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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 89 Band

June 1982 Junie

Pant 3 Deel

REVIEW OF ANOPLOLEPIS

WITH REFERENCE TO MALE GENITALIA,

AND NOTES ON ACROPYGA

(HYMENOPTERA, FORMICIDAE)

A. J. PRINS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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ISBN 0 86813 031 1

Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

REVIEW OF ANOPLOLEPIS WITH REFERENCE

TO MALE GENITALIA, AND NOTES ON ACROPYGA

(HYMENOPTERA, FORMICIDAE)

A. J. PRINS

South African Museum, Cape Town

(With 16 figures)

[MS accepted 18 February 1981]

ABSTRACT

Pugnacious ants are abundant in orchards and vineyards, are predacious and, together with

species of Acropyga Roger, tend hemipterons. Only two species of pugnacious ants, Anoplole-

pis custodiens (Smith) and A. steingroeveri (Forel), and one species of Acropyga are present in

South Africa. Male genitalia are employed in this short review to separate the species. South

African species of Acropyga have four to five segmented maxillary palpi. Anoplolepis braunsi

(Forel) is a synonym of A. steingroeveri (Forel), and Acropyga rhodesiana Santschi a synonym

of A. arnoldi Santschi.

CONTENTS

PAGE

Introd Uctlones pearance tra eed eee eee 215

GenuspAroplolepisisantschie 3:30 no tg hee ne eee 217

Anoplolepis custodiens (Smith)... seston eeke 218

Ano plolepisinuptialisn(Sattschi)\ee ca. sase ele tee ae 222

Anoplolepisssteingroeveri (Forel) ii.s nn 42s. a ose ter 226

DisinibhutioniowAnoplolepisen ware taunt ciate Dill

INaturalicnemicsio& Arioplolepistn sma cias tee ert 231

Key for identification of males and females of pugnacious

An tsioMSouthyA fica. gags eva ctr es alse Aree nl eeae aownomnters 235

GenusvAcropycapROCehem ps Teeter eae erie 235

AcCropyeMarnnoldy Santschiaem reer reac ont er 238

AXCKnOWIEd PEMentSe-r ee rete orci Sota 246

FRELELELICES HY Srey leche 1) sa este Olea tS treks aon besa as i 246

INTRODUCTION

Pugnacious ants are well known for their aggressiveness and fast zig-zag

movements on the ground when disturbed. During the early summer and

autumn labourers in orchards and vineyards are often so molested that they are

forced to stop their work. Fortunately these ants seldom enter houses.

As in the case of the Argentine and brown house-ant, the pugnacious ants

and ants of the genus Acropyga are fond of honeydew, which forms an

important part of their diet. They therefore tend aphids, coccids and their kin

ZAS

Ann. S. Afr. Mus. 89 (3), 1982: 215-247, 16 figs.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

for the sake of the honeydew produced by these insects. The ants soliciting

these hemipterons are not always as innocent as they appear to be, as some of

the trophobionts have to be sacrificed under adverse conditions to provide for

the protein demands of their guardians. Apparently the ants are merely cheated

by their trophobionts, which are mistaken for donor ants (Kloft 1959; Wilson

1971).

The activities of the pugnacious ants are largely influenced by temperature

and relative humidity (Steyn 1954) and they will even visit trees during the

winter months to collect honeydew when the days are warm. The medium or

larger workers store the honeydew in their crops and during cooler weather this

is then apparently metabolized to fat (Smit 1964). However, it was found that

in certain species of ants the honeydew received from foraging workers is

stored by the repletes during cooler conditions. As soon as the temperature

increases, resulting in a higher rate of metabolism, the direction of the flow is

reversed (Wilson 1971). Honeypot ants with extreme forms of repletes are

therefore usually found in arid or semi-arid regions. In orchards and vineyards

where honeydew is fairly easily obtained, it was observed that there are many

more minors and medium-type workers present in a nest than in nests found in

the veld where other food sources have to be utilized.

In the feeding process of the ants the natural enemies of the trophobionts

are disturbed to such an extent that predation and parasitism are obstructed. In

citrus orchards in particular, populations of pugnacious ants feeding on

honeydew from aphids and soft scales usually result in coincident red scale

Aonidiella aurantii (Maskell), infestations, directly as a result of the influence

of the ants on the natural enemies.

We are, however, inclined to look only at the injurious effects of the ants,

whereas they also play an important part in the destruction of noxious insects,

and problems often arise when the natural balance between existing species is

upset. Steyn (1954) lists a whole series of arthropods which form part of the

diet of the common pugnacious ant, and which include the harvester termite,

Hodotermes mossambicus, cockchafers, false codling-moth, and bollworms.

These ants also play an important role as pollinators of flowers. In recent

surveys in fynbos in the southern Cape, for instance, the black pugnacious ant

A. steingroeveri, was found in very large numbers on Protea compacta.

According to their behavioural pattern, the pugnacious ants (and probably

also Acropyga) are unicolonial, although in a few cases observed in northern

Transvaal a whole series of small colonies was found in a rather limited area;

disturbance of these colonies, however, resulted in fierce fighting. Steyn (1954)

and Louw (1968) discuss the biology and ecology of the common pugnacious

ant and information on the other species is given by Myers (1957) and Prins et

al. (1979).

The following abbreviations have been used in the text (for further

reference see Prins (1973)). The nomenclature of the wing venation is accord-

ing to Wheeler (1960).

ANOPLOLEPIS AND ACROPYGA DAG)

m —(median) —cells of front and hind wings

w —(first cubital)

CI —cephalic index (head width/HL x 100)

CL —clypeus length

CLI —clypeal index (clypeal width/CL x 100)

cub —(cubital)

d —(discoidal)

oo ae cxicromedian) —veins of front and hind wings

m+ cua

r | —(marginal or radial)

tr —(transversomedian)

cu —cuspis

di —digitus

ED —distance between compound eyes

HFI —hind femur index (HFL/WL x 100)

HFL—hind femur length

HL —head length

L —length from anterior margin of clypeus to base of abdomen

lab —labrum

MFL—middle femur length

p —penis lobes

PI —petiolar index (petiolar width/PL x 100)

SI —scape index (SL/head width x 100)

SL —scape length

st —gonostipes

—thoracic index (thoracic width/WL x 100)

—total length of body

—volsella

—length of truncus

goes

Genus Anoplolepis Santschi

Arnold (1922) includes the three species of pugnacious ants viz., Anoplole-

pis custodiens, A. steingroeveri, and A. braunsi in the subgenus Zealleyella on

account of their very convex epinotum. The two species A. braunsi and

steingroeveri are distinguished only by the thickness of the scale of the petiole,

the distinctness of the mesometanotal suture, and the shape of the sides of the

head. To date no alates of braunsi have been collected, only workers, which in

all respects agree with the workers of A. steingroeveri. A. braunsi therefore

appears to be a junior subjective synonym of A. steingroeveri; Forel described

the latter species in 1894 and the former in 1913.

There is also a fourth species, viz. A. nuptialis, which is known only from

its alates, and which lives as an inquinilist in the nests of the common

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

pugnacious ant. Both males and females were observed, but workers are

unknown, and they are therefore probably truly inquilinous. It appears that the

alates of this species enter the nests (mating may occur in or near the nest) and

are then accepted by the workers of the common pugnacious ant, which later

destroy the host alates. Only one case is known where they penetrated the nest

of the black pugnacious ant.

Anoplolepis custodiens (Smith, 1858), common pugnacious ant

Worker (Figs 1A-B, 2A-D)

TL 3-10 mm; L 2,0-5,0 mm; HL 0,76-2,2 mm; ED 0,48-1,64 mm; CL

0,24-0,80 mm; SL 0,96-2,44 mm; WL 1,08-3,30 mm; MFL 0,80—-2,56 mm;

HFL 1,02-3,0 mm; PL 0,16-0,40 mm; CI 94,7-104,5; SI 133,3-106,1; CLI

200,0; CTI 70,4-66,7; TI 44,4-44,8; PI 125,0-190; HFI 94,4-90,9.

Polymorphic, colour yellowish-brown to reddish-brown or even somewhat

darker with abdomen dark-brown to almost blackish-brown, therefore easily

confused with workers of the black pugnacious ant. Separated from latter species by

chequered pattern on abdomen, caused by reflection of light on pubescent hairs,

which are arranged in two different directions on each side. Few pilose hairs also

present, particularly on head, apical borders of abdominal segments and on femora.

Anterior border of cypeus angular in the middle, acutely so in some

specimens; in lateral view less convex than in black pugnacious ant. Scale of

petiole rather flattened in larger workers and fairly deeply emarginate above;

scale narrowed dorsally in minors and emargination generally almost obsolete.

The chequered pattern on abdomen and form of the clypeus are not as

distinct as in the majors or media and this caste is therefore difficult to identify

in the absence of larger workers. The convex epinotum is characteristic of the

workers of the subgenus Zealleyella, and seen from the side it is almost as high

as the promesonotum in the majors; in the minors it may be much higher.

Female (Fig. 3A—B)

Wing-span about 27,4 mm; TL 13,3-15,8 mm; L 6,5-7,3 mm; HL 2,2 mm;

ED 1,9 mm; CL 0,7 mm; SL 2,3 mm; WL 4,5-5,2 mm; MFL 2,4-2,6 mm; HFL

2,8-2,9 mm; PL 0,44-0,48 mm; CI 113,6; SI 92,0; CLI 242,8-257,2; CTI

48 ,9-42,3; TI 68,9-67,3; PI 216,7—-241,7; HFI 62,2-55,6.

It has the general features of the females of the subfamily Formicinae, with

a distinct chequered pattern on the abdomen and a wide and fairly deep

emargination on the dorsal edge of the scale of the petiole. Colour more or less

as in the worker, the wings ochreous-yellow with darker subcostal cell; the

abdomen sometimes much darker. Clypeus also angular in the middle as in the

case of the workers.

Wing venation as in Figures 3A, 4A, and although there are minor

variations in the venation of the same individual, the vein m+ cua of the hind

wing much shorter than the preceding cell Ia; in the specimens examined, this

vein is shorter than half the length of the cell.

ANOPLOLEPIS AND ACROPYGA 219

Fig. 1. Anoplolepis custodiens.

A. Major worker, dorsal view. B. Major worker, left lateral view.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

sere we

Fig. 2. Anoplolepis custodiens.

A. Median worker, dorsal view.

B. Median worker, left lateral view.

C. Minor worker, dorsal view.

D. Minor worker, left lateral view.

ANOPLOLEPIS AND ACROPYGA DOM

Fig. 3. Anoplolepis custodiens.

A. Female, dorsal view. B. Female, left lateral view.

hp ANNALS OF THE SOUTH AFRICAN MUSEUM

Male (Figs 4A-C, 9A-B, 10D-F, 11C-D)

Wing-span about 19,0mm; TL 9,9-10,8mm; L 3,6-3,7 mm; HL

1,6-1,64 mm; ED 1,2 mm; CL 0,44-0,48 mm; SL 2,2 mm; WL 3,5-3,8 mm;

MFL 2,5-2,7 mm; HFL 2,8-3,0 mm; PL 0,4 mm; CI 112,5-109,8; SI 12,2; CLI

254,5-233,3; CTI 45,7—-40,3; TI 57,9-62,9; PI 190-200; HFI 80,0-78,9.

Slightly smaller than the female and, as in the case of the latter, they

resemble the males of the subfamily Formicinae. Indistinct chequered pattern

present on the abdomen, although the direction in which the pubescent hairs

are arranged is fairly clearly indicated. Generally darker in colour than female,

more slender, and the clypeus also angular in the middle. Scale of petiole with

indistinct emargination on dorsal edge. Labrum as in Figure 9A.

Although the vein m+ cua of the hind wing (Fig. 3A) is much longer in

most specimens seen, it is not longer than half the length of cell Ia.

External male reproductive organs

The last complete visible segment in the abdomen of the male is the sixth

(the true eighth), the seventh (true ninth) being reduced to a ring-like sclerite

(also known as gonocardo or lamina annularis), which is membraneous dor-

sally, but on the ventral side it is flattened, plate-like and sclerotized. In A.

custodiens the flattened plate is fairly deeply and angularly emarginate behind.

The two lobes formed by the emargination are thinner than the rest of the plate

and are ventrally covered with fine hairs. The plate itself is about twice as wide

as long. Penis or aedeagus situated medially and characterized by its two lobes;

it is somewhat shorter than external lobes, which cover reproductive organs and

are also known as gonostipes (gonoforceps or parameres). In lateral view the

apices of gonostipes are round and broad. On each side and somewhat below

penis is a globular volsella with an outer appendage or cuspis and an inner

shorter digitus. Both digiti and cuspides are devoid of denticles, except in case

of some individuals where there are a few tubercles, particularly on apices of

cuspides. Each volsella has some fine hairs on ventral side.

Material examined

322 Bulawayo, June 1912, G. Arnold; 422 Ellis pass, 2.4.69,

A. Barnard; 3¢¢ Bulawayo, 30.1.1912, G. Arnold; 4dd Ellis pass, 2.4.69,

A. Barnard; 9 8 8 Matopos, 5.11.44, G. Arnold; 12 8 §, Du Toitskloof Pass,

29.5.74, A. J. Prins; 7 8 8 Ellis pass, 2.4.69, A. Barnard.

Anoplolepis nuptialis (Santschi, 1916)

Female

Wing-span about 25,2mm; TL 11,6-13,6mm; L 4,9-5,1mm; HL

1,8-1.9mm; ED 1,6-1,7mm; CL 0,44-0,48mm; SL 1,6-1,9mm; WL

3,6-3,7 mm; MFL 1,9mm; HFL 2,4-2,3 mm; PL 0,36-0,4 mm: €I

115,6-115,8; SI 76,9-86,4; CLI 291,7-318,2; CTI 50,0-51,4; TI 63,9-64,9; PI

150,0-260,0; HFI 63,9-64,9.

ANOPLOLEPIS AND ACROPYGA pep)

Fig. 4. Anoplolepis custodiens.

A. Male, dorsal view. B. Male, left lateral view. C. Male genitalia, seen from the rear.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

As a previous description of the female could not be traced, it is described

here in more detail than in the case of the other species. Very similar to female

of A. custodiens, but lighter in colour, generally yellowish-brown to

brownish-yellow, including the legs and abdomen. Chequered pattern on

abdomen rather inconspicuous, although direction in which the pubescent hairs

are arranged, is fairly distinct. Whole body including legs and antennae covered

with an almost silvery pubescence, hairs on the antennae shorter than on body

and legs. A few pilose hairs present, particularly on the head, posterior borders

of abdominal segments, and on femora.

Head somewhat wider than long, slightly narrower in front than behind,

the sides almost straight. Clypeus angular in middle, however, the angle not as

acute as in A. custodiens. Scape of the eleven-segmented antennae extending

behind hind margin of head by less than half its length. Compound eyes large,

about one-quarter the length of the head and situated far back, on the posterior

half as in other species. Ocelli large, larger in relation to size of head than in A.

custodiens. Frontal carinae short, ending at about middle of compound eyes.

Mandibles as in custodiens, with eight to nine sharp teeth, the apical ones long

and pointed.

Truncus more or less as in A. custodiens, somewhat wider than head, scale

of petiole varied, usually wider above than below, when seen from the rear;

emargination of dorsal edge varying from obsolete to deep and U-shaped. In

some specimens it is even widely emarginate on each side, thus appearing

bilobed on either side of median emargination (Fig. 5D). Wings more or less as

in latter species, but vein m + cua of hind wing long, usually as long as half the

length of cell Ia or even longer. In most specimens seen the exteromedian vein

of front wings, below stigma, more sinuous than in A. custodiens or A.

steingroeveri, and with a thickening in about the third of its length from the

median cell (also present in some males—Fig. 5A). Abdomen only slightly

wider than truncus and therefore smaller than in A. custodiens.

Male (Figs 5A-C, 9C-D, 10A-C, 11E-F)

Wing-span about 19,9 mm; TL 9,1-11,5 mm; L 3,9-4,3 mm; HL 1,4 mm;

ED 1,1-1,2 mm; CL 0,4 mm; SL 1,9 mm; WL 3,0-3,2 mm; MFL 1,8-2,0 mm;

HEL 2,1-235mm; PL 0,)3-0;4mm; Cl 121,4; SI 11158; Cli 27550

43 ,8—46,7; TI 66,7—68,8; PI 280,0-325; HFI 70,0—78,1.

Much lighter or paler in colour than males of A. custodiens, usually

light-brown to pale yellowish-brown, abdomen more robust than in latter

species. Chequered pattern on abdomen almost obsolete due to pubescent

hairs, which are not arranged in regular rows as in A. custodiens. Antennae

twelve-segmented as in the other two species. Scale of petiole shallowly

emarginate above; clypeus not as angular in middle as in A. custodiens, rather

arcuate. Labrum as in Figure 9C. In most specimens examined the vein m + cua

in hind wing longer than half the length of cell Ia, and the stem of marginal (or

radial) and cubital veins in fore wings long (Fig. 5A). In most A. custodiens

ANOPLOLEPIS AND ACROPYGA

” Kew, Tre

WS Neatly

Madd

iN Mii

Bi i i

})

I |

i i

0 ]

ees ere

Fig. 5. Anoplolepis nuptialis.

A. Male, dorsal view.

B. Male, left lateral view. C. Male genitalia, seen from the rear.

D. Scale of petiole seen from the rear.

225

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

males examined there is no stem, the above-mentioned veins arise directly from

first cubital cell (Fig. 4A); in some specimens, however, a short stem is present.

External male reproductive organs

Males of this ant can easily be separated from those of the common

pugnacious ant by the long gonostipes, which are almost triangular in lateral

view and tapering towards the apices which are narrowly rounded, almost as in

A. steingroeveri. Penis lobes much shorter than the gonostipes, but somewhat

longer than appendages of volsellae in most specimens seen. In this case the

digiti and cuspides are of equal length, with the former much more strongly

developed than the latter. Both structures are simple, without any denticles

and, as in A. custodiens, volsellae are somewhat globular and ventrally covered

with a few fine hairs. Annular plate somewhat shorter than in A. custodiens,

but wider, about three times wider than long; posterior emargination wider and

shallower. Ventrally it is covered with fine hairs. ,

Its general characters and external genital organs indicate that this species

is related to both the common and the black pugnacious ant, but it is

apparently closer to the first. It is sometimes extremely difficult to separate the

females of the two species.

Material examined

222 Willowmore, H. Brauns; 522 Du Toitskloof Pass, 15.5.74, V. B.

Whitehead; 422 Du Toitskloof Pass, 29.5.74, A. J. Prins; 4¢¢ Willow-

more, April 1917; 1d Boesmansberg, Feb. 1940, A. J. Hesse; 2d¢ Du

Toitskloof Pass 15.5.74, V. B. Whitehead; 1 d Caledon, 19.3.73, A. J. Prins.

Anoplolepis steingroeveri (Forel, 1894), black pugnacious ant

Worker (Fig. 6A—B)

TL 2,9-8,0 mm; L 1,80-3,90 mm; HL 0,76-1,80 mm; ED 0,52-1,36 mm;

CL 0,22-0,60 mm; SL 0,88-1,64 mm; WL 1,16-2,24 mm; MFL 0,76-1,64 mm;

HFL 1,04-2,08 mm; PL 0,14-0,36 mm; CI 100,0-111,1; SI 82,0-115,8; CLI

213,3-272,7; CTI 65,5-80,4; TI 41,4-55,4; PI 142,9-188,9; HFI 89,7-92,9.

Polymorphic as in case of the common pugnacious ant and usually some-

what darker in colour, varying from reddish-brown to almost piceous-brown or

blackish-brown, and also more shiny. In major workers sides of the head, seen

from above, more convex than in A. custodiens; clypeus in lateral view also

more convex medially, its anterior border, seen from above, more arcuate.

The major workers are fairly easily distinguished from those of A. cus-

todiens by the absence of the chequered pattern on the abdomen due to the fact

that the pubescent hairs are arranged only in one direction on each side of the

median line. On the abdomen of the minors, however, there are less pubescent

hairs and they are also more shiny; they are therefore sometimes confused with

the workers of the black sugar-ant Acantholepis capensis Mayr, or even with

ANOPLOLEPIS AND ACROPYGA DOF),

Fig. 6. Anoplolepis steingroeveri.

A. Major worker, dorsal view. B. Major worker, left lateral view.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

those of the brown house ant, Pheidole megacephala Fabricius. A few pilose

hairs occur on the body of all workers, as in the case of the other species.

Female (Fig. 7A—B)

Wing-span about 24,9mm; TL 11,6-12,6mm; L 5,2-6,4mm; HL

1,76-1,96 mm; EK 1,56-1,70 mm; CL 0,44-0,52 mm; SL 1,6-1,7 mm; WL

3,6-4,6 mm; MFL 1,92-2,1 mm; HFL 2,36-2,6mm; PL 0,4mm; CI

113,6-117,4; SI 73,9-80,0; CLI 300,0-318,2; CTI 42,6-48,9; TI 58,7-63,9; PI

250,0; HFI 56,5-65,6.

Rather similar to female of A. custodiens, but smaller, and usually darker

in colour, mostly brown to dark-brown; abdomen devoid of any chequered

pattern, pubescent hairs arranged in one direction only on each side of median

line. Generally somewhat more shiny than in the latter species and as in the

case of the workers, the clypeus more convex, its anterior border more arcuate

when seen from above.

Scale of petiole deeply emarginate above, emargination V-shaped; hind

wing similar to that of A. nuptialis, vein m + cua long, in most specimens seen,

almost as long as the cell Ia. In specimens at hand, discoidal (or subdiscoidal)

vein (Fig 8A) in front wings more weakly developed than in both A. custodiens

and nuptialis and usually indistinct or even obsolete.

Male (Figs 8A-C, 9E-F, 11A-B)

Wing-span about 15,7 mm; TL 6,6-6,9 mm; L 3,6 mm; HL 1,12-1,16 mm;

ED 0,80-0,84 mm; CL 0,28 mm; SL 1,30-1,36 mm; WL 2,6-2,7 mm; MFL

1,80 mm; HFL 1,92 mm; PL 0,24-0,28 mm; CI 110,7-117,2; SI 100,0-104,8;

CLI 300,0-328,6; CTI 42,9-43,1; TI 61,5-62,2; PI 228,6-250,0; HFI 71,1—73,1.

Smaller than male of A. custodiens, sombre coloured, abdomen without

chequered pattern, pubescent hairs more or less arranged as in female. Scale of

petiole deeply emarginate, emargination V-shaped; clypeus and wings more or

less as in female. In some specimens seen the marginal and cubital veins in

front wings connected by a short stem to first cubital cell, rather similar to that

of A. nuptialis.

External male genital organs

Differ from those of both A. custodiens and nuptialis in the longer penis

lobes, which are longer than volsellae and gonostipes; latter short and seen

from side almost resembling those of A. nuptialis. Both digiti and cuspides

equally developed and more or less of same length. Each digitus with fine

denticles or tubercles on its external side; cuspides each with a row of fine

denticles on its distal half. Volsellae in this case flattened and only slightly

convex on inner side; also covered with a few fine hairs as in case of the other

two species.

ANOPLOLEPIS AND ACROPYGA 229

Fig. 7. Anoplolepis steingroeveri.

A. Female, dorsal view. B. Female, left lateral view.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

v WE Wire

NY Yh Mi)

Fig. 8. Anoplolepis steingroevert.

A. Male, dorsal view. B. Male, left lateral view. C. Male genitalia, seen from the rear.

ANOPLOLEPIS AND ACROPYGA Dail

Material examined

322 Willowmore, Dec. 1912, H. Brauns; 12 Du Toitskloof Pass,

25.1.50, A. J. Hesse; 1d Willowmore, Dec. 1912, H. Brauns; 266 Stan-

ford. 20!3//2) Ae Prins: ll oo (Cape. Towns) 3:4.45-, Acer sllesses S100

Willowmore, Dec. 1912, H. Brauns; 25 $2 Kleinmond, 23.4.60, A. J. Prins.

DISTRIBUTION OF ANOPLOLEPIS

Both the common and the black pugnacious ants are widely distributed

throughout the Republic of South Africa as well as in South West Africa and

Angola. The common pugnacious ant is also found in Zimbabwe and other

central African states, but according to data collected during surveys of ants of

South Africa, it appears as if the black pugnacious ant prefers the drier parts of

the country. It is, therefore, the most common species in the north-west Cape

and certain parts of the Little Karoo; it is particularly abundant in the Hex

River valley, but it is subordinate to the common pugnacious ant in the eastern

Cape and in northern Transvaal. The black pugnacious ant is also common in

western Transvaal, but to date it has not been collected in eastern Transvaal

and in Natal.

Alate forms of the parasitic A. nuptialis were observed only during the late

summer and winter months in the higher parts near Willowmore, near Pletten-

berg Bay, in the vicinity of Caledon and Bredasdorp, and in the Du Toits Kloof

Pass.

NATURAL ENEMIES OF ANOPLOLEPIS

Steyn (1954) gives a list of natural enemies, including ground beetles

(Carabidae), tiger beetles (Cicindelidae) certain flies (Calliphoridae, e.g. Ben-

galia and possibly also Rhina spp. and Asilidae), and spiders as well as different

species of ants.

In their effort to obtain honeydew, these ants are strongly opposed by

other competitors such as the brown house-ant, Pheidole megacephala (Fabri-

cius) and the related P. tenuinodis Mayr. In the south-eastern Cape, P.

megacephala is replaced mainly by P. tenuinodis and the Argentine ant,

Iridomyrmex humilis (Mayr).

Vertebrates also play an important role, particularly geckos, toads, and

lizards. The most important natural enemies, however, apart from other ants

(Forel states that the most dangerous enemies of ants are other ants), seem to

be birds and, in addition to those listed by Steyn (1954), the following, feeding

mainly on pugnacious ants according to crop analysis, may also be included:

Ant-eating chat—Myrmecocichla formicivora (Vieillot)

Black korhaan—Eupodotis afra (Linnaeus)

Blue korhaan—Eupodotis caerulescens (Vieillot)

Crowned plover—Vanellus coronatus (Boddaert)

Coqui francolin—Francolinus coqui (Smith)

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. A. Anoplolepis custodiens, labrum.

B. Anoplolepis custodiens, head of male from above. C. Anoplolepis nuptialis, labrum.

D. Anoplolepis nuptialis, head of male from above. E. Anoplolepis steingroeveri, male

genitalia, dorsal view. F. Anoplolepis steingroeveri, male genitalia, ventral view.

ANOPLOLEPIS AND ACROPYGA 233

Fig. 10. Male genitalia.

A. Anoplolepis nuptialis, dorsal view. B. Anoplolepis nuptialis, ventral view. C. Anoplolepis

nuptialis, annular plate, ventral view. D. Anoplolepis custodiens, dorsal view.

E. Anoplolepis custodiens, ventral view. F. Anoplolepis custodiens, annular plate, ventral

view.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Redwing francolin—Francolinus levaillantii (Valenciennes)

Swainson’s francolin—Francolinus swainsonii (Smith)

Greater blue-eared starling—Lamprotornis chalybaeus (Hemprich &

Ehrenburg)

Helmeted guinea-fowl—Numida meleagris (Linnaeus)—apparently feeds

on a large variety of ants, mainly pugnacious, harvester and ponerine

ants

Kori bustard—Ardeotus kori (Burchell)—feeds mainly on the common

pugnacious ant

Red-billed hornbill—Tokus erythrorhynchus (Temminck)

Yellow-billed hornbill—Tokus flavirostris (Rtppell)—both these birds feed

on a variety of insects, including almost all known ant species

Red bishop—Euplectes orix (Linnaeus)

Social weaver—Philetairus socius (Latham)

Temminck’s courser—Cursorius temminckii Swainson

Steppe buzzard—Buteo buteo (Linnaeus) vulpinus Gloger—in contrast

with the nominate subspecies, which is found in the eastern parts of

the Cape, east Africa, and Ethiopia, this subspecies is a migrant from

Europe and Asia and appears almost throughout South Africa during

the summer; it was found to feed on the common pugnacious ant,

mainly on the workers

The following birds may be regarded as myrmecophagous and usually

include pugnacious ants in their diet:

Bennet’s woodpecker—Campethera bennettii (Smith)—distributed

throughout the northern parts of South Africa

Knysna woodpecker—Campethera notata (Lichtenstein)—distributed

throughout the southern and eastern parts of South Africa

Ground woodpecker—Geocolaptes olivaceus (Gmelin)—widely distributed

in the south-western, southern and south-eastern parts of South

Africa to Transvaal

Red-throated wryneck—Jynx ruficollis Wagler—eastern Province to Trans-

vaal, Botswana, and further north

Pied starling—Spreo bicolor (Gmelin)—widely distributed throughout the

southern parts of South Africa

Various birds are known to utilize ants in their preening activities. This

phenomenon, also known as anting, is, however, rare in South African birds,

although it was observed by Brown & Newmann (1974) and Whyte (1981) in

the case of the previously mentioned greater blue-eared starling and in the

black-eyed bulbul, Pycnonotus barbatus (Desfountaines), and by Colahan

(1981), in the Cape white eye, Zosterops pallidus Swainson.

The common pugnacious ant is apparently generally included in this process

and it is possible that large numbers of the workers may be destroyed in this

way.

ANOPLOLEPIS AND ACROPYGA 235

KEY FOR IDENTIFICATION OF MALES AND FEMALES

OF PUGNACIOUS ANTS OF SOUTH AFRICA

1. Penis lobes of male longer than gonostipes; digiti and cuspides with denticles. Vein m + cua in

hind wing of both males and females long, about as long as half the length of the cell Ia. Abdo-

men of both sexes (in dry specimens) without any chequered pattern ........ A. steingroeveri

— Penis lobes shorter than gonostipes; digiti and cuspides usually without denticles. A more or

less distinct chequered pattern on the abdomen of dry specimens, particularly in females; or

if chequered pattern is indistinct, the vein m+cua in the hind wing of both sexes long,

almost as long as half the length of the cell Ia or even longer

2. Gonostipes long, almost triangular when seen from the side, their apices narrowed or

pointed; the digiti stronger developed than the cuspides; the annular plate about three times

wider than long and widely and shallowly emarginate behind. Indistinct chequered pattern

on abdomen of dry specimens, particularly of female; vein m + cua in the hind wing of both

sexes long, about half the length of the cell Ia or even longer................ A. nuptialis

— Gonostipes shorter, their apices broad and round when seen from the side, digiti and

cuspides equally developed; annular plate longer and narrower, about twice as wide as long;

the posterior emargination deep and V-shaped. Distinct chequered pattern on abdomen of

dry specimens of both sexes, and the vein m + cua in hind wing short, shorter than half the

Fensiho nthe cellar tic dar geri comes oo cia des ecsennvssdore Bere Mahe oe eee A. custodiens

Genus Acropyga Roger

The South African species of this genus are small, seldom seen and

therefore are not as often collected as other ant species. After various efforts,

however, some specimens were found in the western Cape near Gans Bay, near

Saldanha, and near Paternoster. They are hypogaeic and fairly slow-moving

and apparently appear only during the late autumn and winter. Very little is

known about the habits of our species, but information on Neotropical forms

seems to indicate that these ants live on the honeydew produced by hemipter-

ons, which are said to be even carried off by the females during their nuptial

flight (Buinzli 1935); apparently they cause considerable damage overseas to

coffee by transplanting the aphids from infested to healthy plants. It is also

suggested that these ants obtain extra protein by devouring some of their

trophobionts.

A small sample, which includes males and alate females, was collected at

Jacobs Bay near Saldanha during the winter of 1972. Among these specimens

there were some females each with a small white pseudococcid clasped between

the jaws (Fig 15B), which seems to substantiate Biinzli’s theory. Various

specimens of this pseudococcid were collected (Fig 16A—D), including a single

male with the long aedeagus at the tip of its abdomen. This mealy bug

represented a new species and was described by De Lotto (1977) as Xenococcus

scorpioides; he remarked that it is the first representative discovered in the

southern hemisphere of this highly specialized group.

It is apparently common practice among the species of this genus to carry

their ‘cows’ in their jaws, whereas ants of the genus Hypoclinea Mayr (Doli-

choderinae), which is not represented in southern Africa, carrying them on

their backs (Wilson 1971).

According to Emery (1925), Acropyga is divided into four subgenera, viz.

Acropyga s.s., Rhizomyrma Forel (1893), Atopodon Forel (1912), and Malaco-

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

ANOPLOLEPIS AND ACROPYGA 237)

myrma Emery (1922). Only the latter occurs in the Subsaharan region and is

characterized by the long apical segment of the antennae, which is about as

long as the four preceding segments combined, as was depicted in A. (Rhizo-

myrma) palearctica by Menozzi (1936). However, in the latter species the

mandibles are elongate and exhibit a fair-sized space between them and the

clypeus when they are closed, which is not the case in Malacomyrma.

The workers of this subgenus are yellowish in colour, monomorphic, with

very small eyes, consisting of about six to twelve facets. Ocelli are absent. Head

quadrate, sides almost parallel or only slightly convex, hind margin straight or

somewhat concave in the middle, frontal carinae fairly wide apart, and short,

antennal fossae situated close to the posterior border of the clypeus. Mandibles

almost triangular, masticatory margin with six to seven small teeth, the apical

one being slightly larger than the rest; when closed there is only a small space

between them and the clypeus. Antennae eleven-segmented, flagella incrassate

towards their apices. Labrum bilobed angularly and deeply emarginate in the

middle in the specimens examined (Fig. 13C). Gotwald (1969) depicts the

labrum of the Brazilian Acropyga as weakly emarginate. Labial palpi three-

segmented. Clypeus strongly convex in the middle, but not carinate.

Truncus fairly robust, the promeso- and meta-epinotal sutures well deve-

loped, the mesometanotal suture in some specimens distinct in others almost

obsolete. Dorsum of epinotum short, merging into the declivity in a wide angle,

unarmed. Petiole squamiform, the scale inclined forwards. Legs moderately

long.

The median seta on the anterior border of the clypeus is characteristic of

this genus. It occupies a lower position than the other clypeal setae and reminds

one of a trigger hair for setting off the snap action of the mandibles when in the

act of catching prey, or it may even act as a tactile organ, perhaps for detecting

their honeydew producing agents.

Only one species, A. arnoldi, in which the workers have a conspicuous

metanotum, four-segmented maxillary palpi and larger eyes (0,04—0,05 mm in

length, and consisting of about twelve facets) was originally described from the

southern African subregion by Santschi (1926). Two years later he described A.

rhodesiana. In the latter species the workers have a reduced metanotum, and

five-segmented maxillary palpi. A third species, A. silvestrii, was described by

Emery (1915) from Eritrea and it was apparently taken for granted at that time

that the maxillary palpi are one- to two-segmented (Emery 1925; Gotwald

1969). Emery also neglected to mention the number of segments in the

maxillary palpi of silvestrii.

The small collection, made in July 1972 at Jacobs Bay on the west coast

(previously mentioned), contained workers with small eyes (about 0,02 mm

Fig. 11. Male genitalia, cuspides and digiti.

A. Anoplolepis steingroeveri, ventral view. B. Anoplolepis steingroeveri, dorsal view.

C. Anoplolepis custodiens, ventral view. D. Anoplolepis custodiens, dorsal view. E. Ano-

plolepis nuptialis, ventral view. F. Anoplolepis nuptialis, dorsal view.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

long and consisting of about six facets), a distinct metanotum, and five-

segmented maxillary palpi. In this case the males and alate females have

four-segmented and five-segmented palpi respectively. A second collection

made about a year later at Gans Bay on the south coast produced workers very

similar to those found at Jacobs Bay, also having small eyes and five-segmented

palpi, but the mesometanotal suture is almost obsolete. During the winter of

1974, however, collections made at Paternoster, about 12 km north of Jacobs

Bay, revealed workers of both types (some with small eyes, obsolete mesome-

tanotal suture and four-segmented palpi and some with larger eyes, an almost

distinct mesometanotal suture, and five-segmented palpi).

It seems reasonable under these circumstances to assume that most (if not

all) Subsaharan forms have four to five segmented maxillary palpi (the author

has unfortunately not seen A. silvestrii). In specimens with four-segmented

palpi, the apical segment is about as long as the two preceding ones; in the

five-segmented forms, the apical segment is shorter, only about as long as the

preceding one. Unfortunately no males of either the Zimbabwe or Paternoster

forms are as yet known, but on account of the above evidence it is apparent

that A. arnoldi is a variable species and A. rhodesiana should, therefore, be

considered a synonym.

Acropyga arnoldi Santschi, 1926

Acropyga arnoldi Santschi, 1926: 245

Acropyga rhodesiana Santschi, 1928: 211 syn. nov.

Worker (Fig. 12A—C, 15A)

TL 2,40 mm; LI 1,32 mm; HL 0,60-0,62 mm; ED 0,42-0,44 mm; CL

0,14 mm; SL 0,40-046 mm; WL 0,68-0,76 mm; MFL 0,38-0,40 mm; HFL

0,44-0,55 mm; PL 0,10—0,12 mm; CI 90,0—90,3; SI 74,1-82,1; CLI 271,4-300,0

CTT 81,6—88,2; TI 52,9-55,3; PI 150,0-180,0; HFI 64,7-65,8.

Brownish-yellow to yellowish, shiny all over, whole body finely punctured

with piliferous punctures; pubescence moderately long, yellowish and fairly

abundant all over, especially on head and abdomen. Pilosity long, yellowish,

evident only on mandibles, clypeus, apical margins of abdominal segments, and

on scale.

Head almost quadrate about one-ninth longer than wide, sides almost

straight, hind margin very feebly emarginate, or almost straight. Clypeus with

anterior margin arcuate in middle, somewhat sinuate on each side, and with an

angle in between each sinuation and the median area. Frontal area clearly

defined. Eyes small, consisting of about twelve facets, and placed well before

middle of sides of head. Scape about four-sevenths the length of flagellum and

about two-thirds as long as the head, reaching hind margin; second to fourth

flagellar segments about of equal length and shorter than fifth.

Truncus slightly longer than head, pro-mesonotal and meta-epinotal

sutures distinct, mesometanotal suture either distinct or obsolete. Metanotum

ANOPLOLEPIS AND ACROPYGA 239

Fig. 12. Acropyga arnoldi.

A. Dorsal view of specimen described from Zimbabwe. B. Left lateral view of specimen

described from Zimbabwe. C. Left lateral view of truncus of specimen described from

Durban.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Acropyga arnoldi.

A. Dorsal view. B. Left lateral view. C. Head seen from the front, showing the labrum.

ANOPLOLEPIS AND ACROPYGA 241

short, shorter than either meso- or epinotum; latter seen from the side convex

and somewhat higher than promesonotum; epinotal declivity longer than dor-

sum, almost flat, its sides rounded. Scale of petiole wider than long, nearly as

high as wide, rounded dorsally and, seen from behind, wider above than below.

Abdomen nearly as long as the head and truncus together or ohigauly longer,

rounded at base. Legs moderately long.

Workers of this species collected near Gans Bay (Fig. 13A—C) are very

similar to those described above, but the scape of the antennae is somewhat

longer and may extend beyond the hind margin of the head by the width of the

second segment of the flagellum. The truncus in the specimens seen is very

slightly shorter, and in profile the mesonotum is higher than the pro- or

epinotum. The ranges of measurements of twenty specimens are as follows:

TL 2,40-2,96 mm; L 1,26-1,40 mm; HL 0,60-0,64 mm; ED 0,40-0,42 mm;

CL 0,12-0,13 mm; SL 0,46—-0,48 mm; WL 0,60-0,66 mm; MFL 0,38-0,42 mm;

HFL 0,48-0,52 mm; PL 0,11-0,12 mm; CI 87,1-93,3; SI 79,3-85,5; CLI

307,7-333,3; CTI 94,1-103,3; TI 55,9-65,0; PI 150—-163,4; HFI 73,5-83,3.

These ants are also very similar to those collected at Jacobs Bay, but differ

from them by the absence of the mesometanotal suture. They also resemble the

specimens collected at Paternoster, but in the latter case in specimens with

five-segmented maxillary palpi, the eyes are larger and a distinct mesometano-

tal suture is present. The mesonotum is also higher than the pro- or epinotum;

the ranges of the various measurements are:

L 1,30-1,46 mm; HL 0,62-0,68 mm; ED 0,46-0,50 mm; CL 0,15-0,17 mm;

SL 0,48-0,54 mm; WL 0,60-0,70 mm; MFL 0,40-0,46 mm; HFL 0,48-0,58

mm; PL 0,12-0,14 mm; CI 93,8-100,0; SI 78,1-84,4; CLI 270,6-306,7; CTI

97,1-103,3; TI 62,9-70,9; PI 142,9-166,7; HFI 76,2-82,9.

Female (Figs 14A—-B, 15B, E)

TL 3,5-4,0 mm; L 1,80-2,08 mm; HL 0,70-0,76 mm; ED 0,56 mm; CL

0,16 mm; SL 0,5-0,56 mm; WL 1,16-1,22 mm; MFL 0,56 mm; HFL 0,64 mm;

PL 0,12 mm; CI 106,5-108,8; SI 72,7—72,9; CLI 343,8-363; CTI 60,6-62,3; TI

68,9-75,8; PI 200,00; HFI 52,4-55,2.

Brown to piceous-brown in colour, legs paler, antennae yellowish, man-

dibles castaneous. Fairly shiny all over. Finely punctured with piliferous punc-

tures, giving whole integument a superficially and finely reticulate texture.

Pubescence yellowish, moderately long, fairly abundant, especially on

abdomen. Erect pilosity yellowish, sparse, usually present on mandibles,

clypeus, scutellum, scale of petiole and apical borders of abdominal segments.

Head very slightly longer than wide, quadrate, sides very slightly convex,

hind margin almost straight or slightly emarginate. Compound eyes black,

moderately large, occupying nearly one-quarter of the length of the head, and

placed in front of middle of sides. Ocelli large. Frontal area clearly demarcated.

Scapes of eleven-segmented antennae about five-sevenths the length of the

“MOIA [RIUDA ‘a}R]d Je|nUUY “|

‘MOIA [esIop ‘stuod pue Isp ‘sapidsng ‘q ‘“MOIA [esoley YSU ‘eyeyUSs ajepy “D “MIIA [eIO}e] WSU ‘opeulay “q “MOIA JesIOp s[PWIO4{ “V

‘Ipjouan vsddosoy ‘p] “314

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ANNALS OF THE SOUTH AFRICAN MUSEUM

242

ANOPLOLEPIS AND ACROPYGA 243

Fig. 15. Acropyga arnoldi.

A. Worker, left lateral view. B. Female with coccid between jaws, left lateral view.

C. Male, left lateral view. D. Male dorsal view. E. Female, dorsal view.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. Xenococcus scorpioides.

A. Male, right lateral view. B. Male, dorsal view. C. Male, ventral view. D. Female, right

lateral view.

ANOPLOLEPIS AND ACROPYGA 245

head and extending behind hind margin of head by about the width of first

flagellar segment. Apical segment of flagellum about as long as three preceding

ones taken together. Clypeus as in worker.

Truncus about one-quarter or slightly more longer than wide, the sides

somewhat convex, parapsidal furrows clearly defined, scale of petiole almost as

in worker, dorsum slightly emarginate. Legs moderately long. Wings hyaline,

stigma fairly large and slightly darker, venation similar to that of Anoplolepis,

but with large break in transverso-median cross vein (Fig. 15B). Abdomen

about as long as head and truncus taken together, rounded at base.

Male (Figs 14C-E, 15C-D)

TL 2,20 mm; L 1,24 mm; HL 0,43 mm; ED 0,34 mm; CL 0,06 mm; SL

0.48 mm; WL 0,8 mm; MFL 0,56 mm; HFL 0,58 mm; PL 0,08 mm; CI 113,04;

S922 C5333 Ciil-5326) 75.0 PE 187/25: 7255:

Piceous-brown, shiny all over, eyes black. Legs and flagella paler in colour

than body. Pubescence moderately long, adpressed and whitish-yellow. A few

pilose hairs present, particularly on head and posterior margins of abdominal

segments. Sculpture similar to that of female.

Head slightly wider than long, sides and posterior margin convex. Clypeus

rather convex from side to side, also very convex in lateral view, forming a

deep notch between it and junction of frontal carinae; its anterior margin

almost straight. Eyes bulging, large, almost as long as half the length of the

sides of the head. Ocelli large. Scape of twelve-segmented antennae extending

behind hind margin of head by about one-third of its length. Apical segment of

flagellum about as long as three preceding segments taken together. Legs

moderately long. Maxillary palpi four-segmented, apical segment longer than

preceding one and somewhat tapering towards apex and undivided in specimen

seen. Labial palpi three-segmented. Mandibles with five to six small teeth,

apical ones longer.

Truncus longer than wide, somewhat narrower than head, anterior margin

of scutellum indicated by thin black line. Scale of petiole similar to that of

female, thinner above than below, very slightly emarginate in middle of dorsal

edge.

External genital organs

Gonostipes longer than volsellae and penis almost triangular, tapering

towards apices, the latter rounded. Penis lobes broad and also triangular, about

of same length as volsellae; almost spoon-shaped digiti somewhat longer than

short cuspides, which are rounded. Annular plate small, almost trapezoidal,

widely and shallowly emarginate behind, lateral angles acute, each with long

seta; there are also some setae on ventral plate.

Material examined

Holotype ¢, Durban, 29.8.18, C. P. Merwe; another 2 Jacobs Bay,

20.7.72, A. J. Prins; 3, Jacobs Bay 20.7.72, A.J. Prins; syntype 2%, Durban,

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

29.8.18, C. P. Merwe; syntype %9%, Bulawayo, 9.11.24; other 8% Jacobs Bay,

202 Ae J, Prins: ‘Gans Bay 3:32:73, Aj J Enms; Paternoster 9%om/ae

V. B. Whitehead.

ACKNOWLEDGEMENTS

I wish to express my gratitude to Dr V. B. Whitehead of the South African

Museum for his comments. I am also indebted to Dr G. R. McLachlan of the

- South African Museum and Mr R. K. Brooke of the Percy FitzPatrick Institute

of African Ornithology, Cape Town, for checking the lists of birds. I also thank

Mr V. Branco of the South African Museum for the drawings of the female of

Acropyga arnoldi.

REFERENCES

ARNOLD, G. 1920-22. A monograph of the Formicidae of South Africa. Ann. S. Afr. Mus. 14:

403-674.

Brown, L. H. & Newman, K. B. 1974. ‘Anting’ in African passerine birds. Ostrich 45:

194-195.

Bunzu, G. H. 1935. Untersuchungen tiber coccidoplule Ameisen aus den Kaffee felderen von

Surinam. Mitt. schweiz. ent. Ges. 16: 453-593.

CoLaHAN, B. D. 1981. Anting in the Cape white eye. Ostrich 52: 186.

De Lorto, G. 1977. On some African mealy bugs (Homoptera: Coccoidea: Pseudococcidae). J.

ent. Soc. sth. Afr. 40: 13-36.

Emery, G. 1915. Formiche raccolte nell Eritrea dal Prof. F. Silvestri. Boll. Lab. Zool. gen. agr.

Portici 10: 1-26.

Emery, C. 1922. Quelques fourmis nouvelles minuscules. Annis. hist. nat. Mus. natn. hung. 19:

107-112.

Emery, C. 1925. Hymenoptera, Family Formicidae. Subfamily Formicinae. Jn: P. WYTSMAN.

Genera Insectorum 183: 1-302.

ForeL, A. 1893. Formicides del’Antille St Vincent. Récoltés par Mon. H. H. Smith. Trans. ent.

Soc. Lond. 1893: 333-418.

ForeL, A. 1894. Abessinische und andere afrikanische Ameisen, gesammelt von Herrn Inge-

nieur Alfred Ilg. von Herrn Dr Liengme, von Herrn Pfarrer Missionar P. Berthoud, Herrn

Dr Arth. Miiller, etc. Mitt. schweiz. ent. Ges. 9: 64-100.

ForeL, A. 1912. Descriptions provisiones de genres, sous-genres et espéces de Formicides des

Indes Orientales. Revue suisse Zool. 20: 761-774.

Fore, A. 1913. Fourmis de Rhodesia etc. récoltées par M. G. Arnold, le Dr H. Brauns et K.

Fikendy. Ann. soc. ent. Belg. 57: 108-147.

GotwaLp, W. H. 1969. Comparative morphological studies of the ants, with particular

reference to the mouthparts (Hymenoptera: Formicidae). Mem. Cornell Univ. agric. Exp.

Sin. 408: 1-150.

Kiorr, W. 1959. Versuch einer Analyse der trophobiotischen Beziehungen von Ameisen zu

Aphiden. Biol. Zbl. 78: 863-870.

Louw, J. F. 1968. Die ekologie van die malmier Anoplolepis custodiens (Sm.) (Formicidae) in

die sentrale Oranje Vrystaat. Die Bogrondse aktiwiteitspeil van die werkers. J. ent. Soc.

sth. Afr. 31: 241-248.

MeENozzi, C. 1936. Nouvi contributi alla conoscenza della Fauna delle isole italiane dell’egeo.

VI. Hymenoptera—Formicidae. Boll. Lab. Zool. gen. agr. Portici 29: 262-311.

Myers, N. J. 1957. Studies on the biology of ants associated with citrus trees. Unpublished

M.Sc. thesis, Rhodes University, Grahamstown.

Prins, A. J. 1973. African Formicidae (Hymenoptera) in the South African Museum. Descrip-

tion of four new species and notes on Tetramorium Mayr. Ann. S. Afr. Mus. 62: 1-40.

ANOPLOLEPIS AND ACROPYGA 247

Prins, A. J., CILuie, J. J., HONIBALL, F. & DE ViLuisErRs, J. F. 1979. In: BEDForD, E. C. G.

Citrus pests in the Republic of South Africa. Sci. Bull. Dept. agric. tech. Serv. Rep. S. Afr.

391: 203-215.

SANTSCHI, F. 1916. Fourmis nouvelles de la colonie du Cap, du Natal et de Rhodesia. Ann. soc.

ent. Fr. 85: 279-296.

SANTSCHI, F. 1926. Descriptions de nouveaux Formicides Ethiopiens. Revue zool. Afr. 13:

207-267.

SANTSCHI, F. 1928. Descriptions de nouvelles Formis Ethiopiennes. Revue Zool. Bot. Afr. 16:

191-213.

Smit, B. 1964. Insects in southern Africa—how to control them. Cape Town, Salisbury,

Johannesburg: Oxford University Press.

SmitTH, F. 1858. Hymenoptera. Cat. Hym. Brit. Mus. 6: 1-216.

Steyn, J. J. 1954. The pugnacious ant (Anoplolepis custodiens Smith) and its relation to the

control of citrus scales at Letaba. Mem. ent. Soc. Sth. Afr. 3: 1-96.

WHEELER, W. M. 1960. Ants. Their structure, development and behavior. New York: Columbia

University Press.

Wate, I. J. 1981. Anting in Blue: eared glossy Starlings. Ostrich 52: 185.

Wiison, E. O. 1971. The insect societies. Cambridge, Massachusetts: Belknap Press.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

noy., syn. nov., etc. ;

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: SO.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text _

e.g. ‘... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded

by initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a

book or article; such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Pea

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

to initial capital letter, provided the same generic name is used consecutively. ;

Name of new genus or species is not to be included in the title: it should be included in the

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.