire and Cheshire Fauna Society,
95: 23-24.

Shaw, M. R. & Askew, R. R. 1976. Ichneumonoidea (Hymenoptera)
parasitic on leaf-mining insects of the orders Lepidoptera, Hy-
menoptera and Coleoptera. Ecological Entomology, 1: 127-133.

Teodorescu, I. 1970. Contributii noi la studiul proctotrupoideelor
(Hymenoptera — Proctotrupoidea). Studii si cercetdri de Biologie,
Seria Zoologie, 22(3): 189-196.

Teodorescu, I. 1986. Contributii la cunoasterea armaturii genitale
mascule la Ceraphronoidea si Proctotrupoidea. Analele Universitatii
Bucuresti (Biologie), 35: 3-15.

Westwood, J. O. 1832. Descriptions of several new British forms
amongst the parasitic hymenopterous insects. London and Edin-
burgh Philosophical Magazine and Journal of Sciences, 1:
127-129.

INDEX

Synonyms and misidentifications are in italics; principle references are in bold.

antennatus 129, 130

fumipennis 131
Basalys 131

Bibionidae 130 hemipterus 130

crassiclavis 129, 132 Idiotypa 131

Diapriidae 129 integer 129
Diapriinae 129
Drosophila 130
Drosophilidae 130

Juncus 135

Muscidae 130

erythropus 131
Dawe pelion 129, 132

Phoridae 130
Pipunculidae 130

formosus 129, 131, 132, 134
formosus 132

formosus species group 129, 130, 131, 132

Pipunculus 131, 136

Sanbornei 129, 131, 132, 132
Sciomyzidae 130
Spilomicrini 131

Spilomicrus 129, 130
Staphylinidae 130
stigmaticalis 129, 130
stigmaticalis species group 130
Syrphidae 130

Tachinidae 130
Tephritidae 130
tuberculatus 131

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 139

ma

Fig. 1. Spilomicrus stigmaticalis — head, anterior,O’ . Fig. 2. S. crassiclavis — head, anterior, 9 . Figs 3-4. S. formosus — 3, head,
anterior, 2 ; 4, head, dorsal,O’. F igs 5-6. S. sanbornei — 5, head, anterior, paratype @ ; 6, head, lateral, paratype 2

140 D.G. NOTTON

a
NO

10

=X
(Se)
=
O1

—
—k

SNE an gale OR TEE LL!
ara
te)

Figs 7-9. Spilomicrus crassiclavis — 7, left antenna, posterior, Q ; 8, left antenna, fourth segment, posterior, ©’; 9, left antenna, posterior’.
Figs 10-12. S. formosus — 10, left antenna, posterior Q ; 11, left antenna, fourth segment, posterior, © ; 12, left antenna, posterior, 0’.

Figs 13-15. S. sanbornei -13, left antenna, posterior, paratype @ ; 14, left antenna, fourth segment, posterior, paratype’; 15, left
antenna, posterior, paratype O’.

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 141

Fig. 16. Spilomicrus sanbornei — mesosoma, lateral, paratype Q . Fig. 17. S. crassiclavis — mesosoma, lateral, 0’.

142 D.G. NOTTON

Figs 18 & 20. Spilomicrus formosus — 18, mesosoma, lateral, Q ; 20, mesoscutum, dorsal, 9 . Fig. 19. S. crassiclavis — mesoscutum,
dorsal, @ .

REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 143

Fig. 21. Spilomicrus formosus — North-west European distribution.

144 D.G. NOTTON

Fig. 22. Spilomicrus crassiclavis — distribution. Fig. 23. S$. sanbornei — North-west European distribution.

Bull. nat. Hist. Mus. Lond. (Ent.) 68(2): 145-193 Issued 25 November 1999

A review of the soldierless African
termite genus Amicotermes Sands
1972 (Isoptera, Termitidae,
Apicotermitinae)

W.A. SANDS

Department of Entomology, The Natural History Museum, Cromwell Road, London
SW7 SBD, UK.

CONTENTS
(3 STACGY DIST GP apse: iy eee OR Oy cP AU ob Oana or rt SP eee OP rsa ie emenen are Reeree here 145
ICS ERC CTT CEM OEN eee ase ences AEE cca as asco Se cee ed cnepledhat dan cunapRevisaccdtenpiastondnsdisTrensenedeacde 146
INVERN OS fees cixc nse ce atde na sec nd va cuntadgn canes casaza dacarsvsdecksvuach inna aanisesradnscntesancansneaddenestsbaanitsanusiatecadda. 146
RB OMIMBCHISTOULCUANAC ENS SC Oeste cca tancnse sed eces aco ah aean ca ecudas fa atteransncatanadexe asae sans dafacsimauesc ous 147
EAT AGSMCES CLIP MOUS WLLLS CALC iter cecnacessescesee sencstestccstoceeccanet csartordtcsaceavesarsereracevccstessares= 147
COnGenSed catamiatnin, WIth OLE OTOUP lore. ce-cas<ssetceuacexsaveeateestucsher sonar deaceavanevaeeenes-casscescevecseo 150
PAUTEG COLE LTITESS AUS MUON acer tantrescneet car esecns ets cv cate vactnenavendeeaestcetaurtvavasecevucvdstareuhvevesteKersscutsiss 150
IRE WALONLLLE IS PHEC IES OM AIMICOLEN IME Saree crcere neces csccuceuecttauresattcentatensceturastercitstensstaesccw sminctrrtssereas 151
IDESCHIPMOMS!ON SPECIES =.c.ccecceraceceesenen cates eee iss}
PATICOLCTINGS GUIOTMYSIUS SPr ile. cscvestssevassasensxecesssesvevssseaestnadcoces ¥eseesvesaccorsetcoresssuevostueteessss* 153
NTI GCOLETINES GAMEFUNETSIS! SP) aLlstediccsteccesissssvevasaustiacssvassseebece.seuceesvstesecssssaventevdsratveotexsceeesee 154
PATHLCOLEHINESKGON ZOCTISUSHSP Us. ck sarees cats eccnisncetoevucscotuecersereerdaceceenesuotercucctertiectevetsnireetaceys 156
PATI COLENULES GUEST CEILS) SJ gillian averse ors dette vexdsie~sakeveaetdencdeavses: vaxattetiesnsvedusnrsespessccereavecsaetcese 157
PNTTIIC OLE TINE SALD OR ASP NN G2 tec ste eae 2 toa ete? onan usu av eh nce men aetccucanet ar eeteaeeeninranee te cnsarartenwedennves 158
PATIIGOLCN INES GALENUN SAN OS ses sys sacussvssra ccisvascasss seleace nereudeenctaatet tisestdeete an aditeaaoacrtewiees 159
PATFILG OLCTINIES\ OP CLSLEMUDELDSES) 20MM var as aces nen in nah nso Pete ee tat Pee cee eet eee pea aces 160
PAUTILGC OLE IIE SN UVOTENESUS SPs less cvscctrauas caseos gun saneos sane ctothy stds wneannesWeences sae teaeaeck sageseis Res exeb 161
ATRL COLETING SIMAYOMDEL' SP tlre tes ceeseteecen ev sac suse Sncaosihe ete sea eeauneenes waeaseaeitawar ae oseth sacs Lass ave 162
VAVATEG OLETITICS ITLD CUITG OEMS IS SP) AUN ace ss ances scas aes tae auee can SCR a aa cetera ec Peon Se aE 163
PSTITLG OLE TIES TPIAULLSTDLIRTES SP) SDN. g vcza ascents cert ss fnkavsox dees astiversnsbapssieeste} coucuscdsesnceseaytaee¥sns stuustasosvie 164
PATITLC OLE IIE SI SDE GILOTLS GUOs Lc praae tise sete iced seo Saeed sc aya dade Ue Wee diene dhe ce tats cvodecsseccests Sect -Seesncees 165
HEME MeEANG VO PCMELIC ANALY SISter ceete ye seec traces seccnie vac de dt vaccdn vances aactesaconcsbeencaunes Suedpenst iodine 165
PSCEMO WEG SOMACIUS 2. u.sceseesceseupe st ecse coerce dacesesssscenvsaesaents sandvishccsucent snaencsctsusateaienandussscantsesancatés 167
IRE HEL ODIC GS ate re Ne res nate eee ONT PTO cocoa sond Selene Dew nev eck vse JoGe bias suovan caktrenaieasecbvaxanes 167
PS TDSC ING Near aster enue se cstisacn sar au ten cguns tuactcaavavievséadesuecucevessesucesdsst cssvi vas aha its aaesueaeesiefesactst svestss 168

SYNOPSIS. The termite genus Amicotermes is a member of the soldierless Apicotermitinae
(Isoptera, Termitidae), recognised as important constituents of the soil fauna in tropical Africa.
Before the current work was begun, 60 species belonging to 16 genera had been recorded, among
them the single species from the southern Congo for which the genus was named. New intensive
ecological transect sampling in Cameroon and Congo (Brazzaville) by members of the Termite
Research Group at The Natural History Museum has greatly increased the observed species
diversity of the soil. Eleven new species of Amicotermes are keyed and fully described here from
a database compiled in DELTA format in PANKEY, which also allows the matrix to be converted
to HENNIG86 input or similarity coefficients for phenetic analysis. Phenetic and cladistic
analyses of the relationships between the species are provided. All species are illustrated with line
drawings and photographic plates.

© The Natural History Museum, 1999

146

INTRODUCTION

It has been noted before that the soldierless
Apicotermitinae (Isoptera, Termitidae) are very im-
portant constituents of the soil fauna in tropical Africa,
being soil or root feeders, or, some of them, active at
the soil/wood/litter interface. Soil sampling has shown
them to be both abundant and diverse in forest and
savanna ecosystems, accounting for a significant part
of the live biomass of soil organisms and responsible
for a major proportion of soil movement and aeration,
contributing also to water penetration (Sands, 1965,
Eggleton et al., 1996). Before the current study began,
a total of 60 species of soldierless termites belonging
to 16 genera had been recorded from the African
continent by Sands (1972). A few more genera and
species were known from the worker castes alone, but
were not described at that time because their charac-
ters were not considered to provide as reliable
identification as those of the imagos. Since then, fur-
ther studies of the distinguishing features of the worker
castes (Sands, 1992, 1995, 1998) have shown that they
can be identified as readily as the imagos, in many
cases more easily, and that it is no longer logical to
abstain from describing species of soldierless termites
recognisable on the basis of their workers alone.

Among the genera previously known, Amicotermes
was represented by a single species from the southern
Congo. The new species described here were, with
two exceptions, all collected from rain forest, or de-
graded derivatives of it, by the members of the termite
group at The Natural History Museum, either as part of
the NERC-funded TIGER project team in different
selected habitats in Cameroon in the Mbamayo forest
reserve, Or, using a similar transect sampling tech-
nique, in the Republic of Congo (Brazzaville) at
Mayombe. The purpose of the former project was to
examine the effects of different levels of forest degen-
eration and regeneration, resulting from cultivation
and selective logging, on the biodiversity of the soil
fauna and its relationship to the carbon cycle (Eggleton
et al., 1996).

When the first collections from the Mbalmayo area
of Cameroon came to be sorted out, it quickly became
evident that the diversity and abundance of termite
species in the soil, especially the soil-feeders, of which
about half were soldierless Apicotermitinae, was far
greater than had been anticipated for this project or
had been recorded previously from any ecological
sampling, even in rain forests elsewhere. Abundances
up to 10,488m”~ and live weight biomass up to 123 gm~
were recorded. In a significant proportion of the gen-
era examined, new species were found, and indeed,
genera new to science were also present. All of these
will have to be described and properly enumerated in
due course, but the work presented here covers, as a

W.A.SANDS

first instalment, the 11 new species recognised in
Amicotermes, bringing the total of recorded species
for the genus up to 12. This genus was chosen to be
examined first because, in an otherwise very difficult
group, it has an enteric valve armed with spines and
spicules which offer useful taxonomic characters that
define species relatively well. Some other genera that
remain to be worked out lack this feature and will take
much longer to resolve. In the published account of the
ecological sampling (Eggleton et al. 1996) morpho-
species that were thought to be new were designated
numerically, there being three such in the genus
Amicotermes. Further, more careful examination of
the material has doubled this number for the Cameroon,
and four more were found in the Congo. The last new
species was recognised in material acquired earlier
from the Upper Guinean Forest in the Ivory Coast.
Because of the unevenness of the sampling and knowl-
edge of the ecology of the group in the rest of the
Afrotropical zoogeographical region, it is evident that
if the entire continent were to be covered in the same
detail, the number of species and genera would be
multiplied several times over. However, a diligent
search through the unidentified accessions from all
parts of tropical Africa at The Natural History Mu-
seum, including many samples from soil pits collected
in the course of earlier ecological studies in Guinean
savannah in Nigeria (Sands, 1965a), have failed to
produce any further records of Amicotermes. It is
therefore probably safe to conclude that it is confined
to rain forest and this study amounts to a new revision
of a single soldierless genus.

METHODS

In order to speed up the recording of taxonomic data
and facilitate its processing by computer, the character
set was coded and stored in PANKEY (Pankhurst
1986) DELTA [DEscriptive Language for TAxonomy]
(Dallwitz & Paine, 1986, Partridge et al., 1986) for-
mat. In this state, it may be converted by several sets of
programmes into natural language descriptions, di-
chotomous printed keys, analysed by cladistic or
phenetic procedures, or converted into ‘expert sys-
tems’, that is, interactive computerised polyclaves.
Other recently published works (Pankhurst, 1991,
Sands, 1992, 1998) go into the details of how this is
done. The key and descriptions that follow were pro-
duced in this way, as were the comparisons between
related species. In order to incorporate the new species
into the existing genus, the data from Sands (1972)
were coded and added by means of the dedicated
editor, DEDIT, to the DELTA file, a copy of which is
included as an appendix to illustrate the format, be-
cause in the author’s view, too few taxonomists are

REVIEW OF AMICOTERMES

either aware of its value as a tool, or of its relative
simplicity in use once the requirements are under-
stood. The full data matrix itself with ranges, means
and standard deviations of measurements has been
converted to the key and descriptions and is therefore
not repeated here, but it can be supplied on request to
those interested as a copy of the entire DELTA file on
receipt of a single 3.5 inch high density diskette.

COMMENTS ON CHARACTERS
USED

Some of the characters used require further explana-
tion. Apart from A. dibogi, the genus is remarkably
homogeneous in its external features, and a great deal
of emphasis is therefore placed on the enteric valve
armature and on morphometrics. However, it was
thought desirable in constructing a key to give sum-
mary descriptions of the salient externally observable
characters as well as measurements, to supplement the
usually distinctive valve structures, because the latter
require quite a difficult dissection and slide mount. An
annotated character list is given below, followed by a
condensed version of the data matrix as used in the
HENNIG86 analysis, since it includes Acholotermes,
the sister genus of Amicotermes, as an outgroup.

Character descriptions, with states

Some of the characters require quite difficult dissec-
tions to display them properly. Full descriptions of the
techniques involved are published elsewhere (Sands
1998), but summary accounts are given below where
appropriate.

Characters 1-5: The system of coding colour of body
parts as an ordered multi-state character has been used
since 1965. Only the first five grades (Sands 1965) are
required here for the relatively lightly sclerotised
worker caste, being re-numbered for each character.

1. Head capsule colour: 1. yellow-white 2. pale yel-
low 3. yellow 4. pale yellow-brown.

2. Antennal flagellum colour of darkest part 1. yel-
low-white 2. pale yellow 3. yellow 4. pale
yellow-brown.

3. Pronotum colour 1. yellow-white 2. pale yellow 3.
yellow.

4. Legs colour 1. yellow-white 2. pale yellow 3.
yellow.

5. Abdominal tergites colour 1. hyaline 2. yellow-
white 3. pale yellow.

Characters 6-10: Head capsule shape is evaluated in
facial view. Pilosity is best viewed from below and in
front against a lit background; sparse is intended to

147

mean when the setae are spaced further apart than the
length of the longest; two lengths of setae are usually
evident, the shorter, general pilosity being randomly
distributed while the longer, emergent setae are com-
monly paired on each side of the mid-line. The term
‘fontanelle’ is used for convenience although workers
do not have a true fontanelle, which is the opening of
the frontal gland in soldiers and imagos; some species
have a pale spot in this position. The degree of infla-
tion of the postclypeus is estimated partly by its
dimensions, but also in profile by the regularity of
convexity of its outline.

6. Head capsule shape 1. circular 2. short oval near
circular 3. oval.

7. Head capsule setae |. sparse 2. numerous but not
dense.

8. Head capsule emergent, longest setae 1. randomly
scattered 2. regularly and symmetrically arranged.

9. Fontanelle: 1. present as pale spot or small de-
pression 2. absent.

10. Postclypeus contour: 1. clearly inflated 2. strongly

inflated, bulging, length more than half width at
ginglymi.

Characters 11 & 12: The presence of prominent setae
arranged in a line or group on the front surface of the
fore coxa may be difficult to detect because in this
genus they are not particularly “spine-like’. If viewed
in profile against a lit background they can usually be
detected as more robust than normal. Fore tibial apical
spurs always number three in this genus with the third,
outer spur subject to size variation.

11. Fore coxae 1. without prominent larger setae on
anterior surface 2. with one to three prominent
larger setae on anterior surface 3. with four or
more prominent larger setae arranged in line or
group on anterior surface.

12. Fore tibial apical spurs numbering three 1. third
well developed, at least half the size of the others
2. third much smaller than the other two.

Characters 13-17: Mandible characters including
measurements require the cutting edges to be secured
transverse, i.e. at right angles, to the optic axis of the
lens system containing the graticule. This is particu-
larly important in assessing the visibility and
alignment of the left fourth marginal tooth. In order to
see these characters clearly, a simple dissection by
slitting the adductor apodemes is needed, allowing
the mandibles to swing open on their articulations. It
is undesirable to detach them from the specimen,
since their tetrahedral shape makes them impossible
to mount on a slide or secure in a dish in the required
position. The dissection is best done with the speci-
men secured on its back with the finest grade of

148

stainless steel headless entomological points. The
point of a fine scalpel or needle is inserted into the
mouth of the specimen, to push aside the labium and
hypopharynx. The adductor apodemes are usually
visible at the base of the mandibles and can be cut
with a single stroke of a fine blade in each direction.
A gentle push will then swing the mandibles out-
wards in turn, and the labrum can be deflected between
their separated bases to secure them in position; if
necessary the labrum can be pushed out again to
allow the mandibles to return to a natural position.
The designations of the marginal teeth of the left
mandible need further explanation. The first is that
immediately proximal to the apical tooth, but the
second is incorporated into the cutting edge behind it,
forming its curved inner end; the third marginal is
separated from the first-plus-second by a distinct
notch, while the fourth appears out of alignment with
the other marginals, being visible in the gap between
the third marginal and the molar prominence; its proxi-
mal end, which forms the point of the tooth, is
sometimes hidden behind the molar prominence. There
are only two marginal teeth on the right mandible in
this genus.

13. Left mandible, first marginal tooth 1. with ante-
rior edge distinctly longer than posterior 2.
approximately equilateral.

14. Left third marginal tooth with anterior edge 1.
shorter than that of first 2. equal in length to that
of first 3. longer than that of first.

15. Left fourth marginal tooth in front view 1. with
proximal end clear of molar prominence 2. just
reaches side of molar prominence 3. with proxi-
mal end hidden behind molar prominence.

16. Anterior margin of right first marginal tooth 1.
longer than that of second 2. equal in length to
that of second 3. shorter than that of second.

17. Right first marginal tooth with exposed posterior
edge 1. equal to that of second marginal 2. longer
than that of second marginal.

Characters 18—22: The configuration of the parts of
the alimentary canal as seen through the unopened
body wall has been used and illustrated extensively
(Sands, 1972, 1995, 1998) inidentifying worker castes.
The most important features in Amicotermes are the
position of the enteric valve seating and the length and
width of the first section of the proctodeum (P1). The
lines of the heart dorsally and the nerve cord ventrally,
although sometimes hard to see, are used to mark the
positions of features.

18. Junction of mesenteron and proctodeum 1. start-
ing to left of nerve cord in ventral view 2. starting
beneath nerve cord in ventral view 3. starting to
right of nerve cord in ventral view.

W.A. SANDS

19. Proctodeal first segment 1. shorter, length up to
eight times proximal width 2. very long, over
eight times proximal width.

20. Proctodeal first segment beyond junction with
mesenteron |. tubular or conical throughout its
length 2. dilated to about twice its basal width
throughout length.

21. Lateral displacement of enteric valve in unopened
abdomen 1. to left of heart in dorsal view 2. to
right of heart in dorsal or to left of nerve cord in
ventral view.

22. Enteric valve ‘seating’ third lobe 1. smaller than
outer pair 2. approximately equal in size to outer
pair.

Characters 23-45: The enteric valve is variably
sclerotised and armed within its lumen with scales,
spines or spicules. In order to display these a dissec-
tion is required. Having opened the abdomen by pulling
away a section of the body wall with forceps, it is
possible simply to pull on the first section of the
proctodeum near its distal end to extract the enteric
valve from the third (the valve being the second). The
valve then needs to be slit longitudinally by inserting
into the lumen a micro-scalpel, cleaned, separated
where possible from the muscle coats that surround it,
and mounted on a microscope slide. The choice of
mounting medium is dictated by the fragility of the
wisp of tissue that makes up the enteric valve. This
prevents the usual dehydration procedures required by
most permanent mounts, and necessitates the use of a
direct, relaxant temporary medium such as a variant of
‘Swan’s Berlese’.

Within the enteric valve there are six longitudinal
ridges (the ‘bourrelets’ of Grasse and Noirot (1954))
which carry the main sclerotisations, spines and
spicules. The system of numbering the ridges adopted
here is that of Sands (1972), in which the single ridge
nearest to the front of the specimen, or innermost to the
coiling of the gut, is designated ‘position number one’;
the pair on each side of it are numbered ‘position two’,
the pair beyond them ‘position three’ and the distal
singleton, ‘position four’. This is because these
positions describe the most common independent
dimensions of variation of symmetry, sclerotisation
and armature. Because they vary independantly, they
have to be described separately for each character and
its states. This has resulted in a set of characters that
appear repetitive for taxa with hexaradially symmetri-
cal valves; however, to do otherwise would mean
losing a large proportion of the information content of
this important structure. The characters referring to
the positions of the largest spines as proximal, mesal
or distal, are not intended to imply that there are no
spines on the rest of the ridges, merely that where
present, they are smaller than the most elongate and
conspicuous spines. The ridges also carry subsidiary

REVIEW OF AMICOTERMES

armature of scales, spines or spicules. Between and
beyond the ridges the wall of the valve may also bear
scales, sometimes armed with small spicules.

23. Enteric valve ridges, shape 1. more or less taper-
ing ellipsoid to base of apical spines 2. distinctly
‘waisted’ at base of apical spines, then wider.

24. Enteric valve ridge in position one 1. sclerotised
at distal end, pale brown or yellow-brown 2.
weakly sclerotised distally, pale yellow or yellow
3. unsclerotised or colourless throughout.

25, 26, 27, as 24 but referring respectively to positions
two, three and four.

28. Main armature of enteric valve ridge in position
one 1. without large apical spines (retained to allow
forthe outgroup, but does notapply toAmicotermes)
2. with large backwardly directed erect conical
spines 3. with very elongated tapered spines.

29, 30, 31 as 28 but referring respectively to positions
two, three and four.

32. Largest apical spines on ridge one 1. numbering
1-9 2. numbering 10-14 3. numbering 15-20 4.
numbering over 20.

33, 34, 35 as 32 but referring respectively to positions
two, three and four.

36. Longest spines of main armature 1. straight and
evenly tapered 2. curved but still evenly tapered.

37. Longest apical spines |. longer than least width of
ridges in position three at base of spines 2. shorter
than or equal to least width of ridges in position
three.

38. Subsidiary armature on one or more of enteric
valve ridges mainly 1. reticulate 2. with back-
wardly directed scales 3. with backwardly directed
scales fringed with small spines or spicules 4.
with small single backwardly directed spines on
scales 5. with prominent spines different from or
smaller than main armature.

39. Subsidiary armature of enteric valve ridges 1.
with scales or reticulations well defined, distinct
2. with scales or reticulations indistinct, outlines
vague or incomplete.

40. Subsidiary armature of one or more of enteric
valve ridges 1. prominent, covering entire ridge
surface 2. scattered, somewhat sparse 3. very
sparse, few scales fringed with spines or spicules.

41. Subsidiary armature on one or more enteric valve
ridges 1. more or less uniform throughout length
of ridge 2. graduated, spines or spicules becom-
ing longer towards distal end of ridge.

42. Symmetry of enteric valve armature 1. more or
less hexa-radial apart from slight differences in
ridge size 2. more or less tri-radial, alternate
ridges reduced 3. bilateral due to marked differ-
ences in spine and ridge size.

43. Membranous wall of enteric valve between and
beyond ridges 1. smooth 2. scaly.

149

44. Membranous wall of enteric valve between and
beyond ridges 1. armed with minute spines or
spicules 2. armed with short spines or spicules.

45. Spines or spicules on membranous wall 1. very
sparse, scattered 2. numerous but not dense, scat-
tered, fringing scales proximally 3. dense distally,
fringing scales proximally.

Characters 46-60: Some of the listed ratios may
appear unusual in the comparisons they make; they
result from selecting measurements that carried large
contrasting weighting coefficients in principal compo-
nent and canonical variates analyses. Such procedures
are now so much the routine that no extended treat-
ment is given here. The rationale for the use of complex
ratios rather than the actual discriminant functions is
explained in Sands (1972) which also lists the set of
measurements used as standard in the soldierless ter-
mites (and see also Fig. 1). The measurements are
given here to three places of decimals because they
were recorded automatically from an electronic filar
micrometer and processed simultaneously; it seemed
less artificial to give them as recorded rather than to
round them to fewer places arbritrarily. In practice it
has been found that at this level with this equipment
measurements are repeatable within about 3%. The
measurements used are mostly self explanatory being
the maximum distance between identifiable fixed
points. Those that need further explanation are the
following: Postclypeus width, taken between the inner
curves of the ginglymi of the mandibles, which are
always recognisable as strongly sclerotised articula-
tions at the corners of the postclypeus; Left mandible,
La, distance between the points of the apical and first
marginal teeth; L1, distance between the points of the
first marginal tooth and the third marginal, because the
second is often reduced or obsolete; Lm, distance
between the point of the third marginal and the inter-
section of the fourth, subsidiary marginal (some authers
call this the ‘molar tooth’ but its function is marginal)
with the molar prominence, whether this is a notch or
the point where the fourth marginal passes behind the
prominence; Right mandible, Ra, as left; R1, distance
between the points of the first and second marginal
teeth; Rm, distance between point of second marginal
and the intersection of its hind margin with the molar
plate; the front edge of this tooth is also measured, in
order to provide a ‘second marginal index’ to indicate
its degree of reduction. It is important when taking
measurements to ensure that the specimen is held
exactly transverse to the optic axis of the lens tube of
the eyepiece containing the graticule.

46. Head capsule width (range, mean, SD) mm.
47. Postclypeus width (range, mean, SD) mm.
48. Postclypeus length (range, mean, SD) mm.
49. Left mandible index (range, mean).

50. Right mandible index (range, mean).

51. Right second marginal index (range, mean).

52. Right first to second marginal index (range, mean).

53. Pronotum width (range, mean, SD) mm.

54. Pronotum length (range, mean, SD) mm.

55. Hind tibia length (range, mean, SD) mm.

56. Fore tibia length (range, mean, SD) mm.

57. Fore tibia thickness (range, mean, SD) mm.

58. Foretibialindex (length over width) (range, mean).

59. Leftmandible, complex ratio La divided by L1.Lm
(range, mean).

60. Right mandible, complex ratio Ra divided by
R1.Rm (range, mean).

Condensed data matrix with outgroup

The condensed data matrix, outgroup included, is
shown in Table 1. The programmes used to process
descriptions and keys were DESCRIP3 and KCONP
with KCONI, the interactive key construction pro-
gramme and KPRINT from the PANKEY set
(Pankhurst, 1986, 1991). An MS-DOS binary file of
the data to run with PANKEY ONLIN7 that will
provide an interactive but un-illustrated polyclave is
available, for use in conjunction with the illustrations
in this paper, from the author on request and provision
of one MS-DOS-formatted 3.5" or 5.25" HD diskette.
The programme itself must be obtained from EX-
ETER SOFTWARE, 100 North Country Road,
Setauket, New York 11733, USA. The advantage of a
polyclave is that it can be entered with any character
found easiest by the user and any path can be followed
through the data matrix, as opposed to the fixed path
set by a dichotomous key.

The Cameroon material examined was all collected
by members of the TIGER team, some in the period
July to December, 1992 and some later, up to 1997.
They were variously Drs D. Bignell, P. Eggleton, D.
Jones, J. Lawton and T. Wood, and Luc Dibog
(Eggleton et al., 1996). The Congo material was col-
lected later, in the course of a single expedition by Dr
Eggleton and Mr.Davies. It is not practicable to distin-
guish the contributions of individual participants in
the lists of material examined.

AMICOTERMES Sands 1972
(Amicus, L., ‘friend (ly), kind’)
Type-species: Amicotermes galenus Sands 1972

Imago. See Sands 1972. None known apart from
Type-species.

Worker. Small to medium-sized, head capsule width
0.565 to 0.800 mm; head capsule setae usually sparse;
postclypeus strongly inflated, bulging, length more
than half its width at the ginglymi, width 0.3—0.434,
length 0.154—0.25 mm. Fore tibia moderately to strongly

W.A. SANDS

swollen, index (length/width) 2.954 to 4.759. Apical
teeth of mandibles longer than first marginals, left
mandible index 0.697 to 1.205, right mandible index
0.879 to 1.374; left fourth marginal tooth with proxi-
mal point clear or hidden behind molar prominence,
complex ratio La divided by L1.Lm 17.556 to 37.866;
point of first marginal tooth of right mandible behind
line from apical to second marginal, front edge of first
marginal shorter than or subequal to that of second, its
exposed posterior edge longer than that of the second
marginal, right first to second marginal index 1.35 to
2.094; second marginal prominent, second marginal
index 0.788 to 1.262, complex ratio Ra divided by
R1.Rm 20.164 to 34.01. Crop large with hemispheri-
cal ‘bursters’ on front dorsum, gizzard bell-shaped,
not tapered to enter mesenteron. Mesenteric junction
with proctodeum nearly transverse, slightly angled, in
ventral view of unopened abdomen beneath or to right
of malpighian knot, beneath or slightly to left or right
of nerve cord. Enteric valve seating three-lobed, the
third, inner, smaller than or equal to the outer two,
connected to third section of proctodeum (‘pouch’) by
very long neck, left lateral or dorsal in position in
unopened abdomen. Main armature of enteric valve
ridges consists of tapered apical spines, protruding
through valve exit into third section of proctodeum,
subsidiary armature varies from prominent spines dif-
ferent from or smaller than main armature to scales
fringed with minute spicules or bearing single small
spines; membranous wall of enteric valve between and
beyond ridges usually scaly, with spicules.
Amicotermes is a genus easily recognised in the
worker caste by the several features of the abdominal
anatomy visible from the exterior and especially by
the very characteristic internal armature of the enteric
valve, which also supplies many important specific
characters. The workers are small to medium-sized
among soldierless Apicotermitinae, and although fairly
common, are not the most numerous members of this
group to be found in ecological soil samples.
Autothysis (the suicidal rupture of the body wall by
the contraction of abdominal muscles, accompanied
by the discharge of defensive fluids or gut contents) as
a defence mechanism aginst small predators such as
ants is common in the soldierless Apicotermitinae
(Sands, 1972, 1982) and its development may be
responsible for the loss of the soldier caste. In this
genus, extreme autothysis is characterised by a pair of
hemi-spherical ‘bursters’ , apparently inflated from the
front of the crop by pressure from the contracting
abdominal wall muscles, which cause the dorsal body
wall to rupture in front of the first abdominal tergite, so
that few specimens in any series are not so damaged
and most of these have distorted abdomens. . Only in
one species, A. tithasus Sands, 1972 of the sister-
group Acholotermes has a similar bell-shaped
proventriculus been observed and here the rupture of

~~

REVIEW OF AMICOTERMES

151

Table 1 Condensed data matrix, outgroup (in parentheses) included. Continuous variables (measurements) are coded as
means allocated to key states, while variable multi-states are coded as medians. Missing or inapplicable data is shown as a

query.
characters 1-60
(Acholotermes) ZSZ2r2i22) Mie S221 221 3838S i228 22 22s M225 2 23'4 2:7:272222622
autothysius PEA A2MAP 2A M222 ie 2A TTS 33" 321321252 2221321222" 3412211322
camerunensis ECOL AL 222 22222222, 22222222) 232322521 eee 242223) 3422333312
congoensis U2 AU 2A S220 220 2222222333) S2ZU2ZT2252 92221242234 S3822221422
cristatus Z2TAMNZUA U2 S222 2322101 221 ITT 222) 202202) 122343333) 332333 L422
dibogi ASA AN AZ 2M ZZ SAEZ Ala SS S395" S2Z220 25k 2 es 2323 3322331512
galenus SS 122 S2ii22200 92122222333 SlL2ZtLisi2, 23210264456 34343211524
gasteruptus SMe Ze. Al S221 22S SS Sol si2t5 22) 2221221222 Ssierl1 322
ivorensis 22222 AMZ ZZ VIZA2Z 21 2212222338 seo WLS 23 2210 199233. 8423220312
mayombei LN LANA 2 2S IM A2 32202121 PUUS SIMS LSA 251292922 332234' Asti220412
multispinus MA ee Sear See eI AZAZ2395 SSLAUZ522 22222e2223 S2eliris22
mbalmayoensis 1311122222 1121332211 2112222333 3212212422 1121364334 34434427422
spiculatus ZAMMUSIA22 9322032201 23202222 21730125 2092322243823) 3423432412

the body wall in autothysis appears to be mediated
more by pressure from the crop against the underside
of the slightly inflated colon which crosses it at this
point. The proctodeal first segment beyond the junc-
tion with the mesenteron is usually tubular throughout
its length. The mandibles are generally similar to those
of Acholotermes.

In Acholotermes the main armature of the hexa-
radially symmetrical enteric valve ridges is never
more than a few minute spicules fringing scales, but
the membranous wall between and beyond the ridges
is scaly and spiculate like that of Amicotermes.The
_ other genera having enteric valve armature apparently
related to that of Amicotermes are Apagotermes and
Ateuchotermes. The former is smaller and the latter
larger, both having different mandibles from Amico-
termes and both having specialised features of their
valve armature, which, however, do not include a
spiculate and scaly or carunculate inter-ridge mem-
brane (Sands 1972, 1998).

Key to the species of AMICOTERMES
Sands 1972

The disadvantages of dichotomous keys resulting from
forcing on the user a single path chosen by the author
through the data matrix are well-known. In the key
which follows, some couplets appear longer than usual,
especially when nearing an identification. This is ne-
cessitated by the extreme difficulty of identifying
worker castes on the basis of variable characters and
some that require delicate dissections. The solution

offered is a combination of more easily observed
features that are currently diagnostic and may provide
a simple identification but might break down because
the variation is imperfectly known, and more difficult
but possibly more reliable characters. This is an at-
tempt at compromise between the fallible simplicity of
a normal dichotomous key and the greater flexibility
of a polyclave.

1 Head capsule width 0.763 to 0.800 mm, postclypeus
width: OAS toiOAS4 own ie. ses. dea <csensattavasscttvsatevaseeetcs 2

— Head capsule width 0.565 to 0.717 mm, postclypeus
Wena AYO) SOLO Fey(O),3I2)5) Trade eels seepereraccsceceo-oreboakeeeerecece 3

2 Hind tibia length 0.725 mm, fore tibia length 0.563 mm,
fore tibia thickness 0.13 mm, fore tibial index (length
over width) 4.327, left mandible index 1.092, right man-
dible, complex ratio Ra divided by R1.Rm, 31.28. Fore
coxae with four or more prominent larger setae arranged
in line or group on anterior surface, fore tibial apical
spurs numbering three, but third vestigial or much smaller
than the other two. Left mandible, first marginal tooth
with anterior edge distinctly longer than posterior, left
fourth marginal tooth in front view with proximal end
clear of molar prominence; anterior margin of right first
marginal tooth equal in length to that of second. Enteric
valve ridges, distinctly ‘waisted’ at base of apical spines,
then wider, longest apical spines on ridge one numbering
1-9, those of ridges in position three longer than least
width at base of spines, subsidiary armature on one or
more of enteric valve ridges mainly with prominent
spines different from or smaller than main armature,
graduated, spines or spicules becoming longer towards
distal end of ridge, scales or reticulations well defined,
distinct, symmetry of enteric valve armature bilateral due

152

to marked differences in spine and ridge size, spines or
spicules on membranous wall numerous but not dense,
scattered, fringing scales proximally ..............::ceeeeereee
eee galenus (page 159, Figs 76-82, 177, 178)

Hind tibia length 0.779 to 0.82 mm, fore tibia length
0.614 to 0.677 mm, fore tibia thickness 0.159 to 0.173
mm, fore tibial index (length over width) 3.602 to 4.077,
left mandible index 0.896 to 0.978, right mandible, com-
plex ratio Ra divided by R1.Rm, 20.164 to 26.881. Fore
coxae without prominent larger setae on anterior surface,
fore tibial apical spurs numbering three, third well devel-
oped, at least half the size of the others. Left mandible,
first marginal tooth approximately equilateral, left fourth
marginal tooth in front view just reaches side of molar
prominence, or with proximal end hidden behind molar
prominence; anterior margin of right first marginal tooth
shorter than that of second, enteric valve ridges, more or
less tapering ellipsoid to base of apical spines, longest
apical spines on ridge one numbering 10-14, those of
ridges in position three shorter than or equal to least
width, subsidiary armature on one or more of enteric
valve ridges mainly with small single backwardly di-
rected spines on scales, more or less uniform throughout
length of ridge, scales or reticulations indistinct, outlines
vague or incomplete, symmetry of enteric valve armature
more or less hexa-radial apart from slight differences in
ridge size, spines or spicules on membranous wall dense
distally, fringing scales proximally ..............0.:cceeeeee
... mbalmayoensis (page 163, Figs 125-137, 183, 184)

Symmetry of enteric valve armature more or less hexa-
radial apart from slight differences in ridge size ......... 4

Symmetry of enteric valve armature more or less tri-
radial, alternate ridges reduced, or bilateral due to marked
differences in spine and ridge S1Z€ .............:cceseeeeeeeeee 6

Lateral displacement of enteric valve in unopened abdo-
men to left of heart in dorsal view, first section of
proctodeum very long. Main armature of enteric valve
ridges with very elongated tapered spines, ridges
unsclerotised or colourless, spines or spicules on mem-
branous wall very sparse, scattered ............0:e:ceeseeeceeees
beceneusdenetrantea cressonront dibogi (page 158, Figs 62-75, 176)

Lateral displacement of enteric valve in unopened abdo-
men to right of heart in dorsal or to left of nerve cord in
ventral view. Main armature of enteric valve ridges with
large backwardly directed erect conical spines, ridges
sclerotised at distal ends, pale yellow to pale brown or
yellow-brown, spines or spicules on membranous wall
numerous or dense distally, fringing scales proximally

Fore coxae with four or more prominent larger setae
arranged in line or group on anterior surface, fore tibial
apical spurs numbering three, but third much smaller
than the other two. Left mandible, third marginal tooth
with anterior edge equal in length to that of first.
Proctodeal first segment shorter, up to eight times proxi-
mal width, enteric valve ‘seating’ third lobe approximately
equal in size to outer pair. Enteric valve ridges sclerotised
at distal end, pale brown or yellow-brown, largest spines
on ridges numbering up to 12; subsidiary armature on

W.A. SANDS

one or more ridges mainly with backwardly directed,
well defined distinct scales, some fringed with small,
scattered, sparse spines or spicules, more or less uni-
formly distributed throughout length of ridge; membran-
ous wall of enteric valve between and beyond ridges
armed with short spines or spicules, dense distally, fring-
INS SCales proximal ly eee sees se res eec ee eee eee
ote fast teeeres cristatus (page157, Figs 47-61, 173, 174)

Fore coxae without prominent larger setae on anterior
surface, or with one to three at most, fore tibial apical
spurs numbering three, third well developed, at least half
the size of the others. Left mandible, third marginal tooth
with anterior edge shorter than that of first. Proctodeal
first segment longer, over eight times proximal width,
enteric valve ‘seating’ third lobe smaller than outer pair.
Enteric valve ridges weakly sclerotised distally, pale
yellow or yellow, largest spines on ridges numbering 12—
25; subsidiary armature on one or more ridges mainly
with prominent spines different from or smaller than
main armature, borne on indistinct scales or reticulations,
covering entire ridge surface, graduated, spines or spicules
becoming longer towards distal ends of ridges; membra-
nous wall of enteric valve between and beyond ridges
armed with minute spines or spicules, numerous but not
dense, scattered, fringing scales proximally ..................
Sea camerunensis (page 154, Figs 16-28, 175)

Hind tibia length 0.784 to 0.794 mm, fore tibia length
0.620 to 0.625 mm Enteric valve main armature of all
ridges with large backwardly directed erect conical spines,
but ridge in position one unsclerotised or colourless
throughout, ridge two weakly and ridges three and four
fully sclerotised yellow-brown; subsidiary armature of
enteric valve ridges prominent, covering entire ridge
surface ........ spiculatus (page 165, Figs 151-161, 186)

Hind tibia length 0.565 to 0.736 mm, fore tibia length
0.489 to 0.574 mm Enteric valve main armature of all
ridges with very elongated tapered spines, uniformly
sclerotised, yellow-brown or weakly sclerotised distally,
pale yellow or yellow; subsidiary armature of enteric
valve ridges scattered, somewhat or very sparse, few
scales fringed with spines or spicules ...............-..0:20+ 7

Longest apical spines of enteric valve armature shorter
than or equal to least width of ridges in position three,
subsidiary armature of enteric valve ridges with scales or
reticulations well defined, distinct ...............:::::ceeee 8

Longest apical spines of enteric valve armature longer
than least width of ridges in position three at base of
spines, subsidiary armature of enteric valve ridges with
scales or reticulations indistinct, outlines vague or in-
COMPS sos... 2c. cee: vec ceaesveecesetaccetsossecceus She ese eee 10

Left fourth marginal tooth in front view just reaches side
of molar prominence. Symmetry of enteric valve arma-
ture bilateral due to marked differences in spine and ridge
size, membranous wall of enteric valve between and
beyond ridges armed with short spicules ...............:::-0-+
ee mayombei (page 162, Figs 111-124, 181, 182)

Left fourth marginal tooth in front view with proximal
end clear of molar prominence. Symmetry of enteric

REVIEW OF AMICOTERMES

10

Jl

valve armature more or less tri-radial, alternate ridges
reduced, membranous wall of enteric valve between and
beyond ridges armed with minute spicules.................. 9

Head capsule with emergent, longest setae regularly and
symmetrically arranged. Fore tibial apical spurs number-
ing three, but third vestigial or much smaller than the
other two. Enteric valve ‘seating’, third lobe approxi-
mately equal in size to outer pair; enteric valve ridges
weakly sclerotised distally, pale yellow or yellow; spines
or spicules on membranous wall of valve between ridges
numerous but not dense, scattered, fringing scales proxi-
mally ....... congoensis (page156, figs 29-46, 171, 172)

Head capsule with emergent, longest setae randomly
scattered. Fore tibial apical spurs numbering three, third
well developed, at least half the size of the others. Enteric
valve ‘seating’, third lobe smaller than outer pair, enteric
valve ridges sclerotised at distal ends, pale brown or
yellow-brown, spines or spicules on membranous wall
dense distally, fringing scales proximally ...............00
eco eee autothysius (page 153, figs 1-15, 169, 170)

Anterior margin of first marginal tooth of right mandible
equal in length to that of second. Enteric valve ‘seating’
third lobe approximately equal in size to outer pair;
largest apical spines of enteric valve armature on ridges
one and three numbering 1-9; subsidiary armature on
enteric valve ridges very sparse, with a few backwardly
directed scales fringed with small spines or spicules, or
with small single backwardly directed spines on scales;
membranous wall of enteric valve between and beyond
ridges smooth, with very sparse, scattered spines or
SPICULESieerecss--< ivorensis (page 161, figs 99-110, 180)

Anterior margin of right first marginal tooth shorter than
that of second. Enteric valve ‘seating’ third lobe smaller
than outer pair; largest apical spines of enteric valve
armature on ridge one numbering 10—20, those on ridges
three numbering 10 to over 20; subsidiary armature on
one or more of enteric valve ridges with scattered, some-
what sparse prominent spines different from or smaller
than main armature; membranous wall of enteric valve
between and beyond ridges scaly, with numerous but not
dense scattered spicules, fringing scales proximally .....

Head capsule with emergent, longest setae regularly and
symmetrically arranged. Left mandible, third marginal
tooth with anterior edge equal in length to that of first.
Enteric valve ridges weakly sclerotised distally, pale
yellow or yellow, positions one and three with 20 or more
apical spines; membranous wall of enteric valve between
and beyond ridges armed with short spicules ................
Meteneecesieaeees multispinus (page 164, figs 138-150, 185)

Head capsule with emergent, longest setae randomly
scattered. Left mandible, third marginal tooth with ante-
rior edge shorter than that of first. Enteric valve ridges
sclerotised at distal end, pale brown or yellow-brown,
positions one and three with 10—20 apical spines; mem-
branous wall of enteric valve between and beyond ridges
armed with minute spicules ............:::cecceceeceeeeeeeeeneeneees
Tes gasteruptus (page 160, figs 83-98, 179)

153

DESCRIPTIONS OF SPECIES

Amicotermes autothysius sp. n.
(Figs 1-15, 169, 170)

Head capsule pale yellow or yellow, antennal flagellum
colour of darkest part yellow or pale yellow-brown,
pronotum and legs yellow-white, abdominal tergites
hyaline.

Head capsule short oval near circular, setae numer-
ous but not dense, emergent, longest setae randomly
scattered. Fontanelle absent. Fore coxae without promi-
nent larger setae or with up to four or sometimes more
prominent larger setae arranged in line or group on
anterior surface, fore tibial apical spurs numbering
three, third well developed, at least half the size of the
others.

Left mandible, first marginal tooth with anterior
edge distinctly longer than posterior or approximately
equilateral, third marginal tooth with anterior edge
shorter than or equal in length to that of first, fourth
marginal tooth in front view with proximal end clear of
molar prominence. Right mandible, anterior edge of
first marginal tooth equal in length to or shorter than
that of second.

Junction of mesenteron and proctodeum starting
beneath or to right of nerve cord in ventral view,
proctodeal first segment shorter, up to eight times
proximal width tubular throughout its length, lateral
displacement of enteric valve in unopened abdomen to
right of heart in dorsal or to left of nerve cord in ventral
view, enteric valve ‘seating’ third lobe smaller than
outer pair.

Enteric valve ridges, distinctly ‘waisted’ at base of
apical spines, then wider, all ridges sclerotised at distal
end, pale brown or yellow-brown, main armature with
very elongated tapered spines, the largest on ridges
one and four numbering 1—14, those on ridges two, 1—
9, and on ridges three, 10-20, symmetry of enteric
valve armature more or less tri-radial, alternate ridges
reduced; longest spines of main armature straight or
curved but still evenly tapered, shorter than or equal to
least width of ridges in position three; subsidiary
armature on one or more of enteric valve ridges mainly
with scattered, somewhat sparse prominent spines
different from or smaller than main armature, gradu-
ated, spines or spicules becoming longer towards distal
end of ridge, scales or reticulations well defined,
distinct; membranous wall of enteric valve between
and beyond ridges scaly, armed with minute spicules,
dense distally, fringing scales proximally.

Measurements and diagnostic ratios {24 individuals
from 9 series]:

Head capsule width 0.565 to 0.717 mm (mean 0.633
SD 0.039).

154

Postclypeus width 0.300 to 0.395 mm (mean 0.332 SD
0.026).

Postclypeus length 0.154 to 0.233 mm (mean 0.185
SD 0.021).

Left mandible index 0.697 to 1.144 (mean 0.843).

Right mandible index 0.879 to 1.278 (mean 1.040).

Right second marginal index 0.817 to 1.166 (mean
0.982).

Right first to second marginal index 1.432 to 2.095
(mean 1.734).

Pronotum width 0.375 to 0.495 mm (mean 0.424 SD
0.027).

Pronotum length 0.169 to 0.293 mm (mean 0.209 SD
0.024).

Hind tibia length 0.565 to 0.706 mm (mean 0.649 SD
0.032).

Fore tibia length 0.489 to 0.556 mm (mean 0.517 SD
0.018).

Fore tibia thickness 0.131 to 0.144 mm (mean 0.138
SD 0.004).

Fore tibial index (length over width) 3.441 to 3.984
(mean 3.741).

Left mandible, complex ratio La divided by L1.Lm
17.556 to 29.807 (mean 23.846).

Right mandible, complex ratio Ra divided by R1.Rm
20.717 to 34.01 (mean 25.475).

COMPARISONS. The Cameroonian species most
closety related to A. autothysius is A. gasteruptus,
from which it is only distinguished by the shorter
apical spines of the enteric valve main armature and
the more distinct outlines of the scales of the subsidi-
ary armature. Its symmetry in A. autothysius also tends
towards the bilateral rather than being clearly tri-
radial. This pair of species forms a terminal group in
all the trees apart from the Saitou and Nei (1987)
Neighbour-Joining tree. A. camerunensis has the
fourth marginal tooth of the left mandible just reaching
the side of the molar prominence, a proportionately
longer fore tibia, and a hexa-radial enteric valve arma-
ture with large straight conical spines. In A.
congoensis the third fore tibial apical spur is reduced,
being one third or less the length of the others, and the
third lobe of the enteric valve seating is roughly equal
in size to the outer pair; the largest apical spines of the
enteric valve armature are generally longer and the
ridges more clearly triradial. A. cristatus is generally
larger and also shares the features that separate A.
camerunensis. A. dibogi differs in the proportions of
the mandibular teeth, but most notably in the greater
length and inflation of the first proctodeal segment,
hence also the dorsal position of the enteric valve
seating; the valve armature is quite different from any
other member of the genus being armed with very fine
long apical spines and hightly spiculate fringed scales.
A. galenus is, again, larger and the enteric valve
armature differs in the length and straightness of the

W.A. SANDS

apical spines. A. ivorensis has fewer head setae and
lacks prominent fore coxal setae, while its enteric
valve has fewer but longer apical spines and a much
less developed subsidiary armature. In A. mayombei
the fourth marginal tooth of the left mandible just
reaches the side of the molar prominence; the spiculate
fringes of the inter-ridge membrane of the enteric
valve are denser and longer than in A. autothysius and
the symmetry of the valve is more bilateral. A.
mbalmayoensis is considerably larger and its enteric
valve is hexa-radially symmetrical with differently
shaped ridges on which the subsidiary armature con-
sists of single small spines on indistinct scales, more or
less uniform over the entire ridge surface. In A.
multispinus the left fourth marginal tooth just reaches
the side of the molar prominence; the enteric valve
ridges carry longer and more numerous apical spines,
while the scales of the subsidiary armature are very
indistinct in outline. Finally, A. spiculatus has longer
and slightly more slender legs, and the enteric valve is
bilaterally symmetrical, ridges three and four being
differentially sclerotised and three, much larger while
ridge one is colourless and two intermediate; the entire
ridge surfaces are spiny with the subtending scales
very weakly marked.

MATERIAL EXAMINED. [total 13 series]

Holotype, worker and Paratypes from Type series
(field collection number, H,P-2), Cameroon:
Mbalmayo Forest Reserve, Ekombitie July 1996
(Eggleton et al.); other Paratype material, Eboufek,
July—Aug.1992, Ekombitie, Ebogo, Nov. 1995, Ebogo,
Ekombitie, 5 series, Feb. 1996, Ebogo, Mar. 1996,
Ekombitie, Ebogo, 3 series, July 1996, (Eggleton et
al.) (all in BMNH).

Amicotermes camerunensis sp. 0.
(Figs 16—28, 175)

Head capsule yellow-white or pale yellow, antennal
flagellum, colour of darkest part, pronotum and legs
yellow-white, abdominal tergites hyaline.

Head capsule circular or short oval near circular,
setae sparse, emergent, longest setae regularly and
symmetrically arranged. Fontanelle absent. Fore coxae
without or with one to three prominent larger setae on
anterior surface, fore tibial apical spurs numbering
three, third well developed, at least half the size of the
others.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
shorter than that of first, fourth marginal tooth in front
view just reaches side of molar prominence. Right
mandible, anterior edge of first marginal tooth equal in
length to or shorter than that of second.

Junction of mesenteron and proctodeum starting to
left of or beneath nerve cord in ventral view, proctodeal

REVIEW OF AMICOTERMES

first segment longer, over eight times proximal width,
tubular throughout its length or dilated to about twice
its basal width, lateral displacement of enteric valve in
unopened abdomen to right of heart in dorsal or to left
of nerve cord in ventral view; enteric valve ‘seating’
third lobe smaller than outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines, weakly sclerotised distally,
pale yellow or yellow, main armature of all valve
ridges with large backwardly directed erect conical
spines, the largest numbering 10-20, the longest
straight and evenly tapered, shorter than or equal to
least width of ridges in position three, symmetry of
enteric valve armature more or less hexa-radial apart
from slight differences in ridge size; subsidiary arma-
ture on one or more of enteric valve ridges mainly with
prominent spines different from or smaller than main
armature covering entire ridge surface, graduated,
spines or spicules becoming longer towards distal end
of ridge, scales or reticulations indistinct, their out-
lines vague or incomplete; membranous wall of enteric
valve between and beyond ridges scaly, armed with
numerous but not dense minute spines or spicules,
scattered, fringing scales proximally.

Measurements and diagnostic ratios {15 individuals
from 5 series]:

Head capsule width 0.652 to 0.711 mm (mean 0.687
SD 0.016).

Postclypeus width 0.340 to 0.370 mm (mean 0.360 SD
0.009).

Postclypeus length 0.172 to 0.215 mm (mean 0.199
SD 0.012).

Left mandible index 0.731 to 0.902 (mean 0.839).

Right mandible index 0.992 to 1.187 (mean 1.093).

Right second marginal index 0.806 to 1.175 (mean
0.973).

Right first to second marginal index 1.643 to 2.088
(mean 1.820).

Pronotum width 0.411 to 0.468 mm (mean 0.447 SD
0.017).

Pronotum length 0.192 to 0.251 mm (mean 0.218 SD
0.019).

Hind tibia length 0.696 to 0.794 mm (mean 0.74 SD
0.026).

Fore tibia length 0.589 to 0.632 mm (mean 0.607 SD
0.014).

Fore tibia thickness 0.147 to 0.205 mm (mean 0.168
SD 0.013).

Fore tibial index (length over width) 2.954 to
4.121(mean 3.624).

Left mandible, complex ratio La divided by L1.Lm
18.546 to 23.477 (mean 21.018).

Right mandible, complex ratio Ra divided by R1.Rm
22.193 to 29.197 (mean 25.539).

155

COMPARISONS. A. camerunensis has already been
compared with A. autothysius under that species. A.
congoensis differs in having shorter legs; the anterior
edge of the left third marginal tooth is equal to that of
the first and the fourth marginal has its proximal end
clear of the side of the molar prominence; the third
lobe of the enteric valve seating is approximately
equal to the other two, while the spines of the main
armature are curved on a tri-radial valve. In A. cristatus
the third fore tibial apical spur is smaller and the front
edge of the third marginal tooth of the left mandible is
equal in length to that of the first; the apical spines of
the main enteric valve armature are fewer and the
subsidiary armature is much less developed. A. dibogi
is characterised externally by the very long, dilated P1
which pushes the enteric valve seating with its three
equal lobes into a dorsal position to the left of the heart
line; the main armature of the enteric valve is quite
different, with its slender apical spines and the subsidi-
ary armature with well defined scales fringed with
spicules. A. galenus is larger and has the front edge of
the left first marginal tooth longer than the hind edge,
with the fourth marginal clear of the molar promi-
nence; the mandibular indices are higher; the apical
spines of the main enteric valve armature are fewer,
more slender and longer on a bilaterally symmetrical
valve. A. gasteruptus is smaller, with shorter hind
tibiae; its enteric valve ridges are ‘waisted’ apically
with elongated tapered curved spines, fewer in number
on ridges two and four of a tri-radially symmetrical
valve. While about the same size, A. ivorensis has
shorter legs and the left fourth marginal tooth clear of
the molar prominence; the third lobe of the enteric
valve seating is approximately equal to the other two
and the main armature has fewer, longer apical spines
on a tri-radial valve with reduced subsidiary armature.
In A. mayombei the front edge of the left first marginal
tooth is longer than its hind edge and equal to that of
the third, while the front edge of the right first mar-
ginal is shorter than that of the second; the enteric
valve ridges are, again, ‘waisted’ and one, two and
four bear fewer spines on a bilaterally symmetrical
valve. A. mbalmayoensis is larger and the subsidiary
armature of its enteric valve ridges consists of small
single spines on scales, more or less uniform through-
out the length of each ridge. A. multispinus is smaller,
with shorter legs; the front edges of the left first and
third marginals are subequal and that of the right first
marginal is shorter than that of the second; the enteric
valve ridges are again, ‘waisted’ and the longest apical
spines on ridges three are longer than the least width of
these ridges, the symmetry of the valve being tri-
radial. Finally, in A. spiculatus the front edges of the
left first and third marginals are subequal, the proxi-
mal end of the fourth marginal is clear of the molar
prominence and the front edge of the right first mar-
ginal is shorter than that of the second; the enteric

156

valve armature, while superficially similar, is distinc-
tive in being bilaterally symmetrical owing to
differential sclerotisation of the ridges.

MATERIAL EXAMINED. [total, 6 series]

Holotype worker and Paratypes from Type series (field
collection no. Soil#1), Cameroon: Mbalmayo Forest
Reserve, Eboufek July—Aug. 1992 (Eggleton et al.);
other Paratype material, Ebogo, 2 series, July—Aug.
1992, Ebogo, Feb. 1996, Ekombitie, Ebogo July 1996
(Eggleton et al.) (all in BMNH).

Amicotermes congoensis sp. 0.
(Figs 29-46, 171, 172)

Head capsule yellow-white or pale yellow, antennal
flagellum, colour of darkest part, pale yellow or yel-
low, pronotum and legs yellow-white, abdominal
tergites hyaline.

Head capsule short oval near circular, setae sparse
or numerous but not dense, emergent, longest setae
regularly and symmetrically arranged. Fontanelle
present as pale spot or small depression or absent. Fore
coxae with four or more prominent larger setae ar-
ranged in line or group on anterior surface, fore tibial
apical spurs numbering three, but third vestigial or
much smaller than the other two.

Left mandible, first marginal tooth with anterior
edge distinctly longer than posterior or approximately
equilateral, third marginal tooth with anterior edge
equal in length to that of first, fourth marginal tooth in
front view with proximal end clear of molar promi-
nence. Right mandible, anterior edge of first marginal
tooth shorter than that of second.

Junction of mesenteron and proctodeum starting to
left of or beneath nerve cord in ventral view, proctodeal
first segment shorter, up to eight times proximal width,
tubularl throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe approximately equal in size
to outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines or distinctly ‘waisted’ at base
of apical spines, then wider, weakly sclerotised distally,
pale yellow or yellow, main armature with very elon-
gated tapered spines, the largest on ridges one and
three numbering 10-14, those on ridges two and four
numbering up to 10, longest spines curved but still
evenly tapered, shorter than or equal to least width of
ridges in position three, symmetry of enteric valve
armature more or less tri-radial, alternate ridges re-
duced; subsidiary armature on one or more of enteric
valve ridges with scattered, somewhat sparse, promi-
nent spines different from or smaller than main
armature, graduated, spines becoming longer towards
distal end of ridge, scales or reticulations well defined,

W.A.SANDS

distinct; membranous wall of enteric valve between
and beyond ridges scaly armed with minute numerous
but not dense spicules, scattered, fringing scales proxi-
mally.

Measurements and diagnostic ratios {15 individuals
from 4 series]

Head capsule width 0.659 to 0.705 mm (mean 0.684
SD 0.014).

Postclypeus width 0.340 to 0.380 mm (mean 0.365 SD
0.013).

Postclypeus length 0.177 to 0.215 mm (mean 0.197
SD 0.011).

Left mandible index 0.808 to 1.205 (mean 0.940).

Right mandible index 0.971 to 1.374 (mean 1.168).

Right second marginal index 0.885 to 1.174 (mean
1.019).

Right first to second marginal index 1.350 to 1.921
(mean 1.695).

Pronotum width 0.410 to 0.472 mm (mean 0.450 SD
0.019).

Pronotum length 0.184 to 0.236 mm (mean 0.213 SD
0.017).

Hind tibia length 0.670 to 0.710 mm (mean 0.692 SD
0.013).

Fore tibia length) 0.535 to 0.574 mm (mean 0.553 SD
0.011).

Fore tibia thickness 0.136 to 0.150 mm (mean 0.142
SD 0.004).

Fore tibial index (length over width) 3.598 to 4.185
(mean 3.889).

Left mandible, complex ratio La divided by L1.Lm
17.791 to 37.866 (mean 25.316).

Right mandible, complex ratio Ra divided by R1.Rm
21.345 to 32.805 (mean 25.805).

COMPARISONS. A. congoensis has already been com-
pared with the preceding two species; It forms a
terminal pair with A. multispinus in most of the phenetic
trees, but joins with A. autothysius, A. gasteruptus and
A. mayombei at node 16 in the strict consensus tree; the
comparisons with the two latter species are given
below. A. cristatus is larger with longer fore legs and
the left fourth marginal tooth just reaching the side of
the molar prominence; the spines of the main armature
of the enteric valve ridges are large, straight and
conical on a more or less hexa-radially symmetrical
valve. A. dibogi has the front edge of the left third
marginal tooth shorter than that of the first, and the
fourth just reaching the side of the molar prominence;
the main distinguishing feature is again the long di-
lated P1 with the enteric valve seating dorsal, to the
left of the heart; the hexa-radial enteric valve armature
with its fine straight spines and spiculate scales is
unique in the genus. A. ivorensis lacks prominent setae
on the fore coxa, the front edge of the left third

REVIEW OF AMICOTERMES

marginal tooth is shorter than that of the first, and of
the right first marginal, equal to that of the second; the
longest apical spines of the main enteric valve arma-
ture are longer than the least width of ridges three but
all armature, main and subsidiary, is notably sparse. A.
galenus is larger, the front edge of the left third mar-
ginal is shorter than that of the first and that of the right
first marginal equal to the second; the longest apical
spines of the main enteric valve armature are straight,
longer than the least width of ridges three and fewer on
the bilaterally symmetrical valve. A. gasteruptus is
slightly smaller, the front edge of the left third mar-
ginal tooth is shorter than the first and the third lobe of
the enteric valve seating is smaller than the other two;
the longest apical spines of the main enteric valve
armature are longer than the least width of ridges three
and the scales of the subsidiary armature are indistinct.
A. mayombei has the third fore tibial apical spur well
developed, the fourth marginal tooth of the left mandi-
ble just reaching the side of the molar prominence, and
the third lobe of the enteric valve seating smaller than
the outer pair; the enteric valve armature is bilaterally
symmetrical. A. mbalmayoensis is considerably larger;
it lacks prominent setae on the fore coxae and the third
apical spur of the fore tibia is well developed; the
proximal end of the fourth marginal tooth of the left
mandible reaches the side of the molar prominence or
is hidden behind it; the third lobe of the enteric valve
seating is smaller than the outer pair and the longest
apical spines of the main enteric valve armature are
straight on a hexa-radially symmetrical valve. A.
multispinus is slightly smaller and has the third fore
tibial spur well developed; the left fourth marginal
reaches the side of the molar prominence; the longest
apical spines of the enteric valve armature are longer
than the least width of ridges three and more numerous
on one and three. Finally, A. spiculatus is about the
same size but has markedly longer legs, the fore tibiae
with well developed third spurs; the apical spines of
the main armature of the enteric valve are straight and
conical and the symmetry is bilateral largely because
of markedly different degrees of sclerotisation of the
ridges.

MATERIAL EXAMINED. [total 5 series]
Holotype worker and Paratypes from Type series (field
collection no. T,.13), Congo (Brazzaville): Mayombe,
Feb. 1997; other Paratype material, Feb. 1997, 4 series
(Eggleton and Davies) (all in BMNH).

Amicotermes cristatus sp. 0.
(Figs 47-61, 173, 174)

Head capsule pale yellow, antennal flagellum, colour
of darkest part, pale yellow or yellow, pronotum and
legs yellow-white, abdominal tergites hyaline.

Head capsule short oval near circular or oval, setae

iS)

sparse, emergent, longest setae regularly and sym-
metrically arranged. Fontanelle present as pale spot or
small depression or absent. Fore coxae with four or
more prominent larger setae arranged in line or group
on anterior surface, fore tibial apical spurs numbering
three, but third vestigial or much smaller than the other
two.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
equal in length to that of first, fourth marginal tooth in
front view just reaches side of molar prominence.
Right mandible, anterior margin of first marginal tooth
shorter than that of second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width
tubular throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe approximately equal in size
to outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines, sclerotised at distal ends, pale
brown or yellow-brown, main armature of all ridges
with large backwardly directed erect conical spines,
those on ridges one, three and four numbering up tol 4,
those on ridges two, less than 10, straight and evenly
tapered, shorter than or equal to least width of ridges in
position three, symmetry of enteric valve armature
more or less hexa-radial apart from slight differences
in ridge size; subsidiary armature of enteric valve
ridges with scattered, sparse spines or spicules on
scales, scales or reticulations well defined, distinct,
more or less uniform throughout length of ridge; mem-
branous wall of enteric valve between and beyond
ridges scaly, armed with short spicules, dense distally,
fringing scales proximally.

Measurements and diagnostic ratios [8 individuals
from 3 series]:

Head capsule width 0.673 to 0.717 mm (mean 0.690
SD 0.015).

Postclypeus width 0.360 to 0.395 mm (mean 0.379 SD
0.013).

Postclypeus length 0.187 to 0.228 mm (mean 0.204
SD 0.015).

Left mandible index 0.871 to 0.965 (mean 0.915).

Right mandible index 0.968 to 1.265 (mean 1.099).

Right second marginal index 0.861 to 1.113 (mean
0.963).

Right first to second marginal index 1.523 to 1.816
(mean 1.701).

Pronotum width 0.448 to 0.512 mm (mean 0.479 SD
0.021).

Pronotum length 0.221 to 0.265 mm (mean 0.241 SD
0.016).

158

Hind tibia length 0.727 to 0.765 mm (mean 0.742 SD

0.015).

Fore tibia length 0.575 to 0.608 mm (mean 0.589
SD 0.011).

Fore tibia thickness 0.144 to 0.148 mm (mean 0.146
SD 0.002).

Fore tibial index (length over width) 3.974 to 4.165
(mean 4.049).

Left mandible, complex ratio La divided by L1.Lm
21.913 to 26.987 (mean 24.192).

Right mandible, complex ratio Ra divided by R1.Rm
21.978 to 27.22 (mean 24.533).

COMPARISONS. A. cristatus is one of the more dis-
tinctive species with its ‘crest’ of large conical spines
on the enteric valve ridges and has already been com-
pared with the three that precede it. The next species,
A. dibogi, is distinctive in the longer, dilated P1 that
pushes the enteric valve seating around to the dorsal
side of the unopened abdomen; the fine apical spines
of the enteric valve main armature and the highly
spiculate fringed scales of the subsidiary armature are
also different from any other species. A. galenus is
larger with higher mandibular indices and has the front
edge of the left first marginal tooth longer than the
hind edge, the third marginal shorter than the first,
while the fourth marginal is clear of the molar promi-
nence; the front edge of the right first marginal is equal
to that of the second; the ridges of the enteric valve
armature are ‘waisted’ at the base of the apical spines,
which are elongated, straight and longer than the least
width of ridges three, the symmetry of the valve being
bilateral. A. gasteruptus is considerably smaller and
has the third lobe of the enteric valve armature smaller
than the outer pair; the valve ridges are ‘waisted’ at the
base of the apical spines which are elongated, curved,
and longer than the least width of ridges three, the
symmetry of the valve being tri-radial. A. ivorensis is
smaller, lacks prominent setae on the fore coxae, has
the left third marginal tooth shorter than the first and
the fourth marginal clear of the molar prominence; the
front edge of the right first marginal tooth is equal to
that of the second; the apical spines of the main
armature of the enteric valve ridges are straight but
elongated and tapered, longer than the least width of
ridges three, on a tri-radially symmetrical valve with
subsidiary armature and that between the ridges very
sparse. A. mayombei has well developed third fore
tibial spurs, the third lobe of the enteric valve seating
smaller than the outer pair and the left third marginal
tooth shorter than the first; the ridges of the enteric
valve armature are ‘waisted’ at the base of the apical
spines, which are elongated and more numerous on
ridges three on a bilaterally symmetrical valve. A.
mbalmayoensis is considerably larger without promi-
nent setae on the fore coxae and with well developed
third fore tibial spurs; the third lobe of the enteric valve

W.A. SANDS

seating is smaller than the outer pair; the apical spines
of the main armature of the enteric valve ridges are
straight but elongated and tapered, while the subsidi-
ary armature consists mainly of small single spines on
scales. A. multispinus is smaller, with well developed
third fore tibial spurs and the third lobe of the enteric
valve seating is smaller than the outer pair; the valve
ridges are ‘waisted’ at the base of the apical spines
which are elongated, curved, more numerous and
longer than the least width of ridges three, the symme-
try of the valve being tri-radial. Lastly, A. spiculatus
also has well developed third fore tibial spurs, while
the left fourth marginal tooth is clear of the molar
prominence; the enteric valve armature is distinctive
in being bilateral owing to differential sclerotisation of
the ridges, one being colourless and three fully
sclerotised.

MATERIAL EXAMINED. _ [total 3 series]
Holotype worker and Paratypes from Type series (field
collection no. T,.11), Congo (Brazzaville): Mayombe,
Feb. 1997; other Paratype material, Feb. 1997, 2 series
(Eggleton and Davies) (all in BMNH).

Amicotermes dibogi sp. n.
(Figs 62-75, 176)

Head capsule pale yellow or yellow, antennal flagellum,
colour of darkest part, yellow or pale yellow-brown,
pronotum yellow-white or pale yellow, legs pale yel-
low, abdominal tergites hyaline.

Head capsule circular to oval, setae sparse, emer-
gent, longest setae more or less regularly and
symmetrically arranged. Fontanelle absent. Fore coxae
usually with four or more prominent larger setae ar-
ranged in line or group on anterior surface, fore tibial
apical spurs numbering three, third well developed or
much smaller than the other two.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
shorter than that of first, fourth marginal tooth in front
view just reaches side of molar prominence. Right
mandible, anterior margin of right first marginal tooth
shorter than that of second.

Junction of mesenteron and proctodeum starting to
left of nerve cord in ventral view, proctodeal first
segment very long, over eight times proximal width,
dilated to about twice its basal width throughout length,
lateral displacement of enteric valve in unopened ab-
domen to left of heart in dorsal view; enteric valve
‘seating’ third lobe approximately equal in size to
outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines, unsclerotised or colourless
throughout with very elongated tapered spines, the
largest numbering up to14, straight and evenly tapered,
shorter than or equal to least width of ridges in position

REVIEW OF AMICOTERMES

three, symmetry of enteric valve armature more or less
hexa-radial apart from slight differences in ridge size;
subsidiary armature on one or more of enteric valve
ridges with prominent spines smaller than main arma-
ture, covering entire ridge surface, graduated, spines
or spicules becoming longer towards distal end of
ridge, scales or reticulations well defined, distinct;
membranous wall of enteric valve between and be-
yond ridges scaly, armed with very sparse, scattered
minute spines or spicules.

Measurements and diagnostic ratios [12 individuals
from 3 series]:

Head capsule width 0.656 to 0.701 mm (mean 0.676
SD 0.014).

Postclypeus width 0.324 to 0.362 mm (mean 0.347 SD
0.011).

Postclypeus length 0.192 to 0.227 mm (mean 0.206
SD 0.012).

Left mandible index 0.801 to 0.902 (mean 0.852).

Right mandible index 0.973 to 1.295 (mean 1.101).

Right second marginal index 0.788 to 1.262 (mean
0.994).

Right first to second marginal index 1.458 to 1.783
(mean 1.632).

Pronotum width 0.440 to 0.489 mm (mean 0.470 SD
0.015).

Pronotum length 0.192 to 0.227 mm (mean 0.206 SD
0.012).

Hind tibia length 0.659 to 0.730 mm (mean 0.703 SD
0.018).

Fore tibia length 0.576 to 0.618 mm (mean 0.594 SD
0.013).

Fore tibia thickness 0.127 to 0.148 mm (mean 0.139
SD 0.006).

Fore tibial index (length over width) 3.952 to 4.759
(mean 4.272).

Left mandible, complex ratio La divided by L1.Lm
18.009 to 23.312 (mean 20.937).

Right mandible, complex ratio Ra divided by R1.Rm
20.653 to 30.758 (mean 23.927).

COMPARISONS. A. dibogihas already been compared
individually with the species that precede it; however,
it is so distictive with its long, dilated P1 that pushes
the enteric valve seating into a left dorsal position and
the finely spiny and spiculate enteric valve armature,
that it does not need detailed comparisons with the
remaining species under its own heading. Although
distinct from the other species of the genus in the
characters specified, it is not so widely different as to
require separation in another genus.

MATERIAL EXAMINED. _ [total 7 series]

Holotype worker and Paratypes from Type series (field
collection no. H,C-1), Cameroon: Mbalmayo Forest
Reserve, Ekombitie, Aug. 1997 (Eggleton et al.); other

159

Paratype material, Ekombitie, Aug. 1995, Nov. 1995,
July 1996, Aug. 1997, Ebogo, Feb. 1996 (Eggleton et
al.) (allin BMNH).This species is named in honour of
Dr. Luc Dibog, a member of the TIGER team, who
was first to recognise that it was a distinct species on
the strength of its enteric valve armature and long first
proctodeal segment.

Amicotermes galenus Sands, 1972 [Type
species]

(Figs 76-82, 177, 178)

Head capsule yellow, antennal flagellum, colour of
darkest part, pronotum and legs, yellow-white or pale
yellow, abdominal tergites yellow-white.

Head capsule circular, setae sparse, emergent, long-
est setae randomly scattered. Fontanelle absent. Fore
coxae with four or more prominent larger setae ar-
ranged in line or group on anterior surface, fore tibial
apical spurs numbering three, but third vestigial or
much smaller than the other two.

Left mandible, first marginal tooth with anterior
edge distinctly longer than posterior, third marginal
tooth with anterior edge shorter than that of first,
fourth marginal tooth in front view with proximal end
clear of molar prominence. Right mandible, anterior
margin of first marginal tooth equal in length to that of
second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width,
tubular throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe smaller than or approxi-
mately equal in size to outer pair.

Enteric valve ridges, distinctly ‘waisted’ at base of
apical spines, then wider, weakly sclerotised distally,
pale yellow or yellow, main armature with very elon-
gated tapered spines, the largest on ridges one and two
numbering 1—9, those on ridges three numbering 10—
14, and on ridge four, up to 14, straight and evenly
tapered, longer than least width of ridges in position
three at base of spines, symmetry of enteric valve
armature bilateral due to marked differences in spine
and ridge size; subsidiary armature on ridges with
scattered, somewhat sparse prominent spines different
from or smaller than main armature, graduated, spines
or spicules becoming longer towards distal end of
ridge, scales or reticulations well defined; membra-
nous wall of enteric valve between and beyond ridges
scaly, armed with minute spicules, numerous but not
dense, scattered, fringing scales proximally.

Measurements and diagnostic ratios [1 individual]:

Head capsule width 0.763 mm.

160

Postclypeus width 0.425 mm.

Postclypeus length 0.25 mm.

Left mandible index 1.092.

Right mandible index 1.329.

Right second marginal index 0.957.

Right first to second marginal index 1.793.

Pronotum width 0.485 mm.

Pronotum length 0.269 mm.

Hind tibia length 0.725 mm.

Fore tibia length 0.563 mm.

Fore tibia thickness 0.13 mm.

Fore tibial index (length over width) 4.327.

Left mandible, complex ratio La divided by L1.Lm
26.495.

Right mandible, complex ratio Ra divided by R1.Rm
31.280.

COMPARISONS. A. galenus has already been com-
pared with the preceding species. A. gasteruptus is
smaller with lower mandible indices and has the front
edge of the right first marginal tooth equal in length to
that of the second; the largest apical spines of the main
enteric valve armature are curved on a tri-radially
symmetrical valve. A. ivorensis is again smaller with
lower mandible indices and has the left first marginal
tooth roughly equilateral; it lacks prominent fore coxal
setae; the largest apical spines of the enteric valve
main armature are less numerous on a tri-radially
symmetrical valve with very sparse subsidiary arma-
ture. A. mayombei is also smaller, with lower mandible
indices and has the front edge of the left third marginal
tooth equal to that of the first, while the fourth mar-
ginal reaches the side of the molar prominence; the
largest apical spines of the main enteric valve armature
are curved and shorter than or equal to the least width
of ridges three. A. mbalmayoensis is larger, lacks
prominent fore coxal setae and has well developed
fore tibial apical spurs; the left first marginal tooth is
roughly equilateral while the fourth marginal reaches
the side of the molar prominence; the largest apical
spines of the main enteric valve armature are shorter
than or equal to the least width of ridges three and the
subsidiary armature consists of small single spines on
scales, the valve symmetry being hexa-radial. A.
multispinus is smaller with lower mandible indices
and has the left first marginal tooth roughly equilateral
while the fourth marginal reaches the side of the molar
prominence; the front edge of the right first marginal
tooth is shorter than that of the second; the largest
apical spines of the main enteric valve armature are
curved on a tri-radially symmetrical valve and much
more numerous on ridges one and three. A. spiculatus
is smaller, with lower main mandible indices and has
the left first marginal tooth roughly equilateral while
the front edge of the left third marginal tooth is equal
to that of the first; it has well developed fore tibial
apical spurs; the largest apical spines of the main

W.A. SANDS

enteric valve armature are conical and shorter than the
least width of ridges three, while the differential
sclerotisation of the valve ridges is characteristic.

MATERIAL EXAMINED.

Democratic Republic of Congo: Katanga, Keyburg
near Elizabethville, 1948 single Paratype from Type
series, in the Natural History Museum, London
(Emerson). (Holotype imago and other Paratypes in
American Museum of Natural History, examined and
designated at the time of the first description).

Amicotermes gasteruptus sp. n.
(Figs 83-98, 179)

Head capsule yellow-white or pale yellow, antennal
flagellum, colour of darkest part, yellow or pale yel-
low-brown, pronotum and legs yellow-white,
abdominal tergites hyaline.

Head capsule short oval near circular or oval, setae
sparse, the emergent, longest setae randomly scat-
tered. Fontanelle present as pale spot or absent. Fore
coxae without prominent larger setae or with four or
more prominent larger setae arranged in line or group
on anterior surface; fore tibial apical spurs numbering
three, third well developed, at least half the size of the
others or much smaller than the other two.

Left mandible, first marginal tooth with anterior
edge distinctly longer than posterior or approximately
equilateral, third marginal tooth with anterior edge
shorter than that of first, fourth marginal tooth in front
view with proximal end clear of or just reaches side of
molar prominence. Right mandible, anterior edge of
first marginal tooth shorter.

Junction of mesenteron and proctodeum starting to
left of or beneath nerve cord in ventral view, proctodeal
first segment shorter, up to eight times proximal width,
tubular throughout its length; lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view; enteric
valve ‘seating’ third lobe smaller than outer pair.

Enteric valve ridges, distinctly ‘waisted’ at base of
apical spines, then wider, sclerotised at distal ends,
pale brown or yellow-brown; main armature with very
elongated tapered spines, the largest apical spines on
ridges one and three numbering 10-20, those on ridges
two and four numbering 1—9; longest spines curved
but still evenly tapered, longer than least width of
ridges in position three at base of spines, symmetry of
enteric valve armature more or less tri-radial, alternate
ridges reduced; subsidiary armature on one or more of
enteric valve ridges with scattered, somewhat sparse
prominent spines different from or smaller than main
armature, on scales or reticulations with indistinct
vague or incomplete outlines, the spines being gradu-
ated, becoming longer towards distal end of ridge;
membranous wall of enteric valve between and beyond

REVIEW OF AMICOTERMES

ridges scaly, armed with numerous but not dense,
scattered, minute spines or spicules fringing scales
proximally.

Measurements and diagnostic ratios [11 individuals

from 5 series]:

Head capsule width 0.598 to 0.635 mm (mean 0.620
SD .011).

Postclypeus width 0.301 to 0.339 mm (mean 0.322 SD
0.013).

Postclypeus length 0.166 to 0.205 mm (mean 0.184
SD 0.013).

Left mandible index 0.751 to 0.935 (mean 0.818).

Right mandible index 0.895 to 1.228 (mean 1.015).

Right second marginal index 0.796 to 1.097 (mean
0.987).

Right first to second marginal index 1.561 to 1.838
(mean 1.705).

Pronotum width 0.390 to 0.424 mm (mean 0.408 SD
0.01).

Pronotum length 0.168 to 0.246 mm (mean 0.200 SD
0.020).

Hind tibia length 0.597 to 0.6436 mm (mean 0.622 SD
0.014).

Fore tibia length 0.495 to 0.535 mm (mean 0.518 SD
0.011).

Fore tibia thickness 0.131 to 0.154 mm (mean 0.139
SD 0.006).

Fore tibial index (length over width) 3.393 to 3.876
(mean 3.726).

Left mandible, complex ratio La divided by L1.Lm
20.184 to 28.664 (mean 23.214).

Right mandible, complex ratio Ra divided by R1.Rm
20.67 to 30.394 (mean 24.89).

COMPARISONS. A. gasteruptus has already been com-
pared with the preceding species. It forms a terminal
pair with A. autothysius in all of the trees apart from
the Saitou and Nei tree and is close to it in the
ordination. A. ivorensis is a little larger and has the
front edge of the right first marginal tooth equal to that
of the second; the apical spines of the enteric valve
main armature are fewer in number on ridges one and
three, while the subsidiary armature is very sparse. A.
mayombei appears closely related, but the front edge
of the left third marginal is roughly equal to that of the
first; the longest apical spines of the enteric valve main
armature are shorter than or equal to the least width of
ridges three and on ridge one number less than ten, the
valve symmetry being bilateral. A. mbalmayoensis is
much larger and the longest spines of the main enteric
valve armature are shorter than or equal to the least
width of ridges three, while the subsidiary armature
consists of small single spines on scales, the sym-
metry being hexa-radial. A. multispinus is very slightly
larger and has more numerous apical spines on ridges
one and three of the enteric valve main armature.

161

A. spiculatus is much larger with proportionately longer
legs; its enteric valve ridges are ellipsoid, not ‘waisted’
and bear straight apical spines, while the symmetry of
the valve is bilateral owing to differential sclerotisation
of the ridges.

MATERIAL EXAMINED. _ [total 9 series]

Holotype worker and Paratypes from Type series (field
collection no. 38P-1) Cameroon: Mbalmayo Forest
Reserve, Ebogo, Feb. 1996 (Eggleton et al.); other
Paratype material, Eboufek, 5 series, Bilik, July—Aug.
1992, Ekombitie, Noy. 1995, Ebogo, Mar. 1996
(Eggleton et al.) (all in BMNH).

Amicotermes ivorensis sp. n.
(Figs 99-110, 180)

Head capsule, antennal flagellum, colour of darkest
part, pronotum and legs, pale yellow, abdominal tergites
yellow-white.

Head capsule circular or short oval near circular,
setae sparse, emergent, longest setae randomly scat-
tered. Fontanelle present as pale spot or small
depression. Fore coxae without prominent larger setae
on anterior surface, fore tibial apical spurs numbering
three, but third vestigial or much smaller than the other
two.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
shorter than that of first, fourth marginal tooth in front
view with proximal end clear of molar prominence.
Right mandible, anterior edge of first marginal tooth
equal in length to that of second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width,
tubular throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe approximately equal in size
to outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines or distinctly ‘waisted’ at their
base, then wider, all ridges weakly sclerotised distally,
pale yellow or yellow, main armature with very elon-
gated tapered spines, the largest on each ridge
numbering 1—9, straight or curved but still evenly
tapered, longer than least width of ridges in position
three at base of spines, symmetry of enteric valve
armature more or less tri-radial, alternate ridges re-
duced; subsidiary armature on one or more of enteric
valve ridges with backwardly directed indistinct scales,
outlines vague or incomplete, fringed with very sparse
small spines or spicules or small single backwardly
directed spines, the latter graduated, becoming longer
towards distal end of ridge; membranous wall of enteric

162

valve between and beyond ridges smooth, armed with
very sparse scattered minute spicules.

Measurements and diagnostic ratios {3 individuals
from 1| series]:

Head capsule width 0.657 to 0.713 mm (mean 0.684
SD 0.028).

Postclypeus width 0.359 to 0.381 mm (mean 0.368 SD
0.011).

Postclypeus length 0.205 to 0.241 mm (mean 0.227
SD 0.019).

Left mandible index 0.824 to 0.953 (mean 0.889).

Right mandible index 0.947 to 1.149 (mean 1.061).

Right second marginal index 0.839 to 1.136 (mean
0.969).

Right first to second marginal index 1.743 to 1.998
(mean 1.829).

Pronotum width 0.429 to 0.468 mm (mean 0.445 SD
0.020).

Pronotum length 0.206 to 0.293 mm (mean 0.252 SD
0.044).

Hind tibia length 0.665 to 0.692 mm (mean 0.680 SD
0.014).

Fore tibia length 0.525 to 0.549 mm (mean 0.540 SD
0.014).

Fore tibia thickness 0.135 to 0.148 mm (mean 0.142
SD 0.007).

Fore tibial index (length over width) 3.696 to 3.890
(mean 3.818).

Left mandible, complex ratio La divided by L1.Lm
21.554 to 24.308 (mean 22.474).

Right mandible, complex ratio Ra divided by R1.Rm
23.91to 27.495 (mean 25.224).

COMPARISONS. A. ivorensis has already been com-
pared with the preceding species. It stands somewhat
isolated in the ordination, but is closest to A. galenus
and A. mbalmayoensis in the consensus tree while
variously attached in the other, phenetic trees. A.
mayombei has prominent fore coxal setae, the left
third margional tooth with its front edge equal in
length to that of the first and the fourth marginal just
reaching the side of the molar prominence, while the
front edge of the right first marginal is shorter than
that of the second; the longest apical spines of the
enteric valve main armature are curved and shorter
than or equal to the least width of ridges three, the
valve being bilaterally symmetrical. A. mbalmayoensis
is considerably larger with much longer legs, while
the proximal end of the left fourth marginal tooth just
reaches or is hidden behind the molar prominence
and the front edge of the right first marginal is shorter
than that of the second; the longest apical spines of
the enteric valve main armature are more numerous
and shorter than the least width of ridges three, the
valve being hexa-radially symmetrical. A. multispinus

W.A. SANDS

has prominent fore coxal setae, the left third marginal
tooth with its front edge equal in length to that of the
first and the fourth marginal just reaching the side of
the molar prominence, while the front edge of the
right first marginal is shorter than that of the second;
the ridges of the main enteric valve armature have
more numerous apical spines. Finally, A. spiculatus
has much longer legs with prominent fore coxal setae
and the left third marginal tooth with its front edge
equal in length to that of the first, while the front edge
of the right first marginal is shorter than that of the
second; the largest apical spines of the main enteric
valve armature are conical, shorter than the least width
of ridges three, and more numerous on a bilaterally
symmetrical valve with differential sclerotisation of
the ridges.

MATERIAL EXAMINED. [one series]

Holotype worker and Paratypes from Type series,
Ivory Coast: near Ndouci in forest, 1969 (Josens) (all
in BMNH).

Amicotermes mayombei sp. n.
(Figs 111-124, 181, 182)

Head capsule, antennal flagellum, pronotum and legs
yellow-white, abdominal tergites hyaline.

Head capsule short oval near circular or oval, setae
sparse, emergent, longest setae regularly and sym-
metrically arranged. Fontanelle absent. Fore coxae
with four or more prominent larger setae arranged in
line or group on anterior surface, fore tibial apical
spurs numbering three, third well developed, at least
half the size of the others.

Left mandible, first marginal tooth with anterior
edge distinctly longer than posterior, third marginal
tooth with anterior edge equal in length to that of first,
fourth marginal tooth in front view just reaches side of
molar prominence. Right mandible, anterior margin of
first marginal tooth shorter than that of second.

Junction of mesenteron and proctodeum starting to
left of or beneath nerve cord in ventral view proctodeal
first segment shorter, up to eight times proximal width,
tubular throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe smaller than outer pair.

Enteric valve ridges, distinctly ‘waisted’ at base of
apical spines, then wider, sclerotised at distal end, pale
brown or yellow-brown with very elongated tapered
spines, the largest on ridges one, two and four number-
ing up to 10, those on ridges three 10—20, longest
spines curved but still evenly tapered, shorter than or
equal to least width of ridges in position three, symme-
try of enteric valve armature bilateral due to marked
differences in spine and ridge size; subsidiary arma-
ture on one or more of enteric valve ridges with

REVIEW OF AMICOTERMES

scattered, somewhat sparse prominent spines different
from or smaller than main armature, graduated, spines
becoming longer towards distal end of ridge, scales or
reticulations well defined, distinct; membranous wall
of enteric valve between and beyond ridges scaly
armed with short spicules, dense distally, fringing
scales proximally.

Measurements and diagnostic ratios [14 individuals
from 6 series]:

Head capsule width 0.614 to 0.713 mm (mean 0.651
SD 0.028).

Postclypeus width 0.322 to 0.373 mm (mean 0.344 SD
0.014).

Postclypeus length 0.162 to 0.214 mm (mean 0.185
SD 0.014).

Left mandible index 0.772 to 0.984 (mean 0.888).

Right mandible index 0.921 to 1.307 (mean 1.138).

Right second marginal index 0.936 to 1.169 (mean
1.047).

Right first to second marginal index 1.461 to 1.932
(mean 1.638).

Pronotum width 0.382 to 0.448 mm (mean 0.408 SD
0.018).

Pronotum length 0.175 to 0.22 mm (mean 0.191 SD
0.012).

Hind tibia length 0.637 to 0.736 mm (mean 0.683 SD
0.027).

Fore tibia length 0.499 to 0.571 mm (mean 0.542 SD
0.019).

Fore tibia thickness 0.133 to 0.145 mm (mean 0.139
SD 0.005).

Fore tibial index (length over width) 3.759 to 4.028
(mean 3.912).

Left mandible, complex ratio La divided by L1.Lm
18.707 to 26.77 (mean 22.184).

Right mandible, complex ratio Ra divided by R1.Rm
21.009 to 29.644 (mean 25.813).

COMPARISONS. A. mayombei has already been com-
pared with the preceding species. In most of the
phenetic trees it joins A. autothysius and A. gasteruptus
at a slightly lower phenon level and in the strict
- consensus tree it links with them at node 14. A.
mbalmayoensis is considerably larger and lacks promi-
nent fore coxal setae while the left first marginal
tooth is approximately equilateral; The enteric valve
ridges are tapering ellipsoid bearing straight spines
and the valve is hexa-radially symmetrical. A.
multispinus is a little smaller and has the left first
marginal tooth aproximately equilateral; the longest
apical spines of the main enteric valve armature are
more numerous on ridges one and three and longer
than the least width of three, while the symmetry of
the valve is tri-radial. In A. spiculatus the legs are
distinctly longer, the left first marginal tooth is

163

equilateral and the fourth marginal is clear of the
molar prominence; the enteric valve ridges are taper-
ing ellipsoid, not ‘waisted’ and differentially
sclerotised, ridge one being colourless while ridges
three and four are pale brown, two being intermedi-
ate; the largest apical spines are straight and conical.

MATERIAL EXAMINED. [total 5 series]

Holotype worker and Paratypes from Type series (field
collection no. T1.10) Congo (Brazzaville): Mayombe,
Feb. 1997 (Eggleton and Davies); other Paratype ma-
terial, Feb.1997, 4 series (Eggleton and Davies) (all in
BMNH).

Amicotermes mbalmayoensis sp. n.
(Figs 125-137, 183, 184)

Head capsule yellow-white or pale yellow, antennal
flagellum, colour of darkest part, yellow or pale yellow-
brown, pronotum and legs yellow-white, abdominal
tergites hyaline.

Head capsule short oval near circular, setae numer-
ous but not dense, emergent, longest setae regularly
and symmetrically arranged. Fontanelle absent. Fore
coxae without prominent larger setae on anterior sur-
face, fore tibial apical spurs numbering three, third
well developed, at least half the size of the others.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
shorter than or equal in length to that of first, fourth
marginal tooth in front view just reaches side of or
with proximal end hidden behind molar prominence.
Right mandible, anterior edge of first marginal tooth
shorter than that of second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width
tubular throughout its length, lateral dis-placement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe smaller than outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines, weakly sclerotised distally,
pale yellow or yellow, main armature with very elon-
gated tapered spines, the largest on each ridge
numbering 10-14, straight and evenly tapered, shorter
than or equal to least width of ridges in position three;
subsidiary armature on one or more of enteric valve
ridges with scattered, somewhat sparse small single
backwardly directed spines on scales, more or less
uniform throughout length of ridge, scales or
reticulations indistinct, outlines vague or incomplete;
symmetry of enteric valve armature more or less hexa-
radial apart from slight differences in ridge size;
membranous wall of enteric valve between and be-
yond ridges scaly, armed with minute spicules, dense
distally, fringing scales proximally.

164

Measurements and diagnostic ratios {9 individuals
from 3 series]:

Head capsule width 0.765 to 0.800 mm (mean 0.781
SD 0.013).

Postclypeus width 0.413 to 0.434 mm (mean 0.424 SD
0.008).

Postclypeus length 0.199 to 0.239 mm (mean 0.215
SD 0.012).

Left mandible index 0.896 to 0.978 (mean 0.944).

Right mandible index 1.065 to 1.366 (mean 1.174).

Right second marginal index 0.88 to 1.245 (mean
1.013).

Right first to second marginal index 1.606 to 1.987
(mean 1.811).

Pronotum width 0.523 to 0.582 mm (mean 0.545 SD
0.019).

Pronotum length 0.230 to 0.267 mm (mean 0.252 SD
0.013).

Hind tibia length 0.779 to 0.82 mm (mean 0.796 SD
0.014).

Fore tibia length 0.614 to 0.677 mm (mean 0.651 SD
0.019).

Fore tibia thickness 0.159 to 0.173 mm (mean 0.166
SD 0.005).

Fore tibial index (length over width) 3.602 to 4.078
(mean 3.916).

Left mandible, complex ratio La divided by L1.Lm
22.319 to 27.038 (mean 23.941).

Right mandible, complex ratio Ra divided by R1.Rm
20.164 to 26.881 (mean 24.080).

COMPARISONS. A. mbalmayoensis has already been
compared with the preceding species. A. multispinus 1s
much smaller and has prominent fore coxal setae; the
front edge of the left third marginat tooth is equal in
length to that of the first and the right mandible first to
second marginal index is lower; the enteric valve
ridges are ‘waisted’ at the base of the apical spines
which are curved, longer than the least width of ridges
three and more numerous on ridges one and three, the
valve being tri-radial. A. spiculatus is smaller with a
lower left mandible index; the apical spines of the
enteric valve main armature are conical and more
numerous on ridges three which are differentially
sclerotised along with four, while one is unsclerotised
and two intermediate, the valve being bilaterally sym-
metrical.

MATERIAL EXAMINED. _ [total, 3 series]

Holotype worker and Paratypes from Type series (field
collection no. H,P.2) Cameroon: Mbalmayo Forest
Reserve, Ekombitie, Nov. 1996 (Eggleton et al.); other
Paratypes, Ekombitie, Nov. 1995, Ebogo, Mar. 1996
(Eggleton et al.) (all in BMNH).

W.A. SANDS
Amicotermes multispinus sp. n.
(Figs 138-150, 185)

Head capsule, antennal flagellum, colour of darkest
part, pronotum and legs, yellow-white, abdominal
tergites hyaline.

Head capsule short oval near circular, setae sparse,
emergent, longest setae regularly and symmetrically
arranged. Fontanelle absent. Fore coxae with four or
more prominent larger setae arranged in line or group
on anterior surface, fore tibial apical spurs numbering
three, third well developed, at least half the size of the
others.

Left mandible, first marginal tooth approximately
equilateral, third marginal tooth with anterior edge
equal in length to that of first, fourth marginal tooth in
front view just reaches side of molar prominence.
Right mandible, anterior edge of first marginal tooth
shorter than that of second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width,
tubular, throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe smaller than outer pair.

Enteric valve ridges, distinctly ‘waisted’ at base of
apical spines, then wider, weakly sclerotised distally,
pale yellow or yellow, main armature with very elon-
gated tapered spines, the largest on ridge one numbering
15-20, those on ridges two and four numbering 1-9,
those on ridges three numbering 15 to over 20, the
longest, curved but still evenly tapered, longer than
least width of ridges in position three at base of spines,
symmetry of enteric valve armature more or less tri-
radial, alternate ridges reduced; subsidiary armature
on one or more of enteric valve ridges with scattered,
somewhat sparse prominent spines different from or
smaller than main armature, graduated, spines becom-
ing longer towards distal end of ridge, scales or
reticulations indistinct, outlines vague or incomplete;
membranous wall of enteric valve between and be-
yond ridges scaly, armed with numerous but not dense,
scattered, short spicules fringing scales proximally.

Measurements and diagnostic ratios [3 individuals
from | series]:

Head capsule width 0.642 to 0.647 mm (mean 0.645).
Postclypeus width 0.334 to 0.356 mm (mean 0.342).
Postclypeus length 0.183 to 0.191 mm (mean 0.186).
Left mandible index 0.802 to 0.903 (mean 0.852).
Right mandible index 1.049 to 1.163 (mean 1.100).
Right second marginal index 0.924 to 1.021 (mean
0.974).
Right first to second marginal index 1.569 to 1.594
(mean 1.581).

REVIEW OF AMICOTERMES

Pronotum width 0.412 to 0.436 mm (mean 0.427).

Pronotum length 0.183 to 0.191 mm (mean 0.186).

Hind tibia length 0.624 to 0.629 mm (mean 0.626).

Fore tibia length 0.525 to 0.531 mm (mean 0.528).

Fore tibia thickness 0.134 to 0.141 mm (mean 0.138).

Fore tibial index (length over width) 3.732 to 3.974
(mean 3.834).

Left mandible, complex ratio La divided by L1.Lm
23.999 to 27.622 (mean 25.824).

Right mandible, complex ratio Ra divided by R1.Rm
23.347 to 26.271 (mean 24.767).

COMPARISONS. A. multispinus has already been com-
pared with all of the preceding species, only A.
spiculatus remaining. The latter is slightly larger and
has the left fourth marginal tooth clear of the molar
prominence; the enteric valve ridges are tapering ellip-
soid, not ‘waisted’, and differentially sclerotised with
fewer, straight conical spines on ridges one and three,
the valve being bilaterally symmetrical.

MATERIAL EXAMINED. [one series]

Holotype worker and Paratypes from Type series (field
collection no. T1.20) Congo (Brazzaville): Mayombe,
Feb. 1997 (Eggleton and Davies) (all in BMNH).

Amicotermes spiculatus sp. n.
(Figs 151-161, 186)

Head capsule pale yellow, antennal flagellum, colour
of darkest part pale yellow or yellow, pronotum and
legs yellow-white, abdominal tergites hyaline.

Head capsule oval, setae sparse, emergent, longest
setae regularly and symmetrically arranged. Fonta-
nelle absent. Fore coxae with three or more prominent
larger setae arranged in line or group on anterior
surface, fore tibial apical spurs numbering three, third
well developed, at least half the size of the others. Left
mandible, first marginal tooth approximately equilat-
eral, third marginal tooth with anterior edge equal in
length to that of first, fourth marginal tooth in front
view with proximal end clear of molar prominence.
Right mandible, anterior edge of first marginal tooth
shorter than that of second.

Junction of mesenteron and proctodeum starting
beneath nerve cord in ventral view, proctodeal first
segment shorter, up to eight times proximal width,
tubular throughout its length, lateral displacement of
enteric valve in unopened abdomen to right of heart in
dorsal or to left of nerve cord in ventral view, enteric
valve ‘seating’ third lobe smaller than outer pair.

Enteric valve ridges, more or less tapering ellipsoid
to base of apical spines, enteric valve ridge in position
one unsclerotised or colourless throughout, positions

~ two and four weakly sclerotised distally, pale yellow

or yellow, position three sclerotised at distal end, pale
brown or yellow-brown main armature of all ridges
with large backwardly directed erect conical spines,

165

those on ridges one and two numbering up to 14, on
ridges three, 15—20 and on ridge four, 1—9, the longest
being straight and evenly tapered, shorter than or
equal to least width of ridges in position three, symme-
try of enteric valve armature bilateral due to marked
differences in spine and ridge size; subsidiary arma-
ture on one or more of enteric valve ridges with
prominent spines different from or smaller than main
armature, covering entire ridge surface, graduated,
spines or spicules becoming longer towards distal end
of ridge, scales or reticulations indistinct, outlines
vague or incomplete; membranous wall of enteric
valve between and beyond ridges scaly, armed with
short scattered spicules, numerous but not dense, fring-
ing scales proximally.

Measurements and diagnostic ratios [only two speci-
mens available]:

Head capsule width 0.696 to 0.706 mm.

Postclypeus width 0.370 to 0.379 mm.

Postclypeus length 0.203 to 0.206 mm.

Left mandible index 0.855 to 0.88.

Right mandible index 1.069 to 1.149.

Right second marginal index 1.016 to 1.029.

Right first to second marginal index 1.788 to 1.813.

Pronotum width 0.438 to 0.460 mm.

Pronotum length 0.233 to 0.239 mm.

Hind tibia length 0.784 to 0.794 mm.

Fore tibia length 0.620 to 0.625 mm.

Fore tibia thickness 0.15 to 0.152 mm.

Fore tibial index (length over width) 4.119 to 4.135.

Left mandible, complex ratio La divided by L1.Lm
20.586 to 23.569.

Right mandible, complex ratio Ra divided by R1.Rm
24.213 to 25.340.

COMPARISONS. All the necessary comparisons of A.
spiculatus have been made under the preceding spe-
cies.

MATERIAL EXAMINED. _ [one series]

Holotype worker and one Paratype (field collection
no. CHR 35-40) Cameroon: Ebolowa, Akok, June
1997, (Eggleton et al.) (all in BMNH).

PHENETIC AND PHYLOGENETIC
ANALYSIS

The purpose in undertaking these analyses was to
examine the apparent relationships between the twelve
species of the genus now recognised and their geo-
graphical distribution. Phenetics, that is the study of
relationships based on overall similarity between taxa,
has largely gone out of use in favour of cladistics
because the methods take no account of homoplasy.

166

However, in a small, relatively homogeneous set of
species such as this, where convergence was thought
unlikely to be a major feature, they offered an alterna-
tive way of looking at the character matrix. In practice,
it can be seen from the relatively low values of the
consistency and retention indices of the Hennig strict
consensus tree, and the character states marked on it
that there was more homoplasy than might have been
expected. The precise topography of the individual
trees depends on differences in the algorithms that
define them, and there is no clear ‘best’ tree, although
some taxonomists would claim this role for the Hennig
tree. Where a cluster of species is robust through
several methods including the cladistic format, it seems
likely that the underlying relationships are more clearly
illustrated.

The DELTA file of Amicotermes was converted by
means of the PANKEY programmes SC3 and
DELPAUPI respectively into a matrix of similarity
coefficients for input to NTSYS (Rohlf, 1993)
phenetics and Principal Coordinates analyses and a
coded character matrix for HENNIG86 (Farris 1988)
phylogenetic analysis. Acholotermes was included as
an outgroup, being the most closely related other
genus. The first analyses were based on the original
DELTA file, but because that included ranges of vari-
ation in the character codings, many of the characters
were represented by queries in the HENNIG matrix.
Since measurements had proved important in distin-
guishing some of the species, it was undesirable they
should be inactive in the study of relationships. In
order to avoid this, the variable characters were re-
coded to their median (in multi-states) or mean values
(in continuous variables). In the latter case, the DELTA
file included the facility to allocate measurements or
ratios to KEY STATES, equal sub-divisions of the
total range of each variable. It was observed that for
these, closely similar members of the same genus the
variances were roughly equal and it was not thought
necessary to standardise the data. In practice this
amounted to a crude form of gap coding for the
morphometrics. Most of the characters then became
fully active, but the effects of this procedure turned out
to be comparatively slight in terms of the branching of
trees and the positions of terminal taxa. The results of
both phylogenetic and phenetic analysis are shown in
Figs 162-168.

The strict consensus tree derived from the most
parsimonious two Hennig trees (Fig. 162) has essen-
tially the same topology as one of them. The states of
those characters with retention indices greater than 50
are marked on the tree at each node; in addition, the
tree also indicates the autapomorphies of individual
taxa on their stems. It places A. dibogi separated from
the rest of the genus and close to the outgroup,
Acholotermes, with a considerable range of
autapomorphies. The character states shown at node

W.A. SANDS

24 are the plesiomorphic states of the characters con-
cerned on this particular tree. Succeeding single
branches from low on the tree are occupied by
A. camerunensis, A. spiculatus and A. cristatus, which
show progressive development and localisation of
straight spiny armature of the enteric valve. The rest of
the species appeared to divide into two groups, one of
three (A. mbalmayoensis, A. ivorensis and A. galenus)
and one of five (A. multispinus, A. congoensis,
A. mayombei, A. gasteruptus and A. autothysius), the
two last named forming a terminal pair, with
A. congoensis and A. multispinus joining the group at
slightly lower levels. The lower nodes of the tree are
determined largely by features of the main armature of
the enteric valve, while above node 19, measurement
characters predominate. However, the shape of the
enteric valve ridges (‘waisted’, 23a) and the type of
spines in the main armature (elongated, curved taper,
36a) are important synapomorphies at node 18 that
define the group of five species mentioned above; they
also suggest a trend towards certain other genera such
as Ateuchotermes and perhaps Apagotermes.

The Principal Coordinates plot (Fig. 163) shows the
positions of the taxa in relation to the first three
eigenvectors of the similarity matrix transformed by
double centering. A. dibogi is again widely separated
on vectors one and two from the other species and
placed near to the outgroup. A. camerunensis,
A. spiculatus and A. cristatus are separated on vectors
two and three while most of the separation of
A. mbalmayoensis, A. ivorensis and A. galenus 1s ac-
counted for by vector three. A. congoensis is also
separated mainly on V3 while the remaining two pairs
are relatively close together on all three vectors. The
minimum spanning tree is superimposed on the ordi-
nation in order to show up any local distortions. It links
the outgroup to A. dibogi which in turn is linked to
A. autothysius, A. gasteruptus and A. mayombei in
series. It then branches to A. cristatus, A. spiculatus
and A. camerunensis in turn and to A. multispinus and
A. congoensis, with further branches to A. mbalmayo-
ensis and A. galenus, which is finally linked to
A. ivorensis.

The various SAHN phenetic clustering methods,
(Single (Fig. 164), Flexible (Fig. 165) and Complete
(Fig. 166) Linkage methods and Unweighted pair-
group method of averages (Fig. 167)) based on the
matrix of similarity coefficients give comparable re-
sults, allowing for a small amount of branch-swapping,
in terms of the terminal groups of taxa, to the
HENNIG86 (Farris, 1988) tree. A. dibogi is always
separate and near to Acholotermes, the successive
additions at lower levels are largely similar, and the
two terminal groups also retain their cohesion to a
certain extent. The Single-linkage tree, with its ten-
dency to form long chain clusters, has the
A. autothysius—A. gasteruptus—A. congoensis—A. may-

REVIEW OF AMICOTERMES

ombei-A. multispinus group but adds in A. galenus at
about the 0.57 phenon level. The other group has
broken up with A. ivorensis and A. mbalmayoensis
separated; A. spiculatus and A. cristatus unite at the
0.53 phenon level. Both Flexible and Complete link-
age methods keep A. autothysius—A. gasteruptus—
A. mayombei together at about 0.55 phenon level and
also A. congoensis—A. multispinus—A. galenus at 0.50
phenons. A. spiculatus and A. cristatus similarly com-
bine, but Flexible linkage adds A. camerunensis at
about 0.35 phenons and Complete linkage adds
A. ivorensis. The UPGMA tree keeps A. autothysius—
A. gasteruptus—A. mayombei together at about 0.58
phenon level and also A. congoensis—A. multispinus—
A. galenus at 0.55 phenons; A. spiculatus and
A. cristatus combine at 0.53, and A. camerunensis at
0.42. A strict consensus tree of the Complete and
Flexible linkage methods only leaves the A. mayombei—
A. gasteruptus—A. autothysius group recognisable, the
rest being reduced to a comb and nothing remains if
Single linkage is added in, while that of the UPGMA
and Neighbour-joining trees apparently also leaves no
grouping in common.

The only exception to the above arrangement oc-
curs in the Saitou and Nei (1987) Neighbour-joining
tree (Fig. 168), which uses an algorithm based on the
principle of parsimony to produce from the dissimilar-
ity matrix a ‘phylogenetic’ tree. Here the species
closest to the outgroup is A. gasteruptus, while
A. dibogi forms a terminal pair high up the tree with
A. cristatus. Considering how far apart these two ap-
pear in the ordination of the first three vectors, it might
be concluded that they come to be ‘neighbours’ as a
result of closeness on other, lower level vectors that
cannot be plotted; however, this is not borne out by
their positions at almost the extreme ends on the
minimum spanning tree, which agrees with both the
cladistic and the other phenetic classifications in plac-
ing A. dibogi next to the outgroup. None of the terminal
groups recognised in the other methods remains.

The results of the HENNIG, SAHN and ordination
analyses all share aspects with an intuitive view of the
probable relationships of most of the species. Perhaps
the most interesting feature that is found in all of the
trees apart from the Saitou and Nei tree is the distribu-
tion of taxa from each of the two regions sampled
among the terminal groups. Each fauna appears to
include representatives of the different trends within
the genus, and it would be interesting to see whether
this were to be maintained through further sampling
on a wider geographical basis. A surprise is the appar-
ently close relationship of A. dibogi to the outgroup
which must be a reflection of its relatively
unspecialised, hexa-radially symmetrical, finely spiny
and spiculate enteric valve armature. Other, appar-
ently derived features of the gut, such as the elongated
and dilated P1, which pushes the enteric valve seating

167

into a dorsal position, only account for three ‘charac-
ters’ out of the total set of 60 and they appear as
autapomorphies on the tree. The Saitou and Nei is the
only tree to place A. dibogiin a derived position distant
from Acholotermes.

ACKNOWLEDGEMENTS [| am grateful to Dr Paul Eggleton
for reading and commenting on the manuscript of this paper,
and to my colleagues in the Termite Group at The Natural
History Museum for helpful discussions and testing the key to
species. Thanks are also due to the Trustees of The Natural
History Museum for permission to examine specimens in the
collection.

REFERENCES

Dallwitz, M. J. and Paine, T. A. 1986. User's guide to the DELTA
System: A General System for Processing Taxonomic Descriptions,
3rd. edn. Division of Entomology Report No. 13, CSIRO, Can-
berra, ACT 2601, Australia

Eggleton, P., Bignell, D. E., Sands, W. A., Mawdsley, N. A.,
Lawton, J. H., Wood, T. G. & Bignell, N. C. 1996. The diversity,
abundance and biomass of termites under differing levels of distur-
bance in the Mbalmayo Forest Reserve, southern Cameroon.
Philosophical Transactions of the Royal Society of London B, 351:
51-68.

Farris, J. S. 1988. HENNIGS6 Reference, Version 1.5. Photocopied
handbook, author’s copyright, 18 pp.

Pankhurst, R. J. 1986. A package of computer programs for han-
dling taxonomic databases. Cabios, 2: 33-39.

Pankhurst, R. J. 1991. Practical Taxonomic Computing. Cambridge
University Press, Cambridge, 202 pp.

Partridge, T. R., Dallwitz, M. J.,and Watson, L. 1986. A Primer for
the DELTA System on VMS, MS-DOS, and PRIMOS. Division of
Entomology Report No. 38, CSIRO, Canberra ACT 2601, Aus-
tralia.

Rohlf, F. J. 1993. NTSYS-pc Numerical Taxonomy and Multivariate
Analysis System, Version 1.8., Exeter Software, Setauket, New
York 11733 USA.

Saitou, N. & Nei, M. 1987. The neighbour-joining method: a new
method for reconstructing phylogenetic trees. Molecular Biology
and Evolution, 4: 406-425.

Sands, W. A. 1965. A revision of the termite subfamily
Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Re-
gion. Bulletin of the British Museum (Natural History), Entomology
Supplement, 4: 1-172.

Sands, W. A. 1965a. Termite distribution in man-modified habitats in
West Africa with special reference to species segregation in the
genus Trinervitermes. Journal of Animal Ecology, 34: 557-571.

Sands, W. A. 1972. The soldierless termites of Africa (Isoptera:
Termitidae). Bulletin of the British Museum (Natural History),
Entomology Supplement, 18: 1-244.

Sands, W. A. 1982. Agonistic behaviour of African soldierless
Apicotermitinae (Isoptera: Termitidae). Sociobiology, 7 (1): 61—
TANS

Sands, W. A. 1992. The Termite Genus Amitermes in Africa and the
Middle East. Natural Resources Institute Bulletin, 51: 1-140.

Sands, W. A. 1995. New genera and species of soil feeding termites
(Isoptera, Termitidae) from African savannas. Journal of Natural
History, 29: 1483-1515.

Sands, W. A. 1998. The identification of worker castes of termite
genera from soils of Africa and the Middle East. CAB International
Wallingford, Oxon. 500 pp., 1586 figs, 18 pls.

168

APPENDIX

The entire DELTA file is shown below, except that the
character states recorded for the species are omitted, since
they have been converted to keys and descriptions. The
symbols for character types are OM, ordered multistate and
RN, real numbers, the default being unordered multistate.
The format is rigid in that certain lines must end ina slash, and
every line begins with a single space. the numbers of charac-
ters, states and items stated at the beginning must correspond
exactly to those included in the file

*HEADING THE AFRICAN SOLDIERLESS TERMITE
GENUS AMICOTERMES/

*SPECIAL STORAGE

*KEY OPTIONS OLDKEY PARTIAL /

*NUMBER OF CHARACTERS 60

*MAXIMUM NUMBER OF STATES 6

*MAXIMUM NUMBER OF ITEMS 13

*CHARACTER TYPES

1,OM 2,0M 3,OM 4, OM 5,0OM 6, OM 11, OM 14, OM 16,
OM 18, OM 24, OM 25, OM 26, OM 27, OM 28, OM 29, OM
30, OM 31, OM 32, OM 33, OM 34, OM 35, OM 38, OM
40, OM 42, OM 45, OM 46, RN 47, RN 48, RN 49, RN 50,
RN 51, RN 52, RN 53, RN 54, RN 55, RN 56, RN 57, RN 58,
RN 59, RN 60, RN

*NUMBERS OF STATES

1,42,43,34,35,36,3 11,3 14,3 15,3 16,3 18,3 24, 3 25,
3 26, 3 27, 3 28, 3 29, 3 30, 3 31, 3 32, 4 33, 4 34, 4 35, 4 38,
5 40, 3 42, 3 45, 3

*KEY STATES

46, 0.565—0.604/0.605—0.644/0.645—0.683/0.684—0.722/
0.723—-0.761/0.762—0.800

47, 0.300—0.333/0.334—0.367/0.368—0.401/0.402-0.434

48, 0.154—0.178/0.179-0.202/0.203—0.227/0.228—-0.250

49, 0.697-0.781/0.782—0.866/0.867—0.95 1/0.952—1.036/
1.037—1.12/1.121—1.205

50, 0.879-0.962/0.963—1.044/1.045—1.127/1.128—1.209/
1.21-1.291/1.292-1.374

51, 0.788—0.867/0.868—0.946/0.947—1.025/1.026—1.104/
1.105—1.183/1.184—1.262

52, 1.350-1.474/1.475—1.598/1.599—1.722/1.723-1.846/
1.847—1.97/1.971—2.094

53, 0.375—0.426/0.427-0.478/0.479-0.53/0.53 1—-0.582

54, 0.168-0.199/0.2—0.231/0.232—0.262/0.263—0.293

55, 0.565—0.629/0.63—0.693/0.694—0.756/0.757—0.820

56, 0.489-0.536/0.537—0.583/0.584—0.63/0.63 1—0.677

57, 0.127-0.147/0.148—0.166/0.167—0.185/0.186—0.205

58, 2.953—3.255/3.256—3.556/3.557—3.857/3.858—4.157/
4.158-4.458/4.459-4.759

59, 17.556—22.634/22.635—27.712/27.713—32.789/32.79—
37.867

60, 20.164—23.626/23.627—27.087/27.088-30.549/30.55—
34.010

*CHARACTER DESCRIPTIONS

#1. head capsule <colour>/

1. yellow-white/

2. pale yellow/

3. yellow/

4. pale yellow-brown/

W.A.SANDS

#2. antennal flagellum <colour of darkest part>/
1. yellow-white/

2. pale yellow/

3. yellow/

4. pale yellow-brown/

#3. pronotum <colour>/
1. yellow-white/

2. pale yellow/

3. yellow/

#4. legs <colour>/
1. yellow-white/
2. pale yellow/

3. yellow/

#5. abdominal tergites <colour>/
1. hyaline/

2. yellow-white/

3. pale yellow/

#6. head capsule <shape>/
1. circular/

2. short oval near circular/
3. oval/

#7. head capsule setae/
1. sparse/
2. numerous but not dense/

#8. head capsule emergent, longest setae/
1. randomly scattered/
2. regularly and symmetrically arranged/

#9. fontanelle/
1. present as pale spot or small depression/
2. absent/

#10. postclypeus <contour>/

1. clearly inflated/

2. strongly inflated, bulging, length more than half width at
ginglymi/

#11. fore coxae/

1. without prominent larger setae on anterior surface/

2. with one to three prominent larger setae on anterior surface/
3. with four or more prominent larger setae arranged in line or
group on anterior surface/

#12. fore tibial apical spurs/

1. numbering three, third well developed, at least half the size
of the others/

2. numbering three, but third much smaller than the other two/

#13. left mandible, first marginal tooth/
1. with anterior edge distinctly longer than posterior/
2. approximately equilateral/

#14. left third marginal tooth with anterior edge/
1. shorter than that of first/

2. equal in length to that of first/

3. longer than that of first/

REVIEW OF AMICOTERMES

#15. left fourth marginal tooth in front view/

1. with proximal end clear of molar prominence/

2. just reaches side of molar prominence/

3. with proximal end hidden behind molar prominence/

#16. anterior margin of right first marginal tooth/
1. longer than that of second/

2. equal in length to that of second/

3. shorter than that of second/

#17. right first marginal tooth with exposed posterior edge/
1. equal to that of second marginal/
2. longer than that of second marginal/

#18. junction of mesenteron and proctodeum/

1. starting to left of nerve cord in ventral view/
2. starting beneath nerve cord in ventral view/
3. starting to right of nerve cord in ventral view/

#19. proctodeal first segment <length>/
1. shorter, up to eight times proximal width/
2. very long, over eight times proximal width/

#20. proctodeal first segment beyond junction with mesenteron
<condition>/

1. tubular or conical throughout its length/

2. dilated to about twice its basal width throughout length/

#21. lateral displacement of enteric valve in unopened abdo-
men/

1. to left of heart in dorsal view/

2. to right of heart in dorsal or to left of nerve cord in ventral
view/

#22. enteric valve ‘seating’ third lobe <size>/
1. smaller than outer pair/
2. approximately equal in size to outer pair/

#23. enteric valve ridges, <shape>/
1. more or less tapering ellipsoid to base of apical spines/
2. distinctly ‘waisted’ at base of apical spines, then wider/

#24. enteric valve ridge in position one/

1. sclerotised at distal end, pale brown or yellow-brown/
2. weakly sclerotised distally, pale yellow or yellow/

3. unsclerotised or colourless throughout/

#25. enteric valve ridges in position two/

1. sclerotised at distal end, pale brown or yellow-brown/
2. weakly sclerotised distally, pale yellow or yellow/

3. unsclerotised or colourless/

#26. enteric valve ridges in position three/

1. sclerotised at distal end, pale brown or yellow-brown/
2. weakly sclerotised distally, pale yellow or yellow/

3. unsclerotised or colourless/

#27. enteric valve ridge in position four/

1. sclerotised at distal end, pale brown or yellow-brown/
2. weakly sclerotised distally, pale yellow or yellow/

3. unsclerotised or colourless/

169

#28. main armature of enteric valve ridge in position one/
1. without large apical spines/

2. with large backwardly directed erect conical spines/

3. with very elongated tapered spines/

#29. main armature of enteric valve ridges in position two/
1. without large apical spines/

2. with large backwardly directed erect conical spines/

3. with very elongated tapered spines/

#30. main armature of enteric valve ridges in position three/
1. without large apical spines/

2. with large backwardly directed erect conical spines/

3. with very elongated tapered spines/

#31. main armature of enteric valve ridge in position four/
1. without large apical spines/

2. with large backwardly directed erect conical spines/

3. with very elongated tapered spines/

#32. largest apical spines on ridge one <of main armature,
number>/

1. numbering 1—9/

2. numbering 10—14/

3. numbering 15—20/

4. numbering over 20/

#33. largest spines on ridges two <of main armature, number>/
1. numbering 1—9/

2. numbering 10—14/

3. numbering 15—20/

4. numbering over 20/

#34. largest spines on ridges three <of main armature,
number>/

1. numbering 1—9/

2. numbering 10—14/

3. numbering 15—20/

4. numbering over 20/

#35. largest spines on ridge four <of main armature, number>/
1. numbering 1—9/

2. numbering 10—14/

3. numbering 15—20/

4. numbering over 20/

#36. longest spines of main armature <type>/
1. straight and evenly tapered/
2. curved but still evenly tapered/

#37. longest apical spines <length>/

1. longer than least width of ridges in position three at base of
spines/

2. shorter than or equal to least width of ridges in position
three/

#38. subsidiary armature on one or more of enteric valve
ridges mainly/

1. reticulate/

2. with backwardly directed scales/

3. with backwardly directed scales fringed with small spines
or spicules/

170

4. with small single backwardly directed spines on scales/
5. with prominent spines different from or smaller than main
armature/

#39. subsidiary armature of enteric valve ridges/

1. with scales or reticulations well defined, distinct/

2. with scales or reticulations indistinct, outlines vague or
incomplete/

#40. subsidiary armature of one or more of enteric valve
ridges/

1. prominent, covering entire ridge surface/

2. scattered, somewhat sparse/

3. very sparse, few scales fringed with spines or spicules/

#41. subsidiary armature on one or more enteric valve ridges/
1. more or less uniform throughout length of ridge/

2. graduated, spines or spicules becoming longer towards
distal end of ridge/

#42. symmetry of enteric valve armature/

1. more or less hexa-radial apart from slight differences in
ridge size/

2. more or less tri-radial, alternate ridges reduced/

3. bilateral due to marked differences in spine and ridge size/

#43. membranous wall of enteric valve between and beyond
ridges/

1. smooth/

2. scaly/

#44. membranous wall of enteric valve between and beyond
ridges/

1. armed with minute spines or spicules/

2. armed with short spines or spicules/

#45. spines or spicules on membranous wall/

1. very sparse, scattered/

2. numerous but not dense, scattered, fringing scales proxi-
mally/

3. dense distally, fringing scales proximally/

#46. head capsule width <range, mean, SD>/ mm/

#47. postclypeus width <range, mean, SD>/ mm/

#48. postclypeus length <range, mean, SD>/ mm/

#49. left mandible index <range, mean>/

W.A. SANDS

#50. right mandible index <range, mean>/

#51. right second marginal index <range, mean>/

#52. right first to second marginal index <range, mean>/
#53. pronotum width <range, mean, SD>/ mm/

#54. pronotum length <range, mean, SD>/ mm/

#55. hind tibia length <range, mean, SD>/ mm/

#56. fore tibia length <range, mean, SD>/ mm/

#57. fore tibia thickness <range, mean, SD>/ mm/

#58. fore tibial index (length over width) <range, mean>/

#59. left mandible, complex ratio La divided by L1.Lm
<range, mean>/

#60. right mandible, complex ratio Ra divided by R1.Rm
<range, mean>/

*ITEM DESCRIPTIONS
#1.Acholotermes <outgroup>/
#2.Amicotermes autothysius sp. n./
#3.Amicotermes camerunensis sp. n./
#4.Amicotermes congoensis sp. n./
#5.Amicotermes cristatus sp. n./
#6.Amicotermes dibogi sp. n./
#7.Amicotermes galenus Sands 1972/
#8.Amicotermes gasteruptus sp. n./
#9.Amicotermes ivorensis sp. n./
#10.Amicotermes mayombei sp. n./
#11.Amicotermes mbalmayoensis sp. n./
#12.Amicotermes multispinus sp. n./

#13.Amicotermes spiculatus sp. n./

REVIEW OF AMICOTERMES

}
;

171

La i,
4 Oy

Siam

Mesenteron
(Me)

Malpighian
tubules (MT)

~) pee

Proctodeum knot (MK)
first section
(P41)

Enteric valve seating (EV)

Proctodeum
third section
(P3)

Rectum
(Re)

Figs 1-9. Amicotermes autothysius. 1-3, mandibles with surface view of right molar plate; 4, 6, ventral and 5, 7, dorsal
views of abdomen to show gut coiling in situ; 8, gut dissected and uncoiled; 9, crop with ‘bursters’ and proventriculus
[bar, 0.5 mm: top left, mandibles; 1 mm: bottom left, abdomens and uncoiled gut; bottom middle, crop and proventriculus]

Abbreviations in Figs 1 and 2 indicate standard measurements (see text).

W.A. SANDS

172

Amicotermes autothysius. 10-12, enteric valve seating and part of P3; 13-15, enteric valve armature opened

Figs 10-15.

valve seating; 0.5 mm: bottom right valve

>

out to show detail and variation, ridge positions numbered [bar, | mm: middle

armature].

REVIEW OF AMICOTERMES 173

Figs 16-28. Amicotermes camerunensis. 16-18, mandibles with surface view of right molar plate; 19, 21, ventral and 22,
24, dorsal views of abdomen to show gut coiling in situ; 23, gut dissected and uncoiled; 24, crop with ‘bursters’ and
proventriculus; 25—27, enteric valve seating and part of P3; 28, enteric valve armature opened out to show detail, ridge
positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: bottom right, abdomens and
uncoiled gut; bottom middle, valve seating and crop].

174 W.A. SANDS

Figs 29-38. Amicotermes congoensis. 29-31, mandibles with surface view of right molar plate; 32, 34, 36, ventral and 33,
35, 37, dorsal views of abdomen to show gut coiling in situ; 38, gut dissected and uncoiled [bar, 0.5 mm: top left,
mandibles; 1 mm: bottom middle, abdomens and uncoiled gut].

REVIEW OF AMICOTERMES 175

Te eee em

| Figs 39-46. Amicotermes congoensis. 39-40, crop with ‘bursters’ and proventriculus; 41-43, enteric valve seating and part
of P3; 4446, enteric valve armature opened out to show detail and variation, ridge positions numbered [bar, 1 mm: top
left, crop and valve seating; 0.5 mm: bottom right valve armature].

176 W.A. SANDS

49

Figs 47-55. Amicotermes cristatus. 47-49, mandibles with surface view of right molar plate; 50, 52, ventral and 51, 53,
dorsal views of abdomen to show gut coiling in situ; 54, gut dissected and uncoiled; 55, crop with “bursters’ and
proventriculus [bar, 0.5 mm: top left, mandibles; 1 mm: bottom right, abdomens and uncoiled gut; bottom left, crop and
proventriculus].

177

REVIEW OF AMICOTERMES

58

dj
i
Wily

i)

Figs 56-61. Amicotermes cristatus. 56-58, enteric valve seating and part of P3; 59-61, enteric valve armature opened out

to show detail and variation, ridge positions numbered [bar, 1 mm: top left, valve seating; 0.5 mm: bottom left valve

armature].

178 W.A.SANDS

Figs 62-69. Amicotermes dibogi. 62-64, mandibles with surface view of right molar plate; 66, 67, ventral and 66, 68,
dorsal views of abdomen to show gut coiling in situ; 69, gut dissected and uncoiled [bar, 0.5 mm: top left, mandibles; 1
mm: bottom middle, abdomens and uncoiled gut].

REVIEW OF AMICOTERMES 179

71

70

73

~_—""

a)

Figs 70-75. Amicotermes dibogi. 70, 71, crop with ‘bursters’ and proventriculus; 72-74, enteric valve seating and part of
P3; 75, enteric valve armature opened out to show detail, ridge positions numbered [bar, 1 mm: top right, crop and valve
seating; 0.5 mm: bottom left valve armature].

180 W.A.SANDS

78

76 77

79

Figs 76-82. Amicotermes galenus. 76-78, mandibles with surface view of right molar plate; 79-81, enteric valve seating
and part of P3; 82, enteric valve armature opened out to show detail, ridge positions numbered [bar, 0.5 mm: top left,
mandibles and enteric valve armature; 1 mm: bottom right, valve seating].

REVIEW OF AMICOTERMES 181

Figs 83-98. Amicotermes gasteruptus. 83-85, mandibles with surface view of right molar plate; 86, 88, 90, ventral and 87,
89, 91, dorsal views of abdomen to show gut coiling in situ; 92, gut dissected and uncoiled; 93, crop with ‘bursters’ and
proventriculus; 94-96, enteric valve seating and part of P3; 97, 98, enteric valve armature opened out to show detail and
variation, ridge positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: middle left, valve
seating and crop; middle right, abdomens and uncoiled gut].

182 W.A. SANDS

Figs 99-110. Amicotermes ivorensis. 99-101, mandibles with surface view of right molar plate; 102, ventral and 103,
dorsal views of abdomen to show gut coiling in situ; 104, gut dissected and uncoiled; 105, 106, crop with “bursters’ and
proventriculus; 107-109, enteric valve seating and part of P3; 110, enteric valve armature opened out to show detail, ridge
positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: middle right, abdomens and
uncoiled gut; top right, valve seating and crop].

REVIEW OF AMICOTERMES 183

Figs 111-124. Amicotermes mayombei. 111-113, mandibles with surface view of right molar plate; 114, 116 ventral and
115, 117, dorsal views of abdomen to show gut coiling in situ; 118, gut dissected and uncoiled; 119, crop with “bursters’
and proventriculus; 120—122, enteric valve seating and part of P3; 123, 124, enteric valve armature opened out to show
detail and variation, ridge positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: middle

right, abdomens and uncoiled gut; middle left, valve seating and crop].

184 W.A. SANDS

127

Figs 125-132. Amicotermes mbalmayoensis. 125-127, mandibles with surface view of right molar plate; 128, 130, ventral
and 129, 131, dorsal views of abdomen to show gut coiling in situ; 132, gut dissected and uncoiled [bar, 0.5 mm: top left,
mandibles; 1 mm: bottom middle, abdomens and uncoiled gut].

REVIEW OF AMICOTERMES 185

134

135

136

Figs 133-137. Amicotermes mbalmayoensis. 133, crop with ‘bursters’ and proventriculus; 134-136, enteric valve seating
and part of P3; 137, enteric valve armature opened out to show detail, ridge positions numbered [bar, 0.5 mm: bottom left,
valve armature; 1 mm: bottom right, crop and valve seating;].

186 W.A. SANDS

Figs 138-150. Amicotermes multispinus. 138-140, mandibles with surface view of right molar plate; 141, 143 ventral and
142, 144, dorsal views of abdomen to show gut coiling in situ; 147, gut dissected and uncoiled; 146, crop with “bursters’
and proventriculus; 147-149, enteric valve seating and part of P3; 150, enteric valve armature opened out to show detail,
ridge positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: bottom right, abdomens and
uncoiled gut; middle left, valve seating and crop].

REVIEW OF AMICOTERMES 187

153

152

Figs 151-161. Amicotermes spiculatus. 151-153, mandibles with surface view of right molar plate; 154, ventral and 155,

. dorsal views of abdomen to show gut coiling in situ; 156, gut dissected and uncoiled; 157, crop with ‘bursters’ and
proventriculus; 158-160, enteric valve seating and part of P3; 161, enteric valve armature opened out to show detail, ridge
positions numbered [bar, 0.5 mm: top left, mandibles and enteric valve armature; 1 mm: bottom right, abdomens and
uncoiled gut; middle left, valve seating and crop].

188 W.A. SANDS

STRICT CONSENSUS TREE OF AMICOTERMES Length 217 Cl 47 RI 44
from mhennig*;bb*; two trees

i+ SSeS
nea
35 aS teate+s6 galenus
g3 nS coocc oe a c
c ++-20 cs shee +8 ivorensis
aS 17 aE
23 ey eae p
ag + 10 mbalmayoensis
19 eae
Aon
yee
Sey
cro

bcs ers
8P BLP.
e9E BEZ

$+$-11 multispinus
18 3 congoensis
= $+4-9 mayombei
14 = =7,
13

meas gasteruptus
a #1 autothysius

162

a
a

09 6S 99S 4SS eps ees LS EBr Ely IOP PEP OF SE ELE BLE
ROE eG PBZ ALZ 99Z SZ WbZ CZ BLZ EBL PL EL BS &9 S&P

e9p LZ
92 SZ HZ

1

Figs 162, 163. Amicotermes. 162, Strict consensus tree from Hennig most parsimonious two trees, showing states of
characters with retention indices >50 at each node and for individual taxa (states shown alphanumeric, 1, la, 1b. . . ete);
163, Principal coordinates plot of eigenvectors on coordinates 1, 2 & 3, with minimum spanning tree superimposed as a
broken line.

REVIEW OF AMICOTERMES 189

SINGLE-LINKAGE TREE OF AMICOTERMES

0.45 0.50 0.55 0.60 0.65

Acholote
dibogi
gasterup
autothys
mayombel
congoens
multispi
galenus
mbalmayo
Sspiculat
cristatu

164 lvorensl

camer une

FLEXIBLE-LINKAGE TREE OF AMICOTERMES
0.16 ae 2, 0.48 0.64 0.80

| ae II IEE EIT
Acholote

dibogi
1vorensi
gasterup
autothys
mayombel
mbal mayo
congoens
multispi
galenus
camerune
Spiculat

165 cristatu

COMPLETE-LINKAGE TREE OF AMICOTERMES

0.15 0.30 0.45 0.60 0.75

Acholote
dibogi
gasterup
autothys
mayombel
mbal mayo
congoens
multispl
galenus
camer une
lvorensi

166 spiculat
cristatu

Figs 164-166. Amicotermes, phenetic trees. 164, Single Linkage tree; 165, Flexible Linkage tree; 166, Complete Linkage
tree.

190

W.A. SANDS

UNWEIGHTED PAIR-GROUP TREE OF AMICOTERMES

Ohe:

167

0.4

0.5

0.6

OZ

Acholote
dibogi
gasterup
autathys
mayombel
congoens
multispi
galenus
mbal mayo
camer une
spiculat
cristatu
1vorensl

NEIGHBOUR-JOINING TREE OF AMICOTERMES

10

168

Acholote
gasterup
autathys
mayombel
congoens
multispl

lvorensi
galenus

mbal mayo
camer une
spiculat
cristatu
dibogi

Figs 167, 168. Amicotermes, phenetic trees. 167, Unweighted Pair-Group Method of Averages tree; 168, Saitou and Nei

Neighbour-joining tree.

REVIEW OF AMICOTERMES

Figs 169-174. Amicotermes, enteric valve armature, valves slit and opened out; 169, 170, A. autothysius; 171, 172, A.
congoensis; 173, 174, A. cristatus.

192 W.A. SANDS

Figs 175-180. Amicotermes, enteric valve armature, valves slit and opened out; 175, A. camerunensis; 176, A.dibogi; 177,
178, A. galenus; 179, A. gasteruptus; 180, A. ivorensis.

REVIEW OF AMICOTERMES 193

Figs 181-186. Amicotermes, enteric valve armature, valves slit and opened out; 181, 182, A. mayombei; 183, 184, A.
mbalmayoensis; 185, A. multispinus; 186, A. spiculatus.

Bulletin of The Natural History Museum
Entomology Series

Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front
cover). Where the complete backlist is not shown, this may also be obtained from the same address.

Volume 46

No. 1 The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolothripinae). L.A.
Mound & J.M. Palmer. 1983. Pp. 1-174, 413 figs, 14 tables. £24.50

No. 2 A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae). B.C. Townsend. 1982. Pp. 175—
203, 63 figs. £4.20

No. 3 Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new taxa
(Chrysomelidae). S.L. Shute. 1983. Pp. 205—266, 150 figs, 10 maps. £9.00

No. 4 The Afrotropical dacetine ants (Formicidae). B. Bolton. 1983. Pp. 267-416, 81 figs. £21.00

Volume 47

No. 1 A new genus of oriental lacewings (Neuroptera: Chrysopidae). S.J. Brooks. 1983. Pp. 1-26, 95 figs. £4.00

No. 2 The leafhopper genus Batracomorphus (Cicadellidae: Iassinae) in the eastern Oriental and Australian
regions. W.J. Knight. 1983. Pp. 27—210, 977 figs. £25.75

No. 3 The Afrotropical idiocerine leafhoppers (Homoptera: Cicadellidae). M.D. Webb. 1983. Pp. 211-257, 146
figs, 2 tables. £7.00

Volume 48

No. | Gelechiid moths of the genus Mirificarma. L.M. Pitkin. 1984. Pp. 1-70, 112 figs, 2 tables. £10.00

No. 2 Macronematine caddisflies of the genus Amphipsyche (Trichoptera: Hydropsychidae). P.C. Barnard. 1984.
Pp. 71-130, 182 figs. £9.00

No. 3 A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera: Chalcidoidea). J.S. Noyes & M. Hayat.
1984. Pp. 131-395, 450 figs. 22512) 7/5)

Volume 49

No. | Afrotropical jumping plant lice of the family Triozidae (Homoptera: Psylloidea—. D. Hollis. 1984. Pp. 1—
102, 324 figs. £15.30

No. 2 The taxonomy of the western European grasshoppers of the genus Euchorthippus, with special reference

to their songs (Orthoptera: Acrididae). D.R. Ragge & W.J. Reynolds. 1984. Pp. 103-151, 88 figs. | £7.20
No. 3 An historical review of the higher classification of the Noctuidae (Lepidoptera). I.J. Kitching. 1984. Pp.

153-234, 4 figs. £12.00
No. 4 The Pimplinae, Xoridinae, Acaenitinae and Lycorininae (Hymenoptera: Ichneumonidae) of Australia. I.D.
Gauld. 1984. Pp. 235-339, 100 figs, 18 maps. SIGE 7/5)
No. 5 The Palaearctic species of Ascogaster (Hymenoptera: Braconidae). T. Huddleston. 1984. Pp. 341-392, 79
figs. £7.80
Volume 50
No. 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). P.S. Broomfield. 1985. Pp. 1—
152, 501 figs. £22.80
No. 2 Nymphal taxonomy and systematics of the Psylloidea (Homoptera). ILM. White & I.D. Hodkinson. 1985.
Pp. 153-301, 201 figs, 18 tables. £23.00

No. 3 The Whitefly of New Guinea (Homoptera: Aleyrodidae). J.H. Martin. 1985. Pp. 303-351, 48 figs. £8.00

Volume 51

No. 1 The ichneumon-fly genus Banchus (Hymenoptera) in the Old World. M.G. Fitton. 1985. Pp. 1-60, 129
figs. £10.80

No. 2 The phylogeny, classification and evolution of parasitic wasps of the subfamily Ophioninae
(Ichneumonidae). I.D. Gauld. 1985. Pp. 61-185, 52 figs. £21.00

No. 3 A cladistic analysis and classification of trichodectid mammal lice (Phthiraptera: Ischnocera). C.H.C.
Lyal. 1985. Pp. 187-346, 250 figs. £26.00

No. 4 The British and some other European Eriococcidae (Homoptera: Coccoidea). D.J. Williams. 1985. Pp.
347-393, 18 figs. £8.00

Volume 52

No. 1 The sandflies of Egypt (Diptera: Phlebotominae). R.P. Lane. 1986. Pp. 1-35, 80 figs, 4 tables. £5.60

No. 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). G.S. Robinson. 1986. Pp. 37-181, 200

figs. £24.00
No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986. Pp. 183-242, 68
figs. £11.00
No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera. J.
Quinlan. 1986. Pp. 243-366, 359 figs. £21.00
Volume 53
No. 1 A review of Miletini (Lepidoptera: Lycaenidae). J.N. Eliot. 1986. Pp. 1-105, 108 figs. £18.00
No. 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini. I.D. Gauld & G.A. Holloway. 1986.
Pp. 107-149, 65 figs. £8.40
No. 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on the
distribution of the Pseudophloeinae. W.R. Dolling. 1986. Pp. 151-212, 121 figs. £11.50
No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy
(Orthoptera: Acrididae). D.R. Ragge. 1986. Pp. 213-249, 128 figs. £7.50
No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble. 1986. Pp.
251-286, 102 figs. £7.00
Volume 54
No. 1 Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae). S. Okajima. 1987. Pp. 1—
74, 207 figs. £14.00
No. 2 Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae (Lepidop-
tera: Noctuidae). I.J. Kitching. 1987. Pp. 75-261, 465 figs. £36.00
No. 3 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr (Hymenoptera:
Formicidae). B. Bolton. 1987. Pp. 263-452, 100 figs. £36.50
Volume 55
No. | A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain
genera. M.W.R. de V. Graham. 1987. Pp. 1-392, 744 figs. £75.00
No. 2 The songs of the western European grasshoppers of the genus Stenobothrus in relation to their taxonomy
(Orthoptera: Acrididae). D.R. Ragge. 1987. Pp. 393-424, 116 figs. £6.00
Volume 56
No. 1 The legume-feeding psyllids (Homoptera) of the wet Palaearctic Region. I.D. Hodkinson & D. Hollis.
1987. Pp. 1-86, 294 figs. £16.00
No. 2 A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera). D. Hollis. 1987. Pp. 87-127,
94 figs. £6.00
No. 3 A review of the Rhadalinae (=Aplocneminae) (Coleoptera: Melyridae). E.R. Peacock. 1987. Pp. 129-170,
59 figs. £8.00
No. 4 A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). J. Quinlan. 1988. Pp. 171-229, 199
figs. £11.00
Volume 57
No. 1 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special
reference to the fauna of Costa Rica. I.D. Gauld. 1988. Pp. 1-309, 352 figs, 32 maps. £52.00
No. 2 A taxonomic revision of Alabagrus (Hymenoptera: Braconidae). M.J. Sharkey. 1988. Pp. 311-437, 28
figs, 22 maps. £24.50
No. 3 A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae). P. Huemer. 1988. Pp. 439-571, 221 figs.
£25.00
Volume 58

No. 1 The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78, 40 figs.
No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. Arzube &
C.A. Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour).

Volume 59

No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their taxonomy
(Orthoptera: Tettigoniidae). D.R. Ragge. 1990. Pp. 1-35.
A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 1990.
Pp. 37-115.

No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & P.C.
Barnard. 1990. Pp. 117—286.

Volume 60
No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). P.H. Williams. 1991.
Pp. 1-204.

No. 2 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. Pitkin & K.
Sattler. 1991. Pp. 205-241.
A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-288.

Volume 61
No. | Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76.
No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx Warren

(Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148.

Volume 62

No. 1 Caloptilia \eaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. Robinson.
1993. Pp. 1-37.

No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular
reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M. Pitkin. 1993.
Pp. 39-159.

Volume 63

No. | A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the

immature stages of economically important insect species (mainly Hemiptera and Lepidoptera). D.W.
Huang and J.S. Noyes. Pp. 1-135.

No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: Chrysopidae). S.J.
Brooks. Pp. 137-210.

Volume 64

No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M.A. Cook and M.J.
Scoble. Pp. 1-115.

No. 2 Encyrtidae of Costa Rica (Hymenhoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of
mealybugs (Homoptera: Pseudococcidae). J.S. Noyes and H. Ren. Pp. 117-164.

Volume 65

No. | A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae).
Y. Zhang and M.D. Webb. Pp 1-103.

No. 2 Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping
plant-lice (Homoptera: Psylloidea). J.S. Noyes and P. Hanson. Pp. 105-164.

Volume 66

No. 1 Biosystematic studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus. A.J. Shelley,
C.A. Lowry, M. Maia-Herzog, A.P.A. Luna Dias and M.A.P. Moraes. Pp. 121.

No. 2 Microtermes in East Africa (Isoptera: Termitidae: Macrotermitinae) S. Bacchus. 1997. Pp. 123-171.

Volume 67

No. 1 Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus Silvestri associated with the ant genus
Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococcidae).
D.J. Williams. 1998. Pp. 1-64.
Monophyly of the dacetonine tribe-group and its component tribes (Hymenoptera: Formicidae). B. Bolton.
1998. Pp. 65-78.
An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae,
Bombini). P.H. Williams. 1998. Pp. 79-152.

No. 2 A revision of the leafhopper tribe Paraboloponini (Hemiptera: Cicadellidae: Selenocephalinae) in the
Indian subcontinent. C.A. Viraktamath. 1998. Pp. 153-208.
A review of the Palaearctic Neorhacodinae (Hymenoptera, Ichneumonidae) with Eremura Kasparyan,
1995 new to the west Palaearctic. D.G. Notton and M.R. Shaw. 1998. Pp. 209-218.

Volume 68
No. 1 Revision of the Oriental Opostegidae (Lepidoptera) with general comments on phylogeny within the
family. Rimantas Puplesis and Gaden S. Robinson. 1999. Pp. 1-92.

A revision of the African and Malagasy species of the genus
Leptomastix (Hymenoptera, Encyrtidae), parasitoids of mealybugs
(Homoptera: Pseudococcidae)
Jean-Marc Anga and John S. Noyes oe
129 A revision of the north-west European species of the formosus species 2
group of Spilomicrus (Hymenoptera: Diapriidae)
David G. Notton
145 A review of the soldierless African termite genus Amiootennes Sands
1972 (Isoptera, Termitidae, Apicotermitinae)
W.A. Sands

ENTOMOLOGY SERIES

Vol. 68, No. 2, November 1999