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Full text of "Bulletin of the Natural History Museum Entomology"

ISSN 0968-0454 



Bulletin of 
The Natural History Museum 



uiv 



3 ;;ov 198s 






Entomology Series 





VOLUME 68 NUMBER 2 25 NOVEMBER 1999 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British 
Museum (Natural History) ), instituted in 1949, is issued in four scientific series, 
Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Entomology Series is produced under the editorship of the 

Keeper of Entomology: R.I. Vane-Wright 

Editor of Bulletin: Gaden S. Robinson 



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



3 NOV 1999 



Bull. not. Hist. Mus. Lond. (Ent.) 68(2): 93-128 Issued 25 November 1999 

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

JEAN-MARC ANGA 

International Cocoa Organisation, 22 Berners Street, London W1P 3DB, England. 

JOHN S. NOYES 

The Natural History Museum, London SW7 5BD, England. 

CONTENTS 

Synopsis 93 

Introduction 93 

Depositories 94 

Acknowledgements 94 

Terms and measurements 94 

Genus Leptomastix 95 

Diagnosis 95 

Identification 96 

Distribution of species 96 

Biology 96 

Use in biocontrol 96 

Key to African species of Leptomastix 96 

Revision of species 98 

Hosts of African Leptomastix 1 1 1 

References 112 

Index to scientific names 1 16 

Illustrations 117 



SYNOPSIS. The 1 1 species of Leptomastix known from Africa and Madagascar are 
revisedlllustrationsl 17. Three are described as new (herreni, africana, jonesi), three synonymies 
are proposed (lyciae with nigrocincta, phenacocci with nigrocoxalis, superba with dactylopii), 
and a lectotype is designated for algiricu 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. 



Xx/3aofea.7-l 



INTRODI ICTTON about the effect of the indiscriminate use of pesticides 

on the environment. This is especially so in Africa 

In recent years, increasing consideration has been where the continued use of pesticides may be unrealis- 

given to alternative methods to the use of chemicals tic for reasons other than environmental risk alone, eg. 

for pest control resulting from growing public concern poor financial resources or the remoteness of some of 

© The Natural History Museum, 1999 



94 



J-M. ANGA AND J.S. NOYES 



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. 



DEPOSITORIES 



BMNH The Natural History Museum, London, United 

Kingdom 
IEEM Instituto di Entomologfa Espanol, Madrid, Spain 
IITA International Institute of Tropical Agriculture, 

Cotonou, Benin 
INMP Universita Degli Studi Di Napoli Federico II, 

Italy 
MNHN Museum 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.CUSA 
ZAMU Zoological Museum of the Aligarh Muslim 

University, India. 



Acknowledgements We are grateful to the following 
for the loan of material: E. Guerrieri (INMP), I. Macek 
(NMP), J-L. Nieves Aldrey (IEEM), G.L. Prinsloo (PPRI), 
M.E. Schauff (USNM), S. Triapitsyn (UCR) andC. 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 dTvoire 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 



95 



FV 



FWL 



FWW 



GL 

Gono- 

stylus 

HW 

Hypo- 

pygium 

Linea 

calva 

Marginal 



MS 



MT 

MV 
OCL 

OL 
00L 

Post- 
marginal 



PMV 
POL 

Scale-like 
structures 



SL 



and clava. There are no anelli present in 

Leptomastix. 

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. 

Length of forewing excluding the marginal 

fringe and measured from the extreme base of 

the costal cell to the apex of the wing. 

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. 

Relative length of gonostylus. 

Third valvula, or ovipositor sheath, as seen in 

slide-mounted material (Fig. 7). 

Head width measured in facial view. 

Apical, externally visible sternite of abdomen. 

An oblique hairless line extending from the 
stigmal vein to posterior margin of the forewing. 
Measured from the distal margin of the 
subhyaline break at the apex of the submargina! 
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. 

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. 
Length of middle tibia for comparison with 
length of the ovipositor or last tergite. 
Relative length of marginal vein. 
The minimum distance between the posterior 
ocelli and occipital margin. 
Relative length of ovipositor. 
Minimum distance between the eye margin and 
the nearest posterior ocellus. 
Measured from the distal side of the stigmal vein 
where it joins the marginal vein to the apex of the 
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. 

Relative length of postmarginal vein. 
Minimum distance between the posterior ocelli. 
Structures on apical segments of antennae of 
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 ah, 1996). 
Relative length of the scape. This excludes the 



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 2.5^1x 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; F6 without scale- 
like structures ventrally (but see digitariae, Fig. 65); 



96 



J-M. ANGA AND J.S. NOYES 



clava solid and with a line scale-like structures ven- 
t rally in basal half (see Figs 6 1-68); 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 



segments varies from 1 1 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 
appressed hairs 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 
long as broad 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 

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 brown 5 



REVISION OF LEPTOMASTIX 

Table 1. A summary of the worldwide use of Leptomastix species in biocontrol projects. 



97 



Parasitoid Target pest Country/Region 


Year 


Result 


Reference 


dactylopii Planococcus citri Australia (Qd) 


1980 


SCI 


Smith etal. (1988) 


Belgium 


1989 


SCG 


Ronse(1990) 


Bermuda 


1952 


NC 


Cock (1985) 


USA (California) 


1934 


P 


Compere (1939b), Bartlett (1978) 


Canada 


1939-1945 


SCG 


Baird (1938, 1939, 1940a, 1941, 
1942,1943,1944,1946,1947), 
McLeod(1962), 
Tumbull& Chant (1961) 


Chile 


1930-1958 


SC 


Duran (1944), Gonzalez & Rojas 
(1966), Graf Marin & Pefia (1940) 


Costa Rica 


1968 


E 


Molina (1977), Noyes & Hayat (1994) 


Cyprus 


1966-1977 


SC 


Cock (1985), Greathead (1976), 
Krambias & Kotzionis (1980) 


UK 


1982-1987 


SCG 


Copland (1983), Copland & Varley 
(1987), Tingle & Copland 
(1988a,b,1989) 


Georgia 


1961 


E 


Kobakhidze(1965) 


Greece 


9 


P 


Jourdheuil (1986) 


India 


1983-1985 


SC 


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


Israel 


1984,1987 


SCG 


Argov & Rdssler ( 1 988), Rubin ( 1 985) 


Israel 


1941 


NE 


Rivnay (1960) 


Italy 


1956-1980 


SCI 


Greathead (1976), Luppino (1979), 

Viggiani (I975a,c), Barbagallo et 

al. 

(1983), Longo & Benfatto (1982) 








Morocco 


7 


P 


Jourdheuil (1986) 


Netherlands 


1984 


PG 


Hennekam et al. (1987): 
Kole&Hennekam(1990) 


Pakistan 


1985-1985 


P 


CIBC (1985,1986,1987) 


Portugal 


7 


P 


Jourdheuil (1986) 


South Africa 


1935 


NR 


Greathead (1971) 


Spain 


1948,1977 


NE 


Carrero (1980a,b), Greathead (1976), 
Gomez Clemente ( 1 95 1 ) 


Sweden 


1989-1990 


7 


Noyes & Hayat (1994) 


Turkmenia 


1968 


SC 


Niyazov (1969b) 


USA (California) 


1934 


P 


Bartlett (1978) 


USA (Florida) 


1934 


SC 


Watson & Thomson (1940a,b) 


USA (Texas) 


1970-1983 


SCG 


Meyerdirk et al. (1978), 
Summy etal. (1986) 


Uzbekistan 


1959-1960 


E 


Rozanova & Loseva (1963) 


Yugoslavia (former) 


7 


P 


Jourdheuil (1986) 


Planococcoides Ghana 


1949-1955 


NE 


Greathead (1971) 



njalensis 
Planococcus kenyae Kenya 
Pseudococcus USA (California) 

comstocki 
Pseudococcus Chile 

calceolariae 



abyssinica 
nigrocoxalis 



Planococcus citri 



USA (California) 



Maconellicococcus hirsutus 
Nipaecoccus viridis Jordan 



1937,1938 NR Greathead ( 1 97 1 ) 

1 973- 1 975 R *Meyerdirk & Newell ( 1 979) 

1931 SC Marco (1959), Rojas (1967) 



1931 NR Compere (1931) 

Egypt 1934 P Kamal(1951) 

1986 ? Meyerdirk et al. (1988) 



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

? - 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 



J-M. ANGA AND J.S. NOYES 



5 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 
(Fig. 18) 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) 

6 Antenna with Fl at least 4x as long as pedicel (Fig. 22) 

africana Anga & Noyes sp. n. (p. 103) 

- Antenna with Fl not more than 3x as long as pedicel 
(Figs 26, 32, 33, 39, 49, 50, 56) 7 

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

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

8 Distance of posterior ocelli from eye margin (OOL) less 
than 1.4x distance from occipital margin (OCL); ocellar 
area of frontovertex orange, axillae orange 

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 

nigra Compere (p. 103) 

9 Mid coxae completely yellow 



structures (Fig. 61); head and thorax usually largely 
orange herreni (p. 101) 

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

15 Hind coxae brown 16 

- Hind coxae yellow 17 

16 Fore coxae brown africana (p. 103) 

- Fore coxae yellow abyssinica (p. 110) 

17 Temples and occiput yellow tsukumiensis (p. 98) 

- Temples and occiput largely blackish 18 

1 8 Scale-like structures arranged in line at base of clava 
very slender and elongate and difficult to distinguish 
from setae (Fig. 64) jonesi (p. 108) 

- Scale-like structures arranged in line at base of clava 
flattened and short and relatively easy to distinguish 
from setae (Figs 63, 66, 67) 19 

19 Longest setae on Fl at least 4x as long as diameter of 
segment measured in middle (Fig. 63); mesostemum 
dark brown, almost black, mid coxae yellow; forewing 
with linea calva interrupted by not more than two setae 
(Fig. 45) 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 31, 38, 55) 20 

20 Forewing with linea calva interrupted by at most only 
three or four setae (Fig. 55); mandibles and mouth mar- 
gin never darkened nigrocincta (p. 109) 

- Forewing with linea calva interrupted by at least five 
setae (Figs31,38); bases of mandibles and mouth margin 
sometimes blackish 21 

21 Frontovertex and axillae almost completely orange 

nigrocoxalis (p. 103) 

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



dactylopii Howard (p. 106) 

Mid coxae at least partly brown 10 

10 Linea calva interrupted by at least 4 lines of setae; mid 

coxae generally orange and only brown laterally 

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 

11 

1 1 Hind and fore coxae concolorous, yellow 

nigrocincta Risbec (p. 109) 

- Hind coxae brown contrasting with yellow fore coxae 
abyssinica Compere (p. 110) 

MALES 

1 2 Forewings with postmarginal vein not longer than stig- 
mal vein; head mostly dark brown (Figs 14, 20, 21) 

13 

- Either forewings with postmarginal vein clearly longer 
than stigmal vein (Figs 3 1 , 38, 46, 55, 60) or head orange 
without any brown areas 15 

1 3 Tegulae white with brown apex; F6 with scale-like struc- 
tures (Fig. 65) digitariae (p.l 00) 

- Tegulae with base orange or completely dark brown; 
only clava with scale-like structures (Figs 61 , 62) (some- 
times apparently absent) 14 

14 Clava with 5-7 longish, slender, recurved scale-like DIAGNOSIS. Female (length 1.42-2.41): body 



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, 1971:50-51. 

Holotype 9 , India, ZAMU, not examined). 

Synonymized with tsukumiensis by Noyes & Hayat 

(1994). 
[Leptomastix longipennis Mercet; Otanes, 1935:503- 

504. Misidentification.l 



REVISION OF LEPTOMASTIX 



99 



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 Fl 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. Ox 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. 



Material examined. 

Non-type material. Gambia, c?2, Bakau, i. 1978 
(Huggert); 1 cf , Fajara, i. 1978 (Huggert); Ivory Coast, 
1 d", Lamto, 6°13'N 5°02'W, 16. v. 1985 (Rasplus); 8 9 , 
13d", Lamto, 6°13'N 5°02'W, xi.1988 (Noyes); 19, 
Bouake,xi.l981 (Cochereau); R.P. Benin, ld,25Km 
N Ohicon, 4.xii.l988 (Noyes); Nigeria, 119, 20 d\ 
Kaduna St., 20 Km N Kaduna, 8.xi.l987 (Noyes, 
Neuenschwander); Id", 81Km NW Jos, l.xi.1987 
(Neuenschwander); 6 9 , 6d\ Plateau St., 20 Km NW 
Jos, 12.xi.1987 (Noyes, Neuenschwander); 1 9, 6d\ 
Bauchi St., 74 Km W. Bauchi, 14.xi.1987 (Noyes, 
Neuenschwander); 1 9 , Sokoto St., 54 Km E. Sokoto, 
8.xi. 1 987 (Noyes, Neuenschwander); Cameroon, 1 9 , 
Sanyere, 18.xii.81 (Compton); Somalia. 19, Lr. 
Shabelli Valley, Mogadiscio, v. 1977 (Bin); Uganda, 
ld\Serere,22.vii. 1934, BM 1935^59 (Ford); Kenya, 
1 9, Nairobi, Karen, vi. 1982 (Dewhurst); Zambia, 
1 9, 15Km E Lusaka, ll-19.ii.1980 (Beaver); Zim- 
babwe, 14 9, 2o\ Harare, Chishawasha, i.81— xii.84 
(Watsham); South Africa, 39, Cape Province, 
Swellendam, 17.xii.31-18.L32 (Turner), 16 9, 9d\ 
Cape Province, Ceres, various dates iii.1921-iv.1925 
(Turner); 19. Cape Province, Matjesfontein, 7- 
13. xi. 1928 (Turner); 1 9, 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.H. 1987 
(Prinsloo); 1 9 , Natal, Vernon Crookes Nat Res., 
Umzinto, 30°17'S 30°37'E, 443m, 25-26.iii.1985 
(Prinsloo); 1 9 , C.P.. Andries Vosloo Kudu Res. nr. 
Grahamstown, 33°07'S 26°38'E, 30.xi.1983 (Prinsloo, 
Grobbelaar); 2 9 , Tvl, Blyderivierspoortdaam Nat. 
Res., 24°32'S 30°47'E, 25-26.X.1984 (Prinsloo); 1 9 . 
C.P., Grahamstown, 33°19'S 26°32'E, 13. i. 1983 
(Prinsloo, Grobbelaar); 1 9 , Tvl, Hans Merensky nat. 
Res., 23°40'S 30°39'E, 27-30.U981 (Prinsloo); 19, 
Tvl, Mogol Nat. Res., Ellisras District, 23°58'S 
27°45'E., 21. i. 1983 (Prinsloo); Id, Tvl, Pretoria, 
T3754, iii.1971 (Annecke); 1 9,Tvl,Letsitele,xi.l978 
(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. It is 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. lx as long as 
broad whilst in dactylopii the temple and mesoster- 
num are dark brown and the forewings are 2.6-3. Ox as 
long as broad. 



100 



J-M. ANGA AND J.S. NOYES 



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 Fl; 
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; Fl 1.6x as long as pedicel; clava 1.7xaslong 
as Fl and twice as long as F6. Relative measurements: 
HW 85, HL 75, FV 35, POL 15.5, OCL 8, OOL 6, EL 
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. 



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 2 1 .vii. 1988 by JSN and labelled 
'LT 9' on ventral side of card. Paralectotypes: 8 9, 
6d", 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, 2cf paralectotypes in 
BMNH. 

Non type material. Madagascar, 1 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 phy logenetic 
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 



101 



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; Fl 
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 



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; Fl 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,HH92,FV50,POL19.5,OCL5,OOL11,EL58, 
EW37, 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. 1 8): hindwing 4.2x as long 
as broad; gaster as long as thorax, with ovipositor 
hidden. Relative measurements: FWL 230, FWW 90; 
HWL153, HWW36. 

Gaster as long as thorax, with ovipositor (Fig. 19) 
hidden. Relative measurements (paratype): OL 35, 
MT51. 

MALE. See diagnosis. Relative measurements: HW 
45, FV 27, POL 10, OOL 7, OCL 2.5, 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 Fl 
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 Icoma-aurea, and 
lAthenasia sp. 

Distribution. South Africa. 

Material examined. 

Type-material: Holotype 9 , South Africa, C.R, 
Klipheuwel, ex Octococcus sp. on Compositae, T2783, 
x.1968 (Insley). Paratypes: South Africa, 99, 5o\ 



102 



J-M. ANGA AND J.S. NOYES 



C.P., x.1968, ex mealybug on Rehlania ericoides, 
T2779, x. 1968 (Insley); 29, 2cf, C.P., Klipheuwel, 
ex Octococcus sp. on Compositae, T2783, x.1968 
(Insley) 3 9, 2o", same data as holotype; 3 9, 2cf, 
C.P., Tulbagh district, ex mealybug on Rehlania 
ericoides, T2833, x.1968 (Gilliomee); 6 9 , 5d\ C.P., 
Caledon, x.1969, ex Octococcus sp.n. on Oedera 
imbricata, T3219, x.1969 (Insley); 3 9, 10", C.P, 
Villiersdorp District, ex mealybug on Chrysocoma 
Icoma-aurea, T3217, x.1969 (Insley); 1 9 , lcf, C.P, 
Paarl, x.1971, H.PInsley, ex mealybug on Cape 
Macchia, T3971, x.1971 (Insley); 19, 4d\ C.P, 
Worcester, ix.1972, ex mealybug on unknown plant, 
T4525, ix.1972 (Neser); 3 9 , C.P, Windmeul nr Paarl, 
ex Octococcus sp. nr. pentziae on Athenasia trifurcata, 
T4922, 3.X.1974 (Prinsloo) 2 9, ld\ C.P, Ceres, 
9.x. 1974, on Rehlania genistaefolia, T4927, 9.x. 1974 
(Prinsloo); 99, 6cT, C.P, Tulbagh, ex Wctococcus 
sp. on 1 Athenasia sp., T5469, vi, 1978 (Neser); 5 9 , 
9d", OFS, Edenville, OFS, ex Octococcus pentziae on 
Pentzia globosa, T3503, i.1970 (Insley); 29, 3d\ 
C.P, Oudtshoorn, ex Octococcus sp. on 'Mesem', 
T6915, iii.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. 6 1 ) 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 



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 cf (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 Fl ; 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, 1 9, Aures: Ain Zaatout, 1 . — 4.vi. 1 97 1 (Hoffer, 
Horak [Expedition 'SCARABAEUS']) 'Holotypus 
Leptomastix algirica Hoffer 9'. Paralectotypes: Al- 
geria, 19, Biskra, 7.-8.vi.l971 (Hoffer, Horak 
[Expedition 'SCARABAEUS']); Id", Auresi, 
Ainzaatout, 1 .— 4.vi. 1971 (Hoffer, Horak [Expedition 
'SCARABAEUS']. Lectotype in NMP, paralectotypes 
in BMNH. 

Non-type material. Greece, 1 9 , Kos, 4Km W Kos 
Town, 20.viii.1994 (Noyes); Cyprus, 1 9 , 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 



103 



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 Fl 
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 5. 

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. 



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 F 1 ; 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 from a 
mealybug, probably Octococcus sp., on Elytropappus 
rhinocerotis (see below). 

Distribution. South Africa. 

Material examined. 

Type material. Holotype 9 : South Africa, C.R, 
Stellenbosch, Idas Valley, ex lOctococcus sp. on 
Elytropappus rhinocerotis, T5976, ii.1979 (Neser). 
Paratypes: South Africa, 39, lcT, same data as 
holotype; 2cf, C.R, Paarl, ex lOctococcus on 
Elytropappus rhinocerotis, T2737, ex 14.x. 1968 
(Insley); 7 9, 3cf, Cape Province, Ceres, iii.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 9 , South Africa, USNM, not examined. 
Leptomastix phenacocci Compere, 1938:325. Holotype 

9 , 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 



J-M. ANGA AND J.S. NOYES 



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 



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 et al. ( 1 985 ) 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 9, 
5d\ same data as holotype (BMNH, UCR). Holotype 
9 of Leptomastix brevis, India, Kerala, Kalamasseri, 
ex Icerya aegyptica (Douglas), 12.iii.1967 (Hayat) 
(BMNH). Lectotype 9 of Leptomastix longiscapus: 
India, Uttar Pradesh, Garhwal, ex coccid on citrus, 
5.ix.l975 (Khan) (ZAMU); Paralectotypes of 
Leptomastix longiscapus: India, 9 9, 4cf, 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, 5 9 Giza, 
7.ix.l929, ? hibiscus mealybug breeding room, 
(Kamal); Senegal, 12 9, 6o", Louga, ex cochenille sur 
Euphorbia, 1945 (Risbec) (misidentified as 
Leptomastix flava by Risbec); 1 9 , Bambey, sur 
Sourour (misidentified as Leptomastix flava by Risbec); 
South Africa, 3 9 , 5d", Tvl, Dendron, ex, mealybug 
on grass, T6067, i. 1980 (Prinsloo); 1 9 , Tvl, Soutpan, 
Pretoria Dist., 25°24'S 28°06'E, 11-12. i. 1984 
(Grobbelaar); 119, 1 d\ Tvl, Levubu Tvl, ex mealybug 
on grass, T6924, i. 1987 (Prinsloo); 1 9 , Tvl, Entabeni 
For. Res., 33°00'S 30°16'E, 7-1. i. 1987 (Prinsloo); 
19, 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 



105 



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. 1 8). 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 9 , 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 



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 
Fl 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 Stoehe cinerea; 
Octococcus ?minor on Athenasia trifurcata; 
Octococcus pentziae on Metalasia muricata and 
Pentzia globosa; Octococcus on Elytropappus: 
lOctococcus on Stoebe and Elytropappus 
gnaphaloides; Paracoccus burnerae on IDiospyros 
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 9 of Leptomastix nigra. South 
Africa, Tvl, Johannesburg, May 1937 {Compere) 
(BMNH). Paratypes: South Africa, 19, 2d\ Tvl, 
Johannesburg, ex mealybug, i v. 1937 (Compere). 
Other material. Zimbabwe, 59, Salisbury, near Harare 
and Chishawasha, various dates iii.1979-vii.1982 



106 



J-M. ANGA AND J.S. NOYES 



(Watsham); Namibia, 1 9, Helmeringhausen, with 
Parasaissetia nigra on Loranthus elegans, T6925, 
v.1973 (Prinsloo); 29, 2d, Otavi, ex mealybug on 
unidentified plant, T6928, 17.v. 1973 (Prinsloo); 4 9 , 
Karibib, ex mealybug, T6931, 16.ii.1978 (Kok);69, 
Id", Otavi, ex mealybug, T6930, 24.ii.1978 (Kok); 
South Africa, 6 9 and 9d\ OFS, Edenville, OFS, ex 
Octococcus pentziae on Pentzia globosa, T2503, 
i. 1970 (Insley); 17 9, C.P., Jeffreys Bay, ex mealybug 
in axils of Gazania rigens, T6916, 3. i. 1980 (Neser); 
19d\ C.P., Jeffreys Bay, ex mealybug in axils of 
Gazania, T6917, 3.L1980 (Neser); 29, 7d\ Tvl, S. 
of Pongola, 27°23'S 30°38'E, ex Saissetia somereni 
on grapefruit, T6932, vii. 1979 (Kambourov); 1 9 , C.P., 
Witzenberg Vail., 3, 000ft, 19.i.l921 (Turner); 19, 
Id", C.P., Paarl District, ex ^Octococcus on Stoebe, 
T2736, x.1968 (Insley); 2 9 , 2d\ C.P, Du Toits Kloof 
Pass, ex ^Octococcus sp. on Elytropappus 
gnaphaloides, T2795, x.1968 (Insley); 4 9 , Id", Du 
Toits Kloof Pass, ex mealybug on Metalasia muricata, 
T2832, x.1968 (Insley); 2 9 , 2d, C.P, Villiersdoorp, 
ex. mealybug on Stoebe sp., T3205, x.1969 (Insley); 
89, 9d, C.P, Swellendam, T3226, ex Spilococcus 
sp. on Metalasia muricata, x.1969 (Insley); 3 9 , C.P, 
Stellenbosch, ex Octococcus Iminor on Athenasia 
trifurcata, T3231, x.1969 (Insley); 8 9, 15d, C.P, 
Jeffreys Bay C.P, ex Octococcus sp. on Metalasia 
muricata, T3378, iii.1970 (Insley); 3 9 , C.P, Garcia 
Pass, ex Octococcus sp., T3414, iii. 1970 (Insley); 1 9 , 
4d, C.P, Stellenbosch, on Stoebe plumosa, T5383, 
v.1978 (Neser); 49, 3d, C.P, Kleimond, ex 
Octococcus africanus on Stoebe cinerea, T5475, 
viii.1978, (Neser); 2 9 , 4d, C.P, Floraberg, ex prob. 
Octococcus sp. on Metalasia sp., T5793, vi. 1978 
(Neser); 79, Id, C.P, Franschhoek Pass, ex 
Octococcus pentziae on Metalasia muricata, T5883, 
xii.1978 (Neser); 2 9 , 15d, C.P, D.F Malan Airport 
C.P, ex Octococcus prb. pentziae on Metalasia 
muricata, T6114, iv.1979 (Neser); 3d, C.P, D.F. 
Malan Airport, with Ceroplastes sp. on Metalasia 
muricata, 10.iv.1979 (Neser); 4 9 , 1 d, C.P. Goudini, 
ex. prob Octococcus sp., on Stoebe sp., T6914 (Neser); 
5 9, Id, C.P, Goudini, ex prob. Octococcus sp. on 
Elytropappus rhinocerotis, T6920, 25. ix. 1979 (Neser); 
5d, CP, Algeria, Cadarberg, ex prob. Octococcus sp. 
on Elytropappus sp. cf. gnaphaloides, T6921 
16.vii. 1979 (Neser); 4 9 , 1 d, C.P, Sir Lowry's Pass 
rd. nr. Somerset, ex Octococcus sp. on Athenasia sp., 
T6922, 29.viii.1979 (Neser); 6d, Tvl, Irene, ex 
Paracoccus burnerae on IDiospyros lycioides, T6927, 
16.iii.1969 (Insley); 14 9 , 15d, C.P, Capetown, D.F 
Malan Airport, ex Octococcus prb. pentziae on 
Metalasia muricata, T6230, vii. 1980 (Neser); 29, 
Id, C.P, Gamtoos Sta., with Ceroplastes sp. on 
Metalasia muricata, T6933, 22.xi.1983 (Grobbelaar). 
Material in BMNH, PPRI. 



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 herreni) 

Leptomastix dactylopii Howard 
(Figs 39-^16, 63, 70) 

Leptomastix dactylopii Howard, 1885:23-24. 

Lectotype 9 (designated Noyes & Hayat, 1994), 

USA, USNM examined. 
Leptomastix superba Silvestri, 1915:348. Holotype 9 , 

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 9 , Tanzania, examined. Synonymized by 

Noyes & Hayat (1994). 



REVISION OF LEPTOMASTIX 



107 



Paraleptomastixdactylopii (Howard); Mani, 1939:70- 

71. 
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 Fl ; 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 saecharifolii 
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, 1 994), 
Pseudococcus occiduus on Psidium guajava by 
Annecke & Insley (1971), Planococcoides njalensis 
by Strickland (1951), Planococcoides lamabokensis 



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 
lysis lemon. 

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 '1170o Pars on Dactylopius de- 
im/<:7wOct4.[18]84"Type2649U.S.N.M.(USNM)'. 
Paralectotypes lcf, on single point, labelled '1170o, 
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 Nefasif (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' 
(HEM). Leptomastix bifasciata: holotype 9, South 
Africa, Uitenhage, ex Pseudococcus longispinus on 
oleander, 12.U937 {Compere) (BMNH). Paratypes: 
Tanzania (as Tanaganyika), 3 9 . Bukoba, ex 
Pseudococcus on guava, 20.vii. 1935 (Ritchie) 
(BMNH). Leptomastix tambourissae: lectotype 9 , 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: 3 9 , 1 cf (not four as 
stated by Risbec, 1952) remounted from same slide 



108 



J-M. ANGA AND J.S. NOYES 



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); 2 9 , Bambey, ex Pseudococcus on 
Prosopis, 12.x. 1945 (Risbec) (misidentified as L. flava 
by Risbec); Gambia, 1 cf , Bakan, i.78 (Huggert); 2cf , 
Fajara, i.78 (Huggert); Sierra Leone, lcf, Njala, ex 
mealybug on Cassiea javanica; Ivory Coast, 7 9 , 
1 cf , Lamto, 6°13'N 5°02'W, MT/YPT, xi.1988 (Noyes); 
2d\ Korhogo, 1 8-2 l.iii. 1984 (M. Matthews); 19, 
3cf, Gagnoa, 2-5.iii.1984 (Matthews); 19, 3d", 
Bouaffle; R.P. Benin, 5 9, 7cf, 15Km N. Cotonou, 
Abomey-Calavi, MT, xii.1988 (Noyes); Togo: 2 9, 
Kpalime, xii.1988 (Noyes); Nigeria, 3 9 , 1 cf , Kaduna 
St. 20km N., 8.xi.l987, (Noyes, Neuenschwander); 
5 9 , 5cf , Ibadan, IITA compound, MT/YPT, xi.1987 
(J.S.Noyes); 6cf , Zaria, 8.xi.l987 (J.S. Noyes); Cam- 
eroun, 29, 2cf, S.W. Cameroun, Musone, Lake 
Mboandong, ll.xii.1981 (Compton);2Q , 3cf, Victo- 
ria Bot. Gdns,6.xii. 1981, (Compton); 1 9,Nyassosso, 
Mt Koupe, 8.i. 1982 (Compton); Gabon, 29, 2d\ 
Foret de la Mondah, 15-25km N.Libreville, 25. xi- 
3.xii.87 (Noyes); Uganda, 1 9, Bujumbura, iv. 1939 
(Taylor); 3 9, 4cf , ex Pseudococcus kenyae on C. 
arabica, 12.iv.1938 (Melville); Zimbabwe. 69, 3 cf, 
Salisbury, i.1979 (Watsham); 69, 2cf, Harare, 
Chishawasha, xii. 1982 (Watsham); Zambia, 9 9 , 13cf , 
15km E.Lusaka, 27.xi-29.xii. 1979 and 1 1-19.H. 1980 
(Beaver); Kenya, 19, Nairobi, ii.1982, Brit. Mus. 
1982-347; 3cf, Gazi, viii.1982 (Barnett); South Af- 
rica, 18 9, 29 cf, Cape Province, 10-22.xii. 1930, Brit. 
Mus. (Turner); 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, i. 1979 (Prinsloo); 
1 9 , 1 cf Letaba, ex suction trap on citrus, T2504, 
xi. 1966 (Catling); 4 9 , 1 cf Cape Province, Ashton, ex 
mealybug on Ficus carica, T5306, iii.1978 (Neser, 
Kok & Urban); 19, Okahandja, 2-18.iii.1928; 19, 
Transvaal, Roodeplaat, nr Pretoria, ex Allococcus 
quaesitus on Acacia sp., T4292, iii. 1992, (Insley); 1 cf , 
Mogol Nature Reserve, Ellisras Dist. 23°58'S 27°45'E, 
27-29.ii.1984 (Prinsloo); lcf, Entabeni Forest Res. 
Soutlanberg, 25°00'S 30°16'E, 3-7.xi. 1980 (Prinsloo); 
1 cf , C.P., Jeffreys Bay, with unidentified coccoidea on 
Euclea sp., T6918, 13L1980 (Neser); lcf, Natal, 
Umzinto, ex Planococcus sp. on Erythrina lysistemon, 
T4135 i.1972 (Insley); 29, C.P., De Dooms, ex 
Planococcus ficus on Ficus carica, T6919; Mauri- 
tius, 2cf, Rose Hill, 29.ix.32 (Mamet). Material in 
BMNH, PPRI, MNHN. 

Voucher material. Israel, 1 9 , 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; 1 9 , 
lcf, Negev, lab reared, California, Porterville, ex 
comstock mealybug, Warkentin, R76-42, 17.V.1976, 



shipped and released in California, Porterville, by 
Meyerdirk as Leptomastix flavus; 1 cf , 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; 
fore wing 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 Fl ; 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 



109 



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 F 1 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, GL 5.5 [MT 
84]. 

MALE. See diagnosis. 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 1 1 .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: Id", same data as holotype; 19, Cape 
Province, Mossel Bay, 18-30.xi.1921 (Turner); lcf, 
Cape Province, Mossel Bay, vi— vii. 1930 (Turner); 
3 9, Cape Province, Somerset East, 1-26.L1931 
(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 lOx 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 



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, 1 959:27. Lectotype 9 , 
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 Fl ; 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 



J-M. ANGA AND J.S. NOYES 



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 
( 1 959). Recorded in India from Coccidohystrix insolita 
and Phenacoccus sp. (probably misidentification of C. 
insolita) on aubergine, Solarium melongena L. (Noyes 
& Hayat, 1994). 

Distribution. Togo, Zambia, Madagascar, India, 
China. 

Material examined. 

Type material. Lectotype 9 of nigrocincta: 'Elevage 
du 5.12.50 No 810 Eclose du 20.12.50 Parasites des 
cochenilles surle feuilled' Hibiscus sp. deTsimbazaza 
(R.R) Inst. Scient. Madagascar' 'Leptomastix 
nigrocincta', remounted on a card rectangle and la- 
belled 'LT 9 ' on ventral side of card (MNHN). 
Paralectotypes: 6 9, 39, 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, IARI area, x.1979 
(Boucek). Paratypes: 53 9 , 13 o" 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 9, 10km NW Kpalime, 
xii. 1988 (Noyes); Zambia, 1 9, Lusaka, 6-1 2.xi. 1979 
(Beaver); Madagascar, 1 d\ Tananarive, Angavokely 
for sta., 28km E. Angavokely, 24.iv. 1983 (Noyes) 2 9 , 
2d\ Perinet, 27.iv.-3.v. 1983 (Noyes, Day); 519, 
70cf, Tulear, Berenty 12km, NW Amboasary, 5- 
15.v. 1983 (Noyes, Day); 3 9 , 1 1 cf , 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 



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, Fl 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. 1 8). 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 9 , Eritrea, not examined, USNM. 

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



REVISION OF LEPTOMAST1X 



111 



with inner eye margins, genal area and area below 
toruli yellow; antennae dark brown with ventral mar- 
gin of scape yellow or translucent; presternum 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 Fl 
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 lOxaslongasgonostyli. 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, 119, 3o", Nefasit, 
ex Pseudococcus citri on Olea chrysophylla, iii.1930 
and iv. 1930 (Compere); 1 9 , Nefasit, ex Pseudococcus 
citri on Olea chrysophylla, i v. 1930 (Compere); 3 9, 
2d\ Nefasit, ex Pseudococcus No 4630 on Olea 
chrysophylla, 15. iv. 1930 (Compere); 1 d", Nefasit, ex 
Pseudococcus No 4630 on Olea chrysophylla, 
16. iv. 1930 (Compe re); 1 9, Nefasit, on window, out of 
mealybug, 3 l.iii. 1930 (Compere). Material in BMNH 
and UCR. 

Non type material. Eritrea, 5c? , Nefasit, ex 
Pseudococcus No 4630 on Olea chrysophylla, 
1 1 .iv. 1 930 (Compere); 4d", Nefasit, ex Pseudococcus 
No 4630 on Olea chrysophylla, 10.iv.1930 (Com- 
pere); 1 9 , Ezdaclesan, ex Croton macrostachys, 
l.v.1930 (Compere); 59, 1 o\ 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, presternum 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, presternum 
and fore coxae are yellow, the posterior ocelli are 
equidistant from the eye and occipital margins and the 
clava is longer than Fl (Fig. 56), whereas in africana 
the scape is entirely brown, the presternum and coxae 
are dark brown, the posterior ocelli are nearer the 



occipital margin and the clava is shorter than Fl (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) - 

1* 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) - 1* dactylopii 
Coccidohystrix — *nigrocoxaiis 
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 - 

!?* dactylopii 
Phenacoccus madeirensis Green - dactylopii 
Phenacoccus saccharifolii, see Birendracoccus 

saccharifolii 
Phenacoccus solani Ferris - \dactylopii 
Planococcoides njalensis (Laing) - dactylopii 



112 



J-M. ANGA AND J.S. NOYES 



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) - Idactylopii 
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 - l*nigrocoxalis 



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Biological Control. Fontana. Bollettino del Laboratorio di 
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. Ill 
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 

Coccidohystnx 104, 110, 111 

Coffea 108 

coma-aurea (Chrysosoma) 101, 102 

Combretum 107 

comstocki (Pseudococcus) 97. 107, 108. 

112 
concavocerarii (Pseudococcus) 107. 112 
Croton 1 1 1 
cryptus (Pseudococcus) 103 

dactylopii (Leptomastix) 96. 97, 98, 99, 

105, 106-108. 109, III, 112. 123-124, 
127, 128 

Dactylopius 107, 111. 112 

Delotococcus 107. 1 1 1 

destructor (Dacylopius) 107, III, 112 

Digitaria 100 

digitariae (Leptomastix) 96,98, 100-101. 

102, 118-119. 127, 128 
Diospyros 105, 106 
Dysmicoccus 107, 1 1 1 

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. IK). Ill, 119-120, 127 
Hibiscus 104, 110 

hirsutus (Maconellicoccus) 97, 104, 111 
hirta (Hyparrhenia) 104 
histrio (Leptomastix) 95, 105. IK) 
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, 

Coccidohystnx) 104. IK). Ill 
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 



J-M. ANGA AND J.S. NOYES 



manihoti (Phenacoccus) 94 

maritimus (Pseudococcus) 107 

medica (Citrus) 104 

Melia 104 

melongena (Solanum) 110 

Metalasia 105, 106 

minor (Octococcus) 105, 106, 111 

Moras 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 

phenacoeci (Leptomastix) 96, 103, 104 
Phenacoccus 94, 104, 107, 110, 111, 112 
Planococcoides 97, 107, 1 11, 1 12 
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- 



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. Ill, 117-118. 128 

virgata (Ferrisia) 99, 107, 1 1 1 
viridis (Nipaecoccus) 97, 104. Ill 
vitis (Planococcus) 108 
vovae (Planococcus) 107.112 

Wittania 104 

Ziziphus 104 



REVISION OF LEPTOMASTIX 



117 









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



118 



J-M. ANGA AND J.S. NOYES 




' 






8 







V 




Figs 7-12. Leptomastix tsukumiensis - 7, forewing, apex of venation, variant, 9 . Leptomastix digitariae - 8, antenna, 9 ; 
9, forewing, 9 ; 10, forewing base, 9 ; 11, forewing, apex of venation, 9 ; 12, ovipositor. 






REVISION OF LEPTOMAST1X 



119 











• 




JrS 






*2^ 




13 



/ w 1 
















r ' / 

' ' i ' 



14 



■V 




15 







,^ 




y^ 






16 
















/ / 



' / / / < 



18 



:- 



Figs 13-18. Leptomastix digitariae - 13, forewing, d" ; 14, forewing, apex of venation,d\ 15-18, Leptomastix herreni - 15, 
antenna, 9 ; 16, forewing, 9 ; 17, forewing base, 9 ; 18, forewing, apex of venation, 9 . 



120 



J-M. ANGA AND J.S. NOYES 







20 



22 



23 




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



REVISION OF LEPTOMASTIX 



121 







26 




Figs 25-30. Leptomastix africana - 25, forewing, apex of venation, 9 . Leptomastix nigrocoxalis - 26, antenna, 9 ; 27, 
forewing, 9 ; 28, forewing base, 9 ; 29, forewing, apex of venation, 9 ; 30, forewing, d". 



122 



J-M. ANGA AND J.S. NOYES 




i*^ 



33 







^ 


#& 


« / 






^-'/ ''...-■■' 


35 


r *" 




32 






. i 


-1 

; 


a**-.. 




• 

36 



Figs 31-36. Leptomastix nigrocoxalis - 31, forewing base,d". Leptomastix nigra - 32, antenna, large specimen, 9 ; 33, 
antenna, small specimen, 9 ; 34, forewing, 9 ; 35, forewing, apex of venation, 9 ; 36, ovipositor. 






REVISION OF LEPTOMASTIX 



123 





39 



40 




41 





j/"' 




Jff/Tf 

& 42 







Figs 37^12. Leptomastix nigra - 37, forewing, C?; 38, forewing base,d\ Leptomastix dactylopii - 39, antenna, 9 ; 40, 
forewing. variant, 9:41, forewing, variant, 9 ; 42, forewing base, 9 . 



124 



J-M. ANGA AND J.S. NO YES 




<//!& 



^ 



'7j^/ / > 



JW 



/// 



L 



43 




45 





47 




A& 



48 



Figs 43^18. Leptomastix dactylopii - 43, forewing, apex of venation, 9 ; 44, forewing, C? ; 45, forewing base, cf ; 46, 
forewing, apex of venation, cf. Leptomastix jonesi - 47 ', forewing,d"; 48, forewing, apex of venation, cf. 



REVISION OF LEPTOMASTIX 



125 




52 




• - 



53 




54 



Figs 49-54. Leptomastix nigrocincta - 49, antenna, variant, 9 ; 50, antenna, variant, 9 ; 51, forewing, 9 ; 52, forewing 
base, 9 ; 53, forewing, apex of venation, 9 ; 54, forewing, d". 



126 



J-M. ANGA AND J.S. NOYES 




i 



56 



57 




-ff 




■■' 



59 




60 



Figs 55-60. Leptomastix nigrocincta - 55, forewing base,cT. Leptomastix abyssinica - 56, antenna, 9 ; 57, forewing, 9 ; 
58, forewing base, 9 ; 59, forewing, apex of venation, 9 ; forewing, cf . 






REVISION OF LEPTOMASTIX 



127 




Figs 61-65. Leptomastix herreni - 61 , antenna, scale-like structure enlarged, d". Leptomastix algirica - 62, antenna, scale- 
like structure enlarged, Cf . Leptomastix dactylopii - 63, antenna, scale-like structure enlarged, cf . Leptomastix jonesi - 64, 
antenna, scale-like structure enlarged, (S . Leptomastix digitariae - 65, antenna, scale-like sstructure enlarged, d\ 



128 



J-M. ANGA AND J.S. NOYES 






69 



70 




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



I 



Bull. not. Hist. Mus. Lond. (Ent.) 68(2): 129-144 



Issued 25 November 1999 



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 

Synopsis 129 

Introduction 129 

Depositories 130 

Acknowledgements 130 

Spilomicrus Westwood 130 

The formosus species group 130 

Key to north-west European species of the formosus species group 132 

S. crassiclavis Kieffer 132 

S. formosus Jansson 134 

S. sanbornei Masner 136 

References 137 

Index 138 

Synopsis. A revision of north-west European species of theformosus 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 5. crassiclavis and S. formosus 
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. Theformosus 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 



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, 



) The Natural History Museum, 1999 



130 



D.G. NOTTON 



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 



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 ( 1 989) 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, l.l-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 ( 9 ) or 3-7 (cT) 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.fiimipennis, 
B. tuberculatus and B. erythropus which also attack 
pipunculid puparia (Lundbeck, 1 922; 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 ldiotypa; 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 

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 

(Spilomicrus formosus group) 2 

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

Antennal flagellum not clubbed (Figs 9, 12, 15); post- 
petiole with 7 sternites (males) 5 

3 Antennal club 5- to sub-6-segmented (Fig. 1 3); 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 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. 
1 8); medial propodeal keel raised in front to form a right- 
angled to acute projection (Fig. 18); body length 2.2-3. 1 
mm 4 

4 Antennal club 7-segmented, with segments 8- 1 2 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 
(Fig. 2) 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 (Fig. 3) 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. 1 6); medial propodeal keel low, 
hardly raised in front (cf. 9 - Fig. 16); body length 2.0- 
2.3 mm sanbornei 

- In frontal view, apex of medial projection of clypeus 
triangular, pointed (cf. 9 9- 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); 
body length 2.1-3.7 mm 6 

6 Mesopleuron 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. 9 - 
Fig. 19); eye height l.Ox as long as malar space; surface 
of eye with sparse hairs 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 

formosus 

Spilomicrus crassiclavis Kieffer 

(Figs 2, 7-9, 17,22) 

Spilomicrus crassiclavis Kieffer, 1911: 788, 797. 

LECTOTYPE9, 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. TYPE HYM. 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 ( 9 ).] 
Spilomicrus pelionNixon, 1980: 17. Holotypecf, Great 

Britain: Surrey (BMNH) [examined]. Syn. n. 

Holotype labels: Holotype [red BMNH type label]; 

Surrey, Horsley, 14. vi. 1930, G. Nixon; Spilomicrus 

pelion Nixon, Typecf, 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 SPILOM1CRUS 



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 .Ox 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. Ox 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 2.03-3.1 1 mm. 

Material examined. Czechia: 9 , Bohemia, Sumava, 
Hurka v Posum, 20.vii.1954 (Hoffer);(f, Moravia, 
Javorina, 800-900 m, 1 0.viii. 1 99 1 , climax eufagion 
for. (Masner) (CNC). Denmark: 9 , north-east Zea- 
land, Faergelunden, 55°51'N 12°02'E, 21.vi.1988 
{Buhl) (DN);d\ north-east Zealand, Boserup Skov, 
55°40'N 12°0rE, 12. vi. 1988 (Buhl) (DN);cT, 
28.vi.1878 (Schlick) (ZM). Finland: 9 , Pohjois- 
Karjala, Tohmajarvi, 6908:660, 17.vii. 1982 (Koponen) 
(SEL). Great Britain: 9 . Stirling, Mugdock, 
ll.vii.[pre-1910] (Cameron) (BMNH) (lectotype); 9, 
W. Ross, Rassal National Nature Reserve, NG845432, 
l-15.vi.1992, Malaise trap (Brown) (NMS); 9 , West 
Glamorgan, Crymlyn Bog, SS6995, 14. vi. 1989 
(Holmes) (DN); 39, Cheshire, Abbots Moss, 
SJ596680, l-19.vi.1990, Malaise trap/swept (Norton) 
(DN); 9 , Cambridgeshire, Chippenham Fen, 
TL650693, 6.vii.l983, Malaise trap, carr/reed bed 
(Field) (DN); 2 9 , Norfolk, Catfield, TG374204, 2 1 .vi- 
5.vii.l988, water trap (DN);cf, Surrey, Horsley, 



134 



D.G. NOTTON 



14.vi.1930 (Nixon) (BMNH) (holotype);cf , Kent, 
Eynsford, 7.vi.l930 (Nixon) (BMNH) (paratype). 
Japan: 9 , Iwate Prefecture, Mount Hayachie, 400 m, 
19-26. vii. 1989 (Makihara & Sharkey) (CNC). 
Norway: cf, R0yken, Kinnertangen, EIS 28 B0, 
vii. 1993, Malaise trap (Hansen) (DN). Sweden: 9, 
Narke, Orebro, 12.vii.1957 (Jansson) (ZI); 9 , R0rvik, 
Lammhult, 15.vii.1986 (Buhl) (ZM);9, 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 Museum 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 5. 
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 5. 
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 5. formosus was in fact S. 
crassiclavis. Teodorescu ( 1 970; 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 9 , Sweden: Orebro, Oset (ZI), here 
designated [examined]. Lectotype labels: Or. Oset, 
10-8 39 A. J.; Typus [red label]; Spilomicrus 



formosus Janss., 9 ; 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. 9 . 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. 

d\ 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. 1 x 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: d\ Mont Rigi, stat. 
se. Hautes Fagnes, 18-24.viii.1983, Universite Liege 
(CNC). Czechia: 9 , Moravia, Lednice, env., 7- 
9. viii. 1 99 1 , yellow pan trap, riparian forest 
(Masner);cf, Bohemia C, Revnice (Masner) (CNC). 
Denmark: 2 9,d\ north Zealand, i v. 1891, each 



mounted with a pipunculid puparium (Schlick) 
(ZM);d\ v.1889; 49, 2d\ iv. 1890, each mounted 
with a pipunculid puparium (Schlick) (ZM); 9 , 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); 9 , Kemin Lappi, Kittila, 7469:409, 
17. viii. 1983 (Koponen) (DN);d\ Etela-Savo, 
Mantyharju, 6795:508, 1. viii. 1981 (Koponen) 
(SEL);d\ Etela-Pohjanmaa, Toysa, 6948:336, 
23.vii.1987 (Koponen) (SEL);d\ Helsinki, 28.vi- 
1 1 .vii. 1 989, Malaise trap (Goulet) (CNC). Germany: 
3d", Bayern, Oberstdorf, 12-29. viii. 1936 (Nixon) 
(BMNH). Great Britain: 2d\ Elgin, Aviemore, 26.vii 
& 5. viii. 1946 (Harwood) (BMNH); 9, 2c? , W. Ross, 
Rassal National Nature Reserve, NG845432, x.1991, 
Malaise trap (Brown) (DN); 2 9 , Cambridgeshire, 
Chippenham Fen, TL6469, 17-24.vi.1985, Malaise 
trap (Field) (DN),d\ Chippenham Fen, TL650693, 
6-20. vii. 1985, Malaise trap (Field) (DN);d\ 
Pembrokeshire, Brynberian Moor, SN106346, 
20. vii. 1987, pitfall trap, Juncus sp., soligenous flush 
(Holmes) (DN);d\ Wiltshire, Savernake Forest, 
SU21366708, 26.vii-16.viii. 1990, Malaise trap 
(DN);d\ Oxfordshire, Weston Fen, SP526194, 22.vii- 
20. viii. 1987, Malaise trap (DN);d\ Surrey, Oxshott, 
9.viii.l930 (Nixon) (BMNH). Ireland:d\ Co. Wex- 
ford, Benroe, Killybegs district, 18.viii.1933 (Stelfox) 
(BMNH). Japan: 2c? , Hokkaido, Tomuraushi area, 
500 m, 13.viii.1996 (Masner) (CNC). Norway: 
2 9.C?, Baerum, Kjaglidalen, EIS 28 AK, 27.vii- 
ll.viii.1990, Malaise trap (Falck) (DN);c?, Rollag, 
Varviken, EIS 35 BV, viii. 1992, Malaise trap 
(Sagvolden) (DN). Russia: 9 , Moscow, Bitsa Park, 
18. viii. 1993 (Kolyada) (RAS);d\ Moscow, 
13.vii.1993 (Kolyada) (RAS). Slovakia:d\ {illeg- 
ible}, Nizke Tatry, Kysla, 10.viii.1989 (Macek) 
(CNC). Sweden: d", Blekinge, Rodeby Gagnekulla, 
56°16'N 15°34'E, 8.ix.l956 (Sundholm) (ZI); 9, 2d", 
Dalarna, Al, 10 km south east of Leksand, 60°42'N, 
15°02'E, 4. viii. 1968 (Sundholm) (ZI); 3d", Dalarna, 
Jarna, 60°33'N, 14°15'E, 3.viii.l968 (Sundholm) (ZI); 
29, Halsingland, Loos, pre-1942 (Sjoberg) (ZI) 
(paralectotype); 5d", Jamtland, Fors, 63°01'N, 
16°37'E, 7. viii. 1964 (Sundholm) (ZI);d\ 
Lulelappmark, Jokkmokk Messaure, 66°43'N, 
20°20'E, 7. viii. 1966 (Sundholm) (ZI); 9 , Narke, 
Orebro, Oset, 59°17'N, 15°10'E, 10. viii. 1939 
(Jansson) (ZI) (lectotype);cf, Narke, Orebro, Oset, 
59°17'N, 15°10'E, 17.viii.1944 (Jansson) (ZI);9, 
Narke, Orebro, 0. Mark, 59°15'N, 15°12'E, 
20.viii.1944 (Jansson) (ZI);d\ Narke, Orebro, 
22.viii.1937 (Jansson) (ZI) (paralectotype); 9 , Narke, 
Orebro, 20. viii. 1939 (Jansson) (ZI) (paralectotype); 9 , 
Narke, Orebro, 16. viii. 1941 (Jansson) (ZI) 
(paralectotype); 29, lid", Narke, Orebro, 
28.vii.1941-20.vii. 1960 (Jansson) (ZI);d\ Narke, 



136 



D.G. NOTTON 



15.viii.1955 (Sundholm) (ZI);o\ Norrbotten, Edefors, 
66°05'N, 20°54'E, 31.vii.1958 (Sundholm) (ZI);d\ 
Norrbotten, Nedertornea, 65°50'N, 24°07'E, 
24.vii.1966 (Sundholm) (ZI);9, Nu., 15.viii.1955 
(Jansson) (ZI);d\ Oland, Gardby, 56°37'N, 16°05'E, 
7.viii.l967 (Sundholm) (ZI);cf, Ostergothland, 
Skedevi, 58°08'N, 15°42'E, 2.viii.l970 (Sundholm) 
(ZI);cT, Skane, Skaralid, 16.vii.1937 (Jansson) (ZI) 
(paralectotype); 3c? , Skane, Yngsjo, 55°57'N, 
14°H'E, 23.vii.1964 (Sundholm) (ZI);d\ Smaland, 
Halltorp Varnanas, 56°30'N, 16°07'E, 8.viii.l967 
(Sundholm) (ZI);d\ Smaland, 20.ii.1943 (Jansson) 
(ZI); 5d\ Sodermanland, Dalaro, Malmen, viii.1976 
(BMNH);d\ Sodermanland, Lanna, 10 km south of 
Strangniis, 59°18'N, 16°54'E, 9.viii.l958 (Sundholm) 
(ZI);cf, 24.vii.1940 (ZI);9, 25.vii.1943 (ZI);cf, 
l.viii.1944 (Jansson) (ZI). Canada: 9,cf, British Co- 
lumbia, Hollyburn, 24.viii.1930 (Whittaker) (BMNH). 
U.S.A.:d\ Washington, Okanagan C, N. Waconde, 
25.vii.1985 (Tinnamore) (BMNH); 9, Virginia, 
Shenandoah N. P., Big Meadow, 1,300 m, vii- 
viii.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 
(Hellen, 1963; Masner, 1991), Germany (Nixon, 1980), 
Great Britain (Nixon, 1980 - males only), Russia 
(Hellen, 1963), Slovakia (Masner, 1991) and Sweden 
(Masner, 1991). 

BIOLOGY. Eleven Danish specimens of 5. 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 
( 1 976). 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 



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 5. 
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. 
Hellen'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 9 , 
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-l.lx 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 



137 



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. 
cf . 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 
l.lx 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: d", Cheshire, 
Abbots Moss, SJ596680, 1-19.6.1990, Malaise trap 
(Notton) (NMS);9,o\ Cheshire, Abbots Moss, 
SJ596680, 19.6-10.7.1990, Malaise trap (Notton) 
(DN). Canada: 9 , Ontario, Guelph, 8-24. vi. 198 1 , pan 
trap (Barber) (CNC) (paratype);cf , Ontario, Rondeau 
Prov. Pk, 9-26. vi. 1980, interception trap (Goulet) 
(CNC) (paratype);d\ Quebec, Gatineau Pk, 28. vi- 
5.vii.l983, beaver pond, interception trap (Denis) 
(CNC) (paratype); 9 , Quebec, Old Chelsea, 5- 
15.vi.1987, NCC woodpile, pan trap (Masner) (CNC) 
(paratype) ;o\ Quebec, 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 5. abnormis but easily distin- 
guished from it by the presence of a malar sulcus, lack 
of pubescence at the base of metasomal syntergite 
(2+3). 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 scutellarpits 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 



D.G. NOTTON 



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. In: Andre, E. (Ed.) 
Species des Hymenopteres 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.) Opredelitelt 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. 



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, Vm(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. Contribu{ii noi la studiul proctotrupoideelor 
(Hymenoptera - Proctotrupoidea). Studii si cercetari de Biologie, 
Seria Zoologie, 22(3): 189-196. 

Teodorescu, I. 1986. Contribuni la cunoa§terea armaturii genitale 
mascule la Ceraphronoidea^i 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 

Basalys 131 
Bibionidae 130 

crassiclavis 129, 132 

Diapriidae 129 
Diapriinae 129 
Drosophila 130 
Drosophilidae 130 

erythropus 131 

formosus 129, 131, 132, 134 
formosus 132 



formosus species group 129, 130. 131, 132 
fumipennis 131 

hemipterus 130 

Idiotypa 131 
integer 129 

Juncus 135 

Muscidae 130 

pelion 129, 132 
Phoridae 130 
Pipunculidae 130 



Pipunculus 131, 136 

Sanbomei 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 SP1LOMICRUS 



139 




ma 



Fig. 1. Spilomkrus stigmaticalis - head, anterior, Cf . Fig. 2. S. crassiclavis - head, anterior, 9 . Figs 3-4. S. formosus - 3, head, 
anterior, 9 ; 4, head, dorsal, Cf . Figs 5-6. S. sanbornei - 5, head, anterior, paratype 9 ; 6, head, lateral, paratype 9 . 



140 



D.G. NOTTON 





Figs 7-9. Spilomicrus crassiclavis - 7, left antenna, posterior, 9 ; 8, left antenna, fourth segment, posterior. d" ; 9, left antenna, posteriord". 
Figs 10-12. S. formosus - 10, left antenna, posterior 9:11, left antenna, fourth segment, posterior, cT ; 12, left antenna, posterior, cT . 
Figs 13-15. S. sanbomei -13, left antenna, posterior, paratype 9 ; 14, left antenna, fourth segment, posterior, paratypeCf ; 15, left 
antenna, posterior, paratype cT. 






REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOMICRUS 



141 




Fig. 16. Spilomicrus sanbornei - mesosoma, lateral, paratype 9 ■ Fig. 17. S. crassiclavis - mesosoma, lateral. O" . 



142 



D.G. NOTTON 



18 




o 





Figs 18 & 20. 

dorsal, 9 . 



Spilomicrus formosus - 18, mesosoma, lateral, 9 ; 20, mesoscutum, dorsal, 9 . Fig. 19. 5. crassiclavis - mesoscutum, 



REVISION OF THE FORMOSUS SPECIES GROUP OF SPILOM1CRUS 



143 




Fig. 21. Spilomicrus fonnosus - North-west European distribution. 



144 



D.G. NOTTON 




Fig. 22. Spilomicrus crassiclavis - distribution. Fig. 23. S. sanbomei - North-west European distribution. 



Bull. not. 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 5BD, UK. 

CONTENTS 

Synopsis 145 

Introduction 146 

Methods 146 

Comments on characters used 147 

Character descriptions, with states 147 

Condensed data matrix with outgroup 150 

Amicotermes Sands 1972 150 

Key to the species of Amicotermes 151 

Descriptions of species 153 

Amicotermes autothysius sp. n 153 

Amicotermes camerunensis sp. n 154 

Amicotermes congoensis sp. n 156 

Amicotermes cristatus sp. n 157 

Amicotermes dibogi sp. n 158 

Amicotermes galenus Sands 159 

Amicotermes gasteruptus sp. n 160 

Amicotermes ivorensis sp. n 161 

Amicotermes mayombei sp. n 162 

Amicotermes mbalmayoensis sp. n 163 

Amicotermes multispinus sp. n 164 

Amicotermes spiculatus sp. n 165 

Phenetic and phylogenetic analysis 165 

Acknowledgements 167 

References 167 

Appendix 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 






5 ZX>(fi 2.°(- 1, 



146 



W.A. SANDS 



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 etai, 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 
etai, 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 10,488nr 2 and live weight biomass up to 123 gm~ 2 
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 



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 forTAxonomy] 
(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 AM1COTERMES 



Ul 



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 morphometries. 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 Amicotennes, 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 codi ng 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 



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 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 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 



W.A. SANDS 



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) in identifying 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 (PI). 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. 



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 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 
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 



149 



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 not apply 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 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 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. 



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 forthe 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; LI, 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; Rl, 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). 



150 



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 LI .Lm 
(range, mean). 

60. Right mandible, complex ratio Ra divided by 
Rl.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 1 1733, 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. 1 54-0.25 mm. Fore tibia moderately to strongly 



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 LI. 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 
Rl.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) 

autothysius 

camerunensis 

congoensis 

cristatus 

dibogi 

galenus 

gasteruptus 

ivorensis 

mayombei 

multispinus 

mbalmayoensis 

spiculatus 



2322121122 
2311122122 
1111121222 
1211121212 
2211121212 
2312121122 
3311211122 
1311121122 
2222221112 
1111121222 
1111121222 
1311122222 
2211131222 



1111132211 
2111122211 
2121222222 
3212132211 
3222232211 
2121232122 
3211122211 
2111132211 
1221122211 
3112232211 
3122232211 
1121332211 
3122132211 



2213333111 
2121111333 
2112222222 
2222222333 

2211111222 
1213333333 
2122222333 
2121111333 
2212222333 
2121111333 
2122222333 
2112222333 
2113212222 



1??????211 
3213212512 
2323212521 
3212122512 
2111112212 
3222212511 
3112111512 
3313121522 
3111111323 
3113122512 
3314121522 
3212212422 
2113112521 



112125??34 
2221321222 
2121242223 
2221242234 
1122343333 
2121132323 
2321264456 
2221221222 
22111???33 
2322332234 
2222272223 
1121364334 
2322243323 



???????6?? 
3412211322 
3422333312 
3322221422 
3373331422 
3322331512 
3434321524 
3317111322 
3423221312 
4311221412 
3271111322 
3443447422 
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 0.413 to 0.434 mm 2 

Head capsule width 0.565 to 0.717 mm, postclypeus 
width 0.300 to 0.395 mm 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 Rl.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 

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 R 1 .Rm, 20. 164 to 26.88 1 . 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 

... mbalmayoensis (page 163, Figs 125-137, 183, 184) 

3 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 size 6 

4 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 

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 

5 

5 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- 
ing scales proximally 

cristatus (pagel57. 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 

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 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 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- 
complete 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 

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 



153 



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

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 (pagel56, figs 29^4-6, 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 

autothysius (page 153, figs 1-15, 169, 170) 

10 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 
spicules ivorensis (page 161, figs 99-1 10, 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 



1 1 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 

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 

gasteruptus (page 160, figs 83-98, 179) 



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 



W.A. SANDS 



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 Ll.Lm 

17.556 to 29.807 (mean 23.846). 
Right mandible, complex ratio Ra divided by Rl.Rm 

20.717 to 34.01 (mean 25.475). 

Comparisons. The Cameroonian species most 
closely 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 



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 2 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. n. 

(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 



155 



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(mean3.624). 
Left mandible, complex ratio La divided by Ll.Lm 

18.546 to 23.477 (mean 21.018). 
Right mandible, complex ratio Ra divided by Rl.Rm 

22.193 to 29.197 (mean 25.539). 



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 PI 
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. multispimis 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 



W.A. SANDS 



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#l), 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. n. 

(Figs 29^16, 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. 



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 Ll.Lm 

17.791 to 37.866 (mean 25.316). 
Right mandible, complex ratio Ra divided by Rl.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 PI 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 



157 



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./!, 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, . 1 3), Congo (Brazzaville): Mayombe, 
Feb. 1997; other Paratype material, Feb. 1997, 4 series 
(Eggleton and Davies) (all in BMNH). 

Amicotermes cristatus sp. n. 

(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 



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 tol4, 
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 



W.A. SANDS 



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 Ll.Lm 
21.913 to 26.987 (mean 24.192). 

Right mandible, complex ratio Ra divided by R 1 .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 PI 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 



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,. 1 1 ), 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 to 1 4, straight and evenly tapered, 
shorter than or equal to least width of ridges in position 






REVIEW OF AMICOTERMES 



159 



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 Ll.Lm 

18.009 to 23.312 (mean 20.937). 
Right mandible, complex ratio Ra divided by Rl.Rm 

20.653 to 30.758 (mean 23.927). 

COMPARISONS. A. dibogi has already been compared 
individually with the species that precede it; however, 
it is so distictive with its long, dilated PI 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 etal.); other 



Paratype material, Ekombitie, Aug. 1995, Nov. 1995, 
July 1996, Aug. 1997, Ebogo, Feb. 1996 (Eggleton et 
al.) (all in 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 tergiles 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 



W.A. SANDS 



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 Ll.Lm 

26.495. 
Right mandible, complex ratio Ra divided by Rl.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 



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 Elizabeth ville, 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 AM1COTERMES 



161 



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

Measurements and diagnostic ratios [ 1 1 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 Ll.Lm 

20.184 to 28.664 (mean 23.214). 
Right mandible, complex ratio Ra divided by Rl.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. 



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, Nov. 1995, Ebogo, Mar. 1996 
(Eggleton et al.) (all in BMNH). 



Amicotermes ivorensis sp. n. 

(Figs 99-1 10, 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 



W.A. SANDS 



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 0.38 1 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 Ll.Lm 

21.554 to 24.308 (mean 22.474). 
Right mandible, complex ratio Ra divided by Rl.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 



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 



163 



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 0.57 1 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 Ll.Lm 

18.707 to 26.77 (mean 22.184). 
Right mandible, complex ratio Ra divided by Rl.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 



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. T 1 . 1 0) Congo (Brazzaville): Mayombe, 
Feb. 1997 (Eggleton and Davies); other Paratype ma- 
terial, Feb. 1 997, 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 Ll.Lm 

22.319 to 27.038 (mean 23.941). 
Right mandible, complex ratio Ra divided by Rl.Rm 

20.164 to 26.881 (mean 24.080). 

COMPARISONS. A. mbalmayoensis has already been 
compared with the preceding species. A. multispinus is 
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. HP.2) Cameroon: Mbalmayo Forest 
Reserve, Ekombitie, Nov. 1996 (Eggleton etal); 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 1 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 



165 



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. 1 34 to 0. 141 mm (mean 0. 1 38). 
Fore tibial index (length over width) 3.732 to 3.974 

(mean 3.834). 
Left mandible, complex ratio La divided by Ll.Lm 

23.999 to 27.622 (mean 25.824). 
Right mandible, complex ratio Ra divided by Rl.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. T 1 .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, 



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 Ll.Lm 

20.586 to 23.569. 
Right mandible, complex ratio Ra divided by Rl.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 



W.A. SANDS 



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 
DELPAUP1 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 
morphometries. 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 



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 is 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 



167 



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 PI, which pushes the enteric valve seating 



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. dibogi in a derived position distant 
from Acholotermes. 



Acknowledgements I 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. HENNIG86 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. 1 99 1 . 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. 

Rohll, 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 tile 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 Trinen'itennes. 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- 
71. 

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 pis. 



168 



W.A. SANDS 



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 in a 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 

l,OM2,OM3,OM4,OM5,OM6,OMll,OM14,OM16, 

OM 1 8, 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 

1,42,43,34,35,36,311,314,315,316,318,324,325, 

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. 1 54-0. 178/0.1 79-0.202/0.203-0.227/0.228-0.250 

49, 0.697-0.781/0.782-0.866/0.867-0.951/0.952-1.036/ 
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 
56,0.489-0.536/0.537-0.583/0.584-0.63/0.631-0.677 
57,0.127-0.147/0.148-0.166/0.167-0.185/0.186-0.205 

58, 2.953-3.255/3.256-3.556/3.557-3.857/3.858^1.157/ 
4.158-4.458/4.459-4.759 

59, 17.556-22.634/22.635-27.7 1 2/27.7 1 3-32.789/32.79- 
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/ 



#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 



169 



#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/ 



#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/ 



#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/ 



#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/ 



#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/ 



#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 



W.A. SANDS 



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>/ 



#50. right mandible index <range, mean>/ 

#5 1 . 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 Ll.Lm 
<range, mean>/ 

#60. right mandible, complex ratio Ra divided by Rl.Rm 
<range, mean>/ 

*ITEM DESCRIPTIONS 

#l.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./ 
#1 1 . Amicotermes mbalmayoensis sp. n./ 
#12.Amicotermes multispinus sp. n./ 
#13. Amicotermes spiculatus sp. n./ 



REVIEW OF AMICOTERMES 



171 




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). 



172 



W.A. SANDS 










Figs 10-15. Amicotermes autothysius. 10-12, enteric valve seating and part of P3; 13-15, enteric valve armature opened 
out to show detail and variation, ridge positions numbered [bar, 1 mm: middle, valve seating; 0.5 mm: bottom right valve 
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. 29-3 1 , 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 




Figs 39-46. Amicotermes congoensis. 39^0, crop with 'bursters' and proventriculus; 41-43, enteric valve seating and part 
of P3; 44-46, 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 










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]. 



REVIEW OF AM ICOTERMES 



111 





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 




Figs 70-75. Amicoterrnes dibogi. 70, 71, crop with 'bursters' and proventriculus; 12-1 A, 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 










Figs 76-82. Amicotermes galenus. 76-78, mandibles with surface view of right molar plate; 79-8 1 , 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 




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 




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 




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 CI 47 Rl 44 
from mhennig*;bb*; two trees 



-24 



:23 



' j, a j, ' Acholotermes 

- — , co 

fc-frfttlr* 5 dibogi 



«| -k O CD 00 



■±±±22 



-±21 






—£-2 camerunensis 
-+-irt"12spiculatus 



4rir20 



■4*419 



m e> 

er 00 



■tt* 17 



t15 



i-irtrt 4 cristatus 



6 galenus 



S Oi Oi * 



I I I I I I I 



3JkW CO U U' 

A J. J. A i. 8 ivorensis 

<D o> u o o> 



-j-t-10 mbalmayoensis 



►18 



162 



-t16 



4r** 14 

S -j *> 



■±±±13 



-4-+-J-11 multispinus 

3 congoensis 

— j,AA 9 mayombei 

fc-7 gasteruptus 

£-1 autothysius 



PRINCIPAL COORDINATES OF AMICOTERMES 



cristatu 



MINIMUM SPANNING TREE 



163 



AcholotQ 




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, lb . . . etc); 
163, Principal coordinates plot of eigenvectors on coordinates 1, 2 & 3, with minimum spanning tree superimposed as a 
broken line. 



REVIEW OF AM1C0TERMES 



SINGLE-LINKAGE TREE OF AMICOTERMES 



Q.45 



0.5Q 



0.55 



0.60 



164 



0.65 

— Acholote 
— dibogi 
j — gasterup 
— autothys 

— mayombei 
— congoens 
— multispi 
— galenus 
— mbalmayo 
— spiculat 
— cristatu 
— lvorensi 

— earner une 



FLEXIBLE-LINKAGE TREE OF AMICOTERMES 



0.16 



0.32 



0.48 



0.64 



165 



^80 

— Acholote 
— dibogi 
— lvorensi 
— gasterup 
— autothys 
— mayombei 
— mbalmayo 
— congoens 
— multispi 
— galenus 
— earner une 
— spiculat 
— cristatu 



COMPLETE-LINKAGE TREE OF AMICOTERMES 



0.15 



0.30 



0.45 



0.60 



166 



0.75 

— Acholote 
— dibogi 
— gasterup 
— autothys 
— mayombei 
— mbalmayo 
— congoens 
— multispi 
— galenus 
— camerune 
— lvorensi 
— 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 



Q.3 



Q.4 



0.5 



Q.6 



167 



0.7 



•Acholote 
"dibogi 

•gasterup 

•autothys 

■mayombei 

'congoens 

■multispi 

■galenus 

■mbalmayo 

• earner une 

■ spiculat 

cristatu 

lvorensi 



NEIGHBOUR-JOINING TREE OF AMICOTERMES 



168 



10 

mi 

— Acholote 
— gasterup 

— autothys 
— mayombei 

— congoens 
-multispi 
— lvorensi 

— galenus 

— mbalmayo 

— earner 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 



191 




/V ' f i¥ 



/^ villi/' Vlll 



169 



■RHHi 




ft'Wvi 



170 









iW 




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 





%f 






| 



182 



- /^ Ml 






184 












4fL 











186 



Figs 181-186. Amicotermes, enteric valve armature, valves slit and opened out; 181, 182, A. mayombe'v, 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— 

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No. 3 Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new taxa 

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No. 4 The Afrotropical dacetine ants (Formicidae). B. Bolton. 1983. Pp. 267^16, 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. 1 Gelechiid moths of the genus Mirificarma. L.M. Pitkin. 1984. Pp. 1-70, 1 12 figs, 2 tables. £10.00 

No. 2 Macronematine caddisflies of the genus Amphipsyche (Trichoptera: Hydropsychidae). PC. 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. £39.75 

Volume 49 

No. 1 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 Eitchorthippus, 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. 

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No. 4 The Pimplinae, Xoridinae, Acaenitinae and Lycorininae (Hymenoptera: Ichneumonidae) of Australia. I.D. 

Gauld. 1984. Pp. 235-339, 100 figs, 18 maps. £15.75 

No. 5 The Palaearctic species of Ascogaster ( Hymenoptera: Braconidae). T. Huddleston. 1984. Pp. 341-392, 79 

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Volume 50 

No. 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). PS. Broomfield. 1985. Pp. 1- 

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No. 2 Nymphal taxonomy and systematics of the Psylloidea (Homoptera). I.M. White & I.D. Hodkinson. 1985. 
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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 

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No. 2 The phylogeny, classification and evolution of parasitic wasps of the subfamily Ophioninae 

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No. 4 The British and some other European Eriococcidae (Homoptera: Coccoidea). D.J. Williams. 1985. Pp. 

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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 
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No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986. Pp. 183-242, 68 
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No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera. J. 

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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 & GA. Holloway. 1986. 

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No. 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on the 

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No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy 

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No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble. 1986. Pp. 

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Volume 54 

No. 1 Studies on the Old World species of Holoth rips (Thysanoptera: Phlaeothripidae). S. Okajima. 1987. Pp. 1- 

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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. 263^152, 100 figs. £36.50 

Volume 55 

No. 1 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. 393^124, 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. 31 1^437, 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 & PC. 
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. 1 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 leaf-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. 1 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. 1 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. 



93 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 
1 29 A revision of the north-west European species of the formosus species 

group of Spilomicrus (Hymenoptera: Diapriidae) 

David G. Notton 
145 A review of the soldierless African termite genus Amicotermes Sands 

1972 (Isoptera, Termitidae, Apicotermitinae) 

W.A. Sands 



ENTOMOLOGY SERIES 

Vol. 68, No. 2, November 1999