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Full text of "African invertebrates : a journal of biodiversity research"

ANNALS 



OF THE 



NATAL AIUSEITM 



KDITED BY 



ERNEST WARREN, D.S(;.(Lond.), Dieeotor. 




PRINTED BY ORDER OF THE TRUSTEES. 



LONDON: 
ADLARD & SON & WEST NEWMAN, LTD, 

BARTHOLOMEW CLOSE 
1919 



CONTENTS. 



VOLUME III. 

Part 1, issued September, 1914. page 

Descriptions of some South African Marine Shells. By 

Edgar A. Smith, I.S.O. (With Plate I) . .1 

A Case of Hybridism amonj.,^ Cockatoos. By Ernest 

Warren, D.Sc.(Lond.). (With Plate II) . .7 

On South African Ennese, with Descriptions of New Species 
and Varieties. By Henry Clifden Burnup. (With 
Plates III, IV, V, and Text-figs.) . . .29 

On the Development of the Planula in a Certain Species 
of Plumulariau Hydroid. By Ernest Warren, D.Sc. 
(Lond.). (With Plate VI and 4 Text-figs.) . . 83 

Note on the Occurrence in South Africa of a Termitophilous 
Beetle of the G-enus Cor o toe a. By Ernest Warren, 
D.Sc. (Lond.) ...... 103 

Part 2, issued October, 1915. 

Studies on the Carnivorous Slugs of South Africa, including 
a Monograph on the Grenus Apera, and a Discussion 
on the Phylogeny of the Aperidse, T estacellidse, 
and other Agnathous Pulmonata. By Hugh Watson, 
M.A. (With Plates VII-XXIV, and 7 Text-figures) . 1()7 

The Parthenogenetic Tendency in the Moth, Melanocera 
menippe (Westwood). By Ernest Warren, D.Sc. 
(Lond.) . . . . . . .269 

New South African Arachnida. By John Hewitt, Director, 

Albany Museum, G-rahamstown. (With 9 Text-figures) 289 

Oliservations on some South African Termites. By Claude 
Fuller, Division of Entomology, Department of Agri- 
culture, Pretoria. (With Plates XXV-XXXV and 16 
Text-figures) . . . . . .329 



IH'^Z 



IV CONTENTS. 



PAGE 



A Further Note on Hybrid Cockatoos. By Ernest Warren, 

D.Sc.(Lond.) ...... .50.5 

Part 3, issued May, 1917. 

The Plant Ecology of the Drakensberg Range. By J. W. 
Bews, M.A., D.Sc, Professor of Botany, Natal University 
College. (With Plates XXXVI-XXXIX and 3 Text- 
figures) . . . . .. . .oil 

Contributions to a Knowledge of the Terrestrial Isopoda of 
Natal. Part I. By Walter E. Collinge, D.Sc, F.L.S., 
etc., Research Fellow of the University of St. Andi-ews. 
(With Plates XL-XLII) . . . . .567 

South African Bagworms. Editorial Note . . 587 

South African Bagworms ; Notes on the Psychidse and on 
the genera Gymnelema and Trichocossus, with 
descriptions of five new species. By A. J. T. Janse, 
F.E.S.L. (With Plate XLIII) . " . . .589 

Descriptions of Some New South African Microlepidopterous 

Bagworms. By E. Meyrick, B.A., F.R.S. . H15 

South African Bagworms : their Transformations, Life- 
history, and Economic Importance. Part I. By C. B. 
Hardenberg, M.A., Grovernment Entomologist in 
Charge of Wattle Insect Investigations. (With 
Plates XLIV-XLVI and 6 Text-figures) . . 619 

Descriptions of New South African Arachnida. By John 
Hewitt, Director, Albany Museum, Grahamstown. 
(With Plate XLVII and 4 Text-figures) . . 687 

Issued April, 1919. 

Title-page of Vol. Ill . . . . i 

Contents of Vol. Ill . . . . iii 

Index of South African Mollusca .... 713 

General Index ....... 719 



Vol. Ill, part 1, issued September 2dth, 1914. 

SOME SOUTH AFRECAN MARINE SHELLS. ] 



Descriptions of some South African Marine 

Shells. 



By 



Ed^ai- A. <^mitli, I.S.O. 



With Plate I. 



In a collection of shells, mostly from Natal, kindly sub- 
mitted to me for examination by Mr. H. C. Burnup, five 
species appear to be undescribed, and therefore I have drawn 
up the following descriptions of them, and at the same time 
made some remarks on two known forms of the genus 
Tivela. 




Mangilia shepstonensis n. sp. PL I, fig. 1. 

Testa parva, turrita, cerea, circa medium anfractus ultimi 
linea rufa interrupta cincta ; anfractus 6 ; superiores Iseves 
(? detriti), convexi, casteri planiusculi, superne rotunde 
tabulati, costis circiter duodecim leviter obliquis instructi, liris 
spiralibus (in anfr. penult, tribus, in ultimo ad tredecim) 
supra costas subnodulosis cincti, inter liras striis spiralibus 
et lineis increment! minute decussati ; apertura angusta, 
spira paulo brevior; labrum extra valde incrassatum, macula 
rufa ornatum, prope sinum conspicuura superiorem tuberculo 
parvo munitum ; columella rectiuscula, callo tenui induta. 

Longit. 4^, diam. 2 mm. 

Hab. — Port Shepstone (Burnup). 

The apical whorls are large for so small a shell. Two 
specimens have been examined, and one of them has the spire 

VOL. 3, PART 1. 1 



2 EDGAR A. SMITH. 

rather more produced than the other. The fine decussation 
of the surface between the costae and the spiral lirte is only 
visible under a powerful lens or microscope. 

Cerithiopsis (Sella) natalensis n. sp. PI. I, fig. 2. 

Testa subulata, nigro-fusca, ad apicem pallida; anfractus 
circiter decern, convexi, regulariter acerescentes, sutura 
obliqua sejuncti, liris spiralibus ad quinque instructi, liris 
incrementi fortibus oblique arcuatis sculpti, inter liras 
spiraliter tenuiter striati ; ultinius infra concentrice striatus ; 
apertura irregulariter rotundata, intus nigrescens ; columella 
arcuata, callo tenui reflexo induta, ad basim oblique 
truncata. 

Longit. 10 mm., diam. 2"5. Apertura 1"75 longa, 1'25 lata. 

Hab. — Tongaat, Natal (Burnup). 

Very like Sella attenuata Hedley from Port Jackson, 
but with stronger lines of growth and more equal spiral 
cords, and intermediate thread-like lines. On crossing the 
spirals these become almost granulated. The nuclear whorls 
are wanting in the specimen described. 

Grlyphis levicostata n. sp., PI. I, figs. 3-.5. 

Testa oblongo-ovata, autice leviter angustata, mediocriter 
elata, alba, fusco-olivaceo radiata, circa foramen interdum 
pallide livida costulis numerosissimis vix elatis et liris in- 
crementi tenuibus confertis undulatis sculpta ; foramen 
magnum, oblongum, intus callo livido postice truncato 
succinctum ; superficies interna albo-pellucida lineis opacis 
albis radiata, ad marginem lateraliter paulo elatum tenuiter 
crenulata. 

Longit. 35, diam. 23, alt. 14 mm. 

Hab. — Tongaat and Port Shepstone, Natal (Burnup). 

This species is remarkable for the fineness of its sculpture, 
and the elongate cap-shaped form. The radiating riblets are 
very little raised, and separated merely by impressed lines 



SOME SOUTH AFRICAN MARINE SHELLS. 3 

or strias. The concentric lines of growth are crowded 
and wavy, and in the grooves between the riblets there is 
in places a somewhat punctate appearance. Four or five 
concentric waves or ridges at fairly equal distances fi-om 
the foramen may indicate periods of arrested growth. The 
subapical hole is very anterior, oblique in front of the apex, 
and the front slope beneath it is considerably concave- 
Considering the lightness of the external costulation, the 
margin is distinctly denticulate, the grooves between the 
denticles extending within a little distance from the edge. 
The denticles are about eighty in number, and those upon 
the anterior end are often sub-bipartite. G. tenuistriata 
(Soioerhy) is rather like the present species as regards foi-m, 
but the sculpture is not the same. In the largest specimen 
examined the colour rays are very faint, probably through 
fading. A smaller and fresher specimen exhibits seven rays 
— three on each side and one in front ; the two behind slope 
posteriorly, four are inclined towards the front, the seventh 
ray extending from near the foramen to the margin down the 
middle of the shell. 

Tivela compressa Snverhy. PI. I, figs. 6-8. 

Hab. — Port Shepstone, Tongaat, Scottburgh (Burnup) ; 
Isipingo (Miss Lois Trotter) ; Isezela (Miss Chapman). 

T. compressa was originally described from the Cape of 
Good Hope, and it also occurs at Muizenberg, in False Bay, 
and at Port Alfred. The specimens from Port Shepstone, etc., 
differ from the typical form in having the ventral margin less 
roundly curved, so that the valves have a more triangular 
appearance, and the height of the shell is less in proportion to 
the length. The pallial sinus is more acuminate and generally 
extends beyond the middle of the valves, whereas in the normal 
form it hardly reaches half-way across. The reddish-brown 
posterior dorsal area appears to be a constant feature in this 
species. It sometimes attains a length of 60 mm. 



EDGAR A. SMITH. 



Tivela dunkeri {Rome)'). PI. I, figs. 9, 10. 

Venus (Cytherea) (Tivela) dunkeri R'umer, Novitat. Conch. 
Monog. Yenus, p. 5, pi. v, fig. 1. 

Hab. — Port Sliepstone (H. Burnup). 

The locality of this species has not been hitherto recorded. 
It is very donaciform in shape^ the anterior end being longer 
than the posterior. It is described by Ronier as pale violet- 
white, tinged with violet dorsally. His specimen was 25 mm. 
in length, whilst the valves under examination, evidently 
young shells, are only 15. 

The other S. African Tivelas are: T. natalensis Dkr. 
1864 (=alucinans Sow. 1897), T. transversa Sow. 1897, 
and T. polita Sow. 1851 (=dolabella Sow. 1851). 

Tivela rejecta n. sp. PI. I, figs. 11-13. 

Testa trigona, fere asquilateralis, sordida lactea rufo macu- 
lata, distanter livido zonata, lineis incrementi striata; mar- 
gines dorsales utrinque recte declives ; basalis late curvatus ; 
latus anticum acute rotundatum, posticmn magis acuminatum ; 
umbones acuti, purpureo tincti ; lunula vix definita, f usco- 
purpurea tincta ; area postica f usco-purpurea ; pagina interna 
fusco-purpurea, ad marginem basalem albida ; pallii sinus 
angustus, trans medium procurrens. 

Longit. 15'5, alt. 11*5, crass, fere 7 mm. 

Hab. — South Africa. 

The shell here described may not be adult, but it differs 
from the young of all the other South African Tivelas. T. 
polita most nearly approaches it, but that species is higher 
in proportion to its length, flatter, and the pallial sinus is 
shorter and wider. The dorsal margins within the valves in 
T. rejecta are of a deep purplish-brown with pale rays 
beneath them, one on each side, which join the white ventral 
margin. The reddish-brown markings on the outer surface 
are irregular, but disposed more or less in transverse bands. 



SOME SOUTH AFJUCAN MARINE SHELLS. O 

There are also one or two livid zones at distant intervals. 
There are slight differences in the hinge-teeth, which it would 
be difficult to describe,but which can be observed by comparing 
the two forms. 

T. dunkeri, Rnmer, differs in form, the posterior end being 
rounded and not acuminate as in the present species. 

Loripes burnupi n. sp. PI. I, figs. 14-16. 

Testa irregLilariter ovata, mediocriter convexa, subtequi- 
lateralis, tenuis, albo-pellucida, periostraco tenui lutescente 
partim obtecta, lineis incrementi tenuissime striata ; lunula 
parva, pi-ofunda; umbones supra lunulam incurvati ; dens 
cardinalis unicus valvas dextrte crassus, solidus, irregularis, 
infra umbonem situs ; dentes duo valvae sinistree, quorum 
posticus augustus, retrorsum incliiiatus, anterior solidus, 
irregularis, a postico fossa lata sejunctus ; ligamentum 
elongatum, fere internum; pagina interna obsolete radiatim 
tenuiter striata, in fundo cicatricem angustam obliquam 
exhibens ; cicatrix antica, mediocriter elongata, postica 
elongato-piriformis. 

J^ong. 19"75, alt. 20, crass. 10 nnn. 

Hab. — Port Shepstone (Burnup). 

Although the ligament is not so completely internal in this 
shell as is the case in typical forms of Loripes, I do not con- 
sider this a sufficient difference upon which to found a distinct 
genus since the position of the ligament is subject to con- 
siderable variation. 

A much larger shell from the Cape Yerd Islands, described 
by 8owerby ^ as C r y p t o d o n m u r c h 1 a n d i, externally closely 
resembles the present species. The hinge-teeth, hoAvever, are 
absent, or may have become obliterated with age. The form 
of the anterior scar also is somewhat different. 

' Sowerby, G. B.. ' Proc. Maine. Soc.,' vol. vii. p. 303, pi. xxv, fig. 13. 







EDGAR A. SMITH. 



EXPLANATION OF PLATE I, 

Illustrating Mr. Edgar Smith's paper, "Descriptions of 
some South African Marine Shells." 

Fig. 1. — Mangilia sliepstonensis n. si^. 

Fig. 2. — Cerithiopsis (Seila) natalensis n. sj). 

Figs. 3-5. — Glyphis levicostata 7*. sj). 

Figs. 6-8. — Tivela compressa Sotverby. 

Figs. 9. 10.— Tivela dunkeri Rvmer. 

Figs. 11-13.— Tivela rejecta n. sp. 

Figs. 14-16. — Loripes bvirnupi n. sp. 



Ann. Natal Mus. Vol. HI. 



PI. I. 




SOUTH AFPJCAN MOLLUSCA 



J. GresTi. del et hth 



HYBRIDISM AMONG COCKATOOS. 



A Case of Hybridism among Cockatoos. 

By 

Eriieii>t Warren, D.Sc.Loiid. 



With Plate II. 



There has recently occurred an interesting example of 
hybridism among certain tame cockatoos in Pietermaritzburg, 
and since the character of hybrids gives some insight into the 
problems of hereditary transmission it appeared to be desir- 
able to publish an account of the matter, and the living birds 
have been studied as carefully as circumstances would permit. 

The cockatoos belong to Lieut. -Colonel A. T. Wales, resident 
in Pietermaritzburg, who has very kindly allowed me to sketch 
the birds and have them photographed ; my thanks are also 
due to his daughter, Mrs. Brown, who, having considerable 
control over the somewhat wild birds, rendered great assist- 
ance while they were being photographed. My friend, Dr. 
Conrad Akerman, was good enough to undertake the photo- 
graphing, and from the prints of the negatives thus obtained 
the lithographic plate was prepared. The artist who coloured 
the plate was provided with a selection of the variously 
coloured feathers. 

The male-bird is the common sulphur-crested cockatoo, 
Cacatua galerita [Lath.), and the female is the so- 
called slender-billed cockatoo, Licmetis nasica {Temm.). 
The birds were obtained on different occasions at the coast- 
town of Durban, and were doubtless at the time of purchase 
recently imported from Australia. 

The birds live in a garden under fairly natural conditions ; 



ERNEST WARREN. 

they are mostly confined in a capacious wired-in enclosure 
protected from the rain, but quite frequently they escape 
into the garden and climb among the trees. At the present 
time there are five birds in the aviary ; namel}^, tAvo hybrids, 
the parents of the same, and a second specimen of sulphur- 
crested cockatoo. By wire-partitions and wire-cages the 
birds can be separated from one another. This is particularly 
necessary in the case of the elder of the hybrids, which at 
times exhibits a violent disposition. The bird has already 
killed a grey parrot, and has very severely mauled the second 
specimen of cockatoo, which, nevertheless, is a considerably 
larger and more powerful-looking bird. 

The female-bird was purchased some twelve years ago, and 
the male-bird about four years afterwards. Thus the birds 
have been associated together for a period of about eight 
years. During the first two years the birds were not confined 
at all ; they lived among the trees in a garden in a different 
portion of the town. Apparently no pairing occvirred during 
this period, and no eggs were laid. Subsequently the owner 
moved to another residence and the birds were lodged in 
their present quarters. 

It is difficult to say whether the closer association of the 
two birds arising from the confinement favoured the pairing, 
but for the following six years (1908-1913) eggs were laid 
regularly every August. It is possible, however, that the 
production of eggs by the female cockatoo does not necessarily 
mean that copulation invariably took place; for in this connec- 
tion it may be mentioned that a tame owl (Syrnium wood- 
f ordi) in my possession occasionally produces perfectly formed 
eggs, although the bird is unpaired ; also, a certain unpaired, 
tame and unidentified parrot belonging to the Misses Black- 
more of this town has laid eight or nine eggs annually for 
several years. In the case of the birds with which we are 
now concerned, actual pairing has been observed and probably 
it occurred every season. 

The eggs are laid about the middle of August. In 1908 
two «ggs Avere produced, and the owner, under the impression 



HYBRIDISM AMONG COCKATOOS. 9 

that the cockatoos would not give proper attention to incuba- 
tion, placed theui under a hen, but the egg-shell proved to be 
too delicate, and the eggs were soon broken. In 1909 another 
two eggs were laid, and they Avere placed under a pigeon ; 
but the pigeons were not disposed to take any notice of them, 
aud they were almost immediately smashed. In 1910 two 
en-g-s were likewise laid in the middle of August, and the 
cockatoos Avere allowed to take charge of them. The male- 
and female-birds took turns in incubation, and in due time 
one egg hatched, while the other was addled. Colonel AVales 
informs me that he did not take special note of the period of 
incubation, but he believes that it was about twenty -one days. 
The egg which did not hatch was thrown away without being- 
examined. In 1911 the female-bird dropped two eggs from 
her perch and they were smashed. In 1912 two eggs were 
laid and sat upon by the birds, but they were addled. In 
1913 three eggs were laid, and after incubation by the 
cockatoos one egg hatched while the other two were addled. 
Thus in three separate years, 1910, 1912, 1913, out of 
seven eggs laid in all, only two hatched, and the hybrids 
thus arising form the subject of the present observations. It 
cannot be said for certain whether the eggs failed to hatch 
through non-fertilisation, or through the death of the embryos 
owing to constitutional weakness. It is clear that the pro- 
portion of eggs which hatched (two out of seven) was low, 
and it Avould appear probable that this was due to the fact 
that the male- and female-birds were of different species. 

The male-bird (PI. II, A), was a fine specimen of C. 
galerita; and it may be useful to quote here the description 
of the species given in the British Museum Catalogue.^ 

"Adult. — White, crest sulphur-yellow; base of the inner 
web of quills and tail-feathers pale sulphurryellow, like the 
base of the feathers of the head and neck ; ear-coverts some- 
times tinged with sulphur-yellow ; naked skin round the eyes 
white; iris dark-brown; bill black; feet blackish. Total 

' ' Catalogue of the Bii-ds in the British Museum,' vol. xx, p. 117. 



10 ERNEST WARREN. 

lengtli 20-18 inclies, wing 15-13-5, tail 8-5-7-5, bill 2-1-7, 
tarsus 1 '15-1-1. 

''^Hab, — Australia and Tasmania." 

Count Salvadori further remarks : " I have not been able 
to examine specimens from Tasmania, which, on account of 
some little difference in the shape of the bill, have been 
specifically separated as C. licmetorhyncha {Bp.)." 

The present specimen when standing on perch measured 
17-5 inches (445 mm.) from the crown of the head to the tip 
of the tail. 

Sulphur-yellow crest well-developed, and with recurved 
extremity ; the longest feather measured 45 inches (1 14 mm.). 

Prominent projecting forehead. 

The ear-coverts were tinged with sulphui'-yellow. 

Exposed portion of eye, when eye-lids were open, was 
somewhat oval in shape, set horizontally, and measured 
31 sq. mm. Eye-lids, when wide open, exhibited at their 
margin a circle of well-developed, symmetrical, squarish 
granulations. Iris veiy dark brown. 

Naked skin round the eye oval in shape, set horizontally, 
and measured 144 sq. mm. It was irregularly furrowed, and 
both it and the eyelids were of a chalky white. 

The bases of many of the feathers of the head and throat 
were sulphur-yellow in colour. 

Length of tail 8 inches (203 mm.) ; base of inner web of 
tail-feathers resembled that of the quill-feathers in being pale 
sulphur-yellow. 

Bill, black horn-colour, Avitli the appearance of being dusted 
with white powder. The upper mandible from forehead to 
tip of hook measured 1-68 inches (42 mm.) ; depth from fore- 
head to lower inner angle, 1-05 inches (27 mm.). The lower 
edge of the upper mandible, when bill was in a normal 
position of rest, was only about 8° from the horizontal, 
and was nearly parallel to a line passing through the centre 
of the eye and the middle point of the junction of the fore- 
head and bill. There is considerable variation in this matter 
in different individuals, as it has been observed that in some 



HYBRIDISM AMONG COCKATOOS. 



11 



specimens the lower edge slopes upwards towards the mner 
corner at an angle of 10°-15°. The hook of the bill was 
short and perpendicular. There is also much individual 
variation in the length and curvature of the hook ; and from 
Count Salvadori's remarks, quoted above, it would appear 
that in Tasmanian specimens the hook tends to be longer and 
more nearly like that of the genus Licmetis. 

The upper edge of the lower mandible is slightly concave 
for its proximal two-thirds, while towards the tip it is deeply 
concave owing to the curvature of the bill. 

Nostrils surrounded by narrow naked cere at the insertion 
of the bill. 

The female-bird (PI. II, B) was a normal specimen of 
Licmetis n a sic a, and the description of the species given 
in the British Museum Catalogue ^ is as follows : 

"Adult. — White; lores and a narrow frontal band red; 
the feathers of the head, neck and breast are scarlet at the 
base, showing through the white, especially on the breast : 
the under surface of the wing washed with pale brimstone- 
yelloAv, the under surface of the tail also washed with brim- 
stone-yellow, but brighter: iris light brown; bill white; naked 
skin round the eyes light blue ; feet dull olive-grey. Total 
length 1.5 inches; wing 11 ; tail 5 ; bill T95 ; tarsus 0-9.5. 

"Hab. — Australia, from the Gulf of Carpentaria, through 
the interior to N. S. Wales and South Australia." 

The present specimen when standing on perch measured 
13i inches (343 mm.) from the crown of the head to the tip of 
the tail. 

The crest is in the form of a ruif consisting of the front 
white feathers of the crown, which are ei-ected Avhen the bird 
is annoyed. The feathers which correspond to the long- 
recurved sulphur-yellow feathers of the crest of C. galerita 
are further back on the crown, and are short feathers about 
1-5 inches (38 mm.) in length. These have red bases Avhich 
are not ordinarily visible when the bird is quiescent. 

1 Ibid., p. 134. 



12 ERNEST WAEEEN. 

Forehead liigli, but with a receding aspect, and not promi- 
nent as in C. galerita. 

Ear-coverts pale bi'ownish-yellow or bufT. 

Exposed portion of eye, when eye-lids were wide open, was 
circular in shape and measured 25 sq. mm. The edges of the 
eye-lids did not exhibit granulations. Iris, very dark-brown, 
not light-brown as stated in the description quoted above. 

Naked skin around the eye was irregular in shape and 
extended much further below the eye than above it. The 
-area was large, measuring 343 sq. mm., and grooved and 
furrowed ; both it and the eye-lids were of a whitish-blue 
colour. 

Lores red ; a narrow red frontal band was continued from 
the lores above the insertion of the upper mandible, and a 
rather paler red coloration extended backwards both above 
and below the naked area around the eye. 

The bases of most of the features of the head, throat, neck 
and upper chest were bright red in colour. 

Length of tail 4*75 inches (120 mm.). Under surface of 
tail and wing sulphur-yellow. 

Bill whitish, the colour of boiled bone, proximally a slightly 
bluish tinge and distally a yellowish tinge. The upper 
mandible from foreliead to tip of hook measured 1"8 inches (47 
mm.) ; depth from forehead to lower inner angle 0'75 inches 
(19 mm.). The lower edge of upper mandible, when bill was 
in a normal position of rest, sloped upwards towards the inner 
corner at an angle of about 27° with the horizontal. The 
hook was long and conspicuously curved, and the proximal 
portion of the hook projected outwards and not perpendicu- 
larly as in C. galerita. The upper edge of the lower 
mandible was nearly straight proximally for about one-third 
of its length, while the distal two-thirds was very deeply 
concave. 

Cere feathered, and nostrils situated further in from the 
insertion of the bill than in C. galerita. 

The three years old hybrid (PI. II, C), Avhen standing on 



HYBRIDIS-M AMONG COCKATOOS. 13 

perch, measured 15 inches (381 mm.) from the crown of the 
head to the tip of the tail. 

A crest was present consisting of short reddish -yellow or 
orange feathers obviously representing the recurved sulphur- 
yellow feathers of the crest of C. galerita. These feathers 
were only 2'8 inclies (73 mm.) in length, and they were quite 
straight and exhibited no tendency towards the recurved 
condition seen in Cacatua. Thus, in the condition of the 
crest, the hybrid stands between the male- and female-parent, 
but somewhat nearer to the male. The short feathers of the 
female corresponding to the crest-feathers of the hybrid are 
red at their bases, and the colour of the crest of the hybrid is 
clearly a mixture of yellow and red. 

The forehead of the hybrid Avas more prominent than in 
the female- and less so than in the male-parent. 

The ear-coverts were pale yellowish-buff and did not differ 
appreciably in colour from those of Lie metis nasica. 

The eye was large, and when the eyelids were wide open 
the exposed area was 34 sq. mm., thus equalling, or even 
slightly exceeding, the size of the eye in the male-bird. The 
edge of the eyelids when open exhibited granulations, but 
they were smaller and less regular than in C. galerita. Iris 
very dark brown. 

The naked skin around the eye was oval in outline, and 
was more or less intermediate in size (208 sq. mm.) between 
that of the male and that of the female; but in this character 
also the hybrid stands somewhat closer to the male- than to 
the female-parent. The colour of this skin was of the palest 
bluish-white, which was almost exactly intermediate between 
the chalky white of the male and the pale blue of the female. 

The red colour of the lores of the female was converted 
into a yellowish orange in the hybrid ; while the red colour of 
the bases of the feathers of the head and neck was replaced by 
a beautiful gold colour. 

Length of tail 7 inches (178 mm.). Under surface of tail 
and wing sulphur-yellow. 

Bill dark horn-colour, but not quite so black as in C . 



14 ERNEST WAUREN. 

galerita; the difference is more obvious when the bill is 
wetted. The upper mandible, from forehead to tip, measured 
1'7 inches (43 mm.); depth from forehead to lower inner angle 
0'93 inch (24 mm.). The lower edge of upper mandible, when 
bill was in a normal position of rest, sloped upwards towards 
the inner corner at an angle of about 19° to the horizontal. 
The hook was larger than in the male-bird and was more or 
less intermediate in shape between that of the male and 
female. The upper edge of the lower mandible was, on the 
whole, somewhat more concave than in the male-, but less so 
than in the female-bird. 

Cere partially feathered so that the nostrils, which were 
situated just inside the junction of the bill and head, had the 
back edge feathered and the front edge naked, a condition 
strikingly intermediate between that seen in the male- and in 
the female-bird. 

The three months old hybrid (PI. II, D) was not sufficiently 
mature to render it advisable to compare its measurements 
with those of the parents. In all general characters it very 
closely resembled the mature hybrid. 

The orange crest was already rather longer than in the 
older hybrid, but none of the feathers were recurved. 

The area of the lores coloured yellowish-orange Avas rela- 
tively less than in the three years old hybrid, and considerably 
smaller than the red area in L. nasica. Naked skin around 
the eye very pale bluish-white. Iris very dark brown. 

Bill very dark, but not quite so black as in C. galerita. 

The shape of the upper mandible and hook of same Avas 
somewhat nearer to that seen in the male than Avas the case 
in the older hybrid. Also, the curvature of the upper edge of 
the loAver mandible was somewhat closer to that of the male. 

From a Mendelian point of view animal hybrids derived 
from the crossing of species are as a rule singularly unsatis- 
factory. They usually exhibit an intimate blending in the 
majority of the characters of the two parent species, and 



HYBRIDISM AMONG COCKATOOS. 15 

they are mostly sterile. The present case is no exception; 
the characters of C. galerita and L. nasica have very 
obviously blended, and the resulting hybrid stands between 
the two species, but somewhat nearer to Cacatua than to 
L i c m e t i s . 

A simple Mendelian relation would have been shown if, for 
example, the sulphur-yellow of the Cacatua had been 
dominant and the red of Licmetis recessive, and on subse- 
quent crossing of the hybrid (assuming that it was possible) 
if 75 per cent, of the offspring had been entirely yellow and 
25 per cent, entirely red. But in the present case of hybridism 
we do not find any definite evidence of the existence of such 
dominance in any of the characters examined. It is true that 
the colour of the hybrid is nearer to yellow than to red, and 
that in this character there is prepotency on the part of the 
male-bird; but the yellow colour of the hybrid is not the 
sulphur-yellow of Cacatua; the latter colour has become 
blended with red to form an orange crest, orange lores, and a 
beautiful gold under-plumage to the head and neck. 

Again, it would not serve any useful purpose to regard the 
possession of a crest as a dominant character. The short, 
straight, orange crest of the hybrid is obviously a kind of 
mean between the long recurved sulphur-yellow crest of 
Cacatua and the short, white and red feathers of the crown 
of Licmetis. 

A similar blending of characters is seen in the position of 
the nostrils, the condition of the cere with reference to the 
feathering, the colour and shape of the upper and lower 
mandibles, and the shape of the forehead. 

The characters which do not readily admit of numerical 
expression are grouped together in the accompanying table 
(p. 16)._ 

It will be noticed that under " Hybrid " more of the 
characters are found in the column "nearer Cacatua" than 
elsewhere, but it must be remembered that in every case the 
influence of Licmetis is very distinctly obvious, and, except 
perhaps in the character of the possession or non-possession 



16 



ERNEST WARREN, 



Female (Lie metis). 


P3 


a; 
P5 


Light blue. 

Irregular oblong, 

set ijerpendieularly. 

Feathered. 

Receding. 


Scarcely at all 

developed. 
Shape compara- 
ti^ely slender ; 
hook long and 

curved. 
Dirty white. 

Coloured red. 


o 
I— 1 




a 

i 


Coloured orange 

and area smaller 

than in $ 


I— 1 


U 

if "> 


Veiy pale 1)1 ue 

Roughly circular 

Partly feathered 
Medium 


"* 


o 

o 
£ 

03 




2 

o ^ 




Less well-marked 
and regular 

Shape less massive; 

hook less straight 
and somewhat 
longer, curved 
Dark coloured, 
but less l)lack 


o 


Male (Cacatua). 


_o 
'oj 


o 


Chalky white 

Oval, set hori- 
zontally 
Naked 
Prominent 


Well-marked and 

regular 

Shape massive ; 

hook short, 

straight, and set 

perpendicularly 

Black horn 

Not coloured 


o 
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S 

o 

o 


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leathers 
Colour of naked 
area around eye 
Shape of naked 
area around eye 
Feathering of cere 
Prominence of 

forehead 


Granulation of 
eyelid 
Shape of tlie upper 
and lower mandi- 
bles and conditicm 
of hook 
Colour of bill 

Possession of 
coloured lores 


w 

a 

a 

o 

< 



HYBRIDISM AMOxNG COCKATOOS, 



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VOL. 3, PART 1. 



J 8 ERNEST WARREN. 

of coloured lores, it is nowhere possible to speak of a character 
as being dominant. In this case the actual colour of the 
lores is to be disregarded, and only the fact that some of the 
feathers of the lores are wholly coloured and not white in 
their upper portions would have to be taken into considera- 
tion. But the coloured area between the eye and the bill is 
relatively smaller in the hybrids than in Licmetis, and this 
is particularly the case in the younger hybrid. On the whole, 
therefore, it would appear easier to regard the condition seen in 
the hybrids as resulting from a blend of the male and female 
characters with a slight prepotency on the part of the female, 
than to take the view that the presence of the coloured lores 
is an example of dominance in the Mendelian sense. 

Finally, it may be said that out of these ten characters the 
hybrid is nearer to Cacatua in five, nearer to Licmetis in 
one, and almost exactly intermediate in four. 

The characters which lend themselves more readily to 
numerical expression may be summarised in the accompanying 
table (p. 17). 

In the fifth column the measurements in the hybrid are 
given, and in the third and seventh columns the differences 
between these measurements and those of the male (second 
column) and female (eighth column) respectively are shown. 
These differences, expressed in percentages of the measure- 
ments of the hybrid, are given in the fourth and sixth columns. 
Thus, for example, with respect to the length of the tail the 
divergence of the hybrid from the male- and female-parent 
expressed in percentage of the length of the tail in the 
hybrid is 14 per cent, and 32 per cent, respectively ; and 
the hybrid in this character is therefore 18 per cent, further 
removed from the female (L. nasica) than from the male 
(C. galerita). The means of these percentage ratio- 
differences between the hybrid and the parents in the eight 
characters tabulated are 25 per cent, for the male and 32 per 
cent, for the female. In other words, on the average, ratio- 
deviation of the hybrid from the male : ratio-deviation of the 
hybrid from the female : : 25 : 32. 



HYBRIDISM AMONG COCKATOOS. 19 

We see from this that the hybrid in these characters, just 
as in the case of the incommensurate characters tabulated on 
p. 16, stands nearer to the male than to the female. Of 
course, the actual ratio thus arrived at can be regarded only 
as the very roughest appi'Oxin)ation, since we are dealing with 
a single specimen of the hybrid. 

The only examples of hybridism among the Psittacidas 
that have been traced in the available literature are those 
quoted by Hans Przibram.^ 

"Macaws: Ara macao (^ and A. militaris ?, one 
young rather similar to militaris, died early. 

"Parakeets: Platycercus pallidiceps (J and PI. 
eximius (Roselle Parakeet) ?; also, Platycercus exi- 
mius cJ and PI. pennantii ? ." 

From the remark under macaws it would appear that the 
characters of militaris tend to be prepotent over those of 
macao. In this case it will be noticed that it was the 
female which was prepotent, while in the present example of 
the cockatoos it was the male. 

An interesting point in this connection is whether the 
prepotency of the male-parent in this case was due to the 
stronger inheritable nature of the Cacatua characters than 
of the Lie metis characters, or whether the prepotency was 
in some way connected with the male sex or with the indi- 
vidual specimen. To test this point it would be of great 
interest to obtain hybrids with male Lie metis and female 
Cacatua. 

Przibram,'^ in his recent work, has compiled a considerable 
list of cases of hybridism, and has illustrated a good number 
in a series of sketches. In many examples the hybrid is 
represented as being very strikingly intermediate between 
the two parent forms in all the general characters depicted. 
As far as can be seen, a very thorough blending of characters 
has mostly occurred, and from a general inspection it is often 

' Przibram, Hans, ' Experimental-Zoologie. Phylogenese,' p. 89, 
1910. 

2 Ibid., p. 89. 



20 ERNEST WARREN. 

not possible to say that the hybrid as a whole is closer to one 
parent than to the other. 

In the case of hybrid moths, the cross Smerinthus 
ocellatus ^ x Sm. populi ? [Westwood] is nearer to 
the male-parent ocellatus, and the cross Sm. populi (^ x 
Sra. ocellatus ? [Standfuss] is also nearer to the male- 
parent Avhich is populi. 

Similarly in the case of mammals, the cross Equus 
burchelli cJ x E. caballus ? [Ewart] is nearer to the 
zebra in the amount of striping, while the cross E. caballus 
cJ X E. burchelli ? [Ewart] is closer to the horse in the 
comparative absence of stripes in the hybrid. 

Also, the mule is perhaps somewhat closer in general 
character to the male-ass than to the female-horse, and the 
hinny is stated to be nearer the male-horse than to the female- 
ass. 

In these several examples the male appears to be pre- 
potent. 

Nevertheless, no general rule exists, for among fishes the 
cross Acerina cernua (J x Perca fluviatilis ? [Kam- 
merer] is strikingly close to A. cernua, while the cross 
Perca fluviatilis (^ x A, cernua ? [Kammerer] is also 
very near to cernua. 

In the cross of the moths Saturnia pavonivee cJ x 
S. pyvi ? [Standfuss] the hybrid is obviously nearer to 
pyri, the female-parent. 

In the case of recipi'ocal crosses of the moths Deilephila 
euphorbias c^ x D. vespertilio ? [Berge-Rebel] and of 
D. vespertilio (^ and D. euphorbiee ? [Berge-Rebel], 
the hybrids appear to be as nearly as possible intermediate 
between the parent species, and there is very little difference 
between them. 

In the above few examples of the crossing of distinct 
animal species the resulting hybrids exhibit considerable 
diversity with respect to their genei'al average character 
compared with that of the two parent-forms. 

We see that (1) the hybrids may be nearer to the male- 



HYBRIDISM AMONG COCKATOOS. 21 

parent than to the female; (2) the hybrids may be nearer to 
the female-parent than to the male ; (3) in reciprocal crosses the 
hybrids in both cases may be nearer to the male-species than 
to the female-species ; (4) also, it is probable that in some 
reciprocal crosses the hybrids in both cases may be nearer to 
the female-species than to the male-species ; (5) in reciprocal 
crosses the hybrids may be much alike, and nearly inter- 
mediate between the two parents. 

There appear to be no examples of the hybrids between 
distinct animal species being- widely different from both 
parents. 

We already know from Karl Pearson's researches that the 
co-efficient of correlation in man between male-parent and off- 
spring tends to be slightly greater in the inheritance of some 
characters than between female-parent and offspring. For 
example, in the case of stature the co-efficient for father and 
son is '396, and for mother and son *302. 

The few examples of hybridism above quoted do perhaps 
give some slight evidence of the male-parent sometimes exert- 
ing a certain general prepotent influence on account of sex ; 
but the influence of sex is doubtless altogether overshadowed 
by the prepotency of characters which may be regarded 
as phylogenetic. The very fact that a particular character 
occurs in a number of species of a genus is evidence that 
the character has been strongly inherited. In other words, 
the view is, that in blended inheritance if one parent of a 
hybrid has the typical character of the genus, and the other 
parent has a less usual one, then the typical character seen in 
the majority of the species of the genus would tend to be 
more strongly inherited in the hybrid than the more peculiar 
one. 

Possibly, if it is legitimate to surmise at all in the matter, 
the hybrid Licmetis nasica ($ x Cacatua galerita ? 
would not differ greatly from the hybrid above described, 
since the characters ^appertaining to the large genus Cacatua, 
to which galerita belongs, might be expected to conceal any 
small prepotent influence that the male sex might possess. 



22 



ERNEST WARREN. 




HYBRIDISM AMONG COCKATOOS. 23 

From this point of view a glance at Table III may be of 
interest. In this table the characters of the different species 
of Cacatua, as given in the British Museum Catalogue, are 
rather imperfectly summarised; but we are able to see the 
general range of variability in the genus, with a view to 
obtain some insight into the hereditary tendencies of the 
male-parent ; and also of the female-pai'ent, since the genus 
Lie metis to which the latter belongs is closely related to 
C a c a t u a . 

For example, the general coloration in the different species 
of Cacatua is nearly always white, and we find as the result of 
the blended inheritance that the amount of coloration in the 
hybrid is less than half of that of the female-parent. 

The recurved condition of the crest is found in less than 
half of the species of Cacatua, and we notice that the 
recurved feathers of the male-parent are not so strongly 
impressed on the offspring as to counteract the influence of 
the female-parent. Yellow or white crests occur in the 
majority of the species of Cacatua, and the amount of red 
coloration mixed with the yellow of the hybrid is compara- 
tively small. 

The feathered and non-feathered conditions of the cere are 
about equally divided in the different species, and in the 
hybrid the condition with respect to this matter is almost 
exactly intermediate between the two parents. 

The bill is more often black than light-coloured in the 
various species, and in the hybrid it is much nearer to black 
than white. 

In nearly all of the species of Cacatua the naked skin 
around the eye is oval or circular, and in the hybrid it is 
roughly circular. The shade of colour of the naked skin, 
whether Avhite, blue or grey, varies considerably in the 
different species of Cacatua, and the hybrid in this character 
is closely intermediate between the two parents, thus indicat- 
ing that there is no marked preponderating influence in either 
direction. 

The general shape of the upper and lower mandibles is very 



24 ERNEST WAKEEN. 

similar in all the species of C a c a t u a , while it is rather peculiar 
in Licmetis; in the hybrid the general shape is nearer to 
that in the former genus than to that in the latter. 

Lores separable from the surrounding plumage in colour are 
seen apparently in only three or four of the fifteen species of 
Cacatua, and to be comparable with the above results the 
colouring of the lores in the hybrid should be less than half 
the amount seen in Licmetis. The amount of colouring in 
the hybrid is certainly considerably less than in the female- 
bird, but it is greater than one-half, and consequently in this 
character there is some prepotency on the part of the mother. 

With reference to the absolute dimensions of general body- 
length, length of tail and bill, given in the second, third and 
fourth columns of Table III, it will be seen that there is great 
variation in the different species, and this is also the case with 
the relative lengths of the tail and bill. Consequently we 
perhaps should not expect to find any very strong prepotent 
action. As a matter of fact, the absolute dimensions in the 
hybrid are not so very far removed from the means of the 
dimensions of the male- and female-parent. The means for 
the parents of the three dimensions are 15"5, 6'4 and 1*74 
inches respectively, and the measurements in the hybrid are 
15'0, 7"0 and 1"7 inches respectively; the male-parent is 
prepotent with reference to the length of tail and beak, and 
the female-parent with respect to body-length. The mean 

,. length of tail n ^^ .^ ■ • no £ 1 

ratio ■—- — -^ for all the species is '28, tor male 

length of body 

Cacatua "45, for female Licmetis "36, and for hybrid '45. 

Thus, in this case the prepotency of the male-bird has been 

strong enough to counteract the more typical proportion of 

the tail-length seen in the female-bird. 

On the whole it perhaps may be said that old-established 

characters appearing in the majority of the species of a genus 

may from this very fact be regarded as strongly inheritable 

characters, which if present in one parent would tend to be 

impressed on the hybrid to the partial exclusion of any more 

peculiar characters that happened to be present in the other 



HYBRIDISM AMONG COCKATOOS. 25 

parent. Thus, the former parent (Cacatu a in the present 
case) would appear to be prepotent over the latter parent 
( L i c m e t i s ) . 

There is exhibited, as a rule, no simple Mendelian relation- 
ship in the hybrids between animal species. The characters 
of the parents tend to blend, and often they blend very 
thoroughly as in the cockatoos: and although the hybrid 
may be nearer in certain characters to one parent than 
to the other, yet hi no cliaracfer />■• flie direct injhience of 
either parent apparently absent. No character observed 
could be described as "dominant" or "recessive," unless 
these terms are used in a popular sense, and a dominant 
character is simply a prepotent one. It is true that the 
Mendelian relationship concerns characters only find not 
individuals as a whole. Thus, supposing that the hybrid 
cockatoos were capable of crossing among themselves and 
that the Mendelian mode of inheritance occurred, then the 
offspring would not be wholly like C. galerita or wholly 
like L. nasica, but some of them would very strougly 
resemble one or the other of the paient forms in some of 
their characters. According to tlie Mendelian conception of 
hereditai'y transmission by definite unit-characters, ever}' such 
character is capable of being replaced by an alternative, and 
by the discovery of the alternatives (Allelomorphs) the exact 
nature of the units can alone be determined. A character 
like stature or other dimension of an animal is obviously a 
complex one which could not be expected to follow the 
Mendelian mode of inheritance as a whole, unless groups of 
characters are supposed to be capable of remaining connected 
together in some obscure manner and of being transmitted as 
a single character. 

Blended inheritance is the antithesis of exclusive inheri- 
tance : and in the case of the latter the Mendelian relation- 
ship can be readily understood on the simple and ingenious 
supposition propounded by Mendel of the segregation of i he 
gei-minal cells. The extreme jMendelian school tend to deny the 
existence of real blended inheritance ; they attempt to explain 



26 ERNEST WARREN. 

tlie apparent blending as arising through the character being 
a compound one which requires analysing into its component 
parts before the Mendelian relationship can be recognised. 

In breeding experiments for testing the occurrence of the 
Mendelian relationship it is most important, as was pointed 
out by Prof. Weldon some years ago, that the categories into 
which the offspring are grouped should be clearly defined. 
In practice it is sometimes extremely difficult to sort out the 
offspring into clearly defined groups, and unfortunately it 
may become a matter of personal judgment as to whether an 
individual should be placed in one or another category. 

Colour, independently of its distribution, may doubtless be 
regarded sometimes as a simple character, and if the present 
hybrid were fertile, and the Mendelian relationship occurred, 
we should have, assuming that the yellow was dominant, 
25 per cent, sulphur-yellow offspring (pure dominants), 25 
per cent, red offspring (pure recessives), and the remaining 
50 per cent, would be yellow impure dominants. But the 
character of the hybrid of the first generation does not favour 
the view that such a result would be obtained. Neither colour 
is dominant, but the yellow and red have blended to form a 
kind of orange in which there is more yellow than red. I have 
not attempted to obtain a numerical expression for the amount 
of the two colours in the orange of the hybrid, although 
possibly such could be obtained. 

Again, the possession or absence of a sulphur-yellow 
lanceolate recurved feathered crest as seen in C. galerita 
might be conceivably a simple character; but the feathers 
in the hybrid corresponding to the recurved crest-feathers of 
the male are not recurved, they are relatively wider and 
much shorter, and they are in fact more or less intermediate 
in character between the feathers in the male and female. 

Thus in every character examined, with the possible 
exception of coloured and non-coloured lores, there is a very 
obvious blending of the male and female characteristics ; and 
although the external appearance of any hybrid is not to be 
regarded as an absolute guide to its inherent gametic 



HVHKIDIS.M AMONG COCKATOOiS. 27 

cliaracter, the evidence, as far as it goes, would point to a 
real blending of the characters of the parents, and the 
probable absence of so-called gametic purity, or the segrega- 
tion of characters in their sexual elements. 

If characters do not blend, and each hybrid parent, what- 
ever the external appearance may be, produces germ-cells in 
equal numbers capable of producing either one character or 
the other, but not a mixture of both, then the relationship 
found by Mendel and others would naturally follow in the 
offspring by the law of chance in the fortuitous union of 
the male and female elements. 

The hybrids of Mendel, De Viies, Bateson, and other 
observers are more generally crosses of varieties of the 
same species, and the reason why the crosses between 
different species have not been more often employed is 
owing to the fact that these hybrids are frequently sterile 
among themselves ; but it might be reasonably expected 
that the appearance of dominant characters would be more 
often observable in the first generation of hybrids of distinct 
species, if any approach to gametic purity actually occurred 
in their sexual cells. 

If differentiation of the sexual cells with regard to alterna- 
tive characters of the parents really occurs in these hybrids, 
it is highly remarkable that the parent-hybrid itself should 
usually be approximately intermediate in all its characters 
between the two parent-species. The fact indicates that the 
great majority of the characters are capable of blending. 
Should the Mendelian relationship possess that universality 
which some are inclined to attribute to it, the hybrids derived 
from the crossing of distinct species ought certainly to afford 
some decisive evidence in its support. 

It is becoming increasingly obvious that MendeFs law in its 
original sense is of quite limited application, since very many 
apparently simple characters appear to blend intimately, so 
that there can be no question of gametic purity or segregation. 

Into the debatable question that the blending is only 
apparent, owing to the characters being complex ones which 



28 ERNEST WARBEN. 

require analysing into their component parts before the 
Mendelian relation becomes apparent, I cannot enter here, 
and can but remark that this view offers a wide iield for 
experimentation. 



EXPLANATION OF PLATE II, 

Illustrating Dr. E. Warren's paper, " A Case of Hybridism 
among Cockatoos," 

Fig. 1. — X f. Side view of head and neck of Licmetis nasica, 
female-parent. 

Fig. 2. — x f. Similar side view of Cacatua galerita, male-parent. 

Fig. 3. — x f . Hybrid three years old. 

Fig. 4. — x f . Hybrid three months old. 









^ 



CO 



^ 



/ 



cd 
cd 






CO 



ON SOUTH AFRICAN ENNEJi. 29 



On South African Ennese, with Descriptions of 
New Species and Varieties. 

By 

Henry Clifdeii Biiriiiip. 



Witli Plates III, IV. V. and Text-fi^s. 



Contents. 

Introduction 

Systematic Account .... 
Ennea isipingoensis Stur. and its varieties 
Ennea isipingoensis Stur. f . tyj^ica . 

var. discrepans Stur. 

var. stui'anyi n. . 
Ennea elliptica Melv. & Pons, f . typica 

var. manca n. 

var. cselata n. 
Ennea farqnhari Meh\ ScPons.i. typica 

var. bertlise {Melv. & Pons. 

var. avena n. 
Ennea darglensis Melv. & Pons, f . typica 

var. illovoensis n. 
Ennea consobrina Ancey. 
Ennea tlielodonta Melv. & Pons. 
Ennea ninnita Melv. & Pons. 
Ennea sylvia Melv. &■ Pons. 
Ennea melvilli n. sjj. 
Ennea columnella Melv. & Pons. f. typica 

var. vitreola (ilfeZw. & Pons. 
Ennea marise Melv. c{- Pons. 
Ennea niooiensis n. sp. 
Ennea maritzburgensis Melv. d- Pons. f. typica 

var. contracta n. . 
Ennea arnoldi Stur. 



PAGE 

30 
31 
31 
33 
35 
36 
38 
39 
40 
41 
44 
46 
48 
49 
50 
50 
52 
54 
55 
58 
59 
61 
62 
64 
66 
68 



30 HENRY CLIFDEN BUBNUP. 

Ennea connollyi Melv. & Pons. 
Ennea inhluzaniensis n. sp. . 
Ennea jiixtidens Melv. & Pons. 
Ennea premnodes Star. 
Ennea montana Melv. & Pons. 
Ennea ponsonbyi n. sj). 
Explanation of Plates 



69 
71 

72 
73 

75 

78 
80 



Introduction. 

In the following pages an attempt is made to clear up some 
of the difficulties that stand in the way of the student seeking 
an intimate acquaintance with the genus Ennea, as repre- 
sented in South Africa. Special attention is given to the 
comparison of kindred forms and to the consideration of the 
value of the diiferences between one form and another, 
whether they amount, in the opinion of the writer, to specific 
or varietal distinction, or are too trivial to warrant separate 
treatment. 

The result of the investigation, as far as it goes, is the 
recommendation that seven species and two varieties be con- 
sidered synonyms, and two species be degraded to varietal 
rank. It has been found necessary to differentiate six varieties 
of known species, and four species that do not seem to have 
been described. 

The alterations involved are as follows: Ennea laby- 
rinthea, microthauma, hypsoma, oppugnans, and 
periploca of Melv. & Pons, to become synonyms of Ennea 
is,r cinhari Melv. & Fons.i. typica; Ennea callista J/eZv. & 
Pons, to become a synonym of Ennea marine Melv. & Pons.; 
Ennea parallela Melv. & Pons, to become a synonym of 
Ennea montana Melv. & Pons.; Ennea isipingoensis 
Stur. var. cylindrica Stwi'. to become a synonym of Ennea 
isipingoensis f. typica; Ennea isipingoensis var. 
simillima Stur. to become a synonym of Ennea 
isipingoensis var. discrepans Stur. : Ennea berthas 
3Ielv. & Pons, to become a variety of Ennea farquhari 
Melv. & Pons.; and Ennea vitreola Melv. & Pons, to become 
a variety of Ennea columnella Melv. & Pons. 



ON SOUTH AFUICAN ENNE.?':. 31 

The new varieties introduced are as follows : Ennea isi- 
pingoensis Shir. var. sturanyi ; E. elliptica Melv. & 
Pons. var. manca ; E. elliptica ilie/y. & Pons. var. caslata; 
E. farquhari Melv. & Pons. var. avena; E. darglensis 
Melv. & Pons. var. illovoensis; E. maritzburgensis 
Melv. & Pons. var. contra eta. And the species, described 
as new, are : Ennea mel villi, E. mooiensis, E. inhluza- 
niensis, E. ponsonbyi. 

It has also been thoug-ht desirable to re-describe and re- 
figure a few forms whose original descriptions and figures 
iippeared to be erroneous or insufficient in detail, or had been 
published in works not easy of access to South African 
readers. 

The difficulty presented by the writer's inability to examine 
the types in the British Museum has largely been overcome 
by the readiness of friends on the spot to compare them with 
specimens and drawings sent for the purpose, and to report 
upon such coincidence or disagreement of character as might 
be observed. To those ready hewers the writer's most cordial 
thanks are extended, and their special services are duly 
acknowledged under the heads of the several species involved. 
No less are sincere thanks due to those who have supplied 
material for study, prominent among whom are Messrs. J. H. 
Ponsonby, J. Farquhar, and A. J. Taynton. 

The types of the species and varieties described as new, 
together with a few of the other shells figured herein, will be 
deposited in the British Museum. A reference to the explana- 
tion of the plates at the end of this paper will show the 
resting-place of each shell figured. 

Systematic Account. 

Ennea isipingoensis Sturany; and its varieties. 

Through the kindness of Dr. Rudolf Sturany in lending 
me his types of this species and its three varieties described 
at the same time, I have been enabled to acquire a better 



32 HENRY CLIFDEN BUENUP. 

knowledge of the group than has hitherto been possible. 
After a careful examination of the original specimens, and a 
close comparison of them with about 140 other examples, 
collected principally by myself in various parts of the 
province of Natal, and three specimens collected by Mr. 
Farquhar in the province of the Cape of Good Hope, I am 
convinced that two of Dr. Sturany's varieties are superfluous, 
and that another form, hitherto undescribed, requires to be 
differentiated as a distinct variety, in order to make the 
group, so far as it is 3'et known, fully understood. In 
justice to Dr. Sturany, and in order to show how the mis- 
takes have arisen, it is necessary to explain that the original 
material was very meagre, and insufficient for forming a just 
estimate of the value of the apparent differences ; for Dr. 
Penther only found four specimens, which, appearing to 
show sufficient differences, were described provisionally as 
the type and three varieties. Moreover, the shells were very 
dirty, and, possibly through the fear of breaking unique 
specimens, had never been properly washed, for they reached 
my hands much obscured and disfigured by sand and leaf- 
mould. The specimen which was described as the variety 
cylindrica was calcined and had a large hole in the body- 
whorl, and to wash it would probably have led to its 
destruction ; so I was reluctantly obliged to leave it as it 
was, except that with a damp brush I carefully removed 
enough of the mould from its aperture to expose the small 
basal tooth, which is said in the description to be absent, 
and to show more clearly the tooth within the outer lip, and 
the columellar plait, which are said to be respectively weaker 
and not very broad, to be practically identical with those of 
the typical form. The type of var. discrepans had the 
apex crushed in, but its general condition was such as to 
justify washing, though not syringing : the other two 
specimens were sound. 

The fragile condition of the specimen described as var. 
cylindrica has prevented my re-figuring it and checking its 
dimensions, but neAv figures of the other specimens are 



ON SOUTH AFRICAN ENNE^. 33 

given, and their dimensions as taken by me will be found in 
the text. The original figure of the typical form does not 
require to be replaced, except for the convenience of 
having all the figures to be compared of one magnifica- 
tion. 

The original description adequately dilferentiates the 
species, but, for the sake of convenience in comparison, it 
has been deemed advisable to re-describe it here, where also 
will be found an amended description of var, disc re pans 
Stur., and an original description of a new variety with 
which I have much pleasure in associating the name of the 
learned author of the species, who has rendered me such 
matei'ial help in the present study. 

Ennea isipingoensis Sturany, f . typica. PI. Ill, 

fig. 1. 

Ennea isipingoensis Stur., Anz. k. Akad. Wissensch. Wien (1898), 
No. xvi, p. 157 (reprint p. 5) ; Sturany, Siidafrik. Moll. (1898), p. 27, 
pi. i, fig. 19. 

Ennea isipingoensis var. eylindrica Stur., ibid., ibid. , fig. 22. 

Ennea isipingoensis Star.; (sic) Connolly, Ann. S. Af. Mus., vol. 
xi (1912), p. 77. 

Shell small, cylindric-oval, tapering towards the blunt apex, 
neai-ly white, umbilicate ; whorls 7h, very convex, separated 
by very deep, almost channelled sutures, and very strongly 
transversely ribbed except the embryonal whorls, which are 
smooth, and immediately behind the peristome where the ribs 
become finer ; the last whorl is thickened at the base, forming 
a callosity almost encircling the umbilicus, and then becomes 
constricted before expanding to form the peristome ; aperture 
small roundish and furnished with the following plaits : A 
strong parietal plait arising higher on the body-whorl than 
the peristome and passing thence, embracing the parietal 
wall, to the interior of the shell ; a long, strong squarish- 
pointed, tooth-like plait, almost bifid, arising near the margin 
and about the middle of the labrum, extending nearly to the 

VOL. 3, PAET 1. 3 



34 HENRY CLIFDEN BURNUP. 

centre of the aperture, and the lower edge of it penetrating 
furthest inwards ; near the base of the columellar lip, a small 
conspicuous tooth ; at the base, and set further back than the 
two processes last named, an inconspicuous denticle ; and a 
deep-seated, scoop-shaped columellar plait with mamillated 
point on the inner edge presented forwards ; peristome white, 
broad, expanded and reflexed, the ends connected by a slight 
callus. 

Height 2-83, width 1-47 mm. 

Hab. — Isipingo, near Durban (Penther) ; other localities: 
Natal : Kar kloof and Nottingham Road (Taynton), Lower 
Umkomaas, Umbogintwini, Ntimbankulu — Mid-Illovo, Maritz- 
burg, Edendale, Dargle and Inhluzani (Burnup). 

Cape of Good Hope : The Gorge, Van Staaden^s River 
(Farquhar) . 

The umbilicus is rather widely open for so small an 
Ennea, and, as is usual in openly umbilicated shells of the 
genus, has a branch of the perforation passing into the 
columellar fold. In the type the callus connecting the ends 
of the peristome is not very clearly defined, and I can readily 
understand the author having overlooked its presence before 
the shell was cleaned. In most examples it is much better 
developed, sometimes obliterating the sculpture underneath, 
but in many instances, especially in rather young, though 
mature shells, alloAving the ribs to be seen through it. On 
the outside of the shell can be seen, not only a large deep 
cavity corresponding with the strong labral plait, but also 
smaller, shallower pits marking the positions of the basal 
denticle and the basi-columellar tooth. The distribution^ 
including a locality so remote as Van Staaden^s River in the 
Cape Province, is worthy of note : the specimens from the 
last-named locality are quite normal. 

Since I have shown the type of the variety cylindrica to 
be identical in its characters with the type of the species 
except in size and slightly in form, it becomes inadmissible to 
recognise it as a distinct variety. The dimensions given by 
the author show the two shells to be identical in the ratio of 



ON SOUTH AFRLOAN ENNE^. 



35 



width to height, though to me cylindx-ica seems per- 
ceptibly naiTOwei'. The following table of dimensions of a 
few shells that I have measured will indicate the futility of 
any attempt to identify a variety based on size and form 
alone. 



Specimen. 


Height. 


Width. 


Width X 100 
Height. 






mm. 


mm. 




E. 


isipingoeiisis{type). Isipingo 
var. cylindrica (original 


2-83 


1-47 


.51-94 




measui-ement). Isipingo 


3-00 


1-50 


50-00 




Inliluzani . . .J. 


2-74 
3-27 


1-66 
1-69 


60-58 
51-68 




r 


3-18 


1-85 


58-18 




Ntimbankulu . . .< 


3-64 


1-86 


51-10 




I 


3-94 


1-73 


43-91 




Edendale 


2-99 


1-48 


49-50 




Umbogintwini 


2-87 


1-42 


49-48 




Lower Umkomaas . 


3-80 


1-74 


45-79 


Average . . . . 


3-23 


1-64 


51-22 ' 



The ratios given in the fourth column show great variability 
even in specimens from the same locality, e . g. Ntimbankulu. 

Var. discrepans Stnrany. PI. Ill, figs. 2, 3. 

Ennea isipingoensis Stiir. var. discrepans Stiir.. Anz. k. Akad. 

Wissenscli. Wien (1898), No. xvi, p. 157 (reprint j). 5) ; Sturany, 

Sixdafrik. Moll. (1898), p. 27, pi. i, fig. 20. 
Ennea isipingoensis Stur. var. simillinia Stu7\, Anz. k. Akad. 

Wissensch. Wien (1898), No. xvi, p. 157 (reprint p. 5) ; Sturany, 

Slidafrik. Moll. (1898), p. 27, pi. i, fig. 21. 

Shell like E. isipingoensis, typical, but with the following 
differences : The contour is more oval and less cylindrical ; 
the aperture is larger and rather oblique and triangular ; the 
processes of the aperture are similarly placed but differ in the 
following respects : The labral plait is much enlarged, being 
both longer and wider, and bears a distinct, though minute, 
denticle on the upper edge ; the minute basal denticle is 
replaced by an inconspicuous, scarcely raised, minute ridge. 




36 



HENEY CLIFDEN BUENUP. 



whose presence is more easily detected by the corresponding 
depression and callosity behind the basal lip than by its 
appearance within ; and the columellar plait is much swollen. 

The labral and columellar plaits are so long that they 
overlap, the former in front of the latter. 

Height, without apex, which is crushed into the shell, 
2-69, width 1-45 mm. 

Hab. — Isipingo, near Durban (Penther). Other localities : 
Umbogintwini and Equeefa (Burnup). 

This variety, including var. simillima Stur. (fig. 3), which 
is identical in all essential characters, seems to offer nearly 
as much variation in size as the typical form, the dimensions 
of the only five specimens that I have seen being as follows : 



Specimen. 


Height. 


Width. 


Width X 100 
Height. 


Type of var. discrepans from 
Isipingo 

Type of "var. simillima" frora 
Isipingo 

Specimen from Umbogintwini . 

Specimens from Equeefa . J. 


mm. 

2-69 

2-72 
2-78 
3-49 
3-56 


mm. 

1-45 

1-48 
1-47 
1-77 

1-78 


53-90 

54-41 
52-88 
50-72 
50-00 


Average . . ... 


305 


1-59 


52-38 



The relative width to height is much more regular in this 
variety than in the typical form, so far as can be ascertained 
from so small a series. 



Var. sturanyi n. PL III, figs. 4, 5, 6. 

Shell like E. isipingoensis, typical, but differing in the 
following respects : The labral plait is both longer and 
wider and is more compact, ending in a blunted or rounded 
point extending beyond the centre of the aperture; the 
columellar plate is also larger and extends to the centre of 
the aperture, thus passing slightly behind the point of the 



ON SOUTH AFRICAN ENNE^. 



37 



labral plait ; and the basal denticle is replaced by an incon- 
spicuous, scarcely raised, minute ridge like that of var. 
discrepans. It differs from var. discrepans in being 
more cylindrical, in the labral process being less massive and 
more pointed, and in the total absence of the conspicuous 
denticle on the upper edge of the labral plait. 

Height, 2-95, width, 1"55 mm. 

Hab. — Ntimbankulu, Mid-Illovo (Burnup). Other locali- 
ties: Karkloof (Taynton), Maritzburg, Howick and Dargle 
(Burnup) . 

The following dimensions of shells that I have measured 
will give an indication of the extent of variation in size and 
proportion that this form is liable to : 



Specimen. 


Height. 


Width X 100 
^"l*^- Height. 

1 


Type of var. from Ntimbankuhi 

l 

Specimens from Dargle 


mm. 

2-95 
2-57 
2-80 
2-86 
2-91 
313 
3-03 
3-25 


mm. 

1-55 
1-48 
1-52 
1-48 
1-49 
1-52 
1-47 
1-52 


52-54 
57-59 
54-29 
51-75 
51-20 
49-20 
48-18 
46-77 


Average 


2-94 


1-50 


51-44 



It will be seen from the above table that, while this variety 
is less stable in ratio of width to height than the variety 
discrepans, it does not exhibit so much variation in that 
respect as isipingoensis, typical. It will also be seen that,^ 
though some individual specimens of the other forms may be 
smaller than some examples of this, in striking an average of 
the shells measured, sturanyi is the smallest, and isipin- 
goensis, typical, the largest. 

The nature and disposition of the armature of the aperture 
in the several forms remain remarkablv constant, no inter- 



3B HENRY CLIFDEN BUENUP. 

mediate forms having been observed in the very large series 
examined. 

Dr. Sturany has kindly perused my MS. and compared my 
figures with his original shells, and concurs in the elimination 
of the two vai'ieties, simillima and cylindrica. 

Ennea elliptica Melv. & Pons, f . typica, PI. Ill, 
figs. 7, 8, 9, 10, 11, 12. 

Ennea elliptica Melv. & Pons., Ann. and Mag. Nat. Hist., vol. ii 
(1898), p. 126, pi. vii. fig. 2 ; Connolly, Ann. S. Af. Mus., vol. xi 
(1912), p. 73. 

In the original figure, which fairly well represents the 
characters of the peristomatal dentition, omitting, however, 
the internal columellar plait, the aperture is drawn out of all 
proportion to the rest of the shell. It is shown to occupy 
about 35 per cent, of the whole height, whereas in normal 
specimens it does not occupy more than about 28 per cent. 
This would make it appear either that the type is a malformed 
shell or that the figure is bad. As the authors had before 
them, at the time the description was written, four specimens 
agreeing in detail, it is not likely that the type is abnormal ; 
therefore it is to be inferred that the figure is incorrect. In 
any case it seems desirable to publish a new figure, which is 
given on PI. Ill, fig. 7. This drawing represents a normal 
shell from the original locality in the Botanical Gardens,^ 
Maritzburg. Fig. 8, drawn from another specimen from the 
same locality, represents the right side of the shell and shows 
the striate sculpture on the last half whorl. 

This species is very interesting in exhibiting, through large 
series from difl:erent localities, the undoubtedly close relation- 
ship between forms which, but for the intermediates, might 
possibly have been taken to be distinct species. 

' The Botanical Gai'dens, Maritzburg, as a faimistic locality, must not 
he looked vipon with suspicion as indicating the probability of moUuscs 
having been introduced with plants from a distance. The spot indicated 
is a bush-clad hill within the boundaries of the gardens but not dis- 
turbed by cultivation. 



ON SOUTH AFRICAN ENNE.E. 39 

In the bush skirting the beach near the mouth of the 
Tongaat Eiver, the only place near the coast where I have 
met with this species, is found a form, quite typical except 
that it is rather small and has the peristome rather more 
thickened, and the processes yerj strongly developed. Two 
examples of this form are shown in figs. 9 and 10. 

Next comes the typical form from Maritzburg at an eleva- 
tion of a little over 2000 feet (fig. 7). 

At Dargle, at an altitude of about 3700 feet, the shells are 
barely larger than those at Tongaat, and the lower tooth on 
the columellar lip has deteriorated into a mere inconspicuous 
callosity, as seen in figs. 11 and 12. 

The next stage is seen in the shells found at Fort Notting- 
ham and Curry's Post (altitude about 4000 feet), in which the 
height is rather greater, and the lower tooth of the columellar 
lip is suppressed altogether. I think this form may con- 
veniently be regarded as a new variety, differentiated as 
follows. 

Yar. manca n. PI. Ill, figs. 13, 14. 
Shell like el lip tic a, typical, but generally rather larger, 
comparatively narrower, and without the lower tooth on 
the columellar lip. 

Height 3*76, width 1*74 mm. 

Hab. — Fort Nottingham; also Curry's Post, Natal (A. J. 
Taynton). 

In a few specimens from these localities an almost imper- 
ceptible callosity may be detected in the position of the lower 
tooth of the columellar lip of elliptica, typical; but in by 
far the greater number there is no trace of it. There are 
also to be found in some specimens, especially those from 
Curry's Post, slight traces of the sculpture distinguishing the 
variety caelata, hereafter described, for the most part only 
infrasutural, but occasionally extending across the whorls. 
The Curry's Post specimens are rather wider than those from 
Fort Nottingham. 

Yet another form, varying from the type in respects not 



40 



HENRY CLIFDEN BURNUP. 



affecting the gradual evolution from the Tongaat form to the 
variety mane a, that I have endeavoured to trace out above, 
occurs at Eshowe, Zululand. Mr. Ponsonby has placed in my 
hands for determination three examples of this form, Avhich I 
propose to distinguish as a separate variety, c^lata. 

Var. CEelata n. PL III, figs. 15, 16, 17. 

Shell like elliptica, typical, but rather larger, more obese, 
with peristome rather thinner and less reflexed, covered all 
over, except the earlier whorls and the body-whorl above the 
aperture, with most delicate, fine, oblique strife becoming- 
stronger on the last half -whorl, Avhere, however, they are not 
so strong as those in a similar position in the typical form. 

Height 4-06, width 2-11 mm. 

Hab. — Eshowe, Zululand (Ponsonby). 

The other two specimens are not so obese as the type of the 
variety, which was chosen as being the best developed speci- 
men. Thus, while they agree better with elliptica, typical, 
in contour, they are more divergent in their still greater 
height and yet thinner peristome. In all three the distinctive 
sculpture is clearly seen by the aid of a strong lens. 

The following additional measurements will help to convey 
an idea of the variation as to size : 



Specimens from various localities. 


Height. 


Width. 


Figure on 
Plate. 


F. typica, Maritzburg . . 3 

F. typica, Tongaat . . ) 

F. typica, Dargle . . . ) 

Var. mane a, Fort Nottingham 
Var. manca, Curry's Post . - 

Var. cselata. Eshowe . . \ 


mm. 

405 
3-84 
3-75 
3-60 
317 
314 
311 
3-38 
317 
300 
4-12 
3-93 
3-91 
4-44 
4-09 


mm. 

1-82 
1-87 
1-85 
1-73 
1-63 
1-71 
1-65 
1-64 
1-66 
1-66 
1-79 
1-92 
1-90 
2-06 
202 


fig. 7. 
fig. 9. 
fig. 10. 

fig. 11. 
fig. 12. 

fig. 14. 



ON SOUTH AFRICAN ENNE^. 41 

In considering the figui-es illustrating the various forms of 
this species, it should be borne in mind that they are not all 
drawn to the same scale ; therefore it is the more desirable to 
keep in view the dimensions given above. 



Ennea farquliari Melv. & Pons, f . typica. PI. IV, 
fig. 24. 

Ennea farquhari Melv. & Pons., Ann. and Mag. Nat. Hist., vol. xvi 

(1895), p. 478, pi. xviii, figs. 3-5. 
Ennea labyrinthea Melv. Jt Pons., ibid., p. 479, pi. xviii, figs. 7, 8. 
Ennea microtliaiima Melv. & Pons., ibid., vol. iv (1899), p. 194, pi. 

iii, fig. 1. 
Ennea hypsoma Melv. & Pons., ibid., vol. iv (1909), p. 488, pi. viii. 

fig. 7. 
Ennea oppugnans Melv. & Pons., ibid., p. 488, pi. viii, fig. 8. 
Ennea periploca Melv. S: Pons., ibid., p. 489, pi. viii, fig. 10. 
Ennea farquhari M. & P.; Connolly, Ann. S. Af. Mus., vol. xi 

(1912), p. 74. 
Ennea microthauma M. A- P. ; Connolly, ibid., p. 80. 

Shell minute, ovate-cylindrical, with rounded apex, rimate 
and narrowly perforate, thin, vitreous, transparent and 
shining; whorls 5^, moderately ventricose, the first two 
smooth, the rest sculptured with regular transverse stride, 
strong below the suture and evanescent lower on the whorls, 
except the last, on which the sculpture continues round the 
base ; suture moderately impressed ; aperture triangularly 
auriform with thickened, expanded and reflexed, white, 
rather vitreous peristome, and furnished with the following- 
processes : a prominent blade-like, in-running, parietal plait, 
a massive, blunt, lobed, labral tooth, a small internal basal 
tooth, an inconspicuous, broad, slightly raised tooth on, and 
parallel to, the columellar lip, and a deep-seated, rounded, 
flatfish, scoop-shaped columellar plait. 

Height 2-11, width 1-03 mm. 

Hab. — Grahamstown (Farquhar and Langley) ; also High- 
lands (19 miles N. of Grahamstown) and Bathurst 
(Farquhar), all in the Cape Province. The locality, Bosch- 



42 HENRY CLIFDEN BURNUP. 

berg Mountain, Somerset East, given in the description of 
Ennea periploca, is erroneous (see p. 61). 

The right side of the parietal plait, which is very promi- 
nent, arises higher on the body- whorl than the labrum ; the 
upper lobe of the bipartite labral tooth is a little within the 
plane of the aperture, the lower is more internal ; the short, 
broad tooth on the columellar lip is nearly level with the 
general plane of the peristome ; and the small basal tooth 
stands well back from the edge. A narrow channel enters 
the columellar plait from the rimal cavity, and the teeth on 
the labrum. columellar lip and base, have corresponding 
excavations behind the peristome. 

The types of all the so-called species here placed in the 
synonymy, being in the British Museum, are not available to 
me for examination, but I have been fortunate in securing, 
from the collections of Messrs. Ponsonby, Farquhar, and 
Langley, a very large series comprising co-types of every 
form, except that which was described as E. labyrinthea. 
These I have submitted to a searching study and careful 
comparison, and cannot doubt that all belong to one variable 
species, whose divergence from a common type is not wide 
enough to justify the retention of any of the later names in a 
varietal capacity. The form described as Ennea berthas 
Melv. & Pons., as represented by my co-type, ex coll. 
McBean, supported by seven other specimens from three 
localities in Natal, which undoubtedly also belongs to the 
same species, is specially considered hereafter. 

Of the form described as E. labyrinthea, alone, I have 
been unable to obtain any reputed example, and so have only 
had the original description and figure to guide me in uniting 
it with E. farquhari. 

Melvill and Ponsonby not being able to concur in my views, 
and I being unable personally to consult the types, and 
hesitating to make su^ch a sweeping consignment to synonymy 
without reference to them, Mr. Ponsonby most kindly 
arranged, through Major Connolly, Avith Messrs. E. A. Smith 
and G. C. Robson, of the British Museum, that the two last- 



ON SOUTH AFRICAN ENNE^. 43 

named should examine my numerous camera drawings of all 
the forms under consideration (except, of course, laby- 
rinthea), compare them with the types in the British 
Museum, and generally consider the case of synonymy that I 
had inade out. Their deliberations have resulted in the 
following decision, dated at the British Museum (Natural 
History), CromAvell Road, December 6tli, 1912 : 

'' We have examined the seven species named below,^ as 
requested by Major Connolly, and after careful consideration 
have come to the conclusion that, making due allowance for 
variability, the species in question are all indistinguishable 
from E. farquhari save in a few unimportant characters, 
the general characters of the aperture and dentition remain- 
ing the same in all seven. 

(Signed) G. C. Kobson, 
E. A. Smith." 

I am deeply indebted to these gentlemen, for without their 
co-operation my conclusions must have remained unconvincing- 
through the possibility of the material studied by me being 
wrongly identified. 

Major Connolly has been good enough to supply me with 
revised measurements of all the types, which in some cases 
vary considerably from those originally published; but it 
seems to me more useful to give here a selection of dimen- 
sions of specimens from the various localities, than to emend 
those of the specimens originally described under names 
which it is sought to show should drop out of use. To this 
resolve, however, it may be well to make exception in the 
cases of the original type of E. farquhari, and the shell 
originally described as E. labyrinthea — the former because 
it is the persistent type, and the original dimensions are 
manifestly wrong, no specimen in the large series examined 
approaching such an attenuate form as they indicate ; and 
the latter because Major Connolly's measurement, if correct, 

' "E. farquhari, E. microthaiima, E. hypsoma, E. peri- 
ploca, E. oppugnans, E. berthse, E labyrinthea." 



44 



HENRY CLIFDEN BURNUP. 



removes from the record dimensions that are suspiciously 
under- estimated. 

His measurements are as follows : 

E. farquhari M. & P. (type), height 2-80, width 1-50 mm. 

E. farquhari (type of ''labyrinthea"), height 2'Ob, 
width 1'20 mm. 

Both of these shells were collected at Grahamstown. 

The dimensions in the following table are selected from 
those of over forty specimens that I have measured, from the 
various localities in Avhich the species in its typical form is 
known to live : 



Locality. 


Largest. 


Intermediate. 


Smallest. 


Graham stown 

Highlands, about 19 miles N. 

of Grahamstown 
Bathiirst, about 25 miles 

S.E. of Grahamstown 


mm. 

2-35 X 107 
2-14 X 105 
3-19 X 1-36 


mm. 

2-20 X 1-04 
206x1-04 
2-63 X 1-27 


mm. 

1-89 X 0-99 
1-94 X 100 

2 04x101 

1 



As these measurements are taken from the largest, the 
smallest, and an intermediate specimen from each locality, 
they indicate very fully the divergence of size and proportion 
of width to height to which the form is subject, the greatest 
development being reached at Bathurst. 

Var. bertha? {Melv. & Pons.). PL IV, figs. 25, 26, 27. 

Ennea berthas Melv. & Pons., Ann. & Mag. Nat. Hist., vol. viii (1901), 

p. 315, pi. ii, fig. 1. 
Ennea berthse Melv. £• Pons.; Connolly, Ann. S. Af. Mus., vol. xi 

(1912), p. 67. 

Shell like E. farquhari, typical, but elliptical in contour, 
with six, less ventricose, whorls and shallower suture ; the 
striae are rather less regular, shorter, and a little further 
apart ; the armature of the aperture is very similar, but the 
labral tooth is not quite so massive, and is less divided, the 
internal columellar plait is somewhat broader, and the short. 



ON SOUTH AFRICAN ENNE^. 45 

bi'oad tooth on the cohimellar lip is even less conspicuous 
than in the typical form. 

Height 2-84, width 1-50 mm. 

Hab. — Karkloof (McBean & Taynton) ; Nottingham Road 
(Taynton) ; and Ntimbankulu, Mid-Illovo (Burnup), all in 
Natal. 

The dimensions given above are those of a co-type of 
"E. berthas M. & P.," from the original lot collected by Mr. 
McBean at Karkloof^ now in my collection at the Natal 
Museum (Mus. No. 1441), and here figured. Those of the 
type in the original description are manifestly wrong, for 
they give a ratio of width to height of 35"71 per cent., while 
the figure represents a ratio of 50*98 per cent., the latter 
being Avell supported by my shell and new figure with a ratio 
of 52" 11 per cent. It will be seen that if Melvill and Pon- 
sonby's height dimension of 3"5 mm. be a misprint for 2'5, 
and the width measurement of 1'25 mm. be correct, a ratio of 
50 per cent, is established, corresponding very satisfactorily 
with their figure ; but in this case their shell is rather smaller 
than any that I have examined. The suggestion of a mis- 
print, however, is supported by the fact of the dimension line 
alongside their figure being 2"6 mm. long. 

The other known specimens of this variety, excepting the 
two in McBean^s collection, whose dimensions I have not 
been able to ascertain, measure as follows : 

Karkloof: height x width, 3-11 x 1-50,2-85 x 1-55,2-79 
X 1*44 mm. Nottingham Road : 3-25 x 1-52,2-86 x 1-50 mm. 
Ntimbankulu: 3-32 x 1-59, 3-13 x 1-57 mm. 

Although Messrs. Smith and Robson concur in my view that 
this form belongs to the species f arquhari Melv. & Pons., 
they have expressed no opinion as to its claim to varietal 
distinction. I think, however, that such a distinction is not 
only desirable, but inevitable, since it can be distinguished 
from the typical form by the characters above enumerated, 
and inhabits a different district, the variety having only been 
met Avith in the Province of Natal, and the typical form in 
the Province of the Cape of Good Hope. In three of the 



46 HENRY CLIFDEN BURNUP. 

eiglit shells examined the distinction is not quite so fvilly 
maintained as in the other five, as the shells are long-er with- 
out material corresponding* increase in width, and the form 
therefore is less elliptical ; but the other distinctions remain. 
The number of whorls increases with the additional height to 
6^, my largest specimen of E. farquhari, typical, having 
5| whorls. 

Var. avena n. PI. lY, figs. 28, 29, 30, 81. 

Shell small, cylindrical, with rounded apex, rimate and 
most narrowly perforate, thin vitreous, whitish, transparent 
and shining : whorls 6 j, only slightly convex, and, excepting 
for a few irregularly scattered faint transverse scratches, 
smooth all over, except the last half whorl, which is sculptured 
with regular distinct transverse rib-strias continued round the 
base ; suture rather shallow ; aperture somewhat broadly 
triangularly auriform, with well thickened slightly expanded 
and reflexed white porcellanous peristome, and furnished with 
the following processes : a prominent blade-like, in-running 
parietal plait, a massive, rounded, undivided labral tooth, a 
small internal basal tooth, a most inconspicuous, broad, very 
slightly raised tooth on, and parallel to, the columellar lip, 
and a deep-seated, rounded, flatfish, scoop-shaped columellar 
plait. 

Height 3-21, width 1-30 mm. 

Hab. — Maritzburg; also Pinetown and Durban (Burnup), 
Nottingham Road (Taynton), all in Natal. 

The axial perforation rises from a position so deep in the 
rima that it is difficult to detect, but the branch passing into 
the deep-seated columellar plait is more easily seen : depres- 
sions behind the lips correspond with the labral, basal, and 
columellar teeth. 

This variety differs from E. farquhari, typical, in its 
generally greater size, its more cylindrical form with nearly 
parallel sides, less ventricose whorls, shallower suture, absence 
of defined sculpture except on the last half whorl, less 
expanded yet more thickened peristome, more open aperture 



ON SOUTH AFIUOAN ENNE^. 47 

with more superficial, undivided labral process, and columellar 
tooth less developed so showing more of the deep-seated colu- 
mellar plait behind it. From the variety berthse it is 
separable by its more slender, cylindrical form, narrower 
columellar lip with more open aperture and solid, nearly 
superficial labral tooth, and generally the absence of striation, 
or Avhen it is present above the last half -whorl, its extreme 
shortness. 

The twenty-five specimens collected in Durban, one of 
which is shown in fig. 31, and two of those found in Maritz- 
burg, are sculptured immediately below the suture with more 
or less regular, rather coarse, typical striation, which, how- 
ever, soon disappears, leaving the remainder of the whorls 
almost smooth, except the last half-whorl, Avhich is sculptured 
all over. With the presence of this infrasutural striation 
occurs a slight increase in width ; otherwise the shells agree 
in all respects Avitli the smoother shells from Maritzburg and 
those from Pinetown and Nottingham Road, 

There is considerable variation in the height of individuals 
of this variety, though not much in width unless accompanied 
by the variation in sculpture ; with an increase in height there 
seems to be invariably also an increase in the number of 
whorls : thus, the specimens that I have examined vary from 
2'74 mm. to 3*51 in height, and the number of whorls from 
5i to 7. 

The following dimensions show very fairly the extent to 
which the variety, so far as is yet known, differs in size and 
proportion in various individuals : 

The smoother form : 

Maritzburg : 3-32 x 1-27, 3*10 x 1-28, 2-77 x 1-31 mm. 

Pinetown : 3*41 x 1-33, 2-74 x 1-33 mm. 

Nottingham Road : 2-85 x 1*32 mm. 

The form with infrasutural sculpture : 

Durban : 3-51 x 1*45, 3*30 x 1-45 (fig. 31), 3-02 x 1-45 mm. 

Maritzburg : 3-37 x 1-45, 3*33 x 1-47 mm. 

These dimensions, which are chosen to include the most 
divergent forms of the shells examined, Avell show the re- 



"48 HENRY CLIFDEN BURNUP. 

markable stability in width of this variety. Among the 
smooth shells there is a difference in height, between the 
extremes, of "67 mm. ; but in width there is no more than "06 : 
among those with infrasutural striation there is, in height, a 
difference of "49 mm., and in width only '02. 

Ennea darglensis Melv. & Pons, f . typica. PI. IV, 

fig. 32. 

Ennea darglensis Melv. & Pons., Ann. and Mag. Nat. Hist., vol. i 

(1908), p. 130, pi. vii, fig. 1. 
Ennea darglensis Melv. & Pons. ; Connolly, Ann, S- Af. Mus., vol. 

xi (1912), p. 71. 

The original figure not showing clearly the complex 
character of the columellar plait and representing the labral 
tooth as rather too solid and the sculpture too sharply 
defined, the result possibly of over-magnification, a new 
figure is here offered. The magnification chosen, about 
lOf diameters, is the same as that adopted for the figures of 
the new variety hereunder described, so that a fairer 
comparison of the variety with the typical form can be 
made, using the new figure instead of the old. 

The sinuous, duplicate columellar plait is a conspicuous 
characteristic of the species. The upper branch is wholly 
internal with its point produced forward ; and the lower, and 
larger, is less deep-seated, its base arising near the edge of 
the columellar lip with its point curving inward. From the 
rimal cavity an axial perforation passes upwards, and, at a right 
angle thereto, a deep, narrow passage entei-s each branch of 
the columellar fold. The labral and basal processes are also 
indicated by depressions behind the peristome. The labral 
tooth is more distinctly bifid than the original figure suggests, 
the upper branch being longer, and the lower more internal. 

To the localities cited by Connolly, in his most useful 
" Revised Reference List of South African Non-marine 
Mollusca," ^ may be added Bulwer, near which village, in 

1 ' Annals of tlie South African Museum,' vol. xi (1912), pp. 89-306. 



ON SOUTH AFRICAN ENNEiE. 49 

tlie District of Polela, Mr. C. W. Alexander has collected a 
single specimen Avhile these notes were being written. It 
is a good deal smaller than any of the specimens from the 
other localities, but in form is exactly proportionate to the 
shell here figured, and is identical in all detail. 

The locality Gowie's Kloof, Grahamstown, cited in the 
same place, is erroneous, and should be expunged. The 
shell on which the locality was cited is not darglensis, but 
is Ennea ponsonbyi n.sp., described hereafter, p. 78. 

The following are the dimensions of such shells as I have 
measured, besides the type : 

From Inhluzani: height X width, 2-69 x 1-25, 2-62 xM8, 
2-54 X MS, 2-53 x M9 (tig. 32), 2-46 x 1-12. From Bulwer : 
2-23 X 1-07 mm. 

The first and second on the list are picked, as appearing to 
be the largest, from a series of about seventy specimens. 

Var. illovoensis ii. PI. IV, figs. 33, 34, 35. 

Shell small, rimate and narrowly perforate, elongate ovate- 
cylindrical, thin, glossy, transparent, whorls 6j, rather 
convex, almost smooth except the last half-whorl and the 
base which are regularly rib-striate; suture rather shallow; 
aperture rather quadrate and somewhat oblique, with white, 
thickened and reflexed peristome armed with the following 
plaits and teeth : A broad blade-like, squarish parietal 
plait, a deeply cleft labral tooth, a small, sharp in-running 
basal tooth-like plait, and a complicated two-pointed colu- 
mellar plait with detail as in darglensis, typical. 

Height 2-92, width 1-37 mm. 

Hab. — Ntimbankulu, Mid-lllovo (Burnup). 

In form and arrangement of the armature of the aperture, 
this variety almost coincides with the type. The upper 
branch of the labral tooth in the variety is squarish at the 
end instead of pointed, and the complex columellar process is 
situate a little lower; but, though these features seem constant 
in the specimens examined, they are but trifling divergencies 
such as might be looked for in individuals of the same form. 
VOL. 3, PART 1 . 4 



50 HENEY CLIFDKN BUENUP. 

There are, therefore, only the superior size and the smooth 
polished surface to distinguish the variety from the typical 
form; but these are sufficiently marked to warrant, in my 
opinion, the assigning of a varietal name to the form from 
Mid-Illovo, even if intermediates be eventually found. 

I have examined a great number of each form and find 
them very constant. The dimensions of other specimens 
from the same locality, that I have measured, are as follows : 
height X width, 3-07 x 1-40, 2-87 x 1-35, 2-83 x 1-34, 
2-80 x 1-34, 2-72 x 1-38 mm. 

In most specimens there is the appearance of a " margined 
suture " : but I look upon this as an optical illusion rather 
than a shell character, not only in this species, but in most 
South African Enneee where it occurs. The lower sutural 
line cannot be felt with the point of a fine needle, and is, 
without doubt, merely the base of the previous whorl seen • 
through the transparent shell. As the shell becomes calcined 
the illusory line vanishes. 

Ennea consobrina Anceij. PI. TV, fig. 36. 

Ennea consobrina ^Hcey, Brit. Nat. (1892), p. 125; Melv. & Pons., 
Ann. and Mag. Nat. Hist., vol. i (1898), p. 24, pi. viii, fig. 9. 

Through the courtesy of Mr. J. R. le B. Tomlin, in entrust- 
ing to me for examination the type-specimen from his collec- 
tion, I have been enabled to identify further examples of this 
little-known species, collected by Mr. Farquhar at Martindale, 
Bathurst, a division of the Province of the Cape of Good Hope, 
adjoining that of Albany, in which the type was found, I 
have also taken the opportunity of having the type tem- 
porarily in my possession to re-figure the shell, it not having 
been figured in the first instance, and the figure published by 
Melvill and Ponsonby in 1898, though bearing some general 
likeness to the shell, not showing enough detail for purposes 
of comparison. Ancey rightly compares his species with 
E. thelodonta M. & P., but, while stating that the 
denticles in both species . are similar, though somewhat 



ON SOUTH AFEICAN ENNEiE. 51' 

different, does not specify in what respects they differ. 
Melvill and Ponsonby's figure of Ennea thelodonta^ is 
confusing, showing at the same time, to a great extent, a side 
view of the spire, and a front view of the apertvire ; besides, 
the detail is faulty. I therefore present a new figure of their 
species (fig. 37), for comparison with that of consobrina. 
It will be seen from these that the likeness between the two 
species, though striking, is only superficial, the detail of the 
labral tooth and columellar plait in each being of quite a 
distinct character. In consobrina the large labral pi'ocess 
bears on the inner edge two distinct, little in-running plaits : 
in thelodonta a somewhat similar, but smaller, process is 
merely slightly divided into two lobes. In consobrina the 
deep-seated columellar plait is strengthened by a transverse 
rib; in thelodonta it is merely drawn forward at the lower, 
inner corner, into a rounded point. In neither case does it 
seem likely that the difference between these processes is the 
result of the direct evolution of one form from the other, for 
they seem to be constructed on different plans; therefoi-e I 
consider the two species quite distinct. In addition, there is, 
as pointed out by Ancey, the difference in size and number 
of whorls, which is very considerable, though probably not 
quite so great as would appear from a comparison of the 
descriptions; for, while Ancey describes his shell as 8 mm. 
high and 4 mm. wide, or, as he says, twice as large as thelo- 
donta, Avhich is described as 4 mm. high and 2 mm. wide, 
my measurement of Ancey^s type makes it 8 mm. high and 
3*7 mm. wide, while my two specimens of thelodonta 
measure as follows : 

Height X width, 4-87 x '2-26, 4-25 x 2-15 mm. (fig. 37). 

These specimens were collected at Port Elizabeth by Mr. 
-T. Farquhar. 

Other specimeus of E. consobrina examined by me 
measure as hereunder : 

In my collection at the Natal Museum, 7*75 x 4'0 ; in 
collection of J. H. Ponsonby, 7-75 x 3-67, 7-75 x 3-67 mm. 

' ' Ann. and Mag. Nat. Hist..' vol. ix (1892). pi. vi. fig. 4. 



52 HENRY CLIFDEN BUENUP. 

The dimensions of the remaining specimen in Mr.Ponsonby's 
collection were not taken, as, the lip at the base being chipped, 
they would be misleading. 

As regards the difference in the number of whorls, while 
Ancey describes his type as having 8 whorls, I can but count 
7f ; and while Melvill and Ponsonby assign 6 as the nvimber of 
whorls in their type, their figure seems to indicate about 6^ 
or 6J, and my specimens have 6| and 7 whorls respectively. 

Ennea muni t a Melv. & Pons. PL IV, fig. 38. 

Ennea munita Melv. & Pons., Ann. and Mag. Nat. Hist., vol. ix (1892), 
p. 86, pi. vi, fig. 5. 

Shell small, rimate, ovate-cylindrical, whitish, transparent, 
shining, rounded towards the blunt apex ; whorls about 7, 
only slightly ventricose, rather strongly sculptured with 
regular transverse strife, except the first two, which are 
smooth, and the area immediately above the aperture, which 
is nearly smooth and brightly polished ; suture rather shallow ; 
aperture rather long and ear-shaped, with thickened, much 
expanded, and slightly reflexed, white porcellanous peristome 
furnished with the following teeth and plates : a comparatively 
small, blade-like, in-running parietal plait, a massive, rounded 
tooth on the labrum bearing a denticle on the upper edge, a 
small, deep-seated basal tooth, a rather broad, short, expressed 
tooth, tapering below and ending abruptly above, on the 
columellar lip, and a very deep-seated, rounded mammillate 
columellar plait. 

Height 3-58, width 1-86 mm. 

Hab. — Grriqualand East; also Kowie (Ponsonby and 
Langley). 

The original description being rather deficient and the 
original figure indistinct and misleading, I have described 
and figured a specimen in my collection given to me by Mr. 
Ponsonby. Major Connolly, who has kindly compared for 
me the figure with the type in the British Museum, writes 
that the only differences he could find between the type and 



ON SOUTH AFRICAN ENNE.l^. 53 

my figure are that the coluinellar lip of the former is of more 
uniform breadth than that of the figure, and the prominence 
on the deep-set columellar plait is not situate quite so high. 

There is a certain resemblance between this species and 
Ennea crassidens Pfr., but the greater size, the ovoid 
form and conical apex, and the shortness and squareness of 
the labral tooth of the latter are enough to distinguish it at a 
glance. It also seems to bear some resemblance to Ennea 
tharfieldensis, M. & P., a species unknown to me 
except from description and figure. Connolly has kindly 
compared the two types in the British Museum, and con- 
siders the latter sufficiently distinct through the presence 
of a denticle at the junction of the columella and the 
paries, should this feature prove to be constant. The colour 
attributed to the type in the original description probably 
arises through the remains of the animal being seen through 
the transparent shell. The upper whorls of my shell have 
an ochraceous tint from the same cause. 

From the rima a deep narrow channel passes into the deep- 
seated columellar plait, but no axial perforation can be 
distinguished. The processes on the outer and the columellar 
lips and at the base have corresponding depressions on the 
outside of the shell. 

Besides the shell figured I have been privileged to examine 
tAvo specimens in Mr. Ponsonby's collection and three in 
Mr. Langley's, all from Kowie. They are all much larger 
than the specimens from Clriqualand East, but exhibit no 
important variation. A slight difference in the develop- 
ment of the peristome and its processes may be observed, 
but not to any material extent. 

The dimensions of the specimens from Kowie are as 
follows : 

Height X width, 4-94 x 2-18, 4-94 x 2-13, 4-84 x 2-29, 
4-40 X 2-15, 4-34 x 2-14 mm. 



54 HENRY CLIFDEN BURNUP. 

Ennea sylvia Melv. & Pons. PI. Ill, figs. 18, 19, 20. 

Ennea sylvia Melv. & Pons., Ann. and Mag. Nat. Hist., vol. xii (1903). 
p. 599, pi. xxxi, fig. 4; Connolly, Ann. S. Af. Mus., vol. xi (1912), 
p. 86. 

Another species, hereafter described as E. melvilli, from 
Natal, having been mistaken for, and distributed as, E. sylvia 
M. & P., it has become necessary to examine critically the 
characters, description, and figure of the older shell before 
describing the new, and to re-figure the former with a magni- 
fication corresponding to that chosen for the figures of the 
latter, in order to make comparison easier. For this purpose 
I have been able to examine the four co-types in Mr. 
Ponsonby's collection and three shells from the original lot, 
presented to me by Mr. Farquhar ; and from the latter I have 
chosen my type for the new figure, the co-types being unsuited 
to the purpose in that one has a broken peristome, one is 
holed, one is abnormally large, and the fourth has a slight 
extraneous obstruction within the aperture which I failed to 
remove. Though unsuitable for figuring, their characters are 
quite sufficiently revealed to leave no room for doubt that my 
other three specimens belong to the same species. 

In the original description a contradiction occurs in that 
the teeth and plaits are said to be four, while the details of 
five are given. Five is the correct number, of which the 
deep-seated internal columellar plait is more than mammae- 
form, having a strong rib across it, obliquely from left to 
riffht and downwards. The small basal " tooth " is a small 
in-running plait, and that on the columellar lip appears like a 
triangular swelling, but is well excavated from behind. From 
the rima a very narrow perforation rises up the axis Avith a 
wider branch into the deep-seated columellar plait. Besides 
the processes of the aperture detailed in the original descrip- 
tion there is a very inconspicuous thickening of the shell on 
the base forming a slightly raised ridge, hardly amounting to 
a plait, parallel to the basal lip and behind the basal and 
labral processes. A slight corrugation indicates its position 



ON SOUTH AFRICAN ENNE^. 55 

externally, and the outer wall bears a double pit corresponding 
to the bifid labral plait. 

If the dimensions of the type as given in the original 
description, viz. height 1'75, width 0-75 mm., are correct^ 
they cannot be considered normal in view of the measure- 
ments of the remaining available specimens, which are as 
follows : 

Co-types in collection of J. H. Ponsonby : height x width, 
2*58 X r06, 2-26 x TOl. Specimens in my collection : 2*25 x 
1-06, 2-25 X 1-01, 2-06 x 0-96 mm. (figs. 18-20). 

I regard the first of these as being abnormally high. 



Ennea melvilli ii. sp. PL III, figs. 21, 22, 23. 

Shell minute, elongate-oval, rimate, thin, whitish, glossy, 
transparent, apex rounded ; whorls 6, rather ventricose, very 
closely, finely sculptured with delicate transverse rib-striae, 
except the first two, which are smooth ; suture moderately 
deep ; aperture very small, rather roundly triangular, with 
white peristome slightly thickened and reflexed and joined by 
a narrow callus, and furnished with the following plaits and 
teeth; a broad blade-like parietal plait, a squarish, bifid tooth 
on the outer lip, a small basal in-running plait, a broad, 
slightly raised, rounded tooth on the columellar lip, a deep- 
seated, rounded, scoop-shaped plait on the columella, and a 
slightly raised ridge within, and nearly parallel to, the basal 
lip. 

Height 2-12, width 0-91 mm. 

Hab. — Nottingham Road (Taynton) ; also Karkloof 
(McBean), Curry's Post and Fort Nottingham (Taynton), 
Dargle and Edendale (Burnup). 

The parietal plait is very prominent, the right edge standing 
well forward from the general plane of the peristome before 
entering the aperture ; its left edge is shorter and more 
internal. The bifid labral tooth arises very near the edge of 
the peristome, the lower half being rather more internal. The 



56 HENRY CLIFDEN BUBNUP. 

small basal tooth-like plait arises but little within the edge of 
the peristome at the base of the columellar lip and curves 
inwards and upwards slightly to the left. The process on the 
columellar lip, which I have above called " a broad, slightly 
raised, rounded tooth/' consists of an inward bend of the lip 
covered by its outward expansion, and is quite superficial. 
The deep-seated columellar fold, which is wholly internal, is 
rounded in outline, scoop-shaped, and presents its right, lower 
edge furthest forward. The ridge across the base is very 
inconspicuous, and might easily be overlooked ; its position, 
however, can be traced on the outside by a slight corrugation. 
The positions of the other processes, except the parietal, are 
also indicated externally by corresponding depressions. The 
peristome, which is comparatively but little thickened, is 
exceedingly sinuous on the labrum (fig. 22). From the rima 
there is a deep opening passing into the columellar fold, but 
none can be detected into the axial perforation. 

Besides those of the type the following dimensions of 
normal specimens have been taken : Dargle : height x width, 
2-12 X 0-88, 2-10 X 0-88, 2-05 x 0-89. Nottingham Road: 
2-04 X 0-89, 2-07 x 0-89, 2-00 x 0-91, 1-94 x 0-89 mm., and 
the following are the dimensions of the largest and smallest 
examples in my collection, both manifestly abnormalities : 
Nottingham Road, 2-60 x 0-91 ; Dargle, 1-83 x 0-93 mm. 

This shell has been confused with B. sylvia M. & P., but 
though the armature of the aperture is arranged much on the 
same plan, only one plait differing very materially, this fact, 
I think, must be taken as an indication of parallel develop- 
ment rather than close affinity, for in all other important 
respects the two species vary widely. The form of this 
species is elongate-oval or fusiform; that of sylvia cylin- 
drical, with almost parallel sides. In sylvia there is an 
umbilical opening from the rima, fairly wide for so small a 
shell; in mel villi it seems to be closed. Mel villi has 
rather more than half a whorl in excess of sylvia, the whorls 
of the former being more ventricose and separated by deeper 
sutures : with the exception of the embryonal whorls, they 



ON SOUTH AFRICAN ENNE^. 57 

are most finely, delicately, closely and regularly rib-striate ; 
while Sylvia is nearly smooth, except for a few irregularly 
dispersed striae, all over except on the last half -whorl, where 
the stride become more regular and more clearly defined. The 
aperture is smaller, and the peristome much less thickened 
and expanded in m el villi than in sylvia, and the parietal 
plait is larger and more pi'ominent, the labral tooth smaller 
and less distinctly divided, the " tooth " or swelling on the 
columellar lip is less angular, and the internal columellar fold 
is of quite a different nature, being simple instead of crossed 
by an oblique strengthening rib as in sylvia. The profile of 
the outer lip, too, is much more sinuous in mel villi. 

The specimen with the abnormal height of 2"6 mm. is no 
wider than the type, has fully 7 whorls and has the peristome 
scarcely thickened, leaving the sAvelling of the columellar lip 
undeveloped ; the internal columellar fold is small. 

Ennea columnella 31. & P. (PI. Y, fig. 47), is found in 
company with this species at Karkloof, Dargle, and Edendale, 
and, both being much of the same size, a critical comparison 
may here be convenient. 

The form of columnella is rather wider and less fusiform; 
the surface is almost smooth, except immediately below the 
suture, where irregular microscopic striation may be detected 
in a few places, and on the last half-whorl where somewhat 
regular striation appears ; the parietal plait is narrower, the 
labral tooth more widely bifurcate, the basal tooth smaller 
and more central, the slight ridge across the base of mel villi 
is absent, and the columellar plait is of quite different con- 
struction (see postea, p. 58). With the assistance of a 
moderately powerful hand-lens their separation is quite easy. 

I have pleasure in dedicating the species to Dr. J. Cosmo 
Melvill, who has done so much towards the elucidation of 
the South African non -marine molluscan fauna. 

This is the form referred to by Connolly ^ as examples 
from Natal, distinct from E. sylvia, whose habitat is Mae- 
strom Forest, Bedford, Cape of Good Hope. 

• Ibid., p. 86. 



58 HENRY CLIFDEN BUENUP. 

Ennea column ella Melv. & Pons., f . typica. PI. V, 

fig. 47. 

Ennea columnella Melv. & Pons., Ann. and Mag. Nat. Hist., vol. viii 
(1901), p. 316, pi. ii, fig. 2 ; Connolly, Ann. S. Af. Mus., vol. xi 
(1912), p. 69. 

In the original figure the parietal plait appears to be 
wholly internal, which is quite wrong, as its right side arises 
above, and in front of, the aperture. Hence a new figure 
has become necessary in order that comparison with kindred 
forms may be facilitated. In the original description the true 
nature of the very distinctive columellar plait is not very 
clearly set forth. It is not, as is usual in the South African 
Enne«, wholly internal; but, from the lower inner edge of 
a flattish, somewhat scoop-shaped internal surface, a 
strengthening rib curves outwards almost to the outer edge of 
the labium. By this feature the species can be readily dis- 
tinguished from other species of similar size and superficial 
likeness. 

I have not found the peristomatal processes of the many 
examples that I have examined exhibit among themselves con- 
siderable variation, as Melvill and Ponsonby did in the 
original lot ; but, as one of my " co-types " belongs to a 
different species, described in this paper as Ennea mel villi, 
the divergence from the type that they observed seems easily 
explained. 

The figure here presented is drawn from a co-type in my 
collection from the original lot found by McBean in the Kar- 
kloof Bush. The shell measures as follows : Height 2*15 mm., 
width POO mm., and is the lai-gest that I have examined. 

The following measurements of shells collected by me at 
Dargle have also been taken : 

Height X width: 2-08 x 0-89, 2-06 x 0-98, P99 x 0*92, 
1-99 x 0-90, 1-98 X 090, P97 x 0-94, P87 x 0'92 mm. 

The original width dimension of '75 mm. is probably wrong, 
as the original figure, taking the height as 2 mm., represents 
the width to be 1 mm., and Connolly^s measurement of the 
type as communicated in lit., gives height 2, width 0'9 mm. 



ON SOUTH AFRICAN ENNE^. 59 

Since the above was written, my attention lias fallen upon 
two shells collected by me in the year 1900 in the Beach Bush 
at Lower Umkoraaas, which could not be identified at the 
time but undoubtedly belong to this species. Their dimen- 
sions are as follows : 

Height X width: 1-94 x 0-79, 1-82 x 0-83 mm. 

Though among the smallest measured, their ratio of width 
to height corresponds with the typical foi-m rather than with 
the more obese variety vitreola, considered below. In fact, 
the higher of the two is the most attenuate of all. 

This newly cited locality is interesting, as but few species, 
at least in typical form, are found both on the coast and in 
the higher altitudes of the midlands. 

Var. vitreola [Melv. & Pons). 

Ennea vitreola Melv. & Pons., Ann. and Mag. Nat. Hist., vol. i 
(1908), p. 130, pi. vii, fig. 3 ; Coimolly, Ann. S. Af . Mus., vol. xi 
(1912), p. 87. 

I cannot separate this form from the preceding, except on 
the grounds of its more ovoid contour with shallower sutures, 
and the slightly weaker development of the peristome and 
peristomatal processes. The general appearance of the shells 
in other respects, their sculpture, and the arrangement of the 
peristomatal processes, are almost identical. The type having 
been reported lost, I sent the better of the co-types to take 
its place; and this, I believe, is the specimen now in the 
British Museum. The remaining co-type is in my collection 
at the Natal Museum.^ 

Although the exact locality at Hilton Road has been dili- 
gently searched several times since the original shells were 
discovered, no further examples of the variety, and no 
specimen of the typical form at all, have been found until the 
present time (November, 1913), when, a final effort being- 
made, one specimen was obtained confirming the varietal 

' Connolly's statement, in loc. cit., that the type is in Maritzburg, 
is incorrect. 



60 • HENRY CLIFDEN BDRNUP. 

features to a marked extent, it being even shorter than any 
of the others, with a greater relative width. 

In my co-type of this variety and in the newly found 
specimen the two parts of the labral tooth are more distant 
than the original figure shows to have been the case in the 
type^ so bringing them slightly nearer to the typical form of 
E. columnella. The strengthening rib connecting the 
columellar plait with the columellar lip is rather more tor- 
tuous than in the type (of vitreola), but I do not consider 
that an important feature. 

The following are the dimensions of the only four specimens 
known : 

Height. Width. 

Type of E. vitreola (lost) . . 1-87 . 0-94 mm. 

Co-type in British Museum . . 1'82 . 0"92 „ 

„ „ Natal Museum . . I'Sl . 0-93 ,, 

Specimen collected lOthNov., 1913 1-76 . 0-92 „ 

The ratio of width to height in above varies from 50*27 to 
52*27 per cent., while that of the measured specimens of 
columnella, typical, from Dargle, varies from 42*31 to 49*20 
per cent., and that of the higher shell from Lower Umkomaas 
is 40*72 per cent. 

I am indebted to Messrs. E. A. Smith, Connolly, and Preston 
for their views on the relationship of these two forms, 
E. columnella, typical, and var. vitreola, arrived at 
during a discussion over the type of the former and co-type 
of the latter in the British Museum. It is at their suggestion, 
in which I now fully concur, that I have retained the distinction 
of the Hilton Road form under the name vitreola used in a 
varietal sense. 

Although it will be seen that one of the Lower Umkomaas 
specimens of E. columnella is even shorter than the type 
of vitreola, it agrees in every respect except size with 
columnella, typical, its relative proportions being about 
equal to an average example of the Dargle specimens. The 
most stunted example from Dargle approaches the proportions 
of the variety vitreola, but can readily be distinguished 



ON SOUTH AFRICAN ENNE.E. 61 

by its more cylindrical, less ovoid, form, and its deeper 
suture . 

Ennea marine Melv. & Pons. PL V, fig. 48. 

Ennea marise Melv. & Pons., Ann. and Mag. Nat. Hist., vol. ix (1892), 

p. 92, pi. vi, fig. 12. 
Ennea callista Melv. S: Pons., ibid., vol. iv (1909), p. 486, pi. viii. 

fig. 1. 

As neither the original figure of E. maria^ nor that 
published with the description of "E. callista" is di-awn 
in true proportion, or expi"esses clearly the characters of the 
species, I take this opportunity of publishing a new figure. 
For this purpose I have been fortunate enough to secure the 
loan from Mr. Ponsonby of one of the original specimens 
collected by Miss M. Bowker at Somerset East. I have also 
examined one specimen, collected by the late Mr. Crawford, 
in Mr. Ponsonby's collection, three specimens (co-types of 
"E. callista") in Mr. Ponsonby^s collection, and three 
specimens collected by Mr. Farquhar at the same time as 
the type of "E. callista," two of which are in his collection 
and one in mine. All these specimens were found at Somerset 
East, the locality, " Dassy Krantz, Grahamstown," given 
in the description of ^'E. callista," being an error arising 
through the interchange of the contents of two boxes. Thus 
Somerset East remains the only known locality for E . mari^. 
The other shells affected by the interchange were those 
described as "E. periploca"; so Somerset East cannot 
be accepted as a locality for E. farquhari, of which 
"periploca" is a synonym, on our present knowledge 
of its distribution. 

The tooth on the columellar lip is not connected with the 
internal columellar plait, as stated in the original description 
of E. m aria?, but is quite distinct as defined in the description 
of "E. callista." A narrow axial perforation rises from 
the umbilical slit, a branch entering the columellar fold; 
and the three peristomatal processes have corresponding 
depressions behind the lips. 




62 ■ HENEY CLIFDEN BURNUP. 

The dimensions of the various specimens examined are as 

follows : 

Height. Width. 
mm. mm. 

Type of new figure (fig. 48), Miss M. Bowker 3-03 . 1'19 

1-18 

M7 

1-17 

1-17 

1-24 

1-13 

1-09 



Mr. Ci'awford's specimen (in coll. Ponsonby) 2'78 

f 2-79 
Co-types of "callista" (in coll. Ponsonby) . j 2*71 

I 2-71 

Original specimens of "callista" (in coll. f 2*70 

Farquhar). \ 2-70 

Original specimen of "callista'^ (in Natal 2*55 

Museum) . 

■ My examination of these well-authenticated examples 
leaves no room for doubt that they belong to one species; 
indeed, there is little individual variation among them. 
"E. callista'^ must, therefore, become a synonym of E. 
marife. I am indebted to Major M. Connolly, who has 
kindly examined the types in the British Museum for me, 
for the following confirmation of my views: "In E. mar if© 
the trace of a cusp on the upper part of the labral tooth is 
slightly more prominent than in E. callista. The rest of 
the dentition, together with the form and sculpture, is similar 
in both types, and the shells are undoubtedly conspecific." 

He also furnishes me with the following revised dimensions 
of the types as taken by him : 

"Type of E. marine, height 3*00, width 1*35 mm. 
„ „ E. callista, height 2-80, width 1-30 mm." 

Ennea mooiensis n. sp. PI. V, figs. 49, 50, 51. 
Shell minute, elongate-elliptical, deeply rimate, thin, glossy, 
transparent, apex blunt ; whorls 5|, rather ventricose, almost 
smooth except immediately behind the peristome where a few 
transverse strise appear ; suture not deeply impressed ; aper- 
ture roundly triangular, with white peristome thickened 
and reflexed, the ends joined by a thin, narrow callus, and 
furnished with the following processes : a broad, blade-like, 



J 



ON SOUTH AFKIGAN ENNE^, 68 

in-running parietal plait, a sub-central moderately large 
labral tootli slightly bifid, a small pointed tooth near the 
base of the columellar lip, a flattish scoop-shaped internal 
columellar plait whose lower right edge advances forward 
ahnost as a mammillate point, and a slightly raised ridge 
Avitliin the basal lip extending from behind the labral tooth 
downwards and forwards towards the base of the columellar 
lip. 

Height 2'55, width 1'24 mm. 

Hab. — Game Pass, Upper Mooi River, Natal (Burnup). 

The details of the several processes as set forth under the 
description of E. mel villi, herein, Avould apply almost equally 
well to those of this species, but the following differences 
exist : the tooth near the base of the columellar lip is smaller, 
less erect, and less plait-like in form in this species ; the 
broad, slightly raised, rounded tooth higher on the columellar 
lip of melvilli is here absent; and the ridge on the basal 
wall is less parallel with the peristome. Although so much 
of the description of the one species applies to the other, the 
greater dimensions, especially in width, the less fusiform 
contour, the larger aperture and the smoothness of the surface 
of this species distinguish it very readily from melvilli. 
Th^ labral tooth and that near the base of the columellar lip 
have corresponding depressions on the outside, but the ridge 
on the basal wall does not seem to be excavated, though its 
position within can be traced on the outside by a white scar. 
The opening from the rima to the columellar plait is exceed- 
ingly narrow, while that to the axial perforation is even 
narrower, only being discerned by the aid of a very strong 
lens. 

This species may also be compared with E. columnella 
If. & P., whose peculiar columellar plait, however, will always 
readily distinguish it ; besides, the latter is smaller, has 
traces of minute striae below the suture, and much more 
numerous and stronger stride on the last half-whorl, has the 
labral tooth much more widely bifid, and the basal tooth, or 
tooth-like plait, much more central and in-running. 



64 HENRY CLIb'DEM BURNUP. 

The figures of E. sylvia M.&P. (PI. Ill, figs. 18, 19, 20) and 
the comparison between tliat species and E. melvilli (p. 56) 
may here be referred to, for there is also some general like- 
ness between the former species and E. mooiensis, which I 
think an examination of the figures and a perusal of the text, 
above indicated, will show to be only superficial. 

The greater height and narrower, more cylindrical form of 
E. mariae Melv. & Pons., and the prominent tooth situate 
subcentrally on its columellar lip, prevent the possibility of it 
being confused with this species. 

The following are the dimensions of a few specimens of 
E. mooiensis which, in addition to the type, I have mea- 
sured : height X width, 2-65 X 1-25, 2-65 x M6, 2-55 x 1-21, 
2-43 X 1-19, 2-33 x M8, 2-25 x M2, 2-21 x 1-19, 2-18 x 1-18 
mm. 

This species is plentiful in its locality, almost to the exclu- 
sion of other species of the genus, but has not yet been found 
elsewhere. Whilst visiting the district twice, during the 
Christmas holidays 1910-1911 and 1911-1912, I collected 
224 specimens of Enne«, 220 of which belong to this species 
and 4, all distinct inter se, to different, larger species not 
yet identified. 

Ennea maritzburgensis Melv. & Pons f . typica. PI. IV, 
figs. 39, 40, 41, 45, 46. 

Ennea maritzburgensis Melv. & Pons., Ann. and Mag. Nat. Hist., 
vol. xii (1893), p. 107, pi. iii, fig. 11. 

The original description seeming to require some slight 
revision, and the type being unavailable to me, I have chosen 
a normal specimen from the original locality as my type of 
the following emended description and accompanying new 
figures (PL IV, figs. 39, 40, 41). 

Shell small, rimate and narrowly perforate, thin, whitish, 
subdiaphanous, shining ; whorls 7, slightly convex, of which 
the first 2, constituting the protoconch, are smooth all over, 
the next 4^ are delicately, transversely striate immediately 



ON SOUTH AFRICAN ENNExE. 65 

below the suture, and smooth beyond, except for a few irregular 
striae occasionally extending lower, or even passing from 
suture to suture, and the last half-whorl is regularly, similarly 
striate all over ; aperture ovate ; peristome white and shining, 
expanded and reflexed, and furnished with four processes, as 
follows : a moderately large, squarish, blade-like, in-running 
parietal plait, a large labral tooth Avidely bifurcate, the lower 
branch being much the larger, a small tooth near the base of 
the coluuiellar lip, and a columellar plait, wholly internal and 
bearing a conspicuous, strengthening transverse rib. 

Height 4-46, width 2-07 mm. 

Hab. — Maritzburg, Natal, typical ; and, aberrant, Notting- 
ham Road (Taynton) and Rosetta (Hickey, per Ponsonby). 

The axial perforation is very narrow and is easily over- 
looked ; from it a branch passes at right angles into the 
strengthening rib of the columellar plait. The process within 
the labrum, originally described as two simple teeth, is, I think, 
better considered as one bifurcate tooth, as the two points 
spring from a common base, which is indicated on the outside 
by a single depression. The bifurcations, being solid, appear 
in this cavity as two white porcellaneous spots. The basi- 
columellar tooth is barely excavated, and also appears as a 
white porcellaneous spot behind the peristome. 

The dimensions given in the original description seeming to 
indicate a shell of smaller size and much narrower form than 
this species usually assumes, and not agreeing with the pro- 
portions of the original figure, Mr. Robson has kindly re- 
measured the type very carefully for me, and finds that its 
dimensions are, height 3" 9, width 1*8 mm., almost exactly of 
the same relative proportions as the type of the new figure, 
but considerably smaller. On finding the emended dimen- 
sions so much smaller than those of any of my shells collected 
within recent years, I looked up my earliest examples, found 
about the same time as the type, and have picked out two 
nearly as small, measuring respectively 4"05 x 1*83 and 3*94 
X 1"95 mm. This difference in size between the old and 
the new specimens might suggest that recent seasons have 
VOL. 3, PART 1. 5 



6Q HENEY CLIFDEN BUBNUP. 

been more favourable for the growth of the shells, but it 
seems to me more likely to arise through the position in which 
they were collected. Since I have found the species plentifully 
under boxes of plants in the Alexandra Park, watered daily 
in dry weather, I have ceased to look for them in natural 
surroundings, where they are much less plentiful and their 
supply of moisture is restricted to the rain and dew. 

It is noticeable that, as far as is known, the typical form 
has not been found beyond the precincts of the borough of 
Maritzburg. 

The aberrant forms figured on PI. lY, figs. 45, 46, will be 
more conveniently discussed after the introduction of the 
following variety. 

Var. contracta n. PI. IV, figs. 42, 43, 44. 

Shell like E . m a r i t z b u r g e n s i s, typical, with the following 
differences : Shell much smaller, less fusiform and more ovate ; 
whorls 6.\, more finely and more obscurely sculptured; aper- 
ture rounder, proportionately much smaller, and rather less 
obstructed, with peristome much less expanded, and labral 
tooth smaller and less widely bifurcate. 

Height 3"32, width 1*58 mm. 

Hab. — Nottingham Road (A. J. Taynton), very plentiful ; 
also Karkloof and Curry's Post (Taynton), all in Natal. 

The pinched appearance of the relatively small aperture, 
taken in conjunction Avith the much reduced size of the shell, 
would, without further consideration, readily induce one to 
suppose that this form is quite distinct from E. m aritzbur- 
gensis 3Ielv. & Pons. ; but a close study shows both forms to 
be built upon exactly the same plan, such differences in detail 
as exist being merely a matter of degree. The sculpture is 
of the same nature, and the disposition of the apertural 
processes is identical, with the one exception of the labral 
tooth in contracta being less widely ■ cleft. In view of 
such important coincidence, I should have felt disposed, even 
if no further evidence were forthcoming, to consider the new 
form a variety of the older ; but with the support of the 



ON SOUTH AFRICAN ENNEJil. ^7 

aberrant forms from Karkloof and Rosetta, the close relation- 
ship seems manifest. I think the Nottingham Road form, 
which is also found at Karkloof and Curry's Post, showing the 
extreme divei'gence from the type and being well established 
in numbers, may conveniently be distinguished by a varietal 
name, contracta being chosen in reference to the pinched 
appearance of its aperture, while the aberrant forms remain 
undistinguished in name, simply as forms intermediate 
between m a r i t z b u r g e n s i s, typical, and its variety. These 
intermediates have not so far been found plentifully ; from 
Karkloof I have four specimens coi-responding with fig. 45, 
taken with twenty-seven specimens of var. contracta; and 
from Rosetta I have two specimens only, corresponding with 
fig. 46, without representatives of contracta. These, how- 
ever, are from a small parcel, of whose numbers I have no 
record, given to Mr. Ponsonby by Miss Hickey ; but it is 
justly presumable that they are a fair sample of the lot, and 
that var. contracta Avas not collected by Miss Hickey. 

The shell represented by fig. 45 is but little removed from 
m a r i t z b u r g e n s i s, typical, the peristome being only slightly 
less expanded, but the labral tooth is much smaller and its 
bifurcation much less distinct. The Rosetta form, fig. 46, on 
the other hand, is much nearer var. contracta, the shell 
being almost equally small and the peristome much narrowed ; 
but the aperture is relatively large, so avoiding the pinched 
appearance of the named variety. 

The dimensions of other specimens are as follow : 

Intermediate form from Karkloof : height x width, 4*47 X 
1-89, 4-31 X 1-95, 4-09 x 1-97 (fig. 45), 4-03 x 1-94 mm. 

Intermediate form from Rosetta : 3-68 x 1-76, 3-46 x 1-76 
mm. (fig. 46) . 

Var. contracta from Nottingham Road: 3"73 x 1*73, 
3-51 X 1-76, 3-38 x r66, 3-13 x 1-63 mm. 

Var. contracta from Karkloof: 3"60 x 1'66, 3'4() x 1*66, 
3-43 X 1-68 mm. 

A'ar. contracta from Curry's Post: 3-51 x 1"62, 3"40 x 
1"(J2 mm. 



68 



HENRY CLIFDEN BURNUP. 



In general appearance, without regard to detail, var. 
contracta is much like a diminutive E. obovata P/V., 
but the arrangement of the apertural armature is quite 
distinct. 



Ennea arnoldi Stm-any. Text-figs. 

Ennea arnoldi Sturatiy. Anz. k. Akad. Wissensch. Wieu (1898), No. 
xvi, p. 158 (reprint p. 6) ; Stuvany, Siidafrik. Moll. (1898), p. 28, 
pi. ii, figs. 26-30; Connolly, Ann. S. Af. Mns., vol. xi (1912), p. 67. 

From Miss Phyllis Radford I have received two shells. 





A B 

A. Specimen from Tongaat. B. Specimen from East London. 

collected by her at East London, which at first appeared new 
to me, but a closer examination, and comparison with the 
shells in the Natal Museum, have convinced me that they can 
but belong to this species ; and the differences between them 
and typical shells do not seem to be of sufficient importance 
to wai-rant a varietal name. The new locality, East London, 
then, may be added to the localities cited by Connolly in his 
Revised Reference List. 

The chief differences are the less cylindrical form of Miss 
Radford's shells, with sharper apex, the more oblique aperture, 
and the absence of the intense constriction behind the peri- 
stome at the base. The species varies considerably in size 
and in relative width, two of the specimens of which Sturany 
gives the dimensions being even shorter, and one of them 



ox SOUTH AFRICAN ENNE^. 09 

relatively wider, than the East London shells ; therefore, the 
average smaller size and greater obesity of the East London 
shells cannot be considered of importance, since they might 
not be maintained in a series. 

The obliquity of the aperture which inclines from the 
right, above, to the left, below, is conspicuous, and is not 
equalled in any of Stnrany's figures, nor in any Natal 
example that I have examined. The absence of the deep 
constriction behind the labrum is probably the most important 
feature, but is not so easily detected. The accompanying 
figures are designed to illustrate this divergence from the 
type. 

The dimensions of the East London shells are as follows : 
height X width, 2-69 x 1-36, •2-64 x 1-37 mm. 

It will be seen that while the width of these shells exceeds 
that of even Stnrany's largest measured example, the ratio of 
width to height is greater in the specimen of which he gives 
the dimensions as 2*5 mm. higli and 1"3 mm. wide. 

Ennea connollvi 2Ieh-. & Pons. PI. V, fio-. ,52. 

Ennea couuollyi 31elv. it- Pons.. Ann. and Mag. Nat. Hist., vol. iv 
(1909), p. 486, pi. viii, fig. 2 ; Connolly, Ann. S. Af. Mus.. vol. xi 
(1912), p. 69. 

As Connolly indicates, loc. cit., the original figure does not 
adequately represent the shell. Not only is it too wide, but 
the sutures are too shallow, and certain apertural processes 
are not shown. Therefore this opportunity is taken to 
publish a new figure. 

The dimensions set forth in the description, given in 
" round numbers," probably give almost as erroneous an idea 
of the true proportions of the shell as the figure does, for in 
my series, kindly supplied by the discoverer, I have no 
example nearh' so attenuate as the recorded dimensions of 
the type would indicate it to be. 

The following table of dimensions and ratios will indicate 
the amount of variation in those respects which may be 



70 



HENRY CLIFDEN BUENUP. 



looked for in this species, and also the extent of the dis- 
crepancy between the authors' original dimensions and those 
indicated by their figure. To ascertain the dimensions repre- 
sented by the figure, the height is taken from the artist's 
dimension line, and the width is calculated therefrom, pro- 
portionately to the height and width of the figure. 



Specimens from various localities. 


Height. 


Width. 


Width X 100 
Height. 






mm. 


mm. 




Type as per authors' dimensions 




300 


100 


33-33 


Type as per original fig. . 




3-25 


1-56 


48-00 


Type of new fignre from Amajuba 




302 


1-29 


42-71 




/- 


3-19 


1-27 


39-81 


Other specimens from Amajuba 


. J 


316 


1-31 


41-46 




( 


3-06 


1-32 


43-14 






3-28 


1-28 


39-02 


Specimens from Hennop's River 


3-28 
309 


1-22 
1-22 


37-20 
39-48 




2-97 


1-24 


41-75 



Although my examples from Hennop's River tend towards 
a more attenuate form than those from Amajuba, some of the 
narrowest of the latter are, either actually or relatively, 
narrower than the widest of the former ; so a larger series 
than I have been able to examine Avould be necessary before 
it could be seen if the narrower average would be main- 
tained. Coincident with the more attenuate form, a deeper, 
more oblique suture is observable ; but no increase in the 
number of convolutions, beyond, perhaps, a quarter of a whorl, 
can be traced ; therefore it would appear that the increase in 
height is mainly attained by a looser coiling of the whorls. 

In the Hennop's River shells the aperture is less erect than 
in those from Amajuba, sloping, in varying degree, from 
right, above, to left, below; and the upper tooth of the 
columellar lip is uniformly situate higher, being close up to 
the junction of that lip Avith the Avail of the body-whorl. 

Melvill and Ponsonby draw attention to the alliance of this 



OiS^ SOUTH AFRICAN ENNE-E. 71 

species with E. arnoldi Stur., to which it bears a strong- 
resemblance ; but the former is easily distinguished b}^ its 
wider, less obstructed aperture, and the two teeth on the 
columellar lip which are absent in Sturany's species. 



Ennea inhluzaniensis n. sj). PI. Y, figs. 53, 54, 55. 

Shell small, cylindro-elliptical, deeply rimate, thin, shining, 
semi-transparent, apex rounded; w^horls 6^, slightly convex, 
the first 2h smooth, the next 3.^ delicately transversely striate 
immediately below the suture, with occasional stria? passing 
partly or wholly across the whorl, and the last i-whorl more 
strongly, regularly striate across its breadth, round the base, 
and into the rimal depression ; suture moderately impressed ; 
aperture erect, oblong, w^th Avhite peristome slightly 
thickened and moderately expanded, the ends joined by a 
distinct white callus, and furnished with the following pro- 
cesses : A rather narrow, sharp, in-running parietal plait 
arising a little above the suture and in front of the plane of 
the peristome, a rather long, narrow, simple, sub-central 
labral tooth, and a small, rounded, internal columellar plait. 
There is also a slight, svib-central swelling on the columellar lij). 

Height 3'60, width 1*47 mm. 

Hab. — Inhluzani Hill, Dargle, Natal (Burnup). 

Among the forty-five specimens collected, scarcely any 
deviation from the type, except in size, is observable. The 
" columellar plait " of the description is merely the rounded 
termination of the axial column Avithout contortion or 
thickening. The labral tooth is more solid than is usual, the 
excavation behind the lip being very shallow ; but the 
position of the tooth is there very clearly shown by a con- 
spicuous white spot. The swelling on the columellar lip is 
scarcely thickened, the corresponding excavation being very 
deep. 

In the study of this shell I feared, from Melvill and 
Ponsonby's description and figure of Ennea juxtidens, 
and from two very poor specimens, immature, calcined, and 



/Z HENRY CLIFDEN BURNUP. 

holed, in my collection, that the new form might possibly be 
inseparable from their species. Mr. E. A. Smith, therefore, 
most kindly undertook the comparison of my type with the 
type of juxtidens in the British Museum, and has furnished 
me with the following' detailed report : 

"Ennea juxtidens, type, is a larger shell than yours, 
the sculpture is hardly apparent, or at all events much less 
visible than in your shells : the peristome is- stronger, 
especially on the columellar side, where, and at the base, it 
is i-ather more expanded than in inhluzaniensis. The 
parietal lamella, beyond being stronger, offers no difference 
of any importance — it arises higher on the body-whorl than 
the suture, just the same as in your shells. The columellar 
fold is quite the same as in inhluzaniensis. The tooth on 
the labrumis of the same character. In juxtidens there is 
a feeble denticle low down on the columellar side, but not 
quite basal ; it is well within the edge of the peristome. The 
swelling about the middle of the collumellar lip in your 
shells is higher up than the faint tooth in juxtidens. This 
is the most distinguishing feature in M. & P.'s species in 
comparison with yours. 

" On the whole I think it would be safer to regard inhlu- 
zaniensis as a distinct species rather than as a variety of 
juxtidens." 

Since seeking Mr. Smith's opinion, Mr. J. H. Ponsonby has 
generously given me a mature co-type of E. juxtidens in 
excellent condition, from his own collection. Thus I am 
enabled to present new figures of that species (PL V^, figs. 
57, 58, 59) for comparison with those of inhluzaniensis. 
The dimensions are — height 4"53, width 2*05 mm. In con- 
sidering the difference in size, it should be borne in mind 
that the figures of inhluzaniensis are magnified by 9*2 
diameters, and those of juxtidens by 7"45. 

Mr. Farquhar has suggested a relationship between 
inhluzaniensis and elliptica Melv. & Pons., and 
though I think the differences are too many and too great to 
admit of close affinity, the size and general form of the 



ON SOUTH AFRICAN ENNE^. 73 

shells, together Avith the unusual freedom from obstruction in 
the aperture, warrant a comparison. 

The flatter Avhorls, shallower suture, much smoother and 
more highly polished surface, rounder aperture, with more 
widely expanded peristome, shorter parietal plait, and the 
two teeth on the columellar lip (or one in var. mane a) should 
always readily distinguish elliptica from the present species. 
Elliptic a, too, shows a conspicuous opening fi'om the rima 
to the axial perf oi'ation, while none is visible in i n h 1 u z an i e n - 
sis. The swelling on the columellar lip of inhluzaniensis 
cannot be looked upon as representing one or other of the 
teeth on the columellar lip of elliptica, for its position does 
not correspond with that of either. 

The dimensions of such specimens of inhluzaniensis as I 
have measured in addition to the type, are as follows : 
height X width, 3-59 x 1-52, 8-41 x 1-49, 3-85 x 1-49, 3-17 x 
1-42, 2-84 x 1-42 mm. 

From the dimensions of the two specimens figured it would 
appear that juxtidens, besides being larger than inhluza- 
niensis, is also proportionately a wider shell, the ratio of 
width to height of these specimens being respectively 45*25 
and 40*84 per cent. ; and Melvill and Ponsonby's dimen- 
sions of their type uphold the suggestion. The appear- 
ance of my calcined co-types of juxtidens, which are too 
brittle to admit of actual measurement, also seems to support 
it ; but a larger series would be required before the fact could 
be established, for my smallest example of inhluzaniensis, 
manifestly a dwarfed shell, is, with a ratio of 50 per cent., 
proportionately much wider than the figured juxtidens. 



Enn ea pr em nodes, Sturany. PI. V, fig. 56. 

Ennea premnodes Sinr., Ann. Hofnuis. Wien, xvi (1901). pub. 1902, 
p. 69, fig. .5; Connolly, Ann. S. Af. Mus., vol. xi (1912), p. 84. 
•• 1901 '■ [sic). 

Through the kindness of Dr. Sturany in giving me one of 
his co-types, I am enabled to offer a description and figure of 



74 .HENRY CLIFDEN BURNUP. 

this little-known species, which will be easiei- of access to 
South African readers than the orig-inal. 

Shell small, elongate, cylindrical, narrowly umbilicate, thick, 
white, opaque, apex rounded; whorls 7^, slightly ventricose, 
the first 3 smooth, the next 4 nearly smooth, except im- 
mediately below the suture, whei'e they are clearly, rather 
coarsely transversely rib-striate, and the last half whorl simi- 
larly rib-striate all over, except immediately behind the lip, 
where it again becomes smooth ; suture moderately deep ; 
aperture rather large, rounded, with peristome much 
thickened, expanded and reflexed, with well-developed callus 
connecting the extremities, and furnished with the following- 
processes : a strong blade-like in-running parietal plait, a 
large prominent, bluntly pointed labral tooth with corre- 
sponding pit outside, a small tooth near the base of the 
columellar lip, and a medium-aized deep-seated columellar 
plait, flattish and drawn to a small mammillated point at the 
lower right corner. In addition to these regular processes 
there is a very minute denticle on the outer, upper edge of 
the labrum, close to the suture, not shown in the fig. 

Height, 5"89; width (including expanded labrum), 2 '21 mm. 

Hab. — Albany District, Cape of Good Hope (Penther). 

The width of the shell gradually increases with each 
successive whorl after the rapid increase of the apex, so 
giving the spire a somewhat bluntly elongate-conical form. 
To what extent the opaqueness of the shell may be considered 
characteristic of the species, or to Avhat extent it may be due 
to incipient erosion, could only be decided were more ex- 
amples available for examination ; but its solidity suggests 
that in maturity it would always be opaque : my shell, though 
in good condition, does not appear to have been collected 
alive, so slight erosion in it is probable, though it is not 
conspicuous. The minute denticle on the outer, upper edge 
of the labrum, which is very inconspicuous, may not be 
characteristic, and is scarcely worthy of note except as regards 
a comparison of this shell with one of the forms of Ennea 
m on tan a Melv. & Pcma. (PI. V, fig. 60), hereafter discussed. 



ON. SOUTH AFRICAN ENNE^. (D 

E line a montana Meli:. & Pons. PI. \, figs. 60, 61, 62, 63, 

64, 65, m. 

Ennea montana Melv. S: Pons.. Ann. and Mag. Nat. Hist., vol. xii 

(1903), p. 599, pi. xxxi, fig. 15. 
Ennea parallela Melv. &: Pons.. il)id.. vol. iv (1909j, p. 4S9. pi. viii. 

fig. 9. 

On first studying- this species the material at my command 
was small, and showed so much variation, especially between 
my own two shells, ex coll. Farquhar, reputed to represent 
respectively montana and parallela (figs. 61, 62, 63, and 
fig. 60), but also between these two shells and two of the 
co-types of parallela lent to me by Mr. Ponsonby (figs. 65 
and 66), and between those two co-types inter se, that I Avas 
not only prepared to admit the validity of the two species, 
but was endeavouring to define the limitations of intermediate 
forms. As, however, neither of my shells and none of Mr. 
Ponsonby^s co-types of parallela seemed to agree accurately 
with Melvill and Ponsonby 's descriptions and figures, a com- 
parison with the types in the British Museum became essential. 
Mr. Edgar A. Smith most kindly undertook the investigation, 
and, being supplied by me with certain shells and drawings, 
has furnished the following report : 

" Fig. 65 represents the true parallela. Of the shells, that 
figured as 65 and the one unfigured agree with M. & P.^s type. 
1 do not regard the third example (fig. 66) as a distinct species. 
In my opinion it is an older shell with more developed teeth 
and a slightly modified labrum. The shell figured as 60 I 
also consider as parallela with well-developed lip and teeth ; 
the labral tooth is strong, and does not show the tendency to 
be double, as in other specimens. 

"I have compared the types of parallela and montana, 
and cannot regard them as distinct. The latter is an old 
shell with lip and teeth more thickened. Your figure (re- 
drawn as fig. 61) shows a minute columellar tooth which does 
not exist in the type." 

Mr. Smith does not comment upon the presence, in the shell 



7G HENKY CLIFDEN BURNUP. 

drawn as fig. 60, of tlie minute tooth on the outer edge of the 
labruni, close under the suture, so I infer that he attaches no 
significance to it, and it is probably an accidental development. 
It is interesting, however, to note the presence of an equivalent 
tooth in my specimen of Ennea premnodes 8tur. (fig. 56), 
a somewhat similar but much larger and heavier-built shell. 
The position of this tooth in premnodes is a little further 
forward, and so it is lost in perspective and shade in the 
figure. As Sturany does not mention this tooth in his descrip- 
tion of premnodes, it is to be inferred that it is absent from 
the type, and its presence in my specimen is also accidental. 

The minute columellar tooth shown in fig. 61, and, fide 
Smith, non-existent in the type of montana, is, curiously 
enough, shown in the original figure, though not very pro- 
minently, but is not referred to in the oi-iginal description. 
In no specimen that I have examined is it so conspicuous as 
in that shown in figs. 61, 62, 63. In some it is but slightly 
raised (e. g. fig. 60) ; in others it is merely represented by a 
slight swelling (e. g. fig. 66), but in the greater number it is 
entirely absent. 

Since submitting the matter to Mr. Smith, Mr. Farquhar 
has kindly sent me his whole series for examination, con- 
sisting of four specimens of montana collected at and near 
Mountain Drive, Grahamstown, one of them being from the 
original lot, five specimens collected with the original lot 
of parallela near the Asylum, Grrahamstown, and ten 
specimens, unnamed, collected at Gowie's Kloof, near the 
same city. I have also been favoured by an opportunity to 
examine the only specimen in the Alexander McGregor 
Museum, Kimberley, also from Gowie^s Kloof. This large 
series amply confirms Mr. Smith's views that parallela is 
inseparable from montana. It seems to be a species very 
constant in its elongate cylindrical form, in its sculpture con- 
fined (except on the apical whorls, which are smooth, and the 
last half -whorl, in which the sculpture crosses the whole width) 
to the area immediately below the suture, and in the position 
of the important apertural processes, but most variable in the 



ON SOUTH AFRICAN ENNEiE. 11 

development of those processes and in the extent to which the 
peristome is thickened and expanded. 

The divergencies from a conceivable normal type being 
apparently more a matter of degree in development than a 
modification of plan, and no series of specimens exactly sup- 
porting any definite line of departure from such a normal 
type, it would not seem desirable to attempt to define named 
varieties, and the species must remain one diflicult to deter- 
mine. It is hoped that the series of figures chosen to illustrate 
the species may serve as a guide to the amount of variation to 
be looked for. 

From these a summarised definition of the species may be 
drawn up as follows : 

Shell small, elongate-cylindrical, rounded above and below, 
deeply rimate and narrowly perforate, semi-transparent, 
polished; whorls about 7, moderately convex and impressed 
at the suture, sculptured immediately below the suture with 
transverse rib-strin3 of varying strength, except the first 2| 
whorls, which are smooth, and the last half-whorl, in which 
the stria3 are continued across the width of the whorl and 
around the base ; aperture somewhat roundly trigonal, 
broader than high, with more or less thickened and ex- 
panded peristome reflexed on the columellar and basal lips, 
furnished with at least four processes varying much in 
development, viz. a moderate, sometimes heavy, in-running 
parietal plait, a moderate or large, simple or more or less 
divided labral tooth, a small tooth about the base of the 
columella, and a wholly internal columellar plait, large or 
small, more or less thickened below, rounded or produced to 
a mammillate point at the lower right margin, the colu- 
mellar lip sometimes bearing a minute subcentral tooth or 
swelling. 

Height 3'03, width 1"19 mm. (average of 22 specimens 
measured). 

Hab. — Environs of Grrahamstown, Cape of Good Hope. 

The following are the dimensions of the figured specimens 
and a few others : 



78 



HENEY CLIFDEN BUENUP. 



In my collection (figs. 61, 62, 63) 

Ditto (fig. 60) 

In collection of J. H. Ponsonby (co-type 

ofparallela) (fig. 65) 
Ditto, ditto (fig. 66) ..... 
In collection of J. Farquahar (part of 

original lot of parallel a) . 
Ditto, ditto ....... 

In collection of J. Farqnliar (part of his 

series of mont ana) . . . . 

Ditto, ditto ....... 

In collection of J. Farquliar, from Gowie's 

Kloof 

Ditto, ditto . . . . . 

In collection of Alexander McGrregor 

Museum, Kimberley .... 
In collection of J. H. Ponsonby, from 

Hamilton Reservoir, Graliamstown (fig. 

64) 



Height 


Width 


mm. 


mm. 


3-23 


1-27 


3-08 


1-10 


2-93 


M7 


3-06 


1-27 


3-19 


1-24 


3-10 


1-13 


3-05 


1-21 


3-02 


1-19 


3-08 


M3 


2-67 


1-15 



2 '94 



3-03 



M2 



1-24 



Ennea ponsonbyi n. s-p- PI- ^} figs- 67, 68, 69. 

Shell small, ovate-cylindrical, with blunt apex, vitreous, 
semi-transparent, whitish, deeply i-imate and narrowl}" but 
distinctly perforate ; whorls 6, rather convex, the first 2 
smooth, the rest regularly, distinctly, rather strongly, 
sculptured with transverse rib-strise, except a small area on 
the body -whorl, the whole width of the whorl, immediately 
above the columellar lip, which is smooth ; suture rather 
deeply impressed ; aperture rather erect, roundly quadrate 
with whitish peristome much thickened, widely expanded 
and strongly reflexed, the ends connected by an indistinct 
callus, and the apertural pi'ocesses being as follows : A strong, 
in-running, blade-like parietal plait, a large, strong, bi-lobed 
labral tooth, a small in-running basal tooth, a large, fiattish, 



ON SOUTH AFIJICAX ENNE.E. 79 

deep-seated coluniellar plait, but little thickened on the 
lower edge and slightly drawn forward on the right, and a 
swelling on the coluniellar lip. 

Height 2-91, width 1-33 mm. 

Hab. — Gowie's Kloof, Grahamstown, Cape of (xood Hope 
(Farquhar). 

All the specimens that I have seen, namely five in my own 
collection, received from Mr. Farquhar, nine in Mr. Ponsonby\s 
collection, and a large sei-ies in the Alexander McGregor 
^luseum, Kimberley, come from the same locality, and are 
most constant in detail, varying slightly only in size, relative 
width, and in the number of whorls {oh to 6j). 

The following are the dimensions of such examples as I 
have measured, in addition to the type, and fairly repre- 
sent the extent of variation observed : Height x width, 
3-20 X 1-37, 2-94 x 1-37, 2-88 x 1-31, 272 x 1-30, 2-66 x 1-.33 mm. 

A curious feature in this species is the position of the 
smooth area on the body-whorl. In many, if not most, striate 
Enneffi, there is such an area distinct from the callus, more 
or less smooth, immediately above the aperture, in which 
position it is easily conceived to be formed by the constant 
passage of the mantle over the spot while the mollusc is 
active. In by far the greater number of the specimens of 
this species examined, the centre of the smooth area is 
considerably to the left of the aperture, though in a few its 
position is normal. It is difficult to understand why it 
should generally be so far misplaced, unless the animal 
possess some peculiar development of the mantle. 

This species has been tentatively identified as Ennea dar- 
glensis M. & P., and distributed as such ; but the differences 
are so many and so great that the two species cannot be 
considered as closely akin ; a mere glance at the figures of 
the two species should be enough to bring conviction as to 
their wide distinction. 

The general arrangement of the processes of the aperture 
is somewhat similar to that of E. m on tana M. & P., but 
the shorter, stouter form Avith fewer whorls, the continuous 



80 HENRY CLTFDEN BURNUP. 

sculpture, and the much more thickened and reflexed peri- 
stome, readily distinguish the present species. 

I have much pleasure in associating" with this shell the 
name of Mr. J. H. Ponsonby, from whose collection the type 
is chosen, and from whom I have received so much help in 
the form of advice, specimens for study, and the obtaining 
and transmitting of reports upon the types in the British 
Museum, without which much of this paper must have 
remained unwritten. 



EXPLANATION OF PLATES III-A^ 

Illustrating Mr. Henry C. Burnup's paper " On South 
African Enneas, with Descriptions of New Species and 
Varieties." 

PLATE III. 

Fig. 1. — Ennea isipingoensis, Stur., from the type in Natnrlu 
Hofmus., Yienna. 

Fig. 2. — Ennea isipingoensis /Si«r. var.cliscrepans S^itr.,from 
the type of the variety discrepans in Naturh. Hofmus., Vienna. 

Fig. 3. — Ennea isipingoensis Stm: var. discrepans S^zw., from 
the type of the variety simillima Stur., in Naturh. Hofmus., Yienna. 

Figs. 4, 5, 6. — Ennea isipingoensis S^i/r. var. sturanyi n. Type 
of var., from Ntimbankulu, to be j)laced in the British Museum. 

Figs. 7, 8. — Ennea elliptica Melv.&Pons., typical, from specimens 
from the original locaUty, Maritzburg, in the Natal Museum. 

Figs. 9, 10. — Ennea elliptica Melv. & Pons., from specimens from 
Tongaat Beach Bush, in the Natal Museum. 

Figs. 11, 12. — Ennea elliptica Melv. &■ Pons., from specimens 
of aberrant forms from Dargle, to be placed in the British Museum. 

Fig. 13. — Ennea elliptica Melv. & Pons. var. manca n. Type 
of var., from Fort Nottingham, to be placed in the British Museum. 

Fig. 14. — Ennea elliptica Melv. & Pons. var. manca n., from a 
specimen from Fort Nottingham, in the Natal Museum. 

Figs. 15, 16, 17. — Ennea elliptica Melv. & Pons. var. cselata n. 
Type of var., from Eshowe, to be placed in the British Museum. 



ox SOUTH AFRICAN ENNE"^. 81 

Figs. 18, li), 20. — Enuea sylvia Melv. & Pons., from a specimeu 
from the original locality, Maestroom Forest (Farquhar), in the Natal 
Museum. 

Figs. 21, 22. 23. — Ennea melvilli n. .■*^). Type, fi-om Nottingham 
Road, to be placed in the British Museum. 



PLATE IV. 

Fig. 24.— Ennea farquhari Melv. <£• Pons., from a co-type from 
Grahamstown, in coll. J. H. Ponsonby, to be placed in the British 
Museum. 

Figs. 25. 26, 27. — Ennea farquhari Melv. & Pons. var. bertha3 
(Melv. cC- Pons.), from a co-type of Ennea berthas Melv. & Pons., from 
Kai-kloof, in the Natal Museum. 

Figs. 28, 29. 30. — Ennea farquhari Melv. & Pons. var. avena n. 
Type of var., from Maritzburg, to be placed in the British Museum. 

Fig. 31. — Ennea farquhari Melv. & Pons.vav. avena n., from a 
specimen from Durban, to be placed in the British Museum. 

Fig. 32. — Ennea darglensis Melv. & Pons., from a specimen from 
Inhluzani Hill, Dargle, in the Natal Miiseum. 

Figs. 33, 34, 35. — Ennea darglensis Melv. & Pons. var. illovo- 
ensis n. Type of var., from Ntimbankulu, Mid-Illovo, to be i3laced in 
the British Museum. 

Fig. 36. — Ennea consobrina Aaceij, h-om the type, from Albany, 
in coll. J. R. le B. Tumlin. 

Fig. 37. — Ennea thelodonta Melv. d- Pons., from a specimen from 
Port Elizabeth (Farquhar), in the Natal Museum. 

Fig. 38. — Ennea munita 31elv. A- Pons., from a specimen from 
'• ? near McLear" (Ponsonby), in the Natal Museum. 

Figs. 39, 40, 41. — Ennea maritzburgensis Melv. c{- Pons., typical, 
from a specimen from Maritzburg, to be placed in the British Museum. 

Figs. 42, 43, 44. — Enuea maritzburgensis Melv. &• Pons. var. 
contracta n. Type of var., from Nottingham Road (Taynton), to be 
placed in the British Museum. 

Fig. 45. — Ennea maritzburgensis JZe/y. & Pons., from a specimen 
of an aberrant form near maritzburgensis, typical, from Karkloof 
(Taynton), to be placed in the British Museum. 

Fig. 46. — Ennea maritzburgensis Melv. c(- Pons., from a specimen 
of an al)errant form near var. contracta. from near Rosetta (Hickey, 
per Ponsonby), to be i^laced in the British Museum. 

VOL. 3, PART 1. 6 



82 HENRY CLIFDEN BURNUP. 

PLATE V. 

Fig. 47. — Ennea columnella Melv. S: Pons., from a co-tyi^e from 
Karkloof (McBean), in the Natal Museum. 

Fig. 48. — Ennea mariaj Melv. it- Pons., from a co-type from 
Somerset East (Miss M. Bowker), in coll. J. H. Ponsonby. 

Figs. 49, 50, 51. — Ennea mooiensis n. sp. Type, from Game Pass, 
Upper Mooi River, Natal, to be placed in the British Museum. 

Fig. 52. — Ennea connollyi Melv. & Pons., from a specimen from 
the original lot from Amajuba (Connolly), in the Natal Museum. 

Figs. 53, 54, 55. — Ennea inhluzaniensis n. sp. Type, from 
Inhluzani Hill, Dargle, to be placed in the British Museum. 

Fig. 56. — Ennea premnodes Star., from a co-type from Albany 
District (Penther, per Sturany), in the Natal Museum. 

Figs. 57, 58, 59. — Ennea juxtideus, Melv. ct Pons., from a co-type 
from Van Reenan (ex coll. J. H. Ponsonby), in the Natal Museum. 

Fig. 60. — Ennea montana Melv. & Pons., from a specimen of the 
original lot of Ennea parallela, collected near the Asylum, Grahams- 
town (Farquhar), in the Natal Museum. 

Figs. 61, 62, 63. — Ennea montana Melv & Pons., from a specimen 
from Mountain Drive, Grahamstown (Farquhar), in the Natal 
Museum. 

Fig. 64. — Ennea montana Melv. S: Pons., from a specimen from 
near Hamilton Reservoir, Grahamstown (Farquhar). in coll. J. H. 
Ponsonby. 

Figs. 65, 66. — Ennea montana Melv. & Pons., from co-types of 
Ennea parallela Melv. & Pons., from Grahamstown, in coll. J. H. 
Ponsonby. 

Figs. 67, 68, 69. — Ennea ponsonbyi n. sp. Type, from Gowie's 
Kloof, Grahamstown (Farquhar). ex coU. J. H. Pousonljy. to be placed 
in the Bi-itish Museum. 



Ann . Na.t ol Mus . Vol . I II 



PI. III. 




20. 

J. Green. litK. 



SOUTH AFRICAN ENNEvE. 



Ami. Natal Mus.Vol.llI 



PI . rv^. 




SOUTH AFRICAN ENNE^. 



Ann. Natal Mus.Vol.IlI . 



Pl.V. 



.,«<^=^ 




66. 

J. Green litVi. 



SOUTH AFRICAN ENNE^. 



DEVELOPMENT OF FLANULA IN A PLUMULARIAN HYDROID. 83 



On the Development of the Planula in a Certain 
Species of Plumularian Hydroid. 

By 

Ernef^t Wavreii, D. Si*. (Loud.)* 



With Plate VI and 4 Text- figs. 



The development of the planula in this hydroid is note- 
worthy in that the egg never becomes charged Avith yolk. 
The ovum remains small and segments in the midst of a feed- 
ing or placental tissue. Ultimately the embryo grows into a 
well-developed planula with dimensions very greatly exceeding 
those of the original egg. 

Some brief notes on the subject were communicated to the 
South African Association for the Advancement of Science at 
the annual meeting held in Louren^o Marques, July, 1913. In 
the present paper the necessary details with drawings are 
given, and the species of hydroid is described. 

The hydroid was found in January, 1911, in a rock-pool on 
the north side of the mouth of the St, John's River, Pondo- 
land. Only two clusters were gathered, and they both 
occun-ed on the shells of living oysters much incrusted with 
the calcareous tubes of Serpula, The upright pinnate stems 
were about ^ in. in height. 

Trophosome. — Hydrorhiza consists of an irregularly 
branched stolon, which in the present specimens was creep- 
ing among the tubes of Serpula covering oyster-shells (text- 



Text-fig. 1. 




/{[natsc^y-J 



DEVELOPMENT OF PLANULA IN A PLUMULAKIAX HYDROID. 85 

fig 1, A). The perisarc is not so distinctly divided into an 
onter diffuse layer and an inner compact layer as in some 
species of Plum ularia. 

Diaiiieter of hydrorhiza about 0'19 mm. ; thickness of 
perisarc 22*7 /u. 

Hydrocaulus. — Monosiphonic, the hydrorhiza carries up- 
right pinnate stems of various heights. 

The proximal portion of the main-stem, not beai-ing pinnas, 
is about half the length of the distal portion carrying them, 
and it is somewhat irregularly divided into internodes by 
transverse nodes (text-fig. 1, B). The distal portion of the 
stem is regularly divided by oblique nodes. Between the 
two portions of the main-stem there are two very oblique 
joints (text-fig. 1, B ; C, «., n.). 

All the internodes of the main-stem in the distal pinnate 
portion carry a hydrotheca with a mesial sub-calycine nema- 
tophore and a pair of supra-calycine ones. 

'llie pinnaj alternate, and on an ordinary stem there are nine 
to eleven on each side. They usually carry only two hydrotheca?, 
but sometimes there may be three (text-fig. 1, B). The pinnas 
are borne on a short process springing from the main-stem at 
the back of the hydrotheca (text-fig. 1, C). The proximal 
internode is very short, bears no structures, and the upper 
and lower nodes are transverse (w.j, n.i) ; the rest of the pinna 
generally consists of only two alternating non-thecate and 
thecate internodes separated by oblique nodes. These non- 
thecate internodes carry a median nematophore. 

Frequently the pinna?, especially the proximal ones, bear a 
pinnule which is carried on a short process springing from 
the level of the proximal hydrotheca. It consists of a basal 
internode (text-fig. 2, p. t.^^.) with transverse nodes (text-fig. 1, 
C, ?/.2, "Jt-o), followed by a non-thecate and thecate internode. 



Text-fig. 1. — A (nat. size). Colony on oyster shell. By.G. Hydrorhiza 
bearing pinnate stem with pinn* carrying three hydrotheca\ C X 53. 
Piece of pinnate stem with male and female gonangia. D x 55. Side 
view of female gonangium carrying nematophores at the base. 



86 



ERNEST WARREN. 



It appears that the pinnules invariably spring from the lower 
side of the pinna?. They are set at an angle of about 30° to 
the pinna, and project downwards and outwards. These short 
pinnules are characteristic of the species. 

Length of the pinnate-stem with 11 pairs of pinnfe about 
12 mm., proximal non-pinnate portion 4 mm., pinnate part 
8 mm. Diameter of main-stem in non-pinnate region 0"16 mm., 
thickness of perisarc 21*58 /i. Length of thecate internode 
about 0*30 mm., width 0'14 mm. The average length of 

Text-fig. 2. 




Internodes of pinna and base of pinnule, x 200. 



pinna with two hydrothecse is about 0'91 mm., and the 
average length of pinnule is about 0"51 mm. The ccenosarc 
of the internodes exhibits an ectoderm channel on the posterior 
surface (text-fig. 3, A, p.e.c). 

Hydrotheca. — Cup - shaped, rather deep, expanding, 
margin somewhat everted, adcauline surface not adnate for 
whole length, but free above for about one-half of its length. 
Plane of mouth of hydrotheca is set at an angle of about 35° 
to main-stem or pinna. The hydrothecee are placed on the 
front or anterior surface of the pinnte (text-fig. 1, C). 

Height of hydrotheca about 0*16 mm., width at mouth 
about the same. 



DEVELOPMENT OF PLANDLA IN A PLUMULARIAN HYDEOIt). 87 

Xematophores all free and have the typic^-l plumulai-ian 
structure ; there is a median sub-calycine nematophore, and 
a pair of larger supra-calycine ones whi(?li scarcely project 
beyond the level of the mouth of tlie hydrotheca (text-fig. 2). 
The sarcothecae are bithalamic, canaliculate, and tend to be 
narrow at the T^ase. 

Hydranth. — j^bout twenty tentacles in a single verticil. 
Polyp constricted below the whorl of tentacles, dividing the 
coelenteron into an upper and lower division ; the upper part 
is lined by narrow columnar cells, the lower portion by 
vacuolated digestive endoderm (text-fig. 3, A). 

Nematocysts in the ectoderm of the tentacles are small ; 
they measure about 3*2 n in length and I'OQ fx in breadth. In 
the nematophores the nematocysts are of considerable size, 
having a length of 14*8 fx and a breadth of 3*2 /u. 

Gonosome. — The male gonangia are borne on the more 
proximally placed pinna?. The female gonangia occur on the 
proximal portion of the pinnate region of the main-stem ; they 
arise just below the hydrotheca and on one side of the sub- 
calycine nematophore. The niitritive substances are naturally 
more abundant in the main-stem than in the lateral pinnas, 
and from analogy with other hydroids we should expect to 
find the female gonangia in the former rather than in the 
latter position. This is in accordance with the view that the 
development of the female sex is partly induced by a i*ich 
nutritive supply. 

Male Gonangium. — The gonotheca when fully grown is 
cylindrical, and is about 0*36 mm. in length and 0'12 mm. in 
width (text-fig. 1, C). The perisarc is quite thin, and there 
is no distinct operculum. At the base where it springs from 
the pinna there is no jointed stalk, and there are no 
nematophores arising from it. 

In the developing male gonangium the spermatic cells, 
which will subsequently form the generative mass of the 
single reduced gonophore, may be seen embedded in the 
endoderm of the hydrocaulus (text-fig. 3, A, sp.c). In the 
figure the spermatic cells are entering the blastostyle (h.G.) of 



,88 ERNEST WARREN. 

the growing gonangium. Subsequently the genital cells will 
separate themselves from the endoderm on one side of the 
blastostyle, and this may be regarded as representing the 
formation of a gonophore (fig. B, t), which is covered by a 
thin layer of ectoderm cells continuous with the ectoderm of 
the blastostyle. At the apical region the ectoderm (c.^J.) 
of the blastostyle consists of elongated cells constituting the 
" Deckenplatte " of Weismann. 

Female Gonangium. — The gonotheca when mature is 
ovoid and flattened. The plane of flattening is at right 
angles to the antero-posterior plane passing through the 
main-axis. 

The female gonotheca arises on the main-stem just below 
the hydrotheca and on one side of the sub-calycine nemato- 
phore. It possesses a short stalk-segment with transverse 
nodes (text-fig. 1, C, n..^, n...). The stalk and the basal 
portion of the gonotheca are so curved that the main-axis of 
the structure is set at an angle of about 30° to the main-stem 
(text-fig. 1, C and D). The oldest gonangium is the most 
proximal one, and the gonangia are successively younger on 
passing distally. The perisarc is not ver}^ thick, but there is a 
well-defined large opercultmi^ and there the perisarc is stouter. 
At the base of the gonotheca itself and on the flattened 
surface there is a neraatophore on each side (text-tig. 1, D). 

Length about 0*81 mm., greatest lateral width 0"51 mm., 
thickness at right angles to plane of flattening 0*r2 mm. 

The female gonangium bears in an obscure manner a 
single gonophore which becomes provided with a single 
ovum. Before the ovum commences segmentation the gono- 
theca is very hollow, and the blastostyle occurs as a thin 
tubular structure running through the mid-axis. 

Systematic Position.^ — The hydroid possesses the typi- 
cal characters of the genus Plumularia with the 
exception of the presence of the downwardly directed off- 
shoots from the pinnge, and the somewhat unusual occurrence 
of the main-stem bearing hydrothecje. These pinnules are 
similar in every way to the pinna?, and they originate from 



DEVELOPMENT OF PLANULA IN A PLUMULARIAN HYDKOID. 89 

the pinnte just as the latter do from the main-stem ; they 
bear a short basal internode with transverse nodes and no 
nematopliore. 

Bale^ describes both Plumularia campanula Busk and 




Longitudiual sections of male gonaiiginiu. x 250. 

Plumularia buskii Bale as bearing hydrotheca? on the 
main-stem ; and AUman- figures a similar condition in 
Plumularia a r m a t a Alltna n . 



^ Bale, W. M., ' Catalogue of the Australian Hydroid Zoophytes.' 
1884, pp. 124-5. 

- AUman. G. J.. '"Challenger" Reports." vol. vii, 1883. pi. iv. fig. 4. 



90 ERNEST WAREEN. 

In the species Plumularia aglaophenoides Bale,Ba\e^ 
describes the occuiTence of pinnules in the following terms : 

"Hydrocaulus polysiphonic, flexuous, pinnas alternate, dis- 
tant, given off from each flexure of the stem, with two distinct 
oblique joints near the base, and obscurely jointed for the rest 
of their length, bearing hydrothecas as well as pinnules ; pin- 
nules alternate, approximate, one on each internode of the 
primary pinnae, both series borne towards the front and 
supporting a hydrotheca on each internode/' 

According to the definition of the genus Plumularia as 
given by Nutting,^ the pinnte are without accessory branches 
of any kind ; and the only other genera to which the present 
species could be referred are Poly plumularia G. 0. Sars 
(modified by Nutting) and Schizotricha Allman (modified 
by Nutting). Nutting defines Polyplumularia as possess- 
ing pinnae furnished with a hydrothecate ramulus or pinnule 
which springs from the first internode, and is more slender 
than the pinna from which it grows; while in Schizotricha 
the pinnaj bifurcate beyond the first internode at least in the 
mature colony. 

In the present species the ramulus or pinnule springs not 
from the first internode, but from the first hydrothecate inter- 
node. The pinnule arises from the pinna by a short internode 
bearing no structures and with transverse nodes, just as the 
pinna itself arises from the main-stem. This condition is 
shown in Allman's"'' figure of Schizotricha unifurcata 
Allman. Consequently, the lower of the two branches is to 
be regarded as the pinnule, and the upper as the distal 
portion of the pinna, as in the case of the species being 
described. 

It is not very clear that this production of pinnules forms 
a very satisfactory basis for founding a genus. The present 
species is transitional in this respect between Plumularia 

' Bale, W. M., loc. cit., p. 126. 

^ Nutting, C. C, ' American Hydroids. Part I : The Plumularidas,' 
1900, pp. 54, 78, 83. 
^ Allman, G. J., loc. cit., pi. vii, fig. 2. 



DEVELOPMENT OF PLAXULA IN A PLUMULAEIAN HYDEOID. 91 

with no pinnules according- to definition, and Schizotricha 
with many. Id the present species the formation of the pin- 
nules tends to be limited to the more proximally placed pinn«, 
while in Schizotricha it would appear that practically all 
the pinnas bear them. As a provisional designation, however, 
the name Schizotricha simplex is proposed. 



n. 

The Development of the Planula. 

The material was fixed in a warm alcoholic solution of cor- 
rosive sublimate and acetic acid. Several pinnate stems 
with female gonangia were sectioned in different planes, and 
the sections were stained wnth Delafield's hasmatoxylin 
followed by orange. 

Each stem bore gonangia of varying ages, and consequently 
all the different stages of development would appear to have 
been observed. Although such was the case, yet the youngest 
ovum that could be definitely identified as such was already 
in the endoderm situated at a short distance below the level 
of the mesial sub-calycine nematophore. 

Presumably, according to August Weismann and fi-om 
analogy with obsei'vations on other hydroids, the ova first 
arose in the ectoderm, and then migrated into the endoderm ; 
but in the present case the young ova were not definitely 
located in the ectoderm. 

The youngest ovum seen was small, measuring about 14 /u 
in diameter, and surrounded by ordinary endoderm cells. 
The presence of the ovum causes a slight swelling to project 
into the lumen of the internode (PL VI, fig. 1, 0.). 

The ectoderm immediately above the area where the ovum 
is imbedded in the endoderm early becomes slightly modified 
in that the cells are more columnar and regular than ordinary 
ectoderm cells (fig. 1, b. G.). This is the first beginning of 
the future gonangium. 

The perisarc situated just above {d.j).) has become markedly 



92 EKNEST WARHEN. 

thinner, apparently through some dissolving action of the 
subjacent ectoderm cells. 

In the next stage the plate of ectoderm has grown out 
into an ovoid swelling which is covered by a very delicate 
layer of perisarc continuous with the general perisarc. The 
endodemn follows the ectoderm, and the portion of endoderm 
(fig. 2, end.) carrying the ovum (0.) passes into the gonangium, 
and becomes located on one side of the blastostyle. The 
diameter of the ovum has increased to 17 lu. 

It may be mentioned here that all the figures in the plate 
are drawn from sections cut in the plane of flattening of 
the gonangium, with the exception of the following : figs. 7 
and 18 represent sections at i-iglit angles to the plane of 
flattening, and fig. 3 is a transverse section. 

At this time the mesogloea between the ectoderm and the 
area of endoderm in which the ovum occurs becomes thin and 
evanescent. The ovum gradually separates from the endo- 
derm, and the ectoderm above becomes modified, divides into 
smaller cells and forms a kind of cap over it. This cap and 
the layer of endoderm (fig. 2, end., and fig. 4, g. e.) immediately 
below the ovum maybe regarded as representing a rudimentary 
gonophore. 

The distal portion of the ectoderm of the young gonangium 
consists of tall columnar cells with short pear-shaped granular 
cells wedged between. This is the beginning of the covering- 
plate or " Deckenplatte " (fig. 2, c. p.). The coelenteron {C) 
expands distally and becomes somewhat T-shaped. 

The ovum now enters the ectoderm (fig. 3, 0.), and the 
covering cap becomes divisible into an inner columnar layer 
(fig. 4, i. I.) in immediate contact with the ovum, and an outer 
layer (o. /.) of fiat cells. The slight crack-like space (fig. 3, 
c. U.) between the two layers is the equivalent of the umbrella- 
cavity of a medusa, or more complete gonophore, which arises 
as a split in a distally placed thickening of ectoderm. The 
ectoderm at the sides of the gonangium gradually contracts 
away from the perisarc of the gonotheca (fig. 4, .sp.p s^9.o),and 
outside the flat ectodermal layer (o. /.) of the gonophore theie 



DEVELOPMENT OF PLANCLA IX A PLUMULARIAN HYDKOID. 93 

is a narrow space (figs. 3 and 4, .s^^^-)- ^^^'^^ long diameter of 
the ovum has now increased to about 20 ju. 

The gonangium continues to expand, and the ovum grows 
slightly and reaches a diameter of about 2S fi, but there is no 
obvious yolk passed into it. The covering cap of ectoderm 
of two layers also grows, the outer layer (fig. 5, o. I.) becomes 
thinner, and the inner {i. I.) thicker. A delicate mesogloea is 
formed between the ectodermal cap and the endoderm. The 
endoderm cells on Avhich the ovum is seated are granular and 
in an active physiological condition ; they stain more readily 
than the rest of the endoderm. 

The gonangium expands further, and the distal plate of tall 
columnar ectoderm cells (fig. 5, c. p.) begins to secrete a 
thicker layer of perisarc ; this is the beginning of the 
operculum. The blastostyle, except in the region of the 
operculum, has become quite separated from the chitinous 
gonotheca, and lies in the mid-axis. The ectodeimi lining 
the umbrella-cavity of the gonophore, consisting of the two 
layers, becomes separated from the ovum, so that a very 
definite cavity occurs above the egg (fig. 6, c. 0.). This 
cavity may be regarded as a space in the mesogloea of the 
gonophore. The umbrella-cavity (c. U.) of the gonophore 
is very narrow and almost obliterated. 

The ovum is probably fertilised just, before this stage, and, 
presumably after such fertilisation, it appears to secrete a 
kind of vitelline membrane (figs. 6, 7, 8, v. in.) which is 
thicker on the outer than on the inner side. It is possible, 
however, that this membrane is really of the nature of 
mesogloea, and is not actually formed by the egg. The endo- 
derm below the ovum becomes slightly pushed outwards into a 
blunt process (fig. 8, d. C.) consisting of narrow granular 
cells, and the egg is seated on this out-pushing, which may 
be regarded as the spadix of the reduced gonophore. The 
ovum now attains its maximum diameter of about 34 /./. 

In text-fig. 4 a comparison is made between a typical 
gonophore (A) and the highly modified gonophore (B) 
carried by the blastostyle of the present species. 



94 



ERNEST WARREN. 



In B the diverticulum of tlie coelenteron (C.) is directly 
comparable with the endodermal cavity of an ordinary 
gonophore, the granular layer [g. e.) being equivalent to the 
manubrial endoderm. Radial endodermal canals are not 
represented. The cavity (c. 0.) around the outer surface 
of the egg is the space between the manubrial endo- 
derm and ectoderm ; the manubrial ectoderm [M. e.) in the 
modified gonophore consists of a hemispherical layer of 
cubical cells. The umbrella-cavity (c. U.) is a narrow slit-like 
space. The outer ectoderm-lining (e. I. U.) to the umbrella- 

Text-fig. 4. 




Comparison of a typical gonophore (A) and the modified gonophore (B). 

cavity consists of a layer of fiat cells. Outside this ecto- 
dermal lining of the umbrella there is a space {sp.) eqiiivalent 
to the space between the umbrella-ectoderm and the outer 
ectoderm layer of the gonophore. The outermost ectoderm 
{e. I.) of a gonophore is indicated by a few flattened cells 
adherent to the tall attenuated cells lining the upper portion 
of the gonotheca. 

In fig. 7 of the Plate the gonangium is shown in vertical 
section at right angles to the plane of flattening. The 
terminal portion of the coelenteron of the blastostyle is seen 
at C. and the out-pushing to form the spadix occurs at 
d. G. 

The gonangium now grows to its full size, and the cells of 



DEVELOPMENT OF PLANULA IN A PLUMULARIAN HVDKOID. 95 

the covering-plate and of the distal endoderm of the blasto- 
style become very attenuated and full of vacuoles, and 
will subsequently disappear. The space (fig. 8, c. 0.) 
between the egg and the ectoderm of the gonophore in- 
creases in size, and the membrane around the ovupi (r. )ii) 
is very pronounced. The operculum consists of thicker 
perisarc than that of the rest of the gonotheca, and its 
lower edge is sharply marked off from the thinner perisarc 
below. The gonotheca has now assumed its definite form, 
and it does not further increase in size ; its greatest width is 
about 0"51 mm. and length 0"85 mm. 

At a somewhat later stage the outermost gonophore-layer 
of flat ectoderm cells disappears and the iriner layer of 
cubical or columnar cells breaks up to forfti an irregular 
cluster around the ovum (fig. 9, e. G.). The covering plate 
of ectoderm cells below the operculum and the endoderm 
layer of the terminal portion of the blastostyle are still 
faintly visible, but they are on the point of disappearing. 
The out-pushing of endoderm {d. C.) or spadix of the 
gonophore consists of cells which remain in an actively 
living condition. 

The ovum does not grow, the protoplasm is finely granular, 
and there is a large nucleus with nucleolus (fig. 9). There is 
still a pronounced membrane around the egg ; but it is much 
less developed on the inner side against the granular cells of 
the endoderm than on the outer side. 

In the next stage it will be seen that considerable changes 
have occurred in that the whole of the covering plate of 
ectoderm and the distal horizontal portion of the endoderm 
of the blastostyle have entirely disappeared. The out-pushing 
of endoderm (fig. 10, d. C.) or spadix of the gonophore has 
grown upwards, and has pushed the egg, together with the 
surrounding ceils derived from the breaking-up of the 
columnar ectoderm cells of the gonophore, into a more or less 
central position in close contact with the operculum. The 
vitelline membrane or mesogloea-layer round the ovum be- 
comes less distinct. The cells around the ovum arrange 



96 ERXEST WARREN. 

themselves in such a manner that they extend as columnar 
cells from the perisarc to the endoderm on which the ovum 
is still seated. 

Subsequently the ovum becomes quite central at the apex 
of the gonangium, and is closely surrounded by the tissue 
derived from the ectoderm of the rudimentary gonophore. 
The endoderm is now in the form of two symmetrically placed 
lateral lobes, one being the terminal portion of the endoderm 
of the blastostyle (fig. 11, t.j).), and the other is the endodermal 
spadix of the gonophore {cL C). The egg is seated symme- 
trically between the two lobes. 

The ovum segments and a rounded cluster of about sixteen 
loosely attached blastomeres is formed (fig*. 11, M.). The 
cells of the endoderm immediately under the young embryo 
(fig. 12, end.), become taller and more granular than those of 
the rest of the endoderm. It is considered probable that the 
nixtritive substances for the growth of the embryo are passed 
by this endoderm into the ectodermal tissue in which the 
embryo is imbedded. 

The blastomeres increase in number and become smaller. 
A segmentation- cavity soon appears (fig. 13, 8g. C), and 
this is not central, but is nearer to the apex of the gonangium. 
Accordingly the embryo is two or three cells in thickness 
below, while above it is mostly only one cell thick. The cells 
of the placental tissue {jil. t.) increase in size and height ; the 
tissvie is continuous with the thin layer of ectoderm Avhich 
covers the sides of the blastostyle {e.B.). 

The embryo is very obviously supplied with nourishment 
by the cells in which it is imbedded. The diameter of the 
embryo is two and a half times that of the original ovum, 
which never had any appreciable quantity of yolk. 

The embryo continues to expand and the constituent cells 
increase in number and diminish in size. The segmentation- 
cavity becomes larger (fig. 14, Bl.), A differentiation in the 
cells of the embryo can be seen on the lower side Avhere an 
inner layer of flattened cells now becomes evident {end.-^). It 
may be noticed that this inner layer of endoderm lines the 



DEVL:i-or.Mi;xr OK i'l.vnui.a ix a im.i .milaijiax iivkijoii). *I7 

sogmentiition-cavity only on the lowor half of the embryo. 
In the earliest stages of development the inner layer would 
nppear to arise rather hy diffei-ent iatioii tlian by delaniination. 
TIic outer cells or ectoderm {ecf.^) form a, layer one cell thick 
above and several cells thick below. The cells of the placental 
tissue in the space between the cup of the endoderm and the 
lower two-thirds of the embryo also show differentiation. 
Some of the cells are rounded or somewhat stellate ; they are 
further distinguishable by staining much more readily, and 
they appear to consist of protoplasm denser than that of the 
other cells. It is possible that these cells are the more active 
agents in transferring nutriment to the groAving endjryo. 

The end)ry() continues to grow, and the cells of whicli it is 
composed increase greatly in number, but their size remains 
the same as before. The embryo becomes pear-shaped 
(fig. 15). The inner layer of flat endoderm cells {t'll(L'^) is 
beginning to creep upwards, but the upper half of the 
embryo is still only one cell thick and consists of the outer 
ectoderm layer. The embryo as a whole is sinking down- 
wards, and the endodermal cup formed from blastostyle and 
gonophore is becoming deeply flask-shaped. The effective 
placental tissue {2^1. t.) now lies only between the embryo and 
the endoderra-cup. The outer portion of this tissue (e. E.), is 
extending downwards and consists of extremely elongated 
cells. At the mouth of the flask the ectoderm layer has 
separated slightly from the perisarc and forms a definite 
epitlielium adhering to the endoderm, and leaves a slight 
crack-like space {d. S.), extending from the pei'isarc to the 
pointed apex of the embryo. 

In the next stage the embryo has sunk still further towards 
the centre of the gonangium (fig. 10). The ectoderm of the 
embryo has become moi*e than one cell thick all round, and 
the endoderm also is several cells thick below and has com- 
pletely surrounded the blastococl (J5/.). The placental tissue 
(j^l. f-) is well-developed, while the outer portion {c. E.) 
retains its trausparent, attenuated condition. The flask- 
shaped endoderm layer {g. e.) derived from the blastostyle 

VOL. 3, PART 1. 



98 ERNEST WARL'EN. 

and gonophore is more granular than the rest of the endoderm. 
The outer surface of the ectoderm of the embrj'o (fig. 17, ect.^), 
is not sliai'ply marked off from the placental tissue [pi. t.), 
and there appears to be the most intimate contact. Both 
rounded and stellate cells of dense protoplasm are seen in this 
placental tissue. Between the endoderm {end.i) and the 
ectoderm {ect.i) of the embryo a faint line of division can be 
seen, and this is the beginning of the mesogkea. 

Such is the appearance of the embryo in a section cut in 
the plane of flattening of the gonangium. In a vertical 
section at right angles to this plane the planula bears a 
different aspect. The sides of the embryo are squeezed flat 
against the perisarc of the gonotheca with no gonangial 
ectoderm between (fig. 18). Laterally the embryo is very 
thin-walled, and both the ectoderm and endoderm layers are 
one cell thick, while on the other hand these layers are two 
or more cells thick at the distal and proximal ends. The 
blastocccl {Bl.) is roughly rectangular in section. 

Thus the embryo at this stage is of a curious shape. It is 
thick-walled everywhere (fig. 19), except laterally (fig. 18), 
over the flattened sides of the gonotheca, where the walls are 
very thin. 

The material has not permitted an examination of a later 
stage, but it is probable that the planula in the above con- 
dition is nearly ready to burst out of the gonotheca, for the 
gonotheca which Avas immediately below, and therefore 
slightly older, was empty, and the operculum absent, doubt- 
less through the bursting out of the planula. 

Summary. — Here we have an interesting case of the 
nourishing of the developing embryo. In the great majority 
of Invertebrata the ovum becomes charged with sufficient food 
or yolk-material to enable developinent to proceed far enough 
for the young creature to be able to provide itself with food. 
In the present hydroid the egg remains quite small and is 
never pi^ovided with a perceptible quantity of yolk, but it 
segments, and development takes place, in a kind of maternal 
placental tissue which supplies the embryo with food during 



DEVELOPMENT OF PLANULA IN A PLUMTTLARrAX PTYDlfOIO. 00 

the whole development. This phicental tissue arises as a 
modification of a specialised portion of the ectoderm of the 
blastostyle. This portion, as we have seen, forms a kind of 
cap over the young ovum, and doubtless may be regarded as 
representing the manubrial ectodei-m of a rudimentary gono- 
phore which bears one egg. 



EXPLANATION OF PLATE VI, 

Illustrating Dr. Ernest Wnrren's paper, "On the Develop- 
ment of the Planula in a certain Species of Plumularian 
Hydroid." 

Explanatory References for Plate and Text-figures. 

B. Blastostyle. Bl. Blastoewl. h. G. Beginning of gouaugium. 
C. Ccelenteron. c. j). Covering plate (" Deckenplatte '") of ectoderm of 
gouangium. c. 0. Cavity above ovum, wliicli may be regarded as a 
cavity between the manubrial ectoderm and endoderm of gonophore. 
c. U. Cavity of umbrella, d. C. Diverticukim of cceleuteron representing 
tlie endoderm of gonophore. d.jj. Dissolved perisarc. e. B. Ectoderm of 
1 »lastostyle. e. G. Scattered ectoderm cells of gonophore. e. E. Outer 
enveloping ectoderm derived from the scattered ectoderm cells of gono- 
phore. c. I. External ectoderm layer of typical gonophore. e. I. U. Ecto- 
derm layer of imibrella. ect.^. Ectoderm of planiila. end. Endoderm of 
blastostyle and of gonophore. end^. Endoderm of planula. (j.e. Granular 
endoderm. i. I. Inner or manubrial ectoderm layer of gonophore. M. 
Morula. M. e. Manubrial ectoderm, n., n. Oblique nodes at base of 
piunate portion of main-stem, n.^ i;., Transverse nodes at base of pinna. 
n..2, n..2 Transverse nodes at the base of pinnule. 0. Ovum, o. I. Outer 
layer of gonophore, equivalent to the layer of ectoderm lining the um- 
brella-cavity, ojy. Opercuhim. j;. e. c. Posterior ectodermal space of 
liydrocaulus. p. i. j). Proxiuial internode of pinnule. 2d- ^- Placental 
tissue. S(j. C. Segmentation-cavity, s. j). Space between the external 
ectoderm layer of gonophore and the ectoderm of the umbrella-cavity, 
.s^).,, sj)... Spaces between perisarc of gonotheca and blastostyle. sp. c. 
Spermatic cells in endoderm of hydi-ocanlus. t. Spermatic cells (testis) 
of male gouaugium. t. p. Terminal jjortion of the cceleuteron of l)lasto- 
style. V. m. Meuiln-ane around ovum. 

Fig. 1. — X -20. Longitudinal section throtigli the niain-steiu imme- 
diately below a hydrotheca showing the ])egiuniiig of the gouaugium 



100 ERNEST WARRKN. 

(h. G.). the siipei-iniposed attenuated perisarc {d. p.), and the youngest 
observed ovum (0.) embedded in the endoderm. 

Fig. 2. — x 220. Lono-itndinal section tlirough a young gonangium 
showing the ovum (0.) being thrust out into a special thickening of tlie 
ectoderm, whieli early becomes differentiated into an outer io. 1.) and an 
inner layer (/. /.) and represents the ectodermal portion of a reduced 
gonophore. The slight space between the two layers is equivalent to 
the uml)rella-cavity of a medusa. 

Fig. 3. — X 260. Transverse section through a young gonangium, 
showing the ovum surrounded by the ectoderm of the gonophore. sp. is 
the space which arises between the outer ectodei'm of the gonophoi'e and 
the ectoderm of the reduced umbrella-cavity. 

Fig. 4:. — x 220. Longitudinal section through a slightly older 
gonangium. The gonophore, consisting entirely of ectoderm, is more 
developed. The outer layer (o. I.) is now well-defined, while the inner 
layer {i. I.) closely surrounds the ovum (0.). The ectoderm of the gonan- 
gium at the distal surface has thickened considerably and forms the 
"covering cap" {c.f).). The space {sp.) is clearly seen. Spaces (sp.,,sp..,) 
are appearing l^etween the pei-isarc of the gonotheca and the general 
ectoderm. The endoderm immediately under the gonophore ectoderm 
is inoi"e granular than the remainder of that layer. 

Fig. 5. — X 220. Longitudinal section through an older gonangium. 
The outer layer of ectoderm (o. Z.) has become more fiattened through 
pressure exerted l)y the expanding inner layer (/. /.). Beneath the gono- 
phore the endoderm is beginning to grow and to form a diverticulum 
of the ccelenteron which i-epresents the ccelenterou cavity of the reduced 
gonophore. 

Fig. 6. — x 220. In this older stage the longitudinal section shows a 
well-defined space (c. 0.) between the ovum and the ectoderm {i. 1.) of the 
gonophore. The ovum is svirrounded by a membrane which is more 
marked externally than on the inner surface, which is in contact with 
the endoderm. 

Fig. 7. — X 220. Longitudinal section of gonangium at right angles 
to the plane of flattening. Here the cavity (c. 0.) of the gonophore has 
expanded, and the meml)rane around tlie ovum (r. vi.) is thickened. 

Fig. 8. — X 220. Longitudinal section of gonangium in the plane of 
flattening. This is in the same stage of growth as in fig. 7. 

Fig. 9. — X 220. Longitudinal section at a, later stage. The ectoderm 
layer of the gonophore has broken down (e. G.), and forms an irregular 
mass of tissue around the ovum. The endoderm below is granular and 
in an active condition of growth. The cells of the distally placed 



iil:vi<:l()pment of planula in a pi-umulaimax iiydiiMhh. lol 

eududerm and those of tlie " coveriug cap "' have becouie very 
elongated and attenuated. 

Fig. 1(1. — X 220. Longitudinal section at the next stage. The distal 
endoderm and the ''covering cap" have completely disappeared and tlie 
ovum is surrounded by cells which have arisen from the scattered ecto- 
derm of the broken-down gonophore. The ovum is being pushed into 
a central position by the growth of the diverticulum of the coilenteron 
(d. C.) which is the endodermal spadix of the gonophore. 

Fig. 11. — X 220. Longitudinal section showing embi-yo (M.) derived 
from the segmentation of the ovum. It consists of about sixteen cells, 
and is central in position immediately below the middle point of the 
distal surface of the perisarc of the gonotheca. The embryo is sur- 
rounded by the placental tissue. The diverticulum of the ccelenteron 
(d. C.) or gonophore spadix has become equal in size to the terminal 
portion {t. p.) of the ccelenteron of the gonangium. 

Fig. 12. — x 480. Enlarged view of morula imbedded in placental 
tissue {pi. t.). 

Fig. 13. — x 220. Later stage. The eml>ryo. the placental tissue, 
and the ccelenteron pouches have grown. The embryo has developed a 
segmentation space (sg. C). 

Fig. \i. — X 220. In the next stage, a differentiation of the cells of 
the emljryo into an outer ectoderm and an inner endoderm is seen. 
Distally there is only ectoderm {ect.^) which is very thin ; proximally the 
ectoderm is thicker and is more than one cell thick. 

Fig. 15. — x 220. The emliryo becomes pear-shaped, and the endo- 
derm is not continued distally. The maternal ectoderm (e E.), derived 
oi'iginally ivom. the placental tissue, is continued around the outside of 
the coilenteron pouches, and the cells l)ecome very elongated and reach 
the lateral perisarc of tlie gonotheca. 

Fig. 16. — x 220. A later stage where the embryo has grown con- 
siderably and has extended inwards towards the centre of the gonangium. 
The ccelenteron pouches, consisting, on one side, of the spadix of the 
gonophore, and on the other, of the terminal portion of the ccjelenteron of 
the blastostyle, form a deep cup containing the embryo and the true 
placental tissue {pi. t.). The enveloping ectoderm (c E.) has also 
greatly extended. Bl. is blastocrel. The endoderm {g. e.) of the 
gonangium at the base of the embryo is more granular than elsewhere. 
The endoderm of the embryo now forms a contimious layer. It is 
mostly more than one cell thick, as also is tlie outer ectoderm. 

Fig. 17.^ — X 140. Enlarged view of a piece of embryo, the surround- 
ing placental tissue and the endoderm of gonangium. The ectoderm 



102 ERNEST WAUKEN. 

(cci.j) of the embryo is in intimate contact with the placental tissue 
ipl. t). 

Fig. 18. - X 220. Longitudinal section of gonangium at right angles 
to the plane of flattening. The planula is seen wedged between the 
perisarc of the gonotheca. The wall of the planula is very thin laterally. 
Init much thicker proximally and distally. 

Fig. 19. — x 220. The planula in section in the plane of flattening 
of the gonotheca. The wall of the planula is of considei'able thickness 
all round. Both the endoderm (f»fZ.,) and ectoderm (cct.-^) are several 
cells in thickness. 



Ann. Natal Mus Vol, HI. 




\ '''' ^0* 








# 



9 X280 








~'*s»% c. ■■' /#^ — « 



14 X 220. 



17.-* **o. 



A TERMITOPHILOITS P.Kpyrf-K OK 'I'HK GENUS COBOTOCA. 103 



Note on the Occurrence in South Africa of a 
Termitophilous Beetle of the Genus Corotoca. 

By 



Among the most remarkable and interesting of the tei-mito- 
philous insects are the beetk's of the genei\a Spirachtha 
and Corotoca, belonging to the family Staphy linida?. 
They have a much swollen abdomen, with the greater portion 
of it permanently turned forwards over the dorsal surface of 
the thorax. It extends as far as the middle of the pronotum. 
These beetles were first described and figured by J. C. 
Schiodte in 1854 from specimens obtained from termite nests 
in Brazil. 

In 1905, Dr. Ivar Triigardh, of the Upsala University, 
visited South Africa on a collecting trip, and he discovered in 
the nest of a species of Eutermes, in a limited area in Zulu- 
land, a great number of specimens of anew Staphylinid, which 
he has called Termitomimus. This genus is characterised 
by the second and third segments of the abdomen being very 
greatly swollen, and the remainder of the abdomen, which is 
somewhat less swollen, is turned forward over the thorax. 

Dr. Tragardh examined many nests of termites in Natal, 
but no other physogastric Staphylinids were discovered. 

Recently, numerous nests of the common termite Eutermes 
trinervius {Ramhur) have been examined in another connQc- 
tion by Dr. Conrad Akerman and myself in the neighbourhood 
of Pietermaritzburg, and in March, 1913, Dr. Akerman found 
in one of the nests a single specimen of a physogastric 
Staphylinid. Subsequent examination proved that it un- 



104 ERNEST WARREN. 

doubtedly belongs to the g-onus Corotoca, and is closely 
allied to^ altliongli specifically distinct from, Core toe a 
m el ant ho ScJ dixit e of Brazil. 

Since this discovery^ about 120 nests in different localities 
have been examined with a view to obtaining more material 
and ascertaining the life-history of the species, but only three 
other speciiiiens have been found. 

In certain nests a beetle-lai"va with some resemblance to 
the larva figured by Schiodte as being that of Corotoca 
melantho has been found in fair numbers, but all attempts 
to rear it have proved fruitless. The termite Eutermes 
trinervius does not live for any length of time under arti- 
ficial conditions, and the beetle-larva is even more delicate. 
The antennio and mouth-parts are quite like those of the 
larva of C. melantho; the abdominal segments also bear 
lateral processes, but in the present species they carry bristles. 
On the whole it seems probable that the larva is that of the 
Corotoca, but it is remarkable that it is comparatively 
common, while the beetle appears to be excessively rare, and 
this rarity explains the circumstance that apparently it has 
not been previously found. A possible explanation is that the 
termites interfere injuriously with the beetle when in the pupal 
condition, so that only a very few ever arrive at the perfect 
state. 

It is hoped that when further material has been collected, 
a comparison of the anatomy of this genus may be made with 
that of Termitomimus entendvcniensis Tri'ig., which 
was carefully described and figured by Dr. Triigardh^ in 
1907. 

The occurrence of this highly specialised termitophilous 
genus Corotoca both in Brazil and South Africa is interesting 
in indicating its antiquity, and in connection with the fact 
that the similarly modified termitophilous dipteron, Termi- 
toxenia, is found both in South Africa and India. 

' TriigHrdh, Ivav, ' Description of Termitomimus, a New Genus of 
termitophilous physogastric Aleocharini, witli Notes on its Anatomy," 
Upsala, 1907. 



A TEKMITOPIIILOUS BEETLE OF THE GMNUS COHOTOCA. 105 

J. S. Schiodte^ has given a careful definition of the genus 
Corotoca, and also concise diagnoses of two species, C. 
melantho and C. phylo, both from Brazil. 

These diagnoses are quoted below in order that they niay 
be compared with the brief description of the new species 
which is now given, pending a detailed account of the anatomy 
of the insect, which it is proposed to publish hiter. 

I have pleasure in associating this interesting species with 
its discoverer, Dr. Conrad Akerman. 

"1. Corotoca melantho SchvkUe. 

" Fusca, fronte foveolata, pronoto multifoveolato, disco 
bituberculato : tibiis posterioribus fusiformibus, fuscis : scutis 
ventralibus segmenti quarti (juiiiticjue transversis. — Mas, 
Fern. 

" Long, a fronte ad apicem segmenti secundi abdominis, 
2^-3 millim. 

"2. Corotoca phylo Schiodte. 

" Fusca, vertice foveolato, pronoto multifoveolata-, disco 
trituberculato : tibiis posterioribus linearibus, nigrofuscis : 
scutis ventralibus segmenti quarti quintique subquadratis. — 
Fem. 

" Long, a fronte ad apicem segmenti secundi abdominis, 
2^-3i millim." 

3, Corotoca akermani s-p. n. 
Brown, forehead or vertex not pitted, occiput Hat or 
slightly concave, not tuberculated ; pronotum smooth, except 
for a very shallow and rather wide and inconspicuous median 
groove extending along the postei'ior three-quarters of its 
length ; posterior tibia) rather slender, straight or slightly 
curved, not fusiform, but rod-shaped ; sternites of the third, 
fourth, fifth, sixth and seventh abdominal segments appear as 
five isolated, uniform, narrow, transverse, brown strips of chitin 

' Schiodte, J. C. ' Corott)ca og Spiraclitha," Kj^beuhavn. 1854. pp. 
8-9. 

VOL. 3, PART 1. 8 



106 EIINES'I' WARREN. 

on the upper surface, all of which are widely separated from one 
another by intersegmental membranes caused by the swelling 
of the abdomen ; the morphologically dorsal surface of the 
abdomen rather deeply concave, fitting over the thoracic 
region, which is not flattened or otherwise much modified. 

Length from forehead to hinder end of upturned abdomen, 
3 '2-8 "6 mm. 

Loc. — Environs of Pieterniaritzburg : Durban Road (Dr. 
Akerman, March, 1913) ; Botanical Gardens (W. Culling- 
worth, July, 1913) ; Sweetwaters Road (July, 1913) ; Bisley 
Road (December, 1913). 



Vol. Ill, part 2, issued October 22nd, 1916. 

TKE CARNIVOROUS SLUGS OF SOUTH AFRICA. 



107 



Studies on 
The Carnivorous Slugs of South Africa, 



INCLUDING 



A Monograph on the Genus Apera, and a Discussion on 

the Phylogeny of the Aperidae, Testacellidse, and 

other Agnathous Pulmonata. 

By 
Ilii^h \l'at«»oii, ITI.A. 



With Plates VII-XXIV. and 7 text-figures. 



Contents. 

Introduction ...■•• 
The History ob^ the Genus Apera 
A Description of the Genus Apera 

External Characters . . . • • 

Tlie Structure of the Skin .... 

The Shell, the Mantle-cavity, and the Mantle 
The Pedal Gland ..... 

The Nervous System . . . • • 

The Structure of the Upper Tentacles, and the Tentacular 
Retractors . . ■ ■ 

The Digestive System . . . . • 

The Vascular System ..... 
The Excretory System . . . • ■ 

The Reproductive Organs . . . • 

Distribution ....'• 
The Differentiai, Characters of the Species and Sub- 
species of Apera . 
Apera gibbonsi (Binney) 

rubella n. suhsp. 
„ gracilis n. suhsp. 

hipata n. stihsp. 

VOL. 3, part 2. 




PAGE 

. 108 
. 109 
. 115 
. 115 
. 124 
. 127 
. 131 
. 136 

153 
155 
181 

186 
188 
193 

193 
193 
197 
199 

200 



108 HUGH WATSON. 

PAGE 

Apera parva n. sp. . . . . . 202 

Apei'a dimidia n. sp. . . . . 204 

Apera piircelli CoUinge .... 207 

Ajjera bui-nupi Smith . .210 

Apera sexangula n. sp. .... 213 

The Phylogeny op the Species of Apera . . 217 

The Occurrence at Cape Town of Testacella maugei 

F<'r. . ■ .220 

The Affinities of Apera and Test.4CELLA . . 222 

The Evolution of Carnivorous Characters . . . 222 

The Phylogeny of Apera and other naked Carnivorous Slugs 231 
The Phylogeny of Testacella and its possible Allies . 238 

Appendix — Bibliography of Testacella maugei Ft'r. . 256 

Explanation ok Plates ..... 258 

INTRODUCTION. 

Many snails and slugs of great interest inhabit South 
Africa, but very little is known about their anatomy. I have 
therefore been asked to describe some of these animals, 
beginning with those in which the shell is degenerate or 
absent. The present article deals with the slugs belonging to 
the carnivorous gi'oup, and principally with those contained 
in the genus Apera ; and as this genus is a very remarkable 
one and seems to be confined to South Africa, I am endeavour- 
ing to describe it in some detail. Unfortunately, however, 
the material at uiy disposal has been limited to a small 
number of pi-eserved specimens, some of which have proved 
to be immature. More than half of these specimens were 
collected and sent to me by Mr. H. C. Bnruup, of Pieter- 
maritzburg, to whom I am very deeply indebted. I am also 
glad to have this opportunity of expressing my thanks to the 
Rev. Prof. H. M. Gwatkin, to Dr. Doncaster of Cambridge, 
Dr. Peringuey of Cape Town, Dr. Pilsbry of Philadelphia, 
Dr. Simroth of Leipzig, and Dr. Warren of Pietermaritzburg, 
to Major M. Connolly, and to Messrs. Keppel H. Barnard, 
Walter E. Collinge, John Farquhar, John H. Ponsonby, (1. C. 
Robson, Edgar A. Smith, and William Tains. 



TPIE CARNIVOROUS SLUGS OF SOUTH AFRICA. 109 

TRK HISTORY OF THE GENUS APERA. 

Neai'ly forty years ago, when Mr. J. S. Gibbons was visit- 
ing* Natal, he found a sti'ange shig- under a stone in a wood. 
This .-inimal puzzled him greatly ; it seemed more like a 
Testacella than anything else he knew ; yet clearly it was 
not a Testacella, for where the shell should have been 
there was merely a little hole — -the respiratory orifice.^ 
Gibbons therefore sent the slug to Mr. VV. G. Binney, with a 
description of its appearance when alive. Binney. examined 
its radula, and found that its teeth were like those of 
Glandina. He also discovered that it had no jaw, but that 
it possessed an internal shell, Avhich was broken when he 
found it, but which he wrongly assumed to be hexagonal 
when perfect. And Binney ^ established a new genus for this 
peculiar animal, and published a description of it in 1879, 
together w;th Gibbons's account of its appearance when alive, 
a rough drawing of the animal, and a figure of its radula. 
And he named the slug Chlamydephor us gibbonsi. 

In 1884 Binney'^ repeated his description of the radula of 
this slug and reproduced his figure. In the same year Tryon •■ 
reproduced the rough drawing of the animal, and gave a brief 
description of the genus, which he placed in the Testacel- 
lidas; and he also amended the spelling of the generic name, 
changing it to Chlamy dophorus. In the following year, 
however, he reverted to the original sjjelling, when he des- 
cribed the genus and species in the 'Manual of Conchology,' ^ 
and again reproduced the rough drawing of the animal. 

Now Binney named the genus Chlamydephor us because 
he said that the mantle covered the whole back. But in this 
Binney was wrong, as Heynemanu '' was the first to point out, 

' Vide Collinge, W. E., ' Ami. Natal Mas.," 1910. vol. ii, p. 164. 
- ■ Bull. Mus. Comp. Zool. Harvard,' vol. v, p. 331, pi. ii, figs, a, 1). 
^ ■' Notes on the Jaw and Lingual Dentition of Pulmonate Mol- 
lusks." • Ann. N. Y. Acad. Sci.,' vol. iii, p. 81, pi. xvii, fig. a. 
^ • Struct, and Syst. Conch.." vol. iii. p. 13, pi. ci, fig. 47. 
^ Vol. i (2nd ser.). jjp. 7. 17. pi. ii, fig. 95. 
'"' • Jahrb. d. Deutsch. Mai. Ges.,' vol. xii. p. 17, ])]. ii. figs. 5-7. 



110 HUGH WATSON. 

when, in 1885, he published a description and three figures of 
the external characters of a specimen in the British Museum, 
labelled " Cape Colony." He thought that the animal was 
nearly related to Testacella ; and as he showed that it had 
no mantle on its back, he suggested that the generic name 
should be changed to A per a. 

In 1890 Prof. T. D. A. Cockerell ^ published a short descrip- 
tion with measurements of what appears to be the same 
specimen, and he said that Heynemann's name would pro- 
bably have to be used, as Chlamydophorus (differing from 
Binney's name only by one letter) had been employed hj 
Harlan for a genus of Mammalia in 1825. 

In 1892 Mr. E. A. vSmith - described the external characters 
of a new species of A p e r a , very different f I'om A . g i b b o n s i , 
which Mr. H. C. Burnup had found in Natal, and he named 
it Apera burnupi. He also thought that Apera was the 
best name to use, as Chl'.imydophorus had been employed 
in the Mammalia, but he attributed the latter name to Agassiz 
{' Nomenclator Zoologicus, Mammals,' 1842, p. 8), instead of to 
Harlan. Prof. Cockerell ^ then discovered that Apera was 
preoccupied in botany, and therefore thought that it might 
be better to use Binney's name with the original spell- 
ing. He placed the genus in the Testacellidog. On the 
other hand. Dr. Simroth,^ after studying Heynemann's article, 
suggested that the genus might be allied to the Janellida?, 
its carnivorous characters having arisen independently of 
those of Testacella. He thought that the fragmentary 
condition of the shell, as found by Binney, might be natural ; 
but had difficulty in explaining how it was that Apera had 
two pairs of tentacles and Janella only one. In 1895 
Apera and Clilamy de phorus were treated as though 
they were separate genera by Dr. A. H. Cooke.'' 

1 -Ann. Mug. Nat. Hist." (6th ser.). vol. vi. p. 390. 

2 'Ann. Mag. Nat. Hist.' (6th ser.), vol. x. p. 466. 
•■> ' Conchologist; 1893, vol. ii, p. 206. 

■* ' Nacktschnecken, Deutscli-Ost-Afrika.' 189.5. ex. vol. iv, p. 20. 
•' • The Cambridge Natural History,' vol. iii. " Molluscs and Brachio- 
pods,' pp. 333, 334, 440. 



THE CAKNIVOKOUS SLUGS UF SOUIJI AFlilCA. Ill 

Two years later Mr. W. E. Colling-e ^ described and figured 
the alimentary canal, pedal gland, and reproductive organs 
of a specimen of Apera which he had receiv^ed from 
Mr. E. A. Smith. Unfortunately this specimen was not in 
good condition for dissection, and Collinge's description is 
uiaccurate in several respects, and his figures bear but little 
resemblance to the natural condition of the organs that they 
are intended to portray. Moreover the specimen was named 
Apera burnupi, whereas it really belonged to a quite 
distinct species. Collinge stated that he considered that 
Apera had affinities with the Testacellida?, but he thought 
that there was a wide gap between it and either Testacella 
or iJaudebardia, and a still wider gap between Apera aud 
ISchizoglossa. in the following year Dr. Pilsbry published 
a short review of Collinge's article in the ' Nautilus.' ~ 

In 18U9 i-'aul and Fritz Sarasin'' brought forward the theory 
that Apera was aucestral to Atopus. Now Atopus has a 
large mantle covering the whole body, and the brothers 
JSarasin therefore rejected Heyneuiann's interpretation of the 
morphology of Apera, and advocated the use of Binney's 
name Chlamy dephorus. 

In 1900 Collinge* received through Mr. J. H. Ponsonby an 
example of Apera burnupi Smith, found by the Kev. J. K. 
Ward at Kichmond, JS'atal, and belonging to the South 
African Museum, Cape Town. He at once saw that this 
specimen differed from the species which he regarded as 
Apera burnupi, and accordingly he named it Apera 
natalensis n. isp. He made a superficial dissection of some 
of the internal organs, but the specimen does not appear to 
have been m a much better condition than that of the other 
species which he had previously dissected, and his figures of 
its anatomy are in consequence equally faulty. In the same 

' -Ann. Mug. Nat. Hist." (6th aev.), 1897, vol. .xx. pp. 1:21-225, pi. v. 
tigs. 1-6. 

- Vol. xh, p. 12. 

•' ' Die Laiid-Mollusken von Celebes,' p. 112. 

^ ■ Ann. S. Air. Mus.," vol. ii, pp. 'S-b. pi. i, tigs. 3, 4. pi. ii. tigs. 14.. 15 



112 HUGH WATSON. 

paper Colliiige publij^lied two coloured figures of Ijotli species 
by Mr. F. J. Partridge. 

In the following year CoUinge ^ had an opportunity of 
examining a better specimen of the species which he supposed 
to be Apera burnnpi, and he discovered how misleading 
were his former figures of its anatomy. He therefore 
published a more accurate figure and description of the 
reproductive organs of that species. At the same time he 
criticised the theory of P. and F. Sarasin, and said that xA-pera 
was probably closely allied to the Testacellidte, Avhile 
A top us on the other hand seemed to be far removed from 
that family. 

In December, 1901, Collinge ~ described the external 
features of a new species of Apera found by Mr. B. j\l. 
Lightfoot on the slopes of Table Mountain, Cape Town, and 
published coloured figures of it by Partridge. This species 
he named Apera puree Hi. 

Dr. Simroth,'^ in the same month, published a copy of 
Collin ge's figure of Apera natalensis (:= A. burnupi 
iSmitJt), in his interesting article on carnivorous snails and 
slugs. In this paper he unfortunately stated that Apera 
was without a shell, and again suggested that the genus 
might have been derived from the Jan ell id a3, or possibly 
from some other ancient family of herbivorous slugs found in 
the Southern Hemisphere, although he admitted that too little 
was known of the genus to give any value to his hypo- 
thesis. 

In 1902 Collinge^ reproduced on a larger scale his second 
figure of the reproductive organs of the species which he 
erroneously supposed to be Apera burnupi. In the fol- 
lowing year Dr. von Moellendorff ^ removed the genus from the 
Testacellida?, and placed it in a family by itself, which he 

' ' Journ. of Malac' vol. viii. pp. 71, 72. 

2 ' Ann. S. Afr. Mus.,' vol. ii. p. 230, pi. xiv, figs. 1. L'. 

•' ' Naturwiss. Woclienscbnft.* vol. xvii, p. 111. fig. 7. 

"* ' Joiirn. of Malac.,' vol. ix. pi. vi. fig. 66. 

'" ' Conch. -Cabinet (Agnatlia).' p. 5. 



TaE CARNIVOROUS SLUGS OF SOUTH AFRICA. 113 

named the Aperidai. In 1906 Heynemann^ devoted a para- 
graph to the genus in his article on the geographical distri- 
bution of slugs. In 1907 Dr. Pilsbry - followed Moellendorii's 
example aud placed the genus in the family Ape ridte, which 
he included in his super-family Agnathomorpha; and he 
suggested that it might prove to be more nearly related to 
the Rhytididas than to the other carnivorous families. 

In 1909 and 1912 Simroth-^ included in Bronn's 'Tier-Reich' 
some of the information given in Collinge's articles, and 
published copies of four of the figures which had illustrated 
his first paper in the ' Annals of the South African Museum.' 

In 1910 Collinge * gave a short account of the external 
features of the three species of Apera then known to inhabit 
Natal, together with a definition of the genus, and some 
remarks on its habits, distribution, afiinities, and history. 
This paper contains valuable quotations from letters of 
Gibbons and Burnup, but the error about Apera burnupi is 
repeated. In treating of the affinities of the genus, Collinge 
again stated that he considered that P. and F. Sarasin were 
wrong in regarding Apera as in any way related to A topus; 
but he now expressed the opinion that it was not related to 
Testacella either, and thought that it should probably 
stand by itself as a remnant of some very ancient stock of 
carnivorous Pulmonates. Accordingly he placed it in a 
separate family, which he termed the Aperaida? fain, iiov., 
being apparently ignorant of the classification of both 
Moellendorff and Pilsbry. 

Lastly, in 1912, Major Connolly Mncluded Apera in his 
valuable " Reference List of South African Non-marine Mol- 
lusca," following Collinge's nomenclature of the species. 

Such is the story of the treatment that Apera has received, 

' ■ Abhandl. Her. v. d. Senkenb. Naturf . Gesell. (Frankfurt)," vol. 
XXX, p. 23. 

- • Manual of Conchology ' (2nd ser.), vol. xix. pp. ix, xi. 

^ ' Klassen und Ordnungen des Tier-Reichs III. Gaistropoda Pul- 
nionata,' pp. 143, 611, 612, fig. 42c, pi. iv, figs. 9-11. 

* "Ann. Natal Mus.," vol. ii, pp. 163-170. 

■' • Ann. S. Afi-. Mus.," vol. xi, pp. 62, 63. 



114 HUGH WATSON. 

and it is largely a stoiy of mistakes. It will have been seen 
that four species have been described, three from Natal and 
one from the Cape, but that very little is known about the 
anatomy of any of these, as only two reliable figures have 
hitherto been published of their internal features, namely, 
Binney's figure of the radula of A per a gibbon si, and 
Collinge's second figure of the reproductive organs of the 
species which he supposed to be A. burnupi. Nevertheless 
this ignorance does not seem to have prevented malacologists 
from speculating about the affinities of the genus ; for, as we 
have seen, it has been suggested in turn that it may be related 
to the Test ac ell idte, to the Janellidi^, to A top us, or to 
the Rhytididge, forms which differ immensely from one 
another. It is hoped that the information contained in the 
present article may give greater value to such speculations in 
the future. 

In the following pages I am attempting to describe this 
remarkable genus in as great detail as the limited number of 
specimens at my disposal will permit, giving numerous figures 
of its anatomy. Secondly, I am giving a concise description 
of the distinctive characters — both external and internal — of 
each species; that is to say, of Apera gibbonsi, A. pur- 
celli, A. burnupi, the species that Collinge erroneously 
supposed to be A. burnupi, which I am naming A. sex- 
angula, and two new species — A. dimidia from Natal, and 
A. parva from Gi'ahamstown. And at the same time I am 
pointing out that there are some very distinct races of A. 
gibbonsi, which may also prove to be separate species, but 
which I am ])rovisionally regarding as sub-species until more 
material is obtainable. After dealing with the relation of 
these forms to one another, I am describing briefly the species 
of Test ace 11 a that occurs at the Cape, and am then dis- 
cussing at some length the probable relations of Apera and 
Testacella to each other and to the remaining genera of 
carnivorous snails and slug's. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 115 

A DESCRIPTION OF THE GENUS APERA. 

EXTERNAL CHARACTERS. 

General Shape. — The slugs cuntaiued in the genus Apera 
are subcylindrical, tapering gradually towards the head and 
more abruptly at the hind end, the broadest region being 
usually behind the middle of the animal. The height is equal 
to the breadth or sometimes slightly less, while the width of the 
foot varies from three-fifths to scarcely one-half of the greatest 
breadth of the body. Most of the species are rather slender, 
especially A. parva and some forms of A. gibbon si; and it 
is interesting to notice that the latter do not contract when 
about to rest, after the usual manner, but merely bend their 
bodies at various angles, in which state, Mr. Burnup informs 
me, they much resemble the rhizomes of feims. 

Size. — So few specimens of Apera have been examined that 
the mean dimensions attained by each species cannot be stated 
Avith accuracy. It seems probable, however, that the average 
length of each of the three larger species — A. gibbon si, A. 
burnupi and A. sexangula — lies between 60 and 65 mm., in 
the case of specimens preserved in alcohol, although occasional 
examples may reach a length of nearly 80 mm. On the other 
hand adult specimens of A. dimidia preserved in spirit are 
only about 33 mm. long, and A. parva and A. purcelli 
seem to be still smaller species. 

Tentacles. — As usual in the Stylommatophora, there are 
two pairs of tentacles, the upper pair being retractile and 
bearing the eyes.^ The lower tentacles are small, and beneath 
them there is another pair of short processes of a paler colour, 
which doubtless correspond to the laljial feelers found in luost 
of the carnivorous genera. 

Genital Opening. — The common opening of the reproductive 
organs is on the right side of the head, two or three milli- 
metres behind the right lower tentacle, and about half that 
distance from the margin of the foot. 

' See pp. 153, 154 for structural details. 



116 HUGH WATSON. 

Respiratory Opening. — The aperture of the mantle-cavity 
or " respiratory orifice " is situated towards the hind end of 
the back, just to the right of the mid-dorsal line. Its posterior 
position, about four times as far from the head as from the 
hind end of the slug, is due to the lengthening of the anterior 
part of the body and the retrogression of the pallial organs,, 
as in Selenochlamys and Testacella. The dorsal posi- 
tion of the opening is explained by the absence of any external 
mantle or shield in Apera.^ 

The orifice is alwaj-s a little nearer the right side than the 
left, a fact which previous writers seem to have overlooked, 
although it is of considerable morphological importance, as 
may be seen from PL X, fig. 33 (a transverse section through 
the respiratory opening of Apera gibbonsi rubella). This 
illustration also shows that the opening is surrounded b_v two 
lips, one within the other. In A p e r a gibbonsi and A . p a r v a 
the outer lip projects over the inner lip so as almost to con- 
ceal it; but in the other four species the outer lip is much 
narrower, and leaves a wide circular opening, within which 
the light-coloured inner lip is exposed to view (PI. A^II, fig. 
12; PI. VIII, figs. 24, 25, 26). The right anterior edge of 
the inner lip is cleft by an oblique slit, running forwards and 
outwards from the central aperture. 

Keels. — In most of the species of Apera the back is 
rounded, but A. burnupi and A. sexangula both have four 
longitudinal keels, two on each side of the body, which may 
possibly serve to increase the rigidity of the skin. The upper 
keels of A. sexangula are nearly parallel throughout the 
greater part of their length, the distance between them measur- 
ing about three-fifths of the breadth of the body. When followed 
backwards, however, they diverge very slightly just in front 
of the respiratory opening, and then gradually approach each 
other behind it, until they unite to form a single median keel,. 
3 or 4 mm. in length, at the hind end of the animal. In A. 
burnupi the upper keels are further apart, and become very 
prominent towards the hind end, divei'ging considerably and 
' See pp. 130. V.n. 



THE CARNIVOliOUS SLUGS OF SOUTH AFRICA. 117" 

then curving round to meet in a very obtuse angle just 
above the posterior extremity of the foot. 

The lower keels of Apera sexangula extend on each side 
along the whole length of the animal, about half-way between 
the upper keels and the edges of the foot. The slug is thus 
roughly hexagonal in transverse section, but when it con- 
tracts the surfaces between the keels become concave. In A . 
burn up i the lower keels are much nearer the foot than in 
A. sexangula, and become obsolete towards the hind end of 
the slug. 

The other species of i^peraare without any traces of either 
median or lateral keels. They are bluntly pointed at the 
posterior end. 

There is no caudal mucous pore in Apera. 

Dermal GtROGVes. — In all the species of Apera the back 
and sides of the animal are covered Avith a network of grooves, 
dividing the skin into numerous polygonal rugas. These 
grooves are specially deep in A. burnupi. The centres of 
the ruga? are often raised in well-preserved specimens, and 
the skin has therefore a granular appearance. 

Certain of the grooves are larger than the others, and run 
in more definite directions, forming as it were the main 
channels from which branch the smaller grooves that form 
the network. First there are the two dorsal grooves which 
run along the middle of the back from the respiratory 
opening to the head. These are most strongly developed in 
Apera burnupi, and least conspicuous in A. dimidia, in 
which species the main grooves are scarcely more distinct 
than the other grooves of the network. The distance sepa- 
rating the dorsal grooves varies from about one-seventh of 
the breadth of the body in A. dimidia to less than half 
that proportion in some forms of A. gibbon si (PI. VII, 
fig. 4). In A. burnupi, A. sexangula, and A. pur- 
celli these grooves are separate throughout their entire 
length, but in A. gibbonsi, A. parva, and A. dimidia 
they are united posteriorly, and arise from the respiratory 
opening as a single groove which divides between 3 and 



nS HUGH WATSUN. 

5 inm. in front of the opening (PI. VII, figs. 1, 4 ; PI. VIII, 
figs. 22, 24). 

A single median posterior groove runs backwards from 
the orifice. It is usually deep and conspicuous in front, but 
breaks up into finer irregular grooves before reaching the 
hind end of the slug. 

Well-marked radial grooves diverge laterally from the 
region of the respiratory opening. About four of these on 
each side arise from the opening itself, and two or three others 
branch from the median posterior groove behind the opening. 
In Apera purcelli,the most anterior grooves, instead of 
arising from the respiratory opening, branch from the dorsal 
grooves in front of it (PL VIII, fig. 25). As a rule the radial 
grooves extend down the sides of the body, but in A. gib- 
bon si some of tliem usually bifurcate before reaching the 
edge of the foot. The most anterior grooves of A. gibbonsi, 
A. parva, and A. dimidia always divide within a short 
distance of the respiratory opening, the lower branch sloping 
downwards towards the foot, while the other continues forward. 
The latter branch soon divides again in a similar manner, 
and this process is repeated several times, so that an irregular 
lateral groove is formed, which runs forward parallel to 
the dorsal groove and gives off oblique sub-lateral grooves 
parallel to one another. The lateral grooves are seldom well 
developed in A. gibbonsi and A. dimidia, but the oblique 
sub-lateral grooves are rather conspicuous in the former 
species (PL VII, figs. 2, 3, 5, 6). On the other hand, in A. 
parva, A. burnupi, and A. sexangula the lateral grooves 
are more definite and better developed than the sub-lateral 
grooves, of which there are only a snui.ll number (PL VIII, 
fig. 2:3; PI VII, figs. 11, 13). The latei-al grooves of the two 
latter forms are lower down than those of the other species, 
and branch from the most anterior pair of radial grooves at 
a greater distance from the respiratory opening. They run 
along between the upper and lower keels as far as the head, 
the right lateral groove ending in the genital ojiening as in 
Testae el la and other forms. It is probable that the greater 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 119 

development and lower position of the lateral grooves in these- 
two species is due to the pi-esence of the keels. There ai'e 
no definite lateral grooves in A. })urcelli. 

The foot-fringe is bounded above by a peripodial' 
groove, which is usually rather deep (PI. XI, fig. 35). In 
Apera burnupi and A. sexangula there is a second 
groove, parallel to the peripodial groove, but sepai'ated from 
it by a narrow row of rugte, and the radial grooves do not 
extend below this upper groove, but terminate in it. As a 
rule the other species have only the single peripodial groove 
with Avhich the radial grooves unite, but in A. dimidia the 
upper limits of the lowest row of rugfe are sometimes arranged 
so as to form an irregular groove parallel to the peripodial 
groove on each side. 

'J'he foot-fringe itself is narrow and is crossed by a number 
of small vertical grooves, which are often continued across 
the edges of the foot-sole. In contracted specimens many of 
these transverse grooves extend right across the sole, althouo-h 
they are never so numerous in the middle as at the edges. A 
conspicuous but irregular longitudinal groove often runs 
along the centi-e of the foot-sole, but this also is probably due 
to the contraction of the specimens. In some examples two 
grooves diverge from a point at the hind end of the foot-sole 
and extend forward nearh^ parallel to the edges of the foot, 
thus dividing the hinder part of the sole into median and 
lateral areas. Occasionally these grooves may be traced 
forward for about two-thirds of the length of the animal, but 
they are more usually confined to the hind end, as shown in 
PI. VII, fig. 9, and in many specimens they are entirely absent. 
In Natalina I have sometimes found similar grooves towards 
the hind end of the foot ; but none of the grooves on the 
foot-sole seem to have so much systematic value as those on 
the back and sides of the slug, for they may be present or 
absent in different individuals of the same species. 

Colour. — The skin of Apera is coloured, and the colour is 
almost entirely due to pigment. It is true that specimens of 
the more darkly coloured species when preserved in spirit 



120 HUGH WATSON. 

sometimes have a slightly bluish or bliiish-green tinge, and 
that this is probably due not to pigment, but to minute 
particles in the outer layer of the skin which intercept tlie 
blue rays. It is doubtful, however, whether this interference 
effect Avould be as noticeable in living specimens. 

Apera possesses two chief kinds of pigment: the first is 
dark and relatively stable, the second is lighter, very unstable, 
and superposed upon the first. 

The dark pigment is usually brownish-grey, varying from 
dull brown in typical examples of Apera gibbon si to dark 
grey in A. purcelli. It does not spread uniformly over the 
back and sides of the animal, but is more concentrated in 
some parts than in others. In the first place the dark pigment 
tends to be aggregated into small irregular patches, which 
give the slug a mottled appearance. This tendency is least 
apparent in Apera purcelli, and most marked in A. 
dimidia, A. burnupi, and some varieties of A. gibbonsi. 
In these animals the patches of colour are well defined, so 
that the skin is conspicuously mottled. AVhen the dark 
patches are examined through a strong lens they are seen to 
-contain minute lighter specks, which are probably caused by 
the dermal mucous glands. 

Secondly, the dark pigment generally shows a tendency to 
become concentrated along the sides of the dermal grooves. 
Thus in A. gibbonsi the oblique sub-lateral grooves are 
usually rendered more conspicuous by the way in which the 
patches of pigment tend to coalesce along their courses. 

Thirdly, the dark pigment is more abundant on some 
regions of the body than on others. In Apera gibbonsi 
it is concentrated dorsally, although a lighter band is gene- 
rally left along the very centre of the back (PI. VIII, figs. 
14-19). On the sides of the animal the mottling becomes 
sparser, and it dies out before reaching the foot-fringe. 
In the keeled species, as well as in A. dimidia and A. 
purcelli, the dark pigment tends to be concentrated 
laterally, so as to form an irregular longitudinal band on 
each side of the bodv. Between the bands there is a lig-hter 



THE CAHNIVOBOUS SLUGS OF SOUTH AFRICA. 121 

dorsal area, palest at the edges, but a little darker in the 
middle; below them the colour shades off gradually towards 
the foot. This arrangement of the pigment is best developed 
in Apera dimidia (PI. VIII, figs. 20 and 21), but it is also 
fairly conspicuous in A. liurnupi, where the dark mottling 
is concentrated to form a band on each side below the 
upper keels. In A. sexangula and A. purcelli the 
bands, though present, are less noticeable and the colour 
is more evenly distributed, although the edges of the keels 
of A. sexangula are almost free from pigment. 

It will be seen that in these four species the pigment tends 
to form a pattern similar to that found in the genus Arion. 
On the other hand in Apera giV)bonsi the scheme of colora- 
tion is different, and the narrow lighter band which this slug 
usually has along the centre of the back recalls the similar 
band possessed by many species of Veronicella. It should 
be noticed that this difference in the position of the greatest 
concentration of the pigment is not correlated with the 
difference in the position of the lateral grooves ; for in 
A. dimidia the grooves are situated as in A. gibbon si, 
while the dark bands are lower down and occupy a similar 
position to those of the keeled species. 

Apera dimidia is singular in having small patches of the 
dark pigment on the foot-sole (PI. VII, fig. 9). But in some 
specimens of A. sexangula and in at least one example 
of A. gibbon si minute reddish-brown specks are scattered 
over the body, and in the former species these extend on to 
the foot. 

The lighter pigment is usually of some shade of orange, 
but varies from pale orange-yellow to dull red. It is 
generally diffused over the whole surface of the body, 
including the foot-sole, but is often darker above than 
below, the backs of Apera burnupi and A. gibbonsi 
rubella being especially deeply coloured. But this pigment 
is very unstable, and specimens which have been preserved 
in spirit seldom show more than very slight traces of it. In 
the case of an example of A. gibbonsi rubella, which was 



122 HUGH WATSOX, 

di'owned by Mi*. Burnup, tlie red pig^ment came out of the 
skin when the animal was dead, and coloured the water in 
which the slug was immersed. Notwithstanding the dai-k 
grey mottling of this foimi, the living animal is of a reddish 
colour, changing to orange below, as the unstable colouring 
matter is sufficiently abundant to mask the darker pigment 
underneath. 

The coloration of a typical specimen of A per a gibbon si 
increases its i-esemblance to the rhizome of a fern, and the 
chestnut-ln'own colour of A. burnupi may help to render 
the slug inconspicuous among dead leaves. Even the reddish 
colour of A. gibbonsi rubella may serve to conceal the 
animal, although the bright pigment of the red variety of Arion 
ater is supposed to act as a wai-ning colour. Mr. Burnup 
found a specimen of this Apera in the bush at Durban, and 
he noticed that a few of the dead leaves^ under which the 
slug was found were of a dark brownish-red colour, strongly 
veined, and curled up into rolls, and these were so very like the 
slug that it might easily have been mistaken for one of them. 

On the other hand, it seems possible that the vmstable red 
and orange pigments found in Apera and other slugs which 
are often carnivorous, as well as the similar bright colours so 
characteristic of most of the snails belonging to the Strep- 
taxi das, may be partly due to their animal food. It is known 
that this is the case with the red pigment found in the 
flamingos, the scai'let ibis, and the roseate spoonbill ; for the 
colour tends to disappear when the birds are deprived of their 
natural food, which consists chiefly of mollusks and other 
invertebrates. M. F. Woodward- has described the rich 
orange colour of the edge of the mantle in specimens of 
Natalina caff ra which were fed on snails and beefsteak, 
and Miss Davies^ mentions the brilliant orange-red colour of 

' Dr. J. Medley Wood, Director of the Natal Herbarium, has kindly 
examined one of these leaves, and thinks that it prol)a]3ly lielongs to 
Combretum krausii. 

- 'Proc. Mai. Soc.,' 1895, vol. i. p. 271. 

3 'Proc. Roy. Soc. Yietoria.' ^9V.^. vol. xxi. p. 222. 



THE r-ARNIVOROUS SLUGS OF SOUTH AFRICA. 128 

tlie mantle-border and foot in Paryphanta atramentaria 
{Sliuttl.), another carnivorous snail. 

Again, the climate has a marked influence on the colour of 
slugs, moisture being favourable to the development of dark 
colours, while heat increases the production of red pig- 
ments. Now in Apera gibbon si rubella both the dark 
and the red pigments are specially well developed, and this 
form is at present only known from the coastal region of 
Natal, Avhich is both moist and warm ; whereas the typical 
form of A. gibbonsi, as well as the other species inhabiting- 
the Province, extend further inland. 

Dr. Simi-oth^ has maintained that in Testacella the skin 
probably plays a more important part in respiration than the 
lung, and it is suggested that in such slugs as these the chief 
function of the pigment is to assist in the taking up of oxygen. 
But it does not seem likely that this is the case in Apera. 
For in this genus the respiratory tissue of the lung is Avell 
developed ; while, on the other hand, the structure of the 
integument and the arrangement of the blood-vessels which 
it contains do not suggest that the skin plays an important 
part in respiration. 

On the whole it seems probable that the pigments of these 
slugs, like those of many other animals, may be looked upon 
as by-products of metabolism, which are deposited in the skin 
usually in such a way as to help to render the animal incon- 
spicuous or to protect it from harmful rays of light, the 
development of the pigments depending partly on the food, 
but being also influenced by the climate and surroundings. 
Yet it must not be forgotten that the coloration of any 
slug is due not simply to the climate, the food, and the 
colour of the surroundings, but to the power of reacting to 
these influences which the animal possesses through inheri- 
tance. 

' • Nova Acta Acad. Ctesar Leop. -Carol. Germ. Nat. Cur.,' 1891, vol. 
Ivi, J). 245. 

VOL. 3, PART 2. 10 



124 HUGH WATSON. 



THE STRUCTUEE OF THE SKIN. 

Epidermis. — The outermost layer of the skin on the iDack 
and sides of the animal consists of a well-marked cubical 
epithelium, which in Apera dimidia attains a thickness of 
rather less than '01 mm. (PI. XII, fig. 37). The cells composing- 
this epithelium are approximately square in section ; they 
stain rather deeply, and each contains a large rounded 
nucleus. The outer walls of these cells are thickened to form 
a cuticle, while the remaining walls are much thinner. In 
this respect, thei'efore, the epidermis of Apera seems to 
resemble that of Testacella rather than that of D au de- 
bar dia, in which all the walls of the epidermal cells are 
thickened,^ or of Paryphanta, in which a cuticle is said to 
be absent. - 

The epidermis does not change its character on the dorsal 
side of the foot-fringe, as in such forms as Daudebardia* 
and Cystopelta,^ but on the sole of the foot it becomes 
quite different. Here the epithelium is columnar rather than 
cubical, and the inner ends of the cells taper and interdigitate 
with the underlying connective tissue, so that the limits of 
the epithelium are veiy ill-defined (PI. XII, fig. 38^). 
Excepting towards the edges of the foot-sole, the outer end of 
each cell is produced into a number of delicate cilia, which 
in A. dimidia measure about •0035 mm. in length. 

Inner Layers of the Skin. — In the outer part of the con- 
nective tissue, which everywhere underlies the epidermis, are 
embedded the dermal mucous glands and the pigment- 
granules. The mucous glands (PI. XII, fig. 37) are oval or 
pear-shaped, and reach a considerable size, although each is 
formed from a single cell. The protoplasm of the cell is 

' Plate, L. H., ' Zool. Jalirb.,' 1891. vol. iv, p. 529. 
••' Beutler, B., ' Zool. Jahrl).,' 1901, vol. xiv, p. 374. 
' Plate, op. cit., p. 527. 

^ Davies, Miss O. B., 'Proc. Roy. Soc. Victoria,' 1912, vol. xxiv, p. 332. 
•'' I am indebted to Mr. W. Tams for kindly taking this photomicro- 
graph. 



THE CARNIVOROUS SLUGS OP SOUTH AFRICA. 125 

chiefly confined to a thin layer lining the base of the gland 
and containing the rather large nucleus ; the rest of the 
interior is filled with mucous secretion, which can be extruded 
through a narrow pore between the cells of the epidermis. 
Dr. Plate has sepai'ated the unicellular mucous glands of 
Testacella^ and Daudebardia" into two classes, according 
to the nature of this secretion. A similar classification seems 
applicable to the dermal glands of Apera, for in sections some 
of the glands are seen to be entirely filled with a vacuolated 
colourless secretion, while in others the mucus has become 
concentrated into a compact, deeply stained mass in the centre 
of the gland. No dermal glands of either kind occur in the 
foot-sole. In this i-espect Apera resembles Paryphanta,^ 
and contrasts strongly w4th Testacella* and other forms. 
Possibly the great development of the pedal gland may have 
rendered the unicellular glands unnecessary, although in 
Paryphanta the pedal gland is not much more highly 
developed than it is in Testacella. 

The minute granules of dark pigment occur in the con- 
nective tissue between the mucous glands, l)eing chiefly 
aggregated just beneath the epidermis. Some of the granules 
are contained in pigment-cells, others appear to be irregularly 
scattered about, but possibly these may occur inside delicate 
branches of the pigment-cells. 

Beneath this glandular zone the connective tissue contains 
blood-vessels, nerves, and numerous muscle-fibres, longi- 
tudinal, radial, oblique, and circular, the last-mentioned being 
especially abundant near the inner sui'face of the skin.^ In 
Apera dimidia there are more muscle-fibres in the foot 
than in the connective tissue of the back and sides. This 

' Op. cit., p. 530. 

2 Op. cit., pp. 527, 528. 

' Beutler, op. cit., p. 375. 

* Lacaze-Duthiers, H. de, ' Arch. Zool. Exper." (2nd ser.), 1887, vol. v, 
p. 526. 

* I use the term " skiu " in a wide sense to inchide the whole body- 
wall, for no sharp line can be drawn separating the subcutaneous tissue 
with its muscle-fibres from the dermis or cutis vera. 



IBRARYj^i 




126 HUGH WATSON. 

zone of connective tissue is extremely thick, especially 
towards the hind end of the animal; indeed, the most remark- 
able thing about the skin of A per a is its great thickness and 
strength. Only in the region of the head, where flexibility is 
essential, is the skin moderately thin. Its thickness further 
back in A. gibbonsi is well shown in PI. X, fig. 32. On 
the other hand, A. dimidia has a somewhat thinner skin 
than most of the other species, though even in this form its 
thickness is not inconsiderable, as will be seen from PI. XI, 
figs. 34 and 35. This unusual thickness of the skin can easily 
be explained. In Apera the shell is so degenerate that it can 
no longer fulfil its proper functions. Yet the slug still needs 
protection from other animals seeking to prey upon it, and 
from the pressure of the soil if it burrows in the ground like 
Testacella, and also from excessive evaporation. And it still 
needs a firm attachment for its powerful retractor muscles. 
Therefore the skin has taken over the functions of the shell, 
and has become adapted to fulfil these functions adequately. 
The Rev. Dr. Cooke ^ has recently called attention to a case 
among the Poly pi acoph ora in which the protection once 
afforded by the shell has been transferred to the integument. 
But in that case the mantle has taken the place of the shell, 
while in the present instance it is the skin itself which has 
been strengthened. 

The blood-vessels contained in the skin are irregularly 
disposed, though most of them run in a longitudinal direction. 
They communica,te with the body-cavity or hgemocoele by 
minute pores, which can be seen puncturing the inner surface 
of the skin (PI. X, fig. 32). In all parts of the skin these 
vessels are provided at rather frequent intervals with power- 
ful sphincters composed of circular muscles (PI. XIII, fig. 40). 
Similar sphincters have been described by Dr. Simroth and 
P. and F. Sarasin" in Veronicella (a genus in no way allied 
to Apera) ; and it has been suggested that they serve to 
prevent the reflux of the blood when the animal moves. 

1 ' Proc. Mai. Soc..' 1913. vol. x, pp. 321, 322. 

■^ ' Die Land-Molluskeu von Celebes.' 1899. p. 90. pi. xiii. fi^js. 122, 123. 



THE CAKNIVOKOUS SLUGS OF SOUTH AFRICA. 127 

Perhaps iu Apera they also assist the circulation ot the blood 
in a similar manner. But it seems possible tliat their chief 
function may be to enable the animal to increase the turgidity 
of some or all of the dermal vessels when it contracts, and thus 
to give greater rigidity to the skin either as a whole or in part. 
For if the slug by closing the sphincters could thus increase 
the pressure in the dermal blood-vessels, the skin would 
become much more rigid in consequence, just as the strength 
of the stem of an herbaceous plant is largely due to the 
turgidity of its cells. 



THE SHELL, THE MANTLE-CAVITY, AND THE 
MANTLE. 

The Shell. — The small shell of Apera lies deep in a little 
pocket in the skin just behind and to the left of the respiratory 
opening. As a rule its length is only about one-fourteenth 
of that of the animal (iu alcohol). When viewed from above 
the somewhat oval contour of the shell is seen to be inter- 
rupted by a broad sinus which extends backwards from the 
front end more than half way along the right side. The 
posterior part of the shell is thus broader than the anterior 
half, which projects forward on the left side of the respiratory 
opening. The sinus is deepest in Apera gibbonsi, and 
shallowest in A. purcelli and A. dimidia. It will be 
seen from PI. XIII, figs. 42, 45, 48, and 51, that the shell of 
Apera is often much less flattened than is usually the case 
with the iuternal shells of slugs. In A. burnupi and A. 
sexangula it is very convex, and culminates in a conical 
apex, which overhangs the posterior margin of the shell. In 
A. purcelli the apex projects still further, but it is rounded 
instead of conical, as is also the case in A. dimidia, in 
which the apex is less prominent. Lastly, in A. gibbonsi 
and A. parva the shell is more depressed, and the apex is in 
front of the posterior margin. 

The shell is extremely thin and translucent iu Apera 
purcelli and A. dimidia. In the other species it is thicker 



128 HUGH WATSON. 

and more calcareous, especially in full-grown specimens, though 
it is usually very brittle. Indeed in some examples of A. 
gibbonsi its thickness is sufficient to suggest that the chief 
function of the shell in this species and its allies may be to 
enable the animal to get rid of any excess of calcium 
cai'bonate in its blood. It is difficult to see what other 
function it could fulfil ; and if it were to be regarded as a 
purely vestigial structure with no function at all, one would 
have expected it to have become much smaller as well as 
flatter in this species than in the more primitive forms such 
as A. purcelli. Yet, while the shape has degenerated, the 
relative size has remained the same nnd the thickness has 
greatly increased. 

The Shell-sac. — The pocket in which the shell lies has no 
communication with the exterior, but is close to the inner 
surface of the skin (PI. X, fig. 33). In those species in which 
the shell has a prominent apex, the floor of the sac is raised 
towards the hind end to form a papilla fitting into the hollow 
of the shell. Where the margin of the shell rests upon 
the floor of the pocket, there is a fold, Avhich separates the 
central part of the floor underlying the interior of the shell 
from a nari'ow rim beyond it. 

The shell-sac is lined by a well-defined epithelium. On the 
roof of the cavity this epithelium is very thin, being formed in 
Apera dimidia of a compact layer of small cells which are 
slightly broader than they are high. Underneath the shell 
the epithelium is thicker in the same species, and composed 
of larger cells, whose height exceeds their breadth. These 
larger cells are continued over the edge of the fold mentioned 
above, the transition from the thicker to the thinner epithe- 
lium taking place on the outer side of the fold. 

The Mantle-cavity or Lung. — The mantle-cavity occupies 
the region below the shell and the respiratory opening, and also 
extends a short distance further forward. It is, nevertheless, 
mainly behind instead of above the body-cavity or ha3mocoele, 
from which it is separated by a muscular diaphragm sloping 
obliquely downwards and backwards. There is, however, a 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 129 

funnel-shaped prolongation of the body-cavity which extends 
backwai'ds for a variable distance beneath the mantle-cavity, 
and raises its floor along the middle. This feature is well shown 
in PL XI, figs. 34, 35, and PI. XII, fig. 36, for the prolongation 
is unusually large in Apera dimidia and A.purcelli. 

In horizontal section the mantle-cavity or lung is roughly 
triangular, with the large pericardium occvipying the centre 
of the broad anterior part, Avhich is thus divided into a right 
and a left wing (PI. IX, figs. 27-31). The right division opens 
widely behind into the posterior part of the mantle-cavity, but 
the left division is cut off by the kidney, which extends 
obliquely backAvards from the pericardium and fuses with 
the left wall and the roof of the cavity. Thus the left 
anterior division only communicates with the rest of the 
nuintle-cavity by a narrow space round the top and front 
of the pericardium and kidney. It is, therefore, not sur- 
prising that the walls of this part of the cavity are without 
respiratory tissue. But the right anterior division, which 
communicates more freely with the air, and the greater part 
of the posterior half of the cavity have their walls richly 
supplied with pulmonary veins. These vessels branch 
repeatedly, and in the larger species the fine branches form a 
complicated network which divides the surface into numerous 
alveoli. The thinness of the epithelium which separates the 
blood in these vessels from the air in the mantle-cavity will be 
seen from PI. XII, fig. 39. The only portion of the posterior 
half of the cavity that is devoid of respiratory tissue is a 
part of the roof ^ and the immediate neighbourhood of the 
respiratory opening and the anus just below it. 

At the hind end the cavity tapers to a point. In Apera 
gibbon si this point is below the extreme hind end of the 
shell-sac; but in species such as A. sexangula and A. 
dimidia it is within the papilla which is formed by the floor 
of the shell-sac projecting into the hollow apex of the shell. 
Hence in these species the hind end of the mantle-cavity is 
inside the shell itself. 

' See p. 187. 



130 HUGH WATSON. 

I have only been able to examine the minute structure of 
the epithelium lining the mantle-cavity in one species, namely 
A. dimidia. In this species (and probably also in the 
others), that part of the posterior half of the cavity where 
there is no respiratory tissue is lined by moderately large 
cells, slightly higher than they are broad, and strongly ciliate 
near the respiratory opening. These cells are continuous 
through the aperture with those of the epidermis. The whole 
of the remainder of the cavity, including both of the anterior 
divisions, is lined by a pavement-epithelium of cells so thin 
that the nucleus of each usually forms a knob-like projection 
in the centre of its surface. 

The Mantle. — Apera has no external mantle — a remark- 
able fact that was first pointed out by Heynemann.^ Yet it 
cannot be entirely Avithout a mantle, for, as we have just seen, 
it possesses both a mantle-cavity and a shell ; and all Gastro- 
pods that have a shell also possess a mantle, as the 
shell is always secreted by the mantle. I therefore regard 
that part of the integument Avhich forms the roof of 
the mantle-cavity beneath and around the shell as being 
undoubtedly homologous with the mantle. Moreover I 
believe that the mantle is not entirely concealed from view in 
all the species of Apera, for it seems probable that it forms 
the inner lip of the respiratory opening, and that the upper 
surface of this lip may be regarded as part of the upper 
surface of the mantle (PI. X, fig. 33). This theory is confirmed 
by the occurrence of an oblique cleft in the inner lip of the 
opening in a similar position to the cleft which runs from the 
opening to the edge of the mantle in an ordinary slug. 

We may suppose that Apera has been evolved from a 
snail, having on its back a mantle covered by a shell. It 
might well be an advantage to such an animal to get rid of 
this projection, especially if it burrowed in the ground in search 
of worms. Accordingly both mantle and shell seem to have 
sunk into the back and to have become covered by folds of the 
surrounding skin which have grown over it on each side. 
' ' Jahrh. d. Deutsch. Mai. Ges.,' 1885, vol. xii, p. 19. 



THE CAHNIVOKOUS SI-lMiS OF SOU'l'H AFIMCA. lol 

The upward growth of the skin on the right side would tend 
to cover the respiratory opening, which would be in its usual 
position in the right margin of the mantle. To obviate this, 
the mantle edge, with the opening, would have to move up 
towards the middle line, and the growth of the riglit side of 
the shell would be retarded in consequence. And at the same 
time the fold of the skin would be likely to grow more 
slowly immediately opposite the opening than before and 
behind it, thus forming a notch in the edge of the fold. 
Eventually this fold of the skin would unite with the other 
in the middle line excepting at the notch, which would be 
converted into an opening above the original respiratory 
opening in the mantle-edge. 

This hypothesis may not be correct, but it has at least the 
merit of explaining not only the absence of an external 
mantle in Apera, but also the dorsal and slightly asym- 
metrical position of the respiratory opening, the fact that the 
opening has an inner and an outer lip, the former with an 
oblique cleft, and the peculiar sinus in the right side of the 
shell. So far as I am aware the morphology of the mantle 
and shell of Apera differs from that of all other Pulmonates. 

THE PEDAL GLAND. 

The pedal or supra-pedal gland attains a greater develop- 
ment in Apera than in any other genus of snails or slugs 
known to science. It consists essentially of a long duct with 
glandular walls, which opens in front between the mouth and 
the anterior end of the foot, and ends behind in a small vesicle. 

The Position and Length of the Gland. — In the herbivorous 
genera of snails and slugs the pedal gland is usually more or 
less embedded in the muscular tissue of the foot, and this is 
also the case in D audebardia,^ Plutonia,- and the Tri- 
gonochlamin^e^; but in Apera it lies freely in the lower 

' Plate, L. H., ' Zool. Jahrb.,' 1891. vol. iv, p. 525. 
- Simroth, H., ' Nova Acta Acad. Caesar Leop. -Carol. Germ. Nat. Ciir.." 
1891, vol. Ivi, p. 229. 

'■' Simroth, H., • Festschrift Leuckarts,' 1892, pp. 53, 55 ; etc. 



182 HUGH WATSON. 

part of the body-cavity, as in Natalina, Rhytida, Pary- 
phanta, Schizoglossa, Testacella, Streptaxis, 
At op us, and a few other carnivorous genera, and is only 
held in position by the pedal artery which supplies it with 
blood. This artery confines the gland to the anterior part of 
the body-cavity, for it runs back above the gland, giving off 
branches to it as it goes, and, after becoming attached to the 
posterior vesicle for a short space, it dips down and unites 
with the upper surface of the foot about the middle of its 
length. The pedal gland, however, is many times longer 
than this free portion of the pedal artery and is consequently 
thrown into numerous folds and loops. In Apera sexangula 
the gland pursues a comparatively regular zig-zag course, 
bending alternately to the right and the left about half a 
dozen times (PI. XIII, fig. 57). But in A. burnupi and 
A. purcelli, and in the posterior part of the gland in the 
remaining species, the loops are deeper and more complicated, 
twisting about in all directions (figs. 52-56). The gland 
attains its greatest length in A. burnupi, in which its folds 
not only extend up the sides of the body-cavity, but even 
curve over on to the top of some of the other organs (PI. IX, 
fig. 30). 

The Glandular Tissue. — The pedal gland of Apera is not 
only unusually long, it is also often exceptionally broad, as, 
for example, in A. burnupi and in the anterior half of 
its length in A. dimidia (PI. XIII, figs. 56, 54). In 
A. gibbonsi and A. parva it is more slender (figs. 52, 53), 
possibly because the great size of the odontophore in these 
species leaves little room for the growth of any other bulky 
organ in the anterior half of the body-cavity. 

The breadth of the gland is due to an enormous develop- 
ment of glandular tissue on the floor and sides of its duct. 
In Apera purcelli, A. burnupi, and A. sexangula this 
glandular tissue extends along the whole length of the duct 
as far as the terminal vesicle ; but it is less abundant towards 
the posterior end and the gland becomes rather narrower 
behind in consequence. In A. gibbonsi rubella the 



THE CARNIVOEOUS SLUGS OF SOUTH AFRICA. 133 

glandular tissue becomes still scai^cer towards the hind end, 
and disappears entirely just before the vesicle is reached, 
while in the other races of A. gibbonsi and in A. parva 
the last few millimetres of the duct are quite destitute of 
gland-cells. Finally, in A. dimidia the glandular tissue only 
extends along half the length of the duct, the posterior half 
being entirely without it, and therefore much narrower than 
the anterior half (PI. XIII, figs. 54, 59, and 60). 

PI. XTV, fig. 63 shows the structure of the glandular tissue 
near the front end of the pedal gland in A. gibbonsi 
rubella. It will be seen that among the large oval gland- 
cells, which taper towards the duct, there are scattered con- 
nective-tissue cells with conspicuous rounded nuclei, and 
there can be little doubt that it is from these cells that the 
glands are formed. Similar cells occur near the front end of 
the pedal gland in A. dimidia, but further l)ack, near the 
hind end of the anterior half of the gland, no such cells 
occur, and a transverse section has the appearance shown in 
PI. XIII, fig. 59. Possibly the absence of connective-tissue 
cells from this region may be correlated with the absence of 
gland-cells still further back. The posterior part of the 
gland will probably be the first to be formed in development, 
and if all its cells, apart from those which form the duct 
itself, are converted into glands, there will be nothing left to 
replace these when they die and are discharged into the duct ; 
for according to Andre ^ the gland-cells perish after a certain 
time and need to be replaced. 

Mr. Collinge- has stated that the pedal gland of Apera 
sexangula contains "a large series of microscopic chitinous 
(?) dart-like bodies," but his figures leave little doubt that 
these are merely the solidified contents of the glands. In 
specimens preserved in alcohol this congealed secretion 
appears through a simple lens as numerous white specks, 
which often give a glistening appearance to the pedal gland. 

' 'Revue Suisse de Zoologie,' 1894, vol. ii, pj). 310, 311. 
2 ' Ann. Mag. Nat. Hist.' (6th ser.). 1897, vol. xx. p. 223. pi. v, figs. 
4, 5. 



184 HUGH WATSON. 

The Duct. — The duct of the pedal gland is semi-trans- 
lucent, and can be seen running- along the centre of its upper 
surface. Near the opening of the gland the duct usually 
lies more deeply (PI. XIV, fig. 63), but elsewhere the 
glandular tissue is restricted to its sides and lower surface, as 
shown in PI. XIII, fig. 59. Where the glandular tissue is 
less abundant the exposed surface of the duct often shows a 
tendency to have alternate swellings and constrictions. In 
Apera gibbonsi this culminates in a close convolution of the 
posterior part of the duct (PI. XIII, fig. 58), which reminds 
one of the somewhat similar convolution of part of the duct 
in Testacella.^ On the upper sui-face of the anterior half of 
the gland in A. dimidia, two brownish lines can often be 
seen running along, one on each side of the duct, at its junc- 
tion with the glandular tissue. 

In section the duct is approximately circular, without any 
folds on its floor (PL XIII, figs. 59, 60, PI. XIV, fig. 63). In 
this respect it resembles that of Testacella, and differs from 
that of Daudebardia, for in the latter genus, as in most 
herbivorous snails and slugs, the floor of the duct has two 
longitudinal folds enclosing a median groove, and the gland- 
cells only open into this groove, instead of all round the duct.- 
In Paryphanta hochstetteri [Pfr.) traces of the folds 
and median groove seem to occur, but only towards the 
anterior end of the duct."' The duct is lined by an epithelium 
composed of small cubical cells, and this is surrounded by two 
thin muscular layers, the fibres of the inner layer running 
round the duct, while those of the outer layer are longitu- 
dinal. Both layers of muscles occur inside the glandular 
tissue. 

The Terminal Vesicle. — After becoming gradually narrower 
the gland widens again at the hind end to form an oval or pod- 
shaped vesicle (PI. XIII, figs. 52-57). In length this vesicle 

' See de Lacaze-Duthiers, ' Arch. Zool. Exper.' (2nd ser.), 1887, vol. v, 
pi. xl, fig. 95. 

' Plate, L. H., ' Zool. Jabrb.,' 1891, vol. iv, p. 525, pi. xxxii, fig. 16. 
■» Beutler, B., • Zool. Jalirb.." 1901. vol. xiv, p. 376. pi. xxvi, fig. 9. 



THE CARN1V(3R0US SLUGS OF SOUTH AFRICA. 135 

varies from about 1 mm. in the smaller species to more than 
5 mm. in a large specimen of A p e r a b u r n u p i, and its breadth 
is usually about one-third of its length. The walls of the 
vesicle are thin and semi-translucent like those of the duct, 
with which they are continuous. A few gland-cells occur along 
the ventral^ surface in A. burnupi, A. sexangula, and 
A. purcelli, but none at all in the other species. The 
structure of the walls of the vesicle is similar to that of the 
walls of the duct, excepting that the cells of the epithelium 
become somewhat higher on the dorsal side, and among the 
ordinary epithelial cells there are scattered a few rather 
small vacuolated cells. 

From the dorsal wall of the vesicle there projects into its 
cavity a remarkable fold, resembling a typhlosole, which fills 
up a large portion of the interior, so that the cavity is cres- 
centic or horse-shoe-shaped in section (PI. XIII, fig. 62). If 
the wall of the vesicle in Apera sexangula be cut open, it 
will be seen that the fold has a spongy and somewhat lami- 
nated appearance. Serial sections through the vesicle of 
A. dimidia show that in this species the fold is rounded 
excepting at the ends, where it becomes wrinkled and divided 
by furrows (cf . PL XIII, figs. 61, 62, PI. XIV, fig. 66). Fig. 
QQ shows the structure of the fold in A. dimidia. It will be 
seen that it is entirely composed of vacuolated tissue, which 
is not bounded by a distinct epithelium. The pedal artery, 
which always becomes intimately connected with the dorsal 
surface of the vesicle, gives off a branch into the middle of 
the fold (PI. XIII, fig. 62). In A. gibbonsi the structure 
of the fold is different, for in this species it is divided into a 
complicated series of narrow lamellas. Fig. 58 shows the 
appearance of these as dimly seen through the wall of the 
vesicle, Avhile PI. XIV, fig. 65, shows them in section. Each 

' I use the terms '" dorsal " and " ventral " in a morphological sense, 
the dorsal side being that to which the artery is attached. Generally, 
however, the vesicle lies on its side, owing to the twisting of the duct, 
and the attachment of the artery to the floor of the body-cavity beyond 
the vesicle. 



136 HUGH WATSON. 

lamella seems to be composed of a doable layer of peculiar 
rounded or polygonal cells with thick walls. 

The hind end of the pedal gland is also enlarged to form a 
vesicle in Rhytida/ Paryphanta,^ and Schizoglossa/^ 
and indeed this is probably the case in all the genera of the 
RhytididaB, for I have found such a vesicle in Natal in a as 
well. Moreover, Rentier^ has shown that in Paryphanta 
it contains a lai'ge fold similar to that in Apera, but appa- 
rently less highly specialised in structure. In Testacella 
there is also a dorsal fold ■' projecting into the duct at the 
hind end of the pedal gland, but the structure of the fold 
appears to be very different from that of Apera, and there 
is no terminal enlargement of the gland to form a vesicle. 

Various possibilities suggest themselves with regard to the 
function of the terminal vesicle of the pedal gland, and the 
fold which it contains ; but so little is known about the physio- 
logy of the gland in the carnivorous genera that such specu- 
lations can have little value. It is better, therefore, simply 
to state that the function of these structures is at present 
unknown. 

THE NERVOUS SYSTEM. 

The centi-al nervous system of Apera consists of three 
compact ganglionic masses joined by connectives. The first 
of these is formed of the cerebral ganglia, the second of the 
buccal ganglia, and the third of the pedal, pleural, and 
visceral ganglia. 

The Cerebral Ganglia and Commissures. — As in nearly all 
carnivorous snails and slugs, excepting Daudebardia'' and 

' Colliiige, W. E., ' Ann. Mag. Nat. Hist.' (7th ser.), 1901, vol. vii, 
pp. 67, 68, pi. i, figs. 12, 13. 

2 CoUinge, op. cit., p. 70, pi. ii, figs. 21, 24; Min-docli, ' Trans. N.Z. 
Inst.,' 1904, vol. xxxvi, p. 158, pi. vi, fig. 7. 

3 Muvdoch, R., ' Proc. Mai. Soc,,' 1901, vol. iv, p. 170. pi. xvii, fig. 10. 
^ • Zool. Jalirb.,' 1901, vol. xiv, p. 376, pi. xxvi, fig. 7. 

•^ Plate, L. H., ' Zool. Jahi-b.,' 1891, vol. iv, p. 525, pi. xxxii, figs. 17, 18. 
8 Plate, op. cit., pp. 589, 593. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 137 

the Trigone chl am inte/ the cerebral ganglia are situated 
very near together on the dorsal side of the buccal mass. 
Indeed, in Apera gibbon si they are so closely bound to 
each other by connective tissue that they appear as one 
(PI. XIII, fig. 52) ; and it is only in the smaller species, such 
as A. parva (fig. 53) and A. purcelli (fig. 55), that they 
can be clearly seen to be separate ganglia without the help 
of any reagents. When fi*eed from their surrounding tissue, 
the ganglia are seen to be oval structures, nearly twice as long 
as they are broad, and rather nearer together behind than in 
front (PI. XV, figs. 70-72). Their average length in Apera 
is about f mm. Towards the outer side of the front end of 
each ganglion there is a slight eminence formed by the 
accessory lobe. This is best seen in A. dimidia. In this 
species the apex of each lobe is prolonged into a filament, 
which is very like a slender nerve (fig. 71), but is probably 
the remains of the embryonic cerebral tube which originally 
connected the accessory lobe with the exterior. Similar 
accessoiy lobes have been figured by de Lacaze-Duthiers ~ in 
Testacella, l)y Wiegmann'^ in Ennea (Edentulina), and 
by Beutler ^ in Paryphanta, and the last author has 
described them at some length. 

The cerebral ganglia are joined by two commissures, 
namely, the cerebral commissure above the buccal mass, and 
the sub- cerebral commissure below it. The cerebral com- 
missure is very short and broad, and is situated somewhat 
towards the posterior ends of the ganglia (PI. XV, fig. 71). 
The sub-cerebral commissure, on the other hand, is very long 
and extremely slender. It arises from about the middle of 
the outer sides of the ganglia, and encircles the buccal mass 
just in front of the cerebro-pedal connectives and the pedal 
ganglia, being closely bound to these by connective tissue 

' Simrotli, H., ' Festschrift Leuckarts,' 1892, pp. 53, 55. 
' 'Arch. Zool. Exper.' (2nd ser.), 1887, vol. v, pi. xxxviii, figs. 70, 71. 
3 ' Mitt. Zool. Samml. Mns. Nat. Berlin,' 1898, vol. i, pi. iii, fig. 7. 
* ' Zool. Jahvb..' 1901, vol. xiv, pi. xxix, figs. 58, 61. and pp. 400-402 ; 
see also Pelseneer, • Mem. Acad. Roy. Belg.,' 1901, ex. vol. liv, pp. 31-37. 



188 HUGH WATSON. 

(figs. 73 and 74). It thus forms the most anterior element of 
the nerve-collar. Amaudrut ^ has described a similar sub- 
cerebral commissure in Achatina panthera (Fer.), Buli- 
mus funki {Nyst.), Nanina cambodjiensis [Reeve), and 
Helix aspersa Mull.; but hitherto this slender commissure 
has been noticed in only a very few Pulmonates and in none of 
the carnivorous forms. I have found it, however, in Natalina 
quekettiana [M. Sr P.), the only member of the Rhy tididae 
whose ganglia I have examined microscopically ; and I 
believe that if malacologists would examine the central 
nervous system of snails with greater care they would find it 
in many other pulmonate genera. For among the marine 
Euthyneura a sub-cerebral commissure has been observed 
not only in the Pleurobranchid^ and in numerous Nudi- 
branchs,- but also in so primitive a form as Actfeon.'^ 
Moreover, I am inclined to regard the sub-cerebral com- 
missure as homologous with the important labial commissure 
found in the Aspidobranchia, and in Vivipara and 
Ampullaria among the Pectinibranchia, as well as in 
the Amphineura, S cap hoped a, and Cephalopoda."* In 
most of these forms the cerebro-buccal connectives arise 
from the labial commissure instead of from the cerebral 
ganglia themselves, and in Apera the cerebro-buccal con- 
nectives arise from the ganglia very close to the ends of the 
sub-cerebral commissure. 

A little further back arise the thick cerebro-pedal con- 
nectives ; and behind this again, nearly at the posterior end 
of the ganglia, the almost equally broad cerebro-pleural con- 
nectives arise. In the specimen of Apera dimidia that I 

' 'Bull. Soc. Philom. Paris' (7th ser.}. 1885-8(3. vol. x. pp. 107-117: 
'Aim. Nat. Sci., Zool.,' 1898, vol. vii, p. 127. 

- Pelseneer, P., ' Mem. Couronne Acad. Roy. Belg.." 1893. ex. vol. liii. 
p. 69. 

3 Bouvier. E. L.. ' Bull. Soc. Philom. Paris " (8th ser.), 1893, vol. v, p. 67. 

'' Pelseneer has shown that the so-called " labial commissiu-e " which 
Pleurobranchsea and a few other forms possess in addition to the 
sub-cerebral commissure, is merely an anastomosis of two of the nerves 
to the lijDS (op. cit.. p. 33). 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 139 

examined, these two connectives were joined on the right 
side by a short transverse filament not far from their origin 
(PI. XV, %. 71). 

The Nerves of the Cerebkal (xANglia. — From each 
cerebral ganglion arise two large nerves which are as thick 
as the cerebro-pedal and cerebro-pleural connectives. These 
are the olfactory nei've to the upper tentacle, and the nerve 
to the lower tentacle. The olfactory nerve springs from the 
upper surface of the anterior end of the ganglion, towards 
the inner side (PI. XV, figs. 70-72). Near its origin there 
also arise three slender nerves, namely, the two peritentacular 
nerves, and the optic nerve, which is distinct from the 
olfactory nerve even from the ganglion. The nerve to the 
lower tentacle arises nearer the outer edge of the ganglion 
than the four nerves just described, and external to the 
accessory lobe. Near it arise the two labial nerves, which 
are very much narrower than the nerve to the lower tentacle, 
but not quite so slender as the optic and peritentacular 
nerves. As in most genera the nerve to the penis also springs 
from this region in the right cerebral ganglion ; it is of about 
the same thickness as the labial nerves. Beutler ^ states that 
in Paryphanta hochstetteri (P/V.) the penial nerve 
does not arise separately from the cerebral ganglion, but 
seems to branch from one of the other nerves. On the other 
hand, in the Streptaxida3^ the nerve to the penis arises 
directly from the pedal ganglion, instead of its fibres first 
traversing the right cerebral ganglion, as is usually the case. 
Near the origin of the sub-cerebral commissure a very 
slender nerve arises on each side, and runs down with it in 
the connective tissue on the anterior side of the cerebro-pedal 
connective. Above this connective, on the dorsal surface of 
each ganglion, there arises another very slender nerve, which at 
first goes obliquely backwards, but quickly curves outwards, 
and runs down to the ventral group of ganglia between 
the two connectives. This is almost certainly the nervus 

1 Op.cit., p. 400. 

- Wiegmann, F., op. cit., p. 62. 
VOL. 3, PART 2. 11 



140 HUGH WATSON. 

otocysticns, altliough I was unable to find the otocysts 
themselves even in sections of the pedal ganglia. Lastly, a 
slender nerve arises from each ganglion between the two 
connectives, usually neai-er to the cei-ebro-pleural connective 
than to the other. In Apera dimidia, A. pur colli, A. 
burnupi, and A. sexangula these nerves ruii back to the 
anterior end of the buccal retractor, which they innervate. 
Each gives off two branches close to the cerebral ganglia ; 
one of these anastomoses with the cerebro-buccal connective 
not far from its origin ; the other runs down between the 
cerebro-pedal and cerebro-pleural connectives, and divides 
into two about half way to the ventral group of ganglia. 
One of the divisions of this branch leaves the nerve collar 
and innervates the retractor of the lower tentacle.^ The 
other division continues down to the ventral ganglia ; but 
whether it fuses with the pleural ganglion, as one might expect 
from Amaudrut's observations," or whether it merely crosses 
the dorsal surface of the ventral group of ganglia and leaves 
it again in the wall of the buccal artery, I was unable to 
discover with the limited amount of material at my disposal. 
The examination of these slender nerves, deeply embedded 
amongst the connective tissue and blood-vessels which 
surround the larger nerve-cords and ganglia, is a matter of 
considerable difficulty ; but it does not even requii-e a com- 
pound microscope to see that the anterior end of the buccal 
retractor in A. dimidia, A. sexangula, and their allies is 
innervated by nerves arising from the sides of the cerebral 
ganglia. This is one of the very few respects in which 
Apera resembles Daudebardia more than any other 
carnivorous genus with which I am accpiainted.^ In A . 
gibbon si the buccal mass with the odonrophore is much 

1 In Natalina quekettiana (M. & P.) and Rhytida capillacea 
(Fer.) the i-etractors of the lower tentacles are also innervated hj very 
slender nerves issuing from the nerve-collar about half-way down each 
side, and this is prol^ably the case in several other forms as well. 

» ' Ann. Nat. Sci., Zool.,' 1898, vol. vii, pp. 123-126. 

3 See Plate, L. H., ' Zool. Jahrb.,' 189], vol. iv. p. 591. 



THE CARNIVOROUS SLUGS uF SOUTH AFRICA. 141 

larger, and the buccal retractors are more highly developed, 
and it might therefore have been expected that these nerves 
would have been larger and more conspicuous than in the 
other species of Apera. The opposite is the case. In this 
form the corresponding nerves are extremely slender. They 
are joined for some distance with the cerebro-buccal connec- 
tives (PL XIII, fig. 52), and then leave these and unite with 
the sheath of the odontophore at the anterior end of that 
organ. A possible explanation of this apparent anomaly will 
be found in my account of the morphology of the buccal 
retractors.^ 

The Buccal GtAnglia. — The buccal or stomato-gastric 
ganglia are situated close together on the dorsal surface of 
the odontophore, just behind and beneath the front end of 
the oesophagus. In Apera burnupi and A. sexangula 
they are usually in front of the cerebral ganglia, the cerebro- 
buccal connectives curving forwards (PI. XIII, figs. 56 and 
57). In A. dimidia and A. purcelli they are normally 
situated a short distance behind the cerebral ganglia, when 
the odontophore is not protruded (figs. 54, 55). Lastly, in 
A. gibbonsi and A. parva the buccal ganglia are generally 
very far behind the cerebral ganglia, and the cerebro-buccal 
connectives are unusually long in consequence, often attaining 
a length of 5 or 6 mm. in the former species (PI. XIII, figs. 
52, 58, and PI. XV, fig. 70). This posterior position of the 
buccal ganglia and the consequent lengthening of the 
cerebro-buccal connectives is not uncommon among carni- 
vorous snails and slugs, and is entirely due to the increase in 
size of the buccal mass corresponding to the growth of the 
radula. As the buccal mass becomes larger and longer, the 
opening of the oesophagus, with the buccal ganglia behind it, 
is pushed back until it comes to lie posterior to the cerebral 
ganglia, and should this process be continued still further, 
the cerebro-buccal connectives must be correspondingly 
lengthened if the cerebral ganglia are to remain in their 
original position near the sense-organs. Moreover, the 
' See pp. 17.3. 174. 



142 HUGH WATSON. 

longer the cerebro-liuccal connectives, the more freely can 
the odontophore be protruded ; for it is evident that when the 
buccal ganglia normally lie behind the cerebral ganglia, the 
odontophore bearing the buccal ganglia can be brought for- 
ward a distance equal to twice the length of the connectives 
without moving the cerebral ganglia at all. 

The buccal ganglia are rounded or oval, and joined to 
each other by a short buccal commissure. As might have 
been expected, they are proportionately larger and closer 
together in those forms in which the odontophore is very 
large than in such species as A. burnupi, in which the 
odontophore is much smaller. In A. gibbonsi the ganglia, 
when freed from the connective tissue which surrounds 
them, are seen to be almost as near to each other as are the 
cerebral ganglia (PI. XV, fig. 70). 

The Nerves of the Buccal G-anglia. — Three slender 
nerves arise from the anterior part of each buccal ganglion 
(figs. 70-72). One of these becomes attached to the salivary 
duct of the same side, and passes backwards to the salivary 
glands. The other two go to the oesophagus, one bifurcating 
soon after it leaves the ganglion. Two more pairs of 
nerves arise laterally from the ganglia and innervate the 
sides of the buccal mass and odontophore, one pair going 
forwards and the other backwards. These nerves are 
especially large in Apera gibbonsi, in which the parts they 
innervate are so greatly developed. The anterior lateral 
nerve passes forwards to the side of the buccal mass in front 
of the oesophagus, and in this species it is generally united at 
its origin for a very short distance with the cerebro-buccal 
connective on the same side. In A. sexangula, on the 
other hand, this nerve appears to be united near the ganglion 
with the posterior lateral nerve, a fact which is probably 
connected with the anterior position of the buccal ganglia in 
this species. The posterior lateral nerve on each side passes 
to the sheath of the odontophore, and bifurcates as it does so. 
The outer and more slender branch, after passing through 
the external longitudinal muscles, runs down the sides of the 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 143 

odontophore in the circular muscles of the sheath, which it 
doubtless innervates. The inner and larger branch goes 
more deeply. It runs down in the suspensor muscles of the 
odontophoral support and then divides, one division running 
forwards and the other backwards in the angle of the groove 
between the lower ends of the suspensor muscles and the 
origin of the lateral retractors of the radula.^ 

The nerves which innervate the central part of the odonto- 
phore are widely separated from the other buccal nerves in 
their origin. They arise from the inner sides of the buccal 
ganglia, or even from the buccal commissure, and consist of 
two rather large nerves and usually one or two others which 
are very much smaller. These nerves pass through the sheath 
of the odontophore, and run back with the median dorsal 
muscle until it dips down between the lateral retractoi's of the 
radula. They then curve forwards again at a lower level, and 
are distributed where the retractors are inserted on the 
radula-sac. But the most remarkable feature about these 
nerves is their asymmetrical origin. The left of the two 
main nerves always arises nearer the middle line than the 
right ; indeed, it might be said that while the right nerve 
arises from the inner side of the right ganglion, the left nerve 
arises from the commissure, though towards its left end (see 
especially PI. XV, fig. 71). This asymmetry seems to be a 
constant feature of the genus Apera, for I have examined 
microscopically the buccal ganglia of five specimens, belonging 
to three different species, and they all show it. Yet it is 
very surprising, for the odontophore is one of the few organs 
which is generally supposed to have escaped the effects of 
the torsion which has played such havoc with the symmetry 
of G-astropods. And, in fact, the odontophoral muscles of 
Apera do seem to be quite symmetrically arranged, and 
the odontophore occupies an approximately symmetrical 
position, and even the extrinsic buccal retractors show no 
traces of asymmetry excepting towards the hind end of 
the animal. »Since the cause of this asymmetry cannot be 
' See pp. 166, 167. 



144 HUGH WATSON. 

found in Apera itself, we must inquire whether it can 
be discovered in the snails from which the genus may have 
sprung. 

There are certain muscles in the odontophore of Apera 
and Natalina — the flexor muscles of the odontophoral sup- 
port — which, when they are contracted, tend to curve up 
its front end. When this happens the odontophore is likely 
to turn on to its right side, for there is not room for it 
to become curved in a vertical plane above the pedal 
gland, and the reproductive organs will prevent it turning 
on to its left side and curving outwards to the right. And 
in the only specimen of Apera that I have seen in which 
these muscles were contracted, the odontophore was on its 
rig'ht side, curving outwards to the left. Now in carnivorous 
snails with a dextral heliciform or depressed shell, an 
odontophore which curves outwards to the left will lit into 
the body-whorl of the shell much better than one that is 
straight, when the animal retii^es into its shell ; and the 
oblique pull of the retractor muscles arising from the 
columella of the shell will tend to maintain this curvature. 
Thus one might expect to find that in such snails the 
anterior part of the odontophore would be normally lying on 
its right side, with the opening of the oesophagus lateral 
instead of dorsal ; and this is exactly what has been found 
to be the case in Paryphanta hochstetteri [Pfr.)^ and 
Natalina trimeni {M. & P.).~ The nerve-collar Avould not 
be rotated to any extent, as both the cerebral and ventral 
ganglia would be held in position by the nerves which radiate 
from them to the skin, tentacles, etc., on each side ; but 
owing to the odontophore curving outwards to the left, the 
cerebro-pedal and cerebro-pleural connectives would become 
much more lengthened on the left side than on the right 
(see Beutler's fig. 60). The buccal ganglia, however, would 
obviously be involved in the rotation. But owing to the fact 
that the cerebral ganglia are broader than the buccal ganglia, 

' Beutler, B.. ' Zool. Jahrb.,' 1901. vol. xiv. p. 377. pi. xxix, fig. 60. 
■■' Pace, S., ' Proc. Mai. Soc.,' 1895, vol. i, p. 233. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 



145 



the left cerebro-buccal connective would become stretched, 
and would consequently retard the rotation of the buccal 
ganglia to some extent, so that they would come to lie slightly 
to the left of the opening of the cesophagus and the median 
line of the odontophore.^ The result of this would be that 
the nerves from the buccal ganglia would slope obliquely to 

Text-fig. 1. 




-■ Cerebro-buccal connective. 



Opening of cesophagus. 

Buccal ganglia. 

Odontophoral nerves. 

Centre of odontopliore. 



Diagram illustrating the possible origin of the asymmetry of the 
odontophoral nerves of Apera. 

the right, as shown in the accompanying diagram, and this 
would tend to shift their orig-in to the rig-ht. When the shell 
became degenerate and the animal assumed a symmetrical 
form, the odontophore would tend to return to its original 
condition and to lie in a straight line between the mouth and 
the buccal retractor- ; but it might be a long time before the 
buccal nerves moved back again to their symmetrical position. 
Now, as I shall attempt to prove later, it is not improbable 
that Apera may have been evolved from a group of 

' Cf . Plate, L. H.. ' Zool. Jahrb..' 1891. vol. iv, pi. xxxvii, fig. 102. 

- The effect of the slightly asymmetrical origin of the buccal re- 
tractor would be counteracted by the pressure of the reproductive 
organs on the right side. 



146 HUGH WATSON. 

carnivorous snails with dextral heliciforin or depressed shells ; 
therefore it seems possible that the above explanation of the 
asymmetry of the nerves of the odontophore may be not very 
far from the truth. 

The Venteal Group of Ganglia. — Beneath the odontophore 
or the posterior part of the buccal mass, and a little further 
back than the cerebral ganglia, there lie six ganglia very 
close to each other. These are the two pedal, the two pleural, 
and the two visceral ganglia. In A per a purcelli the limits 
of each of these ganglia can be clearly seen, although they are 
very near together ; but in some of the larger forms, such as 
A. gibbonsi, the sepai-ate ganglia can only be distinguished 
with difficulty, so closely are they aggregated (PI. XIV, fig. 
67). These ganglia are joined to the cerebral ganglia by the 
cerebro-pedal and cerebro-pleural connectives, the length of 
which varies directly with the size of the buccal mass and 
odontophore. Thus in A. gibbonsi these connectives are 
very long, so as to permit of the protrusion of the enormous 
odontophore (PI. XIII, fig. 52), while in A. burnupi and A. 
sexangula they are fairly short. The connectives are 
sometimes slightly, but rather abruptly, swollen at their 
junctions with the pedal and pleural ganglia (PI. XIV, fig. 
68). 

The most anterior of these ganglia are the pedal ganglia, 
which are also more ventrally situated than the others. 
These are the largest ganglia in the nervous system, being 
even larger in Apera than the cerebral ganglia. They are 
somewhat oval in shape, and very close together. Two short 
commissures connect them, one anterior and dorsal, the other 
further back and more ventrally situated (PI. XV, fig. 74). 
The first of these is usually termed the pedal commissure, 
and is shown in section in PI. XIV, fig. 68 ; the second is 
knoAvn as the parapedal commissure, and is seen to be rather 
thinner than the other in vertical section. Beutler^ has 
found both commissures in Paryphanta hochstetteri 
{Pfr.), and it is probable that they occur in all the Euthy- 

» Op. cit.. p. 402. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 147 

neura, both Pulmonates and Opisthobranchs.^ The anterior 
commissure appears to be of mixed origin, for an examination 
of serial sections through the region of this commissure in A . 
dimidia seems to show that at least in its upper half it is 
largely composed of nerve-fibres emanating from the pleural 
ganglia. It is interesting to notice that in the Neritidee 
and Helicinidis the commissure connecting the pleural 
g;anglia, instead of traversing the pedal ganglia, as it seems 
to do in most forms, remains quite separate from thein." 

The pleural ganglia are oval structures less than half the 
size of the pedal ganglia. They are situated more laterally 
than the other ventral ganglia, but overlap the posterior part 
of the outer sides of the pedal ganglia, with which they are 
intimately united by extremely short connectives. PI. XIV, 
fig. 68, shows a section of the right pleural ganglion just 
behind the pleuro-pedal connective. 

The two visceral ganglia lie partly between and partl}^ 
behind the pleural ganglia, to which they are joined by very 
short connectives. An equally short visceral commissure 
unites them. They overlap the hind ends of the pedal 
ganglia, but it need hardly be said that they have no direct 
nervous connection with these centres. Both visceral ganglia 
are larger than the pleural ganglia and considerably smaller 
than the pedal ganglia, but the left is always slightly larger 
than the right, and gives off a larger number of nerves. For, 
Avhile the right one is simply the right parietal or supra- 
intestinal ganglion, the left is to be regarded as formed by 
the union of the left parietal ganglion with the median 
abdominal ganglion and is therefore composite. And it is 
only because the right parietal ganglion is usually much 
larger than the left that the difference in size between the 
two visceral ganglia of Apera is not greater. A similar 
fusion between the left parietal and the abdominal ganglia 
occurs in the Helicid^e, but so far as I am aware the only 

' Pelseneer, P.. 'Mem. Acad. Roy. Belg.,' 1901, ex vol. liv, pp. 43-45. 
- Bourne, G. C, 'Proc. Zool. Soc. Lond.,' 1908, pl.lv, figs. 86,87, 
pi. Ivi. fig. 38; and 1911, pp. 791, 792, pi. xxxviii, figs. 49, -50. 



148 HUGH WATSON. 

carnivorous form in which this has hitherto been observed is 
Rhytida ina3qualis [Pfeiffer]} In Sele nochlamy s,~ 
Testacella, Euglandina, Streptostyla,'^ Salasiella,* 
and Paryphanta'^ the three visceral ganglia remain distinct, 
and I have found that this is also the case in Natal in a and 
in Rhytida capillacea [Fer.). In Daudebardia the 
abdominal is fused with the right parietal ganglion.^ Lastly, 
in the Streptaxidte, the three visceral ganglia are all more 
or less fused to form a single mass, which is separated from 
the pleural ganglia by long connectives.^ 

Some of the nerve-cells in the ventral group of ganglia are 
remarkably large, especially those in the posterior parts of 
the visceral ganglia (PI. XIV, fig. 67). One of the cells in the 
right parietal ganglion of a specimen of Apera gibbon si 
rubella measures "IS mm. X '143 mm., and its nucleus is no 
less than '16 mm. x -11 mm. Fig. 69 represents a photo- 
micrograph of a section through one of these large cells in 
the right parietal ganglion of A. dimidia, and shows the 
granular appearance of the large nucleus and its prominent 
nucleolus. 

The Nerves of the Ventkal Ganglia. — The numerous 
pedal nerves arise in an irregular longitudinal line along the 
ventral surface of each pedal ganglion (PI. XV, figs. 73, 74), 

^ Fischer, P.. ' Joiirn. de Concliyl.' (3i-d ser.), 1873. vol. xiii. p. 8. 
pi. iii, fig. 6. 

^Siraroth. H.. 'Festschrift Leucharts,' 1892, p. 55. pi. vi. fig. 15. 
According to Simroth. the three visceral ganglia also remain separate 
in Phrixolestes, but in the other genera of the Trigonochlaminse 
they tend to fuse with one another, and even with the pleui'al ganglia 
in Trigonochlamys itself (seeBronn's 'Klassenii. Ordn. Tier-Reichs 
III, Gastr. Pulmonata,' 1910, p. 257, fig. 80). 

* Crosse and Fischer, ' Mission scientifique au Mexique.' 1878. pi. iv. 
fig 5. 

■* Strebel. H , " Beitrag z. Kenntniss d. Fauna Mexik. L.-n. Si'isswasser- 
Conchylien.' 1878, vol. iii, pi. x, fig. 7. 

" Beutler, op. cit., p. 403, pi. xxix, fig. 59. 

^ Plate, op. cit., p. 590, pi. xxxvi, figs. 94, 97. 

7 Wiegmann, F., ' Mitt. Zool. Samml. Mus. Nat. Berlin." 1898. vol. i. 
p. 61. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 149 

and are distributed to all parts of the foot. The most 
posterior nerves, which arise from the extreme hind end of 
the ganglia, are the largest, and run back for almost the 
whole length of the animal, as will be seen from fig. 75. 
This figure also shows that the other pairs of pedal nerves are 
not arranged quite symmetrically. The pedal gland is inner- 
vated by a pair of nerves which bi'anch from the inner sides 
of the most anterior pair of pedal nerves. The size of 
these nerves to the pedal gland is in direct proportion to 
the size of the gland itself; thus in Apera dimidia, with its 
highly developed pedal gland, the nerve is larger than in A. 
sexangula, as will be seen by comparing figs. 73 and 74. 

From the sides of the ventral group of ganglia a large 
number of nerves radiate to the skin on each side of the 
body and head, passing below the tentacular retractors, but 
above the vagina and the adjacent part of the vas deferens on 
the right side (PI. XIII, fig. 57). These are the nerves that 
de Lacaze-Duthiers termed the "nerves of the neck." The 
most posterior of these nerves can be seen running back on 
each side along the inner surface of the body-wall sometimes 
nearly as far as the diaphragm, thus confirming the hypothesis 
that all this region of the skin corresponds to that of the 
neck of ordinai-y snails and slugs. The posterior ends of the 
common retractors of the tentacles also receive nerves belong- 
ing to this group. All these lateral nerves appear to arise 
by i-epeated branching from three main roots on each side 
of the ventral ganglia. The most anterior root springs from 
the outer edge of the pedal ganglion a little behind the 
cerebro-pedal connective. The second arises from the junc- 
tion of tlie pedal and pleural ganglia — that is to say, from the 
very short pleuro-pedal connective. The third and most 
posterior root arises from the pleural ganglion itself. Fischer's 
figures of the nervous system in Rhytida ina^qualis {Pfi'-) 
and the Oleacinidie also show nerves arising from the 
pleural ganglia, but according to de Lacaze-Duthiers, Plate, 
Wiegmann, and Beutler, no nerves arise from the pleural 
ganglia in Testacella, Daudebardia, Ennea (Eden tu- 



150 HUGH WATSON. 

lina) and Paryphanta.^ It must be remembered that at 
least some of these authors believed that the pleural ganglia 
never did give rise to nerves in the Pulmonata, a belief 
that Amaudrut- has shown to be altogether erroneous. At 
the same time it is unlikely that all these careful observers 
would have figured the "nerves of the ueck" as arising 
solely from the pedal ganglia, if in all carnivorous forms they 
arise partly from the pleural ganglia, as they undoubtedly 
do in Apera. Moreover the fact that the central roots of 
these nerves arise in Apera exactly from the junction of 
the pedal and pleural gauglia also suggests that the more 
posterior " nerves of the neck " arise indifferently either 
from the outer side of the pedal ganglia or from the plem-al 
ganglia. This seems to uphold the theory that the outer 
dorsal portions of the so-called pedal ganglia of most Gas- 
tropods are in reality parts of the pleural nerve-centres 
which have become united with the pedal ganglia, and that 
all the ''nerves of the ueck" therefore spring from the 
pleural division of the central nervous system. The composite 
nature of the pedal ganglia has been shown to be specially 
evident iti some of the most primitive Gastropods, such as 
Pleurotomaria; -^ and the fact that the so-called pedal 
commissure contains, as we have seen, nerve-fibres emanating 
from the pleural ganglia affords additional evidence in favour 
of this theory.'* 

The nerves arising from the visceral ganglia are fewer 

' For references, see p. 148. 

2 ' Ami. Sfi. Nat., Zool.,' 1898, vol. vii, p. 128. 

^ Bouvier and Fischer, ' Journ. de Concliyl.,' 1899, vol. xlvii, pp. lU9- 
143. 

■* It is interesting to note that in Aplysiella and some species of 
Aplysia, as well as in the Gymnosomata, the cervical nei-ves also 
arise partly from the pleural and pai'tly from the pedal ganglia, although 
they anastomose to form a single plexus (Pelseneer, " Mem. Couronne 
Acad. Roy. Belg.,' 1893, ex vol. liii, p. 27, pi. x, fig. 81); while in the 
AuriculidEe these nerves may arise either from the pleural ganglia or 
from the pleuro-pedal connectives (Bonvier, ' Comptes Rendus Soc. de 
Biologic' (9th ser.), 1892, vol. iv. p. 990). 



THE OAENIVORUUS SLUGS OF SOUTH AFRICA. 151 

than ill most genera, probably because they innervate a 
region of the body which is greatly reduced in Apera. The 
origin of these nerves is shown in PI. XV, figs. 73 and 74, 
and their distribution iu A. dim idia will be seen fi'om fig. 75. 

One important nerve arises from the right parietal or supra- 
intestinal ganglion, not far from its junction with the other 
visceral ganglion, and innervates the right wall of the mantle- 
cavity. This may be regarded as the right pallial nerve. 
The corresponding left pallial nerve is more slender, and 
arises from the outer or parietal portion of the left visceral 
ganglion. This nerve innervates the left wall of the mantle- 
cavity. From the median or abdominal portion of the left 
visceral ganglion two large nerves arise close together. That 
to the left becomes closely united for some distance with the 
buccal retractor in Apera dimidia, and then runs back on 
the right side of the rectum, and eventually branches around 
the anus and respiratory orifice. The right abdominal nerve is 
joined to the aorta for almost the whole of its length, but at its 
hind end the nerve leaves the blood-vessel and enters the 
diaphragm, which it probably innervates. Beyond this the 
nerve becomes attached to the adjacent wall of the peri- 
cardium, and can be traced backwards to a point just behind 
the opening of the reno-pericardial duct. The other nerves from 
the visceral ganglia are also attached to the aorta for some 
distance, the left pallial nerve being the first to become free. 
One or two small additional nerves can be seen with a stronsr 
objective to leave the visceral ganglia and run along in the 
tissue surrounding the aorta, but these are so extremely fine 
that I was unable to trace them far. Thus only four nerves 
of any importance arise from the visceral ganglia in Apera — 
two pallial and two abdominal ; and inasmuch as the abdominal 
portion of the left ganglion is probably itself formed from 
the union of the sub-intestinal ganglion with the original 
median abdominal ganglion,^ it follows that there is but one 
nerve corresponding to each original ganglion on the visceral 
loop. 

' Pelseneer. P.. 'Mem. Acad. Roy. Belg.,' 1901, ex vol. liv. pp. 47. 48. 



152 



HUGH WATSON. 



The Nervous System as a Whole. — It will have been 
seen from the above account that the central nervous system 
of Apera may be regarded as formed of five sections, each 



Text-fig. 2. 




f Cerel.ral. ^ Pedal. 

■i Buccal. C] Pleural. 

^ Visceral. 

Diagram of the central nervous system of Apera. 

with a single pair of ganglia. These five sections are dia- 
grammatically represented in the accompanying text-figure. 
There is first the buccal section, which is the most anterior 
only in A. burnupi and A. sexangula. This innervates 
the anterior part of the alimentary canal, including the 
odontophore and salivary glands. Secondly, there is the 



THE CARNIVOROUS SLUGS OP SOUTH AFRICA. 15-'^ 

cerebral division, which is connected with the sense-organs of 
the head/ and is the only section in which the nerve-cells are 
aggregated to form ganglia on the dorsal side of the alimentary 
canal. Thirdly, there is the pedal section, which innervates 
the foot and pedal gland. Fourthly, there is the pleural 
section, innervating the sides of the body and head. And 
fifthly, there is the visceral section, which innervates the 
neighbourhood of the mantle-cavity. 

And just as there are five pairs of ganglia, so also are there 
five commissures ventral to the alimentary canal, as will be 
clearly seen from the diagram. Now some authors have 
thought that there were only three such commissures in the 
P u 1 m o n a t a, and very few have found more than four. Must 
we then regard Apera as exceptional in this respect? I 
think not. For it seems probable that further researches will 
show that there are normally five commissures ventral to the 
alimentary canal in the P^uthyneura, although the slender 
sub-cerebral commissure may have entirely disappeared in 
some Pulraonates, just as in the Strep toneura the corre- 
sponding labial commissure seems to have disappeared in many 
of the Pectinibranchia. 



THE STEUCTURE OF THE UPPER TENTACLES, AND 
THE TENTACULAR RETRACTORS. 

In PI. XVI, the figs. 80-86 represent photomicrographs 
of serial sections through the left upper tentacle of Apera 
dimidia in its retracted condition. It will be seen that 
the olfactory organ is large and extends forwards in front 
of the eye, thus reducing the width of the cavity in the 
retracted tentacle. The eye itself appears to be of the 
usual structure, the small size of the lens being possibly due 
to contraction. The retina is very deeply pigmented. The 

^ It is not improl)able that the motor nerve to the buccal retractor 
should be regarded as belonging to the pleural section, for it arises 
close to the origin of the cerebro-plenral connective. 



154 HUGH WATSON. 

nerves from the different parts of the olfactory organ 
converge and unite behind the level of the eye to form the 
large olfactory nerve, which runs back for a considerable 
distance in the centre of the posterior part of the tentacle. 
The much smaller optic nerve is situated laterally, being- 
embedded in the muscle-fibres on the outer side of the olfactory 
nerve. These longitudinal muscle-fibres make their appear- 
ance just behind the sense-organs, and are chiefly restricted 
to the peripheral part of the retracted tentacle ; but as they 
ai'e traced backwards they gradually increase in number, and 
become united into muscular strands, which coalesce with one 
another to form the retractor of the tentacle . In the spaces 
between these strands, before they all coalesce, large cells 
occur, as will be seen from fig. 84. These large cells stain 
rather deeply, and have rounded nuclei, containing prominent 
nucleoli. They are evidently homologous with the somewhat 
similar cells that Beutler^ found in the tentacles of Pary- 
phanta hochstetteri [Pfr.). In this region the tentacle is 
rather darkly coloured ; yet the pigment-granules are not 
contained in these large cells, but are confined to narrow 
filaments which surround the cells and the different strands 
of muscle. Moreover the pigment extends a little further 
back, even slightly beyond the point shown in fig. 86, where 
the olfactory and optic nerves emerge from the tentacular 
retractor and curve over to the cerebral ganglion (PL XIII, 
figs. 52-55). 

Not more than 2 or 3 mm. further back the retractor of the 
upper tentacle is joined by that of the lower tentacle, which 
is narrower and lies more ventrally. The common retractor 
thus formed is of no ffreat lenarth. It arises from the side of 
the body-wall between a quarter and a third of the distance 
from the head to the hind end of the animal. The common 
tentacular retractors of each side have no connection Avith 
each other or with the buccal retractor ; on the contrary, the 
three retractors have become unusually widely separated in 
their origin. On the right side the retractors of the upper 
' ' Zool. Jahrb..' 19()1. vol. xiv, p. 404, pi. xxix, figs. 63, 64. 



THE CAKNiVOKOUS SLUGS OF SOUTH AFEICLV. 155 

and lower tentacles are separated by tlie penis, which projects 
between them (PI. XIII, figs. 52-57; see p. 190). 

In one specimen of Apera sexangula I found that the 
retracted upper tentacles were joined by a transverse connec- 
tion slightly anterior to their junction with the olfactory and 
optic nerves (PI. XV, fig. 76). The transverse connection 
was pigmented in the same way as the tentacles themselves. 
This remarkable abnormality reminds one of the more striking 
monstrosity figured by Forbes and Hanley/ in which the 
upper tentacles of a specimen of Agriolimax agrestis 
{Lin.) were united throughout their entire length. 

The posterior ends of the common tentacular retractors 
receive nerves arising from the pleural ganglia (or possibly 
from the short pleuro-pedal connectives) ; and the retractors 
of the lower tentacles are innervated, as already described, 
by nerves issuing from the sides of the nerve-collar between 
the cerebro-pedal and cerebro-pleural connectives, but pro- 
bably emanating from the cerebral ganglia by the same roots 
as the nerves to the buccal retractors. 

Beneath the tentacular retractors a few smaller muscles 
arise on each side and run forward to the anterior part of the 
head. These are best seen in PI. XIII, fig. 57. The largest 
of these minor cephalic retractors is inserted in the upper lip 
of the pedal gland. It is doubtless the contraction of these 
muscles, together with the combined action of the buccal and 
tentacular retractors, which causes the front of the head to 
be invaginated in some specimens. 



THE DIGESTIVE SYSTEM. 

The Mouth and the Eadula-sac. — As in most carnivorous 
genera, the mouth has three main lips, one dorsal and two 
lateral, arranged like the sides of an inverted isosceles 

1 ' Hist. Brit. Mollusca,' 1853, vol. i, pi. JJJ, fig. 4. 

VOL. 3, PART 2. 12 



156 HUGH WATSON. 

triangle. But it will be seen from PI. XV, fig. 11 , and 
PI. XVI, fig. 88, that the buccal cavity has four sides, and 
only appears somewhat triangular owing to the narrowness of 
the ventral wall. These figures also show the great thick- 
ness of the sides of the buccal mass — a feature which is still 
better seen in PL XVI, fig. 87. Externally there is a layer 
of longitudinal muscles, which is thin in Apera dimidia 
and A. sexangula, but somewhat thicker in A. gib- 
bons!. Within this there is a very thick layer consisting 
chiefly of circular muscles, but also containing some radial 
and a few longitudinal fibres. The last become more numerous 
towards the inner surface, so as almost to form a third 
muscular layer in which the longitudinal predominate over 
the circular fibres. Lastly, the cavity is lined by a compact 
cubical epithelium which is continuous over the lips with the 
epidermis of the skin. The cells composing this epithelium 
are small, and their nuclei are situated towards their outer 
ends — that is to say, the ends turned away from the cavity. 
Towards the opening the epithelial cells become taller, 
and the translucent cuticle which they secrete is consider- 
ably thickened on the lips. There is, however, no jaw in 
Apera. 

At its hind end the buccal cavity extends into a long 
tubular diverticulum, which opens into the centre of the 
posterior wall of the cavity. This is the radula-sac, and is 
shown in longitudinal section in PL XVII, fig. 92, and in trans- 
verse section in PL XVIII, figs. 116-120. It will be seen that 
a broad fold or cushion projects like a typhlosole from the 
upper wall into the cavity, and renders the latter crescentic 
in section. Towards the hind end of the radula-sac the edges 
of this cushion are spirally rolled (fig. 119). Further forward a 
narrow longitudinal ridge occurs dorsally at each side of the 
cushion. The epithelium on the sides and floor of the I'adula- 
sac consists of small cubical cells, compactly arranged, and 
not unlike those lining the buccal cavity itself ; but the epi- 
thelium on the cushion is of a quite different character, 
being composed of very tall and remarkably irregular cells 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 157 

(PL XVI, fig. 89). The centre of the cushion is loosely 
filled with connective tissue. Immediately surrounding the 
whole radula-sac is a thin layer of circular muscles. 

Around the periphery of the hind wall of the buccal cavity 
there is a circular groove, which is deepened dorsally and 
ventrally (PI. XVIII, figs. 115, 116). The dorsal pocket is of 
no great depth, and is connected with the opening of the 
radula-sac by a median vertical groove. The ventral pocket, 
on the other hand, is much deeper, and contains the anterior 
end of the radula, which is thus folded back below the 
remaining part in the radula-sac (PI. XVII, fig. 92) . Between 
the opening of the radula-sac and that of the ventral pocket 
there is a slight projection formed by the front end of the 
odontophoral support covered by the radula, and it is this 
projection that is the first to appear when the odontophore is 
protruded through the mouth. 

The Radula. — The general appearance of the radula, when 
freed from the surrounding tissue, is shown in PI. XVII, figs. 
97, 98, 99, 102, 107, and 114. It is long and relatively narrow, 
its breadth being sometimes less than one-sixth of its length. 
The teeth are thorn-shaped with narrow bases, and they are 
arranged in rows which diverge obliquely forwards on each 
side of the middle line. In other words, the radula of Apera 
is of the same general type as is found in the Rhytidida3, 
the 01eacinida3, the Testacellidae, and, indeed, in 
all the families of carnivorous snails and slugs excepting 
the Strep taxi das, in which the teeth are of a slightly 
diiferent shape. Nevertheless, the radula of Apera is excep- 
tionally interesting, and shows a diversity which is scarcely 
surpassed in any genus of the Pulmonata containing so few 
species. 

In the first place, the size of the radula varies enormously. 
In Apera gibbonsi and A. parva the radula is extremely 
large ; when flattened out it measures about a third of the 
total length of the animal, and in the former species it is pro- 
portionately broader than in the other members of the genus. 
In A. dimidia the radula is scarcely a sixth of the length 




158 HUGH WATSON. 

of the animal. In A. pure el li it is about a ninth, while in 
A. burnupi and A. sexangula it is only about a twelfth 
of the length of the slug. 

The diversity in the size of the teeth is even greater. In 
Apera burnupi and A. sexangula the animal is about 
550 times the length of the largest tooth in its radula. In A. 
dimidia and A. pur colli the proportion is 335 : 1. In a 
typical example of A. gibbonsi it is about 115 : 1 ; while in 
A. parva and A. gibbonsi lupata the animal is only about 
66 times as long as its largest tooth, which attains a length of 
more than 0*6 mm. in the latter form. 

But the diversity found in the fadula of Apera extends to 
other features besides its dimensions. Even in the closely 
related forms which I am provisionally regarding as subspecies 
of A. gibbonsi, there is a remarkable variation in the form 
and number of the teeth (PI. XIX, figs. 123, 124, PI. XX, 
125, 126, and text-fig. 3, a-e). The typical form has the 
most primitive type of radula, and the others show a pro- 
gressive adaptation to vermivorous habits. A slug that feeds 
on worms — and it is known that this species does so — uses its 
radula, not for rasping off pieces from its victim, but for 
catching hold of it and drawing it back through the mouth 
into the crop. For this purpose the teeth must be large, with 
long sharp points for piercing the worm's skin and firm cuticle; 
and secondly, the teeth must be shaped so as to retain their 
position in the skin of the worm when they are pulling it back 
into the mouth. Accordingly we find, first, a progressive 
increase in the size of the larger teeth compared with the size 
of the animal, and especially in the length of their cusps. 
And in order to make room for these the number of rows is 
diminished, and the smaller teeth towards the outer edges of 
the radula, as well as those down the centre, become still less 
and eventually disappear (cf . PI. XX, fig. 125, and PI. XIX, 
fig. 123, and see also the numbers of the rows, and of the 
teeth in each row, given on pp. 195-201). Again, we find that 
the attachment of the teeth to the basal membrane of the radula 
also increases in length and forms a projecting apophysis, thus 




. g'ibbonsi, s.s. 

. g. rubella. 

. g. gracilis. 

. g. In pat a. 

. g. lupata V. duplex. 



.1 .2 .3 A- .5 b .7 .8 .9 Imm 

1 ' Ill 

Representative teeth fro in the radulse of A per a. 



160 HUGH WATSON. 

making it more difficult for the teeth to be pulled out of posi- 
tion and keeping- them rigidly in place. Lastly, the teeth 
become modified in one of two ways in order to prevent the 
worm that they have transfixed from escaping. In A. gib- 
bonsi rubella the cusps are strongly curved so as to form 
veritable hooks (text-fig. 3, b), and they are also slightly 
broadened towards the points. On the other hand, in A. 
gibbon si lupata the cusps are bai'bed on the lower side, as 
in Testacella — a remarkable case of parallel evolution (cf. 
PL XXIV, fig. 157, and text-fig. 3, d). Still more highly 
specialised is the var. duplex of the same form. In this 
variety all the larger teeth are doubly barbed, there being a 
small barb on the upper side of the cusp nearer to the point 
than that on the lower side (text-fig. 3, e). I do not know 
of any other Pulmonate in which the radula has become so 
highly specialised along these lines as it has in this variety. 
We have to go to the marine carnivorous genera, such as 
Conus, to find such formidable doubly barbed teeth, and here 
also we notice that one barb is nearer the point of the tooth 
than the other. In Conus, however, the barbs are pointed 
instead of being merely square as in Apera, and the distal 
barb appears to have been developed first, for the other is 
often absent or merely represented by a flange. And of 
course the bases and arrangement of the teeth in Conus are 
quite different. 

The radula of Apera parva bears a general resemblance 
to that of A. gibbonsi, but the bases of the teeth are not 
so narrow, the cusps of the inner teeth are slightly shorter, 
and the teeth become relatively larger towards the edges of 
the radula (text-fig. 4, a). None of the transverse rows 
contain more than thirty-five teeth in this species, and some 
have only thirty-four, as the vestigial central tooth is absent 
from some of the rows. On the Avliole the radula of A. 
parva is less unlike the type found in the genus Rhytida 
than are the radula of the other species of Apera. 

In Apera purcelli and A. dimidia the central tooth 
also degenerates. It is present, though small, in A. purcelli 



Text-pio. 4 



A. pai'va. 
(X lUO.) 



A . d i m i d i a . 

(X -M).) 



A . p u r c e 1 1 i . 

(X 4UU.) 



A. l)iirnupi. 

( X -iijo.) 



A. sexangula. 

(X 2U0.) 




Representative teeth from the radul* of Apera. 



162 HUGH WATSON. 

(text-fig. 4, c), but no trace of it remains in A. dimidia 
(PI. XX, fig. 127, and text-fig. 4, b). The outer teeth of 
these two species have rather long, slightly curved cusps, and 
are not very unlike the teeth of the typical form of A. gib- 
bon si on a smaller scale; but the cusps of the inner teeth 
are remarkably short and broad, and the teeth have therefore 
a quite different appearance. In the more primitive species, 
A. purcelli, only the first three or four teeth on each side are 
thus modified, but in A. dimidia half the teeth are of this 
shape. Hence the teeth of these species, unlike those of nearly 
all other carnivorous forms, are differentiated into laterals and 
marginals, although there are one or two on each side which 
might be regarded as transitional. The figures of the radula 
of A. dimidia (PI. XX, fig. 127, and text-fig. 4, b) show that 
in this species the corresponding teeth on each side are not 
opposite to each other, the right half of the row being in front 
of the left. A similar displacement occurs in A. parva 
(text-fig. 4, a), and it is also often observable in A . gibbon si 
(PI, XX, fig. 125) ; but it occurs occasionally in other carni- 
vorous genera, for I have noticed it in Test ace 11a halio- 
ti dea Drap., T. scutulum Soiv., Euglandina truncata 
{Gmel.), E. corneola {Binn.), and Rhytida franklandi- 
ensis {Forhes). 

In Apera sexangula and A. burnupi the difi^erentia- 
tion into lateral and marginal teeth is still more marked (PI. 
XX, fig. 128 ; text-fig. 4, d, e). The marginal teeth are more 
than twice as numerous as the laterals, and their cusps are 
long, slender, and nearly straight, those of one row over- 
lapping the bases of the teeth in the row behind. The 
lateral and central teeth have short broad cusps, and the 
central tooth — which is only slightly smaller than those on 
each side of it — is not unlike the lateral teeth of A. purcelli 
and A. dimidia, though perhaps a little shorter. The cusps 
of the lateral teeth, however, are not only very short, but 
they are bifid and end in two points, the inner of which is the 
longer (text-fig. 4, d-f). Beutler^ has shoAvn that in Pary- 
» ' Zool. Jahrb.; 1901, vol. xiv, p. 380, pi. xxvii, fig. 23a. 



THE C'AENIYOEOUS SLUGS 0¥ SOUTH AFKICA. 163 

phanta liochstetteri [Pfr.) one of the teeth in each row 
may abnormally develop a double cusp ; but I believe that 1 
am right in saying- that in no carnivorous genus excepting 
A per a are some of the teeth normally provided with double 
cusps, for I regard the bicuspid teeth of Pseudosubulina 
lirifera {Morelet) as something quite different. What is the 
reason for this remarkable modification of the inner teeth in 
these two species ? Perhaps it may be found that Apera 
burnupi and A. sexangula live chiefly on some special kind 
of food, and have their radula=i specially modified in conse- 
quence. Or possibly we have here an instance of carnivorous 
slugs reverting to a partly vegetarian diet, and the lateral 
teeth are used in eating vegetable and the marginals in 
eating animal food. The fact that Collinge has found both 
animal and vegetable remains in the intestine of A. sex- 
angula is in favour of the latter view. In case this hypo- 
thesis should prove to be correct, it is interesting to compai-e 
the type of radula found in these species with that which is 
characteristic of the Limacidse, Zonitidee, and other 
omnivorous families. These also have developed thorn-shaped 
marginal teeth, very similar to those in Apera, and here, too, 
we find that the lateral and central teeth are shorter and 
broader with one or two less pointed cusps. Nevertheless, 
these inner teeth are very different to those of Apera 
burnupi and A. sexangula, for they are of the ordinary 
type with broad oblong bases, such as is found in the great 
majority of herbivorous snails, whereas in the species of 
Apera the teeth have narrow bases and have obviously been 
derived from the thorn-shaped type. This difference, how- 
ever, could be easily explained by supposing that in the 
Zonitidse, etc., we have a group of herbivorous snails which 
have become adapted to a partially carnivorous diet, while in 
these species of Apera we are dealing with carnivorous slugs 
which have become partially herbivorous. 

Further details about the radula of Apera will be found 
under the head of the different species. I need only add here 
that the teeth towards the hind end of the radula are more or 



164 HUGH WATSON. 

less tinged witli brown, as is so commonly the case among 
the carnivorous genera. 

I^HE Odontophore. — The hind end of the buccal mass is 
prolonged beyond the opening of the oesophagus to form the 
cylindrical muscular odontophore containing the radula-sac. 
In most carnivorous snails the hind end of the odontophore is 
curved abruptly downwards, but in Apera, as in Testacella, 
it has become quite straight, probably owing to the degenera- 
tion of the spiral shell. The size of the odontophore — and, 
to a less extent, the size of the buccal mass in front of it — is 
proportionate to the size of the radula. In A. burn up i 
(PL XXI, fig. 133) and A. sexangula (fig. 134) it is 
quite small, being onl}^ about 4 mm. long; in A. purcelli 
(PI. XIII, fig. 55) and A. dimidia (Pl.'xXI, fig. 132) it is 
much larger; finally, in A. gibbonsi and A. parva 
(figs. 129-131) the odontophore attains an enormous size, 
being one of the largest organs in the body. Yet the 
structure of the odontophore is remarkably constant through- 
out the genus, the differences between the arrangement of its 
muscles in the various species being quite insignificant. 

The odontophoral support or cai-tilage is a semi- 
cylindrical structure which extends along almost the entire 
length of the odontophore. It is crescentic in section, the 
convex surface being ventral. In the hollow formed by its 
upper surface lie the radiila-sac and the muscles immediately 
surrounding it, while the ventral pocket containing the front 
end of the radula is situated beneath the anterior part of the 
support, the radula being folded over its anterior end (PI. 
XVII, fig. 92, and PL XVIII, figs. 115-122). The upper edges 
of the support are parallel throughout the greater part of its 
length, but anteriorly they curve inwards towards each other, 
and then diverge again and slope downwards at the front 
end, thus leaving an oblique oval aperture where the opening 
of the radula-sac is situated (PL XVII, figs. 95, 100, 105, 112). 
As might have been expected, this aperture is largest in 
Apera gibbonsi, and the front end of the support is 
rounded in this species and in A. parva. On the other 



THE CARNIVOROUS SJ.UGS OF SOUTH AFRICA. 165 

handbill A. burnupi and A. sexangula the aperture is 
very small, and the front end of the support is pointed 
(see fig. 112). The hind end of the odontophoral support is 
rather abruptly truncated. Along the inner surface of the 
support there runs a median longitudinal furrow, which can 
be best seen in transverse sections of the odontophore of 
A. dimidia (PI. XVIII, figs. 116-121). In the other species 
it is not usually so well marked. 

The odontophoral support is flexible, but much firmer than 
ordinary muscular tissue, and it is usually semi-translucent ; 
yet it does not contain any cartilage, but is entirely com- 
posed of long narrow cells stretching radially from the inner 
to the outer surface (PI. XVI, fig. 90). The nuclei of these 
cells are also lengthened, and are rather more numerous 
towards the outer than the inner surface of the support, 
doubtless because the cells are on an avei'age slightly broader 
towards the outside. Most authors have regarded these long 
cells which occur in the odontophoral support of carnivorous 
snails and slugs as being muscle-fibres; but Beutler^ has 
disputed this view, and has maintained that in Paryphanta 
hochstetteri {Pfr.) this tissue is not muscular. Now, I 
should certainly not be inclined to consider these cells as 
being of the nature of ordinary muscu.lar fibres in Apera, 
but they might perhaps be regarded as muscle-fibres which 
have become modified for a supporting or skeletal purpose, 
and, if this is so, it is possible that they have become more 
modified in genera such as Paryphanta and Apera than 
in the types studied by Plate. In Testacella and Daude- 
bardia there are longitudinal muscle-fibres and other cells 
intercalated among the radial elements of the support,'^ and I 
have found that this is also the case in Euglandina 
venezuelensis [Preston); but these are entii'ely absent in 
Apera, as in Paryphanta'' and Natalina.^ 

' Op. cit., pp. 380, 381. 
Plate, L. H., ' Zool. Jalirb.,' 1891, vol. iv, pi. xxxiii, figs. 30, 31, 39, 
41, pi. xxxiv, fig. 51. 

^ Bentler, op. cit., p. 380. 

* Woodward. M. F.. ' Proc. Mai. Soc," 1895, vol. i, p. 273. 



166 HUGH WATSON. 

A longitudinal muscle runs along the upper edge of the 
support on each side towards its anterior end (PI. XVI, fig. 
90; PI. XVIII, figs. 116-118), and probably serves to bend 
up the front of the support, as shown in PI. XVII, fig. 108, 
and may therefore be termed the flexor muscle of the odonto- 
phoral support. In Apera gibbon si there is also a very 
thin layer of longitudinal muscle-fibres covering the ventral 
surface of the support, and possibly the flexor muscles 
might be regarded as a special thickening of this layer. 

The most important muscles in the odontophore are the 
powerful retractors, which arise from the odontophoral 
support and are inserted in the radula-sac. These may be 
divided into lateral retractors, median retractors, and terminal 
retractors. The numerous lateral retractors are attached in 
front to the radula-sac, chiefly around its anterior end, and 
pass obliquely backwards and outwards on each side, curving 
over the edges of the support and becoming attached to its 
outer sides (PI. XVII, figs. 92, 93,96, 101, 106, 109, 111, 113, 
and PL XVIII, figs. 116-121). The median retractors are a 
pair of specially thick muscles, which are attached for some 
distance to the floor of the radula-sac towards its anterior 
end, and pass backwards to the extreme hind end of the 
support (PL XVII, figs. 92, 106, and PL XVIII, figs. 118-122). 
They belong to the same series as the lateral retractors, of 
which they might be regarded as forming the innermost pair. 
No muscles are attached to the posterior part of the radula- 
sac, excepting at its hind end, where there is another pair of 
retractors, which I am calling the terminal retractors. These 
are not so thick as the median retractors, and pass backwards 
above them to the hind end of the support (PL XVII, fig. 92, 
and PL XVIII, figs. 121, 122). All these retractor muscles 
have the same function — that of pulling back the radula. 
In A. gibbonsi some of the more posterior strands of 
muscle are attached to the inner surface of the support, close 
to its hind end, while others are not attached to the support 
at all, but to the hind end of the sheath of the odontophore 
opposite to the insertion of the extrinsic buccal retractors. 



THE CARNIVOROUS Sl-UGS OF SOUTH AFRICA. 167 

Numerous muscles arise in two rows, one on each side of 
the mid-dorsal line of the outer sheath of the odontophore, 
and, passing' downwards outside the lateral retractors, are 
attached to the outer sides of the support just below them 
(PI. XYII, fig-s. 91, 104, and PI. XYIII, figs. 115-121). These 
muscles I am terming the suspensor muscles of the support. 
They are nowhere of any great thickness, but are decidedly 
thinner behind than in front. 

A thin layer of ventral muscles arises from the hind end of 
the odontophoi-al support, and is inserted in the anterior part 
of the radula, or rather in the walls of the ventral pocket 
which contains it (PI. XVII, figs. 91, 92, 101, 104, and PI. 
XVIII, figs. 117-121). These muscles do not only run longi- 
tudinally beneath the centre of the support, but the more 
lateral strands diverge radially from the ventral pocket of the 
radula and unite with the suspensor muscles of the support 
on each side. They serve to pull back the anterior part of 
the radula over the fi-ont edge of the support, and thus to 
pull forward the radula-sac when its retractors are relaxed, 
so that the radula assumes the form shown in PI. XVII, figs. 
92, 97, 102, 1 14. Then, when the teeth have become imbedded 
in the skin of the prey, these ventral muscles will be relaxed, 
and the powerful retractors will draw back the radula until 
it has the form shown in fig. 98. 

A slender median dorsal muscle is inserted in the sheath of 
the odontophore, between the suspensor muscles, a short 
distance behind the opening of the oesophagus, and is 
attached posteriorly to the front end of the terminal 
retractors, its sides also becoming connected with the neigh- 
bouring lateral retractors (PI. XVII, fig. 92, and PL XVIII, 
figs. 116-120) . This is evidently homologous with the muscle 
that Amaudrut terms the "papillaire superieur." It 
may assist the ventral muscle in pulling forward the radula- 
sac, but its chief function is probably to pull back the 
posterior lip of the opening of the oesophagus when the 
retractor muscles are brought into play, thus making a wide 
passage towards the crop. 



168 HUGH WATSON. 

The slieath of tlie odontopliore is composed of three layers 
of muscle-fibres. Externally there is a thin layer of longi- 
tudinal muscles; within this lies a layer of circular muscles, 
which is equally thin throughout the greater part of the 
length of the odontophore, but becomes a little thicker 
toAvards the front end ; lastly, there is a very thin layer of 
longitudinal muscles lining the whole of the sheath excepting 
the dorsal part which lies between the attachment on each 
side of the suspensor muscles of the support. At its hind 
end the sheath of the odontophore becomes intimately united 
with the support and the posterior ends of the retractor 
muscles, but throughout nearly the whole of its length its 
only connection with the underlying tissues is by means of 
the suspensor muscles. 

I have examined the arrangement of the muscles of the 
odontophore in all the species of Apera excepting A. 
purcelli, and the amount of variation is so small, notwith- 
standing the diversity of the radulae, that it must be admitted 
that these muscles are of considerable systematic importance. 
It will, therefore, be of special interest to compare the 
arrangement of the odontophoral muscles of Apera with 
that which is found in other families of carnivorous snails 
and slugs. 

As an example of the Rhytidid^e we may choose 
Natalina. Nineteen years ago M. F. Woodward^ published 
an account of the anatomy of Natalina caffra {Fer.), Avith 
special reference to the structure of the odontophore ; but 
unfortunately his account shows evidence of inaccuracy. I 
shall therefore ignore his description and figures, and compare 
the odontophoral muscles of Apera with those of a species 
of Natalina, nearly allied to N. caffra, which I have 
been able to examine myself, namely N. quekettiana {M. & 
P.). On the whole the odontophore of this species resembles 
that of Apera very closely; it differs, however, in the 
following features. — (1) Instead of being straight the odonto- 
phore is slightly curved towards the left ; moreover its hind 
' ' Proc. Mai. Soc.,' 1895, vol. i, pp. 270-277, pi. xvii. 



THE CAENIVOKOLTS SLUGS OF SOUTH AFRICA. 169 

end is abruptly Lent downwards, and the outer longitudinal 
muscles of the sheath form a pad extending from the recurved 
end a short distance forwards along the ventral surface. (2) 
The lateral retractors towards the posterior end of the 
odontophore are attached to the odontophoral support just 
within its edges ; further forward they are attached to the 
outer sides of the support as in Apera. (3) The median 
dorsal muscle is extremely slender, and the terminal retractor 
is also very weak, being almost devoid of muscular fibres. 
(4) The circular muscles of the sheath are much thicker 
laterally and ventrally, and the suspensor nmscles of the 
support are also much thicker than in Apera. In all other 
respects the structure of the odontojjhore is strikingly similar 
to that found in Apera, and especially to that of A. 
gibbonsi; for in Natalina quekettiana a thin layer of 
longitudinal muscles covers the lower surface of the odonto- 
phoral support, and some of the posterior retractors are not 
attached to the support, but arise from the sheath of the 
odontophore opposite to the insertion of the upper branches 
of the extrinsic buccal retractors — features which are also 
found in A. gibbonsi. 

The chief differences mentioned above may be easily 
explained. The curvature of the odontophore is doubtless 
due to the fact that Natalina possesses a large spiral shell. 
The greater thickness of the constrictor muscles running in a 
circular direction, both those in the sheath and those beneath 
it, is probably connected with the circumstance that Natalina 
feeds on snails rather than on worms ; for, as Woodward has 
pointed out, the contraction of these muscles will press the 
radula against the body of its prey, as it tears the teeth 
through its victim's flesh, a thing that a vermivorous form does 
not do. The reduction of the median dorsal muscle and the 
terminal retractor, which might be regarded as its continua- 
tion, may be due to the fact that Natalina rasps off portions 
of its prey, and therefore does not require to enlarge the 
opening of the oesophagus to such an extent as an animal 
that swallows worms whole. We see, therefore, that the 



170 HUGH WATSON. 

differences between the structure of the odontophore in 
Apera and Natalina are of little morphological impor- 
tance, and are certainly less striking than the differences 
between the raduljB of the two genera, for the radulse of 
Natalina and Apera have become highly specialised in 
different directions. 

Turning now to the TestacellidfC; we find slugs which 
are vermivorous like Apera gibbon si and have radulas 
remarkably similar to the var . lupata of that species. Yet an 
examination of the odontophore ofTestacella maugei Fer. 
reveals many features in which the arrangement of its muscles 
is quite different from anything that we have found in Apera 
or Natalina (PL XXIV, figs. 155, 156). The structure of the 
odontophore ofTestacella maugei differs from that which 
I have described as characteristic of Apera in the following 
respects. — (1) There is no terminal retractor (unless it is 
represented by the small connections between the floor of the 
radula-sac and the median retractor). (2) Perhaps in conse- 
quence of this the radula-sac is shorter than the ventral 
pocket, instead of being much longer. (3) The median dorsal 
muscle is short, very oblique, and divided into three or four 
line strands. (4) The median retractors curve together above 
the radula-sac to form a tube which is continued to the hind 
end of the odontophore. (5) The muscles which form the top 
of this tube are not only firmly attached to the lateral retractors 
on each side, but also to the sheath of the odontophore ; and 
they are continuous with the extrinsic buccal retractors, which 
join the posterior half of the odontophore on each side of the 
mid-dorsal line. (6) The lateral retractors are scarcely 
divided into separate strands. (7) There seem to be no flexor 
muscles along the upper edges of the support, but on its inner 
surface there is a slight median longitudinal ridge which dis- 
appears towards the hind end and is connected with the 
median retractors by a few slender strands. (8) The ventral 
muscles are rather stouter than in Apera and more definitely 
split up into separate strands. (9) Anterior to these the 
suspensor muscles are free, but directed obliquely downwards 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 171 

and backwards from the sides of the roof of the odontophore 
to the support, parallel to the lateral retractors ; further 
back, however, they are either absent or completel}' fused with 
the lateral retractors. Taking into consideration the similarity 
of the radula of Testacella to that of Apera, it is surpris- 
ing that there should be so many diiferences in the muscles 
which control its movements. 

In the Oleacinidas the odontophore resembles that of 
Testacella in some of its features, and that of Natalina in 
others, although it often possesses more primitive characters 
than can be found in either of these genera. StrebeP has 
given a detailed account of the odontophore of Euglandina 
sowerbyana (Pfr.), from which it will be seen that in this 
species the median retractors enclose the hind end of the 
radula-sac and become continuous posteriorly with the 
extrinsic buccal retractor. In this respect, therefore, Eug- 
landina sowerbyana resembles Testacella. But this 
is not the case with many of the other forms, such as fj . 
venezuelensis {Preston) or Streptostyla shuttle- 
worthi {Pfr.)," for in these the radula-sac is long, and its 
posterior part has not yet become entirely surrounded with 
muscles, but projects slightly from the hind end of the odonto- 
phore. In most genera the posterior end of the odontophore 
is curved downwards as in Natalina. The median dorsal 
muscle resembles that of Apera, and the ventral muscles are 
also like those of Apera and Natalina. On the other hand, 
the suspensor muscles are like those of Testacella, and I 
found that Euglandina venezuelensis also resembled that 
genus in having the retractors fused with the sheath of the 
odontophore towards the hind end. No median ridge or 
furrow occurs along the inner surface of the support in 
E. venezuelensis, and I did not find any flexor muscles 
along its edges. Possibly the longitudinal muscle-fibres 
within the support take the place m Testacella and 

' ' Beitrag z. Kenntniss d. Fauna Mexikanischer L. u. Siisswasser- 
Conchylien,' 1878, vol. iii, pp. 40-42, pis. xvii, xviii. 
- Strebel, op. cit., pi. v, fig. 7a. 
VOL. 3, PART 2. 13 



172 HUGH WATSON. 

l^]nglandina of\he flexor muscles in Apei'a and Natalina. 
It is interesting to notice that while the odontophore in the 
Oleacinid^ is usually curved as in the RhytididjB, in 
several features of its internal structure it appears to resemble 
the odontophore of Testacella more than does that of 
A p e r a . 

The Buccal Reteactors and Protractors. — In all the 
species of Apera excepting A. gibbon si and A. parva, the 
extrinsic buccal retractor consists of a long, narrow muscle, 
arising from the right side of the floor of the body-cavity at 
the entrance to its funnel-shaped prolongation beneath the 
mantle-cavity or lung. The origin of the buccal retractor is 
thus nearly as far back as the heart, and is slightly asym- 
metrical (PI. XV, fig. 75). In front the muscle bifurcates just 
before reaching the odontophore, and the two branches are 
inserted one on each side of its posterior extremity (PI. XXI, 
figs. 132-134). As might have been expected, the retractor 
is narrower in A. burn up i and A. sexangula than in A. 
dimidia with its larger odontophore. 

In Apera gibbonsi and A. parva the retractor is split 
up into a number of powerful muscles, which radiate from 
the hind end of the odontophore, and are attached to a 
considerable area of the floor and the lower part of the sides 
of the body-cavity (PI. XXI, figs. 129-131). In A. 
gibbonsi these muscles originate towards the hind end of 
the body-cavity, though further forward than the origin of 
the single buccal retractor in the other species (PI. IX, 
fig. 27), but in A. parva the muscles have become shorter 
and arise from about the middle of the cavity (fig. 28). 
These muscles are often forked and neighbouring strands 
occasionally anastomose, but there is frequently a slight 
gap dividing those on the right side from those on the left. 
It is easy to see that the same carnivorous habits which have 
led to the enlargement of the radula and odontophore in A . 
gibbonsi and A. parva would produce a corresponding 
development and multiplication of the buccal retractors ; and 
it is also evident that it would be an advantage if the ends of 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 17o 

these muscles were distributed ovei* a considerable area, so 
that the strain produced by the contraction of the muscles 
was not concentrated on a small patch of the body-wall. 
But although it is not difficult to derive the many radially 
disposed retractors of A. gibbon si and A. parva from the 
single retractor of the other foi-ms, I would emphasize the 
fact that the difference is a very great one, and not likely to 
have been completed within a short space of time. So far as 
I am aware the only other form which has developed radial 
buccal retractors at all similar to those of Apera is 
S e 1 e n o c hi a my s .^ 

Since the buccal retractor is inserted in the hind end of 
the odontophore, it might have been expected that it 
would receive nerves from the buccal ganglia, which inner- 
vate the odontophore and the posterior part of the buccal 
mass. Yet this is not the case ; for while the posterior end 
of the reti'actor receives nerves emanating from the right 
pedal ganglion, the front end is innervated by a pair 
of nerves from the cerebral ganglia in all the forms 
with a single retractor. This apparently anomalous inner- 
vation may be explained by assuming that the buccal 
retractor belongs to the same series of muscles as the 
tentacular retractors, with which it is united posteriorly 
in the great majority of snails ; and that it was originally 
inserted, like the tentacular muscles, far forward, and in the 
region innervated by the cerebral ganglia. Later the 
anterior part, which consists of a right and left division, 
would become applied to the sides of the odontophore and 
fused with them, so that the retractor would appear to be 
inserted in the hind end of the odontophore. This theory is 
supported by the facts (1) that the nerves to the buccal 
retractor and to the retractors of the lower tentacles arise 
from the cerebral ganglia by the same roots, and (2) that 
the longitudinal muscle-fibres forming the outer layer of the 
sheath of the odontophore are directly continuous on each 
side with those of the buccal retractor behind (PI. XVIII, 
1 Simroth, H., ' Festschrift Leuckarts,' 1892, p. 55. 



174 HUGH WATSON. 

fig". 122). Tliis view would also explain how it is that in A. 
gibbonsi the nerves which innervate the buccal retractor in 
the other species merely unite with the sheath of the 
odontophore towards the anterior end of that organ. For 
we may suppose that in A. gibbonsi and A. parva the 
odontophore has grown back further than in the other species 
between the two halves of the buccal retractor, which have 
thus become applied to its sides, and consequently the free 
radial retractors of these forms are homologous with only the 
posterior half of the long retractor found in the remaining 
species. 

We have seen that the origin of the buccal retractor in such 
forms as Apera dimidia is to the right of the middle line, 
and that its posterior end is innervated exclusively by branches 
of the right posterior pedal nerve (PI. XV, fig. 75). Now the 
foot is a symmetrical organ, and the anterior part of the 
retractor is certainly symmetrical, as is proved by its 
innervation ; it might, therefore, have been expected that 
the posterior part of the buccal retractor would have been 
symmetrical also, unless it had shifted somewhat to the left 
because of the pressure of the reproductive organs on the 
right side, as in the case of Daudebardia saulcyi {Bgt.) 
and Testacella gestroi [Issel)} Why, then, does the 
buccal reti'actor spring from the right side of the floor of 
the body-cavity in Apera? When an animal possesses a 
feature which is asymmetrical for no apparent reason, the 
explanation of that asymmetry is usually to be found by a 
consideration of the animal's phylogeny. It is highly 
probable that Apera has been evolved from a snail possess- 
ing a depressed or heliciform dextral shell. Now when such 
a shell is in its natural position upon the back of the animal, 
its columella is to the right of the aperture. Consequently 
the columellar muscle, from which the buccal retractor 
springs, will be towards the right side of the animal. Then, 
when the shell degenerates, all the upper part of the colu- 
mellar muscle will disappear, and only the lower part, or 
• Plate, L. H., 'Zool. Jahrb.,' 1891, vol. iv, p. 596. 



I 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 175 

" tail muscle/' in connection with the foot, will remain as the 
direct continuation of the buccal retractor, but it will still be 
situated towards the right side of the animal. In Schizo- 
glossa we have a slug in which the degeneration of the shell 
has only partially taken place, and the chief muscle-fibres 
are still attached to the reduced colmnella ; and it would be 
difficult to draw a diagram illustrating the theory which I 
have just set forth more aptly than does Murdoch's figure of 
the retractor muscles of Schizoglossa novoseelandica 

Several pairs of buccal protractors stretch from the outer 
layer of the walls of the buccal mass to the skin of the 
anterior part of the head. The longest and most important 
of these are inserted on each side just in front of the odonto- 
phore and below the opening of the oesophagus (PI. XXI, 
figs. 129-134). These protractor muscles, together with the 
intrinsic muscles of the buccal mass, serve to protrude the 
odontophore, which seems to be protrusible in all the species 
of Apera. But even in A. gibbonsi the protractors are not 
very thick ; and this is not surprising, for it is evident that 
less powerful muscles will be required to protrude the odonto- 
phore than to retract it after the teeth have become fixed in 
the skin of a worm. 

The Oesophagus and Ckop. — In Apera gibbonsi and A. 
parva the oesophagus is very short and broad, and merges 
imperceptibly into the crop (PI. XXI, figs. 129-131). The 
latter is fusiform in these species, and after increasing in 
width it gradually tapers again towards the openings of the 
hepatic ducts, its broadest part being about twice as far from 
these ducts as from the opening of the oesophagus into the 
buccal mass. 

In the remaining species the oesophagus is long and 
narrow, as will be seen from figs. 132 to 134. At its posterior 
end it enlarges rather abruptly to form the crop, which is 
usually widest near the front end, but remains fairly broad 
almost as far back as the ducts of the liver, and then narrows 
' 'Proc. Mai. Soc," 19U0, vol. iv, pi. xvii, fig. 10. 



176 HUGH WATSON. 

rather suddenly ; but, as might have been expected, the 
crop is much more distended in some specimens than in 
others. 

The walls of the oesophagus and crop are very similar 
in structure (PI. XV, figs. 78, 79, and PL XXII, fig. 135). 
The epithelial lining is normally thrown into longitudinal 
folds, and consists of columnar cells without any cilia. Out- 
side the epithelium of both oesophagus and crop there are 
three layers of muscle-fibres. The fibres of the inner and 
outer layers are longitudinal, Avhile those of the intermediate 
layer run in a circular direction. 

The Stomach, Intestine, and Rectum. — Beyond the crop 
in Apera parva the alimentary canal enlarges to form a 
small stomach, into which the hepatic ducts open (PL XXI, 
fig. 131). In the other species of Apera (with the possible 
exception of A. puree Hi) there is no true stomach, but the 
crop passes directly into the intestine at the openings of the 
hepatic ducts. 

At first the intestine bends abruptly upwards and towards 
the right side of the animal. In A. gibbonsi it then 
describes a curve on the upper surface of the liver like a 
reversed S (PL IX, fig. 27; PL XXI, figs. 129 and 130). 
The posterior curve towards the left side is somewhat shallower 
than that towards the right, and beyond it the alimentary 
canal continues backwards low down on the right side of 
the posterior end of the liver. In A. parva the intestine, 
after curving over to the right side of the liver, passes 
straight backwards, the second curve towards the left side 
being absent (PL IX, fig. 28; PL XXI, fig. 131). 

In Apera burnupi and A. sexangula the first curve is 
much deeper, and forms a loop which extends the whole way 
down the right side of the liver, in which it is partially 
embedded (PL IX, figs. 30, 31, and PL XXI, figs. 133, 134). 
The anterior portion of this loop lies further forward than the 
openings of the hepatic ducts. The posterior curve is shallow 
in these species, and the alimentary canal is continued on 
the right side, as in A. gibbonsi and A. parva. 1'he 



'J'HE OAENIVOKOUS SLUGS OF SOUTH AFRICA. 177 

intestine is considerably narrower in A. burnupi and A. 
sexangula than in the other species. 

In Apera dimidia and A. purcelli the first curve of 
the intestine lies in a more nearly vertical plane, and does not 
extend so far towards the right side (PI. IX, fig. 29, and 
PI. XXI, fig. 132). Then, after curving down on the 
left side, the intestine continues backwards on that side of 
the liver instead of on the right (cf. PI. X, tig. 32, and 
PI. XI, figs. 34, 35). 

Beyond the region of the liver the alimentary canal con- 
tinues backwards as the rectum beneath the mantle-cavity. 
It is at first suriounded by the funnel-shaped prolongation 
of the body-cavity, but further back it curves towards the 
right side of the animal, and bends up the right Avail of the 
mantle-cavity on reaching the level of the respiratory opening. 
As will be seen from PI. X, fig. 33, the anus is immediately 
below the respiratory opening, and the space between them 
is partially separated from the rest of the mantle-cavity by 
an oblique fold of the wall of the cavity, which projects 
forward on the left side of the space. 

The epithelium lining the intestine is composed of very 
irregular columnar cells, the appearance of which is well 
shown in PI. XXII, fig. 137. The inner walls of these cells 
are produced into a delicate fringe of extremely minute cilia. 
Outside the epithelium there is a thin layer of circular muscles, 
and external to this a thin layer of longitudinal mu^scles. The 
structure of the rectum is very similar to that of the 
intestine, excepting that the epithelium is normally thrown 
into deep folds (PI. XXII, fig. 136). At the anus, however, 
the epithelial cells become more regular and oblong in section, 
and their cilia become far longer and less delicate. 

The Salivary Glands and Ducts. — The salivary glands 
are fairly compact, and are situated above and on each side 
of the crop near its junction with the oesophagus. There are 
always two glands, but in Apera gibbonsi and A. dimidia, 
and to some extent in A. parva and A. purcelli, they are 
united above the crop, as in so many of the carnivorous snails 



178 HUGH WATSON. 

(PL IX, figs. 27-29, and PI. XXI, figs. 129-132). In A. 
burnupi and A. sexangula the glands are separate, but 
they are joined to each other by blood-vessels in such a way 
as to suggest that they may possibly have been more closely 
united in the ancestors of these species (PI. IX, figs. 30, 31, 
and PL XXI, figs. 133, 134). The glands are never united 
underneath the crop, as in many of the Ole acini dte, but 
remain widely separate below (PL XV, fig. 78). 

The salivaiy ducts are long and very slender, and discharge 
into the buccal mass on each side of the opening of the 
oesophagus. The structure of the ducts is shown in PL XXII, 
fig. 138. It will be seen that the epithelial cells are of a 
peculiar shape and provided with long cilia. The epithelium 
is immediately surrounded by a layer of circular muscles, and 
outside these there is a layer of longitudinal muscles in which 
is embedded the salivary nerve. Within the gland the duct 
splits up into numerous branches, and in these also the 
epithelium is surrounded by circular muscles. One of these 
branches is shown in fig. 139, which also shows the appear- 
ance of the glandular cells of which the gland itself is almost 
entirely composed. 

The Liver. — The greater part of the posterior half of the 
body-cavity is occupied by the liver or digestive gland (PL 
IX, figs. 27-31). In Apera gibbonsi, A. parva, A. 
burnupi, and A. sexangula the liver consists of two very 
distinct divisions, one anterior, dorsal, and to the right, the 
other posterior, ventral, and to the left (PL XXI, figs. 129- 
131, 133 and 134). Not only do these divisions discharge 
their secretions iuto the alimentary canal by quite separate 
ducts, but they are supplied with blood by different arteries, 
the right division being supplied by one or two branches from 
the anterior aorta, while the left division is supplied by the 
so-called posterior aorta. The right division of the liver is 
divided by the intestine into three main lobes, one lyiug 
within the anterior loop of the intestine, another in front of 
it, and the third behind it, as shown in the figures. Of these 
the anterior lobe, lying above the hind end of the crop, is the 



THE CAKNIVOROUS SLUGS OF SOUTH AFRICA. 179 

most distinct. Each lobe is again subdivided into a number 
of lobules, the arrangement of which varies in different 
individuals. The left division of the liver lies entirely to the 
left of the intestine, and is about equal in size to the right. 
It also is divided and subdivided into lobes and lobules, but 
not so distinctly as is the right division. 

In Apera dimidia the liver is also divided into two main 
divisions, one right and anterior, the other left and posterior ; 
and, as in the other species, these divisions have separate 
ducts. They are not, however, quite so distinct from each 
other as in the species mentioned above, and they both 
receive their blood from a single branch of the anterior aorta, 
the so-called posterior aorta being absent. Moreover, the 
disposition of these divisions with regard to the intestine is 
quite different to that which I have described as character- 
istic of A. gibbonsi, A. parva, A. burnupi, and A. sex- 
angula. The anterior or right division lies above the crop, 
entirely in front of the intestine, while the posterior or left 
division, which is divided into two lobes, occupies a position 
corresponding to that held in the other species by the two 
posterior lobes of the right division of the liver (PL XXI, fig. 
132). This fact suggests that possibly the whole of the liver 
of A. dimidia corresponds to the right division of the liver 
in the other species. If this theory be correct, it would 
explain, not only the anomalous position of the liver with 
regard to the intestine in A. dimidia, but arise its still more 
surprising blood-supply. We would have to suppose, however, 
that the origin of the branch of the right hepatic duct coming 
from the anterior lobe had shifted inwards as far as the intes- 
tine itself j but this might have been caused by the greater 
separation of the lobe due to the lengthening of the right 
division of the liver as it came to occupy also the position of 
the left division. And this separation of the tAvo principal 
branches of the hepatic duct would be merely a case of rever- 
sion, for H. Fischer^ has shown that each division of the liver 
was probably originally subdivided into two or more lobes, 
' • Bull. Sci. France et Belg.,' 1892, vol. xxiv, p. 331. 



180 HUGH \VAT,S0N. 

Avhicli opened separately into the alimentary canal. But it is 
more difficult to explain why half the liver should disappear 
in this way. It is known, however, that the redu.ction of that 
part of the body-cavity which usually occupies the spire of 
the shell sometimes produces a corresponding reduction of 
the left lobe of the liver which it contains.^ It seems pos- 
sible^ therefore, that in A. dimidia the left division of the 
liver, instead of merely pushing forward the right, as in the 
other species of Apera, has disappeared altogether, just as 
the right division has disappeared in some of the Pectini- 
branchia. 

On the other hand, it is possible that the two divisions of 
the liver in A. dimidia are actually homologous with the 
two divisions in the other species. Blood-vessels are always 
very subject to variation, and if by some mutation the so- 
called posterior aorta had disappeared, the arteries supplying 
the right division of the liver would probably extend to the 
left also. Further, the posterior aorta passes to the left of 
the intestine, and may to some extent hold it in position 
(PI. IX, fig. 27). When, therefore, this vessel is not present, 
there is nothing to prevent the intestine shifting to the left 
and coming to occupy the position that it holds in A. 
d i m i d i a . 

Possibly an examination of the liver of Apera pur colli 
may show which of these views is correct. In this species the 
intestine seems to occupy a similar position to that of A. 
dimidia, but unfortunately the liver of the only specimen of 
A. purcelli which I have been able to examine Avas in a 
state of partial disintegration. 

Sections through the liver of Apera dimidia show that 
the food is not confined to the alimentary canal itself, but 
passes up the hepatic ducts into the lobes of the liver, and 
probably digestion takes place chiefly inside this organ in 
Apera as in At opus." Perhaps the disappearance of a 
stomach in these genera may be pai^tly explained by this fact. 

' Pelseneer. P., ' Mem. Acad. Roy. Belg..' 1901. ex vol. liv, p. 55. 
Sinu-otli. H., ' Naturwiss. Woohenschr..' 1901. vol. xvii. p. 121. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 181 



THE VASCULAR SYSTEM. 

The Heart and Pulmonary Veins. — The g-eiius Apera is 
opisthopneuniic, the greater part of the respiratory tissue 
being behind the heart. Unlike Dandebarclia, the auricle 
has moved round with the pulmonary veins so as to lie 
obliquely behind and to the right of the ventricle (PI. IX, 
figs. 27-31), but it has not rotated so far as in Testacella. 
The pulmonary veins form a complicated network, variable in 
its structure ; but it is generally possible to distinguish three 
main vessels converging towards the auricle, namely, the right 
anterior pulmonary vein, from the corner of the mantle-cavity 
to the right of the pericardium ; the right posterior pulmonary 
vein, from the region between the pericardium and the anus ; 
and the left posterior pulmonary vein, from the part of the 
mantle-cavity behind the kidney. Owing to the absence of 
the respiratory tissue from the left anterior corner of the 
mantle-cavity there is no left anterior pulmonar}" vein. 

The walls of the auricle are very thin (PI. XI, fig. 85), Ijut 
those of the ventricle are extremely thick and muscular 
(fig. 34). The cardiac muscle-fibres are arranged in bundles 
passing in various directions, and leaving spaces between them 
which communicate freely with the central cavity of the ven- 
tricle (PL XXII, fig. 140). The muscles are thus well supplied 
with blood without the intervention of any cardiac arteries. 

The Arterial System. — The distribution of the principal 
arteries is shown in PI. IX, figs. 27-29 and 31. The single 
aorta leaves the posterior end of the ventricle, passes through 
the diaphragm, and then bends abruptly to the right and 
curves downwards, so that it comes to lie on the right side of 
the liver. In Apera gibbonsi, A. parva, A. burnupi, 
and A. sexangula the so-called posterior aorta branches off 
to the left within 3 or 4 mm. of the diaphragm. This vessel 
passes forwards on the right side of the left division of the 
liver, keeping to the left side of the intestine. It divides into 
two or three branches supplying the left division of the 
liver, but the most anterior branch first passes through the 



182 HUGH WATSON. 

hei-maphrodite gland^ to which it also supplies blood. In A . 
burnupi, but not in the other species, this branch also gives 
off a small artery to the hind end of the right division of the 
liver. In A. parva the posterior aorta divides immediately 
into three branches at the point where it separates from the 
anterior aorta, but in the other species the division of the 
vessel takes place further forward. No posterior aorta is 
present in A. dimidia, and there is probably none in A. 
purcelli either. 

I would suggest that perhaps this so-called posterior aorta 
may be merely the left hepatic artery which has become dis- 
placed, and that the Gastropoda (with the possible excep- 
tions of Haliotis and Fissurella^) resemble the Amphi- 
neura and the more primitive members of the Pelecypoda 
in having no true posterior aorta. The posterior position of 
this artery would be accounted for by the posterior position 
which the left division of the liver has come to occupy owing 
to the torsion of the visceral hump. Its comparatively large 
size in most genera might be due to the fact that the left 
division of the liver is generally much larger than the right, 
since it extends up the spire of the shell. And as the herma- 
phrodite gland is usually more or less embedded in the left 
division of the liver, it would be likely to be supplied with 
blood by the same artery. This view will be made clear by 
the accompanying diagrams a to d. Diagram a represents a 
primitive symmetrical arrangement, and c the condition after 
the torsion has taken place; B shows a hypothetical stage 
between a and c, while d represents the most usual arrange- 
ment of the arteries and liver in Apera. 

The anterior aorta continues forward on the right side of 
the liver, crossing over the intestine in A. gibbonsi, A. 
T)iiriiupi, and A. sexangula. In A. gibbonsi, A. 
dimidia, and probably in A. purcelli, it gives off a single 
artery to the liver on reaching the level of the anterior loop 

' See Wegmann, H., 'Arch. Zool. Exper.' (2nd ser.), 1884, vol. ii, pp. 
352, 353, pi. xviii, figs. 1-4 ; and Boutan, L.. ibid., 1885, ex vol. iii bis 
suppl., pp. 34, 130, pi. xxxiii, figs. 5, 6. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 183 



Text-fig. 5. 



L.L.. 




„..R.L. 




R.L. 



LX... 




LL. 




D. 



Diagrams illustratinfj the hypothetical evolution of the 

" posterior aorta." 

(Diagram D shows the arrangement in Ape r a gibbon si Inpata.) 



L.L. = Left division of liver. R. = Rectum. 
R.L. = Right division of liver. V. = Ventricle. 



184 HUGH WATSON. 

of the intestine. This hepatic ai'tery soon divides into two 
branches. In A. gibbonsi the anterior branch chiefly 
supplies the anterior lobe of the right division of the liver^ 
while the remainder of the right division is supplied by the 
posterior branch. In A. dimidia the anterior branch 
supplies the a,nterior division of the liver, while the posterior 
branch supplies not only the posterior division, but also the 
hermaphrodite gland which it traverses. In A p e ra b u r n u p i 
and A. parva the blood-supply of the right division of the 
liver is similar to that in A. gibbonsi, excepting that 
the two branches arise separately from the anterior aorta, 
though very close together. In A. sexangula these arteries 
also arise separately from the anterior aorta, and their origins 
are much further apart. 

The anterior aorta passes through the loop formed by the 
intestine in Apera burnupi, A. sexangula, A. parva, 
and in A. gibbonsi gracilis and A. g. lupata (PI. IX, 
figs. 28, 31, and text-fig. 5, d) ; but in A. dimidia, A. pur- 
celli, A. gibbonsi s. s., and A. g. rubella, the aorta 
passes straight forward on the right side of the loop (figs. 27, 
29). It is very remarkable to find this important difference 
separating forms which seem to be so closely related that I 
have not ventured to regard them as distinct species. For 
the difference amounts to this : that in the first group the 
aorta, on its way from the dorsally situated heart to the 
ventral ganglia, passes on the left side of the alimentary 
canal, while in the second group the aorta passes on the right. ^ 

The aorta then runs forward between the digestive and re- 
productive organs, being often closely applied to the left side 
of the albumen gland. It gives off to the left three or four 
ai'teries to the crop and salivary glands, and to the i-ight at least 
one artery to the albumen gland and another to the common 
duct, a branch of the latter going forward to the receptaculum 
seminis. The aorta then becomes more ventrally situated and 

^ A similar variation in the course of the aorta lias been found by 
Koliler in the ^^enns Siphon aria ('Zool. Jahrb.,' 1893, vol. vii; compare 
tig. B, p. 27, with figs. A and c, p. 32). 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 185 

runs forward towards the ventral group of ganglia. Just 
before reaching these a pair of arteries is given oif to the 
tentacular retractors (PI. XIII, figs. 52 and 54-57). In most 
of the species these arteries unite with the common tenta- 
cular retractors just behind the points where the latter divide ; 
but in Apera dimidia they unite with the upper tentacular 
retractors, and give oif small branches to the retractors of 
the lower tentacles, which join the nerves from the sides of 
the "nerve-collar" (fig. 54). Finally the aorta fuses with 
the ventral group of ganglia. 

From this point radiate numerous short vessels to the 
various parts of the central nervous system, etc., as well as 
two longer arteries, namely, the buccal and pedal arteries. 

The buccal artery passes upwards and divides into two 
branches, one running forwards and the other backwards. 
In Apera dimidia the posterior branch is about twice as long 
as the other (PL XVII, fig. 103) ; it runs back to the posterior 
extremity of the odontophore, which it enters, and then bends 
round the hind end of the odontophoral support (PI. XVIII, 
fig. 122). The anterior branch runs forward and divides into 
a pair of arteries which apply themselves to the sides of the 
buccal mass, dividing again as they do so into anterior and 
posterior vessels. In A. burnupi and A. sexangula the 
buccal artery bi-anches in a similar manner, but owing to the 
small size of the odontophore, the posterior branch is no 
longer than the anterior branch (PI. XVII, fig. 110). In A. 
gibbon si and A. parva, on the other hand, the posterior 
branch is very long, and in the former species the right and 
left anterior arteries do not branch from a sing-le median 
vessel, but arise separately (PI. XIII, fig. 52). 

In Testacella the artery supplying the odontophore with 
blood behaves differently, for, instead of running straight to 
the hind end of the organ, it divides into lateral vessels which 
pass through the sides of the sheath as shown in de Lacaze- 
Duthier's admirable figui-es.^ On the other hand, in Nata- 
lina quekettiana {M. & P.), this artery behaves in the 
' 'Arch. Zool. Expei-.' (2nd ser.), 1887, vol. v. pis. xxxv. xxxvi. 



186 HUGH WATSON. 

same way as it does in Apera. In Euglandina we 
have an intermediate arrangement, as will be seen from 
Strebel's figures ^ ; for the structure that he described and 
figured as a median odontophoral nerve from the ventral 
group of ganglia, is, in reality, the buccal artery, as I have 
found from an examination of E. venezuelensis {Preston). 

The pedal artery runs backwards above the pedal gland, to 
which it gives off branches on each side. In Apera gib- 
bonsi, A. dimidia, and A. sexangula, I have found a very 
slender vessel uniting the anterior aorta with the pedal artery 
about half way along the pedal gland (PI. XIII, figs. 52, 54, 
and 57). After uniting w4th the terminal vesicle of the pedal 
gland, the pedal artery continues backwards and downwards, 
and becomes more or less embedded in the muscular con- 
nective tissue of the foot. 

The venous system is largely lacunar, apart from the 
pulmonary veins already described. Indeed, the only well- 
defined vessel containing venous blood is the body-cavity itself. 
I have already dealt with the dermal veins and their curious 
sphincters, when treating of the structure of the skin." 

THE EXCRETORY SYSTEM. 
The pericardium communicates with the mantle-cavity by 
means of a single renal coelomoduct, divided into an ascend- 
ing glandular portion, which forms the kidney, and a 
descending non-glandular portion, which forms the ureter. 
It will be seen from PI. IX, figs. 27-31, PI. XI, figs. 34, 35, and 
PI. XII, fig. 36, that the kidney is applied to the left side of 
the pericardium, and also extends over its posterior part 
towards the right side, thus broadly connecting the peri- 
cardium with the Avail of the mantle-cavity, with which the 
kidney fuses a little further back. Ventrally the kidney is 
prolonged backwards in the floor of the mantle-cavity on 

' 'Beitrag z. Kenntn. d. Fauna Mexikan. L.- u. Siisswasser-Conchylien,' 
1878, vol. iii. pi. xvi, fig. 5, pi. xviii, figs. 1, 6. 10; pi. xix, fig. 1, pi. xx, 
fig.l. 

* See pp. 126. 127. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 187 

the left side of the rectum and even beyond the anus (PI. X 
fig. 33). The walls of the kidney are very much folded 
internally (see especially PI. XI, fig. 35), and ai-e lined by an 
epithelium consisting of glandular excretory cells with 
conspicuous round nuclei (PI. XXII, fig. 141). 

The reno-pericardial duct is very narrow, but projects 
some distance into the kidney. Its position is shown in 
PI. XI, fig. 35. The epithelial cells lining the duct are 
provided with veiy long cilia. 

The kidney communicates with the ureter by a minute 
pore situated in the centre of a slightly raised papilla at the 
right end of its dorsal extension (PI. XII^ fig. 36). The 
ureter, as in other sigmurethrous Stylommatophora, is 
doubled back upon the kidney, and curves round its inner 
posterior side. Ventrally it is continued backwards beside 
the ventral prolongation of the kidney, and extends to the 
extreme end of the mantle-cavity, where it bends abruptly 
upwards and opens. The ureter is lined by a cubical 
epithelium, the appearance of which is shown in PI. XXII, 
fig. 141. This epithelium becomes thinner over the papilla in 
which the opening of the kidney is situated ; elsewhere it is 
remarkably like the epithelium which lines the lower surface 
of the shell-sac. 

As in Testacella and the Rhytidid£e, there is no 
secondary ureter; but a rather irregular groove runs for- 
wards from the opening of the ureter along the roof of the 
mantle-cavity. This groove reaches nearly to the point 
where the ureter begins, and then bends round and runs 
obliquely backwards and to the right, to end in the cleft in 
the inner lip of the respiratory opening. The epithelium 
lining this groove is similar to that lining the ureter itself, 
and the groove is probably to be regarded as an incipient 
secondary ureter, such as is found in many other forms. 
The somewhat peculiar course of the groove might be 
explained on the assumption that the respiratory orifice 
was originally situated further forwards. 

VOL. 3, PAKT 2. 14 



188 HUGH WATSON. 

THE REPEODUCTIVE ORGANS. 

The genital or reproductive organs of Apera are of a 
simple generalised type^ at least so far as their external 
structure is concerned (PL XXIII, figs. 143, 144, 146-149). 
As up to the present only a very few fully mature specimens 
have been collected, I have thought it advisable to preserve 
the reproductive organs of these almost intact, and not to cut 
them up in order to examine their histology. The following 
account, therefore, deals chiefly with the external morphology 
of the genital system. The reproductive oi-gans of Apera 
purcelli are unknown, the only dissected specimen being 
immature. 

The Hermaphrodite and Female Organs. — The herma- 
phrodite gland or ovotestis is partially embedded in the 
right side of the posterior division of the liver (see especially 
PI. IX, fig. 31). It is a racemose gland composed of a 
cluster of oval or pear-shaped follicles. These follicles are 
smallest in Apera burnupi and A. sexangula (PL XXIII, 
figs. 148, 149). In the other species they are larger and 
tend to be fewer in number, until in A. parva the entire 
hermaphrodite gland consists of only about four follicles 
(fig. 146). The very slender ducts from the different follicles 
unite to form the hermaphrodite duct, which is long", and 
usually more or less convoluted and slightly swollen towards 
the middle of its length, especially in A. dimidia. But the 
convolution is not so pronounced as in many other genera. 
In front the hermaphrodite duct becomes embedded in the 
albumen gland. At the extreme anterior end it is swollen 
so as to form a minute spherical vesicle, which is doubtless to 
be regarded as a rudimentary vesicula seminalis. 

The albumen gland varies enormously in size according 
to the exact state of maturity of the individual (cf. PL IX, 
figs. 27, 31). In some specimens it reaches a very large size 
indeed (PL XXIII, fig. 144). The common duct, or ovi- 
spermatoduct, is very long and much convohited and twisted 
in Apera burnupi and A. sexangula (PL XXIII, figs. 



THE CARNIVOROUS SLUGS OF SOUTH AFREGA. 189 

148, 149); in A. gibbonsi, A. parva, and A. diniidia 
it is shorter and generally broader (figs. 143, 144, 146, 147). 
In some specimens the female or oviducal portion is greatly 
swollen, as in the example ofA.dimidiaof which the repro- 
ductive organs are shown in fig. 147. The free oviduct is 
rather long and narrow, and is directly continuous with the 
vagina, which is also rather narrow but usually somewhat 
shorter. The vestibule or genital atrium is extremely short. 

The Receptacdlum Seminis and its Duct. — The sperma- 
theca or receptaculum seminis is an oval vesicle, situated, in 
A. parva, beside the anterior end of the free oviduct, the 
receptacular duct in this species being exti-emely short 
(fig. 146). In the other forms the receptaculum seminis 
lies against the anterior end of the common duct, and the 
length of the receptacular duct is therefore about equal to 
that of the free oviduct, since the anterior opening of the 
receptacular duct marks the junction of the free oviduct with 
the vagina. In Apera gibbonsi the receptacular duct is 
a simple narrow tube (figs. 144, 145). In A. sexangula it 
becomes swollen towards its union with the vagina, so that 
the anterior part of the duct is fusiform (fig. 149). In A. 
burnupi the anterior part of the duct is still more swollen 
(fig. 148). Lastly, in A. dimidia the swollen end of the 
duct has become attached to the adjacent part of the body- 
wall by a number of muscles on each side (fig. 147). And 
as we pass up this series the receptaculum itself becomes 
progressively smaller, and it seems as though the receptacular 
duct were taking its place. If this tendency were to proceed 
still further we should have a vesicle which would be indis- 
tinguishable from a sessile receptaculum seminis, although in 
reality it was the swollen anterior end of the receptacular 
duct, the rest of the organ having disappeared. Possibly 
this may have happened in some of the genera which are 
usually described as having a sessile receptaculum. 

The Male Organs.— The vas deferens arises from the 
anterior end of the common duct, and passes forwards close to 
the free oviduct and vagina as far as the vestibule. It then 



190 HUGH WATSON. 

bends round and runs up the side of the penis. In Apera 
gibbon si the vas deferens is rather thick excepting near the 
vestibule, where it has the more usual form of a very slender 
duct. In this species, and also in A. dimidia, before the 
vas deferens unites with the posterior end of the penis, it is 
slightly swollen for a distance equal to about one-third of the 
length of that organ, thus forming a rudimentary epiphallus. 
In A. sexangula this swollen part is very much shorter, 
and bears at its anterior end a minute flagellum, which, like 
the vas deferens itself, is closely applied to the side of the 
penis (PI. XXIII, fig. 149). 

The penis passes between the retractor muscles of the 
right upper and lower tentacles (PL XIII, figs. 52-57). 
In this respect Apera resembles Testacella, Trigono- 
chlamys, Poiretia, Streptostyla, Strebelia, Rhytida, 
and Paryphanta, and differs from Daudebardia, Plu- 
tonia, Selenochlamj^s, Natalina, Delos, andthe Strep- 
taxi dfe, in which the reproductive organs are on the right of 
both retractors, as was first shown by Pfeffer, Simroth, and 
Murdoch. In Apera dimidia the penis is fairly short, but 
a little swollen posteriorly (PI. XXIII, fig. 147). In A. 
parva it is about 4 mm. in length, and somewhat fusiform 
(fig. 146). In typical specimens of A. gibbonsi from Zulu- 
land, the penis is also short (fig. 143), but in the other races 
of that species it is very long and twisted in a peculiar 
manner (fig. 144). In A. sexangula the penis is long and 
very narrow, excepting at the posterior end, where it is broad 
and somewhat flattened (fig. 149). In A. burn up i the 
anterior part of the penis is also narrow, but it is much 
shorter than in A. sexangula, and more or less twisted. 
The posterior broader portion, on the other hand, is longei*, 
and appears to be double, OAving to the presence of a deep 
longitudinal groove on one side ; and the vas deferens dis- 
appears into this groove about 5 mm. in front of the end of 
the penis (fig. 148). Possibly this peculiar arrangement may 
be due to the broad sides of the penis having arched over the 
end of the vas deferens and joined each other above it ; but 



THE CABNIVOEOUS SLUGS OF SOUTH AFRICA. 191 

this hypothesis must remain very doubtful until a sufficient 
number of specimens of this rare species has been collected in 
order to justify the dissection of the penis. In a full-grown 
example of A. gibbonsi rubella from Durban, the penis 
was found to be quite vestigial. 

As so few full-grown specimens of Apera have been 
collected, I have only felt justified in examining the internal 
structure of the male organs in two forms, namely, A. 
dimidia and A. gibbonsi rubella. No penis-papilla is 
present in either of these species, but in both forms the inner 
surface of the muscular wall of the penis is completely covered 
by numerous small papillae, as shown in PI. XXIII, fig. 145. 
These papilla? are diamond-shaped in transverse section, and 
each ends in a short point. The structure of the papillae is 
very similar in the two species, and is shown in PI. XXII, 
fig. 142, which represents a photomicrograph of the inner half 
of a single papilla from the penis of A . gibbonsi rubella. 
In this form the papillse become considerably smaller around 
the opening of the epiphallus, and within the opening they 
become much narrower in proportion to their length, 
changing, in fact, from papillee into filaments. These fila- 
ments occur throughout the entire length of the epiphallus. 
They do not, however, arise from all parts of the inner 
surface of the wall of the epipHallus, but are arranged in 
longitudinal rows, of which there are about fourteen in the 
specimen that I examined. Between the rows the wall of the 
epiphallus is extremely thin. In Apera dimidia the epi- 
phallus has a different structure, being lined by numerous 
minute papilla?, closely arranged, and not unlike the papillae 
lining the penis, only much smaller. It is somewhat remark- 
able that while the inside of the penis is so very similar in 
the two species, the inside of the epiphallus should present 
such a marked contrast. 

The retractor of the penis is attached to its posterior end, 
and arises from the roof of the body-cavity near the mid-dorsal 
line. In Apera burnupi, A. sexangula, A. purcelli, 
and A. parva, it is very long and narrow, arising in A. 



192 HUGH WATSON. 

sexangula and A. parva less than 4 mm. in front of the 
diaphragm, and only a very short distance further forwards 
in the other two species (PI. IX, figs. 28, 30, 31). On the 
other hand, in A. gibbonsi and A. dimidia it is broader 
and very much shorter, the exact position of its origin 
depending on the length of the penis (figs. 27 and 29). 
Probably the retractor originally arose in the neighbourhood 
of the diaphragm ; but owing to the posterior position which 
the diaphragm has come to occupy, the origin of the retractor 
has moved forward. If this view be correct, the condition 
found in A. sexangula and A. parva would be the most 
primitive. 

The Spermatozoa. — PI. XXIII, fig. 151, shows the anterior 
end of a spermatozoon of Ap era dimidia magnified 1200 
diameters. It will be seen that the head is curved into the 
shape of a hook, and is about "0035 mm. in length. The 
'^ middle-piece," or proximal portion of the tail, is surrounded 
by a spiral filament or flange, very loosely coiled, the Avhorls 
being about "004.5 mm. apart. The posterior portion of the 
tail is extremely long. In Apera sexangula and A. 
gibbonsi rubella the head of the spermatozoon is straighter 
and broader, though pointed in front. Fig. 150 shows the 
anterior end of a spermatozoon of the latter variety. It will 
be seen that in this form the whorls of the spiral filament are 
a little nearer together than in A. dimidia, being separated 
by a distance equal to the length of the head, namely, 
•0037 mm. 

The spermatozoa of Testacella are very diiferent from 
those of Apera, as may be seen from PI. XXIV, fig, 160. In 
Testacella the head is globular, and there are two spiral 
filaments surrounding the proximal portion of the tail. 
Moreover the filaments are much more closely coiled than in 
Apera, adjacent whorls being only about "0009 mm. apart 
in T. maugei. In Rhytida capillacea {F<'r.) I find that 
there are also two filaments, but they are not so closely coiled, 
the whorls being separated by about "00165 mm. ; and in this 
species, as well as in Natalina quekettiana (il/. & P.), the 



THE CARNIVOEOUS SLUGS OF SOUTH AFRICA. 193 

liead is not unlike that of Apera sexangula and A. 
gibbon si rubella. On the other hand, in Streptaxis 
funki Pfr. the head is even more strongly curved than in 
Apera dimidia; and, as in the Testacellidas and E.hy- 
tididie, I have found two spiral filaments encircling the 
proximal portion of the tail, the distance separating adjacent 
whorls in this species being about •00115 mm. 
The embryology of Apera is unknown. 

DISTRIBUTION. 

The genus Apera is only known to inhabit the maritime 
provinces of South Africa. One species — A. pur colli — 
occurs on the slopes of Table Mountain close to Cape 'J'own; the 
other forms have only been found in Natal and in the eastern 
districts of the Cape Province. A. parva is at present 
only known to occur near Grahamstown ; A. sexangula 
extends from that neighbourhood as far east as Pieter- 
maritzburg ; the various races of A. gibbon si are known 
to inhabit an area stretching from Pot)doland to Zululand ; 
while A. burn up i and A. dimidia have only been found 
in Natal. 

The animals are usually found in the bush, either amongst 
dead leaves or under stones or logs ; but none of the species 
are at all common. Apera sexangula and A., gibbonsi 
seem to be the least rare. Of the others only six examples of 
A. dimidia, four of A. burnupi, two of A. purcelli, and 
one of A. parva are known to have been collected. It is to 
be hoped that South African naturalists will gather further 
specimens of this interesting genus, which not improbably 
contains more species than are at present known. 

THE DIFFERENTIAL CHARACTERS OF THE SPECIES 
AND SUBSPECIES OF APERA. 

Apera gibbonsi {Binney) . 

Chlamy depliorus gibbonsi Binn., Bull. Mus. Comj). Zool. Har- 
vard. 1879, vol. V, p. 331, pi. ii, figs. (/, h\ Binney, Ann. N. Y. Acad. 



194 HUGH WATSON. 

Sci., 1884, vol. iii, p. 81, pi. xvii, fig. a ; Tryon, Man. of Concli. 

(2nd ser.), 1885, vol. i, pp. 17, 251, pi. ii, fig. 95 ; Cockerell, Ann. 

Mag. Nat. Hist. (Oth ser.), 1890, vol. vi, p. 390. 
Chlamydophorus gibbon si Binn.; Tryon, Struct, and Syst. 

Conch., 1884, vol. iii, p. 13, pi. ci, fig. 47. 
Apera gibbonsi (Binn.); Heynemann, Jalirb. d. Deutscli. Mai. Ges., 

1885, vol. xii, p. 17, pi. ii, figs. 5-7; Collinge, Ann. S. Afr. Mus. , 

1900, vol. ii. p. 4; ColUnge, Ann. Natal Mus., 1910, vol. ii, p. 165; 

Connolly, Ann. S. Afr. Mus., 1912, vol. xi, p. 63. 



External Characters (PI. YII, figs. 1, 2 ; PI. VIII, figs. 14, 
15). — Animal slender, tapering to an acute angle at the hind 
end. Back rounded, without keels. Outer lip of respiratory 
opening almost concealing the inner lip. Skin coarsely 
reticulated. Dorsal grooves usually about 1 mm. apart, 
separated by a single row of rugse, but united for about 4 or 
5 mm. in front of the respiratory opening. Lateral grooves 
irregular, rather near the dorsal grooves, and seldom extend- 
ing as far as the head. Radial grooves conspicuous, some 
being forked. Oblique sub-lateral grooves also conspicuous, 
and rather numerous ; these and the radial grooves termi- 
nating in the single peripodial groove. 

In the typical form the body is mottled with brown, the 
small patches of colour being chiefly concentrated dorsally, 
but leaving a paler band, very sparsely mottled, along the 
centre of the back from the head to the respiratory opening. 
Laterally the brown pigment becomes confined to the sides of 
the grooves in the skin, and dies out completely some distance 
above the peripodial groove. The ground-colour is pale 
yellow in specimens preserved in spirit, but living examples 
are tinged with dull orange, the colour being deepest on the 
back. 

The type was described by Mr. Gibbons as being rather 
more than 3 in. long when fully extended. The following are 
the dimensions (in alcohol) of two specimens that I have 
examined, but the reproductive organs of the smaller one 
were not quite fully developed. 



62 mm. . 


66' 5 mm. 


9-5 „ . 


12-5 „ 


V „ 


9 „ 


5 „ 


5 „ 


9-5 „ 


. 12-5 „ 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 195 

Length . . . . 

Breadth in middle . 

Breadth at respiratory opening 

Breadth of foot-sole . 

Greatest height ... 

Distance from respiratory open- 
ing to hind end . . . 13 „ . ]3'5 ,, 

Internal Characters. — Skin very thick. Shell 4 x 2*5 
mm., usually thick, depressed, with the apex in front of the 
posterior margin ; sinus deep. 

Pedal gland. — Slender; glandular tissue gradually dying- 
out some distance in front of the terminal vesicle; duct closely 
convoluted posteriorly ; fold in terminal vesicle divided into 
narrow lamellge. 

Nervous system. — Cerebral ganglia closely united; 
buccal ganglia large, close together, situated posteriorly, and 
joined to the cerebral ganglia by extremely long connectives ; 
cerebro-pedal and cerebro-pleural connectives also long; 
nerves corresponding to those of buccal retractors in other 
species uniting with anterior end of sheath of odontophore, 
and joined to cerebro-buccal connectives for most of their 
length ; nerves to pedal gland very slender. 

Digestive System (PI. XVII, figs. 91, 92; PI. XX,fig.l25; 
text-fig. 3, A, p. 159 ; PI. XXI, fig. 129).— Radula exceedingly 
large, sometimes attaining a length of 21 mm.; central tooth 
typically present; others not differentiated into laterals and 
marginals, very large excepting near the edges and the 
centre ; cusps in typical form simple, slightly curved ; bases 
very narrow ; apophyses typically not prominent. Formula of 
type : (27 + 1 + 27) X 52, of a Zululand example : (37 -f 1 
-h 38) X 60. Odontophore also extremely large ; posterior 
end joined to the fioor and sides of the body-cavity towards 
its hind end by numerous radial buccal retractors ; odonto- 
phoral support rounded in front. CEsophagus short and 
broad, merging into a broadly fusiform crop. Anterior loop 
of wide intestine not much deeper than posterior loop; 
intestine continued posteriorly on right side of liver. Salivary 



196 HUGH WATSON. 

glands united above crop. Liver of two appi-oximately equal 
divisions, the intestine only traversino- the right. 

Vascular System. — Pulmonary veins greatly branched. 
" Posterior aorta " supplying left division of liver and her- 
maphrodite gland ; two arteries of right division of liver 
arising from anterior aorta as a single vessel ; anterior aorta 
passing to right of intestine in typical form; anterior branches 
of buccal artery separate throughout. 

Reproductive System (PI. XXIII, tig. 143). — Her- 
maphrodite gland with rather large follicles; common duct 
rather short, curving twice to the right and twice to the left; 
vagina narrow ; receptaculuin seminis rather large; recepta- 
cular duct of moderate length and slender throughout. A^as 
deferens rather thick excepting near the vestibule, somewhat 
swollen near the posterior half of the penis to form an 
epiphallus ; penis short and curved posteriorly in specimens 
from Zululand ; penial retractor short. 

Habitat. — Typical form : Umgeni valley, Natal (under 
a stone in a wood ; Gibbons) ; Pietermaritzburg (?, Burnup, 
Ponsonby) ; Hlabisa, Zululand (Burnup). 

Type. — The type is in the Academy of Natural Sciences, 
Philadelphia. Other specimens will be found in the Natal 
Museum, Pietermaritzburg. 

Affinities, etc. — This species is the type of the genus. It 
differs from all the other species in its radula, buccal retrac- 
tors, receptacular duct, etc. At the same time it shows a 
remarkable diversity in some of its organs, specimens found 
in different localities, and even, in one case, in the same 
locality, constituting races which differ markedly from each 
other in their radulte and in a few other important characters. 
Indeed, it is not at all improbable that these forms will prove 
to be distinct species. For the present, however, until further 
specimens liave been collected, I have thought it better to 
treat them provisionally as subspecies of A per a gibbon si; 
for although their differences are undoubtedly of considerable 
importance, in most of their anatomical features they appear 
to be identical, and it is impossible to distinguish them merely 



THE CAIiNIVOROUS SLUGS OF SOUTH AFRICA. 197 

by their external charactei's, excepting in the case of one form, 
of which only a single specimen has been collected. 

Apera gibbonsi rubella n. suhsp. 

External Chaeactbks (PI. VII, figs. 3, 4, 5 ; PI. VIII, figs. 
16, 17). — Animal broader than the typical form of Apei-a 
gibbonsi, tapering more abruptly at the hind end. Back 
usually slightly flattened just behind the respiratory opening. 
Reticulation of skin very similar to that of the typical form 
of A. gibbonsi, but not so coarse, the grooves being more 
numerous and less deep. Dorsal grooves usually less than 
1 mm. apart, and united for about 4 mm. in front of the 
respiratory opening. 

Body thickly mottled with dark grey (which sometimes 
has a bluish tinge in specimens preserved in spirit). The 
small patches of colour are usually ill-defined, and tend ta 
coalesce dorsally, where they are most abundant; but a 
narrow paler band, more sparsely mottled, is generally left 
along the centre of the back from the head to the respiratory 
opening. Laterally the dark pigment is chiefly concentrated 
on the sides of the grooves in the skin, but it dies out completely 
a short distance above the peripodial groove. The ground- 
colour of specimens preserved in spirit is whitish, tinged with 
dull red dorsally, but living examples ai-e deeply coloured with 
an unstable orange-red pigment, so that the animal is usually 
reddish-brown above, merging into deep reddish orange on 
the sides, while the foot- sole is of a paler orange colour. 

The following are the approximate dimensions (in alcohol) 
of the smallest specimen with fully developed reproductive 
organs, and of the largest specimen, which I have examined. 

Length ..... 

Breadth in middle . 

Breadth at respiratory opening- 
Breadth of foot-sole 

Greatest height 

Distance from respiratory open- 
ing- to hind end . .10 ,, .13 



49'5 mm. 


. 67 


10 „ 


. 13-5 


9 „ 


. 11 


5-5 „ 


6 


9-5 „ 


. 13-25 



198 HUGH WATSON. 

Internal Characters. — Glandular tissue of pedal gland 
reaching" nearly as far back as the terminal vesicle. Radula 
(PI. XX, fig. 126; text-fig. 3, b, p. 159) without central tooth; 
cusps of teeth very strongly curved, slightly broadened towards 
the points ; apophyses very prominent ; formula of a speci- 
men from Equeefa : (28 + + 28) X 51; of a young example 
from Equeefa : (24 -f + 24) x 44 ; of a specimen from 
Durban: (34 + + 35) x 48. Liver sometimes shorter and 
broader than in typical form (PL XXI, fig. 130). Penis 
long and twisted (PI. XXIII, fig. 144). The anterior aorta 
passes on the right of the intestine (PI. IX, fig. 27), and all 
the other anatomical characters seem to be identical with those 
of the typical form (PI. X, figs. 32, 33 ; PI. XIII, figs. 41, 
42, 52; PL XIV, figs. 63-65, 67; PL XV, figs 70, 77-79; 
PL XVII, figs. 93-98; PL XXII, figs. 138, 139, 142; PL 
XXIII, figs. 145, 150). 

Habitat. — Equeefa, Natal (Burnup) ; Durban (under dead 
leaves in the bush which fringes the shore; Burnup). 

Type. — The type of subspecies is in the Natal Museum, 
Pietermaritzburg. Another specimen will be found in the 
British Museum. 

Affinities, etc. — The chief characters Avliich distinguish this 
form from Apera gibbonsi s.s. are its broader shape and 
darker colour, its radula, and its penis (although it is not known 
whether the penis of the typical form is always as short as 
it is in the Zululand specimens). It is possible that this may 
be a geographical variety of A. gibbonsi, as the two forms are 
not known to occur in the same district, but the radulte are 
so different as to suggest that this may prove to be a distinct 
species. The single specimen from Durban differs from those 
found at Equeefa in being without a paler dorsal band, in 
having more teeth in each row in its radula, and in the penis 
being abnormally reduced to a minute vestige. The resem- 
blance of this specimen to certain dead leaves amongst those 
under which it was found has already been referred to (see 
p. 122) . Mr. Burnup informs me that this subspecies contracts 
after the usual manner Avhen about to rest, instead of merely 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 199 

bending its body at various angles like the typical form of 
A. gibbon si. 



Apera gibbon si gracilis n. suhsp. 

External Characters (PI. VII, fig. 6). — Animal even more 
slender than the typical form of Apera gibbonsi, tapering 
to a very acute angle at the hind end. Outer lip of the 
respiratory opening not concealing so much of the inner lip 
as in the other forms of A. gibbon si. Eeticulation of skin 
coarse, as in the typical form ; dorsal grooves united for 6 
mm. in front of the respiratory opening. 

Body rather sparsely mottled with brown, as in A. gib- 
bon si, s.s. ; the patches of colour, however, are not concen- 
trated on each side of the mid-dorsal area, but extend across 
it, so that there is no median paler band. 

The following are the measurements of the only specimen 
that has been found : 

Length . . 44 ram. Greatest height . 6 mm. 

Breadth in middle 6 ,, Distance of i-esp. 

Breadth at resp. opening from hind 

opening . . 4*5 „ end . . H „ 

Breadth of foot-sole 3 „ 

Internal Characters. — Shell (PI. XIII, fig. 43) rather 
longer and narrower at the posterior end than in the other 
forms of A. gibbonsi. Radula (PI. XIX, fig. 123; text- 
fig. 3, c, p. 1 59) with fewer teeth than that of any of the other 
forms; central tooth absent; first on each side extremely small ; 
outer teeth larger than in the preceding forms ; cusps only 
slightly curved^ not broadened or barbed; apophyses very 
prominent ; formula : (19 -|- + 20) x 34. Anterior 
aorta passing through loop of intestine; branches of pul- 
monary veins not very numerous. Penis rather long and 
twisted. Other characters agreeing with those of A. gib- 
bonsi s.s. 

Habitat. — Equeefa, Natal (Burnup). 



200 HUGH WATSON. 

Type. — The type of subspecies is in the Natal Museum, 
Pietermaritzburg-. 

Affinities, etc. — The single specimen of this foi'm differs 
greatly from the last subspecies, although it was found in the 
same localit3\ In some respects it is intermediate between 
Apera gibbonsi s.s. and the next subspecies, but it seems 
impossible to associate it with either of these forms, and it 
must therefore stand alone until further examples have been 
collected. It will be noticed that owing to the length of the 
posterior extremity the respiratory opening is proportionately 
further forward than in the other forms. 



Apera gibbonsi lupata n. suhsji. 

External Characters (PL VIII, figs. 18, 19). — Animal 
intermediate between A. gibbonsi s.s. and A. g. rubella 
both in shape and in the reticulation of the skin, being 
usually a little broader and less coarsely reticulated than the 
typical form of A. gibbonsi, and a little narrower and less 
finely reticulated than A. g. rubella. Colour variable; 
sometimes dark like A. g. rubella, sometimes light as 
in A. gibbonsi s.s. In the typical form the mottling is 
inconspicuous and there is only a very slight trace of the 
paler median dorsal band. 

This form (and possibly also the last subspecies) may be 
slightly smaller than A. gibbonsi s.s. The following- 
are the measurements of a specimen which seemed to be 
practically mature : 

Length . . 46 mm. Greatest height . 7 mm. 

Breadth in middle 7'5 „ Distance of resp. 

Breadth at resp. opening from hind 

opening . . 6 „ end . . . 8 „ 

Breadth of foot-sole 4 „ 

Internal Characters. — Shell much depressed. Radula 
(PI. XVII, fig. 99 ; PI. XIX, fig. 124 ; text-fig. 3, d, p. 1.59) 
with a comparatively small number of very large teeth; 
central tooth typically absent ; cusps somewhat curved and 



THK CARNIVOROUS SLUGS OP SOUTH AFRICA. 2Ul 

conspicuously barbed on the lower side ; apophyses promi- 
nent ; formulae of two specimens (the second slightly imma- 
ture) : (20 + + 20) X 44, (21 + + 21) x 38. Anterior 
aorta passing- through loop of intestine (text-fig. 5, d, 
p. 183); pulmonary veins not very numerous. Penis long 
and twisted. Other characters agreeing with those of 
A. gibbon si 6\ .v. 

Habitat. — Port Shepstone, Natal (Burnup). 

Var. duplex n. var. 

Characters. — Back distinctly mottled, with a well-marked 
paler median band, the coloration being very similar to that 
of Apera gibbonsi s.s. Cusps of teeth rather longer than 
in the typical form of the subspecies, and having an additional 
barb on the upper side near the point (excepting in the 
smaller teeth) ; apophyses very prominent; a vestigial central 
tooth is present in most of the rows (text-fig. 3, e, p. 159) ; 
formula: (22 + (1) + 22) x 42. Otherwise similar to the 
Port Shepstone specimens of A. gibbonsi 1 up at a. 

Habitat. — Port St. John's, Pondoland (one specimen ; Mrs. 
E. Warren). 

Type. — The types of subspecies and variety are in the 
Natal Museum, Pietermaritzburg. Another specimen of the 
subspecies will be found in the British Museum. 

Affinities, etc. — This subspecies is especially characterised 
by its large barbed teeth. The course of the anterior aorta 
also serves to separate it from the typical form of Apera 
gibbonsi, as well as from A. g. rubella. But although 
the average shape of this subspecies is slightly different from 
either of these forms, the differences are so small that 
in practice I find that it is quite impossible to distinguish, 
without dissection, light specimens of the present subspecies 
from A. gibbonsi s.)<., and dark examples from A. g. 
rubella, unless, perchance, the odontophore is sufficiently 
protruded to enable one to see the teeth. It follows, there- 
fore, that none of these three forms can be identified with 



202 HUGH WATSON. 

certainty from an examination of their external characters 
only. It will be noticed that this race of A. gibbon si seems 
to have a more westerly distribution than the preceding- 
forms. 



Apera parva n. sp. 

External Characters (PL VIII, figs. 22, 23). — Animal 
very slender, tapering to an acute angle at the hind end. 
Back rounded, without keels. Outer lip of respiratory open- 
ing almost concealing the inner lip. Skin coarsely reticu- 
lated. Dorsal grooves about ^ mm. apart, separated by a 
single row of rug«, but united for 3 mm. in front of the 
respiratory opening. Lateral grooves near the dorsal grooves 
and extending forwards to the head, the right one ending in 
the genital opening. Radial grooves well-marked, but fewer 
than usual, only three arising from the respiratory opening 
on each side (in addition to the two median grooves). 
Oblique sub-lateral grooves less numerous than in A. 
gibbon si, being absent from the anterior third of the 
animal. Both radial and sub-lateral terminate in the single 
peripodial groove. 

Mr. Farquhar informs me that the living animal is fawn- 
coloured ; dark dorsally, but much lighter below. In alcohol 
the specimen has become yellowish brown, as shown in the 
figures. 

The following are the dimensions of the slug in alcohol ; 
but when alive and moving Mr. Farquhar tells me that it 
attained a length of about 35 mm. 

Length . . 25 mm. Greatest height . 3 mm. 

Breadth in middle 3 „ Distance from resp. 

Bi-eadth at resp. opening to hind 

opening . . 2*4 „ end . . . 5*5 „ 

Breadth of foot-sole I'd ,, 

Internal Characters. — Skin very thick. Shell (PI. XIII, 
fig. 44, 45) 1*6 X "9 mm., thick, depressed, with a flattened 



THE CARNIVOROUS SLUGS OF SOUTH AFIUCA. 203 

apex in front of the posterior margin ; sinus of moderate 
depth ; left posterior angle obliquely truncate. 

Pedal gland (PI. XIII, fig. 53). — Slender; glandular tissue 
gradually dying out a little behind the middle of the gland; 
duct less convoluted posteriorly than in A. gibbonsi. 

Nervous system. — Cerebral ganglia clearly sepai-ate ; 
buccal ganglia situated posteriorly and joined to the cerebi'al 
ganglia by long connectives ; cerebro-pedal and cerebro- 
pleural connectives rather long. 

Digestive System (PI. XVII, figs. 100-102 ; text-fig. 4, 
A, p. 161; PI. XXI, fig. 131).— Radula very large, 8 mm. 
long ; central tooth very small, absent in some i^ovvs ; others 
scarcely diiferentiated into laterals and marginals, becoming 
extremely large towards the edges of the radula ; cusps 
simple, very slightly curved, short towards the centre of 
the radula, longer towards the edges ; bases broader than in 
A. gibbonsi; corresponding teeth not opposite to each 
other; formula: (17 -f 1 + 17) x 48. Odontophore also very 
large ; posterior end joined by numerous short radial buccal 
retractors to the sides and floor of the body-cavity about the 
middle of its length ; odontophoral support rounded in front. 
CEsophagus short and broad, merging into a fusiform crop. 
Between the crop and the wide intestine is an oval stomach, 
into which the hepatic ducts open. Second loop of intestine 
absent, the alimentai-y canal curving over to the right side 
of the liver and then passing straight backwards. Salivary 
glands contiguous above the crop, but less closely fused than 
in A. gibbonsi. Liver in two approximately equal divisions, 
the intestine only traversing the right. 

Vascular System (PI. IX, fig. 28). — Pulmonary veins 
not very numerous. " Posterior aorta " dividing at its origin 
into three branches which supply the left division of the liver 
and the hermaphrodite gland ; right division of liver supplied 
by two arteries from the anterior aorta, which arise separately 
but close together; anterior aorta passing through loop of 
intestine ; anterior branches of buccal artery leaving the 
posterior branch as a single median vessel. 

VOL. 3, PART 2. 15 



204 HUGH WATSON. 

Reproductive System (PI. XXIII, fig. 146). — Her- 
maphrodite gland composed of only about four follicles, Avhich 
are relatively very large ; albumen gland in the specimen 
examined extending to the left side of the body-cavity, 
passing between some of the posterior buccal retractors ; 
common duct rather short, curving twice to the right and 
once to the left ; vagina rather narrow ; receptaculum seminis 
further forward than in the other species; receptacular duct 
broad and extremely short. Vas deferens not enlarged to 
form an epiphallus ; penis straight, rather short, and slightly 
swollen in the middle; penial retractor long and narrow. 

Habitat. — Near Fern Kloof, about three miles south of 
G-rahamstown, Cape of Good Hope (in a stony place with 
very little grass, at least 200 yards away from the bush ; 
Farquhar). 

Type. — The type is in the Natal Museum, Pietermaritzburg. 

Affinities, etc. — The only example of this species at present 
known Avas discovered by Mr. John Farquhar, of Grahams- 
town, on October 12th, 1913. In many respects the slug 
bears a close resemblance to Apera gibbonsi, and it might 
easily be mistaken for a young specimen of that species. 
Nevertheless, it diifers from all the forms which I have 
included under A. gibbonsi in so many characters that 
there can be very little doubt that it is specifically distinct. 
The following are the features in which some of the most 
important diif erences occur : the size ; the number of the 
radial and sub-lateral grooves ; the distance between the 
cerebral ganglia ; the size of the teeth towards the edges of 
the radula, and the width of the bases of all the teeth ; the 
position of the origin of the buccal retractors ; the stomach ; 
the course of the intestine ; the division of the arteries to 
both portions of the liver ; the hermaphrodite gland ; and the 
length of the receptacular duct and of the penial retractor, 
these last features being perhaps the most striking of all. 

Apera dimidia n. sp. 
External Characters (PI. YII, figs. 7-9; PL VIII, figs. 20, 



THE CAKNIVOKOUS SLUGS OF SOUTH AFRICA. 205 

21, 24). — Animal sometimes slightly swollen towards the 
middle, tapering to a blunt point at the hind end. Back 
rounded, without keels. Outer lip of respiratory opening- 
narrow, exposing the inner lip. Skin covered with a 
network of grooves, among which the main grooves are less 
conspicuous than in the other species of Apera. Dorsal 
grooves about 1 mm. apart (notwithstanding the small size of 
the species), usually separated by a double row of rugte, 
but united for about 3 mm. in front of the respiratory 
opening. Lateral grooves rather near the dorsal grooves, 
irregular, but sometimes extending as far as the head. 
Oblique sub-lateral grooves also irregular, seldom reaching 
the peripodial groove, and less numerous than in A. 
gibbo n si, being usually absent from the anterior third of 
the animal. Radial grooves fairly well marked dorsally, but 
like the sub-lateral grooves they cannot always be traced as 
far as the peripodial groove. The upper limits of the lowest 
rug^e on each side often form an irregular longitudinal groove 
near the peripodial groove and parallel to it. 

Body conspicuously mottled with greyish-brown pigment, 
which is chiefly concentrated to form an irregular and some- 
what discontinuous dark band along each side of the animal. 
Between these bands is a much paler dorsal area only 
sparsely mottled, darkest around the anterior border of the 
respiratory opening and lightest at the edges next to the 
dark bands. Below these the skin is darker than it is 
dorsally, being more thickly mottled, especially next to the 
dark band, though the pigmentation extends down to the 
peripodial groove. Both dorsally and laterally there is a 
tendency for some of the pigment to be concentrated in the 
dermal grooves, thus emphasizing the reticulation. Small 
patches of the same pigment are sparsely scattered over the 
foot-sole. The ground-colour of specimens preserved in 
spirit is yellowish white. 

The following are the dimensions (in alcohol) of two 
specimens, in both of wdiich the reproductive organs were 
fully developed : 



206 HUGH WATSON. 

Length . . . 32'5 mm. . 38'5 mm. 

Breadth in middle . 6'5 „ . 7'5 ., 

Breadth at respiratory 

opening- ... 4 „ . 4"75 ., 

Breadth of foot-sole . 3-75 „ . 4-25 „ 

Greatest height . 6 „ . 6*5 „ 

Distance from respira- 
tory opening to 

hind end . . 6 „ . 6*25 „ 

Internal Characters. — Skin not quite so thick as in most 
of the species (PI. XI, figs. 34, 35 ; PI. XII, figs. 36-39 ; 
PL XIII, fig. 40). Shell (PI. XIII, fig. 46) 3 x 1-8 mm., 
thin and translucent, convex ; apex rounded, quite at the 
posterior end ; sinus rather shallow. 

Pedal gland (PI. XIII, figs. 54, 59-62; PI. XIV, fig. 66).— 
It has the glandular tissue entirely concentrated in the 
anterior half, which is vei'y broad in consequence ; fold of 
terminal vesicle not divided into nai^row lamellae. 

Nervous System (PI. XIV, figs. 68, 69; PI. XV, figs. 
71, 73, 75; PI. XVI, figs. 80-86).— Cerebral ganglia with 
prominent accessory lobes; buccal ganglia normally situated 
posteriorly, joined to cerebral ganglia by comparatively short 
connectives ; nerves to pedal ganglia large. 

Digestive System (PL XVI, figs. 88-90; PL XVII, 
figs. 103-107; PL XVIII, figs. 115-122; PL XX, fig. 127; 
text-fig. 4,B, p. 161; PL XXI, fig. 132; PL XXII, figs. 136, 137). 
— Padula 4'8 mm. long ; central tooth absent; about thirteen 
laterals with very short curved cusps, and the same number 
of marginals with longer narrower cusps, on each side : 
corresponding teeth not opposite to each other ; formula : 
(25 -t- + 26) X 61. Odontophore rather large, with a 
single buccal retractor. CEsophagus nai*row and as long as 
the salivary ducts ; crop swollen anteriorly ; loops of wide 
intestine nearly vertical, mainly on the left side of the 
posterior division of the liver. Salivarj'' glands united above 
anterior end of crop. Anterior division of liver above crop 
and in front of intestine. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 207 

Vascular System (PI. IX, fig. 29 ; PI. XXII, fig. 140).— 
Pulmonary veins not very numei'ous ; no " posterior aorta " ; 
two main arteries of liver arising from anterior aorta as a 
single vessel, posterior artery supplying hermaphrodite gland 
as well as posterior division of liver ; anterior aorta passing 
to right of intestine. 

Reproductive System (PI. XXIII, figs. 147, 151).— 
Hermaphrodite gland oblong, with moderately large follicles; 
hermaphrodite duct rather more convoluted and swollen in the 
middle than in the other species ; common duct rather short, 
curving twice to the right and twice to the left; vagina 
broader than iu A. gibbonsi; receptaculum semiiiis small; 
receptacular duct much swollen anteriorly, the swollen part 
being attached to the adjacent body-wall by short transverse 
muscles on each side. Vas deferens becoming slightly 
swollen near the posterior half of the penis to form an 
epiphallus ; penis short, broadened posteriorly ; penial 
retractor also short. 

Habitat. — Port Shepstone, Natal (Burnup) ; Town Bush, 
Pietermaritzburg (on a mossy stone ; Mrs. Wan-en) ; Equeefa, 
Natal (Burnup). 

Type. — The type is in the British Museum. Other specimens 
will be found in the Natal Museum, Pietermaritzburg. 

AFFiNrriES, ETC. — This species may be distinguished at a 
glance from young specimens of any of the forms of Apera 
gibbonsi by its dark lateral bands, while the absence of 
keels sepai-ates it from the other species found in Natal. 
Internally it dift'ers widely from A. gibbonsi and A. 
parvain its radula, buccal retractor, oesophagus, liver, 
receptacular duct, etc. ; while in no other species is the pedal 
gland so broad in front and so slender behind. 

This species was fiist discovered by Mr. Burnup in March, 
1903. 

Apera purcelli Collinge. 

Apera puvcelli Cllge., Ann. S. Afr. Mus., 1901, vol. ii, p. 230, pi. 
xiv, figs. 1, 2 ; Connolly, Ann. S. Afr. Mus., 1912, vol. xi, p. 63. 



208 HUGH WATSON. 

External Characters (PL VIII, fig. 25; and 'Ann. S. Afr. 
Mus./ vol.ii, pi. xiv, figs. 1,2). — Animal rather broad anteriorly, 
tapering to a blunt point at the hind end. Back rounded, with- 
out keels. Outer lip of respiratory opening narrow, exposing 
the inner lip. Skin with well-marked reticulation, the rugas 
being rather large for the size of the slug. Dorsal grooves 
less than 4- mm. apart, separated throughout their length by 
a single row of rugee, not united in front of the respira- 
tory opening. Radial grooves well-developed, and extending 
down to the single peripodial groove, which is not very deep. 
The most anterior radial grooves branch from the dorsal 
grooves near the respiratoiy opening, instead of arising from 
the opening itself. There is not even a trace of lateral 
grooves, excepting perhaps for a very short distance ; but 
irregular oblique grooves occur in front of the radial grooves, 
and these correspond to the sub-lateral grooves of other 
species. 

Body dark grey or bluish black, the pigment being chiefly 
concentrated in the dermal grooves and thus emphasizing the 
reticulation. On each side there is a suggestion of a slightly 
darker lateral band, the intervening dorsal area being a little 
paler, although it tends to be darker towards the centre than 
next to the obscure lateral bands. Below these the skin 
becomes much lighter, the pigment dying out completely 
before it reaches the peripodial groove. The foot and the 
ground-colour of the back is whitish yellow in the case of 
specimens preserved in spirit, but in living examples the foot 
is said to be ferruginous. 

According to Mr. Collinge the type (in alcohol) is 2.5 mm. 
long, and its foot-sole is 4 mm. in breadth. I have only seen 
a young specimen, of which the following are the measure- 
ments : 

Length . . .16 mm. Greatest height . 3"5 mm. 

Breadth in middle •3'75 „ Distance from resp. 

Breadth at resp. opening to hind 

opening . . 2*25 „ end . . . 3"75 „ 

Breadth of foot-sole 2 ,, 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 209 

Internal Characters. — Skin thick. Shell (PI. XIII, 
figs. 47, 48) 1*2 X "6 mm. (in the immature specimen whose 
measurements are given above), thin and translucent, very 
convex ; apex rounded, overhanging the posterior margin to 
a considerable extent ; sinus shalloAv. 

Pedal gland (PI. XIII, fig. 55). — Glandular throughout its 
entire length. 

Nervous system. — The various ganglia more distinctly 
separate than in most species of A per a; buccal ganglia 
situated posteriorly, joined to cerebral ganglia by compara- 
tively short connectives. 

Digestive System (PI. XIII, fig. 55; text-fig. 4, c, p. 161). 
— Radula (of immature specimen) 1'8 mm. long; teeth less 
numerous than in most species ; central tooth small ; three 
or four laterals with very short cusps, and about four times 
as many marginals with longer narrower cusps, on each side ; 
formula (of immature specimen) : (18 + 1 H- 18) x 41. Odonto- 
phore with single buccal i-etractor. (Esophagus narrow ; 
loops of intestine nearly vertical, mainly on the left side of 
the liver. Salivary glands more or less united. 

Anterior aorta passing to the right of the intestine, 
"posterior aorta" probably absent. Penial retractor 
long, as in the next species. 

Habitat. — Table Mountain, Cape Town (scarce; Lightfoot). 

'J'ype. — The type is in the South African Museum, Cape 
Town. An immature specimicn will be found in the Cambridge 
University Museum of Zoology. 

Affinities, etc. — This rare species is the only member of 
the genus that is known to occur in the west of the Cape 
Province. In some respects it resembles Apera dimidia 
more than any of the other species, but it diifers from both 
A. dimidia and A. gibbon si in having the dorsal grooves 
separate throughout their entire length, in the glandular 
tissue extending to the terminal vesicle of the pedal gland, 
and in the long penial retractor. In these features the 
present form resembles the keeled species of Apera. Unfor- 
tunately the reproductive organs of this species are practically 



210 HOGH WATSON. 

unknown, and it is to be hoped that further specimens will 
be discovei-ed so that these organs may be described. 

Apera burnupi Smith. 

Apera burnupi Stnith. Ann. Mag. Nat. Hist. (6tli ser.), 1892. vol. x, 
p. 466. 

Apera natalensis CoUinge, Ann. S. Afr. Mus., 1900, vol. ii, p. 3, pi. i, 
figs. 3, 4, pi. ii, figs. 14, 15; Simroth, Naturwiss. Wochenschr., 1901, 
vol. xvii, p. Ill, fig. 7; Simroth, Bronn's Klass. u. Ordn. d. Tier- 
Reichs III, Gastr. Pulm., 1909, p. 143, fig. 42 c, pi. iv, figs. 9. 10 ; 
Collinge, Ann. Natal Mus., 1910, vol. ii, p. 167 ; Simroth, Bronn's 
Klass. u. Ordn. d. Tier-Reichs III, Gastr. Pulm., 1912, p. 611; 
Connolly, Ann. S. Afr. Mus., 1912, vol. xi, p. 63. 

External Characters (PI. VII, figs. 10, 11; PL A^II, 
fig. 26; and 'Ann. S. Afr. Mus,/ vol. ii, pi. i, figs. 3, 4). — 
Animal broad and flattened towards the hind end, tapering 
anteriorly, with four longitudinal keels, two on each side 
of the body. Notwithstanding the two pairs of keels 
the animal is squarish in section, excepting posteriorly, 
as the lower keels ai-e not far from the edges of the foot. 
Upper keels prominent though blunt, nearly parallel to each 
other in the middle of the body and separated by about 
seven- eighths of the breadth of the animal. Anteriorly they 
converge a little towards the head. In the posterior three- 
eighths of the slug the upper keels become very prominent, at 
first diverging and then curving round to meet in a very 
obtuse angle above the posterior extremity of the foot (but 
the angle is probably less obtuse in young than in adult 
specimens). They thus enclose a large flattened oval area, 
sloping downwards and backwards, and having the respira- 
tory opening slightly in front and to the right of its centre. 
Lower keels less prominent, twice as far from the upper keels 
as from the edges of the foot, extending for nearly three 
quarters of the length of the animal but not reaching the 
hind end. Foot rather broad, but tapering behind as in the 
other species, so that the posterior dorsal expansion of 
the body overhangs the edges of the foot on each side. 
Outer lip of respiratory opening rather nai'row, exposing the 



THE CARNIVOliOUS SLUGS OF SOUTH AFRICA. 211 

inner lip. Skin coarsely reticulated, the rug£e being especi- 
ally prominent on the anterior part of the back. Dorsal 
grooves very conspicuous, about 1 mm. apai't (or rather more 
in the centre), separated by a row of rugae which is usually 
double in the middle of the body, converging as they approach 
the respiratoi'y opening, but not uniting with each other. 
Lateral grooves deep but rather irregular, extending along 
each side of the body as far as the head, a little nearer the 
upper than the lower keels, the right one terminating in the 
genital opening. Below each latei^al groove another more 
irregular groove branches from the most anterior radia.l 
groove and runs forward above the lower keel until it unites 
with the lateral groove on the side of the head. There are 
two or three irregular sub-lateral grooves on each side, and 
also three or four equally irregular supra-lateral grooves, 
parallel to the most anterior radial grooves and connecting 
the dorsal grooves with the lateral grooves on each side of 
the back. The upper keels are notched by these supra-lateral 
grooves as they cross them, but the keels are more deeply 
notched by the radial grooves, which are rather numerous 
and very conspicuous. Both radial and sub-lateral grooves 
terminate below in a longitudinal groove, parallel to the peri- 
podial groove, but separated from it by a narrow row of 
rugae. In addition to the usual transverse grooves there is 
an obscure longitudinal groove on the foot-fringe, also 
parallel to the peripodial groove. 

Body mottled with greyish-brown patches of colour, which 
sometimes have a greenish-blue tinge. The mottliug is chiefly 
concentrated on the sides of the body between the keels, but 
patches of colour also occur between the lower keels and the 
peripodial groove, and more sparsely on the dorsal area. The 
ground-colour of specimens preserved in spirit is usually pale 
yellow slightly tinged with reddish-brown dorsally ; but in 
living examples the skin is more deeply coloured, the back 
being reddish-brown, and the sides and foot-sole tinted with 
orange-red. 

The following are the dimensions (in alcohol) of the type 



212 HUGH WATSON. 

of A. natalensis according to Colliiige, and of the still 
larger specimen belonging to the South African Museum 
shown in PL VII, fig. 10 : 

Length . . . . .65 mm. . 77 mm. 

Breadth between upper keels 

in middle . . . . 10 „ . 12'5 „ 

Breadth between upper keels 

behind respiratory opening . 13*5 ,, . 17 ,, 

Breadth betw^een lower keels 

in middle .... — . 14'5 „ 

Breadth of foot-sole . . 9".5 „ . 11 "5 „ 

Greatest height (15 mm. in front 

of respiratory opening) . — .14 ,, 

Distance from respirator}' open- 
ing to hind end ... — .1 8"5 „ 

Internal Characters. — Skin extremely thick, especially 
towards the hind end (PL IX, fig. 30). Shell (PL XIII, 
fig. 49) 5'5 X 3*2 mm., of moderate thickness, convex, w-ith a 
conical apex projecting slightly beyond the posterior margin ; 
sinus rather deep. 

Pedal gland (PL XIII, fig. 56). — Exceptionally large, 
its loops wrapping round the other organs; glandular 
tissue extending throughout its entire length. 

Nervous system. — Buccal ganglia situated anteriorly; 
connectives short. 

Digestive System (PL XYII, figs. 108, 109; PL XX, 
fig. 128; text-fig. 4, D, p. 161; PL XXI, fig. 133).— Radula 
(of large specimen) 6mm. long; central tooth present, with a 
vei-y short single cusp ; eleven laterals with short double cusps, 
and more than twice as many marginals with long, narrow, 
nearly straight cusps, on each side, the transition between 
latei-als and marginals being abrupt ; formula (of large speci- 
men) : (25 -l-11 + l-fll-f 25) X 77. Odontophore small, with 
a single slender buccal retractor; odontophoral support bluntly 
pointed in front. CEsophagus narrow and as long as the 
salivary ducts ; crop cylindrical ; intestine narrow^, first loop 
deep, posterior continuation on right side of liver. Salivary 



THE CAENIVOEOUS SLUGS OF SOUTH AFKICA. 213 

glands separate. Liver of two approximately equal divisions, 
the intestine only traversing the right. 

Vascular System (PI. IX, fig. 30). — Pulmonary veins 
forming a network. " Posterior aorta " supplying left 
division of liver and hermaphrodite gland, and sending a 
small branch to posterior end of right division of liver; 
remainder of right division supplied by two ai-teries from the 
anterior aorta, which arise separately, but close together; 
anterior aorta passing through loop of intestine. 

Reproductive system (PI. XXIII, fig. 148).— Herma- 
phrodite gland composed of relatively smaller follicles than in 
A. diniidia; hermaphrodite duct convoluted; common duct 
very long, rather narrow, and much convoluted and twisted ; 
vagina bent at I'ight angles towards its anterior end ; recepta- 
culum seminis small ; receptacular duct greatly swollen 
towards the anterior end. Penis moderately long ; anterior 
part narrow and more or less twisted, posterior part broad 
and appearing as though it were double owing to the presence 
of a longitudinal groove along one side ; vas deferens dis- 
appearing into this groove about 5 mm. from the posterior 
end of the penis ; penial retractor long- and narrow. 

Habitat. — Chase Bush, Pietermaritzburg (Burnup) ; Rich- 
mond, Natal (nearly in the centre of a rotten log; Ward). 

Type. — The type of A. burnupi is in the British Museum, 
of A. natalensis in the South African Museum, Cape Town. 
Another specimen will be found in the Natal Museum, Pieter- 
maritzburg. 

Affinities, etc. — This striking form differs widely from all 
the preceding species not only on account of its conspicuous 
keels, but also in the double cusps of the lateral teeth of the 
radula and the separate salivary glands. 

Apera sexangula n. s-p. 

Apera burnupi ''Smith" ; Collinge, Ann. Mag. Nat. Hist. (6th ser.), 
1897, vol. XX, p. 221, pi. v, figs. 1-6; Pilsbry, Nautilus, 1898, vol. xii, 
p. 12; Collinge, Ann. S. Afr. Mus.. 1900, vol. ii, p. 4, pi. i, figs. 5. 6; 
Collinge, Journ. of Mai., 1901, vol. viii. p. 71, fig. 1 ; Collinge, Jouvn. 



214 HUGH WATSON. 

of Mai.. 1902, vol. ix, pi. vi, fig. 6G ; Simroth, Bronn's Klass. u. Ordii. 
d. Tier-Reicbs III, Gastr. Piilm., 1909, pi. iv, fig. 11 ; Collinge, Ami. 
Natal Mus., 1910, vol. ii, p. 166; Simrotli, Bronn's Klass. n. Ordn. 
d. Tier-Reichs III, Gastr. Pulni., 1912, p. 611; Connolly, Ann. 
S. Afr. Mus., 1912, vol. xi, p. 62. 

External Characters (PI. VII, figs. 12, 13; and 'Ann. S. 
Afr. Mas.,' vol. ii, pi. i, figs. 5 and 6). — Animal rather slender, 
approximately hexagonal in section, owing to the presence of 
four prominent longitudinal keels, two on each side of the body. 
Upper keels separated by about three-fifths of the breadth 
of the slug, and nearly parallel to each other, excepting pos- 
teriorly, where they diverge very slightly just in front of the 
respiratory opening, and then gradually converge behind it, 
until they unite to form a single median keel, 3 or 4 mm. in 
length, at the hind end of the animal. Lower keels extend- 
ing along the whole length of the slug, about half way 
between the upper keels and the edges of the foot, but sloping 
downwards towards the hind end. The areas between the 
keels are nearly flat when the animal is in motion, but when 
it contracts they become deeply concave. Body, as seen 
from above, tapering to an acute angle posteriorly, but when 
viewed from the side the hind end appears blunter owing to 
the short median keel (though not always so rounded as in 
fig. 13). Outer lip of respiratory opening narrow, exposing 
the inner lip, Rug^e minutely subdivided. Dorsal grooves 
well marked, usually about 1 mm. apart, separated by a row 
of rugfe which is single in front and behind but often 
becomes irregularly double towards the middle, converging 
slightly as they approach the respiratory opening, but not 
uniting with each other. Lateral grooves also conspicuous, 
extending along each side of the body as far as the head 
about half way between the upper and lower keels, the right 
one ending in the genital opening. Sub-lateral grooves few 
and irregular, only two or three being traceable on each hide. 
Radial grooves well marked, forming slight notches in the 
keels where they cross them. Both radial and sub-lateral 
grooves terminate below in a longitudinal groove, parallel to 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 215 

the peripodial groove, but separated from it by a narrow row 
of riigfe. 

Body mottled with greyish-brown pigment, excepting along 
the edges of the keels. The colour tends to be more concen- 
trated towards the hind end, and along the sides of the 
animal just below the upper keels, although it extends down 
to the peripodial groove. Asa rule, minute reddish-brown 
specks are also thickly scattered over the body, and these, 
unlike the greyer patches of colour, occur also on the foot- 
sole, though much less abundantly than on the back. The 
ground-colour of specimens preserved in spirit is yellowish- 
white, very slightly tinged with dull red dorsally ; but in life 
the skin is more deeply stained with dull red pigment, the 
general colour of the living animal being usually chestnut or 
reddish-brown, paler on the keels and f.oot.^ 

The following are the dimensions (in alcohol) of a small 
but mature specimen, whose reproductive organs are shown 
in PI. XXIII, fig. 149, of a slightly larger example in which 
the genital system is not quite fully developed, and of an 
unusually large specimen in the Natal Museum, of which Mr. 
Burnup has kindly given me the measurements. 

mm. mm. mm. 

Length ...... 

Breadth between upper keels in 

middle ...... 

Breadth between upper keels at 

respiratory opening 
Breadth between lower keels in 

middle ...... 

Breadth of foot- sole 
Greatest height (in middle) 
Distance from respiratory opening 

to hind end 11-5 .13 .16 

Internal Characters. — Skin thick, especially towards the 
hind end. Shell (PI. XIII, figs. 50, 51) 4x2 mm., of 

' For a description of the living animal, see ' Ann. Natal Mus.,' 
1910, vol. ii, pp. 166, 167. 



52-5 . 


, 56 


. 77 


4-5 . 


6 


, 10-5 


4 


. 4 


. 8-5 


7-5 


. 10 


. 16 


4 


. 4-5 


. 10-5 


7 


9 


. 15-5 



216 HUGH WATSOX. 

moderate thickness, convex, laterally compressed, Avitli a 
prominent conical apex projecting beyond the posterior 
margin; sinus of moderate depth. 

Pedal gland (lig. 57). — Much smaller than in the last 
species, curving alternately to the right and to the left in a 
comparatively regular manner ; glandular tissue extending 
throughout its entire length. 

Nervous system (PI. XY, figs. 72, 74, 76).— With buccal 
ganglia situated anteriorly; connectives short; nerves to pedal 
gland rather slender. 

Digestive System (PI. XVI, fig. 87; PI. XVII, figs. 
110-114; Text-fig. 4, E, F, p. 161; PL XXI, fig. 134, PI. XXII, 
fig. 135). — Radula 4 or 5 mm. long ; central tooth present, 
with a very short, broad, single cusp ; eight or nine laterals 
with short double cusps, and more than twice as many 
marginals with longer narrower cusps, on each side, the 
transition between laterals and marginals being rather abrupt; 
formula (of a Port Shepstone specimen) : (23 + 9 + 1 + 9 + 
23) X 63. Odontophore small, with a single slender buccal 
retractor. CEsophagus narrow, shorter than the salivary 
ducts ; crop swollen between the salivary glands ; intestine 
narrow, fii'st loop deep, posterior continuation on right side of 
liver. Salivary glands separate. Liver of two approximately 
equal divisions, the intestine only traversing the right. 

Vascular System (PI. IX, fig. 31). — Pulmonar}^ veins 
forming a network. " Posterior aorta " supplying left division 
of liver and hermaphrodite gland; right division supplied by 
two arteries from the anterior aorta which arise at some 
distance from each other ; anterior aorta passing through loop 
of intestine. 

Reproductive System (PI. XXIII, fig. 149). — Her- 
maphrodite gland consisting of a cluster of small follicles ; 
hermaphrodite duct only very slightly convoluted; common 
duct very long and much convoluted and twisted; vagina 
i-ather broad; receptaculum seminis rather small; receptacular 
duct fusiform, swollen towards the anterior end. Penis long 
and narrow excepting at the posterior end, where it is 



I 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 217 

broadened; end o£ vas deferens swollen to foi-m an epipliallus 
about 2'5 mm. long, which bears anteriorly a minute flagellum; 
penial retractor long and narrow. 

Habitat. — Port Shepstone, Natal (Burnup) ; Hilton Road, 
near Pieterraaritzburg (Biirnup) ; Grahamstown, Cape of 
Good Hope (Farquhar, French). 

Type. — The type is in the British Museum. Other specimens 
will be found in the Natal Museum, Pieterniaritzburg, in the 
South African Museum, Cape Town, and in the Academy of 
Natural Sciences, Philadelphia. 

Affinities, etc. — This species resembles Apera burnupi 
in many respects, and was at one time confused with that 
species. Nevertheless, the two forms can be very easily dis- 
tinguished by their external charactei's alone, for in the 
present species the upper keels meet at an acute angle behind, 
Avhere they form a short median keel, and the lower keels are 
equally prominent and extend the whole way to the hind end 
of the animal. The colour of the two forms is also different. 
Internally the most conspicuous features in which the present 
species differs from the last are to be found in the pedal 
gland and the reproductive organs. 

Hitherto this species has usually been known as "Apera 
burnupi," but an examination of the type-specimen of 
A. burnupi leaves no doubt that that name should be 
applied to the preceding species, which Collinge subsequently 
named A. natalensis, and it has therefore been necessary 
to give a new name to the present form. 

THE PHYLOGENY OF THE SPECIES OF APERA. 

It is evident that Apera burnupi and A. sexangula 
are closely related to each other. Of these it is probable that 
the former is the more highly specialised ; for in the pedal 
gland, the reproductive organs, and the keels, A. sexangula 
seems to be rather more primitive than A. burnupi. A wide 
gap separates these two species from those without keels ; 
nevertheless, A. purcelli resembles A. burnupi and A, 



218 HUGH WATSON. 

sexangula in not a few characters, notably the pedal gland, 
the long penial retractor, and the complete separation of the 
dorsal grooves. Moreover, it is probable that the ancestral 
form from which A. burnupi and A. sexangula have been 
evolved resembled A. purcelli still more closely, since the 
keels and the double cusps of the lateral teeth of these species 
cannot be regarded as primitive characters. Yet it is not 
likely that in A. purcelli we have the direct ancestor of 
these two species, for A. purcelli has in some respects 
become modified in a different direction, its central tooth 
having become smaller and the course of the intestine having 
become quite different from what we must regard as the 
more primitive arrangement found in A. burnupi, A. sex- 
angula, and A. gibbonsi. It is therefore probable that 
A. purcelli must be regarded as having diverged somewhat 
from the branch which gave rise to the keeled species. 

Apera dimidia in some respects resembles A. purcelli, 
but it is much more highly specialised. The glandular tissue 
of the pedal gland has become entirely concentrated around 
the anterior half of the duct ; the central tooth of the radula 
has disappeared, and the laterals have become more numerous ; 
the penial retractor has become quite short ; the dorsal 
grooves are united posteriorly: in fact, while A. dimidia 
has probably been derived from the same branch as A. 
purcelli, it has diverged very far from both A. purcelli 
and the keeled species. 

Apera gibbonsi and A. parva differ widely from the 
other species in their enormous radula and numerous radial 
buccal retractors, as well as in the oesophagus, receptacular 
duct, shell, coloration, etc., and apparently in the structure 
of the terminal vesicle of the pedal gland. In some respects 
they resemble A. dimidia most closely, especially in the 
absence of glandular tissue from the posterior end of the 
pedal gland, the short common duct, the union of the dorsal 
grooves in front of the respiratory opening, and the position 
of the lateral grooves ; and it might be possible to regard 
A. gibbonsi and A. parva as being derived from the 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 219 

same branch as A. dimidia, but having become still more 
highly specialised in many ways. On the other hand, we find 
that these species seem to be more primitive than either A. 
dimidia or A. purcelli not only in the uniformity of the 
teeth of the radula and the presence of a well-developed 
central tooth in typical examples of A. gibbonsi, but also 
in the arrangement of the intestine and liver. Now the 
nature of these characters renders it improbable that if they 
had once become so much modified they would revert to their 
original condition ; and I am therefore inclined to regard the 
resemblances between A. gibbon si and A. parva on the 
one hand, and A. dimidia on the other, as due to parallel 
evolution. 

Apera gibbonsi and A. parva are probably even more 
closely related to each other than are A. burnupi and A. 
sexangula. But A. parva is decidedly more primitive 
than A. gibbonsi in its nervous system and long penial 
retractor, and in possessing a true stomach. On the other 
hand, in a few features, such as the buccal retractors and 
hermaphrodite gland, we must regard A. parva as being 
moi-e highly specialised than A. gibbonsi or any other 
member of the genus. 

If these views be correct, the phylogeny of the species of 




Apera might possibly be somewhat as shown in the above 
genealogical tree. 

VOL. 3, PART 2. 16 



220 HUGH WATSON. 

THE OCCURRENCE AT CAPE TOWN OP 
TESTACELLA MAUGEI FEB. 

This well-known species ^ is the only carnivorous slug, in 
addition to those belonging to the genus Apera, which is 
kuown to occur in South Africa, for it is probable that 
Ceratoconchites schultzei {Siviroth) is the larva of a 

The genus Testacella can be distinguished at a glance 
from all the other g*enera of slugs known to inhabit South 
Africa, because in this form alone the hind end of the back is 
covered by an external shell. In some respects the anatomy of 
Testacella resembles that of Apera, but it is so well known 
that it is unnecessary for me to describe it here. More than 
a hundred years ago Cuvier'^ gave an account of the internal 
organs of one member of the genus ; and since then the re- 
searches of Gassies and Pisclier, de Lacaze-Duthiers, Plate, 
Simroth, and others, have given us a fuller knowledge of the 
anatomy of Testacella than of almost any other carnivorous 
snail or slug. 

The following are the principal characters by which 
Testacella maugei can be distinguished from the other 
species of the same genus : (1) the comparatively large size 
of the shell, and its length and convexity; (2) the widely 
separated origin of the lateral grooves ; (3) the presence ol" a 
central tooth in the radula; (4) the small number of the 
buccal retractors ; (5) the unusual length of the tentacular 
retractors, and the fact that they both arise to the left of the 
middle line; (6) the swollen anterior end of the receptacular 

' See Apj)endix for the more important references to Testacella 
maugei. 

■^ Simroth, H., 'Zool. Auz. Leipzig.' 1907, vol. xxxi. pp. 794, etc.; 
Simroth, H., 'Deutsche Siidpolar Exped.," 1910, vol. xii, Zool. iv, p. 172 ; 
Bottger, O., 'Abhandl. Senckenb. Naturf. Ges. Frankfurt,' 1910, vol. 
xxxii, p. 433. 

3 ' Ann. Mus. d'Hist. Nat.,' 1804, vol. v, pp. 435-444. pi. xxix, figs. 
6-11. 



THE CARNIVOROUS SLUGS OF SOUTH AFRFCA. 221 

duct, its comparatively great lengtli, and its lateral junction 
with the receptaculum seminis ; (7) the convolution of that 
part of the vas deferens which lies next to the free oviduct ; 
(8) the length of the narrow anterior part of the penis, and 
its broader posterior end, without any flagellum. 

It will be seen from PI. XXIV that the specimens of 
Test ace 11a from Cape Town which I have examined 
possess all these characters, and I have, therefore, no 
hesitation in assigning them to T. maugei. Nevertheless 
the South African examples of this genus have usually been 
named "T. aurigaster Lai/ard."^ Now, although Major 
Connolly- believes that Layard published a description of 
this supposed species, I have not been able to find any such 
description, nor any type-specimens, and I cannot therefore 
state definitely that T. aurigaster is a synonym of T. 
maugei, as it is conceivable that more than one species of 
Testacella may have found its way to Cape Town durino- 
the last sixty years. At the same time, all the specimens 
that I have seen labelled " T. aurigaster " have proved to 
belong to T. maugei, and I think that T. aurigaster may 
be safely expunged from the list of South African Mollusca. 

The slug- occurs in gardens at Cape Town, and I have little 
doubt that it has been introduced into South Africa by man 
notwithstanding- Dr. Simroth's arguments in favour of the 
possibility of Testae el la being a native of that country.-^ I 
believe that the natui-al distribution of the genus Testacella 
is limited to the western part of the Paltearctic i*egion, from 
Great Britain and Hungary to the Canary Islands, and that 

' The following is a copy of a manuscript note written by Layard 
liimself, for which I am indebted to Major Connolly : " Testacella 
aurigaster Layard. I only found this shelled slug in the Gardens round 
Cape Town. It was common in the grounds of the Sotith African Museum, 
which ivas btiilt at the lower end of the Botanical Garden in Cape Town. 
The belly of the animal was a rich golden yellow, hence the name selected. 
It devours large loorms, & will afsimilate individuals far larger than 
itself r' 

■ -Ann. S. Afr. Mus.,' 1912, vol. xi, p. 64. 

^ ' Zool. Anz. Leipzig,' 1907, vol. xxxi, p. 796. 



222 HUGH WATSON. 

its occurrence in Philadelphia, Cape Town, and Auckland is due 
to the agency of man. Testacella maugei occurs not only 
in the south-west of England, but also in Portugal, Madeira, 
and the Canaries ; and it seems to me that it might easily 
have been transported to South Africa from any of these 
localities, amongst the roots of plants. For the species of 
Testacella are especially common in the rich soil of gardens 
where worms are abundant; and it is known that under 
adverse conditions a specimen can surround itself with a sort 
of cocoon of hardened slime which protects it from drought, 
and may remain dormant in this state for several weeks. 

It will be seen from PI. XXIV, fig. 159, that in some of the 
South African specimens the posterior end of the penis is 
curved round in a peculiar manner, and the retractor muscle 
is attached to the vas deferens, a short distance in front of it.^ 
Possibly this unusual arrangement may have been partly 
caused by the way in which the animal contracted when it 
was killed, for it only occurred in some shrivelled specimens, 
which, in other respects, seemed to be identical with the 
rest. 

Figs. 157 and 158 represent two of the shells labelled 
"T, aurigaster" in the MacAndrew collection at Cam- 
bridge. The first shows the usual shape of the shell in 
T. maugei, while the second may be assigned to the variety 
aperta Taylor,^ a form which was originally described from 
the Azores. 

THE AFFINITIES OF APERA AND TESTACELLA. 

The Evolution of Carnivorous Characters. 

Both Apera and Testacella undoubtedly belong to the 
group of the Stylommatophora comprising the carnivorous 

' The junction of the retractor muscle with the vas deferens is also 
shown in Webb's figure of the genital organs of T. maugei (' Journ. 
of Malac.; 1897, vol. vi, pi. vi, fig. 3). 

■^ ' Monog. L. and F.-W. MoUusca Brit. Isles,' 1902. vol. ii, p. 24, pi. i, 
fitr. 15. 



THE CAKNIVOliOUS SLUGS OF SOUTH AFKIOA. 223 

snails and slugs^ a group which is usually kuown as the 
A gnat ha, since very few of these forms possess a jaw. But 
to state this is to say veiy little about the affinities of these 
genera, for it is doubtful whether the various carnivorous 
snails are at all closely related to one another, some authorities 
asserting that the tribe A gnat ha is polyphyletic. 

Certain eminent malacologists have held that the Stylom- 
matophora may be divided into two groups, one mainly 
herbivorous, the other mainly carnivorous ; and they would 
account for the diversity of the carnivorous forms by 
supposing that they had evolved in a manner parallel to the 
gnathophorous families, which show a similar diversity. 
This view was well expressed by Fischer, who, writing so 
long ago as 1873, said: "Je suis persuade qu'on pourra con- 
stituer, dans quelques aunees, une serie d^Agnathes ou Testa- 
cellidas ayant une grande extension, et dont les genres seront 
representatifs de ceux des Helicida3, ainsi que, chez les Mammi- 
feres, certains groiipes des Marsupiaux i-epresentent d'autres 
groupes des Monodelphes/^^ On the other hand, many 
modern authorities maintain that the various families of 
carnivorous snails and slugs have been evolved independently 
from different families of the Gnathophora, and that they 
are in reality no more nearly related to one another than are 
the European and Tasmanian wolves. The characters which 
the agnathous families undoubtedly possess in common they 
would explain as being due to convergence, brought about by 
their common acquisition of carnivorous habits.- Now there 
is no a priori objection to this second view, for we know 
that many gnathophorous forms occasionally devour animal 
food, and there seems to be no reason why this should not 
have become the usual diet in some cases. The question, 
therefore, can only be settled by a careful examination of the 
comparative anatomy of the various families included in the 

' • Joiu-n. de Conchyl.," vol. xxi, p. 12. 

• See an iuterestiug paper by Dr. Simroth CNaturwiss.Wochensclir.,' 
1901, vol. xvii, pp. 109-114, 121-127, 137-140), m which he has ably 
advocated tlie polyphyletic origin of the carnivorous snails and slugs. 



224 HUGH WATSON. 

Agnatlia. If we find that two agnathous families only 
resemble each other in those features which are likely to 
have been acquired through carnivorous habits^ and are very 
similar to diiferent gnathophorous families in their other 
characters, then we may assume that they are probably of 
different origin ; but if we find that the resemblances between 
two families of agnathous snails cannot all be explained in 
this way, and that these families are in their most important 
features more similar to each other than to any families of 
the Gnathophora, the probability is that they are closely 
related. It is evident, however, that before we can discuss 
the affinities of any particular genus, we must have a clear 
idea as to how a snail is likely to become modified if it adopts 
carnivorous habits, and which of the features characteristic of 
Apera, Testacella, and the other genera included in the 
Agnatha, are likely to be due to their animal food. 

Snails and slugs find their food chiefly by means of their 
sense of smell, and one might expect this sense to be especi- 
ally well developed in the carnivorous forms, because animals 
which move slowly enough for snails to catch them must be 
very much more difficult to find than plants. Now in man}^ 
of the Agnatha, and more especially in the Rhytididas 
and the 01eacinid{«, the olfactory organs at the tips of the 
upper tentacles are so large that the eye comes to occupy a 
position some distance behind the extremity.^ Moreover 
Plate has shown that Testacella, unlike most of the 
Stylommatophora, retains a pallial olfactory organ. The 
unusual development of these sense-organs might be expected 
to lead to a corresponding development and concentration of 
the sensory nerve-centres, and accordingly we find that in 
nearly all the carnivorous forms the cerebral ganglia are 
large and close together, Phrixolestes being perhaps the 
most notable exception. 

Carnivorous snails and slugs prey chiefly upon the herbi- 

' See Strebel, H., 'Beitrag z. Keiintn. d. Fauna Mexikan. L.- u. Siiss- 
wasser-Conchyl.,' 1878, vol. iii, pi. xv, fig. 1 c; Suter, H., ' Jourii. of Mai.,' 
1899, vol. vii, pi. iii, fig. 1 a. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 225 

vorous forms and on worms, for most ai'thropods move too 
quickly and are too well protected by their cliitinous exo- 
skeleton to fall a prey to snails. Now worms burrow in the 
soil, and snails hide themselves in crevices and retire deeply 
within their shells when attacked ; it is therefore evident that 
a carnivorous snail will find the presence of a bulky un- 
yielding shell on its back a great inconvenience when it is 
trying to get near its victim. In order to obviate this diffi- 
culty the shell has become modified in various ways. In the 
first place we find that in Paryphanta it has become more 
or less flexible owing to the degeneration of the inner cal- 
careous layer. Secondly, the shape of the shell has become 
altered in many of the carnivorous genera. In Diplom- 
phalus, for example, the shell has become greatly flattened. 
This will enable the animal to penetrate into crevices, but 
it is obvious that the breadth of the shell will have to be 
reduced as well as its height, if the snail is to crawl into 
narrow holes. Now the only way in which both the height 
and breadth of a depressed or heliciform shell can be reduced 
is by the curvature of the axis or columella, until its 
direction corresponds more nearly with that in which the 
animal moves ; and this is what has occurred in that remark- 
able genus of carnivorous snails, Streptaxis. If, however, 
the shell has a raised spire, the columella naturally takes up 
a position parallel to the direction in which the animal moves, 
and in this case it is only necessary for the height of the 
spire to be increased in ordei- to reduce the diameter of the 
shell. This is what has taken place in the large genus 
Ennea, in which only the young form retains a comparatively 
low spire. In Diaphora this principle is carried to an 
extreme, some species having shells with greatly produced 
spires composed of as many as twenty whorls. Indeed, so 
long does the spire become that the animal can no longer 
occupy the whole of it, and secretes a new internal wall 
cutting oif part of the upper whorls, which may become 
decollated, as in D. teles copium Mlldff. A more efficient 
method of reducing the diameter of the shell (but one 



226 HUGH WATSON. 

which probably necessitates a greater change in the structure 
of the animal) is for the whorls themselves to become laterally 
compressed. This has taken place to some extent in Strepto- 
stele and Obeliscella among the Streptaxidse, but it is 
especially characteristic of the Oleacinidge. In some 
members of this family, such as Streptostyla gracilis 
I*ilsbrij, the whorls have become so narrow that the form 
of the shell closely resembles that of the marine carnivorous 
genus Con us — a striking example of convergence due to the 
acquisition of similar habits. Thirdly, the position of the shell 
may be altered, so that the part of the animal in front of the 
shell becomes lengthened. This is admirably shown in Hedley's 
figure of Rhytida lampra {Pfr.),^ but it occurs to some 
extent in most of the carnivorous genera. A variety of this 
modification is fouud in such forms as Ennea densecostu- 
lata Mlldff. and Diaphora eutrachela Mlldff., in which 
the greater part of the shell has moved further back by the 
elongation of the last whorl towards the mouth. Fourthly, 
the shell may not only be moved backwards, but it may 
become reduced and flattened, until it either disappears 
altogether as in Selenochlamys, or sinks into the skin as 
in Apera. No projection will then remain to impede the 
animal's progress when it follows worms into their burrows, 
and it is more especially in the vermivorous genera that this 
extreme modification seems to have occurred. We see, then, 
that the acquisition of carnivorous habits may lead to a great 
alteration in the structure, form, position, or size of the 
shell. 

The movement of the shell to the hind end of the body, and 
its gradual degeneration, will greatly modify the animal itself. 
Many of the organs will be subject to partial detorsion. The 
lung and respiratory orifice will move back with the shell and 
mantle, so that the animal will become opisthopneumic ; and 
it is probable that it is a direct advantage to a carnivorous 
form to have its respiratory opening near the hind end of the 
back, because this is the part which is least likely to be 
' ' Proc. Linn. Soc. N.S.W.* (2nd ser.), 1891, vol. vi, pi. iii, fig. 3. 



THE CARNIVOROUS Sr,UGS OF SOUTH AFRICA. 227 

covered when the greater portion of the body is under the 
surface ot" the ground, or inside its victim's shell. 'J'he anus 
will move back with the respiratory opening, and. the rectum 
will thus be directed posteriorly (text-fig. 5^ cf. c and d, p. 183) . 
With the degeneration of the shell the pallial organs will 
become modified, and the posterior position of the pulmonary 
veins may pull round the auricle until it lies posterior to the 
ventricle, as in Testae ell a and a few other genera (text- fig. 
6, p. 244). The organs which formerly occupied the viscex*al 
hump will become reduced in size and pushed forward into 
the general body-cavity. The pressure of these will retard 
the backward growth of other organs such as the receptaculum 
seminis, and it is therefore not surprising to find that the 
slugs have as a rule shorter receptacular ducts than the snails 
to which they are probably most nearly allied. The columellar 
muscle will disappear with the degeneration of the shell, and 
the various retractors that originally arose from it will become 
attached to different parts of the skin, which will become 
thickened to take the place of the shell (see p. 126). And 
as the anterior part of the body becomes lengthened, the 
origins of the tentacular and penial retractors will tend 
to move forwards. Thus the mere fact that animal food 
is less easily accessible than vegetable food may lead to 
profound changes in the anatomy of a snail Avhich becomes 
carnivorous. 

When the snail has approached its victim it will require to 
feel exactly where to attack it with its i-adula, and will, if 
possible, try to prevent the animal escaping until the teeth 
are fixed in its flesh. Accordingly we find that nearly all the 
carnivorous genera have a pair of special feelers at each side 
of the mouth beneath the lower tentacles. In Euglandina 
these feelers are very long and are supposed to be used for 
prehension, but according to F. M. Woodward ^ their function 
in Natalina caffra [Fer.) is probably purely tactile. Miss 
Davies"'' has recently discovered that in Paryphanta com- 

1 ' Proc. Mai. Soc.,' 1895, vol. i, p. 271. 

- ' Proc. Roy. Soc. Victoria,' 1913, vol. xxv, p. 225. 



228 HUGH WATSON. 

pacta Cox & Hedley and P. atramentaria {Shuttl.) these 
papillae are glandular in structure. Now in Peripatus, an 
arthropod genus whose habits are not unlike those of the 
carnivorous snails and slugs, there are two large slime-glands 
which open on the oral papillas, and their secretion is used to 
entangle the prey. The carnivorous slug A top us also 
possesses two large glands opening oue at each side of the 
mouth; and Simroth/ who first discovered them, has suggested 
that their function may be similar to that of the slime-glands 
of Peripatus. We have already seen that the pedal gland 
is developed to an unusual extent among the carnivorous 
genera. Now this gland also opens close to the mouth, and I 
would suggest that its secretion may play some part in over- 
powering the prey. The form of the pedal gland in Apera 
has come to resemble in some respects that of Simroth^s 
glands in Atopus, and nearly twent}^ years ago Andre ^ 
suggested that the secretion of the pedal gland in the 
Sty lommatophora might be used in feeding as well as in 
locomotion. We may at least say this : that the remarkable 
development of the pedal gland in the Agnatha, as well as 
the presence of the labial feelers, may not improbably be due 
to their predaceous habits. 

It is unnecessary to attempt to prove that the acquisition 
of carnivorous habits might lead to a modification of the 
radula and the surrounding structures: obviously these would 
be among the first organs to be aifected. It will be sufficient, 
therefore, if I point out the chief ways in which the radula 
has become transformed. In the first place, it has inci-eased 
in size, and especially in length, and become capable of far 
greater- protrusion than in the herbivorous forms. Secondly, 
the individual teeth have become much larger, especially 
those occupying an analogous position in the radula to that 
held by the canine teeth in the jaw of the Mammalia; and to 
make room for these, the central teeth and those at the 
extreme edges of the radula have become vestigial or have 

' ' Naturwiss. Woelieiischr..' 1901, vol. xvii, p. 122. 
^ • Revue Suisse cle Zoologie." vol. ii, p. 332. 



THE CARNIVOEOUS SLUGS OF SOUTH AFRICA. 229 

entirely disappeared. Thirdly, the main cusps of the teeth 
have become far longer and more sharply pointed, so as to 
penetrate the victim's skin, and the secondary cusps have 
completely gone, excepting in some of the more primi- 
tive genera of the Oleacinidse. And as all the teeth have 
become simplified in this manner, the differentiation between 
laterals and marginals has been lost, though it has been re- 
acquired in most of the species of Apera and Natal in a, and 
to some extent in Guestieria. Fourthly, the bases have also 
become lengthened in order to prevent the teeth from being 
bent outwards. Fifthly, the rows of teeth have become 
sharply angled in the centre, owing to the way in which the 
radula is forcibly drawn back over the anterior edge of the 
odontophoral support into the narrow I'adula-sac. 

The muscles of the odontophore have become correspond- 
ingly enlarged. In the more primitive forms the hind end of 
the radula-sac still projects from between these muscles ; but 
in those that are more completely adapted to a carnivorous 
existence the odontophoral muscles have entirely surrounded 
the radula-sac, and form a very large cylindrical structure. 

This growth of the odontophore leaves little room for the 
development of other bulky organs in the anterior part of the 
body, especially as it is an advantage to a carnivorous form 
not to have a very large head and neck. I therefore suggest 
that this is the reason why the genital ducts of carnivorous 
snails and slugs are without large accessory organs. There 
Avould be no room for the development of a dart-sac, for 
example, in a snail with a very large odontophore. 

The growth of the odontophore would also tend to increase 
the size of the body -whorl of the shell. If this were laterally 
flattened as in the Oleacinidae, it is evident that when the 
snail withdrew into its shell, the hind end of the odontophoi'e 
would come to press against the outer side of the shell in the 
neighbourhood of the posterior end of the mantle-cavity. 
The result of this would be that the posterior limit of the 
cavity would be pushed forwards in the middle, and with it 
the heart and the lower end of the kidney. The upper end 



230 HUGH WATSON. 

of the kidney, being near tlie suture of the shell, would I'einain 
in its original position ; and accordingly we find that in the 
Oleacinidae the kidney is obliquely lengthened in a very 
characteristic manner. 

The cerebral, buccal, and ventral ganglia would be pushed 
further apart by the growth of the buccal mass and odonto- 
phore, and thus we find that in many of the carnivorous forms 
the cerebro-buccal, cei'ebro-pedal, and cerebro-pleural con- 
nectives are unusually long (see pp. 141, 146). 

The extrinsic buccal retractors would become strongly 
developed at the same time as the intrinsic muscles of the 
odontophore. So long as they sprang fi'om the columellar 
muscle, they would undergo little change beyond an increase 
in their thickness, although I have found that their anterior 
ends tend to split up into a number of separate strands in the 
Rhytididte. But in those forms in which the shell is 
degenerate and the skin is taking its place, we frequently find 
that the retractors of the odontophore arise from a large area 
of the integument; as, for example, in Apera gibbon si, 
Testacella haliotidea Drajy., and the Trigonochla- 
minse (see pp. 172, 173). 

A jaw becomes superfluous in a snail or slug in which the 
radula is protruded far beyond it, and Simroth ^ has shown 
that it w^ould be a positive disadvantage in a vermivorous 
form, because if it bit off pieces from its prey, the latter 
would escape. Accordingly the jaw is absent or extremely 
degenerate in iiearly all carnivorous genera excepting 
Plutonia, in which Simroth states that the sharp edge is 
covered by a softer downward growth. 

As in other carnivorous animals, the digestive region of the 
alimentary canal becomes reduced in size. This is most 
apparent in the slugs, for in these the pressure of the organs 
which formerly occupied the visceral hump tends to diminish 
anything in the body-cavity that is unnecessarily large. In 
A top us and Apera the true stomach has almost completely 
disappeared, the crop passing straight into the intestine ; and 
1 Op. cit.. p. 113. 



THE CARNIVOROUS SLUGS OF SOUTH AFKICA. 231 

in these forms, and possibly in others also, digestion takes 
place partially within the lobes of the liver. In neai'ly all 
the carnivorous genera the length of the intestine and rectum 
is more or less reduced. 

Finally, Simroth considers certain peculiar connections 
between the genital ducts of a few of the carnivorous genera 
to be modifications to facilitate self-fertilisation, and he attri- 
butes this to the somewhat isolated life which a vermivorous 
slug is likely to lead. But I have not found these modifica- 
tions in the forms which 1 have examined, and it remains to 
be proved whether they are at all general among the 
A g n a t h a . 

Such are the changes which are likely to take place in the 
organisation of a snail when it acquires carnivorous habits. 
We are now in a position to discuss whether Apera, Testa- 
cell a, and other predaceous forms, are really closely related 
to one another, or whether their resemblance is merely due 
to the fact that thev have all become carnivorous. 



The Phylogeny of Apera and other Naked Carnivorous 

Slugs. 

I think that Simroth is the only author who has suggested 
that the genus Apera has been evolved directly from an 
herbivorous slug. As already mentioned, he put forward the 
theory that Apera might have been derived from the 
Janellidas. But Plate^ has shown that the slugs belonging 
to this family differ from most of the Stylommatoph ora, 
not only in being without lower tentacles, but in other 
important characters, such as the respiratory tissue, which 
takes the form of numerous fine tubes radiating from the 
mantle-cavity and recalling the trachea of arthropods. Now 
Apera possesses none of these features. Moreover, it Avould 
be difficult to imagine a radula more unlikely to develop 
carnivorous characters than that of Jan el la, with its 

' ' Zool. Jahvb.,' 1898, vol. xi, pp. 193-280, pis. xii-xvii. 









232 HUGH AVATSON. 

myriads of minute multicuspid teeth. I have therefore no 
hesitation in saying that Apera is in no way allied to the 
Janellidae. 

Nor do I know of any other herbivorous family from which 
Apera is at all likely to have been derived. And the fact 
that the carnivorous characters are so highly developed in 
Apera is against the theory that the genus has been directly 
evolved from any herbivorous form. It seems cei'tain that 
the ancestors of Apera must have been carnivorous for a 
very long time ; and it is not likely that these ancestors 
would all die out without leaving any descendants excepting 
this single genus. The question is whether we can find any 
carnivorous genus of slugs or snails resembling Apera in 
characters which are not likely to have been developed 
independently through the common acquisition of predaceous 
habits. 

Collinge has already pointed out how improbable is the 
theory of P. and F. Sarasin that Apera is closely allied to 
Atopus.^ This genus and the other members of the 
Rathouisiidge differ widely from Apera and every other 
carnivorous form in a number of important characters, such 
as the wide separation of the male and female openings, the 
presence of Sim roth's glands, the structure of the foot, the 
very large mantle, and the structure of the liver ; while they 
only resemble them in the radula, the absence of a jaw, and a 
few other points connected with their carnivorous habits. I 
agree with Simroth in regarding the Rathonisiidas as 
being more nearly related to the Veronicellidfe than to 
any monotrematous carnivorous family ; indeed, I have little 
doubt that, with the exception of the Veronicellidas and 
the Onchidiidse, no family of the Stylommatophora is 
less closely related to Apera than the Rathouisiidee. 

Plutonia, a carnivorous slug found in the Azores, is 

perhaps less unlike Apera than is Atopus; but it differs 

from it in the mantle, the latei'ally compressed form of the 

body, the presence of a jaw, the absence of a penial retractor, 

^ For references, see pp. 111-113. 



THE CARNIVOROL^S SLUGS OF SOUTH AFBICA. 233 

the presence of a secondary ureter, and other characters.^ 
On the whole it is extremely improbable that Apera is 
related to Plutonia. It is possible that the latter genus 
may be allied to the Trigonochlamin^e, but Simroth has 
given weighty reasons in favour of its having been evolved 
fi'om the species of the Vitrininfe inhabiting the shoi'es of 
the Atlantic, and not from any carnivorous group. 

'J'he only other carnivorous forms without external shells 
which are known to science are the genera from Trans- 
caucasia placed in the Trigonochl amiuEe. One of these — 
Selenochl amy s — bears a very striking resemblance to 
Apera, as will be seen from Simroth's figures.^ Almost the 
only external differences between Selenochlamy s and 
Apera are that in the former genus the visible mantle is a 
little larger, the foot is more distinctly tripartite, and there is a 
median dorsal keel extending from the mantle to the hind end 
of the animal. This last difference is the most conspicuous, 
and yet it is not greater than the difference between Apera 
sexangula and A. gibbonsi, dimidia, or purcelli. 
And when we turn to Simroth's description and figures of the 
internal anatomy of Selenochlamys, we at once notice the 
similarity between the radial buccal retractors in this genus 
and those of Apera gibbonsi and A. parva. A closer 
inspection, however, reveals many differences. The nerve 
ganglia of Selenochlamys are all separate ; the pedal gland 
is narrow, straight, and almost entirely embedded in the foot ; 
the vagina is extremely short ; the right tentacular retractor 
does not cross the penis. Moreover there appears to be no shell, 
so that the morphology of the mantle must be quite different 
to that of Apera. Indeed, I think that there can be little 
doubt that we have in the superficial resemblance between 
Apera and Selenochlamys a remarkable instance of con- 
vergence due to the acquisition of similar carnivorous habits ; 
and that the only affinities of Selenochlamys are with the 

* Simroth, H., ' Nova Acta Acad. Caes. Leop. -Carol. Germ. Nat. Ciir.,' 
1891, vol. Ivi, pp. 223-229. 
2 ' Festschrift Leuckarts,' 1892, pi. vi. 



234 HUGH WATSON. 

other members of the Trig'onochlaminas, which Simroth 
has shown to have probably been derived from the Par ma - 
cellinte. This view is confirmed by the geographical 
distribution of the genera. 

We may now consider the carnivorous genera which still 
retain an external shell, for it seems certain that the shell of 
Apera was originally external. 

The New Zealand genus Schizog'lossa resembles Apera 
in mam^ ways, but it differs in its reproductive system, for in 
Schizoglossa the male organs are much reduced, and there 
is no receptaculum seminis.^ These features alone render it 
improbable that Apera has been derived from Schizo- 
glossa. According to Murdoch - the reproductive organs of 
Paryphanta busbyi {Gray) bear a considerable resem- 
blance to those of Schizoglossa, and 1 think that there can 
be little doubt that the latter genus has been evolved in New 
Zealand from that section of the Rhytidida3 to which 
Paryphanta busbyi belongs. 

Strebelia possesses a receptaculum semiuis, but in this 
genus the penial retractor is attached to the vas deferens 
instead of to the penis itself. And, judging from Strebel's 
figures, the salivary glands iire united below the crop, and 
not above it as in Apera.^ Moreover, it does not seem likely 
that a purely South African genus should have been evolved 
from a slug which is only found in Mexico. 

There remain the European carnivorous slugs, Da u de- 
li ardia and Testacella. Of these Daudebardia differs 
widely from Apera in its pedal gland, nervous system, 
reproductive organs, excretory system, etc. There can be no 
doubt that Apera has not been evolved from Daudebardia. 
On the other hand, Testacella has man}'^ points in common 

• Hedley, C, 'Proc. Linn. Soc. N.S.W.' (■2nd ser.), 1893, vol. vii, 
p. 390, pi. ix, fig. 4, pi. X, fig. 9; Collinge, W. E., 'Ann. Mag. Nat. 
Hist.' (7tli ser.), 1901, vol. vii, p. 72, pi. ii, fig. 30. 

2 ' Trans. N. Z. Inst.,' 1903, vol. xxxv, pp. 260. 261, pi. xxvii, figs. 4. 5. 

' ' Beitrag. z. Kenntn. d. Fauna Mexikan. L.-u. Silsswasser-Conchy- 
lien,' 1878. vol. ill, pis. i. ii. 



THE CABNIYOU'OUS SLUGS OF SOUTH AFRICA. 235 

with Apera; indeed, if we take into consideration both the 
internal and external characters, I do not think that any- 
other genus resembles Apera so closely as does Testacella. 
The pedal gland of Testacella lies free in the body-cavity 
as it does in Apera. Both genera are opisthopnemnic, and 
in both the primary ureter opens at the posterior end of the 
mantle-cavity. The reproductive organs of the two genera 
are on the whole very similar, and in both the right tentacular 
retractor crosses the penis. But if we compare the genera 
more closely we find that there are several small differences 
which it is not easy to explain away. In the first place, the 
auricle in Testacella has come to lie directly behind the 
ventricle, whereas in Apera the heart has not been rotated 
so far. Yet Testacella cannot be derived from Apera, 
because it still retains an external shell. Secondly, the foot 
of Testacella contains numerous dermal mucous glands, 
while that of Apera has none. Thirdly, the pedal gland of 
Testacella has no terminal vesicle. Fourthly, the left 
parietal ganglion has not become fused with the abdominal 
ganglion in Testacella, but all the visceral ganglia remain 
separate. Fifthly, the arrangement of the odontophoral 
muscles, and the structure of the odontophoral support, and 
even the blood-supply of the odontophore, show surprising 
differences in the two genera, as we have already seen. And, 
lastly J there is the difference in the geographical distribution 
of the two genera. On the other hand, the resemblances are 
found to be largely of a negative character, if we except 
those which might be due to the common acquisition of 
vermivorous habits by both forms. Therefore it will be well 
to look further before we assume that Apera is related to 
Testacella. It is true that there are no other carnivorous 
genera in which the shell has become reduced, but it is 
possible that the nearest living allies of Apera may have 
quite large shells, for when once the presence of a shell is 
found to be a disadvantage, its degeneration probably takes 
place rather rapidly. 

Pilsbry has suggested that Apera may possibly be allied 
VOL. 3, PART 2. 17 



236 HUriH WATSON. 

to the Rhytididte, ar.d I am inclined to believe thatPilsbry 
is right. Beutler has examined the histology of the skin of 
Paryphanta, and has found that the foot is without dermal 
mucous glands, as it is in Apera. The pedal gland in the 
Rhytidida3 not only lies freely in the body-cavity, but it 
is often somewhat contorted, and, as in Apera, it ends 
in a vesicle containing a broad fold. In Parj-phanta, 
Natalina, and Rhytida capillacea {Fn\), the visceral 
ganglia are all separate, as in Testacella, but in Rhytida 
inaequalis {Pfr.) Fischer has shown that the left parietal 
ganglion is united with the abdominal ganglion exactly as in 
Apera. I have already shown that the arrangement of the 
odontophoral muscles in at least one member of the 
Rhytididse is very similar to that found in Apera, and, 
judging from Beutler's account, the structure of the odonto- 
phoral support is also similar. The reproductive organs of 
Rhytida and Paryphanta are on the whole very like 
those of Apera, and in these genera the right tentacular 
retractor crosses the penis. The members of the Rhytidid^ 
also resemble Apera in having no secondaiy ureter. Indeed, 
almost the only differences that I know of between Rhytida 
and Apera are those which would be likely to be bi'ought 
about by the degeneration of the shell and its further 
retrogression to the hind end of the animal, and we have seen 
that this is a modification which is especially liable to occur 
in carnivorous forms. 

Another fact in favour of the theory that Apera has 
been evolved from the Rhytidid^e is to be found in the 
geographical distribution of that family, for it occurs not only 
in the Australian region, but also in South Africa itself. It 
is true that Apera has almost certainly not been evolved 
from those members of the Rhytidida3 which are now found 
in South Africa : these have retained their separate visceral 
ganglia, but have become specialised in another direction, as 
is shown both by theii- radula and their reproductive system, 
in which the penis lies to the right of the tentacular retractors. 
But the presence of these snails in Africa renders it not at all 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 237 

improbable that forms more like Rhytida intequalis may 
also have ooce extended into that region, and given inse, by 
the degeneration of the shell, to Apera, just as in New- 
Zealand another branch of the family has probably given rise 
to Schizoglossa. 

It is not unlikely that the Rhytidid^ originated in early 
Mesozoic times, or perhaps even before the end of the Pala3o- 
zoic era, in Grondvvanaland — that great Southern continent 
which is supposed to have extended from Australia and New 
Zealand across the Indian Ocean, through Africa, and even 
as far as South America.^ A little later the more highly 
specialised Streptaxidae may have arisen in the same 
region. These did not reach Australia, perhaps because it 
was alread}^ cut off by the sea, but in other regions we may 
suppose that they would enter into competition with the 
Rhytidid^. In South Africa, where only the small pupi- 
form Streptaxidae occur, the Rhytidid^ were able to 
withstand their competition by becoming more specialised 
themselves, either by the degeneration of the shell (Apera) , 
or merely by an increase in the size of their teeth and a slight 
modification of their reproductive organs (Natalina). But 
further north, where we find the heliciform Streptaxidae, 
with their oblique columella, the Rhytididas were almost 
entirely exterminated, only leaving Natalina morrumba- 
lensis [M&P.), N. permembranacea Preston, and pos- 
sibly the species of Tayloria,^ as relics of their former dis- 
tribution. Whether the Rhytididfe ever reached South 
America is at present unknown. They may have done so, 
and have then been exterminated by the heliciform Strep- 
taxidae, which are not uncommon in South America. I 

' Hedley has suggested that the Rhytididse are of Antarctic origin 
C Proc. Linn. Soc. N.S.W.,' 1899, vol. xxiv, p. 398), but, while admitting 
the possibihty of this theory, I agree with Pilsbry in being unable to 
find any evidence in its favour (' Rep. Princeton Univ. Exped. Pata- 
gonia. 1896-1899,' vol. iii, Zool.. 1911. p. 631). 

•■= Thiele, J., 'Deutsch. Zentral-Afrika Exped. 1907-8,' vol. iii, 1912, 
p. 187. 



238 HUGH WATSON. 

would suggest, however, that it is quite possible that some of 
the South American carnivorous genera such as Guestieria, 
which Kobelt places in the Streptaxidte, may pi-ove to 
belong to the Rhytididge when their anatomy has been 
examined. 

Even if this view of the phylogeny of A per a is accepted, 
I would not advocate the placing of the genus in the Rhy- 
tididee. The gap which separates Apera from any known 
member of that family is a very wide one, and the isolation 
of the genus is by no means over-emphasized by placing it in 
a family by itself. 



The Phylogeny of Testacella and its Possible Allies. 

Most modern malacologists are agreed that Testacella is 
allied to Daudebardia, and that these genera have been 
derived fi'om Hyalinia or some closely related form. Now 
it must be admitted that the resemblance between Daude- 
bardia and Hyalinia is very striking. Whether we regard 
the nervous system or the i-eproductive system or the excre- 
tory system, the similarity is equally remarkable. Even in 
the digestive system the diiference is not very great, for 
Hyalinia is frequently carnivorous, and most of its teeth 
have become thorn-shaped, while Daudebardia still retains 
a small jaw, and the odontophoral muscles in this genus do 
not completely surround the radula-sac. Moi'eover, the evolu- 
tion of Daudebardia from Hyalinia is to a great extent 
recapitulated in development, young specimens of Daude- 
bardia having a shell very like that of Hyalinia, into which 
the animal can withdraw itself.^ In my opinion the evidence 
of Daudebardia alone is almost sufficient to prove that the 
carnivorous snails and slugs are not monophyletic; for I think 
that we must admit that Daudebardia has been evolved 
from Hyalinia or some closely allied form, and I do not 

' See Simroth, H., ' Nova Acta Acad. Cses. Leop. -Carol. Germ. Nat. 
Cur.,' 1891, vol. Ivi, p. 270. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 239 

suppose that anyone would maintain that all the carnivorous 
snails and slugs, including" such genera as Atopus and 
Varicella, had been derived from this source. 

But while I agree that Daudebardia has probably arisen 
from the Zonitidse, I canuot admit that Testacella is 
closely allied to Daudebardia and has had a similar origin. 

Testacella differs widely from Daudebardia in many 
respects. In Testacella the apex of the shell is directed 
backwards, the columella being parallel to its greatest length 
(PI. XXIV, fig. 161) ; in Daudebardia the columella is 
nearly at right angles to the length of the animal and the major 
diameter of the shell, as in Hyalinia. In Testacella the 
pedal gland lies freely in the body-cavity, and is very different 
in structure from that of most snails and slugs, a fact that has 
been specially emphasized by Andre ^; in Daudebardia the 
pedal gland is embedded in the foot, and Plate- has shown 
that it has the usual structure, the duct having two ventral 
longitudinal folds enclosing a furrow into which the gland- cells 
open. In Testacella the epidermal cells have their walls 
thickened on the outer side only; in Daudebardia they are 
thickened all round. ^ Testacella has no jaw; in Daude- 
bardia a jaw is present. In Testacella the radula-sac does 
not extend nearly to the hind end of the odontophore (PI. XXIV, 
fig. 156) ; in Daudebardia it projects beyond the odonto- 
phoral muscles. In Testacella the visceral ganglia are all 
separate from one another; in Daudebardia the abdominal 
ganglion is united with the right parietal ganglion, as is the 
case to some extent in Hyalinia also. In Testacella the 
auricle is behind the ventricle ; in Daudebardia the auricle 
is in front of the ventricle, as in Hyalinia. In Testacella 
the heart is to the right of the kidney ; in Daudebardia and 
Hyalinia it is to the left. Testacella has no secondary 
ureter; Daudebardia agrees with Hyalinia in possessing 
one. In Testacella the receptacular duct is moderately long; 

' ' Revvie Suisse de Zoologie/ 1894, vol. ii, pp. 318-321. 
- ' Zool. Jahrb.,' 1891, vol. iv, p. 524, pi. xxxii, fig. 16. 
=* Ibid.,pp. 527, 529. 



240 HUGH WATSON. 

in Dau debar dia it is extremely short. In Testae el la the 
vagina is not surrounded by any glandular tissue ; in Dau de- 
bar dia the anterior end of the vagina is encircled with 
glandular tissue, as in Hyalinia. In Testacella the penis 
passes between the right tentacular retractors; in Daude- 
bardia, as in Hyalinia, it lies outside both retractors. In 
Testacella the genital opening is on the side of the head; 
in Dau debar dia and Hyalinia the opening is further 
back. In short, the two genera only resemble each other 
in a few features which would be likely to be developed in 
any vermivorous form, and they differ in nearly every other 
respect. 

The dissimilarity in the nervous system is alone almost 
sufficient to prove that the carnivorous characters of Testa- 
cella have been acquired independently of those of Daude- 
bardia. It is a general rule in the animal kingdom that 
nerve-ganglia tend to unite and not to separate; therefore we 
cannot derive Testacella, with its distinctly separate 
abdominal and right parietal ganglia, from forms in which 
these ganglia are more or less united ; and this is the case in 
Hyalinia as well as in Daudebardia, notwithstanding 
Plate's statement to the contrary. 

Now, if Testacella is not allied to Daudebardia, there 
is no reason for supposing it to have been derived from the 
Zonitidse. This has only been thought to be the case 
because Daudebardia has almost certainly been evolved from 
that family, and Testacella was supposed to be related to 
Daudebardia. In Testacella, as in Apera, the carni- 
vorous characters have reached a very high state of specialisa- 
tion, and it is therefore more probable that Testacella has 
been evolved from some family of carnivorous snails. 

Beutler^ considers that Paryphanta may be ancestral 
to Testacella. Now Paryphanta certainly resembles 
Testacella much more closely than does Daudebardia. 
Indeed in its visceral ganglia it is more like Testacella 
than Apera. But Testacella dijffers from Paryphanta, 
' ' Zool. Jiilirl)..' 1901, vol. xiv, p. 407. 



THE CARNIVOEOUS SLUGS OF SOUTH AFRICA. 241 

just as it differs from Apera, in the pedal gland having- no 
terminal enlargement, and in the presence of longitudinal 
fibres in the odontophoral support and of dermal glands in the 
foot. And if a shell of the shape of that of Paryphanta 
degenerated, it would assume a form like that of Schizo- 
glossa, and not like that of Testacella. Moreover, while 
it is easy to attach too much importance to geographical 
distribution, it does not seem probable that a w^estern Palas- 
arctic genus of slugs should have been evolved from a snail 
found in New Zealand. 

But is there no family of carnivorous snails inhabiting 
Europe, in which the columella is parallel to the greatest 
length of the shell, and the odontophoral support contains 
longitudinal fibres? The Oleacinid^e possess these charac- 
ters, and in many other ways bear a close resemblance to 
Testacella, and it is from the Oleacinida? that I believe 
Testacella has been evolved. 

Although the majority of the recent species of the Olea- 
cinidfe are found in Central America and the West Indies, 
in Tertiary times the family was represented by many forms 
in Western and Central Europe, and one genus — Poire tia — 
still lingers in the Mediterranean region. We knoAV that the 
shell is liable to degenerate and recede to the posterior end 
of the animal in the Oleacinidte as in other carnivorous 
families, for it has done so in Strebelia. If the degenera- 
tion were to proceed further than it has done in this American 
genus, the shell would assume the form found in Testacella 
m a u ge i . This is seen from the parallel case of the degenera- 
tion of a shell with a pointed spire found in the Succineidffi, 
which is well illusti-ated on plate Ixxiii of H. and A. Adams' 
' Genera of Kecent MoUusca.' Further, we find that the 
abrupt truncation cf the columella which is characteristic of 
Poiretia, Euglandina, Oleacina, etc., occurs also in 
Testacella scutulum Soic. It is significant also that as 
early as the Eocene period the aperture in some of the 
European Oleaci nidge had already become as large as that 
of any of the recent American members of the family 



242 HUGH WATSON. 

excepting Strebelia^; for as a general rule, the lai'ger tlie 
size of the aperture, the smaller is the size of the shell as a 
a whole in proportion to that of the animal. 

It is not only in the shell, however, that the Oleacinidae 
resemble Testacella. In both, a deep cleft in the mantle- 
edge extends forwards from the respiratory opening beneath 
the right lip of the shell ; and Simroth has found that in 
Poiretia this cleft contains an olfactory organ, resembling 
the similarly situated olfactory organ of Testacella. 
Poiretia also has small labial feelers, very like those of 
Testacella. So far as I am aware, the histology of the 
foot and pedal gland in the Oleacinidfe has not been 
described ; but, judging from Strebel's figures," the pedal 
gland lies freely in tlie body-cavity in the more highly 
specialised members of the family, just as it does in Testa- 
cella. A jaw is absent in nearly all the Oleacinidse, as in 
Testacella. We have already seen that the odontophoral 
muscles in the Oleacinidas are tnore like those of Testa- 
cella than are the muscles of any other form which has been 
examined; and I have found that in Euglandina the struc- 
ture of the odontophoral support agrees exactly with that of 
Testacella. In most of the Oleacinidte the salivary 
glands are united, but according to Reymond ^ they are 
separate in the European genus Poiretia, as in Testacella. 
This, however, is not a very important character, for, as we 
have seen, the glands may be separate or united in different 
species of the same genus. The nervous system in the 
Oleacinidge is also identical with that of Testacella, all 
the visceral ganglia remaining separate, although they are 
sometimes veiy closely aggregated in Euglandina. The 
similarity of the reproductive system is equally striking. It 
is true that the receptacular duct of Testacella is shorter 

' See ' Jabresli. Yer. Natui'kunde Wiu'ttemb.,' 1907, vol. Ixiii, pi. ix, 
fig. 8. 

" 'Beitrag z. Keniitn. d. Faima Mexikan. L.-u. Siisswasser-Concbjlien,' 
1878, vol. iii. pi. xix. fig. 1. 

•' ' Joiu-ii. de Conchyl..' 1853, vol. iv, pp. 16-29, ■p\. i, fig. 1. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 243 

than in most genera of the Oleacinidffi excepting Stre- 
belia, but I have already shown that this is what we niiglit 
expect in a genus with a degenerate shell. The other female 
organs seem to be identical. The penis often ends in a caecal 
diverticulum in the Oleacinidfe, which, however, is quite 
short in Poiretia, and absent in Euglandina, Salasi- 
ella, Streptostyla, and Strebelia. In Testacella we 
frequently find a vestige of this diverticulum, especially in 
T. haliotidea DraiJ. In the Oleacinidse an epiphallus 
is usuall}^, though not invariably, inserted between the penis 
and the attachment of the penial retractor ; this is also 
the case in Testacella haliotidea, and sometimes in T. 
maugei (PI. XXIV, fig. 159). Pfeft'er has shown that in 
Euglandina liebnianni (Pfr.) the retractor springs from 
the apex of a flagellum, exactly as it does in Testacella 
haliotidea, and the same author has demonstrated that the 
internal structure of the penis of Euglandina is not 
unlike that of Testacella.^ Indeed, the similarity between 
Testacella and the Oleacinidfe seems to me to be even 
closer than the similarity between Apera and the Rhyti- 
d i d 83 . 

The fact that the heart in Testacella is on the right side 
of the kidney, with the auricle directly behind the ventricle, 
affords further evidence in favour of this view, as will be seen 
from the accompanying diagram. The usual position of the 
heart in Hyalinia, Rh^^tida, and similar genera, is shown 
in text-fig. 6, A — drawn from a specimen of Hyalinia 
d rap am audi [Beck). In Daudebardia, according to 
Plate, the heart is turned slightly to the right as shown in 
diagram B; but it will be seen that the auricle is still in front 
of the ventricle, and the kidney is still mainly on the right 
side of the heart, though the shell no longer extends far 
beyond these organs. In Apera the heart has rotated 
further in the same direction and is not protected by the 
shell (diagram c) ; the auricle has now come to lie further 
back than the ventricle, and the kidney is behind and to the 
' • Jahrb. d. Deiitsch. Mai. Gesell.,' 1878, vol. v, p. 81. 



244 



HUGH WATSON. 
Text-fig. 6. 









Diagrams illustrating the rotation of the heart in the evolution of 
carnivorous slugs from snails. 

A. Hyalinia, Rhytida, etc. B. Daudebardia. c. Apera. 
D. Euglandina, etc. E. Strebelia. F. Testacella 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 245 

left of the heart. Turning now to the Oleacinidge, we find 
that in Euglandina, Sti'eptostyla, etc., the heart already 
occupies a position similar to that which it holds in Apera, 
as is evident from diagram c' This is probably due partly 
to the pushing forwards of the ventricle owing to the pressui'e 
of the hinder portion of the odontophore/ and partly to the 
shape and consequent orientation of the shell.^ Diagram E 
shows the position of the heart and kidney in Strebelia, so 
far as I can judge from Strebel's figures, and it Avill be seen 
that with the reduction in the size of the shell and its retro- 
gression to the hind end of the animal, the heart has I'otated 
a little further. It is but a short step from this to the con- 
dition shown in diagram F, which illustrates the position of 
the heart in Testacella, Avitli the auricle directly behind the 
ventricle. Sixty 3' ears ago Reymond expressed the opinion 
that Testacella was "a Glandina with a rudimentary 
shell," and if we substitute the word " vestigial " for " rudi- 
mentary," I believe tliat Reymond expressed the truth. 

At the same time I am aware that the most eminent modern 
authorities have expressed a contrary opinion. Dr. Pilsbry ■* 
states emphatically that the relationships of the European 
carnivorous slugs (including Testacella) are with the 
Aulacopoda, and not with the other carnivorous families; 
and Simroth entertains the same view. The only facts which 
Pilsbry brings forward in support of this hypothesis are 
(1) that the European carnivorous slugs have lateral and 
pedal grooves, which he states that the other families do not 
possess, and (2) that the cerebral ganglia are generally united 
by a rather long commissure in the European slugs, while in 
the other families the cerebral ganglia are in close contact. 
Let us examine these points. 

In the first place, lateral grooves occur not only in Testa- 

' See also Strebel, op. cit.. vol. iii, pi. v, fig. 5. 

See p. 229. 
^ Cf. Naef, A., ' Ergebn. d. Fortschr. d. Zool.,' 1911, vol. iii, p. 131, 
fig. 19. 

^ ■ Manual of Conch.' (2nd ser.), 1908, vol. xix, p. viii. 



246 HUGH WATSON. 

cella^ Daudebardia, and tlie Trigonochlaminse, but some- 
times also ill the Oleacinidse, Rhytididte, and Aperida3. 
I have already described them in Apera, where they may 
be very conspicuous (PI. VII, figs. 11 and 13). Collinge^ 
shows them in his figures of Schizoglossa novoseelan- 
dica {Pfr.), and they are present on both sides in Natalina 
quekettiana [M. & P.), though rather ill-defined. In 
Euglandina venezuelensis {Preston) I have also found 
both lateral grooves, but the right groove is more conspicuous 
than the left. In these genera, however, the right lateral 
groove ends in the genital opening, whereas in Daude- 
bardia it passes below the genital opening and unites with 
the peripodial groove.- Now, Testae ella differs from 
Duudebardia in this respect, and agrees with the other 
families of carnivorous snails.^ 

In Daudebardia there is a deep peripodial groove, cutting 
off a broad foot-fringe, wdiich is cleft at the hind end by a 
short groove probably representing the lost mucous pore.* 
A peripodial groove is also present in the Aperidse and 
Streptaxida3, and although it is often absent or only slightly 
developed in the Rhytididse and Oleacinidte, Reymond 
states that it is fairly deep in Poiretia. In these forms, 
however, there is no trace of a caudal mucous pore. This is 
also the case in Testacella, which possesses a peripodial 
groove shallower than that of Daudebardia and nearer the 
edge of the foot.''' 

Lastly, Daudebardia resembles the Trigonochlaminte, 
Limacinge, and related groups, in having two conspicuous 
longitudinal grooves on the foot-sole, approximately parallel 
to each other. As a rule no such grooves are found in the 
other families of agnathous snails and slugs, but we have seen 
that occasionally similar grooves occur towards the hind end 

1 'Ami. Mag. Nat. Hist.' (7tli ser.), 1901, vol. vii, pi. ii, figs. 26, 27. 

2 Plate, L. H., • Zool. Jahrb.,' 1891, vol. iv, pi. xxxii, fig. 1. 

' See de Lacaze-Dutliiers, ' Arcli. Zool. Exper.,' 1887, vol. v, pi. xxx. 
fig. 11. 

■» Wiegmann, F., 'Mitt. Zool. Samml. Mus. Berlin.' 1898, vol. i, p. 62. 
* Compare Plate's figs. 3 and 11 on pi. xxxii, op. cit. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 247 

of the foot in Apera and Natalina ; they are, however, very 
inconstant, and instead of being parallel they diverge for- 
wards and seldom extend for moi*e than half the length of the 
animal. Authorities diifer with regard to the foot-sole of 
Testacella: Plate states that longitudinal grooves are 
present; Taylor^ says that they are absent ; Pfeffer- found 
them only in the anterior two-thirds of the foot-sole ; 
Simroth ^ saw traces of them towards the hind end of the foot 
in a few individuals only. I have examined the foot-sole in 
about a dozen examples of Testacella maugei from various 
localities, and I found that about half the specimens showed 
no trace of longitudinal grooves, Avhile in the others the 
grooves were present, but they were much less conspicuous 
than in Daudebardia, etc., and diverged forwards from the 
hind end as in Apera. In other words, the grooves on the 
foot -sole of Testacella, instead of being constant and 
approximately parallel, as in the other European carnivorous 
slugs, are divergent and very inconstant. We see, then, that 
the evidence of the various dermal grooves of Testacella is 
against Pilsbry's hypothesis instead of in favour of it. 

Turning now to the evidence aiforded by the length of the 
cerebral commissure, we find that while in Daudebardia 
and the Tr i go noch laminae the commissure is moderately 
long, in Testacella, as in the OleacinidiTe, Streptaxidas, 
Ehytididae, and Aperida?, the commissure is quite short, 
and the cei'ebral ganglia are in consequence close together. 
This fact was cleai'ly stated by Strebel'^ thirty-six years ago, 
and has been abundantly confirmed by subsequent investiga- 
tions. It is therefore evident that the only features upon 
which Pilsbry seems to base his theory of the affinities of 
Testacella are found on examination to support the contrary 
hypothesis. 

' ' Monog. L. and F.-W. Mollusca Brit. Isles,' 1902, vol. ii, p. 2. 
2 Op. cit., p. 75. 

•* ' Nova Acta Acad. Cses. Leop. -Carol. Germ. Nat. Cur..' 1891. vol. 
Ivi, p. 241. 

* Op. cit., vol. iii, p. 5. 



248 HUGH WATSON. 

If the views whicli I have brought forward are correct, all 
the carnivorous genera of slugs, with the exception of those 
belonging to the T r i g o n o c h 1 a m i n oe and to the Rathouisi- 
idte, have been derived independently fi'ora different snails 
by the pai-allel degeneration of the shell. This will be made 
clear from the following table : 

American Oleaoinidte — > Strebelia. 

European Oleacinidas •-> Testacella. 

Southern Hhytididee — > Schizoglossa . 

Northern Rhytidida3 - — — -^ Apera. 

European Zonitinfe — > Daudebardia. 

Atlantic Vitrininfe > Plutonia. 

Transcaucasian Parmacellinas > Trigonochlamina?. 

Oriental Ditremata > Rathouisiidse. 

But even if we admit that Apera has probably been evolved 
from the Rhytidid^eand Testacella from the Oleacinid as, 
the question of the affinities of these genera is not yet settled ; 
for Pilsbry^ states that the Oleacinidee and Rhytididas 
are closely related to each other, and if this be the case 
Apera and Testacella might still have been deriv^ed from 
a common predaceous ancestor, instead of their carnivorous 
characters having been acquired independently. 

Now it is true that the Oleacinidfe and the Rhytididie 
have many characters in common, but if we except those 
which would be likely to be caused by their carnivorous habits 
the remaining features are chiefly such as are possessed by 
nearly all the more primitive sigmurethrous Stylommato- 
phora. And there are at least two important differences 
between the families — their distribution and their shells. The 
difference in distribution, however, is less important than it 
appears, for, on the one hand, it is quite conceivable that the 
Oleacinidee may have once inhabited tropical Africa, and, 
on the other hand, we have already seen that the Rhytididge 
may occur in tropical America. But the difference in the shells 
cannot be so easily explained away. In the Oleacinidte the 
' Op. cit., vol. xix, p. xiii. 



TEE CARNIVOROUS SLUGS OF SOUTH AFRICA. 249 

shell is elongate, with a pointed spire and laterally compressed 
whorls, and the columella is truncated or folded. In the 
Rhytidid^e, on the other hand, the shell is lieliciforni or 
depressed, with a very obtuse spii'e and laterally expanded 
whorls, and the columella is neither truncated nor folded. I 
fail to see how it is possible to derive the shell of the 
Oleacinidte directly fi-om that of the Rhytididse or vice 
versa. If these families are really related, it can only be 
indirectly, and we must postulate a large number of inter- 
vening forms, with shells intermediate in shape. 

But it might be asked whether we do not find such a series 
of intermediate forms in the Streptaxid^e; and as a matter 
of fact in this family we have every gradation from heliciform 
shells such as Art em on and Imperturbatia to cylindrical 
shells like Ennea, and from these to pointed shells with 
laterally compressed Avhorls such as Streptostele and 
Obeliscella. May it not be that the Streptaxidge have 
been derived from the Rhytididse and the Oleacinidte 
from the Streptaxidfe ? In my opinion the Streptaxid^ 
may possibly have been derived from the Rhytidid^e, for 
I have recently dissected a form which proves to be in some 
ways intermediate between the two families. The great 
majority of the Strep taxidas, however, have come to differ 
widely from both the Rhytidida? and the 01 e acini die in 
their nervous system, their reproductive organs, and even in 
their radula ; and I think that there can be no doubt at all 
that the Oleacinidte have not been derived from that 
family. Therefore, if Pilsbry's view is correct, we must sup- 
pose that all the forms intermediate between the Rhytidida) 
and the Oleacinidte have died out completely, which does 
not seem a probable hypothesis. 

Dr. Simroth has suggested that the Oleacinidte may have 
been derived from the Achatinidse. Now I regard the 
striking resemblance between the shell of the American 
genus Euglandina and the African genus A ch a tin a as 
almost certainly due to convergence; because Euglandina, 
with its long labial papillae and closely aggregated nerve- 



250 



HUGH WATSON. 



ganglia, is probably one of the most recently evolved genera 
of the Oleacinidas. Indeed, its distribution suggests that it 
may not have arisen until after the final separation of the 
West Indian Islands from the mainland. I think that most 
zoologists who have studied the Oleacinidge will agree that 
the most primitive genera in the family are those with long 
spires, namely Spiraxis, Pseudosubulina, and Vari- 
cella. These genera are very unlike Achat in a, but they 

Text-fig. 7. 




2b 



J4. 



A. Curvella caloraplie Presiow, Brit. E. Africa. 
B. "Varicella nemorensis ^cZs., Jamaica. 

Representative teeth from the radula. x 400. 



are exceedingly similar to the more primitive members of 
the A cliatinidai, if we include the Stenogyrinas and 
Coeliaxin^e in that family. It is, in fact, almost impossible 
to say whether some groups of species should be placed in 
the Stenogyrinae or Oleacinid^e until their radula has 
been examined. I believe, therefore, that the Oleacinid^e 
have been derived from gnathophorous snails closely allied 
to the Stenogyrinai or even belonging to that subfamily. 

The chief differences between the internal au atomy of the 
Stenogyrinae and that of the OleacinidaD are to be found 
in the radula. We know, however, that the Stenogyrinas 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 251 

are sometimes predaceous/ and that tlieir teeth are liable to 
alter in form and acquire carnivorous characters. This is well 
shown in text-fig. 7, A, which represents part of the radula of 
an African member of the S tenogyri nse ; and Crosse and 
Fischer - have figured the radula of an American species — 
Leptinaria lamellata {Pot. & 3Iich.)— in which the outer 
lateral teeth have become similarly lengthened. Moreover, 
we find that the radu^la of the more primitive genera of the 
Oleacinidfe is not of the specialised caimivorous type found 
in the higher members of the family and in the Rhytididae. 
Thus in Varicella the radula is usually quite small, the rows 
of teeth are nearly straight, the bases of the teeth are broader 
and shorter than in the usual carnivorous type, and there is 
a small additional cusp outside the main cusp (text-fig. 7, b). 
This last character is especially significant, as the small cusp 
is obviously a vestigial ectocone, such as we find in so 
many herbivorous genera. I am indebted to the Rev. Prof. 
Gwatkin for kindly allowing me to examine the mounted 
radulfe of a large number of carnivorous forms, and I find 
that this additional cusp occurs in all the species of 
Varicella represented in his collection, namely, V. nemo- 
rensis Ads., phillipsi Ads., dissimilis Pilshry (= simi- 
lis Ads.,) and venusta Ads., although it is extremely 
minute in the last species. It is therefore very surprising 
that it has been entirely overlooked by previous observers, 
who have figured the radulse of V. phillipsi and V. 
nemorensis. Unfortunately I have not had an opportunity 
of studying the radula of Spiraxis, a genus which may be 
even more primitive than Varicella; but I have examined 
that of Pseudosubulina lirifera (ilfore^e^), and find that 
in this form the second cusp is also present and is larger than 
in Varicella, although in other ways the radula has become 
more highly specialised. Strebel has shown, however, that in 
Pseudosubulina there is a vestigial jaw, the structure of 
which is not unlike that of the jaw in the Achatinidse. 

' Johnson, C. W., ' Nautilus,' 1900, vol. xiii, p. 117. 
^ ' Mission scientifique an Mexiqiie,' pt. 7, 1877, pi. xxviii, figs. 8-10. 
VOL. 3, PART 2. 18 



252 HUGH WATSON. 

Ill my opinion these facts are sufficient to prove that the 
Oleacinidae have not been evolved from the Rhytididte, 
but have arisen directly from a gnathophoroiis family. 

Perhaps it might be said that the 01eacinid.Ee cannot 
have been evolved from the Achatinidae, because in the 
latter family the central teeth of the radula are almost 
invariably much smaller than the laterals, whereas iu the more 
primitive members of the Oleacinidae the central teeth are 
sometimes nearly as large as those on each side of them, not- 
withstanding that the teeth in the middle of the radula tend to 
diminish in size among carnivorous genera (cf. figs. 7, A and b). 
But I do not maintain that the Oleacinid^ have been 
evolved from any of the recent genera of the Achatinidae. 
It must be remembered that already in Upper Cretaceous and 
Eocene times the Oleacinidge were represented by forms 
which can hardly be regarded as primitive; the family, 
therefore, cannot have arisen very much later than the 
Jurassic period. Now the small central teeth are not an 
absolutely constant feature of the Achatinidee even at the 
present day, and it is not improbable that in Mesozoic times 
the radula was still of the more generalised type found in the 
closely allied family Megas piridfe. Indeed, it is possible 
that in the Jurassic period the Achatinidae may not yet 
have definitely separated from the Megaspiridae. Callio- 
nepion may perhaps be regarded as a descendant of a 
form intermediate between these two families, and in this 
genus the central teeth are nearly as large as the laterals. 
Moreover, the penis has a continuation in Callionepion, 
which reminds us of the similar structure found in Eug- 
landina liebmanni [Pfr.), and other members of the 
Oleacinidae.^ Lastly, the shells of the typical section of 
Spiraxis in some respects resemble the Megaspiridae 
quite as much as the Stenogyrinae, which also suggests 
that the Oleacinidte may have diverged from the Acha- 

' Pilsbry and Vanatta, ' Proc. Acad. Nat. Sci. Phila.,' 1899, pp. 371- 
373, pi. XV, figs. 3, 8, and Pilsbry, 'Man. of Conch.,' 1904, vol. xvi, p. 178, 
pi. xxxi, figs. 7, 8. 



THE CARNIVOROUS SLUGS 01*^ SOUTH AFRICA. 253 

ti nidge Avhen this family was scarcely distinct from the 
Megaspi ridge. Nevertheless, until further anatomical 
investigations have been made, it is impossible to say exactly 
to which group now living the 01eacinida3 are probably 
most nearly related. 

It is equally difficult to form an opinion as to the place 
of oi'igin of the family, for both the Ac ha tin idee and 
the Megaspiridge have an extremely wide distribution. 
Perhaps the Oleacinidse may have arisen near the southern 
or south-eastern shores of that continent which geologists 
believe to have stretched from Western Europe to America 
during a lai'ge part of the Mesozoic era. If this were the 
case, the south-eastern expansion of the area of distribution 
would be prevented by the extensive " Mediterranean Sea " 
which then existed ; the gradual growth of the North Atlantic 
would separate the European from the American forms, and 
the subsequent incoming of the Glacial Period would restrict 
the northern distribution of the family. On the other hand, 
the Oleaci nidge may possibly have arisen further south, 
and have entered America by way of the old land-connection 
which probably extended from Africa to Brazil. The absence 
of the Oleacinidas from the Ethiopian Region is a possible 
objection to this theory ; but perhaps the family may have 
once extended into that area, and may have there been 
exterminated by the elongate Streptaxida3 — Ennea, 
St reptostele, and their allies — which would probably enter 
into competition with them, but appear to have arisen just 
too late to follow the heliciforni Streptaxidse across to the 
Neotropical Region. The limited distribution of the Olea- 
cinid« in South America is, however, another objection to 
the theory of their southern origin, and the first view that I 
have given seems to me to be the more probable. lam there- 
fore glad to see that Pilsbry now upholds the more northerly 
origin of the family,^ although seven years ago he thought 
that the probabilities favoured the hypothesis that the Olea- 

1 ' Rep. Princeton Univ. Exped. Patagonia 1896-1899,' vol. ill, Zool., 
1911, p. 625. 



254 HUGH WATSON. 

cinidse arose in the Brazil- African continent.^ All this is 
highl_y problematical ; but the uncertainty which surrounds 
the precise origin of the Oleacinidfe does not affect the 
general conclusion that the family is probably more nearly 
related to the Achatinidae and their allies than to any of 
the other carnivorous forms excepting Testacella. 

It is not necessary for me to discuss at length the phylogeny 
of the Rhytididas; this family is evidently of very ancient 
origin — possibly it is the oldest of all the carnivorous families — 
and I do not suppose that anyone would maintain that it had 
arisen from the Oleacinidas. I hope to deal in greater 
detail with the affinities of the EhytididEe and Strep- 
taxi dte when treating of the South African members of 
these families ; and I have already said enough to show that in 
all probability the carnivorous characters of the Rhytididfe 
and Oleacinid^, and therefore of Apera and Testacella, 
have been acquired independently. The remarkable resem- 
blance between Apera and Testacella seems only to be 
another example of convergence due to the common acquisition 
of carnivorous habits. 

One thing is quite clear from the preceding argument : the 
tribe A gnat ha is not a natural group, and should therefore 
find no place in the classification of the Pulmonata. The 
Rathouisiida3 should be placed with the Veronicellidse 
among the Ditremata, as some authors have already done. 
The Trigonochlamina3, Plutoniinas, and Daude- 
bardiinae should be placed among the Aulacopoda or 
Oxygnatha, next to the Parmacellinge, Vitrininas, and 
Zonitinge. The Oleacinid^e and TestacellidaB should 
form a group by themselves, near the Achatinid^e and 
Megaspiridfe, if my views are correct; and to this small 
group Morch's term Agnatha may well be applied, for his 
original description is simply: "Agnatha. Ohne Kiefer: 
Oleacina, Testacella. "^ The Rhytidid^e, Aperidae, and 
probably the Streptaxidte, should be placed in another 

^ ' Man. of Concli.,' vol. xix, p. xiv. 
- ' Mai. Blatt..' 1859, vol. iv, p. 109. 



THi] CARNIVOROUS SLUGS OF SOUTH AFRICA. 



255 



group, for wliich we may adopt Pilsbry's term Agnatlio- 
niorphaJ Whether the purely American family Circi- 
iiariida3 should also be placed in this group I am unable to 
say. Pilsbry considers that it is allied to the Streptaxidas, 
but Simroth believes it to be related to the Zonitidfe. There 
are possible objections to both these views, but as I have not 
had an opportunity of examining any members of the Circi- 
nariidie myself, I will refrain from expressing an opinion 
upon its affinities. 

Thus the carnivorous snails and slugs should probably be 
classified as follows : 

JTestacellid^e. 
l01eacinida3. 



S i u" m u !• e t h r a - 



A g n a t h a 
Agnathomorp 



r Aperidas 
r p h a J 



A.ulacopoda 
or 
^ X y g n a t h a 



Rhytididas. 

Streptaxid^. 

Circinariida3. 
' Daudebardiina3. 

( Z o n i t i n a3 ) . 

Plutoniina;. 

(Vitrininte). 

T r i g o n o c h 1 a m i n a3, 

(ParmaceUince, etc.). 
Ditreinata ..... Rathouisiidee. 

While it is hoped that this classification rests on a firmer 
basis than previous attempts of a similar nature, it must be 
remembered that we still know very little of the comparative 
anatomy of the carnivorous suails and slugs. To give a single 
example: about a hundred species of the genus Ennea 
are known to occur in South Africa alone, and I believe that 
I am right in saying that not a single feature of the anatomy 
of any of these species has been described. As we are equally 
ignorant of the anatomy of many other carnivorous genera, it 
is at present impossible to do more than give a rough outline 
of their probable affinities. Let us hope that the time will 
soon come when collectors of shells will cease to throw away 
' ' Proc. Acad. Nat. Sci. Phila.,' 1900, p. 564. 



256 HUGH WATSON. 

the animals when cleaning their specimens, but will study 
their anatomy instead. 



APPENDIX. 
EEFERENCES TO TESTACELLA MAUGEI FEB. 

The literature on Testacella maugei is so extensive that 
it has been deemed more convenient to give the following list 
of some of the principal references to this species in the form 
of an appendix. Those references in brackets relate to fossil 
shells which have been assigned to T. maugei, although it 
is possible that some of them, such as T. asinina, may have 
belonged to species which, while nearly allied to this form, were 
really distinct from it. Most modern writers place the form 
found in New Zealand in this species, notwithstanding that 
Hutton stated that its radula differed slightly from the type 
usually found in T. maugei; I have therefore included the 
references to it in the following list, although I have not had 
an opportunity of examining a New Zealand example myself. 

Testacella haliotoides Lam.. Sys. An. s. Ver., 1801, p. 96; Wood- 
ward, Man. Moll., 1854, p. 169. fig. 94. 

Testacella haliotidea (pars) Drnp., Hist. Nat. Moll. Fr., 1805, pi. 
viii, figs. 46-48 ; Lowe. Rep. Brit. Assoc, 1883, p. 549. 

Testacella haliotidea var. scutuluni Moq.-Tand.,I{.ht. Moll. Fr., 
1855. pi. V, figs. 20, 21. 

Testacella maugei Fer., Hist. Moll., 1819, vol. ii, pp. 94, 95, pi. viii, 
figs. 10-12 ; Miller, Ann. Philos. (new ser.), 1822, vol. iii, p. 380 ; 
Sowerljy, Genera Shells, 1822, Testacella, figs. 7-10; Fleming, 
Hist. Brit. Anim.. 1828, p. 257 ; Desh., Diet. Class. d'Hist. Nat.. 
1830, vol. xvi, p. 179 ; J. D., Mag. Nat. Hist., 1833, vol. vi, p. 45, 
fig. 8, c, d ■ Lnkis and J. D.. Mag. Nat. Hist., 1834. vol. vii, pp. 225. 
229, figs. 40, c, d, 41, /, y ; d'Orhigny. Moll, des lies Canaries, 1834, 
p. 48 ; Gray, Turton's Man. L. and F. W. Shells, 1840. pi. iii. fig. 18 ; 
Encycl. Brit. (7th ed.), 1842, vol. xv, pi. ccclxvii, fig. 5 ; Penny 
Cyclopaedia, 1842, vol. xxiii, p. 246, figs, a, a, b ; Reeve, Conch. Syst., 
1842, vol. ii, pi. clxi, figs. 7-10 ; Morelet, Moll. Terr, et Fluv. Portu- 
gal, 1845, pp. 48, 49 ; Forbes and Hanley. Hist. Brit. Moll., 1853, 



THE CARNIVOKOUS Sl-UGS OF SOUTH AFRICA. 257 

vol. iv, p. 28 ; Albers, Malacographia Maderensis, 1854, pp. 13, 14, 
pi. i, figs. 9-11 ; H. & A. Adams, Gen. Rec. Moll., 1854, vol. iii, pi. 
Ixxii, figs. 7, 7 a ; Grat., Distr. Geogr. des Limaciens, 1855, pp. 15, 
16 ; Tapping, Zoologist, 1856, vol. xiv, p. 5102 ; Woodward, Man. 
Moll., 1856, p. 465, fig. 262; Gassies and Fisclier, Actes Soc. Linn. 
Bordeaux, 1856, vol. xxi, pp. 36-39, pi. i, figs. 2, 5 ; Grat., Distr. 
Geogr. Moll. Girond., 1858, p. 72; Morelet, L'Hist. Nat. A9ores, 1860, 
p. 143 ; Norman, Proc. Somerset Arch, and Nat. Hist. Soc, 1861, 
ex vol. x, pp. 11, 12 ; Bourg., Rev. and Mag. Zool. (2nd ser.), 1861 ; 
vol. xiii, p. 514; Jeffries, Brit. Conch.. 3862, vol. i, pp. 147, 148; 
Reeve. Brit. Moll., 1863, pp. 32, 33; Paiva, Mon. Moll. Ins. Mader., 
1867, pp. 6, 7 ; Massot, Ann. de Malac, 1870, vol. i, pp. 145, 146 ; 
Mousson, N. Mem. Soc. Helv. Sci. Nat., 1873, ex vol. xxv, p. 11 ; 
WoUaston, Testacea Atlantica, 1878, pp. 13, 14, 72, 73, 310, 311 ; 
Tryon. Man. Conch. (2nd ser.), 1885. vol. i, p. 8, pi. i, figs. 1-3; de 
Lacaze-Duthiers, Arch, de Zool. Exper. (2nd ser.), 1887, vol. v, pp. 
469. etc., pi. xxix. figs. 7, 8; PoUon., Boll. Miis. Zool. Anat. Torino, 
1889, vol. iv, No. 57, pi. i, fig. 1 ; Siniroth, Nova Acta Acad. Caes. 
Leop.-Carol. Germ. Nat. Cur., 1891, vol. Ivi, pp. 230-245, 266, etc., 
pi. X, figs. 1, 2, 5-7; Plate, Zool. Jahrb., 1891, vol. iv, pp. 518, 519, 
524. 542-552. 560-562. 566, 588, 594, 595, 614, 618-620, etc., pi. xxxiii, 
figs. 44, 46, 47, pi. xxxiv, figs. 62-65, pi. xxxvi, fig. 91, pi. xxxvii, 
figs. 101, 115, 116 ; Collinge, An. Mag. Nat. Hist., 1893, vol. xii, p. 24, 
pL i, fig. 3 ; Locard, Coquilles Ter. France, 1894, p. 18, figs. 5, 6 ; 
Webb, J. of Malac, 1895, vol. iv, p. 74, pi. ii, figs. 1, 4, pi. iii, figs. 
7, 8 ; L. E. Adams, Coll. Man. Brit. L. and F.-W. Shells (2nd ed.), 

1896. pp. 41, 42. pi. ii, fig. 3, pi. vii, fig. 12 ; Webb, J. of Malac, 

1897, vol. vi. pp. 49, 52, 56, pi. vi, fig. 3 ; Taylor. Mon. L. and F.-W. 
Moll. Brit. Is , 1902. vol. ii, pp. 21-27, figs. 32-42, pi. i. figs. 9-16. 
pi. iv ; L. E. Adams. J. of Conch.. 1911. vol. xiii. p. 212. 

[Testacella asininnm de Serves, Ann. Sci. Nat., 1827, vol. xi, p. 409.] 
[Testacella asinina de Sevres; Gassies and Fischer, Actes Soc. Linn. 

Bordeaux, 1856, vol. xxi, pp. 41, 42, pi. ii, fig. 3.] 
[Testacella lartetii Diipuy, J. de Conch., 1850, vol. i, pp. 302-304. 

pi. XV, figs. 2, a-(Z ; Gassies and Fischer, Actes Soc. Linn. Bordeaux, 

1856. vol. xxi, pp. 40-41. pi. ii. fig. 2.] 
[Testacella bruntoniana de Serves, Mem. Terr. Trausp., 1851, p. 51 ; 

Gassies and Fischer. Actes Soc. Linn. Bordeaux, 1856, vol. xxi, 

p. 42.] 
[Testacella deshayesii Mich., Desc. Coq. Foss., 1855, p. 3, pi. ii.figs. 

10, 11.] 
Testacella burdigalensis Gcnsies <£• Kavlin, Cat. Moll. Ter. and 

Flnv. Fr., 1855, p. 2. 
Testacella oceanica Grat., Distr. Geogr. des Limaciens, 1855, p. 15. 



258 HUGH WATSON. 

Testa cella canariensis Graf., Distr. Geogr. des Liuiaciens, 1855, 

p. 16. 
[Testacella monspessiilana Grat, Distr. Geogr. des Limaciens, 

1855, p. 16.] 
[Testacella browniana Gi-at., Distr. Geogr. des Limaciens, 1855, 

p. 16.] 
[Testacella oecitauige Grat. Distr. Geogr. des Limaciens, 1855, 

p. 16.] 
[Testacella altse-ripse Grat, Distr. Geogr. des Limaciens, 1855, 

p. 16.] 
[Testacella aquitanica Grat, Distr. Geogr. des Limaciens, 1855, 

p. 16.] 
[Testacella nouleti Bourg., Hist. Mai. Collinede Sansau, 1881. p. 15]. 
Daudebardia novoseelandica Hutton, Trans. N. Z. Inst.. 1881, vol. 

xiv, pp. 152, 153, idI. iii, fig. E ; pi. iv, fig. M. 
Testacella vagans Hutton, Trans. N. Z. Inst., 1882, vol. xv, \). 14U ; 

1883, vol. xvi, p. 209, pi. x, fig. T ; Try on, Man. Concli. (2nd ser.), 

1885, vol. i, pp. 11, 12 ; Musson, Proc. Linn. Soc. N. S. W., 1890, 

p. 885 ; Suter, Trans. N. Z. Inst., 1892, vol. xxiv, p. 279. 
Testacella aurigaster Layarcl ; Connolly. Ann. S. Afr. Mus., 1912, 

vol, xi, p. 64. 
According to Gassies and Fischer, and Simroth, Plectropborus 
orbignyi Ffr., 1819, is also probably founded on specimens of this 
species. 



EXPLANATION OF PLATES YII-XXIY, 

Illustrating Mr. Hugh Watson's paper on "The Carnivorous 
Slugs of South Africa," 

PLATE VII. 

Fig. I. — X I'l. Apera gibbonsi s.s.; dorsal view; probably 
fi'om Lower Umfolosi Drift, Zululand. 

Fig. 2. — x 1"1. A. gibbonsi s.s.; side view; Hlabisa, Zululand. 

Fig. 3.— X 11. A. gibbonsi rubella, young; side view; Equeefa, 
Natal. 

Fig. 4.— X 1-1. A. gibbonsi rubella; dorsal view; Equeefa, 
Natal. 

Fig. 5.— X 1-1. A. gibbonsi rubella; side view; Equeefa, Natal. 



THE CARNIVOROUS SLUGS OP SOUTH AFRICA. 259 

YiG. 6. — X I'l. A. gibbousi gracilis, type; side view ; Equeefa, 
Natal. 

Fig. 7.— X II. A. dimidia, type; dorsal view ; Equeefa, Natal. 

Fig. 8. — x 1"1. A. dimidia; side view ; Port Sliep stone, Natal. 

Fig. 9. — X I'l. A. dimidia; ventral view ; Eqiieefa, Natal. 

Fig. 10. — X 1"1. A. burnupi; dorsal view; Richmond, Natal. 

Fig. 11. — X 11. A. burnnj)i; side view; Chase Bush, Pieter- 
maritzburg. 

Fig. 12. — x 1"1. A. sexangula; dorsal view; Port Shepstone, 
Natal. 

Fig. 13.— X 1"1. A. sexangnla ; side view; Port Shepstone, 
Natal. 

PLATE VIII. 

Fig. 14.' — x 1. Apera gibbousi s. .•>•. ; dorsal view ; Zululand. 

Fig. 15. — X 1. A. gibbonsi s. s. ; side view; Zululand. 

Fig. 16. — x 1. A. gibbonsi rubella; dorsal view; Equeefa, 
Natal. 

Fig. 17. — X 1. A. gibbonsi rubella; side view; Equeefa, Natal. 

Fig. 18. — x 1. A. gibbonsi lupata; dorsal view; Port Shep- 
stone, Natal. 

Fig. 19. — x 1. A. gibbonsi lupata; side view; Port Shepstone, 
Natal. 

Fig. 20. — x 1. A. dimidia; dorsal view ; Equeefa, Natal. 

Fig. 21. — x 1. A. dimidia; side view ; Equeefa, Natal. 

Fig. 22. — X Ij. A. parva; dorsal view; near Fern Kloof , Grahams- 
town. 

Fig. 23. — x Ij. A. parva; side view ; near Fern Kloof, Grahams- 
town. 

Fig. 24. — x 3. A. dimidia; dorsal view of hind end; Equeefa. 
Natal. 

Fig. 25. — x S^. A.purcelli; dorsal view of hind end ; Table Mt., 
Cape Town. 

Fig. 26. — x 2. A. burnupi; dorsal view of hind end ; Chase Bush, 
Pietermaritzburg. 

' Figs. 14-23 show the colours of specimens preserved in alcohol. 



260 HUGH WATSON. 

PLATE IX. 

Fig. 27. — X 2A. Apera gibbonsi rubella; general dissection 
from above.' 

Fig. 28. — x 5. A. parva; general dissection from above. 
Fig. 29. — x 45. A.dimidia; general dissection fi-om above. 
Fig. 30. — x 2. A.burnupi; general dissection from above. 
Fig. 31. — x 2f. A. sexangula; general dissection from above. 

PLATE X. 

Fig. 32. — x 25. Apera gibbonsi rubella, young; transverse 
section through the region of the buccal retractors. - 

Fig. 33. — x 25. A. gibbonsi rubella, young; transverse section 
through the region of the anus. 

PLATE XI. 

Fig. 34. — x 25. Apera dimidia; transverse section through the 
region of the heart. 

Fig. 35. — x 25. A.dimidia: transverse section through the region 
of the reno-pericardial duct. 

PLATE XII. 

Fig. 36. — x 25. Apera dimidia; transverse section through the 
region of the anterior end of the ureter. 

Fig. 37. — X 128. A. dimidia; transverse section through the 
skin of the back. 

Fig. 38. — x 300. A. dimidia; transverse section through the 
skin of the foot-sole. 

Fig. 39. — X 118. A.dimidia; transverse section through the right 
corner of the lung. 

PLATE XIII. 

Fig. 40. — x 300. Apera dimidia; transverse section through a 
sphincter of a dennal blood-vessel. 

' The specimens shown in figs. 27-31 have been cut open a little to 
the left of the mid-dorsal line, and, excepting in A. burnupi, the 
digestive and i^eproductive organs have been slightly separated in order 
to show the course of the anterior aorta. 

- The sections rej)resented on Plates X to XII are shown as viewed 
from the front. 



THE GARNI VOEOUS SLUGS OF SOUTH AFRICA. 261 

Fig. 41. — x 3. A. gibbon si rubella; dorsal view of shell; 
Equeefa. 

Fig. 42. — x 3. A. gibboiisi rubella; side view of shell ; Durban. 

Fig. 43. — x 4. A. gibbonsi gracilis; dorsal view of shell. 

Fig. 44. — -X 6. A. parva; dorsal view of shell. 

Fig. 45. — x 6. A. parva; side view of shell. 

Fig. 46. — x 4. A. dimidia ; dorsal view of shell. 

Fig. 47. — x 7. A. purcelli; dorsal view of shell. 

Fig. 48. — x 7. A. purcelli; side view of sheU. 

Fig. 49. — x 2. A. burnupi ; dorsal view of shell. 

Fig. 50. — x 3. A. sexangula; dorsal view of shell. 

Fig. 51. — x 3. A. sexangula; side view of shell. 

Fig. 52. — X 3. A. gibbonsi ritbella; dorsal view of pedal gland, 
central nervous system, etc' 

Fig. 53. — x 8. A. j)arva; dorsal view of pedal gland, central 
nervous system, etc. 

Fig. 54. — x 6. A. dimidia; dorsal view of pedal gland, central 
nervous system, etc. 

Fig. 55. — x Sk. A. purcelli; dorsal view of pedal gland, central 
nervous system, etc. 

Fig. 56. — x 3. A. burnupi; dorsal view of pedal gland, central 
neiwous system, etc. 

Fig. 57. — X 5. A. sexangula; dorsal view of pedal gland, central 
nervous system, etc. 

Fig. 58. — X 6. A. gibbonsi lupata ; posterior end of pedal gland. 

Fig. 59. — X 46. A. dimidia; transverse section through glandular 
part of pedal gland. 

Fig. 60. — X 46. A. dimidia; transverse section through duct of 
pedal gland behind glandular part. 

Fig. 61. — x 46. A. dimidia; transverse section through anterior 
end of terminal vesicle of pedal gland. 

Fig. 62. — x 46. A. dimidia; transverse section through terminal 
vesicle of pedal gland, showing blood-vessel entering the fold. 

PLATE XIV. 

Fig. 63. — X 114. Apera gibbonsi rubella; transverse section 
through one side of pedal gland near its anterior end. 

^ In the specimens shown in figs. 52-57 the oesophagus, penis, etc. 
have been turned aside in order to display the underlying organs. 



262 HUGH WATSON. 

Fig. 64. — x 22. A. gibbonsi rubella; side view of part of pedal 
gland. 

Fig. 65. — x 119. A. gib])onsi rubella ; transverse section through 
fold in terminal vesicle of pedal gland. 

Fig. 66. — ^x 114. A. diuiidia; transverse section through fold in 
terminal vesicle of pedal gland. 

Fig. 67. — X 25. A. gibbonsi rubella; ventral grouj) of nerve 
ganglia. 

Fig. 68. — x 60. A. dimidia; section through pedal ganglia and 
pedal commissure, cut slightly obliquely, and showing pleural ganglion 
on the right and cerebro-pedal connective on the left. 

Fig. 69. — x 120. A. dimidia ; sections of nerve-cells in right 
parietal ganglion. 



PLATE XY. 

Fig. 70. — X 16. Apera gibbonsi rubella; cerebral and buccal 
ganglia. 

Fig. 71. — x 32. A. dimidia; cerebral and buccal ganglia. 

Fig. 72. — x 25. A.sexangula; cerebral and buccal ganglia. 

Fig. 73. — x 30. A. dimidia; left half of ventral group of ganglia. 

Fig. 74. — x 33. A. sexangula: right half of ventral group of 
ganglia. 

Fig. 75. — x 6. A. dimidia; semi-diagrammatic figure showing the 
distribution of the nerves from the ventral group of ganglia. 

Fig. 76. — X 5. A. sexangula; tentacular retractors of an ab- 
normal specimen. 

Fig. 77. — X 26. A. gibbonsi rubella; transver.se section through 
niovith. 

Fig. 78. — x 15. A. gibbonsi rubella; transverse section through 
crop and salivary glands. 

Fig. 79. — x 100. A. gibbonsi rubella ; transverse section through 
wall of crop. 

PLATE XVI. 

Fig. 80. — X 50. Apera dimidia; transverse section through re- 
tracted upper tentacle in front of eye. 

Fig. 81. — X 50. A. dimidia; transverse section through retracted 
upper tentacle, showing anterior part of eye. 



THE CARNIVOROUS SLUGS OF SOUTH AFRICA. 2()3 

Fig. 82. — X 50. A. dimidia; transverse section through retracted 
upper tentacle, showing posterior part of eye. 

Fig. 83. — x 50. A. dimidia; transverse section through retracted 
upper tentacle just behind eye. 

Fig. 84. — x 50. A. dimidia; transverse section through retracted 
upper tentacle behind eye. 

Fig. 85. — x 50. A. dimidia; transverse section through retracted 
upper tentacle considerably behind eye. 

Fig. 86. — X 50. A. dimidia; transverse section through retracted 
upper tentacle at entrance of olfactory and optic nerves. 

Fig. 87. — x 60. A. sexangula; longitudinal section through 
mouth. 

Fig. 88. — x 60. A. dimidia; transverse section through mouth. 

Fig. 89. — x 118. A. dimidia; transverse section through radula 
sac. 

Fig. 90.— X 118. A. dimidia; transverse section through part of 
odontophoral support. 

PLATE XVII. 

Fig. 91. — x 2|. Apera gibbonsi s.s.; side view of odontophore 
without its sheath. 

Fig. 92. — x 21. A. gibbonsi s.s.; longitudinal section of odon- 
tophore. 

Fig. 93. — x 2. A. gibbonsi rubella; doi-sal view of odontophore 
without its sheath. 

Fig. 94. — x 2. A. gibbonsi rubella; ventral view of odonto- 
phore without its sheath. 

Fig. 95. — X 2. A. gibbonsi rubella; dorsal view of odontophoral 
support. 

Fig. 96. — x 2. A. gibbonsi rubella; side view of radula with 
its retractor muscles. 

Fig. 97. — X 2. A. gibbonsi rubella; side view of radula without 
its muscles. 

Fig. 98. — X 2. A. gibbonsi rubella; side view of radula without 
its muscles (retracted). 

Fig. 99. — X 3. jA. gibbonsi lupata; side view of radida without 
its muscles. 

Fig. 100. — X 5. A. parva; dorsal view of odontophoral support. 
Fig. 101. — X 5. A. parva ; side view of radula with its retractor 
muscles. 



264 HUGH WATSON. 

Fig. 102. — x 5. A. parva; side view of radula without its muscles. 

Fig. 103. — x 6. A. dimidia; ventral view of odontophore. 

Fig. 104. — X 6. A. dimidia; side view of odontopliore without its 
sheath. 

Fig. 105. — X 6. A. dimidia; dorsal view of odontophoral support. 

Fig. 106. — x 6. A. dimidia; side view of radula with its retractor 
muscles. 

Fig. 107. — X 6. A. dimidia; side view of radula without its 
muscles. 

Fig. 108. — x 4. A. burnupi; side view of odontophoral support. 

Fig. 109. — X 4. A. burnupi; dorsal view of odontopliore without 
its sheath. 

Fig. 110. — X 6|. A. sexangula; venti-al view of odontopliore. 

Fig. 111. — X fii. A. sexangula; dorsal view of odontopliore with- 
out its sheath. 

Fig. 112. — x 65. A. sexangula; dorsal view of odontophoral 
support. 

Fig. 113. — x 65. A. sexangula; side view of radula with its 
retractor muscles. 

Fig. 114. — x 6h. A. sexangula; dorsal view of radula without its 
muscles. 



PLATE XVIII. 

Figs. 115-122. — x 55. Apera dimidia; serial transverse sections 
through the odontophore. 



PLATE XIX. 

Fig. 123. — x 30. Apera gibbonsi gracilis; front end of radula. 
Fig. 124.- — x 30. A. gibbonsi lupata; front end of radula. 



PLATE XX. 

Fig. 125. — x 25. Apera gibbonsi s. s.; part of radula. 
Fig. 126. — X 25. A. gibbonsi rubella; part of radula. 
Fig. 127. — X 127. A. dimidia; part of radula. 
Fig. 128. — X 127. A. burnupi; part of radula. 



THE CARNIVOROUS SLUGS OP SOUTH AFRICA. 265 

PLATE XXI. 

Fig. 129. — x 2. Apera gibbonsi s.s.; general view of digestive 
system.' 

Fig. 130. — X If. A. gibbonsi rubella; general view of digestive 
system. 

Fig. 131. — x 5. A. parva; general view of digestive system. 

Fig. 132. — x 3A. A. dimidia; general view of digestive system. 

Fig. 133. — X If. A. burnupi; general view of digestive system. 

Fig. 134. — x 2^. A. sexangula; general view of digestive system. 

PLATE XXII. 

Fig. 135. — X 113. Apera sexangiila; transverse section through 
cesophagus. 

Fig. 136. — X 125. A. dimidia; transverse section through rectum. 

Fig. 137. — x 135. A. dimidia; transverse section through wall of 
intestine. 

Fig. 138. — x 135. A gibbonsi rubella; transverse section 
through salivary duct. 

Fig. 139. — x 119. A. gibbonsi rubella; transverse section 
through part of salivary gland. 

Fig. 140. — X 119. A. dimidia; section through wall of ventricle. 

Fig. 141. — X 135. A. dimidia; section through part of kidney and 
wall of ureter. 

Fig. 142. — x 194. A. gibbonsi rubella; side view of the end of 
a papilla from inside penis. 

PLATE XXIII. 
Fig. 143. — x 2. Apera gibbonsi s.s.; reproductive organs. 
Fig. 144. — x 2. A. gibbonsi rubella; reproductive organs. 
Fig. 145. — x 8. A. gibbonsi rubella; interior of penis and 
epiphallus. 

Fig. 146. — x 6. A. parva; reproductive organs. 
Fig. 147. — x 4. A. dimidia; reproductive organs. 

' In the specimens shown in figs. 129-134 the right division of the 
liver, with the adjacent parts of the alimentary canal, is lying to some 
extent on its right side, in order to show the origin of the hepatic 
ducts, etc. (Compare figs. 27-31, in which the lobes of the liver are 
shown more nearly in their natural position as seen from above.) 



266 HUGH WATSON. 

Fig. 148. — X 2 J. A. burmipi; reproductive organs. 
Fig. 149. — X 2i. A. sexangula; rejproductive organs. 
Fig. 150. — x 1200. A. gibbonsi rubella; anterior part of a 
spermatozoon. 
Fig. 151. — X 1200. A. dimidia ; anterior part of a spermatozoon. 

PLATE XXIV. 
Fig. 152. — x 1"8. Testacella maugei ; dorsal view; Cape Town. 
Fig. 153. — x 1"8. T. maugei; side view. 

Fig. 154. — X 3"4. T. maugei; general dissection, viewed obliquely 
from the right side.' 

Fig. 155. — x 3'3. T. maugei; side view of odontophore without 
its sheath. 

Fig. 156. — X 3'3. T. maugei; longitudinal section of odontophore. 

Fig. 157. — X 100. T. maiigei; repi*esentative teeth from the 
radula. 

Fig. 158. — X 4. T. maugei; side view of junction of penis with 
vas deferens and penial retractor in specimen shown in fig. 154. 

Fig. 159. — X 4. T. maugei; side view of junction of penis with 
vas deferens and penial retractor in another specimen from Cape Town. 

Fig. 160. — X 1100. T. maugei; anterior part of a spermatozoon. 

Fig. 161. — x 1"5. T. maugei ; venti-al view of shell ; Cape Town. 

Fig. 162. — X 15. T. maugei. var. aperta; ventral view of shell; 
Cape Town. 



Explanatory References. 

nib. gl. Albumen gland, ant. aor. Anterior aorta, aur. Auricle. 
b. art. Buccal artery, b. c. Buccal cavity, b. gang. Buccal ganglion. 
b. mass. Buccal mass. b. ')nass. n. Nerve to sides of buccal mass. 

b. protr. Buccal protractors. b. retr. Biiccal retractors. b. retr. n. 
Nerve to buccal retractor. 

c.-b. con. Cerebro-buccal connective. cer. gang. Cerebral ganglia. 

c. in. Circular muscles of sheath of odontophore. com. d. Common duct. 
c.-jped. con. Cerebro-pedal connective, c.-pl. con. Cerebro-pleural con- 
nective, c. r. Circular muscles of radula-sac. 



' The liver is shown turned over to the left, and the reproductive 
system (excepting the penis) is displaced to the right; the arteries are 
omitted. 



THE CAKNIVOROUS SLUGS OF SOUTH AFRICA. 267 

diaph. Diaphragm, f d. Hevmaplirodite duct. d. m. Dorsal median 
muscle, d. ped. (jl. Duct of pedal gland. 
ppiph. Epiphalhts. 

/. Flexor muscle of odontoplioral support. /. ov. Free oviduct. 
ijen. atr. Genital atrium. ^ gl. Hermaphrodite gland. 
hep. d. Hepatic ducts. 

i. I. Inner longitudinal muscles of sheath of odontophore. int. Intes- 
tine, i. od. n. Inner odontophoral nerves. 

lah. n. Labial nerves. /. ahd. n. Left abdominal nerve to anus. 
I. hep. d. Left hepatic duct. I. liv. Left liver. /. pal. n. Left pallial 
nerve. I. j)C'>'- + ahd. gang. United left parietal and abdominal ganglia. 
I. ped. gang. Left pedal ganglion. I. pi. gang. Left pleural ganglion. 
I. r. Lateral retractors. I. sal. d. Left salivary duct. /. sal. gl. Left 
salivary gland. I. tent. n. Lower tentacular nerve. /. t. retr. Lower 
tentacular retractor. /. t. retr. n. Nerve to lower tentacular retractoi-. 
m.-cav. Mantle-cavity, m. r. Median retractors. 
n. n. Nerves of the neck. 

od. Odontophore. od. s. Odontophoral support. oss. (Esophagus, 
ces. n. Nerves to cesophagus. o. I. Outer longitudinal miiscles of sheath 
of odontophore. olf. n. Olfactory nerve, o. od. n. Outer odontophoral 
nerve, o-pt. n. Optic nerve. 

ped. art. Pedal artery. j:>efZ. n. Pedal nerves, per?, gl. Pedal gland. 
ped. gl. n. Nerve to pedal gland, pen. n. Penial nerve. jjeH.. retr. Penial 
retractor, peril, n. Peritentacular nerves. 2^c^^- '^'^'''- Posterior aorta. 

r. ahd. n. Right abdominal nerve to pericardium, rd. s. Radula-sac. 
rec. d. Receptacular duct. rec. sem. Receiataculum seminis. r. liv. 
Right liver, r. pal. n. Right pallial nerve, r.par. gang. Right parietal 
ganglion. r. ped. gang. Right pedal ganglion, r. pi. gang. Right 
pleural ganglion. /•. sal. d. Right salivaiy duct. 

sal. d. Salivary duct. sal. gl. Salivary gland. sh. od. n. Nerve to 
sheath of odontophore. sh.-sac. Shell-sac. s. m. Suspenaor muscles of 
support, st. Stomach, suhc. com. Subcerebral commissure. 

tent. art. Tentacular artery. tent. retr. Tentacidar retractor. t. r. 
Terminal retractors. 

ur. Ureter, u. t. retr. Upper tentacular retractor. 
vug. Vagina, vas def. Vas deferens, ventr. Ventricle, ves. ped. gl. 
Vesicle of jpedal gland, v. m. Ventral muscles, v. p. Ventral pocket. 



VOL. 3, PART 2. 19 



Ann. Natal Mus. Vol. I 



PI. VI 





Photo, by Watson. 



Collotype. 



Species of Apera from Natal. x i i- 

Figi. i,z, .i. gihhonsi s.s. ; l-e^, A. g. rubella ; 6, .i.g.grticilis ; j-'.), A.dimiiia i i O, l l,A. htiniupi ; 12, \ l,A. sexivigula 



Ann .Natal Miis .Vol . Ill . 



PI. VIII 



14. 



15. 
A . dibbonsi 



18 



19. 
A . g . lu-patsL 




16 




17. 
A.6. -rubella.. 



20. 



21. 

A. dlTTudicL 



22. 



A.paTn/-a X V/4. . 



23. 




A. dirtudia .X 3 ■ 





A.purcelli.A^5%, 



26. 
Abiirmapi . X 2 . 
Watson, del. J.Green. Chr'omD. 

APERA : EXTERNAL FEATURES . 



Anx\ :\aiai ^vl.^t- v.jMII. 



PI IX- 




A sexangula 



Apera; general dissections. 



PLX. 




32. 




33. 



Ann. Natal Mus. Vol. 11 



PI. X. 










32. 




Photo, by Watson 



33. 



Sections of Jperd gihbonsi ruhelhi. x -5- 



Collotype. 



PIXI. 




34. 




35. 



Ann. Natal Mus. Vol. III. 



Pi. XI, 




34. 




Photo, by Watson. 



35. 



Sections of Apera dim'uiia. ^ -5- 



Collotype. 



pixn 




3S. 



Ann. Natal Mus. Vol. 1 



PI. X 




36, X 25. 













37, X 128. 
Dopsal Skin 









Photo, by Watson & Tarns. 



39, X 118. 
Corner of Lung. 

Sections of Jpeni dimidid. 



Collotype. 



AnnNatal MusVol.lII 



pi.xm. 




55 > ays 
A piaxcelli 







46 <t 
A dunidia. 



47x7, A.p-urcelh. 



49x2 
A"bijmupi 



\ y 51 . 2 

50x3, •A.se:Mii4ula 







58x6, 
d.pecL-gl. - V^A,glupata 



Apera: shell, pedal glan.d, etc. 



62x4.6. 

Huti., London, 



Ann. Natal Mus. Vol. 111. 



1. ped. gang 



PI. XIV. 



1. pi. gang. 



1. par. -f 
abd. gang. 




_...•,■«-,, 67, X 25, 






-#■■ 



"- v?*^' 










63, X 114, 
A. g. rubella. 



r. ped. gang. 



p. pl. gang. 



p. par. gang. 




65, \ 119, 
A. g. rubella. 



e -ped. con. 



ped. gang. _ 












"^■tX-*=^ 



_ V. pl. gan^ 
r. ped. ganj 




69, X 120, A. dimidia. 



64, X 22, A. g. rubella. 

Photo, by Watson. 



66, X 114, A. dimidia. 

Collotype. 



Apera : Pedal Gland (figs. 63-66) and Ventral Nerve-Ganglia (figs. 67-69) 



Ann.Natal Mua, Vol. ID, 



PI. XV. 










djjaphj 



rod 

71 x 32. A^ dimidia. 



al.d-.- 

72 ^ 25 A. sexan^^ula , 











77 ■ , ^ ibella. 



7«xis .Vg.ru'beUa. 




75x6 \ /A.dimidi 

Apera nervous syBtem.etc. 



79.»ioo A.g.rubella, 



Ann. Natal Mus. Vol 



PI. XVI. 




90, X 118, 
Odontophoral Support. 



Figs. 80-86, 88-90, .-1. dtmidi.!. 

88, Mouth. X 60 

Photo, by Watson. 

Jperd : Sections of Tentacle and Buccal Mass. 



Fi^. 87, ^. sexangula. 
Collotype. 



Ann. Natal Mus ., Vol . Ill . 



_.^\ S.7M- cL.m. 



PI XVII. 



A.giblDonsi ^.&^2?/& 




ff 






l.r- 



m 

93. 






i ^ 



95. 96. 

A.^. rubella ^ 2. 



97. 



100. 




^'1 I i 

m 

11 

101. 102 

A.parva •< 5. h.-retr.-\ I-^q^, lO'i 



f1 
I 

r 

99. 
A.g. 
mpata 
x3. 



105. 



l.r.--i\\Vvc-Tn.T: 



106. 107. 



1- 



A. dainidia -< 6. 




108. 




109. 
A., burriupi k 4-. 110 



■$ 



ILii 



I 



m. 



Wats on, del. 



fllJ 

112. 113. 
A. Bexangula >^ qYz. 

Apera: odontopliore. 



U4'. 



lIulh.London. 



Ann. Natal Mils Vol. III. 



pi.x^ni. 



_ - d.m.. -__ 






f. 




121. 

Watson del. 

Apera dimidia: -Lr ouis . s eatijJ'rLS iJrrongrh 

X 55 



Huth, London. 

t7^e odonlophore . 



Ann. Natal Mus. Vol. 111. 



PI. XIX. 




123, A. gibbonsi gracilis, ,• 30. 




121, A gibbonsi lupata, x 30. 



Photo, by Watson 



Collotype. 



Apera : Radulas. 



Ann. Natal Mus. Vol. 111. 



PI. XX. 




125, A. gibbonsi s.s., x 25. 




126, A. g. rubella, x 25. 






127, A. dimidia, x 125. 
r'lfllKlSi'iti 



Photo by Watson. 



128, A. bupnupi, x 125. 

Apera : Radulas. 



Collotype. 



Ann.NatalMus.Vol.lU, 

1 1/ 

\\j iil—-'bprotr. 



PI. XXI. 



c*^ 



rsaZ.d.. 



\----odL. 




crop 1 



6, .V.^^v- P 



ul 



I 12M.—\-, 



129 X 2 A.gib"bonsi s.S- 130 xi^A A gruljella. 

-Ocl. 



crop 




L.sul .gL- 



L.saZ^dL. 



h.reti-. 




^.z^;K__v 



■ r. lix'. 



133 xi2/3A.."bTiniapi. 



13 2 X aVa . A, duTii dia 

Wats on. del. 

Apera- digestive system 



134.xay8\|i: 
A.sexaxLgtLLa 



Hu.th,Loiidcj\ 



Ann. Natal Mus Vol. Ill, 



PI. XXll. 











136, X 125, 
Rectum. 



135, X 113, 
CEsophagus. 







^ ,, ;;.. l&»''.:'^-^,:^gKi. 137, X 1- 



138. X 135, 
Salivary Duet. 







139, X 119, 
Salivary Gland. 



140, X 119, 
Ventricle. 




141, X 135, 
Kidney. 




Photo by Watson. 



Collotype. 



.-/peni : Histology. 

Fig. 1 ^;;, .-J. scxangula ; 136, 137, 140, 141, A. dimidia ; 1 3.S, 139, 142, A. g. iiihilLi. 



Arm NaUil Mus Vol, III 



PI XXIII, 




Apera: reproductive organs 



Ann.NatalMus.Vol.III. 



PIXXIV. 





15 3x1-8. 



15 2 xi-3. 



rd.S. '/'•"'- 




(7^f: 



159x4.. 




•}00O0<XX>C<"C»0«>Oe< 

16 xiioo. 



Wa-tson del. 



161x1-5. 

Testae ellamau^ei; Cape Town. 



16 2xisSl_J 



Huth.L onion. 



PAKTHENOGENETIC TKNDIONCY IN MKLANOCKRA MENIPFE. 269 



The Parthenogenetic Tendency in the Moth, 
Melanocera menippe (Westwood). 

By 

Eriiof^t Warren, D.Sc.(L©nd.). 



Certain attempts have been made recently at tlie Natal 
Museum to cross several species of Saturniid moths for the 
purpose of investigating the relationships of hybrids to the 
parents. It was hoped that if healthy hybrids could be raised 
it might be possible to cross them among themselves and to 
discover if any definite segregation of characters occurred. 
The experiments were effectually closed in connection with 
the species selected owing to the fact that crossing did not 
readily occur, and no healthy offspring were secured. Never- . 
theless, results of some interest were obtained. 

It was found that in one of the species, Melanocera 
menippe ( Westivood) , there is a slight, but definite, tendency 
for parthenogenesis. In this species there occurred some 
indication of an increased parthenogenesis induced by the male 
of another species. In a second case the male of one species 
certainly appeared to influence the female of another, but it 
was not possible to judge whether this was induced partheno- 
genesis or incipient hybridism. 

Material. — In the Black Wattle (Acacia mollissima) 
plantations of Natal the moth Gynanisa maia (Klug) is at 
some seasons very abundant, and the caterpillar is capable of 
doing considerable damage to the trees. During last season, 
October, 1913, to March, 1914, the moth was very common in 
the environs of Pietermaritzburg, and Mr. H. F. Pentland, 
manager of the town wattle plantations, very kindly caused 
about 130 pupa3 to be collected for me from the bases of the 
trees. The pupa3 were obtained in the month of July. 



270 ERNEST WARREN. 

I am mucli indebted to Mr. C. B. Hardenberg, Grovernment 
Kntomologist, Natal, for a supply of pupa3 of Mel an oc era 
menippe bred at New Hanover, Natal, and for other 
specimens; also for kind assistance and co-operation. 

The larvae of menippe are found on Ficus cordata Thh., 
and Mr. Hardenberg's caterpillars, obtained from the eggs of 
several moths, were fed on the same plant. With a view to 
discover a potential wattle pest Mr. Hardenberg experi- 
mented with the wattle foliage and found that the older 
larvae would eat it, but that the newly hatched caterpillars 
were unable to do so. About fifty newly hatched larva? were 
supplied with leaves of the wattle; some refused to feed, 
while others nibbled a little, but all the individuals died in 
the course of several days. 

The change of environment due to domestication or to the 
slightly different climatic conditions of Pietermaritzburg and 
New Hanover would appear to have, however, an influence on 
the larvEe in this connection, as the following experience 
indicates. 

On pairing a male and female menippe, obtained from the 
puptebred by Mr. Hardenberg, fertile eggs were laid, and the 
larv^ on hatching were mostly placed on the leaves of Ficus 
cordata. A few (some seven in fact) were placed on the 
leaves of the Black Wiittle ; but from Hardenberg's experi- 
ment it was confidently expected that they would refuse to 
feed. On the conti^ary, however, those placed on the wattle 
grew much more quickly and appeared healthier than 
those on Ficus cordata. Of sixty-three larvfe placed on 
Ficus forty-four gradually died in the course of three 
weeks, although they were supplied with both young and 
tender leaves, and with older leaves, and were carefully 
attended to. The remaining nineteen scarcely grew at all, 
and, as it was anticipated that all would die, the food was 
changed to wattle. In the next few days eight died, but the 
i-emaining eleven soon assumed a more healthy aspect and 
began to grow. At the present time (Nov., 1914) these 
eleven specimens are in a perfectly healthy condition, but not 



VARTHEXOGENETIG TENDENCY IN MELANOCEfiA MENIPPE. 271 



SO large as the individuals which have been fed on wattle 
from the first. 

Of the seven specimens placed on the wattle at the time of 
hatching* none have died. 

The experiment should be repeated, but the observations, 
as far as they go, show the adaptability of the larva and the 
ease with which the moth might become a wattle pest. 

In the following table is recorded the fate of the pupa? of 
the wild specimens of maia and the bred specimens of 
menippe. 





'S 


Males. 






Females. 








S boo 


dS 






S'S;^ 


OS tt) 


Species. 


OS 

d 

12; 


No. 


T3 

a) 
be 

a 




.-1 

St© . 
S boB 

2 ^. 
00 « g 


No. 

68 


-a 

a 

38=< 


Failed to e 
erge thron 
disease, e 


•- 

02.5 a 


Maia 


128 


60 


40' 


172 


3 


24^ 


6 


Menippe . 


25 


11 


10 


1* 





14 


14 









' Three individuals greatly deformed. ' One individual fully formed in pupa-case. 

' One individual deformed. * Four individuals fully formed in pupa-case. 

^ Fullj' formed in pupa-case. 

In comparing these two series it Avill be at once obvious 
that the bred specimens had been shielded from the adverse 
influences which proved fatal to a large percentage of the 
wild specimens. 

Out of 128 wild pupa3 of maia fifty (39 per cent.) failed to 
produce moths for one reason or another, while with the 
twenty-five bredpupse of menippe only one individual failed. 

On opening the pup^e from which images did not emerge it 
was noticed that generally the contents had mostly dried, 
and it was clear that the development of the moth had not 
proceeded to any considerable extent. The contents were, as 
a rule, thoroughly impregnated with a fungus ; but it is not 
known whether the fungus was the cause of the death of the 
pupa, or whether it appeared later. Quite possibly the cause 
of death was some bacterial disease. In any case in Natal 
the pup8S of Saturniid moths die in great numbers in this way, 



272 ERNEST AYARREN. 

and the percentage of deaths is sometimes much larger than 
in the case of maia cited above. 

In this connection a short digression may be allowable. 
Several years ago some forty full-grown, wild, and apparently 
healthy caterpillars of Nudaurelia belina were collected 
and placed in cages with earth. Pupation occurred in a 
normal manner, but only four moths were ultimately obtained. 
The majority of the pupte dried up without any appreciable 
development of the moth. The question as to the actual cause 
of death requires investigation, but the present point of 
interest to note is the great natural mortality which occurs^ 
and therefore the absolute necessity for great reproductive 
power in the species. 

An analogous case was observed with the so-called Army 
Worm, which is the caterpillar of a Noctuid, Caradrina 
e X i g u a. The caterpillar feeds on grasses, but it is moderately 
omnivorous, and will consume vegetables and other plants. In 
the middle of May, 1914, a plague of these caterpillars 
occurred in certain areas of Pietermaritzburg and its environs. 
From a piece of veld about half a mile from the town 140 
caterpillars were collected from the grasses. These were 
mostly full-grown and were on the point of pupating. They 
were placed in large wooden boxes with a layer of soil at the 
bottom, and were supplied with food in case some of the 
caterpillars had not quite finished feeding. Care was taken 
that the food supplied was dry, as the bad effect of wet food is 
well known. Ninety-two of the caterpillars died without 
pupating, and became covered with a whitish mould; the 
remainder burrowed very superficially into the ground and 
duly pupated. Of these only nineteen emerged as moths, nine 
had been stung by a large dipteron parasite, and the remainder 
(twenty) of the pupa3 dried up. Thus in this case only about 
13 per cent, of the full-grown larva3 produced moths. 

The only other available Saturniid moth in addition to the 
specimens of wild maia and bred menippe for use in these 
breeding experiments were four wild male specimens of 
Nudaurelia belina caught in Pietermaritzburg. 



PARTHENOGENETIC TENDENCY IN MELANOCERA MENIPPE. 273 

To recapitulate, the moths used in these experiments 
included forty male aud thirty-eight female Grynanisa maia, 
ten male and fourteen female Melanocera menippe and 
four male Nudaurelia belina. 

Fairing of the Moths. — The pupas were all carefully 
sexed and buried under soil in muslin cages which were placed 
in a well ventilated shed. At the advent of the wet season 
the soil was occasionally damped. 

The images of maia began to emerge first (September 17th, 
1914), and before any of those of menippe had appeared 
about fifty of the maia moths had become decrepid. The 
last moth to emerge appeared on November 5th, and it was 
deformed. The menippe began to emerge on October 6th, 
and the last moth emerged on October 25th. 

Only fresh moths were used for pairing, and since it fre- 
quently happened that fresh individuals of opposite sex of the 
two species were not available at one and the same time the 
number of possible pairings was greatly reduced. 

The total number of pairings that could be effected were : 

A. 5 (menippe ? x maia c?), 

B. 1 (menippe ? x menippe c?), 
c. 7 (maia ? x menippe S), 

I). 2 (maia ? x belina c?), 

K. 1 (menippe ? x belina d"), 
leaving 7 unpaired ? menippe and 29 unpaired ? maia. 
The paired moths were placed in cubical muslin cages of a 
capacity of about Ij cubic feet. A small branch of wattle 
and a glass pot of diluted honey were placed in each cage. It 
appears that the copulation of Saturniid moths occurs at 
night, and in no case was it observed in these experiments. 
From the account to be given below of the development of the 
eggs laid by the females it would appear that copulation 
occurred in at least four of the pairs menippe ? x maia c?, 
in the pair menippe ? x menippe <S , and in one of the 
pairs maia ? x belina (^. Whether it occurred in any 
of the other pairings is more doubtful. 

Unpaired Maia. — The unpaired maia females laid eggs 



274 ERNEST WAliKEN, 

freel}'. These eggs usually began to shrivel iu about twenty- 
days. If cut open about fourteen days after laying, the 
contents were seen to consist of a fluid, greenish-yellow, 
homogeneous yolk. In about five weeks the yolk had con- 
tracted and dried, but as a rule it did not appear to decompose 
or turn black. The number of eggs laid by an unpaired 
female varied from about 100 to 200, with a mean of about 
150. Eggs laid by the unpaired moths were cut open daily 
and examined with a hand-lens magnifying 20 diameters, but 
no trace of development or segmentation could be detected. 
The unpaired female died in about eleven days, and the un- 
paired male in about seven days. 

Unpaired Menippe. — The behaviour of unpaired me- 
nippe females was markedly different. There was the 
greatest reluctance to lay eggs, and one individual died in 
about twelve days without laying any. The eggs of menippe 
are of somewhat larger size than those of maia, and the 
number produced by a female is about 130. The average 
number laid by unpaired menippe did not exceed thirty- 
two as against 150 of maia. 

The unpaired menippe remained phlegmatic and lived 
about as long as the paired moths. On dying, the abdomens 
were mostly filled with unlaid eggs. A number of these eggs 
Avere opened and examined some five weeks after the death of 
the insect. The egg-shells were thin and not so hard as those 
of the laid eggs. Generally the yolk was still fluid and 
apparently fresh and normal in character, but no trace of 
development could be detected. 

The contents of the laid egg were of a bright yellow colour, 
and in the majority of unfertilized eggs the yolk gradually 
dried, and the egg-shell became somewhat indented in the 
course of four or five weeks. The yolk, as a rule, did not 
appear to decompose in any way. On one occasion artificial 
fertilisation was attempted. The spermatophores removed 
from a fresh male maia were rubbed over the eggs removed 
from a recently emerged female menippe. In about three 
weeks the eggs began to shrivel and darken, and on opening 



PARTHENOGENETIC TENDENCY IN MELANOOERA MENIPPE. 275 



thera the yolk was found to be very fluid, blackish, and 
decomposed; thus the condition of the yolk was in marked 
contrast with that of normally laid unfertilised eggs. 

The eggs obtained from six unpaired female menippe, 
after they had been laid for seven weeks, were cut open with 
scissors and examined with a strong hand-lens. In a small 
percentage of the eggs distinct traces of development could 
be detected. The condition of the egg as seen by a lens may 
be indicated by numbered stages. 

Stage 1, when there is no obvious development. 
Stage 2, when there is some sign of a blastoderm. 
Stage 3, when there is an obvious embryo and a few hairs 
may be visible. 

Stage 4, when the development of the embryo is consider- 
ably advanced and many hairs are visible. 
Stage 5, when the embryo actually hatches. 
Ordinary fertilised eggs of menippe hatch in about 
three weeks after being laid. In the case of the pairing of 
menippe ? x menippe (S, which occurred on October 
22nd, the eggs hatched on November 16th. 

With regard to the eggs laid by unpaired female menippe 
it was quite clear that the maximum development possible had 
occurred at the time of examination, since considerable desic- 
cation had taken place, and a few of the eggs were quite dry. 
In the accompanying table the details with respect to the 
eggs of the unpaired menippe are given. 





Eggs laid by six 


specimens of impaired m e n i p p e. 




Per- 
centage 


Development. 














Total. 
















No. 1. 


No. 2. 


No. 3. 


No. 4. 


No. 5. 


No. 6. 






Stage 1. 


32 


37 


24 


2.5 


20 


43 


181 


93-8 


Stage 2. 


1 


1 


1 


U 


2 


3 


8 


41 


Stage 3. 


1 


1 


1 








1 


4 


21 


Stage 4. 


























Stage 5 . 

















u 








Totals . 


34 


39 


26 


2.5 


22 


47 


193 


100 



276 



ERNEST WARREN, 



The four embi-yos in Stage 3 possessed a few unmistakable 
hairs. 

Thus in 193 eggs laid by six unpaired females 4*1 per cent. 
exhibited some sign of development, and 2'1 per cent, reached 
a further stage in the formation of an embryo. 

There is, therefore, a slight, but quite definite, partheno- 
geuetic tendency in menippe. 

We have seen above in the case of maia that the unpaired 
moths laid eggs freely, and an average number of about 150 
was produced. After the eggs had been laid some four weeks 
about 150 eg-gs from a number of different moths were 
opened in addition to those examined daily, but no trace of 
development could be detected. Thus no parthenogenetic 
tendency was demonstrated in this species. 

Menippe ? x Maia S. — We now return to the paired 
moths (see list, p. 273, a), and the accompanying table gives 
the necessar}^ details. 



Develop- 
ment. 


Eggs laid Ijy 5 jjaired females, menippe ? 
X maia (J . 


Total. 


Percent- 
age (in- 
cluding 
No. 6). 


Percent- 
age (ex- 
cluding 
No. 6). 


No. 3 

(paired 

Oct. 9th.l. 


No. 5 

(paired 

Oct. 9th). 


No. 6 

(paired 

Oct. 13th). 


No. 8 

(paired 

Oct.23rd). 


No. 13 

(paired 

Oct.] 8th). 


Stage 1 
Stage 2 
Stage 3 
Stage 4 
Stage 5 


45 
45 
17 
12 



84 

20 

27 

30 

1 


128 

2 





37 

8 
5 
3 



101 

68 

10 

1 




395 

143 

59 

46 

1 


61-3 

22-2 

9-2 

71 

•1 


51-9 
27-4 
11-5 

8-9 
•2 


Totals . 


119 


162 


130 


53 


180 


644 99-9 


99-9 



On compnring the above two tables it will be seen at once 
that with the exception of the pair No. 6 there is very great 
increase in the degree with Avhich development occurred, and 
also in the number of eggs in which such development was 
present. This is shown in the accompanying comparative table. 



PARTHENOGENETTC TENDENCY IN MELANOOERA MENIPPE. 277 



? m e n i p p e . 


Total. 


stage 1. 


stage 2. 


stage 3. 


Stage 4. 


Stages. 


Eggs of 6 unpaired moths . 
Eggs of 5 paired moths 


193 
644 


181 
395 


8 
143 


4 
59 



46 



1 



It may be noticed that (1) the average number o£ eggs laid 
by the unpaired menippe was only thirty-two, but by the 
paired moths it was 129; (2) with unpaired moths the per- 
centages of eggs reaching the 2nd, 3rd, 4th, and 5th stages of 
development were 4"1 per cent., 2*1 per cent., per cent., and 
per cent, respectively ; while with the paired moths the 
percentages were about 22 per cent., 9 per cent., 7 per cent., 
and 0"1 per cent, respectively. 

On examining these two series it was believed at first that 
the eggs of the paired moths had been duly fertilised with 
the exception of those of No. 6, in which the amount of 
development exhibited by the eggs was almost negligible. It 
was expected that the single larva which hatched would 
exhibit on close examination characters intermediate between 
those of menippe and maia. 

The eggs remaining in the abdomen of some of the paired 
moths were examined five weeks after death. In No. 3 (see 
table, p. 276) sixteen eggs were found ; one close to the 
external aperture coutained an embryo in Stage 3, three were 
in Stage 2, and twelve showed no sign of development. The 
twelve eggs were placed anteriorly in the abdomen. 

In No. 5 three anteriorly placed eggs were found, and these 
showed no sign of development. 

In No. 13 two eggs were in Stage 2, and thirty-six showed 
no sign. 

The eggs which occurred in the middle and front regions 
of the abdomen were thin-shelled ; and in some cases the yolk 
was dry, but in an undecomposed condition. 

It is possible that in No. 3 the eggs which exhibited develop- 
ment had come into contact with the stimulating semen of the 
male in the body of the female, although ihey were not 
actually laid. 



278 EK'NEST WAEREN. 

The Larva. — In the case of No. 5, where one caterpillar 
hatched^ the moths Avere paired on October-9th and the larva 
hatched on November 16th, thus the development occupied 
a period of thirty-eight days, while the time taken by a 
batch of pure-bred eggs was only twenty-five days, a differ- 
ence of thirteen days. Strictly speaking, about four days 
should be deducted from the periods of thirty-eight and 
twenty-five days as representing the time between paii-ing and 
the actual laying of the eggs. 

On hatching, the young larva was singularly lifeless and 
languid. It was carefully placed on the leaf of Fie us 
cordata, but during the first day it refused to feed. Sub- 
sequently it fed to a slight extent, but no appreciable growth 
occurred, and the creature died on November 26th, living just 
ten days altogether. 

The larva passed but very few ftecal pellets, perhaps seven 
or eight altogether, and these did not appear to be normally 
black, but they were greenish brown and irregularly shaped. 
It would seem that the digestive functions were not normal. 

In general aspect the larva was indistinguishable from that 
of a pure-bred menippe, but on closer examination it Avas 
seen that it was not quite normal. Some of the hairs Avere 
less black than in the majority of young menippe larvse that 
I have examined; also several of the tubercles bearing the 
long hairs of the body Avere yelloAvish and translucent instead 
of being black and opaque. The lateral, roundly triangular, 
black, raised areas at the extreme posterior end of the body 
were rather smaller than normal, and the one on the right 
hand side was only black around the periphery, the central 
portion being pale. The posterior right thoracic leg Avas 
pale broAvn instead of dark brown or black. The median 
dorsal raised area over the anus, Avhich bears tubercles and 
hairs, did not project posteriorly as far as usual, owing to its 
antero-posterior median axis being exceptionally short. In 
addition to these peculiarities the penultimate segment of the 
body and the segment in front Avere not separated off from 
one another on the right side of the body, and on this side 



PARTHENOGENETFC TENDENCY IN MELANOCERA MENIPPE. 279 

whei"e there should have been two lateral rows of tubercles 
there was only one. The outer lateral surface of the fourth 
right hand false foot was abnormally swollen and not marked 
off from the body in the usual manner, and the normal 
pigmented bar on the side of this foot was absent. 

Thus, this larva was asymmetrical, there being some 
deficiency in pigmentation, especially on the right hand side, 
and some deformity of the body on the same side. 

Comparison of Larvas. — It will be seen, however, that 
the differences between this larva and a normal larva of 
menippe do not appear to connect in any way the former 
with the caterpillar of maia. 

The larvas of some of the Saturiiiid moths are wonderfully 
alike at the first instar, but there are certain well-marked 
differences between the young caterpillars of maia and 
menippe. 

(1) In maia the general ground-colour is yellowish-brown, 
with a wide hoop-like blackish band in the middle of the 
segment. In menippe the general colour is reddish-brown, 
with sometimes a faintly defined darker transverse band in 
the middle of the segment. In the present offspring the 
general coloration was like that of menippe, but it tended 
to be somewhat paler. 

(2) In maia the tubercles on the body are bright yellow, 
in menippe they are black. In the present offspring they 
were mostly black, but three were pale and two were black at 
the base and pale above. 

(3) In maia the hairs are yellow and translucent. In 
menippe there is some variation in this matter, more usually 
they are black, but some tend to be translucent and brown 
particularly on and around the head. In the present offspring 
a considerable number were translucent and light brown, 
especially towards the apex of the hair. 

(4) In maia the median dorsal posterior raised area above 
the anus carries large tubercles and a number of equidistant 
small tubercles along the posterior margin ; these latter carry 
■one hair apiece. In menippe the area is relatively larger 



280 EKNEST WARREN. 

and projects further back over the anus, the tubercles are 
smaller, and the little marginal tubercles are fewer and not 
regularly arranged as in maia. In the present offspring the 
area was not normal in shape, and perhaps on the whole in 
general outline it Avas intermediate between that of maia 
and menippe. The condition of the tubercles and hairs, 
however, was in no way intermediate ; it was rather peculiar, 
but obviously it was a modification of the menippe type and 
had no connection with the maia arrangement. 

The fact that the arrangement of the tubercles and hairs in 
no way resembled that in maia renders it extremely probable 
that the more or less intermediate shape of the area was a 
quite accidental occurrence, due to the slight deformity of the 
posterior portion of the body. 

(5) The lateral raised area at the sides of the last false foot 
in the present offspring was like that of menippe and not 
like that of maia in general outline and arrangement of hairs, 
but it was rather small. On the right side of the body, as 
already remarked, the area was yellowish except along its 
margin which was dark brown. 

(6) In maia the head is relatively narrower than in 
menippe. In the present offspring the head resembled that 
of menippe. 

(7) In maia the mean lengths of first, second, and third 
joints of the true Avalking legs are in the proportions of 
100 : 84 : 48, in menippe 100 : 94 : 70; in the present offspring 
the proportions did not differ appreciably from those of 
menippe; they were 100 : 105 : 74, thus they showed no hint 
of a shortened terminal joint, which is characteristic of maia. 

(8) In maia the terminal claws of the thoracic legs are 
brighter in colour and considerably larger than in menippe, 
while in the present offspring they did not differ in the least 
from those in menippe. 

(9) In all other characters that were observed no differences 
could be detected between the present offspring and the young 
larvae of menippe. 

On the whole it may be said that no influence of the male 



PARTHENOGENF]'l'IO TENDENCY IN MBLANOCKRA MKNIPPE. 281 

motli could be definitely detected in the ojffspring. There was 
a certain deficiency of pigment, which, however, was doubtless 
associated with the unhealthy state of the larva, and was not 
due, as was at first thought, to the influence of the male. In 
all the more important chai-acters the present larva Avas so 
completely menippe and not maia in character that it is 
extremely doubtful if the offspring can be regarded as a 
genuine hybrid. 

Pseudogamy. — We have already seen that the presence 
of the male maia had an undoubted influence on the female 
menippe; it caused the female to lay a good supply of eggs, 
and these exhibited a greatly increased power of development. 
The only obvious explanation of this rather remarkable result 
would appear to be that copulation of the moths actually took 
place; and that the semen from the male exerted a stimulat- 
ing action on the normal, weak parthenogenetic power without 
real fertilisation, or fusion of male and female pronuclei, 
occurring. This has been termed pseudogamy, audit is the 
explanation offered by Hans Przibram ^ in the case of the 
artificial application of the semen of Mantis religiosa to 
the eggs of female Sphodromantis bioculata. vSome of 
the eggs were rendered fertile, but the offspring appeared to 
be entirely like the mother, and there was no direct influence 
from the male. 

In the present example it is to be regretted that there is no 
cytological evidence to support this view, but careful consider- 
tion of the facts renders any other supposition still more 
difficult to accept. 

If the offspring was a true hybrid the almost complete 
absence of any of the male characteristics is totally unlike 
that which is found in the vast majority of the hybrids 
between distinct species, since these so usually tend to be 
more or less intermediate between the two parents. 

A possible explanation with respect to the absence of a re- 
enforced parthenogenetic tendency in the pair No. 6 (table, 

' Przibram, Hans, ' Experimental Zoologie, Phylogenese,' p. 25, 
1910. 



282 ERNEST WARREN. 

p. 276) is that no copulation occurred, but that the close 
proximity of a male nevertheless influenced the female to lay 
a good supply of eggs. These eggs simply exhibited the 
normal amount of parthenogenesis inherent in the eggs of this 
species. The parthenogenetic power, like any other character, 
would be expected to vary to a certain extent from individual 
to individual ; but the laws of probability are against the 
supposition that the generally much increased parthenogenesis, 
and the freedom of egg-laying observable in the paired moths 
arose purely accidentally through the chance selection of such 
individuals, while none of the unpaired individuals possessed 
these characters in a high degree. It may be added that the 
further evidence that will be adduced from some of the other 
pairings is entirely against such an explanation. 

A few observations on the remaining pairings that were 
eifected may be made here. 

Menippe ? x Menippe c?. — The moths (list, p. 273, b) 
were paired on October 22nd, 1914, and 121 eggs were laid. 
The majority of the eggs were hatched on the morning of 
November 16th, the same day as that on which the partheno- 
genetic individual emerged. It was a damp, warm morning. 
The eggs of one of the batches laid by the female did not 
duly hatch. In this batch there were forty-six eggs. These 
were opened and examined, and only one showed any sign of 
development. Of the remaining seventy-five eggs there were 
twelve irregularly scattered through the different batches 
which did not hatch. On opening them eight showed no 
sign of development and four contained well-formed dead 
embryos. 

The batch of forty-six eggs which did not hatch Avere pre- 
sumably laid before copulation, and the partial development 
which occurred in the case of one egg was due to the normal 
parthenogenesis. 

Maia ? x Menippe S- — In the case of these seven pair- 
ings (see list, p. 273, c) it is less clear that copulation occurred, 
and, if it did, the resulting effect was much less marked than in 
the reciprocal cross of menippe ? x maia c?. Whether 



PAR'l'HENOGENETIC TENDENCY IN MELANOGERA MENIPPE. 283 

paired or not the female of maia lays freely, but more eggs 
were, on the whole, laid by females placed with males. 

It was noticed that the eggs of one of the paired moths were 
very exceptionally slow in shrivelling, and on this account it 
was thought that possibly copulation had occurred and more 
or less normal development was taking place. There was 
ver}" little shrivelling in five weeks after laying, although 
unfertilised eggs are obviously indented after a period of 
fourteen to twenty days. After the eggs had been laid seven 
weeks, shrivelling had become more mai'ked, and seventy-five 
eggs were opened and examined with a hand-lens. In eighteen 
eggs a whitish granular lump or lumps on the periphery of 
the yolk could be seen. An examination with the microscope 
showed that these consisted chiefly of dense clusters of needle- 
shaped crystals. The crystals dissolved in 50 per cent, 
hydrochloric acid. They doubtless ai'ose by the concentration 
of the salts in solution in the egg through evaporation, and 
their formation appears to depend largely on the slowness 
of evaporation. In rapidly drying eggs the crystals were not 
found. In 150 eggs from several unpaired maia females only 
one egg was found containing such ciystals. 

The microscopic examination of the contents of the eggs 
which so successfully resisted desiccation revealed no definite 
sign of segmentation. 

Without much more investigation it cannot be said whether 
the formation of such crystals was in any way connected with 
an unsuccessful attempt at development after copulation. An 
egg stimulated by contact with the male fluid, even without 
actual fusion of nuclei, might be able conceivably to resist 
desiccation much better than an unstimulated egg. The power 
of resisting desiccation is undoubtedly a vital act not directly 
explainable in ordinary chemical and physical terms. 

With respect to the remaining six paired moths, about fifty- 
eggs from each — that is, about 800 in all — w^ere examined, and 
only six exhibited any obvious crystal formation, and even in 
these it was less marked than in the case above described. 
These eggs began to shrivel in about twenty days. 

VOL. 3, PART 2. 20 



284 ERNE8T WARREN. 

The formation of such crystals also occurred in eggs laid 
by a female raaia paii'ed with a male belina, as will be seen 
in the account given below. 

Maia ? x belina c?. — In one of the pairings a female 
maia was placed with a captured, wild, fresh-looking male 
belina (see list, p. 273, d). There is here more evidence 
that copulation occurred, and that the semen exerted an 
influence on the eggs of the female maia. The moths were 
placed together on October 16th, and the female soon began 
to lay very freely. Two hundred and one eggs were laid, and 
these were very slow in shrivelling. On November 10th some 
of these eggs were examined, and on December 3rd, when it 
was obvious that hatching would not take place, all the 
remaining eggs were opened. Altogether forty-eight (i.e. 
about 24 per cent.) showed with a hand-lens an opacity on 
one side of the yolk. Examination with the microscope 
proved that a certain amount of segmentation of the yolk 
had occui'red ; large rounded or polygonal cells crowded 
with yolk-globules were present. On treatment with weak 
methyl blue^ a deeply staining blue spot became defined in 
each area in the course of a day, and there appears little 
doubt that this was the nucleus. There was not much 
difference in the condition of the eggs on November 10th 
and on December 3rd, except that on the latter date drying 
was taking place and death of the large yolk-cells was 
occurring. The nucleus had become less defined and was of 
inflated size and of irregular shape. Portions of the cells had 
apparently disintegrated into an opaque, white, finely granular 
substance. In some of the eggs clumps of needle-shaped 
crystals were found, and they were quite the same as those 
mentioned above, and occurred among, but apparently not 
inside, the yolk-containing cells. 

In the case of the second pairing, a female maia was 
placed with a very shabby, wild male belina. The male soon 
died. Fairly numerous eggs were laid, but these shrivelled 
rather quickly and there was no sign of development. 
Doubtless the male was spent and no copulation took place. 



PAKTHBNOGENEriC TBNDEXOY IN MELANOCERA MENIPPE. 285 



Menippe ? x belina c?. — Only a single pair was avail- 
able (see list, p 273, e). There was probably no copulation, 
or, at any rate, there was no effect ; the moths died in the 
normal time, and no eggs were laid. 

Summary. — To summarise the general differences observed 
in eggs laid by unpaired moths and paired moths, the accom- 
panying table is instructive. 





Unpaired 


Menippe? 


Unpaired 
maia ? 
(9 indivi- 
duals). 


Maia? x 


Maia ? x 


Maia ? x 
belina <J 


Moths. 


menippe? 
1,6 indivi- 
duals). 


X maia S 

(5 pairs). 


menippecJ 
(7 pairs). 


belina (J 
(1 pair) . 


(old shabby- 
male) 
(1 pair). 


No. of eggs 














examined . 


193 


HU 300 


375 


201 


170 


Percentage 












showing 












some de- 










1 


velopment 


6-1 


38-7 





240 


! 



Gerstaecker^ gives a list of species of moths in which 
parthenogenesis has been observed : 

Fam. Sphingidee — Sphinx, Smerinthus. 

Fam. Euprepiadte — Euprepia. 

Fam. Saturniida3 — S.-fam. Saturniina3 — Telea. 

Fam. Bombycidas — Gastropacha, Bombyx. 

Fam. Liparida? — Liparis, Orgyia. 

Fam. Psych id a? — Psyche. 

Fam. Tineidaj - Sol en obi a. 

Melanocera menippe belongs to the family Saturn- 
iidte, and the sub-family Sphingicanipinae, in which the 
larva pupates underground and does not form a silken cocoon 
as in the sub-family Saturniin^. 

It is interesting to note that it is just in these and in 
related families that hybridism between different species has 
been most often observed.- 

' Bronn, H. G.. • Klass. ii. Ord. d. Thier-Reichs, Arthroi)oda,' vol. v, 
p. 166, 1866-79. 

- See Przibram, Hans. • Experimental Zoologie-Phjlogenese,' pp. 
41-51, 1910. 



286 ERNKS'r WARKEN. 

The view that a parthenogenetic tendency is favoui'able to 
hybridism is also suggested by the fact that hybridism 
appears to be especially easy in Echinoderms,^ and it is in 
this group that the parthenogenetic tendency is strongly 
marked and artificial parthenogenesis can be most easily 
effected. Pseudogamy, or the stimulation of partheno- 
gensis by the semen of the male, without real fertilisation, is, 
in the case of insects, a phenomenon based, as far as I am 
aware, on indirect evidence, and not on actual cytological 
observation ; although in Echinoderms some evidence of 
this nature has been adduced. 

In the observations above described there is undoubtedly 
strong evidence that the females which were placed with the 
male of another species behaved differently than when un- 
paired, and the eggs were markedly affected. 

In the case of me nip pe ? x maia c?, the single larva 
which hatched was so fundamentally similar to a noi*mal 
menippe larva, that it would appear extremely improbable 
that it was a hybrid arising from real fertilisation. The only 
other alternative view which appears admissible is that 
copulation occurred and that a spermatozoon entered the egg 
and stimulated the normal parthenogenetic power without 
actual fusion of the male and female pronuclei, or, possibly, 
the mere contact of the male fluid acted as a sufficient 
stimulus. It has been stated, but on whose authority I have 
not ascertained, that painting the eggs of silkworms with 
different chemicals stimulates parthenogenesis, and if such is 
the case it could be readily understood that contact with the 
male fluid might have an analogous effect. 

In one of the pairings, maia ? x belina J, the male 
appears to have had an undoubted influence on the eggs, biit 
without cytological evidence it is impossible to judge Avhether 
or not there was true fertilisation, since development did not 
proceed beyond early segmentation. 

1 See Shearer, C, Morgan. Walter D., Fuchs. H. M., " On the 
Experimental Hybridization of Echinoids," ' Phil. Trans. Roy. Soc. 
Lond.,' vol. cciv. 1914. 



PARTHENOGENETIC TENDENCY IN MELANOCEEA MENIPFE. 287 

Further experimentation with cytological observations 
among- insects might be expected to lead to interesting 
results and to an increase in our knowledge of the inter- 
relationships of normal fertilisation, parthenogenesis and 
hybridism. 



NEW SOUTH AFRICAN ARACHNID A. 



289 



New South African Arachnida. 

By 

Joliii Hewitt, 

Director, Albany Museum, Grahamstown. 



With 9 Text-figures. 



Contents. 

Introduction .... 
Systematic Account 
Ord. ARANE^ 

Fam. AGELENID^ 

MUIZENBERGIA g. n. 

Muizenbergia abrahami sp. n 
Fam. ARGIOPID^ 

Erigonopsis g. n. 

Erigonopsis littoralis sp. 
Fam. CTENIZID^ 

Pelmatorycter dreyeri sp 

Bessia minor Heivitt 

Bessia fossoria Poc. 

Spiroctenus armatus Hewitt 

Acanthodon abrahami (Hewitt) 

Acanthodon ochreolum Poc. 

Acanthodon hamiltoni Poc. 

Acanthodon grandis sp. n. 

Idioi>s pretoriaj (Poc.) 

Idiops astutus sp. n. 

Hermacha mazcena sp. a. 

Hermachola g. n. 

Herniachola grahami sp. n. 

Stasimopus steynsburgensis sp 

Stasimopus gigas s^>. n. 

Stasimopus minor sp. n. 

Stasimopus oculatus l^oc . 



PAGE 

. 290 

. 290 

. 290 

. 290 

. 290 

. 291 

. 295 

. 295 

. 296 

. 299 

. 299 

. 302 

. 304 

. 305 

. 305 

. 306 

. 307 

. 308 

. 310 

. 310 

. 312 

. 314 

. 314 

. 317 

. 318 

. 320 

. 321 



290 JOHN HEWITT. 

PAGE 



Ord. SOLIFUG^ 

Chelypns hirsti sp. n. 
Ord. SCORPIONES . 

O pistil op lit halm us pugnax 
Var. iiatalensis var. n. 



323 
323 
325 
325 
325 



INTEODUCTION. 

The miscellaneous assembly of new Arachnids described 
in this paper are all in the Albany Museum, except when 
otherwise stated. Perhaps the most noteworthy are the small 
marine spiders taken by the Rev. N. Abraham at Muizenberg-, 
near Cape Town (Muizenbergia abrahami and Erigo- 
n op sis littoral is), a discovery of some interest seeing that 
previously only one marine genus, viz., Desis, had been 
known from South Africa. Whether their occurrence on our 
coast has any special zoo-geographical importance it is impos- 
sible to decide at present, as little or nothing is known of the 
Arachnid fauna of other shores ; in any case it seems very 
probable that quite a new field awaits exploration on the coasts 
of the southern hemisphere, in worm-tubes and similar habitats 
between tide-marks. 

Some of the descriptions are based on material which has 
been kindly lent to me by the authorities of the British 
Museum, London, and of the Natal Museum, Pietermaritzburg; 
for this courtesy I am especially indebted to Mr. A. S. Hirst, 
the Arachnologist of the British Museum, and to Dr. E. 
Warren, the Director of the Natal Museum. 

SYSTEMATIC ACCOUNT. 
Oeueu ARANE^. 

Fam. AGELENIDJE. 
Sub-fam. Agelenin^:. 

Group CryphcEce^e. 
Genus Muizenbergia gen. nov. 
This new genus is at once distinguished from any known 
genus of this group through the position of the tracheal 



NEW SOUTH AFRICAN AllAOtlNlDA. 291 

stigma^ which is situated about midway between the genital 
opening and the base of the inferior spinners. The cheliceraj 
have two teeth on each side of the fang-groove. According 
to Mn. Simon's ' key to the genera, Muizenbergia should be 
placed near to Cicurina Menge. 

Muizenbergia abrahami sp. ii.oc. Text- fig. 1, a-g. 

The types of this species, a single adult male and a some- 
what shrivelled female, were collected at Muizenberg, near 
Cnpe Town, in September, 1913, by the Rev. N. Abraham, 
who presented them to the Albany Museum. Mr. Abraham 
found them when searching for Desis, in the interspaces of 
the calcareous masses built up by marine annelids ; they are, 
therefore, marine spiders, probably resembling Desis in habit. 

Colour. — Chelicerte and anterior portion of carapace 
castaneous ; hinder portion of carapace pale brown, like the 
legs and sternum ; distal segments of legs more deeply 
coloured than the basal segments ; on the cephalic portion of 
the carapace there is some slight infuscation, which is only 
strongly pronounced around the eyes. Abdomen uniformly 
dull dirty grey. 

Carapace. — Longer than wide, the cephalic region fairly 
strongly convex, gradually becoming much flatter in the 
thoracic region ; these two regions are not sharply separated 
though their junction is marked on the margin by an obtuse 
angle. Anteriorly the carapace is squarish, the clypeus being 
very wide and very low ; the distance from an anterior 
lateral eye to the anterior margin of the carapace is less than 
the long diameter of the eye. Fovea long and very narrow. 

Ocular Area. — Nearly three times as broad as long. 
Anterior row straight, posterior row slightly procurved. 
Antero-medians smallest, but not very small, a little nearer 
to the antero-laterals than to each other. Eyes of posterior 
row subequally spaced. The laterals of each side very close 

' Simon, Eugene, ' Histoire Naturelle des Araignes," tome ii, p. 2H6, 
Paris. 1897. 



Text-fig. 1. 




■ 'I lU! j, 






Muizenbergia abrahami s^j. jior. 
A. Carapace of male, x 40. b. Tarsusof male jialp with bulb in situ, x 75. 
c. Male j)alp with bulb of tarsus dislodged. X 55. D. Process of the patella 
of the male palp. X 185. E. Posterior half of abdomen of male in ventral 
view showing spinners and tracheal stigma. X 40. F. Spinners of female. 
X 25. G. Ventral view of sternum, labium, coxifi of appendages, and one 
chelicera of the male. X 40. 



NEW SOUTH AFRICAN AKACHNIDA. 293 

together. Quadrilateral formed by the four median eyes 
broader behind than in front, and the posterior breadth 
exceeds its length. 

Chelicerae. — Elongated, a little dilated in the basal half, 
projecting obliquely forwards. Fang strongly curved, reach- 
ing backwards nearly half the length of the chelicera; on 
each side of the fano-.o-roove there are two teeth, of which 
the distal one is large, sharply pointed, and triangular, the 
more proximal one small. 

Pedipalp of Male. — Both the patella and tibia carry a 
strongly projecting process. That of the tibia, which is the 
longer, is a tapering cylindrical process, strongly curved 
backwards and ending in a point ; that of the patella ai'ises 
from a broad base and is expanded over half of its length, 
the expanded portion carrying a group of three bristles, 
distally to which the process suddenly narrows and even- 
tuall}^ ends in a claw-like hook. The tarsus is oval and 
elongated, but not twice as long as broad ; the distal end of 
the bulb is not far from the apex of the tarsus. Numerous 
long, plumose, bristly hairs occur on the tarsus. 

Legs. — All the hairs, bristles, and spines which invest the 
legs and the palps are finely plumose. None of the tarsi are 
spined. The first metatarsus and tibia in the female are 
quite without spines (in male specimen damaged). The 
second metatarsus has a pair of long spines, near the apex, 
and the third metatarsus has two pan-s of long spines below ; 
the third and fourth tibige and fourth metatarsus are beset 
with a number of long and strong spines. The hairs on the 
legs are not very densely arranged. The paired tarsal claws 
of the first leg in the male have nine teeth each, the more 
basal teeth being minute ; the unpaired claw has two teeth, 
but that of the female has only one. The unpaired claw of 
the fourth leg of the male has three teeth, that of the female 
only two. 

Labium. — About as broad as long; broadest near the 
base and tapering gradually towards the anterior margin, 
which is broadly rounded and carries four long bristles; it 



294 JOHN HEWITT. 

reaches about as far as two-thirds of the distance along the 
maxillte. 

Sternum. — Only a trifle longer than wide, broadly 
truncate in front and narrowing to a point posteriorly 
between the hind coxte, which are completely separated 
thereby. It is broadest opposite the cox^ of the second legs. 

Abdomen. — Elongated, covered with soft, slender, plumose 
hairs, all backwardly directed. On its ventral surface the 
tracheal stigma is placed far forwards, being situated only 
very slightly posterior to the mid-point between the genital 
aperture and the bases of the inferior spinners.^ 

Spinners. — Inferior spinners widely separated, about 
twice their own width apart. Superior spinners with the 
apical segment about half the length of the basal one. 
Colulus absent. The median and superior spinners arise only 
very slightly posterior to the inferior spinners, especially in 
the male, where the Avhole six are arranged almost in a 
transverse line ; in the female the arrangement is roughly in 
two transverse lines. 

Total length. — Adult male 3"5 mm. 

This genus in its spinner arrangement approaches the 
group Hahniinae and will probably prove to be closely 
related to some of the little-known genera of that group. 
The tibial process of the male palp is very like that figured 
by Simon- for Hahnia oreophila E. Sim. The position 
of the tracheal stigma is also apparently an extreme condi- 
tion of a tendency obtaining in various genera of this group. 
I have placed Muizenbergia provisionally in the group 

' Since writing the above, I have examined a large female example 
from St. James, near Cape Town (coll. J. H. Power). This clearly shows 
that the median tracheal stigma is double, the actual openings being 
connected by a sliort transverse furrow. The tufts of hairs shown on 
E, text-fig. 1, mark the positions of these openings. Further, the lateral 
eyes of each side are slightly raised on an obliquely disposed prominence. 
Op. cit., p. 273. 



NEW SOUTH AFIUL'AN ARACHNIDA. 295' 

CryphoeceEe mainly on account of the dentition of the 
clielicei-Ee; probably the separation of the two groups is 
artificial. Whether this genus can be regarded as a direct 
derivative of the South i\^frican terrestrial fauna, or whether, 
like Desis, its allies are widespread over the shores of the 
southern hemisphere, cannot be stated at present ; it does 
not seem to be closely related to the various known littoral 
species recorded by Mr. H. R. Hogg.^ 

Fam. ARGIOPID.^. 

Sub-fam. LiNYPHiiN.i<]. 

Genus Erigonopsis gen. nov. 

This generic name is proposed for the reception of the 
species hereafter described, which combines the characters of 
various members included in Simon's world-wide group 
Erigonea3, but does not agree entirely with any one of them, 
and in some respects approaches the group Liny phi a\ The 
characters of the male palp will probably prove sufficient to 
differentiate this from any other known genus. Other 
characters are : carapace of male siinple, precisely like that of 
the female ; no trace of fovea or median sulcus on carapace ; 
length of quadrilateral formed by the four median eyes 
slightly greater than its hind width ; posterior median eyes 
very slightly nearer to each other than to the posterior 
laterals ; sternum with a rather broad posterior prolongation, 
separating the bases of the foui'th legs from each other, 
followed, posteriorly to the coxas, by a small flat quadrangular 
piece ; abdomen without scute or indurated area ; female 
epigyne comparatively simple ; female palp without tarsal 
claws, the maxillge not very broad even in the male (more or less 
intermediate between Erigone and Liny phi a) ; chelicerje 
with four teeth on the anterior margin of the fang-groove and 
thi'ee on the posterior margin in both sexes ; tarsus of first leg 

' Hogg, H. R., " Subantarctic Islands of New Zealand," Article IX, 
Wellington, N.Z., 1909 ; •' Some Falkland Island Spiders." ' Proc. Zool. 
Soc.,' 1913, p. 37. 



'296 JOHN HEWITT. 

very slightly longer than the metatai-sus or subequal thereto, 
and the metatarsns very slightly shorter than the tibia. 

Erigonopsis littoralis sp. nov. Text-fig. 2, a-h. 

The types of this species are one adult male and several 
adult females taken at Muizenberg, near Cape Town, by the 
Rev. N. Abraham in September, 1914. The habitat is the 
same as that of Muizenbergia abrahami sp. nov. 

Colour. — Carapace, sternum, and appendages brownish, 
without strong infuscation except around the eyes : the cara- 
pace and sternum are lightly infuscated, more especially so 
about the middle of the carapace. The skin of the abdomen 
is for the most part infuscated and is covered with black 
rather bristly hairs ; on the hinder half of the abdomen are 
five pale transverse stripes dorsally, and there is a pair of pale 
spots about midway between the anterior stripe and tlie front 
margin of the abdomen. 

Carapace. — Narrowed anteriorly, the front margin broadly 
rounded ; cephalic area fairly strongly convex, gradually pass- 
iug into the slightly convex thoracic portion, the boundary 
between the cephalic and thoracic portions of the carapace 
being very ill-defined both at the margin and elsewhere. 
Clypeus vertical but not greatly elevated, the anterior lateral 
eyes being about three diameters distant from the antei-ior 
margin of the carapace. Anterior margins of anterior row of 
eyes in a distinctly recurved line ; posterior row only slightly 
procurved ; anterior medians smallest, but not very small, 
nearer to each otlier than to the anterior laterals. Along the 
mid-line of the carapace in its cephalic portion there are four 
or five weak bristles, and a few stiffish hairs occur on the 
ocular area, but none on the margin of the carapace. 

Chelicer^. — Fairly stout, directed vertically downwards, 
without spines at the sides and without basal spot ; fang 
strong and rather short ; anterior margin of fang-groove 
carrying four teeth, of which the middle ones are strongest; 
the posterior margin with three small teeth. 



Text-fig. 2. 




Evig-onopsis littoralis sp. nov. 
A. Dorsal view of carapace and abdomen. X 30. B. Tarsal claws of first leg. 
X 375. c. Barbed hairs on margin of fourth tarsus. X 185. D. Dentition 
of cheliceraj. X 100. e. Maxilke, labium, and anterior part of sternum. 
X 90. F. Posterior part of sternum, cox* of fourth pair of legs, and epigyne. 
X 90. G. Distal portion of male palp. X 200. h. Tibia of same showing 
the apophysis. X 250. 



298 JOHN HEWITT. 

Legs. — Cox^ I and IV of equal length. Metatarsus TV 
longer than the tarsus. Apart from a few long spiniform 
seta3 on the femora and tibias, the legs are spineless ; there 
are a few long setiforra spines on the distal segments of the 
palps. The legs are clothed with short stiff setas ; most of 
these are simple, but on the tarsus, and to a less extent on the 
metatarsus, there are some long stiff setj\3, each carrying two 
or three lateral barbules, usually situated on one side of the 
hair about the middle of its length. On tarsus IV the stout 
feathered hairs are arranged in longitudinal rows, about ten 
in a row. The elongated so-called auditory hair on the fourth 
metatarsus is present. On tibia IV dorsally there are two 
projecting hairs, the distal one longer. Onychium present. 
Paired claws of the first tarsus with only five teeth, the distal 
one longest, the others much smaller and gradually decreasing 
in size towards the base ; the median claw is fairly long and 
carries two teeth, the basal one of which is very minute and 
arises from the base of the larger one. On the claws of the 
fourth tarsus the teeth are still weaker, the inferior claw being 
drawn out to a long fine point and carrying only one weak 
tooth. 

Sternum. — Broader than long, quite fused with the labium. 

Labium. — Veiy broad, carrying four bristles anteriorly; 
there is a strongly thickened slightly concave border antei-iorly. 
The maxillae have the long axis very obliquely inclined. 

Palp. — Male palp short, the femur being about twice as 
long as the patella, and their conjoint length about equal to 
the distance from the base of the tibia to the end of the 
conductor of the style of the bulbal organ. The end of the 
conductor somewhat resembles a scoi"pion's vesicle (sting) in 
shape ; immediately beyond the distal extremity of the tarsus 
the style is bent at right-angles on itself, its distal portion 
continuing to the apex as a broad slightly curved lamina. 
Tibial apophysis not bifurcated and not large. External 
branch of the tarsus (paracymbium) slightly arched, but no 
spines nor hairs occur on the concavity. 

Total length. — Female 2^ mm. Male If mm. 



NEW SOUTH AFRICAN ARACHNID A. 299 

Various members of this group of Argiopidse are known to 
live in marslies and on the sides of streams, but no marine 
forms have been hitherto recorded. 

Fam. CTENIZID^. 
Pelmatorycter dreyeri sp. nov. Text-fig. 8, 

The type consists of a single adult male from Bloemfontein, 
presented to the Albany Museum by Dr. T. F. Dreyer, in 
August, 1914. The species is related to P. nigriceps^ 
Purcell, from Johannesberg, but differs therefrom in the 

Text-fig. 3. 




. o o „ 

Pelmatorycter dreyeri sp. nov. 
Eyes of male. X 45. 

ocular arrangement, in the dentition of the claws of the 
fourth leg, and in some minor points. 

Ocular area. — Wider behind than in front, the anterior 
row of eyes strongly procurved, the laterals much larger than 
the medians, the long diameter of the former being quite 
one and two-third times the diameter of the latter. Posterior 
laterals decidedly longer than the anteinor medians, but 
shorter than the anterior laterals. 

Chelicerfe. — With seven teeth on the fang-groove. 

Pedipalps. — Near the base of the tibia on its inner side 
is a spine. 

Legs. — Tarsus II, without spines; III with four spines on 
its anterior side, two or three postero-dorsally situated, and 
one on the posterior side ventrally situated ; IV with no 
spines on its posterior side. Posterior claw of tarsus IV with 

1 Purcell. W. F.. ' Trans. South African Philosophical Society,' vol. 
xi, p. 358. 

VOL. 3, PART 2. 21 



300 JOHN HEWITT. 

one tooth only in the outer row and four in the inner row ; 
anterior claw with three small ones in the outer row and four 
large ones in the inner row. Metatarsus I equal in length to 
the distance from the centre of the fovea to the anterior 
margin of the carapace. Patella IV without an apical spine 
interiorly. The distal group of spines on femur IV, at its 
upper outer edge, is composed mostly of weak spines ; there 
are only very few short, fairly stout, spines. 

Abdomen. — With a few, rather strong bristles near the 
base above. 

Posterior sternal sigilla. — Not very clearly defined, 
but apparently rather more than a length apart, and about 
half a length distant from the sternal margin. 

Posterior spinners. — Apical segment a trifle shorter 
than the penultimate segment. 

Measurements. — Total length 18'5 mm., length of cara- 
pace 5*5 mm., width of carapace 4*4 mm., length of metatarsus 
of first leg 3"5 mm. 

A female specimen taken at Bloemfontein by Dr. Dreyer, 
a few days later, presumably belongs to the same species. 
The characters are detailed below. 

Ocular area. — More than twice as broad as long, the 
posterior row distinctly wider ; anterior row with anterior 
margins moderately procurved, posterior margins in a very 
lightly procurved line, the medians slightly nearer to the 
laterals than to each other, their distance apart being 
distinctly greater than a diameter; posterior row with pro- 
curved anterior margin and recurved posterior margin, the 
laterals oblique and elongated, much longer than the medians 
and as long as the anterior laterals, posterior medians nearer 
to the posterior laterals than to the anterior medians ; distance 
between the laterals of each side about one-third the length 
of the posterior laterals. 

Posterior sternal sigilla. — About half a length apai't 
and almost a length distant from the sternal margin. 

C h el i cerge . — The inner dental series comprises eight teeth. 



NKW SOUTH AFRICAN AKACHNIDA. 301 

Pedi palps. — Coxa with five teeth on the inferior sux-face 
situated far forwards. Tibia with four apical spines inf eriorly, 
also four on the anterior surface below, but none on the 
posterior surface. Tarsus with two spines not far from the 
apex inferiorly, and a longer one on each side near the base. 

Legs. — Tarsi and metatarsi of first two pairs of legs 
scopulate to the base (on the second metatarsus the scopula 
is absent on the posterior side in its basal half). Tarsus I 
without spines, excepting one near the apex inferiorly ; II 
with one or two spines inferiorly on the posterior side; III 
with a row of six spines postero-dorsally, of three antero- 
dorsally, also about seven antero-inferiorly situated ; IV 
strongly aculeate on the anterior surface and with a number 
of spines on the posterior surface in its distal half. Meta- 
tarsus I with one or three apical spines below, and two or 
three along the inferior surface ; II with three apical spines 
beloAv and two along the inferior surface ; III numerously 
spined on both anterior and posterior surfaces superiorly, and 
with four weak spines on the inferior surface, in addition to 
those at the apex, which are long and strong ; IV with 
numerous spines inferiorly, mostly situated on the anterior 
side, superiorly with two spines on the posterior side. Tibia 
I and II each without distinct spines beloAv except one at the 
apex ; III without spines at the apex inferiorly, a supero- 
anterior band of seven spines, also seven dorsal spines, and 
two near the supero-posterior edge distally ; IV with a single 
apical spine inferiorly, and with one spine or none on the 
postero-dorsal edge. Patella III covered with short, stout 
spines on the anterior surface, the dorsal surface with two 
stout spines near the posterior edge ; IV without spinules 
on its anterior surface, except two or three immediately at 
the base of the segment. Femur IV with a dense group of 
short, strong spines anteriorly at the apex and dorsally. 
Tarsal claws of first leg with four teeth in the outer rows, and 
five or six in the inner rows ; anterior claw of fourth leg with 
only one small distal tooth ; the posterior claw with one or 
two distal teeth and one at the base, also a very minute 



302 JOHN HEWITT. 

second basal tooth. Coxa III with a dense tuft of stiffish 
setae on the postero-ventral border. 

Colour. — For the most part castaneous above, chelicer.© 
blackish, patellae and more distal segments of first two pairs 
of legs and of palps pale brown ; carapace pallid in its hinder 
portion at the sides. Abdomen pale except over the median 
area above, where it is infuscated. 

Measurements. — Total length 23*5 mm., length of cara- 
pace 8'5 mm., breadth of same 5*5 mm. 

According to Dr. Dreyer, the nest has two long blind side 
passages leading into the central tube one on each side in the 
upper half of its length ; the upper part of the tube projects 
a little above the siirface of the ground, and the distal end is 
folded inwards, thus closing the entrance to the nest. 

Bessia minor Heivitf. Text-fig. 4. 
Bessia minor Hewitt, Records, Albany Museum, vol. ii, p. 469, 1913. 

An adult male of this species was taken at Alicedale by 
Mr. F. Cruden on March 12th, 1914, and a description of it 
is here given. 

Carapace. — As long as the metatarsus and tarsus of the 
first leg ; the lateral margins fringed with strong bristles, and 
the abdomen also is bristly over its median ai-ea superiorly. 
Fovea lightly procurved. 

Cheliceree. —Dentition somewhat similar to that of the 
female, the teeth not arranged in continuous well-defined 
rows ; the larger teeth are roughly arranged in a double 
series — the inner of which is weaker — in the middle of the 
group, and a single series at each end. 

Pedipalp. — Resembling that of a Spiroctenus. The 
tarsus has numerous short spines distally above; the tibia has 
long spines below ; at the base of the coxa inferiorly is a 
patch of cusps. 

Legs. — Tarsi not spined, all of them scopulate on the 
sides, the fourth tarsus only weakly so ; paired claws with a 
single spirally curved row of numerous long teeth as in 



NEW SOUTH AFRICAN ARACHNIDA. 



303 



Spiroctenus. Metatarsus I longer than the tibia, slightly 
scopulate in its apical third, slightly but distinctly bowed ; 
II with several scopular hairs near the apex ; III and IV not 
scopulate. Apical tubercle of tibia I with a fairly long and 



Text-fig. 4. 




Bessia minor Hewitt. 
Stermim. labium, and basal parts of the appendage of the male. 

X 18. 

stout spur at its apex, and a slightly shorter and less stout 
spur on its base externally ; the proximal tubercle also bears 
a spur and is situated rather less than a quarter of the length 
of the segment from the apical border. Anterior surface 
of patella III with a row of three spines superiorly and 
two longer ones distally below. All the femora are spined 
above. 



304 JOHN HEWITT. 

Labium. — With about twenty-four cusps on its anterior 
half. 

Sternum. — Posterior sigilla elongated, about one and a 
half diameters apart and about half a diameter distant from 
the sternal margin. At the margin of the sternum opposite 
the base of the second leg there is a deep circular depression 
in which is situated the second sigillum ; a similar depression 
also occurs in the female, but is not so noticeable, as the setse 
which fringe its margin are not so strong as in the male. 
Sternal margin fringed with strong bristles, especially pos- 
teriorly. 

Posterior spinners. — Apical segment about three- 
quarters the length of the penultimate segment. 

Measurements. — Total length 15"2 mm., length of cara- 
pace 6"5 mm., breadth of same 4*8 mm., length of first 
metatarsus 4 mm., of fourth metatarsus 5'8 mm. 



Bessia fossoria Foe. 

Bessia fossoria Poc, Ann. Mag. Nat. Hist., vol. vii, 6, p. 820, 

1900. 

I have examined the type, which is very immature, and 
have now no doubt but that the much larger specimen from 
Redhouse, referred to in a previous paper,^ belongs to this 
species. The dentition of the paired tarsal claws was incor- 
rectly described by Pocock, for the inner distal row is repi-e- 
sented by two or three minute teeth on each claw. Moreover 
the tarsus of the first leg and the distal half of the metatarsus 
is rather strongly scopulate, the tarsus of the second leg is 
scopulate at the sides, but the third and fourth tarsi ai'e not 
scopulate. The teeth on the chelicerfe constitute a single 
row, but the line is irregular, not straight. 

The genus is evidently very closely related to Spiroc- 
tenus Sim. 

' Hewitt, J., ' Records, Albany Museum,' vol. ii, p. 471, 1913. 



NEW SOUTH AFRICAN ARACHNIDA. 305 

Spiroctenus armatus Heiclft. 
Spiroctenus armatus Heivitt. Records, Albany Museum, vol. ii, 

p. 467, 1913. 

In drawing up the description of this species the following 
characters were accidentally omitted. 

Colour. — Carapace and legs reddish brown, palps pale 
brown ; abdomen pale with ill-defined dark markings above ; 
ventral surfaces pale. 

Posterior sternal si gill a. — Elongated, oval, about 
one and a half diameters apart and about half a diameter 
distant from the sternal margin. 

Measurements. — Total length 15 mm. ; length of cara- 
pace 6'4 ram. ; breadth of same 5"1 mm. ; length of metatai-sus 
of first leg 4'2 mm., of fourth leg 6*2 mm. 

Acanthodon abrahami (Hewitt). 

Gorgyrella abrahami Hewitt, Records, Albany Museum, vol. ii, 

p. 473, 1913. 

I now prefer to place this species under the genus 
Acanthodon, although formerly I doubtfully referred it to 
Gorgyrella. It is probably closely related to A. ochreo- 
lum Poc, of which only the male is known. The male of 
abrahami seems to differ from that of ochreolum in the 
closer approximation of the frontal eyes and in the less 
strongly bent first metatarsus, but until more Jansenville 
material is available for comparison the specific distinction of 
abrahami should be held somewhat doubtful. Two adult 
males of abrahami were taken by Mr. F. Cruden at 
Alicedale on February 15th, 1914, and I here give a des- 
cription of the sex. 

Frontal eyes quite separated, about one-third of a diameter, 
or slightly less, apart ; the quadrangle formed by the frontal 
and anterior median eyes wider behind, the medians large, 
aboiit three-fifths to three-quarters of a diameter apart ; hind 
margins of posterior row about in a straight line, the medians 
two to two and a half diameters apart and a diameter or 
slightly more distant from the laterals. 



306 JOHN HEWITT. 

Tarsi of all the legs scopulate below throughout their 
length, the first tarsus being only quite thinly scopulate, the 
fourth without setge along its middle. Tarsal claws of first 
leg with three teeth, of fourth leg with two teeth and a third 
tooth may or may not be just indicated. First metatarsus 
slightly bowed, with two spines at the apex inferiorly and 
three along the postero-inferior edge, but none along the 
antero-inf erior edge. First tibia shorter than the metatarsus, 
with four spines along the postero-inferior edge, with the 
usual pair of tubercles near the apex, the more proximal one 
being distant from the apex less than one-sixth the length of 
the segment. Band of spinules on anterior surface of patella 
IV stretching two-fifths to one half of the length of the seg- 
ment and comprising only about eight to ten spinules ; 
III has two to four spines along the distal edge anteriorly, 
and two to nine on the anterior surface, but only one on the 
dorsal surface, in addition to one or two on the distal edge. 

Tibia of pedipalp barely one and two-third times as long as 
deep, the band of spines bordering the excavation complete, 
in three rows proxiniall}^, but elsewhere in a single row, 
except at the distal end ; altogether there are nineteen to 
twenty-four such spines. Tarsus of pedipalp with one long 
spine at the apex superiorly and one or two weaker ones, the 
distal inferior lobe on the posterior side rather strongly 
projecting. Process of palpal organ comparatively short, 
flattened, and twisted. 

Measurements. — Total length 12*5 mm., length of cara- 
pace 4*5 mm., breadth of same 3*5 mm., length of tibia of first 
leg 2*75 mm. 

Acanthodon ochreolum Poc. 
Acantliodon ochreolum Poc, Ann. Mag. Nat. Hist., 7, x, p. 9,1902. 

This species is probably the same as either Gorgyrella 
schreineri Purcell, or G. abrahami Hewitt, for it has 
three pairs of sternal sigilla. 

The following notes on the type specimen will supplement 
Pocock's rather incomplete description. 



NEW SOUTH AFRICAN ARACHNIDA. 307 

Frontal eyes quite separate from each other, about half a 
diameter or a trifle more apart. The quadrangle formed by 
the frontal and anterior median eyes is appreciably wider 
behind. Hind marg-ins of posterior row of eyes in a pro- 
ciirved line. Posterior median eyes three to three and a half 
diameters apart. 

Tarsus of palp witli one rather long- stout spine at the apex 
dorsally, and several much weaker ones. Pocock's figure of 
the tibia of the palp is quite inaccurate in respect to the 
:spinulation of the excavation ; the spinules are not arranged 
in a single row throughout, and are much more numerous 
than represented in the figure ; distally they occupy two or 
three rows, and proximally the spinules are considerably 
longer than elsewhere. 

All the tarsi are scopulate to the base, the fourth being 
without spinif orm seta; mesially below. Tarsal claws without 
a comb of teeth, the first having only one large tooth and the 
fourth one or two. The first tibia has four spines along its 
outer inferior edge, but none otherwise situated, apart from 
the two large tubercles. 

The inner dental series of the cheliceras comprises five large 
teeth. 

The first pair of sternal sigilla is near to the sternal 
margin. 

A cant ho don hamiltoni Poc. 
Acanthodon hamiltoni Poc, Ann. Mag. Nat. Hist., 7, x, p. 320, 1902. 

The type of this species is without doubt very juvenile. 

The inner dental series comprises five strong teeth and a 
few small ones basally situated. The coxfe of the legs are 
without spinules, and alojig the posterior edge of the third 
■coxa inferiorly is a strip of moderately fine hairs. The second 
tibia has a row of eight spines on its anterior side. The band 
of short strong spines on the anterior surface of the fourth 
patella extends two-thirds of the distance along the segment. 
Tlie frontal eyes are very close together; the ocular tubercle 
is deeply grooved above. 



308 JOHN HEWITT. 

A cant ho do 11 grandis sp. iiov. Text-fig. 5. 

This may be the female of A. pectinipalpis Pure. (Annals 
S. Afr. Mus., vol. iii, p. 87) described from Zululand, but I 
think it is probably distinct on account of the wider separation 
of the frontal eyes, those of pectinipalpis being described 
as very close together. It is related to a species described 
by me ^ from Newington, Transvaal, under the name of 
Ctenolophus transvaalensis, but differs in that the third 
coxa is clothed inferiorly with stout bristles, not spinules, and 
in other respects also. 

Text-pig. 5. 








0v///////'/.,.,.., 



Acauthodon grandis sp. Hor. 
The eyes of the female. X 16. 

The type consists of a single adult female in the collection 
of the Natal Museum from Umfolosi, Znluland (F. Toppin 
coll. 1905). 

Colour. — Upper surface of appendages and carapace 
castaneous, the legs with a reddish tinge. Ventral surfaces 
paler; sternum and coxte of last two pairs of legs yellowish- 
brown. 

Carapace. — As long as the tibia, metatarsus and half of 
the tarsus of the fourth leg, or the patella, tibia and three- 
fifths of the metatarsus of the first leg. Ocular area only 
slightly wider than long, its width subequal to the length 
' Hewitt, John, 'Records, Albany Museum,' vol. ii. p. 412. 1913. 



NLIW SOUTH AFRICAN AHACHNIDA. 309 

of metatarsus I, its length almost two-fiftlis of the distance 
from the anterior margin of the carapace to the centre of the 
fovea. Area formed by the frontal and anterior median eyes 
slightly wider in front, the medians small, about two diameters 
apart, the frontals large, about one-third of a diameter apart 
and situated on a common tuljercle which is deeply grooved 
above. Posterior row about equally spaced, the area formed 
by the four medians appreciably wider behind ; posterior 
laterals large, the medians considerably larger than the 
anterior medians, the hind margins in a slightly procurved line. 
Legs. — Metatarsus III with three long weak spines below 
and three stronger ones at the apex inferior! y ; IV with six 
or seven spines along the lower surface and three at the apex 
inferiorly. Band of spines on anterior surface of tibia I 
reaching to the base; of II also reaching to the base and 
including about twenty-two spines ; IV with two inferioi" 
apical spines and one weak one on the lower surface. Band 
of spines along anterior surface of patella III including about 
thirty spines, of which about seven are on the distal edge; on 
the posterior distal edge there are five spines; IV with a band 
of short stout spines stretching the whole length of the 
anterior surface. Coxfe of legs without spinules below, the 
third having a dense patch of short coarse bristles or weak 
spines along its posterior border ventrally. 
Labium. — With two apical teeth. 

ChelicerEe. — Inner row of teeth on fang-groove including 
five large teeth ; the outer group without any lai-ge teeth. 

Sternum. — Three pairs of sigilla, the third pair small and 
not so conspicuous as the preceding pairs ; first pair sub- 
marginal, second pair their own length distant from the 
sternal margin. 

Measurements. — Total length 25mm.; length of carapace- 
10-2 mm., width of same 9 mm.; length of tibia of first leg 
4-25 mm. This is the largest species of Acanthodon 
known to me (excluding Gorgyrella namaquensis and 
G. schreineri, but Gorgyrella is hardly separable from 
A c a n t h o d o n) . 



310 JOHN HEWITT. 

Idiops pi-etorifB (Poc). 
Acanthoclon ijretoriEB Poc, Ann. Mag. Nat. Hist., vii, 1, p. 319, 1898. 

To the description given by Mr. Pocock I can add the 
following noteSj after examination of the type male : 

Chelicerae with seven strong teeth in the inner row and four 
in the outer row ; process of palpal organ strongly flattened 
thi-oughout and twisted ; the excavation on the tibia of the 
palp very shallow and not bordered by spines; band of 
spinules on anterior side of patella IV stretching five-sixths 
of the length of the segment and including about thirty short 
spines; patella III has a continuous strip of spinules anteriorly 
including about thirteen as well as four on the distal edge ; 
frontal eyes one-quarter to one-third of a diameter apart, the 
anterior medians very large, the area formed by the frontal and 
anterior median eyes much broader behind, posterior medians 
very much smaller ; posterior margins of posterior row of eyes 
slightly recurved, the anterior margins slightly procurved ; 
the posterior medians about three diameters distant from the 
posterior laterals, which latter are a little elongated but not 
greatly so. 

Idiops astutus s'p. noc. Text-fig. 6. 

The type consists of a single adult male taken at Bulawayo, 
November loth, 1913, by Mr. G. Arnold, who writes of it 
" hunting insects under the electric lights, waiting for those 
that fall." 

This species can at once be distinguished from I. arnoldi 
Hewitt} which it somewhat resembles, and which also is 
believed to occur at Bulawayo, through the total absence of 
a scopula on the fourth tarsus, whereas arnoldi has a broad 
scopula on the swollen fourth tarsus. 

Colour. — Ui3per surfaces brownish-black, lower surfaces 
somewhat paler, the sternum and coxa3 of the third and 
fourth legs, the genital sternite, lung opercula and spinnerets 
being pale yellowish-brown. 

' ' Records, Albany Museiim,' vol. iii, p. 21, 1914. 



NEW SOUTH AFRICAN ABACHNEDA. 



81 r 



Carapace. — Broad and depressed, its length equal to that 
of the fourth metatarsus together with two-thirds of the 
tarsus. Surface coai'sely granuUxted, except on the ocular 
area and lateral thei'eto. Ocular area very slightly wider 
than long, its length slightly less than one-third of the 
distance from the anterior margin of the carapace to the 

Text-pig. t>. 





A. Idiops astutus sp.nov. b. I. arnoldi Hewitt. 
Distal portions of male palps, a x 14. b x 18. 



centre of the fovea ; frontal eyes quite sepai'ated, about one 
and a quarter diameters apart. Area formed by frontal and 
anterior median eyes very slightly wider in front, its length 
being only slightly greater than its anterior width ; the 
medians about half a diameter apart and much larger than 
the frontals. Posterior margins of posterior row in a pro- 
curved line, the medians about two to two and a half dia- 
meters apart and about a diameter distant from the laterals, 
which are elono-ated but rather small. Posterior medians 



312 JOHN HEWITT. 

about one-third of a diameter distant from the anterior 
medians. 

Pedipalp . — Tibia slightly more than three times as long as 
deep, the excavation completely bordered by a band of spines. 
Tarsus dorsally without spines, except at the apex, where there 
are several; the apical portion strongly projects upwards (see 
text-fig. 6, a). Process of bulb flattened throughout its length, 
obliquely truncate at the apex, curved and slightly twisted. 

Legs. — Tibia I shorter than the metatarsus, with a pair 
of tubercles near the distal end inferiorly, the proximal one 
short and small, the distal one with a large and long black 
projection ; its under-surface without spines along the median 
area, about ten or eleven along the posterior edge inferiorly, 
but none on the anterior edge. Metatarsus I distinctly 
bowed, with seven or eight short spines inferiorly on the 
posterior side and three or four on the anterior side (includ- 
ing two near the apex). Tarsus I with eight or nine spines 
on each side, weakly scopulate in its distal third, II and III 
broadly scopulate from apex to base, IV without trace of 
scopula and not swollen. Claws of all the tarsi with two 
basal teeth, the more distal one larger than the other ; on 
tarsus IV the posterior claw of the left leg (missing on I'ight 
side) carries three teeth, and such is the case on one of the 
claws on tarsus II. Band of spinules on anterior side of 
patella IV extending the whole length of the segment ; 
patella III also has a band of short spines along its anterior 
surface, ending in a row of about live or six longer ones 
along the distal edge. 

Measurements. — Total length 19 mm., length of cara- 
pace 8"8 mm., breadth of same 7"8 mm., length of tibia of 
first leg 5*4 mm., of metatarsus of same 7'4 mm., length of 
first leg 33"3 mm., of fourth leg 31 mm. 

Hermacha mazoena sp. nov. 

The type is a single female specimen from Mazoe, Mashona- 
land (J. Darling), in the collection of the British Museum. 



MEW SOUTH AFRICAuV AKACHNIDA. 313 

Colour. — Almost uniformly pale brown, the abdomen 
superiorly showing indication of a darker tree-pattern. 

Carapace. — Slightly longer than the fourth metatarsus 
and tarsus together. Ocular area a trifle more than twice as 
broad as long. Anterior margins of anterior row of eves in a 
procurved line ; the medians rather small, about three- 
quarters of a diameter apart. Posterior medians rather 
small, oval, and widely separated, about three and a half 
times their long diameter apart, their inner margins about in 
a line with the outer margins of the anterior medians ; pos- 
terior laterals about as long as the anterior laterals. Fovea 
straight. 

Chelicera3. — With seven teeth in the inner row below, 
the distal tooth of the outer group being opposite the fifth 
tooth of the inner row. 

Pedipalps.— CoxfB with about sixty teeth arranged in a 
triangular patch at the base. 

Labium. — Quite without apical teeth. 

Legs. — Tarsus I with dense undivided scopula, II with 
scopula divided by a thin median strip of setas, III and IV not 
definitely scopulate, but carrying a few scopular hairs at the 
sides, in IV only distally so. Metatarsus I and II scopulate 
to the base. III and IV not scopulate ; I with two spines at 
the apex below and two or three along the inferior surface, 
also one or none on the anterior surface. Tibia I with three 
weak spines at the apex below and two on the inferior surface, 
also one on the anterior surface superiorly ; very slightly 
exceeding the metatarsus in length. 

Posterior Sternal Sigilla. — Elongated, almost touch- 
ing the sternal margin. 

Posterior Spinners. — With the apical and basal seg- 
ments subequal in length, being about one and three-fifths 
times the length of the penultimate segment. 

Anterior Spinners. — Not widely separated, less than 
half their own length apart (the abdomen is somewhat con- 
tracted, and in fresh specimens the spinners may perhaps be 
rather more widely separated) ; moderately long, being about 



314 JOHN HEWITT. 

four-fifths the length of the basal segment of the posteinor 
spinners. 

Measurements. — Total length 15"2 mm., length of cara- 
pace 5"6 mm., breadth of same 4'5 mm., length of tibia of first 
leg 2 '5 mm. 

No species of this genus has previously been recorded from 
Khodesia. The described species to which it is most closely 
allied are probably H. caudata Sim. from Delagoa Bay, and 
H. bicolor {Poc.) [Brachytheliscus bicolor Poc.j from 
Durban ; the female of the former is unknown, and the latter 
species is distinguished from mazoena in the following 
characters : ocular area, dentition of chelicerte and of labium, 
and separation of anterior spinners. 

Genus Hbrmachola gen. nov. 
This name is proposed for the reception of a small species 
which has many of the characters of the genus Hermacha 
E. Sim., but differs therefrom principally in the form of the 
male bulbal organ and in the shape of the fovea as hereafter 
described. 

Hermachola grahami sp. nov. Text-fig. 7, a-c. 

The type is a single adult male example collected at 
Grrahamstown by Mr. F. C. Graham, September 23rd, 1914^ 
who presented it to the Albany Museum. 

Colour. — Carapace and appendages pale brown, the more 
distal segments of the first pair of legs with a dull reddish 
tinge ; the hairs which clothe the carapace and appendages 
are blackish . Abdomen pale dorsally, with a variegated dark 
pattern. Lower surface pale, but posteriorly just in front of 
the base of the spinners the abdomen has a dark transverse 
stripe. 

Carapace. — Elongated, about as long as the metatarsus 
and tarsus of the fourth leg. Fovea transverse at the base, 
but with a short backwardly directed median extension, which, 
however, is not so deep as the main portion of the fovea. 
Near the anterior border of the fovea there are two long spini- 



NEW SOUTH AFRICAN AEACHNIDA. 



3] 



form seta3 mesially situated; otherwise no spiniforin sette 
occur on the carapace except a few on the borders, especially 
at the postero-lateral corners. 

Ocular Area. — Not quite twice as broad as long. Ante- 
rior row with their anterior margins strongly procurved, the 
laterals much larger than the medians or than the postero- 
laterals. Posterior laterals subcontiguous with the posterior 

Text-fig. 7. 




Hermachola grahami sp. nov. 
A. Distal portion of male i^alp. X 55. B. Fovea, x 40. c. Ocular 

area, x 65. 



medians and with the anterior laterals. Distance of anterior 
laterals from the mat'gin of the carapace scarcely as much as 
half of a diameter of an anterior median. 

Labium. — With a single small apical tooth. 

Chelicerje. — With seven or eight teeth on the inner row 
below. Anterior surface covered with spiniform seta? which 
are not elongated; rastellum weak, without spines. 

Pedi palps. — Coxa with a small patch of about eighteen 
small teeth at the base. Femur with two or three weak spines 

VOL. 3, PART 2. 22 



316 JOHN HEWITT. 

above ; patella without spines. Tibia with numerous elon- 
gated spiniform setae on its ventral surface on the inner side, 
more strongly developed in the basal half of the segment- 
Tarsus without spines. Bulb small, process coiled up spirally 
like a coi-kscrew ; there are two complete coils in addition to^ 
the expanded and coiled basal portion which invests the bulb 
and the narrowed distal portion which also has a strong spiral 
twist ; the process reaches backwards as far as the basal 
fourth of the tibia. 

Legs. — Tarsi unspined. Metatarsus I almost straight, 
with three sti'ong apical spines below but no other spines 
elsewhere, II also with three strong apical spines inferiorly, 
and in addition with two or three on the inner surface and 
two on the inferior surface posteriorly, III and IV numerously 
spined. Tibia I stout, with three apical spines, none of which 
are enlarged, also with six spines on the lower surface, of 
which one distally situated is specially long and stout ; on the 
anterior surface there is a row of three spines. Tibia II Avith 
two or three apical spines inferiorly, three or four on the 
lower surface, and two or three on the anterior sui'face supe- 
riorly ; III with three apical spines below, four on the inferior 
surface, two on the anterior surface, two dorsally, and two on 
the posterior surface above, all of them being elongated ; IV 
spined like III except that the dorsal sui'face has one or no 
spines. Patella I without spines, II with one on the anterior 
surface near the apex. III with two on the anterior surface 
and one on the posterior surface, IV with one on the anterior 
and one on the posterior surface. Femora armed above with 
long setiform spines. Tarsi I and II scopulate to the base, 
the scopula being entire, though not dense, and composed of 
not very fine sette ; III and IV with a few scopular set^e, but 
mostly the setae are long and subspiniform. Metatarsi I and 
II thinly scopulate from the apex to near the base. Inferior 
tarsal claw of anterior legs small but easily distinguishable. 

Sternal sigilla. — Third elongated, slightly less than its 
own length distant from the sternal margin ; second sub- 
marginal, first indistinct. 



NEW SOUTH AFRICAN ARACHNfDA. 317 

Measurements. — Total length 8*25 mm., of carapace 3 
mm., width of carapace 2'1 mm. 

Stasimopus steynsburgensis sj). nov. 

This species is closely related to S. palpiger Poc. from 
Graaff Reinet and S. schreineri Pure, from Hanover. It 
differs from both in having shorter palps and in possessing a 
scopula on the tarsus of the fourth leg of the male. 

The type is a single adult male in the collection of the 
British Museum from Steynsburg, C.C., taken by Miss Leppan. 

Colour. — Upper surfaces blackish, excepting the tibia and 
tarsus of the palps and the tarsi and metatarsi of the legs, 
which are reddish yellow ; also the patella and tibia of the 
fourth legs are brownish ; sternum and coxfe of appendages 
inferiorly castaneous, abdomen fuscous, the lung opercula and 
genital sternite yellow, the spinners pale. 

Carapace. — Subequal in length to the metatarsus of the 
fourth leg. The three keels well developed anteriorly, but 
absent posteriorly, none of them approaching the fovea ; hairs 
are absent therefrom and from the ocular area; the sides 
of the cephalic area are practically smooth except immediately 
external to the lateral ridges. Anterior margins of anterior 
row of eyes distinctly procurved ; distance between an 
antei'ior lateral and anterior median subequal to the diameter 
of the latter, but less than the long diameter of the former; 
posterior lateral almost as long as the anterior lateral, the 
distance between them about one and three-fifth times the 
length of the latter ; outer lateral margin of anterior lateral 
about in a line with inner posterior margin of posterior lateral. 

Pe dip alps. — Stretched forwards, the apex reaches a point 
three-fifths of the distance along the metatarsus of the first 
leg. Patella longer than that of first leg, but much shorter 
than the tibia of that leg or of the palp. Tibia and tarsus 
together slightly shorter than the carapace. Process of 
palpal organ longer than the bulb, tapering and slightly 
twisted. 

Legs. — Tarsus I absent from the specimen; II a little 



318 JOHN HEWITT. 

swollen, scopulate below, anteriorly with six spines, posteriorly 
with seven or eight spines ; III with about fourteen spines on 
each side, scopulate below; IV numerously spined on both 
sides and scopulate mesially below. Metatarsus I and II 
without trace of a scopula, but with no spines mesially below 
though strong spines occur on each side inferiorly. Tibia I 
shorter than the metatarsus, with only one or two spines 
mesially below and with about eight spines on the anterior 
side and four on the posterior side in addition to those at the 
apex; III with five to seven short spines on or near the 
distal edge on both anterior and posterior sides superiorly. 
Patella I with one spine at the apex inferiorly ; III with an 
anterior band of short spines, about six to eight in number, 
but no distal patch of spinules superiorly ; TV with a patch 
of minute spines extending over about two-thirds to three- 
quarters of the length of the anterior side. Anterior claAv of 
fourth leg with two large teeth basally situated and with no 
small ones distal thereto ; posterior claw with three large 
basal teeth. 

Measurements. — Total length 18'5 mm., length of cara- 
pace 7*8 mm., breadth of carapace 7 mm., length of tibia of 
palp 4'2 mm., of metatarsus of palp 5'6 mm. 

Stasimopus gigas sp. nov. 

The type is a single male example in the collection of 
the British Museum labelled "A^redefort Ed.?, Barrett- 
Hamilton " ; the specimen is in dried condition. 

Colour. — Upper surfaces black, except the tarsi and meta- 
tarsi of the legs, the tibiae and tarsi of the palps, also the distal 
portions of the tibite of the first two pairs of legs, and of the 
patellae of the palps, all of which are reddish. 

Carapace. — Subequal in length to the metatarsus of the 
fourth leg. The three keels are prominent throughout and 
reach back almost as far as the fovea. Anterior margins of 
anterior row of eyes in a very slightly procurved line, sub- 
equally spaced, the distance between the medians about one 



NEW SOUTH AFRICAN AEACHNIDA. 319 

and a half times the diameter of an eye; innei- edges of 
anterior lateral and posterior median practically in the same 
line; distance between anterior and posterior laterals quite 
one and a half times that between an anterior lateral and 
anterior median. 

Pe dip alps. — Patella slightly longer than that of the fii'st 
leg, shorter than the tibia of that leg or of the palp. Tibia 
and tarsus together considerably shorter than the carapace. 

Legs. — All the tarsi scopulate below; no trace of a scopula 
on the metatarsi. Tarsus I with a single spine on its anterior 
side and four on the posterior side (II wanting in the speci- 
men) ; III with numerous spines in a continuous band on both 
anterior and posterior sides; IV with numerous spines on both 
sides, especially anteriorly. Metatarsus I with strong spines 
below, both over the median area and at the sides. Tibia I 
considerably shorter than the metatarsus, with strong spines 
below and at the sides ; III with a few short spines and 
spinules on or near the distal edge on both anterior and 
posterior sides superiorly. Patella I with two spines at the 
apex inferiorly and one on the lower surface; III with an 
anterior band of spines, but no distal patch of spinules above 
apart from those belonging to that band ; IV with an anterior 
patch of spinules extending about three-fifths of the length of 
the anterior side. Paired claws of tarsus of fourth leg with a 
basal comb of four large teeth, distal to which is a very small 
tooth, and on the posterior claw one or two minute teeth occur 
on the proximal side of the comb. 

Measurements. — Length of carapace 11*2 mm., breadth 
of same 10 mm., length of tibia of palp measured from above 
7 mm., of patella of palp 5"5 mm., of fii'st metatarsus 8*4 mm. 
of first tibia 7 mm., of first patella 5 mm., of fourth metatai-sus 
11*5 mm. 

This species is distinct from S. nigellus Poc. (Ann. Mag. 
Nat. Hist., 7, x, p. 319), which also seems to have been taken 
at Vredefort Rd. by the late Capt. Barrett-Hamilton, in the 
following characters : ocular arrangement, keeling of cara- 



320 JOHN HEWITT. 

pace, spinulation of first metatarsus, third tarsus, third patella, 
in the dentition of the fourth claws, and lastly in size, this 
being by far the largest male yet recorded in this genus. 

Stasimopus minor sp. nov. 

The type is a single adult male from Bloemfontein collected 
by Dr. T. F. Dreyer in August, 1914. It was found on the 
open veld. 

This may be identical with 8. oculatus Poc, which is 
based on a female example, or may belong to another species, 
for Dr. Dreyer has taken females of two species at Bloem- 
fontein. It is probably closely related to S. nigellus Poc. 
taken at Yredefort Rd., but differs therefrom in the Avide 
separation of the anterior median eyes. 

Colour. — Jet-black above, the distal portions of the legs 
and of the palps paler ; sternum, coxse of appendages, genital 
plate, lung opercula, and spinners pale. 

Carapace. — A little longer than tibia of first leg or 
metatarsus of palp. Almost quite smooth and shining in 
its anterior half, lightly sculptured (not coarsely rugose) 
in its posterior half. The three keels depressed, only the 
median one reaching back to the fovea. Anterior margins 
of anterior row of eyes in a straight line, the medians much 
nearer to the laterals than to each other, being about one and 
a half diameters apart; anterior laterals subequal in length to 
the anterior medians and separated therefrom by about half 
the diameter of a median. Ocular area sparsely hairy. 

Pedi palps. — Pressed forwards, the tip reaches a point 
three-fifths of the distance along the metatarsus of the first 
leg ; patella considerably longer than that of the first leg, but 
shorter than the tibia of that leg or of the palp ; tibia and 
tarsus together subequal to the carapace in length. 

Legs. — All the tarsi scopulate below, but no trace of a 
scopula on the metatarsi. Tarsus I with two or three spines 
on the anterior side, two or none on the posterior side ; II 
with two anteriorly and two or three posteriorly ; III with 



NEW SOUTH AFRICAN AllACHNlDA. 321 

one anteriorly and one or three posteriorly ; IV with seven 
anteriorly and one posteriorly. Metatarsus I without strong 
spines on the mesial area below. Tibia I subequal in length 
to the metatarsus, with three spines below besides those at 
the apex, but none at the sides excepting one near the apex 
anteriorly ; tibia III with about five short spines on or near 
the distal edge on both anterior and posterior sides superiorly. 
Patella I without spines below; III with a sti'ip of six to nine 
Aveak spines on its anterior side, but no distal patch of spinules 
^bove. Patella IV with an anterior patch of spinules extending 
about three-fifths of the length of the anterior side. Paired 
tarsal claws of fourth leg each armed Avith two large basal teeth, 
Avith four smaller ones more distally situated on the posterior 
<;laAv, but only three on the anterior claw, Avhich are weaker 
than those on the posterior claAv. 

Measurements. — Total length 8'5 mm., length of cara- 
pace 4 nnn., breadth of same S'o mm., length of palp 12 mm., 
of first leg 15 mm. 

Stasimopus oculatus Pac. 

Stasimopiis oculatus Poc, Proc. Zool. Soc, 1897, p. 728, pi. xlii. 

fig. 2. 

It has been pointed out to me by Dr. T. F. Dreyer that at 
least tAvo species of Stasimopus occur at Bloemfontein, and 
Dr. Dreyer has kindly sent to me adult female material of the 
same. 

S. oculatus, the larger species, can be distinguished 
through the presence of a cluster of spines at the apex of the 
third metatarsus inferiorly and the presence of stout spinules 
at the apex of the tibia of the palp superiorly, whereas the 
smaller species has neither oi these characters; there is also 
a difference in coloration, the smaller species having the 
abdomen fairly uniformly infuscated above, Avhilst oculatus 
has the lateral surface of the abdomen and most of the dorsal 
surface comparatively pale but Avith a dark mesial blotch 
anteriorly and some infuscation OA'er the posterior part of the 
dorsal surface. 



322 JOHN HEWITT. 

Female specimens which seem to be identical with oculatus 
are known to me from Redclersbui-g (Dr. Broom), Ladybrand 
(Dr. Dreyer), and Kimberley (Bro. J. H. Power). 

Female specimens which are identical with or very closely 
related to the smaller Bloemfontein species are known to me 
from Vredefort Rd. (Capt. Barrett-Hamilton), Yalsch River 
near Ki-oonstad (Prof. H. H. W. Pearson), and Winburg- 
(Miss S. Brown). 

A third form characterised by the possession of short stout 
spiniiles at the apex of the tibia of the palp but no spines at 
the apex of the third metatarsus occurs at Kroonstad (Dr. 
T. F. Dreyer). 

A fourth form from Jagersfontein (M. Francis) has a group 
of stout spines at the apex of the third i»etatarsus, but no 
stout spinules at the apex of the tibia of the palp. 

Lastly, we have a good series of female specimens from 
Modder River (Bro. J. H. Power) which seems referable to two 
species, all the small and immature examples agreeing with 
oculatus, and all the larger ones being similar to the smaller 
Bloemfontein species in structure, but very much larger than 
that form; in this case it seems just possible that the small 
specimens from Modder River are merely the young of the 
larger ones.^ 

Unfortunately we possess no evidence wliich can serve to 
connect any of these forms with the species based on male 
material. Only four male Stasimopi are known from the 
Free State, viz. the type of S. nigellus Poc. from A^redefort 
Rd., the type and a co-type of S. gigas sp. nov. also from 
Vredefort Rd., and the type of S. minor x/'. noi-. from 
Bloemfontein. 

According to Dr. Dreyei', the nests of the two Bloemfontein 
species are very distinct; the female of oculatus makes 
a D-shaped lid, whilst the other species has a more circular 
lid; the tube of the nest in oculatus is very thickly lined 

' Some of the forms just mentioned will be descrilied l>y me as new 
species in ' Records Albany Museum,' vol. iii, pt. ii. 



NEW SOUTH AFRICAN ARAGHNIDA. 323 

"by felt-like silk, forming a much thicker wall than that found 
in the other species. In a large specimen the longest diameter 
of the lid of oculatus reaches 50 mm., the greatest outside 
diameter of the tube at the top being 60 mm. 

Order SOLIFUG^. 

Chelypus hirsti b-j). nor. Text-figs. 8, 9. 
Text-fig. 8. 




\ \ 




^ ,i 



% 



1 
















% 


1* 


V' 


[m^ 



Clielypus hirsti S2). r.ov. 

Specimen on left is minus the left chelicerii. and is shown in 

ventral view ; specimen on right is in dorsal view, x f . 

The types consist of four male examples in the Albany 
Museum collection, two of which came from Rietfontein, 
Goi'donia, presented by Mr. H. Drew, and two are indefinitely 
located as North- West Gordonia, presented by Mr. C. A. 
Anderson. Female unknown. 

This species is closely related to C. barberi^ Purccll, but 
differs therefrom as follows. 

1 Pm-cell, W. F.. 'Annals S. Afr. Mus.." vol. ii. p. 2-24. ISKJl. 



324 JOHN hj:witt. 

Flagelluin. — Not bifid at the apex, but tapering to a point. 
Near its base the shaft has a ventral, compressed, and keel- 
like extension along the greater part of the length of its pro- 
current portion. 

Chelicerae. — Upper finger furnished internally with 
numerous short stout spines, occupying an elongated obliquely 
arranged area extending from the neighbourhood of the large 
basal tubercle on the inner side of the fang to a point near 
the basal enlargement of the iiagellum. Outer row of teeth 
in the upper jaw comprising six teeth, the fourth one being 

Text-fig. 9. 




Clielypus liirsti sj). nov. 
Left chelicera of male viewed from inner side, x 4. 



minute. Upper surface of cheliceras covered with small 
granules, a few of which are spinuliform. 

Pe dip alps. — The upper surface of the tibia is only finely 
granulated, and in its basal third is almost free of granules ; 
there is some fine granulation and one or two coarse granules 
at the base of the metatarsus above. 

Colour. — Cheliceras without dark marks at the base above. 
Cephalothorax infuscated over its anterior half. 

Total Length. — 27 mm. 

This species is named after Mr. A. S. Hirst, who has made 
important contributions to our knowledge of Arachnida and 
-has described various S. African species. 



NEW SOUTH AFRICAN ARACHNIDA. 



325 



Order SCORPIONES. 

Opistliophthalmus pugnax Ihorell, var. natalensis 



var. nov. 



The types of this variety are two adult specimens, male and 
female, from Estcourt, Natal, collected by Mrs. E. J. Turner 
and by Mr. Gruy Marshall respectively, who presented them 
to the British Museum. The Natal Museum has a female 
example, a trifle larger than the type, from Mooi River, Natal, 
collected by Mr. C. James. 

The variety agrees with the typical form of pugnax, 
as known to me through material from Pretoria, in colour and 
general structure, but diifers as follows. 

No stridulatory lamellee on the chelicerte. 

Median eyes more posteriorly situated, their distance from 
the hind end of the carapace being less than one-third of the 
total length of the carapace (more than one-third in typical 
pugnax). 

Hands much less coarsely granulated, that of the male on 
its inner portion superiorly being covered with numerous quite 
small and isolated round granules, the finger keel with one or 
two coarse pits in its course, but practically continuous 
throughout, the more external secondary keel quite obsolete 
and the inner one, for the greater portion of its length, only 
represented by infuscated scarcely enlarged granules, the 
outer surface of the hand with only one keel ; that of the 
female without coarse granulation on the inner part of its 
upper surface, except quite near the finger, but covered with 
a much flattened meshwork of more or less coalesced ridges 
and granules in which the secondary keels are quite absent, 
the whole surface appearing much smoother than in the 
typical form, the finger keel well defined, continuous in the 
distal half, more or less broken in the basal half (in the Mooi 
River example it is continuous almost throughout). 

Hand of male slightly narrower in proportion to the length 
of the hand back than in pugnax, and the fingers a little 
lono-er. 



326 JOHN HEWITT. 

The granules on the mesial portion of the last abdominal 
sternite o£ the male more numerous and not quite so large as 
in the male of pugnax proper (about ten or twelve rows can 
be roughly counted between the anterior and posterior 
margins) ; in the female this area is covered with low granules 
and irregularly shaped ridges, which occur throughout the 
whole length of the segment, more or less obscuring the keels 
(these are quite obsolete in the Estcourt specimen, but present, 
though indistinct, in the Mooi River specimen). 

Infero-median keels of first caudal segment in the male com- 
posed of about eight to ten coarse granules, and the space 
between them also includes about ten coarse granules, but is 
not obliterated thereby nor are the crests rendered indistinct 
to any great extent ; in the female these keels are pitted and 
more or less coarsely granular (in the Mooi River specimen 
the keels are not definitely broken up into coarse granules), 
with a few granules between the keels. 

The abdominal tergites in the female are all granular 
throughout, coarsely so in the posterior portions of each tergite. 

Pectinal teeth : Thirteen to fourteen male, eleven to twelve 
female (thirteen to foux-teen in the Mooi River specimen). 

Measurements. — Total length, male 78 mm., female 89 
mm.; length of hand, male 17"3 mm., female 18'8 mm.; of 
movable finger, male 11"75 mm., female 12"3 mm.; of hand 
back, male 6 mm., female 7 mm.; breadth of hand, male 8 mm., 
female lO'b mm.; length of cai'apace, male 11*7 mm., female 
18 mm.; distance of median eyes from posterior margin, male 
3"75mm., female 3'7 mm.; width of first caudal segment, male 
5"75 mm.; of fifth caudal segment, male 4*25 ram.; length of 
same measured along the side 10 mm.; breadth of vesicle, 
male 4'25 mm.; length of same 10 mm. 

In respect to the shape of the hand and the granulation of 
the last abdominal sternite and of the infero-median keels of 
the first caudal segment in the male, this form is intermediate 
between typical pugnax and the iorm described by me from 
Keilands under the name of 0. latimanus Koch var. 



NEW SOUTH AFRICAN ARACHNIDA. 327 

keilandsi.^ It resembles that form also in the posterior 
position of the median eyes ; it differs in that the tail of the 
male is much stouter in natal en sis. The fifth caudal 
segment in an adult male of keilandsi measures 10*5 mm. in 
length and 3'75 mm. in width. 

Dr. Purcell " points out in his important monograph on the 
genus Opisthophthalmus that the presence or absence of 
stridu-latory lamellee on the chelicerge is usually of no specific 
value. However, I am inclined to regard the character of 
some importance in the various forms which range around and 
between pugnax and latimanus. Such lamellas are 
present in all our examples of latimanus, 132 specimens of 
all sizes, from Grahamstown, Highlands, Brakkloof, Fort 
Beaufort, Jansenville, Schurfteberg, Alicedale, Cllen Lynden, 
Mimosa, Redhouse, Alexandria, and Queenstown ; they are 
even present as a row of bristles on the newly hatched young. 
They are present in latimanus var. austeroides mihi and 
in every specimen of our series of typical pugnax; they are 
completely absent in keilandsi. 

It is not impi'obable that intermediates between keilandsi 
and natal en sis will be found, in which case it will no longer 
be possible to maintain pugnax as a species distinct from 
latimanus. With the discovery of new local forms in the 
genus Opisthophthalmus the difi^erences between the so- 
called species ai'e gradually breaking down and the genus is 
seen to be composed of numerous local forms which more or 
less completely grade into each other. 

' ' Records, Albany Musextm,' vol. iii, p. 7, 1914. 
- ' Annals S. Af r. Museum,' vol. i, p. 134, 1899. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 329 



Observations on some South African Termites. 

By 

Ciaiido Fuller, 

Division of Entomology, Department of Agriculture, Pretoria. 



With Plates XXV— XXXV and 16 Text-fiffures. 



Contents. 

I. Introduction 

II. The Abundance of Termites 
III. The Behaviour of Winged Termites 
Hodotermes transvaalensis s|). n. 
Termes natalensis Havlland 
Termes latericius Haviland 
Termes vulgaris Haviland 
Termes incertus Hagen . 
Eutermes bilobatus (Haviland) and 

IV. Observations on the Nesting Habits 
Economy of Certain Species . 
Hodotermes transvaalensis sp. n. 
Hodotermes viator (LatreiUe) 
Calotermes durbanensis Haviland 
Rhinotermes putorius Sjostedt . 
Termes waterbergi sp. n. 
Termes natalensis Haviland 
Termes badius Haviland 
Termes latericius Haviland 
Termes vulgaris Haviland 
Termes incertus Hagen . 
Eutermes parvus {Haviland) 
Eutermes bilobatus {Haviland) . 
Eutermes trinervius (Bambur) 



Eutermes sp 
and General 



PAGE 

330 
381 
333 
335 
336 
339 
340 
341 
346 

349 
349 
356 
358 
360 
361 
364 
377 
385 
391 
393 
396 
398 
402 



330 



CLAUDE FULLER. 



V. Classification 

(1) Introduction 

(2) Systematic Account 

Hodotermes 

Hodotermes transvaalensis sp. n. 
Hodotermes pretoriensis sjj. n. 
Hodotermes karrooensis sp. n. 
Hodotermes mossambicus {Hagen) 
Hodotermes havilandi Sharp 
Hodotermes warreni sp. n. 
Hodotermes braini sp. n. 
Hodotermes viator (Latreille) 
Calotermes durbanensis Havilancl 
Rhinotermes putorius Sjostedt 
Termes 

Termes swazise §2^. /i. 
Termes waterbergi sp. >i. . 
Termes natalensis Havilancl 
Termes badius Havilancl 
Termes later icius Havilancl 
Termes vulgaris Havilancl . 
Termes incertus Hagen 
Enter me s parvns (Haviland) 
Entermes bilobatns (Haviland) 
Entermes liastatns (Haviland) 
Enter mes trinervins {Bambur) 
VI. Appendix — Glossary 
VII. References 
VIII. Explanation of Plates 



PAGE 

. 413 

. 413 

. 423 

. 423 

. 425 

. 429 

. 433 

. 438 

. 441 
444 

. 446 

. 448 

. 451 

. 453 

. 456 

. 462 

. 466 

. 470 

. 475 

. 479 

. 481 

. 483 

. 485 

. 487 

. 489 

. 491 

. 494 

. 496 

. 497 



I. INTEODUCTION. 

These notes do not pretend to be more than fragmentary 
records which have been brought together for publication at 
the express wish of my friend, Dr. Ernest Warren. They 
include observations which have been made or collected only 
during the past two or three years ; for most of them the 
writer is personally responsible; but, for some, which are duly 
acknowledged in the text, he is indebted to his colleagues 
and other friends. Unfortunately, owing to the difficulties 



OBSKRTATIONS ON SOME SOUTH AFIilCAN TERMETPJS. o31 

experienced in determining the material collected during the 
period mentioned, it has not been possible to include all the 
observations made ; these it is hoped to deal with at some 
futui-e date. 

African termites having been described somewhat inade- 
quately, as a whole, and for the most part in foreign 
languages, the opportunity has been taken to redescribe the 
soldier caste of some species discussed, and to furnish some 
comment upon the geoieral characteristics of each. 

As far as possible those speculations, which the study of 
termite-nature excites in every observer, have been avoided ; 
here and there, however, the temptation to speculate has been 
too great to overcome. 

For the sake of completeness, some observations made by 
others are repeated ; these are chiefly derived from the 
Haviland notes (1) or Dr. Sjostedt's monograph (2), and the 
context indicates whether or not they have been confirmed. 
The opportunity is also taken to correct some misapprehen- 
sions the author was under in writing his notes, " White Ants 
in Natal" (3). 



II. THE ABUNDANCE OF TEEMITES. 

Owing to the fact that a number of South African species 
are to a great extent subterranean in their hal)its, little idea 
exists as to the great abundance of termites in general. This 
abundance may not be widespread, but it may be said of 
many parts of the country that the soil teems with termites. 
In some parts many species are found whose subways interlace 
without anastomosing, and all find sustenance in the same 
pastures. Where, in other parts, kinds are fewer, at least 
one sort tends to be particularly abundant. Again, upon the 
rock-sti-ewn hills and mountain-sides a number of species, 
making their nests under stones, find a congenial environ- 
ment. 

As only a few points in the Union have been at all explored 
VOL. 3, PART 2. 23 



'o)\< 



vC^)V>>r A fi &^cv/ 



332 CLAUDE FULLER. 

for termites, and none very thoroughly, these generalities call 
for some verification. 

In the south-west Cape, where termites are regarded as 
rare and Avhere, as at Cape Town, white-ant attacks upon 
wooden structures are practically, if not wholly, unknown, 
Hodotermes viator exists abundantly, and a brief search 
in the neighbourhood of Stellenbosch, made for the writer 
by Mr. C. P. van der Merwe, revealed this and three other 
species. 

In the Grreat KaiTOO Hodotermes karrooensis is almost 
ubiquitous and, in restricted areas. Enter me s trinervius 
also abounds ; builders of large mounds, other than the last- 
mentioned insect, are seemingly non-existent, but there is 
little evidence on this point, and nothing is known of subter- 
ranean and rock-nesting forms. 

Over the whole of the central area of the Orange Free State, 
the abundance of Eutermes trinervius is phenomenal ; 
other forms are reported from various parts. 

In Natal and the Transvaal it is no exaggeration to say that 
the soil is riddled from end to end of the country with termite 
tunnellings, through which an inconceivable host of insects 
constantly passes to and fro. 

In two small areas at Pietermaritzburg (784 square yards 
each), the writer reported (3) finding fourteen and sixteen 
nests respectively, the homes of six dilferent species. This, 
however, was an observation based solely upon surface indi- 
cations. Latterly, a favourable opportunity presented itself to 
observe subterranean conditions at Pretoria in the case of 
extensive excavations which were being made for the founda- 
tions of certain large Grovernment buildings. These works, 
covering two to three acres, were visited almost daily, and 
many of the observations detailed later were made during their 
progress. It was found that the soil was inhabited by eight 
species ; to a depth of 4 to 5 ft. it was riddled with their 
galleries, and not a cubic yard existed which did not contain 
one or more cavities belonging to one species or another. 



OBSERVATIONS ON SOME SOUTH AFRICAN TEKMITES. 333 



III. THE BEHAVIOUR OF WINOED TEEMITES. 

It may be taken as an axiom that every colony of the genus 
Termes is originally founded by the unaided efforts of a pair 
of winged insects. There is no acceptable evidence to the 
contrai'y ; and the oft-quoted presumption that all adult 
termites are helpless and dependent upon adoption by a section 
of wandering workers is based upon the merest conjectui'e. 

Again, the aerial flight is often regarded as having for its 
chief objective the prevention of inbreeding ; that it may fail 
in this connection is indeed very obvious, although it must be 
admitted that the possibility of insects from different colonies 
mating is sometimes present. The real object of the production 
of sexual individuals in such vast numbers and their annual 
«xodus seems only to be for the purpose of perpetuating the 
species ; the countless swarm of emigrants which emerge, most 
of them to meet Avith speedy death, has its parallel in the 
phenomenal number of eggs or of young produced by many 
organisms. 

In the case of two species many individuals have been seen 
to mate and at once to seek shelter in the earth within 6 to 
8 ft. of the spot from which they had just emerged. Again, 
with these two species, and others, it has been noticed on 
occasion that an aerial flight takes place from one nest and not 
from others of a like kind adjoining it, and that insects mating 
far afield were of the same colony. At times, vast numbers of 
this species or that, or several kinds together, take to the wing 
simultaneously over a great area of country ; it is on such 
occasions as these that sexes of different parentage meet. At 
other times vast numbers emerge in restricted localities only, 
or from one nest only. 

The flying powers of winged termites do not appear great, 
and the majority do not fly far. At the same time, even in 
the case of those displaying the smallest powers of endurance, 
some few at least far outdistance the majority. Clumsily as 
they seem to fly many are well able to direct their movements 



334 CLAUDE FULLER. 

and i-apidly avoid obstacles placed suddenly in tlieiv way. 
As might be expected, those which take to the wing after dark 
are readily attracted by light. 

It seems almost certain that this phase in the history of a 
termite is a succession of events which follow upon one another 
in regular order, and, if the chain is broken at any point, it 
remains so. This may be illustrated by several specific cases. 
The night-flying Eutermes trinervius, if attracted by illu- 
minating a white sheet, alights, and both sexes dealate after 
meeting. If some are captured at once before meeting and 
placed in a tube, they do not dealate. Some, so captured, 
have been kept over four days alive in confinement, 
and whilst a few dealated, or to be more correct lost their 
Avings, no attempt at pairing was made. The same holds good 
for the dusk-flying Termes incertus, but to a modified 
extent only. Further, with these two species and three others 
(undetermined) no attempt is made at burrowing if the sexes 
are kept separate. In the case of two males confined together 
it was noticed that one would frequently follow the other as if 
it were a female. The stimulation induced them to burrow 
together, but they soon desisted from doing so, although 
stimulated thereto several times by one another ; ultimately 
they died on the surface of the soil. Again, a number of pairs 
of T. natal en sis were captured one evening and placed in a 
small box ; in the morning it was found that the couples had lost 
all regard for one another ; they were allowed to wander over 
a table, but none again mated voluntarily. When, however, 
the sexes were sorted out and the male placed in the position 
it had been in when captured, the sequence of events was re- 
established. In a further instance six pairs of Hodo ter- 
mes transvaalensis Avere collected (December 30th, 1913) 
from burrows which they were making at the time, the couples 
were then placed in jars of soil, but were found to have lost 
the instinct to start burrowing again. They wandered aimlessly 
about, the males taking no notice of the females. When, 
however, artificial bum'ows were made, and the pairs placed in 
them side by side, the environment reacted upon them at once 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 335 

and they burrowed to the bottom of the jar. In 1914, similar 
results attended the mating and re-mating of this species. 

The act of pairing appeai-s to differ in the details of its 
manifestation with each species, as the following accounts 
concerning several species will show. 

Hodotermes transvaalensis .sp. n. — This species was 
noticed upon the wing after nightfall in Pretoria on December 
30th, 1913, and was again observed at 8 p.m. on December 
7th, 1914. The actual meeting of the sexes was not seen. 

Many gathered about the electric street lamps, and beneath 
these some wei^e seen already 

pairedoff with the females lead- Text-kig. 1. 

ing, whilst other individuals, :;*%v >iiJ^>>,, 

male and female, scurried rapid- e ^ -"r^ , ^^ - "' ^^ ^^ . fc =^ =.^--^g ^ 
ly about, as if in search of one ^ - ~- . — — r -7 

another. As soon as a mated 
female began to dig, the male 
took his place by her side and 
seconded her efforts. Both in- 
sects are very expert excavators, 

J 1 1.1 Section of initial burrow and cell 

and when once started soon niade by a pair of H. t vans- 
construct a burrow\ Much of vaalensis. x ^. 
the loosened soil is scattered by 

the feet, the action resembling a hen scratching. 1 'ebbles of no 
meanmagnitndeai-e loosened, carried outin theiaws,and placed 
well to one side of the pit^s mouth. The larger and grosser 
female displays just as much activity in this work as does the 
lither and less obese male. Their strength and their aptitude 
is best illustrated by the fact that they succeeded in burying 
themselves in the stones and grit of a macadamised roadway. 

If interfered with the insects become most excited, and 
pairs placed on soil in captivity lied around like cockroaches, 
taking no notice of one another. When, however, artificial 
burrows were made, and the insects were dropped head first 
into them, they became normal at once, and just as instan- 
taneously began burrowing. 




336 CLAUDE FULLER. 

Actual burrowing in open land has not been observed, but 
under the uniform conditions of a box of soil and observa- 
tion cells, it was found that the insects did not close in their 
burrow behind them ; instead they placed the loose particles 
regularly around the aperture so as to form a small per- 
forated mound, Avhich persisted for several weeks (text-fig. 
1). Some females laid eggs within the first week, others 
only after the lapse of a fortnight. On May 25th, 1915, one 
pair with young were still alive. 

Termes natal en sis Haviland. — The mating of this 
species has only been observed on one occasion (Pretoria, 
October 28th, 1918, 6-7 p.m.), when it took place in conjunc- 
tion with a flight of T. incertus. The latter was, however, 
quite local, whilst the winged natalensis had come from far 
afield — probably a mile away. The insects flew high with 
well-sustained flight, and many passed overhead. The females, 
as appears usual, alighted first; selecting free and high- 
standing grass spears. Upon gaining a foothold (fifteen to 
twenty inches from the ground) they at once reversed their 
position and stood head downAvards. Then all four wings 
Avere half opened and the apex of the abdomen became 
visibly swollen. Some males were flying low over the her- 
bage when this act took place, and in a very short space of 
time the females were discovered by them. The male 
alights, as a rule, directly on the dorsum of the female, flying 
to her with much precision. He rapidly aligns himself, with 
wings closely folded and head upAvard, and combs across and 
across from one cercus to the other the dilated apex of the 
abdomen of the female. During this the female sits quite 
still, but presently she dealates and gives e\adence of restless- 
ness; Avhen her moA^ements sufficiently stimulate the male, he 
dealates and craAvls from her back. The female then moves 
forwards doAvn the stem, the male close behind, Avith mouth- 
organs ahvays closely brushing the anal plates of the female. 

There is no doubt in connection with this mating that, 
whilst the male is attracted to the female bv the sense of 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 337 

smell, as is the case with T. vulgaris, the directness with 
which he alights indicates further that, within a circum- 
scribed radius, the female is visible to him ; the play of the 
rays of the setting sun upon her half-spread wings render- 
ing her quite a conspicuous object. 

On this occasion a number of dealated pairs were collected 
and confined over-night in a small cardboard box. Several 

Text-fig. 2. 




The mating of Termes natalensis. a. Two views of female iu 
calling attitude on grass-stem. B. Two views of association of 
sexes. X i. 



of these were successfully re-mated the following morning, 
three pairs burrowing into the soil contained in large glass jars. 
All burrowed to the bottom, to a depth of between four and 
five inches ; and, unable to proceed further downwards, 
contented themselves by making a more or less globular 
cell about an inch and a quarter in horizontal diameter and 
three-quarters of an inch high. Owing to this fortunate 
circumstance it was possible, after removing the dark shield 
placed around the jar, to obtain some small view of the 
actions of the insects. The first pair soon died; the individual 



338 CLAUDE FULLER. 

which survived for a time buried its mate. On the four- 
teenth day one of the second pair died, and its mate was 
watched whilst it carefully encoffined the corpse in earthern 
paste. The third pair remained alive, displaying little if any 
anxiety Avhen exposed to the light. On the sixteenth day 
(November 13th) a small bunch of yellowish eggs was 
noticed adhering to the glass side of the cell. On December 
29th the first egg hatched, but whether one of those first laid 
(these appeared to become somewhat desiccated) could not 
be determined. Two days later fourteen young termites 
were counted, and many eggs Avere still unhatched. Before 
they hatched, the eggs were the object of frequent attention 
by both parents, and they often inspected them, feeling them 
over with their palpi. The young insects received constant 
attention from their parents, and apart from being fed by 
them they seemed to be constantly groomed. On the ninety- 
eighth day (February 2nd, 1914) some of the workers 
appeared to be adult, the heads showing yellow chitin. By 
this time the cell was a crowded mass of life and some eggs 
were also seen ; at the same time no enlargement of the cell 
had been made and no galleries driven from it. Naturally, 
neither male nor female had taken any kind of nourishment, 
but they did not appear any the worse for their long fast nor 
emaciated by the feeding and raising of their young. This 
unfortunately is the last note made upon the colony. The 
writer left headquarters the next day, and during his absence 
the soil was kept too moist, and the whole colony succumbed 
and rotted before his return. 

The burrowing of the pairs of this species resembles that 
of the smaller kinds ; but, in common with other larger sorts, 
it is capable of greater exertions, and in beginning the 
operations the insects can be seen scratching the loose 
particles with their feet and throwing them behind them, as 
do most burrowing animals. When a pebble is met with, it 
is not avoided but picked up in the jaws and carried out 
and deposited where it cannot roll back into the excava- 
tion. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 



339 



Terines latericius. — The mating of this species was 
observed in Pretoria at dusk on December 4th, 16th, and 
18th, 1914. The females were seen to alight first, taking 
up an inverted position on pendulous parts of grasses and 
herbage. Immediately on assuming this attitude they begin 
to agitate their wings violently, and keep them in motion 
until a male becomes associated with them. How long a 

Text-fig. 3. 




The muting of Termes latericius. AanclB. The female in the 
calling attitude, violently agitating her wings, c. The sudden 
discarding of the winos when the sexes meet. X |. 



female may go on agitating the wings without a male finding 
her cannot be said, but one watched for twenty minutes 
never stopped, and after dark was still unmated. 

The males seem to have great difficulty in finding the 
females ; they fly low and incessantly over the tops of the 
grasses, and again and again approach quite near to a 
female, and even circle within a few inches around, only to 
fly off again far afield. 

When mating was accomplished during twilight it was 
noticed that the male flew in narrowing circles around the 



340 CLAUDE FULLER. 

female and alighted near to her. Directly the male touches 
the female the wings of both drop Avith inexplicable sudden- 
ness, those of the female seeming to fall whilst still being 
agitated. After this the female leads the way to the earthy 
the male following close behind. Here they soon burrow into 
the soil, and there form a cell. Two pairs which burrowed 
into observation-cells on December 16th had both produced 
fifteen to twenty eggs eight days later. They were trans- 
ferred to glass capsules, and on January 2nd had laid a 
further batch of ten to twelve eggs. Alive May 25th, 1915. 

Termes vulgaris Haviland. — The mating of this species 
has only come under observation once (Pretoria, December 4th, 
1914). The departure of the adults from the nest and their 
mating occurred between the hours of 8 and 10 a.m., imme- 
diately after a heavy and continuous downpour of rain which 
had lasted over three hours. The flight took place from a 
nest in the banks of a stream, and, but for this fact, the nest 
could not have been located. 

The insects emerged from numerous apertures in the bank 
and in the surface overlying the site of the nest. Later in 
the day these were closed and covered with little mounds 
of soil. From the fungus-garden cavities to the apertures 
the insects travelled along specially excavated galleries, an 
inch and a half wide and a quarter of an inch high, driven 
upwards through the soil so as to form an inclined plane. 
These galleries are exactly similar to those made for the same 
purpose by T. latericius, and form another connecting link 
between these two similar but quite distinct species. 

Upon leaving the nest the adults radiated out in all direc- 
tions from it, and mating commenced within the first 100 
yards. The full peripher}' of the flight was not actually 
determined, but in one direction it extended for more than 
half a mile. No wind was bloAving at the time, and there 
was no evidence of an exodus from another nest. Along the- 
radius of flight observed it must be conceded that all the 
mating was between insects of like parentage. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 34T 



Text-fig. 4. 



The females came to rest first, alighting here and there- 
upon grass stems — both long and short — and immediately 
took up a position at the apex of the stem, with head down- 
wards and wings closely folded. No particular inflation of 
the abdomen could be noticed, but it was obvious that some' 
sense other than sight alone guided the male insects. 

When a female alighted, a male was soon to be noticed 
fluttering low over the herbage. 
This flight was seen to be most 
erratic, although generally round 
and about the centre represented 
by the female. Often the male 
would fly close by and attempt 
to alight upon an adjacent stem 
— sometimes even doing so — but 
only to fly off, low across the 
grass tops for ten to twenty feet. 
Ultimately the male manages to 
locate the female, and settles be- 
low her upon the same stem. He 
at once crawls up beside her and 
rapidly passes the mouth-parts 
across the region between the 
cerci and touching the cerci them- 
selves. Almost simultaneously The mating of Termes vul- 
with this action both insects dea- 
late and the female begins to 
crawl downwards, the male following. In this attitude the 
insects progress until the female finds a site suitable for 
burrowing; the male never more than just keeps in touch with 
her. When burrowing commences the male takes his place 
beside the female, and in loose soil the pair soon become lost 
to sight. Pairs of this species did not thrive in the observa- 
tion jars, but they constructed cells, and eggs were laid within 
a feAv days. One pair with a few young May 25th, 1915. 




garis. Female 
attitude, x f. 



in calling; 



Termes incertus Hagen. — It would appear that shortly 



542 



CLAUDE FULLER. 



before the time arrives for the exodus of the winged indi- 
viduals of this subterranean species, a number of wide galleries 
or chambers are excavated within two to three inches of the 
surface of the soil beneath some bare spot, and in these the 
winged forms are assembled, ready to dejjart at the chosen 
moment. When this arrives several (one to five) perforations 
are driven upwards through the surface. From each of these 
first of all adventure 2-300 workers, but never any soldiers ; 

Text-fig. 5. 




The flight of Termes incertns. x x. Underground assembly' 
cavities shown in section, x A. 



these workers swarm around the individual openings out of 
which they have come^ spreading out to form a circular mat 
four to six inches across. In this circumscribed area they keep 
constantly upon the move, and none ever wander away from 
the main body. Their egress is immediately followed by 
that of the winged. 

Although the workers and the few soldiers that remain 
Avithin the galleries hasten about among the images, per- 
chance instilling into them " the spirit of the hive," those 
without pay no attention to them, their function seeming only 
to be the formation of a barrier — a living sacrifice — against 



OBSERVATIONS ON SOME SOU'I'H AFRICAN TKRMITES. 343 

the ants Avhich gather viciously ai-ound. By their action 
they secure to the wing-ed a safe departure. Overcome with 
anxiety to seize upon the winged insects, the excited ants take 
no notice of the worker termites, and it is quite exceptional 
to see one of these carried off until after all the winged 
insects have dispei'sed. The barrier is eifective, however, 
because when an ant steps in among the workers to reach its 
most prized prey, it is so discomfited by the nips of the little 
termites upon its feet that it is compelled to retire. 

At Pretoria, both in 1913 and 1914, the images of this 
species emerged in great abundance on various dates through- 
out the month of November,^ and always between the hours 
of 6 and 7 p.m. or during the brief twilight. The act took 
place almost invariably 10 to 24 hours after a fall of rain, 
the evenings selected being uniformly mild and quiet. Yery 
many instances came under observation, but in none of 
these did one occur at any other time of the day, nor when 
windy, nor when rain was falling. On the other hand an 
emergence may often be followed within an hour or so by 
both strong wind and heavy rain. 

The exodus commences almost suddenly and ceases quite 
abruptly. When darkness actually falls the insects are no 
longer on the wing, so that the period involved is never more 
than 30 to 45 minutes. ISJ^umerous cases came under observa- 
tion in my garden, on the neighbouring kopjes and elsewhere. 
During one evening in 1913, and again in 1914, over a mile 
length of street was traversed, and along the whole length 
this species was emerging from countless places, in every 
garden, and at short intervals along the sidewalks. Such 
instances furnish some idea of how extensively this species 
inhabits the soil of Pretoria. 

The winged incertus issue forth from the soil very rapidly 
and jostle and crowd upon one another, in the narrow confines 
of the apertures, to such an extent that very often they are 
so densely packed as to form small upright columns from the 

' The actual period is the last week in October into the first week in 
December. Another emergence occurred in March, 1914. 



344 CLAUDE FULLER. 

apex of wliicli those ahead take flight. As the insects all 
stream away in one direction, carried on some gentle air 
current, the emergence as a whole resembles a cloud of smoke 
issuing from a chimney. 

If one of the holes from which they are emerging is 
covered by a glass tumbler the action at once ceases and the 
workers retire. If a series of holes are so covered the effect 
is the same, but, very rapidly, fresh openings are made near 
by and the exodus proceeds. 

The females are the first to alight, and they do so directly 
on to the soil surface, always selecting a spot — large or small 
— which is bare of vegetation. On alighting they immediately 
dealate ; occasionally they then run forward an inch or two, 
but more usually stand still and, puffing up the apex of the 
abdomen, elevate it until one half of 
EXT-pi . . ^1^^^ region is at right angles to the 

rest of the body. Without doubt some 
perfume is wafted upwards, for males 
flying low overhead are immediately 

T. incertus . A female attracted and drop to the ground within 
m the calling attitude. . '^ ° 

X 1-5. 6 to 18 inches of the female. Usually 

only one alights, but sometimes 
several do so. The male at once dealates and hurries with, 
comparatively speaking, remarkable precision towards the 
female, even should he fall amidst the grass. In one instance 
where the females were alighting the ground was overrun 
Avith ants to such an extent that as each fell it was captured. 
Over this area males were seen flying low (20 to 30 inches) 
and, to all appearance, systematically searching the area for 
the expected scent. This they did until darkness fell. 

The female does not remain in the calling attitude in- 
definitely, and the period is seldom more than a minute at the 
outside. She usually lowers her abdomen after a short while, 
runs a little Avay and then elevates it again. Directly she 
detects the immediate proximity of the male, and is certain 
that he is about to approach her, she hurries forward, and the 
male, catching up to her and placing his head upon the end 





OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 345 

•of her abdomen, hastens along wherever she may lead. 
Occasionally two or even three males will follow^ one and the 
same female, and later the whole party will proceed to burrow 
into the soil quite amicably. Once the male has taken up his 
position, the female's objective is some crack in the soil or 
some small pebble she can easily pass beneath. This reached 
she immediately begins to burrow, and the male, coming for- 
ward beside her, assists in the excavation. If a scratch is 
made in the soil, so as to surround an advancing couple, the 
female on reaching it will begin to dig ; reacting at once to 
the stimulus of this suitable but artificial environment. 

The burrow is begun by lifting out the earth particles and 
placing them around the 

spot. When it is sufficiently ' ^' ' 

deep to accommodate the full 
length of the body — about 
three-quarters of an inch — 
the entrance is closed by a 
gi-adual process of building 
in the earth particles until 
a dome is made. The gallery T. incertus. A pair begimiing 
is then extended slowly down- *"eir burrow and the same in- 

sects alter about au hour s bur- 
wards, each particle as re- rowing. About natural size. 

moved from the bottom being- 
carried up and inserted and carefully compressed into the 
vault. In doing this the insects work to a very great extent 
in alternate positions, one excavating at the base whilst the 
other is placing its burden into the roof above. Hence it 
comes about that as the burrow is extended downwards it is 
filled in behind. The insects do not appear to moisten or 
cement the earth particles in place, the moisture in the soil 
itself being no doubt sufficient for the purpose. 

Although operations proceed without intermission, and 
both sexes work rapidly, progress is very slow. This is not 
so much due to the difficulty of excavating as to the time 
taken in inserting the particles into the dome. A number of 
couples, whilst succeeding in burying themselves in ten to 




346 CLAUDE FULLER. 

fifteen minutes, liad only reached a depth between 1 and 
1^ in. in two houi's, and 4 in. in eighteen hours. ^ 

On October 28th, 1918, a pair of T. in cert us burrowed 
into the soil of a glass jar. They did not go to the bottom 
but to a depth of 5 in. The jar was left undisturbed until 
December 30th, when the soil was carefully removed and 
portion of the cavity they were occupying exposed. Both 
insects were then quite healthy and were surrounded by a 
brood of callows which they had obviously reared themselves. 
The cavity was closed and the jar refilled. They were not 
again disturbed until the middle of Februaiy, when, during 
the absence of the writer for a fortnight, too much moisture 
was added to the jar and the colony succumbed. 

In November, 1914, attempts were again made to keep this 
species under observation in glass cells. At first, the insects 
took kindly to their artificial environment, and several pairs 
placed in the cells on November 5th had laid five to seven 
eggs on December 2nd and then succumbed ; the dead bodies 
being covered with mould-growth which appeai-ed to be rather 
of a parasitic nature than to be due to excess of moisture or 
other adverse conditions. 

A few pairs remained alive in artificial cells, i. e. glass 
tubes plugged with earth ; their eggs had not hatched on 
Januaiy 11th, 1915. All died early in February. 

Eutermes bilobatus Haviland and an undetermined 
species. — The undetermined species is an insect which is 
related to Eutermes bilobatus but is not known to build a 
superticial mound. Structurally the soldiers of the two species 
are quite distinct ; but for convenience the mating of the 
images of both species may be dealt with together. The 
Haviland notes (1) contain a good description of the mating of 
bilobatus, although it is curious that neither in the description 
of this species nor in that of its mating does Haviland remark 
upon the great disparity in size, as between male and female. 

' The observations were made with couples placed in naiTow vertical 
glass jars of moist soil. 



OBSERVATIONS ON SOMH SOUTH AFRICAN TEli JUTES. 347 

According to the notes, the adults of bilobatns '"fly 
around bushes and ultimately settle upon them. The male 
seizes the ventral surface of the abdomen of the female with 
his wings projecting in front of her head. The female flies 
away with him in the direction of the wind. On settling the 
male loosens his hold and both shed their wings. The female 
then proceeds to look for a place of shelter, the male following 
her." 

No full opportunity to confirm these observations has yet 
presented itself. On several occasions the species was seen 
emerging before seven on sunny surainer mornings ; in one 

Text-fig. 8. 




^S^ 




& i c d 



The mating of species allied to Eutermes bilobatns. a. Male 
attaching itself to female, h. Female dragging off dealated male, 
c. Female flying with male. d. Male and female seeking a spot to 
burrow. (About natural size.) 



case a female was seen to alight on the ground beside a 
dealated male, when the small creature immediately seized her 
abdomen Avith all his feet, and then the female rose into the 
air and flew oif with him. 

The mating of the allied species was more fully noticed in 
Pretoria, November 14th, 1914, between 1 and 2 p.m. Before 
this, the species had been seen several times on the wing 
during light showers of rain and its habit of swarming around 
trees noted. During a shoAver of rain the writer happened 
to be standing under an open umbrella in a field where 
thousands of insects w^ere emerging from the soil. The 
umbrella attracted them, and soon a cloud flew around, 
hundreds alighting upon the wet, upper surface, but none 
attempting to come beneath its shelter. In the wet film 
pairing proceeded rapidly. The females walked about con- 

VOL. 3, PART 2. 24 



348 CLAUDE FULLER. 

tinually, rapidly uplifting and lowering tlie whole of the 
abdomen, and raising the wings in unison. Presently a male 
would run in under the uplifted abdomen and, raising his 
head to its apex, seize it with all six feet ; then, as the female 
pressed dow nward, he would dealate and tumble beneath her. 
This association achieved, the female walked with half -spread 
wings to the edge of the wet field, dragging the tiglitly 
clinging male with her. From here she took to flight, and, 
obviously burdened, soon settled down to earth. Immediately 
on touching the earth, the male-encumbered females dealated .; 
during the action the male released his hold on the ventral 
sui'face and climbed upon the back of his mate, placing the 
head upon the fifth or sixth abdominal segment and clasping 
with both fore and mid-legs ; only the hindmost pair was 
disengaged for walking. It appeared that upon some occa- 
sions the male did not dealate on clasping the female, as a 
winged pair was noticed on the wing ; the male attached as 
Haviland describes in the case ofbilobatus. This pair fell 
into a rain-pool, and on touching the water the female dealated. 
When lifted out the male relinquished his embrace and flew 
away. 

The adults were noticed to emerge from small round aper- 
tures leading from cavities just below the soil. This they 
did in a manner very similar to that of incertus, except 
that never more than forty to fifty workers formed the sur- 
rounding mat. In the assembly-cavities many w^orkers and ■ 
a few soldiers were seen hastening in and out amongst the 
crowded winged forms. 

It may be said, with no small degree of certainty, that all 
imago termites are at first positively hydrotaxic, and a 
captured pair of this species displayed the symptom to a 
remarkable degree. They were placed upon a tray of diy 
soil and wandered disconsolately over its surface. When a 
few drops of water were applied to one spot they soon found 
it ; they wandered about for ten minutes, but returned to the 
small moist area and there buried themselves. They remained 
in the moistened area and burrowed about in it for twenty-four 



OBSERVATIONS ON SOME SOUTH APEIOAN TERMITES. 349 

liours — apparently refusing to enter the dry surrounding 
soil. 



IV. OBSERVATIONS ON THE NESTING HABITS AND 
GENERAL ECONOMY OF CERTAIN SPECIES. 

Hodoterraes transvaalensis sjj. n. PI. XXV, figs. 2-7, 
PI. XXVI, figs. 1-5. 

During the pi'ogress of the excavations, of which mention 
has been made, much care was taken to acquire some know- 
ledge of the subterranean work of this species. The oppor- 
tunity was excellent, as all of the ground involved was 
inhabited by it. Upon the whole, however, the results were 
meagre. 

A great many hives were exposed, but only two of these 
were inhabited. The majority were in various stages of 
dilapidation and others appeared to have had their general 
contours destroyed by flooding'. The evidence collected went 
to show that ordinarily the species voluntai'ily deserts its 
hives and later fills them up with excavated particles of earth ; 
using them, in fact, as dumping grounds. It is quite 
possible that desertion is sometimes due to the collapse of 
the f]"agile interior structure. 

The inhabited hives were both similar, and were in reality 
central granaries in which the hay harvested is finally 
deposited. They are doubtless also nurseries, but neither a 
queen, nor eggs, nor particularly small callows were found in 
either. The surrounding earth was explored in the hope of 
tracing some special compartment in Avhich eggs were 
possibly incubated, but nothing of the sort was found, 
although many cubic yards of earth were removed. 

There was ample evidence that during the excavation 
before the hives were reached the insects left them ; one, in 
fact, was wholly deserted. In the other, which was broken 
into somewhat suddenly, numerous and quite white callows 
wei-e found ; none, however, were particularly small, the 



350 CLAUDE FULLER. 

smallest being 5 mm. long. It is suspected that the hives 
of this species intercommunicate, as do those of Eutevmes 
trinervius, and that the economy of the two is moi'c or 
less analogovis. 

The hive-cavities (Plate XXV, fig. 2, and Plate XXVI, 
fig. 1), with one exception, were all sub-spherical, having a 
horizontal diameter of 24 in. and a perpendicular height of 
18 in. The cavities are partitioned by very numerous hori- 
zontal and close-set shelves (Plate XXV, figs. 4-6). These 
are constructed of a thin and very papery snbstance which 
does not dissolve in water or in alcohol. The shelves lie one 
above another with striking regularity, and are attached to a 
series of clay brackets projecting from the walls. The shelves 
are not equi- distant apai't throughout the cavity, but range 
from 6 to 15 mm. Innumerable little cylindrical columns of 
wooden texture, spread over the field of each shelf, hold the 
whole fabric together (PI. XXV, fig. 4). These little columns 
are not stairways ; the insects pass up and down from storey 
to storey of the hive by short inclines. The hive is not like 
that of any other known termite structure in South Africa. 
Except that the shelving is as br-ittle as charred paper, 
whether moist or dry, the shelves might be described as being- 
more like dark brown paper than anything else, as they are 
scarcely thicker than stout paper. 

A most striking feature is found in a series of eight great 
shafts which, descending more or less perpendicularly from 
the sides and bottom of the cavity to a distance of 2 to 3 ft. 
into the soil, run forward in a horizontal coil and then end 
blindly. Whether these are for the purpose of permitting 
heavy gases developing in the hive to find a lower level, or 
are merely for drainage in the case of soil saturation, is purely 
a matter for speculation. 

The hive is approached and entered by equally large 
galleries which perforate the sides of the cavity. They are 
comparatively few in number, only four being found in the 
nest examined fully. The galleries do not run ofl: in a 
horizontal plane, but first descend to a level below that of the 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 



851 



nest and then rise again (text-fig. 9). Here they become 
greatly reduced in diameter (PI. XXV, fig. 7) and then travel 
in a horizontal plane, but somewhat erratically, away from 
the nest. Owing to the labour involved and the indirect 
route taken, it was found impossible to trace a gallery the 
whole way either from the nest to its exit at the surface or 
from a surface burrow to the 

Text-fig. 9. 



iiliiiiiiAi4[rh 



nest. The greatest length 
of straight gallery exposed 
was about 30 ft. This rose 
from a depth of 4 ft. to 
witliin 2 ft. of the surface, 
runnino- in horizontal lenyths 
of 6 to 8 ft. rising by short 
inclines from one level to the 
next. Because of the few- 
ness of pouches in this par- 
ticular gallery, and its re- 
lationship to one of the hives, 
it was regarded as a middle 
length in a main passage- 
way. 

Upon reaching to within 
a foot or so of the surface 
the galleries branch and in- 
terlace, some naturally con- 
necting up with others; fi-om 
here galleries are driven to 
the surface, and as many as 

twenty-six openings were counted on one feeding ground 
10 ft. by 5 ft. Near to the surface of the soil, cavities are 
made for the temporary storage of the material harvested. 
These ai'e usually to one side of the gallery and in the form 
of Avide, flat pockets. Deeper in and also Avhere the tunnels 
approach the hive there are much lai'ger cavities (6 to 8 in. 
in diameter). The floors of these gallery extensions are all 
broadly convex ; they are quite shallow, however, and the 




H o d u t e r m e s 1 1" a ii s a' a a 1 e ii s i s . 
Section of subteiTaneaii grauaiy 
shcjwing entrance galleries and 
descending blind shafts. X o^r;. 



352 CLAUDE FULLER. 

majority of them fall away on one side of the gallery to a 
lower level (PL XXVI, figs. 2 and 2a). 

It was noticed in all the inhabited and uninhabited hives 
that no passage-way entered the dome of the hive, and where 
a tunnel was found about a foot below ground running over 
the site of one nest it showed no indication of approaching 
the cavity but kept straight and horizontal along all the 
distance (10 ft.) it was exposed. 

Harvesting is conducted in the bright sunshine, and Mr. F. 
Thomson, of the Division of Entomology, informs me that on 
several occasions when bivouacking on the veld he has 
observed Ho do term es s2J. working at night, but only when 
the moon was bright and full. He also states that they quite 
commonly enter stables and feed on the hay litter therein at 
night. 

Ordinarily, however, they work during the daylight, closing 
the entrances to the bui-rows at night-fall with short stoppers 
of clay. Harvesting is conducted all the year round with, 
as far as observations go, a break when Avinter merges into 
spring. From complaints reaching the writer Hodotermes 
spp. seem to be most active, or are noticed to be most 
active, during the autumn and early winter months. In 
Pretoria, however, the activities of transvaalensis Avere 
noted all through the summer and until the end of June. 
Some time subsequently these ceased and remained in 
abeyance from August to the middle of October. Their 
activity in the latter part of the season may be put 
down to the then more favourable condition of the grass for 
storage, and the cessation later on to the fact that their 
granaries are full, rather than to the succulence of the young 
grass. There are several fictions regarding the activities of 
these termites. One is that they are more abundant during 
periods of drought, the other that they are most active just 
before a thunderstorm. Speaking on personal observation 
alone, they have never been seen more busy than on one 
winter's morning between 8 and 9 a.m. when frost lay thick 
near-by under the shade of a Cupressus hedge. They 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 353 

appear more abundant during drought, first because they are 
then more apt to pillage cultivated crops, and secondly 
because the veld is barer and so they are more readily seen. 
The thunderstorm fiction seems wholly due to the fact that, 
in the stillness preceding the breaking of a storm, the noise 
they make in biting through dry stems of grass is distinctly 
heard and draws attention to them. The writer cannot say 
positively that they are not more active at such a time, as it 
is quite possible that they are more excited, responding, as do 
many insects, to storm influences ; they seem to be making 
hurried efforts to get in the last piece of hay before the stomn 
breaks. 

The foraging is done by the workers, both large and small, 
callows also assisting in it. The lengths are carried upright, 
as described and figured for H. havilandi by Sharp, or are 
merely di-agged along to the mouth of the burrow. Here the 
bulk is deposited, but some is always being carried in. When 
a sufficiently large quantity is collected, or towards nightfall, 
harvesting ceases, and such supplies as have not been broad- 
casted by the wind are removed below the ground. The 
soldiers do not accompany the workers, but as a general rule 
one or two are to be found on guard just within the mouth of 
the gallery. 

Sometimes, when a veiy large collection of hay has been 
made, a loose mound of earth has been thrown over it. Such 
an accumulation may remain so weighted down for several 
days (PI. XXVI, fig. 3). 

As no mound of any sort overlies the site of the hive, and as 
no galleries are driven to the surface in its proximity, it is 
interesting to note the disposal of the considerable quantity of 
earth excavated in the making of cavities and galleries. As 
far as possible this soil is utilised for filling up disused galleries 
and deserted hives ; cavities made by other termites are also 
filled. Curiously enough these fillings remain dry where they 
are placed, the soil particles not being easily wetted. 

The following are a few specific instances coming under 
observation : (1) a deserted hive completely filled with earth 



354 CLAUDE FULLER. 

particles, the interioi* partitions being mostly intact; (2) a 
deserted hive, in which the interior had collapsed and fallen 

the bottom, was being filled through openings in the dome 
of the cavity, the earth particles and fragments of dead 
Hodotermes being dropped in; (3) the filling in of the 
small fungus-gardens of T. incertus from holes in the dome 
of the cavity. To these may be added many cases where 
cavities of all soi-ts and sizes were found completely filled up. 

Besides acting in this manner, Hodotermes transvaal- 
ensis brings a great deal of earth to the surface, arranging 
the loose particles in mounds which, when of any size, strongly 
resemble small mole-hills (PI. XXVI, fig. 4). These often 
occur over a fairly wide area, but they do not coincide with the 
nest-site nor are they necessarily adjacent to the openings from 
which the termites emerge when bent on gathering provender. 

A hole is driven to the surface which is about a half or one- 
third of an inch in diameter. The first particles of moist earth 
are built together as a collar around this, and further particles 
are pushed up so as to fall outside it. As the mound rises in 
height so the collar grows into a chimney, the encircling pile 
of earth supporting it upon its weak foundations (PL XXVI, 
figs. 5 and 5a) . Often these progress no further, the holes at 
the top are closed up, and the wind wearing on the hill leaves 
the apex of the chimney exposed. 

Where large quantities of earth have been thrown out, the 
inner chimney is found enlarged and branched and of 
various grotesque forms. It is obvious from the arrangement 
of the branches that they are constructed so that the soil will 
fall away from the point on one side of the moundlet where it is 
discharged (PI. XXVI, figs. 4 and 4a, 4^, 4c). 

Curiously enough the insects do not expose themselves in 
doing this work. When the pellets are brought to the mouth of 
the chimney they are moist and cling together so as to cloak 
it ; and all that one sees is grain after grain taking its place 
at the apex of the mound and then, on drying, rolling down 
the incline. If the mound of loose earth is gently removed, 
the chimney — however bizarre — stands upon its base ; but if 



OBSEHVATIONS ON SOME SOUTH AFRICAN TEKMITES. 855 

touched falls over and ci-umbles readily to pieces. Rain and 
wind both rapidly distribute these heaps of soil. 

A hive of H. havilandi was discovered under rather 
peculiar circumstances at Weenen in Natal. Unfortunately 
when inspected nothing but the cavity and some debris, from 
which dead termites Avere removed, was found. This occvirred 
in the wall of an office in the village, and all the information 
obtainable was that one day a slab of plaster fell away, and a 
mass containing many termites and pieces of grass fell out on 
to the floor. The cavity was about 2 ft. 6 in. from the ground, 
and 15 in. in both diameters. The wall was built of burned 
brick ; but since the bricks lining the cavity were only 
grooved, so as to form brackets for the shelving, one cannot 
help suspecting that the spot in the wall chosen by the 
termites was a part which had been filled with some sun-dried 
bricks. Hodotermes havilandi is a very familiar insect 
in and about the village of Weenen ; because of this, it is 
interesting to add that the Chairman of the Village Board — in 
whose office the event here related happened — had never seen 
the insects feeding nearer to the hive than 100 yds. off. He 
had, however, been long familiar with the fact that they 
were tunnelling in the plaster of the walls of his office. 

H. havilandi appears to have a very definite life-zone in 
Natal, being restricted to the warm and dry river valleys of 
the midlands. 

The method of harvesting of all our Hodotermes is 
uniform with that of Eutermes trinervius. The accumu- 
lation of provender outside the bun-ow entrances has a double 
purpose — it prevents congestion of the tunnels and allows 
any green parts to dry. Doubtless also its storage in pockets 
before being carried into the main hive is to insure sufficient 
dryness so as to avoid the growth of mildew. 

Under date of January 12th, 1915, my colleague, J. C. 
Faure, entomologist at Bloemfontein, sent to me specimens of 
H. transvaalensis with the following notes. 

" Bloemfonteiu, Deceml)er 7th. 1914, 6 p.m. Heavy rains had fallen 
a few days previously. Near the Sewage Farm orchard I was attracted 



356 CLAUDE FULLER. 

to flying Hodotermes by a large number of dragon-flies that were 
hovering around one particular spot. 

"Winged forms. — They were emerging in considerable numbers 
from ordinary foraging holes in the soil. There was a considerable 
number of small mole-hill-like workings near by. The males and 
females ran out of the holes, and took flight almost at once by jumping 
a little distance into the air and then bringing the wings into action. 
The distance flown could not be ascertained because the dragon-flies 
captured a very large percentage of all the termites that took to wing. 
Those that did escape did not fly more than, say, 30 ft. from the flight- 
holes. They got rid of their wings in the ordinary way. The meeting 
of the sexes was not observed. I think it likely, however, that the male 
and female meet on the ground after losing their wings. Only a few 
pairs could be found in the act of beginning to burrow. They were, 
however, entirely alone, no soldiers or workers being present. 

"Workers. — Near the holes from which winged forms were 
emerging there were many workers engaged in their ordinary work of 
foraging. I did not actually see them carrying food into the flight-holes, 
l)ut there were several other openings to galleries near by. There were 
no workers at the flight holes while the males and females were emerging. 

" Soldiers. — A number of soldiers were found near the openings of 
the galleries from Avhich earth-mounds had been thrown up. These 
were about 10 ft. from the flight-holes. I was unable to get soldiers 
from one or two flight-holes that I dug up. At any rate, it was easy 
to get soldiers under the little earth-mounds near by, much more so than 
when no flight is in progress. There were, however, no soldiers on the 
surface of the soil. 

"It was remarkable how large a percentage of the flying termites 
were captured by dragon-flies. They were mercilessly captured, their 
abdomen was devoxu-ed in a very short time, and then the head, thorax 
and wings were dropped. Frequently these " de-abdomened " termites 
were seen frantically digging in the soil, obeying their instinct to fly, 
drop and dig. In these abnormal cases, however, they forgot to search 
for a mate first, and they did not drop their wings, probably because 
they had no alidomen. Needless to say, they soon died. 

" Large black ants prowled around near the flight-holes and carried 
off injui-ed termites or those that were unable to take flight soon 
enough." 

Hodotermes viator [Latreille). PI. XXYI, fig-s. 6, 6a^ 

Qh, and 7. 
A numbei' of images without wings which appear to be of 
this species were collected for me at Victoria West during 



OBSERVATIONS ON SOMK SOUTH AFRICAN TERMITES, 357 

December, 1914, by Mr. F. Jansen, the Resident Magistrate. 
These Avere found issuing from holes in the ground i7i the 
streets and in the country surrounding the township. 

Specimens of soldiers and workers have been collected in 
the Stellenbosch district by Mr. F. W. Petty, Lecturer in 
Entomology at Elsenburg Agricultural College, and by Mr. 
C. P. van der Merwe, of the Division of Entomology. The 
following observations have been communicated by these 
colleagues. 

(a) C. P. van der Merwe. " Insects collected May lotb, 1914, in pine 
forest at StellenLoscli. These termites were observed to collect pine- 
needles into heaiDS varying from 1 to 3 ft. in diameter and from 6 to 
18 in. in height, a few being quite 2h ft. high. The small heaps con- 
sisted only of whole pine-needles, and a fungus was growing through 
the heaps. The large heaps consisted of finely cut-vip needles and 
numerous earthy pellets like fine seeds. 

" The small heaps have a tube of earth running down the middle 
into the soil. In the large heaps there are more than one of these 
tubes, but they appear to run from the surface directly into the ground. 
Pieces of cut-up needles were found in the burrows.'" 

(b) F. W. Petty. "The specimens sent were collected from two types 
of mounds, of which I send you sketches (PI. XXYI, figs. 6, 6«, 6h,. 
and 7) in the barley and oat fields. These mounds are about 2A in.- 
high or less. They are made of small, loose. Ijlack pellets brought up 
from below ground, and the mound has a firm, hard core running up 
through the middle of it. This core never runs up straight, and often 
branches into two tubes, one being always shorter than the other. These 
tubes or cores are not made up of the black pellets, but consist of ordinary 
soil, and have the inner lining smooth and hard. Evidently the pellets 
are brought out of the opening, and, as they pile up, the tube is 
lengthened. The insects also make small mounds of gathered bits of 
grain, leaves, weeds, etc., the particles of vegetation being from ^ in. 
to 3 in. in length. These collections of vegetable matter are simply 
grouped about a hole in the form of a low mound no more than 1 in. 
in height. I also found liurrows which have been closed over by a 
hard roof of earth, forming an irregular low chamlier into which the 
bm-row opened. Surrounding the outside of the roof of this chamber 
were pieces of vegetable matter (PI. XXVI, fig. 7). 

"A striking fact which. I noticed in the grain fields where these 
termites were doing much damage was that many colonies, or rather 
the termites of many exit holes, were dead, and usually the head had 



358 CLAUDE FULLER. 

Ijeen severed from the ])ody. I thought this was due to the ' witgat 
spreuw,' because bird excrement was found near and amongst the dead 
termites. On one occasion a flock of these birds was seen in the field, 
and on reaching the spot where they were evidently feeding numerous 
dead and mangled tei"mites were found and fresh bird excrement as 
well. I was, however, told by Dr. Peringuey that this destruction of 
termites was very likely due to a small beetle (Monoplius inf latus), 
as he had previously observed a similar state of affairs.^ Upon in- 
vestigation I found many beetles and larvae in the affected colonies 
among the termite excreta. There are two kinds of beetles (identified 
as Monoj)lius a^niulator and M . se gill at us), and they are possibly 
attracted by the termite excreta. I have not yet succeeded in obtaining 
any proof that they destroy the termites. I have confined termites with, 
Monoplius, but the one does not disturb the other ; the termites die 
off with no evidence of death from bites. 

" In those colonies where the termites are all healthy and busily 
gathering cut leaves, etc., none of the fine, black pellets mentioned 
above are to be found." 

Ill a recent coniniuiiication under date of January lltli, 
1915, Mr. Petty adds the following. 

" As regards Monoplius beetles. I may say that further observations 
led to finding them in small colonies among termite excreta where no 
dead termites were present. I have found dead termites among the 
excreta where no beetles were present. Without a doubt H. viator 
has the habit of removing dead termites from underground through 
burrows at the surface, since I have found dead ones brought out by 
living ones. The ' witgat spreuw ' kills and eats this termite. 

" You may be interested to know that along roadsides with sloping- 
banks in this district, H . viator makes simple oi3enings on the face of 
the bank, without projecting tubes, and discharges the fine, black particles 
(excreta) together with peculiar flat, black pellets. These roll down the 
l)ank and accumulate ; in some places as much as a bushel may be 
seen. Evidently no projecting tube is made in these cases because it is 
not required." 

Caloterines durbanensis Hav. PI. XXV, fig. 1. 

There is evidence to show that this species occurs very 
generally along the coast of Natal. According to the Havi- 

' L. Peringuey, "Note on le Gen. Monoplius," 'Ann. de la Soc. Ent. 
France,' Ixx, 1901. 



OBSERVATrONS ON SOME SOUTH AFRICAN TERMITES. 859 

land notes, bni-rows are made in the living wood, the invaded 
parts subsequently decaying. It is further recorded as attack- 
ing orange trees. The writer cannot help thinking that 
Haviland was under some misapprehension regarding the 
habit of this species, because those colonies which have come 
under his notice have always been at work in dead 
branches, either still attached to the tree or prone upon the 
ground. The opportunity has not been found to make any 
particular study of its habits. It may be interesting to add, 
however, that a piece of infested wood collected in March and 
left to dry out in a breeding cage, in Pretoria, contained live 
insects, soldiers and nymphs, when opened during the latter 
part of October. And further, nymphs and soldiers placed 
during March in a glass tube with fragments of the wood 
from which they Avere extracted were alive when last examined 
(pJanuary 11th, 1915). During the whole of this period there 
was no evidence to show that the insects had fed to any 
extent. 

Tbe workings of this species in the wood take the form of 
elongate, flattened chambers communicating with one another 
by tubes of a nari'ow diameter. 

The following notes have been received from Dr. Warren 
ou the observations of Dr. Conrad Akermaii of the Natal 
Museum : 

" On March 1st, 191.3, a piece of dead branch (about 10 in. long and 
3 in. in diameter) containing Calotermes was broken off a living tree, 
at a height of about 5 ft., in the bush near Winkle Spruit on the Natal 
coast. The branch was brought to the Natal Museum and sawn 
through longitudinally. The wood had been considei'ably channelled, 
and the insects were numerous : nymphs with wing-pads, nymphs with- 
out obvious wing-pads acting as workers, soldiers and many small 
young could be seen, but no eggs were observed. The two halves of 
tlie branch were tied together with string and placed in an inclined 
position in a glass jar without a lid. About an inch of water was 
poured into the bottom of the jar, and the end of the branch was just 
in contact with the water. The termites flourished, and large quantities 
of faecal pellets continually dropped into the jar. In March and April 
forty, or more, winged imagos emerged. From April 23rd to August 
23rd there was a pause in the emergence of winged forms. 



"860 CLAUDE FULLER. 

" The two halves of the branch were occasionally separated for the 
purpose of inspecting the colony, and they were firmly glued together, 
apparently by means of the fsecal pellets. The so-called workers were 
by far the most numerous of the different castes. No eggs were seen at 
any time. In 1913 emergences occurred on March 18th and 24th, April 
3rd and 23rd, August 23rd, November 11th, 17th, and 21st (numerous), 
December oth (numerous). In 1914. January oth (numerous), February 
(several). In September, 1914, the colony was seen to consist of a few 
.soldiers and of approximately 250 large nymphs with well- developed 
wing-pads. There were no individuals in the working stage, and no 
young of any kind. From these observations it would appear that the 
small young seen when the trunk was collected in March, 1913, had 
grown into nearly full-gi'own imagos by September, 1914. a period of 
eighteen months. The piece of wood had been reduced to little more 
than a shell, and the imagos which emerged amounted to about 300 in 
number. Thus the number of individuals in the original piece of 
wood, assuming that no reproductive female was present throughout 
the time, was 550, which, together with, say, 50 soldiers, gives a total of 
approximately 600. The piece of wood was subsequently sawn into 
small pieces, but no trace of young could be seen, and, since no eggs 
had been noticed at any time, it is doubtful if a reproductive female 
was present when the piece of trunk was removed from the tree." 



Rhinotermes pntorius SJost. 

This species has only been observed in the Durban Botanic 
Gardens where the Haviland material came from. 

No opportunity has yet pi-esented itself for studying its 
nesting habits. On the assumption that its nest was in tlie 
dead timber of the trees up the trunks of which its galleries 
were noticed, the galleries were followed up to the topmost 
branches, and here it was found that the creatures entered 
and burrowed in the dead wood where Calotermes 
durbanensis Hav. was established. The soil around the 
base of the tree was examined by Mr. F. Thomsen, but he was, 
for want of time most probably, unable to locate a nest. This 
species builds long, narrow covered ways up the tree trunks. 
The coverings of these are well cemented together and 
obviously intended to be of a permanent nature. They differ 
from covered ways of other species in being of a carton-like 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITKS. 361 

material mixed with sand, such as E. parvus has, in excep- 
tional cases, been found building. As a rule only one gallery 
is built up the main stem, and a nearly perpendicular pathway 
is made from the level of the soil. If a branch is made in the 
gallery upon the trunk of the tree, it is only in the form of a 
loop. From these galleries major and minor workers, a few 
minor soldiers, and a rare major soldier may be recovered. 

Termes waterbergi sy. n. PI. XXVII, figs. 1-3. 

My attention was fii-st drawn to this insect by Mr. C. A. 
Simmonds, of the Division of Horticulture, who sent to me 
material from a mound found below some thorn trees at 
Warrabaths, in the Waterberg district, during October, 1911. 

Mr. David Gunn, of the Division of Entomology, subse- 
quently obtained for me a large series of major soldiers from 
a single gallery which he found in this mound. Time did 
not, however, permit him to make any particular examination 
of the nest. 

More recently (March, 1915) I have been fortunate in 
finding this species at De Wilt on the northern side of the 
Magalies range. The country here is what is generally called 
middle veld. It is hot and somewhat dry and covered by 
various acacias and a large variety of other bush-veld trees. 
Throughout, at very short intervals, which are studded mostly 
with acacias, are more or less circular clumps of dense set 
trees. The trees in these clumps are of a number of kinds, 
and all are more umbrageous than those between the clumps. 
In a fair number of clumps stands a solitary and large 
Euphorbia, that characteristic feature of the park-formations 
of Zululand. Without exception the soil-level beneath the 
tree groups is higher by 2 to 4 ft. than that around them, and 
has obviously been raised around the stems of all the older 
plants. The better part of three days was spent in examining 
this particular aspect of park-formation, and, except in the 
case of some tree groups inhabited by T. waterbergi, there 
was ample evidence to show that all were the homes of 



362 CLAUDE FULLER, 

Termes badiiis. A week previously the country had been 
subjected to a torrential downpour of rain, and because of this 
all evidence of old Termite work had been washed away, and 
Avhat Avas present was quite recent. Beneath one group of 
trees a colony of badius had in the intervening period 
covered its nest-site, to a depth of 4 to 6 in., with a fresh 
layer of earth over an area of 100 square feet. No evidence 
whatever of the presence of Termes natalensis was found, 
but T. latericius abounded. The nests of this species were 
only occasionally found in the tree-covered nest-sites, but in 
the intermediate spaces they Avere quite numerous, and almost 
invariably indicated by chimneys only, the absence of mounds 
being quite conspicuous. 

The very general abundance of T. badius and latericius 
over the whole area examined made the restriction of the 
series of nests of T. water bergi to a small area surrounding 
a natural spring quite remarkable. On account of the water 
the tree clumps in which they were found Avere larger and 
denser than those further afield. This series of nests beino- 
found somcAvhat late in the inspection, time and labour did 
not permit me to make a full investigation of the nesting 
habits of the species, and only one mound of medium size, 
which happened to lie conveniently to one side upon a smaller 
nest-site some 20 ft. in diameter, AA^as explored. Here a nest- 
cavity AA^as disclosed, Avhich, from the absence of the queen- 
cell and eggs, I can only conclude Avas but a part of the 
Avhole nest. An adjacent nest-site Avas over 200 ft. in length 
and about 60 ft. across. It Avas overgroAvn AA'ith many large 
trees and a thicket of undergroAvth into AAdiich one could 
scarcely penetrate. Throughout it, hoAA^ever, Avere many 
Aveathered mounds of earth Avhich had been throAvn up by this 
species. Nearly all Avere 2 to 4 ft. higher than the surrounding 
soil (itself elevated above the natural le\'el), and the largest 
mound Avas 15 to 20 ft. across. 

The mounds of T. Avaterbergi are not masonry mounds, 
although they have that appearance Avhen beaten doAvn by 
rain. They are simply deposits of loose earth particles Avhich 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 863 

ave not cemented together in any way. All were entirely 
composed of subsoil, and presumably represent the soil removed 
by the insects in making large hive cavities and other low- 
level excavations. But very few galleries traverse the mounds, 
and such as do are purely temporary, being made only for the 
purpose of throwing out the excavated earth. This is discarded 
in a very similar manner to that adopted by Hodotermes, 
but there are no hardened tubes made through the centre of 
the earth piles. The pellets are brought up and pushed out 
loosely without the insects exposing themselves. In this way 
many perfectly conical heaps of soil are formed, and those 
seen were from 6 to 10 in. in height and 12 to 18 in. in dia- 
meter. They were found anywhere upon the surface of the 
larger weathered mounds and also round about them, some 
being even a little outside the fringe of the nest-site. 

The mound examined was oval in outline, 5 to 6 ft. long, 
4 ft. across, and 24 to 30 in. in height (PI. XXVII, fig. 1). It 
was so recently made that no roots had as yet penetrated it 
(fig. 2). Immediately under the centre of this mound was 
found a large sub-globular cavity with a flattened floor and a 
somewhat conical dome (fig. o). A layer of undisturbed soil 
of some 18 to 20 in. in thickness overlaid the apex of the 
dome. The height from the floor of the cavity to the roof 
of the. dome was between 24 and 30 in. and the diameter 
at the floor 30 to 36 in. The walls of the cavity were 
smoothed, but not plastered over Avith a cemented clay 
lining, as is the case with most species, and in the somewhat 
similar cavities made by T. badius, T. vulgaris, T. late- 
ricius, and T. incertus. Access to this cavity was gained 
by the termites through a series of more or less triangular 
and fairly large openings to devious galleries leading away 
from it. 

The cavity was completely filled by a large sponge-like 
formation of claj^-laminte or shelving, remarkable for the 
fact that at no point was it attached to the Avails of the 
cavity as in the case of T. natalensis and T. latericius, 
and also for its simulation of certain fungus-gardens, such as 

VOL. 3, PART 2. 25 




364 CLAUD K FULLER. 

the supplementary gardens of T.badius, latericius, and 
more particularly vulgaris. When sectioned it Avas found 
that all the shelves were close together — seldom more than an 
inch apart — those at the bottom were level, but from one- 
third of the height the shelving sloped upwards from the 
sides to the centre of the construction, the inclines becoming 
more acute towards its apex. 

On one of the lower shelves fragments of dried grass were 
found. Dispersed about other parts were flattened separate 
fungus-gardens, none of which were more than four inches 
across. These fungus-gardens resemble those of T. natal- 
en sis; their upper surfaces are flattened and the lower 
coronetted like the surfaces of the gardens of natalensis, 
but the perforations are arranged more like the cells of a 
honey-comb. The lower surfaces of nearly every garden had 
been recently gnawed away by the termites, but the few places 
that had not been mutilated showed that this surface some- 
what approached the coronetted surface of the garden of 
natalensis. 

The hive was inhabited by major and minor soldiers, 
workers, and by young in many stages. The minor soldiers 
were not as numerous as majors, and were not at all aggres- 
sive ; on the other hand, the major soldiers poured out and 
were aggressive, drawing blood with each bite they managed 
to inflict. The workers retired; but, if one ceased disturb- 
ing the nest for a minute, they came forward with pellets of 
soil in their jaws prepared to repair the breaches. 



Termes natalensis, Havilaud. PI. XXVI, figs. 8-19 ; 
PI. XXVIII, figs. 1-12. 

Mounds and Nest-Sites. 

The mounds of the natalensis series do not agree with 
the historic illustration given by Smeathman, nor are they 
begun and built up after the manner which he describes. In 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 365 

fact, the only ones approaching Snieathman's picture have been 
seen at Winterton, Natal (PI. XXYIII, fig. 1) ; these were 
not more than 5 ft. high and 6 to 8 ft. through at the base. 
Nests may be said to fall into two categories, being either 
of ancient or modern formation ; the ancient types are repre- 
sented both by isolated mounds and by " nest-sites," the 
modern by rounded, pinnacled, pyramidal or conical masonry 
mounds. 

The modern mound, which occurs commonly in the grass 
veld (savannah) around Pretoria, is a wide, low, barren 
mound, regulai-ly domed ; it is built of fine earth particles, 
firmly cemented together, and presents an all but smooth 
surface. Such mounds are 3 to 4 ft. in diameter and 12 to 
16 in. high, and occur within 50 to 100 paces of one another 
(PL XXVI, fig. 8). Among them are to be found solitary 
ancient mounds of a roughly conical form, of 3 to 4 ft. in 
height, which are clothed with small, rough, stunted her- 
bage, the roots of which penetrate into the matrix of the 
mound and are not interfered with by the termites (PI. XXVI, 
fig. 9). 

Isolated modern mounds are not common along the Natal 
seaboard, where the species also abounds. Two, found in a 
wattle plantation at Mount Edgecombe in a light sandy soil, 
were regarded as less than seven years old. These (PI. XXVI, 
figs. 10 and 11), although composed of sand particles, were 
extremely hard, but they showed weathering. One, roughl}^ 
rounded, 2 ft. high and 3 ft. in diameter, was upon the edge 
of a slope and the lower base had extended 18 in. by 
weathering. The other, an acute regular cone, 4 ft. high 
and 4 ft. in diameter at the base, stood upon a flat area and 
was encircled by a small buttress of recently weathered-off 
pai-ticles, which extended 18 in. from its periphery. 

Sheltered in the natural bush at Bellair a recent mound 
was examined which, upon certain information gathered on 
the spotj was estimated at under ten years. This nest was a 
regular dome 6 ft. in diameter and between 2 and 3 ft. high 
(PI. XXVI, fig. 12). It was of extremely tough texture. 



366 CLAUDE FULLER. 

being formed from a clayey mould. Its surface was littered 
Avith dead foliage, and into its matrix the roots of the 
surrounding trees had penetrated, Avhilst from its summit gi-ew 
a sapling 10 to 12 ft. high with a stem 2 to 3 in. in diameter. 
This mound avhs grossly invaded by T. in cert us, whose 
fungus-gardens were in great abundance and seldom more 
than 6 in. apart. Pyramidal, pinnacled, and conical masonry 
mounds are to be met with on the highlands of Natal in great 
abundance; in favoured sites being seldom more than 100 
paces apart and often less than 60 paces (PI. XXVI, figs. 13 
and 14). 

The two Mount Edgecombe nests are described to indicate 
the genesis of those great termite ramparts which are prefer- 
ably referred to as " nest-sites." These, it is thought, are of 
very ancient formation indeed, and owe their huge bulk to 
the work of generations of terinites and the accumulations of 
organic matter from the plants which overgrow them. 

The bush fringing the sea-front of Natal contains an 
extensive flora, but in many parts the so-called wild banana 
(Strelitzia august a) predominates. As the bush extends 
back from the sea it becomes more and more sparse, until the 
glades cease in open country ; this, however, is dotted over 
with cii'cular patches of bush, standing oasis-like in the grass 
and giving the impression of an inland invasion of the coast 
bush ; this is usually alluded to as the " park formation " 
(PL XXVI, fig. 15). These oases are nest-sites. As they 
stand, they are islands of trees, palms, and strelitzias girt 
about by a fringe of dense low-growing plants, all so thickly 
intergrown as to be nearly impenetrable. Cleaned of the 
encumbering scrub, they ai-e found to be circular or oval 
ramparts of earth, 20 to 60 paces across and from 3 to 8 or 
10 ft. in height. Very few present any superficial evidence 
of termite work, although in all there is ample evidence in 
the huge burroAvs of the aardvark (Orycteropus cape n sis) 
that they are or have been recently inhabited by a colony. 
Occasionally atypical mound is found on the rampart, and not 
infrequently a nest-site harbours a huge nest of T. latericius. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 367 

Such nest-sites are even more common in the thick fringe 
of bush bordering the sea-shore, and upon the whole they are 
so abundant up to six or eight miles inland as to alter the 
regular contour of every slope and ridge ; a feature readily 
observed because of the vast sugar-cane fields now covering 
the country, once in part or wiiolly bush. Owing to the 
rapidly undulating nature of the country it follows that these 
nest-sites nearly always occur upon hill-sides, and for this 
reason they appear very much larger Avhen approached from 
below than from above. It has, however, been noticed that 
the upper side of a ridge is more favoured than the crest or 
lower down the slope ; hence, many a long ridge of gentle 
gradient when approached at right angles appears serrate, 
so rapidly does one nest-site succeed another down its length. 
The inhabited portion is iisually at the front apex of the site, 
and, by passing up a slope along the line of nest-sites, one 
progresses over so many huge steps of an earthen ladder 
(PL XXVI, fig. 16). 

The cleai-est cut example of a simple nest-site was found in 
front of a marine residence at Scottburgh. This was originally 
overgrown by strelitzia and phoenix, but had been cleared 
and surmounted by a garden-seat. This mound was regularly 
rounded, with a more or less flattened top 8 ft. in diameter. 
Approached from above, it was 5 ft. high but fell away in 
front 10 to 12 ft. The sides were fairly acute and the 
smallest diameter at the base 15 ft. The presence of a living 
community in this rampart was demonstrated. 

The conclusion arrived at regarding the paucity of isolated 
modern nests of T. natalensis away from old nest-sites 
along the Natal coast is that such is due to the better and 
very ample opportunities for nest-making which these ancient 
sites afford. 

The ordinary or modern mound is always perforated by 
almost vertical galleries which ramify upwards throughout it. 
The arrangement of these is complex and disorderly, they are 
large and either circular or elliptical in section, some of the 
latter presenting a long diameter of 4 to 6 in. All reach 



368 CLAUDE FULLER. 

nearly to the crust of the mound, and at times small tubes 
are driven from them, into the crust. What the purpose of 
the mound is, is one question ; what it is, is another. The 
mounds do not begin in a small way and gradually increase 
in size. No amount of field observation can discover the 
mounds of T. natal en sis in stages of growth below a certain 
minimum size,' and there is every reason to believe that a 
mound of some magnitude is built up suddenly. This may 
synchronise with that moment when the colony has attained 
a certain strength, and an extensive addition to its domicile 
is called for. Even the minimum mound (a primary) contains 
more nniterial than is represented by the nest-cavity below it. 
The material used for the building of a mound is, no doubt, 
obtained b}' increasing the diameter of existing galleries and 
extending them. Whatever it may be that leads to the 
formation of a fairly large primary mound, the subsequent 
growth of the mound is due to the fact that it is the dump 
for the soil removed in driving galleries through the soil ; 
and the bulk of the soil in a mound, of almost any magnitude, 
is the material removed in mining far-reaching stopes. 'J'he 
purpose of the mound is primarily for the protection of the 
nest from subsidence. From the position of the queen-cell it 
is obvious that the extensions of the cavity are for the 
greater part around and above it. This is evidenced by the 
fact that in the high mounds the upper region of the cavity 
is above normal soil level (PI. XXYIII, fig. 2). The ordinary 
mound, therefore, serves a double purpose, protecting the 
cavity from subsidence and submersion on the one hand, and 
jDcrmitting its upward extension on the other. Also the great 
shafts which perforate the mound are utilitarian, permitting a 
multitude of miners to reach the crust rapidly and super- 
impose an extra layer of cement whenever occasion demands. 
At the same time, they aid in aerating the whole nest-system. 

' The smallest nest I kuow of was found by Dr. Warren. This. I am 
informed, had a diameter of about 18 in. and a height of about 
15 in. The nest-cavity was considerably smaller than the mound. 
The (jueen-cell was found, l)ut the nest was entirely deserted. 



OBSERVATIONS ON SOJ[K SOUTH AFRICAN TERMITES. 369 

In this connection it is not urged that the mound-shafts are, 
in any sense, flues through which currents of air are always 
in circulation. 

Strongly as the earth particles are cemented together, 
mounds are subject to weathering, and their surfaces have 
to be repaired. As colonies ai*e subject to decimation, more 
particulai'ly by aardvarks, it follows that the mounds of 
weakened communities, and those uninhabited, weather 
rapidly. In the weathered stage grass creeps over them, or 
seeds of large plants, which have no great opportunity to 
become established in the grass, And a suitable nidus and 
flourish. Then the mound becomes the home of a fresh 
colony which reconstructs to its own accommodation. ' It is a 
succession of such phases which brings about in the course 
of time the huge nest-sites and oases, or park formation. 

The exploration of a large nest-site involves a considerable 
amount of labour. With a view to ascertaining whether such 
a nest-site bore evidence of successive habitation, one below 
the medium size was selected and thoroughly examined at 
Mt. Edgecombe, Natal (PI. XXYI, fig. 17). This stood in a 
wattle plantation, the bush having been removed some seven 
years previously when the land Avas put under wattles. 

The features of this nest-site were as follows. It was a 
large accumulation of soil, rising in the centre to a rough 
cone, and it presented, at two points on the circumference, 
aardvark burrows which penetrated inwards for 4 and 6 ft., 
going down to a depth of 2 and 3 ft. below the natural level 
of the ground. The whole mound, including the central cone, 
was perforated by galleries containing living termites, those 
of the cone resembling the typical ramifying galleries found 
in a normal mound. The whole of this mound was removed, 
and the nest of its occupants was found on the western slope 
well above ground level. 

No other colonies or traces of cavities were found, but 
there can be no doubt that such previously existed and were 
destroyed by the aardvarks. The site of the nest of the 
existing colony indicated that it originated fi'om a pair which 



370 CLAUDE FULLER. 

had penetrated the mound at that point. The mound on tlie 
whole displayed a weathered appearance, but the surface had 
been recently repaired immediately above the cavity, whilst 
parts of the cone had had some attention. 

Inferentially we may assume that it is possible for nests of , 
this species to exist without mounds; but as to whether there 
is any reason to suppose that under certain climatic condi- 
tions — such as along the Witwatersrand (5000 ft.) — the 
species never makes a mound, the writer is not yet prepared 
to make a statement. In this connection it may be men- 
tioned that the species has been taken destroying young- 
gum-trees (Eucalyptus) and feeding upon manure where 
no indication of a mound could be seen for a mile around. 
Again the species has been seen feeding upon grass in 
Pretoria where no mound could be found. This latter instance, 
however, rather indicated the existence of a colony that had 
not as yet built a mound than one that would never do so. 

It is not .yet possible to say for certain how a nest of 
natal en sis as an architectural structure begins; but it is 
not difficult to see how it could originate by observing the 
behaviour of young pairs, and their peculiar adaptability to 
their environment in nests built under houses, or in the 
earth-matrix filled in beneath cemented, bricked, or stone- 
flagged floors. When nesting beneath such floors natal- 
ensis does not need to build a protecting mound. As a rule, 
however, there are dump-heaps ; these may be either in free 
spaces beneath the house or even outside of it (PI. XXYI, 
fig. 18). 

A number of such cases have come under observation, and 
the nests arise out of the entrance of a pair through some 
interstice. In the case chosen for illustration, the queen-cell 
was just below the cement, and tlie hive extensions were 
below it ; the nest arrangement is inverted, the cavity being- 
extended downwards instead of upwards. 

What appears to be the ordinary course of events is for the 
sexual couple to burrow into the soil to a certain depth ; this 
is about 18 to 20 in. There thev form their brood-cell. When 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 371 

the brood matures, and as it increases, the nest is enlarged 
by the making of cavities around the original cell ; and these 
increase in number and size with the growth of the colony, 
whilst the original brood-chamber becomes the queen-cell 
and is enlarged from time to time to accommodate her 
growth. Evidence of this latter contention is supplied in the 
case of a nest examined for me at Barberton. This was 
indicated by a low and comparatively small mound, and the 
queen-cell (PI. XXVIII, ligs. 8 and 4) taken from it is a 
quite small, thin-walled structure, with a long diameter of 
60 mm. and a height of 10 mm. ; it is exceptionally entire, 
having no shelves bracketed to it. The soldiers and workers 
of this colony were normal in size and plentiful, but the 
queen was only 40 mm. long. 

The soil removed by a young colony in ni;iking the hive- 
chamber and driving tunnels is no doubt scattered loosely 
over the surface — this is usually what happens where this 
species is found feeding over the open veld on dry grass or 
the droppings of animals. The mound, it is thought, is only 
built when the hive-cavity has been enlarged upwards so 
nmch as to be threatened by subsidence ; in short, when it 
becomes necessary ; and it is because of this that really 
small and new mounds are never found. It is difficult to 
explain why in certain parts mounds are not built; but, as 
the parts of the country (the Witwatersrand) where there is 
some evidence of this are high in elevation (5 to 6000 feet) 
and bleak in winter, it may yet be shown that the species 
takes advantage of the naturally stony nature of the countiy 
to construct its hives beneath or amidst sheltering rocks, when 
the presence of a mound would become non-essential. In this 
connection it is to be noted that "svlien the species nests under 
tiled, stone, or brick-flagged floors it extends its nest upwards 
to the floors. Where these are not laid upon a strong cement 
bed, subsidence frequently occurs ; indeed it is no exaggera- 
tion to say that tbe characteristic unevenness of old brick 
floors to rooms and stoeps in Pietermaritzburg, w'here 
natal en sis abounds, is entirely due to this termite. 



"372 CLAUDE FULLER. 



'J'he Hive. 



When the nest of Termes natalensis is spoken of, the 
term necessarily includes the cavern in which the colony 
<3ongregates and propagates, together with the mound and 
the great stopes and galleries. Some precise term is required 
to express the chamber in which the community has its 
headquarters, wherein resides its parents, and all matters 
pertaining to the propagation of the young take place. No 
word is more applicable than '^' hive " ; but it must be 
remarked here that whilst the term signifies a part of a 
whole as applied to the nests of certain termites, it applies to 
the whole in others (parvus), and cannot be used, at all in 
connection with some (in cert us). 

Upon these premises the hive of natalensis may be 
described as a single sub-globular (sometimes ellipsoidal) 
cavern, which lies as a rule immediately below the centre of 
the mound. It is like a many storied house, a series of fiats 
rising tier upon tier from its concave floor to its arched 
■ceiling. The shelves of clay are slightly arched and attached 
strongly all around to the walls, being systematically 
bracketed thereto. Besides this they are supported by 
vertical columns so fashioned as to serve this purpose, and at 
the same time form stairways from storey to storey. 

In the lower hemisphere of the cavity the shelves are no 
more than thin laminee of clay, and but shortly removed 
from one another ; then, as storey succeeds upon stoi-ey, the 
space between each is increased, and the most spacious are 
those in the dome. As the distances increase so does the 
thickness of the shelf until it may be as much as a quarter of 
an inch. Similarly the pillars or stairways are increased in 
their dimensions. The Avhole of the partitioning is soft and 
moist and of a friable nature, so that stability is due to the 
architectural features; the Aveight being so distributed and 
the curvature of the shelves such that the higher and more 
ponderous do not fall and crash through the more fragile 
structure below. This feature is perhaps illustrated at its 



OBSERVATIONS ON SGML] SOUTH AFRICAN TERMITES. 873 

best in the case of the Pretoria nest (PI. XXVIIT, fig. 2) 
wherein the cavity was extended right into the dome, and the 
arching of the shelves quite exaggerated.^ 

The royal chamber is externally a flattened, rounded mass 
of clay, the exterior form being occluded by the shelves 
bracketed to it and by the grooves which lead to the tunnels 
through its walls (PI. XXYIII, figs. 5 to 8). It is suspended, 
as it were, at a central point in the lower hemisphere ; but 
not infrequently it partly rests upon some intrusive stone or 
root, so excavated that it projects into the hive. When not 
upon such foundation it seems to be in a very precarious posi- 
tion, but the exact support to such royal cells has not yet been 
sufficiently traced. 

The pillars which support or suspend each shelf are in 
reality stairways, excepting those which connect the top shelf 
with the dome of the cavity ; these latter can only be regarded 
as suspenders. 

Most of the stairways are canted slightly and twisted a little 
spirally. In section they are flattened, oval, and slightly 
grooved on the surface traversed by the termites. This groove 
causes the pillar to appear to be more spiral than it really is. 
The canting of the pillar and the pathway upon it are both 
factors in rendering the ascent easier. 

Upon the upper shelves the individual fungus-gardens are 
arranged, placed like so many loaves of bread upon a larder 
shelf. The lower and more restricted shelves at the basement 
of the hive have so far always been found empty ; and, if 
there is differential feeding of the young, it takes place in 
these compartments. 

The gardens present a granular formation throughout, the 
matrix and the entire surface showing coarse granulations. 
They are of varying sizes and heights, being of greater height 
in accordance with the available perpendicular space above 
them. Irregular as they are in shape, differing in height and 

' This nest was not examined by nie, but the series of insects in the 
collection bearing the same accession number are typical T. natal- 
■ensis. 



374 CLAUDE FULLER. 

in diameter — some being low and spreading, others more 
conical — they present a characteristic likeness one to another, 
and can only be likened to small, rough coronets. In their 
natural state they are soft and friable, but when dried they 
become hard and woody. Viewed from above nearly every 
garden presents a broad oval outline. 

The royal cell is not so spacious an affair as is that of some 
other termites (badius). It is broadly oval in outline with a 
flat floor and shallow concave vault. The distance from floor 
to roof in four examples before me is 18 mm., a space which 
is just sufficient to accommodate the body of the queen ; in 
fact, the queen may be snid to be compressed between the 
root' and flooi- of her citadel. The short diameter is 80 to 
100 nnn., and the long diameter 127 mm. 

This description of the hive is based entirely upon my own 
observations. Because of its incompleteness it is given with 
much reluctance ; and, if adding little that is new, is presented 
with the intention of showing that the structure provides 
many architectural problems well worthy of investigation. 

The Inhabitants. 

(1) The Queen. — Smeathman's picturesque account of the 
live queen of bellicosus is indeed a good one for uatal- 
ensis. The creature, when imprisoned in a box with 
attendants, doubtless behaves quite naturally for some time. 
She is very excitable and constantly changes her position, 
swinging her body from side to side, and progressing not 
forward, but somewhat around a small circle. To achieve 
this, the head and thorax are thrown right round to one side 
or the other until the head is brought in contact with the side 
of the abdomen. All the time the insect pulls forward with 
the full strength of her legs. The massive body soon reacts 
to the strain and pivots on its base. This continuous leverage 
of the body, only momentarily interrupted when a worker 
presents its quota of food, is varied but in direction, and is 
the result of enormous muscular effort. Indeed it mav be said 



OBSKKVATIONS ON SOME .SOUTH AFRICAN TERMITES. 875 

that the characteristic of this queen is that it is, and looks 
like, a superaiiimated muscle. Doubtless the effort to move is 
lessened by the compression of the insect in its shallow cell. 
The flaccid and immobile queens of T. badius lie in hisrh 
domed cells, and so the woi'kers can approach the eggs from 
all directions. The movement of the natalensis queen is 
perhaps not without purpose, as it ensures the depositing of 
the eggs more or less i-egularly around the periphery of the 
cell. 

Egg-laying has for its accompaniment an excessive muscu- 
lar actiou, independent of the movement of the body as a 
whole, and as involuntary as the act of breathing with man. 
This muscular action is complex and represented by the 
bulging, or great lateral expansion, of various regions of the 
nbdomen. The body is invariably very asymmetrical, and 
the bulging is quite erratic ; it is not a regular undulation 
proceeding from the anterior to the posterior region, and any 
inflations upon one side do not correspond or alternate with 
any progressing on the other. The dorsal region — a median 
area wider than the sclerites — remains taut, as does the 
similar ventral region. 

The eggs are produced spasmodically ; they may form into 
a small packet, which clings for a while about the vulva, or 
tliey may be squirted out in a quantity of fluid. Even those 
which are laid separately are embedded to some extent in 
a white matter. Although laid spasmodically, the intervals 
of rest are very short, and upon the average the queen lays 
140 eggs per hour. This is far short of Snieathman's 30,000 
in twenty-four hours; but, upon the other hand, it is in agree- 
ment with observations made upon T. badius, of which 
fuller particulars are given. 

It may be urged that the rate of egg-laying would be 
modified by the abnormal circumstances under which the 
observations were made. Whilst not prepared to concede 
this point, the figures may be doubled or even trebled for the 
purpose of showing how far short of 30,000 they still remain. 

The queen is frequently grossly infested with small and 



376 CLAUDE FULLER. 

very active Thysanura, and at times mites are found fixed 
to her body. 

(2) The King. — The queen is invariably accompanied by 
one male ; more than one has never been noticed. This 
creature, under observation, displays an extraordinary devo- 
tion to his mate. He seldom wanders more than an inch 
away from her ; constantly approaching her head and paying 
apparent court; then inspecting her flanks and eggs with 
preoccupied attention ; behaving, in short, as most male 
termites seem in the habit of doing, but never revealing the 
pui'pose of his existence. The male is always somewhat 
inflated with body-fat ; the conjunctiva of the sides of the 
abdomen being distended, but not that of tlie dorsal and 
venti'al sclerites. 

(3) Soldiers. — The main defence of the nest falls upon 
the minor soldiers, or at least appears to do so, because they 
are more numerous than the major soldiers. The major 
soldiers do certaiul}^ accompany the minors, whenever any 
breach of mound or inner citadel is made, but in small pro- 
portion. Upon the whole the bulk of the major soldiers 
retreat to the inner parts of the nest, the fungus-gardens and 
the neighbourhood of the royal cell. Both larger and smaller 
soldiers bite, and the jaws of the larger will meet and cut 
like a knife through the thick skin of the finger-tip. The 
principal enemy of this species being aardvark, it is extremely 
probable that the function of the greater soldiers is to 
lacerate the creature's nostrils and mouth parts, and by so 
doing guard the queen-cell from demolition. That the part 
played is effectual, in this regard, is doubtful. 

(4) Workers. — The workers, on the whole, retire when 
the nest is molested, many crowding into the queen-cell. They 
gather in force for repairing the breach within a very short 
time after the attack upon the nest ceases. An opening 
(about 2 sq. ft.) made in a nest one day at 3 p.m., in expos- 
ing half a hive, was completely closed by 8 a.m. the next 
morning. 

It is obvious, from the fact that this species permits plants 



OBSERVATEONS ON SOME SOUTH AFRFCAN TEllMETES. 877 

of any sorfc to grow upon and send their roots ramifying' 
tliroug'h the mounds, that it feeds naturally upon dead veget- 
able matter. It is only circumstances that render it a pesti- 
lential creature. Upon the whole it plays a beneficent role 
by soil improvement, and nowhere is this more evidenced than 
in sugar-cane fields, where the cane flourishes amazingly upon 
the nest-sites, giving* a greater yield and withstanding 
drought conditions better. On the other hand, a zone of poor 
cane is often noticed about such a site, which may be due to 
the concentration of org'anic matter by the termites ; or, in 
other words, the constant removal of all the dead parts of the 
gi'ass formerly present. 

Termes b a dins Haviland. PI. XXX, figs. 1-6. 

Termes b a dins is not in the strict sense of the term a 
masonry mound-builder; at the same time, under native con- 
ditions, nests are found in mounds which are so hardened that 
they might be described as masonry. If it were not possible 
to observe in and about Pretoria numerous nests without 
mounds, which are distinctly the work of one colony, it would 
be quite natural to describe the species as a mound-builder; 
at first, I found it possible to locate nests under nature con- 
ditions only when in mounds. Whilst no nests have been 
found in open lands which could, with certainty, be attributed 
to this species, still certain soil subsidences and deserted 
cavities which have been found cannot readily be credited to 
any other termite. Generally speaking, it may be conceded 
that any superficial indication of a nest would be obliterated 
almost as speedily as made in open country. This will be 
better understood from the description of the modern nests, 
of which six have been explored. 

The modern nests are found beneath the shelter of ever- 
green trees, shade-belts, and hedgerows; they are very 
common at Pretoria in the Zoological Gardens, in the principal 
parks, and in the grounds surrounding many residences. Such 
nests are readily detected by the series of close-set hillocks 



878 CLAUDE FULLER. 

which overlie them (PI. XXX, fig. 1). These hillocks persist 
not because of their inherent stability, but because of the 
undistui'bed positions in which they are located, and the 
shelter provided against the violence of rain and wind. The 
trees beneath which they lie being of comparatively recent 
growth (twenty to thirty years) the nests are necessarily 
modern, and it is not difficult to decide that during recent 
times the site they hold has not been previously occupied by 
any mound-building termite. The hillocks are not masonry 
moundlets; they resemble mole-hills except for the fact that 
the earth-particles, of which they are built up, are loosely 
cemented together. They are, therefore, of a temporary 
nature, and weather away more or less slowly. 

In the case of some nests, protected by hedges, mounds 
have already begun to form, and there is evidence that a 
large mound w^ould ultimately accumulate, if the hedges were 
left neglected and plants allowed to grow in the mounds and 
so bind together the earth deposits. 

Incidentally it may be mentioned here that badius, like 
T. natalensis, very frequently forms a nest in the soil below 
stone-flagged floors ; in these cases there are large accumula- 
tions of earth in open spaces below the flooring of the house 
or abutting the outside of the building. The deposit below 
the floor remains as it is placed, and is much traversed by 
tunnellings through which the results of further excavations 
are carried. Outside deposits are scattered or weathered 
away unless placed among plants and creepers growing 
against the exterior walls (PI. XXVI, fig. 18). 

Under nature conditions some nests are found in large 
rounded mounds which form the core of isolated thickets of 
trees or pai*k formations (text-fig. 10). Whether the colonies 
in these mounds are as ancient as the sites they occupy is a 
question which cannot be answered, but what is more than 
likely is that these tree-spots originated long ago upon mounds 
of natalensis or latericius, and the preseuce of the trees 
induced badius to colonise them. Further, it is evident 
that the mounds and thickets have only acquired their present 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 



879 



cliaracter after years of occupation by successive colonies of 
this and perhaps other termites. The thickets in Pretoria are 
composed of several varieties of acacias (Acacia horrida, 
robusta, and caffra predoniinating"), associated with them 
are other thorny plants, the more usual kinds being 
G3Mn nosporia buxifolia, Ehretia hottentottica, and 
the spiny Asparagus. In these mounds, which may reach 
2 ft. above ground-level and be 10 to 20 ft. across, the 
actual nest is above soil-level ; but the whole is generally so 
involved by the intermingling and twisted root-systems as to 
render any account of the interior structure impossible. 

Text-fig. 10. 




T. bad ins. Vertical section of nest in mound, involved in root- 
systems of many plants (diagrammatic), (x J^j.) 



In one instance, however, where a thicket had been burned 
over, the nest-cavity was found baked and preserved, the 
clay composing it being changed into a fairly hard red tile. 

In the case of the modern nests the structure is readily 
observed, and, whilst erratic, keeps to a general plan. The 
external feature is the presence of the series of hillocks 
already referred to. In some cases these are superimposed 
upon the mat of leaf-debris under the trees; when this is the 
case they can be lifted bodily from the ground. They are not 
solid, but are perforated by small tubular, twisting galleries 
which lead from the centre at the base and allow the termites 
to dispose of excavated earth by placing it as a new crust to 
the moundlet. 

The tenuous galleries all enter at ground level into a fairly 
large gallery or vertical shaft, which descends either by a 

VOL. 3, PART 2, 26 



380 CLAUDE FULLER. 

direct or indirect route into one of the extensions of the 
fungus-garden. As some of these shafts widen out consider- 
ably as they descend, it is extremely probable that they have 
much to do with the aerating of the nest, and are not solely 
roadways to the dump-heaps. 

The nest proper takes the form of a very large globular 
cavity with supplementary fungus-gardens which imitate 
those of Termes latericius, except that they are in wide 
communication with the centi'al nest, or it has many large 
irregular extensions around its periphery. 

The dome of the great cavity is within 8 to 10 in. below 
the sui-face of the soil, and as its vertical diameter may be 
as much as 2 ft. and the horizontal diameter 3 ft., and as it 
contains no strong clay-snpports, it is readily understood 
why the species displays such a marked predilection to 
establish itself at the base of a tree. Up to the present no 
nest has been examined in which at least one lai'ge root did 
not pass through the great cavity or sufficiently near to help 
in strengthening it. The interior walls of the cavity are 
always plastered about an inch thick with clay-cement, and 
a consistent feature is the strong cemented clay cupola 
roofing it. This is so strong that it can be completely 
exposed by removing the surface soil, and then, if cut around, 
it can be lifted oif in a solid piece like the lid of a caldron. 
It is not necessarily perforated, but, in the case of the nest 
somewhat diagrammatically illustrated (PI. XXX, fig. 1), 
one circular hole, three-quarters of an inch in diameter, gave 
access to a vertical shaft above it. The cavity is quite filled 
by a fungus-garden; this is like a giant sponge and obviously 
built in level layers, although its full arrangement is occluded 
by the reticulum of a clay frame irregularly disposed within 
it ; a frame Avhich is never attached to the cupola and which 
is not firmly joined to the walls. If the nest displays lateral 
extensions, as shown in text-fig 11, a and b, the fungus- 
garden branches, octopus-like, along the length of these. 
When supplementary cavities are present, they have broadly 
oval, flattened floors and are widely vaulted. The fungus- 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 381 



•gardens in them fill them completely, and if removed resemble 
irregular and more or less flattened loaves. 

The queen-cell (PI. XXX, figs. 2-5) is situated in the 
great cavity and, in those nests explored, in a peculiar posi- 
tion and always on small pedestals. The cell itself is within 
an irregular lump of clay, forming a i"ough column at or 
towards one side of the nest. Its site is indicated in both 
nests, of which diagrams and illustrations are given. 

In one of the two nests illustrated the pedestals of the 
queen-cell rested upon the stones which formed the bottom 

Text-fig. 11. 




T. badius. A. Yertical section of nest lying over boulders, 
showing lateral fungus-garden excavations. B. Ground plan 
of same. ( X -^^.) 

of the cavity, in another it was attached to a shelf which ran 
around the chamber at about half its height. In the thiee 
nests the joining of the upper part of the clay column to the 
cupola was perhaps more indefinite than as figured in the 
diagram, and the cell itself not quite so mncli a part of the 
wall. 

Between the two diagrams a considerable difference appears, 
and the diiference in the nest arrangement displayed is 
thought to be due to the difi^erent nature of the soil. In the 
case of the shallow nest it was found that underlying the 
site there was a layer of stone boulders. To some extent the 
soil between these had been removed and the spaces so made 
filled up with fungus-garden ; it was assumed that as the 
termites had not been able to excavate deeper they had made 
the larger horizontal extensions shown in the diagrams. 



382 CLAUD l!] FULl-EW. 

The nest-cavity shown in PI. XXX, fi^. 1, is to be regarded 
as the more noi-mal, being made in deeper soil, in which 
stones and boulders offered no obstacles to the architects. 
From the depth at Avhich the queen-cell was found it is 
reasonable to assume that the original part of this nest was 
tliat which appears in the section as an upper storey, while 
the lower region was subsequently excavated. Here the 
central cavity seems to have reached its maximum proportion 
of safety, and it would appear tliat the supplementary cavities 
were constructed to meet the full demands of the great brood 
of tei'mites being raised. 

The striking feature of the nest of this species may be said 
to lie in the fact that tlie cavern is not developed around the 
queen-cell, but to one side of it. The fungus-garden is 
closely applied about the queen-cell, and eggs are scattered 
freely in tlie centre of the fungus-mass for incubation ; there 
also young termites in all stages of growth abound. 

The king and queen are found in a chamber hollowed in 
the clay -mass. This is of large proportions, and characterised 
by the high dome. In three examples the long diameters are 
130-140 mm., the shorter 75-90 mm., and the heights 28-38 
mm. It will be seen that the cells agree with tliat figured by 
Sjostedt for T . t r a n s v a a 1 e n s i s. The cells are ver}^ different 
to those of T. natal en sis, and the great space above the 
queen is peculiar, seeing she possesses so flattened and flaccid 
a body. The king is active, but never departs far from the 
queen, even when the two are confined in a wider space than 
that of the cell. The queen, on the other hand, cannot move, 
and her body is an almost inert mass. She pulls and tugs 
with her legs, and the head and thoracic parts are as lively 
as those of the queen of natalensis. The involuntary mus- 
cular movements of the abdomen proceed as in the case of 
natalensis, but more slowly. When opened the queen-cell 
contains a fair proportion of soldiers, a few major workers 
and a multitude of minor workers. If a queen is transferred 
to an observation box with workers, the process of egg-laying 
goes on uninterruptedly, whilst the workers, after making 



OBSEliVATlONS ON SOME SOUTH AFRICAN TERMITES. 



383 



tunnels in the soil, carry off the eggs and place them in these 
prepared recesses; they will also start enclosing the queen in 
a new cell. The rate of egg-laying is the same as that given 
for T. natal en sis, and averages two to three per minute. 

The following is the record of egg-laying for a queen 
immediately after removal from a cell into the observation 
box. It is complete except for one short break of 20 minutes, 
when the eggs were lost, and gives a reasonable figure of 
4000 odd eggs for 24 hours. 



Date. 


Time, 


Period 


Actual 


Average 








count. 


per mm. 


Sept. 3rd. 1914 


4 p.m.: fpieen placed 
ill observation box 








Sept. 4th, 1914 


9 a.m. : eggs removed 


17 hi's. 


2360 


2-31 




9.2U a.m.: eggs lost 


20 mius. 


— 


— 




9.20 a.m to 9.35 a.m. 


15 mills. 


160 


10-6 




9.35 a.m. to 3.45 p.m. 


6 hrs. 10 mills. 


1683 


4-5 




3.45 p.m. to 








Sept. 5th, 1914 


9 am. 


17 lirs. 15 mills. 


2611 


2-5 




9 a.m. to 9.12 a.m. 


12 mills. 










9.12 a.m. to 9.23 a.m. 


11 mills. 


72 


6-5 


Totals 


41 hrs. 23mins. 


6886 


2-4 per 










mm. 



The king is not inflated with body-fat as is the king of 
natalensis. The queen resembles that of T. latericius, 
but is not quite so flaccid. The general colour of the body is 
cream, stippled with reddish brown^ except for the smooth 
wide bands down the middle of the dorsum and ventrum. 
The bands are partitioned by the sclerites and ornamented 
by an irregular broad brown strip. The stipplings of the 
remainder of the integument take the form of elongate stars 
arranged along the long axis of the body. Length of largest 
queen (alive) 90 mm., width of abdomen 24 mm., height 
9 mm. 

In examining the supplementary gardens of this species it 
was noticed that the lower portion was concave and somewhat 



384 CLAUDE FULLER. 

old in appearance. There is no doubt that in the case of 
these gardens enlargement is effected by the application of 
additional layers. It is reasonably supposed that a cavity is 
first made of some minimum dimension and filled Avith a 
fungus-garden ; when necessity arises, it is enlarged by the 
lateral extension of the floor and by a general heightening of 
the vault. At periods the core deteriorates and crumbles 
away, and so a concavit}' is formed. There is some evidence, 
however, that the worn-out part of the fungus- bed is 
renewed. 

T. badius shows a decided preference for the bark of 
trees, but only the dead bark is removed. To accomplish this 
the termites swathe the tree trunks with a thin shell of clay, 
carrying the shell to a height of even 20 ft. It feeds upon 
dead grass and the droppings of animals, and it is also a pest 
to wooden structures, wherever it can work undisturbed. 
Unlike other species it does not tunnel into the wood, but 
removes the surface, layer by layer, working as on tree trunks 
under a shell of clay. In houses it may destroy the flooring 
boards, removing the wood-fibre piecemeal from beneath, 
working along the length of the boards, but always working 
under a canopy of clay. 

One very unusual nest of this species was found in a dis- 
used stable. Here several sheets of corrugated iron had laid 
for some time upon some litter. On lifting the iron it was 
found that the concavities had been freed of debris and 
long, narrow fungus-gardens built within the spaces. In 
addition to this the colony had taken advantage of a space 
behind a sheet of iron leaning against the wall, and by 
building up clay partitions it had converted the space into 
a roomy cavity and filled it with a large fungus-garden. 

Mr. J. B. Gordon has sent to me a queen-cell (PI. XXX, 
figs. 4 and 5), with its living contents taken from a nest near 
Pretoria. This particular cell is not only remarkable for its 
size but also because it differs from any other in the collection 
in having a well-defined exterior. It can only be inferred 
that the cell was removed from an older nest than any 



OBSERVATIONS ON SOME SOUTH AFEICAN TERMITES. 385 

examined by the writer. The soldiers and workers were 
normal, and the queen smaller than others from very much 
smaller cells. Owing to the fact that the cell had been 
broken open at one end it was not possible to take its exact 
measurements ; but for all practical purposes it was 200 mm. 
long, 110 mm. wide, and 32 mm. high. 

Termes latericius Haviland. PI. XXIX, figs. 1-8; 
PI. XXX, fig. 11 ; PI. XXXI, fig. 6. 

The juvenile nests of this species are moundless and do not 
present the characteristic funnels or air-pits of older ones ; 
further, there is much reason to believe that colonies of some 
magnitude are established long before an}^ surface indication 
marks the site of the nest. Surface indications first take the 
form of simple crusts composed of clay particles cemented 
together fairly strongly. Such crusts present the character- 
istic cobble-stone appearance of all new termite work of this 
nature (PI. XXVI, fig. 19) and are spread more or less flat 
over the ground with but occasional and inconspicuous, 
conical elevations. Several of such were noticed in Pretoria 
and its environs during May and June (1914), and two 
occurred in a footpath; these latter did not develop, as the 
constant destruction of the crust seemed to disturb the 
makers. If the moi'e elevated portions of the crust are 
removed, they will be found to cap shafts of J to 1 in. 
in diameter ; possibly the beginnings of the air-pits. 

Lai-ge nests in Natal, the Transvaal and Orange Free State 
are always readily recognised either by the air-pits or by 
them and their chimneys (PI. XXXI, fig. 6). T. latericius 
is a true mound-maker, but air-pits protected by large 
chimneys have been observed without the least elevation of 
the soil level, as described by Haviland. There can be no 
doubt that the mounds represent the soil removed from the 
pits as well as from the extensive caverns made to accommo- 
date the main and supplementary fungus-gardens. Where 
mounds are absent the reason is not at all apparent. 



386 CLAUDE PULLER. 

The commonest type of mound is a wide, low elevation, 
usually from 10 to 20 or more ft. across, and of an eleva- 
tion of less than 2 ft. At irregular intervals on this mound 
are the apertures of the air-pits, which may or may not 
be surrounded by an elevated rim. Occasionally a conical, 
non-perforated, super-mound also marks the nest-site ; this 
may have a diameter at the base of 3 to 4 ft., and an 
elevation of 2 to 3 ft. (PI. XXIX, fig. 1, a). Some of the 
air-pits are roofed over v^ith a clay cupola, and, as far 
as observations go, remain permanently in this condition 
(PI. XXIX, figs. 1, d, and 2). Then they appear as low 
conical protuberances on the mound surface. It is certaiidy 
not the case, as given by Haviland, that the air-pits are 
roofed over in winter and opened in summer; moreover one 
rarely finds less than three pits, and over eleven have been 
recorded (3). 

The elevated rims arouud the mouths of the pit are usually 
flattened and rounded by weathering (PI. XXIX, fig. 1, b), but 
in the first place are constructed by the insects as regular 
turrets or chimneys ; the large super-mounds are, without 
doubt, the outcome of combined turret building and weather- 
ing. Equally, no doubt, the whole mound is the result of 
these two factors acting together over a long period of time. 

There can be no doubt that the large nest-sites of T. 
latericius are of very old origin. This is often evidenced 
by the trees sometimes growing upon them, which must be by 
their nature of no mean age. There is also evidence that the 
hives progress within the large mounds ; in other words, as a 
part becomes stale it is neglected and the main hive driven 
further afield through the soil. Very striking evidence of 
this was obtained in examining a large nest in Pretoria, the 
inhabited cavity being 12 ft. removed from the further 
extremity of the collapsed and deserted region it abutted 
upon. 

Termes latericius may often be seen at Avork in broad 
daylight renewing or extending the rims of the air-pits ; this 
they do by continuing the margin in the form of a thin 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMJTKS. 387 

wall in a nearly regular manner all round. The building- 
insects move up the walls of the air-pit in great numbers, 
like so many bricklayers, eacli adding its quota of material 
to the structure. Whilst this work is proceeding, the 
thickening of the rim is done by insects working within the 
chimney wall, and although the edge of the ring is quite 
thin whilst the work is in ])rogress, yet, when the required 
height is reached, the wall of the chimney is thick through- 
out. Chimney building is not conducted all the year round, 
but is frequently to be seen in progress during spring and 
summei". 

In most cases the chimney is seldom more than 6 to 8 in. 
high. The highest observed in Natal was 20 in., but in the 
Transvaal and Orange Free State some chimneys are carried 
up to between 2 and 3 ft. (PL XXXI, fig. 6). 

It has yet to be shown by exact experiment that currents 
of air pass up the taller of these structures ; but that the pits 
of which they are the chimneys are for the aeration of the 
nest there can be no doubt. Nor is there any doubt that the 
chimneys, or rims as the case may be, are for the purpose of 
preventing any inrush of water. It is seldom that the air- 
chambers are filled with soil by weather effects ; but instances 
have been noted — the nests so affected being on footpaths 
worn in the veld by the natives — where tori-ents of water 
during rain storms have poured into them. 

The air-chambers may be of great size and flask-shaped, or 
they may be but deep vertical shafts, in which case they have 
equally large branches (PI. XXIX, fig. l,a). They are 
associated with the large main hives made by this species and 
connect with them by only a very few small galleries, just 
sufficiently large for two insects to pass one another. It is a 
feature of the mound and super-mound of latericius that 
no galleries are driven through them, and this fact strongly 
suggests that the great shafts in the mounds of the 
bellicosus group act largely in aerating the hives, althouo-h 
they have no actual apertures. The main hives contain a 
vast extent of fungus-bed; but, as they are more than half 



388 CLAUDE FULLKR. 

filled with a complex array of clay-girders, brackets, shelves 
and stairways in and amongst which the fungus-beds lie, they 
baffle concrete description. Flung out all around the main 
hive a foot to six feet away are many supplementary fungus- 
gardens in which young are also raised ; a state of affairs 
almost suggesting differential feeding. These gardens are 
placed in typical high domed cavities (usually 6 in. in 
diameter and 4 to 5 high) with flattish and circular floors. 
The fungus-beds in these annexures are regularly built 
and always present an agreeable symmetry (PI. XXX, fig. 11). 
Their general appearance may be likened to large rosettes,, 
and they are noticeably built up tier upon tier ; each tier, 
decreasing in diameter from the base upwards, is circular and 
emargined with triangular, slightly deflected, rigid flaps. The 
fungus-beds are quite distinct from the spongiform loaf made 
by T. vulgaris and do not at all resemble those of the other 
species here described. 

Communication between the cavities containing the small 
supplementary fungus-gardens and the main hive is estab- 
lished by a single and simple, cylindrical tunnel, sufficiently 
large for a couple of insects to pass one another. There are 
no large main pathways leading out from the smaller cavities 
or even from the hive-cavity, and it is only with difficulty 
that the small tubes are displayed. These leave the hive at 
or near its base and, running almost horizontally throughout 
the greater part of their length, rise by short steps in the 
same manner as do the foraging galleries of Hodotermes 
transvaalensis. 

Whilst there are no conspicuous galleries radiating directly 
out from the hive, somewhere near the periphery subterranean 
runways are to be found. These seem to be a distinct 
feature and to be for the purpose of giving the insects ready 
and rapid access to the neighbourhood of the nest on the one 
hand, and the surface soil on the other, as well as to connect 
up with the system of peculiar outlying granaries which this 
species builds. As far as could be ascertained, communica- 
tion with the hive-cavity from a main runway is onl}^ through 



OUSERVA'I'lONS ON SOME SOUTH AFRICAN TERMITES. 389' 

a Dumber of small tubes ; and similar small tubes lead from, 
the main runway to the granaries. These main runways, as 
far as followed, were perfectly uniform. They lay at a depth 
of 18 in., went quite straight and did not deviate in their 
level. They had a flat pathway about a third of an inch wide 
and a vaulted roof, never more than a quarter of an inch at 
its highest point above the floorway (PL XXIX, fig. 3). 
From the vault of these tunnels an inclined and slightly wider 
gallery occasionally leads upwards to within a few inches of 
the soil surface; from its terminus a number of small 
cylindrical galleries ramify through the grass roots (PI. 
XXIX, fig. 4). 

The granaries excavated by this species are very interesting 
structures. Those that are small are easily described as 
globular cavities divided into two parts by a horizontal shelf 
(PL XXIX, figs. 5 and 6). In the larger granaries the 
shelf is also present, but the cavity is su fantasticall}^ 
partitioned in other respects that it baffies description 
(PL XXIX, fig. 7). Except in one instance where a few 
short lengths of green grass were found in the upper storey 
of the cavity, all of those inspected were empty. Never- 
theless, embedded in the soil all around the cavity (in some 
cases to a depth of 4 in.) were thousands of grass seeds 
perfectly preserved, as evidenced by the fact that they 
germinated readily when tested. It is more than likely that 
these granaries are destined to be filled with supplementary 
fungus-gardens, and the storing of the seed in the adjacent 
soil is perhaps to facilitate the rapid building of the nidus for 
the fungus. What the diaphragms are for one cannot even, 
speculate. 

The food of latericius may be described as very varied. 
The collection and storing up of grain does not appear to 
have been ever suggested For termites, although a feature in 
the economy of ants. T. latericius feeds commonly upon 
the droppings of aiiimals and upon dead wood, bark, and 
dried grass. It has also been taken destroying acacia seed- 
lings, and harvests green grass after its own fashion. In> 



390 CLAUDE FULLEE. 

harvesting grass it displays a remarkable predilection for a 
-variety known as Florida Grass, a fine and tender variety 
of Cynodon dactylon. Consequently it is a frequent pest 
to lawns in Pretoria and Johannesburg, where this grass is 
favoured. If not interfered with, latericius will establish 
itself in a lawn, and not only will keep it constantly mown, 
but will disfigure it utterly with the clay canopies beneath 
which it works. The insects first form a small canopy not an 
inch in diameter aiul cut the grass off beneath it ; as the 
herbage is removed the margin of the canop}^ is extended 
until it may be 6 to 10 in. in diameter. As soon as the 
canopy is fairiy large, the insects lose their timidity and 
venture out two to three inches from beneath its protection, 
mowing the grass evenly all round as if with a scythe. If 
disturbed, the workers desist and hasten for shelter. A few 
lose their way. Long after the workers have disappeared, 
the soldiers (about 1 per cent, of the herd) remain, obviously 
on the defensive, and their movements and attitudes convey 
the impression that they are trying to shepherd the stragglers 
back to safety. 

An interesting feature is found in connection with the 
•departure of the winged insects in the early summer. For 
this purpose special exits from the nest are prepared, which 
take the form of inclined oval galleries radiating from the 
hive (PI. XXIX, fig. 8, a, b, c, d). These have a wide dia- 
meter of 1 to 2 in. and a height of i in., dimensions which 
are retained until the inclines ypproach the surface of the 
ground, where they narrow down sharply to an elongate 
oval (1 in. by \ in.), and then taper narrowly into the exit. 
Just above the soil-surface is built an oval and expanded rim 
of clay ; this rests obliquely and forms a vantage from which 
the insects can take flight. 

Before and after the imagos emerge the position of these 
shafts is indicated by very small moundlets of hardened 
•clay. 



OBSERVATIONS OX SOME SOUTH AFRICAN TERMITES. 391 

Termes vulgai'is Hcwiland. PI. XXX, figs. 7-10. 

Mounds. — In Natal the masonry mounds of this species 
are common, and more nearly approach mounds of the 
natalensis group than do those of any other, being formed 
of earth particles strongly cemented together, and presenting 
a hard exterior and an interior with more or less vertically 
placed channels. Often the mounds are very inconspicuous, 
and the abode of a fairly large community may be overlaid 
by quite a small mound which looks no more than a hard 
clod of earth inset in the soil. A rough idea of the form of 
these and the nests below them has already been given (3) ; 
they need, however, more thorough investigation when oppor- 
tunity offers. Although nests of 3 ft. in elevation have been 
recorded (2), the highest met with at Pietermaritzburg was 
18 in., and as this took the form of a low cone, it was first 
thought to be a small nest of T. natalensis. Ordinarily 
the large mounds of this species are circular, widely arched, 
with a diameter of 2 ft. 6 in. to 3 ft. 6 in. and an elevation of 
4 to 6 in. No mounds of this species have been noticed in 
Pretoria, but, as mentioned elsewhere, a moundless nest has 
come under observation, and similar ones have been found at 
Platrand, iVansvaal, and Tylden, Cape. 

Nests. — Unlike the nests oi the natalensis series, which 
can be considered under three headings — mound, hive, and 
galleries — the nest of vulgaris does not lend itself to such 
treatment. One has to consider the whole and the relation- 
ship of the various parts to one another. This is perhaps 
best revealed by the description of a nest explored at Bellair 
(PL XXX, figs. 8-10). In this case the whole nest arrange- 
ment may be said to centre upon the fungus-garden, which 
took the form of a large, flatly pyriform loaf, resting upon a 
broad base and crowned with pinnacles, the central being 
the highest. It measured 12 in. in height and 18 in, throuo-h 
its greatest diameter. In common with other fungus-beds 
of this species, it was more spongiform than any of the 



392 CLAUDE FULLER. 

fungMis-gardeus of the other species under discussion, and con- 
sisted of a series of layers 1^ in. thick enwrapping a central 
core. Fragile as the garden is these layers separate readily. 
The chamber in which this mass lay conformed roughly to its 
outline ; the floor was overlaid with a clay mesh-work, bone- 
like in structure, which formed a raised skeleton platfoi-m. 
From this platform arose upright columns of clay — most 
reminiscent of arm and leg bones — which converged and 
connected indirectly with the roof of the cavity. This 
peculiar frame-work of clay, both in horizontal and perpen- 
dicular directions, was the scaffolding or skeleton supporting- 
the fungus-garden, although the latter was not in any way 
cemented to it. 

Immediately above the fungus-garden there was a narrow 
conical dome, hollow except that from it depended numerous 
thin clay arms ; these being the indirect connections of the 
thicker vertical columns which penetrate the fungus-garden. 
Some of these miiior arms had living grass roots at their 
■ cores, the roots penetrating the matrix of the mound from its 
periphery. As many hang rigid, like stalactites, it was 
conjectured that either the grass root was preserved by the 
application of a layer of clay, when the dome was excavated 
and during an enlargement of the nest cavity, or the roots 
were encouraged, by applications of clay to their growing- 
points, to descend through the hollow dome into the main 
upright and so strengthen them ; a conjecture supported by 
the discovery of roots in some of these latter. From the 
dome itself several wide shafts (1^ in. in diameter) led 
upwards into the mound, there broadening out, an inch 
below its surface, into conspicuous cavities. Apart from these, 
other shafts led up from the base of the nest around the cavity 
to the crust of the mound there tei-minating in a similar 
manner. Means of access to the cavity and other galleries from 
these shafts were provided by apertures at their bases and 
along their length, the openings being both large and small. 

The mound of vulgaris appears purely protective, and 
although repaired in the same manner as in natal en sis it 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 393 

represents but little more than the earth removed to form 
the cavity, and in many cases not as much. 

Although in most nests, examined somewhat cursorily, no 
supplementary fung-us-gardens were noticed, such were 
certainly present in the one discovered by accident in 
Pretoria. 

Termes incertus Hcujen. PI. XXIX, figs. 9-10 ; 
PI. XXXI, figs. 4-5. 

The Haviland notes give the followino- account of the nest 
of T. incertus. 

" This species generally forms its nest in the nests of the large 
species of fungus-growers, but it is sometimes found independently. 
In the latter case the nest may occur at a variable depth below the 
surface, from a few inches to several feet. Owing to the freedom with 
which kings and queens wander about when the nest is opened, it is 
difficult to determine how many there are in a colony or what are the 
limits of a colony. Occasionally three or four queens are found not far 
apart ; kings are less often found." 

This account rather tends to convey the impression that 
T. incertus prefers to nest in the matrix of the masonry 
mounds of fungus-growing termites and only occasionally to 
make nests independent of mounds. Far from this being the 
case it may be said, with a great deal of assurance, that wher- 
ever this species abounds it is just as common to the surround- 
ing soil as to any mound present. The masonry mounds, 
however, undoubtedly provide congenial quarters, as very 
few indeed are not riddled by i n c e r t u s . What is remarkable 
is that the larger termites tolerate their presence ; that they 
do so can only be because the presence of the smaller species 
and its tunnellings make no difference to the actual purpose of 
the mound. 

When incertus lives in the mound of another species, 
owing to the narrowness of the parts inhabited, its galleries are 
tortuous and no definite plan can be recognised. Nests in 
the field, however, although intricate, do present a general 



394 CLAUDE FULLER. 

plan which, whilst difficult to expose, has very constant 
features. 

As Haviland states, it is not possible even in the field to 
define the limits of a colony. In the account given of the 
behaviour of the winged imagos it is indicated that at Pretoria 
the earth is thoroughly occupied by the species, and it is no 
exaggeration to say that to a depth of 4 ft. the soil is pregnant 
with its fungus-gardens. 

In order to obtain some idea of the main features of the 
underground occupations of T. incertus numerous galleries 
were traced through the soil for considerable lengths, and it 
is upon the result of this examination that the foregoing con- 
clusions have been arrived at. 

Termes incertus feeds upon dead grass and will often 
attack plants which have died and may damage newly set out 
seedlings, especially when planted in virgin soil that has 
recently been broken up. It is also known to destroy fruit 
ti-ees and rose bushes. In this work, however, it retains all 
its subterranean characters and is very rarely found above 
soil-level, unless in a dead post. 

In grass lands it makes a network of thin tubular galleries 
through the soil amongst the roots, and secures its food under 
cover of inconspicuous clay canopies. From this upper net- 
work of galleries equally tenuous shafts are sunk deep into 
the soil. The tracing of these is very laborious owing to the 
difficulty of following so small a gallery, but many interesting- 
features are presented every inch or so along the length 
followed. These features are illustrated in a sketch (PI. 
XXIX, fig. 9) made of a portion of such a gallery whilst 
being examined. Before summarising these, however, it is 
necessai'y to say that the long lateral galleries subsequently 
referred to generall}^ wander through the soil for several feet, 
and connect with other descending shafts without displaying 
any such features. 

The descending shaft figured Avent downward at a fairly 
regular angle of 45°. Here and there, for an inch at most, 
it would fall almost perpendicularity, but this feature was 



OBSERVATCONS OX SOME SOUTH AFRICAN TERMITES. 395 

sufficiently rare to be conspicuous when it did occur. The 
chief feature to be noticed in the sketch is that, at every few 
inches, the gallery opens out into well-defined chambers 
[a, a, a). All of these chambers are elongated; and, being 
vaulted and having flat floors, they much resemble the 
■queen-cells of other termites (vulgaris, latericius, etc.) ; 
ordinarily they are 1 to Ij in. long, one half inch wide and 
a quarter to a third of an inch high. Some of them, even 
those within a few inches of the surface, contained young 
insects which were attended by workers. Such young being 
far removed from the fungus-gardens may have been under- 
going differential feeding ; but no evidence on that point 
was secured. Some of these chambers are junctions from 
which lateral galleries, or galleries leading to the fungus- 
gai'den cavities, go out. The majority, however, have but 
one entrance and one exit — the main down-shaft entering 
above at one end and continuing its course thiTjugh a hole in 
the floor at the opposite end of the cavity. These elongated 
cavities are orientated in various directions and occur in the 
compressed nests found in mounds. The second striking 
feature is the little pocket-caverns which subtend the de- 
scending gallery {h, h, h) and open directly into it ; they are 
domed, have circular flat Boors and seem to be rest-houses. 
These are the lenticular cells of Haviland. A further feature 
are slight fusiform enlargements (c, c) which occur wherever 
the short, radiating and always descending galleries to the 
fungus-cavities join the down-shaft, or where lateral galleries 
join it. 

Among the irregular features are cavities such as d, d, and 
the cavities e, e ; the latter are, however, regarded as the 
beginning of excavations for the accommodation of fungus- 
beds. 

The fungus-garden cavities are placed around the down 
shaft (PI. XXIX, fig. 9,/, and PI. XXXI, fig. 4), usually two 
to three inches away from it, or even more. These cavities 
are, as a rule, globular, although the floors generally tend to 
be flattened. With them the whole system looks like nothing 

VOL. 3, PART 2. 27 



396 CLAUDE FULLEK. 

else than the underground stems of a plant with many tubers 
hanging to them. There is only the one aperture to these 
fungus-garden cavities, and, whenever a tunnel leading to 
one is bi^anched, the branch leads to the cavity of another 
fungus-garden. These galleries always slant downwards and 
fall acutely on reaching the vicinity of the cavity, entering it 
at the side. Fungus-garden cavities do not vary much in 
size ; the largest have usually a perpendicular diameter of 
Ij to 2 in. and a horizontal diameter of 3 to 4 in. 

The fungus-garden nearly fills the cavity (PI. XXXI, figs. 
4 and 5). It is distinctly granular and crumbles easily, the 
pellets of which it is composed separating. All are coral-like 
in appearance, and may be compared, as regards folds and 
involutions, to the kei-nel of a walnut. In the case of most 
termites when, in digging, a gallery is broken through, the 
aperture is speedily closed with pellets of moist clay. T. 
incertus does not do this, but endeavours to bridge the 
break by building out a pipe of clay from the disturbed 
surface, constructing the pipe in the direction the lost part 
took (PI. XXIX, fig. 10). In trenches recently cut through 
soil inhabited by this species, thousands of these projecting 
pipes are to be seen. As a rule the structure is not continued 
more than an inch, but some pipes projecting 2 to 3 in. 
have been noticed, and some were even branched. They 
are quite tenuous, with the outer surface roughly granular 
and inside quite smooth. 

Eutermes parvus [Haviland). PL XXIX, figs. 11, 12; PL 
XXXI, figs. 1-3. 

According to the Haviland notes E. parvus is found in 
Natal " from the sea-coast to an altitude of 5000 ft., and the 
winged forms have been taken on the tops of hills of 6000 ft." 
The species is only known to me from nests collected in the 
bush-lands of the Natal coast, and from specimens kindly 
collected for me by j\Ir. F. W. FitzSimmons at Port 
Elizabeth, where the species appears to be abundant. 



OBSERVATIONS OX SOME SOUTH AFRICAN TERMITES. 897 

Haviland noted furthei" that ''the nests are sometimes built 
practically on the surface of the ground, but they are often 
some little distance below, occasionally several feet, the nest 
beinfy an irregular cellular structure." Up to the present 
only the surface nests have been found, but somewhat similar 
nests of a closely allied species abound in the soil of the 
Pretoria district ; these range from the size of a large egg to 
that of a large cocoanut, and may be either superficial, 
beneath stones, inset in the mounds of other termites, or 
buried to some little depth in the surface soil. 

Without doubt sandy dunes, covered with scrub, are the 
sites in which parvus flourishes best of all. At Scottburgh, 
Natal, several nests were found amongst beach plants within 
20 ft. of high-water mark, and one, but 15 ft. off, overhung 
the beach (PI. XXIX, fig. 11). This was 11 in. in diameter 
and quite globular. Originally built in the loose sand and 
involving in its matrix several stems of beach plants, under 
the stress of wind and storm it had become quite exposed. 
The insects inhabiting it established communication with the 
sand dune, from which it depended, by means of covered 
ways along the supporting tangle of roots, as well as through 
the interior of those that were dead. All other nests were 
found loosely embedded in the soil ; so loose that they were 
easily extracted whole The top of these nests usually pro- 
jects a little above the surrounding soil, and has the appear- 
ance of a weathered and rotten piece of black wood. Domed 
nests are sometimes to be found, and these are obviously of 
recent structure (PI. XXIX, fig. 12, h). The domes may be 
3 to 4 in. above soil-level and have a cobble-stone surface 
of an ashen grey colour (PI. XXXI, fig. 1). The underground 
part of every nest found tended to become obconic, some 
being elongated and others quite squat. In one instance a 
domed nest when extracted resembled in shape a huge carrot, 
18 in. long and 4 in. through at its widest diameter. 

The nests of parvus contain no provisions of an}' sort, but 
the workers are animated food-reservoirs. The structure of 
the interior is cellular ; the cells being for the most part 



398 CJ.AITDE FULLER. 

uniform in size and irreg'ularly oblate. Each cell lias, as a 
rule, two apertures. In the core of the nest is a large 
flattened and horizontal queen-cell (PL XXXI, fig. 2), and, 
almost invariably, above this is a series of large inclined cells, 
which can be seen in the vertical section illustrated in 
PI. XXXI, fig. 3. These cells form a kind of stairway, up 
which the queen can and does progress. Whilst she has not 
been detected laying her eggs in these superior cells the 
quantities of eggs located there indicate that she migrated 
to them for ovipositing. If this is the case it maybe assumed 
to overcome the difficulty which would arise from too great 
an accumulation of eggs in the queen-cell. The largest nests 
found had a surface diameter of 12 in. and extended 8 to 
10 in. deep into the soil. 

In the scrub-lands parvus feeds upon decaying wood; 
the allied species in the Transvaal feeds upon grass, and will 
also attack young trees, hollowing out the roots and other 
underground parts. 

The first impi'ession which the nests of parvus gives is 
that they are formed by the conversion of dead stumps, and 
whilst I am inclined to think that this sometimes occurs there is 
sufficient evidence to show that it is not necessarily the case. 

Eutermes bilobatus [Haviland). PI. XXXI, figs. 7-9. 

This species normally inhabits a clay hive, portion of 
which is thrust above ground in the form of a low rounded 
mound 3 to 5 in. high and 5 to 7 in. in diameter; the whole 
interior is cellular; and there is no differentiation or partition 
between that portion in the confines of the mound and the 
subterranean region which it overlies. Its striking external 
feature is the resemblance borne by it to a water-woi'n 
boulder inset in the soil ; a feature which is not only one of 
appearance but also of hardness as well (PI. XXXI, fig. 7). 
The crust of the dome is nearly always one-half to three- 
quarters of an inch in thickness; under this the structure 
is cellular, and in this form the hive is continued into the soil 



OBSERVATIONS uN SOME SOUTH AFKICAX TERMITES. 6^^ 

to a depth of 6 to 8 in., the diameter of the subterranean 
part being invariably greater than that of the base of the 
mound. All the cells are of about one dimension except for 
one, or perhaps two, central cells, which lie two inches below 
soil-level. Tiiese are wide and shallow, and are the head- 
quarters of queen and king. The rest of the cells are very 
irregular, but tend to become flattened spheres ; each foi-ms a 
separate compartment communicating with adjoining com- 
partments by two or three small perforations in the walls. 
The walls between the cells may be described as strong, and 
of a fairly uniform thickness. The perforations have a small 
diameter of 2 mm., and are counter-sunk into each wall. 
There is no series of large permanent galleries leaving the 
nest, and such galleries as do lead away are fine and not 
easily traced. In the large series of nests (both normal and 
abnormal) examined nothing in the nature of stored pro- 
visions has been met with. Upon the other hand, the 
multitude of workers, always present, invariably have their 
abdomens so grossly distended that they can only be looked 
upon as animated food-reservoirs. 

When observations were first begun upon this species the 
frequent intermingling of its mounds with those of E. triner- 
vius, and a general resemblance in the relation of the mound 
to the subterranean part together with other broad structural 
features, led to a suspicion that bilobatus invaded small 
deserted nests of trinervius, as indicated by Haviland, and 
converted them into a structure to their own liking. Then 
the discovery of a small mound inhabited in part by the two 
species, one half built to the bilobatus pattern, the other to 
that of trinervius, led to a series of observations which 
showed that bilobatus simply expels young trinervius 
colonies from their own mounds. The process is nut, so far 
as can be seen, one of decimation : bilobatus simply gaining 
an access to one point of the trinervius mound, and then by 
gradually converting the more open trinervius galleries 
into cells, it slowly builds the original artificers out. This 
process often results in the building of a new mound some 



400 CLAUDE FULLER. 

little distance off by the trinervius community ; in several 
instances this has been begun before the colony has been 
altogether dislodged. 

It is not argued that bilobatus raids the nests of trinei'- 
vius, although upon two separate occasions correspondents in 
Natal have sent to the writer specimens of bilobatus, which 
they had taken from columns of workers found migrating in 
daylight over the surface of the ground. It is rather thought 
that just as pairs of trinervius will burrow into a hive of 
bilobatus so couples of the latter will bui^row next to a hive 
of trinervius, and when the latter is not exceptionally 
strong their progeny will evict it. 

As nests of bilobatus are also found under stones it is 
concluded that they take equal advantage of the nest struc- 
tures of a nasutu Eutermes, closely allied to trinervius, 
which nests in such situations. 

One extraordinary and large nest with a globular mound 
found in a wattle plantation at Mt. Edgecombe cannot be 
accounted for unless it was originally a large clod of earth, 
left when the land was broken up for planting, which had 
been converted into a nest and enlarged. Probably the 
abnormal conical mound, of which form only two examples 
have been observed, shown in PI. XXXI, figs. 8 and 9, is only 
abnormal because rare. 

E, bilobatus nests have been found completely surround- 
ing those of the undetermined species allied to E. parvus. 
These are similar in architecture to those of bilobatus, but 
being built of carton, and owing to their ligneous nature, 
bilobatus does not invade them. Moreover, the other 
species possesses passage-ways through the bilobatus nest; 
using the cells and communications of the latter but lining 
them with carton. Whether this inference is correct, or 
whether the undetermined species builds its nests in those of 
bilobatus, gradually replacing the clay by their own building 
material, it is impossible to say upon the evidence at hand. 

Here it may be mentioned that the process of dislodging 
carried out by bilobatus upon E. trinervius has its 



OBSERVATIONS ON SOME SOUTH AEKIOAN TERMITES. 401 

parallel in the attack of true ants upon them and other 
similarly nestiag species. It is scarcely possible to find a 
normal mound of bilobatus about Pretoria which has not a 
nest of a true ant in part of it ; the ants capture the nest in 
the course of time by taking possession of it cell by cell. The 
process is not, however, a rapid one, as the termite can build 
up quicker than the ant can break down, and the ver}' art 
which enables this termite to supplant another of its kind is 
its most potent means of defence Avhere the ant is concerned. 
When a hive of bilobatus is broken into the ants 
excitedly swarm in at once and seize and catry off the ter- 
mites ; they will not attack an injured nest of trinervius 
with like avidity, and seldom make tlieir own domiciles near 
to the mounds of this latter species. Indeed they evince 
great circumspection, and always endeavour to capture the 
trinervius nasuius from behind. 

The queen of bilobatus has the abdominal region vastly 
•eidarged, but it is somewhat vermiform, presenting a number 
of irregular bulges of the fai'-stretched counecting membrane. 
Although so much enlarged and cumbersome the queen is 
able to make rapid progress, and when exposed essays to 
escape deeper into the nest. The apertures in the cell-walls 
are far too small to allow her abdomen to pass through 
unimpeded, but it is drawn through by the strength of her 
exertions, being constricted in the process to half its 
diameter. The eggs are found in packets in different parts 
of the nest, to which they are carried by the workers. 

The workers comprise the bulk of the colony except when 
the nymphs of winged adults are present. When the nest 
is broken some hide away, but the majority of those exposed 
make no attempt to escape, crawling aimlessly about. Later, 
if unmolested by ants, they will start and repair the structui-e 
with pellets of clay -cement voided from their bodies. 

The soldiers are always very few in number — often no 
more than a dozen can be found in a nest, at times they are 
entirely absent. This is especially the case when the imagos 



402 CLAUDE FULLER. 

are present in the nest. Since after the nuptial flight has 
taken place callow soldiers are soon to be found^ it is assumed 
that for some reason the soldiei's are destroyed at this peiiod. 
The soldiers are extremely cowardly, and always retreat 
rapidh^ when a nest is broken into. This, together with 
their scarcity, seems to indicate that bilobatus,asa species, 
is not in any way dependent upon the soldier caste, and the 
representatives which do occur are but relics of a former 
economy. 

The winged insects are of two sizes, the male being very 
much smaller than the female. When the adults are present 
ill the nest it has a foetid and disgusting odour — a feature 
never to be noticed at any other time. 

Eutermes trinervius [Rambur). PI. XXIX, figs. 13-17; 
PI. XXXII, figs. 1-2. 

Apart from South Africa, the species ranges far afield, and 
it is very widespread throughout the sub-continent, being 
the commonest and most prevalent kind. 

It occurs all over Natal from within 100 yds. of high- 
water mark to altitudes of 5000 to 6000 ft. It is, however, 
much more abundant in the drier parts of the midlands (Klip 
River County) than elsewhere, and is least common on the 
higher altitudes. 

It is very common in the Transvaal, and abounds on high 
altitudes (5500 ft.) under fairly severe winter conditions, 
with the difference that these altitudes receive less rain and 
mist than regions of the sa.me altitude in Natal, and are not 
mountainous. In the Cape Colony it ranges from the Orange 
River to the southern coast, but does not seem particularly 
common in the extreme south-west. In the Karroo it is to be 
met with, but always more abundantly in areas where 
summer grasses gi'ow. 

In the Orange Free State the species abounds from east to 
west and north to south. Through the middle region of the 
plateau, which has a mean altitude of 4500 ft. (ranging from 



OBSERVATIONS ON SOMK SOUTH AFRICAN TERMITES. 403 

4000 to 4900 ft.), it is incredibly abundant; so much is it so 
that one may travel by train for a whole day through country 
so thickly studded over with mounds as to look as if overrun 
by ii multitude of grazing- sheep. This extreme abundance 
on this wide plain, hot in summer and bleak in winter, 
indicates that the species flourishes best in a dry atmosphere. 
Here the denudation of grazing land caused by the presence 
of these numerous colonies, proceeding year by year, must 
impoverish the land, but the species woi'ks in so insidious 
a manner that it is extremely doubtful whether those most 
affected — the stock owners — have the slightest idea of the 
loss entailed. 

The nest of trinervius is a "hive-nest" assuming the 
form of a globular mass, the lower quarter of which is 
embedded in the soil. In Natal the mounds are symmetrical 
half-spheres, whilst the base inset in the soil is a low 
in\erted cone. Regularly rounded domed mounds may also 
be found in parts of the Transvaal and Orange Free State 
Avhere moister conditions obtain, and this is the prevailing- 
shape in the southern region of Cape Colony. Where 
conditions are drier and the summer rains come in fierce 
tropical showers, regular contours are uncommon, and the 
mounds are roughly shouldered and tend to become conical. 
This alteration of the contour is due to spasmodic additions 
to the nest, coupled with weathering ; the colonies under 
these conditions adding humps to the nest and not making a 
crust which envelopes at least two-thirds of the surface, as 
they do under milder and more uniform climatic conditions. 
Along the high ridge (5000 ft.) of the Witwatersrand 
(Heidelberg to Germiston) about half the mounds have a 
somewhat bizai-re appearance (PI. XXIX, fig. 17), and on 
the summit of some of these small or quite large tapering- 
cones are developed. There seems to be nothing to account 
for this local eccentricity. It may be said of this species 
that it does not make bricks without straw, as a considerable 
amount of grass is always incorporated into any new addition, 
of the nest. 



404 CLAUDE FULLER. 

There are several classes of mounds. The first of these is 
the juvenile mound^ obviously the domicile of a young colony. 
These nests appear as collections of clay amidst a tussock of 
:grass. From this stage the nest develops into a small 
domed domicile with the soil about the periphery of the 
mound perforated in every direction by a mesh of galleries, 
much like those of the hive itself. From this second stage 
develop the larger mounds. Around the large and old 
mounds the network of galleries is absent, but replaced by 
.radiating trunk routes to feeding grounds awaj' from the 
nest. The remaining class is the mound built up by a colony 
after having been expelled by bilobatus, or by the destruc- 
tion of the mound, when it is frequently erected on the same 
site or hard by. Under such circumstances a supplementary 
mound may be formed. These mounds are built quite 
rapidly ; in one instance, where a series of small nests was 
exposed for queens, it was found that those colonies, in which 
the queen had escaped detection, had, a week later, built 
mounds equal in size to those which the}^ were previously in 
occupation of. 

Owing to the difficulty in detecting queens in large nests 
it was thought for some time that a queen did not inhabit 
every mound, but this point was kindly cleared up for me 
by Mr. Leonard Bagshawe-Smith, of Platrand, Transvaal, 
upon whose farm the investigation, to be detailed below, 
into the connections between the mounds was carried out. 
Writing to this point under date of August 25th, Mr. 
Bagsh;iwe-Smith sa.ys : " Yesterday, I took out fourteen nests. 
In thirteen I found queens, and in the fourteenth, a fairly big 
mound, I found a king. In one I found both king and queen. 
If you dig the big mounds out very quickly you find the 
queen; if you take time you will not. The queen gets away 
very quickly ; with a quick movement she slides and falls 
from gallery to gallery and gets below soil level. I had a 
good opportunity of watching one sliding and falling, with a 
crowd of workers after her." Again, under date of Septem- 
ber 2nd, he writes: "I took queens from each of six nests 



OBSERVATIONS ON SOME SOUTH AFKICAN TERMI'l'KS. 405 

that were connected by runways. In the first series we 
examined I had great difficulty in recovering some of the 
queens; again and again I was about to give it up, bi^it kick 
favoured and I found one queen over 12 in. below soil- 
level." 

Haviland states that there is only one king and one 
queen to a nest, that they generally live near the centre, and 
when disturbed may escape into the surrounding soil. As a 
matter of fact, the king is very seldom detected, whilst the 
queen moves with such agility thac even when one has seen 
her enter a lump of nest this often has to be broken to 
fragments to capture her, so quickly are her efforts at con- 
cealment manoeuvred. Placed upon a level surface the queen 
cannot get along at all rapidly, but she can climb a wooden 
upright. From this and the foregoing observations it is clear 
that the insect is adapted to rapid progress in the peculiar 
and involved galleries of the nest. Two queens have been 
taken in one nest, but this is unusual. 

If queens are removed directly from nests and placed with 
nest fragments and workers in a glass jar, egg-laying can be 
readily observed, and the workers attend to the queen as if 
nothing had happened. Feeding is continuous, and it is 
seldom that there are not more than two workers in atten- 
dance. The queen is fed with some reddish fluid of which 
but a minute quantity is passed from the mouth of the worker 
to that of the queen. An individual worker may give one, 
two or three sips before retiring, and the queen may take two 
or three sips from one attendant before turning to the other, 
■or she may sip from them alternately. A succession of 
workers attend to the feeding, and when one retires another 
takes its place. 

The eggs are extruded one by one and stick together in a 
little parcel about the vulva. Never inore than ten to twenty 
seem to accumulate, as workers aie constantl}' removing 
them. The eggs do not separate away from one another 
readily, and the workers endeavour to remove them with the 
maxillte and not with the jaws ; it is only when the maxillte 



406 CLAUDE FULLER. 

fail that the jaws are used. When the egg is sepai'ated it is 
carried off in the mouth, projecting out between the jaws. 

The celluUxr arrangement of the whole of the nest, whilst 
appearing in section (PI. XXXII, fig. 1) as a series of layers, is 
so intricate that it baffles concise description. Such words as 
" honeycombed " or " spongiform " do not express it. It is 
better described as a labyrinth of galleries which cross and 
recross one another and go in all directions. It has been 
found that, starting from a given point, a termite can 
progress to any other given point by a multitude of routes. 
Thus a termite at one point can reach any other by an 
almost direct route straight through the nest, or it can take 
a curving or zig-zag route to right or left, restricted only by 
the outer confines of the hive ; equally it can take either of 
such routes by a series of ascents and descents or vice versa. 
There is, in fact, no limit to the possible ways of journeying 
from one part to another. In its arrangement the whole 
nest can be likened to the fungus-beds of such species as 
T. badius and vulgaris. The nest structure throvighout 
is formed of gritty earth particles firmly cemented together, 
but the hardness of the nest is largely dependent upon the 
nature of the soil from which it is built, those in sandy loams 
being much the softer. 

From the larger mounds numerous galleries are driven 
out through the soil at one inch below the surface; these 
are quite straight for some distance, and resemble the 
spokes of a wheel, with the mound for the hub. Their 
number is variable, but as many as twenty have been found 
radiating from a mound having a diameter of 8i ft. Many 
of them anastomose within a few feet of the mound. For 
great lengths these galleries and their branches are of a 
permanent nature, and form a succession of straight short 
lengths or very wide curves. They are uniformly an inch 
below the surface, and rise and fall most regularly with the 
details in the contour of the surface. They have a uniform 
width throughout, have hardened floors, and are vaulted 
above (PI. XXIX, fig. 14). They connect either directly 



OCSEKVATIOXS ON SOME SOUTH AFIUOAN TERi\[ITES. 407 

Avitli other moinuls or lead to feeding grounds. At irregular 
intervals along their length, seldom at a distance of less 
than two inches apart, pouches are excav;ited on both sides 
{PI. XXIX, figs. 15, 15a, 15?), 15c, Ibd) ; these extend outwards 

Text-fig. 12. 




Eutermes trinervius. Sketch plan of system of galleries 
showing nest connections and palmate termini. All the 
galleries in the triangle formed by nests b, k. and G are ex- 
posed. Only around nest B are all the radiating galleries 
exposed. Actual distances apart of nests — A to E. 120 ft. ; a to 
K, 102 ft. ; K to E, 102 ft. ; B to C, 6 ft. 



and downwards, and usually undercut the path. Where, by 
chance, two such pouches are opposite each other the path- 
way is often completely undercut, so that it forms a bridge 
across a small chasm. These pouches are all snb-globular, 
and, as a rule, measure 1 in. in the long diameter, and bulge 
out about three quarters of an inch from the edge of the 
pathway. The apertures into them, fi-om the side-walls of 



408 CLAUDE FULLER. 

the tunnel, ai*e always an elongate oval. Their purpose is 
for the temporary accommodation of the night's harvest. 

A peculiar feature of the galleries is that at every minor 
branch the connection of the branch with the main gallery is 
an indirect one, and occurs between two and three inches deep 
in the soil ; this connection can only be compared to a knot. 
Galleries which approach a feeding ground suddenly taper 
and branch in a palmate form, each branch running out to 
the surface. The surface exits are small round holes of a 
fairly permanent character which have been noticed to remain 
open for over a week. There is some evidence that feeding- 
grounds are more or less permanent, and that continued 
feeding about one point results in barren or bare patches. 

The accompanying sketch illustrates one of a series of 
observations upon galleries and nest-connections. It is drawn 
approximately to scale, and shows, as nearly as possible, the 
directions taken by the various galleries. Tlie mounds 
A, B, c, D, F and K were inhabited by active colonies, and in 
each a queen was recovered. Unfortunately this series was 
rather near to the homestead, and some of the nests had 
been removed for chicken food. The sites of these are indi- 
cated at G and h ; the galleries ran through them, descended 
below, and rose again on the other side of the site ; from this 
it is concluded that the nests when standing formed part of 
the series. It will be obvious that only a few of the galleries 
have been exposed, but the galleries are not more abundant 
than indicated in the rough ti'iangle formed by nests b, k, g, 
which were all exposed. If the evidence revealed is followed 
to its natural conclusion it would appear that all the nests 
except E formed a complete series. Nest e appears to belong 
to a different series. Point is given to this by the obvious 
avoidance of the galleries of the nest d by that from e. Again 
the partly exposed gallery x x, obviously leading to the 
gallery connecting nests c and d, was deliberately under- 
mined by the second gallery from nest e. 

It is to be noticed in the foregoing account that all the 
nests explored harboured colonies ; this was fortunate, as it is 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 409^ 

seldom there is not quite a large percentage of deserted 
mounds. Why so many mounds are deserted is very baffling. 
Dr. Warren tells me that, in the course of his study of the 
biology of this species, he endeavoured to sustain his colonies 
by watering the nests through the dry winter; but, not- 
withstanding this attention, many became deserted. Dr. 
Warren was fortunate enough to see the action of desertion, 
and by his express wish his account is incorpoi^ated here. It 
should be stated, however, that the observation was made 
prior to our full acquaintance with the radiating ga.Ueries. 

One day during the winter of 1913, whilst visiting the 
ground where he had a series of nests under frequent 
observation, he noticed a series of small holes 4 to 6 iu. apart 
and a quarter of an inch in diameter arranged in straight 
lines all radiating from the mound (PI. XXIX, figs. 16, 
a, h). Upon looking into these holes it was seen that the 
termites were hurrying backwards and forwards in an agitated 
stream. Quite unexpectedly, the insects — nymphs, workers, 
and soldiers — poured out of the radiating channels, scattered 
over the surface, and before long died. Some that were 
taken and placed in test-tubes with moist earth lived a fort- 
night longer. In addition to this abnormal behaviour, small 
holes about a quarter of an inch in diameter vvere formed 
irregularly over the mound and more particularly on the 
sides. These holes might be plastered up again with moist 
pellets ; when open they were always fringed by a border of 
soldiers within, whose closely placed heads, projecting beyond 
the edge of the hole, formed a couspicuous reddish border to 
the aperture, as viewed from the outside. 

Subsequently at the sources of the Crocodile River, and 
again at Platrand and at Pretoria, the writer found, during 
the winter of 1914, many nests showing these radiating liues 
of holes. The natural inference was that they were simple 
perforations in the roof of the galleries ; but upon investiga- 
tion it was found that the galleries had been deliberately 
altered and the floor there raised towards the soil-level, the 
apertures being immediately over the raised part of the runway. 



410 CLAUDE FULLER. 

The reason for this extraordinary manifestation, limited as 
it is to some nests only in a series, is very elusive. It can 
scarcely be due to the queen going astray in the galleries or 
to her gaining access to another nest, although it is possible 
for her to do this. If it is due to her death, one would 
expect that the inhabitants of the one mound would simply 
augment the colonies of those with which it is in such intiu;ijite 
communication and not commit suicide. The circumstance 
caniiot be credited to drought or cold, for many neai'-by 
colonies no better circumstanced continue to thrive ; nor is it 
from shortage of food, as Dr. Warren found ample provender 
in the deserted nests, whilst a similar state of affairs obtained 
at Plati-and and Crocodile River. 

The speedy death of the insects on outpouring from the 
nest is equally remarkable ; not alone because those captured 
and placed in a test-tube lived for so long, but also because, 
if a nest be bi'oken and the insects exposed to the glaring 
sunlight, they do not die. The workers simply seek shelter 
under the broken fragments strewn around or in the recesses 
of the nest, whilst the soldiers expose themselves for hours 
unmindful of danger from foe and uninjured by the sun- 
shine. 

On the face of it, this extraordinary action and voluntary 
death seems to be the reflex of some unknown stimulus ; and 
perhaps Ave may assume that the few insects set aside lived 
because their capture and removal to a fresh environment 
broke the influence of the stimulus. 

The two agarics (Podaxon pistillaris and P. carsino- 
malis) which grow out of the mounds of trinervius have 
nothing to do apparently with the economy of the termites 
(PI. XXXII, fig. 2). The mycelia from which these arise 
ramify throughout the earthen structure of the mound. 
Mycelial swellings form in cells just beneath the crust of the 
mound and the agaric breaks through the crust. It must be 
conceded that the mycelium subsists upon the organic matter 
in the matrix of the mound. It is to be remembered that 
there is a good deal of grass incorporated in the substance of 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. -ill 

the mound, and a thin layer of black, and no doubt organic, 
matter on the walls of the galleries, for the fungus to subsist 
upon ; this wall-deposit is more conspicuous when a red or 
grey coloured soil has been used to build the fabric. Although 
the termites will destroy the base of the agaric if the stalk is 
deep set in the mound, they never attack the superior portion ; 
indeed, in the case of the larger agaric, the termites con- 
struct what appear to be protecting clay collars around it 
when in the sprouting stage. Whilst several (one to five) 
agarics may grow upon one mound, it is rather extraordinary 
that only a small percentage of mounds have the fungus 
associated with them. 

The galleries of the mounds are throughout summer and 
winter choked to a great extent with short grass-lengths or 
hay. This is sometimes noticeably more abundant in the outer 
galleries of the nest, those under the crust, and the larger 
cells (or cellars) at the floor of the hive ; but, as a rule, the 
provisions are distributed fairly evenly throughout the whole 
nest. It is only in the spring of the year (November) that 
there is a marked depletion in the hay contents of the 
hive. 

According to Haviland's notes the workers cut the grass 
during the wet season (summer), working at night or in the 
early morning, and even in the middle of the day during dull 
weather. He further states that, " like Hodotermes, they 
stand head downwards ^ on a stem of grass, and with their 
mandibles gradually cut ai'ound the stem until it is sawn 
through." It has only been my good fortune to observe these 
insects foraging in the twilight, when their activities are 
difficult to follow. The workers seem to cut the grass blades 
thi'ough as best they can and stand in any attitude. They also 
gather green grass seed, and in the early evening this is piled 
on the surface of the ground ready for removal into the 
burrows before daylight. 

It has been recorded that the images reach maturity 
during September and October in Natal, issuing after night- 
^ Hodotermes does not necessarily adopt this attitude. 
VOL. 3, PAUT 2. 28 



412 CLAUDE FULLER. 

fall from the nest during November. This is also the case for 
the Transvaal. It has also been noticed that such swarming- 
takes place after rain has fallen, and that the full complement 
does not leave the nest at one and the same time. The 
crescent-shaped openings in the crusts of the mound through 
which the winged insects emerge, and the facts that these 
are closed up aiid subsequently persist for quite a long time 
afterwards, have also been repeatedly observed. 

When a nest is broken open suddenly numerous represen- 
tatives of all four castes are exposed ; but shortly such 
workers as can get out of sight do so. The soldiers, however, 
swarm over the broken surface, the smaller predominating, all 
lifting their heads aggressively and discharging viscid fluid. 
So eager are these little creatures that, where a perpendicular 
break is made, hundreds of them fall and form a mass at the 
base. After nightfall the breach is repaired by the workers, 
and in a little time the mound may be built up to its full 
propoi'tion again. When a calamity like this befalls the 
nest, the bulk of the workers attended by some soldiers, and, 
if present, all the nymphs, retreat into the radiating galleries ; 
hastening away from the zone of danger. 

Upon the whole this species is very immune from attack, 
and it is extremely rare to see nests much despoiled. In 
some, small holes are burrowed, 3 or 4 in. in diameter, most 
probably by meerkats ; but whether as a foe of the termite, 
or simply for the sake of making a lair, has not been deter- 
mined. Haviland^s notes state that jackals appear to feed 
largely on the species. This seems merely a conjecture. It 
has also been said that the aardvark destroys the nest, but if 
this is really so it is extraordinary that I have seen absolutely 
no evidence of it. 

The species to some extent escapes the attack of true ants. 
In certain parts of the Pretoria district where many nests of 
E. bilobatus are intermingled with those of trinervius, 
whilst every nest of the former species has one ants' nest 
associated with it, and presents ample evidence of the 
constant siege going on, nothing of the sort obtains with the- 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES, 41 

latter. If by chance an ants' nest is somewhere near to a 
trinervius mound and the mound is broken the ants will 
swarm around ; they do not, however, display that delirious 
blood-lust exhibited, under like circumstances, against bilo- 
batus, but behave with the greatest circumspection, never 
facing the soldier termite, but awaiting an opportunity to 
seize it from behind, in the region of the nape. 

As the viscid fluid ejected by the soldier of this species, 
considered as a missile of defence, can only be most 
ineffectual against a large marauder of the nest, it must 
be regarded as their chief form of defence against true 
ants. 

In the foregoing account it is assumed that each triner- 
vius colony inhabiting a mound is a separate colony, despite 
the fact that its mound may be connected by undergTound 
galleries with another mound or series of mounds. No experi- 
ments have been made to ascertain how far the insects of 
adjoining mounds are antagonistic or friendly, but Mr. Bag- 
shawe-Smith assui-es me that whenever he introduces a 
foreign queen to a mound the soldiers attack her, sliming her 
over with their viscid secretions. 



V. CLASSIFICATION. 

1. Introduction. 

This contribution is designed to fix the identity of the 
species whose habits have formed the subject of the preceding- 
discussion and that of several allied species. That some species 
are presented as new is incidental. 

For the purpose in view the soldier caste has received chief 
considei'ation, but worker characters, essential for their deter- 
mination or for the qualification of those of the soldiers, are 
also given. The images have not been described. Particular 
attention has been given to the full characters of the soldier 
mandibles and more especially to the basal regions which are 
oi'dinarily hidden by the labrum ; these charactei's ai-e as 



414 CLAUDE FULLER. 

important as those that are moi-e in evidence. The various 
antenna3 of the different adult castes have much specific vahie 
provided the limitations of the caste variation and their mode 
of growth are known. The colour effect of the chitin as 
overlying muscular tissue is given without attempting 
diffei'entiation j this seems to be the usual practice. Unless 
otherwise stated, they are taken from material pi'eserved 
in 75 per cent, alcohol. In certain species the colour tones 
and the colour patterns of the head are fairly constant, in other 
species they are not. Wliilst of assistance in diagnosis neither 
should be regarded as of great specific value. In general 
the length of the head with the mandibles and its greatest 
breadth are secondary aids to diagnosis, Avhilst the total length 
of the insect is a very deceptive measurement, as illustrated 
in the subsequent discussion of Term es natalensis. Here 
it may be mentioned that the various measurements given in 
the description are only to be regarded as approximate, 
whether given in mm. or ju. They have one value insomuch 
as all have been taken by the same means. Total length, 
head with mandibles and head-width have been determined 
by placing insects which have died in a more or less extended 
position npon a plane surface cross-ruled to half a millimetre 
and then reading off the length with a magnif3dng glass of 
low power. This method permits the taking of long series of 
approximate measurements rapidly ; by various checks I 
have found the results quite as reliable as measurements taken 
by more laborious and elaborate methods. Whilst the lengths 
of the soldiers are for fully extended insects, or those in which 
the long axis of the head is continuous with that of the body, 
the lengths of the workers are for insects with the head in 
the natural position at right angles to the long axis of the 
body. 

The lengths for the antennal segments are approximately 
the length of the chitinised*portion of the joint and are taken 
along the middle line of the joint. This omits the conjunctiva, 
and it does not give the full length of such joints as have a 
soft telescopic apex — all but two or three of the basal joints 



OUSERVATIOXS OX SOME SOUTH AFEICAN TERMITES. 415 

and the apical joint. However, it is a far more satisfactory 
method than endeavouring to trace elusive apical margins. 

Terms. — VV^ith regard to the descriptive terms these will 
be found to follow very closely those of Comstock and Kellog 
(5). The general terminology also follows these authors, 
Comstock and Chujiro Kochi (6) and J. B. Smith's ' Glossary 
of Entomology ' (7). Perhaps the only term used at variance 
with these authorities is " f rons/' which is given preference 
to " front.'' Not unnaturally some descriptive terms have 
been used in a restricted sense, a few terms have been 
borrowed from allied natural sciences, and some at least have 
been invented. To avoid any confusion a short glossary is 
given (see Appendix) so that it will be only necessary to 
indicate here the application of the terms '' frontal area," 
" frontal folds" and " ventral genee." 

" Frontal area" is an inclusive term embracing the areas of 
the frons and the clypeus in those cases where the Y-shaped 
suture of the epicranium and the clypeo-frontal suture are 
both obsolete. Thus "frons" is only used when the area is 
demarcated by a decided Y-suture or by some iutra-chiti- 
nous trace of it. 

The suture (clypeo-frontal) dividing the frons from the 
clypeus is present in workers, other than those of H o do- 
te rmes, and in all imagos ; but in all the soldiers it is to be 
regarded as obsolete, except in the case of Rlii note rmes 
putorius, where it is fully expressed. Its probable position 
is indicated to one degree or another in all soldiers by the 
features to which I have applied the term " frontal folds." 
These folds arise at the point of articulation of the mandible 
and usually extend obliquely backwards more or less into the 
median region of the frontal area. They are striking features 
in the soldiers and workers of Hodotermes, and in all cases 
it appears as if the lateral portions of the original suture were 
overgrown by a fold of the frons. The folds are actually the 
points of attachment of the props of the tentorium, and they 
are heavily marked owing to the thickness of the chitin, and 
a greater depth of colour occurs in the form of paired oblique 



416 CLAUDE FULLER. 

bands. To define these the term " frontal folds '' has been 
introduced. 

In certain species, especially in Hodotermes, the post- 
genfe are very large and profoundly differentiated from the 
geufe. In most other species, however, all trace of differen- 
tiation is either lost or but slight indications are present ; 
as in the transverse wrinkles exhibited by Termes swazife 
and the colour differentiation in Termes waterbergi. 
Owing to the flatness of the head it is very desirable to 
describe the ventral aspect, and, as an aid towards this end, 
the term " ventral-genae " has been improvised to indicate 
the whole of the cheeks lateral to the gula. 

The Head. — In the soldier caste it may be said that the 
head is a simple cylindrical capsule composed of two pieces — 
the epi-cranium and the gula — with a supporting tentorium. 
As a consequence, the regions to which the names of sclerites 
are apportioned must be regarded as indefinite and merging 
into those adjoining tlieni. 

The accompanying diagrams of heads of Hodotermes 
and Termes will suffice to illustrate the approximate areas 
to which sclerite names have been applied. As a matter of 
convenience, the term '' epistome " has been retained and 
applied in the restricted sense of a separate piece behind the 
labrum attached to the front margin of the clypeus. In the 
soldiers this is but a thin plate-like extension of the clypeus 
and is included as a part of the clypeus in the descriptions. 
In most of the workers discussed (and in the imagos) it is, 
hoAvever, a large and well-defined region calling for a special 
and particular definition. 

The labrum of the soldiers and workers of Hodotermes 
appears to be always simple. There are, however, many 
instances in which the labrum of common acceptance is in 
part the epipharynx. Thus in the soldiers of Termes 
swazias, waterbergi and natalensis the labrum termi- 
nates in a fleshy cordiform tip to which I have applied the 
term " lingula." This is simply a prolongation of the epi- 
pharynx. Again in Eu termes bilobatus, the labrum of 



Text-fig. 13. 




Termite head. A. Dorsal aspect of the head of Hodotermes 
soldier, b. Ventral view of the same. c. Dorsal aspect of the 
head of T er m e s soldier, d. Yentral view of the same. E. Dorsal 
aspect of the head of Termes worker. Ep. Epistome. 



418 CLAUDE FULLER. 

Avhich is said to be a forked sti'ucture, I can only think that 
the true labrum is a very atrophied sclerite, and that the 
so-called labrum is a chitinised extension of the epipharynx^ 
In nearly all the workers of the Termes species the labrum 
is distinctly bi-articulate and can only be considered as the 
labrum-epipharynx. In the Eutermes species there is a 
pseudo-articulation near to the apex of this sclerite which 
indicates that its structure is the same but simpler. 

Because it seems the proper thing to do, I have endeavoured 
to the best of my ability to describe the impressions, grooves,. 
keels, mounds, etc., of the frons. But it is a wearying matter 
in most cases, as, with every shifting illumination and every 
alteration of the point of vision, the aspect changes. Hence 
my introduction of " kaleidoscopic " into some of the 
descriptions of the frons and my appreciation of HavilandV 
remark : " I have not attempted to outdo Nature in distinct- 
ness; indeed, in this respect I am conscious of shortcomings." 

Upon the whole the frons when impressed and embossed 
is very variable in the degree of its sculpturing, and the 
details are so minute, so variable, so indescribable and of sO' 
little specific value that no more than a loose reference to the 
nature of this area is really necessary. Indeed, I have found 
by experiment on my colleagues that no two of us could 
agree upon the exact nature of the frons-sculpture of either 
Termes natalensis or T. badius. 

Legs. — In certain species the tibia of each leg appears to 
be two-segmented; the differentiation being more or less 
distinct in different species. This arachnoid feature may 
possibly be looked upon as indicating a primitive character. 

Antennae. — The growth of the termite antenna after the 
hatching of the young termite may be said to be due to the 
production of a series of intercalated joints in a vegetative 
manner. This development has not been actually traced, but 
it may be deduced with some confidence. If, for example, 
T. badius is examined it is found that the imagos have 
antennge of 19 joints whilst the adults of other castes may 
have less; the major workers presenting 19, 18, and 17 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 419' 



segments; major soldiers 17 and 16 segments, and the- 
minor soldiers 17 segments. The XlX-jointed worker is rare,. 
and the XYIIl-jointed is more common than the XVII-jointed 
form, a circumstance which, in combination with other 
features, illustrates that the worker-caste is due to arrested 
development. 

Text-fig. 14. 




IV I V I VI I VII J VIII I IX 
Imaffo •. X\X Jointed 




3*4 



Worker X\X Jointlo 
5 6 7 S 9 10 

IV I V 1 VI I VII 




^' Worker XVIII Jointed. 
i 
5+-6 




^^y' Worker &< Soldier : XVII Jointlo. 
5M^5-^6 7 8 9 10 II 

IV 




Soldier :)WI JoiNTtO. 



Diagrams of different antenna; of various castes of Termes 
badius to illustrate liowtlie variations exhibited arise through 
the moi'e ov less complete failure to separate of the segments 
immediately distal to joint II. 



If the several forms of the worker antennte are compared^ 
it is seen that the rare XlX-jointed form is very similar in its 
pattern to the adult imago form ; the chief difference being 
in the generally smaller size of the joints. This is illustrated 
in the antennal curves to which reference will presently be 
made. It is further illustrated in the accompanying diagrams. 
Starting from the XlX-jointed form it is not difficult to show 
how the XVIII-jointed form arises out of fusion (or, more 



420 CLAUDE FULLER. 

correctly speaking, the failure to separate) of joints III and 
IV ; because, if a series is searched for, every degree of non- 
separation can be found from joints with two whorls of bristles 
and distinct pseudo-articulations through joints with two 
whorls of bristles and no trace of articulation to simple uni- 
whorled segments giving no direct evidence of how they 
orig'inated. That the XVII-iointed antenna is formed bv 
the non-separation of joints V and VI of the XlX-jointed 
form can be demonsti^ated just as readily. 

Turning to the soldier caste ample evidence can be found 
to show that the XVII-jointed form originates as does the 
XVI I- jointed major and minor worker form ; that is, by 
finding pseudo-articulations and double whorls of bristles on 
joint IV (or 5 4- 6 of the XlX-jointed antenna). In certain 
nest-series there are soldiers which present XVI segments to 
the antenna and all joints are complete and uni-whorled. 
However, amongst them can be found (a) normal soldiers 
with XVII-jointed antenna, {h) sometimes an abnormal 
soldier with both a XVI- and a XVII-jointed antenna, und 
(c) an abnormal soldier with both antennJB XVI-jointed, in 
some of which III shows an intermediate ai'ticulation ; in 
other words the joint III of the XVI-jointed antenna has 
arisen through the non-separation of joints III, IV, V, and 
VI of the XlX-jointed form. 

In this connection the segmentation can be deduced from 
the larval forms of T. badius. The youngest condition 
observed is the larval stage (1 to I'S mm. long) having ten 
joints clearly expressed (Text-fig. 15, a). Joints I and II have 
already acquired their fixed expression, and seven apical joints 
are distinctly separate and joint III is elongate. Joint III 
-then subdivides into three segments and gives a XII- 
joitited antenna {h). It then elongates (c) and later subdivides 
to give the XlV-jointed antenna (cZ). This shows that the 
XlX-jointed antenna arises out of the growth and sub-division 
of joint III. If there is no check in development, the extra 
joints appear between III and IV of the X-jointed antenna 
until the XlX-jointed form is fully expressed, and this, at 



OBSEKVATIONS ON 80ME SOUTH AFKICAN TERMITES. 421 

one stage, involves the separation of lit into two instead of 
three segments. Consequently the final condition of III 
varies according to whether the final IV is completely 
articulated or not. If no complete fissure occurs the XA^I- 
jointed antenna results. On occasion, too, the most recently 
formed segments, i.e. Y and VI, appear to fuse instead of 
becoming definitely separated, and this circumstance, together 
Avith the compound nature of III, results in the XVII-jointed 

Text-fig. 15. 




T. badiiis; auteiina; of larva?, a. X-joiiited expression with a 
loug joint III. b. The X-jointed changed to a Xll-jointed 
expression by the subdivision of III. c. The Xll-jointed 
expression prior to the subdivision of the long joint III to give 
the XlV-jointed expression, d. The XlV-jointed expression 
before the enlargement of III. 

form. The XVI-segniented type is obtained by an arrest of 
development which must be considered to involve III as 
comprising what would really have been III, IV, V, and VI 
of the XlX-jointed type. 

In short, the XlX-jointed antenna is produced from a form 
with fourteen segments by a process of proliferation or fission 
in joint III. 

Apart from this process other factors seem to come into 
play to influence the form ot certain joints. In badius 
joint V is a relatively small joint in the XlX-jointed antenna 



IujI LIBRARY 






422 CLAUDE FULLER. 

of both the imago and worker castes ; and, as joint IV of the 
XVIII-jointed form, it is also small. In the XVII-joiuted 
form the fifth joint is practically the smallest of the series ; 
and, as this repi-esents the relative large joint VII of the 
XlX-jointed form, it gives evidence that the shape and size 
of the joint is influenced by the distance it is removed from 
the head. 

A somewhat similar state of affairs prevails in the antennal 
pattern of T. natal en sis, where in the XlX-jointed imago 
antenna V is narrower and smaller than IV and VI. It is 
reproduced as joint IV in the XVIII-jointed ; but, in the 
XVII-jointed expression, it is again V that is smaller than IV 
or VI. In the XVIII-jointed antennae. III and IV are small 
and about equal, whilst Vis always larger than VI ; froni this it 
is inferred that the XVIII-jointed worker antenna is due to the 
failure of V and A^I of the full expression of XIX to separate, 
and that the XVII-jointed form of both major and minor 
soldier is due to the fusion of III with IV of the XVIII-jointed 
form ; by this means the small joint VII of the imago becomes 
the small joint V of the soldier. A pseudo XVII-jointed 
worker-major antenna is due to the fusion of the joints V 
and VI of the XVIII-jointed form. Upon the other hand, 
the XVII-jointed antenna of the worker-minor is not due to 
the failure of joints to separate after being produced, but 
rather due to the fact that joints have not been produced. 

As it was thought that a series of antennal formulte, such 
as have been used by workers with coccids, might be useful 
in fixing species, a number of exact measurements of many 
antennae of T. natalensis and T. badius were made. 
Although of a constant general pattern, the variability in the 
actual length of the joints was such that it Avas rare that 
more than two formulae in ten coincided. My colleague, 
Mr. C. K. Brain, of the Union Division of Entomology, having 
met with similar difficulties with the formulee for coccid 
antennas, had some time ago devised a means of reflecting the 
antennal patterns by charts. This he has improved to show 
the range in the lengths of the individual segments. By 



OBSEBV^ATIONS ON SOME SOUTH AFRICAN TERMITES. 423 

applying- his method to the antennee of termites some very 
satisfactory results promise to follow, as is indicated in the 
provisional antennal charts now submitted for T. natal- 
ensis, T. badius, and Eutermes trinervius (PI. XXXIII). 
I do not anticipate equally satisfactory results from its 
application to the genus Hodotermes, although I do for 
most genera. 

Mr. Brain has been kind enough to permit me to include 
here a note by him on his use of antennal charts. 

"The most useful arrangement of antennal data seems to 
be ari'ived at by giving the range of variation in measurements 
of the different segments with the addition perhaps of the 
mode of each. After working over a large series of slides 
one is impressed with the characteristic appearance of different 
coccid antennse ; but this diiference is difficult to express. 
The nearest approach is obtained by charting a polygon 
showing both the mode and the range of variation in the 
segment lengths. This supplies a most useful aid for the 
preliminary location of coccids from slide specimens. 

" Whenever possible, the range of measurement should 
represent at least ten measurements, twenty to thirty being- 
made where material is sufficiently plentiful to allow such to 
be done. The measurements are then plotted in the centre 
of each antennal column and the polygon completed. Thus 
it appears that where any segment is constant the charted 
area approaches a simple line, while a wide band represents 
a wide range of variation. 

" Similarity of antennal chart, whilst indicating- similarity 
of antennal formula, does not of necessity indicate identity 
of coccid species; but it does give a clue to work upon, and 
possibly at times indicates relationship." 

2. Systematic Account. 

hodotebmes. 
I have ventured to split into four species a certain group 
of specimens in the collection which, in part, has been identified 



424 CLAUDE FULLER. 

as Hodotermes mossambicus Hagen. In this I think I 
am right, although it must be admitted that the characters 
are ehisive. I have considered it safer to regard them, for 
the time being, as different species, because the terms sub- 
species, variety, and race are either quite opinionative, and 
therefore involve the personal equation, or they imply a 
biological knowledge, and are only correctly used Avhen 
founded upon a comprehensive study of an extensive series 
with full details of range, local conditions, and life-history. 
In the case of each, however, I have been able to examine one 
or more good series from separate colonies and localities, and 
have found that, when compared side by side, the soldier caste 
of each would seem to represent different species. 

The insect which I have specified as H. mossambicus is 
that to which the descriptions of Hagen^s species, which are 
available to me, apply best. Further, it is that which, in 
the collections of the Division of Entomology and the Natal 
Museum, has been obtained from localities approximating the 
range of mossambicus. It is represented by material from 
the moist coast region of Zululand and the dry table-lands 
of the interior (Kimbei'ley, Marico). In dealing with the 
series the description of the soldier of H. transvaalensis 
sp. n. is given as a standard for comparison. Photo-micro- 
graphs (PI. XXXY) are given of the labrum of each species 
recorded, and, in order to fix the elusive variations of man- 
dibles of similar dentation, a series of camera-lucida outlines 
of these is also furnished (PI. XXXIV). This series shows 
the mandibles selected from the largest soldiers equally 
niao"nified. In the illustrations it is seen that the different 
sets have been variously orientated, but all are arranged 
so that the basal processes are level. Fui-ther to this the 
appearance is largely due to the fact that mandibles are 
articulated in different species in slightly different attitudes. 

I have not regarded any of the Hodotermes as having 
major and minor castes, either among the soldiers or workers, 
because in any extensive series of South African forms all 
o-rades are to be met with between two extremes — the smallest 



OBSERVATIONS ON" SOME SOUTH AFRICAN TERMITES. 425 

and the largest. In the species specified as Termes badius 
Hav. the same course has been followed with the soldier caste 
for similar reasons; but badius does possess well-defined 
major and minor workers. 

The worker caste of sevei'al species has been described in 
detail, but these descriptions, as far as they go at present, 
can be applied for all pi-actical purposes to the largest 
workers of any species except viator and, perhaps, kar- 
rooensis. Certain peculiai-ly obese and bright yellow, 
worker-like insects, taken foraging in company with adult 
and callow workers of H. karroo ens is s^j. n. and H. 
warreni 6'j9. ii., have not been described because it has not 
been possible to decide whether these are aberrant workers 
or juvenile images. 



Hodotermes transvaalensis sp. n. PL XXXIV, figs. 
1, la; PI. XXXY, fig. 1. 

SOLUIEE. 

Measurements.^ — Total length 15 to 20 mm.; head with 
mandibles 6 to 8 mm., head-width 4*5 to 5"5 mm., head-length 
4' 5 to 5' 2 mm. 

Head. — Vertex pale brown, or reddish- or yellowish- 
brown ; f rons conspicuously mottled with light whitish yellow ; 
of this colour three triangular pennant stripes extend back- 
wards, their narrow wavy apices reaching well over the 
vertex ; clypeus wholly dark, or dark red-brown or purple, 
with two faint patches of paler hue. Surface polished ; vertex 
quite smooth : frons wrinkled ; clypeus strongly chitinised ; 
the whole with short scattered hairs, those on the frons being 
the stronger. Dorsal outline broadly U-shaped, not wider 
behind than in front; Y-suture distinct ; vertex flatly ax'ched ; 

^ The figures given in this and the following descriptions are with a 
few exceptions based on the measurement of at least ten insects, but 
in some cases many more were measured. The range is not that of a 
nest series, but represents the difference between the largest and smallest 
of the measured insects. 



426 CLAUDE FULLER. 

sides convex, parallel and straight ; genas faintly concave ; 
trochantin of mandible strong and prominent. 

Frons deflected and twice depressed ; depressions separated 
by a sliort, distinct, transverse ridge which forks at both 
ends, the caudal prongs merging into the vertex and the 
anterior prongs passing within the ocelli and terminating at 
the mandible; caudal depression shallowing into vertex; 
cephalic merging into clypeus and with a low pale-coloured 
median mound which appears triangular and tapers back- 
wards, the apex merging into the ridge separating the two 
depressions. 

Eyes black, sub-renifoi-m, facing more outward than 
forward. 

Ocelli distinct, white, surrounded by a reddish aureole ; 
viewed from the side they appear inset in the top of low 
mound. 

Clypeus actually oblong but with corners so obliquely 
deflected as to give the cephalic margin (the epistome) a 
three-sided appearance, swollen behind cephalic margin so 
as to form a transverse, rounded ridge ; a distinct furrow or 
wrinkle connecting frontal folds (= the clypeo-frontal 
suture) . 

Labrum pale brownish-yellow ; somewhat polished ; with 
yellow bristles ; short and broad ; arched and very broadly 
cordiform ; distal mai'gin flatly brace-shaped or roundly 
acute. 

Mandibles black ; dorsal mounds very dark reddish-brown ; 
broad and flattened ; with wide bases and teeth ; lateral 
margins shouldered near the base and then broadly curvate ; 
the doi-sal surface of each with an elevated ridge which is 
hiirh at the basal knob and extends forward to the second 
tooth as a flattened punctate mound having a rounded 
margin. Apical points broad, much incurved and deflected ; 
the inner margin sinuate, at first incurvate then curvate and 
tapering into acute incision above the first tooth ; the left 
point longer than the right, but not quite as wide and more 
-deflected. The left mandible longer and wider than the 



OBSERVATIONS ON SOME SOUTH AFRIOAN TERMITES. 427 

right ; with thi-ee teeth, of which the first is acute and the 
largest, the third the smallest with a faintly sinuate, tri- 
curvate edge; also with a strong basal process, below which 
the margin tapers and is incurvate and then curvate. The 
right jaw with two teeth, both of which are larger than tlie 
first two of the left jaw, the cutting edge of the second long 
and wavy; margin below the second tooth with three incisions 
and two well-defined scallops ; margin extending obliquely 
outwards. 

Antennae pale yellow-brown throughout; whip-like; joints 
I and II forming the handle, the rest a fusiform thong ; 
joint I inset below the hood of the froutal carina ; of about 
XXVIII, XXIX, or XXX joints ; III, IV, and V usually sub- 
equal and the narrowest and shortest of the series ; VI to 
apical increasing and then decreasing in length ; from VII 
or VIII to the fourth to sixth apical joint the width is 
generally constant, the apical series tapering slightly ; from 
VI the lengths gradually increase to a maximum which may 
be reached between XIII and XVI ; apical joint always 
shorter than the penultimate and often decidedly narrower ; 
maximum length of longest joints about equal to that 
of II. 

I, strong, cylindrical, apex with a distinct whorl of bristles ; 
more than twice the length of II (range 400 to 500 fx) . 

II, strong, cylindrical, nearly as wide as long, wider than 
any succeeding joint ; with an apical whoi'l of bristles and 
scattered bristles on apical half (range 200 to 300 /u) . 

III, IV, V annular; less than half the length of II; III 
anchylosed to II ; each with an apical whorl of bristles 
(range 70 to 100 /z). 

VI annular ; longer than V ; with an apical whorl of 
bristles (range 100 to 130 /x). 

VII longer than VI (100 to 150^) ; often with two Avhorls 
of bristles. 

VIII and onwards, the first few joints globose, the middle 
series obconic, the apical series elongated with almost parallel 
sides and two whorls of bristles. 

VOL. 3, PART 2. 29 



428 CLAUDE PULLER. 

Apical with convex-truncate apex and with two whorls of 
bristles. 

Post-genee differentiated from the genae by a sharp carina ; 
sloping down to the gula ; transversely and regularly rugose. 

Gula scoop-shaped ; free blade broad, with sides bent up 
and parallel, distal margin curvate ; inserted stem much 
constricted behind blade, and widening gradually into blade; 
one third as long as blade. 

Thorax. — Median area of each shield bright yellow, 
broadly and sharply bordered with smoky brown ; meso- 
and metanotum with paired and very distinct dark brown 
dots. 

Pronotum sellate ; narrower than the head ; partitioned 
before half its length by an open transverse furrow which 
debouches on either side into cup-like hollows ; quartered by 
a faint median line; cephalic area arched, but with its 
median field hardly raised above level of caudal field ; 
cephalic margin arched, arch rising almost perpendicularly 
from the lateral processes and bending in a flat curve to its 
apex where there is a shallow and small notch ; caudal area 
with the median field arched, the side-plates sloping gently 
outwards; sides curvate, tapering slightly, margins not 
upbent; lateral processes salient, spout-like (like gargoyles), 
large, projecting outwai-ds and forwards as well as pointing 
downwards, incised at base before and behind, much hollowed, 
hollows shallowing into transverse furrow ; caudal margin 
broadly incurvate, almost straight. 

Mesonotum with median field convex ; sides broadly 
curvate and with edges a little bent up iu front ; caudal 
margin faintly, bilobed, nearly straight; as broad as pro- 
notum less the processes. 

Metanotum a little narrower than that of mesonotum ; 
arched ; sides curvate, diverging sufficiently to form distinct 
caudo-lateral angles with hind margin, edges as with meso- 
notum ; caudal margin bilobed, median portion prolonged, 
laterad of lobes very incurvate. 

Legs brown ; apex of tibia of I with 2 to 4 spurs, of II 



•OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 429 

with 3 to b, of III with 2 to 4 ; in any insect there is always 
a greater number on II than on I and III. The tarsus of 
III is the longest, of II a little longer than that of I, the 
ratio is 7 : 5 : 4; in all, joint IV has a length which is con- 
stantly equal to the length of 1 + 2 + 3. 

Abdomen. — Pale brown. Styli with rather long but not 
numerous bristles. 

This soldier is closely allied to that of pretoriensis, but 
diifers chiefly (a) in its comparatively broader mandibles, (6) 
its less acute labrum, (c) in the basal scallops of the right 
mandible, (d) in the styli both species are entirely diiferent 
from karroo en sis. 

Worker. 

Measurements. — Total length 7"5 to 12 mm.; head- 
width 2'5 to 4 mm. 

There is no great difference between the worker of this 
species and that of pretoriensis except that it seems to 
show a lower length range, and the yellow spot is not so 
prominent on the frons. 

H a b. — Transvaal ; Pretoria. 



Hodotermes pretoriensis sp.n. PI. XXXIV, figs. 2, 2a; 
PI. XXXV, fig. 2. 

Soldier. 

Measurements. — Total length 16 to 20'2 mm.; head 
with mandibles, 7 to 8 mm. ; head-width 4*7 to 5'5 mm. 

Head. — Vertex pale brownish-yellow; frons a little more 
yellow, but not conspicuously so, with short pennant stripes 
running a little backwards over the vertex, and with two 
conspicuous copper-brown bands extending inwards obliquely 
from the bases of the mandibles and indicating the frontal 
folds ; clypeus reddish or pale brown-yellow, except that the 
lateral processes behind mandibles ai'e black. Surface dulled; 
vertex sparsely punctate ; frons finely wrinkled ; whole with 
conspicuous scattered hairs, which appear stouter on the 



430 CLAUDE FULLER. 

frons. Dorsal outline broadly U-shaped; almost as long as 
wide, not broader behind than in front; Y-sutvire distinct; 
vertex roundly arched, more arched than transvaalensis, 
and occipital region inflated ; sides nearly sti-aight, slightly 
curvate; trochantin of mandible strong and prominent. 

Frons deflected ; without transverse ridge or brow ; almost 
flat with paired median impressions between ocelli; merging 
into depressed region of clypens; frontal folds very distinct. 

Eyes black, sub-reniform, facing more forwards than out- 
wards. 

Ocelli represented by distinct, irregular-fusiform pits. 

Clypeus with epistome : margin truncate, sides more or less 
obliquely rounded. 

Labrum broad, longer, and with a more acute apex than 
in transvaalensis, and longer and a little less tapering 
sides. 

Mandibles light i-ed with teeth and apical points black. 
Dentation and form very similar to that in transvaalensis 
but more highly polished ; longer and narrower ; margin 
below second tooth of left with two incisions and one distinct 
scallop. Apical point and distal tooth of both mandibles 
sometimes duplicated. 

Antenna yellow; whip-like; as with transvaalensis; 
joints III to V very variable, sometimes III, IV, and Y equal 
and small, sometimes III larger than TV, or IV than III, in 
which case the larger joint usually presents a pseudo-articula- 
tion and two whorls of bristles ; number of joints ranging 
from about twenty- seven to thirty. 

Post-genee and gula much as with transvaalensis, but 
inserted stem of gula longer and widening more sharply 
into blade ; blade more rectangular with sides faintlj^ in- 
curvate. 

Thorax. — Almost uniformly dull yellow ochre, with quite 
inconspicuous paired dots on shields of meso- and metanotum. 
Agrees in structural details with transvaalensis, except 
(1) the lateral processes do not project so much, and, as a 
consequence, the mesonotum appears as wide as the pronotum ; 



OBSERVATIUNS ON SOJJE SOUTH AFEICAN TEKMITES. 431 

{2) there is only one basal scallop on tlie right mandible. 
Much like that of mossambicus, except that (1) the cephalic 
margin of the pronotum is distinctly notched, (2) the caudal 
margins of all three shields are more retuse, (3) the basal 
scallop of mossambicus is not curvate. 

Legs pale brown ; in the majority of cases I with 3, II 
with 4, and III with 3 spurs; occasionally I with 2, 11 with 
5 to 6, and III with 4. 

Abdomen.— Whitish brown, sordid, paler than that of 
111 ossambicus. 

Worker. 

Measurements. — Head-width 2'7 to 4'2 mm. ; totallength 
8 '5 to 13 mm. 

Head. — Vertex a dark choculate-bruwn or paler, dark 
colour extending forward over median area of frons; mediad 
and laterad of antennal fossae pale sordid brown ; incised 
portion of clypeo-frontal suture indicated by well-defined 
dark brown bands ; epistome translucent ; clypeus usually 
pale or basal area darker, when pale the ridge brown; labrum 
translucent^ a sordid yellow brown ; trochantin of mandible 
pallid ; there is always a small white median dot in the apex 
of a pale V line in the caudal darkened area of the frons. 
Dorsal outline open (J-shaped; cephalic extremity broadened; 
sides convex, tapering sharply and merging into caudal margin, 
not bending in mai'kedly to trochantin of mandible ; Y-suture 
distinct ; vertex flatly arched. 

Frons with cephalic area very flatly depressed, with a very 
faint median carina in depression. 

Eyes black, sub-reniform. 

Ocelli quite obsolete, represented by faint shallow punc- 
tures. 

Clypeus with median length of clypeo-frontal suture 
obsolete; with a ti'ansverse cephalic ridge; epistome plate- 
like, broad, sides tapering; cephalic margin straight with a 
small median lobe. 

Antenna? pale or dark brown ; apical joints tapering. 



432 CLAUDE FULLER. 

Joints III, IV, V sub-equal in the lai'gest insects ; IV the 
largest ; in the smallest, III, IV, V, VI sub-equal. 

Thorax. — Constricted; each shield with a well-defined 
dark brown area bordered with a sordid white. Pronotum 
with cephalic margin white ; arched, with a small median 
notch. 

Mesonotum longer and narrower than metanotum ; both 
with faintly curvate, almost straight, diverging sides, then 
rounded and tapering incurvately to narrow straight caudal 
margins. 

Meta- more prolonged than mesonotum. 

Legs with tibia and tarsus pallid, rest brown. 

Abdomen. — Convex above; dorsal sclerites with dark 
chocolate-brown bands which diffuse to pale brown towards 
both margins; anal appendages long, tipped with brown; 
cerci translucent. 

Hab. — Transvaal; Pretoria. 

This species is closely allied to transvaalensis, and has 
similar habits, and occurs in one and the same locality. The 
most striking differences are mostly those of colour, and the 
thorax of transvaalensis is ornate in comparison with the 
dull mono-tint of that of pretoriensis. The styli of the two 
insects are essentially different, those of pretoriensis being- 
more bristly — the proportion being apparently four bristles 
to every three of transvaalensis. The styli of the workers 
differ in the same respect, but the difference is not so striking 
and can only be seen when a series of these organs from the 
two species are microscopically compared. On two occasions 
it has been taken mining in the plaster of inner walls of 
houses in Pretoria, and is reported to have destroyed books and 
starched lace curtains. The more serious complaint against 
it was the noises made by the rappings of the soldiers at night. 
In one series the majority of the soldiers have the apical point 
and distal tooth duplicated, so that, when viewed from the 
inner side, the apical points and the teeth below them are 
furcate. 



OBSERVATIONS ON SOME SOUTH AFRICAN TEEMITKS. 433 



Hoclotermes karrooensis sj). n. PL XXXIV, figs. 3, 3a; 
PL XXXV, fig. 3. 

Soldier, 

Measm-ements. — Total length 15 to 19 mm.; head with 
mandibles 7'5 to 8'5 mm. ; head- width 5"2 to o"5 mm. 

Head. — Vertex bright yellow or reddish-yellow, broadly 
striped with diffuse copper-brown ; frons pale yellow-brown 
with a conspicuous yellow triangular patch in the foi'k of 
the Y-suture ; mediad of antennal fossse large diffuse patches 
of yellow, and a triangular yellow patch caudad of base 
of clypeus ; clypeus copper-brown. Surface polished ; vertex 
and frons equally faintly wrinkled and equally clothed with 
very scattered short hairs. Dorsal outline broadly oval and 
noticeably wider than long; Y-suture distinct; sides broadly 
curvate, in part almost straight and parallel ; trochantin of 
mandible well expressed. 

Frons defiected and twice depressed ; depressions not 
separated by a distinct ridge; caudal depression very faint, 
level, merging into deflected cephalic depression ; presenting 
two pits, and between them a median elevation which has the 
form of a low triangular pyramid. 

Eyes black, sub-reniform, facing obliquely upwards. 
Ocelli obsolete, indicated by shallow depressions centring 
a small diffuse reddish spot. 

Clypeus oblong; cephalic margin (epistome) faintly incnr- 
vate, thin and deflected anterior to a distinct transverse 
ridge ; clypeo-frontal suture indicated by wrinkles connecting 
frontal folds. 

Labrura bearing a strong resemblance to that of p re- 
tor ien sis, but less acute, and sides less tapering ; more acute 
than in transvaalensis. 

Mandibles reddish-yellow, apical points and teeth black ; 
much coarser than in transvaalensis or pretoriensis ; 
lateral margins irregularly curvate (wavy), more incurved 
than in either transvaalensis or pretoriensis, especially 



434 



CLAUDE FULLER. 



the apical point of right; dentation as with transvaalensis, 
but teeth more separated ; basal margin of process of left 
acute ; margin below second tooth of right with two shallow 
obtuse incisions and no distinct scallop between them. 

Antennae yellow; whip-like; much as with trans vaal- 
ensisj but joint III usually the smallest, with IV larger 
than III, V larger than IV, and VI larger than V ; sometimes 
with IV and V equal ; sometimes with III larger than IV, and 
when so presenting a double whorl of bristles ; number of 
joints about twenty-eight to thirty-one. 

Post-gense concolorous with genae or bi-ighter yellow. 

Gula scoop-like ; edges of free blade tapering towards the 
apex; distal margin faintly incurvate; inserted stem short 
and wide. 

Thorax. — Median area of each shield bright golden 
yellow bordered by pale brown ; with paired and distinct 
pale brown dots on meso- and metanotum ; agreeing in struc- 
tural details with transvaalensis, except that (1) the 
segments are proportionately broader, (2) the processes not 
so incised behind, and (3) the metanotum is as wide or wider 
than the mesonotum. Much like that of mossambicus, 
except that (1) the shields are relatively broader, (2) the 
pronotum presents a faint notch on the cephalic margin, and 
the lateral processes project more outwards, (3) the mesonotum 
is more angled behind. 

Variation in Number of Spurs. 



Specimen. 


1 


2 


3 


Tibia. 


I 


II 


III 


I 


II 


III 


I 


II 


III 


Ou the right 


4 


4 


* 


3 


1 


* 


2 


4 


4 


On the left 


2 


y 


3 


2 


4 


2 


2 


4 


2 



* Spurs broken ofE. 



Legs yellow brown ; in the majority of insects tibia I with 
3, II with 4, and III with 3 strong spurs. 
Abdomen . — Bright golden yellow. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 435 

WOKKER. 

Measureineiits. — Head-widths 2*5 to 4*5 nnn. ; total 
length 9*5 to 14 mm. 

Head. — Vertex intensely black, colour extending over 
median frontal area towards base of clypeus ; frontal folds 
<3hestnut brown ; yellow patches median and lateral to 
antenna] fossa3 on cephalic margin of head ; clypeus with 
epistome ti'anslucent, with a broad dark median stripe 
bordered with white patches; labrum yellowish; trochantin 
of mandible white. Dorsal outline open U-sh^ped ; cephalic 
extremity broadened ; sides convex, slightly curvate, tapering, 
tapering more decidedly in small workers than large, bending 
in to trochantin of mandible ; Y-suture distinct, especially 
in small Avorkers where it is pale in colour; vertex flatly 
arched. 

Fronswith cephalic area presenting a small median mound 
and lateral to this two depressions ; separated from clypeus 
by a groove connecting two lateral incisions (an incomplete 
clypeus-frontal suture). 

Ocelli represented by pale spots. 

Eyes black and sub-reniform. 

Clypeus large, broad ; epistome plate-like and prolonged 
with a narrow straight margin and incurvate oblique sides; 
apex of clypeus elevated and forming a distinct transverse 
ridge reaching almost from side to side. 

Labrum large ; cephalic margin broadly rounded; anterior 
lateral corners lobed ; sides incurvate and tapering. 

Mandibles reddish-yellow or chestnut with black points. 

Antennae a pale sordid brown; HI, lA^, V annular, sub- 
equal, IV generally the largest of the three in the largest 
workers and the smallest in the smallest workers. 

Thorax. — Constricted, nari'ower than head; dark black- 
brown, except that the cephalic margin of the pronotum is a 
paler sordid brown; pro-, meso- and metanotum with a pale 
■median line. 

Pronotum with cephalic margin rounded, entire, arched, 



436 CLAUDE FULLER. 

and somewhat prolonged over head; processes deflected and 
promment; caudal margin straight, sides curvate, tapering 
acutely. 

Mesonotum long and broad ; sides diverging ; caudal 
margin broadly curvate or faintly bilobed. 

Metanotum shorter and wider than mesonotum ; caudal 
margin with three edges, laterals straight and converging, 
median faintly incurvate. 

Legs with coxa dark brown, femur pale brown, tibia and 
tarsus pallid. 

Abdomen. — Broad and flat; colour always pale; dorsal 
sclerites pale, sordid brown ; colour not contrasting sharply 
with conjunctiva ; ventral plates a sordid white ; styli long 
white and black tipped ; cerci pallid. This species can always 
be separated from the others by the characteristic colour of 
the worker abdomen. 

Hab. — Cape Colony; Middelburg, Steynsburg, Victoria 
East (The Karroo). 

The habits of this species agree very well with those of H. 
trans vaalensi s. Field collections made in August, 1913, 
contained many large bright yellow insects. These were 
quite as large and as brightly coloured as the soldiers and 
somewhat obese. They have the worker form, except that 
the heads are less flattened, and they forage with the workers 
and immature images. The images which I consider to be of 
this species are of a bright yellow. These were collected on 
the same date with soldiers and in the same place as this 
species abounds in. This species is known to be very 
destructive to growing crops and to tunnel through the clay 
plaster of inside walls. 

It was regarding this species that Mr. Albert Rubidge, 
farm Emerald, Middelburg, wrote to me the accompanj^ing 
letter under date October 3rd, 1912. As no large species of 
the genus Termes have been noticed at Middelburg, it is 
extremely probable that Mr. Rubidge^s reminiscences refer to 
a Hodotermes. 



OBSEKVATIONIS ON SOME SOUTH AFRICAN TERMITES. 437 

■' I note that you say the iiest of the marchiug termites has not been 
discovered. The following may, therefore, be of interest to you. As 
far back as forty years ago, a couple of Bushmen arrived at my father's 
place in Graaff Reinet and asked to be employed in taking out the 
nests of the " rijsmier.' As these little insects were then troublesome 
in a cottage my father set them to work near it. As a child I Avas 
much interested and watched the process ; it consisted of putting a 
wheat straw (or several in different passages in a small area) down the 
passage from the surface, then excavating to near the end of the straw ; 
then again inserting the straw as far as it would go ; and so on. As 
the passages frequently have short curves it is easy to lose the run of 
them. Thus these Bushmen dug several large holes to the depth of ti 
to 8 ft. without success ; but ultimately they succeeded in reaching 
the nest at about 8 ft. down. This consisted of a circular hole or 
well from 18 to 24 in. in depth by about 10 to 12 in. in diameter 
— beautifully smoothed and cemented. From this was taken about a 
bucket to a bucket and a half full of insects together with the queen — 
about 2^ in. in length and about If in. to 2 in. in circumference, in 
appearance very similar to your illustrations (3) and of a dirty white 
colour. 

" The entrances to the passages were to he found over an area of 
about 50 yards square, and the actual nest was found about 15 
yards away from the building. 

"The unsuccessful holes dug by the Bushmen were 8 to lU yards 
from the successful one. You wull note that I aai not very definite in 
my figures, but must bear in mind that I am speaking from impressions 
on the mind of a child forty years back. 

" These termites were very ti'oiiblesome in my house at one time, so 
I mixed some white arsenic with bran, filled the round of a magnum 
Ijonum pen, and blew the mixture into the entrances to their passages. 
This proved efi'ectual ; as they eat their dead the poisoning continued, 
and they came out to die, so we removed them by plates full (as they 
died in the old-fashioned wall cupboards)."' 

(As Hodoterines discai-d their dead, it is more probable 
that those which died were thrown out by the insects from 
the outlets in the cupboards. — C.F.) 

In a further letter Mr. Rubidge writes : 

'• The damage done in my house referred to was not to the wood work, 
but rather to the plaster and paper. They tunnelled along under the 
paj)er in all directions and ate the paper from the walls. Moreover they 
destroyed books and other papers, and carved out extra patterns in the 
window-curtains, even eating linen clothing." 



438 CLAUDE FULLER. 

Hodotermes mossambicus {Hageii). PI. XXXIV, figs. 4, 
4a ■ PL XXXV, fig. 4. 

Termes mossambicus Hagen. Monatl. Bericht. d. K. Akad. der 
Wiss, Berlin, p. 480, 1853. 

Soldier. 

Measurements. — Total length, 14 to 15"5 mm.; head 
with mandibles, 5'7 to 6"5 mm. ; head-width, 4 to 4*5 mm. ; 
head-length, 4 to 4*5 mm. 

Head. — Vertex reddish-yellow; clypeus reddish, darker 
in the middle; frons yellow, mottled as with transvaalensis, 
but not so conspicuously. Surface polished; vertex not 
quite smooth; with few and inconspicuous scattered hairs; 
frons faintly wrinkled ; clypeus not strongly chitinised. 
Dorsal outline broadl}^ or roundly oval, a little longer than 
wide ; Y-suture distinct ; vertex flatly arched ; sides convex 
and curvate ; trochantin of mandible not so prominent as in 
transvaalensis. 

Frons deflected and once dej^ressed ; depression cephalic ; 
fairly deep, merging into clypeus, bordered behind by a low 
transverse brow, the median region of which projects a little 
forward. 

Eyes black, sub-reniform, facing obliquely upwards. 

Ocelli in the form of two minute, oval, white spots. 

Clypeus as with transvaalensis. 

Labrum with flatly brace-shaped cephalic iDargin ; distal 
corners somewhat lobed, then tapering somewhat acutely; 
like that of transvaalensis but distal margin more flattened, 
very different from that of havilandi. 

Mandibles at base dark red, the rest black ; dentation and 
form as with transvaalensis, but with apical points much 
incurved, and more slender and both about equally coarse ; 
not so highly coloured as in havilandi and coarser; with 
margin below basal process of left irregular and not incurvate; 
with margin below second tooth of right with two short 
incisions and only a suppressed scallop between them ; cephalic 
edge of second tooth of right distinct W bi-curvate. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 439 

AntennjB yellow ; whip-like ; distal series not tapering as 
conspicuously as in transvaalensis ; composed of 27, 28, 
29, or 30 joints ; III, IV, V subequal, annular, and the three 
narrowest and smallest; VI and onwards increasing in length 
and width ; width from VII or VII [ constant up to the last 
4 to 6 apical joints which taper veiy little ; lengths reaching 
maximum at the twelfth to fourteenth joint from apex, apical 
joint always shorter than penultimate and often decidedly 
narrower. 

I strong, cylindrical, furnished at apex with distinct whorl 
of spines ; more than twice the length of II and one third 
wider (length range 400 to 500 fx). 

II strong, cylindrical, nearly as wide as long; wider than 
any succeeding joint ; furnished with a whorl of bristles at 
apex and with scattered bristles over apical half (200 to 230 fi) . 

III short, annular, less than half the length of II ; with a 
single apical row of bristles; anchylosed to II (70 to 100 /u). 

IV and V similar to III and 

VI annular; longer than V; with apical bristles (100 to 
ISO fi). 

VII larger than VI ; with apical bristles and often with a 
second whorl (110 to 150 fx) ; first few succeeding segments 
globose ; rest obconic ; apical series more elongate, with 
almost parallel sides (130 to 210 /x) ; VIII onwards, with two 
whorls of bristles and scattered bristles. 

Apical joint shorter than penultimate, sides parallel, apex 
convexly truncate (175 to 200 /li). 

Post-gena3 as with transvaalensis. 

Gula scoop-shaped; as with transvaalensis, but free 
blade with noticeably chitinised and much up-turned sides ; 
concolorous with post-genas. 

Thorax. — Almost uniformly reddish-brown, paler than 
head ; with dark paired brown spots upon meso- and 
metanotum ; agreeing in structural details with transvaal- 
ensis except that (1) the processes tend to be more rounded 
apically and more lateral in direction of cephalic edge, and 
(2) that the anterior margin of the pronotum is entire. 



440 



CLAUDE FULLER. 



Legs brown ; spurs of tibia very variable, I with 1 to o, II 
with 2 to 5, III with 3 to 4; an analysis of four insects 
gives : 



Specimen. 


a. 


2. 


3. 


4. 


Tibia . . 


I 


II 


III 


I 


II 


III 


I 


II 


III 


I 


II 


III 


On the i-ight 


2 


4 


3 


2 


4 


3 


1 


4 


4 


2 


3 


3 


On the left 


3 


4 


3 


3 


4 


3 


2 


2 


4 


2 


5 


3 



It will be observed that out of eight cases for four insects 
recorded the same sequence only occurs twice. 

Abdomen. — Whitish brown, sordid, darker than in 
pretoriensis. 

Worker. 

Measurements. — Head- width 2*2 to 4 mm. ; total length 
8"0 to 13 mm. Antennas 31 to 34 joints. 

Head. — Vertex black-brown ; the greater part of the frons 
yellow or white, sometimes with median area dark- or black- 
brown ; clypeus pale yellow at sides, dark in the middle ; 
epistome white ; labrum yellowish with a greenish tinge. 
Trochantin not projecting beyond margin. 

Ocelli, each surrounded by a diffuse ring of yellow. 

Antennte pale-brown ; whip-like ; joints I and II large and 
strong; III to V short, annular and sub-equal; III usually 
"the smallest ; remainder increasing in length to about one- 
third the distance from base to apex ; apical series elongate 
and narroAving; apical joint shorter than penultimate and 
elongate oval ; of 31, 33 and 34 joints. 

'J'horax. — Shields in the middle black-brown with a pale 
median stripe ; side-plates p^de brown ; cephalic margin of 
pronotum with a mottled white or pale yellow band. 

Pronotum much narrower than the head. Mesonotum as 
broad as pronotum with processes; metanotum broader than 
mesonotum. 

Legs tarsi and tibia pallid, femur dark brown ; I with 3, 
II with 4, III with 3 spurs to tibia. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 441 

Upon the whole agreeing witli the worker of pretori- 
ensis. 

Abdomen. — Dorsal sclerites each with dark and pale 
brown bands, paler caudad ; conjunctiva forming white 
transverse bars between sclerites ; frequently the caudal half 
of the abdomen much paler than towards thorax; venti-al 
sclerites paler than dorsal. , 

Hab. — Zululand; Somekele. Transvaal; Marico. Cape 
Colony; Kimberley. 

The habits of this species seem to agree generally with 
those of the others. Mr. David Gunn, of the Division of 
Entomology, collected the species at Marico and states that 
it seemed to be confined to the irrigated lands there. 



Hodotermes havilandi Sharp. PI. XXXI Y, figs. 5. .5a ; 
PI. XXX Y, fig. 5. 

Hodotermes liavilaudi Shdrp, The Cambridge Nat. Hist., vol. v, 
p. 384, 1895 ; Haviland, Jom'ii. Linn. Soc, vol. xxvi, p. 372, 

1898. 

Soldier. 

Measurements. — Total length 11'5 to 14"5 mm.; head 
with mandibles 5'5 to 7 mm. ; head-width 4 to 4*5 mm. ; head- 
length 3'5 to 4 mm. 

Head. — Yertex bright yellow, with three broad bands of 
a warm mahogany-red, one median and two lateral ; the 
gense yellow ; clypeus pale reddish ; frons light yellow. 
Surface polished ; vertex not quite smooth ; frons as smooth 
as vertex and without conspicuous hairs. Dorsal outline, 
round-oval, a little longer than wide ; sometimes almost 
circular ; Y-suture distinct ; vertex flatly arched ; sides con- 
vex and curvate; trochantin of mandible not very prominent 
and not projecting outwards. 

Frons deflected and twice depressed, depressions ver}^ 
shallow and separated by a simple but distinct transverse 



442 CLAUDE FULLER. 

ridge which is interrupted in the middle and has a faint 
elevation anterior to the break. 

Eyes black, sab-reniform, facing obliquely outwards. 

Ocelli obscurely represented by pale spots. 

Clypeus with a triple edged cephalic mai-gin (the epi- 
stome) ; median edge extending over labrum as a thin pale 
plate ; without a transverse ridge, but with a faint median 
elevation. 

Labrum with lobed sides as in mossambicus, but with a 
flatly convex and not brace-shaped or acute distal margin, as 
with the other allied species. 

Mandibles yellow-red over greater area ; apical points and 
teeth black. Much lighter in colour and with a greater 
coloured area than in mossambicus. Dentation and form 
as in transvaalensis but with outer margins moi*e curvate ; 
apical points sub-lunate, fine, long, slender and much 
incurved ; that of left but a little more slender than of right ; 
incisions above first tooth very acute. Both mandibles 
distinctly narrower and more incurved than those of 
mossambicus; the third tooth of the left more suppressed 
than in either mossambicus or transvaalensis, the 
process below it more acute ; the margin below process 
bicurvate, less oblique than in transvaalensis. Right 
mandible with cutting edge like that of transvaalensis 
and with three incisions and two scallops below second 
tooth. 

Antennae whip-like ; joints III, TV, and V sub-equal ; III 
the smallest, or IV sometimes smaller than III and V, or IV 
longer than III and wider than V. Number of joints 31. 

Post-gen^ as with transvaalensis. 

Gula much paler than post-genfe ; scoop-shaped ; free 
blade sub-oblong with straight sides and straight distal 
margin ; sides not conspicuously chitinised ; inserted stem 
short. 

Thorax. — Pale yellow suffused with mahogany-red, 
cephalic border of pronotum pale and almost Avhite. Struc- 
tural details in general ae'reement with those of trans- 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 443 

vaalensis except that (1) the anterior margm of the 
pronotum is almost semi-circular and entire, (2) the pro- 
cesses more like sessile bat's ears, (3) the mesonotum 
distinctly broader than the pronotum and processes, (4) the 
metanotum is broader than the mesonotum, (5) the colour is 
different. 

Legs pale reddish-yellow ; having 2 or 3 spurs to tibia of 
I ; 2 or 4 to that of II ; and always 3 to that of III. Analysis 
of 14 gives : six with 2-4-3, two with 2-3-3, three with 
3-3-3, three with 3-4-3. 

Abdomen. — Bright yellow. 

The soldier of havilandi is distinctly different from that 
here described as mossambicus; the chief points of differ- 
ence being (a) the more circular head outline, [h] in colour, 
(c) in size-range, (d) in the character of the mandible. 

Worker. 

Measurements. — Head-widths 2 to 3'5 mm. : total leng-th 
7 to 10"5 mm. 

Antennte about 29 to 33 joints. 

Except that (1) the prevailing colour of the workers of this 
species is chestnut brown and not black-brown, and (2) the 
largest workers do not reach the maximum size of those of 
mossambicus, and the smallest are much smallei-, there 
appears to be little difference between the workers of the 
two species. The description for the worker of preto- 
riensis applies in genei'al, but the V mark of the frons is 
absent. 

Hab. — Natal; Weenen, Colenso (the valley of the Tugela 
River) , 

The habits of this species agree with those of H. trans- 
vaalensis. It is often very destructive to lucerne crops. 
When infested lands are irrigated the insects suspend opera- 
tions. 

The species is very local and not found far from alluvial 
soil. 

VOL. 3, PART 2. 30 



444 CLAUDE FULLER. 

Hodotermes wari'eni sp. n. PI. XXXIA^, figs- ^j ^^^ > 
PI. XXXV, fig. 6. 

Soldier. 

Measurements. — Total length 8 to 14 mm.; head with 
mandibles 3'5 to 6*5 mm. ; head-width 2 to 3*7 mm. 

Head. — Resembling dully polished tortoise-shell; some- 
times a rich reddish-brown with colour bands ill-defined. 
Vertex with a bright orange-yellow ground and three deep 
red-brown furcate-lanceolate sti-ipes; median stripe fusiform, 
extending along stem of Y-suture ; sub-median stripes extend- 
ing from caudad of eyes over occipital region, then forming a 
median line on the ventral slopes of the gen^ ; clypeus 
reddish in the middle, bright yellow at sides ; small lateral 
prolongation behind mandibles black. Surface polished; the 
whole finely wrinkled ; cephalic depression of frontal area 
distinctly wrinkled ; without conspicuous hairs. Dorsal out- 
line broadly U-shaped; longer than wide; sides parallel, 
nearly straight or flatly curvate ; Y-suture distinct; vertex 
broadly arched ; trochantin of mandible not very con- 
spicuous. 

Frons not conspicuously impressed. 

Eyes black, sub-reniform. 

Ocelli small, indistinct, and not bordered b}^ a darker 

colour. 

Clypeus with the cephalic edge (epistome) rounded and 

deflected anterior to a transverse ridge; frontal folds distinct, 

oblique, extending well backwards. 

Labrum pale yellow, very broadly cordate, as in 
transvaalensis, but a little more roundly acute and 
proportionately longer. 

Mandibles in form and dentation as with transvaalensis; 
apical points acute and narrow, but not as slender as in 
havilandi; first tooth of left elongate, acute, and directed 
somewhat towards apical point ; basal process acute and very 
prominent ; second tooth of right comparatively large, margin 
below wavy and not incised. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 445 

Antennas yellow; whip-like; much as with transvaal- 
•ensis ; about 25 to 29 joints ; joint III larger or smaller than 
IV. 

Post-genee as with transvaalensis, but mottled with 
brown. 

Cxula scoop-shaped ; with sides of free blade much upturned, 
straight ; distal margin straight, anterior lateral corners 
oblique. 

Thorax. — Pale yellow or yellow with median patches of 
red-bi*own on each shield. 

Pronotum arched ; cephalic margin entire ; processes 
deflected, projecting forward and not outward, not incised 
behind; sides straight; tapering; caudal margin slightly 
incurvate, nearly straight. 

Mesonotum broadly oval, narrower than pro- and meta- 
notum ; caudal margin slightly truncate. 

Metanotum short and broad ; broader than pronotum ; 
widest behind ; sides nearly straight, oblique and diverging ; 
caudal margin three-sided, all three edges straight or broadly 
incurvate. 

Legs pale brown-yellow. Tibia of I with 2 to 3, II with 3 
to 4, and III with 3 spurs. 

A bdomen. — Yellow. 

Worker. 

Measurements. — Head-width 2 to 3*5 mm.; total length 
6 to 10 mm. 

The worker of these species is much like that of pretori- 
ensis, but presents a lower length-range, and the frontal 
depression of the frons is very shallow. 

Hab. — Orange Free State; Lovat, Tha'banchu. 

Reported as very destructive to veld grass during the 
drought of 1914. The field series collected at Tha'banchu in 
August, 1914, contains an obese worker-like creature, larger 
than the soldiers with a rounded yellow head and pale yellow 
body. This is probably an immature imago. 



446 CLAUDE FULLER. 



Hodotermes braini -sp. n. PI. XXXIV, figs. 7, 7a; 
PL XXXV, fig. 7. 

Soldier. 

Measure m e n t s . — Total lengtli 1 2 mm. ; head with man- 
dibles 5 mm. ; head-width 3 mm. 

Head. — Vertex marked by a broad median, parallel-sided 
stripe of pale brown which seems to extend from the 
clypeus to the occiput, and is about half the breadth of 
the head ; lateral to this the field is dark brown to the sides. 
Clypeus reddish-yellow. Frons yellow with a dark, ill-defined 
median area. Surface dulled ; frons and clypeus faintly 
wi-inkled. Dorsal outline distinctly oblong, U-shaped and 
flatly rounded behind ; Y-suture distinct ; vertex flatly arched 
and not depressed; sides parallel and straight, somewhat 
constricted, tapering from laterad of ej'es to trochantin of 
mandible ; trochantin well defined, concolorous with adjacent 
parts. 

Frons deflected slightly and but faintly impressed. 
Eyes black, sub-reniform. 

Ocelli small, but distinct, oval, white, surrounded by a 
narrow ring of dark red-brown and a difi^use band of pale 
yellow. 

Clypeus oblong ; with cephalic corners of epistome rounded 
and much deflected^ giving it a three-sided appearance ; with 
a median, cephalic, flat, pyramidal elevation; frontal folds 
indistinct. 

Labrum mu^ch like that of transvaalensis, but more elon- 
gate and front margin more acute. 

Mandibles black, bases suft'used with dark red ; dentation 
as with transvaalensis ; comparatively slender and 
straight ; apical points long and not couspicuously incurved, 
that of left nearly straight ; on the dorsal surface, at the 
base of each, a small punctate area ; process of left acute and 
deflected; margin below second tooth of right, twice incised 
and bicrenulate between the incisions. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 447 

AnteniiaB with basal half yellow, rest smoky ; apical 
ses^raents not decidedly tapering ; last segment short, very 
little longer than wide ; joints of one complete specimen 
23. 

Post-genas and gula much as with transvaalensis, but 
gula more elongate and much paler than post-gena3. 

Thorax. — Pale yellow-brown with pale red-brown patches 
in median field of each segment. 

Pronotum nnich like that of havilandi; cephalic margin 
arched and entire; processes acute. 

Mesonotum large, as wide, if not wider, than pronotum and 
as long ; sides diverging with curvate margins ; caudal 
margin broadly and distinctly bilobed. 

Metanotum as wide as mesonotum, but shorter; caudal 
margin three-sided, lateral edges curvate, posterior straight. 

Legs yellow ; tarsi of I with 2 to 3 spurs, II with 3 to 4, 
III with 3. 

Abdomen. — Golden yellow. 

WOEKEB. 

Measurements. — Total length 7 to 8 mm.; head-widths 
1'8 to 3 mm. 

The salient features agree with those of the workers of 
pretoriensis but without V mark or spot on frons. The 
size of the workers is consistently small and presents the 
smallest range of measurements for our Hodotermes. 

Hab. — Transvaal; Pretoria district. 

This species is erected upon a good series of workers, but 
unfortunately only one soldier has been found ; it is, however, 
sufficiently characteristic, especially as regards the alignment 
of the mandibles, to warrant this description. The oppor- 
tuuiry of investigating its habits has not yet presented itself, 
but it may be stated that numerous workers and callows 
were found in small mole-hill-like deposits of earth particles. 



44S CLAUDE FULLER. 



Hoclotermes viator {LatrieUe). PI. XXXIY, figs. 8, 8a; 
PL XXXY, fig. 8. 

Termes viator LatrieUe. Ji\&i. Nat. des Crast. et des Ins., xiii. p. 
51, 1805. 

Soldier. 

Measurements. — Total length 7 to 13 mm.; head with 
mandibles 3 to 6 mm. ; head-width 2"3 to 3 mm. 

Head. — The whole dark red-brown, cephalic half very 
dark in the larger and paler in the smaller soldiers. Surface 
polished. Dorsal outline U-shaped, as broad as long, widest 
at cephalic edge; Y-suture faint or distinct; vertex flatly- 
arched ; sides straight, parallel, tapering in lateral to eyes, 
often asymmetrical, often flatly cnrvate or incurvate ; trochan- 
tin of mandible strong and distinct. 

Frons deflected and twice depressed ; depressions separated 
by a wide, rounded transverse ridge running between the 
ocelli ; caudal depression shallowing into vertex. 

Eyes black, facing forward and outward. 

Ocelli white and defined in small soldiers ; in large masked 
by chitin ; represented by pale aureolate spots which under a 
low magnification appear as dark raised points. 

Clypeus with anterior margin sti-aight and usually bordered 
with a pale yellow, somewhat inflated, brace-shaped sti-ap (the 
epistome). 

Labrum bright but opaque, its orange yellow colour in 
strong contrast to surroundings; large, spatulate and arched; 
distal edge convex with a broad convex median prolongation 
quite different from that in the foregoing species of 
H o d o t e r m e s. 

Mandibles black, except for a small reddish patch at inner 
bases ; this is covered by the lateral lobes of the labrum, 
both comparatively straight ; dentation as with other species, 
but teeth sharp and more on a median alignment, so that the 
lower margin forms a cutting edge ; lower margin of each 
tooth on left distinctly lobed ; first and second tooth on both 
attenuated and sharply pointed ; basal process of left having 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 449 

a strong, blunt, conical tooth ; margin below second tooth on 
left once incised. 

Antennas opaque yellow-brown throughout ; not conspicu- 
ously haired; much like that of transvaa lensis ; about 
26 to 27 joints ; I particularly long, wider at the apex than 
the base and with concave sides ; II strong, cylindrical, about 
one-third the length of I ; III very small ; IV (in some) much 
larger tlian III and larger than V ; V larger than III, a little 
shorter and narrower than IV ; VI, VII, VIII equal and the 
same as VI; basal joints from III onwards are first annular, 
then moniliform or globose, then short obconic ; the apical joint 
is elongate oval and the three preceding it elongate obconic. 
(This pattern is also present in some of the smallest soldiers, 
but occa^sionally joint III is quite large, larger than IV.) 

Post-gense as in transvaalensis. 

Grula much darker than the post-genae ; scoop-shaped ; free 
blade with up-curved, parallel sides and convex median 
region ; distal apex soft incurvate, normally white ; inserted 
stem half cylindrical, narrower than the blade ; very short 
and constricted. 

Thorax. — Brownish or reddish-yellow or concolorous with 
abdomen. 

Pronotuin narrower than the head, in general resembling 
that of transvaalensis; cephalic margin acutely elevated 
(45°), almost semicii-cular, entire; sides upbent, margins 
curvate and tapering behind, hind margin broadly incurvate ; 
lateral processes somewhat deflected, projecting outwards 
from bases of cephalic arch at right angles to median line of 
body ; front margin straight, hind margin curvate, curve 
continuous- with that of sides. 

Mesonotum narrower than pronotum ; almost oblong ; 
caudal margin almost straight, but slightly concave at 
middle ; sides bent, curving inwards in front, outer edges 
straight and parallel, behind bent inwards at an obtuse 
angle. 

Metanotum narrower than mesonotum, caudal margin 
straight. 



450 CLAUDE FULLER. 

Legs concolorous with abdomen ; last joint always longer 
than the remaining three taken together. Tibia of I with 3, 
II with 4, and III with 2 spurs. For a series this 3, 4, 3 
feature is fairly constant. 

Abdomen. — A bright golden yellow and always in sharp 
contrast with head. 

Worker. 

Measurements. — Total length 6 to 8'5 mm. ; head-width 
1*6 to 3 mm. 

Head. — Reddish-yellow or chestnut-red; when pale, 
colour uniform, Y-suture paler, eyes black and in strong 
contrast, aerolate yellow patches at ocelli ; when dark, colour 
uniform but Y-suture not pale, ocelli areas pale, clypeuspale. 
Surface highly polished. Dorsal outline broadly U-shaped ; 
cephalic extremity broadened; sides nearly straight and 
parallel, bent in sharply to trochantiu of mandible ; vertex 
rather roundly arched. 

Frons twice depressed ; caudal depression very shallow ; 
cephalic depression merging into clypeus and with a median 
elevation ; cephalic and caudal depressions separated by a 
low transverse ridge. 

Eyes dark chestnut brown, sub-reniform. 

Ocelli obsolete, represented by pale spots. 

Clypeus differentiated from frons by a transverse furrow ; 
cephalic margin straight and elevated, forming with the 
epistome a transverse ridge ; epistome translucent, cephalic 
margin narrow, faintly curvate, sides oblique. 

Labrum a bright orange-yellow. 

Antennae pale yellow throughout ; joints annular to half the 
length, then broad and short, then elongate. Apical portion 
tapering very faintly. 

Thorax. — Concolorous with abdomen; cephalic margin of 
pronotum arched, acutely curvate and eutire ; processes 
deeply incised at junction with arch of cephalic margin ; 
caudal margin widely bilobed. 

Mesonotum long and broad ; caudal margin broader than 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 451 

pronotum ; sides straight, diverging outwards ; caudal margin 
faintly bilobed. 

Metanotura nearly as long as mesonotuni and wider; sides 
similar ; caudal margin flatly curved. 

Legs uniformly concolorous with abdomen. 

Abdomen, — Bright red-gold; somewhat wide towards 
extremity ; not banded. 

Hab. — Cape Colony; Stellenbosch,Elsenberg, Victoria East. 

This is a well-known and destructive insect in the older 
settled regions of the Cape, and is that to which the term 
" hout-kapper " was first applied. It is strikingly different 
from any of the other species mentioned in this paper, but 
much like aurivellii, another distinct species, a description 
of which is deferred. 

Calotermes. 

Calotermes durbanesis Haviland. PI. XXXIV, fig. 9. 

Calotermes durbanensis Haviland, Journ. Lmn. Soc. Lond., 
xxvi, p. 377, 1898. 

Soldier. 

Measurements. — Total length 7 to 10 mm. ; head Avith 
mandibles 3"5 to 3'7 mm. ; head-width 1"5 to 1'7 mm. 

Head . — Vertex dark brown, clypeus black, labrum reddish. 
Surface polished, cylindrical. Viewed dorsally, elongate, 
sub-oblong ; sides straight and parallel ; vertex with a distinct 
shallow median depression which extends backward from the 
frons over half the length of the head; Y-suture absent; 
occipital region not indented ; trochantin of mandible pro- 
jecting conspicuously. 

Frons precipitous, depressed, wrinkled ; cephalic margin 
straight ; parted from the clypeus by a transverse ridge wliich 
generally presents paired median points. 

Eyes almost obsolete, pale. Ocelli absent. 

Clypeus pale, sub-oblong, with a straight cephalic margin 
and oblique sides. 

Labrum short, almost circular. 



452 CLAUDE FULLER. 

Mandibles black, faintly tinged with maliogany-red, bases 
reddisli ; straight, comparatively slender, right the stouter, 
both equally long, with incurved points and with very con- 
spicuous, shoi't, caudo-lateral shoulders. Left mandible with 
two teeth upon distal half and two proximal to these, the pairs 
being separated by an acute incision ; below the lower pair 
an acute basal process ; the distal tooth is lobe-like and con- 
stricted at its base, the second formed like a cleaver, with a 
bicurvate blade, the third convex, the fourth blunt, conical 
and more salient than any; fourth separated from the third 
by a shallow concave incision; margin below fourth incurvate 
and deeply incised distal to the process. Right with two 
teeth on proximal half ; below the second tooth a sharp 
incision, the base then broadening; the distal tooth is 
cleaver-shaped with a blunt conical apex and bi-curvate 
blade ; the second cleaver-shaped and with both margins 
bi-curvate. 

Anteunte 12 to 13 joints; arising from below a con- 
spicuous hood ; short; IV and onwards moniliform ; charac- 
terised by the long highly chitinised clavate joint III and 
the erratic fusion of more distal joints; II a little longer than 
wide ; III twice the length of II and clavate ; apical joint 
oval. 

Gula with free apex very much broader and shorter than 
inserted portion, with a rounded apex and tri-curvate, sub- 
parallel sides. Inserted portion elongate narro-w ; caudal 
extremity narrow; sides at first parallel and then diverging 
outward ; not constricted as in Hodotermes. 

Thorax. — Sordid yellow-brown; much arched, with 
margins of pro-, meso-, and metanotum much deflected and 
introse. 

Pronotum large ; broader than head ; cephalic margin 
incurvate Avith a very faint median notch ; sides curvate, 
caudal margin broadly curvate; with paired elongate pyri- 
form dark brown spots in cephalo-lateral corners, which are 
connected by a wavy ti-ansverse furrow imuiediately posterior 
to cephalic margin. 



OBSEEVATIONS ON SOME SOUTH AFIUOAN TERMITES. 453 

Meso- and metanotum much narrower than, and about half 
the length of, the pronotum. 

Legs pale and dark brown ; short and stout ; more or less 
highly chitinised ; femur broad. 

Abdomen. — Sordid yellow-brown; concolorous with 
thorax. 

Hab. — Natal; Durban, Bellair, Winklespruit. 

The species here discussed agrees with the Haviland 
material, and some specimens were obtained from the same 
source. If durbanensis is synonymous with C. mada- 
gascarensis Wasm., as stated (2 and 4), the description 
of the mandibles of the latter is as incomplete as was Havi- 
land's description of those of durbanensis. 

Rhinoteumes. 

Rhinotermes putorius {Sjiistedt). PI. XXXIV, fig. 10.. 
Termes putorius Sjustedt. Ent. Tidskr., xvii, p. 298, 1896. 

Major Soldier. 

Measurements. — Total length 7-5 to &6 mm. ; head wdth 
mandibles 2'5 mm. ; head-width 2 mm. 

Head. — Reddish or golden red; bases of frontal folds 
chocolate brown. Dorsal outline short-ovate, somewhat 
cordate ; sides tapering and constricted lateral to antennal 
fossa3; Y-suture absent; frontal folds clearly expressed in 
the form of a V with carved sides ; vertex faintly wrinkled ; 
area of front indicated by several strong wi-inkles which run 
parallel with suture and are V-shaped. 

Eyes minute and pale. Ocelli and fontanelle obsolete. 

Clypeus with cephalic margin straight with a thin plate 
(epistome) ; sides diverging to margin of head ; limited 
caudally by the two frontal margins of the frons, which 
converge to a large white foramen ; the foramen seems to be- 
placed in the clypeo-frontal suture and lies in the foi'k of the 
frontal folds, well in the middle of the anterior region of the- 



454 CLAUDE FULLER. 

head ; from the foramen a distinct, grooved ridge slopes 
down to the cephalic margin. 

Labrum short and broad, distinctly divided into two convex 
regions by a median groove, which forms a continuation of the 
clypeus groove and permits any secretion from the clypeo- 
frontal foramen to flow to the apex of the labrum ; cephalic 
margin of chitinised portion distinctly bilobed, and with a 
distal plate-like extension whose curvate margin is fimbriate ; 
on the disc behind the fringe are several (six to eight) stout, 
spine-like bristles ; sides bicurvate, at first diverging then 
converging. 

Mandibles deep red-brown in sharp contrast with head 
colour ; short and stout with long incurved apical points ; 
left with two acute teeth ; right with one ; neither with a 
basal process ; incisions above teeth very acute. 

Antenna? with I stout and cylindrical ; II cylindrical, 
shorter than I ; III elongate, sub-obconic, much narrower at 
base and longer than II ; IV and onwards moniliform ; IV 
large and broader than III and longer and broader than V ; 
V-XVI sub-equal; XVII not longer than XVI, tapering a 
little, and roundly truncate, XV to XVI-jointed ; IV and V 
fused in XV-jointed. 

Grula with free apex short ; distal margin incurvate ; nearly 
straight ; sides diverging and incurvate. Inserted stem 
narrow, long, sides slightly incurvate; concolorous with 
ventral gense, with dark stripes in lateral sutures. 
Thorax. — Concolorous with head ; polished. 
Pronotum broadly sub-oval; median region convex ; cephalic 
margin broadly curvate ; sides tapering and merging into 
caudal margin which is faintly bilobed. 

Mesonotum large ; sides broadly curvate, diverging widely ; 
caudo-lateral corners rounded ; caudal margin as wide or 
faintly wider than pronotum, curvate and faintly bilobed. 

Metanotum with oblique, slightly cui'vate sides ; caudal 
margin not as wide as that of pronotum, curvate, nearly 
straight ; caudo-lateral corners rounded, but more acute than 
those of mesonotum. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 455 

Legs pale yellow. 

Abdomen. — Concolorous with thorax; styli and cerci 
prominent. 

Minor Soldier. 

Measurements. — Total length 4 to 5 mm.; head and 
mandibles 1"5 mm.; head-width 1 mm. 

Resembling major soldier, but having a more elongate sub- 
pyriform head and proportionately longer jaws and labrnm. 
The labrura tapers fi-om its base to apex, and the sides are 
bicurvate, curvate at the base and incurvate towards the apex, 
so forming with the cephalic margin small lobed corners. 

Antennae shorty 15 or 16 joints. 

Worker. 

Measurements. — Total length 4'5 mm.; head-width 1*2 
to 1*5 mm. 

Head. — Pallid, yellow; smooth; Y-suture distinct with 
triangular splash of white in fork ; two small pennant strips 
extending caudad from arms of suture at half their length ; 
at lateral extremities, behind antennal fosste, large blotches 
of white ; cephalic part of frons with two square patches 
of yellow just lateral to median line, lateral to these and 
medial to frontal cai-inas two irregular yellow patches ;. 
frontal carinse with fine brown lines which merge into 
two concolorous spots at superior attachments of mandibles ; 
clypeus, epistome and labrum concolorous with head ;. 
mandibles yellowish with white spots ; teeth of mandibles 
dark brown. 

Antennae pallid, moniliform, XVI or XVII-jointed. 

Hab. — Natal; Durban. 

This is the insect which Haviland noted in the Botanic 
Gardens, Durban, and hesitated to describe in the absence of 
imaffos. 



456 CLAUDE PULLER. 



Termes. 



A study of a lono- series of soldiers and workers from 
separate nest-series and different localities seems to indicate 
•that the various insects known as Termes monodon 
GerstacJcer (1891), badius Haviland (1898), t err i cola 
Sjostedt (1904) and transvaalensis Sjostedt (1904) are all 
one and the same insect; if so, the last three fall as 
-Synonyms to monodon. The reasons for this supposition 
are as follows : 

(a) The soldier of T. monodon is said to differ from 
badius (1) in being larger (monodon, total length 7 to 11 
mm., head with mandibles 4 to 5"2 mm.; badius, total 
length 7 to 10 mm., head with mandibles 4 to 4*5 mm.) ; (2) 
in that the eyespots appear to l)e sometimes absent ; (3) in 
the presence of a rounded median (?) keel upon the frons. 
But (1) in a nest-series of badius before me the full length 
range of spirit material is 9 to 12 mm., with head and 

mandibles measuring 4*5 to 5 inm., (2) when killed in spirit 
many soldiers of badius have the muscles of the head densely 
stained by the red-staining fluid they eject when attacked and 
-then the eyespots are always seemingly absent; (3) badius 
presents a low-rounded transverse keel, and in certain lights 
and on certain insects even a low median keel can be 
detected ; (4) I have before me workers of T. badius which 
have been determined through Dr. Sjostedt as those of T. 
m o n o d o n . 

(b) The description of T. terricola is the best given in 
the monograph for the series under review, and, as far as it 
goes, it applies wholly to badius. It is said to differ (1) in 
point of size, and (2) in certain impressions or markings on 
the frons. But (1) it is well within the badius range, and 
(2) these markings (more or less obsolete ocelli) are always 
present and, with the minute fontanelle, are quite distinct 
features in any cleared preparations of the heads of T. 
badius. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 457 

(c) The soldier of T. transvaalensis is said to differ from 
badius (1) in being smaller aud (2) in having XVI-jointed 
antennae. But (1) the size of transvaalensis comes 
within the badius range; (2) I have nest-series of badius 
fi'om Natal, the soldiers of which are less (7 to 8 mm.) than 
the transvaalensis range, but unmistakably badius; in 
another nest collection of badius from Pretoria this range 
is, total length 7 to 11 nnn., head with mandibules 3'6 to 4*9 
mm. ; (3) the autotype material shows both XVII and 
XVI-jointed antennae (in one instance a soldier having the 
typical XVII-jointed antenna on the left and a pseudo 
XVI-jointed on the right) ; further, the XVI-jointed antennge 
are all obviously due to the non-separation of joints III and 
IV ; in a field collection made in Pretoria the several soldiers 
present have perfect XVI-jointed antennae and conform in 
size to that given for transvaalensis, but some show the 
pseudo-ai'ticulation and moreover the workers display the 
several antennal patterns of badius and agree in all other 
features; further, in a nest collection from the same locality, 
soldiers possessing XVI, XVII, and pseudo XVI-jointed 
antennas are present whilst the worker chai'acters conform to 
those of badius; (4) the transvaalensis autotypes have 
the same peculiar inverted rose-thorn-like tooth near the 
base of the right mandible; (5) the imagos of T. badius 
conform in every particular with the description of the 
imagos of T. transvaalensis. 

Apart from Haviland's material of badius and Sj6stedt^s 
autotypes of transvaalensis, the series examined comprises 
twenty-two collections of soldiers and workers ; of these ten 
were obtained from separate nests and four are field collec- 
tions from in and around Pretoria ; five are field collections 
and one a nest collection from various other points in the 
Transvaal ; one is a field collection from the Orange Free 
State, another from Pitermaritzburg, Natal. With few 
exceptions the collections comprise a long series of soldiers 
and workei'S; in five, kings and queens are represented. 
There are also three nest collections and two field collections 



458 CLAUDE FULLER. 

of winged adults made in Pretoria, and one nest collection 
from Alberton, Transvaal. 

With regard to T. natalensis Haviland, it is necessary to 
point out that the Haviland material ^ contains major (and 
minor) soldiers of several forms. One series (a) possessing 
large and thick heads and flattened frons, and of which 
certain members of a nest-series display a very aberrant 
meso-thorax ; aiiother (b) having a rather flattened fi-ons, the 
sides of the meso-thorax rounded and the heads large and 
thick," and (c) a third series in which the frons is deeply 
impressed and grooved. The prevalent South African form 
is T. natalensis Haviland, and it seems to be T. belli- 
cosus Smeathman ; but, being unable to decide this point, I 
have grouped, for the present purpose, the whole collection of 
nest-series studied as T. natalensis Hav. 

I have relied upon the constant features of the mandibles 
and the antennse of the Haviland material. 

I have come to the conclusion that the aberrant forms 
which happen to predominate in the portion of Haviland's 
material before me represent a locality variation. The pre- 
vailing form, in my collection, comes from the coast, from the 
thorn country at Kimberley, and from the thorn country of 
the tablelands of the Transvaal. The several nest-sei-ies from 
the midlands of Natal illustrate both the typical and aberrant 
forms of T. natalensis. The latter form is at once recog- 
nisable by the presence of certain soldiers (major or minor, or 
both), of which the meso-thorax presents caudo-lateral lobes, 
and the sides, instead of being rounded, are oblique and 
divergent. That Haviland regarded this as only an aberra- 
tion is obvious from the fact that he does not mention it in 
his description. As to whether natalensis is synonymous 

' The Haviland material in the Natal Museum comprises a number 
of insects from different nest-series collected and labelled by Haviland 
himself ; some of those before me were actually measured l^y liim and, 
no doubt, for the purpose of his original description. 

^ Like the types, as far as one can jiidge from Professor Sjostedt's 
discussion. 



OBSEEVATIONS ON SOME SOUTH AFRICAN TERMITES. 459 

with belli cosus, or whether under the circumstances one 
can select a portion of the Haviland series as a variety^ I am 
not in a position to say; but it can be said that the differences 
cited for the two species in the Monogi-aph (2) are not of 
specific value. 

Some particulars regarding my examinations of T . natal- 
ensis may not be out of place here. 

P'orty series examined composed as follows : 

Fourteen nest collections represented by soldiers, workers, 
and imag-os (kings and queens). From Natal, three (Mount 
Edgecombe, Bellair, Winterton). From the Transvaal, 
eleven (Pretoria, Pietersburg, Barbei-ton). 

Seven nest collections represented by soldiers and workers 
only. From Natal, four (Pietermaritzbiirg, New Hanover, 
Weeueu). From the Transvaal, three (Pretoria, Tzaneen). 

Twelve nest collections represented by two or more soldiers 
from each (Haviland material from the district of Plstcourt, 
Natal). 

Seven field collections represented by soldiers and workers. 
From Natal, two (Bellair, Pietermaritzburg). From the Trans- 
vaal, five (Pretoria and district). 

An examination of the series of major soldiers before me 
brings out one or two peculiar features. The first is that, as 
regards colour, markings, and general appearance, the longer 
material has been in alcohol the more do the specimens 
harmonise. Thus whilst Haviland's insects, which have been 
in spirit since 1898, are all agreeably alike ; colour notes, 
made from certain material, when freshly preserved twelve 
luonths ago, do not wholly apply at the present moment. 
Again, many colour markings and features are more readily 
seen upon submerged specimens than dry ones. On the other 
hand certain other features may be obscured, more particularly 
the sculpturing of the frons. The appearance of the insects 
is changed in alcohol as regards shape as well as colour ; the 
whole insect becomes clarified, and the inflation of the abdo- 
men, so marked in living material, disappears, probably by 
the slow dissolving of certain body-contents and the escape 
of gases and air. 

VOL. 3, PART 2. 31 



460 



CLAUDE FULLER. 



A study of both major and minor soldiers shows that the 
length of the insect has but little relation to the length of 
the head. 

With regard to the total length of major and minor soldiers, 
the major soldiers in my collection have a range of from 13 to 
18 mm. and the minors of from 8 to 12 mm. A purely mis- 
cellaneous series of measurements of 155 major and 68 minor 
soldiers gives the following analysis : 



Major soldiers. 


Minor .soldiers. 


Total length. 


Number. 


Total length. 


Number. 


Mm. 
18-4 - 18-0 
17-9 - 17-5 
17-4 - 17-0 
16-9 - 16-5 
16-4 - 160 
15-9 - 15-5 
15-4 _ 150 
14-9 - 14-5 
14-4 _ 14-0 

13-9 - 13-5 
13-4 - 130 


1 



5 

6 

32 

14 

51 

12 

23 

7 

4 


Mm. 

11-9 - 11-5 

11-4 - 110 

10-9 - 10-5 

10-4 - 100 

9-9 - 9-5 

9-4 - 9-0 

8-9 - 8-5 

8-4 - 8-0 


1 

4 

4 

• 12 

13 

23 

3 

3 


Total . . 


155 


Total . . 


63 



The following tables, showing the range of variation, are 
based upon the measurements of 10 major soldiers and 10 
minor soldiers, or 100 insects in all, from five widely separated 
nests in regions having very different climatic conditions 
(comprising at least four of the floral regions pi'oposed by 
the late Dr. Bolus). 

No. 1 nest : Mount Edgecombe, Natal, at an elevation of 
201 ft. and in coast bush country. 

No. 2 nest : Winterton, Natal, elevation 3354 ft., in grass 
veld. 

No. 3 nest: Pretoria, Transvaal, elevation 4471 ft., located 
below a house. 



OBSERVATIONS ON SOMh] SOUTH AFBECAN TERMITES. 461 

No. 4 nest: Barbertoii, Tivmsvaal, elevation 2877 ft., in 
middle veld bush countiy. 

No. 5 nest : Kimberley, Grricjualaud West, elevation 4012 ft. 
arid Karroo veld. 



Major soldiers. 


Nest. 


No. 1. 


No. 2. 


No. 3. 


No. 4. 


No. 5. 


Total length 
Head with 

mandibles 
Head-width 


Mm. 
14 to 16 

6-5 to 8 

3-8 to 4-2 


Mm. 
14 to 15 

7-5 to 8 

4-2 to 4-5 


Mm. 
14-2 to 17-5 

7-2 to 8 

4 to 4-5 


Mm. 
14-2 to 16 

7 to 8 

4 to 4-5 


Mm. 
14-5tol5-2 

7 to 8-4 

4 to 4-8 


Minor soldiers. 


Total length 
Head v/ith 

mandibles 
Head-width 


8 to 9-7 
4 to 4-5 

2-1 to 2-5 


8-2 to 10 
4 to 5 

2 to 2-5 


9 to 12 
4-7 to 5 

2-5 to 3 


9 to 10 
4-2 to 5 

2-1 to 2-7 


9 to 10 
4-2 to 5 

2-2 to 2-5 



Of the major soldiers enumerated in the first table the 
length of the head with the mandibles was taken from 
97 individuals, and it showed a range of from 6'5 to 
8 mm. Of the whole, only 3 measured less than 7 mm., 
32 measured between 7"0-7'4 mm., 38 betAveen 7"5-7"9 mm. 
and 25 measured 8*0-8"4mm. 

In these cases it was found that the ratio of the length of 
the head with the mandibles to the length of the thorax + 
abdomen varied widely. The distribution of these ratios is 
given in the accompanying table : 



Head + mandibles 


133 


125 
1 


114 

19 


111 
1 


107 

4 


100 

24 


98 

6 


94 

2 


90 

3 


87 

14 


82 

19 


79 
1 


68 

2 


Thorax + abdomen ^ 100 . 


No. . . . . 


1 



462 CLAUDE rULLER. 

The curvature of the head varies within certain limits 
sufficiently to give that organ quite different appearances ; 
this occurs in the case of both major and minor soldiers. 
The dorsal outline is always horse-shoe shaped, but may be 
attenuated, in which case the length exceeds the width ; or it 
may be more clumsy, in which case the length more nearly 
approaches the width, and the thickness of the head is 
increased. In the major soldiers the usual proportion of 
length to width is 5 to 4, and this may vary to 4 to 4. The 
shortening of the head adds to its thickness and the alteration 
of its outline, and the measurements taken appear to indicate 
that the head of the major soldier tends to reach a certain 
capacity, and the shortening of one dimension is compensated 
for by an increase of another dimension. 

It should be added that the greatest deviation from the 
normal, e.g. from the attenuated and more or less flattened 
head, was found in the case of certain Pietermai'itzburg and 
Weenen material, the soldiers presenting a strikingly angled 
meson otum. 

Termes swazi^e, sj^. n. PI. XXXIV, fig. 11. 

Major Soldier. 

Measurements. — Total length, 20 to 22-2 mm.; head 
with mandibles, 9 to 9"5 mm. ; head-width 5'6 to 6 mm. 

Head. — Massive; rich dark chestnut-brown with a long 
median and dark stripe; generally paler on gence and ventral 
genas ; distal margin of clypeus obscurely yellow (the colour 
of the head of the major soldier in sharp contrast with that of 
the minor). Vertex broadly arched and somewhat flat; dully 
polished, with scattered, fairly stout reddish bristles, a pro- 
minent group on the frontal area. Dorsal outline somewhat 
jar-shaped; caudal margin very flatly curvate with rounded 
corners; sides sub-parallel or tapering and flatly curvate, 
constricted and incurvate a little posterior to antennal fossae. 

Fontanelle very distinct, black, with a few radiating 
wrinkles ; Y-suture absent. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 463 

Frontal area much depressed with a median, elongate, 
triangular and transversely wrinkled mound ; mound with its 
apex a little anterior to the fontanelle and its base upon the 
cephalic margin of the clypeus; lateral to this mound are 
fairly deep converging grooves; lateral to its apex and in the 
depression paired suppressed mounds. 

Eyespots pale and insignificant. 

Ocelli represented by a mark which has the appearance of 
a closed eye. 

Clypeus with a cephalic margin (epistome) three-sided, 
distal margin straight or a little incurvate ; distal corners 
oblique and straight ; sides of clypeus incurvate and parallel. 

Labrum concolorous with head ; sides curvate, tapering, 
but almost parellel, apex with a large white cordate lingula 
(epipharyngeal ) . 

Mandibles black throughout; powerful; simple; long and 
tapering; incurved and upcurved ; with tapering bases which 
penetrate deeply into epicranium. Left broader than right, 
larger and longer (L. 8*5 mm. ; R. 3"2 mm.) ; distal half of 
inner margin of left faintly crenulate, proximal half with a 
large produced piece having two rounded points at either end 
and the margin between these incurvate ; proximal to this a 
small process. Right with inner margin entire and distinctly 
elbowed at one-third its length. 

Antennas XVII-jointed ; I and II reddish, the rest dark 
brown ; each with a distal ring of pale colour and with yellow- 
brown bristles ; length about 6 mm. ; joint I cylindrical, about 
twice the length of II (8 : 4), with a whorl of small spines; 
II anchylosed to I, cylindrical, short, nearly wide as long, with 
minute almost obsolete spines ; III clavate and long, as long 
as LI X 2*5 (9 : 4) and longer than I, bristled ; IV as long as 
I, bristled ; V to XII about equal and a little shorter than IV 
(7:6); XIII to XVI tapering and becoming shorter ; XVII 
longer than XVI (6:5); IV to XVI elongate, obconic, about 
half as wide as long, bristled. XVII elongate, oval ; apex 
convex, truncate ; bristled. Joints III and IV of the worker 
XVIlI-jointed antenna fusing to form III of the XVII-jointed. 



464 CLAUDE FULLEli. 

Gula darkei" than and about half the width of ventral 
genge ; sides parallel, incurvate ; free apex narrowed, distal 
margin straight, sides oblique with distinct caudo-lateral 
lobes. 

Ventral gena9 exhibit transverse wrinkles on that portion 
which would be delimited as the post-genae of Hodotermes 
spp. There is no indication of the suture. 

Thorax. — A very dark brown, shields outlined Avith 
black. 

Pronotum sellate, comparatively short and wide, not as 
wide as head but nearly so; with cephalic and caudal 
margins distinctly bilobed, the incisions between lobes rather 
acute ; sides curvate, tapering roundly to hind margin. 

Mesonotum as wide as pronotum, about half its length ; 
caudal margin broadly bilobed ; sides irregularly rounded, 
frequently asymmetrical, often angled and edges a little 
bent up. 

Metanotum a little wider and shorter than mesonotum ; 
caudal margin broadly incurvate ; sides acutely curvate, 
diverging and edges a little bent up. 

Legs yellow-brown. 

Abdomen. — Somewhat polished; with reddish bristles; 
dorsal sclerites with caudal margins black-brown diffusing to 
a dark yellowish-brown ; ventral sclerites paler. 

Minor Soldier. 

Measurements. — Total length 12 to 13 mm. ; head with 
mandibles 6 ram.; head-width 3"2 mm. 

The minor soldier displays most of the characteristics of 
the major soldier, but the head is pale-ochraceous in both 
living and spirit material ; the dorsal outline more rounded 
ovate, and not constricted lateral to antennal fossas ; the 
fontanelle is equall}^ well expressed but not black ; the epi- 
stome is pale and larger ; the frons not so deeply impressed 
and with a very faint median carina; the abdomen is dark 
and banded as in the major but is not so highly chitinised; the 
mandibles are more incurved and more slender, left 2'7 mm., 



OBSEKVATIONS ON SOME SOUTH AFRICAN TERMITES. 465 

right 2*2 mm. ; the left mandible is almost acicular ; the 
antenna are XA^II-jointed, joints III and IV of the worker 
XVIII-jointed fusing to form III of the XVII -jointed form. 

Worker. 

Measnrements. — Total length 7 to 11 mm.; head-width 
3'1 mm. 

There appears to be only one caste, but the workers 
may be gi-aded into three series upon the lengths of their 
abdomens. 

Head. — Dark brown or dark reddish-brown, with clypeo- 
frontal suture black, clypeus pale brown, epistome Avhite, 
labrum-epipharynx yellow-brown. Dorsal outline U-shaped. 
Vertex flatly arched, median region quite flat ; Y-suture 
almost obsolete with paired elongate ocellus-like spots imme- 
diately posterior to sutui-e and towards apices of arms (caudo- 
mediad of antennal fossae). 

Frons with caudal area flat, with a round pit in fork of 
Y-suture in which is a bright yellow spot ; cephalic area much 
depressed posterior to clypeus ; this depression with a faint 
median carina and a few transverse wrinkles. (If a submerged 
specimen is examined, the face of a fox is distinctly pre- 
sented on the frontal area, the pale spot in the pit in the fork 
of the suture representing the nose.) 

Clypeus transversely elliptical, oval and bombous. 

Epistome plate-like, large with distal margin widely 
rounded and with a large median lobe. 

Mandibles diffuse yellow-brown, teeth black. 

Antennas XVIII-jointed; joints II and III sub-ec^ual. 

Thorax and abdomen brownish-black. 

Hab. — Transvaal; Tzaneen, Barberton. 

A nest of this species was examined for the writer at 
Tzaneen by Messrs. A. 0. D. Mogg and W. F. Behrens of 
the Division of Botany. The king and queen together with 
fungus-gardens were collected. The J" measures 13 mm. in 
length, and has a head-width of 3 mm. ;' the female is 100 mm. 
long, and both are characterised by crimson-brown heads ; the 



466 CLAUDE FULLHE. 

antennee of both have been shortened to (king) 13^ (queen) 
13 and 10 joints respectively. 

The nest-site was indicated by a bare patch about which 
were disposed several rough hillocks about 1 ft. high and of 
1 to 2 cubic ft. in bulk. 

Later a second nest was found and examined for me at 
Barberton by Mr, -Kenneth Munro, of the Division of Ento- 
mology. The following note is by Mr. Munro. 

" This nest was found in a private garden in town. The surface indi- 
cations were as follows. There was a cleared oval space some 15 ft. 
long and 6 ft. wide. This was divided into two raised portions. On 
the one half was a low flat mound about 2 ft. high, on the other a 
more conical mound, 3 ft. high, and beside it a smaller subsidiary mound. 

" I dug down beside the higher mound and found the nest-cavity 
3 ft. down. The mound was very sohd, having only two or tlu-ee 
galleries one quarter of an inch in diameter which widened to about 
one inch lower down. 

" As soon as I broke into the cavity, large and small soldiers poured 
out ; the smaller in the majority and no workers were to be seen. 

" The nest was arranged in layers, with the fungus-gardens like those 
of natalensis . The fungus-gardens were much larger and looser than 
those of natalensis. 

" I dug underneath and downward as far as I could, but did not find 
the queen-cell nor any trace of it. The nest seemed to extend a good 
way in all directions with a radius of quite 5 ft. around the hole I made. 

" In the upper part of this mound I found T. incertus. 

" The cavity has a very nauseating smell, very much like the aroma 
of E. trinervius ; I did not notice this with the natalensis nests I 
exposed. 

" This insect is well known to the inhabitants of Barberton and feeds 
upon any wood or dead timber it can obtain. It is said of it that the 
woi'kers and soldiers emerge in droves after dark and forage freely on the 
surface as do Hodotermes." 

Termes waterbergi, sp. n. PL XXXIV, fig. 12. 

Majoe Soldier. 

M e a s u r e m e n t s . — Total length 1 5'5 to 1 8 mm. ; head with 
mandibles 8'5 to 9 mm. ; head-width 4"5 to 5 mm. 

Head. — Pale reddish-brown, ventral gente ochraceous, 
broadly arched ; dully polished ; with short inconspicuous 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 467 

scattoi'ed hairs. Dorsal outline very evenly horseshoe-shaped ;, 
cephalic region much narrowed. 

Fontanelle minute (diameter 11 /u), generally in the apex of 
minute V groove and with a parallel series of short curved 
wrinkles immediately posterior to it. 

Frontal area not deeply depressed ; with a medium trian- 
gular mound, the base of which may extend to the cephalic 
margin of the clypeus or may be interrupted at a wrinkle 
which represents the clypeo-frontal suture ; in the apex of 
this mound, a little anterior to the fontanelle, a black, imper- 
forate, chitinised spot, like an obsolete foramen. 

Eye-spots large, oval, pale, vaguely facetted ; diameters- 
170 X IbSju; surrounded by concentric wrinkles. 
Ocelli quite obsolete. 

Clypeus somewhat convex with a pale plate-like extension 
(the epistome) ; cephalic margin curvate ; sides straight. 

Labrum concolorous with head and with a larg-e white 
cordate lingula. 

Mandibles black throughout ; large (3'3 to 3*4 mm. long) > 
simple ; both incurved and upcurved. Left the more up- 
cui'ved; straighter and broader than right, proximal third 
with a produced piece, the distal end of which very short, 
incurved and pointed ; proximal to this a small conical 
process. Right mandible attenuated ; more incurved than 
left ; distinctly elbowed at half its length and presenting a 
small protuberance on the inner margin a little below the 
elbow. 

Antenna? XVII-, XVIII- or XlX-jointed; the XVIll- 
jointed antennae formed by the failure of either III and IV 
or V and VI to separate. The XVII-jointed through the 
failure of III-VI to separate. The following is an analysis of 
26 iintennse from 13 insects : 
7 v/ith 19 joints. 
11 „ 19 „ V and VI fused but showing pseudo- 
articulation. 
1 „ 18 „ III and IV fused but showing pseudo- 
articulation. 



468 CLAUD K FULLER. 

5 with 18 joints, V and VI fused and showing no pseudo- 
articulation. 

2 „ 17 ,, III, IV, V, VI fused with a pseudo- 
articulation between III and IV. 

Length ranges of joints, when not fused, of four antenna : 

\> I 400 ,1 constant. VI 250-290 /x. XI 320-350 /i. XVI 280-290 fi 

II 180-220 fx. VII 310-340 /«. XII 310-330 /t. XVII 270-280 /. 

111260-280^. VIII 330-360//. XIII 310-320/4. XVIII 260-270;* 

IV 230-270 ». IX 340-360 /i. XIV 310-320 fi. XIX 270-280 ,i 

V 220-250 ^i. X 340-360 /t. XV 290-300 ^l. 

Colour chestnut-red, banded ; I stout, cylindrical and the 
longest, with a few short, stout spines ; II stout, cylindrical 
and the shortest, with a few quite inconspicuous spines; III- 
XVIII attenuated obconic ; III with one whorl of bristles, 
rest with two ; joint XIX oval and comparatively small, as a 
rule as long as III. In the XlX-jointed antenna3, III is 
longer than II or IV, V shorter and narrower than IV and 
much shorter than VI. 

Gula a little darker than ventral genge ; lateral sutures 
brown ; less than one-quarter the width of the head ; sides 
pai-allel, slightly incurvate. Free apex narrowed with distal 
margin straight and oblique incurved sides. 

Ventral genee with post-gense represented by pale ochra- 
■ceous areas. 

Thorax. — Diffuse ochraceous-brown, each shield with a 
I)lack outline ; sides of shields somewhat up-bent. 

Pronotum sellate, narrower than head; median cephalic 
margin bilobed and acutely elevated; cephalo-lateral pro- 
cesses rather acutely rounded; sides straight converging; 
caudal margin broadly bilobed ; caudo-lateral corners broadly 
rounded. 

Mesonotum not quite as wide as pronotum ; sides angled, 
at first curvate and diverging, then converging and straight 
or faintly incurvate and merging broadly into caudal margin ; 
■caudal margin incurvate, nearly straight. 

Metanotum shorter and wider than mesonotum : sides 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 469 

extended, curvate ; sharply angled at caudo-lateral corners ; 
caudal margin straight or faintly incurvate. 

Abdomen. — Polished, diffuse ochraceous-brown. 

Legs ochraceous. 

Minor Soldier. 

Measurements. — Total length 8 to 9 mm.; head with 
mandibles 4 to 4'5 mm.; head-width 2'3 mm. 

Compared to major soldier very small ; head ochraceous ; 
mandibles slender, 2 mm. long ; antennae XVII-jointed ; 
agreeing in other features. 

Major Worker. 

Measurements. — Total length 8 to 9 mm.; head- width 
2'! mm. 

Head. — Reddish-ochraceous, rather pale with a white spot 
in fork of Y-suture and paired ocellus-like spots on caudal 
side of arms of suture as in T. swazise; mandible reddish ; 
abdomen a dark sordid brown. 

This worker is characterised by its distinct, convex, pale 
yellow eye-spots and the peculiar pattern of the antenna?. 

Antennae XIX- j ointed ; joint VI broader than V or VII and 
very conspicuous ; II elongate, cylindrical, constricted at half 
its length and swollen at apex ; joints III and VI are both 
striking; III is elongate cylindrical, constricted at half its 
length and swollen at its apex, it is either nude or with short 
spines ; VI is obconic and very broad, much broader than any 
joint other than I ; V is a nari-ow joint compared to IV and 
VI. 

I the longest (260 fx) broad cylindrical with a whorl of 
short spines. 

II cylindrical, with a whorl of bristles a little shorter than 

I (200 fjL). Ill (as above) as long as II. 

IV short obconic, broad, somewhat globose ; as broad as 

II or broader, not as broad as VI, nearly half the length of 

III (120;,). 

V conspicuously narrow, longer than IV and shorter than 
VI (150^). 



470 CLAUDE FULLER. 

VI as above and of the same length as II, III and A'^II 
(200^). 

VII-XIV elongate, sub-obconic ; about equally wide. 
VIII-XIV sub-equal (250 n). 

XY-XIX shortening faintly and tapering slightly. 
XIX elongate oval. 

Minor Worker. 

Measurements. — Total length 7 mm.; head-width 1'7 mm. 

Head. — Pallid yellow and body generally paler than that 
of major. 

Antennas XVII-jointed ; joints I, II, III as with major 
worker; IV a long joint constricted at half its length Avith 
two whorls of bristles and a well-defined pseudo-articulation, 
into this joint is compressed the characteristic IV and VII 
series of the XlX-jointed major worker antennae. 

Hab. — Transvaal ; Warmbaths (district of the Waterberg), 
De Wildt (district of the Magaliesberg). 

In the material from Warmbaths, collected in October, 
1913, are several large and obese white nymphs measuring 
20 mm. long, also one imago. The imago is somewhat like 
that of 'J \ natal en sis but longer, the measurements are a 
little below those of T. goliath 8jdst. Total length of body 
20 mm. ; head-width 4'5 mm. ; ocelli with a diameter of 40ju ; 
length of insect and Avings, wings folded, 42 mm. ; fore-wing 
36*5 mm. long and 9 mm. wide ; hind wing 34 mm. long and 
9'5 mm. wide. 



Termes natal en sis Haviland. PI. XXXIV, fig, 13; 
PI. XXXIII, fig. 3. 
Termes natalensis ifav., Journ. Linn. Soc. Lond., xxvi, 1898. 
? Termes bellicoaus Smeathmau. Fliil. Trans. Lond.. vol. 71,1781. 

Major Soldier. 
Measurements. — Total length 13 to 18 mm. ; head with 
mandibles 6*5 to 8 mm. ; head-length without mandibles 
4" 5 to 6 mm. ; head-width 3 to 5 mm. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 471 

Head. — Above pale yellow, or pale reddish-yellow, or 
pale reddisli-brown ; cephalic extremity always the darker, 
paler beneath except gula. Vertex with three parnllel 
brown stripes below chitin, one median and two sub-median ; 
median stripe often shortly bifurcate at fontanelle and some- 
times extending- over occiput to occipital foramen ; submedian 
stripes sometimes confluent in frontal area from which they 
diverge and running parallel to the median stripe pass 
over occipital region and traverse the middle line of each 
ventral gena (indicating the obsolete demarcation of the 
post-genge from the genas) ; all three stripes strongly or 
faintly indicated, or only partly indicated; the median 
frequently faint and often abbreviated, only extending from 
fontanelle towards occipital region; submedian stripes 
frequently abbreviated and then only expressed upon occipital 
region ; submedian stripes infrequently branched, forming a 
veined pattern and then upon both dorsal and ventral surfaces. 

Elongate and somewhat flattened ; surface smooth, 
polished, with scattered hairs, dorsal outline horseshoe- 
shaped, broadest in caudal region ; caudal margin broadly 
curvate ; cephalic width a little more than half greatest 
width (but variable) ; sometimes faintly compressed at the 
level of the antennal fossas. 

The rang-e of the measurements made on ten insects from 
three nests is given in the following table : 



Dimensions. 


Nest 1. 


Nest 2. 


Nest 3. 


Gi-eatest widths . 
Cephalic widths . 


4 to 4"4 mm. 
2-3 to 2-8 mm. 


4"3 to 4"5 mm. 

2-4 to 2-8 mm. 


4 to 4'4 mm. 

2-8 to 3 mm. 



Fontanelle always present, often black, often indistinct. 

Frontal area depressed and sloping, features kaleidoscopic. 
Depression bordered by frontal grooves, these grooves origi- 
nate at the caudo-lateral corners of the clypeus, they con- 
verge shortly and then diverge and shallow-out at the level 
of the antennal fossaj (or recurve and embi'ace paired, low, 



472 CLAUDE FULLER. 

oval mounds which lie lateral to the caudal apex of a raised 
median area of the frontal area). Median area of depression 
generally more or less elevated ; elevation more or less trian- 
gular with its apex a little anterior to the fontanelle and its 
base demarcating the caudal extremity of the clypeus ; at the 
base of the triangle, and between the superior articulations 
of the mandible, there are several transverse Avrinkles which 
represent the obsolete clypeo-frontal suture (triangular 
median area often suppressed). Frontal carina black, 
chitinous, widely diverging, crenulate ; forming the lateral 
margins of paired, well-defined, diverging ridges ; these 
ridges originate at the superior articulation of the mandibles 
where they ai'e narrow, mediad of the antennal fossse they 
widen and laterad of the fontanelle merge into the vertex ; 
median slopes of ridges falling more or less abruptly into 
frontal g\*ooves. 

Bye-spots inconspicuous, pale. Ocelli absent. 

Clypeus oblong, wider than long, slightly arched, sides 
straight and parallel ; cephalic margin with a thin plate (the 
epistome) ; faintly curvate or straight Avith slightly oblique, 
cephalo-lateral corners. 

Labrum elongate, tapering, slightly arched ; sides curvate ; 
distal margin of chitinised region more or less incurvate, 
bearing a white cordate epipharyngeal lingula. 

Mandibles black, strong, forcipiform, more or less simple, 
incurved ; apices upbent. Left mandible stouter than right ; 
distal half of cutting margin finely wavy ; proximal half pro- 
duced into a series of irregular processes, one of which is 
particularly prominent and like the blade of a small scalpel. 
Right mandible with cutting margin entire ; base faintly 
wavy with two insignificant sub-conical protuberances. 

Antennae XVII-jointed, dark bi*own-red, ringed. Joint I 
always the longest, II nearly always the shortest, III always 
the second longest joint, V always narrower and shorter 
than IV and VI, generally longer than II. VI-XVI sub- 
equal, or gradually increasing in length from VI to X and 
decreasing from XII-XVI ; XVII always slightly longer and 



OBSEKVATIONS ON SOME SOUTH AFRICAN TERMITES. 47^1 

narrower than XVI. Joint I cylindrical, with an apical whorl' 
of small bristles ; II coarse, sub-clavate, with a whorl of almost 
obsolete bristles ; III elongate clavate, with an intermediate 
and distal whorl of bristles ; IV onwards with one whorl of 
bristles; IV sub-obcoiiic ; V sub-globose; VI-XVI elongate, 
sub-obconic ; XVII short-oval ; apex convex, truncate. 

Cliila elongate, less than half the width of ventral gena ; 
sides parallel and faintly incurvate. Free apex short, nar- 
rowed ; distal margin incurvate, sides straight, oblique. 

Thorax. — Yellowish or yellowish-red; shields frequently 
deeply bordered with smoky brown ; margins slightly up- 
bent, 

Pi'onotum sellate, narrower than head, about half as long- 
as wide ; median part of cephalic margin more or less ele- 
vated and distinctly bilobed, incision notched ; cephalo-lateral 
corners broadly rounded ; sides curvate, tapering roundly into 
caudal margin ; caudal margin broadly bilobed. 

Mesonotum about half as long as pronotum and slightly 
narrower ; caudal margin nearly straight, broadly incurvate ; 
sides parallel, curvate, or with sides angled variously, or 
straight, diverging widely and forming with caudal margin 
distinct caudo-lateral lobes. 

Metanotum a little shorter than pronotum and a little 
wider ; sides more or less acutely curvate ; caudal margin 
sinuate. 

Legs pale brownish-yellow, generally paler than body. 

Abdomen. — Bright yellow and pallid yellowish-brown, 
flattened or arched. 

Minor Soldier. 

Measurements. — Total length 8 to 11 '5 mm. ; head with 
mandibles 4 to 5 mm. ; head-length less mandible 2-4 to 
3"3 mm. ; head-width 2 to 2"5 mm. 

The minor soldier agrees very generally with the major 
soldier, but the head is usually paler, the frontal area less 
depressed and the mandibles proportionately more slender. 

The antennas of the minor soldier, presenting the same 



474 CLAUDE rULLER. 

numbei'. of joints and a similar pattern to tliose of tlie major 
soldier. Joints II and XVII often as long as tliese joints of 
the major soldier antenna. Joint III often sub-equal to II, 
but generall}' longer. Joint V generally shorter than II. 

Major Worker. 

Measurements. — Total length 7 to 9" 5 mm. ; head- width 
2-1 to 2-6 mm. 

Head. — Dark brown or dai'k red-brown, somewhat mar- 
morate ; occipital region and genas paler, stem of Y-suture 
pale, with a bright white or bright pale yellow spot in fork of 
suture ; eye-spots pale, translucent yellow ; clypeo-frontal 
suture darkened; clypeus paler than frons with a median 
brown stripe ; epistome white with brown diffuse stripes. 
Labrum-epipharynx translucent, brown or tinged with green. 

Mandibles at base pale yellow-brown merging thi'ough 
diffuse brown to the black teeth. 

Dorsal outline U-shaped; vertex polished. Frons with 
a small circular pit in fork of Y-suture and with an oblong 
transvei'se depression posterior to clypeus. Clypeus elliptical 
oval ; epistome with distal edge brace-shaped. 

Antennas dark brown to smoky-black, ringed with white ; 
XVII or XVIII-jointed ; in XVIII-jointed form III and IV 
sub-equal and V always larger than VI; in XVII-jointed form 
III is longer than IV and V smaller than VI. 

Thorax. — Sordid, yellow-brown. Pronotuin narrower 
than head ; (quartered by a median and a transverse pale line. 
Mesonotum half the length of pronotum and narrower. 
Metanotum wider than mesonotum, but not longer. 

Legs pallid ; apex of tibia and claws pale red-brown. 

Abdomen. — White, transparent contents visible. 

Minor Worker. 

Measurements. — Total length, 5 to 6 mm. ; head-width, 
1"5 mm. 

Like major worker but paler; with ocellus-like, pale, oval 
spots near cephalo-lateral extremities of arms of Y-suture. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 475 

AnteniitB 17 joints; dark brown ringed with white. Ill 
and V smaller than IV. 

Hab. — Natal; Bellair, Mt. Edgecombe, Pietermaritzburg, 
New Hanover, Weenen, Estcourt, Winterton. Transvaal ; 
Pretoria, Barberton, Tzaneen. Cape ; Kimberley. 

The imago is not being here described, but as a point of 
interest it may be mentioned that in examining the kings it 
was found that in nearly every case the antenna? had been 
reduced to XIII joints and never more than XIV were present. 



Termes bad ins HavUand. PI. XXXIV, fig. 14; 
PI. XXXIII, figs. 1, 2. 

? Termes monodon GerstilcJcer, Jabrb. Hamb. Wiss. Aust., ix, p. 

185, 1891. 
Termes badius Hav.. Journ. Linn. Soc. Lond.. xxvi. p. 385, 1898. 

Soldier Series. 

M e a s nr e m e n t s . — Total len gth 7 to 1 2 mm . ; head with 
mandibles 3'5 to 4*5 mm. Soldiers of one caste only but in 
three or four grades. 

He;id. — Pale, translucent, honey- or reddish-yellow, darker 
towards mandibles ; (spirit material often yellow-brown, or 
reddish-brown or intense black-brown) ; without stripes 
showing through chitin (or with 4 or 5 short dai"k lines ex- 
tending over occipital region, 1 median, 4 sub- median, median 
frequently absent); smooth; shining; somcAvhat thick; broad- 
ovate ; sides curvate, tapering to mandibles, also tapering a 
little towards and merging into caudal margin; broadest at 
about three-quarters the full length of the epicranium. 

Fontanelle indicated by a minute dot, or apparently absent; 
actually always present and always perforate. 

Frontal area faintly impressed, sloping a little and then 
precipitous to cephalic margin of clypeus ; divided into two 
regions which are separated by a low more or less arched 
transverse ridge; appearance otherwise variable and elusive, 

VOL. 3, PART 2. 32 



4i76 CLAUDE FULLER. 

varying- with illumination, sometimes appearing as with a 
median keel. Frontal caringe not strongly elevated. 

Eye-spots very distinct in living insects (distinct in pale 
spirit specimens) ; white or pale yellow with a black centre 
in the form of a tear-drop ; paleness of colour due to thinness 
of chitin ; pigmentation apparently upon muscular tissue. 

Ciypeus very short, precipitous, Avith pale, nearly straight 
margin (epistome), overhanging labrum like a pent-house, with 
several projecting bristles. 

Labrum deflected, triangular, acute ; covering one-third 
the length of the mandibles; apex thin and without a lingula, 
with three superior groups of large bristles, a distal group of 
six and two sub-marginal rows of four each. 

Mandibles black over greater part, bases red or reddish- 
yellow ; strong ; about equal ; elevated, but not up-bent ; 
apices incurved ; each with a strong salient tooth inter- 
mediate between base and apex, tooth on the left being a 
little more distal, larger, and more acute than that on the 
right ; the distal half of the catting margin of each minutely 
and bluntly serrulate. Left mandible with a sinuate, produced 
cutting-margin or blade below tooth, and below that a large 
conical basal protuberance. Right mandible with a minute, 
very acute, persistent, characteristic, and delicate tooth which 
resembles a reversed rose-thorn ; this tooth is placed inter- 
mediate to the large tooth and the base. 

Antennse pale reddish-brown, distal half darker; 2"5 mm. 
long; XVII-jointed (normal) or pseudo XVI-jointed and 
(rarely) with sixteen entire articles. When XVII-jointed, 
I is the longest and XVII nearly as long; III and V are 
short joints placed between the longer joints II, IV, and VI ; 
III is slightly shorter and narrower than V ; IV is always 
shorter than either II or VI, but it is wider than III or V ; 
VI, and often VII, about equal to II ; VIII-XVI sub-equal. 
Joint I cylindrical, robust ; II stout, wider than long, narrower 
than I; III- VI annular, sub-globose; VII-XVI globose, sub- 
obconic; XVII fusiform or elongate-oval. Joints III-XVII 
Avith conspicuous apical whorls of bristles; I and II with incon- 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 477 

spicuous almost obsolete whorls. When pseudo XYI-jointed, 
joints III and IV fused, with a pseudo-articulation. When 
XVI-jointed, joints III and IV of XVII form completely fused. 

Gula much darker than ventral genge ; broad, elongate, 
wider than ventral gente ; sides parallel, curvate, faintly 
sinuate. Free apex short, narrowed, distal margin straight, 
sides diverging and incurvate ; usually with a wide, shallow 
concave depression immediately posterior to the distal margin. 

Thorax. — Tinged with yellow, semi-transparent; pronotum 
paler than head but more coloured than the meso- and meta- 
notum ; meso- and metanotum concoloi'ous with abdomen. 

Pronotum sellate ; cephalic margin bilobed, somewhat con- 
vex and prolonged over head ; cephalo-lateral corners broadly 
rounded ; sides tapering, faintly incurved, merging roundly 
into the iu curvate caudal margin, with a distinct incurvate 
transverse furrow debouching into two depressions ; in each 
depression a low rounded mound ; posterior to furrow are 
paired, convex humps. 

Mesonotum sub-elliptical, more than half the length of 
pronotum and not as wide ; sides parallel, curvate ; caudal 
margin incurvate. 

Metanotum sub-elliptical, shorter and wider than meso- 
notum; sides parallel, curvate; caudal margin straight or 
flatly incurvate. 

Legs pallid. 

Abdomen. — Oval, arched, transparent, slightly tinged 
with yellow ; clothed with yellow-brown bristles. Contents 
of abdomen visible ; contents opaque white and black, 
arranged in a pattern so as to show a black median stripe, 
which is always interrupted on the fifth dorsal sclerite and 
always at its fullest expression on the sixth sclerite. 

Major Worker. 

Measurements. — Total length 4*5 to 7 nun. ; head-width 
1"5 to 1*8 mm. 

Head. — Pale straw-yellow or pale yellow tinged with 
brown; Y-suture with a white spot in fork; superior 



478 



CLAUDE FULLER. 



articulation of mandibles forming- two dark spots ; lateral 
edges and teeth of mandible dark brown ; cephalo-f rontal 
suture a little darker; eyes distinct, with black centres, as 
with soldiers. 

Antennas smoky black, ringed with white ; 19, 18 or 
17 joints; the XVIII-jointed antennas the most fi-equently 
met with, the XlX-jointed appears to be due to the sub-division 
of III of XVIII, the XVII-jointed to the fusing of III and 
IV of XVIII. In the XlX-jointed form. III, IV and V are 
the smallest, V being a little shorter than III or IV. In the 
XVIII-jointed form IV is the smallest and represents the 
unaltered V of the XlX-jointed form. In the XVII-jointed 
form IV is a large joint between two smaller joints. The 
following approximate measurements (in /u) of the three forms 
of antennas from insects of the one nest-series will illustrate 
the reduction in the number of joints.^ 



Joints. 


I. 


II. 


III. 


IV. 


V. 

55 
50 


VI. 

85 
85 


VII. 

95 
95 

75 


VIII. 


JX. 


X. 


XlX-jointed . 
XYIII-jointed . 
XVII-jointed . 


220 
220 
210 


135 
140 
135 


60 


60 


115 


125 


150 
150 
135 


95 
90 


115 125 
105 115 


115 


Joints — (continued). 


XI. 

150 
150 
150 


XII. 


XIII. 


XIV. 


XV. 


XVI. XVII. 


XVIII. 


XIX. 


c 

■ la 


XlX-iointed . 
XVIII-jointed . 
XVII-jointed . 


150 
150 
150 


150 
150 
150 


150 
150 
150 


150 
150 
140 


140^ 140 
140 1 140 
140 130 


140 
140 
130 


175 

175 
175 



Minor Worker. 

Measurements, — Total length 3 to 4*5 mm. ; head-width 
I'l to 1'3 mm. Very much like the major worker ; generally 
with XVII-jointed antennae ; antennge simulating the XVII- 
jointed form of the major worker. 



^ See also introdxactiou to Section Y, p. 419. 



OBSERVATIONS ON SOME SOUTH AFEIOAN TERMITES. 479. 

Hab. — Natal; Pietermaritzburg. Transvaal; Pretoria, 
Alberton, Krugersdorp, Johannesburg, Pietersburg^ Pyramids. 
Orange Free State ; Dealesville. 



Termes latericius Raviland. PI. XXXIV, fig. 15. 

Termes latericius Hav.. Jouru. Linn. Soc. Lond., xxvi, p. 386, 
1898. 

Soldier. 

Measurements. — Total length 5 to 6 mm.; head with 
mandibles 2 5 to 2*6 mm.; head-width 1*4 mm.; mandibles 
1 mm. 

Head. — Pale reddish-yellow or dark red-brown. Ovate 
tapering markedly to mandibles, somewhat roundly arched. 
Fontanelle absent. Frontal area scarcely or not impressed ; 
with an obscured Y-shaped mark indicating the Y- suture. 
Eye-spots absent. 

Clypeus continuous with frons ; distal margin (epistome) pale 
and straight or slightly curvate ; sides straight and parallel or 
slightly diverging. 

Labrum elongate-linguiform ; with three groups of superior 
bristles, apical of 8, sub-marginals of 5 to 6. 

Mandibles with distal two-thirds black, bases reddish; 
apical points sharp and incurved ; cutting margins remark- 
ably tine, entire and blade-like. Left a little broader than 
right, with a sharp tooth below an acute incision and placed 
two-thirds towards the apex ; margin below tooth produced 
as a faintly sinuate, entire blade ; at the base a strong, rounded, 
conical protuberance ; directed forwards. Right with an 
almost obsolete tooth, protuberance salient and opposite to 
that on the left and a minute, acute, conical salient, tooth, 
placed a little distal to the base. 

AntenntB, with 16 or 17 joints, usually with 16; bases 
pallid, distal two-thirds smoky; joints globose or sub-obconic, 
except I and II which are cylindrical. In the XVI- jointed 
form III is usually propoi'tionately large and IV is the 
smallest joint ; in the XVII-jointed form III is the smallest and 



480 CLAUDE FULLER. 

V next. In the XVI-jointed form III frequently shows a 
pseudo-articulation, and its comparatively large size is due to 
the fact that it represents III, IV and V in the XVIII-jointed 
expression found in the worker castes ; the proportion of III 
to IV is not constant because the proliferation of the joint is 
arrested to varying degrees. 

Gula much darker than ventral gen?e ; very broad, nearly 
half the breadth of the head ; sides straight parallel. Free 
apex short, narrowed ; distal margin straight, sides diverging 
and faintly incurvate. 

Thorax. — Transparent, concolorous with abdomen. 

Pronotum sellate ; cephalic mai-gin bilobed ; sides curvate, 
tapering roundly into incurvate hind margin ; cephalo-lateral 
corners rounded. 

Mesonotum sub-elliptical ; shorter and narrower than 
pronotum ; sides curvate, more or less parallel ; caudal margin 
incurvate. 

Metanotum shorter and wider than mesonotum ; sides 
curvate, diverging ; caudal margin straight or slightly incur- 
vate. 

Legs pallid or pale broAvn-yellow. 

Majoe Worker. 

Measurements. — Total length 5'5 to 7*5 mm.; head- 
width 1*7 to 1'9 mm. 

Head. — Pale red-brown ; Y-suture darker, Avith an insig- 
nificant pale spot in fork ; clypeus paler than epicranium, with 
a fine median brown line ; epistome pale yellow-brown ; 
clypeo-frontal suture dark ; labrum-epipharynx pale brown; 
eye-spots insignificant. 

Antennae XVII- or XVIII-jointed, more frequently XVII- 
jointed. In the XVIII-jointed antenna, IV is the smallest 
joint ; in the XVII-jointed, III is the smallest and V next 
because IV and V of the XVIII-jointed form have failed to 
separate ; this failure is often indicated by pseudo-articulation 
in IV of the XVII-jointed form. 

Legs pallid. 



•OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. -iSl 

A b d o m e n. — Transparent. 

Minor Woeker. 
Measurements. — Total length 4to4'7omni. ; liead-witlth 
1'2 mm. 

Similar to 'major worker, antenna XYII- or XVIII-jointed. 

Hab. — Transvaal; Pretoria, Barberton. Natal; Pieter- 
maritzburg, Durban, Scottburgh, Mt. Edgecombe. 

Termes vulgaris Havilmul. PI. XXXIV, fig. 16. 

Termes vulgaris Hav., Journ. Linn. Soc. Loud., vol. xxvi, p. 387, 
1898. 

SOLDIEK. 

Measurements. — Totctl length 6 to 7 nnn. ; head with 
nirindibles 2'5 to 8 mm.; head-width, I'd mm. 

Head. — Reddish-yellow; Y-suture faintly delineated; 
vertex with paired but very faint sub-median stripes. Dorsal 
outline somewhat elongate, broadly oval; roundly yrched ; 
frontal area sloping and somewhat precipitous at clypeus; 
with faint transverse wrinkles. Fontanelle and eye-spots 
absent. 

Clypeus short, insignificant ; cephalic margin pale. 

Labrum linguiform, reaching to half the length of the 
mandibles (more obtuse than that of latericius). 

Mandibles yellow-brown to dark chestnut-brown or black, 
paler at the bases; apical points sharp, incurved. Left 
mandible a little broader than the right, with a salient tooth 
two-thirds towards the apex; cutting margin above and below 
tooth with a moniliform edge, the beads being more prominent 
below the tooth ; at the base a strong, blunt, triangular and 
salient protuberance. Right mandible with a distinct tooth, 
moie intermediate than that on the left and less pronotmced ; 
the cutting margin above and below the tooth is minutely 
irregular; below the tooth towards the base there is a 
distinct curved excision, the proximal end of the excision 



482 CLAUDE FULLEE. 

being produced as a small, salient, thorn-like tooth ; at the 
base a rounded, strong- protuberance. 

Antennas XYII-jointed or pseudo XYI, usually pseudo 
XVI-jointed ; pale, bases pallid, distal two-thirds reddish. 
Joints I and II cylindrical, III and IV irregular, always 
more or less fused, V annular and narrower than IV or VI, 
VI and VII tending- to become obconic, VIII to penultimate, 
short-obconic and only faintly wider than VI and VII, apical 
joint elongate oval. Joint I is the longest (160 /i) but only a 
little longer than the apical joint, II is two-thirds the length 
of I (100 /i), III when separate or fused to IV {30 fi) ; IV 
when separated (40 to 50^), wlien fused always appearing- a 
little longer than III, always wider than III (III and IV 
when fused 70 to 90 fi). V equal in length to IV (40 to 50 /x) 
but always narrower; VI and VII sub-equal (60 to 70 /u) ; 
VIII longer than VII and shorter than IX (90 /m) ; IX to 
penultimate, equal (100^) ; apical joint always half as long 
again as penultimate and equal to II (150/.(). 

On antennal features T. vulgaris differs from the closely 
allied T. latericius insomuch as the Avorker of the latter 
has XVIII-jointed antennae and those of vulgaris XVII- 
jointed. 

Gula a shade darker than the ventral genas and as wide or 
faintly wider; sides parallel. Free apex short, tapering. 

Thorax. — Slightly paler than head. Pronotum sellate^ 
Mesonotum more or less elliptical, sides and caudal margin 
curvate, narrower than pronotum. Metanotum short and 
wide, more or less oblong, wider than mesonotum. 

Legs very pale yellow. 

Abdomen. — Polished; pale yellow (in life the transparent, 
milky-white contents visible) ; ovoid. 

WOEKEK, 

Measurements. — Total length 4*5 to 5 mm.; head-width 
1*5 mm. 

Head. — Pale yellow-brown. Y-suture a little ptiler, with 
an obscure pale spot in fork ; clypeo-frontal suture dark,. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 48o 

almost black ; clypeus with a faint median line ; superior 
articulations of the mandibles appearing as dark red-brown 
spots. Mandibles narrowly outlined and tipped with black. 
Antennae pale reddish. Dorsal outline of head sub-circular ;. 
front with a shallow impression at fork of Y-sutui'e. 

Antennae XVII-jointed, characterised by the insertion of 
the broad joint IV between the small III and Y, otherwise as 
with soldier. 

Thorax. — Pale yellow. Pronotum with two small dark 
spots sub-marginal to cephalo-lateral cornei's. 

Legs pale yellow. 

Abdomen. — Pallid; translucent. 

Hab. — Natal j Mt. Edgecombe, Bellair; Pietermaritzburg. 
Transvaal ; Pretoria. 

Termes incertus Hagev, PI. XXXIV, fig. 17. 

Termes incertus Hagen, Bericht d. Koigl. Preuss. Akad. d. Wiss._ 
Berhn, p. 481, 1853 ; Haviland, Journ. Linn. Soc, xxvi, p. 388^ 
1898. 

Soldier. 

Measurements. — Total length 3"6 to 5 mm. j head with 
mandibles 1*4 to 5 mm. ; head-width 0'7 to 0"8 mm. 

Head. — Vertex pale brownish-yellow, frontal area with a 
whitish-yellow sub-quadi-ate blotch which is usually more or 
less divided by a darker median line and with its caudal 
boundary irregularly W-shaped, the dark colour of the apical 
points of the mandibles and the superior articulations in 
sharp contrast with head. Sub-ovate, tapering to mandibles;, 
somewhat hairy. Frontal area not impressed, somewhat 
precipitous. Fontanelle absent. Eye-spots absent. 

Clypeus broad, front margin incurvate, sides curvate,, 
diverging, more or less lobed. 

Labrum linguiform, elongate, almost lanceolate ; extending 
over two-thirds to three-quarters the length of the mandibles;, 
with an apical group of six conspicuous bristles and paired 
sub-marginal groups of inconspicuous bristles. 



484 CLAUDE FULLEE. 

Mandibles like calipers ; more or less slender with broad 
bases ; apical points up bent and strongly incurved : cutting 
marg'ins simple, entire or with a faint wire-edge. Left 
mandible not larger than right, but with a conspicuous 
basal projection, the distal margin of which is at a right 
angle to the cutting margin and has a small salient tooth 
above it. 

Antennas moniliform ; pallid, translucent ; XlV-jointed, 
characterised by the narrowness and often incomplete separa- 
tion of joints III, IV, and V. Joints I and XIV sub-equal 
(100 to 120 ju) ; II usually the second longest joint (60 to 70^) 
sometimes nearly as long as III + IV + V, sometimes half or 
less than half as long as III + IV -f V (III -h IV -h V 70 to 
130 ju); VI and VII sub-equal (50 to 60 fx) generally a little 
shorter than II ; VIII-XIII sub-equal (60 to 90^) generally 
longer than VI and VII. (When possible to measure III, 
IV, V separately : III (20 to 40 fx), IV (20 to 30 u), V (40 to 
50 ju.) Joints I and II cylindrical, II tending to be obconic ; 
III-IX more or less globose ; IX-XIII globose sub-conical 
(moniliform) ; XIA^ oval, wider than penultimate ; IV the 
narrowest joint, always the shortest; V-VIII gradually increas- 
ing in width. 

Gula broad, as wide as ventral genae. Free apex propor- 
tionately long, narrowed, distal margin straight; sides straight 
-and oblique. 

Thorax. — Concolorous with abdomen, transparent, with 
white and brown body-contents visible. 

Pronotum sellate. Mesonotum much narrower than pro- 
notum. Metanotum broader than pronotum. 

Legs pallid; somewhat compressed ; tibia appearing two- 
segmented and with a patella. 

Abdomen. — Haired; cerci with conspicuous bristles. 

Major Worker. 
Measurements, — Total length 3*5 to 5 mm. ; head-width 
I'O mm. 

H ead .^ — Pale yellow-brown ; haired ; dorsal outline elongate 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 485 

U-shaped ; Y-suture distinct; frontal ai-ea whitish yellow with 
ii pale stripe on genaj; clypeo-frontal suture brown ; superior 
articulations of mandibles dark brown; clypeus yellow; labruni- 
epipharynx yellow or tinged with g-reen ; mandibles yellowish. 

Anteniia3 XIY-jointed^ moniliform ; joints III, IV, V more 
distinctly separated than in soldiers ; apical joint elongate- 
ovate. 

Minor Worker. 

Measurements. — Total length 3'5 to 4*5 mm.; head-width 
0-7 to 0-8 mm. 

The minor workers are much like the major workers and 
have a length-range nearly as great. 

Hab. — Natal; Pietermaritzburg, Mt. Edgecombe, Bellair, 
Winterton, Weenen. Transvaal; Pretoria, Tzaneen, Warm- 
baths, Barbertou. 

EUTERMES. 

Eutermes parvus {Haviland). PI. XXXIV, fig. 18. 
Termes parvus Hav., Journ. Linn. Soc, xxvi, p. 404, 1898. 

Soldier. 

Measurements. — Total length 4 to 5"5 mm. ; head with 
mandibles 2*0 to 2'6 mm. ; head-width 0'8 mm. ; head less 
mandibles 1'6 mm. ; mandibles 0'5 to 0'6 mm. 

Head. — Vertex pale yellow-brown, faintly smoky, outlines 
dark ; frontal area diffuse smoky brown, somewhat maculate. 
Elongate, cylindrical; apex truncate; sides parallel; caudal 
margin rounded and convex ; surface hairy. Fontanelle 
absent. Frontal area sloping and then precipitous; cephalic 
margin between bases of mandibles curvate. Frons convex, 
distinctly bilobed. 

Clypeus produced, tricuspidate, insignificant. 

Labrum short ovate, obtuse-linguiforui, extending to one- 
third the length of the mandibles. 

Mandibles very long and narrow like shear blades; equal; 



486 CLAUDE FULLEK. 

somewhat straight ; points incurved ; taken together much 
narrower than head ; with bases diverging widely to side 
margins of head ; cutting margin very minutely jagged, not 
toothed or serrulate but resembling the " wire-edge " of a 
razor ; left mandible with a delicate, rounded, salient tooth at 
base. 

Antennae pallid; almost moniliforrn ; Xlll-jointed; charac- 
terised by the smallness of III and its incomplete separation 
from IV. I and II cylindrical ; III-VII globose ; VIII-XII 
more or less obconic ; XIII elongate-ovate ; joints widening 
from III onwards. XIII the longest joint and I the second 
longest ; VII to XII longer than II ; VI sub-equal or shorter 
than II ; IV longer than III and a little more than half the 
length of II ; III half the length of II ; III + IV = II. 

Gula, free apex short, narrowed ; distal margin straight, 
sides oblique. Inserted region elongate; widest at cephalic 
end; sides incurvate and tapering almost to occipital foramen,, 
then diverging widely. 

Thorax. — Pronotum sellate ; meso- and metanotum dis- 
tinctly bilobed. 

Legs pallid ; tibia appearing 2-segmented and with a 
patella. 

Abdomen. — Elongate, oval, transparent. 

Worker. 

Measurements. — Total length 3'.5 to 5 mm.; head-width 
0-7 to 1-0 mm. 

Head. — Vertex and clypeus yellowish-white. Frons milky- 
white. Mandibles yellowish-white, finely outlined with dark 
reddish-brown ; superior articulations of mandibles forming 
two dark brown spots. 

Legs transparent ; tibia appearing 2-segmented and with a 
patella. 

Abdomen. — Transparent, contents visible. 

Hab. — Natal ; Durban, Bellair, Scottburgh. Cape Colony;. 
Port Elizabeth. 



OBSERVxVTIONS ON SOME SOUTH AFRICAN TERMITES. 487 

A closely allied but distinct species, making' similar but 
quite subten'anean nests, is common to the Pretoria district, 
Transvaal. 



Eutermes bilobatus [Haviland). PI. XXXIV, %. 19. 
Termes bilobatus Kav., Jonrn. Limi. Soc, Lond., xxvi, p. 411, 1898. 

Soldier. 

Measurements, — Total length 6"5 to 7*5 mm.; head with 
mandibles 3 to 3*5 mm. ; head-width 1'3 to 1*5 mm. 

Head. — Vertex pale, translucent, brownish -yellow ; frons 
paler. Viewed from above short U-shaped, somewhat longer 
than wide ; Y-suture distinct; sides almost parallel ; tapering- 
very faintly to mandibles ; cephalo-dorsal margin curvate. 
Viewed from the side the head is seen to be short and thick, 
the caudal end convex, the frontal area swollen and capistrate; 
below the capistrum the cephalic face is hollowed; a little 
beneath the hood is a thick group of short spines which 
surround the aperture of a gland. E3^e-spots and ocelli 
absent. 

Clypeus short. 

Labrum (so-called) pallid; strongly forked ; stem of fork 
flattened; arms pointed somewhat acutely, sloping upwards ; 
extending to nearly half the length of the mandibles. The 
true labrum is probably atrophied, represented by a thin 
plate superior to a forked, chitinised epipharynx. 

Mandibles long, flat, equal, straight, slender, tapering, 
points slightly incurved ; lateral margins not widening 
greatly at base ; inner margins boldly shouldered at base ; 
cutting margins minutely serratulate. Left mandible with a 
blade-like jagged process on proximal third, which is broadest 
towards its base ; basal process large, salient, conical ; distal 
margin of basal process at right angles to cutting mai'gin and 
lower margin parallel with long axis of mandible. Right 
mandible with an acute process like a reversed rose thorn 
on pi'oximal third ; base like that of left but without process. 



488 CLAUDE FULLER. 

Antennas pallid, slightly suffused with red; 15 joints, or 
with joints III and IV fused, 14 joints. Joint I longest 
(200 n) ; II half as long as I (100 w) ; III, IV, V smallest 
and about equal (50 fx) ; VI-IX increasing in length ; VI 
(80 ,x); VII (100 fx) ; VIII (110 n) ■ IX (140 n)] IX-XI 
about equal (140 ^) ; XII-XIV a little shorter (130 fj) ; XV 
longer than XIV (150 ^u). Joint III narrowest; IV-VII 
increasing in width ; VII-XIV equally wide. Joints III, 
IV, V sub-annular ; VI-XIV elongate obconic ; XV elongate 
oval. 

Grula slightly darker than ventral gence ; sutures brown ; 
elongate ; sides sinuate ; widest towards apex ; constricted 
towards foramen. Free apex very short; distal margin 
straight; sides oblique, continuous with sides of inserted 
portion. 

Thorax. — White, semi-transparent. Pronotum sellate, 
short, narrow, antei-ior part elevated. Mesonotum a little 
shorter and wider than pronotum ; sides diverging ; caudal 
margin broadly curvate. Metanotum shorter than meso- 
notum and wider ; sides diverging ; caudal margin lightly 
ineurvate. 

Legs pallid; tibia appearing 2-segmented and with a 
patella. 

Abdomen. — Transparent, gritty contents visible; ovoid. 

Worker. 

Measurements. — Total length 4*5 to 5 mm. ; head-width 
0"8 mm. 

Head. — Vertex creamy-white ; frons milk-white ; clypeus, 
epistome and labrum creamy-white ; superior articulations of 
mandibles showing as two distinct deep brown spots ; man- 
dibles white with dark brown teeth. Dorsal outline of head 
short U-shaped ; with scattered long pale hairs. 

Clypeus salient, bombous. 

Antennae 14 joints ; as with soldier, but III somewhat large, 
clavate, with a pseudo-articulation and IV the smallest joint. 

Thorax. — White, transparent; much as with soldier. 



OBSERVATIONS ON SOxME SOUTH AFRLCAN TBRMrTES. 4(S9 

Leg's trauspai'ent ; tibia appearing" 2-segineuted and with a 
patella. 

Abdomen. — Transparent, with contents and convolutions 
of viscera visible (when alive a greasy slate-blue) ; elongate- 
ovoid. 

Hab. — Natal; Mt. Edgecombe, Bellair, Pietermaritzburgv 
Freve. Transvaal ; Pretoria. 



Eutermes hasta,tus [Eaviland). PL XXXIV, fig. 20. 

Termes liastatus Hav., Jouni. Linn. Soc Loncl., xxvi, p. 410, 
1898. 

Soldier. 

Measurements. — Total length 4 to 5 mm. ; head with 
mandibles P7 mm. ; head-width PO to PI mm. 

Head. — Pale creamy- white with a median pearly- white 
stripe which is about one-third the width of the head ; colour 
of mandibles in strong contrast. Viewed from above, elongate 
oval ; sides nearly parallel, tapering* to mandibles. Viewed 
from the side ; thick ; roundly arched ; caudal extremity 
convex; frontal area sloping, then precipitous. With 
scattered, long, translucent bristles. 

Labrum obtuse-triangular ; apex with long bristles. 

Mandibles yellowish-red ; falcate ; very much incurved ; 
equal ; similar ; apical points very sharp both with acute 
prominent barbs at half their length. Left mandible with a 
somewhat salient protuberance at base. 

Antennae pseudo XIII- or pseudo XlV-jointed, occasionally 
appearing XV-jointed. Usually pseudo XlV-jointed when 
III is as long as II and nearly twice the length of IV ; III 
composed of III and IV of XV-jointed expression, always 
with a pseudo-articulation and two whorls of hairs. When 
pseudo Xlll-jointed, III as in the XlV-jointed expression, 
with IV longer and larger ; IV composed of IV and V of 
XV-jointed expression with a pseudo-articulation and two 
whorls of hairs. I cylindrical, longer and wider than II ; 
proximal series cylindrical and the same width as II. Distal 



490 CLAUDE FULLER. 

series obconic, increasing in length and width to penultimate 
joint ; penultimate longer than ante-penultimate. Apical 
joint narrow, elongate-fusiform. 

Gula concolorous with ventral genge ; with very short free 
apex. Sides parallel, straight but tapering to apex ; not as 
wide as post-geofe. 

Thorax. — Concoloi-ous with head. Pi'onotum sellate. 
Mesonotum wider than pronotum. Metanotum wider than 
mesonotum. 

Legs pallid, tibia appearing 2-segmented and with a 
patella. 

Abdomen. — Transparent, contents visible ; broad oval. 

Worker. 

Measurements. — Total length 3"5 to 5 mm. ; head-width 
1 mm. 

Head. — Vertex pale creamy-white; frons milk-white; 
superior articulations of mandibles forming dark brown spots ; 
clypeus and labrum creamy-white, mandibles pallid, teeth 
red-brown. 

Antennas pallid, translucent ; 15 joints. III and V small; 
III very small ; otherwise much as with soldier. 

Legs pallid, tibia appearing 2-segmented and Avith a 
patella. 

Abdomen. — Broadly ovate ; transparent, contents visible. 

Hab. — Cape Colony; Port Elizabeth, Stellenbosch district. 

Nest-series of this species have been collected for me by 
several of my colleagues, amongst whom Mr. C. P. van der 
Merwe has sent the largest. In several series there are 
soldiers of an undetermined kind, probably a social species. 
Mr. van de Merwe informs me that a great many of the nests of 
E. hastatus are occupied in part or wholly by the Argentine 
ant (Iridomyrmex humilis Mayr.). This ant has been 
introduced comparatively recently into the Cape, and has 
now acquired a strong foot-hold in the Stellenbosch district. 
It will be interesting to note, as time proceeds, whether 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 491 

I. huuiilis spreads far afield and, as it progresses, annihi- 
lates the termite colonies in the country' it invades. 

The nest of E. has tat us is very much like that of E. 
trinervius, and is constructed of earth particles; the cellular 
spaces are, however, smaller, and no provisions are stored in 
the nest. The queen is active and deposits her eggs in various 
parts of the nest. 

Eutermes trinervius (Ramhur). PI. XXXIII, fig. 4. 
Termes trinervius Bamhio; Hist. Nat Neuvopt., p. 308, 1842. 

Ma.jor Nasutlt. 
Measurements. — Total length 5 to 5'5 mm.; head with 
rostrum, 2'3 to 2"5 mm. ; head-width 1"5 mm. 

Text-fig. IB. 




Baseline of Gena 



Head. — Mahogany-red or aurantiaceous ; very highly 
polished ; rostrum dark brown to black from base or from 
half its length to its apex. Occipital view balloon-like, out- 
line almost circular. Dorsal view turbinate, vertex sometimes 
with a shallow median depression. Ventral surface flat. 
Viewed in profile the epicranium is bombous, short, thick ; 
the genae faintly incurvate, almost straight ; the cephalic 
margin below the rostrum oblique ; the rostrum is attenuated, 
cylindrical, its apex perforated, and with its long axis parallel 
to the base-line of the genae ; from the apex of the rostrum to 
its oblic(ue base (e.g. a point above the antennal foveolae) it 
is shorter than the rest of the head-length, sometimes as much 
as 5 to 6. The superior margin of the apex of the rostrum 

VOL. 8, PART 2. 33 



492 CLAUDE FULLER. 

is slightly deflected. The rostrum has the appearance of 
being elevated above the vertex of the epicraniuni, but this 
is not actually the case. Y-suture, fontanelle, eyes, ocelli 
absent. 

Clypevis, labrum, mandibles, insignificaut. 

Mandibles rudimentary ; plate-like and tinangular with 
broad bases ; length to breadth 5 : 4. 

Antennae yellowish-red ; 14 joints ; I the longest joint ; 
cylindrical ; length constant (200 fi) ; II about half the length 
of I ; cylindrical ; the shortest joint (100 to 110 ju) ; III always 
longer than IV (140 to 180 /u) ; IV equal to or shorter than 
V (120 to 150 ;x) ; V shorter than VI (120 to 150 ,i); VI and 
VII sub-equal (150 to 170 ^u) ; VIII, IX sub-equal, generally 
a little longer than VII (160 to 170 ;x) ; X generally a little 
shorter than VIII and IX (150 to 170^); XI, XII, XIII, 
XIV, decreasing; XII and XIII (180 to 160 |u) ; XIV (130 to 
140 |u). 

Gula minute, quadrate. 

Thorax. — Small, much narrowed; yellow or red-brown. 

Pronotum very short ; with deep transverse incision ; 
cephalic and caudal margins elevated ; caudal margin broadly 
rounded and notched. Mesonotum sub-elliptical, narrower 
than pronotum. Metanotum broader than pronotum, very 
short ; sides very oblique, diverging. 

Legs ample ; pale red-yellow ; tibia appearing 2-segmented 
and with a patella. 

Abdomen. — Elongate-oval ; dorsum very arched; sclerites 
diffuse yellow and brown ; conjunctiva forming marked white 
bands. 

Minor Nasutu. 

Measurements. — Total length 4 to 4*7 mm. ; head with 
rostrum 17 to 2'1 mm. ; head-width 0*7 to 1*0 mm. 

Similar to major nasutu, but more slender ; head less 
bombous ; rostrum proportionately longer ; antenme as with 
major nasutu. 



OBSERVATIONS ON SOME SOUTH AFlilCAN TERMITES. 498 

Major Worker. 

Measufemeats. — Total length 4"5 to 5 inui. ; head-width 
1*5 to 1*7 inm. 

Head. — With a bright, white, cruciform symbol, involvino- 
the Y-suture and having its median line extended to the 
clypeus ; at the intersection a brighter white, fusiform flare 
radiating over the frons. On the vertex immediately lateral 
to the stem of the cross two sub-quadrate brown patches 
bordered by the very pale yellow geua3; mediad of each 
patch a pale sinuate line. Clypeus and epistome pale yellow 
or translucent and tinged with green. Mandibles very pale 
yellow, outlined with dark brown; teeth dark brown to black. 
Dorsal outline of head short, open U-shaped, widest across 
cephalic margin; genas flattened and extended lateral to 
antennal fossae ; vertex quite flat ; clypeus bombous. Labrum 
almost, if not quite complete. 

Antennae XV-jointed ; the extra joint due to proliferation 
of III in the XlV-jointed fixed soldier pattern ; the length of 
III in the XV-jointed worker antennge is variable in accor- 
dance with how much it has enlarged after proliferation ; 
it is either a very long almost clavate joint nearly twice 
as long as II or IV, or it is equal to II and IV or a trifle 
longer. 

Thorax.— Pallid. 

Legs pallid ; tarsi appearing 2-segmented and with a patella. 

Abdomen. — Very arched; translucent with almost chai*- 
acteristic brown and white convolutions of viscera visible. 

Minor Worker. 
Measurements. — Total length 3*8 to 4*5 mm.; head- 
width 1*0 mm. 

Resembling major worker, but head markings generally 
paler. 

Hab. — Cape Colony; Elsenberg, Mossel Bay, Port Eliza- 
beth, Middelburg, Kimberley. Orange Free State, Transvaal 
and Natal — general. 

VOL. 3, PART 2. 33§ 



494 CLAUDE FULLER. 

VI. APPENDIX. 

Significance of Terms generally used in Descriptions. 

Anchylosed. — G-rown together at a joint. 

Antennal carinse. — The ridges or brows superior to the antemial 
fossae. 

Antennal foveolaj. — The pits in which the antenna; are inserted, 
as a feature apart from the antennal fossae. 

Apical points. — The piercing tips of the mandibles of soldier 
termites. 

Arched. — Applied to a curvate margin which rises more or less per- 
pendicular to the plane of the body. 

Aureole. — A ring of colour which is usually diffuse outwardly. 

Barb. — (Restricted). As the barb of a fish-hook. 

Bi-curvate. — With an in-and-out curve (sinuate). 

B i-lobe d. — (As applied to margins) . With two rounded processes, not 
necessarily thickened. 

Brace-shaped. — Shaped like the brace or bracket (/ — '^ — \) of 
printers. (Ornithological.) 

Capistrate. — Hooded or cowled. 

Capistrum. — A hood or cowl, like that of certain birds. 

Caudal, Caudad, Caudo. — Pertaining to the anal extremity and 
more generally the direction (see Cej)halic). 

Cephalic, Cephalad, Cephalo. — Belonging or attached to the 
head and more generally the direction indicated by a line drawn from 
the centre of the animal to the head, but at an indefinite distance in 
that direction. Used in refen'ing to both head and tail (Comstock and 
Kellogg). 

Clypeo-frontal suture. — The suture separating the clypeus from 
the frons. 

Concave. — Hollowed out, as opposed to convex. 

Cordate. — Heart-shaped in outline. 

Cordiform. — Heai-t-shaped in outline and convex above. 

Convex. — The outer curved surface of a segment of a sphere. 

Curvate. — Curved. 

Deflected. — Bent downwai'd, sloping. 

Diffuse. — Spreading out, without distinct edge or margin. 

Distal, Distad. — Toward or at the extremity used in reference to 
appendages or more or less free parts to indicate the end or nearness to 
the end which is free ; opposites ; proximal ; proximad. 

Epistome. — (Restricted). A distinct piece behind or above the 
labrum or a plate-like extension of the clypeus. In the latter sense 
always bracketed thus (epistome). 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 495 

Fimbriate. — Set with a fringe of hair closely placed. 
Foramen. — Applied to gland orifices of the head of certain soldier 
termites which are situated elsewhere than where the fontanelle is. 

Fontanelle. — Particularly applied to a small gland orifice of the 
epicranium of certain soldier termites in which the Y-suture is absent 
and found at a spot which coincides with the fork of the suture in other 
soldier termites. (The fenestra of Haviland.) See Foramen. 

Frons. — = Front; the anterior portion of the head cephalic of the 
Y-suture (see Frontal area). 

Frontal area. — Used as an inclusive term to describe the front 
region of the head (frons and clypeus) when the Y-shaped suture 
and the clypeo-frontal suture are absent. 
F u r c a t e. — ^Forked. 

Gen 86. — -The whole of the sides of the head from the cephalic ex- 
tremity to the occipital region. 

Incurvate. — Bowed, or curved inwards. 

Insignificant. — (Restricted). Not essential to the diagnosis. 
Intermediate. — (Restricted). To indicate the point or region at 
oi" about half way between apex and base of an appendage, or generally 
the second member of a series of three similar parts. 
Intro se. — Directed inward toward the body. 

Labrum. — Used in the proper sense for the unsegmented flap-like 

sclerite articulated to the cephalic margin of the clypeus (see Labrum- 

epipharynx), or loosely for the labrum-epipharynx or labrum + lingula. 

Labrum-epipharynx. — The long articulated labrum of the workers 

of many species. 

Lingula. — A cordate prolongation of the epipharynx which is white 
and not chitinised, and the dorsal surface of which is continuous 
with that of the labrum (found in the soldier termites of the group 
represented by T. natalensis). 

Obsolete. — Inconspicuous or apparently absent. 
Occipital region. —An indefinite area forming the convex caudal 
extremity of the head. 

P a t e 1 1 a. — An apparent segmentation of the tibia of certain species, 
resembling the patella of the legs of spiders. 

Pennant stripes. — Narrow, tapering, and somewhat long stripes 
like the pennant of a war- vessel. 

Proximal, Proxima d. — Used in reference to appendages (mandibles, 
autennaj. legs, etc.) and more or less free parts (clypeus, gula) to 
indicate nearness to the end which is attached to or inserted into the 
body. 

Salient.— (Restricted). Jutting out more or less conspicuously. 
Scallop. — (Restricted). A small, obtuse, and rather roundly 
curved tooth. 



496 OHAUDE FULF-EK. 

Scoop-shaped. — In the form of a shovel with a shoi't, broad handle- 
and with upturned sides to its blade. 

Sellate.— Saddle-shaped, used in the restricted sense of the peculiar 
broad and convex shape of the pronotum of termites. 

Serratulate. — With little teeth or serrations. 

Side-plates. — Thin lateral extensions of the pro-, meso-, and meta- 
notuni. 

Sinuate. — Applied to lines and margins with an in-and-out curve. 

Sub-reniform. — Almost reniform or irregularly oval. 

Tricurvate. — A line or margin with two out and one in curve or 
two in and one out curve. 

Trochantin of mandible. — A narrow, and often salient, sclerite 
between mandible and gena. 

Turbinate. — -Top-shaped, or napiform (Botanical). 

U-shaped. — Having the outline of the letter U, applied to the heads 
of many termites, although the arms are always a little in-bent at their 
apices. 

Ventral genge. — The two cheeks lateral to the gula composed of 
the genfeand post-gense, but with the line of demarcation qviite obsolete. 

Wavy. — Sinuate, with several in-and-out curves. 

Whorl. — A ring of long hair, or hair-like bristles, arranged around 
a centre like the spokes around the hub of a wheel. 

Y-sviture. — The Y-shaped suture or the eijicranial sutu.re which 
divides the vertex into two sclerites, the arms separating the vertex 
from the frons. 



VII. REFERENCES. 

(la) Haviland. G. D. — " Obsei-vations on Termites," " Journal of the 
Linnean Society,' vol. xxvi, 1898. 

(lb) Warren, Ernest. — " Notes on the Life-IIistories of Natal Ter- 
mites based on the observations of the late Gr. D. Haviland," ' Annals of 
the Natal Museum," vol. ii, Pt. 1, 1909. 

(2) Sjostedt, Y. — ' Monograph of African Termites * and Supplement, 
1901. 

(3) Fuller, C— " White Ants in Natal," ' Agricultural Journal of the 
Union of South Africa,' 1912. 

(4) Desneux, J. — " Family Termitida?," ' Genera Insectorum,' 1904. 

(5) Comstock and Kellog. — ' Elements of Insect Anatomy,' New York, 
1902. 

(6) Comstock and Chujiro Kochi. — " The Skeleton of the Head of 
Insects," ' The American Naturalist,' vol. xxxvi, No. 421, 1902. 

(7) Smith, J. B.— ' Glossary of Entomology,' New York, 1906. 



OBSERVATIONS ON SOxME SOUTH AFIUCAN TERMITE.S. 497 



EXPLANATION OF PLATES XXV-XXXV, 

Illustrating Mr. Claude Fuller's paper, " Observations on 
Some South African Termites." 



PLATE XXV. 
Calotermes durbanensis Hav. 
Fig. 1. — X 5. Tunnellings and chambers in dead wood, in vertical 
■sections at right angles to each other. 

Hodotermes transvaalensis sjj. h. 
Fig. 2. — x -j\j. Hive-cavity with shelving removed. 
Fig. 3. — x f. Fragment of wall of hive-cavity. 
Fig. 4. — x 2. A separated carton column for supporting shelving. 
Figs. 5 and 6. — Nat. size. Fragments of shelving showing inclines 
and columns. 

Fig. 7. — Nat. size. Soil fragment perforated by a foraging gallery. 

Termes swazias sjj. n. 
Fig. 8. — x h. Large fungus-garden (120 mm. across) viewed from 
the side. 

PLATE XXVI. 
Hodotermes transvaalensis sp. n. 

Fig. 1. — X yV. Vertical section of hive-cavity with two vertical 
shafts. 

Figs. 2, 2a. — X ^V. Long and cross vertical sections of a large 
gallery-pocket. 

Fig. 3. — X j^. Vertical section of pile of grass-lengths weighted 
down with soil, covering mouth of gallery. 

Fig. 4.— X }. Moundlet of earth particles. 

Figs. 4a, 46, 4c. — x j. Cores or tubes of moundlets like fig. 4. 

Fig. 5. — x y^. Moundlet with simple core partly exposed by wind. 

Fig. 5rt. — x Jg. Core of moinidlet shown in fig. 5. 

Hodotermes viator [Latreille). 

Figs. 6, '6a, 6b. — x |. Vertical sections of moundlets of excreta 
pai'ticles with tubular extensions of gallery (F. W. Petty). 

Fig. 7. — x ^. Vertical section of chambered moundlet covered with 
l^ieces of vegetation (F. W. Petty). 



498 CLAUDE FULLEK. 

Termes uatalensis Haviland. 

Fig. 8. — x g\;. Low, rounded, recent mound ; Pretoria. 

Fig. 9. — Rough, conical mound, with herbage ; Pretoria (compare 
with figs. 3 and 4, PL XXXII). 

Fig. 10. — Acutely conical and recent mound, showing recent and 
uniform extension of base from weathering ; Mount Edgecombe. 

Fig. 11. — Rounded and recent mound on hillside, showing recent 
extension of lower base from weathering ; Mount Edgecombe. 

Fig. 12. — Vertical section of domed and recent mound showing 
relative position of hive-cavity and galleries ; in dense scrub ; Mount 
Edgecombe. 

Figs. 13 and 14. — Vertical sections of mounds showing shouldered 
sides; higher altitudes in Natal. 

Fig. 15. — Nest-sites or oases with fringe of coast bush in back- 
ground ; sketched at Mount Edgecombe, Natal. 

Fig. 16. — A series of nest-sites on gentle slope and to one side of 
ridge ; sketched at Mount Edgecombe, Natal. 

Fig. 17. — Vertical section of large nest-site (marked x fig. 16) ; show- 
ing position of hive-cavity. 

Fig. 18. — In vertical section, a stone-flagged, earth-filled veranda 
with nest-cavity immediately below cement reinforcement; with 
exterior dump-heap and one beneath flooring of interior of house (T . 
badius nests under similar conditions). 

Fig. 19. — x ^. A break in the weather-worn surface of a mound 
in course of repair, showing the apertures through which the earth- 
pellets are placed outside ; these apertures are subsequently closed. 
Sketched from nature. 



PLATE XXVII. 

Termes waterbergi sp. n. 

Fig. 1. — X tj'tt- Mound on periphery of nest-site (oasis of park 
formation). 

Fig. 2. — X gV- Section of mound shown in fig. 1 to illustrate the 
absence of vertical permanent shafts and showing roots in undisturbed 
soil, but not in mound. 

Fig. 3. — x ^g. View of large cavity in half-section underlying site 
of mound. Photograph olitained by directing camera downwards at 
an angle of 45°. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 499 

PLATE XXVIII. 

Termes natalensis Hav. 

Fig. 1. — X 3TJ. Large mound, lateral galleries cut through by trench. 

Fig. 2. — x xV- Vertical section of large and unusually hollowed 
mound, Pretoria. (Photo by F. Thomsen.) 

Figs. 3 and 4. — Nat, size. Small queen-cell, without brackets, and 
with large entrances. 

Figs. 5 and 6. — x \. Large queen-cell ; fig. 5, exterior view ; fig. 6, 
vertical long-section. Cavity 125 mm. long and 18 mm. high. 

Figs. 7 and 8. — x i. A large queen-cell ; fig. 7, exterior view ; fig. 
8, vertical cross-section. Cavity 90 mm. broad and 18 mm. high. 

Figs. 9-12. — x f. Coronetted fungus-gardens; fig. 9, tall form 
(50 mm. high, 80 mm. across) ; fig. 10, short form (25 mm. high, 95 mm. 
across) ; fig. 11 short form from above ; fig. 12, short form from below. 



PLATE XXIX. 
Termes latericius Haviland. 

Fig. 1. — Vertical section of nest-site (diagrammatic) showing 
(a, b, c, d) various types of air-pits, (e, e) supplementary fungus-garden 
cavities ; and general relation of hive-cavity to air-pits. 

Fig. 2. — x i. Vei'tical section of roofed air-pit (sketched from 
nature, Pretoria). 

Fig. 3. — X f . Vertical cross-section of a main-runway. 

Fig. 4. — x f. Vertical section of a main-runway, showing inclined 
vertical shaft to soil-surface. 

Fig. 5. — x 1. Vertical section of small granary-cavity, with simple 
horizontal diaphragm. 

Fig. 6.— X |. Vertical section of small granary-cavity, with simple 
sub-horizontal diaphragm, showing : a, single entrance ; b, a very small 
fungus-garden ; c, a small collection of fresh grass-lengths deposited on 
floor of upper chamber. 

Fig. 7. — x i. Vertical section of medium large granary-cavity, 
showing conspicuous upper cavity, complex interior of lower region, 
and grass seeds embedded in surrounding eai'th (from nature, Pretoria). 

Fig. 8. — x i. Exit galleries made for imagos ; a, front view of van- 
tage ; b, vertical longitudinal section ; c, vertical section showing con- 
dition preceding and following flight ; d, cross-section of gallery two 
inches from surface. 



500 CLAUDE FULLKR. 

Termes incertus Hagen. 
Fig. 9. — A twelve incli length of a down-gallery. For explanation see 
letter-press, p. 394. 

Fig. 10. — X ^. The extensions made when galleries are severed. 

Eutermes parvus (Haviland). 

Fig. 11. — X 5ij. Globular nest overhanging sea-beach at Scottburgh, 
Natal. 

Fig. 12. — a, b, c. Various types of nests. 

Eutermes trinerviiis (Bamhur). 
Fig. 13. — x g-. Vertical section of mound with a lateral gallery 
passing under wheel ti'ack, showing how gallery follows contour of 
siiiface. 

Fig. 14. — x ^. Vertical cross-section of lateral gallery. 

Fig. 15. — x ^. Vertical section of a galleiy and side-pouch. 

Fig. 15«. — x ^. Vertical cross-section of gallery, with opposite 
side-pouches not anastomosing. 

Fig 15&. — x ^. Vertical cross-section of gallery, with opposite 
side-pouches anastomosing. 

Fig. 15c. — x h. A side view of 156. 

Fig. 15d. — x h. Apertiu-es of side-pouches on opposite sides of 
pathway, viewed from above. 

Fig. 16. — x ~. Deserted mound showing apertures on surface and 
radiating series in soil-surface. 

Fig. 16fl. — x i. Vertical section of lateral gallery showing how 
pathway is elevated towards aperture in soil- siai'f ace. 

Fig. 16b. — x h. Aperture as seen when looked into from above. 

Fig. 17. — X^Vi • Bizarre type of mounds ; Benoni, Transvaal. 

PLATE XXX. 

Termes badius Haviland. 

Fig. 1. — X ■j'g. Modern nest, in vertical section and perspective, 
showing trees crusted by tei'mites with clay ; small surface moundlets 
and descending shafts ; great cavity and supplementary cavities filled 
with fungus-garden ; queen-cell attached to wall of cavity (left side) 
and great radiating galleries. (Somewhat diagrammatic.) 

Fig. 2. — x i. A queen-cell from modern nest showing pedestals 
on which it stands ; its rough outer surface with stones attached. 
Total length 170-180 mm. 



OBSEliVATJONS ON SOME SOUTH AFRICAN TERMITES. 501 

Fig. 3. — x i. A queen-cell fi-om a modern nest showing in cross- 
section highly arched cavity, characteristic thin floor and pedestal. 
Width of cavity 85 mm., height 32 mm. 

Fig. 4. — x }. A very large cell, with more or less regular exterior, 
showing where clay arms were attached, and entrance holes. Total 
length about 260 mm. 

Fig. 5. — x J. A three-quarter view of fig. 4, showing cavity and 
pedestals. 

Fig. 6. — Nat. size. Portion of upper surface of great fungus-garden. 

Termes vulgaris Haviland. 
Fig. 7. — Nat. size. Portion of outer surface of great fungus-garden. 
Fig. 8. — x to- "Vertical section of nest described in text, p. 391. 
Fig. 9. — x 1. Great loaf-like fungvis-garden from nest illustrated 
in fig. 8. 

Fig. 10. — X i. Fungus-garden, in section, showing layer formation. 

Termes latericius Haviland. 
Fig. 11. — X 4. — A typical supplementary fungus-garden. Length 
110 mm., height 50 mm. 



PLATE XXXI. 
Eutermes parvus {Haviland). 
Fig. 1. — X ^. Nest viewed from above; the margin represents 
ground level. Diameter 160 mm. 

Fig. 2. — x i. Horizontal section of lower apex of nest, ex^wsing 
queen-cell. (Photo, by Dr. Conrad Akerman.) 

Fig. 3. — X 4. Vertical section of nest showing the queen-cell in 
base; the ground level is represented by a dotted line. (Photo, by 
Dr. Conrad Akerman.) 

Termes incertus Haviland. 

Fig. 4. — Nat. size. Earth fragment showing a very small fungus- 
garden cavity and the young fungus-garden. 

Fig. 5. — x f. A large fungus-garden viewed from above, 70 mm. 
across. 

Termes latericius Haviland. 
Fig. 6. — x j\,. Nest site with one particularly high chimney at 
Pienaar's River, Transvaal. Chimney 2-3 ft. (Photo by F. Thomsen.) 



502 CLAUDE FULLER. 

Eutermes bilobatus (Haviland). 

Fig. 7. — x i. Common type of weather-worn mound associated 
with Eutermes trinervins mounds (140 mm. broad). 

Fig. 8. — x i. Abnormal and recent conical mound, not weather- 
worn. (Photo by F. Thomsen.) 

Fig. 9. — x i. Yertical section of fig. 8, showing typical cellular 
structure. (Photo by F. Thomsen.) 



PLATE XXXII. 

Eutermes trinervius (Ramhur). 

Fig. 1. — x To- "Vertical section of mound. (Photo by F. Thomsen.) 

Fig. 2. — x -^. Mound beax'ing Podaxon sp. (Photo by F. 
Thomsen.) 

Fig. 3. — x ^V. Termes sp. Mound with Cocculus villosus J' 
growing on apex. Koodoos River, Zoutpansberg, Transvaal. (Photo by 
A. O. D. Mogg.) 

Fig. 4. — x aV- Termes natalensis Haviland. Mound in low- 
veld overgrown by bush. (Photo Ijy F. Thomsen.) 



PLATE XXXIII. 

Fig. 1. — Termes badius. Antennal chart (solid black) showing 
the range of variation for each joint of the XlX-jointed imago in ten 
individuals, and chart (double line) of one XlX-jointed worker antenna. 

Fig. 2. — Termes badius. Antennal charts for the XlX-jointed 
soldier, worker major and worker minor. 

Fig. 3. — Termes natalensis. Antennal charts of the major and 
minor soldiers. 

Fig. 4. — Eutermes trinervius. Antennal chart of the major 
nasutu. 

Fig. 5. — Hodotermes spp. Diagram to illustrate the ranges of 
the measurements given for the described species of Hodotermes. 
a. Total length ranges for soldiers, a'. Head with mandible ranges for 
soldiers, a^. Head-width ranges for soldiers, h. Total length-ranges 
for workers. 6'. Head-width ranges for workers. 

Note. — In the case of all five diagrams the figures are based on the 
measurement of at least ten individuals except when otherwise stated. 



OBSERVATIONS ON SOME SOUTH AFRICAN TERMITES. 503 

PLATE XXXIV. 
Hodotermes transvaalensis sj). n. 
Fig. 1. — X 9. Mandibles of soldier. 1«. — x 6. Gula and ventral 
aspect of head. 

Hodotermes pre tori ens is s^. n. 
Fig. 2. — x 9. Mandibles of soldier. 2a. — x 6. Gula and ventral 
aspect of head. 

Hodotermes karrooensis sp. «. 
Fig. 3.— X 9. Mandibles of soldier. 3a.— x 6. Gula and ventral 
aspect of head. 

Hodotermes mossambicus (Hagen). 
Fig. 4.— X 9. Mandibles of soldier. 4a. — x 6. Gula and ventral 
aspect of head. 

Hodotermes havilandi Sharp. 
Fig. 5. — x 9. Mandibles of soldier. 5rt. — x 6. Gula and ventral 
aspect of head. 

Hodotermes warreni sp. n. 
Fjg. 6. — X 9. Mandibles of soldier. 6a. — X 6. Gula and ventral 
aspect of head. 

Hodotermes braini sp. n. 
Fig. 7.— X 9. Mandibles of soldier. 7a. — x 6. Gula and ventral 
aspect of head. 

Hodotermes viator (Latrielle). 
Fig. 8.— X 9. Mandibles of soldier. 8a.— X 6. Gula and ventral 
aspect of head. 

Calotermes durbanensis Haviland. 
Fig. 9.— x 33. Mandibles of soldier. 

Rhinotermes putorius Sjostedt. 
Fig. 10. — X 33. Mandibles of major soldier. 

Termes swazise sp. n. 
Fig. 11, — X 9. Mandibles of major and minor soldier. 

Termes waterbergi sp. n. 
Fig. 12. — x 9. Mandibles of major and minor soldier. 



504 CLAUDE FULLER. 

Termes natalensis Haviland. 
Fig. 13. — x 9. Mandibles of major and minor soldier. 13a. 
Thorax and body showing rounded mesothorax. 136. Thorax and body 
showing lobed or skirted mesothorax. 13o and 13b x 3"5. 

Termes badius Haviland. 
Fig. 1-i.— X 33. Mandibles of soldier. 

Termes latericius Haviland. 
Fig. 15.— x 33. Mandibles of soldier. 

Termes vulgaris Haviland. 
Fig. 16.— x 33. Mandibles of soldier. 

Termes incertus JTa^en. 
Fig. 17. — x 33. Mandibles of soldier. 

Eutermes parvus (Haviland). 
Fig. 18.— X 33. Mandibles of soldier. 

Eutermes bilobatus (Haviland). 
Fig. 19.— X 33. Mandibles of soldier. 

Eutermes h a s t a t u s {Haviland) . 
Fig. 20.— X 33. Mandibles of soldier. 



PLATE XXXV. 

Labrum of soldier of (1) Hodotermes transvaalensis sp. n., 
.(2) pretoriensis sj). »., (3) karrooensis sp. n., (4) mossambicus 
.{Hagen), (5) havilandi Sharp, (6) warreni sp. n., (7) braini sp. n., 
.(8) viator (Latreille) . 1 to 8 — x 28. 



,\js. Natal Mrs., Vol.. IH. 












t'ALOTUKM KS D U K B A N K N S 1 S. Fl"i. 1. H n DOT K K M Ks T R A N S V A A L E N SI S . Fl(J 



Ti;itMEs swAzr.K. Fic s. 



Ann. Nat. Mus., Yol. III. 



Pl. XXVJI. 




TeRMES WATERBEKGI SJ;3. //. X 3\ 



PAo/o. Chiude Fuller. 



Adlard 4' •S'o)?, Inijir. 



Ann. Natai. Mcs., Vol. III. 



*■ .»v 








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Ann. Natai, Mus., V,n.. III. 



I'L. xx^a. 




EfTERMES PARVUS. FlO.S, 1-3. TeuMES INOEKTUS. FlOS, 4-5. T. LATEBICIUS. FiG. ti. E. BlhOBATUS. Fl 



Aitlayd * Soo, Im 



VxN. Natai. Mlis., Vol. III. 



I'l,. XXXII. 





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1 



A FURTHER NOTE ON HYBRID COCKATOOS. 



505 



A Further Note on Hybrid Cockatoos. 

By 
Ernest IkVaiToii, D.Sc.Lond. 



In 1914 I described in this journal^ a case of hybridism 
between the male of Cacatua gelerita {Lath.) and the 
female of Licmetis n a sic a (Temm.) There are now a few 
additional particulars to be given. 

During the last season three eggs were laid ; of these one 
duly hatched, Avhile one was addled, and one was broken. 
The young bird appeared quite healthy and grew rapidly, 
but unfortunately it died when about three months old. 
Subsequent examination suggested that the cause of death was 
peritonitis. Mrs. Brown, who owns the cockatoos, very kindly 
presented the bird to the Natal Museum, and it has now been 
mounted. The growth of the bird appeared to be exceptionally 
rapid, since at the time of death the standing length was not 
less than in the two living hybrids. 

The time between the laying of the first egg and the com- 
mencement of sitting varied from five to six days. In 
the former account it was stated that probably the period of 
incubation was about twenty-one days ; but Mrs. Brown 
informs me that a special note had been taken of the dates of 



Hybrid. 



Date of commence- 
ment of sitting. 



Sept. 23rd, 1910 
Aug. 15th, 1913 
Sept. 8th, 1914 



i 

Date of hatching, i Period. 



Oct. 20th, 1910 
Sept. 11th, 1913 
Oct. 4th, 1914 



27 days 
27 „ 
26 „ 



Approximate age 
at Dec. 31st, 1914, 



50 months 
15 
3 ., 



' ' Annals of the Natal Museum,' vol. iii, pt. 1, p. 7, 1914. 



506 ERNEST WARREN. 

the commencement of sitting and of hatching in the case of 
all three hybrids. These dates are now given, and it will be 
■seen that the period of incubation varied from twenty-seven 
to twenty-six days. 

As I am unaware of the normal incubation periods of 
Cacatua galerita and Licmetis na sic a it is not possible 
to make an interesting comparison as to whether hybridisa- 
tion affects this period. It may be noticed that there is a 
difference of only about twenty-four hours in the incubation 
period of the three hybrids. It also must be noted that out 
oi the two eggs (omitting the broken one) only one hatched, 
■:and in conjunction with the results on the former occasions 
we must regard the large percentage of failure in incubation 
as being due to the hybridisation. 

At the time of writing the account of the first hybrid, the 
second hybrid was only three months old, and it did not 
seem to be fully grown. Owing to these circumstances 
details of the measureinents were not given. 

The second hybrid is now fifteen months old, and the bird 
which died was three months old. It is interesting to ascer- 
tain how far the brother hybrids resemble one another, and 
for this purpose comparative tables are given for the non- 
measurable and for the measured characters. 

From these two tables it will be at once clear thatalthongh 
the hybrids bear a sti'ong resemblance to one another yet 
they exhibit certain differences. 

By reference to Table I it will be seen that the second 
hybrid on the whole does not come so near to the male- 
parent in its non-measurable characters as do the first and 
third hybrids. 

In hybrids 1 and 3, five of the characters are nearer to 
those of Cacatua, four are more or less intermediate 
between those of the two parents, and one is closer to that 
of Licmetis. 

The various characters, however, are not all similar to each 
other in the two hybrids. The colour of the naked skin 
around the eye is very pale blue in hybrid 1, and ahnost, 



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508 



ERNEST WARREN. 



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A FUKTllEK NOTE ON IIYJJKID COCKATOOS. 509 

aUhougli not quite^ white in hybrid 3. In hybrid 1 the 
naked area around the eye is roughly circular in ^^hape, and 
is about intermediate in size; while in hybrid 3 it is nearer 
to that in Cacatua. In the granulation of the eyelids 
hybrid 1 approaches Cacatua, nnd hybrid 3 is more or less 
intermediate. In the colour of the bill hybrid 3 is more 
distinctly intermediate than is hybrid 1. 

Hybrid 2 is less like the male-parent than is the case in 
the other two hybrids ; the naked area around the eye is 
distinctly blue and its shape approaches that in Licmetis. 
The prominence of the forehead is medium, but it is less 
marked than in the first hybrid. Thus in five characters 
the second hybrid approaches Cacatua, in two characters 
it is intermediate, and in three characters it is nearer to 
Licmetis. There is still prepotency on the part of the 
male-parent, ])ut it is less pronounced than in the first and 
third hybrids. 

The greater divergence of the second hybrid from the male- 
parent is confirmed by an examination of Table II showing 
the characters which were measured. Here the mean of the 
percentage divergences of the second hybrid from the father, 
expressed in terms of the characters in the former, is 29 per 
cent., as compared with 25 per cent, in the case of the first 
hybrid and 23 per cent, in the case of the young bird. 

The means of the percentage divergences from the male- 
parent (Cacatua) and the female-parent (Licmetis) are 
respectively for the first hybrid 25 per cent, and 32 per cent., 
for the second hybrid 29 per cent, and 33 per cent., for the 
third hybrid 23 per cent, and 35 per cent., and the averages 
for the three hybrids are 26 per cent, and 33 per cent. In 
all the hybrids there is a general average prepotency of the 
male-parent, but it is lest obvious in the second hybrid. 

Thus on the whole the same order of magnitude is main- 
tained in all three hybrids, although there is considerable 
variation in the expression of the different characters. 

With only three individuals to deal with it is not possible 
to draw any very definite conclusions, but the differences 



510 ERNEST WARKEN. 

between the brother hybrids would seem to be greater than 
might be expected to occur between three pure-bred 
brethren. 

Another point of interest may be mentioned. The eyes of 
the hybrids, as represented by the area of the eyeball 
exposed when the eyelids are wide open, are distinctly larger 
than in either the male- or female-parent, and this condition 
is especially marked in the second hybrid where the area is 
equal to 51 sq. mm., while in the male Cacatua it is 31 sq. 
mm., and in the female Licmetis 25 sq. mm. The appear- 
ance in a hybrid of a new character not clearly connected 
with either parent would seem to be especially worthy of 
investigation, but the data at present to hand are too scanty 
for any detailed examination. 



Vol. Ill, part 3, issued May ISth, 1917. 

THE PLANT ECOLOGY OF DRAKENSBERG RANGE. 511 



The Plant Ecology of the Drakensberg Ran ge 

By 

J. \V. Bews, ]?I.A., D.Sc, 

Professor of Botany, Natal University College. 



With Plates XXXVI-XXXIX, and 3 text-figures. 



T. 

II. 

III. 



Contents. 

Introduction 

Geology, Topography, and Soil Conditions 

Climate 

Plant Formations and Associations 

1. Veld Formation . 

2. Protea Veld Formation 

3. Rocky Senile Formation . 

4. Scrub Formation 

5. Bush Formation . 

6. Stream Bank Formation . 

7. Vlei Formation . 

8. Vegetation of the Movmtain Top 

9. Clii: Vegetation . 
10. Fynbosch or Mtiquis Formation 

IV. Succession and Inter-relationships 
Formations 
Explanation of Plates 




OF THE Plant 



562 
565 



Introduction. 
The study of the vegetation of Natal from the ecological 
standpoint was commenced by the writer in 1911, and three 
papers ^ on the subject have already appeared. Important 

' Bews, J. W.. " The Vegetation of Natal," ' Annals of Natal 
Museum," vol. ii, pt. 3, 1932. "An Ecological Sui-vey of the Midlands 
of Natal," ibid., vol. ii, pt. 4, 1913. "The Growth Forms of Natal 
Plants," • Trans, of Roy. Soc. of S.A.,' 1916. 

VOL. 3, part 3. 34 



512 J. W. BEWS. 

problems continue to present themselves, and the ecology of 
the Drakensberg Range has proved as interesting as its 
scenery is magnificent. 

While increasing knowledge of the flora of Natal and its 
ecology has helped to lighten the task in some respects, yet 
the immense distances and genei-al inaccessibility of most 
parts of the Drakensberg presented fresh difficulties. 

A month (Januaiy, 1914) was spent by the writer at Van 
Reenen's Pass, which is a convenient centre for investigating 
the lower portion of the range and the character and relation- 
ships of the main plant formations, the bnsh and veld. Sub- 
sequently shorter visits were paid to the Newcastle district, 
and the Groodoo Pass and Mont aux Sources region. 

The last mentioned will doubtless in time become much 
better known, since it is being set aside as a National Park. 
It is certainly worthy of the distincton. 

The writer desires to acknowledge generous assistance from 
the following: Mrs. R. Putt, Botanist to the Transvaal Museum, 
who sent a list of plants which she had named for Mr. R. E. 
Symons from Giant's Castle ; Mr. T. R. Sim for identifying the 
mosses and hepatics and other valuable assistance ; Dr. E. P. 
Phillips, of the South African Museum, who undertook the 
naming of various collections made by the writer, and who also 
supplied other lists ; Mr. J. S. Henkel, Conservator of Forests 
for Natal, for obtaining specimens through his forest officer 
stationed at Olivier's Hoek ; and Senator F. F. Churchill, Avho 
was visiting the Groodoo and Mont aux Sources at the same 
time as the writer. 



I. GEOLOGY, TOPOGRAPHY, AND SOIL 
CONDITIONS. 

The Drakensberg range of mountains forms the western 
boundary of Natal. It is, properly speaking, the elevated 
escarpment of the great inland plateau. It is only when 
viewed from the Natal side that it appears as a lofty range of 
mountains, the ascent from the other side being much more 



THK PLANT ECOLOGY OP DRAKENSBKRG BANGE. 513; 

gradual. On the borders of Basutoland it reaches its highest 
altitude^ the Mont aux Sources being given on the maps as 
11, 170; ft. (3405 m.), Cathkin Peak (Champagne Castle) as 
10,357 ft. (3157 m.), and Giant's Castle as 9657 ft. (2944 m.), 
but probably tliese heights are exaggerated. The general 
altitude reached, by this part of the range may be taken as 
approximately 3000 m. At the point where Basutoland 
joins Griqu^land E., the main range bends south-westward,, 
but a lower range passes along the boundai'y between. 
Griqualand E. and Natal, terminating in the Ingeli 
Mountains. 

• ., From the Mont aux Sources northwards along the boundary 
of .the Free State, the altitude gradually decreases to Vaix 
Reenen's Pass, this portion forming a range nearly at right 
angles to the higher part, which continues in line to form the 
boundary between Basutoland and the Orange Free State. 

From Van Reenen the portion bordering Natal rises again 
slightly and forms a long escarpment facing Newcastle, which 
has an altitude of 6500 to 7500 ft. (1981 to 2286 m.). 

This northern portion is much more accessible than the part 
bordering Basutoland. 

The geology of the Drakensberg is fairly simple, owing to 
the absence of folding. The rocks lie mors or less horizontally 
and may be clas.sified in descending order as follows : 

'Amygdaloidal, basaltic, and rhyolitic 

lavas. 
Cave Sandstones. 
.Red Beds. 
^Molteno Beds. 
Beaufort Series. 

Ecca Series (not including the glacial 
conglomerate). 

The amygdaloidal lavas in some parts form practically 
sheer cliffs, but there are numerous buttress-slopes covered 
with a shallow layer of soil, in which tussock veld is formed, 
and in places this reaches almost to the top even of the 



Upper Karroo 



Lower Karroo 



514 J. w. Bbiws. 

highest portions. The two sections given in text-figs. 2 and 
3 (pp. 516, 517) illustrate the two types. 

Immediately below the araygdaloids, a conspicuous sheet of 
dolerite or basalt (it is difficult to say which it is), varying 
from 50 to 300 or more feet in thickness, occurs ; and this 
hard bed forms a protective cap for many of the upper foot- 
hills (text-fig. 1, p. 515). The lavas have been totally 
denuded from every part except the highest portion, Avhich 
extends from the Mont aux Sources to the southernmost 
boundary of Natal. 

The Cave Sandstones, Red Beds and Molteno Beds, the 
downward succession of the Storniberg Series, form the 
northern portion and are exposed in the same order on 
the slopes facing Natal, below the amygdaloids. Sandstones 
alternate with thin beds of shale. The Cnve Sandstones 
usually form conspicuous cliffs, covered with alga3 and 
lichens. 

The lower foothills which extend about thirty miles from 
the main range consist of reptilian sandstones and mudstones 
of the Beaufort Series. Below these again are the Natal 
Coal Measures, which are usually considered to belong to the 
upper Ecca Series. There is complete conformity between 
all the divisions from the Cave Sandstones downwards — and 
only a slight unconformity between the Cave Sandstones and 
the overlying lavas. 

From the main range of mountains the chief rivers of 
Natal have their source, and they have cut through the out- 
crops of the various geological formations nearly at right 
angles. The result is that Natal, as a whole, rises from the 
sea to the Drakensberg in a series of terrace-plateaux. The 
Drakensberg range forms the highest plateau. The vege- 
tation on the steep edge of this plateau, which faces Natal, 
differs considerably from the vegetation on the plateau itself, 
i. e. in the interior (cf. the differences between the edges 
and the surfaces of the lower terraces in Natal, as described 
by the writer in the previously cited papers). Between the 
rivers or between their main tributaries we have a system of 



THE PLANT ECOLOGY OF DKAKENSBERG RANGE. 515 



ridges, spurs, and foothills, pi-ojecting more or less at right 
angles to the main escarpment. The rivers and their tribu- 
taries have cut their way down very steeply, leaving usually 
sheer walls of rock at their upper ends and steep slopes 



Text-pig. 1. 



.Tussoch Veld 




^ Grey i a Scrub 

Prolea Veli 



River Bed with 
„ Leiicosidea . 
__^ Scrub 



Section tbrough a typical foothill of the Drakensberg escarpment, 
sliowing the relationship of plant formation to the geological strata. 

where the valleys begin to broaden. In spite of the fact, 
therefore, that the greatest portion of the main range faces 
south-east, the exposures vary exceedingly and the climate, 
soil conditions, and plant formations vary accordingly. 

This fact is well illustrated as one passes from Bei'gville 
towards the Mont aux Sources up the great valley of the 
Tugela. 



<516 



J. W. BEWS. 



On the left the Drakensberg faces north-east, and the 
■tributaries of the Tugela flow down between foothills in 
•valleys which also open to the north-east. A good example 
;is the valley of the Sanyati River, a grassy type devoid of 



Vegelalion of 
MountcLin Top 



Text-fig. '2. 



^Liihophyies 

and 
Chomophyles 



Fyniosch or Mciqui 




Lilhophyles 

CtTld 

Chomophyles 
Proiea Veld 

Greyia Scrub 



Section through the steep part of the main Drakensberg 
, escarpment. 

bush and shrub. On the right the range faces south-east, 
and here there are several fairly large patches of the mountain 
type of bush. 

; The rate of denudati,on is everywhere: at a maximum. 
After the winter dry season the Tugela (like the other main 



THE PLANT ECOLOGY OF DltAKENSBEIiG RANGE. 517 



rivers) often rises in flood and comes down in the form of a 
wall of water several feet high, which sweeps before it the 
accumulated debris of the river-bed and rolls along large 
stones, leaving the river-channel clean. 

Text-fig. 3. 



Vegelalion of 
MountaiTt Top 



Liihophyles 

and 
Chomophyles 



Fynbosch 



Tussoch Veld 




Liihophyles 

and 
Chomophyles 



Prolea Veld 
Greyiu Scrv,b 



Section through a buttress-slope of the main Drakensberg 
escarpment. 

Where soil is able to accumulate it is quickly leached and 
is usually very poor in chemical salts. The shales are of a 
very friable character and are easily eroded. The sand- 
stones among which they occur are more resistant. 
Frequently, therefore, the shale beds are hollowed out under- 



518 



J. W. BEWS. 



neath overlying sandstones. In many places tlie process lias 
doubtless been assisted by the Bushmen^ who used the caves 
thus formed for shelter. These caves occur not only in the 
Cave Sandstones but in the other beds as well. Their chief 
interest to most people lies in the Bushman paintings, which 
they often contain, but they also afford shelter for many 
interesting shade-loving plants. In such caves, large and 
small, soil is able to accumulate, and if, as often happens, the 
type of plant formation is bush or scrub, a rich leaf-mould 
gathers and the undergrowth is relatively a rich and varied 
one, ferns and brj^ophytes in particular being abundant. 

At the top of the lower portion of the range (A^an Reenen's 
Pass, etc.) we find conditions of greater stability than on the 
Natal face. 

Here there are vleis of considerable size, Avith a varied 
and interesting flora, and comparatively stable veld. The 
soil, however, even in such places, is a poor one, the soluble 
salts being quickly dissolved out of it. The following is an 
analysis of a sample of soil from Botha's Pass, and for pur- 
poses of comparison there is added the average of twenty- 
seven analyses (already published) for different parts of 
Natal : 



Locality. 


Percentages of constituents. 


Moisture. 

Organic 
and vola- 
tile. 




o 

o . 

§* 

.a 

ft, 


1 

o 


s 
13 


.2 

Q> 

a 


* a 

U 

534 


Botha's Pass 


1-77 5-22 86-89 


0-04 


0-18 004 


0-05 


Average of 27 analyses 
for lowlands of Natal 


4-82 7-63 


70-09 


0-09 


019 


0-23 


0-20 


11.64 



All Natal soils show extreme poverty in soluble salts, there 
being great scarcity particularly of lime and phosphates. 
The soil of the Drakensberg, however, is still poorer, having 
a high proportion of insoluble minerals. Tlie water of the 



THE PLANT ECOLOGY OF DK'AKENSBERG RANGE. SIO" 

moimtain streams, which is clear as crystal, is very soft,, 
showing absence of lime. 

On the top of the upper portion, from the Mont aux Sources 
southwards, a distinct type of soil and plant formation 
occurs — the mountain-top detritus. This will be described in 
detail later, when the vegetation is dealt with. 

II. CLIMATE. 

A. Atmospheric Pressure and Insolation. 

The feature of mountain climate which shows the greatest 
regularity is the decrease of pressure with increasing altitude. 
It must naturally come first in our consideration, since so many 
of the other features depend on it. It varies with tempera- 
ture and therefore with latitude, and consequently the decrease 
is not quite so great in Natal ns in colder regions. The 
irregular changes in barometric pressure decrease with alti- 
tude in the same ratio as the pressure itself decreases. On 
the summit of the DraUensbeig the pressure is from 150- 
250 mm. less than at sea-level, where it may be assumed to- 
be 762 mm. (30 in.). 

As a result of the decrease of pressure there is increased 
intensity of insolation, since the thickness of absorbing atmo- 
sphere is less. Water- vapour, which is a better absorber of 
solar radiation than dry air, decreases with altitude more 
rapidly than pressure decreases (vide infra). Atmospheric 
dust, which aifects chiefly the shorter wave-lengths, is com^ 
paratively small in amount, consequently there is a marked 
increase in the more chemically active rays. The great 
intensity of the ultra-violet rays at high altitudes has been 
demonstrated by many observers. Bunsen and Roscoe many 
years ago gave the following figures regarding the chemical! 
intensity of sunlight at different altitudes, expressed in per- 
centages of the intensity just outside the earth's atmosphere,, 
which they computed to be equal to 35*3 " light units." 



520 



J. W. BEWS. 



Chemical Intensity of Sunlight (in percentages 
of the maximum),^ 



Pressure 
(mm.). 


Altitude above sea 
level (m.). 


Sun's altitude. 


90 
degrees. 


70 
degrees. 


50 
degrees. 


30 
degrees. 


10 
degrees. 


750 
650 

550 .. 
450 ' 
350 > 


130 
1270 

,: , , . 2600 
" 4200 
, 16200 


44 
49 

55 
61 

68 


42 
47 
53 
59 
67 


34 
39 
46 
53 
61 


19 
24 

30 
37 

46 


1 

2 
3 

6 
10 



According to this table, when the sun is low, the chemical 
intensity of sunlight on the Drakensberg is three or four 
times greater than on the coast-belt, but the difference 
decreases as the sun rises to the zenith, when it is only 11 or 
12 per cent, greater. 

The remai-kable colour changes in the flowers of species 
which extend from the coast to the higher altitudes is one of 
the many physiological effects of this factor. Helichrysum 
adenocarpnm has flower-heads white or pale pinkish on 
the coast, but brilliant crimson in upper districts. Dierama 
pendula has pure white flowers on the coast, pink in the 
midlands, and deep purple on the mountains. Moraea 
spathacea shows similar variations. 

A relatively high surface temperature of the soil is a direct 
result of the increased insolation. The temperature of the 
soil to a depth of several centimetres is often much higher 
during the daytime than that of the atmosphere. On the 
other hand, owing to the rarity of the air and the decrease 
in the amount of water-vapoui% radiation of heat at night is 
increased, the temperature of the surface falls, and there is 
thus a much larger range of surface temperatures' on the 
Drakensberg than on the lower terraces. The exposure is of 



1 Hann, J., ' Handbook of Climatology," Trans, by D. C. Warfl. 
New York, 1903. 



THE PLANT ECOLOGY OF DRAKENSBEEG RANGE. 521 

great importance in connection witli insolation. A large 
portion of the main Drakensberg faces the rising sun, but it 
will be seen from the description given of the topography 
that there are minor slopes and innumerable steep descents 
into the deep valleys, that are shaded much longer, some- 
times throughout the greater part of the day. This fact, 
taken in conjunction with the distribution of water-vapour 
and precipitation, has an important effect on the vegetation. 

B. Temperatuke. 

Next in point of regularity to the decrease of pressure 
is the decrease of air temperature with rise of altitude. 
This has been taken* to average 0"57°C. for every 100 m. 
There are slight seasonal variations, of course, but here 
again the effect of the topography is most marked. At night, 
•owing to radiation, which, as we have seen, increases greatly 
with altitude, the ground cools rapidly and the lower atmo- 
sphere is also cooled. The cold air, being heavier, settles to 
the lowest point ; it flows fx'om the hillsides down into the 
valleys, and it follows the course of the main valleys down- 
ward, exactly like the rivers. We thus have an inversion of 
temperature, higher altitudes being warmer than lower. This 
is seen regularly during the winter anticyclone, when calm 
weather prevails, when also the general temperature is lower. 
The water-vapour of this colder air is condensed into a fog, 
which settles into the valleys. In the early morning the 
mountain-tops rise above the dense white mist-cloud like 
islands in a sea. The rising sun gradually disperses the 
fog, and here again the exposure determines whether the 
sun's influence will soon make itself felt, or whether the cold 
air and fog Avill linger. The effect of the cold air drainage 
at night is seen most clearly at points where the current is 
moi-e rapid, as where a narrow " nek " opens into a broader 
valley. At such a point the vegetation suffers severely from 
ifrosts. 

' Hann, J., loc. cit. 



522 J. w. BEWS. 

The cold air, which thus flows down to lower altitudes, has 
to be replaced by air from above, which has not been cooled 
by radiation and is further warmed by compression as it 
descends. Consequently, the air on the mountain sides and 
even at the top may be relatively wai'm in winter. At the 
same time it is very dry. The valleys, therefore, are damp 
and cold at nig'hts, the mountains dry and warm. 

On the other hand, during the day-time, and particularly 
during summer, the temperature of valleys is relatively high, 
owing to the heat resulting from insolation being reflected and 
radiated from the sides of the valleys. The foot of the main 
escarpment, for the same reason, is relatively Avarm. The 
valleys and lower altitudes generally, therefore, show a much 
greater range of temperature than the enclosing mountains. 
A decrease of the annual and diurnal range of temperatures 
is a general feature of high altitudes, but it depends to a 
large extent on the topography. A valley at high altitude 
will show a much greater range than a hill of the same 
altitude. 

A common feature of mountain ranges in higher latitudes, 
where there is a covering of snow, is the retardation of the 
time of occurrence of maxima and minima temperatures and 
of the seasons generally. This does not apply to the Drakens- 
berg. It is true that snow may lie on the summit for several 
weeks, but its melting does not require a very large amount 
of ins